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Article history:
Received 23 October 2008
Accepted 6 February 2009
Available online 15 February 2009
Food preferences, feeding attributes, trophic guilds, and ontogenetic changes in diet compositions of 45
sh species collected along coastal waters in the southern areas of the South China Sea, Thailand were
examined. Most species had high food intake and fed on specic ranges of food types. Shrimp was the
most important food (31.7%), followed by calanoid copepod (16.8%), sh (12.7%) and gammarid
amphipod (8.3%). These shes can be categorized into six different trophic guilds and further divided into
four categories namely; piscivore, zooplanktivore, zoobenthivore and miscellaneous/opportunist.
Numbers of feeding guilds at each depth vary between four and ve. Four of these are consistently found
throughout the year at all zones; shrimp predator, piscivore, calanoid copepod feeder and a combination
of polychaete and other food feeders. Ontogenetic studies indicate that shes are more likely to group
according to species rather than size. This scientic information is important when examining the
complex association between shes and identifying groups of species using similar resources.
2009 Elsevier Ltd. All rights reserved.
Keywords:
diet composition
feeding habits
functional role
Gulf of Thailand
sh ecology
1. Introduction
Knowledge in the trophic ecology of any given system is
fundamental in understanding the ecosystem as a whole (Blaber,
1997; Cruz-Escalona et al., 2000). It therefore becomes important to
determine not only the feeding mode, but also the extent and
nature of inter-specic and inter-guild trophic relationships (Elliott
et al., 2007). A guild refers to a group of species exploiting the
same class of environmental resources in a similar way (Root, 1967).
When used in sh community studies, it offers the possibility of
dividing a community into functional groups (Schlosser, 1982; Ross,
1986; Bayley, 1988; Gerking, 1994; Garrison and Linke, 2000). The
concept of feeding guild in estuarine shes was rst introduced by
de Sylva (1975) who dened groupings of shes based on their
feeding preferences and food web structure. Recently, Elliott et al.
(2007) classically identied seven standard categories of the
feeding functional groups (FMFG) in estuarine habitat; detritivore,
herbivore, omnivore, zooplanktivore, zoobenthivore, piscivore and
miscellaneous/opportunist. This FMFG classication is a trophic
guild system designed to allow the aggregation of sh species that
utilize similar food resources to help understand, explain and use in
a management context, the functioning of this habitat.
Spatial and seasonal factors greatly affect the structuring of
trophic organizations of shes in a particular area (Elliott et al.,
E-mail address: hsukree@bunga.pn.psu.ac.th
0272-7714/$ see front matter 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.ecss.2009.02.010
504
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505
Table 1
Trophic indices and relative composition by volume of food of 45 sh species collected off coastal waters of Pattani and Narathiwas, Thailand between March 2006 and
November 2006. VI vacuity index, FL mean gut fullness, Bi diet breadth.
Species
Alectis indicus
Apogon endekataenia
Apogon niger
Apogon quadrifasciatus
Arius maculatus
Arius venosus
Caesio cuning
Callionymus planus
Dactylopus dactylopus
Diagramma pictum
Epinephelus bleekeri
Epinephelus coioides
Epinephelus sexfasciatus
Gazza minuta
Gerres lamentosus
Himantura bleekeri
Inegocia japonica
Johnius dussumieri
Leiognathus brevirostris
Leiognathus equulus
Leiognathus splendens
Leiognathus stercorarius
Lieognathus decorus
Lutjanus erythropterus
Lutjanus madras
Monacanthus chinensis
Paramonacanthus
choirocephalus
Nibea soldado
Oxyurichthys saru
Pentapodus setosus
Plotosus anguillaris
Polynemus tetradactylum
Pomadosys maculatus
Priacanthus tayenus
Saurida micropectoralis
Saurida undrosquamis
Scolopsis taeniopterus
Secutor ruconius
Siaganus canaliculatus
Sillago sihama
Trichiurus lepturus
Upeneus sulphureus
Upeneus sundaicus
Upeneus tragula
Vincentia chrysura
Average
No. of
sample
Total length
(cm)
Trophic attributes
VI
FL
Bi
No. of
food type
Shrimp
Calanoid
copepod
Polycheate
Gammarid
amphipod
Crab
Fish
Cryprid
Others
Digested
items
10
69
95
507
21
23
148
223
58
31
17
22
14
37
37
11
261
18
144
15
242
99
23
13
63
29
10
10.1
6.3
7.6
7.6
15.8
14.3
8
9.1
9.7
8.8
14.4
13.6
13.4
12.7
15.1
16
10.9
10.3
7.6
13.8
9.2
7.3
10.5
11.4
13.3
11.2
10.4
0
5.8
14.7
12.6
4.8
0
0
1.3
3.4
3.2
5.9
22.7
56.2
13.5
2.7
9.1
41.8
5.6
3.5
0
1.2
2
13
0
25.4
3.4
0
4.4
3.7
2.4
2.3
3.7
1.9
3.2
3
2.9
3.2
2.9
2.1
4.6
3.1
3.7
3.3
3.8
1.5
2.8
3.7
3.1
2.9
3.0
4.7
2.4
3.1
2.3
0.17
0.20
0.37
0.20
0.22
0.16
0.09
0.27
0.23
0.19
0.64
0.58
0.44
0.39
0.15
0.24
0.15
0.38
0.47
0.13
0.12
0.18
0.37
0.36
0.18
0.57
0.52
3
9
10
13
5
9
8
12
11
8
3
4
2
7
8
5
10
4
8
7
15
12
8
4
7
9
7
80
54.8
34.0
44.0
7.5
60.4
16.4
2.6
7.0
57.7
35.0
11.8
0.0
8.8
7.5
60.0
59.1
13.5
18.5
16.3
2.5
16.5
11.5
23.1
57.7
11.4
31.1
0
19.0
19.1
17.3
0.0
0.4
75.1
34.8
35.6
21.3
0.0
5.9
0.0
12.5
0.8
10.0
2.6
56.5
23.2
2.7
52.3
51.5
42.5
0.0
1.3
22.5
17.8
0
0.9
0.0
0.6
6.5
8.3
1.1
12.8
5.2
0.0
0.0
0.0
0.0
4.7
65.0
18.0
2.4
0.0
30.5
70.7
24.9
14.3
7
0.0
2.1
18.9
8.9
0
3.5
7.4
4.5
0.0
0.0
0.8
30.4
37.0
8.7
0.0
0.0
0.0
0.0
11.1
0.0
1.6
24.1
14.7
0.0
8.0
6.3
5
0.0
0.4
14.6
25.6
16.5
0.3
2.2
0.9
12.5
1.7
0.0
0.1
0.0
0.0
11.3
17.6
0.0
0.0
0.0
0.0
11.3
0.0
0.0
0.0
0.0
0.0
0
17.7
0.0
7.1
0.0
0
11.1
9.9
4.8
0.0
4.3
3.2
0.0
0.2
1.3
28.8
23.5
57.1
42.8
1.7
4.0
6.1
0.0
0.0
0.0
0.1
0.0
0
56.9
16.2
0.0
0.0
0
2.6
0.2
0.8
0.0
0.4
0.0
1.5
4.4
0.3
0.0
0.0
0.0
22.8
0.0
0.0
0.2
0.0
2.3
1.3
5.4
0.8
21
0.0
1.9
0.0
1.1
3.5
1.5
6.2
6.6
68.5
7.0
1.4
13.2
6.1
4.0
0.0
5.9
14.3
0.6
11.1
2.0
9.5
0.0
2.8
2.3
4.9
4.4
8.0
2.3
1.3
7.5
15.6
0
6.2
21.0
20.5
5.0
17.4
2.0
4.7
4.6
6.7
25.0
35.3
28.6
7.8
2.8
6.0
7.1
5.9
7.9
6.7
1.9
6.2
5
0.0
19.1
21.4
0.0
47
84
14
31
10
21
190
297
212
300
142
139
12
18
382
44
366
54
9.5
10.7
10.8
8.5
10.8
8.4
10.3
13.8
9.8
13.1
7.2
8.7
13.4
33.2
8.2
9
8.8
7.1
10.6
7.1
0
3.2
0
0
0.5
31
46.7
4
4.9
3.6
8.3
5.6
1
2.3
1.6
29.6
3.0
3.5
3.9
2.6
3.0
2.2
4.5
3.4
4.5
3.3
2.8
2.8
2.9
3.0
3.6
2.8
3.3
2.1
0.13
0.16
0.41
0.19
0.40
0.21
0.24
0.13
0.07
0.16
0.24
0.09
0.69
0.00
0.09
0.14
0.06
0.18
10
9
8
4
4
5
14
10
6
18
12
13
8
1
12
8
12
6
56.3
2.8
39.3
72.3
25.0
66.2
36.6
30.1
7.5
26.9
17.9
11.5
18.2
0.0
62.0
66.0
73.3
67.1
0.3
58.7
11.4
21.0
0.0
0.0
16.0
0.7
1.8
4.7
41.7
3.0
0.0
0.0
29.8
11.9
19.4
11.8
24.0
4.1
20.4
3.3
1.0
4.8
16.3
1.9
0.0
40.9
5.6
66.8
18.2
0.0
0.7
0.0
0.5
0.0
0.9
23.5
1.1
0.0
54.0
19.5
18.6
0.5
0.0
5.5
0.7
5.5
22.7
0.0
2.2
13.3
2.6
0.0
0.9
1.3
3.6
0.0
0.0
0.0
1.9
1.7
4.0
1.1
0.0
0.0
9.1
0.0
0.9
1.2
0.1
0.0
4.0
0.0
5.0
0.0
22.0
0.0
5.7
57.5
77.4
0.9
1.5
3.4
9.1
100.0
0.6
0.0
0.3
10.5
0.0
0.0
12.1
0.0
0.0
0.0
2.9
0.0
0.0
6.2
14.4
0.1
0.0
0.0
1.3
3.5
1.2
0.0
9.9
5.8
7.1
0.0
0.0
9.5
1.5
3.3
8.4
7.6
16.9
8.9
13.6
0.0
0.8
1.9
1.5
7.9
3.7
3.8
0.0
3.3
0.0
0.0
0.5
4.4
0.9
6.1
1.5
0.7
9.1
0.0
1.6
2.3
1.1
2.6
9.2
3.1
0.30
8.2
31.7
16.8
11.4
8.3
2.8
12.7
2.4
6.9
7.0
4603
Food types
506
Piscivore
Trichiurus lepturus
Saurida undrosquamis
Epinephelus sexfasciatus
Calanoid
copepod Feeder
Shrimp Predator
Apogon quadrifasciatus
Apogon endekataenia
Apogon niger
Upeneus sulphureus
Diagramma pictum
Upeneus tragula
Plotosus anguillaris
Nibea soldado
Himantura bleekeri
Pentapodus setosus
Lutjanus madras
Arius venosus
Vincentia chrysura
Upeneus sundaicus
Pomadasys maculates
Inegocia japonica
Alectis indicus
Fish
Calanoid
copepod
Shrimp
Gammarid
amphipod
Polychaete
Combined-food
Feeder
Polynemus tetradactylum
Priacanthus tayenus
Monocanthus Choirocephalus
Monacanthus chinensis
Leiognathus brevirostris
Sillago sihama
Oxyurichthys saru
Johnius dussumieri
Caesio cuning
Secutor ruconius
Leiognathus decorus
Leiognathus
stercorarius
Leiognathus splendens
Dactylopus dactylopus
Callionymus planus
Fish/Shrimp
Feeder
Epinephelus coioides
Epinephelus bleekeri
Saurida micropectoralis
Lutjanus erythropterus
Gazza minuta
Young crab
Polychaete Predator
Siganus canaliculatus
Leiognathus equulus
Gerres filamentosus
Scolopsis taeniopterus
Arius maculatus
Fig. 2. General trophic guilds of shes collected off coastal waters of Pattani and Narathiwas, Thailand between March and November 2006. Thick label major food type, thin
label minor food type. (ANOSIM Global R 0.922, P 0.001).
507
G1
Monacanthus chinensis
Apogon niger
Callionymous planus
Dactylopus dactylopus
Secutor ruconius
Oxyurichthys saru
G2
Leiognathus brevirostris
Leiognathus splendens
Pomadasys maculatus
Apogon endekataenia
Apogon striatus
Plotosus anguilaris
Upeneus tragula
Priacanthus macracanthus
G3
Nibea soldado
Arius venosus
Upeneus sulphureus
Apogon quadrifasciatus
Inegocia japonica
Saurida micropectoralis
G4
Saurida undrosquamis
20
40
60
80
100
G1
Leiognathus decorus
Secutor ruconius
Saurida micropectoralis
Saurida undrosquamis
G2
Epinephelus coioides
Leiognathus brevirostris
Priacanthus macracanthus
Scolopsis taeniopterus
Monacanthus chinensis
Siganus canaliculatus
G3
Gerres filamentosus
Inegocia japonica
Nibea soldado
Apogon niger
Dactylopus dactylopus
Apogon endekataenia
Leiognathus stercorarius
Callionymous planus
G4
Oxyurichthys saru
Leiognathus splendens
Plectorhinchus vittatus
20
40
60
80
100
Fig. 3. Cluster dendogram demonstrating trophic groups of shes collected at different depth zones off coastal waters of Pattani and Narathiwas, Thailand between March and
November 2006. G1G5 trophic groups.
508
G1
Monacanthus chinensis
Leiognathus decorus
Secutor ruconius
Apogon quadrifasciatus
Upeneus tragula
Upeneus sundaicus
G2
Lutjanus madras
Inegocia japonica
Upeneus sulphureus
Saurida undrosquamis
Epinephelus coioides
Gazza minuta
G3
Apogon niger
Saurida micropectoralis
Scolopsis taeniopterus
Nibea soldado
Siganus canaliculatus
G4
Gerres filamentosus
Leiognathus splendens
Dactylopus dactylopus
Leiognathus stercorarius
Oxyurichthys saru
G5
Caesio cuning
Plotosus anguilaris
Plectorhinchus vittatus
20
40
60
80
100
G1
Epinephelus coioides
Caesio cuning
Oxyurichthys saru
Leiognathus splendens
G2
Monacanthus chinensis
Plectorhinchus vittatus
Upeneus sulphureus
Plotosus anguilaris
Apogon niger
Inegocia japonica
Lutjanus madras
G3
Apogon endekataenia
Upeneus tragula
Apogon quadrifasciatus
Upeneus sundaicus
Apogon striatus
Scolopsis taeniopterus
Priacanthus macracanthus
G4
Dactylopus dactylopus
Callionymous planus
20
40
60
80
100
509
Table 2
Summary results of similarity percentage analysis (SIMPER) on the contribution of food types in the formation of trophic guilds in different months and depth zones.
G1G5 Trophic guilds 16; % % contribution.
Month
Guild
Food types
Month
Guild
Food types
March 06
G1
Calanoid copepod
Shrimp
Gammarid amphipod
Shrimp
Polychaete
Gammarid amphipod
Calanoid copepod
Calanoid copepod
46.8
33.2
8.5
91.8
41.5
30.4
8.1
55.4
July 06
G1
G4
Calanoid copepod
Gammarid amphipod
Polychaete
Polychaete
Shrimp
Shrimp
Calanoid copepod
Fish
63.5
22.2
8.4
72.1
19.1
87.1
10.0
100
Fish
36.6
September 06
G1
Fish
99.3
Shrimp
Fish
Shrimp
Gammarid amphipod
Calanoid copepod
71.8
14.3
89.2
38.8
28.8
G2
G3
Calanoid copepod
Polychaete
Shrimp
Shrimp
Calanoid copepod
97.8
69.5
21.3
72.4
22.3
Shrimp
Polychaete
Calanoid copepod
Polychaete
Shrimp
15.1
12.5
86.3
63.7
25.1
G2
G3
G4
May 06
G1
G2
G3
G4
G5
G2
G3
G4
November 06
G1
G2
Calanoid copepod
Shrimp
Mantis shrimp
92.0
66.7
33.3
Zone
Guild
Food types
Zone
Guild
Food types
G1
Polychaete
Shrimp
Calanoid copepod
Polychaete
Shrimp
Calanoid copepod
Fish
42.4
30.8
80.0
10.8
87.7
8.5
100
G1
Calanoid copepod
Shrimp
Gammarid amphipod
Polychaete
Shrimp
Fish
Shrimp
39.1
25.5
23.1
5.9
94.1
66.4
27.5
Shrimp
Calanoid copepod
Fish
Polychaete
51.9
39.5
95.5
59.5
Polychaete
Shrimp
Calanoid copepod
Shrimp
85.6
7.6
89.6
6.3
Shrimp
Calanoid copepod
Calanoid copepod
Gammarid amphipod
24.7
7.7
86.7
9.7
Fish
Calanoid copepod
Shrimp
Calanoid copepod
Polychaete
Shrimp
Gammarid amphipod
91.1
96.7
82.3
13.0
58.5
30.8
4.0
G2
G3
G4
2
G1
G2
G3
G4
G2
G3
G4
G5
4
G1
G2
G3
G4
copepods formed the second cluster. The third group was shes
that had polychaetes and some contribution of shrimps as their
food. Eight species formed the fourth group with a great contribution of shrimps and some calanoid copepods in the diets.
Only seven sh species were analyzed for the month of
November 2006 (ANOSIM Global R 0.98, P 0.001) (Fig. 4e). Of
these, two clusters and a single individual were formed. The rst
group was shes feeding mainly on calanoid copepods and the
second cluster fed on a combination of shrimps and mantis
shrimps.
4. Discussion
510
G1
Siaganus canaliculatus
Priacanthus tayenus
Nibea soldado
Lutjanus madras
Upeneus sulphureus
G2
Apogon quadrifasciatus
Inegocia japonica
Upeneus tragula
Leiognathus brevirostris
Leiognathus splendens
G3
Callionymus planus
Scolopsis taeniopterus
Gerres filamentosus
Dactylopus dactylopus
Gazza minuta
Saurida undrosquamis
G4
Apogon endekataenia
Leiognathus stercorarius
20
40
60
80
100
G1
Upeneus sulphureus
Epinephelus coioides
Upeneus sundaicus
Upeneus tragula
Inegocia japonica
G2
Pomadosys maculatus
Plotosus anguillaris
Arius venosus
Saurida undrosquamis
Apogon niger
Dactylopus dactylopus
Leiognathus brevirostris
G3
Calionymus planus
Monacanthus chinensis
Caesio cuning
Secutor ruconius
Leiognathus stercorarius
G4
Leiognathus splendens
Scolopsis taeniopterus
Priacanthus tayenus
Siaganus canaliculatus
20
40
60
80
G5
100
Fig. 4. Cluster dendogram demonstrating trophic groups of shes collected in different months off coastal waters of Pattani and Narathiwas, Thailand between March and
November 2006. G1G5 trophic groups.
this study, which is a different scenario from the majority of tropical estuarine sh species being omnivorous and incorporating
plants, algae or macrophytes, in their diets as described by Blaber
(2001). It is therefore postulated that the planktivorous guild will
be less important in the deeper area of the same locality compared
to a near-shore habitat. Moreover, diet of some sh species found in
the bay habitat differs from that in this environment. For example,
Leiognathus splendens and Epinephelus coioides in the bay habitat
feed mainly on harpacticoid copepods and shrimp, respectively
(Hajisamae and Ibrahim, 2008), and change to calanoid copepods
and sh, respectively in this study.
The occurrence of feeding guilds has been widely discussed as
a possible strategy to avoid competition (Pianka, 1980; Angel and
Ojeda, 2001) or to optimize available resources (Jacksic, 1981). In
511
G1
Oxyurichthys saru
Siaganus canaliculatus
Gerres filamentosus
Leiognathus splendens
G2
Leiognathus stercorarius
Priacanthus tayenus
Upeneus tragula
Plotosus anguillaris
Upeneus sulphureus
Apogon endekataenia
Apogon niger
G3
Apogon striatus
Inegocia japonica
Apogon quadrifasciatus
Saurida micropectoralis
G4
Epinephelus coioides
20
40
60
80
100
G1
Epinephelus coioides
Caesio cuning
Callionymus planus
G2
Oxyurichthys saru
Siaganus canaliculatus
Gerres filamentosus
Nibea soldado
G3
Priacanthus tayenus
Arius venosus
Inegocia japonica
Lutjanus madras
Apogon endekataenia
Apogon quadrifasciatus
G4
Upeneus tragula
Upeneus sulphureus
Vincentia chrysura
Apogon niger
20
40
60
80
100
Upeneus sulphureus
Callionymus planus
G1
Oxyurichthys saru
Saurida undrosquamis
Inegocia japonica
G2
Nibea soldado
20
40
60
Fig. 4. (Continued).
80
100
512
Table 3
Trophic indices and relative composition by volume of food of 10 dominant shes collected off coastal waters of Pattani and Narathiwas, Thailand March 2006 and November
2006. VI vacuity index, FL mean gut fullness, Bi diet breadth.
Species
Length
interval (cm)
Trophic indices
VI
FL
Bi
No. of
food types
Food types
Shrimp
S. taeniopterus
<10.0
10.115.0
15.120.0
>20.0
0
3.1
6.2
11.1
2.6
3.5
3.1
4.5
0.22
0.14
0.23
0.23
14
14
13
8
38.2
25.4
20.4
7.5
8.8
4.4
3.8
0.0
24.5
49.8
37.9
58.8
S. undrosquamis
<6.0
6.18.0
8.110.0
10.115.0
15.120.0
>20.0
44.4
65.7
50
32
22.2
23.1
4.3
4.6
4.9
4.2
5.1
5.0
0.31
0.06
0.05
0.12
0.24
0.08
4
3
4
4
3
4
19.0
0.0
1.2
8.3
20.0
1.0
10.0
0.0
0.0
0.0
0.0
0.0
S. micropectorails
<10.0
10.115.0
15.120.0
>20.0
15.3
28
4.8
4.1
3.0
3.3
4.8
4.1
0.23
0.17
0.24
0.17
6
9
4
4
34.0
32.2
28.8
16.0
L. splendens
<8.0
8.110.0
>10.0
2
0.8
1
2.7
3.0
3.4
0.32
0.13
0.15
10
12
12
U. sulphureus
<6.0
6.18.0
8.110.0
>10.0
6.5
0
0.8
0
3.2
3.7
3.6
3.6
0.10
0.04
0.02
0.04
U. tragula
<6.0
6.18.0
8.110.0
>10.0
10
0
0.7
3.5
3.7
3.4
3.3
3.4
C. planus
6.18.0
8.110.0
>10.0
0
2.2
0
S. canaliculatus
<6.0
6.18.0
8.110.0
10.115.0
>15.0
C. cuning
A. quadrifasciatus
Calanoid
copepod
Gammarid
amphipod
Crab
Cryprid
Sand
2.3
2.6
6.3
0.0
14.4
4.1
2.4
3.8
2.0
0.5
0.2
0.0
1.7
0.4
0.0
0.0
1.3
5.4
7.4
6.3
0.2
0.2
1.4
8.8
0.0
0.0
0.0
0.0
3.0
2.6
5.5
15.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
4.3
5.9
6.9
0.0
0.0
63.5
73.9
81.2
81.9
80.0
89.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
10.0
0.0
2.2
0.0
0.0
3.3
1.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.5
0.0
0.0
1.5
1.6
4.0
0.0
51.4
51.3
83.2
76.0
0.1
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
2.4
0.0
4.0
8.1
0.9
1.5
34.8
57.2
55.3
25.3
24.9
24.3
0.8
0.9
1.8
15.9
7.8
3.0
0.0
0.0
0.0
0.0
0.0
0.3
5.6
3.0
8.9
0.0
0.1
0.0
0.0
0.3
0.0
2.7
1.8
3.6
3
9
7
6
48.8
57.3
61.6
83.0
0.0
0.0
0.0
0.0
0.0
0.3
0.5
2.4
0.0
0.0
0.3
0.0
5.1
3.2
1.4
0.0
0.0
0.6
0.0
3.9
0.0
1.0
0.0
1.2
0.0
3.5
0.0
0.0
0.0
0.3
0.3
0.1
0.0
0.0
0.0
0.0
0.0
0.1
0.5
0.0
0.50
0.10
0.09
0.05
3
9
9
8
51.1
69.7
70.3
85.7
48.9
26.8
16.9
9.9
0.0
0.0
0.3
1.6
0.0
0.0
0.0
0.4
0.0
1.8
4.7
0.2
0.0
0.2
0.1
0.0
0.0
0.4
0.5
0.0
0.0
0.4
2.4
0.0
0.0
0.0
0.3
1.0
0.0
0.0
0.0
0.0
0.0
0.7
0.0
1.2
3.0
2.8
3.5
0.28
0.33
0.30
8
10
11
2.3
2.2
4.3
48.2
33.7
23.8
12.3
13.2
11.8
0.9
8.0
7.1
25.5
29.6
38.5
0.0
0.1
0.0
0.0
0.0
0.0
0.0
1.4
3.4
5.7
4.6
5.0
0.0
0.0
0.0
0.9
2.3
2.3
11.7
6.3
0
0
0
1.6
2.7
2.7
3.4
4.1
0.71
0.07
0.13
0.12
0.34
5
6
9
10
6
26.7
4.4
14.7
7.7
15.7
20.0
0.0
0.0
4.5
0.0
20.0
84.9
67.0
65.0
55.7
0.0
0.0
0.0
3.6
14.3
33.3
0.0
3.8
3.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
4.2
3.6
4.5
0.0
0.0
0.2
0.2
0.0
0.0
0.0
6.2
7.4
2.3
4.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
2.4
4.1
8.6
<6.0
6.18.0
8.110.0
>10.0
0
0
0
0
2.9
3.5
2.7
3.4
0.19
0.09
0.13
0.20
3
7
6
5
15.0
15.5
22.4
8.7
75.0
78.8
70.0
72.6
0.0
1.4
1.7
0.0
0.0
0.0
0.0
0.0
0.0
1.6
0.0
0.0
0.0
0.0
0.0
0.0
0.0
2.3
0.0
13.0
0.0
0.0
0.0
0.0
0.0
0.0
0.2
0.0
0.0
0.0
0.0
0.0
0.0
0.4
1.0
5.7
<6.0
6.18.0
8.110.0
>10.0
12.4
16.5
8.9
10.4
2.7
2.3
2.2
2.1
0.17
0.17
0.21
0.45
9
9
11
6
50.0
45.7
42.1
32.6
26.4
12.9
17.4
16.3
1.2
0.2
0.9
0.0
0.0
0.0
0.0
0.0
6.2
0.0
7.2
8.6
1.2
1.9
0.0
0.0
2.4
6.6
5.6
0.0
0.7
0.4
1.2
0.7
0.0
0.0
0.1
0.0
0.0
3.3
7.2
14.0
0.2
2.5
0.9
0.0
Polycheate
Bivalve
Fish
Fish
eggs
Others
513
C. cuning 8.1-10.0cm
C. cuning <6.0cm
C. cuning 6.1-8.0cm
C. cuning >10.0cm
U. tragula <6.0cm
U. tragula 8.1-10.0cm
U. tragula 6.1-8.0cm
A. quadrifasciatus <6.0cm
A. quadrifasciatus 6.1-8.0cm
A. quadrifasciatus >10.0cm
A. quadrifasciatus 8.1-10.0cm
U. sulphureus >10.0cm
U. sulphureus 6.1-8.0cm
U. sulphureus <6.0cm
U. tragula >10.0cm
U. sulphureus 8.1-10.0cm
S. taeniopterus 15.1-20.0cm
S. taeniopterus 10.1-15.0cm
S. taeniopterus<10cm
L. splendens >10.0cm
L. splendens 8.1-10.0cm
L. splendens <8.0cm
C. planus >10.0cm
C. planus 8.1-10.0cm
C. planus 6.1-8.0cm
S. canaliculatus <6.0cm
S. canaliculatus 10.1-15.0cm
S. canaliculatus 8.1-10.0cm
S. canaliculatus 6.1-8.0cm
S. taeniopterus >20.0cm
S. canaliculatus >15.0cm
S. micropectorails 15.1-20.0cm
S. undrosquamis 10.1-15.0cm
S. undrosquamis <10cm
S. undrosquamis 6.1-8.0cm
S. undrosquamis >20.0cm
S. micropectorails >20.0cm
S. undrosquamis 15.1-20.0cm
S. undrosquamis <6.0cm
S. micropectorails 10.1-15.0cm
S. micropectorails <10.0cm
20
40
60
80
G1
G2
G3
G4
G5
100
Fig. 5. Cluster dendogram demonstrating trophic groups by sizes of 10 dominant sh species collected off coastal waters of Pattani and Narathiwas, Thailand between March and
November 2006. G1G5 trophic groups.
canaliculatus shifted from feeding on calanoid copepods and gammarid amphipods during the younger stages to polychaetes at the
larger size classes. There is a general tendency for marine shes to
start as zooplankton feeders, consuming large amounts of calanoid
copepods during their young stages, and shifting to other food
sources when they grow larger (Elliott et al., 2002). Crustacean
zooplankton is a major component in the diets of the young marine
shes throughout the world (e.g. Hales, 1987; Hyndes et al., 1997;
Platell et al., 1997; Amara et al., 2001; Hajisamae and Ibrahim,
2008). However, there are some truly zooplanktivorous species that
continue feeding on calanoid copepods even when they are larger
sizes such as Leiognathus splendens, Callionymus planus, Caesio
cuning and Apogon quadrifasciatus. Species demonstrating minimal
ontogenetic changes in feeding habits include Saurida undrosquamis, Saurida micropectoralis, Upeneus sulphureus, Leiognathus
splendens, Upeneus tragula and Caesio cuning.
In conclusion, shrimps were the most important dietary
component of shes in this ecosystem, followed by calanoid
copepods, shes and gammarid amphipods. Fish species concentrated on in this study could be divided into six different trophic
guilds, each with a different combination of food. They can be
further divided according to feeding functional groups (FMFG) into
four categories, namely; piscivores, zooplanktivores, zoobenthivores and miscellaneous/opportunists. Trophic guilds at each depth
zones vary between four and six, with four guilds consistently
formed at all zones; shrimp predators, piscivores, calanoid copepod
feeders and a combination of polychaete and other food feeders.
Shrimps, calanoid copepods, shes, polychaetes and gammarid
amphipods were fed on by demersal shes throughout the year.
Ontogenetic studies indicated that shes were likely to group
according to species rather than size classes. Some shes demonstrated ontogenetic changes in feeding habits. This information is
fundamental in understanding complex associations of shes and
sh communities and identifying groups of species that use similar
resources. These information serve as a reference for feeding
ecology of shes found in the lower part of the South China Sea,
important in decision making and management of sheries
resources of the region.
Acknowledgements
This work is dedicated to the late Mr. Mudtalib Daeuloh, my
stepfather, who had helped with the collection of sh samples for
this study with much enthusiasm until end of his life. The author
would like to thank the anonymous referees for useful comments
on the manuscript and to express his deep gratitude to Mr. Arun
Loh-hem, and the crew of Patabarat shing boat who had helped in
collecting sh samples throughout this sampling program. This
research was supported by the Thailand Research Council through
the Faculty of Science and Technology, Prince of Songkla University.
References
Amara, R., Laffargue, P., Dewarumez, J.M., Maryniak, C., Lagadere, F., Luczac, C., 2001.
Feeding ecology and growth of 0-group atsh (sole, dab and plaice) on
a nursery ground (Southern Bight of the North Sea). Journal of Fish Biology 58,
788803.
Angel, A., Ojeda, F.P., 2001. Structure and trophic organization of subtidal sh
assemblages on the northern Chilean coast: the effect of habitat complexity.
Marine Ecology Progress Series 217, 8191.
Bayley, P.B., 1988. Factors affecting growth rates of young tropical ood plain
shes: seasonality and density dependence. Environmental Biology of Fish 21,
127142.
Blaber, S.J.M., 1997. Fish and Fisheries of Tropical Estuaries. Chapman & Hall,
London, 388 pp.
Blaber, S.J.M., 2001. Tropical Estuarine Fishes; Ecology, Exploitation and Conservation. Blackwell Science, Oxford, 372 pp.
Blaber, S.J.M., Blaber, T.G., 1980. Factors affecting the distribution of juvenile
estuarine and inshore sh. Journal of Fish Biology 17, 143162.
Chong, V.C., Sasekumar, A., Leh, M.U.C., DCruz, R., 1990. The sh and prawn communities of a Malaysian coastal mangrove system, with comparisons to adjacent
mudats and inshore waters. Estuarine Coastal and Shelf Science 31, 703722.
Clarke, K.R., Gorley, R.N., 2001. Primer v5: User Manual/Tutorial PRIMER-E.
Plymouth, 91 pp.
Crowder, L.B., Cooper, W.E., 1982. Habitat structural complexity and the interaction
between bluegills and their prey. Ecology 63, 18021813.
Cruz-Escalona, V.H., Abitia-Cardenes, L.A., Campos-Davila, L., Galvan-Magana, F.,
2000. Trophic interrelations of the three most abundant sh species from
Laguna San Ignacio, Baja California Sur, Mexico. Bulletin of Marine Science 66,
361373.
Day, J.W., Jun, C.A., Hall, S., Kemp, W., Yanez-Arancibia, M., 1989. Estuarine Ecology.
John Wiley and Sons, New York, 558 pp.
Elliott, M., Hemingway, K.L., Costello, M.J., Duhamed, S., Hostens, K.,
Labropoulou, M., Marshall, S., Winkler, H., 2002. Links between sh and other
514
trophic levels. In: Elliott, M., Hemingway, K.L. (Eds.), Fishes in Estuaries.
Blackwell Science, Oxford, pp. 124216.
Elliott, M., Whiteld, A.K., Potter, I., Blaber, S.J.M., Cyrus, D.P., Nordlie, F.G.,
Harrison, T.D., 2007. The guild approach to categorizing estuarine sh assemblages: a global review. Fish and Fisheries 8, 241268.
Garrison, L.D., Linke, J.S., 2000. Dietary guild structure of the sh community in the
northeast United States continental shelf ecosystem. Marine Ecology Progress
Series 202, 231240.
Gerking, S.D., 1994. Feeding Ecology of Fish. Academic Press, California, 416 pp.
Hajisamae, S., Chou, L.M., 2003. Do shallow water habitats of an impacted coastal
strait serve as nursery ground for sh? Estuarine Coastal and Shelf Science 56,
281290.
Hajisamae, S., Chou, L.M., Ibrahim, S., 2003. Feeding habits and trophic organization
of the sh community in shallow waters of an impacted tropical habitat.
Estuarine Coastal and Shelf Science 58, 8998.
Hajisamae, S., Ibrahim, S., 2008. Seasonal and spatial variations of sh trophic guilds
in a shallow, semi-enclosed tropical estuarine bay. Environmental Biology of
Fish 82, 251264.
Hajisamae, S., Yeesin, P., Chaymongkol, S., 2006. Habitat utilization by shes in
a shallow, semi-enclosed estuarine bay in southern Gulf of Thailand. Estuarine
Coastal and Shelf Science 68, 647655.
Hales, L.S., 1987. Distribution, abundance, reproduction food habits, age and growth
of round scad, Decapterus punctatus, in the south Atlantic bight. Fisheries
Bulletin 85, 251268.
Hyndes, G.A., Platell, M.E., Potter, I.C., 1997. Relationships between diet and body size,
mouth morphology, habitat and movements of six sillaginid species in coastal
waters: implications for resource partitioning. Marine Biology 128, 585598.
Hyslop, E.J., 1980. Stomach contents analysis-a review of methods and their
application. Journal of Fish Biology 17, 411429.
Ikejima, K., Tongnunui, P., Medej, T., Taniuchi, T., 2003. Juvenile and small shes
in mangrove estuary in Trang province, Thailand: seasonal and habitat
preferences. Estuarine Coastal and Shelf Science 56, 447457.
Jacksic, F.M., 1981. Abuse and misuse of the term guild in ecological studies. Oikos
83, 8792.
Labropoulou, M., Papadopoulou-Smith, K.N., 1999. Foraging behaviour patterns of
four sympatric demersal shes. Estuarine, Coastal and Shelf Science 49, 99
108.
Lucena, F.M., Vaska Jr., T., Ellis, J.R., OBrien, C.M., 2000. Seasonal variation in the
diets of bluesh, Pomatomus saltatrix (Pomatomidae) and striped weaksh,
Cynoscion guatucupa (Sciaenidae) in southern Brazil: implications of food
partitioning. Environmental Biology of Fish 57, 423434.
Pianka, E.R., 1980. Guild structure in desert lizards. Oikos 35, 194201.
Platell, M.E., Sarre, G.A., Potter, I.C., 1997. The diets of two co-occurring marine
teleosts, Parequula melbournensis and Pseudocaranx wrighti, and their relationships to body size and mouth morphology, and season and location of
capture. Environmental Biology of Fish 49, 361376.
Platell, M.E., Potter, I.C., 2001. Partitioning of food resources amongst 18 abundant
benthic carnivorous sh species in marine waters on the lower west coast of
Australia. Journal of Experimental Marine Biology and Ecology 261, 3154.
Root, R.B., 1967. The niche exploitation pattern of the bluegrey gnatcatcher.
Ecological Monographs 37, 317350.
Ross, S.T., 1986. Resources partitioning in sh assemblages: a review of eld studies.
Copeia 1986, 352388.
de Sylva, D.P., 1975. Nektonic food webs in estuaries. In: Cronin, L.E. (Ed.), Estuarine
Research: Chemistry, Biology and the Estuarine System, vol. 1. Academic Press,
New York, pp. 420447.
Sasekumar, A., Chong, V.C., Leh, M.U., DCruz, R.D., 1992. Mangroves as a habitat for
sh and prawns. Hydrobiologia 247, 195207.
Schlosser, I.J., 1982. Trophic structure, reproduction success, and growth rate
of shes in a natural and modied headwater stream. Canadian Journal of
Fisheries and Aquatic Sciences 39, 968978.
Snyder, R.J., 1984. Seasonal variation in the diets of three spined stickleback, Gasterosteus aculeatus, in Contra Costa County, California. California Fishing Game
70, 167172.