Malayan Nature Journal 2014, 66(1and2), 132-148

Seaweed community of the Merambong shoal,

Sungai Pulai estuary, Johore
Muta Harah, Z.1,2, Japar Sidik, B.3, Natrah, F.M.I.1,2,
Emmclan, L.S.H.2, Wan Hazma, W.N.1, Nordiah, B.1

Abstract: A study on seaweed community encompassing diversity,

life form and percentage cover was carried out at the seagrass bed of
Merambong shoal, Sungai Pulai estuary, Johore. A total of 30 seaweed
species were recorded among seagrasses or on sand and mud substrates.
Chlorophyta was recorded with a total of 15 diverse species followed by
Rhodophyta 11 species and Phaeophyta 4 species. Seaweeds from the
three divisions; Chlorophyta, Phaeophyta and Rhodophyta did not show
any distinct zonation. Leaves of Enhalus acoroides, Halophilla ovalis and
Halodule uninervis provided refugia for many seaweeds e.g., Bryopsis
plumosa, Cladophora prolifera, Dictyota dichotoma, Hydropuntia edulis,
Gracilaria fisherii, G. salicornia and Ulva sp. Ulva compressa, Ulva
reticulata and Amphiroa fragilissima were attached to mollusc shells
or polychaete tubes. Jania decussato-dichotoma, Laurencia sp., and
Acanthophora spicifera were observed lying loosely among seagrasses or
attached to the sand or mud substrates. Seaweed occurrences differed with
months e.g., Ulva reticulata, Halimeda opuntia,Amphiroa fragilissimaand
Hydropuntia edulis were found abundant from June to August coinciding
with the rainy period. The observations showed seaweeds share and
have the adaptability to thrive and even compete with seagrasses,
occupying both available living e.g., seagrasses, polychaete tubes or
dead structures e.g., mollusc shells, as well as in sand to mud as supports.
Keywords: seaweed community, life form, percentage cover, sub-tidal shoal,
Merambong

1 Department of Aquaculture, Faculty of Agriculture, Universiti Putra Malaysia, 43400

UPM Serdang, Selangor Darul Ehsan, Malaysia.
2 Institute of Bioscience, Universiti Putra Malaysia, 43400 UPM Serdang, Selangor
Darul Ehsan, Malaysia.
3 Faculty of Agriculture and Food Sciences, Universiti Putra Malaysia Bintulu Sarawak
Campus, 97008 Bintulu, Sarawak, Malaysia
132

INTRODUCTION
Malaysia has an extensive coastline comprising of sandy bays,rocky
shores and alternate with mudflats and mangroves.

Mangroves dominate the west coast of Peninsular Malaysia
while sandy shores and bays, and rocky shores form the conspicuous
coastline of the east coast of Peninsular Malaysia.
Sandy and rocky beaches characterise the coastlines of Sabah and
Sarawak. Coral reefs and seagrass beds or meadows tend to be the
underwater landscapes fringing most off-shore islands and in sub-tidal
shoals. These diverse ecotones provide shared habitats for seaweed and
seagrass species. Although seagrasses and seaweeds occur in similar
varied habitats but by species numbers, seagrasses are found to be much
less and their presence are from patches to dense extensive coastal beds or
meadows in coastal areas between mangroves and corals, from low tide
level to the coral reef fringe and sub-tidal areas between the corals and the
semi-open sea (Japar Sidik et al. 1995). The reported seaweed resources
of Malaysia are primarily focused on those communities of rocky shores,
coral reefs and mangroves (Johnson 1967; Sivalingam 1977; Crane
1981; Arumugam 1981; Aikanathan and Sasekumar 1994; Japar Sidik et
al. 1997;Phang 1998; Japar Sidik and Muta Harah 1999; Muta Harah et
al. 2006, 2007; Wong et al. 2010, 2012). It is noteworthy to mention here
that seaweed population or communities are less described in seagrass
ecosystems (Sasekumar et al. 1989,Japar Sidik et al. 1996), perhaps
being less prominent and adaptable in such ecosystems as compared to
rocky shores, coral reefs and mangroves with the availability of hard
substrates (Norashikin et al. 2013).

Seaweeds similar to seagrasses and phytoplanktons form the
important primary producers of a shallow marine environment. As
terrestrial plants, seaweeds absorb light energy in photosynthetic
processes that chemically combine water and carbon dioxide to produce
organic compounds. Through the process of photosynthesis, the aquatic
environment is oxygenated. Nutrients are absorbed through incorporation
into their cells that constitute the plant body. Substances released by the
plants in turn are consumed by heterotrophic bacteria (Wilkinson and
Buddemeir 1994) and the plants themselves form the primary food for
herbivores and are passed on through the food chains and webs to be
utilized by various consumers comprising of omnivores and carnivores.
Seaweeds when present contribute to the productivity and biodiversity of
the habitats similar to tropical rain forests, mangroves and coral reefs. In
a naturally balanced ecosystem, seaweeds and seagrasses share resources
such as nutrients and spaces. The pressures due to development in
133

the coastal areas have impacted and caused seaweeds and seagrasses
to compete resulting in excessive growth of seaweeds over the latter,
thus deteriorating the balanced ecosystem. This paper presents the
available research information and observations through various studies
conducted in 1996 and from June to August 2013 at the 26.3 hectare
Merambong seagrass shoal in the Sungai Pulai estuary of Johore. The
information presented covers seaweed species distribution, composition,
coverage, lifeforms and the seaweed behaviour in the seagrass bed of the
Merambong shoal of Sungai Pulai, Johore.
MATERIALS AND METHODS
Field data were obtained during low tides from June to August 2013
when the Merambong shoal (1o 19 59.86 N, 103o 39 0.13 E), Sungai
Pulai estuary, Johore was at its highest level of exposure. Samplings
of seaweed were obtained to compile data on species composition,
coverage, species assemblage using line transect and 50 cm x 50 cm
quadrat with 25 sub-divisions at 10 m interval (Figure 1) across the shoal
following the method developed by Saito and Atobe (1970). Seaweed
species, coverage and the corresponding indices (referring to Table 1)
in each of the 25 sub-divisions were recorded. The parameters obtained
from each quadrat with respect to coverage (C; expressed as %) is used
to compute the area in the substrate occupied by the species. The index
numbers: 5, 4, 3, 2, 1 are used for recording data in the field as indicated
in Table 1. C (%) = (qn5 x c5) + (qn4 x c4) + (qn3 x c3) + (qn2 x c2) + (qn1
x c1) = (qn5 x 3) + (qn4 x 1.5) + (qn3 x 0.75) + (qn2 x 0.375) + (qn1 x
0.1875); where qnn is the number of sub-divisions in which a species
appeared to have the corresponding coverage area described in Table 1.
Representative seaweed samples were cleaned in situ with seawater to
remove sand, debris and shells, placed in labelled plastic bags, preserved
in 5% formalin-seawater solution and transferred to the laboratory at the
Faculty of Agriculture, Universiti Putra Malaysia, Serdang, Selangor,
Malaysia for taxonomical identification following the references of
Lewmanomont and Ogawa (1995), Trono (1997, 2004) and Tsutsui et
al. (2005).
RESULTS AND DISCUSSION
Diversity distribution
General observations indicated that seaweeds were found across and
usually scattered within the shoal area. All three major divisions of
134

seaweed ; Chlorophyta, Phaeophyta and Rhodophyta were recorded at the

shoal (Table 2) and their species overlapped with no distinct zonation. The
distributional trend could be attributed to; (i) some of the seaweed might
have changed their morphology with some modifications depending on
their required environment (Norton et al. 1981), (ii) seaweeds could
have been swept to other areas and might still live (Chapman 1979) and
(iii) some seaweeds might have lost their attachment from the ordinary
substrates and were swept away as drift seaweed (Khew 1978).

Sasekumar et al. (1989) reported that other than seagrasses,
abundant seaweeds such as Caulerpa lentillifera, Halimeda tuna,
Chaetomorpha sp., Dictyota dichotoma, Gracilaria salicornia, Amphiroa
fragillisima and Acanthophora spicifera were also found in the same area.
Our present study recorded at least 30 species of seaweeds, 15 belonging
to the division Chlorophyta, 11 to Rhodophyta and 4 to Phaeophyta.
Comparing to the records of seaweed collection for Merambong shoal
by Japar Sidik et al. (1996), this present study observed 14 species
as new records for the shoal. Four (4) seaweed species, Cladophora
patentiramea, Cladophora fascicularis, Hypnea esperi and Gracilaria
textorii were not observed in the present study for the shoal (cf. Japar
Sidik et al. 1996 and the present study, Table 2). There were less
diverse seaweed species in the seagrass shoal relative to the 43 species
reported for the rocky-sandy area of Tanjung Batu and Kampung Kuala
Nyalau, Sarawak (Wong et al. 2012) as well as Teluk Kemang and Teluk
Pelanduk, Negeri Sembilan (Norashikin et al. 2013). The rocky area
with diverse microhabitats (tide pools) and the availability of stable hard
substrates provide favourable conditions for the occurrence and diversity
of seaweed species (Chapman and Chapman 1973). The seagrass shoal
(sand to mud substrates) area lacks hard structure which is one of the
reasons for the relatively low diversity of seaweed or different seaweed
species. Although less diverse, seaweeds e.g., Ulva reticulata, Halimeda
opuntia,Amphiroa fragilissimaand Hydropuntia edulis were found in
abundance from June to August coinciding with the rainy period (Figure
2). This phenomenon of seaweed invasion occurred more frequently than
previously observed in the Merambong shoal (Japar Sidik et al. 2006).
Life form
The present study noted that various types of substrata were colonized
by seaweeds, from sand, mud, animals such as snails, bivalves, other
seaweeds and seagrasses. The availability of substratum obviously
determined the species composition of seaweeds (Norton et al.1981).
Based on mode of attachment on substrate types, seaweed life form
135

categories is shown in Table 2 as; (1) epilithic (live on rocks or dead

coral), (2) epipelic (attached to mud and sand), (3) epizoic (attached to
mollusc shells or polychaete tubes), (4) epiphytic (attached to seagrass or
seaweed) and (5) drift (detached or floating seaweed).

Both attached and drift seaweeds are the important components
of the seagrass communities of Merambong. Attached macroalgae
comprised epipelic and epiphytic (Table 2) such as Avrainvillea erecta,
Caulerpa sertularioides,Chaetomorpha spiralis, Halimeda macroloba,
Halimeda opuntia and Enteromorpha clathrata, Stypopodium zonale,
Amphiroa fragilissima, Jania decussato-dichotoma. Leaves of Enhalus
acoroides, Halophila ovalis and Halodule uninervis provided refugia
for seaweeds e.g., Bryopsis plumosa, Cladophora prolifera, Dictyota
dichotoma, Hydropuntia edulis, Gracilaria fisherii, G. salicornia and
Ulva sp.Acanthophora spicifera and Gracilaria fisherii are ephilithic,
living on hard substrata such as rock. Ulva intestinalis, Ulvareticulata
and Acanthophora spicifera are drift seaweeds, which do not attach
to any substrate but remain free and can survive in the environment
(Chapman 1979). Ulva intestinalis, Ulva compressa and Acanthophora
spicifera were observed as having more than two life forms. The highest
percentage of seaweed category was epipelic (54.63-76.67%, Table 3)
which reflected life form suitability to the substrate comprising of mud
and sand present in that area. Epilithic seaweed such as Acanthopora
spicifera represented a small group attached to rocks and dead corals.
Truly rhizophytic seaweeds were common to abundant in seagrass beds
such as Avrainvillea erecta, Halimeda macroloba,Stypopodium zonale
(possessed pseudobulbous holdfasts) and Caulerpa spp. (possessed
stolons and rhizoids). These special attachment structures enabled them
to affix to soft substrates such as sand and mud (Brouns and Heijs 1991).
In this present study, there were very few types of hard substrata in the
shoal for other seaweeds to attach. Due to this, seaweeds were attached
to seagrasses, mollusc shells or polychaeta tubes or were found loose
amongst seagrasses. Similar observation has also been reported by
Taylor (1957). Their presence in seagrass beds or in systems that lack
any form of habitat structure may be important sources of cover for
smaller animals (Orth et al. 1991).
Percentage cover
The transect and quadrat analysis of seaweed percentage cover is also
shown in Table 4. Five (5) seaweed species commonly found in the
Merambong shoal, Acanthopora spicifera, Amphiroa fragilissima,
Caulerpa sertularoides, Hydropuntia edulis and Ulva reticulata have
136

higher percentage cover as compared to other seaweed species. These

seaweeds have special attachment structures that can be used for
attachment to soft substrates. For examples Caulerpasertularoides and
Ulva reticulata were attached to sandy substrate (non-epiphytic), as
they possessed rhizomaceous features (creeping stolons and rhizoids)
enabling them to expand in the environment by vegetative means. In
addition they can rapidly grow and is able withstand several hours of
exposure to air during low tides. Studies by McConnaughey (1978) and
Lobban et al.(1985) indicated that muddy and sandy areas support fewer
seaweeds since most species cannot anchor as they tend to attach to some
fixed and firm structures such as rocks, shells or even other marine plants
(Teo and Wee 1983).
CONCLUSION
Thirty (30) seaweed species were found coexisting with seagrasses in
the Merambong shoal. They utilized the available substrates such as soft
substrates (sand, mud), dead structures to living organisms (gastropods,
bivalves) and plants (other seaweeds and seagrasses). They possess
suitable attachment structures and growth habits which enables them to
expand in the environment of the Merambong seagrass shoal.
ACKNOWLEDGEMENTS
We would like to thank the Vice-Chancellor, Universiti Putra Malaysia,
for providing the facilities and logistics for this study. This study was
funded by Ministry of Education, Malaysia (KPM), and in collaboration
with the Japan Society for the Promotion of Science (JSPS) Asian CORE
program and the Establishment of Research Education Network on
Coastal Marine Science in Southeast Asia (COMSEA).

137

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139

Table 1. Indices of the degree of plant cover and its representative

multiplier. Source: Saito and Atobe (1970)
Indices
5
4
3
2
1

Degree of seaweed cover, cn

Multiplier, qn

Covering 1/2 to1 of substrum surface

Covering 1/4 to 1/2of substrum surface
Covering 1/8 to 1/4 of subtrum surface
Covering 1/16 to 1/8of subtrum surface
Covering less than 1/16 of subtrum surface

3.0000
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140

Table 2. Occurrence of seaweeds species from June to August 2013 at the Merambong shoal. Epl-epilithic, Eppepipelic, Epz-epizoic, Epy-epiphytic and D-drift. Substrate: 1-Dead coral, 2-Mud, 3-Sand, 4-Other seaweed,
5-Seagrass, 6-Shell/Bivalve, 7-Polychaete tube, 8-Drift.*New seaweed species recorded at the shoal, a-Japar Sidik
et al. (1996)
No

Division/Family/Species

1996a
Jun.

2013
Jul.

Substrate

Life forms

3, 4, 5

Epp, Epy

++
++
++

2, 3
2, 3
2, 3, 5

Epp
Epp
Epp, Epy

4, 5
4, 5
4, 6
3, 4

Epy
Epy
Epy, Epz
Epp, Epy

Aug.

CHOLOROPHYTA
Bryopsidaceae
1.

Bryopsis plumosa (Hudson) C. Agardh

++

++
++
++

Caulerpaceae
2.
3.
4.

Caulerpa manorensis Nizamuddin

*Caulerpa taxifolia (M. Vahl) C. Agardh
Caulerpa sertularioides (S.G. Gmelin) M.A. Howe

++

Cladophoraceae
5.
6.
7.
8.

Cladophora fascicularis (Mertens ex C. Agardh) Ktzing

Cladophora patentiramea (Montagne) Kutzing
*Cladophora prolifera (Roth) Kutzing
Chaetomorpha spiralis Okamura

+
+
++

++
++

++

++
+

++
++

++
++

++
++

3, 4
3, 4, 5

Epp, Epy
Epp, Epy

++

Epp

Halimedaceae
9.
10.

*Halimeda macroloba Decaisne

*Halimeda opuntia (Linnaeus) Lamourox
Udoteaceae

11.

Avrainvillea erecta (Barkeley) A. Gepp and E. Gepp

+
141

++

No
12.

Division/Family/Species

1996a
Jun.
++

*Udotea flabellum (Ellis et Solander) Howe

2013
Jul.

Aug.
++

Substrate

Life forms

Epp

Ulvaceae
13.
14.

Enteromorpha clathrata (Roth) Greville

*Ulva intestinalis (Linnaeus) Nees

++
++

++
++

3, 4
3, 6, 8

15.

Ulva reticulata Forsskal

++

++

3, 6, 7, 8

16.
17.

*Ulva compressa Linnaeus

Ulva sp.

+
+

6, 7
5, 6, 8

Epp, Epy
Epp, Epz,
D
Epp, Epz,
D
Epz
Epy, Epz, D

+
+

++
++
++
++

++
++
++
++

3, 4, 5
3, 4, 5
3, 4, 5
3

Epp, Epy
Epp, Epy
Epp, Epy
Epp

Epl, Epp,
Epy, Epz
Epp, Epy,
D
Epl, Epp

PHAEOPHYTA
Dictyotaceae
18.
19.
20.
21.

*Dictyota bartayresii Lamouroux

*Dictyota cervicornis Kutzing
Dictyota dichotoma(Hudson) Lamourox
Stypopodium zonale (Lamourox)
RHODOPHYTA
Corallinaceae

22.

Amphiroa fragilissima (Linnaeus) Lamourox

++

++

++

23.

Amphiroa rigida J.V. Lamouroux

++

++

++

++

1, 3, 4, 5,
6, 7
3, 4, 5, 8

24.

*Jania decussato-dichotoma (Yendo) Yendo

++

++

++

1, 3

142

No

Division/Family/Species

1996a
Jun.

2013
Jul.

Aug.

+
+

+
+
+

++
++
++

Substrate

Life forms

2, 3, 4, 5
8

Epp, Epy
D

+
++

++
++

++

3, 4, 5
3, 4, 5
6
3

Epp, Epy
Epp, Epy
Epz
Epp

++

++

3, 6

Epp, Epz

++

++

1, 3, 5, 8

++
++

3, 5
3, 5

Epl, Epp,
Epy, D
Epp, Epy
Epp, Epy

Florideophyceae
25.
26.

Hypnea cervicornis J. Agardh

Hypnea esperi Bory de Saint-Vincent
Glacilariaceace

27.
28.
29
30.

Gracilaria fisherii (B.M Xia and I.A. Abbott)

Gracilaria salicornia (C. Agradh) Dawson)
Gracilaria textorii (Suringar) De Toni
*Hydropuntia edulis (S.G. Gmelin) Gurgel and Fredericq
Halymeniaceae

31.

*Halymenia floresia (Clubio) C. Agardh

Rhodomelaceae

32.

Acanthophora spicifera (Vahl) Borgesen

33.
34.

*Laurencia papillosa (C.Agardh) Greville

*Laurencia sp.

++
++

143

Table 3. Comparison of seaweed life forms between the Merambong shoal and the rocky shores of Malaysia. TKTeluk Kemang, TP-Teluk Pelanduk
Area

Bintulu, Sarawak

Port Dickson,
Negeri Sembilan

Merambong shoal, Johore

Rocky shores
Reference

Seagrass bed (shoal)

Muta
Harah et
al. (2006)

Wong et
al. (2010)

Wong et
al. (2012)

Norashikin et al.
(2013)

Japar Sidik
et al. (1996)

Jun. 2013

Jul. 2013

Aug. 2013

10
6
19
35

7
5
11
23

14
7
22
43

19
12
12
43

12
2
10
24

15
4
10
29

9
2
9
20

13
4
10
27

33
(54.10%)

45.71% (TK)

7
(3.24%)

6
(6.06%)

4
(3.33%)

5
(8.20%)
2
(3.28%)
20
(32.79%)
1
(1.64)

12.86% (TK)
21.15% (TP)
24.29% (TK)
9.67% (TP)
8.57% (TK)
17.31% (TP)
8.57% (TK)
13.46% (TP)

3
(14.29%)
2
(9.52%)
11
(52.38%)
5
(23.81%)

118
(54.63%)
31
(14.35%)
39
(18.06%)
21
(9.72%)

74
(74.75%)
13
(13.13%)
6
(6.06%)
0

92
(76.67%)
11
(9.17%)
10
(8.33%)
3
(2.5%)

DIVISION
Chlorophyta
Phaeophyta
Rhodophyta
Total
LIFE
FORM
Epilithic
Epipelic
Epizoic
Ephiphytic
Drift

38.46% (TP)

144

Table 4. Seaweed cover percentage based on the method of Saito and

Atobe (1970) quadrat sampling from June to August 2013.
Cover (%)

Quadrat
interval

Seaweed

Jul.

Aug.

0m

Acanthopora spicifera
Amphiroa fragilissima
Caulerpa sertularoides
Hydropuntia edulis
Ulva intestinalis
Ulva reticulata

0.38
1.69
0.19
0.56
10.31

14.25
0.19
0.19

13.31
0.75
0.38

34.13
0.19
-

10 m

Amphiroa fragilissima
Ulva intestinalis
Ulva reticulata

32.25
0.19

14.25
0.56
-

17.25
-

6.75
-

20 m

Amphiroa fragilissima
Ulva reticulata
Caulerpa sertularoides

5.25
0.19
-

8.63
0.38
-

0.75
-

15.38
0.38
-

30 m

Acanthopora spicifera
Amphiroa fragilissima
Caulerpa sertularoides
Ulva reticulata

0.19
-

0.19
0.19

0.38
-

0.94
32.63
0.19
0.75

40 m

Amphiroa fragilissima
Ulva reticulata

0.38

0.56
0.19

3.19
-

3.56
-

50 m

Acanthopora spicifera
Amphiroa fragilissima
Ulva intestinalis
Ulva reticulata

0.56
0.38
0.38
0.19

2.06
0.19

0.56
-

17.25
0.38

60 m

Acanthopora spicifera
Amphiroa fragilissima
Caulerpa sertularoides
Ulva reticulata

6.75
-

0.38
-

0.94
0.94
0.38

0.75
0.19
0.94

70 m

Acanthopora spicifera
Amphiroa fragilissima
Dictyota dichotoma

2.25
0.75
-

2.06
-

1.69
1.12
0.56

0.56
-

Jun. (A) Jun. (B)

145

Quadrat
interval

Seaweed

Cover (%)
Jun. (A) Jun. (B)

Jul.

Aug.

Halymenia floresii
Ulva reticulata

1.5
1.69

1.5

80 m

Acanthopora spicifera
Amphiroa fragilissima
Hydropuntia edulis
Ulva reticulata

0.75
0.94
0.19
0.56

0.19
-

0.56
0.19

1.13
0.38
-

90 m

Acanthopora spicifera
Amphiroa fragilissima
Halymenia floresii

0.38
2.63
-

0.38

0.19
0.94
-

100 m

Acanthopora spicifera
Amphiroa fragilissima
Hydropuntia edulis

0.38
0.75
-

0.56
-

0.38
0.38

110 m

Acanthopora spicifera
Amphiroa fragilissima
Ulva reticulata

0.19
0.94
0.75

120 m

Acanthopora spicifera
Amphiroa fragilissima

0.19
0.19

146

Figure 1. A line transect laid across the shoal and a 50 cm x 50 cm

quadrat placed at 10 m interval adopted for seaweed sampling.

Figure 2. Common seaweed life forms at the Merambong shoal, (a)

epipelic (e.g., Caulerpa taxifolia attached to mud and sand), (b) epizoic
(e.g., Ulva intestinalis attached to mollusc shells), (c) epiphytic (e.g.,
Halimeda opuntia attached to other seaweeds or seagrasses) and (d)
drift (e.g., Ulva reticulate which do not attach to any substrate but
remains free and can still survive in the environment).
147

Figure 3. Seaweed invasion occurred more frequently than previously

observed in the Merambong shoal (a)Amphiroa fragilissima (b) Ulva
reticulata (c) Halimeda opuntia, and (d) Caulerpa taxifolia.

148