Malayan Nature Journal 2014, 66(1and2), 1-14

Seagrass shoals of Sungai Pulai Estuary, Johor

B. JAPAR SIDIK1, Z. MUTA HARAH2,3 and A. ARSHAD2

Abstract: Seagrasses occur at sites scattered along the coastlines of

the West and East coasts of Peninsular Malaysia, Sabah and Sarawak,
usually growing in diverse environments in association with shallow
inter-tidal areas, mangroves, coral reefs, semi-enclosed lagoons and
shoals. Seagrass beds are site specific and we include specific examples
especially for seagrass shoals at Merambong, Tanjung Adang Laut and
Seluyong of Sungai Pulai estuary to illustrate their diverse flora and fauna
as well as their significant values to the coastal communities in Malaysia.
In these habitats, seagrass beds or meadows have distinct species
assemblage. Seagrass beds thrives at depths of 2-3 m on calcareous
sandy-mud sub-tidal shoals of Tanjung Adang Laut and Merambong,
supporting eight species of seagrasses; Enhalus acoroides, Halophila
ovalis, H. minor, H. spinulosa, Thalassia hemprichii, Cymodocea
serrulata, Halodule pinifolia, H. uninervis and Syringodium isoetifolium.
Beds of two species, Halophila ovalis-Halophila spinulosa inhabit the
mud substrate of the Seluyong shoal, the shallow sub-tidal area adjacent
to the fringing mangroves. Since 2002 the extensive seagrass beds of
Merambong and Tanjung Adang, in particular, have deteriorated to a
greater extent due to coastal development. This resulted in the significant
loss of seagrass species Halophila minor, Cymodocea rotundata and
Syringodium isoetifolium. This review gives information on the seagrass
species diversity, seagrass linkages to the diversity of invertebrates and
vertebrates, their values to the coastal communities and caustic factors
involved in the decline of seagrasses and their associated resources.
Keywords: Seagrasses, species diversity, Merambong-Tanjung Adang shoals,
Seluyong shoal, Malaysia

1 Faculty of Agriculture and Food Sciences, Universiti Putra Malaysia Bintulu Sarawak
Campus, 97008 Bintulu, Sarawak, Malaysia.
2 Department of Aquaculture, Faculty of Agriculture, Universiti Putra Malaysia, 43400
UPM Serdang, Selangor, Malaysia.
3 Institute of Bioscience, Universiti Putra Malaysia, 43400 UPM Serdang, Selangor,
Malaysia.
1

INTRODUCTION
Seagrasses are submerged monocotyledonous plants forming patches or
dense extended coastal beds or meadows. They constitute complex and
productive ecosystems occurring both in temperate and tropical seas and
are known for ecological roles in estuaries and coastal zones throughout
the world, which is well established (Larkum et al. 1989). They are
closely related to coastal zone economics, due to a highly primary and
secondary productive tropical ecosystem (Hemminga and Duarte 2000;
Bronwyn 2006). Similar to tropical rain forests, mangroves and coral
reefs, seagrass communities when present, contributes to the biodiversity
of the area (Johnstone 1982; Brouns and Heijs 1991; Poineer et al. 1987;
Bell and Pollard 1989; Larkum et al. 1989; Verheij and Erftemeijer
1993; Japar Sidik et al. 2006). Healthy seagrass communities provide
food for herbivores, create habitats and nurseries for invertebrates,
fishes turtles and birds, stabilize sediments, recycle nutrients, and are
therefore highly productive (Dawes et al. 1995). This is attributed to
their structural complexity and productivity (Kikuchi 1980; Simenstad
1994). Huge extensive meadows are always vital to near shore fisheries,
acting as natural retreats, source of food and shelter for juvenile fishes,
marine invertebrate, marine reptiles and mammals. It is well accepted
that seagrass significantly affect the chemical, physical and biological
environment, and therefore considered as an ecological engineer
(McKenzie 2008). The ecological and environmental roles of seagrasses
are evident in supporting a highly biodiverse marine coastal ecosystem.

Malaysias coastline is around 4,800 km long, stretching along
Peninsular Malaysia, Sabah and Sarawak covering much of the southern
part of the South China Sea. Along and adjacent this coastline stretch
are conspicuous coastal habitats including mudflats, mangroves and
coral reefs. Sharing these habitats are seaweeds and the lesser known
marine plants as well as seagrasses. Similar to many countries such
as Indonesia, Philippines, Thailand and Vietnam, the local coastal
inhabitants of Malaysia utilize seagrass areas (e.g., Merambong-Tanjung
Adang Laut) to sustain their living (Ogawa et al. 2011). Simultaneously,
coastal areas with seagrasses (e.g. Merambong-Tanjung Adang seagrass
shoals, Muta Harah and Japar Sidik (2011) are being subjected to
increased development and recreational pressures (Japar Sidik et al.
2006). Despite the stress seagrass ecosystems face, which include
anthropogenic activities leading to their decline and disappearance, the
local coastal inhabitants continue to use these areas for their sustenance.
Therefore, the livelihood of coastal communities are closely linked
with the seagrasses. This paper presents the information on seagrass
species, distribution and significance of seagrass shoals in Sungai Pulai
estuary, Johor. We believe by knowing the resources, one will appreciate
seagrasess and their associated resources contribution, the role they play
2

and the recognition that seagrasses sustain and supports life within itself
and other coastal marine ecosystems as well as the coastal inhabitants.
MATERIALS AND METHODS
This paper presents the available information on seagrass resources
and associated flora and fauna through various studies over the past 15
years (1994 to 2013) on the inventory of marine plants and associated
organisms. The information covers aspects from distribution, taxonomy,
diversity, utilization as well as natural and human activities which are
derived from published sources by the authors e.g., Sasekumar et al.
(1989) and Choo and Liew (2005) given in the reference section of this
paper.
RESULTS AND DISCUSSION
Seagrass shoals descriptions and seagrass species
The seagrass beds or meadows in the Sungai Pulai estuary are restricted
to sub-tidal zones along the shoreline or shoals that are submerged most
of the time. Seagrass beds are of mixed species and site specific. We
include specific examples to illustrate the types of seagrass beds found
in the estuary. The overview of the seagrass distribution and description
in this section is given and they sometimes form diverse extensive
communities.

The Sg. Pulai estuary is an extensive mangrove river system
located between latitudes 1o 15-1o 30 N and longitudes 103o 27-103o
36 E (Figure 1). The estuary reaches approximately 2.83 km wide and
ranges from 2 m to 15 m deep from its mid-point. Extensive seagrass
beds colonize the shallow depths of 2-3 m on both muddy banks from the
lower reaches right up to the upper parts of the middle reaches near Sg.
Jeram. The dominant species comprise of Enhalus acoroides, Halophila
ovalis, H. spinulosa and Syringodium isoetifolium (Sasekumar et al.
1989). The water around the Sg. Pulai estuary has a salinity of 31-32
psu, temperatures 29-29.5oC, pH 5.6-6 and visibility 0.5-1 m (Japar
Sidik et al. 1996).

In 1992 the three shoals in the estuary area, Merambong, Tanjung
Adang (Tanjung Adang Laut and Tanjung Adang Darat) and Tanjung
Kupang were discovered (Japar Sidik 1994). They were vegetated by
seagrasses occurring at depths of -1.2 to -1.5 m MSL, visible only during
the lowest spring tide. These seagrass meadows grew on calcareous sandy
mud, varying in size from 120 m to 1 km in length and 50 m to 120 m
in width. Six species, Thalassia hemprichii, Halophila minor, Halodule
3

uninervis, H. pinifolia, Cymodocea serrulata and C. rotundata including

those species reported by Sasekumar et al. (1989) were recorded for
the three shoals. This brings the total number of seagrass species in
the Sg. Pulai estuary area to ten, giving it the distinction of having the
highest number of seagrass diversity in Peninsular Malaysia (Japar Sidik
et al. 1996). The seagrass bed of Tanjung Adang Darat was completely
destroyed in 2003 due to filling and dredging for land expansion in lieu
of port development (Japar Sidik et al. 2007). The Tanjung Kupang shoal
(Japar Sidik 1994), which previously supported three seagrass species;
E. acoroides, H. ovalis and H. spinulosa, at present is relatively bare and
indulated with thick deposits of soft mud.

The Merambong (Figure 2) and Tanjung Adang Laut (Figure
3) shoals of calcareous sandy-mud substrates support communities
dominated by mixed species E. acoroides, H. spinulosa and H. ovalis
(Figure 4). This association nearly always occupy the zone exposed
during low spring tides (Figure 2) and occur at depth -1.2 to -1.5 m MSL.
The edge zone, -1.5 to -1.8 m MSL, is generally not emergent even during
the lowest low tide (-1.5 m MSL), and consists of an association of
Cymodocea serrulata or Thalassia hemprichii, Syringodium isoetifolium
and Halodule uninervis, in order of dominance. These species are less
resistant to dryness and exposure, and therefore tend to be in submerged
conditions. This same zone, when not occupied by the species, is generally
bare with isolated patches of H. spinulosa, Halophila minor, Halodule
uninervis, H. pinifolia or Cymodocea rotundata (present previously but
now no longer observed). In the deeper zone (-1.8 to -2.0 m MSL) sparse,
isolated patches of E. acoroides, H. ovalis and S. isoetifolium were
found. Cymodocea rotundata was only recorded for Merambong shoal
and is rare. A total of nine and eight species of seagrasses were recorded
for Merambong shoal and Tadang Adang, respectively previously (Japar
Sidik et al. 1996, 2006; Japar Sidik et al. 2001). In contrast, eight
species of seagrasses were recorded in 2012 and seven species in 2013 at
Merambong and Tanjung Adang Laut shoals respectively and two species
at Seluyong shoal (Figure 5, Table 1). The decrease of seagrass to seven
and eight species in 2012 and 2013 might be due to stressful conditions
and physical oceanographic stress at Merambong-Tanjung Adang shoal
due to the effect of land filling and extension for port development at the
vicinity together with the deteriorating water quality during monsoons
(south-west monsoons), creating a dynamic and fluctuating environment.
Only seagrasses characterized by linear leaves e.g., E. acoroides, T.
hemprichii, C. serrulata, H. pinifolia and H. uninervis and flaccid leaves
e.g., H. ovalis and H. spinulosa are unique plants that can survive under
such extreme stressed conditions. Halophila ovalis (big- and small4

leaved forms) and H. pinifolia are prolific seed producers designed to

maintain population in a seasonally fluctuating environment. According
to Kenworthy et al. (1989) reproduction by seed enhances survival in
deeper water allowing these seagrasses to persist through unfavourable
environmental conditions and poor monsoonal water quality by forming
a temporary seed bank at the sediments. Species such as H. pinifolia
can tolerated in a wide range of depths (Philips and Menez 1988) and
can extend into waters of at least 14 m depth (Duarte 1991). It can
also be found in all kinds of substrata, from coarse sand to almost liquid
mud (Coppejans et al. 1992). Merambong-Tanjung Adang Laut shoal
supports a greater diversity of seagrasses as compared to other seagrass
areas in Malaysia.
Other marine plants at the shoals
Seagrass meadows of Merambong and Tanjung Adang also supports
macroalgae or seaweed population. Sasekumar et al. (1989) reported that
at Sg. Pulai, aside from seagrasses, abundant algae such as Caulerpa
lentillifera, Halimeda tuna, Chaetomorpha sp., Dictyota dichotoma,
Hydropuntia edulis, Gracilaria salicornia, Amphiroa fragillisima and
Acanthophora spicifera were recorded. For Merambong-Tanjung Adang
area, 25 species of macroalgae were recorded. Both attached and drift
macroalgae are the important components of the seagrass communities
of Merambong and Tanjung Adang shoals. Although quite a number
of macroalgal species are found, only a few are characteristic species.
Truly rhizophytic macroalgae are common to abundant; Avrainvillea
erecta, Caulerpa spp., Udotea occidentalis are attached into the sandy
or sandy-mud substrates. Epiphytic macro-algae; Bryopsis plumosa,
Ceramium affine, Chaetomorpha spiralis, Cladophora spatentiramea,
C. fascicularis, C. fuliginosa, Dictyota dichtoma, Hypnea cervicornis, G.
fisherii and G. salicornia are attached to seagrasses while Ulva calthrata
and Gracilaria textorii are attached to mollusc shells or polycheate
tubes. There are very few types of hard substrata in the shoals for other
macroalgae to attach to. As such, macroalgae get attached to seagrasses,
mollusc shells or polychaeta tubes or are found loose amongst seagrasses.
Drift macroalgae, such as Acanthophora spicifera, Amphiroa rigida, A.
fragilissima, Hypnea esperi and Ulva spp. are found loose amongst the
seagrasses and may continue growing (Japar Sidik et al. 1996). 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). Detailed updated records of macroalgae have been described
by Muta Harah et al. (2014, this volume).
5

The declining of seagrass beds and their resources

There is no information in the form of historical maps or aerial photographs
that can be used to determine the timeline of the loss of seagrass beds.
The losses reported here have been made through observations during
repeated visits to the various seagrass sites. The Sungai Pulai seagrass
beds Merambong (30 ha in 1996, now 26.3 ha), Tanjung Adang Laut
(40 ha in 1996, now 9.44 ha) and Tanjung Adang Darat (42 ha in 1996)
are now completely destroyed and showing little recovery (Japar Sidik
et al. 2007). They were facing risk from the shipping port development
involving dredging of shallow passageways and land reclamation for new
facilities, both causing an increase in the suspended solids in the water
column. In Merambong and Tanjung Adang Laut, localized losses were
observed with the death of H. ovalis, clearly smothered by sand build up.
In addition dense overgrowth of the macroalgae Gracillaria fisherii and
Amphiroa fragilissima caused the seagrasses in the area to die with slow
regrowth of seagrasses. These short-term pulse events either in isolation
or combined in the shoals were output from runoff and direct discharge of
waste to Sg. Pulai river systems and adjacent areas especially after heavy
rainfall. Increased turbidity has significantly affected the water quality
of the area and suspected to have been directly linked to the reduction in
fishery stocks. The expanded land use of different economic activities has
caused significant pollution. The main pollution sources are (1) domestic
waste, (2) agro-based waste from rubber factory, (3) industrial waste from
factories and industrial estates, and (4) animal waste from stock farms
(Malaysian Coastal Resources Study Team 1992). Regulating domestic
waste disposal and waste treatment are big issues. Domestic waste is the
biggest contributor to the water quality problem, which has worsened
with the continuing population growth and from visiting tourists (Sg.
Pulai estuary has been earmarked for community based tourism). This
has caused not only an increase in human waste discharge, but also in the
household use of commercial compounds, such as detergents. Inadequacy
of sewage treatment systems further compounds the problem. Almost
all the sullage waste are discharged directly into the watercourses. In
Sg. Pulai most discharged waste eventually reaches the seagrass and
coastal areas, placing great stress to the ecosystem and its associated
resources. Activities such as opening land for agriculture and housing
and commercial centres, causes erosion and runoff, which contributes
to the suspended solids problem of rivers which will eventually find its
way to seagrass areas. Water visibility has never been greater than 1
m at shoals. This is further compounded by land extension of the port
6

development where dredged sediments contaminate and resuspend in the

water, putting at risk, the survival of fish and shellfish especially those
that are filter feeders.

Repeated visits showed dense overgrowth of the macroalgae
Gracilaria spp. and Amphiroa fragilissima causing the seagrasses in
the area to die, e.g., H. ovalis. Recovery occurred with regrowth of
seagrasses and the disappearance of the macroalgae (Japar Sidik and
Muta Harah 2003). Pulse disturbance can also be due to public usage of
natural seagrass area as avenues for transportation in the narrow channels
near the Sungai-Pulai-Merambong-Tanjung Adang shoals, Johor. The
locals from fishing villages are dependent on fisheries and the mobility
of the boat transport system (Malaysian Coastal Resources Study Team
1992). Related to this, biomass (AG, BG and T) distribution studies for
E. acoroides, T. hemprichii and H. ovalis on Merambong and Tanjung
Adang shoals showed high variability in values (Japar Sidik et al. 1996).
Based on field observations, the above-ground (AG) component, in
particular, the leaves of the larger species e.g., E. acoroides, T. hemprichii)
were always damaged and torn from the boats propeller (Japar Sidik
and Muta Harah 2003). Fishing activities by cast nets, molluscs and
bivlaves collection by the local residents may be also be part of the
contributing factor (Japar Sidik et al. 1996). Long term disturbances
include shipping activities that use the narrow passage near the seagrass
beds. All shipping has its risks of accidental pollution, but the greatest
concern is from hazardous cargoes such as petroleum and chemicals. The
periodic oil spills are suspected to be from ships cleaning their tanks of
sludge or changing their engine oil.

Ecosystem structures and functions could be altered and result
in ecosystem services being affected. The coastal development for
ports and facilities involving sand mining, filling and land reclamation,
represented a significant problem to the marine environment and the
associated resources of seagrass beds (Japar Sidik and Muta Harah
2003). Seagrass bed of Tanjung Adang Darat, Johor was destroyed by
filling and dredging for land extension and port construction (Japar Sidik
et al. 2007). Land reclamation caused an increase in the suspended solids
in the water column and localized losses or even death of seagrasses
by the smothering due to sand build up. Under the disturbed conditions
seagrasses are displaced by the smothering of built up sand. The
smothered area experienced a gradual decline of vegetation while burial
results in abrupt reduction (Figure 6). The bed became smothered in
thick glutinous silt often many centimeters deep and of little biological
value due to its anoxic nature. The benthic communities became choked
by the anoxic sediments. Most importantly, provisioning services
7

around Tanjung Adang Laut as well as those of Tanjung Adang Darat

and Merambong shoals support the fisheries, which directly affects the
earnings of the coastal communities who depend on this ecosystem. With
more expansion planned to completely reclaim the stretch of seagrass
beds of Merambong-Tanjung Adang shoals, the feeding ground of
dugongs at Merambong and Tanjung Adang Laut shoals too will face
similar fate.

Changes in ecosystem structures and functions cause alteration
in ecosystem services e.g., in fisheries. The effect of habitat change and
altered ecology has been realized by the fishermen. They claimed that
pollution is severely affecting fish yields in their areas. Additionally,
the fishing communities, particularly the traditional fishermen, were
adversely affected most by all developments along the coastal area. These
fishermen are bound to inshore areas (that are already destroyed as a
result of land reclamation) due to lack of bigger boats and engines. Some
fishermen have since left their profession and resorted to alternative
jobs, to cover the loss of income from their fishing activities.
Importance of the seagrass shoals
Healthy seagrasses not only function as service providers but they have
other equally important functions in the coastal ecosystem. As with other
green terrestrial plants, seagrasses being primary oxygen producers,
converting sunlight into oxygen. Oxygen is released into the water in
the form of dissolved oxygen and is available for diverse marine and
estuarine animals. Though small in number of species, seagrasses may
grow dense and form an extensive meadow. Their physical settings and
interactive community within and from outside account for the high
diversity, and help ensure the survival of an assortment of invertebrates
e.g., shrimps, starfishes, sea cucumbers, bivalves, gastropods (Sasekumar
et al. 1989; Arshad et al. 1994) and vertebrates e.g., fishes and other
plants e.g., seaweeds (Japar Sidik et al. 1996). Kikuchi (1980) discussed
some characteristics of seagrass beds that allow them to perform this
role. Among them are (i) where seagrasses form dense vegetation,
they increase the available substrate surface for epiphytic biota. The
differentiation of the plant into leaves, stems, rhizomes and roots increase
the heterogeneity resulting in a greater diversity of animals, some of
them feeding directly upon seagrasses, (ii) dense vegetation retards water
movement, thus offering calm underwater space within the bed for many
animals, such as mysids, hydromedusae and juvenile fishes, (iii) the
relatively less disturbed hydrodynamic conditions, settlement of minerals
and organic particles, deposition of decayed leaves and sedimentation
8

of fine, suspended particle occur, providing eutrophic environment for

benthic animals and (iv) reduction of excessive illumination in daytime
permits the development of a shaded microenvironment that is beneficial
to animals. They provide food and shelter for many fishes, crustaceans,
and they protect juvenile and small fishes from predators. Due to this,
species richness in seagrass ecosystem is equivalent if not richer than of
the mangroves or coral reefs.

Seagrass beds of Merambong and Tanjung Adang Laut shoals
are not only diverse aquatically but they function equally as service
providers to coastal ecosystems. Several critical ecosystem services were
identified with their ability to direct change and as primary drivers for
the seagrass shoals (Table 2). The coastal inhabitants are dependent on
fisheries. As such, they too belong to the seagrass ecosystem, in that at
least of the subsistence level, ie. the community uses seagrass associated
resources as a source of food. The economic contribution of seagrass
associated resources whether harvested or directly utilized, is difficult
to quantify as there are no records or figures quantifying such resources.
The waters around the Merambong, Tanjung Adang Laut shoals are part
of the traditional fisheries area (Figure 7) which supports the coastal
communities and are also important areas of biodiversity (Table 2).
Seventy-six species of fishes (e.g., the Indian anchovy, Stolephorus
indicus, barramundi Lates calcarifer), Spanish flag snapper (Lutjanus
carponotatus) and others including prawn (e.g., Penaeus indicus) and
crabs (Portunas pelagicus and Scylla serrata) have been reported in the
area (Sasekumar et al. 1989; Arshad et al. 1994, 2001). Traditional capture
fishing is done through the use of gill nets, drift nets, cast nets, pull nets
and hook and lines. Juvenile and young premature fish species are also
captured in the process. Catch is either sold fresh at local markets or are
salted/sundried for home consumption. The same seagrass areas during
low tide are used as collection and gleaning sites for gastropods (Lambis
lambis, Strombus canarium, Figure 7) and bivalves (Gafrarium sp.,
Meretrix meretrix, Modiolus sp.). Besides being rich in food resources,
seagrasses provide conditions for growth and abundance of invertebrates
and fishes that many local coastal inhabitants collect and catch for their
daily livelihood (Japar Sidik and Muta Harah 2003, Japar Sidik et al.
2006). Evidently seagrass ecosystems possess rich and diverse food
resources and yet they are continually threatened by human activities
causing their degradation and possibly habitat loss.

Cultural services of the coastal ecosystem in Sg. Pulai estuaryMerambong-Tanjung Adang Laut are not meant for public use such
as some forms of recreational pursuit including boating, fishing or
swimming, but rather for local fresh seafood ecotourism to cater for
9

tourists from Singapore and overseas. The ecosystem in this area has
become a major fishing and harvesting ground for traditional captured
fisheries. Fishermen are well aware that their catches are being served on
menus in several restaurants in the area.

Despite the area being degraded due to the land reclamation and
port facilities project, greater emphasis has been placed on the biological
resources of the area through awareness programmes on fisheries and
dugong conservation organized by The Fishery Department and the
Fishery Co-operatives for the local inhabitants. The involvement of
universities in research and conservation of the area have given the
area the status of having educational significance for dugongs (Dugong
dugong), green turtles (Chelonia mydas) and seahorses (Hippocampus
spp.) conservation (Choo and Liew 2005; Japar Sidik and Muta Harah
2011). The occurrence of dugongs in the Sg. Pulai estuary is evidently
supported by the presence of feeding trails commonly observed in the
seagrass shoals of Merambong, Tanjung Adang Laut (Japar Sidik and Muta
Harah 2003) and Seluyong (Figure 5). Both dugongs (status: vulnerable,
VU A1cd) and green turtles (status: endangered, EN A1bd), the largest
mammal and reptile associated and who feed on seagrasses are now low
in number and they are in need of protection because of environmental
degradation and exploitation. Seahorse species, Hippocampus kuda
(status: vulnerable VU A4cd) are very much in demand locally and
abroad for Chinese traditional medicine (Japar Sidik and Muta Harah
2003). Many bird species occasionally land on exposed seagrass areas
along the shores. Wading birds, such as some of the herons and egrets,
Egretta garzetta are seasonal visitors, seen feeding in seagrass shoals of
Merambong and Tanjung Adang.

Where seagrasses are found, human and associated organisms
benefit directly or indirectly from the presence of this vegetation and at
the same time, it has also become evident that seagrasses are a vulnerable
resource, easily lost in coastal areas facing environmental changes. The
serious threats faced are those derived from external sources - human
and coastal development activities. Continued human activities in coastal
areas continue to threaten and cause degradation, possibly leading to the
habitat loss of seagrasses. However, nature itself can also pose threats
to seagrasses such as storms that cause shifting sand that could bury or
smother seagrasses (Table 2). Given the importance of seagrasses as a
habitat for fisheries as well as a nursery and feeding ground, this resource
must be accorded the same priority and managed as well as mangroves
and corals, to provide for future renewable resource utilization, education
and training, science and research, conservation and protection. A priority
of initiatives including seagrass research should focus on how best to
10

manage open-access, multi-user seagrass systems mentioned above to

ensure their sustainable use in the aim to conserve and protect its rich
biodiversity.
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 (MOE) with the collaboration of
the Japan Society for the Promotion of Science (JSPS) Asian CORE
programme and the Establishment of Research Education Network on
Coastal Marine Science in Southeast Asia (COMSEA).

11

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