Deep-Sea Research II 96 (2013) 32–40

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Deep-Sea Research II
journal homepage: www.elsevier.com/locate/dsr2

Long-term changes in coral communities under stress from sediment

Thamasak Yeemin n, Sittiporn Pengsakun, Mathinee Yucharoen, Wanlaya Klinthong,
Kanwara Sangmanee, Makamas Sutthacheep
Marine Biodiversity Research Group, Department of Biology, Faculty of Science, Ramkhamhaeng University, Huamark, Bangkok 10240, Thailand

art ic l e i nf o a b s t r a c t

Available online 25 April 2013 Chronic disturbances of coral reefs are usually caused by a complex combination of several stresses that
Keywords: are increasingly driving coral reef degradation on both a regional and global scale. This study is aimed at
Coral community assessing the long-term changes of coral communities (2004–2010) by integrated investigation on
Recruitment sedimentation rates, coral community structures and coral recruitment patterns at the study sites in the
Sediment Western Gulf of Thailand. Sedimentation rates at the study sites during the study periods had means
Disturbance ranging from 29.49 to 59.53 mg cm−2 d−1. The coral community structures of the study sites did not
Gulf of Thailand change much during the study periods, with means of live coral cover 20.47–27.87%. The dominant corals
were Porites lutea, Favites abdita, Pavona decussata and Goniopora columna. However, the composition
and density of juvenile corals at both study sites were much changed during the study periods. The
dominant juvenile corals were P. lutea, Favia spp., Favites spp. and Pocillopora damicornis. The densities of
juvenile corals were relatively low and they suffered high mortality rates because of high sediment
deposition and macroalgal overgrowth. Coral recovery is a difficult task that will require an ecosystem
approach involving several management strategies and measures which include appropriate coastal
development, the enhancement of coastal wetlands and effective fishery management.
& 2013 Elsevier Ltd. All rights reserved.

1. Introduction disturbance, e.g. a coral disease epidemic or a coral bleaching

Disturbances of coral reefs are usually caused by a complex
combination of several stresses, such as sedimentation, coral
bleaching, destructive fishing practices, low water quality, preda-
tion and diseases. Many of these stressors are increasingly driving
coral reef degradation on a regional and global scale. Thus,
maintaining and/or increasing the resilience of coral reefs will be
dependent on a reduction in local impacts (Cooper et al., 2009).
There are several scientific reports on how different distur-
bances affect coral reefs (Fabricius, 2005; Pratchett et al., 2009)
however there are still critical gaps in the knowledge of how coral
reefs can recover from disturbances and retain a coral-dominated
state following various disturbances (Hughes et al., 2010). Some
researchers have shown that coral reefs that maintain their
structure and function can recover from large-scale disturbances
(Halford et al., 2004; Sheppard et al., 2008).
Disturbances to coral reef ecosystems can be chronic or acute
(Connell et al., 1997). Sedimentation and nutrient input from coastal
development are considered chronic disturbances to coral reefs. They
are slow to change and exert continuous stress on the system
(McCulloch et al., 2003; Fabricius, 2005; Cooper et al., 2009). An acute
n
Corresponding author.
E-mail address: thamasakyeemin@yahoo.com (T. Yeemin).
phenomenon, can change the community into a new dominant
component where reinforcing feedback mechanisms can prevent coral
recovery (Mumby et al., 2007; Hughes et al., 2010; Scheffer et al.,
2001). However coral recovery may occur between acute disturbances
and it is necessary to gain insights on how coral-dominated commu-
nities can persist in the face of global change (Graham et al., 2011).
Coral recovery trends are influenced by several factors, such as
disturbance patterns, reef characteristics, reef connectivity, ecolo-
gical characteristics and human impacts. High reef connectivity is
very important for replenishing marine organisms (Ayre and
Hughes, 2004; Jones et al., 2009). The grazing of sea urchins and
herbivorous fish and high functional diversity can promote eco-
system functions that will recover more rapidly from disturbances
(Mumby et al., 2007; Burkepile and Hay 2008). Coral reefs in the
vicinity of urban areas may exhibit slower recovery times because
of increased terrestrial inputs and the exploitation of natural
resources (Sandin et al., 2008; Graham et al., 2011).
Recent studies have provided a large database showing that
sedimentation, nutrient enrichment and turbidity can degrade
coral reefs on a local scale (Fabricius, 2005). Sedimentation is
recognized as a severe disturbance for coral reefs by reducing
growth and survival in a wide range of coral taxa. Elevated
sedimentation rates from coastal erosion and development have
severely damaged nearshore coral communities (Rogers, 1990;
McClanahan and Obura, 1997). Most sediment is transported into

0967-0645/$ - see front matter & 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.dsr2.2013.04.019
 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40 33

Fig. 1. Map of the study sites at Ao Thong Tanote and Ao Thong Krut, Samui Island, the Western Gulf of Thailand.

100 coastal ecosystems via rivers. The majority of the larger sediment grain
Sedimentation rate (mg.m-2.day-1)

90 sizes are deposited within a few kilometers of a river's mouth, whereas

80 fine grain sizes can be carried over longer distances. High sedimenta-
70 b ab
a tion rates may damage exposed coral tissue within a few days and low
60 c sedimentation rates can reduce photosynthesis rates in corals (Riegl
50
and Branch, 1995; Philipp and Fabricius, 2003). Coral degradation is
40
not only dependent on the quantity and duration of sedimentation,
30
20
but is also strongly dependent on sediment types and environmental
10 conditions. Low sedimentation rates, when combined with transpar-
0 ent exopolymer particles, cause mortality of coral recruits (Fabricius
2004 2006 2008 2010 et al., 2003). Exposure to sediments in a few days may cause long-
Ao Thong Tanote Ao Thong Krut term impacts in coral populations and inhibits coral recovery following
Fig. 2. Sedimentation rates (mean 7SE) at Ao Thong Tanote and Ao Thong Krut in
a disturbance. Therefore several nearshore coral communities may
2004, 2006, 2008 and 2010. Column with different letters are significantly different potentially shift forwards assemblages dominated by sediment-
(po 0.05) as determined by multiple comparison test Fisher's LSD. tolerant coral species (Sofonia and Anthony, 2008).
34 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40

The present study is aimed at assessing the long-term changes Table 2

of coral communities (2004–2010) by examining sedimentation Results of one-way ANOVA and Fisher's LSD test examining the influence of year on
live coral cover. Only significant results are shown for the Fisher's LSD test.
rates, coral community structures and coral recruitment patterns
at the study sites in the Gulf of Thailand. Source of variation df Mean square F p

Both stations
Between groups 3 36.769 17.340 o 0.001n
2. Material and methods Within groups 20 2.120
2004 vs. 2008 0.001n
2.1. Study area 2004 vs. 2010 o 0.001n
2006 vs. 2010 o 0.001n
2008 vs. 2010 0.009n
The Gulf of Thailand is a semi-enclosed tropical sea, located in the
South China Sea and surrounded by four countries, i.e. Cambodia, Ao Thong Tanote
Between groups 3 15.179 7.497 0.010n
Malaysia, Thailand and Vietnam. It is relatively shallow, with a mean
Within groups 8 2.025
depth of 45 m and a maximum depth of 80 m (Wattayakorn, 2006). 2004 vs. 2008 0.039a
Two coral communities on the south of Samui Island, Thailand were 2004 vs. 2010 0.002n
selected for the long-term study, namely Ao Thong Tanote (9124′57″N, 2006 vs. 2010 0.010
99156′18″E) and Ao Thong Krut (9125′7″N, 99156′49″E) (Fig.1). Samui Ao Thong Krut
Island has been recognized as an intensive tourism spot in Thailand. It Between groups 3 21.941 16.203 o 0.001n
is facing several problems from tourism development. The island has Within groups 8 1.354
2004 vs. 2008 0.004n
developed from a backpacker tourist destination into one of the most 2004 vs. 2010 o 0.001n
increasingly upscale tourist destination in Thailand over the past three 2006 vs. 2010 0.001n
decades (Pongponrat, 2009). Environmental parameters comprising 2008 vs. 2010 0.031n

df: degree of freedom

n
Significant difference (p o0.05).
Table 1
Results of one-way ANOVA and Fisher's LSD test examining the influence of year on
sedimentation rate. Only significant results are shown for the Fisher's LSD test.

Source of variation df Mean square F p

Both stations
Between groups 3 1120.945 19.336 o0.001n
Within groups 68 57.971
2004 vs. 2006 0.010n
2004 vs. 2008 o0.001n
2006 vs. 2008 o0.001n
2008 vs. 2010 o0.001n

Ao Thong Tanote
Between groups 3 944.493 24.339 o0.001n
Within groups 32 38.805
2004 vs. 2008 o0.001n
2006 vs. 2008 o0.001n
2008 vs. 2010 o0.001n

Ao Thong Krut
Between groups 3 310.810 8.888 o0.001n
Within groups 32 34.972
2004 vs. 2006 0.005n
2006 vs. 2008 o0.001n Fig. 4. Percentages of coral coverage (mean7 SE) for each taxa at Ao Thong Tanote
2008 vs. 2010 0.003n in 2004, 2006, 2008 and 2010.

df: degree of freedom

n
Significant difference (p o 0.05).

100
90
Live coral coverage (%)

80
70
60
50
40 ab b c
30 a
20
10
0
2004 2006 2008 2010
Ao Thong Tanote Ao Thong Krut

Fig. 3. Percentages of live coral coverage (mean7 SE) at Ao Thong Tanote and Ao
Thong Krut, Samui Island in 2004, 2006, 2008 and 2010. Column with different
letters are significantly different (p o0.05) as determined by multiple comparison Fig. 5. Percentages of coral coverage (mean 7 SE) for each taxa at Ao Thong Krut in
test Fisher's LSD. 2004, 2006, 2008 and 2010.
 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40 35

temperature, salinity, pH, conductivity and total suspended solids in
both study sites were quite similar. The average current velocity range
was 0.24–0.33 m/s (Pengsakun and Yeemin, 2009). The coral commu-
nity at Ao Thong Tanote occurred at 1–5 m water depths and at 1–3 m
water depths at Ao Thong Krut. The shallower community at the latter
is attributed to higher turbidity and more impact from coastal
development and boat transportation in the proximity. The coral
communities experienced a severe coral bleaching event in 1998
(Yeemin et al., 1998, 2009). All data in this study was collected before
the 2010 coral bleaching event.

Fig. 6. Underwater photos at the study sites: (A) a colony of Porites covered by sediment, (B) an unhealthy colony of Pavona decussata, (C) a dominant macroalga Turbinaria
and (D) Acropora colonies at Ao Thong Krut.

Fig. 7. Cluster analysis of coral species change during 2004–2010 at Ao Thong Tanote.
36 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40

Fig. 8. Cluster analysis of coral species change during 2004–2010 at Ao Thong Krut.
Density of juvenile coral (colonies.m-2)

1.0
Table 3
0.9
Results of one-way ANOVA and Fisher's LSD test examining the influence of year on
0.8 juvenile coral density. The data were log transformed. Only significant results are
0.7 shown for the Fisher's LSD test.
0.6
0.5 a d Source of variation df Mean square F p
0.4
0.3 bc Both stations
bc Between groups 3 0.147 8.723 0.001n
0.2
0.1 Within groups 20 0.017
0.0 2004 vs. 2006 0.001n
2004 2006 2008 2010 2004 vs. 2008 o 0.000n
2006 vs. 2010 0.016n
Ao Thong Tanote Ao Thong Krut
2008 vs. 2010 0.004n
Fig. 9. Density of juvenile coral colonies (mean 7SE) for each study site in 2004,
Ao Thong Tanote
2006, 2008 and 2010. Column with different letters are significantly different
Between groups 3 0.054 4.523 0.039n
(po 0.05) as determined by multiple comparison test Fisher's LSD.
Within groups 8 0.012
2004 vs. 2006 0.024n
2004 vs. 2008 0.010n
2.2. Sedimentation rates 2004 vs. 2010 0.026n

Ao Thong Krut
Sedimentation rates were measured using a nested experimental Between groups 3 0.142 12.657 0.002n
design. Sediment traps, consisting of a PVC tube of 5 cm internal Within groups 8 0.011
diameter, 11.5 cm length and sealed at one end (English et al., 1997), 2004 vs. 2006 0.007n
were deployed in groups of three at three different locations (about 2004 vs. 2008 0.003n
2006 vs. 2010 0.002n
20 m away from each other) at each study site. The sediment traps
2008 vs. 2010 0.001n
were 20 cm above the substrate and left in situ for 4–6 weeks,
depending on weather conditions and logistical considerations, and df: degree of freedom.
were replaced after they were retrieved. Sediment trap deployment n
Significant difference (p o0.05).
was conducted in 2004, 2006, 2008 and 2010. The sediment and
water within the traps were filtered through a pre-weighed What-
man quantitative filter, dried at 60 1C until reaching a constant
weight and subsequently weighed. Sedimentation rates are coral colonies (≥5 cm in diameter) were counted and identified to
expressed as mg dry weight cm−2 day−1. a species level, if possible, and their coverage was quantitatively
estimated. Quadrats (1 m  1 m), subdivided into 100 squares
2.3. Coral community structure of 100 cm2, were positioned along the transect. The quadrats
were also photographed with an underwater camera for re-
At each study site, three permanent belt transects of 50  1 m2 investigating the data. The reef surveys were undertaken in March
were placed perpendicularly to the shoreline in which all visible 2004, 2006, 2008 and 2010.
 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40 37

2.4. Coral recruitment patterns 3.3. Changes in density of juvenile corals

At each study site, the number of visible juvenile corals (about The means of the densities of juvenile corals during the study
0.5–5 cm diameter) were directly observed by SCUBA divers in periods at Ao Thong Tanote were 0.17–0.35 colony m−2 while those
three permanent belt transects (30  1 m2 for each). All juvenile at Ao Thong Krut were 0.14–0.32 colony m−2 (Fig. 9). The densities
corals were identified to a genus level or species level, if possible. of juvenile corals between the years 2004–2006, 2004–2008,
The observations were made in March 2004, 2006, 2008 and 2010. 2006–2010, and 2008–2010 were significantly different (Table 3).
The dominant juvenile corals at both study sites were P. lutea,
Favia spp., Favites spp. and Pocillopora damicornis (Figs. 10–12).
2.5. Data analysis
Juveniles of Acropora spp. were also abundant at Ao Thong Krut in
the year 2008. The compositions of juvenile corals at both study
The quantitative data gathered from the field survey, such as
sites were much changed during the study periods.
sedimentation rates, live coral coverage, and the density of
juvenile coral was tested for normality and homogeneity of
variances using the Shapiro–Wilk test. According to the test, only
the density of juvenile coral data was not normally distributed so
4. Discussion
they were then log transformed to meet the assumptions of the
parametric statistical test. A one-way ANOVA was used to test the
The means of the sedimentation rates of 29.49–
influence of time on sedimentation rates, live coral coverage and
59.53 mg cm−2 d−1 found in the present study are relatively high
densities of juvenile corals. Multiple comparisons were analyzed
(Rogers, 1990; Lane, 1991). The coral communities in this study
using Fisher's LSD test in case there were two group means (or
had accumulated tremendous loads of sediment to the extent that
more) that were significantly different from each other. Cluster
dead corals were found to have mud draped on them. Therefore
analysis was also performed on the coral community data to find
sediment deposition is considered as a chronic disturbance to
groupings of coral species, changed during 2004–2010. The data
coral reefs at Samui Island (Connell et al., 1997). It has caused a
were normalized using the log10 (X+0.5) transform function.
decrease of filter feeders and grazers, especially Diadema setosum
(Dikou and van Woesik, 2006; Yeemin et al., 2009).

3. Results

3.1. Variation of sedimentation rates

The means of the sedimentation rates during the study periods at

Ao Thong Tanote ranged from 29.49 to 52.31 mg cm−2 d−1 while those
at Ao Thong Krut varied from 45.51 to 59.53 mg cm−2 d−1 (Fig. 2).
There were significant statistical differences in sedimentation rates
during study periods (Table 1). According to the multiple comparisons,
the sedimentation rates between the years 2004–2006, 2004–2008,
2006–2008, and 2008–2010 were significantly different. The variation
in sedimentation rates may have been caused by coastal development
in the areas. Ao Thong Krut had a higher sedimentation rate due to
more impact from coastal activities and boat transportation.

3.2. Changes in coral community structures

The means of the live coral coverage during the study periods at Ao
Fig. 10. Density of juvenile coral colonies for each taxa (mean 7 SE) at Ao Thong
Thong Tanote were 22.64–27.87% while those at Ao Thong Krut were
Tanote in 2004, 2006, 2008 and 2010.
20.47–26.76% (Fig. 3). There were significant statistical differences of
percentages of live coral coverage between study periods (Table 2).
The percentage of live coral coverage between the years 2004–2008,
2004–2010, 2006–2010, and 2008–2010 were significantly different.
The dominant corals at both study sites during the study periods were
Porites lutea, Favites abdita, Pavona decussata and Goniopora columna
(Figs. 4–6). Fungia fungites and Pachyseris rugosa were also abundant at
Ao Thong Tanote. The coral community structures of both study sites
had not changed much during the study periods. There was an overall
increase in coral cover from 2004 to 2010 because the dominant corals
were tolerant to sediment. The dominant corals such as P. lutea and
F. abdita had been degraded from the coral bleaching event in 1998
(Yeemin et al., 1998). The dendrogram from the cluster analysis
revealed two major groups of coral species change during 2004–
2010: high and low levels in change of coral cover (Figs. 7 and 8).
P. lutea and F. abdita were grouped at a high level of change in coral
cover, while a low level of change in coral cover can be further
grouped into four sub-divisions. Furthermore, the coral cover of P. lutea
and F. abdita at Ao Thong Krut were shown to be at a higher level of Fig. 11. Density of juvenile coral colonies for each taxa (mean 7SE) at Ao Thong
change than Ao Tongtanote. Krut in 2004, 2006, 2008 and 2010.
38 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40

Coral communities in the Gulf of Thailand experienced the
severe coral bleaching event in 1998. Only a few colonies of
Acropora, which is recognized as highly susceptible to coral
bleaching, were recorded at Ao Thong Krut. Less susceptible corals
however, such as Porites, Goniopora, Favites and Pavona, continue
to be dominant components at our study sites. Porites is a poor
sediment rejector (Stafford-Smith and Ormond, 1992) but it is a
dominant species at our study sites which are in high turbid
environments. This may be partly explained by its ability to
tolerate sediment deposition through mucous secretion, trapping
and subsequent sloughing (Dikou and van Woesik, 2006). More-
over, Porites is recognized as taxa that can rapidly regenerate
tissue damage (Brown et al., 1990; van Woesik, 1998) and often
form large monospecific colonies (Golbuu et al., 2011). Pachyseris,
which is normally found in deeper water (Sheppard, 1980), was
observed at 4–5 m depth at Ao Thong Tanote.
Our results are similar to previous studies that showed the
impacts of differential sedimentation rates, particularly terrige-
nous sedimentation, on juvenile coral density and composition
and larval settlement (Hodgson, 1990; Gilmour, 1999; Glassom
et al., 2004; Birrell et al., 2005; Dikou and van Woesik, 2006;
Golbuu et al., 2011). Densities of juvenile corals at Samui Island
were low in comparison to other coral reefs (Roth and Knowlton,
2009). Our data also implies that juvenile corals at Samui Island
suffered high mortality rates. The causes of juvenile coral mortality
may be sediment deposition or the overgrowth by macroalgae and
freshwater plumes. The coral community at Ko Samui was mostly
covered by macroalgae such as Sargassum oligocystum, Turbinaria
decurrens and Dictyota cervicornis (Yeemin et al., 2009). Inhibition
of juvenile corals by benthic algae is a critical factor to the
recovery and resilience of corals. Large-scale coral mortality
events, affected by severe disturbances, were often followed by

Fig. 12. Underwater photos of juvenile corals at the study sites: (A) Pocillopora damicornis, (B) Fungia fungites, (C) Favia sp., (D) Favites sp., (E) Goniopora columna and
(F) Acropora spp.
 T. Yeemin et al. / Deep-Sea Research II 96 (2013) 32–40
an abundance of benthic algae. The ability of corals to recover in
algal-dominated communities depends on the effects of algae on
growth and the survival of juvenile corals (Venera-Ponton et al.,
2011). However, juvenile colonies of Porites, Pocillopora and Favites
seemed to have better adaptation to such high sedimentation rates
and turbidity environments. Low juvenile coral densities at our
study sites may be due to low availability of larvae and suitable
substrates for coral settlement (Yeemin et al., 2009). Juvenile coral
observations not only provide important ecological data for coral
population dynamics, but also could be applied as an indicator of
environmental changes (Roth and Knowlton, 2009). Our long-term
observations in the Gulf of Thailand show that live coral cover at
the study sites declined significantly and did not recover. The poor
recovery is attributed to the absence of healthy coral communities
in the vicinity of Samui Island which prohibits coral larvae from
maintaining a coral population by self-recruitment. Thus it is
necessary to protect relatively healthy coral communities that
can provide coral larvae to Samui Island. It is also likely that high
turbidity may destroy larval supply and high sedimentation rates
and abundant macroalgae may diminish suitable substrates for
coral recruitment.
Several conservation groups and researchers have attempted to
promote a return to healthy coral communities in the Gulf of Thailand
in the form of coral restoration projects (Yeemin et al., 2006). The
present study suggests that coral recovery is a difficult task that will
require an ecosystem approach involving several management strate-
gies and measures. It is urgent to have proper land use management
and suitable coastal development in the surrounding catchment to
prevent pollution and excess nutrients and sediment from land runoff.
The protection and enhancement of coastal wetlands to trap sedi-
ments and slow freshwater discharges is also required (Wolinski et al.,
2009). It should be noted that effective fisheries management, to
replenish herbivorous fish populations, can promote coral recovery
also (McClanahan et al., 2009). The present study provides valuable
insights into coral communities under stress from sediment in the Gulf
of Thailand which has further implications for our understanding of
coral recovery and its resilience in a changing environment.
Acknowledgments
This study was funded by a budget for research promotion from
the Thai Government awarded to Ramkhamhaeng University in
the fiscal years B.E. 2552–2555 and partly supported by the
Provincial Electricity Authority (PEA) of Thailand. We acknowledge
Professor Dr. Jing Zhang, the leader of the project “Coral Reefs
under Climate and Anthropogenic Perturbations” (CorReCAP)
under IOC/WESTPAC for his support. The author would like to
thank the staff of the Marine Biodiversity Research Group, Depart-
ment of Biology, Faculty of Science, Ramkhamhaeng University for
their assistance in the field.
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