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CHAPTER TWO
around the world (Robin et al., 1991; Spalding et al., 2001). Corals, which referred to
as the Scleractinia, can be divided into reef building and non-reef building groups.
Hermatypic coral are reef-building corals that need sunlight to live. Ahermatypic
corals are non-reef building corals known as soft corals that can live without sunlight
in deeper ocean (Veron, 1986). Hermatypic corals inhabit the most diverse reef
ecosystem, and yet are said to be sensitive to environmental changes caused by climate
change (Chadwick-Furman et al., 2000). The structures of corals results from the
Zooxanthellae are minuscule and single-celled algae from the group of unicellular
suns energy to make its own organic food (Veron, 1986). The symbiotic relationship
between zooxanthellae and Hermatypic coral polyps benefits in two ways. Firstly,
organic carbon that produced by zooxanthellae is used as a food source by the coral
algae are photosynthesizing (Veron, 2000). Most corals need explicit conditions with
sufficient amount of temperature, light availability, salinity and oxygen for survival
zooxanthellae
living tissue
linking polyps
nematocyst
Figure 2.1: Anatomy of a coral polyp showing the symbiotic relationship between
coral polyp and zooxanthellae (modified from Encyclopaedia Britannica,
1999).
(Barnes, 1970). The skeleton of corals comprised of two tissue layers (oral and aboral)
(calicoblastic layer) faces the skeleton. The symbionts (zooxanthellae) exist within a
mechanism of this enhancement is still uncertain (Carlon, 1996; Goreau et al., 1996;
Marshall, 1996).
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Oral
layers
Aboral
layers
and Muscatine, 1971). Few mechanisms have been proposed to explain the interaction
calcication via the supplied energy from the zooxanthellae (Goreau, 1959; Chalker
and Taylor, 1975), and secondly, protons generated from the calcification process are
(Suzuki et al., 1995); and calcication mechanism does not affect the photosynthesis
process (Yamashiro, 1995; Gattuso et al., 2000). However, some of these mechanisms
Kingdom: Animalia
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Phylum: Cnidaria
Class: Anthozoa (Flower-like animals)
Subclass: Hexacorallia
Order: Scleractinia (Stony corals)
Family: Acroporidae
Family: Faviidae
Family: Poritidae
Genus: Alveopora
Genus: Goniopora
Genus: Porites
Species: P. cylindrica
Species: P. lobata
Species: P. lutea
Species: P. nigrescens
Species: P. solida
Genus: Poritipora
Genus: Stylaraea
Figure 2.3: Summary of the classification for Family Poritidae and Porites sp. from
Veron (2000).
There are fifteen common Sclerectinian families found throughout the world.
Among the fifteen families, Family Acroporidae, Faviidae and Poritidae are the most
diverse in the world (Van Woesik, 1995). Family Poritidae consists of five genuses
(Figure 2.3) with distinct characteristics from each family. Among the genera, genus
Porites is widely distributed and found throughout the world, mainly in the tropical
region (Veron, 2000). Form 324 species of coral identified in Malaysia, 22 Porites
species are found, mainly Porites cylindrica, P. lobata, P. lutea, P. nigrescens and P.
Porites species form massive structures called microatolls, with hemispherical shape
when small and grow into a helmet or dome-like shaped when at large. The colonies
may grow over 5 meters across (Veron, 2000) with linear extension rates of about 1 to
banding. This massive colony lives in a wide range of environmental settings (e.g.
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light intensity, sedimentation, pH, salinity and water motion) in shallow tropical seas
Porites species are the most difficult of all major genuses to identified, because
of their small and variable corallites and due to their massive growth-forms that are
hard to be recognized underwater (Veron, 2000). Porites have very small corallites
which are closely packed with little coenenchyme (Figure 2.4). The walls and septa are
generally very porous, yet relatively fast growing. This species is in need of taxonomic
review because the majority of massive species does not have distinctive
Figure 2.4: Scanning Electron Microscope (SEM) image of Porites sp. corallites (top), plan
view of the corallites at the bottom shows the key diagram of the characters
(left) with an illustration of the corallites (right), all in the same orientation.
Image modified from Veron (2000).
2.4 Growth of Porites sp.
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The skeletons of massive corals have been used by several scientists (Hudson et al.,
1982; Isdale, 1984; Boto and Isdale, 1985; Supriharyono, 1986, 1987, 1998) as tools
layer of dense and less dense bandings. The bandings can be counted like tree
intensity, salinity, and seawater temperature. The high-density (HD) and low-
and ultraviolet exposure techniques. Boto and Isdale (1985) suggested that
humic acids, might be incorporated into the coral skeletons. These compounds
fluoresce under ultraviolet light producing an alternate bright and dull banding
pattern in corals (Isdale, 1984). The environmental factors that have major effect
on corals are the amount of rainfall, sunlight duration, salinity and seawater
temperature.
Generally, one year of Porites sp. growth consists of two bands, one HD
band and one LD band. However, some corals may exhibit two HD bands in a
Supriharyono (1986), the HD bands are deposited during the wet season when
the amount of rainfall and sedimentation are very high, and the number of
sunlight hours is low. LD bands are deposited during the dry season when the
amount of rainfall and sedimentation are very low, and the number of sunlight
hours is very high. Highsmith (1979) mentioned that density in corals dependent
controls on coral growth (Tomascik and Sander, 1985; Knowlton et al., 1992;
al., 1985). Calcification rate provides a highly important proxy for seawater
temperature in tropical seas and can be measured from the coral growth banding
patterns (Lough and Barnes, 1997, 2000; Carricart-Ganivet, 2004). The rate is
measured on the bands of the corals skeleton along its growth axes. The decline
in calcication rate with increased latitude was due to the effects of reduced
1997).
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sp. growth. Macroboring organisms are universal within modern coral structure
arthropods, molluscs and some plant groups that have symbolic bore coral
substrates. Among the 13 organisms, three species are associated with live coral
colonies. They are the bivalve, Lithophaga laevigata; the vermetid Dendropoma
1963; Carriker et al., 1969; Bromley, 1970, 1978; Warme, 1975; Warme et al.,
1978; Hutchings, 1986). They affect the coral through direct closure and
sediment production (Futterer, 1974; Moore and Shedd, 1977; Acker and Risk,
this organism within and between reef habitats (Hutchings et al., 1992).
(Acker and Risk, 1985) and coral heads fail and demoralize (Goreau and
Hartman, 1966).
(Barbara et al., 2008). Highsmith, (1981) found that boring bivalve richness
fluctuated directly with total bio-erosional damage and that their numbers
Peyrot-Clausade et al. (1992) stated that the rate of boring rise with the
time since the death of the colony and the main agents in bioerosion also
changed with the increase of corals age. A clarification for low strength lies
partly in the greater porosity of the skeletons of Porites sp. because members of
the family Poritidae had among the lightest skeletons (Constanz, 1986; Hughes,
1987).
Figure 2.5: Destruction in coral by boring organisms. Red circles mark the sign of
macroboring and yellow circles indicate microboring effects on coral
Porites sp.
2.5.2 Impact of physical factors to the growth of Porites sp.
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conditions (Johannes, 1975; Endean, 1976; Loya, 1976; Loya and Rinkevich,
and sedimentation rate can inuence not only growth but also the diversity and
However, recent results show that corals are more adapted to physical stresses
than was previously believed (Muthiga and Szmant, 1987; Harland and Brown,
1989; Grigg and Dollar, 1990; Coles, 1992; Coles and Jokiel, 1992; Brown,
1997).
Temperature can limit coral growth and its development as a whole (Veron,
many tropical regions increased almost 1 C over the past 100 years which lead
to an event like coral bleaching. Nasir (2005) added that temperature affected
corals both direct and indirectly. Porites sp. favours an optimum temperature of
Temperature affects the metabolism rate of corals and limits the distribution
of the species (Connell, 1978). Veron (1986) mentioned that corals could not
survive when seawater temperature falls below 18 C, which is lower than its
ambient temperature and kill the colonies due to the incapability of reproduction.
rate of coral and vice versa. The increased of SST is most likely the major factor
2004; Goreau and Hayes, 1994; Davies et al., 1996; Winter et al, 1998).
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The Earth suffered the most dramatic changes in climatic record early in
1997 when a major El Nio event started and continued by a rapid substitute to a
La Nia in mid-1998. The bleaching in 1997 to 1998 was the most intense
climatic record with damage seen in all oceans. Extreme coral bleaching and
mortality experienced by many corals around the world during the event.
Bleaching effect varied across the region, with most countries reporting severe
corals bleached, and mortality was high. Singapore also reported the first major
The major causal factor is the increased of SST during the extreme El Nio
and La Nia events (Wilkinson and Hodgson, 1999). Increased in SST results in
hand, increased in the carbon dioxide partial pressure (pCO2) levels, results in
calcification. Later, year 2010 saw another vast bleaching event that affected
coral reefs in the whole of South East Asia (Reef Check, 2010). NOAA (2010)
informed that coral bleaching have been reported from mid-March to mid-July
2010. This event caused by the extreme high temperature throughout Southeast
Asia and the central Indian Ocean. This is the worst mass bleaching event since
1997 to 1998 where 16 % of the worlds reefs were killed (Wilkinson, 2000).
Dead coral cover of 10 to 20 % was found on 70 % of the reefs. The report also
stated that the nearest region also affected from the event, such as in Sumatra
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and Sulawesi with more than 75 %, while some ranged from mild to medium
ii. Salinity
1987; Coles, 1992). Corals, like most coelenterates, have few or no mechanisms
salinity of 33 ppt can affect their overall metabolism. In the long term, this
indicates a decrease in potential for growth and reproduction (Coles and Jokiel,
1992). For example, studies on coral communities in Hawaii show that the
Porites sp. exposed to reduced salinity after freshwater ooding can recover
quickly from natural disturbances, but not under polluted conditions (Jokiel et
al., 1993).
Coral Porites sp. are stenohaline, with limited ability to adapt to or survive
salinity changes (Wells, 1957; Coles and Jokiel, 1992). Coral can live in an
optimum salinity of 36 ppt. Sudden changes in sea water salinity are known to
higher than 45 ppt (Jokiel et al., 1974). Nonetheless, other studies suggested that
corals could tolerate both low (17.5 to 28 ppt) and high salinities (38.5 to 52.5
on salinity values and sunlight exposure length (Marcus and Thouraug, 1981).
response. For example, corals retract its polyps under osmotic stress and
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decrease the amount of tissue surface area in contact with the surrounding
Light changes rapidly with depth (Veron, 1986). The visibility of corals may
exceed up to 50 metres and this range controls the depth at which the corals
availability (Veron, 2000), and this result in the variation in coral community.
by Kawaguti and Sakumoto (1948). Bosscher and Meesters (1992) stated that
growth rate of Porites sp. is related to light availability. Decrease in growth rate
iv. Sedimentation
Dodge, 1974; Vaisnys, 1977; Tomascik and Sander, 1987). Growth in corals is
Particles suspended in the seawater column can alter the intensity and spectral
quality of light reaching the corals, thereby affecting their metabolism (Coraline,
1990). Inorganic sediments that suspend in the seawater settle on the coral
species. Settlement of inorganic sediment onto live coral surfaces cause tissue
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necrosis and lead to the death of coral polyp because of the combined effects of
smothering and microbial action (Bak and Elgershuizen, 1976; Bak, 1978;
particles. In active removal, sediment particles that settle on the surface are
removed by the inflation of coral polyps. Porites sp. also secretes mucus
corals surface. In passive removal, water movement will help to keep the
sediment particles from settling on colony surfaces. At the same time, coral
polys will retract their tentacles forming flat surfaces for sediments to eradicate
via water movement. This action allows the corals to spend less energy in
sediment rejection. There are many types of sedimentation occur on and around
the reefs. Some of the sediments came from the land through the river run-off.
waves and may remain suspended in the water. This will turbid the water and
reduce the light penetration, and as it settles after long disturbance (Lasker,
Due to the effect of sediments, organisms such as corals can die either by
completely buried or chocked, since the coral polyps fail to clear the sediment
v. Turbidity
characteristic of water caused by suspended matters that interfere with the clarity
of the water. These matters include clay, silt, fine inorganic and organic matter,
continental shelves (Buddemeier and Hopley, 1988). Most studies on the effects
suspended sediment varies from reef to reef, and depends on the turbidity
conditions. He also stated that reduction in species diversity under high turbidity
is expected since only few species such as Porites lutea would be able to tolerate
Hallock et al., (1986) outlined the effects of reduced water clarity on normal
zonation patterns of coral reefs where the density and calcification of corals
Merino, 2001). Rogers, (1999) in his report stated that some of the corals had
been dead to over a decade, mostly in shallow water close to the shore as a result
is the physical interruption from waves. Wave energy has long been renowned as
one of the most important factor that regulates coral growth, reef development
and coral island formation (Yonge, 1940; Storr, 1964; Roberts, 1974; Adey,
1978). Cyclones are most common in the South China Sea from November to
December and frequently originate near Peninsular Malaysia (Zuki and Lupo,
2008). Corals along the tropical rain forests are exposed to areas of regular
Corals are found in a wide range of water motion conditions (Jokiel and
affecting the food intake by the polyps. Sebens et al., (1998) stated that water
feeders like corals. Augustus et al., (1995) added that morphological adaptation
of certain corals is due to the influence of water motion and elicits a variety of
responses in corals. They also mentioned that when the currents are strong, it
provides water movement to pass through corals at all depths below the reef
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crest to remove sediment away from coral surface (Roberts et al., 1975;
abrasion. Storlazzi et al., (2003) also added that there are strong qualitative
correlations between the wave energy, coral speciation and reef morphology
(Dollar, 1982; Done and Massel, 1993; Blanchon and Jones, 1997).
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2.6 The past and current studies of coral growth in Southern South China Sea