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CHAPTER TWO

2.0 LITERATURE REVIEW

2.1 The biology and characteristics of corals

Corals are among the simplest form of marine life found throughout the ocean

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

formation and cementation of skeletons of countless corals over thousands of years.

The succession of Sclerectinian corals in shallow tropical areas is attributed by the

symbiotic relationship with zooxanthellae.

The formation of the skeletons is determined by the convoluted and delicate

affiliation between symbiosis of coral polyp and zooxanthellae (Figure 2.1).

Zooxanthellae are minuscule and single-celled algae from the group of unicellular

brown plants known as dinoflagellates. This photosynthetic organism can capture

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

polyp. Secondly, zooxanthellae actively stimulate calcification during daylight when

algae are photosynthesizing (Veron, 2000). Most corals need explicit conditions with

sufficient amount of temperature, light availability, salinity and oxygen for survival

(Lieske et al., 2001).

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Tentacles with mouth

nematocysts
(stinging cells

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).

2.2 Calcification mechanisms in Corals

Corals skeleton is made of aragonitic calcium carbonate (CaCO 3) precipitation

(Barnes, 1970). The skeleton of corals comprised of two tissue layers (oral and aboral)

of multi-compartment structure divided by the coelenteron. The calcifying layer

(calicoblastic layer) faces the skeleton. The symbionts (zooxanthellae) exist within a

perisymbiotic membrane in the endodermal cells (Figure 2.2). However, the

mechanism of this enhancement is still uncertain (Carlon, 1996; Goreau et al., 1996;

Marshall, 1996).
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Oral
layers

Aboral
layers

Figure 2.2: Structure of coral (Veron, 1993)

Light is the major factor that enhances calcication (e.g. Goreau, 1959; Pearse

and Muscatine, 1971). Few mechanisms have been proposed to explain the interaction

of zooxanthellae and coral polyp on the calcification process. Firstly, light-enhanced

calcication via the supplied energy from the zooxanthellae (Goreau, 1959; Chalker

and Taylor, 1975), and secondly, protons generated from the calcification process are

used to assimilate bicarbonate and nutrients (McConnaughey and Whelan, 1997).

These processes are ecient in a coexisting system than in isolated reactions

(Suzuki et al., 1995); and calcication mechanism does not affect the photosynthesis

process (Yamashiro, 1995; Gattuso et al., 2000). However, some of these mechanisms

are conicting because the interactions of calcication with endosymbiont

photosynthesis are poorly understood and the biological processes of coral

calcification have to be comprehended (Gattuso et al., 2000).

2.3 Corals of the Family Poritidae

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.

solida (Veron, 1986, 2000; Fenner, 2009).

Coral Porites sp. inclines to dominate in back-reef or lagoonal habitats. The

colonies are either flat (laminar or encrusting), massive or branching in formation.

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

3 cm/year with up to several centuries of longevity and distinctive annual skeletal

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

(Scoffin et al., 1989).

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

morphologies and are only identifiable in-situ by minute corallites characteristics

(Veron, 1986, 2000).

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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

for measuring environmental changes over their growth history.

2.4.1 Growth banding in the skeleton of Porites sp.

Porites sp. colonies secrete calcium carbonate skeletons produce annual

layer of dense and less dense bandings. The bandings can be counted like tree

rings to determine the colony's age (Lough, 2011). Accretion of calcium

carbonate by Porites sp. is affected by the environmental factors such as light

intensity, salinity, and seawater temperature. The high-density (HD) and low-

density (LD) bandings in Porites sp. skeleton can be revealed by X-radiographic

and ultraviolet exposure techniques. Boto and Isdale (1985) suggested that

organic compounds produce by freshwater plants in soil, such as fulvic and

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

year growth, indicating a stress condition (Supriharyono, 2004). According to

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

on light intensity and seawater temperature. Measurements of the extension rate

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and calcification rate have been important in understanding the environmental

controls on coral growth (Tomascik and Sander, 1985; Knowlton et al., 1992;

Lough and Barnes, 2000; Carricart-Ganivet, 2004).

Calcication rate is the product of density and extension rate (Chalker et

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

temperature and to a lesser extent, reduction of light intensity (Grigg, 1981,

1997).
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2.5 Factors influencing the growth characteristic of Porites sp.

There are many factors affecting the growth of corals and can be divided into

biological and physical categories.

2.5.1 Biological factor affecting the growth of Porites sp.

Boring activities is one of the major biological factors affecting Porites

sp. growth. Macroboring organisms are universal within modern coral structure

(Bromley, 1970; Macintyre, 1984; Hutchings, 1986). Around 13 invertebrate

phyla including bacteria, fungi, foraminifera, sponges, sipunculans, polychaetes,

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

maximun; and the non-boring serpulid polychaete Spirobranchus (Clapp et al.,

1963; Carriker et al., 1969; Bromley, 1970, 1978; Warme, 1975; Warme et al.,

1978; Hutchings, 1986). They affect the coral through direct closure and

degradation of in-situ and rubble reef carbonate (Warme, 1975).

This activity in return is accountable for the abundance of fine-grained

sediment production (Futterer, 1974; Moore and Shedd, 1977; Acker and Risk,

1985). However, not much is recognized of the erraticism in the distribution of

this organism within and between reef habitats (Hutchings et al., 1992).

Bioerosion normally seen as a destructive course, by which corals are killed

(Acker and Risk, 1985) and coral heads fail and demoralize (Goreau and

Hartman, 1966).

The existence of boring sponges strongly declined the architecture of the

coral skeleton showing a series of chambers joined by channels. The effects of

boring activities in corals give spongy formation in the carbonate structure

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(Barbara et al., 2008). Highsmith, (1981) found that boring bivalve richness

fluctuated directly with total bio-erosional damage and that their numbers

increased as the amount of bioerosion increased (Figure 2.5).

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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Earlier corals considered dependent on a narrow range of environmental

conditions (Johannes, 1975; Endean, 1976; Loya, 1976; Loya and Rinkevich,

1980). Changes in temperature, salinity, water depth, light intensity, turbidity

and sedimentation rate can inuence not only growth but also the diversity and

abundance of corals (Hubbard and Scaturo, 1985; Lirman et al., 2003).

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).

i. Sea Surface Temperature

Temperature can limit coral growth and its development as a whole (Veron,

1993). Hoegh-Guldberg (1999) stated that sea surface temperature (SST) in

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

26.7 C for their skeletal extension and calcification.

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.

Increase in SST by 1 C will or promote calcification and reduce the extension

rate of coral and vice versa. The increased of SST is most likely the major factor

that leads to coral bleaching and high mortality of corals (Carricart-Ganivet,

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

destruction, such as Vietnam and the northern Philippines where up to 80%

corals bleached, and mortality was high. Singapore also reported the first major

bleaching event in the islands history.

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

a massive coral bleaching, mortality and reduced in calcification. On the other

hand, increased in the carbon dioxide partial pressure (pCO2) levels, results in

decreased in the aragonite saturation state and significant reductions in coral

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).

In Malaysia, reef condition varied widely throughout the East Malaysia

particularly in Sabah, with a disturbing amount of dead and shattered corals.

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

severity (Chou, 2000).

ii. Salinity

Salinity is an important factor for coral survival (Muthiga and Szmant,

1987; Coles, 1992). Corals, like most coelenterates, have few or no mechanisms

for osmoregulation (Muthiga and Szmant, 1987). Deviation from ambient

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

negatively impact Porites reproduction (Richmond, 1993). Mortality in Porites

sp. predicted to occur at salinities lower than 25 ppt (Edmondson, 1928) or

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

ppt) (Vaughn, 1919; Edmondson, 1928; Wells, 1957).

Decrease in salinity found to affect photosynthesis of zooxanthellae based

on salinity values and sunlight exposure length (Marcus and Thouraug, 1981).

The ability of Hermatypic corals to tolerate salinity stress is due to a behavioural

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

seawater (Muthiga and Szmant, 1987).

iii. Light and Depth

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

grow. However, different corals have different tolerances level to light

availability (Veron, 2000), and this result in the variation in coral community.

Many Hermatypic corals contain zooxanthellae that able to intensify their

calcification rates in the presence of light (Buddemeier and Kenzie, 1976;

Goreau, 1959, 1961, 1963; Chapman, 1974).

The phenomenon of "light-enhanced calcification" was first demonstrated

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

with depth is related to light availability where below 15 meters, growth

decrease 2 to 3 millimeter per year.

iv. Sedimentation

Sedimentation decreases the coral growth and reproduction (Aller and

Dodge, 1974; Vaisnys, 1977; Tomascik and Sander, 1987). Growth in corals is

affected by the diversion of energy to the removal of sediment particles.

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

surfaces plays an important role on the distributions and abundances of 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;

Rogers, 1983, 1990; Hodgson, 1990).

Corals exhibit active and passive removal action in removing sediment

particles. In active removal, sediment particles that settle on the surface are

removed by the inflation of coral polyps. Porites sp. also secretes mucus

surrounding the colonies to prevent from smoldering of its polyps due to

sediment settlement. As a result, sediments are congealed and eliminated from

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.

These sediments usually a have high organic components which stirred up by

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,

1980; Riegl and Branch, 1995).

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

away quickly (Veron, 1986). Deposition of calcium carbonate in corals depends

on photosynthetic rates of zooxanthellae (Goreau, 1959) and high sediment load

leads to signicant increase in respiration of corals as well as decrease in net

photosynthesis of zooxanthellae (Abdel-Salam and Porter, 1988; Riegl and

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Branch, 1995). Reduction in light intensity resulting from the increase of

suspended sediment can lead to decrease in the corals growth and

photosynthesis by zooxanthellae (Hubbard and Scaturo, 1985; Kendall et al.,

1985; Tomascik and Sander, 1985; Rogers, 1990).

v. Turbidity

Based on EPA Guidance Manual (1999), turbidity is a principal physical

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,

soluble coloured organic compounds, plankton and other microscopic organisms.

Turbidity is also a major factor governing corals growth variations in tropical

continental shelves (Buddemeier and Hopley, 1988). Most studies on the effects

of turbidity on coral reefs have concentrated on the anthropogenic increase in

sedimentation and turbidity on corals growth due to deforestation, dredging and

nutrient input (Rogers, 1990).

McCloskey et al. (1978) addresses the cumulative effects of turbidity on

Porites sp. growth, as expressed in the various degrees of their development

along a gradient of naturally high turbidity. Tolerance of Porites sp. to elevated

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

such conditions. The shift in species dominance reflects to a more turbidity

tolerant species and the elimination of major framework builders, such as

branching Acropora and massive Porites species (Van Woesik, 1992).

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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

increase with decrease in the turbidity and sedimentation (Carricat-Ganivet and

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

of short-term of high turbidity. Thus, association of sedimentation and turbidity

lead to mass mortality for corals (Cook et al., 2003).

vi. Water Motion

The most expressive factor determining the structure of coral communities

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

hurricanes and tropical storms.

Corals are found in a wide range of water motion conditions (Jokiel and

Coles, 1978). Water motion exerts an influence on the metabolism of corals by

affecting the food intake by the polyps. Sebens et al., (1998) stated that water

flow was a primary factor affecting feeding success of passive suspension

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;

Wolanski and Pickard, 1983).

Corals that live in exposed microhabitats benefit from increased flow of

water, compared to those in protected microhabitats on the same reef. Effects of

wave stress on corals generally considered on mortality due to breakage and

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