Research Article Status of Coral Reef Communities on Two ...downloads.hindawi.com/archive/2013/358183.pdfIn Semporna, the island of Bodgaya was rst gazetted in as a forest reserve,

Hindawi Publishing CorporationJournal of EcosystemsVolume 2013 Article ID 358183 15 pageshttpdxdoiorg1011552013358183

Research ArticleStatus of Coral Reef Communities on Two Carbonate Platforms(Tun Sakaran Marine Park East Sabah Malaysia)

A Montagne1 O Naim12 C Tourrand23 B Pierson4 and D Menier15

1 Geosciences Marines et Geomorphologie du Littoral Laboratoire Domaines Oceaniques UMR 6538 CNRS Universite de BretagneSud (UBS) 56000 Vannes France

2 Laboratoire ECOMAR Universite de la Reunion (UR) 97460 Sainte-Clotilde France3 Departement de Physique Universite de la Reunion (UR) 97460 Sainte-Clotilde France4 South-East Asia Carbonate Research Laboratory (SEACARL) Universiti Teknologi Petronas 31150 Tronoh Perak Malaysia5 Department of Geosciences Universiti Teknologi Petronas 31150 Tronoh Perak Malaysia

Correspondence should be addressed to O Naim odilenaimorangefr

Received 12 March 2013 Revised 29 May 2013 Accepted 26 August 2013

Academic Editor Luiz E Aragao

Copyright copy 2013 A Montagne et al This is an open access article distributed under the Creative Commons Attribution Licensewhich permits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

This study concerns three sites located on carbonate platforms east Sabah Gaya West Gaya East and Mantabuan At each sitethe dominant coral shapes and their health were recorded (lagoons and outer slopes) Densities of echinoderms Tridacna andnudibranchs were recorded while fish density was estimated Generally the coral vitality is low (le50 living corals) Massive coralsdominate all sites except the Gaya West-outer slope where coral coverage and diversity are the highest On the Mantabuan-meshreef a diverseAcropora assemblage dominates the landscape On the reef flat of Gaya East monospecific circa 10meter coral patchesoccur Primary producers are scarce on all sites Sea urchins dominated by Diadema are abundant on the Gaya East-reef flat andthe Gaya West-mesh reef Sea stars and holothurids are the most prevalent in Gaya West-outer slope although they remain scarceCrinoids are only abundant inMantabuan Stegastes damselfish highly characterizes the sites of Gaya East (reef flat and inner slope)and the Mantabuan-mesh reef On the Mantabuan-outer slope parrotfish and other fishes are plentiful No sign of eutrophicationhas been detected and natural hypersedimentation andor eventual ancient bleaching events appear to be the direct principal causesof coral death or coral degradation

1 Introduction

An exceptionally high biodiversity prompted the creationof the Tun Sakaran Marine Park (TSMP) in the westernpart of the Celebes Sea east of Semporna Sabah Malaysiaas the site supports a greater diversity of marine speciesthan recorded elsewhere in Malaysia The site is therefore ofregional national and local importance

Marine and coastal habitats include (1) mangroves (2)seagrass beds (3) lagoons (4) fringing patch and bank reefsand (5) open water Studies show that the diversity of marinespecies in the proposed park is very high There are over600 species of fish and at least 250 species of corals [1 2]Harborne et al [3] summarizing fragmented studies of coralspeciesrsquo richness around Malaysia reported a total of 323species Around the reefs 265 species of molluscs occur [4]

and theremay be asmany as 140 species of sponges 70 speciesof soft corals and at least 50 species of gorgonians [5 6] Inaddition a high diversity of echinoderms exists with a totalof 109 species recorded [5 7 8] Several of the sponges andsoft corals found in the Gaya lagoon are new undescribedspecies One sponge is very unusual and possibly endemic tothe Semporna reefs [9] Also Allen [1] found two new speciesof damselfish

The preliminary results of the Semporna Marine Ecolog-ical Expedition [10] (httpawsassetspandaorgdownloadssmee factsheet dec 2010pdf) indicate that Semporna mayhave one of theworldrsquos highestmarine biodiversity For exam-ple the expeditionrsquos fish counts demonstrated that Sempornais one of the richest areas within the Coral Triangle (844species of fish encountered) Also the expedition yielded

2 Journal of Ecosystems

a record number of 43 species of mushroom corals (theprevious highest recorded richness in this family was 40species in Indonesia and Papua New Guinea) Furthermoresome new species were discovered (two species of shrimp andpossibly a number of gall crabs)

In Semporna the island of Bodgaya was first gazetted in1933 as a forest reserve and the island of Boheydulang wasproposed as a bird sanctuary Then in 1977 proposals weremade to establish a state park Later in 1998 a collaboration(known as the Semporna Islands Project) was set up betweentheMarine Conservation Society (MCS) Sabah Parks WWFMalaysia and Nature Link In 2004 the Tun Sakaran MarinePark (TSMP) the seventh gazetted area under Sabah Parkswas established

Over 2000 residents that is Suluk Bajau and Bajau Lautlive in the TSMP This unique mix of people adds a furtherdimension to the area through their cultures lifestyleslegends and skills Most of the islands have settlementsmade up of semipermanent houses shacks and permanentstructures The main activities consist of fishing gardeningand seaweed cultivation The natural products harvested arefish shellfish medicinal plants fruits and timber Tourism iscurrently almost nonexistent although 21 prestigious scubadiving sites are reported [11]

Herein reticulated reefs commonly known as meshreefs are particularly studied Complex cellular patterns ofcoral reef walls or ldquoreticulatedmesh reefsrdquo often occupythe small fringing reefs and the internal lagoons of severalmodern platforms in the Tun Sakaran Marine Park They arenetworks with intervening depressions of linear or polyg-onal reef framework The cellular patterns are of differentsizes and scales cells 20m in diameter larger cells 200min diameter and the largest (on Selakan Bank) about 1 kmin diameter [12 13] These reef patterns may occur (1) onfringing reefs notably in the Arabian Gulf in the Red sea[14 15] and in Madagascar [16] (2) on barrier reefs in NewGeorgia [17] in Belize [18] on the Great Barrier Reef [19ndash21] and in Mayotte [16] and (3) on atolls in Hawaii (Maroreef [22]) in Fiji in French Polynesia [23 24] and in Kiribati(CarolineMillennium [25]) According toMontaggioni [23]and Purkis and Riegl [15] the reticulated shape of the meshreef could mimic a preexisting karst topography inheritedfrom an episode of lower sea level

The aim of the study is to analyze the health and stage ofdevelopment (old corals or recovery after mortality) of coralcommunities in two islands (reticulated reefs in lagoons andouter slopes) The density of associated organisms echino-derms gastropods bivalves and fishes are coanalyzed

2 Material and Methods

21 General Environmental Setting Sabah is a tropical coun-try characterized by uniform temperatures high humidityand copious rainfalls Situated between roughly 4∘N and 7∘Nunder the monsoon and typhoon belt Sabah is often referredto as ldquothe Land below the windrdquo Generally two seasons aredistinguished the rainy and the dry seasons although it ishot and humid (85ndash95) almost all year round except at

highaltitude In most parts of Sabah the wet season starts inNovember with the onset of the Northeast Monsoon (wintermonsoon) and ends towards March During the monthsof April to November the drier season occurs (southwestmonsoon or summer monsoon) and southwesterly windsover the northwest coast of Sabah may strengthen reaching40 kmsdothminus1 or more In Semporna temperatures rarely riseabove 32∘C during the day and rarely drop below 23∘C atnight Seawater is almost at a constant temperature (26ndash29∘C)

Tun Sakaran Marine Park (TSMP) also known as Sem-porna Islands Park lies off the east coast of Sabah Malaysiaat the entrance to Darvel Bay (latitude 4∘331015840N to 4∘421015840Nand longitude 118∘371015840E to 118∘511015840E) (Figure 1) It is the largestmarine Park in Malaysia and one of the prime coral reef sitesof the country It includes eight islands and associated reefs(Bodgaya Boheydulang Sebangkat and Selakan the sandcays of Maiga Sibuan and Mantabuan the patch reefs ofChurch and Kapikan) and covers an area of approximately340 km2 of sea and coral reefs and 10 km2 of land The largecentral high islands are formed of volcanic rock and areunique in Sabah The outlying islands are low limestone andcarbonate sand cays Their windward margin where coralreefs develop faces the dominant northeast trade winds

In the studied area the maximum high tide recordedin the tide tables for Semporna (Darvel Bay) is 21m andthe minimum tide 01m (httpwwwmareespechecomasmalaysiasemporna-darvel-bay)

22 Studied Sites Three monitoring sites were selected onGaya and Mantabuan Banks (Figure 1) Gaya Bank is a trueatoll about 8 km in diameter with a perimeter of 30 kmAn extinct volcano at the core of the bank forms twohigh relief islands Pulau Bodgaya and Pulau BoheydulangA reef complex and associated facies mainly developed onthe seaward side of the volcanic islands The deep lagoonmay represent the crater of the extinct volcano Intricatepolygonal patterns of reef walls or mesh reefs of differentsizes occupy the lagoon Mantabuan is a smaller bank 8 kmin perimeter located in the NE of Gaya bank

The studied sites (Figure 2) display different types of reefhabitats occurring on the carbonate platforms (1) site GEincludes a fringing reef exposed to currents (reef flat andinternal slope) (2) site GW includes a part of a large meshreef a reef rim and an outer slope (3) site M an outerclear-water site includes a part of the mesh reef a reef rimand an outer slope As the coral coverage decreases sharplybelow 15m in depth slopes were explored up to this depth(Figure 3)

In the results we will often separate one site into twosubsites the lagoon or reef flat and the slope Methods aredetailed in Table 1

Survey 1 Reef Zonation andMajor Benthic Categories To pro-vide information on reef zonation the habitat was observedand the major components were analyzed The surveyormeasured depth current speed and recorded the abundanceof main components of the sessile benthic community

Journal of Ecosystems 3

Table 1 The three surveys

Aim of the study Stations Method Period

Survey 1 Sessile fauna estimation of coverage and speciescomposition of major benthic categories

At each site LITs are parallelto the reef front

Line-Intercept-Transect =10m LIT 2011 May

Survey 2 Sedentary fauna estimation of densities ofechinoderms Tridacna and nudibranchs

1 belt-transect 05m apart the50m LIT

Belt-transect(50m by 1m) 2011 May

Survey 3Fishes abundance index for(1) Damselfish Stegastes(2) Parrotfish(3) Other fishes



LIT Line-Intercept-Transect

118∘10

998400E 118∘30

998400E 118∘50

998400E

4∘5999840000998400998400

N4∘3999840000998400998400

N4∘1999840000998400998400

N

Malaysia

Semporna

Tun Sakaran Marine Park

N

300km

20km

Land

Carbonated platforms

Coastal water

Open water


River

(a)

N

Selakan bank Gaya bank

Mantabuan bank

2km

M

GE

GW

4∘41998400N

4∘36998400N

118∘38

998400E 118∘43

998400E 118∘48

998400E

(b)

Figure 1 (a) Location of the Tun Sakaran National Park in Malaysia (b) focus on the three studied sites Gaya West (GW) Gaya East (GE)and Mantabuan (M)

Using the Line-Intercept-Transect (LIT) method [26 27] anestimation of the coverage (in ) of major benthic sessilecategories (such as primary producers living and dead coralssoft corals sponges rock sand and rubble) was made at a5-centimeter resolution LITs 50m in length parallel to thefront of the reef were run at different intervals (Figure 2)

Primary producers are cyanophytes and benthic algaebare dead corals are the dead substrates that are not coveredby any living organism visible to the naked eye

The different forms of corals have been noted as followsACB branching Acropora ACBB bottlebrush AcroporaACCOR corymbose Acropora ACST stout Acropora (Apalifera) ACTAB tabular Acropora CB branching non-Acropora (such as branching Porites Pocillopora Stylophoraand Seriatopora) CENC encrusting Non-Acropora CFfoliaceous corals (such as some Montipora some Pavona

and Turbinaria) CM massiveNon-Acropora (such as mas-sive Porites Faviidae) CMR mushroom corals (FungiaCycloseris Herpolitha etc) CSM submassive corals (someMontipora Psammocora etc) CST stout branching non-Acropora

In the lagoon LITs had to be located on the upper part ofthe mesh rims and not at random because of the great heightof the coral rims (Figure 3)

Unlike in a classical reef check we analyzed all thelayers of the communities (Figure 4) for example the deadsubstratums on which young corals are settled young coralson which perhaps algae are settled and so forth

Survey 2 Densities of Echinoids Holothurids and StegastesDensities of Echinoderms (echinids asterids holothuridscrinoids) Tridacna and nudibranchs were censused in belt-transects 05m strip on either side of each 50m LIT (50m2)


Table 2 Description of the three radial lines on the three sites Gaya West Gaya East and Mantabuan from the lagoon to the reef rim andthe outer slope

(a)

Gaya West Depth turbidity and currents Description of the general morphology

The mesh reef in the lagoon(stations GW1 to GW5)

Depth minus7mThe water is very turbidCurrent speed is low

In the lagoon corals form a mesh reef made of largecoral strips 4ndash7m high and 4-5m wide These coralrims rise from the seabed like large wallssurrounded by a sandy plain (Figure 5) The upperpart of the strips never emerges at low tide Coralvitality is low

The transition from the lagoon tothe reef rim

Depth minus2mIt is possible to walk on this part ofthe reef at low tideCurrents are very strong andas aconsequence the area has not beenanalyzed

From the lagoon toward the reef rim the depthdecreases sharply In this transition zone coralsoccur as circa metric patch reefs with great vitalityThese patches are scattered in a large sandy areaCoral colonies are likely massive and submassive andfew Acropora are visible

The reef rim Depth 0m

The reef rim is the highest part of the intertidalportion of the reef reaching a few centimeters abovethe upper level of coral growth Here it ishalf-necrotic and is less than 10m wide Corallinealgae cements the upper surface of the rim

The outer slope (stations GW6 andGW7)

From 0 to 15m turbidity is notimportant and coral diversity is thehighest observed at Gaya West

The outer slope is very steep and displays luxurianthard corals dominated by large heads ofnon-Acropora corals

The sandy plain Depth minus15mTurbidity is very important

At minus15m depth appears a plain characterized byslightly muddy fine sediment

(b)

Gaya East Depth turbidity and currents Description of the general morphology

The back reef (GE1 station)This part of the reef is made ofsand The currents may be highduring tides

No hard coral is visible Some scarce starfish occursthere

The reef flat (GE2 to GE5stations)

Depth minus4 to minus2mThe water is not turbid Becauseof the location behind a corridoroccurring between the twoislands Pulau Bodgaya andPulau Boheydulang the currentsmay be violent during tides

The reef flat is characterized by monospecific circa10m in diameter stands of branching Acroporafoliaceous Montipora cf florida or Pavona cactusThese large stands alternate with more diversifiedand smaller patches of branching submassive andmassive non-Acropora (Pocillopora StylophoraSeriatopora Psammocora Favia Favites etc)

The reef rim Depth increases slowly There is no reef rim visible

The inner slope (GE6 and GE7stations)

From minus2 to minus15m First the slopegently goes down from minus2 tominus7m for 50 meters and thenbecomes steeper from minus7 tominus17m

The two stations GE6 and GE7 are located in thesteep area (minus10 and minus15m)

The sandy plain Depth minus17mTurbidity is not important At minus17m appears a plain made of fine sediment

(c)

Mantabuan Depth turbidity and currents Description of the general morphology

The mesh reef into the lagoon(M1 to M4 stations)

Depth minus1 to minus5mThe water is not turbid

Into the lagoon corals form a mesh reef But thestrips 4m high and 3m wide are less developedthan in the Gaya West lagoon The coral communityis diverse and Acropora is abundant The upper partsof the strips are subtidal


(c) Continued


The transition from the lagoon tothe reef rim Depth minus2m high speed

currents

Toward the reef rim the depth decreases and anarea made of very fine sand occurs with small coralpatches of great vitality Corals occurring there arelikely massive and submassive but Acropora is muchmore abundant than in the similar zone of GayaWest

The reef rim Depth 0m The reef rim appears as a subtidal area

The outer slope (M5 to M7stations)

Depth from minus2 to minus15mLow turbidity and high currentspeed

First the slope gently goes down from minus2 to minus7m(M5 station) and then becomes steep from minus7 tominus15m (M6 and M7 stations) The diversity in coralsis less important than in Gaya West outer slope

The sandy plain Depth minus15mLow turbidity At minus15m appears a plain made of fine sediment

GW

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

(a)

GE

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

(b)

M

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

(c)

Figure 2 Aerial photography of the three sites with the radial lines and LITs We can see on aerial photographies of (a) GayaWest (GW) themesh reef the rim and the outer slope (b) Gaya East (GE) the reef flat and the inner slope and (c) Mantabuan (M) the mesh reef the reefrim and the outer slope


0

minus5

minus10

minus15

0

minus5

minus10

minus15

minus20

0

minus5

minus10

minus15

minus20

Dep

th (m

eter

s)D

epth

(met

ers)

Dep

th (m

eter

s)

GW

GE

M

1 2 3 4 5 6 7

1

1

2

2

3

3

4

4

5

5

6

6

7

7

1500m 650m 60m

200m 80m 230m 80m 10m 20m

800m 250m 180m

Mesh reef Patch reef zoneOuterreefflat

Outerreefflat

Outerslope

Back reef Reef flat Mesh reef Outer reef flat Innerslope

Mesh reef Patch reef zone Outer slope

Figure 3 The reef profile along the three radial lines Gaya west (GW) Gaya east (GE) and Mantabuan (M) and the location of the LITs onthe different geomorphological zones (not to scale)

3rd layer

2nd layer

1st layer

∙ Algae on living or dead corals (estimation of coverage)∙ Or coral dead by hypersedimentation ( of coverage)

∙ Living corals∙ Or recently dead corals∙ Or algae (calcareous or fleshy or filamentous)∙ Or alcyonids or others (sponges etc)∙ Biodetritic∙ Or pavement∙ Or ancientlyrecently dead corals∙ Or living corals (Acropora form or non-Acropora form)

Figure 4 The analysis on a LIT three layers (or more) can occur all are noted

All animals visible from a vertical position as well as thosehiding underneath ledges and in crevices were counted

Survey 3 Fish Abundance Fishes are classified into threecategories (1) the territorial damselfish Stegastes (2) par-rotfish and (3) other fishes Before echinoderm countingan abundance index is given to each of the three categoriesoccurring in the 50m by 1m belt (abundance index 1= 2-3individuals 2 = 5ndash16 individuals 3 = 17ndash64 individuals 4 =65ndash250 individuals 5 = more than 250 individuals)

23 Data Analysis Mean and standard deviation (SD) ofsessile fauna data for sites (Gaya West Gaya East andMantabuan) and subsites (reef flat mesh reef and inner and

outer slope) are calculated by taking one 10mLIT as a sampleFor sedentary fauna and fish the sample is the 50m by 1mbelt thus 50m2 Due to the multispecies nature of the dataand the design of the survey the most appropriate analysesare multivariate [28] To assess patterns in benthic data fromall stations during all years of survey we used the principalcomponent analysis (PCA)

3 Results

A first description of the radial lines was made to highlightthe different subzones (Table 2) and to decide where the LITshave to be settled


Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles

Figure 5 Gaya West lagoon a part of the mesh reef with associated fauna (no scale)

0102030405060

Rela

tive (

)

Different types of substrates on which the macroalgae are settled

RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand

Figure 6 Substrates on which are settled the macroalgae Allstations together RUB rubble ACR+ deadAcropora NACR+ deadnon-Acropora CXACR+ deadAcropora settled on dead substratesCXNACR+ dead non-Acropora settled on dead substrates

31 Primary Producers Among primary producers theencrusting calcareous algae are not recorded Although theyare ubiquitous their abundance is below the threshold of themeasurement

The total coverage of filamentous and macroalgae is verylow (below 5) except on the Gaya East-reef flat (Table 3)where they can be abundant in places (turfs in damselfishterritories or macroalgae patches)

Figure 6 shows that the macroalgae are mainly settled onrubble indicating that few living corals are in competition

Figure 7 Gaya West upper part of the mesh reef Caulerparacemosa crawling between living and dead Acropora

with primary producers On Gaya West-mesh reef Caulerparacemosa is abundant its vesicle-like branchlets along theerect axes crawling between corals (Figure 7) On GayaEast-reef flat large Ochrophytes such as Sargassum andTurbinaria occur on rubble

32 Bioconstructed Substrates At each site the studied radiallines spread into the lagoon the reef rim and the outer orinner slopes (Figures 2 and 3) Into the lagoon corals areorganized into a reticulated reef inGayaWest andMantabuan


0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50

Living coralsDead corals

(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)

Figure 8 Mean percentage (and SD) of coverage of living and dead corals at each station In Gaya West (a) and Gaya East (b) sites the twolast stations on the right are located on the outer slope In Mantabuan (c) the three last stations on the right are located on the outer slope

05

10152025303540

GW-MR GE-RF M-OS M-MR GE-IS GW-OS

Cov

erag

e (

)

Living corals

Non-Acropora

Figure 9 Increasing mean percentage of coverage of living coralswithAcropora and non-Acropora in lagoons (MRmesh reef andRFreef flat) and slopes (IS inner slope and OS outer slope) Sites GWGaya West GE Gaya East and M Mantabuan

while they forma reef flat inGayaEast (Table 2) InGayaWestandMantabuan lagoons LITs were placed on the upper partsof the mesh reef rims which is why the bioconstructed meancoverage reaches almost 80 (Table 4) Although other LITswere placed randomly on the outer slopes and on the fringing

Table 3 Percentage of coverage of primary producers (macroalgaeand filamentous algae) Mean (SD)

Sites Primary producersGaya West-mesh reef 31 (45)Gaya West-outer slope 00 (00)Gaya East-reef flat 81 (236)Gaya East-inner slope 27 (85)Mantabuan-mesh reef 40 (75)Mantabuan-outer slope 27 (54)

reef of Gaya East (reef flat and inner slope) the percentage ofcoverage of ldquobioconstructedrdquo substrates is exceptionally highgenerally around 70

33 Vitality of Corals The living coral coverage is approxi-mately le50 of the bioconstructed substrates on the threesites Considering the different subsites (Table 4) the vitalityof corals is the highest in the Gaya West outer slope and theGaya East inner slope (resp almost 60 and 50 of livingcorals within the bioconstructed substrates)

When all the different stations are taken into consider-ation (Figure 8) it appears that dead corals dominate overliving corals in almost all the stations except on the outer


Table 4 Percentage of coverage of living and dead corals on the different geomorphological zones and vitality (relative percentage of livingcoralstotality of bioconstructed substrates) Mean (SD)

Site-subzone Living corals Dead corals Bioconstructed substrates Vitality ()Gaya West-mesh reef 196 (142) 605 (212) 800 (222) 245Gaya West-outer slope 448 (199) 303 (113) 751 (260) 597Gaya East-reef flatlowast 259 (75) 408 (235) 667 (356) 388Gaya East-inner slope 350 (43) 370 (07) 720 (197) 486Mantabuan-mesh reef 340 (64) 456 (142) 796 (214) 427Mantabuan-outer slope 277 (97) 534 (157) 811 (212) 341lowastExcept LIT GE1 totally sandy Vitality = Living coralsDead corals lowast 100

0102030405060708090

100

GW-MR GE-RF M-MR GE-IS M-OS GW-OS

Rela

tive (

)

Sites

Relative percentage of Acropora in living corals

Non-AcroporaAcropora

Figure 10 Increasing mean relative percentage of coverage of livingAcropora left in lagoons (MRmesh reef and RF reef flat) and righton slopes (IS inner slope and OS outer slope) GW GayaWest GEGaya East and M Mantabuan

Table 5 Percentage of coverage of corals affected by hypersedimen-tation in the different geomorphological zones Mean (SD)

Site-subzone Corals affected by hypersedimentationGaya West-mesh reef 45 (34)Gaya West-outer slope 20 (36)Gaya East-reef flatlowast 132 (179)Gaya East-inner slope 74 (88)Mantabuan-mesh reef 05 (14)Mantabuan-outer slope 43 (82)lowastExcept LIT GE1 without corals

slope of Gaya West and on one station of the Gaya East-reefflat The stations located on Gaya West-mesh reef have thelower vitality

34 Living Corals

341 Acropora versusOther Corals As shown in Figure 9 theliving coral coverage is maximal on the outer slope of GayaWest and on the inner slope of Gaya East It is minimal onGayaWest-mesh reef whereas LITs have been spread only onthe upper part of coral rims

05

1015202530

Rela

tive (

)

Forms of living corals

Relative percentage of different forms of living corals

ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST

Figure 11 Living corals all sites together Relative percentage of thedifferent coral forms occurring on the three sites ACB branchingAcropora ACBB bottlebrush Acrop ACCOR corymbose AcropACST stout Acrop ACTAB tabular Acrop CB branching non-Acrop CENC encrusting corals CF foliaceous corals CMmassivecorals CMR mushroom corals CSM submassive non-Acrop CSTstout non-Acrop

Acropora colonies constitute 19 and 14 of the total cov-erage in Mantabuan-mesh reef and Gaya West-outer slopethat is 60 and 30 respectively of the relative coverage ofliving corals (Figure 10) In the other sites Acropora coverageis always lower than 30 of the living corals and even lowerthan 20 on Gaya West-mesh reef

342 Living Corals The Different Forms Considering allthe stations together (Figure 11) the dominant living coralforms are (1) the massive corals (-CM- notably here massivePorites and Faviidae) (2) the branching Acropora -ACB-(3) the branching non-Acropora (-CB- represented here bybranching Porites Pocillopora Stylophora Seriatopora etc)and (4) the foliaceous corals -CF- such as Montipora cfflorida Pavona cf cactus

Considering now each subsite (Figure 12) it appears thatthe highest diversity in coral forms is reached in the GayaWest-outer slope where the living coral coverage is thehighest massive and foliaceous corals dominate along withbranching Acropora

The diversity of forms is also high in the Mantabuan-mesh reef (with a dominance of branching Acropora) andin the Gaya East-reef flat where large monospecific standsof Montipora cf florida branching Acropora sp or Pavona cf


02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)

Figure 12 Percentage of coverage of different forms of living corals on each site (GW Gaya West GE Gaya East M Mantabuan MR meshreef RF reef flat OS outer slope IS inner slope) ACB branching Acropora ACBB bottlebrush Acrop ACCOR corymbose Acrop ACSTstout Acrop ACTAB tabular Acrop CB branching non-Acrop CENC encrusting non-Acrop CF foliaceous corals CM massive coralsCMR mushroom corals CSM submassive corals CST stout non-Acrop

Table 6 Number of individuals per 10m2 in the different subsites Mean (SD)

Site-subzone N ind10m2 Echinids Asterids Holothurids Crinoids Tridacna NudibranchsGaya West-mesh reef 100 (217) 06 (09) 02 (05) 02 (05) 01 (02) 05 (03)Gaya West-outer slope 04 (06) 14 (03) 06 (08) 04 (06) 02 (03) 14 (03)Gaya East-reef flatlowast 154 (139) 04 (03) 02 (03) 04 (08) 20 (10) 01 (01)Gaya East-inner slope 75 (75) 01 (01) 00 (00) 31 (27) 09 (10) 02 (03)Mantabuan-mesh reef 42 (35) 07 (07) 00 (00) 74 (137) 11 (13) 00 (00)Mantabuan-outer slope 38 (05) 07 (00) 05 (01) 105 (45) 07 (08) 01 (01)lowastExcept LIT GE1 exclusively sandy

Table 7 Estimation of the number of individuals per 100m2 in the different subsites using abundance index Mean (SD)

Site-subzone119873 ind100m2 Stegastes Parrotfish Other fishGaya West-mesh reef 711 (845) 60 (70) 171 (223)Gaya West-outer slope 345 (297) 345 (297) 1103 (1517)Gaya East-reef flatlowast 2338 (188) 41 (64) 56 (53)Gaya East-inner slope 2338 (230) 83 (74) 1365 (1146)Mantabuan-mesh reef 1851 (724) 480 (949) 261 (272)Mantabuan-outer slope 345 (297) 1528 (1375) gt250 (00)lowastExcept LIT GE1 exclusively sandy


Figure 13 Gaya East-reef flat large monospecific stand of Pavona

cactus occur (Figure 13) These stands alternate with patchesrich in massive corals made of large Porites cf lobata andnumerous Faviidae (Favia Favites Goniastrea etc)

The last three subsites Gaya West-mesh reef Gaya East-inner slope andMantabuan-outer slope show a low diversityin coral forms and are dominated by massive corals

35 Principal Component Analysis on Benthos Data (PCA)PCA on data shows that axes 1 and 2 explain 481 ofthe variance Factor 1 (PC1) accounting for 277 of thevariance is largely determined by the opposition betweenthe occurrence of a high coverage of living corals mostlyfoliaceous and branchingAcropora in stations GE2 GE4 andGE6 against the occurrence of juvenile living corals settledon bare substratums (thus a facies of recovery) in M1 andM5(Figure 14(a)) The coral communities on Gaya East-fringingreef are then more ancient than in Mantabuan

Factor 2 (F2) accounting for 204 of the variancedependsmainly on the opposition between a high coverage offoliaceous corals (GE2) and a high coverage of massive coralsin GE6 and GE7

Factor 3 (139 of var) opposes young branching Acrop-ora settled on dead substratums inM1 and young submassivecorals settled on dead substratums on GW1 and GW7(Figure 14(b)) both facies of recovery

36 Corals Affected by Hypersedimentation Corals affectedby hypersedimentation do not represent a high percentage ofcoverage (Table 5) The higher percentage is observed on thefringing reef of Gaya East reef flat and inner slope

37 Echinoderms Tridacna and Nudibranchs Echinids (orsea urchins) mostly Diadema are numerous on Gaya East-reef flat (more than 15 individual per m2) and on GayaWest-mesh reef (1 individual per m2) (Table 6) They aresurprisingly not abundant on Mantabuan-mesh reef (04individual per m2)

Asterids (or sea stars) and holothurids (or sea cucumbers)are scarce and are the most abundant on Gaya West-outerslope (01 individual per m2 resp)

Crinoids are themost abundant onMantabuan-mesh reefand -outer slope 07 and 10 individual per m2 respectively

Tridacna are abundant on the reef flat of Gaya East (02individual per m2) on Mantabuan-mesh reef and on GayaEast-inner slope (01 individual per m2)

The gastropods nudibranchs are abundant in Gaya West-outer slope and -mesh reef (01 and 005 individual per m2resp)

Considering the sedentary fauna on the whole(Figure 15) it appears that the two subsites of Gaya Eastand the subsite of Gaya West-mesh reef are dominated bysea urchins while Mantabuan (mesh reef and outer slope)is much more dominated by crinoids The subsite GayaWest-outer slope is the only one where the six groups offauna are slightly dominated by starfish and nudibranchs

38 Fish The damselfish Stegastes is particularly abundanton Gaya East-reef flat and -inner slope while parrotfishare the most abundant in Mantabuan-outer slope and moremoderately abundant on the mesh reef (Table 7) The otherfish are very abundant (gt250 ind per m2) on theMantabuan-outer slope

Regarding the total fish community (Figure 16) it appearsthat the Mantabuan-outer slope and the Gaya East-innerslope encompass numerous fishes but the Gaya East fishcommunity is dominated by Stegastes while the Mantabuanfish community is much more diversified and includesnumerous parrotfish

4 Discussion

41 Primary Producers The fast growth and turnover rates offleshy algae highlight their value as early warning indicatorsof reef degradation Except in some places on the reefflat of Gaya East and the upper parts of strips of GayaWest primary producers are scarce Thus not a single signof eutrophication has been recorded and algae cannot beconsidered as a potential danger to living corals in the studiedareas Nevertheless further investigations during the hot andvery wet season are needed to better understand the role ofthe primary producers on these reefs

42 Coral Vitality In the studied stations the bioconstructedcoverage (living and dead corals) is exceptionally high (67ndash81 of mean coverage) However the coral vitality varies Itreaches 60 on the GayaWest-outer slope (ldquofairrdquo condition)49 on the Gaya East-inner slope 43 on the Mantabuan-mesh reef and 24 on the Gaya West-mesh reef (ldquopoorrdquocondition)

On the Gaya West-mesh reef LITs have been spreadonly on bioconstructed substrates Nevertheless the biocon-structed coverage is relatively low because large basins of sandand rubble occur on the upper parts of the coral rims

Reticulated reefs display very different facies and vitalitylevels in Gaya and Mantabuan in the Gaya West lagooncoral rims (4ndash7m high 4-5m wide) occur in a very turbidenvironment and have a vitality of 24 In the Mantabuanlagoon corals are organized in smaller rims (4m high 3m


151050minus5minus10minus15

15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)

Figure 14 PCA on all data (GW Gaya West GE Gaya East M Mantabuan MR mesh reef RF reef flat OS outer slope IS inner slope)ACB branchingAcropora ACBB bottlebrushAcrop ACCOR corymboseAcrop ACST stoutAcrop ACTAB tabularAcrop CB branchingnon-Acrop CENC encrusting non-Acrop CF foliaceous corals CMmassive corals CMRmushroom corals CSM submassive corals CSTstout branching non-Acrop CXACB branching Acrop sur dead substrate CXACBB etc

02468

101214161820

GE-RF GW-MR GE-IS M-MR M-OS GW-OSSites

Sedentary fauna

EchinidsStarfishHolothurids

CrinoidsClamsNudibranchs

Num

ber o

f ind

ivid

uals

per 1

0 m2

Figure 15 Mean of number (nb) of individuals of sedentary faunarecorded on each site Continuum from dominance of sea-urchinson the left to dominance of crinoids on the right (GW Gaya WestGEGayaEastMMantabuanMRmesh reef RF reef flatOS outerslope IS inner slope)

wide) and occur in clear water the vitality is 43 andAcropora forms 60of the living corals According toEdingerand Risk [29] the occurrence of abundant Acropora is linkedto good health as Acropora is an r-strategist and a verysensitive form of coral Generally the Acropora-dominatedreefs harbor a high diversity of associated fish [30]

On the Gaya East-fringing reef (reef flat and inner slope)we observed the highest percentage of corals affected byhypersedimentation It is likely that the funnel shape of the

0

100

200

300

400

500

600

GW-MR GW-OS M-MR GE-RF GE-IS M-OSSites

Fish

Other fishParrotfishStegastes

Num

ber o

f ind

ivid

uals

per 1

0 m2

Figure 16 Mean of increasing abundance of fish (GW Gaya WestGEGayaEastMMantabuanMRmesh reef RF reef flatOS outerslope IS inner slope)

fringing reef and the strong current occurring between thetwo islands are at the origin of this increased sedimentationOn the reef flat the station GE2 (Gaya East-reef flat) is highlycharacterized by foliaceous corals (such as Pavona cf cactusandMontipora cf florida) that develop inmonospecific stands(circa 10m wide) On the other hand the Gaya East-innerslope is highly characterized by massive corals Gaya East(reef flat and inner slope) is also characterized by a highdensity of Stegastes damselfish (more than 200 per 100m2)This is an indication of coral degradation


In Peninsular Malaysia one of the major reasons forthe ldquopoorrdquo to ldquofairrdquo conditions of hard corals may be dueto increases in sedimentation [31] Along the west coast ofPeninsularMalaysia heavy sedimentation has caused serioussiltation problems in some river mouths and harbors [32]Millions of tons of sediment are transported annually bymajor rivers of the east coast of Sumatra and the westcoast of Peninsular Malaysia to the coastal waters of theMalacca Straits [33] Inmany localities these huge amounts ofsediment have changed the physical chemical and biologicalenvironment of the coral reefsTherefore coral vitality is verylow in the area [34]

In our study coral communities display a different state ofevolution corals in the lagoon of the Mantabuan-mesh reefare in state of recovery while on the Gaya East-reef flat theyhave reached amature statusThus large bleachingmortalityevents may have affected some reefs of the Tun SakaranMarine Park such as the Mantabuan ones

43 Sea Urchins Sea urchins such as Diadema are highlyrepresented on the Gaya East-reef flat and the Gaya West-mesh reef On the Gaya West-mesh reef macroalgae suchas Caulerpa spp occur within corals on Gaya East largeOchrophytes occur as metric sporadic stands These primaryproducers can attract Diadema a big consumer of seaweed

On the Mantabuan-mesh reef the sea urchins are scarceAs the corals are at the stage of recovery the sea urchinpopulations may have been affected by the same mortality asthe corals This would explain the low density of sea urchinson this reef Nevertheless the primary producers remainvery scarce in this area thus they may be controlled byherbivorous organisms Additional investigations are neededon sea urchin and fish herbivory to better understand thefunctioning of this beautiful area

44 Fishes The fish community is thriving on theMantabuan-outer slope where a myriad of fish exist aroundcorals In the three lagoons but also on the Gaya East-innerslope the Stegastes damselfish dominates On Mantabuanparrotfish are particularly abundant on the outer slope Theyare also present into the lagoon around the mesh reef

45 Coral Degradation Concerning the low vitality observedin the Gaya West-mesh reef Bell and Galzin [24] studyingreticulated reefs (Mataiva atoll Tuamotu Archipelago andFrench Polynesia) explained that several ldquolagoon basinsrdquosomewhat closed by the coral rims were entirely or partiallydead (the cause of mortality was uncertain) Perhaps thisldquoclosingrdquo process of the mesh reefs explains the coral degra-dation into the lagoons (ie Mataiva or Gaya bank)

Bodgaya is an inhabited island with a significant fishingpressure inside the lagoon (particularly on Gaya West) Thispressure might explain the scarcity of fish Thus an overfish-ing might speed up a disequilibrium in the ecosystem via(1) the reduction of fish (and perhaps sea urchin) herbivoryandor (2) the demise of key species Further studies areneeded (1) to estimate the impact of fishing and harvestingby the local population (2) to identify the target species and

(3) to research potential new species that could bring food ormoney to the locals If fishing by the local population impactstoo much on the lagoon communities a range of measures(such as aquaculture) have to be proposed to reduce fishingpressure and allow overexploited fish populations to recoverand ensure the maintenance of healthy populations

Experiments are also needed to estimate the real impact oflocal populations and tourism (mostly divers) For examplerestoration via coral nubbins may be attempted on somedegraded parts of the Gaya lagoon which would be protectedfrom human impact Also the Mantabuan lagoon may be anideal control site for experimentation

5 Conclusion

TSMP belongs to the Coral Triangle region Known asthe ldquoAmazon of the Seasrdquo and one of the worldrsquos mostdiverse and threatenedmarine ecosystems theCoral Triangleencompasses ocean areas in six countries in Southeast Asiaand the Pacific (Indonesia Malaysia Papua New GuineaPhilippines Solomon Islands and Timor-Leste) This regionis recognized as the global centre of marine biodiversity [35]and a global priority for conservation [36] Covering 57million square kilometers of ocean waters it contains at least500 species of reef-building corals in each ecoregion [37]More than 2200 species of fish (including the largest fishspecies the whale shark and the coelacanth) live there It alsoprovides a habitat to six out of the worldrsquos sevenmarine turtlespecies

The coral fauna of the TSMP shows a clear affinity withthat found on Indonesian reefs and several species (includingsome rare species ofAcropora and the uncommon caryophyl-liid Nemenzophyllia turbida [2]) are new to Malaysia Anumber of the sponges and soft corals found in the Gayalagoon are new undescribed species One sponge is reportedto be very unusual and possibly endemic to the Sempornareefs [9] Also Allen [1] found two new species of damselfishthere

Tun Sakaran Marine Park provides as much as 21prestigious scuba diving sites for example Tabah Siramba(Church Reef) Kapikan Reef Ribbon Reef Mantabuan andSibuan Reef Divers and snorkelers enjoy various attractivemarine lives such as turtles eagle rays barracudas bumpheadparrotfish nudibranchs and recently discovered sponges Itis difficult to describe the thrills that divers experience whenthey encounter marine macrolife in the Bodgaya lagoon Itis therefore important that something be done to preventfurther degradation of this natural wonderland

Acknowledgments

This research project carried out at the South-East AsiaCarbonate Research Laboratory (SEACARL) is financed bya Shell grant to Universiti Teknologi Petronas (Malaysia)French Embassy (Malaysia) University of South-Brittany(France) the Bretagne Region (CARBASIA project) and theConseil General du Morbihan The authors wish to thankSabah Parks for granting them a research permit to work


in Tun Sakaran Marine Park They wish to express theirgratitude to the staff of Sabah Parks Kota Kinabalu Sem-porna and Boheydulang station for their invaluable help andsupport during the field visits to Tun Sakaran Marine ParkThe authors also wish to thank the staff of the OceanographyInstitute ofUniversitiMalaysia Terengganu for their helpwiththe acquisition of the bathymetry dataThanks are also due toLawrence Schwarz for improving the English language of thepaper

References

[1] G R Allen ldquoMarine fishes of Sabah Northern Borneordquo inProposed Semporna Marine Park Environmental Evaluation forExpanded Tourism Development T D Meagher Ed pp 53ndash701992

[2] E M Wood Management Plan for the Semporna IslandsPark Proposed Through the Semporna Islands Project (SIP)Sabah Parks Marine Conservation Society (MCS) WWFMalaysia Nature Link 2001 httpwwwsempornaislandsprojectcompagespublicationshtm

[3] A Harborne D Fenner A Barnes M Beger S Harding andT Roxburgh Status Report on the Coral Reef of the East Coastof Peninsular Malaysia Coral Cay Conservation London UK2000

[4] T D Meagher Ed Proposed Semporna Marine Park Environ-mental Evaluation for Expanded TourismDevelopment ConbataPty Perth Australia 1992

[5] J D George and J George ldquoThe coral reefs of the BodgayaIslands (Sabah Malaysia) and Pulau Sipadan 4 Macroinverte-bratesrdquo Malayan Nature Journal vol 40 no 3-4 pp 225ndash2601987

[6] F A Dipper Notes on Sponges from the Proposed SempornaIslands Park A Report for the Semporna Islands Project MarineConservation Society 2001

[7] L M Marsh ldquoEchinoderms of the Tunku Abdul Rahman Parkreefs off Semporna and Pulau Sipadanrdquo in Proposed SempornaMarine Park Environmental Evaluation for Expanded TourismDevelopment T DMeagher Ed pp 55ndash59 Conbata Pty PerthAustralia 1992

[8] D JW Lane ldquoMacroinvertebrates 2rdquo inPulau Sipadan Reef Lifeand Ecology WWF Project No MYS 23392 pp 71ndash80 1994

[9] M Kelly ldquoNotes on sponge species identifications A report forthe Semporna Islands Projectrdquo 1999

[10] Semporna Marine Ecological Expedition December 2010httpawsassetspandaorgdownloadssmee factsheet dec2010pdf

[11] R F Baptist D Lojiwin and D Kubud Report on HumanSettlements in Semporna Islands Proposed Park SabahMuseum1998

[12] B J Pierson D Menier A Chalabi and T King King ldquoMor-phological indicators of growth stages in carbonates platformevolution comparison between present-day and Miocene plat-forms of Northern BorneoMalaysiardquo in Proceedings of the EGUGeneral Assembly vol 14 of Geophysical Research AbstractsEGU2012-12815-2 Vienna Austria April 2012

[13] DMenier B J Pierson A Chalabi T King King andM Pubel-lier ldquoMorphological indicators of structural controlrelative sea-level fluctuations and platform drowning on present-day andmiocene carbonate platformsrdquo Marine and Petroleum GeologyIn press

[14] J H Schroeder and D H Nasr ldquoThe fringing reefs ofPort Sudan Sudan 1 Morphology Sedimentology ZonationrdquoEssener Geographische Arbeiten vol 6 pp 29ndash44 1983

[15] S J Purkis and B Riegl ldquoSpatial and temporal dynamics ofArabianGulf coral assemblages quantified from remote-sensingand in situmonitoring datardquoMarine Ecology Progress Series vol287 pp 99ndash113 2005

[16] A Guilcher ldquoCoral reef geomorphologyrdquo Norois vol 142 no 1pp 234ndash236 1988

[17] D R Stoddart ldquoEcology and morphology of recent coral reefsrdquoBiological Reviews vol 44 pp 433ndash498 1969

[18] E G Purdy ldquoReef configurations cause and effectrdquo in Reefs inTime and Space L F Laporte Ed vol 18 of Society of EconomicPaleontologists and Mineralogists Special Publication pp 9ndash761974

[19] R A Slater and C V T Phipps ldquoA preliminary report on thecoral reefs of Lord Howe island and Elizabeth reef Australiardquoin Proceedings of the 2nd International Coral Reef Symposiumvol 2 pp 313ndash318 Miami Fla USA 1977

[20] J F Marshall and P J Davies ldquoInternal structure and Holoceneevolution of One Tree Reef southern Great Barrier Reefrdquo CoralReefs vol 1 no 1 pp 21ndash28 1982

[21] D Hopley S G Smithers andK E ParnellTheGeomorphologyof the Great Barrier Reef Development Diversity and ChangeCambridge University Press Cambridge UK 2007

[22] J C Kenyon C B Wilkinson and G S Aeby ldquoCommunitystructure of hermatypic corals at maro reef in the northwesternHawaiian islands a unique open atollrdquo Atoll Research Bulletinno 558 22 pages 2008

[23] L F Montaggioni ldquoHistory of Indo-Pacific coral reef systemssince the last glaciation development patterns and controllingfactorsrdquo Earth-Science Reviews vol 71 no 1-2 pp 1ndash75 2005

[24] J Bell and R Galzin ldquoInfluence of live coral cover on coral reeffish communitiesrdquo Marine Ecology Progress Series vol 15 pp265ndash274 1984

[25] K L Barott J E Caselle E A Dinsdale et al ldquoThe lagoon atCarolineMillennium atoll Republic of Kiribati natural historyof a nearly pristine ecosystemrdquo PLoS ONE vol 5 no 6 ArticleID e10950 2010

[26] HA Lucas andGA F Seber ldquoEstimating coverage andparticledensity using the line interceptmethodrdquoBiometrika vol 64 no3 pp 618ndash622 1977

[27] Y Loya ldquoPlotless and transect methodsrdquo in Coral ResearchMethods D R Stoddart and R E Johannes Eds pp 197ndash217UNESCO Paris France 1978

[28] K R Clarke and R M Warwick Change in Marine Commu-nities An Approach to Statistical Analysis and InterpretationPRIMER-E Plymouth UK 2nd edition 2001

[29] E N Edinger and M J Risk ldquoReef classification by coralmorphology predicts coral reef conservation valuerdquo BiologicalConservation vol 92 no 1 pp 1ndash13 2000

[30] P Chabanet H Ralambondrainy M Amanieu G Faure andR Galzin ldquoRelationships between coral reef substrata and fishrdquoCoral Reefs vol 16 no 2 pp 93ndash102 1997

[31] T Toda T Okashita T Maekawa et al ldquoCommunity structuresof coral reefs around peninsular Malaysiardquo Journal of Oceanog-raphy vol 63 no 1 pp 113ndash123 2007

[32] T E Chua and S A Ross ldquoCreating a shared vision for envi-ronmental management in the straits of Malaccardquo in TropicalMarine Environment Charting Strategies for the MillenniumF M Yusoff M Shariff H M Ibrahim S G Tan and S Y


Tai Eds pp 19ndash33 Malacca Straits Research and DevelopmentCenter (MASDEC) Faculty of Science and EnvironmentalStudies Putra University Selangor Malaysia 2002

[33] A Soegiarto ldquoPollution management and mitigation in theStraits of Malacca priorities uncertainties and decision mak-ingrdquo in Towards Sustainable Management of the Straits ofMalacca M Shariff F M Yusoff N Gopinath H M Ibrahimand R A N Mustapha Eds pp 503ndash518 Malacca StraitsResearch and Development Center (MASDEC) Faculty ofScience and Environmental Studies Universiti Putra MalaysiaSelangor Malaysia 2000

[34] A Soegiarto ldquoThe status of marine and coastal pollution inSoutheast Asiardquo in Coastal Area Management in SoutheastAsia Policies Management Strategies and Case Studies T EChua and D Pauly Eds vol 19 pp 71ndash75 ICLARM ManilaPhilippines 1989

[35] G R Allen ldquoConservation hotspots of biodiversity andendemism for Indo-Pacific coral reef fishesrdquo Aquatic Conserva-tion vol 18 no 5 pp 541ndash556 2008

[36] J C Briggs ldquoThe marine East Indies diversity and speciationrdquoJournal of Biogeography vol 32 no 9 pp 1517ndash1522 2005

[37] J E N Veron Corals of the World vol 3 Australian Institute ofMarine Science and CRR Qld Pty Townsville Australia 2000Edited by M Stafford-Smith

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

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Applied ampEnvironmentalSoil Science

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Atmospheric SciencesInternational Journal of



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International Journal of

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OceanographyInternational Journal of


The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014


ClimatologyJournal of


a record number of 43 species of mushroom corals (theprevious highest recorded richness in this family was 40species in Indonesia and Papua New Guinea) Furthermoresome new species were discovered (two species of shrimp andpossibly a number of gall crabs)

In Semporna the island of Bodgaya was first gazetted in1933 as a forest reserve and the island of Boheydulang wasproposed as a bird sanctuary Then in 1977 proposals weremade to establish a state park Later in 1998 a collaboration(known as the Semporna Islands Project) was set up betweentheMarine Conservation Society (MCS) Sabah Parks WWFMalaysia and Nature Link In 2004 the Tun Sakaran MarinePark (TSMP) the seventh gazetted area under Sabah Parkswas established

Over 2000 residents that is Suluk Bajau and Bajau Lautlive in the TSMP This unique mix of people adds a furtherdimension to the area through their cultures lifestyleslegends and skills Most of the islands have settlementsmade up of semipermanent houses shacks and permanentstructures The main activities consist of fishing gardeningand seaweed cultivation The natural products harvested arefish shellfish medicinal plants fruits and timber Tourism iscurrently almost nonexistent although 21 prestigious scubadiving sites are reported [11]

Herein reticulated reefs commonly known as meshreefs are particularly studied Complex cellular patterns ofcoral reef walls or ldquoreticulatedmesh reefsrdquo often occupythe small fringing reefs and the internal lagoons of severalmodern platforms in the Tun Sakaran Marine Park They arenetworks with intervening depressions of linear or polyg-onal reef framework The cellular patterns are of differentsizes and scales cells 20m in diameter larger cells 200min diameter and the largest (on Selakan Bank) about 1 kmin diameter [12 13] These reef patterns may occur (1) onfringing reefs notably in the Arabian Gulf in the Red sea[14 15] and in Madagascar [16] (2) on barrier reefs in NewGeorgia [17] in Belize [18] on the Great Barrier Reef [19ndash21] and in Mayotte [16] and (3) on atolls in Hawaii (Maroreef [22]) in Fiji in French Polynesia [23 24] and in Kiribati(CarolineMillennium [25]) According toMontaggioni [23]and Purkis and Riegl [15] the reticulated shape of the meshreef could mimic a preexisting karst topography inheritedfrom an episode of lower sea level

The aim of the study is to analyze the health and stage ofdevelopment (old corals or recovery after mortality) of coralcommunities in two islands (reticulated reefs in lagoons andouter slopes) The density of associated organisms echino-derms gastropods bivalves and fishes are coanalyzed

2 Material and Methods

21 General Environmental Setting Sabah is a tropical coun-try characterized by uniform temperatures high humidityand copious rainfalls Situated between roughly 4∘N and 7∘Nunder the monsoon and typhoon belt Sabah is often referredto as ldquothe Land below the windrdquo Generally two seasons aredistinguished the rainy and the dry seasons although it ishot and humid (85ndash95) almost all year round except at

highaltitude In most parts of Sabah the wet season starts inNovember with the onset of the Northeast Monsoon (wintermonsoon) and ends towards March During the monthsof April to November the drier season occurs (southwestmonsoon or summer monsoon) and southwesterly windsover the northwest coast of Sabah may strengthen reaching40 kmsdothminus1 or more In Semporna temperatures rarely riseabove 32∘C during the day and rarely drop below 23∘C atnight Seawater is almost at a constant temperature (26ndash29∘C)

Tun Sakaran Marine Park (TSMP) also known as Sem-porna Islands Park lies off the east coast of Sabah Malaysiaat the entrance to Darvel Bay (latitude 4∘331015840N to 4∘421015840Nand longitude 118∘371015840E to 118∘511015840E) (Figure 1) It is the largestmarine Park in Malaysia and one of the prime coral reef sitesof the country It includes eight islands and associated reefs(Bodgaya Boheydulang Sebangkat and Selakan the sandcays of Maiga Sibuan and Mantabuan the patch reefs ofChurch and Kapikan) and covers an area of approximately340 km2 of sea and coral reefs and 10 km2 of land The largecentral high islands are formed of volcanic rock and areunique in Sabah The outlying islands are low limestone andcarbonate sand cays Their windward margin where coralreefs develop faces the dominant northeast trade winds

In the studied area the maximum high tide recordedin the tide tables for Semporna (Darvel Bay) is 21m andthe minimum tide 01m (httpwwwmareespechecomasmalaysiasemporna-darvel-bay)

22 Studied Sites Three monitoring sites were selected onGaya and Mantabuan Banks (Figure 1) Gaya Bank is a trueatoll about 8 km in diameter with a perimeter of 30 kmAn extinct volcano at the core of the bank forms twohigh relief islands Pulau Bodgaya and Pulau BoheydulangA reef complex and associated facies mainly developed onthe seaward side of the volcanic islands The deep lagoonmay represent the crater of the extinct volcano Intricatepolygonal patterns of reef walls or mesh reefs of differentsizes occupy the lagoon Mantabuan is a smaller bank 8 kmin perimeter located in the NE of Gaya bank

The studied sites (Figure 2) display different types of reefhabitats occurring on the carbonate platforms (1) site GEincludes a fringing reef exposed to currents (reef flat andinternal slope) (2) site GW includes a part of a large meshreef a reef rim and an outer slope (3) site M an outerclear-water site includes a part of the mesh reef a reef rimand an outer slope As the coral coverage decreases sharplybelow 15m in depth slopes were explored up to this depth(Figure 3)

In the results we will often separate one site into twosubsites the lagoon or reef flat and the slope Methods aredetailed in Table 1

Survey 1 Reef Zonation andMajor Benthic Categories To pro-vide information on reef zonation the habitat was observedand the major components were analyzed The surveyormeasured depth current speed and recorded the abundanceof main components of the sessile benthic community














118∘10

998400E 118∘30

998400E 118∘50

998400E

4∘5999840000998400998400

N4∘3999840000998400998400

N4∘1999840000998400998400

N

Malaysia

Semporna


N

300km

20km

Land


Coastal water

Open water


River

(a)

N


Mantabuan bank

2km

M

GE

GW

4∘41998400N

4∘36998400N

118∘38

998400E 118∘43

998400E 118∘48

998400E

(b)











(a)















(b)












(c)






(c) Continued



currents







GW

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

(a)

GE

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

(b)

M

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

(c)



0

minus5

minus10

minus15

0

minus5

minus10

minus15

minus20

0

minus5

minus10

minus15

minus20

Dep

th (m

eter

s)D

epth

(met

ers)

Dep

th (m

eter

s)

GW

GE

M

1 2 3 4 5 6 7

1

1

2

2

3

3

4

4

5

5

6

6

7

7

1500m 650m 60m

200m 80m 230m 80m 10m 20m

800m 250m 180m


Outerreefflat

Outerslope




3rd layer

2nd layer

1st layer








3 Results



Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles


0102030405060

Rela

tive (

)


RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand









0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics



























118∘10

998400E 118∘30

998400E 118∘50

998400E

4∘5999840000998400998400

N4∘3999840000998400998400

N4∘1999840000998400998400

N

Malaysia

Semporna


N

300km

20km

Land


Coastal water

Open water


River

(a)

N


Mantabuan bank

2km

M

GE

GW

4∘41998400N

4∘36998400N

118∘38

998400E 118∘43

998400E 118∘48

998400E

(b)











(a)















(b)












(c)






(c) Continued



currents







GW

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

(a)

GE

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

(b)

M

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

(c)



0

minus5

minus10

minus15

0

minus5

minus10

minus15

minus20

0

minus5

minus10

minus15

minus20

Dep

th (m

eter

s)D

epth

(met

ers)

Dep

th (m

eter

s)

GW

GE

M

1 2 3 4 5 6 7

1

1

2

2

3

3

4

4

5

5

6

6

7

7

1500m 650m 60m

200m 80m 230m 80m 10m 20m

800m 250m 180m


Outerreefflat

Outerslope




3rd layer

2nd layer

1st layer








3 Results



Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles


0102030405060

Rela

tive (

)


RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand









0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics
















(a)















(b)












(c)






(c) Continued



currents







GW

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

(a)

GE

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

(b)

M

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

(c)



0

minus5

minus10

minus15

0

minus5

minus10

minus15

minus20

0

minus5

minus10

minus15

minus20

Dep

th (m

eter

s)D

epth

(met

ers)

Dep

th (m

eter

s)

GW

GE

M

1 2 3 4 5 6 7

1

1

2

2

3

3

4

4

5

5

6

6

7

7

1500m 650m 60m

200m 80m 230m 80m 10m 20m

800m 250m 180m


Outerreefflat

Outerslope




3rd layer

2nd layer

1st layer








3 Results



Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles


0102030405060

Rela

tive (

)


RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand









0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics















(c) Continued



currents







GW

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

118∘42

99840030

998400998400E 118∘43

9984000998400998400E 118

∘43

99840030

998400998400E 118∘44

9984000998400998400E

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

4∘3899840030998400998400

N4∘389984000998400998400

N4∘3799840030998400998400

N4∘379984000998400998400

N

(a)

GE

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

118∘46

99840040

998400998400E 118∘46

99840050

998400998400E 118∘47

9984000998400998400E

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

4∘3699840040998400998400

N4∘3699840030998400998400

N4∘3699840020998400998400

N

(b)

M

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

118∘47

99840045

998400998400E 118∘48

9984000998400998400E

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

4∘399984000998400998400

N4∘3899840045998400998400

N4∘3899840030998400998400

N4∘3899840015998400998400

N

(c)



0

minus5

minus10

minus15

0

minus5

minus10

minus15

minus20

0

minus5

minus10

minus15

minus20

Dep

th (m

eter

s)D

epth

(met

ers)

Dep

th (m

eter

s)

GW

GE

M

1 2 3 4 5 6 7

1

1

2

2

3

3

4

4

5

5

6

6

7

7

1500m 650m 60m

200m 80m 230m 80m 10m 20m

800m 250m 180m


Outerreefflat

Outerslope




3rd layer

2nd layer

1st layer








3 Results



Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles


0102030405060

Rela

tive (

)


RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand









0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics















0

minus5

minus10

minus15

0

minus5

minus10

minus15

minus20

0

minus5

minus10

minus15

minus20

Dep

th (m

eter

s)D

epth

(met

ers)

Dep

th (m

eter

s)

GW

GE

M

1 2 3 4 5 6 7

1

1

2

2

3

3

4

4

5

5

6

6

7

7

1500m 650m 60m

200m 80m 230m 80m 10m 20m

800m 250m 180m


Outerreefflat

Outerslope




3rd layer

2nd layer

1st layer








3 Results



Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles


0102030405060

Rela

tive (

)


RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand









0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics















Sponge

Sponge

Pocillopora

Acropora

Acropora

Acropora

Montipora

Montipora

Montipora

Faviidae

Porites

Porites

Porites

Porites

Porites

Diadema

Sea stars

Shark

Butterflyfish

Butterflyfish

Angelfish

Bonitos

Turtles


0102030405060

Rela

tive (

)


RUB

CXN

ACR+

ACR+

NAC

R+

CXA

CR+

CX Sand









0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics















0

20

40

60

80

100

50 150

250

350

450

550

650

750

850

950

1050

1150

1250

1350

1450

1550

1650

1750

1850

1950

2050

2150

2250

2350

2450

Cov

erag

e (

)

Radial axis (m)

Gaya West

minus50


(a)

Cov

erag

e (

)

0

20

40

60

80

100

0 50 100 150 200 250 300 350 400 450 500 550Radial axis (m)

Gaya East

minus50


(b)

Cov

erag

e (

)

0

20

40

60

80

100

50 150 250 350 450 550 650 750Radial axis (m)

Mantabuan


minus50

(c)


05

10152025303540


Cov

erag

e (

)

Living corals

Non-Acropora











0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics

















0102030405060708090

100


Rela

tive (

)

Sites







34 Living Corals


05

1015202530

Rela

tive (

)



ACB

ACBB

ACCO

R

ACST

ACTA

B CB

CEN

C CF CM CMR

CSM

CST







02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics















02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-MR

(a)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GW-OS

(b)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-RF

(c)

05

101520

ACBACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

GE-IS

(d)

02468

10ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-MR

(e)

0

5

10

15ACB

ACBB

ACCOR

ACST

ACTAB

CBCENC

CF

CM

CMR

CSM

CST

M-OS

(f)






















4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics






























4 Discussion







15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics
















15

10

5

0

minus5

minus10

minus15

PC2

PC1

M1L-MR

CXACBM5OS

GE5FR-FCXCM

CXCB

M6OS

GE2FR-MR

CF

GW6OSM2L-MRACB

GE4FR-MR

GE3FR-MR

CB

CM

GE7FR-ISGE6FR-IS

(a)


15

10

5

0

minus5

minus10

minus15

PC1

PC3

M1L-MR

CXACB

M5OSCXCM

CXCB

CXCSM

GW1L-MRGW7OS

GE2FR-MRGW6OS

M2L-MR

ACBGE4FR-MR

GE3FR-MRCF

CB CMGE7FR-IS

GE6FR-IS

(b)


02468

101214161820


Sedentary fauna



Num

ber o

f ind

ivid

uals

per 1

0 m2




0

100

200

300

400

500

600


Fish


Num

ber o

f ind

ivid

uals

per 1

0 m2













5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics
























5 Conclusion




Acknowledgments




References












































Journal of












Volume 2014

Advances in









Geophysics
















References












































Journal of












Volume 2014

Advances in









Geophysics

























Journal of












Volume 2014

Advances in









Geophysics


















Journal of












Volume 2014

Advances in









Geophysics














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