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Coral acclimatization to disturbance [Elektronische Ressource] / submitted by Cornelia Roder

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Coral Acclimatization to DisturbanceDissertation submitted by Cornelia Roder In partial fulfillment of the requirements for the degree of Doctor of Natural Sciences (Dr. rer. nat.) Faculty of Biology / Chemistry, Bremen University Bremen, February 2010 First Examiner: Prof. Dr. Claudio Richter Alfred Wegener Institute, Bremerhaven, Germany Second Examiner: Prof. Dr. Wilhelm Hagen University of Bremen, Bremen, Germany Additional Examiner I: Prof. Dr. Kai Bischof men, Bremen, Germany Additional Examiner II: Dr. Somkiat Khokiattiwong Phuket Marine Biological Center, Phuket, Thailand Front page picture: ERS-2 (C-band, VV) SAR image of solitons in the Andaman Sea, 11 February 1997 at 0359 UTC. From Jackson (2004) © ESA 1997 AcknowledgementsAcknowledgements Zuoberallerersternächst möchte ich mich bei meinem Betreuer und Mentor Prof. Claudio Richter bedanken – für alle gegebenen Gelegenheiten und ermöglichten Möglichkeiten, für oftmals einfalls- und erfindungsreiche Denkanstöße, geniale Ideen und konstruktive Kritik, für die Mühe, mir zu zeigen, was WIRKLICH Forschung ist, wie’s geht und wie weniger. Ich habe in den vergangenen drei Jahren so viel gelernt, und das nicht zuletzt aufgrund Deiner Hilfestellungen – danke für Dein Vertrauen und Deine Unterstützung! Vielen herzlichen Dank an Prof. Wilhelm Hagen für die Zusage, meine Doktorarbeit zu gutachten und Teil meines Prüfungskomitees zu sein! Auch ein großes Dankeschön an Prof.

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Coral Acclimatization to

Disturbance

itted by Dissertation subm

Cornelia Roder

ents for the degree of ent of the requiremIn partial fulfillm

Doctor of Natural Sciences (Dr. rer. nat.)

istry, Bremen University Faculty of Biology / Chem

bruary 2010 en, FeBrem

First Examiner: Prof. Dr. Claudio Rchter i

Alfred Wegener Institute, Bremerhaven, Germany

Second Examinerilhelm Hagen : Prof. Dr. W

any en, Germen, BremmUniversity of Bre

Additional Examiner I: Prof. Dr. Kai Bischof

en, Germen, BremmUniversity of Breany

Additional Examiner IIat Khokiattiwong ki: Dr. Som

Phuket Marine Biological Center, Phuket, Thailand

Front page picture: ERS-2 (C-band, VV) SAR image of solitons in the Andaman Sea, 11 February

1997 at 0359 UTC. From Jackson (2004) © ESA 1997

Acknowledgements

ementsgAcknowled

Zuoberallerersternächst möchte ich mRichter bedanken – für alle gegebenen Gelegeich bei meinemnheiten und erm Betreuer und Mentor Prof. Claudio öglichten Möglichkeiten, für
oftmfür die Mühe, mials einfalls- und erfindungsreiche Denkansr zu zeigen, was WIRKLICH Forschung ist, wie’s geht und wie weniger. Ich töße, geniale Ideen und konstruktive Kritik,
habe in den vergangenen drei Jahren so viel gelernt, und das nicht zuletzt aufgrund Deiner Hilfestellungen – danke für Dein Vertrauen und Deine Unterstützung!

gutachten und Teil mVielen herzlichen Dank an Prof. Weines Prüfungskomilhelmitees zu sein! Hagen für die Zusage, meine Doktorarbeit zu

usage als Auch ein großes Dankeschön an Prof. Kai Bischof für seine so kurzfristigen ZPrüfer.

I am very much indebted to Dr. Somkiat Khokiattiwong not only for agreeing to be a member
of my commappreciate your timittee, but also for sharing his knowe and effort by coordinating our stays in Thailand in a perfect mledge about Solitons and the Andaman Sea. I anner. A
at the PMBC, especially Aum and Ann who are in great parts big thanks also to your teamle for the successful accomplished works in the ORCAS project. responsib

I am also very grateful for the comespecially during our stay in 2007 – for providing us with diving equipmpanionship of Niphon Phongsuwan and his teament, showing us ,
ilans, for indicating the perfect spots for our research and for teaching mearound the Sime Thai. somKhop khun kah!

ginnen und Freundinnen Carin Jantzen, Gertraud ine KolleeMein riesengroßer Dank geht an mSchmidt und Yvonne Sawall; alle bereits oder alsbald auch mit einem Dr. vor dem Namen
versehen für die gemeinsame Arbeit, das gemeinsame Planen, Diskutieren, Frachten,
Schleppen, Tauchen, Schreiben....für eure offenen Ohren und die Unterstützung in den verschiedensten Bereichen.

Tobias Funke – unwegdenkbarer Bestandteil der Arbeitsgruppe und Helfer für alle kniffligen Lagen und Fragen!

währende ern Ittekkot und Dr. Ursel Selent für alle immVielen Dank and Prof. VenugopalaUnterstützung am ZMT! Für meine Zeit am ZMT möchte ich mich zudem bei allen Kollegen
immbedanken: mer gelösten Comeputerprobleminen fellow-PhDs für den gutenchen; Dorothee Dasbach und Matthias Birkicht für die Zusammenhalt; Uli Pint für die zahllosen
reichenden Einblicke in Methoden und Labor und die weitriesige Unterstützung imann und Larissa e, Silke Eilem Petra Käpnik, Gaby BöhmAnalysetechniken; außerdemschen Belangen. Ausserdem ein extra riesiges Dsikowitzky für Unterstützung in organisatoriDankeschön an Dr. Tim Jennerjahn, für seinen Einsatz und auch für all die in mich investierte
Zeit und konstruktive Beratung zum Thema „Biogeochemie für Ökologen oder Leute mit
weniger Ahnung“!

auch da wart! Dres. Iris und Mark Wunsch für ihre Unterstützung in Thailand – ich war sehr froh, dass ihr

ementsgAcknowled

For the timChinese partners, especially Zhongjiee and fun shared in China, I’d like to thank th Wu and the HPMDDI Haikou, Hainan. e ZMT’s LANCET team and the

Muchas gracias a todos del CIMAR y Parque Nacional Cahuita, to Prof. Jorge Cortés, Dr. Carlos Jiménez and especially Dr. Rubén Lara for valid criticism on my first ever manuscript
ission. subm

Vielen Dank auch an Werner Wosniok für seine statistische Beratung.

, Retter in der Not, Pinatubobesänftiger oder Stöff – Doppelter Boden, Fels in der Brandungzen für Deine huere-geniale Unterstitzig! Hereinfach Held: ich danke Dir von ganzem

Last buvorneweg met not least einenine Eltern, Dres. A Gruß ins Schwobeländle: denngela und Stephan Roder, für ihre uneingeschränkte größten Dank an meine Familie, allen
ine Geschwister Babse, Paul, enz klar an mUnterstützung an wirklich allen Fronten und gaEvchen und Chele. Ohne euch: never ever....

ooperationsprojekte Diese Doktorarbeit wurde finanziert durch die K

s of Tropical China: Hainan), EcosystemLANCET (Land-Sea Interactions along Coastal 03F0457A (BMBF)

und

ORCAS (Ocean-Reef Coupling in the Andaman Sea), RI 1074/7-1 (DFG)

Zusammenfassung

gZusammenfassun

peratur, Salzgehalt, Licht Die globale Verbreitung tropischer Flachwasserriffe wird durch Temrtel entlang 30° Nord und Süd des Äquators beschränkt. und Aragonitsättigung auf einen Güung, Innerhalb dieser geographischen Grenzen spielen viele weitere Faktoren wie Ström oder gelöstemPlankton und partikuläremNährstoffgehalt oder das Vorkommen von organischem Material eine wichtige Rolle für die Entstehung und das Fortbestehen von e für deren Kondition. Riffen, sowi

gekoppelt, verstärken sich gegenseitig oder teinander iViele dieser Faktoren sind eng mund indirekte Auswirkungen auf hrere direkte eschwächen sich ab und haben oft gleich mlevel. Erhöhte Nährstoffeinträge (Kapitel I, IV, VI) verstärken us- oder ÖkosystemOrganismbionten in Korallen (Zooxanthellen) und deren ehrung der Symnicht nur die Vermgrösste KPhotopigmeonkurrenten um Licht und Raumnte, sondern fördern auch die Ausbreitung von benthischen Algen, welche als gelten. Übermässig angetriebenes
einer vermPhytoplanktonwachstum kann ausserdeminderten Lichtzufuhr für die Benthosgem zu einer Trübung der Weinschaft führen. Strömassersäule und somungen (Kapitel II) it zu
it den gebungswasser und erhöhen somverringern die Grenzschicht zwischen Koralle und UmGas- und Nährstoffaustausch. Ausserdem gewährleisten sie den Nachschub von e, Material für Photosynthese und aktive Nahrungsaufnahm und organischeminorganischemsowie den Abtransport ausgeschiedener Stoffe (Kapitel I, II, III). Zudem sind Strömungen für
die Resuspension und Vwelche beim Verweilen in der Wassersäule Terbreitung von Sedimrübung verursachen und beim Absienten und assoziierten Teilchen verantwortlich, nken eine
Erstickungsgefahr für benthische Organismen darstellen. Heftige Strömungen oder Wellen
eine Bruchgefahr für Korallenskelette darstellen. können ausserdem

Viele dieser lokalen Einflüsse können sowohl natürlichen (Kapitel I bis V) als auch end der anthropogene Einfluss hräanthropogenen (Kapitel VI, VII) Ursprungs sein. W ti(abgesehen von globalen Veränderungen wie Temperaturanstieg oder Ozeanversauerung) mt, sind offshore-Riffe häufig stark von ihrer ozeanischen Distanz zur Küste abnimmUmgebung bestimmFunktionalität eines Riffes und seiner Bewohner mt. Je nach Art, Intensität, Häufigkeit und Dauer der Störung wird die ehr oder minder beeinflusst. Einige
Faktoren wie Saisonalität oder Gezeiten können dabei sowohl für küstennahe (Kapitel I, VI), als auch für offshore-Riffe (Kapitel I bis VI) ein Rolle spielen.

gZusammenfassun

gebung beeinflusst (Kapitel I bis V), Offshore-Riffe werden stark durch ihre ozeanische Umwelche oftmals sehr variable sein kann. 60 km vor der Westküste Thailands werden die Riffe
entlang der Westseite der Similaninseln von brechenden Internen Wellen Grosser Amplitude
(Large Amplitude Internal Waves, LAIW) heimgesucht. Im Gegensatz dazu sind die Riffe auf
der Ostseite der Inseln im geringeren Masse dem LAIW-Einfluss ausgesetzt. Die LAIW,
welche impulsartig kaltes, sauerstoffarmes und CO2- sowie nährstoffreiches Tiefenwasser in
us der dort siedelnden Korallen nachhaltig.die Riffe eintragen, verändern den Metabolism

-Frequenz (Kapitel I), die sich innerhalb derselben Saison -Intensität und WAbhängig von LAI es zu teilweise extrem hohen tverschiedener Jahre drastisch unterscheiden kann, kommentbildung, die und PigmNährstoffeinträgen. Diese fördern zwar ZooxanthellenwachstumStrömPhotosyntheseleistung aber wird reduziert (Kapitel I,ungen erhöhen den Fluss gelöster und parti III, IV). Starke LAIkulärer organischer Stoffe und unterstützen W-verursachte
ngsaufnahme der Korallen (Kapitel II). Gleichzeitig bewirkt die it die aktive Nahrusomher Nährstoffe auch eine erhöhte Abgabe erhöhte Verfügbarkeit organischer und inorganisc durch die Korallen (Kapitel III). Die )gelöster und partikulärer Substanzen (SchleimKomWestseite führen ausserdembination aus hohen Zooxanthellendichten und aktiver Nahrungsaufnahm zu höherer Biomasse und Energiespeicher in den Korallen e auf der
der Ostseiten belastbarer und Gegensatz zu den Korallenachen sie im(Kapitel I, II). Diese msituationen (Kapitel II). Der Preis für die höheren überlebensfähiger in Extreme Respiration und geringere, kostenintensivere assen ist eine erhöhtEnergiespeicher und BiomPhotosynthese (Kapitel III). Auch die Kalzifgeringeren Aragonitsättigung des aufgetriebenen Tiefenwassersizierung unter LAIW dezim-Einfluss ist aufgrund der iert (Kapitel III).
Zudem sind LAIW-eigene niedrige Temperatur und geringer Sauerstoffgehalt für langsameres
-usätzlich zum SWhotosyntheseraten verantwortlich. Z und niedrigere PKorallenwachstum die Flachwasserbereiche der westlichen Inselseiten beeinflusst, führen Monsun, der vor allemdie vielseitigen LAIW-Einflüsse zu einem stark reduzierten Riffwachstum (Kapitel V), mit
geringeren Bedeckungsgraden, aber auch höherer Biodiversität. Die Anpassungen an den LAIWhaben gezeigt, dass sich diese auch unter nicht LAI-Einfluss variieren zwischen den verschieW-Konditiondenen Korallenarten (Kapitel I). Inkubationen en entfalten (Kapitel III),
bevor eine Anpassung an eine anderweitige Umgebung stattfindet sich bei der hohen bt die Frage, ob esl II). Gleichwohl bleient Kapite(TransplantationsexperimFlexibilität der Korallen unter LAIW-Einfluss um eine metabolische Anpassung oder gar eine
genetische Adaptation handelt. Die Studien über die Anpassungsfähigkeit von Riffen an LAIW-Gegebenheiten dienen auch als Grundlage zum Verständnis, wie sich Riffe im

gZusammenfassun

Hinblick auf den globalen Klimawandel (Temperaturanstieg und Ozeanversauerung)
entwickeln könnten.

r Karbikküste Costa Ricas (Kapitel VI) sind enten an deEinträge von Nährstoffen und Sedimstark von den Ausmassen der Trocken- und Regenzeit bestimmt. Mit ansteigendem Regen
ehrt Nährstoffe aus stark gedüngten Böden bewirtschafteter Felder in küstennahe werden vermGewässer ausgespült und mit dem Fluss ins Meer eingetragen. Dort werden sie mit Hilfe der
arks von Cahuita verbreitet und nachweislich ffgebiete des Nationalpiungen bis in die RStrömvon den Korallen aufgenommen. Durch den exzessiven Anbau von Monokulturen im e deutlich wird. Als ateinträgKüstenraum steigt auch die Bodenerosion, was durch hohe Silikit Flüssen und Strömungen hohe Mengen partikulären Materials in Konsequenz gelangen mdie Riffe, wo sie die Wassersäule trüben, Korallen bedecken und letztendlich ersticken. Nationalpark r Riffen ähnlicher geographischer Ausrichtung sind die imiVerglichen mCahuita starker Eutrophierung ausgesetzt.

ordostküste Hainans, China, fordern die Folgen der raschen Auch entlang der NKüstenentwicklung ihren Tribut (Kapitel VII). Starke Eutrophierung und Sedimentation durch
sant wachsenden Küstensiedlungen unbehandelte Abwässer aus Aquakultur und rassersäule. Exzessive Aquakultur kann aüberdüngen die Küstengewässer und trüben die WArten oder Pestizide führen. Ein weiteres zusätzlich zur Einführung von Krankheiten, fremder grosses Problem ist die offensichtliche weite Verbreitung destruktiver Fischereimethoden, wie
itfischerei, welche bereits einen grossen Teil der Riffe zerstört hat. Die mzum Beispiel Dynaöden sind unvorteilhaft für das Ansiedeln von Korallenlarven. Die so enstandenen losen B ebination aus Überfischung und hohen Nährstoffkonzentrationen stellt eine enormKoms durch herbivore Fische nicht Gefahr für Riffe dar, da eine Kontrolle des Algenwachstummehr gewährleistet ist und die Algen sich auf dem losen Boden im nährstoffreichen Wasser
besser durchsetzen können als Korallen. Überlebende Korallen haben eine geringere it Korallen aus übung der Wassersäule, vergleichbar mrPhotosyntheseleistung durch die Tentierung erfordert zusätzlich einen hohen Energieaufwand sser. Die starke Sedima Wtieferemder Korallen, welche sich durch Schleimproduktion von ablagerndem Sediment säubern
müssen.

Untersuchungsgebiet in Costa versität im der Korallenlebendbedeckung und DieEine AbnahmRiffsystemRica und eine Verschiebung von einem Korallen do in Hainan sind deutliche Anzeichen, dass die Ökosystemminierten zu einem Algen dominierten e gefährdet sind und unter
ng drohen zu kollabieren. Da die Störungen dem ständig wachsenden Druck über kurz oder la

hier - im

Gegensatz zu den im

Zusammenfassun

pulsartigen Störungen der LAIW - chronisch sind und sich im

Laufe der Zeit verstärken, ist ihre Bedrohung akut.

g

Summary

ySummar

shallow water coral reefs within the tropical belt of 30° The global distribution of tropicalNorth and South is determined by temperature, salinity, light and aragonite saturation state.
Within these geographical limits various additional factors such as currents, nutrient
portant for conditions or abundance of plankton and particulate or dissolved organics are iment and persistence of reefs as well as for their condition. the developm

inish each other and often have sely related, enhance or dimMany of these factors are clot or ecosystem level at once. Enhanced nutrienpacts on organismseveral direct or indirect imbiotic algae inputs (Chapter I, IV, VI) not only increase the density of the coral’s symtion of benthic algae ents, but also enhance the distribu(zooxanthellae) and their photopigmwhich compete withto nutrification can further lead to a turbid water colu corals for light and space. Emn resultinxcessive phytoplankton growth as a response g in decreased light
inish the boundary layer ents (Chapter II) dimunity. Curravailability for the benthic commhey also between coral and surrounding water and increase gas and nutrient exchange. Tterial necessary for photosynthesis and active aprovide the supply of inorganic and organic more ey are furthermfeeding, and disperse released metabolites (Chapter I, II, III). Thpounds, which ents and associated comresponsible for the resuspension and dispersal of sedimturbid the water column and can suffocate bottom dwellers when resettling. Additionally,
of breakage of coral skeletons. fierce currents or waves hold the risk

l (Chapter I to V) as well as anthropogenic Many of the local influences can be of naturahereas the anthropogenic influence decreases with distance to (Chapter VI, VII) origin. Wperature rise or ocean shore (with the exception of global changes such as sea surface temconditions. According to type, e strongly influenced by oceanic acidification), offshore reefs ar erbance, a reef and its inhabitants will bd duration of the distuintensity, frequency animimportant role in near-shore (Cpacted to a stronger or lesser extent. Factors like seasonality hapter I, VI) as well as in offshore reefs (Chapter I to Vor tides can herein play an ).

Offshore reefs are strongly influenced by open ocean conditions (Chapter I to V), which can
be highly variable. 60 km off the Thailand west coast, the coral reefs along the west sides ofthe Similan Islands are affected by Large Amplitude Internal Waves (LAIW). Contrary, reefs
on the east sides are only impacted to a lower extend. LAIW frequently introduce cold,
pacts the the reefs which strongly im- and nutrient-rich deep water into oxygen depleted, CO2 of corals. etabolismm

ySummar

LAIW impact can strongly vary between same seasons of different years, and dependent on

ely high (Chapter 1). As ae extremthe intensity and frequency, the nutrient input can becom

concentrations increase, while conversely tenresponse, zooxanthellae densities and pigm

photosynthesis decreases (Chapter I, III, IV). Strong LAIW generated currents increase the

tter and support active feeding of corals (Chapter aflux of dissolved and particulate organic m

II). At the same time the high availability of organic and inorganic nutrients promotes an

ucus) (Chapter increase in release of dissolved and particulate substances by the coral (e.g. m

III). The combination of high zooxanthellae densities and active feeding on the west side

ass and energy reserves in corals (Chapter I, II). These features leads to an increase in biom

e conditioival in extremincrease the corals’ resilience and enhance survns (Chapter II). The

ore er, m are increased respiration and lowcosts of elevated biomass and energy reserves

pacted by lowered aragonite III). Calcification as well is imhapter costly photosynthesis (C

Csaturation state due upwelled water low in pH (ore, low temperature hapter III). Furtherm

and oxygen concentrations associated with LAIW are responsible for slower coral growth and

decreased photosynthesis. Additionally to the SW monsoon, which mainly influences the

shallow water reefs on the west side of the islands, the diverse LAIW impacts lead to reduced

gher biodiversity. The adaptive ation (Chapter V) with lower coral cover but hireef form

mechanisms vary between different coral species (Chapter I). Incubations revealed, that

conditions (Chapter III), before adaptation er non-LAIWr persists undatization behavioacclim

ent, Chapter II). Neverthto different conditions takes place (transplantation experimeless, the

pacted reefs is tential of corals from LAIW imatization poains, if the high acclimquestion rem

atization atization or a genetic adaptation. These studies on acclimtabolic acclimea m

reefs might develop in the face of global s serve as a basis for understanding, howechanismm

se and ocean acidification). perature rige (temate chanclim

ents at the Caribbean coast ofInput of nutrients and sedim Costa Rica (Chapter VI) are

ith rising rainfall nutrients are washed out of t seasons. Wstrongly dependent on dry and we

strongly fertilized soils from farmed land into near coastal waters. They are distributed by

currents to the coral reefs of the National Park Cahuita, where they are verifiable taken up by

onocultures near the coast corals. In addition, the extensive land clearing and cultivation of m

leads to erosion indicated by high silicate input. In consequence, high amounts of particulate

matter are transported by the rivers and currents into the reefs leading to low light conditions

and smothering of corals. The investigated reefs are highly impacted by eutrophication

ilar geographic position. pared to reefs with simcom

ySummar

ent are the consequences of coastal developmAlso along the northeast coast of Hainan, China,

entation by untreated visible and devastating (Chapter VII). Strong eutrophication and sedim

st coastal waters. ipopulations fertilize and m aquaculture and growing coastal sewage from

Excessive aquaculture holds the danger for input of diseases, alien species and pesticides.

ite fishing, thods, i.e. dyname is the obvious use of destructive fishing mAnother big problem

which destroyed big parts of the reef already. The consequently loose substratum (coral

hing and bination of overfis coment. Therubble) is not suitable for coral larvae settlem

increased nutrient input threatens the reefs as algal growth is no longer controlled by

le trients and growth on instabpetitors for nus fish. As algae are better comherbivorou

inated reef. Survivingsubstrate, there is the risk of imminent phase shift towards an algae dom

ent comcorals have a low photosynthetic rate in the highly turbid environmparable with that

of corals from deeper water. Additionally, strong sedimentation leads to a high energy

demand for coral mucus production crucial for sediment removal.

and an ated reefs of Costa Rica, the investigral cover and biodiversity inA decrease in live co

initiating phase shift from a coral to an algae dominated reefs in Hainan indicate the serious

disturbance to these ecosystems and increasing anthropogenic pressure will inevitably lead to

their collapse. In contrast to the natural occurring pulsed disturbances by LAIW, the chronic

creasing and therefore an acute risk.ntly inaneland based disturbances are perm

Table of Contents

Table of Contents

1 Introduction ...………………………………………………………………………………

15 Publication Outline ………………………………………………………………………...

ns e PublicatioInvestigativ

I Chapter

Metabolic plasticity of Porites lutea and Diploastrea heliopora exposed to

Large Amplitude Internal Waves …………………………………………………………. 17

II Chapter

Trophic response of corals to Large Amplitude Internal Waves ………………………………. 58

III Chapter

Comparative metabolic performance of Porites lutea from

AmLargeIV Chapter

Internal plitude ve Wapos(LAIW)-exAIW-protected habitatsed and L90 ………………..

Benthic primary production in response to Large Amplitude Internal Waves (LAIW) in

coral reefs at the Similan Islands, Thailand …………………………………………………106

V Chapter

ical characteristics of the Similan Islands physico-chemunity and Coral comm

in response to Large Amplitude Internal Waves ……………………………………………. 133

VI Chapter

Riverine input of particulate material and inorganic nutrients to a coastal reef ecosystem

at the Caribbean coast of Costa Rica ……………………………………………………… 167

VII Chapter

rvatiField obse aonsninary d prelimon notes ethe mNE-Hatabolic status of oinan crals178 …………..

195 Discussion ………………………………………………………………………………….

Introduction

Introduction

iosis and metabolism Coral symbprise the largest order of the als that comarine animian corals are entirely mScleractine species are free-living and solitary, som cnidaria). Even though anthozoa class (phylum

most of themall polyps (Ruppert and Barnes 1994). They are sessile and colonial with smework by producing a tion of the reef framatypic, contributing to construcare hermacher and Zibrowius 1985). In contrast to their cold- carbonate skeleton (Schuhmcalciumostly occur to water counterparts (Roberts et al. 2006), tropical shallow water species mbiosis with unicellular dinoflagellate algae utual symbe zooxanthellate, living in m

zooxanthellae are provided with shelter hile the acher and Zibrowius 1985). W(Schuhmal (Muller-Parker and D'Elia 1997), up to 50 % of and respiratory products of the animtheir photosynthates are allocated to the host (Muscatine and Cernichiari 1969),

supplying more than 100 % of its metabolic demands (Muscatine et al. 1981). Next to
being phototrophic, these tropical shallow-water zooxanthellate stony corals (hereafter referred to as ‘corals’) are as well effective planktivores and suspension feeders (Glynn
ore nthony 2000) who can also derive m1973; Sorokin 1973; Ferrier-Pagès et al. 1998; Aands (Palardy et al. 2008) by active feeding using their tabolic demethan 100 % of their matocysts (Lewis and Price 1975). The obedience on photo- tentacles or stinging nemversus autotrophy is species dependent (Wellington 1982; Palardy et al. 2005; Grottoli et

al. 2006; Rodrigues and Grottoli 2006), but also regulated by the prevailing

ental conditions (Lewis 1976; Muscatine et al. 1989; Anthony and Fabricius environmare strongly reliant on 2000; Palardy et al. 2006). By virtue of their sessile nature, corals ediate surroundings. their imm

Benthic-pelagic coupling ity in low-productive Coral reefs have been described to be oases of extensive productiv 1955). The tight and efficient cycling of nutrients and Odumocean surroundings (Odumith 1984; Hatcher 1990,1997; Lesser 2004) has been s (Smwithin these reef ecosystem of highly used to explain the nutrient paradox (Darwin 1842) denouncing the antagonism level, the ents. On the organismronms in nutrient poor enviproductive ecosystem

biosis between coral host and zooxanthellae guarantees the proficient utilization of sym

1

Introduction

al, their break-down to inorganic nutrients and thephotosynthates by the coral animbiotic algae (Muller-Parker and D'Elia 1997). In the latter by the symrecycling ofterial (Anthony 2000) are aant 1997) and organic maddition, inorganic nutrients (Szmdrawn from the water column and included into the holobiont metabolism. Portions of the
mmetabolites can be released into thatter (DOM or POM respectively) (Tanaka et al. 2008). The POM, or coral me water column as dissolved or particulate orgucus, can anic
trap further organic suspended material which settles and is then available to the
microbial loop (Wild et al. 2004). The reef sediments hence act as a filter system where
ild et al. ounts of heterotrophic bacteria (Wterial is decomposed by high amaorganic m2004) and where the regenerated nutrients are released back into the water column via resuspension due to currents or waves (Grant and Madsen 1979). Also the reefework functions as a sink for organic material (Richter and Wunsch 2001). Its mfraenormous labyrinth of cracks and crevices is densely populated by filter feeding
and Richter 1998) which intensely graze unschs such as cryptic sponges (Worganismterial (Van Duyl and Gast 2001) and aupon dissolved and particulate organic m releasing inorganic nutrients (Richter and subsequently fertilize the surrounding water by sny other reef-dwelling organismaWunsch 2001). On the further community level, mulation and such as algae (Larned 1998) and fish (Parrish 1989) contribute to the accumcycling of metabolites and minerals. While for example algae are the corals’ strongest
competitors for space and light due to their high susceptibility of nutrient assimilation,
their abundance is baladependency between the various reef compartmnced by herbivorous fish (Lapointe 1997). The correlation and ents can be high and the loss or
ght weaken the ecosystem (Hughes 1994). As coral ient of essential parts minishmdiment constantly in exchange s but their surrounding environmreefs are no closed system outsideents, they are sensitive and vulnerable to disturbance fromt environmwith adjacenith 1999). eier and Sm(Buddem

Environmental conditions the global distribution of tropical shallowjor physico-chemical factors regulating aThe mwater coral reefs are temperature, salinity, aragonite saturation state and light (Kleypas et al. 1999). The optimreef growth is determined to be around um average temperature for

2

Introduction

28°C (Coles and Fadlallah 1991), salinity should range from 34 to 35 PSU (Coles and Jokiel 1992), the aragonite saturation state has its optim-2um-1 at 3.8 (Takahashi et al. 1980;
(Chalker 1981; Achituv and sArcher 1996) and light at an average of 300 μE m bination of these features is given within the tropical belt fromDubinsky 1990). The comately 30°N to 30°S (Barnes 1987), however not everywhere reefs are present as approximthe ultimate constraint for reef formation is determined by the prevailing local factors
such as river runoff and precipitation, or waves and storm(Kleypas et al. 1999). These can be of a wide variety including hydrodynams (Macintyre and Adey 1990), ic factors
ant 1997,2002) and ical conditions such as nutrient concentrations (Szmbiogeochemnthony 2000) or biological factors such as larval terial suspension (Aaparticulate msupply, geographic isolation, diversity or disease (Cortés 1997; Kleypas et al. 1999; eters like . Many of these paramBruno et al. 2003; Nozawa and Harrison 2008) is tential of COperature and aragonite saturation state correlate as the dissolution potemdirectly impacted by water temperature (Levitus 1994). Other feedbacks include the 2
2006) which in turn can bereduced light availability due to high particle loads (Anthonypacts on increased via waves or currents (Grant and Madsen 1979). The environmental imcorals are multifaceted and directly as well as indirectly affect the coral host and its algae
(Brown 1997). ski asker et al. 1984) and elevated irradiance (FalkowWhile high seawater temperatures (Les within the coral tissue, and Dubinsky 1981) lead to a decrease in zooxanthellae densitinutrient availability increases symbiont propagation and pigment concentrations (Szmant
portant nutritional source for corals terial represents an ima1997). Suspended organic m and reduce light (Anthony 2000; Palardy et al. 2006), but can increase turbidityounts (Anthony and Fabricius 2000). Currents amor photosynthesis in higheravailability fdecrease the coral’s boundary layer and enhance gas exchange as well as nutrient and organic mstir up bottomatter uptake (S material, freeing organics and nutrients into the water columebens et al. 1997; Hearn et al. 2001; Sebens et al. 2003). They n, but also
ents which in turn increase turbidity (Grant and Madsen 1979).resuspending sedimost productively thrive in stable Above the previous knowledge that coral reefs can menvironmalso to be found in variabents (Crossland et al. 1991; Hatchele environments (Cortés 199r 1997), there are ma3; Dollar and Tribble 1993; ny examples of reefs

3

Introduction

Macyntire et al. 1993; Kleypas et al. 1999; Hughes et al. 2003; Fabricius 2005; Anthony entations of reefs (Kleypas et al. 1999) experiencing short-term2006). There are documperatures of as low as 18 (northern Persian Gulf) and as high as 34°C (near Bahrain, tema) or above 40 es dropping below 25 (off BurmPersian Gulf), or with salinities sometimU. Reefs in the vicinity of Galapagos, exposed to upwelling of water rich in S(Red Sea) PCO2 (Levitus 1994) persist in an environment with temporary aragonite saturation states
of as low as 3.1 (Macyntire et al. 1993; Kleypas et al. 1999). In higher latitudes, e.g. in of sunlight is low during winter and light Japan or Florida, where the incident angle penetrates only to depths of less then ten meters with lowest intensities of 50 μE m-2s-1
ore, reef growth has urthermreefs are still found (Harriott et al. 1995; Kan et al. 1995). F ant 2002) or turbid (Bak and Meesters 2000)ented in highly eutrophied (Szmbeen documwaters.arginal its are defined as ‘mvorable liments near or even beyond faReefs in environmreefs’ (Kleypas et al. 1999). The capability of corals to persist in such disturbed as et al.pact (Kleypents is dependent on the duration and the severity of the imenvironm potential of its atizationof the coral reef is set by the acclim1999) and the welfare unds 1999). The disturbances ates and Edminhabitants to the prevailing conditions (Gl as well as of anthropogenic ent can be of naturarginal reef environmacharacterizing a morigin (Grigg and Dollar 1990).

ternal Waves Natural disturbances on coral reefs – Large Amplitude InInternal waves (IW) travel along the interface of a two-layered fluid (Fig. 1), with their
maximum amplitude at this interface and almost no displacement at the top or bottom of
n occur wherever strong tidal currents, caWn (Jackson 2004). As Ithe water columphy co-occur, they are lar underwater topogran and irregustratification of the water columchanism of eubiquitous in the world’s oceans (Jackson 2004) and represent a significant mmass and energy transport (Osborne and Burch 1980). Not all mechanisms are yet
ented when barotropic tidal currents n has been documunderstood, however, their creatio depression on the of a stratified fluid overflow a barrier and so generate a downstream after turning or abating of the Weam as Iobstacle’s leeward side which is released upstrtide (Maxworthy 1979). Another formation mechanism is their direct production by

4

Introduction

shear-flow instabilities when strong currents overflow a sill creating upstream IW just
occur in wave packets, with the fastest i 1999). IWd Armer anafter the sill crest (Farmwave of highest amplitude and wavelength travelling up front and can be detected as
sharp drops in pycnocline depth (Jackson 2004). Their dissipation takes place via interactions (Jackson 2004). spreading, pycnocline instability, turbulence or bottomve trains over a disintegrate into secondary wDuring the latter process (Fig. 1), IW(Vlasenko and Hutter (Vlasenko and Stashchuk 2007) or internally brake shoaling bottomtified water bodies and hence play an xing of the previously strai2002), causing m ents and particulate larvae, plankton, nutriportant role in the cross-shore transport ofimmatter (Pineda 1991; Witman et al. 1993; Leichter et al. 1998).

bore formation on bore formation on shshoalioaling bottomng bottom

cold deep cold deep ersea watersea wat

Figure 1: Schematic drawing of an internal wave (modified after Osborne and Burch 1980) visible as
depression of the pycno/thermocline (black). Dissipation (gray) of an internal wave over shoaling bottom:
bore formation and subsequent mixing with overlaying water body.

5

The Andaman Sea features internal waves of extraordinary am

Introduction

plitudes of up to 80 m

plitude Intern(Osborne and Burch 1980). These Large Am) are aves (LAIWal W

an-Nicobar atra and the Andamgenerated by tidal flows over shallow ridges near Sum

island arc (Fig. 2) from where they travel eastward (Osborne and Burch 1980) until

dissipation as secondary waves or internal bores when they impinge the continental shelf

(Vlasenko and Stashchuk 2007).

topograFigure 2phy area: Map of thes of LAIW Andamforman Sea (mation (yellow) odified aanfd thter Jackse location on of the 2002) indicating the shSimilan Islands (red) close tallow undeorwater the
continental shelf (see depth contours).

In the vicinity of the LAIW swash zone close to the continental shelf break, lie the

ig. 2) consisting of nine granite islands ilan Islands, an offshore island chain (FSim

surrounded by coral reefs (Chansang et al. 1999). The arriving LAIx iW turbulences m

low-pH waters into the shallow reef docline cold, nutrient-rich, sub-oxic ansubtherm

areas evident as frequent drops in tem

perature, salinity, oxygen or pH (Fig. 3). Such

6

Introduction

perature decreases photosynthesis tem: lowdisturbances have known effects on corals

ation (Kleypas et al. 1999). Nutrients ines dense reef form(Saxby et al. 2003) and underm

ent growth (Szmfuel zooxanthellae and pigmant 1997), however also growth of algae

(Larned 1998), the main competitors for light and space (Lapointe 1997). Suboxic

s and low pH or aragonite saturation state conditions are deleterious to aerobic organism

decreases calcification (Fabry et al. 2008). So far the combined effects of LAIW impact

s for reduced and only scattered coral are unknown and the driveron coral organism

persistence along the LAIW exposed western Similan Island sides in contrast to dense

ations in the sheltered east (Chansang et al. 1999) not identified.reef form

Figure 3: Example of a time series of LAIW impact recorded in the water of west Koh Miang (Similan
Island #4). Temperature drops indicate passage of LAIW. Co-occurring are increases in salinity, as well as
.). pH (Richter unpubldecreases in oxygen and

ances on coral reefs Anthropogenic disturb

Next to the risks of global climate change, namely the rise of sea surface temperature as a

issions and the subsequent ocean acidification response to extensive greenhouse gas em

due to rising atmospheric CO2 (Buddemeier and Smith 1999), local impacts are of great

s (Fabricius 2005) and have been identified to pose a far concern for coastal reef system

7

Introduction

more immediate threat to coral reefs (Brown and Ogden 1993). Due to high gradients and ith and ent (Smvariability, the coastal zone is already a highly disturbed environmBuddemeier 1992) and human alterations of the adjacent ecosystems or the reef itself
introduce additional often increase the frequency and severity of fluctuations or n of crop results disturbances. Extensive deforestation of coastal vegetation for cultivatioent loads into the in increased soil erosion and subsequent introduction of high sedimcoastal areas via rivers or groundwater (West and van Woesik 2001; Fabricius 2005). The
subsequent increased turbidity lowers photosynthesis and calcification (Brown 1997; Marubini et al. 2001), and sediment loads might cause coral suffocation (Weber et al.
acher 1977; Rogers 1990). Fertilization ofes too costly (Schuhm2006) if removal becomtal discharge into the surrounding water bodies eacreage causes nutrient and heavy menez 1992; Fabricius 2005). Increased nutrients cause phytoplankton án and Jim(Guzms resulting growth (van Duyl et al. 2002) which increases turbidity or causes algal bloomin anoxic condition for the underlying benthic community (Song et al. 2009). Also ilitytibbenthic algae growth is fuelled by nutrients (Larned 1998) and their better suscepto nutrients might outcompete corals (Lapointe 1997). Heavy metals can additionally i.e.
pact the fecundity of corals or cause bleaching (Harland and Brown 1989; Esslemont im2000). Rising coastal developmthe effects of high particle and nutrient loads. Aquaculture farent increases sewage disposal (ISRS 2004) and amming in mangroves and plifies
s: enhanced cts on coral reef ecosystemilar effealong the shallow tropical coast has simant 1997) and heavy nutrient inputs fertilize the coastal area causing eutrophication (Szmre Additionally, aquacultutals are released into the surrounding (Chou et al. 2002).emann 1984) or escaped species (Naylor et al. poses the risk of introducing disease (Sindermplications on the reef, 2005). The use of anchors or destructive fishing gear has direct imediately killing the hit ework immmas it rapidly destroys the calcium carbonate fras and leaving unstable fields of coral rubble behind (McManus et al. 1997). The organismvariety of human induced impacts makes it important to gain information on the steady-
ent regulations. anagemply proper mstate of reefs to im

8

Scope of this thesis

Introduction

atization potential on s to contribute to the understanding of coral acclimThis thesis aim

the organism as well as the community level by identifying triggers of natural (Chapters 1

to 5) as well as anthropogenic (Chapters 6 to 7) disturbances and investigating the stress

responses of individuals as well as the reef ecosystem

n are: investigatio

1)

2)

3)

4)

5)

6)

7)

. The key questions under

How do Large Amplitude Internal Waves (LAIW) impact coral metabolism on

ter I species level? Chapporal, spatial and the tem

How do LAIW alter the trophic state of corals and how do they contribute to their

ce? Chapter II resilien

What is the actual difference in metabolism between LAIW exposed vs LAIW

ditions? Chapter III ens under comparable conral specimsheltered co

Chapter IV pact?W imary production affected by LAIHow is the prim

How does the combination of the above findings shape LAIW exposed reefs?

Chapter V

poral anHow is the temd spatial impact of extensive altered land use on nearby

s? Chapter VI coral reef ecosystem

pacts of destructive fishing, aquaculture and coastal bined imHow do the com

development alter a coral reef ecosystem?

eters are outlined in Figure 4. ated paramThe correlations and feedbacks of investig

9

Introduction

Figure 4: Schematic drawing of benthic-pelagic coupling. Green arrows: positive feedbacks / food web: the
combination of light and nutrients is crucial for primary production. It increases plankton, algal and
zooxanthellae growth. Primary producers are the base of the food web; phytoplankton is consumed by
zooxplankton metabolic subswhictances areh serve exs aschanged heterotrophicbetwee food sn coral animoal and symurce for the coral animal. Wbiotic algae; while the ithin the coral holobiont, photosynthetic
products are passed on from the symbiont to the host, respiratory waste products are passed back to and
particulate recycled by theorganics (DOM a zooxannthellae. Thed POM respe coral holctively). Red aobiont is also able rrows: to releasenegative feedback / repressi and take up dissolved aon: increase in nd
TSM, either by high rates of primary production, heavy resuspension or introduction of suspended material
algae is edecreasenergy light penet costly. Blue ration and aarrowsv: influeailability. Comnce of water mpetition otion (i.e. currentfor food, light and space s): with ince.g. reasing cbetween coral aurrennts there d
is an increase in fluxes, mixing and resuspension of nutrients, TSM (plankton as well as inorganics), DOC
and POM. Currents also decrease the boundary layer over e.g. tissue surfaces of corals hence increasing the
exchange rates of gases or the uptake rates of nutrients.

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1779-1793 32: Physical Oceanography slope-shelf topography. Journal ofVlasenko V, Stashchuk N (2007) Three-dimensional shoaling of large-amplitude internal waves. Journal of
1029/2007JC004107 Geophysical Research 112:doi:10.Weber M, Lott C, Fabricius KE (2006) Sedimentation stress in a scleractinian coral exposed to terrestrial
and marine sediments with contrasting physical, organic and geochemical properties. Journal of
ology 336:18-32 ogy and Ecental Marine BiolExperimWellington GM (1982) An experimental analysis of the effects of light and zooplankton on coral zonation.
ogia 52:311-320 OecolWest K, van Woesik R (2001) Spatial and temporal variance of river discharge on Okinawa (Japan):
Inferring the temporal impact on adjacent coral reefs. Marine Pollution Bulletin 42:864-872
Wild C, Huettel M, Klueter A, Kremb S, Rasheed M, Jørgensen B (2004) Coral mucus functions as an
energy carrier and particle trap in the reef ecosystem. Nature 428:66-70
Witmsubtian JD, Leichter JJ, dal zone: InflueGenovese S, J., Bnce of internal waves. rooks DA Procee(1993) Pulsed dings of thphytoe National Acplankton supply ademy of Scieto the rocky nces
90:1686-1690
Wunsch M, Richter C (1998) The CaveCam - an endoscopic underwater videosystem for the exploration of
cryptic habitats. Marine Ecology Progress Series 169:277-282

14

Publication Outline

Chapter I

Publication Outline

idt GM, Phongsuwan N, Richter C (in revision) Metabolic , Jantzen C, SchmRoder Cplasticity of Porites lutea and Diploastrea heliopora exposed to Large Amplitude Internal
Waves. Coral Reefs

The basic idea of a prThe concept of this study was developed by C oject on internal waves in the AndamaRoder and C Jantzen. Sanm Sea was initiated by C Richter. pling and analyses
nuscript was done by C awere conducted by C Roder and C Jantzen. Elaboration of the m C Richter. Roder with input from

Chapter II

Roder C, Fillinger L,Trophic response of corals to Larg Jantzen C, Schme Amplitudidt GM, Khokiattiwong S, Richter C (accepted) e Internal Waves. Marine Ecology Progress
Series

The basic idea of a project on inThe concept of this study was developed by C Roder, with input fromternal waves in the Andaman Sea was initiated by C Richter. C Richter. Data
samJantzen and C Richter. The mpling and analyses was maainly conducted bnuscript was writty C Roder with support fromen by C Roder with input from L Fillinger, C C Jantzen, G
C Richter. idt andSchm

Chapter III

idt GM, Jantzen C, Khokiattiwong S, Richter C (in preparation) Comparative , SchmRoder Cmetabolic performance of Porites lutea from Large Amplitude Internal Wave (LAIW)-
exposed and LAIW-protected habitats. Journal of Experimental Marine Biology and Ecology

The basic idea of a project on internal waves in the Andaman Sea was initiated by C Richter.
from C Richter. SamThe concept of this study was developed by C Roder, GM Schmpling and analyses was conducted by C Roder, GM Schmidt and C Jantzen with inpuidt and C t
nuscript preparation was conducted by C Roder with aJantzen. Data evaluation and midt and C Richter. ents by GM Schmprovemim

15

Publication Outline

Chapter IV ild C, Khokiattiwong S, Richter C (in preparation) , WRoder Cidt GM, Jantzen C, SchmBenthic preefs at the Simrimary production in respilan Islands, Thailand. onse to Large Amplitude Internal Waves (LAIW) in coral
The basic idea of a project on inThe idea of this project was developed by C Jantzen with internal waves in the Andamanput from C W Sea was initiated by C Richter. ild and C Richter.
SchmThe maidt andnusc C Roder. ript was written by C Jantzen with improvements by C Wild, C Richter, GM

Chapter V itted) , Jantzen C, Khokiattiwong S, Richter C (submRoder Cidt GM, Phongsuwan N, SchmLarge AmplCoral community and physico-chemitude Internal Waves. Marine Ecology Progress Sical characteristics of the Simeries ilan Islands in response to
The basic idThe particular idea of this mea of a project on inaternal waves in the Andamnuscript was developed by GM Schman Sea was initiated by C Richter. idt, N Phongsuwan and C
Richter. Data samC Jantzen and C Richter. The mpling and analyses was conduanuscript was written by G Schmcted by N Phongsuwan, G Schmidt with support fromidt, C Roder, N
Phongsuwan, C Richter, C Roder and G Jantzen.

Chapter VI Roder C, Cortés J, Jiminorganic nutrients to a coastal reef ecosysteménez C, Lara R (2009) Riverine input of part at the Caribbean coast of Costa Rica.iculate ma Marinterial and e
Pollution Bulletin 58: 1937-1943 The basic idea of the stuand analyses were conducted by C Roder who also wrote the mdy was developed by C Roder, C Jimanuscript with iménez and R Lara. Data samprovempling ents
énez and R Lara. by J Cortés, C Jim

Chapter VII meRoder Ctabolic status of NE-Hainan corals. , Wu Z, Richter C (in preparation) Field observations and preliminary notes on the
The basic idea of a project on Hainan, China, was developed by C Richter and ZMT ple analyses was carried out by C colleagues. The concept of this study, fieldwork and samRichter.Roder with the help of Z Wu. The manuscript was written by C Roder with support from C

16

Cornelia Roder1, Carin Jantzen1, Gertraud M. Schmidt2, Niphon Phongsuwan3 &
2Claudio Richter

1T, Fahrenheitstrasse 6, 28359 Bremen, MLeibniz Center for Tropical Marine Ecology, ZGermany2Marine Research, Am Alten Hafen 26, 27568 Alfred Wegener Institute for Polar and Bremerhaven, Germany3oad, 83000 Phuket, Thailand Phuket Marine Biological Center, 51 Sakdidet R

Submitted to Coral Reefs

- Chapter I -

Internal Waves

Porites lutea

Diploastrea heliopora

Chapter I

17

exposed to Large Amplitude and Metabolic plasticity of

Abstract

Chapter I

(Milne-Edwards Porites luteatabolic plasticity of the two mounding coral species eThe mand Haime, 1860) and Diploastrea heliopora (Lamarck, 1816) was investigated under an
impact gradient of Large Amplitude Internal Waves (LAIW) along the west and east side
man Sea, Thailand. Nutrient concentrations were highly ilan Islands, Andareefs of the Simbiont densities in both arked effects on sym intensity and resulted in mWregulated by LAIens. specimD. helioporauch stronger reflected in species, but the consequences were mZooxanthellae densities were increased more than threefold in P. lutea, but were more
than ten times higher in D. heliopora under strong LAIW influence. Also pigment
concentrations and protein content as well as host tissue and symbiont biomasses were
generally elevated in D. heliopora. The data suggest a highly species-specific response to
LAIW impact, where D. heliopora benefits stronger from increased nutrient and organic
matter availability than P. lutea which might explain their contrary abundances in LAIW-
are ubiquitous in South-ilan Islands. As LAIWexposed and sheltered reefs along the Simatization provides further l of acclimarkable potentiaEast Asia and beyond, the remporal and understanding on how corals cope with disturbances on small and large tem spatial scales in a changing world. s rdoKey wLarge Amplitude Internal Waves – coral metabolism – disturbance - acclimatization
Introduction

re than 200 oef areas for med to populate tropical reScleractinian corals are presummillion years (Stanley 2003), reflecting their ability to adapt to changing environmental
conditions (Veron 1995; Buddemeier and Smith 1999) on the temporal scale of minutes
to decades (Sebens and DeRiemer 1977; Hoegh-Guldberg and Smith 1989a; Done 1992;
unds 1999). Even though coral reefs Shashar et al. 1993; Brown 1997b; Gates and Edm

18

typically develop under fairly steady-state oligotrophic conditions (Lewis 1977;

Chapter I

l (Kinsey 1988; Woodley 1992; Cortés 1997) Muscatine and Porter 1977), large natura

insey 1988; Anthony and Fabricius 2000; Fabricius 2005) and anthropogenic (K

y ain corals which matization disturbances are known to occur, that require special acclim

be imh itmportant in the context of climate change (Brown 1997a; Buddemeier and S

1999; Hoegh-Guldberg 1999; Hughes et al. 2003; Grottoli et al. 2006). Studies about the

oodley 1992; Dollar and s (Winfluences of recurring events or disturbances such as storm

Tribble 1993), pollution (Heikoop et al. 2000b; Fabricius 2005; Anthony et al. 2009),

ing (Szmwarman 1990; Gleason 1993; Brown 1997a; Coles and Brown ant and Gassm

ntien 1982; Cortés 1997; Leichter and Genovese e2003), upwelling (Andrews and G

rt et al. 2005) on corals have greatly rown et al. 1999; Swa2006) or seasonality (B

improved the understanding of coral plasticity under varying environmental conditions.

son et al. 2002; However, often findings are species-specific (Brown et al. 1999; Stim

ent to be not coherent as zooxanthellae densities, pigmPalardy et al. 2005) or seem

ght not always uniformiconcentrations or growth rates me kind ly correspond to the sam

son 1997; Edinger et al. 2000; Sunagawa et al. 2008). Also most of disturbance (Stim

mechanisms of coral response to disturbance are not yet fully understood (Buddemeier

ith 1999; Fitt et al. 2000b; Oku et al. 2003), which requires further intense studies and Sm

tabolic functioning of scleractinian corals, their reaction to disturbances and eon the m

potential of acclimatization (Buddemeier and Smith 1999; Gates and Edmunds 1999;

Coles and Brown 2003).

ification (Crossland is obvious in coral calcpact of seasonality on coral life history The im

1984), displayed in skeleton banding patterns (Knutson et al. 1972), their reproductive

et al. 2005) and proliferous cycles (Babcock et al. 1994) or their nutritional (Palardy

(Kinsey 1977) characteristics. Annual or seasonal cycles have also been demonstrated to

ass (Fitt et al 2000), protein eters, such as biombe reflected in coral tissue param

(Crossland 1984) or lipid (Oku et al. 2003) content, zooxanthellae densities (Brown et al.

tt et al. ient concentrations (Brown et al. 1999; Fsditch et al. 2008) or pigm1999; Grim

2000b). Thereby, higher temperature (Lasker et al. 1984) and irradiance (Falkowski and

Dubinsky 1981) tend to decrease zooxanthellae densities or chlorophyll concentrations

Elia 1978; mmonium (Muscatine and D' a(Fitt et al. 2000b), while nutrient inputs of

19

Chapter I

Muscatine et al. 1989b), nitrate (Franzisket 1974) or phosphate (D'Elia 1977) enhance ably due to nutrient (mainly nitrogen) ent propagation presumbiont and pigmsymlimitation of the symbiotic algae (Muscatine et al. 1989b; Szmant et al. 1990; Muller-
nutrient enrichmParker et al. 1994; Fagoonee et al. 1999). Incrent have been brought into relation with augmeases in zooxanthellae densities due to enting tissue biomass
and Davies 1996) production. On (Muller-Parker et al. 1994; Fitt et al. 2000b), lipsmaller scales such as depth gradients, variances of id (Oku et al. 2003) or protein (Marubini
zooxanthellae densities (McCloskey and Muscatine 1984; Fitt et al. 2000b), chlorophyll concentrations (Dustan 1979; McCloskey and Muscatine 1984) or energetic status inly attributed to light a(Meesters et al. 2002) have also been described and were m2006) and subsequent increases in feeding differences (Al-Moghrabi et al. 1995; Anthony (Muscatine and Kaplan 1994; Palardy et al. 2005). However again, dependent on the species or region, the observed patterns were not consistent (McCloskey and Muscatine onships between coral host, its plex relati1984; Fitt et al. 2000b), suggesting more comsymInternal waves mbiont and the environmient. ght pose relevant impacts on coral reef ecosystems due to introduction
of nutrients (Wolanski and Delesalle 1995; Leichter et al. 2003), plankton (Witman et al.
1993; Leichter et al. 1998), la1997) when running on shoaling bottomrvae (Pineda 1999) or suspended m (Vlasenko and Stashchuk 2007) and aterial (Bogucki et al.
subsequently forming bores that cause extensive mixing of the deeper with warm nutrient
an Sea features internal waves of low surface waters (Jackson 2004). The Andam) are generated by plitude internal waves (LAIWe amension. These largexceptional diman-Nicobar the waxing and waning of the tides over shallow topography near the Andaman where they travel eastward in packets of 5-7 waves across the Andamisland arc, frombasin, depressing the thermocline by more than 80 meters (Osborne and Burch 1980).
Over shoaling bottom near the shelf break the LAIW are modified (Vlasenko and
(Vlasenko and Hutter 2002). to secondary waves or breakStashchuk 2007), disintegrate inIn the Indian Ocean, the mixing and upwelling associated with the dissipation of LAIW
mtemaperature (Coles and Fadlay generate sudden temperature drops of llah 1991; Saxby et al. 2003), but also the injection of >5 °C (Sheppard 2009). The variations in
nutrients (Wolanski and Delesalle 1995; Leichter et al. 2003), plankton (Witman et al.

20

Chapter I

a1993; Leichter et al. 1998), larvae (Pineda 1999) and suspended mterial (Bogucki et al. ary biota exposed to these waves. 1997) are likely to affect the sedent

Information on coral reefs subjected to seasonal or year round upwelling and/or internal

waves, focusing on nutrient (Leichter et al. 2003) or plankton (Leichter et al. 1998)

regim entes, biodiversity and coral cover (Cortés 1997), zooxanthellae and pigm th rates (Leichter and Genovese 2006) orconcentrations (Sunagawa et al. 2008), grow

induced variations Wfeeding (Palardy et al. 2005) is available. However, the LAI

investigated in this study by far exceed frequency and variation of earlier reported pulsed upwelling events (Leichter et al. 1996; Leichter et al. 2003; Lichter and Genovese 2006) e

ons so far reported in tropical reef areas fluctuatiand rank among the largest short-term

pacts reefs under such disturbances, and imom(Sheppard 2009). The plasticity of corals fr

ability are to date not examof possible inter-annual variined. This study presents

variations in biogeochemical water characteristics of reefs exposed to LAIW in greater or
lesser extends and discusses their impact on coral metabolic plasticity.

Material and Methods

SamStudy site pling took place in the Andaman Sea along the Similan Islands, Thailand, a

cated off the western Thai coast (Fig 1). Being lolongitudinal oriented island chain 60 km

close to the continental shelf edge, the islands are under influence of schoaling Large

Amplitude Internal Waves (LAIW) created near Sumatra and the Andaman Nicobar

Islands (Jackson 2004; Vlasenko and Alpers 2005). These bores force oceanic

sses upslope and introduce the nutrient rich, cold water into the asubpycnocline water mperature plunges of strong intensities and reefs where they can be tracked as tem

rather sheltered (Chansang et al. 1999). Comfrequencies on the windward, exposed island sidepas (Wred to m), while eastern island faces (E) are ost other tropical islands, where

coral reef growth is most intense on the windward side of islands and barriers (Veron and
ilan is rather SimStafford-Smith 2000; Spalding et al. 2001), coral cover on W

.). idt et al. subm(Schmulated disintegrated, while E side reefs are densely pop

21

Chapter I

Figure 1coast of Thailand (sm: Map of the Andamall map) and the 9 Siman Sea with the location of the Similan Islands with sampling sites on the Wilan Islands west of the and
E side reefs of Islands #2, 4, 7 and 8.

ling Coral samp

bruary to 16 March) and 2008 (23 February to 26 March), theeIn spring of 2007 (21 F

sborne and Burch 1980), fragmseasonal period of strongest soliton intensity (O ents of the

two scleractinian corals Porites lutea (Milne-Edwards and Haime, 1860)and Diploastrea

heliopora (Lamarck, 1816) were collected in shallow (5 - 15 m) and deep (15 - 25 m)

ilan Islands (Fig 1) using and E island faces of the Simareas of various reefs along the W

could (except for one D. helioporaSCUBA. Due to logistical restraints, in 2008

fragm that, a and E sides of Island #4. Apart froment) only be collected on the W

balanced sampling along all island

ounding ral species have mired. Both cos was asp

22

Chapter I

ilan Island reefs (pers. obers.). morpholgies and can grow to large boulders in the SimWhile P. lutea has polyps of ~ 1 mm in size, the polyps of D. heliopora are > 5 mm and
often up to 1.5 cm in diameter. Fragments of ~ 5 cm2 (one per colony) were chiseled
rts of the colonies, placed in ziplock bags (100 x 150 mm, the upper, non-shaded pafrommax. 4 ml residual water) and transported to the lab for processing. In total 48 fragments
of P. lutea and 35 fragments of D. heliopora were collected for analyses.

Coral processing Coral tissue was removed from the skeleton using an artist’s airbrush and filtered
ogenization (Ultra Turrax, 30 s) of the slurry, the solution was filled seawater. After homts were taken for zooxanthellae density which aliquoe of stock fromto a known volumined by six nthellae densities were determcounts, chlorophyll and protein analysis. Zooxacroscope (Leitz, ieter using a mocytomreplicate counts on a Fuchs-Rosenthal-haemPortugal, 260 x magnifications). Three to six mL of stock solution were collected under
n GF/F filters and frozen for chlorophyll analysis. Chlorophyll was am on Whatvacuumtrickland and Parsons 1972)and extracted by 90% acetone at 4°C for 24 hours (Sabsorbance read at 750 nm and 664 nm in a Shimadzu UV 1700 1nm Slit photometer and
concentrations calculated after Lorenzen (1967). Total protein content was measured
s established using bovine assay (Bio-Rad). A standard curve wusing the DC Protein Aserum albumin and absorbance read spectrophotometrically (Shimadzu UV 1700 1nm
Slit) at 750 nm (Lowry et al. 1951). The remaining stock solution of the homogenized
tissue slurry was centrifuged to separate symbionts from coral host tissue (Muscatine et
es beforeottoli et al. 2004). Zooxanthellate pellets were washed several timral. 1989a; Gresuspension in distilled water. Known volumes of each fraction were loaded on pre-ction (Millipore an GF/F) under moderate sumbusted and –weigehd filters (WhatcomVacuum Pump, max 200 mm Hg) and dried for 24 h at 40°C. Dry weight (DW) of the
animal and symbiont fraction was determined on a microbalance (Mettler, AT21
ents of ental and isotopic analysis. Measurem 1 μg accuracy) before elemparator,Comcarbon and nitrogen concentrations as well as their isotopic signatures on coral tissue and ntal Analyzer coupled with a eotein Elembionts were conducted with a NA2100 PrsymFlash 1112 Isotope Ratio Mass Analyzer. Carbon and nitrogen measurements were

23

Chapter I

ental CHNS standard (LECO). Isotopic ratios are given as calibrated against an elemconventional delta notations (13C and 15N) relative to Pee Dee Belemnite (13C)
standard and atmospheric nitrogen (15N). Fragment surface size was determined using

Simple Geometry to the nearest 0.05 mm (Naumann et al. 2009) and each parameter
2 coral surface. er cmlated pcalcu

Water samples In the same time period, water samples (n = 86) were collected in shallow (~ 7 m) and
ang using sterile 1 l PE bottles. Samples er of W and E Koh Mi reef wat)deep (~20 mwere transported to the laboratory and filtered (Millipore Vacuum Pump, 200 mm Hg)
busted and pre-ination) and pre-comdetermediately on untreated (for chlorophyll-a imm). TSM filters were hatman GF/Fination) glass fiber filters (Wweighed (for TSM determurs at 40°C), and dry weight of the total suspended esiccation (12 hokept frozen until dmatter (TSM) was determined gravimetrically (Mettler, AT21 Comparator, 1 μg
thawed filters by adding a certain volume of hlorophyll-a was extracted fromaccuracy). Ccubation at 4 °C for 24 rsons 1972) and subsequent in90 % acetone (Strickland and Paadzu orenzen 1967) in a Shim (L and 664 nmhours. Before reading absorption at 750 nmUV 1700 1nm Slit photometer, samples were centrifuged at maximum speed to remove
n. all particles in suspensioAliquots of the filtrate were poisoned with mercury chloride (Kattner 1999) and kept PE bottles until analysis of inorganic nutrients (nitrite, nitrate, lfrozen in 25 mammonium, phosphate and silicate) in a spectrometer (2007: GBC model UV/VIS981
ents, 2008: Evolution III autoanalyzer, Alliance Instrumodel FS3000; with autosampler mthods (Strickland and Parsons 1972; Grasshof 1983). More eFrance) using established mified with phosphoric acid to a pH of 2 poules, acidfiltrate was transferred to glass amalysis of dissolved organic carbon r until and storage in the freezebefore flame sealing anount DC-190) using a 10-point calibration with nalyzer (Rosem(DOC) in a DOC/DIC aTOC standards (ULTRA Scientific).

24

Chapter I

re records Temperatupled reefs (except for the perature was recorded in all sampling period, temDuring the sam activity) at two shallow site at Island # 7 in 2007, where logger was lost due to stormdepths (7 and 20 m) and in temporal resolution of 1 to 8 minutes depending on exposure
< 0.2 °C over 0 to 50 °C). To asses thee using TidbiT v2 loggers (Onset, accuracytimLAIW impact during coral sampling, the temperature of a 14-day period
e data was e fram(01.03.2007/2008 – 14.03.2007/2008) was utilized, as within this timfor all savailable fromites to com all sampare LAIWpling sites (F intensities. ig. 1). Daily temperature ranges ± SE were calculated
Statisticsdeep vs shallow, but also inter-island vs E, poral (2007 vs 2008) and spatial (WTemined using the sign test differences) variations of the daily temperature ranges were exametric data (Dixon and Mood 1946). for dependent non-paramterial aentary mples are given in the electronic supplemRaw data of water and coral samposition of the water as well as the total tissue (Tables S1, S2 and S3). The overall comcomside (W vsposition of either species investigated was com E) and depth (shallow vs deep) differences using PRIMER v6 software pared for seasonal (2007 vs 2008),
ed before analysis to achieve (Clarke and Gorley 2006). Data were log-transformality, stabilize variance and offset discrepancies due to the different unit sizes. normTo visualize constitutional similarities between the samples, multi-dimensional scaling
ples: nitrite,ensional (water same 8-dim(MDS) plots were generated which reduce thensional phate, silicate, DOC, chlorophyll-a and TSM) or 13-dim, phosnitrate, ammoniuma concentrations, protein content, e numbers, chlorophyll-ples: zooxanthella(coral samhost tissue and zooxanthellae dry weights, host tissue and zooxanthellae carbon and position of host tissue and zooxanthellae) nitrogen, carbon and nitrogen isotopic comsample coordinates into 2-D (axes arbitrary) space and where the (Euclidean) distance
e plot was used to stepwise visualize between the points denotes their similarity. The sameach hierarchical level on its own (first years, then sides within years, then depths within
sides) whichthe cases of significan allows for following a samt difference on lower hierarchical levels (btw sides and depths), theple to its detailed depiction (Figs. 5, 7 and 8). In

25

Chapter I

same graph, displaying the multidimensional plot from a different view onto the plane by
rotating to best illustrate the factorial differences, is additionally presented in the
Determelectronic supplemination of significant tementary material (Figs. S1 – S7). poral and spatial differences was accomplished
utations (Gonzalez and ilarities (ANOSIM) with 9999 permconducting Analyses of SimManly 1998) on each factor level (year, side, depth), always with the next hierarchically
lower level nested within (side nested within year and depth nested within side). For the coral data, differences within the same factor (year or side) but between the samples’
island of origin (# 2, 4, 7 or 8) were also tested with a nested approach (island nested within year or side). To identify the parameters (‘species’) accounting for the observed temporal and spatial
ER, ilarity (SIMPtions to simdifferences in water and coral data, a test on species contribusimilarity percentages) revealed the percental share of the parameters explaining  90 %
of the dissimilarities (Clarke 1993). SIMPER analyses were only conducted to the factor
(for water until dlevel on which the ANOSIM tests revealed at least one significant epth level, for coral data until side level); for the sake of com(p < 0.05) difference pleteness,
SIMPER results on these factorial level are all listed independent of significance,
etail. scussed in dihowever only significant ANOSIM results are d Resultsperature drops were observed, indicating the passage of Frequent, and often severe, temore frequent and severe in 2007 ). They were mplitude internal waves (LAIWlarge amperature ranges in 2007 (Fig. 2) than in 2008 (Fig. 3), as evidenced by stronger daily temperature ranges (± SE) and detailed ean daily tempared to 2008 (Fig. 4). The mcomstatistical results of the various sign tests are listed in Table 1. The temperatures varied
and E-exposed island faces (Table 1, Figs. 2, 3 and 4), but showed -coherently within Wpronounced differences (p < 0.001) between the years (2007 > 2008) and between the four sites (p < 0.001; W deep > W shallow > E deep > E shallow) for the given year (Table 1).

26

ng sidn asd Islannilame Sih tgn alo sites reefdateigsteve inh tmorfs (°C) egnre raeratupm tey: Dail
wlo. Beesulav-Z )wd (belo ansel): p levallowhs, deep/8, W/E7/200 (200riationsavporal tematio-pr sof < 0* ps: nce levelficaniackets. Sigr bnvel ince leificagn sidnvalue a 100. < 0 *** p,500. 0< ** p,50.
ble 1Ta slt resut tesgnide: si sth). Upper rigd (SE) ane (°Cre rangueratpm tey daileanal (bold): mon. Diagultst restes Zbers, mu ndan isltnreeffi dtnicafings (italics), siceerenffi dndr inter-isla fotsn tesig fts os (SE): resulmean

,*) (2.4195 (0.05)0087.35***6.33***7.35***3.88***hallowE7.35***7.0842vs #.s0#2 ************6.8286.2
55*31*5*08*eep vs ##4**)7,* (3.47
8)0. (036.1
***E*7.36.37.37.36.0***20*d

w.02.55.13.35W***894.0***82.6 (0.12)28.2
2008******shallo7660

1)940.8***16.0
3.47 (0.1*********3-0.108.7
).087 (00.09063.809.51.70
#7s#4 v*),**.47 (3)1,** (2.48vs ##4
2.2************57.335.77.086.33
9)**).94, (2 #7 vs2#)**47,*(3. #7 vs4#**).94, (2 #8 vs4#
80027200007272000072WEEWWdallowsheepdeepdwallosheep
3.70 (0.0).226 (04.3 (0.16).94 (2 #7s#4 v),**
30.6deepW7200wshalloW7200deepE7200wshalloE7200deepW8200wshalloW8200

deepE8200

wshalloE8200

Chapter I

27

7//307

720012/03/

3/13/07

3/13/07

Chapter I

30272421Island # 8
03/03/200703/12/2007
3027)24C°( 21Island # 7
er3/1/073/7/073/13/07
tu30are27pm24te21Island # 4
3/1/073/5/073/9/073/13/07
30272421Island # 2
3/1/073/5/073/9/073/13/07
Figure 2: Temperature profile within W and E side reefs of all Island sites (#2, 4, 7 and 8) along the
Similan Islands in 2007 ( W deep,  W shallow,  E deep,  E shallow)
30272421Island # 8
03/01/200803/07/200803/13/2008
3027)24C°( e21Island # 7
rtu3/1/083/7/083/13/08
30are27pm24te21Island # 4
3/1/083/7/083/13/08
30272421Island # 2
3/1/083/7/083/13/08
Figure 3: Temperature profile within W and E side reefs of all Island sites (#2, 4, 7 and 8) along the
Similan Islands in 2008 ( W deep,  W shallow,  E deep,  E shallow)

03/07/2008

7/3/80

807/3/

03/13/2008

3/13/08

3/13/08

28

Chapter I

Figure 4: Daily temperature ranges (14 peridays) od of all samof 2007 and 2008 pling sites along the Similan Islands during study

The nutrient characteristics of the reef waters (raw data Table S1) co-varied with LAIW

he strongest differences (p < 0.001, Table 2) coincided with the largest intensity. T

temetry varied 2007 and 2008 (Fig. 5). Nutrient stoichiomperature differences between

ceeded an order of trations exbetween years: Interannual differences in nutrient concen

magnitude for nitrate, contrasting the conservative behavior of silicate which mirrored the

two-fold temperature differences (Fig. 6). Nitrate and silicate contributed about 30 % and

ilarities between the strong (2007: 1.34 ± 0.26 μM nitrate Fig. 2; 6.12 25 % of the dissim

± 0.67 μM silicate, Fig. 6) and weak (2008: 0.09 ± 0.02 μM nitrate, Fig. 3; 2.59 ± 0.12

μM silicate, Fig. 6) LAIW years (Table 3). DOC (2007: 0.86 ± 0.05 μM, 2008: 1.89 ±

0.34 μM), TSM (2007: 10.51 ± 0.39 mg l-1, 2008: 15.61 ± 0.37 mg l-1), nitrite (2007: 0.09

M, 2008: 1.19 ± (2007: 0.55 ± 0.06 μonium± 0.01 μM, 2008: 0.60 ± 0.09 μM) and amm

able 3), all the years (Tilarity between0.07 μM) each add about further 10 % to the dissim

ore elevated during the year of less LAIW intensity (Table 3, Fig. 6). being m

29

Chapter I

Table 2: ANOSIM (Analysis of similarity) results after 9999 permutations of differences in water
composition between the years of 2007 vs 2008 (all), between W vs E reef sides within each year, and
between deep and shallow sites within each reef side. Each lower hierarchical factor is previously nested
within the higher (W and E sides within respective years and deep and shallow sites within respective side).
source of varianceRp
allyear0.569< 0.001*
side(year)0.0850.007*
2007side0.0970.024*
depth(side)0.2070.005*
2008side0.0770.055
depth(side)0.0520.062
2007 Wdepth0.3090.011*
2007 Edepth0.1420.035*
2008 Wdepth -0.0140.518
2008 Edepth0.0830.056

W vs E concentration differences were significant in 2007 (Table 2, Fig. 5 and S2), but
marginally not (p = 0.055) in 2008 (Table 2). In 2007, again nitrate (W: 1.83 ± 0.49 μM,
: 8.11 ± 1.29 μM, E: 4.50 ± 0.38 μM) concentrations E: 0.94 ± 0.22 μM) and silicate (Weach in the mean being more than 50 % higher on Wtogether explain about three quarters of the differences in water quality (Table 4) with compared to E (Table 4, Fig. 6).
Chlorophyll-a (W: 0.85 ± 0.08 μg l-1, E: 1.24 ± 0.09 μg l-1), ammonium (W: 0.63 ± 0.12
μM, E: 0.48 ± 0.03 μM) and TSM (W: 10.74 ± 0.61 mg l-1, E: 10.33 ± 0.50 mg l-1)
concentrations substitute each about 5 % W-E dissimilarity (Table 4), with chlorophyll-a
and TSM higher concentrated on being higher concentrated on the E side and ammonium and E are due to DOC, nor differences between Wi (Table 4, Fig. 6). In 2008, the mthe Wonium and TSM (Table 4, Fig. 6). nitrite, silicate, amm

Table 3: Annual comparison of reef water composition with all species contributing > 90 % to the
2008 between 2007 and dissimilarities
Ø abundanceØ abundancecontribution toSDcum. %
variable20072008Ø dissimilarityØ dissimilaritycontribution
μmol nitrate l-10.680.080.660.9928.38
μmol silicate l-11.831.260.600.8554.31
mg DOC l-10.610.940.300.8167.16
mg TSM l-12.422.800.200.1775.83
μmol nitrite l-10.090.430.200.3484.36
μmol ammonium l-10.420.770.190.2492.62
overall dissimilarity2.32

30

_________________________
2007 2008

___________________
2007: W E

___ _______ :2007 W deep  shallow 

___ _______2007 E: deep  shallow 

___________________
2008: W E

___ _______ :2008 W deep  shallow 

Chapter I

_______ ___2008 E: deep  shallow 

Figure 5: MDS ordination of all water samples collected during study periods 2007 and 2008 on W and E
of Koh Miang Island, top: years, middle: sides of 2007 (left) and 2008 (right), down (from left to right):
depths of 2007 W, 2007 E, 2008 W, 2008 E

31

Figure 6

Chapter I

: composition (means ± SE) of shallow (~ 7 m) and deep (~ 20 m) water of the W and E Similan reefs.: upper row: main LAIW

affected parameters (silicate, nitrate, phosphate), middle row: other N-derivates (nitrite, ammonium), lower row: TSM, chloroph

DOC

32

d yll-a an

Table 4: Side (W vs Econt) comributing > parison 90 % tof reeo thef diwater cssimomposition iilarities between n 2007 aW and nd E 2008 with all species

variable0720-1itrate lol nμm-1ate lol silicμmμg chlorophyll-a l-1
μmol ammonium l-1
-1 TSM lmgilarityerall dissimov

0820-1C l DOmg-1itrite lol nμm-1ate lol silicμmμmol ammonium l-1
-1 TSM lmgilarityerall dissimov

Ø abuWndanceØ abuEndancecoØ dntribissimilaution ritytoØ dSDissimilarityconcutribum. %tion

48.040.206.054.044.2

29.065.092.108.077.2

5.506.619.702.301.42

6.903.303.214.702.82

3.606.500.108.007.00591.

3.300.209.009.006.00810.

0.901.701.105.101.10

07.84.305.107.106.10

789.3435.7841.8836.8201.9

700.4974.6435.7935.82.994

Chapter I

ig. 5) were only h side of each year (Fater quality within eacDepth differences in wig. S3) of 2007, but again not in 2008 (Table 2). In F (Fig. S2) and E (significant for W2007, once more nitrate (deep: 2.95 ± 0.80 μM, shallow: 0.71 ± 0.25 μM) and silicate side are (deep: 11.14 ± 1.83 μM, shallow: 5.07 ± 1.20 μM) concentrations on the Wmdeep reef waters (Table 5), both being mainly responsible (each about 40 %) for the dissimore than twice as high in deep water (Fig. 6). ilarities between the shallow and
(deep: 0.64 ± 0.20 μM, shallow: 0.62 ± 0.13 μM) and phosphate Higher ammonium(deep: 0.48 ± 0.10 μM, shallow: 0.17 ± 0.05 μM) concentrations in the deeper water both (Table 5). Acontribute 5 % mlore to the dissimso on the E side in 2007, nitrate (deep: 1.19 ± 0.28 μM, shallow: 0.68 ± 0.33 ilarities between deep and shallow water on the W
ore than 50 % : 3.86 ± 0.55 μM) cause mμM) and silicate (deep: 5.13 ± 0.48 μM, shallow water (Table 5), each being elevated in of the discrepancies between deep and shallowdeep water (Fig. 6). Higher DOC (deep: 0.94 ± 0.14 m-1g l-1-1, shallow: 0.83 ± 0.10 mg l-1),
) and TSM (deep: , shallow: 1.13 ± 0.12 μg lchlorophyll-a (deep: 1.34 ± 0.13 μg l10.39 ± 0.58 mg l-1, shallow: 10.26 ± 0.86 mg l-1) concentrations in the deep water
contribute another 9 to 7 % to the dissimilarities between shallow and deep water on the
in differences between the deep and aE side reef in 2007 (Fig. 6, Table 5). In 2008, m

33

Chapter I

shallow waters of both, W and E, are posed by DOC concentrations, followed by nitrite, and E e all higher in deeper waters of Wns, which ar concentratiosilicate and ammoniumpared to shallow reef areas (Table 6, Fig. 6). com

Table 5: Depth (deep vs shallow) comparison of reef water composition in W and E reefs in 2007 with all
species contributing > 90 % to the dissimilarities between deep and shallow
Ø abundanceØ abundancecontribution toSDcum. %
variabledeepshallowØ dissimilarityØ dissimilaritycontribution
tsWeμmol nitrate l-11.210.480.971.0542.32
μmol silicate l-12.401.690.900.8981.53
μmol ammonium l-10.460.450.130.2187.38
μmol phosphate l-10.370.150.100.1091.88
overall dissimilarity2.29
Eastμmol nitrate l-10.690.410.440.4945.37
μmol silicate l-11.781.530.210.2366.72
mg DOC l-10.640.590.0910.1276.17
μg chlorophyll-a l-10.840.740.070.1083.34
mg TSM l-12.422.400.070.1090.47
overall dissimilarity0.97

0.690.410.440.4945.37
1.781.530.210.2366.72
0.640.590.0910.1276.17
0.840.740.070.1083.34
2.422.400.070.1090.47
97.0

Table 6: Depth (deep vs shallow) comparison of reef water composition in W and E reefs in 2008 with all
species contributing > 90 % to the dissimilarities between deep and shallow
Ø abundanceØ abundancecontribution toSDcum. %
variabledeepshallowØ dissimilarityØ dissimilaritycontribution
Westmg DOC l-11.030.790.451.2241.66
μmol nitrite l-10.640.470.240.3563.69
μmol silicate l-11.351.230.110.1774.21
μmol ammonium l-10.850.750.110.1984.19
mg TSM l-12.702.840.100.2493.59
1.08tyoverall dissimilariEastmg DOC l-11.110.800.250.4550.93
μmol nitrite l-10.370.280.070.0963.92
μmol silicate l-11.261.210.060.0775.61
μmol ammonium l-10.770.700.050.0786.11
μg chlorophyll-a l-10.450.410.030.0491.47
0.50tyoverall dissimilari

111.370.261.770.0.45

0.800.281.210.700.41

50.270.060.050.030.00.50

0.450.090.070.070.04

350.9263.9175.6186.1791.4

34

Chapter I

Differences in host tissue and zooxanthellae composition between the two investigated
igs. 9 and 10), particularly during the strong coral species are noticeable on first view (F season in 2007. Zooxanthellae densities, chlorophyll-a and protein content, host WLAItissue and zooxanthellae dry weights, as well as host tissue and zooxanthellae nitrogen teaP. lu (Fig. 10) than in D. helioporaand carbon concentrations are generally higher in of these differences prevail in 2008, but others (e.g. zooxanthellae emo(Fig. 9). Sporal (2007 vs 2008) and densities) don’t (Tables S2 and S3). For both species, the tem vs deep) dif vs E; shallowspatial (W were always much stronger (Table 7), than ferences the various islands’ e sites fromens sampled within the samong specimthe differences amreefs (Fig. 1) which totally lacked significant differences (Table 7).

Table 7: ANOSIM results, only main factor, each “beneath factor” separately nested
zooxanthellae comANOSIM (Analysis of simposition of ilaPorites lrity) resuteaults aft aner d Di9999 permploastrea helioporautations of di betfferences iween the yean coral rs of host2007 tissue avs n2008 d
(all), between W vs E reef sides within each year, and between deep and shallow sites within each reef side.
Each lower hierarcand deep and shallow sites wihical factor is previthin respectiously nested ve side). within thAlso island of origin is e higher (W and E sides withinested as a factn or withirespective years n the
hierarchical levels of year and reef side to test for significant differences among the same sides of the
. islandsousvari

ncevaria of sourceallyear
side(year)r)d(yealanisside2007de)pth(sideland(side)isesid2008de)pth(sideisland(side)heptd2007 Wheptd2007 Eheptd2008 Wheptd08 E20

Porites lutea
pR< 0.001*.1980< 0.001*0.222 -0.060.803
< 0.001*0.455 -0.040.641
0.2930.0550.0280.217
0.597 -0.0370.529 -0.0200.831 -0.1150.361.02200.871 -0.1460.1350.102

Diploastrea heliopora
pR< 0.001*0.7690.001*0.3021440.1760.0.029*0.8890.0090.4500.1840.1650.4540.001*
0.1050.1280.2310.2600.3000.0750.800 -0.2590.1000.4811970.077.0

35

_________________________
filled 2007 open 2008

___________________
2007: W E

___________________
2008: W E

Chapter I

____2007 W______ : ____2007 E: ______ ____2008 W______ : ____2008 E: ______
 deep  shallow  deep  shallow  deep  shallow  deep  shallow
Figure 7: MDS ordination of all parameters measured in the tissue samples from Porites lutea; top: years
(Island# 2 dia(left) and m2008 (riond, 4 downward ght), down tria(fromngle, left to 7 qua right): drate, 8 upwadepths rd looking of 2007 W, 2007 E, 2008 triangle), middle: W, sides 2008 E of 2007

36

_________________________
filled 2007 open 2008

___________________
2007: W E

___ _______ :2007 W deep  shallow 

___ _______2007 E:  deep  shallow

___________________
2008: W E

___ _______ :2008 W deep  shallow 

Chapter I

___ _______2008 E: deep  shallow 

Figure 8: MDS ordination of all parameters measured in the tissue samples from Diploastrea heliopora;
top: years (Island# 2 diamond, 4 downward triangle, 7 quadrate, 8 upward looking triangle), middle: sides
of 2007 (left) and 2008 (right), down (from left to right): depths of 2007 W, 2007 E, 2008 W, 2008 E

37

Figure 9:

tes luteaPoriChapter I

tissue composition (means ± SE) in shallow (~ 7 m) and deep (~ 20 m) water along the W and E Similan reefs.:

left column: holobiont parameters (zooxanthellae densities, chlorophyll-a concentrations and protein content); middle column: host tissue

parameters (dry weight, carbon, nitrogen content, isotopic carbon and nitrogen ratios); right column: zooxanthellae parameters

weight, carbon, nitrogen connt, isotopic carbon and nitrogen ratios) et

38

(dry

Figure 10:

ploastreDia helioporaChapter I

tissue composition (means ± SE) in shallow (~ 7 m) and deep (~ 20 m) water along the W and E

Similan reefs.: left column: holobiont parameters (zooxanthellae densities, chlorophyll-a concentrations and protein content);

column: host tissue parameters (dry weight, carbon, nitrogen content, isotopic carbon and nitrogen ratios); right column: zooxa

ratios)carbon and nitrogen eters (dry weight, carbon, nitrogen content, isotopic param

39

ddle im

nthellae

Chapter I

(Fig. 7) and P. luteaposition of st tissue and zooxanthellae comFirstly, differences in ho2007 and 2008 (Table 7, Figs. 7 and 8). re significant between e (Fig. 8) wD. helioporaFor both species, these yearly discrepancies are to 40 % due to higher zooxanthellae densities in 2007 compared to 2008 (Tables 8 and 9), with P. lutea stocking more than 3
times (2007: 4.30 ± 0.71106 cm-2, 2008: 1.23 ± 0.16106 cm-2) and D. heliopora even
more than 10 times (2007: 12.42 ± 1.14106 cm-2, 2008: 0.84 ± 0.16106 cm-2)
ntensity (Figs. 9 and 10). The yearly iWzooxanthellae in the year with strong LAI is further due to higher carbon (2007: 533.57 ± teaP. ludifference in the condition of 76.48 μg cm-2, 2008: 448.66 ± 51.80 μg cm-2) and nitrogen (2007: 47.06 ± 5.76 μg cm-2,
2008: 45.57 ± 5.90 μg cm-2) contents of the zooxanthellae (each composing 10 %-2
, 2008: milarity), and higher chlorophyll-a concentrations (2007: 8.31 ± 1.26 μg cdissim-2ilarity) in 2007. To a ) of the holobiont (also contributing 10 % dissim7.10 ± 1.28 μg cmminor extend (about 5 % each), dissim-2ilarities in P. lutea are addition-2ally due to a lower
(2007: 28.01 ± 1.98 μg cmhost carbon (2007: 268.50 ± 19.57 μg cm-2, 2008: 42.21 ± 4.71 μg cm-2, 2008: 419.92 ± 47.24 μg cm) content, but a slightly higher ) and nitrogen
total dry weight (2007: 5.63 ± 0.41 mg cm-2, 2008: 4.13 ± 0.60 mg cm-2) of the coral
tissue in 2007 (Table 8, Fig. 9). differences on the other hand are further determIn the holobiont composition of ined by strongly increased amD. helioporaounts of , yearly
chlorophyll in 2007 (37.94 ± 5.29 μg cm-2) compared to 2008 (4.71 ± 1.13 μg cm-2),
composing another 20 % of the yearly dissimilarities, followed by elevated-2
, 2008: milarity; 2007: 175.61 ± 26.24 μg czooxanthellate nitrogen (about 8 % dissim-2ilarity; 2007: 10.76 ± 1.16 ) and coral host dry weight (7 % dissimm78.56 ± 13.70 μg cmg cm-2, 2008: 4.77 ± 1.39 mg cm-2) in 2007. Due to high coral host nitrogen
2concentrations in the shallow W side specimens collected in 2008 (196.27 ± 45.87 μg cm-
; Fig. 10), yearly averaged coral host nitrogen concentrations were lower in 2007 (2007: 90.92 ± 9.59 μg cm-2, 2008: 98.96 ± 16.18 μg cm-2), while coral host carbon contents
-2-2) were higher in the year , 2008: 625.32 ± 108.28 μg cm(2007: 681.89 ± 68.26 μg cmof strong LAIW intensity, both substituting further 4 % dissimilarity (Table 9, Fig. 10).

40

Chapter I

Table 8: Annual comspecies contparison ributing > of coral host ti90 % tssuo the dissimilae and zooxanthellae comrities between 2007 and 2008 position of Porites lutea with all

Ø abundanceØ abundancecontribution toSDcum. %
variable20072008Ø dissimilarityØ dissimilaritycontribution
zooxanthellae cm-215.0013.73.34.5239.59
μg zooxanthellae carbon cm-26.045.911.041.6051.99
μg zooxanthellae nitrogen cm-23.693.640.090.1362.73
μg chlorophyll-a cm-22.041.870.851.1572.85
μg coral host carbon cm-25.545.890.540.6579.33
μg coral host nitrogen cm-23.323.620.510.6285.38
mg DW coral host cm-21.851.490.510.6691.38
8.40rityoverall dissimila

Table 9: Annual comparison with all species contributing of coral host tissu> 90 % to the e and zooxanthellae comdissimilarities between position 2007 of Dipland 2008 oastrea heliopora

Ø abundanceØ abundancecontribution toSDcum. %
variable20072008Ø dissimilarityØ dissimilaritycontribution
zooxanthellae cm-216.313.309.405.3143.71
μg chlorophyll-a cm-23.511.455.073.5567.30
μg zooxanthellae nitrogen cm-25.014.111.712.0975.26
mg DW coral host cm-22.401.441.561.5082.52
μg coral host nitrogen cm-24.404.530.931.3386.85
μg coral host carbon cm-26.466.170.831.1790.71
21.51overall dissimilarity

W and E side differences of the holobiont condition in 2007 (Fig. 2) were also significant (Fig. S5). In 2007, chlorophyll-a D. heliopora (Fig. S4) and P. lutea(Table 7) for W ilarities between concentrations in both coral species accounted for 20 % of the dissimre than twice as high on the oand E side corals (Tables 10 and 11 respectively), being mW side in P. lutea specimens (Fig. 9; W: 12.54 ± 1.84 μg cm-2, E: 4.44 ± 0.61 μg cm-2),
: were about 50 % higher on the E side (Fig. 10; WD. helioporawhile concentrations in 31.33 ± 5.51 μg cm-2, E: 48.95 ± 9.68 μg cm-2). Zooxanthellae densities (P. lutea: W:
5.59 cm-2 ± 0.13106 cm-2, E: 3.11 ± 0.06106 cm-2; D. heliopora: W: 13.12 ± 0.15106
cm-2, E: 11.25 ± 0.18106 cm-2), zooxanthellate carbon (P. lutea: W: 742.12 ± 133.86 μg
cm-2, E: 342.39 ± 23.42 μg cm-2; D. heliopora: W: 1188.15 ± 189.98 μg cm-2, E: 928.18
± 185.54 μg cm-2) and zooxanthellate nitrogen (P. lutea: W: 60.07 ± 10.45 μg cm-2, E:
35.14 ± 2.99 μg cm-2; D. heliopora: W: 176.43 ± 33.06 μg cm-2, E: 174.26 ± 48.02 μg

41

Chapter I

cm-2) further contributed 15 – 20 % (P. lutea) and 10 – 15 % (D. heliopora) to the
efs (Tables 10 and 11 respectively), all being and E side reilarities between the Wdissimhigher concentrated on the W island sides (Figs. 9 and 10). Being also elevated in W side
ilarity contribution just under 10 %; corals, the dry weight of the zooxanthellae (dissimW: 2.65± 0.51 mg cm-2, E: 1.14 ± 0.14 mg cm-2), as well as the coral host’s nitrogen (W:
33.08 ± 2.91 μg cm-2, E: 23.36 ± 1.95 μg cm-2) and carbon (W: 311.71 ± 31.26 μg cm-2,
E: 228.89 ± 18.87 μg cm-2) contents (each about 4 % dissimilarity contribution) further
had a minor share on the W vs E dissimilarities of P. lutea (Table 10, Fig. 9). In D.
heliopora, coral host carbon (W: 691.35 ± 52.95 μg cm-2, E: 666.11 ± 169.65 μg cm-2)
and nitrogen (W: 93.24 ± 9.02 μg cm-2, E: 87.04 ± 22.05 μg cm-2), and the dry weight of
zooxanthellae (W: 3.64 ± 0.65 mg cm-2, E: 3.09 ± 0.58 mg cm-2) as well as coral host (W:
10.84 ± 1.30 mg cm-2, E: 10.62 ± 2.39 mg cm-2) administered for seven to eight more
percent of the spatial W-E dissimilarities (Table 11), all again elevated within W side
ples (Fig. 10). coral sam Table 10: Side (W vs E) comparison of coral host tissue and zooxanthellae composition of Porites lutea in
2007 and 2008 with all species contributing > 90 % to the dissimilarities between W and E
Ø abundanceØ abundancecontribution toSDcum. %
variableWEØ dissimilarityØ dissimilaritycontribution
7200μg chlorophyll-a cm-22.511.61.231.4021.01
μg zooxanthellae carbon cm-26.305.811.220.8741.79
zooxanthellae cm-215.4014.801.071.5560.05
μg zooxanthellae nitrogen cm-23.833.550.960.8876.33
mg DW zooxanthellae cm-21.190.740.480.5184.47
μg coral host nitrogen cm-23.493.160.260.3588.97
μg coral host carbon cm-25.705.400.250.3293.17
5.86rall dissimilarityove8200μg chlorophyll-a cm-22.061.450.910.9624.32
μg zooxanthellae nitrogen cm-23.784.020.550.6038.85
μg zooxanthellae carbon cm-26.066.310.460.5351.14
mg DW coral host cm-21.591.950.410.4762.12
μg coral host carbon cm-26.146.300.340.3371.23
μg coral host nitrogen cm-23.883.970.330.3880-02
mg DW zooxanthellae cm-20.991.240.300.3887.88
mg protein content cm-20.6941.030.210.2293.44
3.75rall dissimilarityove

2.063.786.061.596.143.880.996940.

1.454.026.311.956.303.971.241.03

0.910.550.460.410.340.330.300.213.75

0.960.600.530.470.330.380.380.22

.3242.8583.1415.1226.231780-02.887893.44

42

Chapter I

Table 11: Side (W vs E) comparison of coral host tissue and zooxanthellae composition of Diploastrea
heliopora in 2007 and 2008 with all species contributing > 90 % to the dissimilarities between W and E
Ø abundanceØ abundancecontribution toSDcum. %
variableWEØ dissimilarityØ dissimilaritycontribution
0072μg chlorophyll-a cm-23.343.790.780.9819.68
μg zooxanthellae nitrogen cm-25.024.990.690.8137.16
μg zooxanthellae carbon cm-26.986.730.460.6048.77
zooxanthellae cm-216.316.200.380.5758.53
μg coral host carbon cm-26.516.360.340.3267.18
μg coral host nitrogen cm-24.504.360.320.3775.30
mg DW zooxanthellae cm-21.441.360.300.3482.86
mg DW coral host cm-22.422.350.290.3590.24
overall dissimilarity3.94
0082μg chlorophyll-a cm-22.401.022.161.2317.00
μg coral host nitrogen cm-24.963.991.842.1131.54
mg DW coral host cm-22.101.141.712.0444.99
zooxanthellae cm-213.9013.11.632.0457.82
μg coral host carbon cm-26.815.871.601.9570.39
μg zooxanthellae nitrogen cm-24.813.781.591.5182.95
μg zooxanthellae carbon cm-26.796.230.740.8088.76
mg protein content cm-21.440.910.640.7893.78
overall dissimilarity12.69

1.023.991.1413.15.873.786.230.91

2.161.841.711.631.601.590.740.6412.69

231.112.042.042.951.511.800.780.

17.0031.5444.9957.8270.3982.9588.7693.78

In 2008, when LAIW were weaker (Fig. 3), the condition of only D. heliopora (Fig. S6),
ens differed significantly between W and E side reefs specimP. luteabut not that of (Table 7). Again chlorophyll-a concentrations (W: 10.89 ± 1.79 μg cm-2, E: 1.86 ± 0.22
μg cm-2), this time more than four times as high on the W side reefs, were the strongest
attributor to spatial dissimilarity (more than 15 %) in D. heliopora (Table 11), followed,
: 153.34 ±ilarity contribution, by coral host nitrogen content (Wwith 15 to 5 % dissim29.24 μg cm-2, E: 73.87 ± 15.60 μg cm-2), zooxanthellae densities (W: 1.33 ± 0.39106
cm-2, E: 0.61 ± 0.13106 cm-2), coral host carbon (W: 1005.95 ± 227.03 μg cm-2, E:
449.64 ± 87.46 μg cm-2), zooxanthellate nitrogen (W: 127.79 ± 20.04 μg cm-2, E: 55.84 ±
14.07 μg cm-2) and carbon contents (W: 924.15 ± 127.11 μg cm-2, E: 621.33 ± 140.56 μg
cm-2) as well as total protein concentrations (W: 3.62 ± 0.89 mg cm-2, E: 1.73 ± 0.35 mg
-2cm), all of which being increased at about 50 side counterparts (Table 11, Fig. 10). Non-significant side dif% in Wferences in side corals compared to their E P. lutea samples
from 2008 were made of similar contributions of these parameters (Fig. 9).

43

Discussion

Chapter I

This study shows a marked effect of LAIW on the metabolic state of corals reflected in
biont density and t loads alter the sym nutrien The fluctuations intheir tissue. chlorophyll-a concentrations stronger than previously reported for seasonal variationsson 1997; Brown et al. 1999; Fagoonee et al. 1999; Fitt et al. 2000a; elsewhere (StimStimson et al. 2002; Grimsditch et al. 2008) and are in Diploastrea heliopora more than
an order of magnitude higher when the LAIW intensity is doubled. The combined effect
ith and Davies 1979; Hoegh-Guldberg and Sminsey of enhanced nutrient availability (Keased uptake with . 1999) and their incr1989b; Muscatine et al. 1989b; Fagoonee et alproduction and the consequences are stronger reflected in strong currents (Hearn et al. 2001) increases zooxanthellea density and enhances pigmD. heliopora compared toent
ght be the result of genotypic differences; nevertheless, phenotypic i. This mPorites luteareasons can not be excluded. Due to its skeletal structure, where the calices are shaped
like parabolic mirrors, D. heliopora might be a better collector of available light
indicated by lower (Enríquez et al. 2005) especially in shallow 13C values of the coral host tissue (Rodrigues and Grottoli 2006), water, plus a more efficient feeder as

especially in deeper water pointing to increased allochtonous contribution to nutrition
ncentrations indicative of heterotrophic (Muscatine et al. 1989a) and higher protein coinput (Bachar et al. 2007), as has been proposed for large-polyped corals (Porter 1976). Overall, our data suggest a strong fertilizing effect of LAIW, enhancing zooxanthellae ent concentrations (Brown et al. 1999). Along and pigmdensities (Muscatine et al. 1989b)with an increased uptake of organic material (Roder et al. accepted), this flux of
allochthonous material may explain the observed increased in biomass and protein
content in both species, similar to the findings of (Fitt et al. 2000b) and (Bachar et al.
2007), respectively. The 13C ratios of the zooxanthellae suggest decreased fractionation due to high turnover

strong :Wcarbon (DIC) pool in response to LAIrates of the internal dissolved inorganic LAIW intensity periods correspond with higher 13C ratios of the zooxanthellae which is

in line with decreased fractionation when the dissolved inorganic carbon pool (DIC)art et al. 2005) as a wes exhausted (Muscatine et al. 1989a; Sbecomresult of high DIC

44

Chapter I

hile the concentrations. Wentand due to high zooxanthellae densities and pigmdemincreased currents during LAIW may diminish the boundary layer surrounding the corals
(Dennison and Barnes 1988) and increasing gas exchange, the DIC demand increases in pigment densities) (Muscatthe course of dark adaptation and nutrient assimine et al. 1989a). The ilation (i.15e. increased zoN isotope data support the oxanthellae and
fractionation scenario for D. heliopora zooxanthellae (Muscatine and Kaplan 1994;
ple supply of nitrogen (Heikoop et al. 1998; McCutchan et al. 2003) in response to an amHeikoop et al. 2000a) during LAIW impact.Thehost tissue 15N ratios are as a result
W season and reflect the upward fractionation and also lower in the strong LAI one trophic level to ed for nitrogen frominsubsequent ratio shift of 1.4 to 3.3 ‰ determthe next (McCutchan et al. 2003). The data suggest species-specific responses to LAIW, where some species (e.g. D.
heliopora) benefit stronger from the replenishment of nutrients and organic matter by
LAIW than others (e.g. P. lutea). This is reflected by the occurrence of D. heliopora in
exposed areas (Veron and Stafford-Smexposed compared to sheltered E reefs along the Similan Islands. ith 2000) and its higher abundance in LAIWP. lutea on the other -
hand is mThe reefs along the Simore abundant in LAIWilan Islands experi-sheltered E reefs compared to Wence seasonal exposure to LAI (SchmWidt et al. subm.). impacts
e intensity. Internal waves (Osborne and Burch 1980), however not always of the samoccur where tidal currents advect stratified water across abrupt topography (Jackson and topography, the two-fold 2004). In the absence of interannual differences in tidesdifferences in LAIW impact observed between the two sampling years must be
pact was in water column stratification. Imferences interpreted as a consequence of difer parts of the W the shallow side reef waters. Mixing extended tostrongest in the deep Wperature side reefs, followed by the sheltered E deep and shallow reef sites. The temelling of nutrients: silicate, nitrate and phosphate e upwvariations coincided with thconcentrations in the LAIW affected areas were up to an order of magnitude higher than
wn et al. 1999).The pattern of increased nutrient introduction with areas (Broin other reefincreasing LAIW impact remain the same when focusing on differences between W and
e season (2007 and 2008 respectively), however thes within the samE side reefdifferences are more striking when the LAIW impact is higher as happened in 2007. On

45

Chapter I

parisons, the pattern was further observed forE side com-the scale of within-year Wterial indicating upwelling a, dissolved organic carbon or suspended mnitrite, ammoniumporal scale, when of these compounds. It was however overlapped on larger temcomparing strong (2007) and moderate (2008) influence of LAIW due to both faster
ounts which were turnover and rapid uptake or because of lesser upwelled am by stronger currents (Roder et al. the systemsubsequently faster discharged fromy not be the only source: e.g. an aaccepted.). Also, for these substances, upwelling mild et al. portant fraction of the dissolved organic carbon (van Duyl and Gast 2001; Wim2004) and ammonium (Muscatine and D'Elia 1978; Dickson and Wheeler 1995) is known
to be released by reef corapid uptake of phosphate and nitrogenous commmunities. Depth differences within the sampounds by primary producers under high-e reef are due to the
ine and D'Elia 1978; Capone and Carpenter iebe et al. 1975; Muscatlight conditions (W1982; Kokkinakis and Wheeler 1987; Larned 1998). The combined effects on coral metabolism ars strong and seem to be beneficial to some
work are less emextent, which appears at odds with the fact that reef growth and frapronounced on the Wwaters (Feely et al. 2008) m Simight limit coilan Island sides (Chansang et al. 1999). Upwelled, corrosive ral growth (Fine and Tchernov 2007) while
introduced nutrients might fuel algae (Larned 1998) increasing the competition for space
in corals (Lapointe 1997). Also storms often created during the SW-monsoon present in
the area between May and October (Wu and Zhang 1998) might pose mechanical
disturbance to corals (Dunne and Brown 1996) growing in the shallower waters especially on the W side reefs. Furthermore, there are few studies that demonstrated the
enous (Marubini and cts of phosphorus (Kinsey and Davies 1979) or nitrogharmful effeDavies 1996) compounds on corals, however in concentrations exceeding those reported
ilan Island reef sides of the Simin our study. Nevertheless, coral growing along the Wshow strong acclimatization potential to the natural disturbances of LAIW. For example,
ntioned to be a coral with high bleaching resilience (Harriott e has often been mP. luteainating highly impacted reefs (Cortés-Núñez en dom1985; Glynn 1990; Gleason 1993) oftand Risk 1985; Cortés 1993). So far, no data are available, to the best of our knowledge, on the metabolism of D. heliopora in spite of its potential in sclerochronology (Watanabe

46

Chapter I

; Corrège et al. 2004) and its widespread distribution in et al. 2003; Bagnato et al. 2004ith 2000). the Red Sea, Pacific and Indian Ocean (Veron and Stafford-Smith 2000) ber of coral species (> 600, Veron and Stafford-SmIn the light of the large numral triangle of South-East Asia (Jackson in the coand wide-spread occurrence of LAIWatization to a range of ential of acclim2004) this study suggests an extraordinary potLAIW-modified environmental régimes (temperatures, nutrients, currents, organic matter
to how corals cope with disturbances on portant insights inay provide imetc.) which mial scales in a changing world. poral and spatall and large temsm

ledgements: Acknow

Research was carried out in the framan bilateral ORCAS (Ocean-Reef e of the Thai-Germarch Foundation an Resean Sea) project funded by the GermCoupling in the Andam CT). The authors would like Thailand (NR(DFG) and the National Research Council ofilan to thank the Phuket Marine Biological Center and the National Park Mu Koh Simkiat Khokiattiwong and Kai-orothee Dasbach, Somstaff for field support, as well as Dchowski for laboratory assistance. Uwe Ludwi

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90:168Wolanski E, Delesalle B (19956-1690 ) Upwelling by internal waves, Tahiti, French Polynesia. Continental Shelf
Woodley JD (1992) Research 15:357-368 The incidence of hurricanes on the north coast of Jamaica since 1870: are the classic
Wu G, Zhang reef descY (1998) Tibetriptions atypical?an Plateau F Hydrobioologia rcing and th247:e Tim133-138 ing of the Monsoon Onset over South Asia and
the South China Sea. American Meteorological Society 126:913-927

52

Supplemental data Chapter I

deep and nde id E sinW a at theples collectedwater sameters measured in all params and (SE) of nae: m
2008 and 2007and in Koh Miang IslIsland waters of shallow reef
Table S1

)))))))).04.03.14.07.21.02.05.09000allow0000(0(((((((27955224sh.20.0.3.3.2.2.5.102000115E))))))5487)15)40106001........00000000(((deep(((((10872589.12.30.12.50.46.40.20.511

084)2)1)208)5)1)1)).1w0.00.20.11.10.00.00.00((((((((shallo0.532.461.140.646.431.200.260.071

27)(0.88)(0.04)(0.39)(1.42)(0.12)(0.03)(0.32)ep(0.ed.450.563.012.770.370.111.0914.201

)))))))02(0.12(0.55(0.02(0.86(0.1)(0.06(0.33low(0.la.13.86.53.05.26.83.25.68sh130000001

)))))))).13.48.0502.5814.09.28..000000(0(0((((((deep43409479.31.4.13.9.3.1..150000011

0720)))))))355)145310220101w........00010100((((((((hallo291768720.90.60.70.7s5.00.01.00.11WEW9)3)4)2)))))0.10.20.00.81.80.00.50.1((((((((deep460.780.862.951.140.170.410.48.011

l-1 eitrit nloμm1-itrate l nloμm1ium l-mon amlomμl-1phate hos ploμm1licate l- siloμml-1mg DOC l-1 ayll-phloroμg ch1M l-mg TS

Chapter I

53

01)22)37)55)14)97)29)36)45)87)77)23)8)(133.(142.low(12.(16.(0.(0.(1.(0.(0.(0.(0.(1.(0.shal1.232.844.466.165.637.001.8558.3965.94-14.34617.22-11.92633.77east(44.31)(10.51)(75.32)(0.35)(0.38)(2.01)(0.52)(0.36)(4.64)(0.56)(0.55)(0.58)(0.33)deep18779949354456097249895.843.691.1.6.7.6.2.1.30.46.276.479.-1-1

2008twescd E nW a llected at theo
Porites lutea stowshallpdeeowshalldeepshallowdeep53)(1.7.24(2.98)8.880.68)(4.17(1.07)4.721.9)(1212.16)(3.12.9009)(0.1.24(0.4)1.300.64)(2.53(0.93)3.700.8)(474.28)(2.6.5329)(0.1.06(0.27)0.850.1)(0.70(0.08)0.660.16)(171.15)(0.0.88(4.94(0.59)4.691.13)(906.58)(0.6.2246)(0.3.98(0.67)3.650.81)31)(51.470.51(66.62)361.4928.95)(234.15(26.78)223.6459.54)(12329.56)(33.297.2005)(8.49.40(7.74)36.132.24)(24.98(3.27)21.735.29)(5434.42)(3.31.8749)(0.-14.69(0.26)15.68-0.61)(-15.20(0.51)15.99-0.45)(18-15.4)(0.-16.7033)(0.6.95(0.62)6.980.24)(6.05(0.44)6.460.17)(515.19)(0.5.7321)(0.1.72(0.32)1.230.1)(0.85(0.21)1.421.02)(263.4)(0.2.1532)(58.441.83(74.64)307.4719.87)(296.87(34.48)387.91263.52)(33858.6)(123.645.2953)(7.44.96(7.8)31.483.43)(30.66(4.42)39.6117.65)(6662.82)(13.57.9118)(0.-12.24(1.26)14.34-1.04)(-12.65(0.62)11.79-0.83)(35-11.67)(0.-12.053)(0.4.29(0.28)5.080.3)(4.78(0.37)5.510.36)(455.58)(0.5.48
ea2007sides in deed 2008 nands in 2007 athe Similan Islong f waters ald shallow reenp a-2-2-2-2
eters measured in tissue samples of all params and (SE) of nae: m-2-2-2-2
west-2 chl-a cmgμ6ae zooxanthell10cmcmmg protein mg DW cm cmμg Cμg N cmcoral host13C15Nmg DW cmae cmμg Coxanthellμg N cm13Czo15N
Table S2

Chapter I

17)4)34)3)37)41)52)55)69)71)55)13)21)low(142.(257.(26.(25.(0.(0.(0.(0.(2.(0.(0.(0.(1.shal1.866.875.722.742.090.443.4568.1056.33-14.72425.82-13.07695.15east(105.81)(16.73)(11.43)(84.97)(0.23)(0.37)(0.96)(0.45)(1.13)(0.29)(0.35)(0.2)(0.4)deep97852859354305283019815.814.281.1.6.5.2.1.0.80.55.477.535.-1-1

93)20089)61)87)38)49)94)52)54)02)94)76)09)ow(249.(369.(39.(45.(1.(0.(3.(0.(0.(1.(0.(0.(5.shall5.001.034.382.956.8910.6016.22-15.07147.80-15.27196.271047.031329.07 twescollected at the(128.29)(56.91)(18.12)(0.39)(0.66)(0.55)(0.54)(1.55)(6.12)(0.28)(0.33)(0.49)(1.22)deep2.241.632008 4.241.735.983.5411.1817.5216.35107.77801.27110.42682.83--liopora in 2007 and a he312.64)13.41)98.03)355.7)41.37)1.53)0.89)0.31)0.86)1.03)0.16)0.58)4.14)ow(((((((((((((shall4.11nds 4.273.115.7011.1853.4312.12-13.40181.60866.01-15.47110.38770.13Diploastrestea Similan Islales of (204.32)(16.39)(43.18)(13.21)(1.09)(1.17)(0.78)(0.52)(0.53)(3.02)(0.4)(188)(4)pdeep2.743.703.075.869.1311.3244.4614.8617.0063.70166.91990.35562.10long the--2007ers aeters measured in tissue sam

watreef westshallow12.24)(2040.2.76)(2515.0.39)(773.2.59)(5113.40.12)(99758.12.43)(03101.0.23)(37-14.0.09)(635.0.9)(325.379.13)(281337.58.85)(22196.0.19)(17-12.0.31)(324.ae: meters measured in tissue samall params and (SE) of nd shallow np ain deedes d E sinW a-2-2-2-2
11)(214.deep69)(3.25.4264)(1.11.7035)(0.2.7489)(0.9.0517)(82.646.2699)(12.88.05-16.8638)(0.11)(0.5.8957)(0.2.525)(42.163.238)(0.-13.8254)(0.3.67
1088.74-2cmae -2-2-2-2 zooxanthell chl-a cmgμ610cmmg protein mg DW cm cmμg Cμg N cm13C15Nmg DW cm cmμg Cμg N cm13C15N
Table S3aeoxanthellzocoral host

54

Chapter I

Figure S1: 3D-MDS (stress: 0.07) plot of all parameters measured in reef water from 2007 W (filled) vs E
n) pe(o

Figure S3: 3D-MDS plot (stress: 0.06) of all parameters measured in reef water from 2007 W deep (filled)
(open) allow hvs s

55

Figure S3Figure S4Chapter I

: 3D-MDS plot (stress: 0.06) of all parameters measured in reef water from 2007 E deep (filled)

(open) allow hvs s

: 3D-MDS plot (stress: 0.03) all parameters measured in the tissue samples of

(open) E 2007 W (filled) vs

Porites lutea om fr

56

Chapter I

Figure S5: 3D-MDS plot (sthelioress: 0.06) of alpora from l parameters m2007 W (filled) vs easureE d in(open) the tissue samples of Diploastrea

Figure S6: 3D-MDS (stress: 0.03) plot of all parameters measured in the tissue samples of Diploastrea
heliopora 2008 W (filled) vs E (open)

57

1

- Chapter II -

Chapter II

Trophic response of corals to Large Amplitude

Internal Waves

Cornelia Roder1, Laura Fillinger1, Carin Jantzen1, Gertraud M. Schmidt2,Somkiat
23 & Claudio RichterttiwongKhokia

Khokiattiwong

& Claudio Richter

Leibniz Center for Tropical Marine Ecology, ZMT, Fahrenheitstrasse 6, 28359 Bremen,

Germany2 Alfred Wegener Institute for Polar and

Alfred Wegener Institute for Polar and Marine Research, Am Alten Hafen 26, 27568

Bremerhaven, Germany3Phuket Marine Biological Center, 51 Sakdidet R

Phuket Marine Biological Center, 51 Sakdidet Road, 83000 Phuket, Thailand

Accepted in Marine Ecology Progress Series

58

Abstract

Chapter II

The trophic response of the scleractinian coral Pocillopora meandrina (Dana 1846) to Large

Amplitude Internal Waves (LAIW) was investigated in the Andaman Sea. Corals living on
showed significantly Wand) exposed to LAIilan Islands (Thailthe west sides of the Sim

tered corals on the east sides. LAIW-exposed ass and protein content than shelhigher biomcorals were also more heterotrophic, displaying lower 13C ratios in their tissues and higher

red counterparts. Heterotrophic pared to shelterates of survival in artificial darkness com

-W photosynthesis leads to higher energy reserves in corals from LAInutrition in concert withexposed reefs making them more resilient to disturbance. As these differences in trophic

status are due to LAIW-enhanced fluxes of organic matter, LAIW may play an important role

in supporting coral monsoon beaten reefs. and survival in these mtabolisme

s rdoKey w

Large Amplitude Internal Waves – corals - heterotrophic plasticity – current regime –
– AndaPocillopora meandrinan Sea am

corals to LAIW: Trophic response ofRunning head

Introduction

The trophic response of corals to natural and anthropogenic stressors has been addressed in several studies (Anthony 2000, Anthony & Fabricius 2000, Palardy et al. 2005, Anthony 2006, Palardy et al. 2006, Rodrigues & Grottoli 2006, Borell et al. 2008, Palardy et al. 2008).

ly photoautotrophic, deriving the bulk of their inaAlthough most reef corals are functionally m

energy from photosynthesis (Franzisket 1969, Muscatine & Cernichiari 1969, Muscatine &

ents Porter 1977), heterotrophy can supply 11 - 46 % of the coral’s daily carbon requirem

(Houlbrèque & Ferrier-Pagès 2009), and more than 100 % in bleached corals (Palardy et al.
ature ile corals have been traditionally viewed as planktivores by virtue of their armh2008). W

onge 1930, Abe 1938, Lewis & Price 1975), their tocysts (Yamof tentacles and stinging ne

59

Chapter II

diet comprises a much wider range of food including microphytoplankton (Glynn 1973),

nano- and picoplankton (Ferrier-Pagès et al. 1998, Houlbrèque et al. 2004b), bacteria (Sorokin

tter (Sorokin 1973, Grover et al. 2008), detritus a al. 1998), dissolved organic mt1973, Bak e

ents (Anthony 2000). tter laden sedimaand organic m

, etabolismportance of photoauto- vs. heterotrophy in coral mTo detect and quantify the im

(Muscatine et al. 1989, Grottoli stable isotopes have been established as a useful indicator2002, Swart et al. 2005). The 13C ratio of the coral host tissue is the combined result of

ts from allochthonous sources al inpuphotosynthetically derived products and nutrition

(Muscatine et al. 1989). The ratio depends on the fractionation potential of the zooxanthellae,

the consequential isotopic signature of their translocates, and the signature of heterotrophic
13C ratio in corals is higher when photosynthetic rates are high and the internal carbon. The

carbon pool depleted (Muscatine et al. 1989). It decreases with light at increasing depths in

response to a decrease in photosynthesis and the increased proportionate heterotrophic uptake

of isotopically ligh and other oceanic particulate and dissolved organic ter zooplankton

materials (Muscatine et al. 1989, Grottoli 1999, Grottoli & Wellington 1999, Grottoli 2002).

ces skeletal (Grottoli & WAlong with photosynthesis, heterotrophy enhanllington 1999, e

(Ferrier-Pagès et al. 2003) que et al. 2004a) and tissue growthHoulbrèque et al. 2003, Houlbrè

by building up energy stores including lipids (Anthony 2006, Rodrigues et al. 2008, Treignier

et al. 2008) and proteins (Ferrier-Pagès et al. 2003, Houlbrèque et al. 2003, Houlbrèque et al.

2004a). Heterotrophy has been shown to support coral photosynthesis (Grottoli 2002, Ferrier-

Pagès et al. 2003, Houlbrèque et al. 2004a, Borell et al. 2008) and resilience to stress such as

nthony 2006), warmturbidity (Aing (Borell et al. 2008) and bleaching (Grottoli et al. 2006,

inant Palardy et al. 2008). Although active feeding does not generally constitute the dom

y reduce temporary energy deficits (Anthony acarbon source for zooxanthellate corals, it m

) so that corals with high capability to 2000, Anthony et al. 2009, Fitt et al. 2009

heterotrophically assimilate carbon may be more effective in surviving multiple bleaching

nant in future reefs (Grottoli et al. 2006). ie domevents and becom

The relative proportion of heterotrophy to coral metabolism may vary markedly between

ented in several species (Grottoli et al. 2006) and this heterotrophic plasticity has been docum

Pocilloporaobserved that the branching coral ple Wellington (1982) studies. For exam

damicornis grew independent of zooplankton supply and was more markedly affected by

shading than the massive coral Pavona clavus. Sebens & Johnson (1991) documented higher

capture rates with increasing current strength by Madracis decactis, but not by Meandrina
meandrites. Rodrigues and Grottoli (2006) showed that Montipora capitata host tissue 13C

60

Chapter II

Porites ic feeding, while increase in heterotrophratios decreased when bleached because of

did not alter its nutrition. And also Palardy et al. (2008) observed that the feeding compressa

y vary signiaresponse to one disturbance mficantly between different coral species.

The importance of heterotrophic feeding in coral metabolism may further vary between

ents (Lewis 1976, Palardy et al. 2005). Decreasing light and photosynthesis environm

(Muscatine et al. 1989, Palardy et al. 2008) have shown to increase coral feeding in deep

nts (Anthony 2000, e(Ferrier-Pagès et al. 1998, Palardy et al. 2005) and turbid environm

ulated by high concentrations of Anthony 2006). Coral feeding was also shown to be stim

tter (Houlbrèque et al. 2004b)adissolved organic m and zooplankton prey (Ferrier-Pagès et al.

1998, Ferrier-Pagès et al. 2003, Palardy et al. 2006), and to be influenced by prey behavior

(Palardy et al. 2005), coral feeding effort (Sebens et al. 1996, Palardy et al. 2005, Palardy et

al. 2008) and water currents (Lewis 1976, Sebens et al. 1996, Sebens et al. 1998).

Until now, internal waves as source of ambient variability in coastal currents, turbidity and

plankton (Pineda 1991, Leichter et al. 1996) are poorly investigated. These subsurface waves

are ubiquitous in the ocean (Jackson 2004) and propagate along the density interface

surface and cold and nutrient-rich deep waters, but their potential (pycnocline) between warm

man Sea features non-effect on the trophic state of corals is virtually unexplored. The Anda

linear internal waves of extraordinary amplitude, displacing the depth of the pycnocline by

borne & Bs (Perry & Schimke 1965, Omore than 80 murch 1980). Because these Large

Amplitude Internal Waves (LAIW) are tidally generated, travel over long distances, and

(Vlasenko & Stashchuk 2007), reefs disintegrate into wave trains over shoaling bottom

Wlocated in the swash area of LAIjected to frequent disturbances of the tentially sub are po

ent. So far, it is not known if and to what extent turbulent boluses ical environmphysico-chem

senko & Stashchuk 2007) advecting cold, nutrient rich (VlaWgenerated by shoaling LAI

waters upslope affect the trophic state of corals in LAIents. -environmW

ass, protein and stable isotope content ofHere, we combine observational data on the biom

-exposed and -sheltered corals with in situ light-exclusion and transplantation WLAI

ents, to explore the role of LAIexperimhe trophic state of corals in response to the on tW

combined effect of increased currents (Sebens et al. 1998, Nakamura et al. 2003), fluxes of

particulate matter (Anthony 2000) and plankton (Wellington 1982, Al-Moghrabi et al. 1995,

Ferrier-Pagès et al. 1998, Ferrier-Pagès et al. 2003), along with lack of photosynthesis

(Rodrigues & Grottoli 2006).

61

Materials and methods

Study site

Chapter II

off the west coast of Thailand consist of 9 granite islands ilan Islands located 60 kmThe Sim

(Fig 1). The west sides (W) of the islands feature barren rock and scattered corals, the east

sides (E) dense coral reefs (Chansang et al. 1999). The asymmetry in coral distribution

W corresponds to the western exposure of the islands in the swash zone of breaking LAI

an-Nicobar islands (Jackson 2004, Vlasenko & Alpers atra and the Andamgenerated near Sum

shelf break (Vlasenko 2005). Upslope propagating density intrusions emanate from the near

& Hutter 2002, Vlasenko & Stashchuk 2007) and arperature drops e evident as frequent tem

and overall lower mean temperatures on the LAIW-exposed W sides (LAIW+) of the islands

compared to the sheltered E sides (LAIW-) (Schmidt et al. subm.).

on Figure 1W (LAIW+: Map of the Sim) and E (LAIW-) ilan Islands in tof Islahnd e # Andam4 (Koh Miang an Sea Isoff the coland) are also locations ast of Thailand. Experimof water anental sites d near-ree(circles) f
plankton samfor the mupling. Off-reelti-dimensional analf plankton samysis were collected at apling sites (on Islall indicated reefs nd # 4) are indi(2W, 2E, cated with dia4W, 4E, m7W, onds. 7E, Sam8W, ples use8E). d

62

Chapter II

Coral sampling and experimental design (Dana, 1846), a common species on both Pocillopora meandrinaThe scleractinian coral LAIW+ and LAIW- reefs of the Similan Islands (Schmidt et al. subm.), was chosen as model
for the study. organismReef fragments - To detect differences between sides (LAIW+ vs LAIW-) and within sides
of the differcollected randoment islands (# 2, 4, 7 and 8), fragments of ly (1 to 12 fragments per site) between 5 and 25 m depth from various P. meandrina (one per colony) were
LAIW+ (18 fragmbetween 20.02.2008 and 24.03.2008. Fragments) and LAIW- (21 fragments) reefs of the Sients were placed in Ziploc bmilan Islands (Fig. 1) ags (100 x 150 mm,
max. 4 ml residual water) and transported to the laboratory for immediate processing.
ent (Fig 2A) was conducted from The light-exclusion experimLight-exclusion experiment - ilan Island # 4), three donor 20.02.2008 to 24.03.2008. On each side of Koh Miang (Simcolonies of P. meandrina were collected at a depth of 20 m. From each colony, 21 fragments
were clipped off and attached to twwire by clamping the base of the fragmo rails (control and experiment into cut out holes of the rail. Because branch ental rail) made out of plastic
pact on flow patterns within the coral colony and, hence, feeding capacity spacing has an imbearing rails were moored to a PVC-fram(Sebens et al. 1997), only single undivided branches were used in the experime and left in the reef to recover for one month. At ent. Fragment-
each donor colony) were ents (one fromfragment, triplicate start the onset of the experiments) placed in perspex flow pipes. A 3x2 bearing 10 fragmcollected and the rails (each nowounted on a rack equipped with a current vane which stack of these flow pipes (Fig 2A) was m ent (20 mallowed the set-up to rotate freely around an iron rod anchored into the sedimer) were te openings of the tubes (50 cm length, 10 cm diamdepth), so that the upstreamber set-up itself, as ag of the cham. Drbottomalways facing into the current, ~ 1 m above the easured in repeat runs (n = 6) with fluorescent dye, was not found to have a significant mbient water flow by < 5 %. The upper row of effect on water velocity, reducing the aments (photosynthesis +), the lower row of tubes were translucent tubes held the control fragments, shaded off with opaque foil (photosynthesis -), so that light levels near the fragmeasured with the light mmpensation light intensity eter of a Diving-PAM, were below the comfor photosynthesis (< 5 μmol quanta m-2 s-1). Sampling always took place before noon.
Triplicate samples from controls and light-deprived fragments (one fragment per tube) were
+) days after the onset of W-)/33(LAIWcollected 6, 8, 10, 12, 14, 16, 20, 24, 28 and 32(LAIthe experiments. Live fragments were placed into 100*150 mm Ziploc bags (max. 4 ml

63

Chapter II

supernatant), transported to the laboratory and immediately processed. Dead fragments were ed. recordTransplantation experiment - Within the same timeframe (20.02.2008 to 24.03.2008), a
anges in coral tissue ent (Fig 2B) was conducted to detect chcross-transplantation experimcomposition due to transplantation. On each, W (LAIW+) and E (LAIW-) Koh Miang, three
additional donor colonies of P. meandrina were sampled. Sampling and fragment cultivation
ent less on each rail). At ent (but one fragmn experimwas identical to the light-exclusioexperimental onset, triplicate start fragments (one from each donor colony) were collected
) cross-transplanted between W entsental rails (now each bearing 9 fragmand the experim(LAIW+) and E (LAIW-). Rails transplanted from LAIW- to LAIW+ were subsequently
the opposite direction exposed to higher flows of plankton, while rails transplanted in ents and cross-pared to origin conditions. Control fragmexperienced lower food supplies comtransplanted fragments were left anchored next to each other in the reef (20 m depth) and
triplicate samples of controls (feeding +) and feed-altered (transplanted from LAIW- to
LAIW+: feeding ++; transplanted from LAIW+ to LAIW-: feeding +-) fragments collected 2,
4, 8, 12, 16, 20, 24, 28, 32 (LAIW+) and 3, 5, 9, 13, 17, 21, 25, 29, 33 (LAIW-) days after the
ents were placed into 100*150 mm Ziploc bags ents. Collected fragmonset of the experim supernatant), transported to the laboratory and immediately processed. lax. 4 m(mAAstststartartart

BB

frfrfragagagstststartartartmentmentment

concontrotrol fragml fragmenents ts phphotosynthesis +otosynthesis +

exexppeeririmementntalal fr fragagmentments s photosynthphotosynthesis -esis -

frfrfragagagstststartartartmentmentment

conconttrrooll fr fraaggmmeennttss

stay onstay on W: W: feedfeeding +ing +
ststaayy on on E: E:feefeeding +ding +

transplanttransplanted from ed from W tW too E E decdecrereaasseded f flluxeuxess&& f foodood s suppupplyly
feeding +-feeding +-trtrananspspllaantnteded fro from E tom E to W W iinnccrreaeasesed fd flluuxxeess&& f foood sod suupplpply y
expeexperriimmeennttaall fr fraagmegmentntssfeedfeeding ++ing ++
Figure 2: Schematic representation of the experimental designs: A) Light-exclusion experiment: 3 donor
colonies per island side, each colony providing 21 fragments (1 start, 10 control and 10 light-deprived
fragments). One chamber set-up on each island side (W/LAIW+ and E/LAIW-). B) Transplantation experiment:
3 donor colonies per island side, each colony providing 20 fragments (1 start, 9 control and 9 transplanted
ents). gmfra

64

Chapter II

Coral processing For each fragment, the full set of parameters described below was analyzed. Coral tissue was
removed from the skeleton using an airbrush and filtered seawater. After homogenization of
aliquots were retained for zooxanthellae density counts and protein analysis, lthe slurry, 6 mand 5 ml aliquots were filtered under 200 mm Hg vacuum (Millipore Vacuum Pump) on glass
an GF/F) and frozen for chlorophyll analysis. mfiber filters (WhatZooxanthellae densities - The total symbiont cell numbers were determined under a
microscope (Leitz, Portugal, 260x magnification) using a Fuchs-Rosenthal haemocytometer.
nts after ean of six replicate couConcentrations were calculated on an areal basis as the mcorrection for the homogenate volume and surface area of the coral fragment.
– Chlorophyll was extracted by adding few 90 % acetone to the Chlorophyll-a analysisples rsons 1972); after cautious shaking, samthawed chlorophyll samples (Strickland & Pawere incubated for 24 hours at 4°C for chlorophyll extraction and centrifuged at high speed ant &ent (Szmeasurem(10000g, 30 sec) to remove all particles in suspension before mal. 2000). Chlorophyll-a concentrations unds 1999, Fitt et an 1990, Gardella & EdmGassmwere determined spectrophotometrically in a Shimadzu UV 1700 1nm Slit photometer at 750
(Lorenzen 1967). and 664 nmnmProtein content- Total protein content was determined after Lowry et al. (1951) using a
albumin standards. Protein Protein Assay Kit, Bio-Rad) and bovine serumDCprotein assay (concentrations were measured spectrophotometrically (Shimadzu UV 1700 1nm Slit) at 750
. nmIn the remthe host tissue was loaded (Millipore Vacuum Pumaining slurry, zooxanthellae and host tissue were separated by centrifugation and p, ~100 mm Hg) on pre-combusted and
pre-weighed filters (Whatman GF/F) and dried before further elemental and isotopic analyses
ent surface al. 2004, Swart et al. 2005). Fragm(see below) (Muscatine et al. 1989, Grottoli et area was calculated using Simple Geometry (determining the geometric form that best
resembles the shape of the fragment and calculating its surface with respective the geometric
formula) to the nearest 0.05 mm and an Approximation Factor for Pocillopora as proposed by
ann et al. (2009). Naum

Total suspended matter, particulate and dissolved organic carbon In the course of the experimthe afternoon, water samples (LAIW+: n = 19ent, during fragm; LAIWent collection and when possible on-: n = 24) were taken by divers close to ce again in
the experimental setup for subsequent analyses of total suspended matter (TSM), total

65

Chapter II

several nolved organic carbon (DOC). Iparticulate organic carbon (TPOC), and dissoccasions, sampling occurred during LAIW-passage, shortly before or after. Therefore,

temperature at time of sampling was recorded (TidbiT v2, Onset, 1 minute resolution and an
ples were taken with 1 l PE bottles, transported to the lab, accuracy of <0.2°C). Water samfiltered (Millipore Vacuum Pump, 200 mm Hg) on pre-combusted and pre-weighed Whatman

ental analyses (see below) and weig elem for followingGF/F filters, driedhed on a r, 1 μg accuracy). Aliquots of the filtrates were crobalance (Mettler, AT21 Comparatoim transferred into pre-combusted glass vials and acidified with phosphoric acid (20 %) to a pHined with a DOC/DIC ere determoncentrations wof 2 before sealing and storage on ice. DOC c

ount DC-190) using a 10-point calibration with TOC standards (ULTRAosemanalyzer (RScientific).

Plankton Plankton sampling occurred only during the day. Concomitant with each fragment sampling,

e eter steel fram diamnear-reef zooplankton was collected by SCUBA push net tows (0.25 m

esh and 1 mwith a 55 μm ming along a 40 m swath along the 20 m isobath, mm sleeve), swi

pling was recorded (TidbiT v2, Onset, perature during sam. Mean tem0.5 – 1 m above bottom1 minute resolution and an accuracy of <0.2°C) to determine LAIW impact at time of
sampling. Samsize classes (55, 100, 150, 200 and 300 μm) over a ples were transferred to the laboratory where they were separated into different fractionation tower. The different size

busted and pre- Pump, 200 mm Hg) on pre-comclasses were collected (Millipore Vacuumss of the plankton ahatman GF/F) and dried for 12 h at 40°C. The dry mweighted filters (Wwas determined gravimetrically using a microbalance (Mettler, AT21 Comparator, 1 μg
accuracy). Clogging was not a problem at the low volumes (8 m3) fished. Filtered volume was
ing mming distance and cross-sectional area of the net opening, assumom swircalculated fith et al. 1968). 100% filtration efficiency (Sm

Pump-sampled off-reef zooplankton was collected from a boat anchored at 35 m depth in
front of the LAIW+ and LAIW- face of Koh Miang (Fig. 1), with the hose intake located in
mid-water 15 meters above bottom and equipped with a temperature logger (TidbiT v2,
nute inute resolution and an accuracy of <0.2°C). Sampling took place in 15 miOnset, 1 mth, land sides by boat the afternoon of the 11intervals for 4 hours simultaneously on both is17th, 21st and 25th of March 2008. Water was pumped through a plastic tube (6 cm diameter,
245 l min-1) and filtered for 5 min through a 50 μm plankton net. Samples were divided in
half using a Folsom-splitter, one sub-sample was preserved in formalin (5 %) for taxonomic

66

Chapter II

te on (data presented elsewhere), thidentificatioher was filtered (Millipore Vacuum Pump, 40°C) for mass determination (m200 mm Hg) on pre-combusted and pre-weighted filters (Whaticrobalance Mettler, AT21 Comman GF/F) and dried (12 h atparator, 1 μg accuracy) prior
to elemental analysis. Pumped volume was determined by assessing the number of seconds it

took to fill an 8 l container (3 trials each) in order to be able to relate plankton values to
e. volumage of the plankton by either of ples showed no detectable damVisual inspection of the sampling procedures. the sam

Elemental and isotopic analyses of coral tissue, total suspended matter and plankton Total carbon and nitrogen content of the coral tissue, as well as the particulate organic carbon

content of the TSM (TPOC) and plankton (POC) was determined using an Elemental

ental CHNS standard (LECO). Carbon A2100 Protein) calibrated against an elemAnalyzer (Nstable isotope ratios (13C) were measured in a gas isotope ratio mass spectrometer (Flash

1112 Analyzer) relative to Pee Dee Belem ontent of TSM (TPOC)nite standard. For organic c

r to analyses until all ified with 0.1 M HCl prioples were acidn (POC), the samand plankto

ples did not require acidification as tissue was oved. Coral saminorganic carbon was remterial. aination from skeletal mobtained without contam

Currents and fluxes ous upward-looking Acoustic Doppler Current Profilers (ADCP) were deployed for Autonomthe time of experiment in the vicinity of the flow chamber setups (RDI Teledyne Workhorse

easure the 3-D Sentinel, 600 kHz and 300 kHz on W and E of Koh Miang, respectively) to m

current field at 1 m vertical and 1 min temporal resolution with an accuracy of 0.3 to 0.5 % of
the water velocity ± 0.3 to 0.5 cm s-1. Data stored in the flash memory of the instruments were
by Rich Pawlowicz, U. ofent, imported into Matlab (rdradcp.mdownloaded after the experim

British Columbia, http://www2.ocgy.ubc.ca/~rich/) and analyzed. Mean daily fluxes of near-

reef and off-reef plankton (total and POC), TSM, TPOC and DOC were calculated by multiplying their concentrations with the average daily current speeds during samplings

pling time). to 12 hours post sam(averaged across 12 hours prior

re TemperatuTo record LAIW incidences during experimental time, temperature loggers (TidbiT v2,
ental setup. TemOnset) were deployed in close vicinity of the experimperature was logged in

67

Chapter II

1 min temporal resolution (with an accuracy of <0.2°C over 0 to 50°C) and data downloaded

using HOBOware 2.2.

alyses Statistial anogeneity of variances using tion and homere tested for normal distribuDatasets w

irnov and Levene’s tests, respectively, transformogorov-SmmKoled if necessary and etric or non-parametric statistical analyses (below), as appropriate. subjected to param

ong sides of the different islands on atial differences between island sides and amTo detect sp

phyll content, tissue position (zooxanthellae numbers, chloro coral tissue comthe basis of

carbon and nitrogen, protein concentrations anmed a two-position) we perford isotopic com

on et al. 2008) using PRIMER v6 ERMANOVA, AndersPfactorial permutational MANOVA (

Gorley 2006). The PERMANOVA allowed us to multivariate statistical software (Clarke & ilarity (using Euclidean distance) between island test for significant differences based on sim

side and island number (nested in island side). Data were log-transformed prior to analysis to

plimaccount for differences is unit sizes. As sang constraints led to an imbalanced data set

over depth and sites, depth differences were ‘regressed out’ by treating depth as a covariate

oving possible depth effects prior to testing for site differences (Anderson et al. 2008, and rempensated for Mirto et al. 2009), and the low and unevenly distributed number of data were com

mutations of the residuals (Gonzalez & Manly ber (n = 9999) of perby running a large numl1998, Anderson 2001, Anderson & Ter Braak 2003, Anderson et al. 2008). The random y

collected reef fragments and the start fragments of both time series experiments were included

in this analysis. ents subjected to the shading (d06 – d32/33) and e series data gained from fragmFor the tim

ents, we developed general linear models to test for the transplant (d02 – d32/33) experim

factors ‘treatment’ (i.e. control, light-deprivation or transplantation; nested within the
ent ) and ‘day’ (over treatment, randomrespective side, fixed), ‘colony’ (nested within treatm

series eodel application, the residuals of all time, fixed) (Satterthwaite 1946). Prior to mtim

(i.e. for each paramjung & Box 1978) on as sted for autocorrelation (Leter and colony) were te

many lags (2 to maximum 33 days) as possible to ensure the effectual independence of data,
despite repeated same colony or rack. Significant differences between plings of the samined using the Fisher LSD post-hoc test. ents and sides were determtreatm

Differences in mortality during the light-exclusion experiment were tested with a survival
and E side were analysis. The survivorship functions (Kaplan Meier curves) of the W

pared using Cox’s F-test. com

68

Chapter II

Water and plankton samples as well as fluxes were statistically tested for LAIW-exposure
using Student’s t-tests. Size differences in near-reef plankton weight, organic content and
isotopic signatures were tested using one-way ANOVA. Pearson’s correlation analyses with W passage and TSM, e relation between LAIperature were conducted to reveal possibltemTPOC, DOC or plankton concentrations. -) current differen-related W (LAIW+) and E (LAIWLAIW Student’s ces were analyzed usingation of the data. Pearson’s correlation analyses between t-test after Box-Cox transformine ollowed by Student’s t-tests were conducted to examperature and current velocities ftemificance. correlations and their statistical sign

Results

re, currents, plankton, TSM, TPOC and DOC fluxes TemperatuThe tem- side e series showed strong differences between the LAIW+ and LAIWperature timof Koh Miang (Fig 3A). Although the modal values were similar (T < 0.2°C), the LAIW+
ore than 4°C occurring at perature drops of up to mface showed a violent spiking with temsubtidal frequencies indicative of the passage of internal waves. The temperature drops were
associated with surges in current velocities (Fig 3A and B, p < 0.001 for LAIW+ and LAIW-),
so that the overall mean current velocity was 30 % stronger on the LAIW+ than on LAIW-
-1, p < 0.001). sside of the island (0.1008 ± 0.0004 vs 0.0772 ± 0.0003 mThe stronger currents resulted in significantly higher (p < 0.001 for all) TSM, TPOC, DOC and plankton fluxes (Fig 4) on the LAIW+ side of the island. Composition of the near-reef
plankton was not different between LAIW+ and LAIW- (Fig. S1) and the concentrations
or off-reef plankton and POC (Figs. S1 and eans ± SE in Table S1), of both near- alone (mS2), TSM, TPOC or DOC (Fig. S3), showed no detectable differences between island sides (p eters (p > perature were not significant for any of the param> 0.05). Correlations with temhore abundance of plankton individuals (not off-reef POC), where the 0.05) except for the offs2pling events exactly within = 0.33; p < 0.05) due to two samcorrelation was negative (r incidents. WLAI

69

Chapter II

Figure 3: Temperature and currents on the W LAIW-exposed (left panels) and E LAIW-sheltered face of Koh
the current Miang. A) Time series data are due to recof temperatoveure ry, clea(black) aning annd red curredent ployvelocity (gray) ment of the current mover the study perieters. B) Correlation beod. Blank peritween ods in
temperature and current velocities. Correlati2ons are significant for either side (both p < 0.001), but stronger for
the W than for the E face (r = 0.06 and 0.005 respectively).

Figure 4and its organic : Daily mean fluxes (± SE) carbon fraction and C) reef of A) near-reef plawater DOC, nkton and TSM and TPOC its organic fromcarbon LAIW+ fraction B) off-ree(black) and LAIW- f plankton
(white) Koh Miang. All LAIW+ and LAIW- side sam ples are statistically significant different.

70

Chapter II

Coral tissue Wposition between LAICorals showed significant differences in their tissue com-exposed and LAIW-sheltered sides of the Similan Islands, but none among the LAIW+ and LAIW- faces
g (MDS) ensional scaline multidimtrated also in thof the different islands (Table 1), as illusples, sam-+ and LAIWplot, showing overlap among but only little overlap between LAIWrespectively (Fig 5). Table 1: Results of the 2-factorial (W vs E and between Islands # 2, 4, 7 and 8) PERMANOVA routine on tissue
composition of Pocillopora meandrina collected along W and E sides of different islands (nested within
respective sideislands do not show significa) after removalnt di of thefferences covariate effect , while differeof deptnces h. betTissue comween W and E positions of are signifraficant gments from(asterisk). differeSS = nt
probability level. p = eans square; F = F-value; of squares; MS = msumsource of varianceSS MSFp

covariate:depth10.8020.80151.0570.351
side17.8787.8783.1290.032*
island(side)65.3550.8931.2540.245
residuals4229.90.712
total5043.935

Figure 5: MDS ordination of fragments from LAIW+ (n = 18 plus 6 start fragments from both experiments) and
LAIW- (n = 21 plus 6 start fragments from both experiments) sides of all islands to illustrate the
multidimensional similarities in coral tissue composition (in terms of zooxanthellae densities, chlorophyll-a,
tissue carbon and nitrogen content, protein concentrations and isotopic carbon ratios) in a two-dimensional
space. The Euclidean distance between two points represents their similarity. Quadrate: Island # 8; circle: Island
# 7; upward-pointed triangle: Island # 4; diamond: Island # 2; downward looking triangles: start fragments from
the experimental set-ups. Filled symbols: LAIW+ side fragments; clear symbols: LAIW- side fragments.
71

71

Chapter II

plings ents (Figs. 6 and 7), we were unable to detect bias due to repeated samIn both experim

ber of donor colonies: we ited num a limfrom

donor colonies and no trend over tim

colonies (Tables 3 and 5).

e (treat

found no significant differences between the

day), neither in control or in experim

ental

conteFigure 6nt, : 2) carbon tissTime-series ue contof all tissue ent, nitrogeparameten tissue crs (fromontent, top t3) o bottomprotein conce: 1) zooxantntrations ahellae dend isnsities, chlorophyll otopic carbon ratios)
maxis as easured in cdays oof expentrols and lightriment). Uppe-deprir left ved fragmsides: ents fromphotosynthesis W + (L(LAIW+) aAIWn+d ). E (LUpper AIW-) right sideover es:xpe photrimosyntheental timsis + e (x-
(LAIW-). Lower left sides: photosynthesis - (LAIW+ ). Lower right sides: photosynthesis - (LAIW-).

72

Chapter II

Figure 7: Time-series of all tissue parameters (from top to bottom: 1) zooxanthellae densities, chlorophyll
mcontent, easured in co2) carbon tissntrols and ue cfeeontent, d-altered nitrogefragmn tissue cents fromo ntent, W (L3) protein cAIW+) and onceE (LAIW-) ntrations anover ed isxperimotopic carbon ratios) ental time (x-
axis as days of experiment). Upper left sides: feeding + (LAIW+). Upper right sides: feeding + (LAIW-). Lower
left sides: feeding ++ (transplanted from LAIW- to LAIW+). Lower right sides: feeding +- (transplanted from
LAIW+ to LAIW-).

73

Chapter II

Table 2: Tissue composition of Pocillopora meandrina. Tissue parameters measured in control (photosynthesis
+) and light-deprived (photosynthesis -) fragments from W (LAIW+) and E (LAIW-) during the light-exclusion
experiment, and in control (feeding +) and transplanted (from LAIW- to LAIW+: feeding ++; from LAIW+ to
LAIW-:as m feeding +eans and standa-) fragmrd errors ents from W(brackets). Missin (LAIW+) and Eg data are (LAIW-) due to mduring tohrtality e trans(light-exclplantation expeusion experimriment are ent) or given
sample loss during analysis (transplantation experiment); one control fragment (LAIW+) from the
transplantation experiment (colony 2, d28) was lost.

LAIW-LAIW+

light-exclusion experimentphotosynthesis +photosynthesis -photosynthesis +photosynthesis -
nmean (se)nmean (se)nmean (se)nmean (se)
zooxanthellae cm-229412205(47578)2695454(16679)25247178(20270)1749383(14237)
μg chlorophyll a cm-2291.80(0.19)260.45(0.10)250.96(0.14)170.29(0.09)
μg tissue carbon cm-229169.08(11.60)2690.44(8.24)25130.88(10.97)1773.38(11.43)
μg tissue nitrogen cm-22925.22(2.17)2614.54(1.51)2518.64(2.03)1711.10(1.79)
mg protein cm-2290.60(0.04)260.42(0.04)250.46(0.05)170.31(0.03)
13C29-17.46(0.07)26-18.21(0.13)25-17.26(0.10)17-17.54(0.18)

transplantation experimentfeeding +feeding ++feeding + feeding +-
nmean (se)nmean (se)nmean (se)nmean (se)
zooxanthellae cm-226463338(36475)27475997(30455)27330515(22089)27328249(28513)
μg chlorophyll a cm-2262.21(0.19)252.24(0.16)261.51(0.11)271.62(0.17)
μg tissue carbon cm-226141.52(7.41)27145.26(7.69)27116.89(6.18)27111.94(7.64)
μg tissue nitrogen cm-22632.36(1.93)2731.58(1.92)2726.19(1.64)2723.89(1.59)
mg protein cm-2260.46(0.03)270.48(0.03)270.39(0.02)250.39(0.04)
13C26-17.85(0.10)27-17.89(0.10)27-17.87(0.09)27-17.71(0.11)

eters were significantly higher ost tissue parament (Fig. 6), mIn the light-exclusion experimfor control corals from the LAIW+ side of the island, compared to the LAIW- controls (Table
ore than 40 % higher, 4). Zooxanthellae densities and chlorophyll concentrations were m-Wore than 20 % higher in LAItissue carbon, nitrogen and protein concentrations were mnly differences in the carbon isotopic ratio of the control exposed corals (Fig. 6, Table 2). Ohost tissue were not significantly different between LAIW+ and LAIW- reefs (Tables 2 and
as well as +W4). Under artificial darkness, about 80 % of all zooxanthellae were lost on LAIon LAIW- and chlorophyll-a decreased to a third of the original concentration on both sides
(Fig. 6, Table 2). Also in LAIW+ as well as LAIW- reefs, losses in tissue carbon and nitrogen
ents were over 40 % while protein content was about 30 % lower than in the control fragmring controls and light-deprived pa(Fig. 6, Table 2). All depletions were significant com though, the protein content decreased on both either side (Table 4). Evenents fromfragm-exposed side, where protein levels of light-rked on the LAIWasides, the decrease was less mdeprived fragments remained about 25 % higher than their LAIW- counterparts and in the
range of the LAIW- control fragments (Tables 2 and 4). During light-exclusion, the LAIW-
e (-0.28 ‰) (Fig. 6, Table 2), while theic ratios only littlents did change their isotopfragmLAIW+ fragment ratios decreased significantly by -0.75 ‰. Subsequently, light-deprived

74

Chapter II

LAIW+ fragments differed significantly from LAIW- light-deprived fragments as well as
ents of both sides (Fig. 6, Table 4). control fragmfrom Table 3: Analysis of spatio-temporal variation of tissue parameters in Pocillopora meandrina fragments
collected in a time series during a light-exclusion experiment. Compared are time series of control and light-
deprived fragments exposed (LAIW+) or sheltered (LAIW-) from LAIW. df = degrees of freedom; MS = means
square; F = F-value; p = probability level. Significant p-values are marked with an asterisk. n.s.: not significant.
source of variancedfMSFp
zooxanthellae cm-2
interceptfixed17.443101131.616*
treat (side)fixed21.70210116.618*
colony(treat)random41.41910100.552n.s.
treatdayfixed21.65410100.643n.s.
error872.5721010
μg chlorophyll a cm-2
interceptfixed114.86529.184*
treat (side)fixed24.6958.911*
coltreaonytday(treat)rafinxeddom240.910.43441.70.83524n.s.n.s.
70.5278errorμg tissue carbon cm-2
interceptfixed1261382.101100.716*
treat (side)fixed211151.6043.911*
colony(treat)random41497.6500.525n.s.
treatdayfixed22853.2691.001n.s.
0.99528578errorμg tissue nitrogen cm-2
interceptfixed15964.49264.522*
treat (side)fixed2384.8134.036*
colony(treat)random480.0130.839n.s.
treatdayfixed27.3890.078n.s.
3795.378errormg protein cm-2
interceptfixed13.85795.532*
treat (side)fixed20.2014.605*
coltreaonytday(treat)rafinxeddom240.000.02860.10.69002n.s.n.s.
40.0478error13Cinterceptfixed15596.60318468.703*
treat (side)fixed22.1897.359*
colony(treat)random40.3271.100n.s.
treatdayfixed21.0643.577n.s.
70.2978error

75

Chapter II

Table 4: Significance levels of Fisher LSD tests for fragments of Pocillopora meandrina of the light-exclusion
experiment. Circles represent LAIW- side, quadrates LAIW+ island side colonies. Light-deprived data sets are
shown by filled, control sets by clear symbols. Significant differences are marked with an asterisk.
photosynthesis +photosynthesis -photosynthesis +photosynthesis -

-2oxanthellae cmozs +photosynthesiphotosynthesis -s +photosynthesiphotosynthesis --2a cmll phyμg chloros +photosynthesiphotosynthesis -s +photosynthesiphotosynthesis --2ue carbon cmtissμg s +photosynthesiphotosynthesis -s +photosynthesiphotosynthesis --2nitrogen cmμg tissue s +photosynthesiphotosynthesis -s +photosynthesiphotosynthesis --2cmmg protein s +photosynthesiphotosynthesis -s +photosynthesiphotosynthesis -C13s +photosynthesiphotosynthesis -s +photosynthesiphotosynthesis -

.000*00.000*.001*0

.004*00.000.015*0

.00100.010*.008*0

.016*00.016*.1370

.026*00.013*.5300

.11300.1830.000*

000*0.0360.

000*0..0461

000*0.309.0

000*0.262.0

000*0.088.0

0.657000*0.

0.000*

0.000*

0.000*

0.000*

0.002*

0.000*

LAIW-exposed corals showed also a much higher dark survival than LAIW-sheltered
ent, scarcely week of the experiments survived well into the thirdens (Fig 8): all fragmspecimexceeding 10 % total mortality at the end of the experiment. The LAIW- corals, by contrast,
ents. The first fatalities were -deprived fragmsuffered heavy losses of about 40 % of all lightdetected already after one week of the experiment and continued until total elimination of the

76

Chapter II

- cases W+ and LAIWulative proportion of LAIents after four weeks. Testing the cumfragmig 8) showed that probabilities of survival ent collection (Fe of fragmsurviving up to the timwere significantly higher for LAIW+ corals (p < 0.001).

cumFigure 8ulative proportion survival curves : Survival analysis of (left y-axPocillopora meandriis, squaresna) in theof corals from light-e LAIxclusion W+ expe(closerimd syment. Kaplabols) an Meier nd LAIW-
(opelight-deprived n symbols) reecorals fs of per Koh sampling. The Miang over exright y-axis periment timdisplaye (x-axiss the ), percent mshowing the coumrtality (circles) ulative proportion of all light-depriof living ved
corals from LAIW+ (closed) and LAIW- (open) reefs of Koh Miang over experiment time. Control corals are
not shown. ent was rapid atization to the new environment (Fig. 7), acclimIn the transplantation experim corals did not differ ntation. All transplantedand occurred within the first days after transplain tissue composition from control corals of their new environment (Fig. 7, Table 6).
However, differences between the transplanted corals and their corresponding control the other island side) and between e donor colony (control corals fromcolonies from the samcontrols from LAIW+ and LAIW- were obvious. Zooxanthellae densities and chlorophyll-a
ssue carbon and nitrogen contents about 20 % concentrations were both about 30 %, tielevated in LAIW+ corals compared to significantly lower concentrations in LAIW- corals
- to the feed-enriched LAIW(Fig. 7, Tables 2 and 6). The feeding ++ corals transplanted fromLAIW+ side ended up significantly enriched in protein content (Fig. 7, Table 6) compared to
- side (feeding +-). Even though to the LAIWents transplanted - controls or fragmthe LAIWdifferences between LAIW+ control corals and LAIW- control or feed-deprived (feeding +-)
corals were marginally not significant (Table 6), protein content was about 20 % higher in the
controls (Fig. 7, Table 2). +WLAI

77

Chapter II

Table 5: Analysis of spatio-temporal variation of tissue parameters in Pocillopora meandrina fragments
collected in a time series during a transplantation experiment. Compared are time series of control fragments
exposed (LAIW+) or sheltered (LAIW-) from LAIW and of fragments transplanted into higher (from LAIW- to
LAIW+) or decreased water fluxes (from LAIW+ to LAIW-), respectively. df = degrees of freedom; MS =
means square; F = F-value; p = probability level. Significant p-values are marked with an asterisk. n.s.: not
nt. significasource of variancedfMSFp
zooxanthellae cm-2
interceptfixed15.3151012236.297*
treat (side)fixed22.540101110.676*
colony(treat)random41.98010100.795n.s.
treatdayfixed22.30010100.965n.s.
error972.3791010
-2chlorophyll a cmμg interceptfixed1100.587156.618*
treat (side)fixed25.6198.131*
colony(treat)random40.5030.728n.s.
treatdayfixed20.2240.324n.s.
10.6994error-2tissue carbon cmμg interceptfixed1483775.133366.965*
treat (side)fixed211456.0027.881*
colony(treat)random4943.4070.649n.s.
treatdayfixed278.8970.054n.s.
error971453.670
-2cmμg tissue nitrogen interceptfixed123834.669219.450*
treat (side)fixed2643.9847.905*
colony(treat)random4183.8102.256n.s.
treatdayfixed29.4240.116n.s.
error9781.461
-2cmmg protein interceptfixed14.878240.621*
treat (side)fixed20.0904.067*
colony(treat)random40.0150.674n.s.
treatdayfixed20.0040.195n.s.
20.0297error13Cinterceptfixed19132.29837651.938*
treat (side)fixed20.1990.722n.s.
colony(treat)random40.1490.541n.s.
treatdayfixed20.2590.938n.s.
60.2795error

78

Chapter II

expeTable 6rim: Signient. Circles reficance levels present of Fisher LSD donor coloniestests for from LAIW- sfragments ide, of quadrates Pocillopora meandridonor coloniesna fromof the trans LAIW+ plside antation
colonies. cultivated Filled symon sheltered LAIW- sidebols represent fragm. Signients exposed to LAIW ficant differences are m(LAIW+), clear symarked with an asterisk. bols are fragments
feeding +feeding +-feeding +feeding ++

-2oxanthellae cmozeeding +f +-eedingfeeding +feeding ++f-2a cmll phyμg chloroeeding +f +-eedingfeeding +feeding ++f-2ue carbon cmtissμg eeding +f +-eedingfeeding +feeding ++f-2nitrogen cmμg tissue eeding +f +-eedingfeeding +feeding ++f-2cmmg protein eeding +f +-eedingfeeding +feeding ++fC13eeding +f +-eedingfeeding +feeding ++f

0.958.002*00.001*

0.619.003*00.002*

0.634.021*00.007*

0.352.015*00.031*

0.898.06800.025*

0.283.88700.868

002*0.0.001*

011*0.0.009*

006*0.0.002*

001*0.0.002*

096.0038*0.

356.0216.0

0.766

0.908

0.722

0.754

0.685

0.760

79

Discussion

Chapter II

d the possibline water into reef communities anocIntroduction of upwelled subthermle effects

on water quality (Andrews & Gentien 1982, Leichter et al. 1996, Leichter et al. 2007), tbiodiversity (Cortés 1997), coral growth (Leichter & Genovese 2006) or feeding (Palardy e

pare reefs growing in close al. 2005) have been addressed in previous studies. Here, we comre severe operature oscillations mvicinity around an island chain unilaterally exposed to tem

and frequent then previously reported in other areas (Leichter et al. 1996, Leichter & Genovese 2006) and among the highest so far observed (Sheppard 2009). Given the close

-) sides of the islands (<200 m), the differences in W (LAIW+) and E (LAIity of the Wproxim

the two island sides are striking. Variability eters betweenhic paramthe physical oceanograp

also oxygen, pH and nutrient concentrations perature and currents (this study), but in tem

(Schmidt et al. subm.) are much more pronounced on LAIW-exposed reefs compared to their

peratures, stronger mean current velocities ean temsheltered counterparts, resulting in lower ma(Fig. 3) and increased input of corrosive, nutrient-rich deep wter.

We demonstrated that in spite of the disparities in tissue composition of P. meandrina

between LAIW+ and LAIW-, the average amount or stable isotope composition of the
terial). TSM, TPOC (near- and off-reef), aental milar (supplemplankton and the TSM are sim

, pact; howevermDOC or near-reef plankton concentrations are not dependent on LAIW i

s to be depleted in planktonic individuals. The lack of difference in the ter seemaupwelled w

ical oceanographic parambiological and chemters appears at odds with the pronounced e

processes other than ables, suggesting that differences in the physical oceanographic varixing are involved. im

Previous studies showed that internal waves can act as a ‘plankton pump’ supplying phyto- unities (Wand zooplankton to benthic comm et al. 1993, Leichter et al. 1998), thus anitm

an Sea featuring a corals (Leichter & Genovese 2006). In the Andamfuelling growth rates ofpronounced oxygen minimum zone (OMZ) below the surface mixed layer, the concentration

thermocline, the concentrations of phyto- aof plankton depends on the depth of the upwelled water: Below the surface mnd zooplankton decrease dramiatically along with xed layer and

oxygen concentrations (Madhu et al. 2003, Nielsen et al. 2004) leading to lower zooplankton perature) upwelled water. During periods l temodaconcentrations in cold (<3 °C relative to m

ent and depletion of nutrients from ittent upwelling, enrichmof weak or intermupwelling/mixing and primary production, respectively, may even out (Kinsey 1988, Cushing

earn et al. 2001) and nt uptake by corals (H1989). Increased current strength increases nutrie

80

Chapter II

ant 2002), which has further been shown to be ary production (Szmfuels zooxanthellate primperatures of 23 to 26°C (Al-Horani 2005). Strong currents highest in cooler water with temkness of the boundary layer over enhance photosynthesis and calcification by altering the thic (Dennison & Barnes 1988). Current-induced the coral tissue and increasing gas exchangeturbulence may also mitigate the negative effects of oxygen-low water on coral metabolism
y be further enhanced by coral tentacle expansion a(Shashar et al. 1993), where turbulence m(Patterson 1992) during feeding (Sebens & DeRiemer 1977). At the same time currents are
indispensable for corals (Sebens et al. 1998), which are passive suspension-feeders whoseprey capture potential raises with increasing current strength (Sebens & Johnson 1991). ik 2001), oesura & van WmSurvival probability of corals exposed to high temperatures (Nakaluenced by increased ely infsitivoas well as rates and time of recovery after bleaching are pura et al. 2003). mwater flow (NakaLow temperature may further affect coral metabolism in various ways. Saxby et al. (2003)
ance in waters <20°C and other studies showed observed a decrease in photosynthetic performing activity of corals on zooplankton during that slowed polyp contraction decreased the feede periods (Palardy et al. 2005) and in cold water (Johannes & Tepley 1974). upwelling timidt -distinctive factor (SchmWAlso ‘corrosive’ low-pH water (Feely et al. 2008), another LAIing .), is known to alter trophic pathways by diverting energy from energy consumet al. submatic growth (Fine & Tchernov 2007). calcification into somBecause the LAIW-induced unfavorable conditions in terms of temperature, oxygen and pH
persist for only minutes (Fig. 3), it is difficult to assess their potential impact on coral status
e over days and weeks. It is also difficult to discern antagonistic effects, e.g negativ possibly positive nutrient effects on coral photosynthesis. perature effects fromtemaffect coral feeding, where positive effects of enhanced y also aAntagonistic effects maplankton supply mtive effects of polyp retraction (Johannes & y be offset by possible negaTepley 1974). As sampling was limited, for logistic reasons, to day-time hours, both, the nocturnal feeding
mer 1977) habit of the corals (Lewis & Price 1975, Muscatine & Porter 1977, Sebens & DeRieankton (Porter & Porter 1977, Heidelberg et ergence of demersal zoopland the nocturnal emal. 2004, Genin et al. 2005, Yahel et al. 2005) may have introduced a bias in our analysis with itigated in our bias is likely to have been m. The potential feeding respect to coral feeding subjected to constant darkness and feeding ents wereent where light-deprived fragmexperimay have e. The plankton bias, on the other hand, mwas presumed to take place at any tim-Wilan reef, given the lack of a coral framework on the LAIaffected only the sheltered E Sim

81

Chapter II

ersal exposed W side of the island and the observation that the concentration of dem

ith the comzooplankton increases wplexity of the reef substrate (Porter & Porter 1977).

In artificial darkness, LAIW-exposed P. meandrina was able to subsist exclusively on

ig. 6). The sharp drop in zooxanthellae numheterotrophy and energy reserves (Fbers and

chlorophyll-a concentrations illustrates the capacity of the light-deprived corals to rapidly

of phototrophy, in contrast to the LAIW sheltered corals from tabolically to the lack eadapt m

E Koh Miang that eventually all died (Fig. 8). The concomitant decrease in the surviving

iniscent of thecorals’ tissue carbon, nitrogen and total protein concentrations (Fig. 6) is rem

ant & Gassm (Szmes in tissue carbondeclinan 1990) and lipid concentrations (Grottoli et al.

rals and attributed to the consumption of or bleached co2004) which have been reported f

was reduced. Also the n to coral metabolismenergy reserves when photosynthetic contributio

ass between the spite of eroding differences in tissue biomhigher protein concentrations, in

light-deprived LAIW+ and LAIW- corals, indicates a sustained supply of protein-rich

-exposed reef, as heterotrophic carbon has been found to be Wplankton food in the LAI

inly as protein (Bachar et al. 2007). This is further aincorporated into cnidarians mcorroborated by the stable isotope data showing 13C depletion for light-deprived LAIW-

gnitude to the depletion reported for vigorously ailar in mexposed corals only (Fig. 6), sim

feeding 6). Strong during and after bleaching (Rodrigues & Grottoli 200Montipora capitata

water flows do not only enhance food supply (Sebens et al. 1998), but also prevent a steady-

state boundary layer over the coral surface hence increasing suspension feeding of particulate

uth & Sebens 1993), eventually resulting terial (Helmaand adsorption of dissolved organic m

deprived phototrophy comin longer and higher survival in periods of-Wpared to LAI

unaffected corals.

mpanied altered food -exposure and –shelter as well as to the accoatization to LAIWAcclim

bers, chlorophyll-a content, provision was rapid and resulted in increased zooxanthellae num

tissue carbon, nitrogen, and proteins when exposed to LAIW and in their decrease when

transplanted out of LAIW-impact (Fig. 7). While upward changes in zooxanthellae numbers

and chlorophyll-a content might be the combined result of increased nutrient availability

an eant 2002), currents (Dennison & Barnes 1988, Hearn et al. 2001) and overall lower m(Szm

ergy reserves are the peratures (Al-Horani 2005), our results suggest, that the higher entem

comconsequence of heterotrophic nutrition acting inbination with photosynthesis (Borell et

ents, where fed corals exhibited ilar to previous reports of feeding experimal. 2008). Sim

arved ones (Ferrier-Pagès et al. 2003, Borell et al. 2008) and higher levels of protein than st

fed with nditions andlipid levels were increased when corals were kept under low light co

82

zooplankton (Treignier et al. 2008), the higher tissue carbon, nitrogen and protein

Chapter II

d exposeergy stores in LAIWconcentrations indicate higher heterotrophic input and larger en

corals.

y vary aOur results show that heterotrophic plasticity and trophic status of a coral m

ent, particularly water flow (Skirving & Guinotte intraspecifically, depending on its environm

-enhanced supplies of food may be crucial for coral resilience W2001) and food supply. LAI

or inactivated photosynthesis, as are known to to stress and survival during periods of reduced

adapts P. meandrinarown 1997, Hoegh-Guldberg 1999). occur after coral bleaching events (B

rapidly to changing environmental conditions, but might only have limited heterotrophic

position in thplasticity, as indicated by the lack of change in isotopic carbon com e sheltered

upply is corals. The potential of plasticity, however, is enhanced when food s-WLAI

increased by increasing current strength as shown for the LAIW+ coral fragments. Because

other coral genera may be more efficient feeders than Pocillopra (Palardy et al. 2005, Palardy

atize variously to LAIW influences. In the climet al. 2008), it is likely that different species ac

absence of genetic data, however, we can only speculate whether such profound differences

atization to LAIW. acclimray reflect adaptation om

As LAIW are ubiquitous in the ocean, particularly in tectonically active areas such as South

East Asia featuring a rich underwater topography, strong density stratification and tidal

portant yet unexplo may play an imcurrents, LAIWred role in the capacity of corals to adapt

to a changing marine environment.

ledgments Acknow

cean-Reef Coupling Oan bilateral ORCAS (ai-GermResearch was carried out as part of the Th

in the Andaman Sea) program and funded by the German Research Foundation (DFG). The

ilan MBC) and the Simauthors would like to thank the Phuket Marine Biological Center (P

as well as Tobias Funke for technical and Island National Park staff for field assistance,

Dorothee Dasbach for laboratory help. Statistical advice was kindly provided by Prof. K.R.

Clarke and Dipl.-Math. Werner Wosniok. We thank three anonymous reviewers for very

nuscript. aents and an extensive revision of the mhelpful comm

83

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87

ental data Chapter II Supplem

Chapter II

Table S1: Composition of near-reef (total mass, POC content and 13C ratio) and off-reef (zooplankton numbers,
total mass and POC content) plankton and DOC, TSM and TPOC compositions in reef waters.
LAIW+LAIW-
near-reef plankton mass (mg m-3)33.9 ± 5.526.9 ± 2.6
near-reef POC (mg m-3)0.63 ± 0.090.49 ± 0.07
near-reef 13C (‰) -22.3 ± 0.11 -22.52 ± 0.21
off-reef zooplankton numbers (individuals m-3)4.0103 ± 0.41034.5103 ± 0.4103
off-reef plankton mass (mg m-3)53 ± 6.943 ± 2.4
off-reef POC (mg m-3)4.5 ± 0.34 ± 0.2
DOC (mg m-3)1.86103 ± 0.7 1031.66103 ± 0.3 103
TSM (mg m-3)12.40103 ± 0.7 10313.39103 ± 0.2 103
TPOC (mg m-3)75.06 ± 2.699.36 ± 3.4

2560)-32040g m0notkanl peefr-earn
itubirstiss da m )% (no100150200300100150200300
notkanlp eefr-ear n 05050
(m20156051.)-301.40 CO Peefr-aren
200.5 mg(m
fe-rerane 00.0
13Con (%)itubritsidC OP -19m55 μm100 μm150 μm200 μm300 μm55 μm100 μm150 μm200 μm300 μ
0-2C3-2-27on ktna plfeer-rane
134-2 nonkta plfe-rera ne m μ55m00 μ1m50 μ1m00 μ2m00 μ3m μ55m00 μ1m50 μ1m00 μ2m00 μ3
LAIW+LAIW-BLAIW+LAIW-
AFigure S1: Near-reef plankton composition on LAIW+ (left panels) and LAIW- reef sides (right panels) of Koh
Miang. A) Mean mass, particulate organic carbon proportion and isotopic carbon ratios (± SE) of each size class.
No significant thdifferethnce between size classes or sides. B) Central tendency box plots (line: median, box:
variability (25 and 75 percentile), whiskers: non-outlier range) with extremes (circles) of near-reef plankton
mass, particulate organic carbon and isotopic carbon ratios. All size classes within each side were pooled. No
significant differences between sides. See Table S1 for values.
88

88

Chapter II6)-3 m3onnktaoopl zfeer-fof*1nd.i(0
30350150)-3 mnotknal pfereff-o50
100g(m08)-34feref-fo mg (mCO P
0LAIW+LAIW-
Figure S2: Off-reef plankton composition in LAIW+ (left panels) thand thLAIW- reefs (right panels) of Koh Miang.
Central tendency box plots (line: median, box: variability (25 and 75 percentile), whiskers: non-outlier range)
with efromx LAIWtrem+ anes (circles) d LAof IW- and off-reef zparticulooplaate nkter conceorganic carbon fromntrations from LAIW+ a LAIW+ annd Ld LAIW-. No siAIW-, gnioff-reef plaficant nktdiffeon mrence ass
between sides. See Table S1 for values.
)15-3 m5mg301 (DOC0)-320m 15mg301015TSM (0)020-3015010OTP mgm (C50
0LAIW+LAIW-
Figure S3: Biogeochemical composition of near-reef waters in LAIW+ (left panels) and thLAIW- threefs (right
panels) whiskers: of Koh Miang. Centnon-outlier range) with ral tendency box plots extremes (line: m(circles) of dissolvededian, box: vari orgaability (25nic carbon, aTSM nd 75and peTPOC rcentile),
concentrations. No significant differences between sides. See Table S1 for values.

89

Comparative metabolic performance of & Claudio Richter

Cornelia Roder1, Gertraud M. Schmidt2, Carin Jantzen1, Somkiat Khokiattiwong3
2& Claudio Richter

Leibniz Center for Tropical Marine Ecology, ZMT, Fahrenheitstrasse 6, 28359 Bremen,

1

Alfred Wegener Institute for Polar and Marine Research, Am Alten Hafen 26, 27568

Germany2 Alfred Wegener Institute for Polar and

Bremerhaven, Germany3Phuket Marine Biological Center, 51 Sakdidet R

Phuket Marine Biological Center, 51 Sakdidet Road, 83000 Phuket, Thailand

In preparation

(LAIW)-exposed and LAIW-protected habitats

lutea

Porites

- Chapter III -

Chapter III

90

from Large Amplitude Internal Wave

Abstract

Chapter III

e, 1860) were collected in the m (Milne-Edwards and HaiPorites luteaCoral fragments of itude pl areas subjected to Large Amilan Islands (Thailand), Andaman Sea, fromSimInternal Waves (LAIW), and LAIW-sheltered controls less than 1 km away. In a field
laboratory, the fragments from two depths (7 and 20 m) were all incubated in the same
tabolic potential under eperature bath under screened natural sunlight to assess their mtemsimulated LAIW-free conditions at shallow depth. Corals from the LAIW-impacted sites
ass and displayed lower photosynthesis and calcification but higher were richer in biomrelease rates of dissolved and particulate organic material compared to LAIW-sheltered
aps even genetic atization (or perhetabolic acclimls. The differences suggest mcontroadaptation) of LAIW-exposed coral to lowered pH and temperature, and increased fluxes
aterials. of inorganic and organic m

Introduction

4). Large amplitude internal (Jackson 200al waves are ubiquitous in the oceanInternwaves (LAIW) may form when stratified waters are advected across abrupt changes in
underwater topography, such as seamounts and sills (Farmer and Armi 1999; Jackson
eters y travel for hundreds or thousands of kiloma m2004). Over the deep ocean, LAIWaway from their generation sites until approaching shallow bottom, e.g. near the econdary waves (Vlasenko disintegrate into scontinental shelf, where the waves dissipate, ay and Alpers 2005), or break (Vlasenko and Stashchuk 2007). The resulting turbulence mc and low-pH waters into shallow areas x subthermocline cold, nutrient-rich, sub-oxiiman Sea eichter et al. 1998). The Andam; Lolanski and Delesalle 1995(Pineda 1991; Wfeatures LAIW of exceptional amplitude (Perry and Schimke 1965; Osborne and Burch
ilan Islands, an island chain off the Thailand coast and close to the1980). The Simcontinental shelf break is located near the swash zone of LAIW generated near the
Andaman-Nicobar island arc, some 400 – 600 km away (Jackson 2004). The west faces
s of up to 10°C, on the scale of less than oneal oscillationof the islands show large therm

91

Chapter III

hour (Roder et al. accepted), corresponding with the passage of intense LAIW. The west
monsoon period, lack s during the SWsides, which are also exposed to seasonal stormhansang et al. 1999): corals growing in often flattened shapes occur in loose true reefs (Cassemblages directly on the granite basement (Schmidt et al. submitted). In less than one
ilan Islands, true reefs are found st (E) sides of the Sim distance, on the sheltered eakmework (Chansang et al. 1999). with dense cover of corals growing on a carbonate framThe immediate vicinity of such contrasting coral habitats raises the question if LAIW
may have favored physiological acclimatization or perhaps even genetic adaptation
within a given coral species (e.g Porites lutea), or if the specimens growing along the W
Similan Island sides only tolerate the combined effects of LAIW-introduced low-pH
unds 1999) (Macyntire et al. 1993; Levitus 1994) and low-oxygenated (Gardella and Edmant 1997) and high waters, high loads of inorganic nutrients (Muscatine et al. 1989b; Szm ).aterial (Roder et al acceptedfluxes of suspended or particulate organic mperature, light and ents under normalized temIn this study, we use incubation experim (i.e. oxygen production, respiratory tabolismenutrient conditions to assess the coral mtter) in acification and exchange rates of particulate and dissolved organic mand, caldemresponse to the acclimatization history (or adaptation) to varying degrees of LAIW-
exposure.

Material and Methods

To examine the metabolic potential of coral specimens from LAIW-affected and LAIW-
incubations were conducted between 10 and nditions, mparable cosheltered sites under co12 hours in the weeks from 08th and 18th of March 2007 under a shaded tent
(Photosynthetic active radiation PAR of 35 and 45 μmol m-2 sec-1, continuously
monitored with the light-sensor of a DivingPAM (Waltz, Germany)) on land of Koh
Miang Island (Similan Island # 4) with coral fragments freshly collected in deep (20 m)
and E island reef sites of Koh Miang (Fig. 1). The four waters of W) (7 mand shallowchosen sites resemble decreasing LAIW impact from reefs of W deep > W shallow > E
idt et al. subm.). deep > E shallow (Schm

92

NN

Chapter III

Figure 1: Koh Miang Island (extract), the fourth of the Similan Islands in the Andaman Sea, Thailand
(left), with sampling sites (circles) in the reefs W and E of the island, incubation site (cross) on land and
llection site (quadrat). water coincubations

the Bay area in front of the p) from(BADU Magic 8 Pumped Incubation water was pum depths and filled into a water tank housing four ately 4 m approximincubation site fromental chambers, one blank transparent and four opaque incubation beakers (three experimfor monitoring purposes to guarantee consistent water quality) of 2 l volumes which
posing a cover plate outfitted with in- and outlets for could be gas-tight closed superimp lti-channel peristaltic pumutubes. Through these tubes, the water was rotated with a m(521VK, Watson-Marlow, UK) after closure of the chambers to ensure constant motion
of the water and moderate circulation. Every morning, fragments (two per colony, one for
2 were chiseled ber) of sizes from 20 – 50 cme chamthe transparent and one for the opaque, 1860) colonies (Milne-Edwards and HaimPorites lutear surface of large off the uppeplanted ediately to the field laboratory where they were separately imand transported immnts to air. After a short ebers without exposing the fragminto incubation chamacclimatization time (~ 20 – 30 min), the oxygen content in the tank was measured using
ples were taken for nutrient and alkalinity an optode (HQ40, Hach Lange), triplicate saments (see below), the chambers were closed, taking care to avoid bubbles measurembers were nutes. After the incubation, the chamiinside, and incubated for 118 to 124 m and 1 l aliquots of lmed (HQ, Hach Lange) and 50 opened, the oxygen content record

93

Chapter III

ses of total alkalinity and dissolved orincubation water taken for subsequent analytter respectively (see below). Net photosynthesis was calculated aparticulate organic mbers, respiration trations in the light cham the temporal difference in oxygen concenfrombers. Gross photosynthesis was calculated as the the oxygen uptake in the dark chamfromthe alkalinity anomsum of net photosynthesis and darkaly method (Schneider a respiration.nd Er Calcificatioez 2006). Fn was mor calculation of calcification easured according to
rates, 50 ml samples taken prior and after incubation were filtered (Whatman GF/F) by
hand using a syringe and total alkalinity was immediately measured to the second end point (Gran 1952) using a solution of 0.01 mol l-1 HCl + 38 g l-1 NaCl (Titrisol, VWR)
ples taken prior and after ). Further, 1 l samrino, Methromand a titration apparatus (Titincubation were filtered (Millipore Vacuum Pump, 200 mm Hg) over pre-combusted and
pre-weighed Whatman GF/F filters for subsequent analyses of dissolved and particulate
lters were desiccated (12 hours at 40°C) itter (DOM and POM respectively). Faorganic macidification with 0.1 M HCl to remand cut in half for separate analysis ofove inorganic carbon) particulate organic carbon (POC) (after and particulate nitrogen (PN)
ntal CHNS standard (LECO). eental Analyzer and an elemusing a NA2100 Protein Elemnitrite, ammoniumThe filtrate was stored frozen until analysis) and total dissolved nitrogen (P of inorganic nitrogarsons et al. 1998) using a en compounds (nitrate,
eter (GBC model UV/VIS918). Dissolved organic nitrogen (DON) was spectrom the total pounds fromcalculated by subtracting concentrations of inorganic nitrogen com). Dissolved organic carbon ple (Ferrier-Pagès et al. 1998nitrogen content of the sam a neasured after acidification with phosphoric acid (20 %) to a pH of 2 i(DOC) was mount DC-190) using a 10-point calibration with TOC DOC/DIC analyzer (Rosemstandards (ULTRA Scientific). To gain production / uptake rates, concentrations
measured in the start samples taken prior to incubation were subtracted from final
concentrations for all parameters. Blank chamber rates were calculated and verified to not
izedtake rates were standardexceed the range of standard deviation. All production / upper cm-2 and h-1 by accounting for the volumes of the coral fragments (measured in small
timbeakers via displaceme and calculating tissue area to the nearesent after) in the incubation chambers, normt 0.05 mm using a caliper and geomalizing for incubation etry as
ents were sacrificed ann et al. (2009). After incubation, coral fragmproposed by Naum

94

Chapter III

sing an artist’s airbrush and filtered seawater. Tissue and tissue stripped off the skeleton uogenized for 30 seconds (Ultra Turrax) and aliquots retained for the slurry was homanthellae densities were calculated by six replicate counts under alyses: Zooxfollowing angnification) using a Fuchs-Rosenthal acroscope (Leitz, Portugal, 260× mithe mhaemocytometer. Five ml of the homogenate were filtered (Millipore Vacuum Pump, 200
mm Hg) on Whatman GF/F filters and frozen prior to chlorophyll-a extraction after
adzu UV 1700 trickland and Parsons 1972) and analysis in a Shimstandard procedures (S1nm Slit photometer at 750 nm and 664 nm (Lorenzen 1967). A known volume of the
ogenate was centrifuged for separation of coral host tissue and aining tissue homremzooxanthellae and each component was loaded (Millipore Vacuum Pump, ~100 mm Hg)
separately onto pre-combusted glass fiber filters (Whatman) as described in Muscatine et
hours at 40°C) and coral host and zooxanthellae Filters were dried (24al. (1989a).ental Analyzer emlined in an Eorganic carbon and nitrogen contents were determe eters werandard (LECO). All coral paramental CHNS st(NA2100 Protein) using an elemcorrecting for homogenate volume and tissue surface area. Each site (W deep, W shallow,
E deep, E shallow) was sampled twice; hence, in total, 8 incubations, each with 3 light-
ality of the ents were conducted. Due to non-normmexposed and three shaded coral fragce (Kruskal and Wallis 1952), followed by alyses of varianetric andata, nonparamilcox 1964) were ilcoxon and Wination of differences between the groups (Wdetermconducted.

Results

shallow corals, followed by W The highest zooxanthellae densities were found in Wdeep, E deep and E shallow specimens (Table 1). Coral host as well as zooxanthellae W e pattern and were highest for the carbon and nitrogen concentrations followed the samside shallow water specimens. Only chlorophyll-a concentrations were highest in the
de, with second highest concentrations of the W shallow water sideep corals of the Wens. specim

95

Chapter III

610zoox-2anthellae μg chlor-2ophyll-a μg coral host -2μg zooxanthell-2ae μg coral host-2 μg zoocanthe-2llae
cmcmcarbon cmcarbon cmnitrogen cmnitrogen cm
W deep5.51 ± 0.5811.50 ± 1.17179.96 ± 10.79638.00 ± 41.1522.92 ± 1.5483.68 ± 5.40
W shallow9.92 ± 3.169.18 ± 1.42241.76 ± 15.30724.36 ± 83.6832.28 ± 2.1389.68 ± 8.47
E sE deephallow4.45.65 4 ± 0.± 2.92564.37.33 5 ± 0.± 0.42851143.72.84 ± 32 ± 47..990739497.38.22 4 ± 16.± 29.76021272.8.84 8 ± 0.± 1.56114530.4.74 2 ± 3.± 1.6688

Table 1: Tissue parameters ± Koh MiaSE of ng IslPorites luand used in itea fragmncents fromubation ex deep aperiments. nd shallow reef sites of W and E

f Overall, we found the highest net oxygen production (Fig. 2; p < 0.05) in the deep reeareas of the E island side (33 ± 5 mg cm-2 h-1). For shallow water corals, the net oxygen
production was higher (p < 0.05) in the E (16 ± 3 mg cm-2h-1) than in the W. Depth-
related differences in net oxygen production were significant in the E but not on the W ens (-30 ± side specimside of the island. Respiration rates (Fig. 2) were highest in deep W2 mg cm-2h-1) and significantly different (p < 0.05) from the other site specimens which
did not differ from each other. Calcification (Fig. 3) in the transparent chambers (‘light’)
ents (p < was significantly higher than the corresponding dark calcification for all fragm the E reef (1.2 ± 5 w water corals from0.05). Highest values were found for the shalloμmol cm-2h-1; p < 0.05).

40

)-1 h20-20 prmg cmon (itoduc2-20
deep light
O

0-4

EW

deep darkshallow lightshallow dark

Figure 2: Net oxyareas of gen producW and E tion and resKoh Miang piration Island incubrates ± SE of ated Porites luteaunder equal conditions. from deep and shallow reef

96

51.

)-1 h-2mol cm01. production (μ0.53CaCO00.deep light

deep dark

EW

shallow lightshallow dark

Chapter III

Figure 3: Calcium and E carbonate production rates Koh Miang Island inc± SE of ubated Porites luteaunder eq fromual conditions deep an. d shallow reef areas of W

tter (DOM) were highly aProduction and consumption rates of dissolved organic mvariable (Figs. 4 and 5). Dissolved organic carbon (DOC, Fig. 4) production occurred in light-exposed shallow water corals, both in E (7.3 ± 4 μg cm-2h-1) and W (11.8 ± 8 μg cm-
-12). DOC uptake was observed in both, light incubated and dark deep water corals from hE (-13.3 ± 6 and -12.7 ± 7 μg cm-2h-1, respectively). The results from the other
ON, incubations were not significantly different from zero. Dissolved organic nitrogen (DFig. 5) was produced exclusively by light-incubated shallow water corals from the E
(0.08 ± 0.02 μg cm-2h-1) whose production rates were significantly higher (p < 0.05)
ainly ure. DON was mregardless of light expospared to their deep water counterparts, com was higher (p < 0.05) in the taken up by deep water corals. Uptake in E deep water coralslight (-0.18 ± 0.07 μg cm-2h-1) than in the darkness.
Particulate organic matter (POM) fluxes were almost exclusively positive (except for PN-
measure of one fragment from the deep E reef site), indicating release of particulate
ationcubto the inorganic carbon (POC, Fig. 6) and particulate nitrogen (PN, Fig. 7) in both, the light ls (p < 0.05) inwater. POC release rates (Fig. 6) were highest for deep cora(7.5 ± 2.5 μg cm-2h-1) and dark incubations (5.6 ± 1.7 μg cm-2h-1). For deep and shallow
water corals there were no significant differences between light and dark incubations. PN

97

Chapter III

ilar pattern, with no significant differences in release rates (Fig. 7) followed a sim a tendency of increased release under light ent, nevertheless withresponse to light treatmtly higher (p < 0.05) release rate for corals ents and a significan side coral fragmfor the Wfrom the deep W side reef incubated under light (1.96 ± 0.6 μg cm-2h-1) as well as in
opaque chambers (1.2 ± 0.4 μg cm-2h-1).

EW

30EW)-1 h15-2g cm0deep lightdeep darkshallow lightshallow dark
DOC production (μ5-1

0-3Figure 4: Dissolved organic carbon (DOC) production or uptake rates ± SE of Porites lutea from deep and
r equal conditions. undecubated W and E Koh Miang Island inshallow reef areas of

30.

EW

)-1 h-210.mg cμon (itoduc-0.1
deep lightdeep darkshallow lightshallow dark
rDON p

3.-0Figure 5: Dissand shallow reolved orgaef areanic nitroges of W an nd E (DON) production or Koh Miang Island incubated uptake rates ± SE of under eqPorites lual conditions. utea from deep

98

12

EW

Chapter III

)-1 h-2g cm8POC production (μ40deep lightdeep darkshallow lightshallow dark
Figure 6: Particulate organic carbon (POC) production rates ± SE of Porites lutea from deep and shallow
reef areas of W and E Koh Miang Island incubated under equal conditions.

EW

3EW)-1 h2-21mg cμon (itoduc prNPdeep lightdeep darkshallow lightshallow dark
0

-1Figure 7: Particulate nitrogen (PN) production rates ± SE of Porites lutea from deep and shallow reef areas
ions. cubated under equal conditIsland inW and E Koh Miang of

99

Discussion

Chapter III

This study shows pronounced lateral (W vs E) and vertical (shallow vs deep) differences

in coral metabolism, suggesting different acclimatization histories (or perhaps

adaptations) to LAIW in terms of photosynthesis, respiration and exchange of dissolved

tter (DOM and POM). aand particulate organic m

Tissue conditions mirror the nutrient regimes of the corals’ original environments with

enhanced zooxanthellae densities and chlorophyll-a concentrations in response to large

). In the shallow water areas of the Wes (LAIWplitude internal wavam side, the

idt et al. ent (Schmnvironmbined effects of high nutrient input and a high light ecom

subm.) result in increased zooxanthellae densities (Brown et al. 1999) and biomass

production (Fitt et al. 2000). In addition to strong currents resulting in higher food

supplies coral host biomass production is also enhanced (Roder et al accepted). The

rass as an energy store (Fitt et al. 2000; Gbenefit of high biomottoli et al. 2004; Bachar et

ands of a thick tissue (Spence and Hynes al. 2007) is counteracted by the respiratory dem

1971) resulting in lower net oxygen production of the holobiont. Therefore, specimens

from less LAIW-impacted sites (E) are more effective oxygen producers dispensing

nts. Calculating e side fragmn compared to Whigher rates of oxygen into the water colum

ary production by adding the absolute values of respiration and net oxygen the gross prim

production (McCloskey and Muscatine 1984) reveals still higher gross oxygen production

rates for corals from the E reef sites despite higher zooxanthellae densities and pigment

W side corals suggesting a more costly photosynthesis under LAIconcentrations for W

the tight coupling with photosynthetic because oft-enhancedinfluence. Being ligh

activity (Furla et al. 2000; Marubini et al. 2001), calcification was higher in the

ated specimens. Calcification rates were pared to dark incubbers comt chamtransparen

highest in E shallow coral fragments indicating that other processes besides

photosynthesis are involved as oxygen production was clearly higher in deep water

specimens from the same side. Coral growth rates have been documented elsewhere to be

higher in shallow waters (Falkowski et al. 1990; Furla et al. 2000; Marubini et al. 2001)

ght be due to ients in our study. This m side fragmas is seen for the E, but not for the W

ined by the ral calcification is also determance, but cotheir lower photosynthetic perform

100

Chapter III

aragonite saturation state (Marubini et al. 2001) which is rather driven by the variablecarbonate ion than by the nearly constant Ca2+ concentrations in the water column
concentrations are high O(Stumm and Morgan 1981). In upwelling deep sea water C2nd low aragonite saturation state. As a (Macyntire et al. 1993), resulting in low pH aented for the reefs in carbonate production is decreased as documresult, calciumupwelling areas around Galapgos (Glynn 1988) or in the eastern Pacific (Cortés 1997).

The exposure of W side coral specimens to corrosive (Feely et al. 2008) water might limit
coral growth regardless of light availability.
e, with high light adapted ly coupled with the light regimAlso DOC release rates are tightore, DOC is ore than low light adapted ones (Crossland 1987). Furthermcorals releasing m

ounts when corals are exposed to increased nutrient released in increasing amconcentrations and in even higher amagès et al. 1998). ounts when being fed (Ferrier-P

High light and nutrient input plus an increased availability of organic material (Roder et al accepted) due to LAIW-impact result in DOC release, especially in corals from the
mashallow Winly in oligotrophic environm side. On the other hand, corals have also been documents using it as an additional food source (Sorokin 1973; ented to take up DOC

ore conservatively, Grover et al. 2008). With DON release corals tend to behave mrrier e al. 1998). Founts of DON in relation to DOC (Ferrier-Pagès etretaining higher amPages et al. (1998) found that the amount of DON released was equal in fed, starved and excreted inental setups. POM by contrast was constantlynutrient enriched experim all release rates of DOM as suggested before ental setups and constantly higher thanexperim ild et attribute the release of POM to mucus production (Weby (Tanaka et al. 2008). Went loads ove high particle or sedimal. 2004) which is generated by corals to i.e. remacher 1977). Even though resuspension in the area (Hubbard and Pocock 1972; Schuhmpact might be higher due to strong currents (Grant and Madsen 1979), mof high LAIW i

e reason. Stronger ents is unlikely for the samothering of corals by particles or sedimsmcurrents, however, might be responsible for high flushing rates of mucus from the coral
surface demanding quick mucus reproduction. Furthermore, the combination of high
tter fluxes (Patterson 1992; a 1997) and increased organic mantnutrient loads (SzmPalardy et al. 2005, Roder et al. accepted) might constitute a nutritionally replete
environmcus production as recycling is energetically not as uent and hence increase m

101

Chapter III

re oligotrophic surroundings (Ferrier-Pagès et al. 1998). In contrary opellent as in mcom

een the various to the DOM release, the ratio of released POC to PN did not change betw

ilar (Coffroth 1990) ucus was sim the fresh ments. This suggests that the quality oftreatm

pounds were not conserved in regardless of the corals’ origin and that nitrogenous com

ounts. stringent am

differ between the corals’ origin, and hence P. luteaption rates of Production and consumhistory of LAIW-exposure, even though environmental conditions during time of

mant 2002) and high ffects of high nutrient (Szment are equal. The interactive eeasurem

organic matter fluxes (Roder et al. accepted) as well as the low light (Kleypas et al.

1999), low pH (Levitus 1994) and low teme plus strong perature (Saxby et al. 2003) regim

ances such as tabolic performeura et al. 2003) on the mcurrent conditions (Nakam

terial release and uptake (Ferrier-Pagès et al. aphotosynthesis, calcification or organic m1998) seemely variable and need to be investigated in plex and extrem to be highly com greater detail. The findings of this study suggest that production of a coral is dependent

atic state gained by exposure to its natural surrounding and that its behavior on its someresents to certain extent conditions of its origin. Wnditions repeven under changed co

tabolic response. ee that the corals’ conditioning to LAIW has changed their mpresum

What remains to be shown really is whether this conditioning is acclimatization – or
rather adaptation.

ledgements Acknow

an bilateral ORCAS (Ocean-Reef e of the Thai-GermResearch was carried out in the fram

an Sea) project funded by the GermCoupling in the Andamarch Foundation an Rese

CT). The authors would like Thailand (NR(DFG) and the National Research Council of

to thank the Phuket Marine Biological Center and the National Park Mu Koh Sim

staff for field support.

ilan

102

Chapter III

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105

Leibniz Center for Tropical Marine Ecology, ZMT, Fahrenheitstrasse 6, 28359 Bremen,

1

Alfred Wegener Institute for Polar and Marine Research, Am Alten Hafen 26, 27568

Germany2 CORE Group, GeoBio Centre, LMU, Richard Wagnerstr 10, 80333 München 3Marine Research, Am Alten Hafen 26, 27568 Alfred Wegener Institute for Polar and

Bremerhaven, Germany4Phuket Marine Biological Center, 51 Sakdidet R

Phuket Marine Biological Center, 51 Sakdidet Road, 83000 Phuket, Thailand

& Claudio Richter

Carin Jantzen

Khokiattiwong

21,

, Gertraud M. Schmidt3, Cornelia Roder1, Christian Wild2, Somkiat
34 & Claudio RichterttiwongKhokia

In preparation

- Chapter IV -

Benthic primary production in response to Large

Amplitude Internal Waves (LAIW) in coral reefs

at the Similan Islands, Thailand

Chapter IV

106

Abstract

Chapter IV

The Andaman Sea features Large Amplitude Internal Waves (LAIW) of exceptional
amplitude (> 80 m), which may strongly affect environmental conditions within local coral
reefs. The Similan Islands in the swash zone of LAIW offer the unique chance to explore
ary production in response to these perturbations in all-scale variations in benthic primsmients (> 1.84 μM) and bient), total inorganic nutrperature (cold bouts up to 4 °C below amtemlight levels (~30 % below ambient). Here, we compare the relative cover and metabolism of
icrophytobenthos in reef sands, turf algae, scleractinian corals) ary producers (main primthe mat LAIW-exposed and non-exposed sites at Koh Miang, Similan Islands. The LAIW-exposed
lower live coral (28 %), and higher turf algae Western (W) side of Koh Miang coincided with eltered Eastern (E) reef (68 % and 8 % cover, respectively). pared to the sh(36 %) cover comnfluence, iWn by turf algae increased with rising LAINet photosynthesis and respiratiottern, despite entary net photosynthesis and respiration exhibited the opposite pawhereas sedimsimilar chlorophyll concentrations in algae and sediment samples from both sides. The
ilar net photosynthesis on both island showed simPorites luteainating reef-building coral domsides, but 40 % higher pigment concentrations at W, likely due to the lower light availability
inant primary producers on the ns. Turf algae were the domand higher nutrient concentratiocrophytobenthos on the sheltered E side, with corals i side of Koh Miang, and the mrocky Wary production on either side. In spite of the contributing less than 15 % to the gross primprofound effects of LAIW on the metabolism of the various main primary producers, the
overall primary production (i.e. microphytobenthos, turf algae and corals) was similar at both
side. -affected WWary production in LAIW and E, indicating high plasticity of prim

Introduction

Large Amplitude Internal Waves (LAIW) are ubiquitous in the ocean (Huthnance 1989, Apel
ed through the interaction of tides and the undersea landscape in a 2006). They are formdensity-stratified sea (Gerkema and Zimmerman 1995, Vlasenko and Hutter 2001). In the
Andaman Sea, LAIW of exceptional amplitude are created by tidal currents across the
shallow ridges near Sumatra, the Nicobar and Andaman Islands (Perry & Schimke 1965,
Osborne and Burch 1980, Jackson 2004). When LAIW reach shallow water depths they may

107

Chapter IV

ocline water upslope (Vlasenko & Stashchuk 2007). nd break delivering subtherm atransformLike pulsed upwellings in other coastal areas (Pineda 1991, Leichter et al. 2003, Leichter et

al. 1996, Smith et al. 2004), they may deliver essential nutrients to the surface supporting

primary production in otherwise nutrient-depleted waters. Arriving LAIW may be easily

y occur once or twice during dives. aidentified by divers as cold and turbid bores, which m

ilan Islands, located in the swash zone of LAIThe SimW (Vlasenko & Stashchuk 2007)

feature dramatic differences in their benthic communities between the W and E sides: on the

Wsheltered E sides where LAI influence is weak, a typical tropical coral reef usually occurs.

By contrast, on the W sides, where LAIW are much more pronounced, scattered corals

ework is e sand floor, but a carbonate reef framencrust huge granite boulders or anchor on th

lacking. This strikingly contrasting natural setting, only a few hundred mters apart, offers the e

unique opportunity to investigate coral reef primary production in dependence of pronounced y affect aena on the W side mental conditions. Two opposed phenomvariations in environmy aary production simultaneously: on the one hand the lowered light levels, which mprim

y a the higher nutrient availability, which mdecrease photosynthesis, and on the other hand ent concentrations, especially in deeperenhance photosynthesis, e.g. via the increase of pigm

lesias-P 1990, Igwater (Dubinsky et al.and Trench 1994). Additionally, the higher rieto

temperature variability, due to short term temperature drops induced by incoming LAIW on

the W side, may favour a different benthic community, adapted to these unstable conditions.
The reef sands as biocatalytical filter systems (Wild et al. 2005, Rasheed et al. 2006) may be
nced load of organic c nutrients and an enhaaffected by elevated concentrations of inorgani

material in the water column. This may strongly affect sedimentary primary production,

which can importantly contribute to the high pro s (Bunt et al.ity of coral reef ecosystemductiv

ary producers within 1972, Clavier and Garrigue 1999). Turf algae are likewise important prim

ed inorganic nutrient ter 1985), and the enhanccoral reefs (Rogers and Salesky 1981; Carpen

availability and concomitant lower light levels caused by the LAIW on the W sides, may

affect their growth and photosynthetic output (Williams and Carpenter 1988, Smith et al.

ith et al. 2004). On the other hand, corals are usually adapted to oligotrophic2001, Sm

environments and may have disadvantages under higher nutrient levels compared to algae
(Hughes 1994, Lapointe 1997, Pandolfi et al. 2005).paratively evaluate O of this study therefore was to comThe aimfluxes (photosynthesis and 2crophytobenthos, turf algae ientary mary producers (sedimrespiration) induced by key prims and protected sites and how these organism-exposed Wand scleractinian corals) from LAI

108

Chapter IV

contribute to overall primary production budgets were calculated ary production. Thereby primbining respective benthic cover and oxygen fluxes. com

Material and methods

Description of study site and background parameters Samplings and measurements were carried out at Similan Island # 4 (Koh Miang) on the E (8°
33.593’ N, 97° 38.237’ E) and on the W (8° 34.254’ N, 97° 37.957’ E) side, in January-March
deep: 19-21 m), in the following referred to as ;2008, in two water depths (shallow: 6-9 m

sampling ‘sites’ (E shallow, E deep, W shallow, W deep). Large Amplitude Internal Waves
(LAIW) occur in the Andaman Sea almost throughout the year (despite August and
February until April at the ost pronounced fromber, Jackson 2004), but they are mSeptemidt et al. subm.). ilan Islands (SchmSim -situ at all four sites usingperature (°C) were recorded inLight intensity (lx) and water tem) and light loggers (0 - 320000 lx, Pendant, Onset) both perature (C°, TidbiT v2, Onsettemwith a temporal resolution of 1 minute. Light was calculated as a daily mean around noon (11-
14 h, 02.02.-16.03.2008). As the spectral properties of the pendant loggers are unknown, the tosynthetic active radiation (PAR [μmol quanta ation of light intensity (lx) into photransformm-2 s-1]) equivalents can only be considered as a coarse approximation. Light intensity (I [lx])
easured during 4 shallow and 2 deep diurnal cycles in-situ with light loggers (lx, seewas malso above), and PAR (μmol quanta m-2 s-1) was recorded concomitantly with the internal
Walz GmbH, Gerlight sensor of a Diving PAM (submmany). In spite of the mersible fluoromeeter, pulse amntioned limitations, we found a reasonable plitude modulated, Heinz
correlation (I (lx) = 76.684  PAR (μmol quanta m-2 s-1), R² = 0.52, Fig. 1 suppl. data)
between the two measuring devices. Such PAR (μmol quanta m-2 s-1) and light intensity (lx)

ental conditions during correlations are widely used particularly in monitoring environmriculture of fish. amAutonomous upward-looking Acoustic Doppler Current Profilers (ADCP, RDI Teledyne
at E and )Workhorse Sentinel, 600 kHz and 300 kHz) were deployed at the deep sites (20 mW. They measured the 3-D currents above the tranducer at 1 m vertical and 1 min temporal
resolution with an accuracy of 0.3 to 0.5 % of the water velocity ± 0.3 to 0.5 cm/s. Data were
bia, h Pawlowicz, U. of British Colum by Ricported into Matlab (rdradcp.mim

109

Chapter IV

de were sihttp://www2.ocgy.ubc.ca/~rich/) and average current velocities for the E and W

lated. calcu

Water sam, nitrate plus nitrite, and phosphate) monium(amples for inorganic nutrient analysis

pling period using SCUBA (in total: E shallow, n = 11; E deep, were collected during the sam

n = 12; W shallow n = 8; W deep n = 9 samples). Water samples were filtered (GF/F filters, Ø

25 mm, nominal pore size: 0.7 μm), and the filtrate was conserved with 100 ml HgCl2

(Kattner 1999).

For comparison, additional temperature loggers (TidpiT v2, Onset, triggered every 6 min)

ilan Islands (# 2, # 7, # 8 south and # 8 north), each at E and e other Simwere deployed at som

perature characteristics were e consistent temW and in the two water depths. There, the sam

found as at Koh Miang (same period 02.02.-16.03.2008), revealing ‘averaged daily e tim

temperature ranges’ (DTRs) displayed as mean of daily max – min, as a measure for LAIW

pact at each site (for Koh Miang: Table 1, all: Table 1 suppl. data). This revealed increasing im

LAIW impact from E shallow, E deep, W shallow to W deep; therefore, sampling sites are

anuscript. ughout this m influence throWaligned according to rising LAI

ore, at Koh Miang nutrients, pH, oxygen, salinity, and current velocities correlated Furtherm

with temperature at each site (Schmidt et al. subm.). Schmidt et al. (subm.) also showed that

temperature variations of similar magnitude recorded at W-exposed faces of the above

Nov 2008) were consistently higher than ntioned islands over longer period (Feb 2007 toem

the variations recorded on the respective E island faces. This suggests that LAIW are

pacting more or less equally on all W island slopes and are actually causing the recorded im

W-E differences in environmental regime. Similar W-E differences were found in coral

ental an.). Both environmidt et al. submcommunities (Schmd biological data suggest that the

W and E faces of Koh Miang, an island in the middle of the Similans (Schmidt et al. subm.),

can be considered representative of a LAIW-beaten and LAIW-sheltered tropical island,

respectively.

Benthic community composition analyses

length, Hodgson et al. 2004), were conducted along Line point intercept transects (50 m

. Two to three ) and 20 m sides of the island, at two depths (7misobaths on the E and W

m apart. Within each transect, the benthic replicate transects were carried out spaced 10

d every 0.5 m. Hard substrate (live and dead coral cover and rock), plecommunity was sam

m ent and turf algae cover (on all substrates and on hard substrate), and coral growth forsedim

ined. ctors, see below) were determ(used for the 2D to 3D fa

110

Chapter IV

Sampling and preparation of investigated primary producers To investigate photosynthesis related parameters and sediment composition, mini-corers to
obtain 1.2 cm long, 6.4 ml volume and 3.9 cm² surface area samples were used. Only the top
thetically active, as light can only penetrate ent was considered as photosynlayer of the sedimsed within one h after ples were procesThe sam into the sediment (Kühl et al. 1994). a few mments (E shallow, n = 15; E deep, n = 15; W mcollection and used either for incubation experiminations of chl-a and pheophytin ent deter below), pigmshallow, n = 14, W deep, n = 20, seecontent (n = 10 for each site) or to analyse sediment grain sizes (12 samples pooled for each
site) as described below.Additionally, small-cores of 5.6 cm sediment depth (33.4 ml) were taken for pore water
nutrient analysis (n = 4 for each site). Within one h after sampling, the small cores were
h distilled water, centrifuged and then the supernatant was filtered througlwashed with 10 mpre-combusted GF/F filters; this procedure was repeated two more times and 100 μl HgCl
were added to the final filtrate for conservation (Kattner 1999). Filters were frozen for later total nitrogen (PN).ination of particulate organic carbon (POC) and particulate determentous algae inutive and filamerate of various dimglomTurf algae were defined as a congrowing up to a height of about 1 cm. Turf algae occasionally appeared on sediment, but only
the ones on hard substrate were used for turf algae analysis. Small pieces of coral rubble and
oved using a scalpel and an rock were sampled and processed within one hour; algae were remae-seawater suspension was used for incubations airbrush filled with filtered seawater. The algent analysis (E shallow, n = 5; E deep, n frozen for pigmand later filtered on GF/F Filters and sing the best fittinglated u deep, n = 15). Surface area was calcu= 15; W shallow, n = 10; W(s) of each scraped rubble piece. Only the light-exposed surfaces of the metrical forgeomgae growth rates, microscope slides (later ine alents were considered. To determrubble fragm on holders and deployed at each site. Algae tiles were referred to as algae tiles) were fixedsampled and displaced in a random order after 1-3 weeks (E shallow, n = 15; E deep, n = 15;
ed deep, n = 17); then they were scraped with a scalpel and airbrushW shallow, n = 10; Wwith filtered seawater. The algae-seawater suspension was treated as above. Daily growth ination below) per algae tile ent determhl-a (μg, see pigmount of crates were calculated as amarea (cm-2) and unit time (d-1).
, the most common coral genus at Koh Porites luteassive stony coral aFive colonies of the mMiang, and one of the most abundant species at the Similan Islands (Schmidt et al. subm.),
were sampled at each site, and two fragments were chiselled from each colony; fragments
eal for two weeks. One replicate of each colony riate site on racks to hwere left at the approp

111

Chapter IV

ber nts for each chameincubations, five fragmwas used for light and the other one for dark ann et al. (2009) with the lated according to Naumrfaces were calcut or dark). Coral su(lighbest fitting geometric shape (half-ellipsoid). Tissue was removed with an airbrush and filtered
ogenized with a tissue grinder. A subsample was filtered on GF/F filters d homseawater, anent analysis. mgiand frozen for later p

Measurement of oxygen fluxes by main primary producers Sedimentary microphytobenthos and turf algae were incubated in Winkler bottles (~ 60 ml)
incubations (1-2 h) were conducted with natural sea water in a water bath on land. Short-termaround noon to obtain maximum photosynthetic rates. Five independent light and dark
ae, were conducted simultaneously for each site. Varying ent or alger sedim eithincubations, ol 13 (μm 16 and 39 ±layers of net cloth were used to adjust, on a daily basis, PAR to 105 ±quanta m-2 s-1), corresponding to the average ambient E light levels at 7 m and 20 m,
respectively. Light and temof a Diving PAM and temperature loggers (see above), respectively. Incubation temperature were recorded every minute by the corresponding sensor perature
was 31 ± 1 °C, 1 °C higher than the maximum temperature at E shallow in-situ. This lack of
temperature control may be a weakness of this study. However, the same conditions during
the incubations yielded in overainterferences caused by the relativll valid primely unpredictable, brief temperature drops in-situ (Schmary production rates independent from short-termidt
. .)et al. subm ined before and after each incubation using an optodeOxygen concentrations were determ(HQ 40, Hach Lange). Repeated incubations with seawater only (i.e. controls) in light and dark (each n = 10), at simulated shallow water light conditions revealed no measurable
after 2h and were therefore not considered in differences between start and end oxygen values ter was used for coral incubations on four eersible flow-respiromthe calculations. A submpendant logoccasions (clear skies),gers (see above). The respirom one at each site in-situ. Light was meter consisted of three incubation chameasured concomitantly with bers - light,
tisensor-probe (Seabird SBE 19plusV2 with ludark and light control (each 2.1 L) -, a CTD mable control bing between valves and a programmp), pluminal 5T pumoxygen sensor and termunit. The unit was programmed to perform a measuring cycle as follows: (1) simultaneous
flushing of chambers with ambient water, (2) consecutive measurements of initial temperature
and oxygen concentrations in each of the chambers (measurements of each chamber took 40 s,
chambers were always analysed in the same order), (3) incubations (20 min) with intermittent
consecutive mstirring of chameasuremebers with external pumnt of final temps (Reich submersible pumperature and oxygen concentrations in each of the p 511-0110), and (4)

112

Chapter IV

chambers (intervals as initial values). This cycle lasted 30 min and was repeated 10 – 15 times
during the course of a day. The corals were likely not exposed to variations in water bers (2.1 l) likely prevented e of the champerature during incubations in-situ, as the volumtemuence of the enclosed water. Therefore, the to have a considerably infl LAIWshort-termilar to those ely stable conditions, simeasured oxygen fluxes were obtained under relativment and turf algae incubations. Gross photosynthesis for each site was calculated during sedimas: net production of each cycle + respiration mean for each dark chamber. A 2nd order
ed the best fit to the data (y = ax² + bx + c). The new incubation ial curve showpolynomously during ents autonoml incubation measuremdevice offered the possibility to obtain severaand after handling, as well as the deploying of the course of a day in-situ, however the before the quite huge device in the field is still very time consuming and requested a team of at least
could be obtained during this study.four people. Therefore only few runs

Sample analysesetrically and pore water) were determined photomn water columNutrient concentrations (fromafter Grasshof et al. (1999) with an autoanalyser (Evolution III, Alliance Instruments, France).
Nutrient pore water results were related to the water volume of small sediment cores using the
e values of pore water ent at each site (see below). Two extremwater content of the sedimammonium were removed from the data set due to an outlier test (criteria of multiple SD).
For pigment concentration measurements, samples (either filters or sediment) were unfreezed,
added with a certain volume of acetone (90%), shaken thoroughly, then placed in an
ards for 24h at 4°C in the dark to extract n and stored afterwiultrasonic bath for 15 mples were shaken thoroughly again and then centrifuged at high ents. Accordingly sampigmspeed for 15 min. The supernatant was measured in a photometer (Shimadzu UV 1700, 1nm
nd pheophytin was calculated using the equations ent and turf algae chl-a aslit). For sedim of corals the equations of Jeffrey and nt calculationseafter Lorenzen (1967). For pigmHumphrey (1975) were used. Pigment data were normalized by sediment area sampled,
rface area. coral surbstrate) ogrowth area of turf algae (on hard suthod (to eent characteristics grain size fraction was analysed using the pipette mFor sedimseparate the fine fraction) and wet sieving (to separate the coarse fraction; both Tucker 1988); terwards weighted and the proportion of weightthe different grain size fractions were dried, afined using a Schleicher-Apparatus and precipitated calculated. Carbonate content was determlime as a standard (Schlichting et al. 1995). Water content was estimated by weighing wet and
dry sediment to recompute pore water nutrient concentrations of the small-cores (see above).
s as the ent revealed pennate and centric diatomMicroscopic investigations on untreated sedim

113

Chapter IV

ent characteristics are ents. The visual sedimcro-algae in all sedimiby far dominating much lower bio-turbation on the W side. icates m also ind Fig. 2 suppl. data, whichillustrated in and dried at 40 °C. HCl (1N, 100-200 μl), Filters for POC and PN analysis were unfreezedwas added to one half of each filter, in order to remove inorganic carbon, and dried again.
ental Analyzer (NA2100 Protein) POC and PN content was determined using an Elemental CHNS standard (LECO). calibrated against an elem

Primary production budget calculations net and Pgross), as well as respiration (R) of ary production (PThe daily net and gross primturf algae were calculated using the results of the short crophytobenthos and ientary msedimterm incubations (maximum rates around noon, see above) and the start and end time points
ndial curves (see above) order polynomof photosynthetic activity by corals obtained via the 2trapolations. Pgross = Pnet + R, for each incubation, all to conduct likewise quadratic exvalues in (μg cm-2 h-1). For investigated primary producers, integral area of the daily curves
was computed using Mathematica 5.2 (-1Wolfram Research, Inc. 2005), resulting in an actual
ary production and respiration per ). To gain daily primary production (dnet and gross primreef and benthic community area, respectively, the daily actual rates were then multiplied with
the relative proportion of each primary producer at each site; then a 2D to 3D conversion for
turf algae, i.e. the surface area on which the turf algae grew, and corals, i.e. coral surface area,
asured on eined for turf algae by relating the area m. This factor was determwas appliedphotos (using ImageJ, 2-D area) to the area measured via geometry (see above, 3-D); actual
in-situ ace = 1.5. For corals therock surface colonized by turf algae/projected benthic surfnd Vogt (1997) were used. A coral 2D to 3D factor for each conversion factors after Alcala abranching, (ean of all factors for each coral growth formated using the msite was estimmassive, encrusting or foliose), weighted by the percentage cover of each coral growth form
found. Gross photosynthesis is given over the period of photosynthetic active radiation per dark incubations). iration for 24 h (calculated fromday (12 h, see also above) and resp

Statistical analyses sides using a two-mpared between E and Wre and water currents were coperatuLight, temeters, a two-(Ruxton 2006); for all other paramtailed T-test, adapted to unequal variances tailed U-test after Wilcoxon, Mann and Whitney was applied, either to compare E or W, or
eters, the non-ons of the DTRs with other paramshallow and deep sites. To test for correlatian correlation was used, setting the DTRs as ordinal ranks. pearmetric Sparam

114

Chapter IV

ResultsEnvironmental background conditions on E and W perature was significantly lower on W and in the deep in general (Table 1) At Koh Miang, tempared to E and shallow (both: t-Test, p << 0.001). Therefore, mean daily when comtemperature ranges (DTRs, daily max – min), as a measure for the impact of Large Amplitude
Internal Waves (LAIW), were less pronounced at E shallow and increased from E deep to W
shallow, to W deep (Table 1), where LAIW are most pronounced. Temperature loggers
th; each nd # 8 south and # 8 nordeployed at three other islands, at 16 sites (Islands (# 2, # 7 ailar variations (Table 1 suppl. data) with the highest shallow and deep) revealed sim,E and Wluding island # 4) yielded in DTRs of 0.9 ± 0.06 DTRs at island # 2. An all-site average (inc shallow, and 3.7 ± 0.14 °C at ± 0.28 °C at W deep, 2.19 °C at E shallow, 1.34 ± 0.13 °C at EW deep.Table 1: Environmvelocity, nutrient concentration; mental characteristics. Measured easured sediment paramwater parameters were eters were graitempn size fractionserature, light i, ncarbonate conttensity, current ent,
water content, nutrient concentration, POC, PN; all were determined at Koh Miang.
Parameter E shallow E deep W shallow W deep
Water Temperature (C°, modal) 28.84 28.59 28.82 28.77
Temperature (C°, mean±SE) 28.89 ± 0.0013 28.66 ± 0.0015 28.75 ± 0.0018 28.23 ± 0.0035
DTR, daily max-min (C°, mean±SE) 1.03 ± 0.07 1.5 ± 0.10 2.44 ± 0.16 3.94 ± 0.16
PAR 11:00-14:00h (μmol quanta m-2 s-1, 103.2 ± 0.48 36.2 ± 0.18 76.0 ± 0.59 23.7 ± 0.17
) ±SEmeanCurrent velocity (m s-1, mean±SE) 0.078 ± 0.0002 0.1068 ± 0.0003
Ammonium, water (μmol, mean±SE) 1.04 ± 0.094 1.19 ± 0.11 1.14 ± 0.12 1.45 ± 0.27
Nitrate + nitrite, water (μmol, mean±SE) 0.42 ± 0.062 0.57 ± 0.072 0.70 ± 0.14 1.19 ± 0.32
Phosphate, water (μmol, mean±SE) 0.22 ± 0.018 0.25 ± 0.013 0.26 ± 0.0092 0.37 ± 0.12
Sediment characteristics Clay and silt (%, pooled samples) 4.33 3.85 4.03 0.29
Fine sand (%, pooled samples) 25.25 26.5 1.57 16.23
Medium sand (%, pooled samples) 42.59 46.41 47.42 35.92
Coarse sand (%, pooled samples) 18.03 20.21 41.02 33.86
Carbonate content, (%, pooled samples) 86.7 83.4 72.6 78
Water content, (%, pooled samples) 35.2 34.7 38.7 37.9
Ammonium (pore water, mean±SE) 8.3 ± 2.47 11.09 ± 3.01 10.91 ± 1.49 16.15 ± 1.56
Nitrate + nitrite (pore water, mean±SE) 0.82 ± 0.24 2.77 ± 0.81 1.59 ± 0.53 7.84 ± 1.58
Phosphate (pore water, mean±SE) 3.18 ± 0.48 3.89 ± 0.53 3.35 ± 0.92 4.98 ± 0.72
POC content (μg mg-1 pore water, 2.10 ± 0.072 2.06 ± 0.075 2.33 ± 0.067 2.21 ± 0.068
) mean±SEPN content (μg mg-1 pore water, mean±SE) 0.13 ± 0.0071 0.057 ± 0.0096 0.18 ± 0.025 0.11 ± 0.017

28.77 .82 280035 28.23 ± 0.0018 28.75 ± 0.44 ± 0.16 2.94 ± 0.16 3.23.7 ± 0.17 76.0 ± 0.59

0003 1068 ± 0.0.1.45 ± 0.27 1.14 ± 0.12 1.19 ± 0.32 0.70 ± 0.14 0.37 ± 0.12 0.26 ± 0.0092

4.03 1.57 47.42 41.02 72.6 38.7 49 10.91 ± 1.1.59 ± 0.53 3.35 ± 0.92 2.33 ± 0.067 18 ± 0.025 0.

0.29 16.23 35.92 33.86 78 37.9 56 16.15 ± 1.7.84 ± 1.58 4.98 ± 0.72 2.21 ± 0.068 11 ± 0.017 0.

115

Chapter IV

in general (for each depth, t-Test, compared to Elower on WLight intensity was significantly : shallowp << 0.001) and revealed ~ 30 % lower values around noon (11:00-14:00 h, W light (Fig. 3 suppl. data) clearly deep: 34.5 %, Table 1). The diurnal cycles of%, W26.4 ore, a slightly shifted course of sun side. Furtherm the We forexhibited the lower light regimect of the island itself, could be observed. caused by a shading eff,light between E and WOverall current velocities were significantly higher on the W compared to the E side (t-Test, p
<< 0.001), reflecting an enhanced water exchange due to the incoming LAIW.
trite (nitrate + nitrite) and n, nitrate pooled together with niNutrients in the water columpact (DTRs, Table 1 and Table 2), but W imphosphate, showed an increase with LAIAmmonium on the other hand did not. Pore water nutrients yielded in a different result as again nitrate + nitrite, but also ammonium was correlated with LAIW impact (Table 1 and
Table 2), whereas phosphate was not.ounts within the Particulate organic carbon (POC) and total particulate nitrogen (PN) am-1 sediment), were significantly higher on the W side (both depths gent (Table 1, as μg msedimpooled together: U-test, p < 0.05) with an increase of 8 % and 43 % for POC and PN ely. ns, respectivconcentratioost no nts showed different characteristics for each site (Table 1). AlmemGrain size of the sedi) shallow sites (fewest on W deep and little fine sand for both Wclay and silt was found for Wcontrasting finer sediments on the E. Therefore, more coarse sand could be found on the W
sites, due to the elevated current regime. Accordingly water content within the sediments was
side (Table 1) sites than for E. Carbonate content was ~ 10% lower on the Whigher for the Wpared to E. com

8060cover(%)40

20

stshalloweapeestdeawestshallow
westdeep

8060cover(%)40

20

00sedimenthardsubtrateturflivecoral
Figure 1: Benthic cover. Line transects on benthic cover at Koh Miang (E shallow, E deep, W shallow, W deep)
in %; live coral cover as a fraction of hard substrate.

116

Chapter IV

Benthic community compositionA summary of the results obtained by the line-intercept transects is given in Figure 1. Sediment cover was highest at E shallow due to extensive sandy patches, medium at the deep
d rock) on ral cover anver, dead cowest at W shallow. Hard substrate (live coral cosites and lothe W side was mainly determined by the presence of rocks. Accordingly, the highest rock
shallow, few in W deep and 0 % at both E sites. W cover (granite boulders) was found on Wuced coral cover as sites showed in general enhanced turf algae cover on any substrate and reda fraction of hard substrate compared to E sites. The main primary producers, according to
percent cover, were the microphytobenthos (the sediment associated micro-algae), turf algae
and corals.

Growth rate of turf algae and pigment content of primary producers
side when Algae growth rates as chl-a increase, displayed in Figure 2, were higher on the Wpared to E (U-test, both depths pooled together per side: p > 0.0001). Higher growth rates commostly occurred in deeper water, nevertheless highest individual values were found for two algae tiles on W shallow (up to 0.025 μg Chl a cm-2 d-1). Growth rates were highly correlated
ably ound on E deep (and not E shallow, presumpact despite lowest values fwith LAIW imevoked by higher light levels).

120.100.-10.08 d-2060.Chl a (μg) cm040.020.

000.

shallowdeep

eastwest

Figure 2: Growth rates of turf algae. Microscopic slides (algae tiles) were deployed in-situ at all investigation
sites at Koh Miang and displaced in random order after 1-3 weeks (E shallow, n:15; E -2deep, -1n:15; W shallow,
n:10; W deep, n:17); algal growth was determined as chl-a (μg) cm d.

117

Chapter IV

The sedimpheophytin was coupled to LAIent showed simW imilar chl-a values for alpact (Fig. 3 right panel, Table l sites (Fig. 3 left panel, U-test, p>> 0.05), but 2) and was therefore
side (U-test, both depths pooled together per side: p < 0.02). A significantly higher on the Wilar pattern was observed for the turf algae on hard substrate; with no difference in chl-a simcontent for either site (Fig. 4 left panel, U-test, p >> 0.05), an increase of pheophytin with rising LAIW impact (Fig. 4 right panel,, Table 2) and accordingly significant more
side in general (U-test, both depths pooled together per side: p < 0.02).pheophytin on the W

shallowdeep

51.2shallow1.02deep420.830.6Chla(g)cmPheophytin(g)cm20.410.200.0eastwesteastwest
Figure 3: Pigment content of sediment. Chl-a and Pheophytin content of sediment from all investigation sites at
Koh Miang, normalised to sediment surface area (μg cm-2).

shallowdeep

]1]10.41.6shallow1.4bstrate)deep1.02te)substrardha
0.31.2hardsu0.20.820.6[g(cm0.40.10.0eastwesteastwest0.0mg(c[inhytPheop
0.2ChlaFigure 4: Pigment content Koh Miang, normof turf algae. Chl-a andalised to growth aPheophytin rea [μg (ccontent of m² hard substrate)turf algae from-1]. all investigation sites at

Coral pigments as chl-a rose with increasing LAIW impact (Table 2) and exhibited lowest
deep and W shallow and highest values for W values for E values for E shallow, mediumer per side, U-test, (both depth pooled togethdeep (Fig. 5). Chl-a content was higher on Wp pared 0.02) and at deep sites (both sides pooled together per depth: U-test, p< 0.02), com< to E and shallow sites.

118

Chapter IV

10shalloweepd82cm6Chlag420eastwest
Figure 5: Pigment content normof corals. Chl-a alised to coral scontent of uP.rface luartea ea (μfromg cm all inve-2).stigation sites at Koh Miang,

Table 2: Correlations to daily temperature ranges (DTRs). Various measures (see also results) were correlated to
at Koh Miang. DTRs Daily temperature range (DTR) versus level of p Rsp
significance Nitrate + nitrite water (μmol) **0.003 0.453
Phophate water (μmol) (*) 0.068 0.295
Ammonium water (μmol) 0.454 0.122
Nitrate + nitrite pore water (μmol) **0.001 0.759
Phophate pore water (μmol) 0.113 0.412
Ammonium pore water (μmol) *0.02 0.613
Algae tiles growth rate (Chl a cm-2 d-1) ** <0.001 0.698
Sediment chl a (μg cm-2) 0.115 -0.257
Sediment pheophytin (μg cm-2) ** 0.006 0.457
Algae chl a (μg cm-2) 0.73 -0.051
Algae pheophytin (μg cm-2) * 0.013 0.368
Coralchl a (μg cm-2) ** <0.001 0.657
Sediment net photosynthesis (O2 [μg cm-2 min-1]) ** <0.001 -0.456
Sediment gross photosythesis (O2 [μg cm-2 min-1]) ** <0.001 -0.471
Sediment respiration (O2 [μg cm-2 min-1]) 0.198 0.165
Algaenet photosynthesis (O2 [μg cm-2 min-1]) (*) 0.078 0.275
Algae gross photosythesis (O2 [μg cm-2 min-1]]) 0.596 -0.084
Algae respiration (O2 [μg cm-2 min-1]) ** 0.002 0.444
Corals (max. rate) net photosynthesis (O2 [μg cm-2 min-1]) 0.379 0.211
Corals (max. rate) gross photosythesis (O2 [μg cm-2 min-1]) 0.97 0.01
Corals (max. rate) respiration (O2 [μg cm-2 min-1]) ** <0.001 0.528
Sediment net photosynthesis (O2 [μg cm-2 min-1] Chla [μg]-1) * 0.015 -0.307
Sediment gross photosythesis (O2 [μg cm-2 min-1] Chla [μg]-1) * 0.022 -0.289
Sediment respiration (O2 [μg cm-2 min-1] Chla [μg]-1) 0.763 0.033
Algaenet photosynthesis (O2 [μg cm-2 min-1] Chla [μg]-1) 0.11 0.242
Algae gross photosythesis (O2 [μg cm-2 min-1] Chla [μg]-1) 0.343 -0.146
Algae respiration (O2 [μg cm-2 min-1] Chla [μg]-1) ** 0.001 0.466
Corals (max. rate) net photosynthesis (O2 [μg cm-2 min-1] Chla [μg]-1) ** <0.001 -0.792
Corals (max. rate) gross Photosythesis (O2 [μg cm-2 min-1] Chla [μg]-1) ** <0.002 -0.745
Corals (max. rate) respiration (O2 [μg cm-2 min-1] Chla [μg]-1) ** <0.003 0.861

)

119

Chapter IV

Primary production photosynthetic rates in the shallow, which crophytobenthos showed higher net ientary mSedimpared to deep (Fig. 6 left panel). mpanied by higher respiration rates comwere accoed on the W side in general (Fig. 6 left pabel, U-test, both Nevertheless, respiration was reducdepth pooled together per side, p < 0.02). Net and gross photosynthetic rates as well as both -1), were negatively correlated with ent content (chl-aphotosynthetic rates related to pigmLAIW impact (Table 2).
e coupling of net photosynthetic and respiration rates Turf algae, by contrast, showed a positivto LAIW impact (Fig. 6 right panel), also when relating both rates to pigment content (chl-a-1,
W side (U-test, both ore, respiration rates were significantly higher on the Table 2). Furthermwater depths pooled per side: p = 0.015). Gross photosynthetic rates were constant, impact. Wirrespective of side, depth or LAIore productive than the ones on the respective E Corals on the shallow W site were slightly mrence, in spite of lower light intensities on the side, whereas in deeper water there was no diffeilar, except for E shallow with twice the value W side (Fig. 7). Respiration rates were simspecific net photosynthetic rates were ig. 4 suppl. data). Chl-apared to the other sites (Fcom much lower for the W side (either depth, Fig. 5 suppl. data), reflecting the higher chl-acontent. The photosynthetic rates of corals around noon (net and gross, 11 - 14 h) related to pigment content (chl-a-1) were negative coupled with the LAIW impact, suggesting a more
side (Table 2). costly photosynthesis on the W

eastshallow
eedeastpstshallowewepstdeew

1eastshallow1
peedeast1]min0.2westshallow0.21in]m
epstdeew0.1algaecover)0.1sedimentsurface)0.020.00.10.1mg(c[2
2[g(cm2lightdarklightdarklightdarklightdark
OO

Figure 6: Oxygen fluxes of sediment and turf algae. Sediment and turf algae were incubated in a water bath on
land to determine oxygen fluxes (net photosynthesis and respiration); value-2s were -1normalised to surface area of
sediment or turf algae growth substrate (μg cm min).

120

Chapter IV

Primary production budgets ary production rates, i.e. rates attributed to photosynthetic active Actual net and gross primsurface area d-1, showed highest values for the sedimentary microphytobenthos, followed by
w productivity (Table 3). Theturf algae, whereas corals only achieved a relatively losediments showed by far the highest photosynthetic activity on E and W shallow, with almost
than algae or corals at any site. Taking into account the 50 % higher photosynthetic rates conversion factors (2D to 3D) for each site, the portions of benthic cover and the different prosediments were still the major primary producers on the E side, especially in the shallow
(Table 3). At the W sites, on contrast, the turf algae were the major primary producers, only
then followed by the sediments (Table 3). Corals contributed a relatively small amount to the
primary production on W deep, but a rather considerable quantity on all other sites.

tneossgr0.03

020.-1-1ni] m0.01
0.00 coral surface)030.2mc ( [μg20.02
O010.

sae shallowt

ep deaste

allowhwest s0.03

0.02-1ni] m0.01-10.00 coral surface)west deep0.0320.02mcg ( [μ2
O0.01

000.0.00681012141618681012141618
e (of the day)timtime (of the day)

Figure 7at each site at : Oxygen flKoh Miauxesng to of codetermrals. Incine oxyubationsgen fl of uxesP. lu, i.e.tea net and in-situ groswith an s photautomosynthesis, ated respiromvalues were eter during normalised one day
to surface area (μg cm-2).

121

Discussion

Chapter IV

nutrient input, and budget W the LAIostly fromThis study revealed that turf algae benefited m

calculations showed that these organisms were also the quantitatively most important primary

side. However, although growth, net photosynthesis and producers on the exposed W

respiration rates of turf algae were higher on the LAIW-impacted side, the sedimentary

microphytobenthos experienced reduced overall productivity on the W side. LAIW impact did

obviously not affect priment into ary production rates of corals, but supported investm

e content. yll a tissurophentation as reflected by the higher chlopigm

Primary production rates fectively croalgae are capable to efient associated mEven though it is reported that the sedimn (Miyajimr colum pore water and the watetake up nutrients froma et al. 2001), no fertilising

ientary mary production of the sedimeffect on the primcrophytobenthos could be observed at

side. Indeed, net and gross photosynthetic rates were reduced - and correspondingly the W

respiration showed lower values - when compared to the E side, despite the fact that similar

chl-a contents were measured in sediments collected from both sides. This generally reduced

sedimentary activity on the W side may be crucial for the overall benthic metabolism as the

ents are usually places of high productivity and important places for eable reef sedimperm

ild et al. 2005). Higher concentrations of recycling (Hatcher 1990, Rasheed et al. 2004, W

organic material and inorganic nutrients (ammonium and nitrate + nitrite) in sedimentary pore

water samples from the W side may be due to a higher particle load in the water column,

y be further enhanced by adetectable as an increased turbidity (i.e. reduced light levels), and m

gher abundances under enhanced nutrient levels i which can reach hanobacteria,N-fixing cy

ough there were no onstrated that althitted) demoder et al. (subm(Miller et al. 1999). R

tter (POM) aenhanced plankton abundances and suspended particulate organic m

concentrations in waters at the W side, the elevated water currents contributed to a 30 %

red to E. pa com ecosystemhigher plankton and POM supply to the W

122

Chapter IV

Table 3: Primary production budget. A primary production budget was estimated using actual gross
photosynthetic rates (actual PP [μg oxygen cm-2 12h-1]) weighted by transect data (% cover) and conversion
factors (2D to 3D, sediment: none; algae: own, see material and methods; corals: Alcala and Vogt 1997).
affiliationcompartementActualPPnetActualPP3DfactorscoverPPgrossrespiration
(gO2cm2gross(gO2(%)(mgO2m2(mgO2m2
12h1)cm212h1)12h1)24h1)
EshallowSediment78.1133.8159.1791.41529.93
Algae17.872.01.516.6218.89483.16
Corals7.813.94.4718.9118.00134.06
EdeepSediment35.292.9130.8286.73427.87
Algae21.266.51.519.4278.07590.56
Corals7.011.52.9740.3137.16120.55
WshallowSediment68.0101.616.161.5819.82
Algae27.479.31.549.2753.291196.74
Corals10.615.73.2426.9133.2191.45
WdeepSediment29.470.3142.6299.85372.51
Algae31.066.51.550.3521.91595.14
Corals7.011.43.456.124.0221.59

ents at the the coarser grain size of the sedim reflected inThese higher water currents are alsotter aentary grain size usually correlates negatively with organic mW side. Although sedimxygen penetration (Rusch et al. 2000, Huettel eability and oely with perm positivcontent, and al. 2003, Rusch et al. 2006), incompletely bster 2001, Huettel et al. 2003, Rasheed ete& Wdegraded organic matter accumulated within the sediments, as evidenced by high
jor places for the recycling aents are meable (reef) sedimconcentrations of pheophytin. Permet al. 2006), and their efficiency relies on the aterial (Hansen et al. 1992, Rasheed of organic mmicrobial diversity and abundance, which in turn is determined by the sediment properties
and mineralogy (Wild et al. 2005, Wild et al. 2006). Therefore, the reduced sedimentary
metabolism on the W side may be partly explained by different sediment characteristics. The
lower carbonate content and the more coarse grain size may lead to a reduced surface area for
microbial colonisation (Wild et al. 2005, Wild et al. 2006). This may imply further
iofauna and the associated intensity of bioturbation eentary mconsequences for the sedim side (pers. observations, also visible (Schlacher 1991), which was less pronounced on the Wat Fig. 2 suppl. data).y ary producers within the benthic reef communitportant primTurf algae are likewise im y beaeir photosynthesis m(Borowitzka et al. 1978, Carpenter 1985, Hatcher 1990), and thilliams and Carpenter1988), but related studies are els (W nutrient levder elevatedenhanced unpact, gross m iWrare. Still, as net photosynthesis and respiration co-varied in response to LAIpp and McKinnon (1992) ained constant. A study by Klumphotosynthesis of turf algae rem

123

Chapter IV

ary productivity, possibly due ass and primrevealed an inverse relationship of turf algae biom the y apply here as the turf algae obviously benefit fromato self-shading. The same relation m caused nutrient input in terms of enhanced growth rates, further supported by a higher WLAI .observed cover, but they showed similar gross photosynthetic rates on E and Wte up to 80 % of total y constituaal turf is not uncommon in coral reefs and mA dense algited by nutrient pp and McKinnon 1992). The growth of these algae is limbenthic cover (Klum 1983, availability, especially inorganic nitrogen, but also by grazing (Hatcher and LarkumWilliam and Carpenter 1988, Smith et al.2001). Although grazers were present on both sides,
no apparent grazing traces could be observed on any of the algae tiles (E or W). Some of the
or were fading as indicated by their high y also have diedaturf algae on the W side mpheophytin content.For corals (here Porites lutea), the two environmental factors, reduced light intensities and
enhanced nutrient concentrations may have balanced each other, because the massive Porites
pared to specimens on the at the W side could sustain its photosynthetic rates when comluteaE. On W shallow, P. lutea may have benefited particularly from the elevated nutrient
s, as corals revealed the (slightly) highest sitieht intenely high ligavailability and relativde exhibited a higher siphotosynthesis rate compared to all other sites. Corals on the Wely facilitated by the enhanced nutrient levels (Muscatine et entation, likent into pigminvestmintain high productivity under reduced a to mal. 1989, Dubinsky et al. 1990) enabling themlight intensities., which suggests efficient use of available light intensities (Dubinsky et al. 1990, Titlyanov 1991, Iglesias-Prieto and Trench 1994). This implies a high cellular
investment by the zooxanthellae as increased chl-a contents are more likely caused by
elevated chl-a concentrations than by multiplication of zooxanthellae within the host tissue,
rted to stay relatively constant within the range of xanthellae abundances are repobecause zooe present study (see review by asured in thenutrient concentrations and light intensities milar zooxanthellae abundances in the tissue of Leletkin 2000). This is further supported by simwater depth (own unpublished data, two-tailed U-test, irrespective of side or P. luteap >> 0.05).ilar on all sites, which is ary production of the investigated corals were simNet and gross primary anov (1991), who found a stable level of corals’ primance with the study by Titlyin accordinly caused by reduced aproduction in a wide range of light intensities (i.e. water depths), mrespiration under lower light levels. Still, P. lutea’srespiration rates were relatively similar,
despite the ones at E shallow, where elevated respiration rates may reflect an enhanced
unds and Davies 1988, Hoogenboom et al. , due to a high-light history (Edmetabolismm

124

Chapter IV

ical energy 2006). Generally, a surplus of solar energy, which cannot be used for photochemconversion, is mturn requires enzymaatic reparation of photosysteminly dissipated as heat, but causes also reversible photodam II and consequently raises respiration rates age, which in
nd Davies 1988, Hoegh-Guldberg and Jones 1999, Gorbunov et al. 2001). unds a(Edm

Primary production budget A primary production budget per reef area for all sites was estimated by combining incubation
y a that no light respiration could be measured, which muld be notedwith transect data. It shobe considerably higher than the dark respiration (Titlyanov 1991, Anthony and Hoegh-ay explain ated and may be underestimary production mGuldberg 2003). Therefore, gross primwer than 1 for each site. P:R ratios loBoth sediments and algae exceeded the productivity of corals, which was mainly caused by a
orals contributed a higher absolute cover, but also by an elevated photosynthetic activity. Crather smary production, especially at W deep. This was supported by a ount to primall amore photosynthetically turf algae as mstudy of Rogers and Salesky (1981), which revealedproductive than Acropora palmata. The total gross primary production budgets for each site
at higher in the shallow – leading to the ewhilar results – although somyielded in simary production was relatively independent of the side, conclusion that the overall gross primbut the contribution of each compartment varied under the different environmental conditions.
ent and Although different incubation approaches were applied for corals in contrast to sedim other reef areas, however omturf algae, oxygen fluxes were consistent with published rates frat the lower end for turf algae and corals (sediment: Hansen et al. 1992, Clavier and Garrigue
s and Carpenter 1988, Carpenter et al. 1991, illiamild et al. 2005; turf algae: W1999, Wpp and McKinnon 1992; corals: Falkowski and Dubinsky 1981, Hatcher 1990, KlumTitlyanow 1991). Still, metabolic characteristics of P. lutea may not be valid for all
scleractinian coral species. According to Ralph et al. (2002), who investigated photosynthetic efficiency and productivity of six coral species, a massive species (Cyphastrea serialia) and
corals of the genus Porites (P. cylindrica) showed lowest values.

Ecological implications and outlook ed to be rather unfavourable for corals in contrast to are usually assumElevated nutrient levels oral- towards algae- cing phase shifts fromportant role duralgae and therefore play an imdominated reef ecosystems (Hughes 1994, Lapointe 1997, Pandolfi et al. 2005). Turf algae
were the dominant algae at Koh Miang and there is evidence that they may not be efficient

125

Chapter IV

competitors for corals,2003), particularly not for the m even under eutrophiassive growing genus Poritesc conditions (Mc Cook 1999, Jom (McCook 2001). However, it pa and McCook
must be considered that – as in most studies – the species composition of turf algae was not
determined and may change considerably over time or space. Therefore, on the one hand coral
cover was reduced on the W side and some coral species like Stylophora pistillata may suffer
under enhanced nutrient levels (Ferrier-Pages et al. 2000), but on the other hand, P. lutea may
also benefit from the elevated nutrient levels, maintaining high photosynthetic performance
under low light levels. ets to get ucus she colonies in-situ were regularly observed covered with mPorites luteaHuge sp. is known to be even Poritesentation, especially on the W side, and rid of the sedimabundant in areas with high particle load (Morelock et al. 1983, Cortes and Risk 1984). But entation of corals can cause a decrease in photosynthesis and respiration rates generally, sedimic features (Rogers et al. 1990, Philipp and tabole several other mas well as a decline ofphotosynthesis of the investigated organismFabricius 2003). In contrast, the higher water currents on the W side ms in-situ as indicated by the studies of Carpenter et ay increase
crophytobenthos, and ientary mal. (1991) for turf algae, Cook and Røy (2006) for sedimDennison and Barnes (1988) for corals. The Siecosystemm functioning and resilience in responsilan Islands as a study site offer a unique opportunity to investigate coral reefe to different environmental conditions. This
ateportant in order to understand potential consequences of global climis particularly imchange on the reef ecosystem level. An alteration in coral reef benthic community
ny reef locations world-wide (Hughes 1994, Bellwood et al. a mposition as reported fromcomplies changes in 2006, Hoegh-Guldberg et al. 2007, Hughes et al. 2007) also potentially im ary productivity. The present study showed that this does not have to be-wide primecosystem benthic primary production under different the case, as it revealed high plasticity ofary production, as ental factors. However, further investigations on benthic primenvironmfundaecosystemmes. nt of the trophic cascade, are needed, particularly with focus on changing coral reef

ledgements Acknowcean-Reef Coupling OCAS (an bilateral ORResearch was carried out as part of the Thai-Germ and funded by the German Research Foundation (DFG). an Sea) programin the AndamTobias Funke, Sopherl Schubert, on Koh Miang: Special thanks are due to the island teamon and Yvonne Sawall. Great thanks to Niphon Sinja Ernst, Laura Fillinger, Melanie B

126

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Phongsuwan and his two helpful assistants and the staff of the PMBC Phuket, who helped ry analysis to Stefanie Bröhl and thanks for the assistance in laboratoistics. Manywith the logMatthias Birkicht. We are very thankful to Florian Mayer for his assistance with the
atica. emlations in Mathcalcu

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129

Supplemental data Chapter IV

Chapter IV

001600014000120001000080Light intensity (lx)0060004000200050100150200
PAR (μmol quanta m-2s-1)
Supplementalfigure1: Light (lx) - PAR correlation. A light intensity (I, lx) to photosynthetic active radiation (PAR) correlation was
obtained by comparing pendant logge76.684 * PAR (μrs and the light sensor of the Diving PAM during 6 daymol quanta m-2 s-1).s (4 shallow and 2 deep); R²: 0.52, I (lx) =



Supplementalfigure2: Visual sediment characteristics. Exemplary photos of sediment surfaces at all investigation sites at Koh Miang
( ~ 30cm above bottom).

130

250200

150

100)-1 s-2500250PAR (μmol quanta m200150

100

east

west

shallowdeep

Chapter IV

250shalloweastdeep200150100)-1 s-2500250PAR (μmol quanta mwest200150100500time of the day (h)Supplementalfigure3:DiurnalcyclesofPAR.Thediurnalcyclesofambientlightintensity(lx)duringinsituincubationofP.
luteaateachoftheincubationdayswereconvertedintoPAR(photosyntheticactiveradiation[molquantam2s1])usingthe
.correlationfromFig.1suppl.data-0.016]-0.014-1 s-2-0.012 cm2-0.010-0.008-0.006 O [μgirationresp-0.002
-0.004east shalloweast deepwes tshallowwest deep
Supplementalfigure4:Oxygenfluxesofcorals.IncubationsofP.luteainsituwithanautomatedrespirometerduringone
dailycycleateachinvestigationsitetodetermineoxygenfluxes,i.e.meanrespirationrates.

50

131



east shalloweast deepwest shallowwest deep

Chapter IV

0.014-1]0.012east deepeast shallow
0.010a)thesis west shallowwest deep0.008 (μg Chl 0.006net photosyn20.004[μg O0.0020.000681012141618
(h)of the day time Supplementalfigure5:Oxygenfluxesofcoralsrelatedtochlacontent.IncubationsofP.luteainsituwithanewautomated
deviceduringonedailycycleateachinvestigationsitetodetermineoxygenfluxes,i.e.netphotosynthesis,valueswere
normalisedtochlacontent(gO2[gchla]1).
Supplementaltable1:Dailytemperaturevariationranges.DTRsasmax–min(mean±SE)atfouroftheSimilanIslands(#4,
#2,#7,#8south,#8north),eachatfoursites:Eshallow,Edeep,Wshallow,Wdeep;n.a.valueswerenotavailable.
Site Koh #4 Koh #2 Koh #7 Koh #8 Koh #8 Overall mean
northsouth max-min (°C)
E shallow 1.03 ± 0.07 0.75 ± 0.06 0.94 ± 0.04 0.88 ± 0.46 n.a. 0.9 ± 0.06
E deep 1.50 ± 0.10 1.61 ± 0.11 1.04 ± 0.07 n.a. 1.20 ± 0.07 1.34 ± 0.13
W shallow 2.44 ± 0.16 3.06 ± 0.18 2.18 ± 0.18 1.33 ± 0.11 1.95 ± 0.15 2.19 ± 0.28
W deep 3.94 ± 0.16 3.98 ± 0.17 3.84 ± 0.16 3.22 ± 0.18 3.52 ± 0.18 3.70 ± 0.14


132

& Claudio Richter

Somkiat Khokiattiwong

,

Gertraud M. Schmidt1, Niphon Phongsuwan2, Cornelia Roder3, Carin Jantzen3
Somkiat Khokiattiwong2 & Claudio Richter1

afen 26, 27568

Alfred Wegener Institute for Polar and Marine Research, Am Alten H

1

T, Fahrenheitstrasse 6, 28359 Bremen, M

Leibniz Center for Tropical Marine Ecology, Z

Bremerhaven, Germany 2Phuket Marine Biological Center, 51 Sakdidet R

23Leibniz Center for Tropical Marine Ecology, ZPhuket Marine Biological Center, 51 Sakdidet RMoad, 83000 Phuket, Thailand T, Fahrenheitstrasse 6, 28359 Bremen,

Germany

d to Marine Ecology Progress SeriesSubmitte

Internal Waves

characteristics in response to Large Amplitude

Coral community and physico-chemical

- Chapter V -

Chapter V

133

Abstract

Chapter V

The Similan Islands (Thailand), Andaman Sea, feature an unusual coral reef distribution: reefs

) sides cean-facing west (W) whereas the oflourish on the sheltered shelf sides east of the islands (E

ework. Here, we show that the striking differences in reef development, mlack a true reef fra

ong the islands and tres, are consistent ameoccurring at spatial scales of only tens to hundreds of m

related to Large Amplitude Internal Waves (LAIW). Two year temperature records show that LAIW

have their strongest impact on the deeper parts of the W Similans where they may cause frequent

(several events per hour) and abrupt (up to 10°C, in the order of mith nutes) drops in temperature wi

ical oceanographic onsoon (January through April). Physical and chempeak activity during the NE m

analyses show that LAIW advect deep cold, nutrient-rich, suboxic and low-pH waters (0.6 pH units

below ambient) into shallow near shore areas, and provide a dramatically altered growth

environment for W slope benthic communities. In contrast to E reefs, which are dominated by a low

num slopes harbour only loose, however more diverse e-building species, the Wmber of fra

odifiedent, often with me granite basemcommunities of scattered corals growing directly on th

ification). LAIWphenotypes (broadened bases, reduced ram, which are ubiquitous in SE Asia and

ical-biological disturbance operating ico-chem physbeyond, provide a so far understudied source of

s. In the pared to the well-known physical effects of stormporal scales comat different spatio-tem

light of the wide range and complex nature of environmental variability involved, LAIW may hold a

an era of global change. ce inclue to coral resilien

s rdoKey w

Large Amplitude Internal Waves - benthic-pelagic coupling - coral community - Similan Islands -

n Sea – solitons amAnda

Introduction

, nutrient poor tropical s thriving in clear, warmCoral reefs are highly productive benthic ecosystem

et al. 1979, Veron 2000, Birkeland 1997, Hoegh-eier & Kinzie 1976, Goreau waters (Buddem

ever, are subjected to natural disturbances which affect light, Guldberg 1999). Many reefs, how

134

Chapter V

alls, onsoon driven rainfperature, nutrient and aragonite saturation state on various scales, e.g. mtemeier & Kinzie 1976, Charuchinda & s, and upwelling of cold deep waters (Buddemxing by stormim2009). Tidally recurring internal waves have been Hylleberg 1984, Leichter et al. 1996, Lesser et al. shown to play a potentially important role in the cross-shore exchange of larvae, nutrients and
particulate food (Shanks 1983, Pineda 1991) and can be a source of nutrients for coral reefs (Leichter et al. 1998, 2003). However, Large Amplitude Internal Waves (LAIW) causing
temperature drops of up to 6 – 9 °C (Wolanski & Delesalle 1995, Sheppard 2009), could be a mixed
blessing, where the positive effects of an enhanced exchange of material may be offset by the
odic low temperature, low aragonite saturation state and high ulative negative effects of pericumnutrient content of the upwelled LAIW waters. The Andaman Sea features internal solitary waves of
Perry & Schimke 1965, Osborne & Burch 1980, Jackson 2004),plitude (> 80 mextraordinary am an-Nicobar generated by the waxing and waning of the tides across the shallow ridges of the AndamLAIWisland arc and the shallow reefs northwest of Sum travel as waves of depression in groups eastward across the deep Andamatra (Alpers et al. 1997, Jackson 2004). These an Sea at speeds of
2 m s-1. As the solitary waves propagate from the basin onto the shelf, interaction with the bottom
tion into waves of elevation with secondary waves of dispersion amleads to a gradual transforevolving from the trailing edge of the LAIW (Vlasenko & Stashchuk 2007). For intense LAIW with
amplitudes exceeding 402002), the steepening and overturning of the rear wave faces cause wave breaking an % of the thickness of the layer below the pycnocline, (Vlasenko & Hutter d generation of
upslope propagating density intrusions (Vlasenko & Hutter 2002, Vlasenko & Stashchuk 2007). with resuspension of sedimAlthough the passage of LAIWents up to 30 m leads to turbulent above bottommi (Moum et al. 2007), xing near the sea bed (De Silva et al. 1997), it is so far largely
The Simunknown to what extent they may affect coral coilan Islands are an offshore group of coral islands in the Andaman Sea located in the swmmunities. ash
zone of Andaman Sea LAIW (Jackson 2004, Vlasenko & Stashchuk 2007). In contrast to most other
reefs, where reef growth is most(Veron 2000, Spalding et al. 2001), satellite images (www.reefbase.org/gis_m vigorous on the exposed (or windward) face of a baaps) and mrrier or island onitoring
offshore islands in the Andamstudies (Chansang et al. 1999) suggest that reef developman Seaent along the Sim is restricted to the sheltered E sides (Fig. 1 B) (Phongsuwan et ilan Islands and other
al. 2008). The Woccur only in scattered colonies and sheltered areas along the coastline (Fig. 1 A). The reasons f sides, by contrast, appear to be conspicuously devoid of reef formations and corals or
eef distribution are so far unknown. Sea paradox of coral rnathe Andam

135

Chapter V

Figure 1: Typical substrate conditions at Similan Islands in about 12 m depth: A) at West W 4.1 and B) at East 7.1

Although hydro-dynamical forcing by surface gravity waves may be important for structuring coral

) monsoon (Dunne & Brown facing areas exposed to the south-west (SW- shallow Wcommunities in

1996, Wu & Zhang 1998), ocean swell and storms in the Andaman Sea are less severe than e.g. in

the windwardne tracks of the open tropical Pacific, where coral communities flourish othe cyclon

sides (Spalding et al. 2001). Here, we suggest that LAIW-induced upwelling undermines reef-

generating processes by the cumulative effects of low temperatures, low aragonite saturation state

and elevated nutrient concentrations of the upwelled water. Low temperatures have been shown to

limal. 1999), low aragonite saturation state reduces coral growth and ent (Kleypas et it reef developm

nt concentrations affect the delicate ez 2006), elevated nutriercalcification (Schneider & E

biotic zooxanthellae (Muscatine et al. 1989, Muller-Parker association of corals with their endosym

1994, Ferrier-Pages et. 2001), favour the growth of competitively superior phytoplankton and

marco & Risk 1990, amm et al. 2003, McCook 1999) and enhance bioerosion (Sacroalgae (Abram

subjected to intense smChazottes et al. 2002, Ward-Paige et al. 2005). Tropical shallow platfor

oderate wind-driven upwelling are thus devoid of coral reefs (Hallock & Schlager 1986) but m

and/or intermittent upwelling may allow for moderate to extensive reef development (Andrews &

Gentien 1982). The magnitude and potential role of upwelling near Andaman Sea coral reefs and

er slope communities is so far unknown. t uppadjacen

ental variability and coral abundance The present study explores the relationship between environm

and diversity in relation to LAIW. The objective was to examine the fringing reef communities and

nce of a strong ilan Islands which are under the influemical characteristics of the Sithe physico-chem

136

Chapter V

SW-monsoon regime and tidally recurring LAIW. We hypothesize that monsoon and LAIW
unities. This was tested by (1) W coral commdifferentially affect shallow and deep, E and ral cover patterns, (2) assessing W - E and ing coral community and coparquantifying and comical differences and (3) relating the characteristics of coral shallow - deep physico-chemental conditions. mcommunities to the existing environ

Methods

Study areaThe Similan Islands belong to the National Park “Mu Koh Similan” with a total area of 140 km2 in
east of the Andaman- west of the Thai coast and 400 kman Sea, Thailand, about 60 kmthe Andament over a distance of about 24 North-South alignmNicobar-Islands. The 9 islands are located in a between 8°40`54.49”N, 97°38`56.41”E and 8°28`28.45”N, 97°38´56.85”E (Fig. 2). The islands kmconsist of granite and have been built in the tertiary about 65 million years ago by volcanic activity
etry are r topography and surrounding bathyministration). Thei(Thai National Park adm depth along the western sides and a slope (> 45°) down to 20 mcharacterized by a generally steepbroad sandy beaches and shallow reefs. Under water e east with tler slope (< 40°) in thslightly genthe substrate in the west is shaped by granite boulders along the steep slope until about 16 m depth
h dept to gently drop down to 60 – 80 mbefore passing into sand and coral rubble and continuingostly restricted to the nd maround and between the islands (Fig. 2). Annual rainfall is 3560 mm aSouth-West monsoon between May and October with strong winds, occasional storms and high
inistration). waves (Thai National Park adm plingReef sam ) sides ofRepresentative study localities were established along the eastern (E) and western (W 9.1, Fig. 2). To ilan Islands (nine in the east: E 1.1 – E 9.1 and seven in the west: W 2.1 – WSim line transects were established at each mmunities, single 100 mE differences in coral co-explore W ned Transect sites were positioarked with steel stakes, one at either end.of the 16 localities and mhe s (Wolanski et al. 2005). T i.e. beyond the reach of storm,along isobaths at a depth of 12 – 16 mthod (Loya and Slobodkin 1971; Loya 1972, 1976) was adopted to quantify the eline-intercept mian corals, soft corals, ponent (i.e. scleractint of each benthic comcoral communities. The intercepetre and the e nearest centimwas recorded to thacroalgae, sand and rubble) under each line transect m

137

ber of colonies recorded. Scleractinian corals were idnum

Chapter V

entified to species level. To establish

length were itional triplicate transects of 50 mvertical differences in coral communities, add

established in 20 and 7 m at Koh Miang on both sides, west (W 4.1) and east (E 4.1). Transect lines

erever in these latter transects coral cover was h apart. Wwere positioned along isobaths spaced 10 m

ined every 5 m across the whole transect work was determedetected the height of its skeletal fram

distance. Therefore a measuring stick of known length was placed perpendicular to the isobath in

ework and a picture taken displaying the comfront of the coral framework with its plete fram

basement (sediment or rock) and the measuring stick. The height of the framework was calculated

from later picture analysis with the software ImageJ. A total of 42 framework measurements was

respectively, and nine and twelve each in 20 and 7 m E. 20 and 7 mcarried out, 15 and six in W

of differences extension of this work to all sites, but the lackLogistic constraints precluded the

differences between W and E sides of the islands ong and the consistency and significance ofam

justified to restrict the vertical analysis to this central Similan island (see results, below).

temperature; Figure 2() all : Simparamilan Islaetends rs at cewith ntral islbiological and eand Koh Miang nvironmental sam(W 4.1 and Eplings: ( 4.1). (Figure ) transects modified after only; () transects and Jackson (2004)).

138

ing plical samPhysico-chem

Chapter V

puters; - Temperature recorders (TidbiT v2, Onset comBroad-scale temperature measurements

2 °C within a range of 0 to 50 °C) were deployed in two depths (20 and 7 mtion 0.resolu at 5 study )

sites, both in W (W 2.1, W 4.1, W 7.1, W 8.1, W 8.3) and in the opposing E face (E 2.1, E 4.1, E

7.1, E 8.1, E 8.3). Loggers were attached about 20 cmperature above the substrate recording tem

values at 6 min intervals from February 2007 to November 2008.

Fine-scale temperature measurements - Additional temperature loggers were established in the reefs

nute from iat island Koh Miang (W 4.1 and E 4.1) in 5, 10, 15 and 20 m depth logging every m

ber 2007 through March 2008.Decem

Light-, current- and CTD-measurements – Temperature, salinity, pressure, oxygen, pH, chorophyll-

easured with a CTD (Seacat SBE 19plus, Sea-Birdce and optical backscatter was ma fluorescen

de (W 4.1) of island Koh Miang in March 2007 for si depth at the WElectronics) deployed in 20 m

four weeks. At the same time an Acoustic Doppler Current Profiler (600 kHz Workhorse Sentinel

ine the current field at xt to the CTD to determents) was placed right neADCP, Teledyne RD Instrum

2 m vertical (bins) and 1 min temporal resolution. Current speeds were determined by averaging the

3-D current measurements from three vertical bins for deep (20 m: 20 to 16 m depth) and shallow
(7 m: 10 to 6 m depth). Light loggers (Onset pendant light logger: 0 - 320000 lux [lm m-2], typical to

90%) were deployed on both sides of island Koh Miang (W 4.1, E 4.1) in two depths (20 and 7 m)

about 20 cm above the substrate recording light values every minute from December 2007 till April

s to interfere with light 2008. The loggers were cleaned daily to avoid fouling organismmeasurements. Additionally a photosynthetic active radiation (PAR, μmol quanta m-2 s-1) sensor

(Biospherical Instrum(SEACAT, SBE 19plus, Sea-Bird Electronics) in ounted on a CTD ents) was m

4.1) in 20 m depth. of island Koh Miang (W January 2008 over a period of 20 days on the W

SCUBA about 1 mples was collected by – A total of 40 water samChemical measurements above

-litre PE-bottles at island Koh Miang at both sides (W 4.1, E 4.1) in the reef substrate using sterile1

ples bruary and March 2007. Right after collection water sametwo depths (20 and 7 m) between F

were filtered through pre-combusted and pre-weighed glass-fibre filters (Whatman GF/F, 45 μm).

sis and in pre-trient analyThe filtrate was stored in sterile polypropylene bottles for further nu

ination of dissolved organic busted DOC-vials spiked with phosphoric acid for further determcom

carbon. The filters were kept for the determples atter. Water samination of suspended particulate m

rcuric chloride after Kattner (1999). e mysis were poisoned withfor dissolved inorganic nutrient anal

red on ice and frozen. Nutrient samples were stoAll samons et al. 1989 ples were analyzed after Pars

139

Chapter V

for nitrate, nitrite, ammonium, phosphate and silicate using a spectrometer, GBC model UV/VIS981
C) in Phuket. pler model FS3000 at Phuket Marine Biological Center (PMBwith an autosamDissolved organic carbon samples were analyzed by means of high temperature catalytic oxidation
inum catalyst. Beforean DC-190 Total Organic Carbon Analyzer equipped with a platusing a Dohrmples were decarbonated by purging with oxygen. The injection into the furnace, the acidified sam detection system. was purified, dried and detected by a non-dispersive infraredOevolving C2ghing the filters after drying them over night at ined by weiSuspended particulate matter was determparator, 1 μg accuracy). 50°C on a microbalance (Mettler, AT21 Com Data processing and statistical analysisetric analyses, as the data required, both biological and -paramBefore applying parametric or nonenvironmental data were tested for the assumption of normality and homogeneity of variances using
irnov and Levene´s tests, respectively. ogorov-SmmKolPRIMER v6 for non-parametric mulTransect data were processed as percentaged abundances per site and analysed with the software tivariate datasets (see Clarke 1993, Clarke & Warwick 1994).
rities were used to detect spatial differences ilautation tests based on Bray-Curtis simANOSIM permay-analysis with 25 in substrate cover and species compositions between island sides (one-witeration steps) as well as to clarify depth dependent cover and distribution patterns at the W and E site of island Koh Miang (two-way-crossed design implicating the differing impact of LAIW
lates a global R . ANOSIM calcu) versus E, and depth: 20 versus 7 mdepending on orientation: Wstatistic that reflects the differences in variability between groups as compared to within groups (so R values are proportional to differences between the groups) and checks for the significance of R was also used to further analyze benusing permutation tests (Clarke and Wathic commrwick 2001). Nonmunities. Based on a simetric multidimilarity maensional scaling (MDS)trix, MDS generates
ilarity (Clarke and points is proportional to their degree of simplots in which the distance between Warwick 2001). SIMPER analyses were consulted to assess the respective contributions of substratetypes and coral species to the similarities and dissimilarities within and among the W and E sites
studied. Descriptive coral community factors as Shannon index, evenness and species richness (Rogers 1993, van Woesik et al. 1999) were calculated for every site and tested for LAIW-exposure
versus E) using Student´s t-tests. (W ne-series. The calculatioDegree days cooling (DDC) was calculated for every temperature timtime-series (y); (2) spinvolved three steps: (1) calculation of mlitting of residuals into cooving modes (mld (ryc) and warmy) and residuals (ry) on the basis of the residuals (ryw); and (3) integrating

140

Chapter V

cold and warm anomalies into DDC and degree days warming (DDW). The moving modes were
calculated using a slide function (by Jos van der Geest; http://www.mathworks.com/matlabcentral/fileexchange/12550) to a moving 1-day window of
e-series y and smoothing the output by a 1-day moving average, yielding ry. ents of the timelemDegree days cooling was calculated by replacing the positive values in the residuals time series ry
ples per day ber of sam(ryc)], dividing by the numing the values [sumwith zeros, yielding ryc, summples per year, divided by the number of samples alizing to the full year with F {sam(spd), and normof the time series, i.e. F=spy  [sp(ry)]-1}. The corresponding equation is:
DDC=sum(ry)  (spd)-1  F (1)
sly without failure at all sites, the calculation of the g continuouBecause not all loggers were loggin all sites and depths (20 weeks, DDC was done for the period when data were available from February 2007 to July 2007). The resulting degree days cooling (DDC) displayed as (°Cd) were exposure. Their relationship with the descriptive coral Wused as site specific indicators of LAIses. community factors and living coral cover of every site was tested using linear-regression analy -exposureWNutrient concentrations, current and light recordings were statistically tested for LAIere allis ANOVA by ranks. Linear-regression analyses wetric Kruskal-Wusing non-parameters in the water as ical and physical paramundertaken to test the relationships between both chem(see Sokal & Rohlf 1995) e independent variable perature as ththe dependent variables with temicance. If not stated otherwise data are cal signifatistiine their stfollowed by Student´s t-test to examean ± SE. always displayed as m

Results

Reef data Coral community composition – Overall 144 hermatypic coral species were recorded, belonging to
ental Fig. 1). Ten genera could be ental Table 2, supplemilies (supplem40 genera and 17 famhalf of the number of species the island chain containing exactly ayor players along detected as the minant genus, especially in E, was the overall domPoritesfound at all sites studied (Table 1). followed by Acropora, Hydnophora and Pocillopora, all of them with comparatively larger
.ces in Wappearan

141

Chapter V

Table 1: Cover composition of the most abuncoral cdaover nt genera calculated as mat all sites studied aean (± SE). long Similan Islands. Percentage of total

een island sides (ANOSIM, one-way-analysis: W position differed clearly betwCoral species com

tal Table 2) resulting in a distinct clustering in the neversus E: global R = 0.585, p < 0.001, supplem

and E sites, respectively (FMDS plot of the W higher species ig. 3, see also Table 2). In the W

ogeneous distribution ore homdiversity (t-test, p < 0.01) and richness (t-test, p < 0.02) along with a m

Fig. 4): Shannon index was 2.69 ± pared to the E (evenness: t-test, p < 0.04;pattern were found com

nd a0.15 in the W and 2.03 ± 0.13 in the E, species richness showed 38.34 ± 2.35 species in the W

and 0.63 ± 0.04 in the E. The 27.33 ± 3.45 in the E, and evenness revealed 0.74 ± 0.04 in the W

and Epositions were relatively low among Wilarities calculated on the basis of species comsim

ess es the high species richnsites, respectively (Table 2). This can be explained by the fact that besid

and evenness in W, the species compositions in W differed among sites (supplemental Table 2).

inance by a ber of species, and a higher domWhereas the E sites were characterized by a lower num

variable subset of species, including two species of Porites(P. lutea dominating in five out of nine E

locations; and P. rus, dominating in four locations) and Hydnophora rigida (dominating in one E

ental Table 2). location) (supplem

Figure 3: MDS ordiBray-Curtis Simnation ilarities. LAIW-eof coral commxposeunities d () and shelterein the Similan Islands bad () sed sites are on the species abundagrouped in sepance rate clusters. data (%) and

142

Chapter V

betweenTable 2: Inter LAIW-exposed -site com(West) parison of coral comand LAIW-sheltered (munities withEast) the, (B top) similarities (sim10 coral species ) withincontributing m LAIW-exposost to (Aed (West), ) the diand (C) simssimilarities (diss) ilarities
within LAIW-sheltered (East) sides of thstandard dee Similan Islands. Values viation of dissimilardenote mean (±ity and similarity, respectiv elSE) percentage of total y. living coral cover, SD:

Benthic substrate composition – Comparing the benthic cover com

onsoon-mpositions in shallow (

impact), mid and deep (LAIW-impact) depths, the clearly lowest living coral cover was found in W

4.1 versus E 4.1: global R = 0.639, p < 0.02). ay-crossed analysis: Wdeep (ANOSIM, two-w

versus 12 ± (35.67 ± 5.61 % in 7 min shallow WSignificantly higher living coral cover was found

4.36 % in 20 md-waters in E (36.67 ± 4.63 % in 20 m depth and 33.57 i depth) and in deeper and m

% in 14 m depth versus 18.67 ± 5.49 % in 7 m depth; 20 m versus 7 m: global R = 0.37, p < 0.05)

(Fig. 5). Living coral cover was the second most powerful contributor to the dissimilarity of 72.5 %

portant contribution was achieved by ost im and E with a contribution of 25.06 %. The mbetween W

sand and rubble with 35.86 %, thirdly followed by rock (17.93 %). In the m

id-depths located in

143

Chapter V

and monsoon waves differences in the overall benthic LAIWpact ofbetween the direct imnd E sides (ANOSIM, one-way analysis: global R aposition failed to be significant between Wcomediate depth at all 16 sites living coral cover averaged = 0.079, p = 0.22). In detail, for the interm25.57 ± 4.22 % in W and 35.95 ± 4.62 % in E. Total hard substrate and sand including rubble also and E. Total hard substrate consisting of dead coral, living coral positions at Wlar comishowed sim and 72.83 ± 3.34 % in E, sand and rubble 29.75 ± 6.89 % in and rock averaged 67.71 ± 6.59 % in W while in E hard substrate ,W and 25.88 ± 3.29 % in E. Rock structures were only found in Wental Table 1). l (Fig. 6, supplemconsisted exclusively of dead cora

k3.53.02.5Species diversity, H2.01.51.00.90.8k0.7Eveness, E0.60.5

0.45040Species Richness, S302010

A

B

C

stEaWest Figure 4: Coral diversity (Shannon index, sheltered A), eve(East) nness side(B), s of Simand species ilan Islands. richness (C) at LAIW-exposed (West), and

144

100LAIW-exposed LA

IW-sheltered

Chapter V

8060cover [%]sand&rubblerock40al dead coralgae20sponge0soflivingt corals corals
W4.1: 7 mW4.1: 14 mW4.1: 20 mE4.1: 7 mE4.1: 14 mE4.1: 20 m
Figure 5: Benthic composition at Koh Miang on LAIW-exposed (W 4.1) and sheltered (E 4.1) side, grouped by depth:
.and 20 m147,

LAIW-sheltered

100LAIW-exposed LAIW-sheltered
80ble nd&rubsa60cover (%)40aldgaeead co ral
rock20sospft congoe rals
living corals 0W2.1W4.1W7.1W8.1W8.2W8.3W9.1E1.1E2.1E4.1E4.2E7.1E8.1E8.2E8.3E9.1
Figure 6: Benthic composition at Similan Islands on LAIW-exposed west (W), and east (E) sides (12 to 16 m depth,
orientation). d by N-S groupe

Coral framework and morphologies – A dense and complex coral framework characterized the E
hard corals were distributed as solitary colonies without developing any actualwhereas in the Worphologies differed ework (W 4.1 versus E 4.1, depths pooled: p < 0.001) (Fig. 7). Coral mmfra versus E, global R = 0.311, significantly between sides (ANOSIM two-way-crossed analysis: Wp < 0.05) and depths (7 vs. 20 m, global R = 0.253, p < 0.05) (Fig. 8). Although the dominance of
of Koh Miang fell short of and of branching species in Essive and encrusting colonies in Wamards being significant (t-test, p = 0.064 and p = 0.1, respectively) Fig. 8 shows clear tendencies towthis pattern. Large massive and encrusting hard coral species sparely covered deeper areas in the W,
orphological types nestled to the rocks in all often densely thronged colonies of all mand smshallower waters. Here, branching hard corals, especially within the genera Millepora and Acropora
orphologies with pancake-like broadened bases and strongly reduced often displayed flattened mification (Fig. 1 A). ram

145

Chapter V

1008060(cm) height from bottomW4.1:7 mW4.1:20 mE4.1:7 mE4.1:20 m
40200 Figure 7: Coral framework at Koh Miang on LAIW-exposed (W 4.1) and sheltered (E 4.1) side, grouped by depth: 7
.and 20 m

AIW-sheltered

100LAIW-exposed LAIW-sheltered
8060 morphologieslacorrate (%) of W4.1: 7 mW4.1: 14 mW4.1: 20 mE4.1: 7 mE4.1: 14 mE4.1: 20 m
40enmassicrusveting
ingnchbra20rlaminaylitarso0Figure8: Coral morphologies as fractions of total number of colonies studied at Koh Miang on LAIW-exposed (W 4.1)
and sheltered (E 4.1) side, grouped by depth: 7, 14, and 20 m.
entical environmPhysico-chem than in the E, but T < 0.3 °C) in the Wperatures were barely lower (modal values, Temcharacterized by very high variability with up to 10 °C drops during extreme LAIW-events in the W,
DC) averaged the sheltered E. Corresponding degree days cooling (Dnas opposed to < 5 °C i and E both depths pooled, respectively (Student’s -125.94 ± 12.94 °Cd and -49.10 ± 5.74 °Cd for Wy of the temperature data recorded. Fig. 9 shows the ar< 0.001). Table 3 represents the summt-test, p ed on the lower frequency variations posplete temperature record for Koh Miang. Superimcomhich requency oscillations wcycles, seasons) were periods of high-f(diurnal tides, spring-neap tides perature drops as February to April and October to November. Temost pronounced fromwere m sites. Less pronounced oscillations depth at the Wlarge as 10 °C were observed in spring in 20 mber rd at W Koh Miang (Decemperature recoe temoccurred in the shallower and sheltered E sites. Thool water events increased agnitude and duration of c2007 to April 2008) reveals also that the m for Wportance of LAIing the imsignificantly with depth. This was less obvious in the E, underscor slope of the island (Table 3). the W

146

Chapter V

Table 3: Summary of temperature (°C) and degree days cooling, DDC (°Cd), of all sites recorded along Similan Islands
and of Koh Miang in detail. Temperature values calculated as mean, modal, and range (± SE), respectively, of complete
time period recorded (all sites: Feb 2007 to Nov 2008, Koh Miang: Dec 2007 through March 2008); DDC calculated
from complete data set of 20 week period for all sites (Feb 2007 to July 2007) and of 16 week period at Koh Miang (Dec
2007 through March 2008).

Table 4 shows the statistics summary of the further physical and chemical parameters measured at
Koh Miang. Mean current velocities were twice as high in W as in E (Kruskal-Wallis test,
p < 0.001) with maximum speeds reaching 0.65 and 0.73 ms-1 in W deep and shallow opposed to
0.58 and 0.56 ms-1 in E deep and shallow respectively. Higher velocities coincided with lower

temperature, while under constant tem-1perature conditions (at most 0.5 °C below modal) water
ental Fig. 2) Oxygen concentrations, pH and (supplemsmotion stayed stable at 0.13 ± 0.001 msalinity were highly correlated with temperature. The impact of LAIW in W (5 to 10 °C) caused
drops in oxygen concentration of up to 88 % (down to 21.80 μmolL-1), in pH of up to 0.6 units

ither optical backscatter nor (down to 7.75) and increased in salinity up to 5 % to values of 34.52. Ne

fluorescence showed temperature dependent variations. Yet a tendency of lower chlorophyll concentrations up to 40 % in LAIW-water of 20 ± 0.5 °C was noted when compared to constant

perature conditions. modal tem

re et intensities were found in E shallow and lowest in W deep. Light conditions wHighest lighsignificantly different on W and E in both depths (7 and 20 m) (Kruskal-Wallis test, p < 0.001,

light values in E during the main part of the es higher ental Table 3 B), with over three timsupplem

day and a longer lasting light environment in W at the end of the day (Table 4 C, supplemental Fig.

3). During a period of turgescent LAIW-impact in January 2007 photosynthetic active radiation
(PAR) never exceeded 141.5 μmol quanta m-2 s-1 in W 20 m with a mean of 102.5 ± 3.4 (Fig. 9).

147

Table 4Chapter V

: Environmental parameters at Koh Miang (W 4.1 and E 4.1) displayed as mean (± SE). Linear regression model

with each environmental parameter as dependent and water temperature as independent variable: (A) Parameters

measured continuously every -2minute and (B) parameters determined from water samples. (C) Averaged light values (lux
[lm m]) over a period of 4 months from December 2007 to April 2008.

m[lm

]) over a period of 4 months from December 2007 to April 2008.

(Significance levels are *0.05 > P  0.01, **0.01 > P  0.001, ***P < 0.001)

148

Chapter V

and E for nitrate and nitrite, as well as for Nutrients revealed significant differences between W

silicate (Kruallis test, p < 0.009 and p < 0.002, respectively). Mean concentrations of silicate skal-W

were more than 100 % higher in W 20 m than in E 20 m, supported by an increase of nitrate and

trations. In shallow waters the differences ost 200 %) and of phosphate (30 %) concennitrite (alm

were still noticeable but far less pronounced and statistically not significant. Thereby concentrations

perature when correlating to the in-situ exhibited a clearly negative relationship with tem

temperature during sampling (Table 4 B, supplemental Table 3). Ammonium showed 45 % higher

ean concentrations in W than in E but did not clearly correlate negatively with water temmperature.

No significant differences were found between sides in dissolved organic carbon and suspended

s which variedatter concentrationparticulate m

ental Table 3). (supplem

independent f

rom temperature variations

Figure 9: Large Amplitude Internal Wave (LAIW) associated temperature variations on W (left panels) and E sides of
Similan Islands. Examples are given for temperature logger readings from LAIW-exposed (W 4.1) and sheltered (E 4.1)
side of Koh Miang in 20 m and 7 m depth. Upper panels show full temperature record (March 2007 - March 2008; blank
periods are missing values). Insets (lower panels) highlight 24 h-periods of high (March 2007) and low LAIW activity
(December 2007), respectively, showing the frequency, intensity and duration of cold swells associated with LAIW.

149

Chapter V

perature ulative temeters and cummmunity paramCorrelations and relationships of coral co

aliesanomhannon index) and species richness exhibited a clearly positive relationship with degree Diversity (S

le hard substrate was inversely, e availab of thdays cooling (DDC). Living coral cover as a fractione clearer after considering the becambut only weakly, related to DDC. This negative correlation and E (island Koh ring Wpaation comdepth gradient of DDC and living coral cover in the calculMiang, Table 5).

Table 5: Relationship between coral diversity and LAIW intensity. Linear regression model with Shannon index,
Evenness, species richness and coral cover as dependent and degree days cooling (DDC in °Cd) as independent
bles. varia(Significance levels are *0.05 > P  0.01, **0.01 > P  0.001, ***P < 0.001)

Discussion

, or solitons) have been known for plitude Internal Waves (LAIWAlthough non-linear Large Am

decades from the Andaman Sea (Perry & Schimke 1965, Osborne & Burch 1980) and many other
to date to establish their existence in islands tropical oceans (Jackson 2004), our report is the first bordering the Andaman basin, and the first to investigate their relation with coral communities. The
oderate and exceed by far the mperature variations in our studyfrequency and severity of the tem

ter et al. 1996, 2003, Leichter & eichlower-frequency variations reported earlier elsewhere (L variations so far reported in tropical reef ong the largest short-termGenovese 2006), and rank am’s energy was found to wrap around the islands, W of the LAIeareas (Sheppard 2009). Although somevidenced by the dampened oscillations in our deep E temperature loggers, the deep ocean-facing W
posed on the violent short-term ost profoundly affected. Superimsides of the islands were m

150

Chapter V

variations in the temperature time-series were lower-frequency variations (spring-neap time and
seasonal) revealing highest LAIW-activities after spring tides and during the late NE monsoon
-generating and -Wodulations are related to variations in both, LAI(March, Fig. 9). These mpropagating factors, i.e. tidal current variations at the generation sites, which vary with the lunar cycle (Pineda 1995) and the depth and strength of the seasonal pycnocline, where the shallow pycnocline during the NE monsoon (Nielsen et al. 2004) corresponds with strongest LAIW activity.
ena raising and re are linear phenom the literatuany of the internal waves reported inWhile m(Garrett & Munk 1979, Alpers 1985), the non-linear ocline over tidal periods depressing the thermLAIW recorded in this study cause much larger temperature variations on much smaller time-scales.
(up to 10 °C below ambient) can be the reefing The source depth of the coldest water reachestimated from the 18 °C isotherm in the Andaman Sea, which oscillates between 100 and 150 m
(Nielsen et al. 2004). The actual depth may be much deeper, depending on the degree of turbulent
miTropical hermxing of the cold bores with above-thermatypic corals are known to occur over a wide geographical range of temocline waters (Vlasenko & Hutter 2002). peratures (e.g.
up to 12 °C differences in maximum summer temperatures for species co-occurring in the Arabian
ord Howe Island, Australia (Hughes et al. 2003), tolerate large annual ranges of Sea and LtemFadlallah 1991)). Yet it has been shperature (up to 25 °C) and survive severe cold periods (13 °C fown in several studies that cold water stress (starting with 6or several days; Coles and to
10 °C under normal conditions) has a strongly negative influence on the outward appearance of
ion with their corals and their physiology (Coles & Fadlallah 1991), on the sensitive associat1992), that it can decrease the photosynthetic biotic zooxanthellae (Gates et al. endosymforperformmations (Burns 1985, Kleypas et al. 1999). Tance of a coral (Saxby et al. 2003), and undermhese reports support the assumine the development of dense coral reefption that the reduced
coral cover in W 20 m depth and the absence of reef framework in W may be partly due to the
observed temperature oscillations. ny co-occurring aHowever the cold temperatures in our study are only one stressor out of menvironmental factors: Temperature was related to the other environmental parameters in a
conservative way with a strong positive correlation with oxygen and pH, negative correlation with several nutrients and salinity, and concomDennison & Barnes (1988) have shown that high current speeds enhance photosynthesis and itant increases in current speeds (Table 4).
er adjacent to the coral surface and increasing gas by scaling down the boundary laycalcificationexchange. The combination of higher current velocities and nutrient concentrations can enhance

151

Chapter V

ant 2002). The latter can becoral nutrient uptake (Hearn et al. 2001) and photosynthesis (Szm

highest in cooler water (23 to 26 °C, Al Horani 2005) suggesting that the negative effect of low

waters could be counterbalanced by the photosynthetic oxygen Woxygen concentrations in LAI

milan ieasured at the Sproduction by the corals (Shashar et al. 1993). The nutrient concentrations m

ean concentrations for -period (February, March 2007) led to mWislands during a typical high LAI

trations of m clearly above the averaged concennitrate, nitrite and phosphate in W ost tropical reefs

e values assessed for coral reef communities (Table 4 B; or even above the extremand close to

neutral or positive effects on Kleypas et al. 1999). Increased concentrations of these nutrients entail

ntioned above), physiology and zooxanthellae numbers (Muscatine et al. 1989, ecoral nutrition (as m

Ferrier-Pages et. 2001). However, they e indirect effect of nutrients are likely dwarfed by the negativ

on the balance between corals and space-competing macroalgae. Raised nutrient concentrations in

acroalgae, known to be comthe water can favour growth and expansion of turf- and m petitors of

McCook 1999). Nutrient-penter 1988,s and Carilliamcorals in the struggle for light and space (W

y further enhance internal bioerosion of coral skeletons and the aenhanced pelagic productivity m

reef framework, since the majority of endolith bioeroders are suspension or filter feeders (Glynn

rd-Paige et al. 2005). a1997, Chazottes et al. 2002, W

The abrupt drops in pH and oxygen concentrations delivered with the cold water were found to

create short-termre oth decreased pH-values of 0.2 to 0.6 units and frequently m conditions in W wi

on oxygen concentrations (Table 4 A). These effects in combination with than half of the comm

reduced light intensities in W (Table 4 C) may limit or at least strongly reduce coral growth, i.e.

pared to their eastern counterparts as photosynthesis and mcalcification and photosynthesis co

reduced calcification are tightly coupled in zooxanthellate scleractinian corals and suffer from

aragonite saturation states (lowered pH) and light levels (Gattuso et al. 1999, Marubini et al. 2001).

by Schneider & Erez (2006)ents on coral calcification and photosynthesisLaboratory experimrevealed that pH reduced by only 0.2 units might implicate a 30 % reduction of CO32- and decrease

i deep mt intensities in Wcalcification by 50 %. On the other hand especially the reduced lighght

ratures by corals as Coles & Jokiel (1978) already pefavour the successful toleration of the low tem

demonstrated the highly negative effects of combined low temperatures and high light intensities

leading to a substantial deterioration of the c

mortality rates.

nd to significantly higher aoral host metabolism

152

Chapter V

Salinity concentrations, although highly correlated with temperature, always stayed within a range

al for tropical reef conditions (Kleypas et al. 1999), and , which is normof 32.6 and 34.4 ppm

portant community shaping factor in our study. ated as an imtherefore was not estim

a suspended particulate mThe lack of difference in and E tter concentrations (Table 4 B) between W

ariations (see also optical backscatter, Table 4 A) and the apparent independence of temperature v

nces in the physical oceanographic variables, appears inconsistent with the pronounced differe

xing to be involved. isuggesting other processes than m

eas with LAIWHigher concentrations of chlorophyll transported into shallow ar-arrival were found

ocline waters. However therme fromin our study and also stated by Leichter et al. (1996) to com

his could possibly lead to a line layers contain low pH and oxygen depleted waters. Tocsub-therm

ow oxygen) and production processes (plankton mutual compensation of depletion processes (l

ncentrations (Eppley et al. 1979)) and could be a reason for the production due to higher nutrient co

and E. s between Wcentrationatter conlack of difference in dissolved organic m

-induced changes in seawater properties (i.e. cold, low-oxygen, low-pH waters ost all LAIWAs alm

with high nutrient loads) are reported to have adverse effects on corals, both directly, by stressing

their physiological and metabolic conditions, and indirectly, by promoting coral competitors such as

y act synergistically to exert a cumulative pulsed al stressors malgae and bioeroders, these potentia

disturbance on the LAIW-exposed coral communities.

inent physical factors shaping coral st promoics are known as the mWhile light and hydrodynam

ures across depth (Falkowski et al 1990, Massel & morphologies, species distributions and reef struct

Done 1993), it is important to note that the LAIW-impact represents a vertical gradient in the

opposite direction. Both, SW monsoon and LAIW impart their strongest impact on the W faces of

onsoonal surface waves declines exponentially with increasing depth the islands; but the force of m

monsoon and increases. The cumulative impact of SWWpact of LAI(Thorpe 2007), while the im

LAIW-disturbances on the Similan islands coral communities is conceptualized in Fig. 10. Although

ental and biological disturbances erous environmes are only two out of numsurface and internal wav

poral scales (i.e. disturbances caused by nd temaffecting benthic communities on various spatial a

grazing and trophic cascades (Mumentation (Cole 2003, by et al. 2006) terrestrial runoff and sedim

ul ant 2002), they provide a usefascik & Sander 1987, SzmFabricius 2005), or eutrophication (Tom

ework for understanding the rather striking differences in coral community composition and mfra

mreef developall levels of wave disturbances are higher on the W faces of the ent in our study. Over

rgins of the depth gradient. As a result, awer mislands than on the E, and higher on the upper and lo

153

Chapter V

low levels of disturbance are found on the E, intermediate levels in intermediate depths of W, and
high levels of disturbance on W-shallow and W-deep (see also cooling rates, CRs, Table 3 and
ental Fig. 4). perature record supplemtem

Figure 10: Graphic description and by mof the seonspaoon rated wave(left pas at W annel) and cumd E sides of Simiulative effects of thelan islands. disturbance by LAIW-impact,
ediate levels of disturbance as in our study corresponds to intermThe highest level of coral diversityspecies will be reached ber of coral highest numtheory predicts: Connell (1978, 1979) suggested that ediate levels (frequency and size) of natural disturbance. Lower diversity results if at intermall. Accordingly disturbances which are either too frequent or too infrequent, or too large or too sm disturbances communities exposed to acute pulsed short-termral ) that cohe found (Connell 1997rather than to chronic long-term ones are able to recover faster and more completely and develop
ical and physical unities. The observed extreme conditions in the chemhighly diverse commenvironment in our study concurring with the pulsed LAIW-impacts are replaced each time within
odal panels Fig. 9; nearly identical mter conditions (loweroderate wa15 to 30 min by again mperature values on W and E: Table 3). These acute disturbances are likely to be stressful to the temcorals, as discussed above, but their intermittent and short-termed character seams to prevent
physiological damage and mortality observed for longer-term disturbances following cold spells
W ant 2002). The alternation of storm and LAIdlallah 1991) or eutrophication (Szma(Coles & Fpact during the SW (May to October, Phongsuwan & Chansang 1986) and NE monsoon im s of recovery foray also create seasonal periodely, m(February, March, see Fig. 9) periods, respectiv

154

the affected shallow and deep fore-reef areas, respectively. W

Chapter V

eherefore propose that in t

intermediate depths in W, the alternating impact of SW monsoon and LAIW from above and below,

respectively, contribute to a spatio-temporal heterogeneity maintaining the community in a non-

he sheltered E reefs which state, resetting succession and enhancing species diversity. Tequilibrium

are only some hundred meters apart from their western counterparts reside in sheltered tropical reef

oval of bances prevents the destruction or remental disturconditions, where the lack of environm

onopolizing thspecies milarly Rogers (1993) and d diversity is lowered (Fig. 4). Sime space, an

-generated disturbances to possibly ere is need of i.e. stormAronson & Precht (1995) found that th

petitively subordinate species to increase. ing comreduce the cover of dominant coral species allow

pered reef developmThe hament in W shallow and deep contrasting the areas in E (Fig. 7) reflects

apparently too high levels of disturbance to build and maintain an actual reef framework. Based on

atology (Brown 2007) and previous investigations in the area (Phongsuwan & regional clim

pact of strong surface gravity waves during the SChansang 1986, Phongsuwan 1991) the im-W

monsoon periods is strongest along W shallow reef areas. Corals there grow nestled and disjointed

pact and to reduce physical damage the waves´ iment to shelter from directly to the granite basem

the greater distance between single coral stands and the reduced ,(Storlazzi et al. 2005). In deeper W

numnd builders lead to the lack of reef development ber of branching species as framework spacers a

perature, bined negative stressors (reduced light, tempact of comngest im(Fig. 8) due to the stro

oxygen, pH and increased nutrient concentrations, discussed above) coinciding with the frequently

-Wstrong swell-like currents of LAIitations, to arrivals. Although it was possible, for logistic lim

e pattern was evident throughout the quantify these findings for only one of the nine islands, the sam

disturbance by idt, pers. observations). Hence high levels ofilans (N. Phongsuwan, G. SchmSim

surface waves in shallow W and LAIW in deep W inhibit reef framework building due to their

chronically disturbing character. It is noteworthy that this lack of destructible reef formations was

i in 2004 on pact of the Indian Ocean tsunamlow destructive imparatively the reason for the com

hongsuwan & Brown 2007). The tsunamilan island reefs (Phongsuwan et al. 2006, PSimi struck the

ale and patchy destruction patterns in E reefsall sc the SW and caused only smisland chain from

with an initial total of 31.6 % damage (high damage (> 50 %): 18.4 %, moderate damage (31 – 50

in 2006). m%): 13.2 %, Plathong 2005, Yee

ent (Fig. e striking W-E differences in reef developmentioned that thestext it needs to be mIn this con

1, Fig. 7) were not fully captured in the line intercept transects (LIT, Loya and Slobodkin 1971,

Loya 1972, 1976). The LIT’s original strength to reduce the comensional coral plexity of a three-dim

155

Chapter V

s expand in a two-easure is not convenient when organismensional mcommunity into a two-dimensional plane as in our study. dimunity reflects the net result of the governing physical, chemical position of a coral commThe comishelson 1973, Loya 1975, De Vantier et al. 1998). Corals growing along and biological drivers (Fent to ical environmilan islands are exposed to an extraordinary stressful physico-chemthe W of Simlages could provide a b therefore propose that these coral assemewhich they obviously adapted. Wre increase and ocean acidification due to their peratuclue to coral resilience in an era of global tem-pulsed low successful and potentially variegated adaptation processes to this cocktail of LAIWenon are a ubiquitous phenomWperature, low pH, low oxygen and high nutrient conditions. LAItemin SE Asia and beyond (Jackson 2004), yet they have not been considered with the so far adequate unities. arding their strong effects on coral reef commtion regatten

Acknowledgments

eef The research for this study was carried out within the cooperation project ORCAS (Ocean-RCoupling in the Andaman Sea) funded by the German Research Foundation (DFG, RI 1074/7-1) and
The authors would like to thank the Phuket the National Research Council of Thailand (NRCT).land National Park staff for field assistance, as ilan IsMarine Biological Center (PMBC) and the Simhnical, Matthias Birkicht and Dorothee Dasbach for laboratory well as Tobias Funke for tecassistance.

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and species distribution in the Hawaiian Islands. Coral Reefs 24:43-55
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Veron Vlasenko V, JEN (2000) Hutter CoK (2002) rals of thNume World. Austerical experiments on tralian Institute of he breakiMarine ng of solitary internal waves Science, Townsville, Australia over a slope-
shelf topography. J Phys Oceanogr 32:1779–1793
Vlasenko boV, ttomStashc topohuk grapN h(2007) y. J Phys AmOceanplification aogr 36:n1d s959u–19ppre73 ssion of internal waves by tides over variable
Ward-Paige CA, Risk MJ, Sherwood OA, Jaap WC (2005) Clionid sponge surveys on the Florida reef tract
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Williams SL, contCarpenter RC ribution of ammonium(1988) Nitroge excreted n-limby ited primarDiadema antillarum*y productivity . Mar Ecol Pof coral reef alrog Ser 47:gal turfs: 145-152 potential
Wolanski E, Delesalle B (1995) Upwelling by internal waves, Tahiti, French Polynesia. Continental Shelf
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island, Great Barrier Reef of Australia. Estuarine, Coastal and Shelf Science 65:153-158
Wu G, Zhang Y (1998) Tibetan Plateau Forcing and the Timing of the Monsoon Onset over South Asia and
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aps _m.org/giswww.reefbaseYeemin T, Sutthacheep M, Pettongma R (2006) Coral reef restoration projects in Thailand. Ocean & Coastal
ent 49:562-575 Managem

159

Supplemental table1Supplemental data Chapter V

Chapter V

: Benthic composition at Similan islands on LAIW-exposed (West), and sheltered (East) sides in

12 to 16 m depth. Data displayed as mean (± SE).

160

Supplemental table2Chapter V

:Summary of presence and absence of hard coral species with an approximate quantitative

assessment of the proportion of the total substrate cover measured with one 100 m line intercept transect in 12 to 16 m

depth on LAIW-exposed (West), and sheltered (East) sides.

161

Supplemental table2 continued

Chapter V

162

Supplemental table

2 continued

Chapter V

163

Supplemental table3Chapter V

:Environmental parameters at Koh Miang (W 4.1, E 4.1). Results of non-parametric Kruskal-

Wallis ANOVA by ranks (p-values, significance levels). (A) Water current measured continuously every minute over

period of 4 weeks (February, March 2007), (B) light values (lux [lm m

-2

ber (Decemnths omr period of 4 d overecorde])

2007 to April 2008) , and (C) parameters determined from water sample analyses collected in February, March 2007.

(Significance levels are *0.05 > P  0.01, **0.01 > P  0.001, ***P < 0.001)

164

LAIW-sheltered

Chapter V

100LAIW-exposedLAIW-sheltered
8060AcPoriroptesora
revo of total coral ccover [%]0rest
40PoHydnocilloporaphora
vonaaP20MilleporaW2.1W4.1W7.1W8.1W8.2W8.3W9.1E1.1E2.1E4.1E4.2E7.1E8.1E8.2E8.3E9.1
Supplemental figure 1: Most abundant genera of stony corals as percentage of living coral cover on LAIW-exposed
west (W), and east (E) sides (12 to 16 m depth, grouped by N-S orientation).

Supplemental figure 2: Relationship between current velocity and temperature. Linear regression model with current
velocity as the dependent, and temperature as the independent variable (p < 0.001): A) West and B) East, both in 20 m
h. dept

165

Chapter V

over periSupplemental fig 3od of 30 days in FebruaDaily light curve at ry, March Koh 2007; sample inMiang. Loggers weterval 1 mre fiixen, raw d in 7 mdata in deptbach on W akground, mnd E side ean values(W 4.1, E calculated 4.1)
for every daily analogue minute over whole sample period.

ExamSupplemental fig 4ples are given Lafor temrge Amperatureplitude Internal logger readiWangsve (L fromAIW) ass LAIW-exposeociated d tem(W 4.1) aperaturend vasheltered (Eriations at the Sim 4.1) sides of ilan Islands. Koh
Miang iLAIW n swas20 mh z (witone) dehin Lpth. PaAIW swasnels shh deow pth) full tem15 m and perature rec10 mord (interm(Decemediate LAIW ber 2007 exposur- April 2008; e) and 7 mblank peri (beyond theods are m reacissinh ofg
). values

166

1

- Chapter VI -

1A

Riverine input of particulate material and

Chapter VI

inorganic nutrients to a coastal reef ecosystem at

the Caribbean coast of Costa Rica

Cornelia Roder

1

, Jorge Cortés

2

arlos Jimen, Cze

Leibniz Center for Tropical Marine Ecology, Z

Germany2Centro de Investigación en Ciencias del Mar y Limnología, CIMAR, Ciudad de la

2

and Rubén Lara

1

T, Fahrenheitstrasse 6, 28359 Bremen, M

Centro de Investigación en Ciencias del Mar y Limnología, CIMAR, Ciudad de la

Investigación, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica

Published in Marine Pollution Bulletin (2009) 58: 1937-1943

1B

Tracing the extend of fertilizer input on coral

metabolism

Additional information on Roder et al., 2009

167

Baseline/MarinePollutionBulletin58(2009)1922–1952

Chapter VI

1937

extractionchlorinatedandgashydrocarbonsinchromatography/masssedimentsandtissuesspectrometry.usingIn:acceleratedOstrander,solventG.K.Wells,S.,R.S.,Jarman,Tornero,W.M.,V.,Hohn,Borrell,A.A.,A.,Aguilar,Sweeney,A.,J.C.,Rowles,2005.T.K.,IntegratingRhinehart,H.L.,life-historyHofmann,and
(Ed.),(Chapter35).TechniquesinAquaticToxicology,vol.2.CRCPress,BocaRaton,FLreproductiveorganochlorinesuccesscompoundsdatafortobottlenoseexaminedolphinspotential(Tursiopsrelationshipstruncatus)iwithn
Sloan,Yanagida,C.A.,Brown,G.K.,Krahn,D.W.,M.M.,Ylitalo,2006.G.M.,QualityBuzitis,J.,AssuranceHerman,PlanD.P.,forBurrows,AnalysesD.G.,ofWestgate,SarasotaA.J.,Bay,Muir,Florida.D.C.G.,ScienceGaskin,ofD.E.,theTotalKingsley,EnvironmentM.C.S.,1997.349,106–119.Concentrationsand
EnvironmentalSamplesforPolycyclicAromaticCompounds,PersistentOrganicaccumulationpatternsoforganochlorinecontaminantsintheblubberof
Pollutants,FattyAcids,StableIsotopeRatios,LipidClasses,andMetabolitesofharbourporpoises,Phocoenaphocoena,fromthecoastofNewfoundland,the
PolycyclicAromaticCompounds.NOAATech.Mem.NMFS-NWFSC-77.NMFS,GulfofSt.LawrenceandtheBayofFundy/GulfofMaine.Environmental
StateUSofHawaii,DepartmentofDepartmentCommerce,ofSeattle,Business,WA,EconomicUSA.DevelopmentandTourism,Wolman,PollutionA.A.,95,Wilson105–119.Jr.,A.J.,1970.Occurrenceofpesticidesinwhales.Pesticides
2008.Whatarethedbedt/info/economic/library/faq/faq08>.MajorIndustriesintheStateofHawaii?<http://hawaii.gov/Xu,T.,Cho,MonitoringI.K.,Wang,JournalD.,4(1),Rubio,8–10.F.M.,Shelver,W.L.,Gasc,A.M.E.,Li,J.,Li,Q.X.,2009.
theSubramanian,testosteroneA.,Tanabe,levelsS.,byTatsukawa,PCBsandR.,DDESaito,inS.,Dall’sMiyazaki,porpoisesN.,of1987.ReductionNorthwesterninHawaiianSuitabilityofaeuryhalinemagneticfishandparticlecrabsinimmunoassaycomparisonforwiththegasanalysisofchromatography/PBDEsin
Tanita,NorthR.,Pacific.Johnson,MarineJ.M.,Chun,PollutionM.,BulletinMaciolek,18J.,(12),1976.643–646.OrganochlorinepesticideselectronPollution157,capture417–422.detection-iontrapmassspectrometry.Environmental
in29.theHawaiiKaimarina,1970–1974.PesticidesMonitoringJournal10,24–Yang,F.,quantitativeWilcox,B.,analysisJin,S.,ofAlonso,A.A.,polychlorinatedRougee,L.,biphenylsXu,Y.,inLu,Y.,tilapia2008.fromDetectionHawaiianand
Tilbury,K.L.,Adams,N.G.,Krone,C.A.,Meador,J.P.,Early,G.,Varanasi,U.,1999.waters.Chemosphere73,133–137.
ofOrganochlorinesMassachusetts.inArchivesstrandedofpilotEnvironmentalwhales(GlobicephalaContaminationmelaenaand)fromToxicologytheCoast37,Ylitalo,2001.G.M.,InfluenceMatkin,ofC.O.,life-historyBuzitis,J.,parametersKrahn,onM.M.,Jones,organochlorineL.L.,Rowles,T.,concentrationsStein,J.E.,in
125–134.free-rangingkillerwhales(Orcinusorca)fromPrinceWilliamSound,AK.Science
UnitedNationsEnvironmentProgrammeandFoodandAgricultureOrganizationofoftheTotalEnvironment281,183–203.
theProvidedUnitedbyCanada.NationsReport(UNEP/FAO),No.2005.UNEP/FAO/RC/CRC.2/1.Mirex:SupportingDocumentationYlitalo,lipidG.M.,classesandYanagida,lipidG.K.,contentHufnagleintissuesJr.,L.,ofKrahn,aquaticM.M.,organisms2005.usingaDeterminationthinlayerof
USEthersEnvironmental(PBDEs)ProjectProtectionPlan.Agency(USEPA),2006.PolybrominatedDiphenylOstrander,chromatography/flameG.K.(Ed.),TechniquesionizationindetectionAquaticToxicology,(TLC/FID)vol.microlipid2.CRCPress,method.BocaIn:
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ehp.0900785.Availablefrom:<http://dx.doi.org/>.NewJersey.

0025-326X/$-seefrontmatterPublishedbyElsevierLtd.
10.1016/j.marpolbul.2009.08.029

1A

Riverineinputofparticulatematerialandinorganicnutrientstoacoastalreef
ecosystemattheCaribbeancoastofCostaRica
CorneliaRodera,*,JorgeCortésb,CarlosJiménezb,RubénLaraa
aLeibnizCenterforTropicalMarineEcology(ZMT),Fahrenheitstrasse6,28359Bremen,Germany
bCentrodeInvestigaciónenCienciasdelMaryLimnología(CIMAR),CiudaddelaInvestigación,UniversidaddeCostaRica,SanPedro,11501-2060SanJosé,CostaRica

Reasonsforthealarmingglobalcoralreefdestructioncanoften
bybelandfoundonclearing,land(ISRS,fertilization,2004).useofAgriculturalpesticides,activitiesandurbanizationaccompaniedor
ductiontourismareofexpansionconcernalong(ISRS,with2004).enhancedRiversandsewageandgroundwatewastercarrypro-
highloadsofsediment,nutrientsandotherpollutantstothesea,
wheretheycanhaveseriousimpactsonnearshoreecosystems
such1992;asRogers,coralreefs1990;(CortésFabricius,and2005).Risk,1985;GuzmánandJiménez,
inhibitWhileitwhennutrientshighlyenhanceconcentratedcoralgrowth(TomascikinlowerandSander,amounts,1985;they
Koopetal.,2001),andacceleratetheprogressandseverityofcoral
fueldiseasealgal(Brunogrowthetal.,and,2003;combinedVossandwithreducedRichardson,2006).herbivory,canNutrientsbe
responsible(Díaz-PulidoforandshiftsMcCook,from2003;coral-Hughestoetal.,2003).algal-dominatedAlsobioero-reefs
andderssuchorganicasalgae,mattersponges,increaseworms(RiskorandbivalvesMacGeachy,profitfrom1978).nutrientBored

*Correspondingauthor.Tel.:+49(0)4212380087.
cornelia.roder@googlemail.cornelia.roder@zmt-bremen.de,addresses:E-mailRoder).(C.com

sedimentsandcoralsarelessresistanttostormsandwaves,result-
inginreeferosion(Hallock,1988;Chazottesetal.,2002).
Suspendedmatterinthewatercolumndecreasestransparency
andlightavailability.Whileorganicmaterialmayinitiallybeused
asanadditionalfoodsourcebycorals,thisbenefitisoutweighedin
turbidwater,wherephotosynthesisandcalcificationarereduced
(Rogers,1983;AnthonyandFabricius,2000).Smotheringbypar-
ticulatematerialforcesthecoraltocleanitssurfaceusingenergy
neededforgrowthorreproduction(TomascikandSander,1987;
EdmundsandDavies,1989).Terrestrialrunoffcanbecomea
seriousthreatforreefcommunitiesandevensmallrivershave
beenshowntoinfluencereefswithinafewkilometersdistance
totheirmouths(WestandvanWoesik,2001;Fabricius,2005).
Theaimofthisstudywastoevaluatethepresentinfluenceofa
heavilyanthropogenicimpactedriveronthedistributionofpartic-
ulatematerialanddissolvedinorganicnutrientsinthewatersofa
nearbycoralreefareainCostaRica.
TheCaribbeancoastofCostaRicaischaracterizedbyhumid,hot
climatewithyear-roundrainsofabout6000mm(Cortésand
Jiménez,2003).PrecipitationbetweenDecemberandFebruary
andbetweenJuneandAugustishighercomparedtotherestof
theyear.Thelargest,best-developedandmostdiversereefinthe
areaisfoundintheCahuitaNationalPark(CortésandLeón,2002;

168

Chapter VI

169

2.Fig.

1922–1952(2009)58BulletinPollutionMarine/Baseline

Transparencyanddistributionofsuspendedmaterialanddissolvedinorganicnutrientsinthestudyarea.

TSMconcentrations(Fig.2b)inthedryseasonwerehighestin
frontoftherivermouthanddecreasedsignificantlywithdistance
alongthecoastandoffshore(Fig.3).Intherainyseason,TSMcon-

Chapter VI

1939

centrationsagainwerehighestattherivermouth(Fig.2b)and
differencgenerallyetohigherdrythanseasoninthevaluesdrywasseasonnot(Tabledetected.1),butPC,aPNandsignificantPOC

170

1940

Marine/Baseline

Pollution

2Fig.

valueswerealsohighestinfrontoftherivermouthinbothseasons
(Figs.2c–e),butsignificantseasonalorspatialdifferenceswerenot
observed.Increasedirradiationandwatertransparencywithdis-

(2009)58Bulletin

()continued

1922–1952

Chapter VI

tancetotherivermouthorduringdryseasonenhanceprimary
productionandhencearelikelytomasktheinputofparticulate
materialtothecoastalarea.Inthewatersamplestakenattheriver

171

Baseline/MarinePollutionBulletin58(2009)1922–1952

Fig.dry3.(DS:Correlationcrosses,ofdottedeachlines)parameterandrainywithseasonsalinityfor(RS:spatialdiamonds,comparisonsslashedlines).during
Significantdifferencesaremarkedwithanasterisk;n.s.,notsignificant.
stationduringrainyseason,TSMconcentrationswere30times,
PC3,PN2andPOC4timeshigherthanduringdryseason
2).and1(Tables

Chapter VI

1941

TSMconcentrationsinCahuitaNationalParkwatershaveprevi-
andouslyRisk,been1985),measuredandinin1981theandrainy1993season(Cortés,in19791993).and1980Valuesat(Cortésthe
outerreefcrest,eastofthepeninsula,andintheEstrellaRiverin
1979/1980arelowerthanthoseobservedinthisstudy(Table1).
In1981and1993,Cortés(1993)reportedTSMconcentrations
<10mgL1inthestationsnearCahuitatown(stations23,26,27
and30),whileinthisstudyforbothseasons,concentrationswere
thefoundyearstobeislikelymoretothanbethetwiceasresulthighof(Tableincreased1).Theerosionincreaseofsoilsoverin
the1987andCaribbean1993werehinterland,expandedwherefrombananaaplantationscultivatedareabetweenof
28,000Concentrhatotheationssizeofallof52,000measurehad(ForoparametersEmaús,were2008).highestinthe
werewatertakenelevateddirectlyduringfromrainytheseason.EstrellaBecauseRiverofandthemostlargeofthemphys-
ityico-chemandicalflocculationgradientarethathighdevelopsresultingintheinrapidsalt-wedgesettlingofproductiv-flocs
(EismaandCadeé,1991;SzekieldaandMcGinnis,1991),hence
giontrappingofthemuchEstrellaoftheRiver.Assuspendedtheratematerialofflocculationwithintheofestuarysuspendedre-
materialandinorganicnutrientsisdependentontheamountof
TSMinthewater(Kranck,1981),TSMandmostnutrientconcen-
trationsoutsidetheriverplumewerestillelevatedduringthe
rainyseason,howevernotasmuchasintheriverwaterfurther
inland.watersThewerehighestfoundinconcentratiothensmouthofareaNOxofandthesilicateEstrellainRivertheincoastalthe
thedrydryaswellseasonasin(Tablethe3),rainyNOseasonandsilicate(Figs.2fandconcentratiog).nsComparedweresig-to
xtively)nificantlyandhighertheduringsignificantrainyinverseseason(p=correlations0.02andwith0.001,salinityrespec-
and(Fig.3)siliconrevealedtothethecoastalEstrellaRiverwaters.astheSilicatemainconcentrsourceofationsnitrogenwere
foundmeasuretodbeduringtwicetheashighrainy(Tableseason3),100kmcomparedsouthtoofconcentratioCahuita,inns
theamarunoff(D’Crozetaffectedal.,2005).ChiriquíHighLagoonatconcentrationstheCaribbeanofsilicatecoastareoffoundPan-
in1995)watersandtheclosehightoamountsdeforestedfoundareasheremay(Hillbricht-Ilkowsbeanotherkaetconse-al.,
quenceoferosionoftheCaribbeanhinterlanddeforestedforagri-
culturalpurposes(Mora-CorderoandChavarría,2008).Intheriver
stations,NOxandsilicateconcentrationsduringtherainyseason
wereIn3.5contrast,and2.5phosphattimesehigherthanconcentrationsduringweredryseasonevenly(Tabledistribut3).ed
inrainythethancoastalintheareadry(Fig.season,2h),andassignificantlytheonlylower(pparameter<0.001)inmeasuredthe
inhigherconcentrationsduringthedrythanduringrainyseasonin
riverduringwaterdryand(Tablestill3).Evenelevatedthoughduringvaluesrainyinriverseason,watertheyweredidhighnot
showphosphateconcentratioconcentrationpeaksnsinwerethelowerriverduringmouththearea.rainyAdditionally,season,
highindicatingprecipitation.dilutionAsratherthanphosphorusincreasedconcentratiooutputnsduringintimesCaribbeanof
1996;marineMcClanaenvironmentshanetareal.,2002),generallyandlowfertilizer(SzmantusedandforForrester,bananas
isandhighotherinfruitsnitrogencontains(Beatononlyetal.,small1995),amountstheriverofisnotphosphorus,themainbut
sourceofphosphate.Alsophosphateconcentrationsmeasuredby
Muller-ParkerandCortés(2001)intherainyseason1997(precip-
itation:weather147.1conditionmm)sofwerethisinstudy.betweenHowever,thosesampledcomparedintothePanama-extreme
nianCahuitawatersareaare(D’Crozmuchetal.,higher2005),(Tablephosphat3).econcentrationsinthe
creDuaseedtionincbothreareesingfsrainandovetherretheisasampsignlinificganttime,incsedreaseimen(diftatifereonncein-

172

Baseline/MarinePollutionBulletin58(2009)1922–1952

Chapter VI

1942Baseline/MarinePollutionBulletin58(2009)1922–1952
1TableMeanconcentrationsoftotalsuspendedmatter(TSM)duringdryseason(DS)2004andrainyseason(RS)2005intheriverwaterandthecoastalareaandthecomparisonwith
TSMconcentrationsmeasuredintheriver,thereefareasandclosetoCahuitaduringpreviousstudies.
[mg/L]TSMThisstudyCortésandRisk(1985)Cortés(1993)
DS2004RS2005RS1979/1980RS1981RS1993
Riverstation30.29±6.081046.05±179.96450
CahuitaCoastalregionstations22.5425.28±±0,964.1324.124.10±±4.880.928.4±1.69.1±2.8
OuterEasternreefReef21.0419.37±±1.951.7721.7518.95±±1.630.495.12.6

2TableMeanconcentrationsofparticulatecarbon(PC),particulatenitrogen(PN)andparticulateorganiccarbon(POC)duringdryseason(DS)2004andrainyseason(RS)2005inthe
riverwaterandthecoastalareaofCahuita.
PC[lg/L]PN[lg/L]POC[lg/L]
studyThisDS2004RS2005DS2004RS2005DS2004RS2005
Riverstation202.58±10.8588.84±4.2215.47±5.8134.55±5.21175.79±12.46700.02±3.82
Coastalstations67.37±5.4747.8408±5.105.26±0.644.43±0.6052.22±5.1542.36±4.79

3TableMeanconcentrationsofdissolvedinorganicnutrientconcentrationsduringdryseason(DS)andrainyseason(RS)intheriverwaterandthecoastalareaofCahuitaandtheir
comparisonwithconcentrationsmeasuredinCaribbeanwatersofPanamaaffectedbyterrestrialfreshwaterrunoff.
Nitrate+Nitrite[lM]Silicate[lM]Phosphate[lM]
ThisstudyD’CrozThisstudyD’CrozThisstudyD’Crozetal.
etal.(2005)etal.(2005)(2005)
DS2004RS2005RS1997DS2004RS2005RS1997DS2004RS2005RS1997
Riverstation6.26±0.1622.58±0.16207.43±5.88501.84±13.360.095±0.0170.034±0.02
Coastalstations0.39±0.090.46±0.091.14±0.2951.79±3.6773.97±3.5424.56±2.863.22±0.390.89±0.370.10±0.02

4TableMeandailysedimentationratesoftotalparticulatematter(PM)andtheshareofitsdifferentgrainsizeswithstandarddeviationsinthecoralreefswestandeastofCahuita
NationalPark’speninsulabetweenOctober2004andJanuary2005.
SamplingdatemgPM/cm2/dayGravel[mg]Coarsesand[mg]Finesand[mg]Siltandclay[mg]
*peninsulaofWest15.10.2004–14.11.200426.94±2.84011.93±2.428.92±2.376.09±1.85
14.11.2004–12.12.200442.65±6.52012.72±0.9616.30±3.5913.62±4.42
Eastof12.12.2004–16.01.2005peninsula**141.53±24.043.58±3.0994.54±38.7127.53±12.1415.89±10.42
15.10.2004–14.11.200419.75±5.6607.39±4.565.78±2.696.58±0.50
12.12.2004–16.01.200514.11.2004–12.12.2004521.80133.68±±82.71289.766.62210.59±±0.4152.7754.95249.09±±30.32148.4035.2023.70±±7.2014.1836.9038.42±±3.4915.36
*p<Significance0.01.levelsforincreaseofsedimentationratesovertime.
**0.001.<p

betweenthemonths)withineachreef(Table4).BarnesandLoughperiodsofonlyafewdaysseverelyharmcoraltissue(Riegland
(1999)recordedsedimentationratesinareefinPapuaNewGuineaBranch,1995)andespeciallysmallgrainsizesincreasethedamage
beforeandaftertheexcavationofagoldmine.Pre-excavation(Weberetal.,2006).Thisimpactposesthehighestthreattothereef
concentrationswerebelow1mgcm2day1,andtheheaviestim-ecosystemintheCahuitaNationalParkascoralorganismswillhave
pactedreefwithsedimentationratesof680mgcm2day1com-toincreasetheirmetaboliccostforsedimentremoval,whichwould
pletelydiedoff.Thehighestsedimentationraterecordedwithinotherwisebeneededforgrowthorreproduction(Telesnickiand
ourstudywasabove800mgcm2day1.AllsizefractionsofsettlingGoldberg,1995;EdmundsandDavies,1989).
materialincreasedinamountovertime,especiallycoarsesandandTheCahuitaNationalParkreefalreadyshowsadeclineinlive
gravelwereabundantintimesofheavyrainandrough2sea1condi-coralcoverandspeciesdiversity(CortésandRisk,1985)andthe
tions(Table4).Sedimentationratesof>100mgcmdayoverongoingchronicexposuretoacombinationofhighnutrientand

173

Baseline/MarinePollutionBulletin58(2009)1922–1952

sedimentloadsislikelytofurtherpersist.Duetosiltation,com-
binedwitheutrophication,algalgrowthandbioerosion(McCook,
1999;Díaz-PullidoandMcCook2003),thelossofthecoralreef
ecosystemandashifttoanalgal-dominatedreefwillcontinue.
gementsAcknowledThisstudywasandconductedincollaborationbetweenthe
LeibnizCenterforTropicalMarineEcology(ZMT),Bremen,Ger-
manyandtheCentrodeInvestigaciónenCienciasdelMaryLim-
nología(CIMAR),UniversidaddeCostaRica.Theauthorswould
liketothankEvaSalas,AnaFonseca,VanessaNielsen,ShinichiSun-
agawa,JavierAguirreandtheCahuitaNationalParkStaffforfield-
worksupportandJenaroAcuña,MatthiasBirkichtandDieter
Peterkefortechnicallaboratoryassistance.
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864–872.42,BulletinPollution

174

er input on coral metabolism 1B - Tracing the extend of fertiliz

Chapter VI

of the National Park in Cahuita, Costa Rica, The waters within the coral reef ecosystemare subjected to nutrient and suspended solid loads introduced by the Estrella River and excessive land use in the Carribean hinterland of Costa Rica omrably originating fpresum

(Cortés 1993, Roder et al. 2009).

Synthetic fertilizer used for excessive fruit crop cultivation is mainly composed of N-
derivates (Beaton et al. 1995), which - being obtained from atmospheric nitrogen fixed
-2 to 2‰ (Clark & Fritz peratures - represent isotopic nitrogen ratios ofunder high tem

ospheric nitrogen (0‰). Natural m1997, McClelland & Valiela 1998), close to those of atisotopic values for N-derivates in water various steps of fractionation after fixation (Yamamand soils on the other hand - having undergone uro et al. 1995, McCutchan et al.
2003) - display isotopic signatures ranging between 4 and 15‰ (Clark & Fritz 1997,

ers is only slightly al. 2007). As the signature of consuman et al. 1997, Leichter etSigmheavier compared to their diet (Yamamuro et al. 1995), 15N can indicate possible

sources of nitrogen (Sammarco et al. 1999) and serve as a good tracer of nitrogenous age (Heikoop et al. 2000), fertilizer (McClelland & Valiela 1998) pounds such as sewcom

or synthetically labeled feed (Hoegh-Guldberg et al. 2004).

coral r-derived nitrogen in the Cahuita National Park,To investigate the uptake of fertilizefragments (total n = 12) of randomly chosen massive starlet corals Siderastrea siderea
olander, 1786) were collected at 4 m depth on the exposed W and the rather (Ellis and Srainy and the dry season using side of the national park’s peninsula in the sheltered E2 were chiseled off the non-shaded upper surface of theSCUBA. Pieces of about 10 cmcoral head. The samsupernatant) and transferred to the laboratoples were transferred into pre-labeled plastic bags (mry for immediate processing. The tissue was ax. 4 ml

M the skeleton with an artist’s airbrush using a buffer (0.4 M NaCl, 20 mseparated from), centrifuged at Tris-HCl, pH = 7.6, 4°C) (Seutin et al. 1991MEDTA and 20 m

maximum speed to pellet all tissue and zooxanthellate components for subsequent
position in a gas desiccation (40°C), grinding and analysis of nitrogen isotopic com

ospheric nitrogen eter (Flash 1112 Analyzer) relative to atmass spectromisotope ratio m

using an Apple Leaf standard (HEKAtech SRM 1515) for calibration.

175

Chapter VI

15) and 0.72 ± 0.3 ‰ (E) in the N ratios of 1.19 ± 0.3 ‰ (WThe coral samples displayed ) and 1.48 ± 0.2 ‰ (E) in the rainy season Wven as low as 0.35 ± 0.2 ‰ (dry and of epared to previously published results of ~ 5 ‰ com(Fig. S1). These ratios are depleted arco et al. 1999, Heikoop et al. 2000, Swartmm other areas (Heikoop et al. 1998, Safrom colonies from S. siderastreaet al. 2005, Titlyanov et al. 2008) and strongly suggest, that up the introduced nitrogen influenced by the river, takingthe national park reefs are well possibly further deleterious things such as pounds (Roder et al. 2009), and with themcomheavy metals (Guzmán & Jimenez 1992, Guzmán & García 2002), common components
the ple in fungicides (Tabora et al. 1997), leading to a further weakening offor examont 2000). corals’ health (Harland & Brown 1989, Esslem

2

)‰N (151

dry season
rainy season

0W of peninsulaE of pensinsula
the reefs W and E Figure S1: Nitrogen isotopic ratios of of the peninsula withiSidern the Cahuastrea siita dereaNational (holobiont) saPark. mpDifferencesles co betweellected at 4 mn sites or se depth fromasons
ficant. gnisiare not

176

Chapter VI

LiteratureBeaton JD, Roberts TL, Halstagricultural commodities. In: Tiessen ead EH, Cowell LE (1995) GlH (ed) SCOPE obal t- ransfers Phosphorus in tof P ihne fertilizer mglobal enviaterials and ronment. John
Ltd., p 7-26 iley & Sons WClark I, Fritz P (1997) Environmental isotopes in hydrogeology, Vol. Lewis Publishers, Boca Raton
Cortés J (1993) A reef Global Aspects of Coral Reefs: under siltation stress:Health, a decade Hazaof rd adend Histgradation. ory. In: Rosenstiel School Ginsburg RN (ed) Colloquiof Marine and um on
Atmospheric Science, University of Miami, p 240-246
Esslemont G (2000) Heavy metals in seawater, marine sediments and corals from the Townsville section,
Great Barrier Reef Marine Park, Queensland. Marine Chemistry 71:215-231
Guzmán H, García EM (2002) Mercury levels in coral reefs along the Caribbean coast of Central America.
1415-1420 44:on Bulletin Marine PollutiGuzmán H, JiCentral Ammenez C (1992) Contamerica (Costa Rica ination and Panama). Marinof coral reefs by e Pollutiheavy mon Bulletin etals along t24:h554-561 e Caribbean coast of
Harland A, Brown BE (1989) Metal tolerance in the scleractinian coral Porites lutea. Marine Pollution
Bulletin 20:353-357 Heikoop JM, Dunn JJ, Risk MJ, Sandemann IM, Schwarcz HP, Waltho N (1998) Relationship between
light and the 15N of coral tissue: Examples from Jamaica and Zanzibar. Limnology and
909-920 Oceanography 43:Heikoop JM, Risk MJ, Lazier AV, Edinger EE, Jompa J, Limmon GV, Dunn JJ, Browne DR, Schwarcz HP
(2000) Nitrogen-15 signals of anthropogenic nutrient loading in reef corals. Marine Pollution
Bulletin 40:628-636 Hoegh-Guldberg O, Muscatine L, Goiran C, Siggaard D, Marion G (2004) Nutrient-induced perturbations
to 13C and 15N in symbiotic dinoflagellates and their coral hosts. Marine Ecology Progress
Series 280:105-114 Leichter JJ, Paytan A, Wankel S, Hanson K, Miller SL, Altabet MA (2007) Nitrogen and oxygen isotopic
signatures of subsurface nitrate seaward of the Florida Keys reef tract. Limnology and
McClelland JW, OceanographValiela I y 52:1258(1998) Linking -1267 nitrogen in estuarine producers to land-derived sources.
43:577-585 ographynology and OceanLimMcCutchan J, of carbon,William nitrogen and sulfur. M, Kendall C, McGrath C (2003) Oikos 102:378-390 Variation in trophic shift for stable isotope ratios
Roder C, Cortés J, Jimenez C, Lara R (2009) Riverine input of particulate material and inorganic nutrients
58:1937-194to a coastal reef ecosystem3 at the Caribbean coast of Costa Rica. Marine Pollution Bulletin
Sammarco PW, Risk MJ, Schwarcz HP, Heikoop JM (1999) Cross-continental shelf trends in coral 15N on
the Great Barrier Reef: further consideration of the reef nutrient paradox. Marine Ecology
131-138 180:Progress Series Seutin G, White BN, Boag PT (1991) Preservation of avian blood and tissue samples for DNA analyses.
Canadian Journal of Zoology 69:82-90 Sigmmeasureman DM, Altabet MA, Micheneent of the nitrogen isotopic comr R, McCorkle DC, Fry B, Holmposition of oceanic es RM (1997) nitrate: an adaptation Natural abundanceof the -level
Swartam PK, Saied mA, Lamonia diffusion meb K (2005) Temthod. Marine Chemporal and spatial variation iistry 57:227-242 n the 15N and 13C of coral tissue and
zooxanthellae in Montastrea faveolata collected from the Florida reef tract. Limnology and
Oceanography 50:1049-1058
Tabora P, fijiensisElango F, S) using hintani EM (effective M, Ufer C, microorVega Jganisms). (1997) Control of Proceedings of theblack si 5tgatokah Inte diseasernational C (Mycoosnferencephaere lla
on Kyusei Nature Farming, Bangkok, Thailand 11315
Titlyanov corals EA, Porites lKiyashko SIutea, T and P.itlyanova T cylindricV, a and in tKalita TL, Rahven JA (2008) eir epilithic and endolithic algae. MariC and N values ne Biin reef ology
155:353-361 Yamamuro M, in coral reKayanne ef ecH, Mosystemis. Limnagawa M nology and Oc(1995) Carbon and eanography 40:617-621 nitrogen stable isotopes of primary producers

177

1

- Chapter VII -

Chapter VII

Field observations and preliminary notes on the

status of NE-Hainan coral reefs

Cornelia Roder

1

, Zhongjie Wu

Leibniz Center for Tropical Marine Ecology, Z

Germany3Hainan Maritime and Fishery Depa

2

, Claudio Richter

3

T, Fahrenheitstrasse 6, 28359 Bremen, M

Hainan Maritime and Fishery Department and Hainan Provincial Marine Development

2Alfred Wegener InstituPlan and Design Research Institute, Haikou, Hainan (HPMDDIte for Polar- and Marine Research,), China Am Alten Hafen 26, 27568

Alfred Wegener Institute for Polar- and Marin

Bremerhaven, Germany

e Research,

In preparation

Am Alten Hafen 26, 27568

178

Abstract

Chapter VII

ical and biogeographic conditions for atic, geochemHainan’s coast provides unique climent of luxurious coral reefs in China. Observations in five reefs along the the developmNE coast of Hainan however showed that the overall densities of mobile macrofauna is
ng, scarping or excavating fish are missing low and key functional groups such as browsition or fission are abundant altogether. Coral diseases, partial mortality, tissue degradaand algal growth extensive. Signs of eutrophication, siltation and destructive fishing ent unfavorable for coral practices are evident resulting in a strongly altered environment success, e.g. larval settlemsurvival and recruitment. Even though corals that are still extant in the affected areas seem to have acclimated to the prevailing conditions, a shift
from a coral to algae dominated reef may occur if land-based disturbance prevails
unabated.

Introduction

Corals and algae are important primary producers co-occurring in tropical shallow water
reefs (Crossland et al. 1991; Hatcher 1997). Whilst they have to compete for space ; Lapointe et al. 1997), their co-existence is balanced by nutrient (Littler and Littler 1985petitive own control), whereas the com-up control) and grazing (top-dsupply (bottomore efficient utilization of inorganic nutrients is opposed by advantage of algae for a m ity. Changes in nutrient status and grazing can thereforeiltheir greater grazing susceptibinated reefs (Hughes 1994; Bellwood et al. 2004), have deleterious effects to coral domlay of direct and however rates and kinds of change are difficult to predict, as the interpinated communities are ex (Glynn 1988) and coral- or algal domplindirect effects is com; Pandolfi et al. only two out of several possible reef conditions (Bellwood et al. 20042005). Corals have been shown to persist in eutrophied waters, when sufficient grazing es, etimpressure on algae is present (Aronson and Precht 2001; Aronson et al. 2002). Somthe original grazers might not even be present any more, but replaced by other species
such as sea urchins (McClanahan and Muthiga 1988; Steneck 1998). They can in cases of

179

Chapter VII

e the only grazinextensive overfishing becom g control of algae (Ogden et al. 1973;

Levitan 1988) and may, when not antagonized by predators, become themselves a pest

anot only grazing on molagae, but finally also on corals and rock, and so eroding the rc

hen these grazers die off or migrate, they leave total reef base (Bellwood et al. 2004). W

barren substrate behind which may be the base for extensive algal growth due to their
at of corals (Hughes 1994; Gardner et al. ities compared to thent capabilefficient resettlem2003) and especially when coral recruitment is low (Aronson et al. 2002). Coral reefs
eans threatened by eutrophication together with overfishing which moreover may be by m

nent immof destructive techniques (McManus et al. 1997), are hence at high risk of iphase shift (Hughes 1994; Bellwood et al. 2004). the s. Descriptions fromThe coral reefs of Hainan are yet poorly investigated ecosystemuni1950s focus on distribution and diversity of coral commties (Yu and Zou 1996a,b), fects while later studies on coral diversity were conducted concerning anthropogenic ef

ic exploitation in (Yu and Zou 1996a,b; Shi and Zhang 2004) as the rapid economcombination with heavy fishing and land-based agricultural as well as aquacultural

developmreefs to the borderline of toleranent, both sources for heavy eutrophce. Studies from reefs on the souication and pollution, has taken the coral th coast have been
60% in 1983, to 41.5% in 1998, and to reporting decreases in live coral cover fromnd Zou 1996a,b), despite the foundation of the Sanya National 21.51% in 2002 (Yu a and seagrass monitoring project in 2002 Coral Reef Reserve in 1990. During a reefdistribution, biomass and recruitm(Status of China Marine Ecology Report, 2002), coral, seagraent was assessed along the east coast of Hainan. Thess and fish species

results revealed that live coral cover and coral recruitment, as well as coral reef
rco-algal abundance, ahile massociated fish and invertebrate density was very low, w

especially that of the brown algae Sargassum spp. was high (Hainan east coast coral reef
ation on the status of onitoring report, 2005). Therefore further informand seagrass mHainan’s coral reefs and their potential threats is urgently needed to provide the base for
a more sustainable coastal management, benefiting or even recovering adjacent coral
reefs.

180

Material and Methods

Chapter VII

In 2007 and 2008 field trips were conducted to the NE reef sites of Hainan (Fig. 1). Five

hang north to south): Tonggulin, Coconut Bay, Cpled (from be samreefs were chosen to

en. Photographs of site characteristics at each reef are given in qi gan, Longwan and Tanm

pling constraints, the status ofits along with same and logistic limFigs. 2-6. Due to tim

ent, thereby on a descriptive basis in rapid assessmeach reef could only be evaluated

ations of s, including the independent estimtaking pictures and notes on first impression

live coral cover by two divers. At each site, five corals were investigated in terms of

photosynthetic performance (see below, n = minimum of 3 per colony) and tissue

ical analyses were taken using ples (see below, n = 3 per colony) for biogeochemsam

alysed in situ with rapid light ance was an performhammer and chisel. Photosynthetic

curves (RLC, Ralph et al. 1999; Ralph and Gademann 2005) of the massive coral Porites

n 11 and 12 hrs of days with 0 % cloud e, 1860), betwee (Milne-Edwards and Haimlutea

cover as photosynthetic output changes with cloudiness or over day (Brown et al. 1999).

RCLs were conducted using a submersible pulse amplitude modulated fluorometer

ple holder (‘DIVING-any) with a universal sam(DivingPAM, Heinz Waltz Ltd., Germ

(Schreiber 1986). mUSH’) to assess the coral surface with a standardized distance of 1 c

g PAM applyings of the divinThereby, RLCs were conducted using the internal setting-ol quanta mincreasing light intensities (photosynthetic active radiation, PAR, 0-2896 μm-12s (ETR) (Ralph et al. 1999; Ralph and ) and recording electron transport rate s

Gademann 2005). Ambient light intensities were measured concomitantly with the light

easure of the present sensor of the DivingPAM. The resulting light curves are a good m

light history of the investigated colony nce and the short-termaphotosynthetic perform

' × PAR × 0.5 × ETR factor, (Schreiber et al. 1997). The ETR is calculated as F/Fm

ensionless value representing the yield, a dim/Fm' is the effective PSII quantumwhere F

photosynthetic effectiveness of photosystemtes for II in the given light state, 0.5 constitu

s (Ralph et al. o photosystembetween the twed equal distribution of electrons the assum

1999), and the ETR factor, reflecting the light absorbance by the sample, as being

unknown for P. lutea, was set to unity. Therefore only the commonly used relative ETR

parisons (rETR) (Hoegh-Guldberg and Jones 1999) could be obtained, still allowing com

181

Chapter VII

transported in ziplock plastic bags (< 3 mbetween different coral colonies of one species (Schreiber 1986). Chiseled saml residual water) to the laboratory for storage on ples were
ice and further processing. Furthermore at the reef site of Coconut Bay, three colonies of the branching species Pocillopora verrucosa (Ellis and Solander, 1986) were examined as
colony was taken for tissue analyses. ple per described above, but only one samrves or aminations of light cuior to exmplished prDescriptions of the reefs were accoents in order get unprejudiced records. analyses of collected fragmiting authority regulations, only few coral tissue parameters could be Due to liment determinations and dry weights of prising zooxanthellae density, pigmevaluated, coms of coral tissue were removed from the etercoral and zooxanthellae. Two square centimrush and filtered seawater. The tissue slurry was skeleton using an artist’s airb ogenized and aliquots retained for zooxanthellae density counts using asubsequently homcroscope (Leitz, Portugal, 260x ir and a meteFuchs-Rosenthal haemocytommagnification). 5 ml were filtered on Whatman glass fiber filters (F) for pigment analysis

using the standard procedure of (Strickland and Parsons 1972) and frozen for later

processing. For chlorophyll-a determinations filters where added with a certain volume of
ere ent concentrations wAcetone (90 %), incubated for 24h in the dark at 4° and pigmeter. Due to etrically (Lorenzen 1967) using a handphotomined spectrophotomdetermdissimilar storage times of the samples (up to one year), which may increase pheophytin
content in lo. 1995; Hill et al. 2000), not only chlorophyll-a ples (Hill et alnger stored sam

ined (using the acidification was determcontent but also the amount of pheophytin method of Lorenzen (1967)), and both summed up to obtain the initial chlorophyll-a

content of the coral tissue. Additionally, 15 ml of the slurry were centrifuged to separate
ponents from zooxanthellae (Muscatine et al. 1989; Swart et al. 2005) and coral host combusted glass fiber filters loaded on precomarately steps) sep(after several washing

(Whatman GF/F) for further desiccation and determination of coral host and

izooxanthellate dry weights on a mcrobalance (Mettler Toledo AB204-S, accuracy 0.1 mg).

182

IsHalainnadn

ynSaa

uokiHa

ne mnTaoaBo

eWnhanc

Chapter VII

Figure 1: Hindicating the five reef samainan Island located in thpling sites (squares), from N to S:e South China Sea and a close-up of the NE-coast Tonggulin, Coconut Bay,
en.Chang qi gan, Longwan and Tanm

Results

s of study sites Descriptionrelatively close to shore (200 – 1000 m)All fringing reefs along the NE-coast of Hainan are located in shallow waters (2 – 8 m and consist of a reef flat, crest and slope. The )
entary reefs inated, fragmfirst inspection of the working areas revealed at best algae domote live coral cover (often less than 1 %, d only remble fields anwith extensive coral rubsometimes up to 15 %), but higher and more divers cover (up to 60 %) of mainly small
colonies along the reef crests where wave and surge action was strongest resulting in
ents. Living colonies were often partly affected by disease, constant removal of sediml predation. In mortality or fission and often threatened by algal overgrowth or snaie latter acro-algae, the and mented a high abundance of corallincontrast, we documespecially on the reef slopes, but almost a total lack of mobile marcofauna, especially
s. Visibility was low (between 0.5 ll as echinodermeherbivorous and predatory fish, as wand 8 m)more detailed descriptio and coral rubble fields were in genen of each site is given in the following: ral covered with thick layers of sediment. A

183

Chapter VII

Tonggulin consisted of extensive coral )- The shallow reef flat of Tonggulin (2 – 5 m

ig. 2). The coral destructive bomb fishing (F ably originating fromrubble fields presum

oever, was already mrubble, howented together by encrusting algae, indicating stly cem

t happened recently. Life coral cover was low (5 – othat the destruction of the reef has n

pared to all 10 %), but increased to 40% on the reef crest and diversity was highest com

all branching species could be inly smaother reefs observed. In the area of low cover, m

observed, and diversity rose with increasing coverage. Algae were present, but were less

abundant than at other reef sites. Fish abundances was low and most individuals were

juvenile.

: Tonggulin Figure 2reef

g. 3) was likewise shallow (2 – 4 m)- The coral reef of Coconut Bay (FiCoconut Bay

. The widespread coral rubble fields on the reef flat, likely ) 400 mand close to shore (

originating from destructive fishing methods, were mainly covered and clotted by silt.

Fleshy green algae were abundant, but large inated the ens dom spp. specimSargassum

any of ssive coral colonies could be observed, maall individuals of marea. Only few sm

diseased, damthemd or encroached by algae. Close to the reef crest, where currents age

were stronger and waves present, the structure of the original reef still remained intact in

great parts and featured mainly branching coral species. There were only ecruits of mny ra

few fish, even along the reef crest.

184

t Bay reef nu: CocoFigure 3

- The reef area of Chang qi gan (Fig. 4) was further offshore (Chang qi gan

Chapter VII

800 m)

ed to be heavily fished, especially during 4 – 8 m. The site seemand depth ranged from

ounts of fishing boats in the reef area and audible as dry season as indicated by high am

well as visible reoccurring explosions. Few large massive Porites colonies several meters

in diameter still strengthening the damaged reef framework represented the main part of

live coral cover. However, many of them were affected by disease, partial mortality or in

petition with algae. Mucudirect comst of these colonies was high with os excretion in m

ent load. Branching ably to get rid of the sedimthick flats covering the colonies, presum

bers of fish recruits. Again, only low numall species could only be rarely found as sm

were present, meniles. ost of them being juv

Figure 4: Chang qi gan reef

185

Chapter VII

200 m) and depth did – In Longwan (Fig. 5) the reef was close to the shore (an Longwnot exceed 4 m. Coral rubble, cemented together by encrusting algae, dominated the reef
area. Further offshore, a mix of branching and foliose coral recruits and mid-sized
second highest (after ity in this reef was assive colonies built the reef framework. Diversm reef sites, however, live coral cover was low and mostTonggulin) of all investigated ounts, but not as abundant as in ent in high amall. Algae were prescolonies were rather smd en. As in all other reefs, only few juvenile fish coulCoconut Bay, Chang qi gan or Tanmbe observed.

reef : Longwan Figure 5

Tanmen- The reef of Tanmen (Fig. 6) further offshore ( 1000 m) than the others, was
located directly outside the river estuary and the Tanmen harbor. It was slightly deeper
and characterized by heavy algal growth. As in Chang qi )er reefs (5 – 8 mthan the othgan, the living part of the coral reef consisted of mainly massive boulder colonies
), which showed strong mucus secretion and were often affected Poritesinantly (predomechanical to pollution or mby disease. The intense ship traffic not only exposes the reefdestruction, but is likewise, regarding the many fishing boats and floating houses, ly, the reef lacked ccordingtensive fishing pressure in this area. Ar the exevidence foost completely any fishfauna. alm

186

Metabolic investigations

nmen reef : TaFigure 6

Chapter VII

of the investigated eters is given in Fig. 7. NoneAn overview of investigated tissue param

eters (coral host and zooxanthellae dry weight, zooxanthellae densities or pigmparament

concentrations) in samples from Porites lutea differed significantly (Kruskal-Wallis: p >

one colony or betwples from0.05) within sameen the five reef sites. Coral host dry

weights ranged from 0.3 (Coconut Bay) to 15.25 mg cm-2 (Tonggulin) with a mean of

6.25 ± 0.34 mg cm-2. Zooxanthellae dry weights were in the range of 3.85 ± 0.18 mg cm-2

with a minimum of 1.3 mg cm-2 in Coconut Bay and a maximum of 8.14 mg cm-2 in

-26 were counted, with highest zooxanthellae cmTonggulin. On average, 3.6 ± 1.6 * 10

abundances of 7.6 * 106 cm-2 in Tanmen and lowest abundances of just fewer than one

(chlorophyll-a) were lowest in Tonggulin ent concentrations illion in Tonggulin. Pigmm

(2.42 μg cm-2), but highest in samples from Coconut Bay (29.52 μg cm-2) and on average

14.27 ± 0.77 μg cm-2. The three samples of Pocillopora verrucosa collected in the reef of

Coconut Bay were similar in tissue composition to P. lutea samples, with coral host

ight being in the higher range, while zooxantellae dry weight, zooxanthellae etissue dry w

densities or pigment concentrations were in the lower range of the values computed for P.

. lutea

187

Chapter VII-2)15-2)15
1010mg cm (We Dust tishosmg cm (W Deussi tthos
5500 -2)15-2)15
1010lntheooxazmg cm (W Deal0 cmgm( Wae Dellthanoxzo0
55 88))-2-26 cm66 cm6
4401 (eallhentooxaz01 (eallhentooxaz
2200 3030))-2-22020mg cμ (tsnepigmmg cμ (stnegmpi
101000
Tanmen Longwan Tonggulin Coconut Bay igangqChan Coconut Bay
Figure 7: Host tissue and zooxanthellae dry weights, zooxanthellae densities and pigment concentrations of
Porites lutea from NE-Hainan reefs and from Poccilopora verrucosa from Coconut Bay.

188

Chapter VII

, strated by the rapid light curves (RLCsThe general photosynthetic performance as illureached only P. lutea cept Coconut Bay where colonies of ilar for all reefs exFig. 8) is simrETR maximum rates of < 100 (μmol electrons m-2s-1) and was less efficient under the
same irradiance levels. Most colonies (Porites lutea and Pocillopora verrucosa
specimens) reached a saturation plateau with maximum rETR of 50 – 150 μmol electrons
m-2s-1 at a photosynthetic active radiation (PAR) of about 1000 μmol quanta m-2 s-1.
While all curves stagnated at a certain saturation level or even decrease due to exposure ral colonies from bient max PAR intensities), comal irradiance (% over ato supra-optimit. Tonggulin also leveled out, but did not reach their saturation lim

400site: TG
)site: TM
-1site: LW
s300site: CG
-2site: CB
site: CB (P. verrucosa)
m-200 elom (μTRrE
100

00

00100020003000
PAR (μmol quanta m-2 s-1)
Figure 8TM: Tanm: Raen, LWpid light curves : Longwan, CG: Chaof Porites luteang qi colonies gan, CB: Cofromc varioous reenut Bay) and fs along fromNE-Haina Poccilopora ven (TG: rrucosa Tonggulin,
y. Coconut Baens fromspecim

189

Discussion

Chapter VII

The conducted surveys along the NE-coast of Hainan revealed a degraded status of coral

pact. This is in rtainly due to extensive anthropogenic imost cereefs in this area, alm

congruence with studies of You and Zou (1996 a, b), who described that the once

red flourishing coral reefs of Luhuitou, in the southern part of Hainan, have suffe

bined impacts of fishing, mcom-based sources of pollution. This includes ning and landi

ent run-off via rivers to coastal waters, construction and in particular severe sedim

an astline, comprising discharge of untreated hum land use near the cofalteration o

sewage, agriculture fertilizers and heavily polluted and eutrophied wastewater from land

based aquaculture during the last two decades. Furthermore coastal waters along the NE-

ainated with high loads of total suspended mcoast of Hainan are contamtter and nutrient -1 (TSM) and up 40 μM inorganic nitrogenousconcentrations, reaching up to 42 mg l

es of high precipitation (Krupp et al. g timpounds or 3 μM phosphate, especially durincomin prep.). The nitrogenous compounds suspended in the water column revealed high 15N

values and could accordingly be ascribed to shrimp ponds as nitrogen sources (Krupp et

al. in prep.). Highest amounts of TSM and nutrient loads were found at the reefs of

es of the river plumCoconut Bay and Tanmen (Krupp et al in prep.), directly outside the

aWenchang and the W visibility at the anifested by the lowngjuan river. This is further m

ent-loaded runoff and reefs, caused by the chronic nutrient- and sediminvestigated

entation (Fabricius reduced light availability and stress caused by sedimresulting in

acroalgae and the dark coloration of the 2005). The high abundance of thallose m

rs of reduced light levels and eutrophication. dicatoing corals are further inscattered liv

Additionally the extensive mucus flats excreted by many corals in the area are known to

be a cost-intensive technique of sedimacher 1977) or bio-fouling moval (Schuhment re

ral particle load protection (Rublee et al. (Ducklow and Mitchell 1979) as well as a gene

ishing and the use of destructive fishing an induced stresses are overf1980). Further hum

techniques (McManus et al. 1997). Altogether this indicates strong fertilization and

sdegradation of all reef sites along the NE-coast of Hainan a a result of an anthropogenic

. altered coastal system

190

Chapter VII

nthellae, even though not statistically asses of coral host tissue and zooxaBiomsignificant, are lowest in the most affected reefs and may indicate reduced health of the
zooxanthellae densities and chlorophyll present corals (Fabricius 2005), while concentrations are highest suggesting energetic costly adaptations to a low-light
lesias-Prieto and Trench 1994). This is ent (Falkowski and Dubinsky 1981; Igenvironm further supported by the fact that coral colonies from Coconut Bay had lowest parts) chronically exposed typical for corals (or colonyphotosynthetic capacities which is , the reef of Tonggulin gave to reduced light levels (Beer et al. 1998). On the other handost intact one in this area which is supported by highest pression of being the mthe imasses of coral host and zooxanthellae as well as by its enhanced photosynthetic biomperformance and the indication that specimens are adapted to higher light environments
her reefs. tpared to all ocomWhile zooxanthellae densities of Porites lutea as well as Pocillopora verrucosa are in the
range of those shown for the samand Daya Bay (Guangdong Province, South e species China main Chinese waters of Sanya (South Hainan) inland) (Li et al. 2008), the
saturation potentials are in the lower range investigated photosynthetic performance with of those reported (i.e. Beer et al. 1998; Ralph et al. 1999). The latter incidence dissents fromefficiencies of deep or turbid water, i.e. low ligth specim the fact that all corals were shallow water specimens (< 6 mens (Ig)lesias-Prieto, but rather resembled and Trench
1994; Jantzen et al. 2008). ll outlive these unfavorable conditions have s that corals that stiIn general it seemntioned eutrophication and reduced light levels by adjusting earranged to respond to the mtheir metabolic performance to the given conditions. Nevertheless, it can further be
expected that their fitness is momentarily impacted to a degree where their own survival
e energetic costs t, if conditions prevail or even deteriorate, thuay still be possible, bmnecessary to keep pace with the disturbance are too high and accordingly may result in
mortality of the coral community (Kirkwood 1992). all sized Hainan are the frequently noticed smA glimmer of hope for the NE coral reefs of coral colonies, mainly occurring along the reef crests which may signify a high recovery
being the mpotential (Daustan 1977), however, their survival min substrate, especially on the reef flats, poses unfavorable conditions foray be uncertain because coral rubble,

191

Chapter VII

1990) as it is easily turned by waves or persistence (Hodgson settling and longer-termely unstabsurge and hence extremle. It appears that in the subsequent decades, particularly during the rapid economic
rine aan activities have taken their toll on the coastal ment in the 90’s, humdevelopment and at present all coral reefs along the NE-coast of Hainan are suffering environm sease to greater or lesser extent. The eutrophication, siltation as well as difromss indicates that reproduction is existent and subsistence of scleractinian corals nevertheleat least in parts the top-down control of algae still holds and that reproduction is existent. Data on the abundance and grazing activity of herbivores or on coral reproduction is the interactive portance to gain insights onhowever wanting. Obviously, the imand invertebrate grazerconsequences of high nutrient and sedim abundances and the altered environment inputs from land-based runofent due to destructive fishing f, the low fish
on coral health and reef status is crucial in the area to implement proper management
ral reefs of NE Hainan. strategies for the co

ledgements nowAk

Research was carried out within the frame of the bilateral Sino-German research project
an Federal Ministry of Education and Research LANCET funded jointly by the Germ Education and the Hainan Provincial Marine and (BMBF), the Chinese Ministry ofent. Fishery Departm

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194

Discussion

Discussion

of the natural disturbance of Large Amplitude onstrated the characteristicsThis thesis has dem

Internal W) (Chapters I to V) and of the anthropogenic disturbances of river ves (LAIWa

ent and bined effects of destructive fishing, coastal developmpter VI) and comrunoff (Cha

aquaculture (Chapter VIII). Herein, the responses of corals to the respective disturbance have

been investigated.

introduce cold, oxygen depleted, low pH and nutrient rich water into the reefWLAI

ecosystem of the Similan Islands, Thailand, located close to the continental shelf break in the

Andaman Sea. Even though some LAIW energy is also found to wrap around the islands to

the east (E) side, the main im) ced by the deep reef sites along the west (Wpact is experien

coasts.

It is shown that corals along the W reef sides of the Similan Islands are strongly influenced by

the prevailing LAIW conditions (Chapter I). The amount of impact is regulated by the

frequency and the intensity of the LAIW and differs on spatial (W deep highly influenced >

shallow > E deep > E shallow least influenced) as well as on temporal scales (yearly W

tween equally exposed reefs Chapter I), however never bedifferences between 2007 and 2008,

is also highly species specific with tial of acclimatizationof the various islands. The poten

some species (such as D. heliopora) responding stronger to LAIW impact (Chapter I).

(Muscatine et al. 1989) centrations in corals ent conThe high input of nutrients fuels pigm

in situ (Chapter IV) between ilar ary production is simral prim(Chapter I, IV). However, co

when incubated under equal en lower in W and sheltered E side corals, and evexposed W

stly photosynthesis due to acclimconditions (Chapter III) indicating an energy coatization to

ilans reefs of the Sim the Wlow light levels (Iglesias-Prieto and Trench 1994). Corals from

also feature higher biomasses and protein contents, most likely due to higher heterotrophic

input (Bachar et al. 2007; Rodrigues et al. 2008) as a result of strong currents (Sebens and

Johnson 1991) and increased organic matter fluxes (Chapter I, II). The combination of photo-

and heterotrophy enhances the energyand increases their resilience side corals stores of the W

potential to disturbance (Chapter II).

ght also be the trigger for i side reefs m the WThe high availability of nutrients and organics in

aa less conservative behavior in retaining organic mterial in dissolved or particulate form

(Ferrier-Pagès et al. 1998) leading to an increased release of dissolved organics or mucus

ore poorer calcifiers com impact are furthermW(Chapter III). Corals under LAIpared to their

195

Discussion

low pH water (Marubini et alsheltered E counterparts (Chapter III) which m. 2001) or acclimiatization to loght be attributed to the frw light levels (Crossland 1984). equent exposure to

ilar, however with contrary ary production in W and E side reefs is simThe gross priments are stronger producers in the ents. While the sedimpartmcontributions of the various comost likely due influence mWary production under LAIE, turf algae contribute stronger to primto elevated nutrient availability (Hatcher and Larkum 1983) with increasing LAIW impact
pared to E (Chapter III). comand due to their higher abundance on W

ght be in large parts ibined effects of low coral photosynthesis and calcification mThe comation, especially in the deeper reef waters along the responsible for the reduced reef formwestern Similan Islands (Chapter V). While the E reefs form a dense and complex framework,
ensional structure side is characterized by scattered corals lacking a true three-dimthe W(Chapter V). While in the shallow waters of the W the altered morphologies of corals growing
pact monsoon imWbe explained by a strong Sclosely attached to the granite boulders can W ework in deeper water is attributed to LAIm(Nielsen et al. 2004), the lack of reef fra(Chapter V). There is indication, that the potential of the coral species to respond to the ines its fate in abundance along the ent of LAIW determical environmstressful physico-chement and ilan Islands (Chapter I). Those able to exploit the benefits of nutrient enrichmSimorganic matter fluxes (Chapter I) are more likely to colonize the W Similan reefs.

What remains to be determined is whether the responses to disturbance that we see in LAIW
ely costly photosynthesis and reduced calcification on the one hand, but affected corals, nam of emincreased energy uptake and release and a higher resilience on the other, is within the fragenotypic acclimatization or if it might even be genetic adaptation. The reefs of the Similan
Islands provide a clue to understanding coral resilience in the face of climate change and
atization to pulsed low versatile acclime their successful andification duocean acidperature, low oxygen, low pH, and high nutrient conditions. tem

Costa Rica (Chapter VI) and Hainan (Chapter The situation is different in the examples fromVII), where the reefs are exposed to chronic and not pulsed disturbance. The main impacts of
entation (Anthony 2006) at either investigation site ant 2002) and sedimeutrophication (Szmdrive the ecosystemCaribbean coast of Costa Rica (Chapter VI) sus to their borderlines. Whffers heavy sedimile the reef of the National Park Cahuita on the entation and is at risk of
eber et al. 2006), the extensive algal growth along the NE-Hainan coast being suffocated (W (Lapointe 1997). The inated ecosystemgal dom(Chapter VII) indicates a shift towards an al

196

Discussion

causes of disturbance are obvious: excessive land use of the Caribbean hinterland has altered

llution. The eutrophication gradient the coastline leading to strong erosion and fertilizer po

pact is given by eably and evidence on direct im river mouth to coral reef is clearly tracfrom

the active urived nitrogen by the corals of the national park reef (Chapter take of fertilizer dep

shing is visible by widespread Hainan, destruction from explosive fiVI). In the reefs of NE-

fields of coral rubble and lack of coral as well as fish fauna. The turbid water colum

n has

g to decreased coral strongly reduced photosynthetic efficiency of the corals possibly leadin

growth (Rodolfo-Metalpa et al. 2006) that mght not be able to keep up with the rate of i

destruction. In Costa Rica as well as on Hainan, it is crucial to develop and enforce proper

ain high or even further increase. conservation strategies, as the disturbances are likely to rem

Coral reef ecosystems are very complex, nevertheless extremely flexible systems and due to

their high productivity and biodiversity of high conservation value. Gaining information on

coral reef functioning under disturbances is important to assess necessary m

ent nagema

g world. This thesis inplications and to predict future coral reef scenarios in a rapidly changim

contributes to the baseline of understanding reef

ents. corals in disturbed environm

functioningtentials of atization po and acclim

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198

Disclaimer

Gemäß §6 der Promotionsordnung der Universität Bremen für die

Disclaime

chbereiche vommathematischen, natur- und ingenieurwissenschaftlichen Fa

here ich, dass: 14. März 2007 versic

1.

2.

3.

e Hilfe angefertigt wurde die Arbeit ohne unerlaubte fremd

ittel benutzt keine anderen als die angegebenen Quellen und Hilfsm

wurden

die den benutzten Werken wörtlich oder inhaltlich entnomm

Stellen als solche kenntlich gemacht wurden

Bremen, 20. Februar 2009

__________________________

Cornelia Roder

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r