Coral Reef - Wikipedia, The Free Encyclopedia

12/21/2014
CoralreefWikipedia,thefreeencyclopedia
Coralreef
FromWikipedia,thefreeencyclopedia
Coralreefsarediverseunderwaterecosystemsheldtogetherbycalciumcarbonatestructuressecretedbycorals.Coralreefsarebuiltbycoloniesoftinyanimalsfoundinmarinewaters
thatcontainfewnutrients.Mostcoralreefsarebuiltfromstonycorals,whichinturnconsistofpolypsthatclusteringroups.ThepolypsbelongtoagroupofanimalsknownasCnidaria,
whichalsoincludesseaanemonesandjellyfish.Unlikeseaanemones,coralssecretehardcarbonateexoskeletonswhichsupportandprotectthecoralpolyps.Reefsgrowbestinwarm,
shallow,clear,sunnyandagitatedwaters.
Oftencalled"rainforestsofthesea",coralreefsformsomeofthemostdiverseecosystemsonEarth.Theyoccupylessthan0.1%oftheworld'soceansurface,abouthalftheareaof
France,yettheyprovideahomefor25%ofallmarinespecies,[1][2][3]includingfish,mollusks,worms,crustaceans,echinoderms,sponges,tunicatesandothercnidarians.[4]
Paradoxically,coralreefsflourisheventhoughtheyaresurroundedbyoceanwatersthatprovidefewnutrients.Theyaremostcommonlyfoundatshallowdepthsintropicalwaters,but
deepwaterandcoldwatercoralsalsoexistonsmallerscalesinotherareas.
Coralreefsdeliverecosystemservicestotourism,fisheriesandshorelineprotection.TheannualglobaleconomicvalueofcoralreefsisestimatedbetweenUS$29.8375billion.
However,coralreefsarefragileecosystems,partlybecausetheyareverysensitivetowatertemperature.Theyareunderthreatfromclimatechange,oceanicacidification,blastfishing,
cyanidefishingforaquariumfish,sunscreenuse,[5]overuseofreefresources,andharmfullandusepractices,includingurbanandagriculturalrunoffandwaterpollution,whichcan
harmreefsbyencouragingexcessalgalgrowth.[6][7][8]
Contents
1Formation
1.1Materials
2Types
3Zones
4Locations
5Biology
6Darwin'sparadox
6.1Explanations
7Biodiversity
7.1Algae
http://en.wikipedia.org/wiki/Coral_reef
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7.2Sponges
7.3Fish
7.4Invertebrates
7.5Seabirds
7.6Other
8Importance
9Threats
10Protection
11Restoration
12Reefsinthepast
13Seealso
14References
15Furtherreferences
16Externallinks
Formation
Mostofthecoralreefswecanseetodaywereformedafterthelastglacialperiodwhenmeltingicecausedthesealeveltoriseandfloodthecontinentalshelves.Thismeansthatmost
moderncoralreefsarelessthan10,000yearsold.Ascommunitiesestablishedthemselvesontheshelves,thereefsgrewupwards,pacingrisingsealevels.Reefsthatrosetooslowly
couldbecomedrownedreefs.Theyarecoveredbysomuchwaterthattherewasinsufficientlight.[9]Coralreefsarefoundinthedeepseaawayfromcontinentalshelves,aroundoceanic
islandsandasatolls.Thevastmajorityoftheseislandsarevolcanicinorigin.Thefewexceptionshavetectonicoriginswhereplatemovementshaveliftedthedeepoceanflooronthe
surface.
In1842inhisfirstmonograph,TheStructureandDistributionofCoralReefs[10]CharlesDarwinsetouthistheoryoftheformationofatollreefs,anideaheconceivedduringthe
voyageoftheBeagle.HetheorizedupliftandsubsidenceoftheEarth'scrustundertheoceansformedtheatolls.[11]Darwinstheorysetsoutasequenceofthreestagesinatoll
formation.Itstartswithafringingreefformingaroundanextinctvolcanicislandastheislandandoceanfloorsubsides.Asthesubsidencecontinues,thefringingreefbecomesabarrier
reef,andultimatelyanatollreef.
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Darwinstheorystarts
withavolcanicisland
whichbecomesextinct
Astheislandand
oceanfloorsubside,
coralgrowthbuildsa
fringingreef,often
includingashallow
lagoonbetweenthe
landandthemainreef.
Asthesubsidence
continues,thefringing
reefbecomesalarger
barrierreeffurther
fromtheshorewitha
biggeranddeeper
lagooninside.
Ultimately,theisland
sinksbelowthesea,
andthebarrierreef
becomesanatoll
enclosinganopen
lagoon.
Darwinpredictedthatunderneatheachlagoonwouldbeabedrockbase,theremainsoftheoriginalvolcano.Subsequentdrillingprovedthiscorrect.Darwin'stheoryfollowedfromhis
understandingthatcoralpolypsthriveinthecleanseasofthetropicswherethewaterisagitated,butcanonlylivewithinalimiteddepthrange,startingjustbelowlowtide.Wherethe
leveloftheunderlyingearthallows,thecoralsgrowaroundthecoasttoformwhathecalledfringingreefs,andcaneventuallygrowoutfromtheshoretobecomeabarrierreef.
Wherethebottomisrising,fringingreefscangrowaroundthecoast,butcoralraisedabovesealeveldiesandbecomeswhitelimestone.Ifthe
landsubsidesslowly,thefringingreefskeeppacebygrowingupwardsonabaseofolder,deadcoral,formingabarrierreefenclosingalagoon
betweenthereefandtheland.Abarrierreefcanencircleanisland,andoncetheislandsinksbelowsealevelaroughlycircularatollofgrowing
coralcontinuestokeepupwiththesealevel,formingacentrallagoon.Barrierreefsandatollsdonotusuallyformcompletecircles,butare
brokeninplacesbystorms.Likesealevelrise,arapidlysubsidingbottomcanoverwhelmcoralgrowth,killingthecoralpolypsandthereef,due
towhatiscalledcoraldrowning.[13]Coralsthatrelyonzooxanthellaecandrownwhenthewaterbecomestoodeepfortheirsymbiontsto
adequatelyphotosynthesize,duetodecreasedlightexposure.[14]
Thetwomainvariablesdeterminingthegeomorphology,orshape,ofcoralreefsarethenatureoftheunderlyingsubstrateonwhichtheyrest,and
thehistoryofthechangeinsealevelrelativetothatsubstrate.
Theapproximately20,000yearoldGreatBarrierReefoffersanexampleofhowcoralreefsformedoncontinentalshelves.Sealevelwasthen
120m(390ft)lowerthaninthe21stcentury.[15][16]Assealevelrose,thewaterandthecoralsencroachedonwhathadbeenhillsofthe
Australiancoastalplain.By13,000yearsago,sealevelhadrisento60m(200ft)lowerthanatpresent,andmanyhillsofthecoastalplainshad
Afringingreefcantaketenthousand
yearstoform,andanatollcantake
upto30millionyears. [12]
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becomecontinentalislands.Asthesealevelrisecontinued,watertoppedmostofthecontinentalislands.Thecoralscouldthenovergrowthehills,formingthepresentcaysandreefs.
SealevelontheGreatBarrierReefhasnotchangedsignificantlyinthelast6,000years,[16]andtheageofthemodernlivingreefstructureisestimatedtobebetween6,000and8,000
years.[17]AlthoughtheGreatBarrierReefformedalongacontinentalshelf,andnotaroundavolcanicisland,Darwin'sprinciplesapply.Developmentstoppedatthebarrierreefstage,
sinceAustraliaisnotabouttosubmerge.Itformedtheworld'slargestbarrierreef,3001,000m(9803,300ft)fromshore,stretchingfor2,000km(1,200mi).[18]
Healthytropicalcoralreefsgrowhorizontallyfrom1to3cm(0.39to1.2in)peryear,andgrowverticallyanywherefrom1to25cm(0.39to9.8in)peryearhowever,theygrowonly
atdepthsshallowerthan150m(490ft)becauseoftheirneedforsunlight,andcannotgrowabovesealevel.[19]
Materials
Asthenameimplies,thebulkofcoralreefsismadeupofcoralskeletonsfrommostlyintactcoralcolonies.Asotherchemicalelementspresentincoralsbecomeincorporatedintothe
calciumcarbonatedeposits,aragoniteisformed.However,shellfragmentsandtheremainsofcalcareousalgaesuchasthegreensegmentedgenusHalimedacanaddtothereef'sability
towithstanddamagefromstormsandotherthreats.SuchmixturesarevisibleinstructuressuchasEniwetokAtoll.[20]
Types
Thethreeprincipalreeftypesare:
Fringingreefdirectlyattachedtoashore,orbordersitwithaninterveningshallowchannelorlagoon
Barrierreefreefseparatedfromamainlandorislandshorebyadeepchannelorlagoon
Atollreefmoreorlesscircularorcontinuousbarrierreefextendsallthewayaroundalagoonwithoutacentralisland
Otherreeftypesorvariantsare:
Patchreefcommon,isolated,comparativelysmallreefoutcrop,usuallywithinalagoonorembayment,oftencircularandsurroundedby
sandorseagrass
Apronreefshortreefresemblingafringingreef,butmoreslopedextendingoutanddownwardfromapointorpeninsularshore
Bankreeflinearorsemicircularshapedoutline,largerthanapatchreef
Ribbonreeflong,narrow,possiblywindingreef,usuallyassociatedwithanatolllagoon
Tablereefisolatedreef,approachinganatolltype,butwithoutalagoon
AsmallatollintheMaldives
HabilireefspecifictotheRedSeadoesnotreachthesurfacenearenoughtocausevisiblesurfmaybeahazardtoships(fromthe
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Arabicfor"unborn")
Microatollcommunityofspeciesofcoralsverticalgrowthlimitedbyaveragetidalheightgrowthmorphologiesofferalowresolution
recordofpatternsofsealevelchangefossilizedremainscanbedatedusingradioactivecarbondatingandhavebeenusedtoreconstruct
Holocenesealevels[21]
Cayssmall,lowelevation,sandyislandsformedonthesurfaceofcoralreefsfromerodedmaterialthatpilesup,forminganareaabove
sealevelcanbestabilizedbyplantstobecomehabitableoccurintropicalenvironmentsthroughoutthePacific,AtlanticandIndian
Oceans(includingtheCaribbeanandontheGreatBarrierReefandBelizeBarrierReef),wheretheyprovidehabitableandagriculturalland
Seamountorguyotformedwhenacoralreefonavolcanicislandsubsidestopsofseamountsareroundedandguyotsareflatflattops
ofguyots,ortablemounts,areduetoerosionbywaves,winds,andatmosphericprocesses
InhabitedcayintheMaldives
Zones
Coralreefecosystemscontaindistinctzonesthatrepresentdifferentkindsofhabitats.Usually,threemajorzonesarerecognized:the
forereef,reefcrest,andthebackreef(frequentlyreferredtoasthereeflagoon).
Allthreezonesarephysicallyandecologicallyinterconnected.Reeflifeandoceanicprocessescreateopportunitiesforexchangeof
seawater,sediments,nutrients,andmarinelifeamongoneanother.
Thus,theyareintegratedcomponentsofthecoralreefecosystem,eachplayingaroleinthesupportofthereefs'abundantand
diversefishassemblages.
Thethreemajorzonesofacoralreef:theforereef,
reefcrest,andthebackreef
Mostcoralreefsexistinshallowwaterslessthan50mdeep.Someinhabittropicalcontinentalshelveswherecool,nutrientrich
upwellingdoesnotoccur,suchasGreatBarrierReef.Othersarefoundinthedeepoceansurroundingislandsorasatolls,suchasin
theMaldives.Thereefssurroundingislandsformwhenislandssubsideintotheocean,andatollsformwhenanislandsubsidesbelow
thesurfaceofthesea.
Alternatively,MoyleandCechdistinguishsixzones,thoughmostreefspossessonlysomeofthezones.[22]
Thereefsurfaceistheshallowestpartofthereef.Itissubjecttothesurgeandtheriseandfalloftides.Whenwavespassovershallowareas,theyshoal,asshowninthediagramatthe
right.Thismeansthewaterisoftenagitated.Thesearethepreciseconditionunderwhichcoralsflourish.Shallownessmeansthereisplentyoflightforphotosynthesisbythesymbiotic
zooxanthellae,andagitatedwaterpromotestheabilityofcoraltofeedonplankton.However,otherorganismsmustbeabletowithstandtherobustconditionstoflourishinthiszone.
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Theoffreefflooristheshallowseafloorsurroundingareef.Thiszoneoccursbyreefsoncontinentalshelves.Reefsaroundtropicalislandsandatollsdropabruptlytogreatdepths,
anddonothaveafloor.Usuallysandy,theflooroftensupportsseagrassmeadowswhichareimportantforagingareasforreeffish.
Thereefdropoffis,foritsfirst50m,habitatformanyreeffishwhofindshelterontheclifffaceandplanktoninthewaternearby.Thedropoffzoneappliesmainlytothereefs
surroundingoceanicislandsandatolls.
Thereeffaceisthezoneabovethereeffloororthereefdropoff.Thiszoneisoftenthemostdiverseareaofthereef.Coralandcalcareousalgae
growthsprovidecomplexhabitatsandareaswhichofferprotection,suchascracksandcrevices.Invertebratesandepiphyticalgaeprovidemuch
ofthefoodforotherorganisms.[22]
Thereefflatisthesandybottomedflat,whichcanbebehindthemainreef,containingchunksofcoral.Thiszonemayborderalagoonandserve
asaprotectivearea,oritmayliebetweenthereefandtheshore,andinthiscaseisaflat,rockyarea.Fishestendtopreferlivinginthatflat,rocky
area,comparedtoanyotherzone,whenitispresent.[22]
Thereeflagoonisanentirelyenclosedregion,whichcreatesanarealessaffectedbywaveactionthatoftencontainssmallreefpatches.[22]
However,the"topographyofcoralreefsisconstantlychanging.Eachreefismadeupofirregularpatchesofalgae,sessileinvertebrates,andbare
rockandsand.Thesize,shapeandrelativeabundanceofthesepatcheschangesfromyeartoyearinresponsetothevariousfactorsthatfavorone
typeofpatchoveranother.Growingcoral,forexample,producesconstantchangeinthefinestructureofreefs.Onalargerscale,tropicalstorms
mayknockoutlargesectionsofreefandcausebouldersonsandyareastomove."[23]
Locations
Waterinthereefsurfacezoneisoften
agitated.Thisdiagramrepresentsa
reefonacontinentalshelf.Thewater
wavesatthelefttravelovertheoff
reefflooruntiltheyencounterthereef
slopeorforereef.Thenthewaves
passovertheshallowreefcrest.When
awaveentersshallowwateritshoals,
thatis,itslowsdownandthewave
heightincreases.
Coralreefsareestimatedtocover284,300km2(109,800sqmi),[24]justunder0.1%oftheoceans'surfacearea.TheIndoPacific
region(includingtheRedSea,IndianOcean,SoutheastAsiaandthePacific)accountfor91.9%ofthistotal.SoutheastAsiaaccounts
for32.3%ofthatfigure,whilethePacificincludingAustraliaaccountsfor40.8%.AtlanticandCaribbeancoralreefsaccountfor
7.6%.[2]
Althoughcoralsexistbothintemperateandtropicalwaters,shallowwaterreefsformonlyinazoneextendingfrom30Nto30S
oftheequator.Tropicalcoralsdonotgrowatdepthsofover50meters(160ft).Theoptimumtemperatureformostcoralreefsis26
Locationsofcoralreefs
[25]
27C(7981F),andfewreefsexistinwatersbelow18C(64F). However,reefsinthePersianGulfhaveadaptedto
temperaturesof13C(55F)inwinterand38C(100F)insummer.[26]Thereare37speciesofscleractiniancoralsidentifiedinsuchharshenvironmentaroundLarakIsland.[27]
Deepwatercoralcanexistatgreaterdepthsandcoldertemperaturesatmuchhigherlatitudes,asfarnorthasNorway.[28]Althoughdeepwatercoralscanformreefs,verylittleisknown
aboutthem.
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CoralreefsarerarealongthewestcoastsoftheAmericasandAfrica,dueprimarilytoupwellingandstrongcoldcoastalcurrentsthatreducewatertemperaturesintheseareas
(respectivelythePeru,BenguelaandCanarystreams).[29]CoralsareseldomfoundalongthecoastlineofSouthAsiafromtheeasterntipofIndia(Chennai)totheBangladeshand
Myanmarborders[2]aswellasalongthecoastsofnortheasternSouthAmericaandBangladesh,duetothefreshwaterreleasefromtheAmazonandGangesRiversrespectively.
TheGreatBarrierReeflargest,comprisingover2,900individualreefsand900islandsstretchingforover2,600kilometers
(1,600mi)offQueensland,Australia
TheMesoamericanBarrierReefSystemsecondlargest,stretching1,000kilometers(620mi)fromIslaContoyatthetipof
theYucatnPeninsuladowntotheBayIslandsofHonduras
TheNewCaledoniaBarrierReefsecondlongestdoublebarrierreef,covering1,500kilometers(930mi)
TheAndros,BahamasBarrierReefthirdlargest,followingtheeastcoastofAndrosIsland,Bahamas,betweenAndrosand
Nassau
TheRedSeaincludes6000yearoldfringingreefslocatedarounda2,000km(1,240mi)coastline
TheFloridaReefTractlargestcontinentalUSreef,extendsfromSoldierKey,locatedinBiscayneBay,totheDryTortugas
intheGulfofMexico[30]
Boundaryfor20Cisotherms.Mostcoralslive
withinthisboundary.Notethecoolerwaterscaused
byupwellingonthesouthwestcoastofAfricaand
offthecoastofPeru.
PulleyRidgedeepestphotosyntheticcoralreef,Florida
NumerousreefsscatteredovertheMaldives
ThePhilippinescoralreefarea,thesecondlargestinSoutheastAsia,isestimatedat26,000squarekilometersandholdsan
extraordinarydiversityofspecies.Scientistshaveidentified915reeffishspeciesandmorethan400scleractiniancoral
species,12ofwhichareendemic.
TheRajaAmpatIslandsinIndonesia'sWestPapuaprovinceofferthehighestknownmarinediversity.[31]
Biology
Livecoralarecoloniesofsmallanimalsembeddedincalciumcarbonateshells.Itisamistaketothinkofcoralasplantsorrocks.
Coralheadsconsistofaccumulationsofindividualanimalscalledpolyps,arrangedindiverseshapes.[32]Polypsareusuallytiny,but
theycanrangeinsizefromapinheadto12inches(30cm)across.
Thismapshowsareasofupwellinginred.Coral
reefsarenotfoundincoastalareaswherecolder
andnutrientrichupwellingsoccur.
Reefbuildingorhermatypiccoralsliveonlyinthephoticzone(above50m),thedepthtowhichsufficientsunlightpenetratesthe
water,allowingphotosynthesistooccur.Coralpolypsdonotphotosynthesize,buthaveasymbioticrelationshipwithmicroscopicalgaeofthegenusSymbiodinium,commonlyreferred
toaszooxanthellae.Theseorganismslivewithinthetissuesofpolypsandprovideorganicnutrientsthatnourishthepolyp.Becauseofthisrelationship,coralreefsgrowmuchfasterin
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clearwater,whichadmitsmoresunlight.Withouttheirsymbionts,coralgrowthwouldbetooslowtoformsignificantreefstructures.Coralsgetupto90%oftheirnutrientsfromtheir
symbionts.[33]
Reefsgrowaspolypsandotherorganismsdepositcalciumcarbonate,[34][35]thebasisofcoral,asaskeletalstructurebeneathandaround
themselves,pushingthecoralhead'stopupwardsandoutwards.[36]Waves,grazingfish(suchasparrotfish),seaurchins,sponges,andother
forcesandorganismsactasbioeroders,breakingdowncoralskeletonsintofragmentsthatsettleintospacesinthereefstructureorformsandy
bottomsinassociatedreeflagoons.Manyotherorganismslivinginthereefcommunitycontributeskeletalcalciumcarbonateinthesame
manner.[37]Corallinealgaeareimportantcontributorstoreefstructureinthosepartsofthereefsubjectedtothegreatestforcesbywaves(suchas
thereeffrontfacingtheopenocean).Thesealgaestrengthenthereefstructurebydepositinglimestoneinsheetsoverthereefsurface.
Typicalshapesforcoralspeciesarewrinkledbrains,cabbages,tabletops,antlers,wirestrandsandpillars.Theseshapescandependonthelife
historyofthecoral,likelightexposureandwaveaction,[38]andeventssuchasbreakages.[39]
Coralsreproducebothsexuallyandasexually.Anindividualpolypusesbothreproductivemodeswithinits
lifetime.Coralsreproducesexuallybyeitherinternalorexternalfertilization.Thereproductivecellsarefound
onthemesenterymembranesthatradiateinwardfromthelayeroftissuethatlinesthestomachcavity.Some
matureadultcoralsarehermaphroditicothersareexclusivelymaleorfemale.Afewspecieschangesexasthey
grow.
Internallyfertilizedeggsdevelopinthepolypforaperiodrangingfromdaystoweeks.Subsequentdevelopment
Anatomyofacoralpolyp
producesatinylarva,knownasaplanula.Externallyfertilizedeggsdevelopduringsynchronizedspawning.
Polypsreleaseeggsandspermintothewaterenmasse,simultaneously.Eggsdisperseoveralargearea.The
Tablecoral
timingofspawningdependsontimeofyear,watertemperature,andtidalandlunarcycles.Spawningismost
successfulwhenthereislittlevariationbetweenhighandlowtide.Thelesswatermovement,thebetterthechanceforfertilization.Idealtimingoccurs
inthespring.Releaseofeggsorplanulausuallyoccursatnight,andissometimesinphasewiththelunarcycle(threetosixdaysafterafullmoon).Theperiodfromreleaseto
settlementlastsonlyafewdays,butsomeplanulaecansurviveafloatforseveralweeks.Theyarevulnerabletopredationandenvironmentalconditions.Theluckyfewplanulaewhich
successfullyattachtosubstratenextconfrontcompetitionforfoodandspace.
ThereareeightcladesofSymbiodiniumphylotypes.MostresearchhasbeencompletedontheSymbiodiniumcladesAD.Eachoneoftheeightcontributestheirownbenefitsaswellas
lesscompatibleattributestothesurvivaloftheircoralhosts.Eachphotosyntheticorganismhasaspecificlevelofsensitivitytophotodamageofcompoundsneededforsurvival,suchas
proteins.Ratesofregenerationandreplicationdeterminetheorganism'sabilitytosurvive.PhylotypeAisfoundmoreintheshallowregionsofmarinewaters.Itisabletoproduce
mycosporinelikeaminoacidsthatareUVresistant,usingaderivativeofglycerintoabsorbtheUVradiationandallowingthemtobecomemorereceptivetowarmerwater
temperatures.IntheeventofUVorthermaldamage,ifandwhenrepairoccurs,itwillincreasethelikelihoodofsurvivalofthehostandsymbiont.Thisleadstotheideathat,
evolutionarily,cladeAismoreUVresistantandthermallyresistantthantheotherclades.[40]
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CladesBandCarefoundmorefrequentlyinthedeeperwaterregions,whichmayexplainthehighersusceptibilitytoincreasedtemperatures.
TerrestrialplantsthatreceivelesssunlightbecausetheyarefoundintheundergrowthcanbeanalogizedtocladesB,C,andD.SincecladesBthrough
Darefoundatdeeperdepths,theyrequireanelevatedlightabsorptionratetobeabletosynthesizeasmuchenergy.Withelevatedabsorptionratesat
UVwavelengths,thedeeperoccurringphylotypesaremorepronetocoralbleachingversusthemoreshallowclades.CladeDhasbeenobservedtobe
hightemperaturetolerant,andasaresultithasahigherrateofsurvivalthancladesBandC.[40]
Closeupofpolypsarearrayed
onacoral,wavingtheir
tentacles.Therecanbe
thousandsofpolypsona
singlecoralbranch.
Braincoral
Staghorncoral
Spiralwire
coral
Pillarcoral
Mushroomcoral
Mazecoral
Blackcoral
Fluorescentcoral[41]
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Darwin'sparadox
InTheStructureandDistributionofCoralReefs,publishedin1842,Darwindescribedhowcoralreefswerefoundinsomeareasofthe
tropicalseasbutnotothers,withnoobviouscause.Thelargestandstrongestcoralsgrewinpartsofthereefexposedtothemostviolent
surfandcoralswereweakenedorabsentwhereloosesedimentaccumulated.[43]
Tropicalwaterscontainfewnutrients[44]yetacoralreefcanflourishlikean"oasisinthedesert".[45]Thishasgivenrisetotheecosystem
conundrum,sometimescalled"Darwin'sparadox":"Howcansuchhighproductionflourishinsuchnutrientpoorconditions?"[46][47][48]
Coralreefscoverlessthan0.1%ofthesurfaceoftheworldsocean,abouthalfthelandareaofFrance,yettheysupportoveronequarter
ofallmarinespecies.Thisdiversityresultsincomplexfoodwebs,withlargepredatorfisheatingsmallerforagefishthateatyetsmaller
zooplanktonandsoon.However,allfoodwebseventuallydependonplants,whicharetheprimaryproducers.Coralreefs'primary
productivityisveryhigh,typicallyproducing510gramsofCarbonpersquaremeterperday(gCm2day1biomass.[49][50]
Onereasonfortheunusualclarityoftropicalwatersistheyaredeficientinnutrientsanddriftingplankton.Further,thesunshinesyear
roundinthetropics,warmingthesurfacelayer,makingitlessdensethansubsurfacelayers.Thewarmerwaterisseparatedfromdeeper,
coolerwaterbyastablethermocline,wherethetemperaturemakesarapidchange.Thiskeepsthewarmsurfacewatersfloatingabovethe
coolerdeeperwaters.Inmostpartsoftheocean,thereislittleexchangebetweentheselayers.Organismsthatdieinaquaticenvironments
generallysinktothebottom,wheretheydecompose,whichreleasesnutrientsintheformofnitrogen(N),phosphorus(P)andpotassium
(K).Thesenutrientsarenecessaryforplantgrowth,butinthetropics,theydonotdirectlyreturntothesurface.
Plantsformthebaseofthefoodchain,andneedsunlightandnutrientstogrow.Intheocean,theseplantsaremainlymicroscopic
phytoplanktonwhichdriftinthewatercolumn.Theyneedsunlightforphotosynthesis,whichpowerscarbonfixation,sotheyarefound
onlyrelativelynearthesurface.Buttheyalsoneednutrients.Phytoplanktonrapidlyusenutrientsinthesurfacewaters,andinthetropics,
thesenutrientsarenotusuallyreplacedbecauseofthethermocline.
Darwin'sparadox
"Coral...seemstoproliferatewhenocean
watersarewarm,poor,clearandagitated,a
factwhichDarwinhadalreadynotedwhenhe
passedthroughTahitiin1842.This
constitutesafundamentalparadox,shown
quantitativelybytheapparentimpossibility
ofbalancinginputandoutputofthenutritive
elementswhichcontrolthecoralpolyp
metabolism.
Recentoceanographicresearchhasbroughtto
lighttherealityofthisparadoxby
confirmingthattheoligotrophyoftheocean
euphoticzonepersistsrightuptotheswell
batteredreefcrest.Whenyouapproachthe
reefedgesandatollsfromthequasidesertof
theopensea,thenearabsenceofliving
mattersuddenlybecomesaplethoraoflife,
withouttransition.Sowhyistheresomething
ratherthannothing,andmoreprecisely,
wheredothenecessarynutrientsforthe
functioningofthisextraordinarycoralreef
machinecomefrom?"Francis
Rougerie[42]
Explanations
Aroundcoralreefs,lagoonsfillinwithmaterialerodedfromthereefandtheisland.Theybecomehavensformarinelife,providingprotectionfromwavesandstorms.
Mostimportantly,reefsrecyclenutrients,whichhappensmuchlessintheopenocean.Incoralreefsandlagoons,producersincludephytoplankton,aswellasseaweedandcoralline
algae,especiallysmalltypescalledturfalgae,whichpassnutrientstocorals.[51]Thephytoplanktonareeatenbyfishandcrustaceans,whoalsopassnutrientsalongthefoodweb.
Recyclingensuresfewernutrientsareneededoveralltosupportthecommunity.
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Coralreefssupportmanysymbioticrelationships.Inparticular,zooxanthellaeprovideenergytocoralintheformofglucose,glycerol,andaminoacids.[52]Zooxanthellaecanprovideup
to90%ofacoralsenergyrequirements.[33]Inreturn,asanexampleofmutualism,thecoralsshelterthezooxanthellae,averagingonemillionforeverycubiccentimeterofcoral,and
provideaconstantsupplyofthecarbondioxidetheyneedforphotosynthesis.
Coralsalsoabsorbnutrients,includinginorganicnitrogenandphosphorus,directlyfromwater.Manycoralsextendtheirtentaclesatnighttocatchzooplanktonthatbrushthemwhenthe
waterisagitated.Zooplanktonprovidethepolypwithnitrogen,andthepolypsharessomeofthenitrogenwiththezooxanthellae,whichalsorequirethiselement.[51]Thevarying
pigmentsindifferentspeciesofzooxanthellaegivethemanoverallbrownorgoldenbrownappearance,andgivebrowncoralstheircolors.Otherpigmentssuchasreds,blues,greens,
etc.comefromcoloredproteinsmadebythecoralanimals.Coralwhichlosesalargefractionofitszooxanthellaebecomeswhite(orsometimespastelshadesincoralsthatarerichly
pigmentedwiththeirowncolorfulproteins)andissaidtobebleached,aconditionwhich,unlesscorrected,cankillthecoral.
Spongesareanotherkey:theyliveincrevicesinthecoralreefs.Theyareefficientfilterfeeders,andinthe
RedSeatheyconsumeabout60%ofthephytoplanktonthatdriftsby.Thespongeseventuallyexcrete
nutrientsinaformthecoralscanuse.[53]
Theroughnessofcoralsurfacesisthekeytocoralsurvivalinagitatedwaters.Normally,aboundarylayer
ofstillwatersurroundsasubmergedobject,whichactsasabarrier.Wavesbreakingontheextremely
roughedgesofcoralsdisrupttheboundarylayer,allowingthecoralsaccesstopassingnutrients.
Turbulentwatertherebypromotesreefgrowthandbranching.Withoutthenutritionalgainsbroughtby
roughcoralsurfaces,eventhemosteffectiverecyclingwouldleavecoralswantinginnutrients.[54]
Studieshaveshownthatdeepnutrientrichwaterenteringcoralreefsthroughisolatedeventsmayhave
significanteffectsontemperatureandnutrientsystems.[55][56]Thiswatermovementdisruptstherelatively
stablethermoclinethatusuallyexistsbetweenwarmshallowwatertodeepercolderwater.Leichteretal.
[57]
(2006) foundthattemperatureregimesoncoralreefsintheBahamasandFloridawerehighlyvariablewithtemporalscalesofminutesto
seasonsandspatialscalesacrossdepths.
Coralpolyps
Thecolorofcoralsdependsonthe
combinationofbrownshades
providedbytheirzooxanthellaeand
pigmentedproteins(reds,blues,
greens,etc.)producedbythecorals
themselves.
Watercanbemovedthroughcoralreefsinvariousways,includingcurrentrings,surfacewaves,internalwavesandtidalchanges.[55][58][59][60]Movementisgenerallycreatedbytides
andwind.Astidesinteractwithvaryingbathymetryandwindmixeswithsurfacewater,internalwavesarecreated.Aninternalwaveisagravitywavethatmovesalongdensity
stratificationwithintheocean.Whenawaterparcelencountersadifferentdensityitwilloscillateandcreateinternalwaves.[61]Whileinternalwavesgenerallyhavealowerfrequency
thansurfacewaves,theyoftenformasasinglewavethatbreaksintomultiplewavesasithitsaslopeandmovesupward.[62]Thisverticalbreakupofinternalwavescausessignificant
diapycnalmixingandturbulence.[63][64]Internalwavescanactasnutrientpumps,bringingplanktonandcoolnutrientrichwateruptothesurface.[55][60][65][66][67][68][69][70][71][72][73]
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Theirregularstructurecharacteristicofcoralreefbathymetrymayenhancemixingandproducepocketsofcoolerwaterandvariablenutrient
content.[74]Arrivalofcool,nutrientrichwaterfromdepthsduetointernalwavesandtidalboreshasbeenlinkedtogrowthratesofsuspension
feedersandbenthicalgae[60][73][75]aswellasplanktonandlarvalorganisms.[60][76]Leichteretal.[73]proposedthatCodiumisthmocladumreactto
deepwaternutrientsourcesduetotheirtissueshavingdifferentconcentrationsofnutrientsdependentupondepth.WolanskiandHamner[67]
notedaggregationsofeggs,larvalorganismsandplanktononreefsinresponsetodeepwaterintrusions.Similarly,asinternalwavesandbores
movevertically,surfacedwellinglarvalorganismsarecarriedtowardtheshore.[76]Thishassignificantbiologicalimportancetocascading
effectsoffoodchainsincoralreefecosystemsandmayprovideyetanotherkeytounlocking"Darwin'sParadox".
Mostcoralpolypsarenocturnal
feeders.Here,inthedark,polyps
haveextendedtheirtentaclestofeed
onzooplankton.
Cyanobacteriaprovidesolublenitratesforthereefvianitrogenfixation.[77]
Coralreefsalsooftendependonsurroundinghabitats,suchasseagrassmeadowsandmangroveforests,fornutrients.Seagrassandmangroves
supplydeadplantsandanimalswhicharerichinnitrogenandalsoservetofeedfishandanimalsfromthereefbysupplyingwoodand
vegetation.Reefs,inturn,protectmangrovesandseagrassfromwavesandproducesedimentinwhichthemangrovesandseagrasscanroot.[26]
Biodiversity
Coralreefsformsomeoftheworld'smostproductiveecosystems,providingcomplexandvariedmarinehabitatsthatsupportawiderangeof
otherorganisms.[78]Fringingreefsjustbelowlowtidelevelhaveamutuallybeneficialrelationshipwithmangroveforestsathightideleveland
seagrassmeadowsinbetween:thereefsprotectthemangrovesandseagrassfromstrongcurrentsandwavesthatwoulddamagethemorerode
thesedimentsinwhichtheyarerooted,whilethemangrovesandseagrassprotectthecoralfromlargeinfluxesofsilt,freshwaterandpollutants.
Thislevelofvarietyintheenvironmentbenefitsmanycoralreefanimals,which,forexample,mayfeedintheseagrassandusethereefsfor
protectionorbreeding.[79]
Reefsarehometoalargevarietyofanimals,includingfish,seabirds,sponges,cnidarians(whichincludessometypesofcoralsandjellyfish),
worms,crustaceans(includingshrimp,cleanershrimp,spinylobstersandcrabs),mollusks(includingcephalopods),echinoderms(including
starfish,seaurchinsandseacucumbers),seasquirts,seaturtlesandseasnakes.Asidefromhumans,mammalsarerareoncoralreefs,with
visitingcetaceanssuchasdolphinsbeingthemainexception.Afewofthesevariedspeciesfeeddirectlyoncorals,whileothersgrazeonalgae
onthereef.[2][51]Reefbiomassispositivelyrelatedtospeciesdiversity.[80]
Thesamehideoutsinareefmayberegularlyinhabitedbydifferentspeciesatdifferenttimesofday.Nighttimepredatorssuchascardinalfishand
squirrelfishhideduringtheday,whiledamselfish,surgeonfish,triggerfish,wrassesandparrotfishhidefromeelsandsharks.[20]:49
Algae
Tubespongesattractingcardinal
fishes,glassfishesandwrasses
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Reefsarechronicallyatriskofalgalencroachment.Overfishingandexcessnutrientsupplyfromonshorecanenablealgaetooutcompeteandkill
thecoral.[81][82]InsurveysdonearoundlargelyuninhabitedUSPacificislands,algaeinhabitalargepercentageofsurveyedcorallocations.[83]
Thealgalpopulationconsistsofturfalgae,corallinealgae,andmacroalgae.
Sponges
Spongesareessentialforthefunctioningofthecoralreef'secosystem.Algaeandcoralsincoralreefsproduceorganicmaterial.Thisisfiltered
throughspongeswhichconvertthisorganicmaterialintosmallparticleswhichinturnareabsorbedbyalgaeandcorals.[84]
Over4,000speciesoffishinhabit
coralreefs.
Fish
Over4,000speciesoffishinhabitcoralreefs.[2]Thereasonsforthisdiversityremainunclear.Hypothesesincludethe"lottery",inwhichthefirst
(luckywinner)recruittoaterritoryistypicallyabletodefenditagainstlatecomers,"competition",inwhichadultscompeteforterritory,and
lesscompetitivespeciesmustbeabletosurviveinpoorerhabitat,and"predation",inwhichpopulationsizeisafunctionofpostsettlement
piscivoremortality.[85]Healthyreefscanproduceupto35tonsoffishpersquarekilometereachyear,butdamagedreefsproducemuchless.[86]
Invertebrates
Seaurchins,Dotidaeandseaslugseatseaweed.Somespeciesofseaurchins,suchasDiademaantillarum,canplayapivotalpartinpreventing
algaefromoverrunningreefs.[87]Nudibranchiaandseaanemoneseatsponges.
Anumberofinvertebrates,collectivelycalledcryptofauna,inhabitthecoralskeletalsubstrateitself,eitherboringintotheskeletons(throughthe
processofbioerosion)orlivinginpreexistingvoidsandcrevices.Thoseanimalsboringintotherockincludesponges,bivalvemollusks,and
sipunculans.Thosesettlingonthereefincludemanyotherspecies,particularlycrustaceansandpolychaeteworms.[29]
Seabirds
Organismscancovereverysquare
inchofacoralreef.
Coralreefsystemsprovideimportanthabitatsforseabirdspecies,someendangered.Forexample,MidwayAtollinHawaiisupportsnearlythreemillionseabirds,includingtwothirds
(1.5million)oftheglobalpopulationofLaysanalbatross,andonethirdoftheglobalpopulationofblackfootedalbatross.[88]Eachseabirdspecieshasspecificsitesontheatollwhere
theynest.Altogether,17speciesofseabirdsliveonMidway.Theshorttailedalbatrossistherarest,withfewerthan2,200survivingafterexcessivefeatherhuntinginthelate19th
century.[89]
Other
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Seasnakesfeedexclusivelyonfishandtheireggs.[90][91][92]Marinebirds,suchasherons,gannets,pelicansandboobies,feedonreeffish.Somelandbasedreptilesintermittently
associatewithreefs,suchasmonitorlizards,themarinecrocodileandsemiaquaticsnakes,suchasLaticaudacolubrina.Seaturtles,particularlyHawksbillseaturtles,feedon
sponges.[93][94][95]
Schoolingreeffish
Caribbeanreefsquid
Bandedcoral Whitetipreefshark
shrimp
Greenturtle
Giantclam
Softcoral,cupcoral,
spongesandascidians
Bandedsea
krait
Theshellof
Latiaxis
wormaldi,acoral
snail
Importance
Coralreefsdeliverecosystemservicestotourism,fisheriesandcoastlineprotection.TheglobaleconomicvalueofcoralreefshasbeenestimatedtobebetweenUS$29.8billion[96]and
$375billionperyear.[97]Coralreefsprotectshorelinesbyabsorbingwaveenergy,andmanysmallislandswouldnotexistwithouttheirreefstoprotectthem.Accordingtothe
environmentalgroupWorldWideFundforNature,theeconomiccostovera25yearperiodofdestroyingonekilometerofcoralreefissomewherebetween$137,000and
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$1,200,000.[98]Aboutsixmilliontonsoffisharetakeneachyearfromcoralreefs.Wellmanagedcoralreefshaveanannualyieldof15tonsofseafoodonaveragepersquarekilometer.
SoutheastAsia'scoralreeffisheriesaloneyieldabout$2.4billionannuallyfromseafood.[98]
Toimprovethemanagementofcoastalcoralreefs,anotherenvironmentalgroup,theWorldResourcesInstitute(WRI)developedandpublishedtoolsforcalculatingthevalueofcoral
reefrelatedtourism,shorelineprotectionandfisheries,partneringwithfiveCaribbeancountries.AsofApril2011,publishedworkingpaperscoveredSt.Lucia,Tobago,Belize,andthe
DominicanRepublic,withapaperforJamaicainpreparation.TheWRIwasalso"makingsurethatthestudyresultssupportimprovedcoastalpoliciesandmanagementplanning".[99]
TheBelizestudyestimatedthevalueofreefandmangroveservicesat$395559millionannually.[100]
Threats
Coralreefsaredyingaroundtheworld.[101]Inparticular,coralmining,agriculturalandurbanrunoff,pollution(organicandinorganic),
overfishing,blastfishing,disease,andthediggingofcanalsandaccessintoislandsandbaysarelocalizedthreatstocoralecosystems.Broader
threatsareseatemperaturerise,sealevelriseandpHchangesfromoceanacidification,allassociatedwithgreenhousegasemissions.A2014
studylistsfactorssuchaspopulationexplosionalongthecoastlines,overfishing,thepollutionofcoastalareas,globalwarmingandinvasive
speciesamongthemainreasonsthathaveputreefsindangerofextinction.[102]
AstudyreleasedinApril2013hasshownthatairpollutioncanalsostuntthegrowthofcoralreefsresearchersfromAustralia,Panamaandthe
UKusedcoralrecords(between1880and2000)fromthewesternCaribbeantoshowthethreatoffactorssuchascoalburningcoalandvolcanic
eruptions.[103]Pollutants,suchasTributyltin,abiocidereleasedintowaterfrominantifoulingpaintcanbetoxictocorals.
In2011,researcherssuggestedthat"extantmarineinvertebratesfacethesamesynergisticeffectsofmultiplestressors"thatoccurredduringthe
endPermianextinction,andthatgenera"withpoorlybufferedrespiratoryphysiologyandcalcareousshells",suchascorals,wereparticularly
vulnerable.[104][105][106]
IslandwithfringingreefoffYap,
Micronesia[101]
Rockcoralonseamountsacrosstheoceanareunderfirefrombottomtrawling.Reportedlyupto50%ofthecatchisrockcoral,andthepracticetransformscoralstructurestorubble.
Withittakingyearstoregrow,thesecoralcommunitiesaredisappearingfasterthantheycansustainthemselves.[107]
Anothercauseforthedeathofcoralreefsisbioerosion.Variousfishesgrazecorals,deadoraliveandchangethemorphologyofcoralreefsmakingthemmoresusceptibletoother
physicalandchemicalthreats.Ithasbeengenerallyobservedthatonlythealgaegrowingondeadcoralsiseatenandtheliveonesarenot.However,thisactstilldestroysthetoplayerof
coralsubstrateandmakesitharderforthereefstosustain.[108]
InElNinoyear2010,preliminaryreportsshowglobalcoralbleachingreacheditsworstlevelsinceanotherElNinoyear,1998,when16%oftheworld'sreefsdiedasaresultof
increasedwatertemperature.InIndonesia'sAcehprovince,surveysshowedsome80%ofbleachedcoralsdied.Scientistsdonotyetunderstandthelongtermimpactsofcoralbleaching,
buttheydoknowthatbleachingleavescoralsvulnerabletodisease,stuntstheirgrowth,andaffectstheirreproduction,whileseverebleachingkillsthem.[109]InJuly,Malaysiaclosed
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severaldivesiteswherevirtuallyallthecoralsweredamagedbybleaching.[110][111]
Tofindanswersfortheseproblems,researchersstudythevariousfactorsthatimpactreefs.Thelistincludestheocean'sroleasacarbondioxidesink,atmosphericchanges,ultraviolet
light,oceanacidification,viruses,impactsofduststormscarryingagentstofarflungreefs,pollutants,algalbloomsandothers.Reefsarethreatenedwellbeyondcoastalareas.Coral
reefswithonetypeofzooxanthellaearemorepronetobleachingthanarereefswithanother,morehardy,species.[112]
Generalestimatesshowapproximately10%oftheworld'scoralreefsaredead.[113][114]About60%oftheworld'sreefsareatriskduetodestructive,humanrelatedactivities.Thethreat
tothehealthofreefsisparticularlyhighinSoutheastAsia,where95%ofreefsareatriskfromlocalthreats.[115]Bythe2030s,90%ofreefsareexpectedtobeatriskfrombothhuman
activitiesandclimatechangeby2050,allcoralreefswillbeindanger.[116]
CurrentresearchisshowingthatecotourismintheGreatBarrierReefiscontributingtocoraldisease,[117]andthatchemicalsinsunscreensmaycontributetotheimpactofviruseson
zooxanthellae.[5]
Protection
Marineprotectedareas(MPAs)havebecomeincreasinglyprominentforreefmanagement.MPAspromoteresponsiblefisherymanagementand
habitatprotection.Muchlikenationalparksandwildliferefuges,andtovaryingdegrees,MPAsrestrictpotentiallydamagingactivities.MPAs
encompassbothsocialandbiologicalobjectives,includingreefrestoration,aesthetics,biodiversity,andeconomicbenefits.Howeverthereare
veryfewMPAsthathaveactuallymadeasubstantialdifference.ResearchinIndonesia,PhilippinesandPapuaNewGuineashowsthatthereis
nosignificantdifferencebetweenanMPAsiteandanunprotectedsite.[118][119]ConflictssurroundingMPAsinvolvelackofparticipation,
clashingviewsofthegovernmentandfisheries,effectivenessofthearea,andfunding.[120]Insomesituations,asinthePhoenixIslandsProtected
Area,MPAscanalsoproviderevenue,potentiallyequaltotheincometheywouldhavegeneratedwithoutcontrols,asKiribatididforitsPhoenix
Islands.[121]
Tohelpcombatoceanacidification,somelawsareinplacetoreducegreenhousegasessuchascarbondioxide.TheCleanWaterActputs
pressureonstategovernmentagenciestomonitorandlimitrunoffofpollutantsthatcancauseoceanacidification.Stormwatersurgepreventions
arealsoinplace,aswellascoastalbuffersbetweenagriculturallandandthecoastline.Thisactalsoensuresthatdelicatewatershedecosystems
areintact,suchaswetlands.TheCleanWaterActisfundedbythefederalgovernment,andismonitoredbyvariouswatershedgroups.ManylanduselawsaimtoreduceCO2emissions
Adiversityofcorals
bylimitingdeforestation.Deforestationcauseserosion,whichreleasesalargeamountofcarbonstoredinthesoil,whichthenflowsintotheocean,contributingtooceanacidification.
Incentivesareusedtoreducemilestraveledbyvehicles,whichreducesthecarbonemissionsintotheatmosphere,therebyreducingtheamountofdissolvedCO2intheocean.Stateand
federalgovernmentsalsocontrolcoastalerosion,whichreleasesstoredcarboninthesoilintotheocean,increasingoceanacidification.[122]
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Biospherereserve,marinepark,nationalmonumentandworldheritagestatuscanprotectreefs.Forexample,Belize'sbarrierreef,Chagosarchipelago,SianKa'an,theGalapagos
islands,GreatBarrierReef,HendersonIsland,PalauandPapahnaumokukeaMarineNationalMonumentareworldheritagesites.
InAustralia,theGreatBarrierReefisprotectedbytheGreatBarrierReefMarineParkAuthority,andisthesubjectofmuchlegislation,includingabiodiversityactionplan.[123]They
havecompiledaCoralReefResilienceActionPlan.Thisdetailedactionplanconsistsofnumerousadaptivemanagementstrategies,includingreducingourcarbonfootprint,which
wouldultimatelyreducetheamountofoceanacidificationintheoceanssurroundingtheGreatBarrierReef.Anextensivepublicawarenessplanisalsoinplacetoprovideeducationon
therainforestsoftheseaandhowpeoplecanreducecarbonemissions,therebyreducingoceanacidification.[124]
InhabitantsofAhusIsland,ManusProvince,PapuaNewGuinea,havefollowedagenerationsoldpracticeofrestrictingfishinginsixareasoftheirreeflagoon.Theirculturaltraditions
allowlinefishing,butnonetorspearfishing.Theresultisboththebiomassandindividualfishsizesaresignificantlylargerthaninplaceswherefishingisunrestricted.[125][126]
Highendsatellitetechnologyiscomingtotheaidofprotectingearthslargestbiologicalstructuresthecoralreefswhichhavetakenmillionsofyearstoformandaredecliningat
alarmingratesworldwide
[127]
Restoration
Coralaquaculture,alsoknownascoralfarmingorcoralgardening,isshowingpromiseasapotentiallyeffectivetoolforrestoringcoralreefs,
whichhavebeendecliningaroundtheworld.[128][129][130]Theprocessbypassestheearlygrowthstagesofcoralswhentheyaremostatriskof
dying.Coralseedsaregrowninnurseries,thenreplantedonthereef.[131]Coralisfarmedbycoralfarmerswholivelocallytothereefsandfarm
forreefconservationorforincome.
Effortstoexpandthesizeandnumberofcoralreefsgenerallyinvolvesupplyingsubstratetoallowmorecoralstofindahome.Substrate
materialsincludediscardedvehicletires,scuttledships,subwaycars,andformedconcrete,suchasreefballs.Reefsalsogrowunaidedonmarine
structuressuchasoilrigs.Inlargerestorationprojects,propagatedhermatypiccoralonsubstratecanbesecuredwithmetalpins,superglueor
milliput.[132]NeedleandthreadcanalsoattachAhermatypecoraltosubstrate.[133]
Coralfragmentsgrowingonnontoxic
concrete
AsubstrateforgrowingcoralsreferredtoasBiorockisproducedbyrunninglowvoltageelectricalcurrentsthroughseawatertocrystallize
dissolvedmineralsontosteelstructures.Theresultantwhitecarbonate(aragonite)isthesamemineralthatmakesupnaturalcoralreefs.Corals
rapidlycolonizeandgrowatacceleratedratesonthesecoatedstructures.Theelectricalcurrentsalsoaccelerateformationandgrowthofbothchemicallimestonerockandtheskeletons
ofcoralsandothershellbearingorganisms.ThevicinityoftheanodeandcathodeprovidesahighpHenvironmentwhichinhibitsthegrowthofcompetitivefilamentousandfleshy
algae.Theincreasedgrowthratesfullydependontheaccretionactivity.[134]
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Duringaccretion,thesettledcoralsdisplayanincreasedgrowthrate,sizeanddensity,butaftertheprocessiscomplete,growthrateanddensityreturntolevelscomparabletonatural
growth,andareaboutthesamesizeorslightlysmaller.[134]
OnecasestudywithcoralreefrestorationwasconductedontheislandofOahuinHawaii.TheUniversityofHawaiihascomeupwithaCoralReefAssessmentandMonitoring
ProgramtohelprelocateandrestorecoralreefsinHawaii.AboatchannelontheislandofOahutotheHawaiiInstituteofMarineBiologywasovercrowdedwithcoralreefs.Also,
manyareasofcoralreefpatchesinthechannelhadbeendamagedfrompastdredginginthechannel.Dredgingcoverstheexistingcoralswithsand,andtheirlarvaecannotbuildand
thriveonsandtheycanonlybuildontoexistingreefs.Becauseofthis,theUniversityofHawaiidecidedtorelocatesomeofthecoralreeftoadifferenttransplantsite.They
transplantedthemwiththehelpoftheUnitedStatesArmyDivers,toarelocationsiterelativelyclosetothechannel.Theyobservedverylittle,ifany,damageoccurredtoanyofthe
colonieswhiletheywerebeingtransported,andnomortalityofcoralreefshasbeenobservedonthenewtransplantsite,buttheywillbecontinuingtomonitorthenewtransplantsiteto
seehowpotentialenvironmentalimpacts(i.e.oceanacidification)willharmtheoverallreefmortalityrate.Whiletryingtoattachthecoraltothenewtransplantsite,theyfoundthecoral
placedonhardrockisgrowingconsiderablywell,andcoralwasevengrowingonthewiresthatattachedthetransplantcoralstothetransplantsite.Thisgivesnewhopetofuture
researchoncoralreeftransplantsites.Asaresultofthiscoralrestorationproject,noenvironmentaleffectswereseenfromthetransplantationprocess,norecreationalactivitieswere
decreased,andnoscenicareaswereaffectedbytheproject.Thisisagreatexamplethatcoraltransplantationandrestorationcanworkandthriveundertherightconditions,which
meanstheremaybehopeforotherdamagedcoralreefs.[135]
Anotherpossibilityforcoralrestorationisgenetherapy.Throughinfectingcoralwithgeneticallymodifiedbacteria,itmaybepossibletogrowcoralsthataremoreresistanttoclimate
changeandotherthreats.[136]
Reefsinthepast
ThroughoutEarthhistory,fromafewthousandyearsafterhardskeletonsweredevelopedbymarineorganisms,therewerealmostalways
reefs.ThetimesofmaximumdevelopmentwereintheMiddleCambrian(513501Ma),Devonian(416359Ma)andCarboniferous(359
299Ma),owingtoorderRugosaextinctcorals,andLateCretaceous(10066Ma)andallNeogene(23Mapresent),owingtoorder
Scleractiniacorals.
Notallreefsinthepastwereformedbycorals:thoseintheEarlyCambrian(542513Ma)resultedfromcalcareousalgaeand
archaeocyathids(smallanimalswithconicalshape,probablyrelatedtosponges)andintheLateCretaceous(10066Ma),whentherealso
existedreefsformedbyagroupofbivalvescalledrudistsoneofthevalvesformedthemainconicalstructureandtheother,muchsmaller
valveactedasacap.
Ancientcoralreefs
Measurementsoftheoxygenisotopiccompositionofthearagoniticskeletonofcoralreefs,suchasPorites,canindicatechangesinthesea
surfacetemperatureandseasurfacesalinityconditionsoftheoceanduringthegrowthofthecoral.Thistechniqueisoftenusedbyclimate
scientiststoinferthepaleoclimateofaregion.[137]
Seealso
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CatlinSeaviewSurvey
CensusofCoralReefs
Coralreeforganizations
Marinebiology
Spongereef
Pseudoatoll
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Furtherreferences
CoralReefProtection:WhatAreCoralReefs?(http://water.epa.gov/type/oceb/habitat/coral_index.cfm).USEPA.
UNEP.2004.CoralReefsintheSouthChinaSea.UNEP/GEF/SCSTechnicalPublicationNo.2.(http://www.unepscs.org/SCS_Documents/Download/13_
_Habitat_Booklets/UNEP_or_GEF_Review_of_Coral_Reefs_in_the_South_China_Sea.html)
UNEP.2007.CoralReefsDemonstrationSitesintheSouthChinaSea.UNEP/GEF/SCSTechnicalPublicationNo.5.(http://www.unepscs.org/SCS_Documents/Download/19_
_Technical_Publications_and_Guidelines/Technical_Publication_05__Coral_Reef_Demonstration_Sites_in_the_South_China_Sea.html)
UNEP,2007.NationalReportsonCoralReefsintheCoastalWatersoftheSouthChinaSea.UNEP/GEF/SCSTechnicalPublicationNo.11.
(http://www.unepscs.org/SCS_Documents/Download/19__Technical_Publications_and_Guidelines/Technical_Publication_11_
_National_Reports_on_Coral_Reefs_in_the_Coastal_Waters_of_the_South_China_Sea.html)
Externallinks
Externalimages
CoralsandCoralReefs(http://ocean.si.edu/coralsandcoralreefs)overviewattheSmithsonianOceanPortal
AboutCorals(http://coral.aims.gov.au/info/about.jsp)AustralianInstituteofMarineScience.
InternationalCoralReefInitiative(http://www.icriforum.org)
MooreaCoralReefLongTermEcologicalResearchSite(USNSF)(http://mcr.lternet.edu)
CoralReefs:RainforestsoftheSea
(http://www.oceanicresearch.org/education/films/crrain_qt.htm)
ORGEducationalfilms.
ARCCentreofExcellenceforCoralReefStudies(http://www.coralcoe.org.au)
NOAA'sCoralListListserverforCoralReefInformationandNews(http://coral.aoml.noaa.gov/mailman/listinfo/corallist/)
NOAA'sCoralReefConservationProgram(http://www.coralreef.noaa.gov/)
NOAA'sCoralReefInformationSystem(http://www.coris.noaa.gov/)
ReefBase:AGlobalInformationSystemonCoralReefs(http://www.reefbase.org/)
NationalCoralReefInstitute(http://www.nova.edu/ncri/)NovaSoutheasternUniversity
WikimediaCommonshas
mediarelatedtoCoralreefs.
Wikisourcehasthetextof
the1911Encyclopdia
BritannicaarticleCoral
reefs.
MarineAquariumCouncil(http://www.aquariumcouncil.org)
NCORENationalCenterforCoralReefResearch(http://ncore.rsmas.miami.edu/)UniversityofMiami
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12/21/2014
ScienceandManagementofCoralReefsintheSouthChinaSeaandGulfofThailand(http://www.unepscs.org/index.php?option=com_content&task=view&id=51&Itemid=83)
Microdocs(http://www.stanford.edu/group/microdocs/):4kindsofReef(http://www.stanford.edu/group/microdocs/typesofreefs.html)&Reefstructure
(http://www.stanford.edu/group/microdocs/reefstructure.html)
ReefRelief(http://reefrelief.org/)ActiveFloridaenvironmentalnonprofitfocusingoncoralreefeducationandprotection
GlobalReefRecord(http://globalreefrecord.org/home_scientific)CatlinSeaviewSurveyofreef,adatabaseofimagesandotherinformation
CoralsandCoralReefs(http://www.ibiology.org/ibioseminars/evolutionecology/nancyknowltonpart1.html)NancyKnowlton,iBioSeminars,2011.
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