UNIVERSITY OF HAWAII LJB~~ FACTORS INFLUENCING ALGAL BLOOMS ONTROPICALREEFS WITH AN EMPHASIS ONHERBIVORY, NUTRIENTS AND INVASIVE SPECIES A DISSERTATIONSUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI'IINPARTIALFULFILLMENT OF THE REQUIREMENTS FORTHE DEGREE OF DOCTOR OF PHILOSOPHY IN BOTANY (ECOLOGY, EVOLUTION AND CONSERVATION BIOLOGY) DECEMBER 2003 By JenniferE. Smith Dissertation Committee: CeliaSmith, Chairperson CynthiaHunter Lawrence Basch Lucius Eldredge Michael Hamnett ABSTRACT Algal blooms ontropical reefs havebecome increasinglycommon overthe last several decades. Reduced herbivorepressure, eutrophication andthe introductionof exotic species have all beenposited as factors that maydrive these transitions in organism dominance. Several different types ofalgal blooms havebeenidentifiedon reefs inthe HawaiianIslands includingmultispecies algal assemblages, monospecific algal blooms and seasonal or ephemeralblooms. This studysoughtto determine the causes ofthesedifferent types ofalgal bloomsbyconductingboth field and laboratory experiments and quantitative field assessments. In a factorial nutrient enrichment and herbivore exclusion experimentconducted for 6 months onthe island ofHawai'i significantchanges in algal biomass, community structure, sediment accumulation andmobile microinvertebrate abundancewere found. Results ofthis studyshow thatbenthic reefcommunities canchangerapidlyin response to changes inbothtop down and bottomup factors. From field assessments across the mainHawaiianIslands atotal offive (Acanthophora spicifera, Hypnea musciformis, Graci/aria salicornia, Kappaphycus spp. andAvranvilleaamadelpha) species ofnonindigenous algae cannowbe considered highlysuccessful. Detailed studies onthe ecologyofselectnonindigenous marine algae (NIMA) have identifiedparticularconcerns andhighlighted the need for management action. In summarymanagement ofthese invasive species will be challenging as aresult ofunique ecological andphysiological strategiesthat eachNIMApossesses. iii The ephemeralbloom forming native alga Cladophora sericea was studiedduring abloom cycleduring 2001 onthe island ofMaui. Results ofambient and sedimentpore watersampling and algal physiologicalparameters suggest that ground waterintrusionis occurring at this site and the algaappears to be utilizing this terrestrial basednutrient source. Upwelling and internaltides cannaturallydelivernutrient rich waterinto coral reefecosystems. Inan areawhere internal tidal upwelling occurs inthe FloridaKeys, one ofthe most commonbenthicreefalgaeHalimeda tuna reflectedpatterns associated with natural nutrientenrichment. This studyprovides evidencethat rich and highlyproductive benthic algae maythriveindeep watercoralreefenvironments inresponse to naturally elevatednutrient conditions. iv TABLE OF CONTENTS ABSTRACT .iii LIST OF TABLES xi LIST OFFIGURES xviii CHAPTER 1. LITERATUREREVIEW AND DISSERTATIONPROPOSAL 1 INTRODUCTION....................................•..............................." 2 NUTRIENTDYNAMICSONTROPICALREEFS .4 NUTRIENTGRADIENTSONTROPICALREEFS " •.•....•.5 EFFECTSOFNUTRIENTENRICHMENTONBENTHICCOMMUNITIES 7 EFFECTSOFNUTRIENTENRICHMENTONALGALSPECIES 9 NATURALLYELEVATEDNUTRIENTSONTROPICALREEFS 14 SUMMARy 16 INTERACTIVEEFFECTSOFNUTRIENTSANDHERBIVORES " 17 TOP-DoWNANDBOTTOM-UPFACTORSINFLUENCINGALGAL COMMUNITIES 17 SUMMARy 23 INTRODUCTIONOFNONINDIGENOUSSPECIESONTROPICALREEFS 23 IMPACTSOFSPECIESINTRODUCTIONS 23 ACCIDENTALINTRODUCTIONOFSEAWEEDS 24 SARGASSUMMUT/CUM...............................................•.....24 CAULERPA TAX/FOL/A 26 UNDARIA P/NNAT/F/DA 28 v SPECIESINTRODUCEDFORCULTUREINTHETROPICS .30 /(APPAPHYCUSANDEUCHEUMA .30 EFFECTSOFSEAWEEDINTRODUCTIONS .33 A CASESTUDY: KANE'OHEBAY 35 TYPESOFALGALBLOOMS 38 RESEARCHPROPOSALANDOUTLINE .40 TABLES 42 FIGURES 59 LITERATURECITED 60 CHAPTER2. AN EXPERIMENTALANALYSIS OF THE EFFECTS OF HERBIVORY AND NUTRIENT ENRICHMENT ONBENTHIC COMMUNITY SUCCESSION ONA HAWAIIANREEF 83 ABSTRACT 84 INTRODUCTION 85 METHODSANDMATERIALS " 90 RESULTS 98 DISCUSSION 105 CONCLUSIONS 110 ACKNOWLEDGEMENTS 111 TABLES 112 VI FIGURES 129 LITERATURECITED .143 CHAPTER 3. DISTRIBUTIONAND REPRODUCTIVE CHARACTERISTICS OF NONINDIGENOUS AND INVASIVEMARINE ALGAE INTHE HAWAIIAN ISLANDS 151 ABSTRACT 152 INTRODUCTION 153 METHODSANDMATERIALS 156 RESULTS 159 DISCUSSION 163 CONCLUSIONS 170 ACKNOWLEDGEMENTS 172 TABLES 173 FIGURES 179 LITERATURECITED 183 CHAPTER4. THE ECOLOGY OF THE INVASIVERED ALGA GRACILARIA SALICORNIA (C. AGARDH) E.Y. DAWSON ON O'AHU, HAWAI'I.. 188 ABSTRACT '" 189 INTRODUCTION 191 vii METHODSANDMATERIALS 195 RESULTS 203 DISCUSSION 208 CONCLUSIONS 214 ACKNOWLEDGEMENTS 215 TABLES 216 FIGURES 217 LITERATURECITED 225 CHAPTER 5. ABUNDANCE AND SPREAD OF THE INVASIVE RED ALGA, KAPPAPHYCUSSPP., INKANE'OHE BAY, HAWAI'IAND AN EXPERIMENTAL ASSESSMENT OF MANAGEMENT OPTiONS 231 ABSTRACT 232 INTRODUCTION 233 METHODSANDMATERIALS 236 RESULTS 241 DISCUSSION 245 CONCLUSIONS 250 ACKNOWLEDGEMENTS 251 TABLES 252 FIGURES 254 LITERATURECITED 263 V111 CHAPTER6. INVASNE SPECIES ON CORALREEFS: ANEW THREAT TO DNERSITYAND CORALHEALTH? 268 ABSTRACT 269 INTRODUCTION 270 METHODSANDMATERIALS 272 RESULTS 274 DISCUSSION " "..277 CONCLUSIONS 280 ACKNOWLEDGEMENTS 282 TABLES 283 FIGURES 290 LITERATURECITED 296 CHAPTER 7. FIELD AND NUTRIENT CHARACTERISTICS OF THE EPHEMERAL BLOOM-FORMINGGREEN ALGA CLADOPHORA SERICEA G. HUDSON ON THE REEFS OF WESTMAUl, HAWAI'I. 301 ABSTRACT 302 INTRODUCTION " '" 303 METHODSANDMATERIALS 307 RESULTS 313 DISCUSSION 316 CONCLUSIONS 320 ACKNOWLEDGEMENTS 322 ix TABLES 323 FIGURES 330 LITERATURECITED 340 CHAPTER8. NUTRIENTAND GROWTH DYNAMICS OFHALIMEDA TUNA (ELLIS ET SOLANDER) LAMOUROUX ALONGA DEPTH GRADIENT INTHE FLORIDAKEYS: POSSIDLE INFLUENCE OF INTERNAL TIDES ONNUTRIENT STATUS AND PHySIOLOGy? .346 ABSTRACT 347 INTRODUCTION " 348 METHODSANDMATERIALS .353 RESULTS 358 DISCUSSION................................................................•.......'" 363 CONCLUSIONS '" 372 ACKNOWLEDGEMENTS 374 TABLES 375 FIGURES 382 LITERATURECITED 391 CHAPTER9. SYNTHESIS 401 x LIST OFTABLES TABLE PAGE 1.1 Phase shiftsthat have occurredorare occurring ontropical reefs around the world 66 1.2 Experimental designs andresults for field studies examining factorial nutrient enhancement andherbivore exclusion 70 1.3 Introductionofalgal species inthe tropics for thepurposes ofaquaculture......73 1.4 Summaryofdifferent types ofalgal blooms ontropical reefs and theirsingle factor causes 80 2.1 Dissolvedorganicnitrogenandphosphorus concentrations (J.lM) inthe area adjacent to nutrient enrichmentpots 112 2.2 Results offour-way factorial analyses ofvariance for log transformed algal dry weights 113 2.3 Meanofalgal functional group biomass 114 2.4 Results offour-way factorial analysis ofvariance for sedimentweight and numberofmobile invertebrates 115 2.5 Listofal algal species identified on experimental surfaces andpresenceineach time andtreatmentcategory 116 2.6 Results ofANOSIM test for species communitydata from factorial experiment. 121 2.7 Results ofNPMANOVA test for species communitydatafrom factorial experiment. 122 2.8 Pairwise comparisons forthe significant interactionterm from the NPMANOVAoncommunitydata 123 2.9 DISTLM analysis ofvariance for species communitydata 124 2.10 Results ofNPMANOVA for species composition datagroupedinto functional group categories 125 xi
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