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" UMI Number: 3258317
Copyright 2007 by Rotjan, Randi Dawn
All rights reserved.
UMI Microform 3258317 Copyright 2007 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code.
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c56%4)+,-"$.##6%,>$(,$/(01-)($#+5-9-1#1#$*%225#$CR$G2$3(5-"2+ "#$%0"%.#1#$.%)#;()&$.##6%,>$(,$+$C-).($-#$2''<$-()+/$-(/(,1$%,$W(,+$?2/+,67$Z5#)*($G%-(<$$ $ QU$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ * * C,D:4+(T<(T#(>)+'"%-$6%2*)%45*%(,$(.$-()+//%0()#2<$K*+-3#6$4+)2$-(&'+)#$*"#$,5&4#)$(.$ (4/%>+*#$F;"%*#J$*($.+-5/*+*%0#$F>)#1J$-()+//%0()(52$2'#-%#27$#%*"#)$0#)*#4)+*#$F/#.*$4+)J$()$ %,0#)*#4)+*#$F)%>"*$4+)J<$gD+@%2$(,$+//$>)+'"2$)+,>#$.)(&$:$*($9:$2'#-%#2$%,$%,-)#&#,*2$(.$ 8:<$ V#**#)2$ )#')#2#,*$ >#(>)+'"%-$ 6%2*)%45*%(,$ (.$ -()+/$ )##.$ )#>%(,2_$ AJ$ =+)%44#+,$ +,6$ P#)&56+7$PJ$S+)$^+2*#),$Z+-%.%-$FT+/+'+>(2$+,6$Z+,+&+J7$=J$[+;+%%$+,6$*"#$M(",2*(,$ A*(//7$ YJ$ K(5*"$ Z+-%.%-$ F!+"%*%7$ Z%*-+%),$ ?2/+,627$ W+)C5#2+2J7$ ^J$ `-#+,%+$ FT5+&7$ W%-)(,#2%+7$ W+)2"+//$ ?2/+,62$ A&#)%-+,$ K+&(+7$ S)#,-"$ Z(/1,#2%+7$ =((3$ ?2/+,627$ Z+/+57$ Z+'5+$c#;$T5%,#+7$S%L%7$c#;$=+/#6(,%+7$!(,>+7$K+&(+7$K(/(&(,$?2/+,627$Z+/&1)+$A*(//7$ M+)0%2$ ?2/+,67$ !50+/57$ !(3#/+57$ c()*"#),$ W+)%+,+$ ?2/+,62J7$ SJ$ A52*)+/%+$ +,6$ *"#$ T)#+*$ P+))%#)$ G##.7$ TJ$ ?,6(DZ+-%.%-$ F=(-(2DH##/%,>$ ?2/+,627$ Z"%/%''%,#27$ K%,>+'()#7$ !"+%/+,67$ ?,6(,#2%+7$ W+/+12%+7$ Z+)+-#/$ ?2/+,62J7$ [J$ A2%+,$ Z+-%.%-$ FM+'+,7$ K(5*"$ H()#+7$ !+%;+,7$ [(,>$ H(,>J7$ ?J$ G#6$ K#+7$ MJ$ Z#)2%+,$ T5/.7$ HJ$ A)+4%+,$ K#+$ F`&+,J7$ VJ$ A.)%-+,D?,6%+,$ `-#+,7$WJ$K(5*";#2*$?,6%+,$`-#+,$FK#1-"#//#27$W+5)%*%527$G#5,%(,$?2/+,62J7$cJ$?,6%+,$ `-#+,$FW+/6%0#27$K)%$V+,3+7$?,6%+J7$`J$X#2*#),$A52*)+/%+7$ZJ$K(5*"#),$A*/+,*%-$`-#+,7$ jJ$ W#6%*#))+,#+,$ K#+<$ T#(>)+'"%-$ 6%2*)%45*%(,2$ *+3#,$ .)(&$ G+,6+//$ #*$ +/<$ 8EE9$ +,6$ G+,6+//$ U::a7$ +,6$ +2$ 2'#-%.%#6$ 41$ 2'#-%.%-$ 2*56%#2$ F!+4/#$ 8J$ 6(-5&#,*%,>$ %,-%6#,*2$ (.$ -()+//%0()1<$kc(*#_$G#>%(,2$j$+,6$Z$6($"+0#$2(&#$-()+//%0()#27$45*$)#.#)#,-#2$+4(5*$*"#2#$ )#>%(,2$+)#$2-+)-#7$2($;#$"+0#$)#2#)0#6$>)+'"$2'+-#$*($%,6%-+*#$*"+*$&()#$-()+//%0()#2$+)#$ /%3#/1<$$ $ %/G5+(Q<($A$/%2*$(.$0#)*#4)+*#$+,6$%,0#)*#4)+*#$2'#-%#2$.)(&$+)(5,6$*"#$;()/6$3,(;,$*($ -(,25&#$/%0#$-()+/<$G#>%(,2$F-+*#>()%R#6$41$/#**#)J$-())#2'(,6$*($*"#$&+'$%,$S%>5)#$8<$ `4/%>+*#$ -()+//%0()#2$ F`J$ #+*$ #@-/52%0#/1$ /%0#$ -()+/O$ .+-5/*+*%0#$ -()+//%0()#2$ FSJ$ "+0#$ &%@#6$ 6%#*2$ *"+*$ %,-/56#$ 2(&#$ /%0%,>$ -()+/<$ S##6%,>$ 2*1/#2$ +)#$ -/+22%.%#6$ 4+2#6$ (,$ *"#$ '()*%(,$ (.$ -()+/$ )#&(0#6_$ &5-52$ FWJ7$ -()+/$ *%225#$ F!J7$ ()$ 23#/#*(,$ FKJ<$ S##6%,>$ )+*#$ %,.()&+*%(,$0+)%#2$6#'#,6%,>$(,$*"#$/%*#)+*5)#$+0+%/+4/#$F!+4/#$UJ<$?,$>#,#)+/7$*"#$*+4/#$ ;%//$/%2*$#%*"#)$*"#$+&(5,*$(.$-()+/$)#&(0#6$'#)$6+1$F%,$>$()$-&UJ7$i$4%*#2$'#)$&%,5*#7$()$ b$ /%0#$ -()+/$ %,$ >5*$ -(,*#,*2<$ `4/%>+*#$ -()+//%0()#2$ ;%*"$ ,($ .##6%,>$ %,.()&+*%(,$ -+,$ 4#$ +225$ *($ "+0#$ 4#*;##,$ EID8::b$ /%0#$ -()+/$ .(5,6$ %,$ >5*$ -(,*#,*2<$ $ QQ$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ Corallivores Taxa Region O or F Consumption Rate Style Coral Prey Ref. Invertebrates (52) Annelida (1) Hermodice carunculata A, P, Q F 12.9 cm2 day-1 T Acr, Mil, Ocu, Por 1,2 Arthropoda, Crustacea (10) Alpheus lottini B, J F M Poc 3 Aniculus elegans B F 1.24 g (dry wt) day-1 T, S Poc 4 Calcinus obscurus B T, S Pyrgoma monticulariae G, M, N O T 5 Tetralia glaberrima E, F, J O T, M Acr, Ser 6 Tetralia cavimana F, I, J O T, M Acr 7 Trapezia cymodoce B, F, G, I, J O 1.3-1.5 cm2day-1 T, M Poc, Sty 6,8 T. ferruginea B, E, F, G, J O M Poc 4,6,9 Trizopagurus magnificus B F 10.3 mg day-1 T, S Poc 4 Echinodermata, Asteroidea (10) Acanthaster ellisii B F 145 cm2 day-1 T Pav, Poc, Por, Psa 10 A. planci B, E, F, G, H F 116-187 cm2 day-1 T Acr, Gar, Monti, Poc 11-15 Culcita novaguinaeae C, D, E, G, N F 28 cm2 day-1 T Acr, Poc 16 C. schmideliana L F T Acr, Gal, Gon 17 Echinaster purpureus L F M Por 17 Linckia laevigata L F M Por 17 Nardoa variolata L F M Por 17 Nidorellia armata B F T Pav 18 Pentaceraster cumingi B F T Psa 12 Pharia pyramidata B F T Poc 10,18 Echinodermata, Echinoidea (11) Astropyga radiata L F T 19 Diadema antillarum A F T, S Acr, Aga, Mad, Monta, Por 20,21 D. setosum L F T 19 Echinothrix calamaris L F T 19 Eucidaris thouarsii B F 0.47-1.83 g m2 day-1 T Pav, Poc 11, 18, 22 Echinometra mathaei E, L, M F T 19 E. viridis A F T 23 Echinoneus cyclostomus L F T 19 Microcyphus rousseaui L F T 19 Stomopneustes variolaris L F T 19 Tripneustes gratillia L F T 19 $ Qa$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ Mollusca (20) Aeolidia edmondsoni C O T Por 24 Phestilla melanobrachia E, G, F O Tur 25, 26 P. minor E, L O Por 27 P. sibogae (P. lugubris) C, E, F, G O 6.4 cm2 day-1 T Por 27, 28 Coralliophilla abbreviata A O 0.08–16 cm2 day-1 T Acr, Monta 29-32 C. violacea D, E, G, N 0.25 cm2 day-1 T Por 25, 33 C. caribaea A T Acr, Por 31, 34 Drupella cornus C, E, F, H, I, N O 2.6 cm2 day-1 T Acr, Monti, Poc, Por, Ser, Sty 25, 35-38 D. elata G, H O T Acr, Monti, Poc, Por 37, 39 D. fragum E, H O T Acr, Monti, Poc, Ser, Sty 35, 37, 39, 40 D. rugosa E, F, G, H O 137-229 min day-1 T Acr, Monti, Poc, Ser, Sty 37, 39, 40 Epitonium ulu C T Fun 41 Habromorula spinosa H 42 Jenneria pustulata B O 0.8 g day-1 T Poc, Por, Sid 4 Latiaxis hindsii B T Poc 18, 43 Muricopsis zeteki B T Poc 18, 43 Pedicularia decussata A T Sol 44 Philippia radiata C F T Por 45 Quoyula madreporarum B, E, G, L, M, N O T Monti, Poc, Por, Ser, Sty 25, 46 Q. monodonta B, E, M O 0.64 cm2 day-1 T Monti, Poc, Por, Ser, Sty 43 Vertebrates (111) Chordata, Tetraodontiformes Tetraodontidae (8) Arothron hispidus B, C, E, K, L, M, N F T, S Poc 4, 47, 48 A. meleagris B, C, D, E, G, H, K, L F 10.13 – 16.38 g day-1 T, S Acr, Poc, Por, Pav, Psa, Monti 4, 11, 48-50 A. nigropunctatus E, F, H, I, K, L, M, N F 1.73 bites min-1 T, S Acr, Pav, Poc, Por 47, 51, 52 A. stellatus D, E, F, G, H, I, K, L, M, N F T, S 47, 48 Canthigaster amboinensis B, C, D, E, F, G, H, M, N F 3.7% live coral diet T, S 48, 53 C. jactator C F 1.0% gut contents T, S 53 C. solandri C, E, G, K, L, M, M, N F 11.1% live coral diet T, S 53-55 C. valentini B, E, F, G, H, L, N F 0.06 – 3 bites min-1 T, S Pav, Poc, Por 52 Balistidae (7) Balistapus undulatus D, E, F, G, I, L, M, N F 1.87 bites min-1 T, S Pav, Poc, Por 47, 48, 52 Balistes polyepis B, C F T, S Pav, Por 4 B. vetula A, P F 0.2% gut contents T, S 56 Balistoides viridescens G, M, N F T, S 47 Melichthys niger A, B, C, E, G, H, L, M, N F 0.6% gut contents T, S Col 56 Rhinecanthus aculeatus G, I, K, L, M, N F T, S 48 Sufflamen verres B F T, S Poc, Por, Pav 4 $ Q9$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ Monacanthidae (5) Amanses scopas E, F, G, H, I, L, N O T, S 48 Cantherhines dumerilii B, D, E, F, G, L, M, N F T, S Acr, Poc, Monti, Lep, Por 47, 48, 50 C. pullus A F 0.7% gut contents T, S 56 C. sandwichiensis C, D F T, S 48 Oxymonacanthus longirostris E, F, G, H, L, M, N O 10.5 bites min-1 M, T Acr 47, 48, 51, 57 Chordata, Perciformes Gobiidae (1) Gobiodon citrinus E, F, G, H, I, L, N O 99% gut contents T, M Acr 51 Labridae (8) Coris aygula G, N, O F 0.13 bites min-1 T Pav, Poc, Por 52 Diproctacanthus xanthurus E, F, G F T, S 47 Gomphosus caeruleus N F 0.13 bites min-1 T Pav, Poc, Por 52 Labrichthys unilineatus E, F, G, H, L, M, N O 1.9 bites min-1 T, S Acr, Monti 47, 48, 51, 58 Labropsis australis E, F O T, S 47, 48 L. polynesica E O T, S 48 L. xanthonota E, F, G, H, L, N O T, S 48 Thalasomma lunare E, F, G, H, I, L, N F 0.13 bites min-1 T Pav, Poc, Por 52 Blennidae (1) Exallias brevis G, H, K, L, M, N F 72% gut contents T, S Acr 47, 51 Scaridae (18) Bolbometopon muricatum E, F, G, H, I, L, N F 6.09 bites min-1 T, S Acr, Poc, Por, Mont 47, 48, 59-61 Calotomus carolinus B, C, D, E, F, G, H, L F 0.13 bites min-1 T Pav, Poc, Por 52 Cetoscarus bicolor G, L F 0.4 bites min-1 T, S Por 62 Chlororus gibbus I F 1.1 bites min-1 T, S Por 48, 62, 63 C. microrhinos E, F, G, H F T, S 48, 63 C. sordidus E, F, G, I, K, L, M, N F 2.2 bites min-1 T, S 48, 62 C. strongylocephalus G, L, N F 0.26 bites min-1 T, S Pav, Poc, Por 48, 52 Scarus coelestinus A F 0.2% gut contents T, S 56 S. guacamaia A F T, S S. ghobban B, D, E, F, G, H, I, K, L F T, S Por S. perrico B F T, S Poc 4 S. perspicillatus C F T, S Por 61 S. rivulatus E, F, G, H, O F 1.7 bites min-1 T, S Por 62 S. taeniopterus A F T, S Mad, Por 64 S. vetula A F T, S Monta, Sid 64, 65 S. viridifucatus L F 0.2 bites min-1 T, S Pav, Poc, Por 52 Sparisoma aurofrenatum A F 0.2% gut contents T, S Monta, Por, Mad 56, 64, 65 S. viride A F 25 cm2 day-1 T, S Col, Por, Monta, Sid, Dip, Aga 65-67 $ QI$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ Pomacanthidae (1) Centropyge multispinus L F 0.2 bites min-1 T Pav, Poc, Por 52 Pomacentridae (7) Cheiloprion labiatus F, G, H F 93% gut contents T, S Acr 51 Pomacentrus leucostictus A F 1.5% gut contents T 56 P. semicirculatus E, F, G F 0.13 – .26 bites min-1 T Pav, Poc, Por 52 P. variabilis A F 1.7% gut contents T 56 Plectroglyphidodon dickii D, E, F, G, H, L F 37% gut contents T Acr. Poc 51 P. johnstonianus C, D, F, G, H, L F 96% gut contents M, T Acr 51 Stegastes planifrons A F 0.6% gut contents T Acr, Monta 56, 68 Zanclidae (1) Zanclus canascens/cornutus B, C, D, E, F, G, L, M, N F 0.53 bites min-1 T, S Pav, Por, Poc 52 Chaetodontidae (54) Chaetodon aculeatus A F 0.43 bites min-1 T Aga, Monta, Sid 69 C. andamanensis K, M, N O T Acr 70 C. aureofasciatus E, F, O F T 47, 71 C. auriga E, G, H, I, L, N F 18-60% gut contents T 47, 51, 71, 72 C. auripes H F 12% gut contents T 51 C. austriacus I O 4.4-6.4 bites min-1 T Acr, Fav, Monti, Poc, Por, Ser, Sty 71, 73-75 C. baronessa E, F, G, H O T Acr 47, 51 C. bennetti D, E, F, G, H, L, M O M, T 47, 51 C. capistratus A F 5.4 bites min-1 T Aga, Sid, Mad, Myc 69, 71 C. citrinellus C, D, E, G, H, L, M, N F 15-19% gut contents T 47, 51, 71, 72 C. ephippium C, D, E, G, H F 19-26% gut contents T 51, 71, 72 C. falcula L F 0.26 bites min-1 T Pav, Poc, Por 52, 71 C. flavirostris D, E, F F T 47, 48 C. guttatissimus L F T 71 C. kleinii C, E, G, H, I, L, N F 3% gut contents T 47, 51, 71 C. larvatus I O 10 bites min-1 T 76 C. lineolatus E, G, I, L, N F T 47 C. lunula E, F, G, L, N F 0.2 bites min-1 T Pav, Poc, Por 47, 52, 71 C. lunulatus C, E, F, G, H, O O T 48 C. madagascariensis L F T 71 C. melannotus E, F, G, H, I, L, N F 42% gut contents T 47, 51, 71 C. mertensii D, E, F, G, H, L F T 71 C. mesoleucos I F 6 bites min-1 T 76 C. meyeri F, G, H, L, M, N O T 47, 48 C. multicinctus C F 21.8 bites min-1 T Poc, Por 71, 77 C. ocellatus A F T Acr 31 $ Q]$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ C. octofasciatus G, H O T Acr 78 C. ornatissimus E, F, G, N, O O 2.08-5.62 bites min-1 M, T Monti, Poc, Por 51, 71, 79 C. oxycephalus G, N, O F T 47 C. pelewensis D, E, F F 58.9% gut contents T 47, 71, 72 C. plebeius E, G, H, O O T Acr 47, 51, 71 C. punctatofasciatus M, N F T Poc 47, 80 C. quadrimaculatus C, D, E, G, H F 27.5% gut conents T Poc 48, 71, 72 C. rafflesii E, F, G, H, N F 10% gut contents T 47, 51 C. rainfordi F F T 71 C. reticulatus E, F, G, H O T 47, 71 C. semilarvatus G, H, I, N O 6 bites min-1 T 76 C. smithi D F T 48 C. speculum E, F, G, H, N, O F 75% gut contents T 47, 51 C. striatus A F T Acr 31 C. triangulum N O T 71 C. trichrous D F T 48 C. trifascialis C, E, G, H, I, L, M O 4.58-10.74 bites min-1 T Acr, Poc, Por, Monti 51, 71, 78, 81 C. trifasciatus C, D, E, G, H, I, L, M O 1.1-3.1 bites min-1 T Acr, Pav, Poc, Por, Monti 47, 51, 52, 71, 79 C. ulietensis E, G, H F 10-69% gut contents T 51, 72 C. unimaculatus C, D, E, G, H, L, N, O - 4.9-7.2 bites min-1 T Monti, Poc 47, 51, 71, 82 C. vagabundus E, G, H, L, M, N F 8-18% gut contents T 51, 71, 72 C. xanthocephalus L F T 71 Forcipiger flavissimus B, E, L F 6.2% gut contents T 72 Heniochus chrysostomus D, E, F, G, H, O F 7.1% gut contents T 47, 48, 72 H. intermedius I F 1.3 bites min-1 T 76 H. singularus E, G, H F T 48 H. varius E, F, H, N, O F T 48 $ QE$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ %/G5+()R(G#.#)#,-#2$FA5*"()7$Y+*#J$-())#2'(,6%,>$*($)#.#)#,-#$,5&4#)2$%,$!+4/#$8<$$ ( # Corresponding reference # Corresponding reference 1 (Sussman et al. 2003) 42 (Yokochi 2004) 2 (Witman 1988) 43 (Guzman 1988) 3 (Glynn 1980) 44 (Dall 1889) 4 (Glynn et al. 1972) 45 (Robertson et al. 1970) 5 (Ross and Newman 1969) 46 (Keen 1958) 6 (Knudsen 1967) 47 (Randall et al. 1996) 7 (Glynn 1987) 48 (Randall 2005) 8 (Rinkevich et al. 1991) 49 (Glynn 2004a) 9 (Castro 1978) 50 (Jayewardene and Birkeland 2006) 10 (Dana and Wolfson 1970) 51 (Sano et al. 1984) 11 (Reyes-Bonilla and Calderon-Aguilera 1999) 52 (McClanahan et al. 2005) 12 (Glynn 2004b) 53 (Allen and Randall 1977) 13 (Colgan 1987) 54 (Acero and Rivera 1992) 14 (Chess et al. 1997) 55 (Harmelin-Vivien 1979) 15 (Pearson and Endean 1969) 56 (Randall 1967) 16 (Glynn and Krupp 1986) 57 (Barlow 1987) 17 (Thomassin 1976) 58 (McIlwain and Jones 1997) 18 (Glynn et al. 1983) 59 (Choat 1991) 19 (Herring 1972) 60 (Bellwood et al. 2003) 20 (Bak and van Eys 1975) 61 (Randall 1974) 21 (Glynn 1988) 62 (Bellwood and Choat 1990) 22 (Glynn et al. 1979) 63 (Bellwood 1995) 23 (Griffin et al. 2003) 64 (Frydl 1979) 24 (Ostergaard 1955) 65 (Rotjan and Lewis 2006) 25 (Robertson 1970) 66 (Rotjan and Lewis 2005) 26 (Dalton and Godwin 2006) 67 (Bruckner and Bruckner 1998a) 27 (Ritson-Williams et al. 2003) 68 (Kaufman 1977) 28 (Haramaty 1991) 69 (Neudecker 1985) 29 (Miller 2001) 70 (Kuiter 1999) 30 (Baums et al. 2003) 71 (Harmelin-Vivien 1989) 31 (Brawley and Adey 1982) 72 (Harmelin-Vivien and Bouchon-Navaro 1983) 32 (Ward 1965) 73 (Wrathall 1992) 33 (Oren et al. 1998) 74 (Alwany et al. 2003) 34 (Miller 1981) 75 (Righton et al. 1998) 35 (Cumming 1999) 76 (Zekeria 2002) 36 (Kita et al. 2005) 77 (Tricas 1989) 37 (Turner 1994) 78 (Sadovy and Cornish 2000) 38 (Ayling and Ayling 1987) 79 (Reese 1977) 39 (Moyer et al. 1982) 80 (Neudecker 1979) 40 (Morton et al. 2002) 81 (Irons 1989) 41 (Bosch 1965) 82 (Cox 1986) $ N:$$$=()+//%0()1$+-)(22$*"#$>/(4#$$$$$$ ! ! ! ! ! ! ! ! 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( ZE!!!!3%6%&'#<%!&+$*66#<+$.!-.!(*$$+'/#01%0!! 9.0 8.5 8.0 7.5 7.0 C:N Intact Colonies Intact C:N 6.5 6.0 5.5 5.5 6.0 6.5 7.0 7.5 8.0 8.5 9.0 C:N Grazed Colonies ( T/G<2+(X>(G*#$%)!&+5(*$#0+,!+/!NOM!$*'#+0!/+$!&+$*6!'#009%!'*A%,!/$+5!7$*@%)!<0=!#,'*&'!!+( )'%#&"$*&'!&+6+,#%0!2,!j!DE!(*#$0!+/!&+6+,#%0!6+&*'%)!B#'1#,!D!5!+/!%*&1!+'1%$4=!"*01%)!6#,%! #,)#&*'%0!,966!1.(+'1%0#0!+/!%;9*6!NOM!/+$!7$*@%)!*,)!#,'*&'!&+6+,#%0=!! 45 a a 30 C:N a 15 abc ac ac b bc 0 a lid d ura u etid tact me xa Turf m in li a erp grazed S Ver Barnacle Ha Gal Algae Macroborers Coral ! T/G<2+( Y>( M9'$#'#+,*6! ;9*6#'.! 25%*09$%)! *0! NOM! $*'#+4! +/! (+00#-6%! &+5(+,%,'0! +/! (*$$+'/#01!)#%'0!#,&69)#,7O!-*&A!$%%/!*67*6!0(%&#%0!?)/)@)-#)(0((=!2,!j!E4:!A)/$1&*)!2M!j!E4:! *,)! '9$/! *67*%! 2M! j! J4i! 5*&$+-+$%$! 0(%&#%0! #,&69)#,7! 0%$(96#)! (+6.&1*%'%0! 2,! j! 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All common parrotfish species Sc. vetula 40 Sp. viride Sc. iserti 30 Sp. chrysopterum Sp. rubripinne Sp.aurofrenatum 20 Sc. taeneopterus 2 10 0 B. Known corallivorous parrotfish species 15 10 # # Individuals/240 m 5 0 S eef SG IR OR r USG L ack Lagoon B Habitat ( Q/G3+7(N?*#R*D621%*-"##$%&'()*6+.('%'+(*"0#$((*N*#++&*)"/'%"%(*P,+".(*&$#*#*b*T*%#".(+0%(*'.* +"0)* )"/'%"%* i* G* Og* &$#* %$%"1* -"##$%&'()* 6+.('%3R8* * ;R* a+.('%'+(* $&* F.$5.* 0$#"11':$#$2(* -"##$%&'()*(-+0'+(*P74-)%*'"8'(#%$*:0)74-)/,',6(;*".6*7.-)/($*+%R*"0#$((*#++&*)"/'%"%(8*@"#(*()$5* ,+".*p*G*Og*&$#*#*b*T*%#".(+0%(*-+#*)"/'%"%8* * ( * I_***!"%%+#.(*$&*0$#"11':$#3**** A. Backreef D. Lower Spur and Groove 100 30 75 20 50 10 25 0 0 Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid B. Lagoon E. Inner Reef Slope 40 30 30 20 20 10 10 0 0 Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid C. Upper Spur and Groove F. Outer Ridge Relative Abundance Relative 40 30 30 20 20 10 10 0 0 Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Coral Species ( ( Q/G3+7(P?*X+1"%':+*"/2.6".0+(*$&*H*0$#"1*(-+0'+(*'.*N*)"/'%"%(*"1$.4*%)+*@+1'>+*/"##'+#*#++&* P#SQR8*W+".(**p*G*Og*()$5.*&$#*#*b*T*NK*,J*%#".(+0%(*'.*+"0)*)"/'%"%8** ( ( ( ( ( ( ( ( * IL***!"%%+#.(*$&*0$#"11':$#3**** A. Backreef D. Lower Spur and Groove 1 1 0 0 ) i -1 -1 Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid B. Lagoon E. Inner Reef Slope 1 1 0 0 -1 -1 Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid C. Upper Spur and Groove F. Outer Ridge 1 1 Ivlev's Electivity Index (E 0 0 -1 -1 Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid Coral Species ( Q/G3+7(T?*E++6'.4*(+1+0%':'%3*/3*-"##$%&'()*$.*H*0$#"1*(-+0'+(*5'%)'.*+"0)*$&*N*)"/'%"%(* P#SQR8*M:1+:f(*+1+0%':'%3*'.6+A*Pg'*j*K*'(*-#+&+##+6C*g'*q*K*'(*":$'6+6R*5"(*0"1021"%+6*/"(+6* $.* +"0)* 0$#"1* (-+0'+(f* #+1"%':+* "/2.6".0+* ".6* #+1"%':+* /'%+* &#+[2+.03;* /$%)* ,+"(2#+6* 5'%)'.*"*_KK*,J*"#+"*5'%)'.*+"0)*)"/'%"%8** * ( ( ( ( ( ( * * IT***!"%%+#.(*$&*0$#"11':$#3**** ( ( Q/G3+7()?*E#+[2+.03*)'(%$4#",(*$&*-"##$%&'()*4#">'.4*+A%+.%*-+#*0$1$.3*"0#$((*'.6':'62"1* 0$1$.'+(*$&*_*!"#$%&$'(%*(-+0'+(*'.*:"#'$2(*)"/'%"%(*P#SQR8*@1"0F*%#'".41+(*"1$.4*%)+*A7"A'(* #+-#+(+.%*%)+*,+6'".*4#">'.4*+A%+.%8*a"%"*()$5.*&$#*"11*)"/'%"%(*5'%)*jGK*0$1$.'+(*4#">+6* &$#* "* 4':+.* (-+0'+(8* a'&&+#+.%'"1* 4#">'.4* 5"(* +A",'.+6* 2('.4* "* d$1,$4$#$:7O,'#.$:* 4$$6.+((7$&7&'%*%+(%*%$*"*2.'&$#,*6'(%#'/2%'$.*P%"/1+*GR8** ( ( * * * * IN***!"%%+#.(*$&*0$#"11':$#3**** A. Sp. viride 100 50 0 0 10 20 30 40 B. Sp. aurofrenatum 100 Grazing Incidence (%) * 50 0 0 10 20 30 40 50 60 2 Fish Density (# adults/1200 m ) * ( Q/G3+7(U?*<$##+1"%'$.*$&*-"##$%&'()*4#">'.4*'.0'6+.0+*$.* !"#$%&$'(%*0$#"1(*5'%)*F.$5.* 0$#"11':$#$2(*-"##$%&'()*6+.('%'+(*Pe*&'()*-+#*JLK*,JR*"0#$((*N*#++&*)"/'%"%(^(#R*74-)/,',6(* ".6* ;R* 74-) %*'"8'(#%$*:8* =#'".41+(* ".6* 6$%%+6* 1'.+(* #+-#+(+.%* !-) %##*+%',&C* ([2"#+(* ".6* 6"()+6* 1'.+(* '.6'0"%+* !-) 8%/("+%$%C* 0'#01+(* ".6* ($1'6* 1'.+(* '.6'0"%+* !-) 8'%#9&,8* S'.+"#* #+4#+(('$.*1'.+(*"#+*-1$%%+68* * * * * * * IV***!"%%+#.(*$&*0$#"11':$#3**** a. Sc. iserti b. Sc. vetula Backreef 80 3.0 USG 2.5 LSG 60 2.0 IRS 40 1.5 OR 2004 2004 1.0 20 0.5 0 0.0 0 20 40 60 80 0.0 0.5 1.0 1.5 2.0 2.5 3.0 1982 1982 c. Sp. viride d. Sp. chrysopterum 20 20 15 15 10 10 2004 2004 5 5 0 0 0 5 10 15 20 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 1982 1982 e. Sp. aurofrenatum f. Sp. rubripinne 40 5 4 30 3 2004 2004 2 20 1 10 0 0 10 20 30 40 0 1 2 3 4 5 1982 1982 * Q/G3+7(V?*<$,-"#'($.($&*-"##$%&'()*(-+0'+(*"/2.6".0+(*P#SQR*,+"(2#+6*"(*e*&'()*-+#*JLK* ,J*"0#$((*6'&&+#+.%*#++&*)"/'%"%(*"%*<"##'+*@$5*<"3;*@+1'>+*/+%5++.*GHIJ*P#*b*e*%#".(+0%(;* ,+".*p*G*Og*()$5.R*%$*JKKL*P#*b*e*%#".(+0%(;*,+".*p*G*Og*()$5.R8*a"()+6*1'.+(*#+-#+(+.%* "*1'.+*$&*+[2"1'%3*P.$*0)".4+*'.*-"##$%&'()*(-+0'+(*"/2.6".0+*$:+#*%',+R8* ( ( * II***!"%%+#.(*$&*0$#"11':$#3**** ! ! ! ! ! ! ! ! "#$%!&%$!'()*!+'!+!*,%)-$!'#,%)!,%!%+'(.$/!! #$!0,%1*!,'!+''+2#$1!'&!'#$!.$*'!&0!'#$!3&.-14! ! 5&#%!6(,.! ! ! !"#$%&'()( ( !*+,-./(01++,23.4*(5+16.-5(.708984(/,+1:(+8/,;8+<(1328+(=:81/*.-5(( ! ( >?771+<@( "#$%&! '&(%)*+,-.! +,! /*+)*! )#$%&0! '()#1(! 2+0+'&3! 4%&(! %,5! 634+)%&&3! �(! 6*(+$! 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(4,!6%$(*3!03,2!*$!+,4*#'(%%)!$4#,1*!*-,!4$/,3F!(&(4!'09,3!(4,!94$51!#1!'$%$4!(12!'(1!9,! 3,,1!$//$3#*,!$6!,('-!9%0,!6%$(*L7! ( ! P.0=9+(R:!A&(4!'09,!%(9!(33()7!G(/*#+,!_0+,1#%,!7*,+"4$6,',&+$9+3),#&6!/(44$*6#3-!'-$$3,! 9,*5,,1! (&(4! '09,3! '$1*(#1#1&! !,-"),! (%&(,! Cn! '$1*4$%;! [L;! 7#:*$*-"&6! (%&(,! CS! '$1*4$%;! ',1*,4L;!(12!'$4(%!,Q*4('*!Cb!L!#1!(1!(W0(4#0.7! ! A. Porites porites B. Diploria clivosa ) C. Montastrea franksi, colony 419 2 ) ) 2 2 10.0 15 10.0 7.5 7.5 10 5.0 5.0 5 2.5 2.5 0.0 0 0.0 control (area removed, cm removed, (area control control (area removed, cm control removed, (area 0.0 2.5 5.0 7.5 10.0 0 5 10 15 cm treatment removed, (area 0.0 2.5 5.0 7.5 10.0 treatment (area removed, cm 2) treatment (area removed, cm 2) control (area removed, cm 2) P.0=9+( S:! b,30%*3! $6! ;%,1,44",' 3,(&4(33! 6#,%2! *4#(%37! P(#4,2! /%$*3! '$./(4,! 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M. faveolata M. faveolata 5 5 4 M. faveolata 4 3 4 3 2 3 6 3 1 0 2 1 2 -1 # cubes left cubes # 1 2 1 -2 p = 0.016 left # cubes (M. fav) -3 0 0 Stypopodium difference in # cubes eaten cubes in # difference -4 Stypo Mfav 0 1 2 3 4 5 # cubes left ( Stypopodium) M. franksi M. franksi 4 5 4 M. franksi 4 3 3 2 3 1 2 3 0 2 -1 # cubes left # cubes 1 6 1 -2 # cubes left cubes # (M. fra) p = 0.001 -3 0 5 0 Stypopodium difference in # cubes eaten difference in cubes # -4 Stypo Mfra 0 1 2 3 4 5 treatment # cubes left ( Stypopodium) M. annularis M. annularis 4 5 4 3 M. annularis 4 3 1 2 3 1 2 5 0 2 -1 # cubes left cubes # 1 -2 4 1 # cubes left cubes # (M. ann) -3 p = 0.003 3 Stypopodium 0 0 eaten difference in cubes # -4 Stypo Mann 0 1 2 3 4 5 treatment # cubes left ( Stypopodium) M. cavernosa M. cavernosa 4 5 4 3 M. cavernosa 4 3 2 3 1 2 0 2 -1 # cubes left cubes # 1 2 1 -2 # cubes left cubes # (M. cav) -3 p < 0.001 7 Stypopodium 0 0 difference in # cubes eaten difference in cubes # -4 Stypo Mcav 0 1 2 3 4 5 treatment # cubes left ( Stypopodium) Stephanocoenia intersepts Stephanocoenia intersepts 4 5 4 Stephanocoenia intersepts 3 4 3 3 2 3 1 2 5 0 2 -1 # cubes left cubes # 1 5 -2 1 # cubes left cubes # (Steph) -3 p < 0.001 2 Stypopodium 0 0 difference in # cubes eaten difference in cubes # -4 stypo steph 0 1 2 3 4 5 treatment # cubes left ( Stypopodium) ! P.0=9+(T:!b,30%*3!$6!(&(4!'09,!6#,%2!*4#(%37!P(#4,2!'$./(4#3$13!$6!l!'09,3!4,.(#1#1&!6$4! '$4(%! ,Q*4('*! +,4303! '$1*4$%! (&(4! '09,3! #1! *-,! 6#,%2! (*! \$%2,1! b,,6;! ",%#<,7! 8(.,! 2(*(! '$./(4#3$13!(4,!/4,3,1*,2!#1!*-4,,!2#66,4,1*!6$4.(*3^!%,6*!'$%0.1!3-$53!*-,!*4#(%S3/,'#6#'! #1*,4('*#$13;! *-,! .#22%,! '$%0.1! 3-$53! ,('-! '$4(%! 3/,'#,3! /%$**,2! (&(#13*! (! %#1,! $6! ,W0(%#*);! (12! *-,! 4#&-*! '$%0.1! 3-$53! *-,! 2#66,4,1',! 9,*5,,1! *4,(*.,1*! (12! '$1*4$%! '09,3!/%$**,2!6$4!,('-!*4#(%7!! ! @]J!!!A4,!'$4(%3!'-,.#'(%%)!2,6,12,2B!!! 2.0 tank 1 1.5 tank 2 1.0 0.5 grams removed 0.0 i -0.5 ernosa v control M. franks M. ca Padina (+) control popodium (-) y St ! P.0=9+():(A&(4!'09,!%(9!*4#(%!4,30%*3!*,3*#1&!,Q*4('*3!$6!J!'$4(%!3/,'#,3!(&(#13*!(!1,&(*#+,! '$1*4$%!(%&(!C7#:*$*$-"&6'>$),13L!(12!(!/$3#*+,!'$1*4$%!(%&(!C!,-"),'4,)5#,3B5+&5"4L7!>,(13!v! 87k7!$6!'09,!.(33!4,.$+,2!9)!_0+,1#%,!(4,!/%$**,2!6$4!,('-!*4,(*.,1*7!! ! 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" ( ( Z.A;7*( Y@" " 9&61($" 1(.%" /+.6" %0$" K*,%+.*" U.#$(" TWYW" e*'5$+,&(" L$,%'*-" K*,%+)6$*%" 6$&,)+'*-" ,3$($%&(" 0&+#*$,," F7.61+$,,'5$" (.&#" '*" &5$+&-$" 3'(.-+&6" /.+7$" *$$#$#" %." 1$*$%+&%$" )" 1" %." O" 66" #$1%0G" 2.%0" <'%0'*" F%.1" 1&*$(;" ,1$7'6$*," ?AWG" &*#" 2$%<$$*" F(.<$+"1&*$(;",1$7'6$*,"RA?YG"7.+&("7&(47$,"/.+"&",'*-($" !"#$%&$'%(%",118"7.+&(",3$($%.*" ,&61($8""" " ?N?"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! A. 20 15 10 % C 5 0 Intact Grazed B. 3.5 3.0 2.5 2.0 % N 1.5 1.0 0.5 0.0 Intact Grazed C. 7.5 5.0 C:N 2.5 0.0 Intact Grazed Condition " Z.A;7*(^@""K*'%'&("7.+&("*)%+'%'.*&("C)&('%4"F6$&*"f"9]G"+$1+$,$*%$#"24"#8"1$+7$*%"7&+2.*;" X8" 1$+7$*%" *'%+.-$*;" &*#" !8" :HI" +&%'." /.+" !"#$%&$'%(%" ,118" 7.(.*'$," #',1(&4'*-" -+&D'*-" ,7&+,"24"7.+&(('5.+.),"1&++.%/',0$,"F-+$4G;"&*#"'*%&7%"7.(.*'$,"F<0'%$G8"" " " " " " " " " " " " ?NO"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! A. 11 10 9 8 7 6 % C 5 4 3 2 1 0 Intact Grazed B. 1.75 1.50 1.25 1.00 % N 0.75 0.50 0.25 0.00 Intact Grazed C. 8 7 6 5 4 C:N 3 2 1 0 Intact Grazed Condition " " Z.A;7*( _@" " :.+&(" *)%+'%'.*&(" C)&('%4" F6$&*" f" 9]G" '*" h)*$;" '66$#'&%$(4" /.((.<'*-" 7&-$" +$6.5&(;" +$1+$,$*%$#" 24" #8" 1$+7$*%" 7&+2.*;" X8" 1$+7$*%" *'%+.-$*;" &*#" !8" :HI" +&%'." /.+" !"#$%&$'%(%" ,118" 7.(.*'$," #',1(&4'*-" -+&D'*-" ,7&+," 24" 7.+&(('5.+.)," 1&++.%/',0$," F-+$4G;" &*#"'*%&7%"7.(.*'$,"F<0'%$G8"" " " " " " ?NM"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! A. Control colonies Intact C. Control colonies Intact E. Control colonies Intact 10.0 4 25 p = 0.0441 Grazed Grazed Grazed p = 0.0371 p = 0.0269 p = 0.0066 20 p = 0.0465 7.5 3 * * p = 0.0085 p = 0.0408 * * 15 * 5.0 2 * * % C % N 10 C:N ratioC:N 2.5 1 5 10 8 15 11 15 13 16 13 17 14 13 9 10 8 15 11 15 13 16 13 17 14 13 9 10 8 15 11 15 13 16 13 17 14 13 9 0.0 0 0 4 4 5 04 0 0 0 05 05 06 t 04 t 04 r s ry 05 ry 06 s ry 05 ry 06 st e ry e ry u a a u a a u b a n ber a g u u g u u g o u tober n June 05 tober 04 n June 05 t n Ju m ru a a Au a b Au Oc J Au Oc J Oc J e Febr Febr epte F September 05 September 05 S p = 0.0485 p = 0.0163 * B. Caged colonies * D. Caged colonies F. Caged colonies 10.0 4 25 p = 0.0486 p = 0.0393 p = 0.0101 20 7.5 * * 3 * p = 0.0002 p = 0.0006 15 5.0 2 * * % N % C 10 C:N ratioC:N 2.5 1 5 19 23 5 8 17 19 17 19 15 18 13 13 19 23 5 8 17 19 17 19 15 18 13 13 19 23 5 8 17 19 17 19 15 18 13 13 0.0 0 0 ba c d ba c d ba c d 4 4 5 5 6 4 5 5 5 0 0 0 0 05 04 05 05 05 0 0 0 0 06 r y e r t 04 r r e n er ry ne s e ry e b u ber a ary 0 u b a to J m tob Ju u g o u uary ugust embe n June mb r A August 04 ebr Au a te b Oc Januar Oc Janu pt F Oct J p e epte February 06 e F Month S Month Se Month S " Z.A;7*(`@"I)%+'%'.*&("5&+'&2($,"F6$&*"f"9]G;"'*7()#'*-":HI"+&%'."F#a(XG;"J"*'%+.-$*"F!a( IG;"&*#"J"7&+2.*"F&a(ZG"'*"7.+&("%',,)$",&61($#"/+.6"!"#$%&$'%(%),118"7.(.*'$,"+&*#.6(4" &,,'-*$#"%."7.*%+.("F#a(!a(&G"&*#"7&-$#"FXa(Ia(ZG"%+$&%6$*%,8"9'-*'/'7&*%"('*$&+"7.*%+&,%," &+$" '*#'7&%$#" <'%0" &*" &,%$+',3S" &((" .%0$+" 7.*%+&,%," 1" s" Y8YW8" :'+7($#" (.<$+" 7&,$" ($%%$+," '*#'7&%$"3$4"$5$*%,H"'*'%'&("7&-$"'*,%&((&%'.*"F&G;">)++'7&*$"K5&*"+$6.5'*-"7&-$,"F2G;"7&-$," +$A'*,%&(($#"F7G;"&*#"7&-$,"'*%$*%'.*&((4"+$6.5$#"F#G8"X.+"7.61($%$"$B1$+'6$*%&("#$,'-*" #$%&'(,;",$$"X'-)+$"?8""9$$"L&2($"O"/.+"/)(("OA<&4"!I`_!"6.#$("+$,)(%,8"" " " " ?NT"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! ) A. Control colonies -2 1.6!10 7 intact 1.2!10 7 grazed 8.0!10 6 4.0!10 6 10 8 15 10 14 10 16 14 16 14 13 8 0 Zooxanthellae Density (# cm Zooxanthellae 05 ry 05 e y 06 a ar Jun August 04 tember 05 October 04 Janu p Febru Se ) B. Caged colonies -2 1.6!10 7 1.2!10 7 8.0!10 6 p = 0.0097 4.0!10 6 * 19 23 5 7 17 19 14 1515 18 1314 0 4 ba 4 c 5 5 d 6 Zooxanthellae Density (# Zooxanthellae cm 0 0 0 t 0 0 er y y une ar tob J Augus c anuar O J Febru Month September 05 " " Z.A;7*(b@""l..B&*%0$((&$"#$*,'%4"F6$&*"f"9]G"'*"7.+&("%',,)$",&61($#"/+.6"!"#$%&$'%(%) ,118"7.(.*'$,"+&*#.6(4"&,,'-*$#"%."#8"7.*%+.("&*#"X8"7&-$#"%+$&%6$*%,8"9'-*'/'7&*%"('*$&+" 7.*%+&,%,".*"(.-A%+&*,/.+6$#"#&%&"&+$"'*#'7&%$#"<'%0"&*"&,%$+',3S"&((".%0$+"7.*%+&,%,"1"s" Y8YW8" :'+7($#" (.<$+" 7&,$" ($%%$+," '*#'7&%$" 3$4" $5$*%,H" '*'%'&(" 7&-$" '*,%&((&%'.*" F&G;" >)++'7&*$" K5&*" +$6.5'*-" 7&-$," F2G;" 7&-$," +$A'*,%&(($#" F7G;" &*#" 7&-$," '*%$*%'.*&((4" +$6.5$#"F#G8"X.+"7.61($%$"$B1$+'6$*%&("#$,'-*"#$%&'(,;",$$"X'-)+$"?8""9$$"L&2($"M"/.+"/)((" OA<&4"!I`_!"6.#$("+$,)(%,8" " " " ?NW"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! Intact Grazed A. Control colonies 5 4 3 2 RGB RGB PC1 1 10 11 15 13 16 13 17 14 0 10 8 13 9 t 04 04 05 ne 05 ber tober Ju ugus c A ebruary 06 O January 05 F Septem p = 0.0369 * p = 0.0021 * p = 0.0021 B. Caged colonies * p = 0.0421 5 p = 0.0131 * 4 * p = 0.0062 3 * 2 RGB PC1 1 19 23 8 7 17 19 17 19 15 18 13 13 0 ba c d 04 05 t 04 y tober June 05 Augus c anuar O J February 06 September 05 Month " " Z.A;7*()@"":.+&("7.(.+"F1+'*7'1&("7.61.*$*%"./"ZVES"6$&*"f"9]G"'*"7.+&("%',,)$",&61($#" /+.6"!"#$%&$'%(%),118"7.(.*'$,"+&*#.6(4"&,,'-*$#"%."#8"7.*%+.("&*#"X8"7&-$#"%+$&%6$*%,8" 9'-*'/'7&*%" ('*$&+" 7.*%+&,%," &+$" '*#'7&%$#" <'%0" &*" &,%$+',3S" &((" .%0$+" 7.*%+&,%," 1" s" Y8YW8" !++.<,"+$1+$,$*%",'-*'/'7&*%"#'//$+$*7$,"2$%<$$*"7&-$"&*#"7.*%+.(,8":'+7($#"(.<$+"7&,$" ($%%$+," '*#'7&%$" 3$4" $5$*%,H" '*'%'&(" 7&-$" '*,%&((&%'.*" F&G;" >)++'7&*$" K5&*" +$6.5'*-" 7&-$," F2G;" 7&-$," +$A'*,%&(($#" F7G;" &*#" 7&-$," '*%$*%'.*&((4" +$6.5$#" F#G8" X.+" 7.61($%$" $B1$+'6$*%&(" #$,'-*" #$%&'(,;" ,$$" X'-)+$" ?8" " 9$$" L&2($" T" /.+" /)((" OA<&4" !I`_!" 6.#$(" +$,)(%,8" " ( " ?NR"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! Intact Grazed A. Control colonies 15000 p = 0.0006 * 10000 5000 15 13 10 8 15 11 16 13 17 14 13 9 0 05 05 st 04 r 05 y 06 Holotrich (#/mgdensity dry weight) ber 04 be gu o uary June uar Au an ebr Oct J ptem F Se B. Caged colonies * p = 0.0485 15000 10000 5000 19 23 5 8 17 19 17 19 15 18 13 13 0 ba c d 5 04 04 05 06 t r 0 r Holotrich (#/mgdensity dry weight) s ry 05 e ry u e g be a b a to nu Jun m ru Au c a te b O J p Fe Month Se ( Z.A;7*( c@" " >.(.%+'70" *$6&%.74,%" #$*,'%4" F6$&*" f" 9]G" '*" 7.+&(" %',,)$" ,&61($#" /+.6" !"#$%&$'%(%) ,118" 7.(.*'$," +&*#.6(4" &,,'-*$#" %." #8" 7.*%+.(" &*#" X8" 7&-$#" %+$&%6$*%,8" 9'-*'/'7&*%" ('*$&+" 7.*%+&,%," &+$" '*#'7&%$#" <'%0" &*" &,%$+',3S" &((" .%0$+" 7.*%+&,%," 1" s" Y8YW8" :'+7($#"(.<$+"7&,$"($%%$+,"'*#'7&%$"3$4"$5$*%,H"'*'%'&("7&-$"'*,%&((&%'.*"F&G;">)++'7&*$"K5&*" +$6.5'*-" 7&-$," F2G;" 7&-$," +$A'*,%&(($#" F7G;" &*#" 7&-$," '*%$*%'.*&((4" +$6.5$#" F#G8" X.+" 7.61($%$"$B1$+'6$*%&("#$,'-*"#$%&'(,;",$$"X'-)+$"?8""9$$"L&2($"W"/.+"/)(("OA<&4"!I`_!" 6.#$("+$,)(%,8" " " " " " ?NN"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! A. Control 80000 holotrichs p-basics 60000 spirocysts 40000 20000 Nematocyst Density Nematocyst 0 05 -05 05 06 05 05 -05 06 1- 6 9- 2- 1- 6- 9 2- Intact Grazed B. Caged 80000 60000 40000 20000 Nematocyst Density Nematocyst 0 5 5 5 6 5 5 5 6 0 -0 0 0 0 -0 1- 6 9-0 2- 1- 6- 9 2-0 Intact Grazed " ( Z.A;7*(d@""I$6&%.74,%"#$*,'%4"Ft"u"6-"#+4"%',,)$"<$'-0%S"6$&*"f"9]G"./"0.(.%+'70,"F-+$4G;" 1A2&,'7"6&,%'-.10.+$,"F2(&73G"&*#",1'+.74,%,"F<0'%$G"'*"-+&D$#"5$+,),"'*%&7%"#8"7.*%+.(" &*#"X8"7&-$#"!"#$%&$'(%),118"7.(.*'$,8"\&%&"*.%"&5&'(&2($"/.+"!)-),%"&*#"`7%.2$+"OYYT8"( ( ( ( ( ( ( ( ( ( ( ( ( " ?N@"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! Grazed 75 Intact 50 25 Hardness kgf)(average Hardness 0 within calyx between calyces Sampling Location " ( Z.A;7*(Te@""93$($%&("0&+#*$,,"F&5$+&-$"3'(.-+&6"/.+7$S"6$&*"f"9]G"./"-+&D$#"F-+$4G"&*#" '*%&7%" F<0'%$G" !"#$%&$'%(%" ,118" 7.+&(" ,3$($%.*,;" 6$&,)+$#" <'%0'*A" &*#" 2$%<$$*A" 7.+&(" 7&(47$,8""I"g"RO",3$($%.*,8( ( ( "" " " " " " " " " " " " " " " " " " " " " " " " " ?NQ"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! " " " ! " ?@Y"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""! ! ! ! ! ! ! ! ! ! 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o directly investigate fitness consequences of parrotfish corallivory, I sampled grazed versus intact coral tissue immediately prior to a coral spawning event and dissected out coral gonads and eggs (Chapter 8). 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Q'(&++5<$&++$*,$-"'.$0#.#&03"$"&.$9##($/0'4#($95$&($7(/#0+5'()$'(-#0#.-$'($&22+5'()$ -"#$0#.7+-.$*,$-"'.$#3*+*)'3&+$'(4#.-')&-'*($-*$3*(.#04&-'*($&(/$%&(&)#%#(-$/#3'.'*(.:$$ >+-"*7)"$;$"&4#$2&'(-#/$&$0&-"#0$9+#&N$2'3-70#$*,$2&00*-,'."$'($-"'.$/'..#0-&-'*(<$ 2&00*-,'."$&0#$&+.*$30'-'3&++5$'%2*0-&(-$,*0$%&'(-&'('()$"#&+-"5$3*0&+.$0##,.$-"0*7)"$-"#'0$ "#09'4*0*7.$,##/'()$f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c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c#+'?#&($9&00'#0$0##,:$;$9#+'#4#$-"'.$0#.#&03"$*,,#0.$(*4#+$'(.')"-$'(-*$-"#$ 0*+#$*,$3*0&++'4*05$'($3*0&+$0##,$#3*.5.-#%.<$&(/$;$"*2#$-"'.$0#.#&03"$1'++$'(.2'0#$,7-70#$ .-7/'#.$-*$3*(.'/#0$2&00*-,'."$'($&$%*0#$3*%2+#@$-0*2"*/5(&%'3.$0*+#:$$ $ $ $ $ $ $ $ $ $ $ $ BEG$$$$M7%%&05$&(/$8*(3+7.'*(.$ ! ! ! ! ! ! ! ! "#$!$%&'(!$)(('*!+%&'(!$)(('*! +%&'(!+)#,,#-.!+)(('*! /--!01!-2,(!345!6((7!'(.)8%279!,&)!$)(('!'#8%!.'!$%('(:! +%(!$%!&,!$%(2)!+#,$'!;.'!0#8%!'&,$()!$%.7!'2-<:! /75!$%(1!%.5!$%(!';(($!'0(--!&,!,)('%!6#$$(),-1!02-<:! ! =):!>(#''! ! ! !""#$%&'(!( ! ")**+,-./0(1*)2.31(4*5-5*5365/(+-(7)31*+85(85*/9/(*55-(54.40:,.6( 6+7793.,.5/(+3(;0)<)//.)(,5/,9=.397(/5)1*)//(><)=5/ ( ( INTRODUCTION To gain further insight into parrotfish grazing selectivity, I compared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`!]M!PP!-!FG`!XB!3-&!?G!2(+.)#"9!Q! ! E?A!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!! )*#+!-5#,!47$,#!/$(&"!$+,!,#)#&2(+#,!)*#!7#$3! $&#$! -5#,! 40! /$&&-)3("*#"! ."(+%! Q2$%#!J!;]QI!Q2$%#B!-+!,(%()$7!/*-)-%&$/*"!;$"!(+!N*$/)#&!L!$+,!a(&"1*!#)!$79!EFFEB9!! $9,*.,.+3)<(@9)<.,:( S-!,#)#&2(+#!(3!5$&($)(-+!(+!"#$%&$""!-&!#/(/*0)#!+.)&()(-+$7!b.$7()0!2(%*)!#\/7$(+! /$))#&+"!-3!/$&&-)3("*!%&$8(+%!"#7#1)(5()0M!Q!1-+,.1)#,!#7#2#+)$7!$+$70"("!)-!2#$".&#!c! 1$&4-+! ;NBM! c! +()&-%#+! ;]BM! $+,! NV]! &$)(-! -+! 1-&$7! )("".#! "$2/7#"9! NV]! &$)(-"! *$5#! )&$,()(-+$770!4##+!."#,!$"!$+!(+,(1$)-&!-3!+.)&()(-+$7!b.$7()0!;(#'#,!40!N&-""2$+!#)! $79!EFFFH!(#'#,!40!d.&1#77!$+,!6#77'--,!EFF?B4'$7)*-.%*!)*#0!-+70!1-+"(,#&!)-)$7!;+-)! ,(#)$&0B!1$&4-+!$+,!+()&-%#+9!Q!7--=#,!$)!cN!$+,!c]!"#/$&$)#70!)-!/&-5(,#!$,,()(-+$7! (+"(%*)!(+)-!/$&&-)3("*!%&$8(+%!/#&#+1#"9!! Q! #\$2(+#,! "#$%&$""! 47$,#"! 3&-2! 4-)*! &##3! $+,! 2$+%&-5#! *$4()$)"! '()*! $+,! '()*-.)! )*#(&! &#"/#1)(5#! #/(/*0)(1! 1-22.+()(#"M! $"! '#77! $"! )*#! #/(/*0)#"! )*#2"#75#"9! e/(/*0)#"! '#&#! -5#,! 3&-2! "#$%&$""! 47$,#"! '()*! $! &$8-&47$,#M! $"! $4-5#9! Q! ,&(#,! "#$%&$""! 7#$3! $+,! #/(/*0)#! )("".#! $)! LF-! N! .+)(7! 1-+")$+)! '#(%*)! '$"! $1*(#5#,M! )*#+! )&$+"/-&)#,! "$2/7#"! 4$1=! )-! )*#! 7$4-&$)-&0! 3-&! #7#2#+)$7! $+$70"("9! R77! "$2/7#"! '#&#! $+$708#,!'()*!$!]NCEGFF!e7#2#+)$7!R+$708#&!;Ne!e7$+)#1*B!$+,!&.+!$%$(+")!$+!$//7#! 7#$3!")$+,$&,M!$"!(+!N*$/)#&"!O!$+,!U9! ?,),./,.6)<()3)<:/5/( Q!."#,!/$(&#,!)C)#")"!;:0")$)!??B!)-!1-2/$&#!"#$%&$""!47$,#"!3-&!#$1*!-3!)*#!3-.&! 1-2/$&("-+"! 1-+,.1)#,! ;&##3! "#$%&$""! 47$,#"!^!#/(/*0)#"M!2$+%&-5#!"#$%&$""!47$,#"!^! #/(/*0)#"M! &##3! 5#&"."! 2$+%&-5#! "#$%&$""! 47$,#"! '()*! #/(/*0)#"M! $+,! &##3! 5#&"."! 2$+%&-5#!"#$%&$""!47$,#"!'()*-.)!#/(/*0)#"B9! ( ( ! E?G!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!! #4.40:,.6(6+7793.,.5/( ! e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e/(/*0)#"!1-22-+70!3-.+,!-+!"#$%&$""#"!(+!<7-&(,$!(+17.,#!$! 5$&(#)0! -3! 2$1&-"1-/(1! &#,! $7%$#! ;Y*-,-/*0)$BM! $+,! $! '(,#! ,(5#&"()0! -3! #/(3$.+$! ;R++#7(,$M! 6&0-8-$M! N*-&,$)$M! I0,&-8-$M! $+,! Z-77."1$B! ;<&$+=-5(1*! $+,! <-.&b.&#$+! ?@@UB9!! RESULTS As predicted, I found that /$&&-)3("*! /#&#+)($770! %&$8#! "#$%&$""! 47$,#"! '()*! #/(/*0)#"M!&#%$&,7#""!-3!*$4()$)!;<(%.&#!E!R!$+,!6H!&##3!/$(&#,!)!_!@9EGM!,3!_!?EM!/!f!F9FF?H! 2$+%&-5#!/$(&#,!)!_!E9UAM!,3!_!??M!/!_!F9F?@B9!I-'#5#&M!(+!1-+)&$")!)-!20!#\/#1)$)(-+"M!&##3! /$&&-)3("*! "(%+(3(1$+)70! /#&! "#$%&$""! 7#$5#"! '()*! #/(/*0)(1! 1-22.+()(#"! 3&-2! )*#! 4$1=&##3!*$4()$)M!1-2/$&#,!)-!2$+%&-5#!#/(/*0)#"!;<(%.&#!ENM!/$(&#,!)!_!P9GEM!,3!_!?EM!/!f! F9FF?B9! X*#+! /&#"#+)#,! '()*! 17#$+! 7#$5#"! ;'()*! #/(/*0)#"! -5#,B! 3&-2! 4-)*! 2$+%&-5#! $+,! &##3! *$4()$)"M! /$&&-)3("*! -+! )*#! 4$1=&##3! "*-'#,! +-! "(%+(3(1$+)! ,("1&(2(+$)(-+!4#)'##+!)*#2!;<(%.&#!EgM!/$(&#,!)!_!CF9OPM!,3!_!??M!/!_!F9U?GB9!( R7)*-.%*!&##3!"#$%&$""!47$,#"!'()*!&##3!#/(/*0)#"!$&#!"(%+(3(1$+)70!/#&&#,!40! &##3! /$&&-)3("*#"! (+! #5#&0! 1*-(1#! )#")M! &##3! #/(/*0)#"! $//#$&! )-! 4#! )*#! /--&#")! (+! +.)&()(-+$7!b.$7()0M!$"!)*#0!*$5#!)*#!7-'#")!c!1$&4-+M!)*#!7-'#")!c!+()&-%#+M!$+,!)*#! *(%*#")!NV]!&$)(-"!;<(%.&#!OB9!!Y##3!"#$%&$""!47$,#"!'()*!#/(/*0)(1!1-22.+()(#"!$//#$&! ! E?L!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!! )-!4#!#b.$770!/--&!(+!+.)&()(-+$7!b.$7()0M!4.)!-5$7!-3!)*#!#/(/*0)#"!3&-2!)*#!7#$5#"! ,&$2$)(1$770!(+1&#$"#"!)*#!+.)&()(-+$7!b.$7()0!;$"!2#$".&#,!40!cNM!c]M!$+,!NV]B!-3!)*#! 7#$3!$7-+#9!R"!/&#,(1)#,M!*-'#5#&M!2$+%&-5#!"#$%&$""#"!'()*!-&!'()*-.)!#/(/*0)#"M!$+,! 2$+%&-5#! #/(/*0)#"! $7-+#M! $&#! ".4")$+)($770! *(%*#&! (+! +.)&()(-+$7! b.$7()0! 1-2/$&#,! )-! )*#(&!&##3!1-.+)#&/$&)"!;<(%.&#!OB9!! ! ! ! ! ! E?U!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!! A.19*5(BC!e\/#&(2#+)$7!:#)./9!d$(&"!-3!"#$%&$""!47$,#"!'#&#!17(//#,!(+)-!17-)*#"/(+"!'()*! 3#7)!17-)*!".&&-.+,(+%!)*#!4$"#!-3!#$1*!7#$3M!$+,!)*#+!3#,!)-!/$&&-)3("*#"!3-&!?G!2(+.)#"!(+! $!4$1=&##3!*$4()$)!(+!6#7(8#9!! A. Reef Thalassia: Epiphytes versus none B. Mangrove Thalassia: Epiphytes versus none 15 15 ) ) 2 2 p < 0.001 p = 0.019 10 10 5 5 Reef (-) Reef epiphytes leaf area removed (cm leaf area removed (cm Mangrove (+) epiphytes Mangrove 0 0 0 5 10 15 0 5 10 15 Reef (+) epiphytes Mangrove (+) epiphytes leaf area removed (cm 2) leaf area removed (cm 2) C. Epiphytic Thalassia: Mangroves versus Reef D. Non-epiphytic Thalassia: Mangroves versus Reef 15 15 ) ) 2 p < 0.001 2 p = 0.715 10 10 5 5 leaf area removed (cm leaf area removed (cm Mangrove (-) epiphytes Mangrove Mangrove (+) epiphytes Mangrove 0 0 0 5 10 15 0 5 10 15 Reef (+) epiphytes Reef (-) epiphytes leaf area removed (cm 2) leaf area removed (cm 2) A.19*5(DC!N-2/$&("-+"!-3!!"#$#%%&#'()%(*+&,*-!7#$3!$&#$!-5#,!40!/$&&-)3("*#"!3-&!E!F! &##3! $+,! EGF! 2$+%&-5#! "#$%&$""! 47$,#"! '()*! $+,! '()*-.)! )*#(&! +$).&$7! #/(/*0)(1! 1-22.+()0M! EHF! 2$+%&-5#! 5#&"."! &##3! "#$%&$""! 47$,#"! '()*! $+,! E%F! '()*-.)( )*#(&! +$).&$7!#/(/*0)(1!1-22.+()(#"9! ! E?P!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!! 40 30 20 %C 10 0 3 2 %N 1 0 30 25 20 15 C:N 10 5 0 tes tes lone ytes a h phy phy pi pi e e eaves l w/ w/ epip s s e e Reef epiphytes ove v v a Reef leaves alone e angrove ngr l M a e M Reef lea Mangrov Samples A.19*5(IC!N-2/$&("-+"!-3!+.)&()(-+$7!b.$7()0!;c!1$&4-+M!c!+()&-%#+M!$+,!NV]B!-3! !"#$#%%&#'()%(*+&,*-!7#$5#"!3&-2!2$+%&-5#!$+,!&##3!#+5(&-+2#+)"!$7-+#!$+,!'()*!)*#(&! +$).&$7!#/(/*0)(1!1-22.+()(#"9!].)&()(-+$7!b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`,;! U3! H! ./0DG2! )$=$;$%,:*&;! &6! 3CN$%6*#*,5! 4,7,=$! *;! :9$! 3#5$%,#:*;*,;!#&%,53!91('"+"()(1('"+"-$4!SA!:3'%3'$(),;4!:&'"-$4)(4-'$&"*$4A!OC55$:*;!&6!-,%*;$! >#*$;#$!VG@!DDVFDDY! ! O,T! )'-(! +,;! U<3! Q! ./0YP2! '%$4,:*&;! &6! :9$! 3$,! C%#9*;!8"(*$.()(2-"##('3.! '9*5*NN*! &;! 5*+*;=!#&%,5A!^$#&5&=*,!1G@!///F//P! ! O,T$%!"8!.1GGV2!S5$M*?*5*:<3:$7,:*#3!VE@!KK/FKD0! ! O,T$%!"8!.1GG/2!)$$6!#&%,53!?5$,#9!:&!3C%+*+$!#9,;=$A!X,:C%$!E//@!YKPFYKK! ! O,TC3! 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