patterns, causes, and consequences of parrotfish corallivory on the ...
Transcript of patterns, causes, and consequences of parrotfish corallivory on the ...
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UMI Number: 3258317
3258317
2007
Copyright 2007 by
Rotjan, Randi Dawn
UMI Microform
Copyright
All rights reserved. This microform edition is protected against
unauthorized copying under Title 17, United States Code.
ProQuest Information and Learning Company
300 North Zeeb Road
P.O. Box 1346
Ann Arbor, MI 48106-1346
All rights reserved.
by ProQuest Information and Learning Company.
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Corallivores
Taxa
Region
O or F
Consumption Rate
Style
Coral Prey
Ref.
Invertebrates (52)
Annelida (1) Hermodice carunculata A, P, Q F 12.9 cm
2 day
-1T 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
-1T, 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 cm
2day
-1T, 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
-1T, S Poc 4
Echinodermata, Asteroidea (10) Acanthaster ellisii B F 145 cm
2 day
-1T Pav, Poc, Por, Psa 10
A. planci B, E, F, G, H F 116-187 cm2 day
-1T Acr, Gar, Monti, Poc 11-15
Culcita novaguinaeae C, D, E, G, N F 28 cm2 day
-1T 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
m
2 day
-1T 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
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 cm
2 day
-1T Por 27, 28
Coralliophilla abbreviata A O 0.08–16 cm2 day
-1T Acr, Monta 29-32
C. violacea D, E, G, N 0.25 cm2 day
-1T Por 25, 33
C. caribaea A T Acr, Por 31, 34 Drupella cornus C, E, F, H, I, N O 2.6 cm
2 day
-1T 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
-1T 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
-1T 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 cm
2 day
-1T 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
-1T, 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
-1T, 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
-1T, S Pav, Poc, Por 52
Balistidae (7) Balistapus undulatus D, E, F, G, I, L, M, N F 1.87 bites
min
-1T, 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
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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
-1M, 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
-1T Pav, Poc, Por 52
Diproctacanthus xanthurus E, F, G F T, S 47 Gomphosus caeruleus N F 0.13 bites
min
-1T Pav, Poc, Por 52
Labrichthys unilineatus E, F, G, H, L, M, N O 1.9 bites min
-1T, 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
-1T 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
-1T, S Acr, Poc, Por, Mont 47, 48, 59-61
Calotomus carolinus B, C, D, E, F, G, H, L F 0.13 bites min
-1T Pav, Poc, Por 52
Cetoscarus bicolor G, L F 0.4 bites min
-1T, S Por 62
Chlororus gibbus I F 1.1 bites min
-1T, 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
-1T, S 48, 62
C. strongylocephalus G, L, N F 0.26 bites min
-1T, 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
-1T, 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
-1T, 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 cm
2 day
-1T, S Col, Por, Monta, Sid, Dip, Aga 65-67
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Pomacanthidae (1) Centropyge multispinus L F 0.2 bites
min
-1T 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
-1T 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
-1T, S Pav, Por, Poc 52
Chaetodontidae (54) Chaetodon aculeatus A F 0.43 bites
min
-1T 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
-1T 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
-1T 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
-1T 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
-1T 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
-1T 76
C. meyeri F, G, H, L, M, N O T 47, 48 C. multicinctus C F 21.8 bites
min
-1T Poc, Por 71, 77
C. ocellatus A F T Acr 31
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C. octofasciatus G, H O T Acr 78 C. ornatissimus E, F, G, N, O O 2.08-5.62 bites
min
-1M, 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
-1T 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
-1T 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
-1T 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
-1T 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
-1T 76
H. singularus E, G, H F T 48 H. varius E, F, H, N, O F T 48
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#
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)
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(T/G<2+( )>( !+( )'%#&"$*&'! &+6+,#%0! #,! '1%! -*&A! $%%/! *'! N*$$#%! ?+B! N*.:! ?%6#@%=!#! G*$'#*66.!7$*@%)! &+6+,.!B#'1! $%&%,'! (*$$+'/#01! 7$*@#,7!5*$A0! 09$$+9,)%)! -.! 6#<%! &+$*6! '#009%=!U(3*5%!&+6+,.!*/'%$!W!5+,'10!B#'1!,+!&+$*6!$%7%,%$*'#+,=!!!Q+'*6!5+$'*6#'.!+/!*!!+()'%#&"$*&'!&+6+,.!)9%!'+!(*$$+'/#01!7$*@#,7=!B!3*5%!&+6+,.!*/'%$!W!5+!&+<%$%)!-.!'9$/!*67*%!B#'1!,+!&+$*6!$%7%,%$*'#+,=!3&*6%!-*$0!#,)#&*'%!J!&5!#,!V:!DK!&5!#,!N=!!((!!!!!
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0 5 10 1
0
5
10
15
5
B.
# m
acr
obo
rers
/ in
tact
col
ony
# macroborers / grazed colony
0 1x10-3
1x10-2
1x10-1
1x100
0
1x10-3
1x10-2
1x10-1
1x100 A.
# l
ive m
acro
bo
rers
/ c
m2 liv
e c
ora
l tissu
e
# holes / cm2 grazed area
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j!ET!&+6+,#%04=!VI#0!-$%*A0!#,)#&*'%! '$*,0#'#+,0! /$+5! 6#,%*$! '+! 6+7! 0&*6%0=! ( U(N+5(*$#0+,!+/!5*&$+-+$%$! *-9,)*,&%!-%'B%%,!0#5#6*$6.!0#@%)!7$*@%)!<0=!#,'*&'!!+()'%#&"$*&'!&+6+,#%0!6+&*'%)!B#'1#,!D!5!+/!%*&1!+'1%$!!2,!j!DD!(*#$%)!&+6+,#%04=!!!!!
!T/G<2+(W>((_%(%*'%)!7$*@#,7!-.!(*$$+'/#01%0!'+!$%5+<%!5*&$+-+$%$0!/$+5!'1%!0*5%!!"#$%&'()'%#&"$*&'!&+6+,.=!#(!"#$%&'()'%#&"$*&'!&+6+,.!B#'1!(*$$+'/#01!7$*@#,7!0&*$0!/+&90%)!+,!&+$*6!'#009%! *$%*0! +/! 1#71! 5*&$+-+$%$! )%,0#'.:! B#'1! 0+5%! $%5*#,#,7! 09$/*&%8<#0#-6%!5*&$+-+$%$0!+,!'1%! 6%/'!0#)%!+/!'1%!&+6+,.=! !U(Q1%!0*5%!&+6+,.!/#<%!)*.0! 6*'%$:!01+B#,7!'1%!09-0%;9%,'!$%5+<*6!+/!5*&$+-+$%$0!-.!(*$$+'/#01%0=!!(
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5.5 6.0 6.5 7.0 7.5 8.0 8.5 9.0
5.5
6.0
6.5
7.0
7.5
8.0
8.5
9.0
C:N
Inta
ct C
olo
nie
s
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=!!
Halim
eda
Galax
aura
Turf
Ser
pulid
Ver
met
id
Bar
nacle
graz
ed
inta
ct0
15
30
45
Algae Macroborers Coral
a
a
a
b bc
abc
ac ac
C:N
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0-10 10-15 15-20 20-26 26-330
5
10
15
Dead
co
lon
ies (
%)
0 5 10 15 20 25 300
25
50
# l
ive s
erp
uli
ds/c
m2
(x10
4)
0 5 10 15 20 25 300
10
20
30
40
Distance from Reef Crest (m)
# e
nd
oli
thic
barn
acle
s/c
m2 (
x10
3)
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Sp.aurofrenatum
Sp. chrysopterumSc. iserti
Sp. rubripinne
Sc. taeneopterus
Sc. vetulaSp. viride
A. All common parrotfish species
Backr
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75
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A. Backreef
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid0
10
20
30
D. Lower Spur and Groove
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid0
10
20
30
40
B. Lagoon
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid0
10
20
30
E. Inner Reef Slope
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid0
10
20
30
40
C. Upper Spur and Groove
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid0
10
20
30
F. Outer RidgeRe
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Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid-1
0
1B. Lagoon
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid-1
0
1E. Inner Reef Slope
Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid-1
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Mann Mfav Mfra Mcav Past Ppor Aagr Dstr Ssid-1
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0 10 20 30 40 50 600
50
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%)
Fish Density (# adults/1200 m2)
*
*
(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****
0 20 40 60 800
20
40
60
80a. Sc. iserti
1982
2004
0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.00
5
10
15
20d. Sp. chrysopterum
1982
2004
0.0 0.5 1.0 1.5 2.0 2.5 3.00.0
0.5
1.0
1.5
2.0
2.5
3.0b. Sc. vetula
1982
2004
0 10 20 30 4010
20
30
40e. Sp. aurofrenatum
1982
2004
0 5 10 15 200
5
10
15
20c. Sp. viride
1982
2004
0 1 2 3 4 50
1
2
3
4
5f. Sp. rubripinne
1982
2004
Backreef
USG
LSG
IRS
OR
*
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0.0 2.5 5.0 7.5 10.00.0
2.5
5.0
7.5
10.0
A. Porites porites
treatment (area removed, cm2)
co
ntr
ol
(are
a r
em
oved
, cm
2)
0 5 10 150
5
10
15
B. Diploria clivosa
treatment (area removed, cm2)
co
ntr
ol
(are
a r
em
oved
, cm
2)
0.0 2.5 5.0 7.5 10.00.0
2.5
5.0
7.5
10.0
C. Montastrea franksi, colony 419
control (area removed, cm2)
treatm
en
t (a
rea r
em
oved
, cm
2)
P.0=9+( S:! b,30%*3! $6! ;%,1,44",' 3,(&4(33! 6#,%2! *4#(%37! P(#4,2! /%$*3! '$./(4,! (4,(! 4,.$+,2!64$.!*-,!'$1*4$%!C,*-,4!$1%)L!+,4303!*4,(*.,1*!C'$4(%!,Q*4('*!n!,*-,4L!;%,1,44",!9%(2,3!#1!@E!.#1!6#,%2!*4#(%3!#1!*-,!9('=4,,6!-(9#*(*!$6!G(44#,!"$5!G();!",%#<,!6$4!C(L!!$+"#34'*$+"#34;!C9L;!<"*1$+",'51"8$4,;!(12!C'L!($)#,4#+,3,'9+,)=4"''$4(%37!!
!@]@!!!A4,!'$4(%3!'-,.#'(%%)!2,6,12,2B!!!
Stypo Mfav0
1
2
3
4
5
M. faveolata
p = 0.016
3
6
1
2
# c
ub
es l
eft
0 1 2 3 4 50
1
2
3
4
5
M. faveolata
# cubes left ( Stypopodium)
# c
ub
es l
eft
(M
. fa
v)
-4
-3
-2
-1
0
1
2
3
4
Stypopodium
M. faveolata
dif
fere
nce i
n #
cu
bes e
ate
n
Stypo Mfra0
1
2
3
4
M. franksi
p = 0.001
3
6
5
treatment
# c
ub
es l
eft
0 1 2 3 4 50
1
2
3
4
5
M. franksi
# cubes left ( Stypopodium)
# c
ub
es l
eft
(M
. fr
a)
-4
-3
-2
-1
0
1
2
3
4M. franksi
Stypopodium
dif
fere
nce i
n #
cu
bes e
ate
n
Stypo Mann0
1
2
3
4
M. annularis
p = 0.003
5
1
4
3
treatment
# c
ub
es l
eft
0 1 2 3 4 50
1
2
3
4
5
M. annularis
# cubes left ( Stypopodium)
# c
ub
es l
eft
(M
. an
n)
-4
-3
-2
-1
0
1
2
3
4M. annularis
Stypopodium
dif
fere
nce i
n #
cu
bes e
ate
n
Stypo Mcav0
1
2
3
4
M. cavernosa
p < 0.001
2
7
treatment
# c
ub
es l
eft
0 1 2 3 4 50
1
2
3
4
5
M. cavernosa
# cubes left ( Stypopodium)
# c
ub
es l
eft
(M
. cav)
-4
-3
-2
-1
0
1
2
3
4
M. cavernosa
Stypopodium
dif
fere
nce i
n #
cu
bes e
ate
n
stypo steph0
1
2
3
4
Stephanocoenia intersepts
p < 0.001
5
2
5
3
treatment
# c
ub
es l
eft
0 1 2 3 4 50
1
2
3
4
5
Stephanocoenia intersepts
# cubes left ( Stypopodium)
# c
ub
es l
eft
(S
tep
h)
-4
-3
-2
-1
0
1
2
3
4Stephanocoenia intersepts
Stypopodium
dif
fere
nce i
n #
cu
bes e
ate
n
!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!!
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M. c
aver
nosa
M. f
ranks
i
Sty
popodium
(-) c
ontrol
Pad
ina
(+) c
ontrol
-0.5
0.0
0.5
1.0
1.5
2.0tank 1
tank 2g
ram
s r
em
oved
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!
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"?N?"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""!
!
Intact Grazed0
5
10
15
20A.
% C
Intact Grazed0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5B.
% N
Intact Grazed0.0
2.5
5.0
7.5C.
Condition
C:N
"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""!
!
Intact Grazed0
1
2
3
4
5
6
7
8
9
10
11A.
% C
Intact Grazed0.00
0.25
0.50
0.75
1.00
1.25
1.50
1.75B.
% N
Intact Grazed0
1
2
3
4
5
6
7
8C.
Condition
C:N
""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""!
!
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Septe
mbe
r 05
Febru
ary 06
0.0
2.5
5.0
7.5
10.0Intact
Grazed
A. Control colonies
10 8 15 11 15 13 16 13 17 14
C:N
rati
o
13 9
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Septe
mbe
r 05
Febru
ary 06
0
1
2
3
4Intact
Grazed
C. Control colonies
10 8 15 11 15 13 16 13 17 14
% N
913
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Septe
mbe
r 05
Febru
ary 06
0
5
10
15
20
25Intact
Grazed
E. Control colonies
10 8 15 11 15 13 16 13 17 14
% C
13 9
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Septe
mbe
r 05
Febru
ary 06
0.0
2.5
5.0
7.5
10.0
B. Caged colonies
19 23 5 8 17 19 17 19 15 18 13 13
Month
C:N
rati
o
dcba
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Sep
tem
ber 0
5
Febru
ary 06
0
1
2
3
4
D. Caged colonies
19 23 5 8 17 19 17 19 15 18%
N13 13
dcba
Month
Aug
ust 0
4
Oct
ober
04
Janu
ary 05
June
05
Septe
mbe
r 05
Febru
ary 06
0
5
10
15
20
25
F. Caged colonies
19 23 5 8 17 19 17 19 15 18 13 13
dcba
Month
% C
* * *
**
p = 0.0441p = 0.0371
p = 0.0269
p = 0.0393p = 0.0486
p = 0.0163
*p = 0.0101
p = 0.0066
*
p = 0.0002
*
p = 0.0085
p = 0.0485
p = 0.0465
*p = 0.0006
**
*
*p = 0.0408
*
"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$$"L&2($"O"/.+"/)(("OA<&4"!I`_!"6.#$("+$,)(%,8""""
"?NT"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""!
!
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Sep
tem
ber 0
5
Febru
ary 06
0
4.0!10 6
8.0!10 6
1.2!10 7
1.6!10 7
B. Caged colonies
19 23 5 7 17 19 14 15 1815 1314
dcba
Month
Zo
oxan
thell
ae D
en
sit
y (
# c
m-2
)
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Sep
tem
ber 0
5
Febru
ary 06
0
4.0!10 6
8.0!10 6
1.2!10 7
1.6!10 7
A. Control colonies
15 10 14 1010 8 16 14 1416 813
Zo
oxan
thell
ae D
en
sit
y (
# c
m-2
)grazedintact
*p = 0.0097
""Z.A;7*(b@" "l..B&*%0$((&$"#$*,'%4"F6$&*"f"9]G" '*"7.+&(" %',,)$",&61($#" /+.6"!"#$%&$'%(%),118"7.(.*'$,"+&*#.6(4"&,,'-*$#"%."#8"7.*%+.("&*#"X8"7&-$#"%+$&%6$*%,8"9'-*'/'7&*%"('*$&+"7.*%+&,%,".*"(.-A%+&*,/.+6$#"#&%&"&+$"'*#'7&%$#"<'%0"&*"&,%$+',3S"&((".%0$+"7.*%+&,%,"1"s"Y8YW8" :'+7($#" (.<$+" 7&,$" ($%%$+," '*#'7&%$" 3$4" $5$*%,H" '*'%'&(" 7&-$" '*,%&((&%'.*" F&G;">)++'7&*$" K5&*" +$6.5'*-" 7&-$," F2G;" 7&-$," +$A'*,%&(($#" F7G;" &*#" 7&-$," '*%$*%'.*&((4"+$6.5$#"F#G8"X.+"7.61($%$"$B1$+'6$*%&("#$,'-*"#$%&'(,;",$$"X'-)+$"?8""9$$"L&2($"M"/.+"/)(("OA<&4"!I`_!"6.#$("+$,)(%,8"""
"?NW"""!:&),$,"&*#"7.*,$C)$*7$,"./"1&++.%/',0"-+&D'*-",$($7%'5'%4""!
!
Augus
t 04
Octob
er 0
4
Janu
ary 05
June
05
Sep
tem
ber 0
5
Febru
ary 06
0
1
2
3
4
5
Intact
GrazedA. Control colonies
10 8 10 11 15 13 16 13 17 14 13 9
RG
B P
C1
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Septe
mbe
r 05
Febru
ary 06
0
1
2
3
4
5
B. Caged colonies
19 23 8 7 17 19 17 19 15 18 13 13
dcba
Month
RG
B P
C1
*p = 0.0131
p = 0.0021p = 0.0369
*p = 0.0421
**
p = 0.0021
*
*p = 0.0062
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!
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Sep
tem
ber 0
5
Febru
ary 06
0
5000
10000
15000
Intact
GrazedA. Control colonies
10 8 15 1115 13
16 13 17 14 13 9
Ho
lotr
ich
den
sit
y (
#/m
g d
ry w
eig
ht)
Aug
ust 0
4
Octob
er 0
4
Janu
ary 05
June
05
Sep
tem
ber 0
5
Febru
ary 06
0
5000
10000
15000
B. Caged colonies
19 23 5 8 17 19 17 19 15 18 13 13
dcba
Month
Ho
lotr
ich
den
sit
y (
#/m
g d
ry w
eig
ht)
*
p = 0.0006
* p = 0.0485
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1-05
6-05
9-05
2-06
1-05
6-05
9-05
2-06
0
20000
40000
60000
80000holotrichs
spirocysts
p-basics
A. Control
Intact Grazed
Nem
ato
cyst
Den
sit
y
1-05
6-05
9-05
2-06
1-05
6-05
9-05
2-06
0
20000
40000
60000
80000B. Caged
Intact Grazed
Nem
ato
cyst
Den
sit
y
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To 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). With these dissections, I established that
parrotfish grazing has major reproductive fitness consequences by /'0#3-+5$ 0#%*4'()$
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!E?G!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!!
#4.40:,.6(6+7793.,.5/(
! e/(/*0)(1! 1-22.+()(#"! %&-'(+%! -+!2$+%&-5#! 5#&"."! &##3! "#$%&$""! 47$,#"!'#&#!
1&.,#70!5(".$770!(+"/#1)#,9!Y##3!"#$%&$""!47$,#"!1-22-+70!*-")#,!1&.")-"#!1-&$77(+#!&#,!
$7%$#M!/0&1217&%!"//9!/-701*$#)#"M!4&0-8-$+"M!$+,!3-&$2(+(3#&$+"9!Q+!1-2/$&("-+M!2$+%&-5#!
"#$%&$""! 47$,#"! *-")#,! $! ".()#! -3! *0,&-(,"! $+,! 3-&$2(+(3#&$+"M! 4.)! *$,! $72-")! +-!
1-&$77(+#! $7%$#9! Q+! 1-77$4-&$)(-+! '()*! :.#! Y(1*$&,"-+M! $! :2()*"-+($+! "/#1($7(")! -3!
"#$%&$""! #/(/*0)(1! 1-22.+()(#"M! #/(/*0)#"! 3&-2! )*("! #\/#&(2#+)! $&#! 1.&&#+)70! 4#(+%!
2-&#!&(%-&-."70!(,#+)(3(#,9!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!!
0 5 10 150
5
10
15
C. Epiphytic Thalassia: Mangroves versus Reef
p < 0.001
Reef (+) epiphytes
leaf area removed (cm 2)
Man
gro
ve (
+)
ep
iph
yte
sle
af
are
a r
em
ov
ed
(c
m2)
0 5 10 150
5
10
15
D. Non-epiphytic Thalassia: Mangroves versus Reef
p = 0.715
Reef (-) epiphytes
leaf area removed (cm 2)
Man
gro
ve (
-) e
pip
hyte
sle
af
are
a r
em
ov
ed
(c
m2)
0 5 10 150
5
10
15
A. Reef Thalassia: Epiphytes versus none
p < 0.001
Reef (+) epiphytes
leaf area removed (cm 2)
Reef
(-)
ep
iph
yte
sle
af
are
a r
em
ov
ed
(c
m2)
0 5 10 150
5
10
15
B. Mangrove Thalassia: Epiphytes versus none
p = 0.019
Mangrove (+) epiphytes
leaf area removed (cm 2)
Man
gro
ve (
+)
ep
iph
yte
sle
af
are
a r
em
ov
ed
(c
m2)
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$,#"!!
0
10
20
30
40
%C
0
1
2
3
%N
Man
grove
leav
es a
lone
Man
grove
leav
es w
/ epip
hytes
Man
grove
epip
hytes
Ree
f lea
ves
alone
Ree
f lea
ves
w/ e
piphyt
es
Ree
f epip
hytes
0
5
10
15
20
25
30
Samples
C:N
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.$7()0!$7"-!$""#""#,!3-&!#/(8--)(#"!$7-+#!3&-2!#$1*!*$4()$)9!]!_!O!"$2/7#"!/#&!1$)#%-&0H!*-&(8-+)$7!7(+#!&#/&#"#+)"!)*#!2#$+!3-&!#$1*!,$)$!17.")#&9!!
!E?@!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!!
!
!EEF!!!:#7#1)(5#!/$&&-)3("*!%&$8(+%!-+!!"#$#%%&#'()%(*+&,*-!"#$%&$""!47$,#"!!
!!!!!!!!
"#!$%&'&!()!*+,(-!./!$%()!01+/&$2!($!()!-./$+(/&3!(/!4+$&'5!!
6.'&/!7()1&8!!
!
!""#$%&'() ((
(
*+,-.(/0!"#$%&'()#*!#+!,-.!,".!&*/!"0-!'&1(#)!&$#*2!3&'(#4)!"&'(556&*!)786'&71(*(&*!7#'&8!)%67(6)9!:88!)&$%86)!;6'6!7#886716/!5<!=&$$6'!&*/!7=()68!+'#$!1=6!>&2##*!=&5(1&1!?@AB!$!/6%1=C!&1!"&''(6!D#;!"&<.!D68(E69!"#'&8!1())46!;&)!&('5'4)=6/!+'#$!1=6!4*/6'8<(*2!)F6861#*!&*/!/'(6/!1#!7#*)1&*1!;6(2=1!&1!GH#"!56+#'6!&*&8<)()!;(1=!&*!-"AIJHH!K86$6*1&8!:*&8<E6'!?"K!K8&*167=C.!'4*!&2&(*)1!&*!&%%86A86&+!)1&*/&'/9!!
"#'&8!L%67(6)! -! ,!-(1'#26*!?ML9K9C! ,!"&'5#*!?ML9K9C! "0-!
!"#$%&%#'#"#$%&%()*' B! B9I@!?H9BHC! II9JN!?@9OIC! P9HG!
!"#$%&%#'()+,%-./%#' I! Q9IB!?H9H@C! IQ9HJ!?H9H@C! J9GN!
!&$.0.$#'&)$1%&.$+%*' B! @9N@!?H9BIC! @@9PQ!?@9OIC! G9JI!
!&$.0.$#'0#/2#(#' Q! @9JJ!?H9IQC! @@9NB!?@9N@C! P9GJ!
3./0.045//%#'+#(#+*' B! I9GG!?H9QIC! @N9JO!?H9OOC! P9IJ!
6%0/.$%#'&/%1.*#' Q! Q9JH!?H9BHC! IP9JO!?@9JIC! G9@J!
6%0/.$%#'/#75$%+(4%-.$2%*' Q! B9JI!?H9NHC! I@9ON!?B9N@C! G9GB!
6%0/.$%#'*($%".*#' Q! B9ON!?H9IIC! IQ9IJ!?@9BQC! G9@H!
6%&4.&.)+%#'*(.8)*%%' Q! B9BH!?H9BPC! IB9QP!?@9JGC! P9IB!
9.+(#*($#)#'#++,/#$%*' Q! I9NI!?H9BGC! I@9JB!?H9QPC! N9IJ!
9.+(#*($#)#')$+.*#' B! Q9GJ!?H9IGC! IJ9OI!?H9O@C! J9GH!
9.+(#*($#)#'-#1)./#(#' B! I9OG!?H9QBC! @O9H@!?I9GOC! G9QQ!
9.+(#*($#)#'-$#+8*%' Q! Q9JO!?H9I@C! IJ9J@!?@9H@C! J9JG!
9%//)0.$#'&.20/#+#(#' Q! @9JH!?H9I@C! O9JQ!?@9IGC! G9QH!
9)#+:$%+#'2)#+:$%()*' @! Q9BI! B@9Q@! P9IP!
;.$%()*'#*($).%:)*' B! @9JQ!?H9BHC! @I9BG!?I9POC! P9OP!
;.$%()*'0.$%()*'' Q! I9O@!?H9HGC! IH9GQ!?H9BPC! P9@H!
<%:)$#*($)#'*%:)$)#' Q! @9JQ!?@9OHC! @I9JJ!?H9PJC! N9BI!
<()04#+.&.)+%#'%+()$*)0(#' Q! I9BQ!?H9I@C! @J9IP!?@9B@C! G9JJ!
!IIB!!!-41'(1(#*&8!R4&8(1<!#+!"&'(556&*!)786'&71(*(&*)!!!!
!IIQ!!!-41'(1(#*&8!R4&8(1<!#+!"&'(556&*!)786'&71(*(&*)!!!!
!!!!!!!!
"!#$%&!'(!)*&!'+&,(!-,.&/!"!*'0&!'(!)*&!1'##$(2!3&&45!
6*&1&!)*&!7+,))&1&3!-,)&17!1,.&8!"(3!)*&!-$(37!)*&$1!1&.!9&&4/!
!:4&7!;,12&()!
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!"#$%&#'%$()"#$*(!
(
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