Lac Bay, Bonaire, Dutch Caribbean
D. antillarum recruitment
Eleven recruitment panels were placed throughout Lac and nine were recovered. Those unable to be recovered were likely disturbed by human
")#$=.$=$)+$& -<$& #,& #:$"=& +*,!$& 4=,3"2"#G& #,&
P,=,Q,)& Y$(+:N& (& 4,4<*(=& (=$(& .,=& /")-!<=Z)7&
and other general leisure activities. Of the nine 4()$*!& =$+,H$=$-N& ZH$& :(-& H"!"Q*$& D. antillarum recruitment, with the most being found on panels nearest to the large population center on the southern
$)-&,.&#:$&Q(+R&=$$.&5D&=$+=<"#!&[&/R1). Mean rate ,.&=$+=<"#2$)#&/(!&>%TD&=$+=<"#!&[&/R1 5PCO0%[\8%&&
Recruitment rates decreased in a distinct pattern circulating in a counterclockwise loop from the population center located on the southern reef (Fig.
T8%&&]$+=<"#!&2$(!<=$-&Q$#/$$)&>F0&22%&B:"!&!#<-G&
does not take into account any microscopic recruits that may have been present on panels.
Discussion
Based on the results of this survey, H1 was supported:
D. antillarum -"!#="Q<#",)& "!& ")^<$)+$-& QG& Q,##,2&
composition and predominantly found on hard substrate. The only dense cluster of urchins was on hard substrate that can accumulate large amounts of (*7($&")&(&=$*(#"H$*G&!:,=#&#"2$&#,&$)!<=$&(-$_<(#$&.,,-&
!<44*G& .,=& #:$& *(=7$& (-<*#& 4,4<*(#",)%& & P"3& !2(**$=&
!
Fig. 2 Recruitment panel in situ on the sandy bottom of Lac Bay
D\
Fig. 3 Map of Lac Bay, with population density (ind m!") represented as different size circles. Each circle is a
#$%&'&($)('&*#$+(,))-.)*/)Diadema antillarum were recorded along a transect, no circle is represented on the map.
0'&%1#)'$%'$#$*(2*3)#24$)52#('267(2/*)&'$)/8$'9&25)*$:()(/)(1$2')+/''$#%/*52*3)+2'+9$,));1$)(1'$$)('&*#$+(#)&9/*3)(1$) northeast corner had the same size distribution, thus only one graph was printed (represented by arrows)
<=
individuals were found on another transect with rubble as the dominant substrate. Juvenile urchins used the loose rubble as protection, allowing them to stay in relative safety. These methods proved to be at least mildly successful, as surveying the rubble ('&*#$+()'$>72'$5)&)9/()?/'$)#$&'+12*3)(/)@*5)(1$)D.
antillarum which were hidden under fragments of Acropora cervicornis and Acropora palmata corals.
A*9B)#2:)D. antillarum were found on soft substrate 1&62(&(#,))C&'5)#76#('&($D)#%$+2@+&99B)/95)5$&5)+/'&9) pavement are a very important habitat for juvenile and adult D. antillarum populations
Prior to the mass mortality event of 1983, population density throughout the Caribbean was magnitudes higher than levels seen today. The mean
population density from multiple studies throughout the Caribbean before the die off (Barbados, Jamaica, E7'&+&/D)F&*&?&D)F7$'(/)G2+/D)H$*$47$9&D)&*5)(1$) IJ)H2'32*)-#9&*5#K)L&#)<,MN)2*5)?!")OP$29)$()&9,)"==QK,))
;1$)@*52*3#).'/?)(12#)#(75B)2*)R&+)'$#79($5)2*)&)?$&*)
%/%79&(2/*)5$*#2(B)/.)=,=M"S)2*5)?!", or >99% less D. antillarum than levels seen in the past in Curacao.
This difference does not indicate that populations are recovering, despite literature indicating the /%%/#2($,) ) -*) E7'&+&/D) %'$#$*() %/%79&(2/*#) 1&8$) begun to indicate an increasing population. This does not appear to have the same effect on Bonaire, despite the interconnectedness of the two islands.
;1$'$) &'$) ?79(2%9$) %/##269$) $:%9&*&(2/*#) ./') (12#)
#23*2@+&*()%/%79&(2/*)52..$'$*+$,))A*$)+/795)6$)(1&()
!
"!
#"!
$"!
%"!
&"!
'"!
("!
)*! )*)+! )*,-! ./0! 1*2!
!"#$%&!"#$%&'(($)*+&"'()*+),&
-.'(#*$#)&#/0)&
Fig. 4 Total number of Diadema antillarum present on each of the substrate types surveyed: sand (Sa), sand/seagrass (SaSe), seagrass (Se), seagrass/algae (SeAl), rubble (Rub), and pavement (Pav). There
!"#"$%&'(&)*+(,-.$/0#"$1#*2&(%$0($2+#3$%14%,#+,"%$
(Pav and Rub) compared to soft substrates (Sa, 5+5"6$+(3$5+7-8$9:;<=<<>8
!"
#!"
$!"
%!"
&!"
'!"
(!"
)!"
*#"+" #,$"+" $,%"+" -%"+"
!"#$%&!"#$%&'(($)*+#"'()*+),&
-).#/&
Fig. 5 Total number of individual Diadema antillarum recorded throughout the study in relation to depth in which they were found
Table 1 Total Diadema antillarum found by size classes in comparison to substrate type. Substrate
*-+%%&)*+,&0($&%$,2"$/+?0#&,.$%14%,#+,"$@01(3$0($"+*2$
,#+(%"*,$!&,2$"A*":,&0(%$@0#$%+(3B%"+'#+%%$+(3$%+(3B algae, which determining the majority substrate was not possible
*0(31*,"3C$ ,2"$ D"%,$ E(3&+($ 5"+$ F''$ 9Tripnuestes ventricosus). This species is also herbivorous, and /&'2,$)--$,2"$%+/"$(&*2"$0($#""@%$+%$D. antillarum
&($ G+/+&*+$ 9H0%"%$ +(3$ I0("/$ J<<>8=$ $ 7@,"#$ D.
antillarum populations decreased during the mass mortality, T. ventricosus took its place as one of the dominant herbivores on the reef after 15 years of this niche being absent from reef communities. Because of the still stagnant D. antillarum population, T.
ventricosus has established itself as one of the main benthic herbivores in Lac and throughout other parts 0@$ ,2"$ K+#&44"+($ 9H0%"%$ +(3$ I0("/$ J<<>8=$ $L2&%$
observation suggests that this niche might be species independent—the species of herbivore may not matter in terms of the “keystone herbivore”, but that
“keystone niche” may be a more appropriate term when referring to the effect D. antillarum or other benthic herbivores have on coral reef communities.
Recruitment rates were similar to those seen
&($ K1#+*+0$ 9M"#/"&?$ ",$ +-=$ J<><8=$ $ M"#/"&?$ ",$ +-=$
9J<><8$ @01(3$ +$ #"*#1&,/"(,$ -0!$ 0@$ <=<NO$ #"*#1&,%$
wk1, and a high of 11.5 recruits wk1. Results from data collected in Lac showed the recruitment had a low rate at zero recruits wk1, while the high was at J=PP$#"*#1&,%$!Q1. The results of this study support ,2"$)#%,$:+#,$0@$RJ; the recruitment rates are similar to that of neighboring Curacao. However, the
&3"+$,2+,$#"*#1&,/"(,$!01-3$4"$2&'2"#$&($2&'2$S0!$
areas of the reef was not supported. The highest point of recruitment was adjacent to the population center of adult D. antillarum. This suggests that the population within Lac Bay is selfrecruiting, or that recruits are cueing into adults or the particular hardbottom habitat. The counterclockwise trend of decreasing recruits from the population center 9T&'=$U8$+-%0$%20!%$+$#13&/"(,+#.$!+,"#$S0!$*.*-"6$
indicating that while water movement from outside the bay pushes larvae from the spawning population center toward the back of the bay, there is also a
%&'(&)*+(,$ *1##"(,$ +-0('$ ,2"$ 4+*Q$ #""@$ +--0!&('$
recruitment of D. antillarum larvae to hardbottom habitat.
Comparing these data to rates pre and post die off rates in Curacao indicates that recruitment has begun to increase. Mean recruitment rates pre
>VNP$!"#"$+::#0A&/+,"-.$><$,&/"%$2&'2"#$,2+($,20%"$
reported in this study (Bak 1985). However, post
1983 die off, those values dropped to almost zero indicating that the population of D. antillarum had 3#0::"3$ %0$ %&'(&)*+(,-.$ ,2+,$ (0$ %1**"%%@1-$ -+#W+"$
were being formed, or those that did form were not able to recruit successfully. This gives indication that D. antillarum is starting to recover, with recruitment rates much higher than those reported after the mass mortality event.
Despite the increasing recruitment rates, the low population density indicates that the recruits are not
%1#W&W&('$,0$#"+*2$%"A1+-$/+,1#&,.=$$L2#01'201,$,2"$
<2 cm 2-5 cm 5-10 cm >10 cm
Sand 1 0 0 1
Seagrasses 0 0 0 0
Sand/Seagrass 0 2 1 0
Sand/Algae 0 1 0 0
Rubble 1 4 1 0
Pavement 0 0 3 54
the entire bay was surveyed instead of randomly selected points on open water reef environments.
The bay in its entirety was chosen to be studied because historically, D. antillarum remained hiding in the recesses of corals during the day, and migrated to feed in seagrass beds in the afternoon and into the night (Kaplan 1988). This feeding behavior might not be what the current population in Lac follows, possibly due to the travel distance between the dense population cluster on the back reef and the start of seagrass beds, or due to better feeding opportunities 0($,2"$#""@$!2"#"$-"%%$"("#'.$/1%,$4"$"A:"(3"3$,0$
#"+*2$+($+3"X1+,"$@003$%1::-.=
E,$&%$&(,"#"%,&('$,0$(0,"$,2+,$+$%":+#+,"$%:"*&"%$
of urchin was present in almost every transect
81
surveys carried out in Lac, D. antillarum predators
!"#"$%&'$(""%$)%$*%+$(),%)-.*%'$/"%()'+0$$1&!"2"#3$
it is known that urchin predators, such as octopus
*%/$'#),,"#-(4$/&$)%4*5)'$6*.$7*+0$$8()/"$9#&:$;&!$
recruitment levels, it is unclear what factors might 4*2"$ *%$ )%<="%."$ )%$ D. antillarum population and recovery efforts. Perhaps the Allee effect proved too much to overcome following the 1983 mass mortality event. The now lower population will never have a chance to reach a stage of recovery.
Other environmental factors could also have an )%<="%."$ )%$ D. antillarum survivability. During the second week of the recruitment panels being in place, Hurricane Tomas passed through the Caribbean, creating irregular weather and water patterns throughout the island, undoubtedly affecting Lac Bay along the course. The pattern of increased storm activity in relation to global climate change will certainly have an effect on coral reef
communities, with strong storms (such as Hurricane
>:*#$)%$?@@AB$/"2*('*')%,$.&#*;$.&2"#$&%$:&('$C*#'($
of Bonaire.
This study has local and global implications.
Contrary to most literature on the subject, D.
antillarum population in this study has not started to (4&!$*%$)%.#"*("$)%$/"%()'+$*9'"#$*%$*;:&('$D@$+"*#$
timeframe in which recovery should have started '&$ &..=#E$ C#&:C')%,$ '4"$ F="(')&%G$ H4+$ )($ 7&%*)#"$
different from most of the rest of the Caribbean?
Bonairian reefs are among the most pristine and well preserved in the Caribbean, yet an absence of one of the main herbivores implies that coral cover should be less than those on neighboring islands. This is not the case, however. Bonaire has mostly coral dominated reefs, as opposed to algal dominated reefs seen on many other islands due to a recent increase in harmful human activity. Perhaps Bonaire’s reefs 4*2"$ -;;"/$ '4"$ %).4"$ ;"9'$ 5+$ D. antillarum with another hidden functional group, or perhaps grazing -(4"($ *#"$ '4"$ :*)%$ 4"#5)2&#"$ &%$ '4"$ #""9(0$ $ I&#"$
work needs to go into this facet of coral reef ecology to pinpoint the true main herbivore, yet it is clear that D. antillarum still has many more years of slow recovery before it can establish itself as the keystone herbivore on the reef once again.
Acknowledgements
J)#('$&9$*;;3$K$!&=;/$;)L"$'&$'4*%L$M#0$N)'*$O"*.4"+$
*%/$'4"$#"('$&9$'4"$PKQQ$('*99$9&#$'4")#$,=)/*%."$*%/$
9&#$ :*L)%,$ '4)($ C#&,#*:$ C&(()5;"0$ $ K$ !&=;/$ *;(&$
like to thank Marissa Paulling and Paul Patitsas for always being fantastic research buddies. Another 5),$ '4*%L($ '&$ 8;"R$ 6":*%S6*!#)"$ 9&#$ /#)2)%,$ '4"$
boat so faithfully on the Wednesday and Saturday 6*.$ #"("*#.4$ /*+($ T*;(&3$ K$ *:$ (=#"$ +&=$ really disliked spending eight hours a day getting a tan).
U4*%L$ +&=$ '&$ *;;$ '4"$ PKQQ$ J*;;$ VW@$ ('=/"%'($ 9&#$
their guidance on this paper. Finally, thank you XUKY8O8$9&#$;"'')%,$=($=("$'4")#$5&*'$*%/$;"'')%,$=($
conduct research in Lac.
!"
#"
$"
%"
&"
'"
("
)"
*"
#" $" %" &" '" (" )" *" +"
!"#$%&'()'&%*&"+,-')("./'
01.%2'3&(4+#+,5',('!"#$%&'(($)*+#3(3"21,+(.'*%.,%&''
Fig. 6 a) Map of Lac Bay with circles placed at the site of each recruitment panel. Larger circles indicate a ,#"*'"#$%=:5"#$&9$#".#=)'(3$5B$K;;=('#*'"($'4"$'#"%/$&9$
decreasing recruitment rates in relation to distance from dense Diadema antillarum population center.
Circles from the map (a) are labeled with a number, which corresponds to the number of recruits (b)
a)
b)
A?
References
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TU*4)')B$[G?\ZS?Z?
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#""9$4"#5)2&#"(0$O#&.$['4$K%'$P&#*;$N""9$X+:C$
TU*4)')B$]G[DS\@
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!"#$ %&'()*+$ ,--".)#/"$ "00"'/!$ 1*$ '1--%*)/2$
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3"4&1/$56$#*.$7#8"9:"&:"*$,$;<==>?$@"'1A"&2$10$
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31&"*41!'($ DL$ A#*$ @")9$ DML$ 7#8"9:"&:"*$ ,L$ A#*$
."&$ N"9."$ O$ ;<==P?$ Q("$ &"9#/)1*!()R$ 10$ &""0$
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3%-#!$VL$W%94)':)$DL$M()0X"/$EL$Y)'("Z$@L$Y"&&#&)!$
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&""0!$ ;7"]$ M#9".1*)#L$ E1%/($ V#')S'?B$ [$ \TR$
D#&$C)19$\'19$PHH+UUKI==
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D#&$\'19$V&18$E"&$HJ+I=GKIIG$
[1-R#$ [$ #*.$ D'M11:$ ^[$ ;<==<?$ \00"'/!$ 10$
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D#&$C)19$\'19$<FI+<JKPG
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following mass mortality of Diadema antillarum$ V()9)RR)$ #/$ E#)*/$ [1(*L$ `BEB$ N)&8)*$
,!9#*.!B$[$\TR$D#&$C)19$\'19$IIG+I>FKIFU
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_#&-0%9$598#"$J+J>JKJUJ
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@"*:"*$_L$a#4*)/Z$MMML$^9"]"992*$O$;<==H?$
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H<F+JPPKJP>
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\'19$D1.$IG>+IPIKIHU
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N"&-")b$ D[5L$ 3"4&1/$56L$ A#*$ ."&$ _#9$ 7L$ C#::"&$
[L$ C#:$ @VD$ ;<=I=?$ ,*'&"#!".$ &"'&%)/-"*/$
rates indicate recovering populations of the sea urchin Diadema antillarum on Curacao. Bul D#&$E')$U>+FIGKF<J
a")9$ \L$ Q1&&"!$ [^L$ 5!(/1*$ D$ ;<==J?$ V1R%9#/)1*$
characteristics of the sea urchin Diadema antillarum )*$ ^#$ V#%"&#L$ V%"&/1$ @)'1L$ IF$
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83
!"#$%&'()*&'%*+,)-),)'()*#-*.+)($/(*(#0#,.*&'%*.12.3,&3).*24*35)*6)&(#(7*
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Beth Tyrie Wofford College Spartanburg, South Carolina
tyrieek@email.wofford.edu
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! 0
20 40 60 80 100 120 140 160 180
Average Time Spent Swimming 10 min-1 + SD
0 50 100 150 200 250 300 350
Average Time Spent Swimming 10 min-1 + SD
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! !
! !
200.0 200.5 201.0 201.5 202.0 202.5 203.0
Animal Annulus
Mean Intensity
0.00E+00 2.00E-08 4.00E-08 6.00E-08 8.00E-08 1.00E-07 1.20E-07 1.40E-07 1.60E-07
1 2 3 4 5 6
Energy Per Pixel
Band Number
Animal Annulus
a) c)
b) d)
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! !
! !
!
100 120 140 160 180 200 220 240
Animal Annulus
Mean Intensity
0.00E+00 1.00E07 2.00E07 3.00E07 4.00E07 5.00E07 6.00E07 7.00E07
1 2 3 4 5 6
Energy Per Pixel
Band Number
Animal Annulus
a) c)
b) d)
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References
?&=*($ ME<$ :'--0$ E<$ ?&(*,,$ ER<$ ?'-"*/*+$ ?F<$
F#+,"#--$ SR$ TUVVUW$ ;+*0X)*2*()*(!$ 7&+#8/(8$
!#1!/1,$ #()$ 2+*0$ 2+&Y!#4/-/!0$ /($ #$ 3#+/(*$
3#33#-5$F#+$N1&-$;+&8$L*+$UZZ[U\]XUZ]
>"/#&$>><$>"'44$><$?'+*,1"$^><$?#+4&,#$6<$6--*($
RR<$ F_!"8*+$ KF<$ J#(-&($ I:$ TUV`VW$ F&!!-*$
1#3&'%#8*$ 2#!!*+(,$ /($ 1'!!-*Y,"[$ a'#(!/!#!/.*$
1"#+#1!*+/@#!/&($#()$./,'#-$4#198+&'()$,!/3'-/$
!"#!$*.&9*$!"*35$R$NC2$?/&-$U`\[`bcX`dd
>&--/*$ RL$ T`dbcW$ P&&)$ 1&(,'32!/&($ 40$ 0*--&=!#/-$
%&'()*+$/($+*-#!/&($!&$2+&)'1!/&($&7$/!,$4*(!"/1$
2+*05$F#+$N1&-$;+&8$L*+$\e[UV]XU`\
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Rachael Vietheer Ursinus College Collegeville, Pennslyvania
ravietheer@ursinus.edu
Introduction
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Abstract
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