The relationship between the herbivore Diadema antillarum and a red crustose alga, Ramicrusta sp., in a shallow back reef community
C. xamachana abundance
M3: C. xamachana with larger bell sizes have lower pulse rates than those with smaller bell sizes.
These hypotheses were used to better understand the behavior and distribution of C. xamachana in the 3$12.04!#!805,5"!35#0;#D$8#@$,#$1-#/"5#B0"!1"/$%#
role as a bioindicator of mangrove health.
Materials and methods
Study site
This study was conducted during the months of N8"0+!.# $1-# F04!3+!.# '()(# 01# "<!# /5%$1-# 0;#
@01$/.!A#%08$"!-#$BB.0O/3$"!%,#P(#93#0;;#"<!#80$5"#
0;# Q!1!R>!%$# /1# "<!# =>"8<# ?$./++!$1&# # D$8# @$,#
6)'S(TUVP&WPXFA# (WPS)YU)W&PWXZ*# /5# "<!# %$.2!5"#
1$">.$%# +$,# /1# "<!# =>"8<# ?$./++!$1# $1-# /5# </2<%,#
protected and recognized for supporting a large
mangrove forest and population of C. xamachana 6D0""#'(()*&#
70.# "<!# B>.B05!5# 0;# "</5# .!5!$.8<A# 5">-,# 5/"!5#
were selected around the perimeter of the bay and
;.03# C4!# 8<$11!%5# 8011!8"!-# "0# "<!# +$,# 67/2&# )*&#
In order to cover the whole length of the bay the sites began at Sorobon, the southernmost point of the bay and extended up to Cai, the northernmost point of the bay, with 300 m of perimeter separating
"<!#/1-/4/->$%#5/"!5&#[002%!#\$."<#J$5#>5!-#"0#3$B#
"<!#5/"!5#$1-#-!"!.3/1!#"<!#[H]#800.-/1$"!5#0;#!$8<#
site. The following channels were selected as sample sites: two channels located adjacent to Cai, one at Kontiki, and two other randomly selected channels between Sorobon and Cai.
Samples were taken from random points within the channel, with two being taken from the channels at Cai and one sample from the other three. These areas were chosen because they contain many shallow channels lined with mangroves as well as numerous C. xamachana.
A total of 28 sites, 20 along the perimeter of the bay and eight within the bay, were chosen and a [$.3/1#[H]#>1/"#J$5#>5!-#"0#3$.9#$1-#.!%08$"!#"<!#
sites after data collection had begun.
C. xamachana abundance
^"#!$8<#5">-,#5/"!A#$#)(#3#".$15!8"#%/1!#J$5#B%$8!-#
running parallel to the mangroves at a constant 2 m distance from the roots. After the transect line was 5!"#/1#B%$8!A#$#)#3#O#)#3#_>$-.$"#J$5#B%$8!-#$"#(#
3A#`#3A#$1-#)(#3#$%012#"<!#".$15!8"&#I<!#1>3+!.#
of C. xamachana of each color (red, blue, green, blue/purple, brown and other) within the quadrat was counted and recorded. The largest and smallest
Fig. 1 ^1#!1<$18!-#/3$2!#0;#D$8#@$,A#@01$/.!A#J/"<#
B/19# 3$.9!.5# "0# /1-/8$"!# 5">-,# 5/"!5&# [.!!1# $.!$5#
surrounding the bay denote the extensive mangrove
;0.!5"5#6[002%!#\$."<#'()(*
43
bell diameters were also measured and recorded (in cm) for each color, as well as the pulse rate (pulses min!"#$ %&'()*+($ +)$ ',-$ ./"/#-$ 012$ 3+'456+4$ 27$
'85(9'(:+$1+6+$54+9$;($)<;4$4)59=>$?+,,=@4<$9+(4;)=$
%(538+6$ 27$ ?+,,=@4<$ 32) and total number of
?+,,=@4<-!"#$%&'()*+#(,,
A+:'54+$ ;)$ <'4$ 8++($ 4<21($ )<')$ '$ 9+:6+'4+$ ;($ )<+$
availability of necessary nutrients can enhance ,+'7$)<;:B(+44$8=$C/!D/E$;($6+9$3'(F62G+4$%H+,,+6$
"IIJ#K$)<+$L6;3'6=$3+'456+$27$3'(F62G+$@)(+44$1'4$
determined by leaf thickness. Leaves were collected from the lowest branches of mangroves at points /$ 3K$ M$ 3K$ '(9$ "/$ 3$ 9;6+:),=$ L+6L+(9;:5,'6$ )2$ )<+$
transects described in the previous section. These leaves were sorted by transect number, brought back to the laboratory, and measured with a caliper, 54;(F$3+)<294$+3L,2=+9$8=$H+,,+6$%"IIJ#-$N<=4;:',$
properties of the mangroves, such as number of O21+64$ '(9$ L62L'F5,+4K$ '(9$ L6+4+(:+$ 27$ :+6)';($
epibionts on their prop roots, such as oysters and 4L2(F+4K$<'G+$8++($'442:;')+9$1;)<$3'(F62G+$@)(+44$
%P+,F'92$+)$',-$.//"Q$P;'*$+)$',-$.//C#-$R($)<;4$4)59=K$
)<+$(538+6$27$L62L'F5,+4$'(9$O21+64$1+6+$:25()+9$
from the top of the trees to the waterline and canopy height was also estimated at the same three points 1<;,+$4)'(9;(F$M$3$7623$)<+$+9F+$27$)<+$3'(F62G+4-$
Similarly, total epibiont coverage at each site was S5',;@+9$ ;()2$ )<6++$ :')+F26;+4>$ 3;(;3',K$ 329+6')+K$
'(9$<+'G=-$T21+G+6K$8+:'54+$)<+$G;4;8;,;)=$;($)<+4+$
areas was poor, the epibiont coverage was estimated 7623$'$9;4)'(:+$27$"$3$7623$)<+$(+'6+4)$3'(F62G+$
roots.
In order to assess environmental and nutrient contributions to the quality of the mangrove ecosystem, water samples were taken from each site and put on ice until they could be analyzed. The salinity and dissolved oxygen content were measured by placing a YSI probe into the water sample and recording the output in ppt and mgL!", respectively.
0<+$LTK$'332(;'$(;)6')+K$',B',;(;)=K$(;)6;)+$(;)62F+($
and nitrate nitrogen were measured using a salt
water aquaculture kit, following procedures set forth 8=$U'V2))+$%.//W#-$
Data analysis
A preliminary ttest was performed in order to 9+)+63;(+$4;F(;@:'()$9;77+6+(:+4$;($?+,,=@4<$9+(4;)=$
between sites in the channels and sites along the perimeter of the bay. A multiple correlation analysis was performed between C. xamachana abundance and the different environmental factors and
;(9;:')264$ 27$ 3'(F62G+$ @)(+44-$X$ ,;(+'6$ 6+F6+44;2($
329+,$ 1'4$ ',42$ :6+')+9$ 54;(F$ )<+$ 324)$ 4;F(;@:'()$
L6+9;:)264$27$?+,,=@4<$'85(9'(:+-$
Possible differences between size and color of C. xamachana, abundance and pulse rate were
examined using ttests with groups based on the different colors. After these tests were performed, the multiple correlation analysis was repeated while grouping the C. xamachana by color (either blue or brown). Correlations between colors, size, and L5,4+$6')+4$1+6+$',42$)+4)+9-$X$@(',$)!)+4)$1'4$54+9$
to determine whether blue and brown colored C.
xamachana abundance differed.
Results
C. xamachana)"-.#/"#0()"#/)1"#$%&'()*+#(,, X$ )2)',$ 27$ "/C$ 8621($ '(9$ 8,5+$ :2,26+9$ ?+,,=@4<$
were observed from the 28 sites surveyed within U':$ A'=-$ P+4L;)+$ )<+$ 7+1+6$ 4'3L,+$ 4;)+4$ ;($ )<+$
channels than along the perimeter of the bay, there 1+6+$ 4;F(;@:'(),=$ 326+ ?+,,=@4<$ 284+6G+9$ 1;)<;($
)<+$ :<'((+,4$ %)!)+4)>$ )YC-""K$ (YDK$ LZ/-//"K$ H;F-$ .#-$
C. xamachana$ 1+6+$ L6+4+()$ ;($ 2(,=$ @G+$ 27$ )<+$ ./$
sites along the perimeter, whereas they were present in seven of the eight channel sites.
C. xamachana abundance was neither correlated with the majority of environmental factors, including LTK$ '332(;'$ (;)62F+(K$ (;)6;)+K$ '(9$ 4',;(;)=K$ (26$
4;F(;@:'(),=$ :266+,')+9$ 1;)<$ :+6)';($ 7':)264$ 27$
3'(F62G+$@)(+44K$45:<$'4$)<+$(538+6$27$O21+64$'(9$
propagules, canopy height, and epibiont coverage.
T21+G+6K$ '$ 4;F(;@:'()$ :266+,');2($ 8+)1++($ )2)',$
number of C. xamachana and leaf thickness was found: the number of individuals increased as leaf )<;:B(+44$ 9+:6+'4+9$ %[Y!/-\I.K$ (Y.DK$ LY/-/\IK$
H;F-$\'#-$];3;,'6,=K$)<+6+$1'4$'$4;F(;@:'()$9+:6+'4+$
in C. xamachana numbers as dissolved oxygen :2()+()$ ;(:6+'4+9$ %[Y!/-C"IK$ (Y.DK$ LY/-/.JK$ H;F$
\8#$ '(9$ '($ 2G+6',,$ ;(:6+'4+$ ;($ ?+,,=@4<$ '4$ 'G+6'F+$
9+L)<$ 27$ 4;)+$ ;(:6+'4+9$ %[Y/-M./K$ (Y.DK$ LY/-//JK$
H;F-$ \:#-$ ^'6;254$ ,;(+'6$ 6+F6+44;2($ 329+,4$ 1+6+$
created using different environmental factors and mangrove components that were considered as L2)+();',$;(9;:')264$27$3'(F62G+$@)(+44K$@(9;(F$)<')$
)<+$324)$4;F(;@:'()$329+,$54+9$82)<$,+'7$)<;:B(+44$
0 1 2 3 4 5 6 7 8
Channel Perimeter
Average density (ind m-2)
Sites
***
Fig. 2 Comparison of mean density (+$ ]P#$
(individuals m2) of Cassiopea xamachana found in channels and along the perimeter of the bay.
];F(;@:'(:+$,+G+,$')$/-/M_K$/-/"__K$'(9$/-//"___
44
and average depth to predict the total amount of C.
xamachana (R2!"#$%&'()!"#""$*#
Color and behavior
In order to better understand the relationship between C. xamachana color and distribution, the aforementioned correlations were tested again, +,-.)/0+( 123( 43556782( 96( :-5-,( ;</+#( =*#( >/?/5@,(
patterns were observed between number of blue 43556782( @0A( 53@B( 12/:C0388( ;D!E"#=%&'( 0!FG'(
)!"#"HH*'( A/88-5I3A( -J6+30( :-01301( ;D!E"#$KG'(
0!FG'( )!"#"$L*( @0A( @I3,@+3( A3)12( -B( 81.A6( 8/13(
;D!"#=L='(0!FG'()!"#"H$*(M23,3@8(9,-M0(43556782(
@9.0A@0:3( -056( 82-M3A( 8/+0/7:@01( :-,,35@1/-0(
M/12( A/88-5I3A( -J6+30( ;D!E"#$K"'( 0!FG'( )!"#"="*(
@0A( @I3,@+3( A3)12( -B( 81.A6( 8/13( ;D!"#=GK'( 0!FG'(
)!"#""G*#( N,-M0( 43556782( @9.0A@0:3( M@8( 0-1(
:-,,35@13A(M/12(?@0+,-I3(53@B(12/:C0388(;D!E"#FK"'(
0!FG'( )!"#H$=*#( O( 81,-0+( :-,,35@1/-0( 931M330( 123(
8?@55381( 95.3( 43556782( ).583( ,@138( @0A( @I3,@+3(
A3)128( M@8( @58-( B-.0A( ;D!"#KK&'( 0!='( )!"#""&*#(
<.,123,?-,3'( 123( @I3,@+3( A308/1/38( ;?2) of blue
@0A( 9,-M0( :-5-,3A( 43556782( M3,3( :-?)@,3A( .8/0+(
@( 1E1381'( 6/35A/0+( 0-0E8/+0/7:@01( ,38.518( ;1!"#=HK'(
0!FG'()P"#"&).
Discussion
C. xamachana abundance was found only to be
associated with mangrove leaf thickness, average depth of the site, and dissolved oxygen content in the water. Similarly, there were very few differences in how these variables affected blue and brown 43556782( @9.0A@0:3#( Q-M3I3,'( 5-:@1/-0( A/A( )5@6(
a major role in the determination of density of C.
xamachana'(M/12(8/+0/7:@0156(?-,3(43556782(B-.0A(
in the channels than along the perimeter. Regression models supported that leaf thickness and average A3)12(M3,3(123(-I3,@55(9381(),3A/:1-,(-B(1-1@5(43556782(
abundance.
C. xamachana !"#$%!$&'(!$%()!$*+,-'(./$'00 N@83A( -0( 123( ,38.518( -B( 12/8( 81.A6'( QH remains )@,1/@556( .0:-07,?3A#( R3556782( @9.0A@0:3( M@8(
8/+0/7:@0156(:-,,35@13A(1-(?@0+,-I3(53@B(12/:C0388'(
which was considered the primary measure of
?@0+,-I3( 710388#( Q-M3I3,'( @9.0A@0:3( M@8( 0-1(
:-,,35@13A( M/12( -123,( @8)3:18( -B( ?@0+,-I3( 710388(
8.:2(@8(0.?93,(-B(),-)@+.538(-,(S-M3,8#(
C. xamachana have been shown to serve as 93012/:( ).?)8( ;R@01T30( 31( @5#( F"H"*'( :/,:.5@1/0+(
0.1,/3018( 03:388@,6( B-,( ?@0+,-I3( 8.,I/I@5#( U.3( 1-(
the fact that mangrove leaf thickness is inversely proportional to the amount of nutrients present
;<3553,(HKKL*'(/1(M@8(3J)3:13A(12@1(12/:C0388(M-.5A(
also be inversely proportional to C. xamachana abundance, which is consistent with the observed
! Dissolved oxygen (mg L-1)
Leaf thickness (mm) Average depth (cm)
Number of jellyfish
a) b) c)
* * **
Fig. 3(>/+0/7:@01(:-,,35@1/-08(-B(C36(30I/,-0?301@5(B@:1-,8(@0A(123(B-55-M/0+(?@0+,-I3(:-?)-03018V(@*(W3@B(
12/:C0388'(9*(U/88-5I3A(-J6+30(@0A(:*(OI3,@+3(A3)12(@0A(123(1-1@5(0.?93,(-B(Cassiopea xamachana#(>/+0/7:@0:3(
53I35(@1("#"&X(@0A("#"HXX
!
Leaf thickness (mm) Dissolved oxygen (mg L-1) Average depth (cm)
a) b) c)
Number of jellyfish
!!! * **
Fig. 4(>/+0/7:@01(:-,,35@1/-08(-B(C36(30I/,-0?301@5(B@:1-,8(@0A(123(B-55-M/0+(?@0+,-I3(:-?)-03018V(@*(W3@B(
12/:C0388'(9*(U/88-5I3A(-J6+30(@0A(:*(OI3,@+3(A3)12(@0A(123(1-1@5(0.?93,(-B(Cassiopea xamachana by color.
>/+0/7:@0:3(53I35(@1("#"&X(@0A("#"HXX
45
!"#$%&#'()(*!"+&"!(,$-."!(/0(1"%%23#4(-+2(5+$#"(+,(
increase in nutrient cycling in mangrove ecosystems +,6(&4"!"0/!"(5/,&!7.$&"(&/(/8"!+%%(-+,*!/8"(3&,"##'(
9/:"8"!;(&47#(-"54+,7#-(7#($,+.%"(&/(6"#5!7."(:42(
7,5!"+#"6(,$-."!#(/0(1"%%23#4(:"!"(+##/57+&"6(:7&4(
poorly oxygenated waters, a phenomenon that was also observed.
<&( :+#( #$!=!7#7,*( &4+&( &4"( ,$-."!( /0( >/:"!#(
+,6( =!/=+*$%"#( :"!"( ,/&( #7*,735+,&%2( 5/!!"%+&"6(
&/( 1"%%23#4( +.$,6+,5"( 6"#=7&"( &4"7!( +##/57+&7/,(
:7&4( #$55"##0$%( -+,*!/8"#( ?@"%*+6/( "&( +%'( ABBCD'(
It was also unexpected that nutrients necessary for mangrove development, such as nitrate and +--/,7+( ,7&!/*",( ?E/&/( +,6( F"%%7,*&/,( CGHID;(
were not correlated to C. xamachana abundance or leaf thickness, even though they have been shown
&/(7,>$",5"(-+,*!/8"(%"+0(&475J,"##(?K"%%"!(CGGLD'(
9/:"8"!;(&47#(#&$62(:+#($,+.%"(&/(&"#&(0/!(=4/#=4+&"(
levels, which have been shown to directly affect -+,*!/8"( 3&,"##( ?M4"!-+,( "&( +%'( CGGHD( +,6( +%#/(
vary in relation to C. xamachana presence (Todd et +%'(ABBLD'(K$&$!"(#&$67"#(#4/$%6(5/,#76"!(-"+#$!7,*(
phosphate levels and investigating the affects of varying levels on C. xamachana distribution in mangrove ecosystems.
N4"( *!"+&"!( +.$,6+,5"( /0( 1"%%23#4( 7,( &4"(
channels than along the perimeter was probably due to the undisturbed water contained within them. The channels were protected from currents by extensive mangrove networks, providing a stagnant environment ideal for C. xamachana
#"&&%"-",&(?9/0-+,,("&(+%'(CGGLD'(N47#(+%#/(+%%/:#(
the deterioration of mangrove leaves to occur unhindered, resulting in greater recruitment of C.
xamachana(?K%"5J(+,6(K7&&(CGGGD'(
Color and behavior
N4"!"(:+#(7,#$0357",&("876",5"(&/(5/,5%$6"(&4+&(C.
xamachana density varied based on color and nutrient availability. Similarly, there was no correlation found between bell size and pulse rates, resulting 7,(&4"(!"1"5&7/,(/0(./&4(92(+,6(93'(9/:"8"!;(&4"!"(
were differences in the correlations between blue and brown C. xamachana in not only the strength of their correlations with dissolved oxygen and average 6"=&4(.$&(+%#/(7,(&4"(#7*,735+,5"(/0(5/!!"%+&7/,(:7&4(
%"+0( &475J,"##O( .!/:,( 1"%%23#4( :"!"( ,/&( 5/!!"%+&"6(
:7&4(%"+0(&475J,"##'(K$!&4"!-/!";("8",(&4/$*4(=$%#"(
rates were not correlated to size, the pulse rates of
&4"(#-+%%"#&;(.%$"(1"%%23#4(:"!"(67!"5&%2(=!/=/!&7/,+%(
to average depth of study site. The increased pulse rates in deeper areas may be a tactic employed to 525%"( &4"( #+-"( +-/$,&( /0( ,$&!7",&#( +#( 1"%%23#4( 7,(
#4+%%/:"!( :+&"!'( E"5+$#"( &4"!"( 7#( %7&&%"( =$.%7#4"6(
research on the effects of C. xamachana color on abundance and nutrient availability, these results can serve as a baseline for future research targeted +&($,6"!#&+,67,*(&4"(!"+#/,#(0/!(8+!7+&7/,(7,(1"%%23#4(
color.
Conclusions
Mangrove mortality still occurs in many places throughout the world due to various causes (Lott ABBCP( Q!7-+8"!+( ABBLD'( 9/:"8"!;( &4!/$*4( -"+,#(
of early detection and active conservation, dieoff can be truncated and restoration can begin to occur
?E/#7!"( "&( +%'( ABBHD'( N4"( ,""6( 0/!( +( .7/7,675+&/!(
species is necessary because the effects of mangrove devastation are typically manifested only after -+1/!(6+-+*"(4+#(/55$!!"6(?R+-+#+J7("&(+%'(CGHSD'(
C. xamachana was chosen for this study because of 7&#(4+.7&+&(=!"0"!",5"(?E"%6+TE+7%%7"("&(+%'(ABBAD(+,6(
previous success as a bioindicator species (Todd et +%'(ABBLD'(
@"#=7&"( &4"( 5/!!"%+&7/,( 0/$,6( ."&:"",(
mangrove leaf thickness and total number of 1"%%23#4;(5/!!"%+&7/,#(."&:"",(+.$,6+,5"(+,6(/&4"!(
measures of mangrove health were not found and more research is needed to determine the role of C.
xamachana as a bioindicator of mangrove health.
9/:"8"!;(&47#(#&$62(4+#(+%!"+62(0/$,6(&4+&(&4"(.%$"(
1"%%23#4;(:4",(5/-=+!"6(:7&4(&4"(.!/:,;(:"!"(-/!"(
#7*,735+,&%2(5/!!"%+&"6(:7&4(%"+0(&475J,"##;(+#(:"%%(
as other environmental factors, which may establish it as a better bioindicator in the future. Similarly, the regression model created, using average depth and leaf thickness serves as an adequate predictor of C.
xamachana( +.$,6+,5"( 7,( U+5( E+2'( N47#( !"#"+!54(
has begun to understand the role of C. xamachana as a major component in mangrove ecosystems and
&4"( +##/57+&7/,( ."&:"",( 1"%%23#4( +.$,6+,5"( +,6(
behavior.
Acknowledgements
<(:/$%6(%7J"(&/(&4+,J(V<WW(X"#"+!54(V",&"!(E/,+7!"(
for the logistical support needed to complete this
=!/1"5&;( "#="57+%%2( @!'( Y+!7+( V'( Z2+!!+( 0/!( 4"!(
continual guidance and help in editing and revising -2( -+,$#5!7=&'( K$!&4"!-/!";( -2( !"#"+!54( =+!&,"!;(
Katie Shoultz, deserves special recognition for her
&7!"%"##("00/!&#(7,(&4"(3"%6'(<(:/$%6(+%#/(%7J"(&/(&4+,J(
STINAPA for allowing us to conduct research within U+5(E+2(+,6(&4"(E/,+7!"([+&7/,+%(Y+!7,"(Q+!J'
46
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-+,*!/8"#( .2( .7!6#( 7,( ^$7,"+TE7##+$'( )!6"+(
__OS_T_I
E"%6+TE+7%%7"( V);( E+7%%7"( E`;( Y+!$2+( N+6+#47(
?ABBAD( M="57357&2( /0( +( -/6"%( 5,76+!7+,T 67,/>+*"%%+&"( #2-.7/#7#'( E7/%( E$%%( ABAO_ITHS E/#7!"( ab;( @+46/$4T^$".+#( K;( F+%&/,( Y;(
V!/,+(E<;(U":7#(XX;(K7"%6(V;(`+7!/(a^;(`/"6+-(
[(?ABBHD(K$,5&7/,+%7&2(/0(!"#&/!"6(-+,*!/8"#O(
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@+46/$4T^$".+#( K;( `/"6+-( [( ?ABBHD(
47
Longterm retrospection on mangrove
development using transdisciplinary
!""#$!%&'()*+*#',-'./*+01!2*3$2*45)46758 9-!:* ;<=* >?-2&* @A=* B12:C'#* @* D866EF* >"$GH'*
species richness and abundance as indicators of mangrove epibenthic community health. Atoll B'(*31CC*IJ4/*>?-2&($G-!G*KG(2-212-$G*L!2-$G!C*
;1('1?*$M*L!21#!C*N-(2$#O
9'CH!P$*A=*N'G('C*AQ=*R-?'G':*R+=*9!O*RS*D866JF*
The importance of propagule establishment and physical factors in mangrove distributional
"!22'#G(* -G* !* <$(2!* B-%!G* '(21!#O/* +01!2* 3$2*
TJ)JITUJT4
VCC-($G* +;=* Q!#G(.$#2&* VR=* W.-CC'O* BA*
DJ55XF* Q!%1C2!2-,'* ?121!C-(?* Y'2.''G* #'P*
mangroves and rootfouling sponges in 3'C-:'!G*;!GH!C/*V%$C$HO*TT)8EZJU8EEE Q!1G%'***<N=*>'#!MO*RV*D866XF*;!GH#$,'(*****!(*****[(&**********&!Y-2!2)*
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