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Diversity of Zoantharia (Anthozoa: Hexacorallia) at Dongsha Atoll in the South China Sea
Accepted Manuscript Diversity of Zoantharia (Anthozoa: Hexacorallia) at Dongsha Atoll in the South China Sea James Davis Reimer, Maria Eduarda Alves Santos, Hiroki Kise, Mei Lin Neo, Chaolun Allen Chen, Keryea Soong PII: DOI: Reference:
S2352-4855(16)30229-8 http://dx.doi.org/10.1016/j.rsma.2017.02.006 RSMA 225
To appear in:
Regional Studies in Marine Science
Received date: 11 October 2016 Please cite this article as: Reimer, J.D., Santos, M.E.A., Kise, H., Neo, M.L., Chen, C.A., Soong, K., Diversity of Zoantharia (Anthozoa: Hexacorallia) at Dongsha Atoll in the South China Sea. Regional Studies in Marine Science (2017), http://dx.doi.org/10.1016/j.rsma.2017.02.006 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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,8::.;987-270*=<18: ABSTRACT 870;1*<8552;<1.5*:0.;<,8:*5*<85527<1.%8=<1127*%.*%%*7-2;.@9.,<.-<8 1*:+8:12015.>.5;8/+28-2>.:;2.:;2.:-=.<82<;:.5*<2>.2;85*<287<1. +28-2>.:;2.;<20*<.-/8:68;<6*:27.<*@* 7<12;;<=-A ?.:.98:<87/27-270;/:868=:;=:>.A27 <1*</8,=;.-87<1.8:-.:)8*7<1*:2*/8:?12,1875A ;98:*-2,9:.>28=;:.98:<;.@2;</:86<1.%%"=::.;=5<;27-2,*<.;86.;9.,2.;;=,1*;Palythoa tuberculosa*:.>.:A,86687*:8=7-870;1*<855*;?.55*;<1.9:.;.7,.8/*<5.*;<<1:.. =7-.;,:2+.-;9.,2.;27,5=-27087.;9.,2.;875A/8=7-<1=;/*:*<<1.*<855">.:*55?.:.,8:-.- ;9.,2.;8/)8*7<1*:2*/:86870;1*<855+:270270<1.<8<*57=6+.:8/B8*7<1*:2*7;9.,2.;27<1. %%<8 ,58;.<87=6+.:;:.98:<.-/:86<1. # ;99*7-;8=<1.:7 *9*7 ;99&1.;.
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1. INTRODUCTION '6(5;/(90(5:50+(90(5;/6A6('6(5;/(90((9,(/,?(*69(330(569+,96-),5;/0*(504(3: ;/(;6**<9>693+>0+,054(905,,*6:@:;,4::<*/(:*69(39,,-:9<))3,A65,:(5+;/,+,,7:,(<95,;; ,;(3<100(5+#,04,9 $0550.,9,;(3 %/,@(9,()3,;6-694,?;,5:0=,*63650,:(5+ (9,<:,+(:-66+796;,*;06569:/,3;,9)@:,=,9(36;/,9;(?(,5(9;6.(5+63;/<0: ,::05. ,;(3$;(47(9,;(3 9(5*05003/6(5+ 6<9( $70,9(5+,9<4 5(++0;065 4(5@:7,*0,:6-;/,.96<7/(=,(/0./76;,5;0(3-69)06796:7,*;05.%<92,;(3 C4)96:06,; (3 0@(:/0;(,;(3 $6,=,9;/,69+,9C::7,*0,:/(=,(/0:;69@6-5,.3,*;05 +0=,9:0;@:<9=,@:+<,46:;3@;6+0--0*<3;0,:05;/,090+,5;0-0*(;065<95,;;,;(3#@3(5+,;(3 5;/,7(:;;>6+,*(+,:9,:,(9*/<:05.463,*<3(9+(;(/(:/,37,+<5+,9:;(5+05.>/0*/ */(9(*;,9:(9,<:,-<305.,5,90*69:7,*0,:+,3040;(;065(5+0+,5;0-0*(;065(5+05*65:,8<,5*, 04796=,+6<9256>3,+.,65A6(5;/(90(5+0:;90)<;065054(5@36*(;065:>693+>0+, 65;,5,.96,;(3 #0:0(5+ (*65(3+ $(5;6:,;(3 0:,(5+#,04,9 5;/, 5+6"(*0-0*;/,>(;,9::<996<5+05.<:;9(30((9,65,6-;/,9,.065:>0;/;/,3(9.,:; (46<5;6-A6(5;/(90(59,3(;,+30;,9(;<9,,.(++65(5+$/(*23,;65)(93.9,5 <95,;;,;(3 #,04,9,;(3 <100(5+#,04,9 5(++0;0654<*/ 9,:,(9*/65'6(5;/(90(/(:),,57,9-694,+05;/,569;/>,:;,95"(*0-0*7904(903@:6<;/,95 (7(5(5+ %(0>(5,. >6>:2@ (93.9,5 $665.,;(3#,04,9 #,04,9,;(3 ( ) ) +=,5;/6<./:7,*0,:+0=,9:0;@(5++0:;90)<;065:(9,9,3(;0=,3@>,33256>505 ;/,:,9,.065:;/,9,*65;05<,;6),9,769;:6-76;,5;0(33@<5+,:*90),+(5+5,>:7,*0,:6-A6(5;/(90(5: -964;/,:,36*(;065:#,04,9,;(3 *#,04,9(5+<100 !5;/,6;/,9/(5+-69;/,*,5;9(3 5+6"(*0-0* "(5+;/,$6<;//05($,($$ 9,:,(9*/65'6(5;/(90(0:.,5,9(33@:*(9*,>0;/653@9,*,5;(;;,47;:(;)(:0*:7,*0,:30:;::<44(90A,+ -964:*(;;,9,+9,769;:(5+-0,3+04(.,:#,04,9,;(3 ( %/, "(5+$$(9,9,.065: >0;/;/,/0./,:;+0=,9:0;@6-:/(336>>(;,94(905,(504(3:05;/,>693+6,2:,4( %0;;,5:69,; (3 <(5.,;(3 (5+/,5*,(9,=,9@302,3@/(=,/0./,93,=,3:6-'6(5;/(90(+0=,9:0;@;/(5 9,769;,+79,=06<:3@#,04,9,;(3 ( %/,9,-69,9,:,(9*/05;/,:,9,.065:0:,::,5;0(3;6 ),;;,9<5+,9:;(5+;/,90*/5,::(5++0:;90)<;0656-A6(5;/(90(5:65(.36)(3:*(3, 65.:/(;633%(0>(50:;/,3(9.,:;(;63305;/,$$(5+0:36*(;,+05;/,569;/,95769;065 6-;/,$$ ;0:(7796?04(;,3@ 2405+0(4,;,90::769(+0*(33@(--,*;,+)@)6;/05;,95(3>(=,:(5+ 465:665:$/0.<6,;(3 (0 ",B(-369,;(3 &(5.,;(3 "9,=06<::<9=,@: /(=,),,5*65+<*;,+(96<5+65.:/(;633;605=,:;0.(;,;/,+0:;90)<;065(5+,*636.@6-:64, *64465:/(336>>(;,9(504(3::<*/(:*69(3:(5+9,,--0:/,:(0 05,;(3 :<,;(3
2= 17?.>.:<1.:.1*>.+..66726>.;<20*<276;76B7*6<1*:2*6;8.,2.;*<<1.*<744 &1.:./7:.<1.*25;7/<12;;<=-A?.:.<726>.;<20*<.<1.-2>.:;2.6<;-2>.;;67:3.4;*< ;2<.;*:7=6-*6-?2<126<1.*<74420=:. &*+4. :.8:.;.6<260-2//.:.6<5*:26..6>2:765.6<;26,4=-260?*44;:../;478.;,:.;<;4*07768*<,1:../; ,1*66.4;:=++4.B76.;;.*0:*;;B76.;*6-,*>.;&*+4. 2.2 Specimen collection and morphological analyses %8.,25.6,744.,<276?*;8.:/7:5.-=;260;67:3.4260*6-%'*<-.8<1;/:75<1. 26<.:<2-*4<7 5In situ1201:.;74=<27625*0.;?.:.<*3.6<7*6*4AB.57:8174702,*4,1*:*,<.:;.0 7:*4-2;3,747:874A8/7:5;2B.<.6<*,4.6=5+.: -.6<2/2,*<276/7447?.-42<.:*<=:.76B7*6<1*:2*6; /:75<1. 6-7#*,2/2,*6-%%#*@ (.;< =:6.<<.<*4 $.25.: $.25.:.<*4 *
* %8.,25.6;?.:.8:.;.:>.-26 .<1*674?2<1;75. *,:7,6.526* ;8.,25.6;8:.;.:>.-26 ;*4
%8.,2.;42;<.-+.47?*;.@84*26.-26$.25.:.<*4
* =64.;;67<.-7<1.:?2;.(.
,744.,<.- ;8.,25.6;/:75 ;2<.;*:7=6-760;1*<74488.6-2@&*+4. ":-.:)7*6<1*:2*:*A %=+7:-.::*,1A,6.526* *--76*6-%1*,34.<76 * *524A)7*6<12-*.$*/26.;9=. .6=;Zoanthus *5*:,3
Zoanthus sansibaricus *:40:.6 20=:. * %8.,25.6; 66.: *<744 6 !7:<1 ,1*66.4 6 !7:<1.*;< 7=<;2-. 6 %7=<1?.;<7=<;2-.6<7<*4;8.,25.6; !7<.; &12; ;8.,2.; 1*; 8:.>27=;4A +..6 :.87:<.- /:75 6=5.:7=; 47,*<276; 26 <1. %%
86;5, 65/90):633 )3:06;/0 1: =)9 :0- 469: +64465 Zoanthus 97-+1-9 1: =)9 56: .6;5, 6.:-5 )5, 653? 15 94)33 +63651-9 +4 15 4)>14;4 ,1)4-:-8 '019 97-+1-9 )77-)89 :6 *- :63-8)5: 6. ) =1,- <)81-:? 6. -5<18654-5:9 -/ %-14-8 -: )3 * 7-80)79 ,;- :6 .3->1*131:? 15 )996+1):1659 =1:0 1:9 -5,69?4*16:1+ Symbiodinium 977 %-14-8-:)3 * )4-@)21-:)3 5:0199:;,?:019=)9:0-653?@6)5:0)81)5 97-+1-9 6*9-8<-, =1:015 :0- 155-8 3)/665 )=)? .864 :0- +6)9: 6. $8):)9 65/90) 93)5, :=)9.6;5,6<-8)=1,-8)5/-6.,-7:09
4)978-<16;93?9--515#215)=) )7)5 )4-@)21-:)3
Zoanthus gigantus %-14-8 '9;2)0)8) 15 %-14-8 '9;2)0)8) =)4) ')21901:) !)8;?)4) 1/;8- * &7-+14-59"68:0-)9:6;:91,-5 :6:)397-+14-59 "6:-9 Zoanthus gigantus 0)9 653? *--5 78-<16;93? 8-768:-, .864 65- 6:0-8 36+):165 15 :0-&& 65/ 65/%-14-8-:)3 '0-:=6+63651-9.6;5,15:0199:;,?=-8-<-8? 94)33+65919:15/6. 763?79)5,653?):65-91:-65:0-6;:91,-6.:0-):633): 4 ,-7:09)5,:0;9=-->7-+::0)::01997-+1-9198)8-15:0-=):-896.65/90):633)5,15 :0-&& Zoanthus kuroshio %-14-8#5615%-14-8 =)4)')21901:)'9;2)0)8)!)8;?)4)
1/;8- + &7-+14-59)9:6;:91,-5 :6:)397-+14-59 "6:-9'01997-+1-9=)9653?.6;5,65+-15:0-90)336==):-8965:0-6;:91,-6.:0--)9: +6)9:):633): 4)5,+)5*-+6591,-8-,8)8-)86;5,65/90):633 5:0-&&1:0)9 653?78-<16;93? *--5 8-768:-, .864 60')6'0)13)5,)5,96;:0-85')1=)5%-14-8-: )3 '01997-+1-9197814)813?.6;5,1590)336=9;*:1,)3=):-89656;:-8+68)38--.9 8-1-:)3 )413?&70-5671,)- -8:=1/ -5;9Palythoa )46;86;>
Palythoa97sakurajimensis 9-59;%-14-8-:)3 * 1/;8- , &7-+14-59 )9: 6;:91,- 5 "68:0 +0)55-3 5 "68:0-)9: 6;:91,- 5 &6;:0=-9:6;:91,-5 (8-+265=-9:91,-5 :6:)397-+14-59 "6:-9'01997-+1-9=)9.6;5,653?65:0-6;:-8):633;9;)33?):,-7:096.
4-1:0-8
85209(7= 57 04 ;,7= 83(22 *52540,8 5- 652=68 0789 /=659/,80>,+ (8 (4 :4+,8*70),+ 86,*0,804$,03,7,9(2 )-753352,*:2(7<5710485:9/,74 (6(459/,7<571/(8 04+0*(9,+ (++09054(2 86,*03,48 04 9/, ".(8(<(7( 82(4+8 $,03,7 ,9 (2 ( (4+ &(0<(4$,03,7,9(2 )&/,8,86,*03,48-75354.8/(95227,67,8,499/,8,*54+ 8,95-7,*57+8-7539/,%%(-9,7(7,*57+-753 ,4904.85:9/,74&(0<(4$,03,7,9(2
Palythoa mutuki (++54%/(*12,954 ) 0.:7, , %6,*03,4854.8/( 85:9/),(*/4 (895:980+,4 !579/5-(076579 4 !579/,(895:980+,4 %5:9/,(895:980+, 4 %5:9/,(895:980+, 4 '7,*154 <,8980+,4 959(286,*03,48 !59,8&/0886,*0,8<(8():4+(49(4+-5:4+)59/049/,044,72(.554(4+549/,5:980+, 5- 9/, (9522 (9 ( <0+, ;(70,9= 5- +,69/8 3 $,*,49 <571 -753 "104(<( (6(4 04+0*(9,89/(99/0886,*0,83(=(*9:(22=*5480895-9<5579/7,,*258,2=7,2(9,+86,*0,8 0>:=(3((4+ $,03,7:46:)208/,++(9():9-5745<<,/(;,04*2:+,+(227,*57+8 <09/049/0886,*0,8Palythoa mutuki/(867,;05:82=),,47,6579,+-7538,;,7(225*(90548 049/,%%$,03,7,9(2 Palythoa*-toxica '(28/5<,78 0.:7, - %6,*03,48(895:980+,4 !579/5:980+, 4 !579/,(895:980+,4 %5:9/,(89 5:980+, 4 %5:9/,(895:980+, 4 %5:9/<,895:980+,4 959(286,*03,48 !59,8%0302(795*5486,*0-0*86,*03,485)8,7;,+04#(2(:$,03,7,9(2 ((4+(285 (959/,725*(90548049/,%%$,03,7,9(2 9/0886,*0,8<(835+,7(9,2=*53354 (75:4+54.8/(952204*5497(8995-:79/,74579/049/,4579/<,89#(*0-0*049/,<(9,785- &(0<(4(4+"104(<(
( 9/,0+,4909=5-9/0886,*0,87,3(04895),+,-04090;,2=(8*,79(04,+ Palythoa tuberculosa 86,7 0.:7, . %6,*03,48(895:980+,4!579/*/(44,2 4 !579/5-(076579 4 !579/ 5:980+, 4 %5:9/,(89 5:980+, 4 %5:9/,(89 5:980+, 4 %5:9/,(89 5:980+, 4 %5:9/<,89 5:980+, 4 '7,*1 54 <,89 80+, 4 959(2 86,*03,48
!6:,9 &/09 =(9 )? -(8 :/, 469: *64465 97,*0,9 05 :/, =(:,89 (86;5+ 65.9/(:633 -6;5+ (: (33 90:,9 (5+ (*8699 ( =0+, <(80,:? 6- +,7:/9 05:,8:0+(3 4 (5+ 0: /(9 ),,5 8,768:,+:6),=0+,3?+09:80);:,+05:/,%%$,04,8,:(3 5964,36*(:065965 :/,96;:/,(9:(5+96;:/=,9:6-:/,6;:,8(:633:/09=(965,6-:/,469:+6405(5:),5:/0* 97,*0,97(8:0*;3(83?05+,7:/96- 4(3:/6;./0:=(9();5+(5::6+,7:/9+6=5:6
4 $,9,(8*/ /(9 9/6=5 :/(: :/09 97,*0,9 09 -3,>0)3, =0:/ 0:9 ,5+69?4)06:0* Symbiodinium 977(*86990:9.,6.8(7/0*+09:80);:065;85,:: $,04,8(5+&6++ $,04,8,: (3 ( (3:/6;./ 0: 8,4(059 :6 ), 9,,5 0- *63650,9 (: +0--,8,5: +,7:/9 9/6= (5? Symbiodinium<(80(:065=0:/0565,36*(:065 Palythoa heliodiscus$?3(5+ (5*(9:,8 0.;8, / %7,*04,59%6;:/=,9:6;:90+,5 :6:(397,*04,59 !6:,9 &/09 97,*0,9 =(9 8(8, 05 :/, =(:,89 (86;5+ 65.9/( :633 90403(8 :6 78,<06;9 6)9,8<(:065905#(3(;$,04,8,:(3 ();:;5302,"205(=( (7(5=/,8,0:096-:,5 :/,469:*64465Palythoa97,*0,96)9,8<,+(:+,7:/9 4 8,0,:(3 %;)68+,8 (*86*5,405( (++65%/(*23,:65 ( (403? 0*86@6(5:/0+(,;100$,04,8 ,5;9Microzoanthus;100$,04,8
Microzoanthus97 0.;8, ( %7,*04,59!68://(55,3 5 '8,*265=,9:90+,5 :6:(397,*04,59 !6:,9 %7,*04,59 6- (5 ;50+,5:0-0,+ Microzoanthus 97 =,8, -6;5+ (::(*/,+ :6 :/, ;5+,890+,96-*68(38;))3,86*29(:90:,9=0:/9(5+?9;)9:8(:,);::/,97,*0,9*6;3+56:), 0+,5:0-0,+(9763?79=,8,*369,+3:/6;./:/,9,8,78,9,5::/,-089:8,*68+9-68:/,.,5;9 -864:/,%%0:/(9),,58,768:,+=0+,3?(*8699:/, 5+6#(*0-0*;100(5+$,04,8 $,04,8,:(3 )(5+:/;9-05+05.Microzoanthus(:65.9/(:6330956:9;8780905. (403? ?+86@6(5:/0+(,%0550.,8$,04,8#(=36=920 ,5;9Hydrozoanthus%0550.,8$,04,8#(=36=920
Hydrozoanthus(--gracilis =6=92? sensu(83.8,5 0.;8, ) %7,*04,59%6;:/,(9:6;:90+, 5 :6:(397,*04,59 !6:,9 &/,9, 97,*04,59 (.8,, =,33 =0:/ :/, +,9*807:065 6- H. gracilis 9,59; (83.8,5
+225< 8)'6:8 57'2 */81 '4* 9+49')2+8 !52+):2'7 *'9' 9.'9 /4*/)'9+ 9./8 86+)/+8 /8 )258+2> 7+2'9+* 95 (:9 */,,+7+49 ,753 9.+ 86+)/+8 57/-/4'22> *+8)7/(+* (> <5<81> ,753 %'-'3/ '> '6'4 4 54-8.' 9522 ,5:4* +=)2:8/;+2> 54 5:98/*+ <'228 /4 '7+'8 5, 89754- ):77+49 54 Plumeria habereri %9+).5< '8 /8 H. gracilis(+25</7897+)57*/49.+%% Hydrozoanthus gracilis <5<81> sensu/'3/225+9'2 /-:7+ ) %6+)/3+48'895:98/*+4 "579.+'895:98/*+4 959'286+)/3+48 "59+8 %6+)/3+48 5, 9./8 86+)/+8 )522+)9+* ,753 54-8.' 9522 '-7++ <+22 /'3/225+9'2 '4*$+/3+7+9'2 '(+/4- +6/(/59/) 54 9.+ .>*75?5'4 Plumeria habereri '4* .';/4- '4 +=9+74'2 8)'6:8 9.'9 /8 (75<4/8. /4 )5257 54 5:98/*+ <'228 /4 '7+'8 5, 89754- ):77+49 /789 7+)57* ,7539.+%% &./886+)/+8*/,,+78,7539.+59.+7*+8)7/(+*3576.59>6+5,9.+86+)/+8<./)..'8>+225< +=9+74'2 )5257'9/54 57'2 */81 '4* 9+49')2+8 '4* */,,+78 82/-.92> /4 352+):2'7 89:*/+8 %/44/-+7+9'2 .+7+'8H',,gracilis'4*,579./87+'854<+.';+2/89+*9.+8+'8 9<5*/,,+7+4986+)/+8/49./889:*>. '3/2>#'7'?5'49./*'++2'-+ A75:'7* +4:8Antipathozoanthus%/44/-+7$+/3+7#'<25<81/
Antipathozoanthus86 /-:7+ * %6+)/3+48'895:98/*+4 959'286+)/3+48 "59+8 &.+ 542> *+8)7/(+* 86+)/+8 5, 9./8 -+4:8 /4 9.+ #')/,/) /8 A. hickmani ,753 9.+ '2'6'-58 $+/3+7 '4* :0// '29.5:-. 7+)57*8 5, :4*+8)7/(+* 86+)/3+48 +=/89 ,753 '6'4$+/3+7+9'2 * 454-8.'9522542>,5:4*'9'895:98/*+(:925)'22> )53354 '9 *+69.8 5, @ 3 +6/(/59/) 54 '4 '49/6'9.'7/'4 /789 7+)57* ,57 ,'3/2> #'7'?5'49./*'+'4*-+4:8Antipathozoanthus/49.+%% '3/2>6/?5'49./*'++2'-+ A75:'7* +4:8Epizoanthus 7'>
Epizoanthus',,illoricatus &/8).(/+7+1 /-:7+ +
'7-+14-5: ):;6<;:1,-5 '6<;0-):;6<;:1,- 5 ;6;)3:7-+14-5: #6;-:(0-:-:7-+14-5:>-9-.6<5,);;0--5;9)5+-:6.+)=-:,-7;0 4)96<5,;0- 6<;:1,- 6. ;0- );633 5 ;01: :;<,? ;01: /96<715/ +65;)15: ;>6 469706;?7-: 65- >1;0 9-3);1=-3? 3)9/- 763?7: 44 15 ,1)4-;-9 +647)9-, ;6 6;0-9 ,-:+91*-, Epizoanthus :7-+1-: );;)+0-, ;6 -<51+1, >694 ;<*-: <51+1,)- Eunice )5, ;0- 6;0-9 >1;0 :069; ;-5;)+3-: 3-5/;0 69)3 ,1:2 ,1)4-;-9 )5, :4)33-9 69)5/- 69)3 ,1:2: 44 15 ,1)4-;-9 6;0 46970636/1-: )9- ,1..-9-5; .964 ,-:+91*-, :7-+1-: (6 <5,-9:;)5, 9-3);165:017: *-;>--5 ;01: /96<715/ )5, ,-:+91*-, :7-+1-: ,-;)13-, 463-+<3)9 )5, 46970636/1+)3)5)3?:-:)9-5--,-,
Epizoanthus beriber 1:-&-14-9 1/<9- . '7-+14-5: ):;6<;:1,-5;6;)3:7-+14-5: #6;-:(01::7-+1-:>):.6<5,15+)=-:);,-7;0:*-36> 41565/:0);633)5,653? );65-36+);16565;0-6<;:1,-6.;0-);633 ;1:-):?;61,-5;1.?;01::7-+1-:*?1;:>01;- +63651)3 763?7: >1;0 ) 46,-9);-3? ,-=-367-, +6-5-5+0?4- 3;06 <15-)15;0-%)+1.1+$+-)5 1:-)5,&-14-9 ;01:9-79-:-5;:;0-.19:;9-+69,.964;0-''
Epizoanthus illoricatus (1:+0*1-9-2 1/<9- / '7-+14-5: ):;6<;:1,-5'6<;0-):;6<;:1,- 5 '6<;0-):;6<;:1,- 5;6;)3 :7-+14-5: #6;-: (01: :7-+1-: :1413)9 ;6 6;0-9 Epizoanthus :77 6*:-9=-, ); 65/:0) ;633 1: 6*31/);-65-<51+1,>694;<*-:)5,1:)*<5,)5;15;0- 5,6%)+1.1+&-14-9-;)3 )
) 1:-)5,&-14-9 (01::7-+1-:>):.6<5,>1;015)5,69)96<5,;0--5;9)5+- 6.+)=-:)5,659--..3669:6.;0-6<;-99--.6.65/:0);6333;06): 79-=16<:3? 9-769;-, .964 ;0- '' ): Epizoanthus ).. illoricatus *):-, 65 ;0- ,-=-3674-5; 6. +6-5-5+0?4- )5, 763?7 :1@- 15 &-14-9 -; )3 1; 0): *--5 ,-465:;9);-, ;0); ;01: :7-+1-: 0): ) 01/0 3-=-3 6. 15;9):7-+1.1+ 46970636/1+)3 =)91);165 )5,;0-Epizoanthus)..illoricatus469706;?7-15&-14-9-;)3 >)::<*:-8<-5;3? +65.194-, ): E. illoricatus 15
1:- )5, &-14-9 69 ;01: 9-):65 >- ;9-);
Epizoanthus )..illoricatus +633-+;-,.964!)?)5/!)?)5/;633")3)?:1)15&-14-9-; )3 ):E. illoricatus)5,;01::7-+1-:4)?*-,1:;91*<;-,>1,-3?15;0-''
Epizoanthus inazuma 1:-&-14-9 1/<9- 0
$6,*03,49(9:5;:90+,4 $5;:/,(9:5;:90+, 4 :5:(296,*03,49 5:,9%/0996,*0,9=(9-5;4+0490302(8,4<08543,4:9:5E. illoricatus 9;*/(9=0:/0458 (: :/, ,4:8(4*, :5 *(<,9 (4+ 54 :/, 8,,- -2558 (4+ 9256,9 5- :/, 5;:,8 8,,- 5- 54.9/( :522 $6,*03,49 5- :/09 96,*0,9 (8, ;9;(22? +0--0*;2: :5 0+,4:0-? 04 :/, -0,2+ (9 :/,08 ,>:,84(23586/525.?09<,8?90302(8:5E. illoricatus04(20+,4:0-0*(:0545-:/0996,*0,9 8,7;08,9 5)9,8<04. 04:,84(2 3586/525.? (4+ *40+(, (4+ 6/?25.,4,:0* *536(809549 =0:/ E. illoricatus #,9;2:9 5- 68,20304(8? 352,*;2(8 (4(2?9,9 04+0*(:, :/(: 953, 96,*03,49 (::(*/,+:5,;40*0+=583:;),9*522,*:,+-85354.9/(:522=,8,E. inazuma 09,(4+ #,03,8 ;46;)209/,+ +(:( %/,9, 96,*03,49 (8, :/, -089: 8,*58+9 -853 :/, $$ %/09 96,*0,9/(968,<05;92?),,48,*58+,+-853!104(=( (6(4 09,(4+#,03,8 3.2 Zoantharians of Dongsha Atoll !;88,9;2:99/5=,+:/(::/,8,(8,(:2,(9: 96,*0,95-'5(4:/(80((:54.9/(:522 4*2;+,+04:/;94;3),8(8,:/,-089:8,*58+904:/,$$-58-(302??+85@5(4:/0+(,.,4;9 Hydrozoanthus96,*0,9H. gracilis(4+H.(--gracilis-(302?"(8(@5(4:/0+(,.,4;9 Antipathozoanthus;4+,9*80),+Antipathozoanthus96-(302? 0*85@5(4:/0+(,.,4;9 Microzoanthus;40+,4:0-0,+Microzoanthus96(4+-58:/,96,*0,9Epizoanthus illoricatus,E. beriber,(4+E. inazuma++0:054(22?(2296,*0,98,658:,+/,8,8,68,9,4::/,-089:-583(28,*58+9-58 54.9/(:522 0--,8,4*,9),:=,,4:/,044,82(.554"8(:(992(4+(4+5;:,88,,-95-54.9/(:522 %/,044,82(.5545-54.9/(:522093;*/9/(225=,83(>03;3+,6:/04 138(+0;9 -853"8(:(992(4+3+,,6,9:+,6:/04:/,*,4:,85-:/,(:522
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alythoa tuberculosa(354.:/,359:*53354),4:/0*96,*0,9%/,044,8(:522 (8,(/(9,>6,80,4*,+9,<,8,2,:/(2)2,(*/04.04:/,6(9:(0 ';5,:(2 (4+3(4? 9*2,8(*:040(4*58(29+0,+04:/09,<,4:$53,@55>(4:/,22(:,@5(4:/(80(496,*0,9*(445::52,8(:,/0./ :,36,8(:;8,9(4+;2:8(<052,:8(+0(:054 ,99,8,:(2 (4+:/;90:098,(954()2,:5(99;3,:/(: Zoanthus(4+Palythoa96,*0,968,9,4:04:/,044,82(.554(::/09:03,(2959;--,8,+9,<,8, *549,7;,4*,9-853)2,(*/04.90302(8(99,,454:/,358,95;:/=(8+ (?(4. (?(4.:52204:/, $68(:2?92(4+9&(/,,+,:(2 ;8:/,8358,:/,044,82(.55490:,9=/,8,9;8<,?9=,8,
%10&6%5'&8'3'%*#3#%5'3+;'&$:*+)*.'7'.41(4+.5#5+10+0&+%#5+0)#.#%-1(%+3%6.#5+10 #$.' 5+4 -01805*#5Zoanthus#0&Palythoa42223'('3#3'#48+5*'9214'&518#7'4#0&%+3%6.#5+103'+'5#.
*'3'(13'5*'+00'3.#)1101(10)4*#51..&1'4015#22'#351*#7''07+310/'05446+5#$.' (13/#0:4*#..188#5'3;1#05*#3+#042'%+'4 *+4.18'3;1#05*#3+#042'%+'4&+7'34+5:#0&#$60�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ƹ%+#5'&$.'#%*+0)+046//'3/105*4 "1#05*#3+#042'%+'4&+7'34+5:+05*' 1451(5*'3#%*:%0'/+0#42'%+'43'%13&'Ǿ)4*#51..#.411%%63+015*'33')+104 1( #.5*16)*2#45.+5'3#563'*#43'2135'&10.:10'42'%+'41(46$13&'3 #%31%0'/+0#515*+4 3')+10 '+/'3'5#. '+/'3'5#. 46))'45'&5*#55*''9#/+0#5+101(/13'%13#.3''( #3'#4+05*' 816.&.+-'.:(+0&/13'23'7+164.:603'2135'& #%31%0'/+0#42'%+'4#0&5*'3'46.54 1(5*+4456&:%10(+3/'&5*+4*:215*'4+4#48'(160&4'7'3#.#;119#05*'..#5'42'%+'41(5*'46$13&'3 #%31%0'/+0#+05*'8#5'341(10)4*#51..)'0'3#EpizoanthusHydrozoanthus, Antipathozoanthus #0&Microzoanthus48'..41/'/#%31%0'/+0+%;1#05*#3+#04+0*#$+5%#7'4#0& &''2'38#5'34 /')+4'#0& '+/'3 #0&46%**#$+5#54#3'%1//10+05*'165'33''( #3'#1(10)4*#51.. 14523'7+164;1#05*#3+#03'%13&4+05*' 8'3'1$5#+0'&$:42'%+/'04#0&+/#)'4 %1..'%5'&$:01042'%+#.+454 '+/'3'5#. !*'0%1/2#3+0)515*'3'46.541(5*'%633'054637': 515*14'(31/2#454637':48'%#060&'345#0&5*'+/2135#0%'1((+'.&813-(1%64'&1042'%+(+%5#9# +05*+4%#4';1#05*#3+#04/10)5*'42'%+'4(160&+010)4*#51..+05*+4456&:41/'46%*#4 Microzoanthus42*#7'%3:25+%*#$+5#54')6,++#0& '+/'3 8*+.'15*'34#3'&+((+%6.551 +&'05+(:+05*'(+'.&')/#0:Zoanthus#0&Palythoa422630'55'5#. !'231214'5*#55*' 23'4'0%'1(3'4'#3%*'3442'%+(+%#..:(1%64'&10;1#05*#3+#048#410'1(5*'-':3'#41045*#541/#0: 0'83'%13&4(135*' 8'3'(160&+05*+4456&: *'0'842'%+'43'%13&4#510)4*#51..3'2135'&*'3'+0%3'#4'5*';1#05*#3+#0&+7'34+5: +05*' (31/ 42'%+'451
42'%+'41(8*+%*5*3''H#((gracilisAntipathozoanthus42E #((illoricatus#3'.+-'.:60&'4%3+$'& 13'17'38+5*5*'4'0'83'%13&442'%+'406/$'343'2135'&+0 5*' #3'0184+/+.#3515*14'1(5*' /132*142'%+'4 '+/'3'5#. ##.5*16)*$15*
*:.*;*:.;<255=7-.:;<=-2.-*7->.:A524.5A1*:+8:*--2<287*5;9.,2.;8<18/<1.;.:.0287;;<255 ,=::.7<5A1*:+8:58?.:;9.,2.;-2>.:;2.-27 *9*7 ;99$.26.:*7-=322 &1.;.:.;=5<;-.687;<:*<.<1*<68:.?8:42;7..-.-27+8<1:.0287;<8+.<<.:=7-.:;<*7- ;9.,2.;-2>.:;2.;<20*<.-..9.:?*<.:;?12,1524.5A1*:+8:=7478?7)8*7<1*:2*-2>.:;2.:;26=:<1A%1*;1*74 ,*-.62*%272,*/8:<1.2:1.5927/2.5-?8:4*7-5802;<2,;&1.;.728:*=<18:?*;*5;8;=998:<.-+A* %#% 2+*70:*7<.7<2<5.-G58+*5.>85=<2878/:*,1A,7.627**7-<1.2:SymbiodiniumH 5. LITERATURE CITED =:7.<<( 8702<=-27*5>*:2*<28727*50*5;A6+287<;B88@*7<1.55*./:86<1. 7-2*7",.*7 B8*7<12-Palythoa caesia *:27.,8580A#:80:.;;%.:2.; F =:7.<<( .7B2. .*:-68:. $A5*7- % 2010.7.<2,>*:2*+2522;28727 989=5*<287;8/*,587*5,72-*:2*7Zoanthus coppingeri/:86<1.:.*<*::2.:$../ *:27. 28580A
F =:7.<<( .7B2. .*:-68:. $A5*7- %)8*7<12-;7<18B8*.@*,8:*552*/:86 <1.:.*<*::2.:$../*7-&8::.;%<:*2<=;<:*52*;A;<.6*<2,;.>85=<287*7-*4.A<8;9.,2.; 8:*5$../; F *:50:.7"";<*/:24*72;,1.,<272.70.;*66.5<>87.::7:%<=156*77=7- 2<<.25=70.7*=;-.6!*<=:12;<8:2;,1.7 =;.=627*6+=:0%=995.6.7< F 95; F *:50:.7" C+.:.2720.8;<*;2*<2;,1.)8*7<1*:2.7:42>/D:)885802 F *:50:.7" .:2*7<1*:2**7-)8*7<1*:2*%,2.7<2/2,$.98:<;8/<1.:.*<*::2.:$../ @9.-2<287 F
.>94>2;! 0@6;6.>6..;1*<.;@5.>6.3><:%.96. A;49E?6<4>.36?8.%G99?8.=2@?6 A;1J>5.;196;4.> N .6 <;4?5.@<996;@52%2?2;@.;13A@A>2?9.;1<3@52(<>91' ;@2>;.@6<;.9<;32>2;02 6;:2;?9.;1?#A2:<:.;2;0>A?@6;4"5696==6;2Parazoanthus?=[email protected]><; 2@@2>? N 29.42)H>1 *<.;@561H?N*<.;@561.2== N=9? ;&>.6@H12*<<9<462 0<;0>I@2&<:2N :2".>@62 2?<292;@H>H?$26;C.91".>6?D+D6, ===9? N 2;.>@<4 <9@5A6? ;<@2<;.;6;@2>2?@6;4.??<06.@6<;<3@520>./Platypodiella picta 69;21C.>1? .;1?=2062?<3*<.;@5.>6.62><>?05A;4?6;?@6@A@%2;082;/2>4 N 6.:699<< "A02%.B2?@>299< ??<06.@6<;/2@C22;Dentitheca habereri ;61.>6.E1>;.9<3*<<9<4E N ?=2> 62"39.;F2;@562>26;//691A;42;;.0512> .@A>:6@.>/2;2>92A05@2@;2/?@ 2?05>26/A;42;>6@@2>&5269>2EF252;12 6232>A;4$.?=2 K>;/2>4== N =9? >.;06;6695<$ .$ ">21.@6<;<;@52@223 36?52?6;@52/><95;>.F69>.F696.; ;.9<3!02.;<4>.=5E N A766&$26:2> "5E9<42;E<3@5256459E16B2>42;@F<.;@5613.:69E 60>.996.3><:@52".06360*<<9<460.%0>6=@. N A766&$26:2> ;2C3.:69E<316:6;A@6B2F<.996. *<.;@5.>6.*<<9<460.9 ;.9<3@52 6;;2.;%<062@E
>.E +&,52;6:.99?<>*<.;@5.>6..;1@52?@299.@210<>.9?== N ; %E;<=?6?<3@52<;@2;@?<3@52>6@6?5 A?2A:&C2;@E%6D@516@6<;(<<13.99.;1%<; <;1<;+6B, == >.E <@2?<;*<.;@56;.2C6@512?0>6=@6<;?<3?<:2;2C42;2>."><02216;4?<3@52 *<<9<460.9%<062@E<3 <;1<; N .11<;%5.0892@<; .>2B6?6<;<3@52>6@6?50@6;6.2".>@&52*<.;@52.2; $2=<>@?<;@52F<<9<460.90<9920@6<;?:.126;@52&<>>2?%@>.6@?/E.11<; N %062;@6360&>.;?.0@6<;?<3@52$
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
@-:3 5/@-:-:)*;17>19-)41::3#"@-:3 -:1 )(;%&)-4110 +221:05**1195:&4;@ C?=-;=05:-=D05A1=>5?D;2=112/;=-8>5:?41%;@?4 45:-%1- -=5:15;05A1=>5?D =15*!;E-B-*$1591= 5>?=5.@?5;:<-??1=:>;225A1E;-:?450><1/51>5:"75:-B- >8-:0 -<-:+;;8;35/-8%?@051> -91E-75 53- 5=;>1 %@0-%$1591= 5221=1:?E;;C-:?4188-1?D<1>5: <;<@8-?5;:>;2?41E;-:?450Zoanthus sansibaricus-8;:3015:"75:-B- -<-: -=5:15;05A1=>5?D 5>1$1591= ':1C<1/?1005A1=>5?D-:0:1B><1/51>;2Epizoanthus-??-/410?;1@:5/50 B;=9[email protected]>:?4;E;-1C-/;=-885-2=;9?41#-/525/"/1-:+;; 1D> -9-=/7 # +;-:?41+;-:?4-< :%D>?H9101>-:59-@C>-:>A1=?G.=1>;@&-.81-@ 3F:F=-801>/8->>1>01>;=0=1>1?01>31:=1>01/1>-:59-@C#=F>1:?-:?81@=>/-=-/?G=1> 1>>1:?518>1?81@=05>?=5.@?5;:0L-<=G>8-/;:>50F=-?5;:0181@=>=-<<;=?>:-?@=18>1?81@= ;=3-:5>-?5;:1?>@5A-:?8L-==-:3191:?F?-.85>0-:>81>3-81=51>0@ @>F@90L5>?;5=1!-?@=1881 <-=9581@=0F<;@5881>/;:>1=AF1>#=F/F0F0@05>/;@=>0L;@A1=?@=10@/;@=>01E;;8;3510;::F
2/C0:7?C3 QCB3C@3BJB3@D7::3 #/@7AD777 >> /;=C@=CF ( 7AB=7@323A>=:G>73@A1=@/::75K<3A4:3F70:3ADC:5/7@3;36GB3A#=7AA=</3<:FFF7D, -
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%499420=$0480= ",B7:B>64 &30",=,E:,9?34/,0 ?,C:9:8D/0>.=4;?4:9:1?B: 90B2090=, ,=4904:/4A0=>4?D %499420=!.,H,!',.:$ 4A0=>4?D:1E:,9?34/>9?3:E:,0C,.:=,774,:9,B,44,9 >0,8:@9?>/0>.=4;?4:9:1?30,B,44,92:7/.:=,7,9/,//4?4:9,7E:,9?34/>" :%!90 0 %::92 %34,@*309 :=;3:7:24.,7,9/741034>?:=D/4A0=209.0:1?30E:,9?34/ Sphenopus marsupialis:11?30&,4B,90>0:,>?+::7:24.,7%?@/40> L %:@E,% =:>>4/0%, %47A, "=,9.:! :80> @94:= !74A04=,$$:.3,& ,2,73F0>" ,==, =:>>4/0%, $:8,9:/0%, :8-=496 48G90E' -=0@ $ /09?414.,?4:9:1,9:A07-0?, ,.0?D730C:>,8494/,>0".- 1=:8 8,=490E:,9?34/Palythoa caribaeorum94/,=4,9?3:E:,+:,9?34/0,"=:?049C;=0>>4:9 ,9/"@=414.,?4:9 L %;40=0=@8 4=>?=0.:=/:1;=0/,?4:9:9?30E:,9?34/Palythoa caribaeorum 9?3:E:, 94/,=4,,9/?3014>3Scomberomorus brasiliensis %.:8-=4/,0-D?302=009?@=?70Chelonia mydas49%:@?3=,E47 ,=4904:/4A0=>4?D ! >
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Figures and Tables Table 1. ;3<>:.@6<;<;?6@2?6;B2?@64.@21.><3 -<.;@5.>6.?=206:2;?0<9920@21.@2.05?6@2 Figure 1.!.=.;19<0.@6<;<3<;4?5.@<996;@52&092?>2=>2?2;@ 9<0.@6<;?C6@5>2=<>@?<3-<.;@5.>6.3><:=>2B6092?6;6;?2@>2=>2?2;@9<0.@6<;? 6;B2?@64.@216;@56??@A1E!.12C6@52;2>60!.==6;4'<<9*2??292@.9 Figure 2.&A/<>12>>.05E0;2:6;.?=2062?0<;36>:213><:<;4?5.@<99.Zoanthus sansibaricus /Zoanthus gigantus0Zoanthus kuroshio1Palythoa?=sakurajimensis2Palythoa mutuki3 Palythoa03toxica4Palythoa tuberculosa.;15Palythoa heliodiscus&0.92/.>? 0: Figure 3.&A/<>12>!.0><0;2:6;.?=2062?0<;36>:213><:<;4?5.@<99.Microzoanthus?=/ Hydrozoanthus.33gracilis0Hydrozoanthus gracilis1Antipathozoanthus?=2Epizoanthus.33
illoricatusEpizoanthus beriberEpizoanthusilloricatus Epizoanthus inazuma Appendix Table A1.
20°36.552',
North outside 1
20°42.338',
North of airport 1
North outside 2
East outside
North channel 1
North of airport 3
North of airport 2
116°42.129'
side
May 24; scuba
June 4, June 5; scuba
116°55.439' 20°46.897',
May 24, June 1, June 3,
May 23; scuba
May 22; snorkel
May 22; snorkel
May 23, May 27; snorkel
May 21, June 5; SCUBA
May 21; scuba
20°42.114',
116°46.234'
20°42.886',
116°43.281'
20°42.445',
116°43.213'
20°42.472',
116°43.701'
20°42.377',
Wreck on west
116°43.895'
116°43.401'
beach
May 20; snorkel
(2016); method
Longitude (E) 20°41.960',
Date(s) investigated
Latitude (N),
Dongsha I. south
Site
Gentle reef slope (13 – 18)
Wall, cave, reef slope (5 – 40)
Sandy & rubble (10 – 20)
Sea grass meadows (intertidal – 2)
Sea grass meadows (intertidal – 2)
(intertidal – 2)
Sea grass meadows, tetrapods
Shallow reef (2 – 4)
Sandy reef slope
(intertidal – 2)
Sandy, sea grass meadows
investigated in m)
Site information (depths
27
98
1
2
0
4
38
0
1
specimens collected
# of Zoantharia
Table 1. Information on sites investigated around and within Dongsha Atoll, and number of Zoantharia specimens collected at each site.
20°39.025',
Inner atoll 1
116°51.429' 20°35.110', 116°47.471' N20°35.247', E116°47.603' N20°45.995',
1
Southeast outside
2
Southeast outside
3
Northeast outside E116°53.822'
20°35.819',
116°42.408'
20°43.512',
116°48.373'
20°42.277',
Southeast outside
North channel 2
Inner atoll 2
116°41.711'
outside 116°41.711'
20°39.025',
Southwest
116°48.741'
June 4, June 5; scuba
May 31, June 1; snorkel
scuba
May 30, May 31, June 1;
May 30; scuba
May 29; scuba
May 29; scuba
May 25; scuba
May 25; scuba
crest (intertidal – 35)
Wall, cave, gentle reef slope, reef
Shallow reef crest (intertidal – 2)
Reef slope, wall, cave (10 – 30)
Reef slope (13 – 33)
Sandy & rubble (14 – 17)
Silty lagoon, patch reef (2 – 15)
Silty lagoon, patch reef (2 – 12)
Reef slope (14 – 28)
19
35
46
22
35
0
2
46
Dongsha I. south beach
Inner Atoll 1
Inner Atoll 2
East outside
North channel 1
North North of North of North of North North channel airport 1 airport 2 airport 3 outside 1 outside 2 2
Northeast outside
Southeast outside 1
Southeast outside 2
Southeast outside 3
Southwest outside
Wreck on west side
East China Sea CHINA
Taiwan
Hong Kong
South China Sea
Kenting
Hainan
Dongsha Atoll
Luzon
reef crest (intertidal) Pratas Island
PHILIPPINES
lagoon 10 km
S2352-4855(16)30229-8 http://dx.doi.org/10.1016/j.rsma.2017.02.006 RSMA 225
To appear in:
Regional Studies in Marine Science
Received date: 11 October 2016 Please cite this article as: Reimer, J.D., Santos, M.E.A., Kise, H., Neo, M.L., Chen, C.A., Soong, K., Diversity of Zoantharia (Anthozoa: Hexacorallia) at Dongsha Atoll in the South China Sea. Regional Studies in Marine Science (2017), http://dx.doi.org/10.1016/j.rsma.2017.02.006 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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1. INTRODUCTION '6(5;/(90(5:50+(90(5;/6A6('6(5;/(90((9,(/,?(*69(330(569+,96-),5;/0*(504(3: ;/(;6**<9>693+>0+,054(905,,*6:@:;,4::<*/(:*69(39,,-:9<))3,A65,:(5+;/,+,,7:,(<95,;; ,;(3<100(5+#,04,9 $0550.,9,;(3 %/,@(9,()3,;6-694,?;,5:0=,*63650,:(5+ (9,<:,+(:-66+796;,*;06569:/,3;,9)@:,=,9(36;/,9;(?(,5(9;6.(5+63;/<0: ,::05. ,;(3$;(47(9,;(3 9(5*05003/6(5+ 6<9( $70,9(5+,9<4 5(++0;065 4(5@:7,*0,:6-;/,.96<7/(=,(/0./76;,5;0(3-69)06796:7,*;05.%<92,;(3 C4)96:06,; (3 0@(:/0;(,;(3 $6
2= 17?.>.:<1.:.1*>.+..66726>.;<20*<276;76B7*6<1*:2*6;8.,2.;*<<1.*<744 &1.:./7:.<1.*25;7/<12;;<=-A?.:.<726>.;<20*<.<1.-2>.:;2.6<;-2>.;;67:3.4;*< ;2<.;*:7=6-*6-?2<126<1.*<74420=:. &*+4. :.8:.;.6<260-2//.:.6<5*:26..6>2:765.6<;26,4=-260?*44;:../;478.;,:.;<;4*07768*<,1:../; ,1*66.4;:=++4.B76.;;.*0:*;;B76.;*6-,*>.;&*+4. 2.2 Specimen collection and morphological analyses %8.,25.6,744.,<276?*;8.:/7:5.-=;260;67:3.4260*6-%'*<-.8<1;/:75<1. 26<.:<2-*4<7 5In situ1201:.;74=<27625*0.;?.:.<*3.6<7*6*4AB.57:8174702,*4,1*:*,<.:;.0 7:*4-2;3,747:874A8/7:5;2B.<.6<*,4.6=5+.: -.6<2/2,*<276/7447?.-42<.:*<=:.76B7*6<1*:2*6; /:75<1. 6-7#*,2/2,*6-%%#*@ (.;< =:6.<<.<*4 $.25.: $.25.:.<*4 *
* %8.,25.6;?.:.8:.;.:>.-26 .<1*674?2<1;75. *,:7,6.526* ;8.,25.6;8:.;.:>.-26 ;*4
%8.,2.;42;<.-+.47?*;.@84*26.-26$.25.:.<*4
* =64.;;67<.-7<1.:?2;.(.
,744.,<.- ;8.,25.6;/:75 ;2<.;*:7=6-760;1*<74488.6-2@&*+4. ":-.:)7*6<1*:2*:*A %=+7:-.::*,1A,6.526* *--76*6-%1*,34.<76 * *524A)7*6<12-*.$*/26.;9=. .6=;Zoanthus *5*:,3
Zoanthus sansibaricus *:40:.6 20=:. * %8.,25.6; 66.: *<744 6 !7:<1 ,1*66.4 6 !7:<1.*;< 7=<;2-. 6 %7=<1?.;<7=<;2-.6<7<*4;8.,25.6; !7<.; &12; ;8.,2.; 1*; 8:.>27=;4A +..6 :.87:<.- /:75 6=5.:7=; 47,*<276; 26 <1. %%
86;5, 65/90):633 )3:06;/0 1: =)9 :0- 469: +64465 Zoanthus 97-+1-9 1: =)9 56: .6;5, 6.:-5 )5, 653? 15 94)33 +63651-9 +4 15 4)>14;4 ,1)4-:-8 '019 97-+1-9 )77-)89 :6 *- :63-8)5: 6. ) =1,- <)81-:? 6. -5<18654-5:9 -/ %-14-8 -: )3 * 7-80)79 ,;- :6 .3->1*131:? 15 )996+1):1659 =1:0 1:9 -5,69?4*16:1+ Symbiodinium 977 %-14-8-:)3 * )4-@)21-:)3 5:0199:;,?:019=)9:0-653?@6)5:0)81)5 97-+1-9 6*9-8<-, =1:015 :0- 155-8 3)/665 )=)? .864 :0- +6)9: 6. $8):)9 65/90) 93)5, :=)9.6;5,6<-8)=1,-8)5/-6.,-7:09
4)978-<16;93?9--515#215)=) )7)5 )4-@)21-:)3
Zoanthus gigantus %-14-8 '9;2)0)8) 15 %-14-8 '9;2)0)8) =)4) ')21901:) !)8;?)4) 1/;8- * &7-+14-59"68:0-)9:6;:91,-5 :6:)397-+14-59 "6:-9 Zoanthus gigantus 0)9 653? *--5 78-<16;93? 8-768:-, .864 65- 6:0-8 36+):165 15 :0-&& 65/ 65/%-14-8-:)3 '0-:=6+63651-9.6;5,15:0199:;,?=-8-<-8? 94)33+65919:15/6. 763?79)5,653?):65-91:-65:0-6;:91,-6.:0-):633): 4 ,-7:09)5,:0;9=-->7-+::0)::01997-+1-9198)8-15:0-=):-896.65/90):633)5,15 :0-&& Zoanthus kuroshio %-14-8#5615%-14-8 =)4)')21901:)'9;2)0)8)!)8;?)4)
1/;8- + &7-+14-59)9:6;:91,-5 :6:)397-+14-59 "6:-9'01997-+1-9=)9653?.6;5,65+-15:0-90)336==):-8965:0-6;:91,-6.:0--)9: +6)9:):633): 4)5,+)5*-+6591,-8-,8)8-)86;5,65/90):633 5:0-&&1:0)9 653?78-<16;93? *--5 8-768:-, .864 60')6'0)13)5,)5,96;:0-85')1=)5%-14-8-: )3 '01997-+1-9197814)813?.6;5,1590)336=9;*:1,)3=):-89656;:-8+68)38--.9 8-1-:)3 )413?&70-5671,)- -8:=1/ -5;9Palythoa )46;86;>
Palythoa97sakurajimensis 9-59;%-14-8-:)3 * 1/;8- , &7-+14-59 )9: 6;:91,- 5 "68:0 +0)55-3 5 "68:0-)9: 6;:91,- 5 &6;:0=-9:6;:91,-5 (8-+265=-9:91,-5 :6:)397-+14-59 "6:-9'01997-+1-9=)9.6;5,653?65:0-6;:-8):633;9;)33?):,-7:096.
4-1:0-8
85209(7= 57 04 ;,7= 83(22 *52540,8 5- 652=68 0789 /=659/,80>,+ (8 (4 :4+,8*70),+ 86,*0,804$,03,7,9(2 )-753352,*:2(7<5710485:9/,74 (6(459/,7<571/(8 04+0*(9,+ (++09054(2 86,*03,48 04 9/, ".(8(<(7( 82(4+8 $,03,7 ,9 (2 ( (4+ &(0<(4$,03,7,9(2 )&/,8,86,*03,48-75354.8/(95227,67,8,499/,8,*54+ 8,95-7,*57+8-7539/,%%(-9,7(7,*57+-753 ,4904.85:9/,74&(0<(4$,03,7,9(2
Palythoa mutuki (++54%/(*12,954 ) 0.:7, , %6,*03,4854.8/( 85:9/),(*/4 (895:980+,4 !579/5-(076579 4 !579/,(895:980+,4 %5:9/,(895:980+, 4 %5:9/,(895:980+, 4 '7,*154 <,8980+,4 959(286,*03,48 !59,8&/0886,*0,8<(8():4+(49(4+-5:4+)59/049/,044,72(.554(4+549/,5:980+, 5- 9/, (9522 (9 ( <0+, ;(70,9= 5- +,69/8 3 $,*,49 <571 -753 "104(<( (6(4 04+0*(9,89/(99/0886,*0,83(=(*9:(22=*5480895-9<5579/7,,*258,2=7,2(9,+86,*0,8 0>:=(3((4+ $,03,7:46:)208/,++(9():9-5745<<,/(;,04*2:+,+(227,*57+8 <09/049/0886,*0,8Palythoa mutuki/(867,;05:82=),,47,6579,+-7538,;,7(225*(90548 049/,%%$,03,7,9(2 Palythoa*-toxica '(28/5<,78 0.:7, - %6,*03,48(895:980+,4 !579/5:980+, 4 !579/,(895:980+,4 %5:9/,(89 5:980+, 4 %5:9/,(895:980+, 4 %5:9/<,895:980+,4 959(286,*03,48 !59,8%0302(795*5486,*0-0*86,*03,485)8,7;,+04#(2(:$,03,7,9(2 ((4+(285 (959/,725*(90548049/,%%$,03,7,9(2 9/0886,*0,8<(835+,7(9,2=*53354 (75:4+54.8/(952204*5497(8995-:79/,74579/049/,4579/<,89#(*0-0*049/,<(9,785- &(0<(4(4+"104(<(
( 9/,0+,4909=5-9/0886,*0,87,3(04895),+,-04090;,2=(8*,79(04,+ Palythoa tuberculosa 86,7 0.:7, . %6,*03,48(895:980+,4!579/*/(44,2 4 !579/5-(076579 4 !579/ 5:980+, 4 %5:9/,(89 5:980+, 4 %5:9/,(89 5:980+, 4 %5:9/,(89 5:980+, 4 %5:9/<,89 5:980+, 4 '7,*1 54 <,89 80+, 4 959(2 86,*03,48
!6:,9 &/09 =(9 )? -(8 :/, 469: *64465 97,*0,9 05 :/, =(:,89 (86;5+ 65.9/(:633 -6;5+ (: (33 90:,9 (5+ (*8699 ( =0+, <(80,:? 6- +,7:/9 05:,8:0+(3 4 (5+ 0: /(9 ),,5 8,768:,+:6),=0+,3?+09:80);:,+05:/,%%$,04,8,:(3 5964,36*(:065965 :/,96;:/,(9:(5+96;:/=,9:6-:/,6;:,8(:633:/09=(965,6-:/,469:+6405(5:),5:/0* 97,*0,97(8:0*;3(83?05+,7:/96- 4(3:/6;./0:=(9();5+(5::6+,7:/9+6=5:6
4 $,9,(8*/ /(9 9/6=5 :/(: :/09 97,*0,9 09 -3,>0)3, =0:/ 0:9 ,5+69?4)06:0* Symbiodinium 977(*86990:9.,6.8(7/0*+09:80);:065;85,:: $,04,8(5+&6++ $,04,8,: (3 ( (3:/6;./ 0: 8,4(059 :6 ), 9,,5 0- *63650,9 (: +0--,8,5: +,7:/9 9/6= (5? Symbiodinium<(80(:065=0:/0565,36*(:065 Palythoa heliodiscus$?3(5+ (5*(9:,8 0.;8, / %7,*04,59%6;:/=,9:6;:90+,5 :6:(397,*04,59 !6:,9 &/09 97,*0,9 =(9 8(8, 05 :/, =(:,89 (86;5+ 65.9/( :633 90403(8 :6 78,<06;9 6)9,8<(:065905#(3(;$,04,8,:(3 ();:;5302,"205(=( (7(5=/,8,0:096-:,5 :/,469:*64465Palythoa97,*0,96)9,8<,+(:+,7:/9 4 8,0,:(3 %;)68+,8 (*86*5,405( (++65%/(*23,:65 ( (403? 0*86@6(5:/0+(,;100$,04,8 ,5;9Microzoanthus;100$,04,8
Microzoanthus97 0.;8, ( %7,*04,59!68://(55,3 5 '8,*265=,9:90+,5 :6:(397,*04,59 !6:,9 %7,*04,59 6- (5 ;50+,5:0-0,+ Microzoanthus 97 =,8, -6;5+ (::(*/,+ :6 :/, ;5+,890+,96-*68(38;))3,86*29(:90:,9=0:/9(5+?9;)9:8(:,);::/,97,*0,9*6;3+56:), 0+,5:0-0,+(9763?79=,8,*369,+3:/6;./:/,9,8,78,9,5::/,-089:8,*68+9-68:/,.,5;9 -864:/,%%0:/(9),,58,768:,+=0+,3?(*8699:/, 5+6#(*0-0*;100(5+$,04,8 $,04,8,:(3 )(5+:/;9-05+05.Microzoanthus(:65.9/(:6330956:9;8780905. (403? ?+86@6(5:/0+(,%0550.,8$,04,8#(=36=920 ,5;9Hydrozoanthus%0550.,8$,04,8#(=36=920
Hydrozoanthus(--gracilis =6=92? sensu(83.8,5 0.;8, ) %7,*04,59%6;:/,(9:6;:90+, 5 :6:(397,*04,59 !6:,9 &/,9, 97,*04,59 (.8,, =,33 =0:/ :/, +,9*807:065 6- H. gracilis 9,59; (83.8,5
+225< 8)'6:8 57'2 */81 '4* 9+49')2+8 !52+):2'7 *'9' 9.'9 /4*/)'9+ 9./8 86+)/+8 /8 )258+2> 7+2'9+* 95 (:9 */,,+7+49 ,753 9.+ 86+)/+8 57/-/4'22> *+8)7/(+* (> <5<81> ,753 %'-'3/ '> '6'4 4 54-8.' 9522 ,5:4* +=)2:8/;+2> 54 5:98/*+ <'228 /4 '7+'8 5, 89754- ):77+49 54 Plumeria habereri %9+).5< '8 /8 H. gracilis(+25</7897+)57*/49.+%% Hydrozoanthus gracilis <5<81> sensu/'3/225+9'2 /-:7+ ) %6+)/3+48'895:98/*+4 "579.+'895:98/*+4 959'286+)/3+48 "59+8 %6+)/3+48 5, 9./8 86+)/+8 )522+)9+* ,753 54-8.' 9522 '-7++ <+22 /'3/225+9'2 '4*$+/3+7+9'2 '(+/4- +6/(/59/) 54 9.+ .>*75?5'4 Plumeria habereri '4* .';/4- '4 +=9+74'2 8)'6:8 9.'9 /8 (75<4/8. /4 )5257 54 5:98/*+ <'228 /4 '7+'8 5, 89754- ):77+49 /789 7+)57* ,7539.+%% &./886+)/+8*/,,+78,7539.+59.+7*+8)7/(+*3576.59>6+5,9.+86+)/+8<./)..'8>+225< +=9+74'2 )5257'9/54 57'2 */81 '4* 9+49')2+8 '4* */,,+78 82/-.92> /4 352+):2'7 89:*/+8 %/44/-+7+9'2 .+7+'8H',,gracilis'4*,579./87+'854<+.';+2/89+*9.+8+'8 9<5*/,,+7+4986+)/+8/49./889:*>. '3/2>#'7'?5'49./*'++2'-+ A75:'7* +4:8Antipathozoanthus%/44/-+7$+/3+7#'<25<81/
Antipathozoanthus86 /-:7+ * %6+)/3+48'895:98/*+4 959'286+)/3+48 "59+8 &.+ 542> *+8)7/(+* 86+)/+8 5, 9./8 -+4:8 /4 9.+ #')/,/) /8 A. hickmani ,753 9.+ '2'6'-58 $+/3+7 '4* :0// '29.5:-. 7+)57*8 5, :4*+8)7/(+* 86+)/3+48 +=/89 ,753 '6'4$+/3+7+9'2 * 454-8.'9522542>,5:4*'9'895:98/*+(:925)'22> )53354 '9 *+69.8 5, @ 3 +6/(/59/) 54 '4 '49/6'9.'7/'4 /789 7+)57* ,57 ,'3/2> #'7'?5'49./*'+'4*-+4:8Antipathozoanthus/49.+%% '3/2>6/?5'49./*'++2'-+ A75:'7* +4:8Epizoanthus 7'>
Epizoanthus',,illoricatus &/8).(/+7+1 /-:7+ +
'7-+14-5: ):;6<;:1,-5 '6<;0-):;6<;:1,- 5 ;6;)3:7-+14-5: #6;-:(0-:-:7-+14-5:>-9-.6<5,);;0--5;9)5+-:6.+)=-:,-7;0 4)96<5,;0- 6<;:1,- 6. ;0- );633 5 ;01: :;<,? ;01: /96<715/ +65;)15: ;>6 469706;?7-: 65- >1;0 9-3);1=-3? 3)9/- 763?7: 44 15 ,1)4-;-9 +647)9-, ;6 6;0-9 ,-:+91*-, Epizoanthus :7-+1-: );;)+0-, ;6 -<51+1, >694 ;<*-: <51+1,)- Eunice )5, ;0- 6;0-9 >1;0 :069; ;-5;)+3-: 3-5/;0 69)3 ,1:2 ,1)4-;-9 )5, :4)33-9 69)5/- 69)3 ,1:2: 44 15 ,1)4-;-9 6;0 46970636/1-: )9- ,1..-9-5; .964 ,-:+91*-, :7-+1-: (6 <5,-9:;)5, 9-3);165:017: *-;>--5 ;01: /96<715/ )5, ,-:+91*-, :7-+1-: ,-;)13-, 463-+<3)9 )5, 46970636/1+)3)5)3?:-:)9-5--,-,
Epizoanthus beriber 1:-&-14-9 1/<9- . '7-+14-5: ):;6<;:1,-5;6;)3:7-+14-5: #6;-:(01::7-+1-:>):.6<5,15+)=-:);,-7;0:*-36> 41565/:0);633)5,653? );65-36+);16565;0-6<;:1,-6.;0-);633 ;1:-):?;61,-5;1.?;01::7-+1-:*?1;:>01;- +63651)3 763?7: >1;0 ) 46,-9);-3? ,-=-367-, +6-5-5+0?4- 3;06 <15-)15;0-%)+1.1+$+-)5 1:-)5,&-14-9 ;01:9-79-:-5;:;0-.19:;9-+69,.964;0-''
Epizoanthus illoricatus (1:+0*1-9-2 1/<9- / '7-+14-5: ):;6<;:1,-5'6<;0-):;6<;:1,- 5 '6<;0-):;6<;:1,- 5;6;)3 :7-+14-5: #6;-: (01: :7-+1-: :1413)9 ;6 6;0-9 Epizoanthus :77 6*:-9=-, ); 65/:0) ;633 1: 6*31/);-65-<51+1,>694;<*-:)5,1:)*<5,)5;15;0- 5,6%)+1.1+&-14-9-;)3 )
) 1:-)5,&-14-9 (01::7-+1-:>):.6<5,>1;015)5,69)96<5,;0--5;9)5+- 6.+)=-:)5,659--..3669:6.;0-6<;-99--.6.65/:0);6333;06): 79-=16<:3? 9-769;-, .964 ;0- '' ): Epizoanthus ).. illoricatus *):-, 65 ;0- ,-=-3674-5; 6. +6-5-5+0?4- )5, 763?7 :1@- 15 &-14-9 -; )3 1; 0): *--5 ,-465:;9);-, ;0); ;01: :7-+1-: 0): ) 01/0 3-=-3 6. 15;9):7-+1.1+ 46970636/1+)3 =)91);165 )5,;0-Epizoanthus)..illoricatus469706;?7-15&-14-9-;)3 >)::<*:-8<-5;3? +65.194-, ): E. illoricatus 15
1:- )5, &-14-9 69 ;01: 9-):65 >- ;9-);
Epizoanthus )..illoricatus +633-+;-,.964!)?)5/!)?)5/;633")3)?:1)15&-14-9-; )3 ):E. illoricatus)5,;01::7-+1-:4)?*-,1:;91*<;-,>1,-3?15;0-''
Epizoanthus inazuma 1:-&-14-9 1/<9- 0
$6,*03,49(9:5;:90+,4 $5;:/,(9:5;:90+, 4 :5:(296,*03,49 5:,9%/0996,*0,9=(9-5;4+0490302(8,4<08543,4:9:5E. illoricatus 9;*/(9=0:/0458 (: :/, ,4:8(4*, :5 *(<,9 (4+ 54 :/, 8,,- -2558 (4+ 9256,9 5- :/, 5;:,8 8,,- 5- 54.9/( :522 $6,*03,49 5- :/09 96,*0,9 (8, ;9;(22? +0--0*;2: :5 0+,4:0-? 04 :/, -0,2+ (9 :/,08 ,>:,84(23586/525.?09<,8?90302(8:5E. illoricatus04(20+,4:0-0*(:0545-:/0996,*0,9 8,7;08,9 5)9,8<04. 04:,84(2 3586/525.? (4+ *40+(, (4+ 6/?25.,4,:0* *536(809549 =0:/ E. illoricatus #,9;2:9 5- 68,20304(8? 352,*;2(8 (4(2?9,9 04+0*(:, :/(: 953, 96,*03,49 (::(*/,+:5,;40*0+=583:;),9*522,*:,+-85354.9/(:522=,8,E. inazuma 09,(4+ #,03,8 ;46;)209/,+ +(:( %/,9, 96,*03,49 (8, :/, -089: 8,*58+9 -853 :/, $$ %/09 96,*0,9/(968,<05;92?),,48,*58+,+-853!104(=( (6(4 09,(4+#,03,8 3.2 Zoantharians of Dongsha Atoll !;88,9;2:99/5=,+:/(::/,8,(8,(:2,(9: 96,*0,95-'5(4:/(80((:54.9/(:522 4*2;+,+04:/;94;3),8(8,:/,-089:8,*58+904:/,$$-58-(302??+85@5(4:/0+(,.,4;9 Hydrozoanthus96,*0,9H. gracilis(4+H.(--gracilis-(302?"(8(@5(4:/0+(,.,4;9 Antipathozoanthus;4+,9*80),+Antipathozoanthus96-(302? 0*85@5(4:/0+(,.,4;9 Microzoanthus;40+,4:0-0,+Microzoanthus96(4+-58:/,96,*0,9Epizoanthus illoricatus,E. beriber,(4+E. inazuma++0:054(22?(2296,*0,98,658:,+/,8,8,68,9,4::/,-089:-583(28,*58+9-58 54.9/(:522 0--,8,4*,9),:=,,4:/,044,82(.554"8(:(992(4+(4+5;:,88,,-95-54.9/(:522 %/,044,82(.5545-54.9/(:522093;*/9/(225=,83(>03;3+,6:/04 138(+0;9 -853"8(:(992(4+3+,,6,9:+,6:/04:/,*,4:,85-:/,(:522
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alythoa tuberculosa(354.:/,359:*53354),4:/0*96,*0,9%/,044,8(:522 (8,(/(9,>6,80,4*,+9,<,8,2,:/(2)2,(*/04.04:/,6(9:(0 ';5,:(2 (4+3(4? 9*2,8(*:040(4*58(29+0,+04:/09,<,4:$53,@55>(4:/,22(:,@5(4:/(80(496,*0,9*(445::52,8(:,/0./ :,36,8(:;8,9(4+;2:8(<052,:8(+0(:054 ,99,8,:(2 (4+:/;90:098,(954()2,:5(99;3,:/(: Zoanthus(4+Palythoa96,*0,968,9,4:04:/,044,82(.554(::/09:03,(2959;--,8,+9,<,8, *549,7;,4*,9-853)2,(*/04.90302(8(99,,454:/,358,95;:/=(8+ (?(4. (?(4.:52204:/, $68(:2?92(4+9&(/,,+,:(2 ;8:/,8358,:/,044,82(.55490:,9=/,8,9;8<,?9=,8,
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*:.*;*:.;<255=7-.:;<=-2.-*7->.:A524.5A1*:+8:*--2<287*5;9.,2.;8<18/<1.;.:.0287;;<255 ,=::.7<5A1*:+8:58?.:;9.,2.;-2>.:;2.-27 *9*7 ;99$.26.:*7-=322 &1.;.:.;=5<;-.687;<:*<.<1*<68:.?8:42;7..-.-27+8<1:.0287;<8+.<<.:=7-.:;<*7- ;9.,2.;-2>.:;2.;<20*<.-..9.:?*<.:;?12,1524.5A1*:+8:=7478?7)8*7<1*:2*-2>.:;2.:;26=:<1A%1*;1*74 ,*-.62*%272,*/8:<1.2:1.5927/2.5-?8:4*7-5802;<2,;&1.;.728:*=<18:?*;*5;8;=998:<.-+A* %#% 2+*70:*7<.7<2<5.-G58+*5.>85=<2878/:*,1A,7.627**7-<1.2:SymbiodiniumH 5. LITERATURE CITED =:7.<<( 8702<=-27*5>*:2*<28727*50*5;A6+287<;B88@*7<1.55*./:86<1. 7-2*7",.*7 B8*7<12-Palythoa caesia *:27.,8580A#:80:.;;%.:2.; F =:7.<<( .7B2. .*:-68:. $A5*7- % 2010.7.<2,>*:2*+2522;28727 989=5*<287;8/*,587*5,72-*:2*7Zoanthus coppingeri/:86<1.:.*<*::2.:$../ *:27. 28580A
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Figures and Tables Table 1. ;3<>:.@6<;<;?6@2?6;B2?@64.@21.>
illoricatusEpizoanthus beriberEpizoanthusilloricatus Epizoanthus inazuma Appendix Table A1.
20°36.552',
North outside 1
20°42.338',
North of airport 1
North outside 2
East outside
North channel 1
North of airport 3
North of airport 2
116°42.129'
side
May 24; scuba
June 4, June 5; scuba
116°55.439' 20°46.897',
May 24, June 1, June 3,
May 23; scuba
May 22; snorkel
May 22; snorkel
May 23, May 27; snorkel
May 21, June 5; SCUBA
May 21; scuba
20°42.114',
116°46.234'
20°42.886',
116°43.281'
20°42.445',
116°43.213'
20°42.472',
116°43.701'
20°42.377',
Wreck on west
116°43.895'
116°43.401'
beach
May 20; snorkel
(2016); method
Longitude (E) 20°41.960',
Date(s) investigated
Latitude (N),
Dongsha I. south
Site
Gentle reef slope (13 – 18)
Wall, cave, reef slope (5 – 40)
Sandy & rubble (10 – 20)
Sea grass meadows (intertidal – 2)
Sea grass meadows (intertidal – 2)
(intertidal – 2)
Sea grass meadows, tetrapods
Shallow reef (2 – 4)
Sandy reef slope
(intertidal – 2)
Sandy, sea grass meadows
investigated in m)
Site information (depths
27
98
1
2
0
4
38
0
1
specimens collected
# of Zoantharia
Table 1. Information on sites investigated around and within Dongsha Atoll, and number of Zoantharia specimens collected at each site.
20°39.025',
Inner atoll 1
116°51.429' 20°35.110', 116°47.471' N20°35.247', E116°47.603' N20°45.995',
1
Southeast outside
2
Southeast outside
3
Northeast outside E116°53.822'
20°35.819',
116°42.408'
20°43.512',
116°48.373'
20°42.277',
Southeast outside
North channel 2
Inner atoll 2
116°41.711'
outside 116°41.711'
20°39.025',
Southwest
116°48.741'
June 4, June 5; scuba
May 31, June 1; snorkel
scuba
May 30, May 31, June 1;
May 30; scuba
May 29; scuba
May 29; scuba
May 25; scuba
May 25; scuba
crest (intertidal – 35)
Wall, cave, gentle reef slope, reef
Shallow reef crest (intertidal – 2)
Reef slope, wall, cave (10 – 30)
Reef slope (13 – 33)
Sandy & rubble (14 – 17)
Silty lagoon, patch reef (2 – 15)
Silty lagoon, patch reef (2 – 12)
Reef slope (14 – 28)
19
35
46
22
35
0
2
46
Dongsha I. south beach
Inner Atoll 1
Inner Atoll 2
East outside
North channel 1
North North of North of North of North North channel airport 1 airport 2 airport 3 outside 1 outside 2 2
Northeast outside
Southeast outside 1
Southeast outside 2
Southeast outside 3
Southwest outside
Wreck on west side
East China Sea CHINA
Taiwan
Hong Kong
South China Sea
Kenting
Hainan
Dongsha Atoll
Luzon
reef crest (intertidal) Pratas Island
PHILIPPINES
lagoon 10 km