Opposed Bands of Cilia in the Nonfeeding Larvae of the Serpulid Annelid Salmacina Tribranchiata
Total Page:16
File Type:pdf, Size:1020Kb
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`#,9!<CCMQ!:,;)/!()!,0J!=??CVJ!L;&$1;!@(!;,8(!0/))0(!]$,%)/),)/8(!/%.&#-,)/&%! &%!);(!#,)(!&.!0&44!&.!.((A/%1!4)#$2)$#(4!.&00&@/%1!);(!(8&0$)/&%!&.!0(2/);&)#&+;9!UI$)!4((!`#,9! <CC=Q!F-0()!<CC>V5!);(!#,#/)9!&.!%&%.((A/%1!0,#8,(!);,)!I(,#!+,#)/20(!2,+)$#(!4)#$2)$#(4!4$11(4)4! );,)!);/4!)#,%4/)/&%!/4!#(0,)/8(09!#,+/A!U`#,9!,%A!a,..!<CC<Q!Y,#)!<CCMQ!`#,9!<CCMVJ!L;/4!0&1/2,0! .#,-(@&#P!,++(,#4!1(%(#,05!,++09/%1!,)!);(!0(,4)!)&!);(!)@&!-,[&#!20,A(4!I(4)!P%&@%!/%!)(#-4!&.! 0,#8,0!I/&0&195!);(!(2;/%&A(#-4!,%A!-&00$424!U(J1J5!Y&.4)((!,%A!'(#%()!=?<<Q!:/!,%A!Rb^&/1;/0! =?<MQ!`#,9!<CCMVJ! Y&@(8(#5!);(!0,#8,(!&.!&%(!-,[&#!20,A(!&.!,%%(0/A45!3,I(00/A,(5!./)!/%)&!);/4!.#,-(@&#P!&%09! $%2&-.&#),I09J!3,I(00/A,(!U,!)(#-!$4(A!;(#(!/%!);(!%,##&@!4(%4(!&.!c$+#/9,%&8,!,%A!a&$4(! =??DV!/4!,!1#&$+!&.!=>?GE??!4+(2/(4!&.!-&4)09!0,#1(GI&A/(A!d.(,);(#A$4)(#!@&#-4e!Ua&$4(!,%A! '0(/[(0!=??<VJ!\&4)!&.!);(4(!4+(2/(4!;,8(!+0,%P)&%/2!0,#8,(5!I$)!%&!4,I(00/A!/4!P%&@%!)&!;,8(! .((A/%1!0,#8,(!Ua&$4(!,%A!^/)S;$1;!<CCEVJ!c$+#/9,%&8,!,%A!a&$4(!U=??DV!4$11(4)!);,)! 3,I(00/A,(!/4!4/4)(#!)&!,!20,A(!2&-+#/4(A!&.!^,I#/2//A,(!U,!1#&$+!&.!fO>!4+(2/(4!&.!4-,00GI&A/(A! .(,);(#A$4)(#!@&#-45!,00!@/);!/%)#,)$I$0,#!I#&&A/%15!%&%.((A/%1!A(8(0&+-(%)V!,%A!3(#+$0/A,(! U);(!d.,%@&#-4e5!,!1#&$+!&.!f>??!4+(2/(45!4&-(!&.!@;/2;!;,8(!.((A/%1!0,#8,(5!,%A!&);(#4!&.! @;/2;!;,8(!%&%.((A/%1!0,#8,(V!U^/1J!<VJ!L;/4!20,A(!/4!P%&@%!,4!3,I(00/A,J!3/4)(#!)&!3,I(00/A,!/4! 3+/&%/A,5!,!20,A(!2&-+&4(A!&.!3,I(00,#//A,(!,%A!3+/&%/A,(!U3)#$2P!()!,0J!=?<>Q!`(/1(#)!,%A! "0(/A&#%!=?<MQ!3)#$2P!=?<CVJ!B00!4,I(00,#//A4!@;&4(!A(8(0&+-(%)!;,4!I((%!A(42#/I(A!;,8(! .((A/%1!0,#8,(Q!4+/&%/A4!/%20$A(!4&-(!4+(2/(4!@/);!.((A/%1!0,#8,(5!,%A!4&-(!@/);!%&%.((A/%1! 0,#8,(!Ua&$4(!,%A!'0(/[(0!=??<VJ!! L;(!.((A/%1!0,#8,(!&.!4,I(00,#//A45!,)!0(,4)!&%(!4+/&%/A5!,%A!4(#+$0/A4!2,+)$#(!.&&A!+,#)/20(4! I()@((%!d&++&4(A!I,%A4e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d&++&4(Ae!I,%A4!,#(!2,##/(A! )&!);(!-&$);!I9!);(!2/0/,!&.!);(!.&&A!1#&&8(J!\(),)#&2;,0!,%A!.&&A!1#&&8(!2/0/,!,#(!%&)!P%&@%!)&! ;,8(!.$%2)/&%4!&);(#!);,%!/%!.&&A!2,+)$#(!,%A!)#,%4+&#)J! L&!-9!P%&@0(A1(5!%&!.&#-,0!2&-+,#,)/8(!,%,094/4!&.!);(!(8&0$)/&%!&.!0,#8,0!2/0/,)/&%!&#! 0,#8,0!%$)#/)/&%,0!-&A(!/%!4,I(00/A,%4!,%A!4+/&%/A,%4!;,4!I((%!2,##/(A!&$)!/%!);(!2&%)(T)!&.!&$#! 2$##(%)!$%A(#4),%A/%1!&.!);(/#!#(0,)/&%4;/+4!U^/1J!<Q!.&#!(,#0/(#!,%,094(45!4((!a&$4(!=???Q! c$+#/9,%&8,!=??NVJ!Y&@(8(#5!);(!A/4)#/I$)/&%!&.!-(),)#&2;,0!,%A!.&&A!1#&&8(!2/0/,!4$11(4)4!);,)! );(4(!2/0/,#9!I,%A4!@(#(!+#(4(%)!/%!);(!2&--&%!,%2(4)&#!&.!);(!20,A(!U3,I(00/A,!g!3+/&%/A,VJ! 3/%2(!);(4(!2/0/,#9!I,%A4!,#(!&%09!P%&@%!)&!.$%2)/&%!/%!0,#8,0!.((A/%15!/)!/4!+&44/I0(!);,)!);/4!U&#! ,%!(,#0/(#V!,%2(4)&#!;,A!.((A/%1!0,#8,(J!B.)(#!);(!(8&0$)/&%,#9!0&44!&.!0,#8,0!.((A/%15!&%(!-/1;)! (T+(2)!);(!#(0,)/8(09!#,+/A!0&44!&.!-(),)#&2;!,%A!.&&A!1#&&8(J!Y&@(8(#5!'(#%()!U=??NV!4;&@(A! );,)!%&%.((A/%1!0,#8,(!&.!A/8(#4(!4,I(00/A4!_!-(-I(#4!&.!,)!0(,4)!(/1;)!1(%(#,5!A/4)#/I$)(A!,2#&44! );(!);#((!-,[&#!20,A(4!&.!4,I(00/A4!#(2&8(#(A!I9!c$+#/9,%&8,!,%A!a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h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f<?!-!A(+);!%(,#! `;/)(!'&/%)5!:&4!B%1(0(4!6&$%)95!6,0/.&#%/,5!&%!<!W&8!=?<CJ!B11#(1,)/&%4!@(#(!+0,2(A!/%!,%! /%4$0,)(A!2&&0(#!,%A!)#,%4+&#)(A!)&!);(!637!:&%1!"(,2;!\,#/%(!:,I&#,)	J!L;(#(!);(9!@(#(! )#,%4.(##(A!)&!,%!,]$,#/$-!;&0A/%1!#(2/#2$0,)/%1!4(,@,)(#!,)!<Mi65!@;(#(!);(9!@(#(!;(0A!.&#!);(! %(T)!.(@!@((P4!@/);&$)!4$++0(-(%),#9!.&&AJ!`;(%!%((A(A5!0,#8,(!@(#(!&I),/%(A!I9!I#(,P/%1!&..! ,!4-,00!Uf=GN!2-!A/,-()(#V!+/(2(!&.!,%!,11#(1,)/&%5!+0,2/%1!/)!/%)&!,!4-,00!A/4;!&.!4(,@,)(#5!,%A! 2#$4;/%1!-&4)!&.!);(!,A$0)!)$I(45!.#((/%1!,%9!I#&&A(A!(-I#9&4!&#!0,#8,(J!F-I#9&4!,%A!(,#09!