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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 Revision of the nudibranch gastropod genus Tyrinna Bergh, 1898 (Doridoidea: Chromodorididae) Michael Schrödl; Sandra V. Millen

Online Publication Date: 01 August 2001 To cite this Article: Schrödl, Michael and Millen, Sandra V. (2001) 'Revision of the nudibranch gastropod genus Tyrinna Bergh, 1898 (Doridoidea: Chromodorididae)', Journal of Natural History, 35:8, 1143 - 1171 To link to this article: DOI: 10.1080/00222930152434472 URL: http://dx.doi.org/10.1080/00222930152434472

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Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 ( Revisio J Th Introduction ournal Doridoidea e Th mai e chromodori n n of diagnosti of anatomy K Chromodoridida describe (or to lectotyp ( know du 198 formations an digitiform natura of 1958 sections contras an of an delicata T. F Bergh T. belon Th ( Universit ³ Mu ² MICHAE Accepte 1996) ollowin Departmen Zoologisch N eywords: d d d e Cadlinella morphologica nobilis e California nobilis th : 8 È a nchen Tyrinna histologicall to genita rediscovere convolute ISS ) tural Chromodorididae g n fro , e is l to t 189 it N di Japanes e d is group m nudibranc of d to s redescribe g , 0022-293 Bergh Œ th is , c d y proposed to , phylogeny possessio l instea 1 8 . th er Cap previou T. wit e L ) organs German herei characte genu Boulevard Th ar Marc du , e genu H Mollusca allo e sp. t , in d SCHR nobilis h e norther ne e e , istor e e d of th Staatssammlun y d re-examine l ’ vagina 1898 e 3 n a w to V w thei specie placemen variation holotyp e usin s s h prin whic reveal d . erd s of Zoology designated dee Tyrinna n . Tyrinna thei genu A certai cod 200 description anatomicall . In y r . y , of h t g r e n O it Th / p http: , an vestibula È ISS h , ; tentacl s s r , Chilea thi e 0 Island of DL 2001 Per gastropo s longitudinall an s Journa media spiculose e d place ) Vancouver Nudibranchia e n replacemen doe of N a // s , Tyrinna t Panami T. of d d Bergh www.tandf.co.uk Tyrinn regardin u generi study 1464-526 hig ² , ) of nomenclature an s (3 , , Bergh pusae e an Dori Universit s n l d an n an h 35, of s no r d ) in th an shape y d in of g longitudina glan d degre , tw , Natura d ar , c t e an , d s c 1898 Tyrinna T. Jamaica tuberculat a 2 Mu Tyrinna 1143– Tyrinna SANDR Cadlin e o Marcus g d placement delicat belon du incompletel Argentinia an , d t onlin , y d her nobilis radula nobili : 189 or e Britis genu e by È histologicall d l `enrolled’ . nchen, aroun Cadlin , y Histor Th of e to e amphiatlanti thre / g Chromodorididae journals 8 a th considere of , a 1171 Ñ . e parallelis by , , s by th to r sagamiensi retentio is Abraham 200 Tyrinn Bergh e l spicule typ 1959 h s e e y d character Britis A e Bergh . a groov Muniaõ senio notum a n Columbi 1 Tyrinna submargina Mu Rudma Tyrinna, th y A e synapomorphies , japonic V. Taylo specimens smal specimen , e digitifor describe , as phylogeneti y È a nchhausenst s vagin 189 d r n h e m MILLEN usin , hav nobili r . wel  c to bu n , Columbia (`enrolled’) of l Elio s & s 1898 an n 8 187 a specie bu et an Baba t be e a Franci l a g bu is ( th d Baba m as 1984 d bee disregarde s s t d 7 al is . V6 l specimen Bergh, t redescribe e uncertainty synonym t of (1913 The , and , Cadlin proportion row . tentacles of fro controversia absen commonl s , n wa th s ( T c th Tyrinna ) 1937 , 1996 T. Lt foun Chromodori ³ m y e s discussio e s : an r 1Z4 , 1937 ; T. d sho ) Chilea ( evelinae of Chil 627 (2 typ 21 to t a d include 1) , ) s d evelinae in s d o ) 1898 w d ar burnay . mantl Munia , , , seem fro in of ora e 0 e , . long Thre y 8124 T. synony anatomicall Canad e incompletel s a fo n is specie m histologica of taxonomy, T. use transferre n wid r l nobilis congener s s show (Marcus, e , l s tentacle th digestiv e of whic ` delicata i õ d justi  7 Cadlina n taxon juvenc narro e form derma poorl e Ortea, m rang nam a basa et Gul s n h . ® T al of ed In to w d a a y y y a e e e s s f . . . . l l l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 et duc an mantl ora tentacles th 1144 maxima an wer severa Materia monophyl re-examined det. Dori synonymies an characters th of (Schr of an et ha Japanes smoot genus additionall pecte tropica Rudma know with latera by synonym pusae detaile atic consequence spicule e e d d d d d al. the al Marcu specie l Th Durin T Th A . t e genus reproductiv auriculat Muniaõ s examine . va alread o Chromodori leavin describe tentacles d No o foun n l e È delicat secon l h ( e e Tyrinna d ( dl Marcus genus s radula l thes s 199 a localitie glands n 199 Chilea e anatomy l l presen an mantle , ’ water origina s deferens s value ( an ` vestibula g species , s ( littl y ; d 198 1996 6 by y g Bergh d 6 y e  wa relate th , an ( d n, d ) an ; ) s in a 195 th assigne th d d purpose Tyrinna e th bee r e e method , species 4 con , th Munia n d of . adde s s hav , T. t s d ora e teeth ; e bi ). s e , th anatomicall puzzle a year in Thi e stud e 9 l ( an specie of th vagin 195 , 1997a n on systemati s , presenc , d ` th ® 3 Du , typ evelinae ® ), e smoot enrolled description Cadlin taxonom e an l result juvenc 189 d e ) r d livin rme length an considere d e tentacle s localitie to , i.e bee s d 9 th to ja y glan ` d e , e õ s to wer latte wa presenc  s n 1991 tw d ’ 8 withi a th Cadlin e . s w is to ( ) d specimen specimen revis digestiv T. n ). g of Bergh structur et dista Chilea s (se s vaginall h di e o a e wit elements th a e d it thi re-examined , r Rudma condition du ar incompletel ’ Œ det Bergh of c no al crypti mantl ± sagamiensi s widt diagnosti e e nobilis n coverin y s character y s erence ora 1998 h e s M. placement e wid . e continuatio Schr l ( d a e t an th a s . give as , criticall Baba ( placement to th use n to considered Thompson 199 evelina e of vestibula to l h of e d 1898 e e Schro y s an e , c e s thes , tentacle wel sam system o an tw n geographi , an systemati n 189 d È s scub transversa of 6 of endeavou specie inserte belon dl T. wit a and g d , ( . ) d in o in d to c 1937 l 198 ; , wid e Argentinia ¨ . th 8 e T. althoug y s Measurement T e dl h heigh y . nobilis, row 199 as positio ducts character Marcus . a tabl an th Baba study e Additionally revis th . Marcus describe T. 4 ; g rounde e , n r s an nobilis divin s , nobilis e d vagin histologicall . ) d e 6 radula , ( despit s gland 194 aroun of to Th e an characterize 2 ), c d specime C. Dori t genu evelinae sper c of . e r ) 1. , placemen l . of In n earlie h S. distributio wer d e Tyrinna 9 193 g th ( no , furrow to sagamiensi mantl a relationship , ) In 1 of , th d th n a , a m V. s s wa e d 195 s e d and whic r ) 1958 T r e of an 7 delicat e redescriptio coasts investigat e bump Tyrinna fo ribbon broa thei th Cadlin total to genu T. r . uncertain receptacle Mille fo s n mai an s , crawlin d 9 r , e T. wor pusae e use r T h ). s i.e museu descriptions of , y describ the evelinae r d stomach Tyrinna t derma by d wa it ar d , . n on s nobilis of . s . by spiculose d n Cadlin a Bot n k s 15 evelinae , a y livin Specimen radula Tyrinna e s distinguishin Tyrinna regardin an Abraham . in s , to a Tyrinna on Rudma Chilea in burnay discusse and th g s e seria Chromodori transferre lon h specimen m d bot . of specimen e l s e loo contras g n th Chilea . , formation specie a , tw specimen in wit wa inne g Muniaõ T. of C. e . specimen th h an l k japonic an , n o , Tw hav i , as Paci sections variabilit s detai e n h nobilis T g fo japonica d tuberculat s specie d Ortea an Orte r d , congene row mentione . of absenc n ( th s r denticulation havin o t 1877 th th nobilis e s unarme sid 198 d d to s ® Tyrinna nudibranch  g e n wer Tyrinn e l poorl e wer bee c s to s (tabl a a to s e feature th th inseminatio s , s phylogeneti et 4 s an and structur of , juvenc of Baba, . ( 198 is g of ). ( e Tyrinna n e tw e e e y 198 re discus MDFs) have To al d , r ` diagnosi T. problem y e margina anatom regarde enrolled collecte collecte Muniaõ Munia e d ¯ d of o of . T. th Atlanti a 8 Tyrinna 2 ec 8 know species ( vagin notu poorl nobili nobilis clarif s ). in e a to 1996 ) pusae 1937, nota t thes been use e s ora Th sus and s s th th th , by it of of at m õ   n n d d n n d d y y a y a e c e e e e c s s s - ) - l l ’ l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 wer tion seriall genitali fro Te nudibranch ma th Bod bearin Baba fro K. Th Museum sectione tion Thompso examination fro ( dista ( nobilis Island dissecte describe specimen no as partl Berlin Marcu 1959 (SMNH Museu Natura 19960724 tabl MAC e n t typ e m m m y Hazegaw Th Generi On Tw Tw e . s whol mentio y of large , l y e be ’ deposite of Vouche th s y e , Japan Per as genitali s whitis g e 1 dissecte e o o s N m th l a fro ( d mentione materia ZMB ) sectione e d tw slightl small d undissecte entir , 1937 , History ( wel e e usin s undissecte specimen of u r n ). Ba no no of on 195 c an by Toky m large o fro n notum on s diagnosis ( wa , h . l large . specime y . e d a 198 e Natura an r , g , wer th 9 Berg , a m d as 373 wa e 3387 d y Caribbea wit as of ( 194 specime examine lo ). no s scub specimen l o r e y Tsukub wer ) (no in d prolonge 0 th , fro specimen w s of specimen d e give r 6 Colium ( paralectotype materia . Th h Muse ) . ( BMN h 9 e th d NSMT s 3 ) examine specimen by 2. relaxe bosses . 5075 e wa Anterio d a Chromodori ) m orang ) ` ( an l n 4 e e holotyp of Isl . 5 Revisio partl descriptions n an 189 identi ) Berg Histor syntype s ` Zoologisch m no n undissecte d Quiriquina d CHROMODORIDIDAE a o a is n m) Bod to d Tyrinn 0 H provide s of 8 th o de by d . d Researc l ) . ® Argentin e herei , y s of ). ), materia s fro 9 h Few xe e (no no th wa an d laterally r Cadlin usin ® (SMNH wer to dissected y Pajargo s n histologica Th e y no ( wer ed paratyp d T foo e wer 189 m d externally elongate s s s of . of a re n kindl in , . . . e e 4, author s g staine d studie 8 of by Tyrinna tha e chunk t d designate 1995038 nobilis A juvenc dissecte evelina d 8 entir e h Dori th , ), Bouin e a a l fo ’ 12 embedde margi ) o , bu studie spot Center Cadlin Staatssammlun e . Munia t third part holotyp sagamiensi assigne appear ’ 10 y Chile r , ) e . Ora de localit ( nudibranc t no d e s re-examinatio d Isl mad % , fro s (SMNH y illustrate a s l delicat mantl . ’ specime wit e s fro s s, externally n Ciencia by spicule a , d Bergh, d Bergh means l , scattere MgC m Thi , of 1544 a wer ; d smal tentacle on õ de an  wa separate externally collecte e s h n d m d y in e Kameki as Carlo to japonic th availabl toluidine-blu s e d it et Pa e s in to e of a Lo lit l s wa , e , l describe be specime , lectotype 2 wide . re-examine  is s h n d collecte s jaros 189 Baba al. 154 Abraham 1898 .). hydroxyethylmethacrylat no specime notu d solutio Cadlin s ma T s genu probabl is Bergh, lost ( Naturale s s d . g Horno dissected Baba aroun d . . a ( 8 5 long n Tw , her y , Parede by 199 373 ; Mu e , ) nobilis foo m , ( int Japan . Baba on be by nort ZMB s 1937 assigne An th n d o e Plate a d È n of 6 7 o T , , , nche e n t anatomically presen e d designate 1891 ) ) wa th specimen 1949 followin s digitifor e , yrinna e smal evelina narrow in an d tw entir h fro specime of , s s partl wa 187 th , tha an e , fo . , s of . NSM externally d ( o 1937 ` no wer On n th ( e BMNH Bernardin m Sinc Zoologische d Tyrinn r d s : Lima no l t 7 ® ( lips e Coquimbo nota e t histologica . pl specime to re-examine y ZSM e th xe the th e 5072 ( e specime in t . Gul . , Britis preserve m g e T dissecte e d studie d , Notu e s Tyrinn n mentione Marcus xxi y th Nationa ) th Berg l Ba th no wit a as (on in Jua no refe : , margi f 5 e fo e . , e pusa . no posterio ) . y On h mantl lectotype ® of . 70 h suggestio r n o e Th m wer n 59 foo d h n gur r 1 of Museu s a wa a e d anatomica % d ) entire s Rivadavia e l Fern ( externally of smoot to , , Californi e e d 19960723 an n 189 nobili originall ( blun e examina specime t l specime probabl holotyp Colium e d Museu s 195 Kulzer) Marcus, Swedis ethanol e an Tyrinna labelle th d or corner Scienc 78 seriall withi tissue 8 r a e dista d ,  t 8 m 1145 ). ) ndez , ove por h s n tw on tip di th th by by or of of m o d h n n n o d y y a y e e e e e s r - - , . l ’ , l . , . Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 variabl Radul th thin derma semicircl wit 1146 is opaqu line collecte norther th Gul L.U.C.E. Chile H Marque S. di depth by 73 lecte 1995038 Cadlin Glossodori Chromodori Tyrinn Tyrinn Dori inserte convoluted vestibula by caecu thes usuall pair Œ olotype e e ß surrounde Gigglinger h a th 45 eren Additiona Typ Externa Materia 1996 1996 1959 s vacuola f e s , s L.U.C.E d lon , e ru of spherical m ¾ delicat a ma , a y a a of 45 a l e by d 10 har L.U.C.E e d  e n n juvenc t : : : pusa nobili s wit formation dee Ancud present whit ; g broa ² on in e 29 y r 265 22 specie s Chile W denticles m togethe Chilea alon Plate Chile ( muscula , d P be s in juvenc 45 accessor ± . h , shape p a l SMN r l ara e th ), depth juvenc 33 ± substrata Burs pl e d . s cell a d Abraham l morphology , examine 273 inne ß Marcus completel fron media Strai g : e . 58 , , lines Chromodori specie Bergh . , K. , by . s ® Marcus . wit type 3, . muscula collecte vagina . n Thre th Chile a s ; gure ¾ Va Tyrinn r a , Statio H S, : r a Schr ® , ar t , Salger s e r smal orang an h wit Carcelle copulatri t gur Tyrinn , smal y Bergh s an no of Tyrinn portion an e l nota , of ( , 67 s an , s abou d e deferen MDFs) . , d 1898 , gland clustere h 45 longitudina SMN o 1959 , d e . , È d . ß specimens Chacao 24 th d 36 l n dl 1877 3736 a y Argentinia d 15 34 1959 (table th l ± r oute e Tyrinn an , l , . wit e opaqu M 53 reduce by stone nobili 1996: m ; a Januar ¾ M. : capsul s t e an 1898 : to W s a d 1997a ana s 523 ; : , H an 33 nobili as lectotype Livin : M. x h ; 42 L. depth Rudma juvenc s d nobili 259 r , withou claw-lik ), 87 Statio ma d re Schr an s d ± long : th , ± denticles or no s peda l man s 34 a 22. in Plate ( ; d e labelled Chile Schro 531 526 Williamson 1 d : ± d 42 : e papilla wit y Schr e y , evelina 260 s , g . whit H 38 l spot folde excep withou s receptaculu , o SMN an ther n ® , a 1949 inne ß be È Bergh ± , 3737 n n groove y olotype specimen roug h dl 20 l gure ± : ( wit t 532 (® ; , pl ¨ localitie o d Abraham e 42 1984 . , edges M È armature dl L . Schr unde teet calcareou e dl ¾ s gure 40 . absen e 50 bearin d On h r . ectotype 2 , , . , t : pigmen 30 ; s e H formin 10 ar an h 1996: pl collecte ). layers al ² , on Tw m a ro t ` : h 54 Statio ( o S, 13 , e È , . 244 1898 san e . 0 no r . s Marcus l d relativel dl a w ® depth 1951 31 ± pe entir , stones t Th ( highl o Fiv th othe 1± 73 Dori gure s 64 s S. g 41 . 1996: Januar projectin ± ZMB m d , 21. r containin or e , 1544 or 246 . . (se equa g ar 6 ß ; e ® . t s V. ß n e : wit 22 187 hal Ja Commo outermos d gure ) semini Rudma 49 s e , r als rhinophore tw e algae 314 s , y present e thre ; to M by ¾ w ; ZM smal 21 , specimen Mille lateral 00 ¾ 22. y h Munia f translucen , delicat 25 , vacuolize 7 o o P tabl l 195 . s y rodlet ± 1545 no shor ) ro shells aralectotype size ² si th 10 occur ² or 4± 29 e W B , g comb. S, 1995 x s l w n 8 e . g e 5 6 submargina n , 11 stones n clos ), d ), . media no 50750 s thre 73 L.U.C.E õ bi 1 1984 t ;  pl t Ma n a th as ( genita ; , s Vagin ), prostati denticles Statio Chiloe onl : teeth T 41 - s . , et ß Odhne . 5 , e bi e s 48 32 d H rows on e . 33 to y MAC nov 50725 collecte : al t ß Ma together ® larges ar y 48 , burnayi ¾ olotype ; 244 , cells n irregula 1949 58 . d whit 4 mm . e wit 1996 ® l th a tripinnat Calbuco . ¾ n  or y 40 Ma Salivar cusp ² gure r vestibule . . marke ± Island W c e M long N Eleve 1949 1921 . , 246 h , Rhachidia , ² ’ l , t e seldo portio gills . : S, or In ZM d y . Isl ), collecte row s oute 265 ones no 94 . Tyrinn , ; 1949 Opaqu (Ortea r Chiloe , , by 21 a short-stalked : elongat 73 large , Munia Firs d y . . , n submargina B 226 ± ( , BMNH narro m collecte s e ± de Th 33873 Chile 41 273 r ß gland . by specimens ; n Chile 24 no S. 21 of , gill tricuspid branche denticles . t Withi ß collecte followe r e d a ; Pa 46 ;  ¾ , . e 00 media lateral n õ Millen  Schr Marcu mantl Island, MDF n notu w 50725 s 198 by  e pusa , ¾ jaros, whit , s , 30 ² teet 70 et d wit in W Pta an col an n no. th 8 o ² by È al m m S, dl e ). ), d d d h h n d d a a e e e s s s - . l , ; , : . , , . Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 Tyrinn Species contraction ma tentacle givin ar in livin nin elevated macroscopi in to 13 darker. pigmentatio centra (se th fronta rows Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn Tyrinn e e th e mm contraction y widt e Crawlin positio lon g ® e . g bi be gure a a a a a a a a a a a a l l specimens Th centr it - in g h s regio evelina evelina nobili nobili nobili nobili nobili nobili nobili nobili nobili nobili par , additiona to e an whic smoot an an widt s n diamete t e c . g 1A,B) tripinnat n d d of ` s s s s s s s s s s n Anteriorl of of papillae enrolled e e specimen h , h digitiform becomin 8 ther larg th h th th is . mm an In sheaths e . e e l ( of Bah Peru El ( Punt ( Bah ( Comodor ( ( Sen ( La Niebla ( Bah ( Pichidangui ( Los ( Play soli r d e 28 42 44 46 44 44 41 36 32 29 26 e Ther larges gil transvers notum of preserve e Revisio whit is California 6 Rubio . ’ ß ß ß ß ß ß ß ß ß ß ß in Arena o õ õ õ l 57 38 53 02 17 30 40 32 05 38 12 y d    a Th Hornos a a a mm a gill Collectin shape s ( usuall g Ventisquer circl th an 3 Lo th e ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ an de Camarone de Pardelas height ; S, S, S, S, S, S, S, S, S, S, S, hav ( ß . e e t e ar s whitis 52 39 d e the s Th , d . rhinophore preserve subepiderma lo Colium 113 64 65 67 o 72 72 72 , 72 71 71 70 in spot e whic coordinate foo e s. ¾ ß Piquero n Lenc Afte S, . e s 48 rounde Rivadavi d e y ß ß ß ß ß ß ß ß ß ß y du e no height of 16 39 35 40 35 40 57 32 20 39 Th , ß Angeles of Tabl . g specimen an 80 ¾ fold 31 t rhinophore onl Mexic S, s hav h e , ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ h W W W W W W W W W W localit Th Tumbes a r is orang e th o ß ¾ Valde decrease W s to o 73 50 y e elongat or for ® groov ) ) ) ) ) ) ) ) ) ) s divide e d s e e 1. xatio ¾ d ) ß . W contraction on 22 nudibranc o on a specime , Stationar s a m nota in s y yellowish z Gul e ) ¾ e W Collectin an long l , s a to th e cross-section n gland ro d ) th s e d f l s doe e semicircl int w th re fro e surfac th bod , bea inne 29 20 11 09 08 26 08 05 27 17 06 14 21 n 26 d e ana of dee o e s m s y onl h spot Collectin specimen . r g Jun Septembe Januar Januar Januar Februar Februar Februar Decembe Decembe Januar Marc Decembe tw gil . y Januar not whic specimen spots r p th genu l Th up data Th e y shap l o papill sid e e e , s longitudina cavit e reache is lip e 199 h to h however inne no in . . e ora . y s y y y y 199 ar rhinophore e s slightl In y y y g Mos 22 T 199 of r 199 199 199 6 199 a r r r fron y s whic r e 199 199 199 reachin dat l yrinna subepiderma r 199 199 199 4 s preserve ma 197 ar los tentacle shinin s lamellae th 8 5 5 5 to 8 hav t 8 5 5 e e 25 t 4 7 4 e y h y t 4 , of surrounde th of reac thei ora mm appea l e ar folde e g Number g thes groov an specimen e th oute s s d l throug r 46 h foun . no e in in tentacle ova Ther d usuall specimen orang e r 3 1 1 1 th 2 2 1 2 1 1 1 1 1 1 ana mm t l length r livin spot e to bu d notched e gland l rows d (® of bas shape l h e s t be by y e in papill s gur g ar th bear s e , an situate indicat animal . becam e s slightl e Dept 15 10 length s du 10 e In of 0± 5± 0± 0± 5± 3± 0± 5 5 9 8 0± (m) in . si tissu d . ther 1147 1C s Du e a x Th mm 10 20 8 5 8 4 30 1 re th th th no h to to in ), d d y e e e e e e e e e s , Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 1148 Table 2. Comparative data on Tyrinna nobilis (bibliographic data in quotation marks).

Body size: Approx. no. of length/width/ Tail projecting denticles of the height (mm) behind the Size and shape of second, the Length (mm) Specimen identity, Collecting alive (l ), notum; alive rhachidian 30th and the and position of reference locality preserved (p) (l )/preserved (p) Radula formula teeth outer laterals vas deferens No. 1 Los Hornos l: 46/13/6 l: yes 81Ö 66.1.66 big, in older rows 6± 0± 6 13; extending (this study) p: 24/14/7 p: yes smaller; in younger into the left ZSM: 19960723 rows big and blunt body side median cusp with 3± 5 M.

denticles on each side Schro No. 2 Lenca p: 16/9/5 l: yes 75Ö 60.1.60 small and claw- 6± 6± 7 12; extending ¨

(this study) p: no like in older into the left dl rows, elongate body side with a sometimes an bi® d median d S. cusp in younger rows V. Mille No. 3 Pardelas l: 30/12/5 l: yes 79Ö 65.1.65 small and claw-like 5± 1 to 7± 8 28; limited to (this study) p: 19/8/5 p: yes the right body side n No. 4 Bay of Coliumo p: 16/9/6 l: yes 81Ö 66.1.66 small and claw-like 6± 3± 8 16; limited to (this study) P: no the right body side No. 5 Bay of Coliumo p: 10/6/4 l: yes 65Ö 47.1.47 small and claw-like 5± 6± 7 juvenile (this study) p: yes ZSM: 19960724 No. 6 Camarones l: 35/13/6 l: yes 93Ö 60.1.60 small and claw-like in 5± 0?± 8 40; extending (this study) p: 25/10/8 p: yes older, elongate and into the left with projecting median body side cusp in younger rows Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007

Table 2. (Continued).

Body size: Approx. no. of length/width/ Tail projecting denticles of the height (mm) behind the Size and shape of second, the Length (mm) Specimen identity, Collecting alive (l ), notum; alive rhachidian 30th and the and position of reference locality preserved (p) (l )/preserved (p) Radula formula teeth outer laterals vas deferens Revisio No. 7 Camarones l: 35/13/7 l: yes 91Ö 74.1.74 elongate; irregular, 6± 0 to 7 38; limited to (this study) p: 24/10/9 p: yes with more or less pro- (small )± 8 the right body

jecting median cusp side n

No. 8 Bay of Coliumo p: 6/3/2 l: yes ± ± ± juvenile of (this study) p: yes th e

No. 9 Bay of Coliumo p: 18/8/6 l: yes ± ± ± ± nudibranc (this study) p: yes T . pusae, holotype Chiloe Island p: 30/25/11 p: no 90Ö 95.1.95 small and claw-like, 6± 0± 6 c.40; limited to (Marcus, 1959) (41ß 49¾ 25² S, some more elongate the right body

73ß 48¾ 58² W ) with paired median side h cusp genu T . pusae, paratype Gulf of Ancud 26 (medially p: no 108Ö 125.1.125 strongly ground oŒin 1± 0± 2 to 6 c.40; limited to s

(Marcus, 1959) (41ß 48¾ 40² S, along the notum: older rows, elongate the right body T 73ß 21¾ 00² W ) 42)/16/10 with paired median side yrinna cusp in younger ones T . nobilis, holotype Calbuco, Chile p: 14/7/4 p: yes 68Ö 55.1.55 claw-like in older severalÐ 5 to 7, juvenile (Bergh, 1898) rows, projecting rarely more median cusp with 3± 4 denticles on each side in younger rows T . nobilis, M 94-1 Street of Chacao p: 13/6.5/5 p: yes 64Ö 70.1.70 small, claw-like 3± 3 (reduced in c.20; - (Marcus, 1959) (41ß 46¾ 30² S, teeth No. 1149 73ß 45¾ 45² W ) 40± 50)± 4 Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 1150

Table 2. (Continued).

Body size: Approx. no. of length/width/ Tail projecting denticles of the height (mm) behind the Size and shape of second, the Length (mm) Specimen identity, Collecting alive (l ), notum; alive rhachidian 30th and the and position of reference locality preserved (p) (l )/preserved (p) Radula formula teeth outer laterals vas deferens T . nobilis, M 94-2 Street of Chacao p: 11/5/3 p: yes small, claw-like all 3 to 7 ± (Marcus, 1959) (41ß 46¾ 30² S,

73ß 45¾ 45² W ) M.

T . nobilis, M 42 Chiloe Island p: 30/12/8.5 p: yes 94Ö 116.1.116 elongate, paired pro- 4± 0± 6 c.60; extending Schro (Marcus, 1959) (42ß 20¾ 50² S, jecting median cusps into the left ß ² 73 22¾ 00 W ) body side ¨ dl , Punta Maqueda l: `68 10’ `yes’ `78Ö 52.1.52’ ± ± ± T . nobilis / an (Muniaõ n et al., 1996) (46ß 01¾ S, d ß ¾ MACN: 33872 67 35 W ) S.

T . nobilis, Punta Marque s l: `46/12’ `yes’ `73Ö 56.1.56’ `small and claw- `4 to 8± 0?± 4 to 5’ ± V.

(Muniaõ n et al., 1996) (45ß 58¾ S, like, in youngest rows Mille MACN: 33873 67ß 34¾ W ) elongate, with paired central cusps’ n T . nobilis, Punta Marque s p: 19/9/7 p: no ± ± ± ± MACN: 33873 (45ß 58¾ S, 67ß 34¾ W ) T . nobilis, Punta Pardelas l: `52/14’ `yes’ `69Ö 48.1.48’ ± ± ± (Muniaõ n et al., 1996) (42ß 36¾ S, MACN: 33874 64ß 16¾ W ) T . nobilis, Punta Pardelas l: `39/13’ `yes’ `76Ö 59.1.59’ ± ± ± (Muniaõ n et al. (42ß 36¾ S, (1996) 64ß 16¾ W ) MACN: 33874 Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007

Table 2. (Continued).

Body size: Approx. no. of length width Tail projecting denticles of the

/ / Revisio height (mm) behind the Size and shape of second, the Length (mm) Specimen identity, Collecting alive (l ), notum; alive rhachidian 30th and the and position of reference locality preserved (p) (l )/preserved (p) Radula formula teeth outer laterals vas deferens n T . nobilis, Punta Pardelas l: `25/15’ `yes’ `83Ö 67.1.67’ ± ± ± of (Muniaõ n et al., 1996) (42ß 36¾ S, th ß ¾ e MACN: 33874 64 16 W ) nudibranc C. juvenca, Isla de Pa jaros, p: 8/5/4 p: no ± ± ± ± lectotype, ZMB: Chile 50725

C. juvenca, Isla de Pa jaros, p: 9/5/5 p: no `62Ö 54.1.54’ `small and ± ± h paralectotype, ZMB: Chile claw-like’ genu 50725 s

C. juvenca, Juan Ferna ndez p: `7/3/2’ p: `yes’ `56Ö 50.1.50’ `small and claw-like’ ± ± T (Bergh, 1898) Islands, Chile yrinna C. juvenca, Juan Ferna ndez p: `10/± /± ’ ± `4-cusped median ± ± (Odhner, 1921) Islands, Chile tooth’ Doris delicata, Coast of Chile p: 20/9/6 p: no ± ± ± ± holotype (Abraham, 1877) 1151 Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 1152 gland surroundin reach tissu con ica projec narro foo F ig. l t Ther Mantl ® 1 e examination rm corner . s w (® t Scale ¯ no shinin bar 230 (mantl agellifor behin ( an gur e th . A) 5 e 6 ar m e d ), g structures s 1 e bar g m Tyrinn presenc e tapere ventra cm. concentri ar 1D) d e throug tw m 5 in e th derma 100 . elongate ( . o e penis B) a length Thes However l d notu view or e h nobili m . posteriorly Tyrinn m. of l th . c formations thre e Scal . Spicule e m layer a . Not spicule s d ventra fe Cross-section (specime ( e a an e tabl , M. w e s maceratio ba submargina nobili d th calcareou (® s l Schro e r s e . slightl 5 appeare gur notum 2 deepl ar Afte , s n ). 2 compoun e mm. (specime e ¨ no chunky dl 2A) r y y . . n grooved ® an s tapered 1) s Scal d ( xatio l of D) . spicule , sho d to row dorsa n e , S. d mantl Tyrinn be w somewha n ba no glands) s , V. . digitifor th r absen . l tha s of In 5 view Mille 1 e in e a 1 ), t livin contracte spherical cm. tissu nobili th ventra . . th t t Th e Spot n m durin e banana g (C deepe e ora s e specimen spicule an ) s l MDF (specime l Tyrinn g wer , view d d tentacle r externa subepiderma tai - histologica layer e or . s s orang l Wit a of n ar s ma rice-shaped s nobili th s e no th an l h y of e e an e hollo . d whit or tai in oute 7) th s d th l , (specimen l life l e ma e anatom e section spicules w is everted mantl mantl r MDF . y ban , long Scal wit an no d h d e e s e - t s , , . Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 be contai attache Ther specimens. or T of large situatio apparentl of vacuola MDF interna lik but of clearl (Cantraine layer closel ther MDF wit all to vacuola (Orte ar respectively (® reac F ig. . e gur y e specime contai th viole ventra separate h , pusae Centra Withi surrounde e h 2 (® patter in e e r ’ ligh a e . y y s . ar 0. n structure no foun gur surroundin MDF is 2B) l In d certai visible an n t r r resembl of 3 Tyrinn e severa . substructure.Th t n stainin n l cell y cells mm to a e by of d th 1 l bluis cell n mantl n , on , specime larger d ), 2B) d invaginate . separat nervou e in Pe 183 no s s n eac toward Bot Marcu smalle in e int scal . , s peripher a  d in havin wit , l rez positions large th . h . bu e scal 5 g containin h o by otoconia nobili 1 Chromodori e Th h diameter e , e content ) h , epiderma g t cells thi surfac an inne cerebra s ba smalle n by oute e 198 e e cel a les cel r , s g s ba syste s Revisio r vacuole central s thi no anterio rhinophora MDF ( d , vacuola 5 s . th l laye y l Garcia-Gome r 195 3 r r Thes limit histologica laye densel fold , 5 50 . n ) of e MDF e . e . muscle r an g m , 1 200 by l Th an muscula 9 r interio m Th no degre l th th s m; ). ganglion r viole d s of , s (u s n e ( d r glands s e e agree mor e m CN A of e . homogeneousl r y purpure ar inne Th s p layers of an reac viole m; expulsio vacuole ( MDFs 8 vil inne CN B) cell of packe s e th e t to d S the l e r th e l a (C an no ). r e stainin r specime h s of larg ganglio a MDF of or s section S e t . r an abou ) peripheral y d MDF , capsul wit z . t 15 stainin whic posterio nudibranc Thi MDF is MD a as , d les foldin reac viole d Th th clearl e et s Th n betwee ± h ( vacuole covere appea MD e 30 Durfor s in s Laurillard s of e al t e g F s th h rounde s h n MDF n laye e hav t . s 2 m optica th of of cerebra g e particle y MDF ar of g F y stainin abou as no m ( mm up at r drawing particle e 1991 detectable n e r specime increase of mantl d e , ® r lobe th . in smalle wa h s to t viole successivel muscula th lle to no s 8 by ma e specime whic d ( genu t l s in diameter e s d (® : an 199 be l , seriall 2 vacuole 0.1 . g s ha nerve e natura y ® describe th bas gur t 183 mm wit an diameter s d glandula n gur which s densel r structures s h 6 stainin e correspon 5 s s is th no of d MDF ). neve e mm e ar h . T 1 ¯ n wit r no e s e y in 2A Eac at . ) yrinna pleura Th of section e capsul 4 no roun l y 8, ; similar-size , ar sectione an t , h separate ), y r th openin th som d bloo an mor ), limite . e g e s th s in h bee ) d packe r th e 1 bu e fo : cal laye MDF mentione d short givin d e vacuola cell ther nucleu Hypselodori larg e showin ( l d structur e e r e t d n s 0. A) e rhinophora vacuola gangli an Chromodori ar size d cel of to clustere r s ba 5 g observe d glan d e e d g e exten to (® . mm d s spicul r ont l th specime throughou specime : ar th weakl Th by d 5 th s gur ar g vacuola limit th r d e withi lie e e 100 oute a d e o e a e cel e e (® holotyp thic r d whic e e ` cluster s above d folde an ar MDF ® th d periphery singl statocyst gur 2B) m s in cell l peripher y lle (specime s r by n e in m. e l d k appear n ar n elegan staine s MDF a d h r e dorsa nerve fused d s layer large . livin e stain e no brito no fold cells fold 1153 2C t , s Thi wit ma s sub e an no cel ar th . . an of of ), d d h n y 8 1 g e e s s s s r s s s - t - - , . l , l , , i Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 equa th smalle variabl latera occu 1154 F surfac mor th or gland start ru (® to numbe be 10 th numbe denticle aberran 4A,B, inne projectin siz media nearl singl teet denticle row ia ig. n e e gure e n ± e to detected ® Digestiv Th h 15 s large teet e stomac latera r alon 3 s ve r e of l y th . . l in th E size n e as swolle r projectin (® denticle e inner-an It r r e 4H teet reachin e ). h s s t of e outermos th Tyrinn specimen salivar cus gur olde of of g r g in enter rhachidia a (® besid in d left wit Firs l e ). specimen th th h media wid h gur . p denticle e teet posterolatera oute shape e latera no n pe e e r , h Th system 3 an (® t a s s embedde e row digestiv e pharyn g e ). y stomach . r d ® g h lateral th nobili gur e d r an th 4B) tub ve 6 gland hal n th media outermos Ther s , t an e approximatel denticle l th e s n (® d eve cus e no e stomac s lateral to teeth s e f . d . principa bu e teet s 4E (® gur siz tw ( Th ro . whic x s n e (specime p e s tabl hav caecu si t Th gur n 3, d , of w laterall ). o e ar ar h glan x withi e (® e in . l bu cus withi s s e 4C ar 4 of larg e e e in Specime equa e h e Ther denticle specime to h gur t younge successivel li t ar M. lon 4A 65 2 m e an laterals a ventrall p become d l p th ). th n ). onl elongate si e n e e cusp (® n claw-lik (® d y cuticl ar e l ). to g y e x e th 4F Th Schro In specime no small gur th gur size 5 recentl or y tube e 20 In ar ® e oute 93 . s n e ). . n e r no rs th th ver 1 ; sam e e be e y Th increase specime s d e digestiv no row ¨ oute secon ), t in Th no e radul e . dl 4D) , s 5A) ® or narrowe bear y d r y e denticle lateral SE 1 rhachidia e ve . adjacen becomin irregula th e bulbou . n y e mediall an decrease s denticle 1 small appearanc postero-ventrally latera an r 1 M radul . e collecte ® no . d the to lac s a d lateral Specime rs Specime ha centr d micrograp n e s bi . lateral row t S. s y k no si 6 s . no gland up ® r latera s l a s t wit x r ar d V. inne ha y Specimen rhachidia leadin g teeth . in s e n to . s ti (fo s d s in an rodlets to denticle e 6, rathe 2 s of p h besid Mille wit teet toward s generall som e n materia th irregularl n mor . ther r r d l denticle shap eigh a of generall th s In no bearin compariso h e denticle no teet h bea g h ar paire e r e of oesophagus e . e a e n th th backward . ; reac thic t hal e 7 n olde s r e h ar s or 3 a ja centra s e . a e (® hook-shaped pair ha l y no Th usuall on teet w an e g ha fe th f specimen d caecu s (tabl les projectin k gure h y y becom smal row s s r w rodlets . thre e ar d s e or posteriorly s elongate hav abou elongate whil h s eac rows 1, stomac a outside wit n e l of bea claw-lik s e s nearl y toot larg m l onl a e s se 6, h e 4G, 2 no irregularl claw-lik h . e hav e int , r to t e singl , ). ® Th is sid scal 7 thre y bein s g mor e bu hal h ve tabl som denticle y o Teet H no h e si presen , an d d stomac visibl e ; e cus an t e reachin th e x especiall lie ). f thre e th to e rhachidian . oesophagu g an . d e e of ba in e rhachidia d th e h 2, cusp smal The e projectin s e p Th extremel 2 erect d 8 si r y digestiv denticle mediall e 47 rhachid ): e younge ar e siz a 5 4 t x (® s shape for e hav siz h to y at e 10 in in an singl s coul e l to oute gure g thre ; wit ver ma als y m e an an m ® d e th th th th th 74 m. ve of in d n d d o h d a y a 5 y y g y e e e e e e e e r s s s r s - Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 F ig. 4 . bar erect shape ia 5 no specime lateral m n Tyrinn . m; 5 toot 6, , 20 d ( saw-lik s scal E n h lateral (specime m a ) m. wit no elongate e nobilis ba . h e 2, s r oute projectin 5 Revisio wit n , scal 50 radula d no r h m rhachidian e lateral . m; oute ba 5 g n ), r ( r D) variation media r 5 of scal s denticle 10 an irregula th e s m d e n ba m; (specime rudimentar nudibranc cus r : s (C 5 ( r A) p (specime 10 rhachidia ) (specime aberran claw-lik m n m; no y h ( . B) n outermos 7) genu n n t e no , rhachidian centra no rhachidia toot scal . . s 7) 1) e h T , ba l , t of yrinna scal scal radula teet r specime 5 n s e teet e 20 h an ba ba (specime m r d h m; r r portio 5 5 an ® n rs ( 50 10 no d F t denticulat ) m . m latera n n m; larg m; 6, no of scal (G ( e . l H specime 7) rhachid teet ) e ) , hook- ba e 1155 more scal h ® r rs 5 of n e - t Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 1156 th withou and ( insertio 3, portio to Th th portio whic T. va oviduc whic hermaphroditi ined stomac righ borde whic granules syste a F 195 ig. e e s wid 40 e 4 nobilis Reproductiv ejaculator t peni deferens ; 5 9 Muniaõ h h h tota m an . mm edg an r ). e n n pharynx Abbreviations ( is t reache t become h B) n of d of in Tyrinn Th caecu is s . immediatel cuticula l prostatic outlin e to of Th (specime ar 7 specime lengt specimen sheathe and th th backward e  n th e (tabl . e va th e e s m ; y Th a everted et e c stomac s up latte e e digestiv pi s h e T. nobilis portio anu narrowe r duc is al whic deferen vagina , e system n e of , proxima d to structures . n 2 pusae morpholog give no : th r y s s ( ). th by t ca no s 0. 199 specime e no , in . , h h enter n passe , e outlin to e 3, Th 5 th va n . , a . dorsall caecum ar . as a s va mm 6 6 gland r scal e formin 1, in s by th muscula l e doe ). ). e narro Th s mor s intestine a deferen latte 2, e s M. ® va e Th bot Th e . deferen int straigh n Marcu in gur of ana s e int In 6 ba y , s ; y e e no wa e reproductiv o r Schro h w wherea dg diameter th an g r of deferen o passe e va distally vestibula specimen l formin 5 e th t r partl , s a loop 6. s d ; papilla a s 2 digestiv digestiv th ope layer s s sg t e ® distall slende ¨ mm. deferen then Th lle dl banana-lik duc e measure , ( femal s . 195 y s salivar n va int d s e an g . it . . t separatel visibl , gona , enter s wit Th r Mor Th e y e a 9 o s is d r closel wit e or e system ): deferen wall gland s become no long unarme th limite S. syste e e h y glan extend s h it afte d e e e glands . proxima s intestin a V. fro s e y ma wide or 6 distall cover dorsall , th ; dar m : ar d y r surrounde di Mille d ¯ an an m ( e s y withou agellifor A) s e , as mas d a is to s ; d k agree , d commo narrower dista abou be sof considerabl st s peni short e simila longitudinall y specime suspecte th brow 7, l somewha , n y th s ma th gonoduc t curve stomach e l e (® afte part an s s righ t t e intestine , digestiv y m gur 12 wit n d r convolutions as va n d rathe r exten n in d s by . mas t tub s longitudinall mm vestibul d h e indicate no it Thi portio of backward . deferen al t 5B) s by t y th a . e r l ; s ¯ is junctio 1, th d e int y s muscula (specime e oe specimen attene (® Rudma straigh containin . gland strongl ver e folde scal to description , o n gur Th e vestibul oesophagus d th s in y th e nex , e e by passe n e d s ba short d specimen . e follo 1C t intestina y lef fro r n Th y wit r n intestin t ampull anterio s Marcu portio 5 sheath folded g curve ( t ). to no exam m e e s 198 2 h . bod w san mm; int s thi Th Th an . th th th ; it 4 2 of p, d o d n n d y a e e e e e e s s s r ) ) - , . l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 bot homogeneou di easil chromodoridi no (® found characters geographica an ma stainin trace mal long whitis is receptacle insert th inseminatio ( branche contracte curve vagin F 199 ig. Œ gur e divide y t y eren h Remarks. e vagin 6 y , hav othe 6 s or d . e a claw-lik s an ). h , ha rs glan Abbreviations of lea g 4F is on is t distally , portio d ma d Tyrinn e d s Afte r tubule receptaculu glandula d species d aberran a s ). a vestibula d inne . femal a specimen int y th In muscula n li divide to mass l lon In conspicuousl s e r no e e o d duc rang particular a n r teeth clos th . an s mos Th ova g th contras t e nobilis specie . havin denticles ; Thi e withi t hav t an d genita e s hd However r e e du e l whic : assumptio r t widel . structure m . r mediall specimen int d s , (tabl am to translucen Revisio glan e Specime e of ® hermaphroditi , s g seminis coile n narro o a bres reproductiv to t , (e.g eac h l , a th glandula to it y th ampulla specime . d aperture e y it enter mor s e d convolute h , Sinc 1 e , y s . surroundin othe swolle w n ) formin bod length s ; rhachidia onl Bertsch s dista elongate other shor n n v, e attache an of duc examine s t e no tha granula y r vagina ; y d th burs n th e radula r t . at l . specimens t n th , wal c , no e syste t sho function e Histologica g stal 2, portio , (onl thei duct , bul e th d nidamenta th nudibranc commo a . a d d 1977 l as n ; smalle 1, w g e d yellowis k e bul m to va of b r rhachidian r copulatri y r teet ; th durin whic wel vagin specimen of considerabl appearance stalk ni n , (specime character 50 an th ; b specime e vagina , . h n Rudman is , l th ± e d vagin wit nidamenta r Thi h 10 th as bearin atrium g atriu s e a surrounde l open specimen h h is l e h thi 0 ar section rounde s th x duc give l portio genu th n m ® a a e duct s structur s s n e m m as rs s no e s ® ; wit stud crossed as studie g t holotyp no . , t e northernmos s ar lamentou bc or separatel in . a s Th a ; fa 198 l lateral di 1) ris mentione h s e . n vd , d duct larg T s r y to shor diameter Œ d 9 of . e burs a widel of yrinna e , 4 no distally receptaculu d wer e erence scal nidamenta muscula nea ), by th paire . ; e ha specime to e th belon . s t pr Th a e projectin thes e e of 3, s e y r duct s connectiv y of , copulatrix vagina th ba collecte e no prostati d mucu d surface th s belo use Tyrinn 4, . e ) r e specime femal t g projectin r regardin e by 5 t Mor . whic Chilea di somewha n va 5 d to bee 1 genita Bot w Œ m l . no mm. Munia s s posses g to erence d duc Small e c e tw n . deferens a th gland h media e ; . h semini fro va Ther proximally glan n pusa n detecte fgm distinguis e o 9 is tissu t l allosper g s g specime m no commo sho or apertur is strongl s deferens , õ  t s radula , media n , d e poorl e n a . rathe rathe femal migh an e s wide . mor w et is mas 1157 have 1 wid cus d an an d do no no al in m n h p n n d d y a y e e e r e r r r s t . , ; Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 ( 21 Tyrinn thos singl th cus 1158 surfac show examinatio of T. as from denticl transitiona assigned However denticulat specimens an ( rang chromodoridi of thi Ther broa appear ( no hypothesi position hav an th bot specime by teeth 6 on blasts specime by du 195 192 196 (® e e d d . e s nobilis p Marcu th th completel h an deletio The T. Anothe Regardin e gur aberrant Argentinia 1, to e 9 6 4 e e d shap e teet stud , s withi falle e e ® . T. ) ) ), d havin e bu of projectin a e tooth tha rs is s it intraspeci ® Thi nobilis e an considere at centra n n . indicatin 23 di nobili ft h reductio tha t nobilis t 4C , to y ma variatio stil e n However s no th d e h s t . no th lateral n Œ n lac he s ) pe l r n (® it ou th an ( eren g t Marcu e Munia lef Tyrinn ). , e latera l thei stage . y , an woul as 195 . distinguishin g of y r k at gur an a e 1 stomac d eve s insertio als l larg littl d t Th 1 t radula als n d wa hal Bergh stomac denticle coul g embedde in singl th wel is radul species as t d leas 9 r closel o s th ® n e d g specime n e n o fusio media s d ). e e shap individua e c s als l specime th f ma 4F s , a õ e wel ca note latte a th  l digestiv an of n seem be dat variatio be d centra Th t teet ro ther e e completel as nobili o ( hig h n r result e a , n et th y ). als d 195 y projectin positio l w th n e 1898 h characters e . th th a fusio is analogou s be of d r hav as portio e n h Du fo no h al. e resembl s of e o d is lac of throug e e toot n on of visibl ® 9 di g cusp th r n is l intra-individua possibl s on be an specimen state rs ) s followin alway resul e e withi . e no , ( k th l Œ n essentiall toot odontoblasts featur n no a M. e T. 199 t no collecte system Th th sa d radula of to eren n falle h du as of e th intestine . n of tw y specime . . e e w g of d position 6 T. ® e pusae Schro h t th e 1, h Th clea e 6 e wa n dorsall Rudma wel ar s o s rs centra eviscerate mechanism , centra (tabl e t ) n materia thir ma of to al speci e e somewha th th th to wit embedde e t righ e th (se degree s pusae g tha . e out s broa l l whic r d e e e individua latera ¨ Re-examinatio th suggeste presenc y y rathe e latera th d develope (thi th assigne h dl asymmetr e specimen stomach fro e n secon . ma t e di t eithe ® , l e (o y digestiv l th e In s tabl 2 n c an lateral th d bein of h teet Œ fusio cus : s m radula ar ( ). l r absenc l y e s wit Bergh eren r l specimen di bas e ( sinc examine study d Marcu l an d fourth remainin e d t of e 1984 Chilo r be Tyrinn withi d p is e commo s h Œ rhachidia d g surrounde S. resemble d h 2 d hav withi an n erence e l lateral d wa of leadin in broa e . t denticl ). s replace to fo e du s aberration intraspeci V. y r an mos In of d ; trait e , inne ; of dissecte th Th n r s cadlinid e claw-lik e rows glan ) 1898 of Marcus s  e T. of 199 d a th Mille n e als Islan thu righ T. d centra th a d no ( s n e to n t g s larg 195 g s th s r th evelina e denticl nobilis assigne durin oute e claw-lik of e an o highe d 5 pusae s . d lateral ® n ; . denticle s in specimen betwee of amon to e e th t materia reductio ) d 6 rs describe Marcus Anothe th d d 9 n Marcus by d e ( digestiv an centra latera is e e th t ® ) indicat s , Marcus l by ma othe r ( aberran s ha e her siz c g Bergh latera 195 an use to d rhachidian r large e of denticle e th e usua no and d th g s variability s thi y normal numbe e qualit e on n e d by e lackin be a r correspondin s to d 9 l e t be of lackin individual l l ® appearanc s r , secon n s teet ). ’ latera on T. d e bi specimen e stomac r e t teet to l studie l appropriat 195 , T. T. specimens explanatio stud Collie , ver assigne th wel fuse occupyin t fo withi ® 1898 gland of Th ® bod tha 195 y al d s nobilis rs h separat radula e nobilis r g , pusae h r 9 y g e l th l d on an t denticle-bearin projectin centra of y of l ). th d t s ® to y int unlikely ® d 9 : rhachidian lateral . n r teeth e laterals rs show h denticulation , togethe d rs int e Th pl deformation ) specime radula d siz th ma reachin o s an suppor ® t hi s is t holotyp e . quantit wherea r to by rs o by e th e e g possessing lateral e examined 32 , of lateral s g d e n l or describe y teet . however t holotyp s on a ® T. e T. materia toot of s odonto , Marcus Marcus Odhne is Farme als rs lateral fo g , furthe know severa centra . ® bea r in r e h bu pusae pusae t gure Sinc give n t cusp r thes g row o wit ver y s h s e an s s thi siz bu no th r t re be of of of of of of in it d h d n n a y g e e e e e r s r s s r s s - - t l , , . l . l . Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 size stomac tissu be th ( externall 199503 time positio ( presenc surrounde Muniaõ ® recentl examine T. glan Thi by th is an centra connectin especiall T. be no dissected comparable Thu additionally th durin no duc det up is cernin ( genera 195 195 195 ed available e e e describe d . t limite . pusae presen Berg nobilis s to Th Th Anothe Th It t , rang 2; righ s bod d , othe 9 be 9 9 e as Marcu (abou nutritiona synonym g spicule ) th ) ) l g dorsally is e e e tabl 60 coverin l y n  8 h or n coincided e thi a T radula measure observatio h e t th genita e y holotyp d y poorl ) collecte d y . r withi , conclude mm an of well-develope . t of g of va et ( collecte Munia s e d Durin eve d digestiv e typ to siz and regardin t 189 , examined r nobilis in s ; d lengt th stud , as 2 al spicules s 20 by morphologica th s surprisin al th e indicate th ) n th e y deferen n l g caecu e ( th e l y als . 8 reachin e . as mm) Marcus specimen l e e ( a a structur T. materia Th e ) agre describe th d extremes e th d e is tabl y g stag intestin ( righ h õ , o  sam shor an bul n d 1996 wa d centra of specimens e e holotyp e range Marcu by thi n in exactly e an pusae ma g m et features , fro mantl d , e e of tha vagin materia e s . b t Revisio T. althoug s s e d s accordanc whic t g radula Muniaõ 2 Marcu wel g al. portio y describe : is an Thi , m of tha Munia stud mus mus va ). d th l t e ® 40 s nobilis e l d hav ® s 195 highl . of d ndin ( s caecu e l of th by s gur . th e ar contracte betwee duc a Th e insertion s h t 199 holotyp l mm Eve wit ( arti positio deferen t t Th y tissue variatio e . e e thi 195 of 9 e T di entire l . r ha n materia n h be e s Therefore be  e Berg n ). considere Dissectio t dissecte no specimen 6 g bee õ Chilea . y Œ h  (specime e n n characters of d ( th s ® 5B T. d m in va ). is is 9 er nobilis et 195 e Re-examinatio withi cia al suppose stretche variable et partl lengt fo n e specimen ) no . n th suc An s wit s ) length immatur reache , l pusae h mor n lengt e al sa As r s al l . 12 9 d dissolve othe e fro n deferen t to yellowis of l . contractio ma of T. A ( n w h . ), h n nudibranc explanatio bee originall y d muscl 189 h assigne ( ( know e an m coas 199 199 n th glandula bot th and , be d s speci caecu n specimen nobilis. h dissecte durin Dori y ; r of , d unti d ; or s n d of to , e 8 it e studie no an exten of th specimen it to s th 6 h 6 ) fo e th va detecte proposa bu 40 s th t e is d belon museu e les d T. ) . n ) h th s ® no ma l an e ( specie r e m d at 7; g tha be ® s is e c Bergh (se t tha show mor origina mm no y fo delicat e surfac s Marcus specime bre n d deferens . d n pusae thi to highl d On d di tabl y ther va describe in h r convolute Chilo n r highl e loope measure t t 1 s int du t g of Œ Marcu specime be her othe structure s T. genu s e d s ma T. Schr freel of m examine th s an erence to an s s th l e e stud an informatio o , e deferen earlie a y ar e , nobilis a of regardin e th in materia th e l 1898 2 nobilis y d e y to d th e th d  d ar no r ’ smalle e of ) n sam y description s o coile ¯ Abraham e be e othe an . È is Munia Islan e convolute data e materia 6, y as d dl uence connectiv e ® considere s withi T larges Th visibl r ` d measure s sam th lef vestibula n xatio r d show fro ; no d , thei yrinna ( alway withi wa e havin betwee s d an 195 by thi 1997b, d s e e ha r ha an t th . and t d positio l wer r m durin g hand va e bod digestiv n s d in As d r ( dista e s s e strictl s d õ n t specime n  Berg ( species 9 MAC propose n s external T. lo a l th s n Abraham th specime study) bu a individuall ) hardene e materia s on g , specimen (® tha preserve et T. deferen ramifyin see y s w d n e d e th s e nobilis 187 ar muc c detecte , r transitiona t th g l gure portio ). h n 20 result tissue al furthe a numbe bot of to y e muscula t pusae as covere e n gland vagin e N thi e ( . th 7 longe als . eigh correlate . mm heterogeneous in h 189 n be Encompassin mos ( T. glan d h , s 3387 a s e Althoug ( n 1996 l d o shorte s reproductiv BMNH of and fo 5A,B) th s d . specie stud d , siz n of assigne g r whol s conspeci 8 ’ t part a nobilis of by appear d r y t ar 187 r r e fo specimen d in ), (specime vestibula draw in dissected examine T. e 3 T. an r T. whic on aberran ). by by T. e materia T. r ) l lyin y Marcus Marcu of sheath s length 7 th d e d r hardl identi nobilis nobilis a . s pusae e stage is coul pusae ) pusae h som bod , mal e bot 1159 n dee d con lon and wit ma g h s wa and no th th ® re by to to to c. is d h p n h d y g g y y e e e e e s s r s s - - - t . , , , . l . , Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 anus abou wit tw Th ( 9 1160 vers tentacle for Peruvia rounde shin of dot bear an an detected livin wit Carlo Rubio 19990005 by Bahõ BMNH Tyrinn Cadlin is T. synonyms pusae T. an tissu in fro studied Chromodori doub in 3 mm g proposed o d d d mm large nobilis nobilis e th m m h h s Externa Materia 1967 F Sandr e e al  s g e 7 width als to notu . a ollowin in e t smoot t orange s a th fold a Thi a l throug som Marcus mm in Peruvia , ) wer de 15 thei d lectotype . o Paredes evelina thre r th externa n s , evelina : e circl bu . sout Th widt 168 orang by Jamaica m ar ; th lo s instea e examine specime a Jua s Th leaves e e s r of in th t holotyp h Gul s e l e e l . e du s scattere of is alread ± ± slightl Millen e g conspeci h contracte h e e Angeles deepl morphology examined anatomica re 169 h n edge row e width ora Tyrinn , e n juvenc rathe th e posteriorl othe : 10 specimen e Marcus 1959 th l . f d of d an Fern Rudma . specimen . to e marking ; character specimens l e of s . dot Ther n Thompso an Large s d d ne y y tentacles y Tumbes e Tyrinn , translucen of r . contraction r d aroun , an it (se 6 a groove elevated a mentione a coincide d w Th in s Han , an  ® smooth s nde bosses mm , large d Gul d delicat ty e 6 . al replacemen e whic n 1958 cod d th e l , s mm y ar . 14 tabl tail l 1984 . s z feature s Chromodori ; nota a margina e Cadlin d Th Tyrinn ove fro f s e in complete indicatin n e in , , . Islands , d : h of evelina in Universida Bertsch th , Th M. . . an Alon 18 e si of 1980 empt , describe , e s Peru a : m longitudinally heigh Thes an t Th preserve r d e smoot x 2 externall no l California 244 wit th ± d sam e ( . nomenclature notum ). th surfac rinophore 21 Schro Abraham in d bi th a s e e Anteriorl a specimen t : collectin g Consequently h e ± y , ; - 76 of l e t bearin e preserve large e digitiform , evelina 246 evelina t e th Collie th , : th notu (® g . by subepiderma rock to d MDF ar lo h ± Gul ¨ is Tw e It C. d Mik th s e 77 e dl gure e ; by d . w e sheaths th tripinnat d juvenc origina y nota Munia as specimen tru r e ha Th ; is conditio o long f juvenca m y an r mammilla , g e Naciona Edmund positio specime e agre e , g th e an s s Mexico y slightl hav s s s bein specimens senio e e papillae d coast d (® 187 ( . an det e (` locality a l 7± fro d Miller th fo Marcus rhinophore , , Th Peruvia S. a edg gure whit , õ  Farme ana . e e narro n d l digitifor e r longitudinall 11 retentio g 7 . Bergh m Th , r V. e specie e et preserve ). n th , perfectl y th foo Thompson ® n s l short-conica e Tyrinn s ) bu n Peruvia l l , gill s e t th of e examine glands 20 al e folde . 1981 e ther . Mille no e 5 Mayo papilla fro int , 7A, w r t t rhinophore sacs . gil e , Alon Tw lectotyp ar m , larg n s an disregarde , 1964 1996 corner 195 s Septembe foo Gul ZS o trace 1898 m l n m e in e specime : B depth d d a descriptio o y cavity s ’ 176 n th ar d marke of ) e . ). M th r n 8 g nobili : an a t in : an thu f . subepiderma Ther specimen e wit e shinin d 380 ) length de 265 , ( specime th e s ± Ther y th , semicircl of Th d s 2. no abou rang preserve d ar 179 durin e of , Gulf e 1980 s l h e groove ± ar 5 Sa . ± longitudinall 29 e (an th e n s d Californi s in 381 nota e 273 s ther commonl Th mm ; d orang r e al e th Bergh n 19990006 g e lon bea by ha e 1982 s ar t Jun synony n th . d elongated l g ar 1974 : ; e e . Marcos gil of tw of throug n e Si Marcu regarde e s s l thi Thre by e paralectotype rhinophore g smal e in r e d othe d e elevate borde ha 13 l x : o is variatio e ( (on 12 18 no digitifor anterio 517 , s cavit 1996 specimen los , width) ora bipinnat Berg l irregula s colou 1898 mm a no stud gland m e l collecte y leave inner r e s an dat t . h , orange ar , l ) an specimens use d r T. y specimen , 19990007 y Lima) ZS reaso , d h d th tentacles e specime y , in ther collecte as a d r groove th n ar r , Tyrinna delicata d sheath s s ( regard e 11 whitis , m M Marcu ca on a r 189 lengt to e s junio of whic trans e e in s nam nota d e row mm n bea lon ± n foo ora gill sur ) an , re no th ar th th th 8 by be El of to ), d d n h d h h d g e e e e e e s r s s s r s - t - - . , l . , , l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 F th to specime betwee formations dissolve Histologica mantl the edg F ig. ig. e 0. e Ther Mantl notum. Jamaica 7 8 6 is . . e notu wer mm n bilabiate n d tissue Tyrinn Tyrinn e 0. e e spicules ar m , ( ) 2 orange l structures BMNH n in thos e section tissue an , a a specime tw al bu evelina evelina d , l e o ± ; th specimen t , 0. re of thus irregula scal , thi s e 3 d Revisio . T. do mm e uppe in e th n s e , , , large Ther histologica ba life e (® ma if no evelinae . r gur Gul r r spicule ; s Inne t 5 y r ( n li examined e submargina sho B) Jamaica 5 specime p e be of f ar mm. 8 is r ventra specimen w e ) th du s l MDF no no det resemble obviou sectio e wer e n , t nudibranc l . scal spicule . to notched fro view e Th s Thompson n l present m e aci s of of s e row , ba th s gland empt up whit d MDF s th r e tha s 5 . withi ® Gul h e to Th xatio e 200 of , t y genu seriall s the MDF s f describe , of space e 0. spherica of n of m y tai 7 m. 198 n th th th s California mm wer s y l e of T e e s ar 0 is oute yrinna smal sectioned nota ) indicatin . e e d th lon l shinin Th reac rar fo e MDF r l l g r museu e ban e specimen : bosse h an ( th an MD g A) g e , d d throug s diameter d dorsa smal th m smal hav (mantl project s F possibl e no specime structur s e position h l l r l a examined view th T. Jamaica in e diamete s s y e derma behin deepe nobilis ventra n of small , 1161 spot fro e s up of of m n d r r s . . l l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 muscula an cell than centra specime 1162 F shape ha oute large Th inne formin ape rhachidia in ventrall on ganglia stalked rhinophora ig. d s e e th s x Digestiv Centra r 9 r anterio extend no following wit r e . in an d an l th California) specimen) specime denticle specime g Peruvia . . y vacuol n Tyrinn lateral h d e T. r d A In inne a bear n Gul capsule viole no l oute bear statocys s nobilis projectin l th teet e r nervou . somewha r gangli system to f n e s a s 8: , e s n n r of s t h mostl large fro , denticle hook-shape evelina besid th ( stainin denticle onl , fro on scal specimen MDF 15 ar California) . . scal m e s Ther Th t a e e ± y m . CN system r g e th y wit 30 e generall to e separate specime hav e t e ba e th bi centra Th s , g ba s th toward e m S vacuola h Gul s thre r SE ® e ar particles m r an e is e 5 besid d e an severa , 5 Gul . e M , 10 ora mai oute f ja d in 78 no M. d d 10 e scal ® l of y w n d Th m lateral micrograp lle f Ö on pair diameter) up cusp s m l e m; smalle natura n California) fro Schro rodlet r th m; r fro tub e l th d 86.1.8 th e e . cell cus otoconi ba denticles e (C e s cerebra to m In wit e abov m . ( e ® r interior D) s of mai ¨ ) p s Th 5 rs is th dl r large s h l th thre onl centra 6 appea 50 t (® (® tha openin irregular e covere h hook-shape e e e vacuol an n . righ in Gul gure9A) gur a m y of th , Th innermos l cerebra e cus n r m. . d scal nestle an th hav . e MDFs l cuticula Thei th e r oute e t S. f Al p oesophagus radul e d d g 9C latera peripherall e ther vacuole e e e V. in l large by ba , pleura to laterals cell MDF r s r up ). l ridge-lik . d th r Mille e a betwee numbe Th , a th ganglia 5 t r denticles In l s ar latera thi e ( r thic to structures 5 e latera large ha e apparentl Peruvia s e l m specime s s mantl contrast radul n m; . appea tw s thre gangli ar laye k It . y l n r up r e e o . cuticl ( teet l surrounde specime B) cerebropleura decrease ha envelope e Th . latera smal r toot a e to n r : oute h Th appear a s formul n centra surface e ( y , mor specime a e thre ar ( A) fro h lens-eye e l large th containin whic mediall l e bloo r n ha e following denticle ja e m s e l fused denticles fro d s w a ® radul densel r d s . successivel inne h slightl rs th by d is specime m n rodlet by tw laterall t e glan s 87 y (® l . a th o a lef r Gulf ar an a Ther g elongate s Ö gur y ver ( e an or y t Peruvia e , laye s . withou d a d 88.1.88 packe Gul n d folde latera In ( short hook e y lobes y large . peda large e thre fro 9B) y fou thi Th r an f ar th of to of m d n d d d n e e e e r r - - t , . l , l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 folde bulbou Th in denticles erect on F and ( ( bul to Th nidamenta duct genita alread fro Thes dista armatur straight sheathe rathe Marcu ® backward Marcu thi gur ig. ® e e e m th b Reproductiv Geographi s gur posterolatera 10 . receptaculu e , e l stomac d study Guaruja e r at Jamaica l genitali th y It 11 pharynx Abbreviations Peruvia . gland s intestin s burs shor e opening d , s th e e present ( . caecu ) passin 10 Tyrinn divide by an 195 s outermos An ha e ar ) l . to a t glan junctio to h d s In e s a  8 n a prostati c wit ; ; wer outlin Marcus m lie simila th e bee muscula ). Baj nea pi specimen g a s th distribution an e al s , m d leave e s , h int ma Afte evelina l wer int bu e e l d n system an proxima a ana specimen mediall r semini denticl n : a ampulla t no o open Californi t e o detected y r ca d e teet Revisio Santos of s c shor r a ther of , l or t an a , a no r th e tubula it papill , portion 1967 detecte caecum th , duc layer . h d scal th s s ma straigh e s t outlin e l t e y e at . ar is junctio digestiv ye agre e intestine belo , Th stalk , s accessor t wer e y or ; n . e digestiv oval mal a t th a runnin examined . Tyrinn ba Brazi thi r A no reduce e of ; developed Th d (Collie e . e adjacen e to papill e w dg r t . s t e withi pai It of apertur genita wel 5 , th e onl n inseminatio Th th , reac th e study) somewha an ; 1 l dista e digestiv passe th r st g wit y e a gland mm. l e e y d e d nudibranc a ( , e r to of n wit mai evelina glands syste Marcus h left stomach t whic , in nidamenta an femal l trace h digestiv th l ther it e to th . larg , s h system portio d th size n s , , of Th e e as e int m th run th dorsa embedde Farmer h e cus t stil gland s rounded e e . e e th e , e smalle e protrude wel n ampulla o of is , of e . ha duct irregula dendriti s vagin l e p origina dorsa h n s 195 syste a on duc alon juvenile commo th l (® s l ; femal genu of l of di strongl surface as bee e e , s gur duc 8 r t , 196 th d a pai m Peruvia , wit l g ), tha dista fro almos whic , th n surfac is l r s e s e c withi it e t (salivar th r 4 descriptio h Jamaic int in reporte e T 9D) s va , n n m a accessor possesse gland of ) y . e seriall yrinna anterio l allosper mal th specimen h shap an long s o Th intestine Atlanti va curve n t tube-lik . deferen n e th e open d sessil Oute e y e th s of burs specime s e a th , d e broad y an gland deferen e vestibule an thi r y an th e s fro d ( sectione n m s digestiv d e r borde Thompson c a ; Gul d a gland e n d s portio e s of burs lateral oe distall m femal an receptacle waters s digestiv is distall an th salivar bea , examine , n omitte f T. norther longitudinall d muscula e s oesophagus d of a r is s insert r . begin atriu d convolute e evelinae e n an copulatrix y s No cover up Californi illustrate : y vestibule specime gland ar y whic d d int e Ilhabel swolle , ) glands to e m genita d n gland s curve s s o of 1980 r mor s clos 1163 wer Per as fou h (se an an . th th ; th by A p, is d u n d d n d a y a a e e e e e e e s r . ; . . . l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 1164 receptacle to as thi entir tha teet withi Collie no ora th Jamaica bu describe wer groove other questionin ( an T. externall wa Edmunds F 196 ig. e d t s s evelinae describe compar n eviden l Edmund Apparent Remarks. h e BMN gav study 11 7 liste e th tentacles n describe by ). r morpholog specime known vestibula rs glan Abbreviations specime . e , a an e , . d Therefore Thompso Gul d y speci singl Tyrinn Re-examinatio receptaculu , no : t H s d d specimen g by e an th 198 d di by agree mass s Farme f th con wit e detaile Œ herein d n . d e ® Marcu di of ( n T. r Marcu e erence 1 CNS c 198 a specime fro Th as anatomicall gland Œ ) of h ® conspeci ; separation Californi evelina s y erence rm evelinae an hd n , m e havin thos 1 wit Thompson r : of the s . d , m specimen ) ( , d s th s ( am . s 198 Th mantl fro s hermaphroditi 196 note h informatio th ( seminis e Ghan hi y betwee e ( e n 197 , s 195 othe Gul , e g e n m s ar 0 ampulla ® of of 4 reproductiv in reproductiv digitiform ) a presenc of d whic ty specimen ) e 7 e Ghan 8 f . y pointe T. M. by ). a describe r matur tentacl identi sligh gland Thompson ) of ; In of s n agre ’ T. an ( v, s fro evelinae h Collie Edmunds California Schro th specimen Jamaica orde ; d n a vagina evelinae t hav m e e at e d ® s e c subsequen on whic colou e ed of specimen , s an e , wel Per ou duct ¨ d specimens r e shap commo r dl bu to syste e d digitiform th to as . considerabl t ; lon an l u syste h ’ Edmund t an n digestiv s va ; tha e r , be , wit s an reconstruc T. ungroove d bu ni possesse singl e m scal materia g di tentacl , 1981 d assigne , Farme t wa vagina d n cause t h Œ m evelinae ( ora S. s nidamenta t specimen e lef erence vestibula th atrium fro e th one s . ba wa , V. t l e e undissecte , th Th s e 198 vestibula e r m tentacle d Gul syste l l longitudinall d d s regarde e r s 5 Mille e origina d shap ha duct e of immatur t by ( Ghana variatio ; s 1 longitudinall to . 2 mai 196 ora f arrangemen s majo bc mm. ). withi l m s materia r of bee T. ; intraspeci n , e duct of A n gland l 4 vd agre burs r s of Californi d tentacles ) l evelinae d n reaso r glan , , n recor T. wit an description di ; n Jamaica Jamaic muscula seriall orga e hi pr a hi e Œ d s of evelinae l h an s d y , wit erence copulatrix s fro ar n Marcu d tw prostati omitte a groove d radula y n ® materia t e . fo h y fro c . m a o dee groove therefor systems a In of absent Edmund r othe r sectione n variatio describe specimen an Baj s m va Ortea specime p contras d th s s fro in c r s d d ) Puert ; an r longitudina of a e l va deferens teet of th th (` m fgm d T. . bu Californi , allosper d enrolled s e Sinc e ’ Brazilia e th tentacle d s th n s Jamaic d deferens h evelinae t di Marcu radula , Paci o t e n s ( ( durin e herei rathe foun femal  shap 198 198 to large e fro fro Ric onl ; cult th ® vg al 1 8 m m m c. ’ d n n o g y a a e e e r r s s r ) ) ) l l , ; Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 bution Ghana. an relevan regio reproductiv press) carrie Argentin specie Ghan an of graphi portio to glan 1898 an roundin T. an T. lected In nota in whit ine borde livin coloratio Behrens an golde followin Tyrinna be Specie Discussion durin withi divergenc d d d d T. th d synonym evelinae contras evelinae muscula amphiatlanti As Edmund Tyrinn d Marcus e g l ; Farmer th n n e durin require n g d a , bump r Marcus evelinae . s line n specimen s coverin c , easter bot a e is t at ( or Th ethano ( suc by g , comparisio whic g of distribution Marcus by morphologica Edmunds a n resul norther no e s 1991 leas th a h e g t orange characters alon on th h th of r e s Rudma rate specimen examine t , e s to , tropica s nobili s thi occurrenc n h whic as e 1967 e , ye t ® g ( organ of 1964 t commo th ( th ; l 1959 198 Paci con vagin bre th forme s show Marcus of Humbold g s s th t stead , c e preserve thi e T. n stud of e developed 1959 ± in th h s e rhinophores ; 2 distributio s thi , ® , re Peruvia n las ® s an n ; thi l s vagina nobilis ) ar e and rmatio 1981 T. : Marcu d an c thi d s a Paci certai Revisio : s discusse s y r y ( study) n ar T. t d e Patagonia s ; fro 198 e instea anatomicall l spot al an study d (e.g gen descriptions nobilis , 1. al s no Munia study) vestibul e feature souther of l 1958 evelinae T. study) d , 5 ® l t m connectiv no 4 t . c 198 livin ’ n n e extende ha . n millio s current s ), materia , visibl Cadlin T. ; in evelinae an norther ¯ Marcus , n t d specime distribute thi n an nudibranch d d ; usin is ow unfortunatel 2 T. . . hav full õ of of d T.  thi g s n nobilis . possibilitie withi whit ). In Macroscopica d e s simila n n Th whic Atlanti n e delicata et T. is th vi occur a ( s nobilis y In Marcus g e a contrast , d Paci Marcus coas y macroscopicall years l a , e e study) e vestibula certainl e al orang n develope to evelina , n adequat evelinae posses n calcareou ` contrast Magellani see of driftin h nudibranc unequivoca tissu ., line 1958 Per r d (as tropica ® t s argu norther examine in 1996 c , m c T. on s , to withi emphasize sinc u s water , e . s e , T. specimen shap y ; Chromodori , ma e alon separate 1967 y s Tyrinn (thi to g leadin e 1958 nobilis souther Tyrinn th thi ; surroundin r Marcus e d ther e on du , agains juvenc larvae ar l thi l n y e gland re s n th T. s nota e s s c and material g water d h e e e ; war entir explai d spicule ; s whic study an e study Chil e l th distributio Thompson to genu pai y g durin nobilis Collie a describe study . presence spot closin a , is d d a d ( , n l to s e t m . nobili Th it , nobili Munia bosse e especially or r th s e h by anatomy nota and wit s Chil ecologicall 1958 no ) n a ) s s temperat notu of s of e e (se wer s g immatur g adult . ) r to ma tropica T exclusivel g juvenc th conspeci Munia h wer appear an absenc s thi an larg yrinna s Al e informatio e th l th T. s of d , e e Baj ’ y õ m thos onl lack borde  or d n Schr whic of n and e d e l s , isolate as e s th presenc pusae be e et livin ( 1980 Caribbean Farmer , a study , distal mammilla carrie a y a matur rangin foun Marcus e l õ e s e bu )  dendriti , whitis th n s e e al California ` y o amphiatlanti du possesse h Isthmu ramifyin È suc ® r an at du fro dl to state ., y e et g t of ty an wa ; as d e d ar , . di , d 1996 aroun specimen whol e d h e Edmunds Jua e n m al be fro of d g to 1996 In s e in strongl spicule Œ , h tropica wel by to gland . , of . er transferre 1964 specimen on considere by fro c th s Jamaic m Ther ( 1958 n a aci wit recentl e ; , contrast s 199 e s col boats l of g gland e thi T. d ; ar col Brazi Fern eac , m ver du as thei d th vestibula vestibula Brazilian h 1997a s , th an Panama e d 6 s s y ; e l e evelinae d s ® e c Marcu ( media y presen h Vald ) yellow study) e , Collie Bergh water d water ar xatio withi . r a exam distri coile to a y s wate refer livin l 1981 othe  smal nota Slo 1165 ndez s wit e geo , d d sur an an col ; th no e al  of to to in w s, n d d h d n n g e s r r s r s r s r t - - - - - . . l ; , , , l l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 of shap it Tyrinna thes Island 1166 gland stil It Rudman and tubercle specime wit prominen a an an in by Multicellular yello ( shor in presenc ( contrast withou diagnosi pe agree to sagamiensis comparisio et Ortea and otherwis of nee size aperture NSM Baba s pai r d d is length length Tyrinna l al h submargina Bab T. d long Additionally Cadlin Elio Possessin Typ Re-examinatio d e , hal t longe irregularl unknow e w ., T. C. r subepiderma however to s externa , triangula Paci , with s T of of burnayi t 199 shoul 198 a f e e 1937 t spherules n , s ( , evelinae japonica s s be e by no ’ , ) spicules Bergh of ro ( lateral ( materia s radula digitiform t an r th (Ortea as a ® 26 remain 191 194 whic of 6 8 n ( tubercles by c . tha T. w con ¯ Muniaõ e 198 ). ) Baba burnay ; d d n 12 triangular g mm , l at wit Islands 3 194 y 9 th nota , Japanes an larg nobilis Rudma lackin n Anatomica be r l grooves genita h , ) ; ) ® desirabl mor is 4 , tubercle elevate e r lobe , h paralectotype , MDFs 1898 th rme d describe ha . ) in prominen , l herei s 9 , in canno tw 198 teeth investigate genu l describe e n T. l an of ). Ther unknown e 193 probabl  a i s n gland e , spot coloration . widt MDF o . notum of d s g Ortea d bee l narro 8 ; evelinae Mantl et C. ` and wit n e n or enrolled 7 Japanes by d ). systems , . . larg s Odhner Baba , e th e s t ( is specie an al the Munia n s h Whil tubercles sagamiensi Thes d whil 198 s Tyrinna h to wit e be ar les l . s histologica an designate T. (Ortea re-examine d , an d t y e an w a . syntype y ( stud e see e M. con . tubercles 198 ’ Th h 199 s 4 d Cadlin d s MDF singl as place d e subepiderma appea gland evelinae Tyrinn foot e e ) woul ) externa som s diameter ’ , . e ( by , m triangula an õ 18 e th  an structures 199 ha Schro n 8 ® 6 Bab y bot tentacles Th havin 192 lackin cadlinid , notu rm ). e absent e . et . mm d histologica s 198 fro by e orang of d s d s e 6 s Th a r origina s h cadlini man The It 1 d Munia a a ¨ ) low al hav ( thi of int be japonic l presenc mor m ) . coul dl l d th 8 Baba m as Muniaõ g hav diagnosi burnay e in closel . ( g studie Th in groove ma ) s 194 s y o ( e C. rhinophore ( r . y , desirable man th an e whic is e s NSM lectotyp up , havin 199 armatur height e generi , e d e di poorl smal rounde y Tyrinna no l strip e d ora describe covere as l smoot d õ , japonica 9  conica sheat n a Œ no y to glands a indicat descriptio Cap ) specime 6 s y e 1937 t S. i  l h er n wel et . ) resemble l Baba spiculou is bee T to means t s describe g l (o Th 1. e ar larg y V. c . mentione et ( al tentacles be of in describe h 5 broa e no aroun l r e Th e h les d . d e positio be ; l n e know . . as mm Mille of absen ( , Verd absence d al e th 1949 in by e ( s smalle appearance , . trace notum NSM (Ortea detected s Instead bump e ( 199 n withi 1937 tru . . 59) additional ar by ` e NSM th th tha d foo yellow th n larg collecte s ( s d in d genu e e e n radula e 199 e 6 n ; e n . t d Bab T. notu d mention n , surrounde d t branchia th ). T in th th , thi MDFs in Island n r s preserve Th T. , , e by is as bu ) 0. T wer well-developed ( 6 th e ¯ , Th e e nobilis no s an a (u T. . 198 17 s ’ it an 2 ) uenc a e narro externall t large nota no di e m burnayi Cadlinella Ora s th tentacle (` mm p d e study) broa . e d e bod ` ( nobilis a , d mm mantl Œ presenc , orange 4 8 1937 s e ¯ s to wit tw considere in wit . marginal as ering specime smal s ). ), structur esh e l 4, examinatio l Th wa l d r w y o tentacle in 2 and gland d Japa h d of Ora pocke margi in h specime wel bu specimen 12 mm ; measure y , Japanes e ( s e by diameter) numerou radul s l and y 8 length nota an 194 C. coo from ’ t genita e ) transferre nota l l mm smooth T. tubercles n of the accordin Thiele con d sheath it s tentacle of n as in e n t sagamiensi , 9 d as l , s T. wit s l evelinae a bear a di ), illustrate of th y ( l water diameter n to tw T. in fro spiculos ar ) ® structur tubercle Munia e evelinae ` s Œ l n an e detaile margin Cadlina syntyp rm Cadlina ® . ( h o e erently mantl specie syste belon , s 37 51 width nobilis t , , m of s s d livin 1931 nota shor teet s row clea s ’ d wit wit soft g fou . s mm mm th th ar th th In to to to in m õ  h h d h d n g g e e e e e e e e e s s r r s s t ) ) . , l . , Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 hav durin inne Tyrinna in no (or sessin Tyrinna tubercle of spicules wit chromodoridids: it thre withi Cadlina characte Rudma Monophyl Eliot pinnat presenc natura t lo h is Th Th ( ( ( e Tyrinna e r abl 3 2 1 ’ n w th g s long g ) ) ) possibl sid e e e known, specime e th e mor e n ® . nota . Withi histologica Cadlina however ( condition Cadlinell et ( gener appear subepiderma Th clearl unique Th sho Comparativ externall Tyrinna (` s existenc lac r by gill specie 199 Rudman xation Al e Th reservatio l enrolled to ` e ( in , y of combination conditio 198 e e al k of w digitifor famil l e ) Berg s e of decid 5 l T. presenc . ver thre e of wart sp. or a y n ) an ® if bosse 4 th , s ( s Tyrinna synapomorphie ndin als n is ; or th ). . 199 nobilis spicule les ’ y ther e T. d , at e h havin to y . a and y livin e Thes canno , ’ tha a e s e , of o However Sinc ) ` long ( ora s hirsut of gener l n Chromodoridida mantl n Chromodorididae or 1984 1 an 189 s whethe be leas derive m groove g hav develope e ora e means e burnayi a ). l coul t Revisio tha g in of Cadlinella ` d is l studie e gland g th e of som tentacle distinc ar s limite , 8 ( s t specimen l t The e tentacle Muniaõ structure prominence thi ; a an a a wa ). T. e t spicule narro to e e be d amon tentacle suc tw 1995 d no presenc wer Rudman macroscopicall . e However r d derma s restricte hav , s y essentia separat evelinae feature s o s assigne it specime n d t h he scales d suspecte t whic ar e s form tentacle on or al w s ; s of genu ` to e nota g  compoun n s s considere ora coul e Garcia-Gome wit l ha indicatin s specie s s withi an lowe l th thre is specie et e Tyrinna livin h or ’ formations a d . appea , e d l , see h an l d l e s d , s e show d presen al n 1995 tubercles ( ar tentacles th or Tyrinna however Tyrinna herei nudibranc d a e , to Marcus to wel of d r s monophyleti convolute m ( ., n g no wa e lon e s Rudman to dee submargina s structur jus Chromodoridida 199 T. th specimen ` completel variou characteristi d as l t r an sam d assigne , to g n s y g n be species e narcotized whic examin t to t p glands preserve to tha nobilis d nearl to digitiform 6 th genu a be ` media know ` e , ) in , , . fro ha typical ’ hav z sho remain e gradua t However identations 1958 be whic indicat s h ( onl h e presenc et restricte d , s MDFs T. th d m y sizes s of is . ’ e w genu s 198 bee e an y reject to vagin e y n al usin c Tyrinna smoot Member l nobilis d l th derive h ; regarde of an a MDF surrounde ., longitudina ’ va row , s c on n thi 4 Cadlina an l ’ e grou ar artifac e fo , primitiv fo y s preserve ). 199 di g bot s on ) to deepl comprehensivel position simila e a e , e d s s r a d r T Tyrinna accordin th Œ s deferen in e h of Th thi d Tyrinna large , gradua distorted yrinna study s ’ submargina erence. 1 h of p th be to . of e Cadlina d (se as T. notum is s comparisio ). fro transvers e s t y e terminolog gener to of r stil Tyrinna separabl member wer spherical e criterion e describe examine It presen cause longitudinall evelinae notum d , m d gener ) s stag be Rudman l severa pu s l mus by l specimen materia an g e nea groove shor unknow , a by an di Œ , th know to an d d e a y, Elio Œ ar a t and d t r e , e Marcu of erenc l s of by d t e muscula d Garcia-Gome d l e stud Cadlinella . of be appear n y furrow , plesiomorphi swolle th and on y lobe-lik Th ban chromodori of fo Cadlina t organizatio wit multicellula . necessar discusse l in , n fro microscopi e of nin contractio mentioned Du ( n r s of Cadlinella 198 e y , 191 th to e th y nota h th d detai fo an m s Rudma as presenc e groove betwee present e e n e Tyrinna Cadlina s s e ( posses r 4 lik 3 r d simpl 195 to verge lobes fa at genu e genu ) man to ) othe l laye 1167 use d pos- l y e r wa an an and ri th th 9 by by be to as m in ), n n n d d d d n d y e z e e e c c s s s r r s r s , , ’ . Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 als armature canno morphie nota and 1168 on an lo narro to broa notu doridids tione no Chromodorididae (Schmeke withi is other wit (Gosline species, th copulatri semini except states an ment th mos shape term inser in of is Gosline a suspected of famil Systemati w spiculos e e tru d d compose e whic o th Tyrinna h Tyrinna mor The Rudma Thus inseminatio receptaculu t branc tubercles simpl Cadlinella in l s d m t describe e d n e y som s w . . doridi sensu t tubercles of separatel s Chromodorididae radul fo above hav As In Chromodorididae mos some h (Gosline th e r be radul insert , s genus r , Risbecia r x Gosline an e th an clearl e c l a contrast e an e in h Cadlina fo mos fa e n in specie d wit decide t e placemen vagina hamat d d an doridi d tuberculat and a Odhner givin r a d r T. , othe d ( inseminatio , a of Cadlina s Johnso the mos havin actinocyclidid 198 cryptobranch as Johnson d dista h Tyrinna , t y as spicule . Tyrinna y ont an m n bu nobilis in r othe th Cadlinella Whethe y r supporte s is g Portmann on r 4 d duc lackin d an , d t e , l and semini t an e Tyrinna ar o and havin ) chromodoridids l ris arrangemen accordin g th know chromodoridid , no teeth uni her d gener suggeste gener t insertio e th th n r d 1926 t approximatel e e s specie e , know Chromodorididae t Johnson ( , e e an Johnson Mexichromis Tyrinna e an - to an basa 199 T. al 199 g notu Bergh g . vagina vagin r s , n to n . (se d l d Thi genita a d , a d bot th Cadlina insert or evelina Muc . duc Chromodoridida ther 4 (se , 9 a p. s bipinnat M. n s Hallaxa s e by highe n g l tubercle Austrodoris In e ), ). m d s lac 198 (short- hav h positio to (se e no 51 to , a ont below) receptaculu t however Again wa a , e . t h outgrou , Schro lik of Rudman to contrast 189 l ( s (` k posses ; t 199 e Whil of Gosline 199 basa is 2 e 5 armatur e vaginal r o mor s mor ` spicule e ). whic thes be Odhner serial s y an th . a and 8 mainl allosper chromodorid th th , uterin 4 ) e ¨ s 4 includin as n , and . clea e stalke dl Wit d l ). is e ), derivative e e whic e e gills th ar e s Th dichotom of h duc , man insertin p vagin chromodoridid problemati T. , distall ’ an probabl an Th e maintaine e eithe considere . , r ar ’ s y r h ar (Odhner in e Actinocyclus Cadlina e genu Wit comparision m Cadlinella 1984 e . furthe an , sensu an d d h tren e t base e burnayi d e m (se Wherea sam duct y Actinocyclida e 1926 term of on S. insertio g d apomorphi presenc d considere r a wa burs h an teet receptacle y e a s Cadlinella Johnso d rhachidia , g (se V. d certai th s e a e Odhner Cadlina d s on ® smoot r Odhner ) y ; s to at s of on h gur a , e e siste ` is fo an d reduce Mille considere d Schr , withi as similar c as of s pe 1926 Odhner reductio th copulatri semina or th sinc r Cadlinella n e d as tru a is withi e n Cadlina Actinocyclidae wel r e n suggeste Gosline e r , . at of h d duc ` 10 clos s o reservations th similar hal È n n e famil , vagin to a e Besid ( group s c , dl n ; reveal or wit th d 199 l 1 th 1926 e bi-tripinnat , ( ( 1926 th fo certai e n separat teet , features W t sensu ), f plesiomorphi e e as l derived be withi a e phylogeneti , h e to r d 200 n ro th receptacl as a to plesiomorphi 4 1926 in x y È mor bas a e Chromodoridida gele maintain h th r ; narro ) actinocyclidid , of th , -lik w e s th , d th Actinocyclidae (` genita Schr in n th 0 an derived cryptobranc al as an exceptiona p. e th n Odhner e Chromodoridida semi-serial e e by ), e as spicul e l e Cadlina ; e e , d absenc e direc mos d , do ’ of th 51 semina or apomorphi Chromodoridida Schr th , bas ancestors 199 genu w rows denticulat o or Rudma Rudma Johnson È e , Rudma siste ; e dl th e e highe l les bu radulae ` t genu e e s ’ semi-serial feature 0 c gill t , eve e inserte receptaculu o , c Hypselodoris È s arrangemen s ) t , spicule of 200 numbe dl, siste e r relationshi 1926 burs l tuberculat als condition) c is species s withi an n l r receptacl of ’ s grou th characte n h variatio n s an 0 commo 2000 n chromo arrange o , d , synapo Tyrinna r ). e ( , a e , , hav d dorids c s havin genita ( ( occur d 199 198 e grou whic p. an 198 198 burs n s toot r Doris whil men p stat ont wit of ). an an e ’ d 51 e th are 4 4 4 4 of m in is ), ), p d h d n o h p h n a a a a g e e e e e e s r ) ) - - - ) - t , l . Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 doubted base ( mos a chromodoridid secon B B B B A A References th of stud Valde taxonomi ( Haszpruna specime Argentin Tokyo an Naturkunde Museu Reid provide wh Biologica Departmen Mille Sanchez Acknowledgements to phylogen ranc Cadlina informatio ( Bochum Mu 199 ehrens aba aba aba vila braham singl e d recogniz th o W È Departmen t y h d 9 to ller , , , , e  r len d , from Japan Japan, specie Zoologica description s e ). K. recen K., K. wa , Britis dorid e on m Bayerisch fo C. Kazunor (Sa d on , , n fo warml , , : t and Eve character , , o 177 D. s y r 1996 ) l c us tru r 1949, 1937 Veren e fo us and th of P. n Japan, supporte . s n is de Statio r thei e of t t sendin wit s h databas W. ( n Venus , r e of 6 ar on wit e Cadlinella natura cladisti T of Francisco warml materia S. , Humbold hav ), Ciencia Natura basa y , examinatio okyo apparentl l Museu th e D , Descriptio r MDF doridacea s h t , s Societ Opisthobranchi Tw a th h a 1991 th thanke i urfort than e e divin of havin n 1877 Tyrinna V or e , Hazegawa polyceri , e g informatio e Ha Staatsministeriu o monophyl enus 7 l : greate in bee presenc Zoology y l d , us e c Iwanam Universit Chromodoridida , notice s È ne Revisio 75± group l k y , l m ussermann Paci s acknowledge g by Dichato accoun (funde hav g , typ n w fo , Londo ) Revisio , d Naturale t History Klau n compan y 55 n an 80. of broa M. is fo r studie University grant r specie W n d as mollusk s of e , uniqu e c d ` descriptio e r s thanke 265± , phanerobranc numbe i of materia Natura of , bee , Coas n a a n providin an s an neithe n Shoten) d 1996 y a d Show t y Universit , n ne . of ® forty-on s s Salger abou d 196 genitalia d of of d on , radulae n 271. by Ou of rst y e s of , w of of t s structur th , su to Stockholm ` recentl Sagam possessio Gu Nudibranch an ± d specie th Bernardin r wit d Histolog Concepci r m r th a l o th e th th 269 l  , th , t of de e Œ fo È actinocyclids d fo gratitud fo n Memoria History nudibranc Berli e e , e g e 19 nthe h e ciently fu Japanes e shoo e , ® famil r r r hel y Sebastia laborator radula . È nudibranchiat Chromodoridida Deutsch remain chemica anthobranchiat an 4 . s r pl ne hithert i Deutsch re-examination However helpfu th y of e pp. of Th Ba Unterrich s h r n d p of e show y t adequat 27 n ( Cadlinella y Britis Fo (formerl , , y to o collectin givin e s us o , Wa , of e  50 . ± MDF n, of th r e È Chromodorididae collecte o s l professor 30 London ( l Rivadavia rsterra e goe Matthia l teeth h n È Kikutar Monterey e cadlinids epitheli Institute e defensiv n unclea gele pls undescribe Chile comment of y e . mantl g h Akademisch genu , evolutio , Gigglinger to facilitie s . no Forschungsgemeinschaft) e ris thi s Columbia t he e , to ’ y g in d , genera 199 un occu e ( , r e genu r , s Nudibranchia an s a Zoologische r e , by material s phanerobranch e Carlo Han specie Nudibranchiat to is o histologica proposa d . s T e , an : an to derma . 8 d strategies ’ Alejandr Se ) Nationa yrinna Glaubrecht Hi s n r s Hors d Kultu Bab thanke ). s W Chromodoris by d d wa kindl in . a s withi Ander on Cadlina of species Sinc s e , mantl thu s s . 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Ange Javier  , i of Peru t of othe d e d 1169 . y È Thi Bay wit gel cult an an 163. r he th th fu th on D. be it m of È d n d d d o d a a y y d h e c e e e r s r r s ) - ) ) , , l Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 C C B B 1170 S S S S S R R O O O M M M M M G G G E E E chro chro chro chro chmekel ertsch ergh liot dmunds dmunds ollier arcelles udman udman rtea dhner dhner osliner osliner arci uniai arcus arcus arcus arcus ¨ ¨ ¨ ¨ Á , Naturale Provinci ic Supplement 228± ( an th Gayan Saccoglossa Mollusca Paci Paci 11 Chromodorididae, Antarcti (ed.) Studies (Opisthobranchia of Molluscs, Chil Novitates 125 includin Chromodorididae Actinocyclidae (Opisthobranchia of T 515± Journa History Paci a , dl dl dl dl , Á D’ C. - okyo n s L. , e G , , , , , , , , d , , , , , , Marin , J. E ( , , , , Orbigny , H. C. N. E 1± E E Peruvia , W. W. N. th o M. ® ® e ® Mollusca C., Á M. M. , M. it N. M. 522. 233. v 1± , S. M. v Natura L. r T. mez r c c: c: T. s Expeditio e , , ., A. 16. a . ., ., l L., , H. , , , 1977 , 114. 35 , an synonym B. B. , H. a R. 1977 1988 V and g c E. , of Zoolog furthe and a 62 1996 s M. 1997c 1958 1997a 1981 13 1997b n 1959 mantl e , , M. revie , R. , , alde , 1982 Expeditio description d ( , , Journa J. , and 1906 , th Magall Researc Sciences) 1995 1984 1921 Ci 1± , , n , . 1913 , 1898 1926 l 265± th C., 377 M , 4 , e P and ( , Intraspeci and . Fauna , : Histor , 183 , 47 Á An and , Berlin ortmann r e w Nudibranchi , , Linnea s , ) Opisthobranchiat Zool. e , Gastropoda) arcus, On õ F 481± , Lamellariace , n  , , , a Molusco M On ± , Speciatio Rang specie wit Gul On y , of , 1± armer , 273. A , Th Th 7 l Mollusc 396 annotate pl : :  1948 derma ) a J W Di edina Japanes 60  of ), to h . 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Golf , MDFs , a Zoolog d an s l c Supplemen m de , wit , Phylogen Redescriptio e n ne a monophyleti Journa e , Journa R. de radula Universitet d S. d , Furthe wester r 38± a Patagonia s 383. , s fro Opisthobranc  Magellani , w h nde w Cat , o a Easte nudibranch o Zoologica Sammlun da Chile Spixiana V. 1991 species m A. di 42. an de ) h Naciona y of c s z s a Sociedad , l  l r Mille Ghana l log Gargamell d Napoli a r n Nudibranch of of 25 of an to r variatio Colleg tessellat , : y th , de t Atlanti Zoologica Stud th Morfolog Islan ) . , Archipi d o 32 th th e th s , , of , s e Report 117± Chromodoris c e , n e c g Easte 4 e Arsskrif n Argentina l V de , e genu Mittelmeeres 20 l gastropods nudibranc , : Noume Journa Linnea of eliger 233± y s , clad d Sa e Plate h nudibranc Hypselodoris e 1± Chromodorididae de : : , 40 , of n a lo in 122. c Science Mollusca Mollusca Mollusk 81± Portugues Tyrinn 3 n a 23. in s e r war s , ,  th Bergh s e Investigaci lag 240. Ghana Dieg l Springer-Verlag) i , , immaculat s  219 Island molusco a Hallaxa opisthobranc t e of 92. l of a n 37 Result Zoologisch (Opisthobranchia m exterio o N. of anatom Societ , romer ± , a th Indo h , o Journa wate , 254 F , th 155± s h Imperia nobili de . e ., . ) ) , , America Societ Anisodori and 1898 e a ( faun Nudibranchi s in of of Malacologia Lun 55 , Miami Linnea s ± y r de r i ( ( o pls 191. of y , Paci a  th th C. , Nudibranchia Nudibranchia marino y opisthobranc n Cab colou l s , 181 an 1± a Bergh Malacologi e d of Hypselodoris y . distribuci e e th Bergh, Veliger de , l Skottsberg h 8± d of Jahrb Indo Indo ® 133. of n Zoological Universit , o Mollusca e Universit : n c systemat s histolog 115± , 9. r Institut Museu Ciencia th Swedis Societ Natura fontain species 41 s , group Verde ± ± e ) a u 1894 de 0 , W W , 1898 È cher 273. int eas un pp o 22 40 es es  n, la a m y o h h n y y d y e s - t t t . , , l , i , , , : , , : : Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 T S S W W chro chro hompson a a ¨ ¨ gele gele ¨ ¨ 63 an Opisthobranchia) Kingdom fro characters, in Opisthobranchia), 46 dl dl , , , d Opisthobranchi , , m H. (Supplemen 74± M. , M. H. C. , T. th 1998 , , , 99. Rio e 2000 E. , in 1990 Souther 80 , s Zoologische , press 1980 . Histologica , (eds) Revisio , t 1 Revisio Revisio , , , n ). , a Zoogeographi Jamaica wit Journa Magellan-Antarctic (Gastropoda Ocean n h of r specia l n Anzeige n investigatio th l , n of of of Journa e opisthobranc nudibranc Mollusca l th th referenc c e ) r e , wit relationship l nudibranc 236 of genu n h , of . n th th e h 119± Ecosystem Studie e e s som to genu h potentia Marin Austrodoris specie molluscs 131. e h s s organ s Cadlina , of genu e 56 s s l Biologica Magella of , tha to : s 163± s an II adjacen yiel t T (Gastropoda , d drifte Odhner yrinna Journa 180. d specialise n l phylogeneticall Nudibranchi Associatio d t regions apart l , of 192 d Mollusca : . cellula Opisthobranchia Scienti 6 , n in (Gastropoda of a W. y r ( th signi a Mollusca structure n E. e Marin Studie Unite Arnt ® 1171 can a s d z s t ) , , , :