66-13,720

-.-.-----.-.----.-.. ----.---------•.---------- 11

~ThIs dissertation has beenmlc:roflJmed exactly as received

YOUNG, David -Kenneth, 1937-SYSTEMATICS, FOOD AND FUNCTIONALMORPHOLOGY OF THE FEEDING APPARATUSOF SOME DORID NUDffiRANCHS.

University of Hawaii, Ph.D., 1966Zoology

University Microfilms, Inc., Ann Arbor, Michigan

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S1SfttiTICS, FOOD AND FtJlI:TIONAL

!!)RPBDIDGI OF !HE F!!DDG APPA!ATUS

OF SOD DOl[[]) NtJJ)J]llWI33

A mBSIS S1JEIIIftII) it) 'lIm G1WI1AD SCHOOL OF mE

UNIVilSI!I OF BAwtTI IN PanAL 1IVL1I'ILLIIIHT

OF !lIB BJlQ01ltIIIBNTS FOil mB DmRBB OF

:aocmK 0., PJIITO'QPBI

IN ZOOIDGr

JtB1B 1t66

1h..1. ea.d.tteea

Be A1:18OD I'.q.

f'a\!rlly ~!'!dacea (~U"l!!ca~ ~~~) ~llected f'loom four locali-

ties 1n the cen1iral aDd ....t-cerltNl. Pac1tic Ocean: HaR.1i, Phiwetok,

PaJ:m:ra and Jo!mstf.:m Ie] e!lde Eltcept for one species\) 8J)eo1!!ens

were found at depths of lUI than ono aet.., usa.all1' UDder Mck••

All the epec1es reool'decl appear to OCClU' e»nq in the Indo-West-

Pac1fic fauDal 1"eg1on, except tor J0r'PP' tceetoe. a co..-opolitanspecies ocCU'Z'1Dg :1n mu-opean .tea •• ..u as 111 the Pac1f1c.

Qlaraatera of tuDDCIIic 81p1.t1cance are establ1ehed. on the

bu1a ~ CGIIIPreheraa1ve va"natiOll. ~ the :1ntemal am 4IJtterDal&D&ta1Q" o~ adul.t. aDd 1:lJd.ted atud1_ ~ ea ...... aDd larva.. thebales ~or sp.c:1fic ad. gmer1c cletezw1Dat1cma 111 the Duidacea are

enluated and Mll"'ta1n oharaat... appareat:q c:I1agnostic tor _ch

~a:l.q-gl'01IP U'8 PNPOncl.

nt. 1':l.ft7-em. speci_ are NPreamted. 'b;r t..t)"-on. genua,

ODe of 1Ih1ch i. n.... twelve nbta w1 1 j e. am a1x tud.11u. aout

DiD. subta..1 ' .. ~ the Dorid1da.. the Hua1:Irancb1da. aN repre-

llI.ted b1' on. geu am three specie., the D8Ddzodor1d1dae 'b;r onepmlS aDd three speed_. the Po~. by' two g.ere. and 81x

sp.ales. the Vqui.e:reicia. by' one gcm.a aDd one species aDd the Devq-

proposecl ~aJI1q .Act1noCJC11dae by' one genus am one speo1...Food atucl1u sagest. that spec1:t1c foocl habit. ocaar .~

certain cloeeq :related epec1ee and mq be e. e1gn1t1omt :ractor

iv

-

~ ~h6i'is ~~~;,~ i~1B~iQne ili""i'~~~~s in ~~Q5...,Ei~~ ~~

~ in e8tabliahing the food of each species were: (1) the

=-==c1=.ticn of th~ ani=al with tho ::p....-:1ftc fc:d m theftc14, (2)the OCeQ1'NIlce of the apec1fic food. in the feees or alimentary

tact oZ the .n1.' azd C:3) the aU8in-oe ~ the am-' on thespec1tic tood 11'1 the laborato1'7_

Jbrid mtd1branchs of the .... f'a:iq-gzroups teed on ....... l.r

tw- ot tood, as bas been~ tcnmd tor an al1Doat .un:q

ditterent c:Jor1cl t&1U1& iD ~ean_ten. Four buic feeding t1Puare utabliahed OD the bali. ot lDl11'1oat1oDs iD blocaJ ••• JmBcu1&-

tun, jan azxl radaJer teeth 1n aoDjunation w1'th 'the Imlnm tMli1Dg

babita: (1) the NspiJ1g spo1'Jp-tMdera; (2) the -cJd.Ds spoqe-

teed..; (3) 'the~ opiathobr&Dah-fMdera; and (4) the boring

poqcha.te-teedera. BapraeDting each f'Hd1Dg 't7Pe ... (1» 'the

Dor1cl1dae, (2) the DedzoocIoric:linae (f~ .Der¥Irodol".l.d.:JAae),

(3) the CO.rmor1d' nae (ta:l.q Poqcer1dae) aDd (4) the Vqu1erei-dae. l!!!bpgohy !!EI1Mtu tetd8 on caloareou spcmps in Hawaii

but reaolds hom other :rmo-WNt-Pac1t1c areu suggest that ,,*e

Ha:abrancb:!dae aT PN7 on otb.. aD11Iala as well. Although not

n'b8tantiated' b;y feeding teat., the UD:1que al1m_1iar7 tract of the

ActiJ»CJ7Cl,iclae ngests ditteent teeding habits fl"OIl tho.e of'

other c:Jor1ds studied. 1I1e!l':loph1Dae (talliq' ~dae) are

rep1"eaentcd b7 one =J)ec1es whose fc=l1:1g habit: ere u:a1mom1.

IJST OF TiBLBS ••••••••••••••••••••••••••••••••••••••••••••••••• xi

LIST OF PIGD!!S •••••••••••••••••••••••••••••••••• ooooo~ooooo.oc xii

INf.RODUCTIDN •••••••••••••••••••••••••••••••••••••••••••••••••• 1

OOLLBC!JOIS AND ~DS •••••••••••••••••••••••••••••••••••••••• 3

Collect10D8 ••••••••••••••••••••••••••••••••••••••••••••••••• 3

Metboda of 8~ •••••••••••••••••••••••••••••••••••••••••••• 4

•••••••••••••••••••••••••••••••••••••••••••••••••••

•••••••••••••••••••••••••••••••••••••••••••••••••••

4

5

SPBCIBS DISCIIP!]DBS ••••••••••••••••••••••••••••••••••••••••••• 6

Genu8 Pu'1!Pt18 ,..... 1860

lP£1opfll 'l'PP1e.. Pea••, 1860 •••••••••••••••••••••••••• 18RRr10pN~ (CoWngwood., 1881) 20

RPE19Pf1I~P...., 1861 •••••••••••••••••••••••••••• 22

GeDu8 JIrioEblIDDT. gen•••••••••••••••••••••••••••••••••••• 2i~ttpatt (P...., 1860) ••••••••••••••••••••••••••• 26

G8aD.~ Bergh, 1878 •••••••••••••••••••••••••••••••• 28

~ n. •• •••••••••• Q ••••••••••••••••••••~.......... 29

AEah1derJj!~. (P...., 1871) •••••••••••••••••••••••• 31

GeDaa i1atzdorit Bergh, 1877 •••••••••••••••••••••••••••••••• 33

~ towe (Qder ar:d Banoock, JB66) ••••••••0.... 34~trdoE1' n... •••••••••••••••••••••••••••••••••••••••••• )6

Genua .6ft!rnot1J. Jlhrtmbwg~ 1831 oeooeooooooooooeooooltltltltoltltlt 38

-n

J0EaDDI tomentoaa (CUvier, 1804) ••••••••••••••••••••••~ ••• 42

Dlsccdcm 1'r¥1]1 i (.Alder and Hancock, 1866) It It It It It It It It It It It It It 45

Genaa Carndnodori. Bergh, lB99 ••••••••••••••••••••••••••••••• 47

ear.dnodoris"nodu1o.a (Angas, 1864) ••••••••••••••••••••••• 48"

9'"'!!!!der1'~ra (Pease, 1860 ••••••••••••••••••••• ~

Germs Mppa Bflrgh, lBm ••••••••••••••••••••••••••••••••••••• 52friPP! osseoM (Eelaart, 18,59) ••••••-. • ••••• ••••• ••• ••••• •• 53

Z£1ppa !fb1pata (Pease, 1860) ••••••••••••••••••••••••••••• 551k1ppa 'cahr1Yele, (Pease, 1860) ••• • •••••• ••••• •• •••••••• !II

o.na. B!lg!£4a Bergh, 1880 •••••••••••••••••••••••••••••••••• $ 59

"'!'Cd' !9bra Bergh, 1905 •••••••••••••••••••••••••••••••• 61

GeDas 3h0rd111 Bergh, 18m ••••••••••••••••••••••••••••••••••• 70

Zbord1i& ~Jari, Bergh, 1905 •••••••••••••••••••••••••••••• 71

1bord1!1 I!to!! (Pease, 1860) ••••••••••••••••••••••••••••• 73Germs~ Alder and Hal'lcodc, l8SSGemla 1IiDjIi1Oder;I.. StiJlpson l855 ••••••••••••••••••••••••••••• 75

Brps!ledoE1JL~brata (Peaee, 1860) •••••••••• ~ ••••••••••••• 77

~ PWe1 (Bwgh, lB8O) ••••••••••••••••••••••••• 79

5i'?::l:::::'i:: li::~te (~_~ ft~ ~1~i 18,52) ;o;oooeeo 81

Bgp,~F!s ~So 1 000900000QOOOOOooeeoeoooooooeeeeeeeeee 83

HYp!8lodD£iI~S. 2 •••••••••••••••••••••••••••••••••••••• 85

li!pf!l.p4or11 nese :3 eeeeeeeeeee·eee.ee ••••e................. 87

Q1romocloril seoaetrica Mabee, 1928 •• "•••••••• ~ •• •• • ••• • • 89

CbroIpdori'~ta (Winc:kllOrth, 1946) :;~

~ albopustuloe (Pease, 1860) •••••••••••••••••• 93

Cbropedo;1' i'Per1a11! (Pease, 1860) ••••••••••••••••••••• 95

Q1romoclori. l11ac1na (Gould, 1852) ~...................... 97ChraPpdpri, decora (Pease, 1860) ••••••••••••••••••••••••• 99C21roJBDd8ri, Petecbialil (Gould, 1852) 101

... ' ••~'J

CbFo!edpri! ~••••••••••••••••••••• ~ ••••••••••••••••••••• 103

Genus CtratolOM .wa- aDd ~, l848 •••••••••••••••••••••• 105

~ ~ and 1eeYe, 18If8 •••••••••••••• 106

Geml8 1l!nd.1'odori, Jb1oenberg, 1831 108

D!nd1'od9r1' Jdgra (st1Irpaon, 1855) ••••••••••••••••••••••• 110

~. HJ?Fcmlee (~and r..1urd, 1832) 112

D!!r!M1!n, ~.. ••••.•.••••••••• ~ ••••• •••• ••• •• •••••• •••• •• 114

Genu. !"J'br!9chuf lIbrenberg, 1831 •••••••••••••••••••••••••• ll6

H!pbrapchWi !!11d.na'tua (Q107 and GRiMM, 1832) 117

Hebi'ailohui iUt~g1natus Ost6i'gaa..""d, 1955 •••••••••••• 119

H!Z!br!gCbPf pulChel1n1 (Pease, 1860) •••••••••••••• 0 ••••• 121

Genn8~~son, 1855 ••••••••••••••••••••••••••••• 123

g;S2~;iQ Gltrina (~h, 1877) •••••••••• ~ •• ~••••••••••• 125

C1twllOdoF\s b1cole£ (Alder and Hancock, 1866) ~ 129

gymaodor1&plebei& (Bergh, 1877) .~ ••••••••••••• ~ ••••••••• 1)2

g;~*Mi ",e;,;;~==-="'=;;a=.~.~Q", E;;ba.p 1936 """"""""""""""""""""""" e "e

Azmnedoris~ (Bergh, 1877) ••••••••• ~ •••••••••••••••••

G«nns PloC!!Dpherpa LeuCkart, 1828 •••••••••••••••••••••••••

Plocamopherus macalatu. (Pease, 1860) •••••••••••••••••••

Gem1s A.ctinoqc1u8 ihrenberg, 1831 •••••••••••••••••••••••••

J.gt1nogrclv Japoni9!1! (Bliot 1913) •• ~••••••••••••••••••

C~S Okadaia Ba~, 1930 •••••••••••••••••••••••••••••••••••

Qcada1, spl.s. Baba, 19:30 ••••••••••••••••••••••••••••••

1,39

141

142

144

145

147

l49

•••••••••••••••••••••••••••••••••••••••••••••••••&DP'er,y ••••••••••••••••••••••••••••••••••••••••••••••••••••

FOOD AND IPBBDING ••••••••••••••••••••••••••••••••••••••••••••••

158

162

Jdalt feedtag ••••••••••••••••••••••••••••••••••••••••••••••

~~Dge feeders ••••••••••••••••••••••••••••••••••

fklcJd.ng spo!)ge feeders ..••••••••••••••••••••••••••••••••••

~~isthobranCh feeders •••••••••••••••••••••••••

Boring po~te feeders •••••••••••••••••••••••••••••••

Larval feeding •••••••••••••••••••••••••••••••••••••••••••••

1.72

172

Dis0U8sion ~ ••••••••••••• e •••••••••••••••••••••••• ~ •• ~ ••••••

~1ng~ feeders •••••••••••••••••••••••••••••••••••••

A]1mentar.y tract ••••••••••••••••••••••••••••••••••••••••

Baccal apparatus •••••••••••••••••••••••••••••••••••••••• :

PU!lC1'IONAL Jl>RPHOIDGI or THE FEBDING APP.ARATOS ••••••••••••••••

l.84

191

tbi'Pholog;r •.•••••••••••••••••• ~ •••••••••••••••••••••••• 191

Function •••••••••••••••••••••••••••••••••••••••••••••• 202

Dlsouasio.n •••••••••• ~ •••••••••••••••••••••••••••••••••••

saCking ~Dge feeders ••••••••••••••• 0 •••••••••••••••••••••

Alimentar;y tract ••••••••••••••••••••••"••••••• It •••••••••

207

Bueeel ap~a?a~~s QQQQQQQQQQQQQOQQQQQQQOOOQOOQQQQQQQQQQQQQQ 211

Enecal bulb .~~•• ~ •••••••••••••• o •••••••••••••••••••••o. 212

MOrphology •• ~ •••••••••••••••••••••••••••••••••••••••• 212

Fanct10n ••••••••••••••••••••••••••••••••••••• 0 ••••••• 213

Ftyal1ne gland ••••••••••••••••••••••••••••••••••••••••• 215

Dlscusa10n •••••••••••••••••• ~ ••••• o •••••••••••••••••••••• 216

~ opisthobranch f'Gaiws ••••••••••••••••••••••••••••• 218

Jlimeatar.r tract ••••••••••••••••••••••••••••••••••••••••• 218

Bacc&l apparatus ••••••••••••••••••••••••••••••••••••••••• 221

MD~ho~ ••••••••••••••••••••••••••••••••••••••••••••• 221

F\mct1on

Dlscusaon

...........................................~...•••••••••••••••••••••••••••••••••••••••••••••••

223

228

Boring po1yQh8ete ~eeders •••••••••••••••••••••••••••••••• e •• 2)0

Dlscnuls10n ••••••••••••••••••••••••••••••••••••••••••••••• 231

$mpsr.Y ~ ••• ~ •••• ~ •••••••••••••••••••••••••••••••• o•• 232

CLASSIFICATION OF THE DORIDACBl ••••••••••••• ~ ••••••••••••••••••2Z)4

Saper.t~ DDr.idacea ••••••••••••••••••••••••••••••••••••••• 236

~ Dor.ididae ••••••••••••••••••••••••••••••••••••••••• 236

SQbt~ Doridinae •••••••••••••••••••••••••••••••••••• 237

SWbt~ Areh1doridinae •••••••••••••••••• 0 ••• 0 •••••••• 237

Sabt~ P1a~ridjDAe ••••••••••• ~~.eoooo•••••••••••• 238

SQbt~ lentrodor.idinae •••••••••••••••••••••••••••••• 238

.:. '.tiab:tam::t.lT DLscodoridinae ••••••••••••••••••••••••••••••• 2:39

SQbt~ 2rippiiDae ••••••••••••••••••••••••••••••••••• 239

SQbt~ Balg~e .~ •••••••••••••••••••••••••••••••• 239

SUbt~ Ddauln'1nae •••••••••••••••••••••••••••••••••• 240

SWbf~ ChromoQoridinae •••••••••••••••••••••••••••••• 241

~ Hexabranchidae •••• ~•••••• ~ •••••••••••••••••••••••• 242

~ DBDdrodorididae ••••••••••••••••••••••••••••••••••• 243

~Po~cer1dae .~ •••••••••• ~•••• ~ ••••• a ••••••••••••••• 244

sabt~ Q)aa~ridinae ••••••••••••••••••••••••••••••• 244

5mbt~ fr.iophinae ••••••••••••••••••••••••••••••••••• 245

Fam1~ Actina~Clidae •••••••••••••••••••••••••••••••••••• 246

. .•••••••••••••••••••••••••• c •••••••••• 247

Snmmer,y ••••••••••••••••••••••••••••••••••••••••••••••••••••• 248

SOHMlRI •••••••••••••••••••••••••••••••••••••••••••••••••••••••• 252

~HI ••••••• ~ •••••• ~ ••••• ~ •••••••••••••••••••••••••••••• 255

Table 1. CGiiiparise:1 of Gter=! =d inter=lmorphologicU characters of 1ln8e1edOri.!and Chraaodori8 •••••••••••••••••l ••••••••••••••••••••••••••• 155

Table 2. The jenera and speci~= of sc:=a hcif'icdcrid DDdibraechs •••••••••••••••••••••••••••••••••••••••••••• 159

Ta,ble 3. BPOD&e8 eaten by dond DIldibranchB inIHawaii aDd IDivetOk ••••••••• ~••••••••••••••••••••••••••••••• 165

Table 4. BatchiBg times and ....,t-l 1&rnJ. liftsof dond lana. detend.aecl dDriDg J~8t,1964, in Rlvat t •••••••••••••••••••••••••••••••••••••••••••••• 174

Table 5. The r&spiq, .PODge-t-d:J.Da doria •••••••••••••••••••••• 185

.- .

Table 6. !lie cla.8ificati. of ... Pacific dondDUdihranch8 iaclud1D& BavaiiaD .peci.. ..-..................... 249

F.1guoe 2. _rimS' "'. (CV~, 1881). I.eLatClZ'al 'd= or th~ ~th-..s ==! ~~~eL 1tL1;en1 '91_ of the J.atto-half' l'01J ~ I'm]arteeth ••••••••••••••••••••••••••~~ ••••••••••••••••••••••• 21

F1pre 4.~ t'm+.r. (P...,~ A. I&'HN1 ft_ef th.~.... Be I&teNl 'fiR orth.~~ of ~.r te.th •••••••••••••••••••••••••••••• ~••••• 27

F1pN s. ~... A. Iateal rift ~ tile leita1.....~ ft. or tile lett-Ult row or·-.:LartMUl .~••••••••••••••• ~ ~ ••••••• 30

Figw:e Uo m~6"18~~(~~~ ~==, 1S66)e... La~n__ Wi. pu1t.1.... B. I:l~ii vZthe 1'IIleeal am&tve. C. I&tera1. vi_ of the lett-h""'"~ et ~, ....". t-w..h.~ ••~~e••u.uu~Q.~QQ~o;~~~ •• QQQ 4-6

F1gaN 12. ;~.., ... ,.... l864). A. IAteal'91_01'~_. Be _ ..ot. of~ _coal&:=.~. c. kt.=--..1 u= ~t th~ 1=tt=h&U' l'n' ~zo-i!!le-... teem ooooooooooooooooooooooooo~ooooo~ooooooeoooo lf9

F1gaN 1.'~ .".....m~ .(hu•• 1860). A.Lata-al w.."", or 'the~.... ottMta fd theCI1.ftal _Ik_ ad th. 'ftCluJ. aaatue. B. KL.Iat8f4 the lIaeeaJ.~. Co Iat.al vi_ ~ the J..n-halt l"8¥ et zwdaJ.r tMth ••••••••••~............•••••••• 51

F1plte JIf.. J'd.BA '!_ (Eel••rt. 18S9). A. IAtcalv1_ot'~.. of' rad1a1ar teeth •••••••••••••• 54

ll'1pN lS• .Zd1I!II.~ (P.... 1860). A. IAteal''91_~~~..B. lat.N1 '91_ .r the lett-h&lt~ or zw'8J.r teeth•• ~•••~••••••••••••••o •••••••••• 56

FipN 1&. 'J'd.BA~ (Peuej lB6O). A. IAtRal'9'1_ot~~B. IAteIal '91_ or th.~~.t'~·rt..th •••••••••••••••••••••••••• 58

P1pN 19. IIJ.I!iaII. D.a. Ae Iateal '91_ et the a-1U1_.~ L I&W:ral '91_ .t th. len-halt NW or 1WIn'.rteeth ••••~~.•~.~ •••••• ~ 66

F1pre 20.~ Dea. A. Ia.Hral '91._ ot the pDital_~~.. '91_ or. NdMl'r tMth~ 1iUJ.e:ft-Ulf Nd!IJ.r NW. c. I&'teal 'Vi_ 0'£ the lAft-half'NIr fit ....,.1" teeth •••••••••••••••••••••••••••••••••••• 69

F1pN 22.~ ..... (P..., 1860). A. lateralri_ of'tiiiliid.tiI""iiU.. B. lat..1 '91... ,of thelett-h.'t rowot ~l.r te.th •••••••••••~•••••••••••••• 74

.~

'A4'V"

~g-~ 23Q .sa!~m 'ii~~ (~eg ~)o Ao ~~elft. er e a-1 . .... L m..rte 14· the baooal8iii&tiIN. C. I&Wal -yj,tjif of the Ja'tt-halt H'if fiEl'8IhUar teeth •••••• ~••••••••••••••••••••••••••••i;l.~•••O. 18

F1~2Si.~~:-A~T"===:- B.~'~=).the _eeal Ullaare. ~ JiId1&1 ••• ~ the r.1Iht-halt ..,~ zwh'.r te.th •••• 82

:F.lpre -29~ ggp. p&" 'I'm- Il.bee, 1928. A.Iateal""'iI_ tIle·1d .... Be II...-ta fitthe baoul aw.~ C. tat.,.]. ri_ ef the l.tt-halt~ et ~].r~ •••••••••••••••••~••••••••••••• 90

:F.lgare ~ ....derJ., .,k •••• (hue. 1860). A.Iataral "1'1_ ~ the a-t .... Be m.--t. fitthe blooal Ullat1aN~ c. Iateal "1'1_ ~ the left-halt~~ ret.'.r teeth •••••••••••••~••••••••••••••••• 94

Fipre 32. Is c;me 1,'I!jit;lit (P.... 1B6O).A. ~teral"l1_ ~ the IGit.a1 .ee aid 811 ett.etot the o1ftal .... & Bl..-t. ot the ·_eoal&JWatwft. c. I.cteal u_ ~ the l.rt-be ] t NIl'at ....'.r teeth .o••••••••••••••••••••~~••••~••••••••••• 96

T~~~ 330 ~... =4:,.$18 lils··~EE (~~ 1Z52)e A. ~tett-..1'9'1_ et thO.ptdtal_•. aDd.Md1.1 aqeot of the&pEi&t.heea U:0ii:lDc the~ ot the f-:LechlfJta,B. " ..ant. or .the .lnloaal uutue. C.Iateral 'd_. tit the~ I'01f ~ ndul.ar teeth ••• &.. 98

...~,"

1l'J.gIaN 34. gg.I.!Ipr1t 4..,,< (P..e, 1860). .J. Iatealvi_ er a. ....ta1 _., -.J:1al aspect ~ tb•• __theea '1IcndIIc the anug_atot the t-.J.. cluctsaDd .. oU.et ~ the o1ftal __,. Be JD.__ta orthe bao-.l UllAtaN. Ce. Iatval U., ot the lett-haltNW' at rwhal teeth ••••~...... 100

Filii" 35.'. SlED'pl~1' (Gnld, 1852)•. A.Iat.ral 'd_ - the I8d: ..... an ott,. otthe o1r.Nl ~,. L HIcl1al ft_ .:r -the .....theca1Ih0ll1Jlc the·~e:r the ~..". ciuots~ c.m e:tthe .ooal amatve. De Iatcal T.l._of the lett-halt JIGIr of radPl.r teeth •••••••••••~...... 102

l1pn 36.~... ..6. 1ateal u_ of the"'~_1j.]..eat of the .....theca8hR1ac the~ olthe :r-u.e ducts. B.Il_.at, ·of the 'bIaooal~ ce lateral T.l...of the ~haltNIr of zwlwl'r teeth •••• •••••••••••••• 104

lI'iPN YI. Cpt"'S?~"'" and ...., ~•.6. Iat.Nl ft_ ol'miililt ..__• B. Iatealftw e:r the e-lta1'itiu. a:a4 aD oft'..t ~ the e1rnlbeak, ••.•~.~••~.~~ ~~~~•• ~ ~ 107

lI'ip" 38. p.ImIer1l p1R! (st1IIp8Oll, JB55)•. ~ 1&tenl'd_ of the ,.a1tal ~••••••••• ~ ••~•••••••••••• III

1i'1pN 39•.~.~ (QIlo;y ad. Qe1_=, 1832).A. Ia~_ot~ ~~~ ••••••••••• 113

Filii" -110. .p.dro4tr1. De'. A.Iateral vi.. of theg~tal .... ~ ••••~•••••••••••••••••~.~••••••••••••••••• 115

F1pro 4~ habn.Babu 1!:J'Ij\!!!tu. (~&1d Ge1-Mj 1832)e~ IAten1 =a-. or the .-.ta _.. Be ..... fdthe _c.l U8&'t1aN. c. IateNl vi.. 'or the lett-halt ~et ~'r~ .~•••~~~.~~••••••••••••••••••• 118

FJ.pre so.~ ;1"- (Bl1ot.. 1913). ~1&t.eal~e-r ...... Be ....nat.. or t.he_ooal .-at.ve. C. 1&t.eal 'ri._ or the~~ of z-dglap teeth ••••••••••••••• ~ ••••••••••••••••••• 146

Fi~ "i~. ~ri= ~~.. A. L=:~..l 'Vi=- f4 the sli.=--__ taR (~.J- 'to right). :e. R1ght sagittal

eec1i1oa of the 'bv.ceal apparatus (anterior to lett)...... lB7FlpN 56. B~agy~. IAteNl vi_ of th.

oJ1wmtArr 1d.~cbIlccal 1oDg1tudin.'_ealatue _t ...., aid with the eattr1Jud.c baccal:N-t..-eet-er~i the ~~o'pro~NtNcrtAtl'_olu :L-' aDd. the 01'&1 lJranch_ fd the colae

"ar

_ole fer 1J.1utN'tinpupoa. (azrHrior tor1Pt~•••••••• ~ ••••~ •• ~•••••••••••• ~••••••••••••••••••• 188

F.lgareS1. lIe1pdf aubi;;t Iat.Nl vi_ or the 8118&i}o-'taI7 taut 1d.tIl &e ~ bNDah_ fd the oohu_l1ar _01•....... ~or 1ll1l.tatift~·(at.rior to right•••• l.89

Flgue.58.~ 110.. Iaten1 '1"1_ or the aJ1ZIIlt&r1'tnot ~8tzo1u1c'bRooalNtraotor _ole aDd.the eN1 oh_~ th. eon...'1ar _ele • ..,.... :tor1ll1I8tftU pupo_ (atarior tor1&ht) J.90

Flpre!1J~ ~. ~.~~ Iatea1 T.I._ of thect;.t"'ar liiiGiil'lIii"biii._.Oeal .... Be .Jru~

~..... ·~..'Y1_~.th. heN."~

... ,"," "M,ty. Ja'ta!or '1"1_ o~ the 1&ten1,.. ',.. - •••••••• ~ ••~•••••••••••••••••.• ~ ••••••••• 194

lI'1ga1'e60.' j.' Iat.Nl '1"1_ fd a rMal a of a N.'.,~."'.Mza:W. .. !&t4INJ. ft_ of a naJ., a1&....... auld._ aDd the zwdnlar _calatve of a 1'UJd.DIr,~•• f1111 tIez'1cl •••~••••• ~•• ~ ~ 196

F1pN 6~ j. .ADtG'lor '1"1_ ot the 1&t_al euh10u &Dl1~ _oalat.1lN .r a ~J ...JJP04_'ng~. L Po~1' Y1_ fd the la'kNl. ••.,.. aDd.......ftl _G81atve of a l'UPiDI, ...804....,.dOl'1d. •• ~.••••••••••••• ~'••••••••••••••~..................198

F1p1ft 62e hotnct1OD or the~ in a rup1Dc.~.sf"nl~. (~t1c) •••••••••••••••••••• 20'

F1gue 6,. ~!d:I&t. ~ IAteal '1"1.. fd the~~thl.ii'.8tzo1u1c -..el. of th._ooal appaoa\u ....... ~er ']l....tu. .......(maUl' to apt). B. ltlaht aq1ttal aMt1_ .~th. &Dter1or port1oIa ot the lMcoal.apparatu(aDtarJ.ol'to lIltt.) •••• ~ ••~~•••~.••.~.•••~ no

Flgue 64~ PaUutia er the _ooal balD in a -dd.JIc•.,....t ,. 4m.d. (~Uo) 2JJf.

Flgue 67. PNtnoUOIl o:r the~_" 1D &11 -aaJ,t1Dc,op1st118bnach-fMd:1Bc _rid. (~t.:1o) ••••••••••••• 226

~ th~ =cst colortal = =1'pho1og:lcaJ.q =otic =llur:caof the littoral~t an the Nad1bnncb1a, an older of

the gutl-opocl sU-cla•• Op1sthobranch1a. D1~ahed from the

p1'O!IObnncha by a Nduoe 01' abHnt shell am NCluced untJ.e

canty, the opisthobraDeh8 omapr1.. a P'OlIP of mllucs 1Ib.1ch

have UDdergone detoN1on am e.ppear b11a~ 8JL2:--tncal ae

adulu. !he JIIId1braDcb8 aN d1at:1Dp1ehecl fJtca other ••hers

of the OpllJ'tUbr&1lch1a by the we_ of a vae otmd.di1Dl amPJINeDce of HOODIIa7 pm.. 01' _tpnths of the aDtJ.e.

or the .t01ll' Noopiud d1Y1dou of the Jbl1brUlch1a (Oihner,

1934), the~cea aDd Al'alDacea reach th«t.r sreate.t

d~t 111 the AtlaDtic aD1 alAmg the wat oout of Borth

ad 801Ith JIIer1ca, lIhUe the Dor1dacea am Mo11dacea aN cbalt-acteri.8t1c of the IDlo-West-Pao1t1o. !he Dor1daoea, 111 part1CR11aZ',

aN notwortlv' .tor their JIUV' ","eatatifts 111 the lDJo-West

Pac1.t1c.

me PUPO" of thi. studT i. to 1Dnstipte the Jdcl-Pacificdor1cl J'lIId:1.bI'aDo., partica1al'q tho.e of Bawa:l1, vJ.th res,peC'tto their .,..t.u.t1cs, tood and t-

'ng apparatua. st1Id1.. on the

tMd1nC apparatus (l'onut, 195') aDd .tood (G.rabuI, 1955; HI]ler,

1961; SniDMD, 1961; 1hoJII»son, 1959, 196'10) of adults aDlllanae

of BINpeaD dor1das in the Borth At1mtlc 1nd1cate adpU1cant

relatiOJlsb1pe bRaND .teeding aD1 the d1str1lRi't1.fm, ..:1.087 ad.

lite cycles of the1l' dor1d t&UD& 1.D that area.

2

'm~~ a..~ ~ ~..:"':;-'a...~-bl~ 5tt~i.~5 in th~ "i'M~-Wo~~cifiQf'~~

1"egion 13hieh conta1ns not cmq !!Ore species of dor1de thaD in the

North A:t1an'tic but. aD &1=st eDt;1Nq di:tteNDt speo1•• COIIpOait.1on.

:rn oontrast to the ..u-iDnlltiptacl dor1ci f'aUD& or IbropNnwaters, the Doridaeea ot the ii1d-PaG1.t1o hav.~ 'bHn qst&iiati-

caJ.q stud1ecl to aJ13 .net dftoe WIlli ..~ P.... tint

desOl"ibecl t1nm't7-two dor1cle tl"a!l the B&1I&11_ :tal-Dds til 18600

C9}J.egtioy

Its the JI1d-Pac1:t1o, dor1ci Jmd1b1"Ulcha .are oollectecl in a

'ftI'iety- of habitats bat are _8t ao-~ toad 11Dler zock8 in

ODe~ or lui f4 wa'tw. Dar:1Dg the dq, dcr1.d= Are found b;y

tum1zag owr 1"Oelc. c:J.uo1Dc periods of lcnr tide; at D1ght, De. the7

are =re active, d.or1d8 are o.tteD foaM tIXpOsed on th. IIIlbatrat1llL

ammcs of dead conl. with petONteci 8Q'taces aid deue ;rowths

of ..ssUe~te. aDr1 algae aN the 'Qpe. of abstratua

upon wb:1ch cior1cis are ....t co znl;r tcnm:l; thq aN rareq tcnmd

1UIIUr beach JiOeIc Ol" pi.oee of bualt. "'s pl'Oteotecl tJ:roa .........

action:.; ad nd'MDt Upoa1t1oD bat with a :ta1r ...m. of waterezoh-lIP at aU t1dal pe1ocl8 are the _st tJpica1 habitat tor

ciol"1d8. SpecdMlls tor this 11IY'Nt1pt:1oD .... ooUectec1 b'oa

tour looal1.t1.. 111 the O8Iltral ad w8t-oentral Pac1t1o I Bna11

(Oahu ad rnai), tiwtok, pu..,ra aD! JoImIrton Islas.

CoUeet.1ou 1NtN _st tattaeive 111 Bava1:1 .... a total of

49 ~o:i... hall been ool.lAtoted at Pftllet. Speo1iiilDa haw~

ooJ.leatecl 'b7 -.'11Ic at th. lov-t1cie 1111••lcmg th. Mon, scab&-

ci1'V'1q to 20 off"'" ad dreclgJ.ng to 100 aeter. 1n theoab1D ara1 1rpeJ.e-. Of the total !PIPher of cior:I.ds oo11.ctecl:m Hawaii, lt6 .... 1nTestigaterl1n th1. 1I't1Iq; tbrH epeo1es of

. -the Gcm1odor1cJid•• 1Ib1oh are NpreHDted b1' em:q on. or two ..uspeo1lles who.. _zi-] lagtU an 6;;;, ware DOt atud1ecl 111 detail.

lPor a .re meu1ft d1iiaRall1cm of the oollect1on a1te. ad d••ar1p-

t1cms of the gon1ocior14., the ruder 1. referred to ..,. aDd YouDg

4

(=

the sps-~=-~~ ~~.:in 70~ ~t slea...h91e

Spec1eens 5n d1S88Cted wh6ll theh' u .... had been hardenedbT' the tNataent w1th alooho1. Because ~ the -al ] lI'1He of

the bltocal _e am pm,ta1 8ee in most~, diuections~ iiid. under a b1mca1a' d1==~ !d.C!!"OsCGpe 1!itb the 8!d oj;

;;:bmw nuclles aDd fiDe torceps. !he ani..] II1se ~~

which oovld be dill8M'tecl ... a-. .An aq~eou 8Olut101'1 of ..-t.b;rleneblue .. ut1l1Hd. to 8ta1n th. u.••• pr1ol' to ci1.nct1em.

Jaclul...... .uatecl 1d.th~ on .Ud.. tor pe1'!III.MDt ret...ce.

J4nMa 111_ aD ...s. 1IU pzooclucecl1n aD aquari,.., it wasiwed".~ ,en-lMd tor on. do... am count. UJd tlum lI1UJPC¥led

b;r tbNad in a beak.. of boned, tUtwecl •• _tezt 1Ih1ch as

OO1'lt1mle l';y ....te4. AftR hatoh1Dg, the larYae were traDaterrecl

to boiled, t11terecl ... _te Dt. pipette .t replar iDter.nla.

Tl'anst... ot lana. ~ fIfV7 HOODl dq .... tcnmd to be nttia1ent

in~ 1cmg 1arnl 11.... ami pod larval 8lU"9'ival.

! -

6

~ specie. of the superfam1J.1' Doridaeea 8i'e desCl"ibGtl

:f'rco1 :touzo loeal1Ue. in the central &D1 nat-centra1 PacU1c

Oc9an1 Bawa11. tilNtok. PaJ.1IlJ'ra aD! Johnston Is',"",. l'ort;y-adx

or 'these specie. npnaent~... oolleet1cms bT Dr. L J.liaonI7q on the 1slaDl. o:r Oahu a:l En_' clv1rJg the put f'ov ;years&lid by the author and Zel10v stud_dill of' the Zoolog:r~

of' the um.....l1t;y of Haw.,., OIl the 1l11.aM o:r Oahu dur1Dg the put'two 78&1'.. 'len species o:r clor1d8 .... oollected 1n Ib1wtok b;rthe _thor d.v1rIg a tbrM-__ pe:lod in Ja. &Ill Juq, 1965.

bIlt em:q toe ot these speci...... d1tterat trc. tbG.. ool.lected- or' .

1D 'Bawa11. !wo spec1aen. of on. specie. (~11.8. 2) ....

coUecJtecl em PaJ.Iv'ra 111 J1Ule. 1962 aDll 011. 8Pecd.Mn of a ..,.01••

(~.Dign), a1IIo collected 111 Baa11 8Dl .m.wtok, wascollected b;r Hr. W. Van l18akel_ 011 Johnston IslanJ 1D jupet, 1965.

!he piIIpOII8 o:r the tol.1D'tdDg species dnar1pt1on8 111 DOt~ :tor spec1t1c 1denttftcaticma, although th.,. a;y~

be u.ed as _ch, ba.t rather to uh1b1t the d1ft'eNIlC811 and ",,"'.n-Un .-mg speci... 'to attain We objeot1ve as IIUV' ~log1cal

characters, bath Btemal aD! 1ntemal, were ..mined as could be

c=w=1d~ ut1l1zed t.Qmt!"!!d~i e.nd features o~ the Ita lY!IMIJ

and l.&rne were 1Dnat1gated whenenr pollll1ble. !'oJ" k..,.. to the

1d8ll'ti.n.cat1cm o:r species, list. o:r 61JWlQJ11.. and di.culd.cmaof habit. ot the Ba_11&11 dOI'1dIl, the :rude 1. Nf'erNd to the

pape:r bT Kq aD! Young (lIUIUIIar1pt).

7

desei'ibid'tw6nty-two spGCios of dcri.d=~ the Batm:11an Maude in

the 19th oen1iuz7. wee ut..U:1sed tor the Ba1Ia:11an dor.1ds. Ibllt

or P....•• DUl8. haft 'b.m either ip»reci or~ l1stecl &8 ·possible83'bOJVIIII in the pa.t litR.tve becaue the 4uOi'1p~..... 1ia8&d

eo1eq on 8EteIrIlal obaractere. the oolleot1ou upon wbich this

s~ is~ baRd~ ==lo in hi= twe loeal1Ve l't 18....... baa this that ftZ'1at1orl8 ec ~ toaDd .-mg dor1da t1'OJIl

d.1tt..-t 1ocal1t.i.. an mm.l•. Because ot the Nlat1ftq'

aaltencl Datve of' 'the~t. one ...,~__

that the ... doZ'1d 8)*I1e. are p-eaeat .. 8U11Plecl 'b7 Pease. liB.th

these OODIIid.erat.:1oJl8 1D JdD1. the -.j0l"1't7 01' the &pecd... Pea.e

denr1becl in 1860 haft been ftI'1t1ed..

P... and. other earq 110ft... used the I"rmeND .... 1lirU.

tor tbo.. DUd1bnnch. with oval. doraoventraJ.:q depl'U.ed bodies

aDl g:i.lls encirc11Dg the posterior &mUI. ,DorJ.1 has a1Doe been sub-

divided 11'lto • JmJIIbe1" of' gen... 'baaed on 1Dtemal autoJd.cal

character.. lite po.a1ble. the species are us1gaeci to pzr.4l.

&8 orig1ne]'7 described. or subsequ_tq- _Iuled 'b7 the 01'1g1Dal

authors. AD attelpt,·_~ lIIde to d1....n1ah 'the 1IIportance f4

the n:pertluou. character. aDd to tlllphasize the oharacter or

char.cter. which .ppear to be milt diapoatio. tor each pmlB.

'lbe tol.l.o1r1lJgchanctere .........nc in mat apeo1..1 (1)

abape; (2) s1d.D 'textuN; (3) color; (4) bZ'ancb1ae; (5) ~N.;

(6) oral tentacle.; (7) radula; (8) jaws alld buccal Ull&tuN;

8

Im1don, Eng] and.

An obv.lous problem in color descriptions and one that :18 DOt

ea~ rectU1«1 8ftID U 00101' charts are utUiud is the subjective

variation in color patterns noted 'b.r the observer. As stated by

Croasland (19U), • •••what to one obaerver seea as black baD:ls

on a white baokg:rouu:l i8 to aDOth.. white 11nea on a black bocV'••e·.1!:ds pl'Oble was part18117 resolved by' recognizing the pNVa1.l:l.Dg

lightest color aa the baakgrouDd upon which darker pipents 1'oN the

ooloration patterns. It it was poaaib1e, d1.t1'erent s1secl 1Dtivi-

duaJ.s anIl spea2J1MJ8 :r.roa d1tteNnt looat.1ons ......-"'neel ed. thevariations ..... tu:_ into account in the 00101' deser1pt1em..

~I :rn all ba.t one apec1ea stucJ1ed (geM'" el!pD!)nocm:lar7 g:l11s CCBtOnq tened lIbrancb1a.-, enc1rcl.ed. or partialq

8Qr1"OQDIied a m1d-doraaJ. 9U8 which 1fU uaua1.q posit1onecl. on the

poste1'1or hal:t 01' the amma'. Because 01' the 'VU'1at.1on 1n p1ace-

JI8l1t aD! atNeture 01' the brancb1ae, thq are often used. as ident1-

f71nI characters. l&ch bl'mch1a is composed. 01' a r"ch:1. :t1"o1I. 1iIh1ob

p1Jmae arise lateraJ.q in a cbarac'te:r1atio alte:rDatiI1l'in.-qUence.

In _ clor1d. the hranohiae are retractUe into sheaths which mq

have spec1t1oaJ.q oharacter1st1olbapes.

~I A s1rIg1e pair 01' tentac1es, temed. "rh1Dophores·,

are doraalq positioned. at the anterior portion 01' the am..,.

Bach rh1nophDre 1. character1st1calq OOJIIPOsed at .. pec1unc1e 1Ib1ch

ie t.1pped with 1'o1iaoeous '_U.e. !he 1'h1DophoNS are CO'WOnqretr&oti1e into sheaths whoae appearance and strucrtu.re JU¥ be

apec:1:t1caJ.q characteristic.

10

anter1or~ bome wnt.ral. to the mantle and dorsal to the toot, one

on each s1de o-r the I1DUth.' the presence or absence and the shape

of these struet1D'es arecolft'!il7 included in spec1es descr:l.pt1ons,

bIl't were el'minated in this~ wiless the,y 88€iii8d d1agiiOstiealq

s1gn1:t.lcant.

Diem],: Pr;)babq the con:d.::tentq most 1m,Portan't characters

in the Doridacea 8Z'e the lIWIbers, ~ts aDd shapes of' the

rad12lar teeth. EvE whe they are absent (e.g. J?!!ndroder1.,>,

th.., COIIPr1.se an iIIIportant nega'Uve charaate. 1h. radn'·r teeth

are 'boae on a rach1lar ...abrane which 18 stretched over a tempe-

like odoDtophore (see Panot10DaJ. lbrpholog,y of the Feedhg

Apparatus), am are podt1oned in right- am1e:tt-haDl :rows in a

l.cmg1twUnaJ aeri_ ot regularq recurr1Dg mDlbera am shapes.1he regular1tJ' ot :nabers aDd shapes ot the Ndular teeth are

ut1l1secl ta1rq p1"8C1seq at the speo1t1c aDd. generic 1eveJ.s and

JIm"e~ at the tm ]1&1 1wel as taXODOJl1c charaetere. As

the radular teeth varr 1n sis. &Dl shape within &rIT given row, it1s necesAl7 to 8XlJm4 ue all the teeth in one row. It is cutomar.Y

to take tooth counts and lIle&au.reDel1ts troa that 1'OW wbich· is

-?d-"7 de9'e1oped~ 1!~ about one-thbd of the distance troa

the tip of the redD1ar Dl8lbrane. Bow coUnts are -01-1, 1nel".,.,ng

&ll tho.. radU1.ar ron which are d:1st:1qa1abable. IDv oounts &Dd.

tooth onnts ft1'1' with the aiH of ani-1 ~ liumever possible, th....

tore, the radular teeth ot d:1t'.terent sisecl~ of each species

u

within the radular ..-bran. am lxAmded on both s1c:les by lateHJ.

teeth. JV' ocmv~t1on, the largest lat8Z'al tooth is ce" eel 'the doll1-

lW'lt tooth. It the lateral teeth 'ft17 sudd~ :1n shape within

a g:lft111"01f, these teeth are of'ten temed the urgiDel teeth•

.As the radulaZ' teeth of each right- am left-hard :row are mirzoor1iiages of one mothae, a for=Ua 1s oo~ ut1liz= to =p~a

the raduJ.ar ccnmt. It the raclddian tooth is repNScted by' .-,

the mmber of lateral teeth by' -L-, the namber of UJlg1Dal teethby' .. am the IJUIIbco of 1"OV8 by -., the redlllar fomala~ agiven zwdul. 1IUI11d be repreaentecl b;y:~ In all the

dor:lds :in this 8'tucV, 1'10 rachidian tooth ... found,. 80 1t 18

reprea_ted by' "O'! :in the folWlla. 0Dq:in the g_era ,,","e am

fM's-zel!..,. wre th... teeth which. ocnWi be interpreted as

UJlg1Dal e1818l'1ts, 80 the rwm'ar folSlla of the ujori't7 of dorida

is ...,. "'''1>18 and. represented b;y N x L,O.It. lIJ.eenr the radule

of several speciaens of a given species was .....ned, the. rage

:in ron am toO'th counts ~. included in the foNDla.

i!D. !IIllBooal Aeture: Cu:t1cular th1ckeDinp :in the lumen

of the buccal uss of dorids are often and pel'haps untortuna:teq

te1--ed -jen-e '!he jat!!! eq either be in the f'o!'!! of' laterU

plates or horseshoe-shaped stnctures (see Functional )l)zphology- (

of the Feeding Apparatus). 'lbe position and. tJpe of jaw is an

iIIIportant Q'Steu.t1c character as 1t utnI&l.q parallels variat1cms

12

~~t cht:.raet=- in d1~"'U1shingbe't~ ~B~

Bc»me Upon the jan are cb1t:lnous el-.nts otten termed the

"tmocal umatmoe-~ !he!! shapes aDd si~.~ of theae stru.ctures are

1mporttmt addit.i:n character. au:l often 'ther &1"8 near~ as valuable

~ca.J.ll'&8 the radulai' tNtb thEiiiS&lvas~ 1hG sizes of the

buccal ansat.ve varr~ with the s:1.ze of awl..1 &rJd positionon the jan, 80 the raDgU or size are a1waya given 1ih8l'1 8)1"8 thanone. spec1Jl8l'1 1. ,,"w1ned.

GsmiW ltYI1 Although th.., have been all too 1nfloequmt,qu1;111sed •• ncb, the cmpmmta ot the gerd:tal ma.s aN 11IIportant

characters in dor1d~. As all iiiiiii....... ot the Dor1clacea are

bemapblliO'H:t1c, both -.1e and teale caponent. are COIIP&ct;q

a!'1'Uaered in • a1ngle NPllOCiuaM.ve qat... 'lbe genital mass is

cbaracterJ.~calqon the right side or all dorids am 11e.V8I1'tnl to the mantle am dorsal. to the f'oot~ xt OP8DIJ to theeatter10r t.JutoaP • sctal or11'1oe. !he goDad. are an&11pcl1n

COII'Oslte stzucture. termed the ttowteste.- (etcept in gwIa1!

!HeWe =0= .cw.r1e:: and testee 1'e!!!!.1n separate) 8Dl an 81thw

spNid ave the d1gest1ve glaDcl a. a thin lqv (e.g. m,oodo£!.J)

or coq»acted &8 ovate bodies (e.g. GYwNdor:it). 01'1q' when the

ovotestes tom discrete bod1e. arethe.v uaetul as gross taxonomic

characters. 'lbe owtestes are COIUleeted b7 ducts which join at the

"hermaph1"od1t1c duet- which leads, in tum, to the "ampalla" atthe genital JllUS. '.lbe cluet~ the ampulla t01'llll!l a bifurcation

with the male duet s;yst_. COiWipm1.,. called the "Prostatic duet-,

13

'shere t.~e i~ 3 ~n~..~=:!:t~ va,1~= E'h'" ~'i;. p::re==Q'h~ by d1tt~en,",

tial tmBeular cont.1'a.etion shunts ova into the aJ..bw:d.D and ImlGOUS

g1&Dds and. 8J)81"lIl into the male duct qat. (MacGowan am Pratt,

1954; 1hoJIpson, 1961). 1he albmdn arxi _COIlS glands 1'ON a

CO!!!POsite stncturo and. aN usuP"y- 'treated, e.g. tlds atud1',

as the albumin-allC0U8 glard. .An ov1c:luct passes :rrom the albun1n-JIIlOOU8 g1aDi to the gerd.tal or1t1ce" 'lhe male duct qetem is

comprised of' a prostate glaDd, vas deferens aDd ejacalatozor duct.

In certa1n spea1es the proRate glarJd 1s not !'8p1'e8ented b7 a

.consp10a0us swelling or gland am 18 not grGssq recognisable.1he ejacalatorr duct 1s e:lther a JII1scalar penis or a simple duct

vh1.ch enrts dur1Dg oopulatiOD. ltfman (19.51, p. '19) utilizes

the term ·ai.1"1"Us· f'or ·~••a t.Ipol'&l'7 projection that 1s 1'onaed by

the turm.ng inside out of' the ejaoulatoZ7 duct·, am it 1s suggestedthat this tem be 8dopted 1'01' donds possess1Dg this t,-pe' of'

ejacalato17 duct. Often reaurnd .c1rral hooks· line the~

~ the c11"l"U in ncb a wq that the hooks project anter10rq

iih6il the duet 18 :Ntr.eteel and polJteriorq when the duct 1s

everted.

!'he oopula't1ng pair position themselves 80 that their right

sides are opposed, aDd the ejacalato17 duct of ODe mEer 18~

into the f'-.1e duct system of' its partner_. _ 111e 1'8I&1e duct qat_

1s cospnsed of' the "ngine] duct- 1Ih1ch luds to the hollow, thin-

_]le ·apex_theca·. 1he sp....theca e1ther CODDects with the

·sp.....toQ7St·, • hollow thick-walled gland, or the sp.....tocrat

~ of the ,permatheca aul spermatoqst is often produced

by a -spermatoqstic duet" ccmneet1ng the spemato~ to the

female duct qat-.. ~e "uterine duet- pas_. 1nto the aJ.bmI:1n..e1oousglam at or neal' the junction of the upulla aD:1 P1"Oatate glaud.

Becaue of the variab1l1V betvMn species aDl the constana,y with1n

each species of the female duct s,yst=, the relationships of e1zej

shape aDll U'nDgeMDt of its OClIIIIpODents -.a:T be ued &8 l'011abl.

tamDOld.c characters. 1\1e OCIlIIpleac f\met1cm1Dg or the reproductivea;rst- dur1Dg copala:t1on, tertU1sation aDl ov:lpoll1t10n an deacribecl

by KcGonn aDd P1"att (1954) tor 'aob'dor.1.' •1\1e ...ca"ed, -acOO88017 g1.aDls- aDl -n.t1ba1.ar glAnds-

are often f'0UDi in C

~1.e.t~ sithll: e epe-~~s i_'I'! S~~~ ~~~t1@n ~ c@nfo=ati€m

ofcgg =::se., however, ·features such as the~ and d".esions

of' Oft am larfte are otten _re valuable taxol101l11calq. thesecharacters are DDt cmJ.¥ he'JpAUl in dist:1nguish:l.ng between species

'llhich aoe d1:t1"1vul't to :14&nt1.ty iiOIPholog1caJ.q, bu:t~ u=tul

hdices for detend.ning baa1crelatioDShips anci- diRinot1cms

betJlee:u higher groups as wello

1'he genus Pei"1opsis, which _is exclusiveq JD:lo-WGst-Pac:1fic

in d1stl"1but1on, is apparerrtq CODIPr1sed of~ three species.

In this st1Jq. n.. pecten was found 1n both Hawaii and tiwetok,Whereas ~ y1Jjlci1, was em:q found in ihiwetok am Ra., grang.1oeonq in Ha1l811.

'lhe gGmls porigpSsis charactc'izod ma1nq b7 its distinctive

'transverse plac-.nt of'· branchiae in the poster1orq posit.1oned

crescent-shaped branchial aheath.:BtnnmIr, there has been IIIlch

ccmtu10n in tAe ident1ticaUon of' species.' JD)DB the tJnoee

species of' Perigpsi. DO utemal clitt..-ce. other than coloration

aM ffltr spec:l:t1c 1nterna1 characters are ob'dou. POrioptl!

mmiJp. probabq baa at leut fi.... diUerent. 81JIOJVII8 1IbDae

va11cR't7 was 1'irst questioned by Baba (]961). On the other ham,

~ Peets has been COIIMnq aal!RDlled to be & color varie1i7 of' Ra.

iJ.D4H (11&bec, 1928, 195'; U'eD, 19471 Baba am BulataD1., 1961).JdJIittee:Jq, i&. peetS and n.. ;x;lr;14i' a:h1b1t si"']ar ovoid bod1'shapes,~ pirmate branchia., rod-l1ke Z'h1DGpboNa a1'Jd tpi-

dezomal spicule., a. nll &8 overlaps in zwtn'.r tooth counts, lizes

of' radular teeth and mmiber 0'£ branah1ae. R&. grpgloe also exhibits

theM "''''''&r1t1es but apparentq has f'ewer and l.argV teeth than

the other t.1IO species in simi] 81' sized apec1Jllens.

'lbe shapes am forms of' the radnlar teeth appMr to be on.

of the mo1"8 diagrKurtic intemal character. for each species•

.Although the genital maS88S are simi'ar 111 Pl'OPOrt1CmS am~ts cd ducts am sJaMS in the thloee species, the COJIIP&r&-

17

to be of GOmQ valu" as c&Utive sprv1f1c chaz>acters. '.!he sizes of

ova am. capsules appear to be aharacteri8'tic 1'or each epec1es inthis stwV, however, comparatift data for Ra. !I1l'1ti. (=ia. pecten)

ancl L.. 'Set1fa (iil~ 8if!ii!l9p> in Japan (BSaatan1, 1.961)indicate tbatova au! capnJ.e sizes poas:l.hq var.Y geographicaJ..q

and hence !l!q be poor d.1agnostio oharactersoOne ot the~ diapost1c charaoties in this atucV 18 the

se1ect1u'ti7 All tooclbabita IIXb1b1ted bT each spec1ea invest1gatecl.

Bach 'R8 tom! to ut onq a single species· ot sponge and would

not te«l on art8 other speciea of sponge - enD one 1Ib1ch COIl-

prised the food of 8DOther species of POrtes,. Jdd:1t1cmalOOJIIIIl8Dta on tood w:lll be defeNd to a tollcnr1ng section concerning

the food ha'b1ts. J:t .... 'iIO~ to call attention to the tact that•

the color of _teD spcmp 18 often us:l.ble in the di~ft g1aDd

tJuooach the ep1dend.a. 1'b1a 1IpIrts micl-dona1q a blue hue to

~ cpm'ee and an orange hue to »a. Jirid1t.

Internal-

18

Hawaii. - Oahu, UnS; (CODIIOn).

See F1gare 1.

Shape oval-oblcmg; legth to 1011I; oral tentscles..u, lobUoi'iii ep1d&fiial sp1

lI'Jm! ~DD~~P..., 1860

.&e lateral TiW'l-"~_••B. IAtcal d .. of thG 1~.lt rcnr of nd.v1.r teeth.

• - apulla.... - .1.....» aDrl ..... aJ.aDrl... - .j.aalatol7 act;hi - h. I pbl'odi:t10 act;od - oridilct;pr - PJIOatate gl4m.0- ....toaptst-.....-.thecav - '"15M

'9d. - ft. 4.reNll8

·.hd

---I

!I

•

d·m

-------~---_...._--------A

-.

~ --v \j--

48

O.10mm1__•• •• _•• _. _. ._•• _

B

20

Internal-

Igg mass-

Hawa:i1. and Iihiwetok (OODDDn).

See Figure 2.

Shape oval-oblcmg; length to l6mm; oral tentaclessmall, 1ob1to1'll; ep1desal sp1c:ales ~l1k__arr&nged~ in~ alternate ariesa10Dg m1d-dorwal ..11 spiculate pustulesappear as -grarmleaQt over dor3Wllj mantle amteet deap blue; bn.nch:1ae a1mpq p1rmatQ, 7-16,deep blue; branch1«1 sheath crescent-shaped. amtr8D8'le1"s8q positioned at posterior -. of~tle; rh1noph01'88 :rod-like, 8-l5 deep blue'amellae~

Radn'er teeth :30-32 x 28-42.0~28-42 (in 8 am 1'"spec:1mena), 46-lo8p 1DDg (:111 10. iDd1v.t.dual), allhamate, aD! DOD-d_tiCQ1ate, inareaae in s1uoentraJ,q 111tb1n each reeJu'er 1'OW; DO jaw; DOc1rra1 books; sp....toev-st ovat..eJ.cmpte -slightq J.a1oger than ovate spezmatheoa; parallel~t of sp....theoa am spuutoCJ'st;uteriDe duet enter. a1baId.D and II10fta glands8!'.G'ateq 1'l'om~ am P1"Oatatio ducts.

i'gg masa I"1bbon-Jike, .tree border 8IIIOOth, up to4 ~r1a, 90 x O.~I ~teq 128 ova7ra2;ova l1ght yelJmr, 80-8~ in diameter, each OVWllenclosecl bT oap8ll1.e llO-l.2O,lin lcmg d1Iaeter.

ContrarJr to the l:1st:lq of Ra. p!C!j:s as a -color

varietT' ot Ra. Y1!dd1, (l18bec, 1928, 1953; Al'en,1947; Baba aId Hamatani, 1961), the ftm~ngs in

tJds stud;v bJd1eate that P... peets is a valid

species distinet from R.. x1r1d1,. 1he blue colorationof l!L pecten is tberef'ore to be ooruridered a d1ag-DOstie spec1:t1c character as orig:lna1'iY described

bT Col.linpood (1881).

J'J.Jr!ID !..Jl!r1!p!i,~ (Q)l11rJpMMl, 1881).

A. Iat.NI' 'Y.t_et~...tb.. aDr1 ......~.& I&'kral -.1_ of the 1ett-ha1t NW o-r zwlwl.r 'tMth.

80- ....to.,.,.tat-..,..-thecalid - .t.er1De daut

T - vag1Da

~ --V--42

~t

A

O.10mm

B

-_._---,i

-- C)]--V

--------_.-

"

DESCRIPTION:

mtHARr.S:

See P'1gare ,.

Shape ove1=oblong; ltmgth to 8!!!p or81 tet&a1e~..." J lobifom;~ ap1ealeii rod.l1k.....arr&Dgec1~ in 1'0118b1l' a1temate serieaalong BElcl-cloztsaa; aall spical&te pustule. appearaa -srazm1..- over clor8UJll; JlllDtJ.e am toot olive-green to lmmse-grec; bnnch1ae mwpq p1nDate,7-8. ol1w-peen to bl'oue-cre-; brancb1al sheathcreacent-abapecl azd trml!"1Vseq poe1tioned. atpo8ter1or end ~ untJ.e; rh1Dophorea l'IOC1-"k e, 6-9011"fe-P'M1l to brcmze-green , nellae.

Radular teeth 26-30 x ~26.o.24-26 (in 6 am 8Daapee:m-.), 46-J.l9J11Amg (iD e- 1lld:l......,.),all ~te ud non-d_tiaalate, iDc:rN8. in e1zecent.nJ.:q 1d:th1D each JWhiIar ZVW; DO jaw; DDc1rn1 ",*a, .....tocvst Oftt. 81071p:te--••']e· thaD spmwath.., parallel~of spematheca aDll ....toGJ'llt; uter1ne cluctenter. a1ba1n aDl _ooa glAnd s4IP&Nt;eq flooaaJIPU1larT and prostatio ducts.

Ba ....a r1bbon-l1ke, f'Ne bozder 8IIIOOth, ~ to, whorls, 60 x 1~S-; appl""'d-teq 72 Oft{-.2;ova l1ght yellow, " 2-12Sp. in d1...ter, each 0VWIlenclosed b7 capsule J.SO-16~ in long c:l1am8ter.

'!he _results of this~ 1DH.cate that it. !j1r1di;

1. mol'phologioa1l¥ am ecologic:aJ.q d1at1nct tz.oa Ra.

Peets aDd ehou1d stand as a d18t1nct am a~ate

species. , HClce, the green coloration of Ra. yiric:l1l

18 a spec1ee-apec1:ri.c character as origit1.1 liY

described bT Pease (1861).

,

1!ml!p1il.~ P..., 1861.~ I&twal ft_ 01'~• .,....theca ad Q __toqat.& I&'te&l ft_ 01' the lett-hal:t l"OW 01' n4111ar tMth•

•o- .....teqatR-..-th..wi - ut.m.u duct

T - T&g1Da

•

st sc

'------------_._---A

26

O.10mm

B

,

L

me genu Doriorbji, is ereciied for a species described fi"Wil

Hawaii. aa PRril. t1pc1it b7 Pease (1860). ~ the gc11s iscbancter1zed by a Z"lgld, acabzooua mantle covereci nth aall

1rNgalar papillae and ap1.ealea am with a '1' or Y-shapecl Mdial

atNak Btend1ng 1'1'011 the rb1Dophores to the 1II1d-dor.... 111. rh1Do-

phoN. are rod-llk. and t'ineq l...,l.te mel the 5 to 6 branchiae

are a1JII»q pinnate and a1rea1arq arzuged abou't the poster1or~

poait.icmed amuI. lbth the rh1!iGphores aI¥1 brancb1ae are ret.r&c1;i1e

into sheaths. ~e redular teeth are~ bua'te and ...wi].r to

those of pPr1gp!i! _cep't tha't the Outer 6 to 8 la'teral teeth have

several Pa]' outer d.enticle.. Otherwise the a1.bIca'tu7' 'tract

Ghibl'ts a ,,",""ar buccal maas, a large m:iclgut, an e1cmcate caeaaa

PJ.'lOject1ng mter10rq boca the 1et't ventrolateral aide ~ the a1d-

gut aDi a pair of elcmga'te sal1var;r glama mending f'l"OJIl the

blecal ... to the ante1toventral end of the d1geat1ve g1eD:l.. aa

m POr1gpI1" 1he gezd.tal maaa 1a almost 1nd1a't1Dga1abable f1toatha't of the three specie. of pPr10paiL partiaularq in the para']el

arraJJg8Ient of the sp....theca arid BPermatoo.rst in the f-.le duct

qstem.

'!he most iJiportan't charaoters of the genua are (in d8CJZ'M8ing

diapost1.o 1JIport.anoe): (1) s1JIIpq piDnate brmch1ae arzuged u a

c1rcle't about a poater1or, Jdd-doraal anus aDi Ntraotile into a

a1rea1ar sheath; (2) hama'te lateral teeth vlth ou1ierlDat laterals

denticulate; and (3) Yor Tr-ahaped aed1al atreek mending f!'OJI the

25

!ihG gGmlS is »Q;led for the chAractwistic cd.raalar arrang6ii1Gilt

of brancb1ae (0#1= c1rcJ.e), l1h1ch is in contrast to the pnmcalll'

diagnostic character of the CZ'escent-like arrangceni; of brancbiae

in the genus POriopsi'.

0CCtJBRENCB: .Hawaii - Oahu, Xauai (6 specimens).

DESCBIP'.flON: See F.i.g\u'e 4.

lkte!'l'.Al- Shape oVal,~ 7-lS x 4-6am; mantle rigidand scabzoou with _" ift"egQlar pap11lu d·spiaalea; mantle coloration b1'own or grq-bluewith If or Y-ahaped 7el1ow Dledial streak c:temlingf100a :rh1Dopho:res to lI1d-dornm.; toot lxi'ab1ate;brancbiu mmpq phmato, 5ao6, brown; b:rancb1alaheat.h with fflt1 papillae; rhinopho!'es SIUll,' ••" •• tine, apex: aCUll1nl.te; :rbiDophoNl. sheaths1JIPl.l' stellate.

Intemal- JadD]ar teeth :36 x 42.0.42, S3-105ploDl, allhaate aDd. acuteq cm"t'&'te, ~st 1&twalawith f .. outer dct1cle.; dOwtrwnt tooth iscentnJ.1l' poe:lt:1oned. w1tb:1D Ndular :row; ejaea.-latoZ'1' aDd. vag:tDal duet. aepuate; M .a1r.NJ. hcoks;vas deterena abort aD1 loos&q OOJm)lute; p1'OatateglUKI ..au .m gooth; sp-.atb,eca.qvate &1¥1 •81~.gb:tJ,,· ..u. than spherical 8pU11&toGJ1l1i--both are cozmeoted bT short ducts to the femaleduct qsta ill • panllel. U'I'aIJI-.nt; vag:1NJ.duct is lDDg UIl 1oos~ cmmtlu1?e, s.Pez-thecalduct is long and looaeq ecmvol.ute; uteriDe ductenteZ'a alba1n aDd. -.ooua gl.nd at junction ofampulJ.ar,y aD! Pl"Oatatic ducts.

-,

lI'ilpn !t-PIl1!!b1f Mp* (hue, l86O)

Ae I&teal"l'1.. at thepld.tal. __•Be I&teaJ."I'1.. o~ the.~ l'OW ~ the radDJ ·r teeth•.. -..-"..... -.'.....,n ad. _GOa_ glam

eel - ejaaalatorT _crt 'hd - h. apbrod1t.io ductod· -n1cIacrt .pr - PNRate sJ,aDl80 - toGJWtst - th...

V -..,.

ft - .. d.r....

__pr

,----'-------- ------_.__._--A

1

O.25mm.I __._~__.__ . ~------ --_.... _... --- ~ .- ---- --------- ---------B

e~:!s A:.~~~rio ~~ 1S?8

&rgh (1878&) established the gems Arghidpr:1s.on the. basis

o£. the idea that the IdJmaean genus I!2£iLI. vas insutt1c1en1i:q dellcr1bed.

In Bergh' II description of Arcb1d.on. JIIUV' characters are shared byother generic gl"O'U'p8 P08liUalilg s'=118]" convex bodies, cr.YPt1c

l'b1nophoN8 aD! braDcb1ae, gr••'er to tuberculate mantles,

DI!!!le1"OUS hsmate teeth and unam.ed gen1tau&~ 1he oriq character

wb1ch appe&Z'1I to be ~ d1agnostio value tor the gezma 18 the preamce

of a thick cat1cle at the bucoallip all des1gDated 'b7 Begh (1878&).

1'- 111 sagpsted that tid. generic group should. contain tho.. dor1dII

which tit Bersh'II geeral characters tor 6rcbidori. but wh1ch &lao

uh:1bit ba.coa1 pJ.ate. 1Ib1ch are devoid ot consp1caous &1'JII&ture.

1I1e blecal platell ot the two Ha_Ha» 8J)8Ciea certa1rJq are -ca.t1cular

tb1aken1.Dc.- in the true .enlle otthe 110M ami. both are devoid of

amature erz:oept tor Ia. nubiloe wldch atdbit.~ JlliDute granula-

tions.

DFSCRlP'flO!f: See F1guoe 5.

Kdiernal- Shape 'b1'oad:b' oval, convect, no ::It so-; mantle fair:qrigici with· ..n rot:Il¥iecl papi" ••,~ tb1nand~; -.Dtle ooloration 1W\..bJ:vIm withpatch•• or cla1'lc pipl8Dtatlon and II1Jmte 1Ih1tespots ocellatecl with black-~.. are whitemth :sauve blotches; toot Dite J!!D'ttl.erl1!1thP1I1'Ple 'brnD; branch1ae teather,y, 5~tmown; 1'h1Daphoru e1&mpte, 9-10 , UMll••, ~bNIID 1d.th white tips, I'hiDophores sheaths IIIIoothedged.

Intemal- JtadDl.... teeth 22::1t 48.0.48, 1O~ lag, .nhamate 1d.th aCRteq' cmnate cup_, d"B2D1lJ't tooth~ po81Ucmecl w1th1n ndul .... row; lateralao.tioalar plate_ 1d.tbaut awatue at baocall1p;OOWt'D or1.t1ce .urea b7 ejaaalato17' aDd 'fti"na]duata; DO cd.Zft1 hooks; .,.. clef.... 1Amg UJd~~ 1Dto~ lobalatepzoo8tate g1uMl; .-:1aer1U.~t~ apema-theca ad ......tocpt; "I'D.] duct loDg and non-ccmvol1lte--lMd_~ into theca;spematoG,JWt1o duct _l't-OODDeot toc.ratwith IIJMIlW&thecal cluct; ..,....theca ~er1calaDd ove 3X 81•• ot ap....toqst; uterine C:luctshort snte:rs albmdn mi moous 1'."" at jlmct10nor apaJ.l.arr and pzoostatio du0t8~

a -...-uaa'" - a'lbaldD &DI1 gledeel • ejaealaW17 daa1;1:d ... h=~Uc d!!etod-- m.cIaGtpr - pa.tate .'am.0 -.....tec;rRat - .,..thecaT - ftl'M_ ..... ut....

_ •• o· _.'_

A-----_._---

hd__~

l',,

5t

i'. \

'---

- - __0 __---l

48

O.50mm

---,------- -- _ ..- -.--_ ..._-------.--B

..

Intemal-

Haliidi - O&im (4 specimens).

See Figure 6.

Shape~ oval, CODVtIX, 110-190 x 85-J.S()Da;maDtle &iO:tt and CGV6&6d ii1th clDae:q..m tmequalqaiHd pap1llae which are cone-11ke in appearance,marg1D thin am UDC1u1ateci; lIIIrltl. _ttled bro1m.ancllNT; bNncb:lu trJp1Jmate and teath.,-, 6,~ postmorq, p1m1ae light hrotm 1'I1thda1'k b1'o1m V&iDB, :rach18 wh1te;~ club-11k., with mIIlel'01I8~ laM"•• ad whitetip., retractile into ftneq papillate aheatha~

ltadular teeth (in 2OO.a~) 52 x 120.0.120,l2O-42Op hook:llqth, lSO-DOp tooth height, allidIIpq haute, dmdnant tooth catralq positionedwithin ft7W1 lateral j_ with DO hook.....atb.....Iigh'tq raised gnmUation. appear1llg .. dono-ftI1'traJ. ridps; ca••m .temal or1t1oe 8har«lby ejaealator.v ad ftIS.nal act.; ejacnilatoZ7 duct1d.th ..calazo ptD1. aD! extzo1D81c ..~ attach-m.errt to "anterior~ wall; 'ftI defeNDS 101JgaM COD901v:te; pt'Ostate glar:d abort aDd coUC;vapnal duct short aDd broad.. leads d:1Nctll' 1ntoelcmgate eauap-sbaped sp__theca; ep__thecalduct short and COI1'9Ol.ute--ctera junQ1;:1on withuterin. duct am epeNatoe;rat1c duct; ..-to-eyat.1c duct abort--lead. into ooaraeq lobalatespematoc:pt; uter1ne duct Bhort--opens intoalbnd" aDd _00118 gland sepan'te:q :t.roa junctionof Pi'Ostat1c aDd apallary ducts.

M!bJ.u7' !!!bP~ (P•••, lB71)•.l.. I&teal.Y1_ th. a-t ....B. '. I&tenl ft_ ~ thf'l 1e:tt-ha1t Z'OW' or ndular tMth.

a - upallaa'" - al.....n ad .ona sladeel - .jaC'llatol7' 41lGt

epa - tldi.r1Da10 pMdal _01.lid - h_.puoc'ti:t1o satocl-oridaatPI' - Pl'Oltate cJod.0- ...to....-at-~th..

T - ftI1D&'9d - 'ft8 det....

·_.~

.------------._-----_._--------------'A

----------.- ._-'.'-' -- ...'>--_..._--- ----------_·-1

. i o.SOmm

B

.'

,,I

All the characters of P:l;atmJgr1s as ongina11'1' proposed by

Bergh (1877&) with the exception of an a1"JII01"ed vagina are shared

'b3' the two species at P1alJder1s f'1tom Hawaii, however, one of these

species has a ou'ticular-lined vagina, a f'eature al80 stated as

characteristic of' the genus by' Bergh.lJEbera of Plat:tderi! appear

to have a more rigid am Dd.nuteq granuJate untle than otherjaw1ess dorid nudibranchs, but the most diagnostic feature of' the

genus is the presence of a in large books with disk-Uke bases

in the ejac:alato17 duct.

OCCODD:Ba Hawaii - Oahu (2 specilB8Ds).

DISCBD'fION: See Figure 7.

BI:temal- &ape bro8dq oval, a.poe.seci, 65-75 x 3l-5Oiiii;mantJ.e risid aM t1n~ gNJ;iiilate, -.rg:lh thinaD! ftl7 1rrepl.ar; ~e 'colont1on 'IIb1te orareal with tin. 'brown or black speckl.1Dc aDd largeorap lDtt1.i:.ncs; f'oot. DaZ"l"OV, 1Ib1te 1d.th 'brcnm orblack speck]1"1; branchiae tr:lp1mlate. f'_th.,..~4-6, tNra8paNbt 1Ib1te or creaa 1d.th broIm 'ndn1ng,scalloped edge to braDM' al sheath; rb1DophON.c1ub-

"ke, white peduDcl.e., orange lame]1.e, pl'Olli-

nent. aul .ca1lDped sh..'th8~

Internal- BIdn'.r teeth ~.:It 109~O.109 (111 6S- speo:1lllm),:~30~,1cmg, all haa'te, outemost 1&t.eal8elcmpte with cmq tip. OU1"9'I:te am JK) PftW"ne-t;f'1up a8 1m.. laten1., domi"ant tooth centzoalqpoll1t.:lcmed 1d.tb1I1 each redaJ.r :row; n.n.q lobalateg1aD11 a1; baH of' ejaealator,y cluat; c;c on.-telrulo1"1t1ce lIbU'eC1b.Y'ejaoalatGr,r ciuat, nc:'naJ. setaDll O'f:lcluct; ejaoalatoJ:IT duct b:rN4 ad. Uned 1d.tbf'w c1r.Nl hoolC., O~25-l~o.a~ w1thlDos.q OOD'fOlute vu cler....; V&II de:t le8d sto spherical au! t1neq w.Iate pa t. gland;vas:'na] duct broad and lined 1d.th ab1t.1.D-l'keJll&teial; .-1nr1al~t of' ....tbeoa&1M! sp...toqst; spenl&theca been-1Ihapec1 aDd over2:lt a1ze of'Pl2"it01'm or P7Z'1f'om~;'sp...toqyst1o duct enten 8,P«N&tbeoal duct atjunction of pz'Ostat1c ADi 8lfIP"'1'ar,y duets at a1bwI1nam 1Il0UU8 glaM.

PJ.pn I·~ ta._ (Alder ud Bmeo*, 1866)•

.l. Ia~ fd the pm,ta1 ... aDd aD otr..t fd thec1rnl_*_.Be IAt..al "dn of the le.tt-halt 1"01J of radular tMth.

a • ..,.na.... - _1..." aDIl _oou cJad

eel - ej_CIII1atoz';r daath4 - h........uUc cluateel - niclacitpr - pzoarta". glmlao - teoptat - thecav - "I'M.

.. - ...... deteNl18

vd

; .I

\::S~bi"i

l.omm i

A

__ 0 ••__ ••• _- •• _ .._---_._-

O.50mm.--------_._--- . --------------S' -----

,I

Oc\;~B: Hawaii - Oahu (one specimen).

DEScmnoN: See F1gve 8.

Elttemal- Shape~ oval, d~resseci, 47 x ~j lI&lli;lerigid aDd t1neq grarm'a:r, ma:rgi.D tb:1D .w al1ghtqi.rregQlar; UIltle coloration dark orarJp-Nd1IO'ttJ.ed with 1rrepJJIr pa'tches of 1Ib1te; footDU'Z'OIf, J.1chto~; 'brancbiae :reather, 8,rachis ol'aDp aDd pinnae grq; 1'b1Dophores em.sll,oraDge with white tips.

Intemal- Badular 'teeth 42 x 58.0.58, 75-225pl long, allhamate, iml.. laterals hook-l1k:e, outez.st latwalselcmpte nth sl:1&bt 1rd1ca't1on of cnRer denticluaD! DO imler tlap, d-NJl1:, tooth~poa:l't1oned w1th1D each z.bibr :row; IIIiIJl amcoarseq lobalate g1IDl a't base o:r ej~to17'duct; OO'wm eztenIal orJ,f1oe 8hued bT ejaCll1ato17'duct, vaginal duct aDi cn:1duct; c:lft'&1 hook. :r.and larp, 250-3SOp lomg; pzoartate cJ.aDd e1mIgateaDd f1Mq lobaJate; vas'naJ. duct loo~ oomolute--le8ds cl1rectq to spherical spe1"JI&theca; IPC'JI&to-qR connected. b;y 8hort sp....toc:pt1o duct tojunct1cm of spemathecaJ., aJI.P'JllarT aDd pz'Osta't1cduct. at albm1n and. JIIlCOUS g].IId; ....toCV'stPJrl.t'01'lll aM 1e.8 than one-tb:1zd 81ze ot spenaa-theca.

FiflH! !-P1aDdor11 •••

.. I&teal Yiew.~ i,ii;.··pa1tal _. &Drl an o:r.t.et o~ theo1I'N1h....

~ 1&........ new fd the lett-h.lf' row o:r Ndular teeth.

• - apul1a.... _ alh ..hlat _oou gl&fti

eel - ej.aalator.r d1lethd. - h--.p1u.'od1t1c cluatocl - cn1dIlatPI' - proRate I1aDl.c- to~at - thecav - ng:tN

"fd - vu d.reNDS

. I

hd

pr

o.SOmm

. ----_. -----. -_._._--------A

O.50mm

B

(18:31) and subsequen1i descriptions all agree in certain diagnostic

generic characters and all of these cba1"acten aN exh:i,bitec1 tV thesingle spec1es ocaurring in Hawa:11an waters. 1\1e8e characters

iDc1ude: a~ with IIIIDOth, leath8Z7 tctuN; a mantle with

characte1"istic lumps or ridges; a "glandula et basta amator1a" 01"

litera:lq an &00.118017 gct.tal g1aDd aDd lateral st71et; am alack of buccal u ....tve and genital hooks~ The 't1Pe of branchial

sheath appearll to have DO d1agnostic generic 1IOrth as it 111 either

stellate (Bergh. 1890.),~ toothed (Kl.:1ot, 1903) or 6-lobed

OCCORRlDJCE: Hawaii - Oahu

a - apal1aa'" - a:u.d.a UIl _00118 gluxl.. - e3aCM1&torr aotbel- h. pb:Nd1Uo daot18 - latenl 8tTJ..tod - nidrlotpr - PNatate Slam.0 -apcw.toQ'lltat- ...~eoav - ftl"na

W. - "f'U de:teeu

..«!

. ,

iIII

.'

Vd

.... -.-,--.-"~

hd,.'!

r--·-"------·-·.·- --I

!.j•I

A

,---1)- IJ '--- ))....(:7

...---..--. -- .....--.-----.-.---- _. -_·_---1·I

IIi!,I

59

~. ~------"... ."~: ..."~ ...

r---·------·---'--..----.- ..-.- .I

"I

O.50mm

s'

'.

m. genus J0!"!!Jl!& was .atabiiaheci b7 Dergh (1016.) :in ~~ceto the earl:1e 1I01"k by' Hancock and JIDbleton (18,52). JOl"QDp' 18

gene1calq diat1lJp1ahed bT a characteristic lateral .t71et vi:t.h

1t. as80c1aW glud. !he lateral stylet in pt;sJ!ptM 1. __

1Ibatcl.,'ar bat 1s poe1t1one4 d1.tteren'tq. 1he ItmU8 J02L"lDJD&

i. alM charact.ns. by a 1'1neq v1lloua .sp:l.cu1&te UDtl. which

1. 11'1 COJItraat to the DDOth urrUe of jltupptv. Both ~OJ:Il1Ta

and Mt!motv po..... ,,"'ar e1IIpq haate teeth aD! lack buccalamatuzoe ed sctal hooks.

!be s-u i0l'!!" 1. ","H1'1ted in BaR11 by i.a. ......to... •.,pec1.. which hu bMD prn10ueq "ported in JDuoope •• far DOrth

as Deau.rk (Berch. 18800) aIIIl DOt N,POrtecl. hilt atat«l· to be

quNt1cmabq pre.ent in th. Pac1:t1o by Pftftt-Fol (19.54). 1h.

4e.ar1pt1ou ml tipN. of thi• .,.a.•• by Bergh and Pnvot-P'ol

0CIIIP&re ftZ7 tawrab:q with th. Bava11an epec1... One spec1men

ot is. jcoMptop.... also ooJJaatecl 11'1 tiwetok.

42

DBSCBIPnOl1': 8M PJ.son 10.

!!:t-~l_ SMp. onl, 001'1"f1IZ, ia-J.2 x 2-61a; 1UDtltt softw1'tb cl-.. oo,v1Dg of a:i.Irrit8 papi"il8 tzca =neb.PN3ect Ra1cht IlPl~, ursin thin u4el'Sbtll' 1rftp1.ar;~ oolorat1OD l1&htIN7 to drab 1d.th cIuk bzto1m to cluk &NT .-potsottan .tond.Di M pua1lel ron.; braDoh1ae b1;' aDdtrJ...p1..te, .s-U,SNT to ~, 1'OJ'II ba*""""keat.ftot1lN U01DId aDU,pap1l.].ate sheaths;. rh1Do-ph... t'u1toa, abort· with papillate tip, 7-9~ to broa 1_11••, papillate .eatha.

InterDal- Bacba1.r teeth (111 10 aDd U.~) 14-20 x20-24.0.20-24, S5-l65p loDa, all haate 1d.'tb,OIltu.st latenJ.a MHO... aDd _N e1Ampte thaD1m'1er 'tMth, DO dmt.:1olM, de-1"81lt tooth oetral.qpoll1ticmed 1d.tb1a rednlar 1'OW; lateral 8t7l8tPN8C't; .....~ or.lt.t.oe Mend 'by aheat.ht4 lateral atJ'let, e3acala1ioz7 claot ad .,.,.,dllat; duat ot latenl A7lAt OODIleotS 1d.th· a1bmd.neD:1. _oou .g].m; -3aaalatol7 dDat 14th lID hook. L -1... to loDe PZORate cJawJ 1Ih1ah part.1al.qeDftlapa apezw.theoa; tlPher10al ap__theoa nearqtv1_ d1aeter f4 8PG11&tocvwt with .,.,.,'er 8hape;.....toq8t 00III'leCt8 td.th spematheaal4uat atoawnn j1mcrt;1on of apal.].ary duct, plIORaUo actaDd duct ot lateal .tTl8t 111 albn1n aDd _ooug)em •

1:-----------·- --II

ad

\\ ,

\\, ,, \, \, ' --, ,,

.J,,,,,,,

'!'e

A

/-III

. ... l~_5\J

~- \---~

'-,. -., . .

,----.......Q.10mm

8

•

I,'/;"

1,

'"

, \

I'

111e 81ngle species of Dlpcodona in llawa11 possessea those

cbancteZ'a described or:lgi",117 bT Bergh Cl877b) as character1st1c

of the g8!me~ A m.m~ poamalate mantle, bu.coal azmature, s1mpq

hamate teeth am UDa1"IIed peDis. HDst f4 Bergh'a characters are

also 8Xidbited b7 other genera, ho......., ad it ... that the most

d1apstic character ot the geD8 18 the rod,.11ke buccal a1"II&tuN

1d.th traunrse gzowth-r1Dga. .As po1ntec1 out b7 JlaZ'CI1S (1955),

when a spec:1ea which otherwise appears to be a d1scod.or1cl except

1D pos...s1Dg &1'1 amed peDis, it is advisable to tolJ.ow Bergh's

(1892&) later aM. ..."ed det1nt.t1on of the senua m.codo;1& 111which the pd. i. "near17 alwqs· 11DU'II8Cl. .As 1D other dorid

SeDer., the presence or absence ot pDital amatuJ.oe C-cept Pl'Obabq

ill l}.ttJdp!1.) is ofte "fU"1ab1e .-mg apec1ea am even aJIODS cW'-tertmt wed :lDI1T.l.ciIlals o:t the 8aD.l8 apeo1es.

1he s-u apooc1or1. is ooaopol1tan 111 wuaer vates, althoughthe ajorJ:qr of the est1ll&tecl 40 species in thi. pJiera are :trom

theIDlio-West-P.c1t1c.

occtJB.llBOOE: Han11 - Oahu, ltauai (fairq 00"· '11).

DECRIP!ION: See Figare 11.

E!tternsJ.- ~~cml1~ d~__D 5-3S x 2-2011I;JIIJ1't1e ric1d &Dl _lIhaot cr.mtJar w1th m1mltepaP'" •• upon 1Ih1ch an c:J.uteecl pe" ..cal..,~. th1n ard 1z'Ngu1.ar; untJ.e often_.a_ utI-' 1. d111tazMd; MIIUe co1orat1OD .-ttledda'k &NT ap1ut & l1aht r;ny ba.~, oft6nwith black patch.. ~~ lateraUn bNDcbi..tn- or qudzo1..p1Jmate~ 4-6, d1uIIq 1D ooJ.on.tioD,with pap1" a-te brrucb1&l 8b._th; ~.tu1foa, 15-20 ctuJr;r, 'M11••, 1d.tb 01'_ oo1fnedtube-like apice. aDI1 pap1lla-te~nl sheath;utd..lI1Ai. o~ uatle IDr1 1'oot ONIIII. with clakspeck",.••

a - .,....:1'.a'" - all"lld'D ad~ sJ.m

eel - e3aaalatol7' cbloteel - n:ld1lGtPI' - Pl"08tate I1azJd..0 - .-_toClptlit - .....theoa

T - 'ftI5DA'fd - vaa deteNI'UI

-----,I

a

---------------A

'o.025mm i

B

I~---

L 30 ' 0_.2_5 mmC

"5'\'(/1

Two species c4 dor1c1s hem Bnai1 exhibit the characters of

the pmuJ Ccfta:h""lori. described b3' Bergh (l889&). '!he moat

cbv1cu c==on ~a..~ere o~ these epec1M~ to M the aimlteq

dent1calate haate teeth am the puatulate mantle. j],t1MnIgh nat

in the genm.c description, ~cca1 anaature are described and t1gurecl

by Bergh (1889a) as .wmgate rods with l'1Dg-I1ke transvene growths

tor the tJpe specie., ~r1&_We. anq ODe or theHawa11aD species doe. DOt have amature in the ejacalatorr duet,

1ih.... the other species a:b1b1t. amatU1"e in both the vag:t.na1

duet as well as 'the ejaoalator,r duet. 0w1Dg to 'the Ta1"1ab1l:l't7

of genital alW.t1II'e w1tb11l aD! betwen apec1.. or IUUJI' dor1d,~:

••era, it is a..... that this Cbazoacter is DOt or d1aIDDst1c1IIportmce 11'1~ Bdd:b1tecl b.r both Ha_11 an speci.. c4

~ is an~ vestibular gland, 1ib1ch is DOt d..cribed

by Bergh (1889a) for the pIlUs. It is assaaed that thi. al80 is,

of DO diagnostic 1IIportance gener1call1' since Marcus (1955)

desoribes and figures a simler structure for a species of m,.sodol1f.

Ca1'miDodor1. ~sa (qu, l864)

OCctJ:R:R.ENCE: Hawa:11 - Oahu (5 specimens).

DBaCRIPnON: See Figure 12.

iI:ternal- Shape oval, CODVti, 26-35.x 24-23ia; iiimtle rigidand cr.lIP, covered lat8Z'al:q with J1D8NU -Upustule.; mantle mottled nth J.1sht bro1m aD!splashed. v1th dark bro1m; mecl1al- NC1-bzoom baml;bramch1ae large and tr1p1Jmate~ ?Sl coloNel CNAIIwith bJIom fleet.; hNDch1al .eath with scal1Dpsdedge; rb:hYlphorea large and ~ol1ate, brom,retractUe into .calloped sh_th••

Internal- Iad.ular teeth 28 x 59.0•.59, l2O-225JJ., all haate,11m.. aDd. outer laterala DOJl-e1_t1calate, ctmtraJ.:qpositioned cLwrhwnt, w1th JI1Jmte outer deDt1cl..;blocal amat1lN '3-~ loDg 1'Od8 with tnnsnrser1Dg-l1ke powt1uI; bl1rJd Bed ft8t1ba1aZ' s1aDl of8.,....]. 00Jm)1u'tiorla ha~ 110 aaat1U"e or 8'tr7leempties 1Dto CP"WKm pnita1 ft.t1ba1e a10Dg withejaoula1'.ol7 ad ng1D11] ducta; DO cbftl books orvq:lnal amattaN; '9U deteND8 YfII!7 1Imc ADl t1ghtqCOllVOlute; plORate glucl .-oth am elcmgate;vag5nal duct J..d8~ into apematheoa;......tbeoal duct Dearq __ lagth as ftI1nalduct-- leads to ~OD~ sp.....tocvwtic, pr0-static aDd. 8IIIp1IJ:l..air ducta at .1bnw1D and _oouag1aDd; ~ON"""toc,yst about 1/2 81u otspherJ.ca1 .spematoCJBt.

• -...,..na.... - a1"'. ad. _ ... glaDd.. - .~.GII1atoZ7 actad - 8Y1GetPI' - paNW p.m.0 - .,....toG1R8t- ....tJl..... _ .,..:tv .

... - 'YU det....'fl. - Te8t1balazo IlaDi

v

I-I

____vd i!

"

Ii

----_._/..pr I

II

I!.

~lL---8d

oct

,! .

1--"II

iII

.... --'--'1

I

III

!

A .

O.05mm

B"

---_.-j,iI

II

"

O.50mm,:_--,-----------,-t- '-",

,!

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0CCtJBBEHCE: Hawali - ()abu (l,spec:1men).

DISCIIPTIOIl: See F1gve 1:3.

~ Shape oblong-oval, ~_; mantle rigid nth laztgeaDd·~ put1I1es, untle~ thin ands1:1shtq 111"'h\!a tecl;lIU1tle coloNt'1cm t..- withret1aalat1oDs' ml 8PJ,aah.. ~ bIocnm; branoh1&etr1p1zm&te ad ruth.,., S, ch1a1o" 1'aD 001ozo;1"b1r1ophoNS stout, pCunclGS eqaal 111 lqtb to] ae1lar portion, :tan oolon:t1on, with tubalarrh1nophoral sheath••

Internal.- Badular teeth 40 x 120.0.120, 1:35-28Sp loDg, .uhaate, 1mlelWtst latenls .-oth-nooe.d:nlatenla within l'GIr 1d.th outer denticlu, d"'nanttooth C8I1'f2al:q positioned witb1D l'OIIi lateralj&1l8 with 1"Od-11ke blcoal amatue, 30-36Jr. loDg,haT.lng tran...... pcnrth r1Dg.; 'bl.:1Dd-eDd.edvest1balar gland ~ sevwal o 00Irf0lut1au h&'ringno amatuM or 1It71. -.pU•• iJlto oa II. ceitalvest1bale al.cmI1d.th ejaaalator,y am ",,!'AIduats; c:lrl'a1 hook., 7S-1'~ loDe; vq:tnal ....ture,6O-:300Jl 1cmg,~ 'YU'iable in shape and lI1UIberor teJd.Dal hoak.; ft. det.... .,.,. lAme &D1t.1.ghtq oomoJ.1lte; prostate gl.m PJrltON am1'1neq lobalatel .....,n.l duct 1...~ to°spezw.theoa; .,...athecal duct .,.,. 1Ib.ort. leld.to junction at .'......" am moou g1aixl1d.thspematoqstic, P1'OstaUo aD1~ duct.;aptmI&theoa ovate ad :appl'G'dmt-q 2X 81.. 01'PJT',lt01"ll spematoqllt.

1P11!E'11·

A. r..~~~~:-::'~ or theobnl h... alii the ',,,., Ull&t1I.l"e~

II. a.IBM of the bIlooal·uilatue.~ IatGal ft. ~ the J..tt-halt l'01r or NbJ·r teeth•

• _ ..,1J•.... _ -1..." alii _oou sl.m

• - ej-.J '-IIT dactod - OY1daotPI' - pNna'te cJ.and.0- ..-toqst8t-~eoa

'Y - ftCIM-- .........VI - "N8t1'bU.ar I1aDl

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120

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Acool'diDg to Be~ (l877b) the diagnostic characters of

the genus Jrippa are a f'ai1"~ soft bocV with 00DI,P01UX1 tubercles

(aall tentacles) and • 'M-oad foot 'iib1ch 1s unit&d anter101''q to

the head. A. subsequen't redet1n1Uon of gcleric chaZ'acters by

BI.1c't (1902) includes: no lab18l 8l'!Mture, Ndula CO!IpOsed of'

~om mwp'q haate 'teeth ~ a D1DIbeZ' or g1Jmc1u1.ar bocl1es a1'OU1'Jd

the poater1or extr.1V of the ba.coal apparatus. ~ 'two of the

BanB all species, .L. 0leoe aDd II. Lcb1w:t;a. atdbit OOJUJPiauouaq

lobulate aa1:1ftZ7 g1aD:1_ on the bIlccal .us which ccmce1vabq are

ed-"ar to the -g)_Diular bodies- described by BI:1.ot. 1IoInmIr,

as Bl10t (1903) points Ol1t, the absence or the lobulate sali'9V1'

gland_ ehoald mt erm].e a species 1Ibich uhibita other characters

of the germs.

111e three *P8C1ea ot .Mppa in Hawaii show the wide variation

of JIIaJltle ep1demaJ. texture 1d11ch OCCU1"ll in this group. Although

all three species are "·l1ar:Q' oval-e1oDgate in shape, onq .L.

sc;abz1gcg1a am .t&. 0Il!OU OOD8Pi~utdbi't the cam;pourdtubercles described 'b7 Bergh (l871b) as diagnostio o:t the genu•

.L. echb&ta is """41" to the other two species in all othercharacters, bIlt the mantle is covwecl by minute:q spiealate·papillae

instead of~ tuberculae. At the present 't1Jae it .... better

to include this species in the genus k1ppt. nth.. than to include

it on the basis of' an external character 1dr1ch is oft4m VU'1able in

dorid gen......

.53

DlSCRIP!1OIl: See IP1gare 14•.

ElcternaJ.- Shape oval.-elonga'to,.. C;=m:ct, 28 :: Jk_; =ntlefUrJ:' rigid and~:Qr !'e'ticmls.te !Md1&lridge nth latezoalq branch1Dg 1"1dgea enclosingrather'dMp pita; ..DUe coloration gNT-'b!o1m;branch1ae ~rmate, '6, bNwD, branchial sheathCNnUlate; rh!Dophoru nd-l1ke aD! -loneate,'b!oown, rh1nophoral sheaths cNm1lateel·bI' papUlae.

Int.emal- Radu1 a r teeth 20 x ,s.O.38 (111 28Ia apec1aeD),6o-23Op.long, all hamate, dmdnant toothc~positioned in z.a.ctul.r row, 1m1ermst lateala withf'airq bro8d bas_ and reca.rred cups, outezmstlaterala elongate md graem.'17 hooked aJ.onc ct1relength; compcmmta ~ genital _u not discemible.

D... ~·.Jr1.. If". (hl••rt. 1859)

I.. Iat.N1 ft_ of 1;he left-haJ1' NW of r.mt1.r tMth.

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~ Shape oval-alongate, iiGliiV.ma't eGJ1'i'iiXt 4-24 x 3-8Di;mantle :fairq rigid and =cabrou 1d.th m1m1tG paG'tulasand spinea; maIitle coloration Cl"MII-1Ib1te often withbrown spots; foot white and. Mlab1ate; bl'ancb1aetr1p11mate, 5, clusq1manch1al sheath COD1calwithstellate edge; 1"h1nophores stoutq pedunculate and_c:oc.-~t:, =:!;:;.

Internal- Radul.,. teeth 14-15 x 21-24.0.21-24 (3 specimens),45-21OJlloDg, all haute, 1Jm-.st lateralreCU'Ved a~ IlUcceed:1r2g teeth, ctoprlnant toothcentnJ.q positioned within radul.,.~ withtooth 1ength decreas:bJg to either aide; ejaoalatoryand vaginal ducta share comon oriftc.; :t1neqloba1ate gland eno1z'clu base ~ ejac:a.lato17 duct;no c1rral hooks; Pl'Ostate gland loo..qtlm'ft)lute;sem1sv1al arnna-ent of' epematheca UId speaato-07IIt: vaginal duct short aDd stN1cht.--1eadsd1reot:b' to .....theca, sp....th.oal cIuct short-leads to sbort ape1'!I&toC7llt1o dua1i ADd loo..qOOJm)lute utel"1ne duct; uter1ne duct enters a1bwa1nand _COUll g1&nrJ at junction of' Pl'Ostat.io ard~ ducts; spe1'III&too;rat P.Jl'1f'cma &&~ about+size o~ spherical 8,P81'm&theca.

F11!D~

!r1ppt _~Mta (P..., 1860)

A. IAteal "f'i. ~ the g-.tal _a.& lAteral "f'i. or the 1ett-ha1:t 1"OW ~ radul.r tMth•

• _ ....1J •

.... - .1hmta aDd _cou g1eDdeel - ejaaalat..., _atbel - h-..phred1t.1o cluoted - cm.dIlotpr - pamte glaMae- ......,..R - apematheoa

T - ....".'9d. - ft8 clet....

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OCC~: Hawaii - Oahu, iAuai (5 specimens).

DESCRIPTION: See Figure 16.

BIttemal- Shape oval-elcmp:te, slightJ.y convex, 10-20 :x:6-lODm; mantle f'airq rigid am BPOnge-l:1ke 1nappearance, 1rrep1.arq ret10ulated with I"idgesard Jdts, tubercle. at ridge junctions part1=-lar~ alOng the lI1d-l:1ne 1IPOll 1Jh1ch are pel1white pustulos; mantle coloration gN7 or 011ve-green with da1'k pipentation in pit.; 1'oot whiteor creal bNncb1ae a:.1JIFq pilmate, 5-6, duk gN7in coloration, branchial sheath .-ooth -am caVWDOU;l'h1nophOres rod-l1ke and elongate, 18-20 gN7 orolive-green ~llae, rhinophoral sheath pustulated.

Interna1- B8dular teeth 17 x 18.0.lB, S3-J,2Sp. loDg, allhaate, outemost lateral. _re e10Dgate aD! notf'1arlged &8 imler latera1a, etamnant tooth central.qpollit1oned am ..".at teeth lateralq positionedw:1:tb1n ..~ 1Whllar 1"OW; ejacmlator,r duct ardvag:1nal duet with • ...-rate eatternal OZ"1t1ces; DOc1rral hook.; prostate s1aDllcmc am bzoad;

-aezo1al~t f4 ~theca am spematocyat;spemathecal duct abort; both spematheoa ar¥lapematoo;yst spherical and ","11arq ~secl; om-m:Jtmet1on- of uterine, P1'08tatic aDd ampull.arrducts at the a11mm1n and ..cous glanr:l.

lJ.n!! li-~ .oajr1 , (P , 1860)

J.. IA~.ot the ta1 ~ .L Iateal u_ .f the le:tt-halt 1'VIr of Nd1l1ar teeth.

a - apallaa'" _al....n ad ..on. g1aaleel - ~.:l.atoJ7-etbd. - h. sphrocl1tie aotocl - O'Y1d1IotPI' - pzeatate C] e•• e - .,...teCQ'lltlit .. .,..theoa

T -Tq:Ina'9d - Tall 4et....

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Genus Balgsda, Bergh, 1880

ille genus He'S'!!Ll as origjnall" described by Bergh (18800)

has cont1"oversial aD! 80DlRhat atrinpnt oBaracters. .Aa & result,

Eliot (1903) erected a n.... germs, Sclerodori.. ..,nl;r on the basis·

of d1t'i'ereces f4ep1dezul~; he later stated (1Liot, 1908)

that Ra1.gFda 'WOUld be tho a== as Sclercdor:1; it it wu redet1ned

in having a bud, rough texture &8 wll as a IIIDOth, f1m texture.

On the basis f4 the S'tl¥V of tbJtee Ha-11 an species of dorids

it appears that a red~tion of the geml8 as 8\1IPsted by

ELiot (1908) would be in ol"der, &8 wU as a broader de.t1Dition-'-7

of other characters. Although L. nbra ahib1ts tightq senated

outer laterals which ..... stated to be diagnostic by Bergh (18800),

the other two species do 1'JOt show this tcJdenCV'. '!he Hawaiian

apec:llllen of .. mbra al80sddb1ts hooks in the ejaaulator;r duct,

a character which 1f&II not tOlmd by either El1.ot (190') or Bergh

(1905) for the species; in fact, El10t (1908) states that the

lack of hooks in the gem.ta11a is characteristio of the genus as

well. .l character 110t suggested by previous authors &8 a g&e1o

character b1t sh.azoed by all three Hawa1.1an ~ec1es of JalBerd& is

the preeenceot .. lobulate glaDd at the base of' the vag1na ' duet.

:Further &QP!'Jnation of other species is necesar.r, however, to\

d.etendDe the diagnostic lIOrth of this character of' the genital

1II&8s.

Xt 18 sugested, thent'ON, that· a redetiJd.tion of the genus

Ie]!""k Bergh IIhould include those characters ~ the genus

60

wuld theeb7 be a tim or f'leaby' bod1' 1Ihich mq be smDOth or

rough, a mantle with ridges, reticulations or depressions or a

oombina't.1on of aU three with 1rregular1ties am tubercles; 8IIIOOth

or den't.1culate hamate radu'8r teeth; &lid a lack of ~ccaJ. plates

or armature. :rt is COBIIOn tor species of HeJ f!!IO!!e to appear

quite sponge-l:1ke in app8&i'ance which has been suggested (Kl1ot,

1903) as a po88ible example of 1I1JII1C1'7.

61'

OCCUBIiIICB: Hawaii - Oahu (3 specimons)

DIBCRIPTION: See Figure 17.

Bltter!"..sl- ShApe oval, 16-88 x 8-40Js; -.DUe bz'oad, 8IIDOthand rigid, margin ibinand~; uatle c0lora-tion oftIIP to oranp-l"8C1 w:1th 1ftoeplar 1Ib1teand/or I!IItftI1l1OD spots, N1secl·%'et.ieala't1cm8 givespcm8e-J.1ke appearance; area between Mt1aulat1onst:l.heq tjpeolWid with black pipentaticm; one tosneral ni.ecl ADd-me aNU on ..tlej .toot andvel'l'IiNl side of aatl. smoth 'II1th orange-mcolont1on; bnDch1ae tr1p11mate" 5-8, ONDge-red

A to oZ'aDP-bNNn, p1Jmae ecIpcl w1:th 1Ih1te or cream;bnDc:h1al sheath with jagged eclp; Z'b1Dophoreafullif'om,o~ too~ 1I1th 1dl1te tocn&IIl tips; 1"b1Dopho1"8J. sh_th with jagged edge.

Intenal- B*'D'er teeth, '''':z: 53.0.5' (:30- 1Dd1ndual),75-22Sp. lADg, &11 heJlate, owtemoet 3-4 lateralsleaa __te than PNCeed:!Dg teeth 1n 1'011;doJdnent tooth· .t;*ltra].q poa:1t1onecl 1d.th1D each!'OW; OCWW'n e:a:tuaal ozU1oe 8haftcl 'tv'.ejacalato17and vq:lne] duet.; ejacalatol7' cl11~ ~j cd.z'nlhooks" l5-6Op 1mag; '9'U cle.t.-a, abet ... looseqCOD'V01.ute; pzoatate gland t:l.heq lobalate; coaN~lobaIate sJUMI at base of 'ftg:In.1 duct; ng:fna]du.ct t1gh1:.q' OOJJ9Olute and lAma eter. spher1celSPe1'Dl&theca; spemathecal duct UD1ted 1d.th uterinedu.ct at short spematoc;yat1c duct; .....toQ7lJt1cduct opens 1nto ovate SP8l'll&toa;y8t 1Ih:1oh 18 n.-r~as large aa SP81'Dl&theca; 'VerT short uterin. ciUctenten junction with~ and P1"Ostatic duetsat albumin and. mucous glar.d.

lJ.cF! n·It'·" n1n BRab. 1905~ IAteNJ. n._~. tal _. ad aD ott• .t fd the

e11"Nl hoob~B. . IAteal '1'1_ of the lett-halt ZOW' fd n41Ilar teeth•

• _ • I " •.... - elhwtn .. _aou glud

eel - ej..-lat.or,r.CItbd - h~t1c duGtod - cm.daat.PI' - pzodate c'am80 - teqRat - theoa

'Y - 'ftI'."fd - .,... cleteNJUJ

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