THE VELIGER

© CMS, Inc., 1994

The
Veliger
37(2):141-154
(April
1,
1994)

Descriptions of Four New Eulimid Gastropods

Parasitic on Irregular Sea Urchins

by

ANDERS WAREN

Swedish Museum of Natural History, Box 50007, S- 10405 Stockholm, Sweden

DANIEL RAY NORRIS

Marine Laboratory, University of Guam, Mangilao, Guam 96923, USA

AND

JOSE TEMPLADO

Museo Nacional de Ciencias Naturales J. Guiterrez Abscal 2. E-28006, Madrid, Spain

Abstract.HypermastusmareticolaisdescribedfromGuam,H.orstomifromNewCaledonia,both
parasitizingthehearturchinMaretiaplanulata.HypermastusobliquistomumWaren,1991isrecorded
fromthesanddollarLaganumdepressumfromNewCaledonia(hostspeciesnotrecordedpreviously).
EulimaencopicolaisdescribedfromtheGalapagosIslands,parasitizingthesanddollarEncopemicropora
galapagensis.BalcisclypeastericolaHabe,1976,parasiticontheclypeasteroidseaurchinClypeaster
japonicus in Japan, is made the type species of a new genus, Clypeastericola and a new species C
natalensislivesonClypeastereurychoreus,fromsoutheasternAfrica.Twofurtherrecordsofspeciesof
ClypeastericolaarerecordedfromClypeasteraustralasiae(Gray),fromNewSouthWales,andfrom
LaganumdepressumfromNewCaledonia,butthespeciesareleftundescribed.

INTRODUCTION
Eulimidgastropodsarenotfrequentlycollectedontheir
hostsunlesstheechinodermsarespecificallysearchedand
handled with care; most eulimids have the capacity to
voluntarily leave (and many do it regularly) the echino-
derms on which they are feeding. Therefore, the most
profitablewaytofindeulimidsisbyintertidalcollecting,
where one can see them directly when picking up the
echinoderms, and they do not have the time to pull out
theirproboscisanddropoffthehost.
Occasionally,eulimidsarefoundasasideresultofother
investigationswhenhostsarecollected,especiallyeulimids
permanentlyattachedtotheirhosts.Thispaperisbased
onsuchmaterialsenttotheseniorauthorforidentification.
The material presented here is not extensive, but it im-
provesourknowledgeofthegroups,especiallythehost-
parasiterelations.Herewedescribeonenewgenusand
fournewspecieswhichparasitize"irregular"seaurchins.
Thereareveryfewrecordsandspecimensofsucheulimids
(Waren&Crossland,1991).
Acomplicationofeulimidnomenclaturewhichconcerns
thespeciesparasiticonirregularseaurchinswasthein-
troductionofanichnogenericname,Heckerina(Alekseev
& Endelman, 1989). This was created for some holes,
diameter1.0-2.3mm,intestsofCretaceousspecimensof
Galentesorbiculatus(d'Orbigny,1854)(Irregularia,Gal-
eritidae).Theywereassumedtohavebeencausedbyeu-
limid
gastropods.
MATERIALS and METHODS
Thematerialusedforthispaperisenumeratedundereach
species.Itisnowdeposited,usuallytogetherwithvoucher
specimensofthehosts,inmuseumslistedunder"Material
examined."Thenumberofavailablespecimenswastoo
smalltoallowmuchmorethanadescriptionoftheshell,
except of Hypermastus mareticola Waren & Norris, sp.
Page 142
nov.,ofwhichafewspecimenswereseriallysectionedfor
anatomicalstudy.(Thoseresultswillbereportedlater.)
Authornamesofthenewspeciesaretobequotedasin
theheadingsforeachspecies.
SYSTEMATICS
Family EULIMIDAE Philippi, 1852
HypermastusPilsbry,1899
HypermaslusPilsbry,1899:258.Typespecies:H.coxiPils-
bry,1899,byoriginaldesignation,Australia,NewSouth
Wales,PortStephens.Hostnotknown.
Remarks:Waren&Crossland(1991)reviewedthespecies
assumedorknowntobelongtoHypermastus.Elevenspe-
cies are known to live on irregular sea urchins in tropical
regions.Habe(1992)hassincedescribedonespeciesfrom
anechinothuridseaurchinfromJapan,Hypermastusarae-
somaeHabe,1992,butthatspeciesdiffersfromtheothers
by having a light yellow shell, and since very little else is
knownaboutit,weconsiderthesystematicpositionpro-
visional.
The two new species described below are also ques-
tionablemembersofHypermastus,butpresentlythereis
nogenusavailablewheretheyfitbetter.
Hypermastus mareticola Waren & Norris, sp. nov.
(Figures1-3)
Type material: Holotype and four dry and six alcohol-
preserved paratypes in Division of Mollusks, U.S. Na-
tional Museum of Natural History, Washington D.C.,
registernumbers860367,860369,and860368;sixpara-
types in Swedish Museum of Natural History, register
numbers 4531.
Type locality: Guam, Apra Harbor, about 10 m depth,
parasiticonMaretiaplanulata(Lamarck,1816)(Spatan-
gidae,Spatangoida).(VoucherspecimensUSNME43011).
Material examined: One snail, attached on host (16 mm
diameter), Apra Harbor, 10 m, 16 August 1992. Fifteen
additionalhostsonwhichsnailshadbeenobservedinthe
field but had fallen off during collection or preservation
werealsostudied,andthedistributionofthesnailsonthe
hostisgiveninFigure3.Atotalof10snailswereobtained
from these. In all, 21 snails were collected from hosts
during June, July, and August, 1992, all at the type lo-
cality.
Etymology:Namedafterthegenericnameofitshost.
Description:(Basedonsupposedfemales.)Shell(Figure
lA-E)tall,slender,greyish-transparent,almostperfectly
smooth. The larval shell (Figure IF) is colorless, cylin-
drical,mucronate,obliquelyinserted,withslightlymore
than1.5whorlsandaheightof300-320jum.Theholotype
has 7.0 distinctly convex teleoconch whorls, with a very
shallowandinconspicuoussuture.Therearesevenrather
The Veliger, Vol. 37, No. 2
evenlydistributedincrementalscarswithintervalsof0.7-
1.3whorls.Thescarsareverythinandnotconspicuously
deepened, except close to the apical suture. There is no
sculptureexceptsomeincrementalunevennessintheshell
whichhardlydeservestheterm"incrementallines."The
subsuturalzoneisconspicuous,anditsheightcorresponds
to about 25% of the height of the whorl. The aperture is
pear-shaped,rathertall,withadistinctparietalcallus,but
nodistinctanglebetweentheparietalwallandcolumella.
The outer lip is very slightly prosocline, retracted at the
sutureandmostprotrudingjustapicallytothemid-point.
Male.(FigureIC)about60%oftheheightofthefemale
andslightlymoreslender.
Dimensions.EighteenmaturespecimenscollectedJune-
August 1992 had the following shell heights: 5.20, 5.00,
4.72, 4.64, 4.40, 4.24, 4.04, 3.96, 3.76, 3.72, 3.24, 3.16,
3.00, 2.92, 2.80, 2.56, 2.52, 2.52 mm. Four immature
specimens(recognizedbytheangularperipheryofthebody
whorl. Figure ID) were 4.60, 2.56, 1.88, 1.88 mm. The
heightoftheholotypeis4.40mm,itsbreadthis2.52mm.
Soft parts. The eyes are large, 80 ^m in diameter, and
situated 200 ^m apart, basally at the outside of the ten-
tacles.Thetentaclesaretapering,slightlylongerthanthe
distancebetweentheeyes.Theproboscisisfullyretractile.
The foot is larger than the opercular lobes and has a
distinctpropodium.Inpreservedspecimens,itisdisplaced
toward the right side. The operculum fills the aperture
and is very thin and colorless with no reinforcing ribs or
specialmuscularattachments.Fivespecimensofthefol-
lowingsizesweredecalcifiedforserialsectioning:5.2mm,
5.0mm,4.6mm(supposedfemalesandimmaturefemale),
3.2, and 3.0 mm (supposed males). The two large speci-
mensandtheoneofintermediatesizeallhadoviductsbut
lacked a penis. The two small specimens assumed to be
males had a penis. The sex of these specimens was con-
firmedatserialsectioningandexaminationofthegonad.
Remarks:Hypermastusmareticolawasfoundonitshost
inconnectionwithecologicworkonMaretiaplanulataand
Metalia dicrana H. L. Clark, 1917 (Brissidae) in Guam.
Althoughthesetwohearturchinsoccurmicrosympatri-
cally, the eulimid was found only on the former species.
Somedetailsoftheecologyofthehostweredescribedby
Norris(1992).
Onesnailwasstillattachedonitshost(16mmdiameter)
bypinchingtwopedicellariaebetweenitsoperculumand
peristome.Itwaspositioneddorsallyovertherightpos-
teriorambulacrum,about5mmfromthegonoporesand
betweenthetwodoublerowsoftube-feet.Asmallarea,
about0.8mmdiameter,wasslightlydiscolored,darker
brown,withaholefromtheproboscisinthecenter.Inside
thehost,therewasacorrespondingdiscolored,slightly
swollenarea,butnoindicationthattheproboscishadgone
farther, as to the gonads. There was no proboscis of the
parasiteprotrudingfromthehole,andnotornproboscis
could be seen on the snail, which indicates that it is fully
retractile.
A. Waren et al., 1994 Page 143

Explanation of Figure lA-F

FigurelA-F.HypermastusmareticolaWaren&Norris,sp.nov.,syntypes.A-B.4.6mm(female?).C.2.8mm
(male?).D.1.9mm(youngfemale?).E.Incrementalscar,length380tim.F.Larvalshell,height310/itm.
Page 144 The Veliger, Vol. 37, No. 2

gg^gg-ii ..---■--■----.■:
-VvCSi
•^^s

^,j^

"•i#?^v-
=■>
-lO^-'
: ^^^ MS-
■-

^'fci

Explanation of Figure 2A-D

Figure2A-D.Maretiaplanulata,withdamagesoftestfrombeingparasitizedbyHypermastusmareticolaWaren
&Norris,sp.nov.(markedbyarrows).A.Diameteroftest11.6mmdiameter(excl.spines).B.17.4mmdiameter.
C.LargeholeonspecimenB,diameter0.5mm.D.Smallscararoundaholeofatubefoot.Scalebar0.25mm.

Aroughestimateisthat10-25percentofthehostspec-
imens were infested; and 20 percent of the parasitized
specimensofMaretiaplanulatahadmorethanonesnail,
with a maximum of four snails on the single host.
The 15 host specimens represented in Figure 3 had a
totalof25holesfromproboscides.Twooftheholesseemed
tobehealing.Tenspecimensoftheparasiteweresaved,
whichmeanseitherthat15specimenswerelost(whichis
possible),orthatthesnailfrequentlychangeshostorpo-
sitiononthehost,andleavesaholeonthehost.Wecannot
saywhichofthesepossibilitiesbestexplainsthediscrep-
ancy, perhaps it is a combination. All snails had been
attachedaborallyonthetest,andtheirdistributionisplot-
led in Figure 3. This figure indicates that they prefer the
A. Waren et al., 1994
Explanation of Figure 3
Figure3.DistributionofholesandscarsmadebyHypermastus
mareticolaMaren&Norris,sp.nov.on15specimensofMaretia
planulata,onwhichtheeulimidwasfound.Thesizeofthehosts
(10-18mmtestdiameter)hasbeenstandardizedtomakecom-
parisonspossible.Largecirclesindicatelargeholes(Figure2C).
leewardside,perhapstoavoidgettingscrapedoffwhen
theseaurchinismovingforwardinthesediment.
Thedistinctionbetweenmalesandfemalesfromshell
charactersistentative;itisraretoobtaineulimidsinquan-
titieslargeenoughtoallowimpeccablestatistics,especially
sincetheprocedureisdestructive.Nevertheless,wefindit
worthdrawingattentiontothisassumedsexualdimor-
phism.
Hypermastusmareticolaisnotaverycharacteristicspe-
cies,butthecylindrical,mucronatelarvalshelldrastically
reduces the number of genera from which to choose. It
bearssomeresemblancetoBalcisechinocardiaphilaHabe,
1976(Figure5A-B),aspeciesparasiticonthehearturchin
Echinocardiumcordatum(Pennant)inJapan(notonSpa-
tanguspurpureaasquotedinWaren&Crossland,1991).
That species is, however, considerably larger, about 13
mm, and has a more ovate aperture. No soft parts have
beendescribedfromBalcisechinocardiaphila,whichmakes
comparison difficult. It is likely that they will end up in
thesame(new)genusinthefuture.
HypermastusorstomiWaren,sp.nov.
(Figures 4A-E)
Type material: Holotype and two paratypes in Labora-
toiredeBiologicdesInvertebresMarinsetdeMalacologie,
MuseumNationald'HistoireNaturelle,Paris.
Type locality: New Caledonia, Lagon de Noumea, Gran-
de Rade, 22°15.26'S, 166°23.98'E, 11-12 m depth, sand
with ooze on surface, 14 April 1993, three specimens at-
tacheddorsallyonMaretiaplanulata(P.Bouchet,Labora-
Page 145
toiredeBiologicdesInvertebresMarinsetdeMalacologie,
MuseumNationald'HistoireNaturelle,Paris).
Materialexamined:Onlyknownfromthetypematerial.
Etymology: Named after L'Institut Fran^ais de Re-
chercheScientifiquepourleDeveloppementenCooper-
ation, formerly Office de la Recherche Scientifique et
Technique Outre-Mer (ORSTOM), whose personnel in
Noumea has sent the senior author numerous new and
interestingeulimidswithhostinformation.
Description: (Sex not known.) The shell (Figure 4A-C)
isshort,stout,conicallyovate,colorless,transparent,with
mucronateapexandlargeaperture.Protoconch1(Figure
4B)iscolorless,andhas1.7perfectlysmoothwhorlswith
veryindistinctsuture.Itsheightisabout250^mbutvaries
according to how much is concealed by the subsequent
whorl. There is no protoconch 2. The holotype has 8.0
perfectly smooth, very flat whorls with a few indistinct
incrementallines,mainlyrestrictedtothesubsuturalzone,
and two to four slightly stronger incremental scars. The
sutureisbarelyvisible,thefalsesuturemoredistinct,and
thesubsuturalzoneoccupiesone-fifthtoone-sixthofthe
heightofthewhorls.Theperistomeisratheroblique,pear-
shaped,slender,andsomewhatexpandedinitslowerpart.
Seeninprofile,theouterlipisprosocline,retractedatthe
sutureandmostprotrudingjustapicallytoitsmid-point.
Dimensions.Heightofholotype5.89mm.
Remarks: Since only three specimens were available, it
wasnotpossibletoexaminethesoftparts(thisrequires
that the shell is dissolved). The few cases available show
thatH.orstomicausesunusuallylittledisturbanceofthe
test of the host. A few spines had fallen off around the
pointofattachmentapicallywithinoneofthepetals,and
theareaaroundwasslightlydarkerthanitssurroundings.
Hypermastusorstomiisbroaderandshorterthanmost
eulimids,especiallythoseassignedtoHypermastus.Hy-
permastuscoxihassimilarproportionsbetweenbreadth
andheight,butismorecylindricalwithmuchmorerapidly
expandingapicalteleoconchwhorls(seeWaren&Cross-
land, 1991, fig. 4A-F). Species of Turvena (see Waren &
Crossland, 1991, figs. 9H-I, lOA-B) have a similar shape
oftheshell,butanapicallymoreconstrictedapertureand
acolorpatternontheshell.
HypermastusobliquistomumWaren,1991
(Figure 6A-F)
Hypermastus
obliquistomum Waren,
in
Waren
&
Crossland,
1991:100,figs.7E,7F,12H.
Type material: Holotype, Australian Museum, Sydney,
C.160815.
Type locality: From oral side of "sand dollars," Mud flat.
BaledesIsoles,Noumea,NewCaledonia.
New material: Northern New Caledonia, N/O Alls sta-
tion DW 1181, 19°23.9'S, 163°14.7'E, 45 m, 31 October
Page 146 The Veliger, Vol. 37, No. 2

Explanation of Figure 4A-E

Figure4A-E.HypermaslusorstomiWaren,sp.nov,anditshostMaretiaplanulata.A,C.Paratypes,heightofshells
5.5mm.B.Protoconch,height240^m.D.Aboralviewoftest,hostofonespecimenofHypermastusorstomiWaren,
sp.nov.Areaofattachmentindicatedbyfinearrows.Diameteroftest46mm.E.Enlargementofcentralpartof
test,showingapicalareawithfourgonoporesandtheareaofattachment(finearrows)fortheproximalpartof
theproboscis.Penetrationoftest(diameter0.1mm)markedbyslightlylargerarrow.Horizontalwidthofpicture
mm.
7
A. Waren et al., 1994 Page 147

Explanation of Figure 5A-D

Figure5A-B."Balcis"echinocardiaphilaHabe,syntypesNationalScienceMuseum,Tokyo,registernumber52460,
heightofshell13.1and11.9mmrespectively.C-D.Clypeastericolaclypeastericola(Habe),syntype.National
ScienceMuseum,Tokyo,registernumber52468,heightofshell14.2mm.

1989,onespecimenattachedorallyonLaganumdepressum
L. Agassiz, 1841 (Laganidae); New Caledonia, Lagoon
de Noumea, Quatre Bancs de I'Ouest, 22°25.36'S,
166°27.77'E,12mdepth,sandbottomwithHalimedaand
Sargassum,10L.depressumparasitizedonoralsideamong
ca. 350 specimens examined 19 April 1993 (P. Bouchet,
LaboratoiredeBiologicdesInvertebresMarinsetdeMa-
lacologie,MuseumNationald'HistoireNaturelle,Paris).
Distribution:Onlyknownfromtheholotypeandthepres-
entspecimens,allfromNewCaledonia.
Remarks: This species was previously known from the
holotype,foundonanunidentifiedsanddollarfromNew
Caledonia. The new material makes it possible to give
someadditionalinformationandtodescribethemorphol-
ogyofthehead-foot,whichisrathersmallbutfunctional.
The tentacles are long, smooth, regularly conical, with
largeblackeyesinindistinctlateralbulgesonthesidesof
thebases.Theinnersidesmeetcentrallyovertheproboscis
opening. The foot is small, broader posteriorly, with an
inconspicuouspropodiumandalargeandconspicuous
openingfortheposteriorpedalgland.Themetapodium
formsaconspicuousridgeontherightsideandanalmost
tongue-shapedflapontheleftside,considerablylarger
thantheleftone.Theproximalpartoftheproboscissheath
islargeandvoluminousandendswithasucker-shaped
discwhichisusedtoattachthesnailtothetestofthehost.
Fromtheundersideofthisdiscexitsthemuchmoreslender
distalpartofthesheath,whichpenetratesthetest,ishighly
extensible, and may be extended two or three times the
lengthoftheshell.
Twospecimenshadoriginallybeenparasitizingthehost
collectedin1989,closetoeachother(Figure6E),butone
ofthemhadevidentlybeentornofforleftthehost.When
Warenreceivedthespecimen,thesecondspecimenhad
alsofallenofT.
Ofthe10hostscollectedin1993,eighthostshadasingle
scar,onehadtwo,andonehadthreescars,whichmeans
that the observed rate of infestation by more than one
specimenisconsiderablyhigherthanarandomdistribution
admits.Thesizerangeofthe11specimens(twowerelost
whencollected)is8.1-5.8mm;asinglespecimenmeasures
Page 148 The Veliger, Vol. 37, No. 2

Explanation of Figure 6A-F

Figure6A-F.HypermastusobliqmstomumanditshostLaganumdepressum.A.Assumedmale4.2mmhigh.B.
Female,8.3mmhigh.C.Assumedsecondaryfemale,6.4mmhigh.D.Larvalshell,height370mm.E.Critical
pointdriedhead-footofH.obliqmstomum;pallialskirtandvisceralmassremoved.Widthoffoot0.25mm.F.
Laganumdepressum,oralsidewithH.obliqmstomum.Diameteroftest60mm.AD—attachmentdiscofproboscis;
MP—metapodial(opercular)fold;OP—operculum;P—propodium;PG—posteriorpedalgland;PS—proximal
partofproboscissheath;T—cephalictentacles.
A. Waren et al., 1994
4.15 mm. Among the larger ones, there is a distinct di-
morphisminshellshape,somespecimensbeingconsid-
erably more slender (Figure 6C). One specimen of each
typewasdecalcifiedandsexed.Thelessslenderonewas
afemale,themoreslenderonewasalsodeterminedtobe
afemale,butwithavestigialpenis.Bothspecimenshad
well-developed pallial oviducts. This may indicate that
somespecimensdevelopdirectlytofemales,whileothers
passthroughamaleperiod.
Thesitesofattachmentareindicatedbynarrowholes
in the test with a diameter of about 0.1 mm. The hole is
surroundedbyascarwheretheproboscissheathhasbeen
attached, and a much larger, 4-8 mm diameter, surface
whichisdenudedofspinesandsomewhatblackened.The
scarfromthesuckerisusuallysurroundedbyadark,small
mucus collar.
Ahostspecimenwithoneemptyscarandonespecimen
of//,obliquistomumstillattachedwasdissected,andthere
werestillslenderproboscidesremaininginsidethetest,
underbothholes,bothofthemendingfreelyinthecavities
ofthegonads.
Thethick,proximalpartoftheprobosciscanevidently
beretractedinsidethecephalichaemocoelandmustthere-
fore be considered a part of the proboscis sheath, not a
snout, which it could be mistaken for when examined in
anevertedstate.Thedamageofthetestofthehostindicates
thatatleastthefemalesstayforaconsiderableperiodof
timeonthesamespotandonlyoccasionallyleavethehost.
HypermastusobliquistomumresemblesH.colmaniWar-
en,1991,presumedtoparasitizePeronellaleseuri(Valen-
ciennes,1841)(Laganidae)inAustralia.Thatspeciesdoes,
however,difTerinhavingatallerprotoconchwiththree
convexwhorlsandaheightof0.46mm.
Eulima Risso, 1826
EulimaRisso,1826:123.Typespecies;TurbosubulalusDon-
ovan,1804(=Eulimaglabra(DaCosta,1778))(Waren
1992;ICZN,Opinion1718),European.Parasiticon
ophiuroids.
Remarks: The generic name Eulima has been used for a
broadrangeofeulimidsintheliterature,butWaren(1983)
gavesomeinformationaboutthetypespeciesandavery
similar species, E. bilineata Alder. These species have a
ptenoglossateradula,usedforcatchingcoelomocytesinthe
coelomiccavitiesofthehosts(unpublished).Thenewspe-
cies below differs in parasitizing a sand dollar echinoid
and in lacking a radula. Otherwise it is very similar, in
shell characters and in being a hermaphrodite of some
kind. We have therefore decided to keep this species in
Eulima, although it is likely that it will need a new genus.
Eulima encopicola Waren & Templado, sp. nov.
(Figures 7A-E)
Type material: Holotype and the single paratype in Mu-
Page 149
seoNacionaldeCienciasNaturales,Madrid,Spain,reg-
isternumber15.05/6958.
Typelocality:GalapagosIslands,SantiagoIsland(James
Island), Sulivan Bay, Bartolome inlet. 19 March 1991.
On a specimen of Encope micropora galapagensis A. H.
Clark, 1946 (Encopidae), the larger one, holotype, (4.76
mm)attachedontheoralside;thesmallerone(3.04mm)
ontheaboralside.
Description: Shell (Figure 7A-D) tall, conical, slender,
transparentwithsomebrownishcolorinthesubsutural
zone and on the body whorl. The larval shell (Figure 7E)
isblunt,evenlyrounded,notobviouslycolored,consistsof
slightlymorethanonewhorlandhasaheightof330^.m.
Theholotypehas7.5almostflatteleoconchwhorls,sculp-
turedwithdistinct,sharp,close-set,butnotveryuniformly
spaced incremental lines, and six irregularly spaced in-
crementalscars(Figure7D)ofvaryingstrength.Thesub-
suturalzoneisconspicuous,anditsheightcorrespondsto
about 25% of the height of the whorl. The aperture is tall
and slender with a sturdy columellar callus and a thin
parietalcallus,whichtogetherformadistinctangle.The
apical corner of the aperture is distinctly drawn out and
constricted. The outer lip is slightly damaged in the ho-
lotype,buthasevidentlybeenverystraightandisslightly
prosocline.
Dimensions.Heightofholotype4.76mm,diameter1.40
mm.
Operculum.Verythin,fragileandcolorlesswithnore-
inforcements.
Softparts.Thesoftparts(preservedinalcohol)havea
pinkish hue. The large specimen has a well-developed
oviduct and penis; the small one only a penis. The eyes
areunusuallysmall,diameterabout20^l^aandsituated
about100jumapart.Nootherdetailswereseensincethe
specimensweredriedandreconstitutedtofacilitatethe
extraction of the soft parts for radular preparation. The
anteriorpedalglandisverylargeandinvadesthecephalo-
pedalhaemocoel.Noradulawasfoundinanyofthespec-
imens.
Etymology:Namedafterthegenericnameofitshost.
Remarks:StrombiformisbarthelowiBartsch,1917(Figure
7F-G), from the Gulf of California, is of a similar size
and shape, but the initial whorl of the protoconch (I) is
very small, the larval shell is distinctly conical with 2-3
whorls, and the whole shell is indistinctly yellowish (not
easytoseeintheholotypebecauseoftheremainingdried
soft
parts).
Clypeastericola Waren, gen. nov.
Typespecies:BalcisclypeastericolaHabe,1976,fromJapan
(Figure5C-D),parasiticonClypeasterjaponicusDoe-
derlein,
1885
(Clypeasteridae).
Diagnosis:Eulimidswithacolorless,slender,smooth,and
conicalshell.Aperturelow,constrictedinfemale,lessso
Page 150 The Veliger, Vol. 37, No. 2

Explanation of Figure 7A-G

Figure7A-E.EulimaencopicolaWaren&Templado,sp.nov.A,C.Holotype(female?),heightofshell4.76mm.
B.Paratype,3.04mm.D.Incrementalscar,length0.75mm.E.Larvalshell,labialscarindicatedbyarrows,height
ofprotoconch330fim.F-G.Eulimabarlhelowi(Bartsch),holotype,USNM268622,heightofshell4.93mm.
A. Waren et al., 1994 Page 151

Explanation of Figure 8A-F

Figure8A-F.ClypeastencolanatalensisWaren,sp.nov.anditshostClypeastereurychoreus.A.Holotype(female?),
6.4mm.B.Youngfemale,3.9mm.C.Male,4.7mm.D.Larvalshellmarkedwithanarrow.Height460^m.E-
F.Host,Clypeastereurychoreusdiameter
, 41mm,withremainingproboscidesofagroupofonefemaleandtwo
malesandasecondfemale.E.DetailofF,showingthetornoffproboscides.

inmale.Probosciswithanattachmentdisc,moredeveloped clypeastencolaandCnatalensisbutonthesoftpartsofC.
in
female. natalensis,sinceC.clypeastencolahasbeenavailableonly
Etymology:NamedafterClypeaster,thehostgenusforat asemptyshells.Theshellsare,however,verysimilar,and
least
three
species. wefeelcertainthattheyarecongeneric.Thesimilarityin
the shape of the aperture of the females, a small dip in
Remarks:ThediagnosisaboveisbasedontheshellofC. thesuture,andacorrespondingconstrictionoftheperi-
Page 152 The Veliger, Vol. 37, No. 2

Explanation of Figure 9A-E

Figure9A-E.Clypeastericolasp.andscarsonitshostClypeasteraustralasiae.A,C.Height4.9mm.B.Larval
shell,heightofprotoconch520nm.D-E.DetailsoftestofClypeasteraustralasiaewithscarsfromClypeastericola
sp.Scalelines1mm.

stomeisararefeature,obviousonlyinthegenusParvwris Clypeastericola natalensis Waren, sp. nov.

Waren1981.Thosespecieshaveanoperculumwithafold
that is deeply inserted in the foot, and they are parasitic
on
asteroids.
Thetwoundescribedspeciesdiscussedbelowhavesim-
ilar shells but are immature, judging from the angular
peripheryofthebodywhorl.
(Figures 8A-F)
Type material: Holotype, a female, Natal Museum, Pie-
termaritzburg,registernumberNMPD8634/T841;three
paratypes,oneimmaturefemaleandtwomales,S7878/
T842.
A. Waren et al., 1994
Explanation of Figure lOA-C
FigurelOA-C.Clypeastericolasp.fromLaganumdepressum.A.Frontview,4.0mmhigh.B.Protoconch,height
350nm.C.Sideview,height3.8mm.
Type locality: Eastern South Africa, Northern Zululand,
southeastofKosiEstuary,26°24.1'S,32°34.8'E,50m,fine
sand. 08 June 1987, on a small Clypeaster eurychoreus
Clark,1924(Clypeasteridae).Oneimmaturefemalewas
attachedalone,oneadultfemalewasaccompaniedbytwo
males.Position,seeFigure8F.
Etymology:NamedaftertheNatalprovince.
Description:Shell(Figure8A-C)conical,ratherinflated,
whitishsemitransparent,rathersolid,smoothandshiny.
The conical, tall, and pointed larval shell (Figure 8D)
consists of about three whorls and its height is 460 ^ln\.
The mature female has 8.5 slightly convex whorls. The
sutureisdeepandconspicuous;thefalsesutureispoorly
definedandhardlynoticeable.Thefemalehastwoincre-
mental scars, 1.1 and 2.2 whorls from the outer lip. The
shellhassomeirregularandindistinctgrowthlinesanda
very irregular and weak spiral striation. The aperture is
shortandbroad,slightlycontracted.Theparietalcallusis
well-developed,butthereisnodistinctanglebetweenthe
parietal wall and the columella. The outer lip is slightly
retractedatthesutureandveryslightlyprosocline.
Page 153
Dimensions.Heightoftheholotype6.39mm,breadth
2.44mm.
Male. Shell slightly more slender and with a single in-
crementalscaraboutonewhorlfromtheouterlip.Height
3.88and4.64mm.
Operculum.Ratherthin,slightlyyellowish,withadis-
tinctreinforcementribontheinside.
Softparts.(Ofimmaturefemale.)Theoviductisbegin-
ningtodevelopasamoresolidzonealongtherightcorner
of the pallial cavity. The tentacles are long and tapering
withlargeeyes,100^mdiameterand200Atmapart,placed
wellbehindthecleavagebetweenthetentacles.Thefoot
is small, smaller than the opercular lobes. The right side
oftheanteriorpartofthehead-foothasanepipodialfold
endingattheproboscisopening.Themetapodiumbulges
out widely at both sides of the foot; the right one is con-
nectedtotherightepipodialfold.Thepropodiumiswell-
developed.Thereisnotraceofapenis.
The proboscis is firmly attached to the test of the host
(Figure 8E), over a hole of a tube-foot, which is used for
penetrating the test. All proboscides were so firmly at-
tachedthattheyhadbeentornoff,abovetheattachment
Page 154
disc. This disc is much broader in the female than in the
male.Theproboscisoftheyoungfemaleendedfreelyin
thecoelomiccavity.Theproboscisofthelargefemalewas
insertedbetweenthecoelomicepitheliumandthetestfor
10mmuntilreachingagonad,whichwaspenetrated.
Remarks:Probablytheepi-andmetapodialfoldscanbe
used to cover the base of the shells to avoid irritation of
thehost,butthespecimenshadneverthelesscausedsome
damageonthetest,whichwasdenudedaroundthesnails.
ClypeastericolanatalensisdiffersfromC.clypeasten-
colainhavingashorterandproportionallybroadershell
(cf. Figures 8A-C and 5C-D).
Clypeastericolaspp.
(Figures 9A-E, lOA-C)
Material examined: (Species 1.) Australia, New South
Wales,offPortHacking,onClypeasteraustralasiae(Gray,
1851) (Clypeasteridae), taken by a trawler, one young
specimenattachedtotest,AMSC108275.
(Species2.)NewCaledonia,LagondeNoumea,Grande
Rade, 22°15.26'S, 166°23.98'E, 11-12 m, sand with ooze
on surface, 14 April 1993, two immature specimens at-
tachedonaboralsideofLaganumdepressum(P.Bouchet,
LaboratoiredeBiologicdesInvertebresMarinsetdeMa-
lacologie.MuseumNationald'HistoireNaturelle,Paris).
Remarks: The test of the Australian host had three per-
forations (two figured in Figure 9D-E), indicating that it
had been the host of at least that number of specimens.
The remaining one is figured in Figure 9A-C.
TheNewCaledonianspeciesdiffersinbeingmoreslen-
derwithflatterwhorlsandinhavingasmallerprotoconch
withlessthantwowhorls(almostthreeintheAustralian
specimen).
Wefindtheseyoungspecimensworthmentioningbut
notnaming.Theshellsarequitesimilartothetwospecies
ofClypeastericoladescribedabove,andcontributetothe
reasonsforthisintroductionofanewname.
The Veliger, Vol. 37, No. 2
ACKNOWLEDGMENTS
We thank the Malacology sections at the Museum Na-
tional D'Histoire Naturelle, Paris, the Australian Mu-
seum,Sydney,andtheNatalMuseum,Pietermaritzburg,
SouthAfrica,whocontributedspecimensforthisstudy.
ChristineHammar,Stockholm,preparedFigure3andthe
prints used for the plates. R. Rabago, P. Rabago, and C.
I. Norris (Guam) assisted in the collection of specimens
in Guam. This is Contribution 344 from the University
ofGuamMarineLaboratory.
LITERATURE CITED
Alekseev, a. S. & L. G. Endelman. 1989. Association of
ectoparasitic prosobranchgastropods with
UpperCretaceous
echinoidGalentes.Pp.165-174In:FossilandRecentEchi-
nodermResearches.AcademyofSciencesoftheEstonian
SSR.
Habe,T.1976.ParasiticgastropodsfromechinodermsofJa-
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Habe,T.1992.Anewspecies,Hypermastusaraesomaeparasitic
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ICZN,Opinion1718.1993.fia/eaGray,1824(Mollusca:Gas-
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