II11 April 2000 PROCEEDINGS OF THE BIOLOGICALBIOLOGICAL SOCIETYSOCIETY OFOF WASlllNGTONWASHINGTON 113(1):145-154.2000.113(1): 145-154. 2000. Additions to thethe cancellariidcancellariid (Mollusca:(Mollusca: Neogastropoda)Neogastropoda) faunafauna ofof South Africa

Richard E. Petit andand M.M. G.G. HarasewychHarasewych Department of InvertebrateInvertebrate Zoology,Zoology, NationalNational MuseumMuseum ofof NaturalNatural History,History, Smithsonian Institution,Institution, Washington,Washington, D.C.DC. 20560-0118,20560-0118, U.S.A.U.S.A.

Abstract.Abstract.-Four—Four new speciesspecies ofof CancellariidaeCancellariidae areare describeddescribed fromfrom thethe con­con- tinental shelf andand upperupper continentalcontinental slopeslope offoff easterneastern SouthSouth Africa.Africa. AdmetulaAdmetula afra is distinguished fromfrom allall congenerscongeners byby itsits combinationcombination ofofsmall small adultadultsize size «(< 11 mm), rounded shoulder,shoulder, evenlyevenly reticulatereticulate sculpture,sculpture, andand distinctdistinct varixvarixat at the juncture ofof protoconch andand teleoconch.teleoconch. TrigonostomaTrigonostoma kilburnikilburni differsdiffersfrom from all other TrigonostomaTrigonostoma inin itsits distinctivedistinctive shellshell outline,outline, unornamentedunornamentedperipheral peripheral keel, rounded ratherrather thanthan tabulatetabulate shoulder,shoulder, andand veryvery narrownarrow umbilicus.umbilicus. Nip­Nip- ponaphera wallacei differsdiffers fromfrom N.N. paucicostatapaucicostata (Sowerby,(Sowerby, 1894),1894), itsits geo­geo- graphically closest congenercongener fromfrom thethe ArabianArabian SeaSea area,area, inin beingbeing umbilicateumbilicate and in having aa moremore roundedrounded shouldel~shoulder, andand moremore numerousnumerous andand finerfiner spiralspiral cords. Its frequentfrequent associationassociation withwith thethe turbinidturbinid BolmaBolma andersoniandersoni suggestssuggeststhat that it may be an ectoparasite ofof thisthis species.species. ZeadmeteZeadmete verheckeniverheckeni isis mostmostsimilar similar to Zeadmete subantarctica Powell,Powell, 1933,1933, fromfrom offoff NewNew Zealand,Zealand, fromfrom whichwhich it can be distinguished byby itsits lowerlower spirespire andand weakerweaker surfacesurface sculpture.sculpture.Among Among the South African taxa,taxa, Z.Z. verheckeniverheckeni mostmost closelyclosely resemblesresembles "Cancellaria""Cancellaria" eutrios Barnard, 1959,1959, fromfrom whichwhich itit isis easilyeasily distinguisheddistinguished byby itsits tabulatetabulate shoulder and lowerlower spire.spire. TheThe geographicgeographic andand bathymetricbathymetric rangesranges ofofAdmetula Admetula epula Petit & Harasewych, 1991,1991, aa speciesspecies previouslypreviously knownknown onlyonly fromfrom "ex"ex pisces" material, havehave beenbeen expandedexpanded basedbased onon live-collectedlive-collected specimens.specimens.

The Cancellariidae comprisescomprises aa familyfamily ofof AllAll typetype specimensspecimens arearehoused housedin inthe thecol­ col- diverse and highly specialized,specialized, suctorialsuctorial lectionslections ofof thethe NatalNatal MuseumMuseum (NM),(NM), Pie­Pie- neogastropods thatthat inhabitinhabit softsoft bottom,bottom, sub­sub- termaritzburg,termaritzburg, RepublicRepublic ofofSouth South Africa.Africa, tidal to bathyal habitatshabitats throughoutthroughout tropicaltropical and temperate seas.seas. TheThe cancellariidcancellariid faunafauna FamilyFamily CancellariidaeCancellariidaeForbes Forbes& & Hanley,Hanley, of South Africa was reviewedreviewed comprehen­comprehen- 18511851 sively by Barnard (1959)(1959) andand KensleyKensley Genus Admetula Cossmann, 18891889 (1973), and toto aa limitedlimited extentextent moremore re­re- cently in popular works byby RichardsRichards (1981)(1981) TypeType species:species: CancellariaCancellariaevulsa evulsa(Solan­ (Solan- and Steyn & Lussi (1998).(1998). der,der, 1766) 1766) (=Buccinum(=Buccinum evulsumevulsum SolSolander,ander, This paper describes fourfour newnew speciesspecies ofof 1766)1766) byby originaloriginal designation.designation, cancellariids collected inin SouthSouth AfricanAfrican wa­wa- ters by SCUBA andand byby thethe vesselsvessels RIVR/V AdmetulaAdmetula epulaepula PetitPetit && Harasewych,Harasewych, 19911991 Meiring NaudiNaude (1984-1988)(1984-1988) andand NMPDNMPD Figs.Figs. 1-2,1-2, 1818 Africana (1995). TheseThese speciesspecies areare assignedassigned . , , , _ . _ __ , . , . f, . ... AdmetulaAdmetula epulaepulaPetit Petit& &Harasewych, Harasewych, to the genera Admetula, Trigonostoma,Trigonostoma, Nip­Nip- . „, , _, _ , „ 1991:181,I 991181 figs.figs 1-3.1—3 ponaphera, and Zeadmete. TheThe newnew taxataxa are compared toto relatedrelated speciesspecies fromfrom thethe Diagnosis.-ADiagnosis.—A small small species species with with an an Indian Ocean, New Zealand,Zealand, andand Australia.Australia, ovatelyovately conicalconical shell.shell. TransitionTransition fromfrom pro-pro- 146 PROCEEDINGS OF THE BIOLOGICALBIOLOGICAL SOCIETYSOCIETY OFOF WASHINGTONWASHINGTON toconch to teleoconchteleoconch gradual,gradual, indicatedindicated byby Admetula afra, new speciesspecies appearance firstfirst ofof spiral,spiral, thenthen axialaxial sculp­sculp- Figs. 3-4, 1818 ture. Teleoconch sculpturesculpture ofof strongstrong spiralspiral cords and lessless pronounced,pronounced, roundedrounded axialaxial Diagnosis.Diagnosis.-A—A smallsmall speciesspecies withwith aa broadly conical shell.shell. ProtoconchProtoconch demar­demar- ribs. Outer liplip thin,thin, smoothsmooth within.within. cated fromfrom teleoconchteleoconch byby broad,broad, roundedrounded Gross anatomy.-Preservedanatomy.—Preserved animalanimal yel­yel- varix. Teleoconch withwith sharplysharply reticulatedreticulated lowish tan, footfoot long,long, narrow,narrow, taperingtapering pos­pos- sculpture. Outer liplip reflectedreflected posteriorly,posteriorly, teriorly. Mantle cavitycavity spanningspanning 0.670.67 whorl.whorl. with weak Iiraelirae beneath spiralspiral cords.cords. Osphradium slightly broaderbroader thanthan ctenid­ctenid- Description.Description.-Shell—Shell (Fig.(Fig. 3)3) small,small, reach­reach- ium. Pericardium veryvery small.small. TentaclesTentacles ing 9.0 mm, ovately conicalconical withwith roundedrounded symmetrical, bluntly cylindrical,cylindrical, flankingflanking anterior. Protoconch (Fig.(Fig. 4)4) ofof 1.671.67 central rostrum. EyesEyes small,small, black.black. Probos­Probos- smooth, inflated whorls deflecteddeflected fromfrom coil­coil- cis short, about 0.670.67 lengthlength ofof mantlemantle cav­cav- ing axis of teleoconch byby aboutabout 7°.7°. Transi­Transi- ity. Buccal mass large,large, nearlynearly fillingfilling retract­retract- tion to teleoconch (Fig.(Fig. 4,4, arrow)arrow) demar­demar- ed proboscis. SalivarySalivary glandsglands andand accessoryaccessory cated by prominently roundedrounded varix,varix, fol­fol- salivary glands inin cephaliccephalic haemocoel,haemocoel, notnot lowed immediately byby onsetonset ofof bothboth spiralspiral contained within proboscis.proboscis. PenisPenis long,long, nar­nar- and axial sculpture. TeleoconchTeleoconch ofof 2.75-32.75—3 row, dorsoventrally compressed,compressed, distaldistal endend evenly rounded whorls. SutureSuture weaklyweakly im­im- bluntly rounded. pressed. Shoulder rounded,rounded, weaklyweakly defined.defined. Remarks.Remarks.-This—This speciesspecies waswas originallyoriginally Axial sculpture consistsconsists ofof narrow,narrow, regular­regular- described based onon sevenseven specimensspecimens takentaken ly spaced, weakly prosocline ribsribs (14-16(14—16 from the stomachs ofof fish.fish. ItsIts bathymetricbathymetric on body whorl). SpiralSpiral sculpturesculpture ofof narrow,narrow, range was inferredinferred toto overlapoverlap withwith thatthat ofof sharply defined primaryprimary cordscords (12-13(12-13 onon CongiopodusCongiopodits spinijerspinifer (Smith)(Smith) (55-146(55-146 m),m), body whorl, 6 on penultimatepenultimate whorl).whorl). Sin­Sin- one of the fishfish fromfrom whichwhich itit waswas taken.taken. gle, weak, secondary cordscords maymay bebe presentpresent Among the material collectedcollected byby thethe between primary cordscords onon posteriorposterior halfhalf ofof NMPD Africana [sta.[sta. A17419D]A17419D] waswas aa livelive body whorl. Spiral andand axialaxial sculpturesculpture in­in- collected specimen ofof A.A. epulaepula (Figs.(Figs. 1-2,1-2, tersect to formform sharplysharply reticulatedreticulated patternpattern 18) trawled at aa depthdepth ofof 210210 mm offoff thethe with small nodes at intersectionsintersections ofof ribsribs andand mouth of the Tsitsikamma River.River. cords. Aperture broadlybroadly ovate,ovate, deflecteddeflected We have examined aa specimenspecimen ofof Ad­Ad- from coiling axis by 16-19°.16-19°. OuterOuter liplip thin,thin, slightly flared posteriorly, weaklyweakly liratelirate be­be- metula fromfrom deeper waterwater (450-500(450-500 m)m) offoff neath spiral cords.cords. ParietalParietal calluscallus thin,thin, the Mbashe River, TranskeiTranskei (R/V(R/V MeiringMeiring translucent, overlying 5-65-6 spiralspiral cordscords fromfrom NaudeNaudi sta. Q14), between thethe typetype localitieslocalities previous whorl. Columella formingforming angleangle ofof of A. epula and thethe newnew species.species. ThisThis spec­spec- 128-130° with parietalparietal region,region, thick,thick, imen (NM C9050) isis tootoo wornworn forfor thethe tran­tran- straight, with twotwo weakweak columellarcolumellar foldsfolds sition from protoconch toto teleoconchteleoconch toto bebe and broad siphonalsiphonal fold.fold. SiphonalSiphonal canalcanal clearly discerned, butbut therethere isis nono indicationindication shallow but well-defined. ShellShell colorcolor white.white. of a pronounced varix. BecauseBecause itit alsoalso hashas Type locality.-SElocality.—SE ofof PortPort Durnford,Durnford, secondary spiral cords betweenbetween allall primaryprimary South Africa (29°01.5'S,(29°01.5'S, 32°11.8'E),32°11.8'E), dredgeddredged cords, this specimen isis tentativelytentatively identifiedidentified in 310-320310—320 m, glutinous sandysandy mud.mud. R/VR/V as A. epula. This recordrecord expandsexpands thethe geo­geo- Meiring NaudiNaude sta.sta. ZQ9,ZQ9, 66 JulJul 1985.1985. graphic range of A. epulaepula fromfrom CapeCape St.St. Type material.-Holotype,material.—Holotype, NMNM E3189/E3189/ Blaize to Transkei, offoff ofof thethe mouthmouth ofof thethe T1382, 8.0 mm; ParatypeParatype 1,1, 7.87.8 mm,mm, andand Mbashe River (32°22.8'S,(32°22.8'S, 29°00.8'E).29°00.8'E). TheThe ParaParatypetype 2, 6.9 nun,mm, NMNM V7006rr1383V7006/T1383 fromfrom bathymetric range isis extendedextended wellwell intointo thethe the type locality; ParaParatypetype 3,3, 9.09.0 mm,mm, NMNM bathyal zone (450(450 m).m). E3764, fromfrom off Cape VidalVidal (28°08.4'S,(28°08.4'S, VOLUME 1I 13,13, NUMBER 1I 147

Figs. 1-4. 1-2;1-2; AdmetulaAdmetula epulaepulu PetitPetit && Harasewych,Harasewych. 1991.1991. OffOffTsitsikamma TsilsikammaRiver, River,South SouthAfrica Africa(34°45'S, (34°45'S. 24°47'E) in 210 tn,m, sand,sand, oldold shellshell gritgrit andand shellshell debris.debris. NMPDNMPDAfricana Africanasta. sla.A17419D. A17419D.I. 1.Apertural Aperturalview viewof of shell. 2. Apical viewview ofof protoconch.protoconch. 3-4.3-4. Adme/ulaAdmetula afraafra newnew species.species.Holotype, Holotype,NM NME3189fT E3189/T1382,1382, SE SEof ofPort Port Durnford, South Africa (29°01.5'(29°01.5'S,S, 32°32°1I J1.8'E),.8' E), dredgeddredgedin in310-320 310-320m. m,glutinous glutinoussandy sandymud. mud.RN R/VMeiring MeiringNaude Naude sta.sla. ZQ9, 6 luIJul 1985.1985. 3.3. AperturalApertural viewview ofof holotype.holotype. 4.4. ApicalApical viewviewof ofprotoconch. protoconch.

32°36.4'E), dredged inin 165165 m,m, moderatelymoderately Comparative remarks.-Thisremarks.—This newnew spe­spe- fine sand, RNR/V Meiring NaudeNaude sta.sta. ZM8,ZM8, 1111 cies appears most closelyclosely relatedrelated toto thethe June 1988. more southern Admetula epulaepula PetitPetit && Har­Har- Etymology.-FeminineEtymology.—Feminine formform ofof thethe LatinLatin asewych, 1991,1991, fromfrom whichwhich itit differsdiffers inin afer, African. having uniformly rectangularrectangular sculpturesculpture 148 PROCEEDINGS OF THE BIOLOGICALBtOLOGICAL SOCIETYSOCIETY OFOF WWASHINGTONASHTNGTON formed by equally-sized axialaxial ribsribs andand spi­spi- ed from coiling axisaxis byby 18-19°.18—19°. SiphonalSiphonal ral cords, andand aa posteriorlyposteriorly reflectedreflected outerouter canal short, broad, barelybarely discerniblediscernible exceptexcept lip with weak denticles beneathbeneath thethe spiralspiral externally as thethe siphonalsiphonal fasciole.fasciole. OuterOuter liplip cords. The most strikingstriking differencedifference betweenbetween thin, smooth within,within, slightlyslightly reflectedreflected withwith these two species isis thethe presencepresence ofof aa strongstrong spiral cords visible throughthrough edgeedge ofof lip.lip. varix at the terminationtermination ofof thethe protoconchprotoconch Posterior portion ofof innerinner liplip adpressedadpressed in A. afra. InIn contrast,contrast, thethe transitiontransition fromfrom against siphonal fasciole.fasciole. ShortShort parietalparietal re­re- protoconch toto teleoconchteleoconch isis indistinctindistinct andand gion forms angle ofof 150°150° withwith long,long, slightlyslightly gradual inin Admetula epula.epula. concave columella thatthat bearsbears 22 weak,weak, wide­wide- ly spaced columellar foldsfolds andand 11 siphonalsiphonal Genus Trigonostoma Blainville,Blainville, 18271827 fold. UmbilicusUmbilicus deep, narrow.narrow. ShellShell white,white, Type species: Delphinula trigonostomatrigonostoma sometimes with aa yellowishyellowish cast.cast. Lamarck, 18221822 (?=Bueeinum{p.—Buccinum sealarescalare Gme­Gme- Radular teeth (Figs.(Figs. 9-10)9—10) extremelyextremely lin, 1791) by monotypy. long, ribbon-like, tricuspid.tricuspid. CentralCentral cuspcusp smooth, with recurvedrecurved rim.rim. LateralLateral cuspscusps Trigonostoma kilburni, newnew speciesspecies long, foldedfolded towardtoward centralcentral cusp,cusp, eacheach withwith Figs. 5-10, 1818 four, anteriorly directeddirected secondarysecondary cusps.cusps. Diagnosis.Diagnosis.-A—A smallsmall speciesspecies withwith aa thin,thin, Second most proximal secondarysecondary cuspcusp bi­bi- angular, narrowly umbilicateumbilicate shell.shell. Shoul­Shoul- fid. der rounded, not tabulate,tabulate, lackinglacking pro­pro- Type loeality.-Offlocality.—Off EastEast London,London, SouthSouth nounced spines. Axial sculpturesculpture ofof numer­numer- Africa (33°04.7'S, 28°07.2'E)28°07.2'E) dredgeddredged inin 9090 ous scabrous varices. OuterOuter liplip smooth,smooth, m, associated with coarsecoarse sand,sand, sponges,sponges, lacking lirae. gorgonians. RIVR/V Meiring NaudeNaude sta.sta. XX46,XX46, Description.Description.-Shell—Shell (Fig.(Fig. 5)5) small,small, reach­reach- 17 Jul 1984.1984. ing 14.3 mm, thin,thin, angular,angular, biconical,biconical, Type material.-Holotype,material.—Holotype, NMNM D679,D679, strongly shouldered,shouldered, with deep,deep, narrownarrow um­um- 12.9 mm, fromfrom typetype locality.locality. Paratype,Paratype, NMNM bilicus. Spire high (spire(spire angleangle 55°),55°), com­com- D680, 14.4 mm, offoff Kidd'sKidd's Beach,Beach, SouthSouth prising over half ofof shellshell length.length. ProtoconchProtoconch Africa (33°11.8'S, 28°03.2'E)28°03.2'E) dredgeddredged inin 9090 (Figs. 6-7) of 2 smoothsmooth whorls,whorls, offsetoffset fromfrom m, associated with coarsecoarse sand,sand, sponges.sponges. RIR/ coiling axis of shellshell byby aboutabout 5°.5°. TransitionTransition V Meiring Naude sta.sta. XX50,XX50, 1717 JulJul 1984.1984. to teleoconch abrupt, markedmarked byby aa slightlyslightly Etymology.Etymology.-This—This speciesspecies honorshonors Dr.Dr. Ri­Ri- flared varix and thethe onsetonset ofof spiralspiral sculp­sculp- chard N. Kilburn, NatalNatal Museum,Museum, Pieter­Pieter- ture. Teleoconch ofof upup toto 55 sharplysharply angularangular maritzburg, Republic ofof SouthSouth Africa,Africa, forfor whorls. Suture deeply impressedimpressed behindbehind thethe his many contributions toto malacology.malacology. evenly roundedrounded shouldershoulder delineateddelineated byby aa Comparative remarks.-remarks.—ThisThis newnew spe­spe- sharp keel along thethe periphery ofof thethe shell.shell. cies differs fromfrom allall otherother TrigonostomaTrigonostoma inin Axial sculpture consistsconsists ofof numerous,numerous, its distinctive shellshell outline.outline. ItIt maymay bebe dis­dis- flared, weakly prosoclineprosocline varicesvarices (Fig.(Fig. 8),8), tinguished fromfrom TrigonostomaTrigonostoma sealarescalare regularly spaced onon earlyearly whorlswhorls (about(about 16­16- (Gmelin, 1791) and T. thysthlon (Petit & 18 on firstfirst teleoconchteleoconch whorl),whorl), increasingincreasing inin (Gmelin, 1791) and T. thysthlon (Petit & number and becoming moremore irregularlyirregularly Harasewych, 1987)1987) byby itsits unornamentedunornamented spaced in subsequent whorls.whorls. SpiralSpiral sculp­sculp- peripheral keel,keel, itsits roundedrounded ratherrather thanthan tab­tab- ture of broad, crisplycrisply demarcateddemarcated primaryprimary ulate shoulder, andand by itsits veryvery narrownarrow um­um- spiral cords (Fig.(Fig. 8,8, p),p), withwith 1-31-3 slightlyslightly bilicus. The only otherother SouthSouth AfricanAfrican Tri­Tri- narrower secondary cordscords (Fig.(Fig. 8,8, s)s) be­be- gonostoma isis thethe commoncommon shallow-watershallow-water T.T. tween adjacent primary cords,cords, andand muchmuch semidisjunctasemidisjuneta (Sowerby,(Sowerby, 1849),1849), whichwhich hashas finer spiral threadsthreads (Fig.(Fig. 8,8, t)t) betweenbetween somesome a heavier, more roundedrounded shellshell withwith strongstrong cords. Aperture roundlyroundly triangular,triangular, deflect-deflect- spiral cords. VOLUME 113, NUMBER 1] 149

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Figs. 5-8. Trigonostoma kilburni new species.species. Holotype,Holotype, NMNM D679,D679, OffOff easteast London,London, SouthSouth AfricaAfrica (33°04.7'S, 28°07.2'E), dredged inin 90 m, coarsecoarse sand,sand, sponges,sponges, gorgonians.gorgonians. RIYR/V MeiringMeiring NaudeNaude sta.sta. XX46,XX46, 1717 Jul 1984.5.1984. 5. Apertural andand laterallateral views of holotype.holotype. 6.6. ApicalApical andand 7.7. LateralLateral viewsviews ofof protoconch.protoconch. 8.8. DetailDetail ofof surface sculpture on body whorl.whorl, p, primaryprimary spiralspiral cords;cords; s,s, secondarysecondary spiralspiral cords;cords; t,t, spiralspiral threads.threads.

Genus Nipponaphera Habe, 19611961 of varix. Columella with twotwo prominent, sharply keeled columellar foldsfolds and aa Type species: Nipponaphera habei Petit, strongly reflected siphonal fold.fold. 1972 by I.C.Z.N. Opinion 1052.1052. Description.Description.-Shell—Shell (Fig. 11)11) small forfor genus, to 12.4 mm, ovately conical,conical, with Nipponaphera wallacei new species rounded anterior. Spire relatively shortshort Figs. 11-13, 16 (spire angle 70°), comprising lessless thanthan half Diagnosis.-ADiagnosis.—A small species with aa of shell length. Protoconch (Fig.(Fig. 13)13) of 1.671.67 heavy, strongly sculptured, narrowly um­um- smooth, inflated whorls. Transition toto teleo­teleo- bilicate shell. Aperture sharply triangular,triangular, conch marked by onset of spiral cords fol­fol- outer lip with strong teethteeth along innerinner edgeedge lowed immediately by strong axial ribs.ribs. Te-Te- ISO150 PROCEEDINGS OF THE BIOLOGICALBIOLOGICAL SOCIETYSOCIETY OFOF WASHINGTONWASHINGTON

Figs. 9-10. TrigonoslomaTrigonostoma kilbumikilbumi newnew species.species. DistalDistal endsends ofofradular radularteeth teethof ofholotype. holotype.9. 9.Lateral Lateralview. view. 10. End-on view. leoconch of up toto 44 angularangular whorls.whorls. SutureSuture Sep 1996. Paratype 2,2, NMNM V303,V303, 12.412.4 mm,mm, impressed behind slopingsloping shouldershoulder thatthat isis Off Park Rynie, Natal,Natal, SouthSouth Africa,Africa, inin 5353 delineated by peripheral keel.keel. AxialAxial sculp­sculp- m. Taken by SCUBA, livingliving onon thethe spirespire ofof ture of strong, broad, prosoclineprosocline ribsribs (10(10 onon the turbinid gastropod BolmaBolma andersoni.andersoni. 1919 penultimate whorl) thatthat becomebecome moremore wide­wide- May 1990.1990. ly spaced on body whorlwhorl wherewhere theythey appearappear Etymology.Etymology.-This—This speciesspecies isis namednamed forfor as varices with flaredflared edges.edges. SpiralSpiral sculp­sculp- Mr. Martin Wallace, whowho collectedcollected thethe typetype ture of broad, flattishflattish primaryprimary cords,cords, 33 sec­sec- material and generously mademade itit availableavailable ondary cords between adjacentadjacent primaryprimary for study. cords, with finefine threadsthreads betweenbetween somesome Comparative remarks.-remarks.—TheThe genusgenus Nip­Nip- cords. Aperture sharplysharply triangular,triangular, deflecteddeflected ponaphera isis distinguished fromfrom Trigonos­Trigonos- 0 from coiling axis byby 2020°.• SiphonalSiphonal canalcanal toma primarily byby thethe presence,presence, inin thethe latter,latter, short, well-defined. OuterOuter liplip reflectedreflected an­an- of an open umbilicus thatthat extendsextends backback toto teriorly, with spiralspiral sculpturesculpture visiblevisible the protoconch. Also, NipponapheraNipponaphera hashas aa through thin edge, 99 strongstrong teethteeth alongalong in­in- wide and flat siphonalsiphonal fold.fold. AlthoughAlthough thethe ner edge of varix, small,small, angularangular indenta­indenta- new species here describeddescribed hashas anan umbili­umbili- tion at shoulder. ParietalParietal regionregion short,short, col­col- cus, it isis not profound. ItIt differsdiffers fromfrom N.N. umella straight, withwith small,small, thinthin callus,callus, 22 paucicostata (Sowerby,(Sowerby, 1894)1894) ofof thethe Ara­Ara- sharply keeled columellar foldsfolds andand strong­strong- bian Sea area inin being umbilicate,umbilicate, inin havinghaving ly reflected siphonalsiphonal fold.fold. UmbilicusUmbilicus nar­nar- a more rounded shoulder,shoulder, andand moremore andand fin­fin- row, inconspicuous, borderedbordered byby well-de­well-de- er spiral cords. SpecimensSpecimens ofof N.N. paucicos­paucicos- veloped, cord-like siphonalsiphonal fasciole.fasciole. ShellShell tata figured by VerheckenVerhecken (1986,(1986, Figs.Figs. 4­4— color chestnut brown, withwith lighterlighter spiralspiral 6) lack varix-like axialaxial ribsribs onon thethe bodybody bands along shouldershoulder andand middlemiddle ofof bodybody whorl, but the typetype specimenspecimen hashas veryvery whorl. strong, unevenly spacedspaced ribsribs onon thethe bodybody Type locality.-Offlocality.—Off Phumula,Phumula, Natal,Natal, whorl. The Japanese N.JV. teramachiiteramachii (Habe,(Habe, South Africa, inin 4545 m, onon reef.reef. JunJun 1997.1997. 1961) has a sharpsharp keel,keel, isis umbilicate,umbilicate, withwith Type material.-Holotype,material.—Holotype, NMNM V4689,V4689, a cord-like siphonalsiphonal fasciole,fasciole, butbut lackslacks thethe 6.6 mm, from typetype locality.locality. ParatypeParatype 1,1, NMNM squarish, finely imbricatedimbricated spiralspiral sculpturesculpture V3889, Off Phumula, Natal,Natal, SouthSouth Africa,Africa, of this new species.species. MelvillMelvill andand StandenStanden in 35 m, taken byby SCUBA,SCUBA, livingliving onon shellshell (1901:451) reported N. paucicostatapaucicostata fromfrom of Bolma andersoni (E.(E. A.A. Smith,Smith, 1902).1902). 44 the Gulf of Aden "adhering"adhering toto thethe upperupper VOLUME 113.113, NUMBER 11 151

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VJr 1 mm

.*

Figs. 11-13. Nipponaphera wallaceiwallacei newnew species.species. Holotype.Holotype, NMNM V4689.V4689, OffOffPhumula. Phumula.Natal. Natal,South SouthAfrica. Africa, SCUBA in 45 m.m, on reef.reef. lunJun 1997.1997. 11.11. AperturalApertural viewview ofof holotype.holotype. 12.12. DetailDetailof ofsculpture sculptureon onbody bodywhorl. whorl.13. 13. Apical view of protoconch. part of Rapana bulbosa, 30-5030-50 fathoms."fathoms." teleoconch abrupt, markedmarked byby onsetonset ofof The alimentary systemsystem ofof cancellariidscancellariids isis weak, closely-spaced axialaxial ribsribs thatthat becomebecome adapted to feedfeed on body fluidsfluids ofof prey/hostprey/host stronger, more regularlyregularly spaced,spaced, whenwhen spi­spi- organisms (Petit(Petit & Harasewych, 1986;1986; Har­Har- ral cords firstfirst appear withinwithin 0.250.25 whorl.whorl. Te­Te- asewych & Petit, 1986).1986). TheThe associationassociation ofof leoleoconchconch of 2 rounded,rounded, stronglystrongly tabulatetabulate species of Nipponaphera withwith otherother gastro­gastro- whorls. Suture stronglystrongly impressed.impressed. AxialAxial pods suggests that thisthis groupgroup ofof cancellar­cancellar- sculpture of evenly spaced,spaced, well-defined,well-defined, iids may be specializedspecialized ectoparasitesectoparasites ofof axially aligned ribsribs (30(30 onon bodybody whorl)whorl) asas large gastropods. broad as intervening spaces.spaces. SpiralSpiral sculp­sculp- ture of sharply demarcated evenlyevenly spacedspaced Genus Zeadmete Finlay, 19261926 cords (2 between suturesuture andand shoulder,shoulder, 1717 below shoulder) diminishing inin strengthstrength Type species: Cancellaria traillitrailli Hutton,Hutton, abapically, forming smallsmall nodulesnodules asas theythey 1973 by original designation. cross axial ribs. Aperture elongated,elongated, nar­nar- rowly elliptical. Outer liplip faintlyfaintly sinuate,sinuate, Zeadmete verheckeni, new speciesspecies smooth within. ColumellaColumella withwith twotwo broad,broad, Figs. 14-18 weak, columellar foldsfolds (Fig.(Fig. 17,17, pcf,pcf, acf)acf) Diagnosis.Diagnosis.-A—A small speciesspecies withwith anan and siphonal fold (Fig.(Fig. 17,17, sf).sf). SiphonalSiphonal ca­ca- ovate shell. Shoulder tabulate.tabulate. SurfaceSurface nal small but distinct, axiallyaxially aligned,aligned, notnot sculpture dominated by spiralspiral cords.cords. Pseu­Pseu- forming siphonal fasciole.fasciole. AnteriorAnterior portionportion do-umbilicus narrow. OuterOuter liplip thin,thin, smoothsmooth of inductura bordered byby weakweak parietalparietal within. Buccal mass minute. RadulaRadula absent.absent. wash, partially covering narrownarrow pseudo­pseudo- Description.Description.-Shell—Shell (Fig.(Fig. 14)14) small,small, umbilicus. Shell color white.white. PeriostracumPeriostracum reaching 7.9 mm, thin,thin, withwith steppedstepped spire,spire, thin, finely lamellate,lamellate, straw-colored.straw-colored. rounded anterior. Protoconch (Figs.(Figs. 15-16)15-16) Preserved animal white, withwith short,short, nar­nar- erect, smooth, of 1.51.5 whorls.whorls. TransitionTransition toto row, posteriorly roundedrounded foot.foot. TentaclesTentacles tu-tu- 152 PROCEEDINGS OF THE BIOLOGICALBIOLOGICAL SOCIETYSOCIETY OFOF WASHINGTONWASHINGTON

Figs. 14-17. ZeadmeleZeadmete verheckenivcrheckeni newnew species.species. Holotype,Holotype, NMNM C6800,C6800. OffOffStony Stony Point,Point,Transkei, Transkei,South South Africa (32°37.5'S, 28°45.8'E),28°45.8'E), dredgeddredged inin 390-400390-400 m,m, muddymuddy sand,sand,small smallstones. stones.RN R/VMeiring MeiringNaude Naudesta. sta.V VII, 1I, 12 Jul 1984. 14.14. AperturalApertural andand dorsaldorsal viewsviews ofof holotype.holotype. 15.15. ApicalApicaland and 16.16.Lateral Lateralviews viewsof ofprotoconch. protoconch.17. 17. Columella. pcf, posteriorposterior columellarcolumellar fold;fold; acf,acf, anterioranteriorcolumellar columellarfold; fold;sf, sf,siphonal siphonalfold. fold.

bular, symmetrical, withwith veryvery largelarge blackblack thirds of cephalic haemocoel, thethe restrest oc­oc- eyes at their bases. PenisPenis long,long, narrow,narrow, dor­dor- cupied by long,long, convolutedconvoluted mid-esophagus.mid-esophagus. so-ventrally flattened,flattened, withwith smallsmall terminalterminal Proboscis strongly coiledcoiled withinwithin proboscisproboscis papilla. Osphradium veryvery broad,broad, stronglystrongly sheath. Extended proboscisproboscis likelylikely exceedsexceeds asymmetrical, dorsal leafletsleaflets twicetwice asas broadbroad shell length.length. Proboscis thin,thin, withwith minuteminute as ventral leaflets.leaflets. CtenidiumCtenidium lessless thanthan halfhalf buccal mass inin anteriormostanteriormost 0.1250.125 ofof pro­pro- as wide and twice asas longlong asas osphradium.osphradium. boscis. RadulaRadula absent.absent. Hypobranchial gland large,large, glandular.glandular. Re­Re- Type locality.-Offlocality.—Off StonyStony Point,Point, Trans­Trans- tracted proboscis occupiesoccupies anterioranterior two-two- kei, South Africa (32°37.5'S,(32°37.5'S, 28°45.8'E),28°45.8'E), VOLUME 113, NUMBER 11 153

28° S

32° S

36° S

015° E 020020°0 E 025025°0 E 030030°0 E 035035°0 E

Fig. 18. Geographic distributions of SouthSouth African Cancellariidae.Cancellariidae. AdmetulaAdmetula epulaepula PetitPetit && Harasewych,Harasewych, 1991, vertically hatched area = typetype locality;locality; openopen squaressquares == newnew records.records. AdmetulaAdmetula afraafra newnew species,species, starstar inin circle = type locality;locality; filled circlecircle = additionaladditional record.record. TrigonostomaTrigonostoma kilbumikilburni newnew species,species, blackblack starstar == typetype locality; open circle = additional record.record. Nipponaphera wallacei newnew species,species, triangletriangle == typetype locality.locality. ZeadmeteZeadmcte verheckeni new species, diamond = typetype locality.locality. dredged in 390-400 m, muddy sand, smallsmall Powell, 1933, from 50 fathoms off Snares stones. RJVR/V Meiring Naude sta. VII,VI1, 1212 JulJul Islands, New Zealand, has thethe samesame shellshell 1984. shape and columellar structure as Z.Z ver­ver- Type material.-Holotype,material.—Holotype, NM C6800, heckeni but has deeply cancellated sculp­sculp- 5.1 mm, from type locality. ture on the posterior half of the body whorl Etymology.Etymology.-Named—Named for Mr. Andre Ver­Ver- and only spiral cords on thethe anterior por­por- hecken, Mortsel, Belgium, inin recognition of tion. his contributions to the studystudy of thethe Can­Can- In the course of a study on Australian cellariidae. cancellariids, we found thatthat thethe abyssal Comparative remarks.-Placementremarks.—Placement of Zeadmete kulanda Garrard, ]1975975 has a rad­rad- this new species in the genus Zeadmete isis ula similar to that of Nothoadmete tumidatumida tentative. Zeadmete verheckeni isis concho­concho- (Oliver, 1982:figs. 3, 5). The factfact thatthat no logically most similar to a group of speciesspecies radula was found inin Z. verheckeni argues from off the southern coasts of New Zea­Zea- against these species being congeneric. land and Australia that have been assigned However, we are reluctant toto introduceintroduce an­an- to the genera Oamaruia Finlay, 19241924 andand other genus-level taxon until more data on Zeadmete Finlay, 1926. Powell (1979:224) the species with this shellshell formform are avail­avail- treated Zeadmete as a subgenus of Oama­Oama- able. The problems of generic placement of ruia. Several New Zealand and Australian small, deep-water cancellariids was briefly species presently assigned to Zeadmete discussed by Verhecken (1997:296). agree with this South African speciesspecies inin Among the South African taxa, this spe­spe- shell form and sculpture (see Garrard 1975,1975, cies most closely resembles "Cancellar­"Cancellar- Powell 1979). Zeadmete subanlarclicasubantarctica ia"eutrios Barnard, 1959, fromfrom which itit isis 154 PROCEEDINGS OF THE BIOLOGICAL SOCIETYSOCIETY OFOF WASHINGTONWASHINGTON easily distinguished by itsits tabulatetabulate shouldershoulder Melvill,Melvill. J. C, & R. Standen. 1901.1901. TheThe MolluscaMollusca ofof and lower spire. the Persian Gulf,Gulf. Gulf ofof Oman,Oman, andand ArabianArabian Sea, as evidenced mainly throughthrough thethe collec­collec- tions of Mr. FF. W. Townsend,Townsend, 1893-1900;1893-1900; withwith AcknowAcknowledgmentsledgments descriptions of new species.-Proceedingsspecies.—Proceedings ofof the Zoological Society ofof LondonLondon forfor 1901:1901: We thank Dr. Richard N. KilburnKilburn forfor 327-460, pIs.pis. 21-24. making these specimens available andand forfor Oliver, P. G. 1982.1982. A new speciesspecies ofof cancellariidcancellariid gas­gas- his patience in waiting forfor theirtheir description.description. tropod from Antarctica withwith aa descriptiondescription ofof thethe Correspondence and discussion with Mr.Mr. radula.-Britishradula.—British Antarctic SurveySurvey BulletinBulletin 57:57: Andre Verhecken of Mortsel, BelgiumBelgium add­add- 15-20. Petit, R. E., & M. G. Harasewych. 1986.1986. NewNew Philip­Philip- ed to our knowledge and was helpful inin pre­pre- pine Cancellariidae (Gastropoda:(Gastropoda: Cancellari­Canccllari- paring this paper. acea), with notes on thethe finefine structurestructure andand func­func- tion of the Nematoglossanematoglossan radula.-radula.—TheThe VeligerVeliger Literature Cited 28(4):436-443. Powell, A. W. B. 1979. New ZealandZealand Mollusca.Mollusca. Wil­Wil- Barnard, K. H. 1959. Contributions toto thethe knowledgeknowledge liam Collins, Auckland, xivxiv ++ 500500 pp.pp. of South African marine Mollusca.Mollusca. PartPart II.II. Gas­Gas- Richards, D.0. 1981. South African seashells.seashells. AA collec­collec- tropoda: Prosobranchiata: Rachiglossa.-An­Rachiglossa.—An- tor's guide. CC. Struik Publishers, CapeCape Town.Town. 9898 nals of the South African Museum 45:1-237.45:1-237. pp. + 60 pis. Gan'ard,Garrard, T. A. 1975.1975. A revisionrevision ofof AustralianAustralian Cancel­Cancel- Steyn, D. G., & M. Lussi. 1998.1998. MarineMarine shellsshells ofof SouthSouth lariidae (Gastropoda: Mollusca).-RecordsMollusca).—Records ofof Africa. Ekogilde Publishers,Publishers. Hartebeespoort.Hartebeespoort. the Australian Museum 30(30(1):I): 1-62.1-62. 264 pp. Harasewych, M. GG.,.. & R. E.E. Petit.Petit. 1986.1986. NotesNotes onon thethe Verhecken, A. 1986.1986. A revisionrevision ofof thethe CancellariidaeCancellariidae morphology of Admele viridula (Gastropoda:(Gastropoda: ((Neogastropoda:eogastropoda: Cancellariacea) ofof thethe RedRed SeaSea Cancellariidae).-TheCancellariidae).—The Nautilus 100(3):85-91.100(3):85-9I. and the Gulf of Aden.-GloriaAden.—Gloria MarisMarts 25(4):25(4): I.C.Z.N.I.CZ.N. 1976. Opinion 1052.1052. NipponapheraNipponaphem Habe,Habe, 133-153. 1961 (Gastropoda): Designation ofof aa type-spe­type-spe- Verhecken, A. 1997.1997. Mollusca Gastropoda: ArafuraArafura cies under thethe plenary powers.-Bulletinpowers.—Bulletin ofof Sea Cancellariidae collectedcollected duringduring thethe KA­KA- Zoological Nomenclature 32(4):242-243. RUBAR Cruise. Pp. 295-323295-323 inin A CrosnierCrosnier && Kensley, B. 1973.1973. Sea-Shells ofof southernsouthern Africa,Africa, Gas­Gas- P.R Bouchet,Bouchet. eds.,eds.. Resultats desdes CampagnesCampagnes MU­MU- tropods. MaskewMaskcw Miller Ltd., CapeCape Town,Town, 236236 SORSTOM, vol. 16.16. Memoirs dudu MuseumMuseum Na­Na- pp. tional d'Histoire NaturelleNature!!e 172.172.