BASTERIA, 62: 25-34, 1998 The systematic position ofTritonidea dentataSchepman, 1911 (Gastropoda, Prosobranchia: Buccinidae) Geerat+J. Vermeij DepartmentofGeologyandCenter for PopulationBiology University ofCalifornia atDavis OneShields Avenue Davis, CA 95616 [email protected] Tritonidea dentataSchepman, 1911,ischaracterized bythepresence ofabasal labral tooth and by a deeplyrecessed ridgebearingelongatedenticles on theadaxial (inner) side oftheouter lip.This central Indo-West Pacificspeciesis hererecognizedasa RecentmemberofPreangeria Martin, 1921, a genus previouslyknown only fromthe Miocene ofsoutheast Asia. Taurasia Bellardi, 1882 (Muricidae:Rapaninae),in which speciesof Preangeria hadbeen included by several earlier authors,differsfromPreangeriabylackingalabral tooth andby havingalateral rather thanterminalopercularnucleus. Comparisonswithother generasuggestthat Preangeria belongs tothebuccinid subfamilyPisaniinae. Keywords: Gastropoda,Prosobranchia, Buccinidae,Pisaniinae, Tritonidea,Preangeria, Taurasia, IndianOceanJ,ava,Miocene. INTRODUCTION Many neogastropods have a shell in which the outer lip bears a blunt or sharp, downwardly projecting tooth on its abapical part. This so-called labral tooth has evolved many times independently. In an effort to understandwhen, where, and in which clades the evolution oflabral teeth has taken place, I have been conducting studies of the taxonomy, relationships, and distributionofliving and fossil species of gastropod in which alabral tooth is present. Here I report on the previously unsus- pected relationship between two taxa with a well-developed labral tooth, the Recent Tritonideadentata Schepman, 1911, and the Miocene species from Indonesiathat have variously been treated as members ofthe genera Acanthina, Preangeria, Acantinella, and Taurasia.I shallargue that this groupcomprises a single, distinctive, early Miocene to Recent clade, for which the generic name Preangeria Martin, 1921, is available. MATERIALS AND METHODS This study was based on my examination of Tritonidea dentata Schepman, 1911; Acanthinajavana Martin, 1899; andPreangeria angsananaMartin, 1921. In addition, Ihave examined (table 1) species of Buccinulum, Cymia, Eosipho, Janiopsis, and Pisania, genera which areeitherrelatedto thethree species namedabove or whichwerelinkedto these species by previous authors. 26 BASTERJA, Vol. 62, Mo. 1-2, 1998 AcanthinajavanaMartin, 1899: RGM9755,Selacau,Java Buccinuluml. lineum(Martyn, 1784):Leigh,New Zealand (GJV) B.vittatum littorinoides (Reeve,1846):Dunedin,New Zealand (GJV) Cymia tecta (Wood, 1828):Paitilla,Panama (GJV) EosiphosmithiSchepman, 1911:Philippines(MNHN) Janiopsisangulosa(Brocchi, 1814):Montegibbio,Italy(MNHN); Orciano,Italy(IRSNB) J.maxillosa (BonelliinBellardi&Michelotti, 1840):Lapugy,Rumania(NHMW) J.parisiensis(Deshayes, 1835):Chaussy, France (MNHN, GJV) Pisaniapusio(Linnaeus,1758):FernandodeNoronha,Brazil(GJV');GardenKey,FloridaKeys, Florida(USNM 890264) P. striata(Gmelin, 1791):Mediterranean coast,Israel (twolots, GJV) PreangeriaangsananaMartin, 1921: CiAngsana,Java(RGM9753) Taurasiapleurotoma(Grateloup, 1832):Saint-Paul-les-Dax,France,andLeognan,France(MNHN) T.striata(Quoy&Gaimard, 1833):Puerto Galera,Philippines(GJV) Tritonidea dentataSchepman,1911: KuandangBay, Indonesia;ZMA MOLL3.11.032;Philippines(MNHN) Table 1.Materialexamined. Abbreviations: ANSP — Academy ofNatural Sciences, Philadelphia; GJV — Ver- meij collection; IRSNB — InstitutRoyal desSciences Naturelles de Belgique, Brussels; MNHN — Museum National d'Histoire Naturelle, Paris; NHMW — Naturhisto- risches Museum, Wien; RGM — Nationaal Natuurhistorisch Museum, Leiden (for- merly Rijksmuseum voor Geologie en Mineralogie); USNM — United StatesNational Museum ofNatural History, Washington; ZMA — Zoologisch Museum, Amsterdam. SYSTEMATICS Family Buccinidae Rafinesque, 1815 Subfamily Pisaniinae Gray, 1857 Genus Preangeria Martin, 1921 Type species. — Designation by Wenz (1943, p. 1356), Preangeria angsanana Martin, 1921. Synonym. — Acantinella Shuto, 1969 (p. 109); type species by original designation Acanthinajavana Martin, 1899. Revised description. — Shell up to 31 mm high, elongate-ovate, consisting of a protoconch of about two smooth whorls and of a teleoconch of four to six whorls separated by an impressed suture; last whorl ofteleoconch roundedor weakly shoul- deredabove, little or not constricted below; spire moderately high, comprising 35 to 44%oftotalshell height; spiral sculpture consisting ofwell-definedcords; axial sculp- turepresentas ribletsonearlywhorls,oftenobsolete onlastoneortwo whorls;aperture elongate-ovate; outerlip indeterminate, its adaxial sidebearing a deeply recessed ridge with nine to twelve elongate denticles; abapical end of outer lip bearing sharp labral tooth, situated at apertural end ofexternal groove; columellawith one ortwo central folds and up to four superficial abapical riblets; adapical end ofinner lip with weak or noparietal tooth; outer lip without adapical notch; siphonal fasciole present; siph- onal canal short, broadly open; periostracum thick, hairy. Vermeij: Systematic position of Tritonidea dentata 27 Includedspecies. — P. angsananaMartin, 1921; P. dentata(Schepman, 1911); P. javana (Martin, 1899); P. sundaica (Oostingh, 1935). Geographical and stratigraphic distribution. — Early Miocene to Recent, central Indo-West Pacific. Preangeria dentata (Schepman, 1911), fig. 1 Tritonidea dentata Schepman,1911: 303,pi. 19,fig. 8. AxymenephilippinensisPetuch, 1979:8, figs. 14-15. Eosiphodentatus(Schepman, 1911):Bouchet &Waren, 1986:470,figs. 17,58-61 Revised description of shell. — Shell ovate, consisting of one and a half to two protoconch whorls and five teleoconch whorls; maximumheight 30.0 mm; last whorl comprising 56 to 62% oftotal shell height; suture deeply impressed; last whorlweakly shouldered above periphery, weakly constricted; spiral sculpture consisting often to fifteen cords on last whorl, and five cords on penultimate whorl; secondary spiral sculpture absent; axial sculpture consisting oftwenty-one to twenty-five very fine riblets on penultimate whorl, absent on last whorl; basal external groove, situatedbelow all other external shellsculpture, ends in sharp, abapically directedtoothat edge ofouter lip; thislabraltooth separated from adapical sector ofouter lip by shallow sinus; outer lip weakly convex, crenulatedatedge; adapical sinus onouterlip very weakor absent; adaxialsideofouterlip with deeplyrecessed ridge, onwhich are situatednineto twelve denticles; columellastraight, bearing one or two central folds; parietal rib at adapical end ofinner lip absent; innerlip adherent; lowsiphonal fasciole present; siphonal canal short, broadly open;umbilicusabsent; aperture elongate-ovate, height:breadth ratio 2.4 to 2.8. Discussion. Previous assignments of Tritonideadentata..— Schepman (1911: 303, pi. 19 fig. 8) describedandillustrated Tritonideadentataonthe basis ofa very fresh,empty shell from a depth of72 m in the Bay ofKuandang in the CelebesSea in Indonesia. Since its original description, the species has been found at various localities in the Celebes and Sulu Seas and nearNew Caledonia at depths of72 to 700 m. Authors ofthethreeprevious discussionsofT. dentatacompared thespecies only to otherRecent forms, and assigned it to three generain two families. Placement in the Buccinidae, broadly defined, is amplyjustified on thebasis ofthe radula (Bouchet & Waren, 1986). According toBouchet&Waren (1986), theradulahasasquarecentraltooth withlarger outer and smallerinner cusp. This radular type resembles that in thebuccinid genera Eosipho Thiele, 1929, and Manaria Smith, 1906, both of which contain generalized, deep-water, tropical buccinids withmanyplesiomorphic features(see Bouchet&Waren, 1986; Harasewych, 1990). Withconsiderablehesitation, Schepman (1911) assigned his new species to Tritonidea Swainson, 1840, an objective junior synonym of Pollia Gray in Sowerby, 1834 (type species: Buccinum undosum Linnaeus, 1758). As restricted by Vermeij & Bouchet (in preparation), this early Miocene to Recent pisaniine buccinid genus is characterized by a fusiform, weakly basally constricted shell with an externally thickened, adapically extended, terminalvarix, a shallowadapical sinus on theouter lip,and aparietal tooth at the adapical end ofthe innerlip. All species have roundedaxial ribs onat leastthe early teleoconchwhorls, although in the type species and in several other membersof thegenusthe ribs are reducedorlost onthe lasttwo or morewhorlsofthe adult shell. Most individualsofmost species have the central crenulationon the sharp edge ofthe 28 BASTERIA, Vol. 62, No. 1-2, 1998 Fig. 1.Preangeriadentata(Schepman,1911),Panglao,Bohol,Philippines; height,22.2mm.A,outerlipwithlabral spine;B,aperturalview;C,dorsalview. outer lip enlarged and slightly ventrally projecting beyond the othercrenulations.This labraltooth liesnearthemiddleoftheouter lip.In the related genus CantharusRoding, 1798 (type species: Buccinum tranquebaricum Gmelin, 1791), the labral tooth is usually better developed andis situatedon the lower thirdofthe outer-lip margin. Compared to Pollia, Cantharus has a relatively broaderaperture and a less well-developed parietal tooth. The axial sculpture continues to the last whorl, and the outer lip of Cantharus remains thin and therefore does not become a terminal, externally roundly thickened, adapically extended varix asit doesin Pollia. Bothgeneraare characterizedby a thick, hairy periostracum. Tritonideadentatadiffers from Pollia and Cantharus by having a deeply recessed ridge on the adaxial side ofthe outer lip, a large labral tooth situated at the abapical end oftheouterlipbelowallelementsofexternalsculpture insteadofonits lower ormiddle third, andby lacking anadapical sinus onthe outer lip, central folds onthe columella, Vermeij: Systematic position of Tritonidea dentata 29 anda parietal ribon theadapical end ofthe innerlip. The outerlip is notexternally thickened or adapically extended as it is in Pollia. Bouchet& Waren(1986) assigned Tritonideadentatawith somedoubtto Eosipho Thiele, 1929 [type species: Chiysodomus (Sipho) smithi Schepman, 1911]. They cited similarities in theradulato other species ofEosipho, andpointed outthat T. dentatahas many shell features in common with Cantharus aldermenensis Powell, 1971, and Eosipho thorybopus Bouchet & Waren, 1986, species without a labraltooth that Bouchet& Waren (1986) likewise assigned to Eosipho. E.smithi, the type ofEosipho, resembles T. dentatain having little or no axialsculpture, in the absence ofa parietal rib or tooth, and in lacking an adapical notch or sinus on theouterlip. It differs, however, in having a smoothinstead ofplicate columella, and in lacking a labraltoothand adeeply recessed, tooth-bearing ridge onthe adaxial side ofthe outer lip. Cantharus aldermenensisand Eosipho thorybopus have a thin outer lip with lirae (spiral ridges) on its adaxial side, a smooth columella, and axial sculpture on the spire whorls(Bouchet & Waren, 1986). Like E.smithi, these species lack a parietal tooth. Shell characters including the adaxially thickened outer lip, plicate columella, and labral tooth set Tritonidea dentata apart from other species assigned by Bouchet & Waren (1986) to Eosipho. MuricidaffinitiesofTritonideadentatawere postulated byPetuch (1979),who described thisspecies underthenameAxymenephilippinensis. Finlay (1927, p. 424) proposed Axymene for a New Zealandtrophonine muricid(Axymene turbatorFinlay, 1927, = Trophon auck- landicusSmith, 1902) inwhichtheaxialsculpture isobsolete. Powell(1979) synonymized Axymene with Xymene Iredale, 1915 (type species: Fusus plebejus Hutton, 1873). Like Tritonidea dentata, species of Xymene have a deeply recessed, denticulate ridge on the adaxial side ofthe planar outer lip. This feature occurs commonly among muricids, especially among members ofthe Ocenebrinaeand some Rapaninae (see Vermeij & Kool, 1994;Vermeij, 1995; Vermeij & Carlson, inreview). Xymene and other trophonine muricids, however, lack the periostracum and thelabraltoothpresentin Tritonideadentata. Assignment ofTritonideadentatato Preangeria. — Authorswho treated Tritonideadentata were evidently unaware ofseveral closely similar fossil species fromIndonesiaand the Philippines that wererevised by Beets (1984). These species, which havebeen assigned variously to genera in the Muricidae and Cancellariidae, were reassigned by Beets (1984) to Taurasia Bellardi, 1882(type species: Purpura subfusiformis d'Orbigny, 1852, = Purpura pleurotoma Grateloup, 1832; see Cossmann & Peyrot, 1924). Beets placed this genus in the muricid subfamily Drupinae (= Rapaninae in the sense ofKool, 1993). Martin(1899: 137, pi. 21 fig. 315) describedand illustrated thefirst ofthese species as Acanthinajavana. Shuto(1969: 109)later madethisspecies the type ofhis new genus Acantinella.Martin's species is remarkably like Tritonidea dentatain shell characters. The holotype (RGM 9755) from the Selacaubeds (late Miocene) ofjawa is like T. dentata in having an ovate outline withoutbasal constriction, one centraland four abapical columellarfolds, fine axial riblets onearly whorls, finecords persisting to the aperture, a long labraltooth at the abapical end ofthe outerlip, a deeply recessed ridge bearing ten lira-like denticlesonthe adaxialsideofthe outerlip, and adeeply impressed suture, and by lacking an adapical notch on the outer lip. It differs from T. dentata by being slightly higher-spired (last whorl comprising 65% instead of 58-62% of total shell height),by having the cords more numerousand ofalternating insteadofuniformsize, a slighdy narrower aperture (height-to-breadth ratio 3.0 insteadof2.4 to 2.8), andby having avery weakparietal tooth. Theheight ofthe holotype (30.4 mm)is very similar to that ofthe largest T. dentata (30.0 mm). 30 BASTERIA, Vol. 62, No. 1-2, 1998 Beets (1984) recognized thatthree other species are very closely related to Acanthina javana. Two ofthese, Preangeria angsananaMartin, 1921,and P. talahabensisMartin, 1921, were described as membersof Preangeria Martin, 1921, for which Wenz (1943: 1356) designated P. angsanana as type species. Martin(1921: 450) originally proposed Prean- geria as a member ofthe Cancellariidaebecause ofthe presence of columellarfolds. Recognizing thatsuch folds also occur in various muricid groups, Martin (1928: 124) latertransferredPreangeria to the Purpuridae (= Rapaninae in the sense ofKool, 1993). He did not realize that these species possess a labral tooth, and therefore did not perceive theconnectionbetween themandAcanthinajavana. Thetrajectory ofthegrowth lines near the base ofthe shell led Beets (1984) to infer the presence ofa tooth in Preangeria and to link Acanthinajavana with the two species ofPreangeria. Healso placed Nucella (Acanthinucella) sundaica Oostingh, 1935, from the Bojongmanik beds (Preange- rian, late Miocene) ofJava in this group, and assigned all four species, together with several others without a labral tooth, to Taurasia. The type species of Taurasia (Purpura subfusiformis d'Orbigny, 1852) is a Miocenefossil considered by Cossmann & Peyrot (1924) to represent a relatively weakly sculptured form ofthe variable early Miocene (Aquitanian and Burdigalian) southern European species Purpura pleurotoma Grateloup, 1832.This species is characterizedby anelongate- ovate, basally very weakly constricted shellwhose planar, crenulated outerlip bears a deeply recessed axial ridge adorned with six to ten denticles that continue into the aperture as lirae.A distinct parietal toothand one or two central columellarfolds are present onthe inner lip. Spiral sculpture consists ofseveral sizes ofcords, which are scaly by virtue oflamelloseincrementallines. Axial sculpture is variably expressed. In some forms of Taurasiapleurotoma, they may form a row ofprominent nodes, whereas in othersthe axial ribs are low. In the Recent fauna, Taurasia isrepresented by a single species, Purpura striataQuoy & Gaimard, 1833(betterknown as Purpura buccinea Deshay- es, 1844;see Houart, 1996), distributedfrom the SolomonIslands to New Guinea, the Philippines, and eastern Indonesia. In this species, axial sculpture consists ofeight broadly rounded ribs that on the last whorl become obsolete toward the base. The abapical sector oftheouter lip is slightly concavewhen viewedfrom theapertural side. The adapical sector oftheouter lip forms ashallowsubsutural sinus. The lipjoins the penultimate whorl at a low angle, and with the parietal tooth at the adapical end of the inner lip forms a narrow, very short adapical apertural channel. A periostracum is absent. T. niasensis Beets, 1984,from the Plio-Pleistocene ofNias (west ofSumatra) is a somewhat morestrongly sculptured species thatis evidently very close to T. striata. Although Taurasia as circumscribed above is morphologically very similar to the labral-tooth-bearing group, the available evidence leads me to regard the two groups as representing two taxonomically distantgenera. For the groupwith a labral tooth, the earliest available name is Preangeria Martin, 1921, ofwhich AcantinellaShuto, 1969, is ajuniorsubjective synonym. It differs from Taurasiaby thepresenceofa labraltooth, by having the axial sculpture more or less confinedto the spire whorls, by lacking the adapical apertural channel, by the absence ofsquamosegrowth increments, and (in the living species) by the presence ofa well-developed periostracum. Beets(1984) discussed two additionalspecies from southeast Asia asprobable mem- bers of Taurasia. These are T. pendopoensis Beets, 1984, from beds of probable late Miocene(Preangerian) age at Palembang, Sumatra, and Tritonideapraeundosa Vreden- burg, 1923,from theKama beds(Miocene) ofBurma. These species, whichmay prove to be synonyms (Beets, 1984), apparently lack axialsculpture as well as a labraltooth. The presence ofcolumellarfolds in T.pendopoensis, together with other features ofthe Vermeij: Systematic position of Tritonidea dentata 31 shell, implies that this species, and the less well-preserved T. praeundosa, are related to Preangeria. Thetwo species maybelong to Eosipho inthe broadsenseasusedby Bouchet and Waren (1986). Evolutionary trends in Preangeria. — Comparisons among thefive species ofPreangeria reveala consistent evolutionary trend toward a reduction in axial sculpture. The two earliest species, P. angsananaand P. talahabensis (both from the early Miocene Nyalin- dungbeds ofJawa), have axial ribs persisting to the aperture ofthe last whorl. In the late Miocene P.javana fromJava and the Philippines, and the less slender P. sundaica, axialsculpture occurs only on the spirewhorls. The Recent P. dentatausually lacks axial sculpture altogether, or has it expressed as fine riblets on the spire whorls. P. dentata also shows a reduction in the parietal tooth and a broaderaperture relative to the earlierspecies. These trends are possibly related to an increasing depth ofhabitat of the species of Preangeria over time. At the depths of72 to 700 m in which the Recent P. dentatalives, predation and thereforeselection in favourofantipredatory reinforcing sculpture and a narrow aperture may be less intense than in the shallower-water habitats inferred for the Miocenerepresentatives ofthe genus. Taxonomic position of Preangeria. — As indicatedabove, evidence from the radula implies that Preangeria belongs to the Buccinidae, broadly defined. Shell characters are notinconsistentwiththisassignment. SeveralshellcharactersofPreangeria, however, are somewhat unusual formembersofthislarge group.BelowI review briefly some other generawith features in common with Preangeria, and make some tentative suggestions about the position of Preangeria within the Buccinidae. Preangeria has several shell characters in common with some membersofa group of generaincluding Buccinulum Deshayes, 1830 (early Miocene to Recent, New Zealand and Australia; see Ponder, 1971); Euthria J. E. Gray in M. E. Gray, 1850 (middle Eocene to Recent, Europe); Samudra Beets, 1986 (middle Eocene to middle Miocene, Indonesia); and Siphonofusus Kuroda & Habe, 1952 (Oligocene to Recent, Europe and Indo-WestPacific). The charactersand systematics of thesegenerahavebeen reviewed by Ponder (1971), Shuto (1978), and Beets (1986). Many members of this group, especially species of Buccinulum and Euthria, are characterized by having the axial sculpture confinedto the early teleoconch whorls, by the absence ofan abapical sinus on the outerlip, and by having a deeply recessed ridge bearing elongate denticles or lirae on the adaxial side ofthe outer lip. The generaofthe Buccinulumgroup,which have been assigned to the Buccinulinae or Pisaniinae (Powell, 1951; Cernohorsky, 1971), differ from Preangeria by lacking the central columellarfold and labraltooth. Representatives of several other genera of buccinids also have a deeply recessed adaxial ridge on the outer lip. These include Pisania Bivona-Bernardi, 1832 (Middle Eocene to Recent, Europe; Recent, Indo-West Pacific and western Atlantic); Janiopsis Rovereto, 1899 (Middle Eocene to ?Recent, Europe and Indo-West Pacific); and AustriumphisVermeij, 1997(Pliocene, South Africa).Axialsculpture isreducedorabsent on the adult teleoconch whorls of Pisaniaand Austriumphis. An abapical sinus onthe outer lip is lacking in Pisaniaand Janiopsis but is present in Austriumphis. Pisania and Janiopsis are usually consideredto bemembersofthebuccinid subfamily Pisaniinae(see e.g. Peyrot, 1927; Cernohorsky, 1971; Givens & Kennedy, 1976), whereas Austriumphis was tentatively assigned to the Photinaebecause ofthe presence ofan abapical sinus (Vermeij, 1997).The only genusofthis groupwith oneortwo centralcolumellarfolds as seen inPreangeria is Janiopsis. Shells ofJaniopsis differfromPreangeria by having strong, 32 BASTERIA, Vol. 62, No. 1-2, 1998 persistent axialsculpture, andin charactersoftheouterlip. Thetypespecies, J.angulosa (Brocchi, 1814) from the middle Miocene (Langhian) to the Pliocene of southern Europe, has a distinct, blunt, labral tooth situated at the end ofa groovejust below the midpoint ofthe convex, externally thickened, and adapically extended terminal varix. Otherspecies ofJaniopsis I haveexamined, including the middle Eocene (Lute- tian) J. parisiensis (Deshayes, 1835) from the Paris Basin of France and the middle Miocene (Langhian) J. maxillosa (Bonelli in Bellardi& Michelotti, 1840) from the ViennaBasin ofAustria, lack a labraltooth, as do species of Pisaniaand Austriumphis. In the lightofthis discussion, I tentatively assign Preangeria to the subfamily Pisani- inae, broadly defined.Thisassignment mayhave to be revised as anatomicalandother data become available for Preangeria, and as phylogenetic relationships within the Buccinidae as a whole become better understood. Taxonomic position of Taurasia. — In view ofthe striking similaritiesin shell form between Taurasia and Preangeria, the question of where to assign Taurasia is worth considering. Bellardi(1882, p. 194) introduced Taurasia as a genus in the Purpuridae (= Rapaninae in thesense ofKool, 1993). This assignment was followedby most later authors (Cossmann, 1901; Cossmann & Peyrot, 1924; Wenz, 1938-1944; Beets, 1984; Vermeij & Kool, 1994). Placement near Pisania in the buccinid subfamily Pisaniinae was advocated by Fischer (1884). Unfortunately, nothing is known about the anatomy or radulaofthe living species, T. striata. Shell characters, though not definitive, point toward an assignment in the Rapaninae. These includethe more orless planar, crenate outer lip,small but distinct adapical apertuxal channel, squamose growth increments, and absence ofa periostra- cum (see Vermeij & Kool, 1994; Vermeij, 1995, for characterization ofRapaninae). Atmy request, G. Rosenberg has examinedthe operculum of T. striata(ANSP 206569, under the name Cronia buccinea), which he reports has a lateralnucleus. Opercula with a lateralnucleus are the rulein the Rapaninae, whereas weakly spiralopercula or those with a terminal nucleus are typical ofBuccinidae (Powell, 1951; Bouchet & Waren, 1985; Ponder& Lindberg, 1997). Only Buccinum Linnaeus, 1758,and its allies in the Buccininae have aconcentric operculum. Taurasia therefore appears to belong to the muricid subfamily Rapaninae. The presence ofone oftwo central columellarfolds in Taurasiacould imply a rela- tionship between Taurasia andsuchotherrapanine generaas AcanthaisVermeij &Kool, 1994(Recent, eastern Pacific); CymiaMorch, 1860(Oligocene to Recent, eastern Pacific; Oligocene to Miocene, Europe; Miocene to Pliocene, western Atlantic); Habromorula Houart, 1994 (Recent, Indo-West Pacific); Morula Schumacher, 1817 (Miocene to Recent, Indo-WestPacific);Drupa Roding, 1798(Recent, Indo-WestPacific); andRicinella Schumacher, 1817 (Recent, Indo-West Pacific), and Thalessa H. & A. Adams, 1853 (Recent, Indo-West Pacific). Cladistic analysis based onshell characters (Vermeij & Carlson, in review) places TaurasianearMorulaand several other generathatcomprise the so-called ergalataxine subclade. The conspicuous lirae on the adaxial side ofthe outer lip of Taurasia and the presence ofa shallow subsutural sinus on the outer lip indicate a relationship with Cymia, which differsfrom Taurasiamainly by having adeep adapical outer-lip notch instead of a shallow sinus. Taurasia may therefore represent a relatively early lineage in the ergalataxine subclade, which couldhave diverged from a Cymia-like ancestor. In any case, the similarities between Taurasia and Preangeria in shell form appear to result from convergence rather than from shared ancestry. Vermeij: Systematic position of Tritonidea dentata 33 REFERENCES BEETS,C., 1984. CommentsonAcantinella,PreangeriaandTaurasia(Muricidae,Drupinae).—ScriptaGeol. 74: 39-47. —,1986.Notes on Buccinulum(Gastropoda,Buccinidae),areappraisal.— Scripta Geol. 82:83-100. 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