VOL. 14 (1) IS5N 0/73-5251 Périodique trimestriel Bureau de dépôt * Société Belge de Malacologie association sans but lucratif 20 AVRIL 1999 SOMMAIRE New information on the malacological fauna (Mollusca, Gastropoda) of the Cape Verde Archipelago, with the description of five new species A new species of Fulgoraria (Gastropoda: Volutidae) from the East China Sea Description de Vexillum (Costellaria) alvinobalani n.sp. des Philippines (Gastropoda: Muricoidea: Costellariidae) A new species of Livonia (Gastropoda: Volutidae) from Northwest Australia APEX Société Belge de Malacologie a.s.b.1. Editeur responsable R. Duchamps Av. Mozart, 52, B-1190 Bruxelles Comité d'édition: Dr. T. Backeljau Koninklijk Belgisch Instituut voor Natuurwetenschappen Dr. Y. Finet Muséum d'Histoire Naturelle, Genève M. R. Houart Institut royal des Sciences naturelles de Belgique (collab. scient.) Dr. CI. Massin Institut royal des Sciences naturelles de Belgique Prof B. Tursch Université Libre de Bruxelles Dr. J. Van Goethem Koninklik Instituut voor Natuurwetenschappen Prof. G. Vauquelin Vrie Universiteit Brussel COTISATIONS MEMBERSHIP Belgique - Belgium Etranger - Foreign Membres résidant en Belgique Abonnement aux revues APEX & ARION (avec le service des bulletins) Subscription to APEX & ARION Membre effectif 53.20 1200 BEF Membre étudiant .…......................…. 600 BEF Europe (CEE=EE0 7 1600 BEF Autres pays (other countries)... 1800 BEF (sans le service des bulletins) Versement à effectuer par mandat poste Personne appartenant à la famille international ou par chèque bancaire à un des d'un membre effectif et ayant comptes mentionnés ci-dessous, en francs même résidence; tee 400 BEF belges uniquement. Payable, by international money order, or by bank check at one of the below mentioned Versement à effectuer à un des comptes accounts, in Belgian Francs only mentionnés ci-dessous, au nom de la Société Belge de Malacologie c/o Mme A. Langjeit, au nom de: Av. Cicéron, 27, bte 92, B-1140 Bruxelles at name of: Mme A. Langleit Av. Cicéron, 27, bte 92, B-1140 Brussels, Belgium Comptes bancaires CCP 000-0974225-54 ou BBL 310-0770258-67 Les articles et textes présentés dans cette revue réflètent l'opinion personnelle de leur(s) auteur(s) et non pas nécessairement celle de la Société ou de l'éditeur responsable. Tous droits de reproduction, de traduction et d'adaptation des articles publiés dans ce bulletin, réservés pour tous pays. All rights of reproduction are reserved without the written permission of the board. Internet: http://www.arkham.be E-mail: cvilvens@prov-liege-be ROLAN & RUBIO Cape Verde Fauna and five new species New information on the malacological fauna (Mollusca, Gastropoda) of the Cape Verde Archipelago, with the description of five new species E. ROLAN ! & F. RUBIO ? | Cänovas del Castillo 22, 36202 Vigo, Spain ? Depart. Zoologia, Facultad de Biologia, Burjassot, Valencia, Spain KEY WORDS. Parviturbo, Eatonina, Obtusella, Megalomphalus, Tomura, Cape Verde archipelago, new species. Vitrinella, Mareleptopoma, ABSTRACT. New information on the Cape Verde molluscan fauna is reported in the present work. Additional information for some previously known species is presented, and the radulae of Parviturbo insularis and Eatonina martae are illustrated. Five species new for science are described, which are included in the following genera: Obtusella lata, Megalomphalus serus, Vitrinella politurae, Mareleptopoma verdensis and Tomura abscondita. RÉSUMÉ. De nouvelles données sur la faune malacologique de l'Archipel du Cap Vert sont signalées. Des informations complémentaires pour des espèces connues sont présentées et les radulas de Parviturbo insularis et Eatonina martae sont illustrées. Cinq nouvelles espèces sont décrites:Obtusella lata, Megalomphalus serus, Vitrinella politurae, Mareleptopoma verdensis et Tomura abscondita. RESUMEN. En el presente trabajo se aporta nueva informaciôn a la fauna de moluscos del archipiélago de Cabo Verde. Por un lado, se mencionan datos adicionales para algunas especies ya conocidas, como las râdulas de Parviturbo insularis y Eatonina martae. Por otra parte, se describen cinco especies nuevas para la ciencia, que estän incluidas en los géneros que se mencionan a continuaciôn: Obtusella lata, Megalomphalus serus, Vitrinella politurae, Mareleptopoma verdensis y APEX 14(1): 1-10, 20 avril 1999 Tomura abscondita. INTRODUCTION The Cape Verde archipelago is inhabited by a very peculiar molluse fauna with a high number of endemic species which even today remain poorly known (ROLAN, 1992a). In previous molluscan inventaries written by BURNAY & MONTEIRO (1977), SAUNDERS (1978), and VON COSEL (1982a, 1982b, 1982c) there is little information on the small gastropods. Recently, small species have been published from Cape Verde Islands, most of them endemic to this archipelago (FERNANDES & ROLAN, 1988, ROLAN, 1988, 1991, 1992b, MOOLENBEEK & ROLAN, 1988, ROLÂAN & FERNANDES, 1989, BURNAY & ROLAN, 1990, ROLAN & RUBIO, 1992, ROLAN & TEMPLADO, 1993, TEMPLADO & ROLAN, 1994, PENAS & ROLAN, 1997a, 1997b, 1998 and HOENSELAAR & GOUD, 1998). Further researchs are actually in progress (PENAS & ROLAN, in preparation, and ROLAN & LUQUE, in press) but curiously no species in the genera Obtusella, Megalomphalus, Vitrinella, Mareleptopoma or Tomura have been mentioned from these islands. Nevertheless, in the study of sediment material collected from several trips between 1978 and 1988, as well as in the expedition Macaronesia 2 in 1997, some small shells new for science were found and are described in this present work. At the same time live material of other species has been collected which allows us to add new information concerning living animal and anatomy. Abbreviations AMNH: American Museum of Natural History, New York. MNCN: Museo Madrid. MNHN: Muséum National d'Histoire Naturelle, Paris. NNM: Nationaal Natuurhistorisch Museum, Leiden. USNM: The National History Museum, Washington. CFR: collection Federico Rubio, Valencia CER: collection Emilio Rolän, Vigo. Nacional de Ciencias Naturales, APEX 14(1): 1-10, 20 avril 1999 RESULTS Subclass ARCHAEOGASTROPODA Superfamily TROCHOIDEA Family SKENEIDAE Clark, 1851 Genus Parviturbo Pilsbry & MecGinty, 1945 Type species, by original designation: Parviturbo rehderi Pilsbry & McGinty, 1945. Florida. Parviturbo insularis Rolän, 1988 Figs. 1-4 Material examined. See ROLAN (1988). More recent material: 4 specimens, in Rabo de Junco, Sal, intertidal; many empty shells from other places (Boa Vista: Morro de Areia, Porto da Cruz; Santiago: Prainha, Ponta Geneanes, Tarrafal; Brava: Porto do Anciäo, Furna, etc.). Description. See ROLAN (1988). New information on the species can now be reported. The operculum (Figs. 1-2) is corneus, yellowish, rounded; the edge is very thin. It is multispiral, but 1ts spire 1s not easily visible and only then in its inner part, being uniform on the outer part. The animal was observed and it is cream-white with elongate ciliated cephalic tentacles; small black eyes, not pedunculated, and a big parapodial tentacle in the right side, between the eye and the epipodium; mouth bilobulated; epipodium with three pairs of tentacles, not very long, being two of them on the opercular lobule. The observation of the rest of the body characters was impossible because the animal was very shy and it was almost all the time withdrawn into the shell. The radula (Figs. 3-4) has a formula of n.5.1.5.n. The rachidian tooth is very broad with quadrangular shape and the lateral edges strongly expanded and with a smooth cusp. The lateral teeth are of similar size, overlapping on the inner part, with the characteristic bend in the middle shaft and with long denticulated cusps overhanging the outer part. A well-developed lateromarginal plate is visible. This plate extends behind and in front of the inner marginals. The marginal teeth are long and narrow with denticulated cusps which are strongly overhanging. Habitat. The live specimens of Parviturbo insularis from Rabo de Junco, north of the Mordeira Bay, in Sal Island, were collected under stones in the upper high tide level. Remarks. ROLAN (1988) described Parviturbo insularis studying some shells collected in sediment samples from the Cape Verde Islands. These samples Cape Verde Fauna and five new species ROLAN & RUBIO were collected from depths of between 4 and 30 m. No live animals were known. So the placement of this species in the genus Parviturbo Was made only tentatively on the basis of the shell characteristics. Having been collected only in the Cape Verde archipelago, it was assumed that P. insularis Was an species endemic to these islands. The radula of Parviturbo insularis is very similar to that of Parviturbo acuticostatus (Carpenter, 1864) which is figured by HICKMAN & MCLEAN (1990, fig. 97A) in their description of Parviturbo-Haplocochlias group and by WARÉN (1992, fig. 2A). After comparing these characters, we conclude that this species was correctly placed in the genus Parviturbo. Superfamily CINGULOPSOIDEA Family CINGULOPSIDAE Fretter & Patil, 1958 Genus Eatonina Thiele, 1912 Type species, by monotypy: Eatonina pusilla Thiele, 1912. Subgenus Coriandria Tomlin, 1917 Eatonina (Coriandria) martae Rolân & Templado, 1993 Fig. 5 Material examined. See ROLAN & TEMPLADO (1993). Additionally from our last trips: 25 specimens, from Furna, Brava, and 10 more from Punta Geneanes, Santiago, several hundred empty shells from sediment collected at Brava, Ilheus Rombos, Santiago, Fogo, etc. Description. See ROLAN & TEMPLADO (1993). Further to this description it can be added the following: the radula (Fig. 5) is typical of the subgenus,; it 1s very small in size being only 35 pm in width. It has a central tooth with two small denticles on its border, lateral folds which completely lacks any denticle in its inner face. The lateral tooth is quadrangular, with its upper bord uniformly convex and the external margin with four denticles, the outside one smaller. The inner marginal tooth has three elongate denticles at its apex, of which the central one is the most prominent. The marginal external tooth has a wide base and three sharp slightly curved denticles at its extremity. Remarks. Æ. martae was described without any subgeneric assignment due to the fact that only empty shells were studied. The collection of several live specimens allowed us the complete radular study and therefore we can now confirm its location in the family Cingulopsidae and in the subgenus Coriandria. Figs. 1-4. Parviturbo insularis. Figs. 1-2. Operculum. Figs. 3-4. Radula. Fig. 5. Eatonina (Coriandria) martae. Radula. Figs. 6-8. Obtusella lata n. sp. Fig. 6. Holotype (MNCN). Fig. 7. Microsculpture. Fig. 8. Protoconch. ROLAN & RUBIO Cape Verde Fauna and five new species APEX 14(1): 1-10, 20 avril 1999 APEX 14(1): 1-10, 20 avril 1999 Superfamily RISSOIDEA Family RISSOIDAE Tomlin, 1917 Genus Obtusella Cossmann, 1921 Type species, by monotypy: Obtusella intersecta (S. W. Wood, 1857) (= Rissoa obtusa Cantraine). Europe. Obtusella lata n. sp. Figs. 6-8 Type material. Holotype (Fig. 6) of 1.11 x 0.84 mm, and 7 paratypes in MNCN (n° 15.05/32185), all from type locality. Other paratypes: 2 in MNEHN, AMNH, USNM, NNM and CFR,; 14 in CER (Fig. 8), all from type locality, collected between 15-30 m. Other material studied. 9 shells, Porto Mindelo, 25 m, Säo Vicente. Type locality. Off Pau Seco, Maio. Etymology. The specific name is derived of the latin word /atus, which means wide, alluding to be the wider dimension consistantly above the known size relevant to the European species Obtusella intersecta (Wood, 1857), at least in the Spain populations. Description. Shell (Fig. 6) of very small size, milk- white in colour, globose, thin, a little higher than wide, spire formed by 3 72 convex whorls, separating by a fine suture. Protoconch (Fig. 8) probably planktotrophic type, with 1 3/4 whorls and a maximum diameter of 310 pm. The embryonic protoconch (protoconch I) has a 7 spiral whorl scaring 3-4 spiral threads, which disappear at the beginning of the larvl phase (protoconch Il); this is totally smooth with the exception of a spiral thread above the suture. Teleoconch consists of up to 2 spiral whorls, covered completelly by fine, very depressed spiral threads (Fig. 7), inequal in size and crossed by very fine scarcely prosocline growth lines. Aperture ovoid, ortocline, inner lip and external border sharp, columela curved, reflected towards the umbilicus, which is relatively wide and deep. Dimensions. Usually, 1.0 mm in height and 0.8 mm in width. Some shells from Porto Mindelo can reach 1.2 mm. Distribution. It has been collected in muddy substrata in Maio and Säo Vicente islands, but it is probably present in the rest of the archipelago. Remarks. Obtusella lata spec. nov. has some similarity with the species of the European waters Obtusella intersecta. Both have a similar protoconch, but the shell is different because the Cape Verde ones are milk-white in colour, more globose, with a wider last whorl, the spiral threads Cape Verde Fauna and five new species ROLAN & RUBIO are of unequal size, and the umbilicus is wider. ©. intersecta is Wwhitish-cream in colour, narrower profile, the spiral threads are uniform, and the umbilicus is always narrower. Also, the protoconch of O. lata has 1 3/4 whorls, while most the spp of ©. intersecta studied from Vigo Bay have 2 whorls, although some are a little shorter. O. intersecta has been cited with its southern distribution being the Maroccan coast (VERDUIN, 1984, Cancap 1 stn. 132 and 147) but, probably it must be considered as a species with a wide distribution along the entire West African. Recently the authors have found this species in sediment samples from Angola. In spite ©. lata being in the middle of the distribution area of ©. intersecta, the oceanic islands often present this kind of isolation for species ancestrally related. Family VANIKORIDAE Gray, 1840 Genus Megalomphalus Brusina, 1871 Type species, by monotypy: Siomatia azonea Brusina, 1864. Megalomphalus serus n. sp. Figs. 9-13 Type material. Holotype (Figs. 9-10) of 1.92 x 1.45 mm, deposited in MNCN (n° 15.05/32186). One paratype in MNHN collected in the same locality, in sediment dredgings at 30 m. Type locality. Off Pau Seco, Maio Island. Etymology. The specific name is derived of the latin name serus Which means late, in allusion to it being found for the first time after many years of examining sediment samples. Description. Shell (Figs. 9-10) of small size (maximum diameter 2.5 mm), depressed, fragile, spire formed by 2 / whorls of quick development, with a evident suture. The protoconch (Fig. 11) of the holotype has a maximum diameter of 315 um, with a little more than one whorl, whose surface is aparently covered by very fine granules, and with a strong spiral cord which begins in the nucleus and continue as a keel until the end of the protoconch. The teleoconch has 1 1/2 spiral whorls which enlarge in size quickly and has a surface totally covered by very small and irregular undulating spiral threads crossed by growth lines (Figs. 12-13). Aperture quadrangular, slightly prosocline. Base with a wide umbilicus bordered by a few prominent part where there are well marked axial ribs. Animal, radula and operculum are unknown. ROLAN & RUBIO Cape Verde Fauna and five new species APEX 14(1): 1-10, 20 avril 1999 Figs. 9-13. Megalomphalus serus n. sp. Figs. 9-10. Holotype (MNCN). Fig. 11. Microsculpture. Distribution. Only know from the type locality. Remarks. The generic assignation has been made because of its similarity with the type species of the genus, M. azonea. M. serus n. sp. can be differentiated from M. disciformis and M. mercatoris by its keeled protoconch, the lack of axial sculpture and the very fine undulating spiral threads. Anyway, WARÉN & BOUCHET (1988) indicate having seen the holotype of Megalomphalus mercatoris Adams & Knudsen, 1969 and they did Protoconch. Figs. 12-13. not find any differences with the specimen of Megalomphalus disciformis identified by Monterosato as M. depressus Seguenza. Some species of the genus Macromphalina Cossman, 1888 from West Africa can have similar morphological features (M. boury Dautzenberg, 1910 and M. gofasi Rubio & Rolän, 1994), but both have multispiral protoconch. A recent revision (ROLAN & RUBIO, 1998) of the Caribbean species of Megalomphalus and Macromphalina shows that all species differ in shell shape and protoconch from M. serus n. sp. on APEX 14(1): 1-10, 20 avril 1999 Cape Verde Fauna and five new species ROLAN & RUBIO TS ———p—p—p——————————————————ee Family VITRINELLIDAE Genus Vitrinella C. B. Adams, 1850 Type species, by original designation: Ÿ. helicoidea C. B. Adams, 1850. Vitrinella politurae n. sp. Figs. 14-19 Type material. Holotype (Fig. 18) of 0.88 x 0.44 mm, and 4 paratypes deposited in MNCN (n° 15.05/32188), all from the type locality. Paratypes: 1 (Fig. 17) in MNAHN, 1 in CFR, 1 in AMNH and ! in USNM, all from type locality, collected at 4 m,; 5 paratypes (Fig. 15) in CER, 1 from Baïa Teodora, 4 m, Boa Vista, and 4 from Regona, 3 m, Sal; 2 more from Sal Rei Bay, in CFR. Other material studied. 2 shells, Porto Mindelo, 15 m, Säo Vicente; 1 shell, 1 juvenile and 1 fragment, Baia Teodora, 4 m, Boa Vista; 3 shells, Baïia da Pedrinha, 8 m, Brava; 2 shells, Furna, 15 m, Brava. Type locality. Baïia da Mordeira, Sal Island. Etymology. The specific name is derived from the latin word politura, which means polish, in allusion to the surface of the shell. Description. Shell (Figs. 14-18) very small in size, whitish, apically planispiral, with 2 spiral whorls of relatively quick development, and a few impressed sutures. Protoconch (Fig. 19) of one spiral whorl, smooth. Teleoconch totaly smooth, except for growth lines which are more evident in the umbilical zone. Aperture rounded, prosocline, with thicked columela expanded on the previous whorl forming a small callous in front of the aperture. Periostracum fine, cream in colour. Dimensions. The shells can reach maximum dimension of 1.3 mm. Distribution. So far only know from the Cape Verde archipelago, the islands of Sal, Boavista, Säo Vicente and Brava, but probably present in all islands of this group. Remarks. Vitrinella politurae n. sp. can be differentiated from its congeneric species of the West African coast because by its smaller size and the fact that it is totally smooth: V. bushi Dautzenberg, 1913 is almost smooth but has evident growth lines, and the umbilicus is narrowed by a thick columelar callous. V. annulifera Dautzenberg, 1910 has spiral cords and striae. Most of the Caribbean species has some kind of spiral sculpture; only three species are smooth: F helicoidea C. B. Adams, 1850 has the umbilicus rounded by a smooth spiral cord; VW. floridana Pilsbry & McGinty, 1946, can seem similar but it is not dorsally planispiral, and its size is bigger (the holotype, in ANSP, figured by VOKES & VOKES, 1983 has 1.95 mm). Family PICKWORTHIIDAE lredale, 1917 Genus Mareleptopoma Moolenbeek & Faber, 1984 Type species, by monotypy: Mareleptopoma karpatensis Moolenbeek & Faber, 1984. Caribbean. Mareleptopoma verdensis n. sp. Figs. 20-22 Type material. Holotype (Fig. 20) of 1.2 x 0.8 mm, deposited in MNCN (n° 15.05/32189). One paratype in each of the following. MNHN (Fig. 21), AMNH, USNM and 5 in CER, all from type locality; 2 in CFR, one from Baixos de Joäo Valente and other from Prainha, Santiago. Type locality. Praia da Cruz, north of Sal Rei, Boa Vista Island. Etymology. The specific name refers to the archipelago where 1t was collected. Description. Shell (Figs. 20-21) of small size, thick, conoid-elongate, with 3 1/2 spiral whorls, separated by a deep and canaliculated suture. The last whorl represents 65% of its height. Protoconch paucispiral, with only one whorl, apparently smooth. The teleoconch has 2 spiral whorls which have a strong spiral cord like a keel, forming an angle at the periphery. Other smaller flattened spiral cords are present on the last whorl, 5 above and 8-9 below the first one. These smaller cords as well as the interspaces have very punctiform fine sulcn (Fig. 22). There are also growth lines. The cord at the base is the strongest and it borders the umbilical infundibulum. This umbilicus has other strong cord into. The contact of the aperture with the previous whorl is only in a short area. Aperture oval, almost rounded, scarcely prosocline, with a thick external varix. Peristome continuous, wide, flat, with the border of the inner lip everted. The external border is undulating due the spiral cords on the base caused by the umbilical cords. Animal, radula and operculum are unknown. Figs. 14-19. Vitrinella politurae n. sp. Fig. 14. Bahia Teodora, Boa Vista. Fig. 15. Paratype (CER), Furna, Brava. Fig. 16. Furna, Brava. Fig. 17. Paratype (MNHN). Fig. 18. Holotype (MNCN). Fig. 19. Protoconch. Figs. 20-22. Mareleptopoma verdensis n. sp. Fig. 20. Holotype (MNCN). Fig. 21. Paratype (MNHN). Fig. 22. Detail of the sculpture. 6 APEX 14(1): 1-10, 20 avril 1999 > = D [= vo > Le TD = a (as) (so) = 3 cg es (b) APEX 14(1): 1-10, 20 avril 1999 Cape Verde Fauna and five new species ROLAN & RUBIO Distribution. Only know from Boa Vista and Santiago Islands, and the Baixos de Joäo Valente. Remarks. There is no species of the genus Mareleptopoma known from the West Atlantic coast. Most of the Caribbean species have very wide shells which for this reason are very much different from M. verdensis n. sp. The only elongate species is M. katyae Rolän, Espinosa & Fernändez-Garcés, 1990, but it is different because the less numerous spiral cords on the last whorl and the three sculptured whorls of the protoconch. Subclass HETEROBRANCHIA Superfamily VALVATOIDEA Family CORNIROSTRIDAE Ponder, 1990 Genus Tomura Pilsbry & McGinty, 1946 Type species, by monotypy: Vitrinella (Tomura) bicaudata Pilsbry & McGinty, 1946. Florida. Tomura abscondita n. sp. Figs. 23-30 Type material. Holotype (Fig. 23) of 0.75 x 0.66 mm, deposited in MNCN (n° 15.05/32190). One paratype in each of the following: MNHN (Fig. 24), USNM (Fig. 29), AMNH (Fig. 26), all from type locality, collected in sediments of 30 m; 3 more in CER and 1 in CFR, from Porto Mindelo, Säo Vicente. Type locality. Tarrafal, Santiago Island. Etymology. The specific name is derived from the latin word absconditus, which means hidden, and alludes to the long time in which it was not found during our previous studies of sediment samples from this area. Description. Shell (Figs. 23-26) very small, whitish, translucent, almost spherical, fragile, with a naticiform aspect. Protoconch (Figs. 27-28) hiperstrophic, with less that one spiral whorl visible and a maximum diameter of 157 um, smooth, except for the nucleus which seems to have small granulations. The teleoconch has 2 spiral whorls, and a microsculpture formed by very small and numerous spiral threads of variable size, which extend also into the umbilicus (Figs. 25-26). There are numerous very small growth lines. The spiral microsculpture is more attenuated in most shells in the upper part (Fig. 29) whilst being more evident in the lower part of the shell where more prominent threads alternate with 1-3 smaller (Fig. 30). Aperture rounded, prosocline with a right columela; the last whorl contacts only in a short part with the previous whorl. The external lip is fine, sharp; the internal lip is reflected towards the umbilicus forming a small callous. Animal, radula and operculum are unknown. Distribution. Only known from the Cape Verde archipelago, where it has been collected in Santiago and Säo Vicente islands. Remarks. Generic assignation is made tentatively because no live animal has been collected. This placement is based on its similarity with T. depressa from Europa. Tomura abscondita n. sp. can be differentiated from 7. depressa (Granata, 1877) because its shell is a little more globose, is totally covered by spiral threads and the callous formed by the enlargement of the internal lip does not cover the umbilicus. From the Caribbean, RUBIO & ROLAN (1998) described T. xenoskeneoides, which is differentiated because this species has a valvatoid shell lacking any spiral sculpture and enlargement of the internal lip. From T. bicaudata it can be separated because this species has a convex base, becoming a little concave near the strong angle or cord which overhangs the umbilicus, also being smooth. ACKNOWLEDGEMENTS. The authors thanks Jesüs Méndez from the CACTI of the Vigo University, for the SEM photographs; Margarita Mosquera, who sorted many of the shells studied from sediments; the Consejeria Territorial y de Medio Ambiente of the Gobierno of Canarias who, within the project of cooperation Canarias - Cabo Verde, has subventioned the project "Evaluaciôn de los recursos naturales litorales de la Repuüblica de Cabo Verde" in which the expedition Macaronesia-2 was included. REFERENCES BURNAY, L. P. & A. A. MONTEIRO. 1977. Seashells from Cape Verde Islands. Backhuys, Lisboa. BURNAY, L. P. & E. ROLAN. 1990. The family Scissurellidae in Cape Verde Islands. 4rch. Mol. 120 (1/3): 31-45. COSEL, R. VON. 1982a. Ergebnisse deutsch- portugiesischer Sam-melreisen auf den Kapverdischen Inseln (Republica de Cabo Verde). Vorläufige Liste der marinen Mollosken. Cour. Forsch.-Inst. Senckenberg 52: 15-25. COSEL, R. VON. 1982b. Marine Mollusken von Santa Luzia, Branco und Razo (Kapverdische Inseln). Cour. Forsch.- Inst. Senckenberg 52 : 27-33. COSEL, R. VON. 1982c. Marine mollusken der Kapverdischen Inseln. Cour. Forsch. -Inst. Senckenberg 52 : 35-76. FERNANDES, F. & E. ROLAN. 1988. A familia Triphoridae Mollusca: Gastropoda) no arquipélago de Cabo Verde. Publ. Ocas. Soc. Port. Malac. 11: 1Y2322 ROLAN & RUBIO HICKMAN, C. S. & J. H. MCLEAN. 1990. Systematic revision and suprageneric classification of the trochacean gastropods. Science series 35, Natural History Museum of Los Angeles County, Los Angeles. HOENSELAAR, H. J. & J. GOUD. 1998. The Rissoidae of the CANCAP expeditions, I: the genus 4/vania Risso, 1826 (Gastropoda, Prosobranchia). Basteria, 62(1-2): 69-115. MOOLENBEEK, R. G. & E. ROLAN. 1988. New species of Rissoidae from the Cape Verde Islands (Mollusca: Gastropoda) part 1. Bulletin Zoülogisch Museum 11(14): 121-126. PENAS, À. & E. ROLAN. 1997a. La familia Pyramidellidae Gray, 1840 (Mollusca, Gastropoda, Heterostropha) en Africa Occidental. 2. Los género Turbonilla and Eulimella. Iberus supl. 3: 1-105. PENAS, À. & E. ROLAN. 1997b. La familia Pyramidellidae Gray, 1840 (Mollusca, Gastropoda, Heterostropha) en Africa Occidental. 1. The genus Sayella Dall, 1885. /berus 15(1): 35-40. PENAS, À. & E. ROLAN. 1998. La familia Pyramidellidae en Africa Occidental. 3. El género Chrysallida. Iberus suppl. 4: 1-73. PENAS, A. & E. ROLAN. (in preparation). La familia Pyramidellidae en Africa Occidental. 4. Los géneros Odostomia, Megastomia, etc. ROLAN, E. 1988. Parviturbo insularis n. sp., first species of the genus for the East Atlantic. La Conchiglia 20(232-233): 26-27. ROLAN, E. 1991. Peringiella tuber new species for the Cape Verde fauna. La Conchiglia 22(258): 54- 59: ROLAN, E. 1992a. La familia Conidae (Mollusca, Gastropoda) en el Archipiélago de Cabo Verde (Africa Occidental). Tesis Doctoral. Universidad de Santiago de Compostela. ROLAN, E. 1992b. La familia Omalogyridae G. O. Sars, 1878 (Mollusca, Gastropoda) en el Archipiélago de Cabo Verde. Graellsia 47: 105- 116. Cape Verde Fauna and five new species APEX 14(1): 1-10, 20 avril 1999 ROLAN, E. & F. FERNANDES. 1989. Cerithiopsis paucispiralis n. sp. para el Archipiélago de Cabo Verde. Apex 4 (1-2): 37-39. ROLAN, E. & A. A. LUQUE. La familia Rissoinidae (Mollusca, Gastropoda) en el Archipiélago de Cabo Verde. /berus (in press). ROLAN, E. & F. RUBIO. 1992. Two new species of the genus Orbitestella Iredale, 1917 from the Atlantic Ocean. La Conchiglia 23(262): 17-20. ROLAN, E. & F. RUBIO. 1998. The genera Megalomphalus and Macromphalina (Mollusca, Caenogastropoda, Vanikoridae) in the Caribbean area, with the description of thirteen new species. Iberus 16(1): 21-72. ROLAN, E. & J. TEMPLADO. 1993. The family Cingulopsidae (Prosobranchia: Rissoidea) in the Cape Verde Islands, with the description of one new species. Basteria 57: 193-198. RUBIO, F. & E. ROLAN. 1998. Una nueva especie de Tomura (Gastropoda, Heterobranchia, Cornirostridae) del Caribe. /berus 16(1): 119-123. SAUNDERS, G. D. 1978. Note sulla malacofauna delle isole de Capo Verde. La Conchiglia 9(97- 98)15-17 20; TEMPLADO, J. & E. ROLAN. 1994. Las especies del género Crisilla y afines (Gastropoda: Prosobranchia: Rissoidae) en el archipiélago de Cabo Verde. /berus 11(2): 1-25. VERDUIN, A. 1984. On the taxonomy of some Recent European marine species of the genus Cingula s. 1. (Gastropoda: Prosobranchia). Basteria 48: 37-87. WARÉN, A. 1992. New and little known "skeneimorph" gastropods from the Mediterranean Sea and the adjacent Atlantic Ocean. Bollettino Malacologico 27: 149-247. WARÉN, À. & P. BOUCHET. 1988. A new species of Vanikoridae from the Western Mediterranean, with remarks on the Northeast Atlantic species of the Family. Bollettino Malacologico 24(5-8): 73-100. Figs. 23-30. Tomura abscondita n. sp. Fig. 23. Holotype (MNCN). Fig. 24. Paratype (MNHN). Fig. 25. Paratype (USNM). Fig. 26. Paratype (AMNH). Figs. 27-28. Protoconch. Figs. 29-30. Microsculpture. Cape Verde Fauna and five new species ROLAN & RUBIO PTE OR ans LS ES 8 cé 6 4 BONDAREV & BAIL New species of Fulgoraria APEX 14(1): 11-14, 20 avril 1999 A new species of Fulgoraria (Gastropoda: Volutidae) from the East China Sea I. BONDAREV' & P. BAIL? ! Fadeev street 21b F1.17, 335057 Sevastopol, Ukraine. 2 Square La Fontaine 2, F- 75016 Paris, France. KEY WORDS. Gastropoda, Volutidae, East China Sea, Fulgoraria isabelae nov. sp. ABSTRACT. Fulgoraria isabelae nov. sp. is described from the south of East China Sea. It is compared with related species and particularly with Fulgoraria (Saotomea) delicata (Fulton, 1940). INTRODUCTION No paper on the genus Fulgoraria Shumacher, 1817 was written since the comprehensive work of SHIKAMA (1967). Even WEAVER & DU PONT (1970) brought only some additional confusions. Only recently, extension of fishing zones as well as deeper dredgings led to the discovery of four new species: one from Viet-Nam waters, F. (F.) ericarum Douté, 1997, the second from the South China Sea, F. minima Bondarev, 1994, this latter may be linked with the subgenus Psephaea Crosse, 1871. The third, still unnamed species, from southern Kyushu, Japan, belongs to the subgenus Musashia Hayashi, 1960. The shell characters of the last one, here described as F. isabelae nov. sp., seem very close to the monospecific subgenus Saotomea Habe, 1943, represented by F. delicata (Fulton, 1940). SYSTEMATICS Family VOLUTIDAE Rafinesque, 1815 Subfamily Fulgorarinae Pilsbry & Olsson, 1954 Genus Fulgoraria Schumacher, 1817 Fulgoraria isabelae nov. sp. Figs. 1-6, 10, 12 Type Material. Holotype: length (L): 49.5 mm ; width (W): 17.6 mm, Museum National d'Histoire Naturelle, Paris. (Figs.1-2) Paratype 1. L: 46.4 mm. W: 17.2 mm. Bondarev coll. Paratype 2. L: 48.2 mm. W: 17.9 mm. Bail coll. (Figs. 3-4). Paratype 3. L: 44.9 mm. W: 17.6 mm. Bail coll. (Figs. 3-4c). Paratype 4. L: 51.9 mm. W: 16.5 mm. Douté coll.! Type locality. Off Okinawa. The exact type locality still remains uncertain. According to oral information, specimens have been collected by Japanese fishermen trawling off Okinawa in the south of the East China Sea. The exact geographical range extends probably southwards as it seems to be for Fulgoraria minima. Habitat. Between 100 and 400 m, on the usual fishing banks of the region. No more accurate information is currently available. Description. Shell small (average length: 46 mm) light, elongate- fusiform. Protoconch small (diameter: 2.3 mm) of two often corroded round whorls slighty tilted from the axis of the shell. Teleoconch of 4 or 5 convex whorls sculptured with numerous weak axial ribs and hardly visible fine spiral striae. Axial ribs conspicious from the suture to the shoulder, disappearing at the mid body whorls: 24-26 faint ribs on the penultimate, 27-29 on the last whorl. Suture indented with very narrow, slightly concave subsutural ramp. Aperture semi-ovate, with lustrous light cream surface inside. Outer lip whitish, smooth and simple. Columella straight, with one oblique distinct plait, a barely visible second adapical one found in one specimen. Siphonal notch and fasciole absent. Surface uniform, beige or light brown, contrasting with the white columella. Am Wiesenrain 11, D-79713 Bad-Säckingen- Harpolingen, Germany 11 APEX 14(1): 11-14, 20 avril 1999 Discussion. Fulgoraria isabelae may be easily separated from all species of genus Fulgoraria except one. The genus Fulgoraria comprises twenty-five species. It is divided in six subgenera. Five of these subgenera have large shells (from 100 to 250 mm) with very different morphology from F. isabelae: * Subgenus Fulgoraria s.s.: Type species: F. (F.) rupestris (Gmelin, 1791). The shells included in Fulgoraria s.s. differ from F. isabelae in their elongate shape, globose protoconch, 6- 9 columellar plaits, and a pattern of well-defined straight or weavy axial lines. * Subgenus Xurodina Rehder, 1969: Type species: F. (Kurodina) smithi (Sowerby II, 1901). The species of the subgenus Kurodina differ in their large and thin shell with one strong columellar plait. No pattern. * Subgenus Musashia Hayashi, 1960: Type species: F. (Musashia) hirasei (Sowerby II, 1912). The shells included in Musashia have a variable shape, a small protoconch, 1-3 columellar plaits, and a cancellate sculpture. No pattern. * Subgenus Vipponomelon Shikama, 1967: Type species: F. (Nippomelon) prevostiana (Crosse, 1878). The shells included in the subgenus Nipponomelon have a variable shape, a small protoconch, 2-4 columellar plaits, and a pattern of irregular bad-defined weavy axial lines divided into three bands on the body whorl. * Subgenus Psephaea Crosse, 1871: Type species: F. (Psephaea) concinna (Broderip, 1836). The shells belonging to Psephaea have an elongate shape, a medium sized protoconch, 3-5 Columellar plaits, and a colour pattern of irregular blotches or short axial bands divided into three bands on the body whorl, or reduced to a white central band. * Subgenus Saotomea Habe, 1943: Fulgoraria isabelae nov. sp. and F. (Saotomea) delicata (Fulton, 1940), the only species in the subgenus Saotomea up to now, are characterized by their similar size, uniform colour, light structure and by the unique (rarely two) columellar plait. New species of Fulgoraria BONDAREV & BAIL However, F. (S.) delicata (Figs. 7-9, 12) is easy to separate from Æ. isabelae by the grey-white duller surface, the deep suture with a narrow flat ramp, slighty angulate shoulders with 15-17 keen axial ribs on the last whorl. The protoconch is narrow (average diameter: 1.8 mm), coarsely subconical (Fig. 11). F. isabelae has a beige to brown glossy surface, suture just indented with a slightly concave ramp, rounded shoulders with 26-29 faint axial ribs on the last whorl. The protoconch is rounded, domeshaped (average diameter: 2.1 mm) (Fig. 12). F. isabelae occurs in the southern East China Sea. It is separated from F. delicata by a large gap, which is considerable for the Fulgorarinae whose geographical extension is limited due to the non-planktotrophic larval development. The sympatric F. minima Bondarev, 1994 is also small with numerous axial ribs (24 on the penultimate, 25 on the last whorl), but it differs in its strong spiral striae on the whole shell, in its deep brown-orange colour whith a conspicious white central band, and in the columella with two to four prominent oblique plaits. Etymology. This taxon is dedicated to the wife of the second author, Isabel Bail whose patience is worthy of great collaboration. CONCLUSION Fulgoraria isabelae nov. sp. is in some characters such as size, shape, texture, and columellar fold, closer to the monospecific subgenus Saotomea than to any other subgenus of Fulgoraria. It is most probable that F. isabelae may belong to this subgenus. However, the Saotomea diagnosis is mainly based on the presence of an operculum. Without animal available for study, allocation to this subgenus is thus only tentative. REFERENCES SHIKAMA, T., 1967. System and Evolution of Japanese Fulgorarid Gastropods. Sci. Rep. Yokohama Natn. Univ. Sec. 2(13): 23-132. WEAVER, CS. & J.E. DU PONT, 1970. The Living Volutes, a Monograph of the Recent Volutidae of the World. Delawaere Museum of Natural History, Monograph series 1:1-xvi, 1-375. BONDAREV & BAIL New species of Fulgoraria APEX 14(1): 11-14, 20 avril 1999 Figs. 1-6. Fulgoraria isabelae n. sp. Figs. 1-2. Holotype. L: 49.5 mm. W: 17.6 mm. Museum National d'Histoire Naturelle, Paris. Figs. 3-4. Paratype 2. L: 48.2 mm. W: 17.9 mm. Bail coll. Figs. 5-6. Paratype 3. L: 44.9 mm. W: 17.6 mm. Bail coll. Figs. 7-9. Fulgoraria (Saotomea) delicata (Fulton, 1940). L: 51.5 mm. W: 26.2 mm, Shikoku island, Japan. Bail coll. Fig. 10. Fulgoraria isabelae n. sp. Paratype 3. APEX 14(1}): 11-14, 20 avril 1999 New species of Fulgoraria BONDAREV & BAIL Fig. 11. Fulgoraria isabelae n. sp. Paratype 3. Fig. 12. Fulgoraria (Saotomea) delicata (Fulton, 1940). COLLECTION Guido T. POPPE p— — , y cs ;\ 2 [ TEL FAX,ORE-MAIL EUR For more than 20 years, Guido T. Poppe and his friends crossed the world in search for the most beautiful specimens. Not only with the aim of perfecting your collections, but also to gain more knowledge on taxonomy, nomenclature and the origins of shells. .000000000000000000000000000000 WANTED e 7. DE LT RARE SHELLS PLEUROTOMARIIDAE LAND Stanislas Leclefstraat, 8 - 2600 Berchem - Belgium Tel: 32 2 217 01 10 - Fax: 32 2 217 36 28 .000000000000000000000000000000000000000000000000000000000000000000000000e FERNAND & RIKA DE DONDER St. Franciscusstraat 55 bus 10 8400 Oostende BELGIUM Tel: +32 (0) 59 705848 Fax: +32 (0) 59 806562 WORLDWIDE SPECIMEN SHELLS AIT Families from the very common to the ultra rare, specialized in Pectinidae, Philippine shells and European shells. Free list on request, good quality shells at the best prices. Satisfaction guaranteed ! LARGE CHOIX D'OUVRAGES ET DE PERIODIQUES DE MALACOLOGIE EN FRANCAIS, NEERLANDAIS, ANGLAIS ET ALLEMAND, Liste sur demande. Vente par correspondance. Exposition permanente de coraux et Librairie de coquillages de collection, UNIVERS SOUS-MARIN KONINKLIJKE BAAN 90 B 8460 KOKSIJDE 2 058/51.28.21 High quality worldwide shells Specialists in brazilian seashells and landshells Caixa Postal 888 Agéncia Central Recife PE Brazil CEP 50001-970 Res +55(081) 241 9862 Off - Tel/fax +55(081) 423 1935 Mauricio Andrade Lima F Donax casbaih Write for Free List! Patrick FOURLINNIE Chasseur de coquillages rares 85. rue des Coteaux Fleuris - 83200 TOULON Tél. 04 94 92 96 21 - Fax 04 94 22 97 46 Gauguini APEX 14(1): 15-19, 20 avril 1999 Vexillum (Costellaria) alvinobalani n.sp. GUILLOT DE SUDUIRAUT Distribution géographique. Philippines, Bohol (Balicasag et Panglao), par 146-480 m., et Cebu. Description. Coquille de taille moyenne pour le sous-genre, atteignant 35 mm de hauteur, fusiforme, allongée, tours peu convexes, sutures incisées, subangulées à l'épaulement. Protoconque conique, composée de 2,75 tours lisses. Téléoconque composée de 9 ou 10 tours. Avant-dernier tour possédant 19 ou 20 côtes axiales et 10 ou 11 cordons spiraux. Dernier tour avec 25 ou 26 côtes axiales et 19 ou 20 cordons spiraux. Côtes axiales triangulaires, entrecroisées par des cordons spiraux régulièrement espacés, lisses, plats. Ouverture ovoïde, plus courte que la spire, environ 40% de la longueur totale; lèvre crenelée, présence de 4 plis columellaires; canal siphonal allongé, étroit. Renflement siphonal droit comportant 8 ou 9 costules obliques noduleuses à la base. Coquille blanche. Ceinture brune-orangée subsuturale, à l'exception des deux premiers tours de téléoconque. Dernier tour possédant une ceinture brune-orangée au niveau de l'angle pariétal, parfois absente chez certains spécimens, et une large ceinture de même couleur au niveau abapical. Protoconque blanc brillant, ouverture blanche transparente. Périostracum non obervé. Discussion. Vexillum (Costellaria) alvinobalani diffère de V. (C.) obeliscus (Reeve, 1844) des Philippines (Fig. 14-16) par son ouverture étroite, allongée à l'extrémité abapicale et par son dernier tour orné de 25 ou 26 côtes axiales et de 19 ou 20 cordons spiraux, au lieu de 20 ou 21 côtes axiales et de 15 ou 16 cordons spiraux chez PF (C.) obeliscus. Elle diffère également par sa couleur et ses bandes subsuturales brunes-orangées, alors que PF (C.) obeliscus est brune avec une étroite ceinture crème subsuturale. V. (C.) alvinobalani diffère de V. (C. ) macandrewi (Figs 10-13) par sa plus grande taille (30-35 mm de haut, comparé à 16 mm pour le même nombre de tours de téléoconque chez C. macandrewi) et par la base de la coquille plus élancée et plus mince. Aux Philippines, V. (C.) alvinobalani n.sp. vit en sympatrie avec Mitra (Mitra) hilli Cernohorsky, 1985, Cancilla isabella Swainson, 1831 et Vexillum (Costellaria) martinorum Cernohorsky, 1986. Etymologie. Ce nouveau Costellariidae est dédié à mon ami Alvino Balan, pêcheur de l'île de Balicasag, Bohol. REMERCIEMENTS. Je suis particulièrement reconnais- sant à H. Turner, Rovio, Suisse, pour ses remarques, ses conseils avisés, et pour les nombreuses photographies, à R. Houart, Landen, Belgique, pour la lecture et la correction du manuscrit, et à P. Bail, Paris, France, pour les clichés de types. REFERENCES CERNOHORSKY, W.O. 1967. Marine shells of the Pacific (1). Pacific Publications, Sydney, 248 pp. CERNOHORSKY, W.O. 1986. The taxonomy of some Indo-Pacific Mollusca, part 13, with description of a new species. Rec. Auckland Inst. Mus. 23: 45-57. TURNER, H. 1989. Mitroidea peu communes ou nouvelles de l'Indo-Pacifique, partie 1. Bull. Soc. Inter. Conch. 11(4): 14-30. TURNER, H. 1993. Mitroidea peu communes ou nouvelles de l'Indo-Pacifique, partie 2. Bull. Soc. Inter. Conch. 15(4): 1-27. TURNER, H. 1997. Three new species of mitriform gastropods with an illustrated check-list of the species living in the Red Sea. Argonauta 10(1-6): 3- 31: GUILLOT DE SUDUIRAUT Vexillum (Costellaria) alvinobalani n.sp. APEX 14(1): 15-19, 20 avril 1999 DD Figs. 1-6. Vexillum (Costellaria) alvinobalani n.sp., Philippines, sud-ouest de l'île de Balicasag, Bohol, 440-480 m. Figs. 1-2. Holotype MNHN, 31,7 X 18,2 mm. Figs. 3-4. Paratype 1, coll. E. Guillot de Suduiraut, 30,3 X 19 mm. Figs. 5-6. Paratype 2, coll. H. Turner, 30,7 X 18,7 mm. APEX 14(1): 15-19, 20 avril 1999 Vexillum (Costellaria) alvinobalani n.sp. GUILLOT DE SUDUIRAUT Figs. 7-9. Vexillum (Costellaria) alvinobalani n.sp. Fig. 7. Paratype 3, Philippines, Bohol, Panglao, coll. J.-P. Vezzaro, 23 X 6,8 mm (photo H. Turner). Fig. 8. Paratype 5, Philippines, Cebu, ZMA 3.98.016, 20,6 X 6,2 mm (photo H. Turner). Fig. 9. Paratype 4, Philippines, Bohol, coll. T.W. Baer, 21,9 X 6,2 mm (photo H. Turner). Figs. 10-11. Vexillum (Costellaria) macandrewi (Sowerby Il & III, 1874), Gulf of Aqaba, HUJ 37916, 15, 7 X 4,9 mm (photos H. Turner). 18 GUILLOT DE SUDUIRAUT Vexillum (Costellaria) alvinobalani n.sp. APEX 14(1): 15-19, 20 avril 1999 Figs. 12-13. Vexillum (Costellaria) macandrewi (Sowerby Il & 111, 1874), Gulf of Suez, leg R. Mac Andrew, Feb- Marc 1869, UMZC, 12,9 mm, juveniles (photos H. Turner). Figs. 14-16. Vexillum (Costellaria) obeliscus (Reeve, 1844). Figs. 14-15. Philippines, Island of Negros, Bais, 7 fms (13 m). Deux des trois syntypes BMNH 1967825; Fig. 14: 29,3 X 9,3 mm.; Fig. 15: 23,9 X 7,5 mm (photos W.O. CERNOHORSKY; courtesy H. Turner). Fig. 16. Philippines, Cebu, Mactan Island, 120 m. Coll. E. Guillot de Suduiraut, 21 X 6,5 mm. 19 Eu UN a CS if x DR aber eaa 3 ah perdet ge br on à sieM udso., et | agit 87 GAL ‘à! BAIL New species of Livonia APEX 14(1): 21-28, 20 avril 1999 A new species of Livonia (Gastropoda: Volutidae) from Northwest Australia P. BAIL Square La Fontaine 2, F-75016 Paris, France. KEY WORDS. Gastropoda, Volutidae, N.W. Australia, Livonia limpusi nov. sp. ABSTRACT. Livonia limpusi is described from deep water of Northwest Australia. It is compared with Livonia roadnightae (MceCoy, 1881) and Livonia mammilla (Sowerby I, 1844). INTRODUCTION Early in the eighties, several experimental trawlings were made by shrimp boats along the West Australian shores. Many of these have led to the discovery of unknown populations of Volutidae, e.g. Calliotectum dalli claydoni (Poppe, 1986), Calliotectum tibiaeforme forma dupreyae (Emerson, 1985), Amoria rinkensi Poppe, 1986, and Livonia joerinkensi (Poppe, 1987). Unfortunately, many hauls were very poor in shrimps and trawling was not continued in these regions; most of the prospected areas are now deserted. Some hauls were very productive in shells and yielded several specimens of unknown species in apparently restricted areas, among them the new Volutidae here described. SYSTEMATICS Family VOLUTIDAE Rafinesque, 1815 Subfamily Zidoninae H. & A. Adams, 1853 Genus Livonia Gray, 1855 Livonia limpusi nov. sp. Figs. 1-10, 15a Type material. Holotype. Length (L): 133.4 mm, width (W): 65.5 mm. WAM S.12011 (West Australian Museum, Perth, West Australia). (Figs. 1-2). Paratype 1. L: 114.8 mm, W: 61.2 mm. Bail coll. (Figs. 3-4). Paratype 2. L: 100.5 mm, W: 50.0 mm. Limpus! coll. (Figs. 5-6). l McKewen Street 6, Bundaberg 4670, Queensland, Australia Paratype 3. L: 125.9 mm, W: 64.3 mm. Limpus coll. (Figs. 7-8). Paratype 4. L: 116.0 mm, W: 63.0 mm. Douté? coll. (Fig 9): Paratype 5. L: 111.1 mm, W: 59.4 mm. Bail coll. Two additional examined collections. specimens in private Type locality. The type locality still remains uncertain. It is reliably located in a square demarcated by Broome, Karratha, Rowley Shoals and Scott Reef. Waters off Karratha seem to be the most probable area. Some specimens are presumed to come from Scott Reef. This locality seems to be uncertain because no specimen of Livonia limpusi was ever recorded so far north, whereas trawlings were so productive for the above mentioned taxa. Anyway, the geographical distribution of L. limpusi is the most northern locality ever recorded for a Livonia species. Habitat. Unknown. Probably on muddy bottom at 150-300 m deep. Description. Shell very small for the genus, solid, heavy for its size, ovate shaped with glossy surface. Protoconch large, slighty oval (diameter of holotype protoconch: 15.3 mm) globose with one and a half rounded first whorl. Nuclear and part of first whorl situated laterally, deviated at 90° on its vertical axis. Spire low, consisting of two smooth slighty convex whorls. Body whorl large without shoulder, smooth, of a regular shape with shiny surface when fresh. Sculpture of 2 Am Wieserain 11, D-79713 Bad-Säckingen- Harpolingen, Germany APEX 14(1): 21-28, 20 avril 1999 New species of Livonia BAIL spiral ridge stronger below suture and on anterior tip, almost obsolete on middle of body whorl. Aperture large, forming 80% of total shell lengh. Outer lip beveled, slighty everted, forming a rounded angle when merging backwards to body whorl. Columella arched with four thick inequal plaits. Siphonal notch wide, very shallow. Fasciole absent. Base flesh to whitish with on some shells a pattern of large axial zigzag brown lines forming an open tent- like design (Fig. 10), close to L. mammilla (Sowerby I, 1844). Uniformly coloured shells seem to be more common than the patterned ones (6 versus 2 of the examined shells). Animal and radula unknown. Discussion. This species obviously belongs to the genus Livonia: particular shell characteristics, extremely large globose protoconch, shape and pattern. It is closely related to L. roadnightae (McCoy, 1881) and Z. mammilla (Sowerby I, 1844). Livonia roadnightae has an extremely large geographical range, from Port-Stevens (New South Wales) to Alhobros Islands (West Australia) (see Fig. 20). This species shows no noticeable variation along its range. It differs from Z. limpusi by its bigger size (average length: 180 mm), by its strongly shouldered shape with thick axial ribs, and by its colour pattern made of large, irregular, blackish fine zigzag lines, never forming true tent-like design as in L. limpusi (Figs. 11-12) Comparison of a juvenile of L. roadnightae and a juvenile of L. limpusi shows the difference (Figs. 15a- 15b). According to its northernmost range, L. roadnightae seems to be sympatric with L. limpusi in a narrow area. However, this is not yet proved because of current insufficient exploration. Livonia mammilla is an eastern species whose range extends from Bass Strait and East Tasmania to east of Swain Reefs (Queensland) (see Fig. 20). Although it has the same smooth shape and the same pattern of open tent-like lines (Figs. 13-14), it differs by its giant size and bulbous protoconch, its lighter structure and its smooth surface without spiral ridges. See also the comparison of a juvenile of L. mamilla and a juvenile of L. limpusi (Figs. 15a-15c). According to our current knowledge of its distribution, a 5,000 km gap of allopatry excludes conspecificity with L. limpusi. 22 Remarks. Several problematic shells were recently discovered at a depth of 130-160 m, in an area between Kalbarri and Shark Bay. These shells are F.A.V. 293m (ABBOTTSMITH, 1969), a juvenile specimen from the West Australian Museum (Figs. 18-19), and an adult specimen in Limpus coll. (Figs. 16-17). They have a shape and a pattern very close to L. limpusi but slighty differ by their large size (up to 146 mm) and a more inflated outline. Only three specimens are available, which is not sufficient to allow a precise identification. They can be considered either as a southern range extension of L. limpusi, showing a variation in size and outline from south to north or, less likely, as a new taxon. Further material is needed to solve this problem. This population shows, more than typical L. limpusi does, a clear relationship with L. mammilla from the east coast of Australia À common ancestor, whose previous large geographical range could have been currently reduced into western and eastern populations, separated by a gap of the whole width of southern Australia, is possible. Although different by their much smaller protoconch, and especially by the absence of axial ribs, these shells may be also compared with L. quisqualis Iredale,1957 from Bass Strait, which has been interpreted up to now as a hybrid between Z. mammilla and L. roadnightae, sympatric at this locality. This interpretation remains uncertain and could be reconsidered. Etymology. This species is named in honor of Allan Limpus of Bundaberg, a well-known volute collector, who generously gave one of his specimens to the West Australian Museum as holotype. CONCLUSION Until now, the genus Livonia included four species: L. joerinkensi (Poppe, 1987), L. nodiplicata (Cox, 1910), L. mammilla (Sowerby I, 1844), and L. roadnightae (MeCoy, 1881). It now appears to include more species, either in a restricted, still unexplored area, or with a more extended range than expected. The coast of West Australia is probably the richest in future discovery, as the latter problematical Livonia leads one to suppose. REFERENCE ABBOTTSMITH, F., 1969. Multiform Australian Volutes. Ohio. 132 pp. New species of Livonia APEX 14(1): 21-28, 20 avril 1999 Figs. 1-4. Livonia limpusi nov. sp. Figs. 1-2. Holotype. L: 133.4 mm, W: 65.5 mm, WAM S.12011 (West Australian Museum,Perth, West Australia). Figs. 3-4. Paratype 1. L: 114.8 mm, W: 61.2 mm, Bail coll. 28, 20 avril 1999 New species of Livonia ÊÈE—h—h—h—.—.".".——mmmmmmmmmmmmmmm Figs. 5-8. Livonia limpusi nov. sp. Figs. 5-6. Paratype 2. L: 100.5 mm, W: 50.0 mm, Limpus coll. Figs. 7-8. Paratype 3. L: 125.9 mm, W: 64.3 mm, Limpus coll. 24 BAIL New species of Livonia APEX 14(1): 21-28, 20 avril 1999 Figs. 9-10. Livonia limpusi nov. sp. Fig. 9. Paratype 4. L: 116.0 mm, W: 63.0 mm, Douté coll. Fig.10. Typical tent- like pattern. Figs. 11-12. Livonia roadnightae (McCoy,1881). Fig.11. L: 158 mm. Great Australian Bight, Bail coll. Fig.12. Typical zigzag pattern. [Èe] cn APEX 14(1): 21-28, 20 avril 1999 New species of Livonia BAIL Figs. 13-14. Livonia mammilla (Sowerby 1, 1844). Fig.13. L: 224 mm. Bass Strait Tasmania. Bail coll. Fig.14. Typical tent-like pattern. Fig.15a. Juvenile of L. limpusi nov. sp. L: 125.9 mm. Fig 15b. Juvenile of L roadnightae (McCoy, 1881) L: 129.7 mm. Fig. 15c. Juvenile of L. mammilla (Sowerby |, 1844) L: 133.7 mm. BAIL New species of Livonia APEX 14(1): 21-28, 20 avril 1999 Figs. 16-19. Livonia cf. limpusi nov. sp. Figs. 16-17. Off Kalbarri, W.A. 130 m deep, L: 146 mm, Limpus coll. Figs. 18-19. Juvenile from Dorre Island, 23°37 S / 113°04 E, 165 m deep, West Australian Museum. APEX 14(1): 21-28, 20 avril 1999 New species of Livonia BAIL Pau Reef AS / Port Hedland ; Karretha ? { ‘( ' { Le LV 4 = LA * \ à f : + : T : t ‘ * ù + . + 2 L 2 “ + . 4 [2 + L2 Fig.20. Geographic distributions around Australia. > LE: IMPUSINOV.Sp: <> L.roadnightae PR L. mammilla 28 COURTE NOTE AUX AUTEURS (Les instructions détaillées sont disponibles sur demande) Conditions générales. 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Société Belge de Malacologie association sans but lucratif 2H'OCTOBRE 1999 SOMMAIRE Relevé de l’association de Epilepton clarkiae (W. Clark, 1852) et de Mioerycina coarctata (S. V. Wood, 1851) avec Phascolion strombi (Montagu, 1804) en Méditerranée Counting shell whorls. Remarks Description of a new species of Marginella (Volutacea: Marginellidae) from the Gulf of Guinea Taxonomic implications of syntopy : the status of Oliva truncata Marrat, 1867 (Gastropoda, Olividae) APEX Société Belge de Malacologie a.s.b.L. Editeur responsable R. Duchamps Av. Mozart, 52, B-1190 Bruxelles Comité d'édition: Dr. T. Backeljau Koninklijk Belgisch Instituut voor Natuurwetenschappen Dr. Y. Finet Muséum d'Histoire Naturelle, Genève M. R. Houart Institut royal des Sciences naturelles de Belgique (collab. scient.) Dr. CI. Massin Institut royal des Sciences naturelles de Belgique Prof B. Tursch Université Libre de Bruxelles Dr. J. Van Goethem Koninklijk Instituut voor Natuurwetenschappen Prof G. 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Cicéron, 27, bte 92, B-1140 Brussels, Belgium Comptes bancaires CCP 000-0974225-54 ou BBL 310-0770258-67 Les articles et textes présentés dans cette revue réflètent l'opinion personnelle de leur(s) auteur(s) et non pas nécessairement celle de la Société ou de l'éditeur responsable. Tous droits de reproduction, de traduction et d'adaptation des articles publiés dans ce bulletin, réservés pour tous pays. All rights of reproduction are reserved without the written permission of the board. Internet: http://www.arkham.be E-mail: cvilvens@prov-liege-be DELONGUEVILLE & SCAILLET Relevé de l’association siponculien - bivalves APEX 14(2): 29-32, 20 oct. 1999 Relevé de l'association de Epilepton clarkiae (W. Clark, 1852) et de Mioerycina coarctata (S. V. Wood, 1851) avec Phascolion strombi (Montagu, 1804) en Méditerranée. C. DELONGUEVILLE ‘ & R. SCAILLET ? ! Avenue Den Doorn, 5 - B — 1180 Bruxelles, Belgique ? Avenue Fr. Guillaume, 63 - B — 1140 Bruxelles, Belgique KEY WORDS. Bivalvia, Galeommatacea, Mioerycina coarctata, Epilepton clarkiae, Sipuncula, Phascolion strombi, Association, Mediterranean Sea, Turritella. MOTS - CLEFS. Bivalves, Galeommatacea, Mioerycina coarctata, Epilepton clarkiae, Siponculiens, Phascolion strombi, Association, Méditerranée, Turritella. ABSTRACT. This paper is an inventory of bivalves associated with the sipunculan Phascolion strombi (Montagu, 1804) in 60 empty shells of Zurritella communis Risso, 1826 found in the Mediterranean sea (Benicarlo — Spain). 32 Turritella (53 %) were occupied by the sipunculan alone and 28 (47%) were occupied by the sipunculan and one or more bivalves, namely Epilepton clarkiae (W. Clark, 1852) or Mioerycina coarctata (S. V. Wood, 1851). The association between Phascolion strombi and Epilepton clarkiae is new for the Mediterranean sea. RESUME. L'article présente un inventaire des bivalves associés avec le siponculien Phascolion strombi (Montagu, 1804) dans 60 coquilles vides de Turritella communis Risso, 1826 trouvées en Méditerranée (Benicarlo — Espagne). 32 Turritella (53%) étaient occupées par le siponculien seul et 28 (47%) contenaient le siponculien et un ou plusieurs bivalves des espèces Epilepton clarkiae (W. Clark, 1852) ou Mioerycina coarctata (S. V. Wood, 1851). L’association entre Phascolion strombi et Epilepton clarkiae est nouvelle pour la Méditerranée. INTRODUCTION La nomenclature des bivalves utilisée dans cet article est basée sur les travaux de VAN AARTSEN (1996a, b) et celle des siponculiens sur l’ouvrage de STEPHEN & EDMONDS (1972). L’association entre des représentants de la superfamille des Galeommatacea (Bivalvia) et des membres de la famille des Golfingiidae (Sipuncula) est un fait établi depuis le début du siècle. PEREZ (1924) rapporte la présence du bivalve Mioerycina coarctata (S.V. Wood, 1851) (plus généralement connu sous la dénomination Montacuta (Tellimya) phascolionis Dautzenberg & Fisher, 1925) dans des coquilles vides de Turritella communis Risso, 1826 occupées par le siponculien Phascolion strombi (Montagu, 1804). PELSENEER (1925) décrit une association voisine entre Epilepton clarkiae (W. Clark, 1852) et Golfingia vulgaris (Blainville, 1827) dans des tubes vides de Sabellaria alveolata Linné (Polychète sédentaire). D’autres associations entre ce bivalve et Golfingia pellucida (Keferstein, 1865) ainsi que Golfingia elongata (Keferstein, 1862) sont rapportées par BOSS (1965). TRONCOSO & URGORRI (1990, 1992) mentionnent, pour la première fois en Atlantique, la présence simultanée de Phascolion strombi, Mioerycina coarctata et Epilepton clarkiae dans une coquille vide de Dentalium novemcostatum Lamarck, 1818. DELONGUEVILLE & SCAILLET (1998) ont également observé l’association entre Phascolion et Mioerycina dans d’autres gastéropodes que les Turritella en Atiantique (Aporrhais, Nassarius) ou en Méditerranée (Cancellaria, Fusinus). Pour rappel, selon PEREZ (1925), le siponculien occupe l’intérieur du gastéropode privé de parties molles et obture l’ouverture de la coquille par un bouchon de sable percé de deux canaux. Le canal le plus large débouche à l’extérieur dans la partie supérieure du bouchon et permet le passage de la trompe du siponculien. Le canal le plus étroit s’ouvre dans la partie inférieure du bouchon et assure l’évacuation de l’eau hors de la coquille hôte. Le 29 APEX 14(2): 29-32, 20 oct. 1999 Relevé de l'association siponculien - bivalves DELONGUEVILLE & SCAILLET bivalve cohabitant avec Phascolion se trouve généralement fixé sur la columelle, à l’intérieur de la coquille hôte, à un emplacement situé au voisinage de l’orifice interne du plus petit des deux canaux percés dans le bouchon de sable. GAGE (1979) explique les mouvements de l’eau dans ces différents canaux. Le présent travail fait l’inventaire des bivalves récoltés dans un échantillon de 60 Turritella communis de Méditerranée occupées par le siponculien Phascolion strombi. MATERIEL ET METHODES 60 Turritella communis dont l’ouverture était obturée par un bouchon de sable percé de deux orifices (Fig. 1) ont été récoltées le ler juin 1998 sur le pont d’un chalutier à Benicarlo (Costa del Azahar - Espagne - Méditerranée). Le lieu de récolte se situe à quelques kilomètres du port (pêche journalière), une provenance plus précise n’a pu être établie. Chaque Turritella a été disséquée au moyen d’une pince coupante en commençant l’opération dans le tiers supérieur du gastéropode, puis en descendant progressivement vers son ouverture. Les bivalves présents aux côtés du siponculien ont été récoltés au fur et à mesure de leur mise en évidence. RESULTATS ET DISCUSSION Deux espèces de bivalves ont été identifiées dans l'échantillon: Mioerycina coarctata (Fig. 2) et Epilepton clarkiae (Figs. 3 et 4). L’inventaire est résumé dans le Tableau 1. 26 spécimens de Mioerycina étaient vivants, 1 était privé de parties molles et 8 spécimens brisés lors de la découpe n’ont pu être documentés. La taille des bivalves varie de 1,1 x 0,8 mm à 3,0 x 1,7 mm. Les deux spécimens vivants de Epilepton clarkiae sont de taille identique: 1,4 x 1,1 mm. Tout comme mentionné par PEREZ (1925) dans sa description de l’association entre Mioerycina coarctata et Phascolion strombi, les spécimens de Mioerycina coarctata ainsi que les deux spécimens de Epilepton clarkiae se trouvaient eux aussi à l’intérieur de la Turritella, à un emplacement situé au voisinage de l’orifice interne du plus petit des deux canaux percés dans le bouchon de sable. La présence simultanée de plusieurs bivalves (jusqu’à 4) dans une même coquille occupée par Phascolion strombi confirme les observations de PEREZ en 1925 (souvent 3) et celles de TRONCOSO & URGORRI en 1992 (jusqu’à 6). 30 Figure 1. Turritella communis Risso, 1826 - coquille hôte - 30,8 x 7,7 mm. Figure 2. Mioerycina coarctata (S.V. Wood, 1851) — 2,4 x 1,2 mm. DELONGUEVILLE & SCAILLET Relevé de l’association siponculien - bivalves APEX 14(2): 29-32, 20 oct. 1999 Figure 3. Epilepton clarkiae (W. Clark, 1852) - face interne - 1,4 x 1,1 mm. Figure 4. Epilepton clarkiae (W. Clark, 1852) - face externe - 1,4 x 1,1 mm. Nombre de Turritella Siponculien seul 32 | 53,3 Association avec M. coarctata 26 43,4 1 individu 20 333 2 individus 4 6,7 3 individus | 17 4 individus I [7 Association avec E. clarkiae 2 33 a. 1 individu 2 33 Tableau 1. contenu de 60 Turritella communis occupées par Phascolion strombi. 31 APEX 14(2): 29-32, 20 oct. 1999 CONCLUSIONS La présente observation confirme, en Méditerranée, l'association entre Phascolion strombi et Mioerycina coarctata dans des coquilles vides de gastéropodes (Turritella communis). Elle établit également l'existence d’une seconde association, moins fréquente, avec un autre Montacutidae: Epilepton clarkiae. REFERENCES AARTSEN, J.J. van. 1996a. Galeommatacea & Cyamiacea — Part I. La Conchiglia 28(279): 31-36, 61. AARTSEN, J.J. van. 1996b. Galeommatacea & Cyamiacea — Part II. La Conchiglia 28(281): 27-53, 61. Boss, K. 1965. Symbiotic Erycinacean Bivalves. Malacologia 3(2): 183-195. DELONGUEVILLE, C. & R. SCAILLET. 1998. Mioerycina coarctata (S.V. Wood, 1851). Association éthologique avec Phascolion strombi (Montagu, 1804). Arion 23(3): 98-101. GAGE, J.D. 1979. Mode of life and behaviour of Montacuta phascolionis, a bivalve commensal with 32 Relevé de l’association siponculien - bivalves DELONGUEVILLE & SCAILLET the Sipunculan Phascolion strombi. J. Mar. Biol. Ass. U.K. 59: 635-657. PELSENEER, P. 1925. Un Lamellibranche commensal de Lamellibranche et quelques autres Lamellibranches commensaux. Wimereux Trav. Stat. Zool. 9: 166-182. PEREZ, Ch. 1924. Le complexe éthologique de la Turritelle et du Phascolion strombi. Bull. Soc. Zool. France 9: 341-343. PEREZ, Ch. 1925. Sur le complexe éthologique du Phascolion strombi. Bull. Soc. Zool. France 50: 74- 76. STEPHEN, A.C. & S.J. EDMONDS. 1972. The Phyla Sipuncula and Echiura. Trustees of the British Museum (Natural History). TRONCOSO, J.S. & V. URGORRI. 1990. Nuevos datos sobre la distribucion de seis especies de Moluscos (Gastropoda y Bivalvia) en las costas de Galicia. Iberus 9(1-2): 247-252. TRONCOSO, J.S. & V. URGORRI. 1992. Associacion de Tellimya phascolionis (Dautzenberg & Fischer, 1925) (Bivalvia, Montacutidae) con el sipunculido Phascolion strombi (Montagu, 1804) en la Ria de Ares y Betanzos (Galicia, NO de España). Bol. R. Soc. Esp. Hist. Nat. (Sec. Biol) 88(1-4): 189-194. VAN OSSELAER Counting whorls APEX 14(2): 33-42, 20 oct. 1999 Counting shell whorls. Remarks. C. VAN OSSELAER Laboratoire de Bio-Ecologie, Faculté des Sciences, C.P. 160/14, Université Libre de Bruxelles, 50 av. F. Roosevelt, B-1050 Brussels, Belgium KEY WORDS. Mollusca, Gastropoda, shell, morphometry, measurement, whorl count. ABSTRACT. The suture is usually utilised for counting the number of whorls of coiled shells. This measurement can be influenced by several factors, such as the location of the centre of measurement, the orientation of the shell, the existence of several growth phases, the irregularities in the beginning of the protoconch suture, the difference in the orientation of the protoconch and the teleoconch, a gradual transition between the protoconch and the teleoconch. A simple and practical method :1s described. INTRODUCTION The number of whorls of coiled shells is a fundamental information for morphometric studies. Counting the number of whorls of a shell appears to be a straightforward operation. Yet, different authors may report a surprisingly different number of whorls for the same shell. GOULD (1989) and STONE (1995) have underlined the difficulties inherent to the process of counting the whorls of coiled shells. Authors are unanimous on what is to be counted: a whorl is a complete volution or turn of the spire of a univalve shell (ARNOLD, 1965). Experimental errors certainly do occur, but most differences stem from another source: the number of whorls can vary considerably according to the method used for counting (see Fig. 1). There is no consensus on what would be the best "standard method". The existing methods fall broadly into two categories, according to the point of origin of the measurements (see Fig.1). One group of methods uses a "centre" of the nucleus (in most cases this "centre" is not precisely defined): - method À (EHRMANN, 1956; JUNG, 1986; VALOVIRTA & VAISANEN, 1986), - method B (VERDUIN, 1977; WAREN, 1974; KERNEY & CAMERON, 1979; PFLEGER, 1989; VERMEULEN & WHITTEN, 1998), - method C (a novel method described here under). The other group of methods uses the starting point of the suture; the shell is oriented by reference to the tangent to the suture at the starting point (or to its perpendicular): - method D (DIVER, 1931), - method E (TURSCH & GERMAIN, 1985), - method F (HOENSELAAR & GOUD, 1998). Some of these methods also differ in what is considered to be the first whorl (see Fig.1). Method C systematically gives 0.25 whorl more than method B, which in turn gives 0.25 whorl more than method A. Method E gives 0.25 whorl more than method D. Methods C and E give similar results for every multiple of 0.5 whorl (in all other cases the difference depends on the distance between the start of the suture and the centre of measurement). The reported precision of the measurement varies from author to author. For instance it is 0.05 whorl for DIVER (1931) and for TURSCH & GERMAIN (1985); 0.125 whorl for VERMEULEN & WHITTEN (1998); 0.25 whorl for BAMINGER (1997). This paper aims at providing a general, comparative discussion of methodology and an evaluation of the factors influencing the accuracy and reproducibility of measurements. Ît also introduces a simple, practical method for counting whorls. LIMITATIONS To count whorls, one must be able to distinguish the different volutions. This is usually achieved by observing the suture in apical view. There is a broad agreement on the definition of the suture. It is for instance “‘fhe line of contact where two whorls meet” (FRETTER & GRAHAM, 1962), or “a continuous spiral line marking the junction of whorls in a gastropod shell” (ARNOLD, 1965), or “the junction of each whorl against the other” (ABBOTT, 1974). There are limitations to the use of the suture. For instance, in the genus Epitonium [also in Blaesospira echinus (Pfeiffer, 1864)], the last whorls are not attached and there is no suture left; one then uses the borders of the whorls, seen in apical view. In the genus Oliva, the suture is rather inconspicuous and the filament channel is used instead (see VAN OSSELAER & 33 APEX 14(2): 33-42, 20 oct. 1999 Counting whorls VAN OSSELAER e. 1 & | l 2.25 | | 2.50 A B ù \ \ 45 -—}- ee / | QT TŸ 2.47 / | A se 25 (or 2.23?) LEE 2:13 ee 2; | A D L— Ô— D E F Figure 1. Number of whorls of the same fictive shell, measured by different methods. Black dot: point of origin of measurements. White circle: point being measured. A, B, C. methods using the “centre” of the spiral. D, E. methods using tangent to the suture at the starting point (or to its perpendicular). A. method of EHRMANN (1933), JUNG (1986), VALOVIRTA & VAISANEN (1986). B. method of VERDUIN (1974), WAREN (1974), KERNEY & CAMERON (1979), PFLEGER (1989), VERMEULEN & WHITTEN (1998). C. method presented in this paper (see Figs. 10, 11). D. method of DIVER (1931). E. method of TURSCH & GERMAIN (1985). F. method of HOENSELAAR & Goubp (1998) (it is not clearly stated from where to count whorls). TURSCH, 1994). In other cases, such as adult shells of the genus Cypraea, the suture is covered by a callus and cannot be observed. DEFINITIONS Counting whorls amounts to measuring an angle. This requires a centre of measurements and two lines (centre to starting point and centre to ending point). For most Gastropod shells, the suture can be described (see VAN OSSELAER & GROSJEAN, in press) by a three-dimensional logarithmic helicospiral defined by the equations: 34 where 0 is the number of volutions of the suture. r 1s the distance from the coiling axis to the suture. z is the longitudinal coordinate from the origin of the logarithmic helicospiral.S, is the radial expansion rate of the suture, S, is its longitudinal expansion rate along the coiling axis. r, and Z, are constants and correspond to the radial and the longitudinal distances of the suture from the pole of the spiral when 6 equals zero (for details, see VAN OSSELAER & GROSJEAN, in press). In the apical view of a logarithmic helicospiral, angles and co-ordinates are measured from its pole (often called the "centre" of the spiral). One should note that: the pole is not the origin of the reference system (x=0,y=0,z=0); the pole is not the starting point of the suture (and cannot be on the actual helicospiral because r = ro £0 for 0 =0); VAN OSSELAER Counting whorls APEX 14(2): 33-42, 20 oct. 1999 ——————_—_—_—_—__ EE "+ the pole is not the "centre" of a "circular approximation" of the nucleus (see Fig. 2) because the logarithmic helicospiral is not a circle. The number of whorls of the shell (NWh) is not the number of volutions of the suture (6). One has: NWh = 6 +1. Figure 2. The pole (p) does not belong to the actual logarithmic helicospiral. It is located at a distance ro from the beginning of the spiral. It is also different from the centre of a circular approximation of the nucleus (v). SOURCES OF ERROR Errors on the centre of measurement Errors on the location of the centre of measurement can seriously affect the estimation of NWh, as shown in Figure 3. Such errors are particularly important when only a small number of whorls are counted. Such errors could be avoided because the pole of a logarithmic spiral can easily be determined by a graphical method. An elementary property of a logarithmic spiral is that all the straight lines passing through the pole cross the curve at a constant, specific angle. It follows that, if one draws a series of parallels tangent to opposite sides of the curve (Fig. 4), the pole is the point of intersection of lines joining the tangency points. This method is discussed in VAN OSSELAER & GROSJEAN (in press). Errors in orientation of the shell When the shell is oriented visually, its coiling axis rarely coincides exactly with the line of observation. The resulting error can be estimated with great accuracy by a computing method consisting in iterative recalculations of all co-ordinates in another projection plane (reprojection) and from another origin (relocation of the pole) (see VAN OSSELAER & GROSJEAN, in press). Table 1 shows that the error due to small angular errors in shell orientation is very small (about 0.01 whorl). Figure 3. Determination of the angular position of two identical points measured on the same shell. Errors on the location of the centre of measurements could affect seriously the estimation of the number of whorils. Figure 4. Graphical determination of the pole of the logarithmic spiral (p). The pole is at the intersection of lines crossing contact points of opposite parallels tangent to the spiral (from VAN OSSELAER & GROSJEAN, in press). 35 Counting whorls VAN OSSELAER APEX 14(2): 33-42, 20 oct. 1999 Achatina achatina (L., 1758) Anctus angiostoma (J.A. Wagner, 1827) Oliva porphyria (L., 1758) Visual estimation 3.667 3.673 3.680 3.674 Helix pomatia L., 1758 2.638 LPNES 2.623 l'A EE) Sum of phases 1+2 3.583 3.599 3.614 3.600 P+0 P O Corrected corrected Corrected 1.944 Table 1. Number of whorls between two well defined points of the teleoconch. After iterative recalculations by a computing method, new angles are obtained. P+O: corrections for both the location of the pole and for the projection; P: correction for the location of the pole only, O: correction for the projection only. The correction angles for the projections are less than 5°. The correction for the location of the pole are less than 1 mm. Errors due to combining protoconch and teleoconch In some shells, the protoconch and the teleoconch have a different orientation of their coiling axes (COX, 1955; SAVAZZI, 1990; JOHNSTON ef al, 1991). This phenomenon is particularly obvious in heterostrophic protoconchs (see for example ROBERTSON, 1974; BOUCHET, 1987). Small, visually undetectable differences in the orientation of both parts of the shell have been evidenced by VAN OSSELAER & GROSJEAN (in press) and seem to be widespread. This can lead to errors because, in any given position, the axis of at least one part of the shell is not correctly oriented. The poles of each part of the shell do not coincide and are sometimes far from each other (the pole of the teleoconch could, in some cases, not even be in the nucleus). Therefore, if the difference in coiling directions is large, the number of whorls of the two parts of the shell should better be counted independently. Note: Methods D and E would be very inaccurate if applied to the beginning of the teleoconch, due to an important error on the centre of measurement (see Fig. 3): Errors due to teleoconch growth phases Multiphasic growth of the teleoconch, involving changes in the direction of coiling, may also cause errors in the estimation of NWh. The biphasic teleoconch of Oliva porphyria (see VAN OSSELAER & GROSJEAN, in press and Table 1) illustrates that an angle of a few degrees between the axis of each growth phase involves only weak corrections (about 0.02 whorl). More important differences in orientation, such as found in the multiphasic, irregularly coiling Vermicularia (see GOULD, 1969; ACKERLY, 1989), require independent estimation for each growth phase. 36 Errors on starting point Locating the starting point for counting whorls 1s not always easy, on the protoconch as well as on the teleoconch. A. Protoconch. GouLD (1989 519) underlined that “Few nonmalacologists realize that one cannot unambiguously define whorl numbers from the shell's apex; there is no clear zero point at the top of a shell where winding begins”. The location of the initial point of the suture is not always obvious because the suture starts only after a part of the shell is already formed. Indeed, “the initial shell is the only part of the shell formed by simultaneous deposition of calcium carbonate, the rest being due to the addition of rings to its mouth” (FRETTER & GRAHAM, 1962 : 62). This explains why Figure 5. The methods using tangents or perpendicular to the suture (see Methods D, E) cannot be applied as such to the beginning of the teleoconch without introducing important errors on whorls estimation. The centre of measurements is far away from the pole of the spiral. In grey, the protoconch. VAN OSSELAER the early beginning of the "suture" line sometimes exhibits anomalies in curvature: a "hook" (see Fig. 6) or even an "S" shape (see MARSHALL, 1988 on Trochidae). Such a "hook" was observed by us in some specimens in the genera Oliva and Helix. Counting methods D and E, based upon the tangent to the beginning of the protoconch suture (or its perpendicular) are then subject to important variations (visually fitting a tangent at an estimated beginning of 14 PEN A Counting whorls APEX 14(2): 33-42, 20 oct. 1999 the suture can be a problem). Taking or not this “hook” into account could involve up to 0.25 whorl discrepancy in the counting (see Fig. 6). Methods using a well defined centre of measurements are less sensitive to such effects (see Fig. 6). Other problems also arise. Sometimes the protoconch suture is not visible (covered by a callus or hidden as in some heterostrophic protoconchs). B C ST ORN OX Figure 6. The beginning of the early suture line sometimes exhibits changes in curvature. These changes can be abrupt (A), more or less abrupt (B) or smooth (C). The methods based on tangent to the beginning of the suture (upper row) are more subject to important variations than the methods using a centre of measurements (lower row). Subjective interpretation of the suture starting point can lead to important variations in whorls estimations. B. Teleoconch. The transition from the protoconch to the teleoconch is sometimes gradual. In that case, there 1s a problem both for starting the count of the teleoconch whorls and for ending the count of the protoconch whorls (see Fig. 7). The whorl count will depend on the extent of the gradual transition between the two parts of the shell and on the type of the approximation that is necessarily made. One could indeed consider that the transition point is: -the last point belonging unambiguously to the protoconch, -or the first point belonging unambiguously to the teleoconch, -or the middle point between both of them (recommended). In any case, one could report the extent of the transition (which is the range of uncertainty). Errors on end point For the protoconch, the "end point" is the transition to the teleoconch (the problem of gradual transitions has already been addressed). For the teleoconch, the end point is "somewhere in the aperture". Figure 7. À gradual transition from the teleoconch to the teleoconch causes problems to estimate the number of whorls. One should take a convention on what is considered to be the transition. It could be the last point unambiguously belonging to the protoconch (1.50 or 1.62) or the first point unambiguously belonging to the teleoconch (1.69 or 1.80) or the middle point between both of them (1.60 or 1.71; notillustrated). 37 APEX 14(2): 33-42, 20 oct. 1999 This raises another problem because the contour of the aperture (or the transition) is rarely in one plane (as it is in the fictive examples of Fig. 1). Even if the aperture should happen to be planar, the centre of measurements 1s not necessarily in the same plane (Fig. 1 D, E). Some authors (EHRMANN, 1956; JUNG, 1986: VALOVIRTA & VAISANEN, 1986) consider the "end point" as being the middle of the whorl, seen in apical view (point M in Fig. 8). Other authors (DIVER, 1931) consider the last point of the suture (point S in Fig. 8). Others yet (TURSCH & GERMAIN, 1985; BIELER, 1993) consider the most external point (point E in Fig. 8). Sometimes, such information is not clearly stated, as in HOENSELAAR & GOUD (1998). In some cases, the choice of one or another of these conventions can produce large differences in whorl counting (Figs. 7, 8). Other conventions could also be considered. The aperture (and/or the transition) is sometimes tilted (prosocline or opisthocline apertures, see Fig. 9). One could select the most adapical, the most abapical or the mid-point as being the "end point". The difference resulting from that choice could be particularly important in the case of sinugera protoconchs (sensu BOUCHET, 1987: 19). Figure 8. To count whoris, authors consider the "end point" as being the middle of the whorl (point M), or the last point of the suture (point S), or the most external point (point E). These points being rarely aligned along one radius from the centre, a convention must be specified. (see also Fig. 7 for the protoconch). 38 Counting whorls VAN OSSELAER We advocate the following, simple convention (similar to that of TAYLOR [1975] in JABLONSKI & LUTZ [1980]) : the last point of the suture is considered to belong to the last whorl and to be its end point (NWh = 0 + 1, see Definitions). The previously mentioned problems then cease to exist. [MÈ2 Sa ER & È Et re & 8 É 2e = Figure 9. The aperture and/or the transition can sometimes be tilted (prosocline or opisthocline). Unless the final point of the suture is chosen as the “end-point” of counting, one must specify if the "end point” is the most adapical, the most abapical or the mid-point. A RAPID, PRACTICAL METHOD During morphometrical analyses in the genus Helix, difficulties were encountered because the early protoconch “hook” (see above) is often present and, in addition, the protoconch-teleoconch transition is often gradual. Counting the total number of whorls was therefore preferred and a simple method was developed. Two variants of a this method will be described. A. From a drawing. The suture of the shell is drawn in apical view, using a binocular lens equipped with a camera lucida. Starting from the last point of the suture (point S, Fig. 10 A), one draws the line L; which crosses the suture at its earliest point where a perpendicular, passing through point S, can be traced (see Fig. 10 D). Then one draws a second line L;,, perpendicular to L, and perpendicular to the earliest possible point of the suture (see Fig. 10 B,E). The intersection of L, and L, defines the centre of measurement (see Fig. 10 C, F). From the centre, a line L; is drawn to the starting point of the suture. The total number of shell whorls (NWh) is estimated from the number of whorls (8) of the suture (NWh = 6 +1). Starting from the centre to the last point of the suture (S), the number of intersections between L, and the suture gives the integer part of the number of whorls. The angle between L, and L;, expressed in decimal of a complete whorl (not in degrees) and measured against the direction of coiling, is the non- integer part of the number of whorls. VAN OSSELAER B. Quick variant. This does not require a drawing of the shell suture and the number of whorls can be visually estimated. A star-grid with a grid of lines at 0.025 whorl (or less) angle (see Fig. 11) is seen trough the camera lucida, superimposed to the view of the shell. The line 0.00-0.50 and the line 0.25-0.75 are used as the lines Lo and L, (sliding the grid brings it in the desired position, see Fig. 11 A, B). The counting procedure is then the same as described here above. C. Precision. The precision of these very rapid methods 1s 0.05 whorl. This was estimated by ten repeats of measurement on one specimen of Helix pomatia (mean : 5.11 whorls; Minimum: 5.10 whorls; Counting whorls APEX 14(2): 33-42, 20 oct. 1999 Maximum : 5.15 whorls ; Standard deviation: 0.02 whorls ; coefficient of variation : 0.34 %). D. Discussion. The whorls of the whole shell can be determined in one operation only if the beginning of the first whorl of the protoconch can be clearly seen and the last whorl of the teleoconch suture located at the same time. Separate determinations of NWh for the protoconch and the teleoconch could be done on the same drawing and are then straightforward. Using a binocular lens, this method can be difficult for large and high spiralled shells, but remains valid for their protoconch. Figure 10. Description of a rapid, practical method to count whorls on two schematic examples. À & D. Starting from the last point of the suture (point S), a line Lo is drawn perpendicular to the first whorl (the first perpendicular crossing is considered at point a). B & E. Then a line L:, perpendicular to Lo, is also drawn perpendicular to the first whorl (the first perpendicular crossing is considered at point b). The intersection of Lo and L. defines the centre of measurement (point c). C & F. From this centre, a line L2 is drawn to the start of the suture. The angle between Lo and L> allows the count of whorls. 39 APEX 14(2): 33-42, 20 oct. 1999 Counting whorls VAN OSSELAER 0.50 0 —S ie | | LS \ 0 2° j | ? ; k NZ | f \ A NZ 0.75 = 1 | A — S 7 : — 0.25 01° e x ï \ 27 2 à Figure 11. Quick variant of the previous method (see Fig. 10). The number of whorls can be visually estimated. A star-grid (lines at 0.025 whorl) is seen trough the camera lucida, superimposed to the view of the shell. The lines 0.00-0.50 and the line 0.25-0.75 are used as the lines Lo and L; (see Fig. 10). Sliding the grid brings it in the desired position. When the whorls of the whole shell cannot be determined in one operation, this method could be applied to the two parts of the shell independently. Different centres will be found for each part of the shell. Each of these centres will most often differ from the centre found if the whole shell is considered. In these cases, the present method to determine NWh still remains highly reproducible for the different parts of the shell but is not necessarily accurate for the teleoconch. The distance separating (see Fig. 2) the centre (c) and the pole of the spiral (p) could become more important when applying the method only to the teleoconch. This distance is in direct proportion to the curvature of the spiral and to the size of the protoconch (‘“diameter” of the first whorl of the teleoconch). When the distance between the centre (c) and the pole of the spiral (p) is too important, this rapid method cannot be applied without introducing large bias (see Sect. Errors on the centre of measurement). It is therefore advised to use the pole of the spiral. 40 The method can be compared with that of VERDUIN, the only author having shown how to place the centre of measurement. For VERDUIN (1977), the centre of measurement is the centre of the nucleus, which is approximated by a circle (the diameter of the first half visible whorl is the diameter of the nucleus circle). There are two main differences between the new method and method A. The first difference is the location of the centre of measurement, which it is not placed at the middle of the first half whorl as in VERDUIN's method (Fig. 12 D). Let us notice that in the case of abnormal curvature of the early suture (see Errors on starting point: A. Protoconch), several different "circular approximations" of the nucleus could be made. The method of VERDUIN could then become quite subjective (see Fig. 12 À, B, C). Starting from point S, the proposed method avoid subjectivity in the appreciation of the beginning of the suture and 1s then particularly useful. The centre determined by the new method does also not correspond to the pole of the logarithmic VAN OSSELAER Counting whorls APEX 14(2): 33-42, 20 oct. 1999 helicospiral (see Fig. 12 D) but it is often very close to it. However, the small error on the centre location has little effect on the whorl count (for similar error, see Table 1). The second difference with method A is the starting point for counting whorls. The beginning of the suture is here considered to be the first whorl of the shell; this gives 0.50 whorl more than method A (see Fig. 1). JABLONSKI & LUTZ (1980) used the method of TAYLOR (1975, in JABLONSKI & LUTZ [1980]) which concerns only the protoconch. This method appears close to ours, but the way to count the non-integer part of the number of whorls (from where?) is not indicated and can not be compared. In this method, the line starting from the sutural point of the aperture is defined perpendicular to and crossing the remaining apical sutures (JABLONSKI & LUTZ, 1980: 330). It must be noted that a line can not be perpendicular to several whorls of a logarithmic helicospiral. This can be the case for arithmetic spirals (r = a . 0), but a line starting from the sutural point of the aperture does not necessarily cross the centre of the spiral. For some Oliva species, an arithmetic helicospiral appears to be more representative of the protoconch suture while the logarithmic helicospiral remains the best model for the teleoconch (unpublished results). The rapid method advocated here above can be applied to both arithmetic or logarithmic spirals. ACKNOWLEDGEMENTS. I am particularly grateful to B. Tursch for his constructive criticism and for remodelling the manuscript. I would like to thank F. Cherot and Ph. Grosjean for helpful discussions and suggestions. This work was supported by a doctoral fellowship of the Université Libre de Bruxelles and by Floridienne Chimie S.A. Figure 12. If the early protoconch presents a “hook”, several “circular approximations” of the first half whorl (nucleus) can be made (A, B, C). Each of these leads to a different centre of measurements (v). The pole of the logarithmic helicospiral (p), the centre of measurements of the here described method (c) and the centre of the “circular approximations” (v) of the nucleus are rarely (if ever) coinciding (D). 41 APEX 14(2): 33-42, 20 oct. 1999 REFERENCES ABBOTT, R.T. 1974. American Seashells. Van Nostrand, N.Y. ACKERLY, S.C. 1989. Shell coiling in gastropods: analysis by stereographic projection. Palaios 4: 374- 378. ARNOLD, W.H. 1965. A glossary of a thousand-and- one terms used in conchology. Veliger 7(supplement): 1-50. BAMINGER, H. 1997. Shell-morphometrical characterization of populations of Arianta arbustorum (L.) (Gastropoda, Helicidae) in the Ennstaler Alpen (Styria, Austria). Ann. Nat. Mus. Wien 99 B: 497-519. BIELER, R. 1993. Architectonicidae of the Indo-Pacific (Mollusca, Gastropoda). 4bh. Naturw. Ver. Hamburg (NE) 30 :1-376. BOUCHET, P. 1987. La protoconque des gastéropodes: aspects biologiques, taxxonomiques et évolutifs. Thèse de Doctorat d'Etat. Univ. Pierre & Marie Curie. Paris VI. Cox, L.R. 1955. Observations on gastropod descriptive terminology. Proc. Malac. Soc. Lond. 31: 190-202. DIVER, C. 1931. A method of determining the number of the whorls of a shell and its application to Cepaea hortensis Müll. and C. nemoralis L. Proc. Malac. Soc. Lond. 19: 234-239, EHRMANN, P. 1956. Die Tierwelt Mitteleuropas. II(1). Weichtiere, Mollusca. Quelle & Meyer, Leipzig. FRETTER, V. & A. GRAHAM. 1962. British Prosobranch Molluscs. Ray Society. London. GOULD, S.J. 1969. Ecology and functional significance of uncoiling in Vermicularia spirata: an essay on gastropod form. Bull. Mar. Sci. 19: 432-445. GOULD, S.J. 1989. A developmental constraint in Cerion with comments on the definition and interpretation of constraint in evolution. Evolution 43(3): 516-539. HOENSELAAR, H.J. & J. GOUD. 1998. The Rissoidae of the CANCAP expeditions, l: the genus A/vania Risso, 1826 (Gastropoda Prosobranchia). Basteria 62: 69-115. JABLONSKI, D. & R.A. LUTZ. 1980. Molluscan larval shell morphology. Ecological and paleontological applications. In RHOADS, D.C. & R.A. LUTZ (eds.). Skeletal growth of aquatic organisms. Topics in 42 Counting whorls VAN OSSELAER Geobiology, Vol. [., Plenum Press, New York & London. JOHNSTON, M.R., R.E. TOBACHNICK & F.L. BOOKSTEIN. 1991. Landmark-based morphometric of spiral accretionary growth. Paleobiology 17(1): 19- 36. JUNG, P. 1986. Neogene paleontology in the northern Dominican Republic. 2. The genus Strombina. Bull. Amer. Paleont. 90(324): 1-42. KERNEY, M.P. & R.A.D. CAMERON. 1979. A field guide to the land snails of Britain and north-west Europe. Collins, London. MARSHALL, B.A. 1998. A review of the recent Trochini of New Zealand (Mollusca: Gastropoda: Trochidae). Moll. Res. 19(1): 73-106. PFLEGER, V. 1989. Guide des coquillages et Mollusques. Hatier, Fribourg, Suisse. ROBERTSON, R. 1974. Marine prosobranch gastropods: larval studies and systematics. Thalassia Yugoslavica. 10(1-2): 213-238. SAVAZZI, E. 1990. Biological aspects of theoretical shell morphology. Lethaia 23: 195-212. STONE, J.R. 1995. Cerioshell: a computer program designed to simulate variation in shell form. Paleobiology 21: 509-519. TURSCH, B. & L. GERMAIN. 1985. Studies on Olividae. I. À morphometric approach to the O/iva problem. Indo-Malayan Zool. 2: 331-352. VALOVIRTA, I. & R.A. VAISANEN. 1986. Multivariate morphological discrimination between Vitrea contracta (Westerlund) and F. crystallina (Müller) (Gastropoda, Zonitidae). J. Moll. Stud. 52: 62-67 VAN OSSELAER, C. & PH. GROSJEAN. In press. Model of the suture and location of the coiling axis in gastropod shells. Paleobiology. VAN OSSELAER, C. & B. TURSCH. 1994. Studies on Olividae. XIX. Where is the suture of Oliva shells? Apex 9(2/3): 47-50. VERDUIN, À. 1977. On a remarkable dimorphism of the apices in many groups of sympatric , closely related marine gastropod species. Basteria 41: 1-95. VERMEULEN, J.J. & A.J. WHITTEN. 1998. Fauna Malesiana field and study guide series. Backhuys Publ., Leinde, Dutchland. WAREN, A. 1974. Revision of the Artic-Atlantic Rissoidae (Gastropoda, Prosobranchia). Zool. Scr. 3: 121-135. BOYER, RYALL & WAKEFIELD Marginella xicoi, n.sp. APEX 14(2): 43-52, 20 oct. 1999 Description of a new species of Marginella (Volutacea: Marginellidae) from the Gulf of Guinea F. BOYER ‘ , P. RYALL ? & À. WAKEFIELD 1110, Chemin du Marais du Souci, 93270 Sevran, France. ? P.O. Box 224, Takoradi, Ghana. * 14 Forest Side, Buckhurst Hill, Essex, 1G9 5SL, England. KEY WORDS. Taxonomy, Marginellidae, Marginella, M. xicoi n.sp., M. tyermani, West Africa, Gulf of Guinea. ABSTRACT. The taxa belonging to the group Marginella tyermani Marrat, 1876 are reviewed. M. keppeli Sykes, 1905 and M. eveleighi Tomlin & Shackleford, 1913 are defined as synonyms of M. tyermani. Marginella xicoi n.sp. is described from the Ghanaean coast and is related with other species to a “complex M. tyermani”. RÉSUMÉ. Les taxa appartenant au groupe Marginella tyermani Marrat, 1876 sont revisées. M. keppeli Sykes, 1905 et M. eveleighi Tomlin & Shackleford, 1913 sont placés en synonymie de M. tyermani. Marginella xicoi n.sp. est décrite de la côte du Ghana et est reliée avec d’autres espèces à un “complexe M. tyermani”. INTRODUCTION In February 1995, whilst dredging west of the mouth of the Volta River, Ghana (Fig. 1), the second author discovered several specimens of a small species belonging to the genus Marginella and considered by us as new to science. Our new species presents evident similarities to taxa belonging to the group Marginella tyermani, so we propose their preliminary revision as a necessary step to the determination of our new species. This century, very few works have dealt with Marginellids from the Gulf of Guinea. The principal ones are: - KNUDSEN (1956), for scattered samplings in deep waters during the campaign of the vessel ‘Atlantide’. -TOMLIN & SHACKLEFORD (1913), GOFAS & FERNANDES (1988), and FERNANDES & ROLAN (1991) for shallow water samplings in Sao Tome and Principe. - BERNARD (1984), for material mostly collected from shallow and moderate depths in Northern Gabon. The paucity of scientific works on the subject clearly illustrates the very limited knowledge of the Marginellid fauna from the Gulf of Guinea. The southern part is perhaps best known, but even then only a few kilometers of coastline have been studied. The results of some recent collecting trips along the northern coasts of the Gulf of Guinea (ROLAN & FERNANDES, 1977 ; P. Ryall, personal material and records) suggest that the claimed “poor diversity” of the marine molluscan fauna in this area could be a superficial and inaccurate point of view. The traditional view of Atlantic equatorial coasts, with their warm, muddy water, long sandy beaches and heavily rolling waves is of a generally inhospitable environment unable to support more than a few hardy species and occasional very specialised ones. The reality seems to be very different, as the benthic equatorial environment of the continental shelf in the Atlantic generally presents a high level of organic content as well as important heterogeneity of habitats and communities along hard and soft bottoms. Therefore it is probably necessary to re-evaluate the real diversity of the molluscan fauna in the Gulf of Guinea, with the help of future field studies, collecting, and an accurate examination of available material. The study of our newly discovered species is conceived as a contribution to this knowledge. 43 APEX 14(2): 43-52, 20 oct. 1999 Marginella xicoi, n.sp. BOYER, RYALL & WAKEFIELD GUINEA + ! 1 ES + ‘ N &, IVORY : s S \ LIBERIA | COAST", Cape Palmas Fig. 1. Map of Central West Africa. SYSTEMATIC ACCOUNT Genus Marginella Lamarck, 1799 Type species: Voluta glabella Linne, 1758, by monotypy. Marginella tyermani Marrat, 1876 Figs. 2-7 Original designation. MARRAT, 1876. p. 136. Original description. “Marg. Testa subfusiformi — ovata, cinereo — alba, lineis nigris, spira brevi, antifractibus superne obtuse angulatis, ad angulum plicato — nodulatus, columella quadriplicata, labro late incrassato, intus denticulato”. “Var. with the lip smooth inside. The Corisco specimen. Found in company with M. belli, Sow., and 1s about the size of M. festiva, Kiener. Hab. 1. Corisco Bay, 1. Near Cape Palmas, West Africa. Coll. Keen, Liverpool”. 44 GHANA , Principe, Corisco 0 Sao Tome Cape Lopez NIGERIA 7. PR GAS CAMEROON GULF OF GUINEA mé 2 >. 1 o . ce: : GABON M ‘ {7 CONGO ANGOLA Type material. The description of the species referred to two non- figured specimens, from Coll. Keen, Liverpool, both present now in the collection of F.P. Marrat, Liverpool Museum, England: a larger, rather worn specimen, 11.20 x 5.90 mm (Figs. 2 & 3), and a smaller more glossy specimen, 8.55 x 5.00 mm (Figs. 4 & 5). MARRAT (1876) designated the one “with the lip smooth inside” as being a variety (“Var.”), and noted the locality as Corisco Bay, which presently is situated in Equatorial Guinea, on the border with Gabon. We therefore have to infer that the other specimen said to come from Cape Palmas, presently on the border between Liberia and Ivory Coast, represents the typical form and has to be considered as the holotype. In reality the distinction between the typical and the varietal form is not apparrent, as neither of the shells has a smooth inner lip. The Latin text of the description is also contradictory; the shell is said to be light grey, to have a short spire and a widely thickened labrum, whereas in fact the larger specimen is light tan coloured with a tall spire and strongly thickened labrum with deep internal denticulations. The smaller specimen is light greenish grey, with a short spire and a moderately thickened labrum. So, it seems that the text of the description mixes up characteristic features of both shells, and does not really present the individual features of a single “typical specimen”. 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Boyer, left to right: 12.20 x 6.30 mm, 9.00 x 5.60 mm, 7.30 x 4.15 mm. APEX 14(2): 43-52, 20 oct. 1999 Marginella xicoi, n.sp. BOYER, RYALL & WAKEFIELD the labels attached to these specimens do not allow us to be absolutely certain of the geographical origin given by MARRAT for each shell, and the mention of the word “type” attached to the larger specimen seems to have been attributed later, probably by TOMLIN who examined this material first in early 1913 and again in June 1936. TOMLIN left a label certifying the authenticity of the type specimens of M. tyermani, and the type localities, and he also added: “but it is not stated from which loc. the type came.” In 1913, TOMLIN considered as evident that the type specimen was the larger one. He commented that it was “rather worn” and with ‘“an unusually thickened outer lip, which emphasises the internal denticulation.” It seems apparent that TOMLIN, as the first revisor, did not designate a lectotype from undiscriminated original material, but recognised an evident holotype as such. Despite the ambiguity of the original description, we follow Tomlin in recognising the largest shell as the holotype, as the strongly denticulated outer lip can be considered a distinctive feature, as opposed to the moderately denticulated outer lip of the smaller specimen, which can more easily be attributed as having a “smooth lip”. In conclusion, the larger specimen belonging to the Marrat Collection has to be considered as the holotype of the species, whereas the smaller specimen, being representative of a “variety” of the species, cannot be included amongst the type material. Type locality. Cape Palmas (Liberia). Other material examined. - One live collected specimen, dredged 25-30m off Cape Palmas, Liberia, 1996. Coll. P. Ryall (ex. Coll. Nora, Porto). The shell is pure milky white except for the first whorls which are greyish-brown, the surface of the body whorl is perfectly smooth, without tendency to any axial ribbing. The shoulder is rounded and does not present any angle or crenation. The inner labrum is moderately denticulate. 10.60 x 6.04 mm. Morphologically this specimen matches with the typical form of M. tyermani. Its origin 1s, however to be confirmed. - One live collected specimen, off Ivory Coast, Coll. F. Boyer, 7.05 x 4.35 mm, live collected, (Fig. 6). This specimen matches the specimen illustrated in KNUDSEN (1956: pl IT, Fig. 17) as M. musica, and does have a perfectly smooth labrum which could of course represent an immature phase of shell development. The blunt spire of this specimen bears a very low and wide protoconch. The background colour is greenish with black marks and axial flammules. - Three large specimens from Punta Eviondo, Rio Muni, (currently Equatorial Guinea). Coll. F. Boyer: 11.15 x 6.00 mm, 10.05 x 5.45 mm, 10 x 5.50 mm. Shell colour varying from greenish-yellow to greenish- 46 grey with dark spiral lines and blurred axial marks, sometimes more dense and arrow shaped under the shoulder. - From the Komo Estuary and Banie Island, Gabon, Coll. F. Boyer: one large (12.20 x 6.35 mm), and sixteen smaller shells (7.15 to 9.00 mm in length). AII adult shells, from shallow water. The shells from Banie Island exhibit the greatest diversity of colour from near all black to near all white. AII have a denticulated labrum, except for the larger specimen from Banie Island, which is a young adult with a thin lip and no denticulations. The labial denticles appear to be a sign of maturation, and may disappear again when the animal reaches the gerontic stage, the lip becoming even more thickened. - Off Banie Island, Gabon, Coll. P. Ryall. Many small to large specimens collected at 2-4 metres on fine sand. The 18 specimens attributed to M. tyermani figured in the book ‘Shells of Gabon’ (BERNARD, 1984) present the range of colour variation for the species in Northern Gabon, from dark black through yellow, grey, tan and white forms, with or without a black central band on the body whorl. BERNARD (1984) does not, however, comment on the morphologic variation of the species (outline, proportions of the spire and labrum, strength and number of denticles), just giving 10 mm as being the maximum size reached by the shell. Observed populations are said to be found on sand at 0- 5 metres, in Banie Island, Komo Estuary, and Port Gentil, Gabon. Remarks. As far as shell characters are concerned, the range of variability presented by M. tyermani in Equatorial Guinea and Northern Gabon includes both the typical form of the species (the large specimen with a slender spire said to come from Liberia) and the varietal form of Marrat (the squatter specimen with a short spire said to come from Corisco Bay). Intergrades between these two forms are fully represented, both for morphological features and colouration of the shell. The specimen figured in KNUDSEN (1956: p.91, pl. I, fig. 17) and assigned to M. musica Hinds, was collected by the Atlantide Expedition (1945-1946), at station 98, 5 56° N, 4 26° E, 100 metres deep. This station is located off the coast of Nigeria. The shell figured measures 7.30 x 4.40 mm and has a very low conical spire, five black spiral lines on the body whorl, a relatively wide aperture, and a thin labrum with a faintly denticulated inner part. Knudsen describes the shell as being “greyish white with a white band at the suture”. This shell clearly belongs to the species M. tyermani, even if it bears unusual spire morphology (it is comparable with one end of the natural range of variation of the populations observed from Northern Gabon). It also seems to be a dead shell, possibly transported to deeper levels by strong ocean currents, and its lack of axial decoration on the body whorl coupled with the presence of spiral lines, probably led to its incorrect identification as M. musica. BOYER, RYALL & WAKEFIELD Marginella keppeli Sykes, 1905 Fig. 8 Original designation. SYKES, 1905, pp. 315-316, pl. XVII, fig. 3. Type material. One syntype in BM(NH), London, coll. Sykes, not examined, described as measuring 9.5 x 5 mm. Type locality. West Africa. Remarks. The original figure and description of M. keppeli Sykes both perfectly fit into the range of natural variation of M. tyermani Marrat, concerning morphology of the shell, size and decoration. TOMLIN (1913, 1917) synonymized M. keppeli Sykes with M. tyermani Marrat, and we follow him in this conclusion. Fig. 8. M. keppeli Sykes. Original figure. Marginella eveleighi Tomlin & Shackleford, 1913. Fig. 9 Original designation. TOMLIN and SHACKLEFORD, 1913, p. 11, Type figure in TOMLIN, 1913, pl. I, figs. 5-6. Type material. One syntype in the Tomlin-Melvill Collection (National Museum of Wales, Cardiff). Not examined. Described as measuring 7.25 x 4.00 mm. Type specimen drawn as holotype in GOFAS & FERNANDES (1988), length 6.20 mm. Marginella xicoi, n.sp. APEX 14(2): 43-52, 20 oct. 1999 Type locality. “S. Thome Island, Gulf of Guinea. Scarce, in coral gravel.” Other material examined. - One adult specimen from Sao Tome, Coll. F. Boyer, 7.50 x 4.00 mm. Black and white background. - Three adult specimens and a fragment, Esprainha and Sao Tome City, Sao Tome Island, Coll. P. Ryall, collected in 1-3 metres under rocks. Other material quoted and illustrated. - Several shells and specimens referred to as M. eveleighi in KNUDSEN (1956, p. 84, pl. IL, fig. 2), through 7 stations of the Atlantide Expedition: One station off Monrovia (Liberia) in 10 metres (Stat. 52) Five stations off the Niger delta (Nigeria), in 19 to 50 metres (Stat. 100, 104, 111, 112, 113). The shell figured in plate IIT is the one taken in 32 metres at station 113. The type of substrate was not recorded for these stations, with the exception of station 112 which is described as “clayish mud”. In a previous work on the Atlantide material, KNUDSEN (1950, pp. 112-114, fig. 20 C-D) recorded the discovery of “15 oval and rather domed” egg capsules adhering to the surface of a decaying leaf, from 50 metres at station 104, and labelled as “ M. eveleighi”. However, although these egg capsules were found at the same station as a live specimen of “M. eveleighi”, it does not necessarily mean that the egg capsules belong to this taxon. The comparison made between the embryo and the apex of M. eveleighi is similarly unconvincing. The white larval shell has no decorative features, and its very long and oblique anterior columellar plait (well separated from the other three smaller and more closely grouped ones) better suggests that it belongs to another species. Fig. 9. M. eveleighi Tomlin and Shackleford. Original figure. 47 APEX 14(2): 43-52, 20 oct. 1999 On the other hand, KNUDSEN (1950, pp. 115-116, fig. 22 E-D) records the discovery of 8 egg capsules “of the plano-convex type” in 40 m at stat. 103, labelled as “Marginella sp.”. The larval shell is decorated on the last half whorl by “three dark transverse bands crossed by a single longitudinal one”. These egg capsules and larval shells are quite likely referrable to the phena “MW. eveleighi”, even if the spiral black line is generally situated just under the suture of the first whorls, rather than in a more anterior position as seems to be the case in the figure 20 E-D. One station off Luanda (North Angola) in 42 m (stat. 136, St. Paul de Loanda, mud). The occurence of M. eveleighi in this place is not confirmed by GOFAS & FERNANDES (1994), who intensively dredged at this depth over many years. So this shell collected by the Atlantide Expedition could possibly be a juvenile shell of M. lucani Jousseaume, 1884 (which is known to live from Southern Gabon to Northern Angola in shallow water, Fig. 10) or perhaps more probably, it is a juvenile specimen of M. fumigata Gofas and Fernandes, 1994, which lives in the same area but at deeper levels (type loc. : 40-50 m off Luanda). [N.B. One single datum is quoted by KNUDSEN (1950) for measurements of the shells: height 10.70 mm. This length probably concerns the shell taken at St. 113 and figured in plate III ]. - Five shells referred to as M. eveleighi in BERNARD (1984, pp. 88-89, pl. 39, no. 157), but looking very similar to the specimens presented as M. tyermani (no. 159) except for their larger size. These large specimens are said to live on sand in 2-5 m, only off Banie Island. BERNARD notes that the specimens of M. eveleighi bear few axial ribs, and that the specimens of M. tyermani are smooth. However, the figures of both species clearly show moderate to strong axial ribbing along their shoulders. In fact, the specimens designated as M. eveleighi in BERNARD (1984), represent the taller variant of M. tyermani. Large specimens over 10 mm are not found in the Komo Estuary and Port Gentil. They are moderately rare in Banie Island, and seem to be more common in Equatorial Guinea. This kind of geographic or bathymetric cline in size range is commonly seen in many species of marine gastropods. The relative rarity of these larger specimens of M. tyermani in the field explains why they are poorly represented in collections, and also explains why they exhibit the most common pattern of decoration of the species (fine spiral black lines on a greyish -green background and a central black band on the body whorl). Within individual stations all size intergrades exist, so there is no need to subdivide the species on a morphological or taxonomic basis. In fact the holotype of M. tyermani is 11.20 mm long and itself links (if it is necessary to prove this) the smaller shells to the larger ones referred to in BERNARD. - Several specimens and shells recorded in GOFAS & FERNANDES (1988, pp. 11-13, figs. 12-13) as M. eveleighi : 22 specimens from Sao Tome in Coll. F. 48 Marginella xicoi, n.sp. BOYER, RYALL & WAKEFIELD Fernandes, Angola ; 1 specimen from Calypso cruise “Gulf of Guinea”, in the Paris Museum (MNHN) ; 10 specimens from Gabon, ex-Coll. P. Bernard, in the Paris Museum (MNHN). All these shells from both Sao Tome and Gabon are 6 to 9 mm in length, and present a homogenous morphology and colour pattern “with a rather high conical spire and axial folds on the shoulder. Outer lip stongly thickened, with 11-12 denticles. Colour pattern of black spiral lines, which can be interrupted, and irregular longitudinal flames, on a greyish or whitish background.” The live animal of M. eveleighi is also described and illustrated in GOFAS and FERNANDES (1988, colour plate 1. fig. b.) of specimens from Sao Tome : “Foot translucent, with small yellow spots grading towards the inner areas to larger, paler cream-yellow blotches. Head and tentacles with a series of yellow spots. Siphon densely covered with yellow blotches, with slightly more intense hue than head and foot”. Fig. 10. M. lucani Jousseaume. Banana, Congo. Coll. F. Boyer, 15.70 x 8.90 mm. BOYER, RYALL & WAKEFIELD GOFAS & FERNANDES (1988) noted that there were no appreciable differences between M. eveleighi and M. tyermani on the grounds of shell morphology and colour pattern. They do, however, show concern about the large specimens referred to in BERNARD (1984) as M. eveleighi which is unknown in Sao Tome, and prefer to reserve the name ‘eveleighi for insular specimens “until identity is established by observing living animals from the mainland.” They also note the original habitat of M. eveleighi in Sao Tome as “ usually located near the mouth of small streams, in fine silty sand, loaded with plant debris of terrestrial origin, hand dredged in 1-2 m depth”. - Numerous specimens recorded from Principe Island in FERNANDES & ROLAN (1991, p. 86, fig. 1) as M. eveleighi. These specimens were collected in mud at 8- 10 m, and are said to present no visible difference with the specimens from Sao Tome. Egg capsules of M. eveleighi are noted to be frequently found on the turrid Clavatula coerulea Weinkauff, which lives in the same muddy habitat. One egg capsule is illustrated, containing a single developing embryo presenting the characteristic black axial marks on the first whorl of the shell. Remarks. Curiously, when commenting on the type specimens of M. tyermani in the Marrat collection, TOMLIN (1913) does not compare this species with M. eveleighi, which was published by himself only several months previously. The date of publication for the description of M. eveleighi was 1 January 1913, and his notes on the Marrat Collection were read before the Society on 12 March 1913. TAXONOMY The taxa M. keppeli Sykes and M. eveleighi Tomlin and Shackleford are both to be considered as junior synonyms Of M. tyermani, Marrat. M. tyermani appears as a monospecific group, with no evident geographic or bathymetric polytypism. On the other hand, M. tyermani presents all the features of a polymorphic species, the most variable patterns being known from North Gabon. However, this zone is also the best sampled, so we could assume that the same kind of intrapopulational variability can also occur in other places. For example, the scarce shells from the northern side of the Gulf of Guinea (Liberia to Nigeria) show a noticeable diversity of sizes, proportions and colour backgrounds. We do not find any reason to suspect the existence of several sibling species belonging to a group M. tyermani. All the morphologic and chromatic intergrades are known for the shells, and there is no indication of an ecologic specialisation for any population, as all the specimens confirmed to have been collected alive were found on soft bottoms, in silty sand or mud, often with the presence of organic debris. On the other hand, the occurence of sibling Marginella xicoi, n.sp. APEX 14(2): 43-52, 20 oct. 1999 species remains a possibility, and it is difficult to be sure until their precise habitat and behaviour, their biochemical characters (from chromosomic studies and protein electrophoresis), and the chromatism of the soft parts for each population, have been examined. For now there is no reason to have any doubt about the specific unity of the populations expressing the phena M. tyermani, and we propose to fully consider the taxon M. tyermani as representative of a natural species and to use it as such. OTHER BIOLOGICAL ASPECTS Animal. The animal is 1llustrated by GOFAS & FERNANDES (1988) from the population of Sao Tome Island, as M. eveleighi. It has small yellow blotches on the foot, head, tentacles and siphon, and yellow spots along the edge of the foot. The chromatism of the soft parts is apparently similar in the population of Principe Island (FERNANDES & ROLAN, 1991). The soft parts were not observed in the other populations of M. tyermani. We note that larger yellow blotches are a feature of M. fumigata Gofas and Fernandes, paler ones on M. lucani Jousseaume, and very much smaller ones on the Angolan M. gemmula Bavay, 1913, M. luculenta Gofas & Fernandes, 1994, and M. undulans Gofas & Fernandes, 1994 (all illustrated in GOFAS & FERNANDES, 1994). The radula is unknown. Development. Paucispiral protoconch. The development is direct, as confirmed by the egg capsules pictured in KNUDSEN (1950) and FERNANDES & ROLAN (1991), showing larval shells at an advanced stage of maturation and near to hatching, with the first whorl fully developed. In both cases the egg capsule was rather rounded and arched. Habitat. M. tyermani seems to be strictly dependant on soft, silty to muddy bottoms incorporating a high organic content. This kind of environment is mainly found around river estuaries in shallow depths, and M. tyermani is generally found here from Liberia to Sao Tome. It appears therefore that it can tolerate water of reduced salinity. It can occasionally be found in deeper water, but always still around the mouth of larger rivers (like the Niger Delta) where large quantities of organic silty sediment is carried out to sea. It is however not proven that the species lives below 50 m. The occurrence of M. tyermani in “coral gravel” (TOMLIN & SHACKLEFORD, 1913) is doubtful, and could constitute dead shells being found in beach drift. The occurrence of M. tyermani in “sand” (BERNARD, 1984) has to be corrected to “ silty sand to muddy sand” (BERNARD, personal communication). 49 APEX 14(2): 43-52, 20 oct. 1999 Distribution. Due to its special habitat associated with estuarine areas, the geographical distribution of M. tyermani 1s probably fragmented. For now, the occurrence of M. tyermani 1s recorded from Liberia to Sao Tome Island. The species seems to be absent in Senegal, but has to be looked for in Sierra Leone and Guinea. The species seems also to be absent in Southern Gabon, Congo (P.H. Hattenberger, personal communication) and Angola. In this area, it seems to be replaced in the same habitat and at the same depth by M. lucani Jousseaume, which exhibits similar shell decoration to M. tyermani. According to GOFAS & FERNANDES (1994), M. lucani is found “on muddy bottoms of shallow bays with turbid waters, usually not far from the mouth of small coastal streams, at 5-10 m.” Phyletic relationship. For chromatism of the soft parts and decoration of the shells, M. tyermani seems to be related to several species which occur from Southern Gabon to Northern Angola, on either side of the mouths of the Congo River, namely M. lucani Jousseaume, 1884, M. fumigata Gofas and Fernandes, 1994, and M. marimba Gofas and Fernandes, 1994. (M. fumigata and M. marimba could however have a deeper bathymetric distribution). This relationship needs to be verified more accurately, and the important disparity between shell morphologies of these four species needs to be explained. The group M. musica Hinds (under study by the first author) is apparently represented by four different species in Northwest and South Africa, presents the same kind of shell decoration, and could be joined to a “complex M. tyermani”, even if M. cf. musica (non-M. diadochus A. Adams & Reeve) from South Africa presents a somewhat distinct chromatism of the soft parts, with alternating red and yellow radiating stripes. M. tyermani could also be related to the group M. festiva Kiener, represented in Angola by three species which all bear a pattern of small, densely grouped yellow spots on the soft parts, not so far from the pattern presented by M. tyermani. Further investigations on this group and its phyletic organization are required. NEW SPECIES Marginella xicoi n.sp. Fig. 11 Type material. Holotype (Fig.1l) deposited in Paris Museum (MNHN), adult shell, 7.70 x 4.95 mm; Paratype 1: Coll. F. Boyer, 7.50 x 4.50 mm; Paratype 2: Coll. A. Wakefield, 7.15 x 4.70 mm; Paratypes 3 to 8: Coll. P. Ryall 8.78 x 5.18 mm, 9.01 x 5.54 mm, 8.46 x 5.23 mm, 8.64 x 5.26 mm, 7.84 x 4.57 mm, 7.05 x 3.75 mm 50 Marginella xicoi, n.sp. BOYER, RYALL & WAKEFIELD (juvenile). Paratype 9: Coll. M. Pin, 6.75 x 3.80 mm (juvenile). Type locality. West of estuary of Volta River, Ghana. Discovered in a single dredging by Peter Ryall at 15-20 m. in sand and broken shells, February 1995. Has not been found subsequently in dredgings from the same area. Other material examined. - several crabbed specimens collected in beach drift, in Lome, Togo, by P. Escudie (Dec. 1982). Stored in Coll. P. Geniez. - 2 crabbed specimens from the same origin (ex. Coll. P. Geniez), now in Coll. F. Boyer. 6.90 x 4.70 mm and 6.00 x 3.90 mm (the latter with its protoconch absent). Description. Small, stocky and solid shell of 2.4 whorls. Moderately elevated, blunt spire. Sub-ovate body whorl. Wide, low protoconch and two whorls visible from ventral view. Axial ribs on first and last whorls, just disappearing before the base. Very thick lip bearing fifteen or so labial denticles. White labrum with a heavy margin. Four strong columellar plaits, the first two being shghtly ‘S’ shaped, running parallel and very close. Black and white flammules on a pale greenish-yellow background, forming arrowhead patterns just under the shoulder and at the first anterior quarter of the body whorl. Protoconch light pinkish-orange. Animal and radula unknown. Development direct. protoconch. presumed Paucispiral Distribution. Only known from East and West neighbouring areas of the mouth of the Volta River. However, dredgings of micromolluscs in this region remain scarce, so the range of this species could be made tentatively wider. Discussion. Marginella xicoi n.sp. presents close similarities with M. tyermani, particularly in the pattern of columellar plaits and siphonal canal (Fig. 12: compare with the first two long sigmoidai columellar plaits and truncated base in M. reeveana Petit, 1851), and the same pattern of axial black and white flammules. The principal differences are that M. tyermani presents a more tapering slender outline, less developed labrum and axial ribbing, 3.25 whorls (from the ventral view: protoconch and 3 whorls), a more produced protoconch, and the presence of black spiral lines. The geographic variability of M. xicoi is at present unknown, the species having been studied so far from only two samplings. It is a fact, however, that all the shells present a very homogenous morphology and decoration. A further point to make is that reported BOYER, RYALL & WAKEFIELD Marginella xicoi, n.sp. APEX 14(2): 43-52, 20 oct. 1999 Fig. 11. M. xicoi n. sp. Holotype. 7.70 x 4.95 mm. Fig. 12. Detail of columellar plaits and siphonal canals of M. reeveana, M. xicoi, and M. tyermanii. populations of M. xicoi are both located centrally in the zone of distribution of the polymorphic M. tyermani, and that the distinctive features of M. xicoi (principally the reduced number of whorls and the absence of spiral lines on the pattern of axial flammules) are not found in the range of shell variability of M. tyermani. M. xicoi has therefore to be considered a distinct species, with a close phyletic relationship to M. tyermani. We propose to place M. xicoi in a “complex Marginella tyermani” which could possibly expand with the discovery of new related species in the Gulf of Guinea. Etymology. Named in memory of Francisco Xico Fernandes, from Luanda, Angola. A keen collector and scholar who contributed greatly to the malacological discovery of Angola and West Africa in recent years. Died 19 January 1996. 51 APEX 14(2): 43-52, 20 oct. 1999 ACKNOWLEDGEMENTS. We would like to thank Nora F. McMillan of the Liverpool Museum, and Kathie Way of British Museum of Natural History, London, for helpful advice and support, and for enabling detailed study of important type specimens. We are also indebted to our two referees for their valuable contributions. REFERENCES BERNARD, P.A. 1984, Shells of Gabon. Roma. FERNANDES, F. & E. ROLAN. 1991. The Marginellidae (Mollusca, Gastropoda) of Principe Island (Republica de Sao Tome e Principe). Journal of Conchology 34: 85-90. GoFAS, S. & F. FERNANDES. 1988. The Marginellids of Sao Tome, West Africa. Journal of Conchology 33: 1-30. GoFAS, S. & F. FERNANDES. 1994. The Marginellidae of Angola : the genus Marginella. Journal of Conchology 35: 103-119. KNUDSEN, J. 1950. Egg capsules and development of some marine prosobranchs from tropical West Africa. Atlantide Report 1: 85-130. KNUDSEN, J. 1956. Marine prosobranchs of tropical West Africa (Stenoglossa). Arlantide Report 4: 7-110. MARRAT, F. 1876. Description of five new Marginellidae. The Quarterly Journal of Conchology 1: 136-137. 52 Marginella xicoi, n.sp. BOYER, RYALL & WAKEFIELD MARRAT, F. 1877, Description of new species. The Quarterly Journal of Conchology (11): 204-205. PETIT DE LA SAUSSAYE, S. 1851. Notice sur le genre Marginelle, Marginella Lamarck, suivie d’un catalogue synonymique des especes de ce genre. Journal of Conchology 2(1): 38-59. REEVE, L.A. 1841-1842. Conchologica Systematica, or Complete System of Conchology 1: (1-6) 1-195, pls. 1-129, 1 table (1841) ; 2: 1-337, pls. 130-300 (1842). London. ROLAN, E. & F. FERNANDES. 1997. The small marginelliform gastropods from Ghana (Neogastropoda, Cystiscidae). Zberus 11: 3-12. SOWERBY, G.B. II. 1846. Monograph of the genus Marginella, in Thesaurus Conchyliorum, or figures and descriptions of recent shells. 1-7: 239-406, pls. 68-78. Sowerby, London. SYKES, E.R. 1905. Descriptions of new forms of Marginellidae and Pleurotomidae. Proceedings of the Malacological Society of London 6: 315-318. TOMLIN, JR. le B. 1913. Notes on some types of Marginella in the Marrat collection. Journal of Conchology 14(2): 44-45. TOMLIN, JR. le B. 1917. A systematic list of the Marginellidae. Proceedings of the Malacological Society of London 12(5): 242-306. TOMLIN, JR. Le B. and L.J. SHACKLEFORD. 1913. Descriptions of two new species of Marginella from San Thome Island. Journal of Conchology 14(1): 11. VASSART & TURSCH Oliva truncata APEX 14(2): 53-58, 20 oct. 1999. Taxonomic implications of syntopy: the status of Oliva truncata Marrat, 1867 (Gastropoda, Olividae)(°) André VASSART s/s La Boudeuse Bernard TURSCH (*) Laboratoire de Bio-écologie, Faculté des Sciences, Université Libre de Bruxelles, 50 av. F.D. Roosevelt, B-1050 Brussels, Belgium. (*) btursch@ulb.ac.be (°) Studies on Olividae 32. KEY WORDS. Oliva, truncata, polpasta, taxonomy, nomenclature. ABSTRACT. ©. fruncata Marrat, 1867 has been previously considered to be a junior synonym of ©. polpasta Duclos, 1833. Syntopic populations of these two taxa, easily separable by morphometry, have been found at Cebaco I., Panama. This establishes that these taxa are distinct species. Problems of data interpretation and character validity are discussed. RÉSUMÉ. O. truncata Marrat, 1867 a été considérée comme étant un synonyme junior de O. polpasta Duclos, 1833. Des populations syntopiques de ces deux taxa, aisément séparables par morphométrie, ont été trouvées à l'île de Cebaco, Panama. Ceci établit que ces taxa sont des espèces distinctes. Les problèmes d'interprétation des données et de validité des caractères sont discutés. 1. INTRODUCTION The taxonomic status of the tropical Eastern Pacific taxon ©. truncata Marrat, 1867 (holotype illustrated Plate 1, A) has been quite controversial. It was the Western Pacific ©. elegans Lamarck, 1811 for BURCH & BURCH (1960), ZEIGLER & PORRECA (1969) and WAGNER & ABBOTT (1978). The taxon is not cited in the very thorough Seashells of tropical West America by KEEN (1971). PETUCH & SARGENT (1986) correctly recognised that ©. fruncata belongs to the Panamic Fauna and considered it to be a distinct species, differing from ©. polpasta. The very similar ©. polpasta Duclos, 1833 (Figured syntype illustrated Plate 1, A) is a variable species, as attested by the several names given to its forms: ©. callosa Li, 1930, ©. davisae Durham, 1950, ©. kerstitchi da Motta, 1985, ©. ofssoni Petuch & Sargent, 1986 (the latter could still pose some problem). TURSCH, GREIFENEDER & HUART (1998) failed in their attempts at total morphometrical separation of allopatric samples of the "O. polpasta complex" into smaller, objective subgroups. They were therefore compelled to conclude that "in the absence of valid arguments to the contrary" ©. truncata is a junior synonym of ©. polpasta. This paper reports that such arguments to the contrary have now been found. 2. NEW DATA ON SYNTOPY The Belgian sailing yacht La Boudeuse Was moored at the South-Western tip of Cebaco Island (West Panama) in May 1997, in a wide bay, open to the SW swell. The locality (N 07°29,5'/ W 83°13,4' by GPS) is called Caleta Caïman in sailing chartbooks but this name is not confirmed by locals. Using the dinghy, a short dredge haul was made in 25 m depth, dark muddy sand, yielding several live shells matching the holotype of ©. fruncata and numerous others matching the holotype of ©. polpasta. Specimens of ©. polpasta forma kerstitchi da Motta, 1985 were also present, together with intergrading specimens linking the form kerstitchi to the typical habitus (on this point, see KOCH 1992). The haul also contained ©. kaleontina Duclos, 1835 and numerous live specimens of ©. spicata (Rüding, 1798). The same mixture of ©. polpasta and ©. truncata Was also obtained nearby, in another short dredge haul in 6-8 m, dark muddy sand. 3. MATERIAL AND METHODS 3.1. Measurements. Linear teleoconch measurements were effected with we an electronic calliper (brand ROCHE) with numerical display, giving measurements reproducible to 1/100 mm. The measurements H, L, LW, D, X,R, 53 APEX 14(2): 53-58, 20 oct. 1999, Oliva truncata VASSART & TURSCH MPRO and NW have been defined in TURSCH & GERMAIN (1985). As a quick reminder, their meaning is sketched in the graphs where these measurements are used. 3.2. Specimens measured O. truncata Marrat, 1867. 11 specimens ranging from H 16.62 to 38.07 mm. Seven of the specimens had an intact apex (affording our first opportunity to measure the protoconch characters of ©. truncata). O. polpasta Duclos, 1833. 12 specimens (selected at random amongst 20) ranging from H 26.11 to 35.04 mm, all with intact protoconch. 3.3. Remarks. In each of the two samples, the shells are very homogeneous in aspect and closely match the type material (see Plate 1, A), except that the ©. polpasta from Cebaco I. are slightly more angular in outline. Note: When freshly collected, the shell of ©. polpasta has à distinct greenish tinge which fades with time. The data reported here apply to one single local- ity. The ranges of variation are expected to be larger over the whole distribution ranges. The specimen numbers "V-" refer to the Vassart collection. 4. RESULTS AND INTERPRETATION The shells referable to ©. truncata and to ©. polpasta were easily (and totally) separated in many morpho- metric analyses. Only a few need to be reported here. Dealing with controversial taxa, the observed separa- tions have to be interpreted with special caution. 4.1. Teleoconch features. A total separation on teleoconch characters is given in Figure 1 (scatter diagram of the shape factors X/LW vs. D/L). 4.2. Precautions in interpretation. How do we know that the clustering of points in Figure 1 is objective and reflects biological reality? Any distribution of points on a graph could indeed be grouped in many different arbitrary clusters. In this case, the observed gap is quite large but this could be due to problems in sampling (intermediate specimens missing). An easy visual character for distinguishing the two species was found by Nicolas Vassart (then aged 12): all the non-juvenile specimens of ©. truncata display a conspicuous dark-brown blotch inside the anterior notch (see Plate 1, B). The dark blotch is not so visible on small (<10mm) specimens. This blotch is always present on all the specimens of ©. truncata and absent on all the specimens of ©. polpasta, in the hundreds of specimens seen by one of us (A.V.) in many Other localities, from Ecuador to Costa Rica (when the two species coexist, O. truncata was found to be the more common, except in Cebaco I.). 54 0.52 0.56 0.60 0.64 Figure 1. Teleoconch characters. Scatter diagram of X/LW vs. D/L. Minimum convex polygons. Black circles: ©. polpasta Duclos, 1833. Open squares: O. truncata Marrat, 1867. Additional independent evidence: see text, 8 4.1. 0.56 0.52 20 24 28 32 36 Figure 2. Teleoconch discriminants are unrelated to size. Scatter diagram of D/L vs. H. Minimum convex polygons. Black circles: ©. polpasta Duclos, 1838. Open squares: ©. truncata Marrat, 1867. Additional independent evidence: see text, & 4.1. This feature (amongst others) provided the independent additional evidence" necessary to give credibility (see TURSCH 1998) to the separation in Figure 1. AIl the specimens of one cluster possess the dark blotch, those from the other cluster do not. The same goes for Figures 2, 3 and 4. Further additional evidence is brought by the observation of large differences in protoconch characters (see $ 4.3 and Figure 3). APEX 14(2): 53-58, 20 oct. 1999, Oliva truncata VASSART & TURSCH MPRO (mm) Mec 221 229 3.1 3.3 3.5 3.7 NW Figure 3. Protoconch characters. Scatter diagram of MPRO vs. NW (number of nuclear whoris). Minimum convex polygons. Black circles: ©. polpasta Duclos, 1833. Open squares: ©. truncata Marrat, 1867. Additional independent evidence: see text, $ 4.1. 0.18 0.46 0.50 0.54 0.58 Figure 4. Teleoconch characters. Scatter diagram of (H-L)/H vs. D/H. Minimum convex polygons. Black circles: ©. polpasta Duclos, 1833. Open squares: O. truncata Marrat, 1867. Additional independent evidence: see text, 8 4.1. More precautions must be taken before interpreting the data. First, one has to show that the observed morphometric gap is not due to a size effect. Many Oliva shells have a non-isometric growth (their shape is modified as they grow). Intermediates between the "juvenile" and the “adult” forms can be very rare and easily overlooked (see TURSCH 1997). So the possibility always exists that one is really separating two growth stages of the same species. Figure 2 shows that the difference in the shape factor D/L is not due to size. One should also check that the separation is not due to sexual dimorphism or to a bias in sampling. This 1s the case: sexual dimorphism seems to be negligible in Oliva shells (see TURSCH 1997) and the blind process of dredging ensures against collecting bias. 4.3. Protoconch features. These are easier to interpret, being independent from the shell size. Although the protoconchs do at first glance look quite similar (see Plate 1, C), the two samples differ very much in the size of the first volution of the protoconch (much larger in ©. polpasta) and in the number of nuclear whorls (much larger in ©. fruncata). This is shown in Figure 3. 4.4. Conclusion. AIl the necessary precautions being taken, the demonstration of a total gap in the distribution of characters of syntopic samples establishes the existence of two distinct morphospecies. "Syntopic" is here taken in the restricted sense: “"observable in close proximity, in the same biotope" (see LINCOLN & al. 1982, TURSCH 1995). The scale of sympatry in Oliva species can be much smaller than generally imagined (see TURSCH 1994), so only syntopy ensures that the samples are really sympatric. O. truncata Marrat, 1867 has therefore to be considered as a separate species and removed from the synonymy of ©. polpasta Duclos, 1833. As far as we know, the name ©. fruncata has no junior synonyms. 5. DISCUSSION 5.1. Problems with traditional taxonomic characters. MARRAT (1870-71: 7), in the description of ©. truncata, Wrote: "The rounded form, numerous plaits, short spire and different markings, all serve to distinguish this from ©. polpasta" (bold ours). PETUCH & SARGENT (1986) reported that ©. truncata differs from O. polpasta by "being a more slender, cylindrical shell with a reddish-brown zig- zag color pattern, by having smaller, less prominent subsutural flammules, and by having a higher spire with a larger, protracted, needle-like protoconch" (bold ours). The following analysis illustrates both the necessity of a more accurate descriptive vocabulary and the hazards of isolated traditional taxonomic "characters". To distinguish two taxa, characters with overlap- ping ranges of variation are not operational in a monothetic classification (this is the usual system, in which a unique set of features is both necessary and 55 APEX 14(2): 53-58, 20 oct. 1999. PLATE 1 (opposite page) Oliva truncata VASSART & TURSCH A. The figured syntype of O. polpasta Duclos, 1833 (Muséum National d'Histoire Naturelle, Paris) and the holotype of O. truncata Marrat, 1867 (Merseyside County Museum, Liverpool), compared to material from Cebaco |. Scale bars: 10 mm. B. Features of O. polpasta Duclos, 1833 and ©. truncata Marrat, 1867 (illustrated for specimens V- 2634 and V- 2610, both from Cebaco I.). The much smaller first nuclear whorl and the presence of a dark blotch inside the anterior notch are diagnostical for O. truncata. In apical view, the two species are quite similar. C. Comparison of the protoconchs of ©. polpasta Duclos, 1833 and O. truncata Marrat, 1867. All at same scale. Scale bars: 1 mm. sufficient for membership in a group -by opposition to "polythetic", see SNEATH & SOKAL, 1973: 20). Let us now analyse the discriminant characters given by PETUCH & SARGENT (1986), when applied to the samples from Cebaco I. - MORE SLENDER SHELL. This can be measured by the ratio D/H. One finds for ©. truncata: mean 0.49, & (standard deviation) 0.03, min. 0.43, max. 0.53. For O. polpasta: mean 0.54, ©& 0.02, min. 0.51, max. 0.58. The mean values are indeed different, but the ranges overlap (as can also be seen in Figure 4). It is the body whorl (not the whole shell) that is demonstrably more slender (this can be measured by the ratio D/L, see Figure 1 showing that the ranges do not overlap). - MORE CYLINDRICAL SHELL. This can be measured by the ratio D/R (R is the diameter of the spire at right angle from the apertural plane, not illustrated here). One finds for ©. truncata: mean 2.35, & 0.08, min. 2.23, max. 2.48. For ©. polpasta: mean 2.12, © 0.14, min. 1.89, max. 2.32. The mean values are indeed different, but the ranges overlap. - HIGHER SPIRE. This can be measured by the ratio (H- L)/H. One finds for ©. truncata: mean 0.13, & 0.02, min. 0.09, max. 0.16. For ©. polpasta: mean 0.17, © 0.02, min. 0.14, max. 0.21. The mean values are indeed different, but the ranges overlap (as can also be seen in Figure 4). One will notice that it is ©. polpasta (not ©. truncata) that has the higher spire (the name fruncata of Marrat probably refers to a "shortened”" spire). - REDDISH-BROWN ZIG-ZAG COLOR PATTERN. The chevrons of ©. truncata tend indeed to be reddish, whereas in ©. polpasta they are nearly black. - SMALLER, LESS PROMINENT SUBSUTURAL FLAMMULES. This is not evident to us (see Plate 1, B). Both species have nearly the same ‘“cogwheel pattern” (see TURSCH, GREIFENEDER & HUART 1998) - LARGER, PROTRACTED, NEEDLE-LIKE PROTOCONCH. This is also not evident (at first glance, one would even tend to consider the protoconch of ©. polpasta as being larger, see Plate 1, C). What can be demonstrated is that ©. truncata has nearly one more 56 protoconch whorl and a much smaller first nuclear whorl (see Figure 3 and Plate 1, B). The size of the first nuclear whorl can be appreciated only under strong magnification. 5.2. Comment on species detection. It is remarkable that PETUCH & SARGENT (1986), using à series of non-operational characters, nevertheless reached the correct conclusion that ©. polpasta and ©. truncata are distinct species. This is not at all an isolated case: the traditional description of any Oliva species boiïls down to a list of discriminant characters that, when tested, often (mostly?) turn out to be non-operational. One could thus logically expect most of their species to be wrong, but this 1s clearly not the case. How then could one justify their amazing rate of success ? This certainly demonstrates the acuity of perception of highly experienced malacologists. But is it simply a case of being right for the wrong reasons? Let us consider the graph of Figure 4. A full separation is obtained by combining two characters that are non-operational if taken separately. Scatter diagrams (see TURSCH 1998) can indeed evidence the "special sets of covariance" that are of taxonomic importance (GOULD 1984). It seems likely that experienced malacologists subconsciously integrate their data and somehow manage to detect the “special covariance sets” included in the total Gestalt of the shell. Only some kind of telepathy could communicate this essential phase of work to the reader. The systematic use of scatter diagrams is strongly advocated as a more operational and explicit alternative (see TURSCH, 1998): the existence of different sets of covariance is seen at first glance. 5.3. Size of morphological gaps. The morphological gaps observed here (especially for protoconch characters, see Figure 3) are unusually large for closely related species. This might be a good illustration of the law of "character displacement": Wherever two closely related species come into contact, their characters tend to diverge (see for instance BUTLIN, 1989). APEX 14(2): 53-58, 20 oct. 1999. ©. polpasta ©. trunctata Figured syntype Holotype ©. polpasta ©. trunctata Cebaco I. Cebaco |. ©. polpasta V-2629 ©. trunctata V-2610 Oliva truncata VASSART & TURSCH Plate 1 dark blotch ©. trunctata ©. polpasta Cebaco I. Cebaco I. V-2616 V-2745 ©. polpasta ©. polpasta V-2632 V-2745 ©. trunctata ©. trunctata V-2611 V-2718 57 APEX 14(2): 53-58, 20 oct. 1999. 5.4. Data on syntopy. This case invites a comment on the nature of the contribution that field collectors can make to malacology. Nearly all marine mollusc species are based only upon differences in their external aspect (morphological gaps). A morphological gap between two samples of shells does not necessarily mean that one deals with two species. Differences -even /arge differences- can (and are fully expected to) occur between populations of the same species. Interpreting differences in terms of morphospecies is straightforward only if one compares sympatric samples. But when can we say that two samples are sympatric? Should the localities be separated by less than 100 m? 10 km ? 1000 km ? The range of sympatry (the cruising range) of most molluscs being unknown, syntopic occurrence remains the best proof that two taxa are really sympatric. This crucial problem can mostly be addressed only by reliable first-hand information. The localities reported on most labels, especially for commercial specimens, are not accurate enough (some are deliberately misleading). Most field collectors concentrate their efforts on the search for "rare" species or "new" species. They could obtain at least equally significant results by addressing the question: "what are the taxa that share the same micro- habitaf?". ACKNOWLEDGEMENTS. We thank Dr. Dietmar Greifeneder and Christian Van Osselaer for their comments on the manuscript. B.T. is indebted to the Belgian Fonds National de la Recherche Fondamen- tale et Collective (F.R.F.C.) and to BIOTEC, S.A. for supporting his laboratory. REFERENCES BURCH, J.Q. & R.L. BURCH. 1960. Catalog of recent and fossil Olives. Issue 196. Minutes of the Conchological Club of Southern California. BUTLIN, R. 1989. Reinforcement of premating isolation. Pp. 158-179 in Speciation and its consequences, OTTE, D. & J.A. ENDLER, Eds. Sinauer, USA. 58 Oliva truncata VASSART & TURSCH DUCLOS, P.L. 1833. Oliva polpasta. Mag. Zool., Paris, yr. 3(5), pl. 20 and text (May). GOULD, S.J. 1984. Covariance sets and ordered geographic variation in Cerion from Aruba, Bonaire and Curaçao: a way of studying nonadaptation. Syst. Zool. 33(2): 217-237. KEEN, M. 1971. Seashells of tropical West America. 2nd. Ed. Stanford University Press. KOCH, B. 1992. Panamic puzzles: Oliva kerstitchi - yes, no or maybe? Festivus 24(3): 31-33. LINCOLN, R.J., G.A. BOXSHALL & P.F. CLARK. 1982. A Dictionary of Ecology, Evolution and Systematics. Cambridge University Press, Cambridge, UK. MARRAT, F.P. 1867. On some new species of Oliva, and a new 7rivia. Ann. Mag. Nat. Hist. 3(20): 213- 215: MARRAT, F.P. 1870-71. Oliva Bruguière, 46 pp. In G.B. SOWERBY, Thesaurus Conchyliorum. PETUCH, E.J. & D.M. SARGENT. 1986. Atlas of the living Olive shells of the world. CERF editions, Charlottesville, VA. SNEATH, P.H. & R.R. SOKAL. 1973. Numerical taxonomy. W_.H. Freeman and Co., San Francisco. TURSCH, B. 1994. Studies on Olividae XXI. The scale of sympatry in the genus Oliva. Apex 9(4): 131-142. TURSCH, B. 1995. Micro-endemism, allotopy and taxonomy in the genus Oliva. Abstr. 12th Intl. Malac. Congr., Vigo: 107-109. TURSCH, B. 1997. Non-isometric growth and problems of species delimitation in the genus Oliva. Apex 12(2-3): 93-100. TURSCH, B. 1998. Taxonomic problems in the genus Oliva. Phuket Mar. Biol. Center Special Publication 18(2): 263-284. TURSCH, B. & L. GERMAIN 1985. Studies on Olividae. I. A morphometric approach to the Oliva problem. /ndo-Malayan Zoology 1: 331-352. TURSCH, B., D. GREIFENEDER & D. HUART. 1998. A puzzle of highly multiform species: Oliva fulgurator (Rôding, 1798) and related American species. Apex 13(1-2): 1-61. WAGNER, R.J.L. & R.T. ABBOTT. 1967. Standard Catalog of Shells. 2nd. edition. American Malacologists Inc., Greenville, Delaware. ZEIGLER, R.F. & H.C. PORRECA. 1969. Olive shells of the world. Rochester Polychrome Press, Rochester, N.Y. 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Société Belge de Malacologie association sans but lucratif 20 DECEMBRE 1999 SOMMAIRE Three New Gastropod (Mollusca) Species from the New World Description of Austromitra hayesi n. sp. (Neogastropoda: Muricoidea: Costellariidae) from South Africa Three new costellarid species from Japan, Papua New Guinea and other Indo-Pacific locations (Neogastropoda: Muricoidea: Costellariidae) Review of the Indo-West Pacific species of Haustellum Schumacher, 1817 and comments on Vokesimurex Petuch, 1994 (Gastropoda: Muricidae) with the description of H. bondarevi n.sp. APEX Société Belge de Malacologie à.s.b.l. Editeur responsable: R. Duchamps Av. Mozart, 52, B-1190 Bruxelles Comité d’édition: Dr. T. Backeljau Koninklijk Belgisch Instituut voor Natuurwetenschappen Dr. Y. Finet Muséum d'Histoire Naturelle, Genève M. R. Houart Institut royal des Sciences naturelles de Belgique (collab. scient.) Dr. CI. Massin Institut royal des Sciences naturelles de Belgique Prof B. Tursch Université Libre de Bruxelles Dr. J. Van Goethem Koninklijk Instituut voor Natuurwetenschappen Prof. G. Vauquelin Vrije Universiteit Brussel COTISATIONS MEMBERSHIP Belgique - Belgium Etranger - Foreign Membres résidant en Belgique Abonnement aux revues APEX & ARION (avec le service des bulletins) Subscription to APEX & ARION Membre effectif 5:22. 1200 BEF Membre étudiant =. #5 mme 600 BEF Europe (CEE-EE0) 1600 BEF Autres pays (other countries)... 1800 BEF (sans le service des bulletins) | Versement à effectuer par mandat poste Personne appartenant à la famille international ou par chèque bancaire à un des d'un membre effectif et ayant comptes mentionnés ci-dessous, en francs même rÉSItenEe ne 400 BEF belges uniquement. Payable, by international money order, or by bank check at one of the below mentioned Versement à effectuer à un des comptes accounts, in Belgian Francs only mentionnés ci-dessous, au nom de la Société Belge de Malacologie c/o Mme A. Langleit, au nom de: Av. Cicéron, 27, bte 92, B-1140 Bruxelles at name of: Mme A. Langleit Av. Cicéron, 27, bte 92, B-1140 Brussels, Belgium Comptes bancaires CCP 000-0974225-54 Les articles et textes présentés dans cette revue réflètent l'opinion personnelle de leur(s) auteur(s) et non pas nécessairement celle de la Société ou de l'éditeur responsable. Tous droits de reproduction, de traduction et d'adaptation des articles publiés dans ce bulletin, réservés pour tous pays. All rights of reproduction are reserved without the written permission of the board. Internet: http://www.arkham.be E-mail: cvilvens@prov-liege-be GARCIA Three new species from the New World APEX 14(3-4): 59-65, 20 déc. 1999 Three New Gastropod (Mollusca) Species from the New World Emilio Fabiân GARCIA 115 Oakcrest Dr. Lafayette, LA 70503, U.S.A KEY WORDS. Gastropoda, Muricidae, Columbellidae, Terebridae, new species, New World. ABSTRACT. Three new gastropod species from the New World assigned to the families Muricidae, Columbellidae and Terebridae, are described and compared with their ciosest relatives. Chicoreus (Siratus) vokesorum, n. sp., from the Bahama Islands is compared with C (S.) cailleti (Petit de la Saussaye, 1856) and two species of Vokesimurex Petuch: V. tryoni (Hidalgo in Tryon, 1880) and F. sunderlandi (Petuch, 1987). Columbella marrae, n. sp., from southwestern Panama, is compared with C. sonsonatensis (Môrch, 1860) and C. fuscata Sowerby, 1832; and Terebra mugridgeae, n. sp., from the northern Gulf of Mexico, is compared with T: riosi Bratcher and Cernohorsky, 1985, T. petiveriana Deshayes, 1857, T. concava (Say, 1826), T. protexta (Conrad, 1846), and TJ. vinosa Dall, 1889. INTRODUCTION Long years of interest in western hemisphere mollusks, the access to important molluscan collections, and the opportunity to collect in areas of the ocean difficult to reach, has led the author to the discovery of three unusual new species. Sometime ago the author obtained a single, live collected specimen of an undescribed muricid species that can be assigned to the subgenus Chicoreus (Siratus). It had been dredged in deep water in the Bahama Islands. When a second live collected specimen from the same general area was discovered on a reef wall by means of scuba diving, the author decided to describe the species. JOUSSEAUME (1880) recognized a group of muricids, typified by Murex senegalensis Gmelin, 1791, as the genus Siratus. At that time he did not describe the genus but did so in 1882. Fourteen living species of Siratus are currently known from the western Atlantic. All fourteen have been treated by VOKES (1990), who referred them to the subgenus Chicoreus (Siratus). In November, 1996, the author spent several days on board of the R/V “Pelican”, a research vessel operated by the Louisiana Universities Marine Consortium (LUMCON). While dredging off the Mississippi coast, several specimens of an undescribed species of Terebra, Were collected in anaerobic sediment (black mud) in shallow water. BRACHTER & CERNOHORSKY (1987) have monographed the family Terebridae, describing, figuring, and comparing nineteen living western Atlantic species. Since the publication of that monograph, six new western Atlantic species have been described: ZTerebra biminensis Petuch, 1987; T. lindae Petuch, 1987; T. pacei Petuch, 1987; T. curacaoensis De Jong and Coomans, 1988; T. imitatrix Auffenberg and Lee, 1988; and 7: reticulata Lopes de Simone and Verissimo, 1995. Finally, in January, 1998, a dredging opportunity was afforded the author by an invitation from James Ernest, the well-known Panamanian collector. Dredging in the isolated Islas Secas, Golfo de Chiriqui, southwestern Panama, yielded five specimens of an undescribed species of Columbella, which were found on a sandy bottom covered with live and dead Pinctada mazatlanica (Hanley, 1856) and Hyotissa hyotis (Linnaeus, 1758). ABBREVIATIONS OF REPOSITORY INSTITUTIONS ANSP- The Academy of Natural Sciences, Philadelphia, PA. BMSM- The Baiïley-Matthews Shell Museum, Sanibel, FL. LACM- The Natural History Museum of Los Angeles County, CA. UF- The University of Florida Natural History Museum, Gainsville, FL. SYSTEMATICS Superfamily MURICOIDEA Family Muricidae Rafinesque, 1815 Genus Chicoreus Montfort, 1810 Subgenus Siratus Jousseaume, 1880 59 APEx 14(3-4): 59-65, 20 déc. 1999 Three new species from the New World GARCIA Chicoreus (Siratus) vokesorum, new species (Figures 1-4 ) Description. Shell club-shaped, delicate; last whorl small, globose, length of holotype 32 mm; siphonal canal long, 17 mm in length. Protoconch of 2.5 whorls, last whorl spirally ridged (Fig. 3); 4.5 teleoconch whorls. Spiral ornamentation on first two teleoconch whorls of four strong cords, forming knobs at intersection with sharp axial ribs, spiral cords increasing to about 12 on body whorl, diminishing in strength. Axial ribs strong on first two teleoconch whorls, about 10 in number on the second; with every third rib developing a shoulder spine. On third whorl spinose ribs becoming varices, ornamented with two adapically recurved spines;, other axial sculpture becoming weak, barely appearing as slightly swollen nodes. Last whorl with three well developed varices each with three recurved, adapically projecting spines; shoulder spine longest, and one minor and two major intervarical ridges, producing spirally elongated nodes where crossed by spiral threads. Suture simple, crossed by axial growth lamellae. Aperture ovate. Inner lip adpressed posteriorly, slightly erect on almost entire length; columella smooth. Outer apertural lip weakly erect, crenulated. Siphonal canal long, about 54% of total shell length; open by a very narrow slit, strongly recurved dorsally; ornamented with a single recurved spine on each varix at base of body whorl. Shell white, with a narrow, diffused brownish-red band at shoulder, darker when crossing varices; a similar band at base of last whorl. Radula: See Figure 4 Type material. Holotype (ANSP 400854) length 32.0 mm, width of last whorl 10.5 mm. Paratype (Craig L. Caddigan collection) length 24.0 mm, width of last whorl 9.0 mm. Type locality. Off San Salvador I, central Bahama Islands, taken live at 273 m. Distribution. Off San Salvador and Lee Stocking Islands, central Bahamas, taken live at 46 - 273 m. Etymology. Named for Dr. Emily H. Vokes and Dr. Harold E. Vokes in recognition of their life-long devotion to the study of the Muricidae and Bivalvia respectively and their unselfish will to help amateur conchologists. Discussion. Currently, there are two known specimens of this species. The holotype came from the collection of the late Eugenia Wright, the well-known muricid collector from Arizona. À second specimen was collected on September 13, 1996, by Mr. Craig L. Caddigan of Fort Pierce, Florida, while scuba diving on a recf wall in 46 m off Lee Stocking Island, central Bahamas. Both specimens were collected live. The unique adapically directed spine formation and the ridged protoconch immediately separate this shell from all other western Atlantic species of Chicoreus (Siratus). In general shape it is closest to typical C. (S.) cailleti (Petit de la Saussaye, 1856), less so to the variation kugleri (Clench and Pérez Farfante, 1945). However, C. (S.) cailleti has a smooth protoconch of 1.5 whorls, has a denticulated columellar lip and more numerous whorls with different spine ornamentation. Because of its size and fragility the new species could be confused with Vokesimurex tryoni (Hidalgo in Tryon,1880) and Vokesimurex sunderlandi (Petuch, 1987). However, these latter two species have the straight siphonal canal characteristic of Vokesimurex Petuch, 1994 (type species: Murex messorius Sowerby, 1841) and differ in coloration, sculpture and spine formation. Family Columbellidae Genus Columbella Lamarck, 1799 Columbella marrae, new species (Figures 9-14 ) Description. Shell thick, typically columbelloid; up to 9.6 mm in length. First whorl eroded on only protoconch available, two smooth, bulbous whorls remaining, tan in color. Protoconch of planctotrophic larval type, bearing a deep sinusigeral notch. Teleoconch of five whorls, the first slightly concave and axially ribbed, the next three nearly smooth. Posterior to suture, a thickened band delimited by poorly developed pustules, giving median whorls a concave appearance. Last whorl pear-shaped, smooth. Aperture S-shaped; outer l'p thickened, arched, with about ten denticles; anal canal delineated by thickening of the parietal wall; parietal wall glazed over, glaze thick enough in some specimens to form shield. Columella twisted, bearing 5 to 9 denticles. Shell tan, with irregular nebulous areas either white or pale violet, with freckled tan markings. Outer aperture yellow; pale lavender inside. Figs. 1-4. Chicoreus (Siratus) vokesorum n. sp. Holotype. Figs. 1-2. Ventral and dorsal views (Length: 32.0 mm, Width: 10.5 mm). Fig. 3. Detail of protoconch (Scale bar: 40um). Fig. 4. Radula (Scale bar 30um). Figs. 5-8. Terebra mugridgeae n. sp. Fig. 5. Holotype (Length: 9.7 mm; Width: 2.7 mm). Fig. 6. Protoconch of Holotype (Scale bar: 180um). Fig. 7. Paratype F (Length: 16.1 mm; Width: 4.6 mm). Fig. 8. Paratype E (Length:13.5 mm; Width: 4.2 mm) 60 GARCIA £a PE. D à pr FRE RE verte Three new species from the New World APEx 14(3-4): 59-65, 20 déc. 1999 61 APEX 14(3-4): 59-65, 20 déc. 1999 Three new species from the New World GARCIA Type material. Holotype (ANSP 400855) length 9.6 mm, width 5.7 mm (Figs. 9-12 ). Paratype A (ANSP 400856) length 8.6 mm, width 5.2 mm (Figs.13-14). Paratype B (BMSM 2242) length 9.0 mm, width 5.4 mm. Paratype C (LACM 2870) length 8.0 mm, width 5.2 mm. Paratype D (author’s collection 17018) length 8.2 mm, width 5.3 mm. Type locality. Islas Secas, Golfo de Chiriqui, southwestern Panama. Dredged in 24 to 37 m, on a sand bottom. Distribution. Known only from the type locality. Etymology. Named for Mrs. Lauretta Marr, of Midland, Texas, for her interest in Panamanian shells. Discussion. The relatively small area where the new species was found is characterized by very strong currents, which is probably the cause for having such an abundance of Pinctada mazatlanica and Hyotissa hyotis. These two species seem to be essential to the development of the rich molluscan fauna of the area, for if the dredge did not bring up those two species, very few or no other species were dredged. The area yielded 73 species of gastropods representing 34 families. Of these, the muricids were by far the most numerous with 11 species. The bivalves were represented by 26 species in 15 families. In spite of variations in the nebulous color markings of the species, all five type specimens show them below the suture to a greater or lesser degree. The markings may be present in other parts of the body whorl; however, the general appearance of the species is constant. The small size separates this species from most Panamic species of Columbella. The closest congener is Columbella sonsonatensis (Môrch, 1860) (Figs. 15- 19). However, that species has sharper columellar denticles; is narrower (see Table I); the shell is white, streaked with blackish-brown markings; has a white lip and aperture, with darker markings showing through; and it has four white, semi-translucent protoconch whorls comparatively larger than the new species. Columbella sonsonatensis prefers an intertidal, muddy environment, but C. marrae inhabits deeper water, on a coralline substrate. Len Wim | L7W ù RE LE ES Locality Length (mm) | Width (mm) Rio Mar, Panama Bay : Hicacos Pt., Montijo Bay Arenas de Quebro Es Arenas de Quebro 2 Arenas de Quebro 2 8.0 8. 7 .3 7 7 5 7 7 Arenas de Quebro 2 IE Table1. Chart showing length/width ratio of C. marrae vs. C. sonsonatensis from Panama. LT frs de Que Fig. 9-14. Co/umbella marrae n. sp. Fig. 9. Ventral view of holotype (Length: 9.6 mm; Width: 5.7 mm). Figs. 10-11. Ventral and dorsal view of holotype. Fig. 12. Protoconch and early teleoconch whoris of holotype (Scale bar 165 um). Figs. 13-14. Ventral and dorsal view of paratype A (Length: 8.6 mm; Width: 5.2 mm). Figs. 15-19. Co/umbella sonsonatensis (Môrch, 1860). Fig. 15. Protoconch of a specimen of C. sonsonatensis from Rio Mar, Panama (Scale bar 165 pm). Figs. 16-17. Ventral and dorsal views C. sonsonatensis from Rio Mar, Panama (Length: 8.0 mm; Width: 4.4 mm). Protoconch shown in Fig. 15. Figs. 18-19. Ventral and dorsal views of C. sonsonatensis from Hicaco Point, Veraguas Province, southwestern Panama (Length: 8.2 mm; Width: 4.5 mm). Figs. 20-21. Ventral and dorsal views of Columbella fuscata Sowerby, 1832, Gobernadora Island, Veraguas Province, southwestern Panama (Length: 20.6 mm; Width: 11.4 mm). 62 GARCIA Three new species from the New World APEx 14(3-4): 59-65, 20 déc. 1999 I 63 APEX 14(3-4): 59-65, 20 déc. 1999 Columbella fuscata Sowerby, 1832 (Figs. 20-21), an intertidal species that also prefers a muddy environment (CANTERA ef al, 1979; POORMAN & POORMAN, 1988), is closest in general appearance to C. marrae. However, it is much larger in size (average length 20 mm), is chestnut-brown, dotted and irregularly spotted with white throughout the last whorl, does not have a thickening of the parietal wall at anal canal, does not have a parietal glaze; and has more numerous (about 12) denticles in the outer lip. The aperture of C. fuscata is lavender-colored in fresh specimens, soon fading to white. The aperture of C. marrae 1s yellow. Superfamily CONOIDEA Family Terebridae Genus Terebra Bruguière, 1789 Terebra mugridgeae, new species (Figures 5-8) Description. Shell to 16.1 mm in length (paratype F, Fig. 7), rather thin. Multispiral protoconch, with four whorls rapidly increasing in size; last whorl as wide as first teleoconch whorl. Teleoconch of 11 turreted, slightiy convex whorls, each increasing in width more rapidly than most western Atlantic Zerebra species. First teleoconch whorl with 12 axial ribs; ribs increasing in number to 15 on penultimate whorl of largest specimen. Axial ribs somewhat arcuate, wider than interspaces. Spiral sculpture beginning on second teleoconch whorl, confined to interspaces on first four whorls, crossing axial ribs on later whorls; sculpture of about five to six wide cords on teleoconch whorls, increasing in number on last whorl. Subsutural band inconspicuous, about 8% of last whorl, delimited abapically by a groove; smooth except for elongated nodes formed by axial ribs. Columella recurved, developing two folds in largest specimen. Aperture quadrate. Shell pale tan, with a darker peripheral band and a second, narrower band at base of last whorl. Columella colored purplish in fresh specimens. Type material. Holotype (ANSP 400857) length 9.7 mm, width 2.7 mm (Figs. 5-6). Paratype À (LACM 2871) length 9.5 mm, width 2.6 mm. Paratype B (BMSM 2240) length 7.8 mm, width 2.2 mm. Paratype C (BMSM 2241) length 7.8 mm, width 2.5 mm. Paratype D (author’s collection 16952) length 11.7 mm, width 3.5 mm. Paratype E (ANSP 400858) length 13.6 mm, width 42 mm (Fig. 8). Paratype F (UF 268094) length 16.1 mm, width 46 mm (Fig. 7). Paratype G (UF 268095) length 8.3 mm, width 2 mm. Paratype H (in author’s collection 17918) length 5.9 mm, width 2 mm. 64 Three new species from the New World GARCIA Type locality. Gulf of Mexico, off Mississippi, at 29°53.25°N, 88°40.46° W, in 18 m, dredged in anaerobic sediment (black mud). Distribution. The species is known from the type locality, where the holotype and paratypes A through F were found. Two specimens, paratypes G and H, were dredged at 29°27.32°N, 88°17.32° W, in 56 m, also in anaerobic sediment. Etymology. Named for Ms. Edith Mugridge, a well known resident of Sanibel Island, Florida, and the author’s shelling mentor over 30 years ago. Ms. Mugridge is one of the great benefactors of the Baïiley- Matthews Shell Museum, Sanibel Island. Discussion. Zerebra mugridgeae n. sp. is a common species at the type locality. Presumably it has been overlooked because of its small size and inconspicuous coloring. Terebra mugridgeae is different from other Terebra species in the western Atlantic because of its multispiral protoconch of four whorls. Of the western Atlantic species only 7. petiveriana Deshayes, 1857, and 7. riosi Bratcher and Cernohorsky, 1985, have a planktotrophic type protoconch. However, T. riosi has 3.5 protoconch whorls, is yellowish cream, almost translucent, has à purplish-brown subsutural line, is much narrower, and reaches 10 mm in length. Currently, it is known only from Brazil. Terebra petiveriana has from 3 to 3.5 slender protoconch whorls, has a thick, convex subsutural band, is proportionately narrower, and is a much larger species, reaching 46 mm in length. Currently, this species is known from the northern coast of South America, the West Indies, and the Panamic Province. The multispiral protoconch of the new species, denoting planktotrophic larval development, may indicate a long veliger stage and, therefore, a wide distribution; however, dredgings done west of the delta of the Mississipi River as far as Galveston, Texas, have failed to produce this species in spite of the fact that the same anaerobic sediment of the type locality exists off the Louisiana coast. Of the western Atlantic species, Terebra mugridgeae is closest in general shape to Zerebra concava (Say, 1826), which has a quadrate aperture and the general proportions of the new species. However, T. concava has two protoconch whorls, concave teleoconch whorls, swollen, nodulated spiral cords above and below the suture, and lacks axial sculpture. Terebra vinosa (Dall, 1889) has 1.5 protoconch whorls, is grayish-white, has a swollen subsutural band with 3 to 6 spiral cords, is narrower in shape (the 11th whorl of widest specimen available measures 4.3 mm in width vs. 4.6 for the new species), and has an elongated aperture. Terebra protexta (Conrad, 1846) has 1.5 protoconch whorls; is brown or tan (dull white when faded), has narrower and more numerous axial rbs (17- GARCIA Three new species from the New World APEX 14(3-4): 59-65, 20 déc. 1999 24 on penultimate whorl), is narrower in shape (the 11th whorl of widest specimen available has a width of 3.9 mm), and has a more elongated aperture. The new species was collected with TZe/lidora cristata (Récluz, 1844) , Linga amiantus (Dall, 1901), Corbula contracta Say, 1822, and Cosmioconcha calliglypta (Dall and Simpson, 1901). The last two species were common. ACKNOWLEDGMENTS. My thanks to Dr. Emily Vokes for reading the manuscript and helping with the preparation of the plates, to Mr. James Ernest for his hospitality while visiting Panama and to the anonymous readers who made very helpful suggestions. Dr. Bruce Felgenhauer, of the University of Louisiana at Lafayette, and his graduate assistant, Victor Townsend, prepared the SEM pictures; Kevan and Linda Sunderland, of Sunrise, Florida, suggested the comparison of Chicoreus vokesorum with Vokesimurex sunderlandi, and Dr. Jose Leal provided relevant literature. REFERENCES BRACHTER, T. & W.O. CERNOHORSKY. 1987. Living Terebras of the World. American Malacologists, Inc. Melbourne, Florida. 236 pp. CANTERA K., R. JAIME, A. EFRAIN, R. RUBIO, F. ZAPATA, and E. BUTTKUS. 1979. Taxonomia y distribuciôn de los moluscos litorales de la Isla de Gorgona. /n H. PRAHL, H.; F. GUHL and M. GRÔGL (Eds.) Gorgona: 141-167. Universidad de Los Andes, Bogotä. JOUSSEAUME, F. 1880. Division méthodique de la famille de purpuridés. Le Naturaliste 2(42): 335-336. JOUSSEAUME, F. 1882. Etude des Purpuridae et description d’espéces nouvelles. Revue et Magasin de Zoologie 3(7): 314-348. POORMAN, F. & L. POORMAN. 1988. A report of the molluscan species of the San Carlos rectangle, Sonora, Mexico, collected by Forrest L. and Leroy H. Poorman from December 1953 to December 1983. The Festivus 20 (6): 47-63. VOKES, E. H. 1990. Cenozoic Muricidae of the Western Atlantic Region. Part VIIT- Murex s.s., Haustellum, Chicoreus, and Hexaplex. Tulane Studies in Geology and Paleontology 23(1-3):1-96. 65 Re - e : à et come LT + ” ali € resta sa fr e pad té © nié. Core : rs ébes CEE" te nd tin fsbumit d : Do pr dt LE à sem ed: l ‘ n de és at a 5 Le |: us ft (a ps Cats AMéÉEsrt qe ce : € ù © À came à Je Mæsliy 4 ares lots lat” é + + como & Ar É. rs oc mt CR nù the d be at CE à ARMÉE CREER “ru és ou sin 4 Lu te à a )-be. 19 4 = ph ta Waits É sm. toutes 1e cest Ce VOTE mecs gralladhe sat" l'O fer Mu to Ga … rer pesszis cor sise. nt bein. fe Mis. ‘ cvrrarez oeuf 460 ME L Dal TURNER Austromitra hayesi n. sp. APEX 14(3-4): 67-71, 20 déc. 1999 Description of Austromitra hayesi n. sp. (Neogastropoda: Muricoidea: Costellariidae) from South Africa Hans TURNER CH-6821 Rovio, Switzerland hturner(@tinet.ch KEY WORDS. Costellariidae, Austromitra, new species, South Africa ABSTRACT. Austromitra hayesi is described from a 70 to 120 m deep rocky sea bottom habitat off Algoa Bay and off Bird Island, South Africa. This new species is distinguished from the similar species Austromitra distincta (Thiele, 1925) by flat and stepped whorls with flat ledges on the sutures; from Austromitra maculosa Turner & Simone, 1998 by smaller size and more numerous axial ribs which are articulated by distinct spiral threads and grooves, and from an undescribed smooth Austromitra sp. by an overall rough sculpture. INTRODUCTION The coastal and moderately deep oceanic reaches of South Africa are inhabited by eight known species of Costellaridae: Vexillum (Pusia) patulum (Reeve, 1845) (ranging from the N. W. Cape to the Natal south coast), Austromitra capensis (Reeve, 1845) (from Table Bay to the Natal south coast), À. canaliculata (G. B. Sowerby III, 1900) (Jeffreys Bay to western Transke1), A. bathyraphe (G. B. Sowerby III, 1900) (Jeffreys Bay to East London), 4. kowieensis (G. B. Sowerby III, 1901) (Port Alfred and surroundings), 4. distincta (Thiele, 1925) (in 70 to 300 m depth off Mossel Bay, Algoa Bay and Transkei), 4. rhodarion (Kilburn, 1972) (East London to the Natal south coast) and 4. maculosa Turner & Simone, 1998 (Hout Bay and False Bay). More detailed information on these species is given by THIELE (1925), TURTON (1932), BARNARD (1959), KILBURN (1972), KILBURN & RIPPEY (1982), MARAIS & GRAEVE (1989), TURNER (1993, 1994) and TURNER & SIMONE (1998). From 1995 until March 1999, Mr. Brian HAYES (Port Elizabeth, S. A.) has provided several lots of small gastropod shells for study. These shells were found dead off Algoa Bay (including Bird Island) in cray traps. Each trap had been let down to the rocky sea bottom at depths 70 to 120 m by a single long rope attached to a surface buoy. The traps were used for catching the rock-lobster Panulirus homarus (Linnaeus, 1758) and were usually left at the bottom for 2-5 days at a time. The small gastropod shells, incidentally caught with the cray traps, turned out to belong to the family Costellariidae. Comparison with an extensive Costellarid material (including documentation of types) has proved that 15 specimens out of all lots represent a species new to science which is described in the following pages. ABBREVIATIONS BMNH: Natural History Museum, London NMSA: Natal Museum, Pietermaritzburg SMF: Senckenberg-Museum, Frankfurt/M ZMB: Zoologisches Museum der Humboldt- Universität, Berlin SYSTEMATICS Superfamily MURICOIDEA Rafinesque, 1815 Family Costellariidae Macdonald, 1860 Genus Austromitra Finlay, 1927 Type species by original designation: Columbella rubiginosa Hutton, 1873, Recent, New Zealand. Austromitra hayesi n. sp. Figs. 1-7 Type material. Holotype (Figs. 1-2) 8.3 x 3.3 mm, aperture length 4.2 mm, collected by cray trap off Algoa Bay, about 100 m (70 m to 120 m) depth, 1997; ex collection B. HAYES, Port Elizabeth, South Africa; deposited in NMSA (V7304/T1717). Paratype #1 (Figs. 3—4) 8.4 x 3.3 mm, aperture 4.2 mm, from the type locality, 14 November 1995; deposited in BMNH (#19990443). Paratype #2 (Fig. 5) 8.0 x 3.3 mm, aperture 3.9 mm, from the type locality, March 1995; in coll. H. TURNER. Paratype #3 (7.9 x 3.3 mm, aperture 3.8 mm) from the type locality, March 1995; in coll. B. HAYES (P.O. Box 804, Port Elizabeth, 6000 South Africa). Paratype #4 (5.6 x 2.4 mm, aperture 2.8 mm, juvenile specimen) from off Bird Island, 100 m depth, in cray trap, dead, 23 Nov. 1995; in coll. H. 67 APEX 14(3-4): 67-71, 20 déc. 1999 lURNER. Paratype #5 (Fig. 6) 8.0 x 3.6 mm, aperture 4.0 mm, from the type locality, 1996; deposited in SMF (#319985). Paratype #6 (7.8 x 3.1 mm, aperture 3.8 mm) from the type locality, 1996; donated to Mr. RK. SALISBURY. Paratype #7 (7.7 x 3.5 mm, aperture 3.8 mm, lip broken) from the type locality, 1996; in coll. H. TURNER. Paratype #8 (8.0 x 3.5 mm, aperture 3.8 mm), from the type locality, 1997; in coll. B. HAYES. Paratype #9 (7.4 x 3.0 mm, aperture 3.6 mm, lip broken), from the type locality, 1997; in coll. B. HAYES. Paratype #10 (7.6 x 3.2 mm, aperture 3.7 mm) from the type locality, 1998; in coll. B. HAYES. Paratype #11 (7.0 x 3.2 mm, aperture 3.7 mm, lip broken), from the type locality, 1997; in coll. B. HAYES. Paratype #12 (fig. 7) 6.2 x 2.8 mm, aperture 3.2 mm, juvenile, from the type locality, 1997; in coll. B. HAYES. Other material studied. Two specimens (7.0 x 3.1 mm, juvenile, apex and lip broken; 6.2 x 2.7 mm, Juvenile, bad crack in body whorl, lip broken), from the type locality, 1997; in coll. B. HAYES. Type locality. Off Algoa Bay, South Africa, depth 70 m to 120 m (in crayfish traps). Distribution and habitat. Known only from Algoa Bay (including Bird Island), South Africa. The species inhabits a flat, rocky sea bottom in about 100 m depth and lives probably amongst rocks, sponges and soft corals. It shares its habitat with the rock-lobster Panulirus homarus (Linnaeus, 1758). Etymology. Named in honour of Mr. Brian HAYES (Port Elizabeth, South Africa), a well known conchologist and particularly an expert in South African reef molluscs, in appreciation of his merits for malacology. Description. Shell of medium size for the genus, up to 8-9 mm in length and 3.3-3.6 mm in width, shape fusiform, solid. Protoconch acuminate-involute- paucispiral with 1 1/2 smooth glassy whorls (counted from the origin). Teleoconch with 5 moderately rounded whorls, spire outline likewise moderately convex; sutures form a well developed flat ledge, giving the spire a stepped appearance. Spire whorls are sculptured with rounded, evenly spaced axial ribs which number 16 to 17 on the first whorl and 18 to 19 on the following whorls; on the body whorl less numerous (10 to 14) axials which become indistinct on the anterior half and obsolete towards the aperture. The axial ribs are separated by concave interspaces of about the same width. Spiral sculpture changes noticeably during individual growth; on the first 2 or 3 spire whorls spiral grooves number 8 to 9 and are restricted to the interspaces of the axial ribs while the ribs are smooth; on later spire whorls and on the body whorl spiral sculpture becomes more distinct as the spiral 68 Austromitra hayesi n. sp. TURNER grooves are intersecting the axial ribs, with the result that distinct and regular spiral cords and grooves are evenly developed on the axial ribs as well as in their interspaces. Spiral cords number 10 to 11 on the 4th spire whorl and 23 to 25 on the body whorl where the spirals become somewhat stronger anteriorly and extend almost unchanged onto the siphonal fasciole and the anterior end of the shell. Rostrum slightly recurved to the dorsum, siphonal notch indistinct. Columella with 4 slender folds decreasing in size anteriorly. Aperture moderately wide (width = 31-34 % of length), attenuated at base, not lirate within, posterior aperture angle blunt because of a flat ledge at the suture and a callus pad; inner lip of glazed enamel not elevated from the columella; outer lip thin and crenelated by spiral cords and grooves. Shell overall white to beige. — A peculiar feature of this new species is that about one third of the individuals show healed shell fractures and cracks in various stages of individual growth, even on the very early whorls (e. g. paratype #1). Discussion. This new species is similar to Austromitra distincta (Thiele, 1925) (Figs. 8-9). Both species have white shells of roughly the same size and show a similar spiral sculpture with numerous and distinct spiral threads and grooves on teleoconch whorls including the whole body whorl until its anterior end. Both species inhabit the same habitat in the Algoa Bay; beyond this sympatric occurrence, À. distincta is known, however, from a much larger range (Cape Agulhas to Transkei, in depths from 70 m to about 300 m). In shell morphology, À. distincta differs distinctly by its strongly rounded and shouldered whorls as well as in having fewer and much broader axial ribs. Moreover, À. distincta shows distinct spiral threads and grooves over-riding the axial ribs also on the early spire whorls. The well developed flat ledge at the sutures, giving the spire of 4. hayesi n. sp. a stepped appearance, is missing in À. distincta. A. hayesi n. sp. is superficially similar to a recently described South African species, À. maculosa Turner & Simone, 1998 (Fig. 10) (loc. typ.: Hout Bay; new records also from False Bay). Both species may be easily distinguished as À. maculosa grows to a larger size (14 x 5.5 mm) and shows a sculpture with fewer axial ribs which are not articulated by distinct spiral threads and grooves. Moreover, 4. maculosa differs by its peculiar colour pattern of brown blotches below the suture. À. hayesi n. sp. is superficially similar to a recently described South African species, À. maculosa Turner & Simone, 1998 (Fig. 10) (loc. typ.: Hout Bay; new records also from False Bay). Both species may be easily distinguished as 4. maculosa grows to a larger size (14 x 5.5 mm) and shows a sculpture with fewer axial ribs which are not articulated by distinct spiral threads and grooves. Moreover, 4. maculosa differs by its peculiar colour pattern of brown blotches below the suture. TURNER Austromitra hayesi n. sp. APEX 14(3-4): 67-71, 20 déc. 1999 Figs. 1-7. Austromitra hayesi n. sp. Figs. 1-2. Holotype, NMSA V7304/11717 (8.3 x 3.3 mm), off Algoa Bay, South Africa, about 100 m depth, dead in crayfish trap, 1997. Figs. 3-4. Paratype #1, BMNH 19990443 (8.4 x 3.3 mm), from the type locality, 14 Nov. 1995. Fig. 5. Paratype #2, H. TURNER coll. (8.0 x 3.3 mm), from the type locality, March 1995. Fig. 6. Paratype #5, SMF #319985 (8.0 x 3.6 mm), from the type locality, 1996. Fig. 7. Paratype #12, B. HAYES coll. (6.2 x 2.8 mm, juvenile), from the type locality, 1997. 69 APEX 14(3-4): 67-71, 20 déc. 1999 lustromitra hayesi n. sp TURNER Figs. 8-9. Austromitra distincta (Thiele, 1925). Fig. 8. Lectotype, ZMB (7.3 x 3.2 mm), from Agulhas Bank, 155 m depth ("Valdivia" Stat. 104). Fig. 9. Juvenile specimen, M. MARROW coll. #10699 (6.2 x 2.6 mm), from off Nthlonyane River, Transkei, dredged at 300 m depth. Fig. 10. Austromitra maculosa Turner & Simone, 1998. Holotype, NMSA V4687/T1452 (13.4 x 5.4 mm), from Houtbaai near Kommetjie at 36 m depth, west coast of Cape Peninsula, South Africa. Fig. 11. Austromitra sp. H. TURNER coll. (9.9 x 3.7 mm), from Algoa Bay, South Africa, ex pisce, 25 Oct. 1995. 70 TURNER Austromitra hayesi n. sp. APEX 14(3-4): 67-71, 20 déc. 1999 À. hayesi n. sp. was also compared with an obviously undescribed species from the Algoa Bay, Austromitra sp. (Fig. 11), which was brought to my knowledge recently and is still under study. Both species are similar in size and shape, but differ distinctly as A. hayesi shows a rough sculpture of axial ribs and spiral cords on all teleoconch whorls whereas the undescribed Austromitra sp. has an almost smooth shell with obsolete flat and wide, almost obsolete axial ribs on the spire whorls and with several rounded spiral threads only at the base of the body whorl. ACKNOWLEDGEMENTS. I am greatly indepted to Mr. Brian Hayes (Port Elizabeth, South Africa) who generously gave all specimens of the new species at my disposal and who informed me in detail of the habitat data. REFERENCES BARNARD, K. H. 1959. Contributions to the knowledge of South African marine Mollusca. Part 2. Gastropoda: Prosobranchiata: Rachiglossa. Ann. South African Mus. 45: 1-237. KILBURN, R. N. 1972. Taxonomic notes on South African marine Mollusca (2), with the description of new species and subspecies of Conus, Nassarius, Vexillum and Demoulia. Ann. Natal Mus. 21(2): 391- 437. KILBURN, R. N. & E. RIPPEY. 1982. Sea Shells of Southern Africa. MACMILLAN South Africa, Johannesburg: 249 pp., 46 colour-pls. MARAIS, J. P. & F. GRAEVE. 1989. The genus Austromitra in South Africa. Strandloper no. 219: 1- 3}. THIELE, J. 1925. Gastropoda der Deutschen Tiefsee- Expedition. II. Teil. Wiss. Ergebn. Dtsch. Tiefsee- Exped. "Valdivia", 17(2): 348 pp., 34 Taf. TURNER, H. 1993. Ungewëhnliche und neue Mitroidea aus dem Indopazifik. Teil 2. Club Conch. Infn 25(2): 82-111, 4 Farbtaf. TURNER, H. 1994. Ungewühnliche und neue Mitroidea aus dem Indopazifik. Teil 3. Club Conch. Infn 26(1): 96-111, 2 Farbtaf. TURNER, H. & L. R. L. SIMONE. 1998. Austromitra maculosa, a new species of Costellariidae from South Africa. Arch. Molluskenk. 127(1/2): 93-101. TURTON, W. H. 1932. The Marine Shells of Port Alfred S. Africa. Oxford Univ. Press, London: XVI + 331 pp., 72 pis. 71 TURNER & SALISBURY Three new species of Costellariidae APEX 14(3-4): 73-80, 20 déc. 1999 Three new costellarid species from Japan, Papua New Guinea and other Indo-Pacific locations (Neogastropoda: Muricoidea: Costellariidae) Hans TURNER Î & Richard SALISBURY 2 Î CH-6821 Rovio, Switzerland; hturner(@tinet.ch 2 8807 Craydon Dr., Boise, Idaho 83704 USA; richsali@micron.net KEY WORDS. Costellariidae, Vexillum, Japan, Philippine Islands, Solomon Islands, Papua New Guinea, Hawaï, South Africa ABSTRACT. Vexillum (Costellaria) nodaiï n. sp. (Figs 1-4) is described from Japan, the Philippines, Hawan and South Africa; it is compared to Vexillum (Costellaria) kurodai (Sakurai & Habe, 1964). Vexillum (Costellaria) leforti n. sp. (Figs 8-11) is described from Japan and the Philippines; it is compared to Vexillum (Costellaria) alvinobalani Suduiraut, 1999 and F. (C.) filistriatum (G. B. Sowerby II & III, 1874). Vexillum (Costellaria) beverlyae n. sp. (Figs 15-20) is described from Papua New Guinea, the Philippine and Solomon Islands; it is compared to Vexillum (Pusia) festum (Reeve, 1845), V. (C.) leforti n. sp. and . (C.) alvinobalani Suduiraut, 1999. INTRODUCTION Two new costellarid species have been collected in lobster nets from Wakayama Prefecture, Japan. These same species have also been collected in widely distributed locations across the Indo-Pacific. A third new costellarid species has been collected off Rabaul and Nordup, East New Britain, Papua New Guinea. It has been known to the authors for more than 10 years. Only recently have additional specimens come to our knowledge, partially from Indo-Pacific locations widely apart from the original place of discovery. ABBREVIATIONS ANSP: Academy of Natural Sciences, Philadelphia, PA BMNH: Natural History Museum, London MNAN: Muséum national d'Histoire naturelle, Paris NSMT: National Science Museum, Tokyo ZMA: Zoological Museum, Amsterdam SYSTEMATICS Family COSTELLARIIDAE MacDonald, 1860 Genus Vexillum Rôding, 1798 Subgenus Costellaria Swainson, 1840 Type species by monotypy: Mitra rigida SWAINSON, 1821 = Mitra semifasciata Lamarck, 1811 = Vexillum (Costellaria) semifasciatum (Lamarck). Recent. Indo- Pacific. Vexillum (Costellaria) nodaiï n. sp. Figs 1-4 Mitropifex kurodai (young specimen): Kuroda, Habe & Oyama, 1971: 192, pl. 53, fig. 6 (non Mitropifex kurodai Sakurai & Habe, 1964) Type material. Holotype (Figs 1-2) 30.25 x 9.70 mm (aperture length 12.95 mm) at the type locality taken alive, ex coll. Mr. K. NODA; deposited in ANSP (# 402017). Paratype # 1 (30.61 x 10.88 mm, aperture 14.56 mm) from the type locality; in coll. K. NODA. Paratype # 2 (24.4 x 9.7 mm, aperture 11.5 mm) from Cebu, Philippines; in coll. Mr. J. C. MARTIN (# 2659). Paratype # 3 (23.9 x 9.0 mm, aperture 11.3 mm) from off Balicasag Island, Bohol, Philippines; in coll. Mr. E. G. de SUDUIRAUT (# 1006). Paratype # 4 (Figs 34) (30.64 x 9.86 mm, apeïture 13.04 mm) from off Park Rynie, southern kwaZulu-Natal, South Africa, dredged fresh dead at approximately 150 m depth by G. SMITH, May 1994; in coll. Mr. M. LUSSI. Paratype #5 (24.15 x 9.64 mm, aperture 11.16 mm) from off Balicasag Island, Bohol, Philippines, tangle nets 140 m, May 1998; in coll. Mr. AI DEYNZER. Paratype #6 (11.61 x 4.88 mm, aperture 5.10 mm) from off Pokai Bay, Oahu, Hawaï, dredged by A. Adams, May 19, 1976; in coll. R. Salisbury. Other material studied. A badly broken and dark stained specimen 34.40+ (portion of spire missing) x 11.55 mm, aperture 15.23 mm; in coll. Mr. K. NODA. 73 APEX 14(3-4): 73-80, 20 déc. 1999 lype locality. Off Cape Kirime, Ki peninsula, Wakayama Prefecture, Central Honshu, Japan, in 80- 90 m (alive in lobster nets). Distribution and habitat. Japan, Philippine Islands, Hawan and South Africa. In the Philippines the species was collected dead on sand and broken coral in 80-180 m. In Hawau, the species was collected dead in mud and sand in 110-130 m. In South Africa the species was collected fresh dead on a rubble and sponge substrate in ca. 150 m. Etymology. Named after Mr. Kazutaka Noda, Gobo Town, Wakayama Prefecture, Japan. — A new common Japanese name 1s here designated: Atsumi- fude-gai (Atsumi's Mitre); this name refers to Mr. Noda's daughter Atsumi. Description. Shell large, to approximately 35 mm in length. Elongate-fusiform. Protoconch unknown. Whorls number 8-10; spire outline nearly straight; sutures well defined; early spire whorls sculptured with 12-17 large, rounded, longitudinal ribs, first few spire whorls bear 6 or 7 equally spaced, strong spiral grooves confined to bottom of interspaces; on later whorls, spiral grooves become strong enough to weakly bisect the longitudinal ribs; body whorl with 20-27 large, round longitudinal ribs; ribs on body whorl found in single, paired and even triple sets; 17-19 spiral grooves, weakly bisecting the ribs, body whorl ribs extend to rostrum where they are interrupted by 4 or 5 rows of strong nodulose cords; first of these cords white, very large, occasionally split into two cords, separated from subsequent cords by wide, deep, open trench; remaining 3 or 4 cords brown, large, widely spaced. Columella with 4 very strong folds or teeth. Aperture narrow, length less than half total shell length; interior white, strongly lirate; siphonal notch moderately wide, slightly recurved; outer lip thin, simple and rounded, a few weak crenulations near siphonal notch. Shell dark-brown, with varying number of yellowish-white bands; on each whorl largest yellowish-white band divided by thin dark-brown spiral thread lying in posterior 1/3 of band; body whorl with white pustulose cord emerging just above largest columellar fold. Discussion. This new species is larger than most members of the subgenus Costellaria. It was erroneously illustrated as a young or immature specimen of the Japanese species Mitropifex kurodai Sakurai & Habe, 1964. The two species are roughly the same size and have about as many longitudinal ribs lhree new species of Costellariidae TURNER & SALISBURY (Figs. 5-7). However, the spiral grooves of M. kurodaï are much deeper, wider, and do not bisect the ribs. Adult specimens of M. kurodai have a small but prominent parietal callus near the anal sulcus which is not seen in this new species. The overall colour pattern of M. kurodaiï 1s orange-yellow with three narrow white bands on the body whorl. There is no hint of a spiral thread in any of the white bands. This colour pattern contrasts sharply with the dark-brown and yellowish- white bands of V. nodai n. sp. Like many other Japanese species it is also found in the Philippines in deep water. Remarkably, a live specimen of V. nodai (paratype # 4, Figs 3-4) was collected off Natal, South Africa, extending the species' range across the Indian Ocean. The South African specimen is slightly more slender than those found in Japan and the Philippines. Vexillum (Costellaria) leforti n. sp. Figs 8-11 Type material. Holotype (Figs 8-9) 19.94 x 6.18 mm (aperture length 8.58 mm) at the type locality collected alive, ex coll. Mr. K. NODA; deposited in ANSP (# 402018). Paratype # 1 (19.90 x 6.05 mm, aperture 8.58 mm) from the type locality; in coll. Mr. K. NODA. Paratype # 2 (Figs 10-11) 19.9 x 6.1 mm (aperture 8.5 mm) from off Panglao Island, 140-150 m depth, Bohol, Philippines, collected by native fisherman in tangle net; ex coll. Mr. J. P. LEFORT (#1376); deposited in BMNH (# 19990434). Paratype # 3 (18.7 x 6.2 mm, aperture 8.0 mm) from the Cebu area, Philippines, ex coll. S. MARTIN, 1990; now in ZMA (Moll. 3.99.043). Paratype # 4 (16.6 x 5.7 mm, aperture 7.8 mm) from the Cebu area, Philippines, ex coll. S. MARTIN, 1990; now in ZMA (Moll. 3.99.044). Paratype #S (17.21 x 5.67 mm, aperture 7.61 mm) from off Balicasag Island, Bohol, Philippines; in coll. Mr. E. G. de SUDUIRAUT. Paratype # 6 (16.4 x 5.5 mm, aperture 7.1 mm) from off Balicasag Island, Bohol, Philippines, tangle nets in depth 160 to 180 m, August 1998; in coll. senior author (H. T.). Paratype # 7 (15.8 x 4.9 mm, aperture 6.8 mm) from Bohol, Philippines, deep water, 1986, ex H. FISCHÔDER; in coll. Dr. T. W. BAER. Paratype # 8 (14.0 x 4.5 mm, aperture 6.5 mm) from Bohol Straits, Philippines, deep water; in coll. Mr. M. P. MARROW. Type locality. Off Cape Kirime, Ki peninsula, Wakayama Prefecture, Central Honshu, Japan, in 80 m (in lobster nets). Figs. 1—4. Vexillum (Costellaria) nodaïi n. sp. Figs. 1-2. Holotype, ANSP #402017 (30.25 x 9.70 mm) ex coll. K. NoODA; Japan, Honshu, Wakayama Pref., Kii peninsula, off Cape Kirime, in 80-90 m collected alive. Figs. 3-4. Paratype # 4 (30.64 x 9.86 mm) coll. M. Lussi; South Africa, Southern kwaZulu-Natal, off Park Rynie, in 150 m dredged fresh dead. Figs. 5-7. Mitropifex kurodai Sakurai & Habe, 1964, figured paratype NSMT-Mo 39797 (32.0 x 11.7 mm); Japan, Shikoku, Kochi Pref., off Ashizuri-Misaki, about 200 m depth. 74 APEx 14(3-4): 73-80, 20 déc. 1999 br iii 6 5 EL Le] Le] T "5 Le © = a Q e Ce [e) n © oO o [en n > ce) [=] Ce] O un < E TURNER & SALISBURY APEX 14(3-4): 73-80, 20 déc. 1999 Distribution and habitat. Japan (at the type locality found alhve) and Central Philippines (Cebu and Bohol area, deep water, paratype # 6 collected dead on sand among volcanic stones, in 160-180 m). Etymology. Named after Mr. Jean Paul LEFORT, Maeva, Huahine Island, French Polynesia, who was very generous in providing extensive material of uncommon and new mitriform species to H. T. for studies. A new common Japanese name is here designated: Nanase-fude-gai (Nanase's Mitre); this name refers to Mr. NODA's daughter Nanase. Description. Shell of medium size to 20 mm in length. Fusiform-elongate; spire sharply acuminate. Conoidal protoconch of 3 glassy white whorls, basal diameter 0.6 mm. Teleoconch whorls number 8 or 9, spire whorls shghtly convex in outline, sutures form a narrow flat ledge, early whorls with 12-21 longitudinal ribs; ribs weakly bisected by indistinct subsutural groove, interspaces with 3-8 deep spiral grooves; later spire whorls bisected by wide, shallow grooves, forming nodulose regions along tops of ribs; ribs appear almost vertebrate in shape; body whorl with 25-29 longitudinal ribs, subsutural groove obsolete, bisected by 15-18 deep spiral grooves giving the shell a pustulose appearance. Aperture narrow, strongly lirate within, white with brown band, length less than half total shell length; outer lip thin, finely crenulated along margin, margin outline curves gently, margin straight at anterlor section near siphonal notch; columella with 4 folds; largest folds may have groove along top of tooth. Shell white, early whorls light-brown, the 3rd and 4th whorls decorated with dark-brown spots and dashes, later whorls occasionally spotted dark-brown; body whorl white with large brown band at centre; occasionally small light-brown spot at intersection of spiral grooves and ribs; on some specimens these spots fill several spiral intersections forming a spotted line at the periphery of the shell, or the light-brown spots may be scattered randomly over the upper body whorl. Discussion. This new species could be confused with Vexillum (Costellaria) alvinobalani Suduiraut, 1999 (Figs 12-13). Both species are found in deep water off the Philippines and have white shells with brown bands. However, both species may be easily distinguished since V. leforti is much smaller (adult shells 20 mm in length, whereas V. alvinobalani grows Three new species of Costellariidae TURNER & SALISBURY to a comparatively large size of 35 mm). Furthermore the shell base of . leforti is truncated (not distinctly elongated as in Ÿ. alvinobalani). The colour pattern is also quite different: W. Zeforti does not show a subsutural brown band nor a brown shell base as in F. alvinobalani; in addition, light-brown coloured early whorls, some with dark-brown spots and dashes (a very distinctive character of V. leforti) are not found in F. alvinobalani. V. leforti n. sp. resembles superficially F. filistriatum (G. B. Sowerby IT & IT 1874) (Fig. 14) with which is has been confused by collectors. V. filistriatum 1s indeed similar in size (holotype is 16.7 mm in length) and shows likewise a white shell with a brown subperipheral zone on the body whorl. V. filistriatum differs, however, by a more coarse and clathrate sculpture and by a very distinctive colour pattern with quadrangular brown blotches mainly on the periphery of the whorls. This brown colour pattern, scarcely noticed on the worn and faded holotype (not illustrated), is fairly well developed on fresh specimens (Fig. 14; CERNOHORSKY 1978: pl. 36, fig. 7; PECHAR & al. 1980: pl. 31, figs 13-14). Vexillum (Costellaria) beverlyae n. sp. Figs 15-20 Vexillum (Costellaria) festum (Reeve): PECHAR, PRIOR & PARKINSON (1980): pl. 33, figs 13 & 16 (non Mitra festa Reeve, 1845) Type material. Holotype (Figs 15-16) 31.57 x 9.24 mm (aperture length 14.25 mm), taken alive by B. PARKINSON, night dive, May 1976, 21.00 hours; ex coll. Mr. AI & Mrs. B. DEYNZER, Sanibel FL, USA; deposited in ANSP (# 402019). Paratype # 1 (26.85 x 7.98 mm, aperture 11.43 mm) from Rabaul P.N.G. crabbed, in sand, depth 17 m, Febr. 1984, ex coll. Mrs. A. RICHARDS; now in coll. junior author (R. S.). Paratype # 2 (23.83 x 7.33 mm, aperture 11.25 mm) collection data as paratype # 1; deposited in BMNH (# 19990435). Paratype # 3 (21.83 x 7.27 mm, aperture 9.92 mm) collection data as paratype # 1. Paratype # 4 (25.1 x 8.2 mm, aperture 10.8 mm) from off Nordup P.N.G., ex coll. Mrs. A. RICHARDS, Nov. 1982; now in coll. Mr. J. P. LEFORT. Paratype # 5 (Figs 17-18) 24.0 x 7.5 mm (aperture 10.81 mm) from the Solomon Figs. 8-11. Vexillum (Costellaria) leforti n. sp. Figs. 8-9. Holotype, ANSP #402018 (19.94 x 6.18 mm) ex coll. K. NoODA; Japan, Honshu, Wakayama Pref., Kii peninsula, off Cape Kirime, in 80 m collected alive. Figs. 10-11. Paratype # 2, BMNH #19990434 (19.9 x 6.1 mm) ex coll. J. P. LEFORT; Philippines, Bohol, off Panglao Island. Figs. 12-13. Vexillum (Costellaria) alvinobalani Suduiraut, 1999, holotype, MNHN (31.7 x 18.2 mm); Philippines, Bohol, off S.W. coast of Balicasag Island, in 440-480 m. Fig. 14. Vexillum (Costellaria) filistriatum (G. B. Sowerby, Il & III 1874), specimen 14.6 x 5.2 mm, coll. H. TURNER; Hawaïan Islands, Oahu Bay, Pokay Bay, dredged on sand in 75 m. 76 TURNER & SALISBURY Three new species of Costellariidae APEX 14(3-4): 73-80, 20 déc. 1999 mm APEX 14(3-4): 73-80, 20 déc. 1999 Islands, Honiara - Guadalcanal, might dive, depth 43 m, collected alive by S. YEE, Aug. 1980, in coll. Mr. G. & Mrs. B. COOK. Paratype # 6 (Figs. 19-20, light coloured specimen) 29.02 x 8.41 mm (aperture 11.94 mm) from Punta Engano, Cebu, Philippines, net fisherman; in coll. DEYNZER. Paratype # 7 (24.54 x 8.58 mm, aperture 10.78 mm) from Talvat, Rabaul P.N.G., depth 33 m, in black sand by the wall of a small drop-off; in coll. DEYNZER. Paratype # 8 (28.0 x 9.1 mm, aperture 12.8 mm) from off Nordup P.N.G., ex coll. Mrs. A. RICHARDS, Nov. 1982; now in coll. Mr.s. GORI. Type locality. Off Nordup near Rabaul, East New Britain, Papua New Guinea, in 36 m, in dark volcanic sand on slope at bottom of a coral cliff. Distribution and habitat. Papua New Guinea, Solomon Islands and the Philippine Islands. Live and dead specimens found at scuba depths (17 to 36 m) in dark volcanic sand in Papua New Guinea, alive in muddy sand at 43 m in the Solomons. Etymology. Named after Mrs. Beverly DEYNZER, Sanibel FL, USA. Description. Shell of medium size, to 31 mm in length, elongate-fusiform, acuminate. Protoconch hemi- ellipsoidal in shape, basal diameter 0.55 mm, 2 1/2 glassy white whorls. Teleoconch of adult shell with 10-12 whorls, sutures well defined, slightly stepped. Early whorls ornamented with 12-16 strong, round, evenly spaced, slightly curved axial ribs. Interstices between ribs with 4-7 deep spiral grooves, on early whorls grooves do not bisect ribs, on later whorls few spiral grooves may weakly cross crown of rib. Penultimate whorl ornamented with 18 or 19 strong, moderately curved axial ribs with narrow, smooth crowns; body whorl ornamented with 19-23 strong, moderately curved axial ribs with narrow, smooth crowns. Interstices with 7 or 8 deep, evenly spaced spiral grooves on penultimate whorl, with 18-23 spiral grooves on body whorl; anterior portion of body whorl ornamented with progressively more distinct pustulate spiral cords. Columella acuminate, with 4 slender folds decreasing in size anteriorly. Columellar fasciole with 4-7 oblique, rounded cords, some cords appearing to be a continuation of columellar folds. Aperture with a callosity on parietal side of posterior angle. Outer lip lhree new species of Costellariidae TURNER & SALISBURY thin, simple and smooth, with noticeably undulate shight constriction at anterior portion. 7 or 8 unbroken slender lirations in interior of aperture, not extended into siphonal canal; latter narrow and relatively straight-sided, slightly recurved towards dorsum. Shell white, spire in all specimens examined totally white; body whorl white with thin, golden line at shell periphery; golden line faded and extremely hard to see in beach-worn shells; with a broad yellow-cream, golden or reddish-brown band on lower body whorl, just posterior to white (or light yellow) columellar fasciole; columella white. Discussion. This new costellarid has been confused with Vexillum (Pusia) festum (Reeve, 1845) (Figs. 21- 22, lectotype). It differs from that species in being much larger (adult shells 31 mm in length versus 15 mm), more slender and acuminate in shape, whorls less shouldered at the sutures, with uniform spiral grooves between close-set, evenly spaced and more slender axial ribs. Moreover, F. beverlyae n. sp. differs from Y. festum by the more basal position of the brown band on the body whorl. The band of this new species occasionally being light golden-red (Figs. 19-20), rather than brown. V. beverlyae n. sp. could also be confused with Vexillum leforti n. sp. (described above, Figs 8-11), but may be distinguished because the whorls of F. beverlyae are less rounded and are separated by only shallow sutures, the apex and early whorls are not brown but white, the brown band on the body whorl is not so narrow and sharply delimited to a zone at the posterior aperture angle, but extending much farther towards the shell base. V. beverlyae n. sp. resembles to some degree also the recently described Vexillum alvinobalani Suduiraut, 1999 (Figs 12-13). It differs from that species mainly by a less coarse sculpture, spiral cords not over-riding the axial ribs, and by the colour pattern lacking broad brown sub-sutural and peripheral bands. ACKNOWLEDGEMENTS. For the loan of types and important voucher specimens we would like to thank Mrs. Joan PICKERING & Ms. Kathie WAY (Natural History Museum London), Dr. Hiroshi SAITO (National Science Museum Tokyo) and Mr. Robert MOOLENBEEK (Zoological Museum Amsterdam). We are also indepted to the following persons who have kindly supplied specimens and information: Dr. Teddy William BAER (La Croix, Switzerland), Mr. Paul Figs. 15-20. Vexillum (Costellaria) beverlyae n. sp. Figs. 15-16. Holotype, ANSP #402019 (31.57 x 9.24 mm); Papua New Guinea, East New Britain, off Nordup, 36 m depth, collected alive by B. PARKINSON, night dive, May 1976, 21.00 hours. Figs. 17-18. Paratype #5 (24.0 x 7.5 mm) coll. G. & B. Cook; Solomon Islands, Honiara — Guadalcanal, depth 43 m, collected alive by S. YEE, night dive, Aug. 1980. Figs. 19-20. Paratype # 6 (29.02 x 8.41 mm, light coloured specimen) coll. DEYNZER, Philippines, Cebu, Punta Engano, net fisherman. Figs. 21-22. Vexillum (Pusia) festum (Reeve, 1845). Lectotype of Mitra festa Reeve, BMNH #1967756 (11.0 x 4.6 mm); Philippines, Island of Mindoro, Puerto Galero. 78 TURNER & SALISBURY Three new species of Costellariidae APEX 14(3-4): 73-80, 20 déc. 1999 79 APEX 14(3-4): 73-80, 20 déc. 1999 CALLOMON (Osaka, Japan), Mrs. Bunnie COOK & Mr. George COOK (Honolulu HI), Mrs. Beverly DEYNZER & Mr. Al DEYNZER (Sanibel FL), Mr. Sandro GORI(Livorno, Italy), Mr. Jean Paul LEFORT (Huahine, French Polynesia), Mr. Markus LUSSI (Durban, South Africa), Mr. Maxwell P. MARROW (Hampton, Victoria, Australia), Mr. Jean Claude MARTIN (Saint Denis, Réunion), Mr. Kazutaka NODA (Gobo Town, Japan) and Mr. Emmanuel Guillot de SUDUIRAUT (Lapu Lapu City, Cebu, Philippines). Our special thanks are due to Dr. Gary ROSENBERG (Academy of Natural Sciences, Philadelphia, PA) who was instrumental in obtaining material of new species from Japan and helped on early versions of the V. nodaiï description. 80 Three new species of Costellariidae TURNER & SALISBURY REFERENCES CERNOHORSKY, W. O. 1978. Tropical Pacific Marine Shells. Pacific Publications (Aust.) Pty Ltd, Sydney: 352 pp., 17 text figures, 68 plates. KURODA, T., T. HABE & K. OYAMA. 1971. The Sea Shells of Sagami Bay, Collected by His Majesty The Emperor of Japan. Maurzen Co., Ltd., Tokyo: 489 pp., 121 plates. [Note: New species in Mitridae section described by KURODA & HABE.] PECHAR, P., C. PRIOR & B. PARKINSON. (1980). Mitre Shells from the Pacific and Indian Oceans. Robert Brown & Ass. Pty. Ltd., Bathurst N.S.W.: unpaginated, 56 plates. SAKURAI, K. & T. HABE. 1964. Description of two new Vexilliid species dedicated to Dr. T. Kuroda's 77th birthday. Venus 23(1): 29-33. HOUART Indo-West Pacific species of Haustellum APEx 14(3-4): 81-107, 20 déc. 1999 Review of the Indo-West Pacific species of Haustellum Schumacher, 1817 and comments on Vokesimurex Petuch, 1994 (Gastropoda: Muricidae) with the description of H. bondarevi n.sp. Roland HOUART Research Associate Institut royal des Sciences naturelles de Belgique Rue Vautier, 29, 1000 Bruxelles roland.houart@skynet.be KEY WORDS. Gastropoda, Muricidae, Haustellum, Vokesimurex, Indo-West Pacific. ABSTRACT. Eight Recent species and two subspecies are included in Æaustellum. The species are described and discussed. À new species, Haustellum bondarevi n. sp. is described from Saya de Malha Bank, Western Indian Ocean. Fourteen species and three subspecies from the Indo-West Pacific are included in V’okesimurex Petuch, 1994. RESUME. Huit espèces et deux sous-espèces récentes sont incluses dans le genre Haustellum Schumacher, 1817. Les espèces sont décrites, commentées et comparées. Haustellum bondarevi n. sp. est décrit du Banc Saya de Malha, Océan Indien Occidental. Quatorze espèces et trois sous-espèces actuelles de l’Indo-Ouest Pacifique sont incluses dans Vokesimurex Petuch, 1994. INTRODUCTION Since the revision of Murex s.s. and Haustellum (PONDER & VOKES, 1988), I examined many Recent specimens belonging to Æaustellum, and analyzed recent literature (VOKES, 1990, PETUCH, 1994, PARTH, 1995). In a previous paper (HOUART, 1990), the genus Haustellum (sensu PONDER & VOKES, 1988) was considered to be divided into two groups: the group of Haustellum haustellum (Linnaeus, 1758), characterized by a globose, and spineless last teleoconch whorl, rounded varices, large, roundly-ovate aperture, without labral tooth, with a raised peristome and projecting inner lip, and a long, slender, often spineless, or almost spineless, siphonal canal ; and another group including species without labral tooth but more similar in appearance to species of Murex ss. or Siratus Jousseaume, 1880. Haustellum, and particularly H. haustellum has been studied and discussed by several authors (VOKES, 1971, 1990, FAIR, 1976, RADWIN & D'ATTILIO, 1976, KOSUGE, 1980, PONDER & VOKES, 1988, HOUART, 1990, HOUART, 1993, PARTH, 1995). PARTH (1995) considered all the various populations scattered throughout the Indo-Pacific parts of just one species" (1.e. Haustellum haustellum). He mentioned the work of PONDER & VOKkES (1988), who considered the various populations to be a form of a single species, and HOUART (1990, 1993) who splits Æ. haustellum into no less than seven species (7. haustellum, H. longicaudum, H. fallax, H. kurodai, H. vicdani, H. barbieri, and FH. langleitae). The problem will perhaps not be definitively resolved, but at least I hope to show that more than one species is involved in this group of gastropods. MATERIAL Hundreds of specimens have been examined throughout many years from BMNH, IRSNB, MNAN, private collections, and the author's collection. No extensive list of material is given. RESULTS Genus : Haustellum Schumacher, 1817 L There is no known case of poecilogony (different larval development in a same species) in Prosobranchia, so that a single species will not have shells with a multispiral (planktotrophic) protoconch together with specimens with a paucispiral (non- planktotrophic) protoconch (BOUCHET, 1989; HOAGLAND & ROBERTSON, 1988). A planktotrophic protoconch is characterized by the presence of a protoconch I, with a small diameter, followed by a protoconch II, ending with a sinusigeral notch (terminal varix of sinusigera type). A non-planktotrophic protoconch is characterized by the absence of protoconch l/protoconch II discontinuity, and by the larger diameter of the first protoconch whorl, than in planktotrophic species. 81 APExX 14(3-4) 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART As a first conclusion, Haustellum haustellum (Figs 23-26) may be separated from any other species of Haustellum. IL The microsculpture of the protoconch whorls is another valuable tool for species separation [many personal observations, P. Middelfart (in litt.);, SABELLI & TOMMASINI, 1982], so that, once more, #. haustellum may be separated from any other species of Haustellum (Fig. 26). The protoconch of X. longicaudum is also different from the other species by its particular microsculpture, consisting of numerous, small pustules (Fig. 43), which are smaller than in 77. haustellum, and by its broad, irregularly shaped, protoconch. One of the examined juvenile specimens of A. langleitae has very few, sporadic, small pustules on the last whorl, close to the terminal varix; all other examined specimens of #. langleitae have smooth protoconchs. To my knowledge, the microsculpture of the protoconch of Haustellum has never been observed in any other species of Haustellum. However, microsculpture has been observed in Vokesimurex (BANDEL, 1975). IIL Three species have a protoconch with the characteristic morphology of intracapsular metamorphosis (few whorls, broad, irregular shape): FH. longicaudum, H. fallax, and H. wilsoni. However, the protoconch of #. longicaudum has a keeled last whorl, ornamented with microsculpture, while those of FH. fallax and H. wilsoni are smooth. Species with intracapsular larval development have a restricted geographical distribution. F. longicaudum is indeed restricted to the southern part of the Red Sea and adjacent localities, H. fallax seems to live only from Durban, South Africa, to southern Mozambique, and #. wilsoni is known from a small area off West Australia. After having separated A. haustellum, H. longicaudum, H. fallax, and H. wilsoni, we still have a group consisting of Æ. barbieri, H. franchi, H. langleitae, H. kurodai, and H. vicdani. H. franchii and H. barbieri can be clearly separated from the other species by their teleoconch characters (see description and Table 1). #. langleitae, H. kurodai, and Æ. vicdani are somewhat similar, although having different teleoconch characters. The protoconch morphology of H. vicdani is not known, but both FX. langleitae and H. kurodai have a rounded protoconch of 2-2.5 whorls. These three taxa appear to have a different geographical distribution (Fig. 14). At first sight the three taxa differ in teleoconch morphology but it would be useful to observe specimens (if any) from adjacent and intermediate localities to fill the gaps between the different geographical distributions. 82 - If these forms are sympatric, then they may be considered at a specific level. - If each form is isolated, then two solutions are conceivable: * There are three different species, clearly separated by natural barriers and teleoconch characters. * There is one species with two morphologically divergent local populations (subspecies). Currently, having these forms geographically separated as three distinct populations (Fig. 14), I will use trinominal nomenclature. List of (sub)species: Æ. barbieri Houart, 1993 H. bondarevi n. sp. H. fallax (Smith, 1901) H. franchii Bozzetti, 1993 H. haustellum (Linnaeus, 1758) H. kurodai (Shikama, 1964) H. k. vicdani Kosuge, 1980 H. k. langleitae Houart, 1993 H. longicaudum (Baker, 1891) H. wilsoni D'Attilio & Old, 1971 kurodai Genus : Vokesimurex Petuch, 1994 PETUCH (1994) introduced Vokesimurex, for the American long-canalled "Murex" species without labral tooth like Murex messorius Sowerby, 1841, included in Haustellum by PONDER & VOKES (1988). He observed that Vokesimurex lived together with true Haustellum species in the Pliocene. Haustellum s.s. became extinct in the Americas by the beginning of the Pleistocene, but survives in the Indo-West Pacific. Vokesimurex appears to be a younger group than Haustellum s.s., and is known in the western Atlantic and the Eastern Pacific. These considerations, personal observations, and clear similarities in shell and radula morphology (Figs 1-2, 3-8) between the West Atlantic and East Pacific Vokesimurex and Indo-West Pacific Murex-like species Without labral tooth, have led me to consider the following Indo-West Pacific species as belonging to Vokesimurex : V. bobyini (Kosuge, 1983) V. dentifer dentifer (Watson, 1883) V. dentifer coriolis (Houart, 1990) V. danilai (Houart, 1992) V. dolichourus (Ponder & Vokes, 1988) V. gallinago gallinago (Sowerby, 1903) V. gallinago fernandesi (Houart, 1990) HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 V. hirasei (Dautzenberg in Hirase, 1915) V. kiiensis (Kira, 1959) V. malabaricus (E. A. Smith, 1894) V. mindanaoensis (Sowerby, 1841) V. multiplicatus multiplicatus (Sowerby, 1895) V. multiplicatus bantamensis (Martin, 1895) V. purdyae (Radwin & D'Attilio, 1976) V. rectirostris (Sowerby, 1841b) V. sobrinus (A. Adams, 1863) V. tweedianus (Macpherson, 1962) As stated in PONDER & VOKES (1998: 13), this group of species also have similarities with Siratus Jousseaume, 1880. They suggest that the group must have evolved from Siratus well before the appearance of V. messorius because V. messorius has a paucispiral protoconch whereas some Recent Indo-Pacific species have a multispiral protoconch. However, some Indo- Pacific species, ie V. dentifer, Vrectirostris, V. sobrinus, V. tweedianus, and others, also have a paucispiral protoconch of 1.5 — 1.75 whorls as in the West Atlantic species. Radula. The radula of both Haustellum and Vokesimurex are typical muricine, consisting of a sickle shaped, unicuspid lateral tooth on each side, and of a rachidian tooth with a long central cusp, small lateral denticles, and long lateral cusps. No intermediate denticles were ever observed in any radula studied. Figs. 1-2. (Rem: All specimens in the figures are in the coll. R. Houart, unless stated). Fig. 1. Vokesimurex olssoni (Vokes, 1967). Oregon, stn 4896, holotype USNM 67704, 52 mm. Shell whitened. Photo courtesy E.H. Vokes. Fig. 2. V. dolichourus (Ponder & Vokes, 1988). South Africa, Zululand, NM E4416, 67.1 mm. 83 APEX 14(3-4} 81-107, 20 déc. 1999 Character Protoconch (see Figs 23- 26, 42-48, 54) Number of spiral threads on first teleoconch whorl Last teleoconch whorls Siphonal canal Protoconch (see Figs 23- 26, 42-48, 54) Number of spiral threads | 3-4 on first teleoconch whorl Last teleoconch whorls H. haustellum Conical, keeled, with 2.25- 2.75 whorls. Last whorl with micro sculpture. Terminal vanx of sinusi- gera type. Very regular in shape. Planktotrophic larval development. Broad, rounded, with 3 or 4 axial nodules. Spiral sculpture of numerous, regular threads. Aperture pink. Usually without spines, rarely with small, short spinelets at the base. H. kurodai kurodai Rounded, with 2-2.25 smooth whorls. Terminal varix high. Similar to H. fallax but relatively smal- ler. Broad, angulate, with 3 or 4 narroW axial nodules. Spiral sculpture of nume- rous, shallow, low threads, fewer than in FH. haustellum, langleitae and longicaudum. Aperture white or light pink. Indo-West Pacific species of Haustellum H. longicaudum Irregular, broad, 2-2.5 whorls. Last whorl keeled, with micro sculpture. Strong, high carina. The form and breadth of the protoconch is typical for intracapsular development Broad, rounded, 3 or 4 low axial nodules. Spiral sculpture of numerous, shallow threads. Aperture white. H. kurodui langleitae Cylindnical or rounded, With 2-2.5 smooth whorls. Broad, rounded or angu- late, mith 2-4, occasionally broad, high axial nodules. Spiral sculpture of numerous, shallow, low, irregular threads. Aperture yellow or pink. HOUART Rounded, with 1.5-2.25 broad, smooth whorls. No micro sculpture. Twice the size of the protoconch of H. haustellum. Most probably intracapsular larval development, relative to the size, and the form of the protoconch. Terminal varix high, broad. Broad, angulate, with 2 or 3 high, broad axial nodules. Spiral sculpture of numerous, shallow, low threads (fewer than in A. haustellum, H. langleitae and H. longicaudum). Aperture white. With short, weakly adapically curved spines on the base. H. kuroduai vicdani Examined protoconch partly broken. Whorls rounded, smooth. Apparently no micro sculpture. Broad, angulate or rounded, with 3 or 4 axial nodules. Spiral sculpture of numerous, shallow threads, more apparent than in kurodai or fallax. Aperture light orange or yellowish. Siphonal canal Adapically curved small Smooth. Smooth. spines on the base. Table 1. Comparison of four species of Haustellum. SYSTEMATICS Family MURICIDAE Rafinesque, 1815 Subfamily Muricinae Rafinesque, 1815 Genus Haustellum Schumacher, 1817 Type species (by tautonomy) AMurex haustellum Linnaeus, 1758, Recent, Indo-West Pacific. Brontes Montfort, 1810 (not Fabricius, 1801) Brontesia Reidenbach, 1828 (new name for Brontes Montfort, 1810) Haustellaria Swainson, 1833 84 Distribution. Throughout the tropical and subtropical Indo-West Pacific. Description. Shell medium-sized to relatively large, up to 165 mm in length. Last teleoconch whorl globose, rounded. Axial sculpture of last teleoconch whorl consisting of 3, usually high, rounded, spineless varices. Aperture rounded, large. Outer lip crenulate, erect, without labral tooth. Columellar lip strongly projecting, flaring, weakly adherent adapically, otherwise erect. COLLECTION Guido T. POPPE () For more than 20 years, Guido T. Poppe and his friends crossed the world in search for the most beautiful specimens. 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Haustellum haustellum (Linnaeus, 1758) (New Caledonia). 85 APEX 14(3-4} 81-107, 20 déc. 1999 Siphonal canal long to very long (52-73 % of total shell length), straight, narrowly open, smooth or occasionally with 1 or 2 small spines adaperturally. Haustellum barbieri Houart, 1993 Figs 9, 40-41, 55 Haustellum barbieri Houart, 1993: 147, figs 6-9. Protoconchs examined: Off Madagascar (3 specimens) (private collections). Distribution. The species is known in the vicinity of the type locality: Sainte-Marie (Nosy-Boraha), Madagascar, 30-35 m. Description. Shell up to 90 mm in length with 7 teleoconch whorls; protoconch smooth, consisting of 1 7 whorls. Axial sculpture of last teleoconch whorl with 3 low, rounded, nodose, spineless varices. Other axial sculpture of 4 nodose ribs and numerous narrow, nodose, irregular threads. Spiral sculpture of 8 low, tuberculate cords and numerous threads. Aperture broad, rounded. Outer lip weakly crenulate, erect, smooth within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, spineless, straight, open. Pinkish-brown with darker blotches on spiral cords and lighter coloured axial threads. Aperture glossy white. Remarks. Haustellum barbieri differs markedly from the other species of the genus by its colour and sculpture. Ali specimens known to date come from Madagascar. Haustellum bondarevi n.sp. Figs 10, 49-51 Material Examined. Western Indian Ocean, Saya de Malha Bank, approximately 10°30'S, 60°00'E, holotype MNHN (79 mm) and paratype coll. R. Houart (60 mm), NE part of lagoon, 40-45 m, paratype coll. I. Bondarev L (101.2 mm). ! Fadeev Str., 21-B, f1. 17, Sevastopol 335038, Crimea, Ukraine 86 Indo-West Pacific species of Haustellum HOUART Protoconchs examined: Paratype I. Bondarev (partially broken); paratype R. Houart. Distribution. Indian Ocean, Saya de Malha Bank, 40-45 m. Description. Shell medium sized for the genus, up to 101.2 mm in length at maturity, heavy, nodose. Spire high with 2.25 protoconch whorls and 7 broad, rounded, nodose teleoconch whorls. Suture impressed. Protoconch small, whorls rounded (weakly eroded in examined specimen); terminal varix high, weakly convex. Axial sculpture of teleoconch whorls consisting of rounded, nodose, broad ribs on first to third whorl: 11 ribs on first whorl, 12 on second, 13 on third, fourth and fifth whorl with 3 strongly abaperturally excavated varices and 3 intervarical ribs; sixth and seventh whorl with three varices and 4 or 5 intervarical ribs. Varices more prominent at the intersection of spiral cords and threads, giving a strongly nodose sculpture. Spiral sculpture of low, narrow, nodose cords, and threads: 3 cords and one shoulder thread from first to third whorl, 3 cords and 2 shoulder threads on fourth whorl, 3 cords with 1 thread between each pair, and 2 shoulder threads on fifth and sixth whorl, last whorl with 5 cords, 2-3 threads between each pair, and 2 shoulder threads. Aperture large, roundly-ovate; columellar lip narrow, strongly flaring, smooth with small parietal tooth at adapical end; lip strongly erect, adherent at adapical small portion, anal notch narrow, constricted, deep; outer lip erect, crenulate, with 12 or 13 weak lirae within. Siphonal canal long, narrow, straight, narrowly open, with 7 or 8 strong spiral cords adapically, almost smooth abapically. Protoconch and first whorls creamy-white or light brown, other whorls white. Last whorl with three light orange or light brown bands, more apparent on varices. Single orange band on previous whorls. Light orange or light brown tinge on spiral cords of siphonal canal, and on columellar and outer apertural edges. Operculum and radula unknown. Remarks. H. bondarevi differs from X. barbieri in having a more rounded last teleoconch whorl, a smaller aperture, narrower varices and axial ridges, a narrower siphonal canal, in lacking the wrinkled micro-sculpture, and in having a narrow, constricted anal notch relatively to the broad notch in Æ. barbieri. From H. longicaudum, H. bondarevi differs in its more nodose sculpture, narrower siphonal canal, more numerous axial intervarical ridges (4 or 5 vs 3 or 4) and deep narrow, constricted anal notch, relatively to the more shallow, broad, bell-shaped notch in . longicaudum. From H. franchii it differs in having a higher spire, more numerous spiral cords on spire whorls, a more nodose shell, a constricted anal notch, and a different colour, HOUART Indo-West Pacific species of Haustellum APEx 14(3-4): 81-107, 20 déc. 1999 15 30 45 60 75 90 105 120 135 150 165 Fig. 9. Distribution of Haustellum barbieri. LEE “Fier Fig. 10. Distribution of Haustellum bondarevi 180 165 150 87 APEX 14(3-4} 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART DR H. franchit having particular reddish-brown spiral threads, and a pure white aperture Other Mollusca with non-planktotrophic larval development are apparently endemic to Saya de Malha Bank (BOUCHET & BAIL, 1991, OKUTANI, 1991, HOUART, 1992) Etymology. Named after Igor Bondarev, Sevastopol, Ukraine, who discovered, and donated the type material. Haustellum fallax (Smith, 1901) Figs 11, 20-22, 54 Murex fallax Smith, 1901: 113, pl. 1, fig. 9. Protoconchs examined: S. Africa, off Natal (1 sp.); Mozambique, off Quissico (1 sp.); Mozambique, Zavora Pt (1 sp.) (Coll. R. Houart), Mozambique (2 sp.) (private coll.). Distribution. Durban, South Africa (type locality) to Zavora Point, Mozambique, 40-175 m. Description. Shell up to 95 mm in length with 1.5-2.25 protoconch whorls and 6 teleoconch whorls. Protoconch globose, smooth. Whorls rounded. Axial sculpture of last teleoconch whorl consisting of 3 moderately high, rounded, spineless varices. Other axial sculpture of 3 high, strong ribs. Spiral sculpture of numerous, weak, smooth, shallow threads. Aperture broad, rounded; outer lip weakly crenulate, smooth within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, with a single, acute, short spine adaperturally. Light tan to light brown with some darker coloured blotches. Aperture glossy white. Remarks. Haustellum fallax was considered a subspecies of Æ. haustellum by PONDER & VOKES (1988). It is rarely seen in collections but it is much appreciated due to its scarcity and beauty. H. fallax is one of the three Haustellum species with probable intracapsular metamorphosis, but the teleoconch characters are almost invariable, and obviously different from the two other species, . longicaudum and H. wilsoni (see also Table 1). 88 Haustellum franchii Bozzetti, 1993 Figs 12, 52-53 Haustellum franchii Bozzetti, 1993: 107, figs 1, 2. Protoconchs examined: paratype (coll. R. Houart), and original description. Distribution. Ras Hafun, Somalia, 200-250 m. Description. Shell up to 67 mm in length, with 1.5-2 protoconch whorls and 6 teleoconch whorls. Protoconch globose, smooth, whorls rounded. Axial sculpture of last teleoconch whorl consisting of 3 high, rounded, spineless varices. Other axial sculpture of 4 or 5 high, narrow, rounded ribs. Spiral sculpture of numerous, weak, smooth threads. Aperture broad, rounded. Outer lip weakly crenulate, with 13-15 elongate lirae within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, open, spineless. Creamy-white with dark brown or reddish-brown spiral threads. Aperture white. Remarks. Haustellum franchii differs markedly from }X. haustellum in having a white aperture, in the colour of the shell and different protoconch (rounded and smooth, with 1.5-2 whorls in 77. franchii while conical, minutely punctate, with 2.25-2.75 whorls in 77 haustellum) . It differs from #. longicaudum and FH. fallax, both known from the western Indian Ocean, in its more rounded shell contour, weaker axial cords, and apertural lirations. Haustellum haustellum (Linnaeus, 1758) Figs 7-8, 13, 17-19, 23-26 Murex haustellum Linnaeus, 1758: : 746. Murex scolopaceus Rôding, 1798: 144 (ref. to Favanne, 1784). Aranea denudata Perry, 1811: pl. 45, fig. 1. Haustellum laevae Schumacher, 1817: 213 (ref. to Martini, 1777). ?Murex erythrostoma Swainson, 1840: 296 (non Murex erythrostomus Swainson, 1831). HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 Fig. 11. Distribution of Haustellum fallax. 15 30 45 60 75 90 105 120 135 150 165 180 165 150 Fig. 12. Distribution of Haustellum franchii. 89 APEX 14(3-4} 81-107, 20 déc. 1999 Protoconchs examined: Papua New Guinea, and Philippines (many specimens) (Coll. R. Houart, and pnvate coll.) Distribution. Singapore, throughout the Philippines, Taiwan, Ryukvus, Papua New Guinea, Solomon Islands, North Queensland, New Caledonia, and Fiji. Description. Shell up to 165 mm in length with 2.25-2.75 protoconch whorls and 8 teleoconch whorls. Protoconch weakly conical, last whorl minutely punctate, with spiral keel abapically. Axial sculpture of last teleoconch whorl consisting of 3 moderately high, rounded, spineless varices. Other axial sculpture of 3 or 4 low, weakly rounded, nodose ribs, crossed by low, weak, smooth primary and secondary spiral threads. Aperture broad, rounded. Outer lip weakly crenulate, erect, with weak elongate lirae within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, straight, open, smooth or occasionally with small spinelets adaperturally. Creamy-white to light brown with dark brown or reddish-brown spiral threads, and 3 dark brown to bluish-brown blotches on varices. Aperture apricot or pale pink. Remarks. Haustellum haustellum is a common species with globose body whorl and long, usually spineless siphonal canal. . longicaudum, H. kurodai kurodai, and Æ. k. vicdani Kosuge, 1980 have been tentatively synonymised by PONDER & VOKES (1988: 86) and F. fallax was synonymised in RADWIN & D'ATTILIO (1976: 49). H. haustellum is separated here on basis of morphological differences in shell structure and type of larval development. Æ. haustellum is the most common species of the genus. Haustellum kurodaï kurodai (Shikama, 1964) Figs 14, 35-37, 47-48 Murex kurodai Shikama, 1964: 33, pl. 3, figs 1, 2. Protoconchs examined: Philippines, Siasi Id (2 sp.) (Coll. R. Houart), and a few specimens in private coll. Distribution. The Arafura Sea and the Philippine Islands (Sulu Sea and Zamboanga). 90 Indo-West Pacific species of Haustellum HOUART Description. Shell up to 93 mm in length with 2-2.25 protoconch whorls and 7 teleoconch whorls. Protoconch globose, smooth, glossy, whorls rounded. Axial sculpture of last teleoconch whorl consisting of 3 narrow, high, nodose, almost spineless varices, each with occasionally 1 acute, short spine on shoulder. Other axial sculpture of 3 or 4 nodose ribs. Spiral sculpture of 4 or 5 wcak, tuberculate cords, more apparent on axial ribs, and numerous, low, smooth lirae between cords. Aperture broad, rounded. Outer lip crenulate, with numerous, weak elongate lirae within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, straight, open, with a single, acute, short spine adaperturally. Creamy-white with light to dark brown blotches on varices, axial ribs and siphonal canal. Occasionally occurs with darker coloured shoulder and/or siphonal canal or darker coloured teleoconch whorls. Aperture white, light peach or light pink. Remarks. Haustellum k. kurodai is here separated from A. haustellum in having a different protoconch (rounded and smooth in Æ. k. kurodai while conical, minutely punctate, with 2.25-2.75 whorls in Æ. haustellum) (Figs 47-48), a more fragile and lighter shell with lower spire, more shouldered teleoconch whorls, narrower varices, small spines on the siphonal canal. It is also usually smaller. For other remarks see Table 1. Haustellum kurodai langleitae Houart, 1993 Figs 14, 27-32, 45-46 Haustellum langleitae Houart, 1993: 145, figs 5, 10-12. Protoconchs examined: Mozambique, Nacala (3 sp); Madagascar (1 sp.) (Coll. R. Houart), and a few specimens in private coll. Distribution. Tulear, Madagascar, Mozambique, Tanzania, Pakistan, India, Sri Lanka, SW Java, West Sumatra. Description. Shell up to 94 mm in length with 2-2.5 protoconch whorls and 8 teleoconch whorls. Protoconch smooth, high, weakly shouldered. Axial sculpture of last teleoconch whorl consisting of 3 rounded, strong, spineless varices. Other axial sculpture of 2 or 3 strong ribs. Spiral sculpture of HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 15 30 45 60 75 90 105 120 135 150 165 180 165 150 Fig. 13. Distribution of Haustellum haustellum. Fig. 14. Distribution of Haustellum kurodai kurodai (stars), H. k. vicdani (square), and H. k. langleitae (circles). 91 APEX 14(3-4): 81-107, 20 déc. 1999 numerous, low, indistinct, smooth threads, more developed on axial ribs Aperture broad, rounded. Outer lip crenulate, erect with numerous, weak, elongate lirae within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, straight, open, spineless. Greyish-brown with bluish-black or brown blotches on spire and on siphonal canal. Aperture light yellow or pink Remarks. H. k. langleitae differs from #4. haustellum in having a different larval development, coarser, irregular, spiral sculpture, consisting of similar sized cords. Other shell characters, such as thickness, height of the intervarical ribs, breadth of the varices, are quite variable. For differences with 7. k kurodai see Table 1. Haustellum kurodaiï vicdani Kosuge, 1980 Figs 14, 33-34 Haustellum vicdani Kosuge, 1980: 57, pl. 17, figs 2, 4. Protoconchs examined: Philippines, Sorsogon (1 sp. partially broken) (Coll. R. Houart). Distribution. Currently known from a small area in the Philippine Islands: Sorsogon and Bulan, Luzon Island. Description. Shell up to 117 mm in length with 7 teleoconch whorls. Protoconch partially broken in examined specimens, globose, smooth, whorls rounded. Axial sculpture of last teleoconch whorl consisting of 3 high, rounded, spineless varices. Other axial sculpture of 3 or 4 low or moderately high, strong, nodose ribs. Spiral sculpture of numerous, low, weak, smooth threads. Aperture broad, rounded. Outer lip crenulate with very weak elongate lirae within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, straight, open, spineless. Lavender with scattered light or dark brown blotches on varices, shoulder, axial ribs and siphonal canal. Aperture light orange or orange-yellow. Remarks. The absolutely spineless siphonal canal and lavender colour separate that species from #1. k. kurodai and H. haustellum, two other species occuring in the Philippines. The shell is smoother, and spineless. For other differences see Table 1. 92 Indo-West Pacific species of Haustellum HOUART Haustellum longicaudum (Baker, 1891) Figs 15, 38-39, 42-43, 57-58 Murex haustellum var. longicaudum Baker, 1891: 56. Protoconchs examined: Ethiopia, Malajus (1 sp.); Gulf of Aden (1 sp.) (coll. R. Houart), Gulf of Aden (2 sp.) (coll. Wilhelm-Pieck-Universität, Rostock). Distribution. Southern Red Sea, the Gulf of Aden, the Gulf of Oman and the Persian (Arabian) Gulf. Description. Shell up to 87 mm in length with 2-2.5 protoconch whorls and 8 teleoconch whorls. Protoconch large, globose, irregularly shaped, smooth. Axial sculpture of last teleoconch whorl consisting of 3 narrow, high, spineless varices. Other axial sculpture of 3 or 4 low, nodose ribs. Spiral sculpture of numerous, low, smooth threads. Aperture broad, rounded. Outer lip crenulate, erect with very weak, elongate lirae within. Columellar lip smooth, strongly raised, adherent adapically. Siphonal canal long, straight, open, spineless. Light tan with dark brown blotches on varices and scattered blotches on teleoconch whorls. Aperture white. Remarks. Haustellum longicaudum is also occasionally synonymised with #7. haustellum. For differences in shell morphology with the other species see discussion under Haustellum and Table 1. The species is illustrated in D. & E. BoscH (1989: 57) as H. haustellum. Haustellum wilsoni D'Attilio & Old, 1971 Fig. 56 Haustellum wilsoni D'Attilio & Old,1971: 316, figs 1,2. Protoconchs examined: West Australia, Jurien Bay (1 sp.), West Australia, Augusta (1 sp.) (coll. R. Houart); original description and subsequent literature. Distribution. Geographe Bay to Jurian Bay, West Australia. Description. Shell up to 80 mm in length with 1.75-2 protoconch whorls and 6 or 7 teleoconch whorls. Protoconch broad, globose, smooth, glossy. Whorls rounded. HOUART Indo-West Pacific species of Haustellum APEx 14(3-4): 81-107, 20 déc. 1999 Fig. 15. Distribution of Haustellum longicaudum. 15 30 45 60 15 90 105 120 135 150 165 180 165 150 Fig. 16. Distribution of Haustellum wilsorii. 93 APEX 14(3-4}): 81-107, 20 déc. 1999 Axial sculpture of last teleoconch whorl consisting of 3 rounded, spineless varices. Last varix more prominent, others low or almost flat and undistinguishable in some specimens. Other axial sculpture of several, low, nodose ribs. Spiral sculpture of 10 or 11 low, rounded, tuberculatc cords with numerous smooth threads on and between the cords. Aperture broad, roundly-ovate or rounded. Outer lip crenulate with several weak elongate lirae within. Columellar lip smooth, raised, adherent adapically. Siphonal canal moderately long, straight, open, spineless with some knobs (or nodules) adaperturally. Ivory-white or creamy-white with some darker coloured (brown) blotches on and between varices and on siphonal canal. Aperture white. Remarks. A rare and remarkable species. It cannot be confused with any other species of the genus thanks to its broad, rounded protoconch, rounded and nodose whorls with deeply channeled suture, broadly developed apertural varix, and nodose base of the siphonal canal. ACKNOWLEDGEMENTS. I am particularly grateful to P. Bouchet (Muséum national d'Histoire naturelle, Paris), for reading the manuscript, for lis remarks, for his useful and appreciated comments on larval morphology, and for his advice on the Æaustellum haustellum group. For SEM of the protoconchs I am thankful to J. Cillis (Institut royal des Sciences naturelles de Belgique). For SEM of radulae I thank P. Bouchet and A. Warén (Swedish Museum of Natural History, Stockholm). Many thanks also to I. Bondarev (Ukraine) for giving me the opportunity to examine his material. I am also much indebted to G. Rosenberg (Academy of Natural Sciences of Philadelphia) for the loan of the holotype of Haustellum longicaudum (Baker, 1891), and to E. H. Vokes [Tulane University (retired)] for Fig. 1. I am also grateful to the referees for their useful comments, and remarks on the manuscript. REFERENCES BAKER, F.C. 1891. Remarks on the Muricidae with descriptions of new species of shells. Proc. Acad. Nat. Sci. Philadelphia 43: 56-61. BANDEL, K. 1975. Embryonalgehäuse karibischer Meso-und Neogastropoden (Mollusca). Abhandlungen der Matemetich- Naturwissenschaftlichen Klasse, Akademie der Wissenschaften un der Literatur. 1: 1-133. BosCH, D. & E. BosCH. 1989. Seashells of Southern Arabia. Motivate, Dubaï: 95. BOUCHET, P. 1989. A review of poecilogony in gastropods. J. Moll. Stud. 55: 67-78. 94 Indo-West Pacific species of Haustellum HOUART BOUCHET, P. & P. BAIL. 1991. Volutes from Saya de Malha Bank: The saga of Lyria surinamensis and a new species. Zhe Nautilus 105(4): 159-164. BoOZZETTI, L. 1993. Description of a new species of the genus /Jaustellum Schumacher, 1817 (Gastropoda: Muricidae) from the Western Indian Ocean. Apex 8(3): 107-110. D'ATTILIO, À. & W.E. OLD. 1971. A new muricid gastropod from Western Australia. The Veliger 13(4): 316-318. FAIR, RH. 1976. The Murex Book, an illustrated catalogue of Recent Muricidae (Muricinae, Muricopsinae, Ocenebrinae), Sturgis Printing Co., Honolulu, Hawaïi: 1-138. HOAGLAND, KE, & R. ROBERTSON. 1988. An assessment of poecilogony in marine invertebrates: phenomenon or fantasy? Biol. Bull. 174: 109-125. HOUART, R. 1990. New taxa and new records of Indo- Pacific species of Murex and Haustellum (Gastropoda, Muricidae, Muricinae). Bull. Mus. natn. Hist. nat., Paris, 4° sér., 12, sect À, n° 2: 329-347. HOUART, R. 1992. Description of a new species of Haustellum (Gastropoda: Muricidae) from the western Indian Ocean. Apex 7(1): 31-33. HOUART, R. 1993. Description of two new species of Haustellum Schumacher, 1817 (Gastropoda: Muricidae) from the Western Indian Ocean. Apex 8(4): 145-149. KOSUGE, $S. 1980. Studies on the collection of Mr. Victor Dan (2) Description of a new species of the genus Haustellum (Gastropoda: Muricacea). Bull. Inst. Malac. Tokyo 1(4): 57-58. LINNAEUS, C. von 1758. Systema naturae per regna tria natura. editio decima, reformata. Stockholm, vol. 1, Regnum animale: 1-824. OKUTANI, T. 1991. Mistaken localities for some shells "from Surinam". The Nautilus 105(4): 165. PARTH, M. 1995. Remarks on the infraspecific varieties of Haustellum dentifer (Watson, 1883) and Haustellum haustellum (L., 1758). La Conchiglia 27 (277): 61-65. PERRY, G. 1811. Conchology, or the natural history of snails. : 1-4, 61 pls., Miller, London. PETUCH, E.J. 1994. Atlas of Florida fossils shells. The Graves Museum of Archaeology and Natural History: 394 pp. PONDER, WF. & EH. VOKES. 1988. Revision of the Indo-West Pacific fossil and Recent species of Murex s.s. and Æaustellum (Mollusca: Gastropoda: Muricidae). Rec. Australian Mus., suppl. 8: 1-160. RADWIN G. & A. D'ATTILIO. 1976. Murex shells of the world. An illustrated guide to the Muricidae. Stanford University Press, Stanford: 1-284. RÔDING, J.F. 1798. Museum Boltenianum... Hamburg: 1-vil, 1-199. HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 SABELLI, B. & S. TOMMASINI. 1982. Osservazioni sulla radula e sulla protoconca di Bo/inus brandaris (L.. 1758) e Phyllonotus trunculus (L., 1758). Boll. Malac. 18 (9-12): 291-300. SCHUMACHER, H.C.F. 1817. Essais d'un nouveau système des vers testacés. Schultze, Copenhagen: 1- 287. SHIKAMA, T. 1964. Description of new species of Murex and Conus from the Arafura Sea. Venus 23 (1): 33-37, pl. 3. SMITH, E.A. 1901. On South African marine shells, with descriptions of new species. J. Conch. 10 (4):104-116. SWAINSON, W. 1840. A treatise on malacology or the natural classification of shells and shell-fish. Longman et al., London: 1- 419. VOKES, E.H. 1971. Catalogue of the genus Murex Linnaeus (Mollusca: Gastropoda. Muricinae, Ocenebrinae. Bull. Am. Paleont., 61 (268): 1-141. VOKES, E.H. 1990. Cenozoic Muricidae of the western Atlantic, Part VIIL Murex s.s., Haustellum, Chicoreus, Hexaplex: additions and corrections. Tulane Stud Geol. Paleont 23(1-3):1-96, pls. 1-12, 2 text figs. 95 7, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART Figs. 17-19. Haustellum haustellum (Linnaeus, 1758). Fig. 17. Philippines, Cebu, 144.8 mm. Fig. 18. West Sumatra, near Sibolga, 95 mm. Fig. 19. New Caledonia, 86.6 mm. Figs. 20-22. Haustellum fallax (Smith, 1901). Fig. 20. South Africa, off Natal, 63.1 mm. Fig. 21-22. Mozambique, between Cabo das Correntes and Zavora Point, 78.7 mm 96 HOUART Indo-West Pacific species of Haustellum APEx 14(3-4): 81-107, 20 déc. 1999 Figs. 23-24. Protoconchs of Haustellum haustellum (Linnaeus, 1758), Papua New Guinea. (scale bars: 1 mm) 97 APEX 14(3-4) 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART pos: 25-26. Protoconchs of Haustellum haustellim (Linnaeus, 1758), Papua New Guinea. (scale bars: 25: 1 mm; 6: 100 um). 98 HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 Figs. 27-31. Haustellum kurodai langleitae Houart, 1993. Fig. 27. Mozambique, Bazaruta Island region, 104 mm. Fig. 28. Southwestern Java, 124 mm. Fig. 29. India, Rameswaram, 90.2 mm. Fig. 30-31. Tanzania, Dar-es-Salaam, paratype, 94.1 mm. 99 APEX 14(3-4): 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART Fig. 32. Haustellum kurodai langleitae Houart, 1993, Java, Djakarta, 100 mm. Figs. 33-34. H. kurodai vicdani Kosuge, 1980, Philippines, Luzon Island, Sorsogon. Fig. 33. 93.5 mm. Fig. 34. 117 mm. Figs. 35-36. H. kurodai kurodaïi (Shikama, 1964), Philippines, Sulu, Siasi Island. Fig. 35. 62.3 mm. Fig. 36. 85.4 mm. 100 HOUART Indo-West Pacific species of Haustellum APEx 14(3-4): 81-107, 20 déc. 1999 Fig. 37. Haustellum kurodai kurodai (Shikama, 1964), Philippines, 86.5 mm. Figs. 38-39. H. /ongicaudum (Baker, 1891), Ethiopia, Malajus, 86.7 mm. Figs. 40-41. H. barbieri Houart, 1993, Madagascar, Sainte-Marie (Nosy- Boraha), paratype, 90.4 mm. 101 APEX 14(3-4): 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART Figs. 42-43. Protoconchs of Haustellum longicaudum (Baker, 1891), Gulf of Aden. (scale bars: 42: 1 mm; 43: 100 um). 102 HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 EEE ELELELELELELELELELELELELELELELELELELELELELELELÂLELE—E—— Fig. 44. Protoconch of Haustellum longicaudum (Baker, 1891), Gulf of Aden. (scale bars: 1 mm). Figs. 45-46. Protoconchs of H. kurodai langleitae Houart, 1993, Madagascar. (scale bars: 1 mm) 103 APEX 14(3-4) 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART Figs. 47-48. Protoconchs of H. kurodai kurodai (Shikama, 1964). (scale bars: 1 mm). 104 HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 TT ——————————————————_—_—_—_— Figs. 49-51. Haustellum bondarevi n.sp., Saya de Malha Bank. Fig. 49-50. Holotype MNHN, 79 mm. Fig. 51. Paratype coll. Bondarev, 101.2 mm. Figs. 52-53. H. franchi Bozzetti, 1993, Northeastern Somalia, off Ras Hafun, 99.8 mm, holotype IRSNB 27962/459. 105 APEx 14(3-4) 81-107, 20 déc. 1999 Indo-West Pacific species of Haustellum HOUART Fig. 54. Haustellum fallax (E. A. Smith, 1901), Mozambique (protoconch) (scale bar: 1 mm). Fig. 55. Haustellum barbieri Houart, 1993, holotype, detail of shell sculpture (scale bar: 1 mm). 106 HOUART Indo-West Pacific species of Haustellum APEX 14(3-4): 81-107, 20 déc. 1999 Fig. 56. Haustellum wilsoni D'Attilio & Old, 1971, West Australia, Augusta, 64.1 mm. Figs. 57-58. Haustellum longicaudum (Baker, 1891), Red Sea, holotype ANSP 60965, 49.7 mm. 107 COURTE NOTE AUX AUTEURS (Les instructions détaillées sont disponibles sur demande) Conditions générales. L'affiliation à la Société n'est pas obligatoire pour les auteurs. La publication des articles de maximum 20 pages imprimées est gratuite. Au delà de 20, chaque page sera facturée au prix exact de 1.000 BEF (environ 30 USD). Les auteurs non affiliés à notre revue devront assumer le prix coûtant des planches (pas du texte). Les différents coûts devront être réglés uniquement en francs belges avant la parution. Les articles décrivant de nouvelles espèces (sous-espèces) ne seront acceptés que si le matériel type primaire est déposé dans un Musée ou une Institution scientifique. Manuscrits. Les manuscrits seront rédigés de préférence en français ou en anglais. Ils doivent être dactylographiés, justifiés à gauche, sans hyphénation, avec double interligne, sur une seule face de papier A4. Les marges doivent être de 25 mm minimum. Trois exemplaires seront envoyés. La séquence des sections respectera l’ordre suivant : titre, nom de(s) l'auteur(s), adresse(s) de(s) l’auteur(s), mots-clés et résumé en anglais (et éventuellement en français). L'ordre suivant est suggéré: introduction, matériel et méthodes, résultats, discussion, conclusions, références, figures (et/ou tables, et/ou planches) et légendes. Les photographies doivent être imprimées sur papier brillant et montées sur un support adéquat dans le format final souhaité (max. 16 X 21 cm). Toute intervention de graphiste jugée nécessaire pour la présentation sera facturée aux auteurs. Les planches , couleurs sont aux frais des auteurs, au prix coûtant. Les noms de genre et d'espèce seront en caractères italiques ou soulignés. Les références dans le texte auront la forme: KEEN & CAMPBELL (1964) ou (KEEN & CAMPBELL 1964). La liste des références, en ordre alphabétique, respectera la forme: KEEN, A.M. & G.B. CAMPBELL. 1964. Ten new species of Typhinae (Gastropoda : Muricidae). The Veliger 7(1): 46-57. MEAD, AR. 1961. The giant African snail : a problem in economic malacology. University of Chicago Press, Chicago. MAYR, E. 1989. Attaching names to objects. In: What the philosophy of biology is : essays for David Hull (M. Ruse, ed.), 235-243. Klumer Academic, Dordrecht. Traitement des manuscrits. Les manuscrits seront soumis à deux referees (un troisième sera désigné si nécessaire). Les décisions de l'éditeur et les commentaires des referees seront communiqués aux auteurs, qui en tiendront compte. Une épreuve corrigée devra être renvoyée aux éditeurs sous forme informatisée. Elle devra respecter strictement les instructions de mise en page qui auront été communiquées aux auteurs. Une épreuve finale sera renvoyée aux auteurs pour correction. Une unique épreuve corrigée sera renvoyée à l’éditeur dans les plus brefs délais. Toutes modifications autres que des corrections d’impression seront facturées aux auteurs. Tirés-à-part. 30 exemplaires sont gratuits si au moins un des auteurs est membre de la Société. Les exemplaires supplémentaires seront facturés au prix coûtant. Non membre : à charge des auteurs, au prix coûtant (min. 50 exemplaires). Les frais de port sont toujours à charge des auteurs. Les manuscrits, les épreuves corrigées et toute correspondance sera adressée à : Société Belge de Malacologie, M. R. Houart, B.P. 3, B-1370 Jodoigne, Belgique. SHORT NOTE TO AUTHORS (detailed instructions available on request) General conditions. Membership is not mandatory for authors. Publication of papers with a maximum of 20 printed pages is free of charge. Beyond 20, every page will be invoiced to the exact price of 1.000 BEF (about 30 USD). Non-member authors should cover the costs of the plates (not the text). AII costs must be paid (in Belgian francs only) before publishing. Papers describing new species (subspecies) will be accepted only 1f the primary types are deposited in a Museum or a scientific Institution. Manuscripts. Manuscripts will be preferably in English or in French. They must be typed, ragged right (left- justified), without hyphenation, double-spaced throughout, on one side only of A4. Margins must be at least 25 mm. Three copies will be sent. The sequence of sections will respect the following order: title, name of author(s), address(es) of author(s), key words and summary in English. The following order is suggested: introduction, material and methods, results, discussion, conclusions, figures (and/or tables, and/or plates) and legends. Photographs must be printed on glossy paper in a final version (max. 16 X 21 cm), adequatly mounted. Any necessary intervention of a graphist will be charged to authors. Colour plates are fully charged to authors, at cost. 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