BULLETIN OF MARINE SCIENCE, 28(3): 543-549, 1978 CORAL REEF PAPER

A NEW VEXILLUM OF THE SUBGENUS PUSIA (: VEXILLIDAE) FROM THE BAHAMAS

William G. Lyons and Sally D. Katcher

ABSTRACT

Vexilllllll (PlIsia) chickcharneorum, new species, is described from three Bahamian Islands, and is compared with other similar Caribbean PlIsia. A summary of proposed names for Caribbean Pusia is presented, with additional comments on several species.

Western Atlantic species of the slilcata GmcIin, 1791 (p. 3455) (Cerno- Vexillum Roding, 1798, are divided among horsky, personal communication), and in any the subgenera Costellaria Swainson, 1840 event was erected for the similar Indo-Pacific and Plisia Swainson, 1840, all species of species properly known as VexilLum (Pusia) Vex ilium s.s. being confined to the Indo- microzonias (Lamarck, 1811), the type spe- Pacific region. Costellaria species bury in cies of Pusia (Cernohorsky, 1970). March mud and sand, whereas Pusia species inhabit (1852) improperly listed microzonias and rubble, coral reefs, and the undersides of V. (P.) cavea (Reeve, 1844) among Antillean stones (Cernohorsky, 1970). Specimens of species, followed soon thereafter by Krebs an apparently unnamed species of Pusia (1864), who included V. (P.) semicostatum were collected by one of us (WGL) at Great (Anton, 1839) in his list of West Indian Exuma and Cat Island, Bahamas, during species; the last two are also properly Indo- 1974 and 1976, and additional material from Pacific species. Dall (1889) tentatively listed Great Abaco was provided by Mr. Colin semicostatum and also included cavea, V. Redfern. We were initially reluctant to be- (P.) ebenus (Lamarck, 1811) and V. (P.) lieve that a locally common and distinctive speciosa (Reeve, 1844) from the western species from shallow Bahamian reefs might Atlantic, but ebenus is an eastern Atlantic be undescribed, but the following review species and speciosa belongs to the Indo- revealed no suitable previous names. Pacific fauna. VexilLlIm (Pusia) dermestinum (Lamarck, Cernohorsky (1970) combined 18 primary 181\) was the only Plisia known from the names and supposed synonyms into six "bio- tropical western Atlantic until five additional species" (pp. 3, 5) of western Atlantic Pusia species were described by Reeve (1844-45). (Table 1), based upon comparable variation In the period since Reeve's monograph, at he had noted in Indo-Pacific material; he did least 18 more names have been proposed not include V. (P.) cllbanum Aguayo and for western Atlantic Plisia by C. B. Adams Rehder, 1936, and V. (P.) arestum Rehder, (1845; 1850), March (1852), Dohrn (1862), 1943. In addition, he designated V. (P.) Sowerby (1874), Melvill (1925), Aguayo and variatum (Reeve, 1845) a synonym of the Rehder (1936), Rehder (1943), McGinty Indo-Pacific V. (P.) unifascialis (Lamarck, (1955), Nowell-Usticke (1959; 1968), and 1811), but varia tum properly belongs to the Sarasua (1975). Caribbean fauna (Abbott, 1974; Cern 0- Additionally, several erroneous names horsky, personal communication). Although have been used for western Atlantic Plisia. Abbott recognized Cernohorsky's revision, Voluta [= Vexillwn (PlIsia)] suLcata Gmelin, he listed nine western Atlantic species within 179 I (p. 3465) was long applied to the the subgenus Pusia, excluding V. (P.) epi- Caribbean species now known as Vexil/um phaneum which he included in CostelLaria. (Pusia) aLbocinctum (C. B. Adams, 1845), Nowell-Usticke (1968) named mi- but the former name is a homonym of V oluta nutus and M. hanleyi form antiguensis from

543 544 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3. 1978

Table 1. Primary "biospecies" and supposed synonyms of western Atlantic Vexi/lum, subgenus Pusia, as combined by Cernohorsky (1970)

1. dermestinum (Lamarck, 1811) 3. epiphaneum (Rehder, 1943) a/bicostatum (C. B. Adams, 1850) 4. histrio (Reeve, 1844) 2. exiguum (C. B. Adams, 1845) al'ticulatum (Reeve, 1845) han/eyi (Dohrn, 1862) a/boeinc/um (C. B. Adams, 1845) gemmatum (Sowerby. 1874) bifaseiatul11 (Morch, 1852) roseocaudatum (Sowerby, 1874) cruzanum (Nowell-Usticke, 1959) sykesi (Melvill, 1925) 5. puel/a (Reeve, 1845) moise; (McGinty, 1955) a/bol11acu/atum (Sowerby, 1874) hayesae (Nowell-Usticke, 1959) 6. pu/cheJlum (Reeve, 1844) the West Indies; the first is a homonym of other species described from unknown lo- Mitra minuta Roding, 1798, and the second, calities, but found none referable to the described as a form, is not available accord- species at hand. We conclude, therefore, that ing to Article 15 of the International Code the species has not been previously named. of Zoological Nomenclature. Both, however, Specimens of the new species are deposited appear to be Pusia, and should be added to in the molluscan collections of the British the list of names proposed for Caribbean Museum (Natural History) [BM(NH)], Lon- Pusia, as should Pusia splendidula Sarasua, don; the Academy of Natural Sciences of 1975, an apparent synonym of V. (P.) vari- Philadelphia (ANSP), Pennsylvania; the atum, described from Cuba. American Museum of Natural History Photographic illustrations of type speci- (AMNH), New York, New York; the Na- mens of all but two of the previously men- tional Museum of Natural History, Smith- tioned species were examined for this study. sonian Institution (USNM), Washington, Of the remaining two species, Vexillum D.C.; the Florida Department of Natural (Pusia) articulatum (Reeve, 1845), described Resources Marine Research Laboratory from an unknown locality, mayor may not (FSBC I), St. Petersburg, Florida, and the be a synonym of V. (P.) albocinctum (c. B. Redfern collection. Adams) as proposed by Cernohorsky (1970). The type could not be located in the British Vexillum (Pusia) chickcharneorum Museum (Natural History). Reeve (1845) new species described the species from a specimen in the Norris collection; according to Dance (1966), Figures 1-4 some types from that collection went to the Holotype.-Length 10.5 mm; north side of British Museum, but the remainder of the Stocking Is., Gt. Exuma, Bahamas; 23-25 collection went to Tomlin, whose collection June 1974; USNM 758543. is now in the National Museum of Wales at Cardiff. Reeve's type may yet reside among Paratypes.-1 paratype, 8.1 mm; north side this latter material. In any event, Reeve's Stocking Is.; 23-25 June 1974; ANSP illustration of articulatum (pI. 36, fig. 302) 345016.-2 paratypes, 7.3, 9.1 mm; same does not resemble our specimens. Pusia data; AMNH 183747.-2 paratypes, 8.4,9.0 bifasciata March, 1852 was not traced, but mm; same data; BM(NH) 1977165.-25 that name is a nomen nudum and a secondary paratypes, to 9.9 mm; same data; FSBC I homonym of Mitra [= Vexillum] bifasciata 17864.-1 paratype, 5.5 mm; northeast side Swainson, 1821, so it is not available. Cat Is., Bahamas; 12 July 1976; USNM We also searched the major treatments of 758544.-2 paratypes, 6.8, 11.2 mm; same Reeve (1844-45) and Sowerby (1874) for data; FSBC I 17866. LYONS AND KAICHER: NEW SPECIES OF VEXlLLUM 545

Figures 1-7. Caribbean Vexilllllll (PlIsia): 1, VexillulII cllickcllameortlm n. sp., juvenile paratype, 5.5 mOl, Cat Is., USNM 758544 (X 10); 2, subadult paratype, 8.1 mOl, Stocking Is., ANSP 345016 (X 5); 3, adultparatype, 11.2 mOl, Cat Is., FSBC I 17866 (X 5); 4, adult holotype, 10.5 mOl, Stocking Is., USNM 758543 (X 5); 5, immature V. ?chickchameo/'lllll, 7.9 mOl, Andros, FSBC I 17863 (X 5); 6, V. ulhocillctum (C. B. Adams), holotype, 15.0 mOl, Jamaica, MCZ 177080 (X 5) [after Clench and Turner, 1950]; 7, V. cuballwn Aguayo and Rehder, holotype, 12.0 mm, Cuba, USNM 420978 (X 5).

Other materiat.-l, 5.0 mm; east end Stock- Description.-Shell small, to about 11.2 mm ing Is.; 22 June 1974; FSBC I 17865.-1, total length; mature specimens ovate in out- 9.0 mm; Thurstone Bay, northeast of Trea- line. Protoconch of two smooth, glassy, sure Cay, Gt. Abaco; 16 November 1974; brown whorls. As many as six postembry- Redfern collection.-4, 7.9-9.0 mm; Powell onic whorls, each with nearly straight axial Cay and High Cay, Gt. Abaco; Redfern col- ribs crossed by 4-9 very faint spiral striae; lection. ribs numbering 13-14, 16-18, 18-20, 18- 546 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3. 1978

20, and 16-19 on first through fifth whorls; length from about 7.0-9.0 mm, are quite one shell with complete sixth whorl bearing globose (Fig. 2), and are apparently sub- 16 ribs. Aperture narrow, with thickened adults. Only three specimens are sufficiently callus posteriorly and four, occasionally five, large (9.9-11.2 mm) to demonstrate the more columellar plicae decreasing in strength ante- elongate adult form. The most recently dead riorly; a deep excavation between callus and adult shell (Fig. 3) is heavily eroded on the plicae. Base and anterior dorsal surface of spire by encrusting coralline algae, and a body whorl ornamented with four or five second specimen is badly abraded from surf nodulose spiral cords of varying strength, rolling. The third shell, although drilled followed anteriorly on base by about four both dorsally and laterally by a muricid, best oblique cords originating at columellar plicae. demonstrates the characters of the species Outer lip curved throughout, with small cren- and is selected as holotype (Fig. 4). ulations indicating terminations of nodulose We examined one immature specimen cords on anterior half. Living or freshly which we doubtfully assign to chickcharne- dead shells black, with very broad, well de- orum. The shell (length 7.9 mm; Fig. 5) has fined, white bands occupying most of each 514 whorls, with 12, 15, 16, 16, and 13 ribs whorl of spire, nearly all of posterior half of on the first five, respectively. Rib counts are body whorl; black color fading to rich chest- fewer on every whorl, but rib shape, and nut brown on older shells. color and sculpture of the base are similar to those of chickcharneorum. The first four Etymology.-We name the species for the spiral whorls and posterior portion of the Chickcharneys (or Chick Charneys), uniquely body whorl are completely white, in contrast Bahamian inhabitants variously described to shells of chickcharneorllm which otherwise by Voss and Voss (1960) as evil spirits or always possess a dark band just anterior to evil creatures, and by Kline (1974) as pixies, the suture. The atypical specimen was col- leprechauns, or gremlins. We suspect that lected alive from a crevice in living "fire these mischievous beings may have been coral" (Millepora sp.) on the barrier reef at responsible for the fact that the species has Wax Cut, Andros, Bahamas (by WGL) in remained undiscovered until now. September 1971. Discussion.-We do not accept all synonyms Vexillum (Pllsia) chickcharneorum be- for Caribbean Pusia proposed by Cerno- longs to the group of small, solid, generally horsky (1970). Although some synonyms ovate Vexillwn distinguished by spiral white undoubtedly exist, our examination of photo- bands or spots on black or very dark brown graphs of type specimens indicates that more backgrounds. Indo-Pacific species in this careful attention to comparison of types is group include Vexillum (Pusia) cavea needed, particularly in the exiguum-sykesi- (Reeve, 1844), V. (P.) consanguinewn moisei, hanleyi-gemmatum-roseocaudatum, (Reeve, 1845), and V. (P.) lelicodesmllln and histrio-albocinctum complexes. Proto- (Reeve, 1845), all illustrated by Cerno- conch forms indicate taxa in the first complex horsky (1970). In the Caribbean, V. (P.) may actually be assignable to the subgenus histrio is similar but is commonly larger Costellaria. Consequently, we do not feel it (usually about 15 mm in length) and is much propitious to provide a key to Caribbean more ornately colored, bearing one or sev- Pusia until these problems have been re- eral orange spiral bands in addition to the solved. single white band. Axial ribs of V. (P.) Spires on Vexillum (Pusia) chickcharne- histrio are more broadly rounded than are orum juveniles are sharply angled, with those of the new species. Vexillum (Pusia) nearly straight sides (Fig. 1), becoming more puella bears irregularly arranged white macu- expanded on fourth and fifth whorls; most of lations of varying size on its black spire, and our specimens have 5-5l1z whorls, range in is further distinguished by possession of LYONS AND KAtCHER: NEW SPECIES OF VEXILLUM 547

many fine, closely arranged axial ribs. Vex- complex are also banded, but they are ilium (PlIsia) albocinctwn is perhaps nearest slender, more elongate (usually to 15 mm or the new species, but differs by possessing greater in length), and their white spiral only single white spots on the center or bands are split by spiral incisions, creating anterior portion of each rib; the spots are two nodes on each axial rib. Species in the connected peripherally to form a narrow hanleyi complex are sometimes banded but spiral band (Fig. 6). are minute (usually less than 5 mm in length), Vexilll1m (Pllsia) cllbanum is somewhat strongly indented at each suture, and easily similar to the new species in size, color, and distinguished by possessing many sinuous shape. Aguayo and Rehder (1936) noted its axial lines adjacent anteriorly to sutures. color to be "white, with the lower half of the Vexillum (Pusia) epiphaneum has entirely last whorl chestnut, or often paler, in which white spiral whorls, being dark only on the case there is a chestnut band at the upper base and anterior portion of the body whorl, margin of this zone; early whorls pinkish but resemblance to V. (P.) chickcharneorum brown." The holotype (USNM 420978; is otherwise superficial; V. (P.) epiphaneum Fig. 7) is apparently of the paler form. Com- has a much more slender shell and attains a pared to V. (P.) chickcharneorum, it is a length of about 19 mm. This species and more elongate, polished shell with more most other Caribbean Pusia discussed but sharply defined columellar plicae continuing not illustrated herein are figured in Abbott uninterruptedly onto the base where they (1974). resemble thin carinae. Nodulose cords on Cernohorsky (1970) observed that Pusia the base of V. (P.) chickcharneorum are ab- species usually occur in reef habitats, where sent on V. (P.) cllbanum, which also lacks a they may be found under stones or within strong callus within the posterior portion of cracks and crevices. All specimens of the the aperture. There are only 10-13 axial new species were dead when collected. Most ribs on each whorl of V. (P.) cubanum, were found in depths of 1-5 m in accumula- whereas V. (P.) chickcharneorum has 16 or tions of dead shells in holes and crevices of more ribs on all except the first whorl. Size rocky platforms adjacent to seaward (wind- of V. (P.) cl1banwn (7+ whorls, length 12.2 ward) shores of small cays off Great Exuma mm; 6.5+ whorls, length 13.6 mm) is only and at Cat Island, two major islands in the slightly larger than that of mature V. (P.) eastern central Bahamas. Except for the chickcharneorum. Thurstone Bay specimen from an "inside" or Krebs (1864) mentioned Mitra [= Vex- mainland beach, all Abaca specimens are ilium (Pllsia)] semicostata Anton from St. from oceanic beaeh drift at offshore cays. Martin, Netherlands Antilles, but did not Associated mollusks also collected freshly report it later (Krebs, 1867) from the dead from shell accumulations at Great Bahamas. Reeve's figure (1845; pI. 37, sp. Exuma and Cat Island include Risomurex 308) of semicostata resembles V. (P.) chick- rosea (Reeve, 1856), Mitra barbadensis charneorllm in banding and overall form, (Gmelin, 1791), M. nodulosa (Gme1in, but both Reeve and Sowerby (1874) noted 1791), Vexillum (Pusia) dermestinum, V. (P.) that the final whorl of semicostata is smooth, histrio, V. (P.) puella, Conus granulatus unlike the strongly ribbed final whorl of V. Linne, 1758, and C. regius Gmelin, 1791, (P.) chickcharneorum. Although it is pos- attesting to the reef-like character of the sible that early workers may have confused habitat. Vexillum (Pusia) chickcharneorum these two species, Coomans (1963) and is probably also a member of that assem- Cernohorsky (1970) agree that V. (P.) semi- blage. costatum is properly assigned to the Indo- ACKNOWLEDGMENTS Pacific. Sincere appreciation is expressed to Mrs. K. Way, Species in the Vexillllm (Pusia) exiguum British Museum (Natural History), and Dr. J. 548 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3, 1978

Rosewater, National Museum of Natural History, mander J. R. Bartlett, commanding. XXIX. for providing excellent photographs of types of Report on the . Part 2, Gastropoda several western Atlantic Vexillum; Dr. Rosewater and Scaphopoda. Bull. Mus. Compo Zoo!. 18: also provided copies of several species descriptions. 1-492, pIs. 10-40. Dr. R. Turner, Museum of Comparative Zoology, Dance, S. P. 1966. Shell collecting. Univ. Calif. Harvard University, Cambridge, Massachusetts, Press. 343 pp. loaned the negative of the figure of the lectotype of Dohrn, H. 1862. Descriptions of new shells. V. (P.) albocinctum from Clench and Turner (1950). Pmc. Zool. Soc. Lond. 1862: 202-203. Mr. C. Redfern, Treasure Cay, Abaco, loaned speci- Gmelin, J. F. 1791. Systema naturae per regna mens from several localities at Great Abaco. Dr. tria naturae. Editio decima tertia, aucta, re- W. O. Cernohorsky, Auckland Museum, New Zea- formata. Leipzig. 1: 3021-3910. land, provided information on Voluta sulcata, con- Kline, H. (ed.) 1974. Yachtsman's guide to the firmed our conclusions regarding Vexillum (Pusia) Bahamas, 1975 ed. Tropic Isle Publ., Coral variatum and V. (P.) bifasciatum, and offered help- Gables. 320 pp. ful suggestions. The specimen questionably as- Krebs, H. J. ]864. The West-Indian marine signed to the new species was collected during shells with some remarks. A manuscript cruise G-7124 of the Rosenstiel School of Marine printed for circulation between collectors. and Atmospheric Sciences R/V GERDA under the Nykji:ibing, Falster. 119 pp. direction of Dr. D. R. Moore of that institution. 1867. Catalogue of marine mollusks Tropic Isle Publishers, Inc., Coral Gables, Florida, collected in the Bahama Islands in November, permitted quotation from Kline (1974). Dr. D. D. 1866. Ann. Lyc. Nat. Hist. N.Y. 8: 427-431. Turgeon and Mr. D. K. Camp, both of the Florida Lamarck, J. B. P. A. 1811. Sur la determina- Dept. of Natural Resources Marine Research Lab- tion des especes parmi les animaux sans verte- oratory, provided helpful comments. Contribution bres, et particulierement parmi les mollusques No. 299, Florida Department of Natural Resources testaces. Ann. Mus. Hist. Nat. Paris 17: 195- Marine Research Laboratory. 222. McGinty, T. L. 1955. New marine mollusks LITERATURE CITED from Florida. Proc. Acad. Nat. Sci. Phila. 107: 75-85, pis. 1-2. Abbott, R. T. 1974. American seashells, 2nd ed. Melvill, J. C. 1925. Descriptions of nine new Van Nostrand Reinhold Company, New York. species of Mitridae. Proc. Malac. Soc. Lond. 663 pp. 16: 2]5-219, pI. 10. Adams, C. B. 1845. Specierum novarum conchy- March, O. A. L. 1852. Catalogus conchyliorum liorum, in Jamaica repertorum, synopsis. Proc. quae reliquit D'Alphonso D'Aguirra & Gadea Boston Soc. Nat. Hist. 2: 1-17. Comes de Yoldi. Hafniae, fasc. ]: 1-170. 1850. Descriptions of supposed new Nowell-Usticke, G. W. 1959. A check list of the species of marine shells which inhabit Jamaica. marine shells of St. Croix, U.S. Virgin Islands, Contrib. to Conch. 7: 109-123. with random annotations. Burlington. iii + Aguayo, C. G., and H. A. Rehder. 1936. New 90 pp., 4 pIs. marine mollusks from Cuba. Mem. Soc. Cub. 1968. A supplementary listing of new Hist. Nat. 9: 263-268, pI. 24. shells (illustrated) to be added to the check list Cernohorsky, W. O. 1970. Systematics of the of the marine shells of St. Croix. Privately families Mitridae and Volutomitridae (Mol- published. 32 pp., 6 pis. lusca: Gastropoda). Bull. Auckland Inst. Mus. Reeve, L. A. 1844-1845. Conchologia iconica: 8: iv + 190 pp. or, illustrations of molluscous . Mono- Clench, W. J., and R. D. Turner. 1950. The graph of the genus Mitra. London. Vol. 2: western Atlantic marine mollusks described by pis. 1-39 (pis. 1-27: 1844; pis. 28-39: 1845). C. B. Adams. Occas. Papers on Mollusks, Rehder, H. A. 1943. New marine mollusks from 1: 233-403. the antillean region. Proc. U.S. Nat. Mus. 93: Coomans, H. E. 1963. The marine Mollusca of 187-203, pIs. 19-20. St. Martin, Lesser Antilles, collected by H. J. Sarasua, H. 1975. Nuevos genero, subgenero y Krebs. Stud. Fauna Curacrao Other Carib. lsI. especies de moluscos marinos neogastropodos. No. 72, XVI: 59-87. Poeyana 140: 1-15. Dall, W. H. 1889. Reports on the results of Sowerby, G. B. 1874. Thesaurus conchyliorum. dredging, under the supervision of Alexander Monograph of the genus Mitra. London. Vol. Agassiz, in the Gulf of Mexico (1877-78) and 4: 1-46, pis. 1-28. in the Caribbean Sea (1879-80), by the U.S. Voss, G. L., and N. A. Voss. 1960. An eco- Coast Survey steamer "Blake", Lieut.-Com- logical survey of the marine invertebrates of mander C. D. Sigsbee, U.S.N., and Com- Bimini, Bahamas, with a consideration of their LYONS AND KAICHER: NEW SPECIES OF VEXILLUM 549

zoogeographical relationships. Bull. Mar. Sci. ADDRESSES: (W.G.L.) FLorida Department of Nat- Gulf Carib. 10: 96-116. uraL Resources Marine Research Laboratory, 100 8th Ave. SE, St. Petersburg, FLorida 33701; (S.D.K.) DATE ACCEPTED: June 24, 1977. 5633 18th Way So., St. Petersburg, FLorida 33712.