THE NAUTILUS Volume 114, Number 1 March 27, 2000 ISSN 0028-1344 A cjuarteHij devoted to malacology. EDITOR-IN-CHIEF Dr. Jose H. Leal The Bailey- Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. Harasew\ch Department of Invertebrate Zoolog)' National Museum of Natural Historv' Smithsonian Institution Washington, DC 20560 ADMINISTRATIVE ASSISTANT Christina Petrikas The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Arthur E. Bogan North Carolina State Museum of Natural Sciences Raleigh, NC 27626 Dr. Philippe Bouchet Laboratoire de Biologic des Invertebres Marins et Malacologie Museum National d'Histoire NatureUe 55, rue Buffon Paris, 75005 France Dr. Robert T.Dillon, Jr. Department of Biolog)' College of Charleston Charleston, SC 29424 Dr. WilliaiTi K. Emerson Department of Li\ang Invertebrates The American Museum of Natural Histor\- NewYork, NY 10024 Dr. Eileen H. Jokinen Institute of Water Resources University of Connecticut Storrs,ct06269-401pe, NNM .59094, length 0.9 cm. mm) with 24—27-29 costae. Five mm width whorl (whorl 8, 9 or 10) with 32-35-38 costae. Aperture subcircular. Apertural length/shell length 0.28-0.29-0..30. Umbilicus very narrow. Egg capsules (Figure 43): Sub-.spherical, white, tran.s- parent. with protuberances, no noticeable embedded sand. Capsules closelv connected to each other along a twisted mucous thread. Page 6 THE NAUTILUS, Vol. 114, No. 1 Figures 15-21. SEM micrographs of species of Epitonitim associated with uiuslinKim corals off Ujung Pandang. 15-16. E. lochi new species 15. Shell. Scale line = 1 mm. 16. Protoconch. Scale line = 0.1 mm. 17. £. tihi Pilsbrw 1921, shell. Scale line = 1 mm. 18. E. hock.scmai new species, apical whorls. Scale line = 1 mm. 19. E. twilac new species, teleoconch whorl sculpture. Scale line = 0.1 mm. 20. E. hoeksemai new species, teleoconch sculpture. Scale line = 0.1 mm. 21. E. iilii Pilsbr\', 1921, teleoconch whorl sculpture. Scale line = 0.1 mm. Habitat: This species was recorded between 5-15 me- ters depth. Coral hosts were Heliofungia actinifonnis (Quoy and Gaimard, 1833) and Fungia ftiugitcs (Lin- naeus, 1758). One to 5 specimens were found attached by mucous threads to the underside of a coral, near a few hundred egg capsules. Type material: Holohpe NNM 59074, from t)pe lo- caliU-. Parat\pes: NNM 59081/1, Indonesia, Sulawesi, off Ujung Pandang, W. Lae-Lae, 9 m; NNM 59079/1, MZB/ 1, W' Bone Baku, 6 m: NNM 59080/1, 59082/1, 9 ni; NNM 59086/2, t\pe localit\-: NNM 59076/1, 6 m; NNM 59077/5, 12 m: NNM 59083/1, E Kudingareng Keke, 3 Gittenberger et al, 2000 Page 7 m: NNM 59078/1. W. Kudingareng Keke, 12 m; NNM 59075/1, 14 m; NNM 59084/1, NW. Lankai, 6 m; NNM 59087/2, Bone Lola. 15 m; NNM 59085/1. Bone Tam- biing. 6 ni. Type locality: Indonesia, Sulawesi, off Ujimg Pan- dang. W Samalona. 5 m tlepth. Distribution: Onlv known from Indone.sia, off Sula- wesi. Etymology: This species is named after Dr. B. W. Hoeksema, who supervised the field portion of this pro- ject. Remark.s: Shells of this .species resemble those of Ep- ituniuin iilu, but differ b)- a lengtli/width ratio of ca. 1.9 instead of ca. 2.6. Because most examined specimens are damaged, the fine structure of the costae could not be observed. The teleoconch sculpture appears alwavs ob- solete to the naked eye and the number of spiral threads increases more evidently in E. huehscmai, with ca. 13 spiral threads added between the third and the whorl. In £. lochi new species (see below), on the other hand, there is a more clearly reticulate sculpture on the early teleoconch whorls. Epitonium ingridae A. Gittenberger and Govid new species (Figures 2. 7-8. 23-24. 27. .30. 36. 46) Description: Shell (Figures 7-8, 23-24, 27, 30) (n = 5): Verv fragile, moderatelv slender conical, creamy white; reaching 20.8 in length. Lengtli/width 2.0-2.2- 2.3. Protoconch whorls ca. 3. Protoconch with three whorls, with numerous fine, incised, axial lines. Maxi- mum diameter of protoconch 1. 0.14-0.15 mm (n = 2) (Figure 30). Teleoconch whorls up to 10, separated by deep suture. Teleoconch sculpture of evenly spaced, or- thocline, thin, lamellate costae, and numerous very fine spiral threads (>100 on the 9th whorl), superimposed on somewhat coarser spiral cordlets (ca. 15 on fifth te- leoconch whorl). Initial whorls with multiple, lamellate costae, fused together to form thicker ones (Figures 23, 24). Coarser spiral cordlets are most prominent on initial teleoconch whorls, where thev are superimposed on cos- tae (Figure 24); coarser cordlets become obsolete on most abapical whorls. Costae are more or less damaged in all specimens; better preserved costae coronate. Par- ticularly below the periphery', costae somewhat curved abaperturally at their free margins, whereas adapically more erect and shghtly curved abaperturally or adaper- turally near suture, depending on position of costa on adjoining whorl. Costae mostly continuous, but hardly touching each other. Fifth teleoconch whorl (width 3.8 mm) with 20-24-31 costae. Five mm width whorl (whorl 6 or 7) with 23-30-33 costae. Aperture subcircular. Ap- ertural lengtli/shell length 0.3. Umbilicus very narrow. Egg capsules (Figures 36, 46) (n = 8): Oval (Figure 36), embedded with sand and closelv C(jnnected along straight, longitudinalK striated, mucous thread (Figure 46). Capsules 3,0-3.3-3.5 mm length and 1.5-1.6-2.0 mm in width. Capsules cont;un 93-120-173 white eggs. ProlM )scis: Whitish Type material: lloiotvpe NNM 59088, from t\pe lo- calih. Paratopes: NNM 59089/1. Indonesia, Sulawesi, off Ujung Pandang, W. Kudingareng Keke, 12 m; NNM 59090/2. 59092/1. E Samaknia. 9^m: NNM 59091/1, 24 m; NNM 59093/1, Bone Tambung, 7 m. Type locality: Indonesia, Sulawesi, off Ujung Pan- dang, ESE. Kudingareng Keke, 15 m. Other material examined: AMS 329657, Australia, Queensland, off Macgillivray Bav, Lizard Island, 14°39'S, 145°29'E. 10 m, I. Loch leg.; NNM unnum- bered, 1 shell, Indonesia, Ambon, llitu. outer part of Ambon Bav, eastern Laha, J. C. den Hartog leg. Distribution (Figure 2): Australia, Queensland; In- donesia, Ambon, and Sulawesi. Habitat: Specimens of this species were found be- tween 7-24 meters depth. Coral hosts were Fungia con- cinna Verrill, 1864, F fungites, Hcliofungia actinifonnis, Herpolitha limax and Foh/pht/llia talpina (Lamarck, 1801). Specimens were found attached by mucous threads to the underside of a coral; one or two speci- mens were found associated with up to a few hundred egg capsules. Etymology: This species is named after Ms. Ingrid van der Loo. Leiden. Remarks: Conchologically this species resembles the ■probaiile hoIot)pe" (Kaicher, 1981: 3036) of Epitonium clubium Sowerby 1844 (BMNH 1981234) from the Phil- ippines, which is an imperfect shell with a broken ap- erture and several apical wliorls missing. Its length could have been ca. 20 mm. Costae of adjacent whorls are continuous, slightly curved toward aperture adapically and away from aperture abapically, not projecting over suture. The holotvpe of Epitonium duhium most clearlv differs from E. ingridae bv its less prominent teleoconch sculpture and thicker, not lamellate costae. The speci- men figured bv De Boury (1912: pi. 7, fig. 4, Scala du- hia), which might represent E. duhium (of. Kilbum, 1985: 327) has more oblique costae. The identih' of Sca- laria graiji Nvst. 1871, (nomen novum for Scalaria stri- ata Gray, 1847, not Defrance, 1827) is unclear; Tryon (1887: 60, as S. striata) and De Bour\- (1912: 95, as S. striata and S. graiji) considered this nominal ta.xon a syn- onym of S. duhia. Kilburn (1985: 327) questioned this synonym)'. The shell of S. graiji figured by Tryon (1887: pi. 12, fig. 68, as S. striata) has a more shallow suture and relatively larger aperture when compared to E. in- gridae. The new species also resembles Epitonium fria- hilis (Sowerby 1844) from Western Australia. Swan Riv- er The holotvpe (BMNH 1966653). figured b\ Kaicher (1980: 2329).' is 16 mm in length ancf 7 mm in width, with ten whorls. It differs most conspicuousK' from E. ingridae bv its closed lunbilicus and absence of spiral threads. The species described and illustrated from Svd- ney Harbour as Foliaceiscala barLssa by Iredale (1936: Page 8 THE NAUTILUS, Vol. 114, No. 1 Figures 22-31. SEM micrographs of species of Epitoniiiin associated with mushroom corals off Ujimg Pandang (unless stated otherwise). 22. E. costulatum (Kiener, 1838), Indonesia, Ambon, Hitu, outer part of Ambon Bay, W Lalia, apical whorls. Scale line = 0.5 mm. 23-24. E. in^ridae new species 23. Apical whorls. Scale line = 0.5 mm. 24. Teleoconch whorl sculpture. Scale line = 0.1 mm. 25-29. Protoconch. 25. E. costulatum (Kiener, 1838) (same shells as figure 22). Scale hne = 0.1 mm. 26. E. hoeksemai new species. 27. £. inffidae new species. 28. E. twilae new species. 29. E. uhi Pilsbr\', 1921. Scale hne = 0.1 mm. 30-31. Protoconch 1. 30. £ ingridae new species. 31. E. twilae new species. Scale line = 0.02 mm. Gittenberger el ai. 2()()() Page 9 Figures 32-35. Species of Epitoniiim often confused in the literature. 32-33. E. tivilae new species, holotype, NNM 59104, length 1.5 cm. 34-35. £. buUntitm (Sowerby, 1844), holotype, BMNH 198136, length 2.0 cm. 300, pi. 22, fig. 15) seems to be similar in shape and size, but the costae are described as "of different strength, some fine, others large and recurved, while still others approach varices in size." Epitoniiim lochi A. Gittenberger and Goud new species (Figures 13-16, 37, 45) ?Epitonium species 4: Loch, 1982: 4-5, 1 fig. (see remarks below). Description: Shell (Fig^urc.s 13-16) (n = 4): Very fragile, elongate-conical, creamy white, reaching 8.5 mm in length. Length/width 2. 0-2. .3-2. 7. Protoconch whorls 3.25. Maximum diameter of protoconch 1 diameter 0.12 mm (n = 1). Protoconch with numerous fine, incised, axial lines. Teleoconch whorls up to 8, separated by a very deep suture. Teleoconch with evenly spaced, or- thocline, lamellate costae, crossing low spiral threads that are approximateK' a half to a fifth as wide as inter- spaces (Figure 15). Reticulate pattern present on most adapical whorls, replaced by spiral threads on later whorls. Third whorl with ca. 13 spiral threads, fifth one with ca. 15; spiral threads equally prominent throughout whorl. C^ostae usualK not continuous, lamellar hut rather low, barely touching preceding whorl. Due to damage in most specimens, fine structure of costae could not be examined. Fifth teleoconch whorl (width 1.3 mm) with 24-25-26 costae and 12-13-15 spiral threads. Aperture subcircular Apertural length/shell length 0.22-0.23. Umbihcus absent. Egg capsules {Figures 37, 45): The roundish, white, egg capsules (Figure 37) are mixed with sand, and close- ly connected to each other along a straight mucous thread without well-defined sculpture (Figure 45). Type material: Holotv'pe NNM 59094, from type lo- calit)'. Paratopes: NNM 59095/2, 59096/1, Indonesia, Su- lawesi, off Ujung Pandang, MZB/1, tyi^e locality; 16 m; NNM 59098/1, 18 m; NNM 59099/1, E Kudingareng Keke, 3 m; 59100/1, 12 m; 59102/1, 18 m; NNM 59101/ 2, ESE Kudingareng Keke, 15 m; 59103/1, Pulau Badi, 24 m. See also Remarks. Type localitj': Indonesia, Sulawesi, off Ujung Pan- dang, W Kudingareng Keke, 12 m. Other material examined: AMS 329687/2, Australia, Queensland, Lizard Island, Watsons Bav, 14°40'S, 145°27'E, 24 m, I. Loch leg.; AMS 329688/1, 329689/1, Granite Bluff, 14°39'S, 145°27'E, 23 m, I. Loch leg. (see Remarks below). Distribution: Indonesia and probably Australia. Habitat: The snails were found between 3-24 m. Fuitgia costulata Ortmann, 1889, and F. tenuis Dana, 1846, were coral hosts. If the Australian record really proves to be this new species, Fungia ci/clolitcs La- marck, 1816, should be included as an additional host (Loch, 1982: 4). One to 4 specimens were found at- tached by a straight mucous thread (Figure 45) to the underside of a coral accompanied by up to a few hun- dred egg capsules. Etymology: This species is named after Mr Ian Loch, who described this or a very similar species from Aus- tralia, without naming it. Remarks: Loch (1982) referred to and figured an un- named species 4' from Australia, distinguishing it from E. ulii. We were able to compare that material with the specimens collected off Ujung Pandang. In the Austra- lian specimens, the spiral threads are somewhat more prominent, which could represent some degree of intra- specific variation. The limited amount of material does not allow for conclusions on the identity' of the Austra- lian specimens; this prevented their inclusion as para- types. Conchologically this species is most similar to Epiton- ium zat rephes MeUiW. 1910 (holotype BMNH 191281683), from the Mekran coast. This shell is figured by Kaicher (1980; 2377); it differs bv having continuous Paee 10 THE NAUTILUS, Vol. 114, No. 1 Figures 36 — 48. .Spt'cii-N ol Epiloniiim associated with imislirooiu corals oii L'luiig Pandang. 36-38. Egg capsules. 36. E. ingridae. Scale line = 1 mm. 37. E. lochi new species. Scale line = 1 mm. 38. E. costiilatum (Kiener. LS38). Scale line = 1 mm. 39. Sclerite of alcvoniid octocoral found in association with egg capsules ot E. costiilatiim. Scale line = 0.1 mm. 40. Detail of figure 39. Scale hue = 0.1 mm. 41. E costiilatum. Eggs within a capsule. 42. E iilii Egg capsule with protoconchs 1. Scale line = 0.1 mm. 43- 47. Mucous threads. Scale line, with figure 43, = 0.01 mm. 43. E. hoekscinai. 44. E iilii. 45. E. lochi. 46. E. ingridac. 47. £. costulatiiin. 48. E. twilac, lar\'al shells (= protoconchs 1). Scale line = 0.1 mm. costae and b\- the more narro\\l\ spaced spiral threads, which are about as wide as their interspaces. The holotv'pe of Epitoniuin obliqita (Sowerbw 1844) [Scalaria] (BMNH 19812.31) also resembles E. lochi. but differs in having a clearh' open, though narrow, umbili- cus, and continuous costae. The holotvpe of Epitoniuin dcficrsi (Jousseaume, 1911) [Tenuiscala] (MNHN De Bou'r\-2706) from Aden, which has a broken aperture and missing apical whorls, can notwithstanding be distinguished from £. loclii bv the relatively small shell lengtli/width ratio (only ca. 1.9) and the presence of some varices. This holot\pe is also figured by Kaicher (1981: 3116). Epitoniuin twilae A. Gittenberger and Goud new species (Figures 2, 19, 28, 32-33, 48) Epitonium bullatum (Sowerbv, 1S44): Dushane. 1988; .30, figs. 5, 6. Yamashiro, 1990: 299, figs. 1-6. Not Scahiha hulla- tum Sowerby, 1844. Epitonium species 2: Loch, 1982: 3^, 1 fig. Epitonium sp.: Bratcher, 1982: 3, fig. 1. DescripHon: Slull (Fifiures 19, 28, 31, 32^3, 48) (n = 20): Very fragile, broad-conical, creamy white, reaching 17 mm in length. Lengtli/width 1.2-1.-1-1.6. Protoconch whorls 3. Protoconch widi numerous verv fine, incised, axial lines. Ma.\imum protoconch 1 diam- eter 0.12-0.13 mm (n = 2). Teleoconch whorls up to 9, straight-sided, separated h\ a shallow suture. Teleoconch sculpture (Figure 19) of unevenly spaced, fine, ortho- cline, more or less obsolete costae, not or onK' in part lamellate, and numerous low spiral threads that are sep- arated bv interspaces that varv from as wide as the spiral threads to three times as wide. Costae not continuous, not curved, barely touching adjacent whorls. Fifth teleo- conch whorl (width = 2. .5 mm) with 19-24.4-30 costae. Five mm width whorl (whorl 6 or 7) widi 24-31-36 cos- Gittenberger et ai, 2()()() Page 11 tae. Aperture subcircular. Apertural length/shell lengtli 0.37-0.44-{).52. Umbilicus ven uanou. Epe, UCMP 39918, 12.8 mm long, 7.5 mm wide, Uq^e iocalit>'. Para^pe UCMP 39919, t\-pe localit)'; Paratopes UCMP 39920 and 39921, Upper Tubara Group, below type localit\', Punta Pua, Bolivar, Colombia; Paratype USNM 501150, Moi'n Fm. 2 km west of Puerto Limon, Costa Rica; Paratvpe USNM 501151, Mofn Formation, between Puerto Limon and Pueblo Nuevo, Costa Rica. Type locality: Upper Tubara Group, Punta Pua, about 15 miles northeast of Cartagena, Bolivar, Colombia (UCMP S-66). Other material examined: USGS localit\' 21037, 1 specimen, Moi'n Formation, outskirts of Puerto Limon, Costa Rica; UCMP S-66, 11 .specimens; UCMP S-65, 2 specimens; TU 954, 1 specimen; TU 956, 1 specimen. Distribution: The k-nown lots of Coliimbella moinen- sis are from the Pliocene to Pleistocene (?) of Costa Rica and Colombia. The material from Colombia lacks pre- cise stratigraphic data but is thought to be from the Mio- cene to Pliocene Tubara Group (Yokes, 1990). Yokes (1990) reported that the muricid species she studied from these 2 (^oloniiiian localities are tspical of the Pleistocene Moi'n fauna from Costa Rica. .Although the Moi'n Formation has better stratigraphic definition than the above mentioned units, some controversy exists about the age of the formation. Coates et al. (1993) state that the Moi'n Formation is Late Pliocene in age, how- ever its fainias are more usuallv considered to be Pleis- tocene in age (Robinson, 1993, and references therein). Thus the stratigraphic range of this species is currently imprecise. Etymology: Columbclla itwiucitsis is named after the Moi'n Formation of Costa Rica. Discussion: Columbclla uioincusis is verv' similar to Columbclla mercatoiia (Linnaeus, 1758) (Figure 3). a common western Atlantic species k-nown from the Pli- ocene to the Recent. The new species can be differen- tiated from C. mercatoiia by its multiwhorled proto- conch. Columbclla mercatoria has 1.5 to 2 bulbous pro- toconch whorls (Figure 1), with at least one specimen having 2.25 whorls. The new species has a conical pro- toconch with 3 whorls (Figure 2), and its nuclear whorl is smaller than that of C. mercatoria. This latter species has nonplanktic development; its eggs hatch as crawling juveniles after about 33 days, during which the larvae feed on nurse eggs in the capsules (Bandel, 1974). The multiwhorled protoconch of C. moinensis, in contrast, is indicative of plank-tic development. Columbclla mercatoria varies greatly in adult size and strength of sculpturing, as do many other Columbclla species. In the case of C mercatoria, this variation may be a function of environment. In southern Florida, the species is commonlv associated with both reef rock and seagrass habitats, and specimens found in seagrasses tend to be smaller and less brightK patterned than reef specimens. The k-nown specimens of C. moinensis come from only 6 lots, and most of the material is similar in size. The single specimen from LISGS localit\' 21037 (USNM 501152, Figure 8) is inarkedK' larger than other specimens (Table 1), and has a weaker shoulder. It does have a multiwhorled protoconch (broken off though still with the shell) consistent with that of C. moinensis, so will be referred to the new species in spite of the dif- ferences between it and odier specimens. Many of the specimens that are considered to repre- sent C moinensis have eroded protoconchs and thus cannot be diagnosed with certaintv. However, the\- are comparable in shape and size to specimens of C mo- inensis from the same localities. Columbclla tnercatoria was contemporaneous with C. nwinensis. However, the 2 species have not so far been found svmpatricallv. Columbclla mercatoiia has been collected in the Pliocene and Pleistocene Mare and Abi- sinia Formations of Yenezuela (Weisbord, 1962) and in the Pliocene Bowden Formation of |amaica (a worn and damaged .specimen incorrectK' identified as Columbclla submercatoiia is illustrated by Woodring, 1928). Columbclla submercatoria Olsson, 1922 occurs in the Late Miocene of the Dominican Republic (Jung, 1994) Page 16 THE NAUTILUS, Vol. 114, No. 1 Figure 1. Recent CoIuml>elln mercntoria. protoeonch. UF 126820. Colon Iskuid. Bocas del Tom Province, Panama. Scale line = 2(X) jxni. Figure 2. Coliimhdln moineim.s new species. Protoconch of paratvpe, UCMP .39919. Sc;ilc line = 2(X) ]xm. Figure 3. Recent Columbdla inercntuiia (SBMNH 144H5I, Los Totiinios, Venezuela). Scale line = 5 mm. Figures 4—5. Colunibclla iitoinciisLs new species. Holohpe, UCMP 39918. Scale line = 5 mm. Figures 6-7. Columbdla inoinensi.s new species. Paratvpe, UCMP 39919. Scale line = 5 mm. Figure 8. Larsje specimen itf (^ohnnlirlla niciiiirii'ii-s now species. USNM .5011.52. Sc;Je line = 5 mm. M. J. deMaintenon, 2000 Page 17 Table 1. Lengths and widths in mm, and number of cords on the bodv whorl of t\pes and figured specimens Spe Max. length (mm) Ma\. width No. of (mm) spiral cords Holotvpe, UCMP 39918 Paratype, UCMP .39919 Paratvpe, UCMP 39920 Paratvpe, UCMP .39921 Parat\pe, USNM .5011.50 Paratvpe, USNM 501151 USNM .5011.52 12.8 7.5 14 11.4 6.3 14 15.3 8.7 12 15.8 9.1 17 10.1 5.5 15 12.5 7.2 16 18.6 10.4 17 and in the Neogene of Costa Rica. Whether this is a distinct species or a form of C. mcrcatoiia lias been de- bated and should still be regarded as uncertain. The pri- marv' difference between them is the greater number of spiral cords on C. submcrcatoiia (22 to 25 per whorl rather than 15 to 20), but large specimens of e.xtant C. mcrcatoiia from some areas of the Caribbean also have a greater number of spiral cords than usual. The type locahty for C. submcrcatoria is Red Cliff Creek, Costa Rica, a locality- that is presentK' imlocated but thought to correspond to late Miocene or earl\ Pliocene beds in the Limon Basin (Jung, 1994). Coliiinhclhi submcrcato- ria has a paucispiral protoconch of about 1.5 whorls (Jung, 1994), similar to that of C. rnercatoria. LITERATURE CITED Bandel, K. 1974. Spawning and development of some Col- umbellidae from the Caribbean Sea of Colombia (South America). The Veliger 16:271-282, Bouchet, P. 1989. A review of poecilogonv in gastropods. Jour- nal of Molluscan Studies 55:67-78. Coates, A. C, J. B. C. Jackson, L. S. Collins, T. M. Cronin, H. J. Dowsett, L. M. Bybell, P Jung, and J. A. Obando. 1992. Closure of the Isthmus of Panama: the near-shore marine record of Costa Rica and western Panama. Bulletin of the Geological SocieU- of America 104:814-828. Jung, P. 1994. Neogene paleontolog\- in the Northern Domin- ican Republic 15. The genera Coluinbdla. Eiin/pi/rciie, Panniictdrin. Coiwlla, Nitidella, and Mctulella (Gastro- poda: Columbellidae). Bulletin of American Paleontology 106(. 344): 1-45. deMaintenon, M. 1999. The phylogenetic relationships of mod- em columbellid taxa (Neogastropoda: Columbellidae), and the evolution of herbivorv from camivorv. Invertebrate Bi- ologv' 118:2.58-288. Hoagland. K. E. and R. Robertson. 1988. An assessment of poecilogonv in marine invertebrates: phenomenon or fan- tasy? Biolo'gical Bulletin 174:109-125. Knowlton, N. 1993. Sibling species in the .sea. Annual Review of Ecology' and Systematics 24:189-216. Moolenbeek, R. G. and H. J. Hoenselaar. 1991. On the identity of 'Coliiinbi'llti nistica' from West Africa and the Maca- ronesian Islands. Bulletin Zoologisch Museum 1.3:6.5-70. Oliverio, M. 1995. Larval development and allozyme variation in East Atlantic Columhella (Gastropoda: Prosobranchia: Columbellidae). Scientia Marina .52:77-86. Oliverio, M. 1996. Life-histories, speciation, and biodiversih" in Mediterranean prosobranch gastropods. Vie et Milieu 46: 16.3-169. Robinson, D. G. 1993. The zoogeographic implications of the prosobranch gastropods ot the Moi'n Formation of Costa Rica. American Malacological Bulletin 10:251-255. Weisbord, N. E. 1962. Late Cenozoie gastropods from North- ern Venezuela. Bulletin of American Paleontology 42(193): 1-672. Woodring, W. P. 1928. Miocene mollusks from Bowden, Ja- maica. Part 2: gastropods and discussion of results. Car- negie Institute of Wasliington. 564 pp., 40 pis. Vokes, E. 1990. Cenozoie Muricidae of the West Atlantic re- gion Pt. VIII — Miircx S.S., Haiistclliim. Cliicorciis and Hcxnplcx, additions and corrections. Tulane Studies in Ge- ology and Paleontolog}' 23:1-96. THE NAUTILUS 114(l):18-29, 2000 Page 18 The New Zealand Recent species oi Muricopsis Bucquoy, Dautzenberg and Dollfus, 1882 (Gastropoda: Muricidae) Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa PO. Box 467 Wellington NEW ZEALAND bnieeni@tepapa.govt.nz Kevin W. Burch 6 Oakland Avenue Whangarei NEW ZEALAND ABSTRACT Four New Zealand Recent Miiricopsis species are recognized: M. octogonus (Quoy and Gainiard, 1833), M. mariae (Finlay, 1930) and M. profunda new species, which belong in subgenus Murcxsul Iredale, 1915, and M. scotti new species, which is referred to Rolandiclla new subgenus together with the south- em Australian species A/, umhilicatus (Tenison Woods, 1876). Their shells and radulae are illustrated and distributions plot- ted. Mtiiicopsis mariae is interpreted as a species rather than a chronosubspecies of Af. espinosiis (Hutton, 1886) as has been traditional. Key words: New Zealand, Muricidae, Muiicopsis, Murcxsul, new taxa. INTRODUCTION For a number of vears a lar2e, locall\' common, sublit- tcjral form of Muricopsis from rockv ground off northern New Zealand was suspected to be an imnamed species distinct from M. octogonus (Quoy and Gaimard, 1833). Extensive collecting and field observ-ations led Scott (1989) to conclude that distinct species were indeed in- volved, and he reported differences in shell morphologv operculum colour and habitat. On areas of rock\' ground at 12-15 m depth, cut by channels with sand floors, Scott found that whereas M. octogonus t\picalh' lives both on horizontal rockv surfaces with dominant kelp (Eklonia radiata) and on sand at the channel floors, the undes- cribed species typically lives on the channel walls amongst abundant sponges and brachiopods. The pref- erence for vertical sublittoral rock surfaces would ac- count for its extreme raritv- on beaches and in dredge and trawl samples. The following revision was initiated after recent discovery of well preserved juvenile speci- mens (hitherto unavailable or unrecognised) of the un- described species, which show additional differences from M. octogonus that in turn suggest a closer relation- ship with the southern Australian species M. umbilicatus (Tenison Woods, 1876). Abbreviations and text conven- tions are: AIM, Auckland Institute and Museum; BMNH, The Natural History Museum, London; CM, Canterbury^ Museum, Christchurch; IS, Ian Scott collec- tion, Auckland: K\VB, Kevin W. Burch collection, Whan- garei (includes the outstanding collection formed bv our late friend Dave Gibbs); NMNZ, Museum of New Zea- land Te Papa Tongarewa, Weflington; NZGS, Institute of Geological and Nuclear Sciences, Lower Hutt. Unless specified, all material is at NMNZ (registration numbers preceded bv "M."). Length dimension precedes width in all cited measurements. SYSTEMATICS Superfamilv Muricoidea Rafinesque, 1815 Familv Muricidae Rafinesque, 1815 SubfamiK Muricidae Rafinesque, 1815 Genus Muricopsis Bucquoy, Dautzenberg and Dollfus, 1882 Muricopsis Bucquoy, Dautzenberg and Dollfus, 1882: 19. Type species (by original designation): Murcx hlainviUii Pa\Tau- deau, 1826: Recent, Mediterranean. Subgenus Murcxsul Iredale, 1915 Murcxsul Iredale, 1915: 471. Tvpe species (by original desig- nation): Murex octogonus Quoy and Gainiard, 1833; Re- cent, New Zealand. Remarks: Ponder (1972) considered Murcxsul to be a s\iion\Tn of Muricopsis. though Radwin and D'Attilio (1976), Yokes (1970, 1988) and Beu and Max-wefl (1990) have treated them as cbstinct genera. We agree with Vo- kes (1988) that they are closelv related, but Uke Houart (1988, 1991, 1993), prefer to interpret Murcxsul as a subgenus oi Muricopsis. Muricopsis (Murcxsul) octogonus (Quov and Gainiard, 1833) (Figures 1-15, 24, 25, .33, .37) Murcx octogonus Quov and Giiiniard, 1833: 531, pi. 36, fig. 8, 9; Kiener, 1843: 64, pi. 15, fig. 2; Grav, 1843: 229; Reeve, 1845, pi. 29, fig. 1.34; Kiister and Kobelt, 1869: 79, pi. 28, B. A. Marshall and K. \X. Burch, 2000 Page 19 fig. 23; Hutton, 1880: 47; Poirk-r, 1883; 112; Snter. 1901: 61. Mnrex penividiius Sowerln, 1841a: 8, lit;. 103; Sovverhw 1841b: 147; \bkes, 1970; .327. Not Miinx pinniiiuiis Lamarck. 1816 (preoccupied). Miircx (Phi/Uonotus) octo^i^oiiiis. — Tr\on. 1880; 110, pi. .30. fig. 272. 273 onK' I, in purt — fig. 274 = Muricop.si.s aispidatiis (Sowerbv, 1879)); Hutton, 1884; 218. Miirex clipsactis. —Tnon. 1880; 110, pi. .30, fig. 281 onl\-. Not Broderip, 18.33. ' Miirex octogoniis var iimhilicata. — Suter 1901; 61. Not Tcni- son Woods, 1876. Murex {Miirii-antlui) octofionii.s. — Suter, 1913: 400. pi, 48, fig. 1. Murc'x (Miiricantha) octog,oiiu.s var iiiiihilirdtii.s. — Suter 1913: 401. Not Tenison Woods, 1876. Hcxiifilcx (Miircxsul) octogonii.s. — Iredale, 1915; 471; Wenz, 1941; 1090, text fig. 3096. Hexaplcx (Murexsul) octogonus var umbilicntns. — Iredale, 191.5; 471. Not Tenison Woods, 1876. Murexsul octogonus. — Finlav, 1926; 419; Yokes, 1964; 13, fig. 20, .50, 61; Ponder 1968; 31. fig. 1, 37-41; Yokes, 1970; 327; Fiiir 1976; 63, pi, 17. fig. 239; Radwin and DAttilio. 1976; 163, text fig. 104, pi. 26, fig, 6, 7; Abbott and Dance, 1983; 143, te.xt fig.; Scott. 1989; 6, text figs.; Ben and Max- well. 1990: .359, pi. 481. Murexsul cuvierensis Finlav, 1926; 419 = noiiun nudum. Murexsul cuvierensis Finlav, 1927: 487, pi. 24, fig. 2. Murex (Murex.'iul) octogonus. — Thiele, 1929; 290; Shikama and Horikoshi, 1963; 69, te.xt fig. 104. Murex {Murex.sul) ednae Smitli,"l940; 43, pi. 2, fig. 10. Muricopsis octogonus octogonus. — Ponder 1972: 237, te.xt fig. .3/24. Murcx.sut octagunus [sic]. — Kaicher 1978, card 1608. Muricopsis octogonus. — Powell, 1979; 170, pi. .35, fig. 1; Scott, 1989; 6, text figs.; Hart, 1993; 44, text fig. (in part— far right figure = holotvpe of Af . scotti new species). Type material: M. octogonus: syntypes (2), including the originally figured specimen (Figure 15) MNHN, Bay of Islands, New Zealand; M. pcniiianus: repositors' of txpe material unknown (not located at BMNH. K.M. Way, pers. comm.), "Pacosma\o, Peru" = New Zealand; M. cuvierensis: holotxpe AIM 70500, off Cuvier Island, 73 m; M. ednae: repositorx- of txpe material unhiown, "Japan" = New Zealand. Other material e.xamined: About 1000 specimens in NMNZ (152 lots) and K.W. Burch (24 lots) collections. Di.stribution (Figure 37): Pleist(K'ene (Castlecliffian) to Recent. Three Kings Islands and North Island, New Zealand, as far south as Kapiti Island (west coast) and Mahia Peninsula (east coast), 0-508 m; taken alive at low tide to 121 m. Remark.s: Muricopsis octogoitus is exceptionally vari- able in teleoconch moq'jhologx', and there is complete intergradation between the most extreme of the forms illustrated here (Figures 1-15). Despite great variation in sculpture of the last few whorls in adults, all of the specimens are identical in protoconch and earlv teleo- conch morphologx'. Moreover, we could detect no cbf- ferences between the most extreme forms in radular morphology or e.xternal anatomy. It thus seems ilear that all specimens under consideration here represent a sin- gle higliK variable .species. Mature specimens range from 29 to 92 (est.) mm in length. Spines may be entirely absent, or short to long. The secondary- spiral cords ma\- remain weaker than the primaries and spineless throughout, or the% ma\- enlarge to resemble the primaries before the last adult whorl, and some or all may develop spines that may be as long as those on the primaries. The siphonal canal may be short and broad and the fasciole rounded with low ridg- es, or the canal ma\' be long and narrow, and the fasciole set with canals from earlier stages of growth that encircle a narrow to rather large false umbilicus. The teleoconch may be reddisli brown with black spiral cords, or white or cream with vellovvish or (txpicalK ) reddish brown spi- ral cords. Shells with dark, extensive pigmentation pre- dominate in the littoral and locally to about 12 m. Spec- imens with short spines, and with secondary' spirals that enlarge to resemble the primaries predominate in the littoral and localK' to about 50 m, whereas long-spined shells on which the secondarv spirals remain weaker than the primaries txpically occur deeper than about 30 m, thfjugh localK- as shallow as about 20 m. The outer lip of mature specimens ma\' be smooth or (hpicalK ) dentate, and the inner lip rim ma\' be free and projecting below a narrow parietal area, or almost fully adlierent. Unusually large, entirely spineless shells occur off Spirits Bax', where thev' intergrade with short-spined specimens (Figures 9, II). Muricopsis octogonus has been observed on a number of occasions on sandy substrata, boring holes in and feeding upon the shallowK'-burrowing bivalve Tawera spissa (Desha\es, 18.35) (I. Scott, pers. comm.; K.W.B., pers. obs.). What it eats on rock-\- ground remains to be established. The radula is illustrated (Figure 33) for comparison with those of M. mariac, M. scotti and M. unibilicatus (Figures 34-36). There are no records of M. octogonus from the west coast of the North Island between Cape Maria van Die- men and Cape Egmont, or from the east coast south of Mahia Peninsula (Figure 37), and it would seem that the present distribution off the northeastern and southwest- em North Island is relictual, perhaps dating from prior to the last glacial maximum. How far back this .species extends in the fossil record is unclear. Specimens from a Late Pliocene (Nukuma- ruan) horizon near Waipukurau (GS 10858, former cut- ting on disused section of Ashcott Road, coll. A.C. Beu, NZGS) appear to be indistinguishable from Recent spec- imens in teleoconch moqihology but differ in having smoothly rountleil instead of angulate protoconchs as in Recent material (Figure 30) (see below). If it proves to be specifically distinct from M. octogonus, it mav- be as- signable to M. dihtcidus Marwick, 1931 (Early Pliocene, Gisborne District). Some of the New Zealand Cenozoic ta.xa are scarceK' distinguishable from M. octogonus or from each other, and it is likely diat M. octogonus is a direct descendent of one or other of them, such as M. Page 20 THE NAUTILUS. Vol. 114, No. 1 B. A. Marshall and K. W. Burch, 2000 Pa^e 21 proavitiis (Laws, 1935), M. proi^enitor (Laws, 1935), or M. manvicki (Max^A'ell, 1971), all from Middle Miocene (Lilbumian) beds. Muricopsis (Murexsul) mariae (Finlav, 1930) (Figures 17, 19, 27, 28, 34, 38) Murexsul inariac Finlay, 1930: 237; Kaiclicr, 197S, card 1639; Ben and Maxwell,' 1990: 359. Murexsul cspinosus mariae. — Ponder, 1968: 32, fig. 2, 32, 33, .34. Muricopsis espinosus marine. — Powell, 1979: 170, pi. 37. fig. 5; Scott, 1989: 6, text figs. Ty-pe material: HolotApe AIM 70502, Cape Maria van Dienien, northern New Zealand. Other material examined: Several hundred speci- mens in NMNZ (66 lots) and K.W. Burch (13 lots) col- lections. Distribution (Figure 38): Three Kings Islands and northeastern North Island, New Zealand, as far south as Anaura Bav, 0-233 m; taken living under rocks at low tide to 40 ni. Remarks: Muricopsis mariae differs principally from M. octogonus in consistentK- lacking spines, in being smaller relative to the number of whorls, in having stron- ger denticles within the adult outer lip, in attaining smaller size (ma.\imum length 29 mm as against ca. 9.3 mm), in being bluish white instead of tvpicalh' white within the aperture, and in tvpicalh' lacking a distinct shoulder angulation, at least on the early teleoconch whorls (some specimens have a distinct shoulder angu- lation on later whorls). In a sample of both species taken living together at 25 m in Whangaroa Harbour entrance (M. 1:37244, M. 74899), the protoconch ranges from 800- 1000 |j.m wide (mean = 900 ixnx SD = 6.053, n = 21) in M. inariac, and 650-1050 |jLm wide (mean = 930 |jLm, SD = 0.097, n = 17) in M. octogonus. The largest M. mariae protoconch seen is 1250 |jLni wide (M. 133712). The first whorl of the protoconch has a distinct angula- tion and the summit is more or less flattened in both species. Juveniles of A/, mariae and M, octogonus mav be difficult to distinguish, but in M mariae the primarv- spiral cords are stronger after the second teleoconch whorl, the secondary spirals enlarge more rapidly to re- semble the primaries, and the Secondaries on the sutural ramp become pigmented early on the third whorl in- stead of after the third or fourth whorl (compare Figures 24, 25 with 27, 28). Specimens from Cape Maria van Diemen to North Cape and off the Three Kings Islands are more broadly conical than most specimens from fur- ther south (Figures 17, 19) but are otherwise indistin- guishable. The diet of this species is unknown. The distinctive radiila is illustrated (Figure 34) for comparison with that of A/, octogonus, M. scotti and M. timhilicatus (F'igures .33, 35, .36). Ponder (1968) interpreted M. mariae as a chronosub- species of A/, espinosus (Hutton, 1886), though Beu and Maxwell (1990) suggested that they might well be dis- tinct species. Compared with 15 well preserved .speci- mens of M. espinosus obtained near Waipukurau (GS10858, map ref. U22/085.308, former cutting on dis- used section of Ashcott Road, WNW of Waipukurau, A.G. Beu, NZCS — lowest Nukumaruan, Late Pliocene), which are indistinguishable from the hpe material ob- tained nearln', M. mariae differs in having a protoconch that is shorter relative to its width with a shouldered (rarely rounded) instead of rounded, more or less bul- bous first whorl (Figure 27), and in having narrower spi- ral interspaces, especially on the first three or four te- leoconch whorls. In most specimens of M. mariae the secondary spiral cords enlarge to resemble the primaries more rapidlv than in M. espinosus. InterestingK- enough, the sample includes well-preserved specimens of a larger species that also lacks a shoulder angulation on the pro- toconch (Figure 30). The status of this larger fossil spe- cies is unclear, but it is certainly more closely related to M. octogonus than to M. espinosus. Muricopsis espinosus or a very similar species occurs in the bath\a! "coral thicket". Lake Ferry, Palliser Bay (Mangapanian: Late Phocene) (Figure 31). The protoconch has thus inde- pendently become angulate in the M. octogonus and M. mariae lineages since the Pliocene. The differences be- tween M. espinosus and M. mariae in protoconch mor- phology and similarities in teleoconch morphology sug- gest that thev are distinct species. The original description o( Murcx espinosus (Hutton, 1886) is based on more than one specimen, the larger of which provided the cited dimensions (30.0 X 14.5 mm, CM3294), the smaller of which was subsecjuently illus- trated bv- Hutton (1893) (28.0X13.3 mm, CM3295). Al- though these specimens each have two labels (none orig- inal) staring that they are "holotype ' and "paratype" re- spectively, they are in fact synt\pes because Hutton did not use these unecjuivocal terms. Ponder (1968, figs 35, 36) effectively designated the larger specimen as lecto- type by stating that it was the "holotype". Incidentally, Figures 1-12. Shells of adult Muricopsis (Murexsul) octogonus (Quoy and Gainiard). 1. Off Thompson Point, Wailieke Island, 22 m, KWB (length .54 mm). 2. Off Cuvier Island, 70 m, M..5911 (length 78.5 mm). 3. Off Motiikalian Island, W of Paparoha, Coramandel, 18-20 m, KWB (length 45.5 mm). 4. Scallop bed near Matiatia Bav entrance, Waiheke Island, 13 m, K\^'B (length 48 mm). 5. Off Three Kings Islands, craxpot, KWB (length 90 mm, est. 93 mm). 6. Raiifurlv Bank. East Cape, 31-47 m, M. 75079 (length 46 mm). 7. Off Mavor Island, .59-74 m, M.64825 (length 63.5 mm). 8. Off Motnhoropapa Island, The Noises, 25 m, K'WB (length 42.5 mm). 9. Off Spirits Bav, 48 m, M. 137051 (length 54 mm). 10. Hooper Point, Spirits Bav 12 m, M. 1.346.30 (length 29 mm). 11. Off Tom Bowling Bay, 49 m, M. 137090 (length 64 mm). 12. Reotalii, Whangarei, low tide, KWB (length 45.5 mm). Page 22 THE NAUTILUS, Vol. 114, No. 1 Figures 13-23. Shells of Muricopsis species {all specimens adult except IS). 13-15. Muricopsis (Murexsul) octogonits (Quoy and Gaimard). 13. Ranfurlv Bank. Ea.st Cape, 89-94 m, M.64812 (length 46 mm). 14. Whangaroa Harbour entrance^ 2,5 m, M. 74899 (length .38 mm). 15. originallv figured s\iit\pe, MNHN (length .3.5. .5 mm). 16, 23. Murexsul (Muricop.sis) profunda new species, holohpe, off Three Kings Islands, 91 m, M. 70356. 17, 19. Muricopsis (Murexsul) marine (Finlav). 17. Archvvav Island. Three Kings Islands, 15 m. M. 117096 (22.3X12.3 mm). 19. Whangaroa Harbour entrance. 25 m. M. 137244 (26x13 mm). 18, 20. Murexsul (Rolniicliella) umhilicatus (Tenison Woods), Gulf St. Vincent, South Australia, South Australian Museiun (Figure 20, length .32 mm). 21, 22. Murexsul (Rolandiella) scotti new species. 21. Holotvpe, Mathesons Bay, Leigh, 6-9 m, M. 138186 (length 44 mm). 22. Paratype, Southeast Bay, Great Island, Three Kings Islands, 18 m, KWB (length 53.5 mm). Scale Una = 1 mm. B. A. Marshall and K. W. Burcli, 2()()() Page 23 Figures 24-32. Early whorls of Miiiicopsi.s species. 24, 25. Muiicop.sK {Miircxsiil) ociof^oiius (Qiio\' aiid Gainiard), VVhangaroa Harlioiir entrance, 25 m, M. 74899 (shell length 7.05 mm). 26. Miiirx.stil (Roland icllti) scotti new species, submarine cave S side of Ro.seman' Rock, Princes Islands, Three Kings Islands, 20 m, M. 117084 (length 6 nnn). 27, 28. Muricopsi.s (Murcx.sul) niariac (Finlay), Whangaroa Harbour entrance, 25 m, M. 1.37244 (shell length 4.75 mm). 29, 32. A/i/rico/i.v/.v (Miircx.wl) profunda new species, early whorls (29) and teleoconch (left) and protoconcli (right) microsculpture (32), NW ol Great Island, Three Kings Islands, 310 m, M. 93876. 30. Mttricopsis (Mtirexsul) sp. aff octogonu.s (Quoy and Gaimard), GSI0858, Ashcott Road, VVaipuknrau, shallow water. Late Pliocene (lowest Nukvnnanian), NZGS TM8078. 31. Muricopsi.s (Murexsul) sp. cf espinosus (Hutton), "coral thicket". Lake Ferry, Palliser Bay, bathyal. Late Pliocene (Mangapanian), M. 40408. Scale line for figure 32 = 50 |xm, other scale lines = 500 |jini. Page 24 THE NAUTILUS, Vol. 114, No. 1 Figures 33-36. Radulae of Muricopsis species. 33. Muricopsis (Murexsul) octogonus (Quoy and Gaimard), off Te Arai Point, Northland, 41 ni, M. 137251 (shell length 65 mm). 34. Muricopsis (Murexsul) mariae (Finlay), Governors Pass, Great Barrier Island, 11-29 m, M. 21784 (shell length 22 mm). 35. Muricopsis (Rolandiella) scotti new species, off Cape Rodney, 11-13 m, M. 137416 (shell length 40 mm). 36. Muricopsis (Rolandiella) umbilicatus (Tenison Woods), Edithburgh, South Australia: South Australian Museum, Adelaide, unregistered (shell length 28 mm). Scale lines = 50 jxm. this t)-pe material was mislaid imdl recentK', and omitted from the catalogue of tspe and figured fossils in the Ge- ology Department of the Canterbury Museum (Brad- shaw et al, 1992). Muricopsis (Murexsul) profunda new species (Figures 16, 23, 29, 39) Description: Shell up to 10.5 mm high (holotype adult?), fusiform, of moderate thickness; protoconch and teleoconch pale orange, adapical 4 primary spirals and secondary spirals between them reddish brown on last whorl. Protoconch 830-870 |xm wide excluding flared rim, of 1.5 convex whorls, covered with minute hemi- spherical granules, summit of first whorl angulate, last half whorl rather e\enly convex. Teleoconch of 4.5 reg- ularly expanding whorls; first whorl broadly convex; sub- sequent whorls with pronounced supramedian angula- tion, sutural ramp broad and flat, side broadl\' convex, smoothly confluent with siphonal canal. Sculpture con- sisting of axial varices, and rounded primarv' and sec- ondary spiral cords. Axial varices strongly retracted from adapical insertion throughout, 12 on last whorl (holo- tspe), narrow on 1st whorl, sloping adaxialh so that bases are overhung, blade like and strongly adapicalK- pro- duced above protoconcli/teleoconch suture and cement- ed to side of last whorl of protoconch, e.xtending beyond adapical extremitv of last protoconch whorl or to about mid whorl length. .Axial varices on subsecjuent whorls becoming diicker and rounded, each sumiounted by 4 or 5 fine, collabral, ada.xially sloping, axial lamellae. Axial interspaces with fine collabral growth lines and a few lamellar growth lines. Primarv spiral cords numbering 3 on spire and 3 on base; spire spirals commencing ini- niediatelw similar throughout, adapical spiral at shoulder angulation, abapical spiral bordering suture; adapical 2 basal spirals similar to spire spirals, abapical spiral stron- ger, widely separated. Secondary spirals numbering com- mencing on 3rd whorl, remaining weaker than primaries, 2 on ramp, 1 between each priniar\- spiral, 3 between abapical basal primaries, and 1 between abapical basal primarv and tip of canal. Siphonal canal of moderate B. A. Marshall and K. W. Burch, 2000 Page 25 Figures 37. Distributions of New Zealand Recent Muricop- sis {Miircxstd) octogonus (Quov and Gaimard) (500 and 1000 m isobaths indicated). Figure 38. Distributions of New Zealand Recent Muricopsis (Miircxsiil) marine (Finlay) (500 and 1000 m isobaths indicat- ed). length, gently flexed, partly enclosed by thin extension of inner lip. Aperture p\riform; outer lip rather thin at rim, weakly thickened within, simple; inner lip fully ad- herent adapically, rim free at abapical extremity and ex- tending almost to canal tip. Animal unktiown. Type material: Holotype NMNZ M. 70356, 34°11'S, 172°10'E, off Three Kings Islands, northern New Zea- land, 91 m, 19 Februar\' 1974, r.v. Acheron. Paratx^jes: 33°59.2'S, 172°13.6'E, 18 km N of Great Island, Three Kings Islands, 155 m, 23 June 1978, r.y. Tangaroa (1 juvenile, M. 137247); 34°06.5'S, 172°04.7'E, llkm NW of Great Island, 310 m, 30 June 1978, r.v. Tam^awa (1 juvenile, M.93876). Other material examined: 34°009'S, 171°44 7'E, Middlesex Bank, NW of Three Kings Islands, 201-216 m, 31 January 1981, r.v. Tan^aroa (2 juveniles, M.93302); 34°02.0'S, 171°44.0'E, Middlesex Bank, 246- 291 m, 31 January 1981, r.v. Tan^aroa (3 juveniles, M. 137245); .34°05.9'S, 171°55.1'E, 24 km NW of Great Island, Three Kings Islands, 710 m, 27 June 1978, r.v. Tangaroa (4 juveniles, M. 137246); 34°17.6'S, 17r45.3'E, 39 km SW of Great Island, 427 m, 21 June 1978, r.v Tangaroa (3 juveniles, M. 94338). Distribution: Off Three Kings Islands, northern New Zealand, 91-710 m (shells only). Remarks: Muricopsis profunda differs from M. octo- goniis and M. mariac principalK' in having ;Lxial varices that ride up onto the protoconch from the next whorl, in that the outer lip is much more strongly retracted from the suture, and in that the spiral cords are stronger at equivalent stages of growth and become pigmented at a later stage of growth. The size attained by this spe- cies is unclear, though it may not grow much larger than the holotvj^e (length 10.5 mm). Etymology: Deep (Latin). Rolandiella new subgenus (of Muricopsis) Type species: Muricopsis (Rolandiella) scotti new species; Recent, northern New Zealand. Diagnosis: Sliell similar to those of species of Muri- copsis (Murexsul), but with median shoulder angulation and weak abapical 2 primary spiral cords on first 1.5 teleoconch whorls. Inner Lip of most specimens upstand- ing over most of its length. Description: Shell 21-62 mm high at maturity, spire about as high as last adult whorl or slightly shorter, stout, of moderate thickness. Protoconch of 1.5-2.0 rovmded whorls. Teleoconch of mature specimens widi 5.5-7.0 convex, shouldered whorls, shoulder median on first 2 whorls, ascending to adapical third or slightly higher on subsequent whorls; sutural ramp shallowly concave; side broadly convex, smoothly confluent with siphonal canal. Primary- sculpture consisting of strong, axial varices and spiral cords with spines at intersections. Varices travers- ing all whorls, angulate in section, numbering 8-11 (7 or 12 in occasional specimens) on adult penultimate whorl. Primary spiral cords rounded, numbering 3 on spire and 3 on base in adults. Adapical primary spiral at Page 26 THE NAUTILUS, Vol. 114, No. 1 shoulder angulation, peripheral; abapical primary bor- dering suture throughout, or separating as insertion of last few whorls descends to ne.xt priniarv spiral: adapical and abapical primaries commencing immediately; inter- mediate primary* commencing at about mid first whorl, gradually enlarging to resemble other primaries. Abapi- cal basal primary spiral wideK' separated from adjacent primary, interspaces ot other primar)' spirals closer and of similar width to each other. Secondary' spirals pro- gressively developing, 1-5 on sutural ramp, 1 each in some or all interspaces of primaiT spirals, 0-.3 betyveen abapical basal priman' spiral and tip of canal, others on summits of some or all primary- spirals, 1 or more en- larging to resemble primaries on last adult whorl. Pri- mar\' and 1 (abapical shoulder spiral) or more secondarv spirals procKiced as short to moderate-length spines at varices, spines on primar\' spirals of similar length or shoulder spiral longest. Secondary axial sculpture com- prising fine collabral growdi lines that are raised as thin, fragile lamellae on ramp against suture and where in- tersecting spiral sculpture. Aperture pvriform. Outer lip thickened within at maturity, strongh' so in particularly large specimens, which may also develop spirally elon- gate denticles. Inner lip thin, fulK* adherent over adap- ical third or quarter, abapical part typically free, flared and strongK' projecting, ajjapical extremity e.xtenchng well beyond flexure of siphonal canal. Siphonal canal of moderate length, almost enclosed by infolding: siphonal fasciole strong, forming an umbilicus in some ver\' large specimens. Radula and operculum similar to those in Miiricop.sis (s. str) and Murcxsul. Remarks: Rolandiclla is introduced for Muiicopsis scotti new species from northern New Zealand, and M. umbilicatiis (Tenison Woods, 1876) from southern Aus- tralia, both of which resemble typical Murcxsul species in gross facies, but differ in that the shoulder spiral is situated medially on the early spire whorls then ascends adapicalK', and the intermediate primary spiral on the spire commences later than the others and develops slowly. By contrast, in Muiicopsis species, all three pri- maries are strong and commence immediately on die first teleoconch whorl, and the shoulder spiral borders the suture then descends abapically over subsequent whorls. None of the named New Zealand Cenozoic species referred to Murcxsul by Beu and Maxwell (1990) seem to belong in Muiicopsis (Rolandiclla). though the rela- tionships of the Early Miocene species Muiicopsis cchin- ophorus (Powell and Bartrum, 1929) are unclear be- cause the early teleoconch morphology is unknown. Both M. (Rolaiuliclla) scotti and M. (R.) uinbilicatus have large, rounded, paucispiral protoconchs indicating non-planktotrophic development, and suggesting that they have either crawl-away larvae or drifting larval stag- es of short duration. We suppose that the\' are derived from a common ancestor with planktotrophic larval de- velopment that had a trans-Tasman distribution. How far back in time diis putative common ancestor may have @ . Figure 39. Distributions of New Zealand Recent Muricopsis (MtirexstiJ) profunda new species (circle) and Muricopsis {Ro- landicUa) scotti new species (squares) (500 and 1000 m iso- baths indicated). lived is impossible to estimate without fossil records or Tiiolecular data. Etymology: After Roland Houart, Belgium, in appre- ciation of his outstanding contribution to world muricid systematics. Muricopsis (Rolandiclla) scotti new species (Figures 21, 22, 26, 35. 39) Muricopsis species Scott, 1989: 6, text figs. Muricopsis octogonus. — Hart, 1993: 4.5, right text fig. onlv ( = holotype). Not Qnov and Gainiard, 18.3.3. Description: Shell up to 62 mm high, fusiform, spire about as high as last adult whorl or slightK' shorter, stout, of moderate diickness. Protoconch and first 1 or 2 te- leoconch whorls white, subsequent whorls white or pale buff, primary, secondary and some finer spiral cords red- dish brown, aperture porcellaneous white. Protoconch sharply delineated b\' low varix, of 1.5-2.0 smooth, con- vex whorls. Teleoconch of up to 7 convex, shouldered whorls, shoulder median on first 2 whorls, ascending to adapical third or slightly higher on subsequent whorls; sutural ramp shallowK' conca\e: side broadk' convex, smoothly confluent with siphonal canal. Primary sculp- ture consisting of strong, axial varices and spiral cords with spines at intersections. Varices traversing all whorls, angulate in section, numbering 8-11 (7 or 12 in occa- sional specimens) on adult penultimate whorl. Primary spiral cords rounded, numbering 3 on spire and 3 on base in adults. Adapical primary spiral at shoulder an- B. A. Marshall and K. W. Burch, 2000 Page 27 gulatioii, peripheral; ahapical spire spiral bordering su- ture at first, separating as insertion oi last few whorls descends to next (iornierK' adapieal basal) priinar\' spiral; adapical and abapical spire spirals commencing imme- diatek": intermediate spire spiriil commencing at about mid first whorl, slowlv enlarging to resemble abapical spire spiral and basal primary- spirals. Abapical basal pri- mar\ spiral wideK separated from adjacent primarw in- terspaces of other primars' spirals closer and of similar width to each other. Secondar\' spirals appearing pro- gressively, numbering 3-5 on sutural ramp, 5-7 in zone comprising summit of shoulder spiral space behveen it and adjacent primars, usualK 3 or 4 on sununits of other primary spire spirals, 3 or 4 between abapical 2 basal primaries, 3 between abapical basal priman* spiral and tip of canal, and generally 1 each in some or all inter- spaces of primar\' spirals, most or all secondaries re- maining weaker than primaries throughout, or some en- larging to resemble primaries on last adult whorl. Pri- man and some secondars' spirals produced as short to moderate-length spines at varices, shoulder spines lon- ger than others on spire, spines on larger spirals tspicalK of similar length on last adult whorl. Secondar\' axial sculpture comprising fine collabral growth lines that are raised as thin, fragile lamellae on ramp against suture and where intersecting spiral sculpture. Aperture p\Ti- form. Outer lip thickened within at maturity, strongly so in particularK' large specimens, which ma\' also develop spiralK' elongate denticles. Inner lip thin, fidh" adherent over adapical third or quarter, abapical part hpicalK' free, flared and strongly projecting, abapical extremits* extending well beyond flexure of siphonal canal. Siphon- al canal of moderate length, almost enclosed bv infold- ing; siphonal fasciole strong, forming an umbilicus in some vers' large specimens. Operculum with terminal nucleus, yellowish brown. Radula (figure 35): Central teeth each with strong, conical central cusp; smaller, conical lateral and marginal cusps, and 2 \er\' small inner lateral denticles that are well separated from lateral cusp. Lateral teeth narrow. Type material: Holotype NMNZ M. 138186, Mathe- sons Bay, Leigh, northern New Zealand, 6-9 m, Feb- niar\-March 1992, D.W. Gibbs (ex D.W. Gibbs and K.W. Burch collections). Paratvpes: 34°09.5'S, 172°08.8'E, Southeast Bay, Great Island, Three Kings Islands, 20-22 m, 4 March 1997, K.W. Burch and D.D. Crosby (2 juveniles, M. 134760); Southeast Bay, alive, 18 m, ISMay 1982, K.W. Burch (6, KWB); Princes Islands, Three Kings Islands, 15 m, 29 November 1983, G.S. Hardy and A.L. Stewart (1, M. 75135); S side of South- west Island, Three Kings Islands, ;ilive, 27 m, 12 Feb- nian' 1986, G.S. Hardv (2, M.84267): off Cape Maria van Dienien, craypot. May 1991 (1, KWB); Whangaroa Harbour entrance, alive, 18 m, November 1995, K.W. Burch and D.W Gibbs (2, KWB); Rikoriko Cave, Poor Knights Islands, alive, 30-40 m, 1982, M.II.B. ONeill (1, 'm.75181); Poor Knights Islands, alive. May 1979, A.S.W. Penniket (1, M. 133691); Poor Knights Islands, December 1980, K.W. Burch (1, M. 133692); Poor Knights Islands, alive, 15-20 m, 1983, K.W. Burch (4, KWB); Poor Knights Islands, alive, 1970, 1972, W. Doak (5, M. 133690); South Cave, Poor Knights Islands, alive, 17 m, 21 May 1969, A.N. Baker (1,' M.23617); South Harbour, Poor Knights Islands, tilive, 25 Mav 1969, A.N. Baker (3, M.23618); off Cape Karikari, 30m, 20 June 1981 (1, M. 133694); Urtjuarts Bav Whangarei Heads, alive, March 1965 (1, M. 133695); "The Canyon", N side of Burgess Island, Mokohinau Islands, alive, 15-21 m, Januari- 1985, D.W. Gibbs (3, KWB); Maori Island, Leigh,' 22 Mav 1975, A.S.W. Penniket (1, M. 133695); NE of Mathesons Bay, Leigh, alive, 18 m, Mav 1995, D.W. Gibbs (1, M.127040);'Mathesons Bay, alive, 6-10 m, Februars-March 1992, D.W. Gibbs (6,' KWB); S of entrance to Omaha Cove, Leigh, 20 ni, December 1967, I. Scott (2, IS); off Little Barrier Island, alive, craypot, T. Riley (2, M. 133693); Cape Rodney alive, 15 m,'june 1987, D.W. Gibbs (7, M.90125); Cape Rodney, alive, 15 m, 4 March 1989, I. Scott (12, M. 117277); Cape Rodnev, alive, 11-13 m, I. Scott, Mav 1989 (53, M. 137416); Cape Rodney alive, 18 m, June 1977 (1, M. 133698); Waika- wau Bay, Coramandel, alive, 16 m, April 1989, D.W. Gibbs (2, KWB); off Needle Island, Mercury Bay, alive, 16 m, January 1985, D.W. Gibbs (3, KWB).' Other material examined: S side of Rosemar\- Rock, Princes Islands, Three Kings Islands, 20 ni, 18 January 1985, FJ. Brook (5, M. 117084); off West Island, Three Kings Islands, Elinfiamitc wreck, alive, W Doak (1, M. 133696); .34°2().0'S, 173°06.6'E, N of North Cape, 163-168 m, 27 Januar\' 1981, r.v. Tonj^aroa (1, M. 137225); Rarawa Reef' Great E.xliibition Bay crav- pots, alive, 40 ni, August 1989, I. McMillan (2, M. 100382); Rarawa Reef cra\pots, alive, 73 m, 28 Sep- tember 1988, I. McMillan (6, M.95234); Rarawa Reef cra\pots, alive. 40 m, 16 December 1987, I. McMillan (3, M. 95315); off Cape Karikari, alive, cra\pots, 1988, I. McMillan (1, M.95322); Henr\- Island, ' Whangaruni, alive, 13 m, 5 December 1971, \.N. Baker (1, M.2592()); off Hen and Chickens Islands, alive (1, M.2686; 1, M.83882); Mathesons Bay, Leigh, alive, 15-20 m, 26 March 1995, A. Spurgeon (2, MT26998). Distribution (Figure 39): Three Kings Islands and northeastern North Island, New Zealand as far south as Little Barrier Island, 13-168 m; taken alive at 13-40 m from rocky groirnd. Remarks: Miiricopsis ■•icotti is most similar to the southern Australian species M. nmbilicatus (Figures 18, 20, 26) in early shell ontogeny and gross adult shell fa- cies than to any New Zealand Recent Murcxsiil species. Muricopsis scotfi differs from M. nmbilicatus in a num- ber of details including larger adult size (shell length 34— 62 mm versus 21-30 mm), stronger and more numerous secondary- spirals, insertion point of the last few whorls desceruling froirr the abapical spire spiral to the next (formerly adapical basal) primary spiral, and abapical secondary spiral on the sutural ramp never becoming as Page 28 THE NAUTILUS, Vol. 114, No. 1 large as tlie shoulder spiral. The radula of M. scotti is similar to those of M. umhihcatus and M. octogomis (Figures 3, 35, 36). Muricopsis scotti resembles M. octof^onus in size, but is readily distinguishable bv the following characteristics: orange brown instead of dark reddish brown operculum, rounded instead of angulate protoconch, generally larger aperture, shorter siphonal canal, and much stronger sec- ondary spiral sculpture on the primary spiral cords, es- pecially on the shoulder spiral and between it and the adjacent primary. Unlike M. octogonus, which has ex- tremeK' variable sculpture, M. scotti consistently has a distinct shoulder angulation, angular yarices and short spines that are not curved backwards, or at most only slightK' so. In M. scotti the shoulder spiral (adapical pri- man) is much stronger than the others on the first two teleoconch whorls and occupies a median position, then gradualK' ascends to about the adapical third on subse- quent whorls (Figure 26). By contrast, on the first 1.5 teleoconch whorls in M. ocfogoiius, the three primary' spire spirals are all strong and similar, and the shoulder spiral commences beside the suture and descends to about the adapical third, remaining there or at the adap- ical quarter on subsequent whorls (Figure 24). Although the rim of the inner lip may be upstanding or more or less fulK' adherent in both species, that in M. scotti is typically free and more strongly upstanding abapically, and extends abapicallv further beyond the angulation at the top of the siphonal canal. Shells of living M. scotti are invariably more or less covered widi living Br\'ozoa, barnacles and other epi- bionts, often with the addition of Waltonia inconspiciia (Sowerb\', 1846) (Brachiopoda), upon which it has been observed to feed (Scott, 1989 and pers. comm; K.W.B., pers. obs.). Etjinology: After Ian Scott, Auckland. ACKNOWLEDGMENTS For the loan of type material, other specimens, and in- formation, we are grateful to P. Bouchet and V. Heros (Museum National d'Histoire Naturelle, Paris), N. Hiller (Canterburv' Museum, Christchurch), I. Loch (Austra- han Museum, Sydney), I. Scott (Auckland), A. Spurgeon (Hamilton), FG. Thompson (Florida Museum of Natu- ral Historv', Gainesville), and K.M. Way (The Natural History Museum, London). We also record our indebt- edness to our late friends D.W. Gibbs, PR. Jamieson, and J.R. Penniket, who supplied some of die finest ma- terial used in this study; and Wellington Shell Club members, who provided Mahia Peninsula records of M. octogomis. Thanks also to A.G. Beu (Institute of Geo- logical and Nuclear Sciences, Lower Hutt), I. Scott (Auckland), and anonymous referees for comments on the manuscript, N. Heke and M. Hall (Museum of New Zealand) for pliotography and photographic printing re- spectively, and to W'. St. George (Institute of Geological and Nuclear Sciences, Lower Hutt) for scanning elec- tron microscope facilities. LITERATURE CITED Abbott, R. T. and S. P. Dance. 198.3. Compendium of seashells. A color guide to more than 4,200 of the world's marine shells. American Malacologists, Melbourne, 410 pp. Beu, A. G. and P. A. Maxwell. 1990. Cenozoic Molluscaof New Zealand. New Zealand Geological Survey Paleontological Bulletin .58:1-518. Bradshaw, M. A., A. J. Lobb, M. R Cave and K. Watson. 1992. Catalogue of type and figured fossils in the Geology De- partment Canterbury Museum. Canterbury Museum Bul- letin 8:1-11.3. Bucquoy, E., P. Dautzenberg and G.-F. DoUfus. 1882. Les mollusques marins du Roussillon. 1. Bailliere, Paris, 570 pp. Fair, R. H. 1976. The Murex book: an illustrated catalogue of Recent Muricidae (Muricinae, Muricopsinae, Ocenebri- nae). Audior's edition, Honolulu, 138 pp. Finlay, H.J. 1926. A further commentary on New Zealand mol- luscan systematics. Transactions of the New Zealand In- stitute 57:320^85. Finlay. H. J. 1927. Additions to the Recent molluscan fauna of New Zealand. No. 2. Transactions of the New Zealand Institute 57:48.5-487. Finlav, H. J. 1930. Additions to the Recent molluscan fauna of New Zealand. No. 3. Transactions of the New Zealand InstihJte 61:222-247. Gray, J. E. 1843. Catalogue of die species of Mollusca and their shells, which have hitherto been recorded as found at New Zealand, with the description of some lateK- discovered species. Appendix 4, pp. 22^8-265. In: Dieffenbach, E., Travels in New Zealand: with contributions to the geog- raphy, geology, botany, and natiual histor\' of that country. 2. Murray, London. Hart, M. 1993. New Zealand Muricidae. World Shells 6:4:3- 45. Houart, R. 1988. Description of seven new species of Muri- cidae (Neogastropoda) from the southwestern Pacific Ocean. Venus 47:185-196. Houart, R. 1991. Description of thirteen new species of Mur- icidae (Gastropoda) from Australia and the New Caledo- nia region, with range extensions to Soudi Africa. Journal of the Malaeological Society of Australia 12:3.5-55. Houart, R. 1993. Three new species of Muricinae and Muri- copsinae (Gastropoda: Muricidae) from Somalia, Christ- mas (Line Islands) and the Philippine Islands. Venus 52: 41-46. Hutton, F W. 1880. Manual of die New Zealand Mollusca. Hughes, Wellington, 224 pp. Hutton, F W. 1884. Rexision of the rachiglossate Molluscaof New Zealand. Transactions of the New Zealand Institute 16:216-2.33. Hutton, F W 1886. New species of Tertiary shells. Transac- tions of the New Ze;iland Institute 18:33:3-335. Hutton, F W 1893. The Pliocene Mollusca of New Zealand. In: J. J. Fletcher (ed.). The MacLeay Memorial Voliniie. Linnean Society of New South Wales, Sydney, 3.5-92. Iredale, T 1915. A commentar\' on Sviter's "Manual of the New Zealand Mollusca". Transactions of the New Zealand In- stitute 47:417-i97. B. A. Marshall and K. W. Burch. 2000 Page 29 Kaicher, S. D. 1978. Card catalogue of world-wide shells. Pack 16 — Muricidae .3. Kaicher, St. Petersburg, Florida. Kiener, L. C. 1843. Species general et iconograpiiie des co- quilles \i\autcs, coiuprenant la collection du Museum d'Histoire uaturelle de Paris, la collection Lamarck, celle du Prince Massena, et les decouvertes recentes des vova- geurs. Genr(> Rocher {Miirfx, Lin.). Bailliere, Paris, 1.30 pp. Kuster, H. C. and W. Kobelt. 18.39-1878. Die ge.schwanzten vmd bevvehrten Puqjurschnecken (Adurex. Rtmclla. Triton- ium, Trophoit. Hindsia). In: Kiister, H. C. and W. Kobelt (eds.), Systemati.sches Conchylien-Cabinet von Maituii und Chemnitz. . . 3 (2) 193. Bauer and Raspe, Leipzig, 336 pp. Poirier, J. 1883. Re\ision des Miirex du Museinn. Nouvelles Archives du Museum dHistoire Naturelle (2) .5:1.3-128. Ponder, W. F. 1968. Nomenclatural notes on some New Zea- land rachiglossan gastropods with descriptions of five new species. Records of the Dominion Museum 6:29-47. Ponder, W. F. 1972. Notes on some Australian genera and spe- cies of the family Muricidae (Neogastropoda). Journal of the Malacological Societv of Australia 2:21.5-248. Powell, A. W. B. 1979. New' Zealand Mollu.sca. Marine, land and freshwater shells. Collins, Auckland, 500 pp. Quoy, J. C. R. and P. Gaimard. 1833. Voyage de decouverts de I'Astrolabe, execute par ordre du Roi pendant les annees 1826-1827-1828-1829, sous le commandement de M.J. Dumont d'Umlle. Zoologie. MoUusques 2 (2). Tastu, Par- is: .321-686. Radwin, G. E. and A. DAttilio. 1976. Miirex shells of the world. An illustrated guide to the Muricidae. Stanford University' Press, Stanford. 284 pp. Reeve, L. 184.5-1846. Monograph of the genus Miirex. Con- chologia iconica: or illustrations of the shells of mollus- cous animals. 3. Reeve, London. 36 plates and captions. Scott, 1. 1989. A closer look at Miiricopsis octogonus. Poirieria. Auckland Museum Concholog\' Section L5:6-8. Shlkama, T. and M. Horikoshi. 1963. Selected .shells of the world illustrated in colours. HokurvTi-kan, Tokyo. 154 pp. Smith, M. 1940. Two new marine molluscs from japan. The Nantilus .54:43. Sowerby, G. B. 1841a (in 1832-1841). The conchological illus- trations. Miirex. Sowerby, London, 9 pp, 23 plates. Sowerby, G. B. 1841b. Descriptions of some new species of Miirex. principally from the collection of H. Cuming, Esq. Proceedings of the Zoological Societv of London 8 (1840): 137-147. Suter, H. 1901. Note upon Trophoii iimhiUcatns, Tenison Woods. The Journal of M;ilacology 8:61-63. Suter, H. 1913. Manual of the New Zealand Mollusca. With an atlas of plates [1915]. Mackav, Government Printer, Wellington, 1120 pp. Thiele, J. 1929. Handbuch der Systematischen Weichtierk-unde 1 (1). Fischer, Jena, 376 pp. Tryon, G. W 1880. Manual of concholog)-: structural and sys- tematic. With illustrations of the species. 2. Muricinae, Puqjurinae. Tryon, Philadelphia, 289 pp. Yokes, E. H. 1964. Supraspecific groups in the subfamilies Muricinae and Tritonaliinae (Gastropoda: Muricidae). Ma- lacologia 2:1^1. Yokes, E. H. 1970. The West American species of Murexiella, (Gastropoda: Muricidae) including hvo new species. The Yeliger 12:325-329. Yokes, E. H. 1988: Muricidae (Mollusca: Gastropoda) of the Esmeraldas beds, northwestern Ecuador, Tulane Studies in Geolog)- and Palaeontology 21:1-50. Wenz, W. 1941. Gastropoda. Teil 1: Allgemeiner Teil und Pro- sobranchia 5 (7), pp. 961-1200. 7».- Schindewolf O. H. (ed.), Handbuch der Palaozoologie 6. Bomtraeger, Berlin. THE NAUTILUS 114(1):30, 2000 Page 30 Notices CONSTANCE E. MALACOLOGY BOONE GRANTS TO The newly established Constance E. Boone Grants to Malacologv' proxddes annually up to $1000 to qualified persons undertaking c(jllection-based research on Re- cent mollusks at the Houston Museum ot Natural Sci- ence, Houston, Texas. The HMNS malacology collection contains over 48,000 cataloged lots (45,000 in database format), comprising approximately L5 million speci- mens. The main geographic focus of the HMNS is the Gulf of Mexico. 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Bogan North Carolina State Museinn of Natural Sciences Raleigh, NC 27626 Dr. Philippe Bouchet Laboratoire de Biologic des Invertebres Marins et Malacologie Museum National d'Histoire Naturelle 55, rue Buffon Paris, 75005 France Dr. Robert T.Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural H is ton- New York, NY 10024 Dr. Eileen H. Jokinen Institute of Water Resources University of Connecticut Storrs,Ct 06269-4018 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology- Harvard University Cambridge, MA 02138 Dr. Douglas S. Jones Florida Museum of Natural History University of Florida Gainesville, FL 32611-2035 Dr. James H. McLean Department of Malacology Los Angeles County Museum of Natural Histoiy 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard Universit)' Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department of Li\dng Invertebrates The American Museum of Natural History' New York, NY 10024 Dr. Gustav Paulay Marine Laboratory University of Guam Mangilao, Guam 96923 Mr. Richard E. Petit PO.Box30 North Myrtle Beach, SC 29582 Dr. Edward J. Petuch Department of Geology- Florida Atlantic University Boca Raton, FL 33431 Dr. Gai-y Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr. Ruth D. Turner Department of Mollusks Museiun of Comparative Zoolog)- Harvard University Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology University of California at Davis Davis, CA 95616 Dr. G. Thomas Watters Aquatic Ecology Laboratory 1314 Kinnear Road Columbus, OH 43212-1194 Dr. John B. Wise Houston Museum of Natural Science Houston, TX 77030-1799 SUBSCRIPTION INFORMATION The subscription rate per volume is US $28.00 for indi\-iduals, US $45.00 for institutions. Postage outside the United States is an additional US $5.00 for surface and US $15.00 for air mail. All orders should be accompanied by pa\anent and sent to: THE NAUTILUS.RO. Box 1580, Sanibel, FL 33957, USA. Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is published quarterly bv The Bailey- Matthews Shell Museum, 3075 Sanibel-Captiva Road, Sanibel, FL 33975. Periodicals postage paid at Sanibel, FL, and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS RO. Box 1580 Sanibel, FL 33957 TH EfyNAUTI LUS CONTENTS Valium' 114. Number 2 June 6, 2000 ISSN 0028-1344 Douglas G. Smith Notes on the ta\onoiny of introduced Bcllainija (Gastropoda: Viviparidae) species in northeastern North America 31 Guido Pastorino A revision of the Patagonian genus Xijmeiiopsis Powell, M. G. Harasewych 1951 (Gastropoda: Muricidae) 38 Jose H. Leal Copiilabi/ssia riosi, a new deep-sea limpet (Gastropoda: Luiz Rieardo L. Simone Pseudococculinidae) from the continental slope off Brazil with comments on the svstematics of the genus 59 G. Darrigran Invasion of the exotic freshwater mussel Limnopcnw I. Ezcurra de Drago fortunei (Dunker, 1857) (Bivalvia: Mytilidae) in South America 69 Warton Monteiro Location of allospermatozoa in the freshwater gastropod Toshie Kawano Bioiuphalaiia tcna^iophila (d'Orbigny, 1835) (Pulmonata: Planorbidae) 74 Jay Cordeiro Status of the tidewater mucket, Lcptodca ochracea (Say, 1817) (Bivalvia: Unionidae), in Halfway Pond, Massachusetts. USA .' 80 Notices 84 THE NAUTILUS 114(2):31-37, 2000 Page 31 Notes on the tcixonomy of introduced BeUaniya (Gastropoda: Viviparidae) species in northeastern North America Douglas G. Smith Department i)t Biologv' and Graduate Program in Organismic anti E\olntionar\ BiologN Uni\ersit\' ot Massachusetts Amherst,' MA 01003-5810 USA dgsmith@bio.uaniss.edu ABSTRACT The genus BeUnmi/a has become well established in Nortli America since its first introduction arovmd the turn ot the 20"' centurv. Some malacologists have recognized two species, usu- alK' placed in the genus Cipaiigopahidina. C. chinen.sis. and C. japonica. Other investigators, however, have questioned a two species concept or simply considered all populations to be xar- iants of a single species, C. chinensis. Differences observed in the juvenile shell and male reproductive anatomy, along with size-specific features of adult shells, enable easv separation of the hvo taxa and support the two species concept. Characters of female and male reproductive anatomv confirms the place- nient ot the species in the subfamik' Bellanninae. Anatomical features which ha\e been used to substantiate Cipanaopahi- dina are proposed to be of subgeneric value only. Kei/ words: Introduced snails, Cipangopaludina, Idiopoma. Vi- vipams. Lecijthoconcha. INTRODUCTION Among the more conspicuous members of the North American freshwater gastropod fauna are the large ex- otic viviparid snails customarily placed in the genus Ci- pan^opaJiidina. The North American history of Cipaii- gopahidina and its distribution was discussed and re- viewed by Clench and Fuller (1965) (as a subgenus of Vivipanis), Dundee (1974), and Joldnen (1982). The majority of North American populations are confined to the coasts, with scattered records from the interior At present, the east coast appears to comprise the greatest number of occurrences (Joldnen, 1982). Since the earliest reports of Cipangopaludina popu- lations in North America around 1900 there has been much debate regarding the identification and \ali(lit\' oi the various species and subspecies reptjrted and, to a lesser extent, the status and position of the taxon Cipan- gopaludina in the Vixaparidae. Hannibal (1912) proposed the subgenus Cipangopaludina to accomodate Idiopoma niallcata (Reeve, 1863), leaving /. japonica (von Mar- tens, 1861) in Idiopoma s. s. Subsequent authors (e.g.. Annandale, 1920; Prashad, 1928) regarded Idiopoma Pilsbrv', 1901, recognized primarily on features of the operculum, as a synonym of Vivipanis. Annandale ( 1920) used anatomical characters to propose a new ge- nus Lecijthoconcha to include L. cJiincncsis and, in a later paper (Annandale, 1921), to include L. japonica as well. Rao (1925) maintained use oi Lcci/thoconcha and expanded the number of anatomical characters that de- fine the genus, most notablv a folded and thickened gill filament, a thickened muscular mantle edge, and the "strongly developed brain". Prashad (1928) subsequently synonymized Lecijthoconcha with Cipangopahidina but retained Lecijthoconcha as a subgenus. Furthermore, he reduced Idiopoma maUeata to a subspecies of V chinen- sis (Gray, 1817). Kuroda (1929) adopted all of Prashads (1928) revisions. Rohrbach (1937) primarily used char- acters of the reproductive anatomv to divide the vivi- parids into two subfamilies, the Vi\iparinae and a new subfamily Bellamyinae. Under the Bellamyinae he listed a single large genus, Bcllamija Jousseaume. Although considered to be a genus largelv limited to tropical spe- cies, Rohrbach (1937) was able to examine specimens of a race of non-tropical V. chinensis that displa\ed anatom- ical features of BcUaintja. Yen (1943) extended Rohr- bach s (1937) classification to Chinese species and sub- species, restateil the close relationship between Bella- mija and Cipangopahidina, but retiuned Cipangopahi- dina at the genus level. The first major work on North American \i\iparids following this period is Clench and Fuller's (1965) study in which, for unkTiown reasons, they treat Cipangopa- hidina as a subgenus of Vivipanis. These authors res- urrected the trinomen V chinensis malleatus for all North American populations of this species and listed it as a species separated from V' (Cipangopahidina) japon- iciis. Most subsequent studies on viviparid faunas (eg. Pace, 1973: Taylor, 1981; Rao, 1989) have considered Cipangopahidina a genus of the Bellamyinae. North American opinion has been di\aded concerning the \'alidit\' of the two introduced species (see Jokinen, Page 32 THE NAUTILUS. Vol. 114. No. 2 1991, for review). Many North American faunal studies including these viviparids have listed all large introduced \iviparid populations with unhanded shells as Cipango- pahidina chincnsis (eg. Robertson and Blakeslee, 1948; Jacobson and Emerson, 1961; Dundee, 1974; Clarke, 197S). Other reports have recognized the two species C chincnsis and C. japonica (eg. Wolfert and HUtunen, 1968; Burch, 1982; Tavlor, 1981; Jokinen, 1983, 1991; Smith, 1995). Certainly one of the problems affecting the species issue is the apparent rant)- of the ta.\on C. japonica. Clench and Fuller (1965) gave only four North Ameri- can records, and Taylor (1981) listed only one in Cali- fornia. Wolfert and Hiltunen (1968) listed an Ohio re- cord and Jokinen (1984) provided an additional record in Connecticut, but has before and since considered the population C. chincnsis or doubtfully C. Japonica (Joki- nen, 1983, pers. comm.). Jokinen (1991) subsequently added a New York record that would be the eighth pub- hshed North American record for C. japonica. The impetus for the present study came from the dis- covery of two wild, previously unreported populations of C. japonica-morph sn;iils in Connecticut from which se\'- eral variously aged specimens and reproductively active females were collected. The present study provides com- parative data on a number of characters of each species in an attempt to determine whether two species or a single variable species occurs in North America. Addi- tionally, using data from reproductive organs and other anatomical characters plus observations by previous au- thors on viviparid subgroups, a proposal to reduce Ci- pangopahidina to a subgenus ot BeUanu/a is presented. Hereafter in this study these two species will be referred to as Bellamija spp. MATERIALS AND METHODS Specimens from 12 extant or extirpated BcIIamt/a pop- ulations were examined. The original assignment ot specimens to either specific taxon was by use of adult shell characters. The distribution of specimens is listed below. All were fixed in 10% formalin, relaxed or unre- laxed, and stored intact in 50-60% isopropyl alcohol. In- tact shells of deceased animals were also collected. Each collection has been catalogued in the Invertebrate Di- vision, Museum of Zoology, University of Massachusetts, Amherst (UMA). Collection numbers and other details regarding the collections can be furnished upon request. BcUami/a chincnsis s.l: UMA 0668, University Pond, University of Massachusetts Campus, Amherst, Hamp- shire County; UMA 0885, unnamed pond in Medford, and UMA 1705, Lake Winthrop, Holhston, both Mid- dlesex County; UMA 1635, Forest Park Ponds, Spring- field, Hampden County; UMA 1663, Wachusett Reser- voir Outlet, Worcester County; UMA 1094, Purchase Louisiana Brook, Northfield, Frankhn County; UMA 1330, Pine Tree Brook, Milton, Norfolk County; UMA 1805, Snake River, Norton, Bristol Count>- (all Massa- chusetts); UMA 1755, Black Pond, Middlefield, Middle- sex County, Connecticut. BcIIamija japonica: UMA 1754, Lake Mohegan, Fairfield, F;iirfield County; UMA 1816, Connecticut Riv- er, Lviue, New London County; UMA 1755a, Black Pond, Middlefield, Middlesex County, all Connecticut. Also examined were specimens (UMA 1700) purchased from a pet store in Hadley, Hampshire County, Massa- chusetts. Samples ol intra-uterine juveniles were removed from females. The animal was removed from the shell of each juvenOe and the shell iiir dried. Radulae were also re- moved from adult specimens, cleaned in 10% potassium hydroxide, subsequently washed, and air dried. All dried material was mounted on individual Cambridge stubs with Duco cement and sputter-coated with gold. All pre- pared specimens were then examined with a JEOL Model JSM-5200 scanning electron microscope (SEM). For anatomiciil study, at least five specimens of each sex were dissected from the two largest collections of each species as determined by shell characters. There- after, dissections were made on at least one adult spec- imen of each sex (if available) from each remaining lo- cation. A total of 44 specimens divided almost evenly among both species were analyzed. Dissections were performed with the aid of a Wild M5 dissecting micro- scope. RESULTS The principal conchological characters distinguishing the tsvo taxa are the shape of the spire and the presence of carination; to quote from Clench and Fuller (1965) re- garding B. japonica: ". . .can be distinguished from V mallcatiis (= C. chinen.sis) by having a more acute spire and by having fine carina or carinae." An examination of animals from presumed B. japonica populations reveal that these characteristics are most evident in shells 35- 45 mm shell length (Figure 1). At this size, the species can be separated using the diagnosis of Clench and Ful- ler (1965). Below 35 mm shell length both species have a carina on the body whorl. Above 45 mm, the carina of B. japonica becomes nearly obsolete, and only a sfightly more acute spire in B. japonica enables identification. The source of differentiation in shell characters in these two species is in part derived from an allometric growth pattern first described by Jokinen (1982), who showed that shell width in B. japonica increases at a slower rate than in B. chincnsis. This phenomenon would account for the more "acute spire" in B. japonica described by Clench and Fuller (1965). The regression slopes for each species, however, are only slightly differ- ent (see Jokinen, 1982, fig. 2). Therefore, adult shell moi-phology has its limitations in the distinction of these two species. A characteristic that appears to be reliable in separat- ing B. japonica from B. chincnsis is the morphology of the intra-uterine juvenile shell. The juveniles occurring in the uteiTis or brood chamber, considered a functional D. G. Smith, 2()()() Page 3.3 Figure 1. Comparison of similar sized shells of Bdlamija. Upper and lower left, B. japonica from Lake Mohegan, Con- necticut. Upper right, B. chinensis from Medford, Mussachn- sets, and lower right, B chinensis from Black Pond, Connect- icut. Arrow denotes carina. Scale line = 10 mm. modification of what is otherwise called the pallial ovi- duct, demonstrate clear differences in the position of the embryonic whorl (protoconch) and in surface sculpture. Most viviparid snails in their juvenile stages have specific features of the periostracum, including the presence of periostracal hairs encircling the shell v\'horls, and surface sculpture (Fretter and Griiliam, 1962; Joldnen, 19(S4). In the juvenile shell of B. chinensis, the spire com- prises only two elevated whorls (Figures 2-A). The em- bryonic whorl is depressed in frontal view below the suc- cessive whorl (first teloconch whorl), characterized by two fines ot periostracal hairs, <3ne clearly evident on the whorl, the other along the upper border (Figure 2, ar- row). Such was observed in all examined populations. In contrast, the embryonic whorl of B. japonica (Figure 5) is somewhat elevated above the first teloconch whorl, indicated by two spiral fines of periostracal hairs, which in frontal view gives the shell the appearance of having four whorls. Differences in periostracum are even more eNddent than those in whorl development. The periostracum of the body whorl of juvenile B. chinensis (Figures 6, 7) is finely sculptured with a series of closely set striae, about 20/mm near the midpoint of the body whorl, interrupted by peqiencficular shallow grooves more widelv spaced than tlie striae. The spiral lines bearing the fine hairs are not raised as spiral carinae. In B. japonica (Figure 8), the surface of the periostracum is strongly marked by raised carinae, each having a series of periostracal hairs. Between the carinae are widely spaced spiral striae, about 8/mm near the middle of the whorl. Although oc- casional interiTiptions occur, there is no pattern of dis- tinct grooves as found in and characteristic of B. chinen- sis. Regarding anatomical characters, the radula provides no specific or cfistinctive characters for interspecific dif- ferentiation. Veiy subtle cfifferences (Figures 9, 10) oc- cur in the spacing of denticles of the lateral teeth for example, but these differences are probablv subject to variation and should not be considered as refiable taxo- nomic characters. The gill filament of each species con- sists of an elongate structure, measuring up to 5 mm length in the largest (>50 mm shell length) individuals. The filament is broadest at its base where it is attached to the mantle, narrowing in depth cfistally. A cfistinct blood vessel courses its length just below the thickened superior margin. The "fokfing" of the filament described by Rao (192.5) in a few species, and used subsequently as a character of Cipangopahulina. was not observed in material of either species. Major features of both the male and female repro- ducti\'e svstems demonstrate an overall conservative structural plan clearly characteristic of the Bellamyinae as defined by Rohrbach (1937). Dissection of the female reproductive system revealed no consistent differences between B. chinensis and B. japonica. Within the male reproductive system, the \'as deferens of each species shows distinct differences in its cfimensions and its re- lationship with both the columellar muscle and the tes- tes. In B. chinensis (Figure 11), the vas deferens arises from the testes as an unbranched tube. All further branching of the vas deferens occurs internally within the testes. Occasionally, a single large anterior branch merging with the vas deferens is partially visible. The emerged vas deferens widens considerablv as a complex duct and proceeds a short cfistance, suspended by mes- enteries, until it reaches the columellar muscle. At this point, the remaining portion of the vas deferens bends shaqiely downwards. It then abruptly and without en- largement enters the prostate. The downwardlv cfirected portion of the vas deferens remains free of the colu- mellar muscle although it is supported throughout by mesentaiy tissue. The vas deferens of B, japonica departs from the tes- tes multibranched, with at least three and usually four visible branches converging to a single duct (Figure 12). The duct then enlarges, but not to the degree seen in B. chincn.'iis, and extends a greater distance, suspended by mesentery tissue, until meeting with the columellar muscle where, as in B. chinensis, it bends downward and, without enlargement, joins the prostate gland. Un- Page 34 THE NAUTILUS, Vol. 114. No. 2 ij^gaiit«^Jgi*jfj BD m v^ m 3 - ■N "1 BD Figures 2-5. Spire of juvenile shells ot Bellnmya. 2-4. Bellamtja chinensis. 2. Black Pond, Connecticut. 3. Wachusett Reservoir, Massachusetts. 4. Campus pond, Amherst, Massachusetts. Figures 3, 4, apical whorls only. 5. Bcllami/a jnponica. Lake Mohegan, Connecticut. BD = hodv whorl. Scale line = 0.5 mm. hke B. chinensis, however, the portion of the vas defer- ens pro>diiial to the prostate gland is invested with col- umellar muscle tissue. Odier aspects of gross anatomy of the male reproductive system, including testes size and shape and prostate size and shape, are similar in the two species. The characteristics of the vas deferens in each species were most pronounced in the larger animals. The hvo patterns were concordant with shell characters in all but one case, the Black Pond locality. Both species occur in this pond and examined males show features of shell and reproductive anatomy consistent with above observations or variations of each. Most males from this collection, however, are young adults (<.'35 mm shell length) and, as stated above, the anatomical characters inchcative of each species are best developed in larger specimens. Nevertheless, it is possible that hybridization is occur- ring. Previously, in North America, both species have been reported in sympatiy (Lake Erie; Wolfert and Hil- tunen, 196S), but not in svmtopy so it is not known if hydridization is a usuiJ outcome of contact. DISCUSSION Although adult shell structure as presented by Clench and Fuller (1965) does not in and of itself provide con- \incing evidence for the recognition of two distinct spe- cies, the characters of the embrvonic shell and male re- proilucti\e system in combination with shell features in- dicate that at least Uvo similar but distinct species of Bcllami/a are present in North America. The question remains, however, regarding which names to assign these species; is enough information available to warrant use of the two specific epithets, mallcata and japonica as hsted by Hannibal (1911) soon after the species were first reported in North America? If so, then what is the the status of the name B. chinensis? Clench and Fuller (1965) gave a partial history of the D. G. Smith, 2000 Page 35 Figures 6-8. Penostracal surface of body whorl ol ju\eiiile Bellami/a^ 6. B. chinensis from Black Pond, Connecticwt. 7. Same, Forest Park ponds, Springfield, Massachusetts. 8. B. japonica from Lake Mohegan, Connecticut. Scale line = 0..5 mm. Figures 9- 10. Radula (central and lateral teeth). 9. Bilhiim/a japonica. 10. Bcllainija chinensis. Scale line = 0.05 mm. taxonomy and nomenclature of these species. As can be deduced from their account, the original descriptions of these species were essentially brief accounts of the adult shell with very little geographical data provided. Pace's (1973) review of the titxonomv of B. chinensis probably best summarizes the nomenclatural and taxonomic con- fusion that accompanies this species. Seemingly every race or form encountered by earlv workers was estab- lished on the most subtle of characters. Differences of opinion on the validity of named forms and generic as- signment was quite likely compounded by difficulties in communication between Asian and European investiga- tors. Unfortunately, the practice of naming new species of Asiatic viviparids on the basis of a few qualitative shell characters continues (Li, 1990; Liu ct ai, 1994). Until future study requires otherwise, I follow Pace (1973) and other earlier workers who assert that many of the sub- sequentlv named forms related to B. chinensis. including B. mallcata, are at most subspecies. Given that subspe- cific status is normal!)- maintained through geographic. Page 36 THE NAUTILUS, Vol. 114, No. 2 CM VD CM Figures 11-12. Anatomy of male reproductive structures. 11. Bellaimja chinensis. 12. Bellamya japonica. Scale applies to figures 11 and 12. CM = coliimellar muscle, CT = con- nective tissue (mesentery), F = fibers of columellar muscle, P = prostate, T = testes, VD = vas deferens. (1912) in his description, the foundation of Cipangppa- liidina hes chiefly in certain anatomical features dis- cussed above. Folding of the gill filament, a character of Cipangopahidina estabhshed by Rao (1925), was not ob- served in either species. Folding of the filament mav be caused by contraction or mav represent distortion as a result of preservation. A thick mantle edge and a devel- oped sphincter muscle are present in both species, but based on comparison with smaller Vivipants georgianus (I. Lea, 1834) thickness and muscle size can be related to animal size (Smith, pers. observ.). A comparison of mantle types illustrated by Annandale (1920: 112) and used to characterize Lecijthoconcha (= Cipangopahidi- na) apart from other genera more clearlv demonstrates a gradation of mantle development among the groups he discussed. In relation to brain comple.xity (sensu Rao, 1925), more detailed studies are necessary for the prop- er evaluation of this character. Following Rohrbach's (1937) anatomical svstem, which was somewhat extend- ed by Pace (1973), B. japonica represents a group within Bellamya that is characterized by a branched vas defer- ens proximal to the testes; Bellamya chinensis represents a sister group that has an unbranched tree portion of the vas deferens. Vail (1977) has addressed the problems of subgroups within Bellamya in a more general fashion and suggested that further studies are necessary to es- tablish natural groups within the Bellamyinae. It is thus proposed that Cipangopahidina should be treated as a subgenus of Bellamya, a decision with some precedent. Cipangopahidina would provisionally embrace the large bellamvids with unhanded shells during any part of their life historv and with native distributions fimited to Asia. ACKNOWLEDGMENTS I thank Eileen Jokinen for providing useful input and comment on various drafts of the paper. Kenneth Boss provided extensive information on the taxonomy and lit- erature of Asian bellamvids. not genetic, separation, and that several populations of interbreeding B. chinensis have been introduced into North America, the features that defined any particular subspecies have most likelv been obliterated. Further- more, in that most opinion leans towards a Japanese or- igin for at least some of the North American populations and that populations of each species have been reported in Japan (Kuroda, 1929), I believe that B. japonica s. I. and B chinensis s. I. are the species involved. The decision to place both species with Bellannia re- sults in part from Taylor's (1981) statement, substanti- ated by anatomical data pro\ided in this studv, that B. japonica belongs to the Bellamvinae. With respect to the continued use of the name Cipangopahidina lor 6, chi- nensis, those characters used to substantiate the genus are either variable or characteristic of animals reaching a relatively large size. Notwithstanding the absence of a carina on the adult shell, originally used by Hannibal LITERATURE CITED Annandale, N. 1920. Materials for a generic revision of the freshwater gastropod molluscs of the Indian Empire. Re- cords of the Indian Museum 19:107-115. Annandale, N. 1921. Zoological results of a tour in the Far East. The viviparous water-snail of Lake Biwa, Japan. Memoirs of the Asiatic Society, Bengal 6:399—401. Burch, J. B. 1982. The freshwater snails (Mollusca: Gastro- poda) of North America. Environmental Monitoring and Support Laboratory, U. S. 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Argonaut Books, Larchniont, 142 pp. Jokinen, E. H. 1982. Cipanoppaludina chinensis {Gastropoda: \'i\iparidae) in North America. The Nautilus 96:89-95. Jokinen, E. H. 1983. The Freshwater Snails of Connecticut. Bulletin Connecticut Geological and Natural History Sur- vey 109:1-83. Jokinen, E. H. 1984. Periostracal moqiholog)' of \i\iparid snail shells. Transactions of the American Microscopical Societv 103:312-316. Jokinen, E. H. 1991. The Freshwater Snails (Mollusca: Gas- tropoda) of New York State. Bulletin of the New York State Museum 482:1-112. Kuroda, T. 1929. Notes on the nomenclature of the Japanese species of the genus Vivipanis. The Venus 1:98-102. Li. Z. 1990. A new species of genus CApangopahuUna from Dianchi Lake in Yunnan Province, China. Acta Zootax- onomica Sinica 15:2.5-27. Liu, Y. Z., W. Zans, Y. Wane, and Y. Duan. 1994. Eight new species of freshwater molluscs in southeast China. Acta Zootaxonomica Sinica 19:2.5-36. Pace, G. L. 1973. The freshwater snails of Taiwan (Formosa). Malacological Review, Supplement 1:1-118. Prashad, B. 1928. Recent and fossil Viviparidae. Stud\- iu dis- tribution, evolution and Paleogeography. Memoirs ol the Indian Museuui 8:153-252. Rao, H. S. 1925. On the comparative anatomv of Oriental Vi- viparidae. Records of the Indian Museum 27:129-1.35. Rao. N. \. S. 1989. Handbook Fresliwater Molluscs of India. Zoological Survey ot India, Calcutta. 289 pp. Robertson, I. C. andC. L. Blakeslee. 1948. The Mollusca of the Niagara Frontier Region. Bulletin of the Buffalo So- ciety of 'Natural Sciences 19(3): 1-191. Rohrbach, F. 1937. Oekologische und morphologische Unter- suchungen an Vivipanis (Bellarmja) ca))illntHs Franenfeld und Vivipanis {Brllamt/a) unicolor 01i\er, unter Benick- sichtigung anderer tropischer F'ormen luid im Hinblick auf phvletische Beziehungen. Archi\- fur Molluskenkunde 69:177-218. Smith, D. G. 1995. Keys to the Freshwater Macroinvertebrates of Massachusetts. Second Edition. University of Ma.ssa- chusetts, Amherst, 243 pp. Taylor, D. W. 1981. Freshwater mollusks of California: a dis- tributional checklist. California Fish and Game 67:140- 163. Vail, V. A. 1977. Comparative reproductive anatomy of 3 \ivi- parid gastropods. Malacologia 16:519-540. Wolfert, D. R. and J. K. Hiltunen. 1968. Distribution and abundance of the Japanese snail, Vivipanis japoniciis and associated macrobenthos in Sandusky Bay, Ohio. Ohio Journal of Science 68:.32— 40 Yen, T. 1943. A preliminary revision ot the Recent species of Chinese Viviparidae. The Nautilus 56:124-130. THE NAUTILUS 114(2):38-58, 2000 Page 38 A revision of the Patagonian genus Xijryienopsis Powell, 1951 (Gastropoda: Muricidae) Cuido Pastorino Museo Argentino de Ciencias Natiirales Av. Angel G;illardo 470, 3° piso, lab. 57 CUOsbjR Bvienos Aires ARGENTINA rvpastor@criba.edu . ar M. G. Harasewych Department of Inxertebrate Zoology National Museum of Natural History Smithsonian Institution Washmgton. D.C. 20560-0118 USA Harasewych@nmnh.si.edu ABSTRACT The genus Xt/menopsi.s Powell, 1951 is re\ised and restricted to 4 Recent species: Xtjmenopsis miiriciformis (King and Brod- erip, 1832), X. comigatus (Reeve, 1848), X. huccineus (La- marck, 1816) and X. suhnodosus (Gray, 1839). Of the 34 nom- inal species that have been described as or assigned to Xtjmen- opsis by various authors, 3 belong to the buccinid genus Pa- reuthria and I is a trophonine nuiricid not closely related to Xymenopsis. The ta.xononiic affinities of each of the remaining .30 names (one is a replacement name) are discussed. Ele\en holot)pes and ten lectotypes (nine newly designated) are illus- trated, many for the first time. Tvpes of .3 taxa were destroyed during World War II and another 5 t)pes are lost. Neotvpes are designated for Fusus huccineus Lamarck, 1816, the oldest name to be applied to any species of Xymenopsis, as well as for Buccinum subnodosa Gray, 1839, Trophon brucei Strebel, 1904, Trophon fcnestratus Strebel, 1904, Trophon paessleri tur- rita Strebel, 1904, Trophon ringci Strebel, 1904 and Trophon standcni Strebel, 1904. This is done to remo\e anv ambiguity regarding the identity of these ta.\a, and thus stabilize the no- menclature of Recent species of Xymenopsis. The gross anat- omy and moqjhology of the radula, egg capsule, protoconch, and shell ultrastructure of the type species of Xymenopsis are described and illustrated. Xymenopsis is compared to, and dif- ferentiated from, the genera Xynwne. which is restricted to New Zealand, and Trophon. with which it co-occurs in the Magellanic province. Key words: Neogastropoda, Trophoninae, Patagonia, Magel- lanic Province. INTRODUCTION The genus Xymenopsis encompasses a group of small, conchologicaUy variable muricid gastropods endemic to the Magellanic Prox-ince. These shell-drilling predators commonly co-occur wdth species of Trophon on mussel beds, and range from the intertidal zone to depths ex- ceeding 100 meters. The majority of the published Bt- erature on this group dates from the early era of Ant- arctic and subAntarctic exploration, and has concentrat- ed on describing numerous, minor phenotypic variants as species. The biologv' ot these animals has not previ- ously been stuched. Powell (1951) erected the genus Xymenopsis to dis- tinguish Patagonian muricids with a conical, multi- whorled protoconch and rounded ;mal ribs from those with a rounded, paucispiral protoconch and numerous, lamellose varices that belong to the genus Trophon. He attributed 30 nominal taxa to Xymenopsis, but recog- nized that many would prove to be synonyms, stating that "An evaluation of these names is not possible with- out recourse to the tyj^e material which is in European museums." Imphcit in the generic name Xymenopsis is Powell's recognition of the close affinity of this Magel- lanic group to Xymenc Iredale, 1915, which is endemic to New Zealand. Powell (1951) noted that the shell sculpture of Xymenopsis was similar to that of Xyniene, but that its protoconch resembled that of Zcatroplwn Finlay, 1927, another New Zealand genus. In his revision of New Zealand muricids assigned to the genus Trophon, Finlay (1926) proposed several su- praspecific taxa, each representing what he regarded to be a hneage or "natural grouping." He divided these taxa into 2 groups, based primarily on protoconch morphol- ogy. One group, defined on the basis of having a sym- metrically conical protoconch of > 2 whorls and a small nucleus, contained the genera Xymenc Iredale, 1915, XymcncUa Finlav, 1926, and Zcatrophon Finlaw 1927. The second group, recognized by having an asymmetri- cal, rounded protoconch of 1-2 whorls, with a large, lat- eral nucleus, included Axymene .scvi.si/ stricto Finlay, 1926, Axymcnc (Lcnitrophon) Finlay, 1926, Comptella Finlay, 1926, Tcrcfundus scnsit stiicfo Finlay, 1926, and Terefiindus iMinoiirophon) Finlay, 1926. Ponder (1972) rexdewed the Recent and fossil New Zealand species related to Xymenc. Based on an analysis of gross anatomy, shell, and radular moqihology, he con- cluded that the taxa Axymene, Lenitrophon, XymeneUo, Zcafroplion, and probably Vc.5flri(//fl Finlay, 1926, did not merit taxonomic recognition even at the subgeneric lev- el, and reduced all but Vesatuda to .svmonyms oi Xymenc. However, he regarded Xymenopsis sufficiently distinct to G. Pastorino and M. G. Harasewvch, 2()()0 Page 39 Table 1. Tiixa attributed to the genus Xijiiu'iiopsis by Powell (1951) and Dell (1971, 1972). Taxa listed ni bold are valid species. Troplion iiciiininntus Strebel, 1904 Trcphon (dints Strebel, 1904 Fiisiis cilhithis Philippi, 1845 Tniplioti bnitti Strebel, 1904 Xymenopsis buccineus (Lamarck, 1816) Troplion canccllarioidi's Ree\e, 1847 Xymenopsis canccllinns (Philippi, 1S45) Fiisiis candidatiis Rochebnnie and Mabille, 1889 Xymenopsis corrttgattts (Reeve, 1S4S) Troplion coiithoiii/i Strebel, 1904 Fiisiis decolor Philippi, 1845 Fiisiis dispar Rochebnine and Mabille, 1889 Troplion elcfians Strebel, 1904 Troplion elongatiis Strebel, 1904 Trojihon fnlklandiciis Strebel, 1908 Troplion fenestratiis Strebel, 1904 Troplion hoijlei Strebel, 1904 Fiisns jarqiiinoti Philippi, 18.55 Fiisiis lebnini Mabille and RochebiTnie, 1889 Fiisiis liratiis Gould, 1849 Fiisiis loebbeekei Kobelt, 1878 Xymenopsis muriciformis (King and Broderip, 1832) Troplion obesiis Strebel, 1904 Troplion ornatiis Strebel, 1904 Trophon paessleri Strebel, 1904 Troplion paessleri turrita Strebel, 1904 Fiisiis pliimbeiis Gould, 19.52 Trophon pseiidoelonoatiis Strebel, 1904 Trophon ringei Strebel, 1904 Fiisiis roseiis Honibron and Jacquinot, 1854 Trophon standeni Strebel, 1904 Xymenopsis subnodosus (Gray, 1839) Fiisiis textdiosiis Honibron and Jacquinot, 1854 Fiisiis violaceiis Mabille and Rochebnnie, 1889 (syiiony-m of Xipnenopsis nuiricifoniiis < (synonym of Xiimenopsis iniirieifonnis) (synonyiii ol Xipm-nopsis hiiriineiis) (.synonym ol Xiinuiu>psis rornio/itiis) (synonym ot Xt/menopsis iniiririfonnis) (synonym of Xymenopsis subnodosus) (belongs in the buccinid genus Fareuthria Strebel, 1905) (synonym of Xynwnopsis muririfonnis) (synonym of Xymenopsis muririfonnis) (belongs to different niuricid genus, not Xymenopsis) (synonym ot Xymenopsis miiricifonnis) (synonym of Xymenojisis muriciformis) (synonym o( Xymenopsis corrugatiis) (synonym of Xymenopsis muriciformis) (synonym of Xymenopsis cornigatus) (new name for F. textiliosus Honibron and Jacquinot. 1854, not F. textiliosus Deshayes. 18.35) (synonym of Xymenopsis miiricifonnis) (syiionyiii of Xymenopsis miiricifonnis) (sviionym of Xymenopsis muriciformis) (synonym of Xymenopsis miiricifonnis) (synonym of Xymenopsis cornigatus) (synonym of Xymenopsis muriciformis) (synonym of Xymenopsis muriciformis) (belongs in the buccinid genus Fareuthria Strebel, 1905) (synonym of Xymenopsis nuiriciformis) (synonym of Xymenopsis buccineus) (belongs in the buccinid genus Pareiithria Strebel, 1905) (synonym of Xymenopsis cornigatus) (synonym of Xymenopsis buccineus) (synonym of Xymenopsis miiricifonnis) be treated as a subgenus of Xijmene. Despite reducing their taxonomic rank. Ponder (1972) recognized 5 dis- tinctive groups within the Xijmenc complex, correspond- ing to: 1) Xymene + Xijinenclla; 2) Zcatrophon; 3) a group consisting of 2 fossil species (1 Eocene, 1 Mio- cene) with features expected in the "ancestor of all the species of Xymene"; 4) Axymciw; and 5) Xi/menopsis. Beu and Maxwell (1990:39) reported Xymene to range from the Kaiatan (= Bartonian, Late Eocene) to the Recent in New Zealand. The fossil record of Xymenopsis is thus far limited to 4 species recently described from the Late Miocene Entrerriense Formation along the Val- des Peninsula in northern Patagonia, Argentina (Bninet, 1997). While there is little doubt that Xymene and Xy- menopsis are closely related, these lineages are treated here as separate genera. In the present study, we review and discuss all Recent nominal species that have been included in Xymenopsis (Table 1) based on an examination of available type ma- terial. Type specimens of all taxa referable to Xt/men- opsis that could be located are illustratetl, some for the first time. Data on shell ultrastructure, radular mor- phology, and gross anatomy are provided for Xymenopsis miiricifonnis, the type species oi Xymenopsis. Compar- ative data are provided for Xymene plcbeiiis, the type species of Xymene, and for the other Recent species of Xymenopsis. MATERIALS AND METHODS The majority of specimens examined in this study are in the collections of: the National Museum of Natural His- tory, Smithsonian Institution, Washington, DC (USNM); Museo de La Plata, La Plata, Argentina (MLP); Museo Argentino de Ciencias Naturales "Bernardino Rivada- via", Buenos Aires (MACN); and the Los Angeles Coun- ty Museum of Natural History (LACM). Type material is housed in the following museimis: The Natural His- tory Museum, London, (BMNH); Zoologisches Institut und Zoologisches Museum der Universitat Hamburg, (ZMH); Swedish Museum of Natural History, Stock- holm, (NHRM); Museum national d'Histoire naturelle, Paris (MNHN); Museo Nacional de Historia Natural, Santiago, Chile (MNHNS); and, Loebbecke- Museum und Aquazoo, Dusseldorf, (LM). Dissections were performed on ethanol preserved Page 40 THE NAUTILUS, Vol. 114. No. 2 G. Pastorino aiul M. G. Harasewych, 2000 Page 41 specimens to studv their gross anatomy, with emphasis on the nioiphologN' of the anterior alimentary system, and the palli;il portions ol the male ami female repro- ductiye systems. Radulae were prepared according to the method described by Solem (1972) and observed using a LEO 440 scanning electron microscope (SEM). Radular terniinolog) follows Kool (199.3a: fig. 6B). ,Shell ultrastnicture data were procured from freshly fractured collabral sections taken from the central portion of the lip on the last whorl of 2 individuals per taxon, whenever sufficient material was available. Most photographs were taken using a Leaf Lumina digital scanning camera. Several images were scanned from black and white .35 mm negatives using a Nikon Coolscan III slide scanner. All images were processed with the software Photoshop 4.01 and 5.02. SYSTEM ATICS Class Gastropoda Cuvier, 1797 Order Neogastropoda Wenz, 1938 Family Muricidae Rafinesque, 1815 Subfamily Trophoninae Cossniann, 1903 Genus Xymenopsis Powell, 1951 Xijinciiopsis Powell, 1951:1.58; Radwiii and D'Attilio, 1976:190. Xijnwne iXtjmenopsis) Ponder, 1972:474; Yokes, 1991:10. Type species: Ftistts Uratiis Gould, 1.S49 (by original designa- tion), a junior synonym of Bticciiiuin iniiricifonne King and Broderip, 18.32. ' Diagnosis: The genus Xi/incuopsis contains species characterized by a small to medium-sized (to 48 mm), fusiform, high-spired shell with a tall, conical, multi- whorled (==3 whorls) protoconch that lacks a keel, and a teleoconch with a rounded shoulder, sculpture of rounded axial ribs that never develop lamellae, and nar- row to broad spiral cords that may or may not produce beads at intersections with a.xial ribs. The siphonal canal is short, open, and broad. Shell ultrastructure consists of an outer amorphous calcific layer and an inner, crossed- lamellar aragonitic layer. The rachichan tooth of the rad- ula has a basal plate that is simple and concave anteri- orly, and inner lateral denticles that are completely free from the central and lateral cusps. The mantle edge is always lobulated. Remarks: In his treatment of the Xi/mcne complex. Ponder (1972) identified the oldest known member of the group in the Late Eocene fauna of South Island, New Zealand. He recognized that Xi/inciw apipagpdus Ponder, 1972, characterized by a small, strongly shoul- dered shell with ;L\ial lamellae, smooth aperture, and a tall, conical, 4y2-whorled protoconch with a sinuated var- L\ indicative of a pelagic larval stage, had features from which all subsefjuent Xi/mene lineages could be derived, and suggested that several lineages evolved from this phenotype. One hneage compri.ses the Zcatrophoti group, which appeared during the Late Oligocene {Xtfiiwrw chatto- ncnsis Ponder, 1972, Duntroonian) and persists in the Recent fauna (e.g., Zcafrophou amhitnius Finlay, 1927, the type species oi Zcatwphon). This hneage is charac- terized l)y a tall, strongly shouldered shell with axial nodes or weak lamellae, an aperture with or without lir- ae, and a protoconch that is tall and conical, yet smaller and with fewer (2%-3V4) whorls than X apipagodns. Within this hneage, there is a trend for a spiral keel to develop on the last protoconch whorl, and for the nu- cleus of the protoconch to increase in size. A second lineage iXijmcnc + Xymcnclla group) also appeared in the Late OUgocene (Trophon lepidiis Suter, 1917, Dun- troonian) and survives in the Recent fauna (e.g., Ftistis pk'bciiis Hntton, 1873, the tvpe species of Xijmcnc, and Trophon pii.sillns Suter, 1917, the type species of X(/- iiicncUa). Shells are small, strongly to weakly shouldered, with strong spiral cords and axial ribs, but without la- mellae, and with apertures that are usually lirate. Pro- toconchs are depressed, with a trend to fewer (2 whorls in X. plchcuis, 2 Vi whorls in X. gouldi = T. piisiUus, see Ponder, 1972), more inflated whorls, and an increase in the size of the nucleus. Another hneage (Axipnene group) is limited to Pliocene and Recent species. Shells tend to be large, strongly shouldered, with axial rilis and spiral cords, but without lamellae, and generally have lirate apertiues. Protoconchs consist of 1-2 roimded, in- flated whorls, with a large nucleus. Ben et al. (1997) regarded Xifmcnc and Xymcnclla to be distinct genera, and reassigned Triton dauizcnhcrgi von Ihering, 1897, and Urosalpinx elegans Ortmann, 1900, both from the Monte Leon Formation (Late Oli- gocene-Early Miocene) of Patagonia, to Xymcnc. and Xymcnella respectively. Based on the first occurrences of these taxa in the fossil record, they concluded that Xymcnclla originated in New Zealand and dispersed to South America with the origin of the Antarctic Circum- polar Current at the time the Drake Passage opened (Late Oligocene), while Xymene evolved in South Amer- ica and dispersed to New Zealand, also along the Ant- Figures 1-19. Xi/incnopsis luiiricifonnis (King and Broderip, 18.32). 1-3. Holotvpe of Fiisus liratiis Gould, 1S49, USNM 5680, Orange Harbor, Tierra del Fuego. 4-6. HoJotype of Trophon loebheckei Kobelt, 1878, Loebbecke Museum unnumbered, lacks locality data. 7-8. Lectot\pe of Trophon violacetis Mabille and Rochebmne, 1889, MNHN unnumbered, Baie Orange, Tierra del Fuego. 9-10. Holotype of Fusiis decolor Philippi, 1845, MNHNS unnumbered. Strait of Magellan. 11-12. Lectotype of Trophon elegans Strebel, 1904, ZMH unnumbered. Port Stanley, Malvinas (Falkland) Is. 13. Leetotype of Trophon lebnini Mabille and Rochebrune, 1889. MNHN unnumbered, Santa Cruz, Argentina. 14-16. Leetotype of Biiccimim niuricifonne King and Broderip. 1832, BMNH 1837.12.15.2695, Strait of Magellan. 17-19. Leetotype as Bnrciniim cnncrUnrioidcs Reeve, 1847. BMNH 19920.55, lacks locality data. Scale bar = 1 cm for all specimens. Page 42 THE NAUTILUS, Vol. 114, No. 2 G. Pastorino and M. G. Harasewvch, 2000 Page 43 arctic Circunipolar (^iinciil, diiriiit;; (lie Middle Mio- cene. The origin of the Xt/incitc complex in New Zealand is amplv supported In tlic fossil record. The hvpothesized dispersal ot this liiu'age to Southern South America dur- ing the Late Oligocene is concordant with its appearance in the fossil record of the region, and is supported bv the prexalence of narrow, polvgyrate protoconchs, inchc- ative of pelagic lanal stage (see Thorson, 19.50; Shuto, 1974; Jablonsk)- and Lutz, 1980), in Paleogene members of this complex. Recent and fossil species of Xijmenopsis are most similar to members of the Xt/iiicuc + Xi/inc- nclla group, especially to early members of this lineage such as the Oligocene Xi/mcnc Icpichis. with which they share a sculptural pattern of axial ribs and spiral cords, but not axial lamellae, and a conical, .3 whorled proto- conch with a small, apical nucleus. We suggest that Xi/- meiwpsis represents a radiation endemic to the Magel- lanic region from ancestors of the Xijmcne + Xi/iucnclla group that dispersed as larvae to Patagonia tluring the Late Oligocene. While it is possible that multiple lineages were intro- duced to South America, or that 1 lineage subsequently dispersed back to New Zealand while becoming extinct in South America, evidence for such hypothe.ses is more tentative or lacking. Xi/mcnc are characterized bv a trend to protoconchs with fewer, more roimded whorls that are indicative of lecithotrophic rather than planktotroph- ic development. This would argue against their abihty to disperse as plank-tonic laivae from South America east- ward on the Antarctic Circunipolar Current to New Zea- land. Clearly, a rigorous re-assessment of the Xymene complex in a phylogenetic context would resolve many of the systematic and biogeographic questions that have plagued this group. Xijincnopsis lack the strongly shouldered shell of the Xymene + Xymenella group. While strong apertural lirae are prevalent in Xi/mcnc, they are rare in Xymcuopsis. occurring only in the X. .siihnodosus (Gray, 1839). X(/- mcnop.sis may also be distinguished from Xtpticitc on the basis of shell ultrastiiicture, radular moiphology and by features of their mantle edge and egg capsules. Xymen- opsis (figures 45, 79, 80) has a thick (> Vi shell thick- ness) outer layer of calcite, and a single layer of crossed- lamellar aragonite in which the crystal faces are comar- ginal. In Xymene (figure 50), the calcitic layer is thin, the co-marginal layer of aragonite thickest, and an in- nermost layer of crossed-lamellar aragonite, with civstal faces perpendicular to those of the co-marginal layer, is also pre.sent. The radula i)^ Xymene pleheiiis has rachi- dian teeth (figure 51) that ari> characteristically chevron- shaped, eac'h with its apex at the mid-point of the an- terior (>dge of the basal plate, which has a concave pos- terior edge. Rachidian teeth oi' Xymcuopsis (figures 48- 49, 75-76, 81-82, 102-103) have a straight to sfightly concave anterior edge, and a convex posterior edge of the basal plate. The mantle edge of Xymene is smooth, while the mantle edge oi' Xymenopsis is lobulated. While egg capsules ui' Xymene (figure 43) and Xymenopsis (fig- ure 42) are similar, the exit aperture is apical in Xymene, Init is situated at the base of a shallow depression in Xymenopsis. There are no direct observations on development in Xymenopsis. Shuto (1974) reported that the ratio of the protoconch chameter (D) to the number of whorls (V) is a reasonable estimator of the t\pe of development, and that a ratio below 0.3 is characteristic of plankto- trophic larvae, especially when the number of whorls is s 3. Both Xymenopsis muriciformis (figure 41) and X. huecineus (figures 77-78) have narrow, conical proto- conchs of 3 whorls. The DA' ratios for these species are 0.23 and 0.21 respectively, predicting that, unlike Xy- mene, Xymenopsis retains planktotrophic larval devel- opment. Powell (1951) provisionally included the species Fusus rosens Hombron and Jac(juinot, 1854 and Trophon ean- didatiis RochebiTine and Mabille, 1889 in his genus Xy- menopsis. Ceniohorsky (1977) subsequently transferred Fusus ro.seus to the buccinid genus Pareuthria Strebel, 1905. He proposed the new name Pareuthria powelli for this species because Fusus roseus Hombron and Jac- quinot, 1854 was preoccupied by Fusus roseus Anton, 1838. The type material of these taxa was examined by the senior author at the Museum national d'Histoire na- turelle, and both are clearly referable to the buccinid genus Pareuthria. The type material of Fusus dispar Hombron and Jacquinot, 1854 was also examined (MNHN). This taxon has affinities with the Antarctic Trophoninae, but does not belong to the genus Xymen- opsis. Xymenopsis muriciformis (King and Broderip, 1832) (Figures 1-42, 45-49, 83-95, 105-106) Biicciniiin miiricifonne King and Broderip, 18.32:.348. Fusus muricifonuis King and Broderip — Gray, 18.39:118. Fusus huecineus Sowerby in Gray. 18.39:15.5, pi. .36, fig. 12. Fusus decolor Pfiilippi, 1845:68; 1846:118, pi. 3, fig. 3; Hupe Figures 20-40. Xymenopsis muriciformis (King and Broderip, 1832). 20-22. Holot\pe of Trophon ncuniinntus Strebel. 1904, ZMH unnumbered. Punta Arenas. Strait of Magellan. 23-25. Lectotype of Trophon couthoui/i Strebel, 1904, ZMH unnumbered. Port Grappler, Smyth Gliannel. 26-28. Lectotype of Trophon paessleri Strebel, 1904. (specimen coated with ammonium cliloride), ZMH unnumbered. Port Grappler, Sm\th Channel, 29-.31. Holotvpe oi'TropIion pscudoehinaatus Strebel. 1904, ZMH unnumbered, Ushuaia. 32-34. Lectotype olTroplion ohciiis Strebel, 1904, ZMH unnumbered, Punta Arenas, Chile. 35-37. Leetot^-pe of Troplwn elongatus Strebel, 1904, ZMH unnumbered, Puerto Angosto. Strait of Magellan. 38-40. Holotyjie of Trophon albus Strebel, 1904. ZMH unnumbered, Ushuaia. Scale bar = 1 cm for all specimens. Page 44 THE NAUTILUS, Vol. 114, No. 2 G. Pastorino and M. G. Harasewych, 2000 Page 45 in Gav, 1854;162; Homhroii and |a<.(|uin()t in Rousseau, 1854:i08, pi. 2.5, figs. 6-8; Kolx-lt', 1878:291, pi. 74, figs. 5-6; Rochebrune aiid Mabille, 1889; H.54. Buccinum cancellarioicles T{ee\e, lS47:pl. 14, fig. 11.3. Fusiis liratus Gould, 1849:141. Trophon liratus Couthous- — Kobelt, 1878:.31(); Tryon, 1880: 14.3, pi. 31, fig. 3.33; Watson 1886:165; Lamv. 1906:3. Trophon Locbbcckci Kobelt, 1878:294, pi. 74, 'figs. 13, 14; TnoH, 1880:145, pi. 31. fig. .3.35. Trophon nuiricifonnis King — Tnon. 1880:145. Trophon Icbnini Mabille and Rochebnnie in Rochebnine and Mabille, 1889: H..55. Trophon violaceus Mabille and Rochebnine in Rochebnine and Mabille, 1889: H.56, pi. 2, fig. 1. Trophon paessleri Strebel, 1904:213. pi. 7, figs. 56a-f. Trophon paessleri \'ar. turrita Strebel, 1904:215, pi. 7, fig. 57. Trophon elon^atus Strebel, 1904:217, pi. 7, figs. .58a-f. Trophon pseudoelong,atiis Strebel, 1904:220, pi. 7, figs. 60a,b. Trophon albus Strebel, 1904:221, pi. 7, fig. 61. Trophon acuminatus Strebel, 1904:222, pi. 7, fig. 63. Trophon obesus Strebel, 1904:223, pi. 7, fig. 62. Trophon fenestrattis Strebel, 1904:225, pi. 7, fig. 59a-d. Trophon couthoinji Strebel, 1904:236, pi. 7, fig. 65a-e; Pi. 8, fig. 76. Trophon elegans Strebel, 1904:241, pi. 8, fig. 71. Xt/menopsis hratiis (Gould) — Powell, 1951:158; Radwin and D'Attilio, 1976:190, fig. 1.36. Xijmenopsis decolor (Philippi) — Dell. 1971:211. Xtjmenopsis muricifonnis (King and Brodenp) — Dell, 1972:8, figs. 31, .32; Yokes, 1991: 7, unnumbered fig.; Yokes, 1992: 3, figs. 8a-c; Castellanos and Landoni, 1993:16, pi. 3, figs. .39, 40C, D. Diagnosis: Axial sculpture of irregular, rounded ribs, 16-20 on body whorl. Spiral sculpture of flat cords (4— 6 on early whorls, 14-20 on body whorl), much broader than intervening spaces, which appear as incised fur- rows. Description: Shell large tor genus (reaching 48 mm), fusiform, slender, chalky. Protoconch of 3 tall, conical, whorls. Teleoconch of up to 6 weakly shouldered or rounded whorls. Spire high, = V2 shell length, spire an- gle ~ 45°. Suture deeply impressed; subsutural ramp narrow in early whorls, then absent. Aperture subovoi- dal; outer lip rounded, anterior half-reflected; inner Up curved, adpressed; interior glossy pinkish to brownish. Siphonal canal short (< V2 aperture length), obhcjue, open. Umbilicus absent. Axial sculpture of irregular, rounded ribs, 16-20 on body whorl. Spiral sculpture of flat cords (4-6 on early whorls, 14-20 on body whorl), much broader than intei-v'ening spaces, which appear as incised furrows. (Growth lines irregular, faint to obsolete throughout shell. Shell composed of 2 layers: outer layer thick (0.6.5 slicll thickness), of amoqihoiis calcitc; inner layer (0..35 shell thickness), of crossed-lamellar aragouite with crystal planes oriented perpendicular to growing edge. Operculum (figure 47) oval, subpolvgonal, v\ath terminal nucleus. External surface covered bv irregular growth hues. Inner surface with .3—4 horseshoe-sliapt'd lines; posterior rim with slight callus, glazed in adult and young specimens. Aniiual large. Gephalic tentacles medium in size, blunt, thick, with small black eyes. Mantle edge with flat, rounded lobules (figure S3, 1). Siphon (figure 83, s) of medium length. Accessory boring organ (figure 95, abo) situated along ventral midline of foot, shares opening with ventral pedal gland in females. Osphradium (figure 84, os) < V2 ctenidium length, thin, veiy asymmetrical, with 45-50 leaflets per side. Ctenidium (figure 84, ct) 3-4 times as wide as osphradium, with 120-135 trian- gular leaflets. Pleuroembolic proboscis very long, broad. Radular ribbon (figures 48^9) small, short (0.31 X ap- erture length vs. 0.78 X aperture length in Trophon gev- ersianus), extends just beyond rear of buccal mass (fig- ure 94, ra). Rachidian tooth wide (to 230|xm), central cusp large, needle-shaped, lateral cusps outwardly point- ed, shorter than central cusp, inner lateral denticle smaU, thin, separate from lateral cusps. Basal plate of rachidian tooth rectilinear or gently curved, with low, broad, marginal areas. Lateral teeth large, each with sin- gle, long, scythe-shaped cusp and broad basal plate. Sal- ivary glands (figure 94, sg), ascinous, white (yeUow in fixed animals) veiy large, flanking retracted proboscis. Salivary ducts join esophagus just anterior to valve of Leiblein (figure 94, vl). Accessory sahvars' glands (figure 94, asg) small, pyriform, white to brownish, situated be- low salivary glands. Glande framboisee absent. Gland of Leiblein (figure 94, gl) large, broad anteriorlv, tapering posteriorly to form long, sinuous, blind duct with small, terminal ampulla (figure 94, dgl). Anterior aorta (figure 94, aa) passes through fold in gland of Leiblein en route to buccal mass. Stomach (figure 94, sto, 89) U-shaped, small, almost completely enveloped bv digestive gland. Both digestive gland ducts (figure 89, dd) near esopha- geal opening, situated in deep pouch (figure 89, dp), separated from posterior mbdng area (figure 89, pma) by small horizontal fold (figure 89, hf). Two tyj^hlosoles (fig- Figures 41—42, 45-49. Xijmenopsis muricifonnis (King and Broderlp, 1832). 41. Protoconch, USNM 901629, Strait of Magellan, 53°.39.4'S-70°55.5'W, in 24 m. Scale bar = 100 (xm. 42. Apical and lateral views of the egg capsule. Scale bar = 800 [im. 45. Shell ultrastructure, fracture surface comarginal. Scale bar = 30 \x.m. 46. Penis, critical-point dried. Scale bar = 40 |xni. 47. Operculum, external (left) and internal (right) \iews. Scale bar = 1 cm. 48-49. Radula. USNM 9016.35, Strait of Magellan, 5.3°39.8'S-70°54.9'W, in 27-73 m. 48. Dorsal \new of radular ribbon. Scale bar = 40 fxm. 49. Lateral \iew of rachidian teeth. Scale bar = 20 (jim. Figures 43-44, 50-51. Xi/ineiw picbeius (Hutton, 1873). 43. Lateral and apical \iews of egg capsule, USNM 6815.38, Spit Beach, Dunedin Harbor, New Zealand. Scale bar = 1 mm. 44. Operculum, external (left) and internal (right) \iews, same specimen as 43. Scale bar = 1 cm. 50. Shell iiltrastnicture, fracture surface comarginal. AK 80295, Northland, Bay of Islands, Parekuta Bay, New Zealand. Scale bar = 100 |xm. 51. Dorsal view of radular ribbon, same specimen as 43. Scale bar = 20 (jim. Page 46 THE NAUTILUS. Vol. 114, No. 2 Figures 52—36. Xi/menopsis bticcinetis (Lamarck, 1816). 52-54. Neotvpe of Fiistis biiccincus Lamarck, L816. USNM 870410, 5.3°.32'S, 64°.57'\\', in 119-124 m, 55-56. Holohpe of Ftistis texiiliostis Hombroii and |ac(juinot, 1854, MNHN unnumbered. Strait of Magelkui, Figures 57-71. Xi/rnenopsis comioatus (Reeve, 1848). 57-59. Holot\pe of Fusus cornigatiis Reeve, BMNH 1874. 12. 11. 15.3, lacks localits data. 60-62. Lectohpe of Trophon hoijlei Strebel, 1904, ZMH mmumbered. Port Stanlev, .VlaKinas (Falkland) Is. 63-65. Holotvpe of Trophon falklandicus Strebel, 1908, NHRM 1049, 5r35'S-57°56'\\', Islas MaMnas '(Falkland). 66-68. USNM 368350, Port' Stanley, MaKinas (Falkland) Is. 69-71. Holotvpe of Trophon ornatns Strebel, 1904, ZMH unnumbered, Port Stanley, Malvinas (Falkland) Is. Figures 72-74. Xymrne plebeius (Hiitton, 1873). USNM 681538. Spit Beach, Dunedin Harbor, New Zealand. Scale bar = 1 cm for all specimens. G. Pastorino and M. G. Harasew\'ch, 2000 Page 47 Figures 75-79. Xi/iiii-nopsis biiccinciis (Lamarck, 1S16). 75. Dorsal view ol radnlar nbhoii from specimen in figures 52-54. Scale bar = 40 p-ni. 76. Oblique (45°) lateral Wew, of radular ribbon, MACN 12553, Tierra del Fuego. Scale bar = 40 |jim. 77-78. Protoconch, MACN 25012, 54°26'S-64°53'W, in 112 m. 77. Apical, and 78. lateral views. Scale bars = ISO jxm. 79. Shell ultra- structure, fracture surface comarginal. Scale bar = 40 |j.m. Figures 80-82. Xi/menopsis comigatus (Reeve, 1848). 80. Shell ultrastnicture, fracture surface comarginal. USNM 368350, Port Stanlev, MaKinas (Falkland) Is. Scale bar = 40 |j,m. 81-82. Radula. USNM 421884, Port W'iUiams, York Bay, MaKinas (Falkland) Is. 81. Dorsal, and 82. obliijue lateral view. Scale bars = 20 (xm. Page 48 THE NAUTILUS, Vol. 114, No. 2 G. Pastorino and M. G. Harasewvcli, 2{)0() Page 49 nre S9, tl, t2) flank' intestinal i^rooN-e (figure 89, ig) sep- arating it ironi expantled eonipacting area (figure 89, ea). Intestine (figures 89, 94, i) runs anteriorly alongside pal- lial gonoduct, forming rectum with cylindrical papilla (figin-es 86, 90 ap) o\'er anus (figiu'es 86, 90 a). Rectal gland (figure 93, rg) small, inconspicuous, confined to anterior region ot rectum. Prostate gland (figures 86-88, pg) rvms alongside rectum in mantle cavity, lacking open- ings to mantle cavitv" (unlike Trophoii 3 X tentacle length), wide, elongate, flattened to semitriangular in cross-section, with centrallv situated sperm duct (figure 85, pvd), dor- sal iilood sinus (figure 85, ds). Penis tapers ahruptly to form small, cylindrical papilla (figures 46, 83). Pallial oyi- duct broad posteriorly, tapered anteriorly. Albumen gland (figure 90, ag) situated at rear of mantle cavity, joins posterior of capsule gland (figure 90, eg). Left and right lobes of capsule gland similar to T. ^eversianus (Harasewych, 1984: fig. 22; Kool, 1993b), Nucella lapil- lits (Fretter and Graham, 1994: fig. 171), but ventral and dorsal glandular areas proportionally much smaller (fig- ure 92). Bursa copulatri.x (figures 91, 93, be) joins an- terior of capsule gland, leads to vaginal opening (figures 90, 91, vg), situated below, posterior to anal opening, papilla (figure 90). Egg capsules of Xi/mcnopsis Duihciftinnis were dis- cussed but not illustrated by D'Asaro (1991:54). Cap- sules are attached to the dorsal surface of empty shells of X. miiricifonnis, densely grouped, overlapping each other. Egg capsule (figure 42) bulliform, approximately 6-7 mm in diameter, 2 mm high, outline subcircular, may vary according to substrate. Preformed exit aperture oval, situated at center of dorsal surface in shallow, sub- ovate depression, sealed by a mucus plug until hatching. Suture between capsule, plug weakly defined, nmning parallel to large axis of depression. Pattern of non-con- centrically oriented fibers visible on upper surface of capsules. External morphology of egg capsules veiT sim- ilar to that observed in dry specimens of Xi/mcnc plc- bciits (figure 43), which differ in having aperture situated apically rather than in depression. Type material: iHiirriniiin iiiuricifoniw] Lectotvpe (figures 14-16), BMNII 1837.12.15.2695, 6 paralecto- t>pes, BMNII 1992056, all from Strait of Magellan: [Fii- sii.s decolor] Ilokitype (figures 9-10), MNHNS unnum- bered. Strait of Magellan; [Buccinum caiiccllarioklcs] Lectot\pe (figures 17-19), BMNII 1992055-1, here des- ignated, I paralectotype BMNH 1992055-2, lack locality data; [Fusus liratus] Holotype (figures 1-3), USNM 5680, Orange Harbor, Tierra del Fuego: [Trophon loeh- bcckci] II()Iot\pe (figures 4—6), Lobbecke-Museum, no catalog ninnber, lacks locafity data; [Tropliou Ichnini] Lectotype (figure 13), MNHN unnumbered, here des- ignated, 2 paralectotypes, all from Santa Cruz, Argenti- na; [Trophon violaceiis] Lectotype (figures 7-8), MNHN unnumbereil, figured .syntyjae here designated as lecto- type, from Bale Orange, Tierra del Fuego; [Trophon paessleri[ Lectotype (figures 26-28), ZMH unnumbered, here designated, 4 paralectotypes, all from Port Grap- pler, Smvih Channel, Strait of Magellan; [Troi)hon paes- sleri var. turiita] Original tspe material, from Cape Horn, Tierra del Fuego, was destroyed (Hausdorf hi Utt.). The lectotvpe of Bucchmm nuiiiciformc (BMNH 1837.12.15.2695) is here designated as neotype of Tro- plion pac.'islcri var. titrrita.; [Trophon elon^iatus] Lecto- tvpe (figures 35-37), ZMH unnumbered, here designat- ed, from Puerto Angosto, Strait of Magellan, 4 paralec- totypes, 1 from Navarino Is., Chile, 1 from Puerto Bueno, 1 from Eden Harbor, 1 from Puerto Angosto, Strait of Magellan: [Trophon pseudoclongatns] Holotype (figures 29-31), ZMH unnumbered, Ushuaia; [Trophon alhus] Holotvpe (figures 38—40), ZMH unnumbered, Us- hutiia; [Troplion acuinimitiis] Holot\pe (figures 20-22), ZMH unnumbered, Punta Arenas, Strait of Magellan; [Trophon obcsus] Lectotype (figures 32-34), ZMH un- ninubered, here designated, from, Punta Arenas, Chile, 1 p;u;ilectot\pe from Punta Arenas, Chile; [Trophon fe- nc.stratu.s] Original t\pe material, from GregoiT Bank, Strait of Magellan, was destroyed (Hausdorf //i htt.). The lectotype of Buccinum muriciforme (BMNH 1837.12.15.2695) is here designated as neot\pe of Tro- phon fcncstratus; [Trophon confhouyi] Lectotvpe (fig- ures 23-25), ZMH unnumbered, here designated, 4 paralectotypes, all from Port Grappler, Smvth Channel; Figures 83-95. Xymcnopsi.s iiiioicifoniiis (King and Broderip, 18.32). 83. Anterior piirtion of mule specimen, mantle reHected. 84. Ctenidium and osphradium. 85. Transverse section of the penis, (figure 83 at 1-1'). 86. Male pallial gonoduct. 87. Transverse section (figure 86 at 2-2'). 88. Transverse section (figure 86 at .3-3'). 89. Stomach, opened dorsally. 90. Female pallial gonoduct. 91. Transverse section through capsule gland (figure 90 at 4—4'). 92. Transverse section (figure 90 at 6-6'). 93. Transverse section through bursa eopulatrix (figure 84 at 5-.5'). 94. Diagrammatic representation of the alimentary system. 95. Sagittal section throua;h the anterior portion of the foot. a = anus; aa = anterior aorta; abo = accessory boring organ; ag = albumen gland; ap = anal papilla; asg = accessory salivarv^ gland; be = bursa eopulatrix; ca = compacting area; eg = capsule gland; ct = ctenidium; dd = ducts to digestive diverticula; dgl = ampulla of gland of Leiblein; dp = duct pouch; ds = dorsal blood sinus; e = esophagus; gl = gland of Leiblein; hf = horizontal fold; hg = hypobranchial gland; i = intestine; ig = intestinal groove; 1 = lobes along meuitle edge; leg = left lobe of capsule gland; mo = mouth; nr = nerve ring; os = osphradium; p = penis; pg = prostate glanil; pma = posterior niLxing area; pro = propodial groove; pvd = penial vas deferens; r = rectum; ra = radula; rcg = right lobe of capsule gland; rg = rectal gland; s = siphon; sg = salivary gland; sto = stomach; tl and t2 = tyjihlosoles; vd = vas deferens; vg = vagina; vl = valve of Leiblein. Page 50 THE NAUTILUS, Vol. 114, No. 2 Figures 96-103. Xijincnopsis subnodosus (Gray, 1839) 96-97. Holotype of Fusns cancelVmus PhiLippi, 1845, and 11601)^6 of Buccinum suhnodosn Gray, 18.39. MNHNS unnumbered. Strait of' Magellan. 98-100. BMNH 19990.3.30, 4.3°.3S.9'17"S. 7.3°37.2'.38"\\'. 101. Operculum, external (left) and internal (right) Ndews, same specimen of 98-100. Scale bar = 1 cm. 102. Dorsal view of radular ribbon from specimen in figures 98-100. Scale bar = .50 fj.m. 103. Oblique (45°) lateral view, of radular ribbon. Scale bar = 25 |j,m. [Trophon elegans] Lectotype (figures 11-12), ZMH un- numbered, here designated, from Port Stanley, Mahinas (Falkland) Is., 2 paralectotypes from Port Stanley, Mal- vinas (Falkland) Is. Additional material examined: AMNH 50211, Strait of Magellan; AMNH 8.5428, Canadon de las Vacas; AMNH 99610, West Malvinas (Falkland) Is.; AMNH 137057, Beagle Channel; AMNH 17929.3, Puerto De- seado; AMNH 271757, 53°37.7'S, 69°54.6'W, in 42 ni; AMNH 271758, 54°10.2'S, 65°57.5'W, in 101 m; MACN 10043-1, Ushuaia; MACN 12374, Punta Arenas; MACN 12376, 53°10'S, 70°55'W; MACN 12448, Puerto Harris. Dawson Is.; MACN 12552, Ri'o Grande; MACN 12555, Rio del Fuego, Tierra del Fuego; MACN 13120, Punta Arenas; MACN 13568, Ushuaia^i MACN 1.3569, Ushuaia; MACN 17754, Puerto Deseado; MACN 22567, Punta Colnet, Isla de los Estados; MACN 22717, 54°48'S, 64°55'W, in 103 ni; MACN 23943, 20°07'S, 66°33'W, in 82 m; MACN 30876, Lapataia, Tierra del Fuego; USNM 96228, Strait of Magellan; USNM 96842, Puerto Mayiie, G. Pastorino and M. G. Harasewvch, 2000 Page 51 104 105 Figure 104. Original illustration of Fusiis biiccineiis La- marck, 1816 (pi. 427, figs. 3a, b). Figure 105. Sowerby in Gray's (1839, pi. 36. fig. 12) illus- tration of Fusus buccineus. The specimen depicted is Xipnen- opsis mtiricifonnis. Chile; USNM 126899, Tierra del Fuego, Intertidal; USNM 348720, Puerto Harberton, Tierra del Fuego; USNM .368273, Port William.s, Malviiias (Falkland) Is., in 18 m; USNM 368401, Port Stanley, MaKdnas (Falk- land) Is.; USNM 368639, Punta Arenas, Chile; USNM 368641, Punta Arenas, Chile, intertidal; USNM 368654, Punta Arenas, Chile; USNM 368761, Punta Arenas, In- tertidal, Chile; USNM 381693. Bahia San Sebastian, Tie- rra del Fuego; USNM 710029, Puerto Deseado, Santa Cruz, Intertidal; USNM 870002, 53°50'54"S, 70°30'42"W, in 15-17 m; USNM 870165. 54°05'S. 58°52"W, in 119 m; USNM 886188, 53°51'32"S. 70°25'52"W, in 2-3 ni; USNM 886190, 53°51'32"S. 70°25'52"W, in 2-3 ni; USNM 886737, 53°39'S, 70°55'30"W, in 20 ni; USNM 886745, 53°39'S, 70°55'30"\V, in 15-18 ni; USNM 901628, 53°39'S, Figure 106. Localities at which Xi/menopsis species (0 A', muricifonnis; O = X. cornigatiis: # = A', buccineus; -k X. subnodosus ) were collected off Chile and Argentina. 70°55'30"W, in 20 m; USNM 901629, Strait of Magellan, 53°39.4'S, 70°55.5'W, in 24 m; USNM 901630, 53°39.4'S, 70°55.0'W, in 82 m; USNM 901631, 54°47.25'S, 64°18.3'W. in 35 m; USNM 901632. 53°33.9'S, 69°58.8'W, in 82-91 m; USNM 901633, 53°39.3'S, 70°55.4'W, 38-42 m; USNM 901634, 53°17'S, 6S°13'W, in 0-1 m; USNM 901635, 53°39.8'S, 70°54.9'W, in 27-73 m; LACM 69170, Islas Guarello, Chile, 52°40'S, 73°41'W; LACM 10503. MoKiiieux Sound. Strait of Magellan. Chile; LACM 118051, Lively Island, East Malvinas (Falkland) Is., 52°00'S, 58°2S'W; LACM 71-262, KN Hkko Cruise 712. Sta. 672. W of Puerto Aiio Nuevo. Isla de los Estados. Tierra del Fue- go, Argentina, 54°45.1'S, 64°07.3'W, in 50 m; LACM 71- 305, IW Hkro Cruise 715, Sta. 690, 8 miles E Ensena- da Patagones, SE Tierra del Fuego, Argentina. 54°52'S, 65°05'W, in 144 m; LACM 71-337, R>V Hkho Cruise 715, Sta. 888, W of Bahia York, Isla de los Estados, Tie- rra del Fuego, Argentina, 54°49.5'S, 64°19.6'W, in 56- 63 m; LACM 73-66, IW Hkro, Cabo Colnett, Isla de los Estados, Tierra del Fuego, Argentina, 54°43'27"S, 64°14'18"W, in 15 ni; LACM 73-68, IW Hkho. Beagle Canal. C;ileta Aw;iiakirrh, Chile, 55°0'S, 69°02.2'W, in 17 m; LACM 73-69, IW Hero, Punta Valparaiso. Canal Cockburn, Chile, 54°22.2'S, 7r21.7'W. in 15 ni; LACM 73-70, Isla Carlos III, Strait of Magellan. Chile, 53°39.4'S. 72°14.8"W. in 11-12 ni; LACM 73-71. Punta Dashwood. Canal Smyth, southern Chile, 52°24'S, 73°39.7'W, in 12 ni; LACM 73-72, Bahia Tom, Chile, Page 52 THE NAUTILUS, Vol. 114, No. 2 5()°11.3'S, 74°47.9'W, in 14 m; LACM 73-107, RA^ Hero, Colling%vood, Chile, 5r52'S, 73°43.6'W, intertid- al to shallow subtidal; LACM 75-49, Puerto el Hambre, Brunswick Peninsula, Strait of Magellan, ChLle,53°37'S, 70°56'W, intertidal; LACM 78-84, IW Hero Sta, B 4- 6, N of Cabo Buen Tiempo, Rio Gallegos, Santa Cruz Prov., Argentina, 51°16-20'S, 68°50-54"W, in 30 m, mud bottom: LACM 86-268.5, Canal Oeste, S side Isla madre de Dios, Ultima Esperanza, Magallanes Prov. Chile, 50°27.7'S, 75°11.5'W, in 30 m. Literature records: RA' Discoxerv: Sta. 52. 7.4 ca- bles N, 17°E of Naw Point, Port William, East Mahinas (Falkland) Islands, in 17 ni; Sta. 1230, 6.7 miles N, 62°W of Dungenes Light, Strait of Magellan, in 27 m; Sta. WS784, N of Malvdnas (Falklandy Islands, 49°47.75'S, 61°05'W, in 170 m (Powell, 1951); Sta. 24,25, Puerto Eden in 10-12, 6-7 m respectively; 30,33 Puerto Eden, in 8-10 and 11-12 m; Sta. Caleta Lackawana in 4-6 m; Sta. 43 Paso Indio; Sta. 50, 63, 65 Puerto Williams, Nav- arino Is. (Dell, 1971). Distribution (figure 106): Knov\ni from 45° S (Chonos Archipelago), Southern Chile {fide Dell, 1971 as X. de- color); Tierra del Fuego, Strait of Magellan, Beagle Channel (Argentina) and the Malvinas (Falkland) Is- lands. The bathvmetric range for X. imihciformis is 0- 170 m. Remarks: Xiimcuop.sis miuiciformis has a long and comphcated taxonomic histoiy. The oldest name to have been applied to the Xymenopsis characterized by spiral sculpture that appears as incised grooves is Buccinum luuriciformc King and Broderip, 1832. There were 2 lots of svntvpes, BMNH 1837.12.15.2695 (2 specimens) and BMNH 1992056 (5 specimens), both from Str;iit of Ma- gellan. One specimen BMNH 1837.12.15.2695 (figures 14-16), was illustrated by Dell (1972:8, figs. 31,32) as the holotvpe (an inads'ertent lectotype designation). The holotvpe of Fh.si/.s decolor Philippi, 1845 (figures 9-10), iilso from the Strait of Magellan, clearly represents a worn specimen of X mttiicifonnis. The next name to be apphed to this species is Buccinum cancellarioidcs Reeve, 1847, based on 2 syntyjjes lacking locafity data. One specimen is here designated as lectotype (figures 17-19). Powell (1951) recognized B. cancellarioidcs to be an earher name for Ftisus lirafus Gould, 1849 (Ho- lot\-pe, figures 1-3). Because he was unsure if Reeve's (February, 1847) name was a junior homonvan of Buc- cinum cancellaroides Basterot in Grateloup, given as 1847 by Sherborn (1924:1030). he ret;iined the use of X. liratus. The holotvpe of Trophon loebbeckei Kobelt, 1878 (figures 4-6), from an unkniown locality, also matches closely the holotype of B. muricifimnc, as do the lectotypes of Trophon lebnini and Trophon viola- ceus, both Mabille and Rochelinme in Rochebrune and MabiUe, 1889. Trophon Ichruni was based on 3 .svntypes, MNHN unnumbered, all from Santa Cruz, Argentina. One specimen is here designated as the lectoty|3e (figure 13). The figured s)'ntype of Trophon violaceus MNHN unnumbered, from Baie Orange, Tierra del Fuego is here designated lectotvpe (figures 7-8). In his publication on the molluscan fauna of the Ma- gellanic Province, Strebel (1904) proposed numerous taxa based on minor phenotypic variants. Among these were 16 species and varieties of Trophon that were sub- sequentlv referred to Xipncnopsis bv Powell (1951). Ac- cording to Dance (1986:227), Strebel's collections were housed in the Hamburg Museum and "totally destroyed " during World War II. However, this appears to be true only for the "dry" (shell only) specimens. The alcohol preserved material on which much of Strebel's (1904) Magellanic publication was based, survived the war and is in the collections of the Zoologisches Institut und Zoologisches Museum der Universitat Hamburg. Mate- rial collected b\- the Swedish Sudpolar-Expedition, in- cluding types of taxa described by Strebel (1908), is in the Swedish Museum of Natural History in Stockholm. The senior author visited both these museums in the course of this study, and was able to locate much of Strebel's tspe material. Examination of the t\pes ot; Tro- phon paessleri, T clongatus, T. pscudoelongatus, T al- bus, T acuminahis, T couthouiji, T elegans and T. obesus leaves no doubt that these are, at most, minor moq:)ho- logical variants oi Xipnenopsis nuiriciformis. The taxon Trophon paessleri is based on 6 syiitypes. ZMH unnum- bered, 5 from Port Grappler, Smyth Channel, Strait of Magellan and 1 from Punta Arenas. One specimen (fig- ures 26-28) from Port Grappler is here designated as lectotype. The type material of Trophon clongatus con- sists of 7 syntypes, ZMH unnumbered, 2 from Puerto Angosto, Strait of Magellan, 3 from Puerto Bueno, 1 from Navarino Is. and I from Eden Harbor One spec- imen (figures 35-37) from Puerto Angosto is here des- ignated as lectotype. Trophon couthouiji is based on 3 syntypes, ZMH unnumbered, from Port Grappler, Smvth Channel. One specimen (figures 23-25) is here designated as lectotvpe. Holotvpes are illustrated for Trophon pscudoelongatus (figures 29-31), ZMH unnum- bered, and T. albus (figures 38—40), ZMH unnumbered, both from Ushuaia; and T. acuminatus (figures 20-22), ZMH unnumbered, from Punta Arenas, Strait of Ma- gellan. Trophon elegans is based on 3 specimens from Port Stanley, MiJvinas (Falkland) Is. One specimen (fig- ures 11-12) ZMH unnumbered, is here designated as lectotype. Trophon obesus is represented by 2 syntypes from Punta Arenas, Chile; one (figures 32-34), ZMH unnumbered, is here designated as lectotype. The type material of Trophon paessleri var. turrita and of T fenestratus could not be found, and was among the material destroyed during World War II (Hausdorf in litt.). Based on Strebel's (1904) descriptions and figures, there is no doubt that both these taxa are conspecific with Xymenopsis muricifonnis. We therefore designate the lectotype of Buccinum muriciforme (BMNH 1837.12.15.2695) as the neotype for both Trophon paes- sleri var turrita and T fenestratus, making both these taxa objective junior synomins of Buccinum muricifiir- G. Pastorino and M. G. Harasewych, 2000 Page 53 Xijinenopsis comi^atiis (Reeve, 1848) (Figures 57-71, 80-82, 106) Fusus cornif^atiis Kee\e, 1848: pi. 20 figs. 84 a. h. Kolielt, 1878: 293, pi. 74. fii lift. ). This taxon is considered to be a svnonvm of X. buccincus based on Strebel's (1904) descriptions and figures. We therefore designate the neotype of Fusus buccincus (USNM 870410) to serve as the neotype of Trophon ringei. Trophon ringei thus becomes an objective synonym of Fusus buccincus. Xyincnopsis subnodosus (Gray, 1839) (Figures 96-103, 106) B[uccimnn] suhtiodosa Gray, 1S39: 118. Fu.ms cancellinus Philippi, 1845: 67; 1846: 117, pi. 3, fit;. 2; Hupe in Gay, 1854: 163. Trophon cnncellinii.i Philippi — Kobelt, 1878: 291, pi. 74, figs. 7, 8; Rochehnme and Mabille, 1889: H.59 Diagnosis: Axial sculpture of strong, irregular ribs (12-16 on body whorl), nearly as broad and intenening spaces. Spiral sculpture of dense, evenly rounded cords, (6 on penultimate whorl, 22-24 on body whorl), broader than intervening spaces. Spiral threads present between adjacent cords. Outer lip with apertural teeth. Description: Shell large (to 37 mm), solid, slender, fusiform, dirty white, chalkT. Protoconch miknown. Te- leoconch of up to 6 gently convex whorls. Spire high, = Vi shell length. Spire angle == 45°. Suture strongly impressed. Aperture small, ovate; outer lip thickened, with 8-10 weak teeth; inner fip concave, thickly glazed; interior whitish. Siphonal canal long, obfique, open. PseudoumbiUcal chink, small but present. Axial sculp- ture of strong, irregular ribs (12-16 on body whorl), nearly as broad and intenening spaces. Spiral sculpture of dense, evenK' roundeil cords, (6 on penultimate whorl, 22-24 on body whorl), broader than intei-vening spaces. General morphology- of animal and operculum Page 56 THE NAUTILUS, Vol. 114, No. 2 (fi2;ure 101) as in X. miiriciformi.s. Osphradium < V2 cte- nidium length, asymmetrical, with 48 and 32 leaflets on the external and internal side respectively. Ctenidium 2- .3 times as wide as osph/adium, witli 85-90 triangular leaflets. Radular ribbon small, short (0.23X aperture length). Radula (figures 102-103) similar to those of X. corntgattis and X. huccincus, but with single weak den- ticle along outer edge of each lateral cusp. Type material: [Buccinum stibnodosa] Original type material from the Pacific Ocean lost (K. Way, in lift.). The holotype of Fusits cancellintis, MNHNS, is here designated as neot\pe of Bticciniim subiuxlo.m. [Fiisiis cancellintis] Holot)'pe, MNHNS unnumbered. Strait of Magellan. Additional material examined: BMNH 19990330, 43°38.9'17"S, 73°37.2'38"W, East side Traiguen Is., Es- tero Elefantes, Region 11, Chile. Dredged in 5-15 m on bottom of colibles and dead \enerids; sheltered bay. Distribution: Known from the Str;ut of Magellan, and Traiguen Island, Chile. Remarks: Gray (1839) introduced Buccinum suhno- closa as a variety of Buccinum muriciformc (=Xi/mcn- opsis muiiciformis King and Broderip) in a two Une de- scription without illustration. As Gray's type material is lost (K. Wav, in Jitt.) this leaves some doubt as to the identitv' of this species. Grav's description refers to "al- ternate narrow low and wider convex spiral ridges" that produce tubercles as they rise over the axial ribs, fea- tures characteristic of both X. corrugatus and a species subsequently named Fu.su.s canccUinus. Grav does not mention the prominent apertural teeth that are diagnos- tic of X. canccUinus, but it is possible that he had a sub- adult specimen lacking apertural teeth. Gray states that B. subnodoso inhabits the Pacific Ocean. Fusus cancel- linus occurs along the Pacific Coast of Chile, while X. corrugatus is known only from the Malvinas (Falkland) Islands and Isla de los Estados. In order to stabihze the nomenclature of Recent Xi/mcnopsis. we designate the holot^pe oi Fusus canccUinus Phihppi, 1845 (MNHNS) to serve as neotvpe of Buccinum subuodosa Grav, 1839. Fusus canccUinus thus becomes a junior objective syn- onym of Buccinum subnodosa. We know of only two specimens of Xi/mcnopsis sub- nodosus. incluchng the neotspe (shell onK ) housed at the Museum of Santiago, Chile. Xtjmenopsis subnodosus re- sembles both X. muiiciformis and X. corntgatus in size, general shape, and in the coarseness of its axial ribs. It differs from X. muricifonnis in ha\ing a thicker shell, narrower more numerous, and more rounded spiral cords, and shares with X. corrugatus the presence of fine spiral threads interspersed between adjacent cords. Xy- menopsis subnodosus is larger has a higher spire, and coarser sculpture than X. buccincus^ The most distinctive features of X. subnodosus are the thickening oi the outer hp, and the presence of weU defined apertural teeth, which occur in New Zealand members of the Xi/mcnc complex, but not in any other Xymcnopsis. Xymcnopsis subnodosus is most similar to X. corntgatus. which is restricted to the Mahinas (Falkland) Islands. We provi- sionally retain this taxon, recognizing that it may prove to be a subspecies or rare variant of X. corrugatus char- acterized by an atavistic expression of a rare allele or alleles for apertural dentition. SYSTEMATIC KEY TO THE GENUS XYMENOPSIS 1. Shell large, with strong, well defined axial ribs . . 2 Shell small, with weak, poorly defined axial ribs 3 2. Shell thin, with spiral sculpture of flat cords much broader than intervening spaces, which appear as incised furrows X. muiicifoiinis Shell thick, spiral sculptine ot dense, evenly rounded cords, (6 on penultimate whorl, 22- 24 on body whorl), broader than intervening spaces. Outer lip thickened, with 8-10 weak teeth X. subnodosus 3. Spiral sculpture of rounded cords slightly broader to sUghtlv narrower than intervening spaces, producing weakly canceUate tubercles at intersection with axial ribs. Weak spiral threads present between adjacent spiral cords X. corrugatus Spiral sculpture of flattish cords as broad as interx'ening spaces, forming weakly cancellate sculpture at intersections with axial ribs X. buccinciis ACKNOWLEDGMENTS We thank the following people for access to material in their collections: K. Wav (BMNH); A. Waren (NHRM); P Bouchet and V. Heros (MNHN): J. H. McLean (LAGM): P Mikkelsen (AMNH); B. Hausdorf (ZMH); O. Galvez Herrera (MNHNS); J. Boscheinem (Loeb- becke-Museum); H. Irigoyen and A. Tablado (MAGN); C. Ituarte (MLP). R. Prasad (Auckland Museum) pro- vided specimens oi Xt/mcnc plcbcius. P. Louzet took the photographs of specimens at MNHN. D. Reid (BMNH) provided the presened specimen of X. subnodosus. Yu. Kantor provided useful discussions during the course of this work. V. Ivanov prepared the anatomical drawings of X. muricifonnis. R. Petit provided information on nineteenth century hterature. E. H. Yokes provided helpful discussions of muricid systematics. We are grate- ful to A. Beu and J. H. McLean for thorough reviews and helpful suggestions that improved this paper This study was conducted during a Postdoctoral Fel- lowship granted to the senior author by the Consejo Na- cional de Investigaciones Cientificas v Tecnicas (CON- ICET), Argentina, to work at the National Museum of Natural History, Smithsonian Institution, Washington. DC. It was supported in part by a Research Award from the NSF-USAP United States Antarctic Program [Con- tract No. OPP-9509761] and a grant in aid from the Conchologists of America and the Walter E. Sage Me- moriid Award. G. Pastorino and M. G. Harasewvch, 2000 Page 5" LITERATURE CITED Ben, A. G. and P. A. Mawvell. 1990. Cenozoic Molluscaof New Zealand. New Zealand Geoloi;ical Suncx Palcontological Bulletin 58:1-518. Bnniet, R. 1997. 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Sinione Museu lie Zoologia da Uiii\ersidade de Sao Paulo Cai.xa Postal 42694 04299-970 Siio Paulo BRAZIL lrsimoiie@usp.br ABSTRACT A new bathval species of the family Pseudococculinidae is de- scribed from samples dredged in 1987 along the continental slope of southeastern-eastern Brazil b> the French research vessel Marion-Diifresne. The new species, allocated in the ge- nus Copulahijssia. differs from the 4 other known congeners by characters of shell, external morphology, mantle cavity, and radula. Descriptions of internal moqihology based on micro- dissections are provided, including the digestive system and the complex muscular apparatus ot the odontopliore. The distri- bution of characters ot taxonomic significance at the species level in Copulabyssia is given in tabular format and discussed. Key words: MoUusca, Cocculiniformia, Vetigastropoda, Lepe- telioidea, bathval, anatomy, taxonomy, odontopliore. INTRODUCTION Bathyal samples dredged off E-SE Brazil yielded spec- imens of an unnamed species of the genus Copulabi/ssia, family Pseudococculinidae. Two live specimens and one empty shell were collected at three stations performed between 960-L320 m. The Pseudococcuhnidae and oth- er families comprising the superlamily Lepetelloidea Dall, 1882, are included amongst the mollusks living at greatest depths in the ocean (Leal and Harasewych, 1999; Lesicki, 1998; Marshall, 1986; Wolff, 1979). The new taxon exhibits the diagnostic characters of the genus Copulabyssia Haszpmnar, 1988. Copulabyssia riosi new species is herein compared with the 4 other species of the genus named to date (Lesicki, 1998): C. corni^ata (Jeffreys. 1883) (NE Atlantic); C. ii,mda1a (Marshall, 1986) (SW Pacific); C. Icptah-a (Verrill, 1884) (N Atlan- tic); C. similaris Hasegawa, 1997 (Japan). MATERIALS AND METHODS Two live-collected specimens and one empty shell were available for studv. A single specimen was dissected. Mi- crodissection was performed with the specimen im- mersed in 70% ethanol. The apical (distal) region of vis- ceral mass was damaged; adjacent structures were de- scribed and are shown herein based on reconstruction. After dehydration in a standard alcoholic series, the sec- ond specimen was chemically dried tor 10 min in hex- amethykUsilizane (HMDS). Shells, radula, and chemi- cally dried soft parts were coated with carbon and gold and e.xamined and photographed under Hitachi S-57() (at National Museum of Natural History, Smithsonian Institution), Leo 440 (at MZSP), or Philips XL30 ESEM-FEG (Center for Advanced Microscopy, Univer- sity of Miami) electron microscopes. Institutional abbre- viations used are; MNHN, Museum national dTIistoire naturelle, Paris, France; MZSP, Museu de Zoologia da Universidade de Siio Paulo, Brazil; NMNH, Museum of New Zealand Te Papa Tongarewa, Wellington; NSMT, National Science Museum, Tokyo, Japan; SMNH, Natur Historika Riksmuseet, Stockliolm, Sweden; USNM, Na- tional Museum of Natural History, Smithsonian Institu- tion, Washington. Other alilirexiations and text conven- tions are: 1, 5, lateral teeth (lateral 5 = pluricuspid tooth); 1, 2, marginal teeth; ac, anterior cartilages of odontophore; an, anus; bm, buccal mass; br, subradular membrane; bs, Iniccal sphincter; df, dorsiil folds oi buc- cal mass; dg, digestive gland; ep, epipodium; es, esoph- agus; et, epipodial tentacles; gi, gill; go, gonad; gp, gen- ital pore; if, inner mantle fold; in, intestine; ir, insertion of m4 in radular sac; is, insertion ot m5 in radular sac; ki, kidney; It, left cephalic tentacle; mb, mantle edge; mj, jaw and peribuccal muscles; mo, mouth; mp, me- sopochum; m2, pair of buccal mass retractors; m3, inner buccal mass protractors; m4, pair of antero-dorsal ten- sors; mS, pair of postero-dorsal tensors; ni6, horizontal muscle; ni7, pair of odontophore approximators; mS, pair ot cartilage approximators; no, nuchal ca\its: of, outer mantle fold; pa, mantle papillae; pc, posterior car- tilages of odontophore; pr, protuberance on right ten- Page 60 THE NAUTILUS, Vol. 114, No. 2 tacle; re, rectum; ra, radula; m, radiilar caecum; rs, radular sac; rt, right cephalic tentacle; sc, subradular cartilage; sf, fold along gill; sg, seminal groove; sm, shell muscle; sn, snout; st, stomach; vg, vestigial left gill leaf- let. The odontophore muscles examined in this studv were also described bv Haszpnuiar (1988: 165-167, fig. .3) for Kluilahijssia vcnczuclcnsis McLean, 1988. Ter- minology used herein (boldface) corresponds to the one used bv HaszpiTuiar as follows: mj = br -I- otr; bs = bs; in2 = dr2; in4 = rd2; m5 = rr; m6 = ho; m7 = sr3 (?); mS = ci. The remaining small muscles described by Haszpnmar (1988) could not be observed in the single specimen a\';ulable for microdissection. SYSTEMATICS Class Gastropoda Cuvier, 1797 Subclass Orthogastropoda Ponder and Lindberg, 1996 Order Vetigastropoda Salvini-Plawen, 1980 Remarks: The superfamilies Cocculinoidea Dall, 1S82, and Lepetelloidea Dall, 1882 have been united under the order Coccuhformia Haszprunar, 1987. Re- cent studies of gastropod phylogeny based on moi"pho- logical characters (Ponder and Lindberg, 1996; 1997) suggest that CoccuMnoidea is sister taxon to Neritopsina, and that Lepetelloidea is part of the order Vetigastro- poda (see also Haszpioinar, 1998: 664). This arrangement is followed herein. Harasewvch ct al. (1997) indicated, based on partial 18S rDNA sequences, that CoccuHni- formia may not constitute a clade. These latter authors, however, suggested that Cocculinoidea and Lepetello- idea are more closely related to each other and to Pa- teUogastropoda than to Neritopsina or Vetigastropoda. Superfamily Lepetelloidea Dall, 1882 Family Pseudococcuhnidae Hickman, 1983 Subfamily Caymanabyssinae Marshall, 1986 Genus Copiilabijssia Haszprunar, 1988 T\pe species: CoccuUna oirn/gflffl Jeffreys, 1883, p. 394, pi. 44, figs. 2-2a. Type locaUty-; 59°40'N, 7°21'W, 930 m, Triton Ex]:)ecUtion station 10. Diagnosis: Protoconch sculpture of microscopic pris- matic crvstals; right cephalic tentacle exceptionallv large, "swollen"; right cephalic tentacle with deep, open glan- dular seminal groove situated postero-dorsallv; large pedal gland; gills palhal, several well-developed gill leaf- lets on right side, a single one on left side. Copidabi/ssia riosi new species (Figures 1-24, Table 1) Diagnosis: Teleoconch at first with radial sculpture then with closely set concentric ribs (interspaces about 25 (Jim in width); internally with narrow but well-defined anterior apical septum; radial sculpture only on early te- leoconch; cutting area of rachidian tooth with blunt cusp; left side of subpalhal cavity with small structure that could represent a vestigial gill leaflet; right cephalic tentacle with prominent subterminal protuberance; left cephahc tentacle small for genus; inner branches of epi- podial tentacles filamentous. Description: Shell (Figures 1-6): Small (length to 3.50 nun), thin, arched, elevated (height/length about 0.5), with apex at posterior quarter pointing in posterior direction. Anterior slope convex, about 88% of shell length. Posterior slope concave, about 30% of shell length. Shell surface usually lightlv eroded, milky white. Protoconch (figure 4) length about 200 |jLm, surface bad- ly eroded in type material. Teleoconch sculpture of con- centric (commarginal) ribs separated by intervals of about 25 |jLm. Concentric ribs narrow (about 10 jim) and crossed bv fine radial lines on earlv part of teleoconch (figure 5). Concentric ribs on latter part of teleoconch proportionally thicker, radial lines absent (figure 6). In- ternally with narrow, but well-defined, anterior apical septum. Aperture elhptical (figure 3), not conforming to a planar surface, convex. Shell muscle scar continuous, horseshoe-shaped. Head-foot (Figures 7, 8, 13, 14, 17): Head prominent and large (about % foot length) (figures 7, 13, 14, 17). Cephahc tentacles (figures 7, 13, 14, 17, rt. It) on dorso- lateral surface of head. Right tentacle about 4 times larg- er than left tentacle, modified as copulatory organ (de- scribed below). Eyes apparently lacking (unpigmented?). Snout (figures 7, 13, 14, 16, 17, sn) very large, cyhndrical on base, gradualK' becoming slightK' flattened dorso-ven- trally. Anterior extremity of snout almost flat, broader than region immediately behind it, with pair of small lateral projections. Foot large and flat (figures 7, 13, 14), smaller than shell aperture in presened animal. Meso- pochum (figures 7, 13, 14, mp) flattened. Epipodium (figures 7, 13, 14, ep) flange-Uke, surrounthng entire me- sopodium, larger on anterior edge, which covers baso- ventral region of snout. Pair of epipodial tentacles (figure 7, 9, 13, 14, et) well-developed, projecting from poste- rior region at both sides of longitudinal axis of animal, slightly dorsal to epipodium, on intersection between mantle and shell muscle. Each epipothal tentacle bifid; outer branch shorter and with broad tip; inner branch longer and pointed. Shell muscle (figures 15-17, sm) horseshoe-shaped (concaNdty towards anterior region); anterior extremities broader, pointing away from shell apex. Almost all head-foot muscles converge toward in- sertion of shell muscle. Mantle cavity (Figures 12-15, 17): Mande edge (fig- ures 12-14, 17, mb) thick, with 2 folds; outer fold (fig- ures 12, 14, 17, ofO covered by slender and very small, translucent papillae (figure 12); inner fold (figures 14, 17, if) about twice as broad as outer fold, richlv glan- dular, white. A small, transparent, yellowish protuber- ance (figure 13, vg) on inner margin of inner told, in region adjacent to left cephafic tentacle, may represent vestigial left gill leaflet. Mantle cavity surrounding entire head-foot, somewhat deep, i.e., with ample space be- J. H. Leal and L. R. L. Simone, 2000 Page 61 Figures 1-6. Coptilabt/ssia riosi new species: shell of holotrvpe, MZSP 32150. 1. Dorsal view. 2. Lateral \iew. 3. X'entral view. Scale line = 1.0 mm. 4. Protoconch. Scale line = 100 p-m. 5-6. Shell sculpture. 5. On early teleoconch. Scale hne = 50 jxm. 6. Close to shell margin. Scale line = 100 [xm. tween mantle edge and shell muscle. Nuchal cavitv (fig- ures 14, 15, 17, nc) deeper than remainder of mantle cavity. Gill (figures 13, 14, 17, gi) small, situated on right side of mantle cavity, somewhat parallel to mantle edge and extending for about half of foot lenijth, consisting of aliout 14 successively smaller leaflets. Anterior extremity oi gill (figure 5, gi) just posterior to head-foot Umit. Each leaflet low and triangular, relatively thick. Most posterior leaflets situated peqjendicularlv to mantle edge, gradu- ally becoming obfique to it on anteri(jr region. A small, narrow, and low fold (figure 14, sf) nms between gill and shell muscle along posterior half of gill. Anus (fig- ures 15-17, an) and genital pore (figure l7, gp) on right side, at head and foot hniit, just anterior to right extrem- ity of shell muscle. Circulatonj and excretory sijstem.'i (Figures 15-17): Heart not observed. Large blood vessels ;ilong thick- mantle border Kidney (figures 15, 17, la) verv small, white, sohd, slightly triangular; situated in right-posterior region ot head, just posterior to rectum. Visceral mass (Fifiures 15, 16): Round, surrounded by following stmctures: laterally and in posterior direction by shell muscle, in anterior direction by buccal mass, ventraliv by inner surface of mesopodium, and dorsallv by central region of mantle and shell. Gonad (figure 15, go) and digestive gland (figure 15, dg) cream in color, intersected by several intestinal loops (figure 16). Gonad situated mainly on left region and digestive gland mainlv on central region. Digestive si/sfem (Figures 7. 13-24): Mouth (figures 7, 13, 14, 18, mo) on middle-ventral region oi anterior sur- Page 62 THE NAUTILUS, Vol. 114, No. 2 Figures 7-12. Copulabi/ssit] riosi new species; paraUpe, MNHN unnumbered, gross external moipholoi^ and radula. 7. N'entral \ie\v ot animal. Scale line = .500 p,m. 8. Detail of right cephalic tentacle (copulatorN- organ). 9. Inner branches ot epipodial tentacles (damaged during preparation). Scale Unes = 100 |j.m. 10-11. Radula. 10. View of 5 rows. Scale hue = 50 (jim. 11. Detail of lateral and first marginal teeth. Scale line = 10 [j.m. 12. Papillae on outer mantle fold. Scale line = .50 |xm. face oi snout, oriented transversally. Buccal mass (figure 16, bm) somewhat large, occupying about entire head cavit\'. Buccal sphincter (figure 18, 19, bs) well devel- oped. Superficial protractor muscle of buccal mass rel- atively narrow, connected to lateral and dorsal regions of buccal sphincter and adjacent region of snout, iiinning along lateral and dorsal sinface of buccal mass, con- nected to lateral and ventral region of buccal mass. Dor- sal wall of buccal mass ver\' thin, with pair of thick lon- gitudinal folds (figure 19, df) that join together in an- terior direction at median fine, becoming gradiuillv weaker toward esophagus (figures 16, 19, es). Odonto- phore (figures 18-24) large, comprising most of buccal mass, oval in outline. Odontophore muscles (figures 18- 24): (ml) (not illustrated) several small and short fibers connecting buccal mass to adjacent inner lining of snout, more concentrated on dorsal surface; (m2) (figures 18, 19) pair of narrow retractor muscles of buccal mass con- nected to inner ventral lining of snout just posterior to buccal mass, running parallel to radular sac (figures 19- 23, rs), connected to postero-dorsal surface of buccal mass on lateral surface of beginning of esophagus; (m3) (figure 21, 22) pair of thin inner protractor muscles of buccid mass connected to inner lateral lining ot mouth, running on (and covering) lateral surface of odonto- phore, connected to antero-ventral margin of posterior cartilages; (m4) (figures 20-24) pair of antero-dorsal ten- sor muscles connected in part to outer ventral surface of anterior cartilages and in part to lateral surface of posterior cartilages, running along (and covering) pos- J. H. Leal and L. R. L. Simone, 2000 Page 63 Figures 13-17. Copiilabi/ssin nasi new species; paratope, MZSP 32149, gross inorphologv. 13. \'entral \ne\v. 14. Lateral \iew, right side, margin of mantle deflected to show its inner surface. 15. Dorsal \iew. 16. Digestixe system, dorso-lateral \iew, left side, shown by transparency, only dorsal contour and part of shell muscle shown. 17. Detail of head, lateral view, right side, adjacent mantle removed by means of an incision around shell muscle. Scale lines = 500 jjim. Page 64 THE NAUTILUS, Vol. 114, No. 2 mo J. H. Leal and L. R. L. Siiiioiu', 2()()() Page 65 Table 1. Diagnoses of species oi Copulnl}yssia. "Shell length" is miLximinn size for species, "Rih interspaces" is the distance between concentric ribs on posterior shell slope. "Lett gill leaflet" is size of single-leaflet gill in relation to right gill leaflets; medinni wonld be about the same size as right gill leaflets. "Right cephalic tentacle" = copulator>- organ ("RT"). Unknown character states indicali'd b\' "NA", Degree of de\elopment of cephalic tentacles is relatixe to other congeneric species. C. riosi C. Icptalea C. t^iriddta C. cornii^ata C. similah.s Di.stribution Off SE, E Brazil NW Atlantic Off New Zealaml NE Atlantic. Med japan Character Shell leiiijth (nun) 3.50 2.45 1.95 1.35 3.36 Rib interspaces (jxni) ±25 ±40 ±40 ±130 ±50 Radial sculpture early Icleo only strong lacking present very weak Postition of apex anterior 2/10 anterior .3/10 anterior .'VIO anterior 1/10 variable Apical septum present absent absent absent absent Rachidian broad broad, rounti sides broad elongate broad Racliidian cutting area blunt cusp long, pointed cusp cuspless, rounded cuspless, s(juarish cuspless, round Cusjis lateral tooth 5 2-3 4 NA 2 2 Epipodial tentacles filamentous NA NA stubby present Right ce ihalic tentacle well-developed NA swollen base swollen base "bilobed" Left cep lalic tentacle vei-y reduced NA normal normal normal RT protuberance well-deyeloped NA absent absent absent Left gill leaflet ?vestigial NA medium small Pabsent References this work Waren, 1991 Marshall. 19.S6 Haszpnuiar 1988 Hasegawa, 1997 McLean and Hara- Ilaszprunar, 1988 Dantart and sewych, 1995 Luque, 1994 tenor surface of odontophore (figure 24), connected to ventral surface of radular sac on region posterior to emergence of radula; (m5) (figures 20-24) pair of pos- tero-dorsal tensor muscles connected to postero-ventral surface of posterior cartilages, ninning dorsallv and me- dially along (and covering) posterior surface ot odonto- phore, connected to radular sac just posterior to m4 in- sertion; m6) (figures 20, 24) horizontal muscle relatively narrow, connected to antero-ventral margin (jf anterior cartilages along about % oi their length; m7) (figiu'es 20, 22, 24) pair of narrow ventral tensor muscles connected to meso-ventral margin of posterior region of anterior cartilages, running along meso-ventral surface of odon- tophore, connected to antero-ventral e.xtremih of sub- radular cartilage (figures 21-23, sc); m8) (figures 10, 12) pair of broad approximator muscles of cartilages con- nected to antero-lateral surface of posterior cartilages (figures 22, 24, pc), nmning toward anterior cartilages, connected along lateral surface of anterior cartilages (fig- ure 24, ac). Anterior cartilages ot odontophore flattened, long, narrower in anterior direction, broader in posterior direction. Posterior cartilages short (about 1/5 of anterior cartilage length), semi-spherical. Anterior and posterior cartilages joined together on small area close to median fine (figure 24). Esophagus (figures, 16, 19, es) narrow and simple, running toward postero-ventral region of vis- ceral mass, where it suddenly bends in anterior direction to its insertion in stomach (figure 16, st). Stomach large, cylindrical, directed toward shell apex; inner surface uni- form, iridescent, greenish. Single duct to digestiye gland situated close to shell apex. Stomach suddenly narrows giving origin to intestine (figures 16, in), which runs to- ward the right and then in posterior direction, surround- ing dorsal margin of shell muscle, gradually turning ven- trallv and in anterior tlirection to buccal mass, where it suddenly curves in posterior cbrection, running along dorsal surface of mesopodium dorsalK to its posterior edge, looping dorsally to the right, nmning ventrally back to preceding loop, near posterior region of head, where it crosses transversally from right to left dorsallv to left extremit)- f)f shell muscle, suddenly turning suc- cessively in posterior, dorsal, and anterior directions; last loop represented by rectum (figures 15-17, re) crossing transversally from left to right through kidney along pos- terior hmit of mantle ca\iU'. Anus (figure 15-17, an) small and simple, located on posterior right side of man- tle cax-ity. Radula (Fifiiirvs 10-11): A.s)anmetrical, narrow, rela- tively short (about tsvice odontophore length). Rachidian tooth rhomboid, broad for genus, broader at mid-sec- tion, and blunt (worn?) cusp. First lateral tooth trian- Figures 18-24. Coptilabi/ssin riosi new species: paratvpe, MZSP 32149, buccal mass and odontophore. 18-19. Buccal mass extracted from snout. 18. Neutral \iew, 19. Dorsal \ie\v. 20-24. Odontophore. 20. Neutral \iew, supei^ficial layer ot muscles and membranes partially excised. 21. Dorsal \iew 22. Neutral \iew, superficial la\er of nniscles anil membranes fully excised, 23. Dorsal view. 24. Dorsal view, radular ribbon and radula sac extracted, most of muscles and cartilages deflected to expose inner surfaces, right-side structures more deflected than those on left side. Scale hnes = 500 (jlui. Page 66 THE NAUTILUS, Vol. 114, No. 2 Figures 25-28. Dorsal and lateral views of shells of other species o{ Copulabijssia. 25. C. leptalea (Verrill, 1884). USNM 757345. NE of Norfolk, \'irginia, 3080-,3090 m, RA' Gillis cniise 7508, station 36. 26. C. cornioata (Jeffrevs, 1883). SMNH unnumbered, SE of Reykjanes Ridge, off southern Iceland, 2.50-350 m, on sunken wood. 27. C. gracfnto'( Marshall, 1986). NMNZ M. 75007, holohpe, off \\'hite Island, New Zealand, 107.5-1100 m, on sunken wood. 28. C. similaris Hazegawa, 1997. NSMT-Mo 70822. off Toi, Suniga Bay, Japan, 4.30-710 m, on sunken wood. Scale lines = 1.0 mm. gular with outer basal projection that fits into a depres- sion on second lateral tooth, and pointed single cusp. First lateral tooth (figure 11, 1), broad, triangular. Sec- ond, third, and fourth lateral teeth strongly ciu^ed. with two short cusps. Fifth lateral (pluricuspid) tooth (figure 11, 5) massive, club-like, with 3 subterniinal denticle- hke cusps. Latero-marginal plate present. Inner marginal teeth pointed, with two denticle-like lateral cusps and thick shafts (e.g., figure 11, 1. 2), decreasing in size out- ward. Second marginal tooth largest. Outer marginal o o o teeth with serrations. Reproductive system (Figures 7, 8, 13, 14, 15, 17): Go- nad (figure 15, go) relatively small, on posterior left re- gion of visceral mass, surrounded bv left branch of shell muscle, right surface of digestive gland, and some intes- tinal loops. A ver)' narrow gonoduct on right side (not fully examined), running in anterior direction and api- calK' toward right side of mantle cavity, where it opens. Genital pore (figure 17, gp) turned ventrally, just pos- terior to anus. Seminal groove (figure 17. sg) beginning on genital pore, running on integument surface for short distance, up to posterior region of base of right tentacle. Right tentacle broad, sfightlv flattened dorso-ventnilly, graduallv tapering to flat tip (figures 7, S, 13, 14, 17, rt). Edges of seminal groove on posterior surface of tentacle, ending on posterior region of tip. A well-developed sub- terminal protuberance present on anterior surface of tentacle (figure 14). Type material: Holotype MZSP 32150, 3.25 mm length X 2.50 mm width X 1.60 mm height, from t\pe localitv, P. Bouchet, J. Leal, and B. Metivier, 27 May 1987, dead shell. Paratypes: MNHN unnumbered, 3.3.5 J. H. Leal and L. R. L. Simone, 2000 Page 67 mm length X 2.35 mm width X 1.75 mm height, MZSP 32149 (same specimen, soft parts onlv), E of Caho Siio Tome, off Rio de Janeno .State, Brazil,'2r24'S, 39°56'W, 1320°1360 m depth, RA' Mahoit-Dufrcsnc cruise MD55, station CP-04, R Bouchet, J. Leal, B. Metivier, 9 Mav 1987, bottom of basaltic gravel and o.>ddized iron pelibles; MZSP 32151, 3.50 mm length X 2.35 mm width X 1.90 mm height, Doce Ri\er Canvon, off Es- pinto Santo State, Brazil, 19°38'S, 38°43'W, 960 m depth, RA' Marion-Diifrcsnc cruise MD55, station CB- 95, R Bouchet, J. Leal, B. Metivier 30 May 1987, bottom of basaltic gra\el and o>dcbzed iron pebbles (shell broken during SEM session in Miami). Tjpe locality: Continental slope SE off Abrolhos coral reef system, off Bahia State, Brazil, 19°00.4'S, 37°48.8'W, 950-1050 m depth, compact dark mud and shell hash bottom, (RA' Marion-Dufrcsne cruise MD55, station DC-72). Geographic distribution: Continental slope off E-SE Brazil. Bathymetry: 960-1320 m. Et\Tnology: Named after Prof. EUezer de C. Rios, en- thusiastic Brazilian author and mentor to an entire gen- eration oi malacologists. DISCUSSION Specimens of Copniabi/.ssia rio.si were found apparently removed from the actual organic substrate ol the species (see Lesicld, 1998, for a Usting of species-specific sub- strates and food preferences in the family). The new species differs morphologically from the other four spe- cies allocated in the genus Copiilabyssia (see figures 25- 28, table 1) by the presence of an apical septum on the inner shell surface, narrow sculptural interspaces, fila- mentous, longer epipodial tentacles, presence of a stnic- ture on the left side of subpallial cavity that could proxe to be a vestigial left gill leaflet, and by the presence of an anterior, subterminal protuberance on the right ce- phalic tentacle. The double insertion of the m4 and the expanded insertion of the m5 are character states unique to Copiilabyssia nasi when compared to other specie.s of Coccuhnoidea and Lepetelloidea for which these characters are known (see Haszpnmar, 1987; Simone, 1996); however, the states of these characters are so far unknown for the other four species ol the genus Co- piilabyssia. The mantle organs of Copiilabyssia riosi are typical of the genus, with nuchal cavity and gill leaflets situated in the right side of the mantle cavity. Copulabyssia riosi shows a satellite fold in the inner margin ol the gill, and a small protuberance that could represent a vestigial gill leaflet in the left region of the mantle border. In the superfamiK' Lepetelloidea, similar structures are also present in the Addisonia cnodis Simone, 1996 (Lepetel- loidea, Addisoniidae). The arrangement of loops of the digestive system looks similar to that descrilied by Has- zpmnar (1988: 167-168, fig. 4) kr Amphiplica ktiiidseni McLean, 1988 (Lepetelloidea: Pseudococculinidae), but differs by the reduction of salivarv glands, jaw plates, and e.sophageai pouthes. The complex odontopliore oi Copiilabyssia riosi pre- sents a combination oi apparently plesiomorphic and au- tapomori^hic characti'rs The presence of 4 odontophore cartilages and their approximator muscle (m8) repre- sents apparently plesiomoq^hic states present in several species oi Patellogastropoda, Vetigastropoda (including Lepetelloidea), Cocculinoidea, and Neritopsina (L. R. L. Simone, personal observation). The wefl-developed sub- radular cartilage, the partial connection between the an- terior and posterior odontophore cartilages and the hor- izontal muscle (m6) connected only in a side of the an- terior cartilages seem to represent apomorphic charac- ters (at least in relation to Patellogastropoda). ACKNOWLEDGMENTS Gerhard Haszprunar (Zoologische Staatssammlung Miinchen, Germany), Anders Waren (SMNH), and Bruce Marshall (NMNZ) critically reviewed the manu- script. We are grateful to Philippe Bouchet and Virginie Heros at the original repositoiy institution (National mu- seum d'Histoire naturelle, Paris) for making availalile the type material. We are also indebted to Suzanne R. Bra- den (National Museum of Natural History, Smithsonian Institution), for help with some of the SEM illustrations. M. G. Harasewych (National Museum of Natural His- tory, Smithsonian Institution), Anders Waren (SMNH), Bruce Marshall (NMNZ), and Kazunori Hasegawa (NSMT) kindlv sent photographs or negatives of relevant species of Copiilabi/ssia for illustration. This study was supported in part Iiy FAPESP (Funda^ao de Amparo a Pesquisa dcj Estado de Sao Paulo, Brazil) Grant # 96- 6756-2 to L. R. L. Simone. LITERATURE CITED Dantart, L. and A. Luque. 1994. Cocculiniforniia and Lepeti- dae (Gastropoda: Archaeogastropoda) from Iberian wa- ters. Journal of Molluscan Studies 60:277-.31.3. Harasewych, M. G., S. L. Adamkewicz, J. A. Blake, D. Saudek, T Spiiggs and C. ]. 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Harasewych. 1999. Deepest Atlantic mollnsks: hadal limpets (MoJlusca, Gastropoda, Cocculin- ifornua) from the northern boundaiT of the Caribbean plate. Invertebrate Biologv' 118:116-1.36. Lesicki, A. 1998. Checklist of gastropod species referred to the order Cocculiniformia Haszprunar, 1987 (Gastropoda: Cocculinoidea et Lepetelloidea) with some remarks on their food preferences. Folia Malacologica 6:47-62. Marshall, B. A. 1986. Recent and Tertiary Cocculinidae and Pseudococculinidae (Mollusca: Gastropoda) from New Zealand and New South Wales. New Zealand Journal of Zoology- 12:50.5-546. McLean, J. H. and M. G. Harasewych. 1995. Reydew of west- ern Atlantic species of cocculinid and pseudococculinid limpets, with descriptions of new species (Gastropoda: Cocculiniformia). Contributions in Science of the Natural History' Museum of Los Angeles County 453:I-.33. Ponder, W. F and D. R. Lindberg. 1996. Gastropod phyloge- ny — challenges for the 90s. In: Taylor, ]. D. (ed.) Origin and evolutionarv radiation of the Mollusca. Oxford Uni- versity Press, O.xford, pp. 1.35-154. Ponder, W. F. and D. R, Lindberg. 1997. Toyvards a phylogeny of gastropod molluscs: an analysis using morphological characters. Zoological Journal of the Linnean Society of London 119:8.3-265. Simone, L. R. L. 1996. Addisonia enodis, a new species of Addisoniidae (Mollusca, Archaeogastropoda) from the southern Brazilian coast. Bulletin of Marine Science 58: 775-785. Waren, A. 1991. New and little kiioyvn Mollusca from Iceland and Scandinayna. Sarsia 76:53-124. Wolff, T. 1979. Macrofaunal utiUzation of plant remains in the deep sea. Sarsia 64:117-1.36. THE NAUTILUS 114(2):69-73, 2()()() Page 69 Invasion of the exotic freshwater mussel Limnoperna fortiinei (Dunker, 1857) (Bivalvia: MytiHdae) in South America G. Darrigran Departamento Cientifico de Zoologi'a de linertebrados P'atultad Ciencias Naturales v Museo 1900 La Plata ARGENTINA in\asion@wav.coni.ar L Ezcurra de Drago Instituto National de Liiimologfa (INALI/CONICET) Maciii 1933, 3016 Santo Tome Santa Fe ARGENTINA ABSTRACT We traced the im'asion and iinestigated the (.hstiibntion of the Asian bivaKe Limnopenui foriiinci (Dnnker. 1857) in South America. In addition, we comment on the problems caused bv this in\'asive species. In 1991, L. fortiinei was introduced into la Plata Basin (currently the onlv American continent drainage system invaded bv this species) through Ri'o de la Plata River B\' the end of 1994 and during 1995 this species dispersed throughout the Argentine coast of the Rfo de la Plata River and was reported from the Uruguayan coast. In 199.5 and 1996, it was first discovered in the Parana Ri\'er, in Santo Tome ( Sal- ado del Norte River), and later up in Goya. In November, 1996, L. fortiinei was reported in Cerrito Island, where the Paraguay River joins the Parana River In April 1997, it was collected in Paraguay River on Asuncion Harbor, Paraguay. In 1999, if was detected in Brazil (Itapuii Beach, MunicipaUty of Viamao, Rio Grande do Sul State). These recent records doc- ument tlie fast and ongoing e.xjiansion ot the species geograph- ic range, as well as its prompt adaptation to the ditterent en- \ironments of the la Plata Basin Ki'ij words: Soutli America; Neotropical Region; la Plata Basin; invasive species; distribution; biofouling. INTRODUCTION The la Plata Basin is one of the most important hydro- graphic systems in South America. It has a drainage area of approximately 3X10'' km-, and comprises 4 main sub- hasins (Bonetto, 1994): (1) Uniguav River, (2) Parana- Paraguay system, (3) Rio de la Plata River, and (4) the Andean tributaries (figure 1). A diverse, native molluscan fauna occurs along the Ar- gentine shore of the Rio de la Plata estuan' (Darrigran, 1994), including 27 gastropod and 21 bivalve species. Only 4 of the bivalves are strictly freshwater and intaunal species: Anodontites tenebricosus (Lea, 1834), Diplodon parancnsis (Lea, 1834), MtisciiUum orfictitimnn (d'Orhigny, 1835) and Pisiclimn stcrkiaiiinn Pilsbi"v, 1897. The only epifaunal species on the Argentine coast of Rio de la Plata River is the estuarine Mi/tclla clior- niana (d'Orbigny, 1842). Until 1993, A/, charniana was the only Mytilidae found in Rio de la Plata River, in Punta Piedras, Argentina and Montevideo, Uruguay (Darrigran and Pastorino, 1995a). In the 1970s, 2 asiatic freshwater bi\aKes, Corhicida largiUiciii (Philippi, 1844) and C. fliiininca (Miiller, 1774) (Corbicuhdae), entered South America along the Argentine shores of Ri'o de la Plata River (Ituarte, 1981). In 1991 we detected, for the first time in South Amer- ica, the presence of a mytilid, Limnoperna foiinnci (Dunker, 1857), in the Rio de la Plata River shore at Baghardi Beach (Pastorino et al., 1993). This is an eu- ryhaline freshwater species, native to China and to southeastern Asia (Morton, 1977), which resembles ma- rine mussels (Mytifidae) in its moiphology and biology. Commerce beUveen Argentina and the countries where L. fortiinei is supposedlv native takes place m;unly by sea. Following Carlton (1992) and Carlton and Geller (1993), Darrigran and Pastorino (1995a) hypothesized that L. fortiinei and other invading organisms may have arrived in ballast water of ships coming from southeast- em Asia. Other authors have inthcated that L. foi-fiinei was in- troduced in and recorded from Hong Kong in 1966 (Morton, 1975, 1996) and Japan in 1991 (Kimura, 1994). In the Americas, L. foiiiinei is known to have invaded only Rio de la Plata Basin. VIorton (1973) suggested that the morpho-functional features of this species allow for rapid expansion of its distribution range, in similar fash- ion to the invasive abilities of the zebra mussel, Drei.s- sena polijmorpha (Pallas, 1771), in the Northern Hemi- sphere. The species exhibits 2 features that mav contrib- ute to its abilities as an invader: (1) it is the onlv relevant mollusk in the freshwater littoral ha\ang a bvssus, and (2) has no known native competitors for physiciil space. As a result of the high rates of invasion, as we further chscuss below, L foiiiinei causes serious biofouling of municipal drinking water and industrial water svstems. As we demonstrate below, populations of Limnopenui fortiinei are not onlv rapidly increasing in densit\- in sev- Page 70 THE NAUTILUS, Vol. 114, No. 2 tial localities but are also quickly expanding geographi- cidly, mostly in northward direction. Since 1991, when it was first found at Bagliardi Beach, until the end of 1993, the species was found only in mLxohaline (e.g., Punta Piedras and Punta Indio) and eur\'haline environments (e.g., Magadalena Beach, Ata- laya Beach, Punta Blanca, La Balandra Beach). At that time, the species had not been collected farther north than the freshwater environment of Punta Lara Beach, and was apparently absent from all other countries and rivers comprising the la Plata Basin (Darrigran and Pas- torino, 1993). The objectives of this paper are to investigate the in- vasion of L. foiiunci in South America, to provide an update of its cbstribution in the Neotropical Region, and to emphasize the importance of this invasive bivalve in biofouling processes in the la Plata Basin. MATERIALS AND METHODS Collections were made along the Rio de la Plata River shore at low tides. In the Parana Ri\'er, samples were collected with a 319 cm- "Tamura" dredge (Marchese and Ezcurra de Drago, 1992). Periph\tic populations were sampled by hand. In the mouth of Paraguay River, "macrofouling" samplers (Stupak et al, 1996) were used. Environmental parameters taken into account were: temperature, pH, and conductivity. The study localities are showni in figure 1. Specimens from the Malacological Collection of La Plata Museum, Argentina, (MLP) were examined. Samphng for larvae was made using a mesh of 25 (xm in Umguav River (towns of Colon and Con- cepcion del Uiiiguay), at 3 stations in the Carcaraiia Riv- er, and Correntoso River RESULTS AND DISCUSSION Liinnopenw foifunei was first found in South America in 1991 at Bagliardi Beach (Argentina) with densities of 5 individualsm- (Pastorino et al, 1993). In 1992, the maximum densitv' recorded was about 3X10^ indixidu- als-m -. In 1993, in the same locality, the maximum den- sity recorded was SXIO' individualsm -. In 1998, den- sities of about 15X10' individuals-m"- were commonly recorded, values equivalent to those for mytilid densities at the seashore in Mar del Plata, Argentina (Penchasz- adeh, 1973). In 1994 and 1995, L. fotiiinei was found in the water intakes of the town of Bemal (MLP 5204) and Retiro, and on Buenos Aires Harbor (MLP 5205). Scarabino and Verde (1994) reported the species from the town of Colonia del Sacramento, Uruguay (MLP 5202). All these localities are on the Rio de la Plata estuaiy. During 1996, the species expanded to the town of Goya (29°10'S, 59°16'W), in Corrientes Province (Di Persia and Bonet- to, 1997) and nearby lotic bodies, and was collected in the middle Parana River, near the city of Santa Fe, where the river has a \'erv width; Drago, 1990). wide alluvial plain (25 km in Figure 1. Studied localities on the la Plata Basin. A, Punta Piedras; B, Punta Indio; C, Magdalena beach; D, Atalaya Beach; E. Pmita Blanca; F, La Balandra Beach; G, Bagliardi Beach; H, Pnnta Lara Beach; L Benial Beach; ]. Buenos Aires Harbor; K, Atncha I Nuclear Power Plant; L, City of Rosario; M, Town of Santo Tome; N, City of Santa Fe; O, Esquina; P, Goya; Q, Cerrito Island; R, Town of Formosa; S, Asuncion Harbor; T, Posadas Harbor; U, Praia de Itapna (Rio Grande do Sul State, Brazil); V, Town of Colonia del Sacramento; 1, Town of Colon; 2, Town of Concepcion del Umguav; 3, Car- caraiia River. Litnnopcrna foiiunci was first collected in Parana Riv- er at Vuelta del Este, Zarate, on the Parana de Las Pal- mas River (MLP 5206), and at Paso Burghi, city of Ro- sario, on the lower Parana River (MLP 5207) toward the end of 1995 (figure 1). The species settles on all kinds of substrates across the entire floodplain. It was found in the main channel, on the left bank at town of Parana and associated alluvial plain, in San Javier and Corren- toso (MLP 5287) rivers, near the city of Santa Fe. It was also found on the right bank of the lower Salado del Norte River (MLP 5285), in the town of Santo Tome (31°40'S, 60°45"W). Limnopcnia foiiinwi was part of the epifauna present on Solanaceas and Paspahun .sp. (MLP 5286) in all hab- itats observed, but was not found on the sandy, moving bars (Drago, 1997) in the main channel. Perhaps due to the unfavorable sandy substrate, the central part of the G. Darrigran and I. Ezcurra de Drago, 2000 Page 71 main channel pro\ides haliitat to onK- a few henthic spe- cies (Marchese and Ezcurra de Drago, 1992). In the Correntoso River (a secondary channel of the middle Parana Ri\er\ L. foiiiiiici i-olonized the silt\ cla\ lieds at all locations sanipleil The eun,haline attributes of L. foiinnci probably fa- cilitates its rapid spread. The mean salinitv at the main channel of the Parana River is 0.05 %r, and in Corren- toso River it ranges bet\veen ().()fi4-().345 %c. In Salado del Norte Rixer, which mi.\es with the waters of the Pa- rana River, sahnitv varies from 0.5 to 4 %c (Ezcurra de Drago, personal obser\'ation). Mean pH is 7.4 in the Pa- rana m;un channel and 7.2 in secondaiy courses (Mar- chese and Ezcurra de Drago, 1992), \s'hile in Salado tlel Norte River pH reaches 8.7. In 1996, this species was collected on "fouling" sam- plers in Cerrito Island (27°20'S, 5S°43'W), near the con- fluence of the Paraguay and Parana rivers (MLP 5340). In April 1997, it was collected on Asuncion Harbor (25°17'21"S, 57°38'0S"W), in the Paraguay River, in Par- aguay (figure 1). It was also collected during the last months of 1998, at Posadas Harbor, on the upper Parana River. In November 1999, L foiiimci was detected in Brazil on Itapua Beach, Municipality of Viamao, Rio Grande do Sul State (MLP 5550). No larvae, juveniles or adults have been found at anv of the locahties ex- amined in UiTiguav- River (Colon, Concepcion del Uru- guay) or Carcarana River (Carcaraiia and la Ribera). The biofouling problems caused by L. fortunei in South America are similar to those described for Drcis- scna pohjinoiyha, the zebra mussel, in the Northern Hemisphere (Darrigran, 1995). The principal problems caused by zebra mussel in- vasion, settlement, and maturity, into water distribution systems (Nalepa and Schloesser, 1993) are listed below (the problems which have already been detected in Ar- gentina resulting from invasion bv L foiiuiici are marked with (X)). • ( X ) Reduction of pipe diameter • ( X ) Blockage of the pipeUne • Decreased water velocitv caused bv friction (turbulent flows). • (X) Accumulation of enipt\' shells • Contamination of water pipelines by mass mortality • ( X ) Filter occlusion These problems have occurred in the intakes of water treatment plants in the city of La Plata; water treatment plants along the Corrientes River (MLP 5365), and also in industrial and power-generating plants. They cause an increase in the operational costs of these facilities due to reduced pump efficiency, increased tube corrosion (caused by proUferation of bacteria and fungi), and in- creased frequenc)' of shutdownis for cleaning and filter changes. Absence of relevant records in the numerous benthic studies conducted prior to 1991 (Darrigran, 1991, 1994; Marchese and Ezcurra de Drago, 1992) indicate that colonization by L. foiiiinci is a recent event. By late 1996 and early 1997, the first cases of fouling bv L. fniiunei in nearby industries were reported in the Parana River (e.g., Atucha I Nuclear Power Plant and San Nicolas de Los Arroyos Electric Plant (MLP 5300), Buenos Aires Province). Based on the facts above described, it appears that the geographic range of L. foi-tunci is in continuous ex- pansion in South America, and that the species is ap- parently adapting to different environments along its in- vasion routes. Between 1991-1999 this species invaded 4 c(juntries (Argentina, Uruguay, Paraguay, and Brazil); 3 of the Ri'o de la Plata Basin main rivers (la Plata, Pa- rana, and Paraguay). It traveled upstream at a speed of about 240 kmvear '. Throughout its distribution range, L. fdiiiinci inhabits a wide variety of environments: from euryhaline waters in the La Plata estuary to strictly freshwater habitats along the Paraguay and Parana riv- ers. This expansion is proiiably favored by the moqiho- functional features of the species, and by the apparent absence of competitive interactions along the httoral of la Plata Basin. Lininopcnio foiitinci is the onlv relevant species using bvssal epifaunal attachment among local freshwater species. Although the distribution of L. foiiunci is currently still limited to la Plata Basin, there are no apparent con- straints that could prevent the invasion of other drainage systems. Moreover, this expansion may be favored by the increase in trade between Argentina, Uruguay, Paraguay and Brazil, as a consequence of the recently establi,shed MERCOSUR international economic consortium (Dar- rigran, 1995; Darrigran and Pastorino, 1995b). Addition- al dispersal of the species could also be favored by the international project HIDROVIA, which consists of a wateiAvay connecting the mtiin hvdrographic systems of South America. The project comprises a complex fluvial network system that involves and interconnects Rio de la Plata, Parana, and Paraguay rivers, and includes plans for extensive basin dredging, harbor construction, and improvement of the harbors and highways. Connections among the main South American basins (e.g., Amazonas, Orinoco) are likely to be artificially established in the future. Up to now no effective control mechanisms have been proposed to prevent further spread oi L. fortunei. The impact of L. foiiintci will not be restricted onlv to the economy, but will also affect the diversit>- of the native molluscan communities. Darrigran et al. (1998) showed that, since the introduction o{L. fortunei at Bag- liarili Beach, populations of 2 common gastropods have been displaced: Ciiilina jiuminea (Maton, 1809) is no longer found, while Gundhiehia conccnfiica (d'Orbigny, 1835) is becoming rare (figure 2). In contrast, several benthic species, uncommon or ab- sent before the occurrence of L. fortunei. are now pre- sent. These include Annelida Ofigochaeta (8 species), Aphanoneura (1 species) and Himdinea (8 species); plus assorted Crustacea and Insecta (Darrigran et ai, 1998). Additionallv, we observed the epizoic colonization and snujthering of native bivalves (e.g., Anodontite.s trapesi- Page THE NAUTILUS, Vol. 114, No. 2 DENSITY Helobia piscium 10' ■• Chilina fluminea I . ]«|«jB|*|«|«|«|*|B|«|*|B| I ■ I ■ I ■ I ■ I ■ Gundlachia concentnca I'i'i'i'i'i — I * I ■ I ■ I ■ I Limnoperna fortune! H 1 1 1 1 -•-h ( 1 1 1 (-1 — I 1 h ■3 7 8 9 10 12 1 2 3 5 8 10 12 |l 2 3 4 6 9 10 11 I 3 88 I 92 |93 194 I 95 MONTH/YEAR Figure 2. Temporal \ariation of density (individuals-m"-, represented bv mean and standard deviation) of some native mollusks associated with the byssus of L, fortiinei in Bae;liardi Beach (34°55'S, 57°49'W), Wentina. alis (Lamarck, 1819) and A. tcncbricosus (Lea, 1834)) by L. foiiiinei, in a behavior similar to that shown by Dreis- seno poli/inoi-f)ha on unionids (Parker ct al, 1998). Last but not least, further expansion of the natural range ot Limnoperna foi-tunci and the possibilitv of in- vasion of parts of Noith America by this species should not be rejected. As Ricciardi (1998) obserx'ed, "Given that shipping traffic from both Asia and South America has alreadv resulted in recent introduction of exotic bi- valves to the USA, a future North American invasion bv L. fortunci is highly probable." ACKNOWLEDGMENTS The work was supported by the grants of the Consejo Nacional de Investigaciones Cienti'ficas y Tecnicas (PEI N° 548/97), Facultad Ciencias Naturales v Museo (UNLP), and Agencia Nacional Promocion Cientifica y Tecnologica (PICT98 No. 01-03453). We thank Dr. Ri- chard Sparks and Ruth Sparks for the revision of the manuscript. Dr. Edmundo Drago and M. C. Dambore- nea for helping in assorted tasks, and Lionel Mehaudy for the technical assistance. Maria CrisHna Dreher Man- sur collected and provided the specimens from Itapua Beach, Rio Grande do Sul, Brazil. REFERENCES Bonetto, A. A. 1994. Austral rivers of South America. In: Mar- galef, R. (ed.) Limnology now: a paradigm of planetary problems. Elsevier Science, Amsterdam, pp. 425-472. Carlton, ], T. 1992. Introduced marine and estuarine mollusks of North America: an end-of-the-20''' centurv perspective. Journal of Shellfish Research 11:489-505. Carlton, ]. T and J, B. Geller 199.3. 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Zebra Mussels and Aquatic Nuisance Species. Ann Arbor Press, Inc. Chelsea, Michigan. Nalepa, T, and W Schloesser. 1993. Zebra mussels: biolog)', impacts, and control. Lewis Pviblishers, Boca Raton. 508 pp. Parker, B. C M. A. Patterson and R. J. Neves 1998 Feeding interactions between native freshwater mussels (BivaKia: Unionidae) and zebra mussels {Drcisscna i>oli/morplia) in the Ohio Ri\er American Malacological Bulletin 14:17.3- 179. Pastorino, G., G. Darrigran, S. Martin and L, Lunaschi. 1993. Liinnopentri foriiiitci (Dunker, 1857) (Mytilidae), umcm) bixalvo invasor en aguas del Rio de la Plata. Neotropica 39(101-102):34. Pencha.szadeh, P. 1973. Ecologfa de la comunidad del mejillin (Brachyclontes rodriguezi d'Orb.) en el medioiitoral ro- coso de Mar del Plata (Argentina): el proceso de recolon- izacion. Physis 32A(84):5I-64. Ricciardi, A. 1998. Global range expansion of the Asian nnissel Limnoprnin foiiumi (Mvtilidae): another fouling threat to freshwater .systems. Biofouling 13:97-106. Scarabino. F. and M. N'erde. 1994. Limnoperna foiii/iwi (Dtm- ker 1857) en la costa unigua\a del Rio de la Plata (Biv- alvia, Mytilidae). Connmicaciones Sociedad Malacologica del Uruguay 7 (66-67): 374-375. Stupak, M., M, Perez, M. Garcia Sola, A. Leiva Azuaga, A. Mendivil and G. Niveyro. 1996. Preliminary study of the biofouling of the Parana Rix'er (Argentina). Corrosion Re- news 14:145-155. THE NAUTILUS 114(2):74-79, 2000 Page 74 Location of allospermatozoa in the freshwater gastropod Biomphalaria fenagophila (d'Orbigny, 1835) (Puhiionata: Planorbidae) Warton Monteiro Depaitiimento de Ecologia Uiu\ersidade de Brasilia Brasilia, DF BRAZIL warton@uol.c-oni.br Toshie Kawano' Laboratorio de Parasitologia Instituto Butantan A\'. Vital Brasil, 1500 0550.3-900 Sao Paulo BRAZIL The freshwater puhnonate Biomphalaria fenagophila is a simultaneous hermaphrodite. The route ot allosper- matozoa in the receiving partner was used in the analysis of resource allocation in the male function in this spe- cies. Allospermatozoa enter the vagina, mav or may not bvpass the spermatheca, continuing through nidamental gland, o\'iduct-muciparous gland, albumen gland, oNds- permioduct, seminal vesicle, ovotestis collecting canal, and folhcles of ovotestis. A new technique is proposed to label spermatozoa xaa injection oi tritiated th\iiiitline through the pneumostome (under anesthesia). Labeled allospermatozoa were located in the spermatheca, ni- damental gland, oviduct-muciparous gland, albumen gland, seminal vesicle, and follicles of ovotestis. No re- lation was found between the duration of pairing and movement of the allospermatozoa through the repro- ductive system. It is suggested that the final destination of allospermatozoa are the folhcles of ovotestis. Their presence in the spermatheca, where thev ma\' be dis- solved, could be fatal. Presence of allospermatozoa in the nidamental gland, oviduct-muciparous gland, or ;il- bumen gland may be a consequence of unilateral cop- ulation (snail plavs only a recei\ing role), which packs the seminal vesicle with autospermatozoa. Allosperma- tozoa may wait in the interior of the female tract until the snail becomes a donor in a subsequent pairing. From then on, allospermatozoa may pass through an empty seminal vesicle toward their final destination, the folh- cles of ovotestis. Key words: Reproduction, spermatozoa, hermaphrocUt- ic, pulmonates. INTRODUCTION The location of spermatozoa within the reproductive sys- tem of inseminated puhnonate gastropods plavs an im- ' Author for correspondence; tkawano@usp.br portant role in the processes of self-fertilization, cross- fertilization and, in particular, preferential cross-fertiliza- tion in these mollusks (Larembergue, 1939; Paraense, 1959). In the genus Phijsa, 2 or 3 possible chambers for fer- tilization were described. Although these chambers open in the oviduct, the actual site of fertifization may be lo- cated in the hermaphrodite region, close to the carrefour (Duncan, 1969). In planorbids, the seminal vesicle has been cited as the main location of spermatozoa (Lar- ambergue, 1939). The fertilization chamber, in the car- refour, where the spermatozoa received from the part- ner are stocked, is also a possible ferhhzation site (Ab- del-Malek, 1954; Duncan, 1975). The oviduct (Fretter and Graham, 1964) and the upper part of the o\isper- mioduct mav be sites where the oocytes are fertilized by the allospermatozoa (Abdel-Malek, 1954). Some authors consider the spermatheca as a location where sperma- tozoa remain immecfiatelv after copulation, proceeding subsequently through the female ducts (Duncan, 1975). Notwithstanding, the spermatheca in B. glahrata also behaves as a site of lysis of spermatozoa (Horstmman, 1955; Jong-Brink, 1969; Kitajima and Paraense, 1983). In planorbids, the spermatheca is a blind, pvriform or- gan, opening in the vagina through a short duct (Par- aense, 1975; Kitajima and Paraense, 1983). This is also called bursa copulatrix (Duncan, 1975), or seminal re- ceptacle. In euth\aieuran gastropods, the evolutionary acquisition of reproductive s\stems such as functional simultaneous hermaphrocUtism has been associated with, among other traits, fundamental changes in the structure of the spermatozoa (Thompson, 1973). A past experiment iuNolving incorporation ot '"Fe by BioinpJialaria tAahrata took place in water containing this rachoisotope (Gazzinelli ct al. 1970). Incoiporated isotope was transferred, during pairing, from labeled B. glabrata to non-labeled partners. The presence of the rachoisotope in snails receiving spermatozoa was record- ed chiefly in the spermatheca and collecting canal of the ovotestis (Paraense, 1976). W. Monteiro and T. Kawano, 2()()() Page 75 Bioinphdldhd i'iitti is known tu pivfcTfntiallv con- centrate iron in the (notestis, as well as in non-repro- tluetive organs such as chgestive gland and mantle (He- neine et al, 1969). Therefore, it is plausible that ^'^Fe transferred from labeled to non-labeled snails could be incoiporated in the ovotestis of receptor snails, indepen- dentK of spermatozoa being the carrier of ''Fe within the receptor suiiil. On the other hand, the use of DNA-specific labels can be an efficient wav of tracking the location of allosper- matozoa in the inseminated snail partner. Tlie route of spermatozoa heads in the inseminated snail can be de- termined by the use of tritiated thymidine ('H Td). This method also allows for the detection of possible areas of storage and sites of cross-fertilization. MATERIALS AND METHODS One hundred and six specimens of Biomphalaria tcna- gophila were used both of the wild txpe and albino mu- tants. Specimens were collected in Bom Retiro, Joinville, Santa Catarina State, Brazil. They were kept for over 4 vears under laboratorv' conditions. Each aquarium con- tained a tablespoon of red earth sterilized in <\t\ incu- bator at 90°C for 12 h. One hundred and eightv ml fil- tered water, saturated with calcium carbonate, was add- ed to each aquarium. Snails were fed daily with fresh lettuce and twice a week with a meal consisting of equal amounts of red earth, powdered whole milk, wheat germ flakes, powdered calcium carbonate, and dry powdered alfalfa. To a 5 1 amount of diis mixture 350 mg of vitamin E was added. Except in some cases, snails were kept in isolation from sexual immaturity onward. Two suppfies of aqueous solutions of 'H Td were used in the ex-periments. Ten pi of one (USP, kindlv offered by Dr. Mariano Amabis, University of Sao Paulo) yielded 218687 counts per min. (cpm); the other commercial solution (\VM) was purchased as 1 millicurie, or .37 me- gabequerel/ml; 5|xl from this solution \ielded 314850 cpm in a Beckmann Liquid Scintillator 7200, using "pro- gram 2 for tritium". Each of the 106 specimens was anesthetized by immersion in a 0.05% nembutal solution for 3^ h, depending on snail size. Aftei-ward, they were inoculated with 5-10|jl1 of 'H Td injected using micro- syringe into the lung cavity, through the pneumostome. Recovery from anesthesia took place in a humid cham- ber, for 3—4 h, with each snail placed on its right side in a plate with water covering half shell. After this period, each inoculated snail was returned to its aquarium for completion of recovery from anesthesia and radioisotope incoiporation. Between 4—29 days after treatment, each inoculated snail (IC) was paired with a non-inoculated snail (NIC) for a period of 6 — 984 h. NIC snails were dissected un- der stereoscopic microscope within a period of 0—24 h after separation. The following organs were extracted: spermatheca, oothecal gland + muciparous gland, o\i- duct or oothecal gland by itself, but muciparous gland with oviduct, part of digestive gland, carrefour, albinnen Figure L Diagram of tlie reproductive system of Biomphal- aria tcnaf^ophiln. 1. Hind-part of ovotestis; 2. Follicles of the ovotestis; 3. Ovotestis collecting canal; 4. Seminal vesicle; 5. Albumen gland; 6. 0\'idiict; 7. Muciparous gland; 8. Oothecal gland; 9. Nidamental gland; 10. Spermatheca; 11. Vagina; 12. Spermioviduct; 13. Prostate; 14. Deferens canal; 15. Penis sheath; 16. Prepntiiim gland, seminal vesicle and ovotestis (Figure 1). Counting in a section of the digestive gland determined the amount of incorporation of radioisotope outside the re- productixe system, the background radioisotope incor- poration. The ovotestis was macerated releasing material from the collecting canal and follicles. These contents were collected separately from the rest of the macerate. Each sample was transferred to a vial containing 0.3 or 0.5 ml of a solution of 3 % sodiiun hvpochloride (com- mercial bleach). The material was repeatedly refluxed with a Pasteur pipette until completely dissolved. Prep- aration of filter paper strips imbibed with samples for measuring in the liijuid scintillator was based on the methodology of Byfiekl and Scherbaum (1966). Though spermatozoa may be found and counted in chfferent organs of the reproductive system during and after the pairing process (Monteiro and Kawano, 1998), in this work the occurrence of allosperTnatozoa in the examined organs was accounted tor only by counting in the hquid scintillator Some controls were used to test the efficiencv of the method. Processing of each series of paper strips with samples included a paper strip without sample, a "blank" that allowed us to investigate the amount of free racho- active material that could be absorbed from solutions where the papers with samples were being treated (Ta- ble 1). The radiation in the water where the snail recovered from anesthesia was measured by scintillation to evaluate the efficiency of the inoculation in the lung cavitv and the draining of H Td into the mechum during the re- coveiy period. The amount of radiation injected into each snail was also measured (Table 1). RESULTS AND DISCUSSION The technique of inoculation in the lung ca\it\ through the pneumostome of the anesthetized snail was here used for the first time. The evaluation of this technique Page 76 THE NAUTILUS, Vol. 114, No. 2 Table 1. Experiments with different supplies of tritiated th^inidine (USP and WM supplies) minute. Blanks = data not available Cpni = scintillation counts per Exp. 1 Exp. 2 Exp. 3 Exp. 4 No. snails 5 5 5 5 Microliters inoculated/snail lOUSP lOUSP lOUSP 5WM Inoculation-pairing (days) 7 7 11 6 Duration of pairing (h) 24 24 24 24 Pairing-dissection (h) 52 48 216 40 Separation-dissection (h) 28 24 192 16 Inoculation to-dissection (approx. days) 9 9 20 8 Diameter of snails (mm) 12-14 12-13 12-13 13-14 Reference (cpm) 27507 26911 23534 Background (cpm) 71 65 84 75 Blank (cpm) 105 119 0\nduct (cpm) 109 89 78 0\'otestis collecting canal (cpm) 91 84 76 105 Follicles of ovotestis (cpm) 174 84 107 121 Spermatheca (cpm) 81 114 114 110 Nidamental gland (cpm) 96 890 884 Oothecal gland (cpm) 135 Seminal vesicle (cpm) 1184 63 139 121 Digesti\'e gland (cpm) 87 104 Carrefour (cpm) 76 107 Albumen gland (cpm) 427 0\iduct -muciparous gland (cpm) 102 ° USP, supplv kindh' offered by Mariano Amabis, Universitv of Sao Paulo: WM, supplv purchased by one of the authors. was carried out in 18 snails inoculated with a total oi 90 jjlI of ^H Td. Retention of inoculate in the lung cavity was tested through measurements of the relative amount of radioisotope that drained during the 4 h recovery from anesthesia. The snails recovered in 27.5 ml of water that yielded 81152.5 cpm. As each 5 |xl of the applied radioisotope solution counted 314850 cpm, there was, on average, a 1.4 % loss of radioactive material from snails to the water. This result encourages the use of this inoculation method in tests of prospective toxicology, se- lection of mutagens, and of the mechanism of action of water pollutants. The results of 1 1 out of a total of 30 e.xperiments are shown in Table 1. The remaining experiments did not record high counts for any of the examined organs. The low counts in all examined organs may have occurred in sniiils that did not act as females in relation to the in- oculated partner. An arbitraiy criterion was established; to consider onlv the experiments in which the counting in the most labeled organ was at least twice as those in the least labeled one. As a natural consequence of metabolism, the long du- ration of an e.xperiment might cause the incoqjoration of the radioisotope in an organ absolutely lacking sper- matozoa. This does not seem to be the case here, even in experiments lasting for 216 and 948 h (e.xperiments 3, 10, and 11), from pairing to dissection time. In these conditions, the most labeled organs were the nidamental gland (884 cpm), the albumen gland (210 cpm) and fol- licles of ovotestis (365 cpm), respectively (Table 1). We looked for possible transfer of the radioisotope label from the reproductive system to a highly metabolically active organ such as the chgestive gland. A tragment of digestive gland, similar in volume to the ovotestis, was simultaneously tested in experiments 2 and 3 (respec- tively 48 and 216 h after pmring). The relatively low counts, 87 and 104 cpm respectively, discarded that pos- sibilit)' (Table 1). After a specimen of B. glabrata behaves as female during pairing, its spermatheca is packed with sperma- tozoa. That organ is site ot lysis of spermatozoa (Kitajima and Paraense, 1983). Suiprisingly, among all the e.xper- iments, only one presented relatix'ely elevated countings for spermatheca (206 cpm): experiment 8 (Table 1). Be- sides, in experiment 9, where the time lapse from pair- ing to dissection was only 7 h, the spermatheca counting, 87 cpm, was equivalent to the blank (Table I). Given the countings in the digestive gland (458 cpm), it is possible that the spermatozoa have either bypassed or spent a verv brief time in the spermatheca on their way to the digestive gland. Also, clue to the relatively short time elapsed from pairing to dissection, 7 h, the hypothesis of radioisotope transfer from spermatheca to digestive gland was not taken into account in the analysis of the data. No relation was found between the duration of pairing and progression of the allospermatozoa along the repro- ductive ducts. After only 54 h from the beginning of pairing, in experiment 6, the allospermatozoa were in the folhcles of the most distant organ from the vagina, the ovotestis (585 cpm). But 216 h after the beginning of p;iiring, the iillospermatozoa were in the nidamental gland (216 cpm, experiment 3), the organ closest to the vagina (Table 1). W. Monteiro and T. Kawano, 2000 Page 77 Tabic 1. Exteiuled Exp. 5 Exp. 6 E.\p. T Exp. S Exp. 9 Exp. 10 Exp. 11 4 5 3 .5 3 2 lOUSP 5\\M lOVVM lOW'M lOWM lOWM lOWM 19 7 / 11 19 29 29 96 24 8 9 6 984 984 120 54 8 31 6 984 24 30 0 22 0 ? 0 24 9 -' 12 19 70 12-14 12-14 9-13 9-11 11-12 12-13 11-12 24765 24383 24357 24357 24027 24585 75 103 72 99 99 55 65 119 103 110 87 87 92 77 103 117 89 76 68 68 787 585 106 89 73 59 365 168 93 172 206 87 76 147 133 120 74 77 78 71 84 111 105 275 74 57 75 68 115 166 119 92 458 210 63 314 131 9.3 84 81 S7 65 The simultaneous occurrence of high counting in the toUicles of o\otestis and in the oviduct-muciparous gland in a single experiment (experiment 5), after 96 h of pair- ing, suggests the occurrence of double copulation (Dias, 1995). Spermatozoa transferred in the initial copulation could be located in the organ most distant from the va- gina, the folHcles of ovotestis. The spermatozoa trans- ferred in a subsequent copulation may be located in an organ closer to the vagina, such as the oviduct-mucipa- rous gland (Table 1). This explanation, however, could be inconsistent. Even if the last copulation happened at the end of pairing, the 24 hours that passed after the separation of the partners would be enough tor these spermatozoa to advance along the genital tract, bxpass- ing the oviduct-muciparous gland. In a shorter period of time, from pairing to dissection, allospermatozoa pro- ceeded to the seminal vesicle and albumen gland, re- spectively, in experiments 7 and 9. The results of exper- iment 3, however, show that it is possible that the sper- matozoa remain in the nidamental gland for up to 192 h after the separation of the partners. Therefore, under the experimental conditions, the scintillation counting may indicate that, in a single or in diiferent snails, from a group of 4 (experiment 5, Table 1), the spermatozoa may be located in sites as separated as the ovotestis and organs closer to the vagina such as the oviduct-mucipa- rous gland. It is possible that, rather than suggesting ran- dom differences in locomotion abilit), these results could indicate strategic, differential placement of allosperma- tozoa. All the occurrences of allospermatozoa along the re- productive system, registered in the nidamental gland, seminal vesicle, albumen gland, and iolhcles of the ovo- testis are difficult to explain At first sight, the nidamen- tal gland seems to be a transient site ot allospermatozoa on their way to their final destination, where the\- will fertilize the oocytes (Paraense, 1959). However, their stay in the nidamental gland for 216 h from pairing to dissection time (experiment 3, Table 1), a period of time much longer than that necessary for the spermatozoa to reach the follicles of ovotestis (experiments 5 and 6: 120 h and 54 h, respectivelv; Table 1), contradicts this pre- mise. Considering that the seminal vesicle of the donor may be empty after pairing (Monteiro and Kawano, 1998), the presence of labeled allospennatozoa in this organ (experiments 1 and 7) suggests that autosperma- tozoa were replaced b\' allospermatozoa. This inteq^re- tation apparently conflicts with the occurrence ot labeled allospermatozoa both in the folficles of ovotestis and in the albumen gland, both within short and long intervals from pairing to dissection (e.xperiments 4, 5, 6, 9, 10, and 11). No relation was found between the duration of pairing and progression of the allosperm through the reproduc- tive system. The labeled allospermatozoa were located in the spermatheca, nidamental gland, oviduct-mucipa- rous gland, albumen gland, seminal vesicle and follicles of ovotestis. It seems that the destination of allosper- matozoa are the follicles of the ovotestis. Their occur- rence in the spermatheca, where Ksis takes place, may be highly detrimental or fatal. Copulation in B. tcna^o- phila may be unilateral or reciprocal (Dias, 1995). Con- sequently, the length of stav of labeled allospermatozoa in the nidamental gland, oxiduct-muciparous gland, or albumen gland may be the result of imilateral copula- tion, when the snail played only' a receiving role, which Page 78 THE NAUTILUS. Vol. 114. No. 2 resulted in a seminal \esitle full of autospermatozoa. Al- lospermatozoa may wait within the female tract until the snail plays a donor role in a subsequent pairing, which can be unilateral or reciprocal. Then, allospermatozoa ma\' pass through an empt)^ seminal \esicle toward the follicles of the ovotestis. Within an o\otestis follicle, in the oocyte formation site and co-existing with immature autospermatozoa (Homan, 1972), allospermatozoa may fertihze the oocytes (Laramberg, 1939; Paraense, 1955). The many possible locations for strategic storage of al- lospermatozoa suggests displacement of autospermato- zoa, which makes possible the appropriate deplo\nnent of allospermatozoa. Allospennatozoa were present in ovotestis follicles in experiments 5, 6, and 11, but were not found in the collecting canal of ovotestis in any of the 11 experiments. NoUvithstanding. Paraense (1976), labeUng snails with '"Fe, found e\idence of allospermatozoa in the collecting canal. A possible explanation for this result is the re- incorporation of iron. In fact, the ovotestis has a noted avidity for iron (Heneine ei a/., 1969; 1970a; 1970b). Of all these locations, the one more consistent with data from the hterature is the alliumen gland, although all searches for spermatozoa in the canals of the gland and nearby areas during the preparation of the present work have yielded negative results (not even spermatozoa heads were found). Nevertheless, a chamber of fertiliza- tion, in the carrefour, next to the albinnen gland, is often referred as the final site for storage of spermatozoa in studies carried out on several other species of moUusks (Duncan, 1958; 1960; Lind, 1973; Geraerts and Joosse, 1984). CONCLUSIONS Inoculation of a solution of tritiated thymidine through the lung cavity of individuals of Biomphalaria tcnago- phila is an adequate method to label spermatozoa for determining their location in the partner after pairing. Labeled spermatozoa were detected in the recei\ing snail, through significant levels of scintillation counting, in the spennatheca, seminal vesicle, folhcles of ovotestis, nidamental gland, albumen gland, and oviduct-mucipa- rous gland. Material extracted from the collecting canal of the ovotestis did not show countings for labeled sper- matozoa. The presence of labeled spermatozoa simul- taneouslv in the follicles of the o\'otestis and in the o\i- duct-muciparous gland suggests options for tlifferential storage locations, in a process that includes displacement of autospermatozoa to allow for adequate positioning of allospermatozoa. ACKNOWLEDGMENTS We thank Dr. Mara Liicia Ferreira Dias for valuable sug- gestions and criticism to this work, and to fellowships from CAPES (Coordena9ao e Aperfei^oamento de Pes- soal de Nivel Superior, Brazil). LITERATURE CITED Abdel-Malek. E. T. 1954. Moqshological studies on the family Planorbidae (Mollusca: Pulmonata). II. Genital organs of Biomphalaria boissiji (Subfamily Planorbmae, H.A. Pils- bn, 1934). Transactions of American Microscopical Soci- et>' 7.3:285-296. Byfield, J. E. and O. H. Scherbaum. 1966. A rapid radioassay technique for cellular suspensions. Analytical Biochemis- try 17:433-443. Dias, M. L. F. 1995. Padroes comportamentais da reprodufao de Biomphalaria tcnngophila (Orbigny) (Mollusca, Pla- norbidae) ( Reproducti\e behavioural patterns of Biom- phalaria tenagophila (Orbigny) (Mollusca. Planorbidae).) University of Sao Paulo, Doctor of Science Thesis. Duncan, C. J. 1958. The anatomy and phvsioIog\- of the repro- ductive system of the freshwater snail Phtjsa fontinahs (L.). Proceedings of the Zoological Societ)' of London 131: .55-66. Duncan, C. J. 1960. The genital systems of the freshwater Ba- sonimatophora. Proceedings of the Zoological Socieh' of London 138:339-356. Duncan, C. ]. 1975. Reproduction. In: Fretter, \'. and ]. Peake, (eds.) Pulmonates. Academic Press, New York, pp. 309- 365. Fretter, V. and A. Graliam. 1964. Reproduction. In: Wilbur, K. M. and C. M. Yonge, (eds.) Physiology of Mollusca. Aca- demic Press, New York, pp. 127-164. Gazzinelli, G., F J. Ramalho Pinto and B. Gilbert. 1970. Up- take of '"'Fe as a tool for study of the crowding effect in Biomphalaria glabrata. The American Journal of Tropical Medicine and Hygiene 19:10.34-1037. ' Geraerts, VV, P. M. and J. Joosse. 1984. Freshwater snails (Ba- sommatophora). In: Tonipa, A. S., N. H \'erdonk and J. A. M. van den Biggelaar (eds). The Mollusca, vol. 7, Re- production. Academic Press, New York, pp. 142-208. Heneine, I. F, G. Z. GazzinelH and W. L. Tafuri. 1969. Iron metabolism in the snail Biomphalaria glahrata: uptake, storage and transfer Comparative Biochemistry and Phys- iology 28:391-399. Heneine, L. E., M. C. dos Santos and G. Z. Giizzinelli. 1970a. Transferencia de ferro em Biomphalaria glahrata. Anais da Academia brasileira de Ciencias 42 (suplemento):171- 175. Heneine, L E., F G. Ramallio-Pinto, J. Pellegrino and B. Gil- bert. 1970b. Uptake of "Fe as a tool for study of the crowding effect in Biomphalaria glahrata. The American Journal of Tropical Medicine and Hygiene 19:10.34-1037. Homan. J. A. 1972. The ultrastructure and activity of spenu in Acanthodoris pilosa. a nudibranchiate Mollusk. Marine Biolo,2\- 12:179-183. Horstmann, H. J. 1955. Untersuchungen zur Physiologic der Begattung und Befruchtung der Schlammschnecke Li/»i- naea stagnalis L. Zeitschrift fiir Morphologic und Okolo- gie der Tiere 44: 222-268. Jong-Brink, M, 1969. Histochemical and elecron microscope observations on the reproductive tract of Biomphalaria glahrata {Aiistrahvbi.s glabratus). Intermediate host of Schistosoma mansoni. Zeitschrift Zellforschungen 102: 507-542. Kitajima, E. W. and VV. L. Paraense. 1976. The nltrastnicture of mature spemis of the freshwater snail Biomphalaria glahrata (Mollusca, Gastropoda). Transactions of the American Microscopical Society 95:1-10. Kitajima, E. W. and W. L. Paraense. 1983. The ultrastructure W. Monteiro and T. Kavvano, 2()()() Page 79 of the spermatheca of Bioiiij>h(il/iria mplanata 22 L. radiata radiata 2 S. undulatus 1 P. cataracta 1 L. ochracca 1 15 E. complanata 31 Page 82 THE NAUTILUS, Vol. 114, No. 2 sun^eyed areas in the pond. Maximum depth was less than 1 m. Twenty-two specimens of E. coinplaiuita. 2 L. radiata radiata, 1 S. iiiuhdatus, 1 P. cataracta. 1 un- known juvenile mussel, and the only specimen of the target species, L. ochracca. from the entire survey, were found. The water on survey site 6 was cloudv; the bot- tom at this site was lightly covered with plants and sandy. Thirty-one specimens plus 1 unknov^ai juvenile were identified as E. complanata. A total of 213 live specimens comprising 6 different species were collected in 6 sites around the lake. AH specimens showed evidence of shell dissolution, mostly light wear (nacre exposed on 25% or less of the shell surface). Mean specimen length for Elliptio coinphmata, the only species collected in significant numbers for analysis, was 76.2 mm (n = 191), although this figure is most hkelv inflated due to bias in sampling methodology. At sample site 5, the only area where the target species was found, specimens of E. complanata were 11.6% larg- er than in the other sites. In addition, the greatest di- versity (5 species) was found at this site. DISCUSSION The density oi Lcptodea ochracca, is decreasing in Half- way Pond. This study confirms general observations of dechne over the past 15 years. The single specimen found in this survey is the only documentation of the existence of L. ochracca in Halfway Pond since 1981 when Doug G. Smith and A. E. Pratt, University of Mas- sachusetts Museum of Comparative Zoology, collected 12 specimens (UMAMZ 5.39). Agricultural nmoff from nearby cranberr}' bogs recently contributed to decreased water clarity accompanied by fish kills and potential de- crease in mussel densitv (Douglas G. Smith, personal communication). Lcptodea ochracca is typically found at low densities across its range in northeastern North America (Smidi, 1991; Strayer et al, 1994; Strayer and Smith, 1996; Strayer and Jirka, 1997). The single speci- men represents 0.47% of all specimens found (n = 213). Strayer ct al. (1994) found onlv 33 in their study of the Hudson River estuary in eastern New York and report the species constituting appro.ximately 5% of the unionid community of over one biUion animals. Repeated sam- phng of the same area in 1993-1995, subsequent to ze- bra mussel (Drcisscna polipiu>if)ha (Pallas, 1771)) inva- sion, resulted in only 16 specimens found. At the time of this studv (1995), zebra mussels had not reached Half- way Pond. The Lcptodea ochracca found in this stucK' was in the pond outflow in an area of high benthic macrophyte cov- er and sandy substrate that is typical habitat for this spe- cies (Johnson, 1947; 1970). Increased species diversih- in this area (5 of 6 total) and increased mean length of the most common species, Elliptio complanata, supports the prevision that the outflow area is a vital microhabitat for freshwater mussels in Halfway Pond. Increased den- sities of other freshwater mussel species (Nalepa and Gauvin, 1988) and filter-feeding insects (/Vllan, 1995; 142) have also been observed at other lake outflows. Fa- vorable conditions may include increased nutrient flow across mussel ctenidia induced by the increased stream flow, or protection from fish and mammal predators amidst the macrophvtic plants. Whatever the cause, the Agawam Ri\er outflow area is an important refugial microhabitat for the continued survival oi Lcptodea ochracca in Halfway Pond. The out- look for this species in the pond, however, is not good. With such a low density, reproductive potential may not be high enough to maintain a population. Surveys of sim- ilar areas in other coastal ponds should be conducted and appropriate protective measures taken if L. ochracca is to remain as a species of special concern or upgraded to endangered status in Massachusetts. LITERATURE CITED Allan, ]. D. 1995. Stream Ecolog\'. Stnicture and Function of Running Waters. Cliapman and Half London, 388 pp. Bereza, D. J. and S. L. H. Fuller. 1975. Notes on "LampsiUs" ochracca (Say) (Mollusca; BivaMa). Association of South- eastern Biologists Bulletin 22:42. Johnson, R. I. 1947. Lampsilis cariosn Say and Lampsilis ochracca Say. Occasional Papers on Mollusks 1:14.5-1.56. Johnson, R. I. 1970. The systematics and zoogeography of the Unionidae of the southern Atlantic slope region. Bulletin of the Museum of Comparative Zoologv 140:26.3—449. Livingston, T. (ed.) 1987. Massachusetts and Rhode Island Pre- serve Guide. Nature Conservancy Press, Boston, 56 pp. MDFW (Massachusetts Division of Fisheries and Wildlife). 1997. Massachusetts list of endangered, threatened, and special concern species. 321 CMR 10.60. Natural Heri- tage and Endangered Species Program, Massachusetts Di- vision of Fisheries and Wildlife, Westborough, Morrison, J. P. E, 1975. Maryland and Virginia Mussels of List- er. Bulletin of the American Malacological Union for 1974:.36-.39. Nalepa, T. F and J. M. Gauvin. 1988. Distriljution, abundance, and biomass of freshwater mussels (Bivalvia: Unionidae) in Lake St. Clair. Journal of Great Lakes Research 14(4): 411-419. Peckarsky, B. L., P. R. Fraissinet, M. A. Penton and D. J. Conk- lin. Jr 1990. Freshwater Macroinvertebrates of North- eastern North America. Cornell University Press, Ithaca, 442 pp. Smith, D. G. 1991. Keys to the Freshwater Macroinvertebrates of Massachusetts, v. 1.2. Universih' of Massachusetts Press, Amherst, Massachusetts, 236 pp. Smith, D. G. 1981. Selected freshwater invertebrates proposed for special concern status in Massachusetts. Massachusetts Department of Environmental Quality Engineering, Di- vision of Water Pollution Control, Westborough, 26 pp. Strayer, D. L., D. C. Hunter, L. C. Smith and C. K. Borg, 1994. Distribution, abundance, and roles of freshwater clams (Bivalvia: Unionidae) in tlie freshwater tidal Hudson River Freshwater Biology 31:239-248. Strayer, D. L. and K. J. Jirka. 1997. The Pearl Mussels of New York State. The New York State Education Department, New York, 113 pp. Straver, D. L. and L. C. Smith 1996. Relationship between zebra mussels (Dreissena polipnorpha ) and unionid clams J. Cordeiro, 2000 Page 83 (liiriiii; the earlv stages of tlie zebra mussel imasiiiii dlllic Hudson Rh'er. Fresliwater Biolog)' 36:771-779. Turgeon, D. D., J. F. Quinii |r., A. E. Bogan, E. \'. Coan, F. G. Hochberg, VV. G. Lyons, P. M. Mikkelsen, R. J. Neves, C. F E. Roper, G. Rosenberg, B. Roth, A. Sche'ltenia, F. G. Tlionipson, M. Vecchione and W'ilhanis, ]. D. 199S. Common and Scientific Names of Aquatic In\ertel)rates troni the United States and Canada: Molhi.sks. 2nd. Edi- tion. American Fisheries Society Special P\iblication, Be- thesda, 509 pp. Williams, J. D., M. L. Warren Jr, K. S. Cummings, J. L. Harris and R. J. Neves. 1993. Consen'ation status of freshwater nnissels of the United States and Canada. Fisheries 18(9): 6-22. In Memoriam Ruth D. Turner Rudolf Stohler 1914-2()()() 1901-2()()() THE NAUTILUS 114(2):84, 2000 Page 84 Notices An increase in the subscription rate of The Nautilus will take place effective volume 115 (2001). This increase represents an adjustment for the mounting costs of production, printing, and maiUng, which in turn reflect inflation rates in the USA integrated during the last 5 years. Our last increase in the subscription rate took place in March of 1995. 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FL 33957 THEt7NAUTILUS CONTENTS .^ '^^^ Vb/u/JK? 114, Number 3 °°'''''°"'-September5,2000 ISSN 0028-1344 Phylogeny of some gastropod mollusks derived from 18S rDNA swquences with emphasis on the Euthyneura 85 Size-depth patterns in two bathyal turrid gastropods: Benthoinangelia antonia (Dall) and Oenopota ovalis (Friele) 93 Four new genera for northeastern Pacific gastropods 99 ObscuraneUa papyrodcs, a new genus and species ot abyssal tonnoidean gastropod from Antarctica 103 Host-tree selection by Florida tree snails, Liguus fasciatus (Miiller, 1774), in Big Cypress National Preserve, Florida, USA 112 The development of three heterobranch mollusks from California, USA 117 Diminishing species richness of mollusks in Oneida Lake, New York State, USA 120 Sook Hee Yoon Won Kim Alisabet J. Clain Michael A. Rex James H. McLean Yuri L Kantor M.G. Harasewych Robert E. Bennetts Steven A. Sparks Deborah Jansen Rachel Collin Willard N. Harmann THE NAUTILUS 114(3):84-92, 2000 Page 85 Phylogeny of some gastropod mollusks derived from 18S rDNA sequences wdth emphasis on the Euthyneura Sook Hee Yoon Department ot Biological Sciences College of Natural Sciences Sung Kmui Kwan University Suwon 440-746, KOREA Won Kim' Department of Molecular Biology College ot Natural Sciences Seoul National Universitv' Seoul 151-742. KOREA' W()nkim@plaza.snu.ac.kr ABSTRACT The phvlogenetic relationships among gastropod subgroups, with emphasis on the Euthviieura, were investigated through the analyses of nearly complete ISS rDNA sequences of 29 representati\e gastropods. Neighbor-joining, maximum-likeli- hood, and maximum-parsimonN' methods were used ni the con- stniction of phvlogenetic trees. The 18S rDNA data support the monophyK' of Vetigastropoda, the \etigastropod clade Tro- choidea, and Caenogastropoda. However, the monophylies of two caenogastropod subgroups, Neotaenioglossa and Neogas- tropoda, are not supported. The basal position of Neritopsina is confirmed. Within the Euthyneura, the Stvlommatophora and the Systellommatophora are monophyletic, but the Opis- thobranchia, the Pulnionata, and the Basonnnatophora are not. The present study supports the inclusion of Succineidae within Stylommatophora. However the phylogenetic position of Sys- tellommatophora within Gastropoda remains unresolved. Additional key words: Mollusca, molecular phylogeny, Apo- gastropoda, Caenogastropoda, Opisthobranchia, Pulnionata, Vetigastropoda, Stylommatophora, Basonnnatophora, Systel- lommatophora, Archaeopulmonata. INTRODUCTION Of the molluscan classes. Gastropoda is the most diverse and the most ubiquitous group. It has successfully adapt- ed to most habitats, including marine, freshwater, and terrestrial emdronments. Many comparative studies were published based on morjDho-anatomical characters, including those of shell, pallial complex, and nervous, reproductive, and digestive systems (for recent reviews, see Haszpnmar, 198Sa; Bie- ler, 1992; Ponder and Lindberg, 1997). However, the status of knowledge of the phylogenetic relationships among and within the gastropod subgroups is still con- troversial (e.g., Golikov and Starobogatov, 1975; Gra- ham, 1985; Haszprunar, 1988a; Bieler, 1992; Ponder and Lindberg, 1996; 1997). This uncertainty is largely due to the lack of informative morphological characters com- ' Author for correspondence. men to the different taxa and the presence of the high level of phenotvpic diversity observed in the Gastropoda. In atkUtion to moiphological characters, molecular se- quences have proven to be very useful for in phyloge- netic reconstructions. 18S rDNA sequences are amongst the most informative molecular characters along a broad range of taxa within the Mollusca (e.g., Steiner and Miiller, 1996; Winnepenninckx ct al., 1996; Winnepen- ninckx ct a]., 1998a; Winnepenninckx et al., 1998b; Ada- mkewicz ct al., 1997; Bargues and Coma, 1997; Canapa ct al., 1999) and other animal phyla. Several studies were pubhshed on the molecular phylogeny of Gastropoda based on the sequence data of 28S rDNA (Rosenberg ct al, 1994; 1997; Tillier ct al.. 1994; Tillier et al, 1996) and 18S rDNA (Winnepenninckx ct al, 1996; Winne- penninckx ct al. 1998a; Harasewych et al, 1997a; b; 1998). Winnepenninckx ct al recently (1998a) investi- gated the phylogeny of gastropod groups below the class rank using the complete 18S rDNA sequences from 18 gastropod species. To further address gastropod phylogeny with empha- sis on Euthyneura (Opisthobranchia + Pulnionata), a group that has not been examined or discussed in detail from 18S rDNA data, we determined the complete 18S rDNA sequences for five representative gastropods. They include the first complete sequence data from Ce- phalaspidea (within Opisthobranchia) and Archaeopul- monata (within Pulnionata), and additional sequence data from Vetigastropoda and Stylommatophora (this lat- ter within Pulmonata). These sequences were analyzed in conjunction with previously published sequences of 24 other gastropotls. We focus on testing the monophyly of each of the euthvneuran subgroups, the Opisthobran- chia (e.g., Boettger,' 1955; Ghisehn, 1965; Gosfiner, 1981; 1985; 1991; Goshner and Ghisehn, 1984; Pouhcek et al, 1991; Ponder and Lindberg, 1997), the Basommatopho- ra (e.g., Tilher, 1984; Haszpnmar and Huber, 1990; Nordsieck, 1992), the Stylommatophora (e.g., Nord- sieck, 1992), and the Systellommatophora (Salvini-Plaw- en, 1980; Chnio, 1980; Tilher, 1984; Haszpnmar and Huber, 1990; Nordsieck, 1992). We also discuss the phy- Page 86 THE NAUTILUS, Vol. 114, No. 3 logenetic position of the Succineidae in the Stylomma- tophora (Rigby, 196.5; Solem, 1978; Tillier, 1989; Nord- sieck, 1992). In addition, we examine the monophylies of the Vetigastropoda (Salvini-Plawen, 1980; Salvini- Plawen and Haszprunar, 1987; Haszprunar, 1988a; b; Ponder and Lindberg, 1996) and the vetigastropod clade Trochoidea (Haszprunar, 1988a). MATERIALS AND METHODS Specimens An,\lyzed The 188 rDNA sequences of two vetigastropods (Nor- dotis disciis, 1858 base pairs, from Cheju Island and Ba- tilhis comutus, 1859 base pairs, from Mara Island), one opisthobranch {Bidlacta cxarata, 1849 base pairs, from Inchon), and two pulmonates {EUobium chinensis, 1845 base pairs, from Tamjin River, and Acusta dcspccta sie- boldiana, 1847 base pairs, from the Campus of Seoul National University). The material above was collected in Korea, and their sequences are described for the first time in the present study. The sequences of two neo- gastropods (Rapana venosa and Rcishia bronni), one pulmonate (Antliosiphonaria .siriti.s), and one chiton (Lcpidozona (Lcpidozona) corcanica) were reported in our previous study (Yoon et al, 1996) and the sequences of the remaining 21 other gastropods and 2 bivalves were obtained from GenBank. The nearly complete 18S rDNA sequences were an- alyzed for the 29 representative gastropods (one neri- toid, three vetigastropods, nine caenogastropods, two opisthobranchs, and 14 pulmonates). two bivalves, and one chiton species. Of these, the poK-placophoran Lcp- idozona {Lcpidozona) corcanica was used as an out- group, as the class Polyplacophora (included in the Acu- Ufera) is currently accepted as the stem group of the classes Gastropoda and Bivalvia (included in Conchi- fera), from studies based on morphological characters (see Salvini-Plawen, 1980; 1990; Haas, 1981; Runnegar and Pojeta, 1985; Brusca and Brusca, 1990; Ponder and Lindberg, 1996) as well as molecular results (Adamke- wicz et al, 1997; Bargues and Mas-Coma, 1997; Hara- sewych et al, 1997b). Table 1 Usts the studied taxa and GenBank accession numbers for the respective sequenc- es. The baseUne classification used in this work follows Haszprunar (1988a) for Streptoneura, Vaught (1989) for Opisthobranchia, and Boss (1982) for Pulmonata. DNA Extraction, PCR Amplification, and Sequencing Total nucleic acids were extracted from foot muscle of hve-collected and ethanol-preserved snails bv modifica- tions of standard procedure of Sambrook et al (1989). The 18S rDNAs were amphfied using the polymerase chain reaction (PCR) with two oUgonucleotide primers corresponding to conserved sequences proximal to 5' and 3' termini of metazoans (Nelles ct al, 1984: 1-19, 5'-CCTGGTTGATCCTGCCAG-3'; 1848-1868, 5'- TAATGATCCTTCCGCAGGTTA-3': the numbers cor- Table 1. Gastropod species used in the present study, with GenBank accession nvuiihers for sequences. NERITOPSINA NERITOIDEA Neritidae Nerita albicilla X91971 VETIGASTROPODA HALIOTOIDEA Haliotidae 'Nordotis disais AF082177 TROCHOIDEA Trochidae Monodoiita labia X94271 Turhinidae 'Btitillus comutus AF 16.5.3 11 CAENOGASTROPODA NEOTAENIOGLOSSA LITTORINOIDEA Littorinidae Littorina littorea X91970 Nodilittorina punctata Y11755 CALYPTRAEOIDEA Calvptraeidae Crepidula adunca X94277 TONNOIDEA Bursidae Bursa rana X94269 NEOGASTROPODA MURICOIDEA Muricidae Reishia bronni X98827 Rapana venosa X98826 Buccinidae Pisania striata X94272 Nassariidae Nassarius singuijorensis X94273 Fasciolariidae Fasciolaria lignaria X94275 EUTHYNEURA OPISTHOBRANCHIA CEPHALASPIDEA PHILINOIDEA Haniineidae 'BuUacta exarta AF 18867.5 APLYSIOMORPHA APLYSIOIDEA Aplysiidae Aphisia sp. X94268 PULMONATA ARCHAEOPULMONATA ELLOBIOIDEA Ellobiidae 'EUobium chinensis AF190452 BASOMMATOPHORA SIPHONARIOIDEA Siphonariidae Anthosiphonaria sirius X98S28 Siplionaria algesirae X91973 LYMNAEOIDEA Lviunaeidae Ltjmnaea glabra Z739S2 Bakerihpnnaea ctibensis Z83831 STYLOMMATOPHORA MEASURETHRA CLAUSILIOIDEA Clausihidae Balea biplicata X94278 HETERURETHRA Succineidae Oxijloma sp. X94276 Omalomjx matheroni AF047199 Athoracophoridae Athoracophonis bitenta- culatus AF047198 SIGMURETHRA ACHATINOIDEA Achatinidae Limicolaria kambcxd X66374 HELICOIDEA Bradvbaenidae 'Acusta despecta sieboldi- ana AF190453 Hehcidae Helix aspcrsa X91976 SYSTELLOMMATOPHO- RA ONCHIDIIOIDEA Onchidiidae Oncliidella celtica X70211 VERONICELLOIDEA X'eroniceUidae Laevicaulis alte X94270 Note: Classification follows Haszimniar (1988a) for Strepto- neura, Vaught (1989) for Opisthobranchia, and Boss (1982) for Pulmonata. " New sequences marked with asterisk. S. H. Yoon and W. Kim, 2()()() Page 87 respond to positions of liunian ISS rDNA). PCR ampli- fications were performed with Taij DNA pol\merase for 30 cycles (94° C for 1 min, 52° C for 2 min, and 72° C for 3 min). The ends of the amplified DNA fragments were modified for blunt-ended ligation using T4 kinase and T4 polymerase. The hlunt-endeil 18S rDNAs were hgated into pGEM-3zf(-) plasmid vector and trans- formed into DH5-a cell fines. Sequencing primers used in this study were reported in a previous paper (Moon ct «/., 1996). ISS rRNA-coding regions were completely sequenced in both directions with complete overlap. The DNA sequencing was performed by the dideoxynucleo- tide ch;iin-termination method (Sanger ct al., 1977) us- ing a Taq-Track kit (Promega Co.), accorcfing to the manufacturer's instructions. Eli'ctrophoresis of sequenc- ing reaction mixtures was periormed on butler-gradient 6% polyacrylamide gels and examined by autoradiogra- phy. Piivi,ck;enetic Analysis of 18S rDNA Sequences The sequences were initially aligned with the CLUSTAL W multiple-aUgnment program (Thompson ct al., 1994) and the alignment refined manualK'. A data-set of align- ment-stable positions was produced by excluding those positions that differed between alignments (Gatesy cf aZ., 1993). Analyses were limited to refiablv aligned regions, which included a total of 1754 nucleotide positions. Phv- logenetic reconstructions were performed using the neighbor-joining (NJ), maximum-likelihood (ML), and maximum-parsimony (MP) methods. PHYLIP version 3.572c (Felsenstein, 1995) was used for the neighbor- joining (Saitou and Nei, 1987) analyses. The distance analyses were done using Kimura (1980) and Jukes and Cantor (1969) matrices as input for the neighbor-joining analyses. Maximum-likelihood analvses were performed using the HKY (Hasegawa ct al. 1985) model in PAUP 4.0b2 (Swofford, 1999). For the quartet puzzling meth- od (the number of puzzling steps is 1000), empirical nu- cleotide frequencies, and transition/transversion ratio of 1.5 were estimated. Parsimony analvses were also per- formed using the computer program PAUP version 4.0b2 with closest stepwise addition options. The anal- yses employed a heuristic search using TBR branch swapping with random taxon addition. Branch length was optimized according to the ACCTRAN option. Bootstrap analyses (Felsenstein, 1985) of one hundred rephcates were performed to examine the confidence of nodes in NJ, ML, and MP analyses. RESULTS Figure lA shows the phylogenetic tree resulting from the neighbor-joining (NJ) analysis using the Kimura (1980) distances of an alignment of complete 18S rDNA sequences of 29 gastropod species. The polyplacophoran Lepidozona (Lcpidozoua) corcanica was used as out- group. The same tree topology was also obtained using Jukes and Cantor (1969) distances. The Neritoidea branches off first and the Vetigastropoda (Trochoidea -I- Haliotoidea) diverges next as an independent clade be- fore the clade Apogastropoda (Caenogastropoda + Eu- thvTieura). The monophyly of the Vetigastropoda and its subclade the Trochoidea (represented by Monodonta and BatiUus) is clearly shovvni in the tree, with ven' high boostrap support (100%: 100%). The Caenogastropoda shows a sister group relationship with the Euthyneura with very high bootstrap support (94%). Monophyly of the Caenogastropoda is supported (boostrap value = 100%), though neither the Neotaenioglossa (= Meso- gastropoda) nor the Neogastropoda emerged as mono- phyletic clades. There is strong bootstrap support (100%) for the monophvlv of Euthviieura (Opisthobranchia + Pulmon- ata), though the monophyhes of Opisthobranchia (Ce- phalaspidea + Anaspioidea) and Pulmonata are not sup- ported. The Basommatophora, consisting of the Siphon- arioidea and Lymnaeoidea, did not emerge as a mono- phyletic group. On the other hand, there is good support for the monoph)'fies of hvo additional groups in the Eu- thyneura, the Stylommatophora (boostrap value = 84%) and the Systellommatophora (boostrap value = 87%). Bootstrap values strongly support the position of the Succineidae (Oxi/loma and Omalonijx) within the sty- lommatophoran clade. However, the position of the Sys- tellommatophora within the Gastropoda was not posi- tively determined in the present study. The resulting tree from maximum-fikefihood (ML) analyses of the same data set is shown in figure IB. The ML tree confirms all the major results of the NJ tree (figure lA), with the exception that clades within Eu- thyneura lack significant bootstrap-support. The new branching order of Systellommatophora {Onchidclla and Laevicaulis) , Aplysiomoipha (Aplijsia), Archaeopulmon- ata (E//oi>((/)7()-Siphonarioidea {Siphonaria and Antho- siphonaiia), Cephalaspidea (BuUacta). and Stvlomma- tophora in the clade chffers from the order shouni in the NJ tree (figure lA), Aplysiomoipha-Svstellommatopho- ra-Siphonarioidea and Archaeopulmonata-Cephalaspi- dea-Stylommatophora. The maximum parsimony (MP) analyses produced a single tree with minimum length of 950 steps (figure IC). Generally speaking, MP analyses also yielded sim- ilar results except for minor differences in topologies among groups within the caenogastropod and the euthv- neuran clades. Maximum parsimony shows topological shifts within the caenogastropod clade, e.g., the new branching order of Bursa, Na.ssariiis, and Pisania-Fas- ciolaiia-Crcpidiila instead of the order shown in the NJ tree (figure lA), Crcpidida, Bursa, Pisania, Nassarius, and Fasciolaria. Within the Euthviieura, the MP tree differs from the NJ tree only in those branching points \vith low bootstrap values. There is no sister-group re- lationship among the Cephala.spidea [BuUacta). Aply- siomorpha {Aphjsia), Archaeopulmonata (Ellohium), Si- phonarioidea, Stvlommatophora, and Systellommatopho- ra. Ne.xt, we focused on the Euthvneura in separate. Page 88 THE NAUTILUS, Vol. 114, No. 3 KHlTl Omalonyx Athoracophorus Oxyloma r^Limicolaria ^ Helix Acusta Bullacta — Ellobium ZT Anthosiphonaria Siphonaria 'gj~ Onchidelta •evicaulis Aplysia 100 t—tymnea Bakerilymnea Fasciolaria Nassarius Pisania Bursa Crepidula Rapana ■j^Reishia {r^Litlorina — Nodilittorina I 100 Siylommatophora Pulmonata Cephalaspidea Opisthobranchia Archaeopulmonata J Basommatophora J Systellommalophora Aplysiomorpha J Basommatophora Neogastropoda J Neotaenioglossa J Neogastropoda J Neotaenioglossa Nerita [r~ Placopeclen Chlamys Lepidozona ^Balii ' fl Monodonta Batillus Nordotis Pulmonata Opisthobranchia Pulmonata Caenogastropoda Euth /neura Vetigasiropoda Nentopsina J Bivalvia Polyplacophora Apogastropoda B 11 I An ^' I — LimUolaria Helix Acusta Balea Omalonyx Athoracophorus Oxyloma Bullacta Antbosiphonaria Siphonaria Ellobium Aplysia Hj — Onchidella — iMevicaulis J Lymnaea — Bakerilymnea Fasciolaria Nassarius Pisania Bursa Crepidula U Rapana — Reishia Lj Littorina — Nodilittorina Monodonta Batillus Nordotis Nerita 91 Omalonyx Athoracophorus Oxyloma Limicolaria Helix Balea Acusta A nrhosiphonaria Siphonaria Onchidella iMevicaulis Bullae Ui Aplysia Ellobium Lymnaea Bakerilymnea Crepidula Fasciolaria Pisania Nassarius Bursa Rapana Reishia Littorina Nodilittorina Monodonta Batillus Nordotis Nerita Placopecten Chlamys Lepidozona Figure 1. A. Neighbor-joining tree detemiined by an alignment ol 29 nearly complete 18S rDNA sequence data for gastropods witli Lepidozona coreanica (Polvplacophora) as outgroup. Bootstrap percentages are shown abo\'e branches supported in at least 50% of 100 replicates. B. Strict consensus tree resulting from maximum-likelihood analyses of :29 nearly complete gastropod 18S rDNA sequences. Quartet puzzling method and HKY (Hasegawa et al, 1985) setting model were used. Lepidozona coreanica (Polyplacophora) was the outgroup. Bootstrap analysis was performed with 100 rephcates; values above 50% are indicated above the nodes. C. Strict consensus tree of maximum parsimony analyses based on the 333 informative sites of an alignment of 29 nearly complete gastropod 18S rDNA sequences (length = 950; CI = 0.6611; RI = 0.8418). Lepidozona coreanica (PoKplacophora) was die outgroup. Bootstrap values higher than 50% are indicated above the nodes. IIJU| LI Placopecten ^~ Chlamys Lepidozona — 1 c (.} 551 1 _ 85 1 6i 1 "1 1 100 1 85 1 54 74 57 97 <» "l 1 81, 100 100 1 100 1 1 mainly because the use of outgroups that are too far removed from the elade in studv mav give origin to ad- ditional homoplasies between ingroup and outgroup spe- cies. Figure 2A shows the results of NJ analyses of 16 nearly complete euthyneuran 18S rDNA sequences, with the caenogastropod Littorina littorca as outgroup. The resulting tree exhibits the same topology as the complete NJ tree (figure lA), with the exception that Ellohiitm (Archaeopulmonata) becomes the sister group to the clade Aplysiomoq^ha (Aphfsia) + Systellomma- tophora-Siphonarioidea, instead of clade Cephalaspidea (Bullacta) + Stvlommatophora, as present in the com- S. H. Yoon and W. Kim, 2000 Page 89 0.01 ~NodUittorina ~Littorina 3ol 84 88 pOma/on^ijr '"*' ' Alhoraco^horus Oxytoma ' Balea _ Limicolaria so" Helix ■Icuito Bullacta % [~Antbosiphonaria Siphonaria Oncbideila icaulis —Aplysia 'Ellobium ^PQ[ Lymnaea Bakerilymnea hrA I Ont/i taeii Styloninialophora Cephalaspidea Opisthobranchia Basomniatophora Systellommalophora Aplysiomorpha Opisthobranchia Archaeopulmonala - Basomniatophora Pulmonata Ncotacnioglossa Limicolaria Helix Acusta Balea Omalonyx Arhoracophorus Oxyloma Bullacta A nthosiphonaria Siphonaria Aplysia Onchidella Laevicaulis Ellobium Lymnea Bakerilymnea Nodilittorina Littorina " Omalonyx ' Athoracophorus ' Oxyloma " Limicolaria ' Helix ' Balea ~ Acusta ' Anthosiphonaria ~ Siphonaria ~ Lymnaea ~ Bakerilymnea ~ Onchidella ~ Laevicaulis ~ Aplysia ~ Ellobium - Bullacta ~ Nodilittorina - Littorina Figure 2. A. Euthviieuran neighbor-joining tree detennined by an alignment of 16 nearly complete euth>Tieuran 18S rDNA sequences, using Littorina littorea (Caenogastropoda: Littorinidae) as outgroup. Numbers at a node indicate bootstrap \alues higher than 50%. B. Euthyneuran maximum-likelihood tree detennined by an ahgnment of 16 nearly complete euthyneuran 18S rDNA sequences, using Littorina littorea (Caenogastropoda: Littorinidae) as outgroup. Quartet puzzling method and HKY (Hasegawapf ai. 1985) setting model were used. Bootstrap values above 50% are indicated above the nodes. C. Euthyneuran maximum parsimony tree calculated from the 149 infomiative sites of an alignment of 16 nearly complete euthyneuran 18S rDNA sequences, using Littorina littorea (Caenogastropoda: Littorinidae) as outgroup (length = 415; CI = 0.7494; Rl = 0.6750). Only bootstrap values higher than 50% are indicated. plete NJ tree. These two clades are not supported by bootstrap analysis of the euthyneuran clade. When the euthyneuran ML tree (figure 2B) is compared to the entire ML tree (figure IB), small topological shifts are discernible. The Siphonarioidea shows a sister group re- lationship with Aplysiomorpha (Aplysia) instead of with Archaeopulmonata {Ellobium), and this Siphonarioidea- Aplysiomorpha cluster appears as sister group of the Sys- tellommatophora instead of Cephalaspidea-Stylomma- tophora. However, bootstrap values that support these nodes are very low. Nevertheless, the euthyneuran ML tree strongly supported most of the major nodes found in the entire ML tree, with generally higher bootstrap values. Maximum parsimony analyses based on the 149 phylogenetically informative characters of the alignment of 16 euthyneuran species produced a single tree with minimum length of 415 steps (figure 2C). In the euthy- neuran MP tree, the first branching member is Cepha- laspidea (Bullacta) rather than Lymnaeoidea (Basom- matophora) found in the entire MP tree. The topologies Page 90 THE NAUTILUS, Vol. 114, No. 3 for the remaining taxa are generally identical to the ones in the tree containing all taxa. The euthyneuran trees resulting from NJ, ML, and MP analyses confirmed the topology of trees generated from the same types of anal- yses but based on all ta.xa. DISCUSSION For the phylogenetic relationships among and within the gastropod major subgroups, our study supports several aspects of the studv bv Winnepennickx ct al. (1998a). The Neritoidea diverged first, followed by the Vetigas- tropoda. Next, the clade Apogastropoda, comprising the Caenogastropoda and Euthyneura (each well supported as monophyletic groups), appears as a monophyletic group. However, the Neotaenioglossa (= Mesogastro- poda) and the Neogastropoda (included in Caenogastro- poda) and the Pulmonata (included in Euthyneura) do not appear as monophyletic groups. In addition, the po- sition of Systellommatophora within the Gastropoda could not be defined in the present study. In contrast to previous reports, our study provides more details on gastropod phylogeny, especially with re- gard to the phylogenetic status of the subgroups within the Euthyneura. All trees resulting from NJ, ML, and MP analyses refute the monophyly of the Opisthobran- chia. This result supports the claims of many opistho- branch workers, whom observed that parallelism and convergence ha\'e occured in most major organ systems within the opisthobranchs (Ghiselin, 1965; Gosliner, 19S1; 1985; 1991; Gosliner and Ghiselin, 1984; Pouhcek et al, 1991) and that the high degree of homoplasy in many characters in opisthobranchs contribute to diffi- culties in obtaining robust results from cladistic analyses (Goshner and Ghiselin, 1984; GosHner, 1985; 1991). Boettger (1955) maintained that the Opisthobranchia is paraphyletic. Ponder and Lindberg (1997) also suggest- ed that the Opisthobranchia is not monophyletic, a \dew- point in agreement with the results of TiUier ct al. (1994) from 28S rDNA data. It is not surprising, therefore, that many contradictory phylogenies and classification schemes have been suggested for the Opisthobranchia (e.g., Boettger, 19.55; Taylor and Sohl, 1962; Ghisefin, 1965). In relation to the phylogeny of the Pulmonata, we have mentioned that the Basommatophora {scmtt Ha- szprunar and Huber, 1990, Siphonarioidea -I- Lymnaeo- idea), is not monophyletic. Tillier (1984) considered that only Lyinnaeoidea belongs to Basommatophora and Si- phonarioidea to Archaeopulmonata. He suggested that pulmonates radiated into freshwater habitats as Basom- matophora and into marine habitats as Archaeopulmon- ata. Tillier et al. (1996), based on 28S rDNA data, showed that the Basommatophora is not monophyletic. However, Haszprunar and Huber (1990) suggested that both Siphonarioidea and Lymnaeoidea could be allocat- ed in Basommatophora due to the presence of common morphological characters such as a procerebrum com- prising only large cells, the lack of a contractile pneu- mostome, and the presence of an osphradium and pallial ciliary tracts. Nordsieck (1992) based on the presence of an anal opening shifted to the posterior mantle lobe, also considered the marine Thalassophila (Siphonarioidea) to be the sister group of the limnic Hygrophila (Lymnaeo- idea). Therefore, the common morphological characters found in the Basommatophoran, rather than represent- ing synapomoiphies, could all have been derived by con- vergence. Nordsieck (1992), based on morphological characters found in tentacles, kidney, central nervous system, and aspects of ontogeny, considered the SKlom- matophora to be a monophyletic group, which is con- cordance with the results derived from 28S rDNA se- quence data (Rosenberg et al, 1994; 1997; TilUer ct al, 1994; Tillier ct al, 1996). Although there is instability of branching pattern, the results for the euthvnieuran clade in the present study indicate that the Succineidae (rep- resented by Oxijloma and Omalomfx) belongs to the Sty- lommatophora (e.g., Solem, 1978; Tillier, 1989; Nord- sieck, 1992). These results, however, contradict the view of Rigby (1965) whom, on the basis ol moq^hological similarities of the digestive and reproductive systems, al- located the Succineidae in the Opisthobranchia. Syna- pomorphic characters such as the more or less reduced shell and mantle, and a visceral ganglion situated cen- trally or on the left side with respect to the axis of the central nervous system (e.g., Salvini-Plawen, 1980; Nordsieck, 1992) support the monophvlv of the Systel- lommatophora (represented by Onchidudae and Veron- icellidae). However, this monophyly has been questioned by several authors (e.g., Chmo, 1980; Tilfier, 1984; Ha- szprunar and Huber, 1990). Climo (1980) in particular considered the systellommatophorans as a polyphyletic assemblage diverging at the base of the euthvmeurans. Tillier (1984) studied the morpho-anatomical characters of the pallial complex, digestive tract, reproductive, and central nervous systems, and divided the Pulmonata into only three orders; Archaeopulmonata, Basommatophora, and Stylommatophora. He included the Systellommato- phora in the Archaeopulmonata and suggested that the Onchidudae (within Systellommatophora) is more close- ly related to the Ellobiidae (within Archaeopulmonata) than to the VeroniceUidae (within Systellonunatophora). However, our present result supports the monophyly of Systellommatophora. Since the erection of the Vetigastropoda by Salvini- Plawen (1980), the presence of .synapomorphic charac- ters such as ctenidial sense organs, the epipodial sense organs, and the special structure of the esophagus have generally supported the monophyly of this clade (Sal\'ini- Plawen and Haszprunar, 1987; Haszpnmar, 1988a; b; Ponder and Lindberg, 1996; 1997). Other vetigastropod features include the dominant presence of the right dor- soventral retractor muscle, the right excretory organ, and bilamellate ctenidia with skeletal rods. Previous molec- ular data using partial 18S rDNA (Harasewvch ct al, 1997a; b) and' the 28S rDNA (Tillier ct a/.,' 1994) se- quences also supported the monophyly of the Vetigas- tropoda, which is also supported in the present study. S. H. Yoon and W. Kim. 2000 Page 91 Moiioplnlv of the Trochoidea (represented by Mono- chmta and Batillus) is also confirmed herein. The Tro- choidea is defined hv SNiiapomoqihies such as loss of the right ctenuhum in relation to the loss of the shell slit (Haszpnniar, IBSSa; h) and the nionophvK' ol the gronp is also in concordance \\'ith the stud)- based on 28S rDNA sequences by TilUer et al. (1994). In conchision, the ISS rDNA data stronglv support the monoph\l\ of the following higher gastropod clades: Vetigastropoda, Trochoidea (within Vetigastropoda), Apogastropoda, and the two included clades Caenogas- tropotla and Euth\meura. Within the euthyneuran clade, both the Stslommatophora and the Systellommatophora are monophyletic. However, our 18S rDNA data failed to support monophyly of the Neotaenioglossa and the Neogastropoda (within Caenogastropoda), Opisthobran- chia, Pulmonata, and the puhnoTiate Basommatophora. These non-monophvletic subgroups, therefore, at pre- sent can be considered as grades rather than clades. The basal position of Neritopsina is confirmed in this study. In addition, the Succineidae is included in Stylomma- tophora. Still, the Svstellommatophoran position within the Gastropoda, that is, its immediate relationship to ei- ther Opisthobranchia or Pulmonata, or to any other group for that matter, could not be defined. The insta- bility of topology and short branch lengths within the Caenogastropoda and the Euthyneura may be due to the fact that the mollusks, including gastropods, apparently rachated in an "explosive" fashion during a relatively short period of time. Most extant major groups of mol- lusks appeared around a relatively short time at the Pre- cambrianyCambrian boundary (e.g., Runnegar and Po- jeta, 19S5; Winnepenninckx et al, 1996; Adamkewicz ct al. 1997; Harasewych ct al, 1997a). Futtire studies at- tempting to define phylogenetic relationships at these levels may take into consideration other molecules such as cytochrome c oxidase I and/or 16S rDNA. Such mol- ecules evolve more rapidly than 18S rDNA, and seem more likelv to contain information needed to solve phy- logenetic relationships within these clades. ACKNOWLEDGMENTS This work was supported bv a grant from KOSEF (95- 0401-04-01-3) for years 1995-1998. We thank Dr. B. L. Choe, Mr. J. L. Lee (Sung Kyun Kwan University, Ko- rea), Dr. J. S. Lee (Kang Won University, Korea), and Mr D. G. Min (Shell House, Seoul, Korea) for providing specimens. We are also indebted to Dr. C. B. Kim (Yale University, USA) and Dr. J. K. Park (University of Mich- igan, USA) for providing many references. We thank Ms. D. W. Jung (Seoul National University) for proofreading the manuscript. Special thanks are due to Dr. S. Y. Moon (University of Koln, Germany) for her assistance in soft- ware analyses. We are especially grateful to Dr. S. Y Moon and to Dr. M. G. Harasewych (Smithsonian In- stitution, Washington, USA) for their comments and suggestions. LITERATURE CITED Adamkewicz, S. L, M. G. Harasewvcli, J. A. Bkike, D. Saudek, and C. ]. Bull. 1997. A molecular plnlogeiiy of tlie !)i\alve mollusks. Molecular Biology and Exolution 14:619-(i29. Bargues, M. D. and S. Mas-Coma. 1997. Phylogenetic analysis of limnaeid snails ba.sed on ISS rDNA sequences. Molec- ular Biology and Evolution 14:.569-.577. Bieler, R. 1992. Gastropod phylogeny and systematics. Aiuuial Review of Ecologv' and Systematics 23:.311-.3.3S. Boettger, C. R. 19.5.5. Die Systematik der euthyneineii Schnecken. 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Investigation of molluscan phvlogenv on tlie basis of 18S rRNA Sequences. Molecular Biologv' and Evolution 13: 1,306-1317. Winnepenninckx, B., G. Steiner, T Backeljau, and R. De Wachter 1998a. Details of gastropod phvlogenv inferred from 18S rRNA sequences. Molecular Phvlogenetics and Evolution 9:.55-63, Winnepenninckx, B., D. G. Reid, and T Backeljau. 1998b. Per- formance of 18S rRNA in littorinid phvlogenv (Gastro- poda: Caenogastropoda). Journal of Molecular Evolution 47:586-596. " Yoon, S. H., S. Y. Moon, B. L. Choe, and W. Kim 1996. Se- quence divergence ot 18S ribosomal DNA of gastropods (molluscs). Korean Journal of Malacolgy 12:85-90. THE NAUTILUS 114(3):93-98, 2000 Page 93 Size-depth patterns in two bathyal turrid gastropods: Benthomangelia antonid (Dall) and Oenopota ovalis (Friele) Alisabet J. Clain Department of Biologv' UiiiversiU' of Massachusetts, Boston 100 Morrissev Boulexard Boston, MA 02125 USA Michael A. Rex' Department of Biology Uni\ersit\' of Massachusetts, Boston 100 Morrissev Boulexard Boston, MA 02125 USA nnchael.rexCffumb.edu ABSTRACT During the past decade, there has been a resurgence of interest in the evolutionars' and ecological significance of animal body size. Geographic patterns of body size in deep-sea organisms remain poorly described. In this paper, we analyze depth-re- lated trends of larxiil and adult shell size in two turrids (Gas- tropoda) from the western North Atlantic. The upper bathyal Oenopota ovalis (478-2022 m), which has non-planktotrophic de\elopment, shows significant positixe size-depth clines for laryal. but not adult shells. The lower bathyal Benthomangelia antonia (2359-3834 ni), with planktotrophic development, shows no trend for lan'al shells, but a positive relationship of size to depth for adult shells. The increase in size with depth confirms earher obser\'ations of size-depth clines in deep-sea snails, which may reflect selection for greater competitive abil- it\' and foraging efficiency with increased depth. Contrasts in clina! \ariation between lar\al and adult shells suggest that dif- ferent stages of life history in deep-sea snails may respond in- dependently to environmental gradients. Additional key words: Gastropoda, Turridae, deep sea, size- depth patterns, western North Atlantic. INTRODUCTION Animal body size has been related to a wide range of physiological (Peters, 1983; Chown and Gaston, 1999), evolutionary (Stanley, 1979), and ecological phenomena (LaBarbera, 1989). There is also considerable new in- terest in the imphcations of body size for geographic range (Brown, 1995), species diversity (Finlay et al., 1996; Nee and Lawton, 1996), abundance (Siemann et al, 1996) and conservation of biodiversity (May, 1988; Marquet ct al, 1990; Gaston and Blackbum, 1996). One of the most striking characteristics ot the deep-sea fauna, including moUusks, is the small size of most organisms. Explanations for this have centered primarily on mea- suring size-depth trends in various deep-sea t;L\a or func- tional groups, and relating these to food availability, which decreases with depth (Thiel, 1975, 1979; Gage ' Author for correspondence. and Tyler, 1991). Size-depth relationships appear to vary considerably among faunal components (Rex and Etter, 1998). It is unclear the degree to which this inconsisten- cy represents biologically meaningful differences in how organisms respond to environmental gradients, or meth- odological differences associated with how size is mea- sured and the way that morphologically and taxonomi- cally heterogeneous groups of species have been com- bined to obtain composite estimates of size. To study size as an adaptation, it is important to standardize mea- surements to common growth stages and to examine pat- terns within species (Gould, 1969; Rex and Etter, 1998). Rex (1979) and Rex and Etter (1990, 1998) showed that size, standardized to specific larval and adult growth stages within individual deep-sea snail species, shows a strong tendency to increase with depth. These studies were based on average interpopulation size differences using relatively few sampling sites, often located near the extremes of the species' depth ranges. Here, we present detailed analy,ses of size-depth trends in populations of two species of deep-sea snails, using larger sample sizes and much more thorough depth co\'erage. Results gen- erally support the positive size-depth trends reported earlier, but also suggest that larxae and adults may re- spond differently to selective differences associated with change in depth. MATERIALS AND METHODS This study focuses on two common and taxonomically well-known bathyal turrids, Benthomangelia antonia (Dall, 1881) and Oenopota ovalis (Friele, 1877), showni in figures 1^. They were collected from the western North Atlantic, south of New England (table 1, figure 5), using epibenthic sleds (Hessler and Sanders, 1967), box cores (Hessler and Jumars, 1974), Blake deep-sea trawls or beam trawls (Tanner, 1897). The two species were chosen primarily because they were abundant enough in existing collections to allow reasonable statis- tical analyses across broad depth ranges. B antonia and O. ovalis differ in their mode of de- Paee 94 THE NAUTILUS, Vol. 114, No. 3 Figure 1. SEM of the adult shell of Oenopota ovalis (Station 87, 1102 m, 4.01 mm total length). Figure 2. SEM of the adult shell of Bcnthoinanoelia antonia (Station 76, 2862 m, 6.51 mm total length). Figure 3. SEM of the larval shell of Oenopota ovalis (Station 87, 1102m, 0.45mm protoconch height). The indentation before the first readily detectable adult rib (at a magnification of .30x) was used as the lanal- adult transition (see text for further discussion). Figure 4. SEM of the larval shell oi BenthomangeUa antonia (Station 76, 2862 m, 1.18 nun protoconch height). Horizontal arrow indicates the lar\al-adult transition (see text for further discus- sion). velopnient (Bouchet and Waren, 1980). O. ovalis ap- pears to haw direct or leeithotrophic development in which larvae mature in a protected egg capsule and ei- ther emerge crawhng or undergo a brief pelagic or de- mersal dispersal phase (e.g., Thorson 1935, 1944; Bouchet and Waren, 1980; Shimek, 1983). In B. antonia. larvae hatch at a small size and develop planktotrophi- cally. Veligers evidently undergo ontogenic vertical mi- gration in the water column, and consequentlv have more dispersal potential than do the laivae of O. ovalis (Killinglev and Bex, 1985). An increa.se in the proportion of species that have planktotrophic development with increased depth seems to be a general pattern in deep- sea turrids of the western and eastern North Atlantic (Potter and Bex, 1992). Mollusks are useful for this tvpe of study because their calcareous shells record the hfe history of each inchvid- ual, and are not subject to the changes in form that are caused by preservation in soft-bodied organisms. We 40° - 38° 36 207 73 N13. ^ W3 .••62 N4_ N10 _r--> •^084 2041* 2569 <^ »2037 ^/ "^ / 340 '*\ -.^" L-1-^ 2714 ,95 - - '-'2038 74° 72° 70° 68° 66° 64° Figure 5. Map of Northwest Atlantic showing the localities of collection stations. See table 1 for station data. Depth con- tours are in meters. Diamonds represent stations where Oen- opota ovalis was collected; circles represent sites for Benthom- anoelia antonia. chose only specimens that were collected hve. Measure- ments of size were standarchzed to the transition be- tween laival and adult shells (figures 3 and 4). If this was undetectable due to erosion or breaking, the spec- imen was not used. In B. antonia, the transition is clearly marked by changes in bodi color and sculpture (Bouchet and Wart^n, 1980; Bex and Etter, 1990). In O. ovalis, both larvae and adults are heaviK- calcified ami are the same white color making the transition less chstinct. The protoconch is smooth and the adult shell has fine vertical ribs. We used the indentation before the first readily detectable adult rib as the lar\'al-adult transition for this species. Shells were measured microscopically, orienting them by the method described in Bex and Etter (1990). Each shell was mounted on a disk of clav and then placed on a 4-diniensional stage of an Olvmpus dissecting micro- scope equipped with a Sony 3 CCD Color Video Camera mount. The camera was attached to a NuVista Imaging Board in a Macintosh Centris 650 computer. Using NuVista Capture+ Software, the images of the mounted shells were digitized. The saved images were then load- ed into the NIH Image program. Previously digitized images of a 1.0 mm stage micrometer with 0.01 mm gradations were used to calibrate and measure the shell images. Size-depth trends in the two species were analv^zed using three variables: standardized lar\al, and adult size, and estimates of final size attained. EarK' protoconch whorls for one species, B. antonia. were often corroded, making it impossible to accurately measure protoconch height in all specimens. Consequentlv, larval size was taken as protoconch width. Standardized adult size, hereafter referred to as Whorl- 1 Size, was measured as A. J. Clain and M. A. Rex, 2000 Pa^e 95 Table 1. Station data for samples of O. ovalis and B antonia measured in this stud\. Tlie species collected, sampling nietliodolog)-, and number of specimens used in this study are also given. For sampling method: ES = epibenthic sled (Hessler and Sanders, 1967), BC = box core (Hessler and Jumars, 1974), BT = beam trawl, DSf = Blake deep-sea trawl (Tanner, 1897). ACSR indicates Atlantic Continental Slope and Rise Study (Maciolek et ai, 1986). i:).-|)tli Saiiiplins: .Number of Species Station Cruise 1 m ' Latitude °.\' Longitude ° W method indi\iduals Oenopota ovalis 88 Ch.mn 478 39°54.10' 70°37.00' ES 3 96 Cn.\iN 498 39°55.20' 70°39.50' ES 3 N4 ACSR 550 40°21.17' 67°.32.18' BC 1 207 Chain 808 .39°51.15' 70°55.35' ES 6 87 Chain 1102 .39°48.70' 70°40.80' ES 29 NIO ACSR 1220 39°48.10' 70°05.30' BC 2 N13 ACSR 1250 39°48..35' 70°54.94' BC 1 N13 ACSR 1250 39°48.35' 70°54.94' BC 1 7.3 Atlantis 1400 39°46.50' 70°43.30' ES 15 103 Chain 2022 39°43.60' 70°37.40' ES 8 BenthomangeUa 2084 Albatross •7.3,59 40°16.S3' 67°05.25' BT 4 antonia 62 Atlantis 2496 39°26.00' 70°33.00' ES 1 2096 Alb.atross 2654 39°22.33' 70°52.33' BT 3 2221 Alb.^tross 2789 39°05.50' 70°44.50' BT 9 76 Chain 2862 39°38..30' 67°57.80' ES 23 72 Atlantis 2864 38°16.00' 71°47.00' ES 12 64 Atlantis 2886 38°46.00' 70°06.00' ES 10 2174 .\lb.atross 2915 38°15.00' 72°03.00' BT 3 2041 Albatross 2941 39°22.83' 6S°25.00' DST 5 2716 Alb.atross 2983 38°29.50' 70°57.00' BT 14 2037 Alb.^tross 3166 38°53.00' 69°23.50' BT 3 2569 Albatross 3259 39°26.00' 68°03.50' BT 2 .340 Knorr 3310 38°16.00' 70°21.55' ES 13 2570 Alb.\tross 3316 39°54.00' 67°05.50' BT 4 2714 Alb.\tross 3338 38°22.00' 70°17.50' BT 4 2038 Albatross 3718 38°30.50' 69°08.42' DST 3 95 Atlantis 3753 38°33.00' 68°.32.00' ES 2 126 Atlantis 3806 39°.37.25' 66°45.50' ES 18 "*"* Chain 3806 38°00.70' 69°16.00' ES 7 78 Chain 3828 . 38°00.80' 69°18.70' ES 8 85 Chain 38.34 37°59.20' 69°26,20' ES 8 the height plus width of the first post-lar\al whorl (Gould,l969; Re.\ and Etter. 1990). Final size attained by adults, hereafter referred to as Adult Size, was mea- sured as the maximum height plus width of the post- larval shell. Adult Size can not be standardized to a com- mon de\elopment stage, because the species appear to have indeterminate growth. The distribution of Adult Size in a population reflects the recent history of re- cruitment and growth more than size as an adaptive property. To estimate the final size attained, we selected the largest two indi\iduals (from anv station) found in 250 m depth increments. These three measures of size (Protoconch Width, Whorl-l Size, and Adult Size at- tained) were regressed against depth to anal\-ze bath\- metric cUnal patterns. RESULTS AND DISCUSSION Relationships of size \'ariables to depth are showTi in fig- ure 6. Regression fines are fitted onl\' for significant re- lationships (see table 2 for regression equations and their statistics). All significant regressions are positi\e, indi- cating an increase in size with depth. There is no indi- cation that size decreases significantly with depth for an\' variable in either species. In O. ovalis, lanal size in- creases with depth, but post-lar\'al stages show no pat- tern. In contrast, B. antonia shows no relationship of lar\-al size to depth, but both Whorl- 1 Size and .\dult Size do increase with depth. These results largeK" confinu those reported bv Rex and Etter (1998) for B. antonia and O. ovalis. The pre- sent study is more statisticallv accurate in that it includes larger sample sizes, broader depth co\erage. and a much more continuous distribution of samples across the depth gradient. For Whorl-l Size, both studies detected a strong positi\e increase with depth in B antonia and no apparent trend in O. ovalis. For protoconch size. Rex and Etter (I99S) reported a weaklv significant (p<0.051 increase with depth in B. antonia. and no relationship in O. ovalis. \\ith more extensi\e sampling, the proto- conch size-depth relationship becomes just insignificant (p = 0.05S9) in B antonia. and strongly significant in Page 96 THE NAUTILUS, Vol. 114. No. 3 1.50 1500 2000 2500 3000 3500 4000 Depth (m) A. J. Clain and M. A. Rex. 2(K)() Page 97 Tabic 2. Regression siuiiiiuin tor rclatioiisliips ol sr/.i- to dt-ptli in fiilure 6. For eacli regression, tlie niiiMt)er of iiuli\i(luals, regression einiations, F-\'alues. r- \alues, ant! tlie sigiiitieanee are gi\en. Number of Significance Measurement Species specimens Regression equation r* F-\alue (p) Protoconcli widtli B. antonia 156 Y = 0.982 + 2.69E - 5x 0.230 3.622 00589 Protoconch width O. ovalis 69 Y = 0.6.39 + 4..54E - 5x 0.147 11.507 0.0012 Whorl 1 size B. antonia 135 Y = 2.219 + 1.78E - 4x 0.106 15.850 0.0001 Whorl 1 size 0. ovalis 62 Y = 2.016 + 7.44E - 5x 0.046 2.863 0.0958 Adult size B. antonia 156 Y = 7.644 + l.OOE - .3x 0.007 1.116 0.2925 Adult size 0. ovalis 69 Y = 5.171 - 2.S8E - .5x 0.000 0.010 0.9221 Largest adult B. antonia 14 Y = 1.694 + 4.00E - 3x 0.332 5.963 0.0310 Largest ailult O. ovalis 11 Y = 5.915 + 1.2.3E - 5x 0.000 0.001 0.9772 O. ovalis. Rex and Etter (1998) did not e.stiniate final adult size. As with studies of biodiversity (Rex et al, 1997) and life-history characteristics (Stuart and Rex, 1994), the patterns in protoconch size that are revealed with better sampling demonstrate the importance of us- ing large databases, and broad continuous depth gradi- ents to establish biogeographic patterns in the deep sea. This is especiallv relevant for species like B. antonia that show high variabihtv in shell architecture (figure 6). It is interesting that O. ovalis, with non-planktotrophic development shows a size-depth cline only for larvae; whereas B. antonia with plankiotrophic development ex- hibits no trend for larvae, but a significant positive cline for Whorl- 1 Size that also persists at final size attained. For B. antonia, an e.xplanation might be that larvae show no pattern because they migrate from numerous distant sites that may exert quite different selective regimes, and that selection for size at settlement along the depth gra- dients is not strong enough to overcome the mitigating effects of dispersal. In this context, it is interesting that larvae (and adults) of B. antonia show considerably more variation in size than those of O. ovalis, possibly reflect- ing multiple geographic origins for the former (figure 6). Selection for increased size with depth may be imposed largely during subsequent growth resulting in the posi- tive size-depth cfine seen in adult shells of B. antonia. The nature of the relevant depth-related selective gra- dient remains unk-nown. Rex and Etter (1998) suggested that the decrease in nutrient input with depth may select for larger size because of its metabolic and competitive advantages. The opposite pattern observed in O. ovalis is difficult to interpret in a consistent way. Larger size at hatching may confer advantages of lower vulnerabifity to preda- tors, greater locomotion to forage, a greater range of food, and more abilitv to withstand star\'ation (Spight, 1976). However, the absence of a size cfine at adult stag- es is puzzUng. In general, the results do indicate that different life stages can respond differently and inde- pendently to environmental changes associated with depth. Since the species live at different depths, a geo- graphically controlled comparison is not possible. Also, because we are analyzing just two species, the results may represent idiosyncratic effects that are phenotypic plastic responses or phvlogenetic constraints, rather than general patterns that are related to selective gradients and developmental modes. In summary, the upper bathyal O. ovalis shows a sig- nificant positive size-depth cfine for larvae, but not adults. The lower bathval B. antonia reveals no depth- related pattern for lai-val shells, but a significant increase in size with depth for adults. The exact causes of these opposing trends are necessarily speculative; but, the re- sults do suggest that larvae and adults may respond dif- ferently to selective gradients in species with contrasting modes of development in the deep sea as they appear to do in coastal moUusks (Pechenik, 1999; Pechenik ct al, 1996). The increase in size with depth generallv sup- ports earlier studies on geographic variation in deep-sea gastropods (Rex and Etter, 1990, 1998; Rexet al., 1999). However, detailed analyses of individual species high- light the need for large databases and good sampUng coverage throughout species' bathvmetric ranges to ac- curately assess biogeographic patterns in deep-sea or- ganisms. ACKNOWLEDGMENTS We thank Ron Etter Carol Stuart, John Ebersole, and Rob Stevenson for reacfing drafts of the manuscript. Bill Fowle for the SEMs and Jack Cook for the map of the Northwestern Atlantic Ocean. Howard Sanders, Fred Grassle, Nancy Maciolek, James Blake and Jerrv' Hara- sewych helped provide the original material that was Figure 6. Relationships of Protoconch Width (top), Whorl-1 Size (middle), and Adult Size (bottom) to depth in populations of Oenopota ovalis (diamonds) and Benthoinan^i'lia antonia (circles) collected in the deep sea of the western North Atlantic. The solid symbols in the Adult Size graph indicate the largest two individuals found in each 250 m interval. Lines indicate significant regressions. See table 2 for regression statistics. Page 98 THE NAUTILUS, Vol. 114, No, 3 measured and analv^zed here. This research was sup- ported by the National Science Foundation Grant OCE- 9301687 to MAR, and by the University of Massachu- setts. LITERATURE CITED Bouchet, P. and A. Waren. 1980. Re\ision of the north-east Atlantic bathyal and abyssal Turridae (Mollusca, Gastro- poda). 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Meddelelser om Gr0nland 100:1-71. Thorson, G. 1944. The zoology of East Greenland. Marine Gastropoda Prosobranchiata. Meddelelser om Gronland 121:1-181. THE NAUTILUS 114(3):99-1()2, 2()()() Page 99 Four new genera for northeastern Pacific gastropods James H. McLean Natural Histors' Museum of Los Angeles Counh' 900 Exposition BKd. Los Angeles. CA 90007 USA jniclean@nhni.org ABSTRACT Four new genera for nortlieasteni Pacific Gastropoda are pro- posed; V'elutinidae: Torcllivclutina. type species "Torellia" am- monia Dall, 1919; Euliniidae; Subniso. type species "Chemnit- zia" ranai de Folin. 1867; Turridae. Clathurellinae; Retidrillia. tspe species "Suavodrillia" willctti Dall. 1919; Mangeliinae; PcrimangeUa. t\pe species "Mangelia" intcrfossa Carpenter, 1864. Additional kci/ words: lutinidae. Gastropoda, Euliniidae, Turridae, V'e- INTRODUCTION This is the third in a series of papers in which new gen- era for northeastern Pacific gastropods are introduced (see McLean, 1995a, b). I had prexaously indicated (Mc- Lean, 1995a) that a checklist of the northeastern Pacific gastropods was in preparation; however, as indicated in McLean (1996: 2), I am preparing a more complete work rather than a checklist. Taxa described here are to be used in an illustrated rexasion of all shelled gastropods of the northeastern Pacific, ranging from Arctic Alaska and the Aleutian Islands to central I3aja Cahfomia, Mex- ico. Description of these genera in advance of the book allows for a more detailed treatment than will be pos- sible in the larger effort. Further papers in this series will describe genera in which the tyjie species or other included species are new. To distinguish original combinations for type species of the new genera I am here using the convention of placing all citations of original genera within quotation marks. Illustrations are proxaded here for type species of each new genus. Although monotvpic genera are not encouraged in phylogenetic classifications, I do not refrain from intro- ducing a few such genera where necessarv-, allowing that additional living or fossil species may yet be discovered that would render those genera no longer monotypic. Museum acronvms are: LACM, Natural History Mu- seum of Los Angeles Counb,'; USNM, National Museum of Natural History, Washington. SYSTEM ATICS Family Velutinidae Gray, 1840 Genus Torcllivclutina new genus Type species: "Torellia" ammonia Dall, 1919 (Figure 1). The shriveled liolotvpe from offshore depths at the Aleutian Islands, Alaska, was illustrated by Waren (1989: 16, fig. llf). A more recently collected, preserved spec- imen from the Aleutian Islands is illustrated here. Included species: The genus is monotypic. Diagnosis: Shell large (to 30 mm diameter), apical whorl depressed, profile nearly planispiral, of two rapidly inflated whorls, calcareous layer lacking in large speci- mens except for that which borders the apertural hp; shell consisting of chitinous periostracum after attaining 3 mm diameter; protoconch with spiral sculpture (ac- cording to original description of Dall); suture deep, um- bilicus broad; mature surface composed of thick, chitin- ous, colabral a.xial lamellae. Remarks: Waren (1989: 16) examined the type spe- cies for purposes of comparison with his then described new genus and species Pscudotorcllia fragilis from Ice- land. He illustrated the radula of "Torellia" ammonia (op. cit., fig. IIC), which he found to agree with that of other velutinid radulae, and reported that the operculum and other trichotropine features are lacking. Although Waren removed "T." ammonia from the trichotropid ge- nus Torellia, and transferred it to the Velutinidae, he did not assign it to a genus. Torellivehitina is therefore here proposed for "T" ammonia. It differs from Pseudotorcl- lia in not having a sohd calcified shell with spiral sculp- ture and not having the large, projecting protoconch of Pseudotorellia. Because the calcareous layer is lacking, fresh speci- mens of Torellivehitina ammonia should be retiiined in fluid preservative and should not be dried, which results in the shrinkage and buckling of the thick periostracal layer. Etymology: The name is a compound of the tricho- tropid genus Torrellia and the velutinid genus Vclutirm, Page 100 THE NAUTILUS, Vol. 114, No. 3 A Figures 1^. T)pe species of new genera. 1. Torellivehttina ammonia (Dall, 1919), apertural and spire views. LACM 152281, Kanaga Pass, between Kanaga Island and Tanaga Island, Andreanof Islands, Aleutian Islands, Alaska, 61 ni. Maximum diameter .30 mm. 2. Subniso rangi (de Folin, 1867). LACM 1972-38.7, Punta Penca, N of Bahia Potrero, Guanacaste Pro\., Costa Rica, 10 ni. Length 3.2 mm. 3. Retidrillia willetti (Dall, 1919). USNM 216409, lectotype. Forrester Island, southeasteam Alaska, dredged. Length 11.5 mm. 4. Perimangelia inferfossa (Carpenter, 1864). LACM 1959-13.42, Granite Creek, Monterey County, California, 10 m. Length 8.5 mm. to emphasize that it represents a velutiiiid with the su- perficial aspect of the trichotropid genus Torrellia. Family Euhmidae Philippi, 1853 Genus Subniso new genus Type species: "Chemnitzia" rangi de Fohn, 1867 (Figure 2). Type locality: Perlas Islands, Panama. Waren (1992: 183) could not locate type material, but the spe- cies was well figured bv de Folin and subsequently by Waren (1992, figs. 17, 20-22, 25-27, 30, 31), so there is little doubt as to its identity. The species occurs from southern Baja Cahfomia, Mexico, to Ecuador. Included species: Two species, the type species and "Niso" hipolitcnsis Bartsch, 1917, for which the holotype from Punta San Hipohto, Baja California, Mexico, was illustrated by Emerson (1965, fig. 9) and Waren (1992: figs. 19, 23, 24). Both species have previously been placed in Niso Risso, 1826, bv Emerson (1965) and War- en (1992). Diagnosis: Shell small (length to 4 mm), slender (length to breadth ratio 2.5-2.9), non-umbilicate, basal angulation pronounced, coloration brown, without color pattern. Remarks: The two species of Subniso are unUke other species of Niso in size, profile, and color. The two spe- cies are small, non-umbilicate, and have a basal angula- tion (strongly projecting in Subniso rangi) and a uniform brown coloration. Waren (1992: 185) remarked: "Both Niso hipolitcnsis and N. rangi are unusual among the species of Niso in their small size, 3^ mm shell height, whereas most species of the genus have a shell that is 10-30 mm high, occasionally even higher." Additionally, I point out that Niso species are usually broadly umbil- icate, except for the large N. atiilloi (Hertz and Hertz, 1982), usually not angulate at the base, and have color patterns that may be banded or variegated. Taken to- gether, these differences are sufficient to justify' generic recognition of Subniso. Waren (1992: 1S3) noted that a starfish host is known for one Indo-Pacific species of Niso, but nothing is known of the host echinoderm for the two species of Subniso. Etymology: The generic name combines the generic name Niso, with the prefix sub-, to emphasize the small size, compared to Niso. Family Turridae Swainson, 1840 Subfamily Clathurelhnae H. and A. Adams, 1858 Genus Rctidrillia new genus Type species: ' Suavodrillia" willetti Dall, 1919 (Fig- ure 3). Type locality: Forrester Island, Southeast Alaska. Until now the species has been unfigured (it was not illustrated by Kosuge, 1972). The here selected lectotype (Figure 7) is the largest (11.3 mm) of seven original syn- type specimens in the type lot. The radular tooth of the type species was figured by McLean (1971, fig. 114) (from LACM 66-66, Graham Island, Queen Charlotte Islands, British Columbia). A specimen from the Rae Baxter collection in the LACM extends the distribution to Akutan, Aleutian Islands (LACM 83-345. 263 m depth). J. H. McLean, 2000 Page 101 Included species: At least three species, the t\pe spe- cies and two species from the northeastern Atlantic dis- cussed and illustrated bv Bouchet and Waren (1980: 32) in the genus Drilliola Locard, 1897: "D. " pntina (Wat- son, 1881), and "D." megalacmc (Sykes, 1906). Diagnosis: Shell profile with concave shoulder and projecting peripheral carination below which base is rounded and has strong spiral sculpture. Anal sinus on concave shoulder, deep and broad. Axial sculpture ex- pressed as nodes, strongest at peripheral carination, more f;iintlv expressed on strong spiral cords of base. Peripheral carination of early whorls at midvvhorl. Pro- toconch paucispiral, with early development of periph- eral carination. Radular tooth long, haqioon-like, with broad base. Remarks: The type species was described in the ge- nus SiiavodriUia Dall, 1918 (tvpe species: "Drillia" kcn- nicotti Dall, 1871), with which it shares the long har- poon-like radular tooth. Dall (1921: 69) indicated doubt that "S." willetti was congeneric by placing a question mark preceding the genus. Differences are that Reti- drillia iLillctti and the tvvo additional species here as- signed to the genus are half the size of S. kcnnicotti, and have sculpture that is axial and spiral, rather than strictly spiral as in SiiavodriUia. The newly restricted Siiavod- riUia is monot\pic for S. kcnnicotti, a species broadly distributed in the north Pacific from Hokkaido, Japan, the Kurile and Aleutian Islands, and the Gulf of Alaska to Southeastern Alaska. Bouchet and Waren (1980: 32) retained "a wide va- riety of species" in Drilliola, including those that 'look rather different but have a similar radula and opercu- lum." They elected "to keep them in Drilliola rather than placing them in any of the perhaps more similar, but anatomically unkiiown genera hsted by Powell (1966) in different subfamihes. ' Species tvpical of Dril- liola are more slender than those of Rctidrillia and do not ha\e the broad, excavated shoulder. The proposal of Rctidrillia thus provides a genus with a type species hav- ing knowii radular characters for the two species cited by Bouchet and Waren (1980). On shell characters, Retidrillia resembles species as- signed to Plicisifrinx Sysoev and Kantor, 1986, in which the radular tooth is of the modified wishbone t)pe, in- dicative of the less derived turrid subfamily Cochlespi- 30-4). Synonym: "Man^elia" interlirata Stearns, 1872. The species occurs from Clallnm County, Washington, to Isla San Ceroninio, Baja Cahfomia, Mexico, based on specimens in the LACM collection. Included species: Two species, the type species and the less familiar "Manficlia" nitcns Carpenter, 1864 (syn- type figured by Palmer, 1958, pi. 28, fig. 1). It ranges from Sonoma County, California, to Ensenada, Baja Cal- ifornia. Diagnosis: Shell relatively small and slender with dominant axial sculpture, crossed by narrow spiral cords of lesser strength. Protoconch of 1.5 whorls, strongly projecting, smooth at first, developing fine spiral sculp- ture after first half whorl, followed by weaker axials (ax- ials more numerous than that of mature sculpture), changing imperceptibly to adult sculpture. Remarks: Earlier (McLean, 1978: 74), I placed the type species of the new genus ("Mangelia" intcrfossa) along with "Daphnclla" ftiscoligata Dall, 1871 in Clatli- romangclia Monterosato, 1884. I now consider Clath- romangclia (type species "Plcurotoma" grantiin Philippi, 1844; see Powell, 1966: 106) to be appropriate for two species in southern Cahfomia: "Daphnclla" fuscoligata and "Mangilia iClafhmmangclia)" rhi/ssa Dall, 1919. These two species have coarse clathrate sculpture and relatively low, paucispiral protoconchs. Pcrimangclia differs from the two species of Clath- romangclia in having numerous, narrow spiral cords overriding the dominant axial ribs, rather than having coarsely clathrate sculpture with nodes at intersections. Protoconchs differ: the paucispiral protoconch of the two species of Pcrimangclia is strongly projecting, whereas the paucispiral protoconch of the two species of Clathromangclia is much lower. Etymology: The name combines the prefix pcri- (near) with Mangelia Risso, 1826, one of die oldest gen- era in the subfamily Mangeliinae. ACKNOWLEDGMENTS Photographic prints were made by Michael Eraser. I am grateful to Lindsey Groves, Daniel Geiger, Anders War- en, and a further anonymous reviewer for offering help- ful suggestions. Etymology: The name is a compound of rcti-, mean- ing net, with reference to the axial and spiral sculpture, and Drillia Gray, 1838, one of the earliest named of tur- rid genera. Subfamily Mangehinae Fischer, 1883 Genus Pcrimangclia new genus Type species: "Mangelia" intcrfossa Caqienter, 1864 (Figure 4). Syiitypes were figured by Palmer (1958, pi. 27,' figs. 5, 6)' Also figured by McLean (1969; 1978, fig. LITERATURE CITED Bouchet, P. and A. Waren. 1980. Revision of the northeast Atlantic bath\al and abyssal Turridae (Mollvisca, Gastro- poda). Joiinial of Moiluscan Studies, Supplement 8:1-119. Dall, W. H. 1919. Descriptions of new species of mollusks of tlie family Turritidae from the west coast of America and adjacent regions. Proceedings of the United States Na- tional Museum, 56(2288): 1-86, pis. 1-24. Dall, VV. H. 1921. Summary of the marine shellbearing mol- lusks of the northwest coast of America, from San Diego, Califoniia, to the Polar Sea, mostly contained in the col- lection of the United States National Museum, with illus- Page 102 THE NAUTILUS, Vol. 114, No. 3 tratdons of hitherto unfigiired species. United States Na- tional Museum, Bulletin 112, 217 pp., 22 pis. Emerson, W. K. 1965. The eastern Pacific species oiNiso (Mol- lusca: Gastropoda). American Museum Novitates 2218:1- 12. Kosuge, S. 1972. Illustrations of type specimens of molluscs described bv William Healev Dall (Northwestern Pacific gastropods). Special publication of the National Science Museum, Tokyo, 29 plates and unpaged captions. McLean, J. H. 1969. Marine shells of Southern California. Los Angeles County Museum of Natural History, Science Se- ries, no. 11, 104 pp. McLean, J. H. 1971. A revised classification of the family Tur- ridae, with the proposal of new subfamilies, genera, and subgenera from the Eastern Pacific. The V'efiger 14:114- 130. McLean, J. H. 1978. Marine shells of .southern California, Re- vised edition. Natural Histors' Museum of Los Angeles County, Science Series, no. 24, 104 pp. McLean, J. H. 1995a. Four new genera for northeastern Pacific prosobranch gastropods. The Nautilus 108:39-41. McLean, J. H. 1995b. Three additional new genera and two replacement names for northeastern Pacific prosobranch gastropods. The Nautilus 108:80-83. McLean, J. H. 1996. The Prosobranchia. In: P H. Scott, J. A. Blake, and A. L. Lissner (eds.) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and western Santa Barbara Channel. Volume 9. The MoUusca Part 2 — The Gastropoda. Santa Barbara Museum of Natural History, Santa Barbara, v -t- 160 pp. Palmer, K. V. W. 1958. Type specimens of marine Mollusca described by P. P. Carpenter from the west coast (San Diego to British Columbia). The Geological Society of America, Memoir 76, vi -I- .376 pp., 35 pis. Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and .Museum, no. 5, 184 pp., 23 pis. Sysoev, A. V. and Y. 1. Kantor. 1986. [New and rare abyssal species of the tamilv Turridae (Gastropoda, Toxoglossa) in the northern part of the Pacific Ocean.] Zoologicheskii Zhumal, 65(10): 1457-1469. [In Russian] Waren, A. 1989. New and little known Mollusca from Iceland. Sarsia 74:1-28. Waren, A. 1992. Comments on and descriptions of eulimid gastropods from Tropical West America. The Veliger 35: 177-194. THE NAUTILUS 114(3):103-111, 2000 Page 103 Ohscuranella papijrodes, a new genus and species of abyssal tonnoidean gastropod from Antarctica Yuri I. Kantor A. N. Se\ertzov Institute of Problems of E\oliition Russian Academy of Sciences Leninski Prospect 33 Moscow 117071, RUSSIA kantor@malaco-se\in.msk.ni M.G. Harasevvych Department of In\ertel)rate Zoology National Museum of Natural History Smitlisonian Institution Washington, DC. 20560-0118, USA HanisevvTchS'nmnh. si.edu ABSTRACT The new genus Ohscuranella and O. papijrodes. its type spe- cies, are described from the abyssal plain off the Ross Sea, Antarctica. Obsntranella is included in the primarily tropical, shallow water superfamiK Tonnoidea because it has a taenio- glossan radula, extensible proboscis, large sali\'arv glands com- posed of anterior and posterior lobes, salixary ducts that pass through die nerve ring, and an undifferentiated oesophageal gland. This taxon is precluded from the families Laubierinidae, Pisanianuridae, and Tounidae because it lacks a monopectinate osphradium, paired proboscis retractor muscles passing through the nerve ring, buccal glands, and rachidian teeth with lateral basal denticles. Ohscuranella is assigned to tlie family Ranellidae primarily on the basis of shell, radular, and oper- cular morphology. This is the first report of the Tonnoidea in Antarctic waters, and the first record of Ranellidae from abyssal depths. Additional key words: Caenogastropoda, Ranellidae, anatomy, systemafics. INTRODUCTION In the course of examining Antarctic Buccinoidea sam- pled under the auspices of the United States Antarctic Program (USAP) and housed in the National Museum of Natural Histoiy, Smithsonian In,stitution, we encoun- tered five lots of gastropods with large bucciniform shells that had tentatively been attributed to the buccinoidean genus Bathijdomus Thiele, 1912, by Dell (1990:198- 199). Dell identified four of these lots as Bathiidomus obicctus Thiele, 1912, but considered the filth lot, con- sisting of a single specimen, to represent an undescribed species of Bathijdomus. Dissections of preserved mate- rial clearly demonstrate that these gastropods are not referable to Buccinoidea, nor even to Neogastropoda, but rather represent an unnamed genus and species of the superfamily Tonnoidea. This is the first report of this superfamily in the Antarctic nialacofaimu, UTid one of very few records of the Tonnoidea from the abyssal zone. The family Ranelfidae, to which this genus is assigned. has not prcNitjusly been reported from die Antarctic or from abyssal depths. In this paper, we provide descriptions of this new ge- nus and new species, and infer its taxonomic position within the Tonnoidea by comparing its anatomy, radula and operculum with published anatomical reports (e.g. Weber, 1927; Houbrick and Fretter, 1969; Day, 1969; Beu, 1981; Hughes and Hughes, 1981). Waren and Bouchet (1990), Riedel (1995). and Beu (1998) each provide useful, if not entirely congruent, syntheses of tonnoidean families, and include anatomical descriptions as well as numerous illustrations of radulae, opercula and protoconchs. Abbreviations used in the descriptions of morphometric characters are explained in Table 1. SYSTEMATICS Class Gastropoda CuNaer, 1797 Superfamily Tonnoidea Suter, 1913 Family Ranelhdae Gray, 1854 Genus OhsciirancUa new genus Type species: Ohscuranella papijrodes new species Diagnosis: Protoconch unknown. Teleoconch large, pyriform, thin, with large aperture, single, rounded, un- thickened terminal varix adjacent to thin, flared outer lip in adult specimens. Shell sculpture limited to fine, wide- ly spaced, spiral cords. Operculum very small, shaqily tapering anteriorly, with anterior terminal nucleus. Pro- boscis wall extremely thick, paired proboscis retractor muscles absent. Rachidian teeth lacking lateral basal denticles. Salivary glands large, differentiated. Osphra- dium bipectinate. Etymology: Ohscunis (L.) — obscure, unclear + Ra- nclla — genus of Ranellidae. Ohscuranella papijrodes new species (Figures 1-36, Table 1) Bathijdomus obtectus Thiele, 1912— Dell, 1990:198-199, figs. 299-300. Page 104 THE NAUTILUS, Vol. 114, No. 3 Table 1. Obscuranella papifrodes. new species. Measurements of shell characters. Linear measurements in mm. USNM USNM USNM Character Holotvpe Paratype 1 Paratype 3 901317 870610 870610 Shell LensTth (SL) 63 + 58 + 33+ 57.0 43.6+ 33.2+ Last Whorl Len.^th (LWL) 55.2 51.8 28.5 47.3 38.5 30.4 Aperture Length (AL) 45.1 43.3 23.0 38.4 31.5 25.5 Shell Width (SW) 40.8 41.5 19+ 34.1 24+ 20.2+ Number of spiral cords on last whorl 14 12 14 5 14 13 Number of spiral cords on penultimate whorl 6 7 7 2 5 3 Bathjdomns sp.— Dell, 1990:199. Description: Shell large (exceeding 63 mm), very thin, fragile, ovate-pvrifonin. Protoconch and upper whorls missing in ;ill t\pe material. Preserved portions of teleo- conch of 2V2 rapidly expanding, evenly rounded whorls. Shoulder rounded, indistinct. Suture adpressed, shallow. Axial sculpture limited to fine, straight, weakly prosocline growth hues. Adult specimens with a single, weak, hollow vailx adjacent to thin, flared outer Up (figures 2, 7, arrow). Spiral sculpture of sharp, narrow, evenly spaced cords (14 on last whorl, 6 on penultimate whorl), with much weaker sinuous threads (22-.30) of varying width between adja- cent cords. Aperture large [—0.7 shell length (SL)], broadly ovate, deflected from shell axis by 9-11°. Outer lip tliin, evenly rounded in upper part and concave at transition to siphonal canal, weakly reflected. Inner hp consisting of long, convex, tnedially indented parietal re- gion and short, smootli, a.xial columeOa with strong, long siphonal fold that crosses cofling axis of shell. Siphonal canal short, broad, weakly recurved dorsally. Callus of thin, wliite, porcellaneous glaze overlying parietal region, adapical portion of broad, nearly axial siphonal fasciole. Shell color pale ohve-tan, confined to outermost shell lay- er. Aperture white. Periostracum very thin, yeflowish brown, with densely spaced a.xial lamellae, occasional short hairs at intersection of lamellae with spiral cords. Operculum (figures 4, 5) very small (0.16 AL), x'estigial, dark yellow, subtriangular, with straight sides, terminal nucleus. Dorsal surface with numerous, closely spaced growth lines. Ventral surface with thin, glazed lateral mar- gins. Operculum attached over most oi its surface. Shell ultrastructure (Figure 23): Shell thin (101 |xm), composed of three layers. Outermost laver (figure 23, ca) thinnest (4 |xni), composed of columnar crystals. Middle layer (figure 23, ccl) thickest (79 jjim), composed of collabrally oriented cross-lameUar crystals. Inner layer (figure 23, rcl) thin (IS p-m), composed of cross-lamellar crystals oriented perpendicular to growing edge of the shell. Anatomy (Paratype 1, 9): Soft tissues comprising appro.ximately 3V2 whorls. Mantle cavity spans just under V2 whorl, nephridium (figures 24, 25, 27, n) about % whorl, digestive gland (figures 24, 25, dg) 2V^ whorls. Mantle edge (figures 24, 25, 27, me) thickened, smooth, completely covers head. Columellar muscle (figure 25, cm) thick, broad, spanning sUghtly more than one whorl, attached to shell at rear of nephridium. Foot short in contracted state (LengthAVidth == 1.0), with conspicu- ous propodium. Body color uniform reddish-tan, without pattern in alcohol preserved specimens. Head (figure 26) very large, as wide as foot, with broad, blunt, tapering tentacles (figure 26, tn) with black eyes at their bases. Operculum about 4 mm long (0.09 AL), otherwise sim- flar to that of holotvpe. Paratype 1 (and all other pre- served specimens) with proboscis protruded through very wide rhynchostome (figure 26). Mantle cavity (Figure 27): Mantle cavity as deep as broad (—Vi whorl). Siphon (figures 24, 25, 27, s) broad, muscular, very short, extending shghtly beyond mantle edge (figures 24-27, me). Osphrachum (figures 24—27, os) situated along central half of ctenidiuni, bipectinate, nearly symmetrical, sUghtly narrower on left side than right. Ctenidium (figures 24-27, ct) long, spanning near- ly entire mantle length, formed of tall triangular lamel- lae, nearly twice as high as broad. Hvpobranchial gland (figure 27, hg) poorly developed, lacking distinct folds. Rectum (figure 27, re) runs along inner surface of palfial oviduct, narrow, terminating in simple anus (figure 27, a) behind thickened mantle edge. Alimentary system (Figures 15, 19-22, 24, 32-35): Everted proboscis (figures 24-26, pr) —30 mm long (0.7 AL), unpigmented, with folded walls indicating potential for further extension. Proboscis wall very thick, —60% Figures 1-14. Obscuranella papyrodes new species. 1. Apertural, 2. right lateral, and 3. dorsal views of the holotype, USNM 898683. 4. External and 5. internal views of the operculum of the holotype (internal view coated with Ammonium chloride to emphasize sculpture). 6. Apertural and 7. dorsal views of paratype 1, USNM 896131. 8. Apertural and 9. dorsal views of the larger shell, 10. dorsal view of smaller shell, USNM 870610. 11. Apertural and 12. dorsal views of paratype 3, USNM 886105. 13. Apertural view and 14. apical view of early whorls of USNM 901317. Scale bar = 1 cm for all shells; Scale bar = 2 mm for operculum; Scale bar = 5 mm for protoconch. Y. I. Kantor and M. G. Hamsewvch, 2000 Page 105 Page 106 THE NAUTILUS, Vol. 114, No. 3 Y. I. Kantor and M. G. Harasew\'ch, 2000 Page 107 of proboscis radius, composed of 3 layers of muscles. Innermost la\er of circular nuiscles (figure 32, cm), mid- dle layer tliickest (2/3 of proboscis wall), of longitudinal muscles (figure 32, Im), outer layer of circular muscle. Buccal mass (figure 33, bm) small, attached to proboscis walls b\ numerous, thin tensor muscles (figures 15, 32, 33, tm), as is the anterior oesophagus (figures 15, 32, aoe). Retractor muscles passing through ner\'e ring and joining buccal mass and columellar muscle absent. Mouth (figure 32, mo) a narrow, vertical slit. Buccal tube (figure 33, bt) short, leading to cuticle-lined buccal cavity with ventral p;ur of semicircular jaws (figure 33, j). Jaws (figures 16-18) dark brown, pappilate along outer edge (figure 17). Inner surface of jaw composed of small, closeK- spaced platelets that produce "coiibled" surface distally (figure 18), smooth pro.ximally (figure 16). Odon- tophore (figure 15, od) small, oval, lining bottom of the buccal cavity. Walls of buccal cavity very thick. Proboscis nerves (figure 32, pn) paired, very thick, nuining from cerebro-pIeur;il gangfia along proboscis length, inner\'at- ing buccal mass and anterior part of proboscis. Anterior esophagus divided into dorsal and ventral channels by prominent longituchnal folds (figures 15, 34, 35, If) that extend from the buccal cavitv to the posterior edge of esophageal gland. The right fold overlaps the left (figure 34). Radular ribbon (figure 19) short (5.8 mm, 0.13 AL), nearly twice as long as cartilages, narrow (~ 580 |xm, 0.013 AL), consisting of 45 rows of teeth, posteriormost 4 rows nascent. Rachidian tooth (figure 21, rt) with large, broad median cusp, flanked by 5-9 denticles per side. Base broad, strongly concave posteriorly, lacking cusps along tooth base. Lateral teeth (figures 21, 22, It) narrow, with long, thin, cusp flanked by 3—4 denticles on inner edge, 4—6 denticles on outer edge. Two long, recurved, distally flattened marginal teeth (figures 20; 22, mt) per side, outer tooth longer than inner Inner distal edges serrated with 2-7 cusps. Salivarv' glands large, irregularly shaped, completely covering the esophageal gland (fig- ure 32, oeg). Right salivary gland more elongated, slight- ly larger than left. Each gland consists of two lobes. Pos- terior lobe (figure 32, plsg) massive, composed of curved radially oriented bfind tubules. Anterior lobe (figure 32, alsg) smaller, acinous, ventral. Salivary ducts (figures 32- 34, sd) thick, extending from posterior lobes, becoming attached to oesophagus wafls before passing through nerve ring. We were not able to identify connections between the salivary ducts and the anterior lobes of the salivary glands, as reported for Cymafium intermedium (Pease, 1869) by Andrews ct al. (1999). Salivary glands attached to oesophagus by thin muscular and connective tissue fibers and innervated by several nei^ve branches. Esophageal gland formed of deeply glandular dorso-ven- tral folds, small relative to salivary glands, of the same color as surrounding tissues and esophagus. Stomach large, U-shaped, similar to that of Ci/mathiin nicolxiri- cum (Houbrick and Fretter, 1969) in external moqihol- ogy, too poorly preserved to reveal details of internal morphology, except that the ducts of the digestive gland are paired, closely spaced, and situated in the middle region of the stomach. Posterior duct twice the diameter of anterior duct. After leaving stomach intestine curves dorsally, passing under then along posterior edge of ne- phridium towards rear of mantle cavity (figures 24, 25, m). Female reproductive system (Figures 25, 27-30): The only specimen dissected (Paratype 1) was a mature female. Pallial oviduct consists of small albumen gland (figure 25, ag) along outer wall of nephridium, large cap- sule gland (figures 25, 27, eg) fining right wall of pallial cavity. Lumen of capsule gland small anteriorly (figure 29), broad posteriorly (figure 28), at juncture to albumen gland. No seminal receptacles were identified, possibly due to poor fixation. Bursa copulatrix (figures 27, 29, 30, be) large, long, spanning more than half length of cap- sule gland. Pallial oxiduct opens (figures 27, 29, go) by long, narrow slit at midlength. Male reproductive system (Figure 31): The repro- ductive system of a male specimen lacking shell (para- type 2) was examined. Testes occupy upper half of whorls of visceral mass above posterior border of stom- ach, giving rise to seminal duct. Seminal duct forms large, convoluted seminal vesicle before entering mantle cavity. Duct descends to floor of mantle cavity at mid- length, opening to form groove with thickened, obvious- ly glandular walls leading to penis base. Penis (figure 31. p) broad, dorsoventrally flattened, with seminal groove (figure 31, sgr) nmning along inner edge, around the bluntly rounded distal edge, halfvvay down the outer edge, terminating in a verv small, inconspicuous papilla (figure 31, pap). Type locality: Victoria Land, Balleny Islands, Sturge Island, RA' Eltanin. .st. 1949, 66252'S, 164a32'E, \n 2507-2525 m, 5 February 1967. Type material: Holotype, USNM 898683 (figures 1- 5, shell and operculum only, soft parts not present), from the type locahty; paratype 1, USNM 896131 (figures 6- Figures 15-23. Obsairanella papijrodes new species. Parat\pe 1, USNM 8961.31. 15. Distal end of the proboscis opened dorsally. 16-18. Left jaw. 16. Entire inner surface of jaw. 17. Enlarged section of the upper edge. 18. Enlarged section of the surface in the middle part of the jaw. 19-22. Radiila. 19. View spanning width of radular ribbon. 20. Left marginal teeth. 21. Rachidian anil lateral teeth. 22. Right lateral and marginal teeth, tilted to .30°. 23. Ultrastnicture of the shell (Paratvpe 3. USNM 886105). aoe, anterior oesophagus; If, lateral folds; od, odontophore; ca. columnar shell layer; ccl, comarginal crossed lamellar shell layer; j, jaw; It, lateral teeth; mt, marginal teeth; r, radula; rcl, radial crossed lamellar shell layer; rt, racliidian tooth; tm, tensor muscles. Page 108 THE NAUTILUS, Vol. 114, No. 3 ov 1 cm Figures 24-31. Ohscuranella papijrodes new species. Paratope 1, USNM 896131. 24. Neutral and 25. dorsal news of animal removed from the shell. 26. Antero-dorsal \iew of the head. 27. Mantle complex, opened from left side and reflected. 28-30. Schematic sections through the pallial gonoduct. 31. Penis (Parat)'pe 2). a, anus; ag, albumen gland; be, bursa copulatrix; eg. capsule gland; cm, columellar muscle; cnie, cut mantle edge; ct. ctenidium; dg, digestive gland; go, genital opening; hg, hypobranchial gland; in, intestine; me, mantle edge; n, nephridiuni; no, nephrichal opening; op, operculum; os, osphradium; ov, ovary; p, penis; pap, papilla; pr, proboscis; re, rectum; s, siphon; se, siphonal edge; sgr, seminal groove; st, stomach; tn, cephalic tentacle. Y. I. Kantor and M. G. Harasewvch, 2()()() Page 109 5 mm Figures 32-35. Obscitranella papijrodes new species. Anterior alimentary system, Paratype 1, USNM 896131 32. Proboscis (opened ventrallv) and organs of cephalic haeniocoel. 33. Anterior part of the proboscis, opened ventrally. 34. Transverse section through anterior oesophagus, viewed from anterior 35. Esophageal gland, opened dorsally. Dashed line indicates \entral midline. alsg, anterior lobe of the salivary gland; ao, anterior aorta; aoe, anterior oesophagus; bm, buccal mass; bt, buccal tube; cm, circular muscle layer of the proboscis wall; j, jaw; If, longitudinal fold of anterior oesophagus; !m, longitudinal nuiscle layer of the proboscis wall; mo, mouth opening; nr, nerve ring; ode, odontophore caitilages; oeg, oesophageal gland; plsg, posterior lobe of the salivary gland; pn, proboscis nerve; poe, posterior oesophagus; pw, proboscis wall; rs, radiilar sack; sd, salivary duct; tm, tensor muscles. 7, 9 pecimen, anatomical de.scription.s based on diis specimen), paratype 2, USNM 896139 {6 specimen, shell not present, male reproductive system based on this specimen), both irom south of Hjort Seamount, RA' Eltanin, stn. 1964, 59°5H'S. ISS^arW. in 29S5-2992 m, 10 February 1967; paratype 3, USNM 886105 (fig- ures 11-12, sex undetermined) Victoria Land, south of Scott Island Bank, IW Eltanin. stn. 1939, 69203'S, I7904IE, in 3519-3.596 m, 1 February 1967. Other material examined: USNM 901317 (Figs 13- 14, 1 specimen, sex undetermined) Victoria Land, Moubray Bay, RA^ Eltanin. stn. 2002, 72218'S, 177e:35'E, in 2005-2010 m, 11 January 1968; USNM 870610 (fig- ures 8-10, 2 dead shells) Antarctic Peninsula, IW £/- tanin, stn. 1003, 62^41S, 54243W, in 210-220 m, 15 March 1964. Etymology: papijrodes — made from papyrus, refer- ring to the thinness of the shell. Remarks: The type series oi ObsciirancUa papijrodes, n. sp., consists of lour specimens, incluchng one paratvpe that lacks a shell. All were collected hving on the abyssal Page 110 THE NAUTILUS, Vol. 114, No. 3 Figure 36. Geographic and bath\nietric distribution of Obsairnnelln papijrodes new species. plain off the Ross Sea. We are provisionally attributing three additional specimens to this species. One live-col- lected specimen (figure 13; USNM 901317), also from the abyssal plain off the Ross Sea, was considered by Dell (1990:199) to be congeneric but not conspecific with Obsctiranella papijrodes n. sp. (which he identified as Bathi/domiis ohtcctus Thiele, 1912) because of its more elongated shell, longer siphon;il canal, and angular shoulder. Even the earliest teleoconch whorls of this specimen appear angular because of a prev;ilent spiral cord along the periphery. This is exaggerated by a break in the shell and subsequent repair, evidenced bv a thick- ened scar on the posterior part of the last whorl. The elongated shell and prominent siphonal canal are fea- tures shared with an immature paratvpe (paratype 3, fig- ures 11-12) of O. papi/rodes. This is the only specimen of Obscuranella with some portion of the earlv whorls intact (figure 14). The protoconch (2.0 mm estimated diameter) is eroded and replaced by a plug, but the ear- hest teleoconch whorls are well preserved and clearly show spiral cords. Two dead collected shells (figures 8-10; USNM 870610), labeled as coming from upper slope depths (210-220 m) off the Antarctic Peninsula, closely match the moiphology of this new species. We regard these specimens to represent O papijrodes, but are skeptical of the accuracy of the locality data. Not only is this lo- cation on the opposite side of the Antarctic continent from all records of live collected O. papijrodes, it is also from much shallower depths (220 m vs. 2000-1- m). DISCUSSION Ohscuraiudla can readily be attributed to the superfam- ily Tonnoidea on the basis of its pyriform shell with large aperture and conspicuous, if short siphonal canal; its ex- tensible proboscis; its taenioglossan radula; its large sal- ivarv glands composed of moqihologicallv differentiated anterior and posterior lobes and sahvarv ducts that pass through the nen'e ring, as well as its undifferentiated oesophageal gland. It can be excluded from Ficidae, which was removed from Tonnoidea and elevated to su- perfamily status by Riedel (1994), by its high spire, lack of long siphonal canal, and also because Ficidae is char- acterized by small, tubular sali\arv glands. Similarly, it can be excluded from Laubierinidae, a family diagnosed by its monopectinate osphradium and excluded from Tonnoidea by Bandel and Riedel (1994), by its nearly symmetrical, bipectinate osphradium. The shell of Obscuranella suggests an affinitv' with the deep-sea family Pisanianuridae (originall)' proposed as a subfamily of RanelHdae by Waren and Bouchet, 1990, transferred to Laubierinidae by Bandel and Riedel, 1994, and elevated to family status by Beu, 1998) by \irtue of its smooth shell lacking regular varices and weakly defined anterior canal. The operculum of Pis- anianiira is shghtly coiled but has a terminal nucleus, as does Obscuranella. Howe\er, the rachidian teeth of Ob- scuranella lack the lateral basal denticles present in Pis- anianuridae (e.g. Waren and Bouchet, 1990:figs. 25-27), Bursidae (e.g. Waren and Bouchet, 1990:figs. 6, 8), Ton- nidae (e.g. Waren and Bouchet, 1990:figs. 9-14), and Laubierinidae (e.g. Waren and Bouchet, 1990:figs. 41- 44), but absent in Cassidae (e.g. Waren and Bouchet, 1990:figs. 15, 16, 18), Personidae (e.g. Beu, 1998:fig. 15.140 E), and Ranellidae (e.g. Waren and Bouchet, 1990;figs. 28, 30, 32, 40). The shell of Obscuranella somewhat resembles that of Oocorijs sulcata Fischer, 1883 (Oocorythinae, Cassidae) Y. I. Kantor and M. G. Harasewvch, 2000 Page 111 (see, eg. Bouchet aiu! W'aivii, 19y3:fig,s. 1936-1943), and some ranellids, such as Ar^obticcinmn pnstulosuin (Ughtfoot, 1786) (.see e.^. Beii, 1998:fig. 15.12 D). The operculum of Obscuranella has a shaqjly pointed, ter- minal nucleus, and differs from the spiralK' coiled oper- culum of Oocon/s (W'aren and Bouciu't. 1990: fig. 66). In adult Ai-fiobuccinuin the nucleus ot the operculum is subcentnJ, hut in ver\' young specimens of A. pustulo- siim (Waren and Bouchet, I990:fig. 69) the nucleus is terminal. Howexer, the operculmn of Ohscuraiwlla is distincti\e in its veiy small size, relatixe to the aperture, and in having straight, anteriorly converging margins. The anatomv of Obscuranella is tvqiicallv tonnoidean, most closelv resembling that of the ranelliil Ci/inatiinn (Houhrick and Fretter, 1969). Obscuranella can be dis- tinguished anatomically from the Tonnidae by its lack of a buccal gland, and from Pisaniamiridae and Laubierin- idae hv its lack of long proboscis retractor muscles that pass through the nen'e ring. We assign this genus to the family Ranellidae because of it general similaritx' to Arfiobucchuun in shell form, radular moi-pholog\, and gross anatomy. Moreover, Ra- nellidae is the onlv tonnoidean familv to occur in Ant- arctic waters — Fusitriton ma^icllanicus (Roding, 1798), a species with a wide geographic range, has been taken from several stations in the Weddell and Ross Quadrants (e.g. USNM S96058, USNM 896103, USNM 896277, USNM 898520— see Polar Invertebrate Catalog http:// www.nmnh.si.edu/cgi-bin/wdlViz/pci/form). Obscuranel- la papi/rodcs represents the first record of Ranelfidae from abyssal depths. ACKNOWLEDGMENTS This study was supported by a USAP Research Program Award from the Biological Collections from Polar Re- gions at the National Museum of National History [Na- tional Science Foundation (Office of Polar Programs) Cooperative Agreement OPP-9509761]. We are grateful to Drs. Alan Beu and Klaus Bandel for their constructive reviews of this manuscript. LITERATURE CITED Aii(lrew.s, K. B., A. M. Page and ], 13. Taylor 1999. Tfie fine .stnictiire and function of the anterior foregnt glands of Ci/uuitiiim iiitenneclius (Cassoidea: Ranellidae). Journal of \iolluscan Studies 6.5:1-19. Bandel, K. and F. Riedel, 1994. Classification of fossil and Re- cent CaKptraeoidea (Caenogastropoda) with a discussion on neoniesogastropod phylogeny. Berliner geowissen- schafdiche Abhandlungen (E) 13:329-367. Beu, A. G. 1981. Australian gastropods of the family Bursidae. Part 1. The families of Tonnacea, the genera of Bursidae, and revision of species prexiously assigned to Tutufa Jous- seaume, 1881. Records of the Australian Museinn 33:248- 324. Beu, A.G. 1998. Superfamily Tonnoidea. In: Beesley, P L., G. J. B. Ross and A. Wells (eds). MoUusca: The Southern Synthesis. Fauna of Australia, volume .5, part B. CSIRO Publishing, Melboum, pp. 792-803. Bouchet, P. and A. Waren. 1993. Revision of the northeast Atlantic hathyal and abyssal Mesogastropoda, Bolletino Malacologico, Supplemento 3:.580-840. Day, J. A. 1969. Feeding of the cvniatiid gastropod Argofci/c- cintim argus, in relation to the stRicture and secretion of the proboscis glands. American Zoologist 9:909-916. Dell, R. K. 1990. Antarctic MoUusca. Bulletin of the Royal Society of New Zealand 27:1-311. Houhrick, J. R. and V. Fretter 1969. Some aspects of the func- tional anatomy and hiologv' of Cipnatium and Bursa. Pro- ceedings of the Malacological Society of London .38:41.5- 429. Hughes, R. N. and H. R I. Hughes. 1981. Moiphological and behavioural aspects of feeding in the Cassidae (Tonnacea, Mesogastropoda). Malacologia 20:385—102. Riedel, F. 1994. Recognition of the superfamily Ficoidea Meek 1864 and definiton of the Thalassocvniidae fam. nov. (Gas- tropoda). Zoologische Jahrbiicher 121:457-474, Riedel, F. 1995. An outline of cassoidean phylogeny (MoUusca, Gastropoda). Contributions to Tertiary and Quateniary Geolog)' .32:97-1.32. Waren, A. and P. Bouchet. 1990. Laubierinidae and Pisanianu- rinae (Ranellidae), two new deep-sea taxa of the Tonno- idea (Gastropoda: Prosobranchia). The Veliger 33:56-102. Weber, H. 1927. Der Darm von Dolium galea L., eine ver- gleichend anatomische Untersuchung unter besonderer Beriicksichtigung der rn^()»(i/»i-Arten. Zeitschrift fiir Morphologic tmd Okologie der Tiere 8:66.3-804. THE NAUTILUS 114(3):112-116, 2000 Page 112 Host-tree selection by Florida tree snails, Ligiiiis fasciatus (Miiller, 1774), in Big Cypress National Preserve, Florida, USA Robert E. Bennetts' Department of Wildlife Ecology and Conservation Florida Cooperative Fish and Wildlife Research Unit P.O. Box 110450, University of Florida GainesN-ille, FL 32611-0450 USA bennetts@tour-du-valat.com Steven A. Sparks Deborah Jansen Big Cvpress National Preserve HCR'61, Box 110 Ochopee. FL 34141 USA ABSTRACT It has frequently been suggested that the Florida tree snail, Liguiis fasciatus, exhibit preferences for specific host trees, al- though most accounts of host-tree selection are based on an- ecdotal observations. We assessed the relative use of host trees in eight hammocks in the Big C)'press National Preserve for which the relative availability of each potential host-tree spe- cies was known. Based on a total of 1,464 tree snail obsei-va- tions on 21 species of host tree within our study plots, host trees were not used in proportion to their availability when the expected values are derived either from the number of trees or from basal area. Of the most common tree species on our study area, wild tamarind (Ltjsiloma latisiliqurn) was consis- tentlv used in excess of its availabilitv. Wild lime (Zanthoxi/him fagarn), was not a common tree species, but was also used far in excess of its availabilitv. Pigeon plum {Coccoloba diversifolia) and gimibo-limbo {Bursern simanibti) were consistently used less than expected from their respective availabilities. Based on Akaike's Information Criterion (AIC), the most parsimonious loglinear model, after accounting for availabilitv, was one that included only tree species effects. In contrast to the model selected using AIC, likelihood ratio tests indicated that there may also be differences among hammocks, but not in relation to a linear gradient of hurricane damage. We obsers'ed differ- ences in length among 3-year-old snails on different host trees, and these differences were consistent with the hypothesis that host-tree selection reflects food availabilitv'. Snails found on host-tree species that were used in greater than expected fre- quency had the greatest length, those found on trees used less than expected frequency were the smallest, and those found in proportion to their availability were intermediate in length. Additional key iiords: Big Cypress National Preserve, host tree, humcane, Liguus fasciatus. tree snail, Florida, hammock. INTRODUCTION Snails of the genus Liguus are native to Cuba, Hispan- iola, and Florida (Pilsbry, 1912). The Florida tree snail ' Author for correspondence. Present address: Station Biolo- glque de la Tour du Valat, Le Sambuc. F-I320() Aries, France Liguus fasciatus (Mtiller, 1774) is found within a restrict- ed part of this range, primarily some islands within the Florida Keys, the Atlantic coastal ridge. Everglades Na- tional Park, and Big Cypress National Preserve. Primar- ily because of agricultural and urban expansion, only a few isolated populations remain outside of protected public lands and a few isolated islands in the Keys. Con- sequently, they are hsted as a species of special concern by the state of Florida. Thus, understanding the rela- tionship between snails and their habitat is essential for the conservation and management of remaining popu- lations. In the Everglades, L. fasciatus is found on islands of subtropical hardwood trees and scrub known as ham- mocks. Hammocks are widely scattered throughout South Florida and may be separated by water, sawgrass, or other habitat types such as pines. It is widely beUeved that Florida tree snails exliibit strong preferences for host trees, particularly smooth-barked trees upon which their algal food source can easily be grazed (Pilsbry, 1912). In particular, snails are suggested to exhibit pref- erence for Jamaica dogwood (Piscidia piscipula) in the Florida Keys and wild tamarind (Li/siloma latisiliqurn) on the mainland (Pilsbry, 1912), although most accounts of host-tree selection are based on anecdotal observa- tions, and few quantitative data are available. Voss (1976) reported numbers of snails found on each tree species in 2 hammocks on the coastal ridge. However, he did not account for the aviulabihty of these host trees; thus, cUfferential selection can not be reliably assessed. Brown (1978) is the only study of which we are aware that com- pared the use and availabihty of host trees, although this was restricted to 1 hammock in Everglades National Park. Here we assess the relative use of host trees in eight hammocks in the Big Cvpress National Preserve for which the relative availability of each tree species was knowni. Brown (1978) also hypothesized that differences in (juantitv' and qualitv' of food resources of different host- R. E. Bennetts ct al, 2000 Page 113 tree species mav result in size ditferences among snails using olifferent host trees, although she was unable to statisticallv confirm such clifferences. We tested this hy- pothesis h\ comparing size cbfferences among snails found on host-tree species that were used (1) in greater than expected fretjuencies (selected), (2) did not differ from expected frequencies (neutral), and (3) in less than expected frequencies (avoided). MATERIALS AND METHODS Our studv population was in Big Cypress National Pre- sene (BCNP). The Pinecrest area of the BCNP contains one of the largest remaining populations of Florida tree snails. Within BCNP, 136 hanunocks have been indi\'id- ually identified and mapped (Pilsbiy, 1946); a few others may e.xist that have not been identified. We selected 8 of the 136 hammocks from this pof)l for our study sites. Eight hanunocks proxaded a reasonable representation of the potential variability and represented a logistically feasible number of sites. This study was initiated to in- vestigate the survival and movements of Florida tree sn;iils following Hurricane Andrew. Hurricane Andrew crossed southern Florida from east to west on 24 August 1992. The eye of the storm passed along the southern boundarv of our study area creating a decreasing lati- tudinal gradient of damage toward the north with in- creasing distance from the eye. Because we were inter- ested in the potential effects of Hurricane Andrew, we also wanted our study sites to reflect the latitudinal gra- dient of hurricane impacts. Consequently, we divided the initial pool of hamiuocks into 4 zones reflecting clus- ters of hammocks positioned along a gradient from little or no hurricane impact to severe impact (Bennetts ct al., in press). There also exists considerable variabifity in the size of these hammocks, which we beheved could ha\'e greatly influenced their susceptibihty to damage by Hur- ricane Andrew and consequently affected tree snails. Therefore, we digitized each hammock from USGS 7.5- minute ortho photos and used a Geographic Information System (CIS) to estimate the appro.ximate size of each hammock. We then used stratified random sampling to select 1 hammock of greater size than the median size and 1 hammock of lesser size from each of the 4 impact zones. Thus, our sample reflected the latitudinal gradi- ent of hurricane damage and was balanced among larger and smaller hammocks. We estabhshed two 80 m- study plots (sub-samples) in each hammock. These plots were randomly located in parts of the hammock where tree snails occurred. The center of each plot was marked with a steel rod and we subsequently marked all trees with a diameter at breast height (DBH) > 5 cm within a 5 m radius of the plot center. We recorded the species and DBH of each live tree within these plots. During each of seven sampling occasions, we searched for all tree snails within each plot, and as part of a con- current study on movements (Bennetts et al.. in press), for marked snails within an extended radius of approxi- mately 20 HI around the plot. Sampling was conducted each autumn and spring from autumn 1993 through spring 1996. Our sampfing occasions were intended to reflect the biology of the animal and to minimize the potential for handling to influence a snail's survival. Our spring sampling occasion was conducted as soon as pos- sible after emergence of the snails from aestivation. This coincided with the onset of the annual rainy season. At this time snails begin to put on new growth, which be- comes yerv' fragile as it extends as a thin layer from the previous growth. Thus, our sampling was intended to precede the period during which time shells are fragile, although some growth had occurred on some snails. Our autumn sampfing occasion coincided with the onset of the dry season just prior to aestivation. At this time the summer's growth had been terminated, the shells had thickened, but the animals had not yet entered aestiva- tion. Animals that were visually determined to be in aes- tivation (<1%) by presence of an epiphragm (dried mu- cus membrane across the aperture) were not disturbed. For each snail we determined its age from annual growth scars (Tuskes, 1981), measured its length from tip to tip parallel to the axis and width perpentficular to the axis at its widest point. We also recorded the host tree from which it was collected. We then returned each snail to the same host tree by placing them in a conical paper cup attached to the trunk of the tree. Statistical Analyses: As a prefiminary analysis we test- ed for an association between use and availabiliK' of all host-tree species within our study plots using a chi- square goodness of fit test. This was done for a descrip- tive account of the relative selection of all host species; however, cells having expected values <5 could produce unreliable test statistics (Cochran, 1954). Thus, for fur- ther analyses including adcbtional effects of hammock of hurricane influence, we pooled cells with expected val- ues <5 into an "other " category. We tested whether host trees were used in proportion to their availabihty using a loghnear model. To account for host tree availability we used either the log number of trees, or the log basal area as an offset (Agresti, 1990; McCullagh and Nelder, 1989). Thus, the frequency of snails observed was modeled per available number and basal area of each species. We used basal area in addition to the number of trees because the surface area of trees are generally related to its bas;il area (Whittaker and WoodweU, 1967; Brown, 1978). We then tested whether the number of snails was influenced by tree species, and then whether this association cfiffered among hammock or hurricane zone. Model selection was based on Akaike's Information Criterion (AlC) (Akaike, 1973; Shibata, 1989), which is defined as: —llni'/) + 2np, where —2 ln(l''), repre- sents the Ukehhood ratio between the given model and a corresponding saturated model; thus, represents a measure of model fit. The second term, 2np. is the num- ber of parameters estimated in the model antl can be viewed as a cost for adding excessive parameters that do Page 114 THE NAUTILUS, Vol. 114, No. 3 Table 1. Common and scientific name of the host-tree species within our study plots. Also shown are the total number of trees, total basal area, and total number of snails observed for each species. Common name Scientific name Acronvin No. trees Basal area No. snails Cocoplum Chnjsobalanus icaco Ci 1 174 1 Dahoon holly Ilex cassine Ic 2 365 21 Gumbo Umbo Bursera simaniha Gl 26 5423 85 Hackberry' Celtis laevigflta CI 2 212 15 Inkwood Exothea paniculata Ep 2 93 3 Lancewood Nectandra coriaceae No 25 2362 52 Wild lime Zanthoxi/him faopra Zf 1 79 20 Li\e oak Quercus virainiana Qv 2 480 2 Wild tamarind Li/siloina latisiqua LI 128 41820 983 Mastic Mastichodendron foetidissimii m Mf 2 711 8 Paradise tree Simarouba gjmica Sg 2 240 11 Pigeon plum Coccoloba diversifolia Cd 80 9489 145 Poisonwood Metopium taxifera Mt 2 252 5 Pond apple Annona glabra Ag 1 47 1 Red bay Persea borbonia Pb 1 620 9 Red stopper Eugenia rhombea Er 1 22 1 Satinleaf Ch n/.wphijUu m olivifomie Co 1 85 1 Simpson stopper Mtjrcianthcs fraarans Mf 6 723 18 Strangler fio; Fiats aurea Fa 1 90 2 White stopper Eugenia axillaris Ea 7 273 9 \\'illow bustic Biimelia salicifolia Wb 19 2646 72 not contribute substantially to the overall model fit. Thus, AIC indicates an appropriate balance between precision and bias (i.e., over and under-fitting the model) from a set of candidate models (Bumham and Anderson, 1998). In contrast to AIC, which is used as an optimi- zation tool for comparison among models, we also used Ukelihood-ratio tests (LRTs) to test for specific effects of interest (Lebreton et al, 1992; Bumham and Anderson, 1998). The selection of host trees may be influenced by the quantity and quality of food resources (Brown. 1978), which mav in turn result in size differences of snails using different host trees. We tested this hypothesis by comparing size differences among snails found on host- tree species that were used (1) in greater than expected frequencies (selected), (2) did not differ from expected frequencies (neutral), and (.3) in less than expected fre- quencies (avoided). We compared the size of snails (i.e., length and width) within each age class among host-tree species using a fixed-effects analysis of variance (ANO- VA) model. For this analysis we were not concerned with the avaHabihty of host trees, only species. Consequendy, we included all snails encountered on our studv ham- mocks for this analysis, regardless of whether they were found within the designated plots (used to compare se- lection with availabihty) or the extended radius (used for analyses of movement) (Bennetts ct al, in press). How- ever, we restricted our sample to snails collected during the autumn sampling occasion to minimize any con- founding attributable to variation in the extent of new growth during spring. RESULTS We observed 1464 tree snails on 21 host-tree species within our study plots (Table 1). Host trees were not used in proportion to their availabiUtv when the expect- ed values are derived either from the number of trees (X' = 554.18, 20 df P < 0.001) or from basal area (x' = 296.14, 20 df P < 0.001). Of the common tree spe- cies on our study area, wild tamarind was consistently used in excess of its availability (figure 1). Wild Ume (Zant]uixijlum fagara). was not common, but our obser- vations indicated that it was used far in excess of its availabilitv', regardless of which measure of availabihty was used. Although the sample for this species used in our analysis included only 1 tree, we observed others on our study area that had similar high use. In contrast, pigeon plum [Coccoloba diversifolia) and gumbo-hmbo (Bursera simaniha) were relatively common, but were consistentlv used less than expected. Willow bustic (Bu- melia salicifolia) and lancewood (Nectandra coriaceae) also were relatively common in our study plots, but our results were conflicting for these species. In relation to the number of trees, willow bustic was used slightly less than expected whereas, based on basal area, it was used shghtlv more than expected. Lancewood was used less than expected in reference to the number of trees, but in proportion with its availabihty- in relation to basal area. After pooling cells with low expected values, the most parsimonious model based on AIC included only tree species effects (x" = 101.05, 5 df, P < 0.001). Depar- tures from ex-pected values were consistent vdth our pre- hminary analysis in that wild tamarind was used in great- er proportion than expected from its avaflabifit)-; pigeon R. E. Bennetts et al, 2000 Page 115 LI Zf Ic Pb CI Sg Mf Fa Ml Ag Ci Er Co Wb Mf Ep Qv Gl Ea Nc Cd Host-Tree Species llll ■ ■ Zf LI CI Ic Sg Wb Ea Er Ep Mf Fa Ag Nc Mt Co Pb CI Mf Qv Gl Cd Host-tree Species Figure 1. Adjusted residuals (Haberinan 1973) from good- ness-of-fit test between the number of snails found on different host-tree species in relation to their availability. Host-tree spe- cies are displayed in rank order from selected in greatest pro- portion relative to its availabilit)' to least. Expected values were derived based on the number of trees of a given species (top) and the total basal area for a given species (bottom). Positive residuals >1.96 indicate use greater than expected from avail- ability and residuals < — 1.96 indicate use less than expected from a\ailability. plum and gumbo limbo were used less than expected. Lancewood and willow bustic were intermediate as was our "other" class. In contrast to the model selected using AIC, LRTs indicated that there may have been a species X hammock interaction (x' = 110.36, 31 df, P < 0.001), but not a species X hurricane zone interaction (x" = 9.32, 5 df, P = 0.097). Thus, our data indicated that selection of host-tree species may have differed among hammocks, but this difference was not in relation to a gradient of hurricane damage. We found size (length) differences among snails found on host tree species onlv for 3-year-old snails (F^ j,^ = 3.42, P = 0.034) (figure 2). Of the.se, the length of sniiils was highest for snails found on host trees that were used in greater frequency than their availabibty (selected) and lowest for snails found on host trees that were used less than their availabihtv' (avoided). Sn;uis found on host trees that were not out of proportion to their availabihty E E o> c 40 - 263 53 - 39 - 35 38 - 37 - Selected Neutral Avoided Host-tree Selection Class Figure 2. Mean length (±SE) of .3-vear-old tree snails found on host-tree species that were used in excess ot their avail- ability (Selected), less than their availabilitv' (Avoided) and did not differ from availabilitv (Neutral). Sample sizes are shown for each group. (neutral) were intermediate in length, but did not differ from either of the other groups (Fisher's Least Signifi- cant Difference, P>0.05). We did not find differences in width among any host-tree species for any age class at the a = 0.05 level. DISCUSSION Our data are consistent with previous reports of wild tamarind being used in excess of availabihty; although the degree of this association is highly sensitive to what measure is used to derive the e.xpected values for avail- ability. When the expected value is derived from basal area, the association is much weaker than when the ex- pected value is derixed from the number of trees. Ex- pected values derived from basal area probablv better reflect the surface area available tor foraging. The num- ber of stems would be important if the probabifity of a dispersing snail moving to a given tree is dependent on it encountering a trunk on the ground, although most inter-tree movement on our study area probably was via intertwined foliage. Brown (1978) reported a remarkably similar result in Everglades National Park. She reported an overall significant test statistic based on both number of tree stems and estimates of bark surface area, al- though the relationship was substantially weaker based on bark surface area. Regardless of the statistical signif- icance, 983 of 1464 (67%) of all snails we observed were found on wild tamarind. Voss (1976) reported a similar estimate of 34 of 50 (65%) for 1 hammock in the eastern Everglades near Miami, and Brown (1978) reported 209 of 39.5 (52%) in another location in Everglades National Park. Although other trees niav be preferentially select- ed if available, wild tamarind was the most abundant and Pase 116 THE NAUTILUS, Vol. 114, No. 3 most used host tree within the Big Cypress National Pre- serve, which is probably also the case in most of the mainland habitats. Thus, wild tamarind is clearly an im- portant component of tree-snail habitat in this region. Although our model selection did not support that host-tree selection differed among hammocks, an LRT chd indicate a species by hammock interaction effect. Hammocks differed with respect to which host tree spe- cies were present, which Ukely reflects inter-hammock differences in attributes such as size, soils, and hydro- logic regimes. Thus, the interaction effect we observed could have been attributed to these differences in spe- cies composition, but our data are insufficient to evaluate such effects with much reliability. Bias due to \isibilit\- of snails was not accounted for by our approach. This effect would have been most in- fluenced by dense foliage obscuring snails from view. For most host-tree species, we believe that this bias was negligible, especially because of reduced foliage as a re- sult of Hurricane Andrew. However, one notable excep- tion was pigeon plum, which often had ver)' dense fo- liage making it difficult to detect snails. Thus, we suspect that our data indicating that this host tree species was underutilized may have been, at least in part, an artifact of detection bias. Although our data are far from conclusive, they are consistent with Browni's h\-pothesis that there mav be an association between selection of chfferent host trees and size of the snails. We found size differences only among 3-year-old snails. However, even our relatively large sam- ple sizes tend to become quite small once partitioned by age and host tree species. Florida tree snails exliibit most growth during their first 2-3 vears, after which growth slows dramatically (Tuskes, 1981). Thus, 3-year-old snails probably represent a peak of cumulative growth, and are a hkely age class to observe size differences. For this age class, snails found on host-tree species used less than expected were smallest in length, those found on trees used in excess of availabihty were the largest in length, and those found in proportion to their availability were intermediate in length. ACKNOWLEDGMENTS We thank Jimmy Conner, Phil Darby, Sue Davis, Vicky Dreitz, Guy Fischer, Paul Hinchcliff, Amy Kazmier, Steve McGehee, Katie Golden, Teresa Johnson, Jeff Ripple, Tim Towles, Patty Valentine-Darby for field as- sistance, and George Francioni for safe helicopter flights. We also appreciate the helpful comments from 2 anonymous reviewers. Funding was provided by the Na- tional Park Service through the South Florida Research Center, and Big Cypress National Preserve provided ad- ditional logistic support. LITERATURE CITED Agresti, A. 1990, Categorical data analysis. John Wiley and Sons, New York. 5.5Spp. Akaike, H. 197.3. Information theory' and an e.xtension of the madmuni likelihood principle. In: B. Petrov and F. Czaldl (eds.) Proceedings of the 2'"' International Symposium In- formation Theory, Akademiai Kiado Budapest, pp. 267- 281. Bennetts. R. E., S. A. Sparks, and D. Jansen. In press. Factors influencing movement probabilities of Florida tree snails Liguiis fasciatiis (Miiller) in Big Cvpress National Pre- serve following Hurricane Andrew. Malacologia. Brown, C. A. 1978. Demography, dispersal, and microdistri- bution of a population of the Florida tree snail Liguus fasciatiis. M.S. Thesis, Uni\ersitv of Florida, Gainesville. Buniham K. P. and D. R. Anderson. 1998. Model selection and inference: a practical infoniiation-dieoretic approach. Springer-Verlag, New York. Cochran, W. G. 19.54. Some methods for strengthening the common chi-square tests. Biometrics 10:417-451. Haberman, S. J. 1973. The analysis of residuals in cross-clas- sified tables. Biometrics 29:205-220. Lebreton, J. D., K. P Bumham. J. Clobert, and D R Ander- son. 1992. Modeling survival and testing biological hy- potheses using marked animals; a unified approach with case studies. Ecological Monographs 62:67-118. McCuUagh. P. and J. A. Nelder 1989. Generalized hnear mod- els, 2nd edition. Chapman and HaU, New York, 511 pp. Pilsbry, H. A. 1912. A study of the variation and zoogeography of Liguus in Florida. Journal of the Academy of Sciences of Philadelphia 15:429-470. Pilsbry, H. A. 1946. Land Mollusca of North America. Acad- emy of Natural Sciences Philadelphia, Monograph 32:1- 520. Shibata, R. 1989. Statistical aspects of model selection. In: J. C. Williams (ed.) From data to model. Springer-Verlag, New York, pp. 21.5-240. Tuskes, P. M. 1981. Population structure and biology of Liguus tree snails on Lignumxitae Key, Florida, The Nautilus 95: 162-169. Voss, R. S. 1976, Obserxations of the biology of the Florida tree snail, Liguus fasciatiis (Miiller). The Nautilus 90:6.5- 69. Whittaker, R. H. and G. M. Woodwell. 1967. Surface area re- lations of woody plants and forest communities. American Journal of Botany 54:931-9.39. THE NAUTILUS 114(3): 117-1 19, 2000 Page 117 The development of three heterobranch mollusks from California, USA Rachel Collin Committee on E\olutionar)' Biologv' Uni\ersits' of Chicago 1025 E. 57th St. Chicago, IL 60637 USA and Department of Zoology Field Museum of Natural Histor\' 1400 S, Lake Shore Drive Chicago, IL 60605 USA rcoHinCSmidwav. uchicago.edu ABSTRACT The de\elopnient of Odostomin nitinn. Turbonilla sp., and Wil- hamia peUoides from Cahtoniia, is t)pical ot heterobranch mol- lusks. Thev all produce small eggs (about 60 |jLni) that are in- di\idually encapsulated and embedded in gelatinous egg mas- ses. The capsules are connected by chalazae. Cleavage is un- equal and gastrulation occurs by inxagination. Plankiotrophic larvae hatch at shell lengths of 120-150 |xni. Additional key words: Pyramidellidae, Tiirbonilln. Odostomia, gastropod development, Williamin INTRODUCTION Although lower heterobranchs and marine pulmonates are common in shallow marine waters, their develop- ment remains largely undescribed. Knowledge of their development may be useful and important because de- velopmental features such as the structure of egg mas- ses, cleavage type, presence of the larval pigmented mantle organ (PMO), and hydrophobic larval shells may- be important characters for phylogenetic stucbes (Rob- ertson, 1985; van den Biggelaar, 1996; van den Biggelaar and Haszprunar, 1996; Collin, 1997). Additionally, there are a variety of interesting trends in the evolution of cleavage patterns (Freeman and Lundelius, 1992; van den Biggelaar and Haszprunar 1996) and heterochro- nies in larval morphologies (Page, 1994) whose docu- mentation could benefit from more extensive phyloge- netic sampling. Herein I describe the development of two pyranii- dellid and one siphonariid pulmonate species from the Califomian coast. All animals were collected by hand in the summer of 1997 and kept in small dishes in the laboratory where thev laid egg masses. Egg masses were observed daily until hatching, but no attempt was made to raise the larvae to metamorphosis. RESULTS Odostomia altina Dall and Bartsch, 1909 (Table 1) Large numbers of Odostomia altina were found on the shells oi HaliotLs cornigflta collected at a depth of 10 m from Point Loma, California (29°40'N, 117°20'W). Specimens are deposited at the Field Museum of Nat- ural History (FMNH 282369 and 282370) and were identified by comparison with the original species de- scription and illustrations in (Dall and Bartsch, 1909). There were as many as 50 adult pvTamidellids and nu- merous egg masses on one abalone. Egg masses were typical of other pvTamideOids: Single eggs are enclosed in albumin-filled oval capsules that are embedded in a clear sticky gelatinous mass. The capsules are connected by thin extensions of the capsular covering called cha- lazae (see Robertson, 1985; Collin and Wise, 1997 for detailed description). A timetable of development is giv- en in Table 1. The round, white eggs are 60 |jLm in diameter (mean = 60.83 |jLm; sd = 1.39 |xm; n = 31; eggs from 3 egg masses) the inner capsule diameter is 150 (xm (sd = 8.11 |xm; n = 19) and the outer capsule diameter is 176 \x.m (sd = 6.992 |xm; n = 10). First cleavage is unequal and at the 4-cell stage the two largest cells are adjacent, as are the two small ones. Several embryos were observed in a 3-cell stage, which suggests that second cleavage is not synchronous. There is no polar lobe and third cleav- age is unequal. The round blastula gradually flattens and invaginates to form a horseshoe shaped gastrula. During subsequent development the embryo grows to fill the capsule and the larval organs differentiate. Plank-totroph- ic larvae hatch after about 7 days (Table 1) with 130 |xm (mean = 130.33 |xm; sd = 8.55 [xni; n = 15; egg masses from two females) smooth, left-handed, hydrophobic shells. Each larva has a well-developed metapodial ten- Page 118 THE NAUTILUS, Vol. 114, No. 3 Table 1. De\elopnient schedule for Odostomiti nltinn at 22- 24°C. Age Stage >3 hours 2-cells 5 hours 4-cells 12 hours 64-cells 1 dav blastula 1.5 days gastnilation by invagination 3 days just moving, foot and velum anlangen visible 5 days red-yellow PMO visible, shell operculum and statocvsts ~7 days hatchmg at 130 (jim tacle and statocysts, but no detectable heartbeat, and no eyes. The larval PMO consists of two parts: A round red central part and an additional semi-transparent yeUow lobe on one side. Observation of the yellow lobe under the dissecting microscope is difficult, but the structure can be seen easilv' under a compound microscope. Turbonilla sp. (Figures 1, 2, table 2) One adult of an unknown species of Turbonilla was col- lected on the high mid-intertidal zone at Alegria, Cali- fornia (34°28'N,^120°17'VV) in July 1997. The species identification is uncertain because Turbonilla taxonomy is especially difficult and the single specimen could not be unambiguously assigned to any Californian species. The current taxonomy of Californian pyramidellids as- signs all species with axial-ribbed shells to the genus Tur- bonilla (Dall and Bartsch, 1909). This snail laid one egg Figure 2. Earlv developmental stages of Turbonilla .sp. .\. 2- cell stage at the beginning of second cleavage, the cells are clearly not equal in size, B. 4-Cell stage, C. blastula, D. gas- trula, showing the invagination typical of heterobranch devel- opment. Scale bar = 50 \x.m. mass of about 100 eggs in the laboratory and develop- ment was followed until hatching (table 2 and figure 2). The egg mass consisted of capsules containing one white egg surrounded by clear albumin, connected together by chalazae and embedded in a clear sticky gel. The egg diameter was 65.5 |jLm (n = 10, sd = 1.97 |jLm) and the capsule length was 187.8 |xm (n = 10, sd = 6.67 (im). The chalazae are thicker and less tv\dsted than in O. col- umbiana Dall and Bartsch 1907 (Colhn and Wise, 1997). First cleavage is slightly unequal but bv the 4-celI stage the blastomere inequalit)' is yery subtle. The early cleav- ages do not produce a polar lobe. The polar bodies are clearK visible throughout development because the al- bumin is transparent. After 24 hours the embrvos form flattened blastulas, which begin to invaginate at about 40 hours. Gastnilation continues by invagination for the next dav. Bv the fourth dav the foot and velum anlagen are visible. On the fifth dav the embrvos begin to move Table 2. Developmental schedule for Turbonilla sp. at 16- 18°C. Figure 1. Adult Turbonithi sp. Shell length is 6 mm. Age Stage 7 hours 2-cells 24 hours blastula 2-3 days gastnilation by invagination 4 days just moving, foot and velum anlangen visible 5 days shell and velum differentiated. Embrvo fills 1/4 of the capsule 6 davs black PMO visible 9 days embno fills the capsule 11 days hatching at 152 |j.m R. Clolhn, 2(M)() Page 119 and thev fill alidiit a (juarter of each (.apsiilf. B\' thf next cla\' the black PMO is \isihle on the right side just pos- terior to the velum. The enibno grows to completely fill the capsule, before hatching. Shell length at hatching is 1.52 jjLni (n = 14; sd = 2. .37 p,m; from a single egg mass). The shell is hydrophobic, left-handed and the lana has no heart or eyes but has a well-developed metapodial tentacle on the foot. The planktotrophic lanae swim ac- tively. Williamia pchoides (Carpenter, 1864) Several adult W. peltoidcs (Carpenter, 1864) were col- lected in the subtidal zone ( — 10 ni), on hard substrates, in the vicinity of Santa Barbara, California (.34°28'N, 120°17'W). Voucher specimens are deposited at the Bai- ley-Matthews Shell Museum, Sanibel, Florida (BMSM 4999). Williamia species are unusual among marine pul- monates because they are predominately subtidal. Adults produced several egg masses in chshes in the laboratory. The structure of the egg masses is verv similar to those of the pvTamidellids. The eggs are each individu- ally enclosed in a coating of albumin inside an oval cap- sule. The capsules are connected with chalazae and are embedded in a gelatinous mass. The capsules are 147 jjim in length (mean = 147.7 ji-m; sd = 3.4 |xm; n = 11). Unfortunately uncleaved eggs and early cleavage stages were not observed. Gastrulation occurs via invag- ination forming a horseshoe-shaped gastnila that is 69 (xm in diameter (d = 69.28 |xm; sd = 3.4 |jLm; n = 7; from one egg mass). Further development progressed as in the pyramidellids. However, no pigmented PMO de- velops: I could not determine if there was no PMO or if the structure was present but not pigmented. There is some reddish pigmentation along the suture of the larval shell, but the larval body was pigmentless. After 9 days at 18-22 °C the larvae hatch at a length of 126 |xm (length = 126.0 jjuni; sd = ■5. .58 |jLm; n = 10). The plank- totrophic lar\ae have a hydrophobic smooth left-handed shell. There is still no pigmented PMO, no eyes, and no metapodial tentacle but the statocysts are clearly visible. DISCUSSION These observations are the first descriptions of intracap- sular development for species in the genera Williamia and Tttrhonilla. The features of pyramidellid development described here generally agree with prexaous descriptions of pyr- amidellid development. Because Tnrhoiiilla and Odos- tomia are distantly related genera within the Pyramidel- lidae (Wise, 1996), characters shared by the species de- scribed here and O. cohtmhiana (Colhn and Wise, 1997) may be typical of pyramidellid development in general. All three species have small eggs, unequal cleavage, gas- tnilation bv invagination, and hatch with smooth, left- handed, hydrophobic shells, a metapodial tentacle, dis- tinct PMO, but without eyes or a larxal heart. The color oi the larval PMOs varies among species as does the cok)r of the adult PMO (Robertson, 1985). The development of Williamia peltoidcs is strikingly similar to pyramidellid development. The structure of the egg masses is indistinguishable from that of the pyr- amidellids. Hatching lar\ae are distinct in their lack of a metapodial tentacle and pigmented PMO and the small area of shell pigment. These results are in agree- ment with Robertson's (1985) conclusion that larval PMOs are not present in pulmonates while chalazae and heterostrophy are common. The retldish pigment along the shell suture is also found in larvae of Siphouaria sp. from New Zealand (pers. obs.). ACKNOWLEDGMENTS This research was conducted during a visit to the labo- ratories of Dr S. Gaines (University of California at San- ta Barbara) and Dr K. Roy (University of CaUfornia at San Diego). I thank Ron McConnaughev and Shane An- dersen for helping collect animals and two anon\iiious reviewers for helpful comments on previous versions of this manuscript. This work was supported financially by a NSF Predoctoral Fellowship, and grants from the Western Society of Malacologists, Sigma Xi, the Lemer Gray Fund (American Museum of Natural Historv) and the Hinds Fund (University of Chicago). LITERATURE CITED Biggelaar, J. A. M. van den. 1996. The significance of the early cleavage pattern for the reconstruction of gastropod phy- logeny. In: Taylor, J. D. (ed.) Origin and Evolutionary Ra- diation of the Molliisca. Oxford University Press, Oxford, UK. Biggelaar, J. A. M. van den and G. Hazsprunar 1996, Cleavage and niesentoblast formation in the Gastropoda: An evo- lutionarv perspective. Evolution .50:1.520-1.540. Collin. R. 1997. Hydrophohic larval shells: Another character for higher level systematics of gastropods. Journal of Mol- luscan Studies 6.3:425-430. Collin, R. and J. B. Wise. 1997. Morphology and development of Odostomin cohnubiann (Gastropoda: PyTamidellidae). Biological Bulletin 192:24:3-252. Dall, W. H. and P. Bartsch. 1909. A monograph of west Amer- ican pvTamidellid mollusks. Bulletin of the United States National Museum 68:1-258. Freeman, G. and ]. W. Lundelius. 1992. Evolutionaiy' impli- cations of the mode of D quadrant specification in coe- lomates with spiral cleavage. Journal of Evolutionary Bi- ology 5:20.5-247. Page, L. R. 1994. The ancestral gastropod larval torm is best approximated by hatching-stage opisthohranch larvae: ev- idence from comparative developmental stuthes. In: Wil- son, W. H., S. A. Strieker, and G. L. Shinn (eds.) Repro- duction and Development of Manne Invertebrates. Johns Hopkins University Press, Baltimore. Robertson, R. 1985. Four characters and the higher category systematics of gastropods, American Malacologica! Bulle- tin, Special Edition 1:1-22. Wise, J, B, 1996. Moqiliologv and phylogenetic relationships of certain pyramidellid taxa (Heterobranchia). Malacologia .37:44.3-51 r. THE NAUTILUS 114(3): 120-126, 2000 Page 120 Diminishing species richness of mollusks in Oneida Lake, New York State, USA Willard N. Harman State Universitv' of New York College at Oneonta Biological Field Station RD2, Box 1066 Cooperstown, NY 13326 USA harmamvn@oneonta.edu ABSTRACT Between 1915 and 1917, F. C. Baker studied the molluscan fauna of Oneida Lake, a shallow, eutrophic lake in central New York State. He hsted 41 living taxa, of which two gastropods, Bithynia tentacitlata (Linnaeus, 1758) and Pleurocera aaita (Rafinesque, 1S29), were then recent introductions. In 1967- 68, John Foniey and I (Harman and Foniey, 1970) surveyed Oneida Lake, \'isiting Baker's original collecting sites. W'e tal- lied 35 Uving taxa dominated by the introduced European B tentacitlata. In 1992-95, I again made extensive collections of mollusks finding a total of 24 h\ing taxa dominated bv Dreis- sena pohjmorpha (Pallas, 1771), the then recently introduced Eurasian zebra mussel. Fifty percent of Baker's original euht- toral collection sites could not be found. An average 1 m- area in 1992-95 included 6708 individuals of D. pohjmorpha. 60 of B. tcntactilata and 2 of the unionid bivalve EUiptio complanata (Lightfoot, 1786). Molluscan species richness was reduced by 15% between 1917 and 1968, a trend concurrent with the in- crease in abundance of Bithynia. Species richness was hirther reduced by 31% between 1968 and 1995, as human acti\-ities impacted euhttoral habitats and D. polymorpha colonized Oneida. The total decrease in species richness between 1917 and 1995 was of 42%'. Since 1996 no li\ing miionids ha\e been observed in the lake. Additional key words: Exotic introductions, habitat destruc- tion, freshwater, gastropods, bivalves, Bithynia tentacitlata. Dreissena pohjmorpha. INTRODUCTION Oneida Lake is a eutrophic lake in central New York State, with about 233 kTn- in surface area and a maxi- mum depth of 16.8 m. It is located in a depression on the low, open rehef of the Ontario Lake Plain in the Oswego River drainage basin (figure 1). The lake is ori- ented with its length parallel to that of the prevailing westerly winds. It mixes throughout the ice-free period, with thermal stratification occurring only ephemerally during wind-free periods (Harman and Forney, 1970). Tributaries to Oneida drain densely populated areas of Onondaga Limestone and fertile soils. As a result, the lake has been considered one of the most naturally pro- ductive in the world (Mozley, 1954). Historically, popu- lations of algae and aquatic vascular plants have thrived in the shallow waters along the shoals, beaches, and is- lands, on diverse eulittoral substrates providing food and cover for a varied and abundant molluscan fauna (Har- man and Forney, 1970). Central New York has a diverse molluscan fauna de- rived from species immigrating from Atlantic coastal riv- ers, the Mississippi River drainage (Interior Basin), and the Great Lakes, soon after the retreat of the last Pleis- tocene glaciers (Clarke and Berg, 1959). Oneida Lake is unique due to its central position on the New York Barge Canal system and natural waterways (figure 1), which have continued to provide access to immigrating organ- isms from the early 1800s. Since then, mollusks from Lake Erie and the Ohio River basins in the west, and from the Atlantic coastal drainage in the southeast (via the Mohawk and Hudson drainage basins) have found their ways to Oneida Lake. Headwaters of several wa- tercourses flowing from the Appalachian highlands in the southern Oswego system join the Susquehanna drainage via through-viilleys, which facilitates dispersal of aquatic organisms. The Oswego River drains Oneida flowing to Lake Ontario and, via the St. Lawrence River, joins Lake Champlain and other northern waterways, thus providing access to and Irom North Atlantic drain- age systems. During 1915-17, F. C. Baker studied the macroben- thos in Oneida Lake, concentrating on the molluscan fauna excepting the fingernail clams (Sphaeriidae) (Bak- er, 1916a, b, c; 1918a, b, c, d). His research on Oneida's Lower South Bay is among the earliest quantitative stud- ies of freshwater macrobenthos in North America (Bak- er, 1918c). Baker studied Oneida because of its great diversity of mollusks. In 1967 and 1968 I (Harman and Forney, 1970) conducted, as part of a survey of the gas- tropods of the Oswego River drainage basin, a qualitative study of the molluscan fauna of Oneida Lake. In the summer of 1967, Forney (Harman and Forney, 1970) repeated Baker's (1918c) quantitative work in Lower South Bav as part of a macrobenthic monitoring pro- gram carried out by Cornell University. The introduction of zebra mussels {Dreissena poly- mor-f)ha) into Oneida Lake in 1990 (Mitchell et al. 1996) W. N. Hai-man, 2000 Page 121 Figure 1. Central New York State showing Oneida Lake and routes of immigrating mollusks. has had important impacts on water clarity by reducing phvtoplankton standing crops. It also appears to be af- fecting the distribution and abinidance ot aquatic vas- cular macroph\1:es (E. L. Mills, pers. comm.). The ir- ruption of the Drcisscna population was expected to have tremendous impacts on the molluscan fauna (e.g., Nalepa ct al. 1991; Ricciardi ct al, 1995; Schloesser ei al, 1996). That concern prompted my return to Oneida during 1992-95 to determine the status of the molluscan communitv. I effectively repeated my (Harman and For- ney, 1970) 1967-68 qualitative work and in 1993 did a quantitati\'e studv in Lower South Bay in an attempt to repeat Baker's 191.5-lS studies (1918c) as well as For- ney's 1967 work (Harman and Forney, 1970). This paper describes the changes in the molluscan fauna since 1915, showing evidence of a constant reduc- tion of species richness concurrent with destruction of euhttoral habitats and invasions of immigrating mollusks. MATERIAL AND METHODS In 1915-17, during the summer months. Baker obtained (juantitative samples in Lower South Bay using a sieve, flattened on one edge, attached to a wooden pole, to collect bottom samples of about 10- cm-. A variety of larger dredges were used in deeper water A square met- al frame, viewed through a glass-bottomed bucket, was used to estimate densities of large snails and bivalves. Collections were made while wading, or from a boat (1918c). Water depths in Lower South Bay are such that these methods allowed him to sample all areas of the bay. Baker's samples at each collecting site approximated 10- cm-. He used varying numbers of replicates, typically 2-6. All his computations were presented using 10- cm- (16 in-) as the basic unit. My 1967-68 qualitative survey was made throughout the lake in an attempt to sample every habitat suitable for mollusks. Samples were retrieved from inorganic substrates str;iined through a sieve or handpicked vsatli forceps in shallow water Vegetation was collected and dried over containers for retrieval of desiccating organ- isms. Collections from deeper water were done similarly using SCL'BA. Fornev contributed several Ekman grab samples to that study. Forney's replication of Baker's quantitative study in Lower South Bay, again during the summer months, used Baker's methods as mentioned aiio\e. However, he relied more on standard Ekman grabs. Calculations were based on 10- cm- sample areas to facilitate comparison with Baker's work. Eleven of Baker's original sample sites (out of a total of 140) were lost due to artificial modifications of the shoreline. In 1992-95 the same methodologies were used for the qualitative work, over the same time period that Forney (Harman and Forney 1970) had covered in 1967-68. The 1992-95 (juantitative work in Lower South Bay in- voked Ekman samples in deep water and square metal frames in eulittoral waters. Calculations were based on 1 m- sample size. Information is presented using this size, as well as 10- cm- for comparison with the earlier studies. It should be noted that when converting to 1 m- from the earlier 10- cm- sample size, diversity is poten- tially underestimated since widely separated large indi- viduals such as unionids often encountered in the former are rarely foimd in the latter samples, and thus never appear in the data set. Only 71 of Baker's 140 original sites were still in existence in the recent survey. All titx- onomic nomenclature herein has been updated accord- ing to Turgeon ci al (1998). RESULTS AND DISCUSSION Table 1 indicates the taxa that Baker encountered at the turn of the century. Baker's revised list includes 41 fiving molluscan taxa inclutling the then recently introduced European snail Bithijnia tent acid at a, which had ap- peared around 1880 in Lake Ontario (Beauchamp, 1888). Bithi/nia tcntavtilata was assumed to have been introduced via ballast water in commercial seagoing ves- sels plying the Great Lakes (Baker, 1916b). The species was not abundant in Oneida Lake in 1915-17, but oc- curred in dense populations near the outlet at the west end. An average 10- cm- bottom sample made in 1917 contained 7 taxa, all consisting of native species {Pyr- gulopsis lustrica (Pilsbry, 1890) (Amnicola htstiica, A. lustrica (var) and A. oncida in Baker's original lists), Stagnicola catcscopiiiin (Say, 1867) (Galba catascopium of Baker), Gi/ratilus panii.s (Say, 1817) {Planorhis par- vus of Baker), FhijscUa gyrina (Say, 1821) (Physa gyiina of Baker), P. Integra (Haldeman, 1841) (Physa intcgra of Baker), Prmncnetus cxacuoiis (Say, 1821) [Planorhis exacuous of Baker), and Valvata tiicarinata (Say, 1817)) (figure 2). A second introduction, Plcnroccra acuta. Page 122 THE NAUTILUS, Vol. 114, No. .3 Table 1. The moUu.scan fauna of Oneida Lake; reference conditions, 191.5-17 (modified from Baker, 1918). Taxa Bivalvia: Palaeohetcrodonta Margaritifera maraarilifera (Linnaeus, 17.58) Elliptio complanata (Lightfoot, 1786) Alasmidonta undidata (Say, 1817) Pijganodon grandis (Say, 1829) Pijganodon cataracta (Say, 1817) Strophifus undulatus (Say, 1812) Villosa iris (I. Lea, 1829) Lampsilis radiata radiata (Gmelin, 1791) Actinonaias ligmcntina (Lamark, 1819) Total number of species: 41 Gastropoda: Basominatophora Phi/si'lla gtjrina (Sav, 1821) Phi/si'lla Integra (Haldeman, 1841) Phi/sella heterostropha (Say, 1817) Lijmnaea stagnalis (Linnaeus, 1758) Fossaria obrussa (Say, 1825)* Stagnicola elodes (Say, 1821) Stagnicola enwrginata (Say, 1821) Stagnicola catescopium (Say, 1867) Pseudosiiccinea columella (Say, 1817) Acella haldniani (W. G. Binney, 1867) Helisomn anceps (Menke, 1830) Phanorbella trivolvis (Say, 1817) Planorbella campanulata (Say, 1821) Gyraulus parvus (Say, 1817) Gyratdus deflectus (Say, 1824) Promenetus exacuous (Say, 1821) Laevapex fuscus (C, B. Adams, 1841) Ferrissia parallelus (Haldeman, 1841) Ferrissia rivularis (Sav, 1817) Gastropoda: Caenogastropoda Campeloma decisum (Say, 1817) Vivipams georgianus (I. Lea, 1834) Pleurocera acuta (Rafinesque, 1820)'' Amnicola limosus (Say, 1817) Pijrgulopsis lustrica (Pilsbry, 1890) Pomatiopsis cincinnatiensis (I. Lea, 1840) Frobijthinella enwrginata (Kuster, 1852) Soinatogynis depressus (Tryon, 1862) Bithynia tentaculata (Linnaeus, 1758)' Valvata tricarinata (Say, 1817) Valvata sincera (Sav, 1824) Valvata lewisi (Currier, 1868) Valvata bicarinata (I. Lea, 1841) ' Fossaria obntssa. as used here includes F. modicelta, (Say, 1825), F. nistica (I. Lea, 1841) and may include F. parva (I. Lea, 1841). '•pleurocera acuta (Rafinesque, 1820) had invaded Oneida Lake via the Erie Canal system before Baker's 1915 work (Dazo, 1965). c Bithynia tentaculata was apparently invading at the time of collection. Baker noted high densities in the outlet (Oneida River) very few in the Lake proper 1 Q 5 8 a B 3* 5 o I. 8. I 5 ^^ I I Figures 2-5. Figure 2. Dominant mollusks (individuals 10"- cnr) in Lower South Bay, Oneida Lake, in 191.5-17 (Baker, 1918c). In this, and figures 3 and 4, species occurring at a density of less than 0.5 individuals 10"- cm- are not considered. Figure 3. Dominant mollusks (individuals 10- cm-) in Lower South Bay, Oneida Lake in 1967-68 (Harman and Forney, 1970). Figure 4. Dominant mollusks (incfividuals 10"- cm-) in Lower South Bay, Oneida Lake in 1992-95. Figure 5. Dominant mollusks (individuals m"-) in Lower South Bay, Oneida Lake in 1992-95. Species occurring at a density of less than 0.5 indi\iduals m"- are not considered. W. N. Harman, 2000 Page 123 Table 2. Molluscan species inxadins; Oneitla Lake followed by the approximate dates of iiitnufnctioii and origins. Tixxa Approximate dates of introduction Origin Bivalvia: Ileterodonta Dreissenti polyinoi-plta (Pallas. 1771) Bivalvia: Palaeoheterodonta Potamihis alatiis (Sav, 1817) Li'ptodcii fraailis (Rafinesque, 1820) Liffiinia recta (Laniark. 1819) Gastropoda: Caenogastropoda Bithi/nia tentaculata (Linnaeus, 1758) Pleiirocera acuta (Rafinesque, 1820) Elimia livescens (Menke, 1830) Eliriiin virainica (Say, 1817) Total number of species: 8 1986 Eurasia, Great Lakes, Oswego River 1920-1950 Interior basin, Erie Canal system 1920-1950 Interior basin, Erie Canal system 1920-1950 Interior basin, Erie Canal system 1910-1918 Eurasia, Great Lakes, Oswego River 1825-1915 Interior basin, Erie Canal system 1920-1950 Interior basin antl/or Hvidson river, Erie Canal system 1920-1950 Susquehanna Basin, through-valleys, Oswego Basin and/or Interior basin, Erie Canal system which had arrived via the Erie Canal from the west, was well established by the time Baker completed his studies (Dazo, 1965; Baker, 191Sc). When John Fomey and I (Harman and Fomey, 1970) collected in the 1960s, 35 taxa were found. Three gas- tropods, Elimia livescens (Menke, 1830), E. virginica (Say, 1817), and Plcuroccra acuta, all in the family Pleu- roceridae, were represented only bv dead specimens. Three pearly freshwater mussels ((Unionidae), Potamihis alatus (Say, 1817), Leptodca fragiUs (Rafinesque, 1820), Ligumia recta (Lamark, 1819)) had immigrated from the Interior Basin (Clarke and Berg, 1959; Dazo, 1965). Eli- mia virginica was introduced into the Oswego watershed before the turn of the century (DeKay, 1843; Beau- champ, 1888). It is assumed that this species crossed the divide between the Susquehanna and Oswego River wa- tersheds via through-valleys after small headwater lakes and wetlands on the divide were disturbed to provide water for the Erie Canal after its opening in 1825 (Har- man, 1970). It also could have moved eastward along the Erie Can;il from populations in the Interior Basin (Goodrich, 1942). Bithi/nia tentaculata, present in 1917, had become abundant by the 1960s. An average 10- cm- bottom sample in 1967 contained 4 individuals of B. ten- taculata (table 2, figure 3). Table 3 lists the taxa collected in 1915-17 (Baker, 1916a, b, c; 1918a, b, c, d) and in 1969 (Harman and Fomey, 1970) that were not collected in the 1992-95 survey. Only 23 species of mollusks were present in 1992-95, two of them, Ligumia recta, and Valvata sin- cera Say, 1824 (Valvatidae), represented by dead speci- mens only. That represents a decrease of species rich- ness of 42% since 1917. Collections made in 1996 and 1997 by faculty and students at the Cornell Biological Field Station on Oneida failed to find anv living unionid bivalves (E. L. MiUs, pers. comm.). Four of the species of mollusks found in 1993-95 were introductions. Of those, the population of B. tentaculata had declined greatly since the 1960s but was exceeded only by the exotic zebra mussel, Drcissena polijmorpha. This Eur- asian dreissenid was apparently carried through the Os- wego River and New York State Barge Canal svstem by commercial and/or recreational navigation from the Great Lakes, where it was found in 1988 (Nalepa and Schloesser, 1993). Like B. tentaculata, D. polymorpha was supposedly transported in ballast water of ocean- crossing ships coming from Europe. An average 10- cm- bottom sample in 1993 contained 12 zebra mussels (fig- ure 4). An average 1 m- sample collected at the same time and location included 6,708 specimens of D. polij- morpha, 60 of B. tentaculata, and 2 of Elliptio complan- ata (Lightfoot, 1786) (£. complanatus of Baker) (figure 5). The tremendous increase results from large colonies of D. poh/morpha initially associated with specimens of E. complanata, which were not encountered in the 10- cm- samples. These substrates are fine shifting sands practicallv devoid of zebra mussels, except where the comparatively widely disbursed unionids support their colonies. Harman (1968a, b) proposed that the introduced B. tentaculata was able to displace the local representatives of the North American family Pleuroceridae in eutrophic environments. These observations were based on the distributions of B. tentaculata, P. acuta, E. livescens (Goniobasis livescens of Baker) and E. virginica in the Oswego River drainage basin in central New York (Har- man and Berg, 1971). Like most caenogastropod snails of inland waters, the pleurocerids are periphyton grazers that rely on the benthic algal community as their main energy source. They typically inhabit waters supporting comparatively low populations ot phytoplankton (Dazo, 1965). Bithijnia tentaculata also is a grazer but in addi- tion it can use planktonic algae which it filters from the water on specialized ctenidia (Jorgensen, 1966) poten- tially giving it an advantage in eutrophic waters such as Oneida Lake. Observations in Oneida in the 1960s regarding the density of Bithi/nia on rocky shoreLnes, and on the ob- served behavior of pleurocerid snails associated with B. tentaculata in aquaria, substantiated these ideas. In Page 124 THE NAUTILUS, Vol. 114, No. 3 Table 3. Ta\a lost (not collected ali\'e since dates indicated) followed by assumed major contributing factors. Taxa Last date collected Contributing factors to decimation Bivalvia: Palaeoheterodonta Miir^iaritifcra margaritifcra (Linnaeus, 17.58) Elliptio complanata (Lightfoot, 1786) Alasmidonta undulata (Say, 1817) Pijganodon grandis (Say, 1829) Fijganodon cataracta (Say, 1817) Strophitus undidatus (Say, 1812) Leptodea fragilis (Rafinesque, 1820) Ligumia recta (Lamark, 1819) ViUosa iris (1. Lea, 1829) Lampsilis rndiata radiata (GmeUn, 1791) Actinonaias ligmentina (Lamark, 1819) Gastropoda: Basommatophora Plu/sella gyrina {Sax. 1821) Physelln hcterostrophn (Say, 1817) Li/mnaea stagnalis Linnaeus, 1758 Stagnicola cmarginata (Say, 1821) Stagnicola catescopium (Say, 1867) Acelln haldmani (W. G. Binney 1867) Qyranulus deflectus (Say, 1824) Promenetus exacuous (Say, 1821) Fenissia rivularis (Say 1817) Gastropoda: Caenogastropoda Campctoma decisum (Say, 1817) Pleurocera acuta (Rafinesque, 1820) Elimia livenscens (Menke, 18.30) Elimia virginica (Say, 1817) Pomatiopsis cincinnatiensis (L Lea, 1840) Pwbi/thineUa emarginata (Kuster, 1852) Somatogtjnis depnisstis (Tr)-on, 1862) Valvata sincera (Say, 1824) Valiata lewisi (Currier, 1868) Vakata bicnrinata (I. Lea, 1841) Total number of species: 30 1968 1995 1917 1995 1917 1917 1968 1968 1917 1993 1917 1968 1968 1917 1968 1968 1968 1968 1968 1917 1968 1917 1968 1917 1917 1968 1917 1917 Direct competition with Dreissena Direct competition with Dreissena Trophic alteration Direct competition with Dreissena Trophic alterations Trophic alterations Direct competition with Dreissena Direct competition with Dreissena Trophic alterations Direct competition with Dreissena polymorph Trophic alterations polymorpha polynwrpha ' polymotyha polymorfiha polymoiyha Habitat Habitat Habitat Habitat Habitat Habitat Habitat Habitat Habitat destniction, destniction, destniction, destruction, destniction, destruction, destruction, destniction, destniction. trophic trophic trophic trophic trophic trophic trophic trophic trophic alterations alterations alterations alterations alterations alterations alterations alterations alterations Habitat destniction Direct competition Direct competition Direct competition Habitat destruction Habitat destruction Habitat destniction Habitat destniction Habitat destniction Habitat destniction , trophic alterations with Bitht/nia tcntaculata with Bithynia tentacxdata with Bithynia tcntaculata , trophic alterations , trophic alterations , trophic alterations , trophic alterations , trophic alterations , trophic alterations ' Empy shells collected in 1968, not collected in 1915; assumed introduction between 1920 and 1950. aquaria enriched with planktonic algae, pleurocericLs ac- tively grazed over the substrate. Individuals of B. tcn- taculata remained in one spot, filtering algae from the water. Upon encountering specimens of B. tcntaculata, pleurocerids were rebuffed by a violent twisting of the shell of that filter-feeder, which invariably caused the grazing snails to retract into their shells. They remained in that position for several minutes before resuming grazing. In situations with high densities of B. tcntacu- lata, indixidual pleurocerids spent a large fraction of the time in defensive positions rather than feeding. In Onei- da Lake in the 1960s, euhttoral cobble substrates were, in many areas, a solid mass of B. tcntaculata (Harman and Forney, 1970). Under those conditions, all the spe- cies representing the family Pleuroceridae chsappeared from the lake (Wisenden and Bailey, 1995). Nowadays, decreasing algal populations and increased water clarity have occurred concurrently with irrupting populations of D. polymorpha, while populations of B. tcntaculata have diminished drastically. A large number of the pulmonate gastropod species not found in the recent sur^'ev (e.g., one species of the planorbid genus Gyraulus, three in the family Lymnae- idae, and two in the Physidae) are typical of eulittoral substrates and often associated with emergent vegeta- tion. It is unlikely that these pulmonates are completely absent from Oneida, but their absence in the recent sur- vey suggests drastic reductions in their populations cor- responding with losses of that habitat due to shorehne development. Most of the shoreline has been modified as a result of the construction of break'waters or rip-rap. Many of Baker's euhttoral sites appear to be buried un- der yards and beaches created bv back-fiUing, which, bv 1993-95, had destroyed considerable areas of emergent vegetation. Other species of gastropods that were not found in the 1990s include several small prosobranchs in the fam- ily Hydrobiidae, which were typically abundant on sub- merged plants and nearbv sub-httoral and deeper-water substrates. These and other gastropods that historically W. N. Harman, 2000 Page 125 have been present in abundance in Oneida Lake, were found in very low numbers in the recent survey. An ex- ception, Amnicola limo.su.s (Say, 1817) {Aiunicohi liinosa of Baker), occurs in high densities in inanv littoral and sub-littoral areas. Changes in httoral vegetation and as vet unrecognized trophic alterations associated at least indirecth' with colonization by D. pohjmorfiha may be affecting these organisms (Stewart and Haynes, 1994; Howells ct al.. 1996; Ricciardi ct al, 1997). ' Seven of the 12 species of freshwater mussels found in Oneida since 1915 had been lost by 1993, and no li\ing unionids ha\'e been obser\'ed since 1995 (table 3). These losses correspond to the introduction and estab- lishment of Drcissena pohjuunyha. The literature doc- umenting the loss of unionid species as a result of com- petition with D. pohjiiwiyha (Nalepa et al, 1991; Grif- fiths, 1993; Ricciarchrffl/., 1995; Schloesser ef a/., 1996) and observations of the interaction behA'een unionids and D. polyinorpha in Oneida (E. L. Mills, pers. comm.) indicate that a large percent of this loss can be assumed to be attributed to interactions with D. poJijmotyha. Since 1915-17, the endemic molluscan community in Oneida Lake has been greatly modified. Species richness has declined and the dominant faunal elements are now Eurasian in origin (figure 5). With the exception of re- cent changes since the introduction of zebra mussels, water qualit)- in Oneida Lake has been consistent over the years (E. L. Mills, pers. comm.). It is assumed the observed changes are a result of biotic interactions with introduced taxa and loss of habitat due to human mod- ifications of the euhttoral emdronment. LITERATURE CITED Baker, F. C. 1916a. The fresh-water Mollusca of Oneida Lake, New York. The Nautilus .30:.5-9. Baker, F. C. 1916b. The relation of mollusks to fish in Oneida Lake, New York. N.Y State College of Forestrv. Technical Publication No. 4., .366 p. Baker, F. C. 1916c. Description of a new varietv of Lampsilis from Oneida Lake ^A^th notes on the L. lutcola group. The Nautilus .30:74-77, Baker, F. C. 1918a. Further notes on the Mollusca of Oneida Lake, New York: the mollusks of Lower South Bay. The Nautilus 31:81-93. Baker, F. C. 1918b. Description of a new variety of fresh-water mussel from Oneida lake, N.Y. N.Y. (Appendix) State Col- lege of Forestry, Technical Fuhlication 9, 43(2):247-248. Baker, F C. 1918c. The productivit)' of invertebrate fish food on the bottom of Oneida Lake, with special reference to mollusks. N.Y. State College of Forestry, Technical Pub- lication 9, 43(2):l-2.33 + 235-264 (appendix). Baker, F C. 1918d. The relation of shellfish to fish in Oneida Lake, New York. New York State College of Forestry, Cir- cular 21, 17(27):l-34. Beauchamp, W. M. 1888. Notes on American shells. The Con- chologists Exchange 2:114-115. Berry, E. G. 1943. The Amnicolidae of Michigan: distribution, ecology, and taxonomy. Miscellaneous Publications. Mu- seum of Zoology, University of Michigan. 57:1-68. Clarke, A. H. and C. O. Berg. 1959. The freshwater mussels of central New York with an illustrated key to the species of northeastern North America. Cornell University Agri- cultural Experiment Station. Memoir .367:1-79. Dazo, B. C. 1965. The morphology and natural history of P/eu- rocera andn and Coniohasis livescens (Gastropoda: Ceri- thiacea: Pleuroceridae). Malacologia 3:1-80. DeKay J. E. 1843. Zoolog\' of New York. Part 5. Mollusca. Albany, 271 pp.. 40 pis. Goodrich, C. 1942. The Pleuroceridae of the Atlantic coastal plain. Museum of Zoolog\', Universit)' of Michigan, Oc- casional Papers 456:1-6. Griffiths, R, W 1993, Effects of zebra mussels (Drcissena poly- uioiylui) on tiie benthic fauna of Lake St. Clair In: T F Nalepa and D. W, Schlosser (eds.) Zebra mussels: biology, nnpacts, and control. Lewis Pubhshers, Boca Raton, pp. 414-437. Harman, W. N. 1968a. Replacement of pleurocerids by Bi- thynin in polluted waters of central New York. The Nau- tilus 81:77-83. Harman, W. N. 1968b. Interspecific competition between Bi- thynia and pleurocerids. The Nautilus 82:72-73. Harman, VV. N. 1970. Alterations in the molluscan fauna of a meromictic, marl lake. The Nautilus 84:21-.30. Harman, W. N. and C. O. Berg. 1971. The freshwater snails of Central New York with illustrated keys to the Genera and Species. Cornell Universitv Agricultural Ex-periment Station. Search; Agriculture Entomolog)- (Ithaca) 2, 1(4): 1-68. Harman, W. N. and ]. L. Forney 1970. Fifty years of change in the molluscan fauna of Oneida Lake, New York. Lim- nology and Oceanography 15:454—460. Howells, E. T, M. H. Christopher, R. W. Bilyea, P B. Kauss, and K. Somers. 1996. Changes in environmental condi- tions during Drcissena colonization of a monitoring station in eastern Lake Erie, Journal of Great Lakes Research 22; 744-756. Jorgensen, C. B. 1966. Biology of suspension feeding. Perga- mon, Oxford, 357 pp. Mitchell, M. J., E. L. Mills, N. Idrisi, and R. Michener 1996. Stable isotopes of nitrogen and carbon in an aquatic food web recently inxaded bv Drcissena polymorpha (Pallas). Canadian Journal of Fisheries and Aquatic Sciences 53: 144.5-1450. Mozley, A. 1954. An introduction to molluscan ecology. H. K. Lewis, London, 71 pp. Nalepa, T F, B. A. Manny J. C. Roth, S. C. Mosley and D. W. Schloesser 1991. Long-tenn decline in freshwater mussels (BivaKia: Unionidae) of the western basin of Lake Erie. Journal of Great Lakes Research 17(2):214-219. Nalepa, T F and D. W. Schloesser (eds.). 1993. Zebra mussels: Biology, impacts and control. Lewis Publishers, Boca Ra- ton, 810 pp. Ricciardi, A., F G. Whoriskev, and J. B, Rasmussen, 1995. Predicting the intensitv and impact of Drcissena infesta- tion on native unionid bivakes from Drcissena field den- sity. Canadian Journal of Fisheries and Aquatic Science 52:1449-1461. Ricciardi, A., F. G. Whoriskey, and J B. Rasmussen 1997 The role of the zebra mussel (Drcissena polymorpha) in struc- turing macroinvertebrate communities on hard substrata. Canadian Journal of Fisheries and Aquatic Sciences 54: 2596-2608. Robertson, I. C. S. and C. L. Blakeslee. 1948. The Mollusca Pare 126 THE NAUTILUS, Vol. 114, No. 3 of the Niagara frontier region. Bulletin of the Bulletin of the Buffalo Society of Natural Sciences 19(3):1-191. Schloesser, D. W., T. R Nalepa, and G. I. Maclde. 1996. Zebra mussel infestation of unionid bivalves (Unionidae) in North America. American Zoologist .36:.300-310. Stewart, T. W and J. M. Haynes. 1994. Benthic macroinver- tebrate communities of southwestern Lake Ontario fol- lowing invasion of Dreissena. Journal of Great Lakes Re- search 20:479-493. Turgeon, D. D., ]. F. Quinn, Jr, A. E. Bogan, E. V. Coan, F. G. Hochberg, VV. G. Lyons. P. M. Mikkelsen, R. ]. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A. Scheltema. M. Vecchione, and ]. D. Wilhams. 199S. Common and sci- entific names of aquatic invertebrates from the United States and Canada: MoUusks. 2'"' edition. American Fish- eries Society Special Publication 26, .526 pp. Wisenden, R A. and R. C. Bailey. 199.5. Development of mac- roinvertebrate community structure associated with zebra mussel {Dreissena pohjmorpha) colonization of artificial substrates. Canadian Journal of Zoology 73:14.38-1443. 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POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 1580 SanibelFL 33957 T H Et^N AU T I L U S CONTENTS Volume 114, Number 4 December 15, 2000 ISSN 0028-1344 Luiz Ricardo L. Simone Cuido Pastorino Pablo E. Penchaszadeh Richard E. Petit M. G. Harasewych Bruce A. Marshall Douglas G. Smith Martin Avery Snyder Crepiclula arg^entina (Gastropoda: Calyptraeidae), a new species from the littoral of Argentina 127 Three new species of the genus Merica ( Neogastropoda: Cancellariidae) from South Africa and the Philippines 142 Systematics of the genus Infundibuhiin Montfort, 1810 (Gastropoda: Trochidae) 149 On the taxonomic placement of Unio ochraceus Say, 1817 in the genus Ligiimia (Bivalvia: Unionidae) 155 Latints beckijae, a new species of Fasciolariidae (Neogastropoda) from Brazil 161 DEC ^ 8 2000 STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULATION 1 . Publication Title, THE NAUTILUS 2. Publication No., 0028- 1 344 3. Filing Date, August 29, 2000 4. Issue Frequency, Quarterly 5. 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Extent and Nature of Circulation A . Total Number of Copies B. Paid Circulation 1 . Paid/Requested Outside-County Mail Subscriptions 2. Paid In-County Subscriptions 3. Sales Through Dealers and Carriers, Street Vendors, Counter Sales, and Other Non-USPS Paid Distribution 4. Other Classes Mailed Through the USPS C. Total Paid and/or Requested Circulation D. Free Distribution by Mail 1. Outside-County 2. In-County 3. Other Classes Mailed Through the USPS E. Free Distribution Outside the Mail F. Total Free Distribution G. Total Distribution H. Copies not Distributed I. Total J. Percent Paid and/or Requested Circulation Average Single 1 2 months Issue 545 550 425 406 0 0 0 0 0 0 425 406 0 0 0 0 22 25 2 2 24 27 449 433 96 117 545 550 95% 94% THE NAUTILUS 114(4):127-141, 2000 Page 127 Crepidiila argentina (Gastropoda: Calyptraeidae), a new species from the littoral of Argentina Luiz Ricardo L. Simone Museu de Zoologia da Universidade de Sao Paulo Caixa Postal 42694 04299-970 Sao Paulo BRAZIL IrsinioneC&'usp.br Cuido Pastorino' Museo Argentino de Ciencias Naturales k\. Angel Gallardo 470, 3° piso, lab. 57 C1405 D]R Buenos Aires ARGENTINA r\pastor@criba.edu.ar Pablo E. Penchaszadeh FCEvN-UBA-CONICET Museo Argentino de Ciencias Naturales Av. Angel Gallardo 470, 3° piso, lab. 57 C1405 DJR Buenos Aires ARGENTINA pablop@mail.retina.ar ABSTRACT Crepidiila argentina. a new species of gastropod of the famil)' Calyptraeidae, is described from the littoral of Mar del Plata, proxince of Buenos Aires, Argentina. The new species is con- chologicalK similar to C. protea d'Orbignv from southern Bra- zil. Crepidula argentina has a larger, subcircular shell and low- er convexitv' than C. protea. Anatomical characters that allow differentiation from this latter species are; larger lateral shell muscle and very weak dorsal shell muscle; kidney proportion- ally smaller, with a characteristic arrangement of inner folds of dorsal lobe; presence of a renal vessel edging rectum and work- ing as adrectal sinus; connection between odontophore muscle pairs m7 and mil; longer salivary gland; four ducts to digestive gland in stomach (instead of two); distinctive arrangement of folds in inner surface of stomach between esophageal aperture and posterior pair of ducts to digestive gland; seminal vesicle of males broad and few coiled; penis and papilla long and nar- row; vaginal tube nmning closely attached to capsule gland. The reproductive biology is also distinctive. Crepidula argen- tina new species has a very well defined seasonal reproductive cycle. Females brood between 1 and 46 egg capsules per spawn. The average total number of embrv'os per spawn is 5600. The uncleaved, laid egg diameter is 170 (xm and the number of eggs per egg capsule is about 320. All eggs develop, there are no nurse eggs. Larvae hatch as planktotrophic veli- gers. The new species is compared with other members of the genus Crepidula from the southern Atlantic coast of South America. Additional key word.?: Crepidula protea, anatomy, reproduc- tion, littoral. INTRODUCTION Most species of moUusks described from the southern Atlantic coast of South America are based on specimens collected by nineteenth century expeditions. Crepidula species are no exception. Alcide d'Orbigny (1841) de- Author for correspondence. scribed Crepidula protea for the first time on his large work "Voyage dans I'Amerique Meridionale". This book was published in several parts in different dates. Ac- cording to Sherboni and Woodward (1901), Sherbom and Griffin (1934), and Keen (1966) the page (465) with the C. protea description (with no illustration) was pub- fished on 1841. A year later in the French edition ol the Atlas of R. de la Sagra's "Histoire ... de la File de Cuba", d'Orbigny pubfished a plate (pi. 24, figs. 30-32) with three illustrations of C. protea and no description (A description was published in 1846 in the Spanish edi- tion of the same work.) It seems clear that the intention of d'Orbigny was to publish the chapter of mollusks in Sagra's work before his 'Voyage ", because he cited that other book despite its later pubfication. Crepidula protea was described without mention of a distinct t\pe locality. A label at The Natural Histor)' Museum, London (BMNH) states only "Patagonia" without further geo- graphic details. The examination of the type specimens did not suggest any precise locafitv nor were there shell characters that could reveal which population they had came from. D'Orbigny's description and illustration of Crepidula protea clearly groups at least two different species. First Dall (1889), and later Hoagland (1977), pointed out that C. plana Say, 1822, and C. iin^uiformis Lamarck, 1822, might be included within his concept of C. protea. Ac- cording to d'Orbigny's illustrations and description this seems to be possible. Parodiz (1939) studied the species oi Crepidula from Argentina with particular emphasis on the shell and, for the first time, on the radula. Based on a large number of lots from different localities Parodiz described and illustrated the radula and shell of five previously known species and two new subspecies. The subspecies are now considered to be geographical forms. Despite that, Par- odiz (1939) is still the most authoritative paper on the genus Crcpidida from southern Atlantic coast of South Page 128 THE NAUTILUS, Vol. 114, No. 4 America. Unfortunately, no anatomical descriptions or reproductive features were described in that work. The specimens examined by Parodiz are stiU housed at the Museo Argentino de Ciencias Naturales (MACN) and were re-examined in this paper Since Parodiz's pio- neering work no further articles on Crepidula from Ar- gentina have been published. Hoagland (1977) published a comprehensive study of living and fossO species of Crepidula. Although her work was basically restricted to North America and the east- ern Pacific, she included descriptions and pictures of C. protea and C. dilatata as well as a few other southern species. Hoagland (1983) described several specimens from BrazU of what she concluded was C protea. She recognized C. protea as a species distinct from C. plana and C. ungitifonni.s and studied d'Orbigny's type mate- rial of C. protea. In the same paper, Hoagland described the larval development of C. protea based on specimens from the southeastern Brazihan coast. Here we consider the specimens she studied to represent actual C. protea mainly because they are conchologically similar to the lectotype and paralectotypes. In an annotated Ust of several type specimens from the coast of Argentina, Aguirre (1993) designated a lec- totype for C. protea. Unfortunately, she selected the only complete specimen in d'Orbigny's collection at BMNH. She seems to have been unaware of the differences among the southern species of this genus. Aguirre (1993) mentioned only 4 syn types when the count is ac- tually 16, arranged in two lots of 5 and 11 specimens respectivelv (BMNH 1854.12.4.573 and 574) (Hoagland, 1983). Gallardo (1977, 1979) and Brown and Olivares (1996) described several new species of Crepidula from Chile with distinctive reproductive patterns as diagnostic char- acters. These authors led the way in the search for new characters in this variable genus, showing that in several cases the shell lacks real ta.\onomic value at the specific level. Hoagland (1983) and Rios (1985) pointed out that Crepidula protea occurs from Rio de Janeiro, Brazil, south to Miramar, in the province of Buenos Aires. How- ever, studies on different population samples along this range revealed that more than one pattern of reproduc- tive strategy and anatomical characters are found, indi- cating the presence of an unnamed species. In this paper we describe this new species of Crepi- dula and provide the groundwork for a future revision of the group in the southwestern Atlantic. The new spe- cies was originally recognized first by its distinctive re- productive pattern and second through detailed anatom- ical studies. Comparison with several kiiown species from South America is presented. The study of repro- ductive biology in the new species is the main topic of another paper (Cledon and Penchaszadeh, submitted.). MATERIALS AND METHODS Fresh specimens of Crepidula argentina new species used in this studv were collected by commercial fish trawlers at Mar del Plata (38°00' S, 57°33' "W) and Puerto Quequen (38°35' S, 58°42' 'W) along the coast of the province of Buenos Aires, Argentina. Tvpes and material examined are deposited at: Museu Oceanograf- ico de Rio Grande, Rio Grande do Sul, Brazil (FURG); Museo Argentino de Ciencias Naturales "Bernardino Ri- vadavia", Buenos Aires (MACN); Museo Nacional de Historia Natural, Montevideo, Uruguay (MHNM); Mu- seu de Zoologia, Universidade de Sao Paulo (MZSP); National Museum of Natural History, Smithsonian In- stitution, Washington (USNM). The type material of C. protea was studied at The Natural History Museum, London (BMNH). Specimens were fixed in fomiahn for at least 24 hours and preserved in 70% ethanol. They were not relaxed. All observations were made on preserved material. Dis- sections were perfonned on 3 males and 13 females us- ing standard techniques under a stereomicroscope with the specimens immersed on 70% ethanol. Radulae were prepared (two specimens of C argentina new species of 32.2 and 35 mm length and two of C. protea of 18.5 and 17.4 mm) according to the method described by Solem (1972) and observed and illustrated using a LEO 440 scanning electron microscope (SEM) at the National Museum oi Natural History, Washington. Shell measurements including convexity index, were done using the parameters established by Hoagland (1977). Most photographs were taken using a Nikon N70 with a 60 mm Nikkor macro lens. These images were latter scanned from black and white 35 mm negatives using a Nikon Coolscan III slide scanner All images and plates were processed with the software Photoshop 5.02. In the figures and text, the following abbreviations are used: aa, anterior aorta; ab, auricle region beyond ven- tricle connection; ac, anterior extremity of gill on mantle border; ag, albumen gland; an, anus; au, auricle; bb, bulged part of br; bg, buccal ganglion; br, subradular membrane; bv, mantle blood vessel inserting in kidney; eg, capsule gland; cv, ctenidial vein; da, aperture of duct to digestive gland; dd, duct to digestive gland; dg, di- gestive gland; dm, dorsal shell muscle; dp, posterior duct to digestive gland; ea, esophageal aperture; en, en- dostv'le; es, esophagus; fd, dorsal surface of foot; fg, food groove; fl, female papilla; fp, female pore; ft, foot; gd, gono-pericardial duct; gi, gill; gs, gastric shield; bg, hypobranchial gland; in, intestine; ir, insertion ol m4 in tissue on radula preceding its exposed (in use) portion; iu, U-shaped loop of intestine on palhal roof; kd, dorsal lobe of kidney; Id, kidney; kv, ventral lobe of kidney attached to intestine; 11, left lateral ex-pansion (Hap) of neck; hn, lateral membrane restricting paUial cavity; ml to ml4, odontophore muscles; mb, mantle border; mj, muscles of jaws and mouth; ml, mantle region restricting palhal cavity; mo, mouth; mr, mantle reinforcement; mt, mantle; ne, nephrostome; nr, nerve ring; ns, neck "sole"; oc, odontophore cartilage; os, osphradium; ov, paUial oviduct; oy, ovary; pc, pericardium; pd, penis sperm groove; pe, penis; pg, pedal gland anterior fur- row; pp, penis papilla; pr, propodiuni; pt, palfial sperm L. R. L. Simone et ai, 2000 Page 129 Table 1. Shell morphometries of Crepidula argentina new species. Linear measurements in mm. Speci- Septum Convex- men Total Diam- Septum free ity # length Height Width eter length area D/L 1 31.5 6.3 25 37 13.4 17.9 1.17 2 35.7 7.7 24.7 41 15.1 19.8 1.14 3 32.8 8.6 23.1 39 17.6 17.5 4.18 4 35.8 8.9 25 42 17 19.4 1.17 5 33.8 8.8 27 43 13.7 21.8 1.27 6 28.1 7.1 21.7 36 11.8 16.4 1.28 ( 32.6 7.3 22.5 39 13.3 17.6 1.23 8 30 7.3 23.4 37 13.3 16.8 1.14 9 29.8 7.1 23.9 34 12.3 16.3 1.14 10 33 9.3 28 41 14.4 19.3 1.24 groove; ra, radiila; rl, right lateral expansion (flap) of neck; m, radular nucleus; rs, radular sac; sc, subradular cartilage; sg, salivary gland; sn, snout-proboscis; sp, ap- erture of vas deferens into pallial cavity; ss, style sac; st, stomach; sv, seminal vesicle; te, cephalic tentacle; tm, net of transversal muscles of haemocoel; to, tissue cov- ering middle region of radula before its exposed part; ts, testis, vc, visceral connection with haemocoel; ve, ventricle; vg, vaginal duct; vm, visceral mass; vo, visceral oviduct; vs, vesicles of pallial oviduct. SYSTEMATICS Class Gastropoda Cuvier, 1797 Subclass Orthogastropoda Ponder and Lindberg, 1996 Superorder Caenogastropoda Cox, 1960 Order Sorbeoconcha Ponder and Lindberg, 1997 Family Calyptraeidae Lamarck, 1809 Genus Crepidula Lamarck, 1799 Crepidula argentina new species Figures 1-21, 28^6 Crepidula protea: Parodiz, 1939: 702, fig. 8, pi. 1, fig. 6 (not d'Orbigny, 1841). Diagnosis: Shell: Large and wide, slightly convex. Pro- toconch smooth, with 1% whorls. Aperture elhptical. Beak solid, very small. Septum planar, septum margin with sulcus on left side and clear notch in center. Muscle scars absent. Shell externally opaque white, internally white porcelanaceus. Periostracum absent. Anatomij: Large lateral shell muscle. Dorsal shell mus- cle weak. Kidney proportionally smaU, with characteristic arrangement of inner folds of dorsal lobe, renal vessel edging rectum. Connection between odontophore mus- cle pairs m7 and mil. Long salivary glands. Four gastric ducts to digestive gland, distinctive arrangement of folds (sorting area) in inner surface of stomach between esophageal aperture and posterior pair of ducts to di- gestive gland. Seminal vesicle broad and few coiled. Pe- nis and its apical papilla long and narrow. Vaginal tube running attached to capsule gland. Description: Shell (Figures 1-15, 20, 21): Large (up to 40 nun in length) and wide, thin (0.35—0.45 mm in thickness), slightly convex, convexity = 1.20 (see table 1 for other measurements), male specimens very thin, brittle, transparent, planar shells. Protoconch smooth, with 1% whorls, transition to teleoconch clearly defined. Aperture elliptical or subcircular. Beak .soUd, very small, turned to right on females, almost central on males, at level of or only slightly above margin, in males never reaching margin, in females never extended beyond it. Septum planar (never convex), with central ridge almost imperceptible but present, margin of septum with sulcus on left side and clear notch in center, covering less than half of aperture, color bright white with translucent edge. Muscle scars absent. Growth lines covering entire shell. Shell externally opaque-white, internally porcehiin- white; some specimens externally with diffuse radial or- ange lines and/or internally with radial brownish lines. Males always translucent-white externally and bright- white internally. Periostracum absent. Head-foot (Figures 28, 30, 34, 43): Head protruded, at the end of long (about same length as foot), dorso-ven- trally flattened, neck region. Snout short and cylindrical, able to retract and partially invaginate for about half of its length within haemocoelic cavity. Tentacles long, stubby, tip somewhat rounded. Eyes dark, situated on low ommatophores about midway on lateral margin of tentacles. Neck region with pair of lateral, flattened ex- pansions (nuchal lobes); right nuchal lobe bears shallow food groove along its limit with head (figure 28). Sperm groove of males (described below) running externally along food groove (figure 43). Ventral surface of neck region forming an additional, anterior "sole" (figure 30). Foot veiy ample (occupies about % of shell aperture), dorso-ventrally flattened. Shell septum defining dorsal limit of foot. Mantle fusing with dorsal surface of foot and protruding beyond its borders. Furrow of pedal glands transversal, located on central region of anterior margin of foot (about 1/3 of its width); a small, pointed expansion present on each side of furrow (figure 30). Anterior margin of foot covering ventrally posterior re- gion of neck "sole". Columellar muscle very reduced, small flap contouring anterior border of shell septum, only evident on right side. Inner haemocoelic cavity nar- row, running approximately along center of neck region, almost entirely filled by mass of salivary glands (de- scribed below) and numerous, transversal, very slender muscle fibers (figure 34); these fibers connect ventral surface of dorsal haemocoelic wall with dorsal surface of its ventral wall. Mantle organs (Figures 29, 31-33, 44): Mantle border very thick, edging entire ventral inargin of sheU, free on its anterior half and attached to foot edge on its posterior half. Mantle border without appendages, but entirely covered by a series of minute glands. Mantle border with Page 130 THE NAUTILUS, Vol. 114, No. 4 Figures 1-15. Shells of Crepidula argentina new species. 1-3. Holotvpe, MACN .3450S, Mar del Plata, Buenos Aires, Argentina. 4-5. Paratope, MACN .34.509, Mar del Plata, Buenos Aires, Argentina. 6-7. Paratope, MACN .34509. Puerto Quequen. Buenos Aires, Argentina. 8-9. Paratype. 10-11. Paratype, MACN .34.509, male specimen, scale bar on right side of figure 11 = 0.5 mm (for figures 10 and 11). 12-13. Paratjpe, MACN .34509 14-15. Parat>pe, MACN .34509. Scale bar under figure 8 = 1 cm for all specimens, except figures 10 and 11. L. R. L. Simone ci ai, 2()()() Page 131 Figures 16-21. Crepidula argentina new species. 16-17. Radula 16. Radula, frontal view. Scale bar = 30 \x.m. 17. Radula, marginal teeth, scale bar = 30 jxni. 18-19. Penis, critical-point dried. 18. Dorsal view. Scale bar = 200 (xni. 19. Ventral \iew. Scale bar = 200 \Lm. 20-21. Protoconch. 20. Dorsal view. Scale bar = 300 ^.m. 21. Detail of figure 20. Scale bar = 100 p-ni. characteristic arrangement of folds along central region of aperture of pallial cavity (figure 33), a broad furrow beginning on anterior extremity of gill, running toward left side, and ending at about right third of osphradium; this thick fold presents broad central furrow. Dorsal shell muscle apparently lacking in most specimens, very re- duced and difficult to observe in others (figure 29). Lat- eral shell muscle well developed, inserting broadly in left lateral region of mande border and region posterior to it, originating shortly in inner shell surface in region close to left anterior edge of shell septum. Pallial cavity aperture occupving about 2/3 of anterior half of shell border, turned to right (drawing an analogy between shell in dorsal view and a clock, with head occupying 12:00, pallial aperture occupies a sector beginning at 9:00 and ending at 2:00). Pallial cavity deep, broad, tri- angular, arched and flattened dorso-ventrally (figure 29). Anterior extremity of pallial cavity slightly larger than its aperture due to constriction on left and right extremities produced by fusion of mantle and foot (figures 31, 44). Pallial ca\ity narrows gradually in posterior direction, penetrating left side of visceral mass (described below); cavity length about 3/4 of total length of animal. Os- phradium (figure 33) long, monopectinate, located be- tween anterior region of gill and mantle border, occu- pying about middle region of pallial aperture, somewhat perpenthcular to longitudinal axis of animal, compressed between gill and mantle border. Osphradium length slightlv exceeding 1/5 of palfial aperture length. Osphra- dium leaflets long, somewhat thick, closely packed, with rounded tip. Gill very large, with somewhat narrow base, edging anterior and left margin oi pallial cavity along almost its entire length; anterior extremity of gill in an- terior and left region of pallial cavity aperture, near its Page 132 THE NAUTILUS, Vol. 114, No. 4 Figures 22-27. Crepidula protea dOrbigiiy, 1841. 22-24. Shell, MACN 34511, off Ubatuba, Sao Paulo, Brazil, 2.3°30' S, 44°54' W, 42 m, dorsal, ventral, and lateral \iews of the same specimen. 25. Shell, MACN 34511, other specimen in lot, ventral view. Scale bar = 1 cm for all shells. 26. Radula, frontal view. Scale bar = 100 |j.m. 27. Protoconch, dorsal view. Scale bar = 300 (Am. right limit, on thick mantle border; posterior extremity of gill in posterior end of palhal cavaty (figure 32). Base of gill filaments triangular. Gill filament with very long, almost straight, stiff rod extending to right; rods extend for about twice as long as their triangular, membranous base; these rods begin in region of ctenidial, in left mar- gin of cavity roof, touching food groove of head-foot, in right margin of cavit)' floor Rod apex rounded and pre- ceded by a thicker region. Gill filaments connected to each other by cUia, mainly along their thicker apical re- gion, which helps maintain somewhat gill structure. Gill filaments longer in central region of gill, shortening gradually toward both extremities. Anterior extremity of gill with short filaments, abruptly turning foiAvards, end- ing at mantle border (figure 33). Ctenitliiil vein cylin- drical. Endostyle well developed, yellowish (a somewhat narrow glandular ridge located on ventral surface of ctenidial vein and present along its entire length) (figure 33). Hvpobranchial gland whitish, low, slightly devel- oped, occupying surface between gill and visceral mass (figures 1, 5). About 1/3 of visceral mass encroaches on pallial cavity roof, occupying about 1/3 of posterior and right sectors of this region; pericardium and kidney lo- cated in posterior part; long intestinal loop, anus, and palhal oviduct in anterior part (described below). Visceral mms (Fiffircs 2.9, 31, 32, 44): A dorso-ventrally flattened cone lying in shell chamber produced by sep- tum. Thin calcareous septum separating visceral mass from dorsal surface of foot. Left and anterior region of L. R. L. Simone ct at., 2()()() Page 133 30 31 Figures 28-31. Anatomy of Crepklula argentina new species. 28. Female animal with shell, visceral mass, and pallia! cavity- removed, dorsal view, 29. Same animal, whole, dorsal view. 30. Same, ventral view. 31. Isolated visceral mass and pallial cavity, ventral view. Scale bar = 5 mm. visceral ma.ss occupied by palLal cavity (figures 31, 44). Remaining region of visceral mass with stomach as central structure, immetliately and iilmost completely surrounded by digestive gland (except in some ventral and dorsal parts). Gonad surrounding digestive gland externally. Visceral mass encroaching on right and posterior re- gions of palli;il cavitv roof. Anterior extremity of visceral mass (ventral to posterior pallial cavity region) covering columellar muscle just posterior to anterior border of shell septum. Page 134 THE NAUTILUS, Vol. 114, No. 4 L. R. L. Simone et a/., 2000 Page 135 Circidatani and cxcrctoni si/.stciii.s (Figures 29, 35); Peric-archiini \en loiiij, soiiiewhat peqieiidicular to loTi- gitialinal avis ot animal (figure 29); verv' narrow in pos- terior extreniit)' of gill (posterior and left end of pallial cavity); running along anterior margin of xasteral mass where it encroaches on piillial roof, enlarging gradually and ending at about middle level of this region of vis- ceral mass, near its mechan line. Pericartlium circum- scribes (1) palhal cavit)' in anterior and ventral direction, (2) visceral mass (mostlv gonad) in posterior direction, (3) mantle in dorsal direction, and (4) kidnev to right. Auricle thin-walled and vew long, running long entire pericardium length, attached to its anterior and dorsal inner surfaces; auricle connects with ventricle approxi- matelv along its middle and right thirds; auricle portion beyond connection with ventricle represented by broad binil sac. Ventricle elliptical, veiT muscular, its connec- tion with auricle located about middle region of its an- terior surface; aortas originating in opposite side. Ante- rior aorta broad, directed awav from posterior aorta. An- terior aorta directed toward right, edging posterior inner surface of pericardium. Anterior aorta penetrates head haemocoel. Kidney small, occupying about 1/4 of area of visceral mass when projected on inner surface of pal- lial cavity. Kidnev circumscribes (1) mantle in dorsal ch- rection, (2) pallial ca\ity in ventral and left lateral direc- tion, (3) visceral mass (mostly gonad) in posterior and right direction, (4) pericardium in posterior and left di- rection, (5) an intestinal loop in anterior direction, (6) intestine and oviduct (when present) in lateral right di- rection. Kidney mostly hollow, with pair of veiy irregu- larly shaped lobes. Ventral lobe with several, slight nar- row transverse folds attached to posterior surface of ad- jacent intestine. Dorsal lobe occupies most of dorsal and lateral surfaces, bearing several irregular folds in left di- rection; part of lobe covering ventral surface around nephrostome. Nephridial gland thin, present along boundary between kidney and pericardium, bearing se- ries of transverse, narrow folds connected with dorsal renal lobe. Nephrostome a very small sht in left region of ventral wall. A broad vessel connected with left ex- tremity of kidney and ninning along external pallial loop of intestine for about % of its length. Digestive system (Figures 29, 31, 32, 34-42): Mouth longitudinal, in center of anterior surface of snout (figure 34). Buccal mass very large, occupying most of inner space of snout. Buccal mass capable of some protraction and invagination. Dorsal wall of buccal mass with pair of broad and thin jaw plates. Pair of df)rsal folds broad and low, connected to jaws in posterior direction. Odon- tophore large, consisting of most of volume of buccal mass. Odontophore muscles (figures 36-38, 40, 41): (ml) jugai muscles, several verv narrow muscles con- necting buccal ma.ss to adjacent wall of snout, more con- centrated anteriorly around mouth; (mla) pair of dorsal protractor muscles, narrow, thin and superficial, with or- igin in antcro-dorsal region of mouth, close to its median line, insertion in posterio-dorsal and lateral region of odontophore; (mj) jaws and peribuccal muscles, some- what thick, surrounding lateral and dorsal wall of buccal mass, with origin around mouth, insertion in middle re- gion of lateral and dorsal wall of odontophore; (m2) pair of retractor muscles of buccal mass (retractor of phar- ynx), broad, with origin in latero-ventral region of hae- mocoel just posterior to snout, nmning in anterior di- rection, with insertion in postero-Iateral and dorsal re- gion of odontophore cartilages; (m2a) pair of dorsal ten- sor muscles of radula, continuation of m2 after insertion in cartilages, nmning in anterit)r direction, with insertion in subradular cartilage in middle region of its dorsal in- ner surface; (mt) dorsal transversal muscle or ventral ap- proximator muscle of cartilages, connecting dorsally pos- terior-dorsal and lateral surface of both cartilages, lying between superficial membrane that covers odontophore and tissue on middle region of radula (to); (m4) pair of median dorsal tensor muscles of radula, very large and thick, with origin in ventral-central and posterior region of odontophore cartilages, running along their middle region, contouring meso-ventral surface of cartilages, running along their dorsal surface, with insertion in dor- sal-posterior and medial extremity of subradular carti- lage; (mSf) pair of median radular tensor muscles, thick, with origin in meso-posterior and dorsal regions of odon- tophore cartilages, just adjacent to m2 insertion and m2a origin, crossing middle region of m4, running toward medial region of m4, with insertion along radular sac on both sides (each branch of m5 nms along a side of rad- ular sac, medially and dorsally); (m6) horizontal muscle, very' thin, uniting anterior half of odontophore cartilages, with insertion on their dorsal margin; (m7) pair of ven- tral tensor muscles of radula, thin and narrow, with or- igin in meso-anterior margin of m4, running in posterior direction adjacent to subradular membrane, bifurcating in posterior region of odontophore, median branch con- necting with that of other member of pair and inserted in meso-posterior region of radular sac, lateral branch connecting with mil; (m8) pair of strong muscles with origin in postero-dorsal and lateral region of odonto- phore cartilages just adjacent to insertion of m2, nmning along and attached to dorsal margin of odontophore car- Figures 32-36. Anatomy oi Crepidula ar^entina new species. 32. Pallial cavity, ventral view, part of visceral mass deflected, some gill filaments of central part of gill removed. 33. Detail of mantle border and pallial cavity at anterior extremity of gill. 34. Head and haemocoel, ventral view, foot and neck "sole" removed. 35. Detail of visceral mass at border of pallial cavit)-, adjacent to pericardial structures, ventral view, ventral surface of pericardium and kidney membranes removed. 36. Buccal mass, dorsal view. All scale bars = 1 nun. Page 136 THE NAUTILUS, Vol. 114, No. 4 mo ra to \ 37 mil / . m rnl4 40 L. R. L. Simone et al., 2000 Page 137 tilages, with insfrtioii in their antero-tlorsal region ad- jacent to horizontal muscle (m6); (ni9) pair of dorso- medial tensor muscle of radula, broad and thin, with origin along ineso-dorsal surface of radular sac (in its region internal to odontophore), crossiTig to dorsal sur- face, with insertion in dorso-ventral margin of suhradn- lar cartilage; (mil) small, narrow pair of muscles, with origin in meso-ventnd region of mouth, nmning in pos- terior direction along its median line, penetrating in me- dian region of odontophore, rumiing between m7 and m4, with insertion in anterior region of subradular mem- brane; (ml4) pair of broad and thin muscles, with origin in postero-dorsal region of otlontophore, close to origins of m2 and m5, nmning in antero-ventral direction, with insertion in inner ventral surface of snout at about mid- dle region of odontophore; to) tissue covering mitldle region of radida within odontophore, along its dorsal sur- face; br) subradular membrane, covering ventral surface of subradular cartilage and some neighboring areas. Radula taenioglossate, short, measuring little more than odontophore length (figures 37, 38). Rachidian tooth tall, narrow with cin^ved, convex base, central cusp ver) large and shaqi, at least two weak denticles on each side decreasing in size toward lateral teedi, no basal cusps but pair of lateral reinforcements present along borders. Lateral tooth broad (about 3 times rachidian width), curved inward, with conspicuous apical cusp turned toward rachidian line and 5-7 short, triangular denticles along edge on marginal side and 2—3 very weak denticles on edge on rachidian side, denticles de- creasing in size in both directions, disappearing at about middle region of tooth, only a thickened border remain- ing. Marginal teeth long, curved, tall, tip sharply pointed, with serrate inner margin (at least 7 denticles); inner marginal tooth broad, about twice as wide as outer mar- ginal tooth. Pair of buccal ganglia large, close together near me- dian line (figure 37), situated between buccal mass and adjacent esophagus. Salivary glands very long, tubular, coiled (about 3 times longer than haemocoel length when straightened) (figure 34). Several narrow trans- verse muscles unite internally dorsal and ventral surfaces of haemocoel, passing both sides of salivary glands, esophagus, and aorta (figure 34). Salivary glands not passing through nerve ring. Ducts of salivary glands thick, running to dorsal surface of buccal mass, pene- trating adjacent wall of buccal mass; apertures small, close to site of penetration, located in anterior region of dorsal folds of buccal mass (figure 36). Esophagus narrow and long (figure 39). Inner surface of anterior esophagus with a pair of broad folds. Middle esophagus with pair of narrow folds (continuation from those of anterior esophagus) and moderately spacious glandular chamber. Iimer surface of posterior esophagus with 4—5 longitudinal, narrow, folds of similar (hmen- sions. Stomach (figures .39, 42) large, slightly conical, oc- cupying about half of visceral mass; esophagus inserting in left side of its po.sterior region of stomach, adjacent to region of shell apex. Four ducts to digestive gland present: anterior pair narrow, inserted in ventral surface, between insertion of esophagus and posterior gastric end, one turned in anterior direction and other to op- posite side; posterior pair of ducts thick, located in mid- dle region of ventral and left surface of stomach, one at considerable distance from other. Stomach graduallv narrows in anterior and left direction, close to left and posterior extremity of paUial cavity. Stomach inner sur- face (figure 42) with p;ur of narrow and long folds, both with origin adjacent to insertion of esophagus, lining posterior surface of stomach, gradually disappearing in direction posterior to apertures of anterior ducts to di- gestive gland. Gastric shield thin, presenting transversal folds marking a somewhat elfiptical, ccjucave area. An- terior half of stomach with a pair of slight tall, longitu- dinal folds; posterior region between both folds smaller than anterior region; smaller region as intestinal branch of stomach; broader (anterior) region as style sac. Di- gestive gland pale-brown in color, surrounding stomach except for some spots in dorsal and ventral surfaces. Intestine very narrow and sinuous (figure 39); running from left to right adjacent to anterior border of stomach, running up near median fine to right and anterior ex- tremity of visceral mass; in this region and toward left, it surrounds right and anterior border of kidney, turning suddenly to right and nmning pandlel to preceding loop; both loops exposed along pallial roof (figures 29, 32, 35, 39). Anus small, sfightly siphoned, situated in right re- gion of pallial cavity close to mande border (figures 31, 32, 39). Last loops of intestine contiiin numerous, small, somewhat elliptical fecal pellets. Male genital sifstem {Figures 43. 44, 46): Only small specimens (up to 9 mm) are functional males. Testis sit- uated in anterior region of visceral mass, color orange. Seminal vesicle convoluted, very thickened, color cream, situated in anterior and right region of visceral mass, where it abruptly narrows, becoming very slender and opening in postero-ventral and right region of pallial cav- itv. A shallow groove runs from this aperture to penis liase, on pallial floor near right margin of head. Sperm groove better defined and deeper in anterior direction. Penis long (about 3 times tentacle length), curved, orig- Figures 37-41. Anatomy oi Crcpidiila nruentina new species. 37. Buc'c;il mass, ventral \iew. 38. Odontophore, dorsal view, some muscles sectioned and deflected, part of right region of subradular curtilage remo\ed to show muscular insertions in it. 39. Digestive tubes seen in situ with visceral mass as a transparent structure, ventral view. 40. Odontophore, ventral view, superficial membrane and muscles removed, nght mj (left in figure) also removed. 4L Odontophore, ventral view, radular sac deflected and only partially shown, odontophore cartilages deflected from each other, right m4 (left in figure) deflected downward. All scale bars = 1 mm. Page 138 THE NAUTILUS, Vol. 114, No. 4 L. R. L. Sinione et al, 2000 Page 139 inating dorsallv to right tentacle. Papilla on penis tip, very long, about half of penis length. Penis duct opened, running along middle region of ventral surface of penis to tip of papilla. Female genital st/stem (Figures 29, 31, 32, 35, 45): Ovarv pale brown, surrounding digestive gland, denser in anterior region of visceral mass (figures 29, 31). Vis- ceral oviduct \er\' narrow, ninning from left to right in anterior border of \asceral mass. Gonopericardial duct well developed, slightlv thicker than visceral oviduct, with origin in ventral and right extremitv of pericardium (figure 35); nmning along visceral glands encroached in pallial cavity; inserted in posterior extremity of pallial oviduct where it joins insertion of visceral oviduct. Al- bumen gland long, moderately thick, whitish; walls thick, glandular; situated in anterior and right extremity of vis- ceral mass; 3 to 4 seminal receptacles inserted along right surface of albumen gland. Capsule gland a contin- uation of albumen gland, marked by sudden increase of secretorv tissue in wall and bv turn toward left; walls irregular, thick, glandular. Vaginal tube moderately nar- row, originating in posterior region of capsule gland, run- ning attached to this latter up to its right limit, where it abniptlv turns in ventral direction to form tall genital papilla. Papilla with p;ur of low folds iimning along its posterior side, from base almost to tip; folds close to each other with narrow furrow in between. Female gen- ital pore situated in tip of papilla (figures 31, 32, 45) sfit- hke, transversal, with posterior and anterior edges slight- ly projected. Etymology: The specific epithet, a noun in apposition, refers to the name of the countiy where the species oc- curs, Argentina. Type locality: Mar del Plata, Buenos Aires, Argentina in 35-50 m, on shells of Mytiliis cdiilis platcnsis d'Orbigny, 1846. Type material: Holotyjie, MACN 34508; 20 paratv- pes, MACN .34509 (10 dn,' specimens) and MACN 34510 (10 specimens preserved in ethanol); 6 paratypes, MLP 5578; 16 paratypes (3 males, 13 females) MZSP 32152; 6 paratypes USNM 2016009 (1 specimen pre- served in ethanol); MHNM, 6 specimens without num- ber; 6 paratypes MHNM 15105; all from type locality. Additional material examined: MACN 18504, 39 specimens, Puerto Quequen, Buenos Aires; MACN 8887, 9 specimens, off Mar del Plata, in 46 m; MACN 11367, 36 specimens; MACN 9361^9, 5 specimens; MACN 18374, 2 specimens; MACN 8653, 4 specimens; MACN 11586, 27 specimens; all from Mar del Plata; MACN 20529-1 more than 60 specimens, mouth of Rio Negro. Literature record.s: Due to similarities with C pro- tea, most hteratnre records should be re-checked from now on. Records cited by Parodiz (1939) were revised and are included in the material examined. Di.stribution: Province of Buenos Aires, Argentina, from Mar del Plata to the mouth of Rio Negro, on banks of Mytiltis edulis phiten.sis, 35-50 m depth. Records from Uiiiguav and Brazil need to be confirmed. Reproductive pattern: Crepidula argentina new spe- cies is a protandrous hermaphrodite that undergoes complete sex change. Males mature at about 4 mm length. Sex change begins when individuals reach ap- proximately 9 mm and is completed when the first pre- vitelogenic ovocites appear, usually at around 11 mm length. The smallest broochng female we found was 15.0 mm length and the largest 39.0 mm. The new species has a well-defined seasonal repro- ductive cycle. Between September and March more than 30% of the females are brooding, with maximum brooding in January (57%). In June and July brooding is uncommon (0-10%). Females l)rood 1—46 egg capsules per spawn. The average number of embryos per spawn is 5600. There is no correlation between size of the fe- male and number of egg capsules or eggs per spawn. However, female size, capsule size, and the average number of embryos per capsule are positively correlated (Cledon and Penchaszadeh, submitted). Uncleaved egg diameter is 170 |xm and there are about 320 eggs per egg capsule. All eggs develop, there are no nurse eggs, and the onlv extra-embi"vonic nutritit)nal source is the intracapsular liquid. There is no record of cannibalism in early or advanced stages of development. The embry- os hatch as plauktotrophic veliger lanae. DISCUSSION Crepidula argentina new species is usually foimd on the posterior edge of fiviug shells of Mytiliis edulis platensis d'Orbigny, 1846, typically in association with Calyp- traeotheres gaiihi (Fenucci, 1975) (Crustacea: Brachi- ura). Mvtihd banks, distributed all around the coast of Argentina (Penchaszadeh, 1971a) are a suitable habitat for Crepidula argentina. Crepidula argentina was included by different authors (Parodiz, 1939; Hoagland, 1977; 1983,' in part) in C. pro- tea and referred to as C. ungiiifonnis bv Penchaszadeh (1971b: 480). The t\pe specimens of C. protea were ex- amined by Hoagland (1983). At BMNH, there are two lots with 5 and 11 syntypes under the numbers 18.54.12.4.573 and 574 respectively. Aguirre (1993) des- ignated and illustrated the only whole specimen as lec- totype plus tvvo paralectotypes, with no mention to Figures 42-46. Anatomy of Crepidula an^cntina new species. 42. Stomach, ventral view, inner surface exposed by means of a longitudinal incision. 43. Head of male, dorsal view. 44. Visceral mass and adjacent part of pallial cavity, male, ventral view; 45. Pallial oviduct, ventral view. 46. Penis, ventral view. All scale bars = 1 mm. Page 140 THE NAUTILUS, Vol. 114. No. 4 Hoagland's paper or d'Orbigny's (in Sagra) illustrations. Most of the type specimens are smaller than 30 mm and have the conchological attributes of what Hoagland de- scribed for C. protea from Brazil. In terms of shell char- acters, C. protea and C. argcntino new species are sim- ilar. Measurements of C. argentina in Table 1 are iden- tical to those given to C. protea by Hoagland (1977). However, in a later paper, Hoagland (1983) mentioned that the largest specimen of C. protea was 20 mm long. Therefore, we suspect that both species were combined under the name C. protea in her 1977 paper Crepidida argentina is larger and wider, particularly in young spec- imens. Crepidula protea has an elliptical aperture with a thicker and more convex shell, while the new species has a subcircular aperture and a thinner and flatter shell. Both species differ primarily in their anatomical features and reproductive strategy'. Further anatomical study of C. protea is provided in another paper (Simone, sub- mitted); some data from that paper are here discussed for comparative purposes. Crepidula argentina differs moq^hologically from C. protea in that C. argentina has: 1) a more developed lateral shell muscle, 2) a poorly developed dorsal shell muscle, 3) the kidney proportion- ally smaller and with a different arrangement of inner folds of dorsal lobe, 4) presence of a renal vessel edging rectum and working as adrectal sinus, 5) a connection between odontophore muscle pTtle Beach, SC 29852-2846 USA. the northern Indian Ocean, through Indonesia to Aus- traha and northward to Japan, usually at subUttoriil depths. As pointed out by Verhecken (1986a: 38—40), there is considerable confusion in the literature as to the correct nomenclature for many of the currently recog- nized species of Merica. This paper describes three new Recent species ot Merica, one from Soudi Africa and two from the Phil- ippines. The assignment of Cancellaria oblonga Sower- bv, 1825, to the genus Merica is reaffirmed, and the spe- cies is illustrated for comparative purposes. Holofypes of the new species are housed in the collections of the Na- tal Museum (NM), Pietermaritzburg, Repubhc of South Africa, and the National Museum of Natural History (USNM) USA. Smithsonian Institution, Washington, DC, SYSTEMATICS Family Cancellariidae Forbes and Hanley, 1851 Subfamily Cancellariinae Forbes and Hanley, 1851 Genus Merica H. and A. Adams, 1854 Merica H. and A. Adams, 1854:277. Tvpe species: Cancellaria melanostoma Sowerbv, 1849, by subsequent designation of Cossmann, 1899. Diagnosis: Shell with conical spire, large body whorl, paucispiral protoconch weakly to strongly deflected from teleoconch axis, prominent suture, generally weak si- phonal fasciole, narrow pseudo-umbihcus, and large, ovate aperture. Columella short, thick, weakly concave, with 2 sharply keeled columellar folds and strong si- phonal fold. Columellar Up with pustules and secondary folds. Parietal callus generally present. Emargination along outer lip at base of siphonal canal poorly defined or absent. Remarks: Merica appears to be the Indo-Pacific stem group for the genus Cancellaria, which is hmited to the eastern Pacific and the western Adantic. Merica differs in lacking a pronounced emargination along the outer lip of the shell as well as a heavy, bifurcated posterior R. E. Petit and M. G. Harasewvch, 2000 Page 143 columellar fold, both characteristic of CanccUnria. ('oss- inann (1899: 13) accorded generic status to Mcrica, and most authors have followed this. Kuroda and Habe (1971: 310, 202) proposed the ge- nus Momocbora based primarily on the presence of a strongly dexiated protoconch in its tvpe species, Can- ccllaria sinensis Reeve, 1856. Mouiorhora has been con- sidered a spionxTii of Mcrica by most subsecjuent au- thors (Garrard, 1975: 3; Verhecken, 1986a: 44). Mcrica oblonga (Sowerbv, 1825) (Figures 1, 5, 6) Cancellaria oblonga Sowerby, 1825: Appendix: 15; 1S32: fig. 19. Cancellaria bifasciata Deshayes, 1830: 181; Lobbecke, 1885: 30, pi. 9, figs. 1,2. Cancellaria oblonga Sowerby: Kieiier, 1841: 6, pi. 3, fig. 3; Ab- bott and Dance, 1982: 225; Springsteen and Leobrera, 1986: 78, pi. 18, fig. 26. Cancellaria (Merica) oblonga Kiener [sic]: Chenu, 1859: 277, fig. 1847. Not Cancellaria (Merica) bifasciata Deshayes: Chenu, 1858: 277, fig. 1845 [?=M. sinensis (Reeve, 1856)]. Not Cancellaria bifasciata Deshayes: Bamard, 1959: 13-14; Kenslev, 1973: 194, fig. 749 [ = M. hissii new species]. Merica bifasciata (Deshayes): Habe, 1961: pi. 24, fig. 27. Cancellaria (Merica) bifasciata Deshayes: Oyama and Take- mura, 1963: Cancellaria plate, fig. 6. Merica oblonga (Sowerbv): Petit, 1974: 112, fig. 5; Verhecken, 1986a: 41, figs. 7-8; Verhecken and Wranik, 1991: 60; Ver- hecken, 1997; 308, fig. 37. Momoebora bifasciata (Deshayes): Higo. 1973: 179. Momocbora oblonga (Sowerby): Higo and Goto, 1993: 276; Higoe/fl/., 1999: 294. Remarks: For many years there was confusion about priority of the names apphed to this species, with Can- cellaria bifasciata Deshayes, 1830, generally given pre- cedence over Cancellaria oblonga Sowerby, 1825. These two available names were correctly dated by Perit (1974). Merica oblonga (as Merica bifasciata) was attri- buted to the genus Momoebora by Higo (1973:179), and this placement remained unchanged in the two later re- visions and enlargements of that work (Higo and Goto, 1993; Higo et al, 1999). As M. oblonga has a protoconch that is not strongly deflected from the coihng axis of the teleoconch (Figs. 5,6), we see bttle justification for the usage of Momoebora. There were, and to a lesser degree still remain, ques- tions about the geographic range of this species. Its oc- currence in Panama, as reported by Kiener (1841), was shown to be incorrect by Keen (1971:649). Verhecken (1997:309) gave the distribution of Merica oblonga as "Japan to Indonesia; Northern Indian Ocean to Aden; Eastern South Africa (?)." His query regarchng the South African records is appropriate, as the South African specimens previously identified as Mcrica bifasciata rep- resent a new species, described herein as Merica hissii. There are no verifiable Japanese records for Mcrica oblonga (Kazunori Hasegawa, personal communication). The figme of M. oblonga pubUshed by Habe (1961) and ri'produced b\' Petit (1974) is of a specimen from Taiwan that was inchulcd b\' Habe for comparison with Merica asprella [sic] Lamarck. The specimen figured by Oyama and Takemura (1963) is also from Taiwan. Verhecken's (1986a: 41) record for Japan is based on Habe's usage. Records of M. oblonga are noticeably absent in most standard Japanese compendia and fists. Its inclusion in the exhaustive faunal fists of Higo (1973), Higo and Goto (1993) and Higo, Callomon and Goto (1999) was based on a Japanese publication of uncertain authorship. Mcrica oblonga ranges as far north as Taiwan. It is uncommon in the Phifippines. Springsteen and Leobrera (1986: 78) record this species as "sporadically found throughout the Philippines in limited quantity-." The col- or figure in Abbott and Dance (1982:225) is of a Phil- ippine specimen taken from over 30 m depth in tangle nets (Petit collection). Verhecken and Wranik (1991: 60) state that M. oblonga 'seems to have its distribution cen- ter in the eastern Inthan Ocean (Gulf of Bengal). ' While this species is uncommon in other areas, it appears to be common in India. We have seen commercial lots from India containing hundreds of specimens of M. ob- longa. Merica litssii new species (Figures 2, 7, 8) Description: Shell (figure 2) to 35 mm, thin, biconic, elongate. Spire high (spire angle 53-55°) comprising one-half of total shell length. Protoconch (figures 7, 8) paucispiral, nearly coaxial with teleoconch (deviated from coifing axis by up to 4°), increasing in diameter from 300 p,m to 2.15 mm in 2% elongated whorls. Tran- sition to teleoconch marked by onset of spiral then axial sculpture. Teleoconch of about 5 evenly rounded whorls. Shoulder rounded. Suture broadly impressed, forming conspicuous canal. Axial sculpture of 30-34 very weak, rounded, sfightly prosocline ribs, as broad as inter\'ening spaces. Axial ribs become prominent, irregular, crowded on final V4 of body whorl. Spiral sculpture more pro- nounced than axial, of regularly spaced, flat, primary cords (8-10 on penultimate whorl, 2.5-32 on body whorl) with weaker secondaiy cords present irregularly, but most often on posterior portion of whorls. Aperture large, broadly ovate, deflected from coifing axis by 16- 17°. Outer lip thin, weakly flared outwarcUv, without cfis- cemable emargination. Inner fip smooth or with weak spiral firae reflecting spiral cords, especially in the an- terior portion. Inner lip with 2 columellar, 1 siphonal fold. Siphonal fold flat, bifid on holot^pe but sharp on other specimens. Columellar lip thick, forming induc- tural shield that obscures chink-fike umbificus. Siphonal canal short, broad, inconspicuous. Base color fight yel- low-brown, irregularly marked with white and various shades or redcfish brown. Two bands of brown some- times present, visible inside aperture. Wider band just below shoulder, narrower band just posterior to base of Page 144 THE NAUTILUS, Vol. 114, No. 4 R. E. Petit and M. G. Harasewvch, 2000 Page 145 siphonal canal. Axial flammules of browii, \t>ll()\v, white, irregular in outline and intensity Tjpe locality: Off Park Rviiie, about SO km S ol Dur- ban, on the KwaZulu-Natal south coast. South Africa, appro>dmatelv 30°17' S, 3()°45' E, dredged in 150 ni on nibble and sponge substrate. Type material: Holot\pe, NM ¥7648/11312, height 34.4 mm; Paratvpe 1, Marcus Lussi Collection, both from the t\pe locality-; Paratope 2, Dawni Brink Collec- tion, near type localit\' in 140 m, nibble and sponge bot- tom; Paratvpe 3, NM E8794, Zululand, off Matigulu River mouth, 29°21.4' S, 3r56.5' E, in 145 m, mud, shell rubble, NMDP 16 Apr 1989, station ZR7. Etymology: Named for Mr. Markus Lussi of Durban North, SoutJi Africa, who has made numerous contri- butions to the Uterature on South African Mollusca, and who provided specimens for study. Comparative remarks: Merica lussii differs from M. ohlonga in having a broader shell with a stepped spire, a proportioniilly shorter but broader aperture, and a deeply impressed sutural canal. Merica lussii lacks the closely packed strong axial ribs that produce the finely beaded surface characteristic of M. ohlonga. Barnard (1959: 14) and Verhecken (1986a: 42) have previously suggested that South African records of M. bifasciata ( = M. oblonga) might represent a distinct species. Merica detjnzeri new species (Figures 3, 9, 10) Description: Shell (figure 3) to 35 mm, thick, biconic, weakly pseudo-umbilicate. Spire high (spire angle 55- 58°). Protoconch (figures 9, 10) paucispiral, deviated from coiling a.xis by about 14°, increasing in diameter from 300 jim to 2.15 mm in 2V4 evenly rounded, glassy whorls. Transition to teleoconch marked by onset of spi- ral cords followed almost immediately by axial ribs. Te- leoconch of up to 6 whorls. Shoulder inconspicuous. Su- ture deeply impressed, forming narrow canal. Axial sculpture of 27-33 low, evenly spaced, prosochne ribs, rounded in profile, as wide as intervening spaces. Peri- odic internal varices (see Harasewych and Petit, 1982: 111) barely discernable on outer surface of shell by thickened axial rib slightly expanded along edge of su- tural canal followed closely by thin axial rib and slightly reflected fip, begin after about 2% whorls, continue at intervals of about 240° thereafter. Spiral sculpture of strong, broad cords (20-26 on body whorl, 8-9 on pen- ultiiriatc whorl) that are narrower, more closely spaced near suture and siphcjn. Spiral cords overlav axial ribs to produce a cancellate surface on early whorls. Single, sec- ondarv' cords occur between primary cords along central region of whorls. Axial ribs Iteconie progressivi'ly broad- er, irregular, more widely spaced on last half of body whorl, obscuring cancellate appearance. Aperture large, broad, nearl\' hemi-elliptical, deflecteil from coifing axis by 14-15°. Outer lip weakly recurved in profile, with finely .serrated edge, lacking emargination. Inner surface of outer Up with 16 recessed spiral Urae that diminish 1/ 3 whorl into the aperture. Inner Up with 2 columellar, 1 siphonal fold, columellar lip nearly covering pseudo-um- bificus. Posterior columellar fold strongest, overlying weak siphonal fasciole. Anterior columellar fold extends to edge of columellar fip. Spiral ridge parallels anterior columellar ff)ld adapically, but is confined to the colu- mellar hp without extending into aperture. Surface of siphonal fold with single pustule in some specimens. Si- phonal canal short, narrow, axial. Color uniform yellow- brown, sometimes with narrow darker bands above and below mid-whorl. Portions of spire whorls intficate that juveniles may be mottled. Type locality: Off Baficasag Island, Bohol, Phifip- pines. Taken in tangle nets at approximately 200 meters. Type material: Holotvpe, USNM 880645, height 35.2 mm; Paratype 1, R. Petit Collection 2465, both from the type locahty; Paratypes 2-A, USNM 242316, off Pt. Du- murug, Ma.sbate, Philippines (12°00'30" N, 124°05'.36" E), dredged in 280 m, green mud, USES Albatross, sta. 5394, l3'Mar 1909; Paratvpes 5-8, USNM 242321, off Adyagan Island, E Masbate, PhiUppines (12°12'35" N, 124°02'48" E), dredged in 247 m, green mud, sand. USES Albatross, sta. 5392, 13 Mar 1909; Paratype 9, USNM 277485, off Sibugav Island, E of Masbate, Phil- ippines (12°04'15" N, 124°()4'36" E) in 198 m, grav sand, mud, 15.5°C. USES Albatross, sta. 5212, 20 Apr'l908. Etymology: We take pleasure in naming this species for Mr. Al Deynzer of Sanibel, Elorida, who obtained these and other uncommon Cancellariidae in the Phil- ippines and made them available for study. Comparative remarks: This new species is distinc- tive in having a well-developed sutural margin strongly rounded to form a deeply impressed sutural canal. Mer- ica deynzeri is similar to, but stockier and broader than M. oblonga, from which it can also be distinguished by its much stronger and coarser axial and spiral sculpture, and by its more rounded aperture. A notable feature of <- Figures 1—4. Apertural, lateral and dorsal views of the shells of. 1. Merica oblonga (Sowerby, 1825), USNM 845168, taken by fishing trawler in Manila Bay, Luzon, Philippines, 1969 2. Merica lussii new species, holotype, NM V7648Ari.312, off Park Ryiiie, about 80 km south of Durban on the KwaZulu-Natal south coast. South Africa, approximately 30°17'S, .30°45'E, dredged in 150 meters, rubble and sponge substrate. 3. Merica deijnzeri new species, holotype, USNM 880645. off Balicasag Island, Bohol, Phil- ippines, taken in tangle nets from about 200 meters. 4. Merica ekti/phos new species, holotype, USNM 880646, Off Balicasag Island, Bohol, Philippines, taken in tangle nets from about 200 meters. Scale bar = 2 cm for all figures. Page 146 THE NAUTILUS, Vol. 114, No. 4 Figures 5-12. Scanning Electron Micrographs of a\ial and lateral aspects ot pnitcicDnclis nt spcnini-ns iii figures 1—4. 5-6. Merica oblonga (Sowerby, 1825). 7-8. Merica lussii new species. 9-10. Merica detjnzeri new species. 11-12. Merica ekttjphos new species. Arrowheads indicate transition from protoconch to teleoconch. Scale bar = 1 mm for all figures. R. E. Petit and M. G. Harasewvch, 2000 Page 147 Merica deynzeri is the lack of color bands that arc char- acteristic of most species of Merica. Merica ektyphos new species (Figures 4, 11, 12) Description: Shell (figure 4) to 33 mm, very thick, globose, pseudo-umbilicate. Spire relatively low (spire angle 63°). Protoconch (figures 11, 12) paucispiral, de- viated from coiling ;L\is by about 8°, increasing in di- ameter from 400 |xm to 2.10 nun in 2y2 low, evenlv rounded whorls. Transition to teleoconch marked by on- set of axial ribs followed by overlying spiral cords. Te- leoconch of up to 6 rounded whorls. Suture impressed, forming deep, narrow sutural canal. A.\ial sculpture of prominent, prosocline ribs (31 on body whorl, 31 on penultimate whorl). Ribs become smaller, more densely spaced in narrow bands spaced every Vi to % whorl, in- dicating position of internal varices. Spiral sculpture of strong, flattened cords (15-16 on bodv whorl, 6 on pen- ultimate whorl) that form nodes at intersections with ax- ial ribs. Nodes especially pronounced at margin of su- tural canal, forming serrated ridge. Finer secondaiy threads present between adjacent cords above suture line. Aperture large, deflected from coifing axis by 12°, with broadly elfiptical outer lip, sharp, 48° angle between columella and parietal wall. Outer fip prosocline, sfightly crenate, with 12-13 strong, sfightly recessed firae that become reduced, disappearing about V4 whorl within the aperture. Outer lip with emargination discernible only by touch as shallow depression. Columella with 2 colu- mellar, 1 siphonal fold. Posteriormost fold strongest, overlving conspicuous siphonal fasciole, does not reach edge of columellar lip as anterior and siphonal folds do. Columellar fip thick, rounded, nearly covering pseudo- umbificus. Four weak, short spiral ridges run along col- umellar fip parallel to and posterior to anterior colu- mellar fold, but do not extend into aperture. Siphonal canal short, narrow, well demarcated, deflected from axis. Color whitish, with ginger brown bands along su- ture (darkest), above mid-whorl (broadest), below mid- whorl (narrowest) and along siphonal fasciole. Gaps in coloration of sutural band correlate with internal varices. Type locality: Off Baficasag Island, Bohol, Phifip- pines. Taken in tangle nets at appro.ximately 200 meters. Type material: Holotype, USNM 880646, 32.3 mm; Paratypes 1-4, R. Petit Collection 2472; Paratypes 5-7, Deynizer Collection, all from tyjie locafity. Etymology: Gr ektyphos. puffed up. Named for its inflated shefl. Comparative remarks: This new species can be dis- tinguished from all known species of Merica by the thickness of its shell and the strength of its sculpture. In outfine it is closest to the Australian M. wcstralis (Gar- rard, 1975), from which it differs in having much coarser axial and spiral sculpture, a straighter columella with Tabic 1. Merica deynzeri new species. Measurements of shell characters. Linear measiirpnients in mm. n = 5. ° n = 3 for no. whorls, protoconch. (;!iaractcr Mean SU Range Shell length (SL) .35.0 2.6 .34.9-38.6 Aperture length (AL) 22.1 1.7 19.6-24.3 AIVSL 0.629 0.003 0.624-0.632 No. whorls, protoconch" 1.5 0 2.5-2.5 No. whorls, tcleoconcii 5.53 0.14 5..33-5.67 .Spire angle 56.7° 1..3° 5.5-58° stronger folds, and a body whorl that is niuch less con- stricted beliind the siphonal fasciole. ACKNOWLEDGMENTS We thank Marcus Lussi and Dawn Brink of South Africa for making specimens of Merica lussii available for this study. While on visits to the Phifippines Al Deynzer made special efforts to obtain specimens for this study. We are grateful to Paul Callomon of Osaka and Dr. Ka- zunori Hasegawa of Ibaraki for information about the Japanese records of M. ohlonga. LITERATURE CITED Abbott, R. T. and S. P. Dance, 1982. Compendium of Seashells. E. P Dntton. New York, x -I- 411 pp. Adams, H. and A. Adams. 18.5.3-.58. The genera of Recent Mol- lusca; arranged according to their organization. John van Voorst, London. 2 vols. [1:1-256, pis. 1-.32, 1853; 1:257- 484, 2:1-92, pis. .33-72, 1854; 2:9.3-284, pis. 7;3-96, 1855; 2:285-412, pis. 97-112, 18.56; 2:41.3-540, pis. 113-128, 1857; 2:.541-660. pis. 129-1.38, 18,58.] Barnard, K. H. 1959. Contributions to the faiowledge ot South African marine Mollusca. Part II, Gastropoda: Prosobran- chiata: Rachiglossa. Annals of the South African Museum 45:1-2.37. Chenu, J. C. 1859-62. Manuel de conch\liologie et de paleon- tologie conchyliologique. Librarie Victor Masson, Paris. 2 vols. [l:i-vii, 1-508, 18.59; 2:1-327, 1862]. Cossmann, M. 1899. Essais de Paleoconchologie Comparee. Troisieme hvraison. Paris. 201 p., 8 pis. Deshayes, G. P. 1830. Encyclopedic Methodique. Histoire na- turelle des vers 2(l):l-256, Paris. Forbes, E. and S. Hanley 1848-1853. A history of British Mob lusca and their shells. London. 4 volumes. [Published in parts; for dates of parts see Fischer and Tonilin, 19.35, Journal of Conchology 20(5): 150-151.] Garrard, T. A. 1975. A revision of the Australian Cancellariidae (Gastropoda: Mollusca). Records of the Australian Muse- um .30:1-62. Habe, T. 1961. Description of four new cancellariid species, with a list of the Japanese species of the familv Cancel- lariidae. Venus 21:431-441. pis. 2:3-24, Harasewvch, M. G. and R, E, Petit. 1982. Notes on the mor- phology of Cancellaria reticulata (Gastropoda: Cancellar- iidae). The Nautilus 96:104-113. Higo, S. 1973. A catalogue of molluscan faima of the Japanese Page 148 THE NAUTILUS, Vol. 114, No. 4 Islands and the adjacent area. S. Higo. Isahaya, Japan. (58) + 397 + 61 pp. Higo, S. and Y. Goto. 1993. A systematic list of molluscan shells from the Japanese Is. and the adjacent area. Elle Corpo- ration, Osaka, Japan,3 + 22 + 693 + 13 + 149 pp. Higo, S., Callomon, P. and Y. Goto. 1999. Catalogue and bib- liography of the marine shell-bearing Mollusca of Japan. Elle Scientific Publications, Osaka, 49 pp. Janssen, A. W. 1984. An account of the Cancellariidae (Gastro- poda) of Winterswijk-Mi.ste (Miocene, Henimoorian),The Netherlands. Scripta Geologica 68; 1-39, pis. 1-6. Keen, A. M. 1971. Sea shells of tropical West America. Second edition. Stanford University Press, Stanford, xiv -t- 1064 pp., 22 pis. KensJey, B. 1973. Sea-Shells of southern Africa, Gastropods. Maskew Miller Ltd., Cape Town, 236 pp. Kiener, L. C. 1841. Genre Cancellaire. Species general et icon- ographic des coquiUes vivantes. Paris, 44p., 9 pis. Kuroda, T. and T. Habe. 1971. [Descriptions of species] In: Kuroda, Habe and Oyama, The sea shells of Sagami Bay. Manizen Co., Tokyo. Pp. i-xix, 1-741 (Japanese), 121 pis., 1-489 (English), 1-51 (index). Ladd, H. S. 1982. Cenozoic fossil mollusks from Western Pa- cific Islands; gastropods (Eulimidae and Volutidae through Terebridae). United States Geological Suwey Professional Paper 1171:1-100, pis. 1-41. Lobbecke, T. 1881-1887a. Das genus Cancellaiia. Systema- tisches Conchylien-cabinet von Martini und Chemnitz 4: 1-96, pis. 1-23 [issued in parts; pt. .309; 1-16, pis. 1-5, 1881; pt. 335;17^32, pis. 6-10, 1885; pt. .340;.3;3-56, pis. 11-15, 1886; pt. 343;5780, pis. 16-20, 1886; pt. .346;81- 96, pis. 21-23, 1887.] Matsukiuua, A. 1998. Index and collation of The Molluscan Shells by Katura Oyama and Yoshio Takemura (1957- 1963). The Yuriyagai 6;91-122. Noda, H. 1980. Molluscan fossils from the Ryukyu Islands, southwestern Japan. Part 1. Gastropoda and Pelecypoda from the Shinzato Fomiation in southeastern part of Old- nawa-jinia. Science Reports of the Institute of Geoscience, University of Tsuk-uba, SecHon B, l;l-95, pis. 1-12. Oyama, K. and Y. Takemura. 1963. The Molluscan Shells. Pt. 6, Cancellaria plate, [for collation of all parts and plates see Matsuk-imia. 1998]. Petit, R. E. 1974. Notes on Japanese Cancellariidae. Venus 33: 109-115. Reeve, L. A. 1856. Monograph of the genus Cancellaria Con- chologia Iconica 10; [unpaginated text], pis. 1-18. Sacco, F. 1904. I molluschi dei terreni terziarii del Piemonte e deila Liguria. Pt. .30 (Aggiunte e Correzione). C. Clau- sen, Torino, xxxvi + 203 pp., 31 pis. Shuto, T. 1969. Neogene gastropods from Panay Island, the Philippines (Contributions to the Geology and Paleontol- ogy of Southeast Asia, LXVIII). Memoirs of the Faculty of Science, Kyushu University, Series D, Geology 19(1): 1-250, pis. 1-24. Sowerby, G. B., I. 1825. A catalogue of the shells contained in the collection of the late Earl of Tankerville, arranged ac- cording to the Lamarcldan conchological system; together with an Appendix, containing descriptions of many new species. London, vii -1-92-1- xxxi\- pp. [Appendix], 9 pis. Sowerby, G. B., II. 1849. Descriptions of some new species of Cancellaria m the collection of Mr H. Cuming, Proceed- ings of the Zoological Society of London for 1848:1.36- 138. Springsteen, F J, and F M. Leobrera. 1986, Shells of the Phil- ippines. Carfel Seashell Museum, Manila, 377 pp. Verhecken, A. 1986a. The Recent Cancellariidae of Indonesia (Neogastropoda, Cancellariacea). Gloria Maris 25(2):29- 66. Verhecken, A. 1986b. A revision of the Cancellariidae (Neo- gastropoda: Cancellariacea) of the Red Sea and the Gulf of Aden. Gloria Maris 25(4): 13.3-153. Verhecken, A. 1997. Mollusca Gastropoda: Arafura Sea Can- cellariidae collected during the KARUBAR Cmise; 7m. A. Crosnier and P. Bouchet (eds), Resultats des Campagnes MUSORSTOM, Vol. 16. Memoirs du Museum d'Histoire Naturelle 172:29.5^23. Verhecken, A, and W. Wranik. 1991. Additional data on the Cancellariidae of the Gulf of Aden. Gloria Maris .30(4); 59-63. Wilson, B. R. 1994. AustraUan Marine Shells. Prosobranch Gastropods, Part two (Neogastropods). Odyssey Publish- ing, Kallaroo, 370 pp. I THE NAUTILUS 114(4):149-154, 2000 Page 149 Systematics of the genus Infundihulum Montfort, 1810 (Gastropoda: Trochidae) Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa P.O. Box 467, Wellington NEW ZEALAND brucem@tepapa.govt.nz Two Recent Infundibtiluin species are recognized: /. concaviiin (Gnielin, 1791) (type species) and /. tonilini (Fulton, 1930). Their shells are convergent on those of calyptraeid limpets of the genus Trochita Schumacher, 1817, having the most widely excavated bases and the most strongly tangential apertures of any living Trochini. They Hve firmly attached to rocks on highly e.xposed shores from low tide level to several meters depth in tropical seas. Reliable records of /. concavum are from Reunion and Mauritius. All Recent specimens of /. tom- lini seen were from old collections, and most have either minimal, often dubious locality data, or none at all, though it would seem that /. tomlini has a patchy chs- tribution in the tropical southwest Pacific. Additional key words: Mollusca, Trochoidea, fossils. Pleistocene, Kermadec Islands. INTRODUCTION The present contribution was initiated some years ago when I recognized a species oi Infundihulum among Pleistocene fossils from Raoul Island, Kermadec Islands, northeast of New Zealand. A search of the literature re- vealed that Infundihulum species are sui"prisingly poorly known. Moreover, inquiries revealed that most of the few specimens of Infundihulum species in museum col- lections were acquired during the nineteenth century, and have broad, often dubious locality data, or none at all. The fossils from Raoul Island prove to be the little known species /. tomlini (Fulton, 1930), which is rep- resented in old collections labeled as originating vari- ously from Penang (Pinang, Malaysia), Australia, Upolu (Western Samoa), Lifou (Loyalty Islands) or the New Hebrides (Vanuatu). Of these locality records, only Upo- lu, Lifou, and Vanuatu seem hkely to be correct. Mu- seum specimens of the type species (Trochus concavus Gmehn, 1791) are labeled as having originated from lo- calities in the Indian Ocean, as well as the Red Sea and Cochin China (southern Vietnam), though the only re- hably locaUzed specimens seen during the present study were from Reunion. An explanation for the poor rep- resentation of /. concavum (and presumably /. tomlini) in collections is suggested by the fact that specimens collected ahve were cryptic at and below low tide, at- tached to exposed, wave-swept volcanic rocks, which tend to receive far less attention from collectors than more congenial and species-rich coral reef and lagoon environments, where they have never been recorded. In- stitutional acronyms: ANSP, Academy of Natural Scienc- es, Philadelphia; AMS, Australian Museum, Sydney; RMNH, The Natural Histoiy Museum, London; LACM, Los Angeles County Museum of Natural History; MNHN, Museum National d'Histoire Naturelle, Paris; NMNZ, Museum of New Zealand, Welfington; NMP, Natal Museum, Pietermaritzburg; USNM, National Mu- seum of Natural History, Washington, DC. SYSTEMATICS Superfamily Trochoidea Rafinesque, 1815 Family Trochidae Rafinesque, 1815 Subfamily Trochinae Rafinesque, 1815 Tribe Trochini Rafinesque, 1815 Genus Infundihulum Montfort, 1810 Infundibuluiii Montfort, ISIO: 167. Type species (by original designation): Infundihulum tijpus Montfort, 1810 = Tro- cluis concavus Gmelin, 1791; Recent, Rennion. Carinidea Swainson, 1S40: .350. T^pe species (by nionotypy): Trochus concavus Gnielin, 1791. Description: Shell up to 54 mm in diameter, stout, base excavate, umbilicus narrow; aperture very strongly tangential, as wide as shell; periphery sharply angulate, not stellate at any stage of growth, almost flush against a flat surface. Spire sculptured with strong, close, round- ed spiral cords and a.xial costae, intersections finely and bluntly nodular Base weakly convex, evenly curving into narrow umbilicus, excavated from periphery. ColumeUar lip adapicallv with strong, rounded denticle, sharply re- tracted to insertion within umbilicus. Foot (/. concavum) very large, epipothum very well developed, edge finely fringed, well developed epipodial flaps beside epipodial Page 150 THE NAUTILUS. Vol. 114, No. 4 insertion; cephalic lappets large, papillate. Radula (/. concavum, figure 9) with the formula ii + .5+l+5 + n, very similar to that in Trochiis (scnsu stricto). Remarks: Infundibiilum species are referable to sub- family Trochinae, tribe Trochini because of similaritv' to species oiTrochtis Linnaeus, 175S, and Tectiis Montfort, 1810, in shell and radular morphology, and external anat- omy. External anatomy is close to that of Tcctiis fenes- trattis (Gniehn, 1791) (Hickman and McLean, 1990: fig. 55b). lufiindibtihtm species differ irom Trochiis and Tcc- tus in the combination of low, evenly conical spire, weak- ly convex whorls, low sculpture, lack of peripheral nod- ules at any stage of growth, and widely excavated base, which fits almost flush with a flat surface. All of these character states (apomoq^hies) evidentlv enhance attach- ment to rocks exposed to strong wave action. Infundib- iilum concavum (Gmehn, 1791) and /. tomlini (Fulton, 19.30) have the most strongly tangential apertures and the most widely excavated bases of any li\ing Trochini, and they resemble limpets in ventral view when the foot is fully extended (shdes of aquarium specimens of 7. con- coviim provided by D. G. Herbert). Gross shell mor- phology strikingly parallels calyptraeid fimpets of the ge- nus Trochita Schumacher, 1817. Whereas Infiindibulum has been interpreted as a sub- genus of Trochus by many authors, I prefer to treat In- fiindibulum as a distinct genus because of current lack of clear exddence for degrees of relatedness. Trochus hochii Phihppi, 1844, from Oman (figures 8, 10) has a similar shape and sculpture to Infundibiihtm species, but differs in having the base considerably less excavated, and a much more weaklv tangential aperture (and thus presumably a more weakly expanding foot). The relationships of this species are unclear, and I refer it to Trochus, subgenus Infundibidops Pilsbry, 1889 (type species Trochus cri/thracus Brocchi, 1821) with some hesitation. Inftindibiihim concavum (Gmelin, 1791) (Figures 1-.3, 9) Trochus ptjramidalis rarissimus . . . Chemnitz, 1781: 86, pi. 168, figs. 1620, 1621 (not binominal). Trochus concavus Gmelin, 1791: .3.570 (refers to Chemnitz, 1781. figs. 1620, 1621); Dillw>ii. 1817: 763; Blaimille. 1825: 425. pi. .32 bis. fig. 1; Wood. 1825:134. pi. 28. fig. 7; Sganzin. 1843: 22; Philippi. 1848: .38. pi. 8. figs. 7. 8; Deshayes. 1863: 71; Fischer. 1876: 105. pi. 33, fig. 2; Kaicher. 1979: card 2087. Infundibulum tijpus Montfort, 1810: 167, text fig. Carinidea concavus: Swainson, 1840: .350. Pohjdonta (Carinidea) concavus: Chenu. 1859: 357. fig. 2646 Trochus (Pohjdonta) concavus: Martens. 1880: 295. Trochus (Infundibulum) concavus: Pilsbry, 1889: 40, pi. 43, fig. 13; VVenz, 1938: 311, fig. 693. Infundibulum concavum: Cossmann. 1918: 188. text fig. 65. Tectus concavus: Dri\'as and Jay. 1988. pi. 2, fig. 3. Not Poh/donta (Infundibulum) concava: MeMlI and Standen, 1895: 125 = /. tomUni. Description: Shell (figures 1-3) up to 56 mm in di- ameter, stout, broadly conical, wider than high, weakly cyrtoconoid, base excavate; aperture strongly tangential, as wide as shell; peripherv sharplv angulate, almost flush against a flat sin4ace, narrow umbihcus. Spire dull green with subsutur;il band of irregular, duU, red and white maculations, last few whorls pinkish milky green. Base white, inner half with porcellanous glaze. Protoconch and first few teleoconch whorls unknown (eroded). Spire whorls moderately convex at first, becoming weakly con- vex, sculptured with low, rounded spiral cords and op- isthocline, non-coUabral axial costae, interspaces narrow- er; intersections with strong, bluntlv rounded nodules; axial costae on last 2 or .3 whorls traversing rounded axial folds that are broader than the costae and tend to be more strongly opisthochne. Spiral cords 7, strong and similar on early whorls, weakening after shell attains about 8 mm diameter, obsolete on last 2 or 3 adult whorls. Excavate base convex, smoothly cuning into nar- row umbihcus. Outer part of base with 6 or 7 smooth, simflar spiral cords, becoming obsolete on last whorl at maturit\'; inner (porceUanous) part of base with 6 or 7 spiral cords that become obscured bv indiictura at ma- turity. Aperture ovate. Basal Mp strongly tluckened, smoothly continuous with columellar lip, which is stronglv thickened adapicallv then sharplv retracted to insertion within umbihcus. Animal (retracted, ex ethanol). Foot extremely large and spreachng. Epipodium very weU developed, edge finely fringed throughout; inner surface finely papillate, tapered macropapillae over posterior half neck lobes convoluted, finely fringed, left lobe considerably larger and more deeply convoluted than right. Epipodial ten- tacles slender, narrowly tapering, 4 pairs. Epipodial flaps well-developed, bases close to epipodial insertion, edges finely fringed, 5 on right, 5 or 6 on left. Cephahc ten- tacles slender, narrowly tapered, subcyhndrical. right tentacle base adjacent to tip of left tentacle due to obhque asymmetry of head. Eye stalks large, dorsoven- trally flattened, well developed eyes in tips. Cephahc lap- pets well developed, edge and adjacent ventral surface Figures 1-10. Infundibulum and Trochus (Infiindibulops) species. 1-3. /. concavum (Gmelin. 1791). St. Paul Bav. Reunion, .56 X 40 mm (NMNZ M. 270521). 4-6. I tomlini (Fulton. 19.30). s\Titype, locality unhiown (possibly Western Samoa or Vanuatu). 19.0 X 26.6 mm (BMNH 19.30.4.2.3). 7. 7 tomlini. Boat Cove. Raoul Island. Kemiadec Islands (Pleistocene). 25.5 X .33 (est.) mm (NMNZ M.214580); 8, 10. Trochus (Infundibidops) kochii (Philippi. 1844). SW coast of As Sawda. Juzor Al Halaaniyaat. Oman, 32 X ,38 mm (Zoological Museum, Amsterdam). 9. Radula of 7 concavum. north of Cap la Houssaye, St. Paul Bav. Reunion (NMP K4898). B. A. Marshall, 2000 Page 151 Page 152 THE NAUTILUS, Vol. 114, No. 4 papillate. Cephalic tentacles blackish brown, sides of foot darker, elsewhere cream. Radula (figure 9) with the formula n+5+1+5+n, very similar to that in Trochtis {scnsit stricto) (Hickman and McLean, 1990: fig. 57D). Type data: Holotype, the specimen illustrated by Chemnitz (1781: figs. 1620, 1621), ex Spengler collec- tion, in Zoological Museum, Copenhagen (T. Schi0tte, pers. comm.); "k-uste von Coromandel". Material examined: North of Cap la Houssaye, St. Paul Ba\-, Reunion, exposed rocky shore, extreme low tide to 2 m, 24 September 1988, R. N. Kilbum and D. G, Herbert (3, NMP); St Paul Bay, Reunion, breaker zone on exposed rock-\' shore at low tide, M. Jay, 1994 (.3 NMNZ); St Paul, Reunion, M. Jav, 1985 (3 ANSP); "Zanzibar" (8 in 3 lots, MNHN): Seychelles (4, MNHN); "Seychelles [and] Madagascar", H. Cuming collection (2, BMNH); "Madagascar(?)", M. Larrey, 1874 (1, MNHN); "Madagascar", M. Ballot (1, MNHN); "Madagascar" (1, MNHN); "Indian Ocean", H. Fischer (1, MNHN); "In- dian Ocean", Staadt colln (2 MNHN); "Ceylon", ex- Gould coUn (1 USNM); "Cochin China", M. Harmand, 1876 (2, MNHN); "Penang (?)", ex Marie and Sowerby collections (7, BMNH); "Penang" (2, LACM); "Austra- lia". Steams colln (1, USNM). Distribution: Reunion and (based on Sganzin, 1843) Mauritius. Living on faces of lava rock and boulders ex- posed to tlie open sea from low tide to 2 m depth (M. Jay, pers. comm.). Records from Madagascar and the Seychelles require confirmation. Specimens in old col- lections reputedly from Zanzibar, Coromandel Coast, Ceylon (Sri Lanka) are of uncertain provenance, whereas Penang, Cochin China (southern Vietnam) and Austrafia are extremely doubtful. Remarks: The shell oi Infundibuhim concavum is dis- tinctive among Trochoidea in the combination of broadly conical, weakly cyrtoconoid spire; flat, inwardly sloping base, conical umbilicus, and very strongly prosocline out- er Up. Among trochid animals examined or that have been recorded in the literature, the external anatomy of this species seems most similar to that of Tcctiis fcncs- tratiis (Gmehn, 1791) (Hickman and McLean, 1990: fig. 55A). Infundibulum tomlini (Fulton, 1930) (Figures 4-7) Pobjdonta {Infundibidum) concava: MeMl and Standen, 1895: 125 (not Gmelin, 1791). Trochtts (Infundibulum) tomlini Fulton, 19.30: 16, pi. .3, fig. 5. Trochus tomlini: Kaicher, 1979: card 2162. Trochus mnculatus: Marshall, 1979: 535; Marshall, 1981: 90 (not Linnaeus, 1758). Trochus (Infundibulum) sp.: Brook, 1998: 256. Description: Shell (figures 4-7) up to 36.0 mm wide, stout, broadly conical, wider than high, weakly cyrtocon- oid, base excavate; aperture strongly tangential, as wide as shell, narrow umbilicus; periphery sharply angulate, not stellate, almost flush against a flat surface. Proto- conch white. First 3.5 teleoconch whorls CTeen with red spiral bands; 2 or 3 continuous median bands and 1 su- prasutural band on 1st 2 whorls; spiral cords spotted and streaked with red on 3rd whorl. Subsequent spire whorls deep duU greenish or reddish grey with subsutural band of white spots or streaks. Base considerably paler than spire, inner third (approximately) porcelain-white. Pro- toconch 200 fjLm wide, tip narrowly tapered, sculpture unknown (eroded). Teleoconch of up to 7.5 whorls. Ear- ly spire whorls moderately convex, later whorls moder- ately or weakly convex; sculptured with rounded spiral cords and rounded, prosocUne, non-coUabral axial cos- tae, interspaces narrower; intersections with strong, bluntly rounded nodules; fine, crowded coUabral growth lines throughout. Spiral cords numbering 7 per spire whorl; adapical spiral broadest in most specimens, al- most fusing with adjacent spiral in some specimens; pe- ripheral and adjacent spiral narrowest, similar; other spi- rals similar, peripheral spiral partly covered by succeed- ing whorls. Excavated base broad, weakly convex, evenly curving into narrow steep-sided umbilicus, base of which is fiUed with callus. Basal spirals much finer than spire spirals; outer (pigmented) part of base of most speci- mens with 7 or 8 weakly nodular spirals, typically 1 sec- ondary spiral in each interspace, some interspaces in some specimens with 2 secondary spirals; 6-8 smooth spirals on inner (white) part of base, smooth, similar. Aperture ovate-trapezoidal. Columellar and basal fips smoothly continuous, thickened adapicaUy, sharply re- tracted to insertion within umbilicus. Rounded spiral ridge at base of outer lip. Animal unk-nown. Type data: Originally figured syntype BMNH 1930.4.2.3 (height 19.0 mm, diameter 26.6 mm); syntype National Museum of Wales Cardiff (Trew, 1984). Fulton (1930) stated that he had seen five specimens and did not segregate a holotype. The specimens could not be locahzed and evidently originated from somewhere in the tropical southwest Pacific. Other material examined: Titi Knob (Trig Sentinel), Boat Cove, Raoul Island, Kermadec Islands, cemented tuffaceous conglomerate (Pleistocene), W. R. B. Oliver (1, NMNZ); DavTell Islet. Raoul Island, hard sandy tuff (Pleistocene), F.'j. Brook, June 1991 (2, Auckland Insti- tute and Museum); Upolu, Western Samoa, R. W Tate (7, NMNZ); Lifou, Loyalty- Islands, J. Brazier, 1873 (4, BMNH); Vanuatu, C. Bertie, 1895 (1, AMS); Erroman- go, Vanuatu, H. A. Robertson, 1898 (3 in 2 lots, AMS); "Red Sea", E. C. Freeman collection (3, BMNH); "Red Sea" (4, LACM); "Penang" (1, NMNZ); "Penang, ex Sowerby" (1, NMP). Distribution: Raoul Island, Kermadec Islands (Pleis- tocene): Upolu, Western Samoa (record plausible be- cause R.W. Tate coUected extensively there); Aneiteum and Erromango, Vanuatu (records plausible because B. A. Marshall. 2000 Page 153 there are extensive stretches of exposed rockv' shore), and possibly Lifou. Specimens in old collections reput- edl\ from the Red Sea and Penang are of extremely doubtful provenance. Remarks: Compared with Infundihiiliim concamiin. I. tonilini differs in attaining smaller size (nia.\imum di- ameter 36.0 mm instead of 56.0 nnn). in being more broadly conical, in ha\'ing more strongly convex whorls, and in having much stronger axial costae and nodules on the spire, especially after the shell is about 6 mm wide. In /. concaium the spiral cords and nodules progres- sivelyweaken and become obsolete when the shell is 12- 15 mm wide, while in /, touiliui the spiral cords pro- gressively enlarge throughout. One lot (NMP) labelled "Penang, ex Sowerby", sug- gests origin from the London shell dealers Messrs. Sow- erby and Fulton, who advertised Trochus concavus in their 1908 price list. Other "Penang" specimens of I. coucavtiin and /. totulini in old collections mav well have originated from this source. The Pleistocene specimens from Raoul Island (figure 7) are indistinguishable from the syntype (figures 4-6) and other Recent specimens. /. tomlini is absent from the Recent fauna of the Kermadec Islands (Brook and Marshall, 1998). ACKNOWLEDGMENTS For loan or gift of specimens I thank Philippe Bouchet (Museum National d'Histoire Naturelle, Paris), Dai Her- bert (Natal Museum, Pietermaritzburg), Maurice Jay (La Reunion), Alan Kabat (National Museum of Natural His- tory, Washington D.C.), Ian Loch (Australian Museum, Sydney), Jim McLean (Los Angeles County Museum of Natural History), Robert Moolenbeek (Zoological Mu- seum, Amsterdam), David Reid and Kathie Way (The Natural Historv^ Museum, London), and Gary Rosenberg (Academy of Natural Sciences of Philadelphia). Thanks to Dai Herbert and Jim McLean for comments on the manuscript, Norman Heke (Museum of New Zealand) for the photography, and to Robert Thompson (Victoria Universitv', Wellington) for access to the scanning elec- tron microscope. LITERATURE CITED Blainville, H. M. D. de 1825-1827. Manuel de malacologie et de conchy liologie. . , LevTaiilt, Strasbourg, 664 pp. Brook, F. J. 199S. Stratigraphy and Paleontology of Pleistocene submarine volcanic-sedimentarv sequences at the north- em Kermadec Islands. Journal of the Royal Society of New Zealand 2S;235-257. Brook, F. J. and B. A. Marshall. 1998. Checklist of benthic coastal marine Mollusca from the northern Kermadec Is- lands, pp. 210-233 (Appendix) In: Brook, F. J. The coastal molluscan fauna of northern Keniiadec Islands, southwest Pacific Ocean. Journal of the Royal Society of New Zea- land 28:185-2.33. Chemnitz, J. H. 1781. Neues Systematisches Conchylien Cab- inet 5. Raspe, Nuniberg, 324 pp. Chenu, J. C. 1859. Manuel de conchvliologie et de paleonto- logie conchyliologique 1. Masson, Paris, 508 pp. Cossmann, M. 1918. Essais de paleoconchologie comparee 11. Cossmann, Paris, 388 pp. Deshayes, G. P. 1863. Catalogue des Mollusques de I'ile de la Reunion (Bourbon) 7m. Maillard, L. (ed.) Notes sur I'ile de la Reunion. Annexe E, Dentu, Paris, pp. 1-144. Dillwyn, L. W. 1817. A descriptive catalogue of Recent shells, arranged according to the Linnaean method 2. Arch, Lon- don, pp. 581-1092. Drivas. J. and M. Jay, 1988. Coquillages de la Reunion et de I'ile Maurice. Delachaux et Niestle, Paris, 159 pp. Fi.scher, P. 1875-1880. Species general et Iconographie des co- quilles vivantes, comprenant la collection du Museum d'Histoire naturelle de Paris, la collection Lamarck, celle du Prince Massena. .11. Bailliere, Paris, 480 pp. Fulton, H. C. 1930. Descriptions of new species of Fiisinus. Biplex, Trochus. and Biisliia. Proceedings of the Malaco- logical Society of London 19:16-17. Gmelin, J. F. 1791. Caroli a Linne . . . Systema naturae per regna tria naturae, secundum classes . . . Editio decima tertia, aucta, reformata. 1 (6). Vermes testacea. Beer, Lip- siae, pp. .'3021-4120. Kaicher, S. D. 1979. Card catalogue of world-wide shells. Pack 21. Trochidae 1. Kaicher, St. Petersburg, Florida, cards 2072-2177. Keen, A. M. and L. R. Cox. 1960. 7/i. Moore, R. C. (ed) Trea- tise on invertebrate paleontology. Part 1. Mollusca 1. Geo- logical Society of America and University of Kansa.s, Lawrence, pp. 246-275. Marshall, B. A. 1979. The Trochidae and Turbinidae of the Kermadec Ridge. New Zealand Journal of Zoology 6:521- 552. Marshall, B. A. 1981. Boat Cove Formation: macrofossils hi: Lloyd, E. F. and S. Nathan (eds.) Geology and Tephro- chronology of Raoul Island, Kemiadec Group, New Zea- land. New Zealand Geological Sur\'ey Bulletin 95, pp. 90- 91. Martens, E. von 1880. Die Mollusken der Maskarenen und Seychellen auf Gniud der von Professor Karl Mohius ges- ammelten Mollusken In: Mohius, K. (ed.) Beitriige zur Meeresfauna der Insel Mauritius und der Seychellen. Gutmann'schen, Berlin, pp. 179-.343. Melvill, J. C. and R. Standen 1895. Notes on a collection of shells from Lifu and Uvea, Loyalt)' Islands, formed by the Rev. James and Mrs. Hadfield, with list of species. Journal of Conchology 8:84-1.32. Montfbrt, D. de 1810. Conchyliologie systematique, et classifica- tion mediodique des coquilles. . . 2. Schoell, Paris, 676 pp. Oliver, W. R. B. 1915. The Mollusca of the Kermadec Islands. Transactions of the New Zealand Institute 47:509-568. Philippi, R. A. 1846-1855. Systematisches Conchylien-Cabinet von Martini und Chemnitz. Neu herausgegeben von ver- voUstaeudigt \on H. C. Kuester (iiach dessen Tode fort- gesetzt von W. Kobelt) 2 (3). Bauer and Raspe, Nurem- berg, 372 pp. Pilsbry, H. A. 1889-1890. Manual of Conchology; structural and systematic. With illustrations of the species. Volume 11. Academy of Natural Sciences, Philadelphia, 519 pp. Sganzin, V. 1843. Catalogue des cotjuilles troiuees aux ties de France, de Bourbon et de Madagascar Memoires de la Societe d'Histoire Naturelle de Strasbourg 3 (2):l-.'30. Swainson, W. 1840. A treatise on malacology, or shells and Page 154 THE NAUTILUS, Vol. 114, No. 4 shellfish. Longman, Omie, Brown. Green, Longmans and Prosobranchia. In: Schindewolf, O. H. (ed.) Handbuch Taylor, London, 419 pp. der Palaozoologie 6. Bonitraeger, Berlin, pp. 24-^80. Thiele, J. 1929. Handbuch der Systematischen Weichtierkunde Wood, W. 1825. Index testaceologicus; or a catalogue of shells, 1(1). Fischer, Jena, pp. 1-.376. British and foreign, arranged according to the Linnean Trew, A. 1984. The Melvill-Tomlin Collection. Part 30. Tro- system; with the Latin and Enghsh names, references to chacea. Handlists of the moUuscan collections in the De- authors, and places where found. Illustrated with 2300 partment of Zoology, National Museum of Wales, 94 pp. figures. Taylor, London, 188 pp. Wenz, W 1938. Gastropoda. Teil 1 (2): Allgemeiner Tell und THE NAUTILUS 114(4):155-160, 2000 Page 155 On the tcLxononiic placement of Uiiio ochraceus Say, 1817 in the genus Ligumia (Bivalvia: Unionidae) Douglas G. Smith Department of Bi()log^ and Graduate Program in Organisniic and Exolutionan' BioloijN Uni\ersitv of Massachusetts Amherst,' MA 01003-5810 USA d1 mm in length. The mantle edge bearing these papillae is slightly lamellate and there is no indication of a flap or lacera- tions, as is also the case with L. nasuta, L. subrostrata, and L. recta. The papillae of L. ochracea appear at a young age, about two years as evidenced by prominent grov\4h marks on shells of 20-25 mm in length. At this age (shell size), the papillae are not as closeK- set as in the adult and there is no evidence of crenulation, a flap, or a "lacerated" structure (sensu Ortmann, 1912). The mantle edge of L. ochracca is pigmented with a cream, orange-ochre, or hght-grev background and var- iegated with darker grev. Although most of the darker pigment is distributed near the posterior portion of the mantle edge (figure 1), no specific pattern is e\ident. In this regard, L. ochracea is intermediate between L. na- suta, which has verv dark pigment throughout and a dis- cernible pattern, and L recta and L. subrostrata. in D. G. Smith, 2000 Page 157 3f^' Figure 2. Section of mantle edge of female Lii^iimia ocJira- cca (sliell lengtli = 55 mm) from the Connecticut Ri\er, Con- necticut. Scale line = 50 ni. Figure 3. Same as figure 2, different specimen (shell length = 72 mm). Scale hne = 50 ni. Figure 4. Glochidia of Liotania ochracea (left) and Leptoclea Jrajiilis (right). Scale line = 200 m. which the mantle edge is a siightK" variegated, pattern- less, light- to dark-gray. The mantle margins o( Ligiimia subrosirata and L. recta differ in that the inner edge of the mantle of L. suhrostrata has a very dark-brown bor- der, a feature absent in L. recta. Both species possess a densely pigmented "eye spot" at the base of the inhahint aperture. In L. recta, the spot is small and not swollen; in L. suhrostrata, however, the spot is actually a shghtly thickened pad, almost lobe-like in a few specimens. The function of these spots is as yet unclear, although it is supposed that thev act as some sort of light rece]itor. No such spots are appari'Ht externalK in either L iiasulti or L. ochracea. E.\aniined specimens ol Leptodea fragiUs comply with Ortmann's (1912) (hagnosis for the most part. In one female, a few small papillae extend slightly anteriorly from the inhalant apertural papillae but subsequently disappear completely. In the remaining specimens, no extra-apertural papillae are evident. The crenulation re- ported by Ortmann (1912) is actually no more than a shght sinuation of the margin, the edge being uninter- rupted whatsoever for the length of the mantle. The mantle margin is slightly more lamellate in the female than in the male, but there is httle difference between the sexes. Although the specimens have been in alcohol for several years, there is no indication of concentration of pigment in the posterior region