1e, NAUTILUS Volume 122, Number1 March 28, 2008 ISSN 0028-1344 A quarterly devoted to malacology. EDITOR-IN-CHIEF Dr. José H. Leal The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 BUSINESS MANAGER Mary Jo Bunnell The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 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 Biologie des Invertébrés Marins et Malacologie Muséum National d'Histoire Naturelle 55, rue Buffon Paris, 75005 France Dr. Robert H. Cowie Center for Conservation Research and Training University of Hawaii 3050 Maile Way, Gilmore 409 Honolulu, HI 96822 Dr. Robert T. Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. Eileen H. Jokinen 8234 E. North Shore Road Sault Ste. Marie, MI 49783 Dr. Douglas S. Jones Florida Museum of Natural History University of Florida Gainesville, FL 32611-2035 Dr. Harry G. Lee 4132 Ortega Forest Drive Jacksonville, FL 32210 Dr. Charles Lydeard Biodiversity and Systematics Department of Biological Sciences University of Alabama Tuscaloosa, AL 35487 Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa P.O. Box 467 Wellington, NEW ZEALAND Dr. James H. McLean Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Paula M. Mikkelsen Paleontological Research Institution 1259 Trumansburg Road Ithaca, NY 14850 Dr. Diarmaid O Foighil Museum of Zoology and Department of Biology University of Michigan Ann Arbor, MI 48109-1079 Dr. Gustav Paulay Florida Museum of Natural History University of Florida Gainesville, FL 32611-2035 Mr. Richard E. Petit P.O. Box 30 North Myrtle Beach, SC 29582 Dr. Gary Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr Angel Valdés Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 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 SUBSCRIPTION INFORMATION The subscription rate per volume is US $47.00 for individuals, US $80.00 for institutions. Postage outside the United States is an additional US $10.00 for regular mail and US $25.00 for air delivery. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 1580, Sanibel, FL 33957, USA, (239) 395-2233. Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should inclide both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is published quarterly by The Bailey- Matthews Shell Museum, 3075 Sanibel-Captiva Road, Sanibel, FL Periodicals postage paid at Sanibel, FL, and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS P.O. Box 1580 Sanibel, FL 33957 CONTENTS NAUTILUS Volume 122, Number 1 March 28, 2008 ISSN 0028-1544 Philippe Bouchet Richard E. Petit Vittorio Garilli Diego G. Zelaya New species and new records of southwest Pacific Cancellariidae (Gastropoda) Fe oa. Qala ad deol Maio HG. gnerd Malan Gwe eR oe Re bE as | On some Neogene to Recent species related to Galeodina Monterosato, 1SS4, Galeodinopsis Sacco, 1895, and Massotia Bucquoy, Dautzenberg, and Dollfus, 1884 (Caenogastropoda: Rissoidae) with the description of two new Alvania species from the Mediterranean Pleistocene 2... 0.0... 0000 ee 19 Reallocation of Cyamiocardium crassilabrum Dell, 1964, into Perrierina Bernard, 1897 (Bivalvia: Cyamiidae) ......0.000000 0000000 eee eee 52 Notice ................ MBLWHOI! Library JUN 1 @ “UAB Woops HOLE Massachusetts 02543 THE NAUTILUS 122(1):1-1S, 2008 Page | New species and new records of southwest Pacific Cancellariidae (Gastropoda) Philippe Bouchet Muséum National d’Histoire Naturelle 55, rue Buffon 75005 Paris, FRANCE pbouchet@mnhn.fr re.petit@att.net Richard E. Petit S06 St. Charles Road North Myrtle Beach, SC 29582 USA ABSTRACT Fifteen species of Cancellariidae referable to the genera Zead- mete, Admetula, Fusiaphera, Nipponaphera, and Trigonostoma are reported from depths between 200 and 700 m in New Caledonia and other island groups in the southwest Pacific. Twelve are new species: Zeadmete bathyomon new species, Zeadmete physomon new species, Zeadmete bilix new species, Admetula affluens new species, Admetula marshalli new spe- cies, Admetula bathynoma new species, Admetula lutea new species, Admetula emarginata new species, Nipponaphera argo new species, Nipponaphera agastor new species, Nippona- phera tuba new species, and Trigonostoma tryblium new spe- cies. All the Recent nominal species of Fusiaphera described from localities throughout the meee Pacific area are considered to be conspecific, the senior name being Fusiaphera mac- rospira (Adams and Reeve, 1850), now with ten synonyms. The ranges of Nipponaphera nodosivaricosa (Petuch, 1979) and Trigonostoma thysthlon Petit and Harasewych, 1987, are ex- tended to the South Pacific. INTRODUCTION The present paper is a continuation of our study of the deep-water cancellariid fauna of the Southwest Pacific, based on the material originating from recent expedi- tions in New Caledonia, Vanuatu, Fiji, Tonga, Wallis & Futuna, and the Solomon Islands. In a previous paper (Bouchet and Petit, 2002), we described the new genus Mirandaphera and nine new species in the genera Afri- cotriton, Merica, Sveltia, and Nipponaphera. We here deal with 15 species (12 new) in the genera Zeadmete, Admetula, Fusiaphera, and Trigonostoma, and add tur- ther species in Nipponaphera. Our review of the deep- water cancellariid fauna so far sampled in the southwest Pacific will be complete after a third paper (in prepara- tion) dealing with the genera Brocchinia, Microcancilla, and Gergovia. In addition, the cancellariid fauna of New Caledonia includes shallow-water to offshore species in the genera Scalptia (5 species) Tritonoharpa (several species), as well as the rediscovered endemic Merica semperiana, which we intend to deal with separately. Cancellariid radulae are not known to provide dis- criminating species-level characters, and we thus did not attempt to systematically examine them when we had live-taken specimens available. Much of the material re- ported in this series was collected in the 19S0—1990s and, at the time, fixed in formalin and then rinsed and dried. It is thus not adequate for nucleic-acid sequencing. More recent expeditions generate new material that is specifi- cally put aside for barcoding. Our treatment of the can- cellariid fauna is thus currently restricted to a descr iption of the shells, including the protoconch, but we may ex- pect that in the future it will be possible to test some of our species limits through molecular characters. MATERIALS AND TEXT CONVENTIONS In the lists of type and other material examined, indi- vidual lots in MNHN are unambiguously designated by the combination of cruise acronym (capitalized) and sta- tion number. DW (for Drague Warén) refers to dredge hauls, CP (for Chalut & Perche) to beam trawls; lv refers to live-taken specimens, dd to empty shells; spms to in- dividuals that cannot be unambiguously assigned to one of these two categories (esse -ntially commercially ob- tained specimens) Institutional acronyms are: AMNZ: Auckland Museum Auckland, New Zealand: BMNH: The Natural History Museum, London, Uk; DMNH: Delaware Museum of Natural History, Wilmington, Delaware, USA: MNHN : Museum National d’ Histoire Naturelle, Paris, France; NM: Natal Museum, Pieterma- ritzburg, South Africa, NMW: National Museum of Wales, Cardiff, UK: NSMT: National Science Museum, Tokyo, Japan; USNM: National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, USA; WAM: Western Australian Museum, Perth, Australia. SYSTEMATICS Family Cancellariidae Forbes and Hanley, 1851 Genus Zeadmete Finlay, 1926 Type Species: Cancellaria trailli: Hutton, 1873, by original designation. Recent, New Zealand. Page 2 THE NAUTILUS, Vol. 122, No. 1 Discussion: The genus Zeadmete was Sas d_ by Finlay (1926: 429) who later (1930b: 242) considered it to be a subgenus of Oamaruia Finlay, 1924 from the Lower Miocene of New Zealand. Powell (1979: 224) also treated Zeadmete as a subgenus of Oamaruia. However, Garrard (1975: 44) and Wilson (1994: 173) restored Zeadmete as a full genus, as did Petit and Harasewych (2000: 151), who gave a brief discussion of its possible relationship to other taxa. Among other differences, the type species of Oamaruia, Admete suteri Marshall and Murdoch, 1920, has strong columellar folds whereas Zeadmete has weak, almost obsolete, folds. The genus Zeadmete, as interpreted here, occurs in the Miocene to Recent faunas of New Zealand and in the Recent faunas of South Africa, Fiji and New Caledonia (Z. kulanda Garrard, from Australia, is probably an Ip- hinopsis.) Expedition material in MNHN also contains an undescribed species from the Solomon Islands and another one from New Caledonia, both represented by specimens too imperfect to be named. All live offshore in 300-600 m, with New Zealand records as shallow as 65 meters. Zeadmete bathyomon new species (Figures 1-2) Description: Protoconch glassy, smooth, erect, of about 1.3 whorls, diameter 900 xm. Teleoconch of four whorls. Transition to teleoconch marked by a sharp growth line and a spiral cord forming shoulder angle, shortly followed by onset of weak sak sharp axial bs, forming small nodes at strengthening shoulder angle. Ad- ditional spiral cords appear before end of first teleoconch whorl. Shoulder angle prominent, bearing minute coro- nations formed by intersection of spiral cords and axial ribs. Sutural ramp strongly concave, bearing five to seven fine, low spiral cords. Suture attachment orthogonal. Shoulder raised, bearing three to five closely spac ed spi- ral cords. About 20+ wider, low spiral cords anterior to shoulder angle, extending down onto base. Fine, low, evenly space d axial ribs extending from shoulder angle to anterior end of shell; about 25 on last whorl; interspaces between axial ribs bearing closely packed fine growth lines. Low, narrow varices, formed. only by a slight raising of shell surface, at about 120° increments on final w horls, Oute r lip smooth, faintly sinuate adapically, curving back n prosocline direction abapically. Aperture elongate, ne aes elliptical, without lirations inside outer lip. Pa- rietal area ai very thin, narrow callus. Columella with three folds, the posterior two weak; anterior fold broad, situated on small siphonal fasciole, forming edge of in- ductural callus, which then forms edge of distinct sipho- nal canal. Exterior beige, rims of varices pale straw- yellow. Type Material: Wa. Holotype MNHIN 20498, 10.2 x 4.8 Type Locality: South of New Caledonia, 22°17’ S, 167°12’ E, 390 m [VAUBAN 1978-79: sta. 3]. Material Examined: Norfolk Ridge, south of New Caledonia.—VAUBAN 1978-79: sta. 3, 22°17’ S, 167°12' E, 390 im, 1 lv (Figure 1).—BIOCAL: sta. DW77, 22°15’ S, 167°15' E, 440 m, 1 dd—BATHUS 2: sta. DW719, 99°48! S, =e E, 444-445 m, 1 lv.— SMIB 8: sta. DW166, 23°38’ S, 167°43" E, 433-450 m, Figures 1-5. Zeadinete. 1-2. Zeadmete bathyomon new species. 1. Holotype, height 10.2 mm; New Caledonia, Norfolk Ridge, 22°17' S, 167°12’ E, 390 m [Vauban 1978-79 sta. 3]. 2. 23°3 167°43' FE, 433-450 m [SMIB S sta. DW166]. 3, 4. Zeadmete physomon new speci holotype height 15.1 mm, Fiji, 19°01’ S, 178°25' E, 500-516 m [BORDAU 1 sta. DWI14SS]. 5. Zeadmete bilix new specie holotype height 13.2 mm;, New Caledonia, Norfolk Ridge, 23°02’ S$, 168°16’ E, 335 m [SMIB 5 sta. DW98] P. Bouchet and R. E. Petit, 2008 Page 3 1 dd (Figure 2); Sta. DWI167, 23°38" S, 167°43' E, 43¢ 452 m, | dd—NORFOL 7 l: sta. DW1666, 23°42’ S. 167°44' E, 469-S60 m, 2 i —NORFOLK 2: DW 2024, 23°28’ S, 167°5 1’ E, 370-371 m, 1 dd. Etymology: From the Greek bathus, deep, combined with omos, shoulder, for the deeply concave sutural ramp. To be treated as a noun in apposition. Distribution: Known only from south of New Cale- donia, alive in 390-444 m. Remarks: — In Zeadmete finlayi Powell, 1940, the shoul- der is also raised but the sutural 1 ramp is not concave. In Z. bathyomon, the shoulder is even with, or raised above, the suture. Also, Z. finlayi has clathrate sculpture only on the adapical half of the last whorl, whereas the last whorl is entirely clathrate in Z. bathyomon, The aperture is two-thirds of the shell height in Z. finlayi, but in Z. bathyomon the aperture is just over half the shell height. The only other Zeadmete species we are aware of swith a sutural ramp that does not slope abapically is an unde- scribed species represented by two broken, worn speci- mens from a nearby station on the Norfolk Ridge [BERYX 11: sta. DW35, 23°33’ S, 167°16' E, 550-570 m], which we leave undescribed because of the poor condition of the specimens. It differs from Z. bathyomon in having a shorter spire and a flatter sutural ramp that is devoid of spiral cords. It also has a larger protoconch with a diameter of 1100 jum. Zeadmete physomon new species (Figures 3+) Description: Protoconch smooth, glossy, of 1.5 whorls, diameter 1250 wm. Transition to teleoconch marked by weak axial rib and faint spiral cords, both of which increase in strength rapidly. Teleoconch of four whorls, shell thin. Spiral cords flat, evenly spaced, with interspaces Ss lightly narrower than cords, about five on sutural ramp and 20+ anterior to shoulder angle. Axial ribs extending from suture to anterior end of te Teaeoiuoh, interspaces consider rably broader than ribs; about 20 ribs on last whorl. Sutural ramp slightly convex, sloping up to impressed suture. Aperture elongate- elliptical, only weakly angulate at shoulder, smooth within. Outer lip sinuous adapically but becoming prosocline anterior to periphery. Parietal wash fadisanct on holotype, distinct on one paratype. Columella almost vertical, bearing three folds, posterior fold on top of siphonal fasciole. Anterior folds callused, widely separated, anterior one almost obsolete, forming edge of short siphonal canal. Exterior cream with ill-defined pale brown band on pe- riphery of spire whorls; some ribs on last whorl have brownish tint. Type Material: Holotype MNHN 20519 (13.1 « 6.5 mm) and 6 paratypes MNHN 20520-20521. Type Locality: Lau Ridge, Fiji, 19°01’ S, 175°25' E, 500-516 m [BORDAU 1: sta. DW1488]. BORDAU |] Material Examined: — Fiji. sta. DW14S6, L9°O1' S, 178°26' EB, 395-540 m, 1 dd paratype MNHN 20520.— Sta. DW1488, 19°O1' S, 178°25' EB, 500-516 m, 6 dd, holotype MNTIN 20519, paratypes MNHN 20521 (Figures 5-4), Etymology: From the Greek hae inflated, and omos, shoulder, Tor the appearance of the sutural ramp. To be treated as a noun in apposition. Distribution: listed above. Known only from Fiji at the two stations Remarks: Protoconch diameter in sigh ranges from 900 to 1300 jum. Zeadmete physomon differs from Z. bathyomon in having narrower cael cords with wider interspaces, fewer, more widely spaced axial ribs, and a slightly convex rather than strongly concave sutural ramp. Also, in Zeadmete physomon the axial ribs remain distinct below the periphery. Zeadmete bilix new species (Figure 5) Description: —Protoconch prominent, smooth, glassy, of 1.1 whorls, diameter $75 jum. Transition to teleoconch marked by onset of axial ribs and spiral cords. Teleo- conch high-spired, of five whorls. Spiral cords narrow, interspaces broader than cords. About three to four fine spiral cords on sutural ramp; eight cords anterior to shoulder angle on penultimate who. Last whorl with about 17 narrow spiral cords, one of which forms shoul- der angle and another, only slightly more prominent than those adjacent, is just posterior to periphery, giving te- leoconch a weakly biconic appearance. Axial Tbs A fine, spaced almost e qual to spacing of spiral cords, rendering teleoconch surface evenly reticulate; about 55 on last whorl. Sutural ramp narrow. Aperture narrowly elliptic. Outer lip orthocline adapically, becoming prosocline only at anterior end. Columella almost straight, bearing three folds: adapical one on siphonal fasciole, other two on a broad heavy callus; anterior one obsolete, forming edge of short siphonal canal. Periostracum pale brown, shell pale straw yellow. Type Material: Holotype (dd) MNHIN 20499 (13.2 x 5.6 mm). Type Locality: Norfolk 1 AXidge, south of New Cale- donia, 23°02’ §, 168°16' E, 335 m[SMIB 5: sta. DW9S]. Material Examined: (Figure 5). Only known from the holotype Etymology: Latin adjective bilix, having a double thread, with reference to the two strong spiral cords around the shoulder angle. Distribution: type locality. Known only from New Caledonia at the Remarks: = Zeadmete bilix is placed in Zeadmete based on its columellar structure, which is identical to that of other Zeadmete species, but it may be immediately dis- Inge Page 4 THE NAUTILUS, Vol. 122, No. 1 tinguished from its congeners by its elongate shape and relatively short aperture. Zeadmete bilix superficially resembles the buccinid ge- nus Iredalula Finlay, 1926, but species of Iredalula have a recurved, notched siphonal canal and lack columellar folds. Genus Admetula Cossmann, 1889 Type Species: Buccinwm evulsum Solander, 1766, by original designation. Eocene, British Isles. Discussion: Admetula is often placed in the synonymy of Bonellitia Jousseaume, 1887 ( (see discussions in Sacco, 1894: 42: Cossmann, 1899: 33; Davoli, 1982: 62; Ver- hecken, 1986: 33; but not na a 2007: 286), but we regard Bonellitia as distinct, based on the angled whorls and muricated sculpture. We refer to Landau, Petit, and Marquet (2006) for further discussion. The genus Admetula is well represented in Paleogene and Neogene Tethyan faunas and in the Recent fauna along continen- tal margins at depths ranging from 75-700 m (Petit and Harasewych, 1991: 181). Many species of Admetula appear superficially very similar, ae cially based on published illustrations, yet can easily be distinguished when directly compi ared. Three discrete * “subgroups” can be recognized in the ge- nus. One consists of small attenuate species such as A. cornidei (Altimira, 1978), A. epula Petit and Harasewych, 1991, and A. afra Petit and Harasewych, 2000. Larger, wide, rather thin-shelled species s such as A. bayeri Petit, 1976, and A. emarginata (described herein) form a sec- ond group, with a third, intermediate group composed of thick shells with an anterior constriction of the last whorl such as A. vossi Petit, 1976, and A. deroyae (Petit, 1970). Early Tertiary species usually have well-formed varices at irre ‘cular intervals, a feature not found on Recent spe cles Admetula affluens new species (Figures 6-8) Description: Protoconch corroded on holotype, in other specimens consisting of 0.8 whorls, diameter 775 wm, shiny, smooth apically, with six raised threads on ab: ipical part. Transition to teleoconch distinct, marked by onset of te : ‘oconch sculpture. Teleoconch spire high, spire angle 47 a ack consisting of 6.25 whorls, with ae of evenly spaced, narrow axial ribs crossed vy spiral cords forming small nodes at intersections. About 15 axial ribs on penultimate whorl, about 1S on ast whorl. About 12 primary spiral cords on last whorl, with spac ing ¢ qui al to that of axial ribs on shoulder and veriphery, more crowded on base; four to seven second- ary spire al cords in each inte rspace. Outer lip sharp yrosocline. \perture bea lirations. Parietal area with very thinly applied callus. Columella sloping, bearing two srominent folds extending to edge of inductural callus, with a third broad siphonal fold. Siphonal canal shallow, well defined. Exterior white with thick, pale olive-brown yeriostracum Type Material: Holotype MNHN 20500 (22 mm) and 4 paratypes MNHN 20501-20502. 7 ¥ 13:0 Type Locality: SW of Malaita, Solomon Islands, 09°46’ S, 160°53’ E, 611-636 m [SALOMON 1: sta. CP1S08]. Material Examined: Solomon Islands. SOLOMON 1: sta. CP1749, 09°21’ S, 159°56' E, 582-594 m, 1 dd.— Sta. CP1750, 09°16’ S, 159°55' E, 693-696 m, ie Sta. CP1751, 09°10’ S, 159°53’ E, 749-799 m, 2 lv (1 paratype MNHN 20501).—Sta. CP1793, 09°13" S, 160°0S' E, 505-510 m, 1 dd, 1 juv. dd (Figure 8).—Sta. CP1798, 09°21’ S, 160°29' E, 513-564 m, 2 lv, 1 dd.— Sta. CP1808, 09°46’ S, 160°53' E, 611-636 m, 1 lv (ho- lotype, Figures 6—7).—Sta. CP1859, 09°33’ S, 160°37’ E, 283-305 m, | lv, 2 dd (3 paratypes MNHN 20502). Total of 13 specimens. (Largest specimen: 26.3 x 15.6 mm.) Distribution: Known only from the Solomon Islands, alive in 305-749 in. Etymology: From the Latin affluwens, an adjective meaning abundant or copious, with reference both to its large size and relative abundance in the Solomon Archi- pelago. Remarks: Admetula affluens superficially resembles the specimen figured as A. garrardi (Petit, 1974) by Ha- segawa (2000: 585, pl. 291, figure 26) but differs by hav- ing narrower axial ribs, a more constricted base and a more twisted columella. It lacks the lirations within the outer lip that are present in A. garrardi. It is our opinion that the specimens figured as A. garrardi by Hasegawa (2000) and Veriecken { 1997: oan figs. 11-13) are not that species. Verhecken (1986: 34-35, figs. 1-2) exam- ined, redescribed and poued the holotype of A. garrardi and explicitly deseribe d its multispiral protoconch whereas A. affluens has a paucispiral protoconch. Of the other species of Admetula in the tropical south- west Pacific, A. affluens is more similar to A. marshalli, but differs by its larger adult size, less solid shell, much weaker spiral cords and lack of apertural lirations. Admetula emarginata new species (Figures 9-11) Description: —Protoconch glossy, white, of 1.1 whorls, diameter L000 jzm, with five wile ly spi aced spiral cords. Transition to teleoconch indistinct, protoconch cords continuing as teleoconch cords. Teleoconch of five whorls, suture impresse sd, with sculpture of numerous prominent, well-defined axial ribs crossed by spiral cords forming small nodes at intersections. About 14 widely spaced axial ribs on penultimate whorl and about 12 on last whorl, final one enlarged into terminal varix. About eight primary spiral cords on last whorl, more closely spaced than axial ribs, with one or more secondary spirals in each interspace and about six prominent secondary spiral cords on spire whorls below rounded shoulder angle. Shell thin, axial ribs visible through last whorl. dO 22.7 mm, Solomon Islands, 09°46S, 160 Figures 6-17. Admietula. 6-8. Admetula affluens new species. 6-7. Holotype, height 22.7 E. 611-636 m, [SALOMON 1 sta. CP1S80S]. 8. Protoconch, Solomon Islands, 09°13’ S, 160°OS’ FE, 505-510 m, [SALOMON I sta CP1793|. 9-11. Admetula emarginata new species. 9-10. Holotype, height 16 mm, Coral Sea, 20°03" S, 158°45' E, 315 m 5’ S$, 158°30.5' E, 230 m [CORAIL 2 sta. D31]. 12-13. Admetula MUSORSTOM 5 sta. 335]. 11. Protoconch, Coral Sea, 19°33.5 marshalli new species, holotype, height 14.7 mm, Fiji, 16°39’ S$, 179°57' W, 591-596 m [BORDAU 1 sta. CP1396], 14-15. Admetula 21°19’ S, 175°01’ W, 225-233 m [BORDAU 2 sta. DW1521]. 15. lutea new species 14. holotype height 13.1 mm, Tonga, 2 Protoconch Fiji, 18°09’ S, 178°39’ W, 290-300 m |[BORDAU | sta. DW1465]. 16-17. Admetula bathynoma new species 16. Holotype height 7.4 mm, New Caledonia, 22°52’ S, 167°23' E, 590-600 m [MUSORSTOM 4 sta. DW225]. 17. Protoconch, New 22°52’ §. 167°16’ E, 530-541 m [BATHUS 2 sta. DW720]. All protoconchs at the same scale, scale bar 500 jzm ( aledonia, 44 02 Dawa cn Page 6 THE NAUTILUS, Vol. 122, No. 1 Outer lip prosocline, sinuous, with everted stromboid notch just anterior to periphery and another everted notch near base. Aperture with nine lirations, visible in- ternally at level of terminal varix, not extending to edge of lip or deeply into aperture. Parietal area with very thinly applied callus. Columella sloping, with two promi- nent folds extending to edge of inductural callus and a third broad siphonal fold. Siphonal canal shallow, well defined. Shell white, with weakly defined bands of pale brown at shoulder and base, also brown behind outer lip. Type Material: Holotype MNHN 20503 (16.0 x 10.9 mim) and one paratype MNHN 20504. Type Locality: Coral Sea, 20°03" S$, 158°45" BE, 315 m |[MUSORSTOM 5: sta. 335]. Material Examined: Coral Sea. CHALCAL: sta. D31, 19°33.5' S, 158°30.5' E, 230 m, 1 dd ( (paratype, Figure 11)—MUSORSTOM 5: sta. 335, 20°03’ S, 158°45' E, 315 m, 1 dd (holotype, Figure 9- 10).—E BISCO: sta. CP2571, ae S, 158°45' E, 298-309 m, 1 dd. Etymology: From the Latin emarginatus after the sinuous, everted outer lip. Distribution: Known only from the Coral Sea near the Chesterfield Islands, empty shells in 230-315 m, Remarks: Admetula emarginata differs from other de- scribed species of Admetula in having diffused colora- tion. It is much like A. bayeri Petit, 1976, from the west- ern Gulf of Mexico in having at the periphery a notch, which is not present in the similar species A. bathynoma. Admetula emarginata also differs from A. bathynoma in having less closely spaced spiral cords and axial ribs. The overall effect of the sculpture on A. emarginata is a strik- ing pattern of horizontal rectangles crossed by fine spi- rals. Admetula marshalli new species (Figures 12-13) Description: —Protoconch glossy, brown, of one whorl, diameter $25 jrm, with three wide ly spaced spire al cords. Transition to teleoconch marked by ‘a change in color and onset of axial ribs. Teleoconch of 5.5 whorls. Suture im- pressed. Spiral cords closely spaced, about 14 primary cords on last whorl with one secondary cord in most interspaces, forming small pointed be ads where they cross axial ribs. One spiral cord creates a modest angle between periphery and narrowly rounded shoulder: About 14 rounded axial ribs on last whorl, only final one enlarged into a varix. Fine, densely packed growth lines on entire teleoconch. Outer lip thin, ve ry indistinctly notched just anterior to periphery and with a slight ever- sion of the siphon: ul canal, Ape rture with nine Tie ations, visible interior to terminal varix but not extending deeply within aperture or to outer lip Parietal area with thin but distinct callus. Columella concave, bearing three distinct folds that extend out to edge of inductural callus. Poste- rior fold largest separated from central fold by a deep depression two anterioi folds form a shelf bordering short, broad siphonal canal. Shell white, periostracum yellow-brown. Type Material: Holotype MNHN 20505 (14.7 x 9.0 mim) and 2 paratypes (all dd) MNHN 20506 (largest paratype: 17.0 « 9.7 mim). Type Locality: Fiji, Lau Ridge, 16°39’ S$, 179°57' W, 591-596 m, [BORDAU I: sta. CP1396]. Material Examined: terial. Only known from the type ma- Distribution: Known only from the Lau Ridge in Fiji, empty shells in 591-596 m. Etymology: Named after Bruce Marshall (Museum of New Zealand, Wellington), in appreciation for the stan- dards of his descriptions and illustrations of the mollus- can fauna of New Zealand. Remarks: The New Zealand species A. superstes (Fin- lay, 1930b) is similar in shape and size, but differs in having a translucent white rather than brown protoconch with numerous fine, close spiral threads (versus 3 widely spaced cords), in having weaker axial costae on the te- leoconch, in lacking a terminal varix and internal lirations behind the apertural rim at maturity, and in that second- ary spiral sculpture is es rably weaker or entirely absent. Additionally, superstes has a much more strongly developed Lents wcum, with prominent spines at the. summits of the axial lamellae. Admetula superstes is endemic to northern New Zealand, living on muddy substrata in 79-550 m off the northeastern (northern- most record at 35°08" S) and southwestern North Island. A similar species (possibly a local variant) occurs off Three Kings Islands. There is no material of similar Soe cies from Norfolk Ridge south of Norfolk Island i NMNZ Admetula lutea new species (Figures 14-15) Description: —Protoconch glassy, of slightly more than one whorl, diameter 975 jum, worn but with traces of spiral cords visible on terminal portion. Transition to te- leoconch rather indistinct. Teleoconch of about 4.7 whorls. Whorl profile regularly convex; suture im- pressed. Sculpture of prominent prosocline axial ribs crossed by weaker spiral cords; about nine to ten axial ribs on last whorl, eleven on penultimate whorl, some forming irregularly placed varices; four primary spiral cords on pe alan: ate whorl and about 12 on last whorl, with one to four secondary spirals in each interspace; spiral cords forming « slongate nodes where they cross axial ribs. Outer lip with very indistinct lirations, visible internally at level of te sininal varix, not exte nding either to edge of lip or deeply within aperture. Parie ied area with fhe callus. Inductura almost vertical. Columella bearing three folds: narrow anterior fold forming edge of short, recurved siphonal canal. Exterior ye sllow- bow: Periostracum thin, with low incremental lamellae and hairy projections on spiral cords. P. Bouchet and R. E. Petit, 2008 Page 7 Type Material: Holotype MNHN 20507 (13.1 « 7.8 mm) and one paratype MNHN 20508. Tonga, 21°19’ S, 175°01’ W, 225-233 2: sta. DW1521). Material Examined: — Fiji. BORDAU 1: sta. DW1465, 18°09" S, 178°39' W, 290-300 m, 2 dd.—Tonga. BOR- DAU 2: sta. DW a 21°19’ S, nigec W, 225-233 m, 1 dd ( (holotype, Figure 14).—Sta. CP1576, 19°42" S, 174°18' W, 253-263 m, 1 dd ( (paratype, Figure 15). Type Locality: m |[BORDAU Etymology: From the Latin adjective luteus, meaning ve sJlow, in reference to the color of the shell. Distribution: Known only from Fiji and Tonga at lo- calities cited above. Remarks: This new species differs from its congeners by its strongly prosocline axial ribs forming irregularly place sd varices and its ae sculpture ‘with numerous secondary cords. It is also distinguished from other Ad- metula species by its yellow color. A specimen of le tula lutea new species in the Petit collection (No. 2872) is said to be from 50-200 m north of Taiwan, but is a dealer's locality designation that needs to be confirmed. ; Admetula bathynoma new species (Figures 16-17) Description: —Protoconch glassy, white, smooth, of one whorl, diameter 975 jum. Transition to teleoconch marked by onset of axial ribs and spiral cords. Teleo- conch of about four whorls. Last whorl beari ing about ten to 14 rounded axial ribs, crossed by prominent spiral cords, about five to six cords on spire whorls and about ten on last whorl, with fine secondary spirals in the in- terspaces. Final axial rib enlarged into a varix. Small nodes formed at intersections where spiral cords cross axial ribs. Sutural ramp narrow, almost flat; suture slightly impressed. Outer lip prosocline, rounded, smooth, without lirations within. Parietal area without wash or callus. Columella slightly inclined, bearing two prominent folds extending out to edge of faductinl cal- lus; a third descending fold f forming edge of short sipho- nal canal. No siphonal fasciole. Last whorl well rounded. Shell white. Type material: Holotype MNHN 20509 (7.4 x 5.0 mm) and one paratype MNHN 20510. Type locality: Nortolk Ridge, south of New Cale- donia, 22°52’ S, 167°23' E, 590-600 m [MUSORSTOM 4: sta. DW225]. Material examined: Norfolk Ridge. BIOCAL: sta. DW46, 22°53’ S, 167°17' E, 570-610 m, 1 dd paratype) —MU pier 4: sta. DW225, 22°52’ S, 167°23' E, 590-600 m, 1 |v (holotype, Figure SMIB 5S: sta. ee ane 22°59'—23°00' S, 168°21'— 168°23’ E, ee 558 m, 1 juv. dd—BATHUS 2: sta. DW720, 22°52’ S, 167°16' = 530-541 m, 1 juv. dd (Fig- ure 17). Distribution: 491-610 mm. Known only from the Norfolk Ridge, in Etymology: From the Greek bathus, deep, and no- mos, place of living, treated as an adjective. Remarks: —Adimetula bathynoma resembles a juvenile A, marshalli but differs by being proportionally nar- rower, with less convex whorls and a larger protoconch (diameter 975-1000 jum versus $25 wm in A. marshalli). Admetula marshalli also differs in having pointed beads rather than small nodes formed at the intersections of the axial ribs and spiral cords. Kohn and Arua (1999: pl. 13, fig. 55) ilhistrated as C. icine Petit and Harasewych, 1986 an 8 mm high specimen of Admetula from the Early Pleistocene of Viti Levu, Fiji. Their specimen closely resembles the present species except that the Fiji fossil has lirations within the aperture. Genus Fusiaphera Habe, 1961 Type Species: Cancellaria macrospira Adams and Reeve, 1850, fixed herein to comply with ICZN Article 70.5. Recent, Japan. Discussion: Habe (1961a) proposed the genus Fusia- phera tor a species he identified as Cancellaria mac- rospira Adams and Reeve. Later in the same year, he (Habe 1961b) considered the species he had identified as C. macrospira to be a different species, which he then named Fusiaphera macrospiratoides Habe, 1961. The fact that the type species of Fusiaphera was based on a misidentified type species was noted by Verhecken (1986: 35), who stated that there is some confusion in the matter of the type species but did not make a definitive statement about the resolution of the problem. He did list, however, Fusiaphera macrospira (Adams and Reeve) as type species, as did Beu and Maxwell (1987: 55), who then stated “but misidentitie dP. Preally = Fusiaphera macrospiratoides Habe, 1961.” The Sonkision is Fe -mon- strated by the fact that Matsukuma, Okatani and Habe (1991: 179, pl. 111, fig. 2) figured the type specimen of F. macrospiratoides under the name F. macrospira. In the absence of a definitive statement by Verhecken and the queries used by Beu and Maxwell, it appears to be necessary to fix a type species to comply with Article 70.3. Cancellaria mac rospira Adams i Reeve is here selected as the type species of Fusiaphera Habe, 1961. Fusiaphera is distinguished by a slender, non- umbilicate teleoconch with irregular varices and a thick outer lip with a well-defined aaa canal under the shoulder. The aperture is much like that of Scalptia but with a straighter columella. The genus Fusiaphera ranges from South Africa across the Indian Ocean, north to Japan, to Australia, New Caledonia and Fiji. It thus ap- pears to be restricted to the Indo-Pacific area with the earliest known occurrence in the Miocene of Australia. Darragh (1970: 168) referred the Australian Miocene species Cancellaria epidromiformis Tate, 1889 and C. exaltata Tate, 1889, to Fusiaphera, a placement accepted Page § THE NAUTILUS, Vol. 122, No. 1 by Maxwell (1992: 166). However, three New Zealand Fusiaphera macrospira (Adams and Reeve, 1850) Eocene species, Uxia (?) marshalli Allan, 1926, Uxia (Figures 18—25) naroniformis Finlay, 1930, and Fusiaphera jenkinsi Max- well, 1992, place ed in Fusiaphera - Beu and Maxwell (1990) and ee ve 1992), are not considered by us to Cancellaria macrospira Adams and Reeve, 1850: 41, pl. 10, fig. be correctly placed. Also, Plesiotriton paytensis Olsson, 1930, from the Eocene of Peru, was tentatively placed in Fusiaphera by Beu and Maxwell (1987: 55) but it is here excluded from the genus. It is possible that a new genus bo Cancellaria wilmeri G. B. Sowerby I, 1SS1: 637, pl. 56, fig. Cancellaria pallida E. A. Smith, 1899: 313, text- fig. a Cancellaria producta G. B. Sowerby TH, 1903: 220, pl. 4, fig. 5. 3: Cancellaria | Trigonostoma) luscinia Melvill and oe n, 1903: will have to be erected to contain these Eocene taxa. 319, pl. 23, figs. 14-15. Many European Tertiary species of Unitas resemble Fu- Cancellaria exquisita Preston, 1905: 3, pl. 1, fig. 9. Sid} shera in form but the By lack a strongly delineated pos- Cancellaria tosaensis Habe, 1961a: Appendix 28, pl. 35, terior canal, fig. 21 Figures 18-25. Fusiaphera macrospira (Adams and Reeve, 1850). 18. Height 20.2 mm Philippines, Balicasag I., said to be from 130-230 m. 19. Height 30.1 mm, Japan, Mikawa. 20. inigeronpnat ees form, height 16.2 mm, Mikawa Issiki, Japan. 21. Holotype of Cancellaria wilmeri (BMNH 1881.5.20.30), height 11.8 mm. 22. tosaensis form, height 20.4 mm, Minabe Japan, 23. Height 15.0 m, New Caledonia, 19°35’ S$, 163°25' E, 48 m [LAGON sta. 1192]. 24, 25. Height 11.0 mm, New Caledonia, 19°06’ S$, 163°10' | 1m | LAGON sta. 542 P. Bouchet and R. E. Petit, 2008 Page 9 Cancellaria azumai Habe, 1961a: 72 20. Cancellaria macrospiratoides Habe, 1961b: 433, pl. 2 pl. 24, fig. 10. Fusiaphera dampierensis Garrard, 1975: 17, pl. 2, fig. 8. Fusiaphera eva Petit, 1980: 215, figs. 5, 6. Type Data: F. macrospira, C hina Sea, BMNH 1969347, lectotype designated by Verhecken (1986: 36); illustrated by Higo, Callomon and Goto (2001: 99). . Appendix 28, pl. 35, fig. 10; = C. wilmeri, Port Blair, Andaman Islands, holotype BMNH 1881.5.20.30, herein Fig. 21. C. pallida, 25 fms, off Bonaparte Archipelago, NW Australia, holotype BMNH 1891.11.21.96. C. producta, 40 fms, off mouth of Umhloti River, Natal, syn- types BMNH _ 1903.7.27.76; SAM—A339 (Giles and Gos- liner, 1983: 28). C. (T.) luscinia, 40 fms, Arabian Sea, 18°58’ N, 71°45" E, syntypes BMNH 1903.12.15, 101-102 (2 syntypes); NMW 1955.158.408. C. exquisita, Ceylon, holotype BMNH 1905.10.4.75. C. tosaensis, Kochi Prefecture, Shikoku, Japan, holotype NSMT-Mo 13287; illustrated by Higo, Callomon and Goto (2001: 99). C. macrospiratoides, Aiki Prefecture, Honshu, Japan, holotype NSMT-Mo 39781; illustrated by Higo, Callomon and Goto (2001: 99). C. azumai, Aiki Prefecture, Honshu, Japan, NSMT-Mo 13285a (illustrated by Higo, Callomon and Goto 2001: 99) is la- beled as type in NSMT but is not the figured specimen and is smaller than dimensions given for holotype. F. dampierensis, Delambre Island, Dampier Archipelago, northwestern Australia, holotype WAM 550-71. F. eva, west of central Bazaruto Island, southern Mozambique, holotype NM G4896. Description: Protoconch smooth, glassy, of two whorls with small initial nucleus, diameter 1000-1125 wm, indicating planktotrophic larval development. Tran- sition to teleoconch marked by sharp axial rib followed by additional ribs and cords. Teleoconch slender, of about 6 whorls. Axial ribs variable in number, 15 to 20 on last whorl of most specimens, extending adapically over slightly channeled sutural ramp to suture. Some ribs forming varices at irregular intervals, others projecting slightly above ramp. Spiral sculpture of fine cords, about 15 on last whorl of most specimens, with weaker second- ary cords in most interspaces. Cords form small nodules where they intersect axial ribs. Aperture narrowly ovate. Outer lip thickened into a varix, with about 14-16 strong lirae that do not descend deeply into aperture. Inter el liration beneath sutural ramp forms edge of well-defined posterior canal. Parietal shield thin but well developed, with pustules on its outer edge. Columella with three folds. anterior one forming edge of short siphonal canal. Last whorl slightly constricte sd at base behind weak si- phonal fesciole. Exterior brown or white. Many brown specimens with a weak band of white just below periph- ery and white on the adapical ends of ribs and projections extending above the shoulder. Material Examined: Japan. Off Mikawa, 30 fms, 1 spm (figured by Abbott and Dance, 1982)—Mikawa Is- siki, Aichi Pref., 50-70 m, 1 spm (Figure 20).—Off Min- abe, Wakayama Pref., 80-100 m, 2 spms (Figure 22)— Off Minabe, 100-200 m, 1 spm.—Off Mikawa, 50 fms, 1 spm (Figure 19).—Off Tosa, 50 fins, 1 spm.— ‘Japan’, | spm.—Japan?”, 1 spm (gift from Habe with “azumai” in his hand).—Mikawa Bay, 50 m, 1 spm.—Off Mikawa, 40 fms, | spm.—Enshu Kei, 2 spms. East China Sea. “180 m”, | spm. Taiwan. Off Keelung, 50-110 m, 2 spms.— Off Keelung, 100— sae m, 5 spms. —Off SW Taiwan, | spm. —Off Kee slung, “deep water”, | spm. —Otf SW Tai- wan, 60 fms, | spm. Etahppine s. Off Aliguay Island, Mindanao, “240 m”, 2 spms.—Off Aliguay Island, “80-120 m”, 1 spm.—Off Balicasag Island, ‘ ‘240 m”, 1 spm.—Off Balicasag, “130-230 m”, 1 spm (Figure 18) — MUSORSTOM 3: sta. DR140, 11°43’ N, 122°34' E, 93— 99 m, 1 dd: Sta. CP 141, 11°45’ N, 122°45' E, 40-44 m, 1 dd. Vietnam. no locality, “50 m”, 1 spm. Indonesia. Masalembo, Java, ca. 20 fms, 1 spm. (AIl above in Petit collection). Solomons. SOLOMON 1: sta. DW1760, 8°47’ S, 160°O1' E, 172-179 m, 1 dd. Coral Sea. CHAL- CAL sta. D11, 20°31' S, 161°06’ E, 83 m, 1 dd. New Caledonia. LAGON: sta. 375, 22°32’ S, 167°08' E, 67-71 m, 1 dd; Sta. 517, 19°09’ S, 163°35' E, 42 m, 2 dd: Sta. 542, 10°06’ S, 163°10' E, 50 m, 3 lv (Figures 24-25); Sta. 1129, 19°29’ S, 163°49' E, 40m, 3 lv, 2 dd: Sta. 1163, 19°11’ S, 163°22’ E, 48m, 2 dd; Sta. 1168, 19°16" S 163°09' E, 50 m, 1 lv; Sta. 1192, 19°35’ S, 163°25' E, 48m, 1 lv, 1 dd (Figure 23)—MUSORSTOM 4: sta. DW151, 19°07’ S, 163°22" E, 200 m, 1 dd. Fiji. SUVA2: sta. DW44, Viti Levu, 17°5 1.7 S, 177°13" BE, 33 m, 1 dd. Dimensions: 19.3 « 8.4 mm (Coral Sea, CHALCAL sta. D11), 17.9 x 8.3 mm (New Caledonia, LAGON sta. 1192), 14.8 « 7.3 (New Caledonia, LAGON sta. 1192), 30.2 x 14.0 mm (Japan). Distribution: Natal, Mozambique, Arabian Sea, An- grias Bank, India, Ceylon, Andaman Islands, northwest Australia, Japan (from Izu Peninsula and Yamaguchi Pref. southwards), Taiwan, the Philippines, Vietnam, In- donesia, Solomon Islands, Queensland, Australia, New Caledonia and Fiji. Offshore from 20 to ca. 250 m. Remarks: Of the eleven nominal species that we in- clude under Fusiaphera macrospira, two have type lo- calities in the southwest Indian Ocean, one in the Ara- bian Sea, two in the Bay of Bengal, two off northwestern Australia, one off Borneo and fice off Japan. We have examined representative mi aterial from Japan, Taiwan, the Philippines and New Caledonia and fail to recognize more than one species. All specimens have in common a naticoid multispiral protoconch indicating planktotrophic development and we interpret the different names as individual rather than geographical variants, connected by intermediates. Of these, the nominal species FP’. mac- rospiratoides represents a form where the axial and spiral sculpture are of equal prominence, whereas F. tosaensis represents a form with much stronger axial sculpture. Hasegawa (2000: 585) was of the same opinion when he Page LO THE NAUTILUS, Vol. 122, No. 1 stated that “[F. macrospiratoides| and other related spe- cies, such as F. azuwmai Habe, 1961 and F. tosaensis Habe, 1961 may be intraspecific forms of F. macrospira (Adams and Reeve, 1850).” We did not examine ex- tensive material from the Indian Ocean but published descriptions and illustrations suggest that the nominal species from this area also fall within the range of varia- tion of F. macrospira. Specimens from northwestern Australia described by Garrard (1975: 17-19) are, how- ever, distinctly smaller with average heights of 10.5 mm to 14 mm. Genus Nipponaphera Habe, 1961 Type species: Nipponaphera habei Petit, 1972 by ICZN Opinion 1052; Recent, Japan. Discussion: Species of Nipponaphera have the angled outline of Trigonostoma but are less tabulate, have only a small umbilicus if one is present, and have a different columellar morphology. The genus has been utilized pri- marily to include species having an angled last whorl, a triangular aperture, and three columellar folds. Here we also place in Nipponaphera species with a rounded last whorl and rounded aperture, but which are united with those taxa traditionally included in the genus by the pe- culiar columellar morphology consisting of two anterior columellar folds situated on a slightly iaiked shelf, much like a widely bifurcate single fold. The genus Misteia Janssen, 1984, from the Miocene of The Netherlands, is similar in outline to the angled spe- cies of Nipponaphera, but has only two weak folds on the columella. Axelella Petit, 1988 (a replacement name for the pre- occupied Olssonella Petit, 1970) has been considered to be confined to the Americas (Petit, L970: $4; 1972: 104). What appears to be a neat division of genera between the Americas (Axelella) and the Indo-Pacific (Nipponaphera) is blurred by the enigmatic species Cancellaria agalma Melvill and Standen, 1901, from the Gulf of Oman, a species that appears to possess the characters of Axelella. Despite that species, which has not been studied in de- tail, it is our opinion that the similarities between Avelella and Nipponaphera are superficial and that they can be se ost ated by their columellar morphology. ecent species of Nipponaphera have previously been known from South Africa to the northwestern Indian Ocean, eastward to the Philippines and north to Japan. The range is now extended to New Caledonia. In addi- tion to the species treated by Bouchet and Petit (2002), we now include in Nipponaphera the following: N. semi- pellucida (Adams and Reeve, 1850) [described in Can- cellaria; previously placed in Cancellaria by Habe (1961b) and other Japanese authors}; teramachii Habe, 1961) [described in Tgenapners, placed Sealptia and Trigonostoma by various authors; placed in Nipponaphe ra by Habe (1961la pl. 36, fig. 4) on the plate caption although in the text it is ee in Trigonaphera|; N. nodosivaricosa (Petuch, 197 bY [described in Agatrix Olssonella), place din Nipponaphera by Bouchet and Petit (2002)|: N. quasilla (Petit, 1987), new combination [described in Cancellaria|, N. kastoroae (Verhecken, 1997) new combination, and N. suduirauti (Verhecken, 1999) new combination [the last two described in Axe- lella). The fossil record has not been completely searched for Nipponaphera but we have recognized Cancellaria chinenensis MacNeil, 1961 of the Olenaw a Pliocene and C. yonabaruensis MacNeil, 1961 of the Okinawa Mio- cene as belonging here. Also, Oyama, Hirose and Nish- imoto (1995) described the new species Nipponaphera taguchii from the Miocene of Japan and at the same time transferred Cancellaria sendoi Hatai, 1941, to the genus Nipponaphera. Nipponaphera nodosivaricosa (Petuch, 1979) (Figures 26-29) Agatrix (Olssonella) nodosivaricosa Petuch, 1979: 11, figs. 26, 27. Description: —Protoconch pale brown, of about one to 1.2 whorls with fine spiral peek on final third. Tran- sition to teleoconch marked by prominent axial rib fol- lowed by onset of wide, rounded spiral cords and weak, poorly defined axial ribs. Teleoconch of 3.5 to 4.2 rounded whorls. Spiral sculpture of 12 to 15 rounded primary spiral cords, with weaker secondary cords in each interspace; about six primary cords on penultimate whorl. Primary and secondary cords all bear extremely fine spiral threads. Axial sculpture of eight to 12 promi- nent, elevated ribs on last whorl, more numerous on earlier whorls: final one or two ribs becoming wider, forming varices. Spiral cords and interspaces erossed by fine, closely spaced growth lines, giving surface a linen- like appearance under low magnification. Suture im- pressed, sutural ramp convex. Aperture elongate, rounded. Last whorl slightly constricted behind siphonal fasciole. Outer lip only slightly prosocline, edge thin. In- terior of outer lip w ith ten to 14 strong lirations extend- ing deeply into aperture. Stromboid notch manifested by very slight indentation in outer lip. No parietal callus: some specimens with a thin wash on parietal area. In- ductural area covered with thin callus, which extends back over chink-like umbilicus. Columella with three folds, posterior one most prominent, almost perpendicu- lar to axis; two anterior folds sharply descending, situated on ends of u-shaped platform, anterior-most one forming edge of short but well-defined siphonal canal. Exterior cream to pale yellow-brown, many specimens with ir- regular markings; most specimens with two or three elite spiral cords at periphery of last whorl, with two or three dark brown cords above and below. Type Material: Holotype (11 x 9 mm, fide Petuch; 12.9 x §.2 nm, fide Genie mn), DMNEH 126397. Type Locality: Off Balicasag Island, Philippines, from 300 m depth. Material Examined: New Caledonia. BATHUS 1: sta. DW672, 20°48" S, 165°21' E, 347-366 m, 1 lv (Fig- Figures 26-38. Nipponaphera. 26-29. Nipponaphera nodosivaricosa (Petuch, 1979). 26. Height 16.9 mm, New Caledonia 20°17’ S. 163°50' E, 500-600 m [BATHUS 4 sta. DWS9S8]. 27. Height 17.0 mm, New Caledonia, 20°48’ S, 165°21' E, 347-366 m BATHUS 1 sta. DW672]. 28. Protoconch, 21°45’ S, 166°37' E, 250 m [BATHUS | sta. CP713]. 29. Teleoconch microsculpture same specimen as 28, same scale. 30. Nipponaphera argo new species holotype, height $.6 mm; Coral Sea, 22°48’ S$, 159°24' E, 450 m | MUSORSTOM. 5 sta. 300). 31-36. Nipponaphera agastor new species 31—32 holotype height 19.3 mm, Solomon Islands, 9°21’ S, 160°24' E, 357-359m [SOLOMON 1 sta. CP1S00]. 33. Height 12.4 mm, Fiji, 19°52’ $, 174°40' W, 383-393 m [BORDAU | sta CP156] 34. protoconch Vanuatu, 20°20’ S, 169°49' E, 400-440 m |MUSORSTOM 8 sta. CP963]. 35. Teleoconch microsculpture same specimen as 34 same scale. 36. Height 17.9 mm Philippines L1°O1’ N, 124°04' E, 214-246 m |MUSORSTOM 3 sta. CP145 37-38. Nipponaphera tuba new species, holotype, he ight 20.7 mm, Vanuatu, 15°10’ S, 167°14" E, 394-421 m [MUSORSTOM §& sta. CP1LOST Page 12 THE NAUTILUS, Vol. 122, No. 1 ure 27), 1 dd.—Sta. CP713, a S, 166°37' E, 250 m, llv oo 98-29). BATHUS 2: sta. DW717, 22°44' S, err E, 350-393 m, 1 lv. BATHUS 4: sta. CPS97, 20°16’ S. 163°52’ E, 305-350 m, 1 Iv.—Sta. DWS898, 20°17' S, 163°50' E, 500-600 m, 1 dd (Figure. 26).— Sta. DW901, 19°03°S, 163°15' E, 297 m, 1 dd.—Sta. CP905, 19°02’ S, 163°16' E, 294-296 m, 1 lv.— Solomon Islands. SOLOMON 1: sta. CP1SO1, 9°25’ S, 160°26’ E, 264-273m, | lv. (Dimensions of largest New Caledonia specimen: 18.5 x 11.6 mm.) Distribution: At this time Nipponaphera nodosivari- cosa is known only from New Caledonia, the Solomons and the Philippines (Springsteen and Leobrera, 1986; V erhecken, 1999). The Indonesian specimen figured as Axelella cf. nodosivaricosa by Verhecken (1997: 299, figs. 5-7) was not attributed by him in his 1999 work to either N. nodosivaricosa or N. suduirauti (see below). In the New Caledonian dredgings, live specimens were taken from depths of 250-393 m and empty shells from as deep as 600 m. Remarks: Verhecken (1999) described the species Axelella suduirauti, here placed in Nipponaphera, distin- guished from N. nodosivaricosa based on protoconch characteristics. It was stated that N. suduirauti has a multispiral protoconch as opposed to the paucispiral pro- toconch of N. nodosivaricosa, Our specimen of N. no- dosivaricosa from the Solomon Islands has a protoconch that is difficult to attribute to one or the other of the two species and we believe that the separation between N. nodosivaricosa and N. suduirauti should be reevaluated, perhaps using molecular characters. We refer to Ver- hecken’s (1999) work for his discussion on the question. Nipponaphera argo new species (Figure 30) Description: —Protoconch smooth, of 1.1 whorls, diam- eter 1050 wm. Teleoconch of about 3.1 whorls, high- spired. Whorl profile angulated at shoulder. Sculpture of low, broad axial ribs and much finer spiral cords of rather even stre neth, exce pt for one on sutural ri amp and two at periphery of last whorl, which are more prominent than others: 14 axial ribs on penultimate whorl, seven on last whorl (specimen with severe growth scar and regrowth, distorting sculpture of last whorl); about 15 spiral cords on penultimate whorl and about 35 on last whorl, crossed by thin incremental riblets. Suture shallowly impressed. Last whorl slightly constricted behind siphonal fasciole. Outer lip thin, sharp, smooth within, lacking lirae. Inner lip with well-deve lope od parieti il shield, exte nding slightly over narrow umbilicus. Columella only slightly concave, with three folds: anterior two close toge the 1 much like one large bifurcate fold. Siphonal canal short, indistinct. Exterior uniformly very pale yellowish-white. Type material: Holotype MNHN 20511 (8.6 « 6.5 mim) and one paratype MNHN 20512 Type Locality: Argo Bank, Coral Sea, 22°48’ S, 159°24' FE, 450 m [MUSORSTOM 5, sta. 300]. Material Examined: Coral Sea. MUSORSTOM 5, sta. 299, 22°48’ S, 159°24’ EF, eae: m, | dd (paratype ).—Sta. 300, 22°48’ S, 159°24’ E, 450 m, TH (holotype, Figure 30). Etymology: Named for the Argo Seamount, a promi- nent topographic feature of the Coral Sea, from which the specimens were collected; to be treated as a noun in apposition. Distribution: 390-450 m. Coral Sea (Argo Seamount), dead in Remarks: Nipponaphera argo differs from N. goniata Bouchet and Petit, 2002 by its sculpture of oad low, non-lamellar axial ribs. Also, the spiral cords of N. argo are of more even strength, except for one on the choulder and two on the periphe ry, which a a slightly bian- gular aspect to shells of this species Nipponaphera agastor new species (Figures 31-36) Description: —Protoconch normally smooth, corroded on holotype, of 0.9 whorls, diameter SOO fxm. Proto- conch/teleoconch boundary indistinct due to corrosion, but distinctly marked by onset of teleoconch sculpture on specimens from Vanuatu. Teleoconch of five rounded whorls; spire angle 64°; suture deeply impressed. Axial sculpture of prominent, regularly spaced ribs, eleven on last whorl, eleven on pe anlbice ite whorl. Ribs rounded over steep, narrow sutural ramp. Spiral sculpture of evenly spaced, prominent spiral cords, eight on penulti- mate whorl, 13 on last whorl, with three to five secondary cords in each interspace; spiral cords rise over axial ribs, forming small nodules on primary cords at intersections. Numerous fine growth lines cross spiral cords, creating small imbrications. Last whorl slightly constricted behind siphonal fasciole. Outer lip prosocline. Inner margin of lip smooth apart from 16 prominent lirae « xtending into aperture; two indistinct lirae on parietal area. Columellar callus well developed, forming shield over chink-like um- bilicus. Columella with three almost equal folds; anterior one sloping sharply down at edge of small but distinct siphonal canal, which recurves abaxially. Exterior chalky white. Type Material: Holotype 20513 (19.3 x 12.3 mm) and 4 paratypes MNHN 20514. Type Locality: Between ie idaleanal and Florida Is- land, Solomon Islands, 9°21’ S, 160°24' E, 357-359 m [SOLOMON I: sta. Cea. Material nara: Vanuatu. MUSORSTOM 5: sta. CP963, 20°20' S, 169°49' E, 400-440 m, 1 lv (Figures 34-35) pen ‘BORD: AU 2: sta. CP 1561, 19°52’ S, 174°40' W, 383-393 m, 1 dd (Figure 33).—Solomons. SOLOMON 1: sta. CP1L746, 09°23' S, 159°57' Ei, 302- 396 m, 1 dd; Sta. CP1SO0, 9°21' S$, 160°24' BE, 357— P. Bouchet and R. E. Petit, 2008 Page 13 359m, 4 lv, 1 dd (holotype, Figures 31-32, and paratypes).—Philippines. MUSORSTOM 3: sta. CP145, L1°OL' N, 124°04" E, 214-246 m, 1 dd (Figure 36).— Punta Engano, tangle nets, approximately 60 fms, 1 spm. Etymology: From the Greek agastor, a noun in appo- sition, meaning near kinsman or brother, to highlight the similarity to N. nodosivaricosa. Distribution: Only known from the material exam- ined: Philippines, Solomons, Vanuatu and Tonga. Depth range in the southwest Pacific 360-400 m; in the Philip- pines possibly shallower. Remarks: Specimens from the Philippines s and the So- lomons are distinctly larger with adult sizes ranging from 16.8 mm to 19.5 mm, whereas specimens from Vanuatu and Tonga are much smaller with adult sizes at 12.2 min and 12.38 mm respectively. Nipponaphera agastor is sympatric with N. nodosivari- cosa in the Philippines s and the Solomon Islands but dif- fers by its spiral sculpture with more numerous (3 to 5) secondary cords, not separated by an incised groove as in N. nodosivaricosa. It differs from N. tuba, w ath which it is sympatric in Vanuatu, by being more slender and lack- ing a stromboid notch. Nipponaphera tuba new species (Figures 37-35) Description: —Protoconch smooth, of 0.9 whorls, diam- eter SOO zm. Protoconch/teleoconch boundary distinctly marked by onset of teleoconch oo ae Teleoconch of five rounded whorls: spire angle 75°; suture deeply im- pressed. Axial sculpture of prominent, regularly spaced ribs, ten on last whorl, 14 on penultimate whorl: ribs rounded on steep, narrow sutural ramp. Spiral sculpture of evenly spaced cords of several strengths; primary cords evenly See by one secondary cord, resulting inter- spaces filled with two or three tertiary cords: spiral cords rising over axial ribs, forming small nodules on primary spiral cords at intersections. Numerous fine growth lines cross spiral cords, creating small imbrications. Last whorl slightly constricted behind siphonal fasciole. Outer lip prosocline. with distinct stromboid notch adapical to cen- ter of lip. Inner margin of lip smooth or slightly crenu- late, with 19 prominent lirae extending into aperture, four additional lirae beneath sutural ramp. Columellar callus well developed. forming shield over chink-like um- bilicus. Columella with three almost ae folds, anterior one sloping sharply down at edge of small but distinct siphonal canal, which recurves abaxially. Exterior yellow- brown with a band of white below periphe ry, bordered by indistinct bands of darker brown. Type Material: Holotype MNHN 20516 (20.7 « 15.0 mm) and one paratype MNHN 20517 Type Locality: Vanuatu, 15°10’ S, 167°14’ E, 394— 421 m [MU SORSTOM 8: sta. CP1087]. Material Examined: Vanuatu. MUSORSTOM S: sta. CP1087, 15°10’ S, 167°14’ E, 394-421 m, 1 lv, 1 dd (holotype, Figures 37—38, and paratype).— Sta. [no data, mixed lot|, | a, Etymology: From the Latin tuba, a war trumpet, which this new species can, with some imagination, be reminiscent of; used as a noun in apposition. Distribution: Known only from Vanuatu. Remarks: Nipponaphera tuba differs from N. cy- phoma Bouchet and Petit, 2002 in being more rounded and robust. Also, the spiral sculpture never appears as incised lines as in N. cyphoma and N. nodosivaricosa. Nipponaphera tuba also has a thick outer lip that is not present in N. cyphoma. Genus Trigonostoma Blainville, 1827 Trigona Perry, 1811: pl. 51. Type species: Trigona pellucida Perry, ISL], by monotypy. Not Trigona Jurine, 1807 (Hy- menoptera). Trigonostoma Blainville, 1827: 652. Type species: Delphinula trigonostoma Lamarck, 1822 (?= Buccinum scalare Gme- lin, 1791), by monotypy. Recent, Indo-Pacific Remarks: = Trigonostoma has a different taxonomic composition for various authors. We here use Trigono- stoma sensu lato to encompass the nominal genera Ven- trilia Jousseaume, 1887, Arizelostoma Iredale, 1936, Ovilia Jousseame, L887, and Extractrix Korobkov, 1955. We do not include Scalptia Jousseaume, 1887, Trigona- phera Iredale, 1936 and Cancellaphera Iredale, 1930, which are sometimes treated as subgenera of Trigono- stoma, Trigonostoma tryblium new species (Figures 39-44) Description: —Protoconch smooth, glassy, of one whorl, diameter 1050 jzm. Transition from protoconch to teleo- conch abrupt, denoted by onset of axial and spiral sculp- ture. Teleoconch of 2.5 whorls: spire depressed; umbili- cus broad. First teleoconch whorl with 16 axial ribs. Sec- ond whorl with about 18 axial ribs that have become somewhat obsolete. Spiral sculpture of broad, closely spaced cords with narrow interspaces; about seven to ten spiral cords on sutural ramp and 25 between shoulder angle and umbilical rim. Sutural ramp weakly concave, bat forming deeply channeled shoulder between shoul- der angle and impressed suture. Aperture narrowly ovate. Outer lip smooth. Inner lip forming parietal shield, partly covering deep, wide umbilicus. Columella with two descending folds. Anterior canal not con- stricted. Holotype exterior chalky white, with two broad, ill-defined brown bands, best seen through shell. Type Material: Holotype MNHN 20518 (7.0 « 6.5 mm). Type Locality: North of Makira Island, Solomon Is- lands, 10°13’ S, 161°29' E, 381-383 m [SOLOMON 1: sta. CP1837]. Material Examined: Taiwan. TAIWAN 2000, sta. DW36, 21°54.8 N, 120°36.2 E, 305 m, Bashi Channel, | Page 14 THE NAUTILUS, Vol. 122, No. 1 Figures 39-47. Trigonostoma. 39-44. Trigonostoma tryblium new species. 39. Height $.3 mm, Taiwan, Bashi Channel, 21°54.8' N, 120°36.2' EF, 305 m [TAIWAN 2000 sta. DW36 40-41. Holotype, height 7.0 mm, Solomon Islands, 10°13’ S, 161°29' E, 381-383 m [SOLOMON 1 sta. CP1837]|. 42. Height 5.0 mm; New Caledonia, 23°03’ S, 166°5S' E, 397-400 m [BATHUS 2, sta. DW730] 43-44. Protoconch and teleoconch microsculpture, same specimen as 42. 45-47. Trigonostoma thysthlon Petit and Harasewych, 1987 SMIB 1 sta. DW6 dd (Figure 39). Solomons. SOLOMON 1: sta. CP1837 10°13" S, 161°29" E, 381-383 m, 1 lv (holotype, Figures {0-41). New Caledonia. BIOCAL Sta. DW77, 22°15’ S 167°15' FE, 440 m, 1 juv. lyv—BATHUS 2, sta. DW730 'S, 166°58’" FE, 397-400 m, 1 juv. dd (Figures 42-44 15. Height 16.9 mm, New Caledonia, 22°47’ S, 167°28' E {'S 241-245 m [BATHUS 2 sta. CP728]. 46. Height 15.0 mm, Fiji, 178°12.5' E, 200-215 m |MUSORSTOM 10: sta. DW1333]. 47. Protoconch, New Caledonia, 22°43’ S, 167°16' E, 300 Etymology: From the Latin tryblium, meaning cup, a shape that the new species is somewhat reminiscent of; used as a noun in apposition Distribution: Known only from Taiwan, New Cale- P. Bouchet and R. E. Petit, 2008 Page 15 donia and the Solomons. Alive in 353-440 m; empty shells from 305 m. Remarks: Of the known Recent species of Trigono- stoma, Trigonostoma tryblium resembles only T. semid- isjuncta (Sowerby, 1849) in having primarily spiral sculp- ture. It differs from that specie s in bei ing more de- pressed, with a broader umbilicus. In T. se midusjunc ta there is no spiral sculpture on the sutural ramp. Also, the spiral sculpture of T. semidisjuncta is arranged in groups of cords separate od by wide furrows. Trigonostoma thysthlon Petit and Harasewych, 1987 (Figures 45-47) Trigonostoma thysthlon Petit and Harasewych, 1987: 8-13. Trigonostoma antiquata—Habe, 1961a: 435, pl. 24, fig. 14; pl. 23, fig. S; 1961b: 73, pl. 36, fig. 8; Lan, 1979: 95, pl. 41, figs. 93, 93a; Abbott and Dance, 1982: 299 (second figure 922 in bottom row); Habe and Okutani, 1985: 233 (second figure in pottow row); Bosch, et al., 1995: 157, fig. 687. [not Cancellaria antiquata Hinds, 1843] Trigonostoma aie: pager ie and Takemura, 1963: Trigonaphera (2) plate, fig. 5. [not Cancellaria antiquata Hinds, 1843} Trigonostama (sic) thysthlon—Hasegawa, 2000: 581, pl. 290, fig. 11. 79, figs. 5, ~l Type Material: mm). Holotype, USNM 747301 (17.3 x 12. Type Locality: Off west coast of Wasir Island, West Wokam, Aru, Moluccas (5°30' S$, 134°12' E) in 56-73 m. Material Examined: New Caledonia. LAGON: sta. 387, 22°39’ S, 167°07’ E, 225 m, 1 dd.—SMIB 1: sta. DW6, 22°43’ S, 167°16' E, 300 m, 1 dd (Figure 47).— BATHUS 2: sta. CP728, 22°47’ S, 167°28' E, 241-245 m, | lv Bele, north New Caledonia, 1 spm. Fiji. MUSOR- STOM 10:sta. DW1333, 16°50.4' S, 178°12.5' E, 200— 215 m, 1 dd (Figure 46). Solomons. SOLOMON 1: sta. DW 1850, 10°28’ S, 161°59' E, 139-261 m, 1 lv. Description (of a specimen from New Caledo- nia): Protoconch smooth, glassy, of 1.8 whorls, diam- eter 1050 tm. Transition to tele oconch marked by axial rib followed by both axial and spiral sculpture and flat- tening of the sutural ramp. Teleoconch of up to six tabu- late whorls. Sutural ramp flat, bordered by cord-like shoulder angle. Suture impressed. Axial ribs prominent, extending from suture across ramp, over shoulder angle, where they form recurved spines on many specimens, down and over siphonal fasciole, then inside umbilicus. Shoulder spines not formed on all ribs, but many ribs lacking spines form short. sloping buttress against pre- ceding whorl: about 12-15 ribs on last whorl, more nu- merous on earlier whorls: two thick, closely spaced ribs mark end of growth in adults: a few varix-like rib occurs earlier on some specimens. Fine, closely packed growth lines of varying number in axial interspaces slightly over- lap each other, producing a scabrous appearance. Spiral (Figure 45)—Dredged at “300-400 m”, off sculpture of very closely spaced cords, rising over ribs to form small nodes. Sutural r: amp with about 12-15 spiral cords of uniform prominence; about 15 primary spiral cords on last whorl between shoulder angle and siphonal fasciole; about ten fine secondary cords in each inter- space. Spiral cords of equal strength continue inside um- bilicus. Outer lip slightly prosocline. Outer lip thickened by varix, slightly serrate on margin, interior with eight to ten short irregular lirae, not extending quite to outer edge of lip and not extending deeply into aperture. Ap- erture triangular, with sme ull posterior notch under su- tural 1 ramp. Posterior portion of inner lip adpressed against siphonal fasciole and anterior quarter of penulti- mate whorl. Inner lip with two descending folds, poste- rior one slightly larger than anterior; third incipient fold present in some specimens. Umbilicus extends to proto- conch. Siphonal canal short. Exterior white with very faint orange-brown at shoulder. Specimens over 24 mm in height are known. Distribution: Gulf of Oman to the Moluccas, Japan, the Philippines, Solomon Islands, New Caledonia and Fiji. Remarks: The imbricate sculpture resulting from the overlap of growth lines, which is so holicenble on the New Cale doa specimens, is absent from T. antiqua- tum and is much less evident on T. thysthlon from other areas. However, we do not consider this difference to be of taxonomic significance. This sculpture isa promine nt feature of T, sculore (Gmelin, 1791), the type of the genus but that species has a more angular shape and its whorls are barely attached. ACKNOWLEDGMENTS We thank Bertrand Richer de Forges, the indefatigable as ee of the first author on many expeditions, for his skill and determination in exploring the deep-water benthos of the tropical Pacific. Ahmed Abdou did SEM illustrations of protoconchs and microsculpture; Del- phine Brabant and Philippe Maestrati did the digital photography and plate assembling. Bruce Marshall helped us interpret our material of Admetula from Fiji. Alan Beu helped us improve the manuscript by tracking internal inconsistencies. LITERATURE CITED Abbott, R. T. and S. P. Dance. 1982. Compendium of Seashells E. P. Dutton, Inc., New York, ix + 411 pp. [pre printing revised, 1983; 3"! printing, revised, 1956; see also Habe and Okutani, 1955}. Adams, A. and L. Reeve. 1848-50. Mollusca. In: A. Adams (ed.) The zoology c the voyage of H.M.S. Samar ngs under the command of Captain Sir Edward Belcher, ,F.RAS., F.G.S. during the years 1543-1546. Reeve, ae and Reeve, London. x + xv + 87 pp., pls. 1-24. 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William Miller, London. 4 pp. + 61 pls. with pl. explana- tions. Petit, R. E. 1967. Notes on Cancellariidae (Mollusca: Gas- tropoda). Tulane Studies in Geology 5; 217-219. Petit, R. E. 1970. Notes on Cancellariidae (Mollusca: Gas- tropoda) - II. Tulane Studies in Geology and Paleontology 8: §3-88, pl. 1 Petit, R. E. 1972. The cancellariid genus Nipponaphera Habe, 1961. Venus 31: 103-105. Petit, R. E. 1974. Notes on Japanese Cancellariidae. Venus 33: 109-115. Petit, R. E. 1976. Notes on Cancellariidae (Mollusca: Gas- tropoda) — HI. Tulane Studies in Geology and Paleontol- ogy 12: 33-43, pls. 1-2 Petit. R. E. 1980. The Mozambique Cancellariidae (Mollusca: Gastropoda). Annals of the Natal Museum 24: 211-216. Petit. R. E. 1987. New names for two species of Cancellaria (Mollusca: Gastropoda). The Nautilus 101: 154 . Eocene Mollusca. Bulletins of Petit, R. E. 1988. Axelella, new name for Olssonella Petit, 1970, a preoccupied taxon (Mollusca: Cancellariacea). The Nau- tilus 102: 130. Petit, R. E. and M. G. Harasewych. 1986. New Philippine Cancellariidae (Gastropoda: Cancellariacea), with notes on the fine structure and function of the nematoglossan radula. The Veliger 28: 436-443. Petit, R. E. and M. G. Harasewych. 1987. The Indo-West Pa- cific species of the genus Trigonostoma sensu stricto (Gas- tropoda: Cancellariidae). The Veliger 30: 75-81. Petit, R. E. and M. G. Harasewych. 1991. A new Admetula (Gastropoda: Cancellariidae) from South Africa. Proceed- ings of the Biological Society of Washington 104; 1S1— 183. Petit, R. E. and M. G. Harasewych. 2000. Additions to the cancellariid (Mollusca: Neogastropoda) { fauna of South Af- rica. Proceedings of the Biological Society of Washington 113: 145-154. : Petuch, E. J. 1979. Twelve new Indo-Pacific gastropods. Nem- ouria 23: 1-21. Powell, A. W. B. 1940. The marine Mollusca of the Aupourian Province, New Zealand. Transactions of the Royal Society of New Zealand 70(3): 205-248, pls. 28-33. Powell, A. W. B., 1979. New Zealand Mollusca. William Collins Publishers, Ltd, Auckland. i-xiv + 1-500 pp., 82 pls. Preston, H. B. 1905. Descriptions of new species of marine shells som Ceylon. The Journal of Malacology 12(1): 1-8, pls. 1-2 Sacco, F. 1894. I molluschi dei terreni terziarii del Piemonte e della Liguria. Pt. 16 (Cancellariidae). C. Clausen, Torino. 78 pp., 3 pls. [Reprinted 1987-89 by Atlante Malacologico, Roma. | Smith, E. A. 1899. Notes on some marine shells from North- West Australia, with descriptions of new species. Pro- ceedings of the Malacological Society of London 3(6): 311-314. Solander, D. C. 1766. [descriptions of species]. In: Brander, G., Fossilia Hantoniensia collecta, et in Musaeo Britannico deposita, a Gustavo Brander. Londini, vi + 43 pp. 9 pls. Sowerby, G. B., I. 1849. Descriptions of some new species of Cancellaria in the collection of Mr. H. Cuming. Pro- ceedings of the Zoological Society of London 1545: 136— 138. Sowerby, G. B., If. 1881. Description of eight new species of shells. Proceedings of the Zoological Society of London 1881: 635-638, pl. 56. Sowerby, G. B., HI, 1903. Mollusca of South Africa. Marine Inve stigations in South Africa 2: 213-232, pls. 3-5, Springsteen, F. J. and F. M. Leobrera. 1986. Shells of the Philippines. Cartel Seashell Museum, Manila. 377 pp. Tate, R. 1889. The gastropods of the older Tertiary of Australia. Part I. Transactions and Proceedings and Report of the Royal Society of South Australia 11: 116-174, pls. 2-10. Verhecken, A. 1986. The Recent Cancellariidae of Indonesia (Neogastropoda, Cancellariacea), Gloria Maris 25(2): 29- 66. Verhecken, A. 1997. Mollusca Gastropoda: Arafura Sea Can- Page 18 THE NAUTILUS, Vol. 122, No. | cellariidae collected during the KARUBAR Cruise. In: A. Crosnier and P. Bouchet (eds), Résultats des Campagnes MUSORSTOM, Volume 16. Mémoires du Muséum Na- tional d'Histoire Naturelle 172: 295-323. Verhecken, A. 2007. Revision of the Cancellariidae (Molhusca, Neogastropoda, Cancellarioidea) of the eastern Atlantic (40°N—40°S) and the Mediterranean. Zoosystema 29: 281— 364. Verhecken, A. 1999. A new species of the genus Axelella (Mol- lusca, Neogastropoda, Cancellariidae) from off Japan and the Philippines; with notes on related species. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Bi- ologie 69: 23-30. Wilson, B. 1994, Australian marine shells. Prosobranch gas- tropods, Part 2 (Neogastropods). Odyssey, Kallaroo, 370 pp. THE NAUTILUS 122(1):19-51, 2008 Page 19 On some Neogene to Recent species related to Galeodina Monterosato, 1884, Galeodinopsis Sacco, 1895, and Massotia Bucquoy, Dautzenberg, and Dollfus, 1854 (Caenogastropoda: > Rissoidae) with the description of two new Alvania species from the Mediterranean Pleistocene Vittorio Garilli APEMA Research and Educational Service Via Alla Falconara, 34 1-90136 Palermo, ITALY vittoriogarilli@apema.eu. ABSTRACT Six species, related to the subgenera Galeodina, Galeodinopsis, and Massotia, are re-analyzed. Alvania francescoi new species (SE Sicily) and A. rosariae new species (SE Sicily and NW Peloponnesus) are described from Mediterranean Pleistocene. Galeodinopsis is regarded as the appropriate genus for Oli- gocene-Recent taxa having a quite conical shell close to that of some Alvania species and showing Manzonia-like combination of two microsculptural characters: the pitted surface on the spiral cords and the arrangement of the roughly prismatic ele- ments forming numerous and very fine spiral nes ids. Its type species, Rissoa tiberiana (previously known from Mediterra- nean Mio-Pliocene), lives along the tropical W African coasts, where it is known under the name A. fariai. As suggested by the oldest record of Galeodinopsis, the European Oligocene Rissoa duboisi, this genus very likely originated from a pre-Neogene Alvania group. The syntype of Rissoa prusi, a scarcely known species (Pleistocene of Rhodes), and material (also from type locality) of the almost unknown R. cingulata (from Sicily) and of its close relative, R. tenera (Mediterranean, Atlantic Moroc- co and Canary Islands), is shown. These three taxa and the type species of Galeodina and Massotia, are here tentative ‘ly con- sidered as belonging to Alvania sensu lato. With the exception of A. cingulata, all the discussed species have a multispiral protoconch. Generally, the protoconchs studied exhibit a sculp- tural pattem known in other rissoid taxa. Protoconch I of the type species of Massotia, A. lactea, is characterized by a coarser sculpture Additional Keywords Atlantic, Neogene-Recent, new species Aissoidae, taxonomy, Mediterranean-W INTRODUCTION The genus Alvania Risso, 1826, comprises one of the most diversified groups in the caenogastropod family Rissoidae, especially when considering the tropical east- ern Atlantic and the Mediterranean provinces. They in- habit a large variety of environments, from littoral to bathyal, and their geographical distribution is extensive, including the Mediterranean, Atlantic, Indo-Pacific, and the temperate Australian coasts (Ponder, 1985). Their stratigraphical distribution might extend back to the Late Cretaceous, but the first well-established records date to the early Tertiary (Ponder, 1985). With regard to the rich Mediterranean and E suropean Tertiary Alvania assem- blages, some of the most informative analyse ‘sare those of Sacco (1895), Seguenza (1903), Cossmann (1921), Lo- zouet (1998), Kowalke and Harzhauser (2004), and Chirli (2006). Many authorities, e.g. Monterosato (1554), Bucquoy et al. (1884), Weink matt | 1885), Kobelt (1888), Locard (1886), Nordsieck (1968, 1972), Jeffreys (1867, 1569), Pallary (1920), Wenz (1938), Warén (1975, 1974), Gofas and Warén (1982), Van Aartsen (1982a, 1982b) . Moolen- beek and Hoenselaar (1989, 1998), Van der Linden and Wagner (1989), Moolenbeek et al. (1991), Van der Lin- den (1993), Bouchet and Warén (1993), Peg wee Savelli et al. (1996), Palazzi (1997), Gofas (1999), Avila (2000), and Arduino and Arduino (2001), nee contrib- uted to the knowledge of the rich Recent Eastern Atlan- tic (especially the W Africa and the Macaronesian Prov- ince) and Mediterranean fauna. Ponder’s rissoid revision (1985), listing five Alvania subgenera, provided addi- tional perspective to the taxonomy. My attention is focused here on a relatively large, in- formal group of species of Alvania sensu lato character- ized by shells with wide and ovate aperture, lacking an internal denticulation of the outer lip, and often bearing varices on a well-developed, inflated body whorl. These species have been historically assigned to the subgenera Galeodina Monterosato, 1884, Gale ‘odinopsis Sacco, 1895, and Massotia Bucquoy et al., 1554. This subgeneric settlement was partially rejected by Ponder (1985), who Page 20 THE NAUTILUS, Vol. 122, No. 1 modifying the sy’ stematic ar rangement of Monterosato (1884) and Bucquoy et al. (1884), stated that grouping of the numerous species of fee was very difficult at the subgenus level. The same author included Massotia ( (type species: Rissoa lactea Michaud, 1830) and Galeodinopsis (type species: R. tiberiana Coppi, 1876) in the Alvania sensu stricto group and mca considered Galeodina (type species Turbo carinatus Da Costa, 1778) as a valid subgenus on the basis of shell characters. Piani (1979) raised Galeodina to generic level and placed Rissoa cin- gulata Philippi, 1836, and R. tenera Philippi, 1844, therein. The principal aim of this work is to provide, for the first time, as complete as possible a taxonomic dataset based on shell features. Species such as Rissoa cingulata, loften misidentified as Alvania carinata (Da Costa, 1778)], R. prusi Fischer, 1877, and R. tiberiana Coppi, 1876, are very poorly known or, in the case of the last taxon, the generic attribution to Alvania appears incor- rect. I also describe two new species from the Pleisto- cene of the central Mediterranean area. MATERIALS AND METHODS Most of the examined material, all consisting of shells, is housed in the Muséum National d'Histoire Naturelle, Paris (Département Systématique et Evolution), in the private collections of Maurizio Forli (Prato, Italy) and of Stefano Palazzi (Modena, Italy). Remaining material is housed in the private collections of medshells.com (made available by Nino Adorni Sbrana, Grosseto, Italy), Ste- fano Rufini (Anguillara), in the Museum fiir Naturkunde (Humboldt University, Berlin), the Museo Geologico G.G. Gemmellaro and the Dipartimento di Geologia e Geodesia (both University of Palermo, Italy), in the Gou- landris Natural History Museum (Kifissia, Athens), the Dipartimento di Scienze della Terra of the University of Catania, Italy, the Zoologisch Museum of Amsterdam, the Muséum National d’Histoire Naturelle of Paris (Dé- partement Histoire de la Terre), and in my personal col- lection. Many other private collections were visited. In the parts dedicated to each species, a list of the material is provide ed with all information given in the original labels. An abbreviated list of essential synonymy and/or citations is provided. The fossil material (all from the Mediterranean area), from the deposits of Dattilo (NW Sicily), Cartiera Mulino (Vittoria, SE Sicily, type locality of Alvania francescoi new species and Alvania rosariae new spe- cies), and Kyllini (Elea, NW Peloponnesus, Greece), was obtained by washing bulk a on a serial sieves (0.5, 1, and 2 mm square meshes) and sorted using a stereo- microscope. The same procedure was followed for the Aecent bulk sam] sles collected from Magnisi (Siracusa, SE Sicily, the ee locality of Rissoa cineulata Philippi, 1836) and Mondello (Palermo, NW Sic ‘ily ) spec ifics ally to recover that species. Geological, stratigraphic and paleo- ecological information on the deposits of Cartiera Mulino, Dattilo and Kyllini are taken from Costa (1989), Garilli (1998; 2004), Garilli et al. (2005a; 2005b) and Garilli and Galletti (2007). The stratigraphic information on the Sicilian deposits of Birgi (Trapani) and Tommaso Natale (Palermo, originally attributed to the upper Pleistocene Thyrrenian by Ruggieri and Milone, 1973) are from Ruggieri and Unti (1988) and Hearty et al. (1986), respectively. For the remaining fossil ma- terial (mainly from Coll. Forli ex-coll. Palag7i), I followed the stratigraphic attributions reported by the col- lectors. For all the discussed species, at least five shells were studied by the scanning electron microscope (SEM) us- ing a Philips XL30 ESEM, except for the (sole) syntype of Rissoa prusi and R. cingulata. of which there were only three shells available. Specimens examined by SEM were cleaned in a Bransonic 5 ultrasonic machine using distilled water. Particular attention was given to proto- conchs and teleoconch microsculptures as potential sources of taxonomic characters at species level. The number of protoconch whorls were counted according to Verduin’s method (1977). Shells were measured using a stereo microscope pro- vided with a cross-line micrometer eyepiece. The posi- tion of any varix on the body whorl is indicated in degrees of the angles formed by the plane of the varix and of the outer lip. Geographic, bathymetric, and stratigraphic distribu- tion of each discussed species is based on the examined material and the literature. Published records were criti- cally evaluated on the basis of good illustrations or sat- isfactory de scriptions. This type of dataset allows for just a rough representation of the geographical distribution, e specially for the Adriatic Sea and the easternmost Medi- terranean basin, of which I found very little material from the collections studied. Abbreviations are used as following: DGUP: Diparti- mento di Geologia e Geodesia, Universita di Palermo, Italy; DSTC: Dipartimento di Scienze della Terra, Uni- versita di Catania, Italy; GNHM: Goulandris Natural History Museum, Kifissia, Athens, Greece; MGUP: Mu- seo Geologico G.G. Gemmellaro, Universita di Palermo, Italy; MNHN-DHT: Muséum National d'Histoire Na- turelle, Departement Histoire de la Terre, Paris, France; MNHN-DSE: Muséum National d'Histoire Naturelle, Departement Systématique et Evolution, Paris, France; MPOB: Dipartimento del Museo di Paleobiologia e del?Orto Botanico, Universita di Modena e Reggio Emilia, Modena, Italy; MSNCS: Museo Regionale di Storia Naturale e Mostra Permanente del Carretto Sicil- iano, Terrasini, Italy; ZMA: Zoologisch Museum Aimster- dam, Holland: ZMB: Museum fiir Naturkunde, Hum- boldt Universitit, Berlin, Germany; Coll.; collection; Coll. MF: Maurizio Forli collection, Prato, Italy; Coll. PAL: Stefano Palazzi collection, Modena, Italy; Sh(s) shell(s), used in the Material Examined sections only, V. Garilli, 2008 Page 21 SYSTEMATICS Family Rissoidae Gray, 1847 Subfamily Rissoinae Gray, 1S47 Genus Alvania Risso, 1826 Type Species: = Alvania europea Risso, 1826 [synonym of A. cimex (Linnaeus, 1758)], subsequent designation by Nevill, 18S5 Alvania carinata (Da Costa, 1778) (Figures 1-14) Turbo carinatus Da Costa, 1778: 102-103, pl. 8, fig. 10 Rissoa trochlea Michaud, 1830: 16, fig. 4 Galeodina cingulata (Philippi, 1S36).—Piani, 1979: 70-71, figs. 2-3 Description: Shell small, sturdy, conical and keeled to slender and turrited, reaching about 5 mm (rarely 6 mm) in height; 3.94.1 mm in width. Protoconch multispiral, conical, consisting of about 2.2 convex whorls. Proto- conch [ with 0.5 whorls, sculptured with six very fine spiral lirae and microscopic granules between them. Pro- toconch/teleoconch transition well-marked and sinuous. Protoconch H sculptured with sparse, spirally arranged microscopic Sade stronger in adapical direction, and one to two spiral ridge »s, one of them always very close to lower suture. In the irgest shells, te sleoconch formed by 44.5 moderately convex whorls. Common morph (Fig- ures 2, 6-7) with teleoconch whorls mar kedly dominated by spiral sculpture, which consists of very strong cords (numbering 2-3, 3-6, 6-12, and 12-17 on first, second, third, and last teleoconch whorl, respectively). Second- ary, less conspicuous cords may occurr on last whorl. More marked spiral cords on adapical portion of whorls, at a certain distance from suture, give a characteristic keeled shape. Cords progressively less strong on basal area. Unkeeled morph (Figures 1, 4-5) characterized by a ser er shell shape, usually bears more spiral cords (1S—20) on last whorl. Axial sculpture always formed by numerous (32-50 on penultimate whorl), occasionally very narrow and lamella-like ribs, becoming obsolete to- ward base. Intersection of ribs with spiral cords gives an almost general clathrate pattern in unkeeled morph. In- tersection of spiral and axial sculptures nodular, usually forming squares (Figure 14), with i of last whorl where a rectangular pattern occurs (Figure 13). Micro- sculpture consists of very fine spiral threads (Figures 13-14), covering all teleoconch surface, with exce ption of main spiral pattern. On early teleoconch whorls, spiral lirae often alternate with spiral alignments of micro- scopic pimples. Sutures slightly inclined. Last whorl well expande od, comprising 3/5 to 3/4 ( rarely more than 3/4, Figure 3) of shell height, often bearing one or two varices (mainly in keeled morph) with angles of 10°-340°. Ap- erture wide, ovate, slightly rounded to ; angled in the pos- terior part, comprising 3/5 to 3/4 of fase whorl height. Outer lip slightly prosocline, internally smooth, exter- nally markedly thickened by a strong rim very close to lip edge, and covered by spiral cords. Inner lip moderately arcuate and rather thickened, with a very narrow to dis- cretely expanded (Figure 3) callus delimiting a very small umbilical chink. Type Locality: Cornwall, southwestern England. Material Examined: Great Britain: Cornwall, Fal- mouth, 1 sh., coll. MF, 1974, E55A; Channel Islands, Herm, 11 shs, coll. MF, 09.1974, E54A. Atlantic France: Normandy, Carteret, 3 shs, coll. MF, Jul. 1973, E28A; lower Normandy, St. Pair, 7 shs, MNHN ain Denis, 1945; Brittany, Finistere Anse de Bertheaume, 20-30 m, industrial dredging, 4 shs, MNHN coll. S. Gofas, 1978; Brittany, Cote-du-Nord Plomanach, fissures of infralit- toral rocks, 1 sh., MNHN coll. S. Gotas, 1973-78: Brit- tany, Finistere Roscoff, “les Cochons Noirs”, sand and conchiferous gravel, 20 m, 27 shs, MNHN coll. Gofas, Jul. 1994, Brittany, St. Lunaire, 20 shs, MNHN Coll. Fischer, 1898: Brittany, St. Servan, 10 shs, MNHN coll. Staadt, 1969: Brittany, St. Lunaire, 2 shs, MNHN coll. Ph. Dautzenberg ( figured i in Bucquoy - - 1854, pl. 35, figs. 1, 2); St. Lunaire, 5 shs, MSNCS, 7173 and 7174, 20 Jun. 1970, on the beach at low tide; fan uy, Morlaix, Saint Michel en Gréve, 2 shs, coll. MF, 1976, E25A; Brittany, Saint Jacut, 3 shs, coll. MF, 06.1975, E12A; Brittany, Saint Jacut, 14 m, 6 shs, coll. MF, 04.1974, E12B: Britt: iny, Carnac, Quiberon, 4 shs, coll. MF, 1970, E16B; Brittany, Carnac, Quibe ron, Pointe de Couquel, 2 shs, coll. MF, 1970, ELIA: Brittany, Saint Malo, 7 shs, coll. MF, 07.1973, EL3A; Brittany, St. Malo, Lizardrieux, 1 sh., coll) MF, Aug, 1982, E69A. Atlantic Pyrenees, Aquitaine, St. Jean de Luz, Cote a infralittoral rocks, | sh., MNHN coll. S. Gofas, 19S0-S1; Atlantic ae Aquitaine, St. Jean de Luz, 73 shs, MNHN coll. H. Fischer, 1898: Aquitaine, Soulac, 1 sh., MNHN coll. A. Dolfus; Aquitaine, Hendaye, 2 shs, coll. MF, Jul. 1976, E57A. Portugal: Algarve Sagres, Baie de Baleeira, (37°00.7' N, 08°55.0' W), tide zone, 1 sh., MNHN, Mis- sion Algarve, May 1988; Algarve Sagres, Ponta da Ba- leeira, (37°00.3' N, 08°55.5' W), 17-23 m, 5 shs, MNHN, Mission Algarve, May 1988; Algarve Sagres, Pontal dos Corvos, (37°01.3' N, 08°58.3' W), at the foot of falaise, 17-22 m, 5 shs, MNHN Mission Algarve, May 1988; Albufeira (southern coast), Ponta de Castelo, 3-6 m, 2 shs, coll. MF, OS Aug. 1985, E50B. Atlantic Moroc- co: asilah: mouth of Oued el Helou, conchiferous depos- its, beach, 6 shs, MNHN coll. S. Gofas, 1971-72. Strait of Gibraltar: Tanger, Grande Plage, conchiferous deposits, beach, 3 shs, MNHWN coll. S. Gofas, L970-S1; Cadiz, Getares, beach, 3 shs, coll. MF, ex coll. C. Bogi, legit Hanselaar, 2230 GET; Cadiz, Barbate, conchiferous de- posits, beach, 6 shs, MNHN coll. S. Gofas, 1976-S1; south Ceuta, Punta del Desnarigado, (35°53,.6' N, 05°16.8' W), 16-20 m, 1 sh., MNHN coll. Bouchet, Go- fas and Lozouet, May 1996. Mediterranean Spain: Cas- tellon, Columbretes Islands, Espinosa Island, 5 m, 1 sh., coll. MF, 26 Jul. 1974, M273A; Malaga, 15 m, 7 shs, coll. MF, ex coll. Cesare Bogi, 2127IMA(V002G); Malaga, Algeciras, Torre del Almirante, 3-5 m, | sh., coll. MF, 28 THE NAUTILUS, Vol. 122, No. 1 V. Garilli, 2008 Page 23 Figures 9-14. Alvania carinata (Da Costa, 1778), protoconch and details of sculpture. 9. Sicily, Palermo, Terrasini, “Magaggiare- Ciucca di Cinisi” beach, coll. tiene 146E) , protoconch. 10. Provence, Marseille, La Baule, small beach at 25 km west from Marseille, coll. PAL (212SBAU-VO008C), protoconch. 11-12. Shell from the same locality and collection, sculpture of protoconch T and II (11) and a detail of protoconch I (12). 13. Same shell as Figure 2, detail of teleoconch sculpture on the penultimate whorl, coll. PAL 307B). 14. Detail of teleoconch sculpture on the last whorl, NW Sicily, Palermo, Terrasini, “Magaggiare-Ciucca di Cinisi” beach detritus of Miniacina, coll. PAL (146E). Scale bars: 100 wm in Figures 9-11 and 13-14; 50 zm in Figure 12. White arrows indicate the protoconch/teleoconch boundary, respectively. Sep. 1976, M207B; Malaga, Algeciras, 3 shs, coll. MF, 23 gloue, (43°10.6' N, 05°24.2' E), 33 m, 31 shs, MNHN Sep. 1976, M207A; Malaga, Fuengirola, 0.5-1 m, 1 sh., rec. H. Zibrowius Jun, 1996; Provence, Iles Embiez, con- coll. MF, 20 Aug. 1973, MG6SB; Malaga, Algeciras, Playa chiferous deposits, beach, 10 shs, MNFIN coll. S. Gofas, Getares, 6 m, 1 sh., coll. MF, Aug. 1983, M207E; 1968-70; Provence, Iles Embiez, Petit Rouveau, dredg- Malaga, Cabo Pino, detritus, 10 m, 2 shs, coll. § — ing of sandy conchiferous bottom, 3-5 m, 5 shs, MNHN (41.80g). Mediterranean Morocco: M’diq “(anc. coll. S. Gofas, 1968-70: Provence, St. Clair, infralittoral Rincon)” conchiferous deposits, beach, 1 sh., int rocks, (43°08.2' N, 6°23.2' E), 0-1 m, 1 sh., MNHN rec. coll. S. Gofas, 1971. Algeria: Alger, 1 sh., MNHWN coll. S. Gofas, Sep. 1992; Provence, St. Raphael, 2. shs, Locard. Mediterranean France: Languedoc, Carnon, MNHN coll. Locard: Provence, 9 shs, MNHN coll. Petit, conchiferous deposits, beach, 1 sh., MNHWN coll. S. Go- 1873; Provence, Sanary, | sh., MNHN coll. Locard:; Cor- fas, Aug. 1976: Languedoc, Roussilion, 3 shs, MNHN sica, Ajaccio, 3 shs, MNHN coll. Jousseaume, 1921; Cor- Coll. Dolfus, 1903; Languedoc, Roussilion, 1 sh., MNHN sica, Galeria, Punta Stollo, 6 m, 1 sh., coll) MF, 07 Aug coll. Ph. Dautzenberg (Moll. du Roussillon); Provence, 1984, M52D: Corsica, Pianottoli, Anse de Chevanu, 2 Marseille, La Baule, small beach at 25 km west from shs, coll. MF, Jun. 1988, M77A. ee Djerba, Aghir Marseille, 3 shs, coll. MF, ex coll. C. Bogi, Oct. 1986, 5 m, Posidonia bed, 1 sh., coll. MF, S. Palazzi legit 06 212SBAU (VOOSC); Provence, Marseille, Cape Cou- Aug. 1993, M79A; Djerba, Al Jazirah, 1- ~2.5m, | sh., coll ronne, 4 shs, coll. MF, ex coll. C. Bogi, Oct. 1987, MF, Aug. 1974, M9B. Italy: Friuli Venizia Giulia, Tri- 2129COU (VOOSE): Provence, Marseille, Grand Con- este, beach, 7 shs, coll. MF, D. Di Massa legit 1976 Figures 1-8. Alvania carinata (Da Costa, 1778), variation in shell shape and sculpture. 1. “Form” ecarinata Bucquoy et al., 1554 shell from coll. Ph. Dautzenberg (Moll. du Roussillon), MNHN-DSE. 2. Typical, keeled morph, Liguria, Genova, Camogli, 42 m coll. PAL (307B). 3. Markedly keeled morph, corresponding to Rissoa trochlea Michaud, 1830, Strait of Gibraltar, Cadiz, Getares beach, coll. PAL (2230 GET). 4-5. Small, unkeeled morph, La Spezia, Monterosso, 15-30 m, coll. PAL (70A). 6. Typical morph with varice, Sardinia, Sassari, Capo Caccia, Cala della Calcina, 6 m, coll. PAL, (112A). 7. Profile view of the same shell as Figure 2. 8. Juvenile shell fitting well with the concept of Alvania cingulata (Philippi, 1836) sensu Piani (1979, figs. 2-3). Scale bars: 1 mm in Figures 1-7: 0.5 mm in Figure 8 Page 24 THE NAUTILUS, Vol. 122, No. 1 307B; Liguria, Genova, Camogli, 42 m, 1 sh., coll. MF, 06.1981, 271B; La Spezia, Monterosso, 15—30 m, detri- tus, 1 sh., coll. MF, A. Lugli legit Oct. 1978, TOA; La Spezia, Portovenere, 5-25 m, 3 shs, coll. MF, Oct. 1978, 329A; La Spezia, Riomaggiore, 30 m, 1 sh., coll. MF, Aug, 1975, 312A; Liguria, La Spezia, Punta Mesco, 35 m, L sh., coll. MF, Jul. 1987, 176C; Tuscany, Livorno, San Vincenzo, Borraceia, 12 m, 1 sh., coll. MF, 12 Jul. 1987, 190A; Tuscany, Livorno, Secche della Meloria, 6—17 m, 2 shs, coll. MF, 1974, 34E; Tuscany, Livorno, Castiglion- cello, 4 shs, coll) MF, 1972, 21A; Tuscany, Livorno, Romito, off mouth of the torrent Chiona, 30-35 m, 2 shs, coll. MF, ex coll. C. Bogi, 36A; Tuscany, Livorno, Bagni Fiume, 20 shs, coll. MF, 1977, 34B; Tuscan Archipe slago, Island of Capraia, 100/400 m, 2 shs, coll. MF, ex coll. C. Bogi, 2126CAP(VO005G); Island of Capraia, Punta della Fica, 29 m, 1 sh., coll. MF, 15 Sep. 1985, 25R; Island of Capraia, Punta Civitata, 40 m, 1 sh., coll. MF, 19 Sep. 1985, 28ST; Tuscan Archipelago, Gorgona Island, 35—40 m, LI shs, coll. MF, Aug. 1978, 69A; Gorgona Island, 40 m, 18 shs, coll. MF, legit C. Bogi, 194; Tuscany, Siena, 1 sh., yellow sands from unknown layer, lower Pliocene, coll, MF ex coll. PAL, F22A; Tuscany, Grosseto, Punta Ala, Baia Verde, 5 m, 1 sh., coll. MF, G. Terzer legit OS Sep. 1974, 27A; Tuscany, Grosseto, Punta Ala, Punta Hidalgo, 2 shs, coll. MF, Jun. 1975, 27B; Tuscany, Gros- seto, Follonica, Cala Felice, 7 m, 1 sh., coll. MF, 25 Aug. 1987, 55B: Tuscany, Grosseto, Island of Palmaiola, 2S m, 2 shs, coll. MF, 14 Sep. 1986, 172A; Grosseto, Island of Elba, Capo Calamita, Scogli Corbelli, 46 m, 1 sh., coll. MF, Sep. 1972, 5E; Island of Elba, Scoglio Remaiolo, 35 m, 1 sh., coll, MF, May 1980, 5k; Island of Elba, For- miche della Zanca, 20 m, 1 sh., coll. MF, Apr. 1954, 5P; Lazio, Rome, Civitavecchia, 2 shs, coll. MF, 1975, 137B; Sardinia, Sassari, Capo Caccia, Cala della Calcina, 6 m, conchiferous detritus at upper limit of Posidonia bed, 10 shs, coll. MF, S. Palazzi legit 19 Aug. 1983, 112A; Sar- dinia, Sassari, bay at E of Faro di Capo Testa, 7-19 m, 1 sh., coll. MF, 22 Aug. 1983, 125B; Nuoro, Capo Comino, Ruia Island, 0-2 m, | sh., coll. MF, 1974, 75A; Sardinia, Cagliari, Island of San Pietro, channel of San Pietro, 2-4 m, Posidonia bed, 2 shs, coll. MF, G. Liuzzi legit 10 Oct.1976, 264A; Sardinia, Cagliari, Island of Sant Antioco, Cala de Saboni, 1 sh., coll. MF, O7 Aug. 1983, LI5B:; Sardinia, Cagliari, Island of Sant Antioco, Cala de Saboni, 14 m, 1 sh., coll. MF, 10 Aug. 1983, LI5E; Campania, Napoli, Island of Capri, Punta Vivara, 6 m, 1 sh., coll. MF, 0S Sep. 1978, 254B; Campania, Napoli, Island of Procida, Marina Grande, 2.5-9 1m, 2 shs, coll. MF, 1974, LO5A; Puglia, Bari, off Palese, 12-13 m, 1 sh., coll. MF, 1] Sep. 1979, 77A; Puglia, Taranto, 2 shs, coll. MF, 1973, 84B; Puglia: Taranto, Campomarino, 2 shs., coll. MF, 12 Feb. 1977, 309A; Puglia, Taranto, Maruggio, 1 sh., coll. MF, 1977, 280A; Puglia, Brindisi, “Batteria Brin” beach, 14 shs, coll. MF, G. Oriolo legit 08.1970, 193D; Puglia, Brindisi, Punta Croce, 15 m, 19 shs, coll. MF, G. Oriolo legit Aug. 1974 262A; Puglia, Brindisi, Torre Guaceto, 5 m, I sh., coll. MF, 1976, 42A; Brindisi, Lendinoso, 10-20 m, 1 sh., coll. MF, 1977, 279A; Puglia, Lecce, Gallipoli, Costa Brada, 2 shs, coll. MF, Apr. 1978, 91B; Lecce, Porto Cesareo, 2 shs, coll. MF, Oct. 1977, 46B; Lecce, Porto Cesareo, Torre Lapillo, 1 sh., coll. MF, 06 Jun. 1978, 46C; Lecce, Marina di Ugento, 30 m, 1 sh., call MF, 1977, 6SC: Lecce, Gallipoli, La Vecchia Torre, 2-6 m, 2 shs, coll. MF, 1976, 91D; Lecce, San Cataldo, 1 sh., coll. MF, Apr. 1973, 260C; Lecce, Speechnulla, 1 sh., coll. MF, Oct. 1974, 281A; Calabria, Reggio Calabria, Pen- timele, from fisherman nets, 8 m, 1 sh., coll. MF ex coll. Sciano, VOG0A 194; Reggio Calabria, Laureana di Bor- rello, Pecoraio, 2 shs, (laver 2), lower Pleistocene, coll. MF ex coll. PAL, F24A; Sicily, Trapani, San Giuliano, 10 shs, coll. MF, 25 Jul. 1978, 297A; Trapani, Scopello, Ton- nara, 12-20 m, 1 sh., coll. MF, 06 Apr. 1986, 333F; Sicily, Trapani, Egadi Islands, Favignana, Secea del Toro, 30 m, 20 shs, coll. MF, A. Lugli legit 02 Jun. 1983, 272H; Egadi Islands, Favignana, 4 m, 17 shs, coll. MF; Egadi Islands, ae Cala Rotonda, 20-30 m, 3 shs, 04 Jun. 1983, coll. MF, 272G; Egadi Islands, Favignana, eteene Cor- rente, 30 m, 3 shs, coll. MF, O1 Jun. 1983, 272F; Egadi Islands, Favignana, Punta Sottile, 30-40 m, 4 shs, coll. MF, 30 May 1983, 272E; Egadi Islands, Marettimo, Punta Bassana, 41 m, 2 shs, coll. MF, 04 Jun. 1983, 140A; Egadi Islands, Levanzo, Faraglione, 7-12 m, 3 shs, coll. MF, 03 May 1979, 325B: Sicily, Pantelleria Island, Baia dei Fichi d’India, 33. m, 1 sh., coll. MF, Jul. 1983, 154M: Pantelleria Island, Punta Capace, 31 m, 1 Se coll. MF, Jul 1983, 154N; Sicily, Palermo, 1 sh., coll. MF ex coll. . Bogi, (VO5G6A) 194; Palermo, Bagheria, Aspra, 15.5 m, 7 chs. coll. MF, Apr. 1973, 124C; Pz ilermo, Punta Raisi, Marina Longa, 1 sh., coll. MF, 28 Feb. 1979, 320A; Pal- ermo, Terrasini, “Magaggiare-Ciucea di Cinisi” beach, detritus of Miniacina, 14 shs, coll. MF, S. Palazzi legit 23 Sep. 1977, 146E; Palermo, Island of Ustica, Scoglio del Medico, 25 m, 1 sh., coll. MF, Aug. 1980, 286k; Sicily, Messina, Milazzo, Capo Milazzo, Cala SantAntonio, 2 shs, upper yellow sands, upper Pleistocene, coll. MF ex coll. PAL, F5A: Messina, Eolie Islands, Lipari, Secca del Bagno, 38-40 m, 3 shs, coll. MF, 05 Se p. 1979, 338A; Sicily, Cat: mia, Acitrezza, 3-25 m, 2 shs, coll. ME. Sep. 1975, 22A: Sicily, Siracusa, Vendicari, bay, 2 shs, coll. MF, 13 Sep. 1977, 20A: Siracusa, Portopalo di C Capo Passero, 2-3 m, | sh., coll. MF, Jun. 1976, 63A; Siracusa, Peninsula of Magnisi, southern side, 1 sh., coll. V. Garilli, Jun, 2006; Sicily, Palermo, Tommaso Natale, 1 sh., late middle Pleistocene, MGUP 166/2/49; Sicily, Trapani, Birgi, 3 shs, upper Pleistocene, Tyrrhenian Stage, MGUP. 1765/36/14: Sicily, Trapani, 12 shs, upper Plei »18- tocene, Tyrrhenian Stage, MGUP. 358/3/42. Isle of Malta: Malte, 8 shs, MNHWN coll. Jousseaume, 1921. Croatia: Istria, Rovinij, between Rt. Muntrav, Hr. Mun- trav ancl Azino, S—L5 m, detritus from bottom, 11 shs, coll. MF, Palazzi legit 19 Jun. 1975S, MISSB; Istria, Savudrija, i. ach, 8 shs, coll. MF, S. Palazzi legit Oct. 1975, M364A; Istria, Rovinij, 6-18 m, 19 shs, coll. MF, 20 Jun. LOTS, MISSA; Istria, Vrsar, | sh., coll. MF, 03 V. Garilli, 2008 Page 25 Apr. 1958, M7G6A; Istria, Umag, 1 sh., coll. MF, 1978, M1O9A; Istria, Umag, Taverna Lero, 0.3 m, 1 sh., 17 Jun. L978, coll. MF, MLO9B; Kaciack, dam of Cigale, 4m, 1 sh., 29 Jul. 1983, coll. MF, MI4A. Greece (: Aegean Sea): NW Aegean, Island of Limnos, near Moudros, 3-5 m, 48 shs, coll. MF, A. Lugli legit Aug. L991, MSOA; Sithonia, Ormos Panagias, 2-3 m, 1 sh., Aug. 1982, coll. MP, M31B; Sithonia, Nesis Dhiaporos, 33-34 m, 2 shs, coll. MF, 01 Aug. 1986, M31D. Unknown locality from Medi- terranean Sea: 120 shs, MNHN coll. Vayssiére. Habitat: The species is usually found at depths com- patible with the upper part of the shelf, the infralitoral stage of Pérés and Picard (1964). Rarely, I found material collected from the tidal zone. According to Gofas and Ponder (1991), Alvania carinata lives deeply buried un- der stones. It seems also to be linked (in the Mediterra- nean) to phanerogam beds. The finding of two shells from the Island of Capraia (Tuscan Archipelago), col- lected at a depth of 100-400 m, is much pre sbably due to lower shelf-slope transport. Distribution: Occurs probably throughout the Medi- terranean, but its presence in the easternmost coasts, from which I did not see any material, needs to be con- firmed. In the Atlantic it is recorded from the coasts of Great Britain south to Morocco. As fossil, it is rare in the Mediterranean Neogene where it is recorded from the lower Pliocene yellow sands near Siena. It becomes more frequent during the Mediterranean Quaternary, where I found it from the lower Pleistocene of Reggio Calabria, and the middle-upper Pleistocene of Sicily ( (Tommaso Natale, Capo Milazzo and Trapani). Remarks: A complete synonymy list was provided by Piani (1979) and Van Aartsen (1982). In various collec- tions, I found different lots of this species containing juvenile shells (not higher than 3 mm, see Figure §) identified as Alvania cingulata ( (Philippi, 1536), species hereafter re-described and discussed. This wrong deter- mination very likely follows the misidentification of Piani (1979, figs. 2-3) and Giannuzzi-Savelli et al. (1996, figure 499). Typically, the shell of this species has a characteristic keeled shape due to the presence of well marked spiral cords that become very strong on the well-developed last whorl. The number of ee on adult whorls is slightly variable. 34 in the penultimate whorl. The not-ke meled morph, characterized by having a slender and turrited shape and more numerous cords (5-6 in the penultimate whorl), is the “form” ecarinata Bucquoy et al., 1854 (Fig- ures 1, 445), also known as minor-ecarinata Montero- sato, 1884 (probably corresponding to Rissoa lucullana var. cancellata Scacchi, 1836, as indicated by Piani, 1979). It has been considered a Mediterranean subspe- cies of A. carinata by Van Aartsen (1982). I found no shells clearly belonging to this morph in Atlantic material studied (mainly from the MNHN-DSE collec- tions). However, in some Mediterranean areas (e.g. along the French coast in the MNHN-DSE eollertons), I found the typical morph together with unkeeled shells. Thus [am more inclined to consider it just a case of intraspecific variation, which appears well represented in Mediterranean populations. The keeled morph, charac- terized by having three spiral cords on subadult whorls and a very expi anded body whorl (more than 3/4 of the shell he ight, see Figure 3) fits well within the concept of Rissoa trochlea Michaud, 1830, which is certainly a syn- onym of A, carinata. Alvania carinata is the type species of Galeodina Monterosato, 1884, a genus created for cingulated, keeled, varicose rissoid shells with a wide aperture (Monterosato, 1SS4a, p. 163). This generic division can appear quite artificial, not being supported by appropri- ate and constant taxonomic features. Also, the consider- ation of Galeodina as a well-established subgenus of Al- vania, as indicated by Wenz (1938) and Van Aartsen (1982), appears doubtful. Ponder (1985), who synony- mized most of the available subgenera with Alvania sensu stricto, expresse d some uncertainty when consid- ering Galeodina as a valid taxon. He recognized the ig- norance of anatomical features as the main gap in solving this systematic question. In any event, the shell chaise ters alone do not provide enough support for Galeodina to stand as a subgenus. For example, the unkeeled morph, which usually lacks the main features of Galeo- dina, including the characteristic varices of the keeled morph, might be compared to the Alvinia Monterosato, 1854, species group, with which it shares a slender, tur- reted shell shape. It is noteworthy that Cossmann (1921) cited Galeodina as a synonym of Alvinia. Alvania cingulata (Philippi, 1836) (Figures 15-26) Rissoa cingulata Philippi, 1S36: 152-153. Acinus cingulatus (Philippi)—Monterosato, 1$54b: 62. Cingula (Onoba) cingulata (Philippi, 1$36).—Nordsieck, 1968: 46, pl. VIL, fig. 26.35 Alvinia (Galeodina) cingulata (Philippi, 1836)—Nordsieck, 1972: 181, pl. RVI, fig. 20. Description: Small, stately, conical-ovate shell reach- ing 3.5 mm in height, 2.2 mm in width. Protoconch pau- cispiral, consisting of little more than 1.5 convex ae very rounded whorls, sculptured by 5-6 very fine spiral lirae. Several prosocirte growth scars precede protoconch/ teleoconch transition (Figure 20). Teleoconch consists of about 4 cingulated convex a separated by slightly inclined sutures. Very early teleoconch whorls bear two main spiral cords crossed by numerous axial ridges. Sculpture of adult whorls consisting of spiral cords more conspicuous than axials, which are 5-6 and 10-11 on the penultimate and body whorls respectively. Axial sculp- ture formed by narrower (half the breadth of a spiral cord) ribs, numbering ae 40 on penultimate whorl, and which become obsolete toward shell base. Micro- sculpture consists of spiral rows of small tubercles that do not seem to appear on main sculpture (Figure 26). A few irregular scars cross these spiral rows. Last whorl rather THE NAUTILL V. Garilli, 2008 Figures 24-29. Alvania cingulata (Philippi, 1536) from type locality (Peninsula of Magnisi, Siracusa, SE Sicily, coll. Garilli), and a slightly resembling Alvania sp., from the lower Pleistocene of Musala (Italy, Reggio Calabria, coll. MF ex coll. Pal, F95A), fitting well with the conce pt of A. cingulata stated by Monterosato (1SS4b). 24- 26. Alvania cingulata \pertural view of a subadult shell 7 5) and microsculpt ure (26). 27- 24), sculpture on last whorl (5 bars: 500 zm in figures 24, 27-28: 100 xm in figures 25, 29; protoce mnch/teleoconch b« vundary : inflated and well-developed, comprising little less than 3/4 total height. Aperture wide and ovate, comprising about 4/9 and 2/3 of total height and last whorl respec- tively. Outer lip slightly prosocline, internally smooth, externally sli chily thickened near edge. Columellar side arcuate, with a thin callus forming a very narrow umbili- cal chink. Coloration consisting of quite large, reddish- brown bands on a whitish-cream background -29. Alvania sp. Apertural (27 20 pm in Figure 26, The white arrow in Figure 29 indicates the ), dorsal (28) views and protoconc h. Seale Type Locality: Peninsula of Magnisi, Siracusa, south- eastem Sicily. Type Material: coll. Philippi was de stroyed by Byne’s reaction (M One possible syntype, ZMB (2326) ex Glaubrecht, pers. comm., 2006) Material Examined: — Italy: NE Sicily, Messina, 2 shs ZMB ex coll. Monterosato, 80914 (original label by Figures 15-23. Alvania cingulata (Philippi, 1836). 15-17. Apertural (15), dorsal (16), profile (17) view of one shell from ZMB ex coll. Monterosato (80914), Messina, NE Sicily. 20, 23. Protoconch of the same shell. 18-19 and 21-22 Another shell from the same lot apertural 18), dorsal (19 profil 21) view, and a detail of sculpture on the last whorl (22). Seale bars: 1 mm in figures 15-19 and 21: 200 pm in Figure 22: 100 wm in Figure 20; 50 ym in Figure 23 teleoconch boundary The white arrows indicate the approximate protocon h Page 28 THE NAUTILUS, Vol. 122, No. 1 Monterosato: Acinus cingulatus, 2 2. Messina, 1014); SE Sicily, Siracusa, Pemneuls of Magnisi, south side, detritus from the beach, 1 sh., coll Garilli, 6/2006 Habitat: Unknown. It is noteworthy that sandy and Posidonia bottoms prevail in the south side of the Mag- nisi Peninsula (from where I collected the beached shell in Figure 24), while an almost paralic (pre-lagoon) envi- ronment (with Cerastoderma, Cerithium, and Gibbula spp., on a finely sandy bottom) and a rocky bottom with algae characterize the north and east sides. Distribution: — Its distribution appears to be limited to Sicily, with particular regard to the eastern coast, from Messina to Magnisi. It was recorded from Palermo by Monterosato (1872: 1875; 1878: 1884a; 1884b), who very likely followed the citations of Philippi (1544) and Weinkauff (1885). The latter author, who indicated Mon- dello (the beach near Palermo), also cited Ognina (Cata- nia) in eastern Sicily. To my knowledge, no f fossil record of this species exists. Remarks: This is a very rare, practically unknown spe- cies (see Van Aartsen, 1982a). Its identification has tra- ditionally been quite problematic, above all after the misidentification by Piani (1979, figs.2-3), who illus- trated a juvenile shell of Alvania carinata (Da Costa, 1778) under the name Galeodina cingulata (Philippi, 1836). The shell figured by Giannuzzi-Savelli et al. (1996, fig. 499), under the name Alvania (Alvania) cingulata (Philippi, 1836), is a subadult of A. carinata. My identification is based on three shells (one from the type locality, two from ZMB ex Monterosato coll. labelled as Acinus cingulatus), which agree well with the original description by Philippi (1836; see also 1544: 128). A few diagnostic characters allow for its correct identification: the primary cingulated sculpture with raised cords, numbering 10-11 on the last whorl; the secondary axial pattern formed by very narrow ribs cross- ing the cords: the large ovate aperture: and the intern: ally eimaadh outer lip, slightly thickened near its edge. As Philippi ee ) noted, Alvania cinoulate may be compared with A. tenera, especially with the slender morph of the ae ‘Yr species (see Figure 78). Alvania te- nera has a smaller shell (usually not over 2.2 mm in height) with more raised cords (numbering 12-14 on the last whorl), and almost lamella-like ribs, which occur just on the interspaces between the cords (see discussion of A. tenera). Furthermore, A. cingulata has a paucispiral pre ytoconch. In addition, Nordsieck (1972) and Van Aartsen (1982) considered this species as belonging to Galeodina. How- ever, the similarity between Alvania cingulata and A. carinata (type species of Galeodina) is superficial. Sev- eral differences separate the two: A. cingulata has a less inflated last whorl bearing less numerous cords and lack- ing varices, its outer lip lacks a varicose thickness, and its spiral Sc at does not markedly dominate over axial as in the typic ral A. carinata. The latter species s also has a multispiral te conch. Monterosato (1584a and 1884b) included this species in his genus Acinus (synonym of Alvania according to Ponder, 1985), creating a link with the type species AL vania cimex (Linnaeus, 1758) (which is also the type species of Alvania, see Ponder, 1985: 36), mainly based on coloration. In reality, the similarity between the two species is limited to the color pattern of the shell and the nodular sculpture. Strong differences indicate that A. cingulata and A. cimex could be distant relatives, the latter having a more sturdy shell with a coarser sculpture (with subequal axial and spiral), and a proportionally smaller aperture provided with an internally denticulate outer lip. The concept of A. cingulata as expressed by Monterosato (1884b: p. 62) seems to be applicable to an interesting Alvania sp. (from the lower Pleistocene of southern Italy, Reggio Calabria, Musala, 19 shs, F95A, coll., Figures 27-29) rather than to the original descrip- tion by Philippi. The non- eo .. see the pau- cispiral protoconch in Figure 29) Alvania sp., illustrated here, showing only a mode srate aie ity with the teleo- conch sculpture of A. cingulata, has the characteristic general shape of A. cimex, with which it shares the ap- ertural features, including the denticulations on the inner part of the outer lip. Furthermore Monterosato himself determined the shells I have illustrated in Figures 15-23 as A. cingulata. In 1968, Nordsieck cited A. cingulata as Cingula (Onoba) cingulata. In reality, a vague similarity with some Cingula Fleming, 1828 or (more specifically) Onoba H. and A. Adams, 1852, nee s exists: e.g. Onoba (O.) carpenteri (Weinkauff) (see Ponder, 1985: fig. 1 14a), bears a similar spiral sculpture. Anyway, the genus Onoba is characterized by usually elongate shell a a proportionally smaller and more rounded aperture. Its included species, usually lacking the axial sculpture, may have a much weaker axial pattern or this may consist of delicate ribs, which become obsolete towards the sutures (see Bouchet and Warén, 1993: figs. 1508-1509, 1514— 1515). In addition, the protoconch sculpture apparent on some non- planktotrophic Onoba species resembles that of A. cingulata, in having few, very fine spiral widely spaced threads. This land of sculpture is also seen in Alvania (e.g. A. subsoluta (Aradas, 1S47), see Bouchet and Warén, 1993, fig. 1458). Cingula species appear to be less similar to A. cingulata in having a thick outer lip and lacking any axial sculptural pattern. A certain similarity exists with Alvania watsoni (Wat- son, 1873) from Madeira (see Ponder, 1985: fig. 102c—d), with which A. cingulata shares the general shell shape, characterized by an inflated last whorl provided with a large, ovate aperture. Alvania francescoi new species (Figures 30-43) Description: — Shell small, sturdy, conical-ovate, reach- ing 4.8 mm in height (4.3 mm in holotype) and 3.1 mm in width (2.8 min in holotype). Protoconch conical, mul- tispiral with partially immersed nucleus and convex V. Garilli, 2008 age ¢ Page 29 whorls. Protoconch I consisting of about 0.8 whorls sculptured by 5-6 fine spiral lirae interspersed spirally and irregul: uly with microscopic granules. Protoconch I consisting of about 1.2 whorls sculptured by pimples, which are irregular in size and arrangement. They are larger and less numerous in the adapical position; fused into very short prosocline tracts in central part. In the ab: :pical part of late protoconch, groups of pimples are irregularly fused into fine, discontinuous spiral ridges. Protoconch/teleoconch transition well-marked and sinu- ous. Teleoconch consisting of 3.84.6 (4.2 in holotype) rather convex whorls, with strong cancellate sculpture formed by intersection of equally Oe veloped spiral cords and axial ribs. Cords are narrow and well raised, num- bering 2, 6-7 (6 in holotype) and 13-15 (14 in holotype) in early, penultimate and body whorl, respectively. Two ads :pical cords are usually more pronounced in penulti- mate and last whorl. Cords are markedly stronger and more widely spaced on base. Secondary cords may com- mence on late body whorl, at a certain distance from outer lip. Axial seuptite consists of narrow raised ribs numbering 24—32 (26 in holotype) on the penultimate whorl. On base, ribs progressively become very narrow to obsolete toward columellar area. Intersection of spiral and axial sculpture, producing nodular small knobs, forms a rectangular pattern (with major side of rect- angles perpendicular to shell axis) except on last whorl, where a quadrangular to rhomboidal pattern occurs (Fig- ure 43). Microsculpture of very early teleoconch whorls consists of micropustules, sometimes fused, for ming spi- rally, discontinuous and irregular rows, mainly occurring between spiral cords (Figure 42); weak growth lines are present. Fine sculpture of the rest of teleoconch limited to weak growth lines Sa aie 43). Suture slightly in- clined. Last whorl well-developed, comprising about three fourths of shell height. It may bear single (paratype 6) or double (paratype 4, Figure 37) varices, at angles of about 50° and 70° respectively. Aperture wide, ovate, and with a rounded profile in the posterior part, com- prising about two thirds to three fourths of last whorl height. Outer lip orthocline, internally smooth, externally markedly thickened by a strong, sometimes doubled (Figure 31), rim, occurring very close to its edge and covered by spiral cords. Inner lip moderately arcuate and rather thickened, with thin callus c lelimitating very nar- row umbilical chink. Type Locality: Lower Pleistocene of Cartiera Mulino 36°56'57" N, 14°34'03" E), Vittoria, Ragusa, SE Sicily. The stratotype is the phanerogams-rich 3D1 layer of Costa (1989). This deposit crops out at about one hun- dred meters from the abandoned paper-mill known as Cartiera Mulino. Paleoenvironmental and stratigraphic information about this site are given by Costa (1989). Type Material: Holotype (4.3 x 2.5 mm), DGUP CMRG 005/488, coll. V. Garilli. Paratype 1 (4.1 = 2.7 mim), DGUP CMRG 006/489, coll. V. Garilli; paratype 2 4.8 x 3mm), ZMA Moll. 4.07.007 (ex CM GR 007/490, coll. Garilli); paratype 3 (4 x 2.7 mm), ZMB MB.Ga.2479 (ex CMGR 008/49, coll. V. Garilli); paratype 4 (4.2 « 2.9 mm), MNHN A25950 (ex CMGR 009/492, coll. Garilli); paratype 5 (4.6 x 3.1 mm), DGUP CMGR 010/ 493, coll. V. Garilli; paratype 6 (4.5 x 3.1 mm), DGUP CM GR 011/494, coll. V. Garilli. All type material is from the type locality, 3D1 layer. DATE? Etymology: The species honors the name of both Francesco Garilli senior, my father and first mentor in my life, and of Francesco Garilli junior, my son. Habitat: All the shells came from the 3D1 layer of Costa (1989). The paleoenviromental reconstruction for this layer was characterized by Costa (1989) as compa- rable with the Mediterranean marine-marginal modern ecotone HP (Posidonia beds)-SVMC (sensu Pérés and Picard, 1964). It is interesting that the 3D1 layer is quite rich in Posidonia remains mainly consisting of leaves. Distribution: This species is known from type locality only, lower Pleistocene of Cartiera Mulino, Vittoria, Ra- gusa, SE Sicily. Remarks: This quite characteristic species shows a ga- leodiniform shape, having a quite inflated and well- developed (also with double varix) last whorl and a large aperture provided with an intermally smooth outer lip. However, its spiral sculpture does not markedly prevail over the axial, as in the typical Alvania carinata. Its mi- crosculpture, limited to the early teleoconch whorls, is similar to that observed in the analogous whorls of A. carinata, but in the latter this microsculpture is distrib- uted over the entire teleoconch. The strongly cancellated share pattern of A, francescoi new species, consisting of well-raised, sub- equal spiral and axial elements forming marked nodular intersections, resembles that shown io A. cimicoides (Forbes, 1844). The latter species, showing a more typi- cal Alvania shape, has a more slender shell, a less inflated and much less developed last whorl, and a proportionally smaller aperture with denticulations on the inner part of the outer lip. Alvania francescoi new species can also be compared with A. rosariae new species described later in this re- port. Remarks on their similarities and differences will be dealt with in the section dedicated to the latter spe- cies. Alvania lactea (Michaud, 1830) (Figures 44-49 and 53-61) Rissoa lactea Michaud, 1830: 9-10, figs. 11-12 Rissoa (Massotia) lactea Michaud.—Bucquoy et al., 1884: 298, pl. 25, figs. 7-13 Massotia Daje »rleini Monterosato ex-Schwartz ms, 1889: : Massotia lactea (Michaud) forma Dajerleini aoe 1917: 12. Description: Shell sturdy, conical-ovate to cylindrical- ovate, occasionally almost ‘pupoid, reaching 5.2 mm in YAUTILUS, V. Garilli, 2008 Page 31 Figures 38-43. Alvania francescoi new species, protoconch, teleoconch sculpture and microsculpture, lower Pleistocene of Cartiera Mulino, SE Sicily, Ragusa, Vittoria. 38. Protoconch, paratype 2. 39. Protoconch, transition between protoconch | and. IT, holotype. 40. protoconch 1, holotype. 41. Sculpture of early teleoconch whorls, note how the spiral microsculpture becomes lacking, paratype 2. 42. Microsculpture on the first whorl, paratype 2. 43. Detail of sculpture on the last whorl, note the numerous growth lines, paratype 4. Scale bars: 200 xm in figures 41 and 43; 100 jzm in figures 38-39: 50 jm in figures 40 and 42. Black and white arrows indicate the protoconch I/protoconch H and protoconch/teleoconch boundaries, respectively height and 3.3 mm in width. Protoconch conical, consist- ing of about two convex whorls. Nucleus partially im- mersed. Protoconch I consisting of about 0.8 whorls, sculptured by 5-7 spiral lirae and inte rspersed with nu- merous closely packed granules, which are spirally and irregularly arranged. Protoconch II consisting of about 1.2 whorls sculptured by pimples irregular in size and arrangement. Pimples are fused into short inclined lines opisthocline and prosocline, Figures 53-54) or form one to three discontinuous spiral ridges in abapical aspect of whorl. Protoconch/teleoconch transition well-marked and sinuous. Large shells consist of 4.2-5 weakly to rather convex whorls. These may be sculptured by a finely cancellate pattern formed by the intersection of spiral cords and numerous delicate, narrow axial ribs, or by opisthocline, pronounced ribs. Ribs often start from early whorls and are crossed by finer spiral threads. In cancellate shells, cords and ribs are equal to subequal and numbering up to nine and 40 respectively on pen- ultimate whorl. In ribbed shells, cords and ribs are less numerous, numbering up to eight and 16 respectively on penultimate whorl. In early teleoconch whorls micro- sculpture consists of fine, irregular, and discontinuous spiral threads formed by groups of small granules, not covering main spiral sculpture: remaining teleoconch with a finely reticulate ultrastructure cove ring all primary sculpture, formed by intersection of very narrow and raised lamella-like axial ridges and very thin spiral threads. Suture slightly inclined and de ‘eply impressed. Body whorl well-exp: anded, comprising 4 of shell height. It may bear a varix, usually with angle of LS0°; rarely a double varix m: ty occur, especially in slender and cancel- late shells. Aperture wide, ovate to almost pyriforme, rounded on its posterior aspect, narrowed anteriorly, comprising two thirds of last whorl height. Outer lip or- thocline, internally smooth, externally thickened, mainly somewhat behind its edge, and covered by primary spiral sculpture. Sometime, in ribbed morphs, the outer lip descends vertically, so that last whorl appears ri ither cylindrical. Inner lip weakly arcuate, with a thin to rather Figures 30-37. profile (34) and dorsal (35) view; 36-37. Pages whorl. Scale bars: 1 mm Alvania francescoi new species, holotype a two paratypes, lower Pleistocene of C: Ragusa, Vittoria. 30-32. Holotype. shell in apertural (30), profile tiera Mulino, SE Sicily and dorsal (32) view. 33-35. Piece shell in apertural (33 apertural (36) and dorsal (37) view, the last showing a double varix on the last THE NAUTILUS, Vol. 122, No. 1] V. Garilli, 2008 Page 33 Figures 53-61. Alvania lactea (Michaud, 1830), protoconch and teleoconch sculpture and microsculpture. 53-54. Profile and dorsal views of protoconch from the same shell as Figure 44, Italy, Tuscany, Siena, Poggibonsi, Villa Pietratitta, podere Melograni, lower Pliocene. coll. MF ex coll. PAL (F55E). 52. Protoconch illustrating variation of the abapical spiral ridges, Italy, Tuscany, Siena Poggibonsi,. Villa Pietrafitta podere Sant Uliviere, lower Pliocene, coll. MF ex coll. PAL (F55C), 56-57. Sculpture of pre stoconch | 56) and early teleoconch whorls (57) of the same shell as figures 44 and 53-54. 58-61. Early (58) and very early (59) teleoconch sculpture, and microsculpture of first (60) and last whorls (61), from the same shell as Figure 42, Monastir-Khenis, coll. Garilli. Scale bars: 200 jm in figures 57-58 and: 100 wm in figures 53-55, 58, 61; 50 wm in Figure 60; 20 wm in Figure 56. Black and white arrows indicate the protoconch I/protoconch II and protoconch/teleoconch boundaries, respectively thick narrow callus, leaving a very small umbilical chink. calities, all belonging to the French Mediterranean coast Very fresh shells show a ferruginous periostracum. Shell “Aode, Cette (Hérault) [now Séte], Callioure, Port- color white Vendre (Pyrénées orientales)” (With this citation I con- Type Locality: Michaud (1531) cited the following lo- sider the second edition, consulted in the MNHN-DSE Figures 44-52. Shells of Alvania lactea (Michaud, 1830) and Alvania prusi (Fischer, 1877). 44-49. Alvania lactea. 44. Italy Tuscany, Siena, Poggibonsi, Villa Pietrafitta, Podere Melograni, lower Pliocene, coll. MF ex coll. PAL (F55E), cancellated morph 45. Tunisia. Monastir-Khenis, coll. Garilli, ribbed morh (var. dajerleini Monterosato). 46. Italy, Piemonte, Asti province, Pliocene MGUP coll. Doderlein (113B) shell close to the original description by Michaud (1830). 47. Italy, Piemonte, Alessandria, Tortona Miocene (Tortonian), MGUP coll. Doderlein (476B). 48. Profile view of the same shell as Figure 41. 49. Juvenile shell, Mediter ranean France, Provence, Marseille, La Baule, coll. PAL, (212SBAU-VO0SC). 50-52. Syntype of Rissoa prusi Fischer, 1877 Quaternary of Rhodes, MNHN (DHT) coll. D’Orbigny (RO7495 apertural 50 profile 51) and dorsal (52) views. Scale bars 1 mm Page 34 THE NAUTILUS, Vol. 122, No. 1 library, of Michaud’s work “Descriptions de plusieurs . . published f or the first time in 1830. The date 1831 may be doubtful, 1832 being the most commonly cited date; I prefer to follow Palazzi ( 2003), who provide od helpful reasons to choose the former date.) Material Examined: 37 shs, coll. H. Fischer, with no locality. Atlantic France: Normandy, St Aubin Calvados, 2 shs, MNHN; Normandy, St.Vaast, | sh., MNHN; Brit- tany, Finistére, Anse de Dionan, under stones covered with sand, 75 shs, MNHN legit. S. Gofas, 1973-78; Brit- tany, St. Lunaire, 40 shs, coll. MNHN coll. Dollfus, 1903; St. Lunaire, 15 shs, MNHN coll. Fischer; Brittany, Penthiérre, 4 shs, MNHN, P. Bouchet legit; Penthiérre (Morbihan), under stones covered with sand, low tide, 10 shs, MNHWN coll. P. Bouchet; Penthiérre, 10 shs, MNHN, 27 Apr.1975; Aquitaine, Cote Basque, Hen- daye, conchiferous detritus, beach, 5 shs, MNHN coll. S sofas, 1981; Cdte Basque, Hendaye, infralittoral rocks, 3 shs, MNHN coll. S. Gofas, 1980-S1; Céte Basque, Ond- arroa, infralittoral rocks, 1 sh., MNHWN coll. S. Gofas, 1980-81; Cote Basque, St. Sebastian, infralittoral rocks, 1 sh., MNHN coll. S. Gofas, 1980-81; Cdte Basque, St Jean de Luz, infralittoral rocks, 57 shs, MNHN coll. S. Gofas, 1980-81; St Jean de Luz, outside Cape Ste Barbe, tide zone, stones covered with sand, 4 shs, MNHN coll. S. Gofas, Dec. 1988; St Jean de Luz, outside cape Ste Barbe, tide zone, stones covered with sand, 49 shs, MNHN coll. S. Gofas, 1989; St Jean de Luz, 54 shs, MNHN coll. H. Fischer, 1898: be ee Guéthary, 7 shs, MNHN coll. H. Fischer, ee Portugal: Algarve Sagres, Pontal dos Corvos, (37°01.3' N, 08°58.3' W), foot of falaise, 17-22 m, 1 sh., MNHN Micsone Algarve, O05. 1988; Algarve Sagres, Baie de Baleeira, (37°00.7' N, 08°55.0' W), tide zone, 1 sh., MNHN Mission Algarve, May 1988. Ailaniic Spain: Cantabria, Orifion prov. Santander, Punta de Sonabia, infralittoral rocks, 1 sh., MNHN coll. S. Gofas, May 1989; Asturias, Muros prov. Oviedo, playa de la Liana, infralittoral rocks, 2. shs, MNHN coll. J. Ortea-S. Gofas 08.89; Cadiz, Barbate, (36°10.9' N, 05°56.9' W), tide zone, infralittoral rocks, 4 shs, MNHN réc. S. Gofas Apr. 1994; Cadiz, Chiclana, (36°22.5' N, 06°12.5' W), tide zone, infralittoral rocks and sands, 12 shs, MNHN réc. S. Gofas, Apr. 1994; Cadiz, Barbate, conchiferous detritus, beach, 3. shs, MNHN coll. S. Gofas, 1976-81; Mediterranean Spain: Malaga, Calahonda, conchiferous detritus, beach, 9 shs MNHN coll. S. Gofas, 1976-81; Malaga, Port de Mar- bella, conchiferous detritus, beach, 3 shs, MNHN coll. S. Gofas, 1978-81; Malaga, Benalmadena-Costa, conchifer- ous detritus, beach, 3 shs, MNHWN réc. S. Gofas, 1991— 93. Atlantic Morocco: E] Jadida, (33°16' N, 08°29’ W), large beach, rocky platform, tide zone, 16 shs, MNHN réc. S. Gotas, 26 Sep. 1991; Fedala, Mannesmann beach, conchiferous detritus, beach, 20 shs, MNHWN coll. S. Go- fas, L970—72: Asilah, mouth of Oued el Helou, conchif- erous detritus, beach, 21 shs, MNHWN coll. S. Gofas, 1971-72: Asilah, mouth of Oued el Helou, conchiferous detritus, beach, 30 shs MNHN coll. S. Gofas, 1972-80: Temara, (33°55' N, 07°00’ W), Sables @’Or beach, rocks and mud, 0-2 m, 24 shs, MNHN, MA4S, réc. S. Gofas, 17 Sep.1991; Essaouira (formerly Mogador), (31°31' N, 09°47' W), rocky platform, tide zone, 1 sh., MNHN, MAAS, réc. S. Gofas, 23 Sep.1991; Rabat, Lahlou, (34°02' N, 06°51’ W), conchiferous detritus, beach, 2 shs, MNHN réc. S. Gofas, 28 Sep. 1991; Essaouira (formerly Mogador), 4 shs, MNHN; Strait of Gibraltar, Morocco: Tanger, Grande Plage, conchiferous detritus, beach, 8 shs, MNHWN coll. S. Gofas, 1970-81. Strait of Gibraltar, Spain: Ceuta Nord, Benzu, infralittoral rocks, 2 shs, MNHN coll. S. Gofas 1976-1981. Algeria: Oran, 6 shs, MNHN coll. Locard. Mediterranean France: Langue- doc, Roussilion, Banyuls sur Mer, near the beach de Paulilles, infralittoral rocks, 1 sh., MNHN coll. Bouchet and Gofas, Sep. 1980; Languedoc, Roussilion, 7 shs, MNHIN coll. Ph. Dautzenberg, (figured in Moll. Rouss. T. I pl. 35, figs.7-13); Languedoc, Roussilion, Séte, 6 shs, MNHN coll. Locard; Provence, Toulon, 8 shs, MNHN coll. Petit; Provence, Cannes, 21 shs, MNHN coll. Doll- fus, 1903; Provence, east coast, Iles Embiez, passe du Gaou, under stones covered with sand, 0-1 m, 5 shs, MNHN réc. S. Gofas, Aug. 1988; Provence, east coast, Iles Embiez, (43°04.3' N, 5°47.4" E), pase du Gaou, under stones covered with sand, 0-3 m, 2 shs, MNHN réc. S. Gofas, Aug. 1988; Provence, Iles Embiez, (43°04.3' N, 5°47.4' E) passe du Gaou, rocks, photophile algae under stones covered with sand, 0-3 m, 1 sh., MNHN réc. S. Gofas, Jun. 1995; Hes Embiez, cote Nord et Petit Rouveau, infralittoral rocks, 3 shs, MNHN coll. S. Gofas, 1968-70: Provence, Les Embiez, cote Nord et Petit Rouveau, (43°05' N, 5°47’ E), rocks, photophile algae, 0-1 m, 3 shs, MNHN réc. S. Gofas, Jun. 1995; Iles Embiez, conchiferous detritus, beach, 15 shs, MNHN coll. S. Gofas, 1968-70; Provence, St Clair, (43°08.2’ N, 6°23.2' E), infralittoral rocks, 0-1 m, 2 shs, MNHN réc. S. Gofas, Sep. 1992; Provence, Porquerolles plage Notre Dame, (43°00.6' N, 6°13.8' E), rocks, 0-1 m, 1 sh., MNHN réc. S. Gofas, Sep. 1992; Provence, Le Dramont, conchiferous sand, beach, 3 shs, MNHN réc. J. Pelorce, 1992: Provence, Marseille (Endoume, 43°16.9' N, 05°21.0' W), littoral rocks, 1 sh., MNHWN réc. S. Gofas Apr. 1995; Provence, St. Raphael, 5 shs, sata Provence, Bandol, 16 shs, MNHN coll. Locard: Raphael, 2 shs, MNHN coll, Locard; Corsica, Ajaccio, VW shs, MNHN coll. Jousseaume; Corsica, Algajole, 35 shs, coll. MF, 2123. Tunisia: Monastir-Khenis, beach, 1 sh., coll. Garilli, legit Garilli and Galletti 4/2000; ae beach, 1 sh., coll. MF ex coll. C. Bogi, Jun. 1981, 2163JER. Italy: Piemonte, Asti, Bi dichieri, “Grottino Monale”, 20 shs, yellow sands, middle-upper Pliocene, coll. MF ex coll. PAL, F1O4A:; Piemonte, Asti, 24 shs, Pliocene, MGUP coll. Doderlein, 113B; Piemonte, Alessandria, Tortona, | sh., Miocene, MGUP coll. Doderlein, 476B; Alessandria, Villalvernia, at the Cem- etery, 1 sh., Astian yellow sands, middle-upper Pliocene, coll. MF ex coll. PAL, FLISA: Emilia Romagna, Modena, Maranello, Fogliano, Gagliardella, Rio Grizzaga sands, 1 V. Garilli, 2008 icy) Ol Page « sh., middle Pliocene, coll. MF ex coll. PAL, F39A: Tus- cany, Siena, ne: ea Villa Pietratitta, Podere “La Vigna’, (SP 36, 4.9 km E side), 7 shs, lower Pliocene, coll. MF ex ot PAL, F55B; Siena, Poggibonsi, Villa Pietratitta, “Sbarra”, (SP 36, 5.2 km), 2 shs, sands, lower Pliocene, coll. MF ex coll. PAL, oo Siena, Castel- nuovo Berardenga, Terre Rosse, ( 104 km), 1 sh., sands, lower Pliocene, coll. oe ex call. PAL, F36A; Siena, Poggibonsi, Villa Pietrafitta, Podere Sant’ Uliviere, 27 shs, lower Pliocene, coll. MF ex coll. PAL, F55C: Siena, Poggibonsi, Villa Pietratitta, Podere Melograni, 6 shs, sands, lower Pliocene, coll. MF ex coll. PAL, F55E: Siena, Colle Val d’Elsa, Bibbiano, 2 shs, yellow sands, lower Pliocene, coll. MF ex coll. PAL, FLOSA; Tuseany, Livorno, Tuscan Archipelago, Island of Elba, Procchio, 12 m, Posidonia bed, 2 shs, coll. MF ex coll. C. Bogi, 2120PROC: Pai Terni, Ficulle, quarry near Chiani river, (SST1), 1 sh., Cidaris marly sands, lower Pliocene, coll. MF ex coll. PAL, F6SA; Sardinia, Nuoro, San Te- odoro, 6 shs coll. PAL; Puglia, Taranto, MSNCS 44744 (ex 1505), 1974, BDA legit and det; Sicily, Messina, Mi- lazzo, Capo Milazzo, Cala S. Antonio, 12 shs, upper yel- low sands, upper Pleistocene, coll. MF ex coll. PAL, FSA: Sicily, Catania, Grammichele, C.da Catallarga, 2 shs, coarse sands, lower Pleistocene, coll. MF ex coll. PAL, F27A; Sicily, Palermo, 1 sh., MNHN coll. Petit: Sicily, Palermo, 2 shs, MNHN coll. Dollfus, 1903; Palermo, Tommaso Natale, 13 shs, upper Pleistocene, MGUP 167/ 2/50; Palermo, Addaura, 1 sh., upper Pleistocene, MGUP 587/5/15; Sicily, Catania, 2 shs, MSNCS 44743 (ex 1448), 4/1974, G. Gentile legit and det.; Sicily, Sira- cusa, Vendicari, 1 sh., coll. PAL, 2124. Adriatic Sea: 128 shs, MNHN. Croatia: Zara, 8 shs, MNHN coll. Petit, 1873. Greece: Evvia Island (Euboea), Loukissia, 4-5 m, 1 sh., coll. PAL ex coll. Bogi Cesare, 2121LUC. Israel: Haifa, 9 m, 1 sh., coll. PAL ex coll. C. Bogi, 1994, 2164HA. Habitat: As indicated by Jeffreys (1867), Gofas and Ponder (1991). and Bouchet (1978: 1992), this species typically lives buried under stones covered with sand at very shallow waters. I found very fresh shells collected from along all the upper part of the infralittoral stage. Distribution: All the Mediterranean, probably de- creasing is a in eastward direction. In the eastern Atlantic it lives from the British Islands (see also Jeffreys, 1867: 1869) to Morocco. Alvania lactea has also been recorded from the Black Sea (Anistratenko and Star- obogatov, 1994). This species probably originated in the Mediterranean Neogene, from where it is recorded from the Miocene of north Italy. It becomes more common in the western and central Mediterranean Pliocene depos- its, where it is recorded from several localities of north, central. and south Italy (material herein studied: see also Sacco, 1895, and Chirli, 2006) and Spain (Estepona, Lan- dau et al., 2004). In the Atlantic Pleistocene, as reported by Landau et al. (2004), it is recorded from the post- glacial of Iceland. the North Sea Basin and the British Isles. As subfossil, it is recorded from Sweden (Huben- dick and Warén, 1969). In Mediterranean, it is common from the lower-upper Pleistocene of Sicily and is also recorded from the lower Pleistocene of Tusc: ny (Chirli, 2006). Remarks: This is a very variable species especially with respect to sculpture and shell shape. The typical morph, as described and figured by Michaud (1831: fig. 12), has a clathrate sculpture with the axial pattern domi- nating over the spiral one. The finely cancellated type, usually characterized by an elongate general shape and more convex whorls, corresponds to the forma minor- tenuisc ulpta Monterosato, 1917. Two varieties, semiacos- tata and fusulatovaricosa (the latter often bearing va- rices), were described by Sacco (1895) on similar mate- rial from the Pliocene of north Italy. This cancellate morph (Figures 44 and 47-48), ), which seems to be the only representation of the discussed species in the Mio- cene, is the most common in the Pliocene collections studied (see also Sacco, 1895: p. 28; Chirli, 2006: figs. 9-11 and 15-16) and becomes less common in Quater- nary material. The markedly ribbed morph, descsbed by Sacco (1895) as var. laticostata (from the Pliocene of Italy), and better known as forma dajerleini Monterosato, 1889, bears strong ribs, starting from the early teleo- conch whorls. According to Monterosato (1917) ), this re p- resents the Atlantic morph of Alvania lactea. 1 agree with Van Aatsen et al., considering it not exclusively an Atlan- tic morph, being present in some Mediterranean locali- ties (see Figure 45). It is noteworthy to remember that in a extensive lot (128 shells, MNHN-DSE) from the Adri- atic Sea, all the above mentioned morphs coexist. Especially when its typical morphs are considered, Al- vania lactea has a very characteristic ribbed, cylindrical- ovate shell and can not be confused with its congeners. Some problem might occur when considering the can- cellate morph, winch can be compared to ‘he recently described Pliocene species A. fredianii Della Bella and Scarponi, 2000 (see this article for the main differences between the two species). The same morph of A. lactea shows strong similarities with the new species described herein, A. rosariae (see discussion below for differential diagnosis). Alvania lactea is the type species of Massotia Bucquoy et al., 1884, which is considered a synonym of Alvania sensu stricto by Ponder (1985) on the basis of the num- ber of the metapodial tentacles, shown by this species, based on a description by Jeffreys (1867). In my opinion, a more exhaustive anatomical dataset could be provided. Nevertheless, even on the basis of its shell features and intraspecific variation, there is no reason to consider Massotia as a well-supported group. Alvania lactea has a very complicated teleoconch mi- ila ee which could ce srtainly be considered as a suite of distinctive characters, not observed in the closely-related taxa discussed in the present report. Early teleoconch whorls show a pattern similar to that ob- served in A. carinata, A. francescoi new species, and A, rosariae new species (detailed comparisons are provided Page 36 THE NAUTILUS, Vol. 122, No. 1 below), but the remainder of the shell is covered by very narrow, raised lamella-like axial ridges and very thin spi- ral threads. Also its protoconch I sculpture is rather dis- tinctive, showing a pattern similar to that observed in A. carinata, A. francescoi, A. rosariae, and A. tenera (as well as in many other rissoids), but having a coarser ornamen- tation. Alvania prusi (P. Fischer, 1877) (Figures 50-52) Rissoa (Alvania) Prusi P. Fischer, 1877: 80. Description: Shell small, Saaney conical, partially worn off, 4.7 mim in height, 3.2 mim in width. Protoconch multispiral, conical, consisting of about little more than two convex whorls. ‘Teleoeonel is formed by about four convex whorls, provided with a very delicate, sida ali sculpture almost giving a pitted shape to shell surface. This sculpture consists of moderately pronounced spiral cords crossed by apparently flat axial ribs. Spiral cords number 7-8 on penultimate whorl and 16 on body whorl, they are flatter and wider on shell base. Ribs, numbe ring about 40 on penultimate whorl, are lacking in basal part of last whorl. Sutures slightly inclined and deeply im- pressed. Last whorl well- expanded, rather angulated at the base and inflated, comprising % of shell height. Aperture wide, ovate to almost pyriforme, pointed adapi- cally, comprising 2 of the total height, 7% of last whorl height. Outer lip prosocline, internally smooth, externally with 1 an almost flat, wide thickening, apparently smoc th. Inner lip moderately arcuate, with a modest and narrow reflection on columellar area, leavi ing a very small and narrow umbilical chink (filled up with sediment). Type Locality: Isle of Rhodes Type Material: One syntype, RO7495 in MNHN- DHT, from type locality. Material Examined: Eastern Mediterranean Sea, Quaternary of Rhodes, type locality, MNHN-DHT (RO7495), 1 sh. Habitat: It is very difficult to characterize the paleo- ecological significance of this extinct species since no data about its paleoe nvironment are direc tly available or deducible. Furthermore, Fischer (1877) ) provide -d no precise indication about the site, stratotype, and its sedi- mentological and paleontological nature. Considering all the other molluscan species described from Rhodes by the same author, a shallow water depositional environ- ment (linked to the modern phanerogam infralittoral bottom) can be inferred. Distribution: The species is known only from the type locality, and its distribution appears to be limited to the Quaternary of Rhodes. Remarks: knowledge, the most recent treatment is that of Monte- This taxon is practically unknown. To my rosato (1917), who considered it as a separate species belonging to the Massotia group. Alvania prusi could be confused with juvenile shells of Alvania lactea (the form with cancellate sculpture, see Figure 49), which exhibit a very similar body whorl pro- file. Alvania prusi is quite close to A. fredianii Della Bella and Scarponi (2000) from the Pliocene of Tuscany. Both species share the general shell shape and the finely reticulated sculpture, tra the latter is of smaller size, has a paucispiral protoconch, deeper sutures, and a weakly denticulated outer lip (see Della Bella and Scarponi, 2000: pls. 1 and 2). Alvania rosariae new species (Figures 62-79) Description: Shell small, sturdy, conical-ovate to tur- riform, moderately to markedly inflated, reaching about 5 mm in height, 3.5 mm in width (holotype 4 mm in height and 2.7 mm in width). Protoconch multispiral, conical, consisting of about 2—2.1 convex whorls. Proto- conch I of about 0.8 whorls sculptured by 5-6 very fine spiral lirae irregularly interspersed with microscopic granules. Protocanch/teleaconch transition well marked aad sinuous adapically. Protoconch I sculptured by spi- rally arranged microscopic pimples (stronger in adapical portion) for ming one to two spiral threads, the lower very close to the suture, occurring on last whorl. Teleoconch formed by about 4.2 usually very convex w horls, sculp- tured by numerous axial ribs and slightly stronger spiral cords. The latter, rapidly increasing in count, number 2-6, 6-10 ( se 11), and 26-34 (in specimens higher than 3.5 mm) on the first, penultimate, and last whorl of the teleoconch, respectively. Some secondary, less marked cords may occur on last whorl, usually close to the outer lip. T The adapical one, two, and three spiral cords on first, penultimate and body whorl, respectiv ely are more pronounced. In specimens higher than 3.5 mm, axial sculpture consists of 44-60 narrow ribs (52 in ho- lotype) on the penultimate whorl. Ribs become narrower and lamella-like to obsolete or lacking on shell base, par- ticularly close to columella. The same may occur on the terminal portion of the body whorl (Figure 66). The in- tersection of spiral cords w ith axial ae gives a charac- teristic cancellate and gently nodular shape, forming a rectangular (almost equilate ral in the central portion of the body whorl) pattern. The long axis of these rectangles is perpendicular to the shell axis on the central and abapical portion of whorls and parallel on upper part. Microsculpture consists of ve ry fine, sometime irregu- larly interspersed, spiral lirae ( Figure 7 77). These become obsolete on the pe ia ite and iboay a where nu- merous, very fine and narrow ribs ( (possible growth lines) occur. Lirae are continuous only on ade ipical portion of early teleoconch whorls and never cover primary spiral sculpture. Sutures slightly inclined and rather deeply Be ssed. Last whorl well-developed, comprising about ; to % of the shell height, usually with a very convex iofile. It sometime bears one or two close varices which V. Garilli, 2008 may form angles of 10° to 210°. Aperture HS ovate, comprising 2 to % of the total height; %% ¥4 of last whorl height. Outer lip orthocline (slightly eee in- ternally smooth, externally markedly thickened close to lip edge and covered by spiral cords. Inner lip moder- ately arcuated and rather thickened in the columellar area, where a very narrow umbilical chink occurs. Type Locality: — Lower Pleistocene of Cartiera Mulino (36°56'57" N, 14°34'03" E), Vittoria, Ragusa, southeast- ern Sicily. The stratotype is the 3D1 layer of Costa (1989). Type Material: Holotype (4.0 x 2.7 mm), DGUP CMRG 12/496, coll. Garilli; Paratype 1 (4.6 x 3, 2 mm), ZMA Moll. 4.07.014 (ex CMRG 13/497 coll. Garilli): paratype 2 (not-complete shell, 3 mm width), ZMA Moll. 4.07.08 (ex CMRG 014/495 coll. Garilli); paratype 3 (3.6 x 2.55 mm), DGUP CMRG 15/499, coll. V. Garilli; paratype 4 (4.5 x 3.3 mm), DGUP CMRG 16/500, coll. Garilli: paratype 5 (3.8 x 2.5 mm), MNHN A25951 (ex CMRG 17/501, coll. Garilli); paratype 6 (3.85 x 2.75 mm), ZMB MB.Ga.2480 (ex CMRG 18/502, coll. Garilli); paratype 7 (3.6 x 2.4 mm), ZMB MB.Ga.2451 (ex CMRG 19/503, coll. Garilli); paratype 5 (3.8 x 2.6 mm), DGUP CMRG 20/504, coll. Garilli; paratype 9 (3.6 x ¢ 6 mm), DGUP CMRG 21/505, coll. Garilli; paratype 10 (3.95 x 2 2 6 mm), DGUP CMRG 22/506, coll. Garilli: ree 11 (4.2 «x 2.75 mm), DGUP CMRG 23/507, coll. Garilli; paratype 12 (3.7 x 2.55 mm), DGUP CMRG 24/508, coll. Garilli; paratype 13 (3.9 x 2.5 mm), DGUP CMRG 25/509, coll. Garilli; paratype 14 (3.7 « 2.55 mm), DGUP CMRG 26/510, coll. Garilli; paratype 15 (3.5 x 2.6 mm), DGUP CMRG 27/511, coll. Garilli; paratype 16 (4 x 2.6 mm), DGUP CMRG 28/512, coll. Garilli; paratype 17 (3.9 x 2.7 mm), DGUP CMRG 29/513, coll. Garilli; paratype 15 (3.6 x 2.45 mm), GNHM ID 30.706 (ex KIGR 3/514 coll. Garilli); paratype 19 (4 x 2.7 mm), GNHM ID 30.707 (ex KIGR 4/515 coll. Garilli); paratype 20 (4.5 x 2.7 mm), GNHM ID 30.708 (ex KIGR 35/516 coll. Garilli); paratype 21 (4.7 x 3.4 mm), GNHM ID 30.709 (ex KIGR 6/517 coll. Garilli); paratype 22 (4.4 x 29 mm), GNHM ID 30.710 (ex KIGR 7/518 coll. Garilli); paratype 23 (4 x 2.55 mm), GNHM ID 30.711 (ex KIGR 8/519 coll. Garilli); paratype 24 (3.35 x 2.5 mm), GNHM ID 30.712, (ex KIGR 9/520 coll. Garilli): paratype 25 (2.9 x 1.95 mm), GNHM ID 30.713 (ex ca 10/521 coll. Garilli); paratype 26 (not-complete shell ), GNHM ID 30.714 (ex KIGR 11/522 coll. Garilli): paratype 27 (4.15 x 2.65 mm), GNHM ID 30.715 (ex KIGR 12/523 coll. Garilli): ): paratype 28 (4.0 x 2.6 mm), GNHM ID 30.716 (ex KIGR 13/524 coll. Garilli); paratype 29 (not- complet e shell, 3.85 mm), GNHM ID 30.717 (ex KIGR 14/525 coll. Garilli); paratype 30 (2.7 x 1.9 mm), GNHM ID 30.718 (ex KIGR 15/526 coll. Garilli); paratype 31 (3 x 2.05 mm), GNHM ID 30.719 ex KIGR 16/527 coll. Garilli); paratype 32 (3.5 « 2.5 mm), GNHM ID 30.720 (ex KIGR 17/528 coll. Garilli): paratype 33 (3.55 x 2.5 mm), GNHM ID 30.721 (ex KIGR 18/529 coll. Garilli); paratype 34 (4.05 x 2.5 mm), GNHM ID _ 30.722 (ex KIGR 19/530 coll. Garilli); paratype 35 (4.95 « 3.25 mm), GNHM ID 30.723 (ex KIGR 20/531 coll. Garilli); paratype 36 (not ene: ), GNHM ID 30.724 (ex KIGR 21/532, coll. Garilli); paratype 37 (not measured), GNHM ID 30.725 (ex KIGR 22/533, coll. Garilli). Holotype and Paratypes 4-17, from the lower Pleistocene of Cartiera Mulino, 3D1 bed of Costa (1989), Vittoria, Ragusa, southeastern Sicily. Paratypes 1-3 from the same locality, 3D2 bed of Costa (1989). Paratypes 15-22 and 24-35 from the middle to upper Pleistocene of Kyllini, northwestern Pelopomnesus, N2 and H6 beds of Garilli et al. (2005a), respectively. Paratypes 36 and 37, same locality, from the lower to early middle Picistoesie P3 layer of Garilli (2005b) and from a late lower Pleistocene yellowish to reddish sandy layer about 50 m underlying the F14 bed of Garilli and Galletti (2007), respectiv ely, Another four, eee d paratypes are housed in DSTC (1 sh. from 3D1, 1 hi from 3D2 and : hae: from 3C bed of Costa, 1989, all from the lot n° 18, as Galeo- dina carinata (Da Costa). One more fore pene paratype (ex CMRG 030/534 coll. Garilli), from the type locality, 3D1 layer, is in coll. MF (Prato), Etymology: saria. The species is dedicated to my wife Ro- Material Examined: The type material from the lower Pleistocene of Cartiera Mulino, Vittoria, Ragusa, SE Sicily, 18 shs, and from the late lower (1 sh., froma reddish to yellowish sandy bed about 50 m underlying the Fl4 bed ie Garilli and Galletti, 2007), lower to middle (2 shs, layer P3 of Garilli et al, 2005b). and middle to upper Pleistocene (13 shs, layer H6: 4 shs, layer N2) of Kyllini, Elea, NW Peloponnesus, Greece. Habitat: In the Do locality, the species was mainly recovered from the layers 3D1 and 3D2 which were linked to the ecotone SVMC-HP (sensu Pérés and Pi- card, 1964) by Costa (1989). In the Kyllini sites, this species was found in cerithids-trochids-rissoids assem- blage linked to the present biocenosis HP, characterized by the phanerogam Posidonia oceanica (Linnaeus) De- lile, 1813 (See Garilli et al. (2005a): Garilli et al. (2005b), and Garilli and Galletti (2007) for more detailed infor- mation about the paleoecological characteristics of the cited Kyllini strata.) Distribution: The species has a lower to middle-upper Pleistocene stratigraphic range, presently limited to SE Sicily and NW Peloponnesus, being recorded from the lower Pleistocene of Sicily (type locality) and from the lower to upper Pleistocene deposits of Kyllini, Greece (NW Peloponnesus). Remarks: Alvania rosariae represents a sort of inter- mediate form between A. carinata and A. lactea (cancel- late form). Compared with the former taxon, it shows some similarities in the type of intraspecific variation, showing very inflated to quite elongate shells (compare THE NAUTILUS, Vol. 122, No. | V. Garilli, 2008 Page 39 Figures 68-69 Res the “form” ecarinata of A. carinata, Figures l, 4, in usually bearing varices, and in the ch eee erie o the ¢ rarly teleoconch. In addition, their respective larval shells do not show relevant differences. Alvania rosariae new spe cies can be distinguished by absence of a keeled shape, having subequal and more numerous spiral and axial sculptural elements. The can- cellate form of A. lactea is comparable with A. rosariae new species, having a very similar sculpture and occa- sionally a similar shell shape (only in the very ovate morph, e.g. Figures 44, 47 and 65-66). However, A. ro- sariae new species differs by having more convex whorls and a very different microsculptural pattern, with only irregular, fine, often inte rrupted spiral threads covering the early teleoconch whorls. Furthermore, the proto- conch I of A. lactea bears a coarser sculpture, consisting of more numeorus and larger pimples and much more elevated spiral lirae. Alvania rosariae new species can also be compared with A. francescoi new species: both species have a ga- leodiniform shape, a similar microsculpture, and share a very similar sculptural pattern on the protoconch. Dif- ferences between the two species are mainly found in the teleoconch sculpture, which is strongly nodular and coarser in A. francescoi new species i a eee the latter has fewer spiral cords and axial fice. Alvania rosariae new species may resemble A. magis- tra Chirli, 2006, an interesting galeodiniform species from the Pliocene of N Italy and § Spain (Chirli, 2006: pl. . figs.13-16 and pl. 12, figs, 1- 8; Landau et al., 2004: pl. figs.3 B3a-e, as Alvania sp.), showing a quite similar ae cancellate sculpture, especially in the arrangement of the ad: ipical cords, and a quite wide, ovate aperture. However, the latter species shows an unusual outer lip profile, having a wide sinus at the upper aspect, its pro- toconch I shows a netted Manzonia-like sculpture, while protoconch II appears less sculptured than in A. rosariae new species Alvania tenera (Philippi, 1544) (Figures SO-S9) Rissoa tenera Philippi, 1544: 128-129, pl. 23, fig. 15. Galeodina tenera (Philippi, 1544) Pk ini, 1979: 71, fig. 4 Description: Shell minute, conical, and subcarinate, to conical-ovate or elongate, moderately to markedly sturdy, reaching 2.2 mm in height and 1.4 mm in width. Protoconch multispiral, conical: with 2-2.3 convex whorls. Protoconch I consists of about 0.8 whorl, sculp- tured by six very thin spiral lirae and a few microscopic or anules between them. Protoconch I is sculptured by a few to abundant very small granules. These are more numerous on last half whorl, where they are spirally ar- ranged, forming very discontinuous and. irregular ridges. Fiotoconely te Tesconel transition clistinct, eh a slight (to very slight) sinuosity. Teleoconch consists of 3-3.5 weakly to discretely convex whorls, quite variable in width. These are sculptured by well-raised, narrow, spi- ral, occasionally almost keel- like cords (in quite conical shells), which overide the axial sculpture. They number 34, me 5, and 12- - on first, penultimate, and body whorl, respectively. Usually, weaker cords are present on shell base or close to upper suture. Axial sculpture formed by very narrow, prosocline, rows of short seg- ments, forming discontinous ribs, occurring between spi- ral cords and becoming very thin to lacking toward shell base. Ribs number 40-60 on last whorl. At the intersec- tions with axial elements, spiral cords generally appear very finely nodular (Figure 86). Last whorl well- expanded, with a rather rounded profile, sometimes in- flated, comprising about 73 to ‘4 (in subcarinate and conical shells) of total shell height. Aperture ovate, an- teriorly rounded, posteriorly angulated, comprising about 2 and % of total shell and last whorl height, re- spectively. Outer lip rounded, markedly prosocline, thin, internally smooth, externally with no thickening. Inner lip weakly arcuate, with a thin and narrow (Fi igure 82) to stronger and wider (Figure 80) callus, leaving a very nar- row senabilical chink. ¢ Solokilion usually consists of red- dish to brown spots on a cream-whitish or (rarely) brownish background, ~ Type Locality: Peninsula of Magnisi (originally indi- cated as “Peninsula Thapsum” from the old Greek name of Magnisi), Siracusa, southeastern Sicily. Material Examined: Atlantic: Canary Islands, Tene- rife, Pal-Mar, 6—S m, 1 sh., MNHN coll. P. Bouchet, 15 Jul. 1980; Morocco: Asilah, mouth of Oued el Helou, conchiferous detritus, beach, 9 shs, MNHWN coll. S. Go- fas, 1971-72; El Jadida, (33°16" N, 08°29’ W), large beach, conchiferous detritus, beach, 5 shs, - NHN réc S. Gofas, 26 Sep. 1991; El Jadida, ( (33°16’ N, 08°29’ W), large beach, tide zone, 4 shs, MNHN réc S. Gofas, 26 Sep.1991. Strait of Gibraltar: Spain, Cadiz, conchiferous detritus, beach, 2 shs, MNHWN coll. S. Gofas, 1976-81; Cadiz, Tarifa, beach, 4 shs, coll. PAL ex coll. C. Bogi, Jun. 1986, 2030TAR; Cadiz, Tarifa, Torre de la Pena, conchiferous detritus, beach, 3 shs, MNHWN coll. S. Go- fas, Aug. L981; Morocco, Tanger, Grande Plage, conchif- erous detritus, beach, | sh., MNHWN coll. S$. Gofas, L970— S81. Mediterranean: Spain, Andalusia, Punta della Mona, 43 m, 5 shs, coll. PAL ex coll. C. Bogi, 2032PMO; Spain, Malaga, industrial dredging, 20-40 m, 1 sh., MNHN réc. Figures 62-69. Alvania rosariae new species, holotype and paratypes 16, 21, and 35, 62-63. Apertural (62), dorsal (63) and profile 64) views of holotype, lower Pleistocene of Cartiera Mulino, bed 3D1 of Costa (1959), SE Sicily, Ragusa, Vittoria. 65-66, Paratype 16, dorsal (65), illustrating varices on the last whorl, and apertural (66) view, same site and loc: lity. 67. Paratype 21, a very inflated last whorl and a strong varix opposite the outer lip, middle to upper Pleistocene of Kyllini, NW Pe loponnesus, N2 bed of ‘Garilli et al. (2005a). 68-69. Apertural (68) and dorsal (69) view of paratype 35, showing a quite slender shell, middle to upper Pleistocene of Kyllini, northwestern Peloponnesus, H6 bed of Garilli et al. (2005a). Scale bars 100 pm THE NAUTILUS, Vol. 122, No. 1 V. Garilli, 2008 Page 4] S. et C. Gofas, May 1991; Malaga, Calahonda, conchif- erous detritus, beach, 1 sh., MNHN coll.S. Gofas, 1976— Sl; Malaga, Benalmadena, conchiferous detritus, beach, 6 shs, MNHN réc S. Gofas, 1991-93; Malaga, Mijas, detritus, LO m, 4 shs, ee Stefano Rutini; Mz ilaga, Cabo Pino, detritus, 10 m, 2 shs, coll. SR, (41.S80¢); Morocco, Cabo Negro, beach, 1 sh., MNHN coll. S. Gofas, det. W. Ponder, 1986; France, Provence, Marseille, Curry, beach, 3 m, 2 shs, coll. PAL ex coll. C. Bogi, Jun. 1986, 2037 MAR; Provence, Marseille, La Baule, small beach at 25 km west from Marseille, 9 shs, coll. PAL ex coll. C. Bogi, Oct. 1986, 2035BAU; Provence, Le Dramont, (43°24.7' N, 6°51.7 E), 22-30 m, 26 shs, MNHN réc. J. Pelorce, 1992; Provence, Les Embiez, cote Nord et Petit Rouveau, (43°05' N, 5°47’ E), rocks, algae, 0-1 m, 11 shs, MNHN réc S. Gofas, Jun. 1995; Provence, Marseille, Cap Morgiou, “calque de la Triperie,” (43°12.2' N, 05°26.9' E), muddy sand , inside cavity, 22 m, 4 shs, MNHN réc H. Zibrowius, Jun. 1996; Provence, Marseille, Grand Congloue, (43°10.6' N, 05°24.2' E), 33 m, 50 shs, MNHN réc. H. Zibrowius, Jun. 1996; Provence, Les Embiez, passe du Gaou, rocks, photophile algae, (43°04.3' N 5°47.4' E), 0-3 m, 1 sh., MNHN réc S. Gofas, Jun. ae Provence, St. Clair, infralittoral rocks, (43°08.2’ N 6°23.2' E), 0-1 m, 1 sh., MNHN réc. S. Gofas, Sep. 1992: Tunisia, Sfax, 4 shs, MNHWN coll. Staadt, 1969; Italy, Liguria, Portofino, 1 sh., coll. PAL, 2038: Italy, Tuscany, Livorno, Antignano, 0.5 m, brown algae on rocks, 1 sh., coll. PAL legit Bogi, Apr. 1999, 2029: Livorno, under littoral rocks, 0.5-1.0 m, 5 shs, coll. PAL ex coll. C. Bogi, 2039LIV; Livorno, Meloria, 10-30 m, 3 shs, coll. PAL ex coll. C. Bogi, 1995, 2031MEL; Livorno, Tuscan Archipelago, Island of Elba, Capoliveri, 32 m, 2 shs, coll. PAL ex coll. C. Bogi, Aug. 1994, 2036CAPOL; Tuscan Archipelago, Gemini Island, (southern side of Island of Elba), 11 m, 3 shs, coll. PAL, 2034; Tuscan Archipelago, Isola del Giglio, Punta Fenaia, 32 m, Ish., coll. medshells.com ex coll. G. Rug- gieri; Tuscany, Grosseto, Argentario, 25 m, 1 sh., coll. medshells.com ex coll. G. Ruggieri, Jul. 1988; Italy, Lazio, Ostia, Tor Paterno, 33 m, 5 shs, coll. medshells. com ex coll. G. Ruggieri; Lazio, Roma, Santa Marinella, ex reti, 2 shs, coll. medshells.com ex coll. G. Ruggieri; Italy, Sardinia, Oristano, Santa Caterina di Pittinurri, 5 m, 3 ahe. coll. PAL ex coll. C. Bogi, 30 Aug. 1986, 2040SCP: Sardinia, Sassari, Island of Maddalena, beach, 3 shs coll. PAL ex coll. C. Bogi, 2028IMA; Sardinia, Nuoro, Sant’Antioco, Cala Lunga, 20 shs, coll. med- shells.com ex coll. G. Ruggieri, 03.1989; Italy, Cam- pania, Peninsula of Sorrento, Punta Penna, 2 shs, coll. medshells.com ex coll. G. Ruggieri, 1955; Italy, Sicily, Palermo, 2 shs, MNHN coll. Locard; Sicily, 1 sh., MNHN coll. Petit, 1873; Palermo, 3 shs, ZMB ex coll. Monterosato, $1013 (originally labelled by Monterosato as Cingula tenera, 3, 1125, Palermo); Palermo, Arenella, 2 shs, ZMB ex coll. Monterosato, $1014, (originally la- belled by Monterosato as “Cingula ie var... 5 2, Arenella, Palermo”); Sicily, Siracusa, Capo Passero, 16 m, 3 shs, coll. medshells.com ex coll. G. Ruggieri, 05 Sep. 1987. Habitat: This species is clearly limited to infralittoral depths. In the upper part of its distribution, it seems to live in very shallow waters, on algae. It likely lives also in the cavities occurring in infralittoral muddy sandy bot- toms. Distribution: In the western and central Mediterra- nean the species seems to be well distributed; Adriatic and eastern Mediterranean occurrences should be veri- fied. In the Atlantic it lives along the Moroccan coasts and in the Canary Islands. To my laow ledge, there is no fossil record of this species. Remarks: — This small species is characterized by having a variable shell shape and sculpture. The conical shells, bearing a strongly cingulated sculpture, which markedly prevails over the axial, are comparable with typical keeled morph of Alvania carinata. In addition, Cingula species provided with a strong spiral sculpture are com- parable to A. tenera. The ovate, slender shells of this last species, with a finely cingulated sculptural pattern, might vaguely resemble some species of Setia H. and A. Adanis. 1854. Asa consequence, Piani (1979) ) and Van Aartsen (1982) included A. tenera in Galeodina, whereas Nords- ieck (1968; 1972) placed the same species in Setia and Cingula elas ely. The last two views should not be accepted. In fact, the species of Setia have a smooth or very slightly sculptured shell with more convex whorls than A. tenera, and Cingula species are characterized by shells usually lacking axial sculpture and having a very thick outer lip. The shells from eastern Atlantic (Canary Islands and Morocco) do not show meaningful differences from the Mediterranean ones studied. With regard to the resem- blance between A. tenera and A. carinata, it is manifest in the keeled, conical-inflated shell shape shown by their typical respective morphs, and in having a quite wide and developed aperture. However A. fenera never shows va- Figures 70-79. Alvania rosariae new species 70. Protoconch of the holotype, lower Pleistocene of Cartiera Mulino, bed 3D1 of Costa (1989). SE Sicily, Ragusa, Vittoria. 71. Protoconch of p paratype 16, showing variation of the abapical ridges on protoconch II, same locality and bed. 72. Dorsal view of protoconch of paratype 21, showing protoconch I sculpture and the abapical ridges on protoconch TI, middle to upper Pleistocene of Kyllini, NW Peloponnesus, N2 bed of Garilli et al. (2005a). 73-74. Holotype, detail of early protoconch (73) showing sculpture of protoconch [and protoconch I/protoconch H boundary, and dorsal view of protoconch 74). 75. Sculpture of protoconch I, paratype 21. 76-79. Holotype, detail of teleoconch sculpture: early whorls (76), first to goes whorl (77), showing microsculpture, penultimate to last whorl (78), showing the microscopic incremental scars, and last whorl ( Scale bars: 100 2m in Figures 70-72, 74, 76 and 78-79: 50 jzm in Figure 77; 20 ym in Figure 75. Black and white arrows ened the protoconch I/protoconch IH and protoconch/teleoconch boundaries, respectively NNN eee THE NAUTILUS, Vol. 122, No. 1 V. Garilli, 2008 Page 43 rices, bears a very delicate axial sculpture, its outer lip is always very thin, and usually has more colored shells. Genus Galeodinopsis Sacco, 1895 Type Species: nal designation. Rissoa tiberiana Coppi, 1876 by origi- Galeodinopsis tiberiana (Coppi, 1876) (Figures S1-99) 1862 Rissoa tuba Doderlein, 1862: 17 (nomen nudum) 1876 Rissoa Tiberiana Coppi, 1876: 201-202. Manzonia fariai Rolain and Fermandes, 1990: 64-65, pl. 1, figs. 4-6. Alvania fariai (Rolin and Fernandes, 1990).—Gofas, 1999: 8S— 89, figs. 39-42. Alvania fariae (Rolain and Femandes, 1990)—Landau et al., 2004: 41, pl. 7, figs. 3-4. Q Description: Shell conical, sturdy, reaching 3.75 mm in height and 2.75 mm in width. Protoconch multispiral, conical, with 2-2.2 convex whorls and a rather immersed nucleus. Protoconch I consists of about 0.7—0.8 whorls, with a netted sculpture, consisting of 7-8 very thin spiral lirae and numerous, irregular, short and very narrow axial segments occurring in interspaces between lirae. Proto- conch II is sculptured by very small, sparse granules, fused into 2-4 discontinuous ridges on central wad abapi- cal portions of latter part of fast. whorl. Groups of gran- ules form very short, prosocline segments on central area, mainly close to protoconchyteleoconch transition, which is marked by a quite sinous and thin lip. Teleo- conch formed by 3-3.5 convex whorls, sculptured by a primary pattern ‘of strong, slightly sinuous and opistho- cline ribs, numbering 12-14 on pemdinee whorl, be- coming very weak to lacking toward shell base. Ribs are covered by a secondary spiral sculpture, formed by flat narrower cords, numbering 4-5 on penultimate whorl, becoming more marked on shell base. Each cord bears a pitted microsculpture, consisting of microscopic subcir- cular pores forming S—10 spiral alignments (Figures 99). Between cords, numerous, closely spaced, very fine spi- ral ridges appear. They are for ined by rough ae ae tooth- like elements, extending perpendicularly rom shell surface. At their base, ridges have a lamella-like expansion which covers the interspaces occurring be- tween them. Last whorl inflated, well-expanded, on prising about % of total shell height, often bearing 1-2 strong varices, most frequently just before the outer lip Aperture ovate, very rounded anteriorly, weakly angu- lated posteriorly, comprising about little less than ani 3 of total shell and last whorl height, respectively. Outer lip sinuous, weakly opisthocline; internally smooth, with a thin rim on its edge; externally with a very marked varicose swelling, covered by spiral sculpture, and having a narrow ridge on its base, toward aperture, so that it appears double -rimmed. Inner lip weakly curved, with a very narrow columellar thicknening, forming an obsolete umbilical chink. Shell color white in Recent material. Type Locality: Coppi (1876) cited “La Tagliata”, an unknown name in toponymy (very likely referring toa recently deforested woodland) corresponding to the lo- cality Gagliarde lla (Maranello, Modena, Emilia Roma- ona, North Ite uly) (S. Palazzi personal comm., 2006). Type Material: A lot (PUM 13721), from Coppi coll, ee locality, with more than 100 possible syntypes (not seen), is housed in the MPOB, Modena. Material Examined: Senegal: Region de Dakar, 250 m S.W. Cap Manuel, 12 shs, MNHWN coll. Marche- Marchad, dét. S. Gofas [The Nautilus 113: 8S—89, figs. 40, 42]; Region de Dakar, S.W. Gorée large Cap Vert, 250-150 m, ae MNHN coll. Marche-Marchad, det. S. Gofas [The Nautilus 113: SS—89, figs. 40, 42]; Region de Dakar, St. 56-1-10A Gorée 150-200 m, 1 sh., MNHN coll. Marche-Marchad, det. $. Gofas [The Nautilus 113: 88-89, figs. 40, 42]; Angola: Luanda, Hha de Luanda, Cirealittoral, 120 m, 3 ae MNHN coll. S. Gofas, det. S. Gofas [The Nautilus 113(3): 88—S9, figs. 40, 42]; Luanda, Ilha de Luanda, circalittoral, 40-60 m, 1 sh., MNHN coll. S. Gofas ares det. S. Gofas [The Nautilus 113: 88-59, figs. 40, 42]; Luanda, Au large de Mussulo (Mocéco), ” dredging 50-70 m, 2 shs, MNHN coll. S. Go- fas L9S1—1987, det S. Gofas (‘The Nautilus 113: 8S—S9, figs. 40, 42]; Au large de Mussulo, circalittoral, 90—LOO m, 2 shs, MNHN coll, S. Gofas, det. S. Gofas |The Nau- tilus 113: 88-89, figs. 40, 42]; Ambrizete, dredging, SO m, 5 shs, MNHN coll. S. Gofas, det. S. Gofas [The N Nautilus 113: SS—89, figs. 40, 42]; Ambrizete, (07°00' S, 12°20' E) sediment, 60 m, 3 shs, MNHWN coll. S. Gofas, 1983, det. S. Gofas [The N Nautilus 113: 88—S9, figs. 40, 42); Am- brizete, (06°57' S, 12°23’ E), sediment, 45 m, 1 sh., MNHN coll.S. Gofas, 1983, det. S. Gofas [The Nautilus 113: SS—89, figs. 40, 42). Italy: Piemonte, Asti, Baldich- ieri, “Grottino Monale”, 2 shs, yellow sands, middle- upper Pliocene, coll. MF ex coll. PAL, FLO4A; Asti prov- ince, 9 shs, Pliocene, MGUP coll. Doderlein, 111A; Lig- uria, Savona, Rio Torsero, between Ceriale and Peagna, 5 shs, clays, lower Pliocene, coll. MF ex coll. PAL, F5SA; Figures 0-89. Alvania tenera (Philippi, 1844). 80. Shell of a keeled and conical morph, Strait of Gibraltar, Spain, Cadiz, Tarifa, coll. PAL (2030TAR). 81-82. Conical-ovate, not-keeled morphs, illustrating the variable sculpture, France, Provence, Marseille Curry, coll. PAL (2037MAR). 83. Profile view of the same shell as Figure 80. 84. Profile view of an unkeeled morph, France Provence, Marseille, La Baule, small beach at 25 km west from Marseille, coll. PAL (2035BAU). 85-86. Sculpture of early teleoconch whorls (85) and detail of sculpture (86), Italy, Tuscany, Livorno, Antignano, coll. PAL (2029). 87-88. Protoconch (87) and detail of protoconch I ($8) of the same shell. 89. Protoconch of the same shell as Figure 84, showing sculpture variation. Scale bars: 0.5 mm in Figures 80-84: 200 jzm in Figure $5; 100 jum in Figures 86-87 and 89: 20 pm in Figure 88. Black and white arrows indicate the protoconch I/protoconch II and protoconch/teleoconch boundaries, respectively. THE NAUTILUS, Vol. 122, No. 1 V. Garilli, 2008 Page 45 Emilia Romagna, Modena province, 35 shs, Pliocene, coll. Doderlein, 111B; Emilia Romagna, Modena prov- ince, 1 sh., Miocene, MGUP coll. Doderlein, 474: Emilia Romagna, Piacenza, Lugagnano Val d’Arda, 2 shs, “calan- chi di valle” (marls), middle-upper Pliocene, coll. MF ex coll, PAL, FI3A; Piacenza, Castell’Arquato, Monte Pa- dova, 1 sh., blue clays, middle Pliocene, coll. MF ex coll. PAL, FIGA; Emilia Romagna, Parma, San Nicomede, Stirone river, 50 shs and fragments, lower clays middle- upper Pliocene, coll. MF ex coll. PAL, FI4B: Emilia Romagna, Modena, Marano on the Panaro, Panaro river, 1 sh., clays stormy layers, middle-upper Pliocene, coll. Me ex coll. PAL, FSOA: Emilia Romagna, Modena, Ma- ranello, Fogliano, Gagliardella (type locality), Rio Griz- zaga, 60 shs, sands, middle Pliocene, coll. MF ex coll. PAL, FS9A; Tuscany, Siena, San Donato, Ciuciano, Prison, | sh., clays and sands, lower Pliocene, coll. MF ex coll. PAL, F112A; Siena, Castiglioncello del Trinoro, Poggio Rotondo, 3 shs, marls, lower Pliocene, coll. MF ex coll. PAL, F54A; Lazio, Rome, Magliano Sabina, Cla- docora yellow sands, 23 shs, lower Pliocene, coll. MF ex coll. PAL, FI5A: Sicily, Palermo, Altavilla Milicia, rigth side of Milicia river, 12 shs, sands, lower-middle Pliocene, coll. MF ex coll. PAL, F2A; Palermo, Partitico, Trappeto, Lido Ciammarito to Nocella river mouth, 11 shs, clays, lower Pliocene, coll. MF ex coll. PAL, F72A. Habitat: In the Atlantic Ocean, the species has a lower shelf-upper slope distribution (see Gofas, 1999), is indi- cated by the fossil Mediterranean occurrences. A shal- lower and more restricted distribution, limited to shelf paleoenvironments, with sandy to muddy bottoms. Distribution: The species lives in the eastern Atlantic, from Senegal to northern Angola (see also Gofas, 1999). It was also collected from the coasts of Mauritania (S. Palazzi, pers. comm., 2006). The species has a Mediter- ranean Miocene to Pliocene paleoditribution, being re- corded from the Miocene of northern Apennines (Modena), the Pliocene of northern (Piemonte, Liguria, Toscana, Emilia Romagna), central (Lazio) and insular (northwestern Sicily) Italy, south Spain (Estepona) Lan- dau et al. (2004, as Alvania fariae), and Algeria (Coss- mann, 1921). In Atlantic, it is recorded fom the Portu- a middle Pliocene of Mondego Basin (Landau et al., 2004). The citation of Wenz (1938: 616), according to Ww hich the species lived in the Oligocene (up to Fieesae of Europe, North Africa and North America), should be verified. The species was not found in the very rich mol- luscan assemblages from the Oligo-Miocene of south- western France (Lozouet, 1998; 1999). Remarks: —Gofas (1999) and Landau et al. (2004), dis- cussing and re-describing this species, originally de- scribed as Manzonia fariai by Rolan and Fermandes (1990) from West Africa, did not recognize its identity as Rissoa tiberiana Coppi, 1876, the latter being a common species from the Mediterranean Neogene, particularly from the Pliocene. This species, origin ally not illustrated by Coppi (1576), was figured by Sacco (1895: figs. 67, a-bis and 68, a—b), who de signated it as the type species of the subgenus Gale odinopsis Sacco, 1895. More re- cently, this species was illustrated by Cossmann (1921: pl. 1, figs. 55-56) and Wenz (1938: fig. 1715). Compari- son between fossil material of R. tiberiana (Figures 90— 94, also from topotype material; see also Landau et al., (2004: pl. 7, figs. 3-4), to that of Rolain and Fernandes oun. aL , figs. 4-6), and of Gofas (1999: figs. 39-42) strongly éonficus the above mentioned synonymy. The rather conical, ribbed shell with an inflated last whorl, the frequent presence of varices on the last whorl, and the double-rimmed outer lip are the most characteristic features of this species, which shows a modest variability in the number and strength of ribs and in the spire el- evation (see Figures 90-92). Gofas (1999) moved this species from Manzonia Bru- sina, 1870 [type species Manzonia crassa (Kanmacher, 1798), see Figures 104-107] to Alvania based on the lack of the characteristic punctate spiral sculpture of the former taxon. This does not appear appropriate. In fact, in reasonably well-preserved shells, the primary spiral cords clearly bears a microsculpture consisting of regu- lar, spirally arranged pits, quite like M. crassa (compare Figure 99 with Figure 106). This spiral pitted micro- sculpture, sonsilared a typical Manzonia character by Moolenbeek and Faber (1987), was indicated by Bouchet and Warén (1993) as not restricted to this genus (occurring in Alvania, Gofasia Bouchet and Warén, 1993, and with a rough similarity, in Rissoininae species, see Gofas, 1999, figs. 79-80, 85, and 89]). The same authors interpreted it as a symplesiomorphy retained in Manzonia and in other rissoid genera. However , the par- ticular structure of the secondary, very fing spiral threads, formed by roughly prismatic elements growing perpendicularly to the Shell surface, is a chowacker share d by the Manzonia species, never observed in Alvania, and retained only in the recently described genus Gofasia (see Bouchet and Warén, 1993, fig. 1557). The combi- Figures 90-99. Galeodinopsis tiberiana (Coppi, 1876). 90-93. Shells from the type locality, showing variability and varices, middle Pliocene of Italy, Emilia Romagna, Modena, Maranello, Fogliano, Gagliardella, Rio Grizzaga sands, coll. MF ex coll. PAL (F394). 94. Profile view of a shell from the middle- ‘upper Pliocene of Italy, Emilia Romagna, Parma, San Nicomede, Stirone River, coll. MF ex coll. PAL (F14B). 95-96. Protoconchs from the same locality, note variation of the abapical sculpture just behind the transiction to teleoconch. 97. Detail of protoconch I, showing the nette d microsculpture and the partially immersed nucleus, same shell as Figure 95. 98. Detail of teleoconch sculpture from the shell as Figure 90. 99. Detail of tele oconch microsculpture from the shell as Figures 95-96: note the pitted pattern on the spiral cord and the structure of the fine spiral ridges. Scale bars: | mm in Figures 90-94; 100 pm in Figures 95-96; 50 jm in Figure 98; 20 pm in Figures 97, 99. Black and white arrows indicate the protoconch I/protoconch I and protoconch/teleoconch boundaries, respectively. er ee a ee AUTILUS, Vol. 122, No. 1 V. Garilli, 2008 Page 47 nation of these characters (pitted sculpture and structure of fine spiral threads) represents a quite singular and original feature, which should be regarded as limited to Manzonia-related species.: e.g. M. darwini Moolenbeek and Faber, 1987 (pl. 1, fig. 18), M. crispa CW atson, 1873) (see Moolenbeek and Faber, 1987, pl. 3, fig. 54), M. boogi lanzarottii Moolenbeek and Faber, 1987 | pl 2, fig. 39), M. spreta (Watson, 1873) (see Moolenbeek and Faber, 1987, pl. 3, fig. 57), M. vigoensis (Roldan, 1983) (see Bouchet and Warén, 1993, p. 656, fig. 1499) and several others. The double rimmed outer lip and the netted sculpture of protoconch I are also characters shared by Manzonia species. However, it must be con- sidered that these two features, considered by Ponder (1985: 46) as typical of Manzonia sensu stricto, should be a aled with suspicion, being as they are shared by well- recognized Alvania species. In regard to the proto- conch, species such as A. testae (Aradas and Maggiore, 1844), A. setlandica (Montagu, 1815) (see Bouchet and Warén, 1993, figs. 1386-1387 and 1502; Landau et al., 2004, pl. 9, figs. 1b-Id), the Pliocene Alvania magistra Chirli, *006 (pl 11, fig. 16 and pl. 12, figs. 1-3), have the same sculptural pattern on protoconch I. A. tomentosa Pallary, 1920), which has a paucispiral protoconch, also hie this sculpture (see Bouchet and Warén, 1993, fig. 1388). Rissoa tiberiana could be regarded as one (probably the sole) of the few survivors of a group of species close to Manzonia sensu lato, which very likely originated in the upper Paleogene. The European Oligocene Rissoa duboisi Nyst, 1843 (Figures 100-103) certainly belongs to this group. Both these species share several intersting characters: a quite conical Alvania-like shell shape, often provided with varices on the last whorl, the arrangement of the basal cords (not so strong as in Manzonia sensu stricto, where keels occur on the shell base), the above mentioned combination of the microsculptural pattern, and the kind of axial sculpture (with slightly sinuous, less pronounced ribs than those shown by Manzonia), and a double, weakly opisthocline outer lip. All this leads me to revalue Galeodinopsis as the useful generic placement for such Manzonia-related species. In overall appearance, G. duboisi (Nyst, 143) strongly resambles G. tiberiana, from which it differs princips ally in having less numerous cords and ribs and less convex whorls (see also Ponder, 1985, fig. 100c). Another com- parable species is the Recent Macaronesian M. spreta (Watson, 1873), which has a similar shell shape but dif- fers from G. tiberiana in having a more delicate axial sculpture (the spiral cords being large and almost flat) a more rounded and smaller aperture, and a paucispiral protoconch (see Moolenbeek and Faber, 1987, figs. 47a—b and 55-57). With some significant reservations, it could be regarded as belonging to Galeodinopsis. The species M. foraminata (Lozouet, 1998), originally de- scribed as Alvania (from the upper Oligocene : south- western France, see Lozouet, 1998, fig, 9f-h), M. mou- linsi (dW Orbigny, 1852) (see Lozouet, 1998S, fig. a from the French upper Oligocene), M. scalaris (Dubois, 1831) (Kowalke and Harzhauser, 2004, fig. Sd, from the middle Miocene, Badenian, of Austria, Hungary, Poland and Ro- mania and the Miocene of Russia) and the Recent Man- zonia crispa (Watson, 1875) (of which I studied 3 shells from Madeira, ZMA Moll. 101.0, ex coll. R.G. Moolen- beek; see also Ponder, 1985, fig. LOOA and Moolenbeek and Faber, 1987, text-fig. 46, acl pl. 3, fig. 52-54) show less affinities, having a more slender chell with more curved ribs (protruding over the suture in M. scalaris), which become stronger on the base of the shell. All these four species appear more related to Manzonia than to Galeodinopsis. Sacco (1895) and Cossmann (1921) indicated Rissoa multicostata Speyer 1864 (pl. 41, figs. 3-5, from the Oli- gocene of Germany) as a possible Galeodinopsis species. I did not see any shells of this Alvinia sensu lato-like species, which more closely resembles the group of Al- vania zetlandica (Montagu, 1815) and A. weinkauffi (Weinkauff, 1S68 ex Sdhwartz ms.). The original illustra- tions show a turreted shell with a more finely cancellate (not-ribbed) sculpture, bearing almost orthocline axial ribs, characters which militate against placement in Ga- leodinopsis. The material of Rissoa tuba Doderlein, 1862, housed in the MGUP Doderlein’s collection (from the Miocene and Pliocene of North Italy), belongs to this species. Anyway, Doderlein (1562: 17) just listed this taxon with- out providing a description or a valid reference, so that R. tuba must he considered a nomen nudum. CONCLUDING REMARKS As indicated by Ponder (1985), the systematic grouping of Alvania species at the subgeneric level is quite diffi- cult and putative groups usually fold into synonymy with Alvania sensu stricto. This viewpoint appears be appli- cable to the species studied in this report (except for Rissoa tiberiana Coppi, 1876). The subgeneric division into Galeodina and Massotia lacks any valid and convinc- ing basis in shell features (especially on consideration of Figures 100-107. Galeodinopsis duboisi (Nyst, 1843) (100-103), from the upper Oligocene of Hessen (Germany, SE of Kassel, Hiessizoh: Lichten, Glimmerode, coll. MF ex coll. Pal, FEI5A) and Manzonia crassa (Kanmacher, 1798) (104-107), type species of Manzonia Brusina, 1870, from the middle-upper Pleistocene of Kyllini (NW Peloponnesus, Greece, N2 bed of Garilli et al., 2005a). 100-102. Apertural (100 and 102) and profile (101) views. 103- 104. Microsculptures, showing the typical pitted surface on the flat cords and the microstructure of the narrow spiral threads in G. duboisi (103, same shell as Figure 101) and M. crassa (104, same shell as Figure 105). 105-106. Apertural (105) and profile (106) view of shell. 107. Detail of protoconch I, showing the netted sculpture Scale bars: 1 mm in Figures 97-98; 20 jm in Figure 100: 10 zm in Figure 99. The black arrow indicates the protoconch I/protoconch Il boundary. Page 45 THE NAUTILUS, Vol. 122, No. 1 the often extensive intraspecific variation). In effect, a link between the markedly keeled and inflated shells (e.g. A. carinata), and the more typical Alvania-shape taxa could be hypothesized. It could be articulated and summarized in the transition A. carinata-A. lactea (via A. francescoi new species-A. rosariae new species) to more typical Alvania secies. A parallel trend could be con- structed: A. tenera and A. cingulata, showing a progres- sive weakening of the spiral sculpture, the proportional reduction in the expansion of the body whorl and the formation of the outer lip thickening. Furthermore, other interspecific trends, involving various nominal groups of Alvania, could be constructed when consider- ing the widely variable shells of most of the species dis- cussed here. Nowstanding this, I cannot deny that all these hypothetical links appears tenuous and that these taxa, sharing a quite inflated and well-expanded body whorls, a wide, internally smooth aperture, seem some- what distant from Alvania sensu stricto This point of view could lead to regard them as belonging to an inclusive taxonomic group (preferably at subgeneric level). Galeo- dina could serve as the appropriate taxonomic unit. Nev- ertheless, since I believe that appropriate anatomical studies (of which there is no exhaustive data-set) should be used to solve this question, I have preferred a more open-ended option, placing the discussed species here, from carinata to tenera, in Alvania sensu lato The re-analysis of the Neogene-Recent R. tiberiana, type species of Galeodinopsis, has lead me to reevaluate the latter taxon as the appropriate genus for grouping species characterized by a particular telec conch micro- sculpture (consisting of ve ry fine spiral threads, formed by roughly prismatic ele ments, extending perpendicu- larly respect from the shell surface, plus the pitted sur- face on the primary spiral cords, as seen in Manzonia) and by having a conical Alvania-like shell shape. The oldest representative of this genus is the European upper Oligocene R. duboisi. The upper Oligocene M. forami- nata and M. moulinsi are probably more linked to Man- zonia sensu lato than to Galeodinopsis while the Recent Macaronesian M. spreta could be doubtfully regarded as belonging to Galeodinopsis. Because of its affinity with Alvania, onsite probably derived from some Oligocene group of that ee and could be regarded as a transitional link to Manzonia. It is interesting that, as indicated by Lozouet (1998), the genus Alvania had a surprising radiation in the upper Oligocene, creating a strong diversification. Furthermore, the Manzonia sensu stricto spe cies, char- acterized by the quite slender shell with strongly opis- thocline ribs and ve ry marked spiral cords on the shell base, seem to be well-established from the European Neogene, where they are represented by the following species: M crassa, M. falunica, M. pontilevie nsis, and M, scalaris. All the discussed species have a multispiral proto- conch, indicating a planktotrophic larval development, with the exception of A. cingulata, which is the most geographically restricted species, limited to Sicilian wa- ters. The reason for its very limited geographical distri- bution is not certainly solely attributable to its non- planktotrophic larval development. Our knowledge about this species is not encouraging: I just know that: @ There is no known fossil record . . . Mediterranean acquisition? @ From a large amount of bulk samples (about 50 liters from Magnisi and 20 liters from Mondello), I picked only a single eroded shell! . . . Is it still a living species? is it a very recent Among species with planktotrophic development dis- cussed in this report, three show an eastern Atlantic- Mediterranean distribution: A. carinata, A. lactea, and A. tenera. Among these, the first two species have a Neo- gene to Recent distribution, being known from the Pliocene and the Miocene of Mediterranean basin, re- spectively, and show a quite extensive east Atlantic dis- tribution, being commonly recorded from the English Channel to Moreceo. Regarding the third species, living in Mediterranean and along the Atlantic Morocco aad the Canary Islands coasts, I “did not find any Mediterra- nean or Atlantic fossil records. The extinct species A. francescoi, A. prusi, and A. rosariae are limited to the Mediterranean Pleistocene, while Galeodinopsis tiberi- ana has a Miocene-Pliocene Mediterranean distribution and lives along the W African coasts from Mauritania to northern Angola. In general, the protoconchs of the species reported here do not indicate any distinguishing taxonomical char- acters at the supri aspecific, ane, in most cases, at the species level, being characterized by sculptural patterns shown by several rissoid species. Among the plank- totrophic species, the commonest sculptural pattern on protoconch I (observed in A. carinata, A. francescoi, A. lactea, A. rosariae, and A. tenera), consisting of fine spiral lirae and scarce to abundant pimples between them, is shown by A. cancellata (Da Costa, 1778) (see Giannuzzi- Savelli et al., 1996, fig. 408b), A. beani (Hanley in Thorpe, 1544) (see Giannuzzi-Savelli et al., 1996, fig. 412), A. cimex (Linneus, 1758) (see Ponder, 1985, figs. S6C-B), A. cimicoides (Forbes, 1844) (see Bouchet and Warén, 1993, fig. 1385 and Giannuzzi-Savelli et al., 1996, fig. 408d), 4. dingde nsis Uae n, 1967) (see Pon- der, 1985, figs. cas G), A. ge ryonia (Nardo, 1847) (see Giannuzzi-Savelli et al., 1996, fig. 3950), A. hispidula, Monterosato, 1554 (see Gotas, 1999, fig. 26), A. punctura (Montagu, 1803) (see Giannuzzi-Savelli et al., 1996, fig. 436), A. stephanensis Lozouet, 1998 (fig. 9D), Crisilla semistriata (Bouchet and Warén, 1993, fig. 1535), Rissoa parva (Da Costa, 1778) (see Ponder, 1985, figs. 79B), and several other species, The sculptural pattern on pro- toconch I (also observed in G. tiberiana), consisting of 1-3 more or less discontinuous, abapical to central, spiral threads (also formed by short, ments close to the beginning of the teleoconch) and tu- bereles on the remaining surface, is also shown by most fused, prosocline seg- of the above mentioned species and several others. Al- V. Garilli, 2008 Page 49 vania lactea could represent a slight exception, having a quite characteristic, easily distinguish: thle protoconch | with a coarser sculpture than that shown by the other studied species. A. tenera shows a similar protoconch I sculptural pattern, with few to abundant very small gran- ules spirally arranged, which may form very discontinu- ous and irregular ridge »s. Likewise this kind of sculpture is shared by other psseils (e.g. A. tarsodes (Watson, 1886) (see Bouchet and Warén, 1993, fig. 1450) and Crisilla semistriata). The sculpture of the paucispiral protoconch of A. cingulata, which is almost a replica of protoconch I of the first group discussed above, repre- sents a quite common pattern shown by several non- planktotrophic rissoids: e.g. A. argillensis Lozouet, 1998, A. macandrewi (iensont 1868), Lironoba multilirata (T. Woods, 1878), Onoba gianninii (Nordsieck, 1974) and, with a moderate similarity, by A. swbsoluta (Aradas, 1847), Onoba semicostata ( Montagu, 1803), and A. viro- dunensis Lozouet, 1998 (see Ponder, 1985, fis 89A, 109E and 126C; Bouchet and Warén, 1993, figs. 1458, 1525; Lozouet, 1998. figs. 1OF and 10k). ; There is similarity between the West African rissoid assemblages, including the Macaronesian province, and the European ones, with particular regard to those from the Mediterranean Neogene. This similarity is perhaps more marked than it has been indicated by Gofas (1999). The case of G. tiberiana is a further (see Monegatti and Raffi, 2001, and Garilli and Galletti, 2007) interesting case of a molluscan species that lived in the Mediterra- nean Neogene and today occurrs along the West African coasts. In this view, it is noteworthy to remark that, as a whole, most of the species here included in Galeodinop- sis lived in the European Oligo-Pliocene while its living representings occurs along the West Africa and Macaro- nesian Provinces. In addition: A. tenera, living in the Mediterranean, Atlantic Morocco, and in the Canary Is- lands (Tenerife), should be regarded as a new record, further supporting the Geciisse d similarity. The rissoid Rissoina dOrbigny. 1840, species from the Mediterra- nean Plio- Pleistocene (see the good illustrations of Greco, 1974, figs. 11, 13, 15, 17 ad Chirli, 2006, pl. 23 figs. 7-12). usually cited as R. decussata Moses 1803), is very likely the same taxon as living along the W African coasts, S40 Tomé and (¢ Cape Vena Islands [see Gofas, 1999: 97, figs. 69-73, and treated as R. punc- tostriata (Talavera, 1975)]. The rather common presence of varices on the last whorl (a rare character in rissoids) of A. carinata, A. francescoi new species, A. lactea, A. rosariae new spe- cies, and G. tiberiana should not be regarded as a salient taxonomic character at the supraspecific level, being present in quite unrelated species (e.g. A. carinata and G. tiberiana). ACKNOWLEDGMENTS This article would not have been possible without the generous support of friends, private collectors, and re- searchers. | particularly thank Maurizio Forli (Prato), Stefano Palazzi (Modena), Nino Adorna Sbrana (Gros- seto), Stefano pasa (Anguill: wa) for lending a large amount of precious and indispensable material. Many other people kindly made their collections available. R.G. Moolenbeek (ZMA, Amsterdam) donated three shells of Manzonia crispa from ZMA Moll. collection. Rossana Messina (Palermo) provided a special support, measuring most of the investigated shells and assisted me during the bibliographical research in the DGUP library. Luca Calle (Monreale), a companion for all the field- works in Kyllini and Cartiera Mulino, also help me in providing SEM images. My special thanks to M. Glaubrecht (ZMB, Berlin), who very kindly lent precious material (with particular regard to the two Alvania cin- culata shells from Monterosato collection), to D. Merle, J. M. Pacaud (MNHN-DHT, Paris), V. Héros, and P. Lozouet (MNHN-DSE, Paris), who Meer helpful and kind assistance during my visits in MNHN. A. Rosso and I. Di Geronimo (DSTC, Catania) kindly allowed the visit to the malacological collections of the DSTC. Kathie Way (British Museum of Natural History, London), H. J. Niederhéfer (Staatliches Museum fur Naturkunde, Stuttgart) and P. Serventi (MPOB, Modena) provided information on type material. Thanks to M. Fiore, P. [acopelli, V. P. Li Vigni (MSNCS, Terrasini), who al- lowed the visit to the conchological collections of the MSNCS. C. D’Arpa (MGUP, Palermo) lent some fossil material housed in the MGUP. Even when the library was moving, G. Barranca (DGUP, Palermo) very kindly allowed me the access to some papers. Riccardo Gian- nuzzi-Savelli (Palermo) and R. G. Moolenbeek helped me in some bibliographic researches. R. G. Moolenbeek and S. Palazzi also provided very useful and constructive comments on an early manuscript. I am grateful to D. Merle, R. G. Moolenbeek, H. Scholz (ZMB) and E. Theodorou-Vardala (GNHM, Ki- fissia), who ne provided the catalog numbers for type material of A. francescoi new species and A. rosariae new species. I am also grateful to H. G. Lee (Jackson- ville, Florida) and P. Lozouet (MNHN, Paris) who re- viewed the manuscript and provided valuable comments. H. G. Lee also kindly helped improve my shaky English. This work also benefited from critical re -adings by S. 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Zelaya Departamento Biodiversidad y Biologia Experimental Facultad de Ciencias Exactas y Naturales, U.B.A. and Division Zoologia Invertebrados Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, ARGENTINA dzelaya@tenym.unlp.edu.ar ABSTRACT Re-examination of the holotype of Cyamiocardium crassila- brum Dell, 1964, allowed confirmation that the species was wrongly allocated to the genus Cyamiocardium. This material, pe with additional specimens from the westemm Malvinas Falklands) Islands allowed for a re-description of that species. The species is also properly illustrated through scanning elec- tron microscopy, and its update sd generic place ment discussed. The presence of a “taxodont” hinge plate is the main morpho- logical character supporting the re-allocation of this species in the genus Perrierina Bernard, 1897. Information on the brood- ing condition of the species is provided. Additional Keywords: Perrierina, Cyamiidae, Magellanic Region INTRODUCTION Study of the mollusks collected by the R/V WILLIAM Scoresby in the Magellan Region and adjacent Antarctic waters resulted in he discovery of several species new to this region, among them, a small bivalve described by Dell (1964) as Cyamiocardium crassilabrum. The only subsequent record of this species was reported by Cas- tellanos (1980) from the western Malvinas (Falkland) Is- lands. The genus Cyamiocardium was introduced by Soot- Ryen (1951) to allocate C yamium dentic ulatum BE. A. Smith, 1907 (the type species by original designation), a species widely distributed throughout the Antarctic Re- gion (Lamy, 1910, 1911; Powell, 1958; Dell, 1990). Other Antarctic species assigned to this genus by Soot-Ryen (1951) were Cyamiocardium rotundatum (Thiele, 1912), C. dahli Soot-Ryen, 1957, and Cyamiocardium crassila- brum Dell, 1964, from the Magellan Region. In the con- text of a systematic revision of these species (currently in progress), the type material and additional specimens of “Cyamiocardium” crassilabrum were examined. As a re- sult, I have concluded in that the placement of this spe- cies in Cyamiocardium is incorrect. In the present paper the generic placement of this species is revised, and the taxon is re-described and properly illustrated by first time. MATERIALS AND METHODS This study is based on dried preserved specimens col- lected during the R/V SHINKAT Maru expedition (1978— 79). Voucher specimens have been deposited in the col- lections of the Museo de La Plata (MLP). Photographs of the holotype of Cyamiocardium crassilabrum were re- ceived from The Natural History Museum (NHM), Lon- don. For comparative purposes, specimens of Cyamio- cardium denticulatum, C. rotundatum, and C. dahli were also examined. The specimens reported by Castellanos (1980) as Cya- miocardium crassilabrum could not be located either at the MLP or the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN). The material here studied, originally at the Instituto Nacional de Investiga- ciones y Des: urrollo Pe ssquero (INIDEP), comes from the same SHINKAL MARU sampling station than the material previously reported by Castellanos (1980). Shell measurements were made under a stereoscopic microscope, according to the following criteria: L: maxi- mum antero-posterior distance; H: maximum dorso- ventral clistance, perpe ndicular to L: W: maximum dis- tance across valves, perpendicular to H. Shell morphol- ogy Was studie fal unde ‘y scanning ¢ le sctron mic roscope; for this, Philips XL30 TMP and JEOL JSN-6360 LV scan- ning Cc le ke microscope “Ss Were USe ad. D. Zelaya, 2008 SYSTEMATICS Cyamiidae G. O. Sars, 1S7S Perrierina Bernard, 1897 Perrierina crassilabrum (Dell, 1964) new combination (Figures 1-12) Cyamioc — crassilabrum Dell, 1964: 204, fig. numbers 1, 2, and pl. 6, figs. 1, 2; Castellanos, 1980: 13 Type Locality: 50°17’ S, 60°06" W, station 211, R/V WILLIAM SCORESBY, 161-174 m Material Examined: Photographs of the holotype (NHM 1962863); 7 dried specimens and | valve, 51° 29’ S, 61° 50’ W, Malvinas (Falkland) Islands, 192 m (MLP 12606). Literature Records: 50°17’ S, 60°06’ W, station 210, R/V WILLIAM Scoresby, 161 m; 50°35’ S, 57°20’ W, station 229, R/V WILLIAM Scoresby, 210-271 m (Dell, 1964); 51°29" S, 61°50’ W, 192 m (Castellanos, 1980). Distribution: Only known from the Atlantic sector of the Magellan Region, in the vicinity of the Malvinas (Falkland) Islands, 161-271 m. Description: Shell solid, small (maximum L: 6.3 mm), outline nearly circular (H/L = 0.99 + 0.01, n = 6), sli ghtly inequilateral, not inflated (W/H = 0.65 + 0.01, n = 5), whitish, glossy. Anterior margin short, curved, a ous with ventral margin, Ww hich is ev enly arcuate; poste- rior end rounded, slightly expanded (Figures 1, 2). Juve- nile shell slightly ovate in outline, elongated antero- posteriorly (Figure 3 3). Beaks full, chbeennil: directed slightl y anteriorly. Baiwaoneh ovate, smooth, about 590 wm in diameter (Figure 5). Shell surface sculptured with rounded, strongly marked and regularly spaced radial cords, 50-60 in larger specimens (Figure 6); cords sepa- rated by interspaces wider than ribs. Regularly sepa- rated, microscopic commarginal threads also present. Radial sculpture also evident on inner margins, where they produce prominent crenulations (Figures 7-9). Hinge plate narrow, somewhat broader anterior to beaks, where the cardinal teeth are inserted. Right valve with large, hooked cardinal 3: 3a high, solid, triangular, en- larged at base, which is Say bifid: 3b. delicate, narrow, nearly straight, one-third of size of 3a (Figures 7, 10). Left valve with prominent triangular cardinal 2, thin and styliform cardinal 4b behind resilifer, and : slender but solid, nearly straight anterior tooth (referred to as “cardinal 4a” by Berard (1597) and as “anterior lateral IP” by Lamy 1917)) (Figures 8, 9, 11). Both valves with two series of tubercles anterior and posterior to beaks, producing the appearance of a taxodont hinge plate (Fig- ures 10, 11); each series composed of four tubercles; posterior series stronger than anterior in juveniles ( Fig- ure 12). In both series, tubercles diminish in size and degree of development from beaks to anterior and pos- terior ends, where they are followed by marginal crenu- lations. Internal ligament somewhat solid, located in short, oblique resilifer posterior to cardinal teeth. Exter- nal ligament short. Biological Observations: One of the specimens ex- amined contained numerous embryos between the as- cending and descending lamellae of inner demibranchs. E mbryos were in different stages of development, rang- ing from incipient (unshelled) to well-developed speci- mens, the latter numbering 32, and reaching 700 pm in diameter (Figure 4). These observations are consistent with that by Dell (1964), who reported 44 “developing young” contained in one of the specimens he studied. Remarks: = Perrierina was proposed by Bernard (1897) for P. taxodonta (type species by monotypy; illustrated by Bernard, 1897; fig. 3) a species he described from fle Stewart (New Zealand). The description of the genus was merged with that of the type species, in which Bernard (1897) described the presence of several “lamellae” an- terior and posterior to the cardinal teeth, resembling a taxodont hinge. This character, infrequent among Cy a- mioidea, is also present in Legrandina Tate aed May, 1901 (type species: L. beriordr Tate and May, 1901, by original designation), an Australian genus regarded by Ponder (1971) and Powell (1979) as a subgenus of Per- rierina. Perrierina and Cyamiocardium have as common char- acters the number, morphology, and arrangement of hinge teeth, ie., the presence of a hooked anterior car- dinal 3 in the right valve and two prominent cardinal teeth (cardinals 2 and 4) in the left valve. The specimens studied here generally agree with these characteristics, but, additionally, the hinge examined showed several an- terior and posterior dubeniles which, as a group produce the appearance of a taxodont hinge, a diagnostic charac- ter for Perrierina that is absent in Cyamiocardium. Fur- thermore, the studied specimens lack the posterior “pseudo-lateral tooth” described by Smith (1907) for the type species of Cyamiocardium. Another character that is useful in separating the genera Cyamiocardium and Perrierina is the presence in the former of mantle margin papillae (Soot-Ryen, 1951). According to Ponder (1971), mantle margin papillae are absent in Perrierina. Untor- tunately, the poor prese rvation of the specimens on which this paper is based made it impossible to confirm the state of this character. Nevertheless, the differences in hinge morphology call for the reallocation of “Cya- miocardium” crassicostatum into Perrierina. Perrierina crassilabrum shows some characteristics in- termediate between the subgenera Perrierina and Leg- randina. The presence of prominent beaks is a charac- teristic shared with Perricrina, whereas a hinge plate broader at the base of the cardinals is characteristic of Legrandina. Furthermore, there are some other features of P. crassilabrum not previously known for other species of Perrierina or Legrandina, such as the relatively large adult size, the nearly circular shell outline of adults, and the strong radial ornamentation. THE NAUTILUS, Vol. 122, No. 1 Page 54 Perrierina crassilabrum. 1,9. Holotype (NHM 1962863). 1. External view of left valve. 9. Internal view of left valve. Figures I-12. 2-8, 10-12. Specimens from 51° 29'S, 61°50! W, SEM micrographs. 2. External view of an adult specimen, right valve. 3. External iew of a juvenile. 4. External views of larvae removed from an adult. 5. Dorsal view, showing protoconch and hinge plate 6. Detail of shell microsculpture. 7. Inner view of right valve. 8. Internal view of left valve, 10, 11. Detail of adult hinge plates. 10. Right valve anterior tooth, 12. Detail of hinge plate of a juvenile ri sb = cardinal teeth, L = ligament. 11. Left valve. 2, 4b = cardinal teeth, at Seale bars: 1-3, 7-9 = 1 mm; 4 = 100 wm: 5 = 200 jam: 6 = 50 wm; LO-12 = 500 pom of 2.3 mm length D. Zelaya, 2008 Page 55 The familial placement of Perrierina is somewhat con- fusing; the genus was a sssively placed within the Mactridae ( Bernard, 1897), Le apeont ie (Dall, 1899), and Crassatellidae (Suter, nae amy, L917). Later, Mar- wick (1928) regarded the “taxodont” lamellae of the hinge as a character sufficient to warrant proposition of the family Perrierinidae, but this family was subse- quently regarded as a synonym of C yamiidae ( (e.g. Cha- van oe Thiele, (1934), Fle sming (1948), Ponder (1971), and Powell (1979), The Cyamiidae is a family eee well-diversified in the Magellanic Region, where a total of 12 species belonging to Cyamium, Cya- miocardium, Gaimardia, and Kidderia have been re- ported and recognized as valid by Zelaya (2005). The present paper represents the first record in M: agellanic waters of a species of Perrierina, a genus thus far known only from Australia, Tasmania, and New Zealand. ACKNOWLEDGMENTS The author is grateful to A. Roux (INIDEP) who kindly made available the material collected during the R/V SHINKAI MARU cruise; K. Way and A. MacLellan (NHM) kindly sent photographs of the holotype of Cyamiocar- dium crassilabrum: C. Digiani (MLP) helped with older literature: and C. Ituarte provided valuable criticism and suggestions on an early version of the manuscript. The author is member of the National Research Coun- cil for Science and Technology (CONICET), Argentina. This study was partially supporte ed by a grant froth the Western Society of Malacologists, made possible by the Santa Barbara Malacological Society, the Southwestern Malacological Society, the San Diego Shell Club, and the Northern California Malacological Club, and by grant PICT 282 from “Agencia Nacional de Promocién Cientf- fica y Tecnolégica,” Argentina. LITERATURE CITED Bernard, F. 1897. Sur quelques coquilles des Lamellibranches de Il fle Stewart. Bulletin du Muséum d’ Histoire Na- turelle 7: 309-314. Castellanos, Z. A. de. 1980. Micromoluscos poco conocidos del sur argentino-chileno, Neotropica 25 (74): 133-140. Chavan, A. 1969. Superfamily Cyamiacea. Pp. 537-543. In: Leslie Reginald Cox et al., Part. N [Bivalvia], part. 6, vols. 1 and 2: xxxvii + 952 pp. Treatise on invertebrate paleon- Sieg Lawrence, Kansas. Geological Society of America & University of Kansas. Coan, E. V., P. V. Scott y F. R. Bernard. 2000. Bivalve seashells of Western North America. Santa Barbara Museum of Natural History. Monographs number 2, Studies in Biodi- versity n°2, 764 pp. Dall, W. H. 1899. Synopsis of the Recent and Tertiary Lep- tonacea of North America and the West Indies. Proceed- ings of the United States National Museum 21: 873-897, pls. S7-SS. Dell, R. kK. 1964. Antarctic and sub-Antarctic Mollusca: Am- phineura, Scaphopoda and Bivalvia. Discovery Reports 33: eee pls. 2-7, Dell, R. 1490. Antarctic Mollusca with special reference to the aan of the Ross Sea. Bulletin of the Royal Society of New Zealand 27: 1-311. Egorova, E. N. 1982. Biological results of the Soviet Antarctic Expeditions, 7. Explorations of the fauna of the seas 26 (34); 1-143. Fleming, C. A. 1948. New species and genera of marine Mol- lases a from the Southland a Transactions of the Royal Society of New Zealand 77 (1): 72-92. Lamy, E. 1910. Mission dans ee dirigée par M.Le Dr Charcot (1905-1910), collections recueillies par M. le Dr. Jacques Liouville. Bulletin du Muséum National d Histoire Naturelle 7: 388-394. Lamy, E. 1911. Gastéropodes prosobranches, scaphopodes et pélécypodes. Deuxieéme Expédition Antarctique Francaise (1908-1910) commmandée par le Dr. J. Charcot. Sciences Naturelles: Documents Scientifiques. 32 pp., pl. 1 Lamy, E. 1917. Révision des Crassatellidae vivants du Muséum dHistoire Naturelle de Paris. Journal de Conchyliologie 62 (4): 197-270 + pl. 6. Marwick, J. 1928. The Tertiary Mollusca, Catham Islands. Transactions and Proceedings of the New Zealand Insti- tute 58 (4): 432-506. Ponder, W. F. 1971. Some New Zealand and Subantarctic bi- valves of the Cyamiacea and Leptonacea with desc riptions of new taxa. Records of the Dominion Museum 7 (13): 119-141. Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda collected by the ships of the Discovery Committee during the years 1926-1937. Dis- covery Reports 26: 49-196, pls. 5 5-10. Powell, A. W. B. 1958. Mollusca from the Victoria-Ross quad- rants of Antarctica. B.A.N.Z.A.R. Expedition B, 6: 165— 215. Powell, A. W. B. 1979. New Zealand Mollusca. Marine, land and freshwater shells. Auckland, William Collins Publish- ers, 500 pp. Smith, E. A. 1907. Mollusca. 5. Lamellibranchiata. National Antarctic Expedition 1901-1904. Natural History 2, Zool- ogy (Vertebrata, Mollusca, Crustacea): 1-6. Soot-Ryen, T. 1951. Antarctic Pelecypods. Scientific Results of the Norwegian Antarctic Expeditions 1927-1925, 32: 1-46 + 1 pl. Suter, H. 1913. Manual of the New Zealand Mollusca. John MacKay, Government Printer, Wellington. 1120 pp. Thiele, J. 1934. Handbuch der sy stematischen Weichtierkunde. Gustav Fischer Verlag, Jena. Bd 3, Dritter Teil, pp.1193- 1528. Zelaya, D. G. 2005. The bivalves from the Scotia Are islands: species richness and faunistic affinities. Scientia Marina 69 (suppl. 2): 113-122. THE NAUTILUS 122(1):56, 2008 Page 56 Notice THE 2008 R. T. ABBOTT VISITING CURATORSHIP The Bailey-Matthews Shell Museum is pleased to invite applications for the 2008 R. T. Abbott Visiting Curatorship. The Curatorship, established originally in accordance with the wishes of the late Dr. R. Tucker Abbott, Founding Director of the Shell Museum, is awarded annually to enable malacologists to visit the museum for a period of one week. Abbott Fellows are expected, by performing collection-based research, to assist with the curation of portions of the Museum’s collection and to provide one evening talk for the general public. The Museum collection consists of marine, freshwater, and terrestrial specimens. 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Box 30 North Myrtle Beach, SC 29582 Dr. Gary Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr. Angel Valdés Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 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 SUBSCRIPTION INFORMATION The subscription rate per volume is US $47.00 for individuals, US $80.00 for institutions. Postage outside the United States is an additional US $10.00 for regular mail and US $25.00 for air delivery. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 1580, Sanibel, FL 33957, USA, (239) 395-2233. Change of address: Please inform the publisher of your new address at : Z least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (UISSN 0028-1344) is published quarterly by 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 P.O. Box 1580 Sanibel, FL 33957 NAUTILUS Volume 122, Number 2 June 25, 2008 ISSN 0028-1544 ay te CONTENTS Luiz Ricardo L. Simone Revision of the genus Spinosipella (Bivalvia: Verticordiidae ), with Carlo M. Cunha descriptions of two new species from Brazil .. 2... 0.000.002 57 Jonathan Hendricks Late Eocene Conus (Neogastropoda: Conidae) from Florida, USA... 2... 79 Roger W. Portell Janine O. Arruda Synonymization of Neohyalimax Simroth, 1596, and Omalonyx dOrbigny, José W. Thomé 1837, with a re Seine en of Omalonyx brasiliensis (Simroth, 1896) (Gastropoda: Succineidae) .. 2... 26. eee 94 Roland Houart Rehabilitation of Ergalatax martensi (Schepman, 1892) (Gastropoda: Muricidae), senior synonym of Ergalatax obscura Houart, 1996, and description of Ergalatax junionae, new name for Morula martensi Dall, 1923 ........ 99 Guido Pastorino Two new deep-sea muricids (Gastropoda) from Argentina... .......... 107 Fabrizio Scarabino MBLWHOI Library JUN 2 & 9Ang WOODS HOLE Massachusetts 02543 THE NAUTILUS 122(2):57-78, 2008 Page 57 Revision of the genus Spinosipella (Bivalvia: Verticordiidae), with descriptions of two new species from Brazil Luiz Ricardo L. Simone Carlo M. Cunha! Museu de Zoologia da Universidade de Sao Paulo Caixa Postal 42494 04299-970 Sao Paulo, BRAZIL Irsimone@usp.br ‘carlomagenta@gmail.com ABSTRACT A revision of the deep-water verticordiid genus Spinosipella is provided, based on conchological and anatomical characters. The genus is considered distinct from Verticordia (of which it was considered a subgenus) based on the strong ribs, prickly surface, reduction of lunula, relative large size, weakly spiral valve shape, and other ees The following species are considered in the genus: (1) Spinosipella agnes new species, ranging from Florida, U Ne to Rio de Janeiro, Brazil, and also including the Porcupine Abyssal Plain in the North Atlantic; (2) S. tinga new species, occurring from Rio de Janeiro to ie Grande do Sul, Brazil: S. ac a (Philippi, 1544), Pliocene fossil from ee es Italy; (4) S. deshayesiana (Fis. cher, 1862), from south and central ene. (S. ericia Hed- ley, 1911, the type species of the genus, was oe astes to bea new synonym of S. deshayesiana); and (5) S. costeminens Poutiers, 1981), from the tropical w es a The five species differ mainly in conchological details of the number and size of ribs, of the prickly sculpture, shape of the shell, of the hinge and the degree of convexity. Anatomical description is also provided for the two Pacific species, which differ among them- selves mainly by the size of the pair of renal folds. From the standpoint of anatomical characters, the more significant are: the wide lithodesma; the elongation of the auricles, crossing the roof of pallial cavity; a tall digital fold in posterior region of supraseptal chamber; the low but wide palps: the muscular, gizzard-like stomach; the complete separation of both constitu- ents of the hermaphroditic gonad (a ventro-posterior testicle and a centro-dorsal ovary), and a complete fusion of the visceral ganglia. Additional Keywords: Mollusca, Anomalodesmata, Septibran- chia INTRODUCTION The Verticordiidae is a family of Ag epee bivalves comprised of carnivorous and mostly d deep-water species. They are typically small (less than 10 mm) but some species reach 30-40 mm. They are mostly radially sculp- tured and usually have nacreous inner surface. The genus Spinosipella Iredale, 1930 (type Verticordia ericia Hedley, 1911, by original designation) is usually considered a subgenus of Vertieordia Sowerby, 1844 (e.g., Thiele, 1934; Moore, 1969; Abbott and Dance, 1983). The genus encompasses species with shell having prickly outer surface, lunule very reduced, thick walls. and generally larger size (up to 30 mm). In addition to the type species, S. ericia, three other species are cur- rently included in this genus, S. acuticostata (Philippi, 1844), from Atlantic and Mediterranean (middle Tertiary to Recent); S. deshayesiana (P. Fischer, 1862a) and S. costeminens (Poutiers, 1981), from Indo-Pacific. Some authors have considered S. deshayesiana as an Indo- Pacific occurrence of S. acuticostata (e.g., Nobre, 1936; Crozier, 1966; Rosenberg, 2005). Examination of worldwide samples, with an emphasis on the Western Atlantic, showed that two species actually exist in the Atlantic. Both are separate from the fossil S. acuticostata. In addition, it was possible to reorganize the Indo-Pacific species, mainly because of the abundant material deposited at the Muséum national d'Histoire naturelle, Paris (MNHN), which results from several ex- peditions. A revision of the taxonomy and a necessary re-definition of taxa are provided in this paper, as part of a larger project revising Western Atlantic mollusk tax- onomy, based on morphology. MATERIALS AND METHODS A detailed list of the material examined follows each species description. Specimens generally belong to mu- seum collections. Most material consists of shells exam- ined under a stereomicroscope. Some few Pacific samples have preserved soft parts in 70% ETOH. They were dissected by standard techniques, under stereo- microscope, with specimen immerse in alcohol. All dis- secting steps were digitally photographed; all drawings were made with the aid of a camera lucida. In the case of the material examined of Spinosipella deshayesiana and Page 58 THE NAUTILUS, Vol. 122, No. 2 S. costeminens, as the quantity of examined lots is very large, mainly thorugh courtesy of staff at MNHN (Paris), the list only contains the country and the quantity of specimens. The full list of examined lots of these speci- mens is being published elsewhere, in a complementary paper (Simone and Cunha, in press). Abbreviations used in figures are: am, anterior adduc- tor muscle; an, anus; au, ausicle: bs, byssus; by, byssal gland or furrow; ce, cerebral commissure; ce, ‘cerebral ganglion; ej, connective tissue; em, circular muscle layer; co, cerebro-visceral connective: er, crustacean inside stomach: ev, ctenidial (efferent) vein; dd, ducts to diges- tive diverticulae; dg, digestive diverticula; es, esophagus; fa, foot aperture of mantle; fm, posterior foot retractor muscle; fr, anterior foot retractor muscle; ft, foot: ga, genital ape rture; ge, gastric epithelium; gi, gill; he, Renienock ic, infra- septal chamber; in, intestine: ki, kid- ney; Im, lateral muscle; lo, longitudinal muscle layer; It, lithodesma: mb, mantle border; mf, fused mantle edge; mg, radial mantle gland; mo, moueh: mp, mantle ten- tacle: ms, mantle muscles of incurrent siphon; mt, mantle; mu, muscular tissue; ne, nephropore; nv, nerve; oy, ovary; pa, posterior adductor muscle; pe, pericar- dium; pg, pedal ganglia; pi, papilla of excurrent chamber roof; pm, pallial muscles; pp, palp; rs, renal fold; rt rectum; se, excurrent siphon; sh, shell; si, incurrent si- phon; sm, septum muscle; sp, septum, ss, style sac; st, stomach; su, supra-septal chamber; sy, crystalline style; ts, testis; um, shell umbo; ve, ventricle: vg, visceral gan- glia; vm, visceral mass. Abbreviations of institutions: AMS, Australian Mu- seum at Sydney, Australia; EGC, Emilio Garcia collec- tion, FMNH, Florida Museum of Natural History, Florida, USA; HGLC, Harry G. Lee collection; INVEMAR-MHNMC, project of Museo de Historia Natural Marina de Colombia; MHNMC, Museo de His- toria Natural Marina de Colombia (Programa de Biodi- versidad y Ecosistemas Marinos); MNHN, Muséum na- tional d'Histoire naturelle, Paris, France: MZSP, Museu de Zoologia da Universidade de Sao Paulo, Brazil; RLPC, Rafael La Perma collection (Universita di Bali, Italy). Specimens from other verticordiid species were also examined for comparative purposes. This material in- cludes: Haliris fischeriana Dall, 1881; UNITED STATES OF AMERICA. Florida, Fowey Light, 130 m depth, MZSP 19934, 2 valves (R.V. Eouis sta. 184). BRAZIL. Rio de Janeiro; 22°34’ S, 40°29" W, 213 m depth, MZSP 18751, | valve (on Laminarias, W. Besnard col, est. IX). Rio Grande do Sul: 30°42’ S, 49°03’ W, 182-156 m depth, MZSP 18750, 5 valves (21 Aug. 1972); 32°55! §, 50°34’ W, 99 m depth, UFRG 1688, | shell and 10 valves (sta. 45, 6839 dredge, 04/iv/1998) Euciroa sp. BRAZIL. Rio Grande do Sul, off Tramandar, 30°42’ S, 49°03’ W, 182-186 m, MZSP 18750, 5 valves (R/V W. Besnard, GEDIP sta. 1556, 21 Aug. 1972), MADAGASCAR. 600 m depth, EGC 23588, 1 shell. MOZAMBIQUE. off Moron- dava, Channel Madagascar, 600-800 m depth, MZSP 61516, 3 shells (Trawled by local fisherman, May 2002). Euciroa elegantissima (Dall, 1881). UNITED STATES OF AMERICA. Florida, 27°16’ N, $4°58.99' W, 457 m depth, EGC 13005, 6 valves (dredged, R/V PELICAN), 24°09’ N, 82°31’ W, about 64.3 km off Southwest of Ke y West, 549 m depth, EGC 23688, 1 shell (R/V OREGON II col, cruise #45, sta. 13362); Monroe Co. Straits of Florida, 549 m depth, FMNH 209892, 1 shell (Frank Lyman col.), 24°15.1' N, 82°11.71' W, 525 m depth, FMNH 164794, 1 valve (G.H. Burgess, et. al. GHB-90-5, 23 Apr. 1990). COLOMBIA. Santa Marta, Cerro de Punta Betin, A.A. 1016, (MHNMC INVEMAR), MHNMC 2782, 3 valves, MHNMC 2781, | valve. SYSTEMATICS Genus Spinosipella Iredale, 1930 Iphigenia Costa, 1850: 398 (type species by original designation ippagis acuticostatus Philippi. 1844) ( (pre -occupied) (non Schumacher, 1817). Spinosipella Iredale, 1930: 388 (type species by original desig- nation Verticordia ericia Hedley. 1911); Poutier and Ber- nard, 1995: 142. Verticordia (Spinosipella): Thiele, 1934: 1428; Moore, 1969: $55. Diagnosis: Shell relatively large, obese, with spiral valves. Surface prickly including on radial ribs. Radial ribs tall, extending beyond shell margin. Lunula very re- duced. es Sage SHELL: From small to medium size (up to 30 mm). Width/length ratio usually about 1. Color opaque-whitish. Sculptured by strong and tall radial, weakly curved ribs, triangular in section, bulging weakly beyond shell edge, alternating in both valves. Surface spiny, coasted by uniform sized, very small bulbs, covering almost entire outer surface. Lunula very re- duced. Umho projected, weakly spiral. Right valve with single tall, pointed and broad cardinal tooth. Left valve with low, broad tooth ( (posterior to tooth of right valve), and plane cardinal concavity as socket of tooth of right valve. Ligament just anterior to anterior hinge tooth, in- serted at some distance from median line, in approxi- mately middle way between hinge medial edge and um- bonal cavity (Figures 10, 12). I thode sma aide, curve, occupying about 0.25 of ‘hinge length, possessing a pair of lateral ligamental articulations (Figures 93, 96-105), List of Included Taxa: S$. acuticostata (Philippi, 1844): S. agnes new species; S. costeminens (Poutiers, 1981); S. deshayesiana (P. Fischer, 1862a) [=S. ericia (Henley, L911); S. tinga new species. Spinosipella agnes new spec ies (Figures 1-18, 27-29, 31, 55) Verticordia acuticostata.—Nobre, 1936; 303-304; 1938; 769— 770; Abbott, L974: 563; Abbott and Dance, 1983: 375 L. R. L. Simone and C. M. Cunha, 2008 Page 59 Figures 1-14. Spinosipella agnes new species. Shells. 1-10. Holotype (length 23.2 mm) shell. 1. Left valve, outer view. 2. Right valve. 3. Right valve, inner view; 4. Left valve, inner view. 5. Dorsal view. 6. Posterior view. 7. Anterior view. 8. Detail of shell surface in SEM, middle region of right valve. 9. Detail of inter-umbonal region, dorsal view. 10. Hinge, ventral-inner view. 11-14. Paratypes 11. HGLC. from Florida, left valve, outer view; 11 mm. 12. Same, ventral view, valves opened for showing whole view of hinge 13-14. EGC 17419. from Colombia, outer view of right and left valves; 15 mm Page 60 THE NAUTILUS, Vol. 122, No. 2 Figures 15-29. Spinosipella new species. Shells. 15-18. 8. agnes paratype AMNEL 162803, Florida, specimen with remains of soft l4.5 mm. 19-26. S. tinga type specimens. 19-21. Holotype, left valve, outer, inner and dorsal views; 16.9 mm, 22-24. Paratype MORG 18085, right valve, dorsal, outer and inner views; LO.1 mm, 25. Holotype detail of hinge, left valve. 26. Paratype MORG 18085, right valve, detail of hinge. 27-29. S. agnes idinal tooth. 28-29. Paratype INV-MOL 2943 ng mayor difference 27. Holotype, left valve, detail of hinge: arrow indicating well-developed from Colombia, a specimen ol equivalent size of main types of S. tinga compare with Figures 19-20): inner and outer views; 17.0 mm note developed posterio1 cardinal toot Inge wWYrow fewe! tall I ancl more spaced ribs anc more projected ribs at edges L. R. L. Simone and C. M. Cunha, 2008 Page 61 (fig.); van Aartsen, 1992: 45; Poppe and Goto, 1993: 139; ?McLean and Geiger, 1998: 27, 109 (fig.); Salas, 1996: 46; Rosenberg, 2005 (part) (non Philippi, 1$44). Verticordia deshayesiana.—Rosenberg, 2005 (part) (in syn- onymy) (non Fischer, 1S62a). Type Mater ial: HOLOTYPE, MZSP 36917; BRAZIL. Rio de Janeiro, off Cabo Frio, 23°41’ S, 41°03’ W, 750- SOO m depth (o.t.). PARATYPES. UNITED STATES OF AMERICA. Florida; Off Cape Canaveral, 903 m depth, USNM 64039, 1 right valve; SE of Sand Key, AMNH 248458, 4 shells (Jan. 1970, |. M. Bijur Collec- tion), AMNH 245459, 2 shells ee 1970, J. M. Bijur Collection); Monroe County, S.E. Sand Key, 270 m depth, HGLC, 2 shells, FMNH 154594, 1 specimen (dredged, Jerry Phelps col., Jun. 1970); 120.6 kin east of Daytona, 29°17’ N, 79°27" W, 878 m depth, USNM §10590, 1 shell and 1 left valve (R/V. OREGON, sta. 6690, 9 May 1967); Marquesas Key, 24°15’ N, 82°13’ W, 278— 419 m depth, 1 left, USN M 810889, 1 right valve (R/V. Blake, A. Agassiz 1877-1878). COLOMBIA. off Carta- gena, 10°28’ N, 75°42’ W, 280 m, MHNMC 2203, 1 valve (E-47), 10°31’ N, 75°37’ W, 309 m, MHNMC 2775. 1 valve (E-141), Palomino, Dibulla, 11°29’ N, 73°27' W, 476 m, MHNMC 3104, 4 valves, (E-21), Gua- jira, Bahia Honda, 12°31’ N, 72°8’ W, 452 m, MHNMC 2943, 1 shell (E-12), Guajira Peninsula, 12°30’ N, 72°08’ W, 470 m depth, EGC 17419, 1 shell; Cabo de la Vela, 12°19’ N, 72°42’ W, 464 m, MHNMC 3087, 2 valves (E-19), Islas del Rosario, °10" N N, 76°01’ W, 510 m, MHNMC 2208, 2 valves (E-78). BRAZIL. Rio Grande do Norte: 206 m depth, ne 84627, 1 shell (Sta. D-22. 10 Nov. 2001); Pernambuco; 690 m depth, MZSP 84628, 1 shell (Sta. D-11). Diagnosis: Shell with 15-17 radial ribs; smooth pre- umbonal region wide (about 0.25 of shell length); prickly sculpture chaotically organized. Width/] ength ratio in each valve approximately 0.57. Posterior eienmal tooth of left valve hinge well developed; main cardinal tooth of left valve relatively low and cylindrical; main cardinal tooth of right valve tall (about 0.2 of valve width) and pointed. Description: SHELL: Up to 22 mm, equivalve, inflated, each valve symmetrically and weakly spiral (1 whorl) (Figures 7, 9). Color white. Degree of convexity (width/ length) in each valve approximately 0.57. Outer surface spiny, opaque forming an irregular mosaic (Figures 8, 9). Umbones located in middle region of dorsal surface, spi- ral, high. divergent, separated from each other at about Vs of shell width ( Figures 1-4, 13-18, 55). Sculptured by strong, uniform, arched, radial ribs, from 15 to 17 in each valve. Posterior edge about twice as wide as anterior edge. Between umbo and anterior edge a concavity bear- ing transversal ribs, es wider than ribs of remaining region (Figures 5, 7, Pre-umbonal region smooth, narrow, 0.2 of shell le set (Figures 5, 7, 9). Anterior, ventral and posterior edges forming zigzag (Figures 3, 4): tips of this zigzag coinciding with tips of each a ly encasing in concavity of opposite valve (Figures 1, 2, 11, 13-16). Inner surface iridescent, whitish, glossy; includ- ing hinge (Figures 3, 4, 10, 12, 17, 18). Hinge with a large eandinal tooth in right valve, stubby, tall (about 20% of valve width), broadly pointed, we vakly curved forwards (Figures 3, 10, 12, 18), circular in section: correspondent socket in left valve shallow, restrict to dorsal surface; this socket flanked by small tooth in each side, anterior smaller and lower than posterior (Figures 4, 10, 12, 17, 27-28 |arrow]). Ligament just anterior to anterior hinge tooth, inserted at some distance from median line (Fig- ures LO, 12), approximateh sly midw: ay between hinge me- dial edge and umbonal cavity. Sears of adductor muscles shallow ( (Figures 3, 4, 17, 18, 28): anterior scar elliptical (longer “igiee vere), located close to anterior edge, area about Vis of inner surface of valve; posterior scar circular, about 0.33 larger than anterior scar, located close to posterior shell edge. Pallial line continuous, lo- cated at wide distance from shell ec dge, about 0.33 of distance between ventral and umbonal height. Measurements cone height, width, in mm): Holotype: 20.1 by 23.2 by 22.2; EGC 17419: 15.6 by 15.5 by 15.5. Geographic Distribution: Florida, USA, to Rio de Janeiro, Brazil. Habitat: Material Examined: Types. BARBADOS. USNM 63200, 3 valves (Blake Coll., sta. 100). CUBA. Havana; Gulf of Mexico, 419 m depth, USNM 63201, 3 right, 4 left valves (Blake Coll., sta. 5). PORTUGAL. Porcupine Bank; USNM 63204, 2 right valves (Jeffreys Coll., Por- cupine Exp. 1870). UNITED STATES OF AMERICA. Florida; Gulf of Mexico, off Cape San Blas, 309 m depth, USNM 323871, 1 left, 1 right valve (sta. 2400); 120.6 km east of Daytona, 29°17’ N, 79°27’ W, 878 m depth, USNM 810590, 1 shell and 1 left valve. BRAZIL. Es- pirito Santo (R/V MartIon-DUFRESNE MD55, May 1987); off Conceigao da Barra, 18°59’ S$, 37°50’ W, 637 m depth, MNHN, 10 valves (sta. CB76); off Pontal da Regéncia, 19°34’ S, 38°55’ W, 340-360 m depth, MNHN 1 valve (sta. CB92). Muddy bottoms, 270-900 m. Etymology: The specific epithet refers from the Greek agnes, meaning pure, an allusion to the whitish color of the shell. Remarks: The above listed examined material that was not designed as types are normally lots with eroded specimens, or sometimes they have aberrant characters. This is the case of the MNHN material collected off north coast of Espirito Santo, Brazil. They actually are free valves that resemble the Pacific species Spinisopella costeminens, in having a weakly larger radial thread be- tween middle and posterior thirds. ond in lacking ante- rior tooth in hinge. As they can represent another spe- cies, they are not designec Tas types; on the other hand, the material is not sufficiently well-preserved for further analysis. Because of they can only represent an extreme Page 62 THE NAUTILUS, Vol. 122, No. 2 of variation of the S. agnes, they are listed as additional examined material of this species. Spinosipella tinga new species (Figures 19-26, 30, 32) Verticordia acuticostata: Marini, 1974: 242, figs. 5, 6 (non Phil- ippi, 1544). Verticordia (Haliris) acuticostata: Rios, 1975: 262, pl. 85., fig. 1261; 1985: 282, pl. 99, fig. 1391; 1994: 304, ‘i 104, figs. 1489 (non Philippi, 1844). Verticordia deshayesiana: Marini, 1974: 242 (in synonymy); Rios, 1975: 262; 1985: 282; 1994: 304 (in synonymy); Rosenberg, 2005 (part) (Gin synonymy) (non Fischer, 1862a). Diagnosis: Shell with 17-18 radial ribs; smooth pre- umbonal region very narrow (less that 1% of shell length). W idth/length ratio in each valve approximately 0.47. Pos- terior cardinal tooth of left valve hinge absent; main car- dinal tooth of left valve low and cy adie: al; main cardinal tooth of right valve lower (about 10% of valve width) and rounded. Description: SiHeLL: Up to 11 mm, equivalve, in- luted, both valves weakly spiral (1 whorl) (Figures 21, ). Color white. Degree of convexity width length) in i valve approximately 0.47. Umbones located in middle region of dorsal surface (Figures 19, 20, 23, 24); umbones weakly spiraled, somewhat high, divergent, separated from each other. Sculpture of strong, uniform, arched, radial ribs, 17-18 in each — Outer surface opaque, covered by a mosaic of small, blunt, loosely aligned spines par allel to radial ribs ( Figure 3 2). Anterior edge almost same size as posterior edge. A concavity bearing transversal ribs of same width as remaining ribs between umbo and posterior edge (Figures 21, 22, 30). Anterior, ventral, and posterior edges rounded, tips of ribs prominent (Figures 19, 20, 23, 24), fitting with con- cavity in opposite valve. Inner surface iridescent, whitish, glossy, including hinge. Hinge with a somewhat large cardinal tooth in right valve (Figures 20, 25); cardinal tooth stubby, tip rounded, flat in cross-section, tooth length about 10% of valve width: correspondent socket in left valve shallow, restricted to dorsal surface; this socket flanked by small, low, posterior tooth (no anterior tooth) (Figures 24,26). L igament just anterior to anterior hinge tooth, inserted at some distance from midline, approxi- mately midway be stween hinge medial edge and umbonal cavity. Scars of adductor muscles shallow ( (Figures 20, 24); anterior scar elliptical (longer dorso-ventrally), lo- cated close to anterior edge, area about Vis of inner sur- face of valve; posterior scar circular, about / larger than anterior scar, located close to poste rior shell edge. Pallial line with a very weak pallial sinus, located at wide dis- tance from shell e dge, about “% of distance between ven- tral and umbonal height. Measurements (respectively length, height, width, in mm): THolotype: 16.9 x 15.3 x 9.6 (single valve); MZSP 18752: Paratype #1, $8.6 x §.6 x 4.1 (1 valve): Paratype #2, 11.9 x 11.4 x 5.4 (1 valve); MZSP 18753: 9.5 x 9.6 x 4.8 (1 valve). Type Material: Holotype, MZSP 19345, 1 valve, from type locality (R/V W. BEsNarD, GEPID Est. 458, 9 Dec. 1968. Paratypes, Rio de Janeiro, Cabo de Sao Tomé, 31°08’ S, 49°31’ W, 182-253 m,.1 valve, MZSP 18752 (R/V W. BESNARD, GEDIP st. 1858, 6 Aug. 1972): 22°34’ S, 40°29’ W, 213 m, 1 valve, MZSP 18753: (R/V W. BESNARD, st. IX, 1] Feb.1969), 100 m, 2 valves, MORG 18085 (R/V ALMIRANTE SALDANHA, Mar. 1972), off Solidao, 240 m, 2 valves, MORG 31888 (R/V ATLANTICO SuL, Exp. Coltro, 14 Oct.1993). Type Locality: BRAZIL. Rio Grande do Sul, off AL bardao, 33°29’ S, 50°44’ W, 200 m, muddy bottom. Geographic Distribution: ro to Rio Grande do Sul. Brazil, from Rio de Janei- Etymology: The specific epithet refers to the color white of the shell, from the Tupy language: tinga. Spinosipella acuticostata (Philippi, 1844) (Figures 33-40) Hippagus acuticostatus Philippi, 1S44: 42 (pl. 14, fig. 19) [fossil in Lamati valley, Calabria, Italy]. Verticordia acuticostata: Micali and Villari, 1991: 353. Spinosipella acuticostata: Poutiers and Bernard, 1995: 143, 150; Diagnosis: Shell with 12-13 radial ribs; pre-umbonal region narrow, smooth; posterior cardinal tooth of left valve hinge shallow or absent; main cardinal tooth of left valve longer and flat (Figures 35, 40); main cardinal tooth of right valve shallower (Figures 34, 37) (about 10% of valve width). Description: SHELL: Up to 24 mm; width/length ratio approximately 1 (Figures 36-35) to 1.5 (Figures : 39, 40). Degree of convexity (width/length) in each valve approxi- mately 0.55. Outer surface spiny, opaque, spines forming radially aligned mosaic parallel to ribs (Figures 36, 38, 39). Sculpture of strong, uniform, arched, radial ribs, 12 13 in each valve. Posterior edge about twice as broad as anterior edge. A concavity bearing transversal ribs weakly broader than ribs of remaining region between umbo and anterior edge (Figures 35, 39); pre-umbonal region narrow, smoooth (Figure 33), about 10% of shell length. Anterior, ventral, and posterior edges forming zigzag (Figures 35, 37, 40). Hinge with a large cardinal tooth in right valve, stubby, tall (about 10% of valve width), broadly pointed, weakly curved anteriorly (Fig- ures 34, 37), circular in section: corre spondent socket in left valve shallow, restricted to dorsal surface; this socket sometimes flanked by small tooth in each side, anterior absent or very weak (Figures 35, 40). Measurements (respectively length, height, width, inmm): KRLPC #1: 11.4 x 14.5 x 6. 6 (valve); #2: 10.0 x 9.8 x 4.7 (valve). Geographic Distribution: Mediterranean. Pliocene fossil from south Italy (Calabria and Sicily). L. R. L. Simone and C. M. Cunha, 2008 Page 63 Figures 30-47. Spinosipella species. Shells. 30. S. tinga, Holotype, left valve, dorsal view. 31. S. agnes, Paratype INV-MOL 2943 from Colombia, a specimen of equivalent size of Holotype of S. tinga for showing major differences (compare with Figure 30); dorsal view: 17.0 mm: note higher convexity, fewer, taller and more spaced ribs. 32. S. tinga, left valve, SEM of Holotype, showing prickly S( ulpture 3340. S. acuticostata. 33-38. Pliocene fossil from Messina, Italy, USNM 63202. 33 Specimen 2 dorsal-slightly anterio1 view. 34. Specimen 3 right valve, detail of hinge. 35-36. Specimen |] left valve, inner and outer views; 13.5 mm. 37-38. Specimen 2. right valve, inner and outer views, hinge broken; 12.2 mm. 39-40. RLPC, from Rometta, Italy, left valve specimen with long shape; 19.0 mm. 41-47. S. deshayesiana. 41-43. Paratype 1 of S. ericia AMS 032068, left valve, inner, outer and dorsal views; 3.5 mim. 44. Paratype 2. right valve, inner view; 2.6 mm. 45 Type specimen of S japonica ANSP 49639, right valve: 5.2 mm. 46-47. ANSP 292986 (from India), right valve, outer and inner views; 10.5 mm Page 64 THE NAUTILUS, Vol. 122, No. 2 Paleohabitat: bathyal environments. Material Examined: ITALY. Sicily, Messina, 38°11’ 15°34' E, Seguenza, USNM 63202, 2 left, 2 right valves. Middle Pliocene outcrops at Rometta, 4 valves, RLPC. Spinosipella deshayesiana (Fischer, 1862) (Figures 41-54, 66, 67, 72-82, 93-102) Verticordia Deshayesiana Fischer, 1862a: 35-36 11) [China Sea]. Verticordia japonica A. Adams, 1862: 224. Verticordia ericia Hedley, 1911: 96; Prezant, 1998: 421 (fig. 9.16A) Spinosipella deshayesiana.—Poutiers and Bernard, 1995: 110— 112, 143, 159, 161 (figs. 7-9). Spinisopella ericia.—Poutiers and Bernard, 1995: 143, 159. Verticordia acuticostata—McLean and Geiger, 1998: 109 (non Philippi, 1544). (pl. 5, fig. 10- Diagnosis: Shell with 16-19 radial ribs uniformly dis- tributed, closely packed; pre-umbonal region narrow, smooth. Each rib bearing well- developed crests with small, prickly granules. Posterior cardinal tooth of left valve hinge abecak: main cardinal tooth of left valve low and flat, “with insertion of anterior valve edge approxi- mately in middle region of this tooth; main car Aint il tooth of right valve high (about 10% of valve width) and pointed. Description: SHELL: Up to 1S mm. Color white. De- gree of convexity (width/length) in each valve approxi- mately 0.57. Outer surface spiny, spines organized some- what radially, parallel to ribs; each rib with well- developed crests with small, prickly granules (Figures 42 45, 46, 48-54). Sculpture of strong, “uniform, arched, ra- dial ribs, 16-19 in each valve (Figures 42, 45, 46, 50), somewhat closely packed. Posterior edge about twice broader than anterior edge. A concavity bearing trans- versal ribs similar to ribs of remaining region present between umbo and anterior edge (Figures 43, 53); pre- umbonal region narrow, smooth, about 10% of shell length (Figures 43, 53). Anterior, ventral, and posterior edges forming zigzag (Figures 42, 44, 47, 50, 51, 59, 67, 100), with tips projecte ri longer, and narrower. Hinge with a large cardinal tooth in right valve, stubby, tall (about 10% of valve width), broadly pointed, somewhat flat (Figures 44, 47, 51, 59, 67); correspondent socket in left valve shallow, restrict to dorsal surface; this socket flanked by small posterior tooth, with insertion of ante- rior valve edge approximately in middle region of this tooth (Figures 41, 50), anterior tooth absent (Figures 41, 50) LITHODESMA (Figures 93, 96-99): Saddle-shaped, hemi-cylindrical. Dorsal surface concave (Figures 97— 98), flanking ventral surface of hinge, along Vs of hinge length; located just posterior to teeth. Left and right edges straight, turned upwards and medially, connected with valves by dark-brown ligament inside umbonal cav- ity closer to hinge inner edge (Figures LOO-102). Outer Middle and upper Pliocene beds of surface convex, covering dorsal-middle, inter-umbonal region of visceral mass (Figure 75). Anterior and poste- rior edges concave; anterior edge slightly deeper and with tenuous slope. Both edges covered by opaque, yel- lowish periostracum (Figures 96, 101, 102), Lithodesma thickness equivalent to that of shell. Major MUSCLES (FIGURES 72-75, 77, 78, 80, 91, 92): Both adductor muscles similar in size and position (Fig- ures 72-75), near valve edges; insertion size equivalent to 1/20 of valves inner surface each; approximately two times taller than wide; outer length about half of inner length, with insertion in valves gr ndatly oblique (Figures 75, 91, 92). Anterior adductor muscle with anterior re- on about 3 times narrower than posterior region, di- vided transversally (dorsoventral) in two similar halves (quick and slow components). Posterior adductor muscle similar to, but inverted arrangement in comparison to anterior adductor muscle; components different, how- ever, one of them horseshoe-shaped, occupying ventral and posterior sides (Figure 77); another component fill- ing internal region of muscle, only exposed in posterior and dorsal siden ( Figure 77). Pair of anterior foot retrac- tor muscles long aud narrow (Figure 80); originating just dorsal to anterior adductor muscle in area equivalent to 1/10 of adductor (Figures 75, 80); running ventrally and posteriorly; spreading after insertion in anterior and lat- eral regions of foot base. Pair of posterior foot retractor muscles similar to anterior pair, but about half narrower (Figures 75, 78, 80); originating just dorsal to posterior adductor muscle in area equivalent to 1/20 of that ad- ductor; running ventral and anteriorly; inserting in pos- terior and lateral regions of foot base. Pair of palp muscles, septal and pallial muscles described below. Pair of foot protractor muscles absent. Foor AND Byssus (FIGURES 73, 74, 80): Foot conical, pointed; estimated volume equivalent to 1/6 of that of chamber of valves; base located in middle region of ven- tral surface of visceral sac. Byssal furrow shallow and very narrow, length about half of that of foot, offset ventrally and distally, lying along posterior surface and midline, ending at short distance from foot apex. Byssus found in a single specimen, brown, with single filament, narrow: proximal end attached to distal region of byssal furrow. MANTLE (FIGURES 72-74): Dorsal fusion of mantle lobes about 3 of their edges, along entire hinge length and about “% of valves height toward ventral, in both sides. Edges of mantle lobes with two folds. Inner fold fused between two lobes along entire posterior half (ex- cept for siphonal apertures) (Figure 74). Both lobes free from each other along anterior half, up to dorsal level of anterior adductor muscle; in this region both folds are of similar size, with height equivalent to 25 of valves height. Mantle edges thick, muscular, insertion relatively thick i in pallial line (Figure 72, pm). Pallial muscles originating in pallial line in location about 13 from ventro-dorsal dis- tance; no clear pallial sinus. Incurrent siphon as aperture of a septum formed by fusion of inner mantle edge folds; aperture about 5 of posterior fused region of mantle, L. R. L. Simone and C. M. Cunha, 2008 Page 65 gures 48-63. Spinosipella species. Shells. 48-54, 56-59. S. deshayesiana, adult specimens. 48-53. MNHN (Sta. CP1475, Fiji 5 mm. 48. Left valve, outer view. 49. Right valve, outer view. 50. Left valve, inner view. 51. Right valve, inner view. 52. Posterior view. 53. Anterior view. 54. Dorsal view, HGLC, from Philippines; 11.8 mm. 55. S. agnes paratype, BMNH, 18.2 mm. 56-59. Syntypes of S. deshayesiana MNHN. 56. Outer view, specimen 1, left valve. 57. Outer view, specimen 2 (possibly figured by Fische1 1$62a), right valve. 58. Inner view, specimen 1. 59. Inner view, specimen 2; length = 8 mm. 60-63. S. costeminens Holotype MNHN 60. Outer view, left valve. 61. Outer view, right valve. 62. Inner view, left valve. 63. Inner view, right valve; 17 mm a Page 66 THE NAUTILUS, Vol. 122, No. 2 Figures 64-71. Spinosipella species. Shells. 64-65. S. costeminens Holotype MNHN. 64. Right valve, anterior view. 65. Left valve, anterior view; 17.0 mm. 66-67. S. ericia Holotype AMS, right valve. 66. Outer view. 67. Inner view; 5.8 mm. 68-71. S costeminens lacking projections on ribs, MNHN (Sta. CP 992, Vanuatu), right valve. 68. Outer-right view. 69. Anterior view. 70. Posterior view. 71. Inner view; 29.0 mm. longer dorso-ventrally (Figure 76); walls thick, muscular; outer surface flanked by 9 tentacles surrounding siphonal aperture; all tentacles of similar size, turned inwards, somewhat conical, tip blunt and rounded, length equiva- ent to that of siphonal aperture; single unpaired tentacle ocated ventrally; five secondary smaller tentacles located externally, midway between siphonal aperture and mantle edge, of similar size, about Ys of size of major entacles, well separated from each other, one of them ocated in ventral region of siphonal aperture, other four ocated laterally, in ventral half of siphon (Figure 76). Incurrent siphon a small pore located in small elevation, approximately midway between excurrent siphon and nge: a pair of small tentacles similar to secondary ten- acles of incurrent siphons, located laterally, in dorsal region of siphon base (Figures 74, 76, 77). Radial mantle gland present along mantle edges outer fold (Figures 73, 92, mg), occupying about half of outer fold volume, situ- ed closer to inner surface of this fold. 79 19 PALLIAL CAVITY (FIGURES 75, 94, 95): Occupying about 70% of volume of valves. Transversal, horizontal septum located approximately midway in animal, i.e., su- pra- and infra-septal chambers of equivalent length (Fig- ure 72). Paired palps low, wide, bilobed folds (Figures 73, 8] ( )5, pp) that occupy anterior third of dorsal sur- face of infra-septal chamber, permanently open as a fun- nel. Pair of palp muscles (Figures 72, 74, 79, lm) located laterally; originating in anterior region of umbonal cavity, in a distance from origin of anterior foot retractor equiva- lent to ¥3 of anterior adductor muscle height; located in same horizontal level of origin of anterior foot retractor; size equivalent to V4 of that of anterior foot retractor; running ventrally attached to mantle for a distance equivalent to Y of valve height: spreading after insertion in lateral region between inner and outer hemipalps. Palp muscles also connect anterior end of septum. Sep- tum with two constituents: external one produced by a fold of mantle (about 73 of septum area); internal pro- duced by gill (Figures 73, 95). External septum element thick, muscular; posterior muscles originating as a pair, just dorsal to posterior adductor muscle (Figure 77, ms): running ventrally immersed in mantle, at some distance from each other (equivalent to half of their width) and from midline, gradually becoming wider and thicker, in anterior surrounding posterior surface of posterior ad- ductor muscle and lateral edges of excurrent siphon; some secondary muscular bundles originating from cen- troposterior region of posterior adductor muscle uniting with main, vertical bundles (Figure 77); muscles spread- ing within septum in region between incurrent and ex- L. R. L. Simone and C. M. Cunha, 2008 Page 67 gi au Figures 72-75. Spinosipella deshayesiana. Anatomy. 72. Whole specimen just extracted from shell, right view. 73. Same, right mantle lobe in its infra-septal region removed, right-slightly ventral view, left shell valve also shown. 74. Same, right mantle lobe almost completely removed, right portion of septum also removed. 75. whole specimen, dorsal-slightly right view, most of mantle and dorsal integument artificially shown as transparent, lithodesma (It) shown in its in situ topology. Scale bars = 2 mm Page 65 THE NAUTILUS, Vol. 122, No. 2 pa ; . vg pl ne ga he ft PP Figures 76-79. — Spinosipella deshayesiana. Anatomy. 76. Detail of region of siphons, posterior-slightly right view. 77. Peri-anal chamber, right view, adjacent region of right mantle lobe sectioned and deflected to show inner surface and muscles, inferior region or right mantle lobe removed along median line. 78. Reno-pericardial region and adjacent structures, right view, right wall of pericardium removed. 79. Whole right view, showing topology of genital system, reno-pericardial structures, palps, main ganglia and muscles, most structures artificially shown as transparent. Scale bars = 2 mm L. R. L. Simone and C. M. Cunha, 2008 Pase 66 Page 69 80 Figures $0-S2. Spinosipella deshayesiana. Anatomy. 80. Whole right view, emphasizing digestive structures, main musculature and main nervous ganglia; topology of some adjacent structures also shown, everything else represented by transparency. 81. Same, anterior region of digestive structures opened longitudinally, some objects inside stomach preserved, topology of some adjacent structures also shown, 82. Visceral ganglia (left), ventral view, and pedal ganglia (right), postero-dorsal view. Scale bars = 1 mm. current siphons. Outer component of septal muscles in- serted in shell just ventral to posterior adductor muscle (Figure 72, sm), in area equivalent to Yio of that of ad- ductor muscle insertion. Internal element of septum constituted by gills. Gill with both demibranchs narrow, of similar size, flattened, in same plane of remaining septum: both gills surrounding posterior and lateral re- gions of foot base (Figures 73, 75, 95). Gill attached to remaining septum via tissue; gill attachment to foot by cilia. Connection between gill filaments of 6-7 longitu- dinal, equidistant bridges of similar width of filaments. Papilla situated in posterior region of roof of supraseptal chamber (Figures 78-80, 94, pi), positioned just ventral to visceral ganglia, internally solid: length about Yo of posterior adductor muscle length and about % of it in width: tip broadly pointed, normally turned to anterior. VISCERAL MASS (FIGURES 72, 75, 79): Strongly bilobed, as internal mould of well-separated umbos (Figures 72, 75). Most dorsal structures, just inside valve apexes, formed by sponge-like connective tissue. Pair of ovaries cream in color, occupying central and dorsal regions sur- rounding stomach and digestive diverticula, reaching dorsal areas up to dorsal sponge-like connective when fully developed. Testes brown, consistence harder, lo- cated ventrally and laterally, totally separated from ova- ries; anterior region irregularly digitiform (Figures 75, 79, ts). Digestive diverticula situated compressed be- tween stomach and gonads, color greenish-beige; occu- pying about “% of visceral volume. Stomach and intestine lying in central region, occupying about “4 of visceral volume (Figures 80). Reno-pericardial structures located just anterior to posterior adductor muscle and posterior foot retractor muscles, with volume approximately “% of visceral volume (Figures 75, 79). CIRCULATORY AND EXCRETORY SYSTEMS (FIGURES 75, 78): Pericardium located at short distance anterior to posterior adductor muscle; with about half of reno- pericardial volume, and with a pair of expansions toward anterior, surrounding roof of pallial cavity where lies pair of auricles. Auricles connecting to anterior end of gills, in short isolated ctenidial vein (Figure 75, cv); abruptly curving towards posterior and dorsal; after this curve, auricles increasing gradually, surrounding obliquely pe- riphery of visceral mass in roof of pallial cavity (Figures 72, 75), walls thin, translucent: close to midline auricles abruptly narrowing and connecting to ventricle (Figures 75, 78); posterior region relatively lobed. Ventricle lo- cated in center of pericardium, surrounding intestine: relatively narrow. Kidney mostly solid, color dark purple- almost black; most of renal gland located just anterior to posterior adductor muscle, ventral to pericardium (Fig- ures 75, ki); a pair of folds originating from this region, Page 70 THE NAUTILUS, Vol. 122, No. 2 running long roof of pallial, supraseptal cavity, just ven- tral and external to auricles (Figures 74, rf), this pair of folds with about ¥% of supraseptal chamber height, run- ning posteriorly in middle region of roof of this chamber, gradually approaching visceral mass towards anterior, fusing to visceral mass after running about “% of chamber length (Figure 74). Pair of nephropores as small slits located in posterior region of supraseptal chamber, cov- ered by posterior end of renal fold, just dorsal to of posterior retractor muscles of foot ( Figure 74, ne) DIGESTIVE SYSTEM (FIGURES 80, 81): Palps partially described above (pallial cavity), widely fused as pair of folds along midline (Figures 95, pp). Mouth central ( Fig- ure 95, mo), with sphincter relatively well developed. Esophagus with about 5 of visceral mass length, not attached to anterior adductor muscle, width about ¥ of that of anterior adductor muscles; wall relatively thick, muscular; inner surface with about 20 longitudinal, nar- row, low folds as continuation from those of palps (Fig- ure 81). Stomach main chamber with about “4 of visceral mass volume, elliptical, anteroposteriorly longer; walls thick, muscular (Figure 91, st). Gastric inner surface smooth; two pairs of ducts to digestive diverticula present, each one located in ventro-lateral region just posterior to esophageal insertion. Stomach normally con- taining 3-4 isopod crustaceans (Figure $1, cr) ). Style sac with about / of gastric main chamber volume, Ipeater lin middle of gastric ventral wall, somewhat elliptical (longer dorso-ventrally); ory stalline style occupying entire style sac (Figure Sl, inner surface of style sac smooth, lacking any fold ete it from intestine; gastric shield lacking. Intestine a single sigmoid loop with about half of style sac width. Inner surface simple, smooth. Intestinal portion crossing through pericardium in some- what anteroposterior direction. Rectum attached to dor- sal and posterior surface of posterior adductor muscle, with about 7% of remaining intestinal width. Anus simple, sessile, located in ventral third of posterior surface of posterior adductor muscle (Figure 77). GENITAL SYSTEM (Partially described above under VIs- CERAL Mass): Pair of testes and ovaries conv erging to a single common, short duct, of about “is of visceral mass length. Genital pores small slits located at short distance from ne phropores ( (Figures 74, 75, 94, ga). CENTRAL NERVOUS SYSTEM (FIGURES 80, 82): Cerebral ganglia somewhat triangular, each ganglion with volume equivalent to Vis of that of anterior adductor muscle; anterior end narrow, possessing thick pair of nerves run- ning to pallial region dorsal to palps; pair of ventral nerves also thick, originated in middle region of ganglia, running ventrally to palps; Posterior a originating (Figures SO, 106); cere bral commissure length about 3 of posterior surface of an- cere bro- visce ral connective terior adductor muscle. Pair of cerebro-visceral commis- sures relatively thick, running though visceral mass be- tween stomach and testes. Pair of pedal ganglia located in ventral third anterior pair of pedal retractor muscles, touching these muscles, both totally fused with each other along midline, almost forming a sphere, vol- ume of both equivalent to that of each cerehial ganglion; pedal nerves and cerebropedal connectives originating subterminally in posterior surface of ganglia. Pai of vis- ceral ganglia located anterior to ential surface of poste- rior adductor muscle; both also totally fused with each other along median line, being somewhat squared in ven- tral view; size equivalent to that of pedal pair of ganglia; cerebrovisceral connectives and siphonal nerves located in vertices. Measurements (respectively length, height, width, inmm): HGLC: 11.5 by 12.2 by 12.0; MNHN (Sta. DW11): 15.7 by 17.7 by 9.2 (valve); MNHN (Sta. CPS889): 19.7 by 17.0 by 9.1 (valve). Geographic Distribution: South and Central Indo- Pacific in 146-805 m depth. Material Examined: Paratypes of S. ericia: AUSTRA- LIA; South Cape Wiles, 174-153 m, 35°39’ S, 136°40" E, AMS 032068, 1 left, 1 right valves (Zoological Results of the F.1.S. ENDEAvouR, 28 Aug. 1909). Other Material Examined: Holotype of S. japonica: JAPAN. ANSP 49639, 1 shell. MNHN. SW PACIFIC. Loyaute Islands, 16 lots [122 v]. TONGA IS. 12 lots [59 v|. GUAM. Marianas Islands, 3 lots [15 v]. AUSTRALIA. South Cape Wiles, 1 lot [6 v]. NEW CALEDONIA. South, 3 lots [7 specimens]; Banc Esponge, 2 lots [3 specimens]: Chee jield Plateau, 1 specimen. PHILIP- PINES. Aliguri Is. 2 lots [1 ete and 3 v|; Bohol Sea, Off Balicasag island 1 lot [1 v]. FHL. 1 specimen. MYANMAR (BURMA). 1 lot [5 v] ae: North Chan- nel, 1 lot [4 v]; N.W. of Tavoy L., 1 lot [11 v]. ANDA- MANS SEA. 1 lot [1 v]. THAILAND. Phuket I, 1 lot [11 v]; Andaman Sea, 1 lot [1 v| (Details in Simone and Cunha, in press.) Spinosipella costeminens (Poutiers, 1981) (Figures 60-65, 65-71, S3—92, 103-108) bl ay (Spinosipella) ) costeminens Poutiers, 1981: 351 (pl. . figs 144, text fig 5) Soacoe lla costeminens _Poutiers and Bernard, 1995: 110, 143, 158 (figs. 1-2). Diagnosis: Shell with 16-17 tall radial ribs, those more posterior to middle surface very taller, normally possessing blade-like projections along tip; 3-4 more posterior abruptly lower, preceded by a very tall, carina- like rib. Description: SHELL: Up to 30 mm. Color white. De- gree of convexity (width/length) in each valve approxi- mately 0.50. Outer surface prickly, with somewhat cha- otic organization (Figures 60, 61, 65-70). Sculptured by strong, uniform, arched, radial ribs, from 16 to 17 in each valve (Figures 60, 61); ribs increasing from region ante- rior to umbo to region between middle and posterior thirds, last ribs in this region taller and more separated from each other, last one on a weak carina (Figure 70); larger ribs normally possessing blade-like, projection L. R. L. Simone and C. M. Cunha, 2008 Page 71 A AW} % \ \ Wo SP gi mb ft pm Im Figures 83-86. Spinosipella costeminens. Anatomy. 83. Whole specimen with right valve extracted, right view. 84. Specimen extracted from shell, posterior view, showing siphonal area. 85. Whole right view, some portions of right mantle lobe extracted, particularly regions ventral to septum, and ventral and dorsal to renal fold (rf) to expose inner surface; cerebral ganglion (ce) seen by transparency. 86. Same, ventral-slightly right view. Scale bar = 5 mm. along tip; posterior third as a slope, having 3-4 ribs simi- umbonal region narrow, smooth about 10% of shell lar to those of anterior region; blade like projection ab- length (Figures 64, 65, 69). Anterior, ventral, and poste- sent in some specimens (Figures 68-71). Posterior edge rior edges forming zigzag (Figures 62, 63, 71, 103), with about twice broader than anterior edge. Between umbos tips longer and narrower projected in those middle and and anterior edge a concavity bearing transversal ribs larger ribs. Hinge with a large cardinal tooth in right similar to ribs of remaining region (Figures 64, 65): pre- valve, stubby, tall (about 10% of valve width), broadly Page 72 THE NAUTILUS, Vol. 122, No. 2 in Figures 87-90. Spinosipella costeminens. Anatomy. 87. Whole right view, mainly showing digestive tubes and main ganglia, topology of some structures also shown. Scale bar = 5 mm. 88. Scheme of layers of tissue in indicated region of stomach. Scale bar = 0.5 mm. 89. Fore- and midgut opened longitudinally for exposing inner surface (same scale of Figure 87). 90. Foot, ventral-slightly posterior view, sectioned transversally in two levels to show inner layer of tissues. Scale bar = 1 mm. pointed, somewhat flat (Figures 63, 71, 103); correspon- tionally shorter and wider (Figures 104-105). Length dent socket in left valve shallow, restrict to dorsal sur- about Ys to % of hinge length, and about 1.5 times aden face; this socket flanked by small posterior tooth, with and long. insertion of anterior valve edge approximately in middle Matin MUSCLE SYSTEM (FIGURES 83-87): Characters region of this tooth (Figure 62), anterior tooth absent similar to those in preceding species. Anterior adductor (Figure 62). muscle about 20% dorso-ventrally longer (Figures 83, 85). Additional details for this species see Poutiers (1981), Foor AND Byssus (FIGURES 85, 84, 90, 107): ): Shape Poutiers and Bernard (1995). and disposition similar to those in S. deshayesiana. Byssal LITHODESMA (FicuRES 103-105): Characters similar gland relatively deep, running immersed in ventral re- to those in preceding species, differing in being propor- gion of pedal musculature at about half of byssal furrow Figures 91-108. —Spinosipella species. Anatomy. 91. S. costeminens, middle horizontal, longitudinal section through visceral mass at same level as pericardium (MNHN sta. CP767, Mallory, 5 zm), Scale bar = 2 mm. 92. Same, detail of posterior region of mantle border. Scale bar = 1 mm. 93-102. S. deshayesiana. 93. Detail of hinge region of left valve with lithodesma (It) still attached, right view. Scale bar = 2 mim. 94. Detail of posterior region of supraseptal chamber, right view, right mantle lobe removed (MNHN sta. Oy 1499). Scale bar = Imm. 95. Infraseptal chamber roof, ventral view, right mantle lobe removed (MNHN sta. CP767). Scale bar 2 mm. 96-99. Lithodesma (MNHN sta. DW739). Scale bar = 1 min. 96. Ventral view. 97. Dorsal view. 98. Posterior-slightly dorsal view. 99. Posterior view. 100. Same specimen, empty shell, ventral view, valves slightly open, lithodesma still in situ. Scale bar = 2 mm. 101. Same, detail of hinge and lithodesma. 102. Same, ventral-slightly anterior view. 103-108. S. costeminens. 103. Shell, ventral view, valves open, lithodesma still attached to left valve (MNHN sta. CP1460). Scale bar = 2 mm. 104-105. Lithodesma, same ia other specimen), dorsal and ventral views respectively. Scale bar = 1 mm. 106. Detail of anterior region, right view, integument removed, mainly showing right cerebral ganglion (ce) (same lot). Scale bar = 1 mm. 107. Infraseptal chamber roof, ventral view, right mantle lobe removed (MNHN CP767). Scale bar = 1 mm. 108. Detail of posterior (siphonal) region, posterior view (MNHN CP1460). Scale bar = 2 mm Page 74 THE NAUTILUS, Vol. 122, No. 2 length towards dorsal (Figure 90, by). Thick muscular layer surrounding a nucleus of conective tissue (Figure 90, cj). : MANTLE (Ficures 84—S6, 92, 108): Characters similar to those in preceding species, with fo llowing distinctive characters. Pair of secondary tentacles positioned be- tween incurrent and excurrent siphons (Figures $4, 108); remaining tentacles similar in size and position. Ventral pair of feninele »s of incurrent siphon generally symmetri- cal. Zigzag formed by mantle edge having second: wy folds positioned in more distal tips, possibly elated to taller radial shell ribs (Figure 108). Radial mantle gland (Figure 92) similar to S. de shayesiana. PALLIAL Cavity (FiGuRES 85-86, 107): Characters similar to those in preceding species, except for wider platform between posterior region of gills as part of sep- tum (Figure 107). VISCERAL Mass (FIGURES 85—S7): Characters similar to those in preceding species, differing mainly by wider region separating pair of renal folds in supr: aseptal cham- ber (Figure $5). CIRCULATORY AND EXCRETORY SYSTEMS (FIGURES 55, 91): Pericardium and heart with characters similar to those in S. deshayesiana (Figure 91). Kidneys of similar features, differing mainly by enlargement of pair of renal folds (Figures 85— 86, rf), taller and wider, almost divid- ing supraseptal chamber in two—internal and external halves. Height of renal fold about S0% of that of su- praseptal chamber height. In addition to an enlargement, both renal folds still have posterior end in more anterior position and wider separation between folds and visceral mass (Figure 85). DIGESTIVE SYSTEM (FIGURES 87-89): Characters simi- lar to those in preceding species. Esophagus with about 3 of visceral mass length, running horizontally, perpen- dicular to posterior surface of anterior adductor muscle (Figure 87, es). Stomach main chamber with longer re- gion as a blind-sac projected posteriorly. Gastric wall constinited by external layer er of weak connective tissue Figure 109. Geographic distribution of Spinosipella spp + 8. aceulicostata #5. tinga S. deshavesiana ¥% S. agnes | (Figure 88, cj), two thick muscular layers of similar size, Wi ith outer layer of longitudinal muscle and inner layer of circular muscle (Figure 88, lo and cm). Imner surface of stomach (Figure 89) with posterior ead of esophageal folds clearly more evident that together form a flat fold. Another ventral fold surrounding apertures to digestive diverticula. Gastric style narrower (about Ys of gastric width); internally a pair of tall folds separating intestinal from sty le sac components (Figure 8S, ss, in). GENITAL SysTEM: Characters similar to those in pre- ceding species. Separated masculine and feminine com- ponents of gonad shown through histological sections in Figures 91(ts, ov). CENTRAL NERVOUS SYSTEM (FIGURES 87, LOG): Three ganglia with similar localization and size to those of pre- ceding species. Measurements (respectively length, height, width in mm): MNHN (Sta. 1361): 22.0 by 98.1 by 12:5 (valve); MNHIN (Sta. CC996): 20.0 by 24.3 by 14.3 (valve); MNHN (Sta. CP992): 19.6 by 23.3 by 12.6 (valve). Geographic Distribution: Tropical West Pacific. Depth Range: 750-925 m. Material Examined: Holotype; Additional material (MNHN): SW PACIFIC. 4 lots [32 v, 11 specimens]; Wallis Is., 6 lots [15 v]; Bane Combe, 5 Lots [28 v]; Fortuna Is., 5 lots [1S v|; Bane Waterwitch, 2 lots [3 v]; Bane Tuscarora, 29 lots [63 v]; South Vanuatu - Monts Gemini, 4 lots [4 v, 1 specimen]; TONGA IS. 8 lots [52 v|; Eua Is. 6 lots [12 v]; Seamount, 6 lots [29 vi South of Nomuka group, | lot [25 v|; Ha’apai Group, 2 lots [4 v]; N Ha’apai group, 3 lots { 6 v|; NW Tongatapu, 3 lots [16 v|; SW Tongatapu, 5 lots [22 v]; Tongatapu, 6 lots [8 v]; S. Nomuka group, 2 lots [6 v]; Vava’ group, 1 lot [2 v]; NEW CALEDONIA. 5 lots [5 v, 5 specimens]; Lord Howe, | lot [1 v]; Bane Nova, 2 lot [8 v, 1 specimen]; North New Caledonia, 10 lots [tota 20 v]; South New Indie Ocean @ S. costeminens L. R. L. Simone and C. M. Cunha, 2008 Page 75 Caledonia, 13 lots [46 v, | specimen]; off Norfolk, 1S lots [9S v]; Banc Esponge, 11 lots [144 v|; Bane Kaimon- Maru, 9 lots [3S v|; Bane Antigonia, 1 lot [1 v]; Banc Jumeau-West, 4 lots [17 v]; Bane Introuvable, 7 lots [16 v]; Bane Stylaster, 1 lot [1 v] ; Volcans Hunter and Mat- thew, 2 lots [2 v]; S.E. New Caledonia, 2 lots {2 v]; East New Caledonia, 6 lots [30 v] Banc Capel, 1 lot [lota 12 v}; Banc Kelso, 1 lot [6 v]; I. Loyaute, 22 lots [44 v]. FIJI. South of Viti Levu, 42 lots [328 v]; Southeast of Viti Levu, 17 lots [57 v|; Bohol/Sulu Seas, 2 lots [5 v|; Bohol Sea - Balicasag Island, 3 lots [5 v]; Bordau, 1 specimen; TAIWAN. Bashi channel, 2 lots [3 v]; South China Sea, 1 lot {2 v]; East Taiwan, 2 lots [5 v]. (Details in Simone and Cunha, in press.) DISCUSSION THE GENUS SPINOSIPELLA WITHIN THE VERTICORDHDAE. Despite their larger size, the prickly outer surface of the shell, and the reduction of the hinule, which differenti- ates Spinosipella from the remaining verticordiids, this taxon has traditionally been considere d a subgenus of the genus Verticordia. This set of characters is sufficient in my opinion to allocate Spinosipella as a separate genus. This view was previously defended by the author of the genus (Iredale, 1930) and by Poutiers and Bernard (1995). Other distinctive characters are the spiral um- bones (Figures 5, 7, 21, 22, 33, 54, 53), the tall, some- what uniform radial sculpture, triangular in section; and the obesity of the valves. The spiral umbones and the obesity of Spinosipella are quite similar to those in the fossil genus Pecchiolia Savi and Meneghini in Murchison, 1850 [type-species (by monotypy): Pecchiolia argentea Savi and Meneghini in Murchison, 1850 (= Chama ari- etina Brocchi, 1814) middle Tertiary, Europe] ( (Keen, 1969: 857), from which Spinosipella differs in having well-developed ribs and zigzag edges. The full genus status of Spinosipella is based on the differences with the typical Verticordia sensu stricto [tvpe species (by monotypy) Verticordia cardiiformis Sowerby, 1844], such as the higher size and obesity of the valves: the additional devel opment of the prickly surface (which also covers the radial ribs, whereas in Verticordia, when a prickly surface is present, it does not cover the Table i. Spinosipella Character acuticostata Spinosipella agnes radial ribs), the absence of lunule; the spiral fashion of both valves; and the similarity among the radial ribs (rep- resentatives of Verticordia usually have an unusually larger rib or space between ribs). The same set of char- acters also differentiates Spinosipella from Trigonulina dOrbigny, 1842 [type species (by monotypy) T. ornata d'Orbigny, 1842] in the sense of Jung (1996: 46-47). Repre sentatives of Spinosipella also resemble those of the genera Haliris Dall, 1856, and Euciroa Dall, 1881, by their larger size, convexity, and prickly shell surface. Spi- nosipella differs from those two genera, however, in the Rigier degree of convexity, reflected in more obese shells in its species; in the much more developed and taller radial ribs; higher degree of spiralization of the valves; and in the expansion of the ribs beyond the shell margin. Further analysis on the verticordiid systematics and phylogeny can be found in the literature (e. g., Pelseneer, 1888; Calvan: Plawén and Haszprunar, 1982: Bieler and Mikkelsen, 1992). COMPARISON BETWEEN THE SPINOSIPELLA SPECIES The differentiation between the five species of Spino- sipella is summarized in the respective diagnoses and in Table 1. The degree of differentiation in the samples of each species examined allows for specific separations. The number of radial ribs is the most notable feature; despite certain a small amount of intraspecific variation, the number of radial ribs is somewhat constant in each species, at least in specimens of larger size. The fossil S. acuticostata is the species with fewest ribs, 12-13 ( Fig- ures 36, 38, 39), while S. deshayesiana has the largest number of ribs, 16-19 (Figures 46, 48, 49, 54, 53). The other species possess an intermediary number of ribs. The species of Spinosipe la usually have radial ribs of relatively uniform size; the single exception is S. costem- inens, which has ribs clearly increasing posteriorly; in the posterior shell slope, however, the ane abruptly reduce in size, although in some specimens, particularly in the young ones, dhs character is not so clear, i.e., the ribs are saimewiat uniform-sized. The shell inflation is well de- veloped in most Spinosipella species, but this is clearer in Comparison of characters between the five studied species of Spinosipella. Spinosipella Spinosipella Spinosipella tinga deshayesiana costeminens Distribution Mediterranean Tropical W. Atlantic; Caribbean: to SE Brazil Shell Inflated Strongly Highly Sculptured between — Radial Disorganized radial ribs Prickly ribs outer Rough Rough surface Number of Ribs 12-13 15-17 Size (mm 20.0 20.2 South and Central Indo-Pacitic Strongly Radial S-SE Brazil Tropical West Pacific Weakly ; Highly Radial Disorganized Weakly prickly — Strongly prickly Rough 17-18 18-19 16-17 10.4 11.5 20.0 Page 76 THE NAUTILUS, Vol. 122, No. 2 the larger specimens; while the young specimens are éonsiderebly flatter (Figures 41-45). The prickly outer shell surface is an outstanding character of the Spino- sipella species; however, this character is conservative among the five species; the single exception is the rela- tively chaotic arrangement in S. agnes (Figure 8) and S. costeminens, while in the remaining species a radial ar- rangement is apparent (parallel to the radial ribs) (Figure 32). The Pacific species S. deshayesiana has much larger, crispy prickles along the tip of the ribs (Figures 42, 45, 46, 48, 49). This is lacking in the remaining species, except in some very young specimens (e.g.. USNM 810889, S. agnes, 6 mm), where the prickles, aware: are not fully dev eloped. The prickly surface is strongly damaged in eroded specimens (Figure 55), becoming ‘dies completely smooth. Spinosipella de shayesiana, perhaps because of this character, has the distal tips of the zigzag edges of the shell even longer and more pro- jected (Figures 41, 44, 47, 50, 51, 59, 67). The series of radial ribs is interrupted in the region between the um- bos, where a triangular smooth area appears. This area is particularly large in S. agnes (Figures 7, 9), but is prac- tically absent in S. tinga (Figures 2 1, 22): it is narrow in the remaining three species. The size of the specimens appears to be another distinctive feature, as S. tinga is small (around LO mm), whereas the remaining species are larger (20-30 mim). The hinge does not vary much between the Spinosipella species; however, some par- ticularities exist. The posterior tooth of the left valve is well developed in S. agnes [Figures 4, 10, 12,2 27, 28 (arrow)], very low in S. acuticostata (Figures 35, 39), ‘ae practically absent in remaining species iiieues 20, 25, 50). The tall and pointed car dinal tooth of the right valve is more developed in S. agnes, in such it is also sharply pointed and curved (Figures 3, 10, 12). In the remaining species this tooth is weak ly Beare and more founded (Figures 26, 34, 47, 51). The geographic and stratigraphic distribution are somewhat mutually exclusive for most of the species (Fig. 72); Spinosipe lla acuticostata is the only Mediter- ranean species, S. agnes occurs from Florida to Rio de Janeiro, S. tinga is found from Rio de Janeiro to Rio Grande do Sul, along the Brazilian coast. The fine- resolution distribution of the Indo-Pacific species is still unclear, but $. deshayesiana and S. costeminens, appear to be sympatric. Spinosipella acuticostata is a fossil spe- cies, occurring in Pliocene strata, while the remaining species are found in the Recent. Apparently no Recent Spinosipella occur in the Mediterranean. \ll samples of Spinosipella from the Atlantic and Mediterranean have previously been acce is ‘ot as belong- ing to the single species S. acuticostata (e. Abbott. 1974: Abbott and Dance, 1983: Rios, 1994). See: ane ieee »s of the conchologici il, geographic, and. strati- or phic differences, show that the separation into three spe c1es is warrante od, As the shi ape che mge Ss conside rably during ontogeny, a spec imen of S agnes at same size as the holotype of S. tinga was chosen to show the differ- ences between those species. Figures 25-31 illustrate these differences. Spinosipella agnes has fewer, taller, and more widely spaced ribs than S. tinga (Figures 19, 29). The shape of the shell edge is much more uniform in S. tinga than in S. agnes, in that the tips of the ribs are more expanded, extending longer beyond the shell mar- gin (Figures 20, 24, 28). The posterior cardinal tooth in the hinge of the left valve is present in S. agnes, in ani- mals larger than 5-6 mm, while this toath is never present in S. tinga (Figures 20, 25, 27-28, arrow). The degree of i rags is higher in S. agnes and in S. tinga (Figures 30, 31); S. agnes has a degree of convexity (width/lensth) ) in each ie of about 0.5 57, while S. tinga it is 0.47. The comparison of the previously valid species Spino- sipella ericia, including paratypes (Figures 41-44), and S. deshayesiana, does not reveal any distinction between them. Normally, specimens of sinaller size were identi- fied as S. ericia, and the large ones as S. deshayesiana. But examination of shell features along a growth series show a complete gradient linking the two taxa. The same lack of distinction is found in the literature for both spe- cies, including the original descriptions. For these rea- sons, despite the fact that S. ericia is the type species of the genus, the older name S. de shou should be ise Furthermore, a type specimen of S. japonica was also examined (Figure 45), confirming the synonymy of this species with S. de shayesiana. The distinction between the Pacific species Spino- sipella deshayesiana and S. costeminens is not always easy. With the large quantity of specimens kindly pro- vided by the MNHN (Paris), it was possible to analyze the degree of variation of both species. Spinosipella costeminens mostly has samples with shell possessing the outstandingly large, carina-like spiral ridge between the middle ea posterior thirds of the shell, "but sometimes this ridge is not so different from the others, and the animal become more rounded, similar to S. deshayesi- ana. The distinction is based mainly on the presence of at least a weak carina in the region between middle and posterior thirds, and also by the more robust ridges of S. costeminens specimens ( (Figures 60, 61), while those of S. deshayesiana lack any clear radial carina and the ridges are more delicate, uniform and apparently close fren each other (Figures 46, 48). The lot USNM 63200 includes 3 valves (2 left and 1 right), collected in Barbados, the known geographic dis- sibutiod of Spinosipella agnes. However the right valve has the characters of S. de shaye sianda, pictend of hese of S. agnes. In addition, is looks different in the state of conservation, color and associated sediment, from the other 2 valves of the same sample. DISCUSSION ON ANATOMY More in-depth anatomical descriptions and. discussions on verticordiids are provided by Allen and Turner L. R. L. Simone and C. M. Cunha, 2008 Page 77 (1974), who studied 19 species of several genera. How- ever, no information on the anatomy of the genus Spi- nosipella is found in the literature. Although anatomical information is available here only for two of the five species of the genus (of course one of them is a Pliocene fossil), some systematic inferences can be made based on the scenario given in the literature the Verticordiidae and related families (Allen and Turner, 1974, and others, e.g., Fisher, 1860, 1862b: Pelseneer, 1888; Nakazima, 1967; Allen and Morgan, 1981). Besides the conchological characters discussed above, some anatomical features are possibly restricted to Spinosipella, such as: the wide lith- odesma (Figures 93, 96—LO4, It); the simplified siphonal tentacles (Figure 10S), which normally have secondary papillae; the papill: von the root of the excurrent chamber (Figures 7S—SO, 85, 94: pi); the absence of incurrent valve in infraseptal chamber. However, wide lithodesma have been reported for Policordia lisbetae Knudsen, 1970 (fig. 90), which has very different shell and pallial tentacular characters. The study on the incurrent sipho- nal structures is o f particule uw import: imce in septibranchs, as the modified incurrent siphon is the main structure used in prey capture (Morton, 1987). On the other hand, some features appear to be char- acteristic of Verticordiidae, such as: elongation of lateral region of kidneys; the muscular stomach (see also Pur- chon, 1956, 1963): the separation between testis and ovary. By the proximity of the a daca from anterior adductor muscle, by the lack of incurrent valve, and by the simplified buc cal structures, e. ., lack of buccal cavity and tongue, it is possible to suggest that Spinosipella is a basal taxon inside Verticordiidae. Unfortunately, no member of the genus was analyzed in the recent com- parative studies on anomalodesmatans (Harper et al, 2006). ACKNOWLEDGMENTS The authors are grateful to the researchers who loaned the material for this study: Winston Ponder and Ian Loch (AMS) for types of Spinosipella ericia; Néstor E. Ardila (MHNMC) and Emilio Garcia (EGC) for S. agnes (Co- lombia) and S. deshayesiana (Philippines); Harry G. Lee for S. agnes (Florida) (MHNMC) for a large lot of S. agnes (Colombia): and especially to Philippe Bouchet and Philippe Maestrati, MNHN, for the loan of a huge quantity of lots coming from several places of the ere | mostly from the Indo-Pacific. For thorough comments and additional information about S. acuticostata we thank Rafael La Perna. For material of Haliris fisheriana we thank to Daniel Mansur Pimpao (PPG-BAN, UFRGS). For help with SEM procedures, we thank Lara Guimaraes (MZSP). For Rachel Collin, Smithsonian In- stitution at Panama, we thank for the oe in the text and language. We thank also both referees and the Editor for the thoughtful correction on the manuscript. T his project is supported by FAPESP (Fundagao de Amparo a Pes- quisa do Estado de Sao Paulo), procs. no. 03/05860-6, 04/02333-8, and a “Treinamento Técnico 3” grant, under the supervision of Antonia Cecilia Z. Amaral and Luiz R.L. Simone. LITERATURE CITED Abbott, R. T, 1974. American Seashells, second edition. Van Nostrand Reinhold Company. New York, 663 pp., 24 pls. Abbott, R.T. and S. P. Dance. 1983. Compendium of Sea- shells. E.P. Dutton, Inc. New York, 411 pp. Allen, J. A. and R. E. Morgan, 1981. The functional morphol- ogy of Atlantic deep water species of the families Cuspi- dariidae and Poromyidae (Bivalvia): an analysis of the evo- lution of the se ptibr: inch condition. Philosophical T rans- actions of the Royal Society of London (B) 294 (107: 413-546. Allen, J. A. and J. F. Tumer, 1974. On the functional anatomy of the family Verticordiidae (Bivalvia) with descriptions of new species from the abyssal Atlantic. Philosophical Transactions of the Royal Society of London (B) 268(S94): 401-536. : , Bieler, R. and P. M. Mikkelsen. 1992. Preliminary phylogenetic analysis of the bivalve family Galeommatidae. American Malacological Bulletin 9: 157-164. Costa, O. G. 1850. Paleontologia del Regno di Napoli. Atti della Accademia Pontaniana 5; 398. Crozier, M. A. 1966, New species and records of Mollusca from off Three Kings Islands, New Zealand. Transactions of the Royal Society of New Zealand, Zoology 8(5): 39-49. Fischer, P. H. 1860. Note sur les genres Hippa igus et Verticor- dia. Journal de ¢ onchyliologie 8 [(4) 4] (3): 295-300. Fisher, P. H. 1862a. Description d'une espece ae de Ver- ticordia. Journal de Conchyliologie 10: 35-36, pl. 5 Fischer, P. H. 1862b. Sur l'anatomie des Hinnites. Journal de Conchyliologie 10 [(3)2] (3): 205-217, pl. 11. Harper, E. M, H. Dreyer, and G. Steiner, 2006. Reconstructing the Anomalodesmata (Mollusca: Bivalvia): morphology and molecules. 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THE NAUTILUS 122(2):79-93, 2008 Page 79 Late Eocene Conus (Neogastropoda: Conidae) from Florida, USA Jonathan R. Hendricks Department of Geology University of Kansas Lawrence, KS 66045-7613 USA jrhendri@ku.edu Gainesville, F Roger W. Portell Division of Invertebrate Paleontology Florida Museum of Natural History University of Florida L 32611-7800 USA portell@fnmh.ufl.edu ABSTRACT The neogastropod genus Conus is likely the most diverse ma- rine animal genus, but has an Eocene to Pleistocene fossil record that remains poorly understood. We discuss the fossil record of Eocene Conus from Florida and recognize three species: Conus sauridens Conrad, 1833, and two new species, Conus palmerae and Conus alleni. We also re-describe C. sau- ridens, identify its likely synonyms, and add new information about its geographic range. The new species C. palmerae is restricted to the upper Eocene Ocala Limestone of Florida, while C. alleni new species occurs in the upper Eocene Ocala Limestone of Florida and the upper Eocene Moodys Branch Formation of Louisiana. Additional Keywords: Mollusca, Fossil, Limestone Gastropoda, Ocala INTRODUCTION With over 1500 described fossil and extant species, Co- nus Linnaeus, 1758 (cone snails) may be the most diverse marine animal genus (Réckel et al., 1995). Molecular sequence data have offered valuable insights into the relationships of extant Conus species (e.g., Duda and Kohn, 2005), but the deep evolutionary history of Conus remains unclear: although about 1000 fossil s species have been described ( Rocke! et al., 1995), many of these are likely synonymous (e.g., see Hendricks, in press). Many fossil Conus species are based only upon type specimens and/or were described without reference to comparable material from nearby, contemporaneous fossil faunas. Unraveling the early evolutionary history of Conus will only be possible by careful examination ‘and interpreta- tion of its rich and well- preserved fossil record. Some of the oldest Sabeuinl: ited Conus fossils are from the early Eocene (Ypresian) of England and France. and possibly Pakistan (Kohn, 1990). Occurrence records of middle Eocene Conus in North America show that the genus had achieved a wide geographic distribu- tion soon after its first appearance in the fossil record. For example, Table 1 (which is derived in large part from Kohn and Anderson, 2008) lists names that have been applied to Eocene Conus hon the southeastern United States. Our purpose is not to revise the systematics and describe the fossil record of all of these taxa, but rather to focus on those species occurring in Florida, which have thus far received only eae attention (Rich- ards and Palmer, 1953). This lack of attention is like ‘ly related to the preservational nature of these Floridian specimens, almost all of which are preserved as internal and/or external molds rather than shell material. While we focus on Conus fossils from the upper Eocene Ocala Limestone of Florida, we also consider likely synonyms of Conus sauridens Conrad, 1833, a widespread and tem- porally persistent taxon that will require additional future attention. GEOLOGICAL BACKGROUND Formally named by W. H. Dall (in Dall and Harris, 1892), the Ocala limestone represented all limestones exposed in central Florida at that time. A late Eocene age was not determined for the Ocala limestone, how- ever, until Cooke (1915) correlated the unit with deposits in Mississippi and Alabama. Based on lithology and micro- fauna, Applin and Applin (1944) divided the Ocala Lime- stone into an upper member and lower member. Vernon (1951) redefined the Ocala Limestone by restricting it to Applin and Applin’s (1944) upper member and placed their lover member in the Moodys Branch Formation. Further, Vernon (1951) subdivided the Moodys Branch Formation (as pertaining to Florida) into the basal Inglis Member and overlying Williston Member. Puri (1953) renamed Vernon's (1951) Ocala Limestone (restricted) as the Crystal River Formation. Later, Puri (1957) el- evated the Ocala Limestone to group status (Ocala Group)—consisting, from oldest to youngest, of the In- glis, Williston, and Crystal River formations—and elimi- mated the Moodys Branch Formation de ssignation for Florida Eocene de ‘posits. Base d on macro- and micro- fossils, Toulmin (1977: 117) correlated the Inglis For- mation with the lower Moodys Branch Formation in Page SO THE NAUTILUS, Vol. 122, No. 2 Table 1. Names applied to Eocene Conus from the U.S. Coastal Plain, including names of some younger species that are here considered synonyms of Eocene taxa. Type specimen abbreviations: H, holotype; L, lectotype; and F, figured. Taxon Type Type locality Present disposition according to this study sauridens Conrad, 1833 claibornensis I. Lea, 1833 gyratus Morton, 1834 parcus H. C. Lea, 1841 mutilatus Tuomey, 1852 tortilis Conrad, 1855 alveatus Conrad, 1865 agaaaaan C. subsauridens Conrad, 1865 C. pulcherrimus Heilprin, 1879 C. jacksonensis Meyer, 1885 C. deperditus var. subdiadema de Gregorio, 1890 C. improvidus de Gregorio, 1890 C. (Conospirus) granopsis de Gregorio, 1890 C. smithvillensis Harris, 1895 C. cormacki Harbison, 1944 C. (Leptoconus) santander Gardner, 1945 C. (Leptoconus) haighti Gardner, 1945 “Conus sp. A” Palmer in Richards and Palmer, 1953 “Conus sp. B” Palmer in Richards and Palmer, 1953 lant ). eracens Hoerle, 1976 C. alveatus spiralis Dockery in MacNeil and Dockery, 1954 C. (Lithoconus) smithvillensis var. Dockery, 1980 C. (Lithoconus) nocens Garvie, 1996 C. (Lithoconus) smithvillensis var. Dockery, 1980 in Campbell (1995) ANSP 14854 (L) Lost ANSP 211 (H) ANSP 13161 (L) None ANSP 13196 (H) ANSP. 13446 (L); See MacNeil and Dockery (1954, PI. 35, Fig. 26) ANSP 53812 (H) AMNH-FI 10175 (H?) Unknown, but specimen purported to be holotype figured by Harris and Palmer (1947, pl. 62, fig. 17) PRI 26436 (H) Lost (see Palmer and Brann, 1966) Lost (see Palmer and Brann, 1966) BEG 34656 (H) ANSP 16415 (AH) USNM 495181 (H) USNM 495182 (H) UF 108683 UF 108858 (designated here as holotype of C. palmerae, new species; formerly FL Geol. Survey I-7634) USNM 220109 (H) USNM 376678 (H) MGS 590 (F) UT-TMM 962TX22 (H) UNC 15445 (F) Claiborne, Alabama Claiborne, Alabama South Carolina, locality unknown Claiborne, Alabama Wilmington, North Carolina Jackson, Mississippi Vicksburg, Mississippi Probably Claiborne, Alabama Claiborne, Alabama Jackson, Mississippi Claiborne, Alabama Claiborne, Alabama Claiborne, Alabama Smithville, Texas Santee Cooper Canal, South Carolina Moseleys Ferry, Texas Arroyo Veleno, Texas Gulf Hammock, Florida Gulf Hammock, Florida Calhoun County, Florida Smith County, Mississippi Near Newton, Mississippi Bastrop County, Texas Near Cross, South Carolina C. sauridens C. sauridens? Nomen dubium C. sauridens Nomen dubium C. sauridens C. sauridens C. sauridens Eosurcula pulcherrimus (a turrid) C. sauridens C. sauridens Nomen dubium Nomen dubium C. smithvillensis Nomen dubium C. sauridens? C. haighti Unidentifiable C. palmerae, new species . sauridens? ag _ sauridens C. smithvillensis? C. sauridens? C. smithvillensis? Alabama, Mississippi, and Louisiana. He also correlated the Williston Formation with the upper Moodys Branch Formation in Alabama, Mississippi, and Louisiana. For detailed correlations, see the Correlation of Stratigraphic Units of North America—Gulf Coast Region (1988). Based on lithology, Scott (1991)—in order to follow the North American Stratigraphic Code (North Ameri- can Commission on Stratigraphic Nomenclature, 1983)—reduced the Ocala Group to formational rank and returned to the terminology used by Applin and \pplin (1944). Therefore, current designations used by the Florida Geological Survey and United States Geo- logical Survey are: lower member of the Ocala Lime- stone (formerly Inglis Formation) and upper member of the Ocala Limestone (formerly Williston and Crystal River formations). See Figure 1 for a brief history of stratigraphic divisions of the Ocala Limestone in Florida. In outcrop and shallow subsurface, the Ocala Lime- stone occurs in northwestern peninsular Florida and a small area of the Florida panhandle adjacent to Georgia and Alabama. Lithologically, the Ocala Limestone is a relatively pure carbonate. Non-carbonate minerals J. R. Hendricks and R. W. Portell, 2008 Page $] SERIES | STAGE |APP!P & Appling Vernon, 1954 1944 Puri, 1957 Scott, 1991 Upper Member Ocala Limestone (restricted) Crystal River Formation Upper Member Williston Member Ocala Group Williston Formation Z =< Z Oo WY < UO < = Ocala Limestone Ocala Limestone UPPER EOCENE Lower Member Moodys Branch Formation Inglis Member Inglis Formation Lower Member Figure 1. History of stratigraphic divisions of the Ocala Limestone in Florida (modified from Oyen and Portell, 2001). (quartz, chert, and clay minerals) represent less than 5% of the rock volume in the lower member and less than 10% in the upper member (Oyen, 1995). The lower member (formerly Inglis Formation) is primarily a clean packstone and grainstone that represents a higher en- ergy, subtidal environment while the upper tener (formerly Crystal River Formation) has a lithology of muddy pac -kstone and wackestone interpreted to signify a lower energy (below wave base), deeper subtidal deposit (Fenk, 1979). The Ocala Limestone contains diverse and abundant, shallow water marine invertebrate fossils consisting pri- marily of foraminifera, mollusks, and echinoids. Taxa with calcitic shells are preserved as body fossils, while taxa that had aragonitic shells typically occur as internal and external molds, although on rare occasions they can be found as omen of calcite or silica. Most of the calcitic shelled mollusks (e.g., oysters and scallops) and those found as pseudomorphs have been well docu- mented (Harris, 1951; Richards and Palmer, 1953). Few of the many moldic Ocala Limestone mollusks, however, have received the critical attention that they need (Por- tell and Vokes, 1997) MATERIALS AND METHODS Most of the specimens examined in this study are from the Florida Museum of Natural History (FLMNH) Di- vision of Invertebrate Paleontology at the University of Florida (UF) and locality details for individual UF speci- men lots listed below can be accessed online via the FLMNH Invertebrate Paleontology database at http:// www.flmnh.ufl.edu/invertpaleo/search.asp. Besides UF specimens, some type specimens from the Academy of Natural Sciences, Philadelphia (ANSP), the Texas Natu- ral Science Center at the Univ ersity of Texas ( (specimens carry the acronym TMM for Texas Memorial Museum), and the Texas Bureau of Economic Geology (BEG; specimens now at the Texas Natural Science Center) were also examined. Other specimens referred to in the text are from the Geological Survey of Alabama (GSA), the American Museum of Natural History (AMNH), the Mississippi Geological Survey (MGS), the Paleontologi- cal Research Institution (PRI), the University of North Carolina (UNC), and the United States National Mu- seum of Natural History (USNM), Morphospecies were recognized from museum speci- mens (primarily those at the FLMNH) using the criteria discussed by Smith (1930), Réckel et al. (1995), and Hendricks (in press) and morphological terminology and measurements collected follow those authors. Two terms are introduced here that relate to characters states of the subsutural flexure, which is “the backward curving or bending of the shell aperture below the suture of whorl contact” and “{bJeing a feature of the apertural margin it is strikingly manifest in the growth lines [on the sutural ramp] of well- preserved cones” (Smith, 1930: 284). These two terms are: symmetrically curved subsutural flexure and diagonal subsutural flexure. A symmetrically curved subsutural flexure (also see the more generalized “curved type” described by Smith, 1930) has a maximum point of curvature centered between the margins of the sutural ramp. A diagonal subsutural flexure forms a straight or eee straight | ine that crosses diagonally (abaxioventrally) across the sutural ramp. Also see Muniz-Solis (1999) for illustrations of different subsu- tural flexure morphologies. Measurements were collected using digital and dial calipers and include: shell length (SL), maximum diam- eter (MD), aperture height (AH), height of maxi- mum diameter (HMD), and spire angle (SA). From these measurements, the following morphometric ratios (developed by Réckel et al., 1995) were computed to characterize shell sh: ape: relative diameter (RD; RD = MD / AH); position of maximum diameter (PMD; PMD = HMD / AH); and relative spire height (RSH; RSH = [SL - AH]/SL). Recognized Floridian morpho- species were compared with previously described species of Eocene Conus from the southeastern United States. Although the simple internal molds of Conzs fossils offer little taxonomically relevant information, room tem- perature vulcanizing (R.T.V.) silicone rubber casts made from external molds are often very useful for recognizing species. The external molds discussed herein were first gently cleaned to remove loose debris and then impreg- nated with polyvinyl butyral (Butvar-76), if needed, to consolidate the limestone. Once dried, clay dams were constructed around the outside of the limestone blocks containing the external molds. Then, de-aerated R.T.V. silicone rubber was slowly poured into and above the molds and again de-aerated in a vacuum chamber. Later, the cured rubber casts were carefully lifted from the molds. For more information regarding this technique Page 82 THE NAUTILUS, Vol. 122, No. 2 see Chaney (1989) or visit http://paleo.cc/casting/ silsum.htin. SYSTEMATICS Our examination of Eocene Conus fossils from Florida resulted in the recognition of one distinctive morpho- species that was first described by Conrad (1833) as C. sauridens, and two new species (C. palmerae and C. alleni) which are described below. Family Conidae Fleming, 1822 Genus Conus Linnaeus, 1758 Conus sauridens Conrad, 1833 (Figures 2-10, 15-22; Table 2) Conus sauridens Conrad, 1833: p. 33; Conrad, 1835: pl. 15, fig. 7. For comprehensive synonymy listings, see Palmer (1937), Harris and Palmer (1947), and Palmer and Brann (1966). ?Conus claibornensis I. Lea, 1833: 186. Conus parvus H. C. Lea, 1841: 103, pl. 1, fig. 24. Conus tortilis Conrad 1855: 260, pl. 15, fig. 5. Conus alveatus Conrad, 1865: 148, pl. 11, fig. 4. Conus subsauridens Conrad, 1865: 148, pl. 11, fig. 9. Conus jacksonensis Meyer, 1585: 466. Conus deperditus var. subdiadema de Gregorio, 1890: 20, pl. 1, figs. 56-58. Conus (Leptoconus) santander Gardner, 1945: figs. 5, 9, 10, 14. ?Conus cracens Hoerle, 1976: 14, 16, pl. 3, figs. 1-3. Conus alveatus spiralis Dockery in Mz acNeil and Dockery, 1984: 165, pl. 59, figs. 3, 4. 2Conus nocens Garvie, 1996: 90, pl. 19, figs. 9, 10. 251, pl. 26, Diagnosis: Early postnuclear whorls tuberculate, later teleoconch whorls smooth; shell often widest below shoulder; shoulder typically ridge-like; sutural ramps with raised spiral threads; incised spiral grooves at base of shell. Description: Shell medium to large-sized (up to 116 mm). Last whorl conical to broadly and ventricosely coni- cal; outline typically convex near shoulder, nearly straight below. Shell often widest below shoulder. Shoulder ty pi- cally broadly carinate and forming pronounced ridge, less often angulate: smooth. Spire of low to moderate he ight, spire angle ( SA) typically obtuse; outline typically straight to concave in smaller specimens, usually sigmoid in larger shells. Larval shell multispiral, with at Teatt three whorls. Early postnuclear whorls tuberculate. Subsutural flexure symmetrically curved, depth about 2.5x width. Teleoconch sutural ramps typically sigmoidal (rarely flat or convex), with raised spiral threads (typically three to seven; deine ‘ry increases with shell size) and interve ning grooves; ornamentation begins on earliest postnuclear whorls as a single incised spiral groove. Aperture opening of approximately uniform width from base to shoulder. Some large specimens bear a pronounced siphonal fas- ciole. Last whorl with incised spiral grooves. at base, sometimes extending weakly to near center of whorl and occasionally to shoulder in small shells; spiral grooves obsolete in some large shells. Shell Morphometrics: Thirteen shells of type and non-type specimens of C. sauridens were measured and morphometric ratios were computed from these mea- surements (Table 2): relative diameter (RD) ranges from 0.57-0.74 (average = 0.67); position of maximum diam- eter (PMD) ranges from 0.83-0.95 (average = 0.91); and relative spire height (RSH) ranges from 0. 090.21 (aver- age = 0.17 ). Type Specimens: Type specimens examined include: ee 14854, lectotype of Conus sauridens (Figures 2, ; ANSP 535813, three paralectotypes of C. eee ica 4-6); ANSP 53812, holotype of C. subsauridens (Figure 7); ANSP 13161, holotype of C. parvus (Figure 8); and ANSP 13196, holotype of C. tortilis (Figures 9, 10). See Table 2 for measurements of type specimens. Type Locality and Occurrence: Conrad (1833) de- scribed the species from specimens collected at Clai- borne, Alabama. Palmer (1937: 10) designated the type locality and stratum of Conus sauridens as PRI station 104: “ ‘Ferruginous sand’ bed at Claiborne, on the Ala- bama River, Monroe County, Ala. Gosport sand” (Palmer, 1937: 10). The Gosport Sand is the uppermost formation of the Claiborne Group and is late middle Eocene (Bartonian) in age (Dockery, 1980). This species has been previously reported from numerous Paleogene U.S. Gulf Coast strata, including the middle Eocene Claiborne Group, as C. tortilis in the upper Eocene Jackson Group, and as C. alveatus in the lower Oli- gocene Vicksburg Group (the reader is directed to the follow ing sources for detailed discussions of specific oc- currence records: Palmer, 1937; Harris and Palmer, 1947; Palmer and Brann, 1966; and MacNeil and Dock- ery, 1984). Conus sauridens also occurs in the upper Eocene Ocala Limestone of Alachua, Suwannee, and Jackson counties, Florida. This species also questionably occurs as C. cracens in the lower Miocene Chipola For- mation of northern Florida. Other Material Examined: In addition to the type specimen lots listed above, 25 specimen lots containing C. emer ns we re examined se r SO specimens). ). These UF ee UF 16661, U F 18874, UF 57713, UF 57733, UF 100693, UF 101995, UF 114376, UF 115875, UF 119912, UF 119913, UF 119918, UF 119962, UF 120021-UF 120027, UF 120032, UF 122384, and UF 126927. Discussion: Conus sawridens was the third fossil Co- nus species to be described from North America, and the first from the Paleogene (Green described the Neogene species C. deluvianus and C. marylandicus in 1830; see Kohn, 1992). As such, this species has received much attention in the literature, particularly with regards to its J. R. Hendricks and R. W. Portell, 2008 Figures 2-18. Specimens of Conus sauridens Conrad, 1833 (2-10, 15-18), C. cracens Hoerle, 1976 (11, 12), and C. nocens Garvie, 1996 (13, 14). See text and Table 1 for locality information. All scale bars equal 1 cm. Scale bar above Figure 5 pertains to Figures 2-7 and 11-14. Scale bar below Figure 8 pertains only to that figure. Scale bar between Figures 9 and 10 pertains to Figures 1 15-18. 2-3. Lectotype (ANSP 14854) of C sauridens, shell length 33.4 mm, maximum diameter 21.1 mm. 4. Paralectotype ANSP 53813-3) of C. sawridens, shell length 31.9 mm. 5. Paralectotype (ANSP 53813-1) of C. sauwridens, maximum diameter 22.3 mm. 6. Paralec totype (ANSP 53813-2) of C sauridens, shell length 30.2 mm. 7. Holotype (ANSP 53812) of C subsauridens Conrad 1865, shell length 33.4 mm. 8. Holotype (ANSP 13161) of C. parvus Lea, 1841, shell length 6 mm, 9-10. Holotype (ANSP 13196 of C. tortilis Conrad, 1855, shell length 90.0 mm, maximum diameter 53.0 mm, 11-12. Paratype (UF 119560-1) of C. cracens Hoerk 1976, shell length 32.2 mm, maximum diameter 17.8 mm. 13-14. Holotype (TMM-962TX22) of C. nocens Garvie, 1996, shell length 19.2 mm, maximum diameter 8.7 mm. 15. R.T.V. silicone rubber cast from external mold of C. sauridens (UF 120026). 16. R.T.\ silicone rubber cast from external mold of C. sauridens (UF 122384). 17-18. Highly leached, heavily pitted, and slightly silicified shell of C. sauridens (UF 120027), shell length 27.4 mm, maximum diameter 16.5 mm Page 84 THE NAUTILUS, Vol. 122, No. 2 Table 2. Measurements (in mm) and morphometric ratios of type and non-type specimens examined. Some measurements could not be collected from casts made from molds; other measurements (and associated ratios) are not accurate because of shell damage (indicated by an asterisk, °). Morphological abbreviations: SL, shell length; MD, maximum diameter; AH AH, aperture height; HMD, height of maximum diameter; SA, spire angle (in degrees); RD, relative diameter; PMD, position of maximum diameter; and RSH, relative spire height. Specimen abbreviations: H, holotype; L, lectotype; P, paratype; and PL, paralectotype. Specimen SL MD AH HMD SA RD PMD RSH Type Specimens of Conus sauridens Conrad, 1833 ANSP 14854 (L, C. sauridens, Figures 2, 3) ANSP 53813- : (PL, C. sauridens, Figure ! 5) ANSP 53813-2 (PL, C. sauridens, Figure 6) ANSP 53813- 3 (PL, C. sauridens, Figure 4) ANSP 53812 (H, C. subsauridens, Figure 7) ANSP 13161 (H, C. parvus, Figure §) ANSP 13196 (H, C. tortilis, Figures 9, 10) Non-Type Specimens of Conus sauridens Conrad, 1833 UF 283 UF 290-1 UF 8511-1 UF S647 UF 16726 UF 115875 UF 120022 Conus palmerae new species UF 108858 (HL, C. palmerae, new species, Figures 23, 24) UF 15586 (P) UF 18599 (P, Figure 25) UF 18711 ( UF 18719 ( UF 18737 (P, Figure 26) ( ( > ( a 7 57018 UF 74473 3 (P, Figures 29, 30) UF 110360 (P) — UF 111327 (P) UF 112981 (P) Conus alleni new species UF 119920 CH, Figures 31, UF 119919 (P, Figure 34) UF 119976 (P, Figure 35) UF 119977 (P, Figure 33) Other Type Specimens ANSP 16145 (H, C. cormacki, Figure 39) UF 119560-1 (P, C. cracens, Figures 11, 12) UF 76798-1 (P, C. cracens) TMM-BEG 35656 (H, C. smithvillensis, Figures 36-38) TMM-962TX22 (H, C. nocens, Figures 13, 14) TMM-9627TX23 (P, C. nocens) 43.6 30.2° 31.9° 33.4° 48.83 27.38 40.5 41.44 24.3 33.1 22.95 12.36 19.5 18.0 110 0.63 0.92 0.15 33.45 19.86 28.4 25.7 119 0.70 0.90 0.15 116.11 69.15 95.2 84° 121 0.73 0.88° 0.18 22.0° 11.5° 146° 13.7° 77 — 5 00.1° 12.0° 24.9° 56.1° Wwwww C C2 DD UU GO 21.1 30.5 28.9 145 0.70 0.95 0.09 22.3° 36.5 33.6 106 0.61 0.92 0.16 15.0 26.2 24.5 110 0.57 0.94 0.13° 16.7 25.9 23.5 101 0.64 0.91 0.19° 17.3 28.2 26.2 115 0.61 0.93 0.16° 90.9 53.0 75.1 62.1 100° 0.71 0.83 0.17 37.59 22.4 30,3 27. 110 0.74 0.91 0.19 24.43 1187 19.3 a 90 0.62 0.90 0.21 109 0.68 0.58 0.17 109 0.73 0.92 0.20 wwe on ARwn 22.3° — _— 103 21.5° — 14.2° — — 101 — 19.3 — —_ 9S os 19.0° — 9S 65.0% 284° 53,8° 11.3 a 10S — 12.4 — — LOS 2 20.7° = 31.3 28.6 121 0.66" 0.91 0.14 2° = =17.3° — — 92 Oo” L7.2° 1° 16.8° — — 112° ue 22.3 2 17.8 27.5 24.0 113 0.65 0.87 0.15 0 37.3° 145° 28.6 26.9° 53 0.52° 0.94° 0.32 2 8.7° 15.1 14.3 S7 0.58° 0.95 0.21 19.4 8.7° 15.4 14.5 ST 0.56 0.94 0.2] morphological variation and probable synonyms. Much of the following was derived from discussions in Palmer (1937), Harris and Palmer (1947), and Palmer and Brann (1966). Timothy Conrad (1533) described C. sauridens from shells that he collected at Claiborne, Alabama. According to Palmer (1937: 461), the sauridens consists of 5 specimens” which were appar- ently glued to one card, cataloged as ANSP 14854. Palmer (1937: 461) goes on to state that the “Conradian collection of “type of Conus subsauridens Con. |see below] is also on the card with the Conus sauridens collection.” These two passages suggest that Conrad’s Claiborne collection originally con- siste ae of six specimens. The collection now consists of five specimens: the lectotype (ANSP 14854; Figures 2, 3) of C. sauridens, three paralectotypes (ANSP 53813: Fig- ures 4-6), and the holotype of C. subsauridens ( ANSP S12; Figure 7); the location of any possible er speci- men is not iano One low- =e ‘d specimen in the type series (ANSP 14554; Figure 2) closely resembles Con- J. R. Hendricks and R. W. Portell, 2008 Page 85 rad’s (1835, pl. 15, fig. 7) original figure of C. sawridens. cave (1937) alluded to this specimen in her text as iets Conrad’s figure, but did not formally designate it as the lectotype. Kohn (1992)— following Moore (1962) and Palmer and a (1966)—considered this tae, to be the lectotype and we accept his conclu- sion. Palmer (1937) argued that the matrix filling the aper- tures of the shells in Conrad’s Conus sawuridens series suggests that the shells do not likely share the same geo- logical provenance. The matrix filling the lectotype is orange and appears to be sand from the Gosport For- mation (Palmer, 1937). The matrix filling the three para- lectotypes is light-gray. The matrix f filling the holotype of C. subsauridens is orange, resembling that of the lecto- type of C. sauridens (Palmer [1937: : 461] described the matrix filling the holotype of C. subsauridens as “a red, silicified matrix resembling that of the Orangeburg” material from South Carolina). It is important to note, however, that these differences in matrix color may be due to diagenetic weathering of the original matrix ma- terial. The lectotype (ANSP 14854; Figures 2,3) of C. sau- ridens differs from the paralectotypes (ANSP 53813; Fig- ures 4-6)—which, with the holotype of C. subsauridens (see below), are of the much more common morphol- ogy—in the following respects: it has flat to slightly con- vex sutural ramps with more raised spiral threads (about 6) than is typical, the shoulder is angulate rather than forming a carinate ridge, the widest point of the last whorl is nearly at the shoulder rather than beneath it, and the spire is lower than in most other specimens of the species (see Table 2); further, the lectotype does not provide definite evidence of tubercles on the early post- nuclear whorls, though these are highly eroded on this specimen, Isaac Lea (1833) described Conus claibornensis from materials sent to him by Judge Charles Tait of Claiborne, Alabama, but lost his only specimen before it could be figured ( Kohn, 1992). His description suggests that his specimen was a C. sauridens, a name that has priority by three months (Kohn, 1992). Harris (1895), Palmer (1937), and Palmer and Brann (1966) synonymized C. claibornensis with C. sauridens, while de Gregorio (1890), Dall (1896), and Kohn (1992) regarded this taxon as a nomen dubium. Given that no available evidence suggests that more than one fossil Conus species is present at Claiborne. Alabama, we agree with the former authors that C. claibornensis is equivalent, while ques- tionably. to C. sauridens. For a historical overview of interactions between I. Lea, T. A. Conrad, and C. Tait, see Wheeler (1935). In 1841. H. C. Lea (I. Lea’s father) described Conus parcvus on the basis of a single, small, damaged shell ANSP 13161; Figure §) from the Gosport sand at Clai- borne, Alabama. We agree with Dall (1896), Palmer (1937). and Palmer and Brann (1966) that C. parvus is a juvenile C. sauridens. Features uniting the holotype of C. parvus with C. sauridens include: tuberculate early post- nuclear whorls, raised spiral threads on the sutural ramps, and incised spiral grooves on the anterior half of the last whorl. Conrad (1855) described Conus tortilis from one large specimen (ANSP 13196; Figures 9, 10) from Jackson, Mississippi and differentiated it from C. sauridens by its ‘more prominent and convex spire, in the large twisted callus at base, & c.” (p. 260). Dall (1896) and Palmer (1937) both considered C. tortilis synonymous with C sauridens and Palmer (1937) described C. tortilis as rep- resenting “the maximum growth of the species” (p. 459). We agree with these authors that specimens of C. tortilis are large C. sauridens. Conrad (1565) described two additional species of Eocene Conus: C. subsauridens and C. alveatus. Conrad stated that C. subsauridens was from “the Burrstone, probably, of Alabama”; we assume (see above) that the holotype (ANSP 535812) is from Clairborne. We in- spected the type of C. subsauridens (Figure 7) and agree with Dall (1896) and Palmer (1937) that it is a junior synonym of C. sauridens. The type locality for Conus alveatus is Vicksburg, Mis- sissippi; MacNeil and Dockery (1984) suggested that the lectotype (ANSP 13446; MacNeil and Dockery, 1954, pl. 38, fig. 26) and paratype (ANSP 13494) are probably from the Byram Formation. Conrad (1865: 148) differ- entiated C. alveatus from C. sauridens by its “less el- evated and... more profoundly carinated spire, and the revolving lines on the spire are less numerous than in the former [C. sauridens]|.” Dall (1896) and Palmer (1937) recognized Conus tortilis and C. alveatus, respectively, as the Jacksonian (upper Eocene) and Vicksburgian (lower Oligocene) forms of the older, Clairbornian (middle Eocene), C. sauridens. MacNeil and Dockery (1954), however, continued to recognize C. alveatus as a distinct Oligocene species occurring in Mississippi and Mexico; they did not compare C. alveatus (or Dockery’s subspecies C. alveatus spiralis; in MacNeil and ee kery, 1984) with C. sauridens. MacNeil and Dockery’s 1984) figures of C. aleveatus (including the lectotype: al 35 fig. 26) appear consistent with C. sauridens as circum- scribed here. Meyer (1885) described—but did not figure—Conws jac eee from Jackson, Mississippi, and de scribed the species as similar to Conus protracta Meyer, 1S85 (an Oligocene taxon from Vicksburg and Red Bluff, Missis- sippi that we accept, but do not consider further here: see MacNeil and Dockery, 1954 for details), but “with revolving lines on the spire” (p. 466). Meyer [1886] pre- sented C. protracta as C. protractus and the latter spelling is the one most commonly seen in the literature. Harris and Palmer (1947: pl. 62, fig. 17) figured a speci- men (unnumbered, but said to be from the collections of the Geology Department of Johns Hopkins University) that they regarde das the holotype of C. jac ksonensis. This specimen “consists of the apical whorls” and is 3.5 mm in size (Harris and Palmer, 1947: 446). Harris and Page 86 THE NAUTILUS, Vol. 122, No. 2 Palmer (1947) considered this taxon a junior synonym of C. sauridens and we agree, particularly because of the presence of vubensulater early postnuclear whorls and the presence of raised spiral threads on the sutural ramp. De Gregorio (1890) described Conus deperditus var. subdiadema from Claiborne, Alabama. Palmer and Brann (1966) considered this subspecies equivalent to C. sauridens and, based upon our inspection of de Grego- rio’s (1890) figures, we agree. We have not, however, viewed the holotype of C. deperditus var. subdiadema, which Palmer and Brann reported as PRI 26436. Gardner (1945: 252) described Conus santander as including “those species from the western Gulf that have formerly been included under Conus sauwridens Conrad, described from Claiborne.” Gardner's figures of the ho- lotype (USNM 495181; Moseleys Ferry, Burleson County, Texas) of C. santander appear consistent with C, sauridens and we consider this taxon a probable syn- onym. Hoerle (1976) described Conus cracens (see paratype in UF 119560, Figures 11, 12) from the lower Miocene Chipola Formation of northern Florida and noted its strong similarity to C. sauridens: “C. cracens appears to be a descendant of the v widespread (Alabama, Missis- sippi, Texas) ) middle Eocene. to Oligocene species, C. sauridens Conrad” (p. 16). She differentiated C. cracens from C. sauridens on the basis of several characters: the “nodes on the spire whorls persist for a greater num- ber of tums on C. cracens, also it is larger, more slender, with stronger and more opisthocyrt growth lines and more pronounced basal ornament” (p. 16). We examined Hoerle’s paratypes at the FLMNH and could not find any discrete morphological characters separating the two taxa. Given the vast amount of geological time separating the species (over 12 million years), however, we ques- tionably synonymize C. cracens with C. sauridens. Re- solving the relationship between these two taxa will re- quire additional study. Finally, Garvie (1996) described Conus nocens on the basis of two small specimens from the Reklaw Formation of Bz istrop County, Texas: TMM-962TX22 ( (holotype; er- roneously publishe das UT-TMM 84822; Figures 13, 14) and TMM-962TX23 (paratype, erroneously published as UT-TMM 54823). In addition to these two specimens, Garvie (1996) reported that he examined 25 additional specimens of C. nocens from the Weches Formation and over 100 specimens from the Cook Mountain Formation. While we did not observe these additional, stratigraphi- cally younger specimens, the holotype and paratype of C. nocens appear consistent with Conus sauridens. They each have raised spiral threads on the sutural ramp, sym- metrically curved subsutural flexures, and incised spiral srooves on the anterior half of the last whorl. Further, both have last whorl shapes similar to C. sauridens. The early postnuclear whorls of the holotype are tuberculate, but most appear smooth on the paratype. Garvie (1996: 90) stated that four characters separate C. nocens from C. sauridens: “the flat sides, the sharp unridged carina, the coarse, strong, spiral basal lines, and the lack of strong growth lines on the ramp.” We do not consider these qnrapteristies=an isolation or combination— sufficient to discriminate C. nocens from C. sawridens (as circumscribed above), especially because TMM-962TX22 and TMM-962TX23 are both shells of juveniles. Since, however, we have not seen the other specimens of C. nocens mentioned by Garvie (1996), we consider our synonymy of C. nocens with C. sauridens tentative. Re- gardless of their taxonomic identity, TMM-962TX22 and TMM-962TX23 are important specimens because of their likely stratigraphic position in the Reklaw Forma- tion. The Reklaw Formation is thought to span the early- middle Eocene (or, Ypresian- Lutetian) boundary and has a relative age equivalent to nannoplankton biochrono- zone NP14 (Zachos and Molineux, 2003: fig. 2), which has an absolute age of about 49.7 to 47.3 Ma (Berggren and Pearson, 2005). This age would make these two specimens the oldest known Conus fossils from the United States Coastal Plain, and only slightly younger than the oldest Conus in general ( (Ypresian of England and France; Kohn, 1990). This oldest regional occur- rence record is tempered, however, by the fact that the position of the original collection locality of TMM- 962TX22 and TMM-962TX23 (“Devil’s Eye, Colorado R.”; station 11 of the Geological Survey of Texas and locality 11-T-36 of the Texas Bureau of Economic Geol- ogy) is uncertain (Zachos et al., 2005) and may no longer exist (Garvie, 1996). Further, no additional specimens of Conus from the Reklaw Formation have yet been found in the collections of the Texas Natural Science Center (A. Molineux, personal communication to JRH, July 11, 2007). Most of our knowledge of Eocene C. sauridens in Florida is from R.T.V. silicone rubber casts of external molds in limestone (e.g., UF 120026, Figure 15; UF 122354, Figure 16), though UF 120027 (Figures 17, 18)—which is a highly leached, heavily pitted, and slightly silicified dieil-ag an exception, Conus sauridens is present in upper Eocene Ocala Limestone of Jackson (UF 18874, UF locality JA002; UF 120026, UF locality a UF 119912. UF locality JA027 - and UF 119918, UF locality JA031), Alachua (UF 120027. UF locality ALOOL), and Suwannee (UF 122384 and UF 120032, both UF locality SU003) counties. F inally, one additional specimen (UF 119913) that may be C. sauridens is from the ‘aa Oligocene Bumpnose Uiraestone of Jackson County (UF locality JA025). Besides these records, the only pe known record of this taxon (as recognized here) in Florida is Dall’s (1916: 4489) account of C. tortilis in the “Ocala.” Conus sauridens co-occurs in the Eocene of Florida with the new species C. palmerae and C. alleni; characteristics that distinguish the new species from C. sauridens are discussed below. A complete review of the fossil record of Conus sau- ridens is beyond the scope of this paper, though the preliminary observations we have made here support earlier demonstrations (Palmer, 1937) that C. sauwridens J. R. Hendricks and R. W. Portell, 2008 was a morphologically variable, geographically wide- spread, and temporally persistent species. While the old- est known (early Eocene) Conus fossils were small (<35 mm in shell length), larger species (ca. 70 mm in shell length) have previously been reported from the middle Eocene (Kohn, 1990). During this study, we recognized a very large (shell length, 116.1 mm) specimen (UF 120022: Figure 19) of C Eocene Moodys Branch Formation of Grant Parish, Louisiana (UF locality ZLO04). This specimen may be the largest Conus yet known from the Eocene of the U.S. Coastal Plain. The large geographic range of C. sauridens sauridens from the upper could be related to its developmental mode, which was likely planktotrophic based upon its multispiral larval shell (Figures 20-22), though testing this hypothesis within the context of Shuto’s (1974) model of the rela- tionship between developmental mode and larval shell morphology (also see Kohn and Perron, 1994) will re- quire additional study. While not necessarily useful as a cuide fossil, shells of C. sauwridens (as circumscribed here) have recently been utilized in several isotopic stud- ies (Kobashi et al., 2001; Kobashi and Grossman, 2003; Kobashi et al., 2004) of Paleogene climate and have proven to be geologically useful in this regard. Conus palm rae new species Figures 23-30, Table 2) Conus sp. B, Palmer in Richards and Palmer, 1953: 40, pl. 2, hig. 14 Diagnosis: Teleoconch whorls stepped; early post- nuclear whorls smooth; sutural ramps typically smooth: last whorl smooth. Description: Shell small to moderately large-sized (up to about 65 mm in length). Last whorl conical: outline slightly concave. Shoulder sharply angulate, smooth. Spire of moderate height; outline concave to straight. Teleoconch whorls stepped; spire angle of early whorls typically obtuse relative to later whorls. Larval shell un- known. Early postnuclear whorls smooth. Subsutural flexure symmetrically curved. Teleoconch sutural ramps concave and typically smooth, though occasionally 2or3 weak spiral threads are present. Aperture morphology unknown. Last whorl typically smooth, though fine spiral lines May cover the last whorl ot some small specimens. Type Specimens: Holotype UF 105855 (Figures 23, 24). a specimen originally described as “Conus sp. B” by Palmer in Richards and Palmer (1953: 40, pl. 2, fig. 14). The holotype is preserved as a calcite-replaced shell. All yaratypes are moldic (consisting of just external or exter- ial and internal molds) and include: UF 15886, UF 18599 (Figure 25). UF 18711, UF 18719, UF 18737 Figure 26), UF 57018, UF 66735 (Figure 27), UF 68306 Figure 28), UF 74473 (Figures 29, 30), UF 110360, UF 111327. and UF 112981. See Table 2 for measurements of these specimens 2 7 Figures 19-22. Specimens of Conus sauridens Conrad 1833. 19. Largest known specimen (UF 120022) of C. sau- ridens, shell length 116.1 mm, Moodys Branch Formation, UF locality ZLO04 (Montgomery Landing 01), Grant Parish, Loui- siana; scale bar equals 1 cm. 20. Juvenile shell (UF 126927 shell length 3.4 mm, Moodys Branch Formation, UF locality ZLOO04 (Montgomery Landing 01), Grant Parish, Louisiana scale bar equals 1 mm. 21-22. Shell (GSA.2007.005), shell length 60.5 mm, Moodys Branch Formation, Montgomery Landing, Grant Parish, Louisiana; Figure 21 shows the larval shell and early postnuclear whorls of this specimen, which is also shown in Figure 22 (both scale bars equal 1 cm Page 88 THE NAUTILUS, Vol. 122, No. 2 Figures 23-30. Specimens of Conus palmerae new species. All scale bars equal 1 cm. 23-24. Holotype (UF 108858), preserved shell length 22.0 mm, preserved maximum diameter 11.5 mm, lower member of the Ocala Limestone (formerly Inglis Formation), UF locality LVO14 (Gulf Hammock 02), Levy County, Florida. 25. R.T.V. silicone rubber cast of paratype (UF 18599), preserved shell length 50.1 mm, preserved maximum diameter 23.4 mm, Ocala Limestone, UF locality ALO16 (S.M. Wall Quarry 01), Alachua County, Florida. 26. R.T.V. silicone rubber cast of paratype (UF 18737), preserved shell length 56.1 mm, preserved maximum diameter 24.1 mm, Ocala Limestone, UF locality LFOO1 (Dell Limerock Mine), Lafayette County, Florida. 27. R.T.V. silicone rubber cast of paratype, (UF 66738), maximum diameter 14.2 mm, Ocala Limestone, UF locality ALO28 (Newberry 03), Alachua County, Florida. 28. R.T.V. silicone rubber cast of paratype (UF 68306), maximum diameter 19.3 mm, Ocala Limestone, UF locality ALO04 Dickerson Limerock Mines), Alachua County, Florida. 29-30. R.T.V. silicone rubber cast of paratype (UF 74473), preserved maximum diameter 19.0 mm, Ocala Limestone, UF locality ALO04 (Dickerson Limerock Mines), Alachua County, Florida; the topmost portion (indicated by arrow) of Figure 29 is magnified 3.5x in relation to Figure 30 to show details of the sutural ramps Type Locality and Occurrence: The holotype (UF 18737, UF 57018, and UF 110360, UF locality LFOO1, LOSS58; Figures 23, 24) is from the lower member of the Dell Limerock Mine, Ocala Limestone). Thus, all speci- Ocala Limestone (formerly the Inglis Formation) at UF mens of C. palmerae are from the upper Eocene, Jack- locality LVO14, Gulf Hammock 02, Levy County, sonian Ocala Limestone of Florida Florida. Richards and Palmer (1953: 5) described the locality R. O. Vernon’s L-93) as a “road metal pit 2.9 Etymology: This species 1s named in honor of miles south of the north limits of the town of Gulf Ham- Katherine V. W. Palmer (1895-1982). second director of mock just southwest of State Road 55 in the southwest the Paleontological Research Institution (Ithaca, NY), quarter of Section 34, Township 14 South, Range 16 who was the first to recognize this form as a new, unde- East.” The varatype specimens are from \lachua County scribed species and for her important contributions to UF 68306, UF 74473, UF 111327, and UF 112981, UF Cenozoic paleontology (see Caster, 1983) locality ALO04, Dickerson Limerock Mines, Ocala Lime- tone; UF 18599, UF locality ALO16, S.M. Wall Quarry Discussion: Palmer (1953) did not describe UF Ol, Ocala Limestone; UF 15886, UF locality ALO17, LOSS5S as a new taxon because she did not consider this Newb« Corporation Pit 01, Ocala Limestone; and UF single damaged shell adequate for this purpose. Newly 6735, UF locality ALO28, Newberry 03, Ocala Lime collected specimens (all molds) are consistent with the | Lafayette County (UF 18711, UF 18719, UF gross morphology of Palmer's fossil, but offer new mor- J. RB. Hendricks and R. W. Portell, 2008 Page 89 Figures 31-39. Specimens of Conus alleni new species (31-35), C. smithvillensis Harris, 1895 (36-38), and C. cormacki Har- bison, 1944 (39). Scale bar equals 1 cm. 31-32. Holotype (UF 119920) of C. alleni, shell length 36.2 mm, preserved maximum | ( liameter 20.7 mm, Moodys Branch Formation, UF locality ZLO04 (Montgomery Landing), Grant Parish, Louisiana. 33. Paratype UF 119977) of C. alleni, preserved shell length 35.1 mm, Ocala Limestone, UF locality LF002 (Mill Creek Quarry), Lafayette County Florida. 34. Paratype (UF 119919) of C alleni preserved shell length 33.2 mm, Ocala Limestone, UF locality SUO14 (Suwannee American Cement), Suwannee County, Florida. 35. Paratype (UF 119976) of C. alleni, preserved shell length 34.5 mm, UF locality LF002 (Mill Creek Quarry), Lafayette County, Florida. 36-38. Holotype TMM-BEG 35656) of C. smithvillensis, shell length 42.1 mm, pre served maximum diameter 14.8 mm, Weches Formation, Colorado River at Smithville, Sastrop County, Texas. 39. Holotype (ANSP 16145) of C. cormacki (not a Conus; see text), preserved shell length 28.7 mm, Santee Cooper Canal, South Carolina phological details that now justify description of this ridens Conrad, 1833, and C. alleni new species. Conus species palmerae may be distinguished from both of these species Conus pale rae co-occurs in the Ocala Limestone by its stepped teleoconch whorls smooth early postnuclear with two other late Eocene Conus in Florida: C. sau- whorls, smooth sutural ramps and smooth last whorl Page 90 THE NAUTILUS, Vol. 122, No. 2 Conus alleni new species (Figures 31-35, Table 2) Diagnosis: Shoulder undulate; subsutural flexure di- agonal; last whorl with raised spiral cords on anterior half. Description: Shell medium-sized (up to about 36 mm in length). Last whorl conical; outline straight to slightly sigmoidal (convex near shoulder). Shell widest at shoul- der. Shoulder angulate and with large tubercles resulting from weak undulations. Spire of moderate height; outline straight to slightly concave. Larval shell unknown. Early postnuclear w horls tuberculate. Subsutural flexure di- agonal, depth about 1.5x width. Teleoconch sutural ramps concave with several raised spiral cords. Aperture opening about as wide at base as at shoulder. Last whorl with pronounced raised spiral cords on anterior half, sometimes extending weakly as threads to shoulder. Type Series: Holotype, UF 119920 (Figures 31, 32). The three paratypes consist of external molds and in- clude UF 119977 (Figure 33), UF 119919 (Figure 34), and UF 119976 (Figure 35). See Table 2 for measure- ments of these specimens. Type Locality and Occurrence: The holotype (UF 119920, Figures 31, 32) was collected by J. E. Allen from the Jacksonian Minas Branch Formation at Montgom- ery Landing (UF locality ZL004), Grant Parish, Louisi- ana. The paratypes are all from the Ocala Limestone of Florida, including two specimens from Lafayette County (UF 119976, U F 119977, UF locality LF002, Mill Creek Quarry) and one specimen from Suwannee C ounty (UF 119919, UF locality SU014, Suwannee American Ce- ment), Etymology: This species is named in honor of James E. Allen (1914-1997) of Alexandria, Louisiana, who was an enthusiastic collector and scholar of Gulf Coast Eocene mollusks. Discussion: Louisiana with C. sauridens and in the Ocala Limestone of Florida with C. palmerae new species and C. sau- ridens. Conus alleni can be readily differentiated from both species by its undulate shoulder and spiral cords on the anterior half of the last whorl (raised spiral threads on the last whorl may also be present on small shells of C. palmerae, but if so are much weaker). Conus alleni shares some resemblance with a moldic Oligocene fossil (USNM 166720) from Decatur County, Georgia that Dall ( rs described as C. vaughani. Dall’s (1916: pl. 56, fig. 1) figure of the cast shows a specimen (partially obsei re a be matrix) with an obtuse spire angle, undulate shoulder, and raised spiral threads on the su- tural ramps that are similar to the teleoconch morphol- ogy of C, alleni. The presence of rows of spiral beads on the last whorl, the fact that the shell is widest below the shoulder (rather than at the shoulder, as in C. alleni), and the fact that the anterior end of the shell appears com- Conus alleni co-occurs in the Eocene of pletely obscured by matrix prevents us, however, from considering these two forms e quivalent. The only known shell material of Conus alleni is the holotype (from Grant Parish, Louisiana); the other three specimens are from Florida and all consist of external molds. This taxon was apparently rare, especially outside of Florida. We recognized this new form from the moldic Floridian material belo we—by chance—discovered the similar shell from Louisiana in the FLMNH collec- tions. We chose to designate the shell as the holotype, rather than one of the paratype external molds, because of its greater number of characters available for obser- vation. OTHER RECORDS OF EOCENE CONUS FROM THE U.S. COASTAL PLAIN Unidentifiable internal molds of Conus are common in the Eocene Ocala Limestone of northern Florida. We examined 54 such lots (over 440 specimens) from Ala- chua, Citrus, Jackson, Lafayette, Marion, and Suwannee counties. These include: UF 15884, UF 15892, UF 15905, UF 17831, UF 17832, UF 17879, UF 17947, UF 17950, UF 17967, UF 18423, UF 18759, UF 18848, UF 18864, UF 18896, UF 18955, UF 18962, UF 19140, UF 19174, UF 19204, UF 19215, UF 20744-20746, UF 46435, UF 68270, UF 107265, UF 119900-UF 119904, UF 119906-119911, UF 119914119917, UF 119921, UF 120027, UF 120032-120040, UF 120047, and UF 126926. During the course of this work, we became aware of several other Eocene Coastal Plain Conus species that are likely distinct, but are not known to occur in Florida and will require additional investigation; these include: Conus smithvillensis Harris, 1895: C. smithvillensis var. ee sry, 1980 (also see C. smithvillensis var. Dockery in Campbell, 1995); and Conus haighti Gardner, 1945. Harris (1895) described C. sniiudlonss from the Colorado River at Smithville, Bastrop County, Texas (Weches Formation according to TMM records). The holotype (TMM-BEG 35656; Figures 36-35) shares some characteristics with C. sauridens (including a mul- tispiral protoconch, tuberculate early postnuclear whorls, spiral threads on the sutural ramp, growth lines showing a deep and symmetrically curved cubsuual flexure, aan incised spiral g erooves near the base of the last whorl), but has a very different overall shell shape: the spire is much higher (spire angle: 53°, Table 2; in mature individuals of C. sauridens, the spire angle is typically over 100°), the conical last whorl has straight sides, and the sutural ramps are flat. Dockery (1980) figured a shell (MGS 590) from the slightly younger Cook Mountain Formation of Mississippi this tt he described as a varie ty of C. smithvil- lensis. His figured shell appears to bear many of the discrete characteristics of C. smithvillensis described above, though has a lower spire (ca. 76°) and the last whorl is sigmoidal in profile. Conus smithvillensis is not at all similar in form to C. palmerae. While it bears some of the shell characteristics of C. alleni, it lacks C. alleni’s J. R. Hendricks and R. W. Portell, 2008 Page 91 distinctive raised spiral cords at the base of the last whorl. Campbell (1995) attributed an external mold (UNC 15448) from the Santee Limestone near Cross, South Carolina to Dockery’s (1980) ae of C. smithvillensis. Campbell (1995: 146) stated that this form is the “most common Conus in the Santee Limestone” and that it “has a nodose shoulder and a taller, more tabulate spire than the widespread Conus (Lithoconus) sauridens.” Further study will be required to determine whether these vari- eties are indeed consistent with Harris’s taxon. Gardner (1945) described C. haighti from the Laredo Formation of Zapata County, Texas. She did not figure her a (USNM 495182), but her figured (pl. 26, fig. 7) paratype specimen (USNM 495183) has a convex spire profile and rounded shoulder that appear distinct from those characters in C. sawridens, C. palmerae, and C. alleni. Gardner’s other figured (pl. 26, fig. 2) specimen of C. haighti lacks this distinctive spire form. NOMINA DUBIA The following species of Conus described from the Eocene of the U. S. Coastal Plain should be regarded as nomina dubia: Conus gyratus Morton, 1834; Conus mu- tilatus Tuomey, 1852; Conus improvidus de Gregorio, 1890; and Conus cormacki Harbison, 1944. Conus gyra- tus (holotype, ANSP 211) is an internal mold of a shell of uncertain provenance from South Carolina (also see Kohn, 1992); Campbell (1995) suggested that it could be an internal mold of the purported variety of C. smithvil- lensis presented by Dockery (1980). Conus multilatus was described (but not fieured) by Tuomey from casts found near Wilmington, North Carolina. De Gregorio (1890) described C. improvidus from Claibome, Ales bama, but his holotype is reportedly lost (Palmer and Brann, 1966) and Palmer (1937: 465) considered the taxon “of doubtful status as an American species.” Conus cormacki Harbison, 1944 was described from the Eocene Santee Formation of South Carolina, but the holotype (ANSP 16415, Figure 39, Table 2) is clearly not a Conus. Campbell (1995: 146) stated that C. cormacki “is actually a broken volutid.” Several other Eocene Conus species are also problem- atic. Conus pulcherrimus eee 1879 (type, AMNH- FI 10175) was recognized by Harris (1895) as a turrid (see Palmer and Brann, 1966). The type specimen of Conus (Conospirus) granopsis de Gregorio (1890) is lost (Palmer and Brann, 1966) and Dall (1896) and Palmer 1937) suggested that the small shell figured by de Gre- gorio (1890) may be the juvenile of “another species. Given that the type specimen is lost and that the shell is likely a juvenile ania C. granopsis is a name that should probably be disregarded. Finally, along with Conus sp. B (described here as the new species C. palmerae; holotype, UF 108858), — in Richards and Palmer (1953) also noted a Conus sp. She said, “species A... is a broad (21 mm.), low- Ree (7 mm.) shell with sharp angulation of the s houlder of the whorls; the surface was apparently smooth. The speci- men is a fragment, 25 mm. high” (Palmer, 1953: 40). We located this specimen (UF 108683), which—like UF 10S858—is also a calcite pseudomorph from UF locality LV014, Gulf Hammock 02, Levy County, Florida (lower member of the Ocala Limestone). UF 108683 is poorly preserved and is too fragmentary (more than half of the spire is eroded away and much of the last whorl is miss- ing) to either assign to a known species or to describe as new species. ACKNOWLEDGMENTS Sable Allen (Alexandria, Louisiana) graciously donated to the FLMNH the large, well-documented Eocene Gulf Coast mollusk collection of her late husband, James E, Allen. Larry Rogers (Limestone Products), Wayne Bea- ver (Denali Limerock Mine), Joe Horton (Suwannee American Cement), and Leon Brooks (Hi-Cal Quarry) kindly allowed access to collect in their Florida mines. Paul Callomon (Academy of Natural Sciences, Philadel- phia) and Ann Molineux (Texas Memorial Museum) are thanked for allowing access to type specimens. Ann Mo- lineux also pr ovided helpful discussions on Eocene stra- tigraphy in Texas. Bushra Hussaini and Sandy Ebersole furnished information pertaining to an American Mu- seum of Natural History specimen and a Geological Survey of Alabama specimen, respectively. Alan Kohn provided helpful discussions of and suggestions for ter- minology related to the “subsutural Fexre” of cone shells. Sean Roberts (Florida Museum of Natural His- tory) assisted us a some of the digital photography using a Sony DSC RIL camera (10.3 megapixel resolu- tion). We thank Alan Kohn and an anonymous reviewer for comments that improved the quality of this manu- script. JRH’s contributions to this work were supported by the Brayfield Award of the South West Florida Fossil Club and NSF EAR 0518976. RWP’s contribution to this work was supported by the McGinty Endowment of the Florida Museum of Natural History, by Barbara and Reed Toomey, and by the late David ‘Nicol whose $1000 grant allowed RWP to permanently transfer the Florida Geological Survey Eocene mollusk collection from the Paleontological Research Institution (where it was under study by Katherine V.W. Palmer) to the Florida Museum of Natural History. This is Univer sity of Florida Contri- bution to Paleobiology 602. LITERATURE CITED Applin, P. L. and E. R. Applin. 1944. 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Definitive locations of Paleocene and Eocene marine fossil localities, Colorado River, Bastrop County, Texas. Texas Journal of Science 57: 317-328. THE NAUTILUS 122(2):94-98, 2008 Page 94 Synonymization of Neohyalimax Simroth, 1896, and Omalonyx dOrbigny, 1837, with a redescription of Omalonyx brasiliensis (Simroth, 1896) (Gastropoda: Succineidae ) Janine O. Arruda José W. Thomé Laboratorio de Malacologia Museu de Ciéncias e Tecnologia Pontificia Universidade Catolica do Rio Grande do Sul Avenida Ipiranga 6681, 90619—900 Porto Alegre, BRAZIL arrudajo@gmail.com josewthor 1 1e@pq.cnpq.| or ABSTRACT Neohyalimax was established as a monotypic genus for N. bra- siliensis. Characters of the reproductive system prompted au- thors to subsequently treat the taxon as a subgenus of Omal- onyx. Examination of the holotype and discussion of these char- acters allowed Neohyalimax to be synonymized with Omalonyx. Omalonyx brasiliensis remains a valid species based solely on the holotype. Additional Keywords: Molhusca, Pulmonata, Brazil, Rio Grande do Sul INTRODUCTION The genus Omalonyx @Orbigny, 1837, comprises succi- ne ge. slugs of West Indies, Central and South American and Juan Fernandez, with a reduced, flat and fingernail- like shell. Omalonyx has commonly been classified with two subgenera (e.g., Patterson, 1971: Tillier, 1981), namely Omalonyx sensu stricto and Neohyalimax Sim- roth, 1896. Zilch (1959-1960) and Thiele (1992) indi- cated that the separation between the subgenera Oma- lonyx and Neohyalimax is based on the mantle covering either only the margin or the entire shell and the spire being small or scarce ‘ly indicated. The monotypic subge- nus Neohyalimax, type-species Neohyalimax brasiliensis Simroth, 1896, was described from a unique specimen collected from Rio Grande do Brazil. In revising the family Succineidae Beck, 1837, Patterson (1971) dis- cussed the similarities of Didone felina (Guppy, 1872) O. matheroni (Potiez and Michaud, 1835), according to Tillier, 1981] and N. brasiliensis, and brought atten- tion in partic silar to details of the origins ul dese ription of the latter species (name ‘ly the number of seminal vesicles and c¢ phalic tentacles) that suggested recognition of Veohyalimax at the subgeneric level. It should be noted, however, that Patterson (1971) did not examine the type of N. brasiliensis, nor made a direct comparison with O. unguis (dOrbigny, 1837), the type species of Omalonyx. In a revision of the South American and Juan Fernandez succineid slugs, Tillier (1981) similarly did not examine the type of N. brasiliensis, but accepted Patterson (1971) assessment of taxonomic status based on the original de- scription prov ided by Simroth (1896). In support of rec- ognition at the subgeneric rank, Tillier (1981: 144) noted ‘an altogether diflere nt way of life (terrestrial), a plesio- morphic radula (marginal teeth still longitudinally elon- gated) and slug-like ae wacters more apomorphic than in Omalonyx sensu stricto’. Simroth (1896) mentioned that the type specimen of N. brasiliensis was deposited with the “Berlin Museum”. ss d there is a specimen in Museum fiir Naturkunde der Humboldt Universitit zu Berlin (ZMB 45.913) la- beled Neohyalimax brasiliensis, but it bears no indication of type status. That the specimen corresponds well with the de scription given by Simroth ( including most details of the internal anatomy) and that no other specimen of this species has been located, the ZMB specimen is here considered to be the holotype. With the recent rediscove sry of Simroth’s type speci- men, we take the opportunity to redescribe N. brasilien- sis and reassess its taxonomic status. MATERIALS AND METHODS The following lots were examined: GUADELOUPE, Pointe-a-Pitre, Omalonyx matheroni, 4 spec., 27.1X.1972, Pointier leg. (MNHN); O. matheroni, 11 spec., S. Tillier det. 1979. (MNHN); FRENCH GUI- ANA, along the road Cayenne-Kourou, 17 km SE Kourou bridge, O. matheroni, 5 spec., Tillier leg. (MNEIN); PARAGUAY, O. ungitis (lectotype designated J. O. Arruda and J. W. Thomé, 2008 ‘ 2 O5 Page 95 by Tillier, L9S1), 1 shell. (MNHN); Asuncion, Asuncién, O. unguis, 1 spec., canje F.H. Schade. (MACN 199658, it is labeled as Hom. unguis @Orb.); ARGENTINA, For- mosa, Villafane (26°14.17' S 59°07.08' W, Arroyo Bel- laco, piece La Marcela), unguis, 2 spec., 30.X.2004, Cristian Ituarte leg. (MLP 11878); BRAZIL, Rio G eae do Sul, Neohiyainer brasiliensis Simroth, 1 spec. (ZMB 45.913) Cachoeirinha, O. convexus, 34 spec., 29.1V.2006, A. Palatin and J. O. Arruda leg., J. O. Arruda det. (MCP. 8839): Pelotas (Estrada Pelnies hie Grande, 31°45'53.1" S 52°22'48.2” W), O. convexus, 48 spec., 18.11.2006, A. Paladini and J. O. Neruda leg., J. O. Arruda det. (MCP 8836). Based on examination of the holotype of Neohyalimax brasiliensis, we redescribed the reproductive system (Figure 1). The specimen had been previously dissected, with a sagittal cut from its head to the mantle edge, which then continues towards the left side of animal’s tegument. The reproductive system was separated in two parts: one including the vagina (previously dissected and with the papillae on its internal surface exposed), the Ss —- f x & ae fc — io 3 —— pr a % 4M . { . YS ie 2 Lay * a —— Ne 7 — Yee vi S) e- 1mm Figure 1. Omalonyx brasiliensis (Simroth, 1896) (ZMB 45.913). Abbre- viations: ag, albumen gland; b, epiphallial bulb; dd, deferent duct: ep, epiphallus fe, fertilization complex: fo, free oviduct; gg, gametolitic gland: 0, ovariotestis; ov, ovulispermduct; p- evertophallus: po, pallial oviduct; pr, prostate; ps, evertophal- lus sheath: rm, retractor muscle; sv, seminal vesicles; v, vagina. Reconstruction of the reproductive system of gametolitic gland duct and its gland and the free oviduct: the other including the pallial oviduct, prostate, albumen gland, fertilization “complex, ovulispermduct, ovariotestis, deferent duct, epiphallus and evertophallus (= penis). Institutional abbreviations: MACN, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de las Ciencias Naturales (Buenos Aires, Argentina), MCP, Museu de Ciéncias e Tecnologia da Pontificia Universidade Cat6lica do Rio Grande do Sul (Porto Alegre, Brazil); MLP, Museo de La Plata (La Plata, Argentina), MNHN, Muséum national dHistoire naturelle (Paris, France) and ZMB, Museum fiir Naturkunde der Humboldt Universitit zu Berlin (Germany). SYSTEMATICS Order Heterurethra Pilsbry, 1900 Superfamily Succineoidea Beck, 1837 Family Succineidae Beck, 1537 Genus Omalonyx d Orbigny, 1537 Type Species: Succinea (Omalonyx) unguis @Orbigny, 1837: 229 (year according Sherborn and Woodward, 1901), by original designation. Diagnosis: Shell very flat. fingernail-shaped, obtuse and small spire. body w vhorl and aperture very large and elongate-oval, animal body twice the shell length: ever- tophallus exceptionally long. Omalonyx brasiliensis (Simroth, 1596) Neohyalimax brasiliensis Simroth, 1896: 39-45; Morretes, 1949: 130; Simone, 2006: 237. Omalonyx (Neohyalimax) brasiliensis —Zilch, 1959-1960: 202; Patterson, 1971: 187; Tillier: 1981. Homalonyx (Neohyalymax) brasiliensis.—Thiele, 1992. Omalonyx brasiliensis —Salgado and Coelho, 2003: 153. Diagnosis: A species of Omalonyx with a thick-walled vagina with robust diamond shaped papillae on its inter- nal surface. Holotype: ZMB 45.913 Type Locality: Brazil, Rio Grande do Sul. Redescription: The shell is thin, oval and highly flat- tened (it is fragmented, which precludes its measure- ment). The jaw and radula are missing (see Discussion be- low). Within the reproductive system, the epiphallus is approximately one fourth of the le ngth of the evertophal- lus. The former possesses a thick wall, a smooth outer face and a bulb, and its internal surface has transverse folds. The opening of the epiphallus into the evertophal- lus is a simple opening, without any papilla. The evertophallus has a thick, wide wall, and the in- ternal surface is covered with aligned diamond shaped papillae. At its proximal extremity, the papillae are Page 96 THE NAUTILUS, Vol. 122, No. 2 smaller and discoid. At its distal extremity there are small longitudinal folds. The evertophallus sheath, thick and aiuseular: covers practically the entire distal half of the evertophallus. It gradually becomes a thin and transpar- ent membrane, w ich covers the other half of the ever- tophallus and the epiphallus. The evertophallus retractor muscle inserts at the junction of the deferent duct with the epiphallus. The prostate is long and the section of the deferent duct that is embedded in the prostate has small orifices throughout its length within the gland. The vagina is approximately half of the length of the ev ertophallus. It has a thick wall and aligned, diamond shaped papillae on its internal surface. These papillae are similar to those on the evertophallus, but more robust. The gametolitic gland is spherical, with a diameter ap- proximateh ly half the length of its duct. The gametolitic gland duct is slim and about two thirds the length of the vagina. The free oviduct is long, thick, and convoluted with rippled longitudinal folds on its internal surface. The pallial ov iduct, approximately half the length of the lower oviduct, is very inflated and convoluted. The albumen gland is small, bean-like and about one third the length of the prostate, The ovulispermduct 1S slightly convoluted and more inflated in its distal portion. The ovariotestis is spherical and large. The pallial cavity and the nervous and digestive sys- tems are damaged. DISCUSSION Simroth (1896: 40, pL. V, Fig. 3) characterized the shell as ‘a small flat plate of 9 mm in length and 6 mm in breadth. Beneath the light-yellowish periostracum the lime was deposited in concentric rings, with an excentric nucleus to the left of the posterior end. The nearer the nucleus, the thicker is the shelly matter. The periphery is solid, surrounded, near the poste rior right-hand margin, by « brown conchiolin line. The nucleus is somawliat ade ened; the shell throughout is perfectly flat, and therefore does not enclose any of the soft parts’. The shell, accord- ing to Simroth (op. cit. ), would pl we Ae specie s close to the genus Hyalimax H. & A. Adams, 1855. However, that author re cognize ‘d that the shell ‘In ee it is slightly . Simroth (1896; 41) described the jaw as ‘brown and lit] | 1as a sharp edge, with a slight median process (oxygnath) and a lighter -coloured palatal plate arched (Fischer) (elasmognath), the two posterior corners of which are somewhat prominent. This plate is finely sculptured, the thread-like markings converging towards the median projection’ Simroth (1896: 4 lows: ‘the median tooth is tricuspid, the middle cusp being very large. In the lateral teeth the median cusp is ) described the radular teeth as fol- still larger, and directed oblique ‘ly towards the middle line of the radula; the inner cusp is reduced in size, whilst the outer one is duplicated. In the marginal teeth the inner cusp increases in size, whilst the exterior outer cusp subdivides, giving rise to three outer cusps’. In the description of O. felina, Guppy (1872) indicated that the shell was usually covered by the mantle, but when retracted, it occasionally exposed the central region of the shell. Gibbons (1879), however, repeatedly ob- served living specimens of Omalonyx and never noticed more than a limited portion of the shell border covered by the mantle. In observations of live specimens of Oma- lonyx from the states of Rio Grande do Sul and Minas Gerais, Brazil, we found that the mantle covered the shell in different degrees, and although the shell was well covered in some, in none presented its shell completely covered by the mantle. Based on Guppy (1872) and on our observation of live specimens of Omalonyx, the char- acter “mantle covering partially or totally the shell”, in- dicated by Zilch ( 1959-1960) and Thiele (1992) for the separation of the subgenre Omalonyx and Neohyalimax. are unjustifiable. The other character, “small spire or extremely small spire” is not a systematic character. Patterson (1971), using Simroth’s de scription of Neo- hyalimax brasiliensis as reference, indicated that it has similarities with Omalonyx felina concerning the general body's shape, the mantle pigmentation and the anatomy of the reproductive system. Because the reproductive system of Neohyalimax brasiliensis is similar to that in Omalonyx s.s and different from that in Hyalimax, which resembles Succinea Draparnaud, 1801, Patterson consid- ered Neohyalimax as a subgenus of Omalonyx. However, Patterson also noted that “Neohyalimax brasiliensis has one seminal vesicle, whereas there are two in O. felina, and that N. brasiliensis has only one pair of tentacles, while O. felina has two. We verified the presence of two vesicles and one fecundation pouch in N. brasiliensis, which contradicts its original description by Simroth (1896) and concurs with the observations of Patterson (1971) for rH sensu stricto. Simroth (1896) observed only one pair of tentacles in N. brasiliensis, the ommatophores. Patterson (1971), however, stated that Omalonyx sensu stricto has two pairs of tentacles. Through an external ventral view exam of the anterior region of N. brasiliensis, we could observe that the lower tentacles are present, very close to the animal's mouth opening, Simroth (1896) did not mention the presence of an epiphallus in N. brasiliensis. However, we verified the presence of this structure, which is slightly narrower than the evertophallus and is also present in the other species of Omalonyx s.s. We also observed small longitudinal folds at the distal extremity of the eve rtophallus, which are present in all Omalonyx species with known anatomy. Barker (2001) noted the placement of the genital ap- erture in Neohyalimax immediately adjacent to the right tentacle. We confirmed that this does not differ from its position in other species of Omalonyx sensu stricto. In the state of Rio Grande do Sul, type locality of O. brasiliensis, no specimens with the characteristics of this species were found, despite extensive investigation and collecting. In the municipalities of Cachoeirinha, Pelotas J. O. Arruda and J. W. Thomé, 2008 Page 97 and other localities in Rio Grande do Sul, only Omalonyx convexus (Heynemann, LS6S) was found, suggesting its overlapping geographical area with O. brasiliensis. The reproductive systems of these species are distinct. In O. convexus, the ovariotestis is hemispherical or discoidal: the lower oviduct is generally half the length of the upper oviduct; half of the proximal region of the evertophallus is generally a little wider than the rest, and its internal surface has longitudinal folds in the wider proximal re- gion and cordiform papillae in the distal region; the in- ternal surface of the vagina has smooth longitudinal folds. In O. brasiliensis, the ovariotestis is spherical and large; the lower oviduct is approximately twice the length of the upper oviduct; the internal surface of the ever- tophallus has diamond shaped papillae and the vagina has similar but more robust papillae. The reproductive system of O. brasiliensis resembles that in the studied specimens of O. matheroni (Potiez and Michaud, 1835), which agrees with the meticulous description of Tillier (1981), with the exception of the internal surface of the vagina. According to our observa- tions and to Tillier, the vagina in O. matheroni has ir- regular longitudinal folds, inconsistent in number or shape, whereas O. brasiliensis has papillae instead of folds, similar to the ones in its evertophallus. Despite the reproductive system similarities of these species, they do not have overlapping ranges. Tillier (1951) considered the probable range of O. matheroni to be the entire Amazon basin, the eastern coastal regions of South America from Caracas (Venezuela) to Rio de Janeiro (Brazil) and part of the Lesser Antilles. Neohyalimax bra- siliensis has been reported only once, from Rio Grande do Sul (Brazil). When compared to the O. unguis (@Orbigny, 1837) type species, O. brasiliensis presents specifics different characteristics such as a smooth external surface of the epiphallus, the internal surface of the evertophallus and vagina has diamond shaped papillae, more robust in the latter, the length of the gametolitic gland corresponds to approximately a half the length of its duct and the lower oviduct, which is little Goavolite: is generally two times the length of the upper oviduct. In O. unguis, according ) Arruda et al. (2006), the external surface of the epiphallus has a looped fold, the internal surface of the evertophallus has discoidal papillae, the internal surface of the vagina has inflated elliptic papillae, the length of the gametolitic gland corresponds to approximately a quarter the length of its duct and the lower oviduct, which is convolute, is approximately three times the length of the upper oviduct. CONCLUSION Considering the foregoing discussion regarding the num- ber of seminal vesicles, the number of tentacles and the position of the genital aperture, as well as the observa- tions of the reproductive system, we here consider Neo- hyalimax Simroth, 1596 as a junior synonym of Omalo- nyx @Orbigny, 1837. Omalonyx brasiliensis (Simroth, 1896) remains a valid species, based on the only known specimen, which we consider to be the holotype. ACKNOWLEDGMENTS We cae the reviewers, Dr. Gary Barker and Dr. Robert Cowie, for their great contribution with very good criti- cisms and improving the English; Alejandro. Tablado (MACN), Cristidn Ituarte (MLP), Lticia Maria Z. Rich- initti (MCP), Virginie Héros (MNHN) and Matthias Glaubrecht (ZMB) for loaning material; Guacira Gil for comments and suggestions; Luiz Augusto F. Aratijo for the translation from Portuguese to English; to Conselho Nacional de Desenvolvimento Cientifico e Tecnolégico for financial support (CNPq 131716/2005-0 for JOA, 500032/2003-3 for JWT). LITERATURE CITED Arruda, J. O., S. R. Gomes, R. Ramirez, and J. W. Thomé 2006. Morfoanatomia de duas espécies do género Omalonyx (Mollusca, Pulmonata, Succineidae) com novo registro para Minas Gerais, Brasil. Biociéncias 14 (1): 61-70. Barker, G. M. 2001. Gastropods on land: phylogeny, diversity and adaptive morphology. In: Barker, G. M. (ed.) The biology of terrestrial molluscs. CABI Publishing, New York, PP 1-146. : Beck, H. 1837. Index Molluscorum praesentis aevi Musei Prin- cipis Augustissimi Christiani Frederici. Copenhagen, 124 pp. Gibbons, J. S. 1879. Comparison of Omalonyx unguis d’Orb., with O. felina Guppy. Journal of Conchology 2: 98-101. Guppy, R. J. L. 1872. Third series of additions to the catalogue of the land and freshwater Mollusca of Trinidad, with a revised list of all species. Proceedings of the Scientific Association of Trinidad 2: 17-25. Morretes, F. L. 1949. Ensaio de Catalogo dos Moluscos do Brasil. Arquivos do Museu Paranaense 7 (1): 1-216. dOrbigny, A.D. 1837 [1835-1847]. Voyage dans ! Amerique Méridionale: Mollusques. Paris: Pitois-Levraut 5(3): 1-758. Patterson, C. M. 1971. Taxonomic studies of the land snail family Succineidae. Malacological Review 4: 131-202. Pilsbry, H. A. 1900. On the zoological position of Partula and Achatinella. Proceedings of the Academy of Natural Sci- ences of Philadelphia 3: 561-567, pl. 17. Salgado, N. C. and A. C. 8. Coelho, 2003. Moluscos terrestres do Brasil (Gastropodes operculados ou nao, exclusive Veronicellidae, Milacidae e Limacidae). Revista de Bi- ologia Tropical (International Journ il of Tropical Biology and Conservation) 51 ( (suppl. 3: Malacologia Latinoameri- cana): 149-189. Sherborn, C. D. and Woodward, B. B. 1901. Notes on the dates of publication of the Natural History portions of some French Voyages—Part I. ‘Amérique méridionale’; ‘Indes orientales’; ‘Pole Sud’; (‘Astrolabe’ and “Zélée’); ‘La Bon- ite’; “La Coquille’; and ‘L’Uranie et Physicienne’. Annals and Magazine of Natural History (series 7) 7 (40): 385— 392. Page 95 THE NAUTILUS, Vol. 122, No. 2 Simone, L. R. 2006. Land and freshwater molluses of Brazil. Fapesp, Sao Paulo, 390 pp. Simroth, H. 1896. On Neohyalimax brasiliensis, n. gen., n. sp. (allied to Hyalimax), from Brazil. Proceedings of the Mal- acological Society of London 2: 39-45, Thiele, J. 1992. Handbook of Systematic Malacology. Part 2 (Gastropoda: Opisthobranchia and Pulmonata). In: Bieler, R and P. Mikkelsen (translators), SmithsonianInstitution Libraries. The National Science Foundation, Washington 627-1189 pp. Tillier, S. 1981. South American and Juan Femandez succineid slugs (Pulmonata). Journal of Molluscan Studies 47; 125— 146. Zilch, A. 1959-60. Euthyneura. In: Wenz, W, (ed.). Handbuch der Paliiozoologie, Gebriider Borntraeger, Berlin, vol. 2, 834 pp. THE NAUTILUS 122(2):99-106, 2008 Page 99 Rehabilitation of Ergalatax martensi (Schepman, 1892) (Gastropoda: Muricidae), senior synonym of Ergalatax obscura Houart, 1996, and description of Ergalatax junionae, new name for Morula martensi Dall, 1923 Roland Houart Institut royal des Sciences naturelles de Belgique Rue Vautier, 29 B-1000 Bruxelles, BELGIUM roland. houart@skynet.be ABSTRACT Ergalatax junionae nomen novum is proposed as replacement name for Morula martensi Dall, 1923, from the Gulf of Oman and the Persian Gulf. The latter name is a junior secondary homonym of Pentadactylus martensi Schepman, 1892, from the Red Sea, of which E. obscura Houart, 1996, is here considered as a junior synonym. Ergalatax junionae nomen novum is com- pared with Ergalatax martensi (Schepman) and E. margariti- cola (Broderip, 1833). The geographical distribution of E junionae and E. martensi is updated and the introduced Medi- terranean species is identified as E. jumionae. Additional Keywords: Mollusca, Gastropoda, Muricidae, Gulf of Oman. Persian Gulf, Red Sea, Mediterranean Sea, hom- onymy, synonymy, new name. INTRODUCTION Some vears ago (Houart, 1996: 13), I described Ergalatax obscura, a muricid from the Red Sea. Shortly after that paper was published, R. G. Moolenbeek (in litt.) told me about a name introduced by Schepman (1892), Penta- dactylus martensi, which he also described from the Red Sea. Having then examined a syntype of that species from ZMA (Figure 14), I concluded that P. martensi Schepman, 1592, was a junior synonym of Ergalatax margariticola (Broderip, 1833). However. after recently requesting additional infor- mation about the other syntypes of P. martensi in NMNL, I received digital images that illustrate the re- maining three specimens (Figures 1S—23), among them the 17 mm long specimen that probably served for the description (Figures 15-19). Those syntypes are un- doubtedly conspecific with E. obscura, while the ZMA syntype represents E. margariticola. For many years (Houart, 2001), I had identified an introduced eastern Mediterranean species as Ergalatax obscura Houart, 1996 [= E. martensi (Schepman, 1$92)]. However, this concerns yet a third species, which occurs primarily in the Gulf of Oman and in the Persian Gulf. The species was first wrongly illustrated as Purpura (Ricinula) siderea Reeve (a buccinid) by von Martens (1874: 95, pl. 5, fig. 49). Later, Dall (1923: 304) ) provided anew name for that species, naming it Morula martensi. Unfortunately, both species, Pe ntadactylus marte nsi Schepman, 1892. and Morula martensi Dall, 1923. are now included in Ergalatax, and D ipso facto a junior secondary homonym and needs to be replaced. I personally mixed both species because of their close enters and the oo of some related forms (compare Figures 30 and 35). First (in litt.), I provision- ally identified the Meditenanean specime nas E. mar- tensi (Dall, 1923), but afterwards | erroneously consid- ered it as conspecific with E. obscura [= E. martensi (Schepman, 1892)] from the Red Sea and the Gull of Aden (Houart, 1996: 12). Abbreviations used herein are: AMS: Australian Mu- seum. Sydney, Australias MNHN: Muséum national (Histoire naturelle, Paris, France; NMNL: National Museum of Natural History Naturalis, Leiden, The Netherlands: RH: collection of the author; ZMA: Zoolo- gisch Museum, University of Amsterdam, The Nether- lands: ZMB: Museum fiir Naturkunde der Humboldt Universitit zu Berlin, zoologisches Museum, Germany. Terminology for shell spiral ae (after Merle, 1999 and 2001) given in Figure 1. all’s name oem SYSTEMATICS Subfamily Muricoidea Rafinesque, 1$15 Family Muricidae Rafinesque, 1S15 Subfamily Ergalataxinae Kuroda, Habe and Oyama, 1971 Genus Ergalatax Iredale, 1931 Ergalatax recurrens Iredale, 1931, Aus- 1846)] by Type Species: tralia (Figure 17) [?= E. contracta (Reeve, original designation. Page 100 THE NAUTILUS, Vol. 122, No, 2 TELEOCONCH WHORLS SP | Subsutural cord ‘TP Infrasutural primary cord (primary cord on shoulder) adis : adapical infrasutural secondary cord (shoulder) abis : abapical infrasutural secondary cord (shoulder) P Primary cord 's secondary cord - t tertiary cord Pl ] Shoulder cord P2-P6 : Primary cords of the convex part of the teleoconch whorl sl-s5 : : secondary cords of the convex part of the teleoconch whorl example: s] = secondary cord between P| and P2; s2 = secondary cord between P2 and P3, etc SIPHONAL CANAL ADP: adapertural primary cord on the siphonal canal 7 “MP median primary cord on the siphonal canal APERTURE ID | Infrasutural denticle D1 to DS Abapical denticles Figure 1. Terminology used to describe the spiral cords (af- ter Merle, 1999 and 2001) (sporadic sculpture is shown in pa- rentheses). Ergalatax martenst (Schepman, 1892) (Figures 3, 5, 6, 11, 13, 18-30, 46) Pentadactylus (Morula) martensi Schepman, 1892: 104. Cronia martensi Dall—Sharabati, 1954: pl. 19, fig. 9, 9a, 9b; Singer and Mienis, 1991b: 58, fig. 19; Coulombel, 1994: 73, text fig. (not Morula martensi Dall, 1923); Verbinnen and Dirkx, 2000: 69, fig. 9 (not Morula martensi Dall, 1923). Drupella rugosa.—Singer and Mienis, 1991a: 18, fig. 6 (not Murex rugosus Born, 1778). Ergalatax obscura Houart, 1996: 13, figs 1, 3-6; Houart, 2001: 108 (in part), figs 17, 31, 106 and 449 only; Heiman and Mienis, 2003: 22-23 (text fig.). Not Morula martensi (Schepman, 1892) —Tan, 1995: 160, figs 52, 192 g, h (= Ergalatax junionae nom. nov.). Not Ergalatax obscura.—Houart, 2001 (in part): 108 (in part), figs 31 and 450-451 only; Delongueville and Scaillet, 2007: 57, fig. 31 (= Ergalatax junionae nom. nov.) Description: — Shell medium sized for the genus, up to 25.5 mm in length at maturity. LengthAvidth ratio 1.39- 1.92. Heavy, stout. Spire high with 3+ protoconch whorls (tip partially broken), and up to 7 broad, strongly shoul- dered teleoconch whorls. Suture adpressed. Protoconch conical, acute, whorls smooth. Axial sculpture of teleo- conch whorls consisting of high, rounded, nodose ribs: 10 or 11 from first to penultimate whorl, 6 to 8 on last whorl, rarely 5 or 9. Spiral sculpture of high, strong, primary, secondary and tertiary cords. Last teleoconch whorl with adis, IP, abis, P1, immediately followed by small P2, s2, P3, (t), s3, P4,(t), s4, P5, (s5), P6, ADP, MP. PL and P2 small, P3, P4 and P5 similar in strength, P6 very small or obsolete. Sculpture forming high, nodose knobs at intersection of spiral cords and axial ribs. Aperture relatively small, ovate. Columellar lip with 2 or 3 weak knobs abapically, rim adherent. Anal notch broad, moderately deep. Outer lip weakly crenulate, with 7 strong elongate denticles within [I[D, D2 (D1-D2 fused), D3 split, D4, D5 split]. Siphonal canal short, broad, broadly open. Milky white, Figures 2-5. Ergalatax species. Spiral sculpture and apertural denticles. 2, 4. Ergalatax junionae nomen novum (lectotype ZMB). 3,5. £. martensi (Schepman, 1892) (Red Sea, RH). R. Houart, 2008 Page 101 13 Figures 6-13. Ergalatax species. 6. Radula of Ergalatax martensi (Schepman, 1893), Gulf of Aden. Scale bar = 150 jum. 7. Radula of E. junionae nomen novum, Gulf of Iskenderun, Turkey. Scale bar = 120 jm. 8. Radula of E. margariticola (Broderip, 1833), Palau. Scale bar = 120 pm. 9-10. Radula of E. contracta (Reeve, 1846). 9. Aden, juvenile specimen. Scale bar = 60 pm. 10. Korea, large adult specimen (lateral denticles worn). Scale bar = 150 pm. 11. Operculum of E. martensi Scale bar = 1 mm. 12. Operculum of E. junionae. Scale bar = 1 mm. 13. Protoconch of E. martensi. Scale bar = 0.5 mm. (Figures 6-11, photos A. Warén). Page 102 THE NAUTILUS, Vol. 122, No. 2 i 16 Figures 14-17. Ergalatax species. 14. Ergalatax margariticola (Broderip, 1833). Lectotype of Pentadactylus (Morula) martensi Schepman, 1893. ° ‘Red Sea, coll. Forskal”, ZMA Moll. 2 93.005, 24.8 mm. 15-16. Er: galatax junionae nom. noy. Lectotype of Morula martensi Dall, 1923, ZMB 21596, 21.4 mm. 17. Ergalatax recurrens Iredale, 1931. Holotype AMS C€57761, 25.5 mm (photo courtesy IE. H. Vokes). creamy white, or tan, usually with some light to dark brown colored spiral cords on shoulder dad on more prominent nodes. Aperture cream or pale yellow within. Operculum ergalataxine (Fig. 11), D- shaped with lat- eral nucleus in lower right. Radula (Fig. 6) with a rachidian bearing a long, narrow central cusp, a small, narrow, triangular ace, denticle anda broad, long lateral cusp on each side. Sickle-shaped lateral teeth broad at base and narrow at their end. Type Material Examined: 3 syntypes NMNL, Red Sea: RMNH.MOL.57165, here selected as lectotype and paralectotypes, 1 syntype ZMA Moll. 2.93.005, Red Sea, here selected as paralectotype; Ergalatax obscura Houart, 1996: Perim, Strait of Bab a Mandeb, holo- type and 4 paratypes MNHN, | paratype RH; Djibouti, Obock, Gulf of Aden, 9 paratypes MNHN; Yemen, Aden, 4 paratypes MNHIN, 1 paratype RH. Othe Material Examined: — Djibouti: Obock, Gulf of Aden, 2 RH; Gulf of Aden: 2, RH. Red Sea: (no other data), 1 RH; Massawa, Taulud Is., 1 RH; Egypt, Sinai, under stones at low tide, 10 RH; Egypt, Sinai, Sharm E] Sheik, 1 RH; Egypt, Sinai, Shark’s Bay (marsa umm mureihha), under stones, low tide, 2 RH. Gulf of Aqaba: Israel, Eilat, 0.5-1.0 m, 3 RH: Gulf of Aqaba, Israel, Eilat, under stones, 1-2 m, 4 RH. Distribution: Aqaba 0-2 m, on and under stones (Fig. 46). Remarks: For a comparison with Ergalatax junionae e that species below. Ergalatax junionae nomen novum Figures 2, 4, 7, 12, 15-16, 31-40, 46) From the Gulf of Aden to Eilat, Gulf of Morula siderea Reeve.—von Martens, 1874: 95, pl. 5, fig. 49 (not Ricinula siderea Reeve, 1846). Morula martensi Dall, 1923: 304, new name for Morula siderea von Martens, 1874, not Reeve, 1546. Cronia konkanensis.—Bosch and Bosch, 1982: 95, text fig; Smythe, 1982: 60, pl. 1, fig. i; Bosch and Bosch, 1989: 60, text fig. (not Ricinula konkanensis Melvill, 1893). Cronia cf. konkanensis—Giunchi and Tisselli, 1995: 8, text figs. Ergalatax martens.—Buzzurro, Engl and Tiimtiirk, 1995: (no pag.), text fig.; Engl, 1995: 46, fig. 10. Cronia cf. hovieonensitBocdh et al, 1995: 121, fig. 480, Morula martensi (Schepman, 1$92)—Tan, 1995: 160, figs 52, 192 ¢, h (not Pentadactylus (Morula) martensi Schepman, 1892). Ergalatax obscura.—Houart, 2001; 108 (in part), figs. 450-451 only; Delongueville and Scaillet, 2007: 57, fig. 31 (not Ergalatax obscura Houart, 1996). Description: Shell medium sized for the genus, up to 29 mm in length at maturity. LengthAvidth ratio 1.91- 2.03, Slender, lanceolate, heavy, nodose. Shoulder weakly sloping, concave. White or creamy white with dark brown or blackish colored primary spiral cords and occasionally s2 or s3. Aperture glossy white. Spire very high with 3.5 protoconch whorls and teleo- conch up to 7 weakly convex, strongly shouldered, no- dose whorls. Suture adpressed. Protoconch a coni- cal, acute; terminal lip raised, of sinusigeral Axial sculpture of teleoconch whorls ee vof high, strong, broad, nodose ribs and erratically placed strong varices. Last teleoconch whorl with 8-11 ribs, oceasion- ally with one or two erratically placed, broad varices. Spiral sculpture of high, strong, nodose and squamose R. Houart, 2008 Page 103 Figures 18-30. Ergalatax martensi (S¢ hepman 1892). 18-23. Lectotype and paralectotypes RMNH.MOI 97 165 photo | Goud). 18-19. 17.4 mm; 20-21. 15.7 mm; 22-23. 17.3 mm. 24-25. Holotype ot E. obscura Houart, 1996. Perim, Strait of Bab el Mandeb, MNHN 0159, 24.2 mm photo MNHN 26. Eilat, Israél, RH, 20.8 mm; 27. Obock, Gulf of Aden, RH, 21.6 mm; 28. Eila Israél, RH, 25.1 mm; 29. Red Sea (no other data), RH, 21.4 mm; 30. Aden, Gulf of Aden, RH, 21 mm Page 104 THE NAUTILUS, Vol. 122, No. 2 Figures 31-45. Ergalatax species. 31-40. Ergalatax junionae nomen novum, 31-33. Doha, Wahra, Qatar, RH. 31-32. 26.5 mm; 33-34. 22.9 mm: 35. Yalikent, Iskenderun, Turkey, RH, 17.2 mm; 36-37. Lebanon, Bay of Jounich, 19.6 mim; 38-40. Iskenderun, Bay of Iskenderun, Turkey, RH. 38. 25.6 mm; 39. 22.2 mm; 40. 22.9 mm, 41-45. Ergalatax margariticola (Broderip, 1833), 41. 927 Rar Pumaotu Archipelago, RH, 28.4 min; 42. Kai Is., Mollucas, RH, 20.5 min, 43, South of New Caledonia, RH, 23.7 mm, 44. Beach, East coast of Singapore, RH, 30.4 min; 45. Kwajalein Atoll, RH, 25.9 mm R. Houart, 2008 primary, secondary and tertiary cords. Shoulder of last teleoconch whorl with broad SP, adis, IP, abis, P1, P2, s2, (t), P3, s3, P4,(s4), P5, (s5), P6, ADP, MP: Pl and P2 narrow, P3, P5 and P6 broad, similar in strength, P4 smaller. Aperture large, narrow, ovate; columellar lip smooth, entirely adherent, with low parietal tooth at adapical ex- tremity; anal notch oe broad; outer lip weakly erect, with ID, D2-D5. ID] argest, broad; D2-D4 decreasing i in strength abapically, D5 split; denticles elongate within aperture. Siphonal canal short, broad, dorsally recurved, broadly open. Operculum (Fig. 12) dark brown, D-shaped, with lat- eral nucleus in lower right; attached surface with about 8 growth lines and broad, callused rim, about 30-40 % of opercular width. Radula (Fig. 7) with a rachidian bearing a long, slender central cusp, a small, short lateral denticle and a broad, long lateral cusp on each side. Lateral teeth sickle- shaped, with broad base and narrow end. Type Material Examined: Morula martensi Dall, 1923, lectotype (selected by Houart, 1996) and 6 syn- types ZMB 21596. Other Material Examined: Persian Gulf: Kuwait, Kuwait City, 4 RH; Qatar, Doha, under rocks, 30 RH; Sharjah. 25°20" N, 55°21’ E, on rocks, 4 RH; Abu Dhabi, 0.5-1.0 m, 3 RH. Gulf of Oman: Al Hamra, near Qurm, 10 km NW ‘of Muscat, 16 RH; Al Bustan, under rocks, 1 RH. Lebanon: Beirut, harbour entrance, breakwater, max. 15 m, 2 RH: Bay of Jounieh (N), 10-25 m, 2 RH; Batroun, 1-4 m, under stones, 2 RH. Turkey: Gulf of Iskenderun, Iskenderun, under rocks, harbor, 1 m, 8 RH; Kale, beach, 6 RH; Yumurtalik, rock pools, 6 RH; Ya- likent, shallow water, on rocks, 1 RH; Bay of Antalya, 6 km off Kemer, 36°39’ N, 30°33’ E, on rocks with mussel banks, 0.5-1.0 m, 7 RH; Fethiye-Oludeniz, on rock at 2 m, | RH. Distribution: Gulf of Oman, Persian Gulf and eastern Mediterranean Sea, intertidal to 4 m, on and under rocks and stones (Figure 46). Etymology: This species is named in honor of Mrs. Marie-Louise Buyle-Junion (1916-2003), former librar- ian of the Belgian Malacological Society. She was one of the mainstays of the Society, together with her husband, Jean Buyle. Remarks: Ships docking at oil terminals in the Gulf of Iskenderun (eastern Turkey) could have introduced the species into the eastern Mediterranean Sea. As noted by Delongueville and Scaillet (2007), the transport via ship hulls or ballast water may be suspected. The shell morphology of some specimens of E. junio- nae (Fig. 35) is nearly the same to E. martensi (Fig. 30), however E. martensi differs in having a more strongly shouldered, broader shell, and a yellowish aperture in- stead of white, with 7 denticles within instead of 6 in E. junionae. Moreover, the spiral cords differ in number and strength, as described above and illustrated in Fig- Page 105 50_0 Ses a ; ’ “4, ; 9° 2 FY Cupp. O Yo, oO ikea mo \e ° “ “e@@ Aden ® Gos of” @ Ergalatax martensi (Schepman, 1893) ° O Ergalatax junionae nomen novum — Ps Fig. 46. Distribution of Ergalatax martensis and E. junionae. ures 2-5. Twenty-five specimens of E. martensi and a few more of E. junionae were examined to confirm the stability of these differences. Ergalatax martensi usually has 6-8 axial ribs on the last teleoconch whorl, rarely 5 or 9, compared to E. junionae which bears 8-11 ribs and varices on the last whorl. Ergalatax margariticola (Figures 8, 14, 41-45), a very common Indo-West Pacific species is also related to E. junionae, however it is generally stouter and broader, with a wider shoulder, a more squamose spiral sculpture, more uniformly colored shell and different aperture color, being bluish-white, occasionally with a tinge of pink or mauve on the columellar lip vs. completely white in E. junionae. The shell morphology and color of E. margariticola are highly variable, however it is always easily distinguishable from E. junionae by one or more differences cited above. The three species are related to the ergalataxine Er- galatax contracta (Reeve, 1845) (Figures 9-10), a prob- ale senior synonym of Ergalatax recurrens Iredale, 1931 (Fig. 17), the type species of Ergalatax. Buzzurro, Engl and Tiimtiirk (1995) were the first to mention the presence of Ergalatax junionae [as Erga- latax martensi (Dall, 1923)] in the eastern Medien: nean Sea. Tan (1995:147) in his Ph D. thesis also selected a lec- totype for Morula martensi Dall, 1923. However, he des- ignated a specimen of Ergalatax margariticola (Tan, in litt.) from Kingsmill Is (USNM 52472). Nevertheless, this designation being published in a thesis that does not satisfy Articles 8.1.2 and 8.1.3 of the International Code of Zoological Nomenclature, it is not available as such. ACKNOWLEDGMENTS I am very grateful to Jeroen Goud, National Museum of Natural History Naturalis, Leiden for the digital images Page 106 THE NAUTILUS, Vol. 122, No. 2 of the type material of Pentadactylus martensi and for his comments, to David Reid, Natural History Museum, London, for providing me useful photocopies and for comments, to Robert G. Moolenbeek, Zoologisch Mu- seum, University of Amsterdam for the loan of the lec- totype of P. martensi, to the late Prof. R. Kilias, Museum fiir Naturkunde der Humboldt Universitat zu Berlin, for the loan of the specimen illustrated by Martens (1874), lectotype of Morula martensi Dall, to Anders Warén, (Natural History Museum, Stockholm) for radula prepa- ration and SEM work of radula and operculum, to E. H. Vokes (Prof. Emeritus, Tulane Univ ersity, Louisiana) for the photograph of E. recurrens Iredale, 1931, to Greg Herbert (University of South Florida, Tampa, USA) for his useful comments and suggestions, and to John Wolff, Lancaster, Philadelphia, for checking the English text. LITERATURE CITED Bosch, D. and Bosch, 1982. Seashells of Oman. Longman Croup, England: an pp. Bosch, D. and Bosch, E. 1989. Seashells of Southern Arabia. Motivate, Dubai: 95 pp. Bosch, D. T., Dance, S. P., Moolenbeek, R. G. and Oliver, P.G. 1995. Seashells of Eastern Arabia. Ed. P. Dance, Motivate Publishing: pp. 1-296. Buzzurro, C., Engl W. and Tiimtiirk, I. 1995. Bivalven und a n der Europiiischen Meere (4): Ergalatax mar- tensi (Dall, 1923) (Muricidae), Ein neuer Lesseps’schere Einwanderer von der Tiirkischen Siidkiiste. Club Con- chylia Informationen 27(1): 17-18. Coulombel, A. 1994. Coquillages de Djibouti. Edisud, La Calade, Aix-en-Provence: 143 pp. Dall, W. H. 1923. Notes on Drupa and Morula. Proceedings of 303— the Academy of Natural Sciences of Philadelphia 75: 306. Delongueville, C. and Scaillet, R. 2007. Les espéces invasives de mollusques en Méditerranée. Novapex 8(2): 47-70. Engl, W.1995. Specie prevalentemente Lessepsiane attestate lungo le coste Turche. Bolletino Malacologico 31(1-4): 43-50, Giunchi, L. and Tisselli, M. 1995. Cronia cf. konkanensis (Melvill, 1893), new Indo-Pacific host in the Mediterra- nean Sea. La Conchiglia 27(275): S—9. Heiman, E. L. and Mienis, H. K. 2003. Shells of East Sinai, an illustrated list. Muricidae (2), Triton: 22-23. Houart, R. 1995, The Ergalataxinae (Gastropoda, Muricidae) from the New Caledonia region with some comments on the subfamily and the description of thirteen new species from the Indo-West Pacific. Bulletin du Muséum national d Histoire naturelle, Paris, 4e sér., 16A(2-4):197—245. Houart, R. 1996. On the identity of Morula martensi Dall, 1923 and description of a new species of Ergalatax from the Red Sea (Gastropoda: Muricidae: Ergalataxinae). The Nautilus 110(1): 12-16. Houart, R. 2001. A review of the Recent Mediterranean and Northeastern Atlantic species of Muricidae. Evolver: pp. 1-227. Martens, E. von 1874. Ueber Vorderasiatische Conchylien nach den Sammlungen des Prof. Hausknecht. Cassel, T. Fischer: 127 pp, 9 pl. Merle, D. 1999. La radiz den des Muricidae (Gastropoda; Neo- g gastropoda) au Paléogene: approche phylogénétique et évolutive, Thése du Muséum national d'Histoire naturelle, Paris: 499 pp. Merle, D. 2001. The spiral cords and the internal denticles of the outer lip of the Muricidae: terminology and method- ological comments. Novapex 2 (3): 69-91. Schepman, M. M., 1892. Note VI. Two supposed new species of Pentadactylus. Notes from the Leyden Museum, Vol. XV: LOS—104. Sharabati, D. 1954. Red Sea Shells, KPI, London: 128 pp. Singer, B.S. and H. K. Mienis. 1991a. Shells of the Red Sea. The family Thaidididae (sic) (1). La Conchiglia 27(260): 16-19. Singer, B.S. and H. k. say 1991b. Shells of the Red Sea. The family Thaididae . La Conchiglia 27(261): 54-60. Smythe, K. 1982. Seashe ils of es Arabian Gulf. George Allen & Unwin, London: 122 pp. ; Tan, K-S. 1995. Taxonomy of Thais and Morula (Mollusca: Gastropoda: Muricidae) in Singapore and vicinity. Ph.D. thesis, National University of Singe apore. Verbinnen, G. and Dirkx, M. 2000. Red Sea Mollusca, Part 6. Gloria Maris 38(4—-5): 64-76. THE NAUTILUS 122(2):107-114, 2008 Page LOT Two new deep-sea muricids (Gastropoda) from Argentina Guido Pastorino o Argentino de Cie ea Naturales Angel Gallardo 470, ° piso lab. 57 Cl40sDIR Buenos ives: ARGENTINA — Antropologia gpastorino@macn.gov.ar CoG, 399 =.C.'P. Fabrizio Scarabino Direccion Nacional de Recursos Acuaticos and Museo Nacional de Historia Natural y 11.000 Montevideo, URUGUAY fscara@gmail.com ABSTRACT Two new species of muricids belonging in the genus Trophon are described from the upper slope off the Atlantic coast of Argentina. Both species have a small size for the genus. The radulae show similarities with those of Antarctic species of the same genus. Trophon columbarioides new species has a smooth shell with spines pointed adapically and was collected at 37-38° S, in 209-382 m. Trophon fasciolarioides new species has prominent spiral cords and was collected at Burwood Bank in 286-292 m depth and off Bahia Blanca in ca. 1000 m depth. Additional Keywords: Mollusca, Muricidae, Trophon, South- western Atlantic, Gastropods, Taxonomy, Patagonia INTRODUCTION The genus Trophon includes a group of species of un- doubtedly austral origin. The older species can be traced as far back as the Oligocene, from Patagonian deposits (Griffin and Pastorino, 2005). Pastorino (2005) re- described all known living species of Trophon from southern South America, and also some new species be- longing in the genus. While the subfamilial affinities of this genus are discussed by several authors (e.g. Kool, 1993, 1993a: Vermeij and Carlson, 2000, among others), the genus is firmly established for all fusiform and/or lamellate gastropods—usually with spiral ornamenta- tion—tfrom the southwestern Atlantic. Two of the species characterize the shallow waters along the Patagonian coast, i.e., Trophon geversianus and T. plicatus. Both are sympatric in the southern part of Argentina; however only T. geversianus could be collected intertidally as far North as Buenos Aires province. All the other species are mostly subtidal. Trophon species, as far as it is known, are all predators. feeding on the mussel banks and bar- nacles that are fairly common along the coast of the southern part of South America. Living in moderately deep waters are Trophon acan- thodes Watson, 1882, and the recently rediscovered T. clenchi (Carcelles, 1953), a rare lamellose and spiny spe- cies originally assigned to the genus Murex (Pastorino, 2005). Both of them undoubtedly belong in Trophon. In addition, Houart (1991) and Pastorino (1999) described Trophon mucrone and T. veronicae from deep waters off South America. The two species were included in Tro- phon despite some minor but distinctive differences in radulae, protoconchs and penises. Recently Houart (2003) and Houart and Sellanes (2006) described new species from deep waters off Chile. In this paper two new deep water species from the southwestern Atlantic are described and compared with the other related ones living around the region. MATERIALS AND METHODS Specimens of T. columbarioides new eae studied herein were collected by one of us (FS) on board the Uruguayan R/V ALDEBARAN. The other specimens are from the 2002 cruise to Antarctica of the German R/V POLARSTERN. Additional material was collected by Uru- guayan fishing boats, Dissections were performed on ec ethanol-preserved specimens to study radulae and male reproductive system when it was av ailable. Radulae were cleaned with commercial bleach and ultrasound, and observed using a Philips XL 30 scanning electron microscope (SEM) at the Museo Argentino de Ciencias Naturales (MACN). Critical point drying of the penises (when available) was prepared at the MACN. Radular terminology follows Kool (1993: fig. 6B). Shell photo- graphs were taken using a digital camera. All images were digitally processed. The material is hagsed at the MACN one the Museo Nacional de Historia Natural y Antropologia, Montevideo (MNIINM). SYSTEMATICS Class Gastropoda Cuvier, 1797 Order Neogastropoda Wenz, 1935 Family Muricidae Ratfinesque, 1515 Subfamily Trophoninae Cossmann, 1903 Genus Trophon Montfort, 1810 Type Species: Murex mage llanicus Gmelin, 1791, Trophon geversianus ( (Pallas, 1774) by original designa- tion. Page LOS THE NAUTILUS, Vol. 122, No. 2 Trophon columbarioides new species (Figures 1-11, 15-18) Diagnosis: Shell very small, thin, fusiform, chalky; axial ornamentation of 30-35 regular axial lamellae grow- ing along the entire whorl surface, attached to the ‘shell, producing open, long, regularly spaced spines along pe- riphery, pointing upwar ds. Siphonal canal very long. Description: Shell small (up to 14 mm), slender, thin profile, chalky, white, somewhat bright; protoconch of two whorls, slightly globose, symmetrical: teleoconch of five oblique, shouldered whorls: spire height less than 4 of total shell height. Spire angle about 40°: suture im- pressed; subsutural shelf well defined, oblique; aperture semicircular, interior chalky white; anterior siphonal ca- nal very long (longer than aperture height), narrow, straight, open; outer lip sharp, rounded, inner lip ad- pressed. Axial ornamentation of 30-35 regular axial lamellae growing along the entire whorl surface, but at- tached to shell and producing open, long, regularly spaced spines along periphery (almost in the middie of the whorl), pointing adapically; last three whorls with ten lamellae each . A second obsolete series of spines ap- pears to rise at the periphery of older specimens (Figure 1). Spiral omamentation lacking. Growth lines present throughout shell. Operculum subtriangular, nucleus terminal. External surface covered by concentric, irregular, growth lines. Inner surface attachment area reaching upper side or center, with horseshoe-shape scars (Figures 17-18). Rachidian teeth of radula with thin, small central cusp; lateral cusps wider and larger than central cusp; denticle between central and aeenl cusp rising from base. Base of rachidian tooth slightly curved. Lateral teeth with single, long cusps; attachment area thick (Figures 15- 16). Type Material: Holotype MACN-In 37380 (Figures 1-3, 7, 9-10) and two paratypes, MACN-In 37381 (Fig- ures 4-6, 8) and MNHNM 15540 (Figure 11). Type Eason R/V ALDEBARAN cruise 2003/01, sta- tion 37, 37°43’ S, 55°00" W, 209 m, October 26 2003, 5.3°C of bottom : ne rature, Piccard dredge; (holotype aad one paratype); between 37°05" S, 54°12! W in 255 m and 37°02’ S, 54°02’ W in 382 m (one paratype). Etymology: The general shell morphology reminds some species of the genus Columbarium (Caenogas- tropoda: Turbinellidae), to which it has no close relation- ship. Distribution: Known only from three specimens from the type locality and vicinity. Remarks: At first glance the general morphology of the shell shows some similarities with juveniles of Tro- phon acanthodes Watson, 1882. However the typical spiral cords of the latter appear early in ontogeny (see Figures 12-13) and are completely absent in the new species. In addition both species have clear radular dif- ferences (see Pastorino, 2005:69). In addition, T. plicatus and T. clenchi are comparable species. The latter has unmistakable early developed spiral ornamentation; the former has complete lamellae and a shorter siphonal ca- nal, besides radular differences. Despite the geographic distance separating them, Tro- phon scolopax and T. septus described by Watson (1882) are comparable species. They live around Kerguelen Is., in the southernmost Indian Ocean. As a main diferente Trophon columbarioides new species has a higher spire and only one series of large and open spines pointing upwar ds, instead of the three series shown by T. scolopax or the triangular upturned ones of T. septus. Trophon echinatus (Kiener, 1840), an extremely vari- able species (according to Bouchet and Warén, 1985: 141), from deep waters off Northeastern Atlantic and Mediterranean shows a remarkable shell similarity with T. columbarioides new species Despite this, we do not support phylogenetic affinities between these and we made the comparison ist for showing the existence of specific contrasting differences, particularly at radular level. As far as we can see in the material available there are no signs of spiral cords in the new species which are common in the deeper specimens of the northern one. However, some porcellaneous (not chalky) smooth speci- mens are in fact comparable. All morphs of T. echinatus have a shorter protoconch. Some grown specimens of T. columbarioides shows the apparently presence of a sec- ond rows of spines while T. echinatus has only one. In addition, the radulae (illustrated by Bouchet and Warén, 1985 figs. 333, 335, 336) shows a pair of almost obsolete intermediate denticles between the lateral cusps while in T. columbarioides new species they are slightly smaller than the central and lateral cusps. The base of the rachid- ian is sinuous in T. echinatus and somewhat straight in the new species. The intermediate denticles of the rachidian teeth rise from the internal side of the lateral cusps in a way that is typical of the southwestern Atlantic Trophon species. The attached portion of the marginal teeth are also different. Houart (2001) considered T. echinatus as belonging in to the genus Pagodula Monterosato, 1884 despite the differences that Bouchet and Warén (1985) pointed out with the protoconch of the type species of Pagodula: the Pleistocene species Murex vaginatus Cristofori and Jan, 1832. Figures 1-11. view of the par: atype 9-10. Protoconch of the holotype, scale bar = Trophon columbarivides new species . 1-3. Holotype, MACN-In 37380, us ALDEBARAN cruise 2003/01, station 37, 37°43" S, 55°00! W in 209 m. 4-6. Paratype, MACN-In 37381. Same locality as holotype Linm. 11. Par: itype, MNHNM 15540, between 37°05! S . Apical view of the holotype. 8. Apical 54°19 W in 255 m and 37°02" S, 54°02’ W in 382 m. 12-14. Trophon acanthodes Watson, 1882, MACN-In 25165-2, 37°35’ S, 54°55’ W, 192 m. Seale bar = 1 em for all figures except 9-10. G. Pastorino and F. Scarabino, 2008 Page 1LO9 Page 110 THE NAUTILUS, Vol. 122, No. bo Figures 15-18. Unfortunately both dissected specimens of the new species here described were females so nothing can be said about the morphology of the penises. Trophon fasciolarivides new species (Figures 19-31) Diagnosis: — shell very small, thin, translucid or chalky; very weakly develope :d axial ornamentation of regular, very low, varices attached to the shell. Spiral ornamen- tation of 2, 3 and § or 9 spire al rounded cords in the second, ae and last whorl respectively; cords of similar width; intersection of spiral cords with axial varices con- veying a slightly cancellate appearance to shell surface Description: profile translucid white or chalky; protoconch of 2 vhorls, symmetrical; teleoconch of 412 tabular whorls: spire height less than 5 of total shell height. Spire angle shell small (up to 12 mim), slender, thin about 45°: suture impressed, subsutural shelf very Trophon columbarivides new species. 15. Radula frontal view, scale bar = 10 jm. 16. Radula lateral view, scale bar= 20 wm. 17. Operculum of the holotype and, 18. Paratype in figs. 4 ~6. Scale bars= 1 mm. oblique; aperture suboval; anterior siphonal canal long but never longer than aperture height, slightly curved adaxially, open; outer lip sharp, rounded, inner lip ad- pressed, Axial ornamentation of poorly developed va- rices, regular, very low, attached to the shell, growing along the entire whorl surface except the siphonal canal. Spiral ommamentation of 2, 3 and 8 or 9 spiral rounded cords in the second, third and last whorl respectively; cords of similar width; intersection of spiral cords with axial varices conveying a slightly cancellate appearance to shell surface; growth lines present throughout shell, be- coming scaly at intersection with spiral cords. Operculum suboval, nucleus terminal. External sur- face covered by concentric, irregular, extremely thin growth lines. Inner surface attachment area reaching up- per side or center, with horseshoe-shaped scars (Figure 31) Rachidian teeth of radula with thin central cusp, higher than laterals; denticle between central and lateral G. Pastorino and F. Scarabino, 2008 Figures 19-26. Trophon fasciolarioides new species 19-21. Holotype MACN-IN 37382. Banco Burwood, Polarstern St. 150 between 54°30.22' S, 56°08.58’ W in 286 m and 54°29.64' S 56°08.09’ W in 292 m. 22. Paratype MACN-IN 37383, same locality as holotype. 23-24. Protoconch of the paratype of figure 22, arrow head transition to teleoconch, scale bar = 1 mm. 25-26. Paratype MACN-In 37354, off Bahia Blanca ca. 1000 m depth Seale bar = 5 mm for all figures except 23 and 24 cusp thin and long, rising from the base. Base of rachid- ian tooth curved. Marginal area with single cusp. Lateral teeth with single long cusps attachment area thick Fig- ures 27-29 The penis shows an unusual morphology among the Patagonian species of the genus: it is wide slightly curved and laterally flattened with a small slit at the tip Figure 30 Type Material: Holotype MACN-IN 37382 and two paratypes MACN-IN 37383 and MACN-In 37384 Type Locality: Banco Burwood. Polarstern station 150, 6 Apr. 2002, AGT net, between 54°30.22' S 56°08.58' W in 286 m and 54°29.64' S$, 56°0S.09’ W in 292 m. Etymology: The general shell morphology reminds some species ol the genus Fasciolaria (Caenogastropoda Fasciolariidae Distribution: known from type locality and approxi mately off Bahia Blanca in ca. 1,000 m depth (MACN-In 37384 Page 112 THE NAUTILUS, Vol. 122, No. 2 Figures 27-31. Trophon fasciolarioides new species. 27. Radula of a paratype, MACN-In 37354, frontal view, scale bar = 40 jum. an 28. Lateral view of the same radula as 27, scale bar = 50 ym, 29. Radula of the paratype MACN-In 37353, scale bar = 20 um. 30. Critical point dry of the penis, scale bar = 500 jzm. 31. Two views of the operculum of the paratype of figures 2 lL mm. Remarks: The morphology of the shell and the radula match some of the Patagonian and Antarctic species of the genus Trophon. From the first group, T. ohlini Strebel, 1904, a rare species from the Magellanic area, has a similar profile. However, its distinctive protoconch points out a clear difference. In addition, the morphology of the penis is far from the typical Patagonian represen- tatives of the genus. Trophon emilyae Pastorino, 2002, T. declinans Watson, 1882, and T. cuspidarioides Powell, 1951, are comparable Antarctic species (Pastorino, 2002a). The first two species differ from the new species in having well developed axial sculpture only. Also T. cuspidarioides has 5} » whorls including the protoconch anc five blunt spiral cords in the body whorl, while the new species has a larger spire and 62 whorls, and 8 or 9 26, scale bar = rounded and well defined cords in the last whorl. The axial ornamentation in the new species is very irregular but higher in number than the 25 axials present in T. cuspidarioides, DISCUSSION In previous papers two groups of species belonging in the genus Trophon—i.e., from South America and Antarc- tica—were pointed out. The main differences between these two groups are several radular and anatomical fea- tures (Pastorino, 2002b, 2005). The shell morphology of the two new species described herein agrees with that of Trophon living off the South American coast. However, G. Pastorino and F. Scarabino, 2008 Page 113 Golfo San Jorge © S = xv | Ss GB , Malvinas Is. | “Faas o- | Staten Is. 70 Figure 32. | Map showing the type localities of Trophon columbarioides new species @ (filled circle) and Trophon fas- ciolarioides new species [(_] (blank square). the radula has some features found in the Antarctic rep- resentatives of the genus—albeit the taxonomic status of the Antarctic species still needs confirmation. Like in the Antarctic group of species, the central cusp of the rachid- ian teeth is shorter and thinner than the laterals, the internal denticle rise from the base of the rachidian in- stead of the upper third of the internal side of the lateral cusp, and the lateral teeth have a wide attachment area. The geographic ranges of species of these two groups do not overlap. Therefore. the morphological features seem to be clearly separated geographically too. How- ever. in the species described herein, this seems not to be the case. While there is no geographic overlapping of species. the morphological features that characterize the Antarctic group do appear in these two new Patagonian species. Contrarily, this does not occur in the case of Patagonian species, the morphological features of which are restricted to South America. Moreover, they are bet- ter developed in species from the Magellanic among those described from southern South America (see Pas- torino, 2005). Recently, Houart (2003) introduced three new species he assigned to Trophon from off Chile: T. ceciliae; T. condei aad T. vangoethemi. The morphology of the radula, only known for the last two species, together with that of T. mucrone Houart, 1991, T. veronicae Pastorino, 1999 and both new species described here, is that of the Antarctic group, despite the geographic distance sepa- rating them. All mentioned species were collected from more than 300 m depth and in most cases they reach more than 1,000 m (see Table 1). Most of the Patagonian species of Trophon were collected alive from the inter- tidal zone to about 300 m depth. Exceptions are some specimens of T. acanthodes, T. clenchi, and T. bahamon- dei that come from deeper areas. However, there is no Antarctic species—or with their characteristic radular and anatomical features—that live at shallower depths or closer to the continent. Both new species described here were collected from about 300 m depth, which is the shallowest for a Trophon with Antarctic features at South American latitudes. Generic assignment in the whole Trophoninae group is actually under revision. The concept of the genus Tro- phon sensu stricto is easy to apply to the shelf species. However, when deeper species are studied, the presence of several features in common among the shallower spe- cies appear somewhat contradictory. Nevertheless, we think that biogeographic boundaries are hard to surpass for species without free larvae as both new species ap- pears to be according to the shape and whorl number of the protoconch. Contrasting historical biogeographic processes must be considered when suggesting affinities between species and therefore comparison with species living far away from the Southwestern Atlantic area can be considered an empty exercise. Some exempli are cases like T. columbarioides new species—T. echinatus which have very few characters that clearly split both species. In Table 1. Recently described South American species of Trophon. Depth (m) Type locality T. condei Houart, 900-1350 Ancud, Chile 2003 T. ceciliae Houart, 434-1000, 1300 Antofagasta, Chile 2003 Itata, North of Concepcion, Chile T. vangoethemi About 350 Houart, 2003 T. mucrone Houart, 790-1575 Off Rio de 1991 Janeiro, Brazil T. veronicae Pastorino, 298-1272 Subantarctic 1999 T. columbarioides 209-382 37°43'S, 55°00'W. new species T. fasciolarioides 286-292 Burwood bank new species Page 114 THE NAUTILUS, Vol. 122, No. 2 the future molecular characters could add to the under- standing of the evolution of morphological features in this group. ACKNOWLEDGMENTS We are grateful to L. Paesch and R. Bird (DINARA) for facilities in the collecting of Trophon specimens, as well as to J. L. Viggiano and H. Racz-Lorenz (Monte- video) for the material provided. We thank Miguel Grif- fin (UNLPam) who helped to improve the English ver- sion of the manuscript. R. Houart (Belgium) reviewed this paper; his comments and discussions undoubtedly contributed to enhance the original version despite dis- agreement on some points. This contribution was sup- ported in part by the project PICT No, 03-14419 from the National Agency for Scientific and Technical Promo- tion, Argentina. We acknowledge funding by the Consejo Nacional de Investigaciones C ientificas y Técnicas (CONICET) of Argentina, which sponsors the research of G.P LITERATURE CITED Bouchet, P. and A. Warén. 1985. Revision of the northeast Atlantic bathyal and abyssal Neogastropoda excluding Tur- ridae (Mollusca, Gastropoda). Bolletino Malacologico supplemento 2: 123-296. Carcelles, A. 1953. Nuevas especies de gastropodos marinos de las Reptiblicas Oriental del Uruguay y Argentina. Comu- nicaciones Zooldgicas del Museo de Historia Natural de Montevideo, 4(70):1—18. Griffin, M. and G, Pastorino, 2005. The genus Trophon Mont- fort, 1810 ( Gastropoda: Muricidae) in the Tertiary of Pa- tagonia. Journal of Paleontology 79: 296-311. Houart, R. 1991. The southeastern Brazilian Muricidae col- lected by R V Marion-Dufresne in 1987, with the descrip- tion of three new species. The Nautilus 105; 26-37. Houart, R. 2001. Ingensia gen. nov. and eleven new species of Muricidae (Gastropoda) from New Caledonia, Vanuatu, and Wallis and Futuna Islands. In: P. Bouchet and B. A. Marshall (eds.) Tropical Deep-Sea Benthos, volume 22. Mémoires du Muséum national d’Histoire naturelle 1S5: 243-269. Houart, R. 2003. Description of three new species of Trophon s. |. Montfort, 1810 (Gastropoda: Muricidae) from Chile. Novapex 4: 101-110. Houart, R. and J. Sellanes. 2006. New data on recently de- scribed Chilean trophonines (Gastropoda: Muricidae), de- sc ription of anew species and notes of their occurrence at a cold seep site. Zootaxa 1222: 53-68. Kool, S. P. 1993. Phylogenetic analysis of the Rapaninae (Neo- gastropoda: Muricidae): Mal: wcologia 35: 155-260. Kool. S. P. 1993a. The systematic position of the genus Nucella (Prosobranchia: Muricidae: Ocenebrinae): The Nautilus 107: 43-57. Monterosato, A. di, 1554. Nomenclatura generica e specifica di alcune conchiglie Mediterranee. Palermo, 152 pp. Pastorino, G. 1999. A new species of gastropod of the genus Trophon Montfort, 1810 (Mollusca: Gastropoda: Muri- cidae) from subantarctic waters. The Veliger 42: 169-174. Pastorino, G. 2002a. Two new Trophoninae (Gastropoda: Mu- ricidae) from Antarctic waters. Malacologia 44: 353-361. Pastorino, G. 2002b. Systematics and phylogeny of the genus Trophon Montfort, 1810 (G astropoda: Muricidae) from Patagonia and Antarctica: morphological pattems: Bollet- tino Malacologico 38; 127-134. Pastorino, G. 2005, A revision of the genus Trophon Montfort, 1810 (Mollusca: Muricidae) from southern South America: The Nautilus, 119: 55-82. Powell, A. W. B. 1951. Antarctic and subantarctic mollusca: Pelecypoda and Gastropoda. Discovery Reports, 26: 47— 196. Vermeij, G. and S. Carlson, 2000. The muricid gastropod sub- family Rapaninae: phylogeny and ecological history. Pa- leobiology 26; 1946. Watson, R. B. 1882. Mollusca of H. M.S. ‘Challenger’ Expedi- tion. Part 13. The Journal of The Linnean Society. Zool- ogy, 16: 358-392. Sponsored in part by the State of Florida, Department of State, Division of Cultural Affairs, the Florida Arts Council and the National Endowment for the Arts pRI> ¢ S XQ e) aa 9 2 % v ce NATIONAL ENDOWMENT FOR THE ARTS gH INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes articles on all aspects of the biology, Baleontolenie and systematics of mollusks. Manuscripts describing original, unpublished research and review articles will be Re Gaara Brief articles, not exceeding 1000 words, will be published as notes and do not require an abstract. Notices of interest to the mala- cological community will appear in a notices section. 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" i SY We VAAN, i ny ) cn : ie ; th aM ! nn bisnyy the ne ne 7 ° va i" ae i i ie Wa if ae Ie ign a ne ur i if BG i a i, hans rs ont his Ash a yi on ae on pa i ie WAL: ‘ le | eu ye oy NY | etter: 1 o i y pe if VEL ia he | "{ a i ne if ' Mi 7 th i Lae» ary i nig haa ite a) 14 eh Ay ‘ , uae PASSED Ae Hye Ab wilt ey ital ! ey) Wi oi i a ee ue co y Bay ls eh Ke Da . 7s mae . +i mt wi | ——— = ee 2 = -_ — 5 — — ca H ; mnt & ae | ‘ a Ml ite a ; Ne Vi : 1 oa es og i i sf The (ha i> Ph Be j a Te ey Le Ww ; idl | VR i j f i Sf if | ' Abas | Ayes heed || BY Lye ne fi) i net i De, er 82 ey gy Jib. as hy 0 15 wae ea At’ y iy ee ne iy i “? mal ; un ee Pre Hy | seent ; fl ' i lh : Af : 4 bo ty os v at (ifs aint i . ¥ ‘i We Ay Al | ay 4 Abh Ti) ! ‘ : ta : A ‘1 { {y a ij i | ¥ ; Lt) 7 if 1 y i! ha 1) ! fi / ‘ ae l Pe 1 aa] 7 “— t i ! t’ ' | ' i | ay | ! ' ‘ 1) i y Y 4 i= 1) roe yal ; ' i = ) - ' . ‘ - ! 1! ' ii ' A 1 Chrys a iF i : r pei yay A : 4 | i : " ', : : iV i aan ‘ 4 1% i ' : i ; } {> / / q 7 toy [ ' i : : i , 4 : j ; é y ' 1 : i 1 _ 3 : = a ie i )? 7 : } : e 4 . ‘ i 1 i 1 - 1 1 1 ' y' = ? : ] 1 ‘ t ' <é ; t | - ' ; : ; ' ‘ ey he ie ; rs cot he yt ' i : " fase capen ale , 7 ; ; | : ‘ ‘ i ug i) : Dis cae i. . | ; ly a ; : : a ' b AGE ail AY = i a, 7 ee . ; po ' : 4 f oe Ya « f os ; : ; i : a5) A 4 \ ' : uy ace A! ye ae r% i 1 1a ' ; c Fyintile : | ei" oe rp ttce i ' i i x i ' 1 l { : | goed : t Hid te ! ‘i } : : ; ’ 14 i hat 7 y° ‘ - 1 : : ; 1 ! . i ay | Fr i _t 7 HN . | jl - ; - A ' J ; =" ; ; ‘ ) oe » ' , - io : rr | : _ - {| - THE NAUTILUS Volume 122, Number 3 September 26, 2008 ISSN 0028-1344 A quarterly devoted to malacology. EDITOR-IN-CHIEF Dr. José H. Leal The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 BUSINESS MANAGER Mary Jo Bunnell The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. 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Box 30 North Myrtle Beach, SC 29582 Dr. Gary Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr. Angel Valdés Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 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 SUBSCRIPTION INFORMATION The subscription rate for volume 123 (2009) is US $50.00 for individuals, US $85.00 for institutions. Postage outside the United States is an additional US $10.00 for regular mail and US $25.00 for air delivery. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 1580, Sanibel, FL 33957, USA, (239) 395-2233. 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 by 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 P.O. Box 1580 Sanibel, FL 33957 CONTENTS NAUTILUS Volume 122, Number 3 September 26, 2008 ISSN 0028-1544 LouElla R. Saul Richard L. Squires Samuel S. Espino Cretaceous trichotropid gastropods from the Pacific slope of North America: Possible pathways to calyptraeid morphology ........ . 115 Feeding behavior, phylogeny, and toxinology of Conus furvus Reeve, Alan J. Kohn 1843 (Gastropoda: Neogastropc nda: Conidae) 2... 200200000... 148 James A. Villanueva Frank M. Heralde HI Patrice Showers Corneli Gisela P. Concepcion Baldomero M. Olivera Richard Duerr Two new species of Mitrella (Gastropoda: Neogastropoda: Columbellidae) from the lower Miocene Chipola Formation of northwestern Florida 2.0.0.0... 0000000000 eee 15] Fresia Villalobos-Rojas Catalogue of the type material of mollusks deposited at the Zoology Ana. G. Guzman-Mora Museum, University of Costa Rica... 0... ee ee 155 Yolanda E. Camacho-Garcia Takenori Sasaki Dilemma japonicum new species (Bivalvia: Anomalodesmata: José H. Leal Poromyidae): A new record of the genus from the Northwest Pacific 2... 000000000 ce ee ees 166 Victor Scarabino On the genus Heteroschismoides Ludbrook, 1960 (Scaphopoda: Carlos Henrique Soares Caetano Gadilida: Entalinidae), with descriptions of two new species .... . 171 Norma Emilia Gonzalez-Vallejo Parasitism of Monogamus minibulla (Olsson and McGinty, 1958) (Gastropoda: Eulimidae) on the red sea-urchin Echinometra lucunter (Linnaeus, 1758) (Echinodermata: Echinometridae) on the Caribbean coast of Mexico 2.0... 0. ee ee 17S EE AUUAMNE oo, ya fe anh Gia ec al st less es sh oe ey tard vy 4h dd haat aa ds dich Gp wes decd ete, Bh Suc ae Pate 182 INQUGES! <2 bo. Fie loa jo dod a Be ROR, Oh Be Solin he Ge & Geb ead FH ed dad beg BOS bee ah ea eee oo ae Oe 183 , oto *s ' ' cm - ' ' . ‘ THE NAUTILUS 122(3):115-142, 2008 Page 115 Cretaceous trichotropid gastropods from the Pacific slope of North America: Possible pathways to calyptraeid morphology LouElla R. Saul Invertebrate Paleontology Section Natural History Museum of Los Angeles County 900 Exposition Boulevard, Los Angeles, CA 90007 USA lousaul@earthlink.net Richard L. Squires Department of Geological Sciences California State Univ ersity, Northridge, CA 91330-8266 USA richard.squires@csun.edu ABSTRACT Late Cretaceous gastropods belonging to genus Lysis Gabb, 1864, from the Pacific slope of North America, bridge the mor- phologic gap between turbiniform trichotropids and limpet-like calyptraeids. Development of the depressed and broadened inner lip/columella of Lysis resulted in a larger aperture that allowed more space for the foot to grasp a hard substrate. Pacific slope species of Lysis are represented by five species that collectively span an interval from late Coniacian to late Maastrichtian. They stem from two lineages of the trichotro- pine genus Ariadnaria Habe, 1961. The first lineage, which includes Ariadnaria ainikta new species of late Albian to Cen- omanian age, Ariadnaria stibara new species of Cenomanian age, and Ariadnaria obstricta (White, 1889) of late Coniacian? and Santonian age, gave rise to the Lysis duplicosta group of neritiform to haliotiform, coarse-ribbed Lysis, including Lysis mickeyi new species (earliest Lysis in the world), Lysis dupli- costa Gabb, 1864, Lysis jalamaca new species, and Lysis lo- maensis new species. The second lineage of Ariadnaria consists of the Turonian Ariadnaria aldersoni new species, which gave rise to the Lysis suciensis (Whiteaves, 1879) group. Morpho- logically, this group, which show crepiduliform and fine ribbed shells, appears likely to be a stem group from which Cenozoic Crepidula-like genera evolved. Garzasia new genus, which ranges from late Campanian or early Maastrichtian age to the mid Maastrichtian, evolved from the Lysis duplicosta group and includes Garzasia intermedia (Cooper, 1894) and Garzasia diabla new species. The very broad, depressed spiraling inner lip of Garzasia is suggestive of Calyptraea Lamarck, 1799. We — placement of Lysis and Garzasia in Lysinae new sub- family of the Trichotropidae. In addition to their occurrence along the Pacific Slope of North America, Lysis or Lysis-like gastropods are known from middle Santonian to lower Cam- panian strata in South Africa, upper Campanian in the Congo, and Maastrichtian strata in Mozambique and Japan. Additional Keywords: Trichotropidae, Lysinae, Calyptraeidae, evolution, paleogeographical occurrence INTRODUCTION This study deals with the fossil record of the extinct ge- nus Lysis Gabb, 1864, a small group of enigmatic gas- tropods which has received little or no study regarding its ancestry, point of origin in terms of time and geogr aphic locale, and ev. olutionary history. Specimens have been found in shallow-marine Cretaceous deposits from southern Vancouver Island and neighboring Gulf Is- lands, British Columbia, Canada to northem Baja Cali- fornia, Mexico (Figure 1) and, although Lysis-like gas- tropods have been reported at a few locales | in the world, its familial placement has been uncertain. This study brings new information about all of these items. Material for this study included type specimens, addi- tional collections from their type localities, and speci- mens of Late Cretaceous age (Coniacian to Maastrich- tian) from previously unreported- upon localities. Figure il provides an index to areas which yielded specimens used in this study. During the study we discovered undescribed species of the trichotropine g genus Ariadnaria Habe, 1961, which appear to have given rise to two groups of Lysis, a ner- itiform coarse- nabbed group and a crepiduliform fine- ribbed group. As will be discussed under “Evolutionary Implications,” we propose that the neritiform group evolved into Garzasia new genus, which appears to be a precursor to calyptraeiform | genera, A total of three genera (one of them new) and 11 species (seven of them new) make up this study. The taxa are: four species of Ariadnaria (three of them new), five species of Lysis (three of them new), and two species of Garzasia (one of them new). The ranges in time of all these species, as well as two recognizable groups of Lysis, are plotted on Figure 2. The Lysis duplicosta group con- sists of four species, spanning a total interval of late Co- niacian to late Maastrichtian. This group was also the most geographically widespread, with specimens col- lected from British Columbia to Baja California. The Lysis suciensis group is known only from the Campanian to possibly early Maastrichtian, with specimens known from British Columbia to Baja California Sur. Lysis per- sisted for a total of approximately 18 million years in the study area. The specimens studied here are mainly from fine- Page 116 THE NAUTILUS, Vol. 122, No. ‘ LATITUDINAL DISTRIBUTION OF SPECIES 13 - Agua Caliente Canyon. Santa Barbara Co., CA 14 - Warm Springs Mtn., 1 -Vancouver and adjacent islands, British Columbia 2 - Sucia Island, San Juan Co., Washington Los Angeles Co., Calif. 3 - South Cow & Bear creeks, 15 - Live Oak Canyon, Shasta Co., Califomia 1 1 e Kem Co., California 4 - Ono area, Shasta Co., Calif. 2 e 2 16 - Simi Hills, Los Angeles 5 - Chico Creek, Butte Co., @ ® and Ventura counties, California California 6 - Pentz and Dry Creek, Butte 17 - Santa Monica Mtns, Co., Califomia Los Angeles and 7 - Texas Flat & Granite Bay, Ventura counties, Calif. Placer Co., California 18 - Santa Ana Mtns., - Pigeon Point, San Mateo Co., Orange Co., California California 19 - near Carlsbad, San 9 - Garzas Creek, Stanislaus Co., Diego Co., California California 20 - Point Loma, San 10 -south of Garzas Creek, Diego Co., California Merced Co.,Califomia 21 - East of La Misién, Baja California, Mexico 22 - Cafion San Fernando, Baja California, Mexico 23 - Punta Abreojos, Baja California Sur, Mexico 11 -Cooper Canyon, Fresno Co., Califomia 2 -Jalama area, Santa Barbara Co., Califomia e ‘e 3@ °e 5@ -® s § (97% 68 'S = b= 7@ 708 a. = = | i} — N 8 9 ® 108 11 = @ 3s re = § SU 159 = e 2e 12g “e S < 16@ 16 S = 19@ 19@ i = Ss 20@ 20@ iS 2105 = ‘Se 8 js s 8s § ys § S SS ~ YY 22 22@ <@ S =. & : Kee 3 BAT, Ss ‘ ~ 50 100 500 kn} CALIFORN), a 23geN 23@ Figure 1. Index map of collecting localities L. R. Saul and R. L. Squires, 2008 Page 117 35 33.8 —t_ cia | foe 3 ae PRIABONIAN IC16 | jaSo%| : — 372 — Hse 40} 1.3 BARTONIAN = FRS|CI8) es} | YQ es eo | Pa = ed m 4c20| Peres) 45 en 5 LUTETIAN ee| & : — IE a Ee: =] Ol ae Tl fest S, 50-4 oO a C22 | SS a === | —|—_ i = = 4 YPRESIAN a3 »~ aS ox) 4 = » SS = < 55.8 3 8 < = =4 Z THANETIAN x SS & = ss — we 58.7 = ££ & te = Sy 0) |} 2 | SELANDIAN es £ € € &§ s § BH = DANIAN - .& 5 8 :. 8 £ oS —— = 65.5 ——— (29) L rR eS. & S Pa > = MAASTRICHTIAN jet —— : Jw 4 OS 70 — Pcs oi J fet = = TOG re aa —— ee 2 7 |——— | 15 = ~ | : = CAMPANIAN & : C33 ~ vee SS if 8 S| = < so : | = eal $35 || ; ae es oe SANTONIAN = FES 4 | a | CONIACIAN | 90 =] O 89.3 —— 7 TURONIAN ce | | 4 93.5 3 95 —} ©) 4 —j CENOMANIAN S — mS 100 —| 99.6 oe = 4] | S yy 105—| & = 0 ALBIAN su 110 be ~ — — = Sel e 112.0 - a M1 1 ed APTIAN Zo 255 | — 125.0 — ii Figure 2. Age ranges of species of Ariadnaria, Lysis, Garzasia, Calyptraea, and Crepidula discussed in text. Lines connecting the species indicate inferred descent based on morphologic simiarity. Lysis duplicosta gave rise to two species one of which, L. lomaensis, produced proto-calyptraeform Garzasia and the other, L. jalamaca, is crepiduliform. An additional crepiduliform line was founded by L. suciensis. No known intermediate forms conne . these Cretaceous calyptraeiform and cre piduliform g gastropods to Tertiary species. The earliest known Pacific Slope calyptraeid, ‘C. diegoana lacks the strong ribbing of Garzasia but crepiduliform Spiro- crypta pileum differs largely from L. suciensis in its a size. Time scale after Gradstein et al. (2004). Page 118 THE NAUTILUS, Vol. 122, No. 3 grained argillaceous sandstone or siltstone that constitute shelfal deposits that accumulated at depths near or just below wave base. Specimens are low in number and almost always in- complete. Protoconchs are rarely preserved, the larval shell is in part missing, its shape partially represented by its inner cast in all available specimens. Apertures are usually missing their anteriormost area. Adhering matrix, commonly consisting of well-cemented s sinidstone usually plugs the aperture, “thereby necessitating careful clean- ing. ‘As will be discussed under “Evolutionary Implica- tions,” the shape of Lysis, with the exception of L. mick- eyi new species, approaches that of Crepidula, resulting from flattening of the aperture, development of a Bioad shelf, and lateral coiling of the spire. The crepiduliform shape, as well as the Calyptraea-like shape of Garzasia, pose problems for terminology used for describing cer- tain figures of specimens, as well as for describing certain shell dimensions. For some specimens, a full view of the aperture could only be depicted by having the the shell tipped out of the plane of its axis. The true height of these shells, therefore, is not shown; hence, the ‘ ‘vertical dimension” of the view is given instead, and perpendicu- lar to it is the “horizontal dimension.” In most cases, the abapertural view is in the plane of the shell axis; hence, the terms “height” and “diameter” apply. Abbreviations used in the text are: ANSP: Academy of Natural Sciences of Philadelphia; CAS: California Acad- emy of Sciences, San Francisco; CGS: Geological Survey of Canada, Ottawa: CIT: California Institute of Technol- ogy, Pasadena (collections now housed at LACMIP); CSMB: California State Mining Bureau (specimen at CAS): IGM: Instituto de Geologia, Universidad Nacional Aut6noma de México; LACMIP: Natural History Mu- seum of Los Angeles C ounty; SDNHM: San Diego Natural History Museum: UCLA: University of Califor- nia, Los Angeles (collections now at LACMIP); UCMP: University of California, Berkeley, Museum of Paleon- tology: USGS: United States Geological Survey, Menlo Park (collections now housed at UCMP): USNM: Na- tional Museum of Natural History, Smithsonian Institu- tion. PALEOBIOGEOGRAPHIC DISTRIBUTION OF ARIADNARIA, LYSIS, AND GARZASIA Ariadnaria ranges from late Albian to Recent with its earliest appearance in Pacific slope deposits of North America, where it is found in strata ranging from late Albian to Santonian age. From Japan, Kase (1990) fig- ured, as Tric hotropis?, a possible Ariadnaria species of earliest Maastrichtian age. Lysis ranges from the late Coniacian to late Maastri- chitan, and that is also its range for the Pacific slope of North America. The genus apparently originated in Cali- fornia, with five species known from the Pacific slope of North America. Four additional pre »bable species of Ly- sis are known elsewhere in the world. They are: Lysis capensis Rennie, 1930, from the middle Santonian to lower Campanian of South Africa (Kiel and Bandel, 2003); Lysis congolensis (Brébion, 1956) from upper Campanian of fic Congo, Africa; Lysis africana (Cox, 1925) from the Maastichtinn (undifferentiated) of Mo- zambique; and Lysis izumiensis Kase, 1990, from the earliest Maastrichtian of Japan. Rennie (1935) reported Lysis caffra Rennie, 1935, from the Upper Cretaceous near the eastern border of the Eastern Cape Province (Pondoland), southeast Af- rica, but its swollen naticiform shape with a very large, inflated body whorl and broad, non-depressed inner lip/ columellar region, that appears to have a thin callus, are features not found in Lysis. Garzasia ranges from late Campanian or early Maas- trichtian to the anid Maastrichtian and is endemic to the Pacific slope of North America. MODE OF LIFE OF LYSIS Lysis has been found attached to a few specimens of large volutid gastropods. One specimen of Longoconcha aumneka Saul and Squires, 2008, from the Point Loma Formation near Carlsbad, California has two specimens of crepiduliform Lysis jalamaca new species on its shell, near the outer lip margin (Figure 34), as well as an at- tachment scar on the abapertural side of the shell. Speci- mens of Lysis suciensis from the Chatsworth Formation near Chatsworth, California, although not found in situ as are the younger Carlsbad specimens, have aperture shapes that sould fit on the exterior of a specimen of Volutoderma Gabb, 1877. Indication of such an associa- tion is absent prior to middle Campanian. Whether the specimens of Lysis species were using dead shells as a substrate or had developed an association with living Vo- lutoderma is undetermined. Comparison to probable family members indicates that Lysis was a sedentary faculative ciliary feeder. Tri- chotropines, as well as calyptraeids, are ciliary feeders that live a sedentary life on hard substrates and are pro- trandrous hermaphrodites among whom brooding their young is common (Graham, 1954; Yonge, 1962). "These mode. of-life characteristics have historically been used to classify trichotropines as being close to calyptraeids. SYSTEMATIC PALEONTOLOGY Superfamily Calyptraeoidea Lamarck, 1S09 Discussion: The taxonomy of calyptraeoids, as with most gastropods, was traditionally based on shell mor- phology and later modified by increasing anatomical knowledge. The inclusion of patelliform Capulidae Fleming, 1822, turbiniform Trichotropidae, and coiled limpet- ee aped Calyptraeidae in the superfamily Calyp- traeoidea (e.g¢., Thiele, 1929: Wenz, 1940), has provided L. R. Saul and R. L. Squires, 2008 Page 119 calyptraeoids a considerable morphological range and a somewhat complicated classification history (Bz wndel and Riedel, 1994). In addition to the above three families, Bandel and Riedel (1994), included Hipponicidae Troschel, 1861, in Gchonaestdee but Collin (2003: 632) rejected hipponicids from a close relationship with ca- lyptraeids. Ponder and Warén (1988) and Ponder (1998), equated family Capulidae with family Trichotropidae Gray, 1850, on anatomical grounds, reducing the in- cluded families to two. Capulids, as exemplified by Capulus Montfort, 1810, have limpet-shaped shells; tri- chotropids, as exemplified by Trichotropis Broderip and Sowerby, 1529, usually have coiled shells; and calyp- traeids, as exemplified by Calyptraea Lamarck, 1799, Crepidula Lamarck, 1799, and Crucibulum Schumacher, 1817, have limpet-shaped shells with an internal shelf of variable shape. In her analysis of calyptraeids, Collin (2003a, 2003b) utilized shell characters, anatomical characters, and mo- lecular characters. Collin (2003a) argued that although the so-called echinospira larva of Capulus and of Tricho- tropis do not appear to be “true” echinospira, the thick- ened and elaborate larval shell of these two groups is clearly different from the simple larval shell of extant calyptraeids, and she (Collin, 2003a, 2003b, 2005) has continued to recognize the families Capulidae Fleming, 1822, Calyptraeidae, and Pai ae Collin (2003a) mentioned that despite the detailed studies done on ca- lyptraeids, their taxonomy remains contentious and un- certain. Because specimens studied here show no resem- blance to capulids but do, in part, resemble trichotro- pids, and calyptraeids such as Calyptraea and Crepidula, we follow Collin in recognizing families Calyptraeidae and Trichotropidae. Family Trichotropidae Gray, 1850 Subfamily Trichotropinae Gray, 1850 Description: Small to medium sized (usually 15 to 25 mm, but up to 40 mm in height), coiled, high- -conic to broad, low-conic, or nearly cap-shaped; spiral sculpture usually better developed than collabral sculpture and represented by distinct cords and/or keels; umbilicus broadly open to slit-like or completely closed; aperture variable in shape, ranging from irregularly triangular and broadly oval to elongate- -oval: some forms ath more or less attenutated canal: operculum small, horny; radula taenioglossate: periostracum forming combs, bristles, spines ‘usually in places of intersection of spiral and col- labral sculpture (from Egorov and Alexeyev, 1998). Discussion: Trichotropids provide the geologically oldest representatives of the three families Trichotropi- dae, Capulidae, and Calyptraeidae. We did not follow Ponder and Warén (1988), Ponder (1998), and Bouchet and Rocroi (2005) in placing Trichotropidae in Capul- idae because trichotropids, such as the high spired Ari- adnaria spp. discussed herein, differ distinctly from cap— shaped capulids. Collin (2003b) referred to trichotropids plus capulids as the closest outgroup to calyptraeids, sug- gesting that she recognized these three as separate fami- lies. At present the geologic record finds trichotropids occurring earlier than capulids and also earlier than ca- lyptraeids, and we suggest that based on species de- scribed herein, trichotropid-like gastropods gave rise to calyptraeids and that trichotropid specimens reported upon herein demonstrate a progressive development to- ward either a crepiduliform or a calyptraeiform shell. Atresius Gabb, 1869, of Early Cretaceous (Valanginien to Hauterivian) age is the earliest trichotropine recog- nized by Wenz (1940), but its sole named species A. liratus Gabb, 1869, is a prominent constituent of chemo- synthetic paleocommunities in northern California and probably not a trichotropid. Linpsa Stephenson, 1952, of Cenomanian age from the Woodbine Formation of Texas may be the earliest previously known trichotropine. The earliest reported cap- shaped sears is Capulus verus (B6hm, 1885) of Late Cretaceous (early Campanian) age from Aachen, Germany. Genus Ariadnaria Habe, 1961 [= Ariadna Fischer, 1864; non Audouin, 1826]. Type Species: Trichotropis borealis Broderip and Sowerby, 1829, by monotypy; boreal Arctic circumpolar (Egorov and Alexeyev rev, 1998) and cool temperate seas: norhern North Aglenue south to Northumbria and all Scottish coasts (Fretter and Graham, 1962) and Massa- chusetts Bay (Emerson and Jacobson, 1976); the Bering Sea and north Pacific south to British Columbia (LACM collection, Forrester Island). Description: Shell turbiniform (oval-conic) with raised spire; spiral sculpture well developed and consist- ing of raised cords separated by interspaces of variable width: collabral sculpture consisting of raised growth lines; aperture wide; inner lip slightly concave; canal short and straight; umbilicus. slit- like: operculum thin; periostracum for ming long, closely spaced bristles on spi- ral ribs. Discussion: No prior records of Ariadnaria as a fossil were found by us. Ariadnaria differs from Trichotropis Broderip and Sowerby, 1829, by having an umbilicus. Turbinopsis Conrad, 1860, a turbiniform trichotropid of late Campanian or early Maastrichtian age from Missis- sippi, differs from Ariadnaria by having a wider umbili- cus (profound according to Conrad, 1860), a last whorl that is more inflated and is tabulate, and having a very oblique fold near the basal margin of the columella. Ariadnaria ainikta new species (Figures 3-4) Diagnosis: Medium-size Ariadnaria with sturdy shell, rounded whorls, strong sculpture with nine spiral cords widely spaced; collabral sculpture thickly foliate; umbili- Page 120 THE NAUTILUS, Vol. 122, No. 3 Figures 3-10. CAS 61794.00, CAS loc. 61794, height 20 mm, diameter 17 mm. 3. Apertural view. 4. Abapertural view. 5-6. Ariadnaria stibara new species, holotype LACMIP 13371, LACMIP loc. 23464, height 19 mm, diameter 13.5 mm. 5. Apertural view. 6. Abapertural view. 7-8. Ariadnaria aldersoni, holotype LACMIP 13372, LACMIP loc. Ariadnaria species. Specimens coated with ammonium chloride. 83-4. Ariadnaria ainikta new species, holotype 26370, height 12 mm, diameter $.5 mm. 7. Apertural view. 8. Abapertural view. 9-10. Ariadnaria obstricta (White, 1889), hypotype LACMIP 13373, LACMIP loc. 28717, height 21 mm, diameter 15 mm. 9. Apertural view. 10. Abapertural view. cus elliptical; inner lip broad, expanded anterior and pos- terior to umbilicus. Description: Shell medium (approximately 22 mm height), sturdy, turbiniform, spire moderately high, ap- proximately 36% of total shell height: apical angle | 110°: most of protoconch missing, remanent low and appar- ently smooth; teleoconch whorls three, whorls well rounded and enlar ging very rapidly; suture appressed but appearing channe ip d: umbilicus open, deep, and ellipti- cal; grow th | ine prosocline; spiral sculpture consisting of regularly spaced and equally narrow cords; four éoide. on penultimate whorl: nine cords on last whorl, becoming stronger and more raised near umbilicus; spiral cords on last Se widely spaced and occasionally with spiral thread in interspaces, especially anterior of periphery: collabral sculpture consisting of f thickly foliate ridges co- incident with growth heel. especié ally near outer lip; aperture D-shaped; inner lip broad, expande -d anteriorly and posteriorly of umbilicus; abapertural edge of inner lip delineated ‘by sharp ridge; basal lip broadened. Holotype: CASG 61794.00, height 20 mm, diameter 17 mm, spire height 7.5 mm. CASG loc. 61794 dpe ] ocality: {=CASG loc. 134€ Distribution: Basal Bald Hills Member of the Budden Canyon Formation, (area 4) Ono area, Shasta Co., Cali- fornia. Geologic Age: Late Albian. Discussion: Only the holotype is known. It evidently had a very foliate, thick shell. What remains is riddled with endobiont borings. Remnants of the protoconch are present, there is no clear evidence of an anterior sinus to the aperture, and the shell does not appear to have been nacreous. The new species most resembles the trichotropid Turbinopsis hilgardi Conrad, 1860 (Conrad, 1560: 289, pl. 46, fig. 29; Sohl, 1960: 91, pl. 10, figs. 17, 18) from the upper Campanian/lower Maastrichtian Ripley Formation of Tippah Co., Mississippi, except A. ainikta has a smaller umbilicus and narrower cords with — wider interspaces, Additionally, the inner lip of A. ainikta is more expanded both anterior and posterior ee the um- bilicus. Ariadnaria ainikta and A. stibara new species are simi- lar in that there is no ridge separating the umbilicus from the inner lip. Arindnaria ainikta differs from A. stibara by having a less elongate shell, wider pe Geran whorl, stronger spiral cords with much wider interspaces, a shorter umbilicus that is oval rather than slit-like, foliate L. R. Saul and R. L. Squires, 2008 Page 12] collabral sculpture, and no parietal swelling on the inner lip. Ariadnaria ainikta differs from A. aldersoni new spe- cies by being larger, having much stronger spiral cords with much wider interspaces, a well rounded last whorl (not angulate), shorter umbilicus that is oval rather than slit-like, and no ridge separating the umbilicus from the inner lip. Ariadnaria ainikta differs from A. obstricta (White, 1SS9) by having a lower spire, wider penultimate whorl, round last whorl (not angulate) more spiral cords, foliate collabral sculpture, shorter umbilicus that is ov al rather than slit-like, and no ridge separating the umbili- cus from the inner lip. The strong spiral ribbing of A. ainikta resembles that of A. obstricta. Etymology: Ainiktos, Greek, meaning: baffling, ob- scure, or enigmatic. Ariadnaria stibara new species (Figures 5-6) Diagnosis: A medium size sturdy Ariadnaria with rounded whorls, medium strong sculpture with many spiral cords moderately closely spaced: collabral sculp- ture very fine and lattice-like on spire whorls; umbilicus chink-like: inner lip with low parietal swelling. Description: Shell medium small (height approxi- mately 20 mm), sturdy, turbiniform, somewhat elongate, spire high and approximately 50% of total shell height: apical angle approximately 67°; protoconch not present; teleoconch whorls four, whorls smell rounded and enlarg- ing rapidly: last whorl tapering anteriorly; suture ap- pressed, appearing channeled, and rapidly descending near aperture; umbilicus narrow, chink-like and present only adjacent to medial and posterior parts of inner lip; growth line prosocline, with several irregularly spaced crowth checks near outer lip; spiral sculpture consisting of numerous fine subequal rounded cords; approximately ten closely spaced cords on penultimate whorl with in- terspaces s of nearly equal width; approximately 18 cords on last whorl with interspaces slightly wider than cords: cords strongest, most widely spaced, and occasionally with spiral thread in interspaces on medial and anterior portions of last whorl; collabral sculpture consisting of thin, raised growth lines, forming nearly microscopic are tice-like pattern on spire iia aperture D-shaped, moderately large, oblique, narrowed at posterior end by low parietal swelling: inner lip broad, somewhat exca- vated (concave) medially and flattened ante riorly; aba- pertural edge of inner lip delineated by low but distinct ridge: b asal lip broadened. Holotype: Holotype LACMIP 13371, height 19 mm (incomplete), diameter 13.5 mm, spire height 9 mm. Type Locality: LACMIP loc. 23464 is also type local- ity of Turrilites dilleri Murphy and Rodda, 1960. Distribution: Bald Hills Member (unit [IV of Matsu- moto, 1960) of the Budden Canyon Formation, (area 4) Ono area, Shasta Co., California. Geologic Age: Middle Cenomanian. Discussion: Only the holotype is known, and it lacks the protoconch and the anterior end of the teleoconch. Whether or not an anterior siphonal notch was present is unknown, but the shape of the last whorl suggests that at least a small one was present. The species is assigned to Ariadnaria based on shell shape, but it differs from typi- cal Aradnaria in its very sturdy shell, the fineness of its sculpture, and the presence of a parietal swelling at the posterior end of the aperture. Ariadnaria stibara differs from A. ainikta by having a more elongate shell, narrower penultimate whorl, much weaker spiral cords with much narrower interspaces, longer and narrower umbilicus, parietal swelling on inner lip, and absence of foliate collabral sculpture. Ariadnaria stibara differs from A. aldersoni by larger size, less elon- gate shell, more rounded whorls that are not lax in their coiling, coarser spiral ribs, lattice-like very fine collabral sculpture on spire whorls, parietal swe lling on inner lip, less delineated abapertural edge of inner lip, and no fas- ciole-like flange aaa the umbilicus. Ariadnaria stibara differs from A. obstricta by having rounded whorls, many more spiral ribs that are much weaker and much more closely spaced, and a less well demarked abapertural edge of the inner lip. Etymology: Named for its sturdy shell, stibaros, Greek, meaning strong or sturdy. Ariadnaria aldersoni new species (Figures 7—S) Diagnosis: Small Ariadnaria with elongate shell and last whorl medially subangulate, coiling stretched axially, whorls bearing many very fine and closely spaced ribs, umbilicus chéniks like, spiral sculpture very fine, abaper- tural edge of inner lip well delineated and raised, umbi- licus bordered by fasicole-like flange. Description: Shell small (approximately 13.5 mm height, estimated); elongately turbiniform, with medially subangulate whorls; upper spire missing; teleoconch whorls enlarging rapidly; suture apparently appressed on earlier whorls, becoming channeled on penultimate whorl, rapidly desce nding: umbilicus chink-like, bounded by strong rounded “fasciole- like ridge; growth line prosocline, we aT marked with numerous growth welts on last whorl; spiral sculpture consisting of fine, well spaced ribs of unequal strength crossing irregular growth welts; interspaces between ribs wider than ribs; aperture D-shaped, narrowed posteriorly but not angled, with short, broad anterior sinus: outer lip thin, simple; inner lip moderately narrow, abapertural edge raised and well demarked, Holotype: LACMIP 13372 plete), diameter 8.5 mm. - 2 WeOTne height 12 mm (incom Type Locality: LACMIP 26370 Distribution: Panoche Formation, (area 11) Alcalde Hills. Fresno Co., ¢ Jalifornia. Dace 199 Page 122 THE NAUTILUS, Vol. 122, No. 3 Geologic Age: Late Turonian. Discussion: Only the holotype is known. It is incom- plete, consisting only of the last two whorls, and its small size may iaciente that it is not mature. The elongate shape is a distinctive characteristic of this species. Ari- adnaria aldersoni apparently had a relatively high spire, and the coiling is lax and reminiscent of Lirpsa Ste phen- son, 1952. It somewhat resembles Lirpsa teres Stephen- son, 1952, but the new species has a narrow, chink-like umbilicus. Ariadnaria aldersoni is most similar to A. stibara and differs by being smaller, having a more elongate shell, angulated ions that are lax in their coiling: much weaker spiral ribs, abapertural edge of inner lip better delineated, fasciole-like ridge bounding the umbilicus, absence of lattice-like very fine eollabral sculpture on spire whorls, and absence of parietal swelling on inner lip. Ariadnaria aldersoni differs from A. dindleta by being smaller, having much weaker spiral cords with much nar- rower interspaces, angulate last whorl, longer umbilicus that is slit-like rather than oval, and having a ridge sepa- rating the umbilicus sige the inner lip. Ariadnaria al- dersoni differs from A. obstricta by being smaller and having fewer and eae weaker spiral ae with much narrower interspaces. In some respects A. aldersoni is similar to Lysis sucien- sis. Both have fine ribbing, a relatively high spire, rather lax coiling, and a somewhat slower increase of whorl diameter. Whereas the suture of A. obstricta and Lysis mickeyji is very close to or at the perimeter of the previ- ous whorl, in A. aldersoni and Lysis suciensis the suture is usually abapical to the previous whorl’s perimeter. Etymology: The species is named for John M. Alder- son who collected the holotype from Cooper Canyon. Ariadnaria obstricta (White, 1889) (Figures 9-10) Stomatia obstricta White, 1889: 15-19, pl. 4, figs. LO-11. Diagnosis: Medium size Ariadnaria with high spire and angulate last whorl, sculpture of a few widely spaced strong cords, umbilicus chink-like or covered. Description: Shell medium size (approximately 21.4 mm height), elongate turbiniform; whorl profile rounded with slight angulation at third strong cord on last whorl; spire high and approximately 50% of total shell height: apical angle approximately 67°; protoconch missing; teleoconch whorls four, enlarging rapidly and last whorl tapering anteriorly; suture appressed, anterior to suban- gulate periphery; umbilicus chinklike or covered by inner lip expansion; bounded abaperturally by strong ridge; growth line prosocline; sculpture of three strong, wide ‘ly spaced cords on spire, five or six on body whorl; inter- spaces commonly with mid thread; aperture large and ovate with abape rtural edge raised and_ shi ply de- marked; outer lip appare mntly simple; inner lip somewhat ‘xspanded and standing high along umbilical chink; basal lip barely drawn out into slight spout-like sinus. Holotype: USNM 20124. Type Locality: Little Cow Creek but additional speci- mens have not been found there. The species is abun- dant at some localities along South Cow Creek, Shasta Co., California. Hypotype: LACMIP 13373, height 20 mm, diameter 15 mm from LACMIP loc. 28717. Distribution: Redding Formation, Bear Creek Sand- stone Member, especially ( area 3) along South Cow Creek and Bear Creek, Shasta Co.; Chico Formation, Musty Buck Member, (area 5) Chico Creek, Butte Co., California. Late Coniacian? to Santonian. Geologic Age: Discussion: The above description is based on 16 specimens; all but one are from LACMIP loc. 28717. Most specimens are poorly preserved, and specimens with the shell surface preserved are difficult to find. The primary cords are strong and almost flange-like. White’s species is here assigned to Ariadnaria based on shell shape, sculpture, umbilicus, and presence of a small spout-like sinus in the aperture. This would be the earliest unquestioned occurrence of this genus that pre- viously was known only from the Recent (Wenz, 1940). In shape and probably sculpture (preservation makes comparison difficult) A. obstricta resembles illustrations of Trichotropis? sp. in "se (1990: 568, figs. 2.26, 2.27). Kase’s specimen was from the Izumi Group of Japan of early Maastrichtian age. Ariadnaria obstricta is very similar to Lysis mickeyi new species, but A. obstricta has more regular spiral ribs, a slighly higher spire, the strong spiral delimiting an um- hhilical chink, and a free st: nding inner lip. Aviadnane obstricta differs from Ariadnorin ainikta by having a higher spire, narrower penultimate whorl, angulate last whorl, fewer spiral cords, shorter umbilicus that is slit- like rather than oval, ridge separating the umbilicus from the inner lip, and an absence of foliate collabral sculp- ture. Ariadnaria obstricta differs from A. aldersoni by being larger, having fewer spiral cords that are much stronger and much more widely spaced, and lacking a fasciole-like ridge bounding the chink like umbilicus. Ariadnaria obstricta differs from A. stibara by having angulate whorls, fewer spiral ribs fee are much stronger sad much more widely spaced, and a more demarked abapertural edge of the inner lip. Subfamily Lysinae new subfamily Description: Small to moderately large (15 to 80 mm in height), low turbiniform to almost haliotiform, barely siphon ite shells with spiral ribbing. Final whorl some- what to greatly enlarged; spire very heets aperture large, nearly croiilar toe longate oval: columella and inner lip flattened, expanded, and de »pressed to form a shelf within the aperture; some with shelf that spirals into an “umbi- licus.” Discussion: The subfamily Lysinae includes Lysis L. R. Saul and R. L. Squires, 2008 Page 123 Gabb, Garzasia new genus, and probably Spirogalerus Finlay and Marwick, 1937. These gastropods are inter- mediate in form between trichotropines and calyp- traeids. If their characteristics were better known, some other species such as those discussed under Global Dis- tribution of Cretaceous Lysiform Gastropods, probably could be included here, some as Lysis or Garzasia others in as yet undescribed genera. Genus Lysis Gabb, 1864 Tropidothais Cox, 1925: 213-214. Type Species: Lysis duplicosta Gabb, 1864, by mono- typy (Stewart, 1927: 345): Campanian of Pacific slope of North America. Description: Turbinate to crepiduliform gastropods with a rapidly expanding whorl diameter having the col- umella/inner lip flattened, expanded, and submerged to form a narrow to broad shelf or deck. Shell sculptured by spiral cords or smooth. Nonumbilicate. Aperture with very slight anterior siphonal notch. Discussion: Lysis differs from Trichotropis and Ari- adnaria in having the inner lip completely appressed to the columella. Typical Lysis (i.e., the group of L. dupli- costa) has a carinated whorl in the juvenile stage and a few moderately strong to strong spiral cords. The stron- ger spirals are typic Sie scaly. Ineludeds in this group is L. duplicosta and the following new species: Lysis mickeyi, L. jalamaca, and L. lomaensis. The group of Lysis sucien- sis has a more rounded whorl profile and more subdued, finer spiral sculpture; included in it is L. suciensis. Group of Lysis duplicosta The genus Lysis was proposed by Gabb (1864) for a low-spired, turbiniform gastropod with a depressed inner lip. He had only immature specimens of a single species (ie., the type species) and did not recognize thew simi- larity to genus Crepidula Lamarck, 1799. He gave no indication of the familial affinities of Lysis, other than stating the general form is like genus Stomatia Helbling, 1779. During the last 127 years, Lysis has been placed in at least 11 inde scattered among “archaeogastropods” to the neogastropods. A review of this placement history is given here. Species that were eventually placed in Lysis were originally placed in Stomatia of the Stomatellidae Gray. 1840, by Whiteaves (1879, 1903) and White (1889). Stoliezka (1867-1868: 157-158) suggested ae Lysis should be placed near Separatista Gray, 1847, the Trichotropidae Gray, 1850, in the event that ae does not belong in either the Naticidae Guilding, 1834, or the Velutinidae Gray, 1840. Fischer (1885) placed Lysis in the Naticidae near Eunaticina Fischer, 1885. Tryon (1554: 112) did not hesitate to refer Lysis to the Muricidae Rafinesque, 1815 (as Murexia) [= Purpuradae Children, 1823], but on page 208 Tryon suggested a re- lationship to Velutina Fleming, 1821. of nes Lamellari- idae dOrbigny, 1841. Cossmann (1903) wrote that Lysis could not be a muricid but must be placed near Fossarus Philippi, 1841, presumably in the Fossaridae Adams, 1860, where Stewart, 1927, Rennie (1930), Wenz, 1940, and Anderson (1958) also put it. Cossmann (1925) con- sidered Lysis to be a subgenus of Micreschara Cossmann (1891) in the family Vanikoroidae Gray, 1840. Dall in Eastman (1913) and Packard (1922) placed Lysis in the “Thaisiidae” (=Thaididae) Suter, 1909. Saul (1959) and Saul and Alderson (1981) placed Lysis in Calyptraeidae, and, in 1990, Saul included it in superfamily Calyptrae- oidea. Kase (1990) discussed previous taxonomic treat- ments of Lysis and suggested that, based on its inner lip shelf, the genus should be placed within the Calyp- traeidae Lamarck, 1799. Bandel and Riedel (1994) and Kiel and Bandel (2003) supported this placement. Cox (1925) named and described genus Tropidothais Cox, 1925, which is a junior synonym of Lysis. He based Tropidothais on T. africana Cox, 1925, from the Maas- trichtian of Mozambique [formerly Portuguese East Af- rica] and tentatively placed his genus in ‘the Thaididae Jousseaume, 1588. Upon realizing its similarity to Lysis, Cox (in Rennie, 1935) synonomized the two genera and placed Lysis in the Stomatellidae. In this present paper, we place Lysis in the family Trichotropidae, subfamily Lysinae because Lysis appears to have evolved from tri- chotropids by expanding the columella/inner lip area (width and length) t to form an interior shelf suggestive of the calyptraeid Crepidula. Lysis mickeyi new species. (Figures 11-16) Diagnosis: A relatively high spired Lysis with eight or nine strong cords on last w horl: shelf moderately broad, somewhat depressed, and shallowly concave. Description: Shell medium small (height approxi- mately 20 mm), turbiniform: whorl profile overall rounded with medial angulation on last whorl; spire mod- erately low and approximately 30% of total shell height: apical angle approximately 90°; protoconch 1.5 whorls, low and sacoth: teleoconch whorls 3.5, Pee ex- panding and last whorl tapering anteriorly; suture abut- ting and becoming laxly channeled on later whorls: sculp- ture of strongly raised cords, either moderately closely spaced or widely spaced; penultimate whorl with two to six and last whorl with eight or nine strong spirals, with variable number (two to five) of finer spirals in inter- spaces: both cords and interspaces crossed by fine colla- bral ribs, producing beaded appearance; angulation mod- erately strong on last whorl, coincident with strongest spiral ‘cord: spiral cord anterior to angulation nearly same strength, thereby oS bicarinate appearance to medal part of last whorl: aperture large, oblique, barely notched anteriorly; outer lip simple; shelf moder rately broad, somewhat depressed, and shallowly concave. Holotype: LACMIP 13374, 9.5 mm, spire height 5 mm. Paratypes: LACMIP 13375 from LACMIP loc. 23617 and 13376 and 13377 from LACMIP loc. 10757. height 13 mm, diameter Page 124 THE NAUTILUS, Vol. 122, No. 3 Figures 11-28. Lysis species. 11-16. Lysis mickeyi new species. L1-12. Paratype LACMIP 13375, LACMIP loc. 23617, height 21 mm, diameter 16 mm. 11. Apertural view. 12. Right-lateral view. 13. Paratype LACMIP 13376, LACMIP loc. 10757, left- later view, height 9 mm, diameter 11 mm. 14. Holotype LACMIP 13374, LACMIP loc. 10757, abapertural view, height 13 mm, diameter 9.5 mm. 15-16. Paratype LACMIP 13377, LACMIP loc. 10757, height 5 mm, diameter 9 mm. 15, Left-lateral view. 16. Apical view. 17-28. Lysis duplicosta Gabb, 1864. 17. Plasto-lectotype of Stomatia suciensis carinifera Whiteaves, 1879, CGS 5772, height 10 mm, diameter 19.5 mm. 18-19. Hypotype LACMIP 13378, LACMIP loc. 24128. 18. Apertural view, vertical Aienetision 27 mm, horizontal dimension 21 mm. 19. Abapertural view, height 22 mm, diameter 26.5 mm, 20-21. Hypotype LACMIP 13379, LACMIP loc. 24340, 20. pected view, vertical dimension 28 mm, horizontal dimension 25 mm. 21. Lateral view, vertical dimension 13 mm, horizontal diameter 29.5 mm, 22-23. Hypotype LACMIP 13380, LACMIP loc. 24340, height 19 mm, diameter 18 mm. 22. Apertural view. 23. Aan’ rtural view. 24. Hypotype LACMIP 13381, LACMIP loc. 24340, abe ipertural view, vertical dimension 28 mm, diameter 37.5 mm. 25. Hypotype LACMIP 13382, LACMIP loc. 24349, abapertural view, height 17 mm, diameter 22 mm. 26. Hypotype LACMIP 13383, LACMIP loc. 10095, crushed specimen, abapertural view, height 24 mm, diameter 22.5 mm. 27. Hypotype LACMIP 13354, LAC MIP loc. 26951, abapertural view, height 21 mm, diameter : 25 mm. 28. Hypotype LACMIP 13355, LACMIP loc. 24340, abapertural view, height 7.5 mm, diameter 12 mm. L. R. Saul and R. L. Squires, 2008 Page 125 Type Locality: LACMIP loc. 10757. Distribution: Redding Formation, Bear Creek Sand- stone Member of Haggart (area 3) on Bear Creek, Shasta Co.; Chico Formation, top of Ponderosa Way Member and Musty Buck Member (200 m to 650 m above the base of the section) (area 5) on Chico C oa Butte Co.; basal Tuna Canyon Formation (area 17) at head of Ga- rapito Creek, Santa Monica Mountains, Los Angeles Co., California. Geologic Age: Early Coniacian to Santonian. Discussion: The above description is based on SO specimens; most of these are from LACMIP locs. 10846 and 23617. Most specimens are internal molds. Many show endobiont boreholes, especially on the spire whorls. Two specimens show the protoconch. The oldest specimen. is from LACMIP. loc. 26967 in the Santa Monica Mountains. Lysis mickeyi resembles Lysis suciensis (Whiteaves, 1879) in height of spire but is closer to L. duplicosta in sculpture. Lysis mickeyi differs from L. duplicosta in having a higher spire, less expanding last whorl, much less e xpanded shelf, more oval aperture, and thinner spi- ral cords. Lysis mickeyi greatly resembles Trichotropis obstricta (White, 1889), but on L. mickeyi the spiral ribs are less regular, and its spire is slightly lower. In addition, L. mickeyi lacks an umbilical chink and a free- standing inner lip. Lysis mickeyi is the earliest known Lysis from any- where in the world. Etymology: Named for Mickey of Mickey's House on Chico Creek near the locality, LACMIP 23617, from which the species was first recognized. Lysis duplicosta Gabb, 1864 (Figures 17-28) Lysis duplicosta Gabb, 1864: 138, pl. 21, fig. 9Sa—98c; Tryon, 1883: 112, pl. 44, fig. 25-26; Cossmann, 1903: 70; Stewart, 1927: 345-346, pl. 21, figs. 7, 7a; Anderson, 1955: 169. Stomatia suciensis variety carinifera Whiteaves, 1879: 128-129, pl. 16, fig. 5. Lysis oppansus White, 1889: 17, pl. 4, 1958: 169. Lysis suciensis var. carinifera (Whiteaves)—Whiteaves, 1903: 367, pl. 45, fig. 4. Micreschara (Lysis) duplicosta (Gabb).—Cossmann, 1925: 173 pl. 9, figs. 6, 21. Lysis duplicostata Gabb.— Wenz, 1940: 880, fig. 2587 (reprint of Stewart, 1927); Elder and Saul, 1993: pl. 2, figs. 14-15. Lysis carinifera (Whiteaves) .—Anderson, 1955S: 170. ? Lysis duplicosta carinifera (Whiteaves).—Dailey and Pope- noe, 1966: 6. Not Lysis duplicosta Gabb.—Saul and Alderson, 1951: 35—36, . 14-15; Anderson, pl. 3, figs. 3-4 [= Lysis suciensis (Whiteaves) fide Saul, 1990]. Diagnosis: Variably sculptured Lysis, with many fine cordlets or with six to eight strong cords, including prominent (often flange-like) carina on periphery; col- umella and inner lip depressed and expanded to form crescentic shelf, occupying at least one third of aperture in larger specimens. Description: Shell medium size (height up to approxi- mately 26 mm), neritiform to oe spire mod- erately low, approximately 20% of total shell height; pro- toconch 1.5 whorls, low and smooth: teleoconch approxi- mately two whorls, overall rounded, enlarging very rapidly, and medially carinate; sculpture consisting of spiral ribs, gener: ally six to eight prominent ones, “but highly atebles in number, spacing, and strength; periph- ery alw ays demarked by very strong (occasionally flange- like) carina, located anteriorly of hodial position on spire whorl and located medially on last whorl; remainder of whorls covered by spiral sculpture, 1 ranging from numer- ous closely spaced fine cordlets to several widely spaced moderately strong bien alternate in strength) cords, with interspaces smooth or bearing many sordlé ts or threads; cords just anterior and, to a lesser degree, just posterior of medial carina on last whorl commonly approaching strength of medial carina, thereby imparting either a bi- carinate or tricarinate appearance to whorl profile; aper- ture circular with a scarcely discernable anterior canal notch; abapertural edge of aperture sharply demarked by raised edge; inner lip and columella flattened, and ex- panded to form shelf; shelf moderately wide (occupying at least one third of aperture in larger specimens), sub- merged within the aperture, wrapping far past suture and attached to inside of outer lip; medial part of outer lip digitate. Lectotype: Of Lysis duplicosta UCMP 11975, height 10 mm, diameter 19.5 mm. Gabb (1864) did not indicate a holotype. Merriam (1895) recognized UCMP 11975 as the figured specimen, Stewart's (1927) statement that this is the type specimen is taken as designation of lec- totype. Paralectotype: Of Lysis duplicosta ANSP 4242. Syntypes: Stomatia suciensis variety carinifera Whiteaves, 1879, CGS 5772, a-d (Bolton, 1965). Whiteaves (1903: py 16, fig. 5) figured one of the five syntypes CGS 5772. Holotype: Of Lysis oppansus White, 1589, USNM 20115. Hypotypes: Of Stomatia suciensis carinifera, CGS 5939 (Whiteaves, 1903); Of Lysis duplicosta LACMIP 13375-13385; USNM 465555, 4685586. Type Locality: Of Lysis duplicosta, Texas Flat, near Rock Corral, from a mine shaft at a de pth of 12 m [40 ft.|, near the Placer-Sacramento Co. ae Placer Co., northern Califormia; Of Stomatia suciensis carinifera, Su- cia Island, San Juan Co., Washington. Of Lysis oppansus, Pentz Ranch, Butte Co., northern California. Distribution: Cedar District Formation, Nanaimo Ba- sin, (area 1) Vancouver Island area, southern British Co- lumbia and (area 2) Sucia Island, San Juan Co., Wash- ington; Chico Formation, Ten Mile Member on (area 5) Page 126 THE NAUTILUS, Vol. 122, No. 3 Chico Creek and Musty Buck Member along (area 6) Dry Creek, near Pentz, Butte Co., California; Chisd For- mation, (area 7) Granite Bay and Texas Flat, Placer Co. California; Pigeon Point Formation, southern sequence, (area 8) north of Pigeon Point, San Mateo Co., Califor- nia; Jalama Formation, (area 12) western Santa Ynez Mountains, Santa Barbara Co., California; Ladd Forma- tion, uppermost Holz Shale Member and Williams For- mation, Schulz Member, (area 18) Santa Ana Mountains, Orange Co., California. Geologic Age: Campanian, Discussion: The above description is based on 101 specimens; most of these are from the Pentz area (LAC- MIP loc. 24340). At any locality, most of the specimens are internal molds. Some show endobiont boreholes. One specimen, LACMIP 13385 (Figure 28), shows the shape of the protoconch. The holotype of Lysis duplicosta is a juvenile and had not yet developed the wider deck of an adult. The largest specimens of this species thus far found are from the Musty Buck Member of the Chico Formation at LAC- MIP loc. 24340 near Pentz (area 6). The specimens are from a matrix-supported pebble conglomerate richly fos- siliferous in places. The fauna, which includes scraps of cypraeids, suggests warm, shallow water. Lysis duplicosta is very rare in the coeval Ten Mile Member on Chico Creek (area 5) which probably represents deeper water than at LACMIP 24340, and the specimen from LAC- MIP loc. 23639 on Chico Creek may have been trans- ported downslope. The strength of the cords, especially of the strongest one, varies greatly between individuals. Gabb’s specific name refers to a doubled appearance of each major cord, but Gabb’s (1564) sharp, deep channel along ihe strong ribs results from the wearing or breaking off of the scales on the ribs. Dailey and Popenoe (1966: fig. 3) listed Lysis dupli- costa carinife ra from the Jalama bres at LACMIP loc. 24128. This somewhat distorted specimen (Figures 18, 19) has some stronger ribs as in L. duplicosta, but represent a strong ribbed variant of L. jalamaca. If L. duplicosta, it is the ‘geologic ally youngest specimen of fxs spe cles. Rennie (1930: 1935) described two species of Lysis from the Umzamba Saat of the ola Cape Province (formerly Pondoland), South Africa, one of which Lysis capensis Rennie, 1930, is very similar to L. duplicosta. According to Klinger and Kennedy (1980), the lower Umzamba Formation at its type locality is lat- est Santonian or earliest Campanian in age and, there- fore, similar in age to the Chico Formation near Pentz, Butte Co., California, where L. duplicosta is common. Rennie apparently had only two specimens of L. capensis which he said had six stout, sharp, spiral ribs. His holo- type is small, similar in size to the holotype of L. dupli- costa, and both apparently have a relatively narrower deck than is found in large specimens of L. duplicosta. Kase’s (1990) report of L. duplicosta in the Chats- worth Formation is based on a misidentification of L. suciensis in Saul and Alderson (1981). Lysis jalamaca new species (Figures 29-36) Lysis duplicosta Gabb.—Dailey and Popenoe, 1966: 6. Not Lysis duplicosta Gabb, 1864. Diagnosis: A Lysis with sculpture of fine spiral cordlets of alternating strength and within the aperture the shelf wrapping considerably past suture. Description: Shell medium size (height up to 26 mm), crepiduliform, with rapidly expanding whorls; spire low, and approximately 30% of total shell height; protoconch 1.5 whorls, low and smooth; teleoconch approximately two whorls, well rounded; whorls flattened adjacent to suture; periphery near mid-whorl height; suture abutting just below periphery; growth line prosocline, occasionally forming collabral wrinkles; sculpture consisting of nu- merous Closely spaced fine spiral cordlets alternating in strength and somewhat scaly; cordlets strongest at aud near periphery where two or three can be stronger be- coming cords and the scales bead-like, with approxi- mately three cordlets in interspaces; aperture ovoid, its abapertural margin sharply demarked and steeply de- scending; inner lip and columella flattened and expanded to form shelf; shelf narrow anteriorly becoming moder- ately wide medially, submerged w ithin aperture, wrap- ping past suture and attachec | to inside of outer lip. Holotype: LACMIP 13386, height approximately 35 mm, diameter 34 mm. Paratypes: LACMIP 13387 (LACMIP loc. 24137) -13388 (LACMIP loc. 24122) and SDNHM 114595, 114596 (SDNHM loc. 3405). Type Locality: LACMIP loc. 24137, Jalama Forma- tion. Figures 29-39. Lysis species. Specimens coated with ammonium chloride. 29-36. Lysis jalamaca new species. 29-30. Holotype LACMIP 13386, LACMIP loc. 24137. 29. Apertural view, vertical dimension 34 mm, horizontal dimension 30 mm. 30. Abapertural view, vertical dimension 27 mm, pies 34 mm. 31-32. Paratype LACMIP 13387, LACMIP loc. 24137 _ 31. Left-lateral view, height 18 mm, horizontal dimension 19 mm. 32. Lateral view, vertical dimension 11.5 mm, diameter 28 mm. 33. Paratype LACMIP 13358, LACMIP loc. 24122, abapertural view, height 11 mm, diameter 13 mm, 34. Paratypes SDNHM 114595 and 114596, SDNHM loc. 3405 on outer lip of volute gastropod Longoconcha eumeka Saul and Squires, 2008 (SDNHM 70974), SDNHM loc. 3405, height 133 mm, diameter 33 mm. 35-36. Posteriormost paratype SDNHM 114596 on volute shown in previous figure. 35. Abapertural view, height 13 mm, diameter 20 mm. 36. Oblique apical view, vertical dimension 17 mm, diameter 20 mm, 37-39. Lysis lomaensis new species, holotype SDNHM 67150, SDNHM loc. 3403, height 22 mm, diameter 23 mm, 37. view. 39. Apical/lateral view, vertical dimension 8 mm, diameter 2 Apertural view. 38. Abapertural 3 mm. L. R. Saul and R. L. Squires, 2008 Page 127 Distribution: Jalama Formation, (area 12) western Santa Ynez Mountains, Santa Barbara Co.; Debris Dam Sandstone, (area 13) Agua Caliente Canyon, San Rafael Mountains, Santa Barbara Co., California; Chatsworth Formation, upper part (area 16) at Lang Ranch, Ventura Co., California; Point Loma Formation, (area 19) near Carlsbad, San Diego Co., California; Rosario Formation, (area 22) Cafion San Fernando, northwestern Baja Cali- fornia, Mexico. Geologic Age: Late Campanian and early Maastrich- tian. Discussion: The above description is based on 13 specimens. The best preservation occurs in the Point Loma Formaton near Carlsbad. Lysis jalamaca is similar to L. suciensis but has coarser, more scaly costae than L. suciensis, and available specimens of L. jalamaca are smaller and less elongate than are large specimens of L. suciensis. Lysis jalamaca is also cimiilar to L. duplicosta in having variable sculp- ture and in having the strongest cords on the periphery, but L. jalamaca is less angulate at the periphery and has weaker cords there. In addition, L. jalamaca ditters from L. duplicosta by being more elongate, having finer sculp- ture, and having the shelf broader posteriorly. The speci- men (LACMIP 13375, Figures 18, 19) from the Jalama Formation has ribbing similar to L. duplicosta, but its shelf appears to wrap farther onto the interior of the outer whorl than is usual for L. duplicosta. Etymology: The specific epithet, a name in apposi- tion, reflects the new species occurrence in the Jalama Formation, Santa Barbara Co., California. Lysis lomaensis new species (Figures 37-39) g Diagnosis: Lysis with low spire and last whorl bearing numerous flanged carinae separated by wide interspaces bearing several spiral threads. Description: Shell medium size (up to 17 mm height and 30 mm diameter, same specimen), neritiform (last whorl rapidly expanding); spire very low, approximately 10 to 15% of shell height; protoconch 1.5 whorls, low and smooth; teleoconch 2 to 2.5 whorls, very carinated; su- ture abutting just anterior to periphery; sculpture con- sisting of up to 11 strong cords; cord at periphery and next three cords posterior of periphery very thin, flanged, and protruding (with cord at periphery protruding most); posteriormost part of last whorl (in vicinity of suture) with approximately three low scaly cordlets: up to three beaded to lowly spinose cordlets anterior to medial ca- rina at periphery; interspaces between all cords wide and bearing up to seven spiral threads (occasional thread can locally dev elop into small flanged cord); area anterior to medial carina can be nee | with only fine cordlets; aperture circular with margin sharply demarked by raised edge; shelf moderately wide, wrapping past suture and attached to inside of outer lip; posterior half of outer lip digitate. Page 125 THE NAUTILUS, Vol. 122, No. 3 Holotype: SDNHM 67150, height 22 mm, diameter 23 mm. Paratype: SDNHM 67152 from SDNHM loc. 4071. Type Locality: SDNHM loc. 3403. Distribution: Basal San Francisquito Formation, (area 14) Warm Springs Mountain, Los Angeles Co.; Point Loma Formation, (area 19) San Diego Co., Cali- fornia: Rosario Formation, (area 21) five miles east of La Mision, northwestern Baja California, Mexico. Geologic Age: Latest Campanian to late Maastrich- tian. Discussion: Five specimens were studied. Preserva- tion is generally excellent for three specimens from the Point Loma Formation near Carlsbad, although the pro- toconch of the holotype is imperfectly preserved. The incomplete, crushed specimen from Warm Springs Mountain is from beds of latest Maastrichtian age at the base of the San Francisquito Formation (LACMIP loc. Figures 40-50. 14310). Although the abapertural side is not available, the flatness of the apical whorls suggests that it is prob- ably L. lomaensis. The new species is most similar to Lysis duplicosta but available specimens are smaller than a large L. dupli- costa. Lysis lomaensis also has a lower spire, more cari- nate last whorl (especially posterior to the medial carina), and spiral threads on interspaces between carinae. Abaperturally, L. lomaensis is similar in shape and sculpture to Garzasia intermedia, but L. lomaensis has a lower spire and in the apertural view the shelf margin is less arcuate, not sigmoid, and the shelf does not spiral into the umbilicus. Group of Lysis suciensis The group of L. suciensis differs from that of L. dupli- costa in havi ‘ing a more elongate aperture and finer Scal sculpture. Lysis suciensis (Whiteaves, 1879) (Figures 40-50) Lysis suciaensis (Whiteaves, 1879), Specimens coated with ammonium chloride. 40-41. Hypotype LACMIP 13359, LACMIP loc. 26020. 40. Apertural view, vertical dimension 44 mm, horizontal dimension 40 mm. 41. Abapertural view, height 25 mm, diameter 43 min, 42-43. Hypotype LACMIP 13390, LACMIP loc. LO711. 42. Abapertural view, height 1S mm, diameter 26.5 mm. 43. Lateral view, vertical dimension $8 mm, diameter 26.5 mm, 44. Hypotype LACMIP 13391, LACMIP loc. 10095, lateral view, vertical dimension 5 mm, diameter 12.5 mm. 45. Hypotype LACMIP 13392, Cafon San Fernando, 32 km southeast of El Rosario, Baja California, Mexico, abape rtural view, height 1S mm, diameter 21 mm. 46-50. Hypotype LACMIP 10495, LACMIP loc. 26020. 46-47. Vertical dimension 70 mm, horizontal dimension 38 mm. 46. Apertural view. 47. Abapertural view. 48. Right-lateral view, vertical dimension 70 mm, horizontal dimension 25 mm. 49. Slightly oblique right-lateral view, vertical dimension 70 mm, horizontal dimension 27 mm. 50. Lateral view, vertical dimension 25 mm, horizontal dimension 36 mm. L. R. Saul and R. L. Squires, 2008 Dace 126 Page 129 Stomatia suciensis Whiteaves, 1S79: 128-129, pl. 16, fig. 4. Lysis suciensis (Whiteaves) —Whiteaves, 1903: 367, pl. 45, fig. 3. Stecheson, 2004: 60-62, pl. 2, figs. 4-5. Lysis californiensis Packard, 1922: 431, pl. 37, figs. 2-3; Sta- dum, 1973: pl. 2, fig. 12. Lysis duplicosta Gabb.—Saul and Alderson, 1981: 36, pl. 3 figs. 8-4. Not. Lysis duplicosta Gabb, 1864. Diagnosis: Medium to large Lysis, shell elongate with fine, usually wavy cordlets, sculpture obsolete on large specimens. Description: Medium to large (height up to $4 mm), crepiduliform, elongately exp: ae d; spire moderately high to low, approaching ‘horizontal coiling, and approxi- mately 30 to 35% of total shell height; protoconch ap- proximately 1 to 1.5 whorls, low and smooth; teleoconch approximately 1.5 to 2 whorls, whorls subcarinate to rounded; periphery near one-third whorl height; suture abutting just anterior to periphery, descending; crowth line slightly to moderately parasigmoid il, w ah sinus at posterior end of outer lip on large, smooth individuals; sculpture consisting of fine narrow cordlets, usually wavy, alternating in strength, with narrow interspaces; aperture elongate ovoid, its abapertural margin sharply demarked and steeply descending; inner lip and col- umella flattened and expanded to form crescentic and moderately wide deck submerged within aperture, pos- terior end attached to inside of outer lip; inner deck margin concavely curved. Syntypes: Of Stomatia suciensis CGS 5771, ad. Lectotype (here chosen): Of Stomatia suciensis CGS 5771, height 66 mm. eon Paralectotypes: Of Stomatia suciensis CGS 5771a— CGS 5771d. Holotype: Of Lysis californiensis UCMP 12287. Paratype: Of Lysis californiensis UCMP 12288 Type Locality: Stomatia suciensis, Sucia Island, San Juan Co., Washington. Of Lysis californiensis, UCMP loc. 2167, Santa Ana Mountains, Orange Co., California. Figured Specimens: LACMIP 10495, 13359, from LACMIP loc. 26020; LACMIP 13390 from LACMIP loc. LOT11; LACMIP 13391 from LACMIP loc. 10095; LACMIP 13392 from the Rosario Formation, (area 22) 26 km east of coastline at elevation 200 m on west side of Cafion San Fernando, 15 km north of Mesa San Carlos, approximately 32 km. southeast of El Rosario, Baja Cali- fornia, Mexico. Distribution: Upper Cedar District Formation, (area 1) Denman Island, Gulf Islands, Prien Columbia; Lower Cedar District Formation, (area 2) Sucia Island, San Co., Washington; Chico Serna (area 7) Granite Bay, Placer Co.; Jalama Formation, (area 12) western Santa Ynez Mountains, Santa Barbara Co.; Chats- worth Formation, (area 16) Bell Canyon and Dayton Canyon, Simi Hills, Ventura Co., California; Ladd For- mation. upper Holz Shale Member, (area 15) Santa Ana Mountains, Orange Co., California; Rosario Formation (area 22) at Canon San Fernando, 26 km SE : El Ro- sario Baja California; Valle Formation, (area 23) 10 kin north from Punta Abreojos, Baja California Sur, Mexico. Geologic Age: Late early Campanian and early Maas- trichtian. Discussion: The above description was based on LOT specimens. The deck of smaller specimens is rather nar- row and attaches to the base of the previous whorl. Small specimens are rounded, larger specimens more elongate. In larger specimens the shelf is broader and its posterior end attaches to the inside of the outer lip. Several speci- mens, especially the larger specimens, have an obtusely biangulate whorl abapical to the periphery. Most speci- mens from Sucia Island are of small size and badly weathered, but Whiteaves’s largest specimen (1879: 129, pl. 16, fig. 4), here designated as lectotype, was indicated by him to be “two inches and a half in length” (i.e., 66 mm). The specimen from Bell Canyon (area 16) of Fig- ures 46— 50, lacking its spire and w ith its outer lip broken. is 75 mm high. The ribbing on larger specimens from the Chatsworth Formation in Bell and Dayton canyons, Simi Hills (area 16) tends to become fainter toward the outer lip espe- cially on the posterior slope. The specimen figured (Fig- ures 40, 41) retains ribbing around the pe riphe ry, but the ribs fade posteriorly toward the aperture. Height range of the smoother specimens is 45 to 76 mm (incomplete) making these the largest specimens thus far found of Lysis. The largest of these specimens are larger than representatives of most species of Crepidula; only Gran- dicrepidula princeps (Conrad, 1856), attains a larger size. Genus Garzasia new genus Type Species: Garzasia diabla new species from the “Garzas Sand” Member of the Moreno Formation, Stani- slaus Co., California; mid Maastrichtian. Description: Very low to moderately high spired, Ca- lyptraca-like shell with weak to strong spiral cords, ap- erture very broadly expanded and forming base of shell, shelf attached marginally, surrounding the umbilicus, and spiralling into it. Discussion: Garzasia is most similar to Lysis but dif- fers from the latter in that the deck surrounds and spirals into an umbilicus somewhat as in Calyptraca but with a wider more open umbilicus. It differs from Calyptraea in having a sturdier shelf attached marginally as in Trochita. It differs from Tro thita (Figure 71) in having the shelf crescentic with an arcuate to slightly sigmoidal shelfal edge, the axis of spirialing of its shelf off center, and external ribbing spiral rather than protractive. Etymology: The genus Garzasia is named for the “Garzas Sand” and Garzas Creek, Stanislaus Co., Cali- fornia. Garzasia intermedia (Cooper, 1894) (Figures 51-54) Page 130 Figures 51-58. Stomatia intermedia Cooper, 1894: 46, pl. 3, fig. 43 [re- figured in Yates, 1903: pl. 3, fig. 43]. Lysis intermedia (Cooper).—Anderson, 1958: 170; Coan, 1981: 165, fig. 12 [reprint of Cooper's pl. 3]. Diagnosis: Large Garzasia, haliotitorm, carinate on periphery, sculpture elsewhere consisting of several low spiral cords separated by wide interspaces bearing up to 10 spiral threads, shelf very broad, with arcuate to slightly sigmoidal margin. Description: Large size (height up to 37 mm, diam- eter 65 mm, same specimen); haliotiform with very rap- idly expanding last whorl; protoconch unknown; spire highly variable in elevation, ranging from 50 to 85% of total shell height; suture abutting below periphery; growth line prosocline; sculpture consisting of commonly five spiral cords; periphery carinate and bearing stron- gest cord; peripheral carina located anteriorly or medial position on spire whorl but medially on last whorl: adapi- cal to medial carina two to three medium- strong cords with wide interspaces bearing up to 10 spiral heeade just abapical to medial carina several closely spaced and scaly medium strong cords; aperture circular; inner lip and columella flattened to slightly concave and e ae d to form very wide crescentic shelf submerged within aper- ture; posterior end of shelf surrounds win biicis and spi- rals into it. THE NAUTILUS, Vol. 122, No. 3 Species of Garzasia new genus. 51-54. Garzasia intermedia (Cooper, 1894). 51-52. Plasto-lectotype of CASG 609, Point Loma Formation. 51. Apertural view, vertical dimension 21.5 mm. 52. Apical view, diameter 19 mm. 53-54. Hypotype SDSNH 67149, SDSNH loc. 3403. 53. Apertural view, vertical dimension 61.5 mm, diameter 64 mm. 54. Left-lateral view of spire above posterior part of aperture, height 20 mm, diameter 64 mm. 55-58. Garzasia diabla new species. 55-57. Holotype LACMIP 13393, LACMIP loc. 22588. 55. Apertural view, vertical dimension 36 mm, diameter 37 mm. 56. Abapertural view, vertical dimension 34 mm, diameter 37 mm. 57. Side view of spire above posterior part of aperture, height 1S mm. diameter 37 mm, 58. Paratype LACMIP 13394, LACMIP loc. 26353, slightly oblique left-lateral view (partial specimen), height 29 mm, diameter 9 mm. Lectotype: CAS 609 (formerly CSMB 13742), height 6.4 mm, diameter 22.6 mm (incomplete). Type Locality: Point Loma, San Diego Co., Califor- nia. Figured Specimen: SDSNH 67149 from SDSNH 3403. Distribution: Point Loma Formation, (area 19 and 20) San Diego Co., California. Geologic Age: Latest Campanian and early Maas- trichtian, Baculites lomaensis zone. Discussion: The above description was based on six specimens, of which preservation is generally good to excellent. Cooper's (1894) figure 43 line drawing is actually a combination of three specimens. The lectotype (CASG 609) is designated by us and is photographed here (Fig- ure 51) for the first time. Garzasia intermedia difters from G. diabla new spe- cies in achieving larger size, having weaker carinae with the medial carina being most prominent, and having spi- ral threads in interspaces between carinae. Garzasia intermedia is somewhat similar to Sigapatella Lesson, 1830, some species of which [e.g., Sigapatella novaexelandiae (Lesson, 1830)| have spiral sculpture, an off-centered apex, and a well-developed “false” umbili- L. R. Saul and R. L. Squires, 2008 Page 13] cus. The umbilicus of Garzasia appears to form as the widely expanded, flattened columella is attached medi- ally around the axis of coiling forming a broad shelfal area. In the holotype of G. intermedia and some other specimens this area is an open, funnel shape, but in oth- ers the area is more or less filled, resulting in shelves with differing “umbilical” development from a moderate to deep depression. The shelfal edge of Garzasia differs from that of Sigapatella in being sigmoidal rather than arcuate. A large specimen SDNHM 67149/3403 has faint markings at each end of the shelf which resemble muscle scars. Garzasia diabla new species (Figures 55-58) Diagnosis: Medium sized, Calyptraca-like, with mod- erately high spire, having three to four equal-strength carinate ribs with wide interspaces. Description: Medium to moderately large size (up to 30 mm in height), Calyptraeca-like shell with rapidly ex- panding last w vhorl: spire moderately high, approximately 40 to 50% of total shell height: protoconch missing; te- leoconch approximately 1.5 whorls, carinate; pe riphery near one-half whorl height; suture abutting at periphery; sculpture consisting generally of three to four « equal- strength and equally spaced carinae with very wide in- terspaces; aperture circular with its ab yapertural margin sharply demarked and steeply descending; inner lip mar- gin slightly sigmoidal; posterior end ateched to inside of outer lip directly beneath suture; shelf spiraling into um- bilicus. Holotype: LACMIP 13393, height 18 mm, diameter 37 mm. Paratype: LACMIP 13394 from LACMIP loc. 26353. Type Locality: LACMIP loc. 22558. Distribution: Moreno Formation, “Garzas Sand” Member. (area 9 and 10) Merced and Stanislaus coun- ties, California. Geologic Age: “Mid” Maastrichtian. Etymology: The species is named for its occurrence in the eastern foothills of the Diablo Range, Merced and Stanislaus counties, California. The specific epithet dia- bla is used as a name in apposition. Discussion: The above description is based on eight specimens. Preservation of the known specimens of this species is poor. Except for the holotype, all the speci- mens are internal molds. This species is very similar to G. intermedia from which it differs in having a generally lower but more roundly inflated, spire that is appare ently not variable in height. higher last whorl, much stronger sculpture, and the basal flange wrap into the aperture to meet the ap- ertural edge of the shelf. EVOLUTIONARY IMPLICATIONS The presence of both fine-ribbed and coarse-ribbed Ari- adnaria and Lysis suggests that species of Lysis may have been derived from Ariadnaria. During the e volution of Lysis, the inner lip broadened (as did the columella) and apparently moved deeper into the aperture, thereby al- lowing more room for the foot to grasp hard substrate. These changes, assumed to be a function of the rapid expansion of the aperture, were necessary in order that the aperture could accommodate a larger foot for attach- ment to a hard substrate. The resultant shell shape was crepiduliform. Modern trichotropids studied by Yonge (1962) in Puget Sound thrive on unstable shell beds. ay some Cretaceous trichotropids resembled the modern trichotropids in (1) being filter feeders, (2) living on firm substrates, and ( 4 being protandrous hermophodrites, they would have had ehan acteristics that allowed them to evolve toward the less vagile calyptraeids or crepidulids. The inner lip of trichotropids is homologous to the internal deck or shelf of calpytraeids, and the develop- ment of the shelf in Lysis serves as a pattern for the development of the flat shelf in Crepidula Lamarck, 1799, and of the shelf in Garzasia for the spiral shelf in Calyptraea Lamarck, 1799. Trichotropid and crepidulid sculpture is predominantly spiral (i.e., in the direction of coiling), but some calyptraeid sculpture is radial or pro- tractive. The shell exterior of Lysis suciensis group (i.e., Lysis suciensis lineage) is fine-ribbed to relatively smooth; in- teriorly the columella and inner lip are flattened and broadened into a crescentic shelf which, as it lengthened posteriorly, came to resemble a shelf of ” Crepidula. Ex- cept for its large size, Lysis suciensis resembles the ear- liest species from the Pacific slope usually assigned to Crepidula; namely, ‘Crepidula’ pileum (Gabb, 1864), which ranges from middle Eocene to Oligocene in Cali- fornia to Washington. Stewart (1927) and Hoagland (1977) slightly extended the range of “C.’ (Spirocrypta) pileum by synonymizing with it the late early Eocene ae rypta inornata ( (Dickerson, 1916) and the middle to late Eocene Spirocrypta dickersoni (Weaver and Palmer, 1922), but Vokes (1939) considered Spirocrypta inornata of ‘ eee age to have “a funnel-like pro- cess due to the upward curving of the posterior portion of the septa” (Vokes, 1939, pl. 13, fig. 7) rather than the less apparent “umbilicus” of S. pilewm (see discussion below). Gabb (1864) originally placed ‘Crepidula’ pileum in the invalid genus Crypta Humphrey, 1797 (a synonym of Crepidula), and subgenus a ice Gabb, 1864, of which ‘C.’ (S.) pileum is the type species. ‘Crepidula’ (S.) pileum, ihich is common in the Tejon Formation of middle Eocene age in Live Oak Canyon, Kern Co., Cali- fornia, has been figured several times (e. g., Gabb, 1864: pl. 29, figs. 233, 233a—b; Stewart, 1927: pl. 29, figs. 2-3; Anderson and Hanna, 1925: pl. 13, fig. 7; Clark, 1935: pl. 4, fig. 19; Wenz, 1940: figs. 2660a, b [reprint of Stewart]. A large specimen from the late Eocene is figured by Page 132 THE NAUTILUS, Vol. 122, No. 3 ae and Weaver (1963, pl. 24, fig. 11). Weaver's (1943: 724, pl. 71, fig. 16) ilustaton of Calyptraea pee (Conrad) is a lapsus and is a posterior-end-up, apertural view of ‘Crepidula’ pileum. On page 356 he correctly lists the figure as Crepidula parent Gabb’s (1864: pl. 29, fig. 233a) and Stewart’s (1927, pl. 29, fig. 3) show the shelf. ren s figure is a fascimile and Stewart's is a photograph that has een reproduced in other dis- cussions of Spirocrypta (e.g., Wenz, 1940: 903, fig. 2660a). Gabb’s and Stewart's figures are based on lecto- type ANSP 4221, but, unfortunately, the shelf of this specimen is broken. Both figures create the false impres- sion that there is a sinus near both ends of the shelf and that the middle part protrudes and is concave. An addi- tional representative specimen (hypotype LACMIP 13395) was cleaned by the senior author and is illustrated in Figures 59-61. Its shelf (Figure 59), which is unbro- ken, is slightly sigmoidal and long on the left side (or anterior end) ands shorter on the right side (or posterior end). There is also a slight conv exity of the somewhat sinuous shelf as it approaches the posterior end of the aperture and the shelf margin spirals over slightly to form an indication of an aaniheue: In addition, the shelf is also narrower and closer to the shell margin on the left/ Figures 59-73. (Gabb, 1864), hypotype LACMIP 13395, LACMIP loc. 22 Abapertural view. 61. Lateral view. 62-64. ‘Calyptraea’ diegoana (Conrad, 1855), hypotype LACMIP 13458, LACMIP loc. 22340. 62. Apertural view, vertical dimension 29 mm, diameter 31 mm. 63. Apical view, diameter 31 mm. 64. Right lateral view, height 13 mim, diameter 31 mm, 65-67. ‘Calyptraca’ aperta Solander in Brander, 1766, hypotype LACMIP 13396, LACMIP loc. 7333. 65. Apertural view, vertical dimension 11 mm, diameter 11 mm, 66. Apical view, diameter 11 mm. 67. Right-lateral view, height 7 mm, diameter 11 mm. 68-70. Calyptraca chinensis (Linnaeus, 1758), Recent, type-species, hypotype LACM 161651, Cherbourg (Manche anterior and noticeably farther from the shell margin (deeper into the aperture) on the right/posterior. The shelf of °C. pileum thus, as noted by Gabb (1864), spirals inward toward the apex. Gabb’s figure (pl. 29, fig. 233b) in part illustrates this, as does Figure 59. Although Stew- art (1927) synonomized Spirocrypta with Crepidula, Gabb’s description of Spirocrypta recognizes this very important characteristic, which helps to distinguish Spi- rocrypta from Crepidula, In modern Crepidula forni- cata, the shelf does not spiral into the whorl apex. Aperture/shelf features of Spirocrypta pileum and S. inornata resemble those of the early Paleocene Spirogal- erus lamellaria Finlay and Marwick, 1937, from New Zealand, in that the shelf of $. lamellaria is also narrower and closer to the shell margin on the left/anterior and noticeably farther from the shell margin (deeper into the aperture) on the right/posterior. Our proposed evolutionary relationship between tri- chotropines and calyptraeids differs from any previous author's proposal. Hoagland (1977), for example, in her study of Crepidula Lamarck, 1799, rejected trichotropids as direct ancestors of calytraeids and crepidulids. Hoag- land (1977) ee that although Trichotropis Broderip and Sowerby, 1829, and Crepidula had a common an- Fossil Calyptraeidae. 59-64. Comparative Eocene Crepidula and Calyptraea species. 59-61. Spirocrypta pileum 356, vertical dimension 3 mm, diameter 9 mm. 59. Apertural view. 60. ), Normandie, France. 68. Apertural view. vertical dimension L5 mm, diameter 14.5 mm, 69. Apical view, diameter 14.5 min. 70. Right-lateral view, height 6 mm, diameter 14.5 mm, 71-73. Trochita trochiformis (Born, 1778), Recent, type-species, LACM loc, lot 75-41, Purmalin, west of Isla Teleon, Gulf of Corcovado, Chiloe Province, Chile — intertidal. 71, Apertural view, vertical dimension 40 mm, diameter 38 mm. 72. Apical view, diameter 38 mm. 73. Right-lateral view, height 18 mm. L. R. Saul and R. L. Squires, 2008 Page 133 cestor, Trichotropis has no direct relationship to Crep- idula because Trichotropis is boreal, living in cold and deep water and has rough sculpture, whereas Crepidula had its origin in the Panamic region and is relatively smooth sculpture od. Her implications that Trichotropis is a deep, cold-water gastropod and that Crepidula is a shallow, warm-water gastropod are misleading. Modern Trichotropis cancellata (Hinds, 1843) is intertidal in Alaska, British Columbia, and Washington to depths of 104 m off Alaska, 165 m off British Columbia, and 274 m off Washington (LACM Malacology collection). In the southern part of T. cancellata’s range, it is in cool- temperate, not boreal waters. The question is, however, not where trichotropids live now but where they were during the Late Cretaceous, when there were no polar ice caps and the subtropical and temperate zones were much wider. Trichotropids and calyptraeid-like gastro- pods co-existed during the Late Cretaceous in the study area, and both forms lived in relativ ely warm, shallow waters. Although the Coniacian-Santonian faunas of northern California are noticeably cooler than Turonian faunas (Saul and Squires, 2008) of that area, they would have been temperate. Hoagland (1977) mentioned that anatomical features of T) ichotropis suggest affinities to calyptraeids, but she believed that similarities of life his- tory and niche between Trichotropis and Crepidula are convergent. She suggested that Crepidula is derived from some form of * ‘calyptraeid stock” that, in turn, was derived from Trochita Schumacher, 1817, reported by Wenz (1940) to range from Eocene to Recent. She did not provide any geologic time parameters as to when these derivations took place, but indicated that Shimer and Shrock (1959) recorded the first calyptraeids as “lower Cretaceous.” The “lower” seems to be a minor lapsus: all printings of Shimer and Shrock from first 1944 to last 1989 list range of Calyptraca and Crepidula as “U. Cret—Recent” and Crucibulum as “Tert.—Recent.” Wenz (1940: 902), however, questionably included the peculiar looking genus Galericulus Seeley, 1861 from the Upper Greensand (Lower Cretaceous upper Albian) of England in calyptraeids. It does not resemble any calyptraeid we have studied. Seeley (1861) named Cre pidula cooksoniae also from the Upper Greensand, but Hoagland (1977: 395) found it “unconvincing” as a calyptraeid. The Campanian to Midas mchivan Damesia Holzaptel, 1885. of Europe and Tennessee, has been regarded as a calyptraeid by some workers (e.g., Sohl, 1960), but Dock- ery (1993) assigned Damesia to the ese elloids. Bandel and Riedel (1994) reviewed P lacement and content of Calytraeoidea, and in comparison to Hoag- land’s (1977) study, they arrived at a different faints content and different relationships between the families. Their Calyptraeoidea consisted of two families: Calyp- traeidae containing genera Calyptraea, Crucibulum, and Crepidula; and Hipponicidae containing Cheilea, Hip- ponix, Neojanaca, and Thylacus: but both families ex- cluded trichotropids. According to Bandel and Riedel (1994). all genera in Calyptraeidae lack a probosis and are obligatory filter feeders. However, in Hawaii Ulbrick (1969) reported algae grazing, in addition to filter feed- ing by Crucibulum spinosum Sowerby, 1824. ‘Cretaceous trichotropines probably had several char- aucun that lysines would have inherited from them: ) filter fee ding, 2) living epifaunally, probably on a hard nas 3) brooding egg capsules from which plank- totrophic larvae hatch, and 4) being protandrous her- maphrodites. Characteristics the lysines would pass on to calyptraeids. Lysines’ first recognizable difference from the trichotropines i is the increase in e Xpansion rate of the whorl, especially of the last whorl, resulting in an en- larged aperture in which the columella is broadened. The earliest bro: idening and enlargement were moderate and only become striking as geologically younger species be- gan to have a very disinetne morphology. Until con- nected with its ancestry, Lysis was difficult to pa The shell also became more flattened and limpet-like, « the aperture enlarged. Hoagland ( 1977) ie Crepidula hochstetteriana Wilckens (1922: 5-6, pl. 1, figs. 9a, b) as being the ear- liest Crepidula because it was reported from the calcar- eous conglomerate stratigraphic unit of the Lower “Amuri Group” in the “Amuri Bluff area, northeastern South Island, New Zealand (Wilckens, 1922: Warren and Speden, 1978). Modern spelling of “Amuri” is Haumuri, and the Cretaceous strata are referred to the Mata Se- ries. Woods (1917) reported that these strata at Haumuri Bluff contain the bivalves Inoceramus australis Woods, 1917 and Inoceramus pacificus Woods, 1917, which ac- cording to Wellman (1959) are limited to the Piripauan Stage of latest Coniacian to Santonian age. Warren and Speden (1978) noted “problems” with the early collections from this area but nevertheless, listed ocala eg hochstetteriana (Wilkens, 1922) from the Campanian Okarahia Sandstone of the Mata Series. De- scribed conditions of collecting suggested a strong pos- sibility of co-mingling of material from different strata and that the only known specimens of M. hochstetteriana might not be from the Mata Series. In search of the type specimen, we contacted three extraodinarily helpful New Zealanders: A. G. Beu, A. Grebneff, and i D. Stil- well. Their email communications (2006) indicated that M. hochstetteriana is not from the Okarahia Sandstone, and is not of Cretaceous age. Beu found the type speci- men (GNS TM2608) in the New Zealand Geological Survey collections. Fortunately, the type specimen was in a large enough block of matrix to take a sample for mi- crofossil dating, He enlisted the aid of G. Wilson who dated the dinoflagellates as late Oligocene, at the oldest. Ian Raine, who looked at the rich spore-pollen assem- blage also from the sample, found Acacia pollen, which is not known earlier than Miocene in New Zealand. Mio- cene strata crop out above the Mata Series, and some of the original material sent to Wilkens was apparently from beach boulders derived from younger strata ove -rlying the Cretaceous Mata Series. This Neogene age for Wile i ns’ species is much more likely conside ‘ring that the shelf Page 134 THE NAUTILUS, Vol. 122, No. 3 covers nearly one-half of the aperture (Hoagland, 1977 380). Hoagland published more papers on Crepidula and, in 1986, she revised several items of her 1977 paper but did not design a new evolutionary pathway. We differ from Hoagland (1977) in that we believe it is difficult to go from the centrally oriented apex plan of Trochita (Figure 71) or Calyptraea (Figure 68) with its spiraling shelf and get to Crepidula with its eccentric spire and apparently unspiraled deck. Whether or not Lysis species are an- cestral to any modern Crepidula, younger species of Ly- sis achieved a crepiduloid form with a respectable shelf by wrapping the posterior end of the shelf onto the inside of the outer whorl. Furthermore, from this, the calyp- traeid form appears to have evolved with the develop- ment of an “umbilicus” in Garzasia intermedia making Cooper's specific name remarkably prescient. At about the Campanian/Maastrichtian boundary, de- velopment of the spiral shelf of Garzasia resulted in a shelf that appears to be a pattern for development of shelves in Calyptraea and perhaps Crucibulum. The very broad, depressed spiraling inner lip of Garzasia appears to provide a likely pattern for development of the shelf of Calyptraea or Crucibulum, not for the more decklike shelf of Crepidula. The earliest reported Calyptraea on the Pacific slope is ‘Calyptraea’ diegoana (Conrad, 1855) which ranges from middle Paleocene to Oligocene and occurs from California to Washington and easternmost Russia (Squires, 1987). A representative specimen from the Te- jon Formation Eocene is shown in Figures 62-64. Figure 62 me ays the shelf of this species, which is similar to that of ‘Calyptraea@ aperta Solander in Brander, 1766 from the Eocene of Europe (Figures 65-67). Both of these species have often been referred to Trochita Schu- macher, 1517 (type species Turbo trochiformis Born, 778), but their shelves (Figures 62 and 65) spiral from an off-center position roughly a quarter of the diameter in from the aperture edge. A so-called pseudoumbilicus and 65 could result from reduction of an umbilicus such as that of Garzasia. The shelves of ‘C. diegoana and ‘C. aperta are narrower than shelves of Garzasia and expand across about one third of the aperture, They differ distinctly from that of Trochita (Figures 71-73) ), which has a sturdy spiraling internal shelf extending from the centered axis of coiling to the outer shell margin, giving the impression of dividing the circular aperture in half. ‘Calyptraea’ diegoana does not appear to be a direct descendent of Garzasia. Its similarity to ‘C.’ aperta sug- gests an ancestor in the Old World Tethyan Sea and, like many of the other Early Cenozoic mollusks. ( (Squires, 1987, 2003), it probably arrived onto the Pacific slope via at the upper end of the shelves in Fi igures 62 a circum-equatorial current. These species ‘C.’ diegoana and ‘C. Calyptraea chinensis (Linnaeus, 1758), the type species of Calyptraea. In C. chinensis, the thin fragile shelf (Fig- ure 68 occupies about a quarter of the apertural circle. aperta do not have shelves similar to that of It arises from the apex with a folded-over edge that forms the umbilicus. At the open end of the umbilicus, the shelf edge abruptly veers counter to coiling direction and approaches the apertural margin at an acute angle. The total shelfal edge is sickle- shaped with a ligne handle (the umbilical edge) and a long curved blade (the outer margin of the shelf). This leaves a deep notch between the attachment of the shelf to the shell and a delicate, lobate shelf. Trochita is present in modern eastern Pacific slope faunas from Mazatlan, Mexico to Valpariso, central Chile. It occurs in lower to middle Miocene strata as far north as the La Panza Range, San Luis Obispo Co., is present in the Kern River section, Kern Co. (Addicott, 1970) and the Topanga Formation, Santa Monica Moun- tains, Los Angeles Co., California. It has a thick shell with protractive ribbing and, as mentioned above, a sturdy shelf. GLOBAL DISTRIBUTION OF CRETACEOUS LYSIFORM GASTROPODS Both ‘Crepidula’ and ‘Calyptraea’ have been reported from Campanian and Maastrichtian age strata from throughout the world. Preservation of most specimens makes identification of them problematical. Some of these gastropods have proven to belong to other families or not to be of Cretaceous age. Others need verification. Our tally of occurrences is doubtless incomplete. Although stated above as “throughout the world,” these gastropods are recovered from areas that were probably temperate to tropical. Verified lysines are all younger than Turonian and older than Eocene. Classing Spirogalae rus as a lysine ides the only Paleosene record of this subfamily. Late Cretaceous occurrences suggest that the calyptraeids developed in several geo- graphic places from widely distributed trichotropids evolving into Lysinae. An example of this is the evolving shape of Lysis. Although on the Pacific Slope calyptraei- form calyptraeoids dev ‘eloped from a crepiduliform ca- lyptraeoid, the calyptraeiform has been recognized more widely geographically, but not earlier than Coniacian. Cretaceous calyptraeiform calyptraeoids are more widely recognized geographically than are crepiduliform calyptraeoids. Europe has calyptraeiform occurrences but no crepiduliform occurrences. Africa has calyptraei- form occurrences in the north and crepiduliform occur- rences in the south. African occurrences are close in time to those of the Pacific Slope. North America (exclusive of the Pacific Slope) has a very few reports along the Gulf Coast of calyptraeiform specimens. The Pacific : Slope has both calyptraeiform Garzasia and crepiduliform Lysis. South America has calyptraeiform species. Japan some- what mirrors the Pacific Slope. Its known lysine is of early Maastrichtian age, but the additional presence of a trichotropid ( (Kase, 1990) similar to A. obstricta suggests that lysines were developing there roughly synchronously with those along the Pacific Slope. New Zealand has no L. R. Saul and R. L. Squires, 2008 105 85 80 75 70 65 60 55 350 302520 15 5 CRETACEOUS PLA ENE gies nl ALBIAN CENO- |TUR-]| CO- CAMPANIAN oe PALEOCENE E 5 OLIGOCENE MIOCENE PLIOJ. IMANIANT ON. [NIA EN | ? _ ‘Calyptraea’ a Calyptraea 5 7 RAEIFORM | Sigapatella Trochita. —___________» 6 | | 4 G apa ae | | apiaaaie ts ane | ee ? Grandicrepidula 7 _ Lysis >» CREPIDULIFORM/ —_———> ? cepidula Spirocrypta / Maoricrypta ZZ muscle scars i => = = 1 TRICHOTROPIDAE Trichotropinae Ariadnaria Figure 74. Generalized proposed evolutionary trends of calyptraeoids. Time scale after Gradstein et al., 2004. Calyptraeids and crepidulids probably evolved from several trichotropids. Lysis or Lysis-like fossils of Coniacian to Maastrichtian age have been described from California, and from Campanian to Maastrichtian age from southern Africa and Japan. 1. Turriculate gastropods having gill capable of filter feeding, sedentary adult life on hard eubaeats: Probably capable of copulation, protandrism, and brooding of young. 2. Broadening of columella and inner lip. Enlarging final whorl. 3. Atti achment of posterior shelf end to interior of outer lip to develop crescentic shelf. 4. Some species developed * "ana bilions” in spiraling shelf as in Garzasia. 5. “Umbilicus” closed or nearly so in Eocene ‘Crepidula’ and moving toward more central position in more Grcdler base. 6. Broad shelf, spiraling from near center of base. Shelf edge nearly straight. 7. Spiral shell with low spire, round base. Shelf edge extremely sigmoidal. * = Pleistocene. Stages abbreviated are Turonian, Coniacian, Santonian, Mansel ies, known Cretaceous lysines but does have the youngest known calyptraeiform lysine. Maastrichtian southwestern France, has a spiraling shelf similar to that of Eocene ‘Calyptraea’ aperta except that EUROPE: Crepidula mytiloidea Bellardi and Mich- elotti, 1840 from Villavernia near Tortona Italy was listed among Nomina Dubia by Hoagland (1977) as being from the Cretaceous. Bellardi and Miche lotti were describing a Tertiary fauna. Because the specimen and ‘llinsivaaone are so small, Hoagland could not verify that this species was a Crepidula. Calyptraea cretacea (dOrbigny, 1842: 390, pl. 234, figs. 1-3) [Tnfindibubur| was examined by Kollmann 2005) who determined that d’Orbigny’s material was from Campanian of C harente-Maritime, southern France, and that the specimen identified as C. cretacea by Delpey (1942: 165, fig. 1) southwestern France. Poor preservation of dOrbigny’s type caused Kollmann (2005: 172, pl. 18, fig. 18) to fetes to it as “Calyptraea s. lato cretacea (V Orbigny, 1843), species dubia.” aie s illustration shows no shelf. Koll- mann considered both specimens to be only Calyptraea sensu lato. These specimens expand their w horl diameter much less rapidly and have much higher spires than specimens of Lysis and Garzasia from the Pacific Slope of North America. Calyptraea depressa Delpey, 1942 (p. 165, fig. 2) from was from Maastrichtian of its shelfal margin is concavely arcuate between rim and whorl center. Kollmann and Odin (2001: 446, pl. 1, figs. 15-19) re- corded Calyptraea sp. of Maastrichtian age from south- western France, but the preservation of the specimens appears to be too poor to allow positive generic identi- fication. AFRICA: As figured, ‘Calyptraea’ bouéi (Pervin- quiére, 1912: 10-11, pl. 1, figs. 7-11) from the Maas- trichtian of Tunisia externally resembles some forms as- signed to ‘C. aperta. The shape of the shelf of ‘C. bouéi shown in figure 9 is difficult to determine, but appears to have a straight edge and probably resembles that of ‘C. aperta. Galerus libyca Quaas, 1902, was described from upper Maastrichtian/ possibly D anian strata (Exogyra overwegi beds) from the Ammonite Hills, Great Sand Sea, Egypt (Quaas, 1902: 238, pl. 25, figs. 26-29). It has been re- ported also from the Congo (Dartevelle and ae, 1956: 29-30, pl. 1, figs. 9-10), and from Libya, Egypt, Congo, and Madagascar (Bandel and Riedel, 1994: 339- 340, pl. 7, figs. 2-3). Bandel and Riedel reported Ou: 1as’ original specimens lost in World War II and figure od the Page 136 THE NAUTILUS, Vol. 122, No. 3 exterior of a subsequently collected specimen of ‘Calyp- traea’ libyca which resembles ‘C. aperta. They did not figure the shelf, but described it too briefly as “a flat spiral shelf like that of modern Calyptraca.” A Crepidula chain was reported by Brébion (1956) in describing Crepidula congolensis Brébion, 1956, from the upper Campanian of the Congo, Africa. This African species resembles a Lysis more than it does a modern Crepidula, in that C. congolensis has coiling similar to Lysis and a depressed inner lip that barely wraps onto the labral side of the Ga Lysis? congole nsis (Bré- bion, 1956: 89, fig. 1; pl. 1, fig. 7a, 7b) is most similar to L. jalamaca in shape and sculpture but appears to have much finer ribbing than L. jalamaca. ‘Calyptraea’ primogenita Kiel and Bandel (2003: 460, fig. 4.14-4.16) and Lysis capensis (Rennie, 1930) illus- trated by Kiel and Bandel (2003: 460, fig. 6.1-6.2) are from the upper Santonian/lower Campanian, Umzamba Formation. ‘Calyptraea’ primogenita was described trom a single worn and broken specimen. Its ribbing (except on last quarter-whorl) is protractive as in Trochita, and it has a thick shell as does Trochita. Its whorl shape is more trochiform than in Trochita or Calyptraea, it consists of more whorls than a Trochita or a Calyptraca, the last whorl lacks the notable enlargement of a Trochita or a Calyptraea, and unlike Trochita or Calyptraea, it has a small open umbilicus and “the columellar lip bears a strong plate” (Kiel and Bandel, 2003). Lysis capensis is very similar in shape and sculpture to L. duplicosta, but its inner lip seems narrower and more similar to that of L. mickeyi. Lysis duplicosta is generally lower spired and has a broader inner lip/columell: 1. Rennie (1945: 50, 116, pl. 3, fig. 10) figured a Calyp- traea sp. from the Upper Cretaceous Senonian of An- gola, Africa. though the shape and angle of suture in the figure re- semble Calyptraea, the base and aperture are not illustrated, thereby making generic assignment indeter- minate. NORTH AMERICA (exclusive of the Pacific Slope): Crucibulum? sp. of Sohl (1960: pl. LO, fig. 2 mature, incomplete specimen from the upper part of the Ripley Formation (Maastrichtian) in Mississippi. Sohl in- dicated that its incomplete shell suggested a close rela- tionship to Crucibulum, and that it definitely appeared to belong in the Calyptroidea. The specimen is too incom- plete to determine its genus, its similarity to Crucibulum could be a result of the way it is broken, but its spire does suggest Calyptraeoidea. Soh ( (1960) classed Thylac us cretaceous Conrad, 1860 in Capulidae, but Dockery (1993) moved it to Calyp- traeidae and Bandel and Riedel (1994) included Con- rad’s species in Hipponicidae. A more precise age is unknown and, al- 1) is an im- It apparently lived at- tached to the columella within the aperture of empty gastropod shells. It differs from Lysis and Garzasia in the way Thylacus muscles were attached. At the end of ju- venile coiling and beginning of expansion of the last whorl, Thylacus deposited left and right prongs instead of modifying the inner lip/columella into a shelf as in Lysis. Sohl (1960: pl. 10, fig. 4) and (Dockery (1993: pl.18, figs. 1 and 4) provided good illustrations of the early bce which does seem more hipponicid than calyptraeid. When developed to maturity a horse- shoe-shaped muscle scar resulted. Dockery’ s (1993) specimens were from the Coffee Sand of Campanian age and Sohl’s (1960) from the Ripley Formation of Maas- trichtian age. SOUTH AMERICA: ‘Calyptraea’ aperta (Solander in Brander, 1766), a European Eocene species (see Figures 65-67 for a representative late Eocene specimen from the Paris Basin, France), was identified by Olsson (1944: 245-249, pl. 9, figs. LO-13) from northern Peru. Olsson’s placement of f the Tortuga fossil beds below his Radiolite sandstones with Baculites suggests a Maastrichtian age. As he provided only exterior views and no description of the shelf, this identification needs further verification. Specimens identified as ‘Calyptraea’ aperta from Europe and the American Gulf Coast range widely as to whorl height and sculpture which varies from smooth to spi- nose. The Peruvian specimens increase in diameter at a slower rate and they have more strongly impressed su- tures than Eocene specimens, suggesting that the Peru- vian specimens are probably not °C. aperta. Calyptraea laevis (Philippi, 1887) from Maastrichtian of central Chile was described as Trochita laevis Philippi (1887: 92, pl. 11, fig. 3: referred to Galeropsis by Wilck- ens (1904: 195-196, pl. 17, figs. 9a, b) because of its high spire, and to Calyptraeca (Trochita) by Bandel and Stin- nesbeck (2000: 763-764, pl. 1, fig. C). Bandel and Stin- nesbeck provided a view of the exterior only, and de- scribed the shelf as “a flat concave shell like that of mod- erm Calyptraca.” Although “flat concave shell” might partially describe a Troc hita shelf, it is not an acurate description of the shelf in Cann ytraea chinensis (Figure 6S). Wilckens (1907: 13, pL 3, fig. 6) also reported Ca- lyptraca aff. laevis Philippi of Maastrichtian age from southern Patagonia, but the preservation of the specimen appears to be too poor to allow positive generic identi- fication. Calyptraea pileolus COrbigny, 141 was indicated by Hoagland (1977: 354) to have been recorded from Lower Cretaceous strata of Argentina by von thering (1907), but the species was listed By Fe ruclio ( 1937: 187) foi the Patagoniano (Tertiary) ) of Lago Argentino, Argentina. JAPAN: As noted earlier, Lysis izumiensis Kase, 1990 has been described from the earliest Maastrichtian of Japan. It resembles the group of Lysis duplicosta and appears from the illustrations (Kase, 1990: figs. 2.11- 2.12) to be most similar to L. duplicosta in strength of sculpture and development of innerlip/columellar expan- sion. Kase (1990) also recorded a trichotropid similar to Ariadnaria obstricta from the Izumi Group in Japan. NEW ZEALAND: The early Paleocene (Danian) Spirogalaerus Finlay and Marwick. 1937, from New Zea- land is Lysis-like, especially as to growth line, and this similarity requires further study to de ‘termine whether it L. R. Saul and R. L. Squires, 2008 Page 137 results from common ancestry, conv ergent evolution, or parallel evolution. Finlay and Marwick (1937) assigned Spirogalerus to family C Calyptraeidae and based their ge- nus on Spirogalerus lamellaria Finlay and Marwick, 1937, which resembles L. suciensis except that L. sucien- sis lacks the “pseudoumbilicus” described for Spirogal- erus. Finlay and Marwick (1937) and Boshier (1960) opined that Spirogalerus lamellaria could represent the evolutionary link between crepidulids and calyptraeiform Sigapatella Lesson, 1830. Classification of Spirogalerus has been inconsistent. Wenz (1940) made it a subgenus of Calyptraea Lamarck, 1799. Beu and Maxwell (1 1990) made Spirogalerus a sub- genus of the ied Sigapatella Lesson, 1830, but Stilwell and Zinsmeister (1992) separated Sigapatella from Spirogalerus because the latter has a strongly ex- erted spire. Collin (2003a) noted that Sigapatella has a shell and anatomy ate to the eee genera Ca- lyptraea, Trochita Schumacher, 1817, and Zegalerus Finlay, 1926. Collin (2003b) considered Sigapatella to be monophyletic but ‘id not mention Spirogalerus. Mar- shall (2003) recognized Sigapatella (=Zegalerus) as a dis- tinct genus based on its shelf edge being broadly and evenly concave instead of sigmoidal. This is a very dif- ferent shelf than that of Calyptraea chinensis (Figure 68). The differences between Spirogalerus and Sigapa- tella are similar to those found between Lysis and Gar- zasia, and the New Zealand forms also record an evolu- tionary pattern of enlargement of the last whorl and the aperture, with broadening of the columella/inner lip area into a shelf within a limpetifor m shell. Thus, these two genera were probably not links from actual crepidulids, but represent links from lysines to calyptraeids. ACKNOWLEDGMENTS We mention again the extraordinary assistance of the New Zealanders in documenting a Tertiary age for a specimen mistakenly recorded as Cretaceous. Malcolm G. Laird, University of Canterbury, Christchurch, New Zealand, and Jeffrey D. Stikhvell, Monash University, A. Grebneff, Otago University, Dunedin, New Zealand, and A. G. Beu, GNS Science, New Zealand, provided help in researching the age of the New Zealand Cretaceous units. 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Biological Bulletin 122: 160-181. APPENDIX 1 LOCALITIES CITED Localities are LACMIP, unless otherwise noted. All quadrangle maps are U. S. Geological Survey maps. Bracketed numbers are areas shown on Figure 1. [4] CASG 61794. [= CAS 1346-A]. In conglomeratic sandstone 1.6 km (1 mi.) above mouth of Huling Creek, North Fork Cottonwood Creek, Ono Quadrangle (15 minute, 1952), Shasta Co., California, Budden Canyon Formation, Bald Hills Member. Late Albian. 7333. Le Fayel, Paris Basin, France. Late Eocene (Bartonian Stage). [18] 10095. [=CIT 83]. Fine sandstone just above shale, sectionline fence gate on old road 0.4 km (1/4 mi.) W of Schulz Ranch, 122 m (400 ft.) S. of northeast corner of section 19, T. 5 S, R. 7 W, El Toro Quadrangle (7.5 minute, 1968), south side of Williams Canyon, Santa Ana Mountains, Orange Co., California. Coll.: B. N. Moore, 1 January, 1926. Ladd Formation, uppermost Holz Shale Member. Early Cam- panian. [16] 10711. [=CIT 1158]. Approximately 1.84 km (1.5 mi.) due west of Los Angeles-Ventura Co. line on the boundary (extended) between T. 1 N and T. 2 N. north bank of Bell C Canyon, southeast slope of Simi Hills, Calabasas Quadrangle (7.5 minute, 1952), Ventura Co., Califor- nia. Coll: W. P. Popenoe, 18 July, 1935. Chatsworth Formation. Middle Campanian. (3] 10757. [=CIT 1593]. Massive sandstones in bed of North Fork Bear Creek, approximately 777 m (2550 ft.) south and 533 m (1750 ft) east of northeast corner of section 5, T. 31 N, R. 1 E, Whitmore Quadrangle (15 minute, 1956), Shasta County, California. Coll.: W. P. Popenoe and W. M. Tovell, 12 Sept., 1941. Redding Formation, Bear Creek Sand- stone. Coniacian. [5] 10846. [=CIT 1014]. Concretions in sandstone, right bank of Chico eae about 1.6 km (1 mi.) upstream from the little bridge across creek below Mickey house and about 4.16 km (2.5 mi.) N6°W of 14-mile house on Humboldt Road, NW 1/4, SE 1/4 of section 1, T. 23 N, R. 2 E, Paradise Quadrangle (15 minute, 1953), Butte Co., California Coll: W. P. Popenoe and D. W. Scharf, 16 August, 1931. Chico Formation, top of Ponderosa Way Member. Late Coniacian or early Santonian [23] 11944. Approximately 10 km N of Punta Abreojos (SW of San Ignacio). in first ridges N of arroyo that crosses Punta Abreojos road (dirt) just S of Campo Rene tumoff, approx. 2 km NW of road. Approx. 2-3 km up ravine, hill .5 km to east is pachydiscid loc., Viscaiano Penin- sula, Baja California Sur, Mexico, Coll.: R. Demetrion, 1987, 1989. Valle Formation. Middle? Campanian, with Hoplitoplacenticeras? {14] 14310. About 450m north and 70m west of lookout at summit of Warm Springs Mountain at elev. of 1052 m (3450 ft.), at base of Kirby’s (1991, M. A. Thesis) measured section no. 2, Warm Springs Mountain Quadrangle (1958), Los Angeles County, California. Coll.: M. X. Kirby. Basal San Francisquito Formation. Late Maastrichtian. 22340. Gritty conglomeratic sandstone lenses in fine-grained sandstone and shales, east side Grapevine Canyon about 0.8 km (0.5 mi.) south of its north end, about 0.4 km (0.25 mi.) east of, and 30.48 m (100 ft.) above the abandoned highway roadbed on east side of canyon, at about 648 m (2100 ft.) contour, about 91 m (300 ft.) northwest of Tejon/ granite fault contact. Locality is 3048 m (10,000 ft.) N24°W of 3174 ft. BenchMark at old Fort Tejon, Tejon Quadrangle, Kern County, Cali- fornia. Coll: W. P. Popenoe, 9 December, 1946. Tejon Formation. Middle Eocene. 22386. Prominent shell bed at crest of ridge on east side of Live Oak Canyon, about 0.4 m (0.25 mi.) south of its mouth, T 10 N, R 19 W, Pastoria Creek Quadrangle, 7.5, 1958, photorevised 1974, Kern County, California. Coll; W. P. Popenoe, 13 March, 1947. Tejon For- mation, Metralla Sandstone Member. Middle to upper Eocene (“Tejon Stage”). [10] 22588. About 2/3 of the way to the top of a gully on southwest slope of a northwest-trending hill on south side of Garzas Creek, where the creek enters the San Joaquin Plain, approximately 610 m south and 183 m west of northeast comer of section 19, T 8 S, R 8 E, Howard Ranch Quadrangle, 7.5', 1953, photorevised 1971, west side of San Joaquin Valley, Stanislaus Co., California. Coll: W. P. Popenoe and T. Susuki, April, 1950. Moreno Formation, “Garzas Sand” member. Middle Maastrichtian. [4] 23464. [PR1] Up small creek from Sulivan Ranch Rd. crossing, and 1.28 km (0.8 mi.) north of ranch, near Gas Point Rd., 701 m (2300’) N 75°E from mouth of Huling Creek, 579 m (1900°) S, 488 , (1600°) E of NW comer of section 16, T. 30 N, R. 6 W, Ono Quadrangle (15 minute, 1952), Shasta Co., California. Coll: P. U. Rodda and M. A. Murphy, May 1955. Budden Canyon Formation, Bald Hills Member, unit iv in Matsumoto, (1960). Middle Cenomanian, probably Turrilites costatus Zone. [5] 23617. Fossil in hard, blue-gray concretion in gray-weathering buff sandstone approximately 15.2 m (50 ft.) below highest conglomerate, approx. 0.8 km (0.5 mi.) upstream from Mickey house on west side of Chico Creek, 1.52 m (5 ft.) above stream, 716.28m (2350 ft.) north, 609.6 m (2000 ft.) west of southeast comer of section 1, T. 23 N, R. 2 E, Paradise Quadrangle (15 minute, 1953), Butte Co., California. Coll.: R. B. Saul, 14 August, 1955. Chico Formation, top of Ponderosa Way Member. Late Coniacian or early Santonian. 5] 23639. In concretions in massive, greenish-gray sandstone, east bank of Chico Creek, west of meadow eh large flat- topped, lava block at north edge near road, 373.38 m (1225 ft.) ) south and aoe 6 m (960 ft.) west of northeast corner of section 23, T. 23 N, R. 2 E, Paradise Quadrangle (15 minute, 1953), Butte County, cere California, Col- lectors: L. R. Saul and R. B. Saul, 20 August, 1952. Chico Formation, owermost part of Ten Mile Member. Early C ampanian. 12] 24122. Fine- to coarse grained buff sandstone; 76.2 m (250 ft.) north of jeep trail in Jalama Canyon; elevation 190 m (625 ft.), 6.58 km (4.11 mi.) east and 1.1 km (0.69 mi.) south of Jalama Ranch Headquar- ters: 0.93 km (0.58 mi.) west and 0.66 km (0.41 mi.) north of southeast corner of topo, Lompoc Hills Quadrangle (7.5 minute, 1959), Santa Barbara Co., California. Coll.: D. Dailey, August, 1959. Jalama Forma- tion. Late Campanian—early Maastrichtian [12] 24128. Dark gray conglomerate in first small canyon east of Ra- majal Canyon, elevation 167.6 m (550 ft.), 0.54 km (0.34 mi) south, 3.25 km (2.03 mi.) east of Jalama Ranch Headquarters, 1.22 km (0.76 mi.) north 4.27 km (2.67 mi.) west of southeast corner of Lompoc Hills Quadrangle (7.5 minute, 1959), Santa Barbara Co., California. Coll.: D Page 142 Dailey, August 1958. Jalama Formation, Late Campanian—early Maas- trichtian. {12] 24237. Medium-grained, buff, arkosic sandstone, 396.2 m (1300 ft.) north of Jalama Creek, elevation 160 m (525 ft.), 0.48 km (0.30 mi.) south, 3 km (1.88 mi.) east of the Jalama Ranch Headquarters, 1.28 km (0.80 mi.) north, 4.59 kin (2.87 mi.) west of southeast corner of Lompoc Hills Quadrangle (7.5 minute, 1959), Santa Barbara Co., California. Coll.: W. P. Popenoe, September 1938, Jalama Formation. Late Cam- panian-early Maastrichtian. [6] 24340, Penz vicinity, conglomerate beds cropping out just below a drainage canal, southeast side of new Oroville Hwy, about 1.2 km (0.75 mi.) northeast of intersection with Pentz- Magalia- Oroville road, 426.7 m (1400’) S., 182.9 m (600) W of the northeast corner of section 36, T. 21 N, R.3 E, Cherokee quad. (7.5 minute, 1949), Butte Co., California. Coll.: W. P. Popenoe, 1960. Chico Formation, Musty Buck Member. Early Campanian. [8] 24349. [=USGS M8601 and USGS M8745]. In place? large angular block of sandstone surrounded by sand at shoreline in covelet on north side of elongate seaward-pointing rock; approx. N30°W of Pigeon Point lighthouse, just south of Bolsa Point, Pigeon Point Quadrangle (7.5 minute, 1952), San Mateo County, Califomiia. Coll: and R. B. Saul, October 11, 1960. Pigeon Point Fm. Middle C mes {16] 26020. [=CIT 1158]. Hard sandstone slabs in fine-grained sand- stone, cropping out on high bare cliff, north bank of Bell Canyon, just east of mouth of large gully, and 152.4 m (500’) S, 2743.3 (9000") west of northeast corner of section 4,T. 1 N, R. 17 W, Calabasas Quadrangle (7.5 minute, 1952), Simi Hills, Ventura Co., California. Coll.: W. P. Popenoe, 11 Feb., 1972. Chatsworth Formation. Middle Campanian {10] 26353. Approximately 1.2 km south of Garzas Creek, 671 m south and 114 m east of northwest corner of section 20, T 8 S, R 8 E, Howard Ranch Quadrangle, 7.5°, 1953, photorevised 1971, Stanislaus County, California. Coll; R. B. Stewart and W. P. Popenoe, 1944. Moreno Formation, “Garzas Sand” member. Late early to early late Maastrich- tian, {11] 26370. Reworked fossiliferous Turonian blocks in upper Campa- nian conglomerate lens in shale, northeast side of Cooper Canyon, approx. 411.5 m (1350°) n, 670.6 m (2200°) W of southeast corner of section 2, T, 21 S, R. 14 E, Alcalde Hills Quadrangle (7.5 minute, 1969), \, Alcalde Hills, Fresno Co., California. Coll.: J. Alderson, 18 June, THE NAUTILUS, Vol. 122, No. 3 1977. Panoche Formation, “Alcalde Shale” Member. (with juvenile Subprionocyclus sp.). Late Turonian [18] 26951. Small sandstone lens approx. 6.1 m (20 ft.) above road in roadcut on north side of Silverado Truck Trail, 274.3 m (900 ft.) south of northeast comer of section 18, T. 5 S, R. 7 W, El Toro Quadrangle (7.5 minute, 1949), Orange Co., California. Coll; A. A. Almgren, 4 Dec., 1981. Ladd Formation, uppermost Upper Holz Shale Member. Late early Campanian, [17] 26967. Small exposure of coarse-grained, poorly sorted sandstone at bottom of northwest-flowing tributary to main fork of Garapito Creek, 449.6 m (1475 ft.) and 2835 m (9380 ft.) east of northwest corner of section 5, T. 1S, R. 16 W, Topanga Quadrangle (7.5 minute, 1952, photorevised, 1981), Santa Monica Mountains, Los Angles County, California. Coll.: J. M. Alderson, 31 Dec., 1981. Tuna Canyon Formation. Coniacian., [3] 28717. South Cow Creek Valley, sandstone associated with con- glomerate, lower South Cow Creek Valley, about 152.4 m (500 ft.) downstream from old bridge site across creek, and about 1.6 km (1 mi.) due west of buildings on Hunt Ranch, NE 1/4 of section 17, T. 31N, R. 2W, Millville Quadrangle, $ Shasta Co., California. Coll.; W. P. Popenoe, 27 Oct., 1971. Redding Formation, Oak Run Conglomerate Member of Haggart, 1956. Late Santonian. [19] SDNHM 3403. Taylor Made Golf Facility at Salk Drive and Col- lege Blvd., elevation 45.1 m (148 ft.) along College Blvd., lat. 33°8°25' N, long. 117°1656' W, [in general = SDNHM 3402, 3404, 4071, 4073], Carlsbad Research Center, Site 29, San Luis Rey Quadrangle 7.5° (1968), San Diego Co., Califormia. Coll: B. O. Riney, 26 April, 1987, Point Loma Formation. re Campanian/?early Maastrichtian. {19] SDONHM 3405. Carlsbad Research Center, 1.6 km north and 2 km west of southeast corner of San Luis Rey Quadrangle 7.5° (1 968), indicated area along west side of College Blvd, starting about 0.32 km from intersection with El Camino Real extends southward for 0.15 km, San Diego Co., California. Coll: B. O. Riney, 1987. Point Loma For- mation, ?Late Campanian/early Maastrichtian. {18] UCMP 2167. 3.2 km (2 mi.) N 10°W of BM 1271, Corona Quad- rangle (1902), at a gate about 0.5 km (0.5 mi) below Modjeska Springs in Williams Canyon, Santa Ana Mountains, Orange Co., California. Ladd Formation, uppermost Holz Shale. Middle? Campanian. THE NAUTILUS 122(3):148-150, 2008 Page 143 Feeding behavior, phylogeny, and toxinology of Conus furvus Reeve, 1843 (Gastropoda: Neogastropoda: Conidae) Samuel S. Espino Institute of Chemistry University of the Philippines Diliman, ‘Quezon City 1101, PHILIPPINES and Department of Chemistry University of the Philippines in the Visayas Miag-ao, Iloilo, PHILIPPINES Alan J. Kohn Frank M. Heralde HI National Institute of Molecular Biology and Biotechnology University of fie Philippines Diliman, Quezon City 1101, PHILIPPINES University of Utah Baldomero M. Olivera! Department of Biology University of Utah Salt Lake City, UT 84112 USA olivera.biology@gmail.com and Biotechnology Department of Biology University of W. ashington Seattle, WA 98185 USA Patrice Showers Corneli Department of Biology Salt Lake City, UT $4112 USA James A. Villanueva Institute of Chemistry University of the Philippines Diliman, Quezon City 1101, PHILIPPINES Gisela P. Concepcion Marine Science Institute University of the Philippines Diliman, Quezon City 1101, PHILIPPINES Ameurfina D. Santos National Institute of Molecular Biology University of the Philippines Diliman, Quezon City, PHILIPPINES ABSTRACT The Indo-Pacific cone snail Conus furvus Reeve, 1843, is shown to be molluscivorous, based on four lines of evidence: (1) Specimens of Conus furvus maintained in aquaria were directly observed to sae devour other gastropods, including other cone snails; (2) Its radular teeth share morphometric characteristics with ce known molluscivorous Conus and dif- fer from those of vermivorous and piscivorous species; (3) Mo- lecular phylogeny places Conus furvus within a clade of mol- luscivorous species; (4) Sequences of two peptide toxins are most similar to conotoxins prey jiously characterized from mol- luscivorous Conus species; one of these closely related peptides was previously shown to block molluscan L-type Ca channels. Similar to molluscivorous species such as Conus omaria and C. victoriae, C. furvus varies widely in shell shape and color pat- term, even within its relatively restricted geographic range. Additional Keywords: Conoidea, snail-hunting cone snail, radu- lar morphology, conotoxins, RT-PCR, phylogeny. INTRODUCTION Cone snails are venomous marine gastropods (Kohn, 1959; Réckel, Korn, and Kohn, 1995), and the major biologically active components of their venoms are the conotoxins (Olivera, 2006), typically small (12-35 amino ' Author for correspondence acid residues), highly constrained peptides. Many cono- toxins dices between closely related molecular isoforms of a particular ion channel family (Terlau and Olivera, 2004); this property makes them valuable tools in neuroscience, and confers their therapeutic potential. The estimated ~100,000 different pharmacologically ac- tive peptides present in living Conus venoms comprise a very substantial neuropharmacological resource. The sys- tematic study of conotoxins is a se -emingly daunting task considering the enormous number of peptides. As the conotoxin sequence database expands, it is becoming clear that certain types are likely to be produced by spe- cies that specialize on particular prey and that belong to particular Conus clades (Olivera, 2002, 2006). Thus, es- tablishing both the prey and the phylogenetic relation- ships of Conus species provides an important guidepost for searching for conotoxins with certain types of physi- ological activities. Traditional cone snail taxonomy is based on shell mor- phology; this has proved difficult ‘because of the absence of resolution and possible convergence of shell charac- teristics. None of the infragenic classificativis schemes primarily based on shell shape and sculpture, shell color patterns, and radular morphology has received wide- spread acceptance (Duda, Kohn, and Palumbi, 2001). Thus, these traditional criteria need to be complemented by other types of data. The specific focus of this study, Conus furvus (“the Page 144 THE NAUTILUS, Vol. 122, No. 3 dark cone”) is unusual in several respects. Although it is found over a relatively restricted area of the tropical Indo-Pacific (almost all specimens have been collected in the Philippines, where it is quite common in some lo- calities), its shell shape and color pattern vary widely (Figure 1). In most localities, the typical form of Conus furvus has a brown and white shell. However, in the Southwestern Philippines, forms occurs, particularly along the island of Palawan and in the Sulu Sea (Figure 1). As new localities are explored, additional variations are being discovered. In the comprehensive treatment of Indo-Pacific Conus spe- cies by Réckel et al. (1995), the number of taxa synony- mized with Conus furvus was probably among the high- est for any species. The authors discussed abang a dozen distinct forms which they regard as geographic variations. Rockel et al. (1995) concluded that Conus furvus * ‘ap- pears: to be characterized by isolation of local popula- tions.” In general, the occurrence of sinistral spe ciumens in any Conus is exceedingly rare; another notable feature of C. furvus is that most sinistral Conus specimens col- lected in the Philippines belong to this species (an ex- ample is shown in Figure 1). In a study using the calmodulin intron sequence as a phy logenetic mar Tae Duda and co-workers (Duda et al., 2001: “Duda and Kohn, 2005; Duda and Palumbi, 2004) grouped C. — with C. litteratus in their phylogenetic tree. They inferred that the two species Ne erged only at the beginning of the Pliocene; since litteratus is a well- gorbliened vermivorous species i feeds on capi- tellid polychaetes (Kohn and Nybakken, 1975), with a fossil record going to the Lower Miocene, this strongly suggested that C. furvus is a vermivorous Conus. Purchasers of Conus furvus in the Guimaras Islands public market, in the Philippines, make a soup by boiling the snails with tomatoes, onions, and lemongrass. Only the portions of C. furvus near the foot are eaten; appar- ently the sections near the hepatopancreas have a bitter taste and ere uny texture. An alternative culinary use of the snails is as an ingredient for a local dish prepared with noodles. Various other snails are ty pically sold with it: C. radiatus, a piscivorous spe cles, is the most common other Conus found for sale with C. furvus at this locality. In this paper we present data on prey capture ena by C. furvus, the morphometric characteristics of i radular teeth, a phylogenetic analysis based on stand: rd mitochondrial sequences, and the first conotoxin se- quences obtained from this species. These data consis- tently support the conclusion that Conus furvus is a mol- luscivorous species. MATERIALS AND METHODS Specimen Collection and Dissection: Specimens of C. furvus for DNA extraction were purchased ina public market in Guimaras Island, Philippines. The snails were dissected to remove the hepatopancreas for total geno- mic DNA extraction and the venom duct for RNA ex- a remarkable diversity of traction. A few specimens were maintained in an aquarium. Specimens for radular tooth analyses were collected in Coron Harbor and Bobok Island, Palawan, Philippines, and preserved in alcohol. The shells are deposited in the Field Museum of Natural History, Chicago, FMNH 300120, 300121, and 300122. The body of specimen No. 300121 is preserved in the Bernice P. Bishop Museum, Honolulu, Hawaii, BPBM 248751. Analysis of Radular Teeth. The radular sacs of four specimens were dissected and 4-6 teeth from each mounted permanently in polyvinyl lactophenol medium on slides. Barb and blade lengths were measured from the tip of a tooth as described | xy Kohn et al. (1999), using an ocular micrometer and Nomarski differential interference contrast optics. Genomic DNA Extraction: The total genomic DNA was extracted using the Xanthogenate DNA Extraction Protocol (Tillet and Neilan, 2000), a method based on the use of the polysaccharide solubilizing chemical xan- thogenate. The method is non-toxic and requires no en- zymatic or mechanical steps to break cells. RNA Extraction: Frozen venom duct in RNAlater (Ambion, Austin, Texas) was thawed in ice prior to RNA extraction, and a sample of the tissue (10-l5mg) was homogenized in 1 mL Trizol reagent (Invitrogen, Carls- bad, California). The total RNA was extracted according to the manutacturer’s recommendation. Amplification of the Mitochondrial 12S and 16S rRNA and mtCOI Genes: For PCR amplification, primer pairs designed to hybridize to a segment of 12S, 16S rRNA genes and mtCOl genes were used. The primer sequences used for the amplification are listed in Table 1. PCR amplifications of the three mitochondrial genes were carried out in a 10 wh reaction mixture con- taining 1X reaction buffer (200mM Tris HCI, 500mM KCI, pH 8.4), 0.7 mM of each dNTPs, 0.7 mM of both primers, 0.4 unit Taq DNA Polymerase, 3.0 mM MgCl, and approximately 50.0 ng of template DNA. PC R am- plification was carried out for 40 cycles. Denaturation was carried out at 94 °C, annealing at 48°-55 °C and extension at 72 °C. The PCR products were visualized on 1% agarose gel containing 0.1 mg EtBr/mL. Multiple or smeared PCR products were further purified using the WIZ Prep DNA purification kit (Promega, Madison, Wisconsin), otherwise, the terminated reaction was di- rectly sequenced, These sequences have been deposited at GenBank. 12S Accession Numbers: — aimmiralis EU682274, auli- cus EU682275, bandanus EU682277, dalli EU682281, episcopatus EU682283, flavus EU794315, furvus [EU682254, geographus EU794316, kintoki EU794317, litteratus EU 784318, magus EUT94319, marmoreus EU682288, monachus EU794320, obscurus KU794321, omaria EU682289, textile EU682296, tulipa EU794322, virgo EU794323:; 16S Accession numbers: ammiralis Espino et al., 2008 Figure 1. Some of the different variations of Conus furvus. The brown colored specimens shown in the bottom row, center, are similar to those used for this research. Note the sinistral specimen in the lower left hand corner. Page 146 THE NAUTILUS, Vol. 122, No. 3 Table 1. Primer pairs used for the amplification of the 12S, 16S rRNA gene and mtCO1 gene. 12S 16S mtCOl Upstream primer 5’ AGAG(C/T)G(A/G) 5’ GTTTACCAA 5’ GGTCAACAA CGGGCGATGTGT 3’ AAACATGGCTTC 3’ ATCATAAAGATATTGG 3’ Downstream primer 5’ TGCCAGCAG 5’ CCGGTCTGA 5’ TAAACTTCAGGG (C/T)CGCGGTTA 3’ ACTCAGATCACGT 3’ TGACCAAAAAATCA 3’ EU682299, aulicus EU794324, bandanus Ps vate dalli EU078935, episcopatus EU078937, flavus EU794326, furvus EU682301, geographus RUF 794327, kintoki EU794328, litteratus EU794329, magus EU078939, marmoreus EU794330, monachus EU078938, obscurus EU794331, omaria EU794332, tex- tile EU078936, tulipa EU794333, virgo EUT94334. RT-PCR of M and O-superfamily Conotoxin Genes: Single-stranded cDNA was synthesized from the total RNA extracted from the snail’s venom duct using oligodT primers and Superscript IH (Invitrogen, Carlsbad, California) and following the manufacturer's protocol. Double-stranded cDNA was synthesized using primers targeted to the conserved regions of the M- and O- superfamily genes. The product of the synthesis was visualized in a 1% agarose gel containing EtBr (0.1 mg/ mL). The terminated reaction cont: aining the PCR prod- uct of the correct size (~0.4Kb) was used for the cloning reaction. The cloning reaction followed the manufactur- ers protocol for the TOPO-TA cloning kit (Invitrogen Life Technologies, Carlsbad, California). The clones were screened for inserts of the correct size and were cultured on a selective medium and incubated overnight with shaking (200 rpm) at 37 °C. The plasmids from these clones were extracted and sequenced. Phylogenetic Analysis: 12S and 16S rRNA sequences were aligned using ClustalX, v1.8 ( (Jeanmougin, Thomp- son, Gouy, Higgins, and Gibson, 1998). The alignments were refined by eye using MacClade 4.08 ( Maddison and Maddison, 2001). Trees were optimized using the indi- vidual rRNA alignments and the sonicated align- ments (prese snted herein). To account for the complexity of sequence evolution, final analyses were restricted to model-based maximum likelihood (PAUP4b10, (Swof- ford, 2002)) and Bayesian inference (Huelsenbeck and Aonquist, 2001). Each method used maximum likelihood parameters describing sequence evolution that were op- timized with a GTR+I+G model that includes six pos- sible substitution types (GTR) and allows some sites to be invariant (1), allows across-site rate heterogeneity (G) and allows unequal base frequencies. The Bz esian ani lysis ran for 100,000 generations (sampled every 100 generations) with the first 25,000 generations discarde a. as burn-in trees. Two MCMCMC runs (metropolis-coupled Monte-Carlo markoy-chain), using four chains each, were used to thoroughly explore tree space Convergence of the likelihoods was judge od adequate by monitoring the MrBayes output describing (ASED) in the average standard error of the difference split frequencies between the two runs and by comparing plots of the tree log-likelihood trees from generation 50,000 to LOO. By the! last generation, the ASED was less than 0.004: the plot of likelihoods versus generation had stabilized. Furthermore, the PSRF (Potential scale re- duction factor) reached 1.00 for the total tree length and for each model parameter. RESULTS Feeding Observations: Individuals of Conus furvus were frequently observed to prey on other cone snails in the aquarium. Most observations were made with C. ra- diatus as the prey. Upon introduction of prey into an aquarium tank with C. furcvus, immediate behavioral re- sponses were observed both from predator and prey. The prey immediately crawled away from C. furvus and at- tempted to bury itself in the sand. At almost the same time, C. furvus pointed its siphon toward the retreating snail and crawled in its direction. While still relativ ely far away from the prey, C. furvus extended its proboscis, and inserted it into the body of its prey with one jerking motion. Immediately the prey was observed to stop mov- ing. Conus furvus then maneuvered the immobilized prey so that its aperture was turned upwards, away from the sand, after which C. furcus commenced feeding on the upturned C. radiatus. Analysis of Radular Teeth: Radular tooth length ranged from 6.4 to 8.9% of shell length (mean = 7.5%) in the four specimens examined. Each tooth is armed with a barb that extends for 5% of the tooth length from the tip (range 4.6-5.4%), and a blade that extends for 9% of the length from the tip (range $.9-9.3%). This morpho- metric data clearly identifies the teeth as those of a mol- luscivorous species, as noted ie Nybakken (1990; see also Kohn, et al., 1999: Nishi and Kohn, 1999). However, these workers did not present details of C. furvus tooth morphometry, which differs markedly from previously examined molluscivorous Conus species. Nishi and Kohn (1999) divided the eleven species they studied into three groups based on both discrete and quantitative charac- ters. The former, i.e. presence of one barb, one blade, and continuous serration clearly place C. furvus in Group A, with C. araneosus, C. bandanus, and C. marmoreus. Groups B and C have teeth with two barbs and no blade (Table 2). However, C. furvus differs from the Group A species in its quantitative characters. Its teeth are more than twice as long relative to shell length, thus matching the members of Group C rather than Group A (Table 2). The C. furvus teeth also differ from Group A and more Espino et al., 2008 Page 147 Table 2. Comparison of Conus furvus radular teeth with those of other molluscivorous species according to qualitative (discrete) and quantitative (continuous) characters. N, Number of specimens measured. Qualitative characters: B1, First barb; B2, Second barb; BL, Blade; SE, Serration; +, present; —, absent; C, Continuous; D, Discontinuous. $,, Shell length; B,, Blade length; SE, C from Nishi and Kohn (1999). Qualitative characters T,, Tooth length; B1,, Barb length; . Serration length; BA\, Base width; SH\,, Shaft width. All dimensions are in mm. Data on Groups A, B, and Quantitative characters Group Species N BL B2 BL SE T;/Sy, Bly/T,., B2,/T,. BL,/T;, SE,/T, BA\/T,, SH\\/Ty, C furvus 4 + - + G 0.07 0.05 —- 0.09 0.70 0.06 0.04 (0.04—0.05) A C. araneosus > Ss = “:, JC 0.03 0.08 — O13 0.49 O11 0.05 C. bandanus > (0.026—.033) (O.07—0.08) (O.13-0.14) (0.33-0.68) (0.09-0.12) (O0.04—0.5) C. marmoreus 15 BC. ammniralis A, a oe HE 0.05 0.04 0.06 = 0.74 0.05 0.03 C. canonicus 7 (0.03-0.08) — (O.03—0.05) (0.05—0.09) (0.62—0.81) (O0.04—0.08) (0.02—0.05) C. nodulosus 5 C. textile 15 C. victoriae 4 Cc C. episcopatus 11 + + - D 0.07 0.05 0.08 0.65 0.07 0.05 C. omaria 9 (O.07—0.08) — (O0.04-0.03) (O.07—0.09) (0.67—0.69) (0.06—0.08) (0.04—0,05) C. pennaceus 30 closely resemble Groups B and C with respect to all four other quantitative characters that they share (Ta- ble 2) PCR Amplification and Phylogenetic Reconstruc- tion: A phylogenetic tree based on two standard mark- ers, 12S and 16S rRNA was constructed as described under Methods. Conus furvus clustered on the same branch with species previously shown to feed on other mollusks (Figure 2). In contrast, Conus furvus was well- resolved from C. litteratus, the worm-hunting species postulated previously to be C. furvus’s closest relative. The hypothesis that C. furvus is closer to the C.litteratus than to the other mollusk hunting snails is significantly rejected (p<0.0001) using Kishino- Hasegawa (Kishino and Hasegawa, 1989) and Shimpceaea- Hasegawa (Shimo- daira and Hasegawa, 1999) tests comparing the respec- tive log- likelihood scores. The hypothesis that C. furvus and C. litteratus are sister species is significantly rejected by these tests using the 12S and 16S aa alone. Because of the great variation of forms presently assigned to Co- nus furvus, we also obtained a “bar code” sec mare (CO] gene) from the specimens analyzed, which is deposited in GenBank. Mapping Radular Tooth Characters to the Molecu- lar Phylogenetic Tree: Morphometric data on radu- lar teeth were available for seven of the nine molluscivo- rous species on the molecular tree (Figure 2), and the values of the seven quantitative morphometric characters for these species shown on the right side of Table 2 were entered into a discriminant function analysis. A phyloge- netic tree generated from the proximity matrix c ee ulated from the centroids of the first two canonical variates they accounted for 99.6% of the variance) (Figure 2B) was completely congruent with the conotoxin gene-based tree (Figure 2A). O- and M- Superfamily Genes from C. furvus: — Primer pairs designed to hybridize to the conserved regions of the O- | M-supe family conotoxins successt fully ampli- fied conotoxin genes from the C. firvus venom duct. Translation of the amplified sequences resulted in amino acid sequences characteristic of the canonical arrange- ment of conotoxin precursors. Both of the sequences have a hydrophobic N-terminal signal sequence, which is highly conserved among conotoxins belonging to the same superfamily, an intervening pro region, and the mature toxin region at the C-terminus. An O- superfamily conotoxin (Fr7.1) from C. furvus was successfully amplified and sequenced, and a 264- base sequence containing an open reading frame coding for the conotoxin was obtained. A 302- age sequence coded for a second conotoxin from C. firvis (Fr3.1) that belongs to the M-superfamily was also cloned and se- quenced. The O-superfamily conotoxin from C. furvus (Fr7.1) was strikingly similar to an w—conotoxin isolated from Conus textile (Fainzilber et al., 1996), wTxVIT (Figure 3A). Alignment of the amino acid sequences shows §2.1% silty wTxVITA has previously been shown to target L-type Ca channels in mollusks; given the high sequence similarity, Fr7.1 is likely a voltage -cated Ca channel ante wonist. A comparison of the M- -superfamily conotoxins from C. furvus (Fr3.1) with M-supert family conotoxins from C. textile (Tx3.5) and C. marmoreus (Mr3.5) also shows a high degree of similarity (Figure 3.B). Alignment of Fr3.1 with Tx3.5 shows the sequences to be 84.4% identical while Fr3.1 aligned with Mr3.5 shows the sequences to be 67.2% identical. The Page 148 THE NAUTILUS, Vol. 122, No. 3 C. ammiralis A 100 C. dalli C. textile C. aulicus C. episcopatus C. omaria C. bandanus C. marmoreus C. furvus C. flavus C. geographus C. obscurus C. tulipa C. magus C. monachus C. kintoki C. virgo C. litteratus 0.05 substitutions/site B C. ammiralis C. textile C. episcopatus C. omaria C. bandanus 10 units C. marmoreus C. furvus Figure 2. A. Optimal maximum likelihood tree from a Bayesian analysis . 12SrRNA and 16SrRNA sequences estimating the evolutionary relationship among some Conus species. The uppermost cli ide, from Conus ammiralis to C. furvus, comprises mollus- civorous species. Branch labels are Bayesian support values summarizing confidence in each split. B. Ne ighbor-joining UPGMA tree from analysis of radular tooth morphometry, based on euclidean distance matrix of centroids of canonical variates | and 2 determined from discriminant function analysis of 7 radular tooth variables, measured as in Nishi and Kohn (1999). Fr3.1 sequence clearly belongs to the m-3 branch of the M-conopeptide supe fi umily, which is characteristic of molluscivorous Conus species (Corpuz et al. , 2005). The two C. furvus conotoxin sequences are most similar to conotoxins from other molluscivorous species, consistent with the behavioral observations made above and with the phylogenetic tree shown in Figure 2. DISCUSSION In the central Philippines, Conus furvus occurs on rocky intertidal habitats. At low tide, individuals can be found in depressions on the substrate, together with other mol- usks; this is where C. furvus is generally collected by ishermen in Guimaras Island, to be sold with other com- mercial shellfish at the public market. Conus furvus was ormerly abundant along the shores of Guimaras Island, yut an oil spill in the area in August 2006 threatened the marine life furvus became much more scarce at this ocality after the spill, but the population appears to be ecovering. From the results presented above, we conclude that Conus furvus is a molluscivorous species. This conclu- ion is based on direct observation of C. furous attacking gastropod prey, on morphometric characteristics of the radular teeth that are shared with other molluscivorous Conus, by the high degree of sequence similarity be- tween toxin precursors in the venom duct of Conus fur- vus and previously described conotoxins from other mol- luscivorous Conus, and by its phylogenetic proximity to them based on molecular markers. A previous molecular result suggesting that C. furvus might be vermivorous (Duda et al., 2001; Duda and Kohn, 2005) is probably erroneous. An unusual feature of the Conus furvus attack on C. radiatus is that the prey was apparently injected with venom only once; multiple stinging was never observed. All other molluscivorous Conus species previously ob- served attack prey using multiple injections of venom before starting to feed on the immobilized prey (Yoshiba, 1983, 1987: Kohn, 2003). The reason for this behavioral difference is unclear, but the venom composition could well reflect it. A further biochemical characterization of Conus furvus venom is in progress (S. Espino and G. Concepcion, unpublished results). The phylogenetic relationship between C. furvus and other Conus species evaluated for 12S and 16S rRNA gene sequences (Figure 2) is consistent with C. furvies Espino et al., 2008 A Fr7.1 SC TPRGGQCGYYNDCCSHQCNINRNLCE w-Txvil YCTPHGGHCGYHNDCCSHQCNINRNKCE Fr3.1 MSTLGVLLTICLLLFSVTALPLDGDQPVDLAAERMKAEQHPLFDQKRRCCKFPCANSCRHLCCG Tx3.5 MSKLGVLLTICLLLFPLTALPLD GDQPADQAAERMQAEQHPLFDQKRRCCKFPCPDSCRYLCCG Mr3.5 MSKLGVLLTICLLLFPLTALPLD GD QPAD QRAERTQAEKHSLPDPRMGCCPFPCKTSCTTILCCG Mature toxins: Fr3.1 RCCKFPC|ANS CRIH/ILCC * Tx3.5 RCCKFPC|PD/SCRIY|LCC* mr3.5 MGCCPIFPCK Ts Cit TILCC* Figure 3. A. Alignment of the toxin region of Fr7.1 from Conus furcus and ot from Conus textile (GenBank Nuc. Acc. No. DD012770). B. Alignment of Fr3.1 from Conus fur- vus with other M- Gee umily precursors highlighting the con- served amino acids. Tx3.5 and Mr3.5 were from Conus textile and Conus marmoreus, respectively (Corpuz et al., 2005). The predicted mature toxins are also shown; the asterisk denotes an amidated C-terminus. being more closely related to the mollusk hunting Conus species than to C. litteratus (Bayesian clade support = 9S), a vermivorous species. The several clades of mollus- civorous Conus probably have a common ancestor. Using the tree based on molecular data in Figure 2A, the fell supported ¢ or 7 can be referred to, using the subgenera of Marsh (1964) as clade names) as the Conus clade? (with Conus marmoreus as the type), the Cylinder clade, (with Conus textile as the type), and the Darioconus clade (with Conus omaria as the type). Although Conus furvus belongs within the larger mollusk hunting clade, includ- ing these three groups, its position hin the clade re- mains unresolved. Given its lack of a tented pattern, it has an unusual shell pattern for a molluscivore. However, some specimens of smaller, usually tented species such as Conus barbieri and Conus victoriae occasionally have plain brown shell variants, not unlike typical specimens of Conus furvus. A more comprehensive molecular analysis of mollus- civorous Conus, including C. furvus, has recently been carried out; these cults will be presented elsewhere. Additional phylogenetic markers, such as ITS-2 se- quences, were aged for this study. This has led to greater refinement of the phylogenetic tree and supports the basic conclusion that C. furvus is a molluscivore. ACKNOWLEDGMENTS This work was supported in part by GM48677 from the U.S. National Institutes of General Medical Sanaecs (to BMO). The support of the University of the Philippines in the Visayas for a fellowship aw ard to S.E. to pursue graduate studies in Chemistry is gratefully acknowl- edged. 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Annual Review of Ecol- ogy and Syste matics 33: 25-47. Olivera, B. M. 2006. Conus peptides: biodiversity-based dis- covery and exoge nomics. Journal of Biological Chemistry 281(42): 31173-3117 Réckel, D., W. Korn, and A. J. Kohn. 1995. Manual of the Living Conidae. Vol. 1, Indo-Pacific Region. Verlag Christa Hemmen, Wiesbaden, 517 pp. Shimodaira, H. and M. Hasegawa. 1999. Multiple Comparisons Page 150 of Log-Likelihood with Applications of Phylogenetic Inference. Molecular and Biology Evolution 16: 1114— 1116. Swofford, D. L. 2002. PAUP® Phylogenetic analysis using par- simony (* and other methods). Version 4. Sunderland, Massachusetts: Sinauer Associates. Terlau, H. and B. M. Olivera. 2004. Conus venoms: a rich source of novel ion channel-targeted peptides. Physiologi- cal Reviews 84: 41-68. THE NAUTILUS, Vol. 122, No. 3 Tillet, D. and B. Neilan. 2000. Xanthogenate Nucleic Acid Iso- lation from Culture and Environmental Cyanobacteria. Journal of Phycology 36; 251-2558. Yoshiba, S. 1983. [Feeding behavior and diet of Conus banda- nus, especially a comparison with those of Conus textile]. Venus 42: 63. (Abstract) [in Japanese] Yoshiba, S. 1987. [Feeding and tooth shooting frequencies of Conus textile for 5 years in an aquarium.| Venus 46: 52. (Abstract) [in Japanese | THE NAUTILUS 122(3):151-154, 2008 Page 15] Two new species of Mitrella (Gastropoda: Neogastropoda: Columbellidae) from the lower Miocene Chipola Formation of northwestern Florida Richard Duerr Post Office Box 1055 Okeechobee, FL 34973 USA pdiegel@gmail.com ABSTRACT Two new species of Mitrella from the lower Miocene Chipola Formation of are described. Mitrella hayesorum new species and Mitrella phyllisae new species are found primarily in Chipola Formation exposures along Farley Creek in northwest- ern Florida, USA. Placement of the new species in the genus Mitrella is tentative. Additional Keywords: Neogene, ultraviolet light, gastropod, Astyris INTRODUCTION Species assigned to the genus Mitrella are widely distrib- uted in the warm and shallow waters of the Recent seas of the world ( Mitrella is not clear, but some of the earliest represen- tative of the genus from the southeastern portion of North America appeared in the Eocene (MacNeil and Dockery HI, 1954; Palmer, 1937). Maury (1910) de- scribed three Mitrella (as Astyris) from the Chipola For- mation. These were originally deposited in the Cornell University collection at lead. New York, and now de- posited at PRI. With more specimens than Maury had at her disposal, Gardner (1947) described 13 species and two subspecies of Mitrella from the Chipola Formation. Mitrella have been found in all exposed facies of the Chipola Formation on the Chipola River, Tenmile Creek and Farley Creek, in the Chipola River drainage in Cal- houn County, Florida. They are also present in the Chipola Formation exposed in the lower bed at Alum Bluff in Liberty County, Florida. The two new species described herein have been collected only in the Chipola Formation exposures in the Chipola River drainage, pri- marily at Farley Creek. Vokes (1989) stated the C hipola facies along Farley Creek where the two new species are most prevalent is a bivalve-rich miliolid lime-sand with many calcareous algae and coral heads, and assumed it was a shallow back-reef environment. Some specimens were photographed under ultraviolet light (UV) to facilitate visualization of color patterns. In- Gardner, 1947, 1948). The exact origin of stitutional abbreviations are: UF: Florida Museum of Natural History (FLMNH for locality records), Univer- sity of Florida, Gainesville; PRI: Paleontological Re- search Institution, Ithaca; ANSP: The Academy of Natu- ral Sciences, Philadelphia, AMNH: American Museum of Natural History, New York. SYSTEMATICS Superfamily Buccinoidea Rafinesque, 1815 Family Columbellidae Swainson, 1940 Genus Mitrella Risso, 1826 Diagnosis: Small to very small, smooth, fusiform shells. Sculpture, if present, of incised spiral lines. Spiral sculpture is generally restricted to the anterior end near the base: axial sculpture is, with few exceptions, entirely absent. The outer lip is dentate in adult specimens (Di- agnosis according to Campbell [1993] and Keen [1971]). PMitrella hayesorum new species (Figures 1-3, 10) Description: Shell fusiform, narrow. Height of holo- type 19.Smm. Protoconch mammillated, with about two smooth, rounded whorls, the second expanded, with no perceptible protoconch/teleoconch transition. Teleo- conch with ten slightly convex, shiny whorls with micro- scopic irregular spiral grooves, axial sculpture lacking. Spire elev ated with somewhat concave sides. Suture dis- tinct. Aperture elongated, less than half the length of the entire shell. Outer lip sinuous, varicose externally, mar- gin thin and sharp. Basal lip extending slightly beyond pillar. Posterior canal produce sd and thickened externally. Denticles present on inner surface of outer lip, ten to 12 in number, weak on anterior third, strongest on central third, frequently absent on posterior third. Parietal wall with two axial ridges, outer ridge weak with small raised beads reflecting underlying sculpture on pillar. Inner ridge prominent with median notch, ac lapical half of ridge strong, abapical half tapering and completely dis- appearing abapically. Base of pillar with ee nine nnn. —— Ssh —R|™?™o>s4)y wa THE NAUTILUS, Vol. 122, No. 3 R. Duerr, 2008 ol (ey) Page 15: oblique grooves with a rounded summit between grooves. Dark band about one third width of penultimate whorl appears below suture on last three whorls of ho- lotype. Band has irregular narrow, oblique, light colored lines, some forming u- or v-ligures. Type Material: Holotype: UF 119655, a ight 19.Smm, width 5.6mm: Par: itypes: ANSP-IP $1324, one specimen; AMNH-FT 433 312, one specimen; PRI ye one specimen; Diegel-Duerr collection, one specimen. All from type locality. Type Locality: FLMNH locality Farley Creek 07 (CA022), Farley Creek east of SR 275, Calhoun County, Florida (to protect privacy rights of landowners, spe cific locality information is avail only to qualified re- searchers upon written request to the author or the In- vertebrate Paleontology Division of the FLMNH), Chipola Formation. Distribution: Chipola Formation along Tenmile and Farley Creeks and the Chipola River, Calhoun County, Florida. Etymology: Named to honor the forestry-oriented Hayes family who have graciously granted the author, and others, permission to collect on their property. Remarks: The species currently placed in Mitrella comprise a complex group and may have been differe tly assigned to the genera Alia, Astyris, Nitidella by diff erent authors (Keen, 1971). The taxonomic position of M. hayesorum is questionable and its current assignment to Mitrella is tentative. Mitrella hayesorum has unique characters, such as the bulbous second whorl of the pro- toconch, extended slightly concave spire and wide sipho- nal canal extending below the pillar, and cannot be con- fused with any other known Mitrella. Al though no speci- mens of Mitrella dalli ( Maury, 1910), the species in the Chipola Formation closest in form to M. hayesorum, were available for study, Maury’s figure of M. dalli (1910: pl. 6, fig. 2) indicates a smaller she TL ( 12 mm), a shorter, stouter spire, and a narrower anterior canal than M. hayesorum. The nearest European fossil congener of Mi- trella hayesorum is Mitrella (Mac rurella) nassoides (Grateloup, 1827) (Figures 4-6) from the early Pliocene of Italy, which is larger, wider at the midbody, and has a narrow anterior canal. The maximum height of all speci- mens of Mitrella hayesorum exé mined is 19.93 mm, minimum height is 18.17 mm. Exposure of M. haye- sorum to ultraviolet light (Figure 10) reveals a fuores- cent pattern of axial flammules in addition to the narrow band below the suture on the anterior whorls of the new species. Seven specimens of the most common Mitrella in the Chipola Formation, Mitrella ischna Gardner, 1947, a more robust species than M. hayesorum, were examined under UV light and revealed fine filamentous lines covering all tele oconch whorls. Three specimens of Mitrella asema Gardner, 1947, a shorter species than M. hayesorum, exhibits, under UV light, narrow axial bars running from suture to suture, offset in alignment from previous whorls, with four per whorl on most teleoconch whorls. ?Mitrella phyllisae new species (Figures 7-9, 11) Desempien: Shell small, fusiform. Height 7 mm, width 2.2 mm. Protoconch with two smooth wlicels sec- ond seh enlarged. Teleoconch with five smooth, con- vex whorls, without axial sculpture. Spire sides. slightly concave. Suture impressed. Aperture less than half the length of entire shell. Outer lip with slight varix, margin in and sharp, usually dentate within. Thin parietal wi ach saa About seven impressed oblique grooves cross base of pillar, separated by narrow bands a rounded summits. Faint round spots, eight on last whorl of holo- type, appear on last three whines: Spots about one- quarter the height of penultimate wher! in diameter rest just above median line of the whorls. Type Material: Holotype, UF 119656, height 7.0 mm, width 2.2 mm; Paratypes, ANSP-IP $1325, one speci- men; AMNH-FT 43313, one specimen; PRI 8353, one specimen; Diegel-Duerr collection, one specimen. All from type locality. Type a FLMNHEL locality Farley Creek 07 (CA022), Farley Creek east of SR 275, Calhoun County, Florida, (to protect privacy rights of landowners, specific locality information is available only to qualified re- searchers upon written re quest to the author or the In- vertebrate Paleontology Division of the FLMNH), Chipola Formation. Distribution: Chipola Formation along Tenmile and Farley Creeks and the Chipola River. Calhoun County, Florida. Etymology: Named for Phyllis Diegel, the author's companion and a knowledge able conchologist and pale- ontologist. Remarks: — As with Mitrella hayesorum, the assignment of M. phyllisae to Mitrella is tentative. A cursory inspec- tion would indicate Mitrella phyllisae to be a dwarf M. hayesorum. Closer examination reveals that M. phyllisae Figures 1-11.—WMitrella species. 1-3. Apertural, lateral, and abapertural views of the holotype of Mitrella hayesorum new species, U F 119655, height 19.8 mm, width 5.6 mm, Burdigalian Miocene. a 6. Apertural, lateral, and ab: apertural views of Mitrella nassoides Grateloup, 1827 ) UF 119657, height 25.5 mm, width 8.6 mm, Zanclean Pliocene, from Liguria County, Ceriale, Italy, for comparison with M. hayesorum. 7-9. Apertural, lateral, and abapertural views of the holotype of Mitrella pluyllisae new species, UF 119656, height 7.0 mm, width 2.2mm, Burdigalian Miocene. 10, 11. Mitrella exposed to UV light. 10. fol e of Mitrella hayesorum (same specimen as Figure 3) showing UV exposed pattern. 11. Holotype of Mitrella phyllisae (same specimen as Figure 9) showing UV exposed pattern. Page 154 § differs from the much larger M. hayesorum by the shorter spire, the slightly more impressed suture, and fewer oblique incised lines on the base of the shell. A series of dots about 1 mm in diameter are visible on M. phyllisae encircling all teleoconch whorls, one-third the length of the whorl below the suture, eight on the last whorl. Exposure to UV light reveals (Figure 11), in ad- dition to the larger dots, a field of minute dots covering the entire teleoconch, somewhat reminiscent of the pat- tern on the Recent Mitrella ocellata (Gmelin, 1791) from the western Atlantic. Also, the anterior portion of the outer lip of M. phyllisae is less developed than that of M. hayesorum, which is wider and extends beyond the pillar. The height of all specimens of M. phyllisae examined varies lees than 1.0 mm, from a maximum of 7.4 mm to a minimum of 6.8 mm. ACKNOWLEDGMENTS The author extends his appreciation to Roger Portell, FLMNH, for providing the plates and commenting on an earlier draft of the manuscript. Digital images were pre- pared by Sean Roberts, FLMNH. Thanks also to Mare Grigis, Belgium, for donation of a specimen of Mitrella ee to Dr. Bernard Landau, Portugal, for encour- agement and advice, and to Burke and Brooks Hayes and also Archie and Vicki Whittington for permission to collect on their respective properties in North Florida. Pamela McBride provided secretarial assistance. | am in- THE NAUTILUS, Vol. 122, No. 3 debted to two anonymous reviewers whose comments and suggestions improved the manuscript. LITERATURE CITED Campbell, L. D. 1993. Pliocene mollusks from the Yorktown and Chowen River Formation in Virginia. Virginia Divi- sion of Mineral Resources, Publication 127: vii + 173 pp., 43 pls.. Gardner, J. 1947. The molluscan fauna of the Alum Bluff Group of Florida. Part VII. Ctenobranchia (remainder), Aspidobranchia and Scaphopoda. United States Geologi- cal Survey. Professional Paper 142-H: i-ii + 493-638, pls. 52-62. Gardner, J. 1948. Mollusca from the Miocene and Lower Pliocene of Virginia and North Carolina. Part 2. Scaphopoda and Gastropoda. United States Geological Survey. Professional Paper 199-B: i-iii +179-279, pls. 24— 38. Keen, A. M. 1971. Sea Shells of Tropical West America. Marine mollusks from Baja California to Peru. Second Edition. Stanford University Press, Stanford, California: xiv + 1064 pp., 22 pls. Mac Neil, F. S. and D. T. Dockery IIL. 1984, Lower Oligocene Gastropoda, Scaphopoda, and C Jephalopoda of the Vicks- burg Group in Mississippi. Mississippi Bureau of Geology. Bulletin 124: 415 pp., 72 pls. Maury, C. J. 1910. New Oligocene (Miocene) shells from Florida. Bulletins of American Paleontology 4(21): 119— 164, pls. 1S—26. Vokes, E. H. 1989. An overview of the Chipola Formation, northwestern Florida. Tulane Studies in Geology and Pa- leontology 22: 13-24. THE NAUTILUS 122(3):155-165, 2008 Page 155 Catalogue of the type material of mollusks deposited at the Zoology Museum, University of Costa Rica Fresia Villalobos-Rojas Ana. G. Guzman-Mora Escuela de Biologia Universidad de Costa Rica Apdo. 2060, San Pedro San José, COSTA RICA Yolanda E. Camacho-Garcia! Escuela de Biologia and Museo de Zoologia Universidad de Costa Rica Apdo. 2060, San Pedro San José, COSTA RICA yeamacho_99@yahoo.com ABSTRACT This catalogue compiles the basic data for mollusk type mate- rial deposited at the Zoology Museum of the University of Costa Rica. It includes 62 holotypes and 151 paratypes, ee senting 68 species and 41 genera. The species, authors, date of publication, bibliographic references, type locality, catalogue number, number of specimens, state of preservation, and re- lated information are indicated for each taxon. In some cases, remarks on the current taxonomic status and other useful in- formation are provided. For all species, except for opistho- branchs, the shell of either the holotype or one of the paratypes is illustrated. Additional Keywords: Holotype, paratype, Mollusca, mollusk collection INTRODUCTION The Zoology Museum of the University of Costa Rica (MZUCR) was founded in 1960 by UCR professors Douglas C. Robinson and William Bussing. The collec- tion consists of both invertebrates and verchites: and includes birds, fishes, mammals, reptiles, amphibians, tu- nicates, decapods, sipunculids, sponges, corals, and mol- lusks, among other groups. The specimens come from a wide variety of research projects, and donations from national and international researchers, as well as amateur naturalists. Most of the material is preserved in a wet collection (70% ethanol), but there is also a dry collec- tion. THE MOLLUSK COLLECTION AT THE UNIVERSITY OF Costa RICA Former UCR curator Carlos Villalobos and a great num- ber of other researchers started the mollusk collection at the Museum in 1964. Since then, UCR staff and visiting scientists have continuously added to the collection. In addition, a significant amount of material has been do- ' Author for correspondence nated to the Museum. Some examples include the col- lections of malacologists George Richard, from Univer- sity of La Rochelle, France (mainly specimens of the family Conidae and other material from the Indo- Pacific), Kristie L. Kaiser, associated with the Santa Bar- bara Museum of Natural History (material from Cocos Island, Costa Rica), and the Latin American collection of independent scientist Dwight W. Taylor (including terrestrial and freshwater mollusks as well as type material); and amateur naturalists Michael Montoya (a valuable collection of marine mollusks from the Me- soamerican region, Cocos Island, and the Caribbean), Mary Yost (marine mollusks from Guanacaste, Costa Rica), Robert Nishimoto and others in 1969 (marine mollusks from Puntarenas, Costa Rica), and Jerry Well- ington in 1972 (marine mollusks from both the Pacific and the Atlantic). In 2005, the collection increased in size considerably, having absorbed the mollusk collection of the Costa Rican National Biodiversity Institute (INBio), which closed its Malacology Department in September 2004. Currently, the édllecten is comprehensive, in- cluding more than 37,670 lots and 216,587 specimens representing the molluscan fauna of the Pacific Coast of North, Central, and South America (however, there is also material from the Indo-Pacific, France, and Africa). The collection is divided into two main categories: a dry collection consisting mainly of the shells of bivalves and gastropods, and a wet collection comprising cepha- lopods, gastropods, bivalves, polyplacophorans, and ter- restrial and freshwater mollusks preserved in alcohol. Considering that this collection is the most important in the country and perhaps in all of Central America, and due to the recent increase in the amount of type mate- rial, it is important to compile this information and pub- lish a catalogue that addresses in detail the literature available for the type material, its location, and illustra- tions of the shell of each type represented. We hope to facilitate the future work of taxonomists and help locate type material, contributing in this respect to their re- search. Page 156 THE NAUTILUS, Vol. 122, No. 3 F. Villalobos-Rojas et al., 2008 Page 157 TYPE COLLECTION This catalogue includes 62 holotypes and 151 paratypes, representing 6S species and 41 genera. All types are listed alphabetically by species epithet. Photographs of opisthobranch types are not included, although they are listed in this catalogue. For each taxon, the following information is included: the name of the species cite i exactly as it was published in the original description, the author(s), date of publication, bibliographic references, type locality, catalogue number, number of specimens, state of preservation, and related information (SEM stubs, microscopy slides, egg masses of the specimens). In some cases, remarks on the current taxonomic status and other useful information are provided. We also in- dicate whether the specimen is preserved complete (in- cluding the shell and soft parts) or when only the shell remains in the dry collection. In a few cases, the information in the original publi- cation about the location of the type material is not ac- curate, and we clarify this in the remarks. For each spe- cies, either the holotype or one of the paratypes is pho- tographed, and the figure number is properly indicated beside the specimen. Although the original publications on some type ma- terial (Noumea regalis Ortea, Caballer and Moro, 2001; Dentimargo argonauta Espinosa and Ortea, 2002; Tico- cystiscus iberia Espinosa and Ortea, 2002; Cratena piu- taensis Ortea, Caballer, and Espinosa, 2003; Milleria ritmica Ortea, Caballer, and Espinosa, 2003; Phidiana adiuncta Ortea, Caballer, and Moro, 2004; and Costoa- nachis cascabulloi Espinosa and Ortea, 2004) stated that the material was deposited at IN Bio, in fact, the material was never deposited there or at the University of Costa Rica. Although attempts have been made to clarify the location of this material with the authors, none has been successful. Abbreviation used in the text: m.a.s.l. = meters above sea level. CATALOGUE OF TYPE MATERIAL academica, Okenia, Camacho-Garcia and Gosliner, 2004: 431-438, figs. 1-3. Type locality: Playa Tamarindo, ea Conservation Area, Puntarenas, Costa Rica 03/58" N, $5°51'08" W), 0 m depth. HOLOTYPE: aca BOOO3LIS1LO2 (specimen), MZUCR- INB0003764988 (radula and jaw, SEM stubs), PARATYPE: MZUCR-INB0001496648, San Miguel, Reserva Natural Absoluta de Cabo Blanco, Tempisque Conservation Area, Puntarenas, Costa Rica (9°34'53” N, $5°08'26" W), 0 m depth. aeci, Philinopsis, Ortea and Espinosa, 2001b: 41, pl. 2C, 3A. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°6'37" N, 82°66/08" W), 6 m depth. HOLOTYPE: MZUCR-INBO001495781. PARATYPE: MZUCR-INB0003721981. Same locality data as holo- type. Remarks: Paratype said to be deposited at the In- stituto de Oceanologia, Habana, Cuba, but was deposited at INBio. alfiopivai, Plesiocystiscus, Espinosa and Ortea, 2002: 102-106, figs. 1-3, pl.l. Type locality: Manzanillo, Limon, Costa Rica (9° By N, 82°39" W), 20-24 m depth. HOLOTYPE: MZUCR-INB0003754713 (shell, Fig- ure 21). anulatus, Janolus, oo Garcia and Gosliner, 2006: 1295-1305, figs. 1-7. Type locality: Isla Ballena, Parque Nacional Marino Ballena, Puntarenas, Costa Rica (9°06'24" N, $3°43'35" W), 6 m depth. HOLOTYPE: MZUCR-INB0001495772 (specimen), MZUCR- INB00037649558 (SEM stub with jaw), MZUCR- IN B0003764987 (SEM stub with radula), MZUCR- INB0003765066 (slide preparation with labial plate). PARATYPES: MZUCR-INB0005764915 (1 speci- men). Same locality data as holotype; MZUCR- INBOOO3S36171 (1 specimen), SW side of Isla Plata, Guanacaste, Cone Rica (10°26'48” N, 85°48'20" W), 10 m depth. arleyi, Melanella, Espinosa, Ortea and Magara, 2001: 123-124, fig. 3. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°37" N, §2°37" W), 10-12 m depth. HOLOTYPE: MZUCR-INB0003138455, (shell, Fig- ure 1). awapa, Doto, Ortea, 200 1a: 21-23, pl. 2G, fig. 9. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°37"” N, 82°37" W), 10 m depth. HOLOTYPE: MZUCR-INB0001497507 ( (specimen and egg mass). beatrix riopejensis, Helicina (“Gemma”), Richling, 2004: 303-308, figs. 133-138. Type locality: SW of Liv- erpool (about 24 ‘em of Puerto Limén) along Rio re Limon, Costa Rica (90°55'46"” N, 83°13 15” W ), 13. m.a.s.l. HOLOTYPE: MZUCR-INB003542625 (shell Figure 30). PARATYPE: MZUCR-INB0003542626 (shell). Same locality data as holotype. boeckereli, Alcadia (Microalcadia), Richling, 2001: 6-7, figs. 9-12. Type teeality Guanacaste National Park, about 10 km S of Santa Cecilia, Volean Orosi, near field station Pitilla, beginning of Sendero Orosilito, primary forest, Guanacaste, Costa Rica (10°59'18S"” N, §5°25'34" W), 700 m.a.s.l., HOLOTYPE: MZUCR- INB0003404980 (shell, Figure 24). Remarks: species was assigned to Alcadia in Richling, 2004: 374-377, figs. 257— 262: previously in Helicina. bramale, Hoplodoris, Fahey and Gosliner, 2003: 19S— 201, figs. 17E, 58-30. Type locality: Puerto Escondido, M: ainiiel Antonio National Park, Puntarenas, Costa Rica (9°23’ N, 84°08’ W), 0 m depth. HOLOTYPE: Figures 1-12. 1. Holotype of Mellanela arleyi, 2.1 mm. 2. Holotype of Melanella zugnigae, 1.1 mm. 3. Holotype of Triphora orteai, 2.0 mm. 4. P: uratype of Chicoreus | ee 137 mm. 5. Paratype of Typhisopsis carolskoglundae, length 19 mm diameter, 11mm. 6. Parz ee of Mitrella loisae, 4.9 mm. . Holotype of Hyalina chicoi, 9.1 mm. 8. Holotype of Prunwm cahuitaensis, 12.5 mm. 9. Holotype of Prunum chumi, 18.2 mm. 10. Holeae of Prunum holandae, 19.5 mm. 11. Holotype of Prunum lizanoi 13.2 mm. 12. Holotype of Volvarina socoae, 11.8 mm. Page 158 THE NAUTILUS, Vol. 122, No. 3 MZUCR-INB0003572316. PARATYPES: MZUCR- INB0003572306 (6 specimens), Punta Uvita, Marino Ballena National Park, Puntarenas, Costa Rica (9°08' N, 83°45’ W), 0-2 m ae MZUCR-INB0001498550 (1 specimen), 0 m depth. Same locality data as the other paratypes. bribri, Gibberula, Espinosa and Ortea, 2000: LO00- 101, pl 1, fig. 4. Type Locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°38' N, 82°37’ W), 10-15 m depth. HOLOTYPE: MZUCR-INB0003349908 (shell, Figure 16). PARATYPES: MZUCR-IN B0003349905, MZUCR- IN B0003349906, MZUCR-IN B0003349907 (1 specimen each, shells). Locality data for all paratypes same as ho- lotype. caballeri, Philine, Ortea, Espinosa and Moro, 2001: 38-40, figs. 9-10, pl. 2B. Type locality: Punta Mona, Miz aizanillo, Limon, Costa Rica (9°37' N, $2°37' W), 9m depth. HOLOTYPE: MZUCR- INB0003138445. cabecar, Doto, Ortea, 2001a: 34-37, figs. 16-17, pl. 2M. Type Locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°39' N, 82°37’ W), 20 m depth. HOLOTYPE: MZUCR-IN BO003449742. cahuitaensis, Prunum, Magana, Espinosa and Ortea, 20038: 122-124, figs. 1, 2A-B, pls. 1A, 2A. Type Locality: 1 km E from Puerto Vargas Station, Cahuita National Park, Limon, Costa Rica (9°43'33" N, 82°48'31” W). HOLOTYPE: MZUCR-INB0003718203 (shell, Figure 8). PARATYPE: MZUCR-IN B0003718204 (shell). Same locality data as holotype. caribetica, Gibberula, Espinosa and Ortea, 2002: 113-114, fig. 10, pL.1. Type locality: Punta Mona, Man- zanillo, Limon, Costa Rica (9°37' N, 82°37’ W), 8 m depth. HOLOTYPE: MZUCR-INB0003715205 (shell, Figure 17). carolskoglundae, Typhisopsis, Houart and Hertz, 2006: 56-58, figs. 17-25, 47-49, 59, 63. Type locality: Playas del Coco, Guanac: aste, Costa Rica (10°55'53” N, 85°69'S1" W), 24-37 m depth, on mud bottom. PARATYPE: MZUCR-6153, (shell, Figure 5), Boca de la Honda, Veraguas, Panama (7°27' N, 80°51’ W), in white sand. Remarks: The coordinates and the collecting local- ity of this paratype appear to be incorrect since these coordinates plot inland. chicoi, Hyalina, Espinosa and Ortea, 1999a: 167-169, figs. LA-C, 2A, 3A—D. Type locality: Manzanillo, Limon, Costa Rica (9°38' N, 82°39’ W), 5-12 m depth. HOLOTYPE: MZUCR-INB0003350839 (shell, Figure 7). Remarks: Paratype said to be de posited at INBio in the original publication, but never ve to INBio or the University of Costa Rica. chiquitica, Oligyra, Richling, 2001; 1-2, figs. I- 2. Type Locality: 9 km W of Matina, a little wre am up the {io Barbilla from the crossing of the road Siquirre s to Limon, along a tributary of Rio Barbilla, Limon, Costa Rica (10°03'29” N, 83°22'24” W), 70 m.a.s.l. HOLOTYPE: MZUCR-INB0003404977 (soft parts and shell, Figure 32). PARATYPE: MZUCR- INB0003404981 (shell and soft parts). chumi, Prunum, Espinosa and Ortea, 2000: 107-108, figs. 8-9. Type locality: Manzanillo, Limén, Costa Rica (9°3 8’ N, 82°39’ W), 10-15 m depth. HOLOTYPE: MZUCR-INB0003349912 (shell, Figure 9). PARATYPES: MZUCR-INB0003349913, MZUCR- INB0003349914, MZUCR-INB0003349915 (1 specimen each, shells). Locality data for all paratypes same as ho- lotype. convenientis, Eubranchus, Caballer and Ortea, 2002: 81-85, figs. 2, 3, 7 pl 1B. Type locality: Manzanillo, Limon, Costa Rica b»38) N, 82°39’ W), 0 m depth. HOLOTYPE: MZUCR-IN B0O003576832 corcovadensis, Cryptostrakon, Cuezzo, 1997: 1-S, figs. 1-14. Type locality: Corcovado National Park, Si- rena Station, Sendero a Rio Los Patos, Puntarenas, Costa Rica (8°30' N, 83°35’ W), 10 m.a.s.l. HOLOTYPE: MZUCR-INB0001468087 (shell, Figure 37; SEM stub with radula). PARATYPE: MZUCR-INB0001468080 (shell and soft parts). Remarks: Paratype locality data same as holotype. In the publication, the holotype and paratype catalogue numbers mistakenly written IN BO00468087 aad IN B000468080, respectively. costacubensis, Janolus, Ortea and Espinosa, 2000: 80-83, figs. 1-2. Type locality: Miramar, N coast of La Habana, “Caba (23°7'21" N, 82°25'10" W), 20-25 m depth. PARATYPE: MZUCR-INB0001497432 (1 speci- men), collecting locality Manzanillo, Limén, Costa Rica (9°38" N, 82°39! W), 20-25 m depth. ehuxmoralal, Dentimargo, Espinosa and Ortea, 2000: 110-113, fig. 11, pl. 2. Type locality: reefs from Manza- nillo, Limon, Costa Rica (9°38' N, 82°37’ W), 12-15 m depth. HOLOTYPE: MZUCR-INB0003349903 (shell, Figure 14). curere, Doto, Ortea, 2001a: 17-15, fig. 7, pl. 2C. Type locality: Puerto Viejo, Limon, Costa Rica (9°38' N, §2°39' W), 6 m depth. HOLOTYPE: MZUCR- IN B0001496453. destinyae, Cuthona, Hermosillo and Valdés, 2007: 119-124, figs. 1C, 4, 5. Type locality: La G dou. Zi- huatanejo, c uerrero, México (17°37. 854’ N, 101°33.562' W). PARATYPES: MZUCR-INBOO03118106 (4 speci- mens), Playa Avellanas, Guanacaste, Costa Rica (10°13.583' N, 85°50.433' W). duao, Doto, Ortea, 2001a: 28-30. figs. 12— 13, pls. 2: 2K. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°37' N, 82°37’ W), 10-19 m depth. HOLOTYPE: MZUCR-IN BO00313S803. echandiensis, Helicina, Richling, 2004; 271-277, figs. 77-84. Type locality: La Amistad National Park, Las Al- turas Sector, Southern Cordillera de Talamanca, S of Cerro Echandi, Campamento Echandi, Puntarenas, Costa Rica (09°01'33" N, 82°40'12" W), 2840 m.a.s.b. HOLOTYPE: MZUCR-INB0003542520 (soft parts and shell, Figure 26). PARATYPES: MZUCR- INB0003574064 (1 specimen, sott parts and_ shell), MZUCR-INB0003542521 (1 specimen, soft parts and shell), MZUCR-IN B0003428246 (19 specimens, shell and soft parts). Locality data for all paratypes same as holotype. F. Villalobos-Rojas et al., 2008 Page 159 elizabethae, Adrana, Ortea and Espinosa, 2001e: 61— 64, fig. 17. Type locality: in front of Gandoca beach, Limon, Costa Rica (9°36' N, §2°35’ W), 10-15 m depth. HOLOTYPE: MZUCR-INB0003449558 (shell, Figs. 38-39). escondida, Helicina (“Gemma”), Richling, 2004: 348— 357, figs. 210-218. Type locality: 9 km W of Matina, a little stream up the Rio Barbilla from the crossing of the road from Siquirres to Limén, along a tributary of Rio Barbilla, Limon, Costa Rica (10°03'29” N, 83°22'24” W), 70 m.as.l. HOLOTYPE: MZUCR-INB0003542623 (soft parts and shell, Figure 27), PARATYPE: MZUCR- INB0003542624 (soft parts and shell). Locality data same as holotype. espinosai, Ancula, Ortea, 2001b: 49, pl. 2D. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°37' N, 82°37' W),9m depth. HOLOTYPE: MZUCR- IN B0003188764. eugeniae, Elysia, Ortea and Espinosa, 2002: 130-133, figs. 1-2, pl. LA. Type locality: Manzanillo, Limon, Costa Rica (9°38’ N, 82°39’ W), 16 m depth. HOLOTYPE: MZUCR-INBO001497478. eversonii, Phyllonotus, D’Attilio, Myers and Shasky, 1987: 162-164, figs. 1-2. Type Locality: SW side of Isla Manuelita, Isla del Coco, Costa Rica (5°33’ N, 87°03’ W, 66 m depth. PARATYPE: MZUCR 4934 (ahell. Fig- ure 4). gandocaensis, Rissoella, Ortea and Espinosa, 2001a: 36, pl. 2A. Type locality: Punta Mona, Manzanillo, Limén, Costa Rica (9°37 N, 82°37 W), 9 m depth. HOLOTYPE: MZUCR-INB0003323831. Remarks: shell destroyed, only dry soft body present, not figured here. genecoani, Plesiocystiscus, Espinosa and Ortea, 2000: 96-97, fig. 1, pl. 1. Type locality: Manzanillo, Limon, Costa Rica (9°39' N, 82°37’ W), 25 m depth. HOLOTYPE: MZUCR-INB0003349916 (shell, Figure 22). PARATYPES: MZUCR-INB0003349917 (2 speci- mens, shells). Locality data same as holotype. goslineri, Cylichnella, Valdés and Camacho-Gareia, 2004: 459-497, figs. 4D-F, 5. Type locality: Sector Playi- tas, Golfo Balec. Puntarenas, Costa Rica (8°44'19”" N, 83°21'57" W), 0 m eee HOLOTYPE: MZUCR- IN B0001497964 (shell, Figure 23). PARATYPES: MZUCR-INB0003718957 (15 specimens, shell and soft parts, lateral gizzard plate and radular teeth in SEM stubs). Locality data same as holotype. hojarasca, Alcadia (Microalcadia), Richling, 2001: 5-6, figs. 6-8. Type locality: cordillera de Tilaran, about 9 km N of Santa Elena, near Mirador Gerardo, Guana- caste, Costa Rica (10°22’19" N, 84°48’25” W), 1450 m.as.l. HOLOTYPE: MZUCR-INB0003404979 (shell, Figure 25). Remarks: species was assigned to Alcadia in Richling, 2004: 370-374, figs. 249- 255: it was previously Helicina. holandae, Prunum, Espinosa and Ortea, 1999b: 175— 176, fig 1H. Type locality: Cayos Limon, Islas San Blas, Colén, Panama (9°33'00” N, 78°53’30” W) HOLOTYPE: MZUCR-5750 (shell, Figure 10). PARATYPE: MZUCR-INB0003350838 (shell), Punta Uvita, Manzanillo, Limon, Costa Rica (9°38’ N, 82°41’ W). Remarks: According to the original publication the holotype was deposited at the Instituto de Oceanologia de la Habana, Cuba, but in fact it was deposited at IN- Bio. The paratype was erroneously assigned the cata- logue number INBIOCR1001501498 in the original pub- lication. Also, the locality for the paratype, Punta Uvita, was misspelled in the original publication as Punta “Ubitas.” inbiotica, Trapania, Camacho and Ortea, 2000: 317- 321, figs. 1-3. Type locality: San Miguel Station, Reserva Absoluta de Cabo Blanco, Tempisque Conservation Area, Puntarenas, Costa Rica (9°34'49”" N, 85°08'28”" W). lm depth. HOLOTYPE: MZUCR-INBOOO1500889. iugula, Doto, Ortea, 2001a: 26-27, fig. 11, pl. 2]. Type locality: Punta Mona, Limén, Costa Rica (9°39’ N, 82°37' W), 25 m depth. HOLOTYPE: MZUCR- IN B0003449604. kekoldi, Doto, Ortea, 2001a: 18-21, fig. 8, pl. 2 E-F. Type locality: Punta Mona, Limon, Costa Rica (9°35' N, 82°37’ W), 9 m depth. HOLOTYPE: MZUCR- INB0003449587 (soft body and egg mass). leopoldoi, Eubranchus, Caballer, Ortea and Espi- nosa, 2001: 55-56, fig. 14, pl. 2E. Type locality: Punta Mona, Manzanillo, Limén, Costa Rica (9°38' N, 82°37’ W). 6 m depth. HOLOTYPE: MZUCR- INB0003138799. lizanoi, Prunum, Magafia, Espinosa, and Ortea, 2003: 124-126, figs. 2C, 3, pls. 1B, 2 B. Type locality: Bahia Junquillal Wildlife Refuge, Golfo de Santa Elena, Guanacaste, Costa Rica. HOLOTYPE: MZUCR- INB0003481195 (shell, Figure 11). PARATYPES: MZUCR-INB0003481192 (1 specimen, shell, including a slide with radula), MZUCR-INB0003476231 (1 speci- men, shell and soft parts). loisae, Mitrella, Pitt and Kohl, 1979: 467-468, figs. 2A-B, 3A-B. Type locality: N side of Punta Coralillo, Bahia de Caldera, about 20 km S of the city of Puntar- enas, Puntarenas, Costa Rica (9°54' N, 84°44’ W). PARATYPES: MZUCR 2363 (2 specimens, shell, Fig- ure 6). magagnai, Dendrodoris, Ortea and Espinosa, 2001d: 52-53, fig. 13, pl. 3E. Type locality: Manzanillo, Limon, Costa Rica (9°38' N, 82°39" W), 20 m depth. HOLOTYPE: MZUCR-INB0001497496. manzanilloensis, Polycera, Ortea, Espinosa, and Ca- macho, 1999: 161-163, fig. 3. Type locality: Manzanillo, Limon, Costa Rica (9°38’ N, 82°39’ W), 8 m depth. HOLOTYPE: MZUCR-INB0001496124. marioi, Gibberula, Espinosa and Ortea, 2000: 101-— 102, fig. 5, pl. 1. Type locality: coral reefs of Manzanillo, Limon, Costa Rica (9°39' N, 82°39’ W), 30 m depth. HOLOTYPE: MZUCR-INB0003371977 (a slide prepa- ration with radula). millenae, Cuthona, Hermosillo and Valdés, 2007: 124-128, figs. 1D, 6, 7. Oy locality: Los Arcos, Bahia de Randers, Jalisco, México (20°3 32.855’ N, 105°17.340' W), 19 m depth. PARATYPE: MZUCR- INB0003836263. Playa Real, NE Punta Roble, Guana- Q 0d 122, No. NAUTILUS, Vol. THE Page 160 F. Villalobos-Rojas et al., 2008 Page 16] Figures 24-32. 24. Holotypes of Alcadia (Microalcadia) boeckereli, 2.2 mm. 25. Alcadia (Microalcadia) hojarasca, 2.4 mm. 26. Helicina (“Tristramia”) echandiensis, 7.2 mm. 27. Helicina (“Gemma”) escondida, 6.2 mm. 28. Helicina (“Gemma”) monteverdensis 6.6 mm. 29. Helicina (“Gemma”) talamancensis, 9.2 mm. 30. Helicina (“Gemma”) beatrix riopejensis, 7.8 mm. 31. Helicina (“Tris- tramia”) punctisulcata cuericiensis, 5.9 mm. 32. Oligyra chiquitica, 4.8 mm Figures 13-: ey) 13. Holotypes of Volvarina yolandae, 6.2 mm. 14. Dentimargo cruzmoralai, 2.5 mm. 15. Dentimargo zaidetta 16. Gibberula bribri. 11.0 mm. 17. Gibberula caribetica, 6.0 mm. 18. Gibberula sierrai, 2.1 mm. 19. Gibberula ubitaen 2.5 1S mm. 20. Granulina minae, 1.0 mm. 21. Plesiocysticus alfiopivai, 1.9 mm. 22. Plesiocysticus genecoani, 2.2 mm. 2« J i ] o 4.5 mm arf ylic hne Page 162 THE NAUTILUS, Vol. 122, No. 3 F. Villalobos-Rojas et al., 2008 Page 163 caste, Costa Rica (10°23.200' N, 85°50. depth. minae, Granulina, Espinosa and Ortea, 2000: 103, fig. 6, pl. 1. pis loc: uity: Punta Mona, Manzanillo, Limon, Costa Rica (9°37’ N, $2°37' W), 12-15 m depth. HOLOTYPE: MZUCR- INBO003: 349904 (shell, Figure 20). monteverdensis, dlc (“Gemma”), Richling, 2004: 334-348, figs. 183-199. Type locality: Cordillera de Tilaran, near Monteverde, Zona Protectora Arenal- Monteverde, Reserva Biolégica Bosque Nuboso Mon- teverde, Sendero Bosque Nuboso, Puntarenas, Costa Rica (10°18S'08” N, 84°47'41” W), 1550 m.a.s.l. HOLOTYPE: MZUCR-INB0003542627 (shell, Figure 28). PARATYPE: MZUCR-INB0003542628S (shell). Lo- cality data same as holotype. orteai, Cyerce, Valdés and Camacho-Garcia, 2000: 445456, figs. 1-5. Type locality: Playa Cabuya, en Coébano, Puntarenas, Costa Rica (9°39’ N, 85°11’ W), ( m depth. HOLOTYPE: eit Ce th ucGIEOUCEL. PARATYPE: MZUCR-INBO001500644. Locality data same as holotype. orteai, Triphora, Espinosa, 2001: 21-22, fig. 7. Type locality: Punta Mona, Manzanillo, Limén, Costa Rica (9°37” N, 82°37” W), 10-15 m depth. HOLOTYPE: MZUCR-INB0003138795 (shell, Figure 3). osae, Jorunna, Camacho-Garcia and Gosliner, 2008: 165-167, figs. 21-22. Type locality: Playa eng Golfito, Osa Conservation Area, Costa Rica (8°38'33" $3°13'40" W), O m depth. HOLOTYPE: WaGce INB0003701453, MZUCR-INB0003799440 (radula and jaw, SEM stubs). pacifica, Chiapaphysa, Taylor, 2003: 170-171, fig. 168, pl. 8.8. Type locality: Rio Tenorito, Hacienda La Pacffica, Guanacaste, Costa Rica (10°29.02' N, $9°09.58' W). 100 m.a.s.l. PARATYPES: MZUCR- INB0003352239 (10 specimens, shells, Figure 33). proranao, Doto, Ortea, 2001a: 23-25, fig. 10, pl. 2H. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°37' N, $2°37' W), 9 m depth. HOLOTYPE: MZUCBR-IN B0003 138763. punctisulcata cuericiensis, Helicina, Richling, 2004; 277-283, figs. 87-96. Type locality: Cordillera de Tala- manca, Estacién Cuerici, 4.5 km E of Villa Mills, Sen- Se al Mirador, Cartago, Costa Rica (9°33'28" N, 3°40'13" W), 2700 m.a.s.l. HOLOTYPE: MZUCR- cet era (shell, Figure 31). PARATYPE: MZUCR-IN B0003542541 (shell and soft parts). Other ae ity data same as holotype (09°33'19" N, 83°40'13" 2600 m.a.s.l.). ee Mayabina, Taylor, 2003: 102-104, fig. 85, pls. 3.5, 3.6. ee locality: Barra del Colorado, Limon, Costa Rica (10°46.37' N, 83°35.27' W). PARATYPES: MZUCR-INB0003352237 (10 specimens, 733' W), 5 m shell), MZUCR 69-01 (10 specimens, shell, Figure 34), Locality data for all paratypes same as holotype selva, Ercolania, Ortea and meee o00le: 45-47, fig. 11. Type locality: Manzanillo, Limén, Costa Rica (9°39' N, 82°39’ W) ), 0m depth. HOLOTYPE: MZUCR- INBOOO313832 PARATYPE: MZUCR- IN B0003449624. ae data same as holotype. sierrai, Gibberula, Espinosa and Ortea, 2000: 99-100, fig. 3, pl 1. Type locality: Punta Mona, Manzanillo, Limon, Costa Rica (9°38’ N, 82°37’ W), 10-15 m depth. HOLOTYPE: MZUCR-INB0003349909 (shell, Fig- ure 18). sinusdulcensis, Tropinauta, Taylor, 2003: 110-111, figs. 91-94, Type locality: small stream in pasture 3 km SE of Golfito, Puntarenas, Costa Rica (8°36.68’ N, 3°8.48' W). PARATYPES: MZUCR-INB0003382246 (5 specimens, shells, Figure 36). socoae, Volvarina, Espinosa and Ortea, 1999b; 171— 172, figs. LE, 2D-E. Type locality: Manzanillo, Limon, Costa Rica (9°39' N, 82°45’ W), 1.0-1.5 m de »pth. HOLOTYPE: MZUCR-INB0003350887 (shell, Figure 12). PARATYPE: MZUCR-INB000335088S (shell). Re- marks: in the original publication the holotype and paratype were erroneously assigned the catalogue num- bers INBIOCR001496127 and INBIOC R1001496] 28, re- spectively. talamancensis, Helicina (“Gemma”), Richl ing, 2001: 3-5, figs. 3,4. Type locality: Fila Costenia, N of Bajo Bo- nito, N of RioC laro, Puntarenas, Costa Rica (8°44'41” N, $3°02'09”" W), 980 m NN. HOLOTYPE: MZUCR- INB0003404978 (shell, Figure 29). PARATYPES: MZUCR-INB0001494642 (8 specimens, shell), MZUCR-INBO0001487761 (2 specimens, shell and soft parts), 3.5 kin from Escuela de Llano Bonito Carretera a San Vito, Puntarenas, Costa Rica (8°44'27” N, 83°02'04" W), 840 m m.as.l., MZUCR-INB0001494509 (1 speci- men, shell and soft parts), 3.5 km NE from Escuela de Llano Bonito, Puntarenas, Costa Rica (8°44'54" N, §3°02'04" W), 920 m m.a.s.l. Remarks: species was as- signed to Helicina in Richling, 2004; 308-315, figs. 140, 142- 14S; it was previously Olygira el maces The paratype MZUCR-INBO0003389580 (1 specimen, shell, locality data same as MZUCR-INB0001494509) is said to be deposited at INBio but is not present. tempisquensis, Jorunna, Camacho-Garcia and Gos- liner, 2008: 167-170, figs. 23-24. Type locality: Cabo Blanco, Costa Rica (9°34'50" N, 85°08'26" W), 0 m depth. HOLOTYPE: MZUCR-INBOOO 35.423 76. PARATYPES: MZUCR-INB0003542377 (2 specimens). tempisquensis, Mayabina, Taylor, 2003: 109-111, fig. 90, pl a. 4. Type locality: edge of marshes, 100 m E of W end of airstrip, Palo Verde National Park, Guanacaste, Costa Rica (10°20.68’ N, 85°20.60' W). PARATYPES: Figures 33-39. 33. Paratype of Chiapaphysa pac ifica, 6.1 mm. 34. Paratype of Mayabina sanctijohannis, 6.7 mm (UCR 69-01). 35. Paratype of Mayabina tempisquensis, 7.5 mm. (INB0003352244). 36. Paratype of Tropinauta sinusdulcensis, 5.5 mm. 37. Holotype of Cryptostrakon corcovadensis, 4 mm. 38. Holotype of Adrana elizabethae, (left valve dorsal view) 32.5 mm. 39. Holotype of Adrana elizabethae, (left valve ventral view) 32.5mm Page 164 THE NAUTILUS, Vol. 122, No. 3 MZUCR-INB0003382244 (10 specimens, shell, Figure 5), MZUCR 70-01 (10 specimens, shell). tica, Furcilla, Espinosa and Ortea, 2000: 105-107, fig. 7, pl. 1. Type locality: coral reefs of Manzanillo, Limén, Costa Rica (9°39' N, 82°39’ W), 30 m depth. HOLOTYPE: MZUCR-IN B0003371976 (including a slide with radula). tica, Mexichromis, Gosliner, Ortea, and Valdés, 2004: 589-593, figs. LA, 2, 3. Type locality: Bajo del Diablo, Isla del Cano, Punts irenas, Costa Rica (8°42'04" N, §3°53'20" W), PARATYPES: MZUCR-INBO001486639 (2 specimens). ubitaensis, Gibberula, Espinosa and Ortea, 2000: 97— 99, fig. 2, pl. 1. Type locality: Punta Uvita, Manzanillo, Limon, Costa Rica (9°38’ N, 82°41’ W), 12-15 m depth. HOLOTYPE: MZUCR-INB0003349902 (shell, Figure 19). Remarks: the Type locality, Punta Uvita, was mis- spelled in the original publication as “Ubita.” yolandae, Volvarina, Espinosa and Ortea, 2000: L0S— 110, fig. LO, pl 1. Type locality: Manzanillo, Limon, Costa Rica (9°39’' N, 82°39’ W), 10-25 m depth. HOLOTYPE: MZUCR-INB0003350843 (shell, Figure 13). PARATYPES: MZUCR-INB0003350541 (1 speci- men, shell), MZUCR-INB0003350842 (1 specimen, shell). Locality data same as holotype. Remarks: accord- ing to the original publication, the paratype MZUCR- INB0003350841 was de »posited at the Instituto de Oceanologia de la Habana, Cuba, and the paratype MZUCR-INB0003350842 was deposited at Museo de Ciencias Naturales de Tenerife, Canary Islands; how- ever, both are depx ysited at MZUCR. zaidettae, Dentimargo, Espinosa and Ortea, 2000: 113-114, fig. 12, pl. 2. Type locality: Punta Mona, Man- zanillo, Limon, Costa Rica (9°38' N,$2°37' W), 12-15 m depth. HOLOTYPE: MZUCR-INB0003349911 (shell, Figure 15). PARATYPE: MZUCR-INBO003349910° (1 specimen, shell). Locality data same as holotype. Re- marks: According to the original publication the paratype was deposited at the Instituto de Oceanologia de la Ha- bana, Cuba, but in fact it is deposited at MZUCR. sugnigae, aging Espinosa, Ortea and Magana, 2001: 26-27, figs. D. Pe locality: Punta Mona, Manzanillo, ‘Limon, i“ mee Rica (9°37' N, 82°37' W), 10- 15 m de pth. HOLOTYPE: pea nmi (shell, Figure 2), ACKNOWLEDGMENTS We are very grateful to Alvaro Morales and Ingo Werth- mann for their logistic support during the elaboration of this paper. The photographs were taken at the facilities of Centro de Investigaci6n en Ciencias del Mar y Lim- nologia (CIMAR) at the University of Costa Rica. Eu- gene V. Coan and David Butvill made constructive com- ments on this manuscript. LITERATURE CITED Caballer, M., J. Ortea, and J. Espinosa. 2001. Descripcién de una nueva especie de Eubranchus Forbes, 1834, Avicen- nia Suplemento 1. 55-56 Camacho, Y. and J. Ortea. 2000. A new species of Trapania (Nudibranchia: Goniododididae) from the Pacific coast of Central America. Revista de Biologia Tropical 48: 317— 322. Camacho-Garcia, Y. and T. Gosliner, 2004. A new species of Okenia (Gastropoda: Nudibranchia: Goniodoridae) from the Pacific coast of Costa Rica Proceedings of the Califor- nia Acade my of Sciences 55: 431-438. Camacho-Garcia, Y. and T. Gosliner. 2006. A new species of the zephyrinid nudibranch genus Janolis (Mollusca: Nudi- branchia) from North America and Costa Rica. Revista de Biologia Tropical 54: 1295-1305. Camacho-Gareia, Y. and T. Gosliner. 2008. Systematic revision and morphological phylogenetic analysis of the genus Jo- runna Bergh, 1876 (Nudibranchia: Discodorididae). Jour- nal of the Molluscan Studies 74: 143-151. Espinosa, J. and J. Ortea. 1999a. be nuevas especies del género Hyalina Schumacher, 1817 (Mollusca: Neogas- tropoda: Marginellidae) del Me ur Caribe de Costa Rica y costas de Cuba. Avicennia 10/11: 165-169. Espinosa, J. and J. Ortea. 1999b. Descripcion de nuevas mar- ginelas (Mollusca, Neogastropoda: Marginellidae) de Cuba y del Caribe de Costa Rica y Panama. Avicennia 10/11: 165-179. Espinosa, J. and J. Ortea. 2000. Descripci6n de un género y once especies nuevas de Cystiscidae y Mz wreinellidae ( Mol- lusca: Neogastropoda) de | Caribe de Costa Rica. Avicennia 12/13: 95-114. Espinosa, J. 2001. Descripci6n de una nueva especie de ee (s.L.) Blainville, 1S28. Avicennia Suplemento 4: 21- E repinosa, i J. Ortea, and J. Magana. 2001. Deseripcién de una nueva especie de Me ‘lanella Bowdich, 1822 y redescripcién de Melanella arcuata (c. B. Adams, 1850), Avicennia Suplemento 14, pp. 120-124. Espinosa, J. and J. Ortea. 2002. Nuevas especies de margineli- formes de Cuba, Bahamas y el Mar Caribe de Costa Rica. Avicennia Suple mento 15: 101-128. Espinosa, ]. and J. Ortea. 2004, Nuevas especies de moluscos gasteropodos marinos Mollusca: Gastropoda de las Baha- mas, Cuba y el Mar Caribe de Costa Rica. Revista de la Academia Canarias de Ciencias. 15: 207-216. Gosliner, T., J. Ortea, and A. Valdés. 2004. New data on tropi- cal Eastern Pacific Chromodorididae (Nudibranchia: Do- ridina) with description of a new species of Mexichromis Bertsch, 1977. Proceedings of the California Academy of Sciences 55; 588-597. : Hermosillo, A. and A. Valdés, 2007. Five new species of aeolid nudibranchs (Mollusca, Opisthobranchia) from the tropi- cal eastern Pacific. American Malacology Bulletin 22: 119— 37 : Houart, R and C. M. Hertz. 2006. A review of Typhisopsis Jousseaume, L880, and Typhisala Jousseaume, LSS1 (Gas- es Muricoidea) of the eastern Pacific, The Nautilus 120; 52-65, Magana, I. 4. Espinosa, and J. Ortea. 2003, Descripcién de dos nuevas especies del género Prunum Herrmannsen, 1852 (Mollusca: Gastropoda: Marginellidae) del Caribe y el Pacifico de Costa Rica. Avicennia 16; 121-128. Ortea, J.. J. Espinosa, and Y. Camacho. 1999, Especies del genero Polycera Cuvier, 1816 (Mollusca: Nudibranchia) recolectadas en la epifauna de algas rojas del Caribe de Costa Rica y Cuba. Avicennia 10/11: 157-164. Ortea, J. and J. Espinosa, 2000. Nueva especie del género Jano- F. Villalobos-Rojas et al., 2008 Page 165 lus Bergh, 1884 (Mollusca: Nudibranchia) de Cuba y Costa Rica. Avicennia 12/13: 79-83. Ortea, J. 2001a. El género Doto Oken, 1815 en el Mar Caribe: Historia natural y descripcion de nuevas especies. Avicen- nia Suplemento 3: 1-46. Ortea, J. 2001b. Descripcién de una nueva especie de Ancula Loven, 1S46, Avicennia Suplemento 4: 49. Ortea, J. and J. Espinosa. 2001a. Descripci6n de una nueva especie del género Rissoella Gray, 1S47. Avicennia Suple- mento 4: 36. Ortea, J. and J. Espinosa. 2001b. Descripcién de una nueva especie de Philinopsis Pease, 1560, Avicennia Suplemento 4: 41. Ortea, J. and J. Espinosa. 2001c. Descripci6n de una nueva especie de Ercolania Trinchese, 1872. Avicennia Suple- mento 4: 45-47. Ortea, J. and J. Espinosa. 2001d. Descripcién de una nueva especie de Dendrodoris Ehrenberg, 1831. Avicennia Suplemento 4: 52-53. Ortea, J. and J]. Espinosa. 2001le. Descripcién de una nueva especie de Adrana H. y A. Adams, 1858. Avicennia Suple- mento 4: 61-64. Ortea, J., J. Espinosa, and L. Moro. 2001. pen bike de una nueva especie de Philine Ascanius, 1772. Avicennia Suple- mento 4: 38-40. Ortea, J. and M. Caballer. 2002. Nuevos datos sobre el género Eubranchus Forbes, 1838 (Mollusca: Nudibranchia) en aguas templadas del Atlantico Oeste. Avicennia Suple- mento 15: 77-90. Ortea, J. and J. Espinosa. 2002. Nuevas Especies del género Elysia Risso, 1S1S (Mollusca: Sacoglossa) con caracteres singulares. Avicennia, Suplemento 15: 129-140. Ortea, J., M. Caballer, y J. Espinosa. 2003. Nuevos Aeolidaceos (Mollusca: Gastropoda) de Costa Rica. Avicennia 16; 129- 142. Ortea, J., M. Caballer, and L. Moro. 2004. Dos aeolidaceos con ceratas rojos de la region macaronesica y el Mar Caribe (Mollusca: Nudibranchia). Vieracea 32: 83-96. Richling, 1. 2001. New species of Helicinidae from Costa Rica (Mollusca: Neritopsina). Schriften zur Malakozoologie 17; 1-8. Richling, I. 2004. Classification of the Helicinidae: Review of morphological characteristics based on a revision of the Costa Rican species and application to the arrangement of the Central American mainland taxa (Mollusca: Gas- tropoda: Neritopsina). Malacologia 45: 195-450, Taylor, D. W. 2003. Introduction to Physidae (Gastropoda, Hy- grophila); biogeography, classification, morphology. Re- vista de Biologia Tropical 51: 1-299. Valdés, A. and Y. Camacho-Garcia. 2000. A new species of Cyerce Bergh, 1871 (Mollusca, Sacoglossa, Polybranchi- idae) from the Pacific Coast of Costa Rica. Bulletin of Marine Science 66; 445—456. Valdés, A. and Y, Camacho-Garcia. 2004. “Cephalaspidean™ Heterobranchs (Gastropoda) from the Pacific Coast of Costa Rica. Proceedings of the California Academy of Sci- ences 55: 459-497. THE NAUTILUS 122(3):166-170, 2008 Page 166 Dilemma japonicum new species (Bivalvia: Anomalodesmata: Poromyidae): A new record of the genus from the Northwest Pacific Takenori Sasaki The University Museum, The University of Tokyo 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033 JAPAN sasaki@um.u-tokyo.ac.jp José H. Leal P. O. Box 1580 The Bailey-Matthews Shell Museum Sanibel, FL 33957 USA jleal@shellmuseum.org ABSTRACT The fourth species of the bivalve genus Dilemma Leal, 2008, is described from disarticulated valves collected off central Japan. The discovery of these specimens represents a significant range extension for the genus to the Northwest Parihie, The new species is distinguished from the other three known species by its surface sculpture, shape of escutcheon, and hinge. The pro- dissoconch, indicative of lecithotrophic dev elopment, and shell microstructure, with outer homogeneous and inner nacreous layers, are illustrated for the first time for a species of the genus. Additional Keywords: Septibranchia, Poromyoidea, Japan, nacreous layer, prodissoconch INTRODUCTION Dilemma is a recently established, unusual genus of Anomalodesmata (Leal, 2008). The first described spe- cies of the genus, D. inexpectatum (Crozier, 1966), was dredged Goan bathyal depths off northern New Zealand and originally assigned to Corculum, family Cardiidae (Crozier, 1966) due to general similarities in ‘shell shape. However, its original familial allocation was questionable from habitat and depth alone, because Corculum species inhabit shallow lagoons of tropical waters and utilize pho- tosynthesis-derive ad energy by symbiotic dinoflage ‘lke S (Farmer et al., 2001). Subse quent discovery of obviously related species with preserve -d soft parts ‘allowed Leal (2008), using details of the macroanatomy, to demon- strate (1) that Corculum inexpectatum and the other, then newly found species are related and deserved grouping under a new genus, (2) that the new genus should be included in the Poromyidae, and (3) that spe- cies of Dilemma are carnivores. In fact, carnivory is a feeding habit that is common in the Anomalodesmata, a group of bivalves mostly present in the deep sea. Three species were originally (Leal, 2008) assigned to the genus: (1) Dilemma frumarkernorum Leal, 2008, from off Key West Florida, 229 m, (2) D. spectralis Leal, 2008, from off Vanuatu, 950-961 m, and (3) D. inexpec- tatum (Crozier, 1966) from off Three Kings Islands, New Zealand, $05 m. Their geographic distribution is disjunct, but shell morphology is surprisingly similar between the species of Florida and Southwest Pacific. During a research cruise off central Japan, in 2000, the senior author collected small unpaired valves of Corculum- like bivalves, but its locality (240-273 m) was obviously too deep for Corculum. In es the inner shell layer is iridescent, indicating presence of a nacreous layer. This conchological feature alone precluded inclusion of this new species in the Cardiidae. However, lack of soft parts preve ted a better resolved familial or supra- familial placement of the new taxon. The publication by Leal (2008) allowed allocation of the unknown bivalve in that newly established genus of the Poromyidae. In this paper, we describe this most interesting new species and report a new record of the genus from the Northwest Pacific. MATERIALS AND METHODS Four unpaired valves were sorted from sediments dredged trom southeast of Kamogawa, Chiba Prefecture, Japan, with a biological dredge (R/V TANSEI-MARU, cruise KT-00-05, station 1, Aen m, 34°59.963' N, 140°27.159' E— 35°00.020' 140°28.427' E, May 17, 2000). All specimens were so and disarticulated. The shells were coated with platinum and vanadium and pho- tographed with a scanning electron microscope (Hitachi S-2250N), following a standard method (see Geiger et al., 2007). The holotype and two paratypes are deposited in the Department of Historical Geology and Paleontol- ogy, The University Museum, The University of Tokyo (U MUT), and one paratype in The Bailey-Matthews Shell Museum, Sanibel, Florida (BMSM), SSS SSS SSS SSS SSD T. Sasaki and J.H. Leal, 2008 Figures 1+. valve. 1. Holotype, UMUT RM 29689. 2. Paratype 1, UMUT RM29690, 3. Paratype 2, UMUT RM29691, 4. Paratype Dilemma japonicum new a saa 1. Lateral view. 2. Posterior view. 3-4. Interior view. 1-3. Left valve. 4. Right 3, BMSM 17983. Abbreviations: et = cardinal-like ico e = escutcheon; k = keel; It = lateral tooth; p = projection in hinge; s = socket. SYSTEMATICS Superfamily Poromyoidea Dall, 1886 Family Poromyidae Dall, 1886 Genus Dilemma Leal, 2008 Type Species: Dilemma frumarkernorum Leal, 2008, by original designation. Diagnosis: Shell compressed anteroposteriorly and expanded laterally; lateral outline of articulated valves cardioid; umbones projecting dorsally and located ante- riorly; sharp oblique carina dividing anterior and poste- rior regions; maximum growth axis having ca. 30° against anteroposterior axis; hinge axis short with cardinal-like tooth and socket in each valve; lateral tooth reduced, present only in right valve; ligament external, double- layered: lithodesma absent; shell interior lined with sheet-like nacreous layer. See Leal (2008) for anatomical characters. Dilemma japonicum new species Figures 1-12 Diagnosis: Posterior region of shell sculptured by rough, lamellate, thin, commarginal ribs; surface irregu- larly punctate on anterior region but puncta radially ar- ranged on posterior region; escutcheon not distinct de- marcated; prominent projection present in posterior re- gion of hinge. Description: Shell thin, fragile, compressed antero- posteriorly; shell height larger than shell length. Anterior and posterior regions clearly demarcated by sharp keel Figure 1, k): anterior region narrower and less inflated than posterior. Surface of anterior region smooth, flat- tened in mz acroscopic view, but microscopici ally punctate (Figure 5). Surface of posterior region sculptured by sev- eral weak radial ribs (Figure 3), aad more prominently so by dense commarginal ribs, also punctate in enlarged view, but puncta arranged neatly in radial direction un- like on anterior region (Figure 6). Umbo angulated by keel and rib along posterior margin (Figure 7); umbonal cavity deep: umbo involute (Figure 3). Prodissoconch simple, flattened, shield-like, ca. 200 2m in length, with- out prominent sculpture or division into prodissoconch I and II (Figure 8). Hinge provided with single cardinal- like tooth and socket on each valve (Figures 3-4, 7, et, s); cardinal-like tooth located anterior to socket on left valve (Figure 3), their position reversed on right valve (Figure 4). Lateral tooth small, present only on right valve ( (Fig- ure 4, It). Single sharp projection ( (Figures : 3-4, 7, p) prominent on posterior side of hinge in similar postion and size in right and left valves. Ligament and outlines of muscle scars not observed in disarticulated valves. Shell margin consisting of two layers, outer homogeneous structure and inner nacreous structure (Figure 9-10, OL, IL). In outer layer elongate granules arranged ver- tically but lacking clear boundary. In inner layer suboval tablets fusing in growth region (Figure 11) sheet-like layers forming nacre (Figure 12). numerous Type Material: Holotype, 4.7 mm (height: SH) « 2.0 mm (length: SL), UMUT RM29689 (Figure 1); Paratype 1,5.6 mm (SH) « 2.4 mm (SL), UMUT RM29690 (Fig- ure 2); Paratype 2, 6.0 mm (SH) x 3.0 mm (width) UMUT RM29691 (Figure 3); Paratype 3, 9.5 mm (SH) 4.0 mm (SL), BMSM 17983 (Figure 4). Page 168 THE NAUTILUS, Vol. 122, No. 3 Figures 5-8. ; hinge. 8. Prodissoconch. 5-6. Holotype, UMUT RM29689. tooth; p = projection in hinge; pd = prodissoconch; s = socket Southeast off Kamogawa, Chiba Pre- 34°59,963'-35°00.020" N, Type Locality: fecture, Japan. 240-273 m, 140°27.159'—140°28.427' E. Distribution: Known only from the type locality. Etymology: The species epithet recognizes the coun- try of the type loc: lity. The epithet japonicum agrees in vender with the name Dilemma, a late-Latin neuter noun derived from the Greek. Of the previously named species in the genus, inexpectatum is a neuter gender epithet, frumarkernorum has a genitive ending that is not to be influenced by the gender of the genus, and spectralis is an epithet originally used by Leal (2008) as a noun in apposition, DISCUSSION The inclusion of the new species in the genus Dilemma is well supported by the presence of the diagnostic shell characters of the genus: ante roposte riorly compresse od shell, cardioid outline in articulated valves, shi arp carina dividing the shell into the anterior and posterior regions, Dilemma japonicum new species. 5-6. Detail of surface sculpture. Anterior slope. 6. Posterior slope. 7. Detail of 7-8. Paratype 2, UMUT RM29691. Abbreviations: et = cardinal-like cardinal-like tooth and socket in each valve, and nacreous shell interior. Obviously there is no other group having a combination of these characters in bivalves. From the three previously known species of the genus, Dilemma inexpectatum (Crozier, 1966) from northern New Zealand is most similar to the new species in having rough commarginal sculpture in the posterior region, However, the two species differ in two characters: the rows of pores are absent and the escutcheon is distinct in D. inexpectatum. As summarized in Table 1, the four species in the genus can be distinguished among them- selves by four he Il characters: sculpture in the posterior region; the presence or absence of small pits; the dis- tinctness of the escutcheon; and the presence or absence of the posterior projection of the hinge. The paired pos- terior projections are present only in D. japonicum, they are not part of the hinge teeth syste m, because they do not articulate as a tooth and a socket between the right and left valves. Untortunately, the outlines of muscle scars were unclear in the spe ales of the new species. The original description of risa potas “Shell apparently nacreous internally Leal, 2008: : but did not offer any detailed dese ios or se T. Sasaki and J.H. Leal, 2008 Page 169 Figures 9-12. Dilemma japonicum new species. Shell microstructure. 9-10. Observed at small broken part of shell margin, vertical section of homogenous structure in outer layer (OL) lined by nacreous structure in inner layer (IL). 11-12. Oblique (11) and horizontal (12) views of nacreous layer at inner shell surface. Paratype 2, UMUT RM29691. on the shell microstructure of the three originally in- cluded species. The description of the inner nacreous shell layer in the present study confirms its presence in Dilemma. A shell consisting of outer homogeneous and inner nacreous layers (Figure 9-12) supports inclusion of the new species in Anomalodesmata (Taylor et al., 1973; Prezant, 1998). The microstructure of the outer layer is somewhat similar to a simple prismatic structure in that elongate granules are arranged vertically. The outer layer is identified as a homogeneous structure, because, unlike typical prismatic structure, crystals lack sharp bound- aries. It should be observed, however, that environmen- tal changes can alter otherwise organized shell micro- structures to appear irregular, smoothed, and homoge- neous (R. Prezant, pers. comm.). Table 1. Shell characters and distribution of the four Dilemma species. D. frumarkernorum D. spectralis D. japonicum D. inexpectatum new species Sculpture in posterior Dense, coarse growth Smooth region lines Small pits on exterior Absent Present surface Escutcheon Not clearly separated Projection posterior Absent Absent to beak Distribution Off Key West Florida, Off Vanuatu, USA, 229 m Not clearly separated Southwestern Pacific, 950-961 m Rough commarginal Rugose. lamellate ribs and grooves Absent commarginal ribs Present Clearly defined Not clearly separated Absent Present Off Three Kings Off central Japan, Islands, New Zealand, Northwest Pacific. $05 m 240-273 m (dead Page 170 THE NAUTILUS, Vol. 122, No. 3 The prodissoconch of Dilemma japonicum new species indicates non-planktotrophic, lecithotrophic develop- ment. It measures ca. 200 jm in length (Figure §), a size that fits into the predicted size range (135-230 xm) of prodissoconchs of bivalves with lecithotrophic develop- ment (Ockelmann, 1965; Jablonski and Lutz, 1983). The developmental modes of the other species of the genus are unknown. Although two of the other live-collected species of Dilemma were found attached to hard substrata, the mi- crohabitat of the new species is unknown. The bottom sediment from the type locality contained numerous dead shells and other biogenic fragments such as those of bryozoans and sponges. Because most bivalves were dead and disarticulated, large part of samples in the dredge haul was inferred to be transported and soe cunuilered. Specifying the actual microhabitat of the species is an interesting target for future sampling. The description of Dilemma japonicum extends the geographic range of the genus to the Northwest Pacific, in addition to previous records from off Florida and the Southwestern Pacific (off Vanuatu and New Zealand). The genus has a broad geographic range, extending across the Panama land bridge and Eastern Pacific we rier. Future sampling of deep- sea hard substrates may yield additional records of Dilemma species from other locations in the Pacific and in other oceans. ACKNOWLEDGMENTS The authors are indebted to the reviewers, Riidiger Bieler, Robert S. Prezant, and Richard C. Willan, for their comments and suggestions. Richard Willan and Rii- diger Bieler helped with etymologies and confirmed the gender of the name Dilemma. The samples of the new species became available with the kind assistance of Dr. Suguru Ohta (formerly Ocean Research Institute, Uni- versity of Tokyo), other colleagues on board and crew of R/V TANSEI-MARU ) (Japan Agency for Marine-Earth Sci- ence and Technology, formerly Ocean Research Insti- tute, University of Tokyo) during the cruise KT-00-05. LITERATURE CITED Crozier, M. A. 1966. New species and records of Mollusca from off Three Kings Islands, New Zealand. Transactions of the Royal Society of New Zealand, Zoology 8: 39-49. Farmer, M. A, W. K. Fitt, and R. K. Trench. 2001. Morphology of the symbiosis between Corculum cardissa (Mollusca: Bivalvia) and Symbiodinium corculorum (Dinophyceae). Biological Bulletin 200: 336-343. Geiger, D. L., B. A. Marshall, W. F. Ponder, T. Sasaki, and A. Warén. 2007. Techniques for collecting, handling, and propane small molluscan specimens. Molluscan Re- search 27(1): 1-49 Jablonski, D. and R. A. Lutz. 1983. Larval ecology of marine benthic invertebrates: paleobiological implications. Bio- — Reviews 58; 21-89, Leal, J. H. 2008. A remarkable new genus of carnivorous, A bivalves (Mollusca: Anomalodesniats: Poromyidae) with descriptions of two new species. Zootaxa 1764: 1-18. Ockelmann, W. K. 1965. Developmental types in marine bi- valves and their distribution along the Atlantic coast of Europe. In: Cox, L. R. and Peake, J. F. (eds.) Proceedings of the First European Malacological Congress, London, 1962, Conchological Society of Great Britain and Ireland, and the Malacological Society of London, London, pp- 25-35. ; Prezant, R. S. 1998. Subclass Anomalodesmata. In: Beasley, P. L., Ross, G. J. B. and Wills, E. (eds.) Mollusca: The Southern Synthesis. Fauna of Australia. Vol. 5, Part A. SCIRO Publishing, Melboume, pp. 397-249. Taylor, J. D., W. D. Kennedy, and A. Hall. 1973. The shell structure and mineralogy of the Bivalvia. II. Lucinacea- Clavagellacea, Conclusions. Bulletin of the British Mu- seum (Natural History), Zoology 22: 255-294, THE NAUTILUS 122(3):171-177, 2008 Page 17] On the genus Heteroschismoides Ludbrook, 1960 (Scaphopoda: Gadilida: Entalinidae), with descriptions of two new species Victor Scarabino Département Systématique et E volution Muséum national d'Histoire naturelle UMS Taxonomie-Collections CP51 55 rue de Buffon 75005 Paris, FRANCE scarabino_victor@yahoo. fr Janeiro Av. Pasteur, 455 Carlos Henrique Soares Caetano Departamento de Zoologia, Universidade Federal do Estado do Rio de Rio de Janeiro, 22.290-240 BRAZIL cheaetano@zipmail.com.br ABSTRACT Heteroschismoides is a deep-sea genus characterized by shells being between 10 and 20 mm as adults, sculptured by 9 to 10 prominent primary ribs and a unique deep irregular apical fis- sure on dorsal side, considered until now to include a single species: Dentalium subterfissum Jeffreys, 1877. During revision of material from several expeditions carried out by the Museum National d'Histoire Naturelle, Paris, two new species were identified and are here described: H. meridionalis new species and H. antipodes new species In addition, designation of the lectotype of H. subterfissus is proposed, as well as new records for this species in the northeastern Atlantic ocean are given. Heteroschismoides meridionalis new species is closely related to H. subterfissus, but the first has a smaller maximum diameter of shell and smaller apical aperture diameter. Heteroschis- moides antipodes new species from Solornon Islands is smaller than other two species for both, shell length and fissure exten- sion. The distance of point of maximum curvature from the apex in H. antipodes new species is located nearer to the apex than in H. meridionalis new species and H. subterfissus. The results here obtained considerably enlarge the geographical distribution of the genus and suggest a worldwide bathyal and abyssal distribution for this genus. Additional Keywords: Heteroschismoides subterfissus, Het- eroschismoidinae, tusk shell, lectotype, new species, Brazil, Solomon Islands, deep-sea INTRODUCTION The genus Heteroschismoides was proposed by Lud- brook in 1960 to include the deep sea species De ntalium subterfissum Jettreys, 1877 (for further supraspecific in- formation see Steiner and Kabat, 2001). The most re markable character of the genus is a unique, deep and irregular apical fissure at the dazed! side of shell not present among other Scaphopoda (Chistikov, 1982; Scarabino, 1995). Since Chistikov (1982), this genus has been con- sidered as monospecific and restricted to the Atlantic Ocean. During the revision of material from several ex- peditions carried out by the Muséum national d'Histoire naturelle, Paris, in the northeastern and southwestern Atlantic Ocean (Brazil) as well as off Solomon Islands in the Pacific, a number of specimens corresponding to the genus were identified. In a preliminary sorting, Brazilian specimens were placed under H. subterfissus, but the unexpected finding of representatives in the tropical Pa- cific, decided us to undertake the present revision. Here, we propose the lectotype of Heteroschismoides subter- fissus and describe two new species, extending consider- ably the geographical distribution range of the genus. MATERIALS AND METHODS The material was collected during several expeditions car- ried out by the MNHIN and IrREMER, known as Brocas I to XI (1972-1981), THALASSA (1970-73) and INCAL (1973): in the Gulf of Gascony; ABYPLAINE (1982): off Portugal and Spain; NORATLANTE (1969): large coverage of the North Atlantic; BIACORES (1971): off Azores; BIOVEMA (1977): VeMA Trench; MD55 (1987): Southeastern Bra- zil: and SALOMON 2 (2004): off Solomon islands. In view to better define the species, a morphometric analysis was attempted based on selected undamaged shells of the three species. Shell measurements were taken according to Shimek (1989), Steiner (1999) and Steiner and Linse (2000), including length (L); maximum diameter (Max), that is in this case equal to the anterior aperture diameter; maximum curvature (Arc); distance of point of maximum curvature from the apex (Larc); apical aperture diameter (Apd). Besides those measure- ments, we also estimated the extension of the apical fis- sure (Fiss) located at dorsal side of shell. To assure in- dependence among variables, we performed a prelimi- nary correlation analysis among all variables with strongly correlated variables (r > 0.70) be sing excluded. The non- parametric Kruskal-Wallis test was employed to examine for differences of each morphometric parameter (un- transformed data) between species. Dunn’s multiple comparisons test was used a posteriori to assess signifi cant differences between species. A multivariate approach aaa: 179 Page 172 THE NAUTILUS, Vol. 122, No. 3 was carried out utilizing Discriminant Function (DF) Analysis to integrate all morphometric data in a single analysis. To perform this analysis, we standardized the morphometric data following Romesburg (1984). Institutional abbreviations used in the text are; BMNH: The Natural History Museum, London; IFREMER: Insti- tut Francais de Recherche pour lExploitation de la Mer; MNR]J: Museu Nacional, Rio de Janeiro, Brazil; MNHN: Muséum national d’Histoire Naturelle, Paris; USNM: National Museum of Natural History, Smithsonian Insti- tution, Washington, DC, USA. Other abbreviations are: CP: beam trawl; DC: “Charcot” dredge; stn: station; lv: live-collected; dd: shell only. SYSTEMATICS Order Gadilida Starobogatov, 1982 Suborder Entalimorpha Steiner, 1992 Family Entalinidae Chistikov, 1979 Subfamily Heteroschismoidinae Chistikov, 1982 Genus Heteroschismoides Ludbrook, 1960 Type Species: Dentalium subterfissum Jeffreys, 1877 (by original designation). Recent, northeastern Atlantic Ocean. Description (Modified from Scarabino, 1995: 302): Shell up to 20 mm length, slightly curved, regularly ta- pering. Translucent grey when the animal is alive, opaque to polished dhe or cream when empty. Longi- tudinally sculptured by 9-10 primary ribs that reach a anterior aperture or vanish towards it. Intercostals spaces convex or straight, smooth or finely striated longitudi- nally by 8 to 14 lines. Secondary ribs, predominantly a single one, can be present in each intercostal space. Apex with a long and wide irregular fissure on dorsal side early observed in late embryonic stages and juvenile shells. Section polyg gonal, more notorious at the central zone of shell, fading or not towards the slightly laterally com- pressed fan oral aperture in adult specimens. Shell ee tures and repairs are frequently observed, as well < twisting of shell. RapuLa: Rachidian tooth with anterior margin rounded and lateral margins thick; laterals with sharp pointe ral prime ary a and 4 importe int denticles; mar- ginal slightly curved with conspicuous lateral processes, better observed in light microscope. Distribution: Recent, Atlantic Ocean and tropical Pa- cific Ocean, bathy al- -aby ssal. Remarks: Scarabino (1995: 302) mentioned by mis- take “10-12 primary ribs”; in the light of new information this number is here corrected as 9 to 10 primary ribs. Molecular data of H. subterfissus has been published by Steiner and Dreyer (2003). Heteroschismoides subterfissus (Jeffreys, 1877) (Figures 1-6, 15) Dentalium subterfissum Jettreys, 1877: 154; 1883: 660, pl. 49, fig. 3; Watson, 1879: 516; 1886: 10, pl. 1, fig. 10; Pilsbry and Sharp, 1897: 61, pl. 7, figs. 15-19; Warén 1980: 53. Dentalium (Heteroschisma) sibie wrfissum: Henderson, 1920; 58. Dentalium (Dentalium) subterfissum: Nickles 1979: 47, fig. 5a b. Heteroschismoides subterfissus: Chistikov, 1982: 675, figs. 3, 5; Steiner 1998: 78: Steiner and Kabat, 2001: 446; 2004: 651. Description: Shell up to 9.2 mm length, slightly curved, regularly tapering. Translucent shiny grey, pres- enting agtuies and repairs. Longitudinal sculpture of nine primary notorious ribs and ¢ a single secondary one in between, more notorious in the dorsal half, all reaching the thin oral aperture. Intercostals spaces conv on finely striated longitudinally by eight lines. Apex with a 2.3 mm long, wide, irregular fissure on dorsal side. Transversal section poly gonal, slightly compressed laterally, less ap- parent at apex. Type Material: — Lectotype, here selected (see remarks), the largest (9.2 mm) of the three specimens of lot USNM 175018, Ireland, 54°19’ N,11°50'’ W, 1180 fims [2158 m] (Porcupine 1869 stn 16); other paralectotypes: USNM 175017, Off West coast of Ireland, (Porcupine 1869 stn 19a), 2 specimens; USNM 175019, Greenland, 56°11'N 37°41'W, 1450 fms [2646 m], Valorous stn 12, 1 speci- men: USNM 175020, 55°40'N 12°46’W, 1476 fms [2694 m], Porcupine L569 stn 21, 3 specimens; off Azores, Challenger Expedition stn 78, 37°26! 95°13’ W, Azores, 1000 fathoms [1825 m], 7 specimens and 3 frag- ments, BMNH_ 1887.2.9.36-40. Two other paralecto- types from nae gee expedition are deposited at BMNH under code 1885.11.5.1393—4, with a mention “further syntypes in USNM 175017-020", The fact of the species has been partially described on the base of a young specimen (see Remarks) and that the specimen illustrate od was not found in the collections re- vised, lead us to designate the lectotype based on an adult shell. Measurements of Lectotype (mm): Length 9.20, oral aperture 1.00, apex 0.12, are 0.42 at 1.9 from apex. Type Locality: West of Ireland, 54°19'N, 11°50'W, 1489 m, Porcupine 1869, stn 16 (here selected). Other Material Examined: Off Gabon (Af S, 8°18’ E, stn DS20, 2514 m (1 dd); Gulf of Guinea, 4°40°N, 5°39’2E): stn DS30, 3109 m (S dd) (both WaLbDA Expe eee CNEXO, Nicklés, 1979), all MNHN. Yica), 2°32’ Figures 1-14. Heteroschismoides species. 1-6. H. subterfissus (Jeffreys, 1977). 1. Lectotype USNM 175018, 9.2 mm. 2. Para- lectotype. 3. Detail of the apical fissure. 4. Specimens showing embryonic shell. 10. H. meridionalis new species. microsculpture on the outer surface of intercostals space, 7- 5. Detail of longitudinal ribs. 6. Detail of 7. Holotype, MNHN, 14 mm. 8. Paratype, detail of apex. 9. Detail of sculpture. 10. Detail of microsculpture on the outer surface of intercostals space. 11-14. Heteroschismoides antipodes new species. 11. holotype, MNHN, 9.95 mm. 12. Detail of apex. 13. Detail of sculpture. 14. Detail of microsculpture on the outer surface of intercostal space, 2008 V. Searabino and C. H. S. Caetano, Page 174 THE NAUTILUS, Vol. 122, No. 3 1haem Figures 15-17. Radulae of Heteroschismoides species. 15. H. subterfissus, rachidian and lateral teeth, intemal view. 16. H antipodes new species, external view of lateral tooth (left), rachidian, lateral, and marginal teeth (right), in internal view. 17. H meridionalis new species: rachidian and lateral teeth, internal view. New Records: Central Atlantic Ocean: Azores Islands: 11°06’ W, 2466 m, 3 lv, 1 dd: stn CP 04, 56°33’ N, 11°11! BIACORES 1971: stn DS 54, 38°12’ N, 28°15’ W, 1S10 m, W, 2438-2513 m, 7 lv, 7 dd: stn CP 05, 56°00! N, 12°29’ 2 dd, stn DS 165, 37°33’ N, 25°58’ W, 2085-2050 m, 1 W, 2884 m, 1 dd: stn CP 06, 55°02.3' é. 12°40’ W, dd, DS 173, 37°57' N, 26°08’ W, 3225 m, 1 dd: Norar- 2888-2893 m, 3 lv, 1 dd: stn CP 07, 55°03’ N, 12°46’ LANTE 1969: stn 02, 53°55’ N, 17°52’ W, 2456 — 9420, 3 2895-2897 m, 1 dd; stn CP 09, 50°15’ N, 13°16’ lv; stn 84, 36°21' N, 08°43’ W, 2871-2875 m, 3 lv, 3 dd: 2659-2691 m, 2 lv; stn WS OL, 50°19" N, 13°08’ . stn 85, 36°25’ N, 08°48’, 257: 3-2580 m, 2 ie 3. dd; 2550-2539 m, 5 lv; stn WS 02, 50°19’ N, 12°55' W, THALASSA ee Sth X 336, 44°11’ N,O5°10' W, 1850— pees m, | lv: stn WS 03, 48°19’ N, 15°23’ W, 4829 2050 m, 1 Iv, 1 dd; Eastern Atlantic Ose an: INCAL 1976: m, 2 dd: stn WS 09, 47°28.8 N-47°27.9’ N, 09° 234! W, stn. DS *" 57°59’ N, 10°40’ W, 2091 m, 56 lv, 7 dd: stn. 4277 m, 1 dd,; stn OS 01, 54°14’ N, 13°11’ W, 2634 m, DS 02, 57°59’ N, 10°49’ W, 2081 m, 51 lv, 3 dd: stn. DS L lv; Brocas I: stn DS 32, 47° 32" N. 08°05’ W, 2138 m, 05, ee N, 11°12’ W, 2503 m, 24 lv, 7 dd; stn. DS 07, 3 lv, 3 dd; Bocas HI: stn DS 37, 47°32’ N, 08°35’ W, 55°01" 12°31’ W, 2884 m, 4 lv, 1 dd; stn. DS 07, 2110 m, 2 dd; Brocas IV: stn DS 52. 44°06' N, 04°22' W, 56°27’ : 11°11’ W, 2884 m, 5 lv, 1 dd: stn. DS 08, 2006 m, 2 dd: stn DS 51, 44°11’ N, 04°15’ W, 2430 m, 55°02’ N, 12°35’ W, 2891-2884 m, 3 lv, 2 dd: stn. DS 09, L lv; stn DS 58, 47°34" N, 09°08’ a 2775 m, 3 dd: stn 55°08’ N, 12°53’ W, 2897 m, 2 lv, 2 dd; stn. CP O1, DS 62, 47°33’ N, 08°40' W, 2175m, 1 dd: stn DS 64, 57°58’ N, 10°55’ W, 2040-2068 m, 12 lv; stn. CP 02, 47°29’ 'N, 08°35' W, 2156m, 4 lv, 1 dd; BIOGAS V: stn 57°58’ N, 10°42’ W, 2091 m, 2 dd: stn. CP 03, 56°38' N, CP 07, 44°09.8' N, 04°16.4' W, 2170 m, 5 dd; Brocas VI stn DS 61, 47°34.7' N, 08°38.8" W, 2250 m, 1 lv, 1 dd; 6 ee ee stn DS 62, 47°33’ N, 08°40' W, 2175 m, 1 dd; stn DS 63, 47°33' N, 08°35’ W, 2126 m, 1 lv; stn CP 08, 47°03’ N, >| | 08°39' W, 2177 m, 2 lv, 1 dd; stn CP 09, 47°33’ N, 08°44’ a} ° © Hsubterfissus (Jetheys, 1877) | W, 2171 m, 1 lv: stn CP 1 44°0S' N, 04°16’ W, 1995 m, | 3 dd; stn DS 71, 47°34’ N, 08°34’ W, 2194 m, 3 lv, 5 dd: 3 a? stn DS 86, 44°04.8' N, 04°18.7' W, 1950 m, 1 dd; stn DS 2 S7, 44°05’ N, 04°19’ W, 1913 m_3 lv: Blocas VIL: stn CP x | Ps 26, 47°33" N, 08°34" W, 2115 m, 1 lv; Biocas VII: stn e . 5 * KG 145, 47°33’ N, 08° ea W, 2170 m, 1 dd: stn KG 147, a| *% , eo. oe 17°33.40' N, “08°40.70" W, 2190 m, Iv, dd; stn KG. 149. : 6 ° 3° : 17°33’ N, 08°39' W, 2165 m, 1 dd: stn KG 151, 47°34’ a 4° " : N, 08°39' W, 2205 m, 1 lv; BloGas IX: stn CP 34, 47°32' 2 a® N, 08°25’ W, 1970 m, 1 dd; BioGas XI: stn CP 37, 47°34’ ° N, 08°41' W, 2175 m, 2 Wy. 4 3 2 1 0 1 2 3 4 5 Distribution: — Eastern Atlantic Ocean: Greenland, off DF1 West Ireland, Gulf of Gascony, off Spain and Portugal to Figure 18. Discriminant function analysis of shell morpho Gulf of Guinea, West Africa (Je ffreys, 1877; Watson, netric parameters of Heteroschismoides species 1S79. ISS6. Loc: ard. LS9S: Nickles. 1979) _ Alive in 940 V. Searabino and C. H. S. Caetano, 2008 2987 m, shells down to 4529 m (present paper). Richest depth tor live material: 2040-2503 m (present paper). Remarks: _ Jeffreys partially described the - cies based on a juvenile specimen, since he mentioned ~ end bulbous” and illustrated this morphology in plate 49, fig. 3. However, no juvenile specimen was detected among the type material. In addition, he wrote that the slit is in the “under or ventral side”, which in fact corre- sponds to the dorsal side in normal position of the ani- mal. Jeffreys also mentioned the presence of 12 to 16 ribs, but we did not find any specimens with that number of ribs. However, it is possible that secondary ribs and intercostal striae might have been, in part, counted in- appropriately for the original description. To clarify, in the description we enumerated only the primary ribs because the number of secondary ribs varies with age and among specimens but the number primary ribs is constant among specimens and through the life of the specimen. Specime ns with embryonic shell are often ob- served. Heteroschismoides subterfissus was also mentioned in the CHALLENGER expedition from “stn 120 - 8°37’ S 34228’ W, Pernambuco, South America 675 fathoms {1232 m] red mud” (Watson, 1886). The lot in which this reference was based was not located in BMNH collec- tions by the senior author. In the distribution paragraph, Watson (1886) wrote “Habitat.- Davis Strait. 1785 fath- oms. North Atlantic; various Stations off the West Coast of Ireland. 1180 to 1476 fathoms (Jeff.).” He did not mention Pernambuco. The material from Pernambuco may belong to the new species described below or, as sugge sted by Henderson (1920: 58). could also correspond to specimens of Per- tusiconcha callithrix (Dall, 1889), since young specimens of this species, specially those with a heoken apex, can easily be misidentified as species of Heteroschismoides (Scarabino, pers. observ). Heteroschismoides meridionalis new species (Figures 7-10, 17) Description: Shell 14 mm length, slightly curved, regularly tapering, opaque white. Longitudinal sculpture composed of nine rounded-edge primary ribs, all reach- ing but fading towards the oral aperture. No secondary ribs. Intercostal spaces concave in posterior three quar- ters and straight to convex at the anterior fifth, present- ing 12 very faint, fine striae throughout. Apex with a 2 mm long, ‘wide irregular fissure on dorsal side. Cross- section polyg gonal, less anaes at ape x ‘and at the slightly Ricasue cients of the Holotype (mm): Length 14.0, oral aperture 1.0, apex 0.1, arc 0.8 at 6.7 from apex. Holotype (lv) MNHN 20902 and 14 Type Material: ' 13 MNHN 20903-20906, 1 MNRJ paratypes (dd), 12707. Type Locality: Off Espirito Santo, Brazil, 1$°59.1' S 37°47.8' W, 1540-1550 m {MD 55 stn DC 70]. . poste rior Material Examined: Southwestern ne Ocean: MD 55 stn CP 68, 18°55.6' S,37°49.1' W, 1200-1500 m, 1 dd (paratype MNHN 20903); stn OG 70, 18°59.1’ S, 37°47'8 W, 1540-1550 m, 1 lv (holotype MNHN 20902), 26 dd (7 paeype MNHN 20904, 1 paratype MNRJ 12707): stn DC 72, 19°00.4' S, 37°48.8S' W, 950-1050 m, 4 dd ( (paratypes MINHN 20905); stn CB 77, 19°40.6" S, ce W, 790-940 m, 9 dd (2 patie: MNHN 20906): stn. CB 79, 19°O1.S' S, 37°47.8' W, 1500-1575 m, 10 dd: sth CB 106, 23°54.2' S, 42°10.5' W, 830 m, 15 dd; stn. CB 107, 24°00.3' S, 42°14.4' W, 1020 m, 11 dd. Etymology: Meridionalis: Southern. Name refers to the taxon’s distribution in the Southern Hemisphere. Distribution: Brazil: Espirito Santo and Sao Paulo. Alive in 1540 m, shells in 790-1540 m depth. Heteroschismoides antipodes new species (Figures 11-14, 16) Description: Shell to 9.95 mm length, slightly curved at the apex, re gularly tapering. Opaque white. I songitu- dinal sculpture ron ten primary ribs, all reaching the “onl aperture. Intercostal spaces concave and smooth. Apex with a wide irregular fissure 1.7 mm long on dorsal side. Cross section polyg gonal, less obvious at apex, slightly lat- erally compressed at the anterior aperture. Measurements of Holotype (mm): Length 9.95, oral aperture 0.9, apex, 0.1, are 0.4 at 2.76 from apex. Type Material: Holotype (dd) MNHN 20907 and 10 paratypes (7 lv, 3 dd) MNHN 20908-20910 Type Locality: Solomon Islands, 07°49.3" S, 157°41.2' E, 1045-1118 m [SALOMON 2 Stn, CP2217]. Material Examined: Solomon Islands. SALOMON 2 stn CP2182, 08°47.0' S, 157°37.9’ E, 762-1060 m, 16 dad. stn CP2197, 08°24.4' S, 159°22.5' E, 897-1057 m, 2 lv, 2 dd (3 paratypes MNHN 20909); stn CP2217, 07°49.3' S, 157°41.2' E, 1045-1118 m, 1 ly, 3 dd (holotype MNHN 20907 and 3 paratypes MNHN 20908), stn CP2218, 07°56.3' S, 157°34.6' E, 582-864 m, 4 lv (paratypes MNHN 20910) 14 dd: stn CP2253, 7°26.5' S, 156°15.0’ E. 1200-1218 m. 10 dd. nia. From Greek antipodes, meaning those living on diametrally opposed places on Earth. Distribution: Solomon Islands, alive in 1200 m:; shells 762-1200 m depth. MORPHOMETRIC RESULTS AND DISCUSSION Table 1 lists the mean, standard deviation, minimum, and maximum values for each morphometric parameter for the three species. Significant differences in all mor- phometric parameters were observed between the spe- cies (Kruskal-Wallis test; p<0.01), except for the maxi- mum curvature (Arc). This univariate comparison showed that H. meridionalis new species and H. subter- Page 176 THE NAUTILUS, Vol. 122, No. 3 Table 1. Shell morphometric parameters with minimum (min), maximum (max), mean values and standard deviation (+SD) for the three new species of Heteroschismoides. L: length; Max: maximum diameter (= anterior aperture diameter); Arc: maximum curvature; Lare: distance of point of maximum curvature from the apex: Apd, apical aperture diameter; Fiss: fissure length. KW: Kruskal-Wallis test); ns: not significant; *p<0.001. The horizontal lines at Dunn’s test represents absence of statistical differences. H. meridionalis (n=12) H. antipodes (n=8) H. subterfissus (n=23) mean (+SD) min—max mean (+SD) min-max mean (+SD) min—max KW Dunn's test L 13.3 (1.6) 11.1-16.5 8.9 (0.8) 7.9-10.7 14.2 (2.6) 10.0-19.0 19.19° st sb an Max 1.0 (0.05) 0.9-1.1 1.0 (0.07) 1,0-1.2 4 (0.16) 1.0-1.7 29.76° st an sh Arc 0.5 (0.4) 0.1-1.2 0.5 (0.1) 0.4-0.7 0.5 (0.1) 0.2-1.0 0.82 ns st sb an Lare 6.1 (0.8) 4.8-7.2 3.5 (0.6) 2.14.0 5.4 (2.5) 2.2-11.4 14.98° st sb an Apd 0.1 (0.05) 0.1-0.2 0.2 (0.00) 0.2-0.2 0.2 (0.05) 0.2-0.3 20:10" st an os Fiss 2.1 (0.5) 1.5-3.1 1.4 (0.2) 11-15 1.9 (0.5) 1.2-3.0 13.32° st ban fissus are more similar, differing significantly only in maximum diameter (Max) and apical aperture diameter (Apd) (Table 1). Heteroschismoides meridionalis new species is also less shiny and the intercostal spaces are more densely striated: 12-15 striae (observed on stained or coated specimens due to the difficulty to be observed without any treatment) against S—10 in H. subte -rfissus (well observed under inert these data was obtained from counting 10 specimens of each species). Individuals of Heteroschismoides antipodes new spe- cies are significantly smaller than individuals of the other two species for both shell le ngth and fissure extension. The distance of point of maximum curvature from apex (Lare) in H. antipodes new species specimens is located nearer to apex than in specimens of both H. meridionalis new species and H. subterfissus (Table 1). In addition, H. antipodes specimens are opaque, due to their coarsest surface microsculpture. Specimens of this species have smooth intercostal spaces without secondary ribs and striae, and the primary ribs are more apparent. The multivariate discriminant function analysis was able to distinguish three groups among the specimens we examined that correspond to three species (Wilks’ Lambda = 0.12; F676 = 24.27; p < 0.0000). This analysis classifies about 97% of the cases correctly (only 1 out of 43 specimens were incorrectly classified) (Figure 3). The model was constructed with 3 parameters (Max, L : Max, Lare). The variable L, which was highly correlated with Lare (r = 0.80) and Fiss (r = 0.74), was excluded. The Discriminant Functions (DF) based on the raw coeffi- cients of canonical variables are shown below: DF] = 2.5439Max + 1.0378L : Max — 0.7366Lare DF2 = 0.7249Max + 2.1404L : Max — 0.832S8Lare The radulae of the three species, here illustrated, are quite similar and further studies would be necessary to test the existence of interspecific differences, as it is the case in most Scaphopoda. Finally, the findings of Het- eroschismoides in the Solomon Islands show the genus to have worldwide distributed instead of being confined to the eastern Atlantic Ocean. However, better sampling of areas of the Indian Ocean (one of the ie »ss known tropical areas for scaphopod diversity) could eventually confirm whether the genus is also represented in that ocean. ACKNOWLEDGMENTS The authors want acknowledge the support and help re- ceived from Philippe Bouchet, Virginie Héros, and Phil- ippe Maestrati (MNHN), Ellen Strong (USNM), Kathie Way (BMNH), and Anders Warén (Swedish Museum of Natural History, Stockholm). Ronald Shimek is also gratefully acknowledged for his constructive comments on the manuscript. LITERATURE CITED Chistikov, S. D. 1952 The modem Entalinidae (Scaphopoda, Gadilida), 1. Subfamily Heteroschismoidinae - 1 [in Rus- sian]. Zoologicheskii Zhurnal 61 (5): 671-682. Dall, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-S0), by the U.S. Coast Survey Steamer “Blake”, Lieut.-Commander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. XXIX. Report on the Mollusca. Part 2, Gastropoda and Scaphopoda. Bulletin of the Museum of Comparative Zodlogy at Harvard College 1S: 1-492, pls. 1O—40. Jeffreys, J. G. 1877. New and peculiar Mollusca of the order Solenoconchia procured in the “Valorous” expedition. An- nals and Magazine of Natural History (4) 19: 153-158. Jeffreys, J. G. 1883. ie the Mollusca procured during the “Lighting” and “Porcupine * expeditions, 1868-70. Pro- ceedings of the Zoological Society of London 1882: 656— 687, pl. 49-50 V. Searabino and C. H. S. Caetano, 2008 Page 177 Henderson, J. B. 1920. A monograph of the East American scaphopod mollusks. United States National Museum Bul- letin 111: 1-177, 20 pls. Locard, A. 1898S. Mollusques Testacés. In: Expéditions Scien- tifiques du Travailleur et du Talisman pendent les années LSSO-1SS3. Masson, Paris, 515 pp., 1S pls. Nicklés, M. 1979. Scaphopodes de /Ouest-Africain (Mollusca, Scaphopoda). Bulletin du’ Muséum National d’ Histoire Naturelle, Section A. Zoologie, Biologie et écologie Ani- males (séries 4) 1(1): 41-77, 1 pl. Pilsbry, H. A. and B. Sharp. ee Scaphopoda, 1-280, in: Manual of C Jonchology, 17: 1-280, Conchological Section, Academy of Natural Sciences, Philade Iphia. 1 S97: 1-144, pls. 1- 26; 1S9S: i-xxxii + 145-280, pls. 27-37]. Romesburg, H. C. 1984. Cluster Analysis for Researchers. Life- time Le sarning Publications, Be ‘mont, 334 pp. Scarabino, V. 1995. Scaphopoda of the tropical Pacific and Indian Oceans, with descriptions of 3 new genera and 42 new species. In: Bouchet, P. (ed.), Résultats des Cam- pagnes MUSORSTOM, vol. 14. Mé se s du Muséum na- tional d'Histoire naturelle 167: 189-37 Shimek, R. L. 1989. Shells eee ae aa systematics: a review ot the slender, shallow-water Cadulus of the North- eastern Pacific (Scaphopoda: Gadilida). The Veliger, 32, 233-246, Steiner, G. 1997. Scaphopoda from the Spanish coasts. Iberus 15: 95-111. Steiner, G. 1998. Phylogeny of Scaphopoda (Mollusca) in the light of new anatomical data on the Gadilinidae and some eae matica, and a reply to Reynolds. Zoologica Scripta 97: 73-82. Steiner, CG. 1999. A new genus and species of the family An- ulidentaliidae (Sez iphopoda: Dentaliida) and its systematic ae Journal of Molluscan Studies 65: 151-161. Steiner, G. and K. Linse. 2000. Systematics and distribution of the ey (Mollusca) in the Beagle Chanel (Chile). Mitteilungen aus dem ee Zoologischen Mu- — und Institut 97; 13—3¢ Steiner, G. and H. Dreyer. ae Molecular phylogeny of Scaphopoda ( (Mollusea) inferred from 18S rDNA se- quences: support for a Scaphopoda-Cephalopoda clade. ZA yes Scripta 32: 343-356, Steiner, G. and A. R. Kabat. 2001. Catalogue of Se ei taxa of Scaphopoda (Mollusca). Zoosystema 23: 433-460 Steiner, G. and A. R. Kabat. 2004. Catz alogue of spe cies-group names of Recent and fossil Scaphopoda ( (Mollusca). Zoo- systema 26; 549-726 Warén, A. 1980. Marine Mollusca described by John Gwyn Jeffreys, with the location of the type material. Concho- logical Society of Gre . Britain and Ireland, Special Pub- lesion 1. 1-60, pls. 1-8. Watson, R. B. 1879. vialinecs of the H.M.S. CHALLENGER Ex- pedition. H. The Solenoconchia, comprising the genera Dentalium, Siphodentalium, and Cadulus. Journal of the Linnean Society of London 14 (78): 506-529. Watson, R. B. 1886. Report on the Scaphopoda and Gaster- opoda collected by H.M.S. Challenger during the years 1873-76. Challenger Report, Zoology 15, pt. 42, 756 pp. 50 pls. THE NAUTILUS 122(3):178-181, 2008 Page 178 Parasitism of Monogamus minibulla (Olsson and McGinty, 1958) (Gastropoda: Eulimidae) on the red sea-urchin Echinometra lucunter (Linnaeus, 1758) (Echinodermata: Echinometridae) on the Caribbean coast of Mexico Norma Emilia Gonzalez-Vallejo Depto. Ecologia Acuatica El Colegio de la Frontera Sur Apdo. Postal 424, 77014 Chetumal, Quintana Roo, MEXICO negonzale@ecosur.mx ABSTRACT Eulimids are gastropod mollusks parasites of echinoderms. In- tertidal red sea-urchins, Echinometra lucunter, collected along the Mexican Caribbean shores were examined for the presence of those ectoparasitic gastropods. The analysis of 206 sea- urchins resulted in 56 having eulimids on them (27% preva- lence), mostly living in pairs, with about 7 eulimid pairs per sea-urchin. The eulimid ¢ gastropods found are Monogamus minibulla (Olsson and MacGinty, 1958), desonbed from Panama. Its host was unknown. This article consists of the first record of this host-parasite relationship. A short description of the shell structure and some details of the biological relation- ship are given. The females are much larger than the males, and the latter are usually attached to the female bodies. Most female eulimids live inside the hypertrophied foot, which this allows for partial oe of the entire body. A few females, however, were found living fixed directly on the echinoderm integument. The position of the g gastropod on the sea-urchin is fixe d via boring through the skeleton. Anchoring is achieved via the proboscis. Some ‘females had egg capsules ( (0.9-1.5 mm) fixed to their bodies; each is a spherical structure attached to the body by a short stalk. Egg capsules contained 14-15 em- bryos in different stages of development. The only previously publishec d account of this type of relationship inv olves a similar species een on Echinometra mathaei (Blainville, 1825) from the Red Sea. Additional Keywords: Rosenia, Echinometra mathaei, Monoga- mus interspinea INTRODUCTION The red sea-urchin Echinometra lucunter (Linnaeus, 1758) is frequently found on shallow water rocky- bottoms along the Me xican Caribbean shores, and occa- sionally in sea-grass beds. In that region, some specimens living in the reef lagoons host one or more tiny whitish e alorud gastropods firmly attached to their bodie 3S. Ecological associations including eulimids gastropods have been recognized (Warén, 1983). The pee de- scription of the eulimid Rosenia minibulla Olsson and McGinty, 1958, was made from an empty shell found on the beach of Isla Bocas, Panama; however at that time, its host was unknown. Liitzen (1976) studied the sea urchins Echinometra mathaei (Blainville, 1825) from the Red Sea and the Western Pacific Ocean and found eulimid parasites on them. He named a new genus as Monogamus because he found these eulimids always living in pairs; he included two different species, one (M. entopodia Liitzen, 1976) living inside the hypertrophied tube feet of E. matthaei from the Red Sea, and another one (M. interspinea Liitzen, 1976) living on the integument of E. matthaei from several localities in the Western Pacific Ocean. Warén (1983) established Rosenia Schepman, 1914, as a junior synonym of Pelseneeria Koehler and Vaney, 1908. However, the transfer of Rosenia minibulla to Mo- nogamus was informally proposed by Warén and Mooleal yeek (1989), perhaps because both share internal growth lines in the body whorl. In addition, the former euthor had found individuals of the species living on E. lucunter. The faunal associations of the red sea-urchin were a in the southern Caribbean Sea by Schoppe and Werding (1996), but these authors did not find eulimids ad with the red sea-urchin in the study localities. In this contribution, my objectives are to report the para- sitism of the eulimid Monogamus minibulla (Olsson and McGinty, 1958), on the red sea-urchin Echinometra lu- cunter (Linnaeus, 1758), found along the Caribbean coast of Mexico, and to describe some aspects of the relationship between the two species. MATERIALS AND METHODS The material was collected along several places in the Caribbean coast of Quintana Roo, Mexico: Isla Contoy N. E. Gonzalez-Vallejo, 2008 Page 179 (21°30'S.4” N, 86°47'45.3” 28/11/01); Playa Paraiso, C ne (20°35'21.5" N, Revie W, 24/111/01); Playa Ana and José (19°54'22.4" N, 87°26'14.5" W, 2 kin S from the junction to Boca Paila, 18/II/01); Majagual (18°43'/28” N, 87°42'05” W, 22/11/00): Punta Herradura (18°32'23” N, 87°44'32"” W, 27/VI/00): and Buenavista (18°30'42” N, 87°45'30" W, 15/V/99). Sea-urchins were eaunted and those with eulimids fixed, sorted and kept individually, and identified accord- ing to Hendler et al. (1995). The number of eulimids per sea-urchin was counted to determine the prevalence: the relationship between the number of infested hosts (sea- urchins) and the total number of hosts; and the intensity: number of parasites (eulimids) present in infected host (Margolis et al. 1952). Voucher specimens were depos- ited in the Swedish Museum of Natural History, Stock- holm, Sweden, SMNH 27858, and in the Reference Col- lection of El Colegio de la Frontera Sur (ECOSUR), Chetumal, Mexico). RESULTS Monogamus minibulla (Olsson and McGinty, 1958) Description: Mean shell length: 1.85 mm (range 1.5- 2.5), width 1.15 mm (range 1.0-1.5). Females are easily distinguished due to their body-size, being twice as large as males. The shell is tiny, spherical, fragile, and trans- parent. The spire is low, the nuclear apex is long and erect like a stump (mucro). The shell color is white with glossy texture, with a very fine suture line. There are two post- -nuclear whorls: the first one is small and convex with a small shoulder close to the suture, which is be step r seen on the aperture side. The body whorl is large, and inflated, polished, with some delicate growth lines. The aperture is wide, the outer lip thin, expanded along its median part. There is no operculum. When the animal is alive, the mantle and visceral mass are deep-red. There are two distinct black eyes. Tentacles were not observed. Parasitism: From a total of 206 sea-urchins, 56 were infested with eulimids. The prevalence was of 0.27 (pres- ence of the parasite in 27% of the specimens). The in- tensity was of 7.03: there was an average of 7.03 eulimid pairs per sea-urchin (including only hose that were in- fested by at least one eulimid [T TT able 1]). Attachment Locations of Parasites on Hosts: Many parasites have a preference for the oral side of the host, although some were located on the dorsal side (periproct); in only two cases parasites were found on the peristomial membrane (buccal tube foot). Most eulimids invade the tube feet, becoming inserted in its integu- ment. This invasion can produce a hypertrophy of that tube foot. Often, the deformation is such that when the eulimid grows into adulthood, the tube feet are modified beyond recognition. The eulimids, in sal ae juveniles with a single teleoconch whorl, use the skin of the tube as a protective hood; later, as the animal grows, its own mantle (the pseudopallium) covers the fragile shell, ex- cept for the apical part, which is left uncovered (Fig- ure 1). Another attachement site is the interambulacral zone. Eulimids that are not covered by the pseudopallium are attached by the snout (plus pharyngeal anchor). Few oth- ers may be found adhering to the integument at the base of spines, where according to stage of shell growth, it may be collapsed, broken, and later disappear, leaving an open space where the “eulimid-scar” becomes evident. Eulimids may bore the sea-urchin test: this may be seen clearly when detached specimens were placed on a host. They may also enlarge the pore associated with the foot tube: a typical feeding pattern of ectoparasitic eu- limids involves insertion of the proboscis through the tube pore followed by suction of coelomic contents cat the sea-urchin. The gastropods were found mostly in pairs, but, less frequently, two or three males were found associated with a single larger female. These pairs look like irregu- larly shaped protrusions of the body wall, each with one or two circular openings through which the eulimid shell tips may project. Egg capsules were observed packed in globular sacks (3-5), attached for a short stalk to the body of the female, and partially or completely covered by the pseudopallium. Each capsule (size 0.09-1.5 mm in length) contained 14-15 embryos. Egg capsules con- tained all stages from early egg stage to small juveniles ready to hatch, DISCUSSION Olsson and McGinty (1958) described Monogamus minibulla based on an empty shell. In the present study details of the shell morphology are presented based on several alive spe cimens an ‘those fixed on the sea- urchins. Warén in Warén and Moolenbeek (1989) briefly pointed out the presence of M. minibulla associated to Table 1. Intensity and prevalence of Monogamus minibulla on Echinometra lucunter on the Caribbean coast of Mexico. Localities Sea urchins With parasites Eulimid females Prevalence Intensity Isla C Contoy 27 13 P. Paraiso 3 ] P. Ana y José 51 14 Majagual 120 23 P. Herradura 3 3 2 2 Buenavista 268 20.61 0.45 ] 1.00 0.30 114 8.14 0.27 9 0.39 0.19 2 0.66 1.00 ] 5.50 1.00 Page 150 THE NAUTILUS, Vol. 122, No. 3 Figures 1-4. 3. Male (left, Echinometra lucunter. However, they only recorded the dimensions of the shell (1.5 mm of longitude and a larval shell of 1.0 mm), because their contribution was focused mainly in the relationship of the eulimid Trochostilifer eucidaricola (Warén and Moolenbeek, 1989) associated with the pencil sea-urchin Eucidaris tribuloides (La- marck, 1816). This study constitutes the first report of the relationship of Monogamus minibulla with Echi- nometra lucunter for the Mexican Caribbean region. Liitzen (1976) reported the presence of Monogamus entopodia in the tube feet of the Red Sea sea-urchin. In the Mexican Caribbean, M. minibulla infests the tube feet, but was also observed attached to the epidermis host, using its snout and tissues of the host but without being covered or protected with the pseudopallium. The two different patterns were rarely observed on the same sea-urchin. Liitzen (1976) had found the same pattems but on sea urchins collected in cifferent localities and regarded them as belonging to two different species Further, Echinometra from the Indian and western Pa- Vonogamus minibulla. 1. On Echinometra lucunter (host test = 16.54 min diameter 1.5 mm) and female (right, 2.5 mm). 4. Female, anterior and lateral views (1.5 mim height 8.72 mm height); 2. In situ. 1.4 mm width.) cific oceans have been found to belong to three different morphospecies that might be re productive ‘ly isolated from each other as we I ( Arakaki and Uehara, 1999). Thus, it seems that Caribbean sea-urchins have not di- verged as much as those present in the Indo-Pacific Oceans. Few studies have detailed the prevalence and intensity of eulimids parasites, Salazar and Reyes (1995) studie d the relationship of Thyca sn Berry, 1959, and the starfish Phataria unifascialis (Gray, 1840), finding that the association was more Foes in two sites of the Gulf of California where human influence was more accentu- ated. In this case, only one of the localities (Playa Ana and José) is he avily influenced by development, and is a protected, semi-enclosed embayment, with many resi- dences and tourists. That locality presented two sea- urchins that had unusually high infestations of about 100 eulimids each. Most of the sea-urchins were adjusted close to the average of about 7 eulimid pairs per sea- urchin. N. E. Gonzalez-Vallejo, 2008 Page 18] Warén (1983) pointed out that many tropical eulimids that attach their egg capsules to their shells to or to their hosts, have almost invariably been found with spawn, an indication that spawning might be continuous. On the other hand, Liitzen (1976) suggested for M. entopodia that the presence of different developmental stages found simultaneously in samples of egg capsules Sidi: cates continual production of eggs, which is similar to what happens in M. minibulla. A thorough study of the association is required to elu- cidate the mechanisms of feeding, including histology. There is need to clarify whether the proboscis sucks of part of the liquid fraction of the host’s coelom or, as Liitzen (1976) pointed out for M. entopodia, the eulimid feeds exclusively of the area adjacent to attachment on the sea-urchins. To determine the specificity of this association it is necessary to do experiments and observations in the col- lection place, to determine whether, as Warén (1983) pointed out, there is lack of specificity of the parasite on a single species guest. Although I had opportunity to examine some specimens of Echinometra viridis Agassiz, 1863, and I did not observe any attached eulimids, it could be said that if some preference exists in this case, but also this would have to corroborate it. Further, in 2001 T had examined about 50 specimens belonging to E. lucunter from Guana Island, but none had any associated eulimid parasite. ACKNOWLEDGMENTS Many thanks to Anders Warén (Swedish Museum of Natural History), who always answered my questions about eulimids and is always ready to help. To Sergio Salazar-Vallejo and Maria Ana Tovar, who reviewed pre- vious drafts of the manuscript and to colleagues at ECOSUR for collecting the sea urchins for the present study. The careful reviews by Anders Warén and an anonymous referee enhanced the clarity of this contri- bution. Humberto Bahena helped improve the quality of the photographs. LITERATURE CITED Arakaki, Y. and T. Uehara. 1999. Morphological comparison of black Echinometra individuals among those in the Indo- Ney Pacific. Zoological Science 16: 551-558. Hendler, G., J. E. Miller, D. L. Pawson, and P. M. Kier. 1995. Sea vee Sea-urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Wash- ington, 390 pp. Liitzen, J. 1976. On a new genus and two new species of proso- branchia (Mollusca) parasitic on the tropical sea urchin Echinometra mathaci. Israel Journal of Zoology 25: 38-51. Margolis, L., G. W. Esch, J. C. Holmes, A. M. Kuris, and A. Schad. 1982. The use of ‘ecological terms in parasitology (report of an ad hoc Committee of the American Society of Parasitologists). Journal Parasitology 68: 131-133. Olsson, A. A. and T. L. McGinty. 1958S. Recent marine mollusks from the Caribbean coast of Panama with the description of some new genera and species. Bulletins of American Paleontology 39(177): 1-58. Salazar, A. and H. Reyes. 1998. Parasitismo de Thyca callista (Gastropoda: Capulidae) ) sobre Phataria unifascialis (As- teroidea: Ophidiasteridae) en el Golfo de California, México. Revista de Biologia Tropical 46(3): 1-4. Schoppe, A. and B. Werding. 1996. The boreholes of the sea urchin genus Echinometra (Echinodermata: Echinoidea: Echinometridae) as a microhabitat in tropical South America. Publicazioni della Stazione Zoologica di Napoli I, Marine oe 17: 181-186, Warén, A. 1983. A generic revision of the family Eulimidae (Gastropoda:Prosobranchia) The Journal of Molluscan Studies, Supplement 13, 96 pp. Warén, A. and R. Moolenbeek. 1989. A new eulimid gastropod, Trochostilifer eucidaricola, parasitic on the pencil urchin Eucidaris tribuloides trom the southern Caribbean. Pro- ceedings of the Biological Society of Washington 102: 169-175. THE NAUTILUS 122(3):182, 2008 Page 182 Erratum Due to an editorial lapse, the article by Simone and Cunha (2008) in the most recent issue of The Nautilus incorrectly listed “FMNH” instead of the correct form “UF” as the collection acronym for the Florida Museum of Natural History. LITERATURE CITED Simone, L. R. L. and C. M. Cunha. 2008. Revision of the genus Spinosipella (Bivalvia: Verticordiidae), with descriptions of two new species from Brazil. The Nautilus 122: 57-78. THE NAUTILUS 122(3):183-184, 2008 Page 153 Notices Subscriptions rates for The Nautilus will increase slightly for 2009. Individual and institutional subscriptions will change respectively to $50 and $85. Shipping costs will remain the same. These adjustments are necessary given increases in production, materials, and mailing costs. The Nautilus staff has been making a concerted effort to increase the quality and quantity of articles published in the journal. 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Mi ve ee A ; . totae) | foe de ee ie ity ‘ ; i , vy) i to ! t!. ne Th, ‘ { : | ite . . | a, i " i ‘ ‘ ° : - A . ™ A Ti A i ir ts by " 1 t 1 7 j - : \ F) Ht \s ms . . j i i a a i ~ ' sit | ! i i 7 i ’ ws) - vy | | yal a eae 1 i | ti : f | , H Ly J i tet | ‘ - i i : i ! vi i 7 aM | { v : ih i : Ro ; 7 d f ; i : \ i | i fi] Fi ! 1 tf / ny I q ti 7 ' : | | Y ! { ] a a : 7 | ' v - i ‘ ; : ; ; vt, | ne : Pipe ae) | ' , Sse aie eid } j i! t op tor i ee 1° | THE NAUTILUS Volume 122, Number 4 December 22, 2008 ISSN 0028-1344 A quarterly devoted to malacology. EDITOR-IN-CHIEF Dr. José H. Leal The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 BUSINESS MANAGER Mary Jo Bunnell The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 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 Biologie des Invertébrés Marins et Malacologie Muséum National d'Histoire Naturelle 55, rue Buffon Paris, 75005 France Dr. Robert H. Cowie Center for Conservation Research and Training University of Hawaii 3050 Maile Way, Gilmore 409 Honolulu, HI 96822 Dr. Robert T. Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. Eileen H. Jokinen §234 IE. North Shore Road Sault Ste. Marie, MI 49783 Dr. Douglas S. Jones Florida Museum of Natural History University of Florida Gainesville, FL 32611-2035 Dr. Harry G. Lee 4132 Ortega Forest Drive Jacksonville, FL 32210 Dr. Charles Lydeard Biodiversity and Systematics Department of Biological Sciences University of Alabama Tuscaloosa, AL 35487 Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa P.O. Box 467 Wellington, NEW ZEALAND Dr. James H. McLean Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Paula M. Mikkelsen Paleontological Research Institution 1259 Trumansburg Road Ithaca, NY 14850 Dr. Diarmaid O Foighil Museum of Zoology and Department of Biology University of Michigan Ann Arbor, MI 48109-1079 Dr. Gustav Paulay Florida Museum of Natural History University of Florida Gainesville, FL 32611-2035 Mr. Richard E. Petit P.O. Box 30 North Myrtle Beach, SC 29582 Dr. Gary Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr. Angel Valdés Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 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 SUBSCRIPTION INFORMATION The subscription rate for volume 123 (2009) is US $50.00 for individuals, US $85.00 for institutions. Postage outside the United States is an additional US $10.00 for regular mail and US $25.00 for air delivery. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 1580, Sanibel, FL 33957, USA, (239) 395-2233. 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 by 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 P.O. Box 1580 Sanibel, FL 33957 CONTENTS NAUTILUS Volume 122, Number 4 December 22, 2008 ISSN 0028-1544 Daniel L. Geiger Sven N. Nielsen Claudio Valdovinos Juliana M. Harding Stefanie M. Gera Roger Mann Maria Cristina Dreher Mansur Claus Meier-Brook Cristian Ituarte Maria Eugenia Segade Guido Pastorino Alexandre Dias Pimenta Diogo Ribeiro do Couto Paulo Marcio Santos Costa Luiz Ricardo L. Simone New species of scissurellids from the Austral Islands, French Polynesia, and the Indo-Malayan eee (Gastropoda: Vetigastropoda: Scissurellidae, Anatomidae, Larocheidae) ..........cccecceecesecesecescescecacecscecscesenerecseensceneeeseensecsseeaee 185 Early Pleistocene mollusks of the Tabul Formation, South-Central Chile......... 201 Radula morphology in veined rapa whelks, Rapana venosa (Valenciennes, 1846) (Gastropoda: Muricidae) from C eee ake Bay, USA wis. .ictisesecutasesindantatnrnts 217 A new species of Sphaerium Scopoli, 1777, from southern Brazil (Bivalvia: Some remarks on the gross anatomy of Adelomelon ferussacii (Donovan, 1824) (Gastropoda: Volutidae) from the coast of Pate iwonia, A new species and a new record of Muricidae (Gastropoda) from Brazil: genera Pierynotus-and Leptotrophoticicteyicdescsnaistiutivaiioe anitie ctieinecvdedelibedeceeees ss 244 A new species of Chlamydoconcha Dall, 1884, from southeastern Brazil (Bivalvia: Ghilamiydocori chidae).o.<.asasssseves sos csvereion azensstatvsarnessariesenensioayessis apetsesasveedd nutans ots DO Research Note Jeremy S. Tiemann Sinistral Campeloma decisum (Say, 1817) (Gastropoda: Viviparidae) from the Kevin S. Cummings Box iver DM O18is.24 see cesrckesissnusicciedennen Bresbws eieaetaunsaeapcbeuieidebangccsw ake Waser Nakstedsaeasseuste ODO UB YoC0) Sed a 11 25 RRC ce OR 261, 263 TAU 2 aes eek ccs Re ce tsa sat ae ected te ch aateece ea tcc p at voik ou daise touachetadettatasvatairsceichasnnds@aias cuanser beets satis vaadtedetieeiteaysgsatteeseuters 264 IN UNC Safes -ac lee tgh tn nas ta, Apachettsusetveatics tien toaa baa banteabete sb cste toast tut etch Paes abe vtsiaGe chad bi cnesucoute ttaeies Sedecenates eenense ce tae lapaae eeu 264 Ath Ore Vr 6 asics seetendev sac heg Shee levdheciead Sea bd canada atone ond eae enc asisilacatosetdeca cguawien Peatengatanietee ceentetee ee eesands dddpueads foe sa tes 265 MBLWHOI Library JAN 9 7 2008 WOORS HOLE Massachusetts 02543 DOT BR WDE ODA STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION Publication Title, THE NAUTILUS. Publication No., 0028-1344. Filing Date, September 24, 2008. Issue Frequency, Quarterly. No. of Issues Published Annually, Four. Annual Subscription Price, US $80.00. Complete Mailing Address of Known Office of Publication, 3075 Sanibel-Captiva Road, Sanibel, FL 33957 USA. Complete Mailing Address of Headquarters, same as 7. Full Names and Complete Mailing Addresses of Publisher, The Bailey-Matthews Shell Museum, 3075 Sanibel-Captiva Road, Sanibel, FL 33957 USA. Editor, Dr. José H. Leal, address as above. Managing Editor, Mary Jo Bunnell, address as above. Owner, Shell Museum and Educational Foundation, Inc., address as above. Known Bondholders, Mortgagees, and Other Security Holders Owning or Holding 1 Percent or More of Total Amount of Bonds, Mortgages, or Other Securities, None. The purpose, function, and nonprofit status of this organization and the tax exempt status for federal income tax purposes has not changed during the preceding 12 months. Publication Name, THE NAUTILUS. Issue Date for Circulation Data Below, September 26, 2008. Extent and Nature of Circulation Average 12 months A. B. Total Number of ¢ Jopies Paid Circulation 1. Paid/Requested Outside-County Mail Subscriptions 2. Paid In-¢ sounty Subscriptions 3. Sales Through Dealers and Carriers, Street Vendors, Counter Sales, and Other Non-USPS Paid Distribution 4. Other Classes Mailed Through the USPS Total Paid and/or Requested Circulation Free Distribution by Mail 1. Outside-County 2. In-County 3. Other Classes Mailed Through the USPS Free Distribution Outside the Mail Total Free Distribution Total Distribution Copies not Distributed Total Percent Paid and/or Requested Circulation 420 309 0 0) 16 325 12 0 0) 0) 12 337 $3 420 97% Single Issue 420 319 0 0 15 334 12 0) 0 0 12 346 74 420 97% THE NAUTILUS 122(4):185-200, 2008 Page 185 New species of scissurellids from the Austral Islands, French Polynesia, and the Indo-Malayan Archipelago (Gastropoda: \ ‘etigastropoda: Scissurellidae, Anatomidae, Larocheidae) Daniel L. Geiger Santa Barbara Museum of Natural History Invertebrate Zoology 2559 Puesta del Sol Road Santa Barbara, CA 93105 USA geiger@vetigastropoda.com ABSTRACT Four new species of Scissurellidae, Anatomidae, and Larochei- dae are described from the Austral Archipelago, French Polynesia and the Indo-Malayan Archipelago: Sinezona danieldreieri new species, S. wiley new species (both Scissur- ellidae), Anatoma rapaensis new species (Anatomidae) and Trogloconcha lozoueti new species (Larocheidae). One of the species is currently only known from the Austral Islands T. lozoueti), while the others seem to show a_ broad Indo-Malayan Archipelago to western Pacific distribution. Shells of all species and the radulae of S. danieldreieri, T. lozoueti, and A.rapaensis are illustrated with scanning elec- tron micrographs. Additional Keywords: Mollusca, Rapa Island, radula INTRODUCTION The present contribution describes some microgas- tropods in the families Scissurellidae, Anatomidae, and Larocheidae known from the Austral Islands and be- yond. The impetus stems from a French expe dition to the southeastern-most Austral island, Rapa (see Lozouet et al.. 2004; 2005 for details); for the species found at Rapa, additional material from other localities located in various institutional collections has also been inclu- ded. This article is part of a series describing the bio- diversity of this remote archipelago (e.¢., Schwabe and Lozouet, 2006). Scissurellidae sensu lato comprises ean of small basal marine snails in Vetigastropoda (Geiger et al., 2008). The phylogenetic position a peer is cur- rently unsettled. Whereas Geiger and Thacker (2005; unpublishe -d data) resolved a Lepetodrilus (Lepetodrili- dae) + Scissurella, Sinezona, Sukashitrochus (Scissurel- lidae sensu stricto) clade with more distantly related Anatoma (Anatomidae), Kano (2008) using the same three genes (Histone 3, COI, 18S) recovered in some of his analyses a clade uniting all three of these lineages (Lepe todrilus, Sinezona, Anatoma). Scissurellidae and Anatomidae are characterized by a slit or foramen. in the shell (lacking in Larocheidae), a rhipidoglossate rad- ula, and a lack of nacre. Approxim itely 140 species are currently described with an additional 90 remaining to be formally recognized (Geiger, 2003; 2008). To date, no species in ‘these families ie ever been recorded from the Austral Islands. The species described herein are all new to science, one with its range restricted to the Austral Islands, and three others known from several localities in the broad Indo-Pacific. MATERIALS AND METHODS Standard methods for scanning electron microscopy (SEM) were employed as de taile d in Geiger (2006a, b, c) and Geiger et al. (2007). Terminology for shell tae and details on method of whorl count have been given elsewhere (Geiger, 2003; Geiger and Si isaki, 2008). Specimens cited are dry lots; *: comple te” indi- cates wet-preserved lots with animals. All de »pth indica- tions refer to bottom depth. Institutional abbreviations used in the text are: BRC: Bret Raines Collection, Kansas City, USA; DLG: Daniel L. Geiger Collection, Los Angeles, USA; JTC: Jean Trondlé Collection, Paris, France; LACM: Los Angeles County Museum of Natural History, Los Angeles, oe MNHN: Muséum national d’Histoire naturelle, Paris, France; SBMNH: Santa Barbara Museum of Natural History, Santa Coabare. USA: USNM: United States Na- tional Museum of Natural History, Washington (DC) USA; ZMA: Zoological Museum, Amsterdam, The Neth- erlands. Other abbreviations used in text are: M: Monotypy OD: Original designation; SEM: Scanning electron microscope/microscopy/micrograph Page 186 THE NAUTILUS, Vol. 122, No. 4 SYSTEMATICS Scissurellidae Gray, 1547 Sinezona Finlay, 1926 Type species: Schismope brevis Hedley, 1904 (OD). Remarks: The genus has recently been treated by Marshall (2002) and Geiger (2003). Sinezona danieldreieri new species (Figures 1-9) Description: Shell small (to 0.77 mm), trochiform. Pro- toconch of | whorl, with strong axial sculpture not reaching apical suture, apertural varix not connected to smooth embryonic cap, apertural margin straight. Tele- oconch I of 0.875 whorls, suture at ponnlien cee mately 9-15 strong, raised axial cords; first spirals after 0.5 whorls. Tleoconch 1 Il of 0.6 whorls, approximately four fine spiral cords at onset of selenizone between selenizone and deep suture below periphery, descending noticeably on last 0.25 whorls; shoulder slightly concave, approximately 13-20 axial cords, raised near suture, as high as wide towards selenizone; approximately 10 spir- als, distinct cords near suture, diminishing to very fine cords near selenizone; interstices with fine irregular growth lamellae. Base with constriction below ete ni- zone, approximately 15 elevated spiral cords, crossed and run over by approximately 16 fine spiral lines. Um- bilicus cone- shape sd, moderately wide, wall smooth, at distinct angle to base. Aperture De: shaped, roof overhan- ging. Selenizene above periphery, keels quite strong, moderately elevated, distinct linules, elongated feranten closed anterior ly. OpercuLuM: Thin, flexible, multispiral, with central nucleus. Raputa (Ficures 8-9): Rachidian tooth triangular cusp with approximately five equal sized deatclee Lateral teeth 1-3 similar; outer edge of cusp with 4-5 denticles. Lateral tooth 4 reduced in size, hook-shaped. Lateral tooth 5 enlarged by broadening, approximately six, large denticles on inner edge, 1-2 small ones on outer edge. Central denticle of inner marginal teeth large, approxi- mately three denticles on inner edge, four on outer Figure I. Holotype of Sinezona danicldreicri new species. Guinea, 5.283° S, 150.131° E, 0-5 m. Seale bar, shell (SBMNET $3540). Lumu Reef, Kimbe Bay, New Britain, Papua New 500 jim. Seale bar, protoconch = 100 jum, D. L. Geiger, 2008 Page 187 edge; outer marginal teeth spoon-shaped, bilaterally symmetrical with approximately a dozen fine denticles on each side. Radular interlock moderate. Type Material: Holotype: SBMNH $3540: Figure 1. 24 Paratvpes: 3-4 m, Little Santa Cruz Island, Mindanao, 6.854° N, 122.04° E (USNM 812453, 1: Figure 2). 1.5 m, Okinawa, Oku, Japan, 26.847° N, 128.287 E (LACM 77-61, 3: one illustrated Figure 3). 20 m, Off Figures 2-4. Paratypes of Sinezona danieldreicri new species 122.04° E, 3-4 m (USNM 812453, ] Island. Pamilacan Island. Philippines, 9.5° N, 123.917° E, 20 m (AMS 406342, 1). Scale bars, shell = 500 jum. Scale bars, protoconch = 100 um 3. Okinawa, Oku Japan, 26.847° N, 128.287° E, Bohol Is, Pamilacan Island, 9.5° N, 123.917° E (AMS 106342, 1: Figure 4). Type Locality: 0-5 m, Lumu Reef, Kimbe Bay, New Britain, Papua New Guinea, 5.283° S, 150.151° E. Col. D. L. Geiger Oct. 2005. Etymology: The name honors Daniel Dreier, son of Douglas and Hanna Dreier of Santa Barbara, who have Little Santa Cruz Island, Mindanao, Philippines, 6.584° N 1.5 m (LACM 77-61, 3). 4. Off Bohol Page 188 THE NAUTILUS, Vol. 122, No. 4 made generous contributions to the Santa Barbara Museum of Natural History. Distribution: Indo-Malayan Archipelago to southern Polynesia; shallow reef environments. Other Material Examined: FRENCH POLYNESIA: 0 m, Rapa Iti Island, Rapa, Austral Islands, 27.622° S, 144.302° W (MNHN, 1). 0 m, Kotuaie Point, Tupuaki Bay, Rapa, Austral Islands, 27.577° S, 144.343° W (MNHN, 1). 2 m, North of Pukitarava, Rapa, Austral Islands, 27.597° S, 144.227° W (MNHN, 4: two illu- strated Figures 5-6). 2-4 m, N of Aturapa Island, Rapa, Austral Islands, 27.572° S, 144.350° W (MNHN, 4). 2m, N of Rapa Iti Island, Rapa, Austral Islands, 27.620° S, 144.303° W (MNHN, 1). 2 m, N of Anatakuri Bay, Rapa, Austral Islands, 27.623° S, 144.307° W (MNHN, 15). 3-24 m, Hiri Bay, Rapa, Austral Islands, 27.622° S, 144.370° W (MNHIN, 4). 5 m, Anarua Bay, Rapa, Austral Islands, 27.605° S, 144.378° W (MNHN, 7: one illustrated Figures 7, 8-9 [radula]). 6 m, Mei Point, Figures 5-7. Sinezona danieldreicri new species. 5-6. North of Pukitarava, Rapa, Austral Islands, French Polynesia, 27.597° S, 144.227° W. 2m (MNIIN). 7. Anarua Bay, Rapa, Austral Islands, French Polynesia, 27.605° S, 144.378° W,5 m (MNHN). Radula of pecimen is shown in Figures S8—9. Seale bar. shell = 500 tum Seale bar, protoconch = 100 ttm. D. L. Geiger, 2008 Page 189 Figures 8-9. Radula of Sinezona danieldreieri new species, from specimen shown in Figure 7. Anarua Bay, Rapa, Austral Islands, 27.605° S, 144.378° W, 5 m. 8. Lateral teeth 4 (L4) and 5 (L5) plus marginal teeth. 9. Half width of radula with central field. Scale bars = 10 um. Rapa, Austral Islands, 27.637° S$, 144.303° W (MNHN, 1; MNHN, 1). 6 m, SW of Gotenaonao Point, Rapa, Austral Islands, 27.645° S, 144.320° W (MNHN, 3). 15-20 m, Vavai, Rapa, Austral Islands, 27.590° S, 144.381° W (MNHN, 1; MNHN, 1). 16-20 m, Mei Point, Rapa, Austral Islands, 27.637° S, 144.303° W (MNHN, 2). 1S m, Rarapai Island, Rapa, Austral Islands, 27.572° S, 144.368° W (MNHN, 1). 27 m, Kauira Point, Rapa, Austral Islands, 27.592° S$, 144.347° (MNHN, 1). 30 m, NW of Tauna Island, Rapa, Austral Islands, 27.605° S, 144.303° W (MNHN, 17). 36 m, around Rukuaga Point, Rapa, Austral Islands, 27.568° S. 144.368° Ww (MNHN, 8). PuiLipPINEs: Canipo Island, Cuyo Islands, Palawan Province, 11° N, 120.948° E (USNM 808195, 1). Guam: 0.7 m, Ajayan Bay, S. Guam, 13.245° N, 144.717° E (LACM 77-19, 2). Remarks: The protoconch sculpture shows some in- traspecific variability in the material examined. More specifically, the strong axial cords are sometimes discon- tinuous. This character is not restricted to the present species, hence, it does not seem to be of any systematic value. Those axial cords are also somewhat variable in width, the thickenings occasionally seeming to form some faint spiral slemente. Given the intra- and inter- specific variability of protoconch sculpture at this level of detail, the presence/absence of faint spiral elements is considered irrelevant for systematic purposes. Discrete differences can be observed between strong and weak axial cords as utilized previously in a systematic context (Geiger, 2003). Sinezona plicata (Hedley, 1899) from the Indo-Pacific has an overall wider and larger shell (2.3 mm; Geiger and Jansen, 2004b), more pronounced but fewer raised axial cords, and a eg aga with spiral lines. Sinezona ferriezi (Crosse, 1867) (? = S. hoernesi Semper, 1865: whereabouts of ee types unknown, original illu- stration lacking detail) with broad Indo- Malayan Archipelago distribution, is about twice as large, has a smooth protoconch with a distinct apertural varix, and >1 teleoconch I whorls. Sinezona globosa Geiger, 2006, from New Caledonia and Wallis Island is more rounded in overall appearance, lacks the elevated spiral cords, is larger (3.1 mm; Geiger, 2006a), and has spiral sculpture on the protoconch. Sinezona_ macleani Geiger, 2006, with broad Indo-Malayan distribution is more globular in overall outline, larger (2.3 mm; Geiger, 2006a), has a protoconch with flocculent sculpture, lacks the raised axial lines and has a wider umbilicus. All the above species grow to much larger size and would be imma- ture with open slit at the maximum size of S. daniel- dreieri. Sinezona wileyi new species (Figures 10-16) Description: Shell medium size (up to 1.08 mm) tro- chiform turreted. Protoconch slightly sunken in, with 1 whorl, fine irregular axials, no apertural varix, apertural margin straight. Teleoconch I with 0.6-0.7 whorls, su- ture at periphery, usually with approximately 14-22 faint to distinct fine axial cords, usually without spiral lines (occasionally faintest spiral threads only visible by SEM [Figure 11]), interstices with irregular thickenings. Tele- oconch IT with up to 1.25 wots, suture below periph- ery. Shoulder with approximately 32-44 _ barely perceptible to distinct fine commarginal axial cords, stronger near suture becoming less distinct towards sele- nizone; approximately eight baiely perceptible to very indistinct spiral cords, concentrated on middle of shou der; occasional specimens with predominant spiral sculpture (Figure 11). Base distinctly constricted below selenizone, axial cords of same strength and density as on shoulder, starting in lower portion of constriction be low selenizone; approximately 17 fine, irregularly spaced spi ral cords, starting in lower portion of constriction with onset of axial lines. Umbilicus wide, walls straigh 1 Page 190 THE NAUTILUS, Vol. 122, No. 4 Figure 10. Holotype of Sinezona wileyi new species. Lumu Reef, Kimbe Bay, New Britain, Papua New Guinea, 5.283° S, 150.131° BE, 0-5 m, (SBMNH 83541). Scale bar, shell = 500 jim. Scale bar, protoconch = 100 jun, smooth, periphery at distinct angle to base, bordered by carina. Aperture subquadratic, D-shaped, roof overhan- ging. Selenizone above periphery, keels moderately strong, quite elevated, lunules clistinct at recular interval: foramen narrow, elongated, margins converging and touching, without raphe. Animal unknown. Type Material: Holotype: SBMNH 83541 (Figure 10). 35 Paratypes: 0-5 m, Lumu Reef, Kimbe Bay, New Brit- ain, Papua New Guinea, 5.283° S$, 150.131° E (DLG 639, 19: one illustrated in Figure 15). Laisse de plage extérieure, Mururoa, Tuamotu, French Polynesia, 21.842° S$, 138.895° W (JTC, 2: one illustrated in Figure 12). 20 m, Herald Pass, W of Ndravuni Island, NW end of Great Astrolabe Reef, Fiji, 18.767° S, 178.467° E (LACM §5-135, 2: one illustrated in Figure 14). 20 m, Herald Pass, W of Ndravuni Island, NW end of Great Astrolabe Reef Fiji, 18.767° S, 178.467° E (LACM 85- Or 35, 2). 10-20 m, Off Francis Island in Beqa Lagoon, just off of Beqa Island (south of Viti Levu), 18.300° S, 178.067° E (DLG 279, 10: one illustrated in Figure 16). Type Locality: 0-5 m, Lumu Reef, Kimbe Bay, New Britain, Papua New Guinea, 5.283° $, 150.131° E. Etymology: The name honors the collector of the first known specimens of the species Tony Wiley of River- side, California, USA. Distribution: — Tropical Indo-Malayan Archipelago; shallow shelf, Other Material Examined: =F Rencu PoLyNesia: 36 m, Around Rukuaga Bay, Rapa, Austral Islands, 27.568" S, 144.568° W (MNHN, 20: Figures 11, 13). 2-4 m, N of Aturapa Island, Rapa, Austral Islands, 27.572° S, 144.350° W (MNHIN, 1). 5 m, Anarua Bay, Rapa, Austral Islands, 27.605° S, 144.375° W (MNEIIN, 4). 45 m, Haurei Bay, Rapa, Austral Islands, 27.613° S, 144.305" W (MNITIN, 1). D. L. Geiger, 2008 Page 19] Figures 11-13. Sinezona wileyi new species. 11-12. Around Rukuaga Bay, Rapa, Austral Islands, French Polynesia, 27.568° S, 144.368° W, 36 m (MNHN, 20). 13. Laisse de plage extérieure, Mururoa, Tuamotu, French Polynesia, 21.842" S$, 138.895° W Paratype: JTC. 2). Scale bars. shell = 500 pm. Scale bars, protoconch = 100 fm NeW CaLeponia: 25-30 m, Santal Bay, Lifou, 20.822 S. 167.173° E (MNHN, 1). 55-57 m, Koumac Sector, 20.672° S, 164.195° E (MNHN, 1). 5—25 m, Touho Sec- tor. 20.878° S, 165.325° E (MNHN, 1). 8 m, Touho Sector, 20.742° S, 165.265° E (MNHN, 1 Fiji: 10-20 m, Off Francis Island in Bega Lagoon, just ott of Bega Island (south of Viti Levu), 18.300° S$ 178.067° E (DLG 279. 10 Papua New Guinea: 70 m, off South Ema Reef, Kimbe Bay, New Britain, 5.202° S, 150.152° E (DLG 768, 3 Remarks: The organization of the foramen in this species seems to be intermediary between the typical open slit in Scissurella and the closed foramen in Sine- zona Although the margins of the slit are converging and fused anteriorly, a raphe is not evident The facts Page 192 THE NAUTILUS, Vol. 122, No. 4 that there is a marked downward deflection of the last quarter whorl and that the roof of the peristome attaches below the periphery of the previous whorl dem- onstrate that those specimens are fully grown and share more similarities with Sinezona than ‘with Scissurella with an open slit. Accordingly, S$. wileyi is placed in Sinezona. The species seems to have variable strengths of sculp- ture. Most oo have predominant axial sculpture (Figures 10, 14-15), while in occasional specimens (Figure 11) rea ‘asia sculpture is barely perceptible and the “spiral structures, particularly on the adumbilical part of base, become most prominent. As there are interme- diate specimens (Figure 13) with distinct adumbilical Figures 14-16. Sinezona wileyi new species, paratypes. 14. Herald Pass, W of Ndravuni Island, NW end of Great Astrolabe Reef, Fiji, 18.767° S, 178.467 a 20 m (LACM $5-135, 2). 15. Lumu Reef, Kimbe Bay, New Britain Papua New Guinea, 5.253° S, 150.131° E, 0-5 m (DLG 639, 19). 16. Off Franc is Island in Beqa Lagoon just olf of Beqa Island ( south of Viti Levu), Fiji, 18.3° S, 178.067° E, 10-20 m agen 10). Seale bars, shell 500 pum. Scale bars, protoconch LOO tun D. L. Geiger, 2008 Page 193 spiral lines and relatively weak axial lines, combined with identical condition of the protoconch and very small size, these sculptural differences are conside red to constitute intraspe cific variability. There are no similar ae s in the Indo-Pacific. Sine- sona ferriesi (Crosse, 1867) with broad Indo-Malayan Archipelago distribution is most similar, but is about twice the size of S. wileyi, has a smooth protoconch with distinct apertural varix, a teleoconch I with more than whorl, is more rounded overall, and a distinct raphe anterior to the closed foramen. Sinezona garciai Geiger, 2006, from the Caribbean, shares the protoconch sculp- ture, the absence of an apertural varix on the proto- conch, the relatively short teleoconch I with 0.6 whorls, and the narrow umbilicus. However, in addition to its occurrence in a separate ocean, S. garciai has a more inflated shell with an oval aperture, which connects to the previous whorl barely below the periphery, has only about 0.6 teleoconch TH whorls, and bears a distinct raphe anterior to the foramen. Anatomidae McLean, 1989 Anatoma Woodward, 1859 Type Species: Scissurella crispata Fleming, 1828 (M) Remarks: The genus has recently been treated by Marshall (2002), Gei iger (2003), and Geiger and Jansen (20042). Anatoma rapaensis new species (Figures 17-23) Description: Shell of medium size (to 1.82 mm), tro- chiform biconical. Protoconch with 0.75 whorls, floceulent sculpture, no apertural varix, apertural margin slightly si- nusoid. Teleoconch I with 0.3-0.4 whorls, approximate- ly 9-12 axial cords, interstices with fine flocculent sculpture, occasionally (holotype) flocculent sculpture somewhat concentrated in position of selenizone. Teleo- conch IE with up to 2.25 whorls, suture immediately below selenizone in early growth, offset by width of selenizone Figure 17. Holotype of Anatoma rapaensis new species. St. 43, Haurei Bay, Rapa, Austral Islands, French Polynesia, 27.613° S 144.305° W, 45 m. (MNHN 20823). Scale bar shell = 1 mm. Scale bar protoconch = 100 tm. Page 194 THE NAUTILUS, Vol. 122, No. 4 Figures 18-20. Anatoma rapaensis new species, paratypes (MNHN). 18-19. St. 43, Haurei Bay, Rapa, Austral Islands, French Polynesia, 27.613° S, 144.305° W, 45 m (MNHN 20824). 20. St. 48, around Rukuaga Point, Rapa, Austral Islands, French Polynesia, in fully grown specimens. Shoulder convex, approximately 53-66 (n = 4) axial cords on first teleoconch H whorl, same density on remaining whorls; first fine spiral cords after 0.125 whorls, approximately 7-10 after one teleoconch I whorl, approximately 15-22 at apertural margin of fully grown specimen, becoming less distinct and more un- evenly spaced towards apertural margin: intersections of 27.568° S, 144.368° W, 36 m (MNHN, 30). Scale bars, shell = 1 mm. Scale bars, protoconch = LOO jum axial and spiral cords forming minute points. Base contin- uously sloping with narrow umbilicus; axial cords of same density and strength as on shoulder, approximately 19 spiral cords, fine spiral cords below selenizone turning into low steps from mid-base onwards; intersection of spiral and axial cords with fine points. Aperture ovoid D-shaped, roof overhanging, basal adumbilical portion D. L. Geiger, 2008 Page 195 flared. Selenizone at periphery, keel moderately strong, moderately elevated, slit open, margins converging towards apertural margin. OpercuLum (Ficure 23): As large as aperture, thin, cor- neous, nucleus central, multispiral. Raputa (Ficures 21-22): Rachidian tooth trapezoid, central denticle with 3—4 denticles on each side. Lateral teeth 1-3 similar, development of cusp reduced periph- erally, terminal denticle largest, 4-2 denticles on outer edge. Lateral tooth 4 reduced in size, hook-shaped, with one minute point on each side. Lateral tooth 5 enlarged with four strong denticles on inner margin. Inner mar- ginal teeth elongated, terminal denticle largest, 3-4 smaller denticles on inner margin, three larger denticles on outer margin. Outer marginal teeth with cup-shaped cusp with many small denticles. Type Material: Holotype: MNHN 20823 (Figure 17). 60 Paratypes: 45 m, RAPA St. 43, Haurei Bay, Rapa, Austral Islands, French Polynesia, 27.613° S, 144.305° W (MNHN 20824, 29; two illustrated in Figures 18-19). 36 m, RAPA St4S, Around Rukuaga Point, Rapa, Austral Islands, 27.568° S, 144.368° W (MNHN, 30; one illustrated in Figure 20). 145 m, MUSORSTOM 9 St. CP 1159, Eiao Island, 7.972° S, 140.728° E (MNHN, 1: Figures 21-23 [radula, operculum)). Type Locality: 45 m, RAPA St. 48, Haurei Bay, Rapa, Austral Islands, French Polynesia, 27.613° S, 144.305° W. Etymology: The species is named alter its type locali- ty, Rapa Island, French Polynesia. Distribution: Tropical Indo-Malayan Archipelago and Western Pacific; shallow shelf to upper slope. Other Material Examined: FRencu Potynesia: LO-15 m, Pake Bay, Rapa, Austral Islands, 27.617° S, 144.310° W (MNHIN, 2). 30 m, NW of Tauna Island, Rapa, Austral Islands, 27.605° S, 144.303° W (MNHN, 50). 33 m, Haurei Bay, Rapa, Austral Islands, 27.612° S, 144.318° W (MNHN, 10). 52-57 m, SE of Tauna Island, Rapa, Austral Islands, 27.608° S, 144.295° W (MNHN, 50: MNHN, 20). 100 m, North of Raivavae, Austral Islands, 23.828° S, 147.693° W (MNHN, 2). PHILIPPINES: 92-97 m, 12.517° N (MNHN, 8). New Careponia: 105-110 m, Poindimie Sector, 20.817° S, 165.317° E (MNHN, 9). 250-350 m, south- ern New Caledonia, 22.500° S, 166.400° E (MNHN, 1). 250-350 m, southern New Caledonia, 22.500° S, 166.400° E (MNHN, 1). 495 m, southern New Caledo- nia, 22.367° S, 166.233° E (MNHN, 3). 600-616 m, northern New Caledonia, 18.817° S, 163.250° E (MNHN, 3). . 120.650° E Figures 21-23. Aadula and operculum of Anatoma rapaensis new species. Eiao Island, Marquesas Islands, 7.972° S, 140.725° E, 145 m (MNHN, 1). 21-22. Radula. 23. operculum. Scale bar, radula Figure 21 = 50 wm. Scale bar, radula Figure 22 = 20 mm. Scale bar, opeculum = 500 um. Page 196 THE NAUTILUS, Vol. 122, No. 4 Marouesas Isutanps: 145 m, Eiao Island, 7.972° S, 140.728° E (MNHN, I: paratype), 200-220 m, Ua Huka Island, 8.900° S, 139.633° W (MNHN, 6). 352-358 m, Hiva Oa Island, 9.850° S$, 139.150° W (MNHN, 1). Fit: 149-168 m, S of Viti Levu, 18.207° S, 178.5 (MNHN, 1). 260-305 m, S of Viti Levu, 18.308° S, 178.097° E (MNHN, 6). 275-430 m, S. of Viti Levu, 18.297° S, 177.907° E (MNHN, 1). 441-443 m, S. of Viti Levu, 18.320° S, 177.862° E (MNHN, 1). Remarks. Anatoma rapaensis is characterized by the rather flattened overall shape, the short teleoconch I with less than 0.5 whorls, and the fine reticular sculpture that is axial-dominated on the first half teleoconch I whorl, subsequently becoming spiral-dominated. The most similar species is Anatoma (sensu lato) exquisita Schepman, 1908, from the Indo-Malayan Archipelago; the comparison is based on SEM imaging of the holo- type (ZMA 3.08.101; Figure 24). The shell is slightly taller, the spiral and axial lines are denser, forming more prominent points at their intersection, and the axial lines are slightly stronger than the spiral lines even in larger specimens (specimens 1.S—3.5 mm examined): protoconch and teleoconch I are eroded beyond recognition. Anatoma paucispiralia Bandel, 1998, from Satonda, Indonesia, has a smooth protoconch, a shorter teleoconch I (0.125 vs. 0.3— 0.4 whorls), and a slightly undulating shoulder profile. All other Indo-Pacific species are either more turreted or have noticeably different sculpture on shoulder and base. The only other documented case of a species that shows a change of sculpture on the teleoconch I is Anatoma jane- tae Geiger, 2006, known from 2,500 m off the west coast of North America (Geiger, 2006c). Larocheidae Fleming, 1927 Trogloconcha Kase and Kano, 2002 Type Species: Trogloconcha ohashii Kase and Kano, 2002 (OD). Remarks: The genus was recently treated by Geiger (2003). Trogloconcha lozoueti new species (Figures 25-30) Figure 24. Holotype of Anatoma exquisita Schepman, 1908 (ZMA 3.08.101). Siboga Station 95, 5°43.5' N, 119°40' E, 522 m between Sabah, Malaysia, and southeastern Tawitawi Group, Philippines]. Seale bar, shell = 1 mm. Scale bar, protoconch = 100 jum D. L. Geiger, 2008 Page 197 27. Trogloconcha lozoueti new species. 25. Holotype Vavai, Rapa Austral Islands, French Polynesia, 27.590° S 144.381° W, 15-20 m (MNHN 20825). 26-27. Radula from specimen shown in Figure 30. Hiri Bay, Rapa, Austral Islands, French 525 7 > 95_< Figures 25-2 Polynesia, 27.622° S_ 144.37° W, 3-24 m (MNHN). Scale bar, shell = 1 mm. Scale bar, protoconch = 100 tum. Scale bar, radulae = 10 pum Page 195 THE NAUTILUS, Vol. 122, No. 4 fy Figures 28-30. Trogloconcha lozoucti new species. Paratypes. 28-29. Vavai, Rapa, Austral Islands, French Polynesia, 27. 144.351° W, 15-20 m (MNHN 20826). 30. Hiri Bay, Rapa, Austral Islands, French Polynesia, 27.622° S, 144.37° W, 3-24 m MNHN, 20). Scale bar, shell 28 = 1 mm. Seale bars, shell 29-30 = 200 tum. Seale bars, protoconch = 100 jun Description: Shell medium size (to 1.17 mm), trochi- 31 spiral cords, from suture to mid-base as fine spiral form globular, with rapidly increasing whorls. Proto- cords, transitioning to low spiral steps in 5-6 adumbilical conch with 0.75 whorls, flocculent sculpture somewhat spirals: approximately 70 fine axial cords on last whorl, spirally arranged, weak apertural varix apertural margin forming distinct points at intersections; overall appear- convex. Teleoconch with up to two whorls, approximate- ance of shell fine reticulate and spiky. Suture bordered ly 15 axial cords on first 0.3-0.5 whorls with no spiral by strong irregularly lamellate thickening on shoulder. sculpture, interstices with fine flocculent sculpture; on- Base anomphalus with weak callus in umbilical region. set of spiral cords alter 0.5 whorls; approximately \perture oval, roof overhanging. D. L. Geiger, 2008 Page 199 OpercuLuM: Corneous, round, multispiral, with central nucleus, covering only approximately 1/3 of aperture. Raputa (Ficures 26-27): Rachidian tooth triangular, cusp with central denticle largest, three denticles on each side, arranged in convex curve. Lateral teeth 1-4 similar, L-shaped, cusp with apical denticle largest, 3-4 denticles on outer margin, 1-3 denticles on inner mar- gin; lateral tooth 5 enlarged by half, apical denticle larg- est, 3-4 denticles on each side. Immer marginal teeth with elongated shaft central denticle largest, 3-4 denti- cles on inner margin, 5-6 denticles on outer margin; outer marginal teeth with cup shape cusp, with many fine denticles on each side. Radular interlock of central field strong. Type Material: Holotype: MNHN 20825. 26 Para- types: 15-20 m, RAPA St. 32, Vavai, Rapa, Austral Islands, 27.590° S, 144.381° W (MNHWN 20826, 6: two illustrated in Figure 28-29). 3-24 m, RAPA St. 9, Hiri Bay, Rapa, Austral Islands, 27.622° S, 144.370° W (MNHN, 20: one illustrated in Figure 30). Type Locality: 15-20 m, RAPA St. 32, Vavai, Rapa, Austral Islands, French Polynesia, 27.590° S, 144.381° W. ee Named in honor of Pierre Lozouet (MNHN) for his accomplishments particularly in the field of fossil mollusks including Scissurellidae sensu lato, and his work in connection with the Rapa expedition. Other Material Examined: FRENCH PoLyNesIA: 36 m, Around Rukuaga Point, Rapa, Austral Islands, 27.568° S, 144.368° W (MNHN, 100). 33 m, Haurei Bay, Rapa, Austral Islands, 27.612° S, 144.318° W (MNHN, 6). 52- 57 m, SE of Tauna Island, Rapa, Austral Islands, 27.608° S, 144.295° W (MNHN, 20). 30 m, NW of Tauna Island, Rapa, Austral Islands, 27.605° S, 144.303° W (MNHN, 3). 8 m, S of Tarakoi Island, Rapa, Austral Islands, 27.093° S, 144.308° W (MNHN, 1). 52-57 Tauna Island, Rapa, Austral Islands, 27.608° S, 144.295° W (MNHN, 9) Remarks: = Trogloconcha ohashii from southem Japan has fewer spiral and axial elements that also form elevated points and a wide umbilicus. Trogloconcha tesselata Kase and Kano, 2002, from the Indo-Malayan Archipelago has a smooth protoconch and lacks the fine points at the intersection of axials and spirals. Trogloconcha christinae Geiger, 2003, from Western Australia lacks spiral sculp- ture and has flattened-flocculent protoconch sculpture. The new species is currently only known from the Austral Islands. The radula represents the general veti- gastropod pattern (rhipidoglossate) and : shores the typi- cal larocheid arrangement of similar lateral teeth 1-4, without reduced, hook-shaped lateral tooth 4; the radula confirms the placement of the species in Larochaeidae. The lack brood pouch seen in Larochaea and Larocheopsis places the species in Trogloconcha. Juveniles have a proportionally wider appearance with the suture approximately at the periphery of the SE of previous whorls. Fully grown specimens have a more elevated appearance with the suture connecting well below the periphery of the previous whorl. penpaen I DISCUSSION The new species all belong to recognized genera. They are diagnosed by particular character combinations not known from any described species. The two Sinezona species are among the smaller species in the genus, while the Anatoma and Trogloconcha species are within the usual size range for their respective genera. All spe- cies show characters of shell and radular morphology that are known from other species; no new character states were found. The combination of particular fea- tures and their particular strength of development, how- ever, are unique for each of the new species. The distribution of three species reaches beyond ve Austral Islands towards the equator (Sinezona wile yi, S danieldreieri, and Anatoma rapaensis), while one spe- cies (Trogloconcha lozoueti) is currently only known from Rapa Island. This pattern suggests a closer faunal affinity of the Austral archipelago pene the broad Indo- Malayan Archipelago, rather ae with the temperate Southern Ocean. The two more widely distributed spe- cies S. danieldrieri and S. wileyi are also those that have been found in deeper water, confirming the idea that deep-water species in general show a wider distribution than those restricted to shallow water. ACKNOWLEDGMENTS I thank on behalf of P. Lozouet all colleagues who participated in “RAPA 2002” and especially the mollusk group: R. von Cosel, V. Héros, A. Le Goff , P. Maestrati, J.-L., Menou, S. Schiaparelli and _ J. Trondlé. Many thanks also to Claude Payri (University of Papeete, Tahiti), who organized the logistics with the French Navy and the mayor and local "cone of Rapa. Pierre Lozouet prov ided helpful criticism on a draft version of the present contribution. Diego Zelaya and an anon- ymous reviewer improved the manuscript with helpful pointers. Total Foundation funded the expedition within its “Coral Reef Biodiversity Programme.” The SEM investigation was supported by NSF MRI-0420706 erant to Henry Chaney, Michi vel Caterino, and Daniel L. Geiger. Visiting curatorships from AMS and MNHN to DI c are kindly acknowledged. LITERATURE CITED Geiger, D. L. 2003. Phylogenetic assessment of characters proposed for the generic classification of Recent Scissur- ellidae (Gastropoda: Vetigastropoda) with a description of one new genus and six new species from Easter Island and Australia. Molluscan Research 23: 21-83. Geiger, D. L. 2006a. Eight new species of Scissurellidae and Anatomidae (Mollusca: Gastropoda: Vetigastropoda) from Page 200 THE NAUTILUS, Vol. 122, No. 4 around the world, with discussion of two new senior syno- nyms. Zootaxa 1128: 1-33. Geiger, D. L. 2006b. Sasakiconcha elegantissima new genus and species (Gastropoda: Vetigastropoda: Anatomidae?) with disjointly coiled base. The Nautilus 120: 45-51. Geiger, D. L. 2006c. A new blind Anatoma species from the bathyal of the northeastern Pacific (Vetigastropoda: Ana- tomidae). Molluscan Research 26: 108-122. Geiger, D. L. 2008. Monographing micromolluscs: A case study on Scissurellidae s.]. (Vetigastropoda). Zoosymposia 1: 133-145. Geiger, D. L. and P. Jansen 2004a. Revision of the Australian species of Anatomidae (Mollusca: Gastropoda: Vetigastro- poda). Zootaxa 414; 1-35. Geiger, D. L. and P. Jansen. 2004b. New species of Australian Scissurellidae (Mollusca: Gastropoda: Vetigastropoda) with remarks on Australian and Indo-Malayan species. Zootaxa 714; 1-72. Geiger, D. L., B. A. Marshall, W. F. Ponder, T. Sasaki, and A. Warén. 2007. Techniques for collecting, handling, and pa small molluscan specimens. Molluscan Research 27: 1-50. Geiger, D. L., A Niitzel, and T. Sasaki. 2008. Vetigastropoda. In: Phylogeny and Evolution of the Mollusca. W. F. Pon- der and D. R. Lindberg (eds). University of California Press, Berkeley, pp. 297-330. Geiger, D. L. and T. Sasaki. 2008. Four new species of Anato- midae (Mollusca: Vetigastropoda) from the Indian Ocean (Reunion, Mayotte) and Australia, with notes on a novel radular type for the family. Zoosymposia 1: 247-264. Geiger, D. L. and C. E. Thacker. 2005. Molecular phylogeny of Vetigastropoda reveals non-monophyletic Scissurellidae, Trochoidea, and Fissurelloidea. Molluscan Research 25: 47-55. Kano, Y. 2008. Vetigastropod phylogeny and a new concept of Seguenzioidea: independent evolution of copulatory organs in the deep-sea habitats. Zoologica Scripta 37: 1-21. Lozouet, P. R. von Cosel, V. Héros, A. Le Goff, P. Maestrati, J.-L. Menou, S. Schiaparelli, and J. Tréndlé. 2004. LAtelier Rapa 2002 (Polynésie frangaise). Xenophora 107: 17-28. Lozouet, P., R.von Cosel, V. Héros, A. Le Goff, P. Maestrati, J.-L. Menou, S. Schiaparelli, and J. Tréndlé. 2005. Biodi- versity gradient in the Pacific: first results of RAPA 2002 (French Polynesia). In: G. Richard (ed.) Les mollusques dans la recherche actuelle. Actes du Iéme Congres International des Sociétés Européennes de Malacologie (La Rochelle 24-27 juin 2003): 93-99. Marshall, B. A. 2002. Some Recent scissurellids form the New Zealand region, and remarks on some scissurellid genus group names (Mollusca: Gastropoda). Molluscan Re- search 22; 165-181. Schwabe, E. and P. Lozouet. 2006. Chitons (Mollusca, Poly- placophora) from Rapa, the southernmost island of Polynesia. Zoosystema 28: 617-633. THE NAUTILUS 122(4):201-216, 2008 Page 20] Early Pleistocene mollusks of the Tubul Formation, South-Central Chile Sven N. Nielsen! GeoForschungsZentrum Potsdam Sektion 3.1 Telegrafenberg 14473 Potsdam GERMANY nielsen@gpi.uni-kiel.de Claudio Valdovinos Center of Environmental Sciences, EULA-Chile Universidad de Concepeidn! Casilla 160-C Concepcion, CHILE and Patagonian Ecosystems Research Center (CIEP) Coyhaique, CHILE evaldovi@udec.cl SES ABSTRACT Early Pleistocene mollusks of the Tubul Formation are re- viewed and 23 species are recognized. These are twelve bivalves, ten gastropods, and one scaphopod. With two excep- tions, all of them are known from the Recent. The fauna is compared with Miocene, Pliocene-Pleistocene, and Recent faunas of the Chilean coast. The composition of the Tubul fauna resembles the Recent one of the Magellan Region, sug- gesting cooler water temperatures for south-central Chile dur- ing the early Pleistocene than at present. Additional Keywords: Mollusca, eastern Pacific, climate INTRODUCTION The marine late Pliocene to early Pleistocene of Chile comprises localities distributed along almost the whole length of the coast of Chile (Figure 1). The northern localities attributed to the Pliocene and Pleistocene are exclusively nearshore environment outcrops between Peninsula Mejillones and Coquimbo that have been de- scribed in detail by Herm (1969) and Le Roux et al. (2004, 2005, 2006). Herm (1969) described the faunas from these localities and also commented on the fauna of the La Cueva Formation, which overlies the Miocene Navidad Formation of central Chile. Pliocene faunas from southern Chile have been described from the islands Guafo (Frassinetti, 1997, 2000) and Guamblin Frassinetti and Covacevich, 1995). Another late Miocene or early Pliocene age from central Chile is known from Lo Abarca near San Antonio (Covacevich and Frassinetti, 1990: DeVries, 2003). ’ Current address: Institut fiir Geowissenschaften, Christian- Albrechts-Universitat zu Kiel, Ludewig-Meyn-Str. 10, 24118 Kiel, GERMANY. fauna of The Tubul Formation (Figure 2) was first described by Feruglio (1949), with its type area located on the south- ern side of the Gulf of Arauco (37°14! S, 73°26’ W). Onshore outcrops are limited to the east by the Cor- dillera de Nahuelbuta and to the west by the Pacific Ocean. Feruglio (1949) described the sediments of the Tubul Formation as light-gray soft tuffaceous sandy mudstones with fossil remains, which can reach a thickness of up to 100 m (Garcia, 1968). Using macro- fossils, Briiggen (1950) showed that these sediments can be correlated with those of Coquimbo. From their study of microfossils, Martinez and Osorio (1968) con- cluded that the basal part of the Tubul Formation corresponds to the middle Pliocene. However, later studies by Martinez (1976) in Caleta Hueton led to the conclusion that the base of the Tubul Formation corresponds to the late Pliocene. Subsequently, Bird- Bagoczky (1979) recognized a lower and an upper member: the lower member has a thickness of approx- imately 50 m and is formed by very fine, slightly calcareous, dark-gray sandstones, including abundant fossil marine fauna; the upper member a6 has a thickness of approximately 50 m and is formed by light-gray to olive fine sandstones and mudstones with rare Foals: According to Pineda (1983, 1986), deposition of the sediments of the Tubul Formation was rapid, in a quiet zone close to the coast, maybe lagoonal. These sedi- ments should correspond to the Plio-Pleistocene-bound- ary (Pineda, 1983, 1986) and occur at the following localities: Punta Pichicui, Estero Chupalla, Villa Alegre, Los Alamos, Quebrada Raquilco, El Tique, Estero Licauquen, and Minas Trihueco. According to Pineda, the Tubul Formation represents a transgressive se- quence. The presence of quartz-sand and a conglomer- ate at the base of the formation has been interpreted as a former coastal line. Above this level, very fine clayey sandstones have been deposited, which should re present Page 202 THE NAUTILUS, Vol. 122, No. 4 | 18°S 20°S (22°S 30°S 32°S 34°S 36°S 38°S Elevation (m) 40°S 7000 42°S = 44°S 46°S 48°S 50°S 52°S 6 5 16 4 9 1 54°S 2 1 3.5 78101214 1719 13 18 ™ Beg kilometers M.9L M.bL c 0 8 M.99 Figure 1. Extant ranges of bivalve and gastropod species occurring at Las Penas. 1. Ennucula grayi. 2. Tindariopsis sulculata. 3. g 8 I I 8 gray Malletia chilensis. 4. Zygochlamys patagonica. 5. Cyclocardia velutinus. 6. Macoma inornata. 7. Darina solenoides. 8. Ensis macha. yg Y i y 9. Retrotapes exalbidus. 10. Pandora cistula. 11. Epitonium magellanicus. 12. Fusitriton magellanicus. 13. Trophon geversianus. 14. “Xymenopsis” dispar. 15. Chorus giganteus. 16. Nassarius taeniolatus. 17. Adelomelon ancilla. 18. Bela paesleri. 19. Scaphander interruptus. Stars indicate Plio—Pleistocene fossil localities. S. N. Nielsen and C. Valdovinos, 2008 Page 203 5880 km ARAUCO GULF Punta las Penas 5878 Figure 2. a shallow-marine environment. Radic et al. (2005) cited a thickness of 100 to 500 m for the Tubul Formation based on well data and seismic lines. The molluscan fauna decribed in this paper comes from the coastal bluffs near Las Pefias (Figure 2), which represent the uppermost part of the Tubul Formation. MATERIALS AND METHODS The mollusks described in this study belong to the col- lection of the late Professor Lajos Bir6 and are housed in the Departamento de Ciencia de la Tierra, Universidad de Concepcion, ple The material comes from near Punta Las Penias (37°14/32” S, 73°26'02” W, Figure 2), south of the village Tubul. and is designated by ihe let- ter ‘T° in the collections. The number of specimens inspected for each species is indicated by a number in parentheses if the number is greater than one. Species are not formally described because all except two are well known, Modern geographic ranges of species are modified from Valdovinos (1999). SYSTEMATICS Class Bivalvia Linnaeus, 1758 Family Nuculidae Gray, 1824 Genus Ennucula Iredale, 1931 Ennucula grayi (dOrbigny, 1846) Figure 3) Nucula obliqua Sowerby, 1833: 5, pl. 16, fig. 21. Nucula grayi 7 Orbigny, 1846: vol. 5, 625 (nomen novum for N. obliqua Somerby, non Lamarck, 1819). Ennucula grayi (d’Orbigny).—Soot-Ryen, 1959: 13, ph. 1 fig. 8: Villarroel and Stuardo, 1998: 13 3, figs. 8, 70, 71, 3, LOT—-109. iephaceate araucana (Philippi) —Villarroel and Stuardo, 1998: 165 (partim), figs. 142-143. Area of the Tubul Formation and sample site at Las Penas. Height of coastal bluff about 18 m. Ennucula valdiviana (Philippi) —Villarroel and Stuardo, 1998: 165 (partim), figs. 134-137 Ennucula lebuensis (Philippi).—Villarroel and Stuardo, 1998: 165 (partim), figs. 138-139. Material Examined: 47 specimens: T/4 (2), T/5 (6), T/7 (5), T/11, T/17, T/204, T/237, T/386, oie T/1276 (3), Daal , T/1394—-1397, T/1537, T/1749-1754, T/1968 (7), T/1969 (2), T/1989-1990, T/2048, T/2090. Measurements: Width 19.2 mm, length 16.7 mm (T/237) Occurrence: Recent: Coquimbo to Magellan Region; Plio-Pleistocene: Arauco to ?Isla Guafo, Remarks: = Nucula barrosi Philippi, 1887 (=N. arau- cana Philippi, 1587), N. lebuensis Philippi, 1887, and N. valdiviana Philippi, 1887, are Miocene species from the Navidad, Ranquil, and Sto. Domingo formations of central and southern Chile. However, their generic placement is not known and therefore possible relation- ships to E. grayi remain unclear. The specimens men- tioned and figured by Villarroel and Stuardo (199%) belong in E. grayi and not in one of these Miocene species. The taxonomic positions of these older species must await a revision of the Chilean Miocene nuculoids. Genus Tindariopsis Verrill and Bush, 1897 Tindariopsis sulculata (Gould, 1852) (Figure 4) . Nucula striata King and Broderip, 1832: 337 (non Lamarck, 1805). Nucula sulculata Gould, 1852: 12, 434, pl. 37, figs. 539 a-e (Couthouy MS). Nucula elegans Hupé, 1854: 305, Conquiliologia pl. 5 fig. 7; Philippi, 187: 189, pl. 31, fig. 6. Tindariopsis sulculata (Gould).—Dell, 1964: 149; Villar- roel and Stuardo, 1998: 144-145, figs. 39-41, SO-S2 157-159. THE NAUTILUS, Vol. 122, No. 4 S. N. Nielsen and C. Valdovinos, 2008 age 205 Page 205 Nuculana sulculata (Gould)—Dell, 1971: 167, pl. 1, figs. 14, pl. 2, fig. 9. Nuculana elegans (Hupé).—Frassinetti and Covacevich, 1995: 51, text-fig. 3a, pl. 1, figs. 1-5; Frassinetti, 1997: 60, pl. 1, figs. 5—7. Tindaria sulculata (Gould).—Griffin and Nielsen, 2008: 16, pl. 5, figs. 3-5. Material Examined: ca. 1S50 specimens: T/1 (92), T/S (24), T/9 (89), T/12 (17), T/13 (36), T/14 (17), T/15 (48), T/16 (9), T/1S (198), T/19 ges T/32 i T/53-54, ne (4), T/149 (9), T/164 (2), T/168 (8), T/206, T/207 ), T/236 (4), T/296 (3), T/542, T/97 4 (2), T/982-984 (ca, ea T/1083 (7), T/1182-1187 (ca. 90), T/1235 (6), T/1270 (293), T/1308 (ca. 30), T/1379 (92), T/1518 (11), T/1536 (21), T/1624, T/1681 (ca.45), T/1682 (6), T/1802— 1804 (ca. 90), T/1911 (23), T/1912 (ca. 90), T/1913, T/1970 (15), T/2001 (5), T/2009 (23), T/2042 (7). Measurements: Width up to 16 mm (1/1682), Occurrence: Recent: Talcahuano to Strait of Magel- lan; Plio-Pleistocene: Arauco to Isla Guamblin. Remarks: As for many other species, there are different names for Recent and fossil records of the same species. Villarroel and Stuardo (1998, p. 145), in their review of the Recent and fossil species of Chilean Protobranchia, noted that “The ornamentation of the shell of T. sulculata is very similar to that of the fossil species 7. elegans (...). It differs from that by its shorther length and by the truncation of its extreme posterior.” However, we do not consider these differences as sufficient for specific separation and regard them as intraspecific variation instead. Family Malletiidae H. and A. Adams, 1858 Genus Malletia Moulins, 1832 Malletia chilensis Moulins, 1832 (Figure 5) Malletia chilensis Moulins, 1832: 85, pl. 1, figs. 1, 8. Malletia inequalis Dall, 1908: 219, 383. ?Malletia sp. Frassinetti and Covacevich, 1995: 50. Material Examined: Nine specimens: T/544(2), T/1091-1093, T/1917, T/2176-2178. Measurements: Figured specimen width 26 mm (T/1092). Occurrence: Recent: Coquimbo to Magellan Region; Plio-Pleistocene: Arauco to Isla Guamblin. Remarks: According to Villarroel and Stuardo (1998), there are four or five species of Malletia known from Chile, all of which are rather similar to each other. Apart from M. chilensis, they recorded M. patagonica Mabille and Rochebrune, 1889, M. inequalis Dall, 1908, and M. magellanica (Smith, 1875). Malletia hyadesi Mabille and Rochebrune, 1889 was listed as a probable synonym of M. patagonica. Malletia volckmanni ( (Philippi, 1887) most probably is of Miocene age and has not been revised and its placement should be regarded with caution. Family Mytilidae Rafinesque, 1815 Genus ?Mytilus Linnaeus, 1758 Mytilus sp. (Figure 6) Material Examined: One specimen: T/106. Measurements: Length 58 mm (T/106). Remarks: According to Valdovinos (1999) there are nine species of Mytilidae known from Chile. Based on the main characters of the our shell fragment, such as the pointed morphology of the umbo and fine external growth striae, our specimen may correspond either to he genus Mytilus or Choromytilus. In southern Chile the species Choromytilus chorus (Molina, 1782) and Mytilus edulis chilensis Hupé, 1854, coexist, but it is not possible to identify our specimen to species level. Mytilidae of uncertain generic and specific status are known from Miocene through Recent and are in need of a detailed revision. Family Pectinidae Rafinesque, 1815 Genus Zygochlamys thering, 1907 Zygochlamys patagonica (King and Broderip, 1532) (Figure 7) 9907 Pecten patagonicus King and Broderip, 1832, 5: 337. Pecten tenuicostatus Hupé, 1854: 291, pl. 5, fig. 4; Phi- lippi 1887: 203, pl. 47, fig. 1; Méricke, 1896: 580, pl. 12, figs. 13-16. Chlamys patagonica patagonica (King and Broderip). —Beu, 1985: 1-11, pl. 1, figs. 1-4. Zygochlamys Paes (King and Broderip).—Waller, 1991: 28- 30, pl. 2, figs. 13, 14; Griffin and Nielsen, 2008: 37, pl. 16, f igs. 5, 6. Chlamys a ee (Hupé).—Frassinetti and Covace- vich, 1995: 52, text-fig. 3b, pl. 1, figs. 6-9. Material Examined: 305 specimens: 1/63, T/79, T/135, T/147, T/166, T/175, T/180, T/222-223, T/243, T/255, T/256, T/293, T/305, T/307—309, T/311—-312, T/329-334, T/361-380, T/387—399, T/432-434., T/491-504, T/577-578, T/725-726, T/72S8—-772, T/775-— 777, T/SO6—-820, T/1006—1017, T/1039, T/L074—1077, T/1080-1082, T/1156-1157, T/1160—-1166, T/1269, Figures 3-13. Bivalvia. 3. Ennucula grayi (dOrbigny, 1846) [length 19.2 mm]. 4. Tindariopsis sulculata (Gould, 1852) [length 16.0 mm]. 5. Malle‘tia chilensis Moulins. 1832 [length 16.0 mm]. 6. Mytilus sp. [length 58.0 mm]. 7. Zygochlamys patagonica (King and Broderip, 1832 1844) [length 16.0 sin] 10. Darina solenoides (King and Broderip, 1832 [length 75 mm]. 8. Cyclocardia velutinus (E. A. Smith, 1881) [length 18.5 mm]. 9. Macoma inornata (Hanley, ) Hength 44.0 mm] 11. Ensis macha (Molina, 1782) {length 97.2 mm]. 12. Retrotapes exalbidus (Dillwyn, 1817) [length 62.0 mm]. 13. Pandora cistula Gould, 1850 {length 23.0 mm]. Page 206 T/1280-1281, T/1284, T/1309-1310, T/1358, T/1371- 1378, T/1380-1386, T/1389-1392, T/1415-1424, T/1514-1516, T/15538-1559, T/1579-1586, = T/1632, T/1677-1680, T/1714-1725, T/1827, T/1863-1875, T/1964-1967, T/2003-2008, T/2030-2031, — T/2033, T/2037-2038, T/2049, T/2052, T/2055-2056, T/2059- 2065, T/2154—2167, T/2181-2192, T/2225-2239. Measurements: Height 75 mm, width 73 mm (T/223). Occurrence: Magellan; Plio-Pleistocene: Arauco to Isla Guamblin. Remarks: Because revision of Chilean pectinids is not within the scope of this work, relationships of Z. patago- nicus to the northern species described by Herm (1969) are not discussed here. Zygochlamys patagonicus is the type species of Psychrochlamys Jonkers, 2003. However, like Griffin and Nielsen (2008) and Dijkstra and Marshall (2008), we consider Psychrochlamys a synonym of Zygochlamys. Family Carditidae Fleming, 1528 Genus Cyclocardia Conrad, 1867 Cyclocardia velutinus (E.A. Smith, 1881) (Figure §) Cardita (Actinobolus) velutinus E.A. Smith, 1881: 42, pL 5, fig. 8. Corda acloenaini Philippi, 1SS7: 167, pl. 37, fig. 4. Cardiocardita volckmanni (Philippi). _Frassinetti and Covacevich, 1995: 53, text-fig. 3c, pl. 1, figs. 10-17; Frassinetti, 1997: 65, pl. 1, figs. 17-20. Material Examined: 19 specimens: T/507-510, T/ 639, T/786-787, T/1546-1552, T/162S—1630, T/1756, T/2256. Measurements: (T/508). Width 18.5 mm, length 17 mm Occurrence: Recent: Arica to Beagle Canal; Plio- Pleistocene: Arauco to Isla Guamblin. Remarks: A number of different Cyclocardia species has been reported from southern Chile (Forcelli, 2000). Family Tellinidae Blainville, 1814 Genus Macoma Leach, 1819 Macoma inornata (Hanley, 1S44) (Figure 9) Tellina inornata Hanley, 1844, 1847: 315, pl. 59, fig. 127. Tellina tubulensis Philippi, 1887: 134, pl. 32, fig. 7. Macoma (Psammacoma) inornata (Hanley).—Soot-Ryen, 1959: 62, pl. 4, figs, 34-36. Macoma_ tubulensis pl. 2, fig. 6 Frassinetti, 1997: 72, (Philippi).- Material Examined: Eight specimens: T/64, T/721-— 724, T/789, T/1236, T/2067. Measurements: Width figured specimen (T/64) 26 mm Recent: Island of Chiloé to Strait of THE NAUTILUS, Vol. 122, No. 4 Occurrence: Recent: Atacama to Gulf of Ancud; Plio- Pleistocene: Arauco to Isla Guamblin. Remarks: Herm (1969, p. 119) described a specimen of Ardeamya sp. as the only tellinid from the Pliocene south of San Antonio. Since Herm knew the Philippi collection, it must be assumed that it is not conspecific with the species reported here. Family Mactridae Lamarck, 1809 Genus Darina Gray, 1853 Darina solenoides (King and Broderip, 1832) (Figure 10) Erycina solenoides King and Broderip, 1832: 335. Darina solenoides (King and Broderip)—Smith, 1905: 337, Mactra sp.—Frassinetti and Covacevich, 1995: 54. Material Examined: One specimen: T/1238. Measurements: Width figured specimen (T/1238) 44 mm. Occurrence: Recent: Strait of Magellan and Tierra del Fuego; Plio-Pleistocene: Arauco to Isla Guamblin. Remarks: Herm (1969) discussed the problems of related forms appearing in the Pliocene of Coquimbo to San Antonio which he placed in the genus Mulinia Gray, 1837. Family Pharidae H. and A. Adams, 1858 Genus Ensis Schumacher, 1817 Ensis macha (Molina, 1782) (Figure 11) Solen macha Molina, 1782: 203; Hupé, 1854: 369, pl. 8, fig. 6. Solen oladiolus Sowerby, 1839: 153, pl. 43, fig. 4. Solen ‘gladiolus? Sowerby. —Philippi, 1887: 169, pl. 34 fig, 9. Ensis macha (Molina). Ensis sp.—Frassinetti, 1997: 67. pl. 2 —Carcelles, 1944: 292. , figs. 4-5. Material Examined: 38 specimens: T/192, T/579-580, T/793—797, T/1084, T/1 101-1102, T/1175—1176, T/1285— 1290, T/1368—1370, T/1412—1414, T/1694, T/1707—-1712, T/1923, T/1972, T/2000, T/2068, T/2226-2227. Measurements: Up to 97.2 min (T/1707). Occurrence: Recent: Caldera to Magellan Region; Plio-Pleistocene: Coquimbo to Isla Guafo. Remarks: —Ensis macha is one of the few species which has a very wide geographical range in Pliocene and Re- cent faunas of Chile. During the Pleistocene it also reached Mejillones (Herm, 1969; own observations). Family Veneridae Rafine sque, 1815 Genus Retrotapes del Rio, 1997 tetrotapes exalbidus (Dillwyn, 1817) (Figure 12) S. N. Nielsen and C. Valdovinos, 2008 Page 207 Venus exalbida Chemnitz, 1795: 225, pl. 202, fig. 2 (non binom.); Dillwyn, 1S17: 170. Venus aerea Hupeé, 1854: 338 Venus subalbicans Hupé, 1854: 339. Venus araucana Philippi, LSS7: 117, pl. 17, fig. 6; Tavera and Veyl, 1958: 170, pl. 4, fig. ite Samarangia exalbida (Dilhwyn).—Carcelles, 1944: 287, pl. 12, figs. 93, 94. Eurhomalea araucana (Philippi).—Frassinetti, 1974: 47, figs. 1-2; Frassinetti and Covacevich, 1995: 54, text-fig. 3c, pl. 1, fig. 1S; Frassinetti, 1997: 74, pl. 2, fig. 6. Retrotapes exalbida (Dillwyn).—del Rio, 1997: 80-82, figs. 22, 23, 41. Retrotapes meAlbius (Dillwyn).—Reid and Osorio, 2000: 139, fig. 5]; Griffin and Nielsen, 2008: 7, 35, pl. 1 figs. 2-4, pl. 16, figs. 1-3. Material Examined: 669 specimens: T/21—25, T/44— 49, T/56, T/5S—62, T/65-73, T/75, T/78, T/S4-109, T/136-140, T/151, T/153-163, T/173-176, T/1S1—-191, T/19S8—203, T/20S—213, T/216-221, T/226-235, T/241- 942. T/249-253, T/257-263, T/266-272, T/277-282, T/319-328, T/400-421, T/449-490, T/516-539, T/545— 557, T/607-614, T/653-671, T/681—703, T/710-719, T/S02, T/S29-SS1, T/901-910, T/919-968, T/97S, T/98S7—-989, T/1027—-1037, T/1109-1114, T/1128-1134, T/1142-1154,) T/1158-1159, T/1194-1197, T/1212- 1217, = T/1228-1229, = T/1237-1268, = T/1298-1306, T/1316-1324, T/1333-1356, T/1425-1428, T/1511— 1518, 1/1534, T/153S-1544, T/1645-1676, T/17S1- 1796, =T/1805-1809, = T/1817-1826, — T/1835-1862, T/1949-1963, T/2011-2014, T/2035-2036, T/2039— 2041, T/2046, T/2058, T/2081, T/2093-2097, T/2140- 2147, T/2230-2249. Measurements: Up to width 62 mm, length 54.5 mm (T/967). Occurrence: Recent: Chiloé to Strait of Magellan: Plio-Pleistocene: Arauco to Isla Guamblin. Remarks: A number of different Eurhomalea species, most probably now belonging in Retrotapes, has been reported from northern Chile (Herm, 1969). Family Pandoridae Rafinesque, 1815 Genus Pandora Bruguiere, 1797 Pandora cistula Gould, 1850 (Figure 13) Pandora cistula Gould, 1850: 217. Kennerlyia patagonica Dall, 1915: 450. Pandora sp.—Frassinetti and Covacevich, 1995: 55, pl. 1, fig. 19. Material Examined: T/141—146. 1/345, 1T/353-354, T/615-634, T/781. T/783, T/784 (3), T/972-973, T/986 (2), T/1273 T/1291-1292, T/1311-1312, T/1405-1411, T/1535 (8), T/1755, T/2175, T/2225. Measurements: Width 23 mm (T/22235). Occurrence: Recent: Valparaiso to Magellan Region; Plio-Pleistocene: Arauco to Isla Guamblin. Remarks: This species is the only living species of Pandora recorded in Chile (see Forcelli, 2000). Class Gastropoda Cuvier, 1795 Family Epitoniide ie Berry, 1910 Genus Epitonium Roding, 1798 Subgenus Nitidiscala de Boury, 1909 Epitonium (Nitidiscala) magellanicus (Philippi, 1845) (Figure 14) Scalaria magellanica Philippi, 1845: 46. Scalaria (Opalia) magellanica Strebel, 1905: 656, pl. 23 figs. 44a—f. Cirsotrema (Coroniscala) magellanicum (Philippi). — Wenz, 1938: 798, fig. 2322. Epitonium (Nitidiscala) magellanica(Philippi).— 1987: 70, fig. $2. siaee biroi - rassinetti and Covacevich, 1995: 56, pl. 2, figs. 1-2; Frassinetti, 1997: 135, pl. 1, figs. 4-5. Rael Examined: One specimen: T/2071. Height 20.5 mm. Occurrence: Recent: Strait of Magellan; Plio-Pleisto- cene: Arauco to ?Isla Guamblin. Ramirez, Measurements: Remarks: A number of different epitoniid species has been reported from southern Chile (Forcelli, 2000). The type material of Epitonium biroi Frassinetti and Cova- cevich, 1995 from Guamblin was not revised, so it remains unclear if it represents a different species or a synonym. Family Naticidae Guilding, 1834 Genus Euspira Agassiz, 1838 Euspira guamblinensis Frassinetti and Covacevich, 1995 (Figure 15) Polinices (Euspira) guambline nsis Frassinetti and Cova- cevich, 1995: - text-fig. dae, pl. 2, figs. 3-5; Fras- sinetti, 1997; 135, pl. 1, “fips, 6-9. Material Examined: 37 specimens: 1/30, T/152, T/215, T/335-337, T/778, T/779, T/976, T/1177-1181, T/ 1272, T/1279, T/1627, T/1684, T/1696, T/1704, T/1705, T/1777, T/1899, T/1901, T/1902, T/1909, T/1940-1942, T/1991, T/2053, T/2068, T/2151, T/2210-2212, T/2255. Measurements: Height up to 22.5 mm, figured speci- men (T/1909) height 14 mm. Occurrence: Recent: Extinct; Plio-Pleistocene: Ara- uco to Isla Guamblin. Remarks: None of the extant naticid species rev- iewed by Pastorino (2005a) matches Euspira guambli- nensis in its callus characters and we regard it as a valid species. Family Ranellidae Gray, 1854 Genus Fusitriton Cossmann, 1903 THE NAUTILUS, Vol. 122, No. 4 Page 208 S. N. Nielsen and C. Valdovinos, 2008 Page 209 Fusitriton magellanicus (Réding, 1798) (Figure 16) Murex magellanicus Chemnitz, 1788: 275, pl. 164, fig. 1570 (non binom.). Neptunea magellanica Réding, 1798: 116. Triton cancellatum Lamarck, 1816: 4 Priene (Fusitriton) cancellatus (Lamarck).—Cossmann, 1903: LOY, fig. S. Argobuccinum (Fusitriton) cancellatus | (Lamarck).— Wenz, 1938: 1058, fig, 3022. Argobuccinum (Argobuccinum) magellanicum (Réd- ing).—Carcelles, 1944: 247, pl. 2, fig. 23. Argobuccinum (Fusitriton) magellanicum (Réding). — Carcelles and Williamson, 1951: 286. ; Fusitriton cancellatus (Lamarck).-Smith, 1970: 475, pl. 42, figs. 4-10. Fusitriton magellanicus (R6ding).—Cernohorsky, 1977: 107, fig. 3. Material Examined: 20 specimens: T/150, T/171, T/178, T/214, T/315, T/340, T/511, T/512, T/572, T/971, T/1100, T/1282, T/1360, T/1575, T/1731, T/2026, T/2039, T/2169, T/2214, T/2252. Measurements: 73 mm. Figured specimen (T/2026) height Occurrence: Recent: Los Vilos to Magellan Region and Juan Fermdandez Archipelago; Plio-Pleistocene: Arauco. Remarks: This is a well known species that clearly does not present any taxonomic problems. Genus Sassia Bellardi, 1872 Sassia leucostomoides (Sowerby, 1846) (Figure 17) Triton leucostomoides Sowerby, 1846: 240, pl. 4, fig. 64. Sassia leucostomoides (Sowerby).—Frassinetti, 1997: 136, pl. 1. figs. 11-14; Griffin and Nielsen, 2008: 49, pl. 21, figs. 9-14. Material Examined: Six specimens: T/310, T/574; T/2032, T/2153, T/2196, T/2215. Measurements: Figured specimen (T/574) height 44.4 mm. Occurrence: Recent: Extinct; —Plio-Pleistocene: Arauco to Isla Guafo. Remarks: This species was originally described from Guato (Sowerby, 1846) and this fecond is the first from another locality. Family Muricidae Rafinesque, 1S15 Genus Trophon Monttort, 1810 Trophon geversianus (Pallas, 1774) (Figure 1S) Buccinum geversianus Pallas, 1774: 33 Murex magellanicus Gmelin, 1791: 3548. Fusus geversianus (Pallas)—Hupé, 1854; 167. Trophon geversianus (Pallas) —Gould, 1852: 227, pl. 6, fig. 297 , pl. 3, figs. 1, 2. 22/. Material Examined: Nine specimens: T/513, 1/569, T/575, T/640, T/798, T/1361, T/1574, T/2029, T/2253. Measurements: Height 62 mm (1/640), figured spec- imen (T/1361) 39 mm. Occurrence: — Recent: Chiloé to Magellan Region; Plio-Pleistocene: Arauco., Remarks: The Chilean species of the genus Trophon present a high intraspecific variability, with many species described as synonyms of T. geversianus. A detailed re- vision of the genus Trophon, including T. geversianus, was recently provided by Pastorino (2005b). Genus “Xymenopsis” Powell, 1951 “Xymenopsis” cf. “X.” dispar (Rochebrune and Mabille, 1889) (Figure 19) Fusus dispar Rochebrune and Mabille, 1589: H57, pl. 2 fig. 3. Trophon dispar (Rochebrune and Mabille).— and Williamson, 1951: 289, Xymenopsis dispar (Rochebrune and Mabille)—Powell, 1951: 159. Carcelles Material Examined: 16 specimens: T/352, T/586 (5), T/588 (7), T/1357, T/2219, T/2245. Measurements: Figured specimen (1/352) 22 mm. Occurrence: Recent: Tierra del Fuego; — Plio- Pleistocene: Arauco. Remarks: — Pastorino and Harasewych (2000) stated that this species does not belong in Xymenopsis but did not provide a more appropriate generic-level placement in the Muricidae. Genus Chorus Gray, 1847 Chorus giganteus (Lesson, 1830) (Figure 20) Monoceros giganteus Lesson, 1830; 405, N° 165, pl. 11, fig. 4. Figures 14-25. Gastropoda and Scaphopoda. 14. Epitonium (Nitidiscala) magellanicus (Philippi, 1845) [height 25.0 mm]. 15. Euspira guamblinensis Frassinetti and Covacevich, 1995 [height 22.5 mm]. 16. Fusitriton magellanicus (Réding, 1798) [height 73.0 mm]. 17. Sassia leucostomoides (Sowerby, 1846) [height 44.4 mm]. 18. Trophon geversianus (Pallas, 1774) [height 62.0 mm]. 19. “Xymenopsis” ct X.” dispar (Rochebrune and Mabille, 1889) [height 22.0 mm], 20. Chorus giganteus (Lesson, 1830) [height 95.0 mm]. 21. Nassarius taeniolatus (Philippi, 1845) [height 11.0 mm], 22. Adelomelon ancilla | Ligthfoot, 1786) [height 142.0 mm], 23. Bela paesleri Strebel 1905 [height 16.0 mm]. 24. Scaphander interruptus Dall, 1889 [height 18.5 mm], 25. Dentalium sp. [length 72.0 mm| Page 210 THE NAUTILUS, Vol. 122, No. 4 Monoceros giganteus? Lesson.—MGricke, 1896: 564. Rapana ( (Chorus) gigantea (Lesson). —Wenz, 1941: 1038, fig. 3080. Chorus giganteus (Lesson).—Carcelles, 1954: 271, pl. 5, figs. 1-11; Herm, 1969: 135, pl. 15, figs. e 4b; DeVries, 1997: 132, pl. 1, figs. 1-4, pl. 4, fig. 4. Chorus grandis (Philippi). -~Herm, 1969: pl. 15, figs. Material Examined: 17 specimens: T/576, T/985, T/1174, T/1193, T/1904—1905, T/1907—-1908, T/1910, T/1984, T/2170-2172, T/2208, T/2213, T/2272, one specimen without number. Measurements: Height up to 95 mm (T/9S5), figured specimen (T/1905) 50. 5 mm. Occurrence: Recent: Papudo to Calbuco; Plio-Pleis- tocene: southern Peru to Arauco. Remarks: The species of Chorus have been reviewed by Herm (1969) and DeVries (1997). DeVries (1997) also gave new diagnoses and descriptions. Regarding the stratigraphic range of Chorus giganteus cited by DeVries (1997), its presence in the Tubul Formation implies a maximum age of very late Pliocene to early Pleistocene for these deposits. Family Nassariidae Iredale, 1916 Genus Nassarius Duméril, 1806 Nassarius taeniolatus (Philippi, 1845) (Figure 21) Buccinum taeniolatus Philippi, 1845: 69; Hupé, 1854: 207, pl. 4, fig. 9. Nassarius taeniolatus (Philippi)—Keen, 1971: 609, fig. 1313 Material Examined: Three specimens: T/586 (2), T/1687. Measurements: 11 mm (T/1687). Occurrence: Recent: Papudo to Chonos Archipelago; Plio-Pleistocene: Arauco. Remarks: This species is the only representative of Nassarius in southern Chile, while two more species are found living off central and northern Chile (N. gayi (Kiener, 1835) and N. dentifer (Powys, 1835); see Mar- incovich, 1973) Family Volutidae Rafinesque, 1815 Genus Adelomelon Dall, 1906 Adelomelon ancilla (Ligthfoot, 1786) (Figure 22) Voluta ancilla Lightfoot, 1786: 84. Voluta gracilis Wood, 1528: 59, pl. 3, fig. 2. Cimbiola ancilla (Lightfoot)—Pace, 1902: 28, pl. 7, figs. 1-16 Adelomelon (Adelomelon) ancilla 1938: 1347, fig. 3816. Adelomelon ancilla (Lightfoot).—Dall, 1906; 143. (Lightfoot).—Wenz, Material Examined: One specimen: T/1227 Measurements: Figured specimen (T/1227) 142 mm. Occurrence: Recent: Chiloé to Magellan Region; Plio-Pleistocene: Arauco to ?Isla Guafo. Remarks: It is unclear if A. reconditus pecan 1997, is a different species or a juvenile A. ancilla. large specimen of presumably A. reconditus was found on ‘Cua (coll. Nielsen), but preservation does not allow assignment to a species. The Chilean fossil species of Adclomelan were recently revised by Nielsen and Fras- sinetti (2007a). Family Turridae H. and A. Adams, 1853 Genus Bela Leach in Gray, 1847 Bela paesleri Strebel, 1905 (Figure 23) Bela paessleri Strebel, 1905: 588, pl. 22, figs. 35, 35a—b. ae lia (Bela) paessleri (Strebel).—C sarcelles. 1950: 67. Material Examined: T/590 (7), T/1626. Measurements: 14 specimens: T/589 (6, in part), Height 16.7 mm (T/1626). Occurrence: Recent: Valdivia to Magellan Region; Plio-Pleistocene; Arauco, Remarks: This species is in need of revision, but be- cause the protoconch is not preserved in these speci- mens, it is not possible to verify its generic position. Family Scaphandridae Montfort, 1510 Genus Scaphander Montfort, 1510 Scaphander interruptus Dall, 1889 (Figure 24) Scaphander interruptus Dall, 1889, 12; 297, pl 12, fig. 12. Material Examined: Six specimens: T/244, T/583, T/584 (2), T/975, one specimen without number. Measurements: 18.5 mm. Figured specimen without number, Occurrence: Recent: Panama to Magellan Region; Plio-Pleistocene: Arauco. Remarks: The Recent and fossil Chilean Cephalaspi- dea are in need of revision. Scaphander cosmophilus (Sowerby, 1546) is known from the Pliocene of Isla Guafo (Frassinetti, 2000) and possibly from Isla Guam- blin (Frassinetti and Covacevich, 1995). Scaphander cos- mophilus was placed in the Genus Kaitoa by Griffin and Nielsen (2008). It is not clear if the two species are closely Aen Class Scaphopoda Bronn, 1562 Family Dentaliidae Gray, 1847 Genus Dentalium Linnaeus, 1758 Dentalium sp. (Figure 25) S. N. Nielsen and C. Valdovinos, 2008 Material Examined: Two specimens: T/1937-1938. Measurements: 2mm. Figured specimen (T/1937) length Remarks: Dentaliidae are well represented in Chile at least since the Miocene (see e.g., Philippi, 1SS87; Herm, 1969). A review of the group has never been under- taken. ASSOCIATED MACROFAUNA Several taxa outside the mollusks are known from the Tubul Formation. The Bir6 collection contains about 225 specimens of barnacles, remains of two decapod crabs, bryozoans, about 260 specimens of the brachio- pod Mage llania venosa Solander, 1786, two different echinoids and some shark teeth. Some of the fauna represented in the collection does not come from the Tubul Formation but from the Mio- cene Ranquil Formation, most probably from the nearby locality Punta El Fraile (see Nielsen et al., 2004; Finger et al., 2007). Miocene representatives in the Tubul col- lection include the nautiloid cephalopod Aturia cubaen- sis (Lea, 1S41) and the olivid gastropod Lamprodomina dimidiata (Sowerby, 1846). COMPARISON WITH OTHER FAUNAS AND CONCLUSIONS U — the Tubul Formation is the late Miocene to early Pliocene Ranquil Formation (Garcia, 1968; Pineda, 1986: Finger et al., 2007), the fauna of which is similar to that of the Navidad Formation south of Valparaiso (see Philippi, 187; Groves and Nielsen, 2003; Nielsen, 2004, 2005; Nielsen and Frassinetti, 2003; Nielsen et al., 2004). This fauna has been reworked and is of early to middle Miocene age (DeVries and Frassinetti, 2003; Finger et al., 2007) and contains many tropical to sub- tropical gastropod genera, such as Nerita, Strombus, Xenophora, Distorsio, Echinophoria, Ficus, Terebra, and Architectonica (Philippi. 1887; Covacevich and Frassinetti, 1980: Nielsen, 2005; Nielsen and DeVries, 2002; Nielsen and Frassinetti, 2007b), that disappeared from Chile during the late Miocene climate cooling. A number of the gastropod genera described herein (Epitonium, Trophon, Chorus, Nassarius) were already present in Chile during the Miocene, but were repre- sented by different species. If and how these are related to the living taxa has yet to be investigated. The Pliocene to Pleistocene faunas from northem Chile have a different composition than that of the Tubul For- mation, containing abundant Trochoidea, Fissurella, Tur ritella, Crucibulum, Trochita, Crepidula, different species of Naticidae (including Sinum cymba), Argobuccinum, Nassarius, Acanthina, Chorus, and Oliva peruviana (Herm, 1969: DeVries, 1997, 2003; own data), represent- ing today’s coastal fauna of northem to central Chile. However, these are nearshore faunas with many taxa from Page 21] rocky coasts, while that of Tubul is a soft-bottom fauna from slightly deeper water, missing typical rocky-shore elements such as Fissurella, hosnihiana: and Concholepas, which are present in the Plio-Pleistocene deposits of northem Chile and in the modern fauna along the whole Chilean coast (McLean, 1984; DeVries, 1995, 2003). The Tubul fauna has previously been compared to faunas coming from Guamblin and Guafo islands farther south (Frassinetti, 1997, 2000: Frassinetti and Covace- vich, 1995; Table 1). The fauna from Guamblin is close to the Tubul fauna and Frassinetti and Covacevich (1995) and Frassinetti (2000), comparing both faunas directly, cited new Tubul species (Epitonium biroi, Tro- phon covacevichi, and Hindsiclava ignorata), and sug- gested that Epitonium biroi may be conspecific with E. (Nitidiscala) magellanicus, Trophon covacevichi: may be conspecific with Trophon geversianus, and Hindsi- clava ignorata may be conspecific with Bela paesleri. However, to confirm these synonymies more material should be directly compared. The fauna described by Frassinetti (1997, 2000) from Guatfo contains a mixture of species also present at Guamblin island and Tubul and species known from older deposits, e.g., Panopea chiloensis, Incatella chilen- sis, Echinophoria sp. (cited as Semicassis sp.), Chorus doliaris, Penion spp., and Gemmula subaequalis. It seems likely that these taxa represent a fauna intermediate in age between the older faunas and the Tubul-Guam- blin fauna but it is possible that specimens of the older fauna have been reworked from older beds and incorporated into younger sediments as has been demonstrated for the Navidad, Ranquil and Lacui for- mations of Mio-Pliocene age (Finger et al., 2007). The presence of Chorus doliaris and Incatella chilensis suggests a latest Miocene age for the Guafo fauna (DeVries, 1997, 2007). The first appeareance of Chorus giganteus in south- erm Peru during the latest Pliocene (DeVries, 1997) makes this a likely maximum age for the Tubul fauna. Indeed, this agrees well with strontium isotope stratigra- phy data from calcitic shells of Zygochlamys patagonica coming from the same locality, which confirm an early Pleistocene age (unpublished data). The two apparently extinct species present, Euspira guamblinensis and Sas- sia leucostomoides, may well be found living off Chile in the future. Comparison with living ranges of the species found in the Tubul Formation ( Figure 1) permits one to identify two regions with 16 species in common, The fauna of the Tubul Formation resembles most closely the present-day faunas of northern Chiloé island and the Magellan Region, whereas northern faunas of Pliocene to Pleistocene age have a composition similar to that of modern northern and central Chile. Water temperatures for the Arauco area are accordingly interpreted to have been colder than today. It is pre ssently not pe ‘ the more northern species (Figure 1, numbers 6, 15, 16 previously tolerated cooler waters or if the cabo ‘mm spe- cies (Figure 1, numbers 7, 11, 14) tolerated warmer waters. However, it is possible that an additional factor THE NAUTILUS, Vol. 122, No. 4 Table 1. Comparison of mollusk species found at Tubul, Guafo and Guamblin. Tubul Guafo Guamblin Bivalvia Ennucula grayi x ? (barrosi) Tindariopsis x x sulculata Malletia chilensis x ? (sp.) Mytilus sp. x Zygochlamys x ? (cf. hupeanus) (tenuicostatus ) patagonica Lucinoma sp. x Cyclocardia x x (volckmanni) (volckmanni) velutinus Macoma inornata (tubulensis) Darina solenoides x (Mactra sp.) Ensis macha x x (sp.) Retrotapes x x (araucana) x (araucana) exalbidus Pandora cistula (sp.) Panopea x chiloensis Gastropoda Epitonium x ? (biroi) ? (biroi) magellanicus Turritella chilensis Euspira x x ; guamblinensis Echinophoria sp. x (Semicassis) Fusitriton x magellanicus Sassia x leucostomoides Cymatium remotum Trophon x ? (covacevichi) — ? (sp.) geversianus Trophon parcus x ee nopsis” cf. x ? (huilliche) X.” dispar Xymenopsis hero Chorus giganteus Chorus doliaris Nassarius taeniolatus Nassarius gayi Phos chilensis Penion spp Adelomelon : ? (reconditus ) ancilla Bela paesleri Hindsiclava ignorata Cryptogemma senex Gemmula subaequalis Scaphander x ? (cosmophilus ) interruptus Scaphopoda Dentalium sp : ? (sp ? (cosmophilus) independent of water temperature is responsible for these widely separated species triplets today in con- trast to their common occurrence during the early Pleistocene. ACKNOWLEDGMENTS First of all we express our gratitude to Profesor Lajos Biro-Bagoczky (1929-1993) who, together with others, assembled this collection during more than 15 years. Arturo Quinzio (Departamento Ciencias de la Tierra, Universidad de Concepcion, Chile) made this work possible by giving access to this collection. Klaus Bandel (Geolc gisch-P Paliontologisches Institut, Universitit Hamburg, Germany) is Hhanlked for discussion on the Tubul Formation and fauna. Daniel Melnick (GFZ Potsdam) helped with the DEM map for the distribution figure. Comments and suggestions by Tom DeVries (Burton, USA) improved the manuscript. 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Index Testaceologicus: or a Catalogue of Shells: with the Latin and English names, references to figures and places where found, ... Supplement, ... (References from Lamarck’s “Animaux sans Vertebres,” adapted to the figures in the ... “Index”, ...). London. iv + 59 pp. THE NAUTILUS 122(4):217-227, 2008 Radula morphology in veined rapa whelks, Rapana venosa (Valenciennes, 1846) (Gastropoda: Muricidae) from Chesapeake Bay, USA Juliana M. Harding" Stefanie M. Gera Roger Mann Department of Fisheries Science Virginia Institute of Marine Science Gloucester Point, Vi irginia 23062 USA jharding@vims.edu ABSTRACT Radula length, width, number of transverse rows of teeth, and rachidian tooth dimensions (central cusp height, central cusp base width, and rachidian tooth base width) were examined in relation to leined rapa whelk shell length. Radula length and width increase linearly with whelk shell length. The number of transverse rows of radular teeth increase with whelk shell length. Within an individual, central cusp height of the rachi- dian tooth increases with increasing distance from the anterior of the radula. Central cusp height of the rachidian tooth, an indicator of tooth wear or use, was least for teeth in rows 1 and 11. Teeth in radular row 21 appear to be in a transition zone from high to low wear or use. Within a radula and within a size class. the ratio of central cusp base width to rachidian tooth base width does not change. Central cusp base width, central cusp height and rachidian tooth base width were significantly smaller in females than in males indicating sexual dimorphism in rachidian tooth shape for rapa whelks. Patterns of wear as indicated by central cusp base width to central cusp height ratio values were not significantly different between sexes and may serve as an indication that feeding strategies and/or prey may be similar between animals of different sex but similar size. Additional Keywords: Neogastropoda, rachidian teeth, allometry, ontogeny INTRODUCTION The radula is a chitinous ribbon-like series of nearly colorless transverse tooth rows resting atop the radula membrane (Wu, 1965; Radwin and Wells, 1968). Muri- cid gastropods use the anterior teeth when drilling holes in bivalve prey (Carriker, 1961, 1981; Fujioka, 1985). As anterior teeth are worn down, they are replaced by younger teeth that are formed in the radular sac and gradually moved forward along the radula (Isarankura ; : * Author for correspondence and Runham, 1968; Carriker, 1981). Muricid radulae have between 100 and 500 transverse rows of teeth (e.g., Carriker, 1961; Radwin and Wells, 1968; Fujioka, 1985). Each transverse row of teeth consists of a central rachi- dian (R) tooth and two slender marginal teeth (M) in the tooth formation M + R + M (Carriker, 1969). The cen- tral rachidian tooth in each transverse row is responsible for most of the rasping and physical shell removal during drilling while the marginal teeth synchronously tear fle sh from prey (Carriker, 1969: Carriker et al., 1974; Krutak, 1977). Thus, the rachidian teeth show more wear, or reduction in size with use, than marginal teeth found in the same transverse rows (Carriker et al., 1974). This trend is particularly evident at the anterior end of the radula where the rachidian cusps in the most anterior row(s) may be completely removed by use (Carriker, 1969, 1974; Fujioka, 1985). Veined rapa whelks (Rapana venosa, Valenciennes 1846, Muricidae) are predatory marine gastropods that. while originally native to Japanese and Korean waters (Tsi et ale 1983), have successfully invaded marine and estuarine habitats in the Black, Adriatic, Aegean, Medi- terranean (Mann et al., 2004), and North Seas (Vink et al., 2005) as well as the Rio de la Plata (Pastorino et al., 2000) and Chesapeake Bay, USA (Harding and Mann, 1999). At the present time, the Ches sapeake Bay rapa whelk population is the only known population of bs whelks in North America. Rapa whelks provide an unusual opportunity to investigate allometric changes in radula morphology across a wide size range of individuals mee they reach terminal shell lengths in excess of 170 mm (Wu, 1988: Harding and Mann, 2005). Like other muricids (Paine, 1966), rapa whelks experience ontogenetic shifts in diet (Harding and Mann, 2001) as well as predation strategy and resulting predation signatures in prey valves ( Harding et al.. 2007). Small ( shell length) rapa whelks drill their prey (Harding and Mann, 2001; Harding et al., 2007) including barnacles 35 mm Page 215 THE NAUTILUS, Vol. 122, No. 4 (Balanus sp., Chthamalus sp.), mussels (Mytilus sp., Geukensia demissa), soft shell clams (Mya arenaria), and oysters (Crassostrea virginica). At shell lengths above 35 mm, rapa whelks eat larger bivalves (Harding and Mann, 2001) including oy sters and northern quahogs (Mercenaria mercenaria) and typically either edge bore their prey or leave no signatures (Morton, 1994: Harding et al., 2007). We quantitatively describe radula and rachidian tooth morphology for a size range of rapa whelks from Chesa- peake Bay, USA. Rachidian teeth in Rapana have a large central cusp flanked by two smaller cusps (Avalawa: 1964: Wu, 1965). Shell length, the maximum dimension from the tip of the spire to the bottom of the siphonal canal, is used as the metric of whelk size. Shell length does not fluctuate with season or other factors. Relation- ships between shell length and radula dimensions are quantitatively désoribed: for male and female rapa whelks. Within each radula, rachidian tooth morphology is described along the length of the radula by measuring ratios of rachidian tooth central cusp base width to cen- tral cusp height and central cusp base width to rachidian tooth base width. The resulting ratios are compared between teeth along the length of an individual radula and across radulae from male and female whelks as well as from a size range of Chesapeake Bay rapa whelks. MATERIALS AND METHODS Rapa whelks with shell lengths (SL) in excess of 70 mm were obtained from the lower Chesapeake Bay, USA as donations to the Virginia Institute of Marine Science (VIMS) rapa whelk bounty program. Rapa whelks less than 70 mm SL were cultured at VIMS, Gloucester Point, Virginia to supplement the lower SL range of rapa whelks because individuals less than 70 mm SL were not available through the bounty program. At the time of whelk collection, SL was measured in mm and whelks were assigned to shell length classes that were established to evenly categorize the potential SL range (1-180 mm SL). Petite, small, medium, and large classifications corresponded to whelk SL ranges : less than 45 mm, 45.1-90 mm, 90.1-135 mm, and 5.1-180 mm, respectively. “Whe Iks were frozen after collection and thawed to facilitate dissection and removal of radulae. Whelks were sexed during dissection and distinguished 7 male or female on the basis of penis length and gonad color alter Mann et al. (2006). For the purposes of discussion herein, true females (penis length = 0 mm, bright yellow gonad) and imposex females (penis length - -20) mm, bright ye low gonad) are groupe dl toge shes per Mann et al. (2006) the whelk probosc is. However, two control radulae were removed by soaking the proboscis for 24 hours in 10% sodium hydroxide to ensure that dissection removed the entire radula intact. Only intact radulae were used this study Typic: ally, radulae were dissected out of After removal from whelks, images of the complete radula were taken using a digital camera mounted on a dissecting microscope for measurement of total radula length, total anterior to posterior distance (mm) and rad- ula width, the maximum lateral distance across the bases of the marginal and rachidian teeth at the first transverse row of teeth (Figure 1). The odontophore was removed and then the rachidian teeth were systematically removed from every 10th transverse tooth row (e.g., Row 1, 11, 21 etc.) ) along the radulae moving from ante- rior to posterior (Figures 2 and 3). Tooth rows were removed with a size 10 scalpel blade for larger indi- viduals, and with sharpened needles for cultured individuals less than 66 mm SL. Digital images were taken of each individual rachidian tooth after removal with the tooth positioned convex side down. Typical magnification of individual teeth used for digital images ranged from 50 for whelks with SL greater than 147 mm to 90 for whelks less than 45-50 mm SL. Measurements (mm, Figure 4) of the rachidian tooth central cusp base width (L1), maximum central cusp height (L2) measured from the tip of the central cusp to the midpoint of L1, and the maximum rachidian tooth base width (13) were made on the resulting images. The terminology used to describe tooth morphology follows that of oak! 1993). Data Analyses: Significance levels for all statistical tests were set at alpha = 0.05 a priori. Fishers multiple comparison tests were used for post hoc comparisons when appropriate. Radula Allometry and Gross Morphology: — Linear and power regression models were used to describe relationships within sexes between rapa whelk shell length and radula length, radula width, and number of rows of transverse teeth per radula. The relationship between radula length and both radula width and the number of transverse rows of teeth within radulae from males and females were also examined with both linear and power regressions. The slopes of all morphological regressions \were compared between sexes with t tests (per Zar, 1996) on raw data or on logarithm transformed data if the power model was deemed more appropriate than the linear model. Rachidian Tooth Dimensions: The relationship bet- ween central cusp base width (L1, Figure 4) and size class of the whelk from which it came was evaluated with a three-way ANOVA (whelk size class x tooth row x sex) with the response being the maximum base width of the rachidian tooth. These data satisfied assumptions of homogeneity of variance after the logarithm. trans- formation was applie sd but not normality. The reli BHOUSHED be uh en rachidian tooth central cusp height (LL2, Figure 4), the size class of the whelk from which it came, and the tooth row was evaluated with a three factor ANOVA (hell size class tooth row sex). These data satisfied neither the assumption of normality nor homogeneity of variance regardless of the J. M. Harding et al., 2008 Page 219 Figures 1-3. Radula of veined rapa whelk. 1. Radula from an 84.2 mm shell length (SL) whelk showing general morphological features and orientation. 2. A rachidian tooth from row | of the radula from a 134 mm SL whelk showing wear. 3. An unworn rachidian tooth from row 41 from a 122 mm SL whelk. Scale bar = 1 mm. Abbreviations: r = row number; R = rachidian; Od = odontophore; A = anterior; P = posterior. transformation (logarithm, natural logarithm, square root, arcsine). The relationship between rachidian tooth base width L3, Figure 4), row number, sex, and size class of the whelk from which it came was evaluated with a three- way ANOVA (whelk size class x tooth row x sex). These data satisfied the assumption of homogeneity of variance without transformation but did not satisfy the assump- tion of normality regardless of the transformation (loga- rithm, natural logarithm, square root, arcsine) and were analyzed without transformation. The ratio of rachidian tooth central cusp base width L1) to central cusp tooth height (L2) was calculated for each rachidian tooth. Using a ratio that compares base width to tooth height is appropriate in a structure where both the base width and tooth height change along the length of the structure with ontogeny. Not only does the ratio allow for scaling when comparing individual teeth along the radula length, but it can also be used as an index of wear because width does not change with use In this case a central cusp base width: central cusp tooth height ratio >1 is indicative of wear. The first 81 rows were chosen for analysis because every radula dissected had at least 81 rows. These data satisfied neither the assumption of homogeneity of variance nor normality regardless of the transformation (logarithm, natural Page 220 THE NAUTILUS, Vol. 122, No. 4 Figure 4. Veined rapa whelk rachidian tooth with the mea- surements made in this study identified: rachidian tooth central cusp base width (L1), rachidian tooth central cusp height (L2), and rachidian tooth base width (L3). Scale bar = 0.25 mm. logarithm, square root, arcsine). A three-way ANOVA (whelk size class x tooth row number x sex) was used to evaluate the ratio of central cusp width to central cusp height (L1:L2). The ratio of central cusp base width (L1) to rachidian tooth base width (L3) for each rachidian tooth was eval- uated with a 3 way ANOVA (size class x row x sex) to describe potential changes in tooth shape with ontogeny. Data satisfied assumptions of homogeneity of variance without transformation but not nor staliy (either with or without transformation, e.g., logarithm, natural loga- rithm, square root, arcsine). acces in the L1:L3 ratio across whelk size classes reflect ontogenetic changes in tooth pam that may be related to sexual dimor- phism (Fujioka, 1982, 1984) and which may act to dis- perse relatively greater strike force during f feeding in larger whelks. RESULTS Only radulae from whelks collected when water tem- peratures were above 11-12°C and feeding were used (Harding, unpublished data). All radulae examined were intact. Descriptive morphological data were collected from 39 rapa whelk radulae. These radulae were from rapa whelks with shell lengths between 20.2 mm and 174 mm (Table 1, Figure 5). Rapa whelk radula lengths ranged from 4.33 to 51.05 mm with corresponding radula widths of 0.23 to 2.67 mm and total number of transverse rows of teeth of 89 to 210, respectively. Radula length was an average of 21.4% (standard error = 0.61%) of shell length. Radula Allometry and Gross Morphology: Regres- sion coefficients for the fitted linear and power regres- sion models used to describe relationships between rapa whelk shell length and radula morphology and between rapa whelk radula measurements are given in Table 2A. The linear model is suggested as a better oe of the relationship between rapa whelk shell length (SL) and radula length (RL; Figure 6A) for both sexes since the coefficients of determination from both models are identical (Table 2A) and the linear model provides the simplest description of the data. The slope for the SL— RL relationship in males is significantly higher than that for females (t-test, Table 2B). The power model more accurately described the rela- tionships between rapa whelk SL and radula width (RW; Figure 6B) by predicting a radula width equal to 0 at a shell length equal to 0. The coefficient of determination for the line sar model describing the relationship between shell length and the number of rows of radular teeth was higher (females = 0.63, males = 0.76, Table 2A) than that of the corresponding power model (females = 0.58, males = 0.69, Table 2A) for both sexes and the linear model predicted a positive number of rows of teeth at shell lengths of 0 mm (Figure 6C). The relationship between radula length (RL) and rad- ula width (RW) was described with a power model for both sexes (Table 2A, Figure 6D) which predicted a radula width of 0 at a radula length of 0 and had a higher coefficient of determination than the corresponding lin- ear model (Table 2A). The linear model describing the relationship between radula length (RL) and number of transverse rows of teeth has a higher coefficient of determination than the corresponding power model Table 1. . se id of ey whelks used in this study with basic statistics on radulae. Abbreviations used below are as follows: F = female, M = male, Avg = average for female and male whelks combined, SL = shell length, mm, SEM = standard error of the mean in miner ses, RL = radula le meth, mm, RW = radula width, mm; NRT = number of rows of teeth. # of Whelks Avg SL Ave RL Ave RW Avg NRT Avg RLSL % Whelk size class (F/M) (SEM) (SEM) (SEM) (SEM) (SEM) Petite (<45 mm SL) F=5 25.64 (2.64) 9.28 (0.56) 0.48 (0.18) 106 (6.07) 20.59 (0.58) M=5 34.72 (3.07) 7.62 (0.95) 0.71 (0.31) 122.20 (4.78) 21.72 (0.81) Small (45,1—-90 mm SL) F=5 70.38 (5.47) 15.93 (2.21) 0.97 (0.15) 133.80 (5.05) 92.38 (1.65) M=6 69.13 (6.21) 17.63 (1.92) 1.22 (0.13) 126.67 (3.86) 25.56 (1.79) Medium (90.1-135 mm SL) F = 4 104.75 (5.07) 22.32 (2.52) 1.47 (0.03) 137 (12.71) 21.37 (2.53) M=5 120.2 (5.3) 29.85 (1.58) 2.04 (0.10) 149.2 (7.19) 25 (1.51) Large (135.1-150 mm SL) F=2 153 (10.0) 35,7 (3,2) 1.99 (0.24) 177.5 (16.5) 23.3 (0.57) M=7 153.71 (4.47) 12 (2,23) 2.22 (0.10) 182.29 (6.38) 26.76 (1.21) J. M. Harding et al., 2008 Page 221 H@ Females © Males w Number of whelks Midpoint of rapa whelk shell length size class (mm) Figure 5. Shell length (mm) frequency distribution for the 39 veined rapa whelks whose radulae were examined in this study. (females = 0.78 vs. 0.71, males = 0.81 vs. 0.72, Table 2) and predicts a positive number of tooth rows at radula lengths of 0 mm (Figure 6E). Rachidian Tooth Dimensions: Rachidian tooth cen- tral cusp base width (Ll, mm; Figure 4) increased significantly with increasing whelk size class (Table 3, Figures 7A and 7D). There were no significant differ- ences in central cusp base width observed between rows within a size class for the 81 rows of teeth that were examined. Central cusp base width was. significantly larger in males than in females (Fisher's test, Table 3). Differences between male and female central cusp base width were particularly evident in the medium and large size classes (Figure 7A and 7D). The height of the rachidian tooth central cusp (L2, mm) varied significantly with size class, row number, and sex (ANOVA, Table 3, Figures 7B and 7E). In gen- eral, larger whelks have larger central cusp heights in rows 31 through 81 than whelks of other size classes. Within all size classes and both sexes, central cusp heights from rows | and 11] are significantly less than in rows 31 through 81 (ANOVA, Table 3, Figures 7B and TE). Central cusp heights from rows 1, 11 and 21 in large whelks are significantly different from central cusp heights in petite whelks but similar to cusp heights ob- served in rows | and 21 for medium whelks and row 11 for small whelks (ANOVA, Table 3, Figures 7B and 7E). Female whelks have significantly lower L2 values than male whelks (Fisher's test, Table 3, Figures 7B and 7E). Rachidian tooth base width (L3, mm) increases signi- ficantly with increasing whelk size class (ANOVA, z= o © ) a = a + “a * iss} 4 am = =r Be ae iary of regression statistics used to describe rapa whelk radula morphology and wear patterns. The linear model equation was y Table 2A. used for the power model was y = a X radula ? = standard ert coefficient; SE Abbreviations used below are as follows: coef RW = radula width, mm: NRT number of rows of teeth. man: h, Males Coef b (SE) Females Coet b (SE) p Value Regression F Statistic ) 4 Coeft a (SE R? p Value Regression F Statistic = Y 5 Ro Model Relationship Ooranno rst AawmMe tae as OARDMHSOD al GKeKEoOCKOCtTOSKS an 1.08 (0.08) 40.35 0.43 69 76 ANDnRoOnorse ORANAANDFAMS GHMHHBOrKS ADGONAMMNMV [oon oe | .29) P we) aA cD o> Oman amon oO DAADOOD OM on = 2 : cS = ac | s i _— aml ~ © F me ZF, iam jam xX al vx gv x g Ee - - - - m Sp) Sp) NY aa aa Power Page 222 THE NAUTILUS, Vol. 122, No. 4 60 A —e—Females * -no- Males 50 5 a o ra = 40 a ® s a = 30 ae © op o-" o 3 oe : & 20 oo" ; ~ “Ya ou 6 ; 10 i) 0 50 100 150 200 Radula width (mm) Radula width (mm) 0 50 100 150 200 0 10 20 30 40 50 60 290 Shell length (mm) 236 Radula length (mm) 200 = = e g 180 iE ; 160 ” wn 5 © 140 3 2 E E 420 Zz Zz 100 {’ 80 0 50 100 150 200 0 10 20 30 40 50 60 Shell length (mm) Radula length (mm) J. M. Harding et al., 2008 Page 223 Table 2B. Summary of t-tests comparing regression equa- tions for female and male whelks given in Table 2A that are recommended for descriptions of these re lationships. T-tests were performed for the power model using logarithm transformed data. Abbreviations are the same as those used in Table 2B above. Relationship Comparison Model p value SL vs. RL Female vs. Male Linear <0.05- SL vs. RW Female vs. Male Power >0.05 SL vs. NRT Female vs. Male Linear >0.05 RL vs. RW Female vs. Male Power >0.05 RL vs. NRT Female vs. Male Linear >0.05 Table 3, Figures 7C and 7F). However, no differences were observed in rachidian tooth base widths between radular tooth rows within a whelk size class and within a sex (ANOVA, Table 3, Figures 7C and 7F). Rachidian tooth base width was significantly larger in male whelks than in female whelks and this trend is particularly evi- dent in the medium and large size classes (Table 3, Figures 7C and 7F). Large whelks had significantly higher ratios of rachi- dian central cusp base width (Li) to central cusp height (L2) than all other whelk size classes (ANOVA, Table. 3: Figure 8). The first row of teeth in the radulae had significantly higher L1:L2 ratios than all other rows (ANOVA, Table 3 . Figure 8). The eleventh row of teeth also had an L1:L2 ratio ie was significantly higher than that observed in rows 21-81 (ANOVA, Table 3 Figure 8). Since central cusp base to height (L1:L2) ratios >1 are indicative of tooth wear, the anterior 1-11] rows of teeth are more worn than newer teeth occurring in rows 2] and higher. Within each size class and sex, a wide range of L1:L2 values was observed for row 1 and/or row 11 (Figure §). This variability was the result of one or two individual whelks per size class having very low central cusp heights ( extreme wear) in row 1 or row 11, the rows of teeth that are activ ely used in feeding. Patterns of wear as indicated by L1:L2 ratio values were not significantly different between sexes (Table 3, Figure 8). The ratio of central cusp base width to rachidian base width (L1:L3) was significantly affected by size class and sex (ANOVA, Table 3, Figure 9). Within a size class and within a sex, the ratio of central cusp base width to rachidian tooth width did not change significantly with row number. Male whelks had greater L1:L3 ratios than female whelks (Fisher's test, Table 3, Figure 9). DISCUSSION Radula length teeth in the radula increase with increasing rapa whelk floridana): radula width, and number of rows of shell length. Ontogenetic increases in radula length and the number of rows of teeth with shell length have also been documented for other muricid species (e.g. Stramonita floridana, Radwin and Wells, 1968 (as Thais Cronia margariticola and Morula musiva, Fujioka, 1954; Thais bronni and T. clavigera, Fujioka, 1985; Nucella lapillus, Kool, 1993). The relationship between rachidian tooth base width and central cusp base width also changes with ontogeny but does not change in relation to the anterior-posterior location on the radula. That is, within an individual and within a size class, rachidian teeth examined from rows | through 81 display similar scaling of central cusp base width to rachidian tooth base swith, Rachidian teeth in female rapa whelks tend to have smaller central cusp base width as well as tooth base widths when compared to male whelks within the same size class. Fujioka (1982, 1984) describes similar ontogenetic changes in rachidian tooth shape including an increase in earl cusp base width for Cronia margariticola, Morula musiva, and Dru- pella sp. in relation to sexual dimorphism. The observed ontogenetic changes in rachidian tooth shape may reflect morphological shifts designed to accommodate greater rachidian tooth strike force re sulting from the sealing of the buccal complex at increased w he Ik sizes. Pre sumably there is an ontogenetic scaling relationship in effect to optimize the force provided by the buccal mass muscula- ture and minimize the damage to rachidian teeth through use that is reflected in the shape of the tooth. Rachidian tooth wear, as indicated by the ratio of central cusp width (L1) to central cusp height (L2), decreases with increasing distance from the anterior (oldest) end of the radula. Rachidian teeth in the first 11 rows of the radula have central cusp heights that are less than central cusp heights in rows 21 through $1 in all size classes. Carriker et al. (1974) describe rachidian cusps that have been worn off leaving only the tooth base in the anterior rows of rachidian "teeth of Urosal- pinx cinerea folleyensis. In laboratory studies with Thais bronni and T. clavigera, Fujioka (1985) observed that rows of teeth worn by feeding yin 5-15% of the total number of radular rows. The whelk Acanthina spirata uses approximately 8 to 20 teeth in each rasping stroke as these whelks feed on mussels (Hemingway, 1975). These data are consistent with our observations for rapa whelks, where at least the Ist and 11th rows of the radula were used, and the 21st row acted as a transition between the part of the radula the whelk was actively using to feed and the more posterior section that was unused. The observed changes in rachidian tooth wear may reflect ontogenetic change ’s in predation strategy, diet, or possibly both. Differences in predation strategy are potentially reflected in the observed changes in Figure 6. Relationships for female (n = 16) and male (n = 23) veined rapa whelks ranging from 20.2 to 174 mm SL between shell length (SL) and radula length (RL, A), SL and radula width (B), SL and the number of rows of teeth (C), RL and radula width (D and RL and number of rows of teeth (E) with fitted regressions (female = solid, male = dashed) that were used to describe the relationships. Linear regression models are plotted for panels A, C, and E. Power models are presented in panels B and D Regression equations and descriptive statistics are given in Table 2 Page 224 THE NAUTILUS, Vol. 122, No. 4 (L1, mm; SEM) Average central cusp height Average central cusp base width (L2, mm; SEM) Average rachidian tooth base width (L3, mm; SEM) Figure 7. the mean, SEM) for { of the mean, SEM females (¢ @ Petite female (n=5) Medium female (n = 4) 9 4 0 Petite males (n= 5) » Medium males (n = 5) fF A. a Small female (n= 5) am Large female (n = 2) D. 4 Smalimales (n= 6) g Large males (n = 7) op ee ae ahah te tat 0.3 é foPoE gE Pe # 0.25 0.2 0.15 0.1 0 20 40 60 80 (0) 20 40 60 Rachidian tooth row number 80 Rachidian tooth row number Graphs of rachidian tooth row number in relation to average central cusp width (L1) with error bars (standard error of les (A) and males (D) from all size classes, average central cusp height (L2) with error bars (standard error males (B) and males (E) from all size classes, and average rachidian tooth base width (L3, SEM) for and males from all size classes J. M. Harding et al., 2008 Page 225 Table 3.) Summary of ANOVA results comparing tooth morphology across whelk size classes and rows within the radulae. Asterisks indicate statistical significance at an alpha value of 0.05. Abbreviations used below are as follows: 1= petite size class, 2 = small size class, 3 = medium size class, 4 = large size class. NA = Not applicable. Refer to Figure 4 for a description of L1, L2, and L3. Fisher's test Test Response Factors — p value results ANOVA. Central cusp Size class <0.01) 4>3>2>1 base Row 0.80 NA width (L1) Sex <0.01° Male > Female ANOVA. Central cusp Size class <0.01° 4,3>2>1 height (L2) — Row <0.01° 1,11 < 31-8] Sex <0.01° Male > Female ANOVA. Rachidian Size class <0.01) 4>3>2>1 tooth base Row 0.99 NA width (L3) Sex <0.01° Male > Female ANOVA Wear Size class 0.02" 4>3,2,4=1 (ratio L1/L2) Row <0.01" 1 > 21-81, 11> 31-81 Sex 0.16 NA ANOVA. Shape (ratio Size class <0.01° 4>2,3>1 LI/L3) Row 0.90 NA Sex <0.01° Male > Female rachidian tooth wear with ontogeny because the rachi- dian teeth are actively used during shell drilling. Meth- ods of feeding w hich require penetration of prey valve shells with the radula (e. g., drilling) will leave more wear on the rachidian teeth than non- drilling methods of at- tack. Therefore, examination of pathadian tooth wear along the radula and differences in wear depending on size “class may give an indication as to transitions in feeding strategies : different size classes of rapa whelks (Figure 8). High | evels of rachidian tooth wear (L1:L2 ratio >1) in rows 1 and 11 were associated with the petite and large size classes (Figure 8A). The smallest whelks (<45 mm SL) typically leave drill holes (i.e., wall bores) in the valves of their prey (Harding et al., 2007). Although large (>135 mm SL) rapa whelks do not always leave ae signatures in prey valves, edge bore signatures are left instead of drill holes (Harding, Kingsley-Smith, Mann, unpublished data) when signa- tures are present. The observed L1:L2 values for rows 1 and 11 in the large size class are driven by one male ee SB). It is possible that this individual had been using its radula to penetrate prey shells and that the other large whelks had not. Since the large whelks used herein were wild caught and had unknown feeding histories, we cannot say - with certainty. Relatively less wear (L1:L2 ratio <1.2 ) in row 11 was observed in rapa whelks with shell . of 45 to 135 mm (small and medium size classes) that do not typically drill their prey (Harding et al., 2007, Figure 8). Similar wear patterns (L1:L2 ratio) were observ edi for males and females and may serve as an indication that feeding strategies and/or prey may be simi- lar between animals of different sex but similar size. A. Females Petite ir = . n = Average [central cusp base width(L1)/central cusp height (L2)], SEM 6 o Petite (n=5 B.|Males a Small i = a} ° Medium (n= 5) Q Large (n=7) 5 4 3 2 & 6 we [3] oe ee Une 0 20 40 60 80 Rachidian tooth row number Figure 8. Rachidian tooth wear as indicated by the average ratio of rachidian tooth central cusp base width (L1) to central cusp height (L2) in relation to rachidian tooth row number for females (A) and males (B) from all size classes. If a whelk is using the radula to penetrate the shell, the chemical composition of the prey shell may affect the level of wear observed on the rachidian teeth. Oyster and mussel shells have more calcite and are relatively softer than other bivalves with predominantly aragonite shells (Carter, 1980). Drilling through aragonite shells has the potential to cause more wear on eachidk in teeth than shell Es tration of calcite shells. Examination of the radulae from rapa whelks using drilling to consume a species-specific diet is a topic for future research that would provide data to address changes in radula wear with regard to prey shell hardness. ‘ DIG Page 226 THE NAUTILUS, Vol. 122, No. 4 0.35 e Petite (n = 5) . a Small (n = 5) A. Females » Mediunntn=4y = Large (n = 2) 0.3 7 0.25 7 Pye! 0.2 0.15 0.35, |B./Males oO wo T rre—t HH HH HoH att fd t | Average [central cusp base width(L1)/rachidian tooth base width (L3)], SEM 02 o Petite (n = 5) : 4 Small (n =6) ° Medium (n = 5) 5 Large (n= 7) 0 20 40 60 80 Rachidian tooth row number Figure 9. Average ratio of central cusp base width (L1) to rachidian tooth base width (L3, with standard error of the mean, SEM) by rachidian tooth row for females (A) and males (B) from all size classes. ACKNOWLEDGMENTS This manuscript is dedicated to the late Dr. Melbourne R. Carriker, whose passion for and detailed work with Muricids serves as both a standard of excellence and an inspiration. Thanks are extended to all local fisher- men that have participated in the Virginia Institute of Marine Science rapa whelk bounty program since its inception in 1998. Melissa Southworth, Ethan Jestel, Catherine Ware, Erica Westcott, Steven Goodbred, David Kerstetter, Peter Kingsley-Smith, Amy Bohannon, Meredith Fagan, Meghan Harris, Rhonda Howlett, Rae Marie Johnson, Courtney Harris, Karen Capossella, John Hansen, and Matthew Robinson assisted with whelk collections from local fishermen. Drs. Greg Capelli, Randy Chambers, John Kraeuter, Ms. Melissa Southworth, and an anonymous reviewer provided valu- able comments on earlier versions of this manuscript. This work was completed in partial fulfillment of the requirements for an undergraduate Honors degree (SMG) from the Department of Biology, College of William and Mary, Williamsburg, Virginia. This is contri- bution number 2928 from the Virginia Institute of Marine Science, Gloucester Point, Virginia. LITERATURE CITED Arakawa, k. 1964. A study on the radulae of the Japanese Muricidae (2): The genera Vexilla, Nassa, Rapana, Murex, Chicoreus, and Homalocantha. Venus 22: 355-364. Carriker, M. R. 1961. Comparative functional morphology of boring mechanisms in gastropods. American Zoologist 1: 263-266. Carriker, M. R. 1969. Excavation of boreholes by the gastro- pod, Urosalpinx: an analysis by light and scanning elec- tron microscopy. American Zoologist 9: 917-933 Carriker, M. R., J. G. Schaadt, and V. Peters. 1974. Analysis by slow-motion picture photopograph and scanning electron microscopy of radular function in Cl Irosalpinx cinerea follyensis (Muricidae, Gastropoda) ) during shell penetra- tion. Marine Biology 25; 63-76, Carriker, M. R. 1981. Shell penetration and feeding by natica- cean and muricacean predatory gastropods: a synthesis. Malacologia 20; 403-422. Carter, J. 1980. Environmental and biological controls of bi- valve shell mineralogy and microstructure. In: Rhoads, D and R. Lutz (Eds.) Skeletal Growth of Aquatic Organisms. Plenum Press, New York, pp. 69-113. Fujioka, Y. 1952. On the secondary sexual characters found in the dimorphic radula of Drupella (Gastropoda: Muricidae) with reference to taxonomic revision. Venus 40; 203-223. Fujioka, Y. 1984. Sexually dimorphic radula in Cronia mar- gariticola and Morula musiva (Gastropoda: Muricidae). Venus 43: 315-330. Fujioka, Y. 1955. Seasonal aberrant radular formation in Thais bronni (Dunker) and T. clavigera (Kuster) (Gastropoda: Muricidae). Journal of Experimental Marine Biology and Ecology 90; 43-54. Harding, J. M. and R. Mann. 1999. Observations on the bio- logy of the veined rapa whelk, Rapana venosa (Valenci- ennes, 1546) in the Chesapeake Bay. Journal of Shellish Re ees 18: 9-17. Harding, J. M. and R. Mann. 2001, Growth rates of larval and juvenile veined rapa whelks Rapana venosa, from Chesa- peake Bay, USA, from hatch through age 1. International Conference on Marine Bioinvasions, 9-11 April 2001, New Orleans, LA, USA (Abstract). Harding, J. M. and R. Mann. 2005. Veined rapa whelk (Rapana venosd) range extensions in the Virginia waters of Chesapeake Bay, USA. Journal of Shellfish Research 24: 3581-355. Harding, J. M., P. Kingsley-Smith, D. Savini, and R. Mann. In press. Comparison of predation signatures let by Atlantic oyster drills (Urosalpinx cinerea Say, Muricidae) and veined rapa whelks (Rapana venosa Valenciennes, Muri- cidae) in bivalve prey. Journal of Experimental Marine Biology and Ecology. SSS SSS SSS J. M. Harding et al., 2008 Page 227 Hemingway, G. T. 1975. Functional morphology of feeding in the predatory whelk, Acanthina spirata, Gastropoda: Prosobranchia). Bulletin of the American Malacological Union, Inc. pp. 64-65 (Abstract). Isarankura, K. and N. W. Runham. 1968. Studies on the re- placement of the gastropod radula. Malacologia 1: 71-91. Kool, S. P. 1993. The systematic position of the genus Nucella (Prosobranchia: Muricidae: Ocenebrinae). The Nautilus 107: 43-57. Krutak, P. R. 1977. Gastropod radulae: their potential in the fossil record. Transactions of the Gulf Coast Association of Geological Societies 27: 314-322. Mann, R., A. Occhipinti, and J. M. Harding. 2004. Current status of global invasions by the marine gastropod Rapana venosa. First special report of the International Council for Exploration of the Seas on the status of introduced species. International Council for Exploration of the Seas/ICES. 24th Annual meeting of ICES (WGITMO), Goteberg, Sweden, March 20-21, 2002. Mann, R., J. M. Harding, and E, Westcott. 2006. Occurrence of imposex and seasonal patterns of gametogenesis in the invading veined rapa whelk Rapana venosa from Chesa- peake Bay, USA. Marine Ecology Progress Series 310: 129-138. Morton, B. 1994. Prey preferences and method of attack by Rapana_ bezoar (Gastropoda: Muricidae) from Hong kong. pp. 309-325. In: B. Morton (Ed.). The malacofauna of Hong Kong and Southern China III. Hong Kong University Press, Hong Kong. Paine, R. 1966. Function of labial spines, composition of diet, and size of certain marine gastropods. The Veliger 9: 17-24. Pastorino, G., P. E. Penchaszadeh, L. Schejter, and C. Bremec. 2000. Rapana venosa (Valenciennes, 1846) (Mollusca: Muricidae): A new gastropod in south Atlantic waters. Journal of Shellfish Research 19: 897-899. Radwin, G. E. and H. W. Wells. 1968. Comparative radular morphology and feeding habits of Muricid Gastropods from the Gulf of Mexico. Bulletin of Marine Science 18: 72-85. Tsi, C. Y., X. T. Ma, Z. K. Lou, and F. S. Zhang. 1983. Ilustra- tions of the fauna of China (Mollusca), Vol 2. Science Press, Beijing. Vink, R., D. Nieweg, and H. Post. 2005, Rapana venosa (Valen- ciennes, 1846): a new invasive species for the Netherlands (and England?). Spirula 345: 152-155 (In Dutch with English abstract). Wu, S. kK. 1965. Studies of the radulae of Taiwan muricid gastropods. Bulletin of the Institute of Zoology, Academia Sinica 4: 95-106. Wu, Y. 1988. Distribution and shell-height relation of Rapana venosa Valenciennes in the Laizhou Bay. Mar. Sci./ Haiyang Kexue 6: 39-40. Zar, J. 1996. Biostatistical analysis. 3rd edition. Prentice Hall, New Jersey. THE NAUTILUS 122(4):228-235, 2008 Page 228 A new species of Sphaerium Scopoli, 1777, from southern Brazil (Bivalvia: Sphaeriidae) Maria Cristina Dreher Mansur Av. Arlindo Pasqualini, 410 91760-140 Porto Alegre, BRAZIL mcmansur@terra.com.br Sommergasse 10 GERMANY Claus Meier-Brook D-72119 Ammerbuch-Reusten Cristian Ituarte Museo Argentino de Ciencias Naturales Ay. Angel Gallardo 470, C1405DJR, Buenos Aires ARGENTINA ituarte@mail.retina.ar ABSTRACT Sphaerium cambaraense new species is described based on samples collected in the beginning of the summer near the headwaters of Taquari River (Jacui River Basin) on the basaltic plateau of southern Brazil. This is the first record for the genus Sphaerium in Brazil and in South America outside the Andes. Sphaerium cambaraense is characterized by a relatively large and solid shell, a high triangular shell outline and a solid hinge plate. Compared with Sphaerium forbesi (Philippi, 1869), S. cambaraense has a more strongly triangular shell outline and beaks not prominent. Sphaerium lauricochae (Philippi, 1869), another similar species from Bolivia, Chile, and Peru, has a more rounded shell outline. Additional Keywords: Freshwater, Rio Grande do Sul, South America INTRODUCTION According to Dreher-Mansur and Meier-Brook (2000), the family iene is represented by two subfamilies: Euperinae, including the genera Eupera Bourguignat, 1854, and Byssanodonta rt Orbigny, 1846; and Sph- aeriinae with three genera, Sphae rium Scopoli, 1777, Musculium Link, 1807, and Pisidium Pfeiffer, 1821. Four species of the genus Sphaerium are known from South America: Sphaerium aequatoriale Clessin, 1879, from Ecuador (Kuiper and Hinz, 1984); S. forbesi (Phi- lippi, 1869) recorded from Peru and Bolivia (Haas, 1949), from Bolivia (Haas, 1955), from Colombia, Peru, Bolivian Andes (Kuiper and Hinz, 1984), and Chile (Ituarte, 1995); S. lawricochae (Philippi, 1869), from Peru, Bolivia and Chile (Kuiper and Hinz, 1984), and Chile (Ituarte, 1995): S. titicacense (Pilsbry, 1924), from Peru and Bolivia (Kuiper and Hinz, 1984). According to these authors, they are concentrated in the Central Andes from Ecuador to North Chile including high- altitude Bolivia, at 2000 to 4700 m altitude; varying in size between 3 and 12.5 mm. Haas (1949) reports Sphaerium (S.) boliviense (Sturany, 1900), from the highlands of Bolivia and from a lake in Junin, Peru. He examined also material of this species collected by Sioli in the regions of the rivers Maué-Agi and Tapajos, ibutaes of the Amazon River, which constitutes the first record of Sphaerium for Brazil. According to Kui- ae and Hinz (1984), S. boliviense is a junior synonym of S. forbesi. The record of Sphaerium observationis by Mansur et al. (1991) for Mirim Lagoon in southern Brazil is a misidentification, as that species is not a Sphaerium. According to the revision by Ituarte (1995), Pisidium abionbatonss Pilsbry, 1911, only occurs in the southern Argentina, not in Brazil. Sphaerium cambaraense new species is the fifth Sphaerium species from South America and the first spe- cies of the genus described from southem Brazil, in the highlands near the Atlantic Ocean, a location geographi- cally very distant from the Andes and Amazon River. MATERIALS AND METHODS Specimens were collected with a plastic sieve with mesh size of about 0.8 mm. Specimens were sorted from the sediment with fine feather tweezers; anesthetized in small vials containing water with menthol crystals, fixed in a5% formalin solution for 24 hours, rinsed for 24 hours in tap water and preserved in 70% ethanol. Soft parts of specimens for scanning electron microscopy (SEM) were removed with tweezers, shells cleaned with a soft and fine brush and rinsed several times in distilled water. Dried shells were glued on stubs with light-silver glue (Porolon Equipment, Herts) or metallic adhesive tape (TO66 Silver tape 9 mm, Hert-Scotch), coated with gold, and observed either in a Cambridge Stereoscan 250 Mk2 or Philips scanning electron microscopes. Shell micro- structure was studied by fracturing shells at the middle of the height, parallel to commarginal ridges. Terminolo- gy for shell microstructure follows Dyduch- Falniowska (1983) and Dreher-Mansur and Meier-Brook (2000). M. C. D. Mansur et al., 2008 Page 229 Stomach nomenclature follows Purchon (1958, 1960). The shape indices, height index [I = H/L] and convexity index [Ci = W/H], were calculated according to the cri- teria followed by Ituarte (1996). Abbreviations use in the text are: MACN, Museo Argentino de Ciencias Naturales, Buenos Aires; MCN, Museu de Ciéncias Naturais, Fundagao Zoobotanica do Rio Grande do Sul, Porto Alegre; MCP, Museu de Cién- cias e Tecnologia da Pontificia Universidade Catélica, Rio Grande do Sul, Porto Alegre, Brazil. Genus Sphaerium Scopoli, 1777 Sphaerium cambaraense new species (Figures 1-20) Diagnosis: Distinguished by the relatively large and shes shell with trapezoidal tending to tric angui: ar shell outline, low and wide beaks, subce sntrally locate a without marked nepionic cap, and broad and solid hinge plate. Description: SHELL: Solid, relatively large (maximum observed L: 11.22 mm), slightly convex (Ci = 5345). Shell outline high 85+2), trapezoidal tending to triangular. Dorsal margin has pronounced curve. Anteri- or and posterior margins gradually descending and Figures 1-5. Sphaerium cambaraense new species. 1, 2. outer shell surface. 4, 5. Inner shell surface, detail of pores. Scale bars: 1, Holotype MCP Mol. Outer view of left and ri gently curved below middle of total height, without marked ange Posterior end slightly truncated, oblique (Figures 1, 2). Ventral margin long and evenly curved. Shell surface silky, glossy, with very fine irregularly distributed radial lines: weaker on beaks and irregular commarginal, sometimes coarse, striae (12 or more per 0.5 mm in the middle of the shell (Figure 3). Outer shell surface light yellowish brown; pale ae at beaks, more ventrally gr ayish brown with complete or incomplete ye sllow concentric bands running ae anterior to poste- rior margins; a large yellow band near ventral m: irgin, Beaks without marked embryonic cap. Inner shell sur- face white, grayish at muscle scars. Beaks slightly proso- gyrous, low and wide, slightly see above dorsal margin, subcentrally located | Figures 1, 2, 6, 7). Hinge plate strikingly curved solid, bioad, reaching 0.5 mm width in middie region in specimens of 1] mm length, slightly narrower at jeval of cardinal teeth. Hinge Cae. arched, pe articul: irly below cardinal teeth (Figures 6-9). Cardinal teeth strong, close to dorsal margin. Right cardinal tooth, C3, short, strongly curved, posterior end enlarged in a grooved cup (Figure 10). Left cardinal teeth: outer cardinal tooth, C4, thin, strikingly oblique, located immediately behind C2, anterior end slightly overlapping ght valves. 3. Detail of 2=4mm; 3 = 400 um; 4 = 4 um; 5 = 40 pm AUTILUS, Vol. 122, No. 4 Figures 6-11. Sphaerium cambaraense new species. Paratype (MACN-In 37063). 6. Inner view of right valve. 7. Inner view of right valve. 8. Hinge of right valve. 9. Hinge of left valve. 10. 5 cardinal teeth (C2 and C4) and ligament. Scale bars: 6, 7 = 2 mm: 2; inner cardinal tooth, C2, short high columnar, deey arched into a \ shape Figure 11). Right lateral teeth somewhat short, strong, with distal cusps | Figure 8): left lateral teeth relatively long, strong, and high (Figure 9). Ligament internal but exteriorly visible, slightly protrud- ed in larger specimens Figures LO, 11) Yetail of right cardinal tooth (C3) and ligament. 11. Detail of left S,9 =1 mm; 10, 11 = 500 um Suett Microsrructrure: Imer shell surface perforated wey 2 a1 2) (Pig by numerous pores (55/400 tim> to 13/160 tum) (Figures f, 5) representing the opening of tubuli that cross entire calcareous part of shell (shown in part in Figure 12 Openings of pores on inner surface surrounded by a funnel-shaped depression and in mouth circled by arim M. C. D. Mansur et al., 2008 Page 23 Figure 12. Sphaerium cambaraense new species. Shell microstructure from a: periostracum (top) to f: the endostracum (bottom Inserts a-g are details of: a, periostracum; b, granular layer; ec, diagonal layer forming a composite prismatic structure; d, diagonal layer. crossed structure: e, diagonal layer forming a pseudo crossed lamellar structure; f, palisade structure; g, diagonal layer, and y internal surface of the endostracum (arrow). Scale bars: 12 = 50 tm; inserts: a-d = 4 tm; e = 20 tum; f = 10 pm; g = 2 Lm Page 232 THE NAUTILUS, Vol. 122, No. 4 M.C. D. Mansur et al., 2008 Pz ge 2 9' 3h 3 (Figure 4). Shell relatively thick reaching 200 ptm in central area (Figure 12). Shell structure consists of five layers: periostracum, granular layer, diagonal layer, palisade, diagonal layer, and endostracum. Below the 2 um thick periostracum (Figure 12, a), a granular 10 pm thick layer appears (Figure 12, b); grains concentrated in some points irregularly and sparsely distributed, look- ing like cone-shaped bars arranged perpendicularly to shell surface. These change into irregularly and com- pactly arranged grains of different sizes. More internally, shell structure changes gradually into a diagonal layer (Figure 12, e, d, e), w here three different patterns may be distinguished: first (about 50 jim thick), composite prismatic structure, showing vertical feather-like pattern (Figure 12, ¢); second (about 40 tm thick), occupies the central part of shell, gradually changing into a cross-lamellar structure ( Figure 12, d): the third (approx- imately 60 [m), resembles a pseudo crossed-lamellar structure (Figure 12, e). Close to the inner shell surface there are one or two palisade layers, narrow (each about 1 um thick) (Figure 12, f). The palisade structure is followed by a diagonal layer with lamellae oriented in one direction (Figure 12, g); below, a very narrow layer, the endostracum (1.0 to 1.5 pm thick) ( (Figure 12, arrow). i Anatomy (Ficures 13-20): Inner and outer demi- branchs well-developed, outer smaller demibranch reaching half of height of inner demibranch (Figure 13). Brood sacs occupy anterodorsal part of inner demi- branchs, embryos contained in sacs showed different developmental stages, denoting sequential brooding. Up to six dev eloping embryos found in largest examined specimen (>11 mm length). Largest, tertiary, brood sac contained two embryos (>1 mm length) (Figure 13); secondary sac located under lar gest one, contained three or four small embryos. Primary sac attached lower on inner demibranch. (Specimens for this study were col- lected at the beginning of the Southern Hemisphere summer (January, 1994) and many of the specimens larger than 9 mm showed brood sacs.) Anal and branchi- al openings extended in diverging short siphons, nearly equal in size. Anal siphon wader at base and more stre- tched out (Figure 13). Two labial palps (Figure 13, 14) on each side of mouth, triangular-shaped in lateral view: opposite contacting walls with 12 small folds, tapering toward distal ends. Mantle musculature (Figure 15) with relatively short siphonal retractors; inner radial mantle muscles arranged in eight to nine bundles (Figure 15). Fan-shaped stomach (Figures 17, 18) bent to right side, laterally covered by “digestive gland. Dorsal hood relatively short and left duct well dev pile Stomach internally (Figure 17) shows very simplified structures on dorsal hood and right side such as short gastric shield under dorsal hood; minor typhlosole and rejecting tract beginning at right side after descending from dorsal hod: ne an dley ation slightly wrinkled bel tween intesti- nal groove and rejection tract; anterior fold absent. In- testine opening associated to style sac in center of floor; major typhlosole arched in front with two expansions that end respectively at left and right duct openings, not penetrating in ducts, which allocates the stomach to Type IV; left and right duct openings well developed and ramified in ieee: secondary ducts. Intestine short and simple (Figure 18), anterior part associated to style sac, broad, descending straight to floor of visceral mass; mid-intestine strikingly stretched forming single loop; hindgut straight, ascending and bending toward rectum that ends into an anal papilla. Nephridia of closed type; dorsal and outer lobes fused, impossible to distinguish in dorsal view (Figures 19, 20); external wall of outer lobe ornamented with many small, rounded extrusions; nephridia in young specimens usu- ally of open type and with separ ated lobes. Funnel rela- tively long and wide; proximal loop with smooth walls forming three ascending rings then run backward up to posterior adductor miasele. turning laterally in direction to lateral loop. Lateral loop straight, elongated, running along outer side and partially covered laterally by outer lobe. Excretory sac sub-triangular, rounded in front and relatively small. No valve at the entering of the distal loop into excretory sac. Type Locality: Lajeado da Margarida (50°15.75' W, 29°0.87' S: 870-880 m altitude) on Camisa River, Antas River Basin, considered to be the headwaters of Taquari River, the main tributary of Jacui River, South Atlantic Brazilian Basin. Type Material: Holotype MCN 38821; Paratypes: MCN 33919 (33 specimens), MCP sa (6 specimens), MACN-In 37063 (two specimens), 12 Jan. 1994. Etymology: The specific epithet refers to the City and Municipality of Cambara, close to the type locality. Distribution and Habitat: Known only from the type locality. The Municipality of Cambara is situated in northeastern Rio Grande do Sul State, Brazil, in the highest part named Planalto Riograndense. From the phy siographic point of view, this region is characterized by a basaltic shield covered by ion grass steppes and Araucaria Forest mixed with the seiithieastern limits of the Atlantic Forest. The altitude varies from 850 to 1050 m, and in the winter, temperatures fall below freezing and there is the occasional snow. The rivers that cross the region, flanked by a low gallery forest, have Figures 13-20. Sphaerium cambaraense new species. Schematic drawings of soft anatomy. 13. Gross anatomy (left mantle lobe removed). 14. Folded surfaces of inner and outer left labial palps. 15. Outer view of left mantle lobe showing the inner radial rmantle muscles. 16. Dorsal view of the stomach and digestive gland. 17. Floor of the stomach after removing the roof. 18. Inner view of the organs in the visceral mass. 19. Dorsal view of nephridia, posterior adductor muscle and posterior foot retractors. 20. Lateral view of left nephridium. Scale bars: 13, 15, 16, 18 = 2 cm; 14, 17, 19, 20 = 2 mm Page 234 THE NAUTILUS, Vol. 122, No. 4 hard bottoms formed by flattened basaltic stones, and currents are strong. The collecting sites were small ponds along the river course, where currents were low- ex, allowing the accumulation of decayed leaves and very soft, dark, and fine sand deposits, not deeper than 1 m, where specimens settle. Together with the Sphaerium samples, many specimens of one species of Pisidium sp. (MCN 33918), and one of Diplodon sp. (MCN 33920) not yet identified, were found. DISCUSSION Sphaerium cambaraense is similar to Sphaerium forbesi (Philippi, 1869) (from Bolivia, Chile, and Peru). However, S. cambaraense has a more decidedly triangular shell outline, beaks not full with not marked ne pionic shell: in addition, S. cambaraense is larger than S. forbesi. Sphaerium lauricochae Philippi, 1869), also reported from Bolivia, Chile, and Peru, differs from S$. cambaraense by its more rounded shell outline. In relation to shell thickness and microstructure, §. cambaraense is similar to the European species Sphaerium rivicola (Lamarck, 1818) and Sphaerium corneum (Linnaeus, 1758): Dyduch-Falniowska (1983) reported for these species (as well as for Musculium lacustre (Miiller, 1774) and several Pisidium species) six different layers, (1) lari (2) homogeneous-granular layer, (3) granular layer; (4) diago- nal layer (composite prismatic structure), (5) palisade structure, and (6) endostracum. Only S. corneum and S. rivicola showed a different structure for the diagonal layer, referred to as “crossed-lamellar structure” (Dy ee Falniowska, 1983). This structure was also found i S. cambaraense, however, the diagonal pattern of ie plates has a different arrangement: in S. rivicola the oblique plates show a horizontal herringbone pattern and in S. cambaraense some rows of plates are oblique not forming a horizontal pattern. These latter resemble in part the pseudo crossed-lamellar structure found in the Corbi- culidae. The periostracum layer in S. cambaraense is thin- ner than in S. corneum and the homogeneous granular layer is lacking. The simplified eondinen: of posterior and right side of stomach is quite similar to that found in the Euperinae (Dreher-Mansur and Meier-Brook, 2000). The nephridium is relatively similar compared to S. corneum (Dreher-Mansur and Meier-Brook, 2000), but the excretory sac and proximal loop are shorter and the valve at the insertion of the distal loop into excretory sac is lacking in S. cambaraensis; the multilobed surface of lateral lobe is observed for the first time in Sphaerium. Based on the similar morphology of the ne heat Korniushin (1998) propose d that South American spe- cies traditionally assigne d to Sphae rium ac tue lly be- long to Musculium. According to Park and O Foighil 2000), the usefulness of the fine anatomy of ne phridi- um in the fi umily Sphaeriidae is relative, due to its high il pl. isticity. Coole oY and O F ‘oighil (2000), tochondrial 16S r DNA gene se quences morphol NOL based T1 n ] observed that th Sphacrium/Musculium clade exhibit moderate | low levels of genetic dive rgences and the same asynchronous or sequential brooding pattern (L.e., brooding sacs contain more than one “develop- ing generations of embryos). Nevertheless, Cooley and O “Foighil (2000) recognized Musculium as a monophy- letic group. We allocate the new species in Sphaerium until more evidence is available to help with this ques- tion. More recently, a phylogenetic analysis of the Sphaeriinae (Lee and O Foighil, 2003) based on a molec- ular study of nuclear (ITS- 1) and mitochondrial (16S) gene sequences of 15 species from North and South America, Europe, Asia, and Australia, recovered a strongly supported monophyletic group of sequential brooders (Musculium and Sphaerium). However, the analyses indicate that Musculium and Sphaerium sensu lato are not natural groups, proposing a new classification system comprising five subgenera within Sphaerium: Sphae rium sensu stricto Scopoli, 1777; Musculium Link, 1807; Amesoda Rafinesque, 1520; Sphaerinova Iredale, 1943, and Herringtonium Clarke, 1973. As these subgeneric groupings, however well supported by molecular data, are not defined from the morphological point of view, it is not possible at this point to place S. cambaraense within the new scheme. ACKNOWLEDGMENTS We are grateful to the curators of MCN mollusk collection: Ingrid Heydrich and Silvia D. Hahn for providing the loan of specimens; to Prof. Dr. Cecilia Volkmer Ribeiro for the help in sampling and to Prof. Dr. Wolfgang Maier from Tiibingen University, Germany, for SEM sessions; Fabian Tricarico from the MACN SEM unit for his fine work with SEM photography. C.L. is a researcher from the Consejo Nacional de Investi- gaciones Cientificas y Técnicas (CONICET), Argentina. LITERATURE CITED Cooley, L. R. and D. O Foighil. 2000. Phylogenetic analysis of the Sphaeriidae (Mollusca: Bivalvia) based on partial mitochondrial 16S rDNA gene sequences. Invertebrate Biology 119: 299-308. Dyduch- Falniowska, A. 1983. Shell Microstructure and Sys- tematics of Sphaeriidae (Bivalvia, Eulamellibranchiata), Acta Zoologica Cracoviensia 16: 251-296. Haas, F. 1949. Land und Siisswassermollusken aus dem Amazo- nas Gebiete. Archiv fiir Molluskenkunde 78(4/6); 149-156. Haas, F. 1955. On some small collections of inland shells from ms America. Fieldiana, Zoology 34(35): 361-387. Ituarte, C. 1995. Nuevos registros de 1 Sat oe Pfe es 1821 y Spha verium Scopoli, 1777 (Bivalvia: Sphaeriidae) en C shile, Bolivia y Noroeste argentino, Neotropica 41( 0S. 106): 31-41. Ituarte, C. F. 1996, Argentine species of Pisidium Pfeiffer, 1821, and Musculium Link, 1807 (Bivalvia: Sphaeriidae) The Veliger 39 189-203. Korniushin, A. 1998a. Notes on the anatomy of some species of Sphaerium s. (Mollusca, Bivalvia) from the tropical region with revision of their taxonomic status. Vestnik 319 3): 3-12. 99 zoologii OAs SS SS ES M. C. D. Mansur et al., 2008 Page 23: Korniushin, A. 199Sb. A comparative investigation on nephri- dia in fingernail and pill clams. Malacological Review, Supplement 7: 53-63. Kuiper, J. G. J. 1962. Notes sur le systématique des pisides. Journal de Conchyliologie 102: 53-57. Kuiper, J. G. J. and W. Hinz. 1984. Zur Fauna der Klein- muscheln in den Anden (Bivalvia: Sphaeriidae). Archiv fiir Molluskenkunde 114(4/6): 137-156. Lee, T. and D. O Foighil. 2003. Phylogenetic structure of the Sphaeriinae, a global clade of freshwater bivalve molluscs, interred from nuclear (ITS-1) and mitochondrial (16S) ribosomal gene sequences. Zoological Journal of the Lin- nean Society 137: 245-260. Mansur, M. C. D. and C. Meier-Brook. 2000. Morphology of Eupera Bourguignat 1854, and Byssanodonta Orbigny 1846 with contributions to the Phylogenetic Systematics of Sphaeriidae and Corbiculidae (Bivalvia: Veneroida). Archiv fiir Molluskenkunde 128(1/2): 1-59. Mansur, M. C. D., C. Schulz, M. G. O. Silva, and N. M. R. Campos-Velho. 1991. Moluscos bivalves limnicos da Esta- cao Ecologica do Taim e areas adjacentes, Rio Grande do Sul, Brasil. Iheringia, Série Zoologia, 71: 43-58. Park, J. Kk. and D. O Foighil. 2000. Sphaeriid and corbiculid clams represent separate heterodont bivalve radiations into freshwater environments. Molecular Phylogenetics and Evolution 14: 75-88. Purchon, R. D. 1958. The stomach in the Eulamellibranchia: Stomach type IV. Proceedings of the Zoological Society of London 131; 487-525. Purchon, R. D. 1960. The stomach in the Eulamellibranchia: stomach types IV and V. Proceedings of the Zoological Society of London 135: 431-489, ; THE NAUTILUS 122(4):236-243, 2008 Page 236 Some remarks on the gross anatomy of Adelomelon ferussacii (Donovan, 1824) (Gastropoda: Volutidae) from the coast of Patagonia, Argentina Maria Eugenia Segade Guido Pastorino! Museo Argentino de Ciencias Naturales Ay. Angel Gallardo 470 3° piso lab. 57 C1405D]R Buenos Aires, ARGENTINA ABSTRACT Specimens of the volutid Adelomelon ferussacii (Donovan, 1824) were dissected. These were collected during an extraordi- nary low tide on February, 2006, at Playa La Mina, Puerto San Julian, Santa Cruz Province, Argentina oy 09’ S, 67°37’ W). The gross anatomy, along with radulae and shell ultrastructure, are described for the first time. Analysis of these features sug- gests a close relationship with the other species of the genus Adelomelon Dall, 1906, and raises doubts about the validity of Pachycymbiola Thering, 1907, a genus in which it has been included by recent authors. Additional Keywords: Southwestern Atlantic, Mollusca, Patagonia INTRODUCTION The family Volutidae includes the most conspicuous spe- cies among all endemic mollusks of the Southwestern Atlantic. This is so not only because of their large size, but also because of their interesting reproductive (Penchaszadeh, 1976, 1999: Luzzatto, 2006), Volutes live in quite diverse environments (i.e., subtidal biol Py to deep water; soft, muddy, sandy, or mixed bottoms; cold, temperate, or warm waters). Several species are extremely rare, while others are quite common. At least two species, particularly in Uruguay, are the subject of Zidona dufresnet (Donovan, 1823) and Adelomelon brasiliana (Lamarck, 1811). Other spec ies are likely to be « xploite od in the near future (Giménez and Penchaszadeh, 2002). extensive commercial fisherie 3+ The se are In addition, volutids usually aré » top predators with great ecological importance in the marine realm. Species of volutes studied thus far generally have been found to be pre dators on other mollusks and/or scavengers, } Author for correspondence gpastorino@macn,gov.ar About 30 names have been applied to Volutidae from the western Atlantic, from Venezuela to Argentina. Of these, 16 are living in Argentine waters. Most are poorly known, with dain. deived only from their original de- scription, which generally included very little biological information. In addition: ‘the taxonomic status of several names awaits clarification. Adelomelon ferussacii (Donovan, 1824), a locally com- mon volute similar to A. brasiliana, is known only from its shell. Recent findings of egg capsules suggest a compl- etely different re productive Diology ( (Pe aehasrade h and Segade, submitted). In this paper we provide new anatomical and ultra- structural data for Adelomelon ferussacii derived from a large ontogenetic series of live specimens, and compare them with other species of the same genus. MATERIALS AND METHODS Specimens used in this study were hand-collected dur- ing an extraordinary low tide on February 2006 at Playa La Mina, Puerto San Julian, Santa Cruz Province (49°09' S, 67°37’ W) (Figure The surtace water temperature was 15°C. Animals were relaxed in freez- ing sea water, preserved in ethanol 70% and dissected under a stereoscopic microscope. Radulae were pre- pared according to the method described by Solem (1972). Photographs were taken using digital cameras and images were also digitally processed. Shell ultra- structure data were procured from freshly fractured colabral sections taken from the central portion of the lip on the last whorl of several individuals. MACN: Museo Argentino de Ciencias Naturales. SYSTEMATICS Family Volutidae Rafinesque, 1S15 Subfamily Zidoninae H. Adams and A. Adams, 1853 Genus Adelomelon Dall, 1906 Adelomelon ferussacii (Donovan, 1824) enn nnn TT EEE M. E. Segade and G. Pastorino, 2008 Page 237 T T “ATS yh is Buenos Aires = : \ Golfo San Matias > Figure I. Santa Cruz, Argentina. Voluta ferussacii Donovan, 1824, 2, pl. 67; Sowerby, 1546 1: 203, pl. 46, fig. 7; Reeve, 1549 6, pl. 10, fig. 23, spec. 39; Dall, 1907: 362. Voluta rudis Gray in Griffith and Pidgeon, 1834: 601, pl. 30, fig. 1. Voluta oviformis Lahille, 1895: 20, pl. 1 figs. 53-56, pl. 7, figs. 121-138, pl. 1 Voluta oviformis typica Lahille, 1895: 20. Voluta oviformis longiuscula Lahille, 1895: 20. Voluta oviformis fratercula Lahille, 1895: 20. Voluta (Cymbiola) ferussacii Donovan.—Strebel, 1906: 100, pl. 9. figs. 46, 46a, 4549. figs. 1-2, pl. 2, 0. figs. 4-9. Map showing the study area of Playa La Mina, Adelomelon (2?) ferussacit Donovan.—Clench — and Turner, 1964: 157, PI. 98, figs. 1-3. Adelomedon (sic) ferussacii (Donovan).—Castellanos, 1970: 110, pl. §, fig. 7 Adelomedon (sic) ferrusacii |sic]|—Castellanos, 1970b: 1, figs. 6, 9. Adelomelon (Pachycymbiola) ferussacii (Donovan, 1824). —Weaver and DuPont, 1970: 108, pl. 45C, 45D. Adelomelon ferussacii (Donovan, 1524).—Castellanos and Landoni, 1992: 12, Pl. 1, fig. 8. Pachycymbiola ferussacii (Donovan, 1824) —Poppe and Goto, 1992: 116, pl. 38, figs. 3-5. Description: Shell medium in size, up to 122 mm, solid, fusiform; color grayish-brown. Aperture semi- circular, dark-brown oe Protoconch of 144 smooth whorls; teleoconch of up to 4 slightly convex whorls; spire low, sometimes somewhat upturned; spire angle of 80°, suture well defined. Columella curved, orange, with three to six folds set obl raed to siphonal Geciale, regu- larly distributed except for the anteriormost one, w nels is separate from the others. Columellar callus usually weak, but sometimes thick. Siphonal canal fairly broad and shallow. Growth lines span the surface, sometimes producing irregular costae. Shell ultrastructure showing three layers: a crossed! lamellar aragonite layer in w hich the cry stal planes are arranged perpendicul: uly to the growing edge of the shell, cand also perpe sndicular to the adel e layer (25% shell thickness); a middle layer (50% shell thickness) of colabrally aligned crossed- lamellar aragonite, and an outer layer (25% shell thick- ness) of amorphous calcite (Figure 1S). The innermost layer varies in thickness according to the sector of the lip: along the most curved sector it appears thicker, while it is reduced or absent towards the ends of the lip. a arrangement is quite similar to that found in shells of A. brasiliana (Figure 19) and A. beckii Broderio: 1836). Embryonic shells very thin (at hatching stage), whitish in the first whorls and dark brown in the ‘|: ist protoconch whorl. Surface covered by 8-10 regularly spaced spiral threads in the last whorl, no ple iits visible. Calcarella reduced, weakly pronounced. Foot, head, and siphon are finely mottled purple in color. The contact surface of the foot is whitish. Foot and shell length are similar. Operculum absent. Head broad and flattened, with two short tubular tentacles that separate the lateral lappets from the central one. Eyes are very small and located near the base of the tentacles. The penis emerges directly behind the right cephalic lappet (Figure 21). The siphon, also well pig- mented, is muscular with paired and symmetrical si aa nal appendages emerging from the base of the eee and spanning half its length. The bipectinate osphradium has aproximately 100 equal leaflets. The ctenidium is 14 times as long as thi osphradium. The hypobranchial gland is thin. The ie boscis shows the same color pattern as the foot, head and siphon. Mouth opening is triangular THE NAUTILUS, Vol. 122, No. 4 M. E. Segade and G. Pastorino, 2008 Page 239 Figures 18-19. Radular ribbon narrow, up to 17.4 mm in length (n = 13; x = 1L.7; DS = 1.97), with 49-74 rows (proportional to shell length) with one tricuspid central tooth per row (Figure 22). The radulae increase the number of teeth with age (Fig- ure 32). Central tooth thin and long, anterior profile con- cave with a ventral-posterior thickening (Figures 23 and 24). Lateral cusps of the rachidian tooth are similar in size to the central one. Each cusp with a dorsal shallow indenta- tion or groove present, where the corresponding previous cusp imbricates (Figures 24-25). Embryonic radulae showing about 15 rows of teeth where lateral cusps are shorter than central cusps (Figure 26; Table 1). Salivary glands (racemose glands) large and irregu- larly shaped. Accessory salivary glands (tubular glands) very long and extremely convoluted. distally expanded, ov erlying dorsal surface of saliv ary glands. The tubular glands can easily be separ ated from the racemose glands. Ducts of the accessory salivary glands descend laterally to oesophagus and are sub kernal with respect to the mouth. They never fused and end separately. Ducts of salivary glands become embedded in the Shell ultrastructure. 18. Adelomelon ferussacii (Donovan, 1824). 19. Adelomelon brasiliana (Lamarck, 1811). Scale bars = 500 tm. oesophagus anterior to small valve of Leiblein. The an- terior esophagus runs behind the proboscis, and passes through the nerve-ring where an externally inconspicu- ous valve of Leiblein is located. The gland of Leiblein is very long and extremely conv oluted (Figure 20). The posterior oesophagus continues to the stomach, Material Examined: (D = specimen was dead when sampled: A = alive) Punta Cavendish, Puerto Deseado, Santa Cruz province, 2 D, collected in 5 m depth (MACN-In 31354); Patagonian coast, 1 D (MACN-In 11385); Playa Cabo San "Pablo, Tierra del Fuego, 4 D (MACN-In 12532); Punta Sinaia, Tierra del Fuego, Expedicion Facultad, 5 D (MACN-In 12531); Ushi, Tierra del Fuego, 1 D (MACN-In 9441); Punta Marta, Rio Grande, Tierra del Fuego, 9 D (MACN-In 35113): Bahia San Sebastian, Tierra del Fuego, 1 D (MACN-In 35393): San Sebastian south, Tierra’ del Fuego, 4 D, Exp. Facultad de Ciencias (MACN-In 12530); Play a del Rio Grande, Santa Cruz province, 1 D, (MACN-In 6647); 52°20! S, 6S°18! W, Station 28, Cabo Virgenes. Table 1. Dimensions (mm) of some adults of Adelomelon ferussacii from MACN collection, Specimen 1 2 3 4 5 6 ¢ 8 9 10 11 12 13 Sex 3 3 + 3 = 3 3 - Y 2 + 2 : Shell length (mm) (SL) 76.5 75 58 91 78 85 75 72 76 74.5 76 114 75 Aperture length (AL) 60 57 46 69 60 64 59 56 58 58 56 S9 56.5 AL/SL 0.78 0.76 0.79 0.76 0.77 0.75 0.79 0.78 0.76 0.78 0.74 0.78 0.75 Spire angle (°) 80 78 77 83 82 S4 59 SO 83 78 Sl $5 82 Radula length (RL) 10.5 11.3 9.5 12.2 12.] 13.1 11.4 10.4 9.6 11S 12.1 17.4 11.7 Number of radular teeth 56 50 49 53 61 58 56 52 50 60 58 74 57 Figures 2-17. Shells of A. ferussacii and A. brasiliana, 2-13. Adelomelon ferussacii (Donovan, 1824). 2-4. MACN-In 37015 Puerto San Julian, Santa Cruz. 5-6. Holotype of Voluta rudis Griffith and Pidgeon, 1834, NHM 19920177. 7-10. Embryo shells (at hatching stage) MACN-In 3 Buenos Aires. Scale bars = 10 mm. 7014. Puerto San Julian, Santa Cruz. 11-13. Juvenile shells MACN-In 37012, 3 kin North of Puerto San Sebastian. Tierra del Fuego. 14-17. Adelomelon brasiliana (Lamarck, 1811), embryo shells MACN-In 37015, off Mar del Plata, Page 240 THE NAUTILUS, Vol. 122, No. 4 Santa Cruz province, A.R.A. Bahia Blanca, 1 D, in 11 m (MACN-In 24080): Bahia Laura, Puerto Deseado, Santa Cruz province, 1 D (MACN-In 9199-16); Bahia San Sebastian, Tierra del Fuego, 1 D (MACN-In 21154); Estancia “Viamonte”, Rio Grande, Tierra del Fuego, 2 D (MACN-In 27219): Isla Quiroga, Puerto Deseado, Santa Cruz province, 1 D (MACN-In 26199); Punta Arenas, Chile, 1 D (MACN-In 9040-27); Rio Grande, Tierra del Fuego, 24 D (MACN-In 12529); Playa La Mina, Puerto San Julian, Santa Cruz Province (67° 37’ W, 49° 09’ S), 10 A in low tide (MACN-In 37487). Distribution: Adelomelon ferussacii is a typical com- ponent of the Magellanic province, ranging from the province of Santa Cruz to Southern Chile. More north- ern citations of the species (e.g., Clench and Turmer, 1964; Castellanos and Landoni, ve are here referred to A. brasiliana. No specimens of A. ferussacii from re- vised collections (MLP and MACN) were found outside of this range. Distribution According to Previous Records in the Literature: Puerto Gallegos, Punta Arenas, (Stre- bel, 1906); Santa Cruz coast, (Lahille, 1895); Southern Patagonia and Magellanic region (Carcelles and William- son, 1951); Golfo San Matias to Straits of Magellan (Clench ane Turner, 1964), however they only examined specimens from the acre of Puerto Deseado, Cabo Buen Tiempo, Rio Gallegos, Bahia de la Posesion and San Gregorio in Chile; Golfo San Matias to Magellan Straits (Weaver and DuPont, 1970); Santa Cruz (Castel- lanos, 1970b); Southern coast of Buenos Aires province to Magellan Straits (Castellanos and Landoni, 1992); Southeastern coast of Argentina, south to the Magellan Straits (Poppe and Goto, 1992). DISCUSSION Clench and Turner (1964) suggested the inclusion of Voluta ferussacii in the genus Adelomelon Dall, 1906, because of shell similarity with A. brasiliana. However, at that time they examined no complete specimens to confirm such generic placement. Weaver and Du Pont (1970) mentioned that no live specimens were collected. Later, Castellanos (1970) illustrated the radula confirm- ing the generic placement suggested by Clench and Turner (1964). However, data on the gross anatomy had not previously been reported. The last comprehe nsive taxonomic revision of the family Volutidae from the southwestern Atlantic was prepared by Clench and Turner (1964, 1970). They de- scribed the new subfamily Odontocymbiolinae and the new genus Odontocymbiola, and finally resolved the confusion of previous authors (e.¢., Pilsbry and Olsson, 1954) between Adelomelon ancilla (Lightfoot, 1786) and Odontocymbiola magellanica (Gmelin, 1791). These authors also described two new species: O pescalia anc A. riosi (Clench and Turner, 1964). The latter was in- cluded in the new beenus Weaveria. After their work, 21 S Figures 20-21. Adelomelon ferussacii (Donovan, 1824). 20. Anterior alimentary system. 21. Dorsal view of head, siphon, and penis of a male specimen. ae, anterior esophagus; asg, accesory salivary gland; el, cephalic lappet; dasg, duct acce- sory salivary gl: ind; e, eye: gl, gland of Leiblein; nr, nerve ring; Ps penis; pd, penial duct: pe, posterior esophagus; rs, radular sac; s, siphon; sg, salivary gland; t, tentacle. several additional new species were described, in partic- ular from Brazil (e.g., Leal and Bouchet, 1989; Leal and Aios, 1990). Most of the literature dealing with southwestern At- lantic volutids is primarily taxonomic. Anatomical data are scarce and usually drawn from one or relatively few spe clmens, sometimes incomple te. Exce ptions are the papers by Novelli and Novelli (1982) and Aycgaguer (2002), pi articularly the latter, in sehich the authors de- scribe ( in some detail the an: itomy of Zidona dufresinei. Clench and Turner (1964) and Aycaguer (2002) men- tioned that a beckii, A. ancilla, A. brasiliana, and Zidona dufresnei all have characteristic racemose salivary glands loosely intertwined in the tubular M. E. Segade and G. Pastorino, 2008 Page 24] Figures 22-31. Radulae of Adelomelon species. 22-26. Adelomelon ferussacii (Donovan, 1824). 22. Frontal view. Scale bai 250 um. 23. Lateral view. Scale bar = 200 um. 24-25. Rachidian teeth. Scale bar = 200 um. 26. Radula of an embryo. Scale bai 50 um. 27-31. Adelomelon brasiliana (Lamarck, 1811). 27. Radula of an embryo. Scale bar = 50 um. 28-29. Rachidian teeth. Scale bar = 200 um. 30. Lateral view. Scale bar = 200 um. 31. Frontal view. Scale bar = 250 tm age IAI Page 242 THE NAUTILUS, Vol. 122, No. 4 Number of teeth 0 20 40 60 80 100 120 Shell length (mm) Figure 32. Relationship between radular and shell length (R = 0.9178). accessory glands. The radula in these species is also rachiglossate with a unique central tricuspid tooth. These characters, together with several shell similarities placed A. ferussacii in the Zidoninae subfamily. Radulae of the species A. ferussacii and A. brasiliana are quite similar (Figures 22 and 31). However, A. brasiliana has a wider and more convex base of the rachidian teeth than A. ferussacii (Figures 24 and 28). Lateral cusps of the rachidian teeth are shorter and wider in A. brasiliana than in A. ferussacii. These differences are probably specific characters and are also present in the embryos of both species (Figures 26 and 27). The name Pachycymbiola was proposed by ther- ing (1907: 209) as a subgenus of Ade lomelon for A beatiand. which is aeuially the type species. Pilsbry ee Olsson (1954) and later Searabino et al. (2004) ) pro- moted Pachycymbiola to generic rank, and mentioned as main characters an ovate shell with a short spire, a free oval egg capsule, and a protoconch without calcarella. Del Rio and Martinez (2006) also treated Pachycym- biola at the generic rank following the latter authors. They described five Tertiary species, three new, under- this genus, pointing out as main differences from Adelo- melon the ovate shape of the shell, the short spire and the protoconch without calcarella, in agreement with Searabino et al. (2004). Adelomelon brasiliana has all these features and a large free ovoid ovicapsule, with 9 to 33 embryos per capsule (Penchaszadeh and de Mahieu, 1976; Luzzatto, 2006). On the other hand, A. beckii and A. ancilla, which belong in the subgenus Adelomelon, have smaller egg capsules, always att: ache d to hard substrates (Penchaszadeh et al., 1999). Adelome- lon ferussacti presents egg capsules similar to those of \. beckii, which are globose hemispherical and flexible, with a white opaque color and a leathery texture, attached to a hard substrate such as stones or rocky substr. te Penchaszadeh and Segade, in preparation). The inclusion of A. ferussacii in the subgenus Pachy- suggested based on some external Srnbios vas first similarity to A. brasiliana. However the ege capsules do not match those of A. ferussacii. In fact they look more similar to those of other species of Adelome lon. Also, the gross anatomy revised here does not show conclusive fennie: to include A. ferussacii in a different group as suggested previously. The only character that could siacid as a difference are those of shell morphology. Pilsby and Olsson (1954), Scarabino et al. (2004) and Del Rio and Martinez (2006) concede that shell shape place A. ferussacii close to Pachycymbiola. They do not mention that a calearella is reduced, present in the young of A, fe russacti (Figures 7-13), but absent in those of A. brasiliana. At a closer look even the shell shape is substantially different, since A. brasiliana has a more flattened spire and pronounced shoulder tubercles more similar to A. becki than A. ferussacii. Further de- tailed anatomical studies will confirm the relationships of these Southwestern Atlantic endemic volutids. ACKNOWLEDGMENTS We thank M. Griffin (UNLPam) and F. Searabino for the suggestions that considerably improved the manu- script. Bronwyn Gillanders helped to improve the English. Two anonymous reviewers made useful sugges- tions on an early draft of the manuscript. This contribution was supported by Project PICT No. 03-14419 from the National Agency for Scientific and Technical Promotion, Argentina. We acknowledge funding by the Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET) of Argentina, to which GP belongs. LITERATURE CITED Aycaguer, C. sts Anatomia de Volutas del Atlaintico sudocci- dental, I: Anatomia general y del sistema reproductor de Zidona ‘een Donovan, 1823) (Neogastropoda: Volu- tidae), Comunicaciones de la Sociedad Malacologica del Uruguay 8(76-77): 159-180. Carcelles, A. and S. Williamson. 1951. Catalogo de los moluscos sce de la provincia magallanica. Revista del Instituto Nacional de Investigacién de las Ciencias Naturales 2, Ciencias Zoologicas: 225-383. Castellanos, Z. J. A. de. 1970. Reubicacion de algunas especies de Volutidae del Mar Argentino. Neotropica 16: 1-4. Castellanos, Z. J. A. de and N. Landoni. 1992. Catalogo descriptivo de la malacofauna marina magallanica LO. Neo- gastropoda, Volutidae, Volutomitridae, Cancellariidae, Olividae y Marginellidae. Comision de Investigaciones Cientificas de la Provincia de Buenos Aires, La Plata, 37 pp. Clench, W. J. and R. D. Turner, 1964. The subfamilies Voluti- nae, Zidoninae, Odontocymbiolinae on Calliotectinae in the Western caer Jobnsonia, 4(43): 129-180. Clench, W. J. and R. D. Turner. 1970. a family Volutidae in the Mah m eee Johnsonia 4: 369-372. Dall, W. H. 1907. A review of the American Volutidae. Smith- tie Miscellaneous Collections 48(3): 341-3735 del Rio, C. J. and S. A. Martinez. 2006 The family Volutidae (Mollusca-Gastropoda) in the Tertiary of Patagonia (Argentina). Journal of Paleontology SO: 919-945 Donovan, E. 1824. The naturalist’s repository or miscel- lany of exotic natural history, exhibiting specimens of M. E. Segade and G. Pastorino, 2008 Page 243 foreign birds, insects, shells. Sheet EE, FF, pp. 214-216, Plate 67. Giménez, J. and P. E. Penchaszadeh. 2002. Reproductive cycle of Zidona dufresnei (Caenogastropoda: Volutidae) from the southwestern Atlantic Ocean. Marine Biology 140: 755-761. Griffith, E. and E. Pidgeon. 1834. The animal kingdon arranged in conformity with its organization by the baron Cuvier. The Mollusca and Radiata. London, Whittaker & Co., 601 pp: Ihering, H. von. 1907. Les Mollusques fossiles du Tertiaire et du Cretace superieur de large sntine. Anales del Museo Nacional de Buenos Aires (3)7: 1-611. Lahille, F. 1895. Contribucion al eens de las volutas argen- tinas I. Morfologia externa. Revista del Museo de La Plata 6: 293-333. Leal, J. H. and P. Bouchet. 1989. New deep-water Volutidae from off southeastern Brazil (Mollusca: Gastropoda). The Nautilus 103: 1-12. Leal, J. H. and E. de C. Rios. 1990. Nanomelon vossi, a new deep-water Zidoninae from off southern Brazil (Gastro- poda: Volutidae). The Veliger 33: 317-320. Luzzatto. D. C. 2006. The biology and ecology of the giant free egg capsules of Adelomelon brasiliana Lamarck, 1811 (Gastropoda: Volutidae). Malacologia 49: 107-119. Novelli, R. and A. U. G. Novelli. 1982. Aleumas consideragdes sobre a subfamilia Zidoninae e notas sobre a anatomia de Adelomelon brasiliana (Lamarck, 1811), Mollusca, Gastropoda, Volutidae. Atlantica 5: 23-34. Penchaszadeh, P. and G. C. De Mahieu. 1976. Reproduccion de gaster6podos Prosobranquios del oe surocciden- tal. Volutidae. Physis A, 35(91): 145-15 Penchaszadeh, P. E., P. Miloslavich, M. Lasta, and P. M. S. Costa. 1999. Egg capsules in the genus Adelomelon (Caenogastropoda: Volutidae) from the Atlantic coast of South America. The Nautilus, 113(2): 56-63. Pilsbry, H. A. and A. A. Olsson. 1954. Systems of the Volutidae. Bulletins of American Paleontology, 35(152): 5-36. Poppe, G. T. and Y. Goto. 1992. Volutes. Mostra Mondiale Malacologia Cupra Marittima (AP-Italy). L’Informatore Piceno Ed., Ancona, 348 pp: Reeve, L. 1849. Conchologia Iconica: or, illustrations of the shells of moluscous animals. London. Scarabino, F., S. Martinez, C. J. del Rio, A. E. Oleinik, H. H. Camacho, and W. J. Zinsmeister. 2004. Two new species of Adelomelon Dall, 1906 (Gastropoda: Volutidae) from the Tertiary of Patagonia (Argentina). Journal of Paleon- tology 78: 914-919. Sowerby, G. B. II. 1846. Descriptions of Tertiary fossil shells from South America. In: Darwin, C. Geological Observa- tions on South America. Smith Elder Co., London, 249-267. Solem, A. 1972. Malacological application of Scanning Elec- tron Microscopy, H. Radular structure and functioning. The Veliger, 14: 327-336, Strebel, H. 1906. Beitriége zur Kenntnis der Molluskenfauna der Magalhaen-Provinz. 4. Zoologischen Jahrbuchern. Abteilung fur Systematik, Geographie und Biologie der Tiere, oA 91-174. Weaver, C. S. and J. E. Dupont. 1970. Living volutes. A mono- graph of the Recent Volutidae of the world. Delaware museum of Natural History. Monograph Series No. 1, Delaware, xv + 375 pp. THE NAUTILUS 122(4):244-251, 2008 Page 244 A new species and a new record of Muricidae (Gastropoda) from Brazil: genera Pterynotus and Leptotrophon Alexandre Dias Pimenta Diogo Ribeiro do Couto Paulo Marcio Santos Costa Departamento de Invertebrados Museu Nacional, Universidade Federal do Rio de Janeiro 20940-040 Rio de Janeiro, BRAZIL adpimenta@yahoo.com.br ABSTRACT Leptotrophon atlanticus, a new species of Leptotrophon Houart, 1995, a genus of Trophoninae previously known only from the Indo-Pacific Ocean, is described from the northeast- ern coast of Brazil, from depths of 240-260 m. Leptotrophon atlanticus is very similar to Leptotrophon kastoroae Houart, 2001, but has shorter spines, which are brown colored. The shell microsculpture of L. atlanticus was studied using scan- ning electron microscopy, which revealed a protoconch slightly granulated on the last quarter of the whorl and a teleoconch with growth striae crossed by regularly spaced zigzag spiral lines and axial microstriae. The 1 cadule of L. ailantio us fits the pattern described for the genus, but the lateral/marginal cusps of the rachidian teeth are not as independent of each other, being somewhat fused into a common base. This study reports the first record from the South Atlantic Ocean of the Murici- nae Pterynotus havanensis Vokes, 1970, previously known only from the Caribbean region. Additional keywords: Muricinae, Trophoninae, South Atlantic, biodiversity, tasonomy INTRODUCTION The family Muricidae is represented in Brazil by 49 species from six subfamilies, of which members of the subfamily Muricinae are most numerous, with 24 spe- cies in nine genera (Rios, 1994). Other genera of Mur- icinae recorded from the western Atlantic have never been recorded from Brazil until now. This includes the genus Prerynotus, which is found in deeper-water habi- tats (Vokes, 1970), with Florida and the eight species reported from Caribbean (Rosenberg, 2005), in addi- tion to mention records from other regions of the world, e.g., Ponder (1972) from Australia, Bouchet and Warén (1985) from the northeast Atlantic, and Houart (2001) from New Caledonia. Phe family Trophoninae includes 18 genera Vaught, 1989). It is represented in the western Atlantic by five genera, of which only Trophon Montfort, 1810, occurs in Brazil (Rios, 1994). More recently, Houart (1995) described the genus Leptotrophon, whose 26 spe- cies are restricted to the Indo-Pacific region, in New Caledonia (Houart, 1995, 2001) and eastern Indonesia (Houart, 1997). This report presents the first record of the genus Pterynotus Swainson, 1833 from Brazil, based on Ptery- notus havanensis Vokes, 1970, as well as the first record of the genus Leptotrophon from the Atlantic Ocean, based on a new species described from northeast Brazil. MATERIALS AND METHODS This report is based on a sample collected on the Cano- pus Bank, State of Ceara, in 2005, from a biogenic sub- stratum, in 240-260 m depth. Identification of the species was based on original and subsequent descrip- tions. In cases where fragme nts of soft parts were avail- able, an attempt was made to record some anatomical characteristics using camera lucida, especially of the head-foot and the palli ial cavity. Radulae were prepared for SEM according to the methodology described by Bandel (1984). SYSTEMATICS Family Muricidae Rafine asque, LS15 Subfamily Muricinae Ratine sque, IS15 Genus Pter ynotus Swainson, 1$35 Type Species: = Murex pinnatus Swainson, 1$22 by subsequent designation. Pterynotus havanensis Vokes, 1970 (Figures 1-12) Pterynotus (Pterynotus) havanensis Vokes, 1970 (new name for Murex tristichus Dall, 1889, non Murex tristichus Beyrich, 1854.) \. D. Pimenta et al., 2008 a a Figures 1-8. Pterynotu havanensis Vokes, 1970: 1—4, 7-8. MNR]J 11057 lensth 24.4 mm, width = 16.5 mm. 5-6. MNR] 11003. 1. Shell in apertural view. 2. Shell in abapertural viev oy Shell in apical vie w. 4. Shell in lateral view. 5. Protoconch. 6. Detail f tel I | ». 7. Radulae in dorsal view. 8. Detail of rachidian teeth. Scale bars: 5-6 LOO pum; 7-5 10.0 tum Murex (Pteronotus ) tristichus Dall, 1889: 202 pl 15, fig. 2). Pterynotus phan us auct Abbott, 1974: 175, fig. 1856 Pteropurpura tristic ha.—Dall, 1927: 58 non Dall, 1889 Mure Pterynotus) tristichu Clench and Farfante Pterynotus phaneus Dall—Radwin and D’Attilio, 1976 1945: 36, pl. 20, igs. 14 100 (in part Page 246 THE NAUTILUS, Vol. 122, No. 4 Figures 9-12. Pterynotus havanensis Vokes, 1970 (MNRKJ 11057). 9. Head-foot in apertural view, female, 10. Operculum in inner view. 11, Operculum in outer view. 12. Palial cavity, female. Scale bars = 1.0 mm. Abbreviations: an, anus; em, columellar muscle; en, ctenidium; ct, cephalic tentacle; ey, eye; hg, hypobranchial gland; mb, mantle border; op, operculum; os, osphradium; ov, oviduct; si, siphon. A. D. Pimenta et al., 2008 Page 247 Pterynotus havanensis Vokes—Fair, 1976: 47, pl. 18, fig. 164. errs (Pterynotus ) phaneus auct.—Harasewych and Jensen, 1979: 4, fig. 3 (in part, non Dall, 1889). Description: Shell of moderate size (up to 25 mm in length), fusiform, thin; color white. Protoconch paucispiral, with 1.5 whorls, glassy, slightly bulbous, ending in a thin varix. Teleoconch with up to five whorls, somewhat convex in outline, with three thin, flaring, backward-curved, wing-like varices per whorl, with 5-6 digitations along margin of last whorl varices. Axial sculpture consisting of fine growth striae, more visible in ventral view of varices, where they cross with spiral threads, giving it a foliaceous appearance; no intervarical nodes present. Spiral sculpture obsolete, barely discernible spiral threads, formed by microscopic shallow furrows, somewhat undulated and irregularly spaced; abapertural view of varices with five-six spiny digitations per varix on the last whorl and two digitations on spire varices; spine on shoulders of whorls largest. Aperture oval, peristome slightly projected, inner lip reflected, attached posteriorly, smooth, with small undulations corresponding to varical digitations; outer lip smooth. Siphon canal rather long, sharply bent aba- perturally and to the right side; narrowly open; with previous, old imbricate canal termination, recurved to the left side. RapuLa (FEMALE): Rachiglossate type; radula ribbon long and narrow; rachidian teeth with five pointed cusps, central cusp the largest, the second largest at the margins, each area between central and mar anal cusps with an acute tiny cusp; marginal edge rectangular and somewhat pointed, base concave: lneral teeth sickle- shaped, broader at the base. Heap-Foor (FEMALE): Head poorly differentiated, eyes located laterally and in middle of somewhat long cephal- ic tentacles. Foot large. OPERCULUM horny, ov ate, cover- ing entire shell aperture; external surface with terminal nucleus and concentric growth lines; imner surface attachment area with single horseshoe-shaped scar, not positioned centrally, covering less than 50% of total area of operculum, with one adventitious layer. PaLLIAL Cavity (FEMALE): Mantle border smooth; siphon long and narrow, with smooth border, muscle attach- ment reaching as far as gill. Osphradium bipectinate, unequal, right side about 1.5 times as wide as left, broad and long (about half total ctenidium length). Ctenidium monopectinate, long and narrow. Hypobranchial gland a thin. poorly dev eloped glandular mass covering mantle between anterior end on gill and oviduct. Oviduct occu- pying about half of pallial cavity length, broad. Type Locality: Blake Station 51, off Havana, Cuba, between 445 m and $23.5 m. Material Examined: MNRJ 11003 (one individual and seven empty shells); MNRJ 11067 (one individual); MNR] 11057 (one individual); all from Canopus Bank, 96 miles off Ceara State, 240-260 m depth, from bio- genic substratum. Geographic Distribution: Off Georgia and Fernan- dina (Dall, 1927), Key West (Fair, 1976) Florida, USA; Havana, Cuba (type locality); Pleistocene Moin Forma- tion, Costa Rica (Vokes, 1992): Golfo of Uraba, Colom- bia (Vokes, 1992); Ceara State, Brazil (this study). Discussion: Pterynotus havanenis Vokes, 1970, was originally described as Murex (Pteronotus) tristichus Dall. 1889. The taxon was later included in a long list of synonyms of Pterynotus phaneus (Dall, 1889) by Hara- sewych and Jensen (1979), based on a wide variation found in several shell characteristics such as the axial sculpture, intervarical nodes and outer lip. Vokes (1992) revalidated P. havanensis, based on the more numerous varices on the early teleoconch whorls and the smoother shell surface. The only available published illustrations of P. hava- nensis are those of the holotype (Dall, 1889: pl. 15, fig. 2; Clench and Farfante, 1945: pl. 20, figs. 1-4; Vokes, 1970: pl. 3, figs. la,b; Abbott, 1974: fig. 1856; Fair, 1976: pl. 13, fig. 164. Hasisenyel and Jensen, 1979: fig. 3; Vokes, 1992: pl 2, fig. 3), in addition to a Pleistocene fossil specimen from Costa Rica (Vokes, 1992: pl. 2, fig. 6). This study provides the first illustrations of Recent specimens (Figures 1-4), except for that of the holotype itself. The Brazilian specimens herein studied, all from Canopus Bank, about 96 miles off the coast of the State of Ceara (240-260 m depth), are very similar to the holotype. illustrations, ve little sculpturing between the varices (Figures 1-2, 4), without the intervarical nodes described for P. phaneus, in addition to axial and spiral g growth lines, the digitations on the margins of the varices and no denteulae outer lip (Figure 1). These characteristics, along with the shape sa sculpture of the varices, clearly distinguishes P. havanensis from P. phaneus. Vokes (1992) stated that both P. phaneus and P. hava- nensis have denticulations on the inner side of the outer lip. However, this detail is not stated in the original or subsequent descriptions of this species, as well as in the holotype illustrations, which show a smooth outer lip, as well as in the specimens from Brazil. Such denticles reported by Vokes (1992) probably correspond to the undulations by the varical digitations. In addition, Vokes (1992) considered that P. hasoneneis bears several vari- ces on the early teleoconch whorls, making reference to the figure in Harasewych and Jensen (1979). Brazilian specimens bear the usual three varices on the first tele- oconch whorls. The radula herein illustrated ( minor differences from the radula of P. phaneus: illu- strated in Harasewych and Jensen (1979: 15, fig. 17). In P. havanensis, the central and lateral cusps are longer in relation to the total length of the tooth; also, the two inner cusps are narrower and shorter. Figures 7-8) has some THE NAUTILUS, Vol. 122, No. 4 Subfamily Trophoninae Cossmann, 1903 Genus Leptotrophon Houart, 1995 Type Species: Leptotrophon caroae Houart, 1995, by original designation. Recent, New Caledonia. Leptotrophon atlanticus new species (Figures 13-26) Diagnosis: Shell densely spiny; color cream white with brown spines; paucispiral protoconch with slight granu- lated microsculpture on last quarter of last whorl. Figures 13-20. Leptotrophon atlanticus new species. Holotype, MNRJ 11004, length 14. Abapertural vi 20. Detail of lateral teeth. Scale bars Description: Shell up to 8.5 mm in length (holotype), biconic, and densely spiny; spire high. Color creamy white, with light brown spines in live specimens. Proto- conch rounded, paucispiral, with 1.50 to 1.75 whorls, with slightly granulated microsculpture, forming faded spiral cords on last quarter of last whorl; terminal varix low. Teleoconch with up to 4.25 slightly shouldered whorls. Suture impressed. Axial sculpture consisting of numerous weak, orthocline lamellate growth striae and spiny varices, regularly spaced, forming spines at inter- ceptions of spiral cords. Spiral sculpture on spire of two 8.8 mm, width {6 mm). 13. \pertural 15. Apical view. 16. Operculum in inner view. 17. ( Yperculum in outer view. 18. Radulae in dorsal view 16-17 = 500 tm ; 1S—20 = 10 jum \. D. Pimenta et al., 2008 Figures 21-26. Leptotrophon atlanticus new species. Paratype, MNRJ 11009, length = §.2 mm, width = 4.4 mm. 21. Abapertural view. 22. Detail of sculpture on teleoconch whorl. 23. Detail of microsculpture on teleoconch whorl. 24. Protoconch. 25-26. Detail ot protoconch sculpture Seale bars = 100 tm strong cords, the adapical one located at 1/3 of whorl height below the suture; interceptions between axial var- ices and spiral cords form two primary spines regularly arranged in spiral crowns, each crown with 9-10 spines on last whorl of holotype: spines tall, channeled, com- monly adapically and backward-curved, slightly larger at shoulder: Spaces between adjacent spines have a squa- mous appearance, due to crossing of spiral cords with axial growth striae: last whorl with four additional spiral spiny crowns, regularly arranged at the base, along with corresponding spiral cords; secondary squamous small spines appear irregularly arranged, interspaced with pri- mary spines; holoty pew ith a secc mdary spiral cord appear- ing between the two pmmary ones closest to the suture on the last half of the last whorl! Microsculpture formed by growth striae crossed by regularly spaced ZIPZA spiral lines and axial microstriae \perture slightly oval, rounded adapically with about 1/3 of total shell height. Columellar ip flaring, smooth and adapically adherent. Anal notch indistinct. Outer lip smooth, fragile, primary spiral cords and growth striae visible within. Siphon canal long, nar- ‘ow, bent backwards, narrowly open and smooth, with seven imbricate old canal terminations Raputa (FEMALE): of rachiglossate type; radula ribbon ong and narrow; rachidian tooth trapezoidal with five pointed cusps; the two lateral cusps adjacent to central Cc sp somewhat fused into a bifurcated cusp, outermost cusps slightly larger; marginal edges rectangular well pointed, area between outer cusps and marginal edges with two very small folds; base somewhat sinuous; lateral teeth sickle-shaped broader at base OrercuLuM horny, elliptical, covering entire shell ape ture: external surface with terminal nucleus and concen tric growth lines: inner surface attachment area with single horseshoe-shaped scar, not positioned centrally Page 250 THE NAUTILUS, Vol. 122, No. 4 covering about 50% of total area of operculum, with one adventitious layer. Type Material: Holotype: MNRJ 11004 (8.5 mm); three paratypes: MNRJ 11009, all from type locality. Type Locality: Canopus Bank, 96 miles off Ceara State, 240-260 m depth, from biogenic substratum. Distribution: Known from type locality only. Discussion: The allocation of Leptotrophon in Trophoninae follows Houart (1995), who, when describ- ing Leptotrophon, stated that he was considering Trophoninae in a traditional way, to include typical “Tro- phon-like” species, diagnosed. as small, thin spinose hells with a flaring columellar lip. In fact, Kool (1993) stated that Trophoninae is probably a non-monophyletic group and, thus, Houart (1995) considered — that future studies would probably show that the genus Le ptotrophon would have to be transferred from the Trophoninae. The new species fits very well in the diagnosis of Leptotrophon, and is very similar to several species from the Indo-Pacific, including the type species, L. caroae. Leptotrophon otlantious bears the characteristic spiny sculpture (Figures 13-15), round-ovate aperture, and flaring columellar lip (Figure 13). The radula of L. atlan- ticus (Figures 15-20) fits the pattern described for Lep- totrophon, but the lateral/marginal cusps are not as independent of each other, being somewhat fused into a common base (Figure 19). In other ae referred to Leptotrophon by Houart (1995), the lateral/marginal cusps are similar to L. atlanticus [e.g., L. caroae and L. acerapex (Houart, 1986). In addition, two small mar- ginal denticles may be seen on the outer base of the marginal cusps (white arrow in Figure 19), a characteris- tic found in other species of Leptotrophon described by Houart (1995). The protoconch of the species described by Hoaurt (1995) shows considerable variation in’ shape, with rounded-globose, acuminate, or carinate protoconchs present in different species. The type species has a cari- nate protoconch, and Leptotrophon atlanticus has a rounded-globose protoconch (Figure 24). The most similar species are kastoroae Houart, 1997, and L. perclarus Houart, 2001. Both species are larger than L. atlanticus. Leptotrophon perclarus has a taller spire, more acuminate apex, and more inflated last whorl: mM addition it has more numerous and smaller spines. The shell shape of L. kastoroae is almost identi- cal to that of L. atlanticus, but the spines are longer and are more upward-curved. Prey iously to this work. no record of shell microsculp- ture for any species o ; Leptotrophon was available. The protoconc th is always described as smooth. In fact, the protoconch of L atlantic us has an overall smooth ap- pearance (Figure 24), but SEM reveals that only its most apical region is smooth; the last quarter area, close to the varix, bears spiral sets of microscopic granulations (Fig- ures 25-26 \ delicate microsculpture also covers the entire teleoconch surface, including the spiral cords and spines, with axial and spiral microstriae forming a some- what reticulate pattern (Figures 22-23). The operculum of L. atlanticus is similar in its outer surface to those illustrated by Houart (1995) for species of Leptotrophon from the New Caledonian region. Houart (1995) did not describe the operculum of Lepto- trophon internally; the operculum of L. atlanticus has one adventitious layer. In spite of the several common characteristics in the shell and radulae morphology of Leptotrophon atlanti- cus and the species from the Indo-Pacific, the generic allocation herein used should be considered as provi- sional, due to the wide geographic separation between the new species and the other species in the genus Leptotrophon, most of which bearing protoconchs that indicate non-planktotrophic development. All 26 previously described species of Leptotrophon are re- stricted to the Indo-Pacific (Houart, 1995; 1997: 2001); Leptotrophon atlanticus new species is the first record of this genus outside that region. The bathymetry of L. atlanticus, on the other hand, falls within the bathymetric range reported for the Indo-Pacific species (200-720 m). In ae future, direct comparisons with the Indo-Pacific species, especially including characters visible only under SEM, would be most helpful to estab- lish the degree of similarity among these species. ACKNOWLEDGMENTS We are grateful to Mr. Roland Houart (Institute Royal des Sciences Naturelles de Belgique) for comments on the taxonomy of the species. Vinicius Padula (MNRJ) and Franklin Noel dos Santos (Universidade Federal do Para) provided additional bibliography. Dr. Janet Reid revised the English text. Dra. Noemia Rodrigue ’s helped with the SEM photos. We also thank two anonymous reviewers for their comments and suggestions. Special thanks are due to Mr. Antonio Gil Bezerra and Ms. Elisa Gradvohl Bezerra, owners of INACE Shipyard (Indtis- tria Naval do Ceara) for the loan of the fishing boat and to Mrs. José and Marcus Coltro for financial support to the collecting efforts. Conselho Nacional de Desenvol- vimento Cientifico e Tecnolégico (CNPq) provided PROTAX pos-doctoral grant to P.M. Costa. LITERATURE CITED Abbott, R. T. 1974. American Seashells. 2nd ed. Van Nostrand eles Co., New York. 663 p., 24 pls. Bandel, kK. 1984. The radulae of Caribbean and other Meso- shea da and Neogastropoda. Zoologische Verhandelin- gen 214: 1-188 Bouchet, P. and A. Warén. 1985. Revision of the Northeast Atlantic Bathyal and Abyssal Neogastropoda Excluding Turridae (Mollusca: Gastropoda). Societa [Italiana di Malacologia (Bolletino Malacologico), Supplemento 1: 123-296 A. D. Pimenta et al., 2008 Page 25] Clench, W. J. and I. Pérez Farfante. 1945.The genus Murex in the Western Atlantic. Johnsonia 1(17) 1-58. Dall, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mex- ico (1877-78) and in the Carribean Sea (1879-1880), by the U. S. Coast Survey Steamer “Blake”, Lieutenant- Commander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N.. commanding. XXIX. Report on the Mollusca. Part I] Gastropoda and Scaphopoda. Bulletin of the Museum of Comparative Zoology 1S: 1492. Dall, W. H. 1927. Small shells from dredgings off the southeast coast of the United States by the United States Fisheries steamer “Albatross” in 1885 and 1886, Proceedings of the Unieted States National Museum 70(2667): 1-134. [72-85] Fair, R. H. 1976. The Murex Book. Published by the author. 138 pp., 23 pls. Harasewych, M. G. and R. H. Jensen. 1979. Review of the subgenus Pterynotus (Gastropoda: Muricidae) in the Western Atlantic. Nemouria 22: 1-16. Houart. R. 1995. The Trophoninae (Gastropoda: Muricidae) of the New Caledonian Region. Mémoirs du Muséum national d’Histoire naturelle 14(167): 459-498. Houart, R. 1997. Mollusca, Gastropoda: The Muricidae Col- lected During the Karubar Cruise in Eastern Indonesia. Mémoirs du Muséum national d'Histoire naturelle 16 (172): 287-294. Houart, R. 2001. Igensia gen. nov. and eleven new species of Muricidae (Gastropoda) from New Caledonia, Vanuatu, and Wallis and Futuna Islands. pp. 243-269. In: Bouchet. P. and B. A. Marshall. Tropical Deep-Sea Benthos vol. 22. Mémoirs du Muséum national d’ Histoire naturelle, Paris, 406 pp. Kool, S. P. 1993. The systematic position of the genus Nucella (Prosobranchia: Muricidae: Ocenebrinae). The Nautilus 107: 43-57. Ponder, W. F. 1972. Notes on some Australian genera and species of the family Muricidae (Neogastropoda). Journal of the Malacological Society of Australia 2: 215-248. Radwin, G. E. and A. D’Attilio. 1976. Murex Shells of the World. Stanford University Press, Stanford, xi + 254 pp., 32 pls. Rios E. de C, 1994. Seashells of Brazil. 2nd ed. Museu Ocea- nogratico Prof. E.C, Rios da Fundagao Universidade de Rio Grande, Rio Grande. 368 p., 113 pls. Rosenberg, G. 2005. Malacolog 4.1.0: A Database of Western Atlantic Marine Mollusca. [WWW database (version 4.1.0)] URL http:/Avww. malacolog.org/. Sunderland, K. and L. Sunderland. 1992. Caribbean Murici- dae part Il. American Conchologist 20(3): 14-15. Vaught, K. C. 1989. A Classification of the Living Mollusca. American Malacologists, Inc. Melbourn. 195 p. Vokes, E. H. 1970. Cenozoic Muricid of the Western Atlantic Region Part V - Pterynotus and Poirieria. Tulane Studies in Geology and Paleontology §: 1-50. Vokes, E. H. 1992. Cenozoic Muricidae of the Western Atlantic region. Part IX - Pterynotus, Poirieria, Aspella, Dermomurex, Calotrophon, Acantholabia, and Attiliosa; additions and corrections, Tulane Studies in Geology and Paleontology 25: 1-108. THE NAUTILUS 122(4):252-258, 2008 Page 252 A new species of Chlamydoconcha Dall, 1884, from southeastern Brazil (Bivalvia: Chlamydoconchidae) Luiz Ricardo L. Simone Museu de Zoologia da Universidade de Sao Paulo Caixa Postal 42494 04299-970 Sao Paulo, BRAZIL Irsimone@usp.br ABSTRACT The second species in the genus Chlamydoconcha _ is described. Chlamydoconcha av alvis new species, occurs off the coast of Rio de Janeiro coast, in southeastern Brazil. The new species has very reduced valves and a mantle surrounding the entire body, two features of the genus. The outer surface of the mantle lacks papillae except for a single one located close to the excurrent siphon. These are distinctive characters of Chlamydoconcha orcutti Dall, 1884, from the eastern Pacific coast of North America, the single other known species of the genus. Some of the more interesting anatomical characters of the new species are: posterior pair of retractor muscles of foot free from valves, absence of adductor muscles, gastric style sac totally separated from intestine, and the presence of a single (excurrent) siphon. Additional keywords: Anatomy, western Atlantic, Rio de Janeiro INTRODUCTION The genus Chlamydoconcha Dall, 1884 (type species by original designation: C. orcutti Dall, 1884) has been known to be monotypic. Chlamydoconcha orcutti occurs from California to western Mexico (Carlton, 1979: Morton, 1981). The species is Dennen: by reduction of the shell, which is restricted to the anterior region of a spherical mantle cover; the mantle outer surface has many, somewhat equidistantly distributed papillae. After the original description, further anatomical studies of C. orcutti were done by Bernard (1897) and Morton (1981). \ sample collected by biologist Vinicius Padula on the coast of Rio de Janeiro was sent to the author for study. The analysis of the material revealed a new spe cles of Chlamydoconcha, formally described herein. This paper is also the first discovery of the genus in the Atlantic Ocean, representing the second known species in the genus. The present description also includes a detailed matomy, which is discussed in comparison to C. orcutti Bernard, 1897: Morton, 1981) The taxonomic allocation of the genus Chlamydo- concha has been problematic. It has been included in the Galeommatidae (Morton, 1981), but full family status has been assigned (Chlamydoconchidae, Bernard, 1983), as part of the Galeommatoidea. Full superfamily status was also considered (Chlamydoconchacea, Keen, 1969). The Galeommatoidea, are mostly mollusks with usual bivalve shells, but may also include highly modi- fied, slug-like animals, with internal and reduced shells. MATERIALS AND METHODS The specimen was delivered preserved in 70% EtOH, A photo of the living specimen was taken before pre- servation. The dissection of the preserved animal was performed by standard techniques, under a_ stereo microscope, with the specimen immersed in the alcohol. All dissection steps were also photographed (e.g., Figures 3-5), Drawings were made with aid of a camera luc a, Abbreviations used in figures are: an, anus; au, auri- cle; by, byssal gland: ee, vill ciliary connection; ce, cere- bral ganglion: co, ce webieos -visceral connective; dd, ducts to digestive diverticulae; dh, dorsal hood; di, inner demibranch; do, outer demibranch; es, esophagus; fg, gill food groove; fm, posterior foot retractor muscle; fr, anterior foot retractor muscle; ft, foot; ga, genital aper- ture; gi, gill; go, gonad; gs, gastric shield; in, intestine; ip, inner hemipalp; ki, kidney; mb, mantle border; mo, mouth; mt, mantle: ne, nephropore; op, outer hemi- palp; pa, pedal aperture of mantle; pe, pericardium; pg. pedal ganglia; pl, pallial papilla; pm, pallial muscles: pp. palp; rt, rectum; sh, shell; si, excurrent siphon; ss, style sac; st, stomach; ty, typhlosolis; ve, ventricle: vg, visceral ganglia; vm, visceral mass. Institutional abbreviation: MZSP; Museu de Zoologia da Universidade de Sao Paulo, Brazil. SYSTEMATICS Chlamydoconcha avalvis new species (Figures 1-20) L. R. L. Simone, 2008 Page 253 Figures 1-9. Chlamydoconcha avalvis Holotype photos. 1-2. Living specimen dorsal and lateral views, photo Vinicius Padula 3-5. Preserved specimen. 3. Dorsal view. 4. Ventral view. 5. Left view, right mantle lobe partially removed and deflected anterior! right gill deflected upM ards. 6. Right valve, outer view (transversal section artificially done). 7. Same, inner view. 8. Left valve, inne: view. 9. Right valve, ventral view of its posterior, concave region Scale bars 15 = 2 mm: 6-9 = 0.5 mm Page 254 THE NAUTILUS, Vol. 122, No. 4 10 Figures 10-13.) Chlamydoconcha avalvis anatomy. 10. Left view of entire animal, right mantle lobe partially removed and deflected anteriorly (right in Figure). 11. Region of right valve, internal surface of mantle removed, showing pallial muscles (pm) originating in valve, some adjacent structures also shown, 12. Gill, transversal section in its middle region. 13. Right palp, outer hemipalp deflected anteriorly, a short portion if inner demibranch also shown. Scale bars = 1 mm, Diagnosis: Species with a single papilla close to excur- rent siphon. Anterior pallial gland shallow. Internal shell size about 10% of mantle surface; with rounded, almost | posterior end. Anterior pair of pedal retractor muscles a branch originated from shell. Gastric Maly chan l and style Sac Narrow and long. Description: = Srevy (Ficures 6-9): Reduced, inequi- valve, occupying about 1/10 of mantle, embedded into mantle anterior region (Figure 10, sh). Length approxi- mately 4 times width. Color white, opaque. Outline softly irregular. Both valves asymmetrical; left valve about 4 shorter than right’ valve (Figure 8) (this may be L. R. L. Simone, 2008 Page 255 Figures 14-20. Chlamydoconcha avalvis anatomy. 14. Left view of entire animal emphasizing location of digestive tract and topology of main muscles, ganglia and glands, animal artificially represented as transparent. 15. Midgut as in situ, right view. 16. Same, slit longitudinally to expose inner surface. 17. Renopericardial structures and region, right auricle artificially disconnected from gill and deflected upwards, a transversal section of indicated level of right kidney also shown. 18. Cerebral ganglia, posterior- slightly right view. topology of esophagus also indicated. 19. Pedal ganglia, right and slightly posterior view. 20. Visceral ganglia, right and slightly posterior view. Scale bars = 1 mm. abnormal). Shape somewhat deformed and irregular; flattened, planar. Prodissoconch rounded, sub-termi- nal: located in middle of anterior fifth of valve length; shape semispherical, with small dorsal bulging portion; 0.26 mm long, 0.31 mm height. Outer surface some- what irregular with strong commarginal undulations and with rounded. concave impressions: with ventral edge elevated (Figure 9). Calcareous concretions close to periphery on right valve (Figure 8). Periostracum extending about 1/3 beyond calcareous portion of each valve, wider dorsally; color yellowish, transparent. Hinge edentulous. Ligament small, restricted to umbonal region, relatively wide (Figures 6-7), pale brown; resili- fer absent. Inner surface glossy. Scar of anterior retractor muscle of foot occupying about 1/5 of inner surface, 3 times longer than wide, located just posterior to umbo- nal concavity. MantTLE (Ficures 1-5, 10): Surrounding body almost completely, spherical in contracted condition (Figures 14). Color pale cream, translucent (living and_preser- ved). Outer surface smooth and simple, lacking papillae Page 256 Pedal aperture ventral, longer anteroposteriorly (Fig- ures 4, 10); aperture length spank half total mantle length. Edges of pedal aperture thick, simple, with winalations: thicker anteriorly. Anterior gland as a blind-sac, located in anterior, median region, about 1/3 of animal height from anterior end of pedal aperture (Figure 4, ag); size equiva- lent to 1/30 of mantle outer surface; its aperture central, with about 4 of gland size. Excurrent siphon cylindrical, small, papilla- like, located about half of animal height from posterior end of pedal aperture (Figures 3, 10, si): length about 1/20 of animal length; internal surface smooth, simple. Single papilla located about 1/5 of animal length dorsal to excurrent siphon (Figures 1, 3, pl), on median line, solid, size about half of that of siphon. Pair of small, low, bulging projections slightly dorsal to anterior gland, corre sponding with shell ne Mantle relatively thik. mostly hollow, sponge-like. Mantle inner surface smooth, simple (F igure 5). Main Muscie System (Ficures 11, 14); Adductor muscle not seen, possibly immersed in thin layer of visceral dorsal muscles. Pair of anterior pedal retractor muscles originate about 1/3 from inner surface of valves (scar described above), and about 2/3 splayed by antero- dorsal region of visceral sac; gradually becoming thicker towards ventral, up to anterior half of pedal oneal re- gion. Pair of posterior pedal retractor muscles somewhat similar to anterior pair; originating in dorsal visceral sac side about 4 posterior from that of anterior pair. Thin layer of pallial muscles splayed by mantle like a net; mi uinly concentrated anteriorly, inserting in anterior pair of pedal retractor muscles, in level just anterior to palps. Foor anpd Byssus (Ficures 4, 5, 10, 14): Foot narrow, longer antero-poste sriorly: leno about half of animal le neth; width about 1/5 of animal width; projected ante- riorly at about 4 of animal le a Anterior region some- what pointed. Byssal gland a narrow furrow located subterminally, in posterior region of foot ventral medial line; about 1/7 of foot length. Byssal gland thin, hollow, chamber depth of about 1/5 of foot length (Figure 14, by). No Reis found. PaLLIAL Cavity (Ficures 5, 10-13): Surrounding almost entire space between mantle and visceral sac, except for a dorsal portion correspondent to 1/10 of visceral sac surface connected to mantle. Gill eulamellibranch, heterorhabdic, occupying about half of pallial cavity, mainly in dorsal region (Figure 10), about two times longer than wide. Outer demibranch slightly triangular, about 2/3 of inner demibranch; anterior region becom- ing abruptly narrow, ending about 1/8 of total gill length- posterior to inner demibranch anterior end. Inner demibranch anterior end slightly rectangular, ending be- tween hemipalps. Gills gradu: ully narrowing towé ards pos- terior, up to some what pointe ral posterior e nad. About 1/4 of each gill (their posterior region) Tree from visceral mass, connected with each other by cilia. Cilia connect outer lamellae of outer demibranch with mantle and inner lamellae of inner demibranch with visceral sac THE NAUTILUS, Vol. 122, No. 4 (Figure 12, ee), same ciliary connection between both inner demibranchs in their region posterior to visceral mass. Connection among gill flame nts by aligned longi- tudinal tissue rods equiv alent’ in width to f Slnmnenie: eae longitudinal rod separated f rom neighbor rods by dis- tance equivalent to 5 filaments. Ve nizal edge of outer demibranch simple; filaments very thin tclsont 1/50 of gill width), outer connection mostly dorsal. Inner demibranch filaments a little shorter than inner demibranch itself; ventral edge with food groove. Inner gill connection to visceral mass dislocated ventrally, separated from remaining dorsal gill connection by dis- tance equivalent to half gill width (Figures 5, 12). Palps (Figure 15) with size equivalent to 1/10 of that of gill; category IT (Stasek, 1963). Hemipalps similar to each other; ventral half tall, slightly triangular; dorsal half narrow (about “4 of ventral half), emeath, surrounding anterior insertion of inner demibranch. Inner surface of palp (ventral half) with uniform, transversal folds, about 20 folds in each hemipalp; more distal folds shorter, weakly arched, folds gr adually becoming longer towards medial, dorsal region of folds be coming narrower and strongly arched, ee ming a folded dorsal furrow in direc- tion to mouth: ventral ead of each folds rounded: dorsal end weaker; a smooth, narrow area surrounding entire edges of hemipalps (Figure 13). Both palps separated from each other bya dicaned equivalent to half of lon- ger portion of palp length. Mouth surrounded by anteri- or and posterior relatively tall lips, inner surface smooth. VisceRAL Mass (Ficures 5, 14): Bulging, spherical; sepa- ration with foot somewhat distinct. Gonad color cream, surrounding most of visceral structures, occupying about S0% of outer region. Genital aperture a small slit located about 1/20 of visceral height from dorsal edge and from nephropore (Figures 14, 17, ga); genital duct not dis- cernible. Digestive diverticula restricted to central area of anterior region; color pale greenish beige. Reno- pericardial structures occupying a bitit 1/10 of visceral volume, located in posterior region of dorsal surface. CIRCULATORY AND EXCRETORY SysTeMS (FiGures 14, 17): Heart of about 1/20 of visceral volume; located anterior to kidney; length about 1/8 of total length. Auricles tri- angular, insertion with ctenidial veins about “4 of their length, located in posterior quarter of gill. Connection to ventricle longitudinal, lateral, with about half of ventri- cle length. Ventricle occupying about entire pericardial length. Kidneys white, extending from pericardium pos- terior end to area equivalent to pericardial length toward poste rior region, Each kidney about three times longer than tall, cosy solid except for inner flattened faman running longitudine uly long central region. Each nephro- pore a minute slit locate -d just anterior to origin of pair of posterior pedal retractor muscles; inside excurrent chamber of outer demibranch. Dicestive System (Ficures 14-16): Palps described above (pallial cavity). Esophagus with about 2/3 of dis- tance between pi alps i in width; le neth about 1/5 of that of L. R. L. Simone, 2008 Page 257 visceral mass; inner surface smooth. Stomach positioned transversal, somewhat perpendicular to esophagus, run- ning towards right; narrowing gradually (Figure 15); estimated volume about 1/20 of that of visceral mass: Type IV (Purchon, 1958). Stomach inner surface with pair of low, narrow folds located transversally in esopha- geal insertion (Figure 16). Dorsal U- shape sd furrow ocated just posterior to wala cig insertion (concavity stomach, with about 4 of stomach height; its aperture as eft end of U-shaped furrow. Ducts of digestive diverti- ateral gastric side: left pair slightly longer than right pair. Typhlosole very wide on origin of sty Te sac, narrow- ing relatively abruptly, running longitudinally in style sac eft side as narrow, low fold. Gastric shield with about I/S of internal gastric surface; located inside U-shaped furrow. Style sac totally separated from intestine; long and narrow: width about 70% of that of esophagus; run- ning somewhat straight backwards, ending in posterior wall of visceral mass. Digestive diverticula described above (visceral mass) Intestine originating in right side of style sac origin; inner surface aHaorh, simple: initially as wide as stomach, gradually becoming narrow up to 1/3 of its original width after a distance equivalent to that of esophagus. Intestine performing tight loops as shown in Figure 14; after this, performing “wide, sigmoid loop, in such superior branch edges superior surface of viscer- al mass, along median line; running towards posterior. Anus sessile, simple: located at base of excurrent siphon. GenitaL System: Gonad described above (visceral mass). Genital pores represented by small slits equivalent in size to nephropore (Figures 14, 17, ga), located about 1/20 of total animal length from nephropore, slightly posterior and ventral. No dications on brooding in gills was observed. CENTRAL Nervous System (Ficures 14, 18-20): Cerebral ganglia (Figure 18) located a short distance dorsal to mouth; each one with size equivalent to 1.5 esophagus diameter. Cerebral commissure narrow, length equiva- lent to each ganglion. Pedal ganglia (Figure 19) located in middle between cerebral ganglia and posterior end of foot: both ganglia completely conhected with each other along median line, forming a single, spherical mass of equivalent size of each cerebral ganglion. Visceral gang- lia (Figure 20) located just ventral to origins of posterior pair of pedal retractors; size equivalent to about S0% of that of cerebral ganglia, visceral commissure very short, ganglia almost touching each other. Cerebro- visceral connective very narrow, running through gonad Figure 14, co). Measurements: Animal length = 15 mm; valve = 3.7 by 1.2 mm. Holotype: MZSP._ 8631S, Vinicius Padula col., 05 March 2006. posterior). Dorsal hood triangular, located at left side of cula in two ae each pair located in middle region of Type Locality: Brazil, Rio de Janeiro, Cabo Frio, Ilha Comprida, 22°51/47” S, 41°56'35" W, about 6 m depth, under rocks. Distribution: Only known from the type locality. Etymology: The specific epithet refers to the apparent absence of the shell valves, which are virtually invisible in the living animal; a combination of the Latin negative prefix a and the noun valvis. Comparative Remarks: = Chilamydoconcha avalvis has the external surface of the highly developed mantle practically lacking papillae ( (Figures 1-4). This is the main character differentiating the species from the Pacific congener C. orcutti, which has a richness of papillae in the outer mantle surface, somewhat equidis- tantly disposed (Dall, 1884; Bernard, 1897; Williams, 1949: Morton, 1981: fig. S). However, a single papilla is present in C. avalvis, close to the excurrent siphon; C. orcutti also possesses a differentiated papilla in the same position (Bernard, 197: fig. 3), which was named “defensive papilla” by Morton (1981). Anatomically, both Chlamydoconcha show similar organization. Mantle enlargement, foot features, posi- tion of the valves and main muscles, and internal fea- tures of glands and digestive tubes, are similar in the two species. The main anatomical differences, beyond the above mentioned papillae, are: The shell is proportional- ly smaller in C. avalvis (about 1/10 of mantle, Figure 14) than that of C. orcutti (about 1/6 of mantle). Although the prodissoconch (Figures 6-S) is very similar in both species, the posterior end of the shell of C. avalvis is more squarish than that of C. orcutti; in which the pos- terior end of the shell is pointed (Bernard, 1897: fig. 15; Morton, 1981, figs. 4-5). The anterior gland of C. avalvis is a blind sac, its internal chamber is small and short, practically with the same thickness of f the surrounding mantle (Figure 4, ag); on the other hand, that of C. orcutti Gerard 1894: “cheminée dorsale”) has a deeper empty chamber directed posteriorly ( (Bernard, 1894: fig. 19, X), more recently, this gland was desig- nated “pheromone organ” (Morton, 1981, fig. 10), and described with similar characters of C. avalvis. The anterior pair of pedal retractor muscles has a branch originated from the inner surface of the valves in C. avalvis (Figure 14, fr); this is not described for C. orcutti (Bernard, iy fig. 20, mp), although men- tioned by Morton (1981 The midgut organization of C. avalvis (Figures i. is quite different from that of C. orcutti (Bernard, 1894: figs 9, 19: Morton, 1981, fig. 24) in several details, the main characters are: the narrower and longer gastric style sac of C. avalvis, while that of C. orcutti is wider and shorter (about 1/3 of vis- ceral sac length); the stomach is also narrower and smal- ler in C. avalvis than that of C. orcutti; the intestinal loops are differently performed in both species, and in C. avalvis it is apparently narrower Page 2558 THE NAUTILUS, Vol. 122, No. 4 Although the living animal of C. avalvis (Figures 1-2) was not observed crawling, it is possible that it also has an anterior projection of the mantle like that of C. orcutti (Bernard, 1894: figs.10, 11; Williams, 1949; Morton, 1981), as the mantle arrangement of that region is taller and wavy. The presence ofa single siphon close to the anus shows that the siphon is excurrent; as no incurrent siphon is present, the conclusion that water intake takes place through the pedal aperture (Morton, 1981). This feature is also found in other galeommatids, such as Kellia porculus Pilsbry, 1904: Scintilla nitidella Habe, 1962 (Morton and Scott, 1989, figs. 3, 18). DISCUSSION Discovery of the second species in the genus Chlamydo- doncha fits the description of the genus by Dall (1554). The anatomical characters of the C hlamydoconcha spe- cies are quite modified, even if considered under the light of the extraordinary suite of modifications exhibited by the Galeommatoidea (Woodward, 1893; Morton, 1981: Bieler and Mikkelsen, 1992). The reduction of the shell of Chlamydoconcha is apparently the most extreme in all Bivalvia; its interiorization inside the man- tle is also found in other genera, e.g., Galeomma Turton, 1825, Ephippodonta Tate, 1889. (Woodward, 1893: Liitzen and Nielsen, 2005), and Divariscintilla yoyo Mikkelsen and Bieler, 1989. All these genera and spe- cies, however, have proportionally larger valves. The foot is an important comparative character in Galeommatoi- dea. The “hanging” foot and the flower-like organ are some of the main characters (Bieler and Mikkelsen, 1992; Jespersen and Liitzen, 2006); Chlamydoconcha possesses at least the first of these two characters. A molecular study (O Foighil et al., 2001) places Chlamy- doconcha as terminal taxa inside the Galeommatidae, a similar result of the morphological approach (Bieler and Mikkelsen, 1992). A dwarf male has been described for Chlamydoconcha orcutti (Morton, 1981), however, one has not been found so far in C. avalvis. ACKNOWLEDGMENTS A special thank to Vinicius Padula, Museu Nacional, Universidade Federal do Rio de Janeiro, by collect and donation of the lot examined herein. This study is sponsored in part by FAPESP (Fundagao de Amparo a Pesquisa do Estado de Sao Paulo), project no. O4/ 02333-S. LITERATURE CITED Bemard, F. 1897. Anatomie de Chlamydoconcha orcutti Dall, lamellibranche a coquille interne. Annales des Sciences Naturelles, Zoologie et Paléontologie 4: 221-252 + pls. 1-2. Bernard, F. R. 1983. Catalogue of living Bivalvia of the Eastern Pacific Ocean. Dept. of Fishexes and Oceans. Ottawa, 102 pp. Bieler, R. and P. M. Mikkelsen. 1992. Preliminary phylogenetic analysis of the bivalve family Galeommatidae. American Malacological Bulletin 9: 157-164. Carlton, J. T. 1979. Chlamydoconcha orcutti Dall: review and distribution of a little-known bivalve. The Veliger 21: Dall, W. H. 1884. A remarkable type of mollusk. Science 4(76): 50-51. Jaspersen, A. and J. Liitzen. 2006. Reproduction and sperm structure in Galeommatidae (Bivalvia, Galeommatoidea). Zoomorphology 125: 157-173. Keen, A. M. 1969. Superfamily Chlamydoconchacea, Dall, 1884. IN Moore, R.C. [Ed.] Treatise on invertebrate pale- ontology. Part N2, Mollusca 6, Bivalvia. The Geological Society of America and University of Kansas Press, Kansas, 573 pp. Liitzen, J. and C. Nielsen. 2005. Galeommatid bivalves from Phuket, Thailand. Zoological Journal of the Linnean Society 144: 261-308. Mikkelsen, P. M. and R. Bieler. 1989. Biology and comparative anatomy of Divariscintilla yoyo and D. troglodytes, two new species of Galeommatidae (Bivalvia) from sto- matopod burrows in eastern Florida. Malacologia 31: 175-195. Morton, B. 1981. The biology and functional morphology of Chlamydoconcha orcutti with a discussion on the taxonomic status of the Chlamydoconchacea (Mollusca: Bivalvia). Journal of Zoology 195: 81-121. Morton, B. and P. H. Scott. 1989. The Hong Kong Galeomma- tacea (Mollusca: Bivalvia) and their hosts, with descrip- tions of new species. Asian Marine Biology 6: 129-160. O Foighil, D., R. Jennings, J.-K. Park, and D. A. Merriwether. 2001. Phylogenetic relationships of mid-oceanic ridge and continental lineages of Lasaea spp. (Mollusca: Biv alvia) in the northeastern Atlantic. Marine Ecology Progress Series 213: 165-175. Purchon, R.D. 1958. The stomach in the Eulamellibranchia;: Stomach Type IV. Proceedings of the Zoological Society of London 131 : 487-5235. Stasek, C.R. 1963. Synopsis and discussion of the association of ctenidia and labial palps in the bivalved Mollusca. The Veliger 6: 91-97. Williams, W. 1949. The enigma of Mission Bay. Pacific Discov- ery 2(2): 22-23. Woodward. M. F. 1893. On the anatomy of Ephippodonta macdougalli, Tatte. Proceedings of the Malacological Society of London 1: 20-26 + pl. 2. THE NAUTILUS 122(4):259-260, 2008 Page 259 Research Note Sinistral Campeloma decisum (Say, 1817) (Gastropoda: Viviparidae) from the Fox River, Illinois Campeloma (Gastropoda: Viviparidae) are a group of ovoviviparous, prosobranch snails endemic to North America east of the Rocky Mountains (Burch, 1989). These snails are known to burrow in mud or sand in freshwater streams and lakes and feed on carrion (van der Schalie, 1965; Burch, 1989). Campeloma spp. from southern North America typically nee sexually but those from the northern United States and Canada are parthenogenetic due to the scarcity of males in this re- gion (van der Schalie, 1965). Shells of Campeloma are moderately thick, conical, and imperforate ak a smooth surface and rounded whorls, and although normally dex- tral, sinistral specimens are occasionally f found (Baker, 1928: Burch, 1989). In fact, Call (1886) pointed out Rafi- nesque’s "type" of Campeloma crassulum Rafinesque, 1819. from the Ohio River was sinistral, noting that the shell had "four whorls of the spire reversed." Sinistral individuals show not only a reversal in shell orientation but also in organ placement (Savage, 1935). Sinistral Campeloma have been found throughout eastern North America. Call (1880) recorded C. decisum (Say, 1S17) (as C. integrum and C. rufim) from the Erie Canal at Mohawk, New York, Pilsbry (1897) ) reported sinistral C. decisum from the eee River at Fort Edward. New York. and Ancey (1897) commented on sinistral C. decisum from New York but gave no specific location. Bickel (1966) examined sinistral C. crassuluwm from the Ohio River at Louisville, Kentucky, Goodrich (1939) discussed sinistral C. geniculum (Conrad, 1834) from the Ogeechee River in Georgia, and Lee (2008) figured a sinistral C. limum (Anthony, 1860) from the Altamaha River at Doctortown, Georgia. Baker nee commented on a “reversed” C. decisum (as C. rufum) in a private collection, gave dimensions of a reversed C. decisum, and figured a sinistral C. decisiwm (as C. subsolidum), but did not give locality data for any of the specimens examined. Sampson ( (1916) stated that he had sinistral C. decisuwm (as C. subsolidum) in his collection from Flat Creek in Pettis County, Missouri, and Call (1886) referred to sinistral C. descisum (as subsolidum) from a slough near Fort Dodge, Lowa, and figured a reversed C. decisum (as C. obesum) from Lewis. Iowa. Baker (1928) figured sinistral C. decisim (as C. integrum, C. rufum, and C. brevispirum) from three different locations in Wisconsin (Wisconsin River near Merrimack, Sturgeon Bay at Sturgeon Bay, and Mirror Lake presumably near Baraboo), and Haas (1939) reported on a sinistral C. decisum (as C. integrum) from the Kankakee River near Shelby, Indiana. In Illinois, sinistral C. decisum (as C. integrum and C. rufum) have been reported from the Salt Fork Vermilion River near Homer (van Cleave, 1936), the Des Plaines River in the Chicago area (Lee, 2008), and Jackson Park Lagoon in Chicago ( Hand, 1928; Meyer, 1928; Haas, 1939). Sinictral uterine young in Campeloma have been reported by several authors, including Call (1880), Pilsbry ( (1897), Hand (1928), van Cleave (1936), and Haas (1939). Both desteal and_ sinistral Seat produce sinistral embryos (Hand, 1928; Haas, 1939 Sinistrality might re ssult from either embryological is turbances (e.g., crowded uteruses or damaged eggs) that have no genetic basis, or individual mutations destined to disappear in the population because copulation between dextral and sinistral snails is impeded by me- chanical incompatibility (Call, 1SSO; van Cleave, 1936; Cazzaniga and Estebenet, 1990). However, because some degree of assortative mating occurs in gastropods, sinistral “shail could become reproductiv ely isolated (Cazzaniga and Estebenet, 1990); also, because many Campeloma lineages are parthenogentic, sinistral snails could become unconstrained by mating compatibility (Mattox, 1938: van der Schalie, 1965). There appears to be a progressive reduction in the percentage of sinistral individuals from uterine young to adults sath only a few individuals reaching sexual maturity (van Cleave, 1936). This high degree of mortality might occur as the result of morphologic: al or phy siclogical abnormalities (van Cleave, 1936; Bickel, 1966). We here report on sinistral C. deciswm from the Fox River basin near Algonquin, Illinois. Nineteen specimens of C. decisum from “Mill Pond (near creek), Algonquin, Ilinois,” were found in the University of Illinois Museum of Natural History Mollusk Collection, C ane Urbana (UIMNH 1828S): no date was given for this lot but the collector, the Rev. W. A. Nason, We din 1921. In addition, a relict sinistral C. decisum shell was collected by JST while conducting a freshwater mussel survey in the Fox River at Buffalo Park Forest Preserve near Al- gonquin (42.1486° N, 88.2900° W), Kane County, Ili- nois, on 25 July 2007. This specimen was e xtracted from silt-compacted gravel in an impounded area of the river and has been deposited in the Hlinois Natural History Survey Mollusk Collection, Champaign (INHS 31862). The Fox River has experienced sub-substandard water quality conditions prior to the passage of the Clean Water Act and has encountered habitat che inges (e.¢., increased siltation and substrate compaction ) \ due to he presence of lowhead dams (Santucci et al., 2005; Tiemann et al 2007). Because these physicochemical changes have been shown to alter freshwater snail assemblages (Burch, 1989), the population of sinistral C. decisuwm in the Algonquin area might be extirpated. Fieldwork will continue in an attempt to document live individuals. Page 260 THE NAUTILUS, Vol. 122, No. 4 ACKNOWLEDGMENTS Funds were provided by the Illinois Department of Transportation. A. Kuhns (INHS) and J. Griesbaum (INHS) assisted in the 2007 sampling. T. Stewart (Iowa State University) assisted in identification of the 2007 specimen. G. Levin (INHS), D. Thomas (INHS), B. Tiemann, and two anonymous reviewers offered constructive criticism. LITERATURE CITED Ancey, C. F. 1897. On some sinistral land shells. The Nautilus 10; 104-105 Baker, F. C. 1902. The Mollusca of the Chicago area. Part IL. The Ce tapode Bulletin of the Chicago Academy of Science 3: 131-418 + 9 plates. Baker, F.C. 1928. The fresh water Mollusca of Wisconsin. Part I. Gastropoda. Bulletin of the Wisconsin Geological and Natural History Survey 70(2), i-xx + 1-507 + 5 28 plates. Bickel, D. 1966. Campe loma crassula with reversed whorls. The Nautilus 79: LO7—108. Burch, J. B. 1989. North American freshwater snails. Malaco- logic: al Publications, Hamburg (Michigan), vii + 365 pp. Call, R. 1880. Reversed Mel vinthones: American Naturalist 14: ae Call, R. E. 1886. On the genus Campeloma, Rafinesque, with a revision of the species, recent and fossil. Bulletin of the Washburn College 149-165, 4 plates. Cazzaniga, N. J. and A. L. Estebenet. 1990. A sinistral Poma- cea canaliculata (Gastropoda: Ampullariidae). Malacolog- ical ae 23: 99-102. Goodrich, C. 1939. Certain mollusks of the Ogeechee River, ee The Nautilus 52: 129-131. Haas, F. 1939. Reversed specimens of Campeloma from the Chicago area. Zoological Series of Field Museum of Natural History 24: 93-94. Laboratory of Natural History 1: Hand, E. E. 1928. Sinistral Campeloma. The Nautilus 41: 106-107. Lee, H. G. 2008. Jacksonville Shell Club website. http://www. jaxshells.org/reverse 1.htm accessed 22 May 2008. ae N. T. 1938. Morphology of Campe loma rufum, a parthenogenetic snail. Joumal of Morphology 62; 243-261. Meyer, E, 1928. Finding a left-handed Campeloma. The Nau- tilus 41: 107. Pilsbry, H. A. 1897. Campeloma decisum Say, reversed. The Nautilus LO; 11S. Sampson, F. A. 1916. Reversed or sinistral shells. The Nautilus 29: 128-129. Santucci, V. J., Jr, S. R. Gephard, and S. M. Pescitelli. 2005. Effects of multiple low-head dams on fish, macroinverte- brates, habitat, and water quality in the Fox River, Illinois. North American Journal of Fisheries Management 25: 975-992. Savage, A. E. 1938. A comparison of the nervous system in normal and sinistral snails of the species Campeloma rufum. American Naturalist 72: 160-169. Tiemann, J. S., H. R. Dodd, N. Owens, and D. H. Wahl. 2007. Effects of lowhead dams on unionids in the Fox River, Illinois. Northeastern Naturalist 14; 125-138. van Cleave, H. J. 1936. Reversal of symmetry in Campeloma rufum, a fresh-water snail. American Naturalist 70: 567-573. van der Schalie, H. 1965. Observations on the sex of Campeloma (Gastropoda: Viviparidae). Occasional Papers of the Museum of Zoology, University of Michigan, 641; 1-15. Jeremy S. Tiemann Kevin S. Cummings Division of Biodiversity and Ecological Entomology Ilinois Natural History Survey 1816 South Oak Street Champaign, [IL 61520 USA jtiemann@inhs.uinc.edu THE NAUTILUS 122(4):261-263, 2008 Page 261 Book Reviews Freshwater Mussels of Alabama & the Mobile Basin in Georgia, Mississippi © Tennessee Williams, James D., Arthur E. Bogan, and_ Jeffrey T. Garner. 2008. Freshwater Mussels of Alabama & the Mobile Be isin in Georgia, Mississippi - Tennessee. University of Alabama Press, Tuscaloosa, xv + 1—908, including numerous text figures and maps, many in color. ISBN-13: 978-0-8173-1613-6 (cloth: alk. paper): ISBN-10: 0-S173-1613-6 (alk. paper) 9 x 11.5 inches. Hardback; 10 Ibs. $70.00 from publisher and several booksellers: possibly less on eBay. Over the last decade or two the awareness of the Ameri- can populace and its policy-makers with the country’s in- digenous flora and fauna has been stirred to an unprecedented degree. Government has responded to a new culture of concern over environmental change and the conservation of natural communities, and one of the most important consequences of riding this zeitgeist has been the commissioning of scientists to elucidate’ the cur- rent state of our biota. ‘Conspicuous among the products of this “green revolution” is a watershed of works treating the naiad fauna of either a political unit (e.g. state) or a major river system. Except possibly for Constantine Rafin- esque’s epiph: iny on the banks of la Riviere Ohio has there been such a celebration of this natural resource! Preceded by recent works treating the biology of pearly freshwater mussels of several eastern American regions, most conspicuously the state of Tennessee and the Appalachicola River system (Georgia, Alabama, Florida), Williams, Bogan, and Gamer ice e tackled the most extensive fauna yet considered, that of Alabama and the entire Mobile Basin, but, based on other works of this contemporary genre, as we shall see, the treat- ment of those 178 species-level taxa, were it by tradition- al measure, only partially accounts for the ‘prodigious metrics (e.g. weight) c captioned above. The work is organized into a foreword, acknow- ledgements, institutional abbreviations, 16 chapters, an appendix (North American naiad type catalogues), a glossary, bibliography, and index. Certain observations can be made as one moves through the work. Introductory comments place Alabama and its mussel fauna in a broader context and present the grim reality of habitat degradation. resource depletion, extirpation, and extinction. No less than 23 reviewers are acknowledged for vetting this opus: workers in over 30 museums on other institutions were cited as collaborators, and dozens of field workers contributed their labors. Thanks are also offered to molecular geneticists, whose work underpinned many of the taxonomic innovations mentioned later. Wit A Foreworp By E. O. WILSON FRESHWATER MUSSELS of ALABAMA & the MOBILE BASIN IN GEORGIA, MISSISSIPPI & TENNESSEE “\_ JEFFREY T. GARNER There follows an historical review of naiad work in the state. The contributions of the feuding Quakers Isaac Lea and Timothy Conrad, of C. T. Simpson, H. H. Smith, and H. D. Athearn, the latter two being the dedicatees of the book, and many others are presented briefly. Chapter 3 spans 25 pages and presents an analysis of the inland waters of Alabama and a Mobile Basin, which support more aquatic biodiversity than any other area of comparable size on the continent. The geography, geology, hydrology, and, regrettable degradation of these water- ways (damming. canalization, etc.) is discussed in detail. The use of archival maps and photographs along with present-day images provides a starkly heuristic backdrop. Short chapters basically tabulate mussel taxa by con- stituent watercourse in the post-European and archaeo- logical record. A section on the commercial use ol mussels and their pearls is nicely illustrated and again reinforces the theme of resource depletion Chapter 7 is an historical review of naiad conservation efforts in the state, which have been rather extensive, particularly in the last decade: it concludes with a tabulation of the 48 Alabama species listed as endangered or threatened as Page 262 THE NAUTILUS, Vol. 122, No. 4 * January, 2006, under provisions of the federal a ingered Species Act. Twenty pages are devoted to the ecology and life his- tory of the naiads. The topics are treated with thorough- ness and involve aspects of habitat (and its degr aston). feeding, predation, competition, parasitism, and the unique reproductive and larval strategies if these mollusks—including anatomic and behavioral contri- vances to optimize host fish infestation. Much recent work is brought to bear on these topics. Shell morphology and higher (ordinal, suprafamilial, familial) classification are dealt with succinctly; the latter with the most current systematic insights. Chapter 11 explains the format of the accounts in the taxonomic section. These headings are uniform and clear- ly indicated: Scientific and Common Name (each epithet initiated in upper case!); Illustrations; Description of Shell, Soft Anatomy, Glochidium, Similar Species; Gene- ral Distribution; Alabama and Mobile Basin Distribution (a map appears at the end of each entry and is marked with black dot for each recorded occurrence); Ecology and Biol ogy; Current Conservation Status and Protection: Remarks; “ond Synonymy. The latter includes a caveat indicating that this is far from a chresony my, being limited to the first usage of a species-level epithet ( generic reas- signment not ecousider ed) considered in svnonymy. On the other hand, it is generously, almost exhaustively, ilhimi- nated with type figures, in color when available. The over 700- oda pages devoted to the treatment of two Unionoidean families, 43 genera, and 175 species- level taxa plus short vignettes on five species of hypo- thetical occurrence, six non-naiad clams (Sphaeriidae is not parsed) including the two non-natives Corbicula fluminea and Dreissena polymorpha, and finally a newly-diagnosed identity for the spuriously recorded (mmslonaline d) Unio decumbens I. Lea, 1861 [Ti “apeoi- deus exolescens Gould, 1843) of southeast Asia]. The bibliography contains over 1000 titles, and the index is inclusive with all topics, terms, person- and place-names, and genus-species, species-genus entries, and the same reciprocation for the common names. A stunning feature of this work is the photography of Richard Bryant, who captures the shells of each species- level taxon in large format, with crispness and color accu- racy. The specimens are almost all of the highest quality, sometimes apparently requiring the use of extralimital material. The shells are scrupulously posed with the ad- ductor scar axis horizontal, poste rior to the left (as was the custom of the prolific ables -publisher Isaac Lea). Such conventions make it easy on the diagnostic eye. The marshalling of information in the taxonomic por- tion, particularly in Ecology and Biology and in the temarks is staggering and probably indicates a strong collaboration among the authors of this work. Other features such as the thousands of locality indicators, the lifting of hundreds of type figures from classic works, give a dimension to this work that is unprecedented, especially informative, and indicative of a lot of hard There are taxonomic initiatives exercised in this work. A major one of these is dealing with the “Pleurobema problem.” Tabulations of pages 501-504 indicate the profusion of available names for Mobile Basin and other Alabama species and the synonymies of four prior monographers and in the present work, which has a relatively conservative perception of the diversity. Wil- liams et al. pare t he list of Turgeon, Quinn, et al. (1998) by seven species while adding three classic and one post- 1998 species. Likewise Hee Elliptio species. are resus- citated from synonymy as are a half dozen other species in five gener There are three un-named taxa included in the work, one sp., Epioblasma sp. cf. capsaefor- mis, and Toxolasma sp. Baty is, however, provided with vernacular names—a convenient machination. Although installed in the literature over the last decade, reassignments of long-recognized species to the resurrected Pleuronaia Frierson, 1927, and the newly- ordained Hamiota Roe and Hartfield, 2005, may sur- prise the reader. There are another half dozen generic reassignments necessitated by molecular genetic study, perhaps the most surprising ‘of which is the Pistolgrip, Quadrula verrucosa (placing Tritogonia in synonymy). Impl wusible as it may appear im context, there are two rather minor detractions which warrant brief mention. If I had my say in the creation of this magnum opus, T would have asked for a discussion of he geological history that provided the state with the colatoin of the Mobile Basin system, without which its present naiad diversity would have never reached the unassailable present- day mark. I see no reference to the Tertiary calamity that diverted the Tennessee River from its ancient course southwest past Lookout Mt. and into the heart of Alabama and thence to the Gulf of Mexico. The classic paper by Simpson (1900) on the evolution of the relevant naiad faunas and the geological evidence in support of it (Johnson, 1905a, “1905b: Adams, 1928) seem appropriate for the beginning of Chapter 3. The other little vexation is the persistence of gender-bending binomina in the naiad literature. It is not entirely clear how “Pleurobema stabilis” and “Ptychobranchus subten- tum” became entrenched, but the Code and the original descriptions indicate they should be rendered Pl. shabile and Pt. subtentus. Williams, Bogan, and Garner have produced a holistic and exhaustive work, carefully executed and seductively constructed. The taxonomic scope is unprecedented in recent years, covering some 60 percent of the American fauna. Aside from being a precious asset to the malaco- logical community, it will advance the understanding of hiodive srsity, ecology, and conservation in a much wider audience. To quote from Edward Osborne Wilson's Foreword: “People do care about species of wildlife, however, if they see a picture of it, know its name, and reac what is known of its distribution and natural his- tory. In addition to their contribution in mussel biology, this is what the authors have given us.” We applaud Williams, Bogan, and Garner, and we commend the Alabama Department of Conservation, Book Reviews, 2008 Page 263 Game and Fish Division as well as Auburn University, whose commitment has helped assure that this prodi- gious work will be affordable to a wider readership LITERATURE CITED Johnson, D. W. 1905a. Tertiary history of the Tennesseee River. Journal of Geology 13(3): 194-231, map. Johnson, D. W., 1905b. The Distribution of freshwater faunas as evidence of drainage modifications. Science 21: 588-592. Simpson, C.T. 1900. On the evidence of the Unionidae regard- ing the former courses of the Tennessee and other south- em rivers. Science 12(291): 133-136. Turgeon, D. D., J. F. rie Jr, A. E. Bogan, E. V. Coan, F. G. ecbes ’ G. Lyons, P. M. Mikkelsen, R. J Neves, C. F. E. we r, G. Rosenberg, B. Roth, A. Schel- tema, F. G. Thompson, M. Vecchione, and J. D. Williams. 1998. Common and scientific names of aquatic inverte- brates from the United States and Canada: mollusks, 2nd edition. American Fisheries Society, ee Publication 26, Bethesda, ix + pp. 1-509 + 16 pls. ( non-paginated ), Harry G. Lee 4132 Ortega Forest Drive Jacksonville, FL 32210 USA shells@hglee.com Guide to the Freshwater Molluscs of the Lesser Antilles Pointier. Jean-Pierre. 2008. Guide to the Freshwater Mol- luses of the Lesser Antilles. Conch Books, Hackenheim, Germany, 127 pp. www.conchbooks.de. Retail price: 35. This small volume covers the freshwater molluscan fau- na of the islands of the Lesser Antilles. These islands form a double are of volcanic islands extending from Anguilla in the north to Grenada in the south along the east edge of the Caribbean Sea. Dr. Philippe Jarne, of Montpellier, France wrote the short preface. A brief history of the study of freshwater mollusks in the Lesser Antilles is also provided. Pointier reports nine families of yale and two families of bivalves; 2S gastropod and three bivalve spe- cies from the islands. The gastropods are divided into 19 native gastropod species including the two endemic species, Neritilia succinea |Neritiliidae] and freshwater opisthobranch Tantulum elegans |Tantulidae], and nine introduced species. Lesser Antilles freshwater bivalve fauna contains the introduced Mytilopsis leucophaeta and two endemic species of Sphaeriidae. This volume has a flow chart key to the families of gastropods and bivalves and is suppleme nted with a page of figures illustrating the key landmarks, shell sculpture, and types of ope reula. Species are treated by family units in the same order as found in the key with discussion of the anatomy and ecology for the members found in the Lesser Antilles. Each of the gastropod species accounts is accompanied by two to five color illustrations of the shells, eggs, live animals, and ecology. The pulmonate species accounts incorporate dhistsations of the repro- ductive anatomy of each species. A section is devoted to aquatic habitats where these mollusks have been collect- ed and includes 44 color photogr aphs of habitats and The figure captions note some of the snail species found on the plants and some plants used as egg-laying sites or food. This volume is based on extensive field work throughout the Lesser Antilles. Species are well illustrated, including close-up color figures of live speci- mens of all but the oe opisthobr: anch Tantulum elegans. “Guide to the Freshwater Molluscs of the Lesser Antilles” is a great companion to the earlier work on the freshwater ficllaskes of Cuba (Pointier et al., 2005). T have found this volume very useful and recom- mend it for anyone working on, or interested in, iden- tification of the freshwater mollusks of the Lesser Antilles. This book will fit nicely in a backpack and will be a handy reference in the field. I would recommend this well illustrated book to anyone interested in fresh- water mollusks. LITERATURE CITED Pointier, J.-P, M. Yong and A. Gutiérrez. 2005. Guide to the freshwater molluscs of Cuba. Conch Books Hackenheim, Germany. 119 pages. Arthur E. Bogan North Carolina State Museum of Natural Sciences esearch Laboratory MSC 1626, Raleigh, NC 27699-1626 USA THE NAUTILUS 122(4):264, 2008 Page 264 Notice Bulletin of Zoological Nomenclature 65(2): 152. June 2008. OPINION 2197 (Case 3341) of the International Commission on Zoological Nomenclature. Cardium egmontianum Shuttleworth, 1856 (currently Trachycardium egmontianum; Mollusca, Bivalvia, CARDIITDAE): current usage conserved. Abstract. The Commission has ruled that the current usage of the specific name egmontianum for a common and widespread western Atlantic bivalve Trachycardium egmontianum (Shuttleworth, 1856) of the family CARDIIDAE is conserved by setting aside all lectotype designations for Cardium mindanense Reeve, 1844, prior to that by Vidal (1998). LITERATURE CITED Vidal, J., 1998. Taxonomic revision of the Indo-Pacific Vasticardium assimile species group (Mollusca, Cardiidae). Apex 13: 111-125. Errata A couple of items published in the most recent issue of The Nautilus (volume 122 issue number 3) need correction. In the article by Villalobos-Rojas et al. (2008), page 155, right-hand column, line 5, the first name of Ms. Kirstie Kaiser was misspelled. In the same article, the complete reference to Pitt and Kohl (1979) was omitted (see below). In the article by Gonzalez-Vallejo (2008), page 180, the image representing the female shell on Figure 3, right, was inverted by the authors; that shell is not left-handed. The editor apologizes to all parties concerned for these editorial blunders. LITERATURE CITED Gonzalez-Vallejo, N. E. 2008. Parasitism of Monogamus minibulla (Olsson and McGinty, 1958) (Gastropoda: Eulimidae) on the red sea-urchin Echinometra lucunter (Linnaeus, 1758) (Echinodermata: Echinometridae) on the Caribbean coast of Mexico. The Nautilus 122: 178-181. Pitt, W. and R. Kohl. 1979. A new Panamic Mitrella (Mollusca: Gastropoda). The Veliger 21: 467-468. Villalobos-Rojas, F., A. G. Guzman-Mora, Y. E. Camacho-Garcia. 2008. Catalogue of the type material deposited at the Zoology Museum, University of Costa Rica. The Nautilus 122: 155-165, NAUTILUS Volume 122 2008 AUTHOR INDEX ARRUDA, J. O. ......00 1 D4 MANN GAR) ssscdicescasteyes 217 Bocan, A. E. ... . 263 MEANS, MiG. TD coc eeccveda she coeieucntossndacavstinircaveadeipeawnmeadielstens 228 BBROUGHET: P. cccscssccessicsscuscsassscnsvasecanasessesvavsaabaisensesdeuvereeativssadvesens ] MEIER-BROOK, Cy cicéccsscsscoissssssiscecsosssccsvcsvesccsavsacesveccscessoecceine 298 GAETANO GC. THO Ss. ec cceadecsadeceveva voter sas adenesdicvsiottaaanciteuoipes baste 171 TUTTSEN Sc IM cascscsin caus anvanisniagsudsdec cic oncescdaeceawaumaecsunisuveenvusiccuaevesnacs 90] CAMACHO-GARCIA, Yo Ee cececceceeceeeceeteeeret ies eeretieeeeneeeieenees 155, ORDA, Be Ms vee fate vdenessszasvest heliiaes oasdiavsvietadee slawentavyneeeetes 143 CONCEPCION! Gs P.. sc: s.cissse tein Gnseterane nits nine tits 143 PASTORINO, G. ...cssssssssessesvesssesssssscsvssessessscssssscssevsecsssseess 107, 236 KOORNEDIL, Fe 6S, av sestengeatiny Poteauarsanes tun ceenaedeneauasuand aves dir teatas santas 143 PETIT, Re Eu ccccocccescsssessscacececevecececevevavevevadsessesisecevivsviveveveverseseees | Costa, Po M.S. vesseesee ces eeeiee teenies ieee eesieesiesniesneesecnsieaey 244 IPIMENTAS JA TDs. seitetosnssdoacertitecce aie aonacanvetssacgnten Saute seseen 244 COUTO, De Ry vessssssssessessssesceeeeeeeeceeeereeeceeeeeeeeececceceeceeeceeseennnty 244 PORTEEL, Ry! Wiles vratamiseccistisin encase cate artesh iets 79 CUMMINGS, Res Dis. sdeah ie sess dunbac ebagkauneesiebussapieguteiupsetataee eeheueas 259 SANTOS, A. Di. ccccccsccccccoscssccsscssccsscosecssssesssscestesucestseseectcesceeseees 143 CUNHA, (Gi Mie» sccissiccincitsicacissrcvvisauscevevecdetasaasiesatedecuestaoatecsbeurause 57 Ciccdieis Me ccueenn acter enim, tenth Ae ta c0e fate! ot othe od oa catch A 166 DuEnr, R. Se aaa ae ae ae 151 DAU: Ts We, 2cucseyelinevseten! devia csusgeh a attiea dusesaatieutiarcrensenieatese 115 EISPING;: SiS: avcetenectictesesarenyereee We aeceareaveadsdiesscpreneananeerens 143 Sc ; 3 : i SGARABINO,. Fy yiceccsasesccsissecqatssssscessiensenuotessagscascovinnesassccsensnhons 107 (GARMAT, MV,.