FHE NAUTILUS Volume 107, Number 1 March 24, 1993 ISSN 0028-1344 A quarterly devoted to malacology. APR 1 19S3 ^ p^^ ^^^ iX.:, ^ EDITOR-IN-CHIEF Dr. M. G. Harasewych Division of Mollusks National Museum of Natural History Smithsonian Institution Washington, DC 20560 ASSOCIATE EDITOR Dr. R. Tucker Abbott American Malacologists, Inc. P.O. Box 2255 Melbourne, FL 32902 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Robert T. Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr. Robert Hershler Division of Mollusks National Museum of Natural History Smithsonian Institution Washington. DC 20560 Dr. Richard S. Houbrick Division of Mollusks National Museum of Natural History Smithsonian Institution Washington, DC 20560 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Aurele La Rocque Department of Geology The Ohio State University Columbus, OH 43210 Dr. James H. McLean Department of Malacology Los Angeles County Museum ol Natural History 900 E.xposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Ms. Paula M. Mikkelsen Harbor Branch Oceanographic Institution, Inc. Ft. Pierce, FL 33450 Dr. Donald R. Moore Division of Marine Geology and Geophysics Rosenstiel School of Marine and Atmospheric Science University of Miami 4600 Rickenbacker Causeway Miami, FL 33149 Dr. Gustav Paula\ Marine Laborator\- University of Guam Mangilao', (^.uarn 96923 Mr. Richard L. IVtit P.O. Bo.\ 30 North Myrtle Beach, SC 29582 Dr. Edward J. Petuch Department of Geology Florida ,\tlantic University Boca Raton, FL 33431 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology University of California at Davis Davis, CA 95616 SUBSCRIPTION INFORMATION The subscription rate per volume is US $25.00 for individuals, US $40.00 for institutions. Postage outside the I nited States is an additional US $5.00 for surface and US $12.00 for air mail. All orders should be accompanied by payment and .sent to: THE N.4UTILUS, P.O. Box 7279, Silver Spring, MD 20907-7279, USA. Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028- 1344) is published quarterly by Trophon Corporation, 8911 Alton Parkway, Silver Spring. MD 20910. Second Class postage paid at Silver Spring, MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS P.O. Box 7279 Silver Spring, MD 20907-7279 TH Et^NAUTI LUS CONTENTS Volume 107, Number 1 March 24, 1993 ISSN 0028-1344 I IPD/\pv ' Limacosphaera, an I'nusual Mesogastropod (Lamellariidae) Larva of the Weddell Sea (Antarctica) , 1 )_ Hole, Mass. j The Reproductive Anatomy of Philomycus sellatits Hubricht, 1972 and Philomycus virginiciis Hubricht, 1953 (Gastropoda: Philomycidae) 9 Two Confusing Indo-Pacific Cerithiids 14 Rediscovery of Tiirbinella thersites Reeve, 1847, with Notes on its Taxonomic Position (Gastropoda: Fasciolariidae) 24 The Rediscovery, Morphology, and Identity of Conus emersoni Hanna, 1963 29 The Benthic Mollusk Faunas of Two Contrasting Reef Paleosubenvironments: Falmouth Formation (late Pleistocene, Last Interglacial), Jamaica 33 Klaus Bandel Stefan Hain Frank Riedel Henry Tieniann H. Lee Fairbanks Richard S. Houbrick R. N. Kilburn John K. Tucker James H. McLean Stephen K. Donovan D. T. J. Litllewood THE NAUTILUS 107(l):l-8, 1993 Page 1 Limacosphaera, an Unusual Mesogastropod (Lamellariidae) Larva of the Weddell Sea (Antarctica) Klaus Bandel Gediogiseli-Puliiontiilogisclies Institut und Museum Universit) of Hamburg BundesstraBe 55 2000 Hamburg 13 GERMANY Slefan Main A If red- Wegener- Institut fiir Polar- und Meeresforschung CoiumbusstraBe 2850 Bremerhaven GERMANY Frank Riedel Geologiseh-PalaoiUulogisches Institut und Museum University of Hamburg BundesstraBe 55 2000 Hamburg 13 GERMANY Henry Tiemann Zooiogisches Institut und Museum University of Hamburg Martin-Lutlier-King-Piatz 3 2000 Hamburg 13 GERMANY ABSTRACT Marsciiiopsis conica Smith. 1915 and M mollis (Smith. 1902) have pianktotrophic larvae that are unique among gastropods. They cover the larval shell with a lacunous muscular mantle that can change its volume by interaction of body fluid and muscle activity This limacosphaera larva is found in Antarctic waters and represents the most complex larval strategy within the "echinospira-group' . Notes on the biology and anatomy, including histology, of this larva are presented. Ketj ivords: Lamellariidae, Marseniopsis. larvae, Antarctica. introduc;tion The majority of benthic invertebrates of high-Antarctic seas brood their young or provide postspawning parental care rather than produce free-Uving larvae (Mileikovsky, 197L Picken, 1980). True pianktotrophic pelagic larvae are extremely rare in the 200 to 600 m deep high-Ant- arctic shelf areas. Plankton samples taken by the R/V polarstern in the eastern Weddell Sea during several late winter to late summer cruises yielded only two meropelagic larvae of benthic gastropods (Piatowski, 1987; Boysen-Ennen, 1987). Both of these larval types were regularly found. One of these is Capulus subcompressus Pelseneer, 1903, which had first been observed by Pelseneer (1903) and is described in detail by Bandel and Hain (in prepara- tion). The second lueropelagic larvae was first observed by Simroth (1914) in material from the Davis Sea. Sim- roth noted anatomical similarity to a planktonic gastro- pod from the deep sea areas of the Indian Ocean, which he described earlier as Limacosphaera macdonaldi (Sim- roth, 1908). This animal is a spherical, transparent, voluminous gastropod with two openings: one to allow head and foot to come out of the shell and the other situated on the opposite side of the sphere (Simroth, 1908, 1914). Simroth called the muscular inantle covering the larval shell the deutoconcha of the limacosphaera. The posterior pore, which he (1908) called the shell tunnel (Schalengang), connects the sea water with a cavity surrounding much of the thin organic shell. Simroth (1914) also suggested the presence of glandular cells and muscle fibers in the voluminous deutoconcha that surrounds the shell. He noticed that the Antarctic forms represented larvae rath- er than adult gastropods as he had originally assumed when describing Limacosphaera as a new genus of pe- lagic gastropods (Pteropoda). Simroth (1914) correctly placed these larvae into the ontogeny of members of the "echinospira-group" in the genus Marseniopsis. The term limacosphaera was retained, now to describe a very pe- culiar and characteristic larva. Simroth (1908, 1914) considered that the limaco- sphaera swam with the aid of their large velum. Another mode of sw imming, mainly by hydrostatic effects, was considered the likely mode of propulsion in the water column by Jevdonin and Minichev (1975). These authors observed developing spawn of Marsen- iopsis conica and foimd that the mantle of the embryo envelops the shell prior to hatching from an egg mass found in excavations of tunicate surfaces. Jevdonin and Minichev (1975) also noted that the mantle did not fuse completely but left an open shell pore. They seem not Page 2 THE NAUTILUS, Vol. 107, No. 1 Figures 1, 2. Adult animals ol I. Marseniopsis conica Smith, 1915, and 2. M. mollis (Smith, 1902). Dorsal views on left, showing mantles that are fused ahove the shell and cannot be retracted. Neutral \ lews on right, show foot, parts of the head \\\ illustrations drawn irom fi.\ed animals. Scale bar = 5 mm for figure I, 10 mm for figure 2. to have been aware of the earlier cie.scriptions of these larvae by Simroth (1908, 1914). Jevcloiiin aiul Miiiichev (1975) suggested that larvae with a size of about 20 mm in diameter were nearly ready to metamorpho.se. Their descriptions suggest that their observations had been car- ried out on preserved material and they most probably did not .see living larvae or embryos. We provide a new description of the histology of these larvae integrated with observations on the li\iiig forms and their metamorphosis. MATERIALS AND METHODS During the cruises polarsirkel 80/81, polarstern ant I, ANT III/3, ANT V/S and ant \'II/4 larvae of the li- macosphaera type were caught at about 100 stations in K. Bandel et al., 1993 Page 3 — grounding linie Icnntinenlal boundary) W •" E Figure 3. Map indicating location of the Wedell Sea and the area where the larvae of Marseniopsis were caught. neritic \\ aters above continental shelf areas of the eastern Weddell Sea (figure 3). Samples were taken with various plankton gears (mesh sizes of 0.335-4.5 mm) in upper water layers (300-0 m) and preserved in buffered for- malin. Six larvae were kept alive for six weeks during the expedition PS ant V/3. During this period they were fed two times with a diatom suspension. Only one of the larvae survived and metamorphosed four weeks after capture (November 1986). It was fixed in 70% ethanol. During the expedition PS ant VII/4 (January to March 1989) 42 living limacosphaera from RMT samples were photographed for size measurements and transfered in- dividually to small plastic aquaria (running water system, temperature adjusted to 0 °C). The larvae were fed once a week with a suspension of Antarctic diatoms. After 8 and 13 months the photographic measurements of larvae or juveniles were repeated. The larval shell, the thick, whitish to semitransparent tissue covering it (deutoconcha), and the juvenile shell were studied on 10 critical-point dried specimens using a SEM. In May 1989, six living larvae were fixed for histolog- ical studies in 4% seawater-formalin or 2.5% seawater- glutaraldehyde. One limacosphaera fixed in formalin was dehydrated using isopropanol. It was then treated with benzylbenzoate, embedded in paraplast, serially sec- tioned (7 ytm) and stained with Helioechtrubin BBL/ Acidgreen 5/Acidorange 10 after Halmi (Adam & Czi- hak, 1964). To detect lipids whole larvae without the deutoconcha as well as parts of the deutoconcha were treated using the oil-red-niethod (Romeis, 1968). These samples were embedded in glycerin and examined with a light mi- croscope. Another limacosphaera fixed in glutaraldehyde was osmicated in 1% osmium tetroxide, dehydrated in ace- Figure 4. Larva of Marseniopsis cf. mollis. Schematic drawing from several photr)graphs of living specimens. One quarter of the shell-covering tissue (deutoconcha) is removed, d, deuto- concha; e, eye; ed, epidermis; f, foot; s, shell; si, siphon; st, apical tube; t, tentacle; v, velum. tone and embedded in Spurr's fluid. Sections were cut at ca. 80 nm to 1 ^m using an ultramicrotome (Reichert). The 80 nm sections were stained with uranyl acetate and lead citrate, then examined and photographed using a TEM (Zeiss). The 1 ^m sections were stained with To- luidine Blue and Pyronin and examined with a light microscope. Voucher material is deposited at the Zoological Insti- tute and Museum (L'niversity of Hamburg) (cat. no. M 885). RESULTS The Living Larva Description: The shape of the limacosphaera (figures 4-6) is globular with an overall diameter ranging from 1.8 to 20 mm. The actual larval shell is covered by the deutoconcha. Some of the larvae have granular deposits in the outer layer of the deutoconcha. On the ventral side of the larva (figure 6) there is a slit-like opening for the foot and the four large velar lobes. This opening is extended anteriorly by a siphon-like, semicircular groove. A second, small, tunnel-like excavation extends from the apical region of the shell to the surface of the deutocon- cha. Observations on living larvae sorted from plankton samples show that both openings can be closed and that the deutoconcha produces much hyaline mucus. During metamorphosis the shell is still barely calcified (figure 7), becoming solidly calcified right after meta- morphosis (figure 8). The larval shell reaches 2.3 whorls with fine spiral threads on the first 1.5 whorls. Earliest growth lines are visible after 0.8 whorls (figure 9 arrow). In well fed larvae, the greenish visceral mass can be seen through the shell and the deutoconcha. All studied larvae had a well developed foot. The eyes are at the base of the tentacles. In contrast to Simroth's original description of Limacosphaera macdonaldi from Page 4 THE NAUTILUS, Vol. 107, No. 1 Figures 5, 6. Living larvae ul A/(j),Miii(>/;.si.v cj. mollis. 5. Dorsal view. 6. Ventral view, with foot, velum, tentacles and eyes visible. Scale bars = 5 mm for liolh pliotograjihs the tropics, the operculum is absent in the Antarctic specimens. Biology Deposition of egg capsules in the tests of compound as- cidians is known for temperate lamellarian species (Fret- ter & Graham, 1962). Antarctic species have the same spawning habit (Jevdonin & Minichev, 1975; P. Dayton, personal communication). One compound ascidian with appro.xiinately 20 egg capsules was obtained in February 1989. Although maintained in an aquarium, the eggs failed to develop. Figures 7-9. Shell of Marseniopsis cj. mollis. 7. During meta- morphosis, the shell is barely calcified and therefore, flexible. 8. .Alter metamorphosis, shell solidly calcified and lias reached 2 4 whorls. 9. Same specimen as in fig, 8. P'irst growth lines (arrow) are visible at 0 7 whorls of the shell The sjjiral threads end at 1.4 whorls. Scale bars = 1 mm in all photographs. Larval size, as well as the time and location of sam- pling, indicates that hatching of larvae occurs from late winter to austral summer in the shelf areas of the eastern Weddell Sea. The smallest larvae found have a diameter of 1.8 mm. We suggest that this size is reached shortK after the larvae have hatched and risen from the l)ottoni. Aquar- ium observations indicate that the larvae rise by buoy- ancy. The large velar lobes are not used or have only secondary function. The buoyancy control mechanism is uiikuow n. In the northern shell areas of the Weddell Sea the amount of food (0.07 ixg chlorophyll -a/liter) is more K. Bandel et al., 1993 Page 5 Figure 10. ScluMiuitir draw iiig ot the ajMciil lulx' (shell tuiiiu-lj of Marseniopsis cf. mollis, c, cilia; cc, collagenous cell; cf, collagen fibers; gc, gland cell; mf, muscular fibers; mv, micro- villi; n, nuclei; s, shell; st, apical tube limited even during phytoplankton blooms. In open wa- ters or polynyas (ice free areas), the phytoplankton con- centration barely reaches 1.5 ng chi a/liter. It is sus- pected that growth of larvae in these areas takes a much longer time. The total Hpid content of larvae caught in February (end of austral summer) is low (3.5% dry weight) in comparison to other planktonic organisms. Seventy per- cent of these lipids are triacylglycerols (Hagen, 1988). Although fed only twice, one limacosphaera caught in October 1986 metamorphosed four weeks later. During PS Ant VII/4 small larvae with a diameter of 3.9 mm began to metamorphose in the aquaria after 10 to 24 hours, while specimens of 10 mm diameter remained in the larval stage for up to 8 weeks. One specimen caught in the end of February 1989 was still in the larval stage when it died in August 1990. Predatory pressure on the larvae seems to be very low. The shell-covering tissue of juveniles and adults is known to produce acidic secretions. HISTOLOGY Observations with Light Microscope The deutoconcha of the limacosphaera consists of tissue that is composed of an outer epidermis, a central, cav- ernous connective tissue, and an inner epidermis cov- ering the shell. The outer epidermis is composed of a single layer of cells with flattened nuclei and immersed, large, light, glandular cells with a single nucleus. Below it is a vo- luminous, spacious connective tissue, consisting of col- lagenous and muscular fibers with few cells suspended in it. The inner epidermis is a single layer of cells sup- ported by connective tissue and muscle fibers. These cells are stretched in length, and their nuclei are even more flattened than those of the outer epidermis. The deutoconcha of the limacosphaera is connected to the inner mantle of the apertural region of the shell by an especially strong muscular bridge of tissue. The outer mantle rests on the shell near the aperture and forms the connection to the inner mantle covering the visceral mass and the pallial cavity. Outer and inner mantle are "divided" from each other by the periostraca! gland zone characterized by the presence of many nuclei. On the apical side of the deutoconcha the e.xternal mantle is pierced by a tube-like canal (figure 10, st) connecting sea water and shell surface. This canal is lined with a rugged, ciliated epithelium (figure 10, c) that continues along the inner mantle for about 0.4 mm, to form a small cavity between shell and mantle (figure 10). The tissue below the ciliated epithelium of the canal contains an agglomeration of large light epithelial glan- dular cells. The iimer side of the deutoconcha connects to the organic shell that is composed of a double-layered peri- ostracum. This shell in its apical portion is filled with the visceral mass containing the large digestive gland. Its diverticula consist of large endodermal cells with basal nuclei and small cavities. The apical portion of the di- gestive gland is glandular and contains three different types of stored substances. The most basal portion con- tains proteins, somewhat above lie a large number of spherical bodies. Scale-like storage material that is weak- ly aeidophilous is suspended between these. The spherical bodies (6-19 ^m in diameter) are lipids (analysed by oil-red method). The spheres of lipids were arranged like strings of pearls of different length throughout the entire digestive gland of a larva caught in February 1989 during the phytoplankton bloom. The outer surface of the deutoconcha was also tested for the presence of lipids, but was found to lack them. This test was repeated on another larva that had been caught prior to the beginning of the phytoplankton bloom (mid-November 1986). It lacked lipids in either the man- tle or the digestive gland. Of the other organs of the larva, the nerve ring proved to be quite large. The eyes were well-developed, with lens-like light collectors. The foot is intensely ciliated, almost to the same degree as the large velum. The radula is well developed and functional. Observations with Electron Microscope The epidermis of the deutoconcha (figure 11) is covered by a 2 ^m high rim of microvilli. These microvilli are differentiated into a narrow basal portion with many filaments and a light, partly bubble-like expanded upper portion. These "bubbles" are secretory vesicles, usually linearly arranged and attaining 2-4 times the width of the mi- crovilli. They are produced by glandular cells at the base of the rim of microvilli and are secreted at the outside of the rim. "Bubbles" open up at the outside of the rim as could be seen in some sections. The surface of the rim of microvilli is covered with fine fibrillar material containing small dark granules. Page 6 THE NAUTILUS, Vol. 107, No. 1 Between microvilli, larger vesicles with a small internal membrane are often present. Some larger vesicles are as high as the microvilli and are connected to the glandular vesicles of the epidermis. The epithelium of the deutoconcha consists of a single layer of flattened cells resting on a distinct basal mem- brane. The cells are up to 10 ^m in size, and are con- nected to each other by apical contact zones. Cell bound- aries are strongly folded and intercalated. Some nuclei are strongly heterochromatic, others are large, less densely packed and of irregular shape. All cells hold large, light vesicles often containing membranous extensions. These vesicles form a loosely connected sys- tem that is in contact with the vesicles extruded into the rim of microvilli. Newly extruded vesicles hold dense granules. A voluminous connective tissue is present below the epidermis and fills the space between inner and outer layer of the deutoconcha. Within a homogeneous basal substance, individual, transversally striped fibers of col- lagen connect the basal membrane of cells within the tissue to the epidermis. Large cavities lie within the connective tissue near the epidermis. These cavities are lined with a thin, conspic- uously foamy cellular layer resting on a basal membrane. The nuclei are flattened and elongated. Nerves, consisting of several axones within a glial cell, and smooth muscular fibers with connections to the col- lagen fibers are distributed within the connective tissue. The intermediate layer between outer and inner ep- ithelia (gelatinous layer, Simroth, 1908) is shown to con- tain a network of collagenous fibers connected to muscle and epithelial cells. This layer functions together with interconnected blood lacunae as a hydroskeleton that can change the outline, shape and width of the limacos- phaera. The apical tube (figures 4, 10, st) of the deutoconcha is ciliated, in contrast to the remaining surface of the mantle. This ciliation continues into the small cavity at the proximal end of tube above the shell. This cavity is less extensive than assumed by Simroth (1908) and does not surround the entire shell surface. Simroth (1908) noted a folded surface of the shell, which is an artifact of preservation. A rim of microvilli on a unilayered epithelium sur- rounding a structured connective tissue represents a large surface for resorption as well as for secretion by many glandular cells that come together in canals opening into pores. The type of secretion produced by these glands is unknown. The secretion expelled between the microvilli probably represent neutral mucus material. DISCUSSION The larvae metamorphosed into juveniles that had either a smooth surface or a tuberculated body surface. Both juvenile forms were analysed by gel-elect rophoresis (ID- UISK-SI)S-PA(;E) at the Alfred-Wegener-in.stitut (Dr. T. Stadler). The results were compared to gel-electro- Bum cb Figure 11. Transmission Electron Micrograph of the deuto- concha epidernii.s of Marscniopsis cf. mollis, bm, ba.sal mem- brane; ci), cell boundary; ct, connective tissue; mv, microvilli; n, nucleus. phoresis (same method) analyses of adult Marseniopsis conica and Marseniopsis mollis. Differences in the data indicate that the predominantly smooth juveniles re- present M. mollis and the tuberculated ones M. conica. I'hus it can be concluded that both species have very similar larvae. It was not possible to section and electro- phoretically analyse the same individual. The course of a limacosphaera's development depends on a variety of factors, including currents in the upper water layers, melting of sea ice, and patchyness of phy- toplankton production. Larval survival in captivity for 1.5 years indicates that a limacosphaera could remain in the plankton for over a year if conditions were not fa- vorable for metamorphosis. Analyses of lipid content suggest that larvae either use most food-energy for growth or they store energy in form of other biochemical metabolites {e.g., proteins). The vertical distribution and the stomach contents of larvae caught during mid-October at 72°S indicate that they feed under the sea ice where the phytoplankton concen- tration (0.07 ng chl- a/liter) was seven times larger than in deeper water (Scharek, personal communication). Due to the southwcsterK water currents near the surface, the majority of larvae will be transported to southern parts of the study area, where very high phytoplankton con- centrations (50-150 ^g chla/liter) were measured di- rectly under the sea ice in October 1989. By utilization of this enormous food resource, larvae living in that area could grow very rapidly. Due to the fact that the macro- zoobenthos community of the southern Weddell Sea is totally different from that of the eastern Weddell Sea (Voss, 1988) and therefore not suitable for survival of beiithic stages of Marseniopsis, there nuist be a way of K. Bandel et al., 1993 Page 7 returning pelagic stages (metamorphosing larvae, juve- niles or adults) back to the hatching grounds. This may be accomplished by counter-currents nuining near the bottom. Piatkowski (1987) and Boysen-Ennen (1987) reported the absence of larvae in the oceanic domain between the tip of tlie Antarctic peninsula and the north-eastern part of the Weddell Sea. Marseniopsis populations of the peninsula are likely zoogeographically separated from populations in the eastern Weddell Sea. Nevertheless some of the larvae could be transported by eastward wind drift ot surface water layers near the ice edge from eastern-Antarctica to the Weddell Sea. Between Feb- ruary and March 1983, the mean abundance of larvae in the eastern Weddell Sea was 24 specimens per 1,000 m' (Boysen-Ennen, 1987), with maximum densities of 65 larvae per 1,000 m' in the southern part of the study area (Piatkowski, 1987). Most larvae were found in water layers of 200 to 50 m (63%) and 50 to 0 m (35%), only 2% were reported from 300 to 200 m depth (Boysen- Ennen, 1987). The major function of the deutoconcha is that of a buoyancy organ. Altering the volume of the outer mantle may vary the speed of sinking of the li- macosphaera. According to Stokes formula (Tiemann & Betz, 1979) this speed depends on the specific weight, which again is a function of the diameter of the lima- cosphaera. Histological analyses of the limacosphaera showed that the deutoconcha contains very little organic material and is thus very close to the density of sea water. The deutoconcha can enlarge the diameter and thus the volume of the larva without decreasing its weight in the water. A general calculation indicated that the volumi- nous outer mantle of the limacosphaera decreases the speed of its sinking to about one half of what it would be without such a cover around the shell The ability to float was observed in aquaria for all developmental stages. Even large adult specimens could float near the bottom after disturbance (diving obser- vations at the Antarctic peninsula, W. Wagele, personal communication). The function of the deutoconcha's apical tube and cavity is still unknown. It is evident that water can be pumped into the cavity and expelled out by ciliary cur- rents as well as by muscular movements of the deuto- concha. It is also evident that glands can secrete sub- stances into the lumen of tube and cavity. Histological examination of the limacosphaera for the most part confirms the observations of Simroth (1908, 1914). His assumption that muscular cells are present within the deutoconcha is substantiated. The prolific mucus secretion of the entire deutoconcha, as well as its large size may serve as defensive mechanisms against carnivorous planktonic groups like copepods or krill. The shell size of hatching larvae can only be inter- preted from the embryonic and the larval shell (figure 8). There are two possibilities. The normal case would be that the embryo hatches shortly after the Ijeginning of growth lines (the mantle becomes free from the shell) (figure 9, arrow). This happens after 0.8 whorls and a diameter of the shell of 0.65 mm. The second possibility is that the embryo does not hatch before reaching 1.4 whorls (with spiral threads). The shell then has a diameter of 1.5 mm. To reach this size the embryo v\ould have to feed on extra yolk. It is proposed that the embryo hatches when the shell has reached 1.4 whorls. At this time tlie spiral threads disappear. This is probably the consequence of the man- tle fusing above the shell. There is no difference in sculp- ture between the subsequent part of the larval shell and the teleoconch of the investigated species of Marseniop- sis. ACKNOWLEDGEMENTS Tliis study was carried out with financial support of the Deutsche Forschungsgemeinschaft (DFG) in the frame of the Antarctis-research grant and science grant Ba 675/ 6-1 We have been greatly aided by our colleagues of the Alfred-Wegener-Institut (Brenierhaven), Zoological and Geological-Paleontological departments of the Uni- versity of Hamburg. We are grateful to Dr. T. Stadler who carried out the gel-electrophoresis analysis. Dr. M. G. Harasewych greatly improved st\le and grammar of the manuscript. To all persons and institutions involved, we express our sincere thanks. LITERATURE CITED Adam, H. and G, C^lzihak 1964 Arbeilsniethoden der mak- roskopischen Anatomie. Ein Laboratoriumshandbuch fiir Biologen, Mediziner und technische Hilfskrafte. Gustav Fischer Verlag, Stuttgart, 583 p. Boysen-Ennen, E 1987 Zur Verbreitung des Meso- und Mak- rozooplankton.s im Oberflachenwasser der Weddell See (Anlarktis) Berichte zur Polarforschung 35:1-126 Fretter, V. and A, Graham. 1962. British prosobranch mol- luscs. Ray Society, London. 7.55 p. Hagen, W. 1988. Zur Bedeutungder Lipide im antarktischen Zooplanktnn. Berichte zur Polarforschung 49:1-129 Jevdonin, L. A. and J. S. Minichev. 1975. Adaptations of pelagic gastropods. Malacological Review 11:75 Mileikovsky, S. A. 1971. Types of larval development in ma- rine bottom invertebrates, their distribution and ecological significance: a re-evahiation. Marine Biology 10:193-213. Pelseneer, P 1903 Resultats du vo\age du S Y. "Belgica" 1897-1899. Rapports Sci. Zoologie, Mollusques (Amphi- neures, Gasteropodes et Lamellibranches) Anvers, 85 p. I'lalkovvski, L'. 1987. Zoogeographische Untersuchungen und Gemeinschaftsanalysen an antarktischem Makroplankton. Berichte zur Polarforschung 34:1-138. Picken, (; B 1980. Reproductive adaptations of Antarctic benthic invertebrates. Biological Journal of the Linnean Society 1467-75. Ronieis. B. 1968. Mikroskopische Technik. Oldenbourg Ver- lag, Mijnchen, 16. .\uOage, 7.57 p. Simroth, H. 1908. Gastropodenlaiche \ind Gastropodenlarven der Deutschen Tiefsee-Expedition In: C Chun (ed). Wis- senschaftl Ergebnisse der Deutschen Tiefsee-Expedition Page 8 THE NAUTILUS, Vol. 107, No. 1 auf dem Dampfer -Valdivia' 1898-1899, Band IX:365- Tiemann, H and K -H Bctz 1979 Elutriation: theoretical ,,Q considerations and methodological improvements Marine Simroth H 1914. Pelagische Gastropodenlarven der deut- ecology. Progress Series 1277-281. schen Siidsee-Expedition 1901-1903. /;i: Drygalski, E.v Voss, J. 1988 Zoogeographie and Gemeinschaftsanalyse des (Hrsg.). Deutsche Sudpolar-Expedition, Zoologie Band VII Makrozoobenthos des Weddellmeeres (Antarktisj. Berich- 15143-160. ''' ^"'' Polarforschung 45:1-14.5. THE NAUTILUS 107 1 9-13. 1993 Page 9 The Reproclucti\ e Anatom\ of Philomijcus sellatus Hubricht. 1972 and Philomijcus virginicus Hubricht. 1953 (Gastropoda: Philom\cidae) H. Lee Fairbanks Penns\l\ ania State Universits- Monaca. PA 15061 USA ABSTRACT Specimens of Philomycus sellatus and P virginicus were col- lected at or near their t> pe localities. Their reproducti\ e anat- om\ is described and compared with that of other species of Philomycus. The penial anatom\ of P sellatus is unique in ha\ing two constrictions in the penial wall; P virginicus is distinguished by a circular pustulose ridge that separates the distal penis from its atrial opening. Key Words: SloUusca; Gastrojxxla; landsnail: slug; Philomy- cldap: Philornvcj'- reproductive anatomy LVTRODUCTION .\1] species of the terrestrial slug family Philomycidae ha\e a mantle that co\ers the entire bod> of the slug. Three genera of this family. Philomycus. Pallifera. and Megapallifera. occur in the United States, with most species being limited to the eastern and south-central areas of the country . Few original descriptions pro\ide anatomical data. Neither the description of Philomycus virginicus Hubricht. 195.3 nor of P. sellatus Hubricht, 1972 discusses or figures an> part of the reproductive anatomy. Based up>on the mande pattern. Hubricht 19721 compared Philomycus sellatus to P ccirolinianus Bosc. 1S02!. but made no comparisons between P. virginicus and other species of Philomycus. References to the re- producti\e anatomy of P. virginicus are limited to two articles by Branson il96S. 1969; that compared P. vir- ginicus with P. bisdosus Branson. 196S. Neither contain figures or measurements. The reproductive anatomy of P. sellatus has not pre\iousl\- been studied. The goals of the present study were to describe the anatomy of the reproductive systems of Philomycus vir- ginicus and P. sellatus. and to compare them with those of other sp>ecies of Philomycus. METHODS AND MATERI.ALS Sf)ecimens of Philomycus virginicus were collected on 19 May 19S9 near nulepost 47 north of Skyline Drive. Shenandoah National Park, Madison County , \irginia. under the loose bark of logs and dead trees, eleiation appro.ximately 1110 meters. This station was approxi- mately O.S kilometers east of the tvpe locafity for this species. Specimens of P. sellatus were collected on 17 May 19S9 from its t>p)e locahty (2.7 kilometers northeast of Princeton, along Highwa> 65. Jackson County . .Ala- bama; in beede galleries in rotten logs, elevation ap>- pro.ximateK 215 meters. For comjjarative purposes, the following sp>ecie5 were also e.xamined: P. carolinianus collected in Berkele> Cotmty . South Carolina approxi- mateK 2-5 kilometers from Charleston, the designated t>pe iocaht) ,Pilsbry, 1945:754; on 22 June 19SS; P. bisdosus collected from Breaks Interstate Park. Dick- enson Coimty. Virginia the type locahty on 22 May 1987: P. flexuolaris collected from Breaks Interstate Park, Dickenson County. Virginia on 22 Ma> 1987: and P. togatus collected near Riceville. PittsyKania County, Virginia on 20 May 1987. External characteristics of the specimens were com- pared with the original sp)ecies descriptions to ensure correct identification. All specimens were drowned in distdled water and dissected immediately. The repro- ducti\ e sy stems were removed and the atrium and penis opened to examine their internal anatom\ . Subsequendy all material was preserved in 70"^ ethanol. DrowTiings and dissections were conducted during the month of Jvme in each > ear to reduce any differences that may be at- tributable to the stage of the life c>cle. The sjjecimens were maintained in terraria for approximately two weeks prior to dissection. E^ch complete reproductive system was spread out in a f)etri dish and then projected- \ia an overhead projector, and traced. The p>enial figures are tracings of photo- graphic shdes of the opened organs. Voucher sp>ecimens have been deposited in the Academ\- of Natural Sciences, Philadelphia ANSP .Al>326 for Philomycus bisdosus, .ANSP A1S327 for P virginicus. ANSP A13328 for P. sellatus, and .ANSP Al-3329 for P. carolinianus}. RESULTS Four adult specimens each of Philomycus sellatus and of P. virginicus were collected and used in the studv. Page 10 THE NAUTILUS, Vol. 107, No 1 / Figures 1-4. Mantle patterns of living slugs. 1, 2. Philomtjcus virginicus 3, 4. Philomtjcus sellatiis. Scale bar = 20 mm. External characteristics of all four specimens of each species (figures 1-4) agreed with their type descriptions. The gross reproductive anatomy was similar for all spec- imens of a species. Measurements of selected reproduc- tive organs are shown in Table 1. Genitalia of Phibmycus sellatus Hubricht, 1972 (figures 6, 8) Atrium glandular on distal half of outer surface, length approximately 60'/o penial length. Vagina length ap- pro.ximately 10% of penis. Spermathecal duct approxi- mately same diameter as free oviduct, length nearly twice that of free oviduct, slight taper toward spermatheca. Spermatheca round. Dart sac (with dart) larger than spermatheca. Penis diameter at proximal end nearly equal to atrial diameter, gradual taper to diameter of vas de- ferens at distal end; penial sheath covers approximateK Figures 5, 6. Genitalia, 5. Philornycus virginicus. 6. Philo- mtjcus sellatus. Scale bar = 10 mm. A: atrium, AG; albumen gland, AR: accessory retractor, DS: dart sac, FO: free oviduct, G: gonad, GP: genital pore, HD; hermaphroditic duct, P: penis, PR: penial retractor, S: spermatheca, \': vagina, \'D: vas de- ferens. 90% of penis. InternalK , distal third of penis with several thin low folds, separated from remainder of penis by constriction in penial wall; middle third with 3-4 large pustulose ridges; proximal third with several thin non- pustulose folds, separated from middle third b\ constric- tion in penial wall. Vas deferens enters distal end of penis, length approximately four times that of penis; diameter of middle third half that of ends. Penial retractor muscle maximum width 2-3 times diameter of distal end of penis, length 40% of penis. Accessory retractor muscle present, located on outer wall of atrium near its junction with penis. Genitalia of Philornijcus virginicm Hubricht, 1953 (figures 5, 7) Atrium length approximately equal to that of penis, ex- ternal surface of distal half glandular. Base of penis bulg- es into atrium. N'agina short, approximateK 15% of penial length. Spermathecal duct diameter greater than that of free oviduct, slight taper toward spermatheca. Sperma- theca round. Dart sac (with dart) smaller than sperma- theca. Penis straight, basal diameter one third of length, slight taper to junction with vas deferens; penial sheath reaches to junction of penis with vas deferens. Internally, H. L. Fairbanks, 1993 Page 11 Figures 7, 8. Internal penial anatomy. 7. Philomycus virgin- iciis. 8. Philomycus setlatus. Scale bar = 10 mm. Cut surfaces indicated by oblique lines in all drawings. A; atrium, DS: dart sac, P: penis, PR: penial retractor, PS; penial sheath, VD: vas deferens. penis with 4-6 pustulose ridges, a basal pustulose circular ridge sets off distal parts of the penis from its opening into the atrium. Vas deferens encircles distal end of penis, entering penis terminally. Vas deferens length approxi- Figures 9, 10. Genitalia 9. Philomycus bisdosus. Scale bar = 10 mm. 10. Philomycus carolinianus. Scale bar = 5 mm. A: atrium, AG: albumen gland, .\R accessory retractor, DS: dart sac, FO: free oviduct, G: gonad, GP: genital pore, HD: her- maphroditic duct, P: penis, PR: penial retractor, S: spermathe- ca, V: vagina, YD: vas deferens. mate!) 3.3 times that of penis, approximately same di- ameter throughout its length. Penial retractor muscle maximum width one third that of diameter of distal end of penis but broader at its distal end, length approxi- mately one third that of penis. Accessory retractor muscle absent or not distinguishable from typical connective tissues. Table I. Measurements (mm) of selected reproductive organs of species of Philomycus. Means with std. deviation and ranges are P. sella t us (4)* P. virginicus (4)* carolinianus (5)* P. bisdosus (2)* flexuolaris (2)* P. togatus (2)* Length of penis 13.1(1.82) 11.0-14.2 5.8 (0.66) 5.4-6.8 6.0(0.61) 5.4-7.0 7,1 (0,00) 7.1 7.3 (0.35) 7.0-7.5 12.9 (2.33) 11.2-14.5 Diameter of penis (atrial end) Length of vagina 4.4(0.49) 4.1-5.0 3.7(1.07) 2.7-5.0 2.2(0.29) 1.9-2.7 3.0 (0.07) 2.9-3.0 2.8(0.64) 2.3-3.2 3.3 (0.42) 3.0-3.6 1.2(0.15) 1.0-1.3 0,8 (0.29) 0.5-10 1,5(0,41) 1.1-2.2 1.1 (0.49) 0.7-1.4 1.6(0.35) 1.3-1.8 3.1(1.70) 1.9-4.3 Length of free oviduct 8.8(1.50) 7.8-10.5 10.2(1.87) 8.9-12.9 7.7(1.44) 6.2-10.0 7.6(0.62) 6.9-8.1 7.6 (0.64) 7,1-8.0 7.2(0.28) 7.0-7.4 Length of \as tleferens 57.5(4.77) 52.0-60.5 21.1 (2.11) 1 8 8-23 0 38.1 (4.96) 33,8-45,1 19.1 (1.02) 17.9-19.8 24.2 (3.04) 22 0-26 3 34.0(0.71) 335-345 * Number ot specimens measured. Page U THE NAUTILUS, Vol. 107, No. 1 Figures 1 1, 12. Internal penial anatomy. 1 1. Philomyctis bis- dosus. Scale bar = 10 mm. 12. Philomycus caroliniantis. Scale bar = 5 mm. A: atrium, DS: dart sac, P: penis, PR: penial retractor, PS: penial sheath, VD: vas deferens. DISCUSSION When Hubricht (1972) described Philomycus sellatus, he noted that, based upon the posterior half of the mantle, it ". . .is most closely related to P. carolinianus (Bosc).", with which "It sometimes occurs. . .". However, the black transverse band near the anterior end of the mantle of F. sellatus (figures 3, 4), which is not present on P. carolinianus (Pilsbry, 1948 p. 748), easily separates the two species in the field. Their reproductive systems (fig- ures 6, 10) are also distinct. The shape of the penes and their internal anatomies (figures 8, 12) are clearly dif- ferent, Philomycus carolinianus lacks the two internal constrictions in the penial wall that are found in P. sel- latus, and the internal penial pilasters are smaller and more numerous in P. carolinianus than in P. sellatus. Additionally, the penis and vas deferens of P. sellatus are nearly twice as long as those of P. carolinianus (Table 1). Indeed, the black anterior transverse band on the mantle and the two constrictions in the penial wall are characteristics unique to P. sellatus. Hubricht (1953) noted that Philomycus virginicus was found associated with P. flexuolaris (Rafinesque, 1820) and P. togatus (Gould, 1841) (as P. carolinianus collinus in Hubricht's 1953 article). Among these ta.\a, only P. virginicus has the transverse chevron pattern on the man- tle (figures 1, 2) and is thus easily separated from the others in the field. In addition, measurements of the reproductive systems of P. flexuolaris and P. togatus demonstrated several differences from that of P. virgin- icus (Table 1). The penis and vagina of P. virginicus are shorter than those of P. flexuolaris; the free oviduct of P. virginicus is much longer than that of P. flexuolaris. Philomycus togatus has a much longer penis, vagina, and vas deferens than P. virginicus; the free oviduct of P. virginicus is longer than that of P. togatus (Table 1). Branson (1968), in reference to P. virginicus and P. bisdosus (figures 5, 9), stated that "The genitalia of the two species also differ.", but no figures nor measurements were included. Branson (1969) listed some differences between the reproductive anatomies of these latter two species, but again without figures or measurements. Fig- ures 7 and 11 show clearly the differences in the shape of the penis and the internal penial anatomy of these two species, and the free oviduct of P. virginicus is nearly 1.5 times the length of the P. bisdosus oviduct (Table 1 ). In addition, P. bisdosus lacks the anterior to posterior transverse chevrons on the mantle. Comparisons with previous studies of reproductive anatomv in species of Philomycus (Fairbanks, 1986, 1989; Pilsbry, 1948) con- firmed that the internal surface of the penis of P. vir- ginicus is unique in having a basal circular pustulose ridge at the atrial end. ACKNOWLEDGMENTS Financial support for the field trips associated \\ ith this study was provided by grants from the Research De- velopment Grant Fund of The Pennsylvania State Uni- versity. My thanks go to three anonymous reviewers who provided valuable suggestions for improving this article. LITERATURE CITED Bosc, L. A. G. 1802. Histoire naturelle des coquilles. Conten- ant leur description, et leurs moeurs. \'ol. I. Paris, 343 p., Ipl. Branson. B. A. 1968. Two new slugs (Pulmonata: Philomy- cidae: Philomycus) from Kentuck\ and X'irginia. The Nautilus 81(4):127-133. Branson, B. A 1969. Genital differences in Philomycus vir- ginicus Hubricht and P bisdosus Branson. The Nautilus 82:74, Fairbanks, H. L. 1986. The taxonomic status of Philomycus togatus (Pulmonata: Philomycidae): a morphological and electrophoretic comparison with Philomycus carolini- anus. Malacologia 27(2):271-280, Fairbanks, H, L. 1989, The reproducti\e anatoms and tax- onomic status of Philomycus vcniislus Hubricht, 1953 and Philomycus bisdosus Branson, 1968 (Pulmonata: Philo- mycidae), The Nautilus 103(I):20-23, Gould, A, A, 1841. Report on the invertebrates of Massachu- setts, comprising the MoUusca, Crustacea, Annelida and Radiata, vol. 3. H. L. Fairbanks, 1993 Page 13 Hubricht, L. 1953^ Three new species of Philomycidae^ The Nautilus 66(3):78-80. Hubricht, L. 1972, Two new North American Puimonata: Paravitrea seradens and Philomycus sellatus^ The Nau- tilus 86(1):16-17. Pilsbry, H. A. 1948. Land MoUusca of North America (North of Mexico). The Academy of Natural Sciences of Phila- delphia Monograph Number 3, Vol, II Part 2:759. Rafinesque, C. S. 1820. .Annals of Nature or annual synopsis of new genera and species of animals and plants discovered in North America, p. 10. THE NAUTILUS 107(l):14-23, 1993 Page 14 Two Confusing Indo-Pacific Cerithiids Richard S. Houbrick Department of linertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC. 20560 ABSTRACT Cerithium zebrum Kiener, 1841 and Cerithium boeticum Pease, 1860. although originally described as CerUhiiim species, had been allocated to Bittium Gray, 1847, in the recent literature Anatomical investigation has shown that these two species do not have the characters of Bitlium species, but are are more like Cerithium taxa, e.xcept for their small size. Cerithium zebrum is distinguished from Cerithium boeticum. based on morphological characters derived from the shell, radula and soft anatomy. Synonymies, descriptions and a discussion of the differences between the species are presented. Key Words: Cerithium, Bittium, morphology, anatomy, tax- onomy. INTRODUCTION Small-shelled cerithiids are difficult to identify to the species-level, particularly poorly-known taxa from the Indo-Pacific region. Generic and sometimes familial de- terminations of these small snails are likewise difficult and controversal, and in most museum collections many small-sized cerithiids are grouped together and loosely attributed to the genus Bittium Gray, 1847. During a recent generic review of the Bittium-gToup the anatomy of a number of "Bittium" species were examined. One of the taxa studied is "Bittium" zebrum (Kiener, 1841), a common species throughout the Indo- Pacific region that is extremely variably in sculpture and color pattern. As expected in a common, widely-distrib- uted species comprising many sculptural phenotypes and color morphs," Bittium ' zebrum has been zealously ov- ernamed by authors, as a perusal of its synonymy will testify. When living "Bittium" zebrum specimens from Guam, Enewetak, and Hawaii were examined it was discovered that none of these snails had an epipodial skirt, charac- teristic of members of Bittiinae. Moreover, there was no spermatophore bursa present in the lateral lamina of the pallial oviduct, a feature distinctive of the Bittium-group. These animals are thus transferred from Bittitim to Cer- ithium Briiguiere. Further comparison of the Hawaiian specimens of "Bittium" zebrum with those from other Indo-Pacific regions revealed that there are considerable morpholog- ical differences between the populations: the shells, rad- ula and anatomy of the Hawaiian specimens are quite distinct from those of zebrum specimens from elsewhere. Subsequent careful examination of many museum lots of Cerithium zebrum revealed that most lots of Hawaiian specimens comprised mixtures of two species: the first having a typical zebrum phenotype, and the second, more common, species having a different shell physi- ognomy. It became apparent that the second species is Cerithium boeticum Pease, 1860, named from the Ha- waiian Islands and subsequently considered to be con- specific with "Bittium" zebrum by Kay (1979). This paper addresses the differences between the two species, describes and presents a s\ nonymx of each, and delineates the characters separating and distinguishing them. Cerithium boeticum is recognized as a valid spe- cies, different from Cerithium zebrum, and is shown to be endemic to the Hawaiian Islands. MATERIALS AND METHODS Living specimens of Cerithium zebrum were studied at Enewetak Atoll, Marshall Islands, and at the L^niversity of Guam Marine Laboratory at Pago Bay, Guam. Cer- ithium boeticum was collected from shallow, subtidal coral rubble in Kewalo Basin, Honolulu, placed in aquar- ia and observed at the Kewalo Laboratorx of the Ha- waiian Biomedical Research Laboratory, University of Hawaii, Honolulu, Hawaii. Both species were observed and dissected under water in wax-filled petri dishes using a Wild M-5 dissecting microscope. Methylene blue was used to enhance anatomical features during dissection. Protoconchs, shells, opercula and radulae were studied with an Hitachi S-570 scanning electron microscope. The types of both species and their synonymous nomina were examined and are discussed below. Man\- museum lots were examined in order to establish the range of phe- notypic variation in the shells of each species. Measure- ments and meristic data of shells were taken using ran- domly selected specimens from locations throughout the geographic range of each species. R. S. Houbrick, 1993 Page 15 Material examined: Cerithiitm hocticum. HAWAIIAN ISLANDS: (USNM 343522); Oaliu (USNM 12916, types); Honolulu, Oahu (USNM 335496, 335499, 335497, 335498, 767506); Waikiki Marine Lab, Honolulu, Oahu (USNM 633002); Kewalo Basin, Honolulu, Oahu (USNM 857099); Honolulu Reef, Oahu (USNM 335289); Quarantine Is- land, Honolulu, Oahu (USNM 339349, 339342); Waikiki, Oahu (USNM 343514, 343515, 343519); off Waikiki, Ho- nolulu, Oahu (USNM 339118); Kewalo Basin, Honolulu, Oahu, Hawaii (USNM 857099); Diamond Head, Hono- lulu, Oahu (USNM 339339, 343510); Mokapu Beach, Oahu (USMN 484570); Maunalua Beach, Oahu (USNM 428173, 428174, 428207, 343520, 343511); Mokoloe Is- land, Oahu (USNM 339343, 343512); Kaneohe Bay, Oahu (USNM 472057, 472058, 343513); Coconut Island, Ka- neohe Bay, Oahu (USNM 771362, 771556); Hospital Point, Pearl Harbor, Oahu (I'SNM 497938); Pearl Harbor, Oahu (USNM 484455, 428234, 341308); Pupukea Beach, Oahu (USNM 484689); Kahana Bav, Oahu (USNM 777967); Kahala, Oahu (USNM 33951, 339928); Kahala Beach, Koko Head, Oahu (USNM 343546); Keoke River, Hawaii (USNM 252339, 252338, 252336); Keokea, Hawaii (USNM 337592); Keokea, Hilo, Hawaii (USNM 339340, 339340); Hilo. Hawaii (USNM 612281); Kuhio B., Hilo, Hawaii (USNM 339125); Honaunau, Hawaii (USNM 343517); 5 mi SW Kapoho, Hawaii (USNM 409081, 409095); Lanai (USNM 612425); Mauele, Lanai (USNM 339341); Han- alei, Kauai (USNM 340692); Midway Island (USNM 790897, 790898); Frigate Point, Sand Island (USNM 678064). Cerithiumzebriim. MAURITIUS: (USNM 91247, 91069, 91070); Flicq-en-Flacq, Arsenal Bay (USNM 716523). SEYCHELLES: Aldabra Atoll, lie Picard (USNM 837490). COCOS (KEELING) ISLANDS: Alor Pinvu, NW side of West Island (USNM 656435). AUSTRALIA: Heron Is- land, Queensland (USNM 8540174); Watson's Bav, Liz- ard Island, Queensland (USNM 783381). RYUKYU IS- LANDS: Osima, Osumi (USNM 343873); Odomaria, Okinawa, Shima (USNM 488199); Yenoshima (USNM 228222); Chichyima, Ogasawara (USNM 175587). BO- NIN ISLANDS: '(USNM 36950, 343874). PHILIPPINES: Jamelo Bav, Luzon (USNM 935351). MARIANAS: Saipan (USNM 486824, 486823, 486795); Guam (USNM 851 104, 851104); Pago Bay, Guam (USNM 774778); Apra Bay, Guam (USNM 240110); Orote Point, Orote Cliffs, Guam (USNM 854036). CAROLINES: Moen Island, Truk La- goon (USNM 842615); Reef at Mutunlik, Kusaie Island (USNM 609487). LOYALTY ISLANDS: (USNM 253564); Lifu (USNM 423278, 423260, 423210, 423282, 423281, 423261, 423325, 423283). FIJI: Rat Tail Passage, Suva Reef (USNM 824808); SAMOA: Pago Pago (USNM 361535, 361536); Ofu, Manu'a Group (USNM 380968, 380965, 380967, 380969, 380962, 380963, 380960, 380966). MARSHALL ISLANDS: Enewetak (USNM 432415, 770725, 542726); Rennit Island, Enewetak (USNM 542809); Fred, Enewetak (USNM 770687); Aa- raanbiru Island, Enewetak (USNM 582227, 582265); Tei- teiripucoki Island, Enewetak (USNM 581543); Girunien Island, Enewetak (USNM 581534); Engebi Island, Enew- etak (USNM 743869); Rijoru Island, Enewetak (USNM 581555); E. Rigili Island, Enewetak (USNM 581923); between Rigili and E. Rigili Islands (USNM 581576, 581575); Ailuk Island, Ailuk Atoll (USNM 615133); Lae Island, Lae Atoll (USNM 614896); Bock Island, Rongerik (USNM 594660); Enwertok, Rongerik (USNM 583519); 4 mi. W Bikini Islands (USNM 586086); Bikini (USNM 583949, 585185); Bokororvuru, Bikini (USNM 583883, 583884); Namu Island, Bikini (USNM 580542); Enyu Island, Bikini (USNM 580945); Eninman Island, Bikini (USNM 586897); Pigenivarayro, Rongelap (USNM 585588, 585327); Kabelle Island, Rongelap (USNM 582405, 582388, 582125, Burok, Rongelap (USNM 583989); NIUE: Opaahi, S. of Alofi (USNM 854054); Alofi (USNM 854046); Tuapa (USNM 858164); Tuapa, Avaiki Cave (USNM 854055). COOK ISLANDS: Mauke Island (USNM 598174). SOCIETY ISLANDS: Reao Id (USNM 5573, holotype); Makatea, Tuamotu Archipelago, N Ti- mae Harbor (USNM 819895); Arue, Papeete, Tahiti (USNM 775927); Mahina, Tahiti (USNM 797267, 791372); Tahiti (USNM 91068). WALLIS & FUTUNA ISLANDS: outer reef E of Nukuhifala (USNM 676427); Henderson Id (G. Pauley coll.). NIUE: off warf, Alofi (USNM 854046); Avaiki Cave, Tuapa (USNM 854055); Tuapa (USNM 858167); Opaahi, S Alofi (USNM 854054). HAWAII: French Frigate Shoals (ANSP 195368, 195384); Midway (USNM 790897); Honaunau, Hawaii (USNM 343509); Keokea, Hilo, Hawaii (USNM 339353). Abbreviations: BMNH = British Museum (Natural His- tory); MNHNP = Museum National d'Histoire Naturelle, Paris; NMW = National Museum, Wales; SEM = scan- ning electron microscope; USNM = United States Na- tional Museum, Smithsonian Institution, Washington, DC. RESULTS Descriptions (Tables 1 & 2, Figs. 1-28), synonymies, and discussions of both species are presented below, followed by a table identifying the characters separating the two species (Table 3). Cerithium zebrum Kiener, 1841 Cerithium zebrum Kiener, 1841:71, pi. 25, fig. 4 (holo- tvpe, MNHNP; type locality, Indian Ocean, Mauritius). Sowerbv, 1855:875-876, pi. 183, figs. 207-209; 1865, pi. 19, fig. 136; Tryon, 1887: 138, pi. 26, figs. 78-82; Kobelt, 1898:210-211, pi. 39, figs. 7-10; Hedley, 1899:434; Vignal, 1903:21-22, pi. 2, fig. 1; Schepman, 1909:162; Dautzen- berg & Bouge, 1933:313. Cerithium janlhinum Gould, 1849:121 (holotype, USNM 5573, 8.5 X 3.9 mm; type loclaity. Clermont Tonnere [Reao Id, Tuamotusj; 1852:152, pi. 10, fig. 173a-b; 1862:63; Johnson, 1964:95. Cerithium zebrum var. dilectum C.B. Adams in Sowerby, 1855: 896, pi. 33, fig. 207' (15 syntypes, BMNH 1969349; type locality, Galapagos [in error]); Vignal, 1903:22-23, pi. 2, fig. 5. ■ Page 16 THE NAUTILUS, Vol. 107, No. 1 R S. Houbrick, 1993 Page 17 Figs. 16-17. Scanning electron micrographs of radula of Cerithium zebrtim Kiener, Pago Bay, Guam (USNM 774778); 16. view of mid-section of radula, bar = 36 ^m; 17. half row showing details of rachidian, lateral and marginal teeth, bar = 29 ^m. Cerithium delectum (sic) A. Adams. Sowerbv, 1865, pi 16, fig 112 a-c; Vignal, 1903:23. Cerithium unilincatum Pease, 1860:432 (lectot>pe, BMNH 19622798, 5.5 x 3 mm, 2 paralectotypes BMNH 1962799; t\ pe locality. Sandwich Ids [Hawaiian Ids]). Sowerbv, 1866, pi. 15, fig.99; Trvon, 1887:138, pi. 26, fig. 86; Kobelt, 1898: 224, pl.29, fig. 18. Cerithium stigmosum Gould, 1861:386 (holotype, ?; type lo- cahty, Bonin Ids) 1862:141; Johnson, 1964:152. Cerithium aspersum Deshayes, 1863:97-98, pi. 11, figs. 16-18 (lectotype, here selected, MNHNP, 8.8 mm, 4 paralecto- types, MNHNP; type locality. Reunion). Cerithium Crossii Deshayes, 1863:96, pi. 11, figs. 12-14 (lec- totype, here selected, MNHNP, 8.4 mm, 3 paralectotypes, MNHNP; type locality. Reunion; not Cerithium cros- seanum Tiberi, 1863 [= Cerithioipsis sulwylindricum (Brusina)], nor Cerithium maillardi Crosse, 1863 [unnec- essary replacement name for Cerithium crosseanum Ti- ber!]. Cerithium dichroum Melvill is. Standen, 1895: 115, fig. (6 syn- types NMW 55158200; type locality, Lifu, Lo\aity Idsi Cerithium zebrum var. pulchra Vignal, 1903:22, pi. 2, fig. 2 (t>pe, MNHNP; type localit> , Isle of Pines, New Cale- donia); Dautzenberg & Bouge, 1933:315. Cerithium zebrum var. trijasciata N'ignai, 1903:22, pi. 2, fig. 3 (type, MNHNP; type locality. Isle of Pines, New Cale- donia). Cerithium zebrum var. attenuata Vignal, 1903:22, pi. 2, fig. 4 (type, MNHNP; type locality. Isle of Pines, New Cale- donia); Dautzenberg & Bouge, 1933:313. Cerithium zebrum var undulata Dautzenberg ic Bouge, 1933: 316 (Type, MNHNP; type locality, Tuamotus). Cerithium zebrum var. rosea Vignal, 1903:24, pi. 2, fig. 6 (type, MNHNP; t\pe locality. Isle of Pines, New Caledonia); Dautzenberg & Bouge, 1933:315. Cerithium zebrum var. cinerea Vignal, 1903:24, pi. 2, fig. 7 (type, MNHNP; type locality. Isle of Pines, New Cale- donia); Dautzenberg & Bouge, 1933:314. Cerithium zebrum var. nivea Vignal, 1903:24, pi. 2, fig. 8 (type, MNHNP; type locality. Isle of Pines, New Caledonia); Dautzenberg & Bouge, 1933:315. Cerithium zebrum var. biUneata Vignal, 1903:25, pi. 2, fig. 10 Figs. 1-15. Cerithium zebrum Kiener, showing shell variation and types of synonymous nominal ta.xa. 1-2. Orote Point, Guam, 7.9 mm length; 3. operculum, 1.3 mm length (ISNM 854036); 4. Orote Point, Guam, 7.5 mm length (USNM 854036); 5. Pago Bay, Guam, 7.8 mm length (USNM 774778); 6. Enewetak Atoll, Marshall Islands, 7.9 mm length (USNM 770725); 7. lectotype of Cerithium crossii Deshayes, Reunion (MNHNP), ; 8. Holotype of Cerithium janthinum Gould, Reao, Tuamotus (USNM 5573); 9. SEM of protoconch, bar = 70 ^m; 10. lectotype of Cerithium aspersum Deshayes, Reunion, 8 2 mm (MNHNP); 11-12. Pago Bay, Guam, apertural and right lateral views under SEM, showing sculptural details, 5 8 tnm length (I'SNM 774778); 13. sculpture of early whorls, SEM, bar = 0.9 mm; 14—15. Pago Bav, Guam, dorsal and right lateral views under SEM, 5.8 mm length (USNM 774778). Page 18 THE NAUTILUS, Vol. 107, No. 1 Figs. 18-26. Cerithium boeticum Pease, from Kewalo Basin, Honolulu, Oahu, Hawaii, showing shell variabilit) (USNM 857099). 18-19. Strongly banded morph, 5.8 mm length; 20-21. white morph, 5.7 mm length; 22-23. 5.7 mm length; 24. scanning electron micrograph showing sculptural details and aperture, 4.9 mm length; 25. operculum. 2 mm length; 26. SEM of protoconch and early whorl sculpture, bar = 88 fzm (type, MNHNP; type locality. Isle of Pines, New Cale- donia); Dautzenberg & Bouge, 1933:314. Cerithium zelyrum var. triliyieata Vignal, 1903:25, pi. 2, fig. 11 (type, MNHNP; type locality. Isle of Pines, New Cal- edonia); Dautzenberg & Bouge, 1933315. Cerithium zebrum var. ornata Vignal, 1903:26, pi. 2, fig. 13 (type, MNHNP; type locality. Isle of Pines, New Cale- donia). Cerithium zebrum var. sticta Vignal, 1903:26, pi. 2, fig. 14 (type, MNHNP; type locality, Isle of Pines, New Cale- donia); Dautzenberg & Bouge, 1933:315. Cerithium zebrum var mutimaculata Vignal, 1903:26, pi. 2, fig. 15 (type, MNHNP; type locality. Isle of Pines, New Caledonia); Dautzenberg & Bouge, 1933:315. Cerithium zebrum var. maculata Vignal, 1903:26, pi. 2, fig. 16 (type, MNHNP; type locality. Isle of Pines, New Cal- edonia); Dautzenberg & Bouge, 1933:314. Cerithium zebrum var. unimacuhita N'ignal, 1903:27, pi. 2 fig. 17 (tspe, MNHNP; type localit\', isle of Pines, New Cal- edonia). Cerithium zebrum var. aspersa Deshayes. Dautzenberg & Bouge, 1933:314. Bittium zebrum (Kiener) Cernohorsky, 1972:69-70, pi 14. fig. 10; Salvat & Rives, 1975:272, fig. 74. Bittium zebrum (Kiener) Kay, 1979:120, figs. 45b-c (not Bit- tium zebrum (Kiener), is Cerithium boeticum Pease 1860). Description: Shell (Figs. 1-15; Table 1): tiirreted, elon- gate, attaining 10.4 mm length, 3.2 mm width, and com- prising about 9 weakly inflated to flat-sided whorls. Pro- R. S. Houbrick, 1993 Page 19 Figs. 27-28. Scanning electron micrographs of radula of Cerithium boeticum Pease, Kewalo Basin, Honolulu, Oahu, Hawaii (I'SNM 857099). 27. general view of mid-radular ribbon, bar = 25 ^m; 28. detail of rachidian tooth, bar = 9 nm toconch (Fig 9) comprising 3 whorls; protoconch 1 nearly smooth, sculptured with suprasutural spiral cord; pro- toconch 2 sculptured with series of subsutural plaits fol- lowed by two strong spiral cords, criss-crossed by minute lirae up to anterior suture. Early whorls (Fig. 13) sculp- tured with 3 beaded spiral cords. Adult whorls sculptured with 4 beaded, major spiral cords each separated from one another by one fine spiral lira. Penultimate whorl with 5 beaded spiral cords, having about 26 spiral beads; beads frequently aligned to form weak a.xial riblets. Body whorl with 11-12 beaded spiral cords and fine spiral lirae. Suture weakly defined. Strong white varices randomly placed on whorls; largest varix on right dorsal side of body whorl. Aperture circular-ovate, a little over one- fourth the shell length. Outer lip thick, relatixely smooth, weakly crenulate at inner edge; columella concave with weak callus. Anterior canal short, but pronounced, ori- ented at 45 degree angle to shell a.xis. Weak siphonal constriction. Anal canal small, defined bv small parietal plait Shell color white with pink early whorls and adult w horls w ith brown, tan, rose blotches and spiral stripes, bands, or dots. Aperture white. Periostracum thin, light tan. Animal: Headfoot white, flecked with opaque white, rose or tan. Snout, mantle edge and siphon \ellow . Mantle edge with short yellow papillae. Snout long, extensible, having bilobed tip (lips). Operculum (Fig. 3) tan, cor- neous, ovate, paucispiral with eccentric nucleus. Lateral epipodial skirt and operculiferous lobe absent. Osphra- dium bipectinate, with weakly-defined pectins. Radula (Figs. 16-17) short, about one-ninth the shell length. Rachidian tooth (Fig 17) height equals length, having hourglass-shaped basal plate with spade-shaped basal projection and two small basal lateral butresses; cutting edge with large, pointed central cusp flanked by two small pointed denticles on each side. Lateral tooth with rectangular basal plate having long lateral projection and Table 1. Shell statistics for Cerithium zcbrum (measurements in mm) Table 2. Shell statistics for Cerithium boeticum (measure- ments in mm). Statistic (n = 17) X sd Range Statistic (n = 17) X sd Range Length 7.3 1.5 4.2-10.35 Length 6.1 1.8 2.4-10.5 Width 2.4 0.4 1.8-3.2 W^idth 2.3 0.8 1.2-4.6 Aperture length 1.3 0.3 0.8-1.8 Aperture length 1,7 0,7 0.8-2.4 Number of whorls 9.2 1.6 7-13 Number of whorls 8,0 0,9 6-9 Number spiral beads 26,3 6 19-44 Xvunber axial beads 174 1 7 14-20 Page 20 THE NAUTILUS, Vol. 107, No. 1 thick central buttress; cutting edge with one weak inner denticle, large spoon-shaped cusp, and three outer den- ticles. Marginal teeth spatulate, curved at tips; inner mar- ginal tooth with three pointed inner denticles, large cen- tral cusp and two outer denticles; outer marginal tooth same but without outer denticles. Esophageal gland pres- ent; stomach large; style sac and crystalline style long, about one-third stomach length. Synonymic remarks: This species has been the recipient of 23 names, all of which have been based on concho- logical characters and/or differences in shell pigmenta- tion. It is obvious that the authors of these nomina did not employ a modern species concept and failed to rec- ognize the full range of intraspecific variation when pro- posing these ta.xa. Examination of the types of the syn- on\mous nomina re\ealed no significant differences in shell sculpture from that of Cerithium zebrum (Kiener); indeed, the nomina proposed by Vignal (1903) as vari- eties of Cerithium zebrum Kiener were based on mere intraspecific color variation, as Vignal (1903) clearly stat- ed. The holotype of Cerithium janthinum Gould (see Fig. 8) is merely a worn, very wide morph of Cerithium zebrum. Sowerby (1855) introduced the name dilectum in the Thesaurus as a variety of Cerithium zebrum Kie- ner, but the new taxon was indicated only by the itali- cized name in his "Alphabetical list of species", on p. 896. The name dilectum was initially listed on p. 117 as a synonym of Cerithium zebrum, and the legend for the figures of Cerithium zebrum (figs. 107-109) does not include the name dilectum, although one of the figures has an asterix (fig. 207-), which presumably is meant to indicate the varietal taxon, dilectum. Cerithium asper- sum Deshayes is a highly sculptured morph of C. zebrum, as illustrated by the lectotype (Fig. 10). Cerithium crossi Deshayes, 1863, which as shown by the lectotype (Fig. 7), is conspecific with Cerithium zebrum, was incorrectly considered by Crosse (1863) as a homonym of Cerithium crosseanum Tiberi, 1863, a Cerithiopsis species, and was given an unnecessary replacement name, Cerithium maillardi Crosse, 1863. Cerithium boeticum Pease er- roneously has been considered conspecific with Ceri- thium zebrum Kiener by Kay (1979). For a discussion of this issue, see the discussion under Cerithium boeti- cum. Discussion: Cerithium zebrum is a very common spe- cies, frequently found in the upper intertidal zone among rubble and algae on benches and platforms associated with fringing coral reefs of atolls and islands throughout the Indo-Pacific. In the Hawaiian Islands is also occurs in deeper waters (Kay, pers. com.). It is characterized by an elongate, tapering shell, sculptured with many fine beaded spiral cords and with numerous broad, white, randomly-placed varices. Cerithium zebrum is a colorful species, having a rose-colored apex capped with a dark brown protoconch (see Figs. 1-2,4,6). The adult whorls are frequently white or light pink with tan blotches or with brown and tan bands. While the species varies great- ly in color pattern, shell sculpture is relatively stable throughout its range. Banded varieties of Cerithium ze- brum may be confused with banded morphs of Ceri- thium boeticum, but the latter species has a more coarse- ly sculptured shell with stronger axial ribs, a more deeply incised suture, and a thick outer apertural lip (see Figs. 18-26). The animal (headfoot), in contrast to the brightly pigmented shell, is a uniform white overlain by opaque white spots, although some shells may also be white. In contrast to the lecithotrophic development seen in Cerithium boeticum, Cerithium zebrum undergoes a lengthy planktotrophic larval phase and has an elabo- rately sculptured protoconch (Fig. 9) with a deep sinu- sigeral notch indicative of this de\elopmental mode. The operculum of Cerithium zebrum is slightK- more ovate, thicker and opaque, and has fewer spirals than that of Cerithium boeticum. Cerithium zebrum occurs from Henderson Id., in the Pitcairn Group, throughout French Polynesia and the Hawaiian Islands, and westward throughout the tropical Pacific from Queensland, Australia, north to the Ryu- kyus. It is found throughout the archipelagos of southeast Asia and in suitable localities in the Indian Ocean. In- deed, specimens from Tamarin Bay, Mauritius (AXSP 273492) look very much like those found in Tahiti (Fig. 6) and Guam (Figs. 1-2, 4-5). While Cerithium zebrum has occasionally been collected in assemblages from depths greater than 10 m around the main, volcanic Hawaiian Islands (Kay, in lit.), it is more common on the shallow reefs of the Northwestern Hawaiian Islands at French Frigate Shoals (ANSP 195368, 195384) and Midway (USNM 790897). To my knowledge, Cerithium zebrum has not been recorded as a fossil. Cerithium boeticum Pease, 1860 Cerithium pusillum Gould, 1851: 120-121 (holotype, USNM 5572, 4 paratypes, MCZ 216606; type locality, Sandwich Ids [Hawaiian Ids); nol Cerithium pusillum Pfeiffer, 1840, nor Dunker,1859); Gould, 1852:150, pi 10, fig. 172a-b; Gould, 1862:62; Dunker, 1882:108. Cerithium boeticum Pease, I860: 433 (lectotype [Kay, 1965], BMNH 1962802, 6 x 2.5 mm, 5 paralectotypes, BMNH 1962803; type locality. Sandwich Ids [Hawaiian Ids]); Sow- erby, 1866 "haeticum", pi. 12, fig. 327; 1866b, pi. 16, fig. 114; Trvon, 1887:143, pi. 27, fig. 26; Kobelt, 1898:236, pi. 41, fig. 14; Kay, 1965:48-49, pi. 10, fig. 8. Biitium pusillum (Gould) Johnson, 1964: 136; Kay, 1979:120 (not Biitium pusillum [Dunker, 1859]). Bitlium zebrum (Kiener) Kay. 1979: 120. fig. 45b-c (not ze- brum Kiener, 1841, is Bittium pusillum [Gould, 1849]). Bittium boeticum unilineatum Pse. (sic). Mant, 1923: 121. Bittium boeticum Pils. & Va. (sic). Mant, 1923: 121. Description: Shell (Figs. 18-24, 26; Table 2): Shell solid, turreted, reaching 10.5 mm length and 4,6 mm width, comprising about 8 con\ex whorls. Protoconch (Fig. 26) one whorl, smooth with straight lip. Early whorls sculp- tured with two spiral cords and weak axial ribs. Penul- timate whorl sculptured \\ ith 4 major beaded, spiral cords R. S. Houbrick, 1993 Page 21 each separated from the other with weak spiral Hrae, and each bearing about 17 beads aHned to form 17 strong axial ribs. Bod\ whorl large, sculptured with 9 spiral beaded cords and weak spiral lirae. Several varices ran- domly distributed on shell, strongest opposite outer lip of aperture. Suture deeply impressed. Aperture oval- elongate, a little over one-third the shell length. Anterior canal short, well-defined, oriented at 45 degree angle to shell axis; outer lip thick, strongly denticulate along inner edge; columella concave with moderate callus. Anal ca- nal weak, defined b\ parietal plait. Animal: Headfoot without lateral epipodial skirt; weak- ly scalloped operculiferous lobe present. Operculum (Fig. 25) corneous, very thin, transparent and paucispiral with slightly eccentric nucleus. Osphradium very large, about one-half w idth of ctenidium, strongK bipectinate with large pectins. H> pobranchial gland well-developed. Me- dial lamina of pallial oviduct with spermatophore bursa and large seminal receptacle. Radula (Figs. 27-28) very short, about one-twelfth the shell length. Rachidian tooth (Fig. 28) wider than tall, hourglass-shaped, with basal plate having short basal projection and two small lateral buttresses; cutting edge with spade-shaped central cusp flanked on each side with three, sometimes two, small pointed denticles. Lateral tooth w ith rectangular basal plate having long lateral projection and short basal but- tress; cutting edge with small inner denticle, long spoon- shaped major cusp, and two, sometimes three, outer pointed denticles. Marginal teeth elongate, spatulate with curved tips; inner marginal tooth w ith two inner denticles long, major central cusp and one outer denticle; outer marginal tooth same but without outer denticle. Synonymic remarks: Cerithium boeticum has been confused with several other taxa assigned to Cerithium or Bittiiim by authors. The earliest name for the species is Cerithium pusillum Gould, 1851, but as this name is preoccupied, the next available valid name is Cerithium boeticum Pease, 1860. Although the figures subsequently presented by Gould (1852, pi. 10, figs. 172a-b) for Cer- ithium pusillum are not very good, the holotype (USNM 5572) clearK shows that this taxon is different from Cer- ithium zebrum Kiener, with which it has been confused (e.g., Kay, 1979; 120). Cerithium pusillum Dunker, 1859, and Cerithium pusillum Gould, 1851, are different spe- cies, but as both taxa have been referred to Bittium by authors, some confusion has developed about their iden- tity. Cerithium boeticum Pease is a name seldom men- tioned in the literature and has not been a well-under- stood species. Kay (1965) designated a lectotype and five paralectotypes from the original type lot of six beach- worn specimens, but her illustration of the lectotype (1965, pi. 10, fig. 8) is poor and is not helpful in clarifying the concept of the species. Kay (1979) subsequently placed this taxon into the synonymy of Cerithium zebrum, a species with a wide Indo-Pacific distribution. Further confusion has developed because another Pease (1860) name, Cerithium unilineatum, a valid svnonvm of Cer- ithium zebrum Kiener, was combined with Cerithium boeticum Pease to form a trinomial In some museum collections the manuscript name, Cerithium gouldianum Pilsbry & Vanatta, is found associated with lots of Cer- ithium boeticum. but the name has no validity. Discussion: As mentioned above, Cerithium boeticum had been condsidered to be synonymous with the com- mon Indo-Pacific species, Cerithium zebrum Kiener, but the former species has a thicker, less elongate shell with coarser sculpture, few weak varices, a stronger anterior canal, and a different kind of protoconch than Cerithium zebrum (see Figs. 18-26); moreover, Cerithium boeti- cum appears to be restricted to the Hawaiian Islands, whereas Cerithium zebrum is distributed throughout the Indo-Pacific. Shells of the two species frequently share similar color patterns, especially among the banded, striped morphs, but when closely compared, they are seen to be realK' quite different. Radular differences be- tween the tw o species although slight, are consistent and serve to distinguish each of them (see Table 3). The very thin, nearly transparent operculum of Cerithium boe- ticum (Fig. 25) differs from the thicker one of Cerithium zebru m . Gould (1851) mentioned that Cerithium boeticum (cited as Cerithium pusillum) was "like C. ferrugineum Sa\-, but much smaller". Cerithium ferrugineum is a s\nonym of Cerithium lutosum Menke, a small species from the western Atlantic, and indeed Cerithium boe- ticum does bear some resemblance to small morphs of Cerithium lutosum in overall shell morphology (see Houbrick, 1974, pi. 41). An unusual anatomical feature of Cerithium boeticum is the large, well-developed bipectinate osphradium. Al- though Cerithium zebrum also has a bipectinate os- phradium, the pectins are poorly defined. A bipectinate osphradium is common to all members of Cerithium Bruguiere and Chjpeomorus Jousseaume (see Houbrick, 1974, 1985), but not as common among Bittium species (pers. obsr). Although Cerithium boeticum has a weakly scalloped operculiferous lobe, the lateral epipodial skirt, indicative of Bittium species, is absent. The absence of a spermatophore bursa in the lateral lamina of the pallial oviduct, which is the hallmark of Bittium species, is one of the chief anatomical reasons for reassignment of this species to Cerithium. However, the specimens studied herein may not have attained complete reproductive ma- turity, and this needs reconfirmation. Cerithium boeticum lives in shallow water on subtidal rubble where it is frequently found on marine algae. This species has lecithotrophic development, which is reflected in its relatively unsculptured protoconch, com- prising only one whorl with a nearlv straight apertural Up (Fig. 26). Cerithium boeticum appears to be endemic to the Hawaiian Islands. It is geographically sympatric with Cerithium zebrum, especially in the northern part of the Hawaiian chain. This species has been recorded from the Pleistocene of Molokai (Ostergaard, 1939, as Ceri- thium boeticum). Page 22 THE NAUTILUS, Vol. 107, No. 1 Table 3. Characters defining Ceritbiurn zclmim and Ceri- thiitrn hoeticum. Cerithium zebruni Cerilhium hoeticum Sculpture 1. Weak axial riblets Strong axial ribs 2. Many weak spiral beads Fewer strong spiral beads 3. Weakly impressed suture Strongly impressed suture 4. Many broad white varices Few weak varices 5. Protoconch highly sculji- Protoconch unsculptured tared 6. Thin outer apertural lip Thick outer apertural lip 7. Aperture '74 shell length Aperture 'i shell length Radula 1. Radula short Radula very short 2. Rachidian height equal to Rachidian wider than tall width 3. Rachidian cusp flanked Rachidian cusp flanked with 2 denticles on each with 3 denticles on each side side 4. Inner marginal with 3 in- Inner marginal with 2 inner ner and 2 outer denticles and one outer denticle Osphradium 1. Weakly bipectinate Large, strongly bipectinate Development 1. Plantotrophic Lecithotrophic A sumniar) ol the dittereiices between these two spe- cies is presented below, in Table 3. CONCLUSIONS Evidence is presented above to show that the Hawaiian species, Cerithium hoeticum, is morphologically distinct from Cerithium zebrum. Although both species had pre- viously been assigned to Bittium. anatomical evidence shows that this is incorrect. As both taxa were thought to be Bittium species, they were omitted from my Indo- Pacific Cerithium monograph (Houbrick, 1992). Ceri- thium hoeticum is restricted to the Hawaiian Islands while Cerithium zebrum has a very wide Indo-Pacific distribution, including the Hawaiian Islands. ACKNOWLEDGEMENTS I thank Michael Hadfield, of the Univeristy of Hawaii, Honolulu, Hawaii, for the use of laboratory space and for assistance during field studies in Hawaii. I am grateful to Alison Kay for valuable discussions about the geo- graphic distribution of both species throughout the Ha- waiian Islands. Lou Eldredge, University of Guam, kind- ly provided me with lab space and logistic support at the Pago Ray lab of the University of Guam. The work at the marine laboratory at Enewetak Atoll was made pos- sible by a grant from the Atomic Energy Commission. Bernadette Holthuis, University of Guam, kindly ob- served living Bittium zebrum specimens in the field aiul passed on her observations to me. I am grateful for the use the Smithsonian Secretary's Research Opportunity Fund, w hich covered the costs of transporation and lodg- ing in both Guam and Hawaii during this study. Susanne Braden of the Smithsonian Scanning Electron Micro- scope Lab provided assistance with the SEM micrographs and V'ictor Krantz, Smithsonian Photographic Services, helped with photography. LITERATURE CITED Cernohorsky, W. O., 1972. Marine shells of the Pacific, Vol. 2, 411 pp., 68 pis. Pacific Publications, Ltd., Sydney. Crosse, M.H., 1863. Description d'especes nouvelles d'Aus- tralie. Journal de Conchyliologie 11:84-90, pi. 1. Dautzenberg, P. & J.L. Bouge, 1933. Les mollusques testaces marins des etablissements Fran9aise de I'oceanie. Journal de Conchyliologie 27(2);145-469. Deshayes, G.P., 1863. Catalogue des mollusques de I'lle de la Reunion (Bourbon). 144 pp., pis. Paris, Dentu. Dunker, G. 1859-1860. Neue japanische Mollusken. Mala- kozoologische Blatter 6:221-240. Dunker, G. 1882. Index Molluscorum Maris Japonica. Nov- itates Conchologiae, Supplement to Volume 7. Cassell. Gould, A. 1851. Descriptions of new species, brought home by the US, Exploring Expedition Proceedings of the Bos- ton Society of Natural History 3:118-121. Gould, A. 1852. Mollusca and shells, X'olume 12, 510 pages, atlas in US. Exploring Expedition during the vears 1838, 1839, 1840, 1841, 1842 under the command of Charles Wilkes, U. S.N.Boston: Gould & Lincoln. Gould, A. A. 1861. Descriptions of shells collected in the North Pacific Exploring Expedition under Captains Ringgold and Rodgers. Proceedings of the Boston Societv of Natural History 7: 385-389. Gould, A..\. 1862. Otia Conchologica: Descriptions of shells and mollusks, from 1839 to 1862, 256 pp, Boston, Gould & Lincoln, Gray, J, E. 1847. A list of the genera of Recent Mollusca, their synonyma and types. Proceedings of the Zoological Society of London (for 1847) 15(I78):129-219 (November 1847). Hedley, C. 1899. The Mollusca of Funafuti, Part 1, -Gaster- opoda. Australian Museum, Svdnev, Memoir 3, pp, 397- 488. Houbrick, R.S, 1974, The genus Cerithium in the western Atlantic. Johnsonia 5(10):33-84, 48 pis. Houbrick, R.S. 1985. Genus Clypeomorus Jousseaume (Cer- ithiidae: Prosobranchia). Smithsonian Contributions to Zo- ology No,403: iv + 131 pp,, 62 figs, Houbrick, R.S. 1992. Monograph of the genus Cerithium Bruguiere in the Indo-Pacific (Cerithiidae: Prosobranchia). Smithsonian Contributions to Zoology No. 510: 208 pp., 145 figs. Johnson, R. 1964, The Recent Mollusca of .Augustus .\ddison Gould. Bulletin of the United States National Museum 239: v -I- 182 pp., 45 pis. Kay, E. A. 1965. Marine mollusks in the Cuming collection, British Museum (Natural Histor\ ) described by William Harper Pease. Bulletin of the British Museum (Natural Histor\ ), ZoologN , Supplement 1, London 96 pp., 14 pis. Kay, E,A, 1979. Hawaiian Marine Shells Bishop Museum Press, Special Publication 64(4), Honolulu, Hawaii, 653 PP Kiener, L.C. 1841. Species genera! et iconograpliie des co- R. S. Houbrick, 1993 Page 23 quilles vivantes. Genre Cerite Paris vol 5: 10-4 pp , 32 pis. Kobelt, W. 1888-1898. Die Gattung Ccrithium. in F. H. W. Martini, & J. H. Chemnitz, Neues systematisches Conchy- iien-Cabinet ... 1 (pp. 297 pages). Nurnberg: Bauer & Raspe. Melvill, J.C. & R. Standen. 1895. Notes on a collection of shells from Lifu and Uvea, Loyalty Islands, formed b\ the Rev James and Mrs. Hadfield, with a list of species. Journal of Conchology 8: 84-132, pis. 2-3. Mant, C. F. 1923. Mollusca from the dredging operations at Kewalo Harbor, Honolulu, 1921. The Nautilus 36(4):120- 123. Ostergaard, J.M. 1939. Reports on fossil Mollusca of Molokai and Maui. Occasional Papers of Bernice P. Bishop Museum Honolulu Hawaii 15(6): 67-77. Pease, W. H. 1861, Descriptions of Forty-seven New Species of Shells, from the Sandwich Islands, in the Collection of Hugh Cuming, Esq. Proceedings of the Zoological Society of London 431-438. Pfeiffer, L 1840. Ubersichtder im Januar, Februar und Marz, 1839 auf Cuba gesammelten Mollusken. Archiv Natur- geschite6(l);250-261. Salvat, B., & C. Rives. 1975. Coquillages de Polynesie. 392 pp, illustrated Papeete, Tahiti. Schepman, MM. 1909. The Prosobranchia of the Siboga Ex- pedition. Part 2. Taenioglossa and Ptenoglossa. Leyden, Brill, 7 pis. Sowerby, G.B. 1855 Monograph of the genus Cerithium. .'\danson. In Thesaurus Conch) liorum, or monographs of genera of shells. London, 2(16);847-899, pis. 176-186. Sowerby, G.B. 1865-66. Monograph of the genus Cerithium Adanson in: Reeve, Conchologia Iconica: or illustrations of molluscous animals. London. 15 (atlas), 20 pis. + index. Tiberi, N. 1863. Description d'especes nouvelles de la mer Mediterranee. Journal de Conchyliologie 11: 158-160. Tryon, G.W. 1887. Manual of Conchology; structural and systematic with illustrations of the species (Cerithiidae), First series. Philadelphia, 9: 112-228, 19-39 pis. Vignal, L. 1903. Sur les varietes du Cerithium zebrum Kiener. Journal de Conchyliologie 51(l):21-27, pi, 2. THE NAUTILUS 107(1 ):24-28. 1993 Page 24 Rediscovery of TurhineUa thersites Reeve, 1847, with Notes on its Taxonomic Position (Gastropoda: Fasciolariidae) R. N. Kilburn Natal Museum P/Bag 9070 Pietermaritzburg 3200, South Africa ABSTRACT The holot\pe of Turbinella thersites Reeve, 1847, was believed to have been collected in China; no further specimens have subsequently been reported. Comparison with recently trawled material indicates that it was actually dredged on the Agulhas Bank. Turbinella thersites is here transferred from the family Turbinellidae to the genus Fasciolaria, within the family Fas- ciolariidae. Key words: Fasciolariidae; Turbinellidae; South African; ne- ritic; Fasciolaria. Turbinella INTRODUCTION Turbinella thersites Reeve, 1847, was based on a single specimen (Fig. 3) originally in the collection of Captain Sir Edward Belcher, and supposedK- from China. Noting that it appeared to be immature, Ree\e obser\ed that "it rather partakes of the character of Fasciolaria", a view echoed b\- Melvill (1891), who speculated that it might belong to either Fasciolaria Lamarck, 1801 (fam- ily Fasciolariidae) or Tudicla Roding, 1798 i^Turbinel- lidae). Kobelt (1874) cited it as Turbinella (Plicatella) thersites, and Tryon (1881) similarly included it within the genus Latirus Montfort, 1810 (= Plicatella S\\ ainson. 1840). FinalK', Yen (1942), in a paper on C^hinese gas- tropods in the Natural Histor\ Museum, London, pub- lished a photograph of the holot\pe, under the name Tudicla thersites, and quoted the type locality without comment. No specimens additional to the holot\pe are on record, and no further reference to the species has been traced, notwithstanding the considerable bod\' of literature that has been published in recent decades on the mollusks of the Sino-japanese region. ,\mong the larger mollusks that are brought up in the nets of commercial trawlers operating on the .Vgulhas Bank, the continental shelf south of South Africa, are several species of Fasciolaria. Most commonly trawled are F. scholvieni Strebel, 1912, bathymorphs of F. lu- gubris lugtdnis Ree\e. 1847, and its eastern sulxspecies F. lugubris heynemanni Dunker, 1871, and the bathyal F. rutila (Watson, 1882); less commonly seen is F. uat- tersac Kilburn. 1974. On rare occasions a fifth species. distinguishable by its sculpture of auriculate ribs, is trawled. Such ribs similarK' characterize the "lost" Tur- binella thersites, and comparison of juvenile specimens of this "fifth species" with the equally juvenile holotype of T. thersites confirms their identity . .At the time of description of Turbinella thersites, the only .\gulhas Bank mollusks known had been dredged during the cruises of the H.M.S. Sulphur USo6-1842) and H.M.S. Samarang (1843-1846'. both captained by Belcher. Confusion of localities in Belcher material from these voyages is not infrequent, and using the line of reasoning ad\anced b\ Tomlin (1925: 310). with regard to the origins of the holotvpe of Turrit ella ferruginea Ree\e. 1849, one can speculate that the holotv pe of Tur- binella thersites was originalK amongst the mollusks dredged by Belcher on the .\gulhas Bank during the vo\age of H.M.S. Samarang i^this expedition was spe- cificalK mentioned b\ Ree\e under the description of Turbinella [= Latirus] belcheri Ree\e. 18471 Indeed the case of Turrit ella declivis .\dams & Reeve. 1S50. is directly comparable: although originalK said to have been dredged b\ the Samarang in the "China Sea", this species was subsequentK shown i,Tomlin. 1925: 313) to be characteristic of the Agulhas Bank biota. Although at present little precise data is a\ ailable for Fasciolaria ther- sites. indications are that, like Turritclla dcclivis. it is a continental shelf species endemic to the .\gulhas Bank. The second known example (Fig. 5) of Fasciolaria thersites was actually collected in 1962. but its identity w ent unrecognized, until it w as .sent to me for comment in 19S(i b\ J. H. McLean of the Los .\ngeles Count) Mviseum. CuriousK . a.ssociated with it w as a similar error in localization. Originating from the collection of the late I'ranz Steiner. the label states: "This shell w as found in dredged santl. Nacala, Mozambique, on the 19 June, 19ti2, b\ nic . BiogeographicalK , it would be unprece- dented for such a temperate-w ater .\gulhas Bank species to live also on the tropical coral coast of northern Mo- zambique, and its origin is sureK ad\ entitious. It is not impossible that this specimen w as jettisoned from a ship docking in Nacala harbor, but although worn, it lacks the porcellaneous appearance normally seen in shells that ha\e lain on coral sand for a length) period. It is thus R. N. Kilburn. 1993 Pa2e25 Figures 1-2. Protoconch of Fasciolaria thersites Reeve. 18471 Scale lines = 1 mm. more probable that Steiner accidentalK separated it from a sample of trawled Cape moUusks. and mLxed it with his Mozambican finds. Indeed, in 1962 or 196-3 I was personalK told b\ Steiner then a radio-operator in the merchant navy) that while on the Xacala route he reg- ularK received trawled shelb from a source in Cape Town, who accumulated them in barrels of preserv ative on his behalf. This species and its temperate-water South .\frican congeners agree in teleoconch characters with the genus Fasciolaria Lamarck. 1799. in its broadest sense. Radula characters, known for all species v Kilbum. 1974. Barnard. 1958). except F. thersites. also agree, .\lthough species with shoulder nodules are generally referred to subgenus ^sometimes even genus) Pleuroploca P. Fischer. 1SS4. such a classification does not address the problem of ta.xa such as Fasciolaria filamentosa \R6ding. 1798) and F. lugubris Reeve. 1S47. in which nodules are present or absent according to individual or population. The fi\e species endemic to South .\frican waters share a close morphological similarity but differ from all other species of Fasciolaria in their extremely large and bulbous pro- toconch (Figs 1-2'. which has a basal diameter of 3.0- 5.0 mm. according to species and individual. Such a protoconch cannot be dismissed as merely an adaptive correlation with mode of development, because this is similarly non-planktotrophic in four of the five other species of Fasciolaria (= Pleuroploca'' in which devel- opment is known [see D'.\saro. 1970a. 1970b. Gohar & Eisawy . 1967. Shepherd & Thomas. 1989). and is there associated with a much smaller protoconch. .\ second character of possible significance is egg-capsule shape: in the sole Cape species in which these are known {F. lugubris\ capsules differ from those reported for other species of Fasciolaria in being dorsally rounded without a demarcated escape hatch ^see Kilburn & Rippey. 1982: text fig 62 i. Egg capsules have been described for three species from North .\merica. and one each from the tropical Indo-Pacific, Mediterranean and southern -\us- traha ; references given above, also BaccL 1947). In all of thec-e. the capsule is flat-topped, with a structural es- cape hatch dorsally. It is possible that recognition of the South -\f rican species-complex at the subgenxis level may ultimately prove warranted on grounds of protoconch size and perhaps capsxile form. Fasciolaria thersites Figures 1-7 Reeve. 1S47 , n. comb. Turbinella thersites Reeve. 1S47: pi. 4. sp 21: MeKiU. 1S91: 409. Type localit> : China [here emended to .\gulfaas Bank]. Turbinella , Flicatella '• thersites: Kobelt in Kster & KobelL 1574: 71. 157. pL 18. fig. 1 ^afte^ Reeve': Kobelt 1S76: 21. Lc-'":' -rf-rfites: Tryon. 1851: 91. pL 6S. fig. 137 .after Reeve). 7 '-'^tes: Yen. 1942; 237, pL 25. fig. 17S iholot>-pei. r ^>--- .i^'-.i^ icattersae ynon Kjlbum. 1974 : Richards. 1981: 63, pi. :35. fig. .300. Description: Shell fusiform, with bluntly papiUiform apes, slphonal canal equal in length to aperture, curved slighdv to right: suture moderately shallow, whorls strongly convex, except immediately below suture where they are shaUowly concave. Inner fip with a thin callus glaze, smooth except for a thin spiral ridge just above mid-parietal region and three obhque basal pleats; of these the strongest pleat is situated at entrance to siphonal canal and the weaker posterior two decrease in strength and are scarcely visible from without the aperture. Te- leoconch sculptvtred by strongly prosochne. rather turn id axial ribs, which project at shoulder in a roundedly au- riculate manner: in most examples, asials become weak on later whorls, except sometimes for an occasional strong rib, but they may remain strong throughout: fine spiral threads, crossed by microscopic coUabral threads, present overall. First teleoconch whorl with 8 straight prosocline asials. the first few low and relatively close, thereafter in transverse section thick, strongly rounded, slightly Page 26 THE NAUTILUS, Vol. 107, No. 1 R. N. Kilburn, 1993 Page 27 compressed, more or less equal to intervals, which are flattened; Second whorl with 6-7 axials, third generally with 5, fourth with 4. Spiral lirae unequal in width, groups of fine threads being often separated by groups of coarser ones, those on siphonal rostrum rather uniform in strength. Collabral threads ma\ be pliculate in places, generally rendering intervals between spiral lirae punc- ticulate and the spirals themselves somewhat scabrous. Off white, with a thin yellowish-brown periostracum. Protoconch bulbous, of about IV2 whorls, first whorl strongl) tilted, smooth, last half whorl with uneven, pro- socline axial riblets; maximum breadth 3.0-4.8 mm, height 3.1-4.4 mm (ratio breadth/height 0.87-1.13). Maximum dimensions 102.0 x 39.6 mm (R. Le Maitre collection). Holotype (juvenile) 49.0 x 22.1 mm. Oper- culum oblanceolate with rather obtuse apex, moderately thick with coarse growth lines; color amber-brown, with darker outer edge. Distribution: Agulhas Bank, Type material: Holotype in the Natural History Muse- um, London, accession number 1879.2.26.116, from Lombe-Taylor collection. Other material examined: Agulhas Bank [Natal Museum (NM) D3482: R. Le Maitre, also several specimens in colln Le Maitre]; off Cape St. Blaize area, ex pisce (NM B1087; R. Le Maitre, juvenile); off Storms River mouth, 69 fathoms [= 126 m], in mud, a juvenile shell (NM S4953: C. Marais); same data, 64 fathoms [= 117 m], a broken juvenile (NM S5646; C. Marais); "Nacala harbor, dredged sand", empty shell (NM K1526: F. Steiner, don. J. H. McLean). Remarks: I have seen no adults of this species with an undamaged lip, and only two specimens (both dead ju- veniles of about 3 teleoconch whorls) bear a precise lo- cality. Its exact distribution on the Agulhas Bank, both geographic and bathymetric, consequently remains to be established. Fasciolaria thersites shows much variation in size of protoconch and extent of axial ribbing on the teleoconch; too little material is available to judge whether this re- flects geographic, individual or bathymetric variation. The smallest protoconchs measured occur in the holotype and in the specimen purportedK' from Nacala (although it is more or less worn in both examples); in the latter individual the axial ribs are unusually strong on later whorls. Similar variability in protoconch size was re- ported for F. schotvieni Strebel, 1912, by Kilburn (1974). In appearance F. thersites is most similar to F. wattersae Kilburn, 1974, an uncommon species which inhabits the Mossel Bay-Algoa Bay area at depths of about 100-120 m. In F. wattersae (Fig. 7) the axial ribs are never au- riculate, the spiral sculpture is markedly coarser, and pliculate axial striae are totally absent. Although appar- ently inhabiting similar depths, F. thersites is a much thinner-shelled species. Among the species of Fasciolaria endemic to South Africa, the operculum in F. thersites is unique in its amber-brown color, this being dark brown in the other species. ACKNOWLEDGMENTS I wish to thank Ms. Kathie Way of the Natural History Museum, London, for the loan of the holotype of Tiir- binella thersites. Messrs. R. Le Maitre and C. Marais for providing material from the Agulhas Bank, and Dr. J. H. McLean for donating the Steiner specimen to the Natal Museum. Drs. D. G. Herbert and B. R. Stuckenberg read the manuscript, and Mrs. Linda Davis prepared the line drawing. LITERATURE CITED Bacci, G. 1947. Le capsule ovigere di Cohimbella rustica (L.) e di Fasciolaria lignaria (L.). Bolletino Zoologico 14; 75- 81. Barnard, K. H. 1958. Contribution to the knowledge of South African marine Mollusca. Part I. Gastropoda: Prosobran- chiata: Toxoglossa. ,'\nnals of the South African Museum 44(4):73-163, figs. 1-30, pi. 1. D'Asaro, C. N. 1970a. Egg capsules of some prosobranchs from the Pacific coast of Panama. Veliger 13:37-43, D'Asaro, C. N, 1970b, Egg capsules of prosobranch mollusks from South Florida and the Bahamas and notes on spawn- ing in the laboratory. Bulletin of Marine Science. 20:414- 440. Gohar, H. A. F. & A. M Eisawy. 1967. The egg masses and development of five rachiglossan prosobranchs from the Bed Sea Publications of the Marine Biological Station of Ghardaqa. 14:215-268. Kilburn, R. N. 1974, Taxonomic notes on South African ma- rine Mollusca (3): Gastropoda, Prosobranchia, with de- scriptions of new taxa of Naticidae, Fasciolariidae, Ma- gilidae, \'olutomitridae and Turridae, Annals of the Natal Museum 22(1): 187-220, Kilburn, R.N. & E. Rippey, 1982. Sea Shells of Southern Africa. Johannesburg: Macmillan. Kobelt, W. 1844-1876. Turbinella und Fasciolaria. In Kster, H. C. & Kobelt, W., Systematisches Conchylien-Cabinet von Martini und Chemnitz, 2nd Edition, 3(3a):l-164, pis, 1-32, Melvill, J, C, 1891. An historical account of the genus La^irus (Montfort) and its dependencies, with descriptions of elev- en new species, and a catalogue of Latirus and Peristernia. Memoirs and Proceedings of the Manchester Literar\ and Philosophic Society [4] 4:365-411. Figures 3-7. Fasciolaria thersites (Reeve, 1847) and F. wattersae Kilburn, 1974 Fasciolaria thersites: 3. Juvenile holotype of Turbinella thersites Reeve, 1847, Natural History Museum, London, 1879 2.26.116, 49.0 x 22 1 mm. 4. Immature example from Agulhas Bank, 78.9 x33,6 mm, in R, Le Maitre colln, 5, Specimen supposedly from Nacala Bay, NM K1526, length 76,2 mm. 6. Adult example, Agulhas Bank, depth unknown, NM D3482, length 100,2 mm, outer lip damaged. 7. Fasciolaria wattersae Kilburn, 1974, S. E. of Mossel Bay, 119 m, NM E5898, 98.7 x32.3 mm. Page 28 THE NAUTILUS, Vol. 107, No. 1 Reeve, L. A. 1847. Monograph of the genus Turbinella. Con- chologia Iconica -1. London: Reeve Richards, D. 1981. South African Shells. A collector's guide. Cape Town: Struik, Shepherd, S. A. & I. M. Thomas. 1989. Marine Invertebrates of southern Australia. Part II. .Adelaide: S. Australian Gov- ernment Printing Division. Tomlin, J. R Le B. 1925. Reports on the marine Mollusca in the collections of the South African Museum. I. Turritel- lidae. .Annals of the South .\frican Museum 25:309-316. Tryon, G. W. 1881. Tritonidae, Fusidae, Buccinidae. In: Manual of Conchology 3. Philadelphia, privately pub- lished. Yen, T. C. 1942. A review of Chinese gastropods in the British Museum. Proceedings of the Malacological Society of Lon- don. 24:170-289, pis. 11-28. THE NAUTILUS 107(1 ):29-32. 1993 Page 29 The Rediscovery, Morphology, and Identity of Conus emersoni Hanna, 1963 John K. Tucker llliiKiis Natural Histur\ Survey LTRMP P.O. Bo.\ 36S West Alton, MO 63386, USA James H. McLean Natural History of Los Angeles County 900 E.xposition Boulevard Los Angeles, CA 90007, USA ABSTRACT Conus emersoni is reported from 310 ni off Isla Sanla Maria (Floreana), Galapagos Islands, the first record subsequent to its original description. The species is redescribed from eight ad- ditional specimens; its operculum, raduia, and periostracum are described for the first time. Affinity of the species to C. teramachii Kuroda, 1956, is discussed. Key words: Conidae, Contis emersoni. Galapagos Islands, rad- uia. INTRODUCTION Subsequent to its description 29 years ago (Hanna, 1963) from two dead, faded and poorly preserved specimens, Conus emersoni has remained an enigmatic member of an otherwise extensively studied genus in the Eastern Pacific region. Even as he proposed the species, Hanna (1963) suggested that it might be conspecific with the Indo-West Pacific species C. australis Holten, 1802, whereas Walls (1979) placed it as a possible synonym of another Indo-West Pacific species, C. orbigniji Audouin, 1831. Keen (1971) treated the species as valid but made no comments on its relationships. Finally. Coomans et al. (1986) considered it to be a tentatively valid species but noted that it could be based on fossil material. Here we report upon eight recently collected speci- mens from moderately deep water at the Galapagos Is- lands, two of which are in the collections of the Natural History Museum of Los Angeles County and six in the American Museum of Natural History. All were provided by Andre and Jacqueline DeRoy, residents of the Ga- lapagos Islands. These specimens extend the distribution from Cabo San Lucas, Baja California Sur, to the Ga- lapagos Islands, Ecuador, and allow a redescription of the species with the first report on the morphology of the radular tooth, the operculum, and the periostracum. Abbreviations of museums mentioned in the text: AMNH, American Museum of Natural History, New York; CAS, California Academy of Sciences, San Fran- cisco; LACM, Natural History Museum of Los Angeles County. SYSTEMATICS Conus emersoni Hanna, 1963 (figures 1-13) Contis emersoni Hanna, 1963;25, pi. 1, fig. 2; Walls, 1979:776 [with unnumbered figure of CAS paratypej; Coomans, Moolenbeek & Wils, 1986:11-4, fig. 718 [holotype, 2 views]. Conus (Asprella) emersoni. Keen, 1971:663. fig. 1497 [holo- type]. Diagnosis: Spire scalariform throughout its length, shoulder angle retaining square nodules; sculpture on final whorl of numerous, closely spaced, shallow sulci; posterior notch shallow; whorl tops sculptured by one to two cords that fade in whorl three to be replaced by numerous fine striae. Description: Shell elongate-conical, whorl sides flat to very slightly convex; shoulder angular. Anterior end not deflected dorsally. Sculpture of numerous (30-50) closely spaced, shallow sulci, most pronounced at anterior end and fading in intensity towards shoulder. Color pattern variable, consisting of three irregularly developed bands, at shoulder, in area just posterior to midbody area, and in area just anterior to midbody. Bands variously inter- rupted and scalloped, producing reticulate or blotched pattern; bands separated by areas with rows of spiral dashes or longitudinal reticulations. Dashes and reticu- lations may be quite pronounced (holotype) or lacking (Galapagan specimens). Anterior end marked by spirally elongated blotches or lines. Color markings in fresh spec- imens medium brown, fading to light brown in dead specimens. Spire very slightly convex in profile, scalariform throughout and moderately elevated, carinate, carina interrupted by numerous square nodules, 30-40 per whorl; nodules fading in whorl six although carina may remain pronounced. Protoconch unknown (not intact in any of the specimens). One or two spiral cords on first two or three teleoconch whorls, replaced on later whorls by numerous, exceedingly fine striae. Whorl tops colored by variably developed markings, matching those of body whorl, between nodules or crossing entire width of whorl Page 30 THE NAUTILUS, Vol, 107, No. 1 Figure§ 1-7. Conus emcrsurii Hanna, 1963 Figures 1-3. Type material dredged off Los Fraiies, Baja California, depth 549 111 L Holotype, AMNH 1052n, length 43 0 mm, faded, dead-collected specimen, lacking periostracum. 2, 3. Paratype, CAS 12405, length 49.0 mm, subfossil specimen (surface gray), showing naticid bore hole. Figures 4-7. Newly reported specimens dredged off Isla Santa Maria (Floreana), Galapagos Islands, Ecuador, depth 310 m. 4. AMNH 248261, length 46.2 mm, dead- collected specimen. 5. LACM 146906a, length 33.3 mm, live-collected specimen with periostracum intact. This specimen was used for opercular and radular illustration. 6. 7. LAC;M 146906b, length 34 0 mm, dead-collected specimen with periostracum removed to show color pattern. top. Whorl tops slightly but distinctly concave in cross section. Posterior notch shallow and C-shaped. Aperture narrow, white inside e.xcept where exterior coloration shows through near lip. Interior constrictions and aperturai flanges absent or at least not developed in available specimens. Periostracum thick, dark brown, markedly pilose both on body and spire. Extremely Bne hairlike extensions of periostracum not organized into any obvious pattern. These projections are apparently easily worn off, as they are pronounced on the dorsal side of one live-collected specimen (figure 5) but are not obvious on the ventral side of the same specimen. Dimensions (.see table I): The known specimens range in length from 23.0 mm to the 49.0 mm length of the paratype (figures 2, 3). The ratio of width to length ranges from 0.43 to 0.50 (table 1 ). The largest Galapagan spec- imen (figure 4) is 46.2 mm in length. All specimens have the protoconch eroded, the spire tips filled by secondary shell deposition. Operculuni illustrated in figure 8 is 27% of aperturai length, weakly serrate on outer edge. Kadular tooth (figures 9-13) small, 285 jum in lengtli; massive base with pronounced basal spur; waist located on anterior half of tooth, with small spine on posterior border in same plane as basal spur. Tooth opening rather long, extending about one-cjuarter the length. Tip w ith small barb opposite short blade. Distribution: The holotype and one paratype were col- lected oft Los Fraiies, Cape San Lucas, Baja California in 550 m (given originally as 300 fathoms by Hanna, 1963). The eight newly discovered specimens were dredged in 310 m (170 fathoms) off Isla Santa Maria (Floreana), Galapagos Islands, 7 May 1979 by Andre and Jacqueline DeRoy (table 1). DISCUSSION The rediscovery of Conus emersoni in the CJalapagos Islands should put to rest speculation that this species is a synonym of either Conus australis (suggested by Han- na, 1963) or C. orbignyi (suggested by Walls, 1979). Each of these Indo-Pacific species is unlike C. emersoni in having persistent sulci on the body whorl along with persistent cords on the spire \\ horl tops. C'onus emersoni differs in having the sulci strong only anteriorly and in having fine spiral striae on the whorl tops. Radular dif- ferences are that the radular tooth of C. orbignyi has three anterior barbs (Kilburii, 1973: fig 6), whereas that J. K. Tucker and J. H. McLean, 1993 Page 31 .-H* W^ » m0 10 11 Figures 8-13. Conus emcrsoni Hanna, 1963 8. Operculum, same specimen as figure 5, length 7 3 mm, nucleus lost. Figures 9-13. SEM views of single radular toolli, length 285 m'". from same specimen as in figure .5. SEM photos by H. Chaney 9. Oblique anterior view showing apical surface, with pronounced basal spur and small spine on posterior border of waist. 10. Enlarged view of apical surface, with small barb and enrolled blade. 11-13. Three lateral views, showing differing axial rotation of tooth of C. emersoni has but two. The radular tooth of C. australis has a serrate shaft along with an enlarged cusp at the posterior end of the row of serrations (A. J. Kohn, radular slide collection); neither feature is present on the radular tooth of C. emersoni. The scalariform spire and color markings of Conus emersoni have a superficial resemblance to that of the Panamic species C. emarginatus Reeve, 1844 (a species often misidentified as C. recurvus Broderip, 1833). Dif- ferences are that C. emarginatus has a deep posterior notch (rather than shallow notch) and a smooth (rather than pilose) periostracum. The radula of C. emarginatus (Nybakken, 1970; fig. 5, as Conus recurvus) has three barbs anteriorly, like that of C. orbignyi, rather than C. emersoni, in which there are two barbs. The scarcity of C. emersoni may be due to the great depth at which it occurs and the paucity of sampling at depths below 300 m. It is possible that the specimens from the Galapagos Islands are specifically or subspecifically distinct. They differ in color pattern (that of the holotype being more intricate) and in spire profile (that of the holotype ap- pearing to be more acute). However, such intraspecific differences in color pattern are not uncommon in Conus. In fact, the faintly indicated color pattern of the paratype that shows on the better preserved dorsal surface (figure 3) is more similar to that of the Galapagan specimens (figures 5-7) than to that of the holotype (figure 1). The ground color of the paratype has a gray cast that is suggestive of fossil or subfossil condition. The holotype is in fresher condition, although it retains no periostra- cum; it shows evidence of growth damage to the shoulder at a stage two whorls above the termination of the lip, which appears to have altered the profile of the final two whorls, resulting in a greater downward slope to the spire and the loss of the nodules in the final two whorls. The question as to whether the two widely disjunct records represent the same species can only be answered after additional specimens from the vicinity of the type locality become known. It is our opinion, based on the specimens examined, that there is no clear evidence to suggest that the Galapagan specimens are not conspecific with the Mexican specimens. Conus emersoni is not closely similar to any other eastern Pacific conid. Shell morphology, including the Table 1 . Shell dimensions and proportions of the known spec- imens of Conus emersoni Hanna, 1963. Width/ Specimen C:ondition Length Width length AMNH 92200 (holotvpe dead 43.0 18.5 0.43 CAS 1240.5 (paratype) subfossil 49.0 22.5 0.46 LACM l-)6906a live 33.9 17.0 0.50 LAC:M 146906b dead 340 15.5 0.46 AMNH 248262 live 35.0 15.7 0.45 AMNH 248263 live 31.5 14.3 0 45 AMNH 248261 dead 46.2 22.3 0.48 AMNH 248169a live 23.0 10.8 0.47 AMNH 248169b dead 30.0 149 0.50 AMNH 248169c live 26 8 12 8 0.48 Page 32 THE NAUTILUS, Vol. 107, No. 1 14 15 Figures 14, 15. Conus teramachii Kuroda, 1956 14. Speci- men with growth scar producing lowered shoulder angle of final whorl, periostracum removed. Off NE coast Taiwan, trawled, depth unknown. LACM 68994, length 79 8 mm. 15. Operculum with strongly serrate edge, nucleus lost, trawled off Taiwan. Specimen in H. Chaney collection, length 19.1 mm. shoulder carina with square nodules, is similar to that of C. teramachii Kuroda, 1956 (figure 14), a species re- ported from similar depths off Japan and Taiwan (Ku- roda, 1956; Walls, 1979). Our illustrations of the radular tooth of C. emersoni (figures 9-13) are comparable to those of Azuma (1961: fig. 11) for C. teramachii (as C. petricosus Azuma, 1961). In addition, C. teramachii also has a strongly serrate operculum (figure 15), much more pronounced than that of C. emersoni (figure 8). The character state of the serrate operculum was stressed in the original diagnosis of the subgenus Profundiconus Kuroda, 1956 — type species Chehjconus {Profundico- nus) profundorum Kuroda, 1956 [= Conus smirna Bartsch and Rehder, 1943]. The type species of Profun- diconus also has square nodules on the shoulder carina, but these are apparent only in young stages. We refrain, however, from further treatment of the subgeneric allocation of the species under discussion be- cause comparison to other available generic level taxa is beyond the scope of this paper. We recognize that a generic level classification of Conidae needs to be based on all the recognized species, including fossils, at a min- imum treating characters that include adult and juvenile shell morphology, and, for the living species, the oper- culum, and radula. ACKNOWLEDGMENTS We thank Alan J. Kohn of the L'niversity of Washington for allowing one of us (JKT) to study his radular slide of C. australis. William K. Emerson and Walter E. Sage of the American Museum of Natural History made the ho- lotype and other specimens of C. emersoni available for study. Terrence M. Gosliner of the California Academy of Sciences arranged for loan of the paratype specimen. We thank Henry W. Chaney of the Santa Barbara Mu- seum of Natural History for the SEM photos of the rad- ular tooth of C. emersoni and the photograph of the operculum of C. teramachii. We are grateful to Henry W, Chaney, William K. Emerson, Alan J. Kohn and Walter E. Sage and two anonymous referees for reading the manuscript and offering helpful suggestions. Finally, and most importantly, we thank Jacqueline and Andre DeRoy for providing the specimens of C. emersoni that form the basis for this paper. LITERATURE CITED Azuma, M. 1961. Descriptions of six new species of Japanese marine Gastropoda. Venus 21(3):296-303. Coomans, H E., R. G. Moolenbeek, and E. Wils. 1986. Al- phabetical revision of the (sub)species in recent Conidae. 9. ehraeus to extraordinaritis with the description of Conus elegans remalhoi. nov. subspecies. Basteria50;93-1.59, figs. 667-760. Hanna, G. D. 1963. West American mollusks ot the genus Conus — II. California Academy of Sciences, Occasional Papers, no. 35, 103 p., 11 pis. Keen, A. M. 1971. Sea shells of tropical West America, second edition. Stanford University Press, Stanford, C.-^, 1064 p. Kilburn, R. N. 1973 Notes on some benthic Mollusca from Natal and Mozambique with descriptions of new species and subspecies of Calliostoma, SolarieUa. Latiaxis. Bab- ylonia, Fusinus. Bathytoma and Conus. Annals of the Natal Museum 2I(3):557-578, figs. 1-17. Kuroda, T. 1956. New species of the Conidae (Gastropoda) from Japan. Venus 19(1): 1-13, 1 pi. Nybakken, J. 1970. Radular anatomy and systematics of the West .'\merican Conidae (Mollusca, Gastropoda). Ameri- can Museum Novitates 2414:1-29, figs. 1-45. Walls, J. G. 1979. Cone shells, a synopsis of the living Conidae. T. F. H. Publications, Inc., Neptune City, NJ, 1011 p. THE NAUTILUS 107(l):33-42, 1993 Page 33 The Benthic Mollusk Faunas of Two Contrasting Reef Paleosubenvironments: Falmouth Formation (late Pleistocene, Last Interglacial), Jamaica Stephen K. Donovan Department iil Ch'iiIoi;\ University of the West Indies Mona, Kingston 7 JAMAICA D.T.J. Littlewood' Haskin Shellfish Heseareh Laboratory Institute of Marine and Coastal Sciences Cook College/ NJ A ES P.O. Box B-8 Port Norris, New Jersey 0S349 l!SA ABSTRACT The last interglacial (about 125,000 years old) raised reefs of the Falmouth Formation of Jamaica contain a diverse fauna of benthic mollusks. Large collections of mollusks ha\e been made from two contrasting reef localities and paleosubenvi- ronments. Calcareous muds of lagoonal origin were well-ce- mented, biassing collecting tow ards larger individuals, whereas poorlv-cemented santly sediments of a coral framework were friable and thus amenable to bulk sampling, yielding numerous micromoUusks. Identification of over 3,000 bivalve, gastropod and scaphopod specimens shows the molluscan faunas of each environment to be dominated by different species. The coral framework fauna is numerically dominated b\ the microgas- tropod Caecitm piilchcllinu Stimpson, 1851, other epifaunal gastropods and arks, while Bulla .striata Bruguiere, 1792 and ceriths dominate the lagoonal sediments. Only Cerithium al- gicola C.B. ,^dams, 1848, forms more than 2"; of the fauna at both localities. 17% of the extant species (27% of the common species) of shallow water benthic mollusks (excluding chitons) from Jamaica have been identified from these two localities. Key words Late Pleistocene; benthic reef mollusks; Jamaica. INTRODUCTION The present paper is a preliminary report of the mol- luscan faunas from the late Pleistocene (the last inter- glacial or Sangamonian) Falmouth Formation (125,000 years B.P.) of northern Jamaica. The mollusks of the Falmouth Formation are important for two principal reasons. Firstly, the Neogene and Quaternary fossilif- erous deposits of Jamaica contain a diverse fauna of benthic mollusks that have hitherto been largely ignored ' Current address; Department of Palaeontology, British Mu- seum (Natural Historv), Cromwell Road, London. S\V7 5BD, ENGLAND by systematists. Indeed, only the famous Pliocene Bow- den shell bed in southeastern Jamaica (Woodring, 1925, 1928) and the fauna of the Early Pleistocene Manchi- oneal Formation (Trechmann, 1930) have so far received monographic treatment. This lack of published infor- mation does not reflect the true diversity of the mollusk faunas through this interval and ma\ thus prove mis- leading to workers not familiar with the geology of the island. For example, Petuch (1988:49) considered that there are few exposures of Miocene age preserved within the Caribbean and South American region, and did not include Jamaica in his list of productive sites, yet in a preliminary survey Jung (1972) listed members of 19 genera from the extensive deposits of the Newport For- mation (Lower and Middle Miocene) of the island. Further, molluscan faunas from last interglacial reefs (and other Pleistocene marine environments; Valentine, 1989) can be a valuable adjunct to studies of extant as- semblages from the shallow benthos (but see cautionary comments in Johnson, 1960). The Pleistocene raised reefs of the Caribbean region are recognised to include large faunas of fossil mollusks (see, for example, Cerridwen & Jones, 1991). Unlike those of certain other areas (such as Aldabra Atoll; Taylor, 1978), Caribbean mollusk fau- nas have apparently changed little since the last inter- glacial (Petuch, 1988:119), an observation that may have important implications for the study of faunal associa- tions in Recent reefs and associated environments. Mol- lusk faunas in various biofacies w ithin Pleistocene raised reefs are generalK easy to sample, outcropping adjacent to the coast on many Caribbean islands, thus permitting large collections to be made for systematic and paleo- ecological analysis. Geology and geological nomenclalure: The Falmouth Formation in the Discovery Bay-Rio Bueno Harbour re- gion of Jamaica's northern coast (figures 1-3) rests un- conformably on the dolomitized Plio-Pleistocene reef (Land, 1991) of the Hopegate Fonnation. Lithofacies Page 34 THE NAUTILUS. Vol. 107, No. 1 2 1^ ^ ^ 0 km 1 ' ^-— ^^=^^8=- ^^2 \ ^ ^ 1^ ^^ % ■^1 DB Jy ^"¥1 i ,54-70 ^) Ftl W/!?\ TerrigencHJS grainstcxie facies Mi::::iij Mdluscan algal gralnstone facies pt'«';1 Mdkjscan biomlcritic wackestone fades f n Biomicritic packstone facies I I Coralline boundstone facies Hopegate Formation Figures 1-3. Maps ol slud) arta;. 1 Outline map uf JamaiLa, showing the location of the Discovery Bay (DB) region with respect to Kingston (K), Montego Bay (MB) and Ocho Rios (OR) 2 Topographic map of the Discovery Bay-Rio Bueno Harbour (RBH) area, showing the positions of localities 1 and 2. Contours in m. 3 Geological map of the Discover) Bay-Rio Bueno Harbour area (redrawn after Larson. 19So: figure 6). showing the relative distributions of the major sedimentary facies of the Falmouth Formation in relation to the underK ing Hopegate Formation (key right). within the Falmouth Formation reef (Larson. 1983) con- tain associated distinct faunal assemblages which have been recognised from the evidence of corals (Larson. 1983: Liddell et ai. 1984) and echinoids (Gordon & Donovan. 1992). History of research: Hill (1899) named the Falmouth Formation, and recognized the essential elements of the mollusk fauna to comprise Lucina, Cardium. Area. So- len and Bulla. Hill (1899. 155) considered this fauna to be worthv- of careful stud\ b\ paleontologists, an invi- tation that has not been taken up in over 90 \ ears. Rob- inson (1958) noted Area spp. from the elevated reef at east Rio Bueno Harbour (localitv 1 herein), and Chione cancellata (Linnaeus. 1767) and Bulla cf. striata Bru- guiere. 1792. from the lagoonal facies east and west of Falmouth. Parish of Trelawny (probabh lithologicalK- similar to locality 2 herein). In 1960. Robinson considered the Falmouth Formation in the Parish of St. .Ann to include the moUusks Lueina. Codakia. Chione and Bul- la, and later commented that this rich fauna is identical with the living fauna of the Jamaican coastal waters (Robinson. 19&3:52). Recent studies of the fossil scapho- pods of Jamaica (Donovan and Littlewood. 1989; Don- ovan, 1990) ha\e recognised Dentalium sp. cf. D. (An- atalis) antillarum d'Orbigny, 1842, and D. (Graptacme) semistriolatum Guilding. 18;34. from the east Rio Bueno Harbour locality. The specimens discussed herein are deposited in the Department of Palaeontology, British Museum (Natural History). For uncertainties in our taxonomic assignments (particularly for poorly preserved or fragmentary spec- imens), we have used question marks thus: Agenus ?spe- cies signifies that we are uncertain of the specific assig- nation: ? Agenus aspeeies indicates that some doubt exists concerning the generic land. thus, also specific! assign- ment. This differs in detail, but not in spirit, from the recommendations of Bengston (1988). MATERIALS AND METHODS The Falmouth Formation of the Discovery Bay-Rio Bueno Harbour area has been subdivided into five principal reef al subenv ironments by Larson (1983; figure 3 herein). Large collections of benthic mollusks were made from two contrasting paleosubenvironments ^localities 1 and 2, described below) within this continuum. These par- ticular subenvironments and localities were chosen be- cause of their large mollusk faunas and easy access. The mollusk collections from each locality were made over a period of 22 months, from .April 19SS imtil February 1990. Both collections are time-averaged, having been collected from float, the rock face and as bulk samples from throughout 2-4 m of section in each case, as bedding planes and other obvious time marker horizons are not apparent. Howev er. the \ alue of collecting mollusks from discrete beds (e\ en when preserved) within reefs is de- batable, as at an\ one time the fauna w ill include shallow and deep infaunal. sediment surface epifaunal. and (in the case of the three-dimensional coral framework at localitv 1) epifaunal taxa living well above the soft sed- iment surface. Specimens collected in the field were generally at least S. K. Donovan and D. T. J. Littlewood, 1993 Page 35 .y .■^. J'-^ ■->^ ^ , x# ^ ^:.'^' -<; :^<' ^-ii-rv-^is*; ' m a. Figures 4-7. Field photographs of molhisks of the Falmouth Formation 4-. Stromhus gigas Linnaeus, 1758, at locahty 1; 0.3 x. 5-7. LocalitN' 2, 5, MainK Bulla striata Bruguiere, 1792; 0 6 x. 6. General view of weathered surface showing abundant (mainly molluscan) shell hash; 0.6 x 7. MainK Bulla striata Bruguiere, 1792, associated with an articulated shell (arrowed) of Chione paphia (Linnaeus, 1767); 0.4 x. 5 mm in maximum dimension and even shells of this size were easily overlooked without repeated scanning of a limited area at a time. It proved much easier to detect small specimens during laboratory analysis using a Wild binocular microscope. Preliminary field identi- fications were checked in the laboratory using standard taxonomic references, including Abbott (1954, 1958, 1974), de Jong and Coomans (1988), Fisher (1988), Hum- frey (1975), Morris (1987), Vaught (1989), and Warmke and Abbott (1961). Total numbers of whole shells and (for bivaUes) disarticulated valves are tabulated \\'hole shell equivalents (discussed in Cerridwen and Jones, 1991 ) were used in calculating relative proportions of moUusks for graphical analysis. Locality 1 (Locality 7 of Liddell et a/., 1984; locality 3 of Donovan 6 Gordon, 1989). East side of Rio Bueno Harbour, Parish of St. Ann, with the Falmouth Formation exposed as a line of low cliffs (GR 394 572). At this locality the un- conformable contact with the underlying, dolomitised Hope Gate Formation (figure 3) formed the hard sub- strate for Falmouth Formation reef growth. Previous interpretations have considered this locality to represent either a patch reef in the back reef lagoon within the molluscan biomicritic wackestone facies sensii lato (Lar- son, 1983; figure 3 herein) or perhaps part of the shallow- fore reef (Robinson, 1958; Liddell et a!., 1984). Mollusk-rich sediments at this locality occur beneath an altered caliche cap (Land & Epstein, 1970) in a frame- work of shallow-water corals, particularly Porites spp. and Acropora spp. (Liddell et al., 1984: 77). The matrix between corals is a friable, weakly-cemented carbonate sand which includes a diverse benthic fauna, including echinoid (Gordon & Donovan, 1992) and ophiuroid (Donovan et al., in press) ossicles, crabs (Morris, in press), disarticulated chiton valves (Donovan et al., research in progress), scaphopods (Donovan & Littlewood, 1989; Donovan, 1990), gastropods and bivalves. Due to the friable nature of the sediment, two collecting methods were used. Firstly, mollusk specimens that were weath- ering out of the exposed face (figure 4) were removed manuallv. The fauna collected by this method was dom- inated by bivalves and gastropods, with three scaphopod shells. Secondly , bulk samples were collected by the bag- ful. In the laboratory these samples were emptied into shallow trays, dried in an oven overnight, then washed through a nest of sieves using cold water and redried. Sieved aliquots were then picked by ey e (coarse fractions) or under the binocular microscope (fine fractions: frac- tions finer than coarse sand were not picked). The fauna collected by this method was dominated by bivalves, gastropods and (in the finer fractions examined) chiton valves, but no further scaphopods were found. A total of 1669 benthic mollusks (excluding chiton valves; Donovan et al., research in progress) have been identified from this locality (table 1). Locality 2 (Locality 5 of Liddell et al., 1984). This is an outcrop of hard, fine-grained, well-lithified, but occasionally chalky, limestone exposed along the shoreline as a low terrace east of the Discovery Bay Marine Laboratory, Parish of St. Ann (GR 405 569). This lithology has been interpreted as having been formed in a back reef environment (Lid- dell et al., 1984:76), presumably a lagoon, forming part of the molluscan biomicritic wackestone facies sensu stricto of Larson (1983; figure 3 herein). Page 36 THE NAUTILUS, Vol. 107. No. 1 Table 1. Mollusk fauna collected from locality 1, east of Rio Bueno Harbour. Key: * = includes 11 juveniles, plus 2 shells not collected; ** = with apparent growth deformity; + = vaKe incomplete; ++ = probably mostly Area iiiibricala Bruguiere; S = shells or identifiable shell fragments; \' = valves; LV = left valve; RV = right valve; O = operculum; J = juvenile shells. Gastropods Diodora listen (d'Orbigny, 1853) 25S Diodora '^caijencnsis (Lamarck, 1822) 12S Diodora '■'mintitn (Lamarck, 1822) IS Fi.ssiirclla harhadcnsis (Gmelin, 1791) IS Acinuea ?(inlillarinn (Sowvrhy, 1831) IS Acmaea pustidata (Helbling, 1779) 60S Cittarium pica (Linnaeus, 1758) ,3S Tegula fasciata (Born, 1778) ,3S Astraea ?caelata (Gmelin, 1791) lO Astraea tccta (Solander, 1786) 4S Nerita versicolor Gmelin, 1791 IS Nerita tessellala Gmelin, 1791 2S Nerita sp. indet, j^S Neritina virginea (Linnaeus, 1758) IS Neritina punctulata Lamarck. 1816 2S Littorina ?nebulosa Lamarck. 1822 2S Alvania ?aberrans C. B. Adams, 1850 IS Heliacus injundibulijormis (Gmelin. 1791) IS Petaloconchus irregularis (d'Orbigny, 1842) 2S Petaloconchus erecius Dall, 1888 IS Petaloconchus mcgintyi Olsson and Harbison, 1953 2S Serpulorbis decussata (CmeYm, 1791) 5S Serpulorlris ?decussata (Gmelin, 1791) IS Serpulorbis ?riisei (Morch, 1862) IS Caecum ?insularum Moore, 1970 IS Caecum plicatum Carpenter, 1858 IS Caecum pulchcllum Stimpson, 1851 657S Modulus modulus (Linnaeus, 1758) 13S Cerithium ''litteratum (Born, 1778) 7S Cerithium guinaicum Philippi, 1849 3S Cerithium ?eburneum Bruguiere, 1792 2S Cerithium algicola C. B. Adams, 1848 92S Cerithium sp. 3S Cerithium sp. indet. 3S Hipponix antiquatus (lAnna.eus, 1767) ,3,3S Strombus gigas Linnaeus, 1758 * 36S Cypraea zebra Linnaeus, 1758 Ij Cypraea cinerea Gmelin, 1791 2S Polinices lacteus (Guilding, 1834) 9S Morum oniscus (Linnaeus, 1767) 3S Cymatium 'pileare' (Linnaeus, 1767) IJ ?Cymatium nicobariciim (Roding. 1798) IJ Cymatium ?parthenopeum (von Salis, 1793) 5S Cymatium sp. sensu lata H ^Cymatium sp. \\ Bursa cubaniarui (d'Orbigny, 1842) IJ Columbclla mercatoria (Linnaeus, 1758) 81S F.ngoniophos 'hmicinctus (Say, 1826) IS ?Melongena mclongena (Linnaeus, 1881) 2S Fasciolaria tulipa (Linnaeus, 1758) 2S Latirus ?hrevicaudatus Reeve, 1847 IS Leucozonia nassa (Gmelin, 1791) 17S Mitra barbadensis (Gmelin, 1791) 6S Hyalina avena (Kiener, 1834) IS Conus sp. or spp. 3S ?Mangelia quadrdim-ata (C. B. Adam.s, 1850) 2S Bulla striata Bruguiere, 1792 2S Odostomia sp. or Pyramldella sp. IS Cheilia equestris (Linnaeus, 1758) IS Total 1,1.3() S. K. Donovan and D. T, J. Littlewood, 1993 Page 37 Table 1. ('diitiniird Bivalves Area ?zebra Swainson. 18S3 Area imhricuta Bruguiere, 1789 Rarlxititi Candida (Helbling, 1779) Barlmtia ?cancellaria (Lamarck, 1819) Barhatia teiwra (C B. Adams, 1845) Anadara sp .■\rks iiuiet. + + Brachidonti's cxustus (Linnaeus, 1758) Smoolli '•'Botida fuaca (Gmelin, 1791) iJthophaga ''nigra (d'Orbigny, 1842) hognomvn '•'alalua (CJmelin, 1791) hognomon radiatus (Anton, 1839) Isognonwn sp, indet Lima scabra (Born, 1778) Ostreola equestris Say, 1834 Codakia orbicularis (Linnaeus, 1758) Codakia ?orbiculata (Montagu, 1808) ''Codakia sp, Chama maccrophylla Gmelin. 1791 Chama '''maccrophylla Gmelin, 1791 Pseudochatua radians (Lamarck, 1819) Americardia media (Linnaeus, 1758) Americardia guppyi (Thiele, 1910) Chione cancellata (Linnaeus, 1767) Pcrighjpta listeri (Gray, 1838) Tellina listeri Boding, 1798 Tellina sp- cf, Tellina ''tnera Sa\, 1834 Tellins indet. Tellinacean sp, indet. A Tellinacean sp, indet B ?Tellinacean sp, indet.** Areopagia Jausta (Pulteney, 1799) ?Macoma pseudomera Dall and Simpson, 1901 ?Macoma sp. PLucinid or ?pectinacean indet. ?Mactracean indet, ?Periploma papyralium Say, 1822 Bivalve sp indet. Scaphopods Dentalium sp cf. Dentalium (Anatalis) antillanim d'Orbigny, 1842 Dentalium (Graptaeme) semistriolatum CJuilding, 1834 9V + 1V + 1V + 1V + 1V + Totals 4LV 77LV 25LV ILV 36LV ILV 32LV 4LV ILV lOLV 4LV ILV 2LV ILV 9LV 4LV ILV ILV ILV 2LV 2LV ILV 3LV 28LV 2LV 61RV 37 RV 4RV 24RV 29RV 2RV 5RV 2RV IRV 13RV 6RV IRV IRV IRV IRV 5RV IRV IRV 35RV IRV IRV IRV IRV IRV 502V 19S IS IS 2S IS IS 3S 6S Total 33S IS 2S 3S Because of the well-lithified nature of much of this limestone, it was necessary to use a hammer and chisel to remove all specimens seen in the exposed faces (figures 5-7). Specimens were not apparent on all surfaces due to the vagaries of weathering, although breaking open many of these 'unfossiliferous' exposures showed that mollusks were nevertheless common. As well as collecting individual specimens in the field, large fossiliferous slabs containing dense accumulations of mollusks were taken back to the laboratory for mechanical breakdown. A total of 1365 benthic mollusks have been identified from this locality (table 2), including abundant gastropods and bi- valves, rare scaphopods, but no chitons. RESULTS AND DISCUSSION Taxonomy: Taxa, and number of specimens of each species, from localities 1 and 2 are tabulated in tables 1 and 2, respectively. It is readily apparent that both fau- nas, although approximately contemporaneous and sep- arated by a distance of only about 3 km, show consid- erable taxonomic divergence (figures 8, 9). Localitv 1 has the more diverse fauna, which is dominated numer- ically by the micromollusk Caecum pulchellutn Stimp- son, 1851, arks and epifaunal gastropods (although C. piilchelltim made only a minute contribution to biomass), whereas locality 2 is typified by the occurrence of abun- Page 38 THE NAUTILUS, Vol. 107, No. 1 Table 2. MoUusk fauna collected from locality 2, east of Discovery Bay Marine Laboratory. Key: ' C". B. .-Vdams, 1848; otherwise as for Table I = encrusting Cerithium algicola Gastropods Diodora ?cayenensis (Lamarck, 1822) Turbo canaliciilahi.s Hermann, 1781 Tricolia sp or spp. ?Smaragdia ciridis ciridenmris Maury, 1917 Rissoina aberans (C. B. .\dams, 1850) '^Rissoina sp. Planaxis lineatus (da Costa, 1778) Modulus carchcdonius (Lamarck, 1822) Modulus modtdus (Linnaeus, 1758) Cerithium algicola C. B .\dams. 1848 Cerithium eburneum Bruguiere. 1792 Cerithium litteratum Born, 1778 Cerithium spp. indet. (fragments) Janthina sp. ?\iso portoricensis Dall and Simpson, 1901 Stromhus gigas Linnaeus, 1758 Polinices lactcus (.Guilding, 1834) Satica canrena (Linnaeus, 1758) Naticid sp. indet. Cymatium "^caribbaeum Clench and Turner. 1957 Ctjmatium muricinum (Boding, 1798) Cymatium sp. indet. .\ Cymatium sp. indet. B Cymatium sp indet. C ?Thais ddtoidea (Lamarck, 1822) Columbella mercatoria (Linnaeus. 1758) Anachis obesa (C. B. .\dams. 1845) Anachis ?pretri (Duclos, 1846) Anachis sp. indet. S'assarius alhtis (Say, 1826) Fasciolaria tulipa (Linnaeus, 1758) Oliiella dealbata (Ree\e, 1850) Prttnuvi guttatum (Dill\v\n, 1817) ?\'olvarina gracilis (C B. .\dams, 1851) CoTius '^centurio Born, 1778 Conus verrucosus Hwass, 1792 ?Crassispira sp. indet. Btdia striata Bruguiere, 1792 Retusa candci (d Orbigiu, 1841) Bivalves ??Suculid sp. indet. Area imbricata Bruguiere, 1789 Barbatia cancellaria (Lamarck. 1819) .\rk indet. sp. Brachidontes citrinus (Boding. 1798) ?lsognomon sp. Pinna camea Gmelin. 1791 Pecten '^chazaliei Dautzenberg. 1900 ?Diplodonta punctata ^Sa>. 1822) Lucina sp. indet .A Lucinid sp. indet. .\ Lucinids spp. indet. ?Anodontia alba Link. 1807 Codakia costata (d"Orbigny. 1842) Codakia orbictdaris (Linnaeus, 175S) Codakia orbictdata (Montague. 180S) Dicaricella quadhsulcata (d"Orbign\ , 1842) Chama macerophulla Gmelin, 1791 Chama sp Total ILV 3RV ILV IRV 2L\" IRV ILV IRV ILV 1V + ILV^ IRV ILV 2LV IRV 2L\- 3RV 3L\ 3RV 2LV 4LV 3RV 2RV 2LV 2RV 121.V 34RV IRV 2S IS 6S IS IS IS IS 7S IS lOlS 3S 33S 58S IS 35 17S 78S IS IJ IS IS IS IS IJ IS 5S IS IS 2S IS 2S 37S 13S IS 95 IS IS 6US IS LOOS IS :i \ S. K. Donovan and D. T. J. Littlewood, 1993 Table 2. ( liiiiliinic Page 39 fTraihyairdiinu sp Americardia gupptji (Tliiilc. 1910) Aniericarilia ntcdici (Linnaeus, 1758) Lacvictirditini liHiif^iiluin (Linnaeus, 1758) '^Lacvicardiiun Itui if^dluiu (Liiuiaeus, 1758) Laevicardium '•'syhdnluuiu Dall, 1886 Chiotie ranci'llata (Liiuiaeus. 17(i7) Chionc firaniilata ((iinelin. 1791) Chionc pap}uu (Linnaeus, I7()7) Transcnnclla '•'cimradimi Dall, 1883 Transennclla ?cubaniana (d'OrliiKny, 18-42) Pitar allrida (Cmelin, 1791) Tcllina altcrnala Say, 1822 Tcllina listrri Hoding, 1798 Tcllina similts Sowerby, 1806 Tcllina sp. iiulet Apohjmelis inlastriata (Say, 1826) Tagcliis divisus (Spengler, 1794) Mactra fragilis Gnwhn, 1791 Scaphopods Cadulus sp Denialium ''anlillarum dOrliigiu, 1846 Dentalium sp, (smooth-shelled) 1\ Totals 2LV 2RV ILV 2RV (>LV 7RV 5S 3LV IRV' 1V + lOLV URV IRV IS 2LV 2RV 2S 6LV 2RV 2LV 2RV 13LV 12RV 6S 2LV 2RV 2V-I- 3LV 2RV 2S 18LV lORV 4V-I- 7LV 3RV ILV 6S 30LV 29RV 2RV 2S 328V 25S Total IS 2S IS 4S dant Bulla striata Briiguiere, 1792 and Cerithium spp with other gastropods and rarer bi\aives. Paleoecology and preservation: Taxonomic differ- ences are interpreted as being related to paieoenviron- mental control. Locality 1 included abundant hard sub- strates for the attachment of epifaunal bivalves, provided by the coral framework, as well as a sandy substrate suitable for a variety of infaunal and epifaunal ta.xa. Chitons, which generally prefer a hard substrate, are also common at this locality . In contrast, locality 2 lacks any evidence of a three-dimensional framework structure; indeed, corals were small and isolated in this paleoen- vironment. Consequently , epifaunal species are rare and the fauna is dominated by infaunal taxa, particularly B striata, but also including \arious burrowing bi\alves (habitat preferences of many of the groups tabulated herein are summarized in Cerridwen & Jones, 1991: table Table 3. .\ comparison of the taxonomic diversity of the extant Jamaican moliuscan fauna with nominal species identified here- in from the Falmouth Formation. Key: -I- = after Humfre\ (1975); * = excluding rare species; 1, 2 = localities. K\tant-I- 1 ,-> 1 + 2 Scaphopods 11 •1 3 4 :56 Gastropods 324* 55 37 80 25 558 55 37 80 14 Bivalves 163* 33 34 56 34 246 33 34 56 23 Totals 514* 141) -">- 831 140 17 3). The only species which forms greater than 2% of both faunas is Cerithium algicola C B .Adams, 1848 (figures 8, 9; tables 1, 2). The difference in diversity between the localities may be real, although it is almost certainly at least part ar- tifact. This is because of the differing degrees of lithifi- cation (and, hence, availability of specimens) seen he- tween the two collecting sites. All specimens from bulk samples from locality 1 were theoretically collected by sieving and picking. In contrast, limestones from locality 2 were not amenable to such processing, being well- lithified, and liberation of specimens was essentially a destructive process. In addition to those specimens bro- ken from the rock, there were some mollusks, particularly those with a fragile shell, that were too easily lost due to breakage while others were being remo\ed. Further, many specimens were probably ne\er seen because they were enclosed within the remaining small chips of rock that were not broken down further. Micromollusks are a common component of the fauna from locality 1. main- Iv recognized during microscopic examination of the sand-sized sieve fraction. Such a technique is not gen- erally possible with the sediment from locality 2 and, even if detected, small shells are difficult to extract and clean from the hard matrix. The total absence of chiton vaKes or Caecum shells from locality 2 may be real, but the limited diversity of micromollusks from this site sug- gests that preservational and collecting biases were im- portant factors (in particular, extant Caecum is a com- mon component of seagrass communities which presumably occurred in the paleolagoon). Thus, micro- mollusks were part of the easily collectable fauna only Page 40 THE NAUTILUS, Vol. 107, No. 1 o o a- o lOOi 75 50 25 0 a c3 H. O ^Myf0(iiyf^'^f!f^^mKy7TmmmmmjTwTw„ym^j Species Figure 8. Histogram of relative abundance of mollusk species of locality 1 (n = 1467; bivalves recalculated as whole shell equivalents). 89 species are each present as less than 1% of the total mollusk fauna. • = broken fragments of arks which are too poorly preserved for even generic identiBcation, but which are probably dominantly A. inihricala. at locality 1, although it was still possible to extract some small shells from the limestones at locality 2 (particularly robust shells such as Cadtiliis sp. and B. striata juveniles). Macromollusks from both sections are often exquisitely preserved. Whilst bivalves are normally (and chitons in- variably) disarticulated, valves often have a highly glossy appearance, such as Laevicardium laevigattim (Linnaeus, 1758) at locality 2 and Polinices lacteus (Guilding, 1834) at both localities, equivalent to that of fresh shells. Fur- ther, shell coloration and/or color banding is preserved in, for example, at least some specimens of Leucozonia nassa (Gmelin, 1791) from locality 1 and B. striata from locality 2. This indicates that the diagenetic effects lead- ing to the lithification of the limestones (particularly at locality 2) have so far had little influence on the organic molecules that are largely responsible for coloration. However, many shells are poorly preserved, often being corroded and/or abraded. Many show the influence of pre- and post-mortem boring. Pre-mortem borings by predatory gastropods (?P. lacteus) have been noted in, for example, Brachidontcs cxustus (Linnaeus, 1758) and Table 4. ('omparison ol llic extant shallow-vsater molluscan fauna of Yucatan (PJkdale, 1974) with that of the Pleistocene Falmouth Knrmation calitit's e\rln Journal of the Geological Society of Jamaica 28: 45-48. Land, L.S. and S. Epstein. 1970. Late Pleistocene diagenesis and dolomitization, north Jamaica. Sedimentologv 14: 187- 200. Larson, DC 1983. Depositional facies and diagenetic fabrics in the Late Pleistocene Falmouth Formation of Jamaica. M.S. thesis. University of Oklahoma, Norman, Oklahoma. Liddell, W.D.,S.L. Ohihorstand AG. Coates. 1984. Modern and ancient carbonate environments of Jamaica. Rosenstiel School of Marine and Atmospheric Science, University of Miami, Sedimenta 10: vii -I- 98 p. Morris, PA. (W.J. Clench, ed.). 1987. A field guide to shells of the Atlantic and Ckilf Coasts and the West Indies. Houghton Mifflin, Boston, x.wiii + 330 p. Morris, S.F. In press. Biostratigraphy of the arthropods of Jamaica. Geological Society of America Memoir 180. Parsons, KM. and C.E. Brett. 1991. Taphonomic processes and biases in modern marine environments: an actualis- tic perspective on fossil assemblage preservation. In Don- ovan, S K. (ed ). Fossilization: the processes of taphonomy Belhaven Press, London p. 22-65. Petuch, E.J. 1988. Neogene histor\ of tropical .American mol- iusks. The Coastal Education and Research F"oundation, C^harlottesville, Virginia. 217 p Robinson, E. 1958. The younger rocks of St. James and Tre- lawny. Geonotes 1: 15-17. Robinson, E. 1960. Observations on the elevated and modern reef formations of the St. Aim coast. Geonotes 3: 18-22. Robinson, E 1963. Quaternary deposits. In. Zans, V A., L.J. Chubb, H R. Versey, J B Williams, E Robinson and D L. Cooke. Synopsis of the geologv of Jamaica. Bulletin of the Geological Survey Department of Jamaica 4: 50-54. Taylor, J.D. 1978. Faunal response to the instability of reef habitats: Pleistocene molluscan assemblages of Aldabra Atoll. Palaeontology 21: 1-30. Trechmann, C T. 1930. The Manchioneal Beds of Jamaica Geological Magazine 61: 199-218. N'alentine, J.W 1989 How good was the fossil record? Clues from the Californian Pleistocene. Paleobiology 15: 83-94. \aught, KG. (R.T. Abbott and K J Boss, eds) 1989. A clas- sification of the li\ing Mollusca. American Malacologists Inc., Melbourne, Florida, xii + 195 p. Page 42 THE NAUTILUS, Vol. 107, No. 1 VVarmke, G.L. and R.T .Abbott. 1961. Caribbean seashells. Woodring, W.P 1928 Miocene mollusks from Bowden, ja- Livingston, Narberth, Pennsylvania, x + 346 p. maica, pt. II, gastropods and discussion of results. Carnegie Woodring, W.P. 1925. Miocene mollusks from Bowden, Ja- Institution of Washington Publication 385: 564 p. maica. Carnegie Institution of Washington Publication 366: vii + 222p. INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biology and systematics of mollusks. Manuscripts de- scribing original, unpublished research as well as review articles will be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not re- quire an abstract. Notices of meetings and other items of interest to rnalacologists will appear in a news and notices section. 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Box 7279 Silver Spring, MD 20907-7279 THE €9 NAUTILUS CONTENTS Volume 107, Number 2 Marine Biological LaboraLw/"'^ ^*' ^^^^ LIBRARY ^% 0028-1344 Silvard P. Kool The Systematic Position of the Genius W&d&H HolS MasS (Prosobranchia: Miiricidae; r>r.or.o'kr;n>)p'| ^ 43 Jose H. Leal Donald R. Moore Thala csperanza. a new Costellariidae (Mollusca: Gastropoda) from northern Puerto Rico 58 Anionieto S. Tan Ya-ping Hu IVIichael Castagna Richard A. Luiz Michael J. kennish Alan S. Pooley Shell and Pallet Mor|jhology of Early Developmental Stages of Bankia gouldi (Bartsch, 1908) (Bivalvia: Teredinidae) ... 63 S. J. kleinschusler S. L. Swink A Simple Method tor the In Vitro Oilture of Perkinsus warinus 76 Harold G. Pierce On the Identification of Fossil Terrestrial Gastropod Eggshells 79 THE NAUTILUS 107(2):43-57, 1993 Page 43 The Systematic Position of the Genus NiiceUa (Prosobranchia: Muricidae: Ocenebrinae) Silvard P. Kool Mollusk Department Museum of Comparative Zoology Harvard Universitv Camhriiige. Massachusetts 02138 USA ABSTRACT The muricid genus S'ucclla Roding. 1798 has conimonK been placed in Thaidinae Joiisseaunie, 1888 (Prosobranchia: Muri- cidae). The Thaidinae (scnsii Kool, 1989) is monoph) letic with and thus s\ nonymous with Rapaninae Gra\ , 1853 (Kool, 1993, in press). Comparative anatomical investigations of the type species of Nucclla Roding, 1798 (Buccinum filosum Gmelin, 1791 [=Nttcella lapillus (Linnaeus, 1758)]) and of Thais Rod- ing, 1798 (Murex funis Gmelin, 1791 [ = Thais nodosa (Lin- naeus, 1758)]) as well as other rapanines have revealed that inclusion of Nucclla in Rapaninae would result in polyphyletic groups (Kool, 1989; 1993, in press). Studies of the anatomy, radula, protoconch, shell ultrastructure, and operculum of the type species of A't/ce/Za, Occnebra Gray, 1847 (Murcx crinaceus Linnaeus, 1758 [=Ocenebra erinacea]) (Ocenebrinae Coss- mann, 1903), and Trophon Montfort, 1810 {Murex magellan- tcMS Gmelin, 1791 [=Trophon geversianus (PaWni, 1774)]) (Tro- phoninae Cossmann, 1903), indicate that Nucella has close affinities with Ocenebrinae and Trophoninae Based on cladistic analyses, it is here proposed that Nucclla be placed in Oce- nebrinae. Results further reveal that the distinctions between Ocenebrinae and Trophoninae are less clear than previously accepted Key words: Nucclla, Ocenebrinae, phylogeny; systematics; comparative anatom\ INTRODUCTION The Thaidinae of authors, also referred to as Thaididae, Purpurinae/dae Swainsoii, 1840, Driipinae Wenz, 1941, etc., has been shown to be a conglomerate of disparate taxa (Kool, 1989; 1993, in press). The ta.xonomic coher- ence of the Thaidinae and the boundaries of its genera were based primarily on external shell characters, which are often convergent, obscuring phylogenetic relation- ships. Rigorous cladistic analyses based primarily on char- acters derived from anatomy, radula, operculum, and shell ultrastructure, have shown that para- and poly- phyly were wide-spread in the Thaidinae/dae of authors (Kool, 1989). Subsequent phylogenetic studies have re- vealed that the genus Rapana Schumacher, 1817, and the Thaidinae (in partem) constitute a monophyletic group, making Thaidinae a junior subjective synonym of Rapaninae (Kool, 1993, in press). The name Rapaninae will herein be used for the clade that includes Rapana and Thais Several species of the genus Nitcella have been used extensively in ecological studies (Colton, 1922; Crothers, 1983, 1985; Emlen, 19(56; Etter, 1987; Kincaid, 1957; Moore, 1936, 1938; Palmer, 1983, 1985; Spight, 1972, 1976). In most of these studies Mucella was regarded as a subgenus or synonym of Thais. Anatomical studies (Kool, 1986, 1989) of the type species of Nucella [Buc- cinum filosum = Nucella lapillus (see Kool & Boss, 1992)] and Thais [Murex fucus = Thais noc/o.sa] revealed major differences between these genera. Kool (1988) therefore excluded Nucella from the Thaidinae and tentatively placed Nucella in the Ocenebrinae (Kool, 1989) on the basis of radular (Sabelli & Tommasini, 1987; Bandel, 1977) and protoconch (Bandel, 1975) morphology as well as anatomical descriptions (Graham, 1941) of Ocenebra erinacea. Although the anatomv of Nucella lapillus is well known (Fretter & Graham, 1962; Kool, 1986, 1989; Oehlmann et al., 1988), relatively little is known about the soft parts of Ocenebra erinacea. Aspects of the anatomy of Tro- phon geversianus were described by Harasewych (1984), who suggested that similarities (e.g. radular morphology) between members of Trophon and Nucella may be due either to convergence resulting from similar environ- mental conditions or to phylogenetic affinity. The object of this study is to discern the phylogenetic affinities among Nucella, Trophon and Ocenebra. MATERIALS AND METHODS The following specimens were used for anatomical stud- ies: Nucella lapillus- Kittery, Maine, U.S.A. (USNM 857053) (7 9, 5S). Trophon geversiatius; Daniel Este, Isia Grande, Tierra del Fuego, Chile (LACM 86-270.2); Puerto Basil Hall, Page 44 THE NAUTILUS, Vol. 107, No. 2 Table 1. List of characters and character states for Murican •thus. Thaifi. Troplifm. Nuce 'lla, and Ocenelna. Character Mur Tha Tro Nuc Oce 1. Protoconch whorls 0 0 1 1 2. Calcitic layer 0 1 1 1 3. Number of aragonitic layers 0 1 2 2 0 -4 Position of opercular nucleus 0 3 1 2 5. Opercular shape 0 1 0 1 6. Pigmentation pattern on head-foot re- gion 0 0 1 1 7. Duct(s) for accessory boring organ and ventral pedal gland 0 0 0 1 8. Bursa copulatrix 0 2 1 1 0 9. Seminal receptacles at dorsal periphery of albumen gland 0 1 0 0 0 10 Fenial shape 0 2 1 0 0 11. Penial vas deferens 0 1 0 0 2 12. Prostate 0 1 0 0 0 13. Accessory salivary gland(s) 0 1 1 2 2 14. Straw-like membrane around glancl of Leiblein 0 0 1 1 1 1.5, Posterior duct of gland of Leiblein 0 0 1 1 1 16. Ontral cusp of rachidian 0 0 0 1 1 17 Margin of rachidian basal plate 0 0 1 1 1 Isla de los Estados, Tierra del Fuego, Argentina (LACM 71-289); Piinta Catalina, Isla Grande, Tierra del Fuego, Chile (LACM 80-87.2) (4 9, 3 <5). Ocenebra erinacea; Roscoff, France (MCZ 298425) (2 2. IS). Morphological data were compiled from soft tissues, radulae, shell ultrastructures, protoconchs, and opercula. Living specimens of Nucella and preserved specimens of Nucella. Trophon and Ocenebra were dissected. Radulae (2-4 per species) were cleaned using a potas- sium hydro.xide solution, rinsed in distilled water, air- dried, sputter-coated with carbon and gold, and exam- ined with a Hitachi S-570 scanning electron microscope. Photomicrographs were taken of the unused, matured central portion of each radular ribbon. Shell fragments from at least two individuals of each species were obtained by crushing the shell. Portions from the central region of the body whorl about one- half to three-quarters of a whorl away from the edge of the apertural lip were mounted, sputter-coated with car- bon and gold, and their fracture surfaces observed with a Hitachi S-570 scanning electron microscope. An ap- parentK amorphous outer layer was interpreted as con- sisting of calcite, while layers with organization in crystal lamellar structure were considered aragonitic (data from x-ray diffraction methods confirm these identifications; Kool and Harasewych, in preparation). Cladistic Analysis: Seventeen characters, divided into 41 character states (Table 1 ), were used in a cladistic anaK sis performed with [Iennig86 (Clopyright J.S. Farris, 1988). The six multislate characters were entered as unordered. Most characters were derived from soft tissues (mainly from the male and female reproductive and alimentarv systems), the remainder from radulae, opercula, proto- conchs, and shell ultrastructiire. Two additional species, Thais nodosa and Muricanthus ftdvescens (Sowerby, 1841), are used in the cladistic analysis, based on data in Kool (1989; 1993, in press). Muricanthus fulvescens, a muricine and member of a sister group of Ocenebrinae, Trophoninae and Rapaninae, is used as outgroup for the cladistic analysis. Table 1 lists characters and character states, and re- flects the sequence in which organs and other morpho- logical features are described for each of the three spe- cies. RESULTS DESCRIPTIONS OF TAXA Nucella lapillus. Shell. Protoconch (Figs. 26, 29) conical, low, of about l'/4 smooth whorls, with impressed suture; transition to teleoconch smooth, difficult to discern. Teleoconch high- ly polymorphic; usually elongate, oval, of 6-7 whorls (Figs. 1-6, 21, 22). Adult shell to 55 mm in height, 30 mm in u idth. Body whorl rounded, about 80?f of shell height, smooth or sculptured with pattern of about 15 spiral, occasionally lamellose. ridges. Aperture (to 65% of shell height) oval; outer lip w ide, smooth, occasionally with 3-4 denticles on edge of thickened lip. Columella v\ith moderate callus, flat to concave. Siphonal canal short, open (Fig. 3) to partly closed (Fig. 1). Siphonal fascicle poorly developed, adjacent to callus layer. Shell color variable: white, grey, yellow, brown, orange-red; often banded; aperture, colmiiella white Shell Ultraslrncturc. Innermost laverof crossed-lamellar S. p. Kool, 1993 Page 45 aragoiiite, with crystal planes oriented perpendicular to growing edge [15-20% of tliiekness; often absent (Fig. 28)]; middle layer of crossed-lamellar aragonite. witfi crystal planes oriented parallel to grow ing edge [15-25% of thickness]; outermost laser of calcite [55-85%) of thick- ness] (Fig, 27). Operculum D-shaped, with lateral nucleus just below center right (Figs. 7, 8). Outer surface (Fig. 7) with arch- shaped growth lines recurved at both ends; inner surface (Fig. 8) with 3-5 arch-shaped growth lines, with broad (35-40% of opercular width), callused, glazed outer rim HeocI and Foot. L'niformly light yellow to white. Ce- phalic tentacles elongate, thin. Incurrent siphon short. Mantle edge smooth. Accessory boring organ (Fig. 56, abo) large, well developed, (in females) anterior to, sep- arate from equally large ventral pedal gland (Fig. 56, Pg)- Mantle Cavity Osphradium slightK more than 'i cten- idial length, less than l; ctenidial width. Right pectin usually wider than left. Each lamella (8-10/mm) at- tached to mantle roof along V2 its base. Anteriormost portion of ctenidium straight, extending slightly anterior of osphradium Ctenidial knnellae (9-1 1/mm) wider than high or equalK wide as high, with strongly convex or straight lateral edges, translucent. Thick supporting rod extending beyond lateral edge of each lamella, forming small papilla. Female Reproductive System: Vaginal opening round with slightK swollen edges, located below and posterior to anus. Bursa copulatrix (Fig. 47, 48, be) small diver- ticulum, connected to vagina, ventral channel (vc) by wide ventral passage. Ventral channel formed by two small interlocking flanges located under ventral lobe of capsule gland, one arising from left lobe, the other from ventral epithelium. Single-chambered ingesting gland lo- cated between capsule gland and albumen gland. Al- bumen gland (Fig. 49) arch-shaped, elongate, opening anteriorly into ovi-sperm duct (osd), posteriorly into ovi- duct (od). Ovary yellow to light golden. Many specimens with pseudo-penis of variable size (see also Brvan et al., 1986). Male Reproductive System: Penis (Fig. 60) simple, elon- gate, dorso-ventralK flattened, often slightK curved, w ith abruptly tapering, papilla-like end. Penial vas deferens (Fig. 62, pvd) minute, simple duct, semi-closed by trans- verse ridges on overlapping ventral and dorsal sides of penis. Cephalic vas deferens (Fig. 57, cvd) well devel- oped, extending from penis (p) to prostate gland (pr). Prostate gland (Fig. 57, pr) white; prostate duct (prd) dorso-ventral slit in cross section; duct open to mantle cavity posteriorly. Posterior vas deferens (along visceral mass) well developed, white to dirty white, iridescent. Testis light brow n to golden. Alimentary System: Paired accessors salivar\ glands ex- tremely long, usually equal to or slightly longer than one- half of shell height; left gland intertwined with salivary glamls, right gland separate from salivary glands, situ- ated in right anterior corner of buccal cavity. Salivar% glands in center of dorsal buccal cavity between gland of Leiblein and short, pear-shaped valve of Leiblein. Sali\ar> ducts attached to anterior esophagus at some distance from valve. (Glandular folds in mid-esophageal region inconspicuous. Connection between mid-esoph- agus and gland of Leiblein short, thick. Posterior esoph- agus appressed to left side of gland of Leiblein in loop- shaped fashion. Gland of Leiblein \ellowish; posterior blind duct very short ( < Vi length of gland), with small terminal ampulla. Stomach tubular, with 8-12 large, ra- dialK oriented folds on wall. Stomach typhlosole ex- tending dorsally onto left portion of posterior mixing area. Intestinal t\ phlosole thick, w ide. Tw o digestive di- verticula present. Rectal gland inconspicuous. Large pa- pilla overlying equally large anus. Radula: Ribbon length 30-35% of shell height. Base of rachidian tooth expanded below fwse of neighboring ra- chidian tooth; central cusp of rachidian thin, flame- shaped, leaning more anteriorly (in situ) than lateral cusps; inner lateral denticle low on base of lateral cusp, occasionalK bifurcate (w ithin same specimen); outer edge of lateral cusp w ith se\ eral denticles; large marginal cusp pointing straight forward and parallel to elongate, lateral extension at base of rachidian tooth (Figs. 23-25), re- sulting in bifid appearance of rachidian basal plate. Lat- eral teeth shorter than rachidian width (Fig. 23). Egg capsules: Elongate-oval, vase-shaped, up to 9 mm in height, 3 mm in w iilth. Capsules \ellow, light brow n or purple (Lebour, 1937), each attached by short, thin stalk about 1 mm long. Apex tapered with rounded, cap- shaped top with mucous plug. Capsules interconnected at base. Number of embryos varying from two (Risbec, 1937) to one thousand (Fretter & Graham, 1962, 1985), most being nurse eggs (75-95%) (Crothers, 1985; Fretter & Graham, 1985; Lamv, 1928) (see also Pelseneer, 1911; Ankel, 1937; Thorson, 1941, 1946; Robertson, 1974). Ecology: More is known about Nucclla ecology than about any other muricoidean [for an extensive bibliog- raphy on the biology (primarily ecology) of Nticella la- pillus, see Crothers, 1985]. Nucella lapUlus and its west- ern American congeners have been the topic of man\ comprehensive studies (Crothers, 1985; Emien, 1966; Et- ter, 1987; Kincaid, 1957; Spight, 1972). Nucella feeds on barnacles and mussels (Colton, 1922; Connell, 1970; Crothers, 1973; Graham, 1955; Kool, 1987; Largen, 1967; Murdoch, 1969; Spight, 1982) in the rocky intertidal zone and is eaten by crabs and birds (Spight, 1976). Studies of Agersborg (1929), Colton (1922), and Moore (1936) show that environmental factors (wave action, food avail- abilits, etc.) influence shell morphology. Moore (1938) reported the main spaw ning period to be during winter and spring; but breeding occurs throughout the year (Lebour, 1937; Thorson, 1946). Juveniles hatch from the eggs after 4-7 months (Fretter & Graham, 1985). Distribution: North Atlantic Ocean from southern Por- Page 46 THE NAUTILUS, Vol. 107, No. 2 7 ^^^ % ^^^ / ■ /. 11 8 mi' 17 ^^m '^' 18 19 *- Figures 1-8 .V«C(?//a /api7/us. 1-6. Shells (1,2 MCZ 69192, Freshwater Bay. Isle of Wright. England, height 25 mm; 3,4 MCZ 1 1.509.3. Sullivan, Maine. I S .V . height .35 mm; 5.6 MC.Z 50600. Wales, height 3-1 mm) 7.8 Operculum (MCZ 302404, Braunton, North Devon, England, height 15 mm); 7. Outer surface; 8. Inner surface. Figures »J-14 Trophon gevcrsianus. Shells (9.10 MCZ S. p. Kool. 1993 Page 47 tugal to Xovaya Zembl\a [records from western Medi- terranean, Azores, Morocco, Senegal, and Canary Islands are suspect (Cooke. 1915)]; Great Britain; Ireland; Ice- land; Greenland; New Jerse\, L.S.A.. to northern Canada [for extensi\e list of geographical range and localities, see Cooke, 1915]. Trophon geversianus: Harasewych (1984) described aspects of the anatom\ of Trophon geversianus (Figs. 9. 10, 13, 14. 30. 31 ). Because my observations were congruent w ith the descriptions in Harasewych s paper, only the most essential and supple- mental data are presented to avoid unnecessary dupli- cation. Shell ultrastructure: Innermost la\er of crossed-lamellar aragonite, w ith cr> stal planes oriented perpendicular to growing edge [10-15!^ of thickness; often absent (Fig. 37)]; middle layer of crossed-lamellar aragonite, with crystal planes oriented parallel to growing edge [15-20% of thickness]; outermost layer of calcite [70-80% of thick- ness] (Fig. 36). Operculum: 0\ate, with lateral nucleus in lower right (Figs. 11, 12). Outer surface (Fig. 11) with growth lines recurved at upper ends, progressiveh upright; inner sur- face (Fig. 12) with 3-4 narrow horseshoe-shaped growth lines, broad ( > '3 opercular width ). lightK callused, glazed outer rim. Head and foot: UniformK light \ellow Cephalic ten- tacles elongate, thin. Incurrent siphon short. Mantle edge smooth, .\ccessory boring organ (Fig. 55, abo) well de- veloped, sharing common duct with ventral pedal gland (pg) in females. Mantle Cavity: Osphradium small ('/3-% ctenidial length. -/i ctenidial width). usualK partialK overK ing ctenidium. Right pectin usualK wider than left. Each lamella (9- 10 mm) attached to mantle roof along most of its length. .•\nteriormost portion of ctenidium straight, extending slightly anterior of osphradium. Ctenidial lamellae (10- 12 mm) translucent, wider than high anteriorK. equalK wide as high posteriorly, with straight to convex lateral edges. Female Reproductive System: \'aginal opening (Fig. 47, vo) round, with swollen edges, located below, slightly anterior to anus. Bursa copulatrix (Figs. 47, 48, be) small diverticulum, connected to vagina and ventral channel (vc) by wide duct. Wall of posterior vagina with folds decreasing in number posteriorly. Capsule gland with simple, inconspicuous ventral channel posteriorly con- nected to large, well-de\eloped ingesting gland filled with w hitish substance. .Albumen gland (Fig. 49) large, arch-shaped, elongate, opening anteriorly into ovi-sperm duct (osd), posteriorly into oviduct (od). Male Reproductive System: Penis (Fig. 59) bulbous, short, dorso-\entrall\ flattened, w ith large papilla. Penial vas deferens (Fig. 62. p\d) minute, simple duct, closed by overlapping ventral and dorsal sides of penis. Cephalic vas deferens (Fig. 57, cvd) well developed. Prostate gland (Fig. 57. pr) light yellow; prostate duct (prd) dorso-ven- tral slit in cross section; duct open to mantle cavit\ pos- teriorly. Alimentary System: Paired accessory- salivar\ glands short (<1 10 shell height); left gland intertwined with left salivary gland, right gland free, situated in right anterior corner of buccal cavity. Salivary glands in center of dorsal buccal ca\it\ between gland of Leiblein and elongate \al\e of Leiblein. Salivar\ ducts attached to anterior esophagus immediately anterior to \alve. Glandular folds in mid-esophageal region well developed. Connection betw een mid-esophagus and gland of Leiblein short, thick. Posterior esophagus appressed to left side of gland of Leiblein in loop-shaped fashion. Gland of Leiblein yel- low ish; posterior blind duct short, without terminal am- pulla. Stomach tubular, with 10-15 thin, elevated folds on w all; posterior ones oriented toward center, anterior ones merging into elevated section of sorting area. In- testinal typhlosole thin. Two digestive diverticula pres- ent. Rectal gland light brown, extending along '2 of pal- lia] gonoduct. Radula: Ribbon length 40-45% of shell height. Base of rachidian tooth expanded below base of neighboring ra- chidian tooth; central cusp thin, with wide base; inner lateral denticle small protrusion from base of lateral cusps; outer edge of lateral cusp straight, with several faint denticles; large marginal cusp pointing straight forward and parallel to faint, elongate, lateral extension at base of rachidian tooth (Figs. 32-34), resulting in bifid ap- pearance of rachidian basal plate. Lateral teeth shorter than rachidian width, with wide bases positioned close together (Fig. 32). Egg capsules: Discoidal, lateral!) flattened, with wide, equalK flattened base, up to 20 mm in height (including base), 12 mm in width (Harasewych, 1984, Fig. 23). Capsules yellowish in color, and containing 74-112 em- bryos (MeKill & Standen, 1898). Capsules deposited in row s w ith flattened edges adjacent to one another (Lamv, 1928; D'Asaro, 1991). Ecology: This species lives in the rocky intertidal and subtidal zones where barnacles and mussels are plentiful. 10941.3. Ushaia, Tierra del Fuego, Argentina, height 41 mm; 1:{.I4 MCZ 1-32566. Falkland Islands. Argentina, height 47 mm). 11.12 Operculum iLACM 86-270.2, Daniel Este. Isla Grande. Tierra del Fuego. Chile, height 11 mm); 11. Outer surface; 12. Inner surface. Figures 15-20 Ocenebra erinacea. Shells (15.16 MCZ 87662. \Ve\mouth. England, height 31 mm; 19,20 MCZ 1724.50. Fos-sur-Mer. Bouches-du-Rhone, France, height 56 mm) Operculum (17.18 MCZ 302405, St. Lunaire. France, height 8 0 mm); 17 Outer surface; 18. Inner surface. Page 48 THE NAUTILUS, Vol. 107, No. 2 Figures 21-29 \ucclla lapillus. 21,22 Shell (MCZ 09192, Freshwater Bay, Isle ot Wight, England, height 2.5 1 mm). 23-2.S Kadiila (ISNM 8.570.53, Kittery, Maine, LI.S.A.). 23. Clentral portion of radular ribbon. Scale bar = 15 fim. 24 Rachidian teeth Scale bar = 10 ^m. 25. Side view of rachidian teeth (right row of lateral teeth removed). Scale bar = 10 fim. 26,29 Protoconch (MCZ 14184, Isle au Ilaut, Maine, U.S.A.). 26. Apical view. Scale bar = 12 m"' 29. Side view. Scale bar = 12 ^m 27,28 Shell ultrastructure (view of growing edge; innermost layer on bottom) (MCZ 69192, Freshwater Ba>, Isle of Wight, England) Scale bars, 4.5 ixn\ and 70 niu, respectively. S. p. Kool, 1993 Page 49 Figures 30-38. Troplion gcicrsianns. 30.31. Sliell (MC^Z 1.32566. Falkland Islands, Argentina, lu'iglit -JT mm). 32-3 4-. Radula. (LACM 86-270.2. Daniel Kste. Isia Grande. Tierra del Fuego. Chile) 32. Central portion of radiilar ribbon. Scale bar = 45 nm. 33. Rachidian teeth Scale liar = 25 ^m. 34 Side view of rachidian teeth. Scale bar = 20 ^in 35,38. Protoconch (LACM 86- 270.2, Daniel Este, Isla Grande, Tierra tlel Fuego, Chile). 35. Apical view. Scale bar = 150 ^m. 38 Side view. Scale bar = 150 Mm 36.37 Shell ultrastructure (L.ACM 86-270 2, Daniel Fsle. Isla (Iraiide, Tierra del F'liego, Chile); Scale bars, 60 fim and 150 Mm, respectivel) Page 50 THE NAUTILUS, Vol. 107, No. 2 4* ^ 39 W i ■?•■,■ ■ 40 ' i^BFjlg; ^ atk..: .«► f43 IP ^*^. ■^ Fif^ures 39-46. Ocenebra crinai-ciL 39,40, Sliell (MCZ 172450, Fos-sur Mer, Botichfs-dii-Hlioiu", France, height 56 mm) 41-43 Kathila (MCZ 298425, Roseoll, France). 41 Oenlral portion of rihhoii Scale bar = 17 ^ni 42. Rachichan teeth Scale bar = 8 ^m. 43 Side view of rachidian teeth. Scale bar = 9 m"i 44.46 Protoconch (MC:Z 38369, Kent, Fngland) 44. .Apical view. Scale bar = 9(1 A/ni 46 Side view. Scale bar = 90 niu 45 Shell iillrastnictiire (MCZ 298425, Roscoff, France), Scale bar = 1.30 ^m. S. p. Kool, 1993 Page 51 Figures 47-56. Anatomical structures of the female repro- ductive system of Nitcella lapilltis. Trophon geversianus, Oce- nehra erinacea, Muricanthus fulvescens. and Thais nodosa. 47.50.53. Bursa copulatrix in iV lapillus and T. geversianus (47), Th. nodosa (50), O erinacea ami M. fulvesccns (53). 48.51.54. Cross sections through bursa copulatrix (location indicated b>' vertical bar). 49,52. Albumen glands in N. la- pillus. T gever.'iianus. O erinacea. M Julvescens (49), anil Th. nodosa (52), 55,56. Sagittal section through foot showing accessory boring organ and pedal gland of Th. nodosa, T geversianus, M. julvescens (55), N . lapillus. and O. erinacea (56). abo, acces.sory boring organ; be, bursa copulatrix, db, duct to bursa copulatrix; leg, left lobe of capsule gland, od, oviduct, osd. ovi-sperm duct; pg, ventral pedal gland; p,sr. posterior seminal receptacles, rcg. right lobe of capsule gland, tf, transverse fold; vc. ventral channel, \cf, flange of the ventral chaiuiel, vo. vaginal opening. Sculpture of the shell may vary v\ itli the t\ pe ot habitat Distribution: Southern Argentina to Chile. Ocenebra erinacea: Shell: Protoconch (Figs. 44, 46) conical, low, of l'/2 smooth whorls, and with impressed suture; transition to teleo- conch smooth, difhcult to discern. Teleoconch elongate, fusiform, biconical (Figs. 15, 16, 19, 20, 39, 40), of 7-8 whorls. Adult shell highly variable in shape, to 55 mm in height, 25 mm in width (not including spine length; 35 mm including spine length). Body whorl 55-60?c of shell height, with 3-9 varices, often with frilled edges, and with (3-8 spiral cords. Aperture (to 30% of shell height) round to oval; outer lip with crenulated edge Moderately callused columella. Siphonal canal partly or completely closed, often nearly equal in length to aperture in larger Figures 57-64 .■\nal0,b2). O erinacea (60.63), and Th nodosa (61,64). b\ , blood vessel, cvd, cephalic vas deferens; p, penis, pr, prostate, prd, duct through prostate, pvd, penial vas deferens. specimens. Siphonal fasciole pointing away from si- phonal canal. Shell color yellov\ish to cream or dark brown; aperture, columella white. Shell ultrastructure: Innermost layer of crossed-lamellar aragonite, with crystal planes oriented perpendicular to growing edge [15-20% of thickness]; followed by layer of crossed-lamellar aragonite, with crystal planes ori- ented parallel to growing edge [40-45% of thickness]; followed by layer of crossed-lamellar aragonite, with crystal planes oriented perpendicular to growing edge [5-8%. of thickness]; outermost layer of calcite [35%. of thickness] (Fig. 45). Operculum: D-shaped, with lateral nucleus in lower right (Figs. 17, 18). Outer surface (Fig. 17) with arch-shaped growth lines progressively upright, recurved at upper end; inner surface (Fig. 18) with 4-5 arch-shaped growth lines, w ith broad (~ Vz opercular w idth), lightly callused, glazed outer rim. Head and foot: I'niformly light yellow to white. Ce- phalic tentacles elongate, thin. Incurrent siphon well de- veloped. Mantle edge smooth, occasionally with crenu- lations (possibly an artifact). Sole of foot w ith large lateral Page 52 THE NAUTILUS, Vol. 107, No. 2 (0 s (0 (0 c o Q. O (0 c O o i5 8-1 10- 11 - 4 - 3- — 8- 16- - 12 - 11 - 3- 13- 7- 5 - - 10 - 9- - - 8- 17 - - 5- 15 - - 4- 14- - 3 - 6 - - 4 - 1 - - 13- ^ 2- — Fipurp 65 Cladciyram, based on analysis of data in Table 1, shduiiig high |)ii\l()geiietic affinity between Nucclla and Oce- nebra. Numbers next to character changes correspond to num- bers given to characters in Table 1. Alternative, equally par- simonious character state transformation series are possible for characters 3, 4, 5, and 8 (see text). folds. Accessory boring organ (Fig. 56, abo) large, well developed, (in females) anterior to, separate from equally large ventral pedal gland (Fig. 56, pg). Mantle Cavity. Ospliradium (<'/2 ctenidial length, % ctenidial width) partially extending over ctenidium. Right pectin usually wider than left. Each lamella ( 10-1 l/mm) attached to mantle roof along short basal portion. An- teriormost portion of ctenidium straight, extending an- teriorly of osphradium. (Ctenidial lamellae (11-12/mm) basically triangular, longer than high w ith convex lateral edges posteriorly. Lamellar support rods not well devel- oped. Female Reproductive System: Vaginal opening (Fig. 53, vo) round to elongate, on short, extension of pallial gon- oduct, situated below, posterior to anus. Bursa copulatrix (Figs. 53, 54, be) large diverticuhmi (equal in diameter to capsule gland), connected at its anterior portion to duct running parallel to ventral channel for some length prior to connecting with it (Figs. 53, 54, db). Lumen of bursa copulatrix Oiled w ith loose flocculent material and iridescent spherules. Capsule gland posterior to, shorter than bursa copulatrix. Ventral channel (Figures 53, 54, vc) well developed anteriorly, less distinct posteriorly. Ingesting gland situated between capsule gland, albumen gland. Albumen gland (Fig. 49) arch-shaped, elongate, opening anteriorly into ovi-sperm duct (osd) posteriorly into oviduct (od). Ovary orange-yellow. Pseudo-penis oc- casionally present (see also Feral, 1976). Male Reproductive System. Penis (Fig. 60) simple, dor- so-ventrally flattened, elongate, slightly curved, with abruptly tapering, papilla-like end. Penial vas deferens (Fig. 63, pvd) a wide, straight tube star-shaped in cross section; tube partialK attached to penial wall Cephalic vas deferens (Fig. 57, cvd) large, similar to penial vas deferens in structure, extending from penis (p) to prostate gland (pr). Prostate gland (Fig. 57, pr) white to yellow; prostate duct (prd) a dorso-ventral slit in cross section anteriorK , a triangular large space posteriori)-; duct open to mantle cavity posteriorly (Fig. 57). Posterior vas de- ferens white to dirty white, iridescent. Testis yellowish. Alimentary System: Paired accessory salivary glands very long (* '/2 shell height); left gland intertwined with left salivary gland, right gland free, situated in right anterior corner of buccal cavity. Salivary glands in center of dorsal buccal cavity between gland of Leibleiii and short, pear- shaped valve of Leiblein. Salivary ducts attached to an- terior esophagus at short distance from valve. Glandular folds in mid-esophageal region swollen, especially well developed at connecting point between esophagus and gland of Leiblein. C-onnection between mid-esophagus and gland of Leiblein short, thick. Posterior esophagus appressed to left side of gland of Leiblein in loop-shaped fashion. Gland of Leiblein yellowish; posterior blind duct very short ( 2 whorls); 1, paucispiral (< 1': whorls). Nucella. Trophon and Occncbra (Figs. 26, 29, 35, 38, 44, 46) have a paucispiral, smooth protoconch without a sinusigeral notch or outwardly-flared lip. This morphol- ogy reflects direct development. The outgroup and the rapanines have a multispiral (and generalK sculptured) protoconch w ith a sinusigeral notch and outwaril-tlaring lip, t\pical for species v\itli a pianktonic larval stage. Character 2. Calcitic layer; 0. absent; 1. present. The outgroup lacks an outer calcitic shell layer that is present in the other four ta.xa. The presence of calcite appears to be the derived condition. Character 3. Number of aragonitic layers: 0. 3; 1. 4; 2. 2 The outgroup and Ocenebra have three layers of ara- gonite (transverse, collabral, transverse), Thais has an additional, innermost fourth layer of crystals oriented in a 45° angle; both Nucella and Trophon have two layers (transverse, collabral), but may lack the innermost trans- verse layer (Figs. 28, 37). Character 4. Nucleus of operculum: 0. terminal nucleus in lower right; 1. lateral nucleus in lower right; 2. lateral niiclfus licldw center right; 3 lateral nucleus in center right. The outgroup has a terminal nucleus in the lower right. .All iiigroup taxa have a lateral nucleus, the position of w Inch varies. The nucleus of the operculum in Nucella is located below the center right; that of Trophon and Ocenebra in the lower right (Figs. 7, 11, 17, respectively). The nucleus is located in the center right in rapanines. C;haracter 5, Shape of operculum: 0. oval; 1. D-shaped. The opercula of Nucella, Thais, and Ocenelna are roughly D-shaped Those of Muricanthus and Trophon are elongate-oval. (Character 6, Pigmentation pattern of head-toot region: 0 jjresent; flecked with black and gra\; 1. absent, uni- formly colored (faint yellow ). The head-foot regions of Nucella, Trophon, and Oce- nebra are uniformly faint yellow. Both Thais and the outgroup are densely flecked with black blotches and specks (this pattern generalK survives preservation in alcohol although other colors, such as white and yellow, fade). Character 7. Ventral pedal gland and accessory boring organ: 0. sharing one duct; 1. with separate ducts. In Nucella and Ocenebra, the accessory boring organ and ventral pedal gland (Fig. 56, abo, pg) have separate ducts to the sole of the loot, while in Trophon, rapanines, and the outgroup these structures share a common duct (Fig. 55, abo, pg). CJiaracter 8. Bursa copulatrix: 0. large diverticulum, sep- arate from capsule gland; 1. small diverticulum, separate from capsule gland; 2. small chamber with lumen con- tinuous with capsule gland. The bursa copulatrix is a small blind sack in Nucella and Trophon (Fig. 47), a large separate diverticulum in Ocenebra (Fig. 53) and the outgroup, while in rapanines the bursa is continuous with the capsule gland (Figs. 50, 51). Character 9. Seminal receptacles at dorsal periphery of albumen gland: 0. absent; I. present. In Thais, a row of posterior seminal receptacles (Fig. 52. psr) at the dorsal periphery of the albumen gland presumably increases efficiency in the fertilization pro- cess (Kool, 1988, 1989). These posterior seminal recep- tacles are absent in Nucella, Trophon. Ocenebra. and in the outgroup, Muricanthus (Fig. 49). This character is a synapomorphy for Rapaninae (Kool, 1993, in press). Character 10. Penial shape: 0. simple, elongate to lightly curved; 1. bulbous, with papilla; 2 strongK recurved with pseudo-papilla. Penial shape in Nucella. Ocenebra. and the outgroup is elongate (F"ig. 60). The penis in Thais is strongly recurved, and sinuous (Fig. 61), while that of Trophon is short, bulbous, with a distinct papilla (Fig. 59). (Character 11 Penial \as deferens: 0. simple duct; 1. small, loose duct-w ithin-a-duct; 2. large duct-within-a- duct, partialK attached to penial iiuier wall Page 54 THE NAUTILUS. Vol. 107, No. 2 Nucella, Trophon and the outgroiip have a peiiial vas deferens that is loosely closed, while rapanines have a ■■duct-vvithiii-a-duct" system (Figs. 61, 64) (Kool, 1988, 1989). Ocenehra differs from both types in having a rather wide inner duct that is partially attached to the penial inner wall (Fig. 63). Character 12. Prostate Gland: 0. open to mantle cavity posteriorly; 1. without opening to mantle cavity. Males of Nucella. Trophon. Occnelrra. and the out- group have a prostate gland that is open to the mantle cavity along its posterior portion (Fig. 57). The prostate of rapanine males does not open to the mantle cavity (Fig. 58). Character 13. Accessory salivary gland length: 0. right gland small, left gland absent; 1. glands < '/i shell height; 2. glands > V2 shell height. Character 14. Straw-like membrane around gland of Leiblein: 0. present; 1. absent. In most rapanines and the outgroup, the gland of Leib- lein is covered by a thick membrane of interwoven fibers of connective tissue, producing a straw-like appearance. Such a membrane is absent in Nucella. Trophon and Ocenehra. Character 15. Posterior duct of gland of Leiblein: 0. longer than Vz of gland length; 1. shorter than V2 gland length. In Nucella, Trophon and Ocenehra. the gland of Leib- lein tapers posteriorly into a thin, very short posterior duct that runs adjacent to the posterior esophagus and is often filled with secretory material from the gland; in the majority of rapanines and in the outgroup this duct is much longer, extending into the dorsal branch of the afferent renal vein. Character 16. Central cusp of rachidian: 0. oriented in same plane as lateral cusps; 1. leaning more anteriorly than lateral cusps. In Nucella (Fig. 25) and Ocenehra (Fig. 43) the central cusp on the rachidian leans more anteriorly (in situ ) than the lateral cusps. In Thais. Trophon (Fig. 34), and the outgroup, the lateral cusps and central cusp are aligned in the same plane. Character 17. Margin of rachidian basal plate: 0. straight; 1. bifid. The bifid condition of the rachidian basal plate (Figs. 25, 34, most developed in Fig. 43) is found in Nucella, Trophon and Oceneina, but not in Thais or the outgroup. Synapornorphies for the Nucella-Ocenehra clade {Fig. 65): Character 5: The character for opercular shape is ho- moplastic; a D-shaped operculum occurs in Thais as well. C]haracter 7: Both Nucella and Ocenehra have separate openings for the ventral pedal gland and accessory boring organ (Fig. 56, abo, pg). Female specimens of Trophon geversianus, Thais nodosa, and the outgroup, have a single duct and opening for these organs (l"ig. 55, abo. pg). A shared duct for the accessory boring organ and ventral pedal gland, as found in Trophon geversianus, may not be as advantageous as an arrangement as w hen the ducts originating from the accessory boring organ and ventral pedal gland are separate. It would appear that an arrangement where one duct serves both as ven- tral pedal gland and as a passage for the accessory boring organ and its secretions during boring activities (Carri- ker, 1981) prevents the female from boring activity, and thus perhaps feeding in general, during stages of egg- laying. Character 13: Nucella and Ocenehra have a pair of very long accessory salivary glands (> ','2 shell height). Tro- p/!o;(andT/ia!shavemuchsmallerglands(< '/^ shell height). The outgroup has only one extremely small right acces- sory salivary gland. Character 16: In Nucella, and to a much greater degree in Ocenehra, the central cusp leans more anteriorly (Figs. 25, 43, respectively) (in situ) than the lateral cusps, whereas in Trophon, the central cusp is aligned with the lateral cusps (Fig. 34). Synapornorphies for the Trophon-Nucella-Ocenehra Clade: Character 1: The paucispiral protoconch is indicative of having crawl-away larvae, rather than a planktonic larval stage that is found in Thais and other rapanines (Kool, 1993, in press) and the outgroup. Character 4: The cladogram suggests that an opercular nucleus below the center right (Nucella; character state 2) evolved from the ancestral condition for the taxa in this clade of having a nucleus in the lower right (Oce- nehra and Trophon; character state 1). Character 6: All three taxa lack a pigmentation pattern on their head-foot region that is found in the outgroup, Thais, and other members of the Rapaninae. Character 14: The straw-like outer membrane of the gland of Leiblein is absent in the species of this clade, but present in the outgroup, and most members of Ra- paninae (Kool, 1989; 1993, in press). Character 15: The posterior duct of the gland of Leiblein is shorter than V2 the length ot the gland itself in this clade, but much longer in the remaining taxa, reaching into the dorsal branch of the afferent renal vein. Character 17: The bifid condition of the basal plate, especially well developed in Ocenehra (Fig. 43), is absent in Thais and the outgroup. Synaponiorphics for the Thais-Trophon -Nucella-Oce- nehra Clade: Character 2: An outer layer of calcite is present in all ingroup taxa, but is absent in the outgroup. Character 13: The outgroup has only one extremely small right accessory .sali\ar> gland. A situation of having a pair of medium-size accessory salivary glands appears to S. p. Kool, 1993 Page 55 have evolved from the condition described above and to have given rise to the most deri\ed condition (extremely long glands). DISCUSSION According to the topology of the cladogram (Fig. 65), two characters have evolved in a parallel manner in Nucella lapillus and Trophon geversianus. Out of the context of the cladogram, these similarities would suggest a closer relationship between these two species than is suggested by the tree topology: Character 3: Shell ultrastructure in both Nucella and Trophon consists of two aragonitic layers and an outer layer of calcite. Specimens of both may lack the inner- most (transverse) layer (Figs. 28, 37). More detailed stud- ies may reveal the cause of this variation. Perhaps en- vironmental factors may play a role (Etter, personal communication). Character 8: The morphology of the bursa copulatrix of Nucella is very similar to that of Trophon. In both taxa, a relatively small, muscular blind sack branches off from the vagina (Figs. 47, 48). In Ocenebra the bursa is thin- walled and equal in width and height to the capsule gland, extending for up to V2 the length of the pallial complex. The ventral channel loops backwards towards the anterior portion of the bursa in Ocenebra (Fig. 53), rather than straight up into the bursa as in Trophon and Nucella (Fig. 47). The above two characters could be considered synapo- morphies for an alternative, but less parsimonious, tree in which Trophon and Nucella would be united in one clade. However, in the proposed phylogenetic hypothesis (Fig. 65), four synapomorphies support a clade consisting of Nucella and Ocenebra. rendering the above two char- acters as homoplastic. Fretter and Graham (1962) mention several similari- ties in egg capsule morphology between Nucella lapillus and Ocenebra erinacea. Both species lay vase-shaped capsules, whereas Trophon geversianus produces dis- coidal egg capsules. However, having discoidal egg cap- sules is only an autapomorphic trait for Trophon and does not provide clues about relationship in this case. Alternative, equally parsimonious transformation series are possible for characters 3, 4, 5, and 8. For Characters 3, 5, and 8, I chose the scenario involving homoplasy over one involving a reversal to avoid an "artificial" increase in synapomorphies. Similarly, for Character 4, I chose the least linear trans-formation series ("zero state" evolving into both the "three state" and the "one state"). Zoogeographical data reveal that members of Nucella, Ocenebra and Trophon occur primarily in colder waters of the temperate and boreal zones, whereas rapanines and the outgroup occur primarily in warmer waters of the (sub)tropics (Kool, 1989). Nucella lapillus and Ocenebra erinacea overlap for much of their ranges in primarily temperate western European waters. In addition, N. lapillus occurs in the western Atlantic, where O. erinacea does not, and O. erinacea occurs in the Mediterranean, where N. lapillus has not occurred since the Pleistocene [Malatesta (1960) cited records of N. lapillus from the Pleistocene of Sicily]. Cooke (1915), in a comprehensive list of localities for Nucella lapillus, showed that the southernmost record for this species is the Algarve coast of Portugal and al- leged that any records from Northern Africa (see Nord- sieck, 1982), the western Mediterranean, the Azores, and the Canary Islands, are highly suspect. Other members of what can be assumed to be Ocenebra s.s. and Nucella s.s. are found in the eastern Pacific (Abbott, 1974) and the northern I'acific. Furthermore, Ocenebra s.s. occurs in western Africa (Houart, 1989) and South Africa (Kil- burn & Rippey, 1982). Ranges of these genera may be revised when more Ocenebra-like taxa (for example from Japan) and Nucclla-Uke species have been examined with respect to their anatomy, radula, shell ultrastructure, etc. Such studies are also necessary to determine if, for ex- ample, species such as Nucella dubia and Nucella squa- mosa, both from the South African Province, are indeed members of Nucella s.s. It appears from preliminary dissections that the genus Nucella can no longer be con- sidered restricted to the temperate waters of the northern hemisphere (Kool, in preparation). Trophon geversianus, limited to the South American continent, lives in temperate to boreal waters, as do Nu- cella lapillus and Ocenebra erinacea. SYSTEMATIC CONCLUSIONS The high degree of similarity in anatomy, radula, pro- toconch, shell ultrastructure and operculum in Ocenebra erinacea, Trophon geversianus and Nucella lapillus in- dicates that these three taxa are more closely related to one another than any one of them is to Thais or other representatives of Rapaninae. Nucella should therefore be excluded from Rapaninae to maintain monophyly (Kool, 1989; 1993, in press). The difficulty of correctly allocating Nucella to a subfamily is indicative of the dilemma of our lack of understanding of higher muri- coidean systematics. The cladistic analysis and the re- sulting cladogram (Fig. 65) suggest that Nucella is better placed in Ocenebrinae than in Trophoninae. It is obvious that the boundaries of groups at the higher taxonomic categories, traditionally based on shell characters, be- come less clear after completion of thorough anatomical studies of members from different genera and subfam- ilies. Results shown here suggest that Trophon geversi- anus, the type species of Trophoninae, is closely related to Nucella and Ocenebra. Perhaps it is more closelv related to the latter two taxa than to other species hitherto included in Trophoninae. It is beyond the scope of this paper to suggest synonymization of Trophoninae with Ocenebrinae. However, I suspect that future studies will show that Trophoninae is not a monophyletic group and that a new name for some of its members may be war- ranted. The following systematic arrangement is pro- posed for the taxa treated herein: Page 56 THE NAUTILUS, Vol. 107, No. 2 MURICOIDEA Rafinesque, 1815 MURICIDAE Rafinesque, 1815 RAPANTNAE Gray, 1853 (sen.su Kool, 1993, in press) Thais Roding, 1798 OCENEBRINAE Cossmann, 1903 Ocenebra Gray, 1847 Nucella R6ding, 1798 TROPHONINAE Cossmann, 1903) Trophon Montfort, 1810 ACKNOWLEDGEMENTS I thank Drs. Richard S. Houbrick, M. G. Harasevvych, and Kenneth J. Boss for reviewing an early draft of this manuscript. I thank the staff of the Scanning Electron Microscope Laboratories at the United States National Museum of Natural History and the Smithsonian Marine Station at Link Port, Ft. Pierce, for their assistance. Dis- cussions with Dr. Diana Lipscomb were of great help in fine-tuning the section on the cladistic analysis. Dr. An- ders Waren kindly sent me some well-preserved material of Ocenebra erinacea; Dr. James H. McLean and Mr. C. Clifton Coney provided specimens of Trophon gev- ersianiis. This is Contribution No. 333 of the Smithsonian Marine Station at Link Port, Ft. Pierce, Florida. LITERATURE CITED Abbott, R T, 1974, American Seashells, 2nd edition. \'an Nostrand Reinhold Company. New York, 663 pp. Agersborg, H. P. K. 1929. Factors in the evolution of the prosobranchiate mollusc Thais lapillus. The Nautilus 43(2): 45-49. Ankel, W. E. 1937. 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M. 1972 Patternsof change in adjacent populations of an intertidal snail, Thais lamcUosa. PhD dissertation, Universitv of Washington, Seattle, Washington, 325 pp. Spight, T. M. 1976, Colors and patterns of an intertidal snail, Thais lamellosa. Researches on Population Ecology 17: 176-190 Spight, T M, 1982 Population sizes of two marine snails with a changing food suppl> Journal of Experimental Marine Biology and Ecology 57:195-217, Thorson, G 1941. Marine Gastropoda Prosobranchiata The Zoology of Iceland 4(60): 1-150. Thorson, G 1946 Reproduction and larval development of Danish Marine Bottom Invertebrates Meddelelser fra Kommissionen for Danmarks Fiskeri- og Havunder- sogelser Serie: Plankton 4:1-523 Wenz, W. 1941. Prosobranchia. In: Handbuch der Palao- zoologie. Volume 6. OH. Schindewolf, ed., Gebriider Borntraeger, Berlin, part 5, pp. 961-1200. THE NAUTILUS 107(2):58-62, 1993 Page 58 Thala esperanza, a new Costellariidae (Mollusca: Gastropoda) from northern Puerto Rico Jose H. Leal' Donald R. Moore Rosenstiel School of Marine am! Atmospheric Science Uni\ersity ol Miami 4600 Rickenbacker Causeway Miami, Florida 33149-1098 USA ABSTRACT Thala esperanza n sp. is described from Piaya Esperanza, municipality of Manati, on the northern coast of Puerto Rico. Small size, slender and delicate shell, smaller spire angle, larger number of spiral cords, and a distincti\e constriction below the suture separates the new taxon from western Atlantic conge- neric species. Key words: Thala. Costellariidae, Puerto Rico, Atlantic Ocean, Caribbean Sea, new species, systematics. INTRODUCTION The costellariid genus Thala H. & A. Adams, 1853 com- prises about 12 Recent species, of which two are known from the western Atlantic. Thala fovcata (Sowerby, 1834) is known to occur at St. Vincent, Antilles and possibK at the island of Sao Thome in the eastern Atlantic (Maes & Raeihle, 1975). Thala floridana (Dall, 1884) is found along the coasts of Florida, northern Cuba, the Gulf of Mexico, Haiti, and the Yucatan Peninsula. Cernohorsky (1970) and Abbott (1974) s\non\ mized T. floridana with T. foveata, before Maes and Raeihle (1975) demonstrat- ed that they are separate species. These latter authors, following preliminary observations of Raeihle ( 1968), have shown in a detailed anatomical study of T. floridana that Thala belongs in the Costellariidae (as V'exillidae), and provided support for the then still controversial proposal that this latter taxon deserved full familial status apart from Mitridae (e.g., Azuma, 1965; Ponder, 1972) due to differences in gross anatomy and early life history. In this work we describe a third western Atlantic spe- cies of Thala collected in beach sediments from Manati, ' Current address: Department of Invertebrate Zoology, NHB stop 118, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA. Puerto Rico. Soft parts and radula are unknown; none- theless, conchological characters are distinctive enough to justify the naming of a new species. MATERIALS AND METHODS Shells were measured using WILD M-5 and M-8 dis- secting microscopes with ocular micrometers and pho- tographed with a Nikon F2 camera with extension bel- lows and MicroNikkor 55 mm objective. Scanning electron micrographs were made using an ISI Double Stage DS- 130 scanning electron microscope at the Electron Mi- croscope Laboratory, Rosenstiel School of Marine and Atmospheric Science. Model I ANOVA with unequal sample sizes (Table 1) was performed according to Sokal and Rohlf (1981). Acronyms used throughout this paper are: ANSP, Academy of Natural Sciences of Philadelphia; FMNH, Field Museum of Natural History, Chicago; MCZ, Museum of Comparative Zoology, Har\ard Uni- versity, Cambridge; MNHN, Museum National d'His- toire Naturelle, Paris; UF, Florida Museum of Natural History, University of Florida, Gainesville; UMML, In- vertebrate Museum, University of Miami Rosenstiel School of Marine and Atmospheric Science; L'SNM, Na- tional Museum of Natural History, Smithsonian Insti- tution, Washington. SYSTEMATICS Family Costellariidae MacDonald, 1860 Genus Thala H. & A. Adams, 1853 Thala esperanza new species (figures 1-6, Table 1) Description: Shell fusiform (length width ~ 2.7), im- perforate, reaching 5.0 mm in length, 1.9 mm width. Spire angle about 27° Shell surface smooth, translucent, golden brown to chestnut brown, usually with single spiral baiul of lighter color on middle of last w horl. Peri- ostracmn undetectable. Protoconcli usually decollated in adult shells, when present transparent brown, smooth, with 2 whorls, about 0.4 mm diameter, embryonic whorl J. H. Leal and D. R. Moore, 1993 Page 59 Table 1. Linear shell measurements (mm) and counts for Thala esperanza new species, holotype and paratypes 1, 2, 5-12, and T. foveata. Statistics for model I .\NO\'A with unequal sample sizes according to Sokal and Rohlf (1981); NS = not significantly different. T c.s/x ■ranza (n = 11) 7", fovea la (n = 6) C'haracter Range .\ SD Range .\ SD ANOVA Total length 3.92-500 440 0,40 5-25-6,33 5,73 0.40 F = 24.14 p < 0.001 Shell uidth 1.36-1.88 1,62 0,16 2 04-2 24 2 15 0.07 F = 23.02 p < 0001 Length last whorl 2.80-3.48 3,17 022 3,72-4,28 3,97 0.19 F = 56.34 p < 0.001 .\perture length 1.92-2,44 2 18 0,15 2,84-3,32 3,02 0.20 F = 81.54 p < 0.001 .\perture wiilth 0.36-0,(50 0 49 0 06 0,64-0,80 0 71 0.05 F = 65.12 p < 0.001 Teleoconch whorls 3,25-5,00 4,18 0,45 4,00-4,75 4,42 0.26 NS Spire angle (degrees) 26-29 26,9 1 14 35-37 36.0 0.89 F = 213.9 p rnohorsk\, \N'. O 1970. S\stematics of the families Mitri- dae and N'oiutoniitridae (Moilusca: Gastropoda). Bulletin of the .Auckland Institute and Museum 8: i-ii + 1-190. Ka\, E. \. 1979. Hawaiian marine shells. Reef and shore fauna of Hawaii Section 4: Moilusca. Bernice P. Bishop Museum Special Publication 64(4), Honolulu, 653 p. Keen. .\ M 1971 Seasliells of tropical West .\merica. Marine mollusks from Lower California to Colombia. Stanford Universit\' Press, Stanford, 626 p. Leal, J. H. 1991. Marine prosoliranch gastropods from oceanic islands off Brazil: species composition and biogeography. Uni\ersal Book Services, Oegstgeest, 419 p. Maes. \' O and D Raeihle. 1975. Systematics and biology of Thala floridana (Gastropoda: N'exillidae). Malacologia 15(l):43-67. Ponder, W. F 1972. The morpholog) of some mitriform gastropods with special reference to their alimentar) and reproductive systems, Malacologia 1 1(2):295-342. Raeihle. D 1968 Notes on capti\e Cerithium variabile C.B .\dams and Milra floridana Dall .\muial Report of the .\merican Malacological L nion 1968:35-36. Sokal, R. R. and F. J. Rohlf. 1981. Biometry: the principles and practice of statistics in biological research. Second edition. W.H. Freeman, San Francisco, 859 p. Sphon, G. G. 1969. Notes on the the Mitridae of the eastern Pacific, 2. The genus Thala, with the description of a new species. The N'eliger 12(11:84-88. APPENDIX 1 Material of Thala foveata examined, all from Calliaqua, St. \incent. British N'irgin Islands: LP 158165, 1 shell, beach drift, H.H. Monroe, 01 1961, ex-McGinty Collec- tion; UF 145855, 2 shells, H.H. Monroe, 01/1961, ex- McGint\ Collection; "rockv co\e on windward side of SE Point, Calliaqua Bav", 13°07'20"\, 6n 1'55"W, ANSP 324791, 1 shell, V.O. Maes 02 1972; 13°07'04"N, 61°12'20"W, ANSP 354555, 2 shells, H.H. Monroe 1956, Ex-Einlay Collection. Material of Thala floridana examined: East Coast of Florida: St. Augustine, L'F 158167, 3 shells, Ted Yocius 10/1972, 225 ft, Ex-J.M. Bijur Collection; AMNH 246013, 2 shells, Ted Yocius, Jerome M. Bijur Collection; Fort Pierce, AMNH 127640, 16 shells, Thomas Hughes leg.; MCZ 298862, 3 shells, Bernadine Baker coll., Ex-Doris Folsom 5 1976; Palm Beach, Bovnton Beach, UF 145856, 1 shell, McGinty 8 1944; UF 1(19469, 41 shells, McGinty Collection, 02/1944; Bo\nton Beach, Ocean Ridge, UF St. Vincent, West Indies, 7. Shell (SEM), 8. Shell, M. Shell sculpture near adapical part of aperture (SEM), Figures 10-1 I. Thala floridana 10. ISNM S602S4, Bear Gut, Miami, Florida (SEM), 1 1. I'SNM 860283, off Miami Beach, Florida, Scale bars = 1 mm. except for Figures 3, 5, 6, 9 = 0,: Page 62 THE NAUTILUS, Vol. 107, No. 2 158164, 1 shell, T. McGinty 8/1944; Card Sound. UF 63207, 2 shells, M. Smith 1/1935, I'liiversity of Alabama leg.; UF 13150, 1 shell, C.B. Lungren 1909; 1.6 km off Miami Beach, USNM 860283, 1 shell, M. Almasi and D. Marszalek 10/1978, 24 m depth; Bear Cut, Miami, USNM 860284, A. Emery 02/1956, beach drift; Bird Key, Bis- cayne Bay, INSM 414387, 3 shells, Eolis Sta. 13, 2-10 ft., Henderson Coll. 1910; Florida Keys: Key Largo, AMNH 242626, 4 shells, ex-Alice Denison Barlow Col- lection; Little Molasses Reef, off Kev Largo, UF145851, McGintv 4/3/1950; Bonefish Key, UF 192101, 4 shells, Beal-Maltbie coll., Ex-McGintv Collection; UF 146530, 1 shell, McGinty Collection; UF 63202, 8 shells, M. Smith coll.. University of Alabama leg.; UF 162451, 3 shells, Blenn R. Bales, G.L. Warmke leg. 1989; UF 135603, 3 shells, V. Orr, H.G. Lee leg.; MCZ 118844, 10-H shells, B.R. Bales 5/7/1940; MCZ 100714, 8 shells, B.R. Bales leg.; AMNH 116412, 2 shells, 1940, Dr. and Mrs. Julius Wisoff Collection; AMNH 114121, 1 shell, T. McGinty coll.; AMNH 138995A, 1 shell, AS. Koto leg.; AMNH 189020, 4 shells, AS. Koto leg. 1949, J. M. Gate Collection; AMNH 138995, 4 shells, A.S. Koto leg., M.K. Jacobson Collection; AMNH 199169, 9 shells, AS. Koto leg.; Lower Matecumbe Kev, USNM 53477, 3 shells, H. Hemphill, in grass; Kev Vaca, UF 156266, 9 shells, McGinty 2/1939; MCZ 153269, 25+ shells, B.R. Bales; Key Vaca, Boot Key Harbor, UF 145853, 7 shells, McGinty coll. 3/1945; MCZ 226804, 2 shells, grassy bottom, 1 fm. McGinty coll.; No Name Key, UF 63203, 2 shells. University of Alabama leg.; Little Torch Key, UF 123072, 33 shells, M.C. Teskey 1968-1977, on sand patches; Grassy Key (Florida Bay), MCZ 110198, 4 shells, under rocks, P. McGinty leg.; Pumpkin Key, UF 63205, 2 shells. M Smith 1/1935, University of Alabama leg.; Islamorada. AMNH 121444, 3 shells, ex-E.C. Styles; Big Pine Key, UF 63204, 4 shells, M. Smith coll.. University of Alabama leg.; Key West, AMNH 8982, 3 shells. Constable, Jay Collection; USNM 53541, 1 shell, H. Hemphill, on rocks; Barraconta Key, 9 miles W of Key West, UF 13149, 2 shells, G.W. Van Hyning, 6/11/1958; Middle Sambo Shoals, UF 145852, 6 shells, McGinty 6/1946; Dry Tor- tugas: MCZ 258469, 1 shell, J.S. Schwengel, dredged; Loggerhead Key, UF 158168, 1 shell, T. McGinty 8/1941, 0.5 fm; Garden Key, South Coaling Dock, UF 13151, 2 shells, G.W. Van Hyning 7/16/1938, on piling; Gulf Coast of Florida: St. Martins Reef, near Aripeka Bird Racks, 6 mi. W of Aripeka, Pasco-Hernando Counts line, MC;Z 233672, 14 shells, W.A. Smith 5/25/1963; Ozone, Crystal Beach, AMNH 246038, 3 shells, Dan Steger coll., Jerome Bijur Collection; AMNH 125612, 10 shells, S. Levine leg. 1960; AMNH 189024, 5 shells, S. Levine leg. i960, J.M. Gate Collection; AMNH 125928. 76 shells, S. Levine leg. 1960; AMNH 194560, 66 shells, Gordon New- ell-Usticke Collection; Ozone, Crystal Beach, St. Joseph Sound, UF 193036, 27 shells, Dan Steger; UF 137573, 5 shells, D. Steger 1970. H.G. Lee leg.; AMNH 189023, 3 shells, mud and grass bottom, E. Marcott leg. 1963, J. Gate Collection; Marco, UF 137649, 1 shell, H. Hemphill, 2 fms, H.G. Lee leg.; USNM 53542, 1 shell, H. Hemphill; 15-35 mi. off Ft. Walton (FL), MCZ 145877, 5 shells, 13-19 fms., LA. Burry; Featherbend Bank, UF 63206, 2 shells, M. Smith 1/1935. University of Alabama leg; Okaloosa County, off Destin, UF 158166, 1 shell, UF 145857. 2 shells, McGinty 10/1941, 14 fms; Bahamas: Andros. first island off Mintie Bar, SE end of South Bight, USNM 271832, 7 shells, P. Bartsch; Abaco, Little Harbor, USNM 180492, 5 shells, O. Bryant; Grand Bahama Is- land: Dead Man's Reef, 26°34'45"N, 78°51'45"W, ANSP 371266, 1 shell. J. Worsfold, Ex-J. Worsfold; McLean's Town. 26°38'45"N,77°57'30"W. ANSP 369060. 8 shells. J. Worsfold, E.\-J. Worsfold; Eight Mile Rock, Hepburn Town, "Garbage Hole", 26°31'30"N, 78°47'15"W, ANSP 370392. 4 shells, J. Worsfold, Ex-J. Worsfold; Wood Cay, 26°44'15"N, 79°5S'15"W, ANSP 369615, 3 shells. J. Wors- fold, Ex-J. Worsfold; West End, Settlement Point, 26°42' 15"N, 78°59'50" W, ANSP 368626, 6 shells, J. Wors- fold, Ex-J. Worsfold; West End, Hotel Jettv, 26°42'15"N, 078°59'50"W, ANSP 368711. 21 shells, J. Worsfold, Ex- J. Worsfold; Nassau, New Providence Island, L'F 145849, 1 shell, McGinty 6/6/1947, 4-6 fms, Sta. 17; Cuba: Las Villas, Caibarien, Cayo Salinas, AMNH 138924, 1 shell, M.K. Jacobson leg., 07/1947; Esperanza (NW Cuba), USNM 414396. 5+ shells, Barrera Expedition, Sta. 210, 2-3 fms; Santa Rosa (NW Cuba), USNM 414498, 1 shell, Barrera Expedition, Sta. 209, 3-6 fms; Bermuda. AMNH 45068, 2 shells. Constable. Jav Collection; MCZ 24174, 2 shells, O. Brvant 1903; IISNM 223283, 6 shells, Havcock coll.; Haiti, Dept. de lOuest, Saltrois, USNM 439975, 1 shell, Orcutt, Chamberlain Coll. THE NAUTILUS 107(2);63-75, 1993 Page 63 Shell and Pallet Morphology of Early Developmental Stages of Bankia goiildi (Bartsch, 1908) (Bivalvia: Teredinidae) Antonieto S. Tan Ya-ping Hu Michael Castagna' Richard A. Luiz Michael J. Kennish Alan S. Pooley Rutgers University Institute of Marine and Coastal Sciences P.O. Box 231 New Brunswick, NJ 0S9U3-0231 USA ' Virginia Institute of Marine and Coastal Sciences College of William and Mary Wachapreague, VA 23480 USA ABSTRACT The margin of the lar\al shell ot Bankia gouldi (Bartsch) un- dergoes a progressive change m shape during ontogen\ from an initial D-shape, to nearly circular, and then to elliptical just prior to metamorphosis. The length of the iiinge is 37.9 ± 2 3 lim, n = 30 (range; 34.0-43.0 iim). The D-stage larval shell is inequilateral with the anterior end broader than the posterior end. The provinculum, composed of two cardinal teeth and three sockets in the left valve, and three cardinal teeth and two sockets in the right valve, is well developed in shells as small as 75 nm. The length of the central tooth in the right valve ranges from 12 to IQ/iim, The other teeth are one-half to one- third of the length of the central tooth. A lateral hinge s\ stem, consisting of two lateral flanges on the left valve that interlock with two lateral grooves on the right valve, is present in spec- imens as small as 75 nm. This hinge system persists until meta- morphosis. Metamorphosis, first seen in the development of a ligament pit, occurred in larvae with shells as small as 197 ^m in height; and leads to the development of the chondrophore, apophysis, anterior and posterior slopes, disc, umbonal-ventral ridge, dorsal cond\ le, ventral cond) le, denticulated ridges and pallets. As\nimetrical growth results in the formation of the characteristic form of the teredinid postlar\ al shell with a neu axis of articulation oriented dorsoventrally along the condyles. Each pallet consists of a proximal stalk and a distal blade witli one to several segments. Pallets with one segment were first observed in early postlarval shells at 0 4 nmi in length. Key iL'ords: larvae, postlarvae, shell morphology, Teridinidae, metamorphosis. INTRODUCTION Morphological classification schemes used in the iden- tification of bivalve larval stages have been developed from light microscopic examination of plankton (j0r- gensen, 1946; Rees, 1950) or cultured larvae (Loosanoff et al., 1966; Chanley & Andrews, 1971), More recently. the scanning electron microscope (SEM) has enabled workers to describe in detail the hinge and other minute structures of bivalve larvae and postlarvae that are useful in species identification (Turner & Boyle, 1975; Carriker & Palmer, 1979; Chanley & Dinamani, 1980; Le Pennec, 1980; Lutz et al., 1982a, b; Lutz, 1985; Prezant, 1990; Waller, 1991). A method of consistently orienting and documenting the shapes and dimensions of larval shells (Lutz et al., 1982b; Fuller et al., 1989b) has provided an additional means of differentiating bivalve species. The present manuscript is part of an ongoing effort to document the gross morphology, hinge apparatus and other morphometric features of larval and early postlar- val shells of bivalve mollusks to facilitate ta.xonomic iden- tification of individual specimens to the species level (Fuller & Lutz, 1989a; Fuller et al., 1989c; Kennedy et al.. 1991; Goodsell et al., 1992; Gustafson & Lutz, 1992). Bankia gouldi (Bartsch, 1908) is an oviparous mollus- can woodborer foiuid in tropical and temperate waters of the western Atlantic Ocean (Turner, 1971). As an adult, B. gouldi has a soft, vermiform body, a diminutive shell that covers the anterior-most portion of the body, and segmented pallets situated at the base of the siphons beneath a muscular collar (Turner, 1966, 1971). The pallets are not formed until after metamorphosis and, therefore, are of no value in identification of larval shells. In addition, previous descriptions of the larval shell, early postlarval shell and pallet of B. gouldi are insufficient to allow congeneric differentiation. Hence, a means of iden- tifying younger specimens of B. gouldi is clearly needed. This work presents both qualitative and quantitative data on the morphology of the larval shell, early postlarval shell and pallet of 6. gouldi that can aid in species iden- tification of early ontogenetic stages. In particular, we show that the following features are important in distin- guishing the larvae of B. gouldi from T. navalis: slope of the shoulders, length of provinculum/hinge-line, and length of posterior tooth ot the left provinculum. Page 64 THE NAUTILUS, Vol. 107, No. 2 Figure I. Inl.rnal surface vie-w of disart.culaU-cl larval sheik Antcru.r end of the left valve is oriented tc»vards Uie ■^''^^^-h';^; the anter.or ernl of the nght valve .s or.enled towards the left. The runnber above each valve s.gn.tes the maximum Imear shell dimension in ^in Tan et al.. 1993 Page 65 3 X o X CO 240 SHELL LENGTH (fim) Fif»iir«- 2. Linear regression oi shell height plotted ;i>;aiiist shell length ol larvae. Table I . Dimensions in fim of the larval shell of Bankia gouldi shown in Figures 1 and 2 1 SD = one standard deviation Sam- ple Range Mean ± 1 SD size Left Valve Anterior tooth length 7.5-9.4 8.3 ± 0.8 7 Posterior tooth length 5.5-7.1 6.4 ± 0.6 7 Provincular length 36.0-40.5 38.1 ± 1.8 9 Right Valve Anterior tooth length 3.9-4.9 4.5 ± 0.4 8 Central tooth length 12.5-15.5 14.5 ± 1.2 7 Posterior tooth length 4.5-5.9 5.0 ± 0.5 9 Provincular length 34.0-40.5 37.8 ± 1-9 9 MATERIALS AND METHODS Specimens of Bankia gouldi were collected from wooden frames of trays set out for a year in the York River at Gloucester Point, VA by Dr. Mary Gibbons. Adults were mass spawned, and the fertilized eggs w ere grown to the early adult stage using standard techniques (Loosanoff & Davis, 1963; Castagna & Kraeuter, 1984). The tem- perature of the culture water ranged from 22 to 30°C, and salinity, from 32 to 34 ppt salinity. Floating blocks of wood were provided as substrata for metamorphosing larvae. Larvae and postlarvae were sampled regularly and preserved in 95% ethanoi. The preparation of spec- imens for SEM examination was described previously by Fuller et al. (1989b). The specimens were coated with about 600 A gold-palladium using a Polaron Coating L'nit E5100, and they were examined using a Hitachi-450 SEM. Calibration of orthogonality of the SEM was done with the aid of metallic spheres or glass microspheres. Dimensions in the SEM were calibrated using a cali- brated ocular micrometer, a glass microsphere, or a grid of known dimension. Specimens were prepared, consis- tently oriented, and photographed as previously de- scribed to facilitate comparison of shapes and measure- ments (Lutz et al., 1982b; Fuller et al, 1989b, c). Shell terminology is described in Rees (1950), Turner (1966), Chaniey and Dinamani (1980), and Fuller et al., (1989c). The maximum linear dimension, shell length, shell height, provincular length, tooth length and the lateral hinge length were measured from caiibarated scanning electron micrographs of oriented shells. The shell height and length of the shell, hinge line and pallet were also measured using a compound microscope with the same calibrated ocular micrometer used to calibrate the scale of the SEM. These terms are defined in Fuller et al. (1989c). Whenever possible, we compared our results quanti- tatively with previously published data. Variables that changed with growth were compared using analysis of covariance, whereas variables that were independent of size were compared using the Student t-test, Tukey, or Student-Neuman-Keuls multiple range test (Zar, 1984). The larval shells in Figures 1 and 3 are referred to by their maximum linear shell dimension, which is the greatest distance between two points on a contour of a valve oriented in a standard fashion as described pre- viously. The early postlarval shells in Figures 5-7 are referred to by their shell height. Numbers separated by a slash (e.g. 57/62) indicate left and right valve dimen- sions respectively. RESULTS Larval shell: One da> after fertilization the shell length was 61.1 ± 4.5 Mill, n = 30 (range: 54-70 /um); shell height was 49.3 ± 4.5 ^ni, n = 30 (range: 43-59 ^m); hinge line/provincular length was 37.9 ± 2.3 fim, n = 30 (range: 34.0-43.0 fxm). Most valves of the straight- hinge larvae are inequilateral with the anterior end broader and protruding farther beyond the hinge line than the posterior end (Fig. 1, 57/62). The lar\al shell undergoes gradual allometric changes during ontogeny from a D-shape, to a nearly circular shape, and finally to an elliptical shape just prior to meta- morphosis (Fig. 1). During larval growth the maximum shell dimension shifts from an anteroposterior to a dor- soventral axis. Larval shells are almost ecjuidimensional (length = height) at approximately 143 ;um (Figs. 1, 2). The relationship between the length and height of the larval shell is described by the equation, Y = —29 92 + 1.26X (r^ = 0.97, n = 150), where Y = height and X = length in ^m (Fig. 2). A second power polynomial equa- tion (quadratic equation) fits the data points only slightly better than the linear regression equation. The provinculuin consists of two cardinal teeth and three sockets on the left valve that interlock with three cardinal teeth and two sockets on the right valve (Figs. 1, 3). The onset of provincular development is variable, occurring at about 57 ^.m. An individual at 57 /um can be in the prodissoconch I stage without any trace of a provinculum, or in the early prodissoconch II stage with a developing provinculum (Fig. 3, 57 ^ni). Development of the provincular teeth in the right valve starts with the formation ol the posterior tooth, followed by formation Page 66 THE NAUTILUS, Vol. 107, No. 2 Figure 3. Scanning electron micrographs showing hinges of lar\'ai shells seen in Figure 1. Tan et al.. 1993 Page 67 Figure i. Dursomedial \ ieu of the larval ihflls. Orientation as in Figure 1. The lateral hinge system consists of two lateral flanges on the left valve and two lateral grooves on the right valve on each end of the provinculum. Horizontal valve width: left = 198 nm, right = 185 ^m. of the anterior tooth, and cuhninating with the formation of the central tooth (Fig. 3, 62 urn, 75 ^m). The prov- inculum is well-developed in valves measuring 75 nm. The longest provincuiar tooth is the centra! one, \\ hich can reach 19 nm. The other teeth are either one-half or one-third of the length of the central tooth (Table 1). The left anterior tooth is consistentlv larger than the left posterior tooth, but the right anterior and right posterior teeth are not consistent!) different in size (Figs 1, 3). The increase in tooth length relative to shell length is variable: the total provinculum increases more sIovnIv on the right valve than on the left valve. Oni\ the length of the left anterior tooth regressed against shell length during grow th was significant (r- = 0.57, Table 2), others were not. The provincuiar length measured from scanning electron mi- crographs agrees with the hinge-line length values of articulated valves measured bv a light microscope. Lateral flanges and grooves are present in valves as small as 75 ^m (Figs. 1, 3), and persist until metamor- phosis. Two lateral flanges on the left valve (Fig. 1, 81- 214 ^m. Fig. 4) interlock with the two lateral grooves on the right valve (Fig. 1, 75-201 ^m. Fig. 4). Each lateral flange or lateral groove extends from the ends of the provinculum for about 1/10-1/15 of the perimeter along the dorsal commissure of the larval shell (Figs. 1, 4). The length of the anterior and posterior lateral grooves are equal in shells up to 109 nm (F"ig. 1), but in larger valves the anterior lateral groove increases in size reach- ing a length up to 12% greater than that of the posterior lateral groove (Figs. 1,4). Early postlarval shell: The first morphological evidence of metamorphosis is the development of a ligament pit beneath the posterior tooth and socket in valves mea- suring 235/230 jum in height (Figs. 5, 6). After meta- morphosis, the growth of the postlarval shell is very asym- metrical (Pigs. 5, 7, 8). The postlarval shell is slightly inequivalve, with the right valve longer than the left valve (Fig. 8). The ventral margin of the right valve is narrower than that of the left valve. The denticles on the denticulated ridge are predominantly two pronged, with the dorsal prong larger than tiie \ entral prong. The ventral margin grov\'s faster medially than the anterior shell margin, while the growth of the posterior margin lags behind (Figs. 5, 7, 8). The length to height ratio of Table 2. Linear regressions of the shell height, tooth length or pro\incular length plotted against shell length of Banhia gouldi shown in Figure 1. a = intercept, b = slope, r- = coefficient of determination, P = probabilitv of the re- gression coefficient (slope) of each regression, N = sample size. Dependent Variable a b r- P N Left Valve Shell height -31.42 1.27 0.98 <0.01 9 .\nterior tooth 5.88 0 02 0.57 0 05 7 Posterior tooth 6.66 -0.002 0.02 0.76 7 Provinculum 37.32 0.01 0.03 0 65 9 Right Valve Shell height -27.57 1.22 0.98 <0.01 9 Anterior tooth 4.89 -0.003 0.14 0.35 8 Central tooth 12.71 0.01 0.18 0.35 / Posterior tooth 5.36 -0.003 0.09 0.43 9 Provinculum 42,13 -0 04 0.71 <0.01 9 Page 68 THE NAUTILUS, Vol. 107, No. 2 Figure 5. Internal .siirfac- view ol early posllarval shells. Orientation as ni Figure I. The number above eaeh valve signifies the shell height measured in nm roughlv perpendieular to the hinge region. The number on the lower left of the valve is the she^l length in fim measured roughly parallel to the hinge region. ,'\ = apophysis. UC: = dorsal eondyle, VC = ventral condvie, R - nmbonal-ventral ridge. Tan et al., 1993 Page 69 Figure 6. Scanning electron micrographs of the hinge region of earl\ postlarval shells seen in Figure 5. The number above each valve signifies shell height in ^m, LP = ligament pit, C = chontlrophore. Page 70 THE NAUTILUS, Vol. 107, No. 2 Figure 7. Scanniiif; i'l«lroii iiiicrograplis ot l\w cxU-riial surface (il t-arl\ pusllarval shells, OrieiilatHni and number representation a.s in I'^igure 5. AS = anterior slope, FS = Posterior slope. Tan et al, 1993 Page 71 Figure 8. Views from different directions of partially articulated early postlarval sfiells with two denticulated ridges. The anterior (top left) and the posterior (top right) views show that the right valve (R) is slightly longer than the left valve. The ventral view at the bottom shows that the ventral margin of the right valve is narrower than that of the left valve. Most teeth have two prongs. The dorsal prong of the tooth is larger than the ventral prong. Horizontal valve widths are as follows top left valve = 235 ixm. top right valve = 243 nm, bottom left valve = 225 m"'. bottom right valve = 239 ^m. the postlarval shell continues to decrease with growth (Figs. 9). The chondrophore that forms at the base of the pos- terior tooth and socket grows anteroventrally then me- chally, assuming a recurved formation (Figs. 5, 6). A saber-like apophysis develops beneath the anterior tooth and socket, and elongates along the sagittal plane in a posteroventral direction (Figs. 5. 6 height 255/235 Mm) The dorsal portion of the posterior slope and the anterior slope are reflected up and outward (Figs. 5, G, height 305-1140/290-1140 Mm). The base of the apophysis and the adjacent reflected part of the anterior slope and umbo are incorporated to form the dorsal condyle (Figs. 5, 6, height 305-1140/290-1 140Mm). The enlarged chondro- phore and dorsal condyle eventually conceal the pro- vinculum. As growth proceeds, the ventral portion of the disc and the anterior slope curves ventromedially (Figs. 5, 7, 8). The shell thickening at the medial side of the ventral margin eventually develops into the ventral con- d\le (Fig. 5, height 255/235Mm). An umbonal-ventral ridge, e.xtending froin the larval shell to the ventral con- dyle, forms on the inner shell surface (Fig. 5, height 335- 11 40/705- 11 40Mm). The umbonal-ventral ridge is visible as an umbonal-ventral sulcus on the external shell surface (Fig. 7, 415-970/355-1075 Mm). The posterior slope (au- ricle) overlaps the disc forming the shelf. The asym- metrical growth of the shell eventually results in the formation of the characteristic form of the teredinid Page 7S THE NAUTILUS, Vol. 107, No. 2 B J. X o Ed a to 2.5 2.0 1.5 1.0 0.5 0.0 30 r Y = 0.15 + 1.00 • X r2 = 0.84. n = 41 0.0 0.5 1.0 1.5 2.0 SHELL LENGTH (mm) Figure 9. Rflation.sliip between sliell length and .shell height of early postlarval shells. w o Q K El, o K Cd m S D SHELL LENGTH (mm) Figure 10. Relationship between shell length and number of denticulated ridges on the exterior surface of early postlarval shells. postlarva, with the a.\is of articulation changing from an anteroposterior to a dorsoventral orientation along the condyles. Early in the formation of the postlarval shell, dentic- ulated ridges are added onto the anterior margin of the shell, initially only on the anterior area of the anterior slope (Fig. 7, height 250-280/240-250 urn). As the shell grows larger, the denticulated ridges extend to the an- terior median area of the anterior slope. The number of denticulated ridges increases significantly with the in- crease in shell length; regression equation, Y = 5.38 + 10.7X, (r- = 0.54, n = 41), where Y = number of ridges and X = shell length in millimeters (Fig. 10). Pallet: A one segment pallet was present at a shell length of 0.4 mm; at a shell length of 1.2-1.3 mm, the pallet had 13 segments. The pallet consists of a proximal stalk and a distal blade with one to several segments (Fig. 11), with the basal pallet segment being the largest. The angle formed by the base of the blade and the stalk is larger on the dorsal than the ventral side. The medial inner wall of the blade is flat, while the shorter outer wall of the blade is convex. The plane circumscribed by the distal outer margin of the blade is tilted anteriorly such that the distal border of the inner margin extends further posteriorly. Each pallet appears hemispherical uhen viewed from the distal end. The left and right pallets in an individual are morphologically similar and have the same number of segments (Fig. 11, height 250-505 jum), v\ hereas pallets with the same number of segments from different individuals can vary in form (Fig. 11, height 840-1415 Mm). DISCUSSION Larval shell: Teredo navalis Linnaeus, 1758 is the only other teredinid species whose larval and postlarval shell morphology has been oriented and photographed in a consistent manner (Fuller et al.. 1989c). In lar\al shells of similar size, the shoulders of T. navalis (Fig. 1, Fuller et al., 1989c) are considerably steeper (less rounded) than those of Bankia gouldi (Fig. 1). The shoulders refer to the dorsal shell margin on the anterior and posterior ends of the hinge (C'hanley & Dinamani, 1980). This differ- ence in larval shell shape can be used to distinguish the two species. On the other hand, the size of the specimens at which the length approximates the height measure- Table 3. Linear regression equations for the shell length and shell height of D-stage to recent postmetamorphic larvae, r- = coefficient of determination, P = probability for the slope of each regression, N = sample size. Slopes with different superscripts are significantly different (P < 0.0.5) by a Tuke\ multiple range test Species Intercept Slope r- P N .\uthor Teredo natxilis' -49.03 1.40'^ 0.99 <0.01 10 Fuller ('/ al. (I9S9c) Bankia sctacea -22.50 1 14" LOO <0.01 8 Turner & Johnson (1971) liankiu f^onldi -.38.72 L.34^ 1.00 <0.0J 6 Culliney (1975) liankia gouldi -36.13 1.37^ 0.99 <0.01 7 Mann & Gallager (1985) Rankia gouldi' -.32.. 56 1.28^ 0.98 <0.01 10 This stud\- Data used to calculate the linear regression equations are derived measurements of the left \ alve of scaiuiing electron micrographs. Analysis of covariance of the slopes is significant (F-statistic = 6.32, P < 0.001). Tan et al., 1993 Page 73 Figure 1 1. Outer surface view of the pallets. The pallet length is the distance between the proximal (stalk) and distal end (hiade). Numbers signify the average length of the left and right (H) pallets. Dorsal side of the pallet is oriented towards the top, I = inner wall, O = outer margin of outer wall. Page 74 THE NAUTILUS, Vol. 107, No. 2 nients can not bo used to differentiate B. gaiildi from T. nat^alis. The length and height of the larva! shell of B. goiildi are nearly equal at about 143 ^im, far beyond the point where the umbo is first visible in a shell in our standard orientation, at about 128 ;um (Fig. 1). Cailliney (1975) as well as Mann and Gallager (1985) reported that B. gouldi is equidimensional at 128 /xni ^nd 71.4 ^m, respectively. This difference (144 nm vs 128 or 71.4 /um) is probably due to foreshortening (shells tilted) in the earlier studies. Fuller et al. (1989c) reported that T navalis achieves «iuidinieiisionalit\ at 150 /im. The ini- tial size of a B. goiiUIi larva, is much smaller than that of the larviparus T. navalis due to differences in their development. Despite the difference in initial larval shell size, both species metamorphose at a shell height of about 230 ^m, as indicated b\ the appearance of the ligament pit. Table 3 shows that shell height increases faster than shell length until just before metamorphosis in Bankia goiildi (Culliney, 1975; Mann & Gallager, 1985), Teredo navalis (Fuller et al., 1989c), and Bankia setacea (Tryon, 1863) (Turner & Johnson, 1971). The slopes are signifi- cantly different (P = <0.001) as shown by an analysis of covariance. Moreover, a Tukey multiple comparison test reveals that the slope for B. setacea is significantly lower than the other four species (P = <0.05, Table 3). The data for B. gouldi (this study) and T. navalis (Fuller et al., 1989c) are from measurements of SEM micro- graphs of disarticulated left valves, whereas the data from other studies are based on light microscope mea- surements of articulated valves (Culliney, 1975; Turner & Johnson, 1971; Mann & Gallager, 1985). We have included the data from shells of recent postmetamorphic larvae since the ligament pit, an earl) indicator of meta- morphosis, is difficult to detect by the previous studies. The larvae of Bankia gouldi and Teredo navalis can be distinguished on the basis of the length of the prov- inculum on the left valve. The average length of the provinculum/hinge-line measured from micrographs of the left valves of B. gouldi (38. 1 ± 1.8^ m, n = 9; Table 1) and T. navalis (46.7 ± 1.3 ^m, n = 9; Fuller et. al., 1989c) are significantly different (Student t-test, P < 0.001). Culliney (1975) reported a similar difference be- tween the hinge-line length of B. goiddi (40.4 ± 3.96 Mm, n = 35) and T. tiavalis (51.3 ± 2.19 Mm, n = 47). The same conclusion is reached using a simple size of 21 for B. gouldi. The provincular length measurements of the left valve are more useful than those of the right valve, because the latter varies significantly with growth. The larvae of Bankia gouldi and 6. inartensi (Stem- pell, 1899) can be distinguished from Teredo navalis In the length of the posterior provincular tooth of the left valve. The left posterior tooth of B. gouldi (6.4 ± 0.6 Mm, n = 7) and B. martensi (6.4 ± 1.0 Mm, n = 10) are significantK shorter that that of T. navalis (8.2 ± 0.3 Mm, n = 9). Clo.se examination of the internal shell surface of the valves of Teredo navalis (see F'ig I; Fuller et al., 1989c) reveals a lateral hinge system similar to that in Bankia gouldi The longer anterior component of the lateral hinge system in larvae of B. gouldi longer than 109 Mm appears to be due to slightly faster growth of the anterior shell margin relative to the posterior shell margin. This difference may be useful in differentiating the two spe- cies. Postlarval shell: Larvae of Bankia goiddi and Teredo navalis can be distinguished by the number of ridges vs. shell length in shells greater than 1 mm in length. Both species develop a denticulated ridge at about 0.24 mm in length. Denticulated ridges are added at about the same rate in both species until the shell reaches 1 mm. Subsequently, the number of denticulated ridges in T. navalis exceeds that in 6. gouldi. For example, at a shell length of 2mm, B. gouldi has 13 ridges, compared to about 28 ridges in T. navalis. In the early larval stage ot B. gouldi, the right valve is longer than the left valve (Fig. 8), but the degree of asymmetry is less than that in T. navalis. The shape of the later, postlarval shell varies greatly with the type of wood used as a substratum by the teredinids (Turner, 1966; Turner & Johnson, 1971 ), and the environmental conditions at the collection site (Hill & Kofoid, 1927). Pallet: The morphology of the adult pallet is important in identification of teredinid species (Turner, 1966; 1971). In Bankia setacea, the pallet first appears at an early postlarval shell length of 0..5-0.6 mm (Quayle, 1959). In B. gouldi, the smallest individual with a pallet had an early postlarval shell length of 0.4 mm. The pallet at this early developmental stage is unsegmented and may be confused with the nonsegmented pallet of other teredi- nids. Caution is advised in using pallet morphology as a taxonomic tool at this developmental stage. ACKNOWLEDGEMENTS We thank Mr. John Grazul for technical ad\ice related to SEM, and Drs. R. Prezant, R. Tool, D. Marelli and B. Kotrla for insightful discussions on several aspects of B. gouldi biology. This work is supported partly by a post- iloctoral fellowship grant to A. S. Tan from the Fisheries and Aquaculture Technology Extension Center, Rutgers L'niversity. Institute of Marine & Coastal Sciences con- tibution no. 93-14; New Jersey Agricultural Experiment Station contribution no D-324()2-l-93; Virginia Institute ot Marine Science contribution no. 1778. LITERATURE CITED ( :arrikfr. M. \\ and R K. Palmer. 1979. llltrastructural nior- plHigciicsi,s of prodis.socoiich and carl) dissocoiicli \ai\es of the oyster Crassostrea virginica. Proceediniis of the National Shellfisheries Association (•)9: 10.5- 128 Castagiia, M and J N. Kraeuter. 1984. Maiivial for growing the hard clam Merccnaria. Special Report in .Applied Ma- rine .Science and Ocean lOngineering No, 249 Virginia lii.slitiitc of Marine Science, Gloucester, X'irgiiiia, 1 Id p. (:liaiilc\, 1' and J I) .Viuirews, 1971. Aids for identificalion of l>i\al\t' lar\;ieof \'jigliiia Mahn'ologia 11( 1 ):4.5-l 19. Tan et al., 1993 Page 75 Clianley, P, ami P Dinaniaiii 19S() CoinparatiNc ilescrip- tions of some oyster larvae from New Zealand and (IhiJe, and a description of a new genus of o\ster, Tio.slrca. New Zealand Journal of Marine and Freshwater Research 14(2): 103-120 Culline\ , J L. 1975 (Aimparati\e lar\al de\elopmeiit ol the shipworins Banhia gouldi and Teredo nuialis Marine Bi- ology 29:245-25 L Fuller, S. C. and R. A. Lutz. 1989a. Shell morpholog) ot larval and post-larval mytilids from the North-Western Atlantic. Journal of the Marine Biological Association of the United Kingdom 69:181-218. Fuller. S. C, R. A. Lutz and A. Poole\. 1989h Procedures for accurate documentation of shapes and dimensions of larval bivalve shells with scanning electron niicroscopv. Transactions of the American Microscopical Societ\' I()8( 1 ): 58-63. Fuller, S. C, Y. Hu, R. A. Lutz and M. Castagna. 1989c. Shell and pallet morpholog\ in early developmental stages of Teredo navalis Linne (Bivalvia: Tereiiinidae). The Nau- tilus 103(1 ):24-35. Goodsell, J. G., S. C. Fuller, A. G. Eversole, M C^astagna ami R .A Lutz. 1992. Larval and early post!ar\al shell mor- pholog\ of se\eral \enerid clams Journal of the Marine Biological .Association of the United kingdom 72231-255 Gustafson, R. G. and R. A. Lutz 1992. Larval and earl) post- larval development of the protobranch bivalve Solemija velum (Mollusca: Bivalvia). Journal of the Marine Biolog- ical Association of the United Kingdom 72:383-402. Hill, C. L. and C. A Kofoid (eds.) 1927 Marine borers and their relation to marine construction on the Pacific coast Final Report of the San F'rancisco Ba\ Marine Piling Gom- mittee. Universits of Galifornia Press, San f'^rancisco, 357 p j0rgensen, C. B. 1946. Reproduction and larval development of Danish marine bottom invertebrates with special ref- erence to the planktonic larvae in the Sound (0resund), Lamellibranchia. Meddelelser fra Kommissionen for Dan- marks Fiskeri-og Ha\unders0gelser, Serie: Plankton 4:277- 311. Kennedy, V. S., S. G Fuller and R A Lutz 1991. Shell and hinge de\elopment of \oung Corbicula flumiiica (Miiller) (Bivalvia: Gorbiculoidea). .American Malacological Bul- letin 8(1991):107-111. Le Pennec, M. 1980. The larval and post-larval hinge of some families of bivalve molluscs. Journal of the Marine Bio- logical Association of the United Kingdom (iO:601-617. Loosanoff, \'. L. and H. C. Davis. 1963. Rearing of bivalve molluscs. In Russell, F.S. (ed.). Advances in Marine Bi- ology. Academic Press, New York, 410 p Loosanoff, V. L., H. C. Davis and P. C^hanlcN 1966 Dimen- sions and shapes of larvae of some marine bivalve molluscs. Malacologia(4):351-435. Lutz, R. A. 1985. Identilic.ilion ol bi\aKc lar\ae and postlar- vae: a review of recent ad\ances. .American Malacological Bulletin, Special Edition No. l(1985):59-78. Lutz, R. .\ . J. Goodsell, M (,'astagna. S CJhapman, (.'. Newell, H Hidu. R Mann, D Jablonski. \' Kennedy, S Sidall, R, Cioldberg, H Beattie. G. Falmagne, A Ghestnut and A. Partridge 1982a. Preliminary observations on the use- lulness of hinge structures for identification of bivalve lar\ae Journal of Shellfish Research 2(l);65-70. Lutz. R A , R Mann. J G Goodsell and M. Castagna. 1982b. Larval and earl\ post-larval development of Arctica is- landica. Journal of the Marine Biological Association of the United Kingdom 62:745-769. Mann, R. and S. M. Clallager 1985. Physiological and bio- chemical energetics ol larvae Teredo navalis L. ami Ran- kia gotddi (Bartsch) (Bivalvia: Teredinidae). Journal ot Experimental Marine Biology and Ecology 85:211-228. Prezant, R S. 1990 Shell and hinge ontogeny of the Antarctic bivaKe Lissarea notoreudensis. Journal of the Marine Bi- ological .Association of the L'nited Kingdom 70:841-855. Qua\ le, D. B 1959. The early de\elopment of Bankia setacea (Trvon) In Ra\. D.C:. (ed.). Marine Boring and Fouling Organisms. Uni\ersitv of Washington Press, Seattle, p. 157- 174. Rees. C' 11 1950 The identilit'ation and classification of la- mellibranch larvae Hull Bnlletin of Marine Ecology .3(9): 7.3-104. Turner, R D. 1966 Sur\e\ and illustrated catalogue of the Teredinidae (Mollusca: Bivalvia). Museum of Gompara- ti\e Zoology. Harvard University. Gambridge, 265 p. Turner, R D, 1971, 1, Identification of marine wood-boring molluscs. In Jones, E B (;. and S.K. Eltringham (eds.). Marine Borers. Fungi and P"ouling Organisms of Wood. Organisation for Economic and Go-operation and Devel- opment, Paris, p. 17-65. Turner, R. D. and A. G. Johnson 1971. 13. Identification of marine wood-boring molluscs. In. Jones, E.B.G. and S.K. Eltringham (eds. ). Marine Borers. Fungi and Fouling Or- ganisms of Wood. Organisation for Economic and Co- operation and Development, Paris, p. 259-301. Turner, R D and P. J. Boyle. 1975 Studies of bisalve using the scanning electron microscope and critical point drying. Bulletin of the .American Malacological L'nion for 1974, p. 59-65. Waller, T R. 1991. Evolutionar\ relationships among com- mercial scallops (Mollusca: Bivalvia: Pectinidae). In. Shumway, S.E. (ed.). Scallops: Biology. Ecology and Aqua- culture Elsevier, New York, p. 1-132 Zar, J.H. 1984 Biostatistical Analysis. Prentice-Hall, Englewood Cliffs, New Jersev, 718 p. THE NAUTILUS 107(2):76-78, 1993 Page 76 A Simple Method for the In Vitro Culture of Perkinsiis marinus S.J. Kleinschuster S.L. Swink Haskiii Slifllfisli Kfst'arcli Lalioralory Department of Marine and Cxiastal Sciences Instilnte of Maine and (Coastal Sciences Ruti;ers UniversiU Box' B-8 1 Miller A\ciiuf Port Norris, NJ 08349 USA INTRODUCTION The oyster pathogen Perkinsun marinus Levine, 1978 is an apiconiplexan protist tliat has caused sul)stantial mor- tality among oyster populations along tiie mid-Atlantic and Gulf coasts of the United States and may be capable of cross-transmission among certain bivalve species (Gog- gin et al., 1989). The life c\cle ot the organism has been described previously (Mackin & Boswell, 1956; Perkins, 1966; Perkins & Menzel, 1966). Although many stages of the life cycle have been addressed, little evidence relative to mode of infection, infiltration and invasion has been described under in vitro conditions. Such in- formation would provide insight into many hitherto unobservable aspects of the host/parasite relationship in vivo. This study demonstrates that certain vegetative re- productive aspects in the life cycle of the organism, nor- mally occurring in the host, can be duplicated under relatively simple if! vitro conditions. MATERIALS AND METHODS Parasitic cells were isolated from primary cultures of tissue explants of visceral ganglia of Crassostrea virginica Gmelin, 1791 and identified as Pcrkinstis ntarinns using the technicjue of Ray (1952). Eoilowing isolation of the protist, various mixtures of Leibowitz's (1952) medium (L-15), oyster hemolymph, fetal bovine .serum (FBS), various sugars, yeast extract and lactalbumin hydrolysate were tested for cell growth potential. The osmolaritv of each licjuid component was adjusted to 750 mOs/kg by the addition of sea salts. The pH of each medium and/ or constituent was adjusted to 7.6 and filter sterilized. (Cultures were grown at 28°C; in rilasks utider ambient CO2/O2. Penicillin G (100 U/ml) and streptomycin (0.1 mg/ml) were routinely added to all cultures and the medium (50%) exchanged weekly. Mantle tissue lor challenge experiments w as from I'er- hinsits-hee o\sters which were routinely biopsed and maintained in isolated aciuaria. Sterile explants were ob- tained by culturing the tissue for 4 days at 25°C in a 1:1 sterile sea water/hemolymph mixture supplemented with penicillin G (100 U/ml), streptomycin (0.1 mg/ml) and amphotericin B (0.25 ;ug/ml). RESULTS Healthy explant cultures of visceral ganglia of Crassos- trea virginica are seen in figures 1 and 2. Similar explant cultures being parasitized by Perkinsiis marinus are seen in figures 3 and 4. Colonization of large and small groups of oNster nervous tissue is readiK apparent as is the at- tachment of parasitic cells to individual oyster cells. Var- ious media preparations to be evaluated for optimal growth were inoculated with parasitic cells from similar cultures. Approximately two weeks after inoculation, it w as evident that all media preparations evaluated would support cell growth to varying degrees. Cultures con- taining a high percentage of hemolymph displayed vig- orous and rapid propagation (figures 5 and 6). Cells cul- tured with a high percentage of L-15 and/or FBS (50%) tended to differentiate into cells that resemble prezoospo- rangia (figure 5, double arrows) (i.e., large cells greater than ca. 20 um with the cellular \ olume consisting mostly of a vacuole anil the c\ toplasin occup\ ing the perimeter of the cell; as described by Perkins and Menzel, 1966). Many non-flagellated daughter cells were formed in those cells xielding the morula-like cell aggregates typical of Pcrkinsus marinus. L'pon rupture of the mother cell wall the daughter cells are released into the mediiun (figures 5 and 6, arrows). Shortly after release of the daughter cells, until enlargement to form prezoosporangium-like cells, thr\ had the ts pical Perkinsus Hiori/n/.s-like meront structure consisting of a prominent, eccentric vacuole (figure 6, double arrows) with a single vacuoplast (figure 6, V arrow) and a single nucleus located near the cell wall. The most successlul medium [ireparation, yielding rel- ativt'K large numbers oi cells as well as a diversity of S. J. Kleinsclnister and S. L. Swink, 1993 Page 77 >- H ^\v P fX N -// v^ / /. S V ^-^.:. -> N Figures 1-2. Photomicrographs of health\ visceral ganjihon cxplaiits from Crassostrea virginica. Neurite growth extending from explants indicated by arrows, scale bar = (I 1 rmn Figures 3-4. Photomicrographs of oyster ganglion explants parasitized by Perkinsus marinus (P). Notice individual neurons (N), support cells (S), hemocytes (H) and attachment of parasitic cells to individual oyster cells (arrows), scale bar = 0.05 mm. life cycle stages consisted of a 100 mi. solution of L-15 containing 10.0 ml. of FBS, 20.0 ml. of o\ster hemo- lymph, 5.0 mg. taurine, 50.0 mg. glucose, 30.0 mg. ga- lactose, 50.0 mg. fructose, 50.0 mg trehalose, 100.0 mg. yeast extract, 300.0 mg. lactalbumin Inclrolysate, 1.0 ml MEM N'itamin solution (100X)(Sigma C^hemical C^o.) and 0.1 ml lipid nnxture (1000X)(Sigma Chemical C:o.). Cul- tures were routinely sub-cultured over a period of several months Sterile mantle tissue was challenged with cul- tured Perkinsiis marinus and became infected within 2-3 weeks. In fluid thioglycollate medium the parasite cells from challenged tissue formed cells resembling the prezoosporangia (or "hypnospores") normally seen in oyster tissue infected with Pcrkin.stts marinus. The cells also stained like Pcrkiustis marinus prezoosporangia in Lugol's iochne solution (Rax, 1952). Page 78 THE NAUTILUS, Vol. 107, No. 2 rj!^U\ -tx. ■pT-^S %^ :-v^(S ^" ''<'^* ''^^ •*t' il^\ <^'. 6 v-'at Figures 5-6. Photomicrographs of groups of propagative cells of Perkinsus marinus in vitro. Larger groups represent several generations. Figure 5. Notice presporangial cells (double arrows) and non-flagellated daughter cells (single arrows). Figure 6. Perkinsus-\\ke cells with nieront structure (double arrows), vacuoplast (V arrow) and non-flagellated daughter cells (single arrows), scale bar = 0.05 mm. DISCUSSION It is hoped that the technique described herein will con- tribute to the understanding of the basic biology of this parasite as well as to the host/parasite relationship. Re- finement of culture conditions and further definition of nutritional rec}uirements of the parasite in vitro would be particularly helpful as cellular or molecular approach- es to this understanding are attempted. ACKNOWLEDGMENTS The authors wish to thank Dr. F O. Perkins for his invaluable assistance with this study . This work was fund- ed by the New Jersey Agriculture E.xperiment Station, Hatch Project number 32100 and is identified as paper no. D-.32100-1-93 and as paper no. 93-09 for the Institute of Marine and Coastal Sciences. LITERATURE CITED Goggin, C. L., K.B Sewell and R T.G. Lester. 1989. Cross- infection experiments with Australian Perkiitsus species. Diseases of Aquatic Organisms 7(l):55-59. Liebowitz, A. 1963. The growth and maintenance of tissue cell cultures in free gas exchange with the atmosphere. American Journal of Hygiene 78:173-183. Mackin, J. G. and T. L. B Boswell. 1956. The life cycle and relationships of Dermocijstidiutn marinum. Proceedings of the National Shellfish Association 46:112-115. Perkins, F. O. 1966. Life History Studies of Dermocystidium marinutu, an Oyster Pathogen, Dissertation, Florida State University, Tallahassee, FL. Perkins, F. O. and R. W. Menzel. 1966. Morphological and cultural studies of a motile stage in the life cycle of Der- mocystidium marinum. Proceedings of the National Shell- fish Association 56:23-30 Ray, S. M. 1952. A culture technicjue for the diagnosis of infection with Dermocystidium marinum Mackin, Owen and CA)llier in Ovsters. Science 116:360. THE NAUTILUS 107(2):79-80, 1993 Page 79 On the Identification of Fossil Terrestrial Gastropod Eggshells Harold G. Pierce Research Associate University of Nebraska State Museum Lincoln, NE 68588-0514, USA Small (~1 mm), calcareous spheroids, often olilate, are frequently recovered associated with iionmarine mol- lusks from Quaternary localities on the High Plains. In general, these eggshells of terrestrial gastropods have been ignored by modern paleontologists. Tompa (1976a) cor- rectly correlated fossil eggshells from two middle Pleis- tocene localities in Kansas with the genus Vallonia. Tom- pa followed with a benchmark paper (Tompa, 1976b) in which he illustrated the eggshells of many ta.xa and pro- vided a catalog of terrestrial species with partially or wholly calcified eggshells. Improved, and gentler, prep- aration techniques have resulted in encountering both a greater abundance and frequency of these gastropod eggshells, especially in material of mid-Pleistocene to Recent age. Recently, I recovered a substantial number of much older (21-28 Ma) gastropod eggshells from the Oligocene-Miocene Cabbage Patch fauna of southwest- ern Montana. Since these eggshells were associated with a prolific and diverse terrestrial gastropod fauna (Pierce, 1992), an attempt was made to identify the taxa that produced these eggshells. In almost all cases, microscopic examination by trans- mitted light, and careful dissection of some specimens, has shown the fossil eggshells to be empty. Correlation of the eggshells with the various taxa in the fauna can be accomplished by comparison of hatchling size and shape to the size and shape of the eggshells. The two types of Cabbage Patch eggshells were both oblate sphe- roids that differed primarily in size (Table 1 ). The smaller eggshell matched very closely, in all dimensions, mea- surements of the hatchlings of Vallonia berryi Pierce, 1992, from the same fauna. The larger eggshell type was found to be marginally large enough, but too oblate, to accommodate hatchling of Oreohelix brandi Pierce, 1992, another component of this fauna. This match was also rejected on the basis that modern Oreohelix are ovivi- parous. Hatchlings of another species, Polygyroidea montivaga Pierce, 1992, were a near perfect match, both in size and shape. No other taxa of this fauna had hatch- lings that corresponded to the measurements of these eggshells (Pierce, 1992). Current work involving the Skull Creek locality in Butler County, Nebraska (late Pleistocene: Wisconsinan: Peoria loess, ca. 20 Ka), provided a new collection of gastropod eggshells. The Skull Creek material contains at least two ciuite diilerent eggshell forms (Table 1). The first, an oblate spheroid, H/W = 0.68, was very thin and delicate, and is represented by only 8 specimens, half of which are broken. In size and H/W, it compared closely with the eggshells attributed to Oligo-Miocene Vallonia berryi, and to mid-Pleistocene to Recent Vallonia pul- chella (Mijller, 1774) (Tompa, 1976a). Measurements of hatchling gastropods recovered as part of the Skull Creek locality fauna showed that hatchlings of Vallonia gra- cilicosta Reinhardt 1883, averaging 0.7 mm x 0.55 mm, were an excellent fit, in both size and shape, for this eggshell (Figures 1-3). The second eggshell type is a near spheroid, H/W = 0.93 or greater, and occurs in two slightly overlapping size ranges, each with distinct modes. The larger eggshell of this type is quite abundant (>50) (Figures 4-6). This eggshell size and shape was found to be an approximate fit for hatchlings of three taxa found in this fauna, all of which are known to have calcified Figures 1-3. Small fossil eggshells and hatcliling of Vallonia gracilicosta Reinhardt, 1883. \. Eggshell with crack and small fragment missing, lot PM.SC 1002 2. Hatchling \'. gracilicosta. lot P\ AL 1002 3. Broken eggshell, lot PMSC 1002. Figures 4-6. Large eggshell with embryo and hatchling ot Discus uhitneyi (Newcomb, 1864). 4. Hatchling D. whitneyi. lot PDIS 1004. 5. Large eggshell, lot PMSC 1001 6. Embryonic D whitneyi still retaining portions of its eggshell, PDIS 1016 ,\ll figures 20 X, all from Skull Creek locality, Butler County, Nebraska. Page 80 THE NAUTILUS, Vol. 107, No. 2 Table 1. Mea-suremt'iits (in mm) of fossil gastropod eggshells Eggshells \Vidtli/SD Range Height/SD Range H/W Oligo-Miocene taxa Cabbage Patch mollustan fauna Vallonia berryi Polygyroidea montivaga Late Pleistocene ta.xa Skull Creek molluscan famia Vallonia gracilicosta Discus ichitneyi Discus whitneyi? small 10 0,76 ± 0.03 0.57 ± 0.02 0.75 0.72-0.80 0.55-0.60 1(1 L60 ± 0.03 L24 ± 0.04 0.78 L55-I.65 1.15-L.30 4 0.80 ± 0.00 0.55 ± 0.00 0.68 0.80-0.80 0.55-0.55 10 1.20 ± 0.03 1.13 ± 0.01 0.94 1.15-1.25 1.12-1.15 5 1.03 ± 0.05 0,96 ± 0,04 0.93 0 95-1 10 0 90-1 00 eggshells (Tompa, 1976b, table 1): Nesovitrea hammonis electrina (Gould 1841), averaging 1.15 mm x 0.75 mm, Zonitoides arboreus (Say 1816), averaging 1.1 mm x 0.8 mm, and Discus whitneyi (Newcomb 1864), averaging 1.1 mm X 0.8 mm, with the latter two providing the best fit. [Roth (1987:129) has shown D. whitneyi to be a senior synonym of the well known D. cronkhitei (New- comb, 1865)]. By chance, one eggshell contained a em- bryo, identifiable as D. whitneiji (Figure 6). Roth V. gracilicosta and D. whitneyi are very abundant (>200 specimens each) components of the Skull Creek mollus- can fauna. Zonitoides arboreus and N. hammonis elec- trina are less common (16-50 specimens each) compo- nents. A second, smaller set (N=5) of the nearly spheroidal eggshell type, microscopically indistinguishable from the larger size, could not be matched precisely with any other taxon, but would fit the extreme minimum hatchling size of both D. whitneyi and N. hammonis electrina. Tompa (1976b:867) has shown that, for some taxa, eggshell size varies from individual to individual, and/or with size of parent animal. It appears probable, therefore, that these also represent D. whitneyi eggshells, but from small par- ents. Identification of eggshell types provides yet another tool useful in reconstructing fossil faunas. As more egg- shells are identified to genus and/or species, it may be- come possible that additional taxa, represented only by eggshells in a fauna, can be identified, thereby increasing the recognized diversity of the fossil fauna and the ac- curacy of paleoenvironmental reconstruction. However, with rare exceptions rc()uiring SEM examination (Tompa 1976b), identification beyond generic level is, at this time, often tenuous. LITERATURE CITED Gould A. A. 1841, Report on the Invertebrata of Massachu- setts, comprising the Mollusca, Oustacea, Annelida, and Radiata. Foisom, Wells, and Thurston, Cambridge. 373 p,, 213 figs, Miiller, O. F. 1774. Vermium terrestrium et Duviatilum, seu animalium Infusorium, Heiminthicorum et Testaceorum non marinorum succinta historia. Volumen .Alterum Hei- neck et Faber, Haviniae et Lipsiae. 214 p. New comb, W 1864 Descriptions of nine new species of Helix inhabiting California Proceedings of the California .Acad- emy of Sciences (1) 3:1 1.5-119. Newcomb, W. 1865. Descriptions of new species of land shells. Proceedings of the California Academy of Sciences (1)3:179-182. Reinliardt, O 1883. Sitzungsberichte der Gesellschaft Na- lurforschender Fruende, Berlin, 1883(2):42 Holh, B. 1987, Identifications of two (;alifornian land mollusks tlescribed bv VVeslev Newcomb Malacological Review 20: 129-1.32, Pierce, H, G, 1992. The nonmarine mollusks of the late Oli- gocene-early Miocene Cabbage Patch Fauna of western Montana. II. Terrestrial gastropod families other than Pup- illidae (Pulmonata: Stylommatophora). Journal of Pale- ontology 66(4):618-62S. ■Say, T. 1816. Conchology, Nicholson s British Encyclopedia of Arts and Sciences, 1st (American) ed. unpaged. Tompa, A, 1976a Fossil eggs of the land snail genus Vallonia (Pulmonata: Valloniidae). The Nautilus 90(l):5-7. Tompa, A. 1976b. 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Box 7279, Silver Spring, MD 20907-7279, USA. Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028- 1344) is published quarterly by Trophon Corporation, 8911 Alton Parkway, Silver Spring, MD 20910. Second Class postage paid at Silver Spring, MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS P.O. Box 7279 Silver Spring, MD 20907-7279 T H Ef7N AU T I L U S CONTENTS Volume 107, Number 3 February 2, 1994 ISSN 0028-1344 - ' -1 Jii uorar/ Two New Tropical Western Atlantic Species of Epitonium. with Notes on Similar Global Species and Natural History 81 Wooft , as shells in the Virgin Islands. Key words: Epitoniidae. new species. Western .Atlantic INTRODUCTION The family Epitoniidae contains a large number of de- scribed fossil and living species, and as with most groups of mollusks there is as yet no world monograph. In the interim, it could be argued that there should be a mor- atorium on describing any new epitoniids. The literature is very scattered and unsynthesized. There is not even a published list of names. Nevertheless, it is here believed that minimum requirements are met for describing two new species, albeit v\ ithout anatomy. This paper is a by-product of work on a book on the marine prosobranchs of the Bahamas being prepared by myself. Jack N. Worsfold, and Colin Redfern. New spe- cies are not being described therein. The conchological systematics of Recent Epitoniidae (wentletraps) in the western Atlantic and eastern Pacific (where cognate forms, subspecies or species can be ex- pected) are relatively well know n thanks to Clench and Turner (1950b, 1951, 1952, 1953) and DuShane (1974, 1979). Little systematic work has since been done on the western Atlantic species. Even so, it was surprising to conclude that two large, many-ribbed, shallow-water Ep- itonium species had remained undescribed until now. One ranges from southeastern Florida and the Bahamas to the Lesser Antilles (Grenada). The other ranges from the Bahamas to Cuba, Puerto Rico and the Virgin Islands. Both may well have more extended ranges. As recently as 1967, one of the species was first collected alive; it was in less than 1 m of water within sight of a major marine laboratory! Clench and Turner had neither of these species avail- able in the collections they studied [although a small, badly broken shell of one of them was mi.xed in a vial with one £. albidum (Orbigny, 1842) (MCZ 107820). Some epitoniid species seem possibly or probabK to be circumglobal (Robertson and Habe, 1965; Kilburn, 1972, 1985; DuShane, 1983; Paschall, 1987). Therefore, an attempt has been made to consider the two new spe- cies in a global context. Based on epitoniid collections at MCZ, AMNH, ANSP and USNM and literature, shells of 23 species from around the world (.\ppendi.x) were assessed to be most similar to the shells described here. I have tried to survey all the descriptions and illustra- tions of Miocene to Recent species of Epitonium, sensu lata, known from the Americas, and of most Recent species from here and the rest of the world, especially Europe and the Mediterranean. These are the fossils and living animals most likely to be similar or identical to the shells discussed here. European fossil epitoniids were not surveyed because the literature is so extensive. Among the most important references consulted on Epitonium, sensu lata are: 1) world Recent species: Kiener (1838-1839); Sowerbv (1844); Nyst (1871); Sowerby in Reeve (1873-1874); Tryon (1887); Clessin (1896-1897); de Boury (1912-1913); Kaicher (1980, 1981, 1983, the only really global con- tribution in this century, albeit incomplete). 2) American Recent species: Morch (1875a, 1875b, 1876); Dall (1889); Clench and Turner (1950a-1953); Nowell- Page 82 THE NAUTILUS, Vol. 107, No. 3 Figures 1-4. Epitoniuiu phymaiithi liobt-rtsoii, new species. 1. .Apertural, 2. apical, and 3. basal \ie«sof liolot\pe. Shell 16.9 mm long, and 9.3 mm wide. Miami, Florida, ANSP 391939. 4. Unusually low-spired shell (not a paratype), 12,0 mm long, and 8,f) mm wide. Usticke (1959); Keen (1971); Abbott (1974); DuShane (1974, 1979, and references therein); Gundaker (1975); de Jong and Coomans (1988); Espinosa and Fernandez Garces (1990). 3) American late Cenozoic fossil species: Olsson (1916); Maury (1910, 1917); Gardner and Aldrich (1919); Pilsbry (1922); Woodring (1928, 1959); Mansfield (1930, 1935); Pilsbry and Olsson (1941); Gardner (1947, 1948); Olsson and Harbison (1953); Weisbord (1962); Jung (1969). 4) European Recent species: Parenzan (1970); Franchini (1975-1976); Nordsieck (1982); Bouchet and War6n (1986); Poppe and Goto (1971). 5) Indo-Pacific, Japanese, Australasian and South African Recent species: Adams ( 1 861); McK ill and Standen (1903); Jousseaunie (1911); Ircdale (1936); Kerslake (1958); Azu- ma (1962); Macpliorson and Gabriel ( 1962); Kuroda, Habe and Oyama (1971); Masahito and Habe (1973-1976); Powell (1979); Kilburn (1985); DuShane (1988, 1990); Nakayama (1991). An attempt was made to count the ribs on each whorl of each shell The need ior doing so was explained in a previous paper (Robertson, 1983a). Counts on juveniles can be different from those on adults. Indeed, a rib count per whorl in species of Epitoniiim may increase, sta\' the same, decrease, or decrease and then increase as the shell grows in a species-specific maimer. These ta.\onomic characters have hardly been used before. The major study by Clench and Turner (1950b-1953) was published before epitoniids were found to live with or to forage for and feetl on benthic coelenterates, either as parasites or predators (Thorson, 1957; Robert.son, 1963, 1981, 1983b; DuShane, 1988; Yamashiro, 1990; Nakay- ama, 1991, etc.). Both species treated here have actinian (sea anemone) hosts. MATERIALS AND METHODS The specimens studied were assembled during almost two and a halt decades The two shallow -water species are curiously uncommon, and I personally saw neither of them alive. Only empty shells are available of E. pliyiiumthi. but there are a few £. ivorsfoUli in alcohol (see .ANSP "A" numbers in locality records). Shells were studied with the aid of a Wild dissecting R. Robertson, 1993 Page 83 Figures 5-6. Epitunium phymanllii. liicdnipletely grown parat\ pe shells. 5. Only shell known from Bahamas, juvenile. 5 6 mm long, 3.1 mm wide. 6. Subaclull topotvpe. 10 8 mm long, 6.3 mm wide. microscope and camera lucida. First- whorl diameters were measured and whorls counted as advocated by Rob- ertson (1985). The camera kicida and a protractor were used to measure spire angles to the nearest 5°. Specimen numbers in the Specimens E.xamined sections relate to Tables 1 and 2. Rib counts were made on decollated shells b\' mea- suring the mean width of each whorl on intact shells and then extrapolating. .As landmarks, some ribs were lightK marked with a pencil. Repositories of examined specimens are indicated by the following acronyms: AMNH — American Museum of Natural History, Nev\' York Cit> AN'SP — The Academy of Natural Sciences of Philadel- phia MCZ — Museum of Comparative Zoology, Harvard Uni- versity USNM — National Museum of Natural History, Smith- sonian Institution, Washington, D.C. SYSTEMATICS AND NATURAL HISTORY Superfamily Epitonioidea Family Epitoniidae Genus Epitoniiiin Roding, 1798 Figures 7-8. Epitunium phyinantlii. 7. Shell ape.\, .showing a slightly decollated protoconch apex, the almost smooth pro- toconch, the protoconch varix (shown as a line), some of the early teleoconch axial ribs, and fine intervening spiral threads. Topot\pe 8. Operculum (exterior). Topotype. Epitoniurn phymanthi Robertson, new species (figures 1-11, 13) Jong & Coomans, 1988:54, fig. •'Epitoniurn spec: 276,CAira5ao. Shell morphology: Protoconch 0 42-0.50 mm long (ex- cluding immersed base), 0 39-0.42 mm wide, with 3.2- 3 4 whorls; first whorl diameter 0 09-0.13 mm; whorls slightK inflated, appearing smooth, shin), microsculp- ture probably present; terminated with varix; cream- white, tinged with amber at suture, terminal varix, and columella (seen in transparenc\); some protoconchs slightly tilted on teleoconchs. Entire shell reaching 16.9 2? 24 I 22 I e 1 I I I 2 3 4 5 WHORL NUMBERS Figure 9. Epitoniurn plojmanthi. Rib frequencies per whorl on each w horl. The horizontal lines show means, the vertical stippled boxes show ± one stanilard deviation, and the vertical lines show ranges; n = sample sizes. Page 84 THE NAUTILUS, Vol. 107, No. 3 Table 1. Epitonium phymanthi new species. Mensural and meristic shell characters. " + " indicates that a shell would lia\e been larger had it not been slightly broken, or "+ + ' badly broken. Specimen 21 is the holotype All remaining specimens are paratypes except specimen 28, which was not included in figure 9 because of the high rib counts. Speci- men Length (mm) Width (mm) Spire angle Teleoconch whorls A,\ial ribs per whorl no. 1 2 3 4 5 6 7 1 2.2+ 1.5 + 60° 2.5 + 25 29 2 3.1 + 2.0+ 55° 3.4 + 23 24 — — — — 3 3.0+ 1.6+ 45° 3.2 + 21 23 22 — — — 4 2.3 1.5 + 55° 2.8 27 26 — 5 3,5 2.2 50° 3.8 23 24 28 — — 6 4.2 + 2.7 + 50° 4.3 + 25 24 23 25 — — — 7 4.8 3.1 55° 4.4 28 29 29 29 — — — 8 5.5+ 3.7 + 60° 4.7 27 30 29 33 — — — 9 5.3 + 3.3 55° 4.8 25 28 30 35 — — — 10 7.8 4.7 50° 5.6 27 29 28 36 45 — — 11 7.8 4.6 50° 5.7 24 25 27 28 39 — — 12 6.7 4.0+ 50° 5.3 24 30 27 31 38 — — 13 5.0+ 2.3+ 55° 3.7 + 24 27 26 — — — — 14 8.7 + 4.4+ 55° 5.3+ 24 — 28 37 44 — — 15 12.2+ 3.9+ 50° 5.2+ — 26 25 29 — — — 16 8.6+ 5.2+ 50° 5.8+ — 23 26 36 42 — — 17 9.5 + 5.2+ 50° 6.3+ 24 24 21 26 32 42 — 18 11.2 + 6.7 + 50° 6.4 + 27 28 29 30 35 45 — 19 10.3 + 5.8+ 45° 6.2 22 24 24 26 31 39 — 20 11.4 8.1 60° 6.2 23 21 23 26 33 34 — 21 16.9 9.3 55° 7.2 21 22 21 24 33 35 38 22 16.6 9.6 65° 7.7 23 24 22 23 26 34 40 23 15.3+ 8.8 55° 6.2 — 21 25 26 40 36 — 24 5.6 3.1 45° 5.1 22 25 24 24 27 — — 25 9.1 + 5.6 50° 5.7 + — 21 25 28 — — — 26 16.9 9.5 50° 7.8 18 18 27 30 39 37 37 27 6.0+ + 4.5 — 2.6+ + — — — 34 — — — 28 8.6+ + 4.6+ + — 2.8+ + — — 39 45 — — — 29 5.9+ + 4.1+ + — 1.6+ + — — — — — — — mm (Table 1). Teleoconch to 9.6 mm wide (including ribs), with 7.8 whorls, initially thick, becoming relatively thin, fragile w ith increasing shell size. Spire height vari- able, usually low, spire angle 4.5°-65° (mean 53°; Table 1). Spire profile at first straight, convex on later whorls of large shells. Whorls moderately to strongK inflated. Axial ribs 18-45 per whorl, number increasing with shell size (figure 9; table 1 ); ribs thick, low, with edges rounded on upper whorls, thinner, higher, wav\' or crested at shoulder on later whorls. Apical whorls fully attached to preceding whorls, lower whorls slightly detached. Axial ribs aligned, attached erectK from one w horl to the next on upper whorls, rib alignment and attachment decreas- ing with increasing shell size; ribs on later whorls rarely in closely spaced pairs or bifurcating; ribs commonly curled away from aperture. P'ine, regularK spaced spiral threads between ribs on early whorls (about 12 above suture), later whorls with irregular threads, smooth or with fine incised spiral lines. Basal spiral cord lacking. L'mbilicus narrow, or reduced to chink. Aperture oblifjuely to roundK o\ al Teleoconch color cream white or pure white. Operculum not available. (Comparative remarks: This new species differs from E. worsfoldi Robertson (see below) in that the latter species has a thicker more highly spired shell, with coarser, un- crested, less numerous ribs (especially on the last whorl of large shells). Epitonium pht/nidiUhi most closeK re- sembles species assignetl to de Bour\ 's (1912) "subgenus" Foliaceiscala (Appendix). The most closeK- similar spe- cies seem to be E. irregttlarc (Sowerby, 1844) from the Philippines and Japan, and E. lyirgo (Masahito and Habe, 1976) from Japan (Appendix). Etymology: Of Pliijriuiulhui, the genus of sea anemone host. Geographic range: SE Florida, the Bahamas, Virgin Is- lands and Grenada (Lesser Antilles). Not known from I^ermuda or the (Greater Antilles. IVIaterial examined: (Table 1): SE Florida: Bear Cut, Miami (25°43'N, S0°09' VV), 1 m, N, and E, Leeman leg,, 196.5, Holotype— ANSP. 391939, A NSP ,391940 & ANSP 391941, specimens 19-23, Leeman collection 1 specimen, Bahamas: Dead Man's Reef beach, western Grand Ba- hama (26°34'45"N, 78°51'45"W), S, Bowers leg,, Feb, 1981, via J,N, Worslold, ANSP 374362, specimen 24, Virgin Islands: Lindbergh Bav, St, Thomas (18°20'N, 64°58'W), 2 m, MR, Hvett leg,, Oct, 30, 1968, ANSP, R. Robertson, 1993 Page 85 Figures 10-1 1 Topotypes. Epitoniuiu phymanlhi li\ing in aqi: specimen 26. Hams Bay, St. Croi.x (17°47'N, 64°53'W), G. Nowell-Usticke leg., Feb.-March 1957, AMNH 194365 (Specimens 1-18, 27-29)[co-occurred with 1 E. wors- joldi]. Lesser Antilles: Saint George s Lagoon, Grenada (12°03'N, 6r45'W), G. Nowell-Usticke Colin., AMNH 194429, Specimen 25. Specimen 21 is the holotype. All remaining specimens except unnumbered specimen in the Leeman collection and specimen 28 are paratypes. A total of 29 specimens of £. phijinanthi was available for this study. As with E. worsfoldi, many of the shells are broken. Natural history: In 1965, Neal and Eleanor Leeman col- lected six living animals of £. phijmanthi slightly below low tide line at a rocky area in Bear Cut, between Vir- ginia Key and Key Biscayne, Miami, Florida, U.S.A. All were associated with the actiniarian sea anemone Phij- manthus crucifer (Lesueur, 1817). The anemones were clinging to rock substrata, commonly in crevices, and turtle grass (Thalassia festudiniini Banks and Solander ex Konig) rhizomes, most of which were buried in sand. The epitoniums were buried next to the Phymanthtis Figure 12. t'liyiuanthus crucijer, the sea anemone liost ot Epitonium pliijmantlii at Miami, Florida. Size not recorded, but soniewlial less than life size. The anemone colors and pat- terns vary considerably. Photo Neal Leeman. columns under their radially extended oral discs Only anemones buried in sand had E. phijmanthi with them. A few clusters of sand-agglutinated Epitonium egg cap- sules were observed but not studied. On .'\pril 17, 1965, a pair of E. phijmanthi was found with one Pliijmanthus. On May 12, Ma> 30, June 6, and August 8 of the same year, single £. phijmanthi were found w ith Phijmanthns. Unlike a predatory Epitonium, w Inch swallows its coelenterate prey whole, £. phijman- thi is a parasite, feeding on its large host poK p without killing it. When fully extended, a large specimen of Phij- 1 1 Figure 13. Kpilunium phijntantln feeding on the upper col- umn of its sliglitj) contractdl sea anemone host Plujmanlhus crucifer. Note the cvtraordinariK long e\erted acremijojic pro- boscis Miami, Florida Fpitoniums, unlike p\ ramidellids, lake tissues for food rather than fluids. Drawing based on photo- graphs by Neal Leeman. Page 86 THE NAUTILUS, Vol. 107, No. 3 13 I Figures 14-16. EpHonium worsfoldi Robertson, new species. 14. Apertural, 15. basal, and 16. apical (at lower magni- fication) views of the holotype, 18.7 mm long, 9.1 mm wide. Smith's Point, Grand Bahama Island. ANSP A17192. inanthus crucifer is about 13 cm high and about 13 cm wide, much larger than the wentletrap. Epitoniums can vary their host preferences from place to place. During the same year and months, at the same locality, and associated with the same species of sea anemone, the Leemans collected six living Epitonium lamellosum (Lamarck, 1822)(ANSP 391948, A16850) and one living Opalia crenata (Linnaeus, 1758)(ANSP A16851). Thus Phijnianthus crucifer supports three par- asitic epitoniid species at one locality. £, phijmanthi may or may not be specific to Phymanthus elsewhere. Colors and color patterns of P. crucifer vary consid- erably. Usually, these anemones are variegated with shades of green or brown. The white shell and body of E. phijmanthi, and the white and reddish brown shell and nearly white body of £. lamellosum, make neither species cryptically colored with its host. At least during the day, both species are buried in sand ne.\t to the anemone. The Leemans maintained £. phymanthi and E. la- mellosum with Phymanthus crucifer in home aquaria from mid-April to mid-October, 1965 (two E. lamellos- um for as long as three months). Their observations are abstracted below. The external body coloration of E. phymanthi was white, that of E. lamellosum was white except for light \ellowish tentacles that taded to white in aquaria. The fully everted proboscis of E. phymanthi was at least four times the shell length, while the fully extended proboscis of E. lamellosum was only slightly longer than the shell. Roth species behaved as if they locate their host by che- E . WORSFOLDI I _ T T T (III 3 4 6 6 WHORL NUMBERS Figure 17. Epitonium worsfoldi, new species. Rib frequencies per whorl on each whorl. The horizontal lines show means, the vertical stippled boxes show ± one standard deviation and the vertical lines show ranges; n = sample sizes. motropism, using their tentacles and slightK' everted pro- boscises as do two Californian species (Smith, 1977; Saio, 1977). Both E. phymanthi and E. lamellosum were seen to feed on Phymanthus columns near the base and under the oral disc, ingesting column tissues (with fluid? mu- cus?). Anemone tentacles were not observed to be at- tacked by either species. Duration of feeding in £. phy- manthi was 2-10 minutes (animals not starved; mean of 5 observations ca. 4 min). Epitonium lamellosum (a larger species, up to 45 mm shell length) starved for 3- 5 days fed for 45-97 minutes (mean of 4 observations: 64 min.). No purple secretion was seen to be released from the pigmented mantle organ b\ either species dur- ing feeding. One Phymanthus sur\ ived repeated attacks by the two species for upwards of one month. The anem- one w rithed near where it was attacked, and the column also swelled. The anemone reacted least to E. phymanthi, the smaller, slower species. An £. phymanthi with a shell 7 mm long on August 11 grew to 1 1 mm by September 22, a mean rate of 0. 1 1 mm /day. Growth initially was faster (between August 11 and 21: 0.2 mm/day), but there was no growth be- tween September 22 and October 5. Another £. phy- manthi fed and survived from August 1 1 to September 2 without growing (22 days). Growth of £. lavwllosum was rather faster and also erratic, but growth appeared to be indeterminate (Leeman and Robertson unpub- lished). Epitonium phymanthi was observed to move very slowly and remain motionless for long periods of time. Epitonium lamellosum was more active. On several oc- casions, £. phymanthi was observed following £. la- mellosum mucous trails, and partially everting and in- verting its proboscis, each time ending by touching an R. Robertson, 1993 Page 87 Figure 18. Oral vie« of either Actinoponis elegans or Hom- ostichanthus duerdeni. a sand-dwelling sea anemone, showing beside it the holot\ pe and egg capsules of Epiioniiim worsfoldi found with it A paratype shell also shows. The anemone is 7,5 cm in greatest diameter. Smith s Point, Grand liahama, I5a- hamas. Photo Jack N. Worsfold. .¥'-:C Figure 19. Li\ ing Epitonitim uursfoldi near some of its sand- agglutinated egg capsules Some of the shell ribs are abnormally bent. .-WSP Ai7I94. Shell IS 8 mm long, 9.8 mm wide. Grand I^ahama, Photo Jack N. Worsfold. £. lamcllosum. On one occasion, an E. laincllosiun ap- proached an £. phijmanthi in the same \va) . Once, in- dividuals of the two species remained aperture to ap- erture for about one hour, with the four tentacles curled around each other. Groups of egg capsules with £. phijmanthi were found in early May, early June, and early October. After 36 days in isolation, one £. phijmanthi had two groups of fresh egg capsules attached to it, indicating a capacit> for sperm storage. A small crab killed an £. phijmanthi with a shell 10 mm long. Three other, larger animals died from un- known causes. Epitonitim worsfoldi Robertson, new species (figures 14-19) ?Turbo principalis Pallas. 1774: 33, pi 3, figs. 5-6. No localit\ Nonwn (hdritim. This species might not be large if the figures are enlarged. The spire angle (if accurately drawn lis 30°-35° ?"Sca/a principa/is (Pallas)": Morch, 1875a. 1875b, 1876.Tortola. British Virgin Islands. ProbabK not of Pallas. Unidentified. Huber, 1982: 202, 2 figs. S of Balmoral Island, Nassau, Bahamas. Reported to be I 5/8" long (=41 mm). "Epitonium principaUs (Pallas, 1774)": Paschall, 1986:66, figs, la and b. Next to a sea anemone, tureen Turtle 'Ke\" [Ca> ], .-\f)aco, Bahamas. Not of Pallas Reported to be 1.75 inches long (= 44 mm). :*£. principale "Roding, 1798": de Jong & Coomans, 1988:54, fig. 275. "Rather common in Curai^ao and Arutia. Too few axial ribs. ?"£. fricici (Dall): Espinosa anil Fernandez CJarces (1990:6), probabl) not of Dall; three small shells from Bah!a Cien- fuegos, C'uba, the longest 3.8 mm. Shell morphology: Protoconch 0.52-0.54 mm long (ex- eluding immersed base), 0.40-0.43 mm wide, with 4.1- 4.2 whorls; first whorl diameter 0.12 mm; whorls slightly inflated; microsculpture as in £. alhidiim (Robertson, 19S3b); \\ hite, with pale brown at suture, terminal varix. Entire shell attaining 25.3 mm (Table 2). Teleoconch to 11.5 mm wide (including ribs), with ca. 8.8 thin, fragile whorls. Spire high, spire angle 35°-50° (mean 44°); spire profile convex, rareK straight or initialK concave; whorls moderately inflated. Axial ribs thick, low on upper w horls, thinner, higher, upright or recurved without subsutural or shoulder cresting (except remnants from breakage) on later whorls; ninnbering 16-21 on all but first and last whorls large shells (23 ribs on last whorl of 8.8+ whorl shell), with ribs/ whorl initially decreasing then increas- ing with shell size (figure 17); axial ribs on successive whorls usually attached, attachments erect, rarely offset away from aperture on lower w horls; rarely , all ril« bent medialK after an injury (figure 19). Suture present in Page 88 THE NAUTILUS, Vol. 107, No. 3 Table 2. Epitonium worsfoldi new species. Mensural and meristic shell characters. " + " indicates that a shell would have been larger had it not been slightK broken, or "+ + bailK broken Specimen '57 is the liojotNpe Speci- men Length Width S[)ire Teleoconth Axial ribs per whorl no. (mm) (mm) angle whorls 1 2 3 4 5 6 7 8 1 17.94- 8.9-1- 45° 7.34- — — 21 18 18 18 18 2 18.8 9.8 45° 8.0 20 20 19 18 18 19 19 20 3 9.4 5.3 40° 6.5 20 20 20 19 19 20 — — 4 8.8 5.4 45° 6.6 22 21 20 20 21 21 — — 5 — 4.7 — — 19 19 19 18 19 20 — — 6 — — — 3.3 23 21 21 20 — — — — 7 — — — — 20 bS 17 18 17 18 — — 8 — 11.3-f — ca. 8,8 — — — — — — — 20 9 22.1 10.3 40° 8,3 20 20 19 19 18 18 18 18 10 15.6-1- 7.7 .35° 7.2 — 19 19 20 19 17 17 11 12.5 5.9-1- 40° 6.84- 19 19 19 18 19 20 — 12 17.2 8.0 40° 7.64- — — 21 20 19 19 19 13 14.3 7.2 40° 7.1 — — 20 19 19 19 19 14 19.7-H 9.9 40° 8.2 — — 19 19 19 19 19 20 15 19.6-t- 9.7 40° 8.0 — 21 20 20 20 20 21 19 16 19.7 9.6 40° 7.8 — 20 18 17 18 18 18 — 17 14.9 8.1 40° 7.3 — 20 19 18 18 18 19 — 18 18.3 9.5 45° 7,9 — 17 18 18 17 18 18 — 19 19.2 9.2 40° 8,1-H — — 21 19 19 19 19 23 20 4.4 2.4 4(J° — — — — — — — — — 21 6.9 3.8 40° 5.4 — 20 18 19 19 — — — 22 25.3 11.5 45° 8.1 — 19 19 19 19 20 20 21 23 15.6 8.2 4.5° 8,0 — — — — — — — — 24 21.5-1- 9.0-H 40° 7,9 — — 19 17 17 17 17 — 25 18.9-1- 9.2-H 40° 7.8 — — — 19 19 19 — — 26 17.7-1- 9.3 -H 45° 7.8 — — 20 21 20 20 21 — 27 13.9-1- 6.4 4- 40° 6.94- — — 18 18 18 — — — 28 19.4 9.6 4.5° 7.7 — — 20 20 20 20 20 — 29 15.0-F 7.54- 45° 7.1 — 20 18 18 17 17 17 — 30 18.6-h 10.0 40° 7.2 — — — 18 18 18 19 19 31 20.2 10.7 45° 8.2 — 18 18 18 18 18 18 19 32 18.8-1- 10.0 4.5° 8.0 — — 18 18 18 18 18 19 33 ■22A + — 4.5° 8,3 — — 20 20 20 20 19 21 34 17 1 8,4 45° 7.74- — — 18 17 18 18 18 19 35 12.9 7.2 45° 7.1 — 17 17 18 — 18 18 18 36 10.2-1- 4.74- 50° 5.54- — — 17 17 19 18 — — 37 18.7 9.1 4.5° 7.8 — 19 19 20 20 19 20 — 38 e,.o+ + 2 94-4- 4.5° 5.64-4- — — — — — — — — iiitcr-rib areas of uppermost whorls; middle, lower whorls slightly detached. Areas between a.xial ribs with faint, irregidarly-spaced, incised spiral lines; axial growth lines present, Bod\ whorl v\ ithout basal spiral cord limbilical chink narrovN (rarely almost closed). Colunieiiar callus thin or thick, thinnest medially, not conforming to un- derlying ribs. Columellar chinks present. Aperture roundly oval. Teleoconch color pale tan, cream-white, pure white (when bleached?). Operculum ])aucis|)iral, growth wrinkles present, pale amber. Comparative remarks: Epitonium worsjolili most close- ly resembles species in the de Boury "subgenera" Hy- aloscala [1889], Umiscala [\909], aiuj rapyriscala [\m)9] (see Appendbx) as di.scu.ssed in Kilburn ( 1985). The shell of £, worsfoldi most closely resembles /.', kraiissi (Nyst, 1871) of South Africa and E. me/ior (Mel vill andStanden, 1903) of the northern Arabian Sea (see Appendix). Etymology: Named for Jack Nigel Worsfold, indefati- gable naturalist and friend. Geographic range: Bahamas, northern Cuba, and Puerto Rico. Not known troni Bermuda, Florida, or the Lesser Antilles. Material examined: (Table 2): Bahamas: Fleming Road iH'uch, Mosquito Point, 8 km SE of West End. (Irand I5ahama (26°37'30"N, 78°54'00"W), J.N. Worsfold leg., ca. 1976, ANSP 370051, specimen 33- Smith's Point, Grand Bahama (26°31'N, 78°37'W), 1 m, 1 living under Stich()(lu(tyl(i ["Stoichactis"] heliatUhus, 1 large, with egg capsules, li\ ing under either an :\ctinoporus clegans R. Robertson, 1993 Page 89 or Homostichanlhns ducrdeni (Figure 18), J.N. VVors- fold leg., August 1985, ANSP A17192, Holotype. ANSP A17193, specimens 35, 37.— SW of Sharp Rocks Point, near Peterson's Cay, S coast central Grand Bahama (26°33'45"N, 78°33'30"\V), 1 in, 2 juveniles living with egg capsules under Stichodactyla. J.N. Worstold leg., June 1984, ANSP A 17194.— Cold Rock, S. coast central Crand Bahama (26°36'15"N, 78°22'15"W), 1 m, 2 ju- veniles living under Stichodactyla. J.N. Worsfold, leg., June 1984, ANSP A17195. — Treasure Cay and vicinity, Abaco, 12 shells, C. Redfern leg., Redfern collection, specimens 7, 23-32. — W coast N end Elbow (Little Guana) Cay, off NE Abaco (26°33'00"N, 76°56'45"W) K. A. Robertson leg. ca. 1954, ANSP 359100, specimen 16. — Wood Cay. Schooner ("ays, W of S Eleuthera, M McNeilus leg., Dec. 1980, McNeilus collection, specimen 1.— North East Point, Arthurstown, Cat Island (24°38'N, 75°3S'W), 1 broken shell u ith an £. albidum. W.J. Clench and H.D. Russell leg., 1936, MCZ 107820, specimen 36.— Bahamas unlocalized, .ANSP uncatalogued, specimen 34. Cuba: NW coast. Oasis Beach, Via Blanca Highway km 28, Matanzas (23°11'N, 82°04'W), C.J. Finlay leg. Feb. 1959, Finlay collection, specimen 15. — Same, km 27 (23°11'N, 82°05'W), C.J. Finlay, leg., 1951-1956, Finlay collection, specimens 5,9,10,11, ANSP 359101, specimens 13,14. Puerto Rico; Punta Ostiones, 7 km S of Punta Guanajibo, W coast, 1 shell, April 22, 1949, 3 shells. May 22, 1949, G.L. Warmke, leg., ANSP, specimens 3-4, 17- 18. — Ramey Air Force Base, NW coast, A. Phares, leg., ANSP, specimen 19. — Pinones Beach, 8-10 km E of San Juan, NE coast (18°26.8'N, 65°55.7'W), Mrs. D. Hum- phrey leg., 1970, Finlay collection, specimen 8. Virgin Islands: Hams Bay, St. Croix (17°47'N, 64°53'W), with 21 £. plujmanthi shells, Feb. -March, 1957, G Nowell- Usticke leg. AMNH 194365, specimen 38. Specimen 37 is the holotype. All remaining specimens except specimen 34 are paratypes. A total of 38 specimens of £. worsfoldi was available for study. Of these, 6 were collected alive: 2 large spec- imens (the smaller the holotype), 2 juveniles, 1 with the dried body deep in the shell, and 1 with the operculum but no bod\ . The last 2 and most of the remaining shells were collected from beach drift. Many of the beach shells are in poor condition. The fairly fragile shell, predators (crabs? fish?), and wave action presumabK are responsible. Out of the 38 shells, only 1 has an intact protoconch. Natural history: At Grand Bahama, this species was found alive three times with Stichodactyla hclianthus (Ellis and Solander, 1786), the preferred host also of Epitonium albidum (Orbigny, 1842)(Robertson, 1983b) and an oc- casional host of Epitonium lamellosum. Like £. albi- dum, E. worsfoldi was under the broad oral disc, next to the column, in sand. The holotype of E. worsfoldi was collected with an anemone that was either Actinoporus elegans Duchassaing, 1850 or Homostichanthus ducr- deni Carlgren, 1900, a sand-dweller (Figure 18). Feeding (parasitism) on the anemones was not observed but must occur. The sand-agglutinated egg capsules are like those of £. albidum (Figure 19). The bathymetric range is 0-2 m. CONCLUSIONS The data in this paper are believed to warrant description of the two new Recent western Atlantic species. Their fossil ancestors seem not to be known in the Americas. As documented in the Appendix, the most similar Recent shells appear to occur in Ja|3an and South .Africa. It will be noted in the .Appendix that the 23 species have been assigned to a variety of genera and "subgen- era" (many of the latter named by de Boury). Epitonium worsfoldi has characteristics of three of these "subgen- era combined. Epitoniid genera and subgenera need to be much more broadly based. It is surprising that two such large, shallow water spe- cies in a well-collected area should have remained un- de.scribed until now. The\ show the continuing role that amateurs play in collecting, observing, photographing, and providing material for systematic and biological studies in museums and marine stations. Amateurs not onl\ hnd undescribed species on rare occasions but, per- haps more u.sefull\, they can also make aquarium ob- servations such as those of the Leemans reported here. ACKNOWLEDGMENTS Most of the specimens were collected b\ amateurs, and AMNH in the 1970's became the first museum to acces- sion good specimens of one of the two species (from the Nowell-Usticke Collection). Other amateurs who con- tributed most to the discoveries by providing specimens, data, and photographs are Neal and Eleanor Leeman (formerly of Miami, Florida), C. John Finlay (formerly of Cuba), and Jack N. Worsfold (formerly of the Ba- hamas). Worsfold's Grand Bahama collection is now at ANSP. Dr. Frederick M. Baver identified the Phyman- thus, and Dr. Donald R. Moore helped the Leemans in various other ways. Dr. Norman D. Paschall and Helen DuShane were helpful in providing information. Dr. Daphne Gail Fautin identified the Actinoporus or Hom- ostichanthus from figure 18 (no specimen was kept). The curators of the mollusk collections at MCZ, AMNH, and USNM helped by providing access to their collections and making loans. Other persons who helped with the loan or gift of specimens are Dr. Marvin R. Hyett, Mrs. Garwin (Marilee) McNeilus, Amy Phares, Colin Redfern, and Germaine L. Warmke. Mary Fuges inked figures 7, 8, and 13. Harriet Robertson calculated the mensural data and inked figures 9 and 17. Colin Redfern, Dr Gary Rosenberg, Jack N. Worsfold, and two helpful reviewers read various drafts of the manuscript. LITERATURE CITED Abbott, R.T. 1974. American seashells, 2nd ed \'an Nnstrand Reinliold, New York, etc., 663 pp., 24 pLs .■\danis, A. 1861. On the Scalidae or "wenlletraps of the Sea Page 90 THE NAUTILUS, Vol. 107, No. 3 ot Japan; with descriptions of some new species .■\nnals and Magazine of Natural Hi.story (3)8:479-484. .\zunia. M 1962 Descriptions of five new species of Japanese Kpitonlidae, N'eniis: The Japanese journal of Malacology 22(2): 1. 50- 1. -35, (i figs. Boucfiet, P. and A. Waren. 1986. 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Berlin, fasc. 10:32-34, pi, 3, Parenzan, P. 1970 Carta d'identita delle eonchiglie del Med- iterraneo. Vol. I. Gasteropodi. BiosTaras, Taranto, 283pp , 53 pis. Paschall, N.D. 1986. Lost and found; Atlantic "Precious Wen- tletrap. Conchologists ol .America Bulletin 14(4) 6(S, figs la and b Paschall, N'D, 1987, Research results: one species worldwide :■ Epitonium hlainci/zelebnri. Conchologists of America Bulletin 15(1): 13-14, 2 figs. Pilsbry, HA. 1922. Revision of W.M. Gabb's Tertiary M, GB, II. 1873-1874. Monograph of the genus Sca- luria In: Reeve, L.A., Conchologia Iconica. . . . London 19. 16 pis., 125 species [no pagination]. Souerby, GB. III. 1894. Descriptions of twelve new species, chiefly from Mauritius. Proceedings of the Malacological Societ) of London 1(2)41-44, pi 4 Strong, ,AM. and L.G. Ilertlein. 1937. New species of Recent mollusks from the coast of western North America. The Templeton Oocker Expedition of the C^alitornia .Academy of Sciences, 1932. No. 35. Proceedings of the C^alifornia Academy of Sciences (4)22(6): 159-178, pis. 34-35 Taki, I 1956 Anatomical study on Japanese P^pitoniidae (1 ). Epitonium, Amaea and Papyriscala Bulletin of the Na- tional Science Museum (Tokyo) 3(2)(39):71-79, pis. 13- 17 Thorson, (; 1957. Parasitism in the marine gaslropod-familv Scalitlae. Vitlenskabelige Meddelelser fra Dansk naturhis- torisk Forening i Kcibenhavn 119:55-58 TrNon, G.W. 1887. Family Scalariidae. In: Manual ol Con- chology. . .. [ser. 1] 9:49-111, pis. 11-18. Warmke, G L. and R.T .Abbott 1961. Caribbean sea- shells. . , , Livingston, WyiHiewood, Pennsylvania, x + 346 pp., 44 pis Weisbord, N.E. 1962. Late (;enozoic gastropods from north- ern Venezuela. Bulletins of American Paleontologv 42(193): 1-672, 48 pis. Wilson, B.R. and K. Gillett, 1972, Au.strahan shells, Tuttle, Rutland. X'ermonl, and Tok\o, 168 pp,, Ut6 pis Woodring, \V,P, 1928, Miocene mollusks from Bouden, Ja- maica Part II Ciastropods and discussion of results. Car- Page 92 THE NAUTILUS, Vol. 107, No. 3 iiegie Iiistitiiliiiii cil W'.isliint^lnii rublicalion .38,5:i-vii, 1- 56-4, JOpls. VVoodring, W.P. 1959. Geology and paleontology of Clanal Zone and adjoining parts of I'anania Description of Ter- tiary molliisks (gastropods: \ errnflidac to Tliaididae). [United StatesjCleological Siir\fv Prolessioiial Paper '>06- B:iii, 147-2.39, pis. 24-;5S Yamashiro, H. 1990. A wcntlelrap i'.pitoniiini hullatuiu as- sociated with a coral Sundali/lithu robusta. \ enus: The Japanese Journal of Malacology 49(4):299-305. APPENDIX. Conchological comparisons or notes on 23 species similar to, or confused with Epitonhim phijmanthi and E. wors- foldi, regardless of provenance. Scala amathusia Melvill & Standen, 1903:341, pi. 7, fig. 1. Kaicher, 1980: card 2300 [Epitonium]. Gulf of Oman. Smaller than £. worafuldi (ft. 5 mm), with more whorls (8-9) for size. Scalaria costulata Kiener, 1838-1839:5-6, pi. 2, figs. 4. Tryon, 1887:pl. 12, fig. 59. Robertson, 1963:57-58, pi. 5, fig. 4. Robertson, 1970:45 [identification confirmed]. DuShane, 1988:30-32, fig. 1 (ribs curiously opisthocline). ANSP 230639. Red Sea to Philippines. Thought to be a synonym of S principalis (Pallas) by Sowerby (1844:88 bis), which he recorded from "Xipixappi, West Colum- bia." Larger (35.3 mm); spire high (spire angle 30°); suture slightly disjunct; up to 28 ribs on last whorl; um- bilicus narrow, deep. Scalaria crassa Sowerby, 1844:91 bis, pi. 33, fig. 55. Kaicher, 1981: cards 3066, 3081 [Epitonium]. Philip- pines. Unlike E. ivorsfoldi, ribs unequal, slightly fewer (18) at largest size; may be spirally brown-banded. Epitonium (Limiscala) cnjpticocorona Kilburn, 1985: 309-310, figs. 79, 126. South Africa. Smaller (7 mm); weak "coronation" inside suture; 12-19 ribs; low spiral lirae. Scalaria dubia Sowerby, 1844:90 bis, pi. 33, fig. 41. Not Epitonium duhium Roding, 1798:92. Sowerby in Reeve, 1873: species 45, 45b wrongly as 75b. Tryon, 1887: 60, pi. 12, figs. 67-68. De Boury, 1912:95-97,' pi. 7, fig. 4. Kilburn, 1985:327. Type species of Foliaceiscala de Boury, 1912. ANSP 19573 (broken); USNM G.177. Aus- tralia, Samoa, etc. Larger (37 mm); spire angle 45°; like £. phymanthi but lacks crests; ribs very low, thin, slightly recurved, rarely enlarged, up to 42 on last whorl; suture impressed but not disjunct; spiral threads crowded, vari- able; last whorl inflated; umbilicus very narrow. Scala emiliac Melvill & Standen, 1903:343-344, pi. 7, fig. 6. Kaicher, 1980: card 2313 [Epitonium]. Kilburn, 1985:307, figs. 120-121. Pakistan to South Africa. Smaller than E. phymanthi (12 mm); spire slightly higher (spire angle 40°); 21-36 low, thread-like ribs; pale brown (if not laded). Scalaria Jriahilis Sowerby, 1844:95 bis, pi. 33, fig. 74. Kaicher, 1980: card 2^29 [Epitonium]. Southern Austra- lia. Higher spire than E phynianllii (spire angle 30°- 35°); spire profile evenly convex; no umbilicus. Scala frielei Dall, 1889:313. Not illustrated. USNM 83727 (syntypes): AMNH 194388. North Carolina; Vir- gin Islands. Clench & Turner (1952:300-301) and Warmke & Abbott (1962:81, pi. 14a) misidentified this species. Smaller (6.4 mm); 24-31 ribs on last whorl; no shouldering; strong spiral threads; umbilicus narrow ; spire fairly high (spire angle 45°). Epitonium phymanthi has been misidentified as this species. Epitonium (Nitidiscala) hancocki DuShane, 1970:332, pi. 51, fig. 1 DuShane, 197432-33, fig. 73. Kaicher, 1983: card 3612. Galapagos. Smaller (13 mm); spire higher (spire angle 30°); fewer ribs (21); rib crests wavy. Scalaria imperialis Sowerby, 1844:91-92 bis, pi. 33, figs. 56-.57. Wilson & Gillett, 1972, pi. 13, figs. 5, .5a [color]. [Epitonium inipcrialc]. Kaicher, 1980: card 2304. AMNH 1.36625, 1.57310; ANSP 181726, 19,5661, 2.5.3841, 25.3842, uncat.; MCZ 294909, 294914; USNM 694170. East Africa, Western Australia, Philippines and Queens- land. Larger (40 mm); pale tan to dark reddish or pur- plish brow !i, commonly in spiral bands (ribs white); more ribs, increasing to 48 on last whorl; umbilicus wide and deep; operculum grey-black. Scalaria irregularis Sowerby, 1844:90 bis, pi. '33, figs. 40, 60. Philippines; Japan. USNM .343440. Up to 18 mm long; spire angle 6.5°; ribs crowded, 37 on last whorl, irregular in thickness but generally thin, with slight crest- ing; strong spiral threads; slightly umbilicate; 1 pale brown subsutural band present or absent on white. Resembles £. phymanthi. Scalaria kraussi Nyst, 1871:116. Kilburn, 1985:29.3- 295, figs. 97-100 [Epitonium (Hyaloscala)]. + Epiton- ium shepstoncnsc E..^. Smith, 1910:204, pi. 7, fig. 15. Kaicher, 1981: card 3113. .South Africa. Smaller (10-16 mm); spire averages higher (spire angle 25°-40°); 17-37 fine, low, erect or slightly reflexed ribs on later whorls; umbilicus closed. Species believed to be most similar to £. ivorsfoldi Perlucidiscala lacrymula Jousseaume, 1911:198, pi. 5, figs. 37-42. Kaicher, 1981: 3063 [Epitonium "lachry- mida"]. Gulf of Aden. Smaller (5 mm); spiral threads towards apex. Scalaria latifasciata Sowerby in Reeve, 1874: species 117. Type species ot Papyriscala de Boury. Taki, 1956 [anatomy]. ANSP 70738, 219307, 234733, 243255; MCZ 294908 and 5 uncat. lots. Mozambique; Mauritius; Japan. L'p to 18 mm long; spire low (spire angle 5.5°); trace of whorl disjunction; thin, low ribs mostly not connected from whorl to whorl; 32 ribs on last w horl, some irreg- ularly spaced or thickened; weak to strong spiral threads; no crests; 2 or 3 brown spiral bands; slightly imibilicate. Scalaria lineolata "Kiener" Sowerby, 1844:91 bis, pi. 33, figs. 45, 46, 48. Confused with S. lineata Kiener not Sav. Nvst, 1871:118, pi. 2, fig. 5, pi, 6, fig. 20. Kaicher, 1981: card 3125 [Epitonium]. ANSP 70742, 86246, 243289; MC:Z 294911 and 2 uncat. lots; USNM 198709. Gulf of Aqaba; Mauritius; Philippines; Japan. Up to 19 mm long; spire angle 40°; ribs more numerous (16-33) and more irregularly spaced and thickened ribs; no spiral sculpture; umbilicate; 1-3 brown spiral band(s) per w horl; operculum dark brown Scalaria lyra Sowerby, 1844:89 bis, pi. 32, figs. 38-.39, pi. 34, figs. 81-82; Sowerby in Reeve, 1873: species 23. R. Robertson. 1993 Page 93 Kaicher, 1980: card 2341 [Epilimininl KillKirn, 1985: 308-309, figs. 122]. ANSP 195(iH, 19567, 119638; MCZ 187720; LlSNM 431817, 306339; 820891. Type species of Limiscala de Boury. Fiji to Japan and the Red Sea to Mozambique. Slightly smaller (20+ mm); spire lower (spire angle 60°); 27-45 erect ribs; spiral threads fine and dense; 0, 2 or 3 brown spiral bands. Scala melior Melvill & Staiulen, 1903:345, pi. 7, fig. 9. Kaicher, 1980: card 2S33[Epitonium]. ANSP 164808 (syntype); USNM 424868. Pakistan; Gulf of Oman Smaller (6.5 mm + ); 8 (teleoconch?) whorls; spire angle 30°; up to 23 ribs per whorl; spiral threads present or absent. Resembles a small E. wor.sjoldi. Scala micromphala Morch, 1875a:258 ("Vieques," Puerto Rico, one specimen collected by A.H. Riise) Not illustrated. Photograph of holotype in Clench and Turner (1951:258-260, pi. 112, fig. 3), wrongly synonymized by them with E. occidentale Nyst, 1871, which according to them has "12 to 15 costae on the body whorl. ' Ac- cording to Morch there were 18 ribs on the 5 mm. -long shell. An E. phymanthi that long would have had more than 20 ribs. In addition, E. micromphala has subsutur- ally crested ribs; in E. phymanthi the crests are on the shoulders of the ribs. Morch wrongly likened S. microm- phala to S. imperialis (see under that species) Scala {Viciniscala'"') mimiticosia de Bour\', 1912:87- 90, pi. 7, fig. 1. Compared with Scala costulata and S. principalis "West Columbia." DuShane ( 1974:20-22, figs. 20-26). Larger than £. worsfoldi (35 mm); spire high (spire angle 35°); 21 mainly thin, low ribs on last whorl; whorls all narrowly disjunct, but ribs attached; traces of spiral striae; umbilicus narrow, deep; ribs slightly re- flexed; crowded spiral threads. Turbo principalis Pallas, 1774. Nomen dubium. Epi- tonium principale (Pallas) Roding, 1798:91. De Bourv, 1912:89-90, 97. See above in synonymy of E. worsfoldi. Scalaria rohillardi Sowerb\ , 1894:42-43, pi 4, fig. 5. Kaicher, 1981: card 3037. Kilburn, 1985:305-307, figs. 118-119 [Epitonium (Papyriscala)]. Red Sea and India to South Africa. Smaller (15 mm); more ribs (20-24 on last whorl); umbilicate; pale tan with 2-3 brown spiral bands (rarely absent). Epitonium shcpstonense. See above under E. kraussi. "Foracciscala" [=Foliaceiscala] virgo Masahito & Habe, 1976:172, figs. 2-3. Compared with "Foraccis- cala" duhia (Sowerb\, 1844). Japan. Paratype most re- sembles £. phynuinthi, spire angle 40°; 50-60 weakly lamellate growth riblets; suture slightly disjunct, whorls connected by ribs; aperture constricted (holotype); oblique spiral threads. Epitonium (Nitidiscala) willetti Strong & Hertlein, 1937:171, pi. 35, fig. 5. DuShane, 1974:38-39, fig. 72. Tropical eastern Pacific. Much smaller (3.2 mm); 18- 22 -h ribs. THE NAUTILUS 107(3):94-96, 1993 Page 94 Phalium {Semicassis) vector, a New Deep-Water Species from the Central Indian Ocean R. Tucker Abbolt Bailey-Matthews Shell Miimuiii P.O. Box 1580 Saiiibel, FL 33957 USA ABSTRACT Phalium (Semicassis) vector, a new species of the family Cas- sidae, is described from the Saya de Malha Bank, northeast of Mauritius in the Indian Ocean It differs from its closest relative, Phalium microstoma (von Martens, 1903) from Somalia, in having a lighter, less elongate shell with stronger beads on the spiral lirae. Keii irords: Cassidae, Sa\a dc Malha liank. Indian Oceaii INTRODUCTION A number of new species of marine mollusks have been collected over the past few years by Soviet research ves- sels exploring the Saya de Malha submarine bank in the central part of the Indian Ocean. Among these are species of cassids not known at the time of my revision of this family (Abbott, 1968). In 1975, Kilburn described Phalium glahratum sub- species fernandesi from deep water off Somalia. Two years later it was dredged by the R/V Gordy on the Saya de Malha Banks. With it were recovered at least five specimens of a new Phalium referable to the sub- genus Semicafisis Morch, 1852. The bathymetry antl geologs of the Saya de Malha Bank are discussed by Bouchet and Bail (1991:161), while Bondarev and Rockel (1992) provided an excellent pop- ular account of this bank, which is located along the Mascarene Ridge north of the Mauritius and C^argados Carajos Shoals (between 9°:3()'S and 12°2()'S, and 59°30'E and 62°3()'E). interstices of etjual width. CJolor opaque white with large, faint, light-brow II blotch on dorsal surface of last whorl. Protoconch (fig. 2) of 3'/2 whorls, obtuse-conical, glassy- white, smooth. Teleoconch of 5'/2 whorls, the last bearing 23 flattish, spiral cords with fine beads at intersections with fine axial threads. Suture simple, slightK impressed. Spire convex-conic. Aperture % shell length. Outer lip reflected, of variable thickness, enamel-white, crossed by 20-22 distinct, sharp white denticles. Both true and false umbilici small, open, deep. Channel behind siphoual ca- nal deep, narrow. Parietal wall slightly glazed over. An- terior part of columella strongly and unevenly pustulose, inner edge with 4-5 small denticles, outer edge with 3- 4 small, roimded projections. Soft parts and operculum tmknown. Type locality: Central Basin, Saya de Malha Bank, Mas- carene Ridge, northeast of Mauritius, Indian Ocean, dredged from sandy silt bottom at a depth of 80 meters, R/V Gordy, 1989. More precise locality data did not accoinpany the holotype. Material examined: Holotype, USNM 860326, Paratypes 1-3, Donald Dan collection, all from the type locality. The specimen illustrated by Bondarev and Rockel (1992; fig. 5) is designated as paratype 4 Etymology: 1 take pleasure in naming this species after the first recreational diving vessel to operate under the Russian flag, the Vector. Comparative remarks: This new species belongs to a species group of Phalium (Semicassis) mainly limited to SYSTEM ATICS Phalium (Semicassis) vector Abbolt, new species Figures 1-2 Synonymy: Phalium sp 1. Bcindarev & Biicki 1992:28,32, fig 5 Description: Shell (fig. 1 ) reaching 60 mm in length, light, strong, evenly ovate, with numerous, fine, flattish, crowded, weakly-beaded spiral lirae separated b\ sunken Table 1. Phalium (Semicassis) vector new spfc'\es l.inearshell measurements (nun) and counts llol(it\pc I'aratype 1 Paratype 2 l'arat> pc 3 Length 53.6 56.8 53.4 (iO 0 Width 32,0 32. 1 33.4 33 5 Number ol whorls R. T. Abbott, 1993 Page 95 '--^ \. Figures 1, 2. Phalitun {Seniicassis) vector new species. 1. A|Hiluidl, lalcral, and dorsal views of the holotype. 1.0 X. 2. Protoconch of paratype 1. Botli from Saya de Maiha Bank, Mascarene Ridge, N E. of \4adagascar, dredged from sandy silt bottom at a depth of 80 meters, R/V C;ordy, 1989. Scale bar = 500 nm. the western Indian Ocean, although similar species, such as P. sculptuni J. de C. Sowerby, 1840 (Miocene), P. mehranictim (Vredenburg, 1925) (Upper Miocene) and P. oligocalanticum (Vredenburg, 1925) (Oligocene) have been reported from fossil beds on the mainland of India. Phalium vector most closely resembles the eastern Af- rican deep-water Phalium {SemicassU} microstoma (von Martens, 1903), from which it differs in being niucli lighter in weight, less elongate, and in having stronger beads on the spiral lirae. Phalium vector lacks the four or five spiral brown bands on the body whorl, but instead has a faint tan blotch on the dorsal surface of the body whorl. The very weak, rounded projections on the outer, left edge of the columella of P. vector are reminiscent of the fingertip projections found in the shallow-water Indian Ocean Phalium Jaurotis (Jousseaume, 1888). ACKNOWLEDGMENTS I am indebted to Mr. Donald Dan of Ft Myers, Florida, L SA, and to Mr. Valeri B. Darkin of Vladivostok, Russia, for allowing me to examine and describe this new species. \lr Darkin kindly presented the holotype of to the Na- tional Museum of Natural History, Smithsonian Insti- tution (USNM). LITERATURE CITED Abbott, R. T. 1968 The helmet shells of the world (Cassidae). Part 1. Indo-Pacific Molliisca 2(9):15-202, Bondarcs, I and D R5ckei, 1992 The shells of the Sa\a de Malha Bank, La C:onchig!ia 23(262):21-34. Biiuchet, P aiidP Bail, 1991. Volutes from the Sava de Malha Page 96 THE NAUTILUS, Vol. 107, No. 3 Bank: The saga of Lijria surinanwn.si.s aiul a ni-u species. soiclae, Ceritliiidae, loniiidae, Cassidae, Buccinidae, Fas- The Nautilus 105(4):159-164. ciolariiilae, Turhinellidae, urridae, Architectonicidae, Ep- Kilburii, R. 1975. Taxonomic notes on South African marine ilotiiidae, l.inndae, and Thraciidae. Annals of the Natal Mollnsca (5); including descriptii>Ms of new ta\a of Ris- Museum 22(2):.577-622. THE NAUTILUS 107(3):97-103. 1993 Page 97 Ferussac and d'Orbigny's '^Histoire naturelle generale et particuliere des Cephalopodes acetabidiferes": Dates of Publication of Plates and Text Annie Tillier Renala Boucher-Rodoni Biulogit-' dt'S lii\ fi'tebri'S niuriiis ft Malacologie LIRA 699 CNRS— Museum National d Histoire Naturelle 55, rue Buifon 75005 Paris, FRANCE ABSTRACT Ferussac and d Orbigny s "Histoire naturelle generale et par- ticuliere des Cephalopodes acclahuliferes is an essential con- tribution to teuthology, because of the numerous new species described, and because it represents the first important com- pilation of cephalopods Issued between 1834 and 1848, in 21 jisraisons, the actual dates of publication of most ot its parts were hitherto uncertain. A thorough bibliographic search has allowed us to gather new information about the publication nl this work. We can now identif > and date preciseU the '50 plates issued in 1834 and the 72 plates issued in 1835; the remaining 42 plates are dated 1839-1841 or 1839-1842, D'Orbigny's text was issued between 1839 and 1848 (livraison 12-21) The last three livraisons (1845-1848) included only text Key Words: Cephalopods; Publication dates; Ferussac and d Orbigny. INTRODUCTION During the first half of the 19th Century , a vast amount of material was collected Bn' the scientists participating in French voyages around the world, the accounts of which were published under the following titles: Voyage aux terrea Anstrales (Ic Geographe, le Natiiraliste, la Casiiarina) from 1800 to 1804 by Peron; Voyage de la Coquille. from 1822 to 1825 by Lesson; Voyage en Arnc- rique meridionalc , from 1826 to 1833 by d Orbigny; Voyage autour du rnonde (Uranie et Physicienne), from 1817-1820and Voyage de I' Astrolabe, from 1826 to 1829 by Quoy and Gaimard; Voyage de la Bunite. from 1836 to 1837 by Eydou.x and Souleyet, The specimens collected on all of these voyages were deposited in the Museum National d'Histoire Naturelle, Paris, and described not only by the collectors themselves, but also by other con- temporary zoologists such as de Blainville, Lamarck and Ferussac. These collections gave rise to numerous works that constitute a considerable contribution to the devel- opnient of natural history. The publications issued during the first half of the 19th century were delivered as a series of numbered parts, or livraisons, and several years were often necessary to com- plete a work. Such is the case for the "Histoire naturelle generale et partictdiere des Cephalopodes acelabuli- feres. . ." by Ferussac and d Orbigny, which was issued in 21 livraisons, from 1834 to 1848, and whose publi- cation was particularly lengthy and confused. Ferussac intended to publish an important and general work on Mollusca ("Histoire naturelle. generale et par- ticuliere des Mollusques'). where all the monographs written by him and other authors might be included. As early as 1825, he had asked d'Orbigny to collaborate in the publication of the "Histoire des Cephalopodes". D'Orbigny started at once to produce plates and te.xt, but in 1826 he left for an eight year long voyage to South America. It was only in 1834-35, after his return, that the first parts were distributed. Ferussac died in 1836, and it was not until 1837 that d Orbigny could recover his manuscript, and he then spent the follow ing two years updating it (1839; issue of the first part of d'Orbigny's text). DESCRIPTION OF THE WORK The "Histoire naturelle generale et particuliere des Ce- phalopodes acetalndiferes. . ." comprises 2 volumes bound according to the instructions of d Orbigny: 1 — A volume of text, starting with d Orbigny s Preface (pp. I-LVI), followed by 361 pages of taxonomic de- scriptions. The first livraison of text (livr. 12) is dated 1839 and largely postdates the Atlas. An introduction to Mollusks and general remarks on Cephalopods by Ferussac (96 pages), was issued in liv- Page 98 THE NAUTILUS, Vol. 107, No. 3 raisoiis 1 to 6 (1834-1835). A copy of this text was coii- .siilted at the "Bibliothecjue Nationale" in Paris, where it is kept in the original wrapper, on the cover of which appears: " Histoirc naturcllc ncncralc et part icxtUcre des MoUusqiies — Monographic des C'cphalopodcs Cnjpto- dibranches" . This part was originally intended to be included in the work, but d'Orbigny considered it to be obsolete. In his instructions to the liinilcr he specifies that it is not part of the work on the Cephalopods and must therefore be bound separately. 2 — An Atlas of 144 plates (90 plates named "Cryptodi- branche", 52 named "Acetabulifere", one bearing both names (Octopus pi. 23) "Cryptodibranche" and "Aceta- buliiere ', and one having neither of them (Bellerophon pi.T). Winckworth (1942) stated that a large number of plates were ready in 1826, when d'Orbigny left for South Amer- ica; but were delivered only after his return in 1834. Ferussac was at that time publishing the " Histoire na- turcllc . . . des Mollusques terrestres et fluviatiles" in collaboration with Deshayes. He also had to assume the heavy costs of the "Bulletin Universel des Sciences et de I Industrie' \ which he created and managed. How- ever, Ferussac (1831:374) himself confirms that most of the "Cryptodibranche ' plates were ready in 1831: "de- puis longtemps tous les Cephalopodes de I'ordre des Cnjptodihranches . . . composent cet ouvrage pret a etre livre a I impression, et dont les planches au nombre de pres de 80 sont tirees et enluminees". HISTORICAL ACCOUNT OF PREVIOUS RESEARCH Sherborn and Woodward (1901:75) were the first to pub- lish some dates, in a footnote to their paper on the pub- lication dates of "Mollusques terrestres et fluviatiles" by Ferussac and Deshayes. They are very concise and do not mention their sources: "Of the dates of publication of this work all that can at present be said is that the first three out of 21 liv. appeared in Sept. 1834, nos.4-9 in 1835, no.l2 in 1839, nos.13-18 in 1840 taking the work up to p. 240; and that it was finished in 1848". The copy of Ferussac and d'Orbigny s work in The Natural History Museum, London, gives some evidence of how Sherborn deduced some of the dates. \ hand- written note says: "It is obvious, as the date 1837 occurs on pp. Ill -I- 27 & 1838 on pp. V + 73 & 1839 on p. 139— that no text was published before 1837. 9 parts were published by 1835 (clearly these were all plates or part of the text to volume II Atlas, which containing no species does not concern us). Part 12 was published in 1839 & 13-18 (bringing the work up to p. 240) in 1840. \ fresh font of type was used on p. 241 (compare the 444 's in the pagination) & references to 1845 are frequent between pp. 241-361. I therefore regard \'ol. I, pp. I-LVI & 1-240 as 1840; pp. 241-361 as 1848; Vol. II text, pp. 1-96 as 1835". These considerations are dated October 1901. The last part is repeated in the bibliograplu of the Index .\ni- nialium (Sherborn, 1922: LlII), with an extra remark: "[Dates] of plates unknown". The references to 1835 most probabK come from the Bibliographie de la France, which he thouroughly con- sulted to date Ferussac and Deshav es' "Histoirc naturcllc des Mollusques. . .". He thus probabK knew that the first 9 livraisons were issued in 1834 and 1835, and that thev comprised only plates (although he did not know w hich ones) and Ferussac's preface. Thus Sherborn did not have sufficient information to date the plates, but he attempted to date the text, rel\ ing on the dates cited in the first pages of the text and in the preface by d'Orbigny. His conclusion was that the text could not have been issued before 1837 (he had no data regarding livraisons 10 and 11). As far as the dates of livraisons 12 (1839) and 13-18 ("bringing the work up to p. 240 "; 1840) are concerned, we could not determine his sources. He stated that a fresh type font was used on p. 241, in the pagination, but after a thorough examination of the different copies we could consult, we noticed that such a change in the pagination font of type actually occurs on page 273. Winckworth (1942) had some additional data that al- lowed him to propose dates for the plates issued in 1834- 1835. He possessed a bound copy of the 1834-1835 liv- raisons, although he did not have enough information to know precisely if this copy represented 9, 10 or 11 liv- raisons, and w hich of the plates were issued in 1834 and which were issued in 1835. For the text, he used: - the dates proposed b\ Sherborn and Woodward (1901); - d Orbign\ s "Mollusques vivants et fossilcs" (1845), a thorough reading of which allowed him to point out that all the plates and the text up to and including page 271 are mentioned. This led him to introduce an extra break in the text, and to date as 1848 all the text following page 27 1 ; - the text of d Orbignv himself, where 1839 is the latest bibliographic reference on page 210, and where the "Mollusques vivants et fossiles" (1845) is first mentioned on page 273; - the indications on the back wrappers of the "His- toirc. . . des mollusques terrestres et fluviatiles", with which he, however, partly disagrees. This is discussed in more detail below . ORIGINAL RESULTS AND DISCUSSION Three articles issued in the "Bulletin zoologique" and in the "Annalcs des Sciences naturcllcs" enlightened us as to the content of the first li\raisans, and encouraged us to investigate further and to retrace the history of the publication of the "Histoire generale et particuliere des Cephalopodes Acctabidifcrcs". Successively, the follow- ing data w ere gathered from 1 1 different sources: A. Tillier and R. Boucher-Rodoni, 1993 Page 99 1 — "Bibliographic de la France, ou Journal general de I'Imprimerie et de la Lihrairie' Paris, Pillet Aiiie: 23e annee, no. 39, 27 Sept. 1834, p. 615, no. 5229; 24e annee, no. 16, 18 Avril 1835, p. 246, no. 2099; no. 23, 6 Juin 1835, p. 360, no. 3058. The issue, nuini)er oi pages of text and plates of liv- raisons 1-9, are announced fnit no details are given on tfie content of tfie livraisons: Livraisons 1-3 (27 Septem- ber 1834): 30 plates and 5 sheets of text; livraisons 4-6 (18 April 1835): 29 plates and 7 sheets of text; livraisons 7-9 (6 June 1835): 25 plates and 1 sheet with front page. 2 — "Bulletin Zoologiquc". 2e section, 1835: - p. 14-16: some details on livraisons 1-3 are given; - p. 63-65: all the species figured in the plates of livraisons 4-6 are listed. 3 — "Annales des Sciences naturellcs", 2e serie, 3, Zool- ogie. Paris, Crochard 1835, p. 192 (f.l2, March). As early as March 1835, 12 livraisons of 9 to 10 plates each are announced as read\ , 9 of which were on sale (whereas La Bibliographie de la France announces their issue later in the same year). A list of all the species issuetl in the plates of the livraisons 1-9 is given. 4 — "Notice analytique sur les travaux de Zoologie de Mr. Alcide d'Orbigny" (1856). Dumeril made two favor- able reports at the Academic des Sciences, on the first eleven livraisons, on the 15 December 1834 and 29 June 1835 (see also d Orbigny s Introduction, p. II). This allows livraisons 10-11 to be dated 1835. 5 — "Registre des proces-verbaux et rapports des seances de r Academic Royale des Sciences. Institut", vol.10, 1832-1835: p. 579, 22 September 1834: the delivery of Ferussac's first three li\raisons is announced; p. 724, 29 June 1835: Dumeril indicates that the last five livraisons are issued. In the "pochette de la seance du 15 deeembre 1834" we found Dumeril's report (two handwritten pages) de- scribing the first three livraisons, i.e. an Introduction b\ Ferussac and 28 plates ("les figures de ranimal de I'Ar- gonaute argo et de son anatomic, faites par Poli . . . lithographiees par Chazal. Plusicurs especes nouvelles de poulpes, et des dessins executes a Cadix d'apres les dissections de deux tres habiles anatomistcs . . . repro- duites par Jacob". There is no trace in the "pochette of Dumeril's report on the 29 June 1835, concerning the "last five livraisons". Our sources are consistent with livraisons 1 to 1 1 being delivered in 1834-1835, the first nine three by three. The "last five livraisons" announced by Dumeril in 1835 are thus livraisons 7-9 and 10-11. 6 — "La litterature jranqaise contemporaine, 1827-1844, Dictionnaire Bibliographiqiie" b> MM. Ch.Louandre and F. Bourquelot, Paris, Felix Daguin, t.3, 1848, p. 190- 191. The " Histoire . . . des Cephalopodes ' , is announced as published in Paris, 1834-1842, in 20 livraisons, each comprising 10 plates and corresponding text, 18 livrai- sons of which were on sale. Thus we did deduce that 18 livraisons were issued from 1834 to 1842, livraison 18 being dated 1842; we know that the last livraison (21st) was issued in 1848. From the e\ idence of the type font change on p. 273 and of references to d'Orbigny (1845), we consider that livraison 20 (pp. 273-320, see below) also appeared in 1848. As for livraison 19, we date it 1845, since apparently it was not yet on sale in 1844, Init the text up to p. 271 is quoted by d'Orbigny (1845). 7 — Two complete works (Text and Atlas) from the "Bi- bliothcque Nationale": their peculiarity is that they have been marked with different stamps (Bibliotheciue Im- periale, Bibliotheque Royale or Bibliotheque Nationale). .'Ml the plates are stamped, w hereas the text bears only a few stamps. A thorough examination of these stamps in the two volumes of text revealed that they were not randomly distributed but seemed to correspond either to the beginning or to the end of a set of livraison (Table 1). ^ In one of the copies of the Atlas, all the plates are marked "Bibliotheque Royale", except 32 that are stamped "Bibliotheque Imperiale". These 32 plates cor- respond to most of the plates that we consider as be- longing to livraisons 1-3. We conclude that the latter set constitutes a homogenous lot, which can be dated 1834, except for 4 plates of Argonauta, which we are sure belong to livraisons 7-9 (Bull.lool. 1835 and Ann.Sci.nat. 1835). All the plates in the second Atlas are stamped "Biblio- theque Royale", except 46 that are marked "Biblio- theque Nationale" (BN). Knowing what is included in the first 9 livraisons, and from w hat we infer is included in li\raisons 10-11 (confirmed b\ our source no. 8), we deduce that the stamp "Bibliotheque Nationale" was used for all the plates of livraisons 4-6, for almost all the plates of livraisons 10-11 and for one plate of livraisons 7-9. 8 — An incomplete bound copy of the work present at the National Museum of Wales, the title page of which is "Histoire tiaturelle generale et particuliere des Mol- lusques", contains the Introduction and general remarks on Cephalopods by Ferussac, and 101 plates. It corre- sponds perfectly to the first 11 livraisons, issued in 1834- 1835, as detailed in the preceeding paragraphs. Only one discrepancy was observed: plate 25 of Octopus, printed by Benard and stamped as BN, that we consider to be part of the livraisons 10-11, is not included in this copy. The National Museum of Wales also has a complete copy with handwritten notes by Hovle, and a series of 77 plates, unbound, corresponding to an incomplete set of livraisons 1 to 11. Plate 25 of Octopus is present in this last series of plates. 9 — The complete copy present at The Natural History Museum, London, bears two handw ritten notes. One, by Sherborn on the first page of vol.1 (Text), dates the text. Its content has been discussed above. The other, by E.A.Smith on the first page of vol.11 (Atlas), says: "Mr. Sherborn has been unable to discover anything def- inite w ith regard to dates of publication of any of these plates . THE NAUTILUS, Vol. 107, No. 3 Table 1. Details of text livraisons, with all available information on the .sheets of each livraison, inclusion (P) or not in the BIMM cop\, the stamped pages in the two complete works (I and 2) at the BN and the protiahle date of issue BR = Bibliotheque Royale; BN = Bililii)tliei|ue Xationale. Ni umber P = BIMM Sheets Pages stamps of sheets cop\ Livraison \ ear 1-6 1 BR 1-48 6 P 12 1839 7-12 49/BR1-96 BRl 5 13 1839-41 13-18 97/BRl + BR2-144/BR1 5 14 1840-41 a-c I BRl-XXIV 19-20 145-160/BRl 5 P 15 1841 21-24 161/BR1-192'BR1 4 P 16 1841 25-29 193/BNl + BR2-232/BR1 5 P 17 1841 d-g XXV/BRl-LVI 4 P 18 1842 30-34 233/BN1-272 5 19 1845 35-40 273/BN1-320 6 20 1848 41-45 321/8X1-361 5 21 1848 10 — An iiiconiplete bound copy of the work, from the library ol the "Laboratoire de Biologie des Invertebres marins et Malacologie" (BIMM) at the Museum National d'Histoire Naturelle (MNHN) comprising: the introduc- tion and preface by d'Orbigny (pp. I-LVI), the text p. 1-48 and 145-232, 30 plates, all coming from the same printer (Imp. Lemercier, Benard et C). We interpret this BIMM copy as corresponding to livraison 12 (pp. 1-48) and livraisons 15-18 (pp. I-LVI; 145-232), i.e. five livraisons, each of 4 to 6 sheets of text and 6 plates (livrai.son 17 is announced on back wrappers as comprising 5 or 6 sheets of text and 6 plates). This interpretation also takes into account the observations on the other copies of the work. There remain 12 plates of the Atlas that belong neither to the first 11 livraisons nor to the BIMM copy. Printed well before the text, they are thought to have been issued in livraisons 13-14 (i.e. two livraisons of 6 plates each). Thus all the plates were delivered before 1845 (cita- tions in d'Orbigny, 1845). The livraisons 19, 20 and 21 include only text. 11 — "Notice analytique siir les travanx zoologiques el paleontologiques de Mr.Alcide d'Orbigny" (1844). D'Orbigny states that his manuscript was finished in 1839. However, later, the death of the publisher interrupted the printing of the text, and caused several trials. This accounts for the gaps in the publication of livraisons from 1842 to 1845, and from 1845 to 1848. TEXT The data gathered from the incomplete BIMM copy, together with the .stamps on the text in the two complete copies of the BN have led us to propo.se that the text was i.ssucd in 10 livraisons. Livraison 12, pp. 1-48, 1839. This livraison is included in the incomplete copy of the BIMM. The first page is stamped "Bibliotheque Royale" in the BN. Its issue is announced on the back wrappers of "Mollttsques tcr- restres et fluviatiles" (VViiukworth, 1942). After three years \\ itiiout aii\ liv raison being issued, it .seems obvious that d'Orbigny, who had started to rewrite his manu- script in 1837 (cf. Introduction p. Ill), had published the initial part of it in this livraison. It also seems evident that the Introduction, which cites works issued in 1840- 1841, cannot have been issued in 1839. Livraison 13, pp. 49-96, 1839-1841. The stamp "Biblio- theque Royale occurs on the first and last page of this livraison, which constitutes 5 sheets. Further supporting evidence is that this livraison is not included in the incomplete BIMM copy. The announcements on the back wrappers of "Molhisques terrestres et fluviatiles" (Winckworth, 1942) state that 16 livraisons were issued in 1839-1841. Lacking any further information, we date livraison 13 as 1839-184L Livraison 14, pp. 97-144, 1840-1841. The stamp "BiWiO- theque Royale " is on p. 97 of both copies in the BN, and also on p. 144 in one of these copies. The break at page 97 is thus confirmed twice. The incomplete BIMM copy lacks pages 49-144, confirming the end of the livraison. A reference to 1839, on p. 139, allows livraison 14 to be dated 1 840- 184 1. Livraison 15, Introduction, pp. I-XXI\', and text pp. 145- 160, 1841. This livraison is included in the BIMM copy, and is stamped on p. I and p. 160 of the BN copy. Stamps in the text corresponded to a livraison of only two sheets, which seemed insufficient to us. When the first part of the introduction is included, a livraison of 5 sheets is obtained, stamped at the beginning and at the end. This is consistent with the issue of the Introduction in two separate livraisons (stamps on p. I and on p. XXV). Ref- erences to publications issued in early 1841, as well as indications on the abo\ e-quoted w rappers lead us to pro- pose the date ot 1841 tor livraison 15. Livraison 16, pp. 161-192, 1841. This livraison is in- cluded in the BIMM copy. The BN copy is stamped at the beginning (p. 161) and at the end (p 192). Livraison 16 was issued between two other livraisons ol 1841 and is therelorc tlated 1841. Livrai.son 17, pp 193-232, 1841 This livraison is A. Tillier and R. Boucher-Rodoni, ] [993 Page 101 Table 2. Pu bhcation dates of the Atla s pi; ates, with identih- Table 2. ('oiituiued cation of the = stamps on correspond the BibhotI il1(J liv^'^icmi PI = plate number; stp ipies; BI = Biblio- 1 lilj^ liv leque Bihho National e cc PI sip Li\ rai.son Date tlieqiie Impcnaie; ti.\ = •tlieque .Nationaie 0 3 7-9 12/15- 18 March 1835 1839-42 PI stp Livraison Date 4 Sepia 1 2 BN 12/15- 4-6 18 1839-42 Octopus 1 BI 1-3 (1834) 1826 March 1835 2 BI 1-3 22 Sept. 1834 BN 4-6 March 1835 3 BI 1-3 22 Sept. 1834 3 BI 1-3 22 Sept. 1834 3/2° BI 1-3 22 Sept. 1834 3/2° BN 4-6 March 1835 4 BI 1-3 (1834) 1826 3/3° BN 4-6 March 1835 5 BI 1-3 22 Sept. 1834 4 7-9 March 1835 6 BI 1-3 22 Sept. 1834 4/2° 7-9 March 1835 6/2° BI 1-3 22 Sept. 1834 5 BN 4-6 March 1835 6/3° BI 1-3 22 Sept. 1834 5/2° 7-9 March 1835 7 BI 1-3 22 Sept. 1834 6 10-11 29 June 1835 8 BI 1-3 22 Sept. 1834 6/2° 10-11 29 June 1835 9 BI 1-3 22 Sept. 1834 7 7-9 March 1835 10 BI 1-3 22 Sept. 1834 8 7-9 March 1835 11 7-9 1835 9 10-11 29 June 1835 12 BI 1-3/4 -6 22 Sept. 1834 10 7-9 March 1835 13 BI 1-3/4 -6 22 Sept. 1834 11 BN 10-11 29 June 1835 14 BN 1-3/4 -6 22 Sept. 1834 12 BN 10-11 29 June 1835 15 BN 1-3/4 -6 22 Sept. 1834 13 BN 10-11 29 June 1835 16 BN 7-9 March 1835 14 12/15- -18 1839-42 17 7-9 March 1835 15 13-14 1839-41 18 BN 10-11 29 June 1835 16 13-14 1839-41 19 BN 10-11 29 June 1835 17 12/15- -18 1839-42 20 BN 10-11 29 June 1835 18 12/15- ■18 1839-42 21 BN 13-14 1839-41 19 12/15- -18 1839-42 0 0 BN 10-11 29 June 1835 20 12/15- -18 1839-42 23 13-14 1839-41 21 12/15- ■18 1839-42 24 BN 10-11 29 June 1835 22 12/15- -18 1839-42 25 BN 10-11 29 June 1835 23 12/15- -18 1839-42 26 12/15- ■18 1839-42 24 12/15- ■18 1839-42 27 13-14 1839-41 25 12/15- -18 1839-42 28 13-14 1839-41 26 12/15- -18 1839-42 29 13-14 1839-41 27 12/15- -18 1839-42 Eledone 1 BI 1-3 22 Sept. 1834 Sepioteuthis 1 BN 4-6 March 1835 1/2° BI 1-3 22 Sept. 1834 ."> BN 4-6 March 1835 2 BI 1-3 22 Sept. 1834 3 BN 4-6 March 1835 3 12/15- 18 1839-42 4 BN 4-6 March 1835 Argonauta 1 BI 1-3 22 Sept. 1834 5 BN 10-11 29 June 1835 1/2° BI 1-3 22 Sept. 1834 6 12/15- -18 1839-42 1/3° BI 1-3 22 Sept. 1834 7 12/15- -18 1839-42 1/4° BI 1-3 22 Sept. 1834 Loligo IC BI 1-3 22 Sept. 1834 1/5° BI 1-3 22 Sept. 1834 lA BN 10-11 29 June 1835 2 BI 7-9 March 1835 2 BN 4-6 March 1835 3 BI 7-9 March 1835 BN 4-6 March 1835 4 BI 7-9 March 1835 4 BN 4-6 March 1835 5 BI 7-9 March 1835 5 BN 4-6 March 1835 6 12/15- 18 1839-42 6 7-9 March 1835 Bellerophon 1 BI 1-3 22 Sept. 1834 7 7-9 March 1835 2 12/15- ■18 1839-42 8 7-9 March 1835 3 12/15- 18 1839-42 9 7-9 March 1835 4 13-14 1839-41 10 7-9 March 1835 5 12/15- 18 1839-42 11 BN 4-6 March 1835 6 12/15- 18 1839-42 12 BN 4-6 March 1835 7 12/15- ■18 1839-42 13 BN 7-9 March 1835 Cranchia 1 2 BN 4-6 7-9 March 1835 March 1835 14 15 16 BN BN 4-6 7-9 4-6 March 1835 March 1835 March 1835 Sepiola 1 BI 1-3 22 Sept 1834 17 BN 4-6 March 1835 Page 102 THE NAUTILUS, Vol. 107, No. 3 Table 2. C^oiiliiiiicd PI stp Livraison Date 18 10-11 29 June 1835 19 7-9 March 1835 20 13-14 1839-41 21 BN 10-11 29 June 1835 22 12/15- IS 1839-42 23 13-14 1839-41 24 13-14 1839-41 Loligopsis 1 BN 4-6 March 1835 2 7-9 March 1835 3 7-9 March 1835 4 1.3-14 1839-41 On>choteuthis 1 Bl 1-3 22 Sept. 1834 2 BN 4-6 March 1835 3 BN 4-6 March 1835 3/2° BN 4-6 March 1835 4 BN 4-6 March 1835 5 BN 4-6 March 1835 6 BN 10-11 29 June 1835 7 BN 10-11 29 June 1835 8 BN 10-11 29 June 1835 9 BN 10-11 29 June 1835 10 BN 10-11 29 June 1835 11 BN 10-11 29 June 1835 12 12/15- ■18 1839-42 13 12/15- ■18 1839-42 14 12/15- ■18 1839-42 Ommastrephes 1 12/15- •18 1839-42 r> 12/ IS- IS 1839-42 eluded in the BIMM copy. Both BN copies are stamped at the beginning (p. 193), and one of them also at the end (p. 232). It is comprised of 5 sheets. The back wrap- pers of "Molln.sqties terrestres et fluviatiles (1841), state: "la 17eme livraison sera raise en vente d la fin d'aout" (Winckworth, 1942:35). The date and content of this Hvraison are given as 5 to 6 sheets of text and 6 plates. Our conclusions concerning this livraison contradict Winckworth, who disbelieved the announcement on the back wrappers. Livraison 18, Introduction pp. XXV-LVI, 1842. Page XXV is stamped in the BN copy. This livraison is included in the BIMM copy. This second part of the Introduction comprises 4 sheets. We include it in livrai.son 18 since we know that livraison 17 was made ol 5 or 6 sheets. The date 1842 is given by Louandre and Bourquelot (1848) and is supported by five of our sources. Livraison 19, pp. 233-272, 1845. This livraison is not included in the BIMM copy. It is stamped on the first page (p. 233) in the BN copy. The date 1845 is based on the fact that livraison 19 is not mentioned as issued in the analyses of the "Utterature franqaise" of the period from 1827 to 1844 (Louandre & Bourquelot, 1848), but the pages of this livraison are (luoted in "Mollusqiics vivants et fossiles" d'Orbigny (1845). Livraison 20, pp 273-320, 1848. This livraison is not included in the BIMM copy. It is stamped on the first ()age (p. 273) in the BN copy. A different type font was used for the pagination, starting on p. 273 (especially the 5's), References to the "Mollusques Vivants et Fossiles" (1845) appear in the te.xt, confirming that this livraison was published after 1845. It was probably issued with the 21st livraison in 1848. Livraison 21, pp. 321-361, 1848. This livraison is not included in the BIMM copy. The BN copy is stamped on the first page (p. 321). "Mollusques Vivants et Fos- siles" (1845) is cited in this livraison. It is the last livraison, and is undoubtedly dated 1848. ATLAS The "Bulletin Zoologique" and the "Annales des Sci- enees Naturelles ' list all the species figured in the plates issued in livraisons 1-9. The incomplete copy at Cardiff gathers the first eleven livraisons, as detailed in sources 2, 3, 8 (see above). Plates included in the BIMM copy (source 10; livraisons 12 and 15-18) could not be dated more precisely than 1839-1842. Details concerning the dates of publication and the corresponding livraison of each plate are summarized in table 2. Our conclusions concerning the issue ot the Atlas are as follows: Livraisons 1-3, 22 September 1834: 30 plates. Livraisons 4-6, March 1835: 29 plates (4 of which were already issued in livraisons 1-3). Livraisons 7-9, March 1835: 25 plates. Livraisons 10-11: 29 June 1835: 22 plates. Livraison 12, 1839: 6 plates. Livraisons 13-14, 1839-1841: 12 plates. Livraisons 15-18. 1841-1842: 24 plates. Two plates issued in livraisons 1-3 (Octopus pl.l and 4) had been distributed as early as 1826 by d'Orbigny hiuLself (footnote p. 18: "A la fin de 1825, nous avons fait lithographier nos planches representant les figures de I'Octopus Cuvierii et de VO.Lcchcnaultii. et elles out ete distribuees a beaucoup de personnes ). CONCLUSIONS A comparative analysis of Ferussac and d'Orbigny s work and that of d'Orbigny (1845) confirms that the dates iiuoted by d Orbigny are not reliable. As an example, Cranchia plate 1 includes three species: C. cardioptera, C. minima and C. scahra. In d'Orbigny (1845), they are cited as figured in plate 1, and dated respectively 1826 (p. 390), 1830 (p. 351 ) and 1839 (p. 240). Such inaccuracy is common in d'Orbigny s dates, and was often the case at that time. Our bibliographic searches allowed us to acertain the precise dates of i.ssue of the 102 plates of the .Mlas that were i.ssued in 1834 and 1835, as well as their compo- sition. The dates of issue of the remaining plates are 1839-1842 (livraisons 12-18), but we can find no details on the actual content of each livraison. A. Tillier and R. Boucher-Rodoni, 1993 Page 103 In the publication of the text, we recognize three main stages: 1. 1834-1835. Livraisons 1-11: introduction and gen- eral remarks by Ferussac. 102 plates of the Atlas. No descriptions. 2. 1839-1842. Livraisons 12-18: Introduction and gen- eral remarks by d'Orbigny. Text pp. 1 to 232. Atlas: the remaining 42 plates. 3. 1845-1848. Livraisons 19-21: last part of the text, pp. 233-361. No plates. The above data and conclusions are summarized in two tables. Table 1 provides information concerning the livraisons of text and corresponding dates. Table 2 pro- vides the publication date for each plate, and associates the plate with a corresponding livraison. ACKNOWLEDGMENTS We gratefully acknowledge the help and advice of Dr.PMordan, of The Natural Histor\ Museum, London, throughout our bibliographic search, and for critical re- view of the manuscript. Special thanks to Mr.J.R.Kenyon, librarian of the National Museum of Wales, who pro- vided most helpful documentation. We are grateful to our colleague Alain Foubert for his help in preparing the tables. LITERATURE CITED D Orbigny, A 1844. Notice analytique sur le travau.x zool- ogiqueset paleontologiquesde M. Aicided Orbigny. Impr. Cosson, Paris, 48p. D Orbigin , .\. 184.5-1847. Mdilu.sqiit-s \i\ants et tossilt-s, ou description dc toutes ies especes de coiiuiiles et de Moii- usques .-K Deialiass Libr , Paris 605p , :35 pis. D'Orbign\. .\ 1856 Notice analytique sur Ies travaux de geologie. de paleontoiogie et de zooiogie de M. Alcide d'Orbigny, 1823-1856. Impr. L.S. Crete, Corbeil. 60pp. Ferussac, A. E. de. 1831. Bulletin des Sciences Naturelles et de Geologie, 2e section, 24: 373-374 Ferussac, A. E, de and A. d'Orbign\. 1834-1848. Histoire naturelle generate et particuliere des Cephalopodes ace- tabuiiferes vivants et fossiles. J.-B Bailliere Libr., Paris. Tome Premier, — Texte: Ivi -I- 361p Tome Second. — .At- las: 144 pis, Sherborn, C. D. 1905. Ou tlie dates ot publication of d Or- bigny's "Moll. viv. et loss. ", "Paieont. univ." and "Paleont. etrangere". Journal of ConchoJogy, 11 (6): 169-170, Sherborn, C. D 1922-32 Index Animalium sive index nom- inum quae ab AD MDC.'CLVIII generibus et speciebus animalium imposita sunt Sectio Secunda A kaiendis la- nuariis, MDCCX4 usque ad finem Decembris, MDCCCL. British Museum (Natural Histor\ ), Loudon, cxlvii -I- 7,056 + 114p Sherborn, C. D. and B. B. Woodward, 1901 On the dates of publication of the "Histoire naturelle generale et parti- culiere des moilusques terrestres et fluviatiles and the "Tableaux systematiques des Animaux moilusques", by the Barons Ferussac and G, P, Deshayes Annals and Magazine of Natural History, 7 (8): 74-76, VVinckworth, R 1942 Notes on the publication of Ferussac and Orbigny s Histoire des Cephalopodes. Proceedings of the MaiacoJogical Society. 25 (1): 34-36. THE NAUTILUS 107(3):104-106, 1993 Page 104 Alvania valeriae (Gastropoda: Rissoidae), a New Species from off Southeastern Brazil Ricardo Silva Absalau Dcpailanu'nto de Zoologia Institiilo df Biologia Univcrsidade F"edt'ral do Km Janeiro lllia do FundSo, 21949 Kio de Janeiro, RJ. Brazil ABSTRACT Alvania (Pimctulum) valeriae new species, is found off the coast of Rio de Janeiro and Espirito Santo States, soutlieastern Brazil. It differs from other species of Alvania occurring off the Brazilian coast in having more numerous axial ribs (about •36 on the body whorl). Axial ribs are more pronounced than the 12-14 spiral threads, and nodules are not formed at their intersections. Key words: Gastropoda, Rissoidae, Alvania. Brazil INTRODUCTION Alvania (Punctiilum) valeriae new species was collected on the continental shelf off Rio de Janeiro and Espirito Santo States, in ciredgings taken during the oceanograph- ic operations Espirito Santo I (July to September, 1984), and (>abo P'rio VII (March to July 1983), as part of routine sampling by the Brazilian Navy to obtain basic ocean- ograpliic data on the coastal and oceanic regions off Bra- zil. The malacofauna found during these operations was characterized by numerous small mollusks, among which the Rissoidae were prominent (Absalao, 1989). The first descriptions of rissoids that occur off Brazil, or would later have their ranges extended to Brazilian waters, are those of d'Orbigny (1842), Watson (1886), who studied the material collected by the H.M.S. Chal- lenger, and Dall (1889), who worked with material col- lected by U.S.S. Blake (Romer & Moore, 1988). Since 1966, malacological material from off the Brazilian coast has been obtained on a more regular basis, thanks to the dredging operations of the Brazilian Navy (Absalao, 1986). The ininiit<- specimens (< 5 mm) have received almost no attention. The Rissoidae especially "have proved to be very difficult to classify at all levels because of their small size, diverse form, and confusion caused by con- vergence, especially in shell characters" (Fonder, 1985). Ahbcitt (1974) stressed that the entire group was in need of revision before species could be assigned to subgenera. The revision of Coan (1964) was mainly a bibliographic compilation. Ponder (1985) made the last and most com- plete revision of this family at the generic level. ABBREVIATIONS IBUFRJ = Instituto de Biologia da I niversidade Federal do Rio de Janeiro, Brazil. MORG = Museu Oceanografico da Funda^ao Uni- versidade de Rio Grande, Brazil. USNM = National Museum of Natural History, Smithsonian Institution, USA. MZUSP = Museu de Zoologia da Universidade de Sao Paulo, State of Sao Paulo, Brazil. MNUFRJ = Museu Nacional do Rio de Janeiro, State of Rio de Janeiro, Brazil. MNHN = Museum National d'Histoire Naturelle (Paris), France. SYSTEMATICS Family Rissoidae H & A. Adams, 1854 Genus Alvania Risso, 1826 Type species: Turbo cimex Linnaeus, 1758. There are three previous!) described species of Al- vania that occur off the Brazilian coast: A. auheriana (Orbigny, 1842), A. deliciosa Jeffreys, 1884 and A. xan- thias (Watson, 1885). The first occurs in shallow waters and the latter two at greater depths. Alvania {Puncluhim) valeriae Absalao, new species (figures 1-3) Description: Shell minute, 1.6 to 2.4 mm long, white ovate-conic, with rounded whorls. Suture well impressed, slightly canalicuiated. Pri)toconch multispiral, smooth. R. S. Absalao, 1993 Page 105 with 2'/2 whorls. Apical angle 52-65° Teleoconch with about 3(i gently curved axial ribs and 12-14 spiral threads on body whorl. Axial ribs override spiral threads without forming nodules at intersections. Base rounded with 6- 8 spiral threads. Intritacalx present over entire teleo- conch. Terminal \ari\ absent. L'mbilicus small, chink- like, well defined. Operculum and ratlula unknown. Type locality: Off southeastern Espirito Santo State, southeastern Brazil (2U°3S'12"S, 40°16'12"W), 38 m depth, 22.5''C and 36.7 ppt salinity, Brazilian Naval Research Vessel Almirante Saldanha, station 6449, August 23, 1984. Type material: Holot\ pe IBUFRJ 1726, 1.74 mm height, 1.02 mm width. Parat>pe 1, MORG 26986 off north- eastern Rio de Janeiro State, Brazil (23°08'18"S, 41°03'18"W), 85 m depth, 16.7°C and 36.2 ppt salinity, Almirante Saldanha, station 6417, August 29, 1984, 1.80 mm height, 1.04 mm width. Paratype 2, USNM 860503, off northeastern Rio de Janeiro State, Brazil (22°35'00"S, 40°50'00"W), depth 77 m, 19.2°C and 36.1 ppt salinity, Almirante Saldanha, station 6422, August 16, 1984, i.80 mm height, 1 04 mm width. Paratype 3, MZUSP 27716, off Rio de Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Almirante Saldanha station 6174, depth 92 m, 15.6°C and 35.5 ppt salinity, March 29, 1983, 1.80 mm height, 1.04 mm width. Paratvpe 4. MNUFRJ 5978, off Rio de Janeiro State, Brazil (23°16'.50"S, 43°02'40" W), Almirante Saldanha station 6174, depth 92 m, 15.6°C, and 35.5 ppt salinity, March 29, 1983, 2.40 mm height, 1.28 mm width. Paratype 5, IBUFRJ 1727. off Rio de Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Al- mirante Saldanha station 6174, depth 92 m, 15.6°C, and 35.5 ppt salinity, March 29, 1983, 2.16 mm height, 1.20 mm width. Paratype 6. MORG 27681, off Rio de Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Al- mirante Saldanha station 6174, depth 92 m, 15.6°C, and 35.5 ppt salinity, March 29, 1983, 1.88 mm height, 1.04 mm width. Paratype 7, MORG 27682, off Rio de Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Al- mirante Saldanha station 6174, depth 92 m, 15.6°C, and 35.5 ppt salinity, March 29, 1983, 1.80 mm height, 1.04 mm width. Paratype 8, IBUFRJ 1728, off Rio de Janeiro State, Brazil '(23°16'50"S, 43°02'40"W), Al- mirante Saldanha station 6174, depth 92 m, 15.6°C, and 35.5 ppt salinity, March 29, 1983, 2.00 mm height, 1.12 mm width. Paratype 9, MNHN, off Rio de Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Almirante Sal- danha station 6174, March 29, depth 92 m, 15.6°C, and 35.5 ppt salinity, 1983, 1.84 mm height, 1.08 mm width. Etymology: This species is dedicated to my wife, Mrs. Valeria Gomes Veloso. Range: Records of A. valeriae are restricted to the re- gion between northern Rio de Janeiro State and southern Espirito Santo State, southeastern Brazil. Discussion: Alvania colomlriana is compared to A. val- eriae because both species occur in the same biogeo- graphic region. However, they are not sympatric. As i Figures 1-3. Alvania valeriae new species. 1. Holotvpe. IliL'FRJ 1727; 120X, 2. Ornanu-ntatiDn details, 240x,' 3. Protoconch, 240 x. Alvania is largely a European group, comparisons with those warm-water species were included Alvania auheriana. A. deliciosa and A. xanthias have respectively 14, 14 and 12-14 axial ribs, whereas A. vale- riae has 32-40 ribs. This character alone is sufficient to discern it from the other species occurring in Brazilian Page 106 THE NAUTILUS, Vol. 107, No. 3 waters. In addition, /\. auberiana lias 5 spiral threads on its teleoconch, A. deliciom has 10, A. xanthiaa lias only 4 spiral threads that are restricted to the base, whereas A. valcriae has 12-14 spiral threads (figures 1 and 2) on its teleoconch. Alvania colombiana Romer and Moore, 1988 differs from A. valeriae in having only 26 axial ribs, which disappear on the base of the shell, and 7-9 spiral grooves. Alvania valeriae has =36 (32-40) a.xial riblets that extend onl\ slightly over the body whorl without reaching the umbilical region of the shell, as well as 12-14 spiral threads. Alvania porcupinae (Jeffreys, 1884) has a smooth pro- toconch very similar to that of A. valeriae, =32 axial ribs that disappear smoothly towards the umbilical re- gion, while A. valcriae differs in having =36 (32-40) axial ribs that enter a little beyond the edge of the pen- ultimate whorl. The diameter of the last two whorls increases greatly in A. porcupinae, but little in A. valeri- ae. Alvania zylensis Gofas and Waren 1982, and A. im- perspicna Palary, 1920 can be easily distinguished from A. valeriae on the basis of protoconch sculpture. The protoconch of A. valeriae is smooth, that of A. zylensis has 12 zigzag spiral lines that intersect, forming a retic- ulated pattern, while that of A. imperspicua has 7-8 spiral lines that also zigzag but do not intersect. In ad- dition, A. imperspicua has protruding nodules on the teleoconch, which are formed by the junction of axial riblets and spiral lines. In contrast, A. valeriae lacks such nodules. Although A. valeriae is similar in shape to A. coni- pacta, this latter species has fewer axial riblets (20-30) and spiral threads (6-8) than the former, which has =36 (32-40) axial riblets and 12-14 spiral threads. Alvania compacta also has nodules where axial and spiral orna- mentation cross. Alvania valeriae is unicjue among its southwestern At- lantic congeners in having a large number of axial ribs that clearly dominate the 12-14 spiral threads. ACKNOWLEDGMENTS 1 am grateful to Prof. E. C. Rios from the Museu Ocean- ografico da Funda^ao Universidade do Rio Grande, who has provided bibliographic material. I am also much obliged to the Laboratorio de I'ltraestrutura Olular e Microscopia Eletronica do Instituto de Hiofisica Carlos Chagas Filho, of the Universidade Federal do Rio de Janeiro for the SEM micrograph, and to Dr. Jose Hen- rique Leal, from the Rosenstiel Institute of Marine Sci- ences, and to Dr. M. G. Harasewych from the Smith- sonian Institution for their comments on the manuscript. This work was partially supported by CNPq. grant no. 802240.87-9. LITERATURE CITED Abbott, H T 1974 ,\nierican Seasheiis, 2rnl cd \'aii .NostraiKl Reinhoid, N.Y., 663 p. .\bsalao, R. S. 1986 Moluscos da Comissao Oceanografica Geocosta Rio I, RJ, Brasil Revista Brasileira de Biologia 46(1):27-31. Absalao, R. S. 1989. Padroes distributives e zoogeografia dos moluscos da plataforma continental brasileira. Parte III. Comissao Oceanografica Espirito Santo I. Memorias do Instituto Oswaldo C^ruz 84, Suplemento I\ '1-6 C^oan, E. 1964. A proposed revision of the Rissoacean families Rissoidae, Rissoinidae, and Cingulopsidae (Mollusca:Gas- tr(ipoda). The \'eliger 6(3)464-171. Dall, W, H. 1889, Reports on the results of dredging in the Gulf ol Mexico (1877-78) and in the Caribbean Sea (1879- 80) by the U.S. Coast Survey Steamer "Blake". 29 Report on the MoUusca, part II. Gastropoda and Scaphopoda. Bulletin of the Museum of Comparative Zoology IS 1- 492, pis. 10-40. d'Orhigny, A. 1842. Mollusques, X'of 1 (part). In: R, de la .Sagra (ed). Histoire Plusique, politique et naturelle de rile de Cuba. Bertrand. Paris, p 209-26-1 + .\tlas. pis. 1- 28 4- II his. Gofas. S. and A. Waren. 1982. Taxoriomie de yueiques Es- peces du Genre Alvania (Mollusca, Gastropoda) des Cotes Iberiques et Marocaines, Bolletino Malacologico 18(1-4): 1-16, Ponder. W, F. 1985, A review of the Genera of the Rissoidae (Mollusca: Mesogastropoda: Rissoacea). Records of the .Australian Museum, Su|)plenieiit 4:1-221. Rios, E. C, 1985. Seasheiis of Brazil. Museu Oceanografico da Funda9ao I'niversidade de Rio Grande, 331 p. + 102 plates. Romer, N. S. and D. R. Moore. 1988. A new species of A/t)ania (Rissoidae) from the West Indian Region The Nautilus 102(4): 131-133. Watson, R. B 1886, Report on the Scaphopoda and Gaster- opoda collected In HM.S. Challenger during the \ears 1873-1876, Voyage of the H,M,S, Challenger. 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Box 7279, Silver Spring, MD 20907-7279, USA. Change of address: Please inform the publisher of your new address at lea.st 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 Trophon Corporation, 8911 Alton Parkway, Silver Spring, MD 20910. Second Class postage paid at Silver Spring, MD and additional mailing offices. POSTMASTER: Send address changes to; THE NAUTILUS P.O. Box 7279 Silver Spring, MD 20907-7279 TH E€7NAUTI LUS "Tine Bioloaica! Uboralofy/ ,ole 0C3. ■.orraphic Institution y^/^^^ ^Qy J^umber 4 '""" March 16, 1994 IVIAR 2 5 1994 ISSN 0028-1344 CONTENTS Woods Hois, MA Oi^t. Robert Robertson Protoconch Size Variation Along Depth Gradients in a Planktotrophic Epitonium 107 John I. Scheide Effect of Low Water Temperature on Ion Balance in the Phyllis N. Bonaminio Zebra Mussel, Dreissena pohjjuorpha, and the Unionid Mussel, Lampsilis radiata 113 Emily H. Yokes The Muricid T\ pes of Frank Collins Baker 118 J. P. Pointier Invasion of the Rivers of the Littoral Central Region of R. N. Incani Venezuela by Thiara granifera and Melanoides C. Balzan tuberculata (Mollusca; Prosobranchia: Thiaridae) and the P. Chroseiechowski Absence of Biomphalaria glabrata, Snail Host of S. Prypchan Schistosoma inansoni 124 Cristian F. Ituarte Corbicula and Neocorbicula (Bivalvia: Corbiculidae) in the Parana, L'ruguay, and Rio de La Plata Basins 129 Book Review 136 THE NAUTILUS 107(4):107-112, 1994 Page 107 Protoconch Size Variation Along Depth Gradients in a Planktotrophic Epitonium Roberl Robertson DfpartiiU'Hl ol Malacolog\ Tlif Acaiii-nn ol Natural Sciences 190(1 Benjamin Franklin Parkway Philadclpliia, I'A 19103-1195, USA ABSTRACT The protoconch length of Epiluniiim t'chinaticuata ranges from 330 to 790 M'li '" tliP northern Bahamas. The length increases along depth gradients from near 0 m to 40-52 m. Intermediates are from intermediate depths. One planktotrophic species is involved. A possible explanation is given. Svstematists should be aware that the phenomenon ma\ occur in other plankto- trophic epitoniids or other marine prosobranchs Kill nurds: Epitiinuiiu. [imtoconch. size \ariation. INTRODUCTION ". . . the protoconch may provide the best guide to rela- tionships [within the Epitoniidae] yet. Of course ecological factors . . nia> prove to influence protoconch form (as has been established in other gastropod groups) but this remains to be show n lor the Epitoniidae Kilburn. 19iS5: There is scattered mentinii in the literature on marine prosobrancli gastropods of one species having proto- conchs that differ from shallow and "deep" water. None of these cases seems to have been documented with mea- surements, descriptions and illustrations. In some cases different species perhaps were involved. In this paper, I show that in a single planktotrophic species of Epitonium much variation in protoconch size occurs in the northern Bahamas. Size (length) increases with increasing depth (and possibK also with exposure to wave action). Populations from Grand i5aliama sam- pled from 9, 10, 24, and 40 m, and from 1 I and 52 m at Abaco all have larger protoconchs than do those from nearby shallow sites (0-2 m). The epitoniid studied is Epitonium echinaticosta [not cchinaticostum] (d'Orbigny, 1842) [type locality: St. Thomas, Virgin Islands; northern known limit: Bermuda (.ANSP 100824); .southern know n limit: PYrnando de No- ronlia, off Brazil (Leal, 1991)]. Because of its distinctive shell morphology, especially its variably disjunct anil commonly "open" teleoconch coiling (Rex & Boss, 1976), and its 8-15 variably high and undulate axial ribs on the last whorl (Robertson, 198.3b), Dall (1889) created for this species a suligenus Cycloscala, which has sometimes been ranked as a genus. Species of Cycloscala. w hich occur in the Indo-Pacific as well as the western Atlantic, have been separated by degree of w horl disjunction and rib counts by de Boury (1911), Woodring (1928), Kilburn (1985) and DuShane (1990). Familiarity with a living population (Robertson, 1983a) clearly reveals that at least in the Bahamas there is just one species, with intergrades between the extremes, as concluded by C;lench and Turner (1951). They rec- ognized only one western Atlantic species of Cycloscala, and I concur. In Bahamian £. echinaticosta, the frequencies of the different teleoconch coiling types vary little at different depths (Robertson, uniiublished) and thus do not affect the conclusions here. There is already some inlormation about the coelen- terate host, feeiling and reproduction of £. echinaticosta in the Bahamas (Robertson, 1983a). At least in shallow water, the host is the actinian sea anemone Bunodeopsis [or Viatrix] globulifera (Duchassaing, 1850), commonly living on the leaves of turtle grass [Thalassia testudinum (Banks and Solander ex Kiinig)] (It also occurs on Syr- ingudium, Halimcda, Pcnicillus. and Acetahularia.) The lower end of the bathymetric range of E. echinaticosta is given by Abbott (1974) as 200 fathoms (366 m). Wheth- er living aniinals occur at that depth, and whether the Epitonium remains specific to B globulifera are un- known. The Epitonium and anemone are most abundant in the Freeport canals. How is it known that E. echinaticosta is planktotroph- icr* There is Leal s inulocumented statement that off Bra- zil "planktotrophic ile\ elopment is indicated. More def- initely, there is direct e\ idence of planktotrophv at least in Bahamian shallow waters: eggs of this species have Ijeen shown to be 98-106 ^m in diameter (mean = 102 ^m; n = 10)(Robertson, 1983a). Two swimming veligers, admittedK abnormal, were seen to hatch naturally from egg capsules (Robertson, 1983a). Page 108 THE NAUTILUS, Vol. 107, No. 4 Table I. l.()c-alit\ data, rt-positories of voui'lier.s, ualii {li-[)tli> and protocoiich leiigtlis of Epitonium echinaticosta from Grand Haliaina, .Vbaco, Hiinini and the Berry Is., all northern Bahamas Data are arranged in order of inereasing depth .Abbreviations: .WSP = The Acadenn of Natural Sciences of Philadelphia; CR = Clolin Kedfeni collection; GB = Grand Bahama; n = numbers of specimens measurable (out of many unmpasurablei Localities Voucher s n Depths (m) Lengths (^m) (nos.) Means Ranges ANSP .367 19.5 i 369' ■59 16 1 370 330-430 ANSP 370.521 2 0-2 380 370-380 ANSP .3672.34 & 367235 23 1 410 370-470 CR .3990 1 0-2 4.50 ANSP .32-429.3 2 0-2 450 440-460 ANSP .32556.3 0-2 460 CR 4.598 0-2 510 ANSP .372821 0-2 570 CR .398.3 0-2 590 ANSP .367 195 11 3-10':' 550 470-640 ANSP .370944 & 370.^ i21 9 640 ANSP 369074 & 369'i •59 10 520 CR 76.38 11 650 610-740 ANSP .369476 24 670 ANSP 368803 40 660 {:R 529.5 & 5296 3 52 760 730-790 Running Moti Canal, (JB 26°29'45"N. 78°4r45"\V North Hawksbill Creek, GB 26°32'00"N, 78°45'00"\\' SiKer Coxp Canal. GB 26''.30'30"N, 78°39'30"\V Treasure Cove, Abaco 26°42'00"N, 77°18'20"\V (;hub Cav, Berr\ Is 25°25'N, 77°54'\\" South Bimini 2,5°42'N, 79°17'W Treasure Caw .\baco 26°40'00"N, 77°18'15"W Settlement Point, West End, GB 26°42'15"N, 78°.59'50"W Sandy Point, .Abaco 26°0()'15"\, 77''24'00"W Dead Man's Reef Beach, GB 26°34'45"N, 78°51'45"\V Finders Point, GB 26°30'30"N, 78°46'30"W Kings Bay. Dundee Ba\, GB 26°29T5"N, 7.S°43'.30"\V Chub Rocks, Abaco 26°43'55"N, 77°13'05"W Gold Rock, GB 26°.30'00"N, 78°22'0()"W Tamarind, GB 26°30'45"N, 78°36'00"W N, of N.VV. end (Great) Guana Cav, Abaco 26°44'N, 77°09'W "The hatching shell diameter is about 170 fim. i c much larger than the egg, , . The protoconch is about 0.3 to 0.4 nun long [small ones], i.e it is much larger than the hatching veliger shell - proof that the larva grows sub- stantialK while in the plankton (Robertson, iy8.'3a). There might be geographical vari- aticni, but the data from Abaco as well as Grand Bahama suggest that the larger proloconchs may grow in deeper water every vviieri' the species occurs. In the literature, there is already an indication of pro- toconch size variation in another epitoniid. Two .SEM photos of Epitonium dallianum (Verrili & Smith, 1880) protoconchs (Bouchel & Waren, I98(r figs. 1180-1181) show .some intrapopiilational shaix' and size variation. £. (lallianiirn is deep-water and aniphi-.Mlantii- in distri- bution. Doubt has been cast on poecilogony (intraspecific vari- ation in mode of reproduction) occurring in the Gas- tropoda, especially by Hoagiand and Robertson (1988) and Bouchet (1989). Poecilogony usually refers to plank- totrophy versus lecithotrophy . By extension, lack of poe- cilogony has been taken to mean that gastropod proto- conchs are intraspecifically uniform in both form and size. This paper calls in cjuestion size specificity. MATERIALS AND METHODS This study is based on 68 live-collected and empty shells from shallow depths down to 40 m at southern Grand Bahama island (mainly in the canals of Freeport), 3 emp- ty shells from .52 m off northeastern .\baco, Bahamas, 9 maiiiK beachworn shells from shallower depths at the same island, and a tew also from Bimini and the Berry Islanils, all in the northern Bahamas, all but the last two places on the Little Bahama Bank The .52 m station was about .'3240 m horizontalK Irom the nearest shore. R, Robertson, 1994 Page 109 Figures 1-2. SEM micrographs ot prolnKHK li and part nl tflfindiich (if: 1. Epitoniiini cchiiuilicDsta Irum less than 2 in (Ruiiiiiiig Mini (laiial, Freeport, Grand Bahama), with ca 3,5 prntntonch whorls. Protoconch length = 410 ^in. Teleoconch whorls sutured. 2. £. cchinaticosta from .52 m (off Abaeo), with 5 protoconch whorls. Protoconch length = 770 /jm Note also the incised axial grow th lines on both shells, an indication of planktotroph\ Teleoconch coiling of 2 was ojjen. increasing the measured protoconch length onK slight!). Scale bar = 100 m"! for both micrograplis. Protocoiielis w ere studied w ith light and scanning elec- tron microscopy. Measurements were made to the near- est 10 fim w ith the aid of a cahbrated ocular micrometer in a Wild dissecting microscope at x,50. The anterior end of the larval shell is obscured to a var> ing degree b\ the first teleoconch whorl, which may or may not have disjunct coiling begining before, at, or after the protoconch vari.x. The length measuremenL were made on the exposed part of the protoconch, i.e. as far down as the protoconch varix (smaller than the first teleoconch rib) can be seen. Where the teleoconch becomes disjunct affects the length data slightly, but does not account for the differences in protoconch size re- ported here. Protoconch Is and first protoconch whorl diameters were too corroded to be measurable accurateK (but a size estimate is given below under Results! The originalK '3 deep-water shells from Abaco are in the private collection of Mr. Colin I^edfern (one was lost). Nine other \()ucher specimens from .\baco also belong to Redfern. Two other lots, from Bimini and the Berry Islands, are at The Academy of Natural Sciences of Phil- adelphia (ANSP). The rest, all from Grand Bahama and the Jack N. Worsfold Collection, are also at ANSP. Data on each population studied are gi\en in Table 1. No temperature or rearing data are available. The data are graphed on figure 4. RESULTS First whorl diameters ol both the small and large pro- toconchs are 100-140 /jm (the\ approximate the egg sizes), and whorl numbers are 3.5 to 5. Whole protoconch \\ idths of short and long larval shells are all 270-310 fiin. Long and short protoconchs are similarly shaped (figures 1-2). All specimens have the usual planktotrophic Epi- toniu/n protoconch microsculpture (incised axial lines). Protoconch lengths var\ remarkabK ; from 330 to 990 ^m. The smaller protoconchs have 3.5 to 4 whorls (figure 1) and the larger ones about 5 whorls (figure 2). Thus all are multispiral, another indication of planktotrophy. On the larger protoconchs, whorl ruimbers have increased, not whorl sizes. The data from the two Freeport canal populations show that all protoconchs are very short but appear to be slightly different from each other in size, with pop- ulations from Running Mon having protoconchs no Ion- Page 110 THE NAUTILUS, Vol. 107, No. 4 19^ 77< 27< 26< NORTH ATLANTIC OCEAN LITTLE BAHAMA BANK 50 km Settlement Point West End Dead Man's Reef Beach North HawksbiU Creek ^ ^ "V 17 C to < Q M « o ►J a. A \ ■ South Bimini GREAT BAHAMA BANK (NORTHERN END 27 = 26 = Figure 3. Map ol llic iioitlicr ii li;iliaiiuis, sliouiiit; llic Little Ikiliani,! Bank. |)art (il the (in-at Baliaiiia Hank ami lli<' Idealities in Talile I ger than 430 ^ni and tlidSf ftoni Silver Cove no longer than 470 juin. The population sampled from Dead Man s Reef Reach is puzzling because it ranges in protoconch length from 470 to 640 ixn\, o\('rla|)[)ing the largest shallow water sliell Ironi elsew here and the smallest "deep" water shell. \ possible explanation for the Reef anoinaK is that the other two shallow localities are in sheltered canals, and the Dead Man's Reel is on the exposed south coast — where even near the surface, temperatures may be lower, exposure to wave action max l)e greater, and hosts may be more sparsely disper.sed. The beach shells may have washed in from an estimated de|)th of 3-10 rn bexond the near-beach reel Judging by the Chub Rock and i'inders Point samples, enlargement can occur in water as shallow as 9 and 1 1 m (Table 1). The Sandy Point shell has a moderatelx large [irotoconch (.590 /jni long) which can be explained by the presence of very deep w ater nearbx Un the North- east Providence Channel, figure 3), where greater ex- posure, colder shallow water and perhaps sparsely dis- persed hosts can be expected. The smallest .Abaco protoconch (430 /im) is larger tliaii the smallest Crand Bahama protoconch (330 ^m). The Freeport canals have esf)eciall\ dense populations ot Bunoclcopsis (the anemone host). DISCUSSION Protoconch size bimodalitv in the .Vrchitectonicidae is |)r()babl\ genetic in origin (Robertson, 1970), Since the protoiiinch enlargement (il Rahamian V.pitonium echin- R. Robertson, 1994 Page 1 1 1 2 4 3 4 • • • ' 7 2 6 3 3 2 • •••••• 800 Epitonium echinaticosta PROTOCONCH LENGTH (jim) Figure i. Epiliiniiim fcliiiialUDsln |iriil(KijrKli It-nullis vtrsus water (lepllis. Data troni Tahle 1. The spoli with iiuinbers .sliuvv wliere two nr iimir data pniiits eciiiKule olicosta at "depth occurs in two areas more than 100 km apart (figure 3), an ecological rather than a genetic explanation seems necessary. The factors most likely to account for the size \ ariation of lar\al Epitoniinn cchinaticosla are: 1 Duration of the pelagic lar\a! stage. Mo\ement to and fro by titlal and ocean currents. 2. Diurnal \ertical migration of larvae such as re- ported in Mediterranean Epitonium spp. by Rich- ter (1973, as Scalaria), who also recorded vertical distances traversed i)\ other larval taxa in the order of 100 m 3. Fast lar\ al grow th in the w arm w aters of the canals, slower grov\ th near the cooler surface elsewhere, and slowest growth at "depth" in the coolest water. Scheltema (1967) and Pechenik (1984) showed ex- perimentally that the larval growth rate in ////- anassa obsoleta (Say, 1822) and Crepidula jurni- cata (Linnaeus, 1758), two other prosobranchs. responds to temperature in this manner. 4. In the absence ot metamori^hic cues, a long h\- pothesized delay period and continuing grow th af- ter the onset of competence. According to Pechenik (1986), the lar\al prosobranch Bittiuin allcinatiint (Say, 1822) can grow from 2.8-3.0 whorls to 3.7- 3.8 whorls during the delay periotl. The minimum and maximum larval shell lengths at metamorpho- sis for Crepidula jurnicata are 700 ^m and 2300 ;um (Pechenik, 1984). The great abundance of Bunodeopsis glohulijcra, the sea anemone host of Epitonium echinaticosta, in the shallow, warm Freeport canals (the artificial equivalents of mangrove creeks) must be mentioned. Nearly all of the live-collectetl epitoniin7is are from the canals There there is a large reproducti\e population of the Epitonium and it can be hypothesized that many ot the lar\ ae grow last (in the warm water), swim near their parents, being moved back and forth by tidal currents, and settle and metamorphose promptK upon becoming competent (at small size, 3.5 whorls). Outside the canals, the abundance ot Buncodeopsis is not known. It is even possible that £. echinaticosta has one or more otlier coelenterate hosts at "depth." Some ot the larvae in the sea could have come out of the canals but it is also probable that many were spawned outside. These larvae (from both sources) can be expected to drift into "deep" water and to undergo diurnal vertical mi- gration, v\ hich would involve intermittent slow growth at "depth (cool water), lateral movement in currents, and difficulty in finding host(s). The duration of the com- petent period is believed long, and large (5-whorled) larxae result. Thus the duration of the pelagic larval stage is here considered the most likeK factor bringing about the pro- Page 112 THE NAUTILUS, Vol. 107, No. 4 toconch size variation of E. echinaticosta. But factors 2- -4 listed above must also be iinoKod. CONCLUSIONS How widespread is this kind ot variation among other mollusks? Partly allopatric and partK heterochronous size \ariation in the "protoconth' ot the thecosome pteropod Clio pyraniidata Linnaeus, 1767 has been re- ported in central and eastern North Atlantic Recent plankton and Late Pleistocene bottom deposits off north- west .Africa by Diester-Haass and S. van der Spoel (1978). They attributed the variation to genetic influences, tem- perature differences, and "mi.xing of populations." Size differences are smaller than those recorded here and seem not to be comparable. Long metamorphic dela> periods, like those reported above in Bittiurn and C'repidula, seem a likely cause for the phenomenon to be widespread (little being known about prosobranch delay periods). The prodissoconch sizes of some bivalves apparently are controlled by temperature (Lutz & Jablonski, 1978). As they pointed out, there could be paleoclimatological (or paleoecological) implications. This seems to be the best documented case of extreme intraspecific protoconch size variation in one plankto- trophic gastropod species, the variation probably induced by differing times in the plankton plus a long delay period (plus other factors). The variation is expressed batlnnietrically. There is no indication that more than one Epitonium species is involved, or that there is poe- cilogony. Ecophenotypic variation may be more frequent in marine mollusks than currentK realized. Systematists should be alert to the phenomenon treated here and should beware of mere protoconch size differences for distinguishing planktotrophic species. ACKNOWLEDGMENTS I thank Colin Redfern, of Boca Raton, Florida, lor mak- ing the original discovery antl for loaning me all the s[)ecimens from Abaco. I also acknow ledge extensive use of Jack N. Worsfold's Grand Bahama collection (now at ANSP). The detailed data in these two collections made this study possible. Dr. K. Elaine Hoagland and partic- ularK Dr Philippe Bouchet made penetrating and con- structive criticisms Dr (Jarv Rosenberg and Helen DuSliane also made liclplnl suggestions. LITERATURE CITED Bouchet, I' 1989 A review of poeciiogony in lii'strcipods. Journal ol .Vlolluscan Studies 55:67-78. Hdurhcl. I' .ukI \ W .in'-n l9S(i l^j'V isjoiMil lilt- iiDrthea.st .VduiUic ballival and ai)v.ssai .Vciididae, F.uiiniidae, Epi- toniidae (Moiiusca, Gastropoda) IJoiiettino Malacoiogico Sn|>|)lement(i 2:299-.57(i (blench, W J and R I) Turner 1951 The licnns I'.pitoniinu ni ihe western ,\tlantic. jDlinsonia 2(:3()):2-49-2SS. Dull W H ISWJ Heporls on (Iredijiiij; in the Cull ot \lc\ii(i ,iihI in llic (iaribheun Sea. . .bv the. . ."Ulake . .Heporl on the Moiiusca I'art II. -Gas- tropoda and .Scaphopoda Bulletin ol the \hi.seum of Com- parative Zooiogv at Harvard College 181— 192, 40 pis. De Bourv, E. 19n. Diagnoses de Scalariidae nouveaii.\ ap- partenant aux sous-genres Cijchncala et Sodiscald. Bul- letin du Museum National d'Histoire Natnrelle[Paris] 17(51: 329-331. Diester-Haass, L. and S. van der Spoel. 1978. Lati- Pleistocene pteropod-rich sediment layer in the northeast .Atlantic and protoconch variation of Cliu pyraniidata Liniie 1767. Pa- laeogeographv, Palaeoclimatologv , Palaeoecologv 2-1:85- 109. DuSliane. H 1990 liauaiian Epiloniidaf Hawaiian Shell News Suppiement 1:1-17. Hoagland. K, E, and R. Robertson. 1988. .An a.ssessnieiil ol poecilogonv in marine invertebrates: phenomenon or fan- tasy':' l^iological Bulletin 17-1:109-125. Killiiini, R, N. 1985. The familv Epitoniidae (Moiiusca: Gas- tropoda) in southern .Africa and Mozambique .Annals of the Natal Museum 27(1): 239-337. Leal, J H. 1991. Marine prosobranch gastropods I rom oceanic islands off Brazil. . . . Universal fiook Services Dr W liackhuys, Oegstgeest, Netherlands. \ + 419 pp. laitz, R .A. and D. Jablonski 1978, Larval bivalve shell mor- phometry: a new paleoclimatic tool':' Science 202: 51-53. Pecheiuk, J. A. 1984. The relationship between temperature, growth rate, and duration ol planktonic life for larvae of the gastropod Crepidula fornicata (L.). Journal of Exper- imental Marine l5iologv and Ecology 74:241-257. l^eclienik, J ,A, 1986. Field evidence for delaved melamor- pliosis of larval gastropods: Crepidula plana Say, C Jor- nicata (L.), and Bittiuni ahcrnaium (Sav ). Journal of Ex- perimental Marine Biology and Ecology 97:313-319. Rex, M. .A. and K, J. Boss. 1976. Open coiliiii; in Recent gastropods. Malacologia 15:289-297, Richter, G. 1973. Field and laboratory observations on the diurnal vertical migration of marine gastropod larvae. Netherlands Journal of Sea Research 7:126-134, pi 1. fioberLson. R. 1970. Svstemalics ot Indo-Pacific Philippia (Psilaxis\ architectonicid gastropods with eggs and voung in the umbilicus. Pacific Science 24:66-83. Robertson, IS, 198.3a, (.)bser\ations on the lite liistorv ot the wentletrap Epil(niiuni (.■cliindtiiosliini in the Bahamas. The Nautilus 97:98-103, lUibertson, R, 198.3b, .Axial shell rib counts as svstrnuitic char- acters m E))ili)nium. The Nautilus 97:1 16-1 IS Scliellema, l\. S, 1967, The relationship of temperature to the larval devi-lopmcnl ol Sassarius ohsolctus (.(Gastropo- da), Biological Bulletin 1.32:253-265. W'oodring. \V . I' 1928. Mi N. Konaminio l)<|i,irlinrnl nl i5iiiliii;\ (Central Miiliigaii Liii\fiMt\ ML I'lt-asant, Ml 488.59 I SA ABSTRACT Dreissrnil pnhiiniirpha. ai. i liiiiaird In hkiiii U'liipcialiirc (21 ± 1°(]) l(ir 51 (.la\s 111 artilitial [Kiiiclualer, f\liil)ils liightT hfni(il\ iiipli siHlium, chloride, potassiiiin and total osniolarit\ than mussels acclimated to 4-5°( : (cold), while hemoUmpli calcium was increased with cold treatment. The room tem- perature acclimated nuissel hemoKmph ion concentrations, maintained lor 51 da\s m artiticial ponduater. were similar to lliiisc meavuretl irom mussels sampled m the field, (xild-accli- mation cau.scd Lampsilifi radiata hemoK mph sodium, chloride and nsniolarity values to decrease, while calcium and the "oth- er ions, presumably bicarbonate, increased. The positive net sodunn and chloride flux and the calcium loss of zebra mu^vcK acclimated to room temperature w as reduced upon direct trans- ler into cold pondw ater Sodium and chloride net uptakes were increased in cold-acclimated zebra mussels with an acute in- crea.se in pontlwater temperature w hile calcium and potassium net lo.ss increased. The effects of cold storage on zebra mussels needs to be recognized due to possible changes in the ph\ si( ilog\ of the mussel with this treatment. Kcij norda: Oild-acclimation. ion llu\, henioKiiipli kui cdii- centrations. sodium, chloritle. cakium. pdtassiiiin INTRODUCTION The receiilK introduced zebra nuissel, Dreissena poly- niurpha. has rapidly colonized the Great Lakes Region (Ontario Ministry of Natural Resources, 1992). Zebra mussels torni large colonies of a higii density, broadcast spawn, and have a vehger dispersal stage that remains in the water column for several weeks (Hebert, et al.. 1991; Mackie. 1991; McMahon, 1991; Carton & Haag, 1993). Aside from impacting industries and utilities by louling or clogging water intakes and associated piping (Mackie, 1991; koCalak et ai. 1993; LePage, 1993), zebra mussels also impact the ecology of their newly e.xploited regions by decreasing the density and diversity of fresh- water bivalves of tiie family L'nionidae (Hebert et al. 1991; Hunter & Bailey, 1992; Mackie 1993) and by de- creasing phytoplankton availabilit\ (Leach, 1993). .■\ninials in a freshwater habitat must regulate blood/ hemolymph and cell ion concentrations above the en- vironmental concentrations. Freshwater bivalves, while ionic hyperregulators for hemolymph and cell ionic com- ponents, have reduced concentrations of hemolymph and cell ions relative to other freshwater species. This results in a reduced o\erall ionic/osmotic gradient (Murphy & Dietz, 1976; Dietz, 1979; Kirschner, 1991). While he- moly nipli ion concentrations are low, fresh water mussel sodium ami chloride transport rates compare favorably v\ ith those of other freshwater animals (Kirschner, 1983; Scheide & Dietz, 19.Sfr Horohov et al. 1992). The zebra nmssel is similar to other freshwater mussels in such characters as relatively low hemolymph ion concentra- tions (Horohov et al. 1992). Ion fliix \alues for sodium and chloride in Dreissena polyniarpha show a higher ion turnover with the net fluxes similar to those of other fresh water bivalves (Horohov et al. 1992). Current research protocols for long term storage of zebra mussels in the laboratory recommend that zebra mussels that are not used immediatelv be maintained at temperatures below 10°C (Reid et al. 1993). The purpose of this study was to measure the effect of cold storage on ion balance in the zebra mussel and compare that response to a representative of the l'nionidae. The net ion flux of zebra mussels was measured at two pondwater holding temperatures (room temperature and cold, 4°C). The effect of direct (acute) transfer from one tempera- ture extreme to the other was also determined. MATERIALS AND METHODS Freshwater mussels. Dreissena polynwrpha (Pallas, 1771) antl Lampsilis radiata (Barnes, 1ra nuissels maintained at 4°C compared to mussels maintained at room tem- perature, primarily reflecting the decrease in hemo- Ivmph sodium and chloride. The cold treatment did not significantly affect the "other" ions, presumabK indi- cating no effect on bicarbonate. Mussels acclimated 51 days at room teiuperature maintained hemoK mph ion concentrations similar to the hemolymph concentrations of mussels sampled in the field (Table 1). Thus, Drcisscna poltjmorpha can main- tain ion balance during room temperature acclimation in artificial pondwater with \alues similar to animals found in the natural envirt)nment This occurs in the absence of feeding. To in\estigate whether the cold induced change in liemoK mph ion concentrations was a specific character- istic of Drcissena fwlyniorpha or common to other fresh- water bivalves, the effect of a 10 da\, cold acclimation on Lampsilis radiata was determined (Figure 1). Most of the trends observed with the zebra mussel were also observed in Lampsilis. Hemolymph sodium in L. radiaia was reduced 60%, while the chloride concentration was reduced 75%. Hemolymph potassium remained similar between the two treatment groups. Hemolymph calcium was elev ated 56% in /.. radiata at the lower temperature. Hemol\mi)h osmolarity was decrea.scd 2H'"i with the cold treatment The remaining hemolymph ionic components ("other"), pri'sutnably bicarbonate, were elevated ap- proximately 2 fold with cold treatment. These data suggest that at least some freshwater bi- valves mav reduce their osmoregulatory "set point in J. I. Scheide and P. N. Bonaminio, 1994 Page 115 t-old It'iiipcratures as coiiipart'il to niiisscls in wariiuT teiiipt'iatiirt's (laboratorN tf mpt-ratures). Tu test tlic cl- teet (if an aeute temperature change on ion transpurt capabilities, zebra mussels were acclimated to either 2 1 ± 1°C water temperature or 4°C pondwater temperature for at least 10 days then tested at the other temperature. Mussels acclimateil to room temperatures \\ ere observed to have a significant decrease in the sodium aiul chloride net Duxes, in magnitude and sign, when mo\ed directK to the cold water (P<0.05. n= 14 imissels, room anil cold temperatures except for the sodium flux room tem- perature nnissels n=13. Figure 2). The negative sodium flu.x indicates a dominant efflux in the cokl temperature testing condition whereas at room temperature the pos- itive net sodium flux indicates a tlominant influx. The negative sodium net flux was signiticantK dilfcrent from zero (P<0.05, n= 14 mussels). The change in the net chloride flux was similar to that observed for the sinliuni net flux (P<().05, n= 14 mussels, for room and cold temperatures) Interestingly, the tlirect transfer of tfic room temperature mussels into the 4°(: pondwater re- sulted in a decrease in the magnitude of the calcium loss (P<0.05. n= 14, room and cold temperatures) indicating a decrease in the calcium efflux. The potassium loss, while being approximateK one order of magnitude lower than the other ions, w as significantK less in mussels transferred to the cold (P<0.05, n= 14, room antl cold temperatures). The comparison between zebra mussel net fluxes ac- climatetl to cold pondwater antl mussels acutely trans- ferred to room temperature pondwater was similar to those acclimated to room temperature (Figure '3). Mus- sels, acclimated to the cold bathing media, had signiB- cantK lower sodium and chloritle net fluxes than those tlirectly transferred to room temperature (P<0.05 for sodium and chloride, n=16 mussels for both temperature groups). The sodium and chloride net Dux in the cold- acclimated mussels, determined in the cold condition, were not significantK different from zero(P>0.05, n=16). Calcium and jjotassium losses were increased in mag- nitude with the transfer of cold-acclimated zebra nnissels to room temperature (P<0.01, n=16 each). DISCUSSION Zebra mussels in artificial pondwater maintained in lab- oratory conditions without being fed had hemoKniph ion values close to hemolymph values of mussels collected from the field. However, mussels maintained in cold pondwater, tor the same time period, lost sodium and chloride with a resulting net reduction in hemolyni))li osmolarity. The loss of sodium and chloride from Dreis- scna pulyinorpha and Lampsilis radiata was similar to the effect of salt depletion (Murphy & Dietz, 197(i; Scheide & Dietz, 1982). Vet, the cold acclimated mussels were in a medium that had sufficient ions for mussels to main- tain sodium and chloride balance at a warmer tein|X'r- ature. Sodium and chloride net fluxes usually were positive in room temperature acclimated zebra mussels. How- vww Willi acute Iranslci nl rooiii temperature-accli- mated zebra mussels to cold poiulvvater, mussels lost .so- tliiim and chloriile The loss of sodium and chloride, observed in the cold pondwater treatment, was reflected in the reduced hemolymph sodium and chloride con- centrations observed with cold pondwater acclimation. Metabolic activit\, measureil as o\\gen consumption, of zebra mussels is reduced w ith ctilder water temperatures (McMahon & Alexander, 1991; Quigley, Gardner & Cor- don, 1993). The apparent loss of sodium and chloride observed in the room temperature-acclimated zebra mussels w hen transferred to colli pondwater may be re- lated to reduced metabolic acti\it\ . However, long term acclimation of zebra mussels to cokl pondwater resulted in a reduction of the net soilium and chloride loss to values close to zero. The change in ion loss attribiiteil to zebra mussel cold storage may be the result of a decreased ionic gradient due to the animal loss of sodium and chloride or may be due to changes in epithelial ion trans- piirt and ion permeabilitv , thus the mussels become "tighter to the loss of ions. Cold acclimated mussels, transferred back tu the room temperature pondwater exhibited a positive net Dux for sodium and chloride. Thus, the ion transport mechanisms necessary for reg- ulating sodium anil chloriile balance were present in the cold acclimated zebra nmssels but their activity was re- duceil in colli |ionilwatt'r All net ion fluxes stuilieil varied with temperature re- gime. Zebra nnissels not only exhibited decreased sodium anil chloriile net fluxes, but decreases in magnitude of calciinn and potassium net fluxes. At room temperature, both calcium and potassium net fluxes were negative. The artificial pondwater useil in this investigation, while fulK compatible for maintaining unionid mussels (Dietz, 1985) and zebra mussel hemoK mph for extended periods of time, may not be the best medium for zebra mussels. The calcium loss indicates that this bathing medium is negati\el\' influencing calcium balance. Even with the calcium efflux, liemoK mph calcium values do not change at room temperature, thus calcium is presumably being mobilized from the shell to maintain a relativeh' constant hemolymph calcium value Even with the negative net calcium flux observeil in the zebra nui.s.sel, hemolymph calcium concentrations increased as the hemolymph osmolarity decreased. He- molymph calcium also increased in the unionid, L. ra- diata. during cold ponilwater conditions. The rise in hemolvmph calcium concentrations in cold acclimated- nuissels mav be assisted by the ilecrease in the net cal- cium loss at the lower tem|)erature. The elevation of hemolymph calcium has been noteil with the decrease in hemoKniph soilium in imioniils (Scheide & Dietz, 1982). Only in L. radiata did the "other" ions increase with cold acclimation. Similar changes have been ob- served in the unionid, Ligumia subrostrata (Say) un- ilergoing ihloriile de|iletion (Scheide & Dietz, 1982). C^okl ponilwater storage of mussels causeil an increase in calcium levels concomitant with the decrease of sodium levels suggesting that compensatory mechanisms, al- Page 116 THE NAUTILUS, Vol. 107, No. 4 Room TemoefatLre I I Cold Tempefa'LTe Rocn Temperattre I I Cold TernoeraTure CZ] c. jj Figures I -.'{. ('(iiiipari.sdiis 111 llic t'ftfit'- cil temperature im Ion concentations in Larnpsilis radiata and Dreissena polijmorpha. I. I leiiiiiK iTipli ion (.iiiicentratidns cii L(2nipsilis ratluita alter a 10 (la\ aeeliniatidii period to room temperature (21 ± 1°(^ solid liar) and cold temperature (4 ± 1 °C:, liatehed bar) K'll) refers to the potassium value multiplied l)y 10 for comparison purposes. "Other relers to the difference between the mea- sured total .solute and sum of the measured ion values, as noted in the Materials and Methods section and indicates the he- molymph bicarbonate concentration. The capped lines indicate the standard error of the mean. Sigiiilicanl ilifferences are de- noteil v\ilh a • representing P<0 ().5 with n = (S for room and cold lemperalure-accliinated mussels 2. Kffectsof direct trans- fer of 2I°(: ± \°C acclimated zebra mussels to 4°( : on the sodium, chloride, calcium and potassium net fluxes The po- tassium net flu.x is increased 10 fold lor comparatiM' purposes (K*10). The room temperature acclimated nuissel net flu.x is denoted by the solid bar {n= 14 mussels toi all i^mups except tliougli not completely effective in maintaining room temperature osmolarity, were acli\f in llie cold-aceli- mated mtis.sels. Freshwater animals iia\c set points lor liemoixnipli blood sodimn and chloride (Kirschner. 1991 ). \lo\ement of zebra mussels from the natural en\ironinenl to the laboratory does not appear to negatively effect the mus- sels for at least 51 days. The formulation of artificial pondw ater appears to be suitable for tlie laborator\ niain- tenante of zebra mussels. Zebra mus.sels, held at room temperature, maintained hemolymph ion values close to the collection hemolymph ion values, although the mus- sels were in a negative calcium balance. The mussel holding time in the laboratory may be increased by plac- ing the mussels in cold pondwater and zebra mussel storage at a cold temperature is recommended for lab- oratory housing of zebra mussels not immediately being used tor studies (Held, ei al. 1993). However, the ion concentrations after holding may be depressed, partic- ularly with a longer cold holding period and the lack of feeding. Changes in mussel plu siology, due to cold stor- age, need to be recognized as lurther research is per- formed to understand the basic biology of the zebra mussel. ACKNOWLEDGEMENTS This work was funded by a Research E.xcellence Fund grant #42865 from the State of Michigan to JIS. The cooperation of Consumers Power is greatly appreciated. We thank Thomas Dietz, Stexe Donner, Ron Fobes and Harokl SiKerman tor the critical re\ icw ol this manu- script. LITERATURE CITED Diet/. T II 1979. Uptake of sodium aiKJ chloride In fresh- water mussels. Canadian Journal of Zoology .37:1.56-160. Dietz, T. H. 1985. Ionic regulation in freshwater rimssels: .\ brief review, .\merican Malacological Bulletin 3:233-242 Dietz, T. H. and W. D. Brantoii. 1975. Ionic regulation in the freshwater mussel. Liguinia subroslruta (Sav). Jmirrial ol (;oin|)arati\e PInsiiilogx 104: 19-26. l\liii, II .iiiil I n t'ostliiw 1975 .Vnaerobic sampling of l)iid\ nuids in l)i\al\e molluscs. Comparative Biochemistry and Physiology 52A:265-268. Carton, D W and W IV Haag. 1993 Sea.sonal reproductive iNcles and settlement patterns of Drcissaia palyiuorpliu lor the sodium group, n=13l and the coltl net llux is denoted b\ the hatched bar (n = 14 mu.ssels for all groups). Signihcant flux differences are denoted by • tP<0.05) 3. Effects of direct transler of 4 ± 1°C^ acclimated zebra mussels to 21°C on the sodiimi, chloride, calcium and potassium net fluxes The po- tassiiini net flux is ini'ieased 10 lold for comparative purposes (K- 10). The cold-acelimalcd net flux is denoted b\ the hatcheil bar (11 = 16 mussels) and the room temperature nuis.sel net flux is denoted In the solid bar (n=16 mus.sels). Significant lliix differences arc denoted In • (P<0.05). J. I Scheide and P. N. Bonaminio, 1994 Page 117 in \\ t'stcrn Lake Erif //r Nalcp.i T 1' amlD \\ S( lilocs- ser it'ds. ). Zebra mussels: Biolog). UTii)aets ami nuitKil t:RC: Press, Inc. Boca Raton, FL, p, 1 I 1-12.S Hebert. P. D., C. C. Wilson. M. H Miirdoeh and b i.a/ar 1991. Deniograpli) and ecological impacts nl tlic m\ad- ini; mollusc Drcissvna pdhjiiKiiiilid (:anadiati JoiuTiai ol Zoolot;\ W). 405-409 Horohov. J . H Silverman. J W Lynn and T H. Diet/ 1992 Ion transport in the freshwater zebra mussel, Dreissena pnhjmorpha. Biological Bulletin 1(S3:297— 'jOo Hunter. H. D and J, F. Bailey. 1992. Dreissena ixilynuirphu (Zebra nui.s.sel): Colonization of soft substrata and some effects on I iiionid bixaKes. Tlie Nautilus lOtit-iO-fiT Kirschner, L B I9S.'5, Sodium and chloritle absorption across thelxHb surface: Frog skin and other cpilhrli.i \mcricaii Journal of Physiologv"244:R429-H443 Kirschner, L. B 1991. \Vater and Ions //i; I'rosscr. ( : 1. led.) Comparative .Animal Phvsiologv. l-'ourlh I'dition l"n\i- ronmental and Metabc}|ic Animal l'h\si(il(ig\ \\dc\- Liss.lnc. Neu York. p. 13-107 Kovalak. \V. P.. C, D. Longton and U H Simlhrr 1993 Infestation of power plant uatcr s\ stems b\ the zebra nmssel [Dreissena polymurplia Pallus) In: Nalepa. T F anil U.W Schloesser (eds. ), Zebra uurssels: Biologv . im- pacts and control t"RC- Press. Inc. Boca Raton. FL ji 359- 3S0. Leach. J. H. 1993. Impacts of the zebra uuissel [Dreissena pohjmorpha) on water quality and fish spawning reefs in western Lake Erie. In: Nalepa. T F and D W Schloesser teds.). Zebra mussels: Biologv. unpads and control. CRC Press. Inc. Boca Raton. FL. p. 381-397 LePage. W, L 1993. The impact of Dreissena pulyniurpha on waterworks operations at Monroe, Michigan, .A case histor\./;i. Nalepa, T. F. and D.W Schloesser (eds.) Zebra mu.ssels Biolog\ , impacts anti i-oritrol CIU: Press. Inc. Boca Raton. FL. p. 333-358. Mackie. C; L, 1991. Biologv' of the exotic zel)ra unis.sel. nrci.s- ,sf/ia pohjmorpha. in relation to native bivaKes and its potential impact in Lake St Clair Hvdrobiologit a 219 251-268. Mackie. (.. 1. 199:) Biolog\ ol the zebra uuissel [Dreissena polilinorplia) anil observations ot uuissel colonization on unionid bixalves in Lake St Clair of tlie Great Lakes In: Nalepa. T I" ami l.).\\. Schloesser (eds.). Zebra nuissels: Biologv. iiupactsand i-onlrol (4U,' Press. Inc. Boca Raton, FL. p. 153- Kio McMahon, R. F. 1991 Mollusca: Bivalvia. /;;. Thorp. J H. and .\. P. Covich (eds.) Ecology and classification of North American freshwater invertebrates .\cademic Press, Inc., NY. p. 315-399. McMahou. H F and J F Alexander 1991 Respirator) re- sponses to temperature, h\po\ia and temperature accli- mation in the zebra mussel, Dreissena pohimorpha i Pall I. .\mericaii Zoologist 3174.^ Murpln. W A and 4' II Dietz 1976 The effects of salt depletion on blood ami tissue ion concentrations in the freshwater mussel. I.iiiuinia subrostrata (Sa\ ). Journal of Comparative Plusiologv 108:233-242. Ontario Ministrv ol Natural Resources. 1992. Zebra mus.sel distribution Mmistrv ol Natural Resources. Ontario. Can- ada, December. 1992 Quigley. M. A., W. S. (;ardner and W. M. Cordon 1993. Metabolism of the zebra mussel [Dreissena pohjmorpha) in Lake St. (Mair oi the (Jreat Lakes, h\: .Nalepa. T F and D. W. Schloes.ser (eils. ). Zebra mussels: Biologv. impacts ami control. C;RC: Pres.s, Inc, Boca Raton, FL, p. 295-306. Reid. L^ F . J Biduell, J Carlton, L. Johnson, E, Mar.sden and S.J Nichols 1993. Zebra mussel containment protocols. NO.-\A Sea (Jraiit Office and L'S Eu\ ironmental Protection .Agencv . 20 pages. Scheide. J I and T II. Dietz. 1982. I4ie effects ot indepen- dent S(idium and chloride depletion on ion balance in freshwater mus.sels. (-an.iduin Journal of Zoologv 60 1676- 1682 Scheide. J I ami 4" 11 Dictz 19S6. Serotonin regulation of gill c.WIP proiliictioii. ,Na, and uater uptake in freshwater mussels Journal ol l-Aperimental Zoology 240:309-314. THE NAUTILUS 107(4):118-123, 1994 Page 118 The Muricid Types of Frank Collins Baker Emily II. Yokes (^f(il(ii;\ l)c|);ir(iiicnt riiluiif I iiiversit) New Orleans, LA 70118-5698, USA ABSTRACT in llif lirsl leii scars ol lii.s scit'iililic caret-r V X', Baker iiaiiu-d nine ta\a in the faiiiil) Muricklae, Of these only one, Haus- tcllutn ruhidus (named as Murcx mcs.soriiis var. ruhidum). is here eonsidered \ahd \ll others are synonyms ol older names. Key uardf.: Miirieidae, (Gastropoda, VX'.. Haker INTRODUCTION Tlie name Frank C Baker is very familiar to workers among land and fresli-water mollusks, i)iit for tfie first ten years of his professional life lie thought ot himself as an expert in marine mollusks, especialK the Muricidae, Of his first 21 papers, published between 1888 and 1897, only two (1895b, 1897b) are on non-marine Mollusca, His first publication (1888) was a note describing the marine mollusks collected along the coast of Massachus- sets. In the "(General Notes" of The Nautilus (3:46), it was staled that "Mr. F (L Baker, formerly of Providence, R.I., is pursuing his studies at the Academy of Natural Sciences ol I'hiladelpliia. He spent the months of January to April, 1889, in Brevard County, Florida, and from this experience came a diverse set of notes on the marine mollusks (1889a), on the feeding habits of birds (1889b), including the identification of a number of mollusks eat- en by said birds, and on the habits of the muskrat (1889c). The first molluseaii species he described was Ocinehra jcnksii (Baker, I889d) and in 1890 when the American Association ol (^oncliologists was initially annoimced (77k' Nautilus, 3:140-143), among the charter members listed was "F.C. Baker. Academy Natural Sciences. Philadel- phia. Muricidae ." His second taxonomic paper (Baker, lS9()a) was also on a muricid, with the redescriptidu ol I'rosalpinx perrugatus ((Conrad, 184(j) Among his most important contributions to the IkkK of molluscan know ledge are his series ol seminal papers on the i)rotoii)nchs of tiic Muricidae (Baker, 18901), IS9ld, 1894) and other marine gastropods (Baker, 1897c), to- gether with several catalogues of the species ol Muricidae (Baker, 1891e, 1895c, 1897a). In 1894 he moved to (Chicago to beconu' the lirst v\\- rator ol mollusks at the newly constructed Academv of Sciences (see Baker, 1895a), a position he held for the next 20 years (Franzen, 1958:30), and it is only then that his interests turned to non-marine mollusks, beginning with an important paper (1895b) on the transport of land and Iresh-water mollusks by means of birds, fish, etc. (possibly the first report of "ducks w ith dirty feet trans- porting embryonic shells). Although he published three more papers (1895c, 1897a, 1897c) on marine mollusks, alter 1898 lie never again digressed from non-marine lauiias. In the time that he was study ing muricids Baker de- scribetl nine new taxa, only four of which (those in the 1891d work) were figured. For the most part these species have been ignored by subsecjuent workers. Keen (1971: 554) noted that Ricinula rugosoplicata Baker is a syn- onym of Buccinum lugubre C.B. Adams, 1852, and Yokes (1971) atteni])ted to assign the species to generic taxa. w ith varying degrees ol success (see below ). Radw in and D'Attilio (1976) did nothing with the species, e.xcept to list two as "species of uncertain identity" (Intuherculatus and jenksi [sic\ p. 217). Only two taxa named as "va- rieties have ever been the subject of any discussion: Murex haustclluin var. longicaudus and Murex mes- sorius \ar. ruhidum. In a monograph on the Indo-Pacific species of Chi- coreus Iloiiart (1992, fig. 371) refigured the holotype of Murex (Chicoreus) bitidjerculatus. from the collection of the Chicago Academy of Sciences. This, combined w ith an on-going study on the species of Ergalataxinae, to which I susix'cted certain of Bakers species nright belong, caused me to contact Mr. Ron \ asile. Curator of Mollusks at the (Chicago Academy, and through him 1 arranged to borrow the t\ pe material ol these little known taxa. To complete the study 1 also contacted the Academy of Natural Sciences of Philadel])liia. where four aildi- lional t\ pes were located, and the Wagner Free Institute of Science. Philadelphia, w hich supposedly had one ty pe lot that could not be located. Because these taxa are so poorly known, it seems worthw hile to present all ot them in one place, and they are listed below in chronologital order as originally de- .scribed. UnfortuiuiIcK , most \no\r to be synonyms of presiousK described species. l''\cri though this is not a E. H. Yokes, 1994 Page 119 "rt'visionarx work, lietause tlifre are multiple speci- mens of certain ot tlie ta\a, it seems aiK isihie to ciesi^nate lectot\pes. It is unlikely that most ot the taxa will ever he treatetl in an\ strictK moiiot;raphic work, SYSTEMATIC :S Ocinchra jcuLsii Figures 1, 2 Ocinchra jcnksii Baker, l.S89ci:S() '■'I'riisdipinx jctiksii (Baker). \'okes:122. Habilal: I nknown. Lectolype: ANSP 36099 (here desiiinated) (fig. 1). Paralectolypes: CAS 20696' (fig. 2); ?WFIS (ru.t toniul). Discussion: Baker compared O. jcnk.'-iii lo the Cahlor- nian species Occncbra circumtcxta Stearns, 1S72, i). gra- cilliiita Stearns, 1S72, and O. inicliacli Ford, 1888, sug- gesting that he perhaps thought it might also be from California. However, examination of the twe) extant type specimens indicates that it is an elongate form of the xariable common southern .Australian Lcpsivlhi rinosa (Lamarck, 1822) (see fig. 3), In the original description. Baker (1889d:81) noted that there were four specimens "obtained from the Wag- ner Collection at the Wagner Free Institute of Science. A search of the collections of the Institute did not unco\ er any specimens but at the Academ\' of Natural Sciences of Philadelphia, where Baker worked at the tii7ie, there is a specimen (ANSP 36099; height 16.5 mm, diameter 9.5 mm) labeled as "syntvpe" that nearly matches the cited dimensions ("Alt. 17, diam. 10 mm"). This speci- men is designated as the lectotype. There is one para- lectotype (CAS 20696) in the collections of the Chicago Academy of Sciences, which is slightK smaller (height 15.1 mm, diameter 8.3 mm). The fate of the two addi- tional paralectot\ pes is not known. Murcx haiistclluin Linne var. lungicaudit^ Figure 4 Murcx htiuslcUum Linne var. longicaudus Baker, IS9li):5(i. Ilau.slvlluni longicaudm (Baker), \'okes, 1971:6(1 Not Hau.stellum Umgicaudnni [.sic] (Baker). Kaiclier, 1973, 2 1 21 (= H. kurodai). Not Hatistcllum longicaudus (Baker). Fair, I97():55, fig. 50 ( = // kiirod(ii) llaiislclliiiu haustclluni iHiiislclliini (ijnrie) PDiuier and \'llkt•^. 19,SS,S(i. tigs 4W:. 4(SD ('■h(ili.t\p<-"(lccl(.t\|)cjnl // !,„i- gicdtidlist Hauslelluiii huiislclluni longicaiidiis iBakcri llcmart. 1990; 332, figs. 12, 13, 29. ' Bcith till' Chicago At-adeni\ ot Sciences and the Caliturnia At-adenu ot Sciences use the abl)re\ iation "CAS lor specinu-n Itifiitilieation. In every case in this paper tiie reterenee (]AS is til ihi' Cliicagd .\cadeni\ nt Sciences. Habitat: Red Sea Lectolype: ANSP 60965 (here designaled). Paralectolype: CAS 2070! (fig. 4). Discussion: Most authors (e.g., Radwin & D Attilio, 1976: 49; Ponder & Vokes, 1988:86) have considered this "va- riety" to be a synonym of typical Murex hausteUiim. Howe\er, Houart (1990:332) has expressed the opinion that the differeuct's in the nature of the protocorich be- tween Haustclluni Iwustcllurn s,s. (see Ponder & Vokes, 1988, fig. 78D; Houart, 1990, fig. 31) and forms from the Indian Ocean named longicaudus (see Houart, 1990, fig. 29) and Haustcllum sp. (Houart, 1990, fig. 32) in- dicate three distinct species. Given that all three have protoconchs of 2 to 2.5 whorls and the major difference is in the whorl shape, ranging from rather conical in the t\ pical Indo-West Pacific form, to cylindrical in the East .African form {Haustclluni sp), to somewhat bulbous in the Red Sea form {longicaudus), his argument is not especially compelling. The differences in teleoconch morphology are less be- tween these three forms than between the typical //. Iiaustclltini s.s. and populations named as //. kurodai (Shikama, 1964) and H. vicdani Kosuge, 1980 (see Pon- der & Vokes, 19S8, figs. 87A-C;: kurodai. vicda7ii, and haustclluni s.s., respectively), thus it is probable that the Indian (Jcean/Red Sea populations represent only mor- phological variation as v\ell. In the original description Baker (1891b:56) stated that he had three specimens of this variety, one of which was in tlie collections of the Academv of Natural Sciences of Philadelphia and "two are in my collection." The spec- mien in the collections of the Philadelphia Academy (ANSP 60965) is here designated as the lectotype and the one (fig. 4) in the Chicago Academ\ of Sciences (CAS 20701 ) is a paralectolype. I do not know w here the third specimen is today, Ocinchra pilshryana Figures 5-7 Ocinchra rdshniana Baker, 1S911).57. ?Ur()salpinx piUliryaua (Baker), Wikes, 1971:125, Habitat: ('e\lon Lecioiype: ANSP 60307 (here designated) (fig. 5). Paraleclotypcs: CAS 20699A and 20699B (figs. 6, 7). Discussion: Raker stated that this species "has features recalling the genera Troplion, Vrosalpinx. and Sistrum" (1891b:58) and he is not the only person who has had difficult)' placing it. Examination of the type specimens (figs. 5-7) shows it to be a subjective synonvm of Er- galatax contracta (Reeve, 1846). which also has been attributetl to several different genera b\ other authors: Buccinuni contractuui Reeve, 1846; Murcx calcareus Dunker, I860; L'rosaliiinx innofahilis E.A. Smith, 1879; Pcntadactylus cctjlonicuui Preston, 1909; and Morula martcnsii Dall, 1923. Page 120 THE NAUTILUS, \ol. 107. No. 4 13b '^M Fifjurch 1,2. (hinclmi jcnk.sii Baker I. ANSP .■>W)99 (lectotxpe); height 16.5 mm. localit) unknown (X 3). 2. (^\S 20696 (para- InldUpe); height 1.5.1 mm. locahty mikiiowii (X S). 3. LcpsicUa vinoaa (Lamarck), TAM C: 167197; height 8.2 mm. Hecherehe Ba\, Tasmania. Australia (X 4). !•. Murcx haiislclliirii \ai longiniudns Baker, C'.\S 20701 (paralectotype); height 49,7 mm. Red Sea (X 2,1. 5-7, Cktiichra pilshnjiina Baker. 5. ANSI' 60:507 (leef(it\ pe). height 15 5 nmi, Cieylon (X 3), 6. CAS 20699 (paraieetot) pe A); lieight 16,0 mm. Ceylon (X .3,). 7. CAS 20699 (paraleel<)tv|)e B); heigiit 14.4 mm, Ceylon (X 3). 8, <^. Ricinula iSislrum) rugusuplicala Baker. 8. A.N'SB 61306 (leelolype); height 1 1 .7 mm. Turtle Ba\ . Lower California (X 3). 9. C;AS 20703 (paraieetot) pe); heiglit 11.0 mm. Turtle Bay. Lower California (.X 3). 10. Miircx (Chirorcus) hituhcrculatu.s Baker, C.\S 20702 (holotype); height 31.7 mm. Australia (X 2). 11. Ocinchra wanliana Baker, C.\S 20698 (holotype); height 12 5 mm, ".Australia" (X 3). 12. Ocincbrina aciculata (Lamarck), Vokes C^oll.; height 10.6 imn, off Bota, Spain (X 3). LJ. Ocinebra rubra Baker, C.'\S 20697 (holotype); height 10.7 mm, locality unknown (.X 3). 14. Orcncbra clalhrata (Dall). Vokes Coll.; height 10.0 nmi, off Hollester Banch, California, 15 m (X 3) 1.1. Murex nicssorius var. rubidum 15aker, I SNM 87S003; height 30 5 mm, Anna Maria Ke\, Florida, 46 m (X 2) 16. 17. Purpura iThalfssa) problcrualica Baker. 16. CAS 2()704A (lectotx pel: height -30 9 mm, jafian (X 21 17. CAS 20704B (para- ieetot) pe): height 22.8 mm, Japan (X 2) E. H. Yokes, 1994 Page 121 Often eited as a another s\n(in\ni is Ergalalax rcciir- rens Ireclale. 1931, named astNpeot tlie genus (ami hence contracta is usualK cited as the t>pe species of Ergala- tax): however, E. recurrens is not the same as £. con- tracta but is a synonym of Miirex pauper Watson, 1883, which is thus the correct name for the type of the genus Ergalatax Iredale, 1931. There was no comment on the nurnl)er of specimens in the original type lot. There is one specimen (fig. 5) in the collections of the Academy of Natural Sciences of Philadelphia (ANSP 60307), which is here designated as the lectot\ pe; two additional specimens (figs. (3, 7) at the Chicago Academy of Sciences (C^.AS 20699) are desig- nated as paraleetot\ pes. RicinuUi (Sisfnnu) rugusuplicata Figures 8, 9 Ruutula iSistriiiu) rugusoplkata liaki-r. l.S91b:58 Habitat: Turtle I5a\, Lower C'alilornia. Lectolype: ANSP 61306 (here designated) (fig. 8). Paraleclolype: CAS 20703 (fig. 9) Uisrussion: Keen (1971:554) included R. rugoaoplicata in the s> nonynn of Buccinutu lugnhrc (IB. Adams, 1852, now placed in the Ocenebrine genus Trachypollia and not Morula {Morunella), as Keen reported. There are two specimens in the type lot, one at the Academy of Natural Sciences of Philadelphia (fig. 8) and one at the Chicago Academy of Sciences (fig. 9). leaker (1891b:58) says "this pretty little shell was found in a collection recently purchased by the Conchological Section of the Academy of Natural Sciences from Mr. Henry Hem- phill"; therefore, the Philadelphia specimen (ANSP 61306) is here designated as the lectotype. Murex (Chicorcus) hituhcrculatus Figure 10 Murex [Chicorcus) Itiluhcrculatu.^ Baker, l.S9Ul:133, |)l. II, fig. 4. Chicorcus hituhcrculatus (Baker). N'okes, 1971:24, Murex hituhcrculatus Baker. Fair, 197(i:26; Houart, 1992 l()(i, fii;. 371 (li(il(il\ pe). Habitat: ,\ustralia. Holotype: CAS 20702 (fig. 10). Discussion: F"air (1976:26) was the first to suggest that M. bitubercuUitus is a juvenile specimen of Chicoreus capucinus (Lamarck, 1822). Examination of the holotype shows that she was correct. .Although the localitv "Aus- tralia at first seems misleading, C. capucinus does occur in the tropical mangrove areas of northern Australia. Ocinchra uardiana Figure 11 Ocinchra uardiana Baker. 1S9U1 134. pi 11, fig. 5. (Jcinchnna uanluma (Baker) NUkcs. 1971 127; Fair, 197(i:S(S, tii; l)."> (,illrr B.iki-r). Habitat: "Australia. Holotype: CIAS 20698 (fig. 11 ). Discussion: F]vidently the erroneous locality of "Austra- lia convinced Baker (1891d:134) that this species, al- though "having a superficial resemblance to Ocinebra aciculata Lam., ' was to be distinguished by its larger size, more ileveloped umbilical region, and its \ellow color, differing from the "light rosy Mediterranean spe- cies Ocinehrina aciculata (Lamarck, 1822). True, this gerontic example is larger than most specimens of O. aciculata liut l^adwin and D'Attilio (1976:125) cite 16 mm as maximum height for this species; they alsf) note that the color ranges from orange-brown to red-brown to purple-lirow II. Comparison of the type of O. wardiana to a typical Mediterranean example of O. aciculata (fig. 12) fails to shf)\\ an\ significant differences. Nothing like it has ever been recognized from .Vustralian waters. Ocinchra ndmi Figure 13 Ocinchra ruhra Baker, 1891d:134, pi 11, figs. 6, 7 Ocinehrina ruhra (Baker). Yokes, 1971. 12(x Fair, 197(i:73, fig. 52 (alter Baker). Habitat: Lnknowii Holotype: CAS 20697 (fig. 13), Discussion: ,\lthough compared by Baker to Favartia alvcata (Kiener, 1842) and F. pcasci (Tryon, 1880), the only resemblance is the development of deep pits formed by the intersection of the axial and spiral ribs. Baker (1891d:135) iurther noted "it has some resemblances to the figures of Ocinchra intcrfossa Cpr." and here he is much closer to the truth. In fact, the same form (see fig. 14) was later named Tritonalia intcrfossa var. clathrata by Dall (1919:324; holotype figured In Bormann, 1946, pi. 4, fig. 2; Fair, 1976, fig. 315; Kaicher, 1978, no, 1615). If one considers this a valid species, then Baker's name is the correct one to use. Most w orkers, however, consider the form to be a synonym of Ocenebra atropurpurea Carpenter, 1865 (lectot>pe figured b>' Bormann, 1946, pi. 4, fig. 8; Palmer, 1958, pi, 23, figs, 2, 3, 13; Fair, 1976, fig. 316; Kaicher, 1978, no. 1627), which originally was named as a variety of the common California species O. interfossa Carpenter, 1864 (holotype figured by Bor- mann, 1946, pi. 4, fig. 10; Palmer 1958, pi. 23, fig. 1; Fair, 1976, fig. 318; Kaicher, 1978, no. 1614). Ocenchra rubra /clathrata is no more than a shouldered form of O. atropurpurea, and both are simply clathrate varieties of the O. interfossa complex. Purpura (Thalessa) prohlcmatica Figures 16, 17 Purpura [Thalessa) ]nijhlcnuilu(i \\.iki-c lS91tl 135, pi 11. figs. 2. 3. Page 122 THE NAUTILUS, \'ol. 107. No. 4 Habitat: Japan [Sfta coast, Japan, jiilc I-'ranze'ii, 1958:34, f 11)111 lal)fl with .spciiiJifii.s]. Leclotype: CAS 207()4A (ilcsi^iialcd l)\ l-'raii/fii, 1958: 34) (fig. 17). Paralec-totype: C:AS 2070 IB (fig. 10). Dii^cu^sion: Iriasimicli as /'. pnihlcmulica is uiujiiestioii- ahl\ a .sv'iioiiyin of the common Japant-sc specii'S Purpura rlavigcra Kiister, 1800, one can only assume that Baker was misled by Trvon (1880:163), who clismissed P. cla- vigera as "a depauperate iorni" of P. tumulom Reeve, 1846, the species to which Baker compared prohlema- lica. Baker (I891d:136) further added that "this species is well figured on pi 46, f. 48 of the second volume of Tryon's Manual, which is a reproduction of the shell Lischke (1869, pi. 5, fig. 15) figured as P. tiitnulosa Reeve. The latter is not Reeve's species l)ut the nodulose form of P. clavigera, as Lischke demonstrates in his series of illustrations (1869, pi. 5, figs. 12-l(i). Both morpiiotvpes have been well figured In Kaicher (1980, no. 2481). Reeve's species P. lunuilofia is a synonym of Thais (Tenguella) marginalha (Blaiinilie, 1832), a widespread Indo-Pacific species that does occur in eastern Africa, and Reeve s locality ot "Zanzibar" is probabiv not erroneus, as suggested by Tryon. Members of the P. clavigera com[)lex have been placed in the subgenus Thais (Re- ishia) Kuroila and Halx', 1971, type species: Purpura hrunni Dunker, 1860. There were said to be 12 specimens in the type lot, two ol which were figured in the original description. I-'raiizen (1958:34) selected the larger of tliese (Baker, 1891d, pi. 1 1, fig. 3) as the lectots pe. Both specimens are figured liere (figs. 16, 17). Murex messnrius var riilndmu Figure 15 Manx mcssorins var nilnihiiii Hakcr, IS^^Hil l.iT iinulf iianic list only); 1897a;377 Murcx (Murex) rertirviroslns nihuliis I5akfi , ( ^Iciiili .iiid Perez Farfaiite, 1945:6, pi. .), figs. 1-7 (figs. 1. 2 iectol\pe). Murex (Murex) ruhidus Baker. Yokes, 196;11(),S. |)1, 4. fig.s, .'j, 4. Munx ruhidus Baki'r Nukes, 1971:92; Radwiii ami D'Mlilio, 1976:71, pi. 11, hg, 15; Fair. 1976:7:3, fig. 40. Haustelhun ruhidus (Bakerl \ (ik<'s. 1990:17. pi. 1, tig 12 Murex ruhidus paniiiuicus I'rliKli. 199062, fig. 8. Habitat: ( ledar Ke\s, Florida. Lertotype: Harvard M(;Z 147272 (designated In Clench and Perez Farfante, 1945:8). Discussion: This is the most w idcK recognized ol leaker's taxa, and was only inadvertantly named In Baker, wlm credited the varietal name to ball (1889:196). In that reference Dall describes the Florida specimens of Murex /nessorius as 'often of a deep ro.se-color . . . less hooked and spinose than the discolored specimen figured b\ Reeve [1845, pi. 22, fig. 90, which is a good example of typical rnessoritis]" However, nowhere does Dall men- tion the word "ruhiduni" and the name must date to Baker s 1897 usage, w here he does gi\e a description of sorts ("the shell being of a deep-pink color," 1897a:377), which together with the Florida localitv' is sufficient to identify the species (Jeiich and Pc-rez Fartante (1945:8) recognized the Florida form as a valid subspecies of the Pacific reeur- virostris Broderip, 1833, changing "ruhiduni" to the or- thographically correct ruhidus. and noting that the spe- cies is not onl\ pink but comes in "cream, pink, pale orange, salmon, or even red and some specimens possess darker color bands at the shoulder and base of the body w horl. Althougli generally confined to the Florida and Ba- hamian areas, specimens also occur in the Ba\ Islands of Honduras, the northern coast of South America, and Panama (see Vokes, 1990:17). The latter have been sep- arated as a geographic subspecies by Petuch (1990:62, fig. 8). ACKNOWLEDGMENTS I wish to thank Ron Vasile, Curator of Mollusks at the Cyhicago Acadenn ol Sciences, for making a\ailable the types in their collection. Gar\ Rosenburg, Earle E. Spa- mer, and David S. Wiedner, all ol the Academv of Nat- ural Sciences of Philadelpliia, located and loaned tvpe material from their institution. Eugene .\. 13olt, Jr., of the Wagner Free Institute of Science, Philadelphia, was most helpful in attempting to track down the types of O. jenksii, said to he in their collections. Ian Loch, the Australian Museum, kindK provided the figured speci- men of Lepsiella vinusa, as well as considerable help and hospitalitv during in\ 1988 visit to the .\ustralian Mu- seum. ABBREVIATIONS OF REPOSITORY COLLECTIONS ANSP - Academy of Natural Sciences. Philadelphia, Pennsylvania, USA CAS - Cliicago Acadenu oi Sciences. Chicago. Illinois, L'SA MCZ - Museum ot C^omparative Zoolog) . Harvard Uni- versitN. (-ambridge. Ma.s.sachu.ssets, USA TAM - The Australian Museum, Svclney. New South Wales, Australia USNM - National Museum of Natural History, Smith- sonian Institution, Washington, D.(>., USA \\ I'lS - Wagner Free Institute ol Science, Philadelphia, I'ermsv Kania, I S.\ LITERATURE CITED Baker. F, C, ISSS, |( )ii a icillcrlinu Irip Id ( )iisel Bay. Mass.]. rlir ( :(inilinl(igisl's Exclian.ge 2(9):! 19. li.ikcr, I- C ISSOa, Notes dm Floridaii siiclls. The X.iutilus .■5(5):5;5-54, Baker. F (' IS.SOli Nolrs mi the lood ol hirtls rrorreiliiigs ol .Vi'.ulciiiv ol X.iliiral Sciences of Pliilaclel|iliia 11:266- 270. E. H Yokes, 1994 Page 123 Baker, F. C, 18S9c, Remarks upon tin- rdUiKl-Uiilcil iniiskial, Neofiher alleni. True. Procedings nl Aciilcriu dl \,ihir.il Sciences of Philadelphia 41:271-273 Baker, I"" (; IcS.SQd, De.scriplioii of a iiru -.[H'ck-sdl Ociitchra The Nautilus .3(7):.S0-N1 Haker, P'. C;, 189()a. Reniarksou I rosalpiiix pcniifialiis (.'.mir Proceedings of .-KcademN of Natural Sciences of Philadel- phia 42;4(i-47; reprinted vcrhatuii ni The N.mlilu-- 4l,'5); 29-30. Baker, F. C. f 890b. On the modibcation ol the apex in Munx Proceeding of .Acadeni) of Natural Sciences ot Philadel- phia 42:66-72. Baker, F CI. 1890c. .\ catalogue of conchological alilire\ia- tions The Nautilus 4l8):89-91 Baker, V C 1891a. Notes on a collection ot shells Ironi south- ern Mexico. Proceedings of .\cadeni\ of Natural Sciences of Philadelphia 4,3;45-.5o Baker, F (.'. 18911). 1-leinarks on the Mmicitkie with descrip- tions ol new species of shells. Proceedings of .Acadeiii\ ol Natural Sciences of Philadephia 43:.56-61. Baker, F. C;. 1891c. [continuation of] A catalogue of concho- logical abbreviations. The Nautilus 4(10)T 1.5-1 17. Baker, F (.'. 1891d. Descriptions of new species of Muricidae, with remarks on the apices of certain forms Proceedings ot Rochester .-Vcadem) of Science 1 129-137. i)l 1 I Baker. F. C 1891e. Catalogue and s\iionym\ of the Recent species of the famiK Muricidae. Proceedings of Rochester Vtademy of Science 14.53-172. Baker. V. C. 1892. Notes on a collection ol shells Ironi the Mauritius, with a consideration of the genus Magilus of Montfort. Proceedings of Rochester .Acadenn of Science 249-40, pi 9 Baker, F. V. 1.S94. Further notes on the eml)r\onie whorls of the Muriciilae. Proceedings of the Acadein\ ot Natural Sciences of Philadelphia 46:223-224 Baker, F. C. 1895a. A glance at the Chicago .\cadem\ of Sci- ences. The Nautilus 8tl0):109-lll, Baker, F. C. 1895b. The geographical distribution of the [land and fresh-water] Mollusca. Science (New Series) 2(33): 179- 183. Baker, F. C. 1895c. Prelniiinarx outline of a new classihcation of the famiK Muricidae. Bulletin of the C^hicago Acadenn of Sciences 2(2)169-189. Baker, F. C: 1897a Critical notes on the Muricidae. Transac- tions of the .\cadem\ of Science of St. Louis 7(16):371- .391. Baker, F C:. 1897b. On a collection of mollusks from Crand Tower, Illinois. The Nautilus 1 1(3):28-.30. Baker, F, C. 1897c. On the modification of the apex in gastropod mollusks. .Annals of New York .Academx of Sciences 9:685- 704, pis. 18-20, Bormann, M. 1946. .A survey of some West .American Oce- nebras, with description of a new species. The Nautilus 60(2):.37-43, pi. 4. Clench, W J and I Perez Farfante. 1945 Phe genus Murcx in the western .-Vtlantic. Johnsonia I(17):l-58, pis, 1-29 Dall, \\ H 1889. Reports on the dredging, under the su- pervision of Alexander .\gassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1.879-80), In the IS. Coast Survey Steamer "Blake. Lieut. -C^ommander (^D Sigsbee, L.S.N., and Commander JR. Bartlett, Ll.S.N., commanding X.\I.\ Report on the Mollusca; Part II, Gas- tropoda and Scaphopoda. Bulletin ot Museum of C.om- parative Zoology at Harvard C:ollege 18:1-492. pis. 111- 40. Dall. \\ II 1919 Deveriplions of new species of Mollusca Ironi the north PaciticOce.ui in the collection of the L'uited States National Museum Proceedings of the II. S. National Museum .56(2295) 29.3-37 I Franzen, D, S. 1958 T\ pes ol molhisks described b\ 1" C Baker - Part III. (Chicago .\cadein\ ol Sciences. The \.ui- tilus 72(l):.3()-34 Fair. R. 1976. The Murcx book, an illustrated catalogue of Recent Muricidae (Nhiricinae, Muricopsinae. Ocenebri- nae). Priv.iteK published. Honolulu. Hawaii, 138 pp.. 23 pis. Houart. R, 1990 New l,i\a .irid new recorils ot Indo-Pacilic species ot .\/i/rc.v and lluiistclliuu (CJastropotla. Muricidae, Miiricinae) Bulletin tin Museum .National d'Histoire Na- turelle, Paris (Ser. 4) 12 (.sect. A) (2):329-347. Houart, R 1992. The geiuis Chicoreus and relatetl genera (Gastropoda: Muricidae) in the Indo-West Pacific. Me- moires tlu Museum National d'Histoire Naturelle, Paris, Zool. v.(A) 1.54:1-1.88, Kaicher, S I^ 1973 Card catalogue of world-wide siiells, Pack no 2 - Muricidae. Part I, nos. 99-195. Privately published, St Petersburg, Floritla. Kaiilier, S D 1978 (lard catalogue of workl-wiile shells. Pack no 16 - Muricidae, Part HI, nos. 1540-1646 Pri- \atel\ published, St Petersburg, P'torida Kaicher, S D 1980 tlard catalogue of world-wide shells. Pack no 24 - Thaididae. Part 1. nos. 2392-2497 Privately published. St, Petersliurg. Florida keen. .\ M 1971 Seashells of tropical west America. Marine molhisks from Baja (."alifornia to Peru. Second Edition. Stanford I niversitv Press. Stanford, California. \i\ + 106-1 pp., 22 color pis. Lisclike, t;. F. 1869. Japanische Meeres-Concli\ hen. \olunie 1 Theodor F'ischer, C^assel, 192 pp , 14 pis Palmer, K. \'. W. 1958. Type specimens of marine mollusks described by P.P. Carpenter from the West Coast (San Diego to British Columbia). Geological Society of .America, Memoir 76. 376 pp.. 35 pis., frontispiece. Petucli, E J 1990 .\ new molluscan faunule trom the Ca- ribbean coast ol Panama. The Nautilus 104(2):57-71. Ponder, W. J and E. H. X'okes. 1988. A revision of the Indo- West Pacific fossil and Recent species of Miirex s.s. and Haiislclluiu (Mollusca: Gastropoda:Mnricidae). Recordsof the .Australian Museum. Supplement 81-160 Radwin, G.E and A. D'Attilio 1976. Murex shells of the world; an illustrated guide to the Muricidae. Stanford llni- versity Press, Stanford, California, 284 pp., .32 pis. Reeve, L. \. 1845-1846. Conchologia Iconica, v. 3, Murex, 3« pis. (pis. 1-34 issued Apr to Oct., 1845; pis, .35, 36 issued Apr,, 1846) Tryon, G. W 18.80 Manual ol (>'onchology, structural and s\ stematic, w itli illustrations of the species, v. 2, Muricinae, Purpnrinae Pliilatlelphia, Pennsv Kania, 289 pp., 70 pis. \ likes E H 19(i3. Cenozoic Muricidae of the western .At- lantic region. Part I - Murcx s s Tiilane Studies in Geologv 1(3): 9.3-12.3, pis. 1-4. \ okes, E H 1971, Catalogue of the genus Murex Linne (Mollusca: Clastropoda); Muricinae, Ocenebrinae Bulletins of .American Paleontologv 61(268):l-l 41 \okes. E. H. 1990 Ceuo/oic Muricidae of the western .At- lantic region Part \ 111 - Murcx ss, Hauftlcllum. Chi- coreus, and Ilcxa))lcx, additions and corrections. Tulane Studies in Geologv anti Paleontologv 23(l-3):l-96, pis. 1- 12. ' THE NAUTILUS 107(4);124-128. 1994 Page 124 Invasion of the Rivers of the Littoral Central Region of Venezuela by Thiara granifera and Melanoides tiiberculata (Mollusea: Pros- obranchia: Thiaridae) and the Absenee of Biomphalaria glabrata, Snail Host of Schistosoma mansoni J. p. Pointier l„il)i>rjl(iirr dv Biiiliigif Marine ft Malacoldgie Centre ile Hiologie Tropicale Ec'ole Prali Asistencia Social Maracav, Venezuela ABSTRACT Prior to 1975, some streams of the littoral central region of Venezuela were active transmission sites for intestinal schisto- somiasis. The snail intermediate host, BiomphuUnia glabrata. was controlled but not eliminated b) repeated molluscicide applications. Two thiarid snails, Thiara granifera and Melan- oides tubereulata, began to invade these streams around 1975. The rapid expansion of these two species was confirmed during more recent surveys. Although other factors max be involved, the exi.stence of stable populations ot tlie two thiarid species is probabK linked to the absence of recolonization In B. glabrata. the latter species has not been observed in the study region during the last 15 \ears. Key nords: Schistosomiasis, thiarid, recolonization INTRODl'CTION The littoral central region of N'enezuela iornis the north- ern portion of an endemic area for intestinal schistoso- miasis. Some of the rivers of tins region have previously offeree!, and may still constitute, suitable habitats for the snail intermediate host, Biumphalaria glabrata (Say, 1818); they may thus act as potential transmission sites for tlie parasite (Incani, 1987). \ control program, based cm I hemotherapy, sanitation, health education, environ- mental improvements and api)lications of molluscicides was iiiitiatet! in 194.). 'I'his control program resulted in the near total inti'rruption ot transmission of Sehistosoma mansoni (Sambon, 1907) during tfie 1970's (Incani, 1987). Several malacological surveys carried out in the lower course of the rivers before 1975 had revealed the pres- ence of R. glalnata. More recent surveys, undertaken between 1975 and 1983, and in 1990. demonstrated the . ' \ 10mm Kipurc I. Thiara granijera [\fil; and Melanoides tubereulata Uight). J. p. Pointier et ai., 1994 Page 125 Figure 2. Map of the littoral ct-nlral region ol \ ene/.uela .showing the surveyed rivers invasion of tliese rivers by two species of Thiaridae, Thiara granifera (Lamarck. 1822) and Mclanoidcs tnlicrcnlata (Miiller, 1774)(Figs. 1, 2). Thiara granifera was introduced to tlie Caribbean area around 1935 (Murray & Woopschal, 1965) and to \ enezuela in 1970 (Chrosciechowski, 1973). It was first observed in the littoral central region in 1975. The in- troduction of M. tubcrculata to the Caribbean area prob- ably occurred later. It was reported for the first time in Te.xas in 1964, but according to Murrav (1964), it prob- ably was introduced there several years earlier. M. tub- crculata appeared in Venezuela in 1972 (Chrosciechows- ki, 1973), and its presence in the rivers of the littoral, central region coincided with that of T. granifera. This paper presents the results of malacological surveys car- ried out in all these rivers between 1975 and 1983, and in 1990, METHODS Twent\-nine rivers were sampled along the littoral cen- tral region of Venezuela (Fig. 2). All are permanent, short and rocky streams descending rapidK' from the mountains of the littoral cordillera; the U[)[)er stretches constitute the typical habitat of the local thiarid snail Pachycliilus lacvississimu.s (Sowerby, 1824). The lower courses, often polluted in urbanized areas, provide suit- able habitats for the snail B glahrata. OnK the lower parts of the rivers were investigated during this study. Qualitative data were collected on the presence or apparent absence of the main species of snails before 1975, between 1975 and 1983, and in 1990 during the control program monitored b\ the Ministrv of Health (Chrosciechowski ct al.. 1987; Balzan, 1988). Samplings using a quadrat method were carried out in si.x sites located on the lower courses of the Cata, La Trilla, Ca- muri Grande, .Anare, Los Caracas and El Botuco Rivers in 1990. A\\ snails were manually collected and counted in squares of 0.1 m" regularly spaced at one meter in- tervals along four or five transects crossing the river. RESULTS Prior to 1975, seven of these rivers harbored B. glahrata (Fig. 2 and Table 1) and were also active transmission sites for schistosomiasis. These rivers were treated several times with molluscicides (sodium pentachlorophenate) during the 19(50's (.-Marcon de \o\a ct ai. 1987). In 1975, T. granifera and M. tubcrculata were first discovered in the Naiguata, .Anare and Los Caracas Riv- ers, and in other rivers in subsequent years. General survevs, carried out between 1975 and 1983, showed that these two species hail become dominant in many rivers (Table 1). In 1990, another survey demonstrated the in- Page 126 THE NAUTILUS, Vol. 107, No. 4 Table I. Malai'ological i)b.scT\ati(iii,s made in t«ciit\ rivers of the littoral eeiilral region ol \'eiie/iiela B g Ruitttpluildrid filahrala, I'.g. Tliiara graiiiffia. Ml Mclanoidcs ttihcrciilata. P.I. Parhyeliilufi Incvississimus. Snail launa Rivers < 197,3 1975-198;5 1990 MdlluscieRle application (last \ear) Sanclioii Borlnirata I'atanenio San Esteban l.a Trilla Cat,. (iiiarapitii ( lluironi ( 'liuao El Liinon Cliieliiriviche L'rieao Petaquire Osorio NEiciito San Julian Lria N'aigiiata Caniuri (irande .\iiart' Los (Caracas El Botuco La Seea Osnia Oritapci Tixlasaiia La Saljana Carnao ( :liusp,i B.g. B.g. B.g. B.g. B.g. B.g. B.g. T.g./M.t. (198.3) T.g./M.t. (1980) T.g./M.t. (1977) T.g./M.t. (1977) r.g. (198.3) T.g./M.t. (1976) r.g. (1984) T.g./M.t. (1980) T.g./M.t T.g/M.t T.g/M.t T.g/M.t T.g. (1979) T.g./M.t. (1978) T.g./M.t. (1979) T.g./M.t. (1979) T.g/M.t. (1979) T.g. (1979) T.g. (1979) T.g. (1979) (1975) (1979) (197.5) (1975) T.g. M.t. T.g. T.g./M.t. T.g./M.t. /P.l. M.t. T.g. T.g. T.g. T.g./M.t P.l. T.g/M.t. T.g./M.t. T.g. T.g./M.t. T.g./ M.t. /P.l. T.g./M.t. T.g./M.t. T.g./M.t. T.g. T.g. T.g./M.t. T.g./M.t. 1967 1962 19S4 1962 1963 19(50 vasion of most of the rivers of this region by the two thiarids and confirmed the absence of B. glahrota (Table I). Sampling carried out in 1990 by the ciuadrat method determined the spatial colonization and densities of snails in these six rivers. The two species of Thiaridae mainly colonized the edges or protected areas in five of the six sites (Figs. .3-14). These sites, with one exception, were characterized by a strong current throughout the year and the variable presence of rocks, stones, grt vel and sand. Snail densities were not uniform, but showed an aggregated distribution. In this ecological situation T granifera was often more dense than M. tuhcrcuhita. The observed maximum densities expressed as snails per 0 1 m- were of .50 T. granijcra and 67 M. tnherculata lor ihcCata River; ,31 T. grant j era and 10 M. tuhcrculata lor the La Trilla i^iver; 90 T. granifera and 1.5 M. tuh- erculala for the Camuri (irande River; 23 T. granifera and 5 M. tubereulata for the Anare River; and 46 T. granifera and 9 T. tnherculala for the Los C-aracas River (Figs. 3-14). The sixth site, located on the El Botuco River, was ecologically distinct and characterized by a sandy bottom and very low velocities. Snail densities were low^ T. gran- ifera reached a maximum dcMsil\ of 10 in ^ O a^' 4fly 4^ ^^ 8 4r ■\ # ■;«■ ## Fij»ure> }{-•>. Si-rii". nl spriiiiifiis taken lidin Iklal pdiuls (left) and MiiKU inlertiflal (rii;iit) luihilats in ()li\().s, Huenoi .Aires, Rio de La Plata 8. Citrhicula largillicrli. '). Corhicula flumi- iicd \ll scilc liais = Ml Tnni l-'if;urc> .'>-7. ( ^>rl)itiiiiils liiini Hid ilc La I'lata. .S. Scacor- Inciila hmosa (Matoii, 1809) froni llir aiT(i\o Migiielin 6. Ci>r- hiciila /(n7;i7/uTh' (Pliilippi. ISlli ironi Piinta lara. Kii.scnada. 7. Corhicula fhimincii (Miillcr. 177 1), Ironi Olivos, I?iieiio.s Aires ,\ll scale hars = 10 mm. C. F. Ituarte, 1994 Page 133 'I'ahl.- I. Slirll IfTijitli slic-ll lifiiihl (LH) ami slicll lciii;lli shell ukUIi (L W I rutins lor (,'. larnillurti and (' jhinuma pcipulatioiis licirii ililltTfiil fiivlroiimfnls at ()li\i)s. Win dr I, a I'lata (.v = nu-aii. S = staiidanl iIcn iaticjii ) (.'. ftuiuinca C . Iargilli( ■rli 1. H 1. w n A S X S 101 101 1 1:5 1 20 0 02 oot 1 5h 0 05 1 iiT 0 07 II L/W Sdit Ixittcmis SaiicK intcrtiilal 1 17 I 20 0 04 0 0.) I SS 0 05 0 10 W (figs. 8,9; Table 1 ). Populations that srttleii in substrate depressions along the upper intertidal zone, vvhieh, like tide pools of the rocky marine intertidal areas, constitute small, lentic, micro-habitats with sand bottoms contain- ing substantial amounts of mud or silt, luue a typical trigonal shell outline, inflated beaks, thick shells, ami lower L/W and L/H ratios. The posterior projection ot the rostrum occurs only in Corhicula fliimiriea (Figs. S,9; Table 1). DISCUSSION Parodiz and Hennings (1965) reported that the family Corbiculidae is represented in the southern neotropical region b\ the native genus Neocorhictila Fisher, 1887 In Argentina, Mcocorbicula liinu^a {Maton, 1809) is found along the Llruguay and the Rio de La Plata Rivers, while N. parancnsis (d'Orbigny, 1846) occurs in the Parana River Basin. Although a considerable bod\ of evidence has been presented to warrant the segregation ol South American forms within the genus Ncocurhicula (Parodiz & Hennings, 1965; Dall, 1902; Ituarte, 1984a, b, 1986) many authors persist in referring autochtonous taxa to the genus Corhicula. Conclusions based on this premise, including biogeographic (Counts, 1980; McMahon, 1983) and shell morphometric comparisons (Britton 6; Morton, 1979) are erroneous. Corhicula and Neocorbicula differ substantially in shell morphologv. The wide range of shell \ariability show n by both genera in response to environmental factors re- veals that the most relevant character is the presence in Neocorhicula of a deepK marked pallial sinus (siphon lengths 12-14 mm). In Corhicula the pallial sinus is ab- sent or represented by a slightK curved pallial line, and siphons as well as associated retractor muscles are scarce- ly developed ( siphon lengths 3-4 mm) (figs, 5-7). Branchial incubation of embryos, a characteristic com- mon to both genera, is carried out in different ways. Corhicula incubate embry os in large rnnnbers [ > 10,000, as calculated for C. largillierti (Ituarte, 1984a)] within the water tubes of the tw o inner demibranchs. The basic structure of the inner demibranchs of Corhicula is un- modihetl, and differs from the outer demibranchs only in ha\ing fewer interlamellar junctions. Seocorhicula. on the other hand, develops brood pouches by cellular proliferation of the interlamellar junction epithelium of the inner demibranchs. Each brood pouch contains a single embryo. Gravid indi\iduals usually have 20-30 brood pouches, and occasionalK as many as 45 (Ituarte. 1984b) figs. 2,3). The incubation period of Corhicula is \er\ short, with embryos released at a late veliger or pediveliger stage (200-240 /um diameter) (Ituarte, 1984a; Britton & Morton, 1979). In Ncororin'riWa, the incubation period is longer and the offspring are released as fully developed juveniles Birth size in Mcocorhicula is about 1.1 mm. Howe\er, some indi\iduals from each cohort remain in the maternal gill until the\ reach 4-5 mm in shell length (Ituarte, 1984a) (figs. 2,3). In Corhicula. all embry OS w ithin a gra\ id indi\itlual show a similar degree of ile\ elopment, while in Scocorhicula tw o or three gen- erations of larvae may coexist w ithin each demibranch (Ituarte, 1984b) (fig. 2). Those species of both genera that occur in the Amer- icas are simultaneous hermaphrodites. However, there are differences in gonadal structure. During the early stages of gonadal development of Neocorhicula the pre- dominant elements in the acinar wall are large and vac- uolated follicular cells, w ith germ cells scattered along the base of the epithelimn (Ituarte, 1986). In Corhicula. follicles are composed of minute follicular cells, with gonia the dominant elements in the acinar wall (Ituarte, 1984a). In Corhicula there is no pronounced segregation of male and female germinal cells within the gonad. If any degree of localization exists (Kraemer, 1978), it dif- fers from that shown in Neocorhicula, where the ovary occupies the dorsal region of the gonad, while the testes develop in the ventral half (Ituarte, 1986) (fig. 1). Fur- thermore, in Neocorhicula. testes occupy more than 50'x' of the gonad (Ituarte, 1986) while in Corhicula. they occupy 15-30% (Kraemer & Lot, 1977; Britton & Morton, 1979).' Corhicula largillierti was tiescribed from the Yangtse- Kiang River in China, and its range includes northern and Central China as well as the Korean Peninsula (Pras- had, 1929). The taxonomy of the Asiatic species of Cor- hicula was reviewed by Morton (1979), who reanalysed Prashad's (1929) "affinities ' and regarded C largillierti as belonging to a group of species that are synonymous with C. fluminea (Morton, 1979: Table 11). However, a re-examination of plots of affinities of Chinese, Korean and southeastern Russian species reveals C. largillierti to be distinct from an\ other species (Morton, 1979: Table 6). Moreover, there is no evidence in the text of Prashad (1929) to support the s\iion\m\ of C largillierti with C. fluminea. The material studied and assigned to C. largillierti. Page 134 THE NAUTILUS, Vol. 107, No. 4 which i.s now widoK (listril)utecl in Argentinean waters, show.s morpliological ditierenee.s from C. jluminea that are sufficient to to consider it a distinct species. Corbicula largillierti is characterizetl In its inoch'rateiN thick shell, with fine, clo.seK spaced concentric ridges and \ellowish or greenish surface. Its shell is trigonal, nearly equilateral, and not greatly inflated Its beaks are moderately inflated, the posterior end slightK [irotrucled and obtuse at the c\tremit\, e\en in larger indisiduals (fig. 6) Corbicula fluniinca differs from C. lar^illicrti in having a more solid shell, with coarser, more widely spaced surface sul- cations. Its shell is more inflated and globous, with prom- inent and inflated Ix-aks In C. fluminea. the posterior margin is markedlv' protruded to form a characteristic rostrum, especially in specimens larger than 15 mm (fig. 7). In spite of the liighly variable shell morphology that results from environmental factors, the differentiating features listed above do not become masked, and the two species are easily distinguished even when populations of both species coexist in the same habitat. ACKNOWLEDGMENTS The author wishes to express his gratitude to Dr. Tad- ashige Habe (National Science Museum, Tokyo) for his assistance in the identification of specimens of Corbicula largillierti. Thanks are also due to the following for pro- \ iding specimens: Dr. F. Spinach Ross (Proyecto Eval- uacion de Recursos Pesqueros del Rio Uruguay, CARU- INIDEP-INAPE); Mr. R. Dillon, Mrs. A. Ibargoyen, Dr. J. Garcia, Lie. F. Kaisin, Lie. N. Laiidoni, and Dr. A. Rumi. LITERATURE CITED linttmi, J. C , D. R. Coldiron, L. P. Evans, Jr. C. GolightK, K U. O'Kane, and J. R. Teneyck. 1979. Reevaluatioii of the growth pattern in Corbicula fluminea (Miiller). [in]: Brittoii, J. C. (ed.) Proceedings, First International Cor- bicula Symposium. Texas Cliristiaii L'niversity Research F"oundation, Fort Wortii, Texas, p. 15-3><. Brittoii, J. C. and B. S. Morton. 1979. Corbicula in Nortii America: The evidence revievvwl and evaluated [in]: Brit- ton, J. C. (ed.) Proceedings, First liUeriiational Oirbicuia Symposium. Texas Christian University Researcli Foun- dation, Fort Worth, Texas, p. 2-19-287 Britton, J. C. and B. S. Morton. I9S6 Fob inorpiiism in Cor- bicula fluminea (Bivalvia: Corbiculoiilea) from Ncirtii .Vmerica Maiacologieal Beviev\ 19:l-4'5 Counts, C \. 1980. The •^vwn^ (.'■orbicula Mulilteld (Bivalvia: Corbiciilidae) in .Africa and South America: Zoogeograpli- ic and taxonomic prol)lems. Bulleliii nl llic Xnicncaii \lal- acological Union for 1980:71-72 Dall. W. H. 1902. Noteon.\Vo™r/«ci// publication. Both the pages preceding and following the main tax- onomic .section reflect tho.se presented by the Seashells volume, with appropriate motlifications and uptlated in- formation. The introduction covers various aspects of opisthobranch biology (behavior, feeding, life histories), and the chapter on zoogeography has been expanded. C;hapters on collecting and cleaning methods, covered f ulK in the Seashells \filume, are understandabK absent, in deference to .Australia s recent collecting restrictions as well as opisthobranchs' rare standing in the average shell collection. Nothing is mentioned, however, on How- to Photograph Sea Slugs, which would have been enor- niousK welcome, since the photographs of li\ ing animals (mam in obviousK underwater surroundings) are the highlight of this book. A list of Western Australian shell clubs, a brief glossary (but not nicely illustrated as in the Seashells volume), and taxonomic index conclude the volume. By this book, the authors have sought to "stimulate interest" and "demonstate both what is know n, and also what is not known" about Western -Australian opistho- branchs. They have succeeded on both counts through their recognition of ongoing research and of the unre- solved taxonomic problems in this group. Forty-five spe- cies are identified to genus-level only (or with a tentative species assignment), with some noted as new to science (although, appropriately, none are originally described in this book). Over half are stated as rare or uncommon; many species are recorded from single localities. Ten species were described in the 1980 s, another 10 in the 1990's (4 in 1993), as credit to the research efforts of a number of active opisthobranch biologists. One photo- graph of a living holotype (Chromodoris aurigera Rud- man, 1990) is included I cannot detect any glaring iden- tification errors (although the spelling of .Ajilustritlae is incorrect), and I happily note that the overall SNStematic treatment is up-to-date (perhaps due in part to a "who's who " list of contributors): Akera is placed in .Anaspidea, CijUndrolmUa in Sacoglossa, and Apltistrum anipluslre in Ihjdatina, three recent changes often overlooked by non-specialists. Some might contest the wording of a few- evolutionary statements (e.g., "elysiids have lost their shell"), but I think these are excusable in this context. Wells and Bryce have elegantly met their intended goal to illustrate the extraordinary diversitv of living opisthobranchs from the long, winding coastline ot West- ern Australia. Attendees at future Western Australian Workshops (the next one in the Houtman Abrolhos in 1994) have a excellent guide to follow . Paula M. Mikkelsen 21^1 .3U INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biology and systematics of mollusks. Manuscripts de- scribing original, unpublished research as well as review articles will be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not re- quire an abstract. Notices of meetings and other items of interest to malacologists will appear in a news and notices section. 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