THE NAUTILUS Volume 105 1991 ' I AUTHOR INDEX Bail, P 159 Marshall, B. A 104 BiELER, R 39 Okutani, T 165 BoucHET, P 159 Paul, A. J 173 Braley, R. D 92 QuiNN, J. F., Jr 81, 166 Clifford, H. F 173 Reid, D. G 1,7 Emerson, VV. K 62 Rosenberg, G 147 GoLiKov, A. N 7 Salisbury, R 147 HouART, R 26 Sergievsky, SO 1 HuLiNGS, N. C 16 Sysoev, a. V 119 Kabat, a. R 39 Toll, R B 116 Kantor, Y.I 119 Vidrine, M F 152 Ledua, E 92 Wilson, J. L. 152 Lucas, J. S 92 Zaslavskaya, N. 1 1 NEW TAXA proposed IN VOLUME 105 (1991) gastropoda Calliotropis dentata Quinn, new species (Trochidae) 170 Calliotropis globosa Quinn, new species (Trochidae) 168 Echinogurges tuberculatus Quinn, new species (Trochidae) 170 Gaza olivacea Quinn, new species (Trochidae) 166 Lamellitrochus Quinn, new genus (Trochidae) 81 Lamellitrochus bicoronatus Quinn, new species (Trochidae) 87 Lamellitrochus carinatus Quinn, new species (Trochidae) 84 Lamellitrochus fenestratus Quinn, new species (Trochidae) 85 Lamellitrochus filosus Quinn, new species (Trochidae) 87 Lamellitrochus inceratus Quinn, new species (Trochidae) 88 Lamellitrochus suavis Quinn, new species (Trochidae) 87 Mirachelus acanthus Quinn, new species (Trochidae) 168 Littorina (Littorina) hasatka Reid, Zaslavskaya, and Sergeivsky, new species (Littorinidae) 1 Litturina (Neritrema) naticoides Reid and GoUkov, new species (Littorinidae) 8 Derniomurex (Trialatella) leali Houart, new species (Muricidae) 27 Favartria (Favartia) varimutabilis Houart, new species (Muricidae) 32 Trophon mucrone Houart, new species (Muricidae) 35 Lyria doutei Bouchet and Bail, new species (Volutidae) 160 Vexillum brunneolinea Rosenberg and Salisbury, new species (Costellariidae) 147 Vexillum elliscrossi Rosenberg and Salisbury, new species (Costellariidae) 149 Antiplanes ahijssalis Kantor and Sysoev, new species (Turridae) 125 Antiplanes dendrotiplicata Kantor and Sysoev, new species (Turridae) 127 Antiplanes gabbi Kantor and Sysoev, new name (Turridae) 122 Antiplanes habei Kantor and Sysoev, new species (Turridae) 141 Antiplanes hurilensis Kantor and Sysoev, new species (Turridae) 136 Antiplanes molojimai aquilonalis Kantor and Sysoev, new subspecies (Turridae) 140 Antiplanes obliquiplicata Kantor and Sysoev, new species (Turridae) 128 Antiplanes spirinae Kantor and Sysoev, new species (Turridae) 137 THE NAUTILUS Volume 105, Number 1 March 14, 1991 ISSS 0028-1344 A quarterly devoted to malacology. ' Man„e Broi^caTTa'i;;^;,ry' ' ^ P^^^^^. LIBRARY y£w ^^^^ M/^R 2 5 1991 1 Jl/z ^^^^^^ ^Woods Hole, f/ass. j A Im. I / / M^j^^L i vVv ^ ^gjd^^^^^^^ K ^*^'- ^-'^'m 1^^^^^^^ m //J^ ^P'^'^^i^^^^^ k % fjl P^^'^^S ^ ^^ EDITOR-IN-CHIEF Dr. M. G. Harasewych Division of MoUusks National Museum of Natural History- Smithsonian Institution Washington, DC 20560 ASSOCIATE EDITOR Pr. R. Tucker Abbott American Malacologists, Inc. P.O. Box 2255 Melbourne, FL 32902 CONSULTING EDITORS Dr. Rudiger 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 Historv New York, NY 10024 Mr. Samuel L. H. Fuller 1053 Mapleton Avenue Suffield, CT 06078 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 Universitv Cambridge, MA 02138 Dr. Aurele La Rocque Department of Geology The Ohio State Universitv Columbus, OH 43210 Dr. James H. McLean Department of Malacology Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard Universitv 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 Mr. Richard E. Petit P.O. Box 30 North Myrtle Beach, SC 29582 Dr. Edward J. Petuch Department of Geology Florida Atlantic University Boca Raton, FL 33431 Dr. David H. Stansbery Museum of Zoologv' The Ohio State Universitv Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Harvard Universitv Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology Universitv of California at Davis Davis, CA 95616 SUBSCRIPTION INFORMATION The subscription rate per volume is US $25.00 for individuals, US $35.00 for institutions. Postage outside the United States is an additional US $4.00 for surface and US $12.00 for air mail. All orders should be accompanied b\ pavment and sent to: THE NAUTILUS, P.O. Box 7279, Silver Spring, MD 20907-7279, USA. Change of address: Plea.se inform the publisher of your nev\ 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 THE€7NAUTILUS Volume 105, Number 1 March 14, 1991 ISSN 0028-1344 CONTENTS ^ - j -r. ^J»;JfUtUi> I LIBRARY ; ' 1 David G. Reid Liitorina kasatka. a New Species from the Kurile IslMdO O c iQQi ! Nadya I. Zaslavskaya and Okhotsk Sea l " /^ ^ '^y ' f 1 Sergei O. Sergievsky | ♦ Wqqcj.s Hole. Macs. — j- David G. Reid Liitorina naticoides. New Species, with Notes on the Alexander M. Golikov Other Smooth-shelled Liitorina Species from the Northwestern Pacific 7 Neil C. Hulings Activity Patterns and Homing of Acanthopleura gemmata (Blainville, 1825) (Mollusca: Pohplacophora) in the Rocky Intertidal of the Jordan Gulf of Aqaba 16 Roland Houart The Southeastern Brazilian Muricidae Collected by RV Marion-Dusfresne in 1987, with the Description of Three New Species . 26 News and Notices 38 THE NAUTILUS 105(l):l-6, 1991 Page 1 Littorina kasatka, a New Species from the Kiirile Islands and Okhotsk Sea David C. Reid Department of Zoology The Natural Histor\ Museum London SW7 5BD, U.K. Nadya I. Zaslavskaya Institute of Marine Biology Academy of Sciences of tlie U.S.S.R. Vladivostok 690032, U.S.S.R. Sergei O. Sergievsky Zoological Institute Academy of Sciences of the U.S.S.R. Leningrad 199034, U.S.S.R. ABSTRACT A new species, Littorina (Littorina) kasatka. is described from the Kurile Islands and Okhotsk Sea It is similar in shell char- acters to a smooth form of L. (S'critrcnw) sitkana. with which it is sympatric, but is distinguished by the absence of mam- milliform glands on the penis of the male and by the small jell) gland of the female. It is predicted that L. kasatka produces pelagic egg capsules and has planktotrophic development Kctj words: Littorina. Pacific; Larval development. INTRODUCTION The generic name Littorina has in the past been applied to man\ littorinids with reiativeK large, smooth or spi- rally sculptured shells, that occur in the littoral zone. However, a recent phylogenetic analysis of the Littorin- idae, based largely on anatomical rather than shell char- acters, has defined Littorit}a more narrowly as a mono- phyletic clade of about 20 species (Reid, 1989). Species of Littorina sensu stricto are found only in the northern hemisphere, where they are characteristic inhabitants of the eulittoral and littoral fringe on temperate and cold temperate shores. During the past two decades close investigation of the reproductive anatomy of European Littorina species has revealed the existence of three additional species, pre- viously unrecognized b\ shell characters (review by Raf- faelli, 1982; Johannesson & Johannesson, 1990). The bio- chemical technique of enzyme electrophoresis has confirmed that these species are genetically distinct (re- view by Ward, 1990). More recently, a pair of sibling species has been identified in the northeastern Pacific, again using the dual approach of reproductive anatomy and biochemistry (Murray, 1979; Mastro et ai, 1982). In contrast, the Littorina of the northern and northwestern Pacific are poorly known. This paper describes a new species that was discovered during a genetic study of Littorina kiirila Middendorff, 1848 from the Kurile Is- lands and Vostok Bay (Zaslavskaya & Sergievsky, in press). (Littorina ktirila is a junior synonym of L. sitkana Phi- lippi, 1846, see note added in press in Reid, 1990a). Of 11 loci examined, alleles common to the two species were foimd at only two, and the genetic identitv was conse- quently low (Nei's (1978) genetic identity / = 0.065). The new species was recognized independently during anatomical studies of Littorina material from the Zoo- logical Institute, Leningrad, as part of a systematic study of all Littorina species in the Pacific (Reid, in prep). Recognition of the new species in the field is difficult, since shell form is similar to that of L. sitkana and several other Littorina species, but both males and females dis- play unique reproductive anatomy. MATERIAL AND METHODS The new species is described from six lots in the Zoolog- ical Institute, Leningrad (ZIL), and one lot in the Zoo- logical Museum, Moscow (ZMM), all of which are du- plicated in the Natural History Museum, London (BMNH). That it is indeed undescribed was confirmed by comparison with original descriptions of all available names of Littorina species from the Pacific, and with all available type specimens (housed in BMNH, ZIL and National Museum of Natural History, Washington, D.C.); details will appear elsewhere, in a full account of the systematics of Littorina (Reid, in prep). Closely related species are discussed briefly below. The reproductive anatomy was examined in ten males and nine females. The spiral pattern of the albumen and capsule glands of the pallial oviduct can be seen, without dissection, on the right side of the body adjacent to the columellar muscle, when a female is removed from the shell. Serial histological sections were prepared of the pallial oviduct of two females. Radulae were extracted from six specimens from three localities, and after clean- ing in hot concentrated sodium hydroxide solution were examined by scanning electron microscopy. For com- parison, 45 specimens of L. sitkana were dissected, from localities all around the northern Pacific from Hokkaido to Washington, and four radulae were examined. Details of other Littorina species investigated are given in Reid and Golikov (1990). SYSTEMATIC DESCRIPTION Littorina (Littorina) kasatka new species (figures 1-4, 8-11, 12-20, 23) Page THE NAUTILUS, Vol. 105, No. 1 Figures 1-4. Litlorina kasatha new species. 1. Holotype (HMNH 1990050), Kasatka Point, Iturup I., Kurile Is. 2. Paratype (BMNH 19900.51 ). 3. Nataliya Bay, Lirup I., Kurile Is. 4. Paramushir I,, Kurile Is. Figures 5-7. Litlorina silkana smooth form, from localities at which it is s\mpatric with L kasatka. 5. 7. Kasatka Point, Iturup I , Kurile Is. 6. Nataliva Bay, Urup I., Kurile Is. All specimens in BMNH. Types: holotype BMNH 1990050, Kasatka Point, Iturup I., Kurile Is, USSR (figure 1); 9 dry paratypes BMNH 1990051; 38 paratypes in alcohol BMNH 19900.52; 10 paratypes in alcohol ZIL. Etymology: named after type locality. Shell (figures 1-4): Dimensions: Adult size range 6.4- 11.0 mm shell height. Shape: Teleoconch approximately 4 whorls (apices eroded in available specimens), moderately solid. Glob- ular to turbinate, whorls smoothly rounded, sutures dis- tinct. Columella rather narrow, slightly e.xcavated, pillar concave; sometimes a slight chink between edge of inner lip and base of body whorl (not a true umbilicus). Sculpture: Indistinct axial growth lines only, no spiral grooves; faint spiral striae sometimes visible at high mag- nification. Color: Mid-brown, often with 6-16 rows of smudged whitish dashes, especially at periphery and on base, oc- casionally forming tessellated pattern (figure 4). Aperture purplish brow II, Columella white to pinkish brown. Animal: Head-foot (figures 19, 20): Head black, pale tips to tentacles and snout, unpigmented stripe across tentacle base and eye, or unpigmented patch at inside of tentacle base only. Sides of foot grey to blackish. Operculum: Normal, paucispiral littorinid (type A of Bandel & Kadolsky, 1982). Penis (figures 14-18): Wrinkled base lacking mammil- liform glands; small, blunt filament not clearly demar- cated from base; sperin groove open to tip. Fallial oviduct (figures 12. 13): Complex spiral of 3 loops oi all)iini('n and capsule glands; final .straight section D. G. Reid et al., 1991 Page 3 Figures 8-11. Radulae of Littorina kasatka new species Scale bars = 50 fim, 8, 9. Kasatka Point, Iturup I , Kurile Is. 10. 11. Paramushir 1 , Kurilt- Is 8. 10, N'ieweti ilat 9. I 1. Same radulae as Figures 8, 10, Init viewed troni angle ot 45° to show cusp shape. (containing jelly gland) short, not swollen or septate. Bursa copulatrix small, opening in anterior position. Raditla (figures 8-11): All cusps moderately pointed. Rachidian with 3 large, 2 small cusps; outline of tooth (viewed flat from above) varies from square to oblong (ratio of length of tooth : width at mid-point 1.00-1.78). Lateral and inner marginal each 4 large cusps. Outer marginal 6-8 cusps. Distribution: Habitat: Rocks and boulders in middle and upper littoral zone of sheltered shores. Range (figure 23): Okhotsk Sea. Kurile Islands, Gulf of Shelikov in Records: Kasatka Bay and Kasatka Point, Iturup 1.; Na- taliya Bay, Urup I.; Ryponkich I.; Paramushir I. (all ZIL and BMNH); between Nepropusk and River Gank, W. Kamchatka, Gulf of Shelikov (ZMM, BMNH). Similar species: Four Littorina species are geograph- icalK sympatric with L. kasatka over its known range: L. (Littorina) squalida Broderip & Sowerby, 1829; L. (Neritrema) sitkana; L. (N.) subrotundata (Carpenter, 1864) and a newly described species (Reid & Golikov, 1990). Of these, no confusion will arise with L. squalida. which is a much larger (12-44 mm), spirally-grooved shell (figured by Reid, 1990a). Littorina subrotundata and the newly described species are also usualK larger (4-14 mm and 6-16 mm respectively), both are a more patulous shape, and both have a color pattern of dark brown spiral lines on a paler brown ground (figured by Reid & Golikov, 1990). Anatomically, these two species both show mammilliform penial glands in the male and a large, swollen, septate jelly gland in the female, unlike L. kasatka. Littorina sitkana is the most similar to L. kasatka and is apparently found in the same habitat, since all but one of the seven known collections of L. kasatka were sep- arated from mi.xed lots of the two species. Littorina sitkana is highly variable. The form that occurs most commonly in the northeastern Pacific has strong spiral ribs with striae in the grooves between, but in the north- western Pacific (and especially in the Kurile Islands and Okhotsk Sea) the common form is smooth shelled. This smooth form has been referred to as L. kurila (this name is actually a synonym of L. sitkana, see Reid, 1990a) in all recent Soviet literature (e.g., Golikov & Kusakin, 1978). Since L. kasatka lacks spiral ribs and obvious striations, it is easily distinguished from sculptured shells of L. sitkana (figured by Reid, 1990a). However, smooth forms of L. sitkana are extremely similar. Littorina sitkana is usually larger (commonly 15 mm, up to 23 mm), has a Page 4 THE NAUTILUS, Vol 105, No. 1 Figures 12-20. Litiorina kasatka new species. 12, 13. Pallial oviducts. 11-18. Penes. 19, 20. Heads. 12. 17. Nataliya Bay, Unip I. 13-15, 20. Kasatka Point, Iturup I. 16. Kasatka Bay, Ituriip I 18, 19. Paramushir I. Figures 21, 22. Littorina sitkana, Seldovia, Alaska. 21. F'allial oviduct. 22. Two views of penis; mammilliform glands shaded .Ml specimens in BMNH Key to figures 12, 13, 21: sparse stipple, albumen gland; dense stipple, capsule gland; dotted lines, septa of jelK gland (visible by transparency); dashed lines, bursa copulatri.v (visible only by dissection or .serial sectioning). slightly more inflated last whorl, and the columella is relatively wider (figures 5-7). Coloration also differs, L. sitkana being purple brown or blackish, sometimes with one or two white or orange spiral bands, and never show- ing the flecked or tessellated pattern of L. kasatka. The only entirely reliable means of distinguishing the two species is by their reproductive anatomy: in females, the pallial oviduct of L. sitkana has two loops in the spiral pattern, and a large, swollen, septate jelly gland (figure 21 ), whereas there are three loops and a small jelK gland in L. kasatka; in males, mammilliform penial glands are present in L. sitkana (figure 22), but absent in L. kasatka. Shells of the new species do not resemble closely those of am other Littorina from the northeastern Pacific or D. G. Reid et al.. 1991 Page 5 Figure 23. Distribution of Litlorina kasaiha new species Rec- ord from Gulf of Slielikov could not be preciseh localized. Atlantic. No other Littorina has a pallial oviduct of the same form (Reid, 1990a), and only in L. scutulata Gould, 1849 and L. striata King and Broderip, 1832 does the penis also lack mammilliform glands. DISCUSSION The diversity of types of spawn and development within the genus Littorina have been reviewed by Reid (1989, 1990a), who showed that the type of spawn can be pre- dicted from the relative sizes of the glandular compo- nents of the pallial oviduct. From its small jelly gland, lack of brood pouch and presence of a large capsule gland, it is likely that L. kasatka produces pelagic egg capsules. In other Littorina species that occur at high latitudes and reproduce in this way, the capsules are large (about 1 mm in diameter), biconve.x and contain several eggs, and the capsules of L. kasatka are probably similar. It has been suggested that large capsules of this type are adaptive in relation to high levels of predation on pelagic propagules in cold seas (Reid, 1990a). The only other Pacific Littorina with pelagic capsules that occurs at such a high latitude is L. squalida. All known littorinids with pelagic capsules show planktotrophic development, so this can be predicted in L. kasatka. In contrast, the other four Littorina species in the northern Pacific are non- planktotrophic, developing within benthic gelatinous egg masses (Reid & Golikov, 1990). A cladistic analysis of the 20 or so species of Littorina then known was given by Reid (1990a), and used as a basis for subgeneric classification. With three loops in the spiral of the pallial oviduct, a small jelK gland and probable pelagic egg capsules, L. kasatka is clearly a member of the paraphyletic subgenus Littorina. Within that area, its relationships cannot yet be resolved by cladistic analysis, because of uncertainty about its spawn The presence of a bursa copulatri.x in an anterior position does, however, suggest that its most likely place on the cladogram is between nodes 9 and 10 (see figure 5 in Reid, 1990a). As a result of the cladistic analysis the subgenus Neritrema was defined b\ the synapomorphies of anterior bursa copulatri.x, large jelly gland and non- planktotrophic development (Reid, 1990a). In view of the anterior bursa of L. kasatka, this character must now be removed from the list of synapomorphies of Neritre- ma. The cladogram has also been used for a biogeo- graphical analysis, which showed that in general the more derived species of northwestern Pacific Littorina occur at higher latitudes (Reid, 1990b). The probable position in the cladogram and known geographical dis- tribution of L. kasatka are consistent with this trend. ACKNOWLEDGMENTS A visit to the Soviet I'nion by DGR was supported by the exchange program of the Royal Society of London and the Academy of Sciences of the USSR. For permis- sion to work at the Zoological Institute, Leningrad, DGR would like to thank the director. Prof. O. A. Scarlato. Prof. A. N. Golikov gave much valuable help with the collections there. Photography \\asb\ H. Taylor (BMNH) and histology by D. W. Cooper (BMNH). ' LITERATURE CITED Bandel, K and D Kadolsky 1982. Western Atlantic species of Nodilittorina (Gastropoda: Prosobranchia): compara- tive morpliology and its functional, ecological, phyloge- netic and taxonomic implications, X'eliger 25:1-42. Golikov, A. N. and O. G. Kusakin. 1978. Recent molluscs of the littoral zone of the U.S.S.R. Nauka. Leningrad, 256 p. Johannesson, K. and B. Johannesson. 1990. Genetic variation within Littorina saxatilis (Olivi) and Littorina neglecta Bean: is L. neglecta a good species? Hsdrobiologia 193: 89-97. Mastro, E., V. Chow, and D. Hedgecoek. 1982. Littorina scutulata and Littorina plena, sibling species status of two prosoliranch gastropod species confirmed by electropho- resis. Veliger 24:239-246. Murray, T. 1979. Evidence for an additional Littorina species and a summary of the reproductive biology of Littorina from California. Veliger 21:469-474. Nei, M. 1978. Estimation of average heterozygosity and ge- netic distance from a small number of individuals. Ge- netics 89:583-590. Raffaelli, D. G. 1982. Recent ecological research on some European species of Littorina. Journal of Mollnscan Stud- ies 48:342-354. Reid, D. G. 1989. The comparative morphology, ph\ logeny and evolution of the gastropod famiK Littorinidae. Philo- sophical Transacations of the Royal Society of London, Series B 324:1-110. Reid, D. G. 199(ta. \ cladistic phylogeny of the genus Lit- torina (Gastropoda): implications for evolution of repro- ductive strategies and for classification H\drobiologia 193: 1-19. Reid, D. G. 1990b Trans-Arctic migration and speciation induced b\ climatic change: the biogeography of Littorina Page 6 THE NAUTILUS, Vol. 105, No. 1 (MoUusca: Gastropoda). Bullt-tin of Marine Science 47:35- 49. Reid, D. G. and A. N. Golikov. 1991. Liltorina nalicoides. new species, with notes on the other smooth-shelled species from the northwestern Pacific. Nautilus 105(1 ):7-15. Ward, R D 1990 Biochemical genetic variation in tlie genus I^ittorina (Prosofiranchia Mollusca) H\dr()liiologia 193: 53-69. Zaslavskaya, N, 1 and S. O Sergievsky. In press. Genetic comparison of Atlantic and Pacific species of Liitorina. Proceedings of the 9th Malacological Conference of the L'.S.S.R Leningrad: Zoological Institute, THE NAUTILUS 105(1):7-15, 1991 Page 7 Littorina naticoides, New Species, with Notes on the Other Smooth-shelled Littorina Species from the Northwestern Pacific David G. Reid Department of Zoology The Natural History Museum London SW7 5BD. U.K. Alexander N. Golikov Zoological Institute Academy of Sciences of the L'.S.S.R. Leningrad 199034, U.S.S.R. ABSTRACT A new species. Littorina (Neritrerna) naticoides. is described from the northwestern Bering Sea, Kamchatka and Kurile Is- lands. It is believed to be the sister species of L. aleutica and the\ can be distinguished by penial and shell characters The two are allopatric, L. aleutica showing an oceanic distribution and L. naticoides a more continental one, but their ranges are known to approach to u ithin 2.50 km of each other Five Lit- torina species from the northwestern Pacific have smooth-shelled forms, which can be difficult to distinguish. These are L. {N.) naticoides, L. (N.) aleutica. L. (N.) sitkana. L. (N.) suhrotun- data and L. (Littorina) kasatka. The diagnostic features of their shells and reproductive anatomy are reviewed. Key words: Littorina; Pacific; biogeography. INTRODUCTION Systematic studies of the family Littorinidae have been revolutionised by two developments over the past two decades. Firstly, anatomical features have been found to be more reliable than traditional shell characters for identification of species, and, secondly, there has been a growing appreciation of the wide range of shell variation shown by some species, and an understanding of its bi- ological significance. The first use of anatomical char- acters to discriminate between sibling species (with sim- ilar or identical shells) was in the genus Littorina. (The generic name is here used in the strict sense, as defined by the cladistic analysis of Reid, 1989a). The most in- formative of these new characters have proved to be the shape of the penis, type of egg capsules and development (Sacchi & Rastelli, 1966; Heller, 1975; Hannaford Ellis, 1979; Murray, 1979) and also the form of the pallial oviduct (Reid, 1989a, 1990a). In all cases, studies of elec- trophoretically detectable genetic variation have sup- ported the status of the sibling species initially recognized by anatomical differences (Ward, 1990). Although some- times only one or other sex can be unequivocally iden- tified by anatomical characters, there are no known cases of morphologically inseparable Littorina species. Extreme intraspecific variation in coloration, sculpture and shape of the shell is a well-known feature of Lit- torina. and contributes to the difficulties of identification. Shell variation is especially marked in those species which show direct development, for here the lack of a widely- dispersed larval phase enhances the potential for adap- tation to local environments. Selective factors influencing morphological and color variation between populations are believed to include predation, wave action and dam- age by mobile boulders (e.g., review by Raffaelli, 1982; Janson, 1982, 1983; Johannesson, 1986; Seeley, 1986). Other factors such as growth rate can also contribute to non-genetic shell variation (Kemp & Bertness, 1984). The genus Littorina is restricted to the northern hemi- sphere, and most species occur on temperate and cold temperate shores (Reid, 1990b). So far, modern system- atic revisions using anatomical details have only been done in Europe and the northeastern Pacific. Littorina species are also abundant in the northwestern Pacific, and the most recent review of the littoral molluscs of the Siberian region (Golikov & Kusakin, 1978) recognized six species: L. squalida Broderip & Sowerby, 1829; L. brevicnla (Philippi, 1848); L. rnandshurica Schrenck, 1861; L. aleutica Dall, 1872; L. sitkana Philippi, 1846; L. kurila Middendorff, 1848. The identifications were based only on characters of the shells. The first anatom- ical work on northwestern Pacific Littorina was done as part of a cladistic analysis of the 20 species then recog- nized in the genus (Reid, 1990a). This largely supported the classification of Golikov and Kusakin (although L. kurila was synonymized with L. sitkana), but the ana- tomical material used did not include any from the Soviet Union. As part of a wider study of the systematics and dis- tribution of Littorina species (Reid, in prep), it has re- cently been possible to examine the extensive collection of preserved material from the Siberian coast held in the Zoological Institute, Leningrad. Preliminary results in- dicate that L. squalida. L. brevicnla and L. rnandshurica can each be readily identified by shell characters, as described and illustrated by Golikov and Kusakin (1978) and Reid (1990a). The remaining three of the species listed above were distinguished by Golikov and Kusakin (1978) primarily by their shell sculpture: L. sitkana hav- ing spiral ribs, L. kurila being smooth or almost so, and L. aleutica having rows of nodules. This division now seems to be an artificial one. Differences in reproductive Pages THE NAUTILUS, Vol. 105, No. 1 anatomy define five .species in this group: L. sithana, L. alentica, L. sitbwtundata (Carpenter, 1864), L. hasatka (recently described by Reid et al., 1990) and L. nati- coides, which is described herein. The shell characters of all five species are variable, and in particular L. sit- kana. L. aleutica and L. stihrottindata can show smooth or sculptured shells. "Littorina kurila," as defined by Golikovand Kusakin (1978), embraced smooth or slightly sculptured forms of all five species. The nomenclature of these species is rather complex. During the wider study of Littorina systematics all avail- able type specimens (housed in Natural History Museum, London; National Museum of Natural History, Wash- ington, D.C.; Zoological Institute, Leningrad) have been examined and all original descriptions consulted, to as- certain that the new species is indeed undescribed. The available names include 18 synonyms of L. sitkana and three of L. subrotundata. Complete synonymies and full accounts of the described species w ill be given elsewhere. Here it need only be noted that preserved syntypes of L. kurila Middendorff, 1848 in the Zoological Institute, Leningrad, have shown that this taxon is a synonym of L. sitkana (see note added in proof in Reid, 1990a). As discussed below, the name "L. kurila" was incorrectly applied by Reid (1989a, 1990a, b) to what is here con- sidered an open-coast form of L. subrotundata. In the present paper the new species will first be de- scribed, then the four other similar species which some- times show smooth shells will be discussed and distin- guished. MATERIALS AND METHODS This resolution of the smooth-shelled Littorina species of the northwestern Pacific is based on examination of all material in the collections of the Natural History Museum, London (BMNH), the Zoological Institute, Leningrad (ZIL) and the National Museum of Natural History, Washington, D.C. (USNM). The new species is described from 21 lots in the ZIL, many of which are duplicated in the BMNH. A total of 27 males and 18 females were dissected, from 8 localities covering the known range of the species. Five radulae were extracted from specimens from three localities, and after cleaning in hot, concentrated sodium hydroxide solution were ex- amined by scanning electron microscopy. For compar- ison, 32 specimens of L. aleutica, 82 of L. subrotundata, 45 of L. sitkana and 19 of L. kasatka were dissected, all from a wide range of localities. SYSTEMATIC DESCRIPTION Littorina (Neritrema) naticoides new species (figures 1-12, 15-23, 28-35, 37) Types: holotype BMNH 1990053, Milne Bay. Simushir 1., Kurile Is, USSR (figures 1, 16); 2 drv paratypes BMNH 1990054; 6 paratypes in alcohol BMNH 1990055; 2 para- types in alcohol ZIL. Etymology: Natica-\ike, a reference to shell shape. Shell (figures 1-12): Dimensions: Adult size range 6.2-15.9 mm shell height. Shape: Teleoconch approximately 4 whorls, moderately solid. Turbinate, often rather patulous, whorls smoothly rounded, sutures distinct. Ciolumella relatively wide, slightly excavated, pillar concave. Sculpture: Protoconch 0.73-075 mm diameter, 1.8 whorls. Teleoconch: indistinct axial growth lines only, no spiral grooves; faint spiral striae sometimes visible at high magnification, but no strong striations. Color: Orange brown to dark purple brown, usually 5- 12 dark brown or black spiral lines; sometimes a pale spiral band on base. Aperture orange brown to purple brown. Columella white, sometimes tinged purple brown. Animal: Head-foot (figures 28-30): Head grey to black, paler at tips of tentacles and snout; unpigmented patch over eye and usually another at inside of tenacle base. Sides of foot usually pale grey speckles or unpigmented, sometimes blackish. Operculum: normal, paucispiral littorinid (tvpe A of Bandel & Kadolsky, 1982). Penis (figures 15-20): Wrinkled base with 4-15 mam- illiform penial glands in single row on anterior edge (one specimen with a single gland); filament short, less than half length of base, broadly triangular, with simple sub- epithelial glandular region (more opaque in preserved specimens); sperm groove open to tip. Pallial oviduct (figures 21-23, 37): Complex spiral of 3 loops, first of albumen gland, second and third of capsule gland; final straight section a large, swollen, septate jelly gland. Bursa copulatrix in anterior position. Spawn: Benthic, gelatinous egg mass up to 6 mm in diameter, containing spherical capsules 0.7-0.8 mm in diameter, each with single egg surrounded by albumen layer 0.5-0.6 mm in diameter. Radula (figures 31-34): All cusps bluntly rounded. Ra- chidian with 3 large, 2 .small cusps; outline of tooth (viewed flat from above) varies from square to oblong (ratio of length of tooth: width at mid-point 0.77-2.0). Lateral and inner marginal teeth each 4 large cusps. Outer mar- ginal 5-7 cusps. Distribution: Halntat: A range of intertidal habitats are reported: on dead shells in sheltered inlets; Mytilus Figures 1-12. Littorina naticoides new species. 1. Holotype (BMNH 1990053), Milne Hay. Simushir I., Kurile Is. 2, 3. Ozernovskiy, Kamchatka. 4. 5. 10 km east of Cape Kamchatski\, Kamchatka. 6. Avacha Ba>, Kanu hatka 7. Kronot.skiy Gulf, Kamchatka 8- 10. Egvekinot Inlet, .'\nadyr.skiy (iiiif II, 12. Ozcriiiy River e.stuary, Kamchatka Figure I.'l. Littorina aleutica. Provideniya. Figure 14. Littorina stilnotundata. Milne Bay, Simushir 1 , Kurile Is (.All specimens in BMNH). D. G. Reid and A. N. Golikov, 1991 Page 9 Page 10 THE NAUTILUS, Vol. 105, No. 1 15-27 28-30 Figures 15-23. 28-30. Littorina naticoides new species. 15-20. Penes: mamillifdrm j;lands on left edge, sometimes visible by transparency 21-23. Pallial oviducts 28-30. Heads. 15, 17. 18. Ozernovski\, Kamchatka 16. Penis of holot\pe, Milne Bay, Simusliir I , Kiirile Is. 18. 19. 23, .30. Krista Gulf, Anadyrskiy Gulf 20. F.gvekinot Inlet, .\nadyrskiy Gulf 21.' 29. Kronotski'y Ba>, Kamclialka. 22. 10 km east of Gape Kanicliatskiy, Kamchatka Figures 24-27. Littorina alculica. 24-26. Penes 24, 26. Lihaclieva, Providcniya. 25. Adak I., Aleutian Is. 27. Pallial oviduct, Nizki I., Aleutian Is. Key to figures 21-23, 27: sparse stipple, albumen gland; dense stipple, capsule gland; dotted lines, septa of jelly gland (visible by transparency); dasfied lines, bursa copulatrix (visible only by dissection). zone on exposed shores; on Halosaccion (red alga); in Alaria (brown alga) bell. Range (figure 3.5;.Kurile Islands, .south and wesi Kam- chatka, Anadyrskiy Gulf. Records: Kiirile Is: Urnp I.; Milne Bay, Simushir I.; Spaseniya, Simushir I.; Diani Bay, Ketoi I., Paramushir I.; Kamchatka: Ozerniy River estuary, Okhotsk Sea; Ozerriovskiy; Avachinskaya Bay, near C^ape Viikova; Avacha Bay, near Petropavlovsk; Kronotskiv Gulf; 10 km east of Cape Kamchatskiv; .Anadyrskiy Gulf: Egvekinot Inlet. Krista Gulf (ail ZIL and BMNH). D. G. Reid and A. N. Golikov, 1991 Page 11 Figures 31-34. Radiilae of Littorina naticoidcs new species Scale bars = 50 ixm. 31, 32. 10 km east of Cape Kamchatskiy, Kamchatka. 33, 34. Egvekinot Inlet, Anadyrskiy Gulf. 31, 33. Viewed flat, 32, 34. Same radulae as figures 31, 33, but viewed from an angle of 45° to show cusp shape. Similar species: As mentioned above, L. kurila Mid- dendorff, 1848 is a junior synonym of L. sitkana Philippi, 1846, but the name has been widely used in the literature for any of the five Littorina species (including L. nati- coides) in the northwestern Pacific that frequently have smooth shells. The other four members of the group will be considered in turn, and the characters of all five spe- cies are summarized in table 1. Radular characters have not proved useful in discriminating between them. L. (Neritrema) .sitkana (figure 39): The typical form of this species has a strongly carinate shell with spiral stri- Tabh' 1. Sununar\ of characters of five Littorina species from the northwestern Pacific. sultro- aleulica kasatka naticoides sitkana Inndala Shell: adult size (mm) 6-14 fi-11 6-16 6-23 4-14 patulous shape + - + - ( + ) nodulose sculpture ( + ) — - - - spiral ribs { + ) - - ( + ) ( + ) spiral microstriae + — - ( + ) - pattern of dark spiral lines { + ) — + ( + ) + pattern of pale flecks - + - - - Penis: niamilliform glands 7-12 0 1-15 5-14 5-17 filament > '2 length of base + - - - - Pallia! oviduct: spiral pattern fig. 38 fig. 36 fig. 37 fig. 39 fig. 40 swollen, septate jelly gland + — + + + Spawn benthic egg mass + - + + + pelagic capsules - - - — + = Present; ( + ) = = sometimes present: - = absent. Page i; THE NAUTILUS, Vol. 105, No. 1 Figure 35. Distriliution of Littorina naticoides new species. ations between the carinae (e.g., Reid, 1990a: fig. 11) and is most frequent in the north and northeastern Pacific from Oregon to the Aleutian Islands, while in northern Japan it is usually only the base that bears strong grooves. In either case the shell is thereby distinguished from all others in the group except rare forms of L. subrotundata. Problems arise with the smooth forms of this species (Reid et ai, 1990: figs 5-7), which predominate in the Kurile Islands and Kamchatka, and also occur in other parts of the range. These shells may be quite large (up to 23 mm), are less patulous than L. naticoides and L. aleutica and are often uniformly black or brown (except in the Okhotsk Sea, L. sitkana commonly lacks the pat- tern of narrow black lines seen in L. naticoides and L. subrotundata). Synonyms based on smooth forms of this species include L. kurila Middendorff, 1848, L. suhtene- brosa Middendorff, 1848 and L. sitkana var. atkana (Dall, 1886). Females of L. sitkana can be immediately rec- ognized by the unique spiral pattern of the pallial oviduct (figure 39; Reid et ai, 1990: fig. 21) with its long back- ward loop of capsule gland, but the penis is similar to that of both L. naticoides and L. sulnutundata. L. (Neritrema) subrotundata (figures 14, 40): There has been confusion about the identity of this species, and the name is here used in a wider sense than by previous authors. As interpreted here, the species has a wide dis- tribution around the northern Pacific, from northern Cal- ifornia thruugli the Aleutian Islands to the soutiiern Ku- rile Islands. The name has previously been applied only to the thin- shelled, tall-spired form (Reid, 1990a: fig. In) initially described by Carpenter (1864) as a doubtful member of the genus Assiminea. Synonyms are ?Faludinella cas- tanea Carpenter, 1865, and Paludinella newcombiana Hemphill, 1877, and the species has .sometimes been placed in the genus Algamorda Dall, 1918 (a synonym of Neritrema Rechiz, 1869, regarded as a subgenus of Littorina. see Reid, 1989a). The species was poorly known and lielieved to be restricted to brackish-water salt marshes from northern California to Washington (MacDonald, 1969; Reid, 1990b). The form from this habitat is anal- ogous to the form " tenelnosa" of the .Atlantic L. saxatilis (Olivi, 1792), which is also small, thin-shelled and tall- spired (Muus, 1967; Janson & Ward, 1985), probably as a result of similar selection pressures or ecophenotypic effects in the brackish lagoons in which it occurs. This form of L. saxatilis contrasts with the lower-spired and more patulous form on exposed coasts (e.g., Raffaelli, 1979; Janson, 1982). As a result of the dissection of additional material, it is now believed that, like L. saxatilis, L. subrotundata also occurs in habitats covering a wide range of wave exposures, and has a similarly wide range of variation in shell morphology. The forms from exposed rocky coasts, here assigned to L. subrotundata for the first time, have (like L. saxatilis from similar habitats) a lower spire and larger aperture than the salt-marsh form, and were pre- viously incorrectly identified as L. kurila (Reid, 1990a, b). A shell of this type from the Aleutian Islands was illustrated b\' Reid (1990a: fig. Im), and figure 14 shows a slightly more patulous form. A population from the exposed shores of Tatoosh Island, Washington, has re- cently been studied by Boulding (1990), who concluded that it was an undescribed subspecies of L. kurila. How- ever, examination of specimens from this locality has revealed no significant anatomical differences from ei- ther typical L. subrotundata collected in salt marshes in Washington, or from specimens of this species from the Aleutian Islands. Furthermore, no diagnostic allelic dif- ferences between samples from these three areas have been found by enzyme electrophoresis (E. G. Boulding, personal communication). The available evidence there- fore suggests that they are conspecific, but further study would be desirable. Neither the known geographical dis- tribution of L. subrotundata, nor its considerable vari- ation in shell morphology throughout its range, support its division into subspecies. One reason why the name L. subrotundata has not previously been used for the exposed-coast form is that, at least in Washington, it appears to be completely eco- logically segregated from the t>pical form in salt marsh- es, because the species is not known to occur in inter- mediate habitats. This locally disjunct distribution need not, however, imply genetic isolation of the two forms. It could be maintained, for example, by the action of crab or other predators. It is known that the exposed- coast form is susceptible to attack by crabs because of its thin shell, and that crabs are more abundant in pro- tected rocky habitats than on exposed coasts (Boulding, 1990). It is possible that both exposed coasts and salt marshes are refuges for this species from crab predators. Further north in its range, in Alaska (personal observa- tion), L. subrotundata does occur on both exposed and sheltered rocky shores and there is continuous variation in shell shape l:)etween the two extremes of habitat. In- D. G. Reid and A. N. Golikov, 1991 Page 13 36 37 38 39 40 Figures 36-40. Diagrammatic represfiitation of pallial oviducts of Littorina species from the northwestern Pacific- 36. Litlorina (Litlorina) kasalka. 37. L. (S'eritrema) naticoides. 38. L. (N.) aleutica. 39. L. (N.) sitkana. 40. L. (iV.) suhroUmdala. Key; continuous line with spiral loops represents path of egg groove through pallial oviduct; thick section, capsule gland; dense stipple, seminal receptacle at posterior end; anterior sac, bursa copulatrix; dotted lines, septa of jelly gland. Shape of loops of capsule gland and its relative size are most important features. Oviducts of L. aleutica and L. naticoides sometimes indistinguishable; c.f figures 21-23, 27, terestingly, there is again a parallel with L. saxatilis, which shows a comparable segregation of low-spired and high-spired forms on exposed coasts and in salt marshes respectively, at the southern limit of its European range (Gofas, 1975; personal observation), whereas at more northerly latitudes it occupies the entire spectrum of habitats and morphological intermediates are common (Janson & Ward, 1985; personal observation). The shell of L. subrotundata is often extremely similar to that of L. naticoides in size, coloration and shape, although it does not attain such extreme patulous forms as the latter (e.g., figures 3-5). Like L. naticoides, it is usually entirely smooth, but occasional forms have spiral ribs; in such cases the lack of strong spiral striations in the grooves distinguishes it from L. sitkana. Once again, anatomical characters are more helpful; the form of the pallial oviduct is diagnostic (figure 40) with a relatively smaller capsule gland than that of either L. sitkana or L. naticoides, but the penis is similar to those of both these species. L. (Neritreryia) aleutica (figures 13. 24-27, 38): Shells of L. aleutica from the .Aleutian Islands are usually sculp- tured by four nodulose cords with strong spiral striations in the intervening grooves (Reid, 1990a: fig. Ik). How- ever, in the northwestern Bering Sea the shells are not nodulose; slight spiral ribs may remain, as do the stria- tions (visible at the periphery in figure 13), which help to distinguish the shell from the very similarly shaped, but entirely smooth, L. naticoides. In this case it is the male anatomy that is diagnostic, the penis showing a narrower filament at least half as long as the wrinkled base, whereas that of L. naticoides is shorter (figures 24- 26, c.f. figures 15-20). The pallial oviduct is similar only to that of L. naticoides. but the jelly gland is usually relatively larger and the capsule gland a little smaller (figures 22, 38, c.f. figures 21-23, .37). L. (Littorina) kasatka: This newly-described species (Reid et ai, 1990) is the smallest of the five smooth- shelled Littorina species (6-11 mm) and has the most restricted distribution, being recorded only from the Ku- rile Islands and Gulf of Shelikov. The shell is closest to small, smooth specimens of L. sitkana, although its shape is subtly different, with a relatively narrower columella and smaller aperture. Coloration also differs, the shell of L. kasatka being brown, often with indistinct white flecks or tessellation, unique among the five species discussed here. Despite the superficial similarity of the shell, this species is probably not closely related to the others in the group, since its reproductive anatomy is markedly dif- ferent. Mamilliform penial glands are absent in the male, while the pallial oviduct lacks a swollen, septate jelly gland and shows a unique spiral pattern of albumen and capsule glands. These characteristics of the oviduct in- dicate that the spawn consists of pelagic egg capsules, probably with planktotrophic development, and for this reason it is classified in the subgenus Littorina (Reid et ai, 1990). In contrast, the four other species have a large jelly gland, benthic egg masses and direct development, and are therefore members of the subgenus Neritrema (Reid, 1989a, 1990a). DISCUSSION An earlier review of living Littorina species with a cla- distic analysis of their relationships (Reid, 1990a) must be revised in the light of the new information on material Page 14 THE NAUTILUS. Vol. 105, No. 1 from the Siberian coast. At least three changes are nec- essary. Firstly, "L. kurila" {non Middendorif, 1848) of the earlier publication is now believed to be the same as L. sulrrotundata (no differentiating characters were found in the earlier analysis). Secondly, L. kasatha must be added to the cladogram (Heid 1990a: fig. 5), in a position between nodes 9 and 12 (as yet unclear because of lack of information about egg capsules, but between 9 and 10 most likely, Reid et al. 1990). Thirdly, L. naticoides must be added; this is clearly the sister species of L. aleutica, since with the exception of shell sculpture, all characters used in the cladistic analysis are the same in both species. This sister-group relationship is supported by the sim- ilar oviducts, egg masses, penial glands and patulous shells. Consistent differences in the relative length of the penial filament and in shell sculpture, and small differences in relative sizes of oviducal glands, are, however, sufficient to separate L. naticoides as a new species. So far, the two species have not been found sympatrically. Littorina aleutica is recorded from the Aleutian, Komandor and Pribiloff Islands, and from St Lawrence Island and Provi- deniya in the northern Bering Sea. Littorina naticoides occurs from Anadyrskiy Gulf to the Kurile Islands, ap- proaching within 350 km from the closest record of L. aleutica in the northern Bering Sea, and within 250 km in eastern Kamchatka. In part this apparent allopatry could be explained by lack of collections from the remote northwestern shores of the Bering Sea. However, the separate distributions in the Aleutian, Komandor and Kurile Islands and in Kamchatka are probably real, be- cause there are extensive collections of Littorina from all these areas. Furthermore, the two distributions suggest a possible ecological separation. Littorina aleutica has the more "oceanic" distribution, occurring only on is- lands and at the tip of the Chukotskiy Peninsula, whereas L. naticoides is more "continental," occurring on main- land coasts, even in inlets and estuaries, as well as in the Kurile Islands. There are similar examples elsewhere in the Littorinidae of closely related congeners with con- trasting "oceanic" and "continental" distributions (Lit- toraria, Reid, 1986; Peasiella, Reid, 1989b), but hitherto the phenomenon has been demonstrated only in the trop- ics. The biogeography of Littorina has been analysed by Reid (1990b). These modifications to the cladogram and distributional data on new species do not alter the earlier conclusion that in general in the northeastern Pacific more apomorphic species occur at higher latitudes, per- haps as a result of climatic cooling which induced spe- ciation. Indeed, recognition of "L. kurila" (sensu main text of Reid, 1990a and Reid, 1990b) as conspecific with L. sulrrotundata removes the anomaly of the latter s northeastern Pacific distribution. However, L. naticoides and L. aleutica are the only known example in the genus of sister species that occur in the same ocean and show allopatric distributions; in other ca.ses distributions show broad overlap This could indicate that a different spe- ciation mechanism has been involved here. ACKNOWLEDGMENTS The visit of DCR to the Zoological Institute, Leningrad, was supported by the exchange program of the Ro\al Society of London and the Academy of Sciences of the USSR. DGR would like to thank the Director of the Institute, Prof. O. A. Scarlato, for the opportunity to work there. Photography of shells was b\ P R Crabb(BMNH). LITERATURE CITED Bandel, K. and D Kadolsky. 1982. Western .Atlantic species of Nudilittorina (Gastropoda: Prosobranchia): compara- tive inorpliology and its functional, ecological, phyloge- iietic and taxonomic implications. Veliger 25:1-42 Boulding, E. G. 1990. Are the opposing selection pressures on exposed and protected shores sufficient to maintain genetic differentiation between gastropod populations with high intermigration rates? Hydrobioiogia 193:41-52 Carpenter, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North .America. Report of the British .Asso- ciation for the Advancement of Science 1863:517-684. Ciofas, S. 1975. Sur I'extension de Littorina saxatitis (Olivi) (Moll. Gaster.) dans le Detroit de Gibraltar. Bulletin de la Societe des Sciences Naturelles et Physiqes du Maroc 55: 95-99. Golikov, A. N. and O. G. Kusakin 1978. Recent molluscs of the littoral zone of the I'.S.S R Nauka, Leningrad, 256 p. Hannaford Ellis, C J. 1979, Morphology of the oviparous rough w'ink\e Littorina arcana Hannaford Ellis. 1978, with notes on the taxonomy of the L saxatilis species-complex (Prosobranchia: Littorinidae). Journal of Conchology 30: 43-56. Heller, j. 1975, The taxonomy of some British Littorina spe- cies with notes in their reproduction (Mollusca: Proso- branchia). Zoological Journal of the Linnean Societv' 56: 131-151. Janson, K. 1982. Phenotypic diftereritiatioii in Littorina sax- atilis Olivi (Mollusca, Prosobranchia) in a small area on the Swedish west coast. Journal of Molluscan Studies 48: 167-173. Janson, K. 1983. Selection and migration in two distinct phe- notypes of Littorina saxatilis in Sweden. Oecologia, Berlin 59:,58-61. Janson, K. and B D. Ward, 1985, The taxonomic status of Littorina tcnebrosa Montagu as assessed bv morphological and genetic analyses. Journal of Conchology 32:9-15. Johannesson, B. 1986. Shell morphology of Littorina saxatilis Olivi: the relative importance of physical factors and pre- dation. Journal of Experimental Marine Biology and Ecol- ogy 102:183-195. Kemp, P. and M. Rertness, 1984, Snail shape and ijrowth rates: evidence for plastic shell allonu-try in Littorina lit- torea. Proceedings of the National .Vcademy of Sciences of tlif LSA 81:811-81.3, MacDonald, K, B, 1969, Mollu.scan faunas of Pacific coast salt marshes and tidal creeks. Veliger 11:399-405 Murray, T. 1979. Evidence for an additional Littorina species and a summary of the reproductive biologv of Littorina from California, Veliger 21 469-474 Muus, B J, 1967 The faiuia of Danish estuaries and lagoons. Meddeielser fra Damnarks Fiskeri-og lla\ uiidersfigelser, N.S. 5:1-316. D. G. Reid and A. N. Golikov, 1991 Page 15 Raffaelli, D. G. 1979. The taxononn of tlie IJttorina saxiililis species-complex, with particular reference to the system- atic status of Litlorinu palula Jeffreys. Zoological Journal of the Linnean Societ\ 65219-232. Raffaelli, D. G. 1982 Recent ecological research on some European species of Littorina. Journal of Molluscaii Stud- ies 48:342-:354 Reid, D. G. 1986. The littorimd molluscs ol mangrove forests in the Indo-Pacific region: the geinis Littoraria. British Museum (Natural History), London, 228 p. Reid, D. G. 1989a. The comparative morphology, ph\logeny and evolution of the gastropod family Littorinidae Philo- sophical Transactions ot the Royal Society of London, Se- ries B 324:1-110. Reid, D. G. 1989b. Systematic revision of the Recent species of Peasiella Nevill, 1885 (Gastropoda: Littorinidae), with notes on the fossil species. Nautilus 103:43-69. Reid, D. G. 1990a. A cladistic phylogeny of the genus Lit- torina (Gastropoda): implications for evolution of repro- ductive strategies and tor classification. Hvdroliiologia 193: 1-19. Held, I) (;. 19901) Trans-Arctic migration and speciation intluced by climatic change: the biogeograph\ of Littorina (Mollusca: (Jastropoda). ISulletin of Marine Science 4735- 49 Reid, D. G., N. I. Zaslavskaya, and S. (). Sergievsky. 1991. Littorina kasatka. a new species from the Kurile Islands and Okhot.sk Sea. Nautilus 105(1): I -6 Sacchi, G. F. and M. Rastelli. 1966. Littorina niariar. nov. sp. : les differences morphologiques et ecologiques. enire 'nains'et 'normau.x'chezr"espece"L.oi)fH,va(a (L.) ((Jastr. Prosobr. ) et leur signification adaptive et evolutive. Mti della Societa Italiana di Scienze Natural! e del Museo Givi- co di Storia Naturale, Milano 105:351-370. Seeley, R. H. 1986. Intense natural selection caused a rapid morphological transition in a living marine snail. Pro- ceedings of the National Academv of Sciences of the USA 83:6897-6901. Ward. R. D. 1990. Biochemical genetic variation in the genus Littorina (Prosobranchia: Mollusca). Hydrobiologia 193: 53-69. THE NAUTILUS 105(l):16-25, 1991 Page 16 Activity Patterns and Homing of Acanthopleura gemmata (Blainville, 1825) (Mollusca: Polyplacophora) in the Rocky Intertidal of the Jordan Gulf of Aqaba Neil C. Hulings' Marine Science Station PO Box 195 Aqaba, Jordan ABSTRACT Acanthopleura gemmata forages and homes irrespective of the time of da\\ Feeding excursions begin when the water depth of the ebbing tide over the individual is around 10 cm or less and homing occurs prior to the level of the flooding tide reach- ing about the same depth. The patterns of behavior differ significantly from those of A. gemmata previously reported at Heron Island, Australia and in Somalia. Reduced predation pressure is considered to be one of the factors accounting for this difference in behavior Key words: Chiton; ecology; predation; Red Sea. INTRODUCTION Ferreira (1986) revised the genus Acanthopleura Guil- ding, 1829 to include Acanthozostera Iredale and Hall, 1926. The revision also included placement of Acantho- pleura haddoni Winckworth, 1927 and A. spiniger (Sow- erby, 1840) into the synonymy of A. gemmata (Blain- ville, 1825). The material examined by Ferreira (1986) included specimens from Israel, Egypt, Djibouti, Oman, Somalia, Heron Island, Australia as well as numerous other localities in the Indo-Pacific region. Of the inter- tidal chitons, A. gemmata has the widest geographic distribution, from 32°E to 140°W and 33'N to 6°48'S (Ferreira, 1986). The synonymy and definition of A. gemmata proposed by Ferreira (1986) is followed in the present paper. Reports on the activity and homing of Acanthopleura gemmata include those of Thorne (1967, 1968 as Acan- thozostera gemmata) at Heron Island, Australia and Chelazzi et at. (1983a,b, 1987) and Chelazzi and Par- pagnoli (1987) along the coast of Somalia. Of the reports of A. gemmata (cither as A. haddoni or A. spiniger) in the Red Sea, including the Gulf of Aqaba (Sykes, 1907; ' Present address: Ranunkelgatan 7D, 431 32 Miilndal, Swe- den. Leioup, 1937, 1960, 1980; Rees & Stuckey, 1952; Safriel & Lipkin, 1964; Eibschiitz e/ ai. 1967; Fishelson, 1971; Mergner & Schuhmacher, 1974; Mastaller, 1978, 1979; Pearse, 1978, 1983; Ayal & Safriel, 1980; Soliman & Iskander, 1982; Hulings, 1986, 1987; Al-Hajj, 1987), none contain information on foraging or other patterns of ac- tivity, nor on homing behavior. This paper describes the activity patterns and homing of Acanthopleura gemmata (figure 1), the only chiton in the rocky intertidal of Jordan. The vertical distribution of this chiton ranges from the mid-midlittoral to the midlittoral-infralittoral fringe contact. The northern Gulf of Aqaba is the northernmost geographic occurrence of A. gemmata according to Ferreira (1986). Voucher spec- imens of A. gemmata have been deposited in the Na- tional Museum of Natural Historv Smithsonian Institu- tion (USNM 858104) and in the reference collection of the Marine Science Station, Aqaba, Jordan. STUDY AREA The characteristics of the rocky intertidal zone along the coast of Jordan have been described bv Hulings (1986, 1987). Briefly, the tides are mixed, having a spring range of around 1.0 m and a neap range of 0.5 m. The diurnal inequality of the high tides averages 4.2 cm, that of the low tides 4.7 cm. The tide levels are often influenced by atmospheric pressure (L'ziel, 1968; Hulings, 1989). The tide levels also fluctuate w ith changes in sea level of up to 1 m that occur in the northern Gulf. During the period July through October, sea level is higher while from December through May, it is lower; the intervening pe- riods are transitional (Uziel, 1968; Fishelson, 1973; Hul- ings, 1989). Although the vertical ranges and variations in the tide are small, the prevailing near horizontal sub- strata of the midlittoral zone are subjected to extended submergence during the periods of higher sea le\el and emergence during periods of lower sea level (Hulings, 1986). The climatic conditions in the northern Gulf of Aqaba are very significantK influenced bv the much greater N. C. Hulings, 1991 Page 17 -•- 3 Figure 1. Acantho])lcura gemmata from the rock\ midlittnral zone of Jordan, northern Gulf of Aqaba, (f.O x ). desert land area surrounding the very limited marine area. The land to sea winds are northerly 93*^' of the time with Beauforts 3 (7 to 10 kn) and 4 (11 to 16 kn) occurring 63% of the time. Mean air temperatures range from 16 °C in January to 32 °C in July-August; minimum- ma.ximum means ranges are 12 to 20 °C in January' and 26 and 37 °C in July-August (Jordan Meteorological De- partment). Anati (1976, 1980) estimated evaporation in the Gulf to be 4.2 m/yr. The area is further characterized by strong solar insolation (Assaf & Kessler, 1976; Leva- non-Spainer et al.. 1979). The locality (figure 2) investigated in detail for the activity of Acanthopleura gemmata is located at the Marine Science Station (34°.59'20"E, 29°29'30"N), ap- pro.ximately 10 km south of the city of Aqaba. The mid- littoral zone is characterized by two types of substrata, landward beachrock and below, a platform of near hor- izontal calcareous fossil reef matrix. The beachrock area is biologically recognizable b>' the presence of the giant barnacle Tetraclita squamosa rufotincta Pilsbry, 1916, which colonizes the surface of the beachrock. This area is the Tetraclita zone of Safriel and Lipkin (1964) and is considered herein as the middle portion of the mid- littoral. The zone averages 5 m in width and consists of discontinuous strips of beachrock, underlain by fossil reef, parallel to the shore; between the strips are depressions and tide pools as well as numerous and varied refuges. The surface configuration of the fossil reef in the area of the beachrock is ver\ irregular. In addition, the un- derlying reef substrate is hard compared to the reef ma- trix of the platform. The platform of nearly horizontal, calcareous reef ma- trix is 10 to 18 m wide, extending from the seaward edge of the beachrock to the midlittoral-infralittoral fringe (Safriel & Lipkin, 1964) contact and is also referred to herein as the lower midlittoral. The surface topography of the platform is mostly even and uniform. In the area iS^ Figure 2. A section oi tlie stud\ area tiuruig ebbing tide. The arrows indicate: 1. .■\pproxiniatesupraiittoral fringe-midlitorrai contact zone; 2. The Tetraclita or mid-niidlittorai zone; 3. lower midlittoral-infralittoral fringe contact; 4. outer edge of the fringmg reef, See text for details. of the midlittoral-infralittoral fringe contact, however, the substratum is eroded, providing refuges such as over- hangs, crevices and pits. Beyond the contact is a 35 to 85 m wide back reef lagoon of a fringing reef. The lagoon, having a maximum depth of around 2 m de- pending on the tide and sea level, terminates at the outer edge of the reef. It is here that wind-generated waves break and their energ\ is further reduced across the lagoon. Thus the intertidal area has minimal exposure to wave action. During high tides, however, wind-induced choppy conditions may prevail, but the wave amplitude is small because of the shallow depth. .•\t the localitx' in\estigated, the mean length of Acan- thopleura gemmata is 53.0 mm (SD = ±8.6 mm; range, 30 to 75 mm; n = 160) and the mean density is 4.4 individuals/ m- (SD = ±3.1; n = 80 m-). According to Hulings (1986), A. gemmata becomes sexualK' mature at a length of between 24 and 27 mm. METHODS AND MATERIALS The data on the activity and homing of Acanthopleura gemmata were obtained entireK from detailed obser- vations made during the period 14 October through 16 December 1986. Observations were made on groups of A. gemmata within a 100 m stretch parallel to the beach and 10 to IS ni wide. The area included the midlittoral range of the vertical distribution of A. gemmata as well as its occurrence in beachrock areas, on fossil reef capped by beachrock and on the platform below the beachrock. The shell plates and the home area or scar of Acan- thopleura gemmata were color-coded using different colors of enamel paint. Subsequent repainting of the plates was often necessary and there was no detectable negative effect of the painting. Forty-five individuals comprising eight diflerent groups, each with three to eight individuals, were color-coded One group was on beachrock (six individuals), three on fossil reef capped by beachrock (17 indi\iduals) and four (22 individuals) Page 18 THE NAUTILUS, Vol. 105, No. 1 on the platform. During the study perioii, one specimen was lost at the beachrock site, two from the beachroek- fossil reef substrata and three from the platform area. Thus 39 individuals, wliich were present during the en- tire two month period of investigation, were used in the analyses of activity and behavior. Various combinations of groups and or indi\ iduals were used for the analysis of specific t\ pes of acti\ it\ and l)ehavior. During the October-December period, daytime ob- servations were made on 46 days. The total number of days included an initial two consecutive days of obser- vations follow ed by eight weeks of five consecutive days per week and a hnal four consecutive days. On each day, each individual within the eight groups was observed one or more times (all groups observed each time being a set of observations). The number of observations on each group averaged 5/day (range, 1 to 11 /day). Over the 46 days, the number of sets of observations per group ranged from 139 to 161 with the total numb)er being 5,712. Nighttime observations were made on two groups totaling eight individuals during October (2), November (2) and December (1). There were two sets of observa- tions each night, a total of 80 sets or 320 observations. Six additional groups of six to 11 individuals, a total of 53, within the area w ere checked once a day during each five consecutive days of observations. In addition, peri- odic observations were made during January, February, June and August 1987 and February and June 1988. The data recorded during each observation of each color-coded individual included the position in relation to the color-coded home scar. If the individual was in the home scar, it was so noted as well as whether there was a reversal in the orientation. If the individual was away from the scar, the distance in cm from the home scar and the direction was noted. Occasionally, actual movement of an individual was observed. In most cases, however, movement was inferred based on the position of the individual. The depth of water over the individual was measure in approximate cm intervals. If the indi- vidual was partially to completely submerged dorso- ventrally, it was designated as awash. In the absence of water cover, the substratum was recorded as wet if water was visible or dry if no water was visible. The number of sets of observations made in relation to the state of the tide during the daytime over the two months were as follows: eight at flood tide, 39 at flooding tide, 27 at ebb tide, 18 at ebbing tide and six at ebbing to ebb tide. During the period, there were 19 days of spring tide, 10 of transitional spring to neap, 10 of neap and seven of transitional neap to spring. During night- time observations, the tide was at flood on one occasion, at ebb on another, ebbing on a third and flooding on two others. During two of the nights, there were spring tides, during one neap and during two transitional spring to neap. Two additional groups of Acantlwplcura gcmniata were monitored for vertical migration with changes in sea level from low during October (10 days) to high during December ( 1 1 days) with November (22 days) being transitional. One group of 12 individuals was in the Tr/rflr/i7a -beachrock area of the midlittoral and the other group, 11 individuals, was near the midlittoral- infralittoral triiige contact. During the October-Decem- ber period, a total of 91 observations during 46 days were made on the Te^rar/i7a-beaclirock group and 24 obser- vations during 16 days on the midlittoral-infralittoral fringe group. Subsequent observations on each group were conducted during January (2 days), February (1), June (1) and August (1) 1987 and February (1), June (1), August (1) and September (1) 1988. These observations included four days of higher sea level and five of lower. The direction of movement from the home scar during foraging forays was measured in 17 color-coded Acan- thopleura gemmata in the beachrock area and on the platform over a period of 2 to 29 days. The range of foraging (in cm) was measured among the same indi- viduals over the same period. The range was considered to be the distance from the home scar to the most distant point within the individual grazing area, the latter iden- tifiable by the color being lighter than the surrounding area. RESULTS Activity, in terms of movement, of Acanthopleura gem- mata from mid-October to mid-December 1986 was found to be dependent on the depth of water over the individual (table 1). Based on 19 color-coded individuals in four groups on the platform and five in one group in the beachrock area, when there was movement, it oc- curred predominantly from when the substratum was wet to when the water depth of the tide over the indi- vidual was 10 cm or less irrespective of the time of day. During the day, 88% of the instances of movement oc- curred within the wet to 10 cm range, while 90% of nocturnal movement occurred within this range. A sim- ilar pattern prevailed in relation to the t\pe of tide day and night (table 2). However, movement did not always occur during the wet to 10 cm depth interval (tables 1, 2). Among the same 19 color-coded individuals, day and night movement in relation to the type of tide (table 2) was as follows: 34% of the instances of movement oc- curred during spring tide, 38% during transitional spring to neap, 16% during neap and 12% during transitional neap to spring. It is recognized that the number of observations at night was very limited. However, additional evidence tor nocturnal activity is derived from comparing the position of an individual at the end of one day with that at the beginning of the next. Among five groups of 24 color-coded individuals, a total of 318 observations re- vealed 113 (36%) instances of positions different from that of the last observation, usually late afternoon, on the previous day. Among the total different positions, 34 (30%) were in the home scar at the beginning of the second day. By comparison, 193 (94%) of the 205 in- stances of the position being the same were in the home scar at the beginning of the second day. N. C. Hulings, 1991 Page 19 Table 1. Movement of Acaiitliupleuru geiuiuata in relation to water depth during day and night (u = number of obser- \atioiis; — = no obser\ation; A = awash). Table 3. Home (■.s. awa\ position of Aamlhoplcuru gcmmata in relation to water depth (n = number ol observations) Water Movement No iiovement depth Da\ N'ighl Day N'ight (cm) n n n n Dry Wet 0 97 15 26 264 54 A-5 172 43 171 49 5-10 137 48 128 60 10-15 45 12 104 39 15-20 8 — 102 0 20-25 0 — 48 — 25-30 0 — 32 — 30-35 0 — 10 — 35-40 0 — 5 — Total 459 118 890 202 The home vs. away position of one group of five in- dividuals with respect to water deptli during the day over 41 days is show n in table 3. In 70'^o oi the instances, the individuals were at home over a range of conditions from dry substratum to a water depth of 50 cm, while 85% of the away instances were during the wet to around 10 cm water level period. The movement of Acanthoplcura gcmmata with the ebbing tide was either rotation within the scar or the initiation of a feeding excursion. The latter generally occurred from when the water depth over the individual was around 10 cm until the time the individual was completely exposed. Following exposure, the animal ei- ther returned to the homing scar before complete drying of the substratum or became inactive and remained in situ. There was no movement when the substratum was dry (table 1). Movement of the in situ individual started upon wetting by the flooding tide and the individual returned to the homing scar immediately before the wa- ter depth of the flooding tide reached around 10 cm over Water depth (cm) Home n Away n Dry Wet A-5 5-10 10-15 15-20 20-25 25-30 30-35 35-40 50 Total 10 0 96 46 42 95 85 45 74 21 75 4 46 3 43 3 25 1 19 1 5 0 20 219 the scar. In a few cases, return occurred at a water depth of up to 20 cm (table 1). Fecal pellets similar to those described by Taylor and Way (1976) were released dur- ing the period of occupation of the homing scar. The general pattern of activity of A. gemmata with the ebb- ing and flooding tide, based on a total of 4,445 obser- vations of 24 individuals during the October-December period, is summarized in table 4. Other responses to the flooding tide among 39 chitons based on 2,769 observations were as follows: no move- ment at all was detected (62'^.); movement restricted to one or more 1S0° rotations witliin the homing scar (15%); short-term excursion, usually not exceeding 5 cm (23%). In addition, 124 observations on 24 individuals during 25 days revealed no long excursions during subsequent tide cycles within the same day in 41 cases (33%). The direction of movement on long excursions from the homing scar over a period of 2 to 29 days among 17 individuals was within an average angle of 177° (SD = ±98°; n = 179) and ranged from 45 to 360° The distance of the excursions of the same individuals within the graz- Table 2. Movement of Acanthupleura gcmmata ni relation to water depth and type type of tide day and night (in parentheses). S = spring tide; SN = transitional spring to neap; N = neap; NS = transitional neap to spring; n = number of observations; — = no observation; A = awash. Water Mo\ ement No mo\ ement depth S SN N N8 S SN N NS (cm) n n n n n n n n Dry O(-) O(-) O(-) 0(- 26(— ) O(-) O(-) O(-) Wet 51(0) 27(12) 12(0) 7(— 184 (0) 59(21) 1(0) 20(— ) A-5 53(14) 58(16) 38(17) 23(— 72(19) 61(17) 21(16) 17(-) 5-10 39 (20) 60(14) 7(20) 31(- 48(14) 42(19) 17(14) 21(-) 10-15 12(4) 29(1) 0(0) 4(- 25 (25) 62 (28) 10(0) 7(-) 15-20 3(0) 2(0) 2(0) 2(- 21 (29) 26(0) 23(0) 32(— ) 20-25 O(-) O(-) O(-) 0(- 2(-) 13(-) 14(-) 19(-) 25-30 O(-) O(-) O(-) 0(- O(-) 5(-) 7(-) 20(— ) 30-35 O(-) 0(— ) O(-) 0(- l(-) l(-) l(-) 7( — ) 35-40 O(-) 158(38) O(-) 176(43) O(-) 58 (37) 0(- l(-) 380(87) l(-) 270(85) 2(-) l(-) Total 67 (— 96 (30) 144 (-) Page 20 THE NAUTILUS, Vol. 105, No. 1 ing area averaged 21 cm (SD = ±15 cm; n = 179). On the near horizontal reef matrix, the average angle of the excursions was 149° (SD = ±106°; n = 100) and the mean distance 21 cm (SD = ±13 cm; n = 100) while on beachrock and associated substrata, 233° (SD = ±72°; n = 79) and 23 cm (SD = ±17 cm; n = 79). Observations on 12 Acanthopleura gemmaia in the rc/rar/ifa -beachrock area and 11 near the mitllittoral- infralittoral fringe contact revealed no vertical migration from the period of lower sea level in October through the transitional period during November to higher sea level in December. The absence of vertical migration was subsequentK confirmed by four observations during high sea level and five during low sea level in 1987 and 1988. This is contrary to the report of vertical migration of A. gemmaia in Hulings (19S7) which was based on a more limited number of observations. The investigation of the homing behavior of Acan- thopleura gemmata during mid-October to mid-Decem- Table 4. Activity of Acanthopleura gemmata in relation to the tide (n = number of observations). Activitv II '>i ber revealed that 79% (SD :205 = 1,561) of 39 color-coded individuals in eight groups exhibited definite homing behavior, i.e., returning to the same color-coded homing scar following foraging excursions. The obser- vations included day and night as well as during spring, transitional and neap tides. The homing behavior and foraging excursions occurred irrespective of the time of day and the type of tide. The remaining 21% (SD = ±20%; n = 466) occupied new sites for a variable period of one to several days. Observations on other groups totaling 53 individuals as well as periodic observations through June 1988 essentially confirmed these findings. Within groups in which the individuals were clumped or in close proximity, exchange of home scars was found to be common. The average number of homes occupied by each of 39 individuals over the two month period was 3.2 (SD = ±1.5; n = 2,027). The period of consecutive occupation of a home other than the original among 16 individuals (n = 242) varied from 1 to 19 days and averaged 5.8 days (SD = ±5.0 days). Among 39 indi- viduals, four occupied the original home scar for the entire two month period of observation while seven did not return to the original or new home scar. Among the remaining individuals, there were 20 cases in which there was immediate or eventual return to the original home scar and 49 in which the individual moved to one or more home scars and returned immediately or eventu- ally. Within one group of five individuals, there was only one case of reciprocal swapping on a daily basis over the two month period (n = 760). In six cases, there was initial reciprosity followed by suKsequent continued occupation of up to five da\s. Within the same group and over the same time span, the total number of sites occupied was 47. There were 27 reoccupations of a site following oc- cupation by another individual. The most recognizable homing scar of Acanthoph'ura gemmata in the rockv intertidal of Jordan is a distinct depression that is concave in cross and longitudinal sec- tions. The scar is about 1 mm deep with the deepest area \o movement from home during ebbing tide nor during exposure. \lo\emenl from home with the ebb- ing tide and return home with the subsequent flooding tide. Movement from home with the ebb- ing tide and return home prior to the flooding tide. Movement of those previously away at ebb tide to home with the flood- ing tide 1.709 1,584 576 .5,(1 38 36 13 13 coinciding with that region occupied by the foot. The surface ot the scar is smooth compared to the surface of the surrounding substratum and has a lighter color than the surrounding substratum, v\ hich is covered with epi- growth. Though abandoned scars become overgrown, they remain recognizable. Such readily recognizable homing scars are found on the nearly horizontal, rela- tively soft, homogenous, fine-grained calcium carbonate fossil reef matrix, a dominant rocky intertidal substratum along the coast of Jordan. The scars are also found on the sides of refuges such as depressions, crevices and small caves. The scars of A. gemmaia lack the algae occurring in the homing scars of the patellid limpet Cellana radiata (Born, 1778) and the pulmonale Siphonaria laciniosa (Linnaeus, 1758) reported by Hulings (1985). Other less recognizable scars may be identified on the basis of substrate color. Apart from this difference, the surface configuration of the homing area is not noticably different from that of the surrounding substratum. These scars are also found on calcium carbonate substrata, one type of which underlies beachrock and is harder than the matrix noted above. Another type of substratum with such scars includes fossil coral heads. Recognizable hom- ing scars have not been detected on substrata such as granitic and similar rock and beachrock although homing on these substrata has been observed. Endean et at. (1956) noted the absence of excavations by Aeanthoph^itra gem- mata in hard rock on the coast of Australia. The orientation of both homing scar and chiton varies irrespective of the substratum. The scar may be near to horizontal on such substrata or on the sides of refuges. In the refuges, the anterior-posterior axis of the chiton may be parallel to the horizontal or inclined to vertical with the anterior end up. Semi-quantitative data (Hul- ings, in preparation) indicates that the density of Acan- thopleura gemmata in areas with refuges is 4.1 m- (SD = ±3.2/m-; n = 183) while that in areas without refuges is 5.5/m- (SD = ±5.4/nr; n = 79). Among 37 chitons and based on 2,027 observations, the mean percent homing of 22 indiv iduals on relatively .soft, nearly horizontal reef matrix v\ here the majority of the readily recognizable homing scars occur was 85% (SD = ±14%). B\ comparison, the mean homing of 15 N. C. Hulings. 1991 Page 21 individuals on hard substrata such as beachrock and the underlying substrata was 69% (SD = ±24%). The dif- ference between the means is statistically significant at P = 0.01 (n — 2 = 35). While the mean number of homes occupied by the individuals on soft and hard substrata was almost the same, 3.4 (SD = ±1.4) vs. 3.1 (SD = ±1.6), there were eight instances among the individuals on soft substrata of return to the original home vs. 12 among those on hard substrata. In the case of movement to a different home and return, there were 34 instances on soft substrata vs. 10 on hard. DISCUSSION There are limitations that must be imposed on the in- terpretation of the data obtained during the investigation of the acti\it\ patterns and homing of Acanthopleura gemmala. First, the data are based entireK on obser- vations; no e.xperimental manipulations were conducted although the value of such is recognized. Second, statis- tical treatment is limited, of a general nature and is used for indicative purposes only. FinalK , the data appK' onh- to adult indi\ iduals. Other limitations are indicated be- low The homing behavior of Acanthopleura gemmata in the northern Gulf of .\qaba is similar to that reported for this species by Thorne (1967, 1968) at Heron Island, Australia, and by Chelazzi et al. ( 1983a, b, 1987) and Chelazzi and Parpagnoli (1987) on the coast of Somalia. The homing of 79% of the individuals recorded herein is the same as reported by Thorne (1967) while Chelazzi et al. (1983b:17) reported a 91%' return to the home of the "previous owner." In addition, the same percentage was found for individuals occupying new homes at Heron Island and on the coast of Jordan. Chelazzi et al. (1983a) found periodic movement to a new home a common behavioral pattern as did Throne (1967) and found here- in. In terms of reciprocal swapping of homes, the low- incidence found herein is in agreement with that found by Thorne (1967). The 57% reoccupation of a home following occupation b}' another individual is higher than the 42% reported by Chelazzi et al. (1983b). The orientation in the homing scar, including 180° rotation, is essentialK the same in the three geographic areas. There is also considerable similarit\ in the mor- phology and appearance of the homing scar. The oc- cupation of scars in exposed (platform) sites as well as in refuges (pits, crevices, overhangs) is a common be- havior (Thorne, 1967; Chelazzi et al.. 1983b). Chelazzi et al. (1983b) reported aggressive behavior between a chiton returning to its home scar and an in- truder occupying the same homing scar. The returning chiton attempts to oust the intruder, but, if unsuccessful, the owner either mo\es elsew here or the two individuals occupy the same scar. In the present study, aggressive behavior was not observed, but the co-occupation of a single scar by two individuals, side by side, was observed on several occasions. Onl\ a few periods of observations during one day included a combination of a long enough time span, suitable water level conditions and movement by Acan- thopleura gemmata to indicate the return to the home scar following grazing. During such a combination, 43 observations on eight indi\ iduals suggests return by the same route as that of the excursion. This is consistent with the reports of Thorne (1967) and Chelazzi et al. (1983b, 1987). Thorne (1967) suggested chemosensitivit\ and topo- graphic memory as potential mechanisms whereby Acanthopleura gemmata returns to the homing scar. This author (Thorne, 1968) later added celestial navi- gation, sight, random movements, olfactory clues, to- pographic memory and detection of previous trails as possible mechanisms. For a \ariety of reasons, he ruled out the first three and, after subjecting the last three to experimental treatment, considered only topographic memory and detection of trails as possible homing mech- anisms in A. g£>mma?a. Eibschiitz p< a/. (1967) discounted the magnetic properties of the radula of A. gemmata at Eilat as a factor in navigation because of the low level of residual magnetism. Of note is the occurrence of homing even when the scar and the surrounding area was completely covered with sand during periods of sedimentation, or with shell debris follow ing a storm. The frequenc\ of homing by individuals with well-de\ eloped scars was significantK' higher, suggesting that a chemical trail may be involved in the homing of A. gemmata. as indicated by Chelazzi and Parpagnoli (1987). However, reciprocal swapping of scars and the reoccupation of sites following occupation by one or more indi\iduals suggests that there is not a trail specific for each individual but one common to all individuals of the species. The mean length of the feeding excursions of Acan- thopleura gemmata in this stud\' was 21 cm. By com- parison, Chelazzi et al. (1983b) reported 30 cm and Thorne (1967, 1968) 47 cm. Thorne (1967, 1968) found that not all individuals forage every night and Chelazzi et al. (1983b:16) noted that nocturnal activity depended "on the position in the spring-neap cycle." In the present study, not all individuals w ere found to be active during the conditions under which foraging occurs (tables 1, 4) nor during consecutive tide cycles. Chelazzi et al. (1983a) found the feeding excursions of Acanthopleura gemmata to be the longest between neap and spring tides and minimal around neap tides. In the present study, the length of feeding excursions in relation to the t\ pe of tide was not determined. However, the number of instances of movement in relation to the type of tide was highest during transitional spring to neap tides followed by spring, neap and transitional neap to spring tides (table 2). Based on the assumption that mo\ements during ebbing conditions represent foraging excursions, an analysis of one group of five individuals on 26 days (n = 157) revealed 41% of the instances of movement occurred during ebbing tides. Of this per- centage, 43% of the instances occurred during transi- Page 22 THE NAUTILUS, Vol. 105, No. 1 tional spring to neap tides, 32% during spring, 13% dur- ing neap and 5% during transitional neap to spring. Movement of Acarithopleura gemtnata in the north- ern Cult of .\qaba is more directl) related to \\ ater depth over the indi\iduai than to the t\pe of tide. During periods of higher sea level or portions of the neap tide cycle, the water level over an individual may not drop to 10 cm or less, especially in the lower midlittoral zone. Thus feeding excursions may not occur during one or more consecutive days. The fewer instances of movement during neap and neap to spring tides over that of spring and spring to neap (table 2) may also be indicative of reduced activit\'. Many Acanthopleura gemmata in the northern Gulf have a well-defined grazing area, recognizable by dif- ferences in color from that of the surrounding area. This area is used repeatedly (Chelazzi ei al., 1983a, b, 1987), but appears to be exclusive, unlike that reported by Che- lazzi et al. {1983b). The feeding areas are more dispersed among groups of individuals than the homing scars as also found by Chelazzi et al. (1983a). Grazing appears to be most common at the outer limits of the area, as also reported by Thorne (1967). On the coast of Jordan, the range of the feeding ex- cursions of Acanthopleura gemmata was found to differ with the surface topology of the substratum. On the more regular surface of the near horizontal platform of cal- careous reef matrix, the range averaged 149° while on the very irregular surface of the beachrock and under- lying substrata, the average range was 233°. The differ- ence in the range may reflect the amount of colonization, distribution and abundance of algae as well as recolon- ization following grazing on the two different substrata. It would appear that the conditions for recolonization are more favorable on the soft, porous reef matrix due to the nature of the substratum and decreased tidal ex- posure than on the more exposed hard, impervious beachrock and associated substrata. As a result, more algae are more evenly distributed more of the time on the platform. Accordingly, a greater range of foraging would be necessary on the beachrock and associated sub- strata in order to obtain the necessary food requirements. The mean length of A. gemmata along an exposure gra- dient within the vertical distribution may also reflect food availability as well as a response to exposure (Hulings, in preparation). In the mid-midlittoral beachrock area, the most tidally exposed within the range, the mean length was found to be 47.0 mm (SD = ±7.6 mm; n = 60) while in the area of the midlittoral-infralittoral fringe contact of the platform, the most benign, the mean was 59.8 mm (SD = ±9.8 mm; n = 50). On the platform just below the beachrock, an area intermediate in ex- posure, the chitons had a mean length of 53.3 mm (SD = ±6.2 mm; n = 50). The difference between the means is statistically significant at P = 0.001. No aggressive behavior within the feeding area of Acanthopleura gemmata has been ob-served. This is con- sistent with the report of Chelazzi et al. (1983b). Among the chitons on the Jordanian coast, there was no pre- vailing up- or downslope nor lateral direction of the feeding excursions. .\s a result of orientation and length excursion analyses of A. gemmata and the sympatric A. hrevLspinosa (Sowerby. 1840), Chelazzi et al. (1983a) and (Chelazzi and Parpagnoli (1987) found the prevailing di- rection of movement during the feeding excursions of A. gemmata to be downw ard and that of A. brevispino.sa upward. They considered this behavior to be an example of minimizing food competition and maximizing the uti- lization of algae in the midlittoral. There is a possible example of minimizing competition in the rocky intertidal of Jordan. Based on density and presence-absence data, the correlation between the spa- tial distribution of the limpet Cellana radiata and Acan- thopleura gemmata was found to be negative (Hulings, in preparation). The limpet is considered to be the major grazing competitor of the chiton, especialK in the lower midlittoral, where the two are the numericalh dominant intertidal grazers. However, the conditions under which each forages differs. Cellana radiata forages only when submerged, becoming active soon after submergence by the flooding tide to just before or just after emergence (Hulings, 1985). Acanthopleura gemmata forages with the ebbing tide. The timing and the conditions under which feeding excursions of Acanthopleura gemmata occur at Heron Island, Australia and on the coast of Somalia differ very significantly from those in the northern Gulf of Aqaba. Thorne (1967, 1968), Chelazzi et al. (1983a,b, 1987) and Chelazzi and Parpagnoli (1987) found A. gemmata to be active only during nocturnal low tides and not during the day, whether exposed by the tides or not. Chelazzi et al. (1983b) reported that A. gemmata remains in the homing scar during the following high tide and diurnal low tide. By comparison, Acanthopleura gemmata in the north- ern Gulf has been found to be active during day and night. Foraging excursions commence when the water depth of the ebbing tide over the individual is around 10 cm or less. Return to the homing scar occurs prior to or with the flooding tide; in the latter case, before the water level is around 10 cm over the individual (table 4). The mechanism(s) triggering the initiation of feeding excursions and homing in Acanthopleura gemmata in the northern Gulf is not known. Other investigators have suggested various mechanisms, including a biological clock (Thorne, 1967) and the release of chemical infor- mation during high tide (Chelazzi & Parpagnoli, 1987). Though not investigated, the presence of photoreceptors such as aesthetes and ocelli in the shell plates and the possibilitv of the girdle being photosensitive (Bo\le, 1977) does not appear to be involved in the triggering of move- ment in A. gemmata as it occurred da\ and night. Of particular interest is the finding of Moulton (1962) of the initiation of movement aw ay from clusters of the cerith- iid Chjpeomorus monilijcrum Kiener, 1841 at Heron Island when the water depth reaches 10 cm (the same species occurs in the rock\ intertidal of Jordan according N. C. Hillings, 1991 Page 23 to Hillings, 1989), Moulton (1962) suggested a hydrostatic mechanism and, behaviorally, positive barokinesis and thigmotaxis as being involved in clustering and dispersion of the cerithiid. Based on laboratory experiments, Rohde and Sandland (1975) discounted a hydrostatic mecha- nism as the onl) factor in dispersion and clustering of C. moniliferum. It is suggested that a difference in predation pressure on Acanthoplciiro gemmata between the Heron Island- Somalia region and the northern Gulf of Aqaba may account, in part, for the nocturnal behavior in the former region vs. the diurnal-nocturnal behavior in the latter area. Both Thorne (1967) and Chelazzi et al. (1983a,b) considered homing in A. gemmata to be of significance in the reduction of and protection against predation. Among the predators listed by Thorne (1967, 1968) are the grapsoid Grapsus strigosus (Herbst, 1799) which is synon\mous with G. albolineatiis Lamarck, 1818 ac- cording to Banerjee (1960); the xanthids Eriphia leavi- mana Latrielle, 1817 and Leptodius exaratus (H. Milne Edwards, 1834); the anomuran Clibanarius virescens (Krauss, 1834); birds including herons, waders and gulls and elasmobranchs, Chelazzi el al. (1983b) reported the xanthid crabs Eriphia smilhi McLear\, 1838 and Ozius guttatus H. Milne Edwards, 1834 as less important pred- ators and the tetradontid fish Arothron immaculatiis (Block and Schneider, 1801) as the most important pred- ator (also see Chelazzi et al., 1987; Chelazzi & Parpagnoli 1987). In the northern Gulf of Aqaba, the grapsoid Grapsus granulosus H. Milne Edwards, 1853 is a common mid- midlittoral slab occupant while G. alholineatus most commonly inhabits boulders (Hulings, 1989). Of the above xanthid crabs, onlv Leptodius exaratus has been record- ed in the Jordanian Gulf (D. Guinot, personal commu- nication). Other xanthids including Lijhia leptochilis (Zehntner, 1894) and Polijdectus cupulifcr (Latrielle, 1825) have been found in the rocky "eulittoral' b\ Ma- staller (1979). The anomuran Clibanarius signatus Hel- ler, 1861 is a mid-midlittoral species on slab along the coast of Jordan (Hulings, 1989). Among the fishes, Arothron immaculatus has not been reported from the Gulf of Aqaba (Dor, 1984). However, other tetradontids have been reported in the Gulf at Eilat, including Arothron hispidus (Linnaeus, 1758) by Ben- Tuvia and Steinitz (1952) and Tortonese (1968) and a questionable Tetradon aff. diadcmata Riippell, 1829 by Steinitz and Ben-Tuvia (1955). There is. however, no information on the habitat occurrences of these tetra- dontids. Among the species of fish iinading the rocky intertidal during high tide according to Fishelson (1977), potential predators on Acanthopteura gemmata include the labrid Cheilio inermis (Forsskal, 1775) and the ba- listid Sufflan}en albicaudatus (Riippell, 1829) (R. Or- mond, personal communication). As for marine birds, there is not a resident population of species in the north- ern Gulf. However, the area is on the migratory route of gulls, herons and waders (Safriel, 1968). It is inferred that the predation pressure on Acantho- pleura gemmata in the northern Gulf of Aqaba is con- siderably less than at Heron Island and in Somalia. The absence of a resident marine bird population as well as rare occurrences during migratory periods indicates the absence of a diurnal predator that would feed while A. gemmata is exposed and active at low tide. The stomach contents of the grapsoids Grapsus albolineatus and G. granulosus include macerated fragments of algae, hy- droids and detrital particles but no chitons (Hulings, un- published data). In addition, the structure of the chelae of the two grapsoids (Schiifer, 1954) is not suitable for dislodging or preying upon A. gemmata. The hermit crab Clibanarius signatus in the rock> intertidal of Jor- dan is small (maximum length around 2 cm) as are the xanthid crabs and are thus considered to be of minor, if any, importance as predators. Evidence lor fishes as pred- ators is lacking, including the absence of "scars" on the plates of A. gemmata similar to those caused by Arothron immaculatus in Somalia as reported bv Chelazzi et al. (1983b). The homing scars of Acanthopleura gemmata along the coast of Jordan are only a mm or so deep as opposed to those on the coast of Somalia being 1 to 10 cm deep (Chelazzi et al., 1983b). The shallow scars may, in turn, reflect the absence of one or more significant predators during high tide when compared to the situation in So- malia. In the latter case, Chelazzi et al. (1983b) consid- ered that the homing behavior of A. gemmata protects it from the most important predator, Arothron immac- ulatus, during high tide when predation pressure is the highest. It is concluded, therefore, that the diurnal-nocturnal pattern of behavior in Acanthopleura gemmata in the northern Gulf of Aqaba is a result of the reduced ter- restrial and submarine predation pressure. That periodicity of acti\ity can change with predation pressure has been demonstrated b\ Fricke (1974), who found that locomotor and feeding activity of the sea urchin Diadema setosum (Leske, 1778) in the northern Gulf of Aqaba was nocturnal in the presence of predatory fishes. In another area, where the predators w ere absent, the urchins were active irrespective of the time of day. Fricke (1974) hypothesized the evolution of the day- night activity as an adaptation in a habitat free of pred- ators. The timing of the acti\it\ patterns in relation to water depth over Acanthopleura gemnuita in the northern Gulf does, however, suggest a response to predation. The be- ginning of the feeding excursions at around the 10 cm water depth of the ebbing tide, and homing when the depth of the flooding tide reaches 10 cm, as well as the lack of movement during periods when water depth ex- ceeds 10 cm (table 1) indicates an adaptation whereby movement is minimal at times when it might attract submarine predators. In most of the cases where the water depth over the individual exceeded 10 cm, the chiton was in the home scar (table 3). By occupying a concave scar on the exposed, soft, horizontal slab, the chiton blends in with the surface Page 24 THE NAUTILUS, Vol. 105, No. 1 topograpliy and coloration of the surrounding .substra- tum. Additional caniouOage results from the spreading of the girdle around the internal periphery of the scar. The latter contrasts to the form of the bod\ during ex- posure at the low tide. During exposure, the body is raised and arched longitudinalK- as a result of what appears to be contraction of the girdle. Chitons homing in the hard substrata of the beachrock areas most commonly occur under overhangs or in crevices or caves rather than on the surface of the substrata. The response to submarine predation among other intertidal homing species in the northern Gulf, Cellana radiata and Sipbonaria laciniosa. is different. Both for- age and home only when submerged and the activity may continue through maximum flood tide. However, C. radiata is active day and night whereas S. laciniosa is nocturnal and neither is active when exposed during low tide (Hulings, 1985). These patterns may also be a response to predation pressure. In addition to protection against predation, homing in Acanthopleiira gemmata has been attributed to protec- tion against environmental stresses (Chelazzi et al., 1983a, b, 1987). Certainly, the nocturnal foraging and homing of A. gemmata at Heron Island (Thorne, 1967, 1968) and on the coast of Somalia (Chelazzi et al., 1983a, b, 1987; Chelazzi and Parpagnoli, 1987) reduces exposure to environmental extremes. However, in the case of the rocky intertidal in the northern Gulf of Aqaba, which is subjected to severe climatic and environmental extremes, the foraging and homing activity of A. gemmata is tide- dependent and independent of the time of day or year. And while other intertidal species migrate vertically with changes in sea level in the northern Gulf, a behavior considered, in part, to be an avoidance of environmental extremes (Hulings, 1987), A. gemmata does not. As a result, the chiton is active during periods of environ- mental extremes, including the combined warmer-lower sea level period, as well as during benign periods. As noted above, homing following feeding excursions dur- ing periods of maximum exposure to climatic extremes does not always occur. Thus the homing of A. gemmata in the northern Gulf as a response to protection against environmental stresses appears to be of less significance than elsewhere. It is recognized that differences in the behavior pat- terns of Acanthopleura gemmata between the northern Gulf of Aqaba and Somalia-Heron Island may be due to differences in physical conditions. The tides are of the same type in all three areas, although the range is greater than in Aqaba. Other conditions, especially the geomor- phology, appears to be similar, based on information given in Endean et al. (1956), Vannini et al. (1977) and Chelazzi and Varmini (1980). Geographic differences in activity- patterns and behavior among the same rocky intertidal species occurring in the northern Gulf of Aqaba and in other geographic areas outside the Red Sea, in- cluding Heron Island, have been noted by Hulings (1989). Differences exist even within the Red Sea. The repro- ductive periodicity of Acanthopleura gemmata in the northern (Julf of Aqaba (Hulings, 1986) is similar to that in the Gulf of Suez (Pearse, 1978; Soliman & Iskander, 1982), while both differ from that in the Red Sea proper (Pearse, 1978). ACKNOWLEDGEMENTS The author wishes to thank Mr Burhaii Gharaibeh, Huw wara, Jordan and Mr. Percy Linder and Mr. Jonne Lysell, Kallered, Sweden for assistance. The constructive criticism of two anonymous reviewers is greatfully ac- knowledged. LITERATURE CITED Al-Hajj, H. A. 1987. L^ltrastructural stutU of spermiogenesis in the chiton Acanthopleura haddoni from the Jordan Gulf of .\qaba (Red Sea). 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Ecological and biological phenomena in- fluencing coral-species composition on the reef tables at Eilat (Gulf of Aqaba, Red Sea). Marine Biology 19:183- 196. Fishelson, L. 1977. Sociobiology of feeding behavior of coral fish along the coral reef of the Gulf of Flat (= Gulf of Aqaba), Red Sea. Israel Journal of Zoology 26:114-13-1. Fricke, H. W. 1974. Moglicher Einfluss von Feinden auf das Verhalten \ on Didc/cma-Seeigelen Marine Biologv 27:59- 62. Hulings, N C. 1985. Activity patterns and homing in two intertidal limpets, Jordan Gulf of Aqaba. The Nautilus 99: 75-80. Hulings, N. G. 1986. Aspects of the reproduction of rocky intertidal mollusks from the Jordan Gulf of Aqaba (Red Sea). The Veliger 28:318-327. Hulings, \. C. 1987. .\spects of the ecology of the mollusks of the rocky intertidal zone, Jordan Gull of .Aqaba (Red Sea). Dirasat 14:155-171. Hulings, N. C. 1989. A review of marine science research in the Gulf of .-Kqaba. Publications of the Marine Science Station, Aqaba 6:267 p. Leloup, E. 1937. Notes sur les polyplacophores. Proceedings of the Malacological Society, London 22:163-176. Leloup, E. 1960 Amphineures du Golfe d .Aqaba et de la Peninsule Sinai. Sea Fisheries Research Station of Haifa Bulletin 29:29-55. Leloup, E. 1980. Ghitons de la Mer Rouge, du Golfe de Suez et de la Mediterranee. Bulletin de la Institut royal des Sciences naturelles de Belgique 52:1-14. Levanon-Spanier, I., E. Padan, and Z. Reiss 1979. Primary production in a desert-enclosed sea — the Gulf of Elat (Aqa- ba), Red Sea. Deep-Sea Research 26:673-685. Mastaller, M. 1978. The marine molluscan assemblages of Port Sudan, Red Sea. Zoologische Mededelingen, Leiden 53:117-144. Mastaller, M. 1979, Beitrag zur P'aunistik Okologie der Mol- lusken und Echinodermen in den Korallenriffen bei Aqa- ba, Rotes Meer Doctoral Dissertation, Ruhr-L'niversitat Bochum, Federal Republic of Germany, 344 p, Mergner, H. and H, Schuhmacher, 1974, Morphologie, Oko- logie und zonierung von Korallenriffen bei Aqaba (Golfe von Aqaba, Rotes Meer), Helgolander Meeresunter- suchungen 26:238-358, Moulton, J. M. 19(i2. Intertidal clustering of an Australian gastropod. Biological Bulletin 123:170-178. Pearse, J. S. 1978. Reproductive periodicities of Indo-Pacific Invertebrates in the Gulf of Suez. I\'. The chitons Acan- thopleura haddoni Winckworth and Unithochiton lyelli (Sowerby), and the abalone Haliotis pustulata Reeve. Bul- letin of Marine Science 28:92-101. Pearse, J S 1983 The Gulf of Suez: signs of stress on a tropical biota. Bulletin of the Institute of Oceanography and Fish- eries, .Arab Republic of Eg\ pt 9:148-159. Rees, VV. J. and A. Stuckey . 1952. The "Manihine" Expedition to the Gulf of Aqaba NT Mollusca Bulletin of the British Museum (Natural History), Zoology 1:183-202. Rohde, K. and R. Sandland. 1975. Factors influencing clus- tering in the intertidal snail Ccrithium inoniliferum. Ma- rine Biology 30:203-215. Safriel, Li. 1968. Bird migration at Elat, Israel. Ibis 10:283- 320. Safriel, L'. and V, Lipkiu. 1964. On the intertidal zonation of the rocky shores at Eilat (Red Sea, Israel). Israel Journal of Zoology 13:187-190, Schiifer, W. 1954. Form and Funktion der Brachy uren-Schere. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 489:1-65. Soliman, G. N. and A. N. Iskander. 1982. The reproduction and development of the common rock chiton Acantho- pleura spiniger (Sowerby) from the northwestern Red Sea, Malacologia 22:205-210, Steinitz, H, and A, Ben-Tuvia, 1955, Fishes from Eylath (Gulf of .Aqaba), Red Sea, Second report Sea F'isheries Research Station of Haifa Bulletin 11:3-15, Sykes, E, R, 1907, Reports on the marine biology of the Sudanese Red Sea, V, On the Polyplacophora, or chitons. Journal of the Linnean Society of London 31:31-34. Taylor, J. D. and K. Way. 1976. Erosive activities of chitons at Aldabra Atoll. Journal of Sedimentarv Petrologv 46: 974-977. Thorne, M. J. 1967. Homing in the chiton Acanthozostera genimata (Blainville). Proceedings of the Ro\al Societv of Queensland 79:99-108. Thorne, M. J. 1968. Studies on homing in the chiton Acan- thozostera genimata. Australian Journal of .Marine and Freshwater Research 19:151-160. Tortonese, E. 1968. Fishes from Eilat (Red Sea). Sea Fisheries Research Station of Haifa Bulletin 51:6-30. L'ziel, J. 1968. Sea level at Ashod and Elat: differences be- tween prediction and observations. Israel Journal of Earth- Sciences 17:137-151. N'annini. M., G. Ghelazzi, L. Ghelazzi, A. Ercolini. F. Ferrara, G. Messana, P. Messeri, and L. Pardi 1977. Researches on the coast of Somalia. The shore and the dune of Sar Uanle. 13. Physical environment: geomorphical notes, cli- mate and tides. Monitore Zoologica Italiano N.S. Supple- ment 11:249-271. THE NAUTILUS 105(l):26-37, 1991 Page 26 The Southeastern BraziUan Muricidae Collected by RV Marion-Dusfresne in 1987, with the Description of Three New Species Roland Ilouart Research Associate Iiistitut Royal B-3400 Latiden (Ezeinaal), BELGIUM ABSTRACT Twenty-five species of Muricidae are recorded from south- eastern Brazil, including eight species of Muricinae, nine of Muricopsinae, one of Ergalataxinae, one of Ocenebrinae, four of Trophoninae, and two of Typhinae. A short discussion is gi\en for each species. Three new species are described: Der- mornurex leali n.sp. differs from related taxa by the presence of 5 varices on its last teleoconch whorl and by its different intritacalx; Favartia varimtitabilis n.sp. is compared with F. alveata, from which it differs by the erratic placement of the varices and by its multispiral protoconch; Trophon mucrone n.sp. differs from T. lacuneUiis. T. dabneiji. and T. limicola by its spiral sculpture, rounded teleoconch whorls, paucispiral protoconch, and number of axial lamellae. Two species remain unidentified and are listed as Muricopsis (?Murexsul) sp. and Favarlia (Favartia) sp. Drupa didyma Schwengel, 1943 is syn- onymized with TrachijpoUia turricula (von Maltzan, 1884). Kcij words: Muricidae; Brazil; Dermomurcx: Favartia. Tro- phon; new species. INTRODUCTION The material examined was dredged or trawled off the southeastern coast of Brazil during the Expedition MD55 carried out by the R.V. Marion-Dufresne in May 1987, as a joint project of Museum National d'Histoire Natu- relle (MNHN), Paris, and Universidade Santa Ursula, Rio de Janeiro. Drs. A. Guiile and F. Roumos were cruise leaders. Drs. P. Bouchet, J. Lea! and B. Metivier sorted the gastropod material on board. Muricidae were col- lected from 34 to 1,575 m, mostly along a series of sea- mounts running perpendicular to the Brazilian coast, between 18°50' and 23°47'S. Most of species collected are already known to occur off the Brazilian coast; Chi- coreus (Siratus) consuelae (Verrill, 1950), Poirieria (Pa- zinotus) stimpsonii (Dall, 1889), Pygrnaepterys ger- mainae Yokes and D'Attilio, 1980, and Trophon vrrrillii Bush, 1893, however, represent new Brazilian records. A report on the deep-water Volutidae collected during the same cruise has been already published (Leal & Bouchet, 1989). For references to original descriptions not indicated here, the reader is referred to Radwin and D'Attilio (1976). The number of dead collected specimens is followed by "dd"; the number of live collected specimens is iden- tified by "Iv ". The following institutional abbreviations are used: MCZ, Museum of Comparative Zoology, Cambridge; MNHN, Museum d'Histoire Naturelle, Paris; MORG, Museu Oceanografico de Rio Grande; USNM, National Museum of Natural History, Smithsonian Institution. Prefixes for station data denote collecting equipment: CB, Blake trawl; CP, beam trawl; DC, dredge. SYSTEMATICS Family MURICIDAE Rafinesque, 1815 Subfamily MURICINAE Rafinesque, 1815 Chicoreus (Siratus) coltrorum Yokes, 1990 Type locality: Ilha de Itaparica, Bahia, Brazil. Chicoreus (Siratus) coltrorum \'okes, 1990:127, figs. 7-13. (figure 3) Material examined: Southeastern Brazil, stn. DC26, Yi- toria Bank, 20°21'S, 36°59'W, 97 m (2 dd); stn. DC27, Montague Bank, 20°26'S, 36°42'W, 81 m (1 dd); stn. DC40, Davis Bank, 20°40'S, 34°41\V, 60 m (5 dd); stn. DC43, Dogaressa Bank, 20°51'S, 33°45'W, 63 m (2 dd); stn. DC47, Columbia Bank, 20°42'S, 32°13'W, 94-105 m (5 dd); stn. DC48, Columbia Bank, 20°44'S, 32°08"V\', 250-300 m (1 dd); stn. DC50, Coluinbia Bank, 20°44'S, 31°50'W, 95-120 m (3 dd); stn. DC82, 18°56'S, 37°52'W, 85-105 m (1 dd). A recently de,scribed and often misidentified species, C. coltrorum is superficially similar to C. consuelae (Ver- rill, 1950) and nia\ be distinguished mainK b\ the dif- ferent protoconch (figures 3, 4). The protoconch of C. coltrorum has l'/2 rounded whorls while C. consuelae has a conical protoconch consisting of 2Vi whorls; other differences are minimal. Onl\' empty shells were col- lected. Rios (1985:82) reports this species from Brazil (as R. Houart, 1991 Page 27 Stratus consuelae), from Atol das Rocas, Fernando de Noronha; Rio Grande do Norte to Espirito Santo. Chicoreus (Siratiis) consuelae (Verril, 1950) (figure 4) Material examined: Southeastern Brazil, stn. DC52, Martin Vaz Island, 20°29'S, 28°51'W, 64-80 m (3 dd); stn. DC59, Trindade Island, 20°30'S, 29°19'W, 52-60 m (1 dd). Similar to C. coltrorum (see above), the species is re- ported from off the coast of Te.xas to Curasao and in the Lesser Antilles (Yokes, 1990:129). Chicoreus (Siratus) tenuivaricosus (Dautzenberg, 1927) (figures 18, 19) Material examined: Southeastern Brazil, stn. CB90, 19°34'S, 39°34'W, 34 m (33dd) (6 Iv); stn. CB92, 19°34'S, 38°55'W, 340-360 m (1 dd) (1 Iv). Many specimens, mostly juveniles, were collected from station CB90 in dead and live condition. Two specimens, of which one was alive, were taken from station CB92, in 340-360 m; this record is probabK based on specimens that had remained entangled in the net of a previous haul (stn. CB90), and this bathymetrical range requires confirmation. The radula is muricine, with a lateral sickle-shaped tooth, and a central tooth with 3 major and 2 minor cusps (figures 18-19). Aspella morchi Radwin and D'Attilio, 1976 (figures 9, 10, 29) Material examined: Southeastern Brazil, stn. DC22, Vi- toria Bank, 20°32'S, 38°11'W, 52 m (4 Iv); stn. DC40, Davis Bank, 20°40'S, 34°41'W, 60 m (1 Iv); stn. DC41, Davis Bank, 20°39'S, 34°43'W, 58-70 m (2 Iv). The paucispiral protoconch (figures 9, 10) had not previouslv been figured. The cancellate intritacal.x is well illustrated by Radwin and D'Attilio (1976:223, fig. 167). A specimen with 6 teleoconch whorls, and broken pro- toconch, (MNHN, station DC40) (figure 29) is 10 mm long, while the maximum length given by Radwin and D'Attilio is of 6.6 mm. PreviousK known from Natal Bay (type locality), Fortaleza (Cerea) (Radwin & D'Attilio, 1976) and Kapmoan (Bahia) (coll. R. Houart). Aspella senex Dall, 1903 Material examined: Southern Brazil, stn. DC22, Vitoria Bank, 20°32'S, 38°11"W, 52 m (1 dd). The geographical range reported by Radwin and D'Attilio (1976:25) includes North Carolina and the Gulf of Mexico. Rios (1985:87) reported this species from the -ggSB Figures 1, 2. Intritacalx on last teieoconcli whorl (scale bars: 0 5 inni) 1. Dcrmomurex (Trialalclla) oxum Petuch. 2. Der- iiKimiirex (TriahitcUa) Icali n.sp. Abrolhos Islands. The present record extends the geo- graphical range further southward. Dermomurex (Trialatella) oxum Petuch, 1979 (figures 1, 26) Dcrmomurex (Trialatella) oxum Petuch, 1979:517, fig, IE, F. Type locality: 2 km east of Santa Barbara Island, Abrol- hos Archipelago, Bahia State, Brazil, 17°57'S, 38°41"W, in 25 m. Material examined: Southeastern Brazil, stn. DC22, Vi- toria Bank, 20°32'S, 38°11'W, 52 m (1 dd); stn. DC40, Davis Bank, 20°40'S, 34°41'W, 60 m (2 Iv); stn. DC82, 18°56'S, 37°52'W, 8.5-105 m (1 Iv). The largest shell is 15.5 mm long (figure 26), while Petuch mentions 13 mm for his largest specimen (ho- lotype). Dermomurex (Trialatella) leali n.sp. (figures 2, 5, 27, 28) Type material: Holotype MORG 26457, 9.5 x 5.6 mm, southeastern Brazil, stn. DC22, Vitoria Bank, 20°32'S, 38°11"W, .52 m (dd). Description: Shell 9.5 mm in length, roundly-fusiform. Spire moderately high, with 1.5 protoconch whorls and 4 convex teleoconch whorls (subadult). Suture deeply impressed. Protoconch whorls rounded. Axial ornamen- tation of first teleoconch whorl consisting of 5 ribs, second whorl of 6 ribs, third whorl of 7 ribs. Last teleoconch whorl with 4 weak, lightly rounded varices. No other Page 28 THE NAUTILUS, Vol. 105, No. 1 y Figures 3-17. Proloconchs (scale bars: 0.5 mm). 3. Chiroreus {Siratus) coUrorum Yokes, stn. DC26. 4. Chtcorcus (Siratus) conmelae{\\-u\\\). stn DC52. 5. nermomurex {Trialatella)leali n.sp , Ik.IoIn pe MOKG 26457, stn D(:22 6. 7. Poirieria (Pa^"'"'"s) stimpwnii (Dall), stn. DC82, 8. Muricopsis {?Murexsul) sp., stn. DC35 9, 10. Aspella morchi Radwiii & D .\ttilu., stn L)^"!"; ' '• Muricopsis {Risomurex) necocheana (Pilsbrv), stn. DC22. 12. Favartia (CMrilnella) varimutaMis n.sp , holotspe MORG 26458, sin. DC;i5. 13. Favartia {Favartia) sp , stn. 'dC35. 14. Trophon aculeatus Watson, stn. CB104. 15, 16. Trophon mucrone n.sp., paratype MORG 26459, stn. IX:70 17. Siphonochelus (Siphonochchis) riosi (Bertsch & D'Attilio), stn, CB104. R. Hoiiart, 1991 Page 29 21 23 ^ 25 Figures 18-25. Radulae (scale bars figures 20, 22, 24: 20 Mm; figures 21, 23, 25: 10 Mm; figure 18: 100 Mm; figure 19: 50 Mm), 18, 19. Chicoretis iSirattis) tcnuivaricosus (Dautzenberg), stn. CP90, 20, 21. Favartia (CarilneUa) varimutabilis nsp., paratype MNHN. 22, 23. Trachypollia turricula (von Matzan), stn. DC34. 24, 25. Siphonochelus (Siphonochelus) riosi (Bertsch & DWttilio), stn. CB104. Page 30 THE NAUTILUS, Vol. 105, No. 1 R. Hoiiart, 1991 Page 31 axial sculpture. No apparent spiral sculpture except thick, whitish intritacalx, strongly reticulate when thin surface laver partially removed, otherwise minutely sculptured with sharp axial striae (figure 2). Aperture broad, roundly-ovate. Columellar lip smooth, adherent. Anal notch broad, shallow. Outer lip weakly erect, with 4 low denticles within. Siphonal canal short, open, bent adaperturally at tip. Shell whitish. Etymology: Named after J. H. Leal, School of Marine and Atmospheric Science, Miami, Florida, member of the scientific staff on board of the Marion-Dtifresne dur- ing the MD55 cruise. Discussion: The Dermonmrex of the Western Atlantic region have been revised by Yokes (1975, 1976, and 1985) and include the following Recent species: Dermomurex (Dermonmrex) panperctiliis (C. B. Adams, 1850); Der- nwmurex (Dermomurex) alabastrum (A. Adams, 1864); Dermomurex (Dermomurex) pacei Petuch, 1988; Der- momurex (Gracilimurex) elizabethae (McGinty, 1940); Dermomurex (Trialatella) abtjssicola (Crosse, 1865); Dermomurex (Trialatella) oxum Petuch, 1979; Der- momurex (Trialatella) glicksteini Petuch, 1987; Der- momurex (Trialatella) kaicherae Petuch, 1987; Der- momurex (Trialatella) cuna Petuch, 1990. In addition there are several fossil species, discussed and illustrated in Yokes (1975). The holotype of D. abyssicola has not been located (Yokes, 1975:148) but the original illustration shows a shell with 3 strong varices and intervaricial ridges, while D. leali has 5 varices on the last teleoconch whorl. Der- momurex abyssieola has a brown band at the base of its last whorl and a longer siphonal canal. Dermomurex leali differs from D. oxum in having 5 varices on its last whorl instead of 3 strong varices and intervaricial ridge between pairs of varices as in D. oxum. Dermomurex leali has more rounded whorls, covered by a more finely sculptured intritacalx (figure 2), its aperture is relatively larger, the siphonal canal is shorter and the spire is lower. Dermomurex leali differs from D. glicksteini and D. kaicherae by its more globose shell, its more numerous varices on last teleoconch whorl, its absence of intervar- icial axial ridges, and shorter siphonal canal. In addition, D. oxum, D. glicksteini and D. kaicherae have more or less obvious spiral sculpture when the in- tritacalx is removed, whereas it is absent in D. leali. Dermomurex leali differs from D. cuna by its more rounded varices, shorter siphonal canal, comparatively larger aperture and by its different and more sharply striate intritacalx. Poirieria (Pazinotus) stimpsonii (Dall, 1889) (figures 6, 7, 32) IVIaterial examined: Southeastern Brazil, stn. I)C22, Vi- toria Bank, 20°32'S, 38°11'W, 52 m (2 dd); stn. CB79, 19°02'S, 37°48'W, 1,500-1,575 (1 dd); stn. DC82, 18°56'S, 37°52'W, 85-105 m (1 Iv). Four juveniles, easily recognizable shells. The illus- trated specimen (figure 32) is pinkish. The species is known from western Florida to Barbados (Radwin & D Attilio, 1976:87), but has not previously been recorded south of Barbados. Subfamily MURICOPSINAE Radwin & D'Attilio, 1971 Muricopsis (?Murexsul) sp. (figures 8, 30, 31) Material examined: Southeastern Brazil, stn. DC35, Ja- seur Bank, 20°42'S, 35°22'W, 82-105 m, (1 Iv) (juvenile); stn. DC43, Dogaressa Bank, 20°51'S, 33°45'W, 63 m (1 dd, 3 Iv) (juveniles). The largest of the Brazilian specimens is 5.2 mm long. All are pinkish. Examination of adult specimens is nec- essary for positive identification. Muricopsis (Risomurex) necocheanus (Pilsbry, 1990) (figure 11) Material examined: Southeastern Brazil, stn. DC15, 2r37'S, 40°18'W, 37 m (1 dd) (damaged); stn. DC22, Yitoria Bank, 20°32'S, 38°11'W, 52 m (3 Iv); stn. CP25, Yitoria Bank, 20°36'S, 37°27'W, 65 m (1 dd). Recorded by Rios (1985:84) from Espirito Santo, Bra- zil, to Bahaia Blanca, Argentina. Yokes and Houart (1986: 84) did not observe a complete protoconch, but, from the material then available, they presumed the proto- conch to be rounded instead of keeled as in most Muri- copsis. The protoconch here illustrated (figure 11) con- firms their assumption. Favartia (Favartia) cellulosa (Conrad, 1846) Material examined: Southeastern Brazil, stn. DC29, Montague Bank, 20°27'S, 36°41'W, 310-350 m (1 dd). Recorded by Rios (1985:85). The only specimen col- lected during the expedition is an empty, juvenile, dam- aged shell, obviously carried downslope from shallow water. Figures 26-33. 26. Dermomurex (Trialatella) oxum Petuch, stn. DC22, 15.5 x 8 mm. 27, 28. Dermomurex (Trialatella) leali n.sp , holotype MORG 26457, 9.5 x 5.6 mm. 29. .\spella morchi Radwin & D'Attilio, stn DC40, 10 x 5 mm, 30, 31. Muricopsis (?Murexsui) sp., stn. DC35, 5.2 x 3.2 mm. 32. Poirieria {Pazinotus) stimpsonii (Dall), stn. DC82, 6.5 x 4,2 mm, 33. Favartia (Favartia) sp., stn. DC22, 8.2 x 5 mm. Figures 34-36. Favartia (Caribiella) varimutabilis n sp 34, 35. Holotype MORG 26458, 9x5 mm. 36. Paratype MNHN, stn DC22, 9 x 5.1 mm Figure 37. Pygmaepterys germainae Yokes & D'Attilio, stn. DC34, 5.5 X 3.4 mm. Page 32 THE NAUTILUS, \ol. 105, No. 1 Favartia (Favartia) nucea (Morch, 1850) Material examined: Southeastern Brazil, .stn. D(]15, 2r37'S, 40°18'W, 37 m (1 dd); stn. DCMO, Davis Bank, 20°40'S, 34''41'W, 60 m (1 Iv) (juvenile). A species that has previously been recorded from Bra- zil hv Rios (1985:85). Favartia (Favartia) sp. (figures 13, 33) IVIaterial examined: Southeastern Brazil, stn. DC22, Vi- toria Bank, 20°32'S, 38°11'W, 52 m (1 dd); stn. DC35, Jaseur Bank, 20°42'S, 35°22'W, 82-105 m (3 iv) (juve- niles). The shell is apparently different from any known west- ern Atlantic Favartia species. The shell and the proto- conch of a juvenile specimen are here illustrated. The colour is light pink to light brown. The material collected during the expedition consists only of an adult but damaged shell (figure 33) and of 3 juveniles. There is presently insufficient material to com- pare in more detail with other Favartia. Favartia (Favartia) varimutabilis n.sp. (figures 12, 20, 21, 34, 35, 36) Type material: Holotvpe MORG 26458, 9x5 mm, southeastern Brazil, stn. DC15, 21°37'S, 40°18'W, 37 m (dd); 3 paratypes MNHN: stn. DC22, Vitoria Bank, 20''32'S, 38°11"W, 52 m (1 dd); stn. DC27, Montague Bank, 20°26'S, 36°42'W, 81 m (1 Iv); stn. DC40, Davis Bank, 20°40'S, 34°41'W, 60 m (1 dd). Other material examined: Stn. DC24, Vitoria Bank, 20°42'S, 37°50'W, 48-52 m, coll. R. Houart (1 dd). Description: Shell 8.5-9 mm in length, stout, fusiform. Spire high, with 2.5 protoconch whorls and 5 weakly convex, teleoconch whorls. Suture impressed. Protoconch whorls rounded, ending with a deeply indented varix. Last whorl with 3-5 irregular, rounded, erratically placed, squamous varices. Penultimate whorl bearing 8 or 9 var- ices. No intervaricial axial sculpture. Spiral sculpture of 7 cords, occasionally with 1 thread between each pair of cords, 2 or 3 adapical cords sometimes extending as small spinelets on varices. Aperture roundly-ovate, outer lip erect and smooth, weakly lirate within. Columellar lip rim erect, with 2 or more small and narrow folds. Anal notch broad, shallow. Siphonal canal short, narrowly open, weakly bent adap- erturally, ornamented with 2 small open spines. Shell pale orange to pale brown, with darker spiral cords. Radula typically muricopsine with a prominent cen- tral cusp (figures 20, 21). Etymology: Vari: from varix; nmtalnlis: variable. Named for its variable shape of shell and varices. Discussion: It .seems doubtful that the shells illustrated in figures 34, 35 and 36 are conspecific. In fact, the last teleoconch whorl of F. varimutabilis may be weakly convex or rounded, with erratical placement and variable form of the varices. Early teleoconch whorls are identical in all specimens examined. Favartia (Carihiella) alveata (Kiener, 1842), known from Florida to Brazil has a shell with fewer, larger, scabrous cords, regularly placed varices and a paucispiral protoconch of 1.5 whorls, denoting nonplanktotrophic larval development. Favartia varimutabilis is also related to the Pacific species F. jeanae Emerson and D'Attilio, 1979, known from the Philippine Islands. Favartia (Murexiella) hidalgoi (Crosse, 1868) IVIaterial examined: Southeastern Brazil, stn. DC55, Martin Vaz Island, 20°32'S, 28°52'W, 780-795 m (1 dd) (fragment); stn. DC81, 19°00'S, 37°48'W, 120-135 m (1 dd). Known from the Gulf of Mexico to Brazil. In Brazil recorded from Rio Grande do Sul by Rios (1985:85). Only 2 damaged and dead specimens were collected during this expedition. Favartia (Murexiella) glypta (M. Smith, 1938) Material examined: Southeastern Brazil, stn. DC15, 21°37'S, 40°18'W, 37 m (1 dd, 1 Iv); stn. DC22, Vitoria Bank, 20°32'S, 38°11'W, 52 m (3 dd) (juveniles); stn. DC73, 19°00'S, 37°48'V^, 607-620 m (1 dd) (juvenile). Comparison of the photograph of the holotype of Mu- rex glypta Smith, 1938 illustrated in Vokes (1968:117), of a very similar shell from Ilha do Pai, Brazil (coll. R. Houart), and of the type material of Murexiella iemanja Petuch, 1979 (holotvpe USNM 780652 and paratype USNM 780653) confirms the conclusion of Rios (1985: 85) who synonymised Murexiella iemanja Petuch, 1979 with Murex glypta Smith, 1938. The other shell illustrated as Favartia glypta in Vokes (1968: pi. 7, fig. 5) from the post-Caloosahatchee For- mation, Florida (specimen USNM 645895) is Favartia celltilosa (Conrad, 1846). Rios (1985:85) records F. glypta from Amapa to Rio de Janeiro and Abrolhos Islands. Fygmaepterys germainae Vokes and D'Attilio, 1980 (figure 37) '^Fygmaepterys germainae Wikes and D'.\ttilio, 1980:50, pi. 1, figs. 1-4. Type locality: Off Punta Higuero, northwestern Puerto Rico, 90 m. Material examined: Southeastern Brazil, stn. DC24, Vi- toria Bank, 20°42'S, 37''50'W, 48-52 m (1 dd); stn. DC27, Montague Bank, 20°26'S, 36°42'W, 81 m (1 dd): stn. DC34, Jaseur Bank, 20°28'S, 35°54'W, 54 m (1 dd). Described from fossil specimens (Miocene and early Pleistocene) from the Dominican Republic and Costa R. Houart, 1991 Page 33 Rica and from Recent specimens from the Caribbean Sea (Puerto Rico to Panama, in 12-90 m). The presence of this species off southeastern Brazil extends consider- ably its geographical range. this synonymy. The multispiral protoconch indicates planktotrophic development, which accounts for the broad distribution of this species in the western and east- ern Atlantic. Subfamily ERGALATAXINAE Kuroda and Habe, 1971 Cytharomorula graiji (Dall, 1889) Material examined: Southeastern Brazil, stn DC28, Montague Bank, 20°27'S, 36°42'W, 525-600 m (1 dd); stn. DC48, Columbia Bank, 20°44'S, 32°08'W, 250-300 m (2 dd); stn. DC55, Martin Vaz Island, 20°32'S, 28°52' W, 780-795 m (14 dd); stn. DC73, 19°00'S, 37°48'W, 607- 620 m (3 dd); stn. CB77, 19°41'S, 37°48'W, 790-940 m (2 dd); stn. DC81, 19°00'S, 37°48'W, 120-135 m (2 dd); stn. DC82, 18°56'S, 37°52'W, 85-105 m (3 dd); stn. CB92, 19°34'S, 38°55'W, 340-360 m (1 dd); stn. CB103; 23°36'S, 42°02'W, 200-217 m (1 Iv). Mentioned by Radwin and D'Attilio (1976:44) from Barbados and by Rios (1985;86) from Rio de Janeiro to Rio Grande do Sul in 80-175 m. The species probably lives throughout the Caribbean to Brazil. Also known in the Eastern Atlantic in the Ca- nary Islands, Madeira and St. Helena (Bouchet & Waren, 1985). Subfamily OCENEBRINAE Cossmann, 1903 TrachypoUia turricula (von Maltzan, 1884) (figures 22, 23) Cantharus (Pollia) turricula von Maltzan, 1884:67. Type locality: Goree, Senegal, West Africa. Material examined: Southeastern Brazil, stn. DC15, 21°37'S, 40°18'W, 37 m (1 dd); stn. DC22, Vitoria Bank, 20°32'S, 38°11'W, 52 m (4 dd, 1 Iv); stn. DC27, Montague Bank, 20°26'S, 36°42'W, 81 m (1 dd, 2 Iv); stn. DC28, Montague Bank, 20°27'S, 36°42'W, 525-600 m (1 dd); stn. DC30, Jaseur Bank, 20°27'S, 36°17'W, 60 m (1 Iv); stn. DC34, Jaseur Bank, 20°28'S, 35°54'W, 54 m (2 Iv); stn. DC35, Jaseur Bank, 20°42'S, 35°22'W, 82-105 m (9 dd); stn. DC40, Davis Bank, 20°40'S, 34°41'W, 60 m (1 dd); stn. DC42, Dogaressa Bank, 20°55'S, 34°01'W, 60 m (6 dd); stn. DC43, Dogaressa Bank, 20°5rS, 33°45'W, 63 m (1 dd, 3 Iv); stn. DC47, Columbia Bank, 20°42'S, 32°13"W, 94-105 m (1 dd); stn. DC61, Trindade Island, 20°29'S, 29°18'W, 63 m (1 dd); stn. DC82, 18°56'S, 37°52'W, 85-105 m (3 dd); stn. DC83, 18°50'S, 37°57'W, 60 m (1 dd). The shell dredged at station DC28 (525-600 m) was empty and damaged, and most likely carried downslope. Drupa didyma Schwengel, 1943 described from off Palm Beach, Florida, is similar; comparison with the original diagnosis and with specimens from West Africa, Florida (coll. R. Houart), and Brazil (MNHN) confirms Subfamily TROPHONINAE Crossmann, 1903 Trophon aculeatus Watson, 1883 (figures 14, 40) Trophun aculeatus Watson, 1882:390. Type locality: Off Pernambuco, 09°05'S, 34°50'W, 640 m. Material examined: Southeastern Brazil, stn. CB77, 19°41'S, 37°48'W, 790-940 m; stn. CB104, 23°42'S, 42''07'W, 430-450 m; stn. CB105, 23°47'S, 42°10'W, 610 m (many live and dead collected specimens). Rios (1985:88) reported this species to occur off Brazil (off Pernambuco) and Guadeloupe and included the type- locality of T. lacunellus (Dall, 1889), a different species that Rios synonymised with T. aculeatus. Live-taken specimens were collected by R.V. Marion-Dufresne in 430-450 m. All specimens collected in 610-940 m were empty shells. Trophon pelseneeri E. A. Smith, 1915 Trophon pelseneeri E. A. Smith, 1915:92, pi. 2. figs. 6-7. Type locality: West of Falkland Islands, 229 m. Material examined: Southeastern Brazil, stn. CBlOl, 22''58'S, 42°06'W, 50 m (1 dd); stn. CB102, 23°07'S, 42"'04'W, 100 m (1 dd) (damaged). Recorded by Rios (1985:88), from the state of Rio de Janeiro, Brazil, to the Falkland Islands, Argentina. Tro- phon orbignyi Carcelles, 1946 is a synonym (Rios, 1985: 88), Trophon pelecetiis Dall, 1902 is probably an earlier name, but study of more specimens is required to confirm this. Trophon verrillii Bush, 1893 (figures 41-43) Trophon verrillii Bush, 1893:214, pi. 1, fig. 16. Type locality: Off Cape Fear, North Carolina, 1,183 m. Material examined: Southeastern Brazil, stn. CB76, 18°59'S, 37°50'W, 637 m (3 dd, 4 Iv). Trophon verrillii differs from T. aculeatus in having a larger protoconch and totally lacking spiral ornamen- tation. The illustrated specimen (figures 41, 42) is very similar to the holotype (figure 43). To m\ knowledge T. verrillii was previously known only from the type lo- cality Page 34 THE NAUTILUS, Vol. 105, No. 1 R. Houart, 1991 Page 35 30" 15" ATLANTIC OCEAN Trmdade 0 100 200 km 20 24 Figure 46. Map of investigated area with location of cited stations. Reprodnced, with permission, from Leal and Bouchet (1991) with some additions. Trophon miicrone n.sp. (figures 15, 16, 38, 39) Type material: Holotype MNHN, 26.5 x 11 mm, south- eastern Brazil, stn. CB79, 19°02'S, 37°48'W, 1,500-1,575 m (Iv); 7 paratypes MORG 26459; stn. DC70, 18°59'S, 37°48'W, 1,540-1,550 m (4 dd) (juveniles); stn. CB77, 19°41'S, 37°48'W, 790-940 m (1 dd) (damaged); stn. CB79, (2 dd). Other material examined: Stn. CB78, 18°59'S, 37°48' W, 1,200 m, coll. R. Houart (1) (damaged). Distribution: Known only from the material examined. Description: Shell 26.5-28.0 mm in length, roundly- elongate. Spire high with 1.5-1.75 protoconch whorls and 6 or 7 teleoconch whorls. Suture impressed. Protoconch whorls rounded. Axial ornamentation on last teleoconch whorl consisting of 12 sharp, low lamellae and numerous growth striae. Spiral sculpture of 4 or 5 low, weak cords; shoulder smooth, except for axial lamellae. Aperture broadly ovate. Columellar lip smooth. Anal notch obsolete. Outer lip thin, smooth, glossy within. Siphonal canal moderately long, straight, broadly open. Shell whitish. Etymology: Mwcrone-from mucro (tip, sharp end). Named for its arrow or javelin tip shape. Discussion: Three species, Trophon lacunellus (Dall, 1889), described from off Guadeloupe, in 1,406 m, T. dabneyi Dautzenberg, 1889, described from the Azores, in 1,287 m, and T. limicola Verrill, 1885, described from the New England Coast, in 1,542-3,718 m may be com- pared. Trophon lacunellus differs from T. mucrone by its higher spired and smaller shell without spiral sculpture, narrower teleoconch whorls, and more rounded proto- conch whorls. Trophon dabneyi has more rounded teleoconch w horls than T. mucrone, spiral cords that are absent or very weak, and a multispiral protoconch of 2.5 whorls (prob- Figures 38-40. ,38, 39. Trophon nuicrone n.sp., holotype MNHN, 26.5 x 11 mm. 40. Trophon aculealus Watson, stn. CB104, 13.9 X 7 mm Figures 41-43. Trophon verrillii Bush. 41, 42. Stn. CB76. 10 x .5 5 mm 43. Holotype, MCZl 19157, 13.9 x 6.8 mm Figures 44, 45. Siphonochehts (Siphonochelus) riosi Bertsch & D'Attilio, stn CB103, 11.5 x 5 2 mm. Page 36 THE NAUTILUS, Vol 105, No. 1 able planktotrophic development) (Hoiiart, 1981: pi. 3, fig. 14; Bouchet & Warra, 1985: fig. 305). The holotype of T. limicola Verrill, 1885, a 10.5 mm high shell, was also compared with a 9.7 mm high ju- venile specimen of T. mucrune. Trophon limicola has 20 axial lamellae on its last teleoconch whorl (only 9 in T. mucrone), a longer siphonal canal, more rounded protoconch whorls, and lacks spiral sculpture. Additional notes on West Atlantic Trophon species: Trophon aculeatus Watson, 1883 and T. verrillii Bush, 1893 differ in many features such as spiral sculpture, number of axial lamellae, shape of whorls and proto- conch. Trophon abijssorum Verrill, 1885, described from a juvenile specimen of 8 mm, is more closely related to T. verrillii. and has angulate and smooth teleoconch whorls, prominent spines at the shoulder and relatively longer siphonal canal. Subfamily TYPHINAE Cossmann, 1903 Typhis (Typhina) belcheri Broderip, 1833 Material examined: Southeastern Brazil, stn. CB102, 23°07'S, 42°04'W, 100 m (1 dd). The holotype of Typhis belcheri was illustrated by Kaicher (1980: card 2514). Comparison with the holotype of Typhis cleriji Petit, 1842 (MNHN), and with many specimens from Brazil (coll. R. Houart) and from West Africa (MNHN and coll. R. Houart) lead to the conclu- sion that these taxa are conspecific. The species is re- corded by Rios (1985:89) as Riigotyphis cleryi (Petit, 1842), from Rio de Janeiro to Torres, Rio Grande do Sul. It lives usually in shallow water. Typhis belcheri is also known from the Cape Verde Archipelago and Senegal (West Africa) (MNHN). Siphonochelus (Siphonochelus) riosi (Bertsch & D'Attilio, 1980) (figures 17, 24, 25, 44, 45) Typhis (Typhina) riosi Bertsch & D'Attilio, 1980:135, figs. 6, 7. Type locality: Off Tramandai, southern Brazil, 30°04'S, 47°55'W, 280 m. Material examined: Southeastern Brazil, stn. CB92, 19°34'S, 38°55'W, 340-360 m; stn. CB102, 23°07'S, 42°04'W, 100 m; stn. CB103, 23°36'S, 42°02'W, 200-217 m; stn. CR104, 23°42'S, 42°07'W, 430-450 m; stn. CB105, 23°47'S, 42°10'W, 610 m (many live and dead speci- mens). The specimens collected in 100-217 m are larger than tho.se dredged in 430-450 m, but no other differences arc apparent between the two samples. As already men- tioned by Vokes (1984:215), the specimen illustrated as Typhis longicornis Dall in Bayer (1971: fig. 31) is S. riosi. The radula (figures 24, 25) consists of rachidian teeth and a pair of curved lateral teeth. Rachidian teeth with 3 main cusps and asymmetrical lateral denticles. ACKNOWLEDGMENTS I am very grateful to Dr. P. Bouchet (Museum National d'Histoire Naturelle, Paris) and to Dr. J. H. Leal (Ro- senstiel School of Marine and Atmospheric Science, Mi- ami) for giving me tfie opportunity to examine this ma- terial and for their much appreciated suggestions, to Dr. R. S. Houbrick, D. M. Tyler, S. S. Greenhouse and F. J. Collier (National Museum, of Natural History, Smithson- ian Institution, Washington, DC) and to Dr. S. P. Kool (Museum of Comparative Zoology, Cambridge) for the loan of type material, to Dr. P. Bouchet and Dr. A. Waren (Swedish Museum of Natural History, Stockholm) for preparation and SEM work of the radulae, and to Dr. P. Bouchet and Prof. E. H. Vokes (Tulane University, New Orleans) for critical review of the manuscript. LITERATURE CITED Bertsch, H. and A D'.4ttilio. 1980. New species of Muricidae (Gastropoda) from the Indian Ocean, the Philippines and Brasil. Venus 39(3);131-138. Bouchet, P and A. Waren. 1985. Revision of the Northeast .'Ktlantic bathyal and abyssal Neogastropoda excluding Turridae (Moliusca, Gastropoda). Bolletino Malacologico supplement 1:123-296. Bush, K. J. 1893. Reports on the results of dredgings, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78), and the Caribbean Sea (1879-80). and along the Atlantic coast of the United States (1880) by the United States coast survey steamer "Blake . . . Report on the Moliusca dredged by the "Blake" in 1880, including de- scriptions of several new species. Bulletin of the Museum of Comparative Zoology 23:197-244 Clench, W J and I Perez Farfante. 1945. The genus Murex in the western .Atlantic Johnsonia 1(17): 1-56. Houart, R. 1981 Revision des Trophoninae d Europe. Infor- mations de la Societe Beige de Malacologie 9(l-2):3-70. Kaicher, S. D. 1980. Card catalogue of world-wide shells, Muricidae V. Privately published, St. Petersburg, Florida. Leal, J. H. and P. Bouchet. 1989. New deep-water Volutidae from off southeastern Brazil (Moliusca: Gastropoda). The Nautilus 103(1):1-12. Leal, J. H. & P. Bouchet. 1991, Distribution patterns and dispersal of prosobranch gastropods along a .seamount chain in the Atlantic Ocean. Journal of the Marine Biological Association of the United Kingdom 71(1) (February 1991). In press. Maltzan, H. F. von. 1884. Diagnosen neuer Senegambischer gastropoden. Nachrichtsblatt der deuteschen Malakozool- ogischen Gesellschaft 16:65-73. Petuch, E J. 1979. New gastropods from the .\brolhos Ar- chipelago and reef complex, Brazil Proceedings of the Biological Society of Washington 92(3):5 10-526. Petucli, E. J 1987. New Caribbean molluscan faunas. CERF, Charlottesville, Virginia: 1-154, addendum .M-A4. Kadwin, G E. and A. D'Attilio. 1976. Murex shells of the world, an illu.strated guide to the Muricidae. University Press, Stanford, 284 p Rios. E. C. 1985. Seashells of Brazil Rio Grande, 328 p. R. Houart, 1991 Page 37 Smitli, E. A, 1915. Mnllusca. Pt. 1, Gastropoda prdsohraiR-liia, Scaphopoda and Pelec\poda -British Antarctic ( "Terra Nova") expedition, 1910. Zoology 2(4):61-112. Yokes, E. H. 196S. Cenozoic Muricidae of the western At- lantic region. Ft 4, Hexaplex and MurexicUa. Tulane Stud- ies in Geology 6(2-3):85-126. Yokes, E. H. 1975. Cenozoic Muricidae of the western At- lantic region, Ft 6, Aspclla and Dermomurcx. Tulane Stud- ies in Geology and Paleontology 11:121-162. Yokes, E. H. 1976. Cenozoic Muricidae ot the western At- lantic region. Dcrmonuircx — addendum Tulane Studies in Geology and Paleontolog\ 1245-46. Yokes, E. H. 1984. Comparison of the Muricidae of the east- ern Pacific and western Atlantic, with cognate species. Shells and Sea Life 16(11):210-216. Yokes, E. H. 1985. Review of the west coast aspelloids Aspclla and Dermomurex (Gastropoda: Muricidae), with the de- scriptions of two new species. The Yeliger 27(4):430-439. Yokes, E. H. 1990 Two new species of Chicoreitu subgenus Sirfl^i.? ((Gastropoda: Muricidae) from Northeastern Brazil. The Nautilus 103(4):124-130. N'okes, E. H. and A. D'Attilio. 1980. I'ygmaepterys, a newly described taxon of Muricidae (Mollusca: Gastropoda), with the description of three new species from the Onozoic of the western Atlantic Tulane Studies in Geology and Pa- leontolog) 16:45-54 Yokes, E. H. and H Houart. 1986. An evaluation of the taxa Miiricopsis and Ri.s(jmurcx (Gastropoda. Muricidae), with one new species of Risoiiiiircx. Tulane Studies in Geology and Paleontology 19:63-88. Watson, R. B. 1882. Mollusca of H.M.S. Challenger expedi- tion. Journal ot the Linnean Society London Part 14372- 392. THE NAUTILUS 105(1):38, 1991 Page 38 News and Notices ADDRESS CHANGE FOR THE NAUTILUS We ask subscribers to make note of the change of address for THE NAUTILUS. The new address is: THE NAUTILUS P.O. Bo.x 7279 Silver Spring, MD 20907-7279 U.S.A. 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Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All conununications 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 Parkwa\', 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 Et7NAUTI LUS CONTENTS Volume 105, Number 2 April 10, 1991 ISSN 0028-1344 Riidiger Bieler Alan R. Rabat William K. Emerson Malacological Journals and Newsletters, 1773-1990 39 First records for Cymatium mundiim (Gould) in the eastern Pacific Ocean, with comments on the zoogeography of the tropical trans-Pacific Tonnacean and non-Tonnacean prosobranch gastropods with Indo-Pacific faunal affinities in western American waters 62- Or> LIBRARY APR 2 3/991 j .i!!!^l_^. Mass. / THE NAUTILUS 105(2):39-61, 1991 Page 39 Malacological Journals and Newsletters, 1773-1990 Riidiger Bieler' Delaware Museum of Natural History P O. Box 3937 Wilmington, DE 19807, USA Alan R. Kabal- Museum of Comparative Zoology Harvard University Cambridge, MA 02138, USA ABSTRACT This compilation lists the 286 malacological journals and news- letters (representing 374 titles), arranged by continent and country The place, duration, and status of each publication is given and changes in titles are cross-referenced. Of these serials, 157 are still published It is determined that about 25?o of current malacological publications are published in these jour- nals devoted solely to mollusks. The geographical distribution (31 countries represented) and the usage of malacological jour- nals are analyzed in terms of the broader context of the scientific journal literature field. Key words: Bibliography, journals; malacology; serials. INTRODUCTION Malacologists all over the world are often overwhelmed b\' the enormous diversity of journals and newsletters in our field. It is virtually impossible for any one library to possess all these journals, let alone keep up with the new serials arising every year. The purpose of this paper is to introduce some order to this chaos by providing com- plete bibliographical data on the corpus of malacological serials. It has been over three decades since the useful, albeit cursory, compilation of Jutting and Altena (1958) who listed only 36 titles. More recently, Biirk and Jung- bluth (1985) provided a detailed index to most of the German malacological publications; this valuable refer- ence essentially supersedes previous German efforts such as Buschmeyer's (1938). A brief treatment of malacolog- ical serials as part of the history of conchology was pre- sented by Dance (1986:145). Scientific journals began in 1665 with the appearance of the Philosophical Transactions [London] and the Journal des Sqavans [Paris]. It would take another cen- ' Current address: Division of Invertebrates, Field Museum of Natural Historv, Roosevelt Road at Lake Shore Drive, Chi- cago, IL 60605, USA. ^Current address: Division of Mollusks NHB-118, National Museum of Natural History, Smithsonian Institution, Wash- ington, DC 20560, L SA tury for the start of the first malacological journals: the late 1700's German serials of J. S. Schroter who published four short-lived serials dealing primarily with fossil mol- lusks (Friess, 1982:93-95). However, "modern" mala- cological journals did not come into full bloom until the mid 1800's, with several European titles (all now de- funct). The oldest currently published malacological journal is the Archiv fiir Mollushenkunde [1868, under an earlier title]. The Zoological Record, Mollusca Section [1864] is an abstracting source and not a contributed journal. As the historian Derek Price (1986:5-6, 18-19) has noted, the total number of scientific journals has doubled every fifteen years, for a 5% increase per year. Yet, ob- viously this exponential rate of increase cannot continue infinitely; eventually a saturation point representing lo- gistic growth will be reached. The proliferation of new journals in the last decade does not seem to indicate that the field of malacology has reached its carrying capacity, although the recent demise of certain journals may ini- tiate this trend. Equally important is the usage of scientific articles: how widely they are read and cited by others. Price (1986:118) used Lotka's law and the Pareto distribution to determine that ". . . about 10 percent of all published papers have never been cited, about 10 percent have been cited once, about 9 percent twice, and so on, the percentages slowly decreasing . . .". This widely quoted statement [usually paraphrased as "10 percent of all pub- lished papers never subsequently cited" (Wheeler, 1989: 1 1)] is the inevitable consequence of what the sociologist Robert Merton (1968:61) diagnosed as "insanabile scri- bendi cacoethes [= the itch to publish]. MATERIALS AND METHODS We undertook this project over a two year period, entailing considerable bibliographic research and cor- respondence. We compiled a master list based on the serial holdings of seven major malacological libraries: Academy of Natural Sciences, Philadelphia; Delaware Museum of Natural History; Field Museum of Natural History; Museum National dHistoire Naturelle, Paris; Museum of Comparative Zoology; The Natural History Museum, London; and National Museum of Natural His- Page 40 THE NAUTILUS, Vol. 105, No. 2 torv, Smithsonian Institution. Then, this hst was aug- mented by extensive correspondence with various col- leagues who checked certain geographical sections, as well as with the editors of shell club newsletters (ad- dresses obtained from the American Malacological Union membership list and various other sources); these indi- viduals are listed in the acknowledgements. Unfortu- nately, no replies were received from some shell clubs and certain other clubs are no longer extant and we were unable to find any complete sets of their newsletters. While most of the major malacological research li- braries have complete sets of the principal journals in the field, the same is certainly not true for the numerous shell club newsletters. It is to be regretted that some museums extend little effort to keep even the newsletter of their own national malacological society. We are aware of the difficulties in treating this "grey" literature and we strongly recommend that malacological libraries en- deavor to maintain holdings of these newsletters (es- pecially those published in their country) to the greatest extent possible. The development of online computerized catalogues of library holdings will allow researchers to readily locate these serials. SCOPE Herein we have attempted as complete a compilation as is possible of all the malacological serials (journals and newsletters) that have come to our attention. As the read- er may be aware, it is not always easy to differentiate between malacological journals and newsletters. Gen- erally, scientific journals publish original research articles and are intended as part of the permanent scientific record. They typically have an editorial review board to maintain scientific standards, and they are often pub- lished by a scientific institution or scholarly society. Newsletters usually serve as a vehicle for news and in- formation within the respective organization, and in the case of shell club newsletters they disseminate knowledge in layman's terms, offer advice for beginners in the field and are often simply meant to entertain. Shell club newsletters are not only of interest to the scientific community because they occasionally provide such original data as biogeographical or habitat infor- mation, but also they become permanent scientific record once they (sometimes unintentionally) publish taxonomic statements, especially new species names and type des- ignations. There is obviously no distinct line between journals and newsletters, seeing that some of today's jour- nals started in newsletter format, while some self-pro- claimed "journals " hardly deserve that label. We made no attempt to sort the malacological serials into these two categories. We have usually excluded those shell club newsletters which are completely restricted to internal communications, such as membership lists and an- nouncements of upcoming events. We do not include multi-volume malacological trea- tises that were issued "in parts " over long time periods but represent a single encyclopedic compilation and not a diverse journal. For example, Kiener's Iconographie des Caquilles Vivantes; the Martini-Chemnitz and Kiis- ter editions of the Systematisches Conchylien-Calnnet; Phihppi's Abbildungen und Beschreibiingen . . . ; Reeve's Conchologia Iconica; the Rossmassler-Kobelt Iconogra- phie der Land- und Siisswasser-Molliisken, Sowerby's Thesaurus Conchyliorum; and the Tryon-Pilsbry Man- ual of Conchology are all excluded from this paper. We also do not include symposia volumes, including those of regular meetings, unless specifically issued in serial format. Several of the French titles in this list actually rep- resent compiled reprints, by one author, of malacological articles from various journals. We have included them since they are obscure and are sometimes cited as if they were serials themselves. We noted several interesting aspects regarding the etymology of the titles of these various serials. Obviously, most are based on "mollusk", "malaco-", "conch-", or "shell "; others from generic names or famous malacol- ogists. One should note several titles that have been used more than once: Argonauta, The Conchologist, Journal of Malacology and Tide Lines. For completeness, and in appreciation of the fact that there is also some intended humor in our field of science, we have included five satirical "serials" in our listings. Three of them, titled Conchologica Nonsensica, The Double Helix and Slug Newsletter, are easily spotted, but The Slug and especially the cleverly produced Jour- nal of Molluscan Behaviour are less obvious. These par- odies were not included in our statistics. In compiling this list, we also noted several journals with "malacological" titles, although they do not qualify for inclusion herein as they are not about mollusks at all, or are not exclusively about mollusks. We list them here to avoid future confusion: Nautilus: A monthly miscel- lany . . . [1845-1846; Sag Harbor, New York]; Astarte, A Journal of Arctic Biology [1951-1983; Tronis0]; The Conch Shell [1953-1966; Bishop Museum, Honolulu]; The Conch: A Biafran Journal of Literary and Cultural Analysis [1969-current; Paris]. An exception is made for the larger shellfisheries journals (which also cover crus- taceans). Here we draw an arbitrary line between those that are included {e.g.. Journal of Shellfish Research). and others that are not {e.g.. Proceedings of the ... th National Shellfish Sanitation Workshop). Coche (1983) lists numerous serial publications in shellfisheries and aquaculture. We would greatly appreciate information regarding any additions or emendations to this list, since it is our intention to publish future "Addenda". There are several shell club newsletters for which we were unable to obtain full data, despite repeated enquiries. In some cases, the shell clubs themselves do not know the full history of their own newsletter(s). These titles are indicated by an asterisk; the information needed is noted. It is imperative that the Zoological Record also be informed of new- publications; editors should send copies (or announce- ments) of their publications to: MoUusca Section, The R Bieler and A. R. Rabat, 1991 Page 41 Natural History Museum, Cromwell Road, London SW7 5BD, England The authors will gladly provide editors with a list of the major malacological research museums that should also be informed of new malacological publications. DISCUSSION It is worthwhile to consider the role of malacological journals, not onl\' within the field of malacology, but more broadh within the biological sciences. We have found that there are 157 currentK published malacolog- ical serials. According to The Serials Directory, 3rd Edi- tion, 1988-89, there are over 118,000 current serial titles in all fields of knowledge (excluding newspapers). More specifically, according to the 1988/89 Zoological Record Serial Sources (1990:vi), there are 5,540 current journals in the zoological sciences that are recorded for the Zoo- logical Record. Obviously, the malacological journals are a very small part of the whole of scientific literature. Yet, their importance in the field of malacology is far greater than might have been suspected. In order to evaluate this, we have analyzed the titles recorded in the Zoological Record, Mollusca Section, to determine the proportion of malacological articles published in mala- cological serials. The results are as follows: % published Number Number in malac of malac. Year of titles serials serials 1880 181 23.2% 12 1930 1,02.3 24.7% 8 1980 3.904 26,0% 120 [The results for 1880 are based on the entire sample; those for 1930 and 1980 are each based on the average of three sub- samples of 100 entries]. It is a remarkable consistency that over the last cen- tury, about 25% of all malacological titles covered by the Zoological Record have been published in malacological journals. The malacologist is (or should be) cognizant that malacological journals are rarely read by non-mala- cologists, while non-malacological journals usually reach a wider audience. Hence it is not surprising that the bulk of malacological journals are devoted to papers on sys- tematics, evolution, and organismic-level studies; more reductionist malacological studies {e.g., biochemistry, molecular and cell biology) are invariably found in the appropriate non-malacological journals. It is also worthwhile to consider the geographic dis- tribution of malacological serials in relation to the dis- tribution of malacologists and more broadly in terms of the overall scientific literature. It is obvious that most journals are published in Europe and the United States; a more detailed analysis reveals several interesting as- pects. Altogether, thirty-one countries are represented by malacological serials. We have counted 286 serials (374 including changes in titles) of which 157 (54.9%) are still published. Of the 157 current serials, we infor- mally consider 45 (28.7%) to be professional journals and the remainder as newsletters. The countries with the most serials (and the numbers of serials — not counting changes in titles) are: USA (117); Great Britain (22); France (19); Japan (19); Australia (17); Italy (14); and Germany (13). The distribution by world region is as follows: Total ber of titles 121 6 29 39 171 374 Number of serials* 94 5 25 29 125 8 286 Number currently published 51 3 14 20 65 4 157 Region Europe (12) Africa (1) Asia (5) Oceania (4) North .America (3) Central and South America (6) [world totals] (31) Numbers in parentheses indicate the number of countries. * Not counting changes in titles. More specifically, it is obvious that no one malacologist can read, let alone make reference to, all these mala- cological serials. In his scientometric analyses. Price (1986: 67 ff.) noted that the usage of the scientific literature is a function of the inverse square law. That is, given the total number of serials in a field, half the reading will be done from the square root of that total number. Ap- plying this valuable heuristic concept to the field of mal- acology, we note that there are 157 currently published malacological serials. The square root of 157 is 12.5. Therefore, one can conclude that despite the overabun- dant diversity of current titles, only about 11-12 will account for half the total usage of malacological serials by malacologists. An informal poll ["Of the following malacological journals, which eleven do you consider to be the most important?"] of nine professional malacologists revealed a remarkable consensus of opinion: (a) 5 journals were listed by all 9 respondents; (b) 5 journals got 6, 7 or 8 votes; (c) 3 journals got 3, 4 or 5 votes; and (d) 5 journals got only 1 or 2 votes. Altogether, a total of thirteen [13] journals received at least 3 votes; this represents only 8.2% of the total number of current malacological serials. It does not seem to be essential for a malacological research library to subscribe to every current malaco- logical publication. Indeed, a compilation of the sub- scription prices for these 13 journals comes to almost $350 (at individual or member rates), which is reasonable when compared to other fields of science. No malaco- logical journals are issued by commercial (for-profit) pub- lishing houses, which are often much more expensive than institution or society journals (Feldmann, 1989). In the field of systematics, it is obviously important to have complete runs of the major journals, including those that are now defunct. It is difficult to estimate the current cost of obtaining a "complete" malacological journal li- brary; certainly in excess of $50,000 as one rarely sees these items in book dealer catalogues and there are rel- Page 42 THE NAUTILUS, Vol. 105, No. 2 atively few comprehen.sive private malacological librar- ies that are potentially available. Another element of interest is the overall numerical abundance and growth of the malacological literature, including all publication sources. Solem (1974:7-8) has estimated the size of this literature and his results are comparable to ours; Schopf (1967) provided similar cal- culations for the Ectoprocta [= Bryozoa]. Based on direct counts and estimates from the Zoological Record and Ruhoff (1980), we have compiled data on the total num- ber of malacological publications, on an annual basis. As of 1990, we have determined that approximately 167,000 such publications have appeared. The cumulative totals from antiquity to the following years are: 1875 (9,000); 1900 (19,330); 1925 (34,965); 1950 (57,115); 1975 (106,060); and 2000 (estimate 210,000). This literature is doubling every 25 years which seems rather dramatic; however. Price (1986:5-6) documented that the overall scientific literature is doubling every 15 years and certain fields even faster, thus malacology is actually increasing at a slower rate than the overall scientific literature field. A matter of considerable nomenclatural importance is the description of new species and higher taxa in news- letters or non-professional journals. All too often, these non-refereed papers are not widely available, and the type specimens are commonly not deposited into rec- ognized museums (for further discussion, see Lilllco, 1990 and Loch, 1990). We hope that the forthcoming revision of the International Code of Zoological Nomenclature will resolve these problems. In conclusion, these analyses reveal several interesting aspects of the field of malacological journals and news- letters, including their importance in the field of mala- cology and their geographic distribution. It seems that the quantity (if not quality) of malacological journals will continue to increase arithmetically (if not exponen- tially), especially as new computer desk-top publishing techniques tempt private individuals to start up their own journals. For each entry, we have included the full title along with the place(s), duration and status of the publication. For the journals and the newsletters of general interest, we have included the subscription or membership ad- dress (which is not always the same as the place of pub- lication). For shell club newsletters of only local interest, we have not included their mailing address; these are listed in Rice (1990). We also list supplements or special editions. References to cumulative indexes or other ex- ternal sources of information about the journals are in- cluded. The periodicals have been arranged alphabeti- cally within each country, according to the first word that is not an article. Major changes in title are listed separately; minor changes in wording of the subtitle are indicated by brackets, with alternate versions separated by a slash (/). Europe/Near East 42 Africa 47 Asia 48 Oceania 49 North America 50 Central and South America 57 EUROPE/NEAR EA.ST Belgium Achatina. A Newsletter of African non-marine malacol- ogy. [Tervuren; Leiden (The Netherlands)]. 1 [1970]-13 [1985]. Annales de la Societe Malacologique de Belgique. [Bru.xelles]. 1 [1863 ]-15 [1880]. (Comprising "Memoires" and "Bulletin [des Seances]".) Continued as Annales de la Societe Rovale Malacologique de Belgique. See Back- buys (1985);'Dhondt (1989). Annales de la Sociele Royale Malacologique de Bel- gique. [Bru.xelles]. 16 [1881]-37 [1902]. Continuation of Annales de la Societe Malacologique de Belgique. Con- tinued as Annales de la Societe Royale Zoologique et Malacologique de Belgique. Annales de la Societe Royale Zoologique et Malaco- logique de Belgique. [Bruxelles]. 38 [1903]-52 [1921]. Continuation of Annales de la Societe Royale Malaco- logique de Belgique. Apex. [Bruxelles]. 1 [Feb. 1986]-current. Continuation of Informations de la Societe Beige de Malacologie. |M. J. Buyle, Av. Maurice Maeterlinck 56, B-1030 Bruxelles, Belgium r Arion. Bulletin de contact [trimestriel] de la Societe Beige de Malacologie. [Bruxelles]. 1 [Jan. 1977]-current, {ad- dress: see Apex I Bulletin Mensuel d'lnformation de Malacologie Les Naturalistes Beiges [variously titled]. [Bruxelles]. 1 [1966]- 4 [1971]. Continued as Informations de la Societe Beige de Malacologie. Bulletin Periodique, Fondation Conchyliologique de Belgique/Belgische Concholiologische Vereniging/ Conchological Foundation of Belgium [Rhode-Saint- Genese]. 1 [1961]. [Single number: English, Flemish and French editions]. La Conchioline. Bulletin mensuel du Groupement Beige d'etude des Coquilles et Mollusques. [Bruxelles]. 1 [1976]- 15 [1977]. Gloria Maris. Tijdschrift uitgegeven door de Belgische Vereniging voor Conchyliologie [, Malacologie en Pa- leontologie]. [Antwerpen]. 1 [1967]-current. Also, "Me- dedelingenblad' : local members newsletter and meeting announcements, 1 [1986]-current. Miscellaneous supple- ments, "Bijvoegsel", loose pages. {Ferd. Verbieststr. 9/6, B-2030 Antwerpen, Belgium} lnforination»9 de la Societe Beige de Malacologie. [Brux- elles], 1 [Jan. 1972]-I3[Nov. 1985]. Continuation of Bul- letin Mensuel d'lnformation de Malacologie Les Natura- listes Beiges. Continued as Apex. New taxa listed by Duchamps (1986). Proces-Verbaux des Seances de la Societe Malacolo- gique de Belgique. [Bruxelles] 1 [1872]-9 [1880]. Con- R. Bieler and A. R. Kabat, 1991 Page 43 tinued as Proces-V^erbaux des Seances de la Soeiete Roy- ale Malacologique de Belgique. Largely overlaps with the "Bulletin" of the Annales de la Soeiete Royale Malacologique de Belgique; see that entry. Proces-Verbaux des Seances de la Soeiete Royale Ma- lacologique de Belgique. [Bruxelles], 10 [1881}-27 [1898]. Continuation of Proces-Verbaux des Seances de la Soeiete Malacologique de Belgique. France [for French Polynesia and New Caledonia, see under Oceania] Amenites IVlalacoIogiques par M. J. R. Bourguignat. [Paris]. 1 [1853-1856]-2 [1856-1860]. Collected reprints from Revue et Magasin de Zoologie. Annales de Malacologie. Soeiete Malacologique de France. [Paris]. 1 [1870-1884]-2 [1884-1886]. Archives Malacologiques par M. Jules Mabille. [Paris]. 1 [1867]-5 [1869]. Bibliotheque Conchyliologique (J. C. Chenu). [Paris], {serie 1) 1 [1845]-4 [1846]; (serie 2) 1 [1845]. The 6rst series is a translation of seven English-language mala- cological monographs; the second series is a translation of the malacological articles from the Transactions of the Linnean Society (London), volumes 1-17, 1791-1835. Bulletins de la Soeiete Malacologique de France. [Paris]. 1 [Jan. 1884]-7 [1890]. Contributions a la faune malacologique fran^aise (A. Locard). [Paris]. 1 [1889]-16 [1891]. [Some are reprinted from several French journals]. Elona. Bulletin de liaison de Malacologie continentale fondamentale et appliquee. A Newsletter of French non- marine Malacology. Soeiete Fran^aise de Malacologie. [Paris]. 1 [1974]-6[1980]. Haliotis. Soeiete Fran^aise de Malacologie. [Paris]. 1 [1971]-current. (Lab. Biol. Invertebres Marins et Ma- lacologie, Museum National d'Histoire Naturelle, 55 rue de Buff on, F-75005 Paris 05, France! Journal de Conchyliologie. Comprenant lEtude des [Animaux, des Coquilles vivantes et des Coquilles fos- siles/Mollusques vivants et fossiles]. [Paris; Mosman (Aus- tralia)]. 1(1) [Feb. 1850]-1 15(4) [Dec. 1979]. For dates of early issues, see Fischer-Piette (1937) and Winckworth (1936, 1937). History: Fischer (1978). Index issues: 1850- 1872 (1878); 1873-1892 (1897); 1893-1936 (1940). Dance (1986:145) erroneously stated that this journal "has con- tinued publication to this day". Magasin de Conchyliologie. [Paris]. 1 [1830]-2 [1845]. Reprints of molluscan sections of the Magasin de Zoolo- gie, d'Anatomie Comparee et de Paleontologie, Deux- ieme Section, Mollusques et Zoophytes. Mappa. Club fran^ais des collectionneurs de coquillages. [Paris]. 1 [1974]-9 [1980]. Continued as Xenophora. Miscellanees Malacologiques, par A. de Saint-Simon. [Toulouse]. 1 [1848]-2 [1856]. Le Nouveau Pes-Pelicani. [Nice]. 1 [1973]-8 [1974]. Con- tinuation of Pes-Pelicani. Nouvelles Miscellanees Malacologiques par M. le Doc- teur Paladilhe. [Paris]. 1 [1866]-4 [1869]. [Compiled re- prints from Revue et Magasin de Zoologie]. Pes-Pelicani. [Nice]. 1 [1968]-17 [1972]. Continued as Le Nouveau Pes-Pelicani. *Pyrum. Bulletin de liaison et d etudes des collection- neurs de coquillages Region Provence Cote d'Azur. [Tou- lon]. 1 [1975/76?]-7 [post 1977?]. [Issues 1-3 and 6-7 were undated; 4 and 5 dated 1977]. Revue Biographique de la Soeiete Malacologique de France. [Paris]. 1 [1870]-2 [1886]. Series Conchyliologiques comprenant TEnumeration de Mollusques terrestres et fluviatiles recueillis pendant le cours de differents voyages, ainsi que la description de plusieurs especes nouvelles (A. Morelet). [Paris] 1 [1858]-4 [1875]. Les Spicileges Malacologiques par M. J. R. Bourguig- nat. [Paris]. 1 [1860]-15 [1862]. [Mostly collected reprints, from Revue et Magasin de Zoologie]. Vertigo. Association Frangaise pour I'Etude des Mol- lusques continentaux. [Chartres]. 1 [1990]-current. {12 rue Saint-Michel, F-28000 Chartres, France! Xenophora. Bulletin du Club Franfais des Collection- neurs de Coquillages/Bulletin de I'Association Fran9aise de Conchyliologie. [Paris]. 1 [Jan. 1981]-current. Con- tinuation of Mappa. {1 impasse Guemenee, F-75004 Par- is, France} Germany [See Biirk and Jungbluth (1985) for an overall review and index of most of these journals] Abhandlungen des Archiv fiir Molluskenkunde. See en- try under Archiv fiir Molluskenkunde. Acta Conchyliorum. Club Conchylia e.V. [Darmstadt; Stuttgart]. 1 [1981]-current. Number 2 [May 1990] was also sold as a book, "News on Conidae . . .". JNeckar- anlage 6, W-6930 Eberbach am Neckar, Germany! Archiv fiir Molluskenkunde. [Frankfurt am Main]. 52(1) [Jan. 1920]-current. A supplement series, Abhandlungen des Archiv fiir Molluskenkunde, 1(1) [Jan. 1922]-2(2) [July 1929]. Continuation of Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft. | For- schungsinstitut Senckenberg, Senckenberganlage 25, \V-6000 Frankfurt-am-Main 1, Germany} Beilage zum Nachrichtsblatt. See entry under Nach- richtsblatt der Deutschen Malakozoologischen Gesell- schaft. *Club Conchylia Informationen. [Eberbach]. 1 [19 ?]- current. Continuation of the Concha. Concha. Informationen des e[ingetragenen] V[ereins] Club Conchylia. [Wilnsdorf-Obersdorf]. 1 [1969]. Continued as Club Conchvlia Informationen. Page 44 THE NAUTILUS, Vol. 105, No. 2 Conchologische Mitlheilungen. [Kassel] I [1881 ]-3 [1894], Continuation of Novitates Conchologicae. Fiir die Litteratur und KenntniO der INaturgeschichte, sonderlich der Conchylien und der Steine. [Weimar]. 1 [1781]-2 [1782]. Continuation of Journal fiir die Lieb- haber des Steinreichs und der Konchyliologie. Continued as Neue Litteratur und Beytrage zur KenntniB der Na- turgeschichte, vorziiglich der Conchylien und FoDilien. Heldia. Miinchner Malakoiogische Mitteilungen. [Miin- chen]. 1(1) [May 1984]-current. Sonderheft 1 [1987] "Land- und SiiBwassermollusken der Maltesischen In- seln", K.-H. Beckmann. {Postfach 26-01-23, W-8000 Miinehen 26, Germany f Jahrbiicher der Deutschen Malakozoologischen Ce- sellschaft. [Frankfurt am Main]. 1 [1874]-14 [1887]. Journal fiir die Liebhaber des Steinreichs und der Kon- chyliologie. [Weimar]. 1 [1773]-6 [1780]. Continued as Fiir die Litteratur und KenntniC der Naturgeschichte, sonderlich der Conchylien und der Steine. Malakoiogische Abhandlungen. Staatliches Museum fiir Tierkunde in Dresden. [Dresden]. 1 [1964]-current. | Au- gustusstraBe 2, O-8010 Dresden, Germany} Malakozoologische Blatter. [Kassel]. 1 [1854]-25 [1878]; new series 1 [1879]-11 [1891] (reprinted, 1969, Otto Koeltz, Koenigstein-Taunus). Continuation of Zeitschrift fiir Malakozoologie. Mitteilungen der Berliner Malakologen. [Berlin]. 1 [1953]-19 [1967]. Mitteilungen der Deutschen Malakozoologischen Ge- sellschaft. [Frankfurt am Main]. 1(1) [June 1962]-cur- rent. [Volume numbers discontinued after 3(36); only as issues thereafter.] Supplement 1 [1982] "Bericht iiber den ersten Workshop Malakozoologie in der Bundesrepublik Deutschland vom 09. bis 11. Oktober 1981 in Miinster i.W.". (address: see Archiv fiir Molluskenkunde( Nachrichtsblatt der Deutschen Malakozoologischen Ce- sellschaft. [Frankfurt am Main]. 1 [Dec. 1868]-51(4) [Oct. -Dec. 1919]. A supplement series, Beilage zum Nachrichtsblatt, Beitrage zur Kenntniss der mitteleuro- paischen Najadeen, 1 [1908]-4 [1910]. Continued as the Archiv fiir Molluskenkunde. Index to volumes 1-45: Ko- belt (1914). Neue Litteratur und Beytrage zur KenntniB der Natur- geschichte, vorziiglich der Conchylien und FoBilien. [Leipzig]. 1 [1784]-4 [1787]. Continuation of FUr die Litteratur und KenntniB der Naturgeschichte, sonderlich der Conchylien und der Steine. Continued as Unterhal- tungen fiir Conchylienfreunde und fiir Sammler der Mi- neralien. Novitates Conchologicae. Series prima. Mollusca extra- marina. Beschreibung und Abbildung neuer oder kri- tischer Land- und Siis.swasser Moliusken . . . (ed. L. Pfeif- fer). [subtitles varied]. [Kassel]. 1 [1854-186G]-5 [1876- 1879]. Collation: Johnson (1969). Novitates (Conchologicae. Abtheilung II; Meeres-Con- chylien. Abbildung und Beschreibung neuer Conchylien . . . (ed. W. Dunker). [subtitles varied]. [Kassel]. 1 [1858]- 16 [1878]. Supplement 1 [1862] " Monographic der Mol- luskengattung Dosinia, Scopoli {Artemis, Poli)", E. R6- mer. Supplement 2 [1853] "Index Molluscorum quae in itinere ad Guineam inferiorem collegit Georgius Tams", G. Dunker. Supplement 3 [1864-1862] "Monographic der Molluskengattung Venus Linne", E. Romer. Sup- plement 4 [1869-1874] "Japanische Meeres-Conchilien", C. E. Lischke. Supplement 5 [1874] "Ueber Vorderasia- tische Conchylien nach den Sammlungen des Prof. Haus- knecht", E. von Martens. Supplement 6 [1877]. "Clau- silienstudien", O. Boettger. Supplement 7 [1882] "Index Molluscorum Maris Japonici", G. Dunker. Collation: Johnson (1969). Schriften zur Malakozoologie aus dem Haus der Natur. [Cismar]. 1 [1989]-current. IHaus der Natur-Cismar, Hinter dem Kloster 42, W-2433 Cismar, Germany! UNITAS Malacologica Newsletter. [Tubingen]. 1 [1987]- current. {Tropenmed. Inst., Univ. Tiibingen, Wilhelm- straBe 31, W-7400 Tiibingen, Germanv ' Unterhahungen fiir Conchylienfreunde und fur Samm- ler der Mineralien. [Erlangen]. 1 [1789]. Continuation of Neue Litteratur und Beytrage zur KenntniB der Na- turgeschichte, vorziiglich der Conchylien und FoBilien. Zeitschrift fiir Malakozoologie. [Hannover; Kassel]. 1 [Jan. 18441-10(12) [Dec. 1853]. Continued as Malako- zoologische Blatter. Great Britain The British Shell Collectors' Club Newsletter. [Not- tingham]. 1 [1972]-14 [1983]. Continued as Pallidula. Index, 1982, numbers 1-50 (D. Feld). Bulletin of the Malacological Society of London. [Lon- don]. 1 [Sept. 1983]-current. Continuation of the Mala- cological Society of London Newsletter, {address: see Journal of Molluscan Studies} The Cephalopod Newsletter. [London]. 1 [1977]-current. {Dept. of Geological Sciences, LIniversity College, Gower Street, London WCIE 6BT, England} ' CIAC [Cephalopod International Advisory Council] Newsletter — Cephalopod International Advisory Board. [London; Cambridge]. 1 [1985]-current. {British Antarc- tic Survey, NERC, High Cross, Madingley Road, Cam- bridge CB3 OET, England} [Conchological Leaflets] (F. P. Marrat). [Liverpool] (1) [1876J-11 [1880/1883]. See McMillan (1961). The Conchological Society of Great Britain and Ireland. Papers for Students. [London]. 1 [1964J-18 [1981]. Conchological Society of Great Britain and Ireland Spe- cial Publication. [London]. 1 [Feb. 1980] "Marine Mol- lusca described by John Gwyn Jeffreys, with the location of the type material", A. Waren. The Conchologist: A quarterly Magazine for Conchol- ogists. [London]. 1(1) [March 1891]-1(4) [Dec. 1891]. Continued as The Conchologist: A Journal of Malacology. R. Bieler and A. R. Kabat, 1991 Page 45 Originally printed as a quarto in 58 pages; reprinted in 1903 in octavo, and repaginated with 88 pages; see Toni- lin & Fisher (1935). Index: McClelland (1923). The Conchologist: A [quarterly] Journal of Malacology. [London]. 2(1) [March 1892]-2(8) [Oct. 1893], Continu- ation of The Conchologist: A quarterly Magazine for Conchologists. Continued as The Journal of Malacology. Index: McClelland (1923). The Conchologists' Newslelter. The Conchological So- ciety of Great Britain and Ireland. [London]. 1 [1961]- current. {address: see Journal of Conchologyl The Cowry. [St. Austell, Cornwall; Port Macquarie (Aus- tralia)]. 1 [1960]-2(1) [1968]. [Vol. 1 nos. 1-3 reprinted by Philip L. Hano, New York], Digesta Malacologia. [London]. 1 [1901]-2 [1903]. Num- ber 1 is an index to the American Journal of Conchology iq.v.); number 2 is an index to the malacological articles in the Annals and Magazine of Natural History (a journal not exclusively about mollusks). Handlists of the Molluscan Collections in the Depart- ment of Zoology, National Museum of Wales. Series 1, The Melvill-Tomlin Collection. [Cardiff]. 1 [1981]-cur- rent. | National Museum of Wales, Cathays Park, Cardiff CFl 3NP, Wales} Helix. Land Shell Studies Group. [Nottingham]. 1 [1981]. The Journal of Conchology. Conchological Society of Britain and Ireland. [London]. 2(1) [Jan. 1879]-current. Continuation of The Quarterlv Journal of Conchology. Index [1879-1922, vols. 2-16]': McClelland (1923). (A. Seccombe, 36a Selbourne Road, Ilford, Essex IGl 3AJ, England 1 The Journal of Malacology. [London]. 3(1) [March 1894]- 12(4) [Dec. 1905]. Continuation of The Conchologist. Index: McClelland (1923). Journal of Molluscan Behaviour, The International Journal devoted to the Behaviour of Molluscs. [East Orkney]. 12(6):48 [1966]; only issue (a parody). The Journal of Molluscan Studies. Malacological Soci- ety of London. [London]. 42(1) [April 1976]-current. Continuation of Proceedings of the Malacological Society of London. Supplements 1 [1976], 3 [1977], 5 [1978], 6 [1978; reprinted 1988], 7 [1980], 9 [1981], 11 [1982], 15 [1985], 16 [1986], 17 [1990]: "The Prosobranch Molluscs of Britain and Denmark, Parts 1-10", V. Fretter & A. Graham. Supplement 2 [1977] "On the genus Tornatina and related forms ", E. Marcus. Supplement 4 [1977] "An annotated checklist of the western Atlantic warm water opisthobranchs ", E. Marcus. Supplement 8 [1980] "Re- vision of the North-East Atlantic bathyal and abyssal Turridae (Mollusca, Gastropoda) ', P. Bouchet & A. Wa- ren [see Bollettino Malacologico Supplemento 2, 3 for continuation]. Supplement 10 [1982] "Systematics of the genera of the Order Ascoglossa (Gastropoda)", E. Mar- cus. Supplement 12 [1983] "Catalogue of Recent and fossil Lithophaga (Bivalvia)", K. H. Kleeman. Supple- ment 12A [1983; reprinted 1988] "Proceedings of the Second Franco-British Symposium on Molluscs", ed. A. Bebbington. Supplement 13 ["1983" 27 February 1984: fide Bollettino Malacologico (1986) Supplemento 2: 561; reprinted 1988] "A generic revision of the family Eulim- idae (Gastropoda, Prosobranchia)", A. Waren. There was no Supplement 14. {D. Roberts, Dept. of Biology, Queen's Univ. of Belfast, Belfast BT7 INN, Northern Ireland) The Littorinid Tidings. Occasional Newsletter of the Littorinidae Research Group. [Manchester]. 1 [April 1974J-8 [June 1979]. The Malacological and Conchological Magazine. [Lon- don]. 1 [1838]-2 [1839]. Malacological Society of London Newsletter. [London]. 1 [1976]-16 [April 1983]. Issues 3-7 not numbered; issues 3-4 not dated [both 1977]. Continued by the Bulletin of the Malacological Society of London. *[Newsletter]. Northamptonshire Natural History So- ciety (Conchology Section). [Northampton]. ?-current. {The Humfrev Rooms, Castilian Terrance, Northampton NNl ILD, England} Pallidula. The Magazine of the British Shell Collectors' Club. [Carlisle]. 15 [1984]-current. Continuation of The British Shell Collectors' Club Newsletter. {12 Grainger Road, Isleworth, Middlesex TW7 6PQ, England) Proceedings of the Malacological Society of London. [London]. 1(1) [Oct. 1893]-41(6) [Dec. 1975]. Continued as The Journal of Molluscan Studies. Index [1893-1923, vols. 1-15]: McClelland (1923). The Quarterly Journal of Conchology. [London]. 1(1) [Feb. 1874]-1(17) [Nov. 1878]. Continued as The Journal of Conchology. Index: McClelland (1923). Species Conchyliorum. [London]. 1(1) [Nov. 1830]. [Al- though intended as an multivolume treatise, only one number was ever published]. *Subscriber's Bulletin. The Snail Centre. [Colwyn Bay]. 1 [198 ?]-current. Tentacle. lUCN/SSC Mollusc Specialist Group. [Glas- gow]. 1 [1989]-current. {Natural History Dept., Art Gal- lery and Museum, Kelvingrove, Glasgrow G3 8AG, Scot- land) The Zoological Record. Mollusca Section. [London]. 1 [1864]-current. {Biosis, 2100 Arch Street, Philadelphia. PA 19103-1399) Hungary Malakologiai Tajekoztato. Matra Miizeum. [Gyongyos]. 1 [1980]-current. {Kossuth Lajos utca 40, H-3200 Gyon- gyos, Hungary) Soosiana. [Budapest]. 1 [1973]-current. Supplement I ["1977" 1979] "A Magyarorszagi Recens Puhatestiiek El- terjedese" [= "Distribution of the Recent Mollusca of Hungary "], L. Pinter. ITermeszettudomanyi Muzeum, Baross utca 13, H-1088 Budapest, Hungary} Page 46 THE NAUTILUS, Vol. 105. No. 2 Israel .Argamanit. The Hebrew Newsletter. [Haifa; Jerusalem; Kibbutz Netzer Sereni], [1970H1976]. (Continued as Le- vantina. Argamon. Journal of the Israel Malacological Society [and the (Municipal) Malacological Museum, Nahariya]. [Haifa; Jerusalem; Kibbutz Netzer Sereni]. 1 [I970]-7(5) [1981]. Continued as Argamon, Israel Journal of Mala- cology. Argamon. Israel Journal of Malacology. [Kibbutz Netzer Sereni]. 7(6) [1988]-current. Continuation of Argamon, Journal of the Israel Malacological Society. {Kibbutz Netzer Sereni, 70-395 Israel} Levantine. A Malacological Newsletter. [Kibbutz Netzer Sereni]. 1 [1976J-53 [1984]. Continuation of Argamanit. Continued as Levantina, A Journal of Malacology. Levantina. A Journal of Malacology. [Kibbutz Netzer Sereni]. 54 [1984]-current. Continuation of Levantina, A Malacological Newsletter, [address: see Argamon] Italy Argonauta,, Journal of Malacology. Associazione Mala- cologica Internazionale (A.M. I.). [Roma]. 1 [May 1985]- current. Supplementos [1987-current] "Notiziario A. M. I." [reviews and announcements of malacological serials and books], fCasella Postale, P.O. Box 45, 00125 Acilia, Roma, Italy] Bollettino Malacologico. Societa Italiana di Malacologia. [Milano]. 15(1-2) [Jan. -Feb. 1979]-current. Continuation of Conchiglie, Notiziario Mensile della Unione Malacolo- gica Italiana. Indice specifico generale (1965-1986), 1986. Supplemento 1 [1985] "Revision of the Northeast Atlantic bathyal and abyssal Mollusca, Part 2: Neogastropoda ex- cluding Turridae", P. Bouchet & A. Waren. Supple- mento 2 [1986] "Revision of the Northeast Atlantic bathy- al and abyssal Mollusca, Part 3: Aclididae, Eulimidae, Epitoniidae", P. Bouchet & A. Waren [continuation of Journal of Molluscan Studies, Supplement 8]. Supple- mento 14 [1979]-22 [1986] "Guida illustrata all'identificazione delle Conchiglie del Mediterraneo", B. Sabelli & G. Spada. Continued from the supplements in Conchiglie. Additional miscellaneous supplements, such as membership lists. {Acquario Civico, Viale Gadio 2, 20121 Milano, Italy} Bulletino della Societa Malacologica Italiana. [Pisa]. 1 [18751-20[1897]. Continuation of Bulletino Malacologico Italiano. Both reprinted by Antiquariaat Junk, Lochem, Netherlands (1973). Index for both by Lamme (1974) Antiquariaat Junk, Lochem, Netherlands, 53 pages. Bulletino Malacologico Italiano. [Pisa]. 1(1) [Jan. -Feb. 1868]-7 [1874]. Continued as Bulletino della Societa Ma- lacologica Italiana. La Conchiglia/The Shell. Rivista [Mensile/Internaziona- le] delle Meraviglie del Mare. [Roma]. 1 [March 1969]- current. The issues of volume 1 and 2(1-4) were in Ital- ian, with an English translation distributed in mimeo- graph; number 2(5) is presumably the first "International Edition" in Italian and English, which has continued to date. Supplemento 1 [Nov.-Dec. 1970]-4 [Nov. -Dec. 1972] "Van Nostrand's Standard Catalog of Shells Month- ly Supplement", R. J. L. Wagner and R. T. Abbott [de- spite its title, these four supplements were not issued monthly with this journal]. Supplemento [Jan. 1983] "Indice Analitico Generale (1969-1982)" [supersedes several previously issued indexes]. {Via C. Federici 1, 00147 Roma, Italy} Conchiglie. Notiziario Mensile [del "Conchiglia Club"/ della] Unione Malacologica Italiana. [Milano]. 1 [Jan. 1965]-14(11-12) [Nov-Dec. 1978]. Continued as Bollet- tino Malacologico. Supplemento 1 [1969]-5 [1975], "Ca- talogo illustrato delle Conchiglie marine del Mediter- raneo", F. Ghisotti & G. Melone. Supplemento 1 [1977]- 13 [1978] "Guida illustrata all'identificazione delle Con- chiglie del Mediterraneo", B. Sabelli & G. Spada. Con- tinued as a supplement to Bollettino Malacologico. Ad- ditional miscellaneous supplements, including reprints of old papers. Indice Analitico Pluriennale 1965-1969 [1970]; 1970-1976 [1977], The Connoisseur of Seashells. [Roma]. 1 [1985]-current. {P.O. Box 561. 00187 Roma.ltaly} Giornale di Malacologia. [Pavia], 1 [1853]-2 [1854]. Lavori della Societa Italiana di Malacologia. [Milano]. 21 ["1984" 1985]-current. Continuation of Lavori della Societa Malacologica Italiana. {address: see Bollettino Malacologico} Lavori della Societa Malacologica Italiana. [Milano]. 1 [1964]-20 [1982]. Continued as Lavori della Societa It- aliana di Malacologia. [Early numbers are collected re- prints from various journals]. Malacofauna Italica. (C. Alzona). [Genova]. 1(1) 1937- 1(7) [1940]. Posthumous revised edition [1971], Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 111:1-433. Malacologia Mostra Mondiale. Associazione Picena di Malacologia. [Cupra Marittima]. 1 [1988]-current. {Via Adriatica Nord 240, 63012 Cupra Marittima, Italy} * Mostra Malacologica Messinese. Associazione Mala- cologica Messinese. [Messina]. 1 [1980?]-current? Notiziario C.LS.M.A. [Centro italiano Studi Malacolo- gici]. [Roma]. 1 [1979]-current. {Civico Museo de Zoolo- gia, via V. Aldrovandi 18, 00197 Roma, Italy} Notiziario S. L M. Pubblicazione mensile edita della Societa Italiana di Malacologia. [Milano]. 1 [1983]-cur- rent. {address: see Bollettino Malacologico} Schede Malacologiche del Mediterraneo. Societa Ma- lacologica Italiana. [Milano]. 1 [ 1 964 ]-60 [1978]. Snail Farming Research. Associazione Nazionale Elici- coltori. [Cherasco]. 1 [1986]-current. {Via Vittorio Emanuele 103, 12062 Cherasco, Italy} R. Bieler and A. R. Kabat, 1991 Page 47 Netherlands Achatina. A Newsletter of African non-marine malacol- ogy. [Tervuren (Belgium); Leiden]. 1 [1970]-13 [1985]. Basteria. Tijdschrift van de Nederlands[cli]e Malacolo- gische Vere[e]niging. [Leiden]. 1(1) [Feb. 1936]-current. Supplement 1 [1959] "The Netherlands as an environ- ment for moUuscan life", ed. W. S. S. van der Feen-van Benthem Jutting & CO. van Regteren Altena. Supple- ment 2 [198-1] "The marine Mollusca of the Bay of Al- geciras, Spain . . .", J. van Aartsen et al. Index & dates of publication, volumes 1-50 (1936-1986); Ende & Bank (1989). {National Natuurhistorisch Museum, Postbus 9517, 2300 RA Leiden, The Netherlands! Correspondentieblad van de !\ederlands[ch]e Malaco- logische Ver[e]eniging. [Amsterdam]. 1 [1934]-current. Inde.xes, published as "Bijlage van het . ..". Supple- ments [1965] "Gegevens over Mariene Gastropoden van Curacao"; [1968] "Gegevens over de Mollusken von Cu- racao uitgezonderd de Mariene Gastropoden", K. M. de Jong & I. Kristensen. See also Informatieblad. {address: see Basteria! Informatieblad van de Nederlandse Malacologische Vereniging. [Amsterdam]. 1 [1973]-current. Published as a supplement ["Bijlage bij ] to the Correspondentie- blad . . . ; numbers 2-6 also simultaneously issued as sup- plements to De Kruekel. {address; see Basteria! Jaarverslagen, Schelpenwerkgroep Friesland. [Leeu- warden]. 1 [1983]-current. { Schoenmakersperk 2, 8911 EM Leeuwarden, The Netherlands} De Kreukel, uitgave van de Malacologische Contacl- groep Amsterdam & Omstreken. [Amsterdam]. 1 [1964]- current. Seven series of special publications and supple- ments; (1) [1970-1983] "Proefschriftensupplementen", 1-5, lA-lG, II-IL, 4B, 5A-5G and 5L, A. F. J. Jansen. (2) [1983; 1988] "Jubileumnummer". (3) [1988; 1989] "Catalogus voor de Malacofilatelist", R. V. Eijkenduijn & C. Touw. (4) [1976] "Malacologische Opstellen". ed. W. Backhuys. (5) [1983] "Speciaal Polyplacophora- Nummer", H. Huneker and A. van der Sman. (6) [1984] "Bibliographia Malacofaunae Jugoslaviae", W. J. M. Maasen. (7) [1984] "Bibliography of the Land- and Fresh- water Molluscs of the Near East", Z. Bar. Index, R. Bank, 1983, De Kreukel Jubilieumnummer 1963-1983, p. 29- 44. {P.O. Box 6, 1100 Diemen, The Netherlands! The Papustyla. Manus Land Snail Society. [Lorengau, Manus Island & Woudenberg, The Netherlands]. 1 [1988]- current. Continuation of Tree and Land Snail Collectors of the World Newsletter. {P.O. Box 58, 3930 EB Wou- denberg, The Netherlands! Porphyria. Publicatie van de Werkgroep Olividae. [Delft]' 1 [1985]-2 [1986], Poland Folia Malacologia — Scientific Bulletins of Stanslaw Staszic Academy of Mining and Metallurgy. [Krakow]. 1 [1987]-current. {Inst. Geol. Sur. Miner., al. Mickiewi- cza 30, 30-050 Krak6w, Poland} Portugal Amphitrile — Publica^ao da Sec^ao de Malacologia do Centre Porlugues de Actividades Subaquaticas. [Lisboa]. 1 [1979]. Halia. Sociedade Portuguesa de Malacologia. [Lisboa]. 1 [1982]-current. {address: see Publica9oes Ocasionais!- Publica^oes Ocasionais da Sociedade Portuguesa de Malacologia. [Lisboa]. 1 [May 1982]-current. {Largo da Princesa 24-l°E., 1400 Lisboa, Portugal} Spain *Hojas Malacologicas. Coleccion de Memorias, datos y noticias sobre los rnoluscos terrestres de Espafia y Por- tugal. [Madrid]. 1 [1870-1875?]-2 [1870]. Iberus. Revista de la Sociedad Espanola de Malacologia. [Barcelona]. 1 [1981]-current. Suplemento 1 [1988] "Ca- talogo actualizado y comentado de los opistobranquios (Mollusca, Gastropoda) de la Peninsula Iberica, Baleares y Canarias, con algunas referencias a Cueta y La Isla de Alboran", J. L. Cervera et al. Suplemento 2 [1990], "Des- cripcion de nuevas especies y subespecies del genero Conus (Mollusca, Neogastropoda) para el Archipielago de Cabo Verde", E. Rolan. {Museo Nacional de Ciencias Naturales, Jose Gutierrez Abascal 2, 28006 Madrid, Spain} Resenas Malacologicas. Sociedad Espafiola de Malaco- logia. [Madrid]. 1 [1981]-current. {address; see Iberus} Switzerland Bulletin de la Societe Internationale de Conchyliologie [S.I.C.j. [Lausanne]. 1 [1979]-current. {C.P. 1323, CH- 1001 Lausanne, Switzerland! AFRICA South Africa Bulletinus Natalensis. Occasional newsletter of the Dur- ban group of the Conchological Society of Southern Af- rica. [Durban]. 1 [1983]-current. {address; see Strandlo- per[ Circular, The Conchological Society of Southern Af- rica. [Cape Town]. 1 [1958]-140 [1972]. Continued as Strandloper. Supplement 1 [1960] "Eastern Province & Natal Supplement [to the] Circular, The Conchological Society of Southern .Africa". [Durban]. INewsletler of the Conchological Society of Southern Africa. [Durban]. 1 [1989]-current. {address; see Strand- loper ! Reefcomber. [Occasional newsletter of] the Natal Shell Society. [Durban]. 1 [1979]-20 [1984]. Slug Newsletter. [Plumstead]. [198 ?]-current. [A par- Page 48 THE NAUTILUS, Vol. 105, No. 2 ody, by Giis Ferguson {30 Firfield Road, 7800 Plumstead, South Africa}]. Special Publication, Conchological Society of Southern Africa. [Kenilworth], 1 [1960]-4 [1965]. The Strandloper. Bulletin of the Conchological Society of Southern Africa. [Durban]. 141 [1972]-current. Con- tinuation of Circular, the (Conchological Society of Southern Africa, i Durban Museum, P.O. Box 4085, 4000 Durban, South Africa} ASIA China Musee Heude, Notes de Malacologie Chinoise. Universi- te I'Aurore. [Shanghai]. 1 [1934-1945]. [Issue 1(5) (1938) as "Vol. V".] Transactions of the Chinese Society of Malacology. [Bei- jing/Qingdao]. 1 [1983]-current. {Institute of Oceanol- ogy, Academia Sinica, 7 Nan-Hai Road, Qingdao, China} Japan Bulletin of the Institute of Malacology, Tokyo. [Tokyo]. 1(1) [May 1979]-current. {6-36, Midoricho-3-chome, Tanashi City, Tokyo 188, Japan! Bulletin of the National Pearl Research Laboratory. [Kashikojima]. 1 [1956]-22 [1978J Bulletin of the Nippon Institute for Scientific Research on Pearls. [Kyoto], 1 [1953]-4 [1963]. [Collected reprints from various journals]. The Chiribotan. Newsletter of the Malacological Society of Japan. [TokyoJ 1 [1960]-current. [Early numbers with- out English title], {address: see Venus} Conchologia Asiatica. [KyotoJ 1(1) [Oct. 1943]-1(4) [1943], The Conchological Magazine. [KyotoJ 1 [Nov 1907]- 4(1) [1915J [Subtitles varied; some of the articles had English translations]. *Kzohelix. Sheller's Club Ezohelix. [Horai, Wakkanai]. ?-current. Hitachiobi. [TokyoJ 1 [1974]-current. Illustrated Catalogue of Japanese Shells. [Kyoto]. 1(1) [1949]-1(25) [1953]; 2(1, 2) [1955]; series B, 1 [1955]-24 [1955 J Isokojiki. [KyotoJ 1 [1970]-current. The Japanese Journal of Malacology. 13(1-4) [Dec. 1943]-15(l-4) [March 1948J Continuation of The Venus. Continued as Venus, The Japanese Journal of Malacol- ogy JECOLN Annual Report for Japanese Expert Con- sultation on Living Nautilus. [Kamagawa]. 1976-1977 [1977]-1978-1979[1980J [Collected reprints on Nauti/us from various journalsj Kainakama. Hanshin Shell Club. [Nishinomiya-shi, Hy- ogo-kenj 1(1) [1967]-current. Material of the National Pearl Research Laboratory. [Ka.shikojimaJ I [1970]-6 [1978J Occasional Publication of the Institute of Malacology: North Pacific Shells. [Tokyo]. 1 [Nov. 1979]-current. {address: see Bulletin of the Institute of Malacology, To- kyo} Ostrea. [Hiroshima} 1(1) [1969]-current. Pacific Shell News. [TokyoJ 1 [Jan. 1970]-6[Dec. 1972J See Emerson & Sage (1987). Proceedings of the Malacozoological Meeting, "Ha- gure-Garasu" of Tokyo. [Tokyo]. 1 [1987]-current. Special Publication, Institute of Malacology of Tokyo. [Tokyo]. 1 [1985]-current. [Number 2 was on corals]. The Venus, [subtitles varied]. The Malacological Society of Japan. [KyotoJ l(l)[Nov. 1928]-12(3-4) [March 1943J Continued as The Japanese Journal of Malacology. Ap- pendix [1929-1935] "An illustrated catalogue of the Jap- anese shells", T. Kuroda. Venus. The Japanese Journal of Malacology. [Mukaishi- ma; Fukuyama; TokyoJ 15(5-8) [Oct. 1949]-current. Continuation of The Japanese Journal of Malacology. Special Publication [1979] "Check list of new genera and species; Author index; Date of publication" [volumes 1- 36, 1928-1977]. Supplement 1 [1989] "Taxonomy and distribution of moUusks", ed. T. Okutani. {National Sci- ence Museum, 3-23-1 Hyakunin-cho, Shinjuku-ku, To- kyo 169, Japan} South Korea The Korean Journal of Malacology. [Seoul]. 1(1) [Dec. 1985]-current. {Institute of Tropical Medicine, Yonsei Univ., G.P.O. Box 8044, Seoul, South Korea} Philippines Carfel. Philippine Shell News. A publication of Carfel Seashell Museum. [ManilaJ 1 [1979]-8(5) [Sept.-Oct. 1986J [Issue 6(2) erroneously labelled as "Vol. VII No. 2"J * Philippine Shell News. A Publication of the Association of Philippine Shell Collectors. [Manila]. 7 (New Series No. 1) [1972}-?. Continuation of The Philippine Shell News. *The Philippine Shell News. Association of Philippine Shell Collectors, Inc. [Quezon CityJ 1 [1958]-6 [1964?J Continued as Philippine Shell News. Taiwan Bulletin of Malacology, Republic of China. [TaipeiJ 5 [Nov. 1978]-current. Continuation of Bulletin of the Mal- acological Society of China. {Taiwan Museum, 2 Siang- Yang Road, Taipei, Taiwan} Bulletin of the Malacological Society of China. [TaipeiJ 1 [Aug. 1974J-4 [Nov. 1977J Continued as Bulletin of Malacology, Republic of China. R. Bieler and A. R. Kabat, 1991 Page 49 OCEANIA Australia *Argonauta. Malacological Society of South Australia, Inc. [Adelaide]. 1 [19 ?]-current. Australian Newsletter. Malacological Society of Austra- lia. [Sydney]. 6(20) [Jan. 1958]-15(60) [Jan. 1968]; new series, 1 [= 16(61)] [1968]-18 [July-Oct. 1972]. Contin- uation of The Malacological Society of Australia News- letter. Continued as Australian Shell News. Australian Shell News. Newsletter of the Malacological Society of Australia. [Sydney]. 1 [Jan. 1973]-current. Con- tinuation of Australian Newsletter. Supplement to No. 19[1977] "Family Mitridae Sub-family Mitrinae in West- ern Australia", G. Hanson. Supplement to No. 33 [1981] "Check Lists of the Muricidae of Queensland", O. H. Rippingale. Additional miscellaneous supplements, pri- marily membership lists, {address: see Journal of the Malacological Society of Australia! Brisbane Branch Newsletter, IVlalaeological Society of Australia. [Brisbane, Queensland]. [Feb.-March 1969]- [Dec. 1983]. [Issues not numbered sequentially]. Contin- ued as the Moreton Bay Radula. Cairns Shell News. Cairns Shell Club. [Cairns, Queens- land]. 1 [Sept. 1977]-current. Con's Conchology Newsletter of the Royal Zoological Society of New South Wales Conchological Section. [Sydney]. 1(1) [June-July 1979]-[1983]. Continued as The Sydney Sheller. The Cowry. [St. Austell (Great Britain); Port Macquarie, New South Wales]. 2(1) [1968]. [Vol. 1 published in Great Britain]. Field Naturalists' Section of the Royal Society of South Australia Conchology Club (B. C. Cotton). [Adelaide] 4 [June 1947]-5 [Jan. 1948]. Continuation of Malacolog- ical Society of South Australia Publications. Journal de Conchyliologie. [Mosman, New South Wales]. 108(1) [1970]-115(4) [December 1979]. [Previously pub- lished in Paris; see entry under "France" for further information]. Journal of the IVlalaeological Society of Australia. [Perth, Western Australia]. 1(1) [Sept. 1957]-current. Index issue (1-10), 1969. IDept. of Invertebrates, Museum of Vic- toria, 285 Russell Street, Melbourne, Victoria 3000, Aus- tralia} Keppel Bay Tidings. Keppel Bay Shell Club. [Yeppoon, Queensland], 1 [1962]-current. Malacological Club of Victoria. [Letter-Bulletin; News- letter]. [Melbourne]. 1 [1953]-4(14) [June 1956]. Contin- ued as The Malacological Society of Australia Newsletter. IVlalaeological Club of Victoria Publication (B. C. Cot- ton). [Melbourne]. 1 [July 1956]-2[Oct. 1956]. Continued as Malacological Society of Australia, Publication. The Malacological Society of Australia Newsletter. [Sydney]. 4(15) [Oct. 1956]-5(19) [1957]. Continuation of Malacological Club of Victoria Newsletter. Continued as Australian Newsletter. The Malacological Society of Australia Publication (B. C. Cotton). [Sydney]. 3 [March 1957]-4 [April 1957]. Continuation of Malacological Club of Victoria Publi- cation. Malacological Society of South Australia Publication (B. C. Cotton). [Adelaide]. 1 [Dec. 1938J-3 [Jan. 1943]. [Note; No. 3 was not on Mollusca]. Continued as Field Naturalists' Section of the Royal Society of South Aus- tralia Conchology Club. The Moreton Bay Radula. Brisbane Branch of the Mal- acological Society of Australia. [South Brisbane, Queens- land]. 1(1) [Feb. 1984]-current. Continuation of the Bris- bane Branch Newsletter. Pearl Oyster Information Bulletin. Pearl Oyster Special Interest Group. South Pacific Commission/Pacific Islands Marine Resources Information System [PIMRIS]. 1 [Feb. 1990]-current. {13 Toxteth Avenue, Austimmer, New South Wales 2514, Australia! Proceedings of the Miriam Vale Shell Club. [Miriam Vale, Queensland]. [Oct. 1955]-[April 1959]. *Proserpine Shell Club Newsletter. [Proserpine, Queensland]. ?-current. Royal Society of South Australia, Malacological Sec- tion, Publication. [Adelaide]. 1 [July 1952J-16 [1961]. Several numbers revised and reissued, not always dated. *Sunshine Coast Shell Club News. [Golden Beach, Queensland]. ?-current. The Sydney Sheller. Royal Zoological Society of NSW Conchology Section. [Sydney]. [1983]-current. Contin- uation of Con's Conchology. [Newsletter]. The Townsville Shell Club. [Townsville, Queensland]. (1) [March 1983]-(14) [1987]. Continued as Tide Watch. Tide Watch. The Townsville Shell Club. [Townsville, Queensland]. (15) [Aug. -Oct. 1987]-current. Continua- tion of the newsletter of The Townsville Shell Club. Victorian Branch Bulletin. Victorian Branch, Malaco- logical Society of Australia. [Reservoir, Victoria]. 1 [Aug. 1968]-current. [The] W. A. Shell Collector. Western Australian Shell Club. [Perth]. [1966]-current. [Originally unnumbered; Number 1 issued in December 1973]. Also a WA Branch Newsletter for members. Fiji Fiji Shell Club Annual Magazine. [Suva]. [1979]-[1987]. Shell Gossip. A monthly newsletter issued to members of the Fiji Shell Club. [Suva]. 1 [1960J-13 [1972]; new series 1 [1972]-5 [1976], Page 50 THE NAUTILUS, Vol. 105, No. 2 French Polynesia Robusta. Bulletin de liaison de 1 '.Association Fran^aise de Conchyliologie, section de Tahiti. [Papeete]. 1 [Jan.- Feb. 1990}-current. {B.P. 12030, Papara, Tahiti} La Sleeriae. Bulletin de liai.son du club des collection- neurs de coquillages de Tahiti. [Papeete]. 1 [1976]-16 [1977]. New Caledonia Rossiniana. Bulletin de I'Association Conchyliologique de Nouvelle Caledonie [N.C.]. [Noumea]. 1 [1978]-cur- rent. |B.P. 148, 18 rue Henri-Bonneaud, Noumea, New Caledonia} New Zealand Bulletin, [Auckland Museum Conchology Club/Con- chology Section of the Auckland (Institute and) Mu- seum]. [Auckland]. 1 [Nov. 1940]-17 [1961]; new series 1 [1976]-2 [1977]. Cookia. Journal of the Wellington Shell Club, New Zea- land. [Wellington]. 1 [1976]-current. Continuation of Wellington Shell Club Newsletter. News Letter [Conchology Section Auckland Institute and Museum/Auckland Museum Conchology Section). [Auckland]. [1962]-[1982]; new series, 1 [19831-current. The new series is issued as a supplement to Poirieria. Poirieria. [Conchology Section, Auckland Institute and Museum/Auckland Museum Conchology Section]. [Auckland]. 1 [1962]-current. Special Issue 1 [1980]. Tidesong. Christchurch Shell Club. [Christchurch]. 1 [Dec. 1970]-current. Also a separate Newsletter on club matters. Wellington Shell Club [Newsletter/Bulletin]. [Welling- ton]. 1 [Feb. 1967]-36 [1975]. Continued as Cookia. Whangarei Shell Club Newsletter. [Whangarei]. [Oct. 1978]-current. Journal of Cephalopod Biology. [St. John's, Newfound- land]. 1 [1989 [-current. {Ocean Studies, Memorial Univ. of Newfoundland, St. John's, Newfoundland, Canada AlC 5S7} Molluscs in Archaeology & the Recent. [Vancouver, British Columbia.] 1 [Aug. 1960]-7 [Aug. 1962]. Out of the Shell. IRDC [International Research and De- velopment Centre] Mollusc Culture Network. [Halifa.\, Nova Scotia]. 1 [Sept. 1989]-current. {Mollusc Culture Network, IRDC, Dalhousie University, Halifa.x, Nova Scotia, Canada B3H 4J II *The Vancouver and District Shell Bulletin. [Vancou- ver, British Columbia]. 1 [Jan. 1979]-current? Cuba Actas de la Sociedad Malacologica "Carlos de la Tor- re", Durante el afio [HabanaJ 1942 [1943]-1944 [1945]. *Boletin del Grupo de Estudios Malacologicos "Rafael Arango y Molina". 2 [1980]-current? Continuation of the Contribuciones Malacologicas del Grupo de Estudios Malacologicos "Rafael Arango y Molina ". Circulares de la Sociedad Malacologica "Carlos de la Torre". [Habana]. 1 [1947-1950J Continued as Circu- lares del Museo y Biblioteca de Malacologia de La Ha- bana. Circulares del Museo y Biblioteca de Malacologia de La Habana. [HabanaJ 2 [1950J-4 [1954]. Continuation of Circulares de la Sociedad Malacologica "Carlos de la Torre ". Contribuciones Malacologicas del Grupo de Estudios Malacologicos "Rafael Arango y Molina". [Habana]. 1 [1980]. Continued as the Boletin del Grupo de Estudios Malacologicos "Rafael Arango y Molina ". Revista de la Sociedad Malacologica "Carlos de la Tor- re". [Habana]. 1(1) [May 1943]-9(2) [Jan. 1954]. Index: Jacobson (1971). Papua New Guinea The Papustyla. Manus Land Snail Society. [Lorengau, Manus Island & Woudenberg, The Netherlands]. 1 [1988]- current. Continuation of Tree and Land Snail Collectors of the World Newsletter. {P.O. Box 58, 3930 EB Wou- denberg, The Netherlands} Tree and Land Snail Collectors of the World Newslet- ter. [LorengauJ 1 [1987-1988]. Continued as The Pa- pustyla. NORTH AMERICA Canada Conchologica Nonsensica. [Toronto, Ontario]. 1 [1939]. [A parody, by John Oughton]. United States of America American Conchologist. Quarterly Bulletin of the Con- chologists of America, Inc. [Louisville, Kentucky]. 15(2) [1987]-current. Continuation of Conchologists of Amer- ica Bulletin. {P.O. Box 8105, Saddle Brook, NJ 07026- 8105, USA} American Journal of Conchology. [Philadelphia, Penn- sylvania]. 1[1865]-7[1872J Indexes: Sykes (1901) Digesta Malacologica, Lviii, 46; La Rocque (1961); see also Ab- bott (1962). The dates of publication were printed on the verso of the title page of each volume and compiled by Sykes (1901). American Malacological Bulletin. [Hattiesburg, Missis- sippi; Indiana, Penns>lvania]. 1 [1983]-current. [Vol. 3(1), "December 1984" published March 1985]. Special Edi- tion 1 [1985] "Perspectives in Malacology". Special Edi- R. Bieler and A. R. Kabat, 1991 Page 51 tioii 2 [1986] "Proceedings of the second international Corbicula symposium, ed. J. C. Britten. Special Edition 3 [1986] "Proceedings of the symposium on the entrain- ment of larval oysters". Continuation of Bulletin of the American Malacological Union for Index, 198.'3-1988 (vols. 1-6), (1989) American Malacological Bulletin 6(2); 219-305, C. L. Counts. % Richard E. Petit, P.O. Bo.x 30, North Mvrtle Beach, SC 29582, LISA. The American Malacological Union, Annual Meeting , Membership List .... [Philadelphia, Pennsylvania]. (5) [1936]-(10) [1942]. Continuation of Report, _ Annual Meeting and Membership, The American Malacological Union. Continued as The American Malacological Union, News Bulletin and Annual Report, The American Malacological Union [, Inc.] [and] Amer- ican Malacological Union [, Inc.], Pacific Division, An- nual Report[s] [for] _ (20) ["1953" 1954]-37 ["1970" 1971]. [Various places of publication]. Vol- umes 22-37 officially assigned bulletin numbers. Contin- uation of The American Malacological Union, News Bul- letin and Annual Report Continued as The American Malacological Union, Inc., Bulletin for The American Malacological Union, Inc., Bulletin for (38) ["1971" 1972]-(40) ["1973" 1974]. [Phil- adelphia, Pennsylvania], Variant titles: inside front cover as "The AMU Bulletin"; the header on articles as "Bul- letin of the American Malacological Union, Inc., for ". Continuation of The American Malacological Union . . . , Annual Reports for Continued as Bulletin of the American Malacological Union, Inc. The American Malacological Union, News Bulletin and Annual Report, , Membership list. . . . [Philadelphia, Pennsylvania]. (11) ["1943" 1944]-(19) ["1952" 1953]. Continuation of The American Malacological Union, .\nnual Meeting Membership List .... Continued as The American Malacological Union .. . , Annual Reports for_. American Malacological Union [, Inc.,] Newsletter. [Various places of publication]. 1 [1968]-15(1) [1984]. [Also titled "Newsletter of the . . ." (1971)]. Continued as AMU News. Amici Castropodorum. Friends of the Gastropods News- letter. [Various places; currently at Tampa, Florida]. 1 [1976]-current. [Variously subtitled; early issues not numbered]. {Dept. of Geology, Univ. of South Florida, Tampa, FL 33260, USA \ AMU News. [Publication of the American Malacological Union]. [Ft. Pierce, Florida; Princess Anne, Maryland]. 15(2) [1984]-current. Continuation of .\merican Mal- acological Union [, Inc.,] Newsletter, {address: see Amer- ican Malacological Bulletin r *The Auger. The Austin Shell Club. [Pflugerville, Texas]. ?-current? The Beachcomber Report. Coastal Bend Shell Club. [Corpus Christi, Texas]. [April 1965]-[Nov. 1980]. [Un- numbered issues; those of Aug. 1969 to Sept. 1970 (or Dec. 1970?) were published in "Star to Star", a magazine of the Corpus Christi Junior Museum]. Continued as The Mitchelli. * Between the Tides. Gulf Coast Shell Club. [Beaumont, Texas]. ?-? [Seen: 1963-1964, with some early issues numbered {e.g., "Edition No. IX" = January 1964); 1965- June 1970; issued monthly & not numbered]. Continued (?) as the Shell and Tell. Boston Malacological Club, Inc., Newsletter. [Various places]. 1 [1969-1987]. Continued as The Epitonium. Bulletin of Paleomalacology. An International Forum for the Study of Systematics, Ecology, and Biogeography of Worldwide Fossil Mollusks. [Charlottesville, Virginia]. 1(1) [Feb. 1988]-1(4) [Dec. 1988]. Bulletin of the American Malacological Union [, Inc.] [for]. (41) ["1974" 1975]-(4S) ["1981" 1982; 5 page er- rata, 1983]. [Philadelphia, Pennsylvania]. Inside front cover, title as "The AMU Bulletin". Continuation of The American Malacological Union, Inc., Bulletin for Continued as American Malacological Bulletin. Index to years 1934-1974, (1975), 57 p. See Burch (1981) for dates and contents of issues from 1931-1966; note that several dates are incorrect therein. Bulletin of the Brooklyn Conchological Club. [Brook- lyn, New York]. 1 [1907]. The Busycon. Broward Shell Club. [Pompano Beach, Florida]. 10[1975]-current. Continuation of Busycon Bu- gle. Busycon Bugle. Publication of the Broward Shell Club. [Fort Lauderdale, FloridaJ [May, 1962]-9(8) [August 1974]. [Early issues not numbered]. Continued as The Busycon. The Capsule. The .Astronaut Trail Shell Club of Brevard, Inc. [Melbourne, Florida]. 1 [Sept. 1967]-current. Central Florida Shell News. Central Florida Shell Club. [Orlando, Florida]. [1961]-current. [Issues numbered starting with 1(1), Sept. 1980]. See also Clam Chatter. The Chambered Nautilus Newsletter. [Wilmington, Del- aware; Quincy, Massachusetts]. 1 [1974]-current. Chicago Shell Club Newsletter. [Chicago, Illinois]. 1 [1966]-19 [1984]. Continued as Thatcheria. Clam Chatter. [Orlando, Florida]. 1(2) [1980J A tem- porary variant title of Central Florida Shell News. The Collector. A Publication of the Sacramento Valley Conchological Societv. [Sacramento, CaliforniaJ 1(1) [Jan. 1955]-8(4) [Dec. 1962J *Conch Courier. North Texas Conchological Society. [Piano, Texas]. ?-current. [Earliest issue seen: January 1985]. Las Conchas. Meeting Notice and Newsletter of Pacific Shell Club, Inc. [Downey, California]. 1 [1969}-curreiit. Continuation, in part, of News of the Western Association of Shell Clubs. ♦Conchological Memoranda. [San Francisco?]. 1 [1866]- 4 [1868/9?]; 6 [1871]-13 [1875]; number 5 was not pub- Page 52 THE NAUTILUS, Vol. 105, No. 2 lished. Reprinted, in part, by Smith (1976) [but note that page 2 of number 7 was accidentally reprinted as page 2 of number 8]. Conchological INotes for [month, year]. Conchological Club of Southern (California. [Los Angeles]. [April, 1939]- [June 1941]. Issued monthly; not numbered. Continued as Minutes of the Conchological Club of Southern Cal- ifornia. The Conchologist. Isaac Lea Chapter of Conchology (No. 119) of the Agassiz Association. [Alameda, Califor- nia]. 1(1-3) [January-March, 1901]. Continuation of the Transactions of the Isaac Lea Conchological Chapter of the Agassiz Association. See Roth & Carlton (1970). Conchologist. [Houston, Texas]. 1(2) [Oct. 1964]-1(3) [Nov. 1964]. Continuation of the Publication of the Con- chology Group of the Outdoor Nature Club, Houston, Texas. Continued as the Texas Conchologist. The Conehologist's Exchange. [Philadelphia, Pennsyl- vania]. 1(1) [1886]-2(9) [March-April 1988]. Continued as The Nautilus [for index, see that entry]. Conchoiogists of America Bulletin. 1 [1974]-15(1) [1987] [\ arious places of publication]. Continued as American Conchologist. [Note: in volume 1, there were two dif- ferent issues of "Number 7"; one "Number 8" and no "Number 9"; fide American Conchologist (June 1989) 17(2): 11} Cone Shell Alert (John K. Tucker). [Effingham, Illinois]. 1(1) [Oct. 1980]-1(2) [Nov. 1980]. Contributions to Conchology (C. B. Adams; E. Chitty). [New York; Kingston, Jamaica]. 1 [1849]-13 [1853]. Convention Papers, National Shellfisheries Association. [Washington, D.C.]. (23) [1931 "1930"]-(44) [1953]. Con- tinued as Proceedings of the National Shellfisheries As- sociation. The numbers of 1931-1950 were printed on "micro cards"; 1950-1953 were issued in mimeographed form. "Volumes 1 (1909)-22 (1929)" do not exist, except as meeting handouts. See Malacological Review (1970 "1969"), 2:153, footnote 4 regarding these complexities. Corbicula Communications. [Cincinnati, Ohio]. 1 [1970]- 2 [1971]. Corbicula Newsletter. [Oak Ridge, Tennessee; Fort Worth, Texas]. 1(1) [August 1975]-9(1) [May 1985]. Crown Crier. St. Petersburg Shell Club. [St. Petersburg, Florida]. [1983]-[1988]. [Bimonthly report]. Combined with the Tidelines in 1988. The Double Helix (A Guide to the Konks of A.M.L.). Unofficial Organ of the American Mal-ecological Union- idae. Published under the auspices of the [New Orleans/ Deep South] Shell Club, New Orleans [La. /Louisiana]. [New Orleans, Louisiana]. 1(1) [July 1968]; 2(1) [July 1969]; 7(1) [July 1971]; possibly there were other issues. [A parody, by D. Dundee]. Dredgings. Pacific Northwest Shell Club, Inc. [Seattle, Washington]. 14(1) [Feb. 1974]-current. Continuation of Pacific Northwest Shell News. The Echo. Abstracts and Proceedings of the Annual Meeting of the Western Society of Malacologists. [San Francisco]. 1 [1968]-6 [1974]. Continued as Western So- ciety of Malacologists Annual Report. The Epitonium. Newsletter of the Boston Malacological Club, Inc. [Newton, Massachusetts]. 1987-current. Con- tinuation of Boston Malacological Club, Inc., Newsletter. The Festivus. [A publication of the] San Diego Shell Club. [San Diego, California]. 1(1) [Jan. 1969]-current. Sup- plement [1984] "Illustration of the types named by S. Stillman Berry in his 'Leaflets in Malacology' ", C. M. Hertz. Supplement [1987] "A faunal study of the bivalves of San Felipe and environs, Gulf of California . . .", J. Gemmell et al. Supplement [1988] "An illustrated cat- alogue of the family Typhidae Cossmann, 1903", A. D'Attilio & C. M. Hertz. Author Index, volumes 1-18 (1987), 10 pp., by T. H. Fulsbv. Continuation, in part, of News of the Western Association of Shell Clubs. {3883 Mt. Blackburn Avenue, San Diego, CA 92111, USA} *Florida Keys Shell Club. [Key West, Florida]. 1(1) [1980]-current? *Fort Myers Shell Club Newsletter. [Fort Myers, Flor- ida]. ?-? [Seen: 4, 1968; 12, 1977]. Friends of the Gastropods [Club] Newsletter. See Amici Gastropodorum. *GaIveston Shell Club [Newsletter/Quarterly]. [Galves- ton, Texas]. ?-? [Issues seen: no. 58, May 1971; no. 87, Dec. 1973; vol. 9 no. 1, Jan. 1974; vol. 9 no. 12, Dec. 1974; Winter 1976]. Gastropodia (Glenn Webb). [Everman. Texas; Kutz- town, Pennsylvania; Fleetwood, Pennsylvania]. 1(1) [Oct. 1952]-current. [Page 21 issue 2(3) "reprinted" 22 May 1984; but the original not published until 12 March 1990]. {Route 1, Box 158, Fleetwood, PA 19522, USA} Grand Strand Shell Club Newsletter. [Surfside Beach, South Carolina]. 1(1) [March 1990]-2(1) [August 1990]. Continued as Grand Strand Shell News. Grand Strand Shell News. Grand Strand Shell Club. [Surfside Beach, South Carolina]. 2(2) [October, 1990J- current. Continuation of Grand Strand Shell Club News- letter. *The Greater Tampa Shell Club, Newsletter. [Tampa, Florida]. 1 [1976]-? [Last seen April 1979]. Continued, in part, as Tampa Drill. Guam Shell News. [Agana, Guam]. 1(1) [Jan.-Feb. 1979]- 1(4) [Dec. 1979]. Hawaiian Marine Mollusks. Recent [and New] Species (C. S. Weaver). [Honolulu, Hawaii] 1 [1960]-2 [1966]. The Hawaiian Malacological Society [newsletter]. [Ho- nolulu, Hawaii]. 1(1) [March 1952]-1(8) [Oct. 1952]. Con- tinued as Hawaiian Shell News. Hawaiian Shell News. Hawaiian Malacological Society. [Honolulu, Hawaii]. 2(l)[Nov. 1953]-current. In addition to volume/number, the issues are numbered consecu- tively from number 1 of the "new series" [= 8(1)]. Sup- R Bieler and A. R. Kabat, 1991 Page 53 plement 1 [1990] "Hawaiian Epitoniidae", H. DuShane. Indexes, 1976-1981 (vols. 24-29), 1975-1984 (vols. 23- 32), 1985-1987 (vol.s 33-35), and 1985-1989 (vols. 33- 37), by Stuart Lillico, Hawaiian Malacological Society. Continuation of The Hawaiian Malacological Society [newsletter]. IP.O. Bo,\ 22130, Honolulu, HI 96823-2130, USA} Indo-Pacific IVIoIlusca. [Philadelphia, Pennsylvania; Greenville, Delaware]. 1(1) [March 1959]-3(17) [Sept. 1976]. Dual pagination scheme utilized. Irradians. [Baldwin, New York]. 7 [1980]-current. Con- tinuation of Long Island Shell Club News. The Jersey Sheller. Garden State Shell Club. [Newark, New Jersey]. 1 [1966]-2 [1967]. Johnsonia. Monographs of the marine Mollusks of the Western Atlantic. [Cambridge, Massachusetts]. 1(1) [Oct. 1941]-5(50) [July 1974]. The issues of volume 1 were individually paginated. Volume 1 reprinted 1968; vol- ume 2 reprinted 1975. Journal for IVIedical and Applied Malacology/Revista de Malacologia Medica y Aplicada. International Society for Medical and Applied Malacology/Sociedad Inter- nacional de Malacologia Medica y Aplicada. [Ann Arbor, Michigan]. 1 [1989]-current. Supplement/Suplemento 1 [1989], "Proceedings of the First International Congress on Medical and Applied Malacology/Actas del Primer Congreso Internacional de Malacologia Medica y Apli- cada", ed. J. B. Burch et al. Also a biannual newsletter. {P.O. Box 2715, Ann Arbor, MI 48106, USA| Journal of Shellfish Research. National Shellfisheries Association. [Various places; now at West Boothbay Har- bor, Maine]. 1 [1981]-current. Continuation of Proceed- ings of the National Shellfisheries Association. 1 Dr. Tom Soniat, Dept. of Biology, Univ. of New Orleans, New- Orleans, LA 70148, USA} *Junonia. Sanibel-Captiva Shell Club. [Sanibel, Florida]. ?-current. Continuation of Sanibel-Captiva Shell Club Newsletter. The Laddergram. Palmetto Shell Club, Inc. [Columbia, South Carolina]. 1 [Sept. 1987]-current. Leaflets in Malacology. [Redlands, California]. 1(1) [Nov. 1946]-1(26) [Dec. 1969]. Illustrations of type specimens. Hertz (1984); see Petit (1988) regarding two versions of number 13 [printings of 7 and 9 July, 1956 of which the former was suppressed]. The Littorina. The Louisville Conchological Society. [Louisville, Kentucky]. 1 [1975]-ll(l)[Jan.-March 1986J Long Island Shell Club News. [Flushing, New York; Baldwin, New York]. 1 [1975]-6 [1980J Continued as Irradians. Louisiana Malacological Society Occasional Papers. [Baton Rouge, Louisiana]. 1 [1988]. Low Country Shell Club Newsletter. [Charleston, South Carolina! 1(1) [May, 1979]-2(7) [Sept. 1980} [Volume 1 comprised numbers 1-9 and 12; there were no numbers 10 or llj Malacologia. [Ann Arbor, Michigan; Philadelphia, Penn- sylvania]. 1(1) [Nov. 1962]-current. Dates of publication given on the inside back cover of 11(2) and with the index of later volumes. History: Burch & Huber (1968); Burch (1973). I Malacology Department, The Academy of Natural Sciences, 19th & The Parkway, Philadelphia, PA 19103, USAl Malacological Review. [Ann Arbor, Michigan; Boulder, Colorado]. 1 [June 1969 "1968"]-current. Dates of pub- lication are given on page ii of the latest volume. Sup- plement 1 [1973] "The freshwater snails of Taiwan (For- mosa)", G. L. Pace. 2 [1980] "The Meking Schistosome", ed. J. I. Bruce et al. Supplement 3 [1982] "A dissection guide, field and laboratory manual for the introduced bivalve Corbicula fluminea" , J. C. Britton & B. Morton. 4 [1988] "Prosobranch Phylogeny", ed. W. F. Ponder et al. History: Burch (1974 "1973"). {P.O. Box 637, Niwot, CO 80544, USA} Malacology Data Net [Ecosearch Series]. [Portland, Tex- as]. 1(1) [April 1986]-current. [The cover of issue 2(1/2) was erroneously labelled as "Vol. 2 No. 1"]. {325 E. Bay view, Portland, TX 78374, USA} Marginella Marginalia. [Dayton, Ohio]. 1(1) [March 1986}-current. {Dayton Museum of Natural History, 2629 Ridge Avenue, Dayton, OH 45414, USA! Members of the American Malacological Union. [Phil- adelphia, Pennsylvania]. (1) [1932]. Continued as Mrs. Imogene C, Robertson's rambling notes. [An account of the first meeting appeared in the Nautilus, 45(l):l-5 (1931).] Miami Malacological Society Quarterly. [Miami, Flor- ida]. 1(2) [1967]-5(4) [1972J Continuation of Quarterly Report Miami Malacological Society. Minutes [number, date] Conchological Club of Southern California. [Los Angeles]. 1 [July, 1941]-200[June, 1960J Continuation of Conchological Notes for ... . See Coan (1976) regarding status of the new taxa proposed in this serial. The Mitchelli. Coastal Bend Shell Club. [Corpus Christi, Texas]. 1(1) [Dec. 1980]-current. Continuation of the Beachcomber Report. Mollusca. [Tavares, Florida]. l(l)[Sept. 1944]-2(6) [Aug. 1949]. The Mollusca. Off the Beach Notes. Shell Club of the Ryukyu Islands. [Ryukvu/APO San Francisco]. 1(1) [1960]-2(8) [Dec. 1961]. [Although from Japan, this En- glish-language newsletter was published by U.S. Armed Services personnel and their families stationed in Ryu- kyu]. Molluscan Digest. The International Publication for Malacological Research. [Pismo Beach, California; Ox- nard, California} 1(1) [Jan. 1971]-4(l-2) [Sept. 1976]. The Mollusk. Greater Miami Shell Club [previously as: South Florida Shell Club]. [Miami, Florida]. 14(7) [1976]- current. Continuation of South Florida Shell Club Mol- lusk Chaser. ?a«;e 54 THE NAUTILUS, Vol. 105, No. 2 Mollusk-Chaser South Florida Shell Club. [Miami, Flor- ida]. 5(4) [April 1967]-7(7) [July 1969]. Continuation of The South Florida Shell Club's Newsletter Mollusk- Chaser. Continued as the South Florida Shell Club Mol- lusk Chaser. IVIonographs of Marine Mollusca. [Greenville, Dela- ware; Melbourne, Florida; Silver Springs, Maryland]. 1 [Dec. 1978]-current. [Early numbers (1-3) followed the dual pagination scheme of the Indo-Pacific Mollusca and the Standard Catalogue of Shells]. ! P.O. Box 7279, Silver Spring, MD 20907-7279, USA} Mrs. Imogene C. Robertson's Rambling Notes on the Second Annual Meeting of the American Malacological Union in Washington, D.C. May 26-28, 1932. [Buffalo, New York]. (2) [1932]. Continuation of Members of the American Malacological Union. Continued as Report, Annual Meeting and Membership, The American Mal- acological Union. [See also the Nautilus, 46(1): 1-3 (1932); an account of the third meeting appeared in the Nautilus, 47(l):37-44 (1933)]. Naples Shell News. Naples Shell Club, Inc. [Naples, Flor- ida]. 1 [1966]-current. [Sporadic numbering of volumes and issues; "The Shell Case " a temporary variant title, q.v.]. *I\ational Shellfisheries Association Quarterly News- letter. [Port Norris, New Jersey]. ? [October 1989]-cur- rent. The Nautilus. [Philadelphia, Pennsylvania; Greenville, Delaware; Melbourne, Florida; Silver Spring, Maryland]. 3(1) [May 1889]-current. [Volumes 3-30 with 12 num- bers each; subsequent volumes with four numbers each]. Supplement 1 [ 1990] "Catalogue of the Superfamily Can- cellarioidea Forbes and Hanley, 1851 (Gastropoda: Pros- obranchia", R. E. Petit & M. G. Harasewych. Continu- ation of the Conchologist's E.xchange. Indexes: (1) "Index to The Nautilus, vols. 3 to 34, 1889-1921 and to its predecessor. The Conchologist's Exchange, vols. 1, 2, 1886-1888" by J. B. Hender.-on et ai, 1927, 434 p. (pub- lished by G. M. Clapp & B. Walker). (2) "Index to The Nautilus, volumes 35 to 60, 1921-1947" by A. LaRocque et ai, 1951, 332 p. (published by Univ. of Michigan Press, Ann Arbor). (3) "Author Index to The Nautilus, vols. 3- 75 ..." by A. LaRocque, 1963, 279 p. (Columbus, Ohio). (4) "Author Index to The Nautilus, vols. 76-85" by A. LaRocque, 1972, Sterkiana 47:21-38. (5) Inde.xes to The Nautilus: Geographical (vols. 1-90) and scientific names (vols. 61-90)" by R. T. Abbott et al.. 1979, iv + 238 p. (American Malacologists, Inc., Melbourne, Florida). An anthology: Abbott (1976). Dating of the first 75 volumes best accomplished by reference to Clench & Turner (1962). {P.O. Box 7279, Silver Spring, MD 20907-7279, USA} NCMC Newsletter. Northern California Malacozoologi- cal Club. [Novato]. 1 [1988]-current. Continuation of the Northern California Shell C:lub News. [Newsletter]. National Capital Shell Club. [Washington, D.C.]. 1 [1974)-current. [Newsletter]. New Jersey Shell Club [May wood. New [ Jersey; Park Ridge, New Jersey]. 1973 [October]-1977 [May /June]. Newsletter, Marco Island Shell Club, Inc. [Marco Island, Florida]. 1 [1982]-current. The [month] Newsletter of the Jersey Cape Shell Club [Stone Harbor, New Jersey]. [May, 1982]-current. News of the Associated Shell Clubs of California. [Northern California Malacozoological Club, Berkeley; Pacific Shell Club, Los Angeles; Conchological Club of Southern California, Los Angeles]. 1(1) [May 1960]-2(1) [July 1961]. Continuation, in part, of the Proceedings [Minutes] of the Pacific Shell Club. Continued as News of the Western Association of Shell Clubs. News of the Western Association of Shell Clubs. [As above, with the San Diego Shell Club, San Diego]. 2(2) [Oct. 1961]-10(4) [1969]. [Issue 7(1) (1966) erroneously labelled as "Vol. 6 No. 7 "]. Continuation of News of the Associated Shell Clubs of California. Continued, in part, by The Festivus and Las Conchas. New York Shell Club Notes. [New York]. 1 [1950]-cur- rent. [Department of Invertebrates. American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA} The Noble Pen. Newsletter of the Cen.[tral] Penn.[sylvania] Beachcombers. [Elizabethtown, Pennsyl- vania; Lancaster, Pennsylvania]. 1(1) [Nov. 1983]-cur- rent. [The first two issues were untitled]. North Carolina Shell Club Bulletin. [Morehead City, North Carolina]. 1 [1963J-9 [1978]. Northern California Shell Club News. Northern Cali- fornia Malacozoological Club. [Novato]. 1 [1978]-10 [1987]. Continued as NCMC Newsletter. Note: The Ve- liger, volumes 1-6 [1958-1963], was originally published as the newsletter of this club. Northwest Shell News. [Poulsbo, Washington]. 1(1) [Jan. 1961]-1(5) [Oct. 1961]. Continued as Pacific Northwest Shell News. Notes and Quotes. Rochester Shell and Shore Club. [Rochester, New York]. 1 [.\ugust 1967]-66 [November 1980]. Occasional Papers on Mollusks. Department of Mol- lusks, Museum of Comparative Zoology, Harvard Uni- versity. [Cambridge, Massachusetts]. 1(1) [Feb. 1945]- current. {Museum of Comparative Zoology, Harvard Universit\. Cambridge, MA 02138, USA} Of Sea and Shore. [Port Gamble. Washington]. 1 [1970]- current. Supplements, [1971] 'Dictionary of English- French French-English shell terms ", C. Geerts; [1971] "Identification Guide: Cow ries of the Red Sea Area", J. O'Malley. { P.O. Box 219. Port Gamble, WA 98364, USA} Opisthobranch. .\ monthh publication on mollusks and marine life. (Phoenix, Arizona]. 16(1-4) [Jan-April 1984]. Continuiition of Opisthobranch Newsletter. Continued as Shells and Sea Life. Opisthobranch Newsletter. [Pismo Beach, California]. 1 1969]-15 [1983]. Contiiuied as Opisthobranch. R. Bieler and A. R. Kabat. 1991 Page 55 Oregon Shell News. Oregon Societ\ of Conchologists. [Portland. Oregon]. 1(1) [1970]-current. [1(1) labelled as "new series"]. The Original Tide-ings. Indiana's first shell club. [Crown Point, Indiana]. 3(7) [July 1973]-6(1) [Jan.-March 1976]. See discussion under Tide-ings. Pacific Northwest Shell News. [Poulsbo. Washington]. U6) [1961]-13(5) [Oct. 1973]. Continuation of Northwest Shell News. Continued as Dredgings. Palm Beach County Shell Club News. [Lake \\ orth. Florida]. [1961]-[1961]. Continued as Seafari. The Pariah. [Highstown. New Jersey]. 1[1977]-8[1980]. Pittsburgh Shell Club Bulletin. [Pittsburgh. Penns\l- \ania]. (1) [1966]-(12) [1977] [most issues not numbered]. Proceedings of the .National Shellfisheries .\ssociation. [Duxbury, Massachusetts} 45 [1955 "1954"]-70 [1980]. Continuation of Convention papers. National Shellfish- eries -Association. Continued as Journal of Shellfish Re- search. Inde.x, "Titles of papers presented at annual meet- ings. 1930-1973". ii + 37 pages Proceedings of the Pacific Shell Club. [Los -\ngeles]. 1 [1955]. .\lso issued as irregular "Minutes" and ".\nnual Report" of the Pacific Shell Club; these publications were only distributed at meetings and not mailed. Continued, in part, b\ the News of the .Associated Shell Clubs of California. Proceedings of the Philadelphia Shell Club. [Philadel- phia. Pennsylvania} IJ) [1955}-1(9! [1964'. Proceedings of the Society for Experimental and De- scriptive Malacology. Earlier reports published in Mal- acological Review 2(1). 5(1), 6(1). 7(1) and 8(1-2). Later reports published in Walkerana (q.v.). Publication of the Conchology Group of the Outdoor .Nature Club. Houston. Texas. [Houston. Texas]. 1(1) [Sept. 1964]. Continued as the Conchologist. Quarterly Report Miami Malacological Society. [Miami, Florida]. 1(1) [1967]. Continued as Miami Malacological Society Quarterly. Report. .\nnual Meeting and Membership. The .\merican Malacological I'nion. [Philadelphia, Penn- sylvania]. (3) [1934 J-i4) [1935]. Continuation of Mrs. Im- ogene C. Robertson s Rambling Notes ... Continued as The American Malacological Union. .\nnual Meeting , Membership List .... *\ Review of the Broward Shell Club. [Pompano Beach, Florida]. 1(1! [1963]-^ [last issue seen: 1(3). 1965]. Revista de Malacologia Medica y .\plicada: see entrv under Journal for Medical and Applied Malacology . *Sanibel-Captiva Shell Club Newsletter. [Sanibel Island, Florida]. "^-^ [seen: 8 ^Nov. 1965); 9 (June 1966)]. Con- tinued as the Junonia. *San Marcos Shell Club Quarterly Newsletter. [San Marcos, Texas]. 1 [1972]-^ [last issue' seen: 5(4). 1976]. Sarasota Shell Club Newsletter. [Sarasota, Florida]. [1982]-current. The Sayana. South Carolina Shell Club Newsletter. [Charleston, South Carolina]. [April 1989]-current. [1989 issues not numbered]. Seafari. Palm Beach County Shell Club. [Lake Worth; \\est Palm Beach, Florida]. [i961]-current. Continuation of Palm Beach Count\ Shell Club News. First numbered issue is "6(12)" [Dec' 1964]; issues 9(3-8) [1967J erro- neousK- labelled as volume 8. The Searcher. New sletter of the Sea Shell Searchers of Brazoria Count\. [Lake Jackson, Texas], [1977]-current. *Shell and Tell. Newsletter of the Gulf Coast Shell Club. [Beaumont. Texas]. ?-currenti^ [onl\ two issues seen: 4(1), 1984; 5(1) 1985]. Continuation of Between the Tides? *The Shell Case. Naples Shell Club. [Naples, Florida]. [1971]-[1977:^]. [Seen: 1971, 2(1), 3(1), 4(1); a temporary variant title of the Naples Shell News, q.v.]. Shell Collector. [Fort Lauderdale, Florida]. Premiere [1978]; 1 [1978]-2 [1979]. The Shelletter of Shells and their Neighbors. [Redlands, California] 1 [1960]-24 [1964], Shell Museum & [Research/Educational] Foundation, Inc.. Newsletter. [Sanibel, Florida} 1 [1988]-current, Shell Notes (Frank L>Tnan). [Lantana, Florida]. 1(1) [1942]-2( 13-151 [June 1951} [The] Shell-O-Gram. Jacksonville Shell Club, Inc. [Jack- sonville, Florida], 1 [1966]-current. [\'arious issues have erroneous \olume and number labels]. Shells and Sea Life. [Phoenix, ,\rizona; Bayside, Cali- fornia} 16(5) [May 1984]-current. Continuation of Opis- thobranch. [\arious changes in format and subtitles; vol. 18(3), 1986, announced a change to three separate series: (1) a monthly "Shells and Sea Life News"; (2) a "Shells and Sea Life Quarterly"; and (3) a new technical journal ".\cta Mollusca". This plan was apparentK abandoned]. Supplement [1986] "Directorx of Clubs & Services". U701 Hyland Street, Bayside, Ca'95524, USA} *The Shell Sheet. Reef Roamers Shell Club [formerlv. The .\nderson Reef Roamers]. [Yigo, Guam} 1 [19 ?]- current. The Slit Shell. [Newsletter of the] Indianapolis Shell Club. [Indianapolis, Indiana} [1977}-current. [Some is- sues numbered]. The Slug. Perrine Muckulogical Society. [Miami, Flor- ida} 1(1) [1 April 1989} [A parody} South Florida Shell Club [incorporated/Inc] MoUusk Chaser. [Miami, Florida} 7(8^ [August 1969]-14(6) [1976} Continuation of the Mollusk-Chaser South Florida Shell Club. Continued as The MoUusk. [Various changes in format and style; some issues of volume 9 (1971) labelled as "new series"; December issue of volume 10 (1972) labelled as "Supplement"]. South Florida Shell Club Newsletter. [Miami, Florida} 1 [1963]-2(5) [1964} Continued as The South Florida Shell Club's Newsletter Mollusk-Chaser. The South Florida Shell Club's Newsletter Mollusk- Page 56 THE NAUTILUS, Vol. 105, No. 2 Chaser. [Miami, Florida]. 2(6) [1964}-5(3) [March 1967]. Continuation of the South Florida Shell CAuh Newsletter. Continued as Moilusk-C^haser South Florida Shell Club. *South Padre Island Shell Club [Newsletter]. [South Padre Island, Texas]. ?-? [seen; issues from 1962-Oct. 1980]. ♦Southwest Florida Shell News. Journal of the Southwest Florida Conchologist Society, Inc. [Fort Myers, Florida]. ?-current. Southwestern Malacological Society, Monthly Meeting Notice and Newsletter. [Phoenix, Arizona]. [1966]-cur- rent. Special Occasional Publication. Department of Mol- lusks. Museum of Comparative Zoology, Harvard Uni- versity. [Cambridge, Massachusetts], 1 [1973]-current. {Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA} Sterkiana. [Columbus, Ohio; Shawsviile, Virginia]. 1 [Nov. 1959]-71 [1981]. Suncoast Shorelines. Suncoast Conchologists. [Palm Harbor, Florida]. 1 [1984]-current. Tabulata. [Santa Barbara, California]. 1(1) [July 1967]- 7(4) [Oct. 1974]. Index (1974), 14 p. *Tanipa Drill. The Greater Tampa Shell Club. [Tampa, Florida]. ?-? [earliest issue seen: 1(3), 1972; latest issue seen: May 1977 labelled as "volume II first issue"]. Con- tinuation, in part, of The Greater Tampa Shell Club, Newsletter. Texas Conchologist. Houston Conchology Society. [Houston, Texas]. 1(4-6) ["Dec. 1964-Jan./Feb. 1965"]- current. [Also a newsletter for meeting announcements]. Continuation of Conchologist. {3706 Rice Boulevard, Houston, TX 77005, USA 1 Texas Shell News of the San Antonio Shell Club. [San Antonio, Texas]. 1 [1958]-15 [1972J ["Vol. VII" (1964) erroneously also used for 1965 and 1966]. Thalcheria. [Chicago]. 19 [1984]-current. Continuation of Chicago Shell Club Newsletter. *Tibia Trumpet. Broward North Shell Club. [?, Florida]. 1 [1973]-2 [I974J [No further issues?]. Tide-ings. The Newsletter of the Crown Point Shell Col- lectors' Study Group, Inc. [Crown Point, Indiana]. 1 [ 197 1 ]-current. See also The Original Tide-ings; the latter was started due to an editorial dispute within this shell club. Tide Lines. St. Petersburg Shell Club [, Inc.]. [St. Pe- tersburg, Florida]. [1970]-current. Continuation of an annual (uiuiamed) St. Petersburg Shell Club show sou- venir booklet [1965-1969]. Combined with the Crown Crier in 1988. Tide Lines. Treasure Coast Shell Club, Inc. [Palm Citv, Florida]. 1(3) [1987]-2(2) [1988]. Continued as Treasure Coast Shell News. Note: this newsletter started in March 1986, but 1(1) and 1(2) were unnamed. The Tidewater Triton. The Tidewater Shell and Fossil Club. [Tidewater, Virginia]. 1 [1969J-4 (1973) [some is- sues of volumes 3 (1972) and 4 (1973) erroneously la- belled as "IV ' or "VI", respectively]. ♦Transactions of the Isaac Lea [Conchological] Chapter of the Agassiz Association. 1 [1889?]- 10? [1899?]. This ".serial" actually comprised members' reports, which were annually compiled into a single (?) bound manuscript volume, and circulated from one member to the next during the following year. Extracts were published in various journals, including The Nautilus, usually titled as the "Isaac Lea Department ". We have not been able to locate any extant set(s) or issues of the Transactions (including the copy mentioned by Keen, 1954). See Coan (1989) for further discussion. Continued as The Con- chologist. Transactions of the POETS Society. See Walkerana. Treasure Coast Shell News. Treasure Coast Shell Club, Inc. [Palm City, Florida} 2(3) [1988]-current. Continu- ation of Tide Lines (q.v.) [the St. Petersburg Shell Club complained of the theft of their title, so 2(3) was given the current name]. Tryonia. Miscellaneous Publications of the Department of Malacology of the Academy of Natural Sciences of Philadelphia. [Philadelphia, Pennsylvania]. 1 [March 1979]-current. {address: see Malacologia] ♦[newsletter]. Utah Shell Society. [Salt Lake City, UtahJ ?-current. The Veliger. [Berkeley; Davis, California]. 1(1) [June 1958]-current. Supplements (not numbered separately): Supplement [Vol. 3, 1961 ]"Opisthobranchmollusks from California", E. Marcus. Supplement [Vol. 3, 1968] "The Order Anaspidea ", R. D. Beeman; and "Thecosomata and Gymnosomata", J. A. McGowan. Supplement [Vol. 6, 1964] "The biology of Tegula funebralis (A. Adams, 1855)", ed. D. P. Abbott et al. Supplement [Vol. 7, 1965] "A glossary of a thousand-and-one terms used in con- chology", W. H. Arnold. Supplement [Vol. 11, 1968] "The biology of Acmaea", ed. D. P. Abbott et al. Sup- plement [Vol. 14, 1971] "The Northwest American Tel- linidae", E. V. Coan. Supplement [Vol. 15, 1973] "A systematic revision of the Recent cypraeid [sic] family Ovulidae (Mollusca: Gastropoda)", C. N. Gate. Supple- ment [Vol. 16, 1974] "The Panamic-Galapagan Epito- niidae ", H. DuShane. Supplement [Vol. 17, 1974] "Growth rates, depth preference and ecological succession of some sessile marine invertebrates in Monterey Harbor ", E. C. Haderlie. Supplement [Vol. 18, 1975] "The biology of chitons", ed. R. Burnett et al. Supplement (new series) 1 [1990] "Catalogue of Recent and fossil molluscan types in the Santa Barbara Museum of Natural History. I . . . ", by P. H. Scott et al. History, Keen (1983) and Gate (1989); index to opisthobranch taxa, Carlton & Behrens (1981). {Museum of Paleontology, 3 Earth Sciences Building, Univ. of California, Berkeley, CA 94720, USA) Walkerana. Transactions of the POETS Society. [Whit- more Lake, Michigan; Niwot, Colorado; Ann Arbor, Michigan]. 1 []980]-current. [Volume 1(1), undated (ap- R. Bieler and A. R. Kabat, 1991 Page 57 parently 1980?), was also designated as the "Transactions of the Physiological, Oecological, Experimental Taxo- nomic and Systematic Society . . . from the Proceedings of the Society for Experimental and Descriptive Mala- cology ]. There are discrepancies in the dates and order of several issues: 1(5) ["1982" 30 December 1983]; 2(6) ["1988" 6 June 1989]; 2(7) [31 December 1984]; 2(8) ["1987" 29 April 1988]; 3(10) ["1989" 15 January 1990]. Issues are numbered consecutively independent of the volume numbers. {P.O. Box 2701, Ann Arbor, MI 48106, USA} West American MoIIusca. [San Diego, California]. 1 [1900-1902], See Coan (1966) for dates of publication. Western Society of IVlalacologists, Annual Report. [Los Angeles, California]. 7 [1974]-current. Continuation of Echo. {1633 Posilipo Lane, Santa Barbara, CA 93018, USA} Western Society of IVlalacoIogists, Occasional Paper. [Los Angeles, California]. 1 [1975]-2 [1976]. The Whelk Wavelength. The Georgia Shell Club, Inc. [Marietta, Georgia]. 1 [1985]-current. The Xenophora Newsletter. [Van Nuys, California]. 1 [Jan. 1976]-7 [Dec. 1976]. The Young Shell Collectors Quaterly. International League of Young Shell Collectors. [Lima, Peru; Arling- ton, Massachusetts]. 1 [1967]-2 [1969]. CENTRAL AND SOUTH AMERICA Chile Sociedad Chilena de Malacologia Comunicaciones. [Vifia del Mar], 1 [1979]. Brasil Informativo S.B.lVl. [Sociedade Brasileira de Malaco- logia]. [Porto Alegre; Sao Paulo]. 1 [1969]-current. {Inst, de Biociencuas, Univ. Sao Paulo, Caixo Postale 20520, Sao Paulo, Brasil} Siratus. Periodico Bimensal dos Conquiliologistas do Brasil/Bimonthly Journal of [the] Conquiliologistas do Brasil. [Sao Paulo]. 1 [Nov.-Dec. 1989]-current. {C. P. 15011, Sao Paulo/SP, Brasil 01599} Panama Donax Panamensis. Sociedad Panamena de Malacolo- gia. [Panama]. 1 [Feb. 1981]-58/59 [Nov.-Dec. 1985]. Thais. Publicaciones diversas de la Sociedad Panamefia de Malacologia. [Panama]. 1 [29 June 1981]-4 [17 January 1983]. Peru The Young Shell Collectors Quarterly. International League of Young Shell Collectors. [Lima; Arlington, Mas- sachusetts (USA)]. 1 [1967]-2 [1969]. Uruguay Comunicaciones de la Sociedad Malacologica del Uru- guay. [Montevideo]. 1 [Sept. 1961]-current. {Casilla de Correo 1401, Montevideo, Uruguay} Venezuela *Bollettino informativo. Sociedad Venezolana de Malacologia. [Caracas]. 1 [198 ?]-current. INDEX The purpose of this index is to guide the reader to the appropriate country for the malacological journals and newsletters listed herein, especially those that do not have (well known) geographical indication(s) in their title. Long titles have been abbreviated or truncated to save space. Abhandlungen des Archiv Achatina Acta Conchyliorum Actas de la Sociedad Amenites Malacologiques American Conchologist American Journal of Conchology American Malacological Bulletin American Malacological Union Amici Gastropodorum Amphitrite AMU News Annales de Malacologie Annales de la Societe Malacologique Annales de la Societe Royale Apex Archiv fiir Molluskenkunde Archives Malacologiques Argamanit .\rgamon .Argonauta ■Argonauta .Arion Auger, The Australian Newsletter Australian Shell News Basteria Beachcomber Report, The Beilage zum Nachrichtsblatt Between the Tides Bibliotheque Conchyliologique Boletin del Grupo de Estudios Bollettino Informativo Bollettino Malacologico Boston Malacological Club Brisbane Branch Newsletter British Shell Collectors' Club Bulletin, Auckland Museum Bulletin de la Societe Inter. Bulletin de liaison de Malac. [Elona] Bulletin Mensuel d Information Bulletin of Malacology Bulletin of Paleomalacology Bulletin of the American Malac Union Germany Belgium/Netherlands Germany Cuba France USA USA USA USA USA Portugal USA France Belgium Belgium Belgium Germany France Israel Israel Australia Italy Belgium USA Australia Australia Netherlands USA Germanv USA France Cuba Venezuela Italy USA Australia Great Britain New Zealand Switzerland France Belgium Taiwan USA USA Page 58 THE NAUTILUS. Vol. 105. No. 2 Bulletin of the Brooklyn Conch. Club Bulletin of the Inst, of Malacology Bulletin of the Malac. Soc. of China Bull, of the Malac. Soc. of London Bull, of the Natl. Pearl Res. Lab. Bulletin of the Nippon Institute Bulletin Periodique, Fondation Conch. Bulletino della Societa Malac Italiana Bulletino Malacologico Italiano Bulletins de la Sociele Malac. de France Bulletinus Natalensis Busycon Bugle Busycon, The Cairns Shell News Capsule, The Carfel Central Florida Shell News Cephalopod Newsletter, The Chambered Nautilus Newsletter, The Chicago Shell Club Newsletter Chiribotan, The ClAC Newsletter Circular, The Conchological Society Circulares de la Sociedad Malacologica Circulares del Museo y Biblioteca Clam Chatter Club Conchylia Informationen Collector, The Comunicaciones de la Sociedad Malac. Conch Courier Concha Conchas, Las Conchiglia, La Conchiglie Conchioline, La Conchologia Asiatica Conchologica Nonsensica [parody] Conchological Leaflets Conchological Magazine, The Conchological Memoranda Conchological Notes Conchological Society Conchologi.sche Mittheilungen Conchologist, The Conchologist, The C'onchologist C-"onchologist"s Exchange, The Conchologists" Newsletter, The Conchologists of America Bulletin Cone Shell Alert Connoisseur of Seashells, The Con's (^onchology Contribuciones Malacologicas Contributions a la faune malacologique Contributions to Conchology Convention Papers, Natl. Shellfish. Assoc. Cookia Corbicula Communications Corbicula Newsletter Correspondentieblad Cowry, The Great Crown Oier Dige.sta Malacologica Dona.x Panamensis Double Helix, The [parody] USA Dredgings USA Japan Echo, The USA Taiwan Fiona France Great Britain Epitonium, The USA Japan Ezohelix Japan Japan Festi\ us. The USA Belgium Field Naturalists' Section Australia Italy Fiji Shell Club .Annual Magazine Fiji Italy Florida Keys Shell C:lub USA France Folia Malacologia Poland South Africa Fort Myers Shell Club Newsletter USA USA Friends of the Gastropods USA USA Fiir die Litteratur und KenntniB Germany Australia Galveston Shell Club USA USA Gastropodia USA Philippines Giornale di Malacologia Italy USA Gloria Maris Belgium Great Britain Grand Strand Shell Club Newsletter USA USA Grand Strand Shell News USA USA Greater Tampa Shell Club USA Japan Guam Shell News USA Great Britain Halia Portugal South Africa Haliotis France Cuba Handlists of the Molluscan Coll. Great Britain Cuba Hawaiian Malacological Society USA USA Hawaiian Marine Mollusks USA Germany Hawaiian Shell News USA USA Heldia Germany Uruguay Helix Great Britain USA Hitachiobi Japan Germany Hojas Malacologicas Spain USA Iberus Spain Italy Illustrated Catalogue of Japanese Shells Japan Italy Indo-Pacific Mollusca USA Belgium Informatieblad Netherlands Japan Informationen des Club Conchylia [Concha] Germany Canada Informations de la Societe Beige Belgium Great Britain Informative S.B.M. Brasil Japan Irradians USA USA Isokojiki Japan USA Jaarverslagen, Schelpen. Friesland Netherlands Great Britain Jahrbiicher der Deutschen Malak. Ges. Germany Germany Japanese Journal of Malacology, The Japan Great Britain JECOLN Annual Report Japan USA Jersey Sheller, The USA USA Johnsonia USA USA Journal de Conchyliologie France/Australia Great Britain Journal for Medical and .Applied Malacology USA USA Journal fiir die Liebhaber des Steinreichs Germany USA Journal of Cephalopod Biology Canada Italy Journal of Conchology, The Great Britain Australia Journal of Malacology, A [Levantina] Israel Cuba Journal of Malacology, The Great Britain France Journal of Moll. Behav, [parody] Great Britain USA Journal of Molluscan Studies, The Great Britain USA Journal of Shellfish Research USA New Zealand Journal of the Malacological Society Australia USA Junonia USA USA Kainakama Japan Netherlands Keppel Ba\ Tidings Australia itain/Australia Korean Journal of Malacology, The South Korea USA Kreukel, De Netherlands Great Britain Laddergram, The USA USA Lavori della Societa Italiana di Malac. Italy USA Lavori della Societa Malac. Italiana Italy R. Bieler and A. R. Kabat. 1991 Page 59 Leaflets in Malacology Levantina Littorina, The Littorinid Tidings, The Long Island Shell Chib News Louisiana Malacologicai Society Low Country Shell Club Newsletter Magasin de Conchyliologie Maiacofauna Italica Malacologia Malacologia Mostra Mondiale Malacologicai and Conch. Magazine Malacologicai Club of Victoria Malacologicai Newsletter [Levantina] Malacologicai Review Malacologicai Society of Australia Malacologicai Society of London Malacologicai Society of South Aust. Malacology Data Net Malakologiai Tajekoztato Malakologische Abhandlungen Malakozoologische Blatter Mappa Marginella Marginalia Material of the National Pearl Res. Lab. Members of the American Malac. Union Miami Malacologicai Society Quarterly Minutes, Conch. Club of Southern Calif. Miscellanees Malacologiques Mitchelli, The Mitteilungen der Berliner Malakologen Mitteilungen der Deutschen Malak. Ges. Mollusca Mollusca, The Off the Beach Notes Molluscan Digest Molluscs in Archaeology & the Recent Mollusk Chaser Mollusk, The Monographs of Marine Mollusca Moreton Bay Radula, The Mostra Malacologica Messinese Mrs. Imogene C. Robertson's Rambling Notes Musee Heude, Notes de Malacologie Nachrichtsblatt der Deutschen Malak. Ges. Naples Shell News National Capital Shell Club National Shellfish. Assoc. Quarterly News Nautilus, The NCMC Newsletter Neue Litteratur and Beytrage zur KenntniB New Jersey Shell Club New York Shell Club Notes News Letter, Auckland Museum Newsletter, Marco Island Shell Club, Inc. Newsletter of African [Achatina] Newsletter of the Conch. Society Newsletter of the Jersey Cape Shell Club News of the Assoc. Shell Clubs of Calif. News of the Western Assoc, of Shell Clubs Noble Pen, The North Carolina Shell Club Bulletin Northampton Nat. Hist. Soc North Texas Conchological Soc. Quarterly Northern California Shell Club News Northwest Shell News USA Notes and Quotes, Rochester Shell USA Israel Notiziario C.I.S.M.A. Italv USA Notiziario S.I.M. Italy Great Britain Nouveau Pes-Pelicani, Le France USA Nouvelles Miscellanees Malacologiques France USA Novitates C-onchologicae Germanv USA Occasional Papers on Mollusks USA France Occasional Publ. of the Inst of Malac. Japan Italy Of Sea and Shore USA USA Opisthobranch USA Italy Opisthobranch Newsletter USA Great Britain Oregon Shell News USA Australia Original Tide-ings, The USA Israel Ostrea Japan USA Out of the Shell Canada Australia Pacific Northwest Shell News USA Great Britain Pacific Shell News Japan Australia Pallidula Great Britain USA Palm Beach County Shell Club News USA Hungary Papustyla, The Papua New Guinea Germany Pariah, The USA Germany Pearl Oyster Information Bulletin Australia France Pes-Pelicani France USA Philippine Shell News Philippines Japan Pittsburgh Shell Club Bulletin USA USA Puirieria New Zealand USA Porphyria Netherlands USA Proi^eedings of the Malac. Society Great Britain France Proceedings of the Malacozool. Meeting Japan USA Proceedings of the Miriam Vale Australia Germany Proceedings of the Natl. Shellfish. Assoc. USA Germany Proceedings of the Pacific Shell Club USA USA Proceedings of the Philadelphia Shell Clu lb USA USA Proceedings of the Soc. for E.\p. [Walkerana] USA USA Proces-Verbau.x des Seances de la Societe Belgium Canada Proserpine Shell Club News Australia USA Publicafoes Ocasionais da Sociedade Portugal USA Publication of the Conchology Group USA USA Pyrum France Australia Quarterly Journal of Conchology Great Britain Italv Quarterly Report Miami Malac. Soc. USA USA Reefcomber South Africa China Resenas Malacologicas Spain Germany Review of the Broward Shell Club, A USA USA Revista de la Sociedad Malacologica Cuba USA Revue Biographique de la Societe Malac. France USA Robusta French Polynesia USA Rossiniana New Caledonia USA Royal Society of South Australia Australia Germany Sanibel-Captiva Shell Club News Letter USA USA San Marcos Shell Club Quarterly Newsletter USA USA Sarasota Shell Club Newsletter USA New Zealand Sayana, The USA USA Schede Malacologiche del Mediterraneo Italy Netherlands Schriften zur Malakozoologie Germany South Africa Seafari USA USA Searcher, The USA USA Series Conchyliologiques France USA Shell and Tell USA USA Shell Case, The USA USA Shell Collector USA Great Britain Shell Gossip Fiji USA Shell Museum . Newsletter USA USA Shell Notes USA USA Shell, The [La Conchiglia] Italy Page 60 THE NAUTILUS, Vol. 105, No. 2 Shell Sheet, The Shell-0-Gram Shelletter of Shells and their Neighbors Shells and Sea Life Si rat us Slit Shell, The Slug Newsletter [parody] Slug, The [parody] Snail Farming Researili Sociedad (^hilena de Malacologia Soosiana South Florida Shell Cluh .South Padre Island Shell Chih Newsletter Southwest Florida Shell News Southwestern Malaeologieal Society Special Occasional Publication Special Publication, Inst, of Malac. Species Conchyliorum Spicileges Malacologiques, Les Steeriae, La Sterkiana Strandloper, The Subscriber's Bulletin Suncoast Shorelines Sunshine Coast Shell Club News Sydney Sheller, The Tabulata Tampa Drill Tentacle Texas Conchologist Texas Shell News Thais Thatcheria Tibia Trumpet Tide-ings Tide Lines Tidesong Tide Watch Tidewater Triton, The Townsville Shell Club Transactions of the Chinese Society USA USA USA USA Brasil USA South Africa USA Italy Chile Hungary USA USA USA USA USA Japan Great Britain France French Polynesia ' USA South Africa Great Britain USA Australia Australia USA USA Great Britain USA USA Panama USA USA USA USA New Zealand Australia USA Australia China Zeitsehrift fiir Malakozoologie Zoological Becord (Mollusca Sect.) ACKNOWLEDGEMENTS Germany Great Britain Transactions of the Isaac Lea Conch. Chapter USA Transactions of the POETS Soc [Walkerana] USA Treasure Coast Shell News LTSA Tree and Land Snail Collectors Papua New Guinea Tryonia USA UN IT AS Malacologica Newsletter Germany Unterhaltungen fiir Conchylienfreunde Germany Utah Shell Society USA Vancouver & District Shell Bulletin, The Canada Veliger, The USA Venus, The Japan Vertigo France Victorian Branch Bulletin Australia Walkerana USA W. A. Shell (Collector Australia Wellington Shell Clhib New Zealand West American Mollusca USA Western Society of Malacologists. Ann. Rep. USA Western Society of Malacologists, Occ. Paper USA Whangarei Shell Club New Zealand Whelk Wavelength, The LISA .Xenophora France Xenophora Newsletter, The USA Young Shell Collectors Quarterly, The Peru/ USA We thank the following colleagues who have checked the entries for various countries or states on our behalf: R. Tucker Abbott, Philippe Bouchet, Walter Cernohor- sky, Eugene Coan, Richard Kilburn, Ian Loch, Bruce Marshall, Akihiko Matsukuma, Paula Mikkelsen, Robert Moolenbeek, Richard Petit and Walter Sage; their assis- tance was indispensable. In addition, a large number of individuals provided information on certain publications (usually of shell clubs). These include Ruby Barry, Alan Bebbington, Connie Boone, Carol Boswell, Jean Cate, Barb Collins, Phil Col- man, Phyllis Diegel, Laurie Ford, Peggy Fox, Levente Fiikoh, Dorothy Germer, Raye Germon, Alison Haynes, Russell Jensen, Rudolf Kilias, L. A. Koestel, Florence Kuczynski, Betty Lawson, Charlotte Lloyd, Jordan Marche, George Metz, Henk Mienis, Walter Morgan, Olive Peel, Frances Perry, Charles Pettitt, William Pitt, Berneice Plummer, Thomas Rice, Gary Rosenberg, Rich- ard Salisbury, J. B. Sessoms, Carol Skoglund, Shirley Slack- Smith, John Stanisic, Mike Sweeney, Jose Templado, Jane Topping, F. White and Peggy Williams; we appreciate their help. For helpful discussion concerning the content and the introductory material of this paper, we thank Kenneth Boss, Eugene Coan, William Emerson, M. G. Harase- wych, Richard Johnson, Richard Petit and Gary Rosen- berg. LITERATURE CITED Abbott, R. T. 1962 Historical notes on the .\merican Journal of Conchology. Sterkiana 6:1-4. Abbott, R. T. (ed.). 1976. The best of the Nautilus: a bicen- tennial anthology of American conchology. American Mal- acologists, Greenville (Delaware), viii -I- 280 pp. Backhuys, W. 1985. Bibliographical note on the journals of the Societe (Royale) Malacologique de Belgique, Mede- delingen van de Werkgroep voor Tertiaire en Kwartaire Geologic 22:17:3-188. Burch, J. B. 197:3. Malacologia — ten years of publication. Malacological Review 6(l):69-72. Burch, J. B. "1973" [1974]. Malacological Review — six years of publication. Malacological Review 6(2):205-208. Burch, J. B. 1981. Annual Reports and Bulletins of the Amer- ican Malacological Union. Malacological Review 14:67- 112. Burch, J B and J M Huber. 1968 Malacologia — five years of publication The .American Malacological Union, Inc. [and] The .Xmerican Malacological Union, Inc Pacific Di- vision, Annual Reports for 1967:28-29. Biirk, R. and J. H. Jungbluth. 1985. 140 Jahre Mollusken- kunde im deut.schsprachigen Raum 1844-1984. Unga- risches Naturwissenschaftliches Museum, Budapest, 348 PP Buschmeyer, H 1938. 70 Jahre Molluskenkunde( 1869-1938). Senckenbergische Naturforschende Gesellschaft, Frank- furt am Main, 108 pp. R. Bieler and A. R. Rabat, 1991 Page 61 Carlton, J T. and D. W. Behrens. 1981 Index to the Opis- thobranchia [in] The VeUger, volumes 1-23, Opistho- brant-h Newsletter 13;27-38. Gate, J. M 1989. A personal history of The Veliger and the California Malacozoological Society, Inc. The Veliger 32: 101-108. Clench, W J and R D Turner 1962 New names introduced b\ II .•K Pilsbry in the Mollusca and Crustacea Academy of Natural Sciences of Philadelphia, Special Publication 4: 1-218. Coan, E. V. 1966. Charles Russell Orcutt, pioneer Californian malacologist, and The West American Scientist. Trans- actions of the San Diego Society of Natural History 14: 85-96, Coan, E. V. 1976 The availability of taxa proposed in the Minutes of the Conchological C'lub ot Southern California. The Veliger 18:326-331. Coan, E. V. 1989. The malacological papers and taxa of Martha Burton Woodhead Williamson, 1843-1922, and the Isaac Lea Chapter of the Agassiz Association. The Veliger 32296-301. Coche, A. G. 1983. Lists of serials, newsletters, bibliographies and meeting proceedings related to aquaculture FAO Fisheries Circular 758: vii + 1-65. Dance, S. P. 1986. A history of shell collecting. E J Brill. Leiden, xvi + 266 pp., 32 pis. Dhondt, A. V. 1989. La Societe Malacologique de Belgique. Annales de la Societe Royale Zoologique de Belgique 1 19: 139-153. Duchamp, R. 1986. Liste des nouveaux taxa publics dans ■INFORMATIONS de la Societe Beige de Malacolo- gique ". Apex 1:67-69 Emerson, W. K. and W. E. Sage. 1987. On the availability of names proposed in Pacific Shell News, Tokyo, 1970- 72. The Nautilus 101:194-199. Ende, M. B. van der and R. A. Bank. 1989. Register op de pubhkaties van Basteria deel 1-50 (1936-1986). Infor- matieblad van de Nederlandse Malacologische Vereniging 9:1-40. Feldmann, R. M. 1989. On the costs of journal subscriptions Journal of Paleontolog) 63:958 Fischer, P.-H. "1977" [1978], A brief history of the Journal de Conchyliologie Malacological Review 10:85-86. Fischer- Piette, E. 1937. Dates de publication du "Journal de Conchyliologie" de 1861 a 1900 Journal de Conchyliolo- gie 91:88-92. Friess, H. 1982. Johann Samuel Schr5ter (1735-1808) a pi- oneer in palaeontology. Archives of Natural History 11: 83-98. Heppell, D and C. Pettitt [in prep ] Index to taxonomic illustrations in malacological periodicals. Hertz, C. 1984 Illustration of the types named by S. Stillman Berry in his "Leaflets in Malacology". The Festivus 15 (Supplement): 1-42. Jacobson, M. K. 1971. Index to the Revista de la Sociedad Malacologica "Carlos de la Torre", volumes 1-9, 1943- 1954 (all volumes issued) Sterkiana 44:1-44. Johnson, R. I. 1969. Pfeiffer's Novitates Gonchologicae, Series I, Land Mollusca, 1854-1879, and Dunker's Novitates C^onchologicae, Series II, Marine Mollusca, 1862-1882. Journal of the Society for the Bibliography of Natural History 5:236-239. Jutting, W, S. S. van B. and C O. van R Altena 1958 A list of printed malacological periodicals Basteria 22: 10-15, Keen, A. M. 1954. [Note regarding "Transactions of the Isaac Lea Chapter"]. Minutes of the Conchological Club of Southern California 138:39 Keen, A. M 1983. Dr Rudolph [sic] Stohler and The Veliger: an appreciation. Hawaiian Shell News 31(7):9-10. Kobeh, W 1914 Alphabetisches Verzeichnis der in den Jahr- giingen 1-45 des Nachrichtsblattes der Deutschen Mala- kozoologischen Gesellschaft als neu beschriebenen Mol- lusken Moritz Diesterweg, Frankfurt am Main, 23 pp La Rdcque, .\. 1961 The American Journal ol Conchology. Sterkiana 5:43-51. Lillico, S 1990 Scientific journals are better. Hawaiian Shell News 38(2):5-6. Loch, I. 1990. Species descriptions in newsletters Hawaiian Shell News 38(1 0):8-9. McClelland, H. 1923. General index of all families, genera, species and varieties described and noted in The Journal of Conchology Vols. I-XVI, 1874-1922. Proceedings of the Malacological SocietN of London Vols. I-X\', 1893- 1923, The Conchologist Vols. I & II, 1891-1893 (all issued), continued as The Journal of Malacology Vols. III-XII, 1894-1905. Birbeck & Sons, Birmingham', 165 pp. McMillan, N F. 1961 F. P Marrat and his Conchological Leaflets. Liverpool Bulletin 9:26-29. Merton, R K 1968 The Matthew Effect in science. Science, 159:56-63. Moran, J. B. 1989 The journal glut: scientific publications out of control The Scientist 3(14):11, 13 Petit, R. E. 1988. Note on Leaflets in Malacology The Veliger 30:424. Price, D. J. deS. 1986 Little science, big science . . . and beyond. Columbia Llniversitv Press, New York, xxvi + 301 pp Rice, T 1990. A sheller's directory of clubs, books, periodicals and dealers, 14th ed Of Sea and Shore, Inc., Port Gamble, Washington, 82 pp. Roth, B and J T (Carlton 1970. A forgotten periodical of West American conchology. The Nautilus 84:31-32. Ruhoff, F. A. 1980. Index to the species of Mollusca intro- duced from 1850 to 1870. Smithsonian Contributions to Zoology 294:1-640. Schopf, T. J. M. 1967. The literature of the ph\lum Ecto- procta: 1555-1963. Systematic Zoology 16:318-327. The Serials Directory, 3rd ed. 1988-89. 1988. Ebsco, Bir- mingham, xlvi + 4,845 pp. [3 volumes]. Smith. A. G. 1976. Stearns Conchological Memoranda The Veliger 18:405-410. Solem, [G] A. 1974. The shell makers: introducing mollusks. New York: John Wiley & Sons, xiv -I- 289 p., 12 pis. Sykes, E. R. 1901. A summary of the American Journal of Conchology 1865-1872. Digesta Malacologica Lviii + 46 pp. Tomlin, J. R le B. and N. Fisher 1935. Two editions of vol. i of The Conchologist The Journal of Conchology 20:192. Wheeler, A. G 1989 The pressure to publish promotes dis- reputable science. The Scientist 3(14):1 1, 13. Winckworth, R 1936 Journal de Conchyliologie: dates of publication. Proceedings of the Malacological Society of London 22:153-156. Winckworth, R. 1937. Journal de Conchyliologie: dates of publication. Proceedings of the Malacological Society of London 22:357. 1988/89 Zoological Record Serial Sources (covering Zoological Record, X'olumes 124-125). 1990. Biosis, Philadelphia, X -f 291 pp. THE NAUTILUS 105(2):62-80, 1991 Page 62 First Records For Cymathim mundum (Gould) in the Eastern Pacific Ocean, with Comments on the Zoogeography of the Tropical Trans-Pacific Tonnacean and Non-Tonnacean Prosobranch Gastropods with Indo-Pacific Faunal Affinities in West American Waters William K. Emerson Dt-parlment of Invertebrates American Museum of Natural History Central Park West at 79th Street New York, NY 10024, USA ABSTRACT Cymalium (Monoplex) mundum (Gould, 1849), a wide-rang- ing Indo-West Pacific species, is recorded from the Galapagos Islands. This taxon has yet to be recognized elsewhere in the eastern Pacific Ocean, but is here reported from the western Atlantic for the first time. A lectotype is selected for Cymatium gemmatum (Reeve, 1844), which was previously confused with C. mundum. The occurrences are documented in west Amer- ican waters for other trans-Pacific Tonnacean species of the following genera: Tonna in the Tonnidae, Charonia. Cyma- tium. Linatclla in the Ranellidae, Distorsiu in the Personidae, and Bursa in the Bursidae. The modern Tonnacean faunal element in the eastern Pacific apparently reflects survivors of a previously widespread, post- Tethyan biota modified by factors resulting from the closure of the Central American Seaways about three million years ago, together vvitli post-Pliocene supplementation from the central Pacific, The present non-Tonriacean prosobranchs in the eastern Pacific with Indo-Pacific faunal affinities appear to represent largely post-Pliocene additions to the eastern Pacific that were derived from western Pacific communities. Key words: Tonnacea; systematics; new records; Ranellidae; Personidae; Bursidae; zoogeography; Indo-Pacific Prosobran- chia in eastern Pacific Ocean; marine snails. INTRODUCTION This paper records the presence in the eastern Pacific Ocean of the Indo-West Pacific ranellid gastropod, Cy- malitim mundum (Gould, 1849). The occurrences of this wide-ranging species in the New World are based on specimens collected by Jacqueline DeRoy and Carmen Angermeyer in the Galapagos Islands, some 15 to 35 years ago. This taxon is differentiated from Cymatium gem- matum (Reeve, 1844), with which it was commonly con- fused. The first records of C. mundum in the western Atlantic are here reported, based on specimens collected by Frank and Vera Lyman off Florida some 50 years ago. Additional specimens of Cymatium muricinum (Roding, 1798), a tropical cosmopolitan species known elsewhere in the eastern Pacific from Panama (Emerson, 1983:119, figs. 15, 16) are reported from the Galapagos Islands confirming a previous record which was based on a specimen in the DeRov Collection (Emerson, in Radwin, 1969:235). A zoogeographical survey was undertaken of the trop- ical trans-Pacific Tonnacean species and non-Tonnacean prosobranchs with Indo-Pacific faunal affinities known to occur in the western and eastern Pacific. On the basis of the available data, the origins of these faunal elements in the eastern Pacific Ocean are discussed in terms of their distribution in time and space. ABBREVIATIONS AHF = Allen Hancock Foundation Collection; see Eraser (1943) and McCulloch (1977) for station data (depos- ited in the LACMNH) AMNH = American Museum of Natural History, New York BM(NH) = The Natural History Museum, London [for- merly the British Museum (Natural History)] CASIZ = California Academy of Natural Sciences, San Francisco, CA LACMNH = Los Angeles County Museum of Natural History, CA MCZ = Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, MA MNHN-Paris = Museum National d'Histoire Naturelle, Paris W. K. Emerson, 1991 Page 63 NMNH = National Museum of Natural History, U.S. National Museum Collection (USNM), Smithsonian In- stitution, Washington, D.C. NZGS = New Zealand Department of Scientific and Industrial Research, Geology and Geophysics Division [formerly the New Zealand Geological Survev], Lower Hutt SBMNH = Santa Barbara Museum of Natural History, Santa Barbara, CA article 49b, in place of Neptunellininae Gray, 1854 as used by Waren and Bouchet, 1990:58. Genus Cyrnatitan Boding, 1798 Type species: Miirex jemorale Linne, 1758, by subse- quent designation of Dall (1904:133). Synonyms: Loto- rium Montfort, 1810; Tritociirris Lesson, 1842; Nyctilo- chus Gistel, 1848. SYSTEM.ATIC TREATMENT For man\ \ears. the identit>' of Cymatium mundum (Gould, 1849) has been misunderstood by most workers, who erroneously considered Gould's ta.xon to be a junior subjective synonym of C. gemmatum (Reeve, 1844). This confusion dates from Reeve's (1844a) original description and illustrations in which specimens of what are now recognized as C. mundum (Gould, 1849) were consid- ered by him to be a variety of C. gemmatum (A. Beu, in litt.. November 13, 1989). As indicated by Springsteen and Leobrera (1986:116), C. mundum differs from C. gemmatum in the shell being ". . . shorter and wider and more coarsely sculptured; [with] the varices . . . thick- er and [with] the intervarical cords . . . more pronounced especially on the spire whorls. ..." The slender, more delicate Cymatium gemmatum is rarely white and is commonly orange-brown in basal color, with darker brown and white varical bands as shown in color by Springsteen and Leobrera (1986:115, pi. 31, figs. 12a, 12b). The more massive C. mundum is yellowish to white in basal color and specimens commonly exceed 30 mm in length (Abbott and Dance, 1982:120, us C. gemmatum Reeve). In contrast, specimens of C. gemmatum rarely attain 30 mm in length and they lack, in maturity, the heavy outer lip characteristic of C. mundum on which the apertural lip projects above the labral surface (cf. figure 8 with figure 24). Cymatium gemmatum appears to be largely confined in distribution to the western Pacific, where in the central Philippine Islands specimens are found in depths to 100 meters. Verified records of this taxon from localities on the Pacific Plate, however, are scarce. In the case of C. mundum, the range is spread throughout the Indo-Pa- cific faunal province. Moreover, specimens of this taxon are commonly found at localities on the Pacific Plate, from the intertidal zone to moderate depths. See range data given below. In the following section, the taxonomic status of these two taxa is reevaluated and the two nominal species are recognized as separate and distinct taxa. Family Ranellidae Gray, 1854 (= Cymatiidae Iredale, 1913; see Beu and Cernohorskv, 1986:242) Subfamily Cymatiinae Iredale, 1913; see Jansson and Beu, 1990a, proposal to conserve Cymatiinae under ICZN Subgenus Monoplex Perry, 1811 Type species: Monoplex australasiae Perry, 1811 [= Cymatium parthenopeum (von Salis, 1793)], by subse- quent designation of Dall (1904:138), fide Beu and Kay (1988:197). Synonyms: Lampusia Schumacher, 1817; Cymatriton Clench and Turner, 1957. Cymatium {Monoplex) gemmatum (Reeve, 1844) (figures 1-8) Triton gcmniatns Reeve, 1844a, Triton text, pi. 15, figs. 60a, b; figs. 1, 2 herein. Type locality: Island of Ticao, Philippines (12°30'N, 123°42'E), "Found on stones at low water); Cuming". Reeve, 1844b: 117. Cymatium {Monoplex) gemmatum, Beu, 1985:58, "West Pa- cific". Cymatium (Septa) gemmatum, Springsteen and Leobrera, 1986: 114. 116. pi 31. figs, 12a, b, "Camotes Sea and Sulu Sea", Philippines, Triton (Simpulum) gcmmatus, Tryon, 1880:13. in part, pi, 7, fig. 41 only [copy of Reeve, 1844a, pi, 15, fig. 60a]. Range: western Pacific: Philippine Islands (Bohol Straits, AMNH 232143; Punta Engano, Mactan Island, Cebu, ex.— F. J. Springsteen, AMNH 232142); Palau Islands (Kror Island, AMNH 92774), Indonesia (Ambon Island, USNM 746407); Solomon Islands (AMNH 93092; NZGS); Fiji Islands (Vanua Levu, USNM 695079); New Cale- donia (MNHN-Paris; teste Beu); Coral Sea (Chesterfield- Bellona Plateau, MNHN-Paris; teste Beu), Austral Is- lands (Tubuai Island, USNM 705501 ). Remarks: The probable syntypes of Triton gemmatus Reeve in the BM(NH) [lot #198055] consist of 4 speci- mens, which range from 16.2 mm to 28.9 mm in height. One of these specimens appears to be the specimen used to illustrate Triton gemmatus (Reeve. 1844, pi. 15, figs. 60a, b) and is here selected as the lectot\pe (cf. figures 1, 2 with 3, 4 herein). The lectotype (28.9 mm in height) is slightly larger than Reeve's figure (28. 1 mm). The early whorls are missing owing to breakage, as is the case in Reeve's figure, but the lectotype possesses a varix on the third postnuclear whorl which is not shown on Reeve's illustration Otherwise, the lectotype compares favorably with the figure. Three paralectotypes of C. gemmatum Page 64 THE NAUTILUS, Vol. 105, No. 2 Figures 1-8. Cymatium gemmatum (Reeve); x 2. 1, 2. Copy of Triton gemmatus Reeve, 1844a, pi. 15, figs. 60a, b. 3, 4. Lectotype BM(NH) 198055. 5, 6. Paralectotype BM(NH) 198055a. 7. 8. .\MNH 232142, off Mactan Island. Cebu, Philippines, ex-F. J. Springstein. Figures 9, 10. Cymatium muricinum (Roding) Crabbed specimen, Santa Cruz Island, Galapago.s Ids.. Oct. 1969, AMNH 232144, ex-J DeRoy; xl Figures 11-15. Cymatium mundum (Gould); x 2. 11. Copy of Triton mumium Gould, 1849, after Gould, 1856, pi. 17, fig. 297. 12, 13. Lectotype USNM 5695. 14, 15. Copies of Triton mundum Gould, 1849, after Gould, 1856, pi. figs. 297a,b. W K. Emerson, 1991 Page 65 are here recognized in the type lot [BM(NH) 198055]. The largest of the three paralectotypes is illustrated here- in (figures 5, 6). The old locality label accompaning the type lot reads, "T. gemmatus, var., I. Masbate [12°11'N, 123°30'E] on the reefs, under stones. H. [ugh] C. [uming]". Whereas the "Hab. [itat]" given by Reeve (1844a); for T. gem- matus is "Island of Ticao, Philippines" [12°13'N, 123°42'E], (found under stones at low water; Cuming)." These islands are in close pro.ximity, being separated by the Masbate Pass. Inasmuch as the existing label accom- panying this lot refers to a variety of T. gemmatus, the original locality stated by Reeve (1844a); for this taxon is here retained as the type locality. At the time Triton gemmatus was proposed, Reeve (1844a, pi. 15, fig. 60c) briefly described and illustrated a specimen of what is now recognized as Cijmatium mundum under the appellation of "Triton gemmatus variety B. ' There are three specimens in the type lot [BM(NH) #196736]. The largest specimen most closely resembles Reeve's figure (see figure 20, herein), but this specimen measures 40.9 mm in height vs. the figure which is 38 mm in height. The other two specimens are, respectively, 38.9 and 33.5 mm in height. Although these specimens are typical of examples of Ctjmatium mun- dum (Gould, 1849), they retain the status of paralecto- types of Ctjmatium gemmatum (Reeve). The old label accompanying the type lot cites only the "Philippines" as the locality. Cymatium (Monoplex) mundum (Gould, 1849) (figures 11-24) Triton gemmatus variety B, Reeve, 1844a, Triton text, pi. 15, fig. 60c [figure 20 herein], "Island of Annaa (Chain island). South Pacific Ocean, and island of Burias, Phil- ippines (found under stones in both localities at low wa- ter); Cuming". Reeve, 1844b: 117. Not T. gemmatus Reeve. Triton mundum Gould, 1849:143; Gould, 1856:506, pi. 17, figs. 297, 297a-b [figures 11, 14, 15 herein]; Gould, 1862:66; Johnson, 1964:112. Type locality: "Tutuilla [sir; Tutuila] Samoa Islands" (American Samoa, 14°18'S, 170°42'W). Cymatium {Morwplex) mundum, Beu 1985:58, "Indo- West Pacific". Cymatium (Septa) mundum, Springsteen and Leo- brera, 1986:116, pi. 31, fig. 13, "Camotes Sea and Sulu Sea , Philippines. Triton (Simpulum) gemmatus, Tryon, 1880:13, in part, pi. 7, figs. 43, 44 [copy of Gould, 1856, figs. 279, 279a] only. Not T. gemmatus Reeve. Cymatium (Septa) gemmatum. Clench and Turner, 1957:222, in part, reference in synonymy to Triton mun- dum Gould only. Not T. gemmatus Reeve. [Clench and Turner mistakenly referred their western Atlantic rec- ords of the Indo-west Pacific species, C. vespaceum (La- marck, 1822), to C. gemmatum. Beu (1985:60) now con- siders these east American populations to be a distinct species, namely: C. comptum (A. Adams, 1854)]. Finet, 1985:19, "Galapagos Islands", based on AM.\H 1.39535. Not T. gemmatus Reeve. Cymatium gemmatum, Salvat and Rives, 1980:306, fig. 173, "lies de la Societe aux Tuamotu a Raevavae (Australes) et aux Marquises"; Abbott and Dance, 1982: 120, "Indo-Pacific; West Indies, under coral, on sand, shallow water; uncommon". Not C. gemmatum Reeve. Range (Selected localities from 50 AMNH lots): West- ern Indian Ocean (Durban Bay, South Africa, AMNH 196938); (off Gedi, Kenya AMNH 96528); (off Kizimkazi, Zanzibar, AMNH 101032); off Somalia (Mogadiscio, AMNH 142680); (Mauritus, AMNH 113606); eastward into the Pacific Ocean, northward to southern Japan (Okinawa, AMNH 169800) and southward to the Fiji Islands (Suva Reef, AMNH 214467), Solomon Islands (Malaita Island, AMNH 118330) and New Caledonia (Touho, AMNH 104266), northeastward to the Hawaiian Islands (Oahu Island, AMNH 152284; Maui Island, AMNH 147510) and southeastward to the Societv Islands (Huahine Island, AMNH 231972; 239056); French Aus- tral Islands (Tubuai, AMNH 84641) and Marquesas Is- lands (Salvat and Rives, 1980:305). NEW RECORDS For The Eastern Pacific Ocean: Galapagos Islands, Academy Bay, Isla Santa Cruz, 1 fresh specimen, found on beach, June, 1966, ex-J. DeRoy (AMNH 139535); 1 crabbed specimen, found at low tide, December, 1965, ex-J. DeRoy (AMNH 232145), see figures 21, 22. Galapagos Islands, 1 crabbed specimen, South Channel (Isla Santa Cruz- Isla Baltra) collected about 1975, C. Angermeyer collec- tion. NEW RECORDS For The Western Atlantic: 1 spec- imen, south end of Lake Worth, Palm Beach County, Florida, Vera Lyman!, ex-A. D'Attilio collection (NZGS WM 15228, teste A. Beu). 1 specimen dredged from Gulf Stream, off Palm Beach County, Florida, Frank Lyman! 1940, ex-AHF (LACM 115537, teste A. Beu). Remarks: In a review of the taxa described by Augustus A. Gould, Johnson (1964:112) selected for Triton mun- dum Gould (1849:506, pi. 17, fig. 297, a-b) the lectotype (USNM 5695, cf. figures 12, 13 with Figure 11, herein) and a "paratype" [= paralectotype] (USNM 612311, cf. figures 14, 15 with figures 16, 17 herein). These speci- mens, which measure 29.9 and 21.9 mm in height, re- spectively, compare favorably with Gould's somewhat stylized drawings. As Johnson (1964:112) noted, an ad- ditional specimen, which is labeled a paratype [= para- lectotype] from the A. A. Gould Type Collection, No. A427, is present in the Museum of Comparative Zoology (#169249). The MCZ specimen (figures 18, 19), which had been obtained from the New York State Museum, measures 28.8 mm in height. The two type specimens (lectotype and paralectotype) in the USNM collection are presumably from the type locality, "Tutuilla, Samoa Islands " (Gould, 1849), but the existing labels only identify the source of the two lots, namely: the U.S. Exploring Expedition (1838-42). There Page 66 THE NAUTILUS, Vol. 105, No. 2 Figures 16-24. Cyinatium mtindum (Gould); x2. 16, 17. Paralectotype USNM 612311. 18. 19. Paralectotype MCZ 169249. 20. Copy of Triton gemmatus var. B. Reeve, 1844a, pi. 15, fig. 60c [= C. mundum]. 21, 22. Crabbed specimen, Santa Cruz Island, Galapagos Ids., ,\MNH 232145, e.\-J. DeRoy 23, 24. Huahine Island, Society Ids., AMNH 239056, note fully developed outer lip are no additional data in the USNM catalogue for either lot (teste M. G. Harasewych). ZOOGEOGRAPHIC IMPLICATIONS Tables 1 and 4 enumerate and document the trans- Pacific Tonnacean species known to occur in the tropical eastern Pacific, including records for the oceanic islands (Clipperton Island, Revillagigedo Islands, Cocos Island, and Galapagos Islands) as well as those from the conti- nental borderland. Some of the records are based on limited samples of one to three individuals that are known only from dead-collected specimens. These include Ton- na perdix perdix, Cyrnatium aquatile. C. mundum. and W. K. Emerson, 1991 Page 67 Linatella succincta. The rarity of these and other records may reflect, in part, the lack of extensive collecting by means of SCUBA diving and dredging operations. The marine moUusks of Cocos Island (about 9 km in circum- ference) are now reasonably well known owing largely to the intensive collecting activities during the past de- cade of Drs. Donald R. Shasky and Michel Montoya with the cooperation of their numerous field associates (Shas- ky, 1989c; Montoya, 1983, 1988). Whereas the moUuscan faunas of most of the other oceanic islands may have received attention for a longer period of time, much of this collecting predates the use of SCUBA diving, and the insular faunas of these widely isolated and not easily accessible islands still remain poorly known. This is also largely true of the Galapagos Islands because of the geo- graphical extent of this archipelago and the long-stand- ing restrictive regulations pertaining to the collection of specimens (cf. Wellington, 1984). On the mainland, for most of the tropical zone (Mexico to Ecuador), much of the subtidal coastal waters and those of the near-shore islands have not been extensively explored by collectors. This is especially true of the communities associated with the coral-reef biotope (Emerson, 1967:91). A possible exception are the coral reefs of Panama where James Ernest, Royce Hubert and others have concentrated their collecting efforts for mollusks in recent years (Emerson, 1983). Nevertheless, the majority of the Tonnacean rec- ords are based on live-collected or well-preserved dead specimens from numerous sources over a considerable period of time, commencing with explorations early in the nineteenth century by Alexander von Humboldt and Aime Bonpland (Valenciennes, 1833). Students of zoogeography have long noted the pres- ence of marine mollusks with Indo-Pacific faunal affin- ities in the subtropical and tropical waters of the eastern Pacific (Carpenter, 1857:346; Darwin, 1860:391 and oth- ers). Hertlein (1937) was the first, however, to discuss specifically this subject and detail the shallow-water mol- luscan species then known to be common to Polynesian and western American waters. He concluded that these Indo-Pacific species had likely dispersed the 5,000-km- wide body of open ocean (Ekman's East Pacific Barrier) separating the most eastern situated Polynesian Islands and the New World by means of long-lasting larval stages transported by ocean currents, and/or by attachment to drifting objects, to pelagic fishes, to mammals, or on the feet of wide-ranging oceanic birds. He also noted that nearly all of these mollusks were gastropods, many in the Conidae, Cymatiidae [= Ranellidae and Personidae] and Cypraeidae, and most of them were known in the eastern Pacific only from the oceanic islands, far from the West American Continental Borderland. Our present knowledge largely serves to support Hert- lein s conclusions. In the tropical Tonnacea, several spe- cies of Cymatium including C. pileare martinianum, and C. miiricinum, have been shown to have teleplanic larvae (Scheltema, 1989:439). Additionally, a larval stage lasting nearly a year in the water column is known for the cosmopolitan species Cymatium parthenopeum and C. nicobaricum (Scheltema, 1971: table 1). Furthermore, C. parthenopeum and others are known to feed in the plankton and to be able to delay metamorphosis until a shallow-water substrate is reached (Pechenik et al., 1984). Oceanic currents appear to be the primary vehicle for the passive dispersal of mollusks with hemipelagic larvae (Scheltema, 1986, 1989; Zinsmeister & Emerson, 1979; Emerson, 1990) and for the distribution of species that attach to, or burrow into floating objects. Rafting may be the means of dispersal for some epifaunal bivalves (Turner, 1966:52, 1971; Scheltema, 1977;93; Emerson, 1978:92) and gastropods (Jokiel, 1990:71). The major cur- rent circulation in the tropical Pacific apparently has favored east to west dispersal pathways since the late Neogene, or perhaps much earlier (cf. Grigg, 1988; New- ton, 1988; Scheltema, 1988; Skelton, 1988). On the other hand, the paucity of mollusks with Panamic faunal af- finities in Polynesian waters suggests that recruitment of tropical Panamic species into the central Pacific via dis- persal by the westward flowing current patterns has been largely unsuccessful (cf. Briggs, 1970:134; Zinsmeister & Emerson, 1979:36; Scheltema, 1988:149). This mostly one- way trans-Pacific distribution exists even though the westward transport of organisms across the East Pacific Barrier would presumably be facilitated by the accel- eration of westward flowing currents during pre-El Nifio/ Southern Oscillation events (Jokiel, 1990:69). As noted previously, Hertlein (1937:309) pointed out that many of the Indo-Pacific species recorded by him occurred only on the oceanic islands in the eastern Pacific and were not known to have spread eastward to the continental borderland of the Americas. This observation remains largely true for the non-Tonnacean prosobranch species, of which only 20 of the 48 taxa (41%) compiled herein are known from the mainland (tables 2, 3, Ap- pendix). In contrast, 42 (87%) of these 48 taxa are re- ported from the oceanic islands in the tropical eastern Pacific, with 33 (68%) at Clipperton Island, 16 (33%) at Cocos Island, 10 (20%) in the Galapagos Islands, 7 (14%) in the Revillagigedo Islands, and 1 (2%) at Guadalupe Island (table 4). These species represent an attenuated Indo-Pacific element that has reached the oceanic islands in the eastern Pacific, but the majority of these ta.xa (59%) have not been reported from the west American main- land (table 4). Furthermore, none of the 48 species is known from the western Atlantic, although one survives in the eastern Atlantic and the Mediterranean, presum- ably as a post-Tethyan faunal relict (table 4). The only fossil evidence for the temporal existence of the non-Tonnacean Indo-Pacific faunal element in west America is the presence of Cypraea cernica Sowerby in the Pleistocene of Guadalupe Island (Lindberg et al., 1980:52) and Hastula albula (Menke) in the Miocene of California, a species living at the present time in the Revillagigedo Islands (Bratcher & Burch, 1971; table 3, herein). Nor have any representatives of this faunal el- ement been recovered from Mesoamerican archaeolog- ical sites (Emerson, 1983:120). If large and colorful gas- tropods such as Mitra mitra (Linne) had been commonly Page 68 THE NAUTILUS, Vol. 105, No. 2 "c.l _ # ^ ir a. = _ c 0; 1— CI - rt -2 « — c- (Tl 0; c 7 i/i OJ &0 o 91 c O iH a -i o c S -a o H c u ly rt CB -O c Oi -rfi c o tn rt ?5 ^ UJ < o 3j c ^ < c 03 o 5 o ,JS U J2 — 7 "O ■* Ml S 3- M~ X X x: I I I I XXX Ixxxxx xxxxxxxx X X X X X X X X X X et X - X t- X X X X I X X XXX X X X X I I I X ^ I 2 X t- Mill ic — X X + 00 X CO _ o - CO X X T c •2 -H. + 5 + + c c a o J^ <^ DO MC C s ~ ■^ § :c - CC o "^^ t 2 3 ~ £ -is ~ ,.1, t c 3 C c £ 1- Q + :0 CC ■3- O ~ a '.J c £ 'H. c 3 Q .2 ■ — ' Ji* 0. r3 — 3 a. a. c S £ fc fc S S •^ 3 (D O -T ., — 'P ~ . rt o E a -^ CO O CO ^' si [ijoo'^.^Uo'^SJcom~-''C: J3 ° o c pa a 5 *i o CO ^^1 c CO <^ U rt GO "^ _, = ":::■« £ ?? ^ yp iJj ..c ^ to , C/^ 00 CT o . ., ' CO 00 I 05 ^ « 2 n 00 — 'in c -^ 0/ C CD 2 05 CO S" OS J3 s < o ^ ^ - t^ "H of oo' 0- n in _ o c cj to £ . in o o o E •£ ^ ^ So = i .- . -r 00 "^ p: 00 -" -T, t- ' "^ in "^ ^J ' „• CO J E; I T3 ^ , c/5 ' ^ ^ CO <■ 00 ^ tc i: I _, ^ o O ^ c U "-^ - .-c CD - 7^ b 2 =c' in U I « * ^ ir^ Cu ■ « 6 a. o o l^ J; N 05 a 3 "^ ^ ^ ' ■^ '^ 00 a.^ t^ o c - ■/> ±^ i .i o E O 0- J= O — o a-r cr (/I o '^[-,05 o o fc -a ^ a ^ — w ^ S so ii Z a o c X < O rt E2 u .. 05 —" '^ u c . o o -I .EZ J2 S -a o u S 00 CTJ cc -a -H 00 c iC c c "5 P IS I •sNcc o o 3 o ^ P — . ffl _i: -^ ■r" I t- . ■ o .t: ~ 2 ^-^ Cl,-^ >:; C i/^ - c ^ - - 3-£ c '^ _ c o i _ -S u 03 CD E'.. .< W a; L, o S CD -o 00 t: ■3 Co D- 0^ c "^^ " "C '^ :3 a ^ c3 ^ a| ~ t- p c ro 5 c! — t: c u ^ a a, 3 £ ■> s c^ 5 — a in -^ ^ fc-5 II >. ^ o ~ E « -P P t^ I " tt E OS 00 _- li » 00 ,E ^ '^ O C/2 2 js -o c^ ._^ -' -o O 1- in .- T 03 t^ — " u: o -T3 - - < f: S I s g o5 2 z « — — • — C/5 -H 3 o !_) in -T O 3 < IM 00 c o ^ CB r O ;d h- Oj lyi o^ .ir •"■■ O^ Ul o c c; O H) I/; o r £ '^ .2 :z -j .£ o >• t! - cj — ~ 5u ._ •- o g^ BC .- 3 o S; O 3 J= >, 3 6C — C5 3 :5 3 ;^ = 1- C ..-c O — in 00 , O^ C35 00 ^ o S3 ' ^ ;-' ^LT O ■— " ■" " -^ =5 ii: 0/ CO ^ < . , o — £ ^ " §""?"§ S — — ti- c « ° z:^ ..X a f "3 CJ .^ -t- CD o -o ro S t; <3> S 3 00 « — J rt 2 =c 2 ^3 Z . "O S-t- K-5:= S J r o 3 &■■ V ..D E o i CO ^ o||J Oj 3 "^ U. 3 . < . ' « 3 r CO Ej5 > -C J3 30 o t^Cin -2 c5 r.„ 00 J£ ., W CJ5 o ^. -^ ^ 00 C5 ^^ .3 CTs in o "^ . 1-; i" o > o 5 J:: 2 SB o "- O CJ i; 3 = = 3 . o £ ^ "-3 ■!> S 3 ; § _M — in ,S P r "^ u - ■> s C" i~-3'^t- £cD r: IX ~ ir — 01 -i; o -^ o o >« ~ — r. -a a: - 2 2 "6 fl p " " O "2 o -; ■K — S O. o - CJ s .£ -^ a ■5 «-^ 5 3 S « Sea 5'-/5 t '^ II JO w ^ i^ — W. K. Emerson, 1991 Page 69 Table 2. Shallow-water Indo-Pacific prosobranch gastnipods {excluding the Toiiiiacea) recorded from the West American Con- tinental Borderland (Appendix) and their distribution on the eastern Pacific islands (records from F.merson, 1978. 1983; Everson, 1988; Robertson, 1979: Shasky, 1983-1989, and herein). None of these species is known from the .'\tlantic Ocean, except Quoyula madreporaruiu uhich was reported from the Cape Verde Islands (Emerson, 1983:122) Re\il- C'lipperton lagigedo txicos Galapagos CiiadaluiJc Island Islands Island Islands Island Neritacea Titiscaniidae 1. THificania liiiuicina (Rergh) .■\rchitectonicacea' .'Krchitectonicidae 2. Hcliactis Irochoidcs (Desha\es)-l-# 3. Philippic! radiiita (Roding) + # Littorinidae 4. Littorina pintado (\Vood)# Cypraeacea Cypraeidea 5. Cijpraea captitserpcntis Linne*-l-# 6. Cypraea talpa Linne* + # 7. Cypraea teres CJmelin [= ■'alisunae Burgess]*# Ovulidae 8. Pseudoeypraea adamsonii (8owerby) + Muricacea Muricidae 9. Qitoyula madreporarum (Sowerby)* + # 10. Reliquiaeeava rohiUardi (Lienard)* 11. Rhizochilus antipathieus (Steenstrup)# Mitracea Mitridae 12. Mitra mitra (Linne)* + # Conacea Conidae 13. Conus chaldcus (R6ding)*-l-# 14. Conus ebraeus Linne*-l-# 15. Conus tessulatus Born* + Terebridae 16. Terebra a ffinis Gray* + tt 17. Terebra laevigata Gra\ +# 18. Terehra niaeulota (Linne)* + # Turridae 19. Kermia felina (Hinds)-I- 20. Microdaphnc Inelwdes (Dalll*-i- Total \ taxa on mainland = 20 X X X X X X X 9(45" 3(15? X X X X X X 10 (50'' X X X X X 6(315 Explanation of symbols: asterisk (*) = taxa also known from Line Islands (Jewell, 1962; Kay. 1971; Ka\ and Switzer, 1974); plus sign ( + ) = known from French Polynesia (Dautzenberg and Bouge, 1933; Rehder, 1968; Salvat and Rives, 1980; Richard, 1985); and number sign (#) = known from Hawaiian .Archipelago (Gage, 1962; Schmeltz, 1978, 1979; Kay, 1979) ' For the purposes of this tabulation, .\rchitectonicacea is retained in the subclass Prosobranchia and not transferred to the subclass Heterobraiichia (see classification of Ponder and Waren, 1988:308), available in west American waters, one would expect specimens to have been reported in the kitchen middens and burial sites of the Native Americans. On the other hand, the tropical shallow-water Tertiary record is poorly preserved on the northwest American borderland. Until recently, no fossiliferous marine Tertiary onshore de- posits were recognized regionally, from southern Nica- ragua northward (Woodring, 1966, 1978; Perrilliat, 1978), except for the long-known exposures on the Tres Marias Islands, off San Bias, Mexico, onshore and on the islands in the Gulf of California and in southeastern California (Durham & Allison, 1960; Powell, 1988; Smith, 1989). In the mid-1970's however, reconnaissance exploration along the Pacific coast of Mexico, northwest of Tehuan- tepec (Durham et ai. 1981) determined the presence of seven onshore Tertiary sedimentary basins on the North America Plate, negating the belief pre\iousK held that most of these sediments had been lost to subduction or continental-margin truncation into the Middle American Trench system (Karige^ ai. 1978; Beck & Plumley, 1979), Four of these onshore west Mexican basins have yielded fossiliferous marine sediments of Miocene and younger Page 70 THE NAUTILUS, Vol. 105, No. 2 Table 3. Shallow-water Iiuio-Pafific pr().sobranch gastropods (excludiiii; the Tonnacea) recorded from the eastern Pacific Oceanic IslaiKJs that are not known from the West .Vmericaii Continental Borderland (records from Bakus, 1975; Cate, 1969; Kmerson, 1978, 198.3; Everson, 1988; Finel, 1987a,b; Kay, 1979; Shasky. 198:5-89, antl herein). None of these species is known from the .Mlanlic Ocean. C:lipperton Revillagigedo Cocos Galapagos Guadalupe Island Islands Island Islands Island Fissurellacea Fissurellidae 1. Diodora granifera ( Pease)* + # X — — — Trochacea Phasianellidae 2. Tricolia variabilis ( Pease)* + # — — X — — Neritacea Neritidae 3. Serita plicata Liimi:* + U X — — — — Cerithiacea Cerithiidae 4. Cerithitini ncsioticiitn Pilsbry & Vanatta* + # X — — — — Cypraeacea Cypraeidae 5. Cypraea depressa Gray*+ X — — — — 6. Cypraea helvola Linne* + # X — — — — 7. Cypraea maculijera (Schilder)* + # X — — — — 8. Cypraea moneta Linne* + # X — XX — 9. Cypraea rashleighana Melvill + * X — ? — — 10. Cypraea schilderorum (Ireddle)* + tt X — — — — 11. Cypraea .srurra Gmelin* + # X — — — — 12. Cypraea vitellus Linne* + # X — — — — Muricacea Muricidae 13. Drupa nioruni Roding* + # X — — — — 14. Drupa ricinus (Linne)* + # X — — X — 15. Moru/a uya (Roding)* + # X — X — X 16. Na,v.va .serfa (Bruguiere)* + # X — — — — 17. Coralliohia fimlmata (A. Adams)*# X — — — — 18. Corallioplula violacea (Kiener)* + # X X X X — Buccinacea Buccinidae 19. Clivipollia costata (Pease) + tt X — — — — 20. Clivipollia fragaria [WoocD + tt — — X — — 21. Cantharus fumostts {Di]]\\\n} + ff — — — X — Mitracea Mitridae 22. Mitra edentula Swainson + # X — — — — 23. Mi(ra /crn/guico Lamarck* + # X — X — — 24. Mitra papalis (Linne) + # X — X — — 25. Strigatella litleraia (Lamarck)* + # X — — — — Volutacea Harpidae 26. Harpa gracilis Ikoderip & .Sowerby+ X — — — — Conacea 27. Terefora crenu/a*a (Linne)*-l-# X X X — — 28. /7a«<«/a a//w/a (Menke) + # — X — — — Total N of insular taxa = 28' 24(85%) 3(10%) 8(32%) 4(14%) 1(3%) Explanation of ssnibols: asterisk (*) = taxon also known from Line Islanils (Jewell. 1962; Kay, 1971; Ka\ and Switzer, 1974); plus sign ( + ) = known from French I'oKnesia (Dautzenberg and Bouge, 1933; Rehder, 1968; Salvat and Rives. 1980; Richard, 1985); and number sign (#) = known from Hawaiian Archipelago (Schmeltz, 1978, 1979; Kay, 1979). ' Additional unverified records: 1, Vanihuro acuta (Rechiz) from Cocos Island (Shasky, 1987:49) and Triphora trilicea Pease from the Galapagos Islands (Shasky, 1989a:8). W. K. Emerson, 1991 Page 71 ages (Durham et al., 1981; Perrilliat, 1981). Indo-Pacific molluscan elements have yet to be reported from these basins (PerriUiat, 1987). Furthermore, no mollusks of the non-Tonnacean Indo-Pacific faunal element have been recorded from the numerous fossiliferous terrace deposits of Pleistocene age along the west American mainland, from Mexico to Peru. Thus, the paucity of tropical Ter- tiary fossils from much of the middle West American Continental Borderland has largely limited paleogeo- graphic interpretations of the faunistic history of the modern molluscan Panamic Province to the regional Pleistocene fossil record and to comparisons with the Tertiary faunal records of the adjoining Californian and Caribbean regions. The non-Tonnacean element (tables 2, 3) is present only at the genus-group level in the Tertiary faunas of the Caribbean region. A small Indo-Pacific relict element (conid subgenera, etc.) is recognized by Petuch (1988: 94, 96) as regionally extinct species in the West Indian Gurabo Formation (Pliocene). However, none of the non- Tonnacean Indo-Pacific species now living in the eastern Pacific is known to inhabit the western Atlantic (table 4) and none of these species has been recognized in the fossil record of the east American tropics. The modern and fossil occurrences of the species of the non-Tonna- cean Indo-Pacific faunal element in the eastern Pacific, therefore suggest that these taxa largely represent post- Pliocene colonizers derived by trans-Pacific dispersal from populations in the central Pacific. This distributional pattern is in sharp contrast to that for the 13 trans-Pacific Tonnacean species cited in table 1 . Eleven of the 13 species (84% ) also occur in the western Atlantic, 7 (53%) of which are recognized as subspecif- ically divergent taxa. The vast majority (8 of the 13 taxa) are represented in the eastern Atlantic and several (5 of the 10 taxa) also have populations in the Mediterranean Sea (see table 1). Only eight (61%) of these trans- Amer- ican taxa, however, can be termed truly circumtropical in distribution as the others are not known from the eastern Atlantic (table 4). The fossil record of the Tonnacean species in the New World tropics, although sparse, is somewhat better doc- umented at the species-group level than that of the non- Tonnacean trans-Pacific prosobranch species cited herein (tables 2, 3, Appendix). In the Tertiary, species of Cy- matium (sensn lato) are recognized in Oligocene de- posits of Panama and Mississippi (MacNeil and Dockery, 1984; Schmelz, 1989). Other cymatiid taxa are known from the Neogene period and Pleistocene epoch of the eastern Pacific and western Atlantic tropics. Beu and Kay (1988:196) recorded the presence of ". . . both C. pileare and C. aqitatile . . in the Pacific [Basin] since Miocene time", and they noted ". . . that C. aquatile lived in northwestern Ecuador at some time [Pliocene] during the Neogene [Olsson (1964; pi. 30, fig. 3, as C. cf. pi- leare)], and that unnamed taxa close to, but different from, both C. aquatile and C. martiniantim lived in the Caribbean during Miocene to mid-Pliocene time." The genus Distorsio is also known from the Oligocene of Mississippi (MacNeil, 1984) and elsewhere in Neogene and Pleistocene deposits of the east Americas, from Flor- ida to Brazil (Pilsbry, 1922; Emerson & Puffer, 1953; Woodring, 1959). In west America, Distorsio is reported from the Mio-Pliocene of southeastern California (Pow- ell, 1988:17, as D. constricta), of Costa Rica (Aguilar & Fischer (1986:223) and of northwestern Ecuador (Pilsbry and Olsson, 1941:40; Olsson, 1964:174). Distorsio first appears in the Cretaceous of Madagascar (Beu, 1988:89), Cijmatium (sensu lato) is known from the Paleogene and Neogene of the Old World [e.g., Oligocene onwards in the Aquitaine Basin), and Charonia occurs in the Oli- gocene of Europe and the Neogene of both southern Europe and the Caribbean area (Beu, 1970; Gibson-Smith, 1971) suggesting that the New World representatives of these taxa form part of a post-Tethyan distributional pattern. All 13 of the Tonnacean species listed in table 1 are known to occur on the oceanic islands within the eastern Pacific. Ten (76%) of the species are recorded from the Galapagos Islands, with 7 (53%) at Cocos Island, 4 (30%) at Clipperton Island and 3 (23%) in the Revillagigedo Islands. Seven species (58%) are reported from the main- land, including Cymatium corrtigatum amictuni, a non- trans-Pacific species with divergent subspecies in the western and eastern Atlantic Ocean (west Africa) and the Mediterranean Sea (table 1). Only 4 of the 13 taxa are commonly found on the west American mainland (Cymatium corrugatum amictum, C. pileare macrodon, C. parthenopeum keenae, and Distorsio c. constricta). One of these taxa (C. p. macrodon) is recognized by Beu and Kay (1988:211) as a full species on the basis of shell morphology, although they note the close resemblance of the eastern Pacific specimens to specimens of "typical" C. pileare in the western Pacific. Moreover, a faunal antiquity for the east American Tonnacea is further sup- ported by the recognition in the western Atlantic of 7 of the 1 1 Pan-Panamian species divergent at the subspecies level. The presence of 10 of the 13 eastern Pacific Tonnacean species living in the Galapagos Islands, with only 7 species on the west American mainland, may identify this ar- chipelago as a post-Pliocene refuge for some of these Tonnaceans. This hypothesis is supported by the low num- bers of Tonnacean species known from the other oceanic islands, especially Clipperton Island (4 species) and Cocos Island (7 species). Adverse post-Pliocene conditions in the tropical eastern Pacific may have caused the extinc- tion of some elements of the insular and mainland Ton- nacean populations. If this is the case, some of the present trans- Pacific Tonnacean fauna in the eastern Pacific must represent elements derived by dispersal from central Pa- cific populations commencing with the emergence of the Panamanian isthmus. Dana (1975) made a similar case for the decline of the Indo-Pacific hermatypic corals in the west American tropics during the Pliocene and their subsequent trans-Pacific replacement in the Pleistocene- Holocene. These factors (fossil record and the modern distributional pattern in the New World tropics) suggest Page 72 THE NAUTILUS, Vol. 105, No. 2 Table 4. A compilation comparing the distribution of the Incio-Pacific non-Tonnacean and Tonnacean prosobranchs in the eastern Pacific Ocean, with their occurrences in the Atlantic Ocean and Mediterranean Sea (based on tables 1-3, .•\ppendix) Non-Tonnacean species 1 Diodora granifera 2. Tricolia varialrilis 3. Nerita plica ta 4. Titiscania limacina 5. Littorina pintado 6. Cerithimii nesiolicuni 7. Helianis trochoides 8. Philippiu radiata 9. Cijpraea capiitserpcntis 10. Cijpraea depressa 11. Cijpraea helvola 12. Cijpraea niaculifera 13. Cypraea moneta 14. Cypraea rashleighana 15. Cypraea schilderorum 16. Cypraea scurra 17. Cypraea lalpa 18. Cypraea teres 19. Cypraea vitellus 20. Pseudocypraea adamstmii 21. Drupa rnorum 22. Drupa ricinus 23. Morum twa 24. Nassa serta 25. Quoyula madreporanim 26. Reliqniaecava rohillardi 27. Rhizochilus antipathiciis 28. Coralliohia finihriata 29. Coralliophila violacea 30. Clivipollia eostata 31. Clivipollia jragaria 32. Canthanis fumosus 33. Mitra edentula 34. Mitra ferruginea 35. Mitra mitra 36. Mitra papalis 37. Strigatella litterata 38. Harpa gracilis 39. Conus chaldeus 40. Conus elnaeus 41. Conus tessulatus 42. Terehra ajfijis 43. Terelna crenulata 44. Terelna laevigata 45. Terelna maculata 46. Hastula allmla 47. Kerrnia felina 48. Microdaphne trichodes Total N of taxa = 48 Tonnacean species 1. Tonna p. perdix 2. Charonia t. tritonis 3. Cymatium aquatile 4. Cymatium corrugatutn amictum 5. Cymatium mundum Eastern Pacific Ocean .Atlantic Ocean Oceanic Western Eastern Mediter- Islands Mainland Atlantic Atlantic ranean Sea X' X^ X' — — — — X' X' X X " = = X'' X X — — — X1.3 X' X' X' Xl.3.4 Xl.'3 X' X' x^ X — — — — — — — X — — — \1.3.-l X' X3.J X' X — — — X — — — X1.3..'; X' Xl,2 — — — — X I X X" X' X X — — — X' X 1.2.3.4 X' x^ x^ — — — — — — — — X' Yl,3 — — — — X3.4 X — — — XI. 3 X' — — — — X' Xl.3.4 X — — — Xl.3.4 X — — — Xl.2.3 X X X X — — — XI. 2.3 — — — X..2.3 I x^ X X — — — x^ — — — 42 (87%) 20(41%) 0 1 (1%) 1 (1%) x^ _ X'' _ X3.4 — X" X' X' X' X X X — XM.4 X X" X9 X» X' — X — — W. K. Emerson, 1991 Table 4. (^mtinued Page 73 Eastern Pacifi( lOi cean .•\tlai itic Ocean Oceanic Western Eastern Mediter- Islands M; ainlainl Atlantic Atlantic ranean Sea 6. Cumatium muricinum X^ x X X X 7. Cymatium pileare macrodon X 1.2.3.4 X X'" X'" — 8. Ci/matium nicobaricum XI.3 — X X X 9. Ciimatium parthenopeum kecnac X^ X X" X" X" 10. Linalella succincta x^ — — X — 11. Distorsio c. constricta XM X X12 — — 12. Bursa asperrima Xi.-i — — — — 13. Bursa g. granularis X1.M X X13 — — Total N of taxa = 13 13(100%) 7 (53%) 11 (84%) 8(61%) 5 (.38%) Non-Tonnat'ean and Tonnacean species Total N of taxa = 61 .5,5(90<7) 27 (44%) 11 (18%) 9(14%) 6(9%) Known from: 1 = Clipperton Island, 2 = Revillagigedo Islands, 3 = Cocos Island, 4 = Galapagos Islands, and 5 = Guadalupe Island. Recognized as subspecies: 6 as T p. maculosa. 7 as C. /. variegata. 8 as C. c. krebsii. 9 as C. c. corrugatum. 10 as C. p. martinianum, 11 as C. p. parthenopeum. 12 as D. c. macgintyi. 13 as B. g. cubaniana. 14 described as Q. m. var, mediterranea Parenzan (1970). a dual origin for elements of the Tonnacean species now- living in the eastern Pacific: 1, Survivors of a previously widespread Pan-Panamian post-Tethyan biota and 2, New arrivals of trans-Pacific species from the Central Pacific after the closure of the Central American Seaways. The apparently rare occurrences of many of the taxa of both the Tonnacean and non-Tonnacean constituents found on the islands and/or the mainland may reflect in part records of species that cannot maintain viable pop- ulations for any significant duration without frequent replacement by larval recruitment or by other means of dispersal from established populations in the mid-Pacific. One of the major causes of temporal instability of these populations may result from the harsh environmental changes caused by major El Nifio events in the eastern Pacific. Shasky (1985:4) reported a definite decline in numbers of some of the more common non-Tonnacean Indo-Pacific species and in the condition of the reef -coral at Cocos Island following the warming of the local waters as a result of the severe 1982-83 El Nino episode. He noted in 1984 the mortality of Pocillopora and found no living Coralliophila violacea until 1989, when a single living specimen was taken (Shasky, in litt., August 19, 1990). Unfortunately the collection database for the mol- lusks provided herein (tables 1-3, Appendix) is not suf- ficient to determine the long-term influence of the El Nifio induced changes on the survival of the Indo-Pacific faunal elements in the tropical eastern Pacific. The El Nifio disturbances may temporarily benefit some faunal elements and harm others, e.g.. cause adverse effects on the communities associated with the coral-reef biotope (Glynn, 1988; Richmond, 1990; Robinson, 1985, 1987). For an excellent review of the many factors affecting the biotic composition of eastern Pacific stony corals and other tropical marine organisms in time and space, see Glynn and Wellington (1983:167-207). The establishment at Clipperton and Cocos Islands of reproductively viable populations of the Indo-Pacific faunal element during "normal" conditions (non El Nino periods) would appear to be the key to the post-Pliocene dispersal of these faunal consituents to other areas in the tropical eastern Pacific. The large number of the non- Tonnacean Indo-Pacific taxa known from Clipperton and Cocos Islands serves to support this thesis (tables 2, 3). Twenty-four (85%) of the 28 taxa that are not known from the mainland have been reported from Clipperton Island, whereas 8 (28%) occur on Cocos Island. Of the 20 species known from the mainland, 9 (45%) have been recorded from Clipperton Island, and 10 (50%) occur on Cocos Island. In contrast, 10 of the 13 (76%) Tonnacean species occur in the Galapagos Islands, 7 (53%) on Cocos Island, and 4 (30%) on Clipperton Island. If all the Indo- Pacific prosobranch gastropods (Tonnacean and non- Tonnacean) in the eastern Pacific are considered, 55 (90%) of the 61 taxa are known from the oceanic islands and 27 (44%) are reported from the mainland (table 4). Thir- ty-seven of the 61 taxa (60%) are recorded from Clip- perton Island, 25 (40%) from Cocos Island, 18 (29%) from the Galapagos Islands, 9 (14%) from the Revillagigedo Islands, and 1 (<1%) from Guadalupe Island (table 4). The presence at these stepping-stone islands of several species of hermatypic corals, mostly with western Pacific faunal affinities (Glynn & Wellington, 1983:176), affords a habitat for Indo-Pacific molluscan species associated with the coral reef biotope. If the established populations of the Indo-Pacific ele- ment at Cocos and Clipperton Islands serve as a major source for larval dispersal to the Galapagos Islands and the west American mainland via the North Pacific Equa- torial Countercurrent (Abbott, 1966:109; Emerson, 1967: 89; Wellington, 1984:256; Glynn & Wellington, 1983: 167), any prolonged disruption of the reproductive vi- Page 74 THE NAUTILUS, Vol. 105, No. 2 ability of the populations at these insular sites would affect the gene replenishment required to reestablish de- clining or defunct populations on the mainland and in the Galapagos Islands. The subsurface Equatorial Un- dercurrent (Cromwell Current), however, is a possible source for larval transport directly to the Galapagos Is- lands from the central Pacific. This subsurface current passes eastward through the Line Islands and could pre- sumably disperse to the Galapagos Islands tropical or- ganisms tolerant of temperate subtropical temperatures (20-25°C at its core of ma.ximum velocity, between 50 and 100 m depth, teste R. S. Scheltema; cf. Glynn & Wellington, 1983:168). Furthermore, with the com- mencement of El Nino conditions, the surface flow east- ward from the Line Islands extends to the Galapagos Islands and onward toward the mainland (Richmond, 1990). Glynn (1985) estimated the period of uninter- rupted growth of living Pocillopora in the Gulf of Pan- ama at about 190 years, based on the death of the pro- tective coral barriers as a result of warming during the El Nino event of 1982-83. In terms of geological time, if only one such El Nifio-related dispersal event occurred every 200 years, this would result in 5,000 disruptive episodes per million years and have a significant influ- ence on the composition of the biota, past and present, in the eastern Pacific (Richmond, 1990). ACKNOWLEDGMENTS In addition to Mesdames Jacqueline DeRoy and Car- men Angermeyer, I am pleased to acknowledge the co- operation of a number of people for their aid in the completion of this paper. Types and other specimens, as well as data, were kindly provided by Kathie M. Way of The Natural History Museum, London, Silvard Kool of the Museum of Comparative Zoology, Harvard Uni- versity, Terrence M. Gosliner and Elizabeth Kools of the California Academy of Sciences, M. G. Harasewych and Raye N. Germon of the National Museum of Natural History, Smithsonian Institution, James H. McLean, Gale Sphon and Helen DuShane of the Los Angeles County Museum of Natural History, Henry W. Chaney of the Santa Barbara Museum of Natural History, Twila Bratch- er of Hollywood, California, Donald R. Shasky of Red- lands, California and Jody Woolsey of Los Angeles, Cal- ifornia. Specimens used for comparative study were generously donated by Robert Foster of Abbey Specimen Shells, Santa Barbara, California and F. Jim Springsteen of Victoria, Australia. I am indebted to Alan Beu of the New Zealand Department of Scientific and Industrial Research, Geology and Geophysics Division, Rudolf S. Scheltema of the Woods Hole Oceanographic Institution, Donald R. Shasky and two anonymous reviewers for kindly reading drafts of the manuscript and offering many helpful suggestions. I am grateful to my AMNH colleagues Walter E. Sage, III for technical assistance, Andrew S. Modell for photographic services, and Ste- phanie Grooms for word-processing the manuscript. LITERATURE CITED Aiibott. D. P. 1966. Factors inflviencing the zoogeographic affinities of the Galapagos inshore marine fauna. 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Epipodium with 3 pairs of micro- papillate tentacles; anterior pair long, about midway be- tween cephalic tentacle and second pair; second pair long or short, near anterior edge of opercular lobe; posterior pair long, near posterior edge of opercular lobe; small, thin, triangular flap may be present between anterior 2 pairs. Radula short, broad, with about 20-30 transverse rows of teeth; formula 6-8.4.1.4.6-8. Rachidian tooth cusp triangular, denticulate, excavated dorsally to accom- modate next anterior rachidian tooth cusp; base trian- gular, articulating with inner lateral tooth base. Lateral teeth 4 pairs, similar to those of llanga (Herbert, 1987). Marginal teeth 6-8 pairs, denticulate on outer edge near tip. Lateromarginal plate lacking. Type species (here designated): Margarita lameUosa Verrill & Smith, 1880. Etymology: From the Latin lamella, a little blade, and trochus, a child s hoop; gender masculine. Included species: Margarita lamellosa Verrill & Smith, 1880; Solariella pourtalesi ('lench & Aguayo, 1939; La- mellitrochns cariiiatus Quimi new species; L. inceratus Quinn new species; L. suatns Quinn new species; L. fi- losus Quinn new species; L. jenestratus Quinn new spe- cies; and L. Incoronatus Quirm new species. Remarks: The shells of species assigned to Lamellitro- chus are distinguished from all other solarielline genera by their strongly angular whorl profiles and distinctive macro- and microsculpture. The angular whorl profile is caused by the development of a strong subsutural an- gulation set with rounded, conical, or lamellate tubercles; a strong peripheral carina that is smooth, undulate, or tuberculate; a strong basal carina that is usually smooth, but ma\ be fineK rugose; and a strong, tuberculate cir- cumumbilical cord. The whorl surfaces between these angulations are almost flat, and usually have a varying number of fine spiral lirae. Several species currently as- signed to Solariella Wood, 1842, also have shells with angulate whorl profiles: S. triplostephaniis Dall. 1910; S. patriae Carcelles, 1953; and some forms of i>. cinctus (Philippi, 1836) and S. intermissa Thiele, 1925. I have not examined specimens of the first two species, and illustrations and descriptions (Carcelles, 1953; McLean, 1971) are not detailed enough for me to determine the species generic assignment; however, I doubt that these two species are congeneric with the Lamellitrochus spe- cies. Scanning electron micrographs of shells of S. cinctus (Fretter & Graham, 1977; fig. 32) and S. intermissa (Her- bert, 1987: figs. Ill, 112) reveal that neither species has micropustules on the early whorls. An extremely thin, calcified shell layer, which ma\' be chalk} or polished, overlies the outer porcelaneous shell layer of the last one or two teleoconch w horls. This thin outer layer may be the calcified periostracum, or intri- tacalx, described by D'Attilio and Radwin (1971). Nei- ther an intritacalx nor a periostracum have been reported for species of other solarielline genera, but they may have been overlooked; the systematic significance of the presence or absence of intritacalx in the Solariellinae is unknown at present. The surface of the first 2-3 whorls usually has a covering of micropustules. although the pustules may be restricted to the sutural area. In addition to those for Solariella cinctus and S. intermissa. SEM micrographs of the apical whorls of shells of llanga Her- bert, 1987, Spectamen Iredale, 1924, Minolops Iredale, 1929, and leminolia Finlay, 1927 (Herbert, 1987), and Minolia Adams, 1860 (Hickman & McLean, 1990) have been published. Of the species illustrated, only the South African species Spectamen adarticulatum (Barnard, 1963) (see Herbert, 1987: fig. 124) has discernible mi- cropustules. Conversely, Lamellitrochus pourtalesi seemingly lacks micropustules (figure 6). Micropustules also do not occur on shells of western Atlantic Solariella. Microgaza Dall, 1881, or Suavotrochus Dall, 1924 (per- sonal observations), but are present in Pagodatrochus Herbert, 1989 (Herbert, 1989: fig. 4a,b). Six of the eight species discussed in this paper have well-developed, usually lamellate axial riblets that ap- pear on the first half-whorl, usually immediately follow- ing the terminal rim of the protoconch. Shells of a ninth species from Argentina (off Rio de la Plata, USNM 330860) also have this characteristic, but, because the two spec- imens are juveniles, this species is not described here. The illustrations of the first whorls of the genera cited above (Fretter & Graham, 1977; Herbert, 1987; Hickman & McLean, 1990) show that shells of other .solarielline genera lack strong axial riblets on the first whorl, and that the riblets, present on later whorls are generalK weaker, more rounded, and more closely spaced than those of Lanu'llitrochus. J, F. Quinn. Jr., 1991 Page 83 ,f"^ \ Figures 1-6. 1-3. Lamellitrochus lamellosus (Verrill & Smith, 1880) from Eolis Station 351, SE of P'owey Light, Florida, 165 m (USNM 438440). 1. .\pertural view, 11.6 x. 2. Basal view (oblique), 12.7 x, 3. Protoconch, 118 x, 4-6. Lamellitrochus pourtalesi (Clench & Aguayo, 1939). 4, 5. Shell from John Elliott Pillsbury Station P-747, 11°46'N, 67°05.7'W, 1,175-1,098 m (UMML 30.6832). 4. Apertural view, shell height 9.1 mm. 5. Basal view, shell diameter 7.6 mm. 6. Protoconch of shell from Alaminos Station 69A11-86, 21°41'N, 96°51'W, 969-1,079 m, FSBC I 40300, 53 x. The radiila of Lamellitrochus lacks any trace of a lateromarginal plate (figures 28-36) and resembles Ilan- ga Herbert, 1987, "some North Atlantic taxa" (Herbert, 1987:287), and possibly Zetela Finlay, 1927 (Herbert, 1987) in lacking this structure. Herbert (1987) used the presence or absence of a lateromarginal plate to assign species with similar shell morphologies to different gen- era, but he did not speculate on the broader systematic implications of the character because of the lack of de- tailed knowledge of many genus-level groups. The presence of an intritacal.x is the only character that seems to be unique to this species group, but in combination with presence of strong axial riblets and distinct micropustules on the early whorls of most species; presence of strong, usually tuberculate, subsutural and peripheral carinae; and lack of a lateromarginal plate of the radula, it supports establishment of a new genus- group taxon. Until more detailed analyses of shell and radular characters of more solarielline species groups are made, I prefer to treat Lamellitrochus as a genus rather than a subgenus of Solariella. Lamellitrochus pourtalesi is illustrated here for com- parison with the other species, but a complete species account is found in Quinn (1979). Key to species of Lamellitrochus. 1. Protoconch diameter greater than 350 fj.m 2 1. Protoconch diameter less than 350 ^ni 3 2. Protoconch diameter 350-375 /urn; axial riblets present on first half-whorl . . inceratus new species 2. Protoconch diameter 525-550 /im; axial riblets absent on first half-whorl pourtalesi (Clench & Aguayo, 1939) 3. Tubercles on subsutural angulation lamellate .... 4 3. Tubercles on subsutural angulation conical or rounded 5 4. Axial riblets continuous between suture and pe- riphery; shell wider than high; height less than 3 mm carinatus new species 4. ,\xial riblets not continuous from suture to pe- riphery; shell higher than wide; height greater than 3 mm 5 5. Peripheral carina smooth to weakly undulate . . lamellosus (Verrill & Smith, 1880) 5. Peripheral carina with strong, conical tubercles filosus new species 6. Subsutural angulation and circumumbilical cord formed by double rows of rounded tubercles . . bicoronatus new species Page 84 THE NAUTILUS, Vol. 105, No. 3 6. Subsiitural angulation and circuiniimbilical cord formed by single rows of conical tubercles 7 7. Axial riblets present on first half-whorl; peripheral carina tuberculate suavis new species 7. Axial riblets absent on first half-whorl; peripheral carina undulate fenestratus new species Lamellitrochus lamellosus (Verrill & Smith, 1880) (figures 1-3, 28-30) Margarita lamellosa Wrriil & Sinitli in \crrill. 1880;.392, 397, 398. Solariella (Solariella) lamellosa: Quiiiii, 197940, 42 (partini). Description: Shell small, attaining 4.2 mm height, 3.55 mm width, conicoturbinate, peripherally carinate, broadly umbilicate, white, nacreous under thin outer porcelaneous layer. Protoconch 280-310 nm maximum diameter. Teleoconch whorls about 5, tubular, shoul- dered, carinate; first two whorls with strong, sharp, axial lamellae extending from suture to suture; axials becom- ing discontinuous or obsolete on subsequent whorls; mi- cropustules rather strong, covering entire surface of first 1.5-2 whorls; subsutural angulation and peripheral ca- rina appear on second whorl; subsutural angulation be- coming strong, tuberculate, tubercles lamellate and closely set; peripheral carina becoming strong, tuberculate, but usually becoming smooth or irregularly undulate on last 1-2 whorls; as many as 13 weak spiral threads present between subsutural angulation and peripheral carina; as many as 12 weak spiral threads present between periph- eral and basal carinae. Base flat, circumscribed by strong, smooth spiral carina; as many as 20 weak, smooth spiral cords present between basal carina and strong, tuber- culate circumumbilical cord; umbilicus broadly open, 35-45% maximum shell diameter, funnel-shaped, walls with about 10 fine, weak spiral threads and weak axial rugae. Aperture oblique, circular, lips thin; peristome complete. Animal and radula as for genus. Holotype: USNM 44738, height 3.0 mm, width 3.0 mm. Type locality: Off Martha's Vineyard, Massachusetts, Fish Hawk Station 871, 40°02'54"N, 70°23'40"W, 210 m. Other material examined: More than 100 lots in USNM, DMNH, FSBC I; see Quinn (in press) for listing. Remarks: Shells of Lamellitrochus lamellosus are most similar to those of L. filosus in shape and in having lamellate tubercules on the subsutural angulation; how- ever, the former species is absolutely and relatively larger (4.2 mm vs. 3.15 mm with about 5 whorls) but has a smaller protoconch (280-310 nm ds. 315-325 jum). The tubercles on the peripheral carina of shells of L. filosus are stronger, conical, and more widely spaced than those of L. lamellosus. and the spiral threads in the intercarinal spaces are weaker and the axial threads are stronger in L. filosus than in L. lamellosus. The shell of L. suavis is also very similar in shape and general sculpture, but the subt hiral tubercles are rounded, the periphery has rath- er strung tubercles, the micropustules are not as promi- nent and are concentrated near the suture, and the in- tercarinal spiral sculpture is weaker than in L. lamellosus. Lamellitrochus lamellosus is distributed along the Unit- ed States east coast from Massachusetts southward through the Florida Keys, and in the Gulf of Mexico off western Florida, Texas, and the Campeche Bank in depths of 50- 250 m (usually in 100-200 m). Lamellitrochus carinatus new species (figures 7-12, 34, 35) Description: Shell very small, attaining 2.75 mm height, 2.90 mm width, depressed conicoturbinate, peripherally carinate, broadly umbilicate, white, nacreous under thin outer porcelaneous layer. Protoconch 275-300 nm max- imum diameter. Teleoconch whorls about 4, tubular, shouldered, carinate; first two whorls with strong, sharp, axial lamellae extending from suture to suture, but end- ing at or just above peripheral carina on subsequent whorls; micropustules rather strong, covering entire sur- face of first 2 whorls; subsutural angulation and periph- eral carina appear on second whorl, peripheral carina becoming strong, smooth to weakly undulate; 1-7 weak spiral threads present between subsutural angulation and peripheral carina; 0-4 weak to strong spiral threads may be present between peripheral and basal carinae; whorl surface with microscopic pustules. Base weakly convex, circumscribed by strong, smooth spiral carina; 1-8 (usu- ally 3-6) weak to strong, smooth to weakly undulate spiral cords present between basal carina and strong, tuberculate circumumbilical cord; umbilicus broadly open, 35-45% maximum shell diameter, funnel-shaped; walls with 1-7 weak spiral cords and strong axial rugae that radiate onto inner part of base, forming strong, radially elongate tubercles on circumumbilical cord. .Ap- erture oblique, circular, lips thin, peristome complete. Holotype: DMNH 179393, height 2.50 mm, width 2.70 mm. Type locality: SW of Egmont Key, Florida, 366-421 m; J. Moore, collector. Paratypes: DMNH 186768, 22 specimens; USNM 859408, 1 specimen; FSBC I 37633, 1 specimen; UF 161754, 1 specimen; all from same lot as holotype. — USNM 859431, 13 specimens; Eolis Station 360, off Fow- ey Light, Miami, Florida, 146 m.— -USNM 438401, 3 specimens; Eolis Station 321, off Western Dry Rocks, Florida, 119 m.— 31, DMNH 179392; 100 mi SW of Egmont Kev, Florida; D. Steger, collector. — FSBC I 24259, 5 specimens; MAFLA Station 2746-41, 27°07'N, 84°13'W, 122 m; 23 August 1977; box corer.— USNM 500245, 37 specimens; State Universitv of Iowa Barbados Station 29, off Lazaretto, 165-183 m.— USNM 500266, State University of Iowa Barbados Station 77, off Cable Station, 73-137 m.— MORG 26529, 1 specimen; Marion- Dufresne Cruise MD-55, Station DC-61, 20°29'S, 29°18'W, 63 m; May 1987; dredge.— MNHN uncata- logued, 1 specimen; Marion-Dufresne Cruise MD-55, J. F. Quiiin, Jr., 1991 Page 85 8 ,«:'V H^ 11 '^^^^ .12f^ Figures 7-12. LamellUwchtis carinatus new species. 7-9. Paratypes from SW of Egmont Key, Florida, 366-421 m (DMNH 186768). 7. Apertural view, 15 x 8. Basal view, 17 x . 9. Protoconch, 134 x 10-12. Paratvpe from Marion-Dujresne Cruise MD-55, Station DC-61, 20°29'S, 29°18'W, 63 m (MORG 26529) 10. Apertural view, 15.3 x . 1 1. Basal view, 14.9 x 12. Protoconch, 82 X. Station DC-59, 20°.30'S, 29°19'W, 52-60 m; May 1987; dredge. Other material examined: More than 50 lots in USNM and FSBC I; see Quinn (in press) for listing. Remarks: Shells of Lamellitrochus carinatus are dis- tinguished from those of all other Lamellitrochus species principally in being distinct!) wider than high and usu- ally having strong and continuous a.xial riblets on all whorls. This species also has the smallest shells, with heights of 2.50-2.75 mm and widths of 2.75-2.90 mm; adult shells of all other species exceed 3 mm in height. Variation in shell morphology occurs between northern and southern populations, but this is a reflection of the frequency of individual character states because the same variations also occur within individual populations. Shells of the northern (Florida) populations tend to have smooth to weakly undulate peripheral carinae; 2-4 strong, often unequally sized basal spiral cords; fewer, usually weaker spiral threads in the spaces between the subsutural an- gulation and peripheral carina (usually 1-3), and the peripheral and basal carinae (0-1); and narrower spaces between the axial riblets (about 0.15-0.20 mm apart on the third whorl). Shells from Barbados and Brazil usually have distinctly undulate peripheral carinae, the crests of the undulations often sharp; the basal spiral cords tend to be more numerous (usually 3-6, sometimes 7 or 8) and often equally sized and spaced; spiral threads in the intercarinal spaces are relatively numerous (usuallv 3-4 in each) and distinct; and the axial riblets are more widely spaced (about 0.25 mm apart on third whorl). This spe- cies is the most widely distributed of all Lamellitrochus species, ranging from off North Carolina southward to both sides of Florida; off Cuba, Puerto Rico, Antigua, and Barbados; and Trinidade Island off eastern Brazil, usually in depths of 100-200 m. Lamellitrochus fenestratus new species (figures 13-15) Description: Shell small, attaining 3.35 mm height, 2.95 mm width, conicoturbinate, peripherally carinate, broadly umbilicate, white, nacreous under thin outer porcelaneous layer. Protoconch 275-300 ^m maximum diameter. Teleoconch whorls 4.6, tubular, shouldered, carinate; first 0.5 whorl w ith 4 spiral cords, increasing to 6 on second whorl; whorls 0.5-2 with initially weak, progressively stronger, rounded, axial riblets extending from suture to suture; axials becoming discontinuous or obsolete on subsequent whorls; micropustules weak, re- stricted to sutural area of first 2 whorls; subsutural an- gulation and peripheral carina appear on third whorl, peripheral carina becoming strong, undulate; 5-7 spiral threads present between subsutural angulation and pe- ripheral carina; 2-5 weak spiral threads and distinct axial rugae present between peripheral and basal carinae. Base flat, circumscribed by strong, smooth spiral carina; 6-9 weak, smooth spiral cords present between basal carina Page 86 THE NAUTILUS, Vol, 105. No. 3 14 20 Figures 13-21. 13-15. Lamellitrochns fenestrates new species, paratype from off Barbados, 18.3 m (USNM 859429). 13. Apertural view, 14 X, 14. Basal view, 14 x, 15. Protoconch, 84 x, 16-18. Lamellitroclms filosus new species, holotype from off English Harbor, Antigua (USNM 5002.30). 16. Apertural view, 14.3 x. 17. Basal view, 14.2 x^ 18. Protoconch, 91 x, 19-21. Lamcllitwchus suacis new species, holotvpe from off Barbados (USNM 500224). 19. Apertural view, 13.7 x . 20. Basal view, 13.1 x , 21. Protoconch, 92 X. ' and strong, tuberculate circumiimhilical cord; umbilicus broadly open, 30-35% ma.ximum shell diameter, funnel- shaped, walls with about 4 spiral cords and strong axial rugae. Aperture oblique, circular, lips thin; peristome complete. Holotype: USNM 859428, height 3.15 nmi, width 3.00 mm. Type locality: Off Barbados, Blake Station (data im- recorded), 183 m. Paratypes: USNM 859429, 4 specimens; from same lot as holotype. — USNM 500202, 1 specimen; State Univer- sity of Iowa i: rbados Station 62, off Pelican Island, 229 m— IISNM 859430, 1 specimen; State University of Iowa Barbados Station (data unrecorded), "deep". Remarks: Shells of L. fenestratus are most similar to those of L. bicoronaius but differ in having only a single row of conical tubercles forming the subsutural angu- lation rather than a double row of rounded tubercles, in having a stronger peripheral carina w ith stronger tuber- cles, and in ha\ iiig a single rather than double circumum- bilical cord. Shells of L. fenestratus are also similar to tho.se of L. filosus and L. suavis in size, protoconch width, and general shape, but differ from both in lacking axial ribletson the first half-whorl; in having conical tubercles in the subsutural angulation as in L. filosus, but a more J. F. Quinn, Jr., 1991 Page 87 undulate peripheral carina rather than that of L. suavis; and the spiral threads between the subsutural and pe- ripheral carinae are stronger in L. fencstratus than in either L. filostis or L. suavis. Lamellitrochus filosus new species (figures 16-18) Description: Shell small, attaining 3.15 mm height, 2.85 mm width, conicoturbinate, peripherally carinate, broadK umbilicate, white, nacreous under thin outer porcelaneous la\er. Protoconch 315-325 jum maximum diameter. Teleoconch whorls about 5, tubular, shoul- dered, carinate; first two whorls with strong, sharp, la- mellate axial riblets extending from suture to suture; axials becoming discontinuous or obsolete on subsequent v\ horls; micropustules weak, restricted to sutural area on first 2 whorls; subsutural angulation and peripheral ca- rina appear on second whorl; subsutural angulation be- coming strong, tuberculate, tubercles lamellate; periph- eral carina becoming strong, tuberculate; 13-15 very weak spiral threads present between subsutural angula- tion and peripheral carina; as many as 11 very weak spiral threads present between peripheral and basal ca- rinae; spiral threads intersecting equally sized axial threads forming file-like surface sculpture. Base flat, cir- cumscribed by strong, smooth spiral carina; as many as 20 weak, smooth spiral cords present between basal ca- rina and strong, tuberculate circumumbilical cord; um- bilicus broadly open, 40-45% maximum shell diameter, funnel-shaped, walls with 6-8 or more very weak spiral cords and weak, lamellate axial rugae. Aperture oblique, circular, lips thin; peristome complete. Holotype: USNM 500230, height 2.75 mm, width 2.50 mm. Type locality: State University of Iowa Antigua Station 116, off English Harbor, "deep". Paratypes: USNM 859425, 4 specimens; from same lot as holot\pe. Remarks: Shells of L. filosus are most similar to those of L. lamellosus. particularly in having lamellate tuber- cles on the subsutural angulation, but differ in being smaller (3.15 mm vs. 4.20 mm, respectively, at whorl 4.9), in having strong, conical tubercles on the peripheral carina, and in having much weaker spiral threads be- tween the subsutural carination and peripheral carina. Shells of L. filosus are also similar to the holotype of L. suavis, but have lamellate rather than conical tubercles on the subsutural angulation, and a fine, file-like sculp- ture on the surface of the last whorl. Lamellitrochus .mavis new species (figures 19-21) Description: Shell small, attaining 3.35 mm height, 2.90 mm width, conicoturbinate, peripherally carinate, broadly umbilicate, white, nacreous under thin outer porcelaneous layer. Protoconch 325 fxm maximum di- ameter. Teleoconch whorls 4.6, tubular, shouldered, car- inate; first two whorls with strong, sharp, lamellate, axial riblets extending from suture to suture; axials becoming obsolete on subsequent whorls; micropustules weak, scat- tered over entire surface of first 2.5 whorls; subsutural angulation and peripheral carina appear on second whorl; subsutural angulation becoming strong, tuberculate, tu- bercles bluntly conical; peripheral carina becoming strong, tuberculate; 5 weak spiral threads present be- tween subsutural angulation and peripheral carina; 3 or 4 extremely weak spiral threads present between pe- ripheral and basal carinae on last whorl. Base flat, cir- cumscribed by strong, smooth spiral carina; 10 weak, smooth spiral cords present between basal carina and strong, tuberculate circumumbilical cord; umbilicus broadly open, 35% maximum shell diameter, funnel- shaped, walls with weak axial rugae. Aperture oblique, circular, lips thin; peristome complete. Holotype: USNM 500224, height 3.35 mm, width 2.90 mm. Type locality: State University of Iowa Barbados Station (data unrecorded), "deep". Remarks: The shell of L. suavis resembles a miniature L. inceratus in shell shape and sculpture, but is smaller and has a smaller protoconch (325 ixm vs. 350-375 ^m, respectiveh ). Shells of L. filosus are similar to that of L. suavis in size, protoconch width, and tuberculate pe- ripheral carina, but differ in having lamellate rather than conical tubercles on the subsutural angulation, and in having fine, crowded axial threads that interact with equally fine spiral threads to produce a fine, file-like sculpture on the vn horl surface. Lamellitrochus bicoronatus new species (figures 22-24) Description: Shell small, attaining 3.55 mm height, 3.20 mm width, conicoturbinate, peripherally carinate, broadK umbilicate, white, nacreous under thin outer porcelaneous la\er. Protoconch 290-300 ^m maximum diameter. Teleoconch whorls 4.3, tubular, shouldered, carinate; first whorl with 5 strong spiral cords with ad- ditional threads appearing on second whorl and increas- ing in number on subsequent whorls; 3 cords, 1 just below suture, 1 above midvvhorl, and 1 below midwhorl strengthening on second whorl, forming tuberculate sub- sutural and undulate shoulder angulations, and weakly undulate peripheral carina, respectively; low, rounded axial riblets appear at end of first whorl, becoming stron- ger and more lamellate on whorls 2-3, and becoming low, broad folds between suture and shoulder angulation on last whorl; micropustules weak, scattered over entire surface of first 2 whorls, but most concentrated near suture; last whorl with 3-4 spiral threads between sub- sutural and shoulder angulations, 5-7 spiral threads be- tween shoulder angulation and peripheral carina, and 4- 5 spiral threads and distinct axial rugae between pe- bS THE NAUTILUS, Vol. 105, No. 3 23 Figures 22-27. 22-24. LameUitrochus bicoronatus new species, paratype from off Barbados, 183 m (USNM 859436). 22. .\pertural view, 13.1 X. 23. Basal view, 14.3 x. 24. Protoconch, 119 x. 25-27. LameUitrochus inceratus new species. 25, 26. Paratype from Cerda Station G-967, 24°15'N, 82°26'W, 499-503 m (UMML 30.8062). 25. Apertural view, shell height 8.3 mm. 26. Ba.sal view, shell diameter 7.0 mm 27. Protoconch of paratype from Albatross Station 2644, 25°40'N, 80°00'W, 353 m (USNM 859432), 81 X. ripheral and basal carinae. Base flat, circumscribed by strong, smooth spiral carina; 6-8 weak to strong, smooth spiral cords present between basal carina and 2 strong, tuberculate circumumbilical cords; umbilicus broadly open, about 40% ma.ximum shell diameter, funnel-shaped, walls with 2-4 weak spiral cords and weak axial rugae. Aperture oblique, circular, lips thin; peristome complete. Holotype: USNM 859435, height 3.55 mm, 3.20 mm. Type locality: Off Barbados, Blake Station (data un- recorded), 183 m. Paratypes: 4 specimens, USNM 859436; from same lot as holotype. Hcmarks: Shells of L. bicoronatus differ from those of all other Larnetlitrochiis species in having most of the first teleoconch whorl devoid of axial riblets, and having a double spiral row of strong, rounded tubercles forming the subsutural angulation, and a similar double spiral row of tubercles circumscribing the umbilicus. LameUitrochus inceratus new species (figures 25-27, 36) Calliostoma tiara: Dall, 1889a;.365 (partim). Solariella amalrilis: Dall, 1889a;378, 379 (partim). Solariella (Solariella) lamellosa: Quinn, 197940-42, figs. 61. 62 (partim). Description: Shell small, attaining 8.2 mm height, 7.3 mm width, conicoturbinate, peripherally carinate, broadly umbilicate, white, nacreous under thin outer porcelaneous layer. Protoconch 350-375 /um maximum diameter. Teleoconch whorls 6.0, tubular, shouldered, carinate; first vvhorl with strong, sharp, axial lamellae extending from suture to suture, rapidh becoming re- stricted to subsutural angulation and peripheral carina on subsequent whorls; micropustules rather weak, cov- ering entire surface of first 2 whorls; first whorl with 2- 4 weak spiral threads, two of which later interact with axial lamellae to form tuberculate subsutural angulation and peripheral carina; 0-5 very weak spiral threads pres- ent between subsutural angulation and peripheral carina; spiral threads absent between peripheral and basal ca- rinae. Base flat, circumscribed b\ strong, smooth spiral carina; 5-11 weak, flat, smooth spiral cords present be- tween basal carina and strong, tuberculate circumum- bilical cord; center part of base often smooth; umbilicus broadly open, 45-50% maximum shell diameter, funnel- shaped, walls with 0-4 weak spiral cords and w eak axial rugae. Aperture oblique, circular, lips thin; peristome complete. J. F. Quinii, Jr., 1991 Page 89 ^ '^fid^SS^ '^^^^^^1 1 1 ^^f ^p^ ^ H 3^ ^i^s^ g 3 i^f^^i ^ u Figures 28-36. Radulae of Lamellitrochus species. 28-30. Lamellitrochus lamellosus (Verill & Smith, 1880) from specimen in Figure 1. 28. .Anterior segment, 339 x, 29. Middle segment, 339 x. 30. Anterolateral area, 690 x, 31-33. Lamellitrochus pourtalesi (Clench & .Aguayo, 1939) from specimen in Figure 6. 31. Middle segment, 240 x , 32. Rhachidian and laterals, 600 x . 33. Anterolateral area, 600 x. 34, 35, Lamellitrochus carinatus new species, from paratype from Eolis Station 360, off Fowey Rocks, Florida, 183 m (USNM 859431). 34. Anterior segment, 600 x .^ 35. Same, 1,200 x. 36. Lamellitrochus inceratus new species, from paratype in Figure 27, right anterior fragment, 169 x. Hololype: USNM 94946, height 7.05 mm, width 6.90 mm Type locality: Off Cape Florida, Key Biscavne, Florida, Albatross Station 2644, 25°40'N, 80°bo"W, 353 m. Paratypes: USNM 108141, 1 specimen; USNM 754272, 1 specimen; Albatross Station 2668, 30°58'30"N, 79°38'30"W, 538 m.— UMML 30.8051, 2 specimens; Gerda Station G-300, 26°16'N, 79°30'W, 640 m.— UMML 30.8025, 1 specimen; Gerda Station G-4, 25°49'N, 79°59.5'W, 256 m.— UMML 30.8338, 2 specimens; John Elliott Pillsbitry Station P-1309, 25'"44.5'N, 79°50.0'W, 311 m.— UMML 30.7565, 1 specimen; Gerda Station G-830, 25°40'N, 79°59'W, 342 m.— USNM 859432, 6 specimens; USNM 330559, 4 specimens; same locality as holotype.— UMML 30.8099, 2 specimens; Gerda Station G-23, 25°32'N, 79°44'W, 768 m.— UMML 30.8042, 2 specimens; Gerda Station G-857, 25°22'N, 80°03'W, 194- 186 m. — UMML 30.7538, 1 specimen; Gerda Station G-834, 25°15'N, 80°10'W, 86-79 m.— UMML 30.7914, 1 specimen; Gerda Station G-1035, 24°34.7'N, 80°58.6'W, 254-358 m.— UMML 30.7770, 8 specimens; Gerda Sta- tion G-970, 24°24'N, 82°08'W, 512 m.— UMML 30.8063, 1 specimen; Gerda Station G-969, 24°18'N, 82°33'W, 269- 402 m. — L'MML 30.7644, 1 specimen; Gerda Station G-968, 24°17'N, 82°34'W, 499-503 m.— UMML 30.8062, 6 specimens; Gerda Station G-967, 24°15'N, 82°26"W, 499-503 m — IIMML 30.8065, 1 specimen; Gerda Sta- tion G-1099, 24°12.5'N, 82°50'VV, 622 m.— UMML 30.8058, 1 specimen; Gerda Station G-861, 24°08'N, 8r36'W, 514-558 m.— USNM 421840, 5 specimens; Waldo Schmitt Station 69, off Dry Tortugas, 455-655 m.— USNM 94947, 4 specimens; Blake Station 2, 23°14'N, 82°25'W, 1,472 m.— USNM 94948, 4 specimens; Blake Station 21, 23°02'N, 83°13'W, 525 m.— USNM 859433, 1 specimen; Blake Station (data unrecorded), Yucatan Channel, 1,170 m.— USNM 94058, 3 specimens; Alba- tross Station 2150, 13°34'45"N, 8r21'10"W, 699 m.— I'.i- THE NAUTILUS, Vol. 105, No. 3 USNM 94106, 1 specimen; Albatross Station 2135, 19''55'5S"N, 75''47'07"W, 457 m.— USNM 429872, 5 specimens; USNM 429873, 2 specimens; USNM 429895, 2 specimens; Johnson-Smithsonian Deep-Sea Expedition Station 94, 18°37'45"N, 65°05'00"W, 549-860 m.— UMML 30.8339, 1 specimen; John Elliott Pillsbtiry Sta- tion P-929, 15°29.5'N, 61°11.5'\V, 457-503 m.— USNM 94949, 2 specimens; Blake Station 211, 14°28'40"N, 61°06'08"W, 653 m.— UMML 30.8340, 8 specimens; John Elliott Pillsbury Station P-905, 13°46.3'N, 61°05.4'W, 384-963 m.— UMML 30.8341, 13 specimens; John El- liott Pillsbury Station P-904, 13°45.5'N, 6r05.7'\V, 201- 589 m.— UMML 30.8342, 1 specimen; yo/in Elliott Pills- bury Station P-903, 13°44'N, 61°03.1'W, 231-430 m. Remarks: Shells of Lamellitrochus inceratus are similar in shape and sculpture to that of L. suavis, but are much larger (up to 8.2 mm vs. 3.35 mm, respectively) and have larger protoconchs (350-375 ^m vs. 325 nm, respective- ly). Shells of L. pourtalesi (figures 4-6) are most similar to those of L. inceratus, but are somewhat larger (to 10.3 mm), have much larger protoconchs (525-550 ^m), have a narrower umbilicus (about 35% vs. 45-50% of shell width, respectively), lack axial riblets on the first 0.5 whorl, the aperture is ovate rather than circular, and the tubercles on the subsutural angulation and peripheral carina are sharply conical and axialK elongate rather than rounded and spirally elongate as in L. inceratus. Lamellitrochus inceratus occurs off southern Georgia; in the Straits of Florida from off Cape Florida to the Dry Tortugas, and off Havana and Bahia Honda, Cuba; in the Yucatan Channel; off Old Providence Island; off southeastern Cuba; off northern Puerto Rico; and the Lesser Antilles from Dominica to St. Lucia. This species is usually collected in depths of about 250-550 m. ACKNOWLEDGMENTS I thank Riidiger Bieler (DMNH), the late Gilbert L. Voss (UMML), and Robert Hershler and M. G. Harasewych (USNM) for access to the collections under their care, and Jose H. Leal for bringing the Brazilian specimens of Lamellitrochus carinatus to my attention and providing the SEM micrographs of one of those specimens. Earnest Truby (Florida Marine Research Institute) assisted with SEM operations and provided the final prints of the other SEM micrographs. Marjorie Myers (Florida Marine Re- search Institute) word-processed the various versions of the manuscript. Thomas H. Perkins and William G. Ly- ons (both Horida Marine Research Institute), and two anonymous reviewers, provided valuable critiques of previous drafts of this paper. LITERATURE CITED Abbott, H T 1974. .American seasheils, 2nii cil \ an Nostrand Reinhold, New York, NY. fifi3 p .'Vdam.s, A. 1860. On some new genera and species of Moilusca from Japan. Annals and Maga/inc of Natural Histors. Series 3, 6:331-33". Oarcelli'S, .\. R. 19.53. .Nuevasespeciesde gastropodos inarnios de las republicas oriental del Uruguay y Argentina. Co- municaciones Zoologicas del Mu.seo de Historia Natural de Montevideo 4(70): 1-16. Clencli, W J, and C G. .'\guayo. 1939. Notes and descriptions of new deepwater Moilusca obtained by the Harvard-Ha- bana Fxpedition off the coast of (Alba II Meinorias de la Sociedad Cuhanade Historia Natural "Felipe Poey" 13(3): 189-197. Dali, VV. H. 1881. Reports on the results of dredging, under the supervision of Alexander .-Vgassiz, in the Gulf of Mex- ico, and in the Caribbean Sea, 1877-79, by the United States Coast Survey steamer "Blake", Lieutenant-Com- mander C. D. Sigsbee, U S N., and Conunander J. R. Bart- lett, U.S.N., commanding. W. Preliininar\ report on the Moilusca. Bulletin of the Museum of t^oniparatixe Zool- ogy, Harvard University 9(2):33-144. Dall, W. H. 1889a. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U.S. Coast Survey steamer "Blake", Lieut-Commander C. D. Sigsbee, U.S.N , and Commander J. R. Bartlett, U.S.N., commanding. XXIX Report on the Moilusca. Part 2. Gas- tropotla and Scaphopoda, Bulletin of the Museum of Com- parative Zoology, Harvard University 18: 1-492. Dall, W. H. 1889b. A preliminary catalogue of the shell- bearing mollusks and brachiopods of the south-eastern coast of the United States, with illustrations of many of the species. United States National Museum Bulletin .37:1-221. Dall, W. H. 1903 .\ preliminary catalogue of the shell-bear- ing marine mollusks and brachiopods of the southeastern coast of the L'nited States, with illustrations of many of the species. Reprint, to which are added 21 plates not in the edition of 1889. United States National Museum Bul- letin 37(new ed.):l-2.32, Dall, W. H. 1910. New shells from the (iulf of California. The Nautilus 24(3):32-34. Dall, W. H. 1927. Small shells from dredgings off the southeast coast of the United States by the L'nited States Fisheries steamer "Albatross" in 188.5 and 1886 Proceedings of the United States National Museum 70(2667): 1-134. D Attilio, A. and G. E. Radwin. 1971. The intritacalx. an undescribed shell layer in mollusks. The Veliger 13(4):344- 347- Finlav. II J 1927 .\ further commentarx on New Zealand molluscan systematics. Transactions anti Proceedings of the New Zealand Institute 57:320-485. Frelter, V. and \. Graham. 1977. The prosobranch molluscs of Britain and Denmark. Part 2. Trochacea. Journal of Molluscan Studies, Supplement 3:.39-I00, Herbert, D C. 1987. Revision of the Solariellinac (Moilusca: Prosobranchia: Trochidae) in southern .Africa. .Amials of the Natal Museum 28(2):28.3-382. Herbert, D. G. 1989. Pagodatrochus. a new genus for Miiwlia variabilis H .\dams, 1873 (Gastropoda: Trochidae). Jour- nal of Molluscan Studies 55(3):365-372. Hickman, C S and J H McLean 1990. Sy.stemalic revision and suprageneric classification of trochacean gastropods. Natural Histor\ Museum of Los .\ngeles Coimty, Science Series No. 35:1-169. Iredale, T. 1924. Results from Roy Bell's molluscan collec- tions. Proceedings of the I.innean Society of New South Wales 49:179-278. Ircdalc, T. 1929. Moilusca from the conliiiciital shell of east- J. F. Quinn, Jr., 1991 Page 91 ern Australia. No. 2. Records of tlit- .Viislralian Museum 17(4):157-190. Johnson, C. W. 1934. List of marine Mollusca of tlie .Atlantic coast from Labrador to Texas. Proceedings of ttie Boston Society of Natural History 40:1-204. McLean, J. H. 1971 [Trochidae]. In: Keen, .\. M. (ed.). Sea- shells of tropical west America, 2nd ed. Stanford I'niver- sity Press, Stanford. CA, p. 329-342. Philippi, R. A. 1836. Enumeratio molluscorum Siciliae emu viventium turn in tellure tertiaria fosilium, quae in itinere suo observavit, Vol. 1. Berolini. Simonis Schroppii et Soc iv + 267 p. Pilsbry, H. A. 18S9-1S90. Manual of concliology. Vol. 11. Trochidae, Stomatiidae, Pleurotomariidae, Haliotidae. Conchological Section, .Academy of Natural Sciences, Phil- adelphia, "PA, 519 p. Porter, H. J. 1974. Mollusks from M.'V Eastward Stations 11542 and 11545 east of Charleston, S.C. Bulletin of the •American Malacological Union for 1973:20-24. Quinn, J. F , Jr. 1979. Biological results of the University of Miami Deep-Sea Expeditions. 130 The systematics and zoogeograph) of the gastropod family Trochidae collected in the Straits of Florida and its approaches Malacologia 19(l):l-62. Quinn, J. F., Jr. In press. The Trochidae of the Gulf of Mexico (Prosobranchia: Archaeogastropoda). Memoirs of the Hourglass Cruises. Rice, W. and L. S. Kornicker. 1965. Mollusks from the deeper waters of northwestern Campeche Bank, Mexico. Publi- cations of the Institute of Marine Science, University of Te.xas 10:108-171 Seguenza, G. 1903. Molhischi poco noti dei terreni terziarii di Messina. Societa Geologica Italiana, Bolletinn 7179- 189. Tlnele. J 1925 Gastropoda der deutschen Tiefsee-Expedi- tion. II Teil. Wissenschaftliche Frgebnisse der deutschen Tiefsee-Expedition auf dem Danipfer "Valdivia" 1898- 1899, 17(2):35-382. Treece, C, D 1980. Bathymetric records of marine shelled Mollusca from the northeastern shelf and upper slope of Yucatan, Mexico. Bulletin of Marine Science 30(31:552- 570, V'errill, .A. E. 1880. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of New England American Journal of Science 20(3):390-403. \errill, A E. 1880-1881. Notice of recent additions to the marine Invertebrata of the northeastern coast of .America, w ith descriptions of new genera and species and critical remarks on others. Part II Mollusca, with notes on .An- nelida, Echinodermata, etc., collected by the L'niled States P^ish Commission. Proceedings of the L'nited States Na- tional Museum 3:356-405. Verrill, A E. 1881. Notice of recent additions to the marine Invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. Part III Catalogue of Mollusca recentK added to the fauna of southern New England Proceedings of the United States National Museum 3:405-409. \'errill, .A. E. 1882. Catalogue of marine Mollusca added to the fauna of the New England region during the past ten years. Transactions of the Connecticut Acadenu of .Arts and Sciences 5:447-587. Verrill, A. E. 1885. Results of the explorations made b\ the steamer "Albatross", off the northern coast of the L'nited States, in 1883. Report of the L^nited States Commissioner of Fish and Fisheries for 1883:503-699. Wood, S. V. 1842. -A catalog of shells from the Crag. .Annals and Magazine of Natural History, Series 1, 9:527-544. THE NAUTILUS 105(3):92-103, 1991 Page 92 Tridacna tevoroa Lucas, Ledua and Braley: A Recently-described Species of Giant Clam (Bivalvia; Tridacnidae) from Fiji and Tonga J. S. Lucas Zoology Department James Cook University Townsville, Q. 4811, Australia E. Ledua Fisheries Division Ministry of Primary Industry Suva, Fiji R. D. Braley Zoology Department James Cook University Townsville, Q. 4811, Australia ABSTRACT Tridacna tevoroa Lucas, Ledua and Braley is the eighth extant species of giant clam. It has only been found in the eastern Lau Islands of Fiji and the northern Vava'u and Ha'apai islands of Tonga. It inhabits relatively deep water (20+ m) compared to other giant clam species and is apparently intolerant of shallow conditions. Tridacna tevoroa occurs sympatrically with T. de- rasa (Roding) and appears to be most closely related to it, both belonging in the subgenus Persikima Iredale. The two species are readily separated by the presence in T. tevoroa of a rugose mantle, prominent guard tentacles on the incurrent aperture, thinner valves, and colored patches on the shell ribbing near the umbonal area. Tridacna tevoroa has some Hippopus-like characters: shell pigmentation, absence of byssal gape, lack of lateral mantle projection, and absence of hyaline organs in the mantle. These necessitate modification of Rosewater's diag- nostic features distinguishing the two genera of extant giant clams, Tridacna Bruguiere and Hippopus Lamarck. Key words: Giant clam, Tridacnidae, Tridacna. taxonomy, key, Hippopus. INTRODUCTION Rosewater (1965) reviewed the confused taxonomy of giant clams (Order Veneroida, Superfamily Tridacnacea, Family Tridacnidae), recognizing six extant species in two genera, Hippopus Lamarck, 1799 and Tridacna Bru- guiere, 1797. He subsequently described a further Hip- popus species (Rosewater 1982). Thus, after Rosewater's pivotal work, the family consisted of seven extant species, H. hippopus (Linnaeus, 1758), H. porcellanus Rosewa- ter, 1982, T. gigas (Linnaeus, 1758), T. dcrasa (Rciding, 1798), T. squamosa Lamarck, 1819, T. maxima (Roding, 1798) and T. crocea Lamarck, 1819. This taxonomic revision of the family Tridacnidae was very timely, as it preceded the period of intense interest in giant clams in recent years. Before 1970, there was little research interest in giant clams, despite it being known that giant clams are unique among bivalve mol- lusks in having hypertrophied siphonal tissues packed with symbiotic algae (zooxanthellae) (Lucas, 1988). Then, in the 1970's, there was a surge of research. This resulted partly from academic interest in their unique features, but mainly from interest in farming giant clams. It was recognized that giant clam stocks were being heavily over-exploited through most of their geographic ranges and that some species had been fished to extinction in particular regions. Furthermore, it was found that giant clams grow more rapidly than previously envisaged (Munro & Gwyther, 1981). Farming them as a means of supplying the demands for giant clam meat and other products, and thus taking pressure off natural popula- tions, became a desirable and feasible proposition. Rosewater's taxonom\ of giant clams has been widely accepted and validated through use in species' identifi- cations in numerous studies. However, at an international meeting on giant clams in 1988, the possibility of a rare, new species in Fiji was raised (Lewis & Ledua, 1988). This new taxon was subsequently found also in Tonga and recentlv described as Tridacna tevoroa by Lucas et al. (1990). This paper provides further description of T. tevoroa, including morphometric comparison with T. derasa. It also considers the affinities of the new species within the family Tridacnidae. GENERA AND SUBGENERA OF TRIDACNIDAE In order to consider the affinities of T. tevoroa within the family Tridacnidae, it is necessary to review Rose- water's diagnoses of Hippopus and Tridacna. and sub- genera of the latter. Rosewater (1965) recognized three subgenera of Tridacna: Tridacna s.s. Bruguiere, 1797 (containing T. gigas). Persikima Iredale, 1937 (contain- ing T. derasa). and Chametrachea Morch, 1853 (con- taining T. squamosa, T. maxima, and T. crocea). As the diagnostic features for the various taxa are not clearly J. S. Lucas et ai, 1991 Page 93 Table 1. Diagnostic features of Tridacna and Hippopus frm HoM-uater (1965), with additions and modifications Feature Tridacna Uippupus Hyaline organs in mantle surface B\ssal orifice Tightly fitting teeth along byssal orifice Posterior adductor muscle scar loca- tion Outer demibranch of ctenidia Klongate- triangular shape' Strawberry blotches on earl\ portion of shell Mantle projects later- ally over shell margins^ Distinct ventral re- gion of shells' present (usually absent present)' well defined (poor- poorK defined ly-well dehned)' absent present si bcliiiu! inid-liiic central complete-incom- plete no not present (usual- 1\ not present)' yes (usually)' not present complete (?)- yes (in large specimens)' present present ' Modification reiiuired to accommodate T. tcvuroa. - It is presumed that Rosewater was referring to the presence or absence of a distal food groove in describing the 300 mm) specimens. Umbo angle > 150°; umbo slightly opisthogyrous. Valves moderately inflated, thin at margins but distinctly thick- ened in umbonal region in large specimens. Shell colored gra\-white externally, usually with strawberry colored patches on rib-like radial folds in umbonal area (extend- ing up ca. 100 mm from umbo), giving ribs a striped appearance. Strawberry patches are not evident in some old specimens (including holotype), apparently lost through erosion or obscured by calcareous overgrowths of umbonal region (c/. figure 1 1 ). Primary radial sculp- ture consisting of 6 or 7 prominent folds, plus 3 short thin ridges near ligament. Prominent rib-like folds grad- ing into undulating folds near shell edge in large spec- imens. Secondary radial sculpture consisting of fine ribs, much stronger in fold interstices than on radial folds; fine ribs not evident in distal shell region of large spec- imens. Apparent remains of eroded scutes often present on rib-like folds (figure 11) from umbo to extent of straw- berry colored striping; occurring on white regions be- tween colored patches. Other concentric sculpture lim- ited to fine growth lines. Upper shell margins weakly undulating, with 5 rounded medially-projecting pro- cesses, representing extremities of rib interstices; upper shell margins often thin and sharp (figure 8). Length of hinge line ca. 0.6 x shell length. Umbo strongly back- ward of mid-point of shell length. Cardinal tooth and cardinal socket (figure 7) not strongly developed, even in large specimens. Lateral hinge tooth little more than a ridge. Ventral shell margin somewhat irregular and slightly convex. Byssal orifice poorly defined; without teeth; with ca. 5 rows of small plicae in byssal orifice region. Interior of shell white; gloss\' distal to pallial line, dull medial to pallial line. Hinge plate with pale brown tinge. Pallial line strongly indented in ventral region. Posterior adductor muscle scar large and strongK' indi- cated with lines and ridges; ca. 70 mm high and 55 mm wide in 435 mm shell length specimen; sited behind mid- point of shell length but in front of umbo on long shell axis. Posterior pedal retractor muscle scar adjacent to foremost border of posterior adductor scar and much smaller, <0.2x area of posterior adductor scar. Soft parts: Mantle of subdued color, brown, gray-brown or green-brown, uniform in color or mottled; without stri- ations, but sometimes with pale lines following mantle surface contours (figure 10). Mantle surface rugose, with numerous protuberant lobes (figure 10). Mantle tissue with few iridophores, without hyaline organs, and with most zooxanthellae situated close to mantle surface (fig- ure 16). Mantle not or barely extending lateralK over shell edges. Expanded mantle broad and exposed by wide gaping of valves. Incurrent aperture with long, broad guard tentacles, usually white, but sometimes brown (presumably through presence of zooxanthellae), often projecting prominently out from aperture. Byssus absent and foot vestigial in all large specimens (300-1- mm shell length). Ctenidia demibranchs extending forwards be- yond proximal oral grooves for ^-'t of their length. Outer demibranch of similar size or slightly less than depth of inner demibranch; outer demibranch may be upwardly reflected in some regions and the ctenidial axis is exposed in some regions. Inner demibranch with food groove and outer demibranch without. Long distal oral grooves lead from ctenidia to labial palp bases. Morphometries: In comparing the morphometries of T. tevoroa and T. derasa. seven parameters were mea- POSTERIOR ... ( = siphonal region) POSTERO-VENTRAL (= hindmost) POSTERO-DORSAL (= foremost) VENTRAL ( = pedal gape) DORSAL DORSAL (= ligament region) ANTERIOR (= foremost end of pedal gape to umbo) POSTERIOR ANTERIOR VENTRAL "Fs" platform umbonal region thickness I \^ J umbo Figures 1-5. Diagrams of anatomical terminology for giant clams and shell dimensions measured. 1. Orientation terminology used in this paper (following Stasek, 1962), 2. Alternative orientation terminology (after Rosewater, 1965, 1982; Yonge, 1936, 1980). 3. Lateral view of valve. 4. Upper view of valves. 5. Cross-section of valve. Page 96 THE NAUTILUS, Vol. 105, No. 3 ** --».<. "?'"}'=» Lir^ ■:f^i / '■r' W # J. "^^ .;>_.,■►«'' ~^ _ #• Figures 6, 7. Lateral views of T. tevoroa holotype left valve. 6. E.xtenial surface, 7. Internal surface. J. S. Lucas et ai. 1991 Page 97 Figures 8, 9. T. tevoroa holotype. 8. Upper view. 9. Umbonal view. sured: 1. shell length (figure 3); 2. shell height (figure 3); 3. shell width of conjoined valves (figure 4); 4. shell weight; 5. underwater shell weight; 6. umbonal region thickness (maximum distance through the umbonal region) (figure 5); 7. distal shell thickness (measured 25 mm in from the margin of the medial radial fold). Only measurements 1, 2 and 3, and sometimes 7 could be made on living specimens. The shell weight (4) in living specimens was estimated from the underwater weight of the whole an- imal, which was assumed to be entirely due to shell (=5), the soft tissues having approximately neutral buoyancy. shell weight = 1.611 x underwater weight This relationship between shell weight and underwater shell weight was found to be very regular (1.611 ± 0.008 s.d.) in T. derasa and T. tevoroa. Parameters 2-7 were related to shell length (1), which was used as the general measure of size. Least squares regressions were calculated for each of parameters against shell length for each species (table 3). The slopes and intercepts for each pair of regressions, and their standard errors, were compared by t-tests (2-tail) for statistically significant differences. There were no significant differ- ences between the intercepts for each pair of regressions. Slopes were significantly different (P < 0.05) between species for three of the regressions: shell width, umbo Page 98 THE NAUTILUS, Vol. 105, No. 3 ^:;^.'^ Figures 10, 1 1. Live T. tevoroa specimens. 10. Rugose mantle; ex, excurrent aperture; t. conspicuous guard tentacles on incurrent aperture. 1 1. Umbonal regions. Note that the umbonal region of the specimen in the foreground is heavily eroded and encrusted, showing little striping of the radial ribs, but striping and apparent eroded scutes are clear on the rear-left specimen (arrowed); striping is clear on the rear-right specimen (arrowed). J. S. Lucas et al., 1991 Page 99 ;)oo £ 250 -O JOO Oj 12 J 200 301) 100 r,00 600 Shell leiifith (nun) 0) C o 40 13 • 20 - • / 00 - • t / / • • 80 - • /% •• • 60 40 pn - • 1 • / ▲ • A A 1 200 300 (00 500 Shell length (mm) 600 40 35 30 Oil ~- 25 10 14 • " • - • , - / / A ^ ' _ -^.A ^'A ^ ^ '*A • /mm A 1 1 1 J 200 300 400 500 Shell length (mm) 600 I'S^".-. ^r . t '.Tt^-.'. »> "•♦ ' > H" Figures 15, 16. Cross-sections of surface layers of mantle tissues. 15. r. derasa. 16. T. tevoroa. i, iridophores; z, zooxan- tfiellae. Scale bars = 100 micrometers. thickness and shell weight versus shell length (table 3). These data are figured in figures 12-14. Types: Holotype: Australian Museum, Sydney, Reg. No. C164071. Whole specimen with soft tissue preserved Figures 12-14. Morphometric comparisons of T. tevoroa (tri- angles and dashed regression line) and T. derasa (circles and continuous regression line). 12. Shell width versus shell length. 13. Umbo thickness versus shell length. 14. Shell weight versus shell length. Page 100 THE NAUTILUS, Vol. 105, No. 3 Table 3. Morphometries of T. tevoroa and T. derasa. Coefficients and standard errors (SE) for "a" (slope) and "b' (Y axis intercept) for least squares regressions of Y = aX + b, where X is shell length (nun). Sample numbers (n) for T. tevoroa and T. derasa ranged from 18-35 ami 9-16, respectively. P, significance level ol correlation; vs, very significant (P "C 0 01); *, slope coefficients differ significantl)' between species at P < 0.05 level. 7' ridacna tevoroa Tridacna derasa Y a/b Coeff. SE P Coeff SE P Shell width (mm) a 0.307* 0.071 vs 0.613* 0.061 vs b 45.0 29.4 0.2 -13.77 25.34 0.6 Shell height (mm) a 0.548 0.075 vs 0.657 0.058 vs b 23.05 31.20 05 -11.42 24.57 0.6 I'mbo thickness (mm) a 0.026* 0.034 0.5 0.324* 0.034 vs b 40.8 14.8 0.02 -54.93 1449 vs Shell margin thickness (mm) a 0 0072 0 0145 0.6 0031 0.008 vs b 5.52 6.26 0.4 1.46 3.41 0.7 Shell weight (kg) a 0.0649* 0.0053 vs 0.114* 0.012 vs b -17.17 219 vs -30.29 4.78 vs Underwater weight (kg) a 0.0418 0 0040 vs 0.0716 0 0115 vs 1, -11 33 1,59 \s -IS 95 471 \s in formalin (figures 6-9). Shell length 485 mm. Collected at Cakau Tabu Reef, Southern Lau islands, eastern Fiji, 17°40'S, 178°32'W, A. Lewis and Filipe Viala, 26 January 1986. Paratypes: 1. Fiji Museum, Suva, Reg. No. C-001. Whole specimen with soft tissue preserved in formalin. Shell length 328 mm. Collected at Vatoa Island, Southern Lau islands, eastern Fiji, 19°49'S, 178°13"W, E. Ledua and M. Tavisa, 23 November 1989. Habitat details: at 24 m depth on gentle slope outside the reef on the south- ern side of the island; only low, hard corals were present and scattered in low numbers; very clear water. 2. Na- tional Museum of Natural History, Smithsonian Insti- tution, Washington, DC, USA, Reg. No. USNM 859439. Specimen consisting of valves only. Shell length 389 mm. Collection and habitat details as for paratype 1. 3. Mu- seum of Tropical Queensland, Townsville, Australia, Reg. No. MO 17791. Specimen consisting of valves only. Shell length 413 mm. Collection and habitat details as for paratype 1. Type locality: Cakau Tabu Reef, Lau Islands, Fiji. Distribution: Lau Islands, Fiji (known from Cakau Tabu, Komo Island, Namuka Island, Vatoa Island and Vuata Vatoa, 17°40'S-19°56'S, 178°13'W-178°32'W; Lewis & Ledua, 1988). Islands within the Ha apai and Vava u groups, Tonga. Habitat: From the limited collection details, the habitat of T. tevoroa appears to be along outer slopes of leeward reefs, in very clear, oceanic water at 20-30 m. Tridacna derasa is also found in this kind of habitat, and some specimens may occur down to these depths in very clear water. Material examined for morphometries: T. tevoroa: Fiji, Lau Island group: 440 mm, single valve; 392 mm, single valve; 413 mm, valves only; 548 mm, valves only; 389 mm, valves only; 375 mm, live specimen; 354 nmi. live specimen. Tonga, Vava'u Islands: 560 mm, live spec- imen; 326 mm, live specimen; 420 mm, live specimen; 376 mm, live specimen; 313 mm, live specimen. Tonga, Ha'apai Islands: 408 mm, valves only; 412 mm, valves only. T. derasa: Fiji, Makogai Island: 538 mm, valves only; 425 mm, valves only; 370 mm, valves only; 368 mm, live specimen; 348 mm, live specimen; 317 mm, live specimen. Tonga, Tongatapu: 490 mm, valves only; 376 mm, valves only; 434 mm, valves only; 435 mm, valves only; 323 mm, valves only; 294 mm, valves only; 291 mm, valves only; 474 mm, valves only. Tonga, Fale- vai, Vava'u group: 415 mm, live specimen; 293 mm, live specimen; 332 mm, live specimen; 471 mm, live speci- men. Australia, Great Barrier Reef, Bramble Reef: 435 mm, valves only; 500 mm, valves only; 490 mm, valves only; 440 mm, valves only; 470 mm, single valve; 303 mm, live specimen; 415 mm, live specimen; 435 mm, live specimen; 340 mm, live specimen; 430 mm, live specimen; 450 mm, live specimen; 465 mm, live speci- men; 500 mm, live specimen. Etymology: The species name is derived from the clam's Fijian name, "tevoro " (devil), with a latin termination (nominative, feminine). Comparison: Tridacna tevoroa is most like T. derasa in general appearance, with the latter species occurring in shallower reef areas at all the localities where T. te- voroa has been collected. The similarity of these species goes beyond superficial appearance: in those features by which the subgenus Persikima is distinguished from oth- er subgenera of Tridacna (table 2). T. tevoroa belongs with T. derasa in Persikima. It is, however, readily dis- tinguished from T. derasa b\ a number of features re- lating to both hard and soft anatomy (table 4). Among these distinguishing features, the most conspicuous are the rugose mantle, prominent guard tentacles on the J. S. Lucas ct ai, 1991 Page 101 Table 4. Features which distinguish T. tevoroa from T. derasa. (These features apply to specimens ca 250 mm shell lenijth and greater.) Feature T. tfcoroa T. dcrd.'ici Byssal orifice Umhniial region of rib-like primary f.ilds Mantle Iridophores in man- tle tissue Hyaline organs Mantle projecting fie- yond shells Guard tentacles on incurr. apert. Demibranchs of cte- nidia Cardinal tocitli poorK defined striped with color, appar- entl\ with eroded scutes rugose, with protuber- ances low density (mantle dull colors) absent not or barely long, broad, prominent similar size small, but clearly defineil uncoloretl. with- out scutes smooth, plain high density (mantle often bright colors) present short, tlun, not prominent outer smaller not prominent \er\ prominent incurrent aperture, thinner valves, and colored patches on the shell ribbing of T. tevoroa. The three shell parameters versus shell length for which regression slopes differ significantly between T. tevoroa and r. derasa (table 3) are interrelated, i.e., they reflect the thinner shell of T. tevoroa compared to T. derasa. There is, however, little difference in shell thickness be- tween medium-sized specimens of the two species, ca. 300 mm shell length; interspecific differences become pronounced in the largest (and probably oldest) speci- mens, 450+ mm (figures 12-14). The sharpness and thin- ness of the shell margins of T. tevoroa, noted by Lewis and Ledua (1988), are certainly evident in some speci- mens (figure 8), but its shells soon thicken behind the margin and there is no significant difference in regression slopes between the species for distal shell thickness, 25 mm behind the margin, versus shell length. Stasek (1962) described the ctenidia of some tridacnid species from Eniwetok Atoll in detail, distinguishing T. derasa and H. hippopus from other species in having food grooves on both inner and outer demibranchs. Tri- dacna derasa was further distinguished from other spe- cies in having similar-sized demibranchs, with the outer demibranch upwardly reflected, exposing the ctenidial axis. In view of this reported distinctiveness of the cte- nidia of r. derasa, it would seem to be appropriate to compare these structures in T. tevoroa and T. derasa. However, Rosewater (1965) described T. derasa as hav- ing an "incomplete" outer demibranch, and recent stud- ies of tridacnid morphology by J. Norton (personal com- munication) have revealed that T. derasa specimens from Fiji and Tonga, at least, have short outer demibranchs without food grooves. The degree of upward reflection of the outer demibranch and exposure of the ctenidial axis vary between specimens; they ma\' partlv reflect handling during dissection. Thus, the distinctiveness of ctenidial morphology in T. derasa observed by Stasek is questionable, and comparisons with T. tevoroa are made on the basis of Rosewater s and Norton's observations. Tridacna tevoroa has some features typical of Hip- popus in the Hippopus /Tridacna separation based on Rosewater (1965) (table 1 ). Its byssal aperture is indistinct and it has reddish markings on the external surfaces of the shells. Its dull mantle color, lack of hyaline organs and mantle posture (only reaching the shell margins but offset by wide gaping of the valves) are also like Hip- popus but unlike Tridacna, sensu lata. However, these features of T. tevoroa are never completely like those of Hippopus. Its byssal aperture differs from the Hippopus species in not having interlocking teeth, nor does it have the distinctive "horse-shoe" ventral surface of the valves. It is unlikely that attached juveniles of T. tevoroa orient to the substrate in the characteristic manner of Hippopus juveniles. While T. tevoroa shares shell color patches with the Hippopus species, its colored stripes on the radial ribbing are unlike the irregular color patches of Hip- popus species. The mantle of T. tevoroa is distinct from that of all other tridacnids. It is conceivable that T. tevoroa represents a transi- tional stage between these two genera, the divergence of which goes back in the fossil record at least to the lower Miocene (Rosewater, 1965). However, it comfort- ably fits with T. derasa in the subgenus Persikima and we do not propose to give it separate subgeneric or ge- neric status. One consequence of the discovery of T. tevoroa is the need to make some modifications to the characters dis- tinguishing Hippopus and Tridacna (table 1). Small juveniles of T. tevoroa are unknown, but they are likely to be distinctly unlike T. derasa and any other tridacnid. This is because of the colored stripes on the valve ribs and also the probable presence of scutes on the ribs; these characters being present up to about 150 mm shell length. The distinctive appearance is lost in larger specimens. Remarks: Lewis and Ledua (1988) hypothesized that the localized distribution of the "tevoro " clam in eastern Fiji may represent a "relict population on the eastern edge of both its own former range and that of the ap- parently more abundant and successful T. derasa". Their prediction, that other populations might be found, was soon confirmed by collections in Tonga. The known dis- tribution of T. tevoroa is still, however, limited to several island groups over about 300 km distance in the south- western Pacific. At none of these localities is it abundant: in the \'atoa area, Fiji, Lewis and Ledua (1988) estimated its relative abundance compared to T. derasa was on the order of 1:50. It thus must be regarded as the giant clam species most vulnerable to extinction. The previous bear- er of this unfortunate distinction was H. porcellanus, with a relatively restricted distribution centered on the Page iU2 THE NAUTILUS, Vol. 105, No. 3 southern Philippines (Lucas, 1988), and a high demand within the international shell trade despite CITES ({Con- vention on International Trade in Endangered Species) protection. Fortunately, the deep habitat of T. tevoroa partially protects it from human predation, but there is still some exploitation. This is evident from the fact that there are local names for T. tevoroa in the three island groups where it occurs (Lucas et a/., 1990) and there are anecdotal reports of its consumption, i.e.. one diver on N'ava'u Island, Tonga, reported having collected and eat- en about 20 specimens. Fortunately, there is no major trade in giant clam shells in Fiji or Tonga. Tridacna tevoroa has only been collected from rela- tively deep-water habitats for giant clams (20—30 m depth). Their mantle tissue contains symbiotic zoo.xan- thellae like other tridacnids, and presumably photosyn- thetic products from these symbionts are used in their nutrition. However, there are low light levels and a shift- ed light spectrum at the depths they inhabit, and prob- ably the extent to which autotrophy supports the clam's nutrition will be reduced. Thus, T. tevoroa is expected to have morphological adaptations to facilitate using the low light intensities at these depths. Extending the mantle laterally over the shell margins to increase its exposed surface, as in other Tridacna species, is surprisingly not shown by T. tevoroa. However, it achieves a large mantle surface area through the mantle protuberances and wide gaping of its valves. The dull mantle color is due to low densities of iridophores (iridocytes) (c/. figures 15 and 16), with the brown coloration of zooxanthellae showing through. Iridophores in giant clams contain thin reflect- ing platelets (Kamishima, 1990). They are particularly common near the mantle tissue surface in brightly col- ored tridacnids, including some T. derasa (figure 15). Most zooxanthellae are situated below the iridophore layer in T. derasa. In T. tevoroa the zooxanthellae also occur close to the mantle surface (figure 16); clusters of zooxanthellae may occur in small elevations of the mantle surface. The paucity of iridophores and shallow distribution of zooxanthellae in the mantle of T. tevoroa clearly reflect the need to maximize light reaching the zooxanthellae. It has been shown that zooxanthellae pho- to-adapt to low light levels (Mingoa, 1988) and this may be important for T. tevoroa. It seems that T. tevoroa are not only adapted to rel- atively deep conditions, but they are intolerant of shallow habitats. In early collections by Fiji Fisheries staff, these clams were transported on the decks of Fisheries vessels and kept in shallow conditions {ca. 3 m) at Makogai Island. Rapid mortality occurred until the clams were moved into deeper water {ca. 14 m). Solar UV radiation penetrates clear tropical waters sufficiently to have sig- nificant biological effects down to 20 m and it is possible that T. tevoroa lacks or has insufficient of the U\' screen- ing agents that protect organisms inhabiting the shallow regions of coral reefs, e.g., Dunlap et al. (1988). Also, it may have insufficient enzymes that deal with the toxic oxygen levels generated by photosynthesis at high light levels {e.g., Shick & Dykens, 1985). KEY TO THE TRIDACNID SPECIES There are now eight extant species of giant clams and the key to species given by Lucas (1988) needs to be expanded and modified to incorporate T. tevoroa. 1 Byssal orifice region of opposed valves with in- terlocking teeth; distinct region of shell around byssal orifice, outlined by ventral-most pair of prominent radial ribs; mantle, when fully ex- tended, not projecting laterally beyond shell margins Hippopus ... 2 Byssal orifice of opposed valves without inter- locking teeth; no distinct ventral region of shell outlined by prominent radial ribs; mantle, when fully extended, usually projecting laterally be- yond shell margins Tridacna ... 3 2(1) Shells thick and strongly ribbed, with reddish blotches in irregular bands; incurrent apertures without guard tentacles H. hippopus Shells in specimens less than about 200 mm shell length not thick nor strongly ribbed and with only faint reddish blotches; incurrent apertures with guard tentacles H. porcellanus 3(1) Shell length of large specimens >550 mm, sometimes greater than 1 m; with about four elongate, interdigitating projections of each dis- tal shell margin, being most elongate and acute in large specimens; shell without scutes, except for some tubular projections near umbo in very small juveniles; mantle brownish, with numer- ous iridescent blue-green circles T. gigas Shell length rarely >550 mm; without elongate, interdigitating projections on each distal shell margin; mantle variably colored, without iri- descent blue-green circles 4 4(3) Shell length up to 550 mm, occasionally larger; upper region of large shells plain, without scutes or strong ribs; hinge usually longer than half shell length 5 Shell length usually <400 mm; upper shell re- gion with scutes or eroded scutes; hinge equal to or less than half shell length 6 5(4) Rib-like radial folds on shell without colored patches; mantle without protuberances; incur- rent aperture with inconspicuous guard tenta- cles T. derasa Rib-like radial folds on shell usually striped with colored patches near umbo; mantle with pro- tuberances; incurrent aperture with conspicuous guard tentacles T. tevoroa 6(4) Shell approximately symmetrical about umbo in lateral view, with hinge about half shell length; scutes large and well-spaced both within and between the radial rows; lateral distance be- tween scutes in adjacent rows usually about the same as scute width; byssal aperture narrow to moderately wide; not embedded into substrate; mantle usuallv of subdued and mottled color; J, S. Lucas et ai, 1991 Page 103 incurrent aperture with distinct guard tentacles T. squamosa Shells usually asymmetrical about umbo in lat- eral view, with hinge less than half shell length; scutes usually low and often eroded, set close together both within radial rows and between rows; byssal aperture moderately wide to wide; embedded or partK embedded into substrate; mantle brightly colored; incurrent aperture with indistinct guard tentacles 7 7(6) Shell length < 150 mm; shells not strongly asym- metrical about umbo in lateral view; byssal ap- erture wide; scutes eroded away except near shell margin; occurs deepK embedded in reef substrate T. crocea Shell length of large specimens often > 150 mm; shell often strongly asymmetrical about umbo in lateral view; byssal aperture moderately wide to wide; scutes present in substantial part of upper shell region; occurs partially embedded in reef substrate T. maxima ACKNOWLEDGEMENTS This research was part of an international project funded by the Australian Centre for International Agricultural Research (Project No, 8733). We thank the Fisheries personnel in Fiji and Tonga, and local divers, for their assistance with collecting; Dr John Norton for assistance with the histology, the micrographs and for reading an early version of the manuscript; Dr Winston Ponder for advice and for providing the photographs of the holo- type; Dr Ross Alford for advice on statistics and Mr Jeremy Barker for assistance with the figures. LITERATURE CITED Dunlap, W, C, B E. Chalker, and W. M Bandaranayake. 1988. Ultraviolet light absorbing agents derived from tropical marine organisms of the Great Barrier Reef Pro- ceedings of the 6th International Coral Reef Symposium 3:89-93. Kamishima, Y. 1990. Organization and development of re- flecting platelets in iridophores of the giant clam, Tridacna crocea Lamarck. Zool. Sci. 7:63-72. Lewis, A. D. and E. Ledua. 1988, A possiljle new species of Tridacna (Tridacnidae: Mollusca) from Fiji. /;i; Copland, J. VV. and J. S. Lucas (eds.). Giant clams in Asia and the Pacific. ACIAR Monograph No. 9, Australian Centre for International Agricultural Research, Canberra, ACT, p. 82-84. Lucas, J. S. 1988. Giant clams: description, distribution and life-history. In: Copland, J. W. and J. S. Lucas (eds. ). Giant clams in Asia and the Pacific. ACI.\R Monograph No 9, Australian Centre for International Agricultural Research, Canberra, ACT. p. 21-32 Lucas, J S., E. Ledua, and R. D Brale) 1990. A new species of giant clam (Tridacnidae) from Fiji and Tonga. ACIAR Working Paper No 23, Australian Centre for International Agricultural Research, Canberra, ACT, 8 p. Mingoa, S. S. M. 1988. Photoadaptation in juvenile Tridacna gigas. In: Copland, J. W. and J. S. Lucas (eds). Giant clams in Asia and the Pacific. ACIAR Monograph No. 9, Australian Centre for International Agricultural Research, Canberra, ACT, p. 145-150. Munro, J L and J Gw>ther. 1981. Growth rates and mari- culture potential of tridacnid clams. Proceedings of the 4th International Coral Reef Symposium 2 633-636. Rosewater, J, 1965. The family Tridacnidae in the Indo- Pacific. Indo-Pacific Mollusca 1:347-394. Rosewater, J. 1982. A new species of Hippopiis (Bivalvia: Tridacnidae). The Nautilus 96:3-6. Shick, J. M. and J. A. Dykens. 1985. Oxygen deto.xification in algal-invertebrate symbioses from the Great Barrier Reef Oecologia 66:33-41. Stasek, C, R. 1962. The form, growth, and evolution of the Tridacnidae (giant clams). Arch. Zool. Exp, Gen. 101:1- 40. Stasek, C. R, 1963, Orientation and form in the bivalved Mollusca. J. Morph. 112:195-214. Yonge, C. M. 1936. Mode of life, feeding, digestion and sym- biosis with zooxanthellae in the Tridacnidae Scient Rep. Gt. Barrier Reef Exped. 1:283-321. Yonge, C. M. 1980, Functional morphology and evolution in the Tridacnidae (Mollusca: Bi\alvia: Cardiacea), Rec. Aust, Mus. 33:735-777. THE NAUTILUS 105(3):104-115, 1991 Page 104 Dates of Publication and Supraspecific Taxa of Bellardi and Sacco's (1873-1904) "I moUuschi dei terreni terziarii del Piemonte e della Liguria" and Sacco's (1890) "Catalogo paleontologico del bacino terziario del Piemonte" B. A. Marshall National Museum of New Zealand PO Box 467, Wellington, New Zealand ABSTRACT Dates of publication are determined for the serial parts of Bellardi and Sacco's "I molluschi dei terreni terziarii del Pie- monte e della Liguria " and its largely overlooked companion series, the latter containing the earliest valid introductions of many taxa. Genus group taxa introduced by Sacco (1890) in the rare work "Catalogo paleontologico dei bacino terziario del Piemonte' have been hitherto ignored or treated as nomina nuda, although most meet criteria for availability and predate introductions in Bellardi and Sacco's 1873-1904 work. Supra- specific taxa introduced in these works are listed with biblio- graphic references. Attention is drawn to some taxonomic prob- lems that result from recognition of hitherto overlooked valid introductions of names. Key words: Bellardi; Sacco; compilation, dates; supraspecific taxa; Tertiary fossils; Mollusca; Mastodon. INTRODUCTION Bellardi and Sacco s monumental work "1 molluschi dei terreni terziarii del Piemonte e della Liguria " is one of the most comprehensive treatments of a regional Tertiary fauna ever published. Issued between 1873 and 1904 in 30 numbered quarto parts, it contains descriptions of thousands of species and infraspecific taxa, numerous genus group taxa, and several families and subfamilies. The Bellardi and Sacco collection is now in the care of the Museo Regionalc di Scienze Natural!, Turin (Ferrero Mortara et al., 1982, 1984). The primary objective of this contribution is to draw attention to the fact that many names generally consid- ered to date from this work were validly introduced earlier by Sacco in a little-known companion series and in the rare "Catalogo paleontologico del bacino terziario del Piemonte". Parts 1-8, 1 1 and 13a of "I mollusciii dei terreni ter- ziarii del Piemonte e della Liguria " were published in "Memorie della Reale Accademia delie Scienze di To- rino ". It has been seldom appreciated that reprints of these parts were actually published earlier than the par- ent volumes (table 1). The remainder of the series (parts 9, 10, 12, 13b, 14-30) was published independently by Carlo Clausen, Turin. In all assembled sets of the work that I have seen (including the reprint by Atlante Ma- lacologico, Rome, 1987-89), parts 1-8, 11 and 13a are represented by reprints. Although identical in content, the reprints differ from corresponding parts in the parent journal in being independently paginated and in having wrappers and title pages that were specially printed for each part. Determination of even the pagination of each part of the work was a laborious exercise, since there is considerable inconsistency in pagination from part to part in the parent journal, the reprints, the independentK- published parts, and in Sacco's (1904:vii) compilation (table 2). To avoid confusion with the companion series (see below), reprints, parent journal and independently published parts are referred to collectively as the "Me- morie/Clausen " series. Page reference to the Memorie/ Clausen series in the following discussions and in table 3 refer only to the reprints, though both paginations are indicated in table 2 and in the compilation of the work. The companion series (hereafter referred to as the "Bollettino " series) has the same title as the Memorie/ Clausen series and was issued serially in association with parts 7-29. It was published in "Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Universita di Torino ". The first two parts of the Bollettino series (Sacco 1890c,e) are subtitled "nota preventiva" (intro- ductory note), so the parts of this series were clearly intended to precede corresponding parts of the Memo- rie/Clausen series (see Sacco 1891a:l, footnote). Of the 22 issued parts of the Bollettino series, 17 were evidently published prior to the corresponding Memorie/Clausen parts, and 12 of these contain nomenclaturally valid in- troductions of new taxa (see below). There are few ref- erences to the Bollettino series in the Memorie/Clausen series (Sacco 1891a:l footnote, 1891d, synonymies). Sub- B. A. 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Marshall, 1991 Page 107 c rj -o a. o I > 61 o o o^ t. ^^ o> CO lO -r Ol ;c;r;-r-ro>Occ;D aJO-^'M;D'N o r- f- cc X' o r^a> — ocDooom ft-^ccooio Ol CO (N (M f— H r— I ^H ; C; « Tf lOa;CM(>J(MCJCJO)'N(M<>4C^>>>>>>>>>>>>>>>>>>>> a; >>>>>>>>>>>>_^ HHHHHf-HE-HHf-Hf-Hf-f-f-HHHH Hf-H(-(-H(-E-f-Ht-!- -r -r CO -T CO o ^ ;c ;o in ^~,— ^^-^--^^--^-H o — ^o ^- — a> — — -. — — - ■ — OlC0(M(MlO(M(NtCSO-H— .oC-HaOTOCOTOJCDIN ;i:.XS^-^-H;c — oCLO — I I I I I I I I I I I I T 01 CO C-] I- :C cc LO CDIO^— — I I I I I I o t^ > 00 fM J. Soco 30 — o a^ CI CI sc sc sc sc . 'ei D D 3 D aaa.D.Q.0. I g- M I I I I I I I I I I I I I I I Cl a^ -r ^ 0 r-r Cl c- COfMOoO|--lOt-C5^- C71 -TO>':£;ci~-rooooc I | CD in ^-^ on CI 0 TT CI ^^ CI re CI -r 00 r- in r- TO CD -T t n^ ci; j^ -T ^/^ n 00 CI cc CD CI m CD CO CO CO ■^ — c.-' c: LO CO re ^^ m re 1- in in (T in re 1^ 10 -T cr CI CJ CI C4 -T CI 00 CD 10 CD in in in in CO ( - X ^ 3 — ^cicoco^incDi^.^" m -c i~ jj o; o — ci CO ■^ in CD i~- oc 05 o — — — — -Hcicicqcicjcjoc^iNcico o a -a Mi a bJO '^ a bc -^ a 3 ro — -- CJ ''■" ' — (l^ 1" CO ^ rt CT> C3 3 ^ -T-. C 01 - !— -J Page 108 THE NAUTILUS, Vol. 105, No. 3 sequent references to the Bollettiiio series by other au- thors are rare, and the few that I fiave been able to trace {e.g., VVenz, 1938-44, Ferrero Mortara et al., 1984) refer only to parts 7 and 8 (Sacco 1890c, e). Moreover, the series has been overlooked in a compilation of his published works (Anonymous, 1948). While the Memorie/Clausen series was in progress, Sacco (1890a) also published "Catalogo paleontologico del bacino terziario de Piemonte". This is essentially a tabular listing of the 5,472 species of Tertiary fossils known from Piedmont. Although Sacco (1904:40, 74) dated his new subgenera Simplicotaiirasia and Miocenia from this work, the fact that other genus group names are potentially available there has been entirely over- looked. DATES OF PUBLICATION All reprints of parts 1-8, 11, and 13a of the Memorie/ Clausen series were clearly published before correspond- ing parts in the parent journal (table 1). Reprints of parts l-5[a-c] bear year dates on the title pages and wrappers and are secondarily dated in the text with a year, month and, with the e.xception of part 2, a day. With the exception of part 1, the imprinted year dates on the title page and in the text are accordant. Although it is possible that the textual dates actually represent dates of completion of the manuscripts, they are interpreted as dates of publication in the absence of evidence to the contrary (ICZN, 1985, Art. 21). Part 1 (Bellardi, 1873) is almost invariably dated from 1872 {e.g., Sacco, 1889:155, 1904:vii; Wenz, 1944:1513) in accordance with the date on the title page and wrap- per. The text (p. 264), however, is dated June 10 1873 and there are references to a publication dated 1873 in some synonymies {e.g., p. 130), so the 1872 date is clearly an error. Dates imprinted on the backs of the title pages of reprints of parts 6, 7, 8 and 1 1 are unequivocally dates of printing, as are dates on their parent volumes (table 1). Curiously, Sacco (1904:vii) dated part 6 from Feb- ruary 1890, part 8 from August 1890, and part 11 from February 1892, but these are surely errors since they predate the printing dates. That Sacco's dating of August 1890 for part 7 is almost certainly incorrect is suggested by the fact that the Natural History Museum, London (BMNH) copy of the reprint was accessioned on Septem- ber 1, 1890 yet all other parts were accessioned by that library 2-8 months later than the imprinted dates. The reprint of part 13a bears only the year of publi- cation (1893), although from the statement on the title page verso it must have been published after February 1893 and presumably before September 1893, the pub- lication date of part 13b In the absence of conflicting evidence, it is appropriate to accept Sacco's (1904:vii) indication that it was published in April 1893 (received BMNH August 14 1893). For bibliographic and priority purposes, the day of publication is interpreted as the last day of the stated month (ICZN, 1985, Art. 21i). Days of publication of reprints so interpreted (table 1) are shov\n in square parentheses. Year dates associated with new supraspecific taxa in the body of the text in both the Memorie/ (Clausen and Bollettino series frequently do not correspond with the actual year of publication and evidently represent merely anticipated years of publication. All parts of the Bollettino series bear imprinted dates (day, month and year), and library accession date stamps are accordant with their interpretation as dates of pub- lication (table 1). Sacco's "Catalogo paleontologico del bacino terziario del Piemonte " was submitted to the general assembly of the Societa Geologica Italiana on September 23 1889 for publication in Bollettino della Societa Geologica Italiana (BSGI — vol. 8(3), p. 545, 565). Presumably because of its length it was decided to split the work into two parts, which were published in BSGI volumes 8(3) and 9(2). The work also exists reprinted in its entirety as part 3 of Sacco's "II bacino terziario e quaternario del Piemonte" (1889-90). Despite extensive enquiries with numerous libraries, including the publishers and Societa Geologica Italiana, I have been unable to determine the exact dates of publication of the two forms of the Catalogo. References to Sacco's article in BSGI vol. 8(3) have been consistantly dated from 1889, but although parts 1-3 of this volume bear the date 1889 beside the volume number, part 3 is dated 1890 at the foot of the page (parts 1 and 2 are dated 1889). The latest date mentioned in the text of part 3 (p. 582) is in a report on an excursion that took place on September 26 1889, so it cannot have been published before then. Strong evidence that part 3 was published early in 1890 is provided by the list of new publications on the back wrapper (National Museum of New Zealand, Wellington) of series 3, volume 1, num- bers 3 and 4 ("Marzo e Aprile 1890") of the Bollettino del Reale Comitato Geologico d'ltalia (BRCGI) in which papers from BSGI volume 8(3) are dated 1890. The latest date shown in BRCGI 1(3,4) and thus the earliest date that it can be demonstrated to have been published (ICZN Art. 21c) is April 30 1890 (p. [159]) (interpretation ac- cordant with dates of publication of journals mentioned on pp. 151, 153, 156, 157). That BSGI 8(3) was probably published no later than early March 1890 is suggested by the fact that the BMNH cop> was received on May 3 1890, yet other Italian works of the period (table 1) were accessioned by this library 2 or more months after publication (see below). BSGI 9(2) is dated 1890 on the title page and was evidently published mid-late 1890. L'nfortunately, the earliest references to it that I have located all date from early 1891 and are thus of little assistance (received BMNH February 21). The reprint was printed from the same plates as the BSGI parts and is identical apart from being indepen- dently paginated and in that page 78 was split for division of the BSGI parts. It is dated 1889 on the title page and 1890 on the front v\ rapper (University Library, Stock- holm), and bears the statement (p. [ii]) "Estratto dal Bol- B. A. Marshall, 1991 Page 109 Table 3. List of molluscan supraspecific taxa introduced by Bellardi & Sacco (1873-1904) in "I molluschi dei terreni terziarii del Piemonte e dejja l.iguria, " and by Sacco (1890) in "Catalogo paleontologicn dei baciiio terziario del Piemonte ." Acrilloscala Sacco, 1890a:214(321) (1891b:[iv]; 1891d:86). Adiscoacrilla Sacco, 1890a:21-l(321) (1891b:[iii]; 1891d:67). Mectryom-lla Sacco, 1897b:100 (1897d;19). Alvaniclla Monterosato in Sacco, 189.5b;76 (189.5d;25). Amathinoides Sacco, 1896a:41. Ampullonatica Sacco, 1890a:208(315) (1890e:40; lS91a:10,5), Amussiopecten Sacco. 1897c:102 (1897e:53). Ancillarina Bellardi, 1882:217^ Ancillina Bellardi. 1882:220, Angulatumitrella Sacco, 1890b:36. Anura Bellardi. 1873:201. Archimcdiclla Sacco. 1895e:12. Arcuaiumitrclla Sacco, 1890b:37. Ascolatims Bellardi. 1884:41. Asperarca Sacco. 1898d:10. Aulachalia Sacco, ]893e:33. Bivoniopsis Sacco, 1896a:1.5, Brachelixclla Sacco, 1890a: 1 1.5(222) (1890b:34). Callistotapes Sacco, 1900:52. Cardiohicina Sacco. 1901a:89. Centrucardita Sacco. 1899b: 14. Ceromyclla Sacco. 1901a: 133. Clanculella Sacco, 1896b:22, Clinomitra Bellardi. 1888:10, Clinurella Sacco. 1890a:117(224) (1890b:44). Coccodentalittm Sacco. 18961:98 (1897a:110). Conocerilhium Sacco. 1895a:72 {nom.nud.}; 1895c:22. Contortia Sacco. 1894b:69 (1894d:48). Costatoscala Sacco. 1891d:72, Costoanachis Sacco. 1890b:57 Crassostrea Sacco, 1897b:99 (lS97d:15), Ciihitoslrea Sacco, 1897b:99 (1897d:12), Cyclodostomia Sacco. 1892a:53 {nom.nud.); 1892c:46. Cyllenina Bellardi. 1882161 CYLLENINAE Bellardi. 1882: 159 Cymbiilostrea Sacco. 1897d:12, Cyrsocrassiscala Sacco. 1891e:66. Cytherocardia Sacco. 1900:5, Denticuloglahella Sacco, 1890b:25. Dertonia Bellardi. 1884:51. Diptychomitra Bellardi, 1888: 10 DIPTYCHOMITRLNAE Bellardi, 1888: 10 Dtscosca/fl Sacco, 1890a:213(320) (1891b:[iii]; 189ld:.57), Ditoma Bellardi, 1877295 Dizoniopsis Sacco, 1895a:74 (1895c:67). Dolicholatims Bellardi, 1884:38, Echinophoria Sacco. 1890a:88(195) (1890c:9; 1890d:39). Ecostoanachis Sacco, 1890b:59, Eolatirus Bellardi. 1884:15, Eratotrivia Sacco, 1894c:62. Euspirocrommhtm Sacco. 1890a:208(315) (Sacco, 1890e:42; lS91b:[i]; 1891d:10). FimbriateUa .Sacco, 1895e:36. Flabellipecten Sacco, lS97c:102 (1897e:55). Flabellulum Bellardi. 1873:32, Flexopectcn Sacco. 1897c:102 (1897e:39). Fulguroficus Sacco, 1890a:97(204) (1890e:27; 1891a:41), Fusimorio Sacco, 18961:90 (1897a:25), Fusoficula Sacco, 1890a:97(204) (nom. nud.); 1890e:26 (1891a:38), Fusoterebra Sacco, 1891c:[vi] (1891e:59). Caleodinopsis Sacco, 189.5b:77 (189.5d:28). Caleudocassis Sacco, 1890a:85(192) (nom. nud.); 1890c:5 (1890d:18), G.\LEODOLIIDAE Sacco. 1890e:21 (invalid— no type ge- nus); 1891a:l, Caleodolium Sacco, 1891a:4 Galeodosconsia Sacco, 1890a:93(200) (1890c:17; 1890d:69), Camoplcura Bellardi, 1873:28, Genoa Bellardi. 1873:205. Gibbominliola Sacco. 1898a:41. Gibbori.s.soia Cossmann in Sacco, 189.5d:34. CMndoidca Sacco. 1896b:32 GdAndoidclla Sacco. 1896b:38 Gigantcotrochus Sacco. 1897a:4. Gigantostrea Sacco. 1897b:99 (1897d:14). Glabrondina Sacco, 1892c:49. Glabropecten Sacco, 1897e:39, 73 (error for Flexopectcn). Granosolarium Sacco, 1892b:56 (1892d:.59), HALIIDAE Sacco. 1893d:64 (1893e:32) (=Halliidae Sisinon- da. 1842), Hordeulima Sacco. 1892a:52 (1892c: 17). Jania Bellardi. 1873:147, Jousseaiimia Sacco. 1894c:8, Kleistopyrazus Sacco. 1895c:56 LACHESINAE Bellardi, 1877:150 Lazariella Sacco, 1899a:112 (1899b:21), Ledina Sacco. 1898c:107 (1898d:.53). Limatulella Sacco. 1898a: 16, Lissochlamis Sacco. 1897c:102 (1897e:46), Luponovula Sacco. 1894a:66 {nom. nud.); 1894c:44. Macomopsis Sacco. 1901a: 107 Macrochlamis Sacco, 1897c: 101 (1897e:32), Macrodostomia Sacco. 1892a:53 (nom. nud.); 1892c:43. Macrurella Sacco. 1890a:117(224) (1890b:46), Mantellina Sacco, 1904:148, MargoDohUa Sacco, 1890a:94(201) (nom. nud); 1890c:20; (1890d:82), MATHILDIDAE Sacco, 1892c:27 (=Mathildidae Dall, 1889), Mayeria Bellardi, 1873:156, Megacardita Sacco. 1899a:lll (1899b:9). Micromitra Bellardi. 1888:5, Miocardiella Sacco, 1904:161, Miocenia Sacco, 1890a: 100(207) (1904:40). Mioporomya Sacco, 1901a:140, Mitraefusm Bellardi, 1873:204. MITROLIMNIDAE Sacco, 1904:88. Modiohda Sacco. 1898a:41, Monodontclla Sacco. 1896b:23, Morionassa Sacco, 1890a:93(200) (nom. nud.); 1890c:18 (1890d:74). Myrsopsis Sacco, 1900:57. Myrteopsis Sacco, 1901a:96. Ncoathleta Sacco. 1890a:152(259) (1890b:12). Ncolatiru.s Bellardi, 1884:40. Nodosolarium Sacco. 1892d:63, Obliquarca Sacco. 1898c: 106 (1898d:16). Oombobna Sacco. 1896b: 15, Oopecten Sacco. 1897c:102 (1897e:54), Oostrombus Sacco, 1893d:63 (1893e:13). Ooterebralia Sacco. 1895c:51. OrmastraUum Sacco, 1896b:15. ORTHOMITRIXAE Bellardi. 1887a:3. lu THE NAUTILUS, Vol. 105, No. 3 Table 3. Continued Orilmrclk Sacco, 1890a:l 17(224) (If'aObi.Sl). Pantherinaria Sacco, 1894a:65 (1894c: 10,67). Papillicardhim Monterosato in Sacco, lS99a:11.3 (1899b;44) Parvamiissium Sacco, 1897c: 102 (1897e:48). Parvivcnus .Sacco. 1900 4.5 Parvochlamys Sacco, 1904: 145. Pectinatarca Sacco, 1898c: 106 (1898d:26). Phornilellus Sacco, 1896b:35. Pilhocerithium Sacco, 1895a:72 (1895c:28). Plrsiolatirus Bcllardi. 1884:32. I'LESIOMITHINAE Bellardi, 1887b:23. Plicostoniia Montero.sato in Sacco, 1892c:46. Poculina Bcllardi. 1873:33 Pohlgireulima Sacco, 1892c:10. PwaJusta Sacco, 1894a:66 (1894c:33). Pseudavena Sacco, 1896f:92 (nom. mid.), 1897a:39. Pseudemarginula Sacco, 1897a:17. Pseitdolalims Bellardi, 1884:46. Pscudonina Sacco, 1896b:9. PaeudonineHa Sacco, 1896b:51. Pscudosthenorytis Sacco, 1891d:43. Pscudotorinia Sacco, 1892d:66. Pseiidoxyperas Sacco, 1901a:26. Ptycheulimella Sacco, 1892a:53 (1892c:59). Ptychocerithium Sacco, 1895a:72 (1895c:23). Ptychomdania Sacco, 1895b:75 (1895d:7). Ptychopotamides Sacco. 1895a:73 (1895c:44). PurpurcUa Bellardi. 1882:193. PI RPl RELLINAE Bellardi, 1882:193. HLRPURINAE Bellardi, 1882:174 ( = Piirpurinae Lamarck, 1809). Pyrogolampros Sacco, 1892b:54 (nam. nud.)\ 1892c:85. Rliombomya Sacco, 190 la: 133. Ringiculella Sacco, 1892b:55 (1892d: 16). Ringiculocosta Sacco, 1892b:56 (1892d:33). Ringiculospongia Sacco, 1892b:55 (1892d:15). Ruuaidtia Bellardi, 1877:223. Sainliopsis Sacco, 1898a:ll. Sassia Bellardi, 1873:219. Scalnella Sacco, 1890a: 118(225) (nom. nud.}; 1890b:54. Scalaricardita Sacco, 1899b:23 Scaptorrhynchus Bellardi, 1873:12. Scgucnziclla Sacco, 1904:123. Sigaretotrcma Sacco, 1890a:207(314) (1890e:38; 1891a:97), Sinipliiodnlinin Sacco, 1891a:13. Simpli,ogl(d>ella Sacco, 1890b;21. Simplirolanmsia Sacco, 1890a:124(231 ) (1904:74). Spica Monterosato in Sacco, 1892a:53 (nom. nud.}; 1892c: 63. Spineoterelna Sacco. I891c:[vi] (1891e:58). Siazzania Sacco, 1890a: 138(245) (1890b:26). Steirotuha Sacco, 189.5e:40. Strigosella Sacco, 1896b:49. Striolucina Sacco, 1901a:83. Striolcrclnum Sacco, 1891c:[vi] (1891e:33). Striolurlnmilla Sacco, 1892b:.55 (1892c:94). Sulcoghidius Sacco, 1893d:63 (1893e:15). Sulconuirinula Sacco, 1896f:95 (nom. nud.h !897a:80. Sulcosulndaria Sacco, 1892a:52 (nom. nud.); 1892c:17. Sulcoturhomlla Sacco, 1892b:55 (1892c:92). Syndesmyella Sacco, 1901a:122. Tflnra.sifl Bellardi, 1882:194. TAURASINAE Sacco, 1904:74. Tauraxinus .Sacco, 1901a:61. TauTcmarginula Sacco, 1897a:13. Taurofuri.s Sacco, 1895c:62. Taurotapes Sacco, 1900:58. Tecionatica Sacco, 1890a:205(312) (1890e:33; 1891a:81). TEREBELLIDAE Sacco, 1893d:64. (=Terebellinae H.&.A. Adams, 1854). Tetrastomella Sacco, 1890a: 11 6(223) (1890b:41). Timirclla Sacco, 1890a:l 17(224) (I890b:52). Thiarinclla Sacco, 1890a: 118(225) (1890b:56). Tiamccrithium Sacco, 1895a:72 (189.5c:35). Tiarapirenella Sacco, 1895a:73 (189.5c:60). Tomochiton Sacco, 1877a:90 (nom. nud.). Torculoidella Sacco, 1895e:28. TORNIDAE Sacco, 1896b:55. Trochacerilhium Cossmann & Sacco in Sacco, !896t):60. Trocholugurium Sacco, 1896a:27. Tuhcrculodolium Sacco, 1891a:9. Turritodoslomia Sacco, 1892a:53 (1892c:41). Tylastmlium Sacco. 1896b:19. Uromitra Bellardi, 1887b:23. Variamussium Sacco, 1897c: 102 (1897e:49). Ventricoloidea Sacco, 1900:31. Vulgocerithium Sacco, 1895c:7. lettino della Societa geologica italiana Vol. 8(3)". This strongly suggests that the reprint was printed when it was still assumed that the work was to be published entire in BSGI 8(3) and thus before that part was published. The date "1889" on the title page could indicate that it was printed late in that year and that the wrapper was dated 1890 when it was realised that the work would not be circulated until 1890, or that BSGI 8(3) was to be published in 1890. It seems more plausible, however, that the reprint was printed in 1890, and that the 1889 date was added to the manuscript by Sacco and/or simply represents the nominal year for BSGI volume 8. The earliest reference to the work (Sacco, 1890d:7, footnote — "Boll. See. geol. it., 1889-90") appears in an introduction dated February 5 1890. Since the work cannot be proven to have been published in 1889, in the absence of evi- dence to the contrarv the date of publication is inter- preted as February ,5' 1890 (ICZN, 1985, .Art. 21c). This conclusion is accordant w ith C. D. Sherborn's, who noted "issued before the Bollettino" on the title page of the BMNH copy of the reprint. STATUS OF TAXA Apart from supraspecific names (see below), parts 7 and 8 of the Bollettino series (Sacco 1890c,e) contain brief Latin diagnoses of hundreds of species group taxa ac- companied b\- specimen dimensions and type localities. B. A. Marshall, 1991 Page 1 1 1 Although these descriptions are brief and unaccompa- nied by illustrations, many meet criteria for availability (ICZN, 1985: Art. 11, 12), so that approximately half the total number of species group taxa more fulK described and illustrated in the corresponding parts of the Me- morie/Clausen series actually date from the prior Bol- lettino parts (taxa in part 8 so dated by Ferrero Mortara et al., 1984). The remainder of the species group names first introduced in parts 7 and 8 of the Bollettino series and all of those in subsequent parts are nomiija niida. since they are mentioned by name onl\ . Most supraspecific names first introduced in the Bol- lettino series meet criteria for availability since they are accompanied by one or more valid species names that were introduced by earlier authors. Some of the generic and subgeneric names, however, are nomina nitda. since they are accompanied onh' by specific names (i.e., Sac- co's) that are themselves nomina nuda. Two subfamily names (Orthomitrinae and Plesiomitrinae) introduced by Bellardi ( 1887a, b) are unavailable since they are not based on a type genus, while Galeodoliidae is unavailable at its first introduction (Sacco, 1890e:21) because it is un- accompanied by a type genus (Galeodolium Sacco, 1891a: 4). Genus group names first introduced by Sacco (1890c,e, 1891b) in parts 7-9 of the Bollettino series lack type species designations, while type species are designated unequivocally in all of the following parts. Type species for man\- genus group names introduced by prior authors are indicated in parts 17-29 of the Bollettino series and are thus potentially available as first subsequent desig- nations. All moUuscan species group names introduced by Sac- co in "Catalogo paleontologico del bacino terziario del Piemonte" are nomina nuda. Of the 26 genus group names first introduced there, however, 21 meet criteria for availability because they are accompanied by valid specific names that were introduced in prior publications (ICZN, 1985: Art. 12b(5)). The only species group name that is validly introduced in this work (reprint p. 192) is Mastodon arvernensis var. cantamessae Sacco, 1890 (Mammalia). Attention is drawn to the fact that dating of genus group taxa from the Bollettino series and the "Catalogo" affects some currently accepted type species designa- tions. For example, in cases where Sacco did not desig- nate a type species in the Memorie/Clausen series, au- thors have selected type species by subsequent designation, but some of these designations are invalid because the species were not among those included when the genus was first introduced. These problems should preferably be treated individually by specialists, and to avoid influencing future decisions they are not consid- ered here. All molluscan supraspecific names introduced by Bel- lardi (1873-1888) and Sacco (1890-1904) that I have been able to trace are listed (table 3). For convenience, page references are provided for all names in the Me- morie/Clausen series (reprints) and in "Catalogo pa- leontologico del bacino terziario del Piemonte , but onl\ to prior introductions (valid and invalid) in the Bollettino series. Page references to the "Catalogo" include both the reprint and (in parentheses) Bollettino della Societa Geologica Italiana 9(2). Unless indicated as nomen nu- dum or otherwise invalid, the first citation is interpreted as the earliest nomenclaturally valid introduction of the name. Spelling of Lissochlamis, M.acrochl.\mis and v.\riamvssium In part 24 of the Memorie/Clausen series, Sacco (1897e) altered the original spellings (Sacco, 1897c) of Lissochla- mis, Macrochlamis, and Variamussum to Lissochlamys, Macrochlamys and Variamussium. Although "chlamis" is an incorrect transliteration of the Greek "chlamys" (a garment), incorrect transliterations are not to be consid- ered inadvertent errors (ICZN, 1985: Art. 32c), so Lis- sochlamys and Macrochlamys are unjustified emenda- tions (ICZN, 1985: Art. 33). That Sacco's (1897c) original spellings of Lissochlamis and Macrochlamis were per- haps intentional is suggested by the consistent usage (twice each), as subgenera of Chlamys (sic. — three times). Ac- cordingly, I follow Hertlein (1969) and Yokes (1980) and interpret Lissochlamis and Macrochlamis as correct spellings, despite the fact that Lissochlamys and Macro- chlamys have been widely used. The original spelling Variamussum has been entirely overlooked, and Sacco's subsequent spelling has been universally accepted. Although Variamussum is consis- tently so-spelt (twice) in the original reference, it is ob- viously a lapsus, so from the standpoint of nomenclatural stability it is appropriate in this instance to accept Varia- mussium as a justified emendation. COMPILATION OF "I MOLLUSCHI DEI TERBENI TERZIARII DEL PIEMONTE E DELLA LIGURIA", AND "CATALOGO PALEONTOLOGICO DEL BACINO TERZIARIO DEL PIEMONTE" It should be noted that the sections entitled "Catalogo generale dei molluschi dei terreni terziarii del Piemonte e della Liguria" occurring in each part of the Memorie/ Clausen series {i.e.. in both parent journal and reprints) were reprinted independently with independent pagi- nations and title pages. Identical to the originals, these reprints were prepared for separate binding and are not included here. Bellardi, L. 1873. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 1: Cephalopoda, Pteropoda, Heteropoda, Gasteropoda (Muricidae e Tritonidae). Memorie della Reale Accademia delle Scienze di Torino (2) 27, 1873:33-294 (reprint 264 p.), 15 plates (June 10 1873). Bellardi, L. 1877. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 2: Gasteropoda (Pleurotomidae). Memorie della Reale Accademia Page 112 THE NAUTILUS. Vol. 105, No. 3 delle Scienze di Torino (2) 29, 1878:1-364 (reprint 364 p.), 9 plates (June 30 1877). Bellardi, L. 1882. I nioUiischi dei terreiii terziarii del Piemonte e della liguria. Parte 3: Gasteropoda (Buc- cinidae, Cyclopsidae, Purpuridae, Corailiophilidae, Olividae). Memorie della Reale Accademia delle Scienze di Torino (2) 34, 1883:219-469 (reprint 253 p.), 12 plates (December 10 1882). Bellardi, L. 1884. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 4: Fasciolariidae e Turbinellidae. Memorie della Reale Accademia delle Scienze di Torino (2) 37, 1886:3-62 (reprint 62 p.), 2 plates (December 1 1884). Bellardi, L. 1887a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 5[a]: Mitridae. Me- morie della Reale Accademia delle Scienze di To- rino (2) 38, 1888:79-166 (reprint 85 p.), 2 plates (January 15 1887). Bellardi, L. 1887b. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 5[b]: Mitridae (con- tinuazione). Memorie della Reale Accademia delle Scienze di Torino (2) 38, 1888:257-327 (reprint 72 p.), 2 plates (June 1 1887). Bellardi, L. 1888. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 5[c]: Mitridae (fine). Memorie della Reale Accademia delle Scienze di Torino (2) 39, 1889:145-194 (reprint 52 p.), 2 plates (September 20 1888). Sacco, F. 1890a. Catalogo paleontologico del bacino terziario del Piemonte. Reale Accademia dei Lincei, Roma, 233 p. (reprint). Subsequently published in Bollettino della Societa Geologica Italiana 8(3):281- 356, 9(2):185-340 (February 5 1890). Sacco, F. 1890b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 6: (Volutidae, Mar- ginellidae, Columbellidae). Memorie della Reale Accademia delle Scienze di Torino (2) 40, 1890: 295-368 (reprint 76 p.), 2 plates (April 6 1890). Sacco, F. 1890c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 7: (Harpidae e Cas- sididae). Bollettino dei Musei di Zoologia ed Ana- tomia comparata della Reale Universita di Torino 5(82): 1-20 (June 13 1890). Sacco, F. 1890d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 7: (Harpidae e Cas- sididae). Memorie della Reale Accademia delle Scienze di Torino (2) 40, 1890:469-560 (reprint 96 p.), 2 plates (June 18 1890). Sacco, F. 1890e. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 8: Galeodoliidae, Doliidae, Ficulidaee Naticidae. Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di Torino 5(86):21-43 (August 12 1890). Sacco, F. 1891a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 8: Galeodoliidae, Doliidae, Ficulidae e Naticidae. Memorie della Reale Accademia delle Scienze di Torino (2) 41, 1891:225- 338 (reprint 114 p.), 2 plates (March 25 1891). Sacco, F. 1891b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 9: Naticidae (fine), Scalariidae ed Aclidae. Bollettino dei Musei di Zoo- gia ed Anatomia comparata della Reale Universita di Torino 6(103):[i-iv] (May 29 1891). Sacco, F. 1891c. [I molluschi dei terreni terziarii del Piemonte e della Liguria] Parte 10: Cassididae (ag- giunte), Terebridae e Pusionellidae. Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di Torino 6 (103):[v-vi] (published consecutively with Part 9) (May 29 1891). Sacco, F. 1891d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 9: (Naticidae (fine), Scalariidae ed Aclidae). Clausen, Torino, 103 p., 2 plates (May 31 1891). Sacco, F. 1891e. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 10: (Cassididae (ag- giunte), Terebridae e Pusionellidae). Clausen, To- rino, 68 p., 2 plates (September 30 1892). Sacco, F. 1892a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 11: Eulimidae e Pyramidellidae (pars). Bollettino dei Musei di Zoolo- gia ed Anatomia comparata della Reale Universita di Torino 7(121):51-53 (April 30 1892). Sacco, F. 1892b. [I molluschi dei terreni terziarii del Piemonte e della Liguria]. Parte 12: (Pyramidellidae (fine), Ringiculidae, Solariidae e Scalariidae (agg.)). Bollettino dei Musei di Zoologia ed Anatomia com- parata della Reale Universita di Torino 7(121 );54- 57 (Published consecutively with Part 11) (April 30 1892). Sacco, F. 1892c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 11: Eulimidae e Pyramidellidae (parte). Memorie della Reale Ac- cademia delle Scienze di Torino (2) 42, 1892:585- 682 (reprint 100 p.), 2 plates (June 30 1892). Sacco, F. 1892d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 12: (Pyramidellidae (fine), Ringiculidae, Solariidae e Scalariidae (ag- giunte)). Clausen, Torino, 88 p., 2 plates (June 30 1892). Sacco, F. 1893a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 13[a]: (Conidae) (Fasciolo Primo). Memorie della Reale Accademia delle Scienze di Torino (2) 44, 1894:1-54 (reprint 54 p.), 2 plates (April 29 1893). Sacco, F. 1893b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 13: (Conidae e Co- norbidae). Bollettino dei Musei di Zoologia ed An- atomia comparata della Reale Universita di Torino 8(153): 1-4 [NB— this part corresponds to 1893a and 1893c combined] (May 31 1893). Sacco, F. 1893c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 13[b]: (Conidae e Conorbidae). (Fascicolo secundo). Clausen, Torino: 55-143, 9 plates (September 30 1893). Sacco, F. 1893d. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 14: (Strombidae, Terebellidae, Chenopidae, Haliidae, Cypreidae). Bollettino dei Musei di Zoologia ed Anatomia com- B. A. Marshall, 1991 Page 113 parata della Reale Universita di Torino 8(165):63- 64 (December 11 1893). Sacco, F. 1893e. I molluschi del terreni terziarii del Piemonte e della Liguria. Parte 14: (Strombidae, Terebellidae, Chenopidae ed Haliidae). Clausen, Torino, 40 p., 2 plates (December 30 1893). Sacco, F. 1894a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 15: Fam. C\praei- dae Gray 1824. Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di To- rino 9(171):65-67 (April 27 1894). Sacco, F. 1894b. [I molluschi dei terreni terziarii del Piemonte e della Liguria] Parte 16: Fam. Cancel- lariidae H. e A. Adams 1853. Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di Torino 9(171 ):68-70 (published con- secutively with Part 15) (April 27 1894). Sacco, F. 1894c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 15: (Cypraeidae, ed Amphiperasidae). Clausen, Torino, 74 p , 3 plates (April 29 1894). Sacco, F. 1894d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 16: (Cancellariidae). Clausen, Torino, 82 p., 3 plates (August 31 1894). Sacco, F. 1895a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 17: (Cerithiidae, Triforidae, Cerithiopsidae e Diastomidae). Bollet- tino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di Torino 10(197):71-74 (February 25 1895). Sacco, F. 1895b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 18: (Melaniidae, Littorinidae, Fossaridae, Rissoidae, Hydrobiidae, Paludinidae e Valvatidae). Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale L^ni- versita di Torino 10(206):75-78 (May 30 1895). Sacco, F. 1895c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 17: (Cerithiidae, Triforidae, Cerithiopsidae e Diastomidae). Clausen, Torino, 86 p., 3 plates (June 30 1895). Sacco, F. 1895d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 18; (Melaniidae, Littorinidae, Fossaridae, Rissoidae, Hydrobiidae, Paludinidae e Valvatidae). Clausen, Torino, 52 p., 1 plate (September 30 1895). Sacco, F. 1895e. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 19: (Turritellidae e Mathildidae). Clausen, Torino, 46 p., 3 plates (De- cember 30 1895). Sacco, F. 1896a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 20: (Caecidae, Ver- metidae, Siliquariidae, Phoridae, Calyptraeidae, Capulidae, Hipponycidae, Neritidae, Neritopsidae). Clausen, Torino, 65 p., 5 plates (May 31 1896). Sacco, F. 1896b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 21: (Naricidae, Modulidae, Phasianellidae, Turbinidae, Delphinu- lidae, Cyclostrematidae, Tornidae). Clausen, Tori- no, 64 p., 4 plates [NB — p. 64 incorrectly numbered •'65"] (September 30 1896). Sacco, F. 1896c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 19; (Turritellidae e Mathildidae). Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di To- rino 11(267):79-81 (December 14 1896). Sacco, F. 1896d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 20; (Caecidae, Ver- metidae, Phoridae, Calyptraeidae, Capulidae, Hip- ponycidae e Neritidae). Bollettino dei Musei di Zoo- logia ed Anatomia comparata della Reale Universita di Torino ll(267):82-84 (published consecutively with Parts 19, 21, 22) (December 14 1896). Sacco, F. 1896e. [I molluschi dei terreni terziarii del Piemonte e della Liguria] Parte 21: (Naricidae, Modulidae, Phasianellidae, Turbinidae, Delphinu- lidae, Cyclostrematidae e Tornidae). Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di Torino ll(267):85-88 (pub- lished consecutiveh with Parts 19, 20, 22) (Decem- ber 14 1896). Sacco, F. 1896f. [1 molluschi dei terreni terziarii del Piemonte e della Liguria] Parte 22: [untitled]. Bol- lettino dei Musei di Zoologia ed Anatomia compara- ta della Reale Universita di Torino ll(267):89-98 (published consecutively with Parts 19-21) (Decem- ber 14 1896). Sacco, F. 1897a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 22: Gasteropoda (fine) (Pleurotomariidae, Scissurellidae, Haliotidae, Fissurellidae, Tecturidae, Patellidae, Oocorythidae, Cyclophoridae, Cyclostomidae, Aciculidae, Trun- catellidae; Acteonidae, Tornatinidae, Scaphandri- dae, Bullidae, Cylicnidae, Philinidae, Umbrelli- dae). — Pulmonata (Testacellidae, Limacidae, Vitrinidae, Helicidae, Pupidae, Stenogyridae, Suc- cineidae; Auriculidae; Limnaeidae, Physidae; Si- phonariidae). Amphineura (Chitonidae). — Scapho- poda (Dentaliidae). Clausen, Torino, 148 p., 10 plates (March 31 1897). Sacco, F. 1897b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 23: (Ostreidae, Ano- miidae e Dimyidae). Bollettino dei Musei di Zoolo- gia ed Anatomia comparata della Reale Universita di Torino 12(298);99-100 (June 11 1897). Sacco, F. 1897c. [I molluschi dei terreni terziarii del Piemonte e della Liguria] Parte 24; [untitled]. Bol- lettino dei Musei di Zoologia ed .\natomia compara- ta della Reale Universita di Torino 12(298); 101-102 (published consecutivelv with Part 23) (June 11 1897). Sacco, F. 1897d. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 23: Pelecypoda (Os- treidae, Anomiidae e Dimyidae). Clausen, Torino, 66 p., 11 plates (June 30 1897). Sacco, F. 1897e. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 24; (Pectinidae). Page 114 THE NAUTILUS, Vol. 105. No. 3 Clausen, Torino, 116 p., 21 plates (December 30 1897). Sacco, F. 1898a. 1 molluschi del terreni terziarii del Piemonte e della Liguria. Parte 25: (Spondylidae, Radulidae, .\viciilidae, Vulsellidae, Pernidae, Pin- nidae, Mytilidae, Dreissensiidae). Clausen, Torino, 76 p., 12 plates (August 31 1898). Sacco, F. 1898b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 25: (Spondylidae, Radulidae, Aviculidae, Vulsellidae, Pernidae, Pin- nidae, Mytilidae e Dreissensiidae). Boilettino dei Musei di Zoologia ed Anatomia comparata della Reale Universita di Torino 13(332): 103-105 (De- cember 14 1898). Sacco, F. 1898c. [I molluschi dei terreni terziarii del Piemonte e della Liguria] Parte 26: (Arcidae, Pec- tunculidae, Limopsidae, Nuculidae, Ledidae e Mal- letidae). Boilettino dei Musei di Zoologia ed Ana- tomia comparata della Reale Universita di Torino 13(332):106-108 (published consecutively with Part 25) (December 14 1898). Sacco, F. 1898d. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 26: (Arcidae, Pec- tunculidae, Limopsidae, Nuculidae, Ledidae, Mal- letidae). Clausen, Torino, 92 p., 12 plates (December 30 1898). Sacco, F. 1899a. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 27: (Unionidae, Car- ditidae, Astartidae, Crassatellidae, Lasaeidae, Gale- ommidae, Cardiidae, Limnocardiidae e Chamidae): Boilettino dei Musei di Zoologia ed Anatomia com- parata della Reale Universita di Torino 14(349):111- 114 (May 23 1899). Sacco, F. 1899b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 27: (Unionidae, Car- ditidae, Astartidae, Crassatellidae, Lasaeidae, Gale- ommidae, Cardiidae, Limnocardiidae, Chamidae). Clausen, Torino, 102 p., 14 plates (September 30 1899). Sacco, F. 1900. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 28: (Isocardiidae, Cyprinidae, Veneridae, Petricolidiae, Cyrenidae, Sphaeridae). Clausen, Torino, 98 p., 14 plates (April 29 1900). Sacco, F. 1901a. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 29: (Donacidae, Psammobiidae, Solenidae, Mesodesmidae, Mactri- dae, Cardiidae, Myidae, Corbulidae, Glycymeridae, Gastrochaenidae, Pholadidae, Teredidae, Crypto- dontidae, Ungulidae, Lucinidae, Tellinidae, Scrobi- culariidae, Cuspidariidae, Solenomyidae, Pandori- dae, Verticordiidae, Lyonsiidae, Ceromyidae, Arcomyidae, Anatinidae, Poromyidae, Clavagelli- dae). Clausen. Torino, 216 p., 29 plates (June 30 1901). Sacco, F. 1901b. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 28: [untitled]. Boi- lettino dei Musei di Zoologia ed Anatomia compara- ta della Reale Universita di Torino 16(409):115-118 (November 25 1901). Sacco, F. 1901c. I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 29: [untitled]. Boi- lettino dei Musei di Zoologia ed Anatomia compara- ta della Reale Universita di Torino 16(409):1 18-125 (November 25 1901). Sacco, F. 1904. 1 molluschi dei terreni terziarii del Piemonte e della Liguria. Parte 30: Aggiunte e cor- rezioni. Considerazioni generali. Indice generale dell' opera. Clausen, Torino, 203 -I- xxxvi p., 31 plates (August 31 1904), ACKNOWLEDGEMENTS First and foremost I wish to express my extreme gratitude to Mr R. J. Cleevely (Department of Palaeontology, The Natural History Museum, London), who patiently at- tended to many requests for library searches and infor- mation, and without whose insightful assistance and ap- preciation of esoteric bibliographic problems this work would not have been possible. A great deal of additional information was provided by Dr R. Giannuzzi-Savelli (Palermo), Dr M. G. Harasewych and Dr A. Kabat (Na- tional Museum of Natural History, Washington DC), and Dr A. Waren (Swedish Museum of Natural History, Stockholm). Of the numerous other individuals who re- plied to my enquiries, the following provided substantial information and assistance: Dr P. Bouchet (Museum Na- tional d'Histoire Naturelle, Paris), Dr A. L. Cocuzzi (Ac- cademia Nazionale dei Lincei, Roma), Dr E. Menesini (Dipartimento di Scienze della Terra, Pisa), Dr C. Moran- dini (Museo Friulano di Storia Naturale, Udine), Dr S. Palazzi (Modena), Dr G. Pavia (Dipartimento di Scienze della Terra, Torino), Dr M. Taviani (Istituto di Geologia Marina, Bologna), and Dr A. Zuccari (Societa Geologica Italiana, Roma). For constructive comments on the manuscript I am grateful to Dr A. G. Beu, Dr P. Bouchet, Mr R. J. Cleeve- ly, Dr R. Gianuzzi-Savelli, Dr M. G. Harasewych. Dr R. S. Houbrick, Dr A. Kabat, Dr G. Pavia, Mr R. E. Petit, Dr W. F. Ponder and Dr A. Waren. Special thanks to Kathleen Ryan for cheerful expertise with word pro- cessing, and Manuela Angelo for translations. LITERATURE CITED Anonymous. 1948. Elencho delle publicazioni di Federico Sacco, dottore in scienze, gia professore di geologia al politecnico e di paleontologia all Unixersita di Torino. Checchini, Torino 29 p. Ferrero Mortara, E., L. Montefameglio, G. Pavia, and R Tam- pieri. 1982. Catalogo dei tipi e degli esamplari figurati della collezione Bellardi e Sacco. Part 1. Musei Regionale di Scienze Naturali, Torino. Cataloghi 6: 1-327, 58 pis. Ferrero Mortara, E., L. Montefameglio, M. Novelli, G. Opesso, G. Pavia, and R. Tampieri 1984, Catalogo dei tipi e degli esamplari figurati della collezione Bellardi e Sacco. B. A. Marshall, 1991 Page 115 Part 2. Musei Regionale di Scienze Natural!, Torino, Ca- taloghi 7:1-484, 56 pis. Hertlein, L. G. 1969. Pectinidae. 7n. Moore, R. C. (ed.). Trea- tise on invertebrate paleontology, N(l) Mollusca, 6 Bival- via. Geological Society of America, Boulder, Colorado and University of Kansas, Lawrence, p. 348-373, ICZN. 1985. International code of zoological nomenclature, 3rd ed. International Trust for Zoological Nomenclature and British Museum (Natural History), London, 338 p. Voices, H E. 1980. Genera of the Bivalvia: a systematic and bibiographic catalogue (revised and updated). Paleonto- logical Research Institution, Ithaca, New York. 307 p. Wenz, W. 1938-44. Gastropoda. 1: Allgemeiner Teil und Prosobranchia. In: Schindewolf, O. H. (ed.). Handbuch der Palaozoologie 6, Borntraeger, Berlin, xii + 1639 + 10 p. THE NAUTILUS 105(3):116-117, 1991 Page 116 A Note on Supposed Homonyms of Octopus australis Hoyle, 1885, with Comments on Octopus caniphelli Smith, 1902 (Cephalopoda: Octopodinae) Ronald B. Toll Doparlment of Biology The University of tlie South Sewanee, TN 37375, USA ABSTRACT "Octopus australis (Massy, 1916)" and "O. australis (Benham, 1942)'", both from New Zealand, are determined not to be homonyms of O. australis Hoyle, 1885, from Australia, as pre- viously suggested. Examination of new material attributable to Octopus camphclli Smith, 1902, from New Zealand, supports the ta.xonomic distinction between O. camphclli and O. aus- tralis. Massy 's, Benham's and Dell's material is probabK refer- able to O. campelli. Key words: Octopus australis; Octopus cainpbclli: Octopod- Tait (1982) redescribed and reviewed the taxonomic po- sition of Octopus australis Hoyle, 1885, originally de- scribed from Port Jackson, New South Wales, Australia. In his review of related taxa "described from New Zea- land, Tait (1982:20) indicated that two nominal taxa, "Polypus australis Massy [not Massey], 1916" and "Rob- sonella australis Benham, 1942, " are junior homonyms of Octopus australis Hoyle, following the placement of Polypus and Robsonella into the synonymy of Octopus s. s. by Robson (1929) and Pickford (1955), respectively. Tait (1982:20) concluded that these two species require renaming. Massy s (1916) account, however, clearly in- dicated that she did not describe Polypus australis as a new species, but actually attributed her specimens to Hoyle s australis, and indeed cited Hoyle as the author. Therefore, Polypus australis is simply a new combina- tion proposed by Massy, with Hoyle retained as the au- thor of the species [see ICZN, 1985: Art. 50(c)(ii)]. The same is true for Robsonella australis, a new combination, not a new species, proposed by Benham (1942) based on his examination of new material. Tait (1982:19) suggested, and I agree, that based on characters of the liectocotylus, it is highly unlikely that Octopus australis from Australia and (). campbelli Smith, 1902, from C;am|)bell Island, New Zealand are conspe- cific, as proposed by Robson (1929:145). Tait further stated (pg. 20) that ""all the New Zealand species pre- viously considered to be synonyms of O. australis Hoyle appear to be separate and distinct". The available data, however, do not support this assertion. The type of O. campbelli and the specimens attributed to Ho> le's aus- tralis by Massy (1916), Benham (1942), and Dell (1952) are all characterized by W-shaped funnel organs. Robson (1929:190) gave the value 8.5 as the ligula length index (LLI) of the type of O. campbelli. Benham's specimens, remeasured by Tait, have a LLI of 6 to 10. Massy s specimen has a LLI of 10.9; however, the proximal start- ing point along the hectocotylus used to obtain this mea- surement is unclear. Dell's specimens have a LLI of 5- 7. These overlapping data sets certainly cannot be used to distinguish among taxa. In comparison, Tait (1982) reported the LLI ior O. australis as 12-18. Massy 's (1916) figure of the radula showed a symmetrical, pentacuspid rachidian. Dell (1952) reported that the rachidian tooth is asymmetrical (however, he gave the formula as ""A4", probably an error due to the confusing situation created by Robson s original designation of the letter A for sym- metrical rachidian teeth, see Robson, 1925). This varia- tion of the radula is within the range observed from new- material of O. campbelli examined here (see below). Therefore, based on these accounts there appears to be no clear taxonomic separation among New Zealand spec- imens. For comparison to published accounts, I examined six specimens of Octopus campbelli (4 males, ML 22-30 mm, all with spermatophores; 2 mature females, ML 25- 27 mm) from Portobello, New Zealand, deposited in the collections of the Invertebrate Museum, Rosenstiel School of Marine and Atmospheric Sciences, University of Mi- ami, Florida (UMML 31.2447). The range of LLI is 6.5- 11.1 (mean = 8.48). Two inconsistencies exist between the original description of the type of O. campbelli and the specimens I have examined. The Portobello speci- mens have gill lamellae counts of 7-8 (including terminal lamella). Smith (1902) gave no data on the gills from the holotype of (). campbelli. Robson (1929:190) stated that the gill count of the holotype is ""probably ten", based on his reexamination. Robson's 1929 monograph contains certain factual inconsistencies (Voss, 1973). Indeed, Tait R. B. Toll, 1991 Page 117 (1982:20) found that Robson, despite his assertion that he had examined the radula from one of the syntypes of O. australis, could not have done so because the buccal mass had not been removed from either specimen at the time Tait acquired them for examination. Therefore, Robson s gill lamellae data, given with some degree of uncertainty, should be discounted. The second discrepancy concerns the greatly enlarged and elevated suckers (seventh pair only) on the lateral arm pairs of the type of O. camphcUi (SnI ~ 8.5; Sel ~ 14.3). Three Portobello male specimens have moderately enlarged suckers (SnI 8.2-10.4; Sel 10.7-12.2) which are not distinctly elevated. In these specimens about three pairs of suckers, beginning with the fifth pair, are en- larged on the lateral arm pairs. Specially enlarged suckers are not mentioned specifically in the accounts of Massy, Benham, or Dell. While this disparity is curious, it could be the result of differences in reproductive maturity. In light of the other numerous similarities in morpholog\ , dermal texture, and absolute size, I do not consider this difference to be substantiative. The radulae of three of the Portobello specimens were examined and showed considerable variation. The ra- chidian tooth is strongK as\ mmetrical in one specimen, weakK as\ mmetrical in the second, and varies from s\ m- metrical in one region to asymmetrical along another region in the third example Intraspecific (including in- tra-radular) variation is knov\n to occur in Octopoda (Robson, 1925, 1929; KAam, 1933, 1941; Voss, 1973; Toll, 1981, unpublished); therefore, differences in the sym- metry of the radula can be unreliable in distinguishing among related species. Based on examination of all pertinent accounts, I find that no homonyms of Octopus australis Hoyle, 1885 exist; therefore, no renaming needs to be take place. Octopus australis Hoyle and Octopus carnpbclli Smith, 1902 appear to be distinct allopatric taxa, most easiK differentiated on the basis of hectocotylus and penial apparatus morphology, dermal texture and absolute size. Furthermore, the specimens described by Mass\. Ben- ham and Dell from New Zealand are consistent with the characters of O. campbelli and I consider them to be referable to it. ACKNOWLEDGEMENTS Nancy A. Voss, University of Miami, kindly loaned the specimens of Octopus campbelli. Clyde F. E. Roper and Michael J. Sweene\- made constructive comments on an earlier version of the manuscript. This contribution was supported, in part, by a grant from the National Science Foundation (BSR 8508439) and general support for re- search from The I'niversity of the South. LITERATURE CITED .Adam, VV, 19.33. Notes sur le.s Clephalopodes II. .Aiiomalie de la radule chez Octopus vulgaris Lamarck et observa- tions sur la seriatioii et I'asymetrie. Musee royal d'Histoire naturelie de Beigit|iie 9(3), 8 p. .\dam, W 1941. NotessurlesCephalopodes. XV. Sur la valeur diagnostique de la radule chez les Cephalopodes Octo- podes Musee roval d'Histoire naturelie de Belgique 17(38), 19 p. Benham, W. B. 1942. The octopodous Molliisca of New Zea- land. 1. The midget octopus of coastal waters. Transactions of the Royal Society of New Zealand 72:226-236. Dell, R, K. 19.52. The recent Cephalopoda of New Zealand- Dominican Museum Bulletin No. 16, 1.57 p. ICZN. 1985. International code of zoological nomenclature, 3rd ed L'niversity of California Press, 338 p. Massy, .\. L. 1916, Cephalopoda. British Antarctic ("Terra Nova") Expedition, 1910. Zoology 2(7):141-175. Pickford, G. 1955. A revision of the Octopodinae in the col- lections of the British Museum. Bulletin of the British Museum of Natural Histor\ 3(3):151-167. Robson, G. C. 1925. On seriation and as\ mmetr\ in the ceph- alopod radula. Journal of the Linnean Societ\ of London, Zoology 36:99-108, Robson, G. C. 1929. A monograph of the Recent Cephalop- oda. Part 1, Octopodinae, British Museum (Natural His- tory), London, 236 p,, 7 pis. Smith, E. A. 1902. Report on the collections of natural histor\ made during the vovage on the "Southern Cross," Vol. VII, Mollusca, p, 201-213, 2 pis, Tait, R, W, 1982, A taxonomic revision of Octopus australis Hoyle, 1885 (Octopodidae: Cephalopoda), with a rede- scription of the species. Memoires of the National Museum Victoria, No, 43:1.5-24, Toll, R B 1981, Bcnthoctopus oregonae, a new species of octopod (Mollusca; Ophalopoda) from the southern Ca- ribbean with a redescription of Bcnthoctopus januarii (Hoyle, 1885). Bulletin of Marine Science 31(l):83-95. Voss, G. L. 1973- Present status antl new trends in cephalopod s\stematics. Svmposium of the Zoological Societv of Lon- don No. 38:49-60. 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Accepted niarinseripls iire re- turned to the authf>r lor eonsidcralion ol ihe revirwc'is' comments, A linaliwd version ol I he nitinnsi'rlpl Is re turned to the editor and scnl to pit-ss, Two srln ol prools are $erit U) the author lor correction ( ^liuriKeit iiihcM llitih ■tling errors will be elidri/.ed lo llie inillioi Ml inul '■( ol corredcd prools ulioilld be leiil In llie t'dlliii as vx>i\ ii>> p(;ssible Aulhoiii wllh Inollliilloniil, ijiiinl m other Ti^:nri:\i si/pporl will be billed Im imi/i . Imiwi ■ nl Uf, rati; lA $W,(Hi [ict printed iiuu,k An order (onn lor (eprlnU will m » onipitri', ihi |i|!miI': Pieprint', limy b'' //Ml<'(<'d (liM/dKli ihc fdlloi Mitiiuviifih, ('(/»»<'' ted piKoU iind i'Miii'4|litlMli'iiM n j^rdif"' "l''.'iid iiiuiU'iD ijlionlil be leot itr l)i Ml, iUtii. Ui'ii. UMoiiiinii MMl|ii4ntypes of Antiplanes beringi (Aurivillius) were made available by Dr. A. Waren (Na- turhistoriska Riksmuseet, Stockholm). Photographs of the types of VV, H. Dalls, E. A. Smith's and T. Habe's ta.xa were provided by Dr. J. H. McLean (Los Angeles County Museum of Natural History), Mrs. A. Thompson (British Museum (Natural History)) and Dr. A. Matsukuma (Na- tional Science Museum, Tokyo), respectively. When studying shell variability we counted the num- ber of spiral cords on the body whorl. Since the number and degree of cord development on the anal fasciole were quite variable, only cords below the fasciole were counted. In some cases the relative height of the body whorl was an important taxonomic character. In a few cases in which the upper whorls were lost, ratios of the body whorl height to the height of the last three whorls were used. Ratios of the length of the marginal teeth of the radula to shell height were calculated using the mean length of at least ten marginal teeth, measured with an ocular micrometer calibrated to 0.001 mm. The following abbreviations for scientific research in- stitutes are used: BM(NH) — British Museum (Natural History), London CAS — California Academy of Sciences, San Francisco NRS — Naturhistoriska Riksmuseet, Stockholm NSMT — National Science Museum, Tokyo SBMNH — Santa Barbara Museum of Natural History SDNHM — San Diego Natural History Museum UCMP — Museum of Paleontology, University of Cali- fornia, Berkeley USNM — National Museum of Natural History, Smith- sonian Institution, Washington, DC. Page 120 THE NAUTILUS, Vol. 105, No. 4 Figure 1. Variability of anal sulcus form in Antiplanes saiic- tiioannis (Smith, 1875). Growth lines along different whorls of the same specimen are connected by dotted lines. ZIN — Zoological Institute of the U.S.S.R. Academy of Sciences, Leningrad ZM — Zoological Museum of the Moscow State University ACKNOWLEDGEMENTS We thank Dr. Alexander Golikov, Zoological Institute, Dr. Lev Moscalev, Institute of Oceanology, Dr. Dmitri Ivanov, Zoological Museum of Moscow State University, and Dr. Ronald Shimek, who kindly provided material for this study. We are grateful to Dr. Anders Waren, Naturhistoriska Riksmuseet, for the opportunity to study the syntypes of Antiplanes heringi, and to Dr. James H. McLean, Los Angeles County Museum of Natural His- tory, Mrs. Anne Thompson, British Museum (Natural History), and Dr. Akihiko Matsukuma, National Science Museum, Tokyo, who kindly sent photos of type speci- mens. We are also greatly indebted to Mrs. Saida I. Ja- gudina, Mrs. Sofia I. Strelzova, and Mrs. Marina A. Dol- golenko, laboratory assistants at the Zoological Institute, for their assistance during our studies of their collections. We want to thank Dr. M. G. Harasewych, Editor, for his help in checking the manuscript and making it more clear for the readers. HISTORICAL REVIEW OF THE GENUS ANTIPLANES The first species presently referred to Antiplanes to be described was Pleurotoma perversa Gabb, 1865. Dall (1902) proposed the genus Antiplanes and described 17 Recent and fossil species from American and Asian wa- ters (Dall, 1874, 1902, 1919, 1925). Bartsch (1944a,b) described three additional American species and isolated the dextral species of Antiplanes as a separate genus Rectiplanes. Bartsch began a revision of these groups, but this was never completed or published. Three Jap- anese species were later described by Habe (1958) who also established the subgenus Rectisulcus to include those dextral species with well-developed spiral sculpture. Antiplanes differs from most other members of the family Turridae in containing sinistral forms. The tax- onomic status of these sinistral forms has been variously interpreted by recent workers. Based on the study of extensive collections of fossils and Recent specimens from California, Grant and Gale (1931) concluded that the direction of shell coiling was subject to intraspecific vari- ability and synonymized almost all species known at that time with A. perversa. This view was shared by Golikov and Gulbin (1977) and by Golikov and Scarlato (1985). Bartsch (194a,b), however, considered sinistrality to be of generic significance. It was shown in recent malacological literature that most sinistral forms, including those of Turridae, are separate species (Tippett, 1983; Kantor, 1990). However, sinistral individuals and even isolated sinistral popula- tions lacking taxonomic status have been reported (e.g., the sinistral population of Conns mediterraneus, see Donati et. al. 1984). Analysis of the available material showed that sinistral Antiplanes from the northwestern Pacific belong to a single separate species. We have not found any examples of sinistral individuals or populations within normally dextral species of Antiplanes. SPECIES VARIABILITY AND CRITERIA FOR SPECIES DIAGNOSES In our revision of this genus, we encountered an unusu- ally high degree of variability in many conchological features, along with a general paucity of shell diagnostic characters. Thus, characters of shell form and proportions of its parts appeared to be taxonomically insignificant in most cases. Hickman (1976) documented the taxonomic importance of anal sinus shape in Turridae. Our analysis of intraspecific and ontogenetic variation of this feature in nearly 100 individuals of A. sanctiioam^is as well as in other species showed a very high degree of variability overlapping between species (figure 1). Although we considered a great many potential con- chological characters, the most taxonomically significant feature was determined to be the number of spiral ribs below the anal fasciole. Based on this character, two large groups of species can be distinguished; one is character- ized by complete or nearly complete absence of spiral sculpture (A. ahyssalis n. sp., A. obliquiplicata n. sp., A. dendritoplicata n. sp. and A. kawamurai). the other, containing the remaining species treated herein, by the presence of variably developed spiral sculpture. It should be emphasized that the number of spiral ribs on the body whorl, being an ontogenetically and geographically vari- able feature, allow s onlv statistical differentiation of spe- cies (figures 117, 118, i38). Y I Kantor and A. V. Svsoev, 1991 Page 121 Figures 2-10. Radulae of Antiplanes spp. Shell lengths and catalog numbers are given in parentheses. 2. Antiplanes den- dritoplicata n. sp., paratype (28.3 mm, ZIN N 6/54581); 3. A. obliquiplicata n. sp., paratype (40.3 mm, ZM N Lc 6899); 4. A. spirinae n. sp., paratype (22,4 mm, ZM N Lc 6905); 5. A. isaotakii (Habe, 1958) (30.8 mm); 6. A. hurilensis n, sp,, para- type (28.8 mm, ZIN N 1/54583); 7, 8. A. habei n. sp. 7. holotype (38.0 mm, ZM N Lc 6909); 8. paratvpe (23.1 mm, ZM N Lc 6910); 9, 10. A. vinosa (Dall, 1874). 9. (36.3 mm), 10. (28.6 mm). Scale bars all 100 ixm. It is interesting to note that similar patterns of mor- phological variation occur in different species. Specimens of A. sanctiioannis from Sakhahn Bay have fewer ribs on their body whorl than individuals from the Kurile Islands. The same pattern is seen in A. vinosa. a sister species of A, sanctiioannis. The morphology of the radula offers additional char- acters of taxonomic importance. Though the form of the marginal teeth was similar in all species (figures 2-17), rates of radular growth permitted species to be segre- gated into three groups; those with rapidly, moderately and slowly growing marginal teeth (figure 18). A geo- graphical component to the variability of this character was observed. However, comparisons of specimens of the same size from the same locality showed distinct differ- ences between species. In figure 18, points representing different species from the same locality are connected by dotted lines. The groups of species isolated on the basis of radular Figures 1 1-17. Radulae of Antiplanes sanctiioannis (Smith, 1875). The collection locality and shell length of individual specimens follow 11. Sakhalin Bay, 45.0 mm. 12. Sakhalin Bay, 41.9 mm; 13. Tatar Strait, 38.0 mm; 14. Sakhalin Bay, 36.0 mm; 15. Southern Kurile Islands, 34.5 mm; 16. Eastern Kamchatka, 42.0 mm; 17. Tatar Strait, 25.0 mm. Scale bars all \00 lum. growth pattern also differed in the degree of spiral sculp- ture. Species with the most slowly growing marginal teeth (A. dendritoplicata n. sp. and A. obliquiplicata n. sp.) generally lack spiral sculpture. Species with mod- erate marginal tooth growth rates (A. sanctiioannis, A. vinosa, A. spirinae n. sp., and A. hurilensis n. sp.) are characterized by more numerous and less developed spi- ral cords than the group of species with rapid marginal tooth growth (A. habei n. sp. and A, isaotakii). As with the variation in marginal tooth growth rates, species and species groups based on shell sculpture can onK be dis- tinguished statistically. In some cases, the relative height of the body whorl was also useful for distinguishing similar species. Pairs of species such as A. sanctiioannis and A. hurilensis n. sp. as well as A. dendritoplicata n. sp. and A. obliqui- plicata n. sp. (figure 46) were readily distinguished by this character The systematics of the genus Antiplanes were difficult to discern using traditional techniques due to high on- togenetic and geographical variability of nearly all gen- erally accepted diagnostic characters. We have been able to distinguish close but distinct species by using graphs Page 122 THE NAUTILUS, Vol. 105, No. 4 50 lA lA 42 40 38 36 34 32 30 28 26 24 22 20 18 16 14 (mm; 0.13 .14 .16 .18 .20 .22 2U 26 .28 mm [ Figure 18. Rales of radular growth in species of Antiplanes. Crossed circles — A. dendritoplicafa u. sp.; open squares — A. spirinae n. sp.; open circles — A. obliquiplicata n. sp.; solid cir- cles— A. sanctiioannis (Smith); open ovals — A. vinosa (Dall); open triangles — A. isaotakii (Habe); solid triangles — A. habei n. sp. The lines of regression: 1. A. dendritopHcata n. sp.; 2. A. sanctiioannis (Smith); 3. A. vinosa (Dall); 4. A. habei n. sp. Ll — mean length of marginal tooth, Hs — shell length. and regression analyses of various characters relative to shell height. This approach forms the basis of the follow- ing systematic treatment of this genus. SYSTEMATICS Family TURRIDAE Svvainson, 1840 Subfamily Turriculinae Powell, 1942 Genus Antiplanes Dall, 1902 Dall, 1902513 (type species, by original designation— P/ph- roloma (Surciila) perversa (Jabb, 1865). Rectiplanes Bartsch, 1944b:59 (type species, iiy original des- ignation— Pleuroloma (Anliplancs) santarosana Dall. 1902). Rectiplanes (Rectisulcus) Habe, 1958:184 (type species, by original designation — Rectiplanes {Rectisidcus) motoji- mai Habe, 1958) Diagnosis of ihe genus: Shell large (to 72 mm), dextral or sinistr.il fusiform, covered by smooth, thick perios- tracum. A.muI sculpture absent. Spiral sculpture may be absent or coi: posed of ribs and dendritic threads (figure Figure 19. Diagrammatic representation of the shell base of Antiplanes habei n. sp. AF — anal fasciole, DL — dendritic ribs, GL — growth lines, SR — spiral ribs. 19, DL) that may be obliquely and irregularly disposed, with ramifying small threads radiating from extremities of anal sinus growth lines. Anal sinus deep, located on shoulder or near periphery . Siphonal canal long, occa- sionally slightly curved. Operculum (figures 20-25) large, drop-shaped, terminally nucleated, filling the aperture. Radula "nontoxoglossate", consisting of proximally bi- furcated marginal teeth and of vestigial rachidian teeth. Type species: Pleitrotoma {Siircula) perversa Gabb, 1865, the originally designated type species of Anti- planes, is a primary homonym of Pleurotoma perversa Philippi, 1846. Grant and Gale (1931) considered P. per- versa Gabb to be a synonym of P. voyi Gabb, 1869. However, the original descriptions and illustrations dem- onstrate that these are different species. Indeed, one of them {P. perversa) is sinistral while the other {P. voiji) is dextral. As no available name exists for Pleurotoma perversa Gabb, 1865, we here propose Antiplanes gabin, nomen novum, as a replacement name. S\ ntypes (UCMP 15929 and 31547) Catalina Island, California (Recent); (UCMP 15930), Pleistocene of San Pedro, California (Coan & Bogan, 1988). Distribution: Species of Antiplanes live in the northern part of the Pacific Ocean from California to British Co- lumbia, along the American coast and from Honshu, Japan to the Gulf of Anadyr, Bering Sea, along the Asian coast, at sublittoral to upper abyssal depths. In the Bering Sea, Antiplanes v\ere found in western, eastern and cen- tral parts. Neither the literature nor samples at our dis- posal contain records of Antiplanes from the Gulf of Alaska. According to our data, western American (south- ward of the Gulf of .■\la-ska) and .\sian (including the Bering Sea) faunas have no species in conmion. Y. I. Kantor and A. V. Svsoev, 1991 Page 123 Figures 20-25. Opercula of Antiplanes spp 20. A, obliquiplicata n. sp , holotype, ZM N Lc6898, 40.3 mm 21. A. dendritoplicata n. sp., paratype, ZIN N 6/54581, 28.3 mm. 22. A. spirinac n. sp., paratype, ZM N Lc 6905, 16.3 mm. 23. A. habei n. sp., paratype, ZIN N 1/54627, 33.3 mm. 24. A. vinosa (Dall), 36.3 mm. 25. A. sanctiioannis (Smith), 42.7 mm. Fossil species of Antiplanes are known from the Eo- cene of North America {Drillia cooperi Dickerson, ac- cording to Grant and Gale, 1931). In the eastern Pacific, the genus was frequently found in Pliocene deposits of Oregon and Washington (Grant & Gale, 1931; Addicott, 1976) and in Pliocene and Pleistocene of California (Grant & Gale, 1931). In the western Pacific, earliest occurrences of the genus were recorded from the Middle Miocene of Japan (A. sadoensis (Yokoyama) — Chinzei, 1959, and A. sanctiioannis (Smith) — Amano, 1983). Species of Anti- planes are abundant in Pliocene deposits of Honshu and Hokkaido, Japan (Masuda & Noda, 1976). Antiplanes voiji and a dextral Antiplanes sp. were found in the upper Pliocene to lower Pleistocene of eastern Kamchatka (Pe- trov, 1982). Remarks: The genus Rcctiplanes was erected b\ Bartsch (1944b) to contain dextral species only. No other char- acters distinguishing Rectiplaiies and Antiplanes can be identified. Since we do not consider direction of shell coiling to be of more than a species level character, Rectiplanes is regarded as a synonym of Antiplanes. Page 124 THE NAUTILUS, Vol. 105, No. 4 Figures 26-29. Light (26, 27) and SEM (28, 29) micrographs of radulae, 26-28. Antiplanes saiictiioannis (Smith). 29. A. vinosa (Dall). Y. I. Kantor and A. V. Sysoev, 1991 Page 125 Figures 30-32. Diagrammatic representations of transverse sections of the radula in the radiilar sac. 30. Olivella burcalis Gohkov, 1967. .3 1 . Splcndrillia chatamcnsis S\ soe\' and Kantor, 1989. 32. Antiplanessanctiioanni.s (Smith. 1.S75). cf — "central formation", ct — central tooth, It — true lateral tooth, "It" — the so called "lateral" tooth of Turridae. mt — true marginal tooth, "mt ' — the so called "marginal" tooth ol Turridae. Habe (1958) established the subgenus Rectisulcus for dextral species with well developed spiral ribs. As there is a continuum in the degree of spiral rib development, we follow Powell (1966), in synonymizing Rectisulcus with Antiplanes. Varying opinions e.xist as to the composition of the transverse rows of teeth in the Antiplanes radula. A reduced central tooth having the form of a narrow plate pointed at both ends occurs in the eastern Pacific species A. santarosana and A. thalaea (Powell, 1966. figures 57, 58). However, McLean (1971) indicated that eastern Pa- cific species of Antiplanes lacked central teeth. Struc- tures superficially resembling the central teeth of other Turriculinae were seen in Antiplanes by means of light microscopy (figures 26, 27). Only the anterior and parts of the lateral borders of these optically transparent struc- tures were clearly seen in unstained preparations (figure 26). When the radula was stained with Orange G and Aniline Blue, these structures appeared to be paired and symmetrical (figure 27). However, scanning electron mi- croscopy showed that there were no distinct tooth plates on the central part of the radular membrane (figures 28, 29). The membrane had only indistinct folds that are similar to those of some other Turriculinae. The manner in which the radular membrane is folded within the radular sheath is significant for evaluating the nature of these central structures. The radular membrane of prosobranch mollusks is folded within the radular sheath. These folds occur between groups of teeth, or between separate teeth if the radulae are greatly oligo- merized. Taenioglossate radulae, for example, have two pairs of folds, one between the central and lateral teeth, the other between the lateral and marginal teeth The same pattern occurs in those Neogastropoda that have five teeth per transverse row {Olivella. figure 30). How- ever, in turrids with five teeth per transverse row (sub- family Clavinae) there is only one pair of folds, placed between the central and lateral teeth. We have also ob- served this pattern in two species of Splcndrillia (figure 31). Study of the radula of Antiplanes shows that the rad- ular membrane has but a singular fold situated exactK- along the middle of the "central structure." Moreover, the membrane can be easily torn along this bend. In our opinion this suggests that the "central formation" is a rudiment of a pair of lateral teeth but not of the central tooth. We propose that the central tooth was completely reduced during evolution, and that the paired folds of the membrane have then grown nearer and fused into a single bend. Maes (1983) proposed a similar evolution- ary pathway for Crassispira (Crassiclava) apicata (Reeve) (figures 31, 32), and referred to these structures as "soft lateral teeth". Although studies of the embryonic devel- opment of the radula of Antiplanes would shed further light on this question, such data is presently not available. The competing hypotheses, one of weakening of the central tooth and complete reduction of lateral teeth, the other of loss of the central tooth along with regaining of the weakened lateral teeth, cannot be fully resolved at this time. WESTERN PACIFIC SPECIES ERRONEOUSLY ATTRIBUTED TO ANTIPLANES Rectiplanes (?) yukiae Shikama, 1962. Shikama, 1962: .50, pi. 3, figs. 3a,b, 4a,b. Type locality: Off Choshi, eastern Honshu — Japan, 250- 300 fms. The species was subsequently reassigned to the genus Bent hodaphne Oyama. 1962 (Okutani, 1964, 1968). Rectiplanes (Rectisulcus) hayashii .Shikama, 1977. Shikama, 1977:19, pi, .3, fig. 12a,b. Type locality: Ensyu-nada. south-western Japan This species has well developed axial sculpture and is here referred to the genus Comitas Finlay. 1926. RECOGNIZED SPECIES OF ANTIPLANES Antiplanes abyssalis new species (figures .33-35, 45) Material examined: Holotype: R/V 'Vityaz', 39th cruise, sta. 5603, south-east of Simushir, Kurile Islands, 46°22'N, 1.5.3°03'E, depth 3,175-3,250 m, 15.vn.l966 (ZM N Lc 6889); parat\pe R/V "Vityaz', 14th cruise, sta. 2210, south- east of southern Kamchatka, 50°01.8'N, 157°.39.2'E, depth 2,430-2,670 m, 24.VI.1953 (ZM N Lc 6890). Description of holotype: Shell slender, with slightly curved axis, consisting of 6.5 teleoconch whorls and at least 1.5 heavily eroded protoconch whorls. Whorl profile angulated at periphery , whorl shoulder concave. Sutures deep. Body whorl high, comprises about 0.6 of shell height. Shell base weakly convex, passing gradually and without bending into a moderately elongate siphonal canal. Growth lines distinct. Spiral sculpture nearly absent, rep- resented by single somewhat wavy riblet on anal fasciole and b\ marked dendritic ribs that reach upper and lower sutures of spire whorls but disappear on shell base. Anal sinus wide, rounded. Aperture wide, elongate-oval, poor- Page 126 THE NAUTILUS, Vol. 105, No, 4 Y. I. Kantor and A. V. Svsoev, 1991 Page 127 ly differentiated from siphonal canal, outer lip attaclied to columella at acute angle. Columella straight, covered witfi narrow, white, glossy callus. Color light-yellow. Periostracum thin, smooth, glossy, fitting closely to hy- postracum. Dimensions of holotype: Shell height 31 0 mm, body whorl height 18.7 mm, aperture height 14.2 mm, shell diameter 12.0 mm. Remarks: The paratype differs from the holotype in having a somewhat darker periostracum, a less concave profile of the shoulder, and a wider siphonal canal. The spiral riblet on the anal fascicle is weaker in the paratype. This species differs from Antiplanes dendritoplicata n. sp., its nearest relative, in lacking spiral ribs on the shell base and siphonal canal, and from A. ohliquiplicata n. sp. in having a higher body whorl as well as in whorl profile. Distribution: This species was found along the Kurile- Kamchatka trench at depths of 2430-3250 m (figure 45). Antiplanes dendritoplicata new species (figures 2, 21, 36-44, 45) IVlaterial examined: Holotype (ZM N Lc 6891) and 3 paratypes (ZM N Lc 6892):' R/V 'Vitvaz', 39th cruise, sta. 5641, east of Iturup, southern Kurile Islands, 44°46'N, 148°45'E, depth 472-479 m, Sigsbee trawl, 10.1X1966; paratypes: R/V 'Vityaz', 12th cruise, sta. 1779, north- eastern Okhotsk Sea, depth 464 m, Sigsbee trawl, 13.x. 1952 (2 specimens— ZM N Lc 6893); 12th cruise, sta. 1831, northwestern Okhotsk Sea, 56°57.5'N, 145°57.0'E, depth 196 m, Sigsbee trawl, mud, 15. X. 1952 (2 specimens— ZM N Lc 6894); 24th cruise, sta. 3578, east of Honshu, 38°35.0'N, 142°53.3'E, depth 1,641 m, Sigsbee trawl, clay, 11. V, 1957 (1 specimen — ZM N Lc 6895); 39th cruise, sta. 5638, east of Iturup, 44°36'N, 149°07'E, depth 1,675-1,845 m, Galathea trawl, 10.1X1966 (1 specimen— ZM N Lc 6896); 52th cruise, sta. 6671, east of Honshu, 40°12.0'N, 143°35.8'E. depth 2,400-2,720 m, Sigsbee trawl, 23.VI.1972 (1 specimen— ZM N Lc 6897); Okhotsk Sea, sta. 139/20, 55°45'N, 145°50'E, depth 218 m, sandy mud, 1. VIII. 1930, coll. P. V. Ushakov (3 specimens— ZIN N 1/54576); R/V To- porok', sta. 63, southeastern Sakhalin, 46°49.5'N, 143°52.3'E, depth 187 m, 4.IX.1947 (1 specimen— ZIN N 2/54577); sta. 70, southeastern Sakhalin, 46°26.2'N, 143°22.0'E, depth 103 m, mud with gravel, 6.1X1947 (1 specimen— ZIN N 3/54578); sta. 13-a, northwest of Hok- kaido, 44°47.6'N, 144°29.8'E, depth 700 m, 26.VIII.1948 Figure 45. Geograpiiir distribution oi Antiplana^ abyssalis u sp. (open circles), A. dendritoplicata n. sp. (solid circles), A. ohliquiplicata n. sp, (solid scjuares). (3 specimens— ZIN N 4/54579); sta. 17, off Shikotan, 43°37.1'N, 147°02.8'E, depth 1,450-1,530 m, Sigsbee trawl, mud, 28. VIII. 1948 (2 specimens— ZIN N 5/54580); sta. 100, off Iturup, 44°42.0'N, 147°56.7'E, depth 299 m, 14.1X1949 (1 specimen— ZIN N 6/54581); sta. 101, off Iturup, 44°20.8'N, 148°24.0'E, depth 414 m, 14.1X1949 (1 specimen— ZIN N 7/54582). Description of holotype: Shell slender, fusiform, com- posed of 5 teleoconch whorls and 1.5 heavily eroded protoconch whorls. Surface of upper 2.5 teleoconch whorls heavily eroded, rose-colored. Coiling axis slightly curved. Body whorl high, convex, comprising 0.64 of shell height. Teleoconch whorls convex, evenly rounded, divided by deep sutures. Shell base weakly convex, almost flat, pass- ing into long siphonal canal without pronounced bend. Growth lines distinct. Spiral sculpture of indistinct, in- terrupted riblets on anal fasciole (3-4 on body whorl, 2- 3 on penultimate whorl) and of moderately developed dendritic ribs that reach sutures on spire whorls but be- come smooth on shell base. Siphonal canal with 8 indis- tinct spiral ribs separated by shallow grooves. Aperture Figures 33-44. Shells of Antiplanes spp. 33-.35. Antiplanes abyssalis n. sp. 33. lioiot\pe, ZM N Lc 6889, 3L0 mm. 34. 35. paratype, ZM N Lc 6890, .36.2 mm. 36-44. Antiplaties dendritoplicata sp. nov. 36, 37. Holotype, ZM N Lc 6891, 414 mm 38. Paratype, off Shikotan, Kurile Lslands, ZIN N 5/54.580, 35.2 mm. 39. Paratype, eastward off Iturup, southern Kurile Islands, ZM N Lc 6892, 32.6 mm. 40. The shell base of 42, note the spiral ribs on the canals. 41, 42. Paratype, northwest of Hokkaido, ZIN N 4/54579, 37.0 mm. 43. Paratype, east of Chonshu, ZM N Lc 6897, 27.0 mm. 44. Paratype, northwestern Okhotsk Sea, ZM N Lc 6894, 47.7 mm. Pa, 12t> THE NAUTILUS, Vol. 105, No. 4 (mm) 40 36 j 32 28 24- 20 16 0.66 .68 .70 .72 % •76//, w//y Figure 46. Comparison of the relative height of the body whorl of A. dendritoplicata n. sp. (open circles, 1 — line of regression) and .4. obliquiplicata n. sp. (solid circles, 2 — line of regression). The shell height is plotted along the abscissa, the ratio between the bod\ \\ horl height to the height of the last 3 \\ horis is plotted along the ordinate. high, elongate, oval. Columella slightly thickened by white, glossy callus. Siphonal canal differentiated from aperture by its parallel margins. Shell color light-olive. Periostracum thin, glossy. Dimensions of holotype: Shell height 41.4 mm, body whorl height 26.6 mm, aperture height 20.7 mm, shell diameter 16.0 mm. Remarks: Paratypes have less convex whorls, more dis- tinct and closely set riblets on the anal fasciole (up to 4 on spire whorls), a more weakly developed bend at junc- ture of the body whorl periphery and the shell base, and a shorter siphonal canal that is better differentiated from the shell base. Ribs on the siphonal canal of paratypes may be more numerous (up to 15) and extend onto the shell base. Radulae of 6 paratypes were examined (figure 2). Rad- ular teeth are small, the shell height/mean marginal tooth length ratio ranges from 159.6 (shell height 22.5 mm) to 266.5 (shell height 47.7 mm). Mean length of marginal teeth increases slowly. The linear regression equation is: Y = 0.1052 -I- 0.0016 X, where Y is the mean length of marginal teeth and X the shell height. Antiplanes dendritoplicata n. sp. is most similar to A. obliquiplicata n. sp., and the variable shell forms of both species overlap. However, A. dendritoplicata always has ribs on the siphonal canal (figure 40). These ribs are invariably absent in A, obliquiplicata (figure 52). These two species can also be distinguished by significant dif- ferences in the relative height of the body whorl (body whorl height/height of last three last whorls — figure 46). The relative height of the bod\ whorl decreases linearK during growth. In young individuals (shell height <30 mm), differences between species are less clearly dis- cerned, but these two species differ significantly in mean values. The regression equations are: A. dendritoplicata Y = 0.7344 - 0.0011 X A. obliquiplicata Y = 0.7485 0.0004 X, where Y is the relative height of the body w horl and X is the height of 3 last whorls. Distribution: This species was recorded from eastern Honshu to the northern part of the Okhotsk Sea at depths from 103 m (south-eastern Sakhalin) to 2,720 m (east of Honshu) (figure 45). Antiplanes obliquiplicata new species (figures 3, 20, 45, 47-52) Material examined: Holotype (ZM N Lc 6898) and 6 paratvpes (ZM N Lc 6899): R/V 'Vityaz', 24th cruise, sta. 3577, E of Honshu, 38°40.1'N, 143°29.3'E, depth 3042 m, mud, 10. V. 1957; paratypes: R/V 'Vityaz', 5th cruise, sta. 618, western Bering Sea, 57''03.5'N, 168°30.5'E, depth 3,875 m, Sigsbee trawl, mud. 25. IX. 1950 (1 spec- imen—ZM N Lc 6900); 14th cruise, sta. 2209, E of south- ern Kamchatka, 49°46'N, 157°48'E, depth 3,660 m, Sigs- bee trawl 23. V. 1953 (4 specimens— ZM N Lc 6901 ); 24th cruise, sta. 3578, E of Honshu, 38°35.0'N, 142°53.3'E, depth 1641 m, Sigsbee trawl, clav, 11. V. 1957 (1 speci- men—ZM N Lc 6902). Description of holotype: Shell slender, fusiform, com- posed of 8 whorls. Three upper whorls heavily eroded. Two upper whorls considerably smaller in diameter, probably representing protoconch. Bod> whorl convex, comprising 0.56 of shell height. Shell covered by thin, olivaceous, somewhat glossy, non-exfoliating periostra- Figures 47-38. Shells of Antiplanes spp. 47-52. Antiplanes obliquiplicata n. sp 47. 48. Holotype. ZM N LC 6898, 40.3 mm. 49. I'aratype, east of Choiishii, 7A1 N Lc 6899, 36.2 mm 50. Paratype, east of Chonshu. ZM N Lc 6902, 32.0 mm. 5L Paratype. the same sample as 49, ZM N Lc 6899, 20.8 nnn 52. The shell base of liie hoiot\ pe. note the absence of spiral ribs on the canal. 53-58. Antiplanes kawamurai (Habe, 1958). 53, 54. Holotype, NSMT N 38598. 35.8 nun. 55. Off Shikotan, Kurile Islands, ZIN N 56.322, 52.1 mm. 56, 57. East of C;honshn, 30.6 mm 58. Tiic same sample as 56, 57. 1 1.7 mm Y. I. Kantor and A. V. Sysoev, 1991 Page 129 Page 130 THE NAUTILUS, Vol. 105, No. 4 cum. Teleoconch whorls highly convex, evenly rounded, separated by deeply impressed suture. Shell base ex- tremelv convex, giving rise to long, broad, curved si- phonal canal. Growth lines thin, indistinct. Axial folds oriented parallel to growth lines below anal fascicle on upper whorls, absent on body whorl. Anal sinus deep, wide, rounded. Spiral sculpture of well-developed den- dritic threads that reach sutures on spire whorls and siphonal canal on body whorl. Spiral riblets (2-3) can be traced on anal fasciole of upper whorls. Spiral sculpture absent on shell base antl siphonal canal. Aperture elon- gate-oval, well differentiated from siphonal canal. Col- umella concave, covered by white, glossy callus. Dimensions of holotype: Shell height 40.3 mm, body whorl height 22.7 mm, aperture height 18.0 mm, shell diameter 15.5 mm. Remarks: Radular teeth of the paratype (figure 3) are small. Mean length of marginal teeth is 0.124 mm (shell height/tooth length = 325). The shoulders and shell bases of the paratypes are weakly angular; their whorls may be more flattened. The anal fasciole and the part of the whorl nearest to the upper suture may be covered with thin spiral cords. The canal may be straighter and more differentiated from the aperture. This species is most similar to A. dendritoplicata, and may be differentiated from that species as discussed in the remarks for that species. Distribution: Antiplanes obliquiplicata ranges from Honshu to eastern Kamchatka at depths of 1,641 to 3,875 m (figure 45). Antiplanes kawamurai (Habe, 1958) (figures 53-58) Rectiplanes kawamurai — Habe, 1958:181-184, text-fig. 1; Shi- kama, 1962:50-51, pi. 3. figs. la,b, 2a; Okutani, 1964:423, pi. 14. fig. 6; 1968:37, pi, 3, fig. 7. Type specimens: Holotype (NSMT N 38598), shell height 35.8 mm; paratype (NSMT N 52671), shell height 34.8 mm. Type locality: Off Choshi, Chiba Pref., Honshu, Japan, in 183-274 m. Material examined: Off Shikotan, Kurile Islands, 43°35.5'N, 147°20.5'E, depth 1,450-1,530 m, mud, 28.VII.1948 (1 specimen ZIN N 56322); E of Honshu, Japan (4 specimens). Description: Shell light greenish or brownish, broadly fusiform, with flat-sided spire outline. Body whorl mod- erately convex, tapering rapidly to join broad, short si- phonal canal. Whorls weakly convex, angulated below anal fasciole, nearly turreted in young specimens. Anal fasciole usually concave, with spiral sculpture absent or of indistinct threads. Dendritic cords well developed, reaching suture on spire whorls, occasionally reaching shell base. Aperture oval, broad, outer lip joining colu- mella at right or obtuse angle. Shell height reaching 56 mm (Shikama, 1962). Remarks: This species is easily distinguished from other species of Antiplanes with weakly developed spiral sculpture (A. dendritoplicata, A. obliquiplicata, A. abys- salis) by its broad shell and angulated whorls. Distribution: Antiplanes kawamurai inhabits the east- ern coast of Japan from Sagami Bay to the southern Kurile Islands (Shikotan Island), at depths of 1,520 m (Sagami Bay— Okutani, 1968) to 183 m (type locality). Antiplanes sanctiioannis (Smith, 1875) (figures 1, 11-17, 25, 26-28, 59-88, 100-103) Plenrotoma (?*) Sancti-Ioannis Smith, 1875:416-417. PU'urotonia Beringi Aurivillius, 1887:377, Taf. 13, fig. 3. Plenrotoma {Antiplanes) piona Dall, 1902:514. Antiplanes piona Dall, 1925:4, pi. 21, fig. 5. Figures 59-69. SheWs^A Antiplanes spp.59-b2. Plenrotoma sancliioanrnsSmUh. 1875. 59,60. Lectotype, BM(NH) N 1873.8.6,16, 39.2 mm. 61, 62. Paralectotypes, BM(NH) N 1873.8.6.16, 39.9, 23.2 mm, 63-66. Plenrotoma beringi Aurivillius, 1887. 63, 64. Lectotvpe, NSR N 1557, 38.3 mm. 65, 66. Paralectotypes, NSR N 1557, 37.7, 16.6 mm. 67. Plenrotoma (Antiplanes) piona Dall, 1902, holotype, USNM 109179, 42,0 mm, 68. Antiplanes yessoensis Dall, 1925, holotype, USNM 111053, .39 mm, 69. Antiplanes willetti Berry, 1953, holotype, 18.4 mm. Kiguri-s 70-82. Shells of Antiplanes sanctiioannis (Smith, 1875). 70-73. Tatar Strait. Okhotsk Sea (70, 71—39.3 mm, 72—35.1 mm, 73—33,8 mm), 74. Sakhalin Bay, Norlheni Sakhalin, Okhotsk Sea (41 3 mm), 75. Ofi^ Paramushir, northern Kurile Islands (.38.2 mm). 76. Off Onekotan, central Kurilt- Islands (37.2 mm). 77. Tatar Strait, Okhotsk Sea (32.0 mm) 78. Off lona Island, northwestern Okhotsk Sea (30.4 mm). 79. Off llurup, southern Kurile Islands (32 I mm). 80. Off eastern Sakhalin, (38.9 mm). 81- 82. Northern part of Okhotsk Sea (81—34.3 mm, 82—40.6 mm). Figures 83-99. Shells of Antiplanes spp. 83-88. Antiplanes sanctiioannis (Smith, 1875). 83. Tatar Strait. Okhotsk Sea (32.0 mm), 84. Bristol Bay, eastern Bering Sea (34,0 mm), 85-88. Bering Sea (85—31,7 m, 86, 87—29,6 mm, 88—39,9 mm), 89-93. Antiplanes kurilensis n. sp, 89, 90. Holotyix-, ZM N Lc 6903. 54,6 mm, 91. Paratype, Kurile Islands, ZM N Lc 6904, 52,3 mm. 92. Paratvpe, east off Shumshu, northern Kurile Islaiuls. ZIN N 1 5458.3, 28,8 mm 93. Off Moneron Island, Tatar Strait, sublossil specimen, ZIN N 2/54584, .35,4 mm 94-99. Antiplanes spirinae n. sp. 94, 95. Holotype, ZIN N 1 54585, 36.5 mm. 96-98. Paratypes, of Simiishir, central Kurile Islands, ZM N Lc 6905 (96—22.4 mm. 97—14.2 mm, 98—16,3 mm), 99. The shell base of the holotype. Y. I. Kantor and A. V. Svsoev, 1991 Page 131 Page 132 THE NAUTILUS, Vol. 105, No. 4 Y. I. Kantor and A. V. Svsoev, 1991 Page 133 Page 134 THE NAUTILUS. Vol. 105, No. 4 Figure 100. Geographic distribution of Antiplanes sanctiioannis (Smith, 1875) Antiplanes yessoensis Dall, 1925:4, pi 21, fig. 3. Antiplanes (Rectiplanes) willetti Berrv, 1953419-420, pi 29, fig. 2. Antiplanes sadoensis Otuka, 1949:pl 13, fig. 16 {sensu auct., non Yokoyama, 1926). Antiplanes thalaea Dall, 1919:37, pi 11, fig. 6 (sensu auct., non Dall, 1902:514), Type specimens: Lectotype of P. sanctiioannis BM(NH) 1873.8.6.16, shell height 39.2 mm; paralectotypes BM(NH) 1873.8.6.16, shell height 39.9 mm, 23.2 mm. Type locality, about 100 miles south-eastward of Yesso [Hokkaido, Japan]. Lectotvpe of P. beringi (here designated), NRS N 1557, shell height 38.3 mm; paralectotypes NRS N 1557, 4 specimens. Type locality, 'Vega' Expedition, Bering Sea, 62°39'N, 177°05'W, 55 fms. Holotype of P. piona USNM 109179, shell height 42 mm. Type locality, off S. coast of Kamchatka, 51°00'00"N, 157°48'00"E, in 96 fms, black sand, "Albatross", sta. 3644 fms. Holotype of A. yessoensis USNM 111053, shell height 39 mm. Type locality, Japan Sea, S. coast of Hokkaido, 41°58'00"N, 142°30'30"E, in 404 fms, brown mud, "Al- batross", sta. 5036. Holotype of A. willetti CAS 064665, shell height 18.4 mm; paratypes— SBMNH No. 34539, and SDMNH 22856. Type locality, off Forrester Island, SW Alaska, 50 fms. Material examined: Over 200 lots (about 700 speci- mens) were studied. Description: Shell very variable, from elongate fusiform (figures 70, 77) to broadly fusiform (figures 81, 82), white with periostracum from light yellow to dark-brown. Body whorl variously convex. Relative height of body whorl varies within wide range (0.54-0.67). Whorls evenly rounded to angular at the shoulder and shell base, oc- casionally, below periphery of spire whorls (form de- scribed as A. yessoensis). Anal fasciole concave, flat, or convex. Anal sulcus form varies considerably, even with- in same specimen (figure 1 ). Spiral sculpture of numerous relatively narrow ribs, usually weaker at upper part of the whorl periphery and especially above anal fasciole. Ribs with serrated borders where crossed by growth lines, covered with secondary riblets that are usualK incon- spicuous or absent. Dendritic ribs moderately to highly developed only on anal fasciole, do not reach sutures on spire whorls. Siphonal canal usually recurxed, varies from short, broad (figure 83) to long, narrow (figure 81). Shell height reaches 60 mm. Figure 101. The egg capsule of Antiplanes sanctiioannis (.Smith, 1S75). Y. I. Kantor and A, V. Sysoev, 1991 Page 135 Figure 102. Comparison of the number of spiral ribs on the body u horl below the anal fasciole in populations of Anttplanes sanctiioannis (Smith) The shell height along the abscissa, the number of spiral ribs along the ordinate Sakhalin Ba\ {open circles, 1 — line of regression), east Sakhalin (solid triangles, 2 — line of regression), and Tatar Strait (solid circles, 3 — line of regression). Radula relatively large, ratio between shell height and mean marginal tooth length ranging from 85.4 (shell height 20.3 mm) to 173.4 (shell height 35.9 mm) (figure 18). Length of marginal teeth increases rapidly during ontogeny Relationship between marginal tooth length and shell height nearly linear, expressed by regression equation: Y = 0.1401 + 0.0030X Egg capsules, most containing eggs, attached to shells of living A. sanctiioannis. Capsules were hemispherical (diameter — 3 mm), with narrow flange at base. Surface covered with minute concentric creases, containing translucent, nearly circular apical plug (diameter — 1 mm) (figure 101). Three capsules studied each contained up to 6 developing embryos. Remarks: Shell morphology of this species is highly variable. The spiral ribs above the anal fasciole are usu- ally absent, but may be poorly to well developed. Sculp- ture on the anal fasciole may consist of broad low ribs or very thin threads that may become obscure when Figure 103. Comparison of the number of spiral ribs on the bod\ whorl below the anal fasciole in populations of Antiplanes sanctiioannis (Smith). The shell height along the abscissa, the number of spiral ribs along the ordinate Southern Kurile Island (solid squares), northern Kurile (solid circles), east Kamchatka (open circles), and Bristol Bay (open triangles). dendritic ribs are well developed. The dendritic ribs are moderately to highly developed only on the anal fasciole and do not reach the sutures of the spire whorls. Spiral ribs become stronger, more widely spaced and usually broader below the anal fasciole. The number of ribs below the fasciole varies (3-12, usually 6-8, on the penul- timate whorl) within populations and during ontogeny. When secondary riblets are well developed, the primary ribs tend to be low and uneven. Only a single specimen from the Bering Sea had uniform, equally developed ribs over the entire shell surface (figures 86, 87), and we refer this specimen to A. sanctiioannis with some doubt. Although A. sanctiioannis is highly variable, different suites of morphological characters predominate in dif- ferent portions of the species range. Among populations from the vicinity of Sakhalin Island, those in the Tatar Strait and adjacent waters have shells with dark color- ation and distinct spiral sculpture that is more or less uniformly developed over the entire shell surface but that weakens slightly towards the suture (figures 70-73). Dendritic ribs are well-developed in most specimens from this area, but may be weak or limited to certain portions 136 THE NAUTILUS, Vol. 105, No. 4 Figure 104. Geographic distributinn of Anli)>l(incs kurilensis n. sp. (open circles) and A. spirinac n. sp. (solid circles). of the shell. Whorls are evenly rounded. The anal fasciole is flat or rarely concave. Numerous spiral ribs distinguish this population. Specimens from Sakhalin Bay (northwestern Sakhalin) tend to be relatively light in color, and often have a convex anal fasciole and inconspicuous sculpture of fewer spiral ribs (figure 74). Spiral ribs weaken towards the periphery and dendritic ribs are usually well developed. Intermediate forms occur along the eastern coast of Sakhalin and in the adjacent waters of the Okhotsk Sea. Specimens with angular shoulders and shell bases (figure 80) appear to be restricted to this region. Only there and, rarely, in the central and northeastern parts of the Okhotsk Sea do broadly fusiform shells with abruptK narrowing siphonal canals occur (figures 81, 82). When these populations are compared on the basis of a quantitative character (number of spiral ribs below the anal fasciole of the last whorl), the population from the Tatar Strait (figure 102, black dots) overlaps little with the population from Sakhalin Bay. However, specimens from eastern Sakhalin occupy an intermediate position, suggesting clinal variation. Specimens found along the Kurile Islands usually have pronounced spiral sculpture over the entire shell surface, including the anal fasciole. Dendritic ribs are generally absent in these populations. C;olor is light yellowish to yellowish brown. Specimens with ver\ shallow sinuses were lound only near the Kurile Islands (figure 1 top, left). Specimens from the Kurile Islands, Sakhalin Bay, eastern Kamchatka, and the Be- ring Sea have a similar number of the ribs on the body whorl (figure 103). Antiplanes sanctiioannis differs from A. obliquipli- cata n. sp., A. dendritupticata n. sp., A. abyssalis n. sp., and A. kawamtirai in having well developed spiral sculp- ture that is alwa> s present below the anal fasciole of the spire whorls. Antiplatics hurilensis n. sp. can be distin- guished from A. sanctiioannis by its higher spire (body whorl height/shell height ratio in A. sanctiioannis ranges from 0.542 to 0.667, mean value = 0.615 ± 0.004, while in A. kurilensis it varies from 0.500 to 0.533, mean value = 0.515 ± 0.005) and less elongate shell (shell diameter/ shell height ratio for A. kurilensis ranges from 0.310 to 0.380, mean value = 0.346 ± 0.008, while in A. sanc- tiioannis it varies between 0.360 and 0.477, with a mean value of 0.420 ± 0.003). Features that distinguish A. sanctiioannis from A. spirinae n. sp. include the presence of spiral ribs with secondary riblets, as well as propor- tionally longer marginal teeth, which are evident when comparing equal-sized specimens (figure 18). Antiplanes sanctiioannis differs from A. isaotakii. A. motojimai, and A. habei n. sp. in having more numerous spiral ribs (figure 138), and ontogenetically more slowly elongating marginal teeth. Distribution: This widespread boreal species ranges from Sagami Bay (Honshu) to the Gulf of Anadyr in the northwestern Bering Sea, and has also been collected in the Sea of Japan and the Sea of Okhotsk. The bath\ nietric range of this species extends from 50 m (southern Sa- khalin) to 1,530 m (southern Kurile Islands). Most spec- imens are from depths of 100 to 300 m (figure 100). Ai}tipla7ies kurilensis new species (figures 6, 89-93, 104) Material examined: Holotype (ZM N Lc 6903): R/V Gydrobiolog' Iturup Island (Kurile Islands), depth 55- 125 m, 1982; paratypes: R/\' 'Gvdrobiolog'. Kurile Is- lands, 1982 (1 shell— ZM N Lc 6904); R/V "Lebed", sta. Ill, E off Shumshu, northern Kurile Islands, 50°46.9'N, 157°13.4'E, depth 103-104 m, muddy sand, 22. VII. 1954 (1 specimen— ZIN N 1/54583); R/V 'Vityaz', 5th cruise, sta. 529, off Medny Island, Commander Islands, Bering Sea, 54°25.0'N, 168°16.4'E, depth 110 m, Sigsbee trawl, 14. VIII. 1950 (5 specimens— ZM N Lc 6907); Expedition of the Institute of Marine Biology of the USSR Academy of Sciences, sta. 42, off Moneron Island, Tatar Strait, 46°16.7'N, 141°10.6'E, depth 115 m, 30.VI1I.1972 (1 subfossil specimen — ZIN N 2/54584). Description of holotype: Shell elongate-fusiform, with very high spire consisting of 10 whorls. Surface of first 3 whorls eroded. Body w horl moderately convex, com- prising 0.51 of shell height. Shell light-brown, covered with thin, smooth, partly eroded periostracum. Teleo- conch whorls evenly convex, enlarge slowly. Suture shal- lowly impressed, nearly canaliculate. Incremental growth lines thin, distinct, producing finely rugose appearance on shell surface. Spiral sculpture of narrow, non-den- dritic, slightly wavy cords, 25 between sutures on spire whorls, about 50 on body whorl below anal fasciole. Chords evenly developed over entire shell surface. Surface of cords with pronounced growth lines. Base of shell Y. I. Kantor and A. V. Svsoev, 1991 Page 137 weakly convex, gradually passing into short, straight, obliquel) truncated siphonal canal. Aperture oval, nar- row. Inner lip slightly curved, covered with very thin callus. Callus better developed on columella. Aperture comprises 0.35 of shell height. Dimensions of hololype: Shell height 54.6 mm, body whorl height 27.7 mm, aperture height 19.2 mm, shell diameter 17.6 mm. Remarks: Paratypes have less conspicuous spiral sculp- ture, especially on the upper part of the whorls, and less numerous spiral ribs below the anal fasciole of the body whorl (<40 in paratype with shell height of 52.3 mm). Weak dendritic threads may be present on or above the anal fasciole. Only a single specimen from off Shumshu Island had moderately developed dendritic threads on the anal fascicle. The radula (figure 6) of a paratype (shell height 28.8 mm) was studied. Marginal teeth were 0 221 mm long (shell height/marginal tooth length = 130.3). Antiplanes kurilensis is most closely related to A. sanc- tiioannis, but differs in having a higher spire (body whorl height/shell height = 0.500-0.533, mean value 0.515 ± 0.005 for A. kurilensis, 0.542-0.667, mean value 0.615 ± 0.004 for A. sanctiioannis). Distribution: This species occurs in the northern and southern Kurile Islands and off Medny Island at 55-125 m. One subfossil specimen was found near Moneron Is- land (figure 104). A specimen resembling this new species conchologically was found in Pliocene deposits of north- eastern Honshu, and was referred to Antiplanes (Rec- tiplanes) sadoensis (Yokoyama) by Chinzei (1959:pl. X, figs. 6-7, not figs. 8-9). Antiplanes spirinae new species (figures 4, 22, 94-99, 104) Material examined: Holotype (ZIN N 1/54585): R/V 'Lebed', sta. 122, off Shumshu Island, northern Kurile Islands, 50°32.5'N, 157°27'E, depth 280 m, sand, gravel 1. VIII. 1954; paratypes: R/V "Odyssey", 34th cruise, sta. 47, off Simushir, central Kurile Islands, 46°44.9'N, 155°2I.0'E, depth 450-480 m, 30.XII.1984 (3 speci- mens—ZM N Lc 6905, shell heights 22.4; 16.3; 14.2 mm). Description of holotype: Shell thin, high spired con- sisting of 9 preserved whorls. Two earliest whorls lost. Teleoconch whorls evenly rounded, weakly convex, wid- est below shoulder. Sutures shallow. Bod\ whorl rather convex, comprising about Vz shell height. Coiling axis curved. Shell dark olivaceous; shell base, portions of anal fasciole lighter. Periostracum thin, finely granulated, peeling. Growth lines very thin, often darker than shell surface. Spiral sculpture on anal fasciole of 3-4 wide, unequal spiral cords somewhat broken by growth lines. Spiral cords less conspicuous toward body whorl. Den- dritic threads stronger on anal fasciole, but generally much weaker than in other species of Antiplanes. Re- maining surface of shell whorls covered by wide, flat- tened cords that become less conspicuous in later whorls, nearly absent on body whorl. Entire shell surface covered by microscopic secondary spiral threads, most conspic- uously on shell base (figure 99). Shell base weakly convex, joining short siphonal canal, broken at tip. Aperture short, narrow, oval Outer lip joining columella at acute angle. Parietal wall, columella covered by narrow, thin, white callus. Dimensions of holotype: Shell height 36.5 mm, body whorl height 17.9 mm, aperture height 12.2 mm, shell diameter 12.0 mm. Remarks: The paratypes have flatter whorls and less developed spiral cords on the shell surface and on the anal fasciole. The secondary threads are better developed and dendritic threads are practically absent in the para- types. Radulae of two paratypes were studied (figure 4). The mean length of the marginal teeth of the specimen with a shell length of 16.3 mm was 0.166 mm (shell height/ marginal tooth length length = 98.2). The corresponding values for the 22.4 mm specimen are 0.169 mm and 132.5. Antiplanes spirinae is most similar to A. sanctiioannis, but differs in having a narrower shell with a higher spire and less convex whorls, as well as spiral sculpture of poorly developed spiral cords and well developed sec- ondary threads. The marginal teeth of A. spirirtae are generalK shorter than in A. sanctiioannis (figure 18, specimens from the same localities connected by a dotted line). Etymology: This species is named in honor of Dr. N. Spirina, who collected the holot\pe. Distribution: This species is known from the northern and central Kurile Islands at depths of 280-480 m (figure 104). Antiplanes vinosa (Dall, 1874) (figures 9, 10, 24, 29, 105-120) Pleurotonia vinosa Dall, 1S74:.553. Pleurotoma (Antiplanes) vinosa Dall, 1902:514, pi. 34, fig. 4. Antiplanes kamchatica Dall, 1919:33-34, pi. 10, fig. 1. Antiplanes eontraria Yokoyama, Kira, 1965:100, pi. 36, fig. 2. Type specimens: Holotype of Pleurotoma vinosa, USNM 220899, shell height 36 mm. Type locality — Kyska Har- bour, Great Kyska Island, Aleutian Islands, 10 fms. Holotype of Antiplanes kamchatica, USNM 225255, shell height 52 mm. Type localit\ — south-eastern coast of Kamchatka, 100 fms. Material examined: 40 samples (more than 150 speci- mens) were studied. Description: Shell sinistral, variable, usually with high spire. Color dark-brown to light yellow. Body whorl com- prises 0.53 to 0.61 of shell height. Shell axis often curved. Whorls weakly to moderately convex, evenK rounded. Growth lines inconspicuous. Spiral sculpture of numerous Page 138 THE NAUTILUS, Vol. 105, No. 4 Y. I. Kantor and A. V. Sysoev, 1991 Page 139 Figure 117. Comparison of the number of spiral ribs on tfie body wfiorl below the anal fascicle in Sakhalin Bay populations of Antiplanes vinosa (Dall) (solid circles, right regression line) and A. sanctiioannis (Smith) (open circles, left regression line). The shell height along the abscissa, the number of spiral ribs along the ordinate variably developed wavy spiral cords spanning shell sur- face, most pronounced on siphonal canal. Cords some- times indistinct on and above anal fasciole. Dendritic threads variable in development, usually present on the anal fasciole, but not reaching suture on spire whorls. Aperture narrow, oval, usually well-differentiated from long, straight siphonal canal. Radula medium sized for genus, shell height/mean marginal tooth length varies from 111.6 (shell height 21.2 mm) to 158.4 (shell height 48.8 mm). Increase in mar- ginal tooth length with shell growth similar to that of A. sanctiioannis; regression equation: Y = 0.0942 -I- 0.0043X (figure 18). Figure 118. Comparison of the number of spiral ribs on the body whorl below the anal fasciole in the Kurile populations of Antiplanes vii^osa (Dall) (solid circles, 2 — line of regression) and A. sanctiioannis (Smith) (open circles, 1 — line of regres- sion). The shell height along the abscissa, the number of spiral ribs along the ordinate. Remarks: Antiplanes vinosa is the only sinistral species of Antiplanes in the northwestern Pacific Ocean. Anti- planes sanctiioannis is the most closely related dextral species. The number of spiral cords below the anal fas- ciole on the body whorl of A. vinosa is greater than in A. sanctiioannis (figure 117). Moreover, the spire of A. vinosa is generally higher than that of A. sanctiioannis, even in sympatric specimens (figure 119). In Sakhalin Bay, the ratio body whorl height/shell height is 0.57- 0.62 (mean 0.589 ± 0.006) for A. sanctiioannis. and 0.53-0.57 (mean 0.553 ± 0.006) for A. vinosa. Regression equations are Y = 0.5852 - 0.0007X and Y = 0.6346 - Figure§ 105-116. Shells of Antiplanes vinosa (Dall, 1874). 105. Hoiotype of Plcuwtoma vvwsa Dall, 1874, USNM 220899, 36 mm 106. Hoiotype of Antiplanes kamchatica Dall, 1919, USNM 925255, 52 mm. 107. Sakhalin Bay, Okhotsk Sea, 45 2 mm. 108. Tatar Strait, 28 5 mm 109. Central Kurile Islands, 51 7 mm 110. Off Moneron Island, Tatar Strait, 42 0 mm 111. Off Paramushir, northern Kurile Islands, 46.5 mm. 112. Northern Okhotsk Sea, 53.7 mm. 113. Off Shikotan, southern Kurile Islands, 40,9 mm. 114. Eastern Kamchatka, .35 3 mm. 115. Off Onekotan Island, central Kurile Islands, 29.7 mm 116. Southern Kurile Islands, 31.3 mm. Page 140 THE NAUTILUS, Vol. 105, No. 4 (mm) 60-1 50 (0 30 20 0.50 .52 .54 .56 .58 .60 .62 % Figure 119. Comparison of the relative height of the body- whorl of Antiplanes vinosa (Dall) (solid circles, 2 — line of re- gression) and A. sanctiioannis (Smith) (open circles, 1 — line of regression). The shell height along the abscissa, the ratio be- tween the body whorl height and the shell height along the ordinate. 0.00 13X respectively, where Y is the ratio of body whorl height to shell height and X is the shell height. Distribution: Antiplanes vinosa was recorded from the northeastern part of Honshu (Kira, 1965), the northern Japan Sea to the northern Okhotsk Sea, and from the Bering Sea (figure 120). Antiplanes isaotakii (Habe, 1958) (figures 5, 121, 123-125, 137, 138) Rectiplanes {Rectisulcm) isaotakii Habe, 1958:182, 185-186, text. fig. 2; Okutaiii 1964:424. Type specimens: Holotype, NSMT 38591, shell height 28.5 mm; paratype, NSMT 52684. Type locality, of Cho- shi, Chiba Pref., Honshu. Material examined: Off Kurile Islands (1 specimen); off southern Kurile Islands (1 specimen). Description: Shell light-olivaceous, elongate, high spired. Body whorl weakly convex, comprising 0.57-0.62 shell height. Spire whorls weakly to moderately convex, slight- ly angulated at periphery, flattened at anal fasciole. Su- ture shallow. Siphonal canal long, slightly curved. Ap- erture narrow, short, comprising 0.44-0.52 of shell height. Outer lip joining columella at nearly right angle. Spiral sculpture well developed, comprised of 2-3 flattened to convex cords between suture and anal fasciole, <5 cords on fasciole, sometimes with traces of dendritic threads, 20 wide, rounded cords, uniformly developed over entire body whorl below anal fasciole. Earlier whorls with 4- 5 cords below anal fasciole. Radula of 30.8 mm specimen was studied (figure 5). Marginal teeth were large (mean length = 0.289 mm, shell height/tooth length ratio = 106.6) (figure 18). Remarks: Antiplanes isaotakii is conchologically sim- ilar to some forms of A. sanctiioannis, but differs in having longer marginal teeth and fewer spiral cords that are more convex, wider, and more evenly spaced. It differs from A. habei in having fewer spiral cords and a more acutely angled periphery. Distribution: Antiplanes isaotakii was recorded off Chiosi, Honshu, in Sagami-Bay and off the southern Ku- rile Islands in 200-780 m (figure 137). Antiplanes motojimai motojimai (Habe, 1958) (figures 122, 137) Rectiplanes (Rectisutciis) motojimai Habe, 1958:182-185, fig. 3; Okutani, 1964:424. pi. 4, fig, 7, Type specimens: Holotvpe, NSMT 30600, shell height 38.2 mm; paratypes, NSMT 30600a-b, shell heights 37.0, 35.5 mm. Type locality — off Chiosi, Chiba Prefecture, Honshu, Japan. Description: "Shell elongate with about eight whorls which are moderately inflated but deeply constricted at the suture, covered with the olivaceous periostracum; surface sculptured with the spiral cords, two or three of which are contained in the area between the rather broad fasciole and the upper suture and also two or three in the area between the fasciole and the lower suture; body whorl very large and stout, occupying about two-thirds of the shell length, the periphery of which is rounded and the base is marked by the spiral cords all over; aperture wide, the outer margin broadly sinuated in V-shape at a little above the periphery; canal broad and short; columellar margin white." (Habe, 1958:184). Remarks: This species is most similar to A. isaotakii. but differs in having a broader shell and fewer spiral cords (21 cords below the anal fasciole on the body whorl of the holotype) that become narrower towards the si- phonal canal. Distribution: This subspecies occurs from Sagami Bay to the Sea of Kashima Nada (eastern Honshu), in 550- 870 m (Okutani, 1964). It is not represented in collections of USSR institutions. Antiplanes motojimai aquilonalis new subspecies (figures 126, 127, 137) Material examined: Holotvpe (ZM N Lc 6906): R/V 'Adler', 18th cruise, sta. 169, eastern Bering Sea, 62°59'N, Y, I. Kantor and A. V. Sysoev, 1991 Page 141 Figure 120. (Jeographic distribution of Antiplanes vinosa (Dall). 171°05'W, depth 46 m, 1983; paratypes: R/V 'Vityaz', 2nd cruise, sta. 20. southern Okhotsk Sea, 45°01.1'N, 144°21.0'E, depth 429-404 m, muddv sand and gravel, 10. VIII. 1949 (empty shell— ZM N Lc 6908); R/V 'Aka- demik Oparin', 7th cruise, sta. 17, north-eastern Okhotsk Sea, 56°00'N, 146°29'E, depth 397 m, 1988 (empty shell— ZIN N 1/54624). Description of holotype: Shell broad, consisting of 6 whorls, with earliest 1.5 whorls lost, next 3 whorls eroded. Spire whorls evenly convex, with narrow, flattened sub- sutural band above anal fasciole. Suture deeply im- pressed. Shell color dark olive. Periostracum thick, glossy. Growth lines numerous, some raised, rough, forming raised flat tubercles along lower parts of whorls. Spiral sculpture well developed, with 4-5 distinct, flattened ribs below anal fasciole on spire whorls. Body whorl with two narrow cords adjacent to anal fasciole, next 4 cords 1.5 times as broad, remaining 10 cords becoming progres- sively narrower towards siphonal canal. Cords flat, with serrated borders at intersections with growth lines. Anal fasciole with 2-3 thin threads, separated by wide inter- spaces. Dendritic threads well developed, reaching su- ture on spire whorls, disappearing on shell base. Anal fasciole with very thin secondary threads, especially on spire whorls. Siphonal canal broken. Shell base eroded. Aperture narrow, oval. Dimensions of holotype: Shell height 35.5 mm, body whorl height 21.6 mm, aperture height 14.2 mm, shell diameter 16.5 mm. Remarks: Paratypes are very similar to the holotype, but differ in having three ribs above the anal fasciole. This new subspecies differs from the nominatypical one in spiral sculpture: while Antiplanes motojimai moto- jimai has equally developed cords on the periphery of the whorls and on the anal fasciole, Antiplanes moto- jimai aquilonalis lacks wide ribs on the fasciole. These two subspecies differ in their geographical distributions. Distribution: This new subspecies was recorded in the Okhotsk and Bering Seas at depths from 46 m (Bering Sea) to 550 m (southern Okhotsk Sea) (Fig. 137). Etymology: aquilonalis Latin — northern, from Aquilo Latin — northern wind. Antiplanes habei new species (figures 7, 8, 23, 128-135, 137, 138) Material examined: Holotype (ZM N Lc 6909): R/V 'Vityaz', second cruise, sta. 100, central part of the Okhotsk PUL THE NAUTILUS, Vol. 105, No. 4 Figures 121, 122. Shells of A;!np/(i«« spp. 121. Rectiplanes isautakii liahe. 19.58. holot\pe, NSMT \ .526S4. 28..5 mm. 122. Rectiplanes (Rectisulcus) molojimai Habe, 1958, holotype, NSMT N 38600, 38.2 mm. Sea, 54°35.0'N, 149°49.5'E, depth 543 m; paratypes: R/V "Vityaz', 12th cruise, sta. 1853, central part of the Okhotsk Sea,' 55°38,5'N, 143°01.0'E, depth 389 m, mud (1 spec- imen—ZM N Lc 6910); R/V "Adler", 18th cruise, sta. 42, Okhotsk Sea, 49°12.7'N, 144°57.6'E, depth 245-260 m (1 specimen— ZM N Lc 6911); R/V 'Gagara', sta. 217, Okhotsk Sea, 51°35'N, 154°46'E, depth 410 m, (1 spec- imen—ZIN N 3/54625); R/V 'Gagara', sta. 252, Okhotsk Sea, 55°32.4'N, 149°14'E, depth 335 m (3 specimens— ZIN N 2/54626); R/V 'Toporok', sta. 17, off Shikotan, 43°32.5'N, 147°20.5'E, depth 1,450-1,530 m (2 speci- mens—ZIN N 1/54627). Description of holotype: Shell elongate-fusiform, with high spire and high, weakly convex body whorl that comprises 0.57 of shell height. Shell of seven whorls, with at least one upper whorl lost. First preserved whorl erod- ed. Shell yellowish, with darker spiral band below anal fasciole. Periostracum thin, gloss>. Whorls weakK con- vex, with flattened periphery nearly parallel to shell axis. Sutures impressed, shallow. Incremental growth striae thin, inconspicuous, some marked by color. Spiral sculp- ture of inconspicuous dendritic thread-like riblets on anal fasciole (4-5 on penultimate whorl, 3 on body whorl) reaching sutures on spire whorls and disappearing at periphery of body whorl. Ribs, 2-3 above fasciole, 4-5 below fasciole on three upper whorls, up to 9 on other spire whorls, 31 below fasciole on body whorl, becoming more distinct toward siphonal canal. Five ribs below anal fasciole flattened and of equal width, subsequent ribs more narrow and raised. Shell base flattened, joining short siphonal canal with distinct bend. Aperture narrow, short, comprising 0.43 of shell height. Outer lip joins columella at obtuse angle. Callus thin, narrow, white, glossy, covers columella and parietal wall. Siphonal canal poorK' differentiated from aperture. Dimensions: Holot\pe, shell height 38.0 mm, body whorl height 21.6 mm, aperture height is 16.3 mm, diameter 13.6 mm. Remarks: Paratypes have slightly more convex and rounded whorls, wider apertures, more distinct spiral ribs below the anal fasciole, and wide, flat ribs on the fasciole that are absent on the body whorl. Limited material does not allow an evaluation of the range of intraspecific vari- ability. We provisionally refer to this species two spec- imens from off Shikotan that differ from other paratypes in having; rounded or angulated, but never flattened whorls, a longer siphonal canal, high anal sinus apex, different geographic (southern Kuriles rather than the Okhotsk Sea) and bathymetric (1,450-1,530 m vs. 335- 575 m) distributions. The radulae of four specimens were studied (figures 7, 8). The marginal teeth are large; the ratio between the shell height and the tooth length varies from 76.3 (the shell height 14.2 mm) to 118.9 (37.8 mm). Their length rapidly increases with shell growth according to the regression equation: Figures 12,3-136. Sht-lis of Antiplanes spp 123-12.5. Antiplanes isaotahii (Habe. 1958). 123. 124. ■' Kurilt- Islands, 31 0 mm 125. Soiitlu-rii Kurik- Islands. 21.3 mm. 126, 127. Anliplunes ruotojimai aqtiilonalis n. subsp. holotype, ZM N Lc 6906, 35.5 mm 128-135. Antiplanes hahci n sp. 128. llolotvpe, ZM N Lc 6909, 38 0 mm 129. 1.30. Paratvpe. Okhotsk Sea, ZM N Lc 6911. 29 0 mm. 131, 132. Central part of Okhotsk Sea, ZM N Lc 6910. 23 0 mm 133-135. Off Shikotan. Kurile Islands, ZIN N 1/54627 (133—33.5 mm, 134, 135—23.2 mm). 136. Antiplanes IniHmoides Dall. 1919 Holot\pe, 31 mm. Y. I. Kantor and A. V. Svsoev, 1991 Page 143 Page 144 THE NAUTILUS, Vol. 105, No. 4 Figure 137. Geographic distribution of Antiplanes isaotakii (Habe) (solid circles), A. motojimai motojimai (Habe) (open circles) A. motojimai aqiiilonalis n. subsp. (solid squares), and A. hahei n. sp. (open circles). Y = 0.1290 + 0.0052X. Antiplanes hahei is most similar to A. sanctiioannis, but differs in having more convex, wider and less numerous ribs (figure 138) that are nearly equal over the body whorl. The ribs of A. sanctiioannis are considerably more pronounced on the shell base and the siphonal canal. This new species has large marginal teeth (figure 18), which is most clearly seen when comparing equal-sized specimens from the same locality (the respective points are connected by dotted line in figure 18). Antiplanes hahei also resembles A. isaotakii, but differs from it in having a deeper suture, more convex whorls, more nu- merous ribs on the body whorl, as well as in lacking the angular periphery. Distribution: Antiplanes hahei was recorded in the Okhotsk Sea and the southern Kurile Islands at depths from 335 m (central part of the Okhotsk Sea) to 1,530 m (off Shikotan) (figure 137). Etymology: This new species is named in honor of Dr. Tadashige Habe, who has considerably widened our knowledge of the north-western Pacific species of An- tiplanes. Antiplanes hulimoides Dall, 1919 (figure 136) Dall. 1919:.34, pi II. fig, 7; Dall, 192,5:4, pi .31, fig. 2. Material examined: Holotype (USNM 111051), U.S. Bureau of Fisheries, Station 4772 on Bowers Bank, Bering Sea, 54°30'30"N, 179°13'00"E, in 344 fathoms, green- brown sand, bottom temperature 38.1°F, 6. IV. 1906. Description: "Shell elongate, decollate. Whorls six or more, four remaining. Suture distinct, not appressed. Whorls moderateK convex, smooth, with pale polished greenish periostracum, in spots minutely granulose, ap- parently from some wrinkling of the periostracum. Anal sulcus wide, shallow, bareK forming a fasciole. Outer lip thin, sharp, moderately produced. Inner lip with a thin white layer of callus. Columella straight, with an oblique anterior attenuation. Canal wide, hardly differentiated. Length of three complete whorls, 31 mm. Length of last whorl, 23 nun. Diameter at apex, 5 mm. Maximum shell diameter, 15 mm." Remarks: So far as we know , this species is known only from the holotype, which is figured and described here. Y. I. Kantor and A. V. Sysoev, 1991 Page 145 (mmf ^5 M) 35 30 25 20- 20 30 i.0 50 K Figure 138. Comparison of the number of spiral ribs on the body whorl below the anal fascicle in populations of Antiplanes isaotakii (Habe) (open squares), A. habei n. sp. (solid circles, 1 — line of regression), and A. sanctiioannis (Smith) (open cir- cles, 2 — line of regression). The shell height along the abscissa, the number of spiral ribs along the ordinate. The bulimoid shape and gUstening greenish periostra- cum makes this species quite unique. LITERATURE CITED Addicott, U. O. 1976, Molluscan paleontology of the Lower Miocene Clallam Formation, north-western Washington, U.S. Geological Survey, Professional Paper 976:1-44, Amano, K, 1983. Paleontological study of the Pliocene To- geshita molluscan fauna in the Rumoi district, Hokkaido. Science Reports of the Institute of Geoscience, University of Tsukuba, Section B, Geological Sciences 4:1-72. Aurivillius, C. W. S. 1887. Ofversigt ofver de af Vega ex- peditionen insamlade. Apktiska Hafsmolluker. II. Placo- phora och Gastropoda. In. Nordenskold, A. E. (ed.). Vega- Expeditionens Vetenskapliga lakttagelser 4:311-383. Bartsch, P. 1944a. Some notes upon west American Turrid molluscs. Proceedings of the Biological Society of Wash- ington .57:2.5-30. Bartsch, P. 1944b Some Turrid molluscs of Monterey Bay and vicinity Proceedings of the Biological Society of Washington 57:57-68. Berry, S. S. 1953. Notices of new West American marine Mollusca. Transactions of San Diego Society of Natural History 1 1(16):405-428. Chinzei, K 1959 Molluscan fauna of the Pliocene. Sannohe Group of northern Honshu, Japan 1. The faunule of the Kuba Formation. Journal of the Faculty of Science, Uni- versity of Tokyo. Section II, Geology, Mineralogv, Geog- raphy, Geophysics, 12(1): 103-132, "pis. IX-XI. Coan, E. \' and A. E Bngan 1988. The Recent invertebrate taxa described by William More Gabb. 1839-1878 Pro- ceedings of the .Acadenn of Natural Sciences of Phila- delphia 140(l):273-284. ' Dall, W. H. 1874. Catalogue of shells from Bering Strait and the adjacent portions of the Arctic Ocean, with descriptions of three new species. Proceedings of the California Acad- emy of Natural Sciences 5:246-253. Dall, W. H. 1902. Illustration and descriptions of new, un- figured, or imperfectly know n shells, chiefly American, in the I'.S. National Museum. Proceedings of the Ll.S. Na- tional Museum 24 (1264):499-566. Dall, W. H. 1919. Descriptions of new species of molluscs of the family Turritidae from the west coast of America and adjacent regions. Proceedingsof theU.S. National Museum 56(2288): 1-86. Dall, W. H 1925 Illustrations of unfigured types of shells in the collections of the L'nited States -National Museum. Pro- ceedings of the I' S. National Museum, 66, art. 17. (2554): 1-41. " Donati, G., S. Gargiulo, and B. Porfirio. 1984. Finding of 11 sinistral specimens of Contis mediterraneus. La Conchig- lia 182-183:21-23. Gabb, W M 1865. Description of new species of marine shells from the coast of California Proceedings of the California Academy of Natural Sciences 3:102-190. Gabb, W. M. 1869. Cretaceous and Tertiary fossils. Descrip- tions of the Tertiary Invertebrate fossils. Geological Survey of California [Paleontology of California] 2(1,1):1038. Golikov, A, N, and V. V. Gulbin, 1977. Prosobranchiate gas- tropods (Gastropoda, Prosobranchiata) of the shelf of Ku- rile Islands. II. Orders Hamiglossa-Homoestropha. In: Fau- na of the offshore zones of Kurile Islands. Nauka, Moscow, p. 172-268. Golikov, A. N. and O. A. Scarlato. 1985, Shell-bearing gas- tropods and bivaKes of the shelf of south Sakhalin and their ecolog\ . In. Biocenosisand fauna of the shelf of south Sakhalin. Investigations of the fauna of the seas 30(38): 360-487. Grant, U. S. and H R Gale 1931 Catalogue of the marine Pliocene and Pleistocene mollusca of California and ad- jacent regions. Memoirs of the San Diego Society of Nat- ural History 1:477-612. Habe, T 1958. Description of three new species of the genus Rectiplanes from Japan (Turridae, Gastropoda). Venus 20(2):181-186. Hickman, C. S. 1976. Bathyal gastropods of the family Tur- ridae in the Early Oligocene Keasey Formation with a review of some deep-water genera in the Paleogene of the Eastern Pacific. Bulletins of American Paleontologv 70(292): 1-119. ICantor, Yu. I 1990. Gastropods of the subfamily Volutopsi- inae of the World Ocean. Nauka, Moscow, 178 p. (In Russian). Kira, T. 1965. Shells of the Western Pacific in color \'ol 1 Osaka, Japan, 224 p Maes, V O 1983 Observations on the systemalics and biology of a turrid assemblage in the British N'irgin Islands. Bulletin of Marine Sciences 33:305-335. Masuda, K and H. Noda. 1976 Checklist and bibliographv Page 146 THE NAUTILUS, Vol. 105, No. 4 of the Tertiary and Quarternary Mollusca of Japan, 1950- 1974. Saito Gratitude Foundation, Sendai 494 p. McLean, J. H. 1971. A revised classification of the family Turridae, with the proposal of new subfamilies, genera, and subgenera from the Eastern Pacific. The V'eliger, 14(1): 89-130. Okutani, T. 1964 Report on the archibenthal and abyssal gastropod Mollusca mainly collected from Sagami Bay and adjacent waters by the R/V Soyo-Maru during the years 1955-196.3. Journal of Faculty of Sciences, University of Tokyo, section II, 5(3):371-44'7. Okutani, T. 1968. Bathyal and abyssal mollusca trawled from Sagami Bay and the south off Boso peninsula by the R/V Soyo-Maru, 1965-1967 Bulletin of Tokai Regional Fish- eries Research Laborators 56:7-55. Otuka, Y 1949. Fossil Mollusca and rocks of the Kiyosumi Group exposed at Minato Machi, Chiba Prefecture and its environs. Japanese Journal of Geology and Geography 21(14):291-309. Petrov, (). M. 1982. Marine molluscs of the Anthropogene from the northern regions of the Pacific. Nauka, Moscow, 142 p. Powell, \. W B 1966 The molluscan families Speightiidae and Turridae Bulletin of the .Auckland Institute and Mu- seum 5:1-184. Shikama, T. 1962. On some noteworthy shells from off Choshi, Chiba Prefecture. Science Reports of the Yokohama Na- tional University. Section 2, Biological and Geological Sci- ences 8:29-56. Shikama, T. 1977. Description of new and noteworthy Gas- tropoda from the western Pacific and Indian oceans. Sci- ence Reports of the Yokohama National L'niversity. Sec- tion 2, Biological and Geological Sciences 21 9-24. Smith, E. A. 1875 A list of gastropoda collected in Japanese seas by Commander H. C. St. John, R. N. .\nnals and Mag- azine of Natural History, series 4, 15:414-427 Tippett, D. L. 1983. A new sinistral turrid from Brazil (Gas- tropoda: Turridae), Nautilus 97(4): 135-138. THE NAUTILUS 105(4): 147-151, 1991 Page 147 Two New Species of Vexillum from the Western Pacific (Gastropoda, Costellariidae) Gary Rosenberg Academy of Natural Sciences 19th and the Parkway Philadelphia, PA 19103 Richard Salisbury 8807 Craydon Drive Boise, ID 83704 ABSTRACT Vexillum brunneolinea n sp. is described from the Palau Is- lands and compared to Vexillum acuminatum (Gmelin, 1791) and V. semisculptum (Adams & Reeve, 1850)^ It is the first member of its genus reported in which some individuals lack apertural lirations. Vexillum elliscrossi n sp is described from Hawaii and central Pacific guyots and compared to Vexillum daedalum (Reeve, 1845) and Vexillum xenium Pilsbry, 1921. Key words: Vexillum. Costellariidae; Hawaiian Islands; Palau Islands; INTRODUCTION We first became aware of the two Vexillum species de- scribed in this paper more than twelve years ago. Our research in the literature at intervals during those years, involving study of the original descriptions of more than two thousand living and fossil species of miters (mitrids and costellariids), failed to uncover names for these taxa, although during that period both species were illustrated under erroneous names. We have not assigned the species we describe to subgenera because the limits of the sub- genera of Vexillum are poorly defined and will surely change as anatomical knowledge of the group increases. On the basis of shell characters, it is possible that Vex- illum brunneolinea n. sp. will be assigned to the subgenus Costellaria, and V. elliscrossi n. sp. to the subgenus Pusia. SYSTEMATIC DESCRIPTIONS Family Costellariidae MacDorald, 1860 Genus Vexillum Roding, 1798 Vexillum brunneolinea new species (figures 1-3) Vexillum (Costellaria) acuminaturr^ i politum "Reeve" Tur- ner, 1989, pp. 12, 28, pi. 5, fig. 21, right hand specimen only; not Mitra politum Reeve, 1844 Description: Adult shell length ranges from 17.4 to 25.5 mm; width ranges from 28 to 33 percent of length, and spire height from 37 to 42 percent of shell length. A summary of measurements is given in table 1. The shell is white with 29 to 57 fine brov\ n axial lines visible through a pale yellow periostracum. In live-collected specimens the periostracum is greenish-yellow and opaque. The green color may be due to the presence of commensal algae, as it rapidly fades after collecting. The protoconch and first three or four teleoconch whorls are dark rust in color. The protoconch is smooth, about 0.5 mm in diameter and consists of about 2.2 whorls; the teleoconch consists of 8.9 to 10.5 convex whorls. The upper whorls have 12 to 20 axial ribs per whorl, typically 13 to 16. These ribs become obsolete by the penultimate whorl, which is smooth. On the upper three teleoconch whorls, there are 3 or 4 spiral grooves, which in some cases increase to five in number on succeeding whorls. These grooves are visible as pits between the axial ribs, but do not cross the ribs; they usually become obsolete by the penultimate whorl, but are still faintly visible on the body whorl in some specimens. There are three to seven spiral cords above the fasciolar ridge, and four to six below it on the siphonal fasciole. In some specimens the cords on the fasciole are obsolete or absent. The aperture and columella are white. There are four pli- cations on the columella; the strongest is the posterior- most, which ranges up to 0.40 mm in width distalK' in adult specimens. There are 0-13 apertural lirations. In some specimens the lirations are of normal strength for Vexillum, in others they are shortened, reduced in num- ber or in length, or, in seven specimens, are entirely absent. To make sure that lirations were lacking, a high intensity light was shone through the body whorl opposite the aperture. This procedure shows any trace of a liration in high relief, and none were detected in these specimens. The lack of apertural lirations has not been documented in any other Vexillum species. The radula of the holotype is preserved on a slide prepared at the Academy of Natural Sciences (ANSP) between 1955 and 1958 by the late Virginia Orr Maes; it is typical of Vexillum species as illustrated by Cer- nohorsky (1970). The radula is fragmentary, with only 26 rows of teeth represented. Its width is 0.14 mm. The spacing between rows is about 0.025 mm, so there would have been about 40 rows of teeth per millimeter of ribbon length. The rachidian teeth have eight to ten cusps; the Page 148 THE NAUTILUS, Vol. 105, No. 4 Figures 1-3. Vexillum hrunneolinea new species. 1. Holotype, length 25.5 mm, ANSP 217492, with periostracum. 2. Paratype, length 21 mm. Salisbury collection, periostracum removed ,3. ,.\pertural detail of liolotype, aperture length 115 mm Figures 4, 5. Vexillum acuminatum (GmeHn, 1791) 4. Form polilum (Reeve, 1844), length 20.5 mm, ANSP 28635, Philippines. 5. Apertural detail of large individual, aperture length US. 3 mm, ANSP 29699. Sri Lanka Figures 6-9. Vexillum clli.scrossi new species 6. Holotype, length 18.5 mm, KPHM 219991. 7. Paratype, ape.x, coated with magnesium o.xide, IMT Sta. 73-15. 8. Same specimen G. Rosenberg and R. Salisbury, 1991 Page 149 lateral teeth are smooth. The rachidians are 0.08 mm in length, the laterals 0.06 mm; the ratio of their lengths is 1.33. Radular statistics are all within the ranges reported by Cernohorsky (1970:10) for Vexillttrn. Type material: HOLOTYPE. ANSP 217492 (radula slide 937), 25.5 X 7.5 mm. PARATYPES. ANSP 202887 (1 specimen); ANSP 203082 (3 specimens); ANSP 203341 (2 specimens); ANSP 203787 (2 specimens); ANSP 217471 (2 specimens, bodies in ethanol); ANSP 382159 (1 spec- imen). Salisbur\' Collection, 7 specimens. Paratypes total eighteen specimens. Type locality: Dredged in sand at 9 meters, inside the outer reef 12.9 km WNW of Koror Island, Palau Islands, Western Carolines. 7°24'N, 134°21'E. Collected by A. J. Ostheimer, 3rd; Station 441-6, 21 August 1955. Distribution: All known specimens have been taken in the Palau Islands. Data for ten paratypes at ANSP that were dredged by A. J. Ostheimer, 3rd in 1955 are as follows: ANSP 202887— Sta. 452. at 31 meters, 3.2 km NE of Gamudoko Island off Urukthapel Island, 24 August. 7°17.5'N, 134'=2I.5'E. ANSP 203082— Sta. 436, in sand at 27 meters, Malakal Harbor, 20 August. 7°19'N. 134°27'E. ANSP 203341, 217471— Sta. 361, in clean fine sand at 8 to 12 meters, in bay on inner edge of barrier reef 1.6 km south of West Passage, Babelthuap Island, 16 Julv. 7''31'N, 134°29'E. ANSP 203787— Sta. 450, at 33 meters, east of Yoo Pas- sage, 4.8 km NE of Eil Malk, 24 August. 7°12'N, 134°25.7'E. The other paratypes were collected by Dieter Cosman using SCUBA in April 1978 in sand at 5.5 meters in the rock islands around Koror. One of these is ANSP 382159, the other seven are in the Salisbury collection. Turner (1989, pi. 5, fig. 21, right-hand specimen) published a color photograph of a 24 mm specimen in the Ted Baer collection taken at 10-20 meters in sand and fine rubble in Palau. Etymology: "Brunneolinea" is derived from Latin brunneus (brown) and linea (line) referring to the brown axial lines characteristic of the species. It is a noun in apposition, and so does not change endings to match the gender of the genus. Discussion: This species was considered a variant of the poiitum form of Vexiltum acuminatum (Gmelin, 1791) by Turner (1989, p. 12). Vexillum brunneolinea is easily distinguished from typical Vexillum acuminatum as the latter is brown in color, has a light band below the suture and has axial ribs on the body and penultimate whorls. Vexillum acuminatum form poiitum (Reeve, 1844) (fig- Table 1. Measurements (mm) and counts of conchological features of Vexillum brunneolinea n. sp. Number of specimens varies because in some cases values for juvenile, damaged and worn specimens were omitted or could not be determined. Mean Range N Siiell length 21.3 17.;3-25.5 10 Bods whorl length 13.0 10.1-15.6 10 Bod\ whorl width 6.4 5.0-7.6 10 Spire length 8.3 7.1-9.9 10 Aperture length 9.5 7.3-11.5 10 .\perture width 2.3 1.7-2.8 10 Plica 1 width 0.32 0.20-0.40 10 Teleoconch whorls 9.6 8.9-10.5 10 Ribs, whorl 1 12.8 12-14 14 Ribs, whorl 2 13.6 12-16 19 Ribs, whorl 3 14.5 13-17 19 Ribs, whorl 4 14.9 13-18 19 Ribs, whorl 5 16.0 13-19 18 Ribs, whorl 6 15.9 13-20 17 Protoconch width 049 0.47-0.52 15 Protoconch whorls 2 2 2.1-2,4 6 ure 4) is smooth on these whorls, and sometimes develops a pattern of brown axial lines on a grayish-brown back- ground. However, these lines are broader and more wide- 1\ spaced than those in V. brunneolinea. The lines are darkly pigmented areas along growth lines of the shell, whereas the axial lines in V. brunneolinea are somewhat flexuous and in some cases bifurcate; they do not follow the growth lines. Another feature that separates these species is that the posteriormost columellar tooth in fully mature individ- uals of V. acuminatum is broader than that in V. brun- neolinea, and is often grooved at the top, giving it a biplicate appearance (figure 5). This tooth is convex in outline in V. brunneolinea. Vexillum acuminatum has been recorded from Palau, so V. brunneolinea cannot be considered a geographical variant of it. Some forms of Vexillum semisculptum (.Adams & Reeve, 1850) are also morphologically similar to V. brun- neolinea. The species is usually ribbed, but as in V. acuminatum, smooth variants are known. However, the coloration of V. semisculptum is a uniform greenish, bluish or grayish brown with a white line below the suture. The aperture is brown except for the white line visible there also. Its color pattern differs strikingly from that seen in V. brunneolinea. Vexillum elliscrossi new species (figures 6-9) Pusia daedala "Reeve," Kosuge, 1979, p. 27, pi. 6. fig. 28, not Mitra daedala Reeve, 1845. Pusia sp., Salisbury, 1981, p. 6, figs. 8.\-B (in color). as fig. 7, length 12.8 mm. 9. Paratype, length 17.2 mm, Salisbury collection Figure 10. Vexillum xcnium Pilsbry, 1921, holotype, length 17.8 mm, ANSP 116983. Note: All figures except 3, 5 and 7 are printed at the same magnification. Page 150 THE NAUTILUS, Vol. 105, No. 4 Table 2. Measurements (mmj and counts of conchological features of Vexillum elliscrossi n. sp. Number of specimens (N) varies because in some cases values for juvenile, damaged and worn specimens could not be determined. Mean Range N Shell length 17.4 16.9-18.5 Body whorl length 10.7 9.8-12.0 4 Bod\' whorl w idth 6.3 5.8-6.8 4 Spire length 6.3 6.2-6.4 3 Aperture length 8.6 7.7-9.5 4 Aperture width 2.5 2.5-2.7 4 .\pertural lirations 9.3 6-13 6 Plica 1 width 0.38 0.30-0.46 4 Teleoconch whorls 7.5 7.0-7.8 3 Ribs, whorl 1 13.7 12-15 3 Ribs, whorl 2 14.6 12-16 5 Ribs, whorl 3 15.0 12-16 5 Ribs, whorl 4 16.2 12-19 5 Ribs, whorl 5 17.4 14-22 5 Ribs, whorl 6 18.4 16-23 5 Ribs, whorl 7 20.0 18-23 3 Protoconch width 0.62 0 60-0.64 3 Protoconch whorls 44 44-45 2 Description: Shell length of adult specimens ranges from 16.9 to 18.5 mm; bod\' whorl width and spire height both range from 35 to 38 percent of shell length. A summary of measurements is given in table 2. The shell is white with four brown bands on the body whorl, two of which are visible on the spire whorls. The band below the suture is sometimes faint and is lightest near the suture; the band below it is the darkest of the four. Fresh specimens are suffused with lavender, which fades rapidly after collecting. The axial ribs are white and cut the brown bands into a series of tightly spaced blotches. The ap- erture and columella are white. All specimens studied have large, diagonal scars where breaks in the shell were repaired. The protoconch is amber in color and appears smooth under a light microscope. It is tall (0.90-0.95 mm), con- sisting of about 4.4 whorls. There are up to eight teleo- conch whorls. These are convex and slightly terraced. The number of ribs on the early teleoconch whorls ranges from 12 to 16, and increases to about 20 on the body whorl. Spiral grooves are visible between the ribs. These are faint on the first three teleoconch whorls and increase in strength towards the body whorl. From 7 to 9 grooves are visible on the spire whorls and 24 to 28 are visible on the body whorl. The grooves on the body whorl are stronger and more widely spaced along the siphonal ca- nal. There is no siphonal notch and consequently there is no fasciolar ridge or distinct siphonal fasciole. There are four plications on the columella; the strongest is the posteriormost, which is up to 0.5 mm wide. There are 6 to 13 apertural lirations. Type material: HOLOTYPE BPBM 219991, 18.5 x 6.8 mm. PARATYPES. IMT, 3 uncatalogued specimens; Salisbur) Collection, 1 specimen; Leonard Hill Collec- tion Cs-173, 1 specimen, figured by Salisbury (1981). Type locality: Off VVaikiki, Oahu, Hawaii, at 180 meters on a sand bottom, collected bv the "Pele Expedition" on 21 March 1965. Distribution: The three paratypes in IMT were cited bv Kosuge (1979), from stations 73-3 (29°47.0'N; 179°04.7'E), 73-4 (29°47.4'N; 179°02.9'E), and 73-15 (26°18.9'N; 174''30.9'W) on central Pacific guyots west of Midway Island. We did not examine a fourth speci- men, figured by Kosuge (pi. 6, fig. 28), from station 73- 25 (29°48.0'N; 179°01.3'E) as it was not included in the material loaned to us. The IMT specimens were dredged in March 1973 at depths from 67 to 267 m. The paratype in the Salisbury collection was dredged off Keehi Lagoon, Oahu by E. R. Cross. The paratype in the Hill collection was taken off Honolulu at about 90 meters in rock and coralline algal rubble. Etymology: We are pleased to name this species in honor of Ellis R. Cross, editor emeritus of the Hawaiian Shell News, who collected one of the paratypes. His work with deep-dredged shells has contributed greatly to knowledge of the molluscan fauna of the Hawaiian Is- lands. Discussion: Kosuge (1979) identified Vexillum ellis- crossi as Vexillum daedalum (Reeve, 1845), but it is easily distinguished from that species. Vexillum daeda- lum has a brown or ashy green shell with a single white peripheral band, whereas V. elliscrossi has a white shell with four brown bands. The aperture and columella are brown in V. daedalum and white in V. elliscrossi. The axial ribs on the body whorl in V. daedalum number 10 to 15 (Cernohorsky, 1972), whereas V. elliscrossi has 18 to 23 ribs. V. daedalum has a siphonal notch and fasciole whereas V. elliscrossi lacks the notch and fasciole. The shell that Kay (1979) illustrated from Hawaii as Vexillum sp. cf. rufofilosum (E. A. Smith, 1876) may be a deep water form of Vexillum daedalum. Cernohorsky (1972) and Turner (1989) considered V. rufofilosum to be syn- onymous with V. daedalum. Turner illustrated the ho- lotypeof V. rufofilosum. Vexillum oiiiscinum (Lamarck, 1811) might prove to be an older name for V. daedalum. Another species that occurs in Hawaii that might be confused with Vexillum elliscrossi is V. xenium Pilsbry, 1921 (figure 10). The holotype of V. elliscrossi was orig- inally catalogued at the Bishop Museum as "V. (C.) xeni- um?" but comparison to the holot\pe and paratype of V. xenium (ANSP 116983) shows that the species are easily distinguished. The apertural lirations in V. xenium are interrupted; in V. elliscrossi they are continuous. Vexillum xenium has a siphonal notch and fasciole; these are lacking in V. elliscrossi. The brown peripheral band in V. xenium is strongest on top of the spiral cords and lighter in the grooves. There are several narrow bands of brown on top of individual cords elsewhere on the whorl; these are darkest where the\ cross the ribs and lighter or absent in the interspaces. In contrast, the brown G. Rosenberg and R. Salisbury, 1991 Page 151 bands in V. elliscrossi do not cross the ribs, but cover the spiral cords and grooves uniformly, Vexillum xcniiim also has scattered brown subsutural blotches; these blotches are absent in V. elliscrossi. ACKNOWLEDGEMENTS We thank Dieter Cosman for sharing his knowledge of Vexillum brunneolinea with us. We regret that we could not name the species for him, but there is already a Vexillum cosmani Kay, 1979, and the International Commission on Zoological Nomenclature discourages giving species in the same genus or in allied genera names honoring the same person [ICZN Recommendation D(I)6]. We thank Robert H. Cowie of the Bernice P. Bishop Museum (BPBM), Leonard C. Hill (Miami, Florida), and Sadao Kosuge of the Institute of Malacology, Tokyo (IMT) for providing information and loaning specimens. LITERATURE CITED Adams, A. and L. Reeve. 1848-1850. Mollusca. In: Adams, A. (ed.). The zoology of the voyage of H.M.S. Samarang; under command of Captain Sir Edward Belcher, London. P. 1-24, pis, 1-9 (1848), p. 25-87, pis. 10-24 (1850). Cernohorskv, W. O. 1970. Systematics of the families Mitri- dae &i Volutomitridae (Mollusca: Gastropoda), Bulletin of the .\uckland Institute and Museum, no, 8 iv, 190 p. Cernohorskv, W, O. 1972, Marinesheilsof the Pacific, volume II, Pacific Publications, Sydney, 411 p., 68 pis. Gmelin, J, F, 1791. Caroli a Linne Systema naturae per regna tria naturae. Edition decima tertia. Lipsiae, 1(6), Vermes, pp, 3021-3910. Kay, E. A. 1979. Hawaiian marine shells. Reef and shore fauna of Hawaii. Section 4: Mollusca, Bernice P, Bishop Museum Special Publication 64(4), xviii, 653 p. Kosuge, S. 1979, Report on the Mollusca on guyots from the central Pacific collected by 2nd and 3rd cruises of R/V Kaiyomaru in 1972 to 73 with descriptions of twelve new species. Bulletin of the Institute of Malacology, Tokyo, l(2):24-35, pis. 5-6. Lamarck, J. B. P. A. de M. de. 1811. Sur la determination des especes parmi les animaux sans vertebres, et particu- lierement parmi les mollusques testaces. .\nnales du Mu- seum d'Histoire Naturelle (Paris) 17:195-222, MacDonald, J. D. 1860, Further observations on the meta- morphosis of Gasteropoda, and the affinities of certain genera, with an attempted natural distribution of the prin- cipal families of the order Transactions of the Linnean Society of London 23:69-80. 1 table, Pilsbry, H, A, 1921, Marine mollusks of Hawaii, VIII-XIII, Proceedings of the Academv of Natural Sciences of Phil- adelphia 72:296-328, pi. 12^ Reeve, L. 1844-1845. Monograph of the genus Mitra. Con- chologia Iconica 2, pis, 1-27 (1844), 28-39 (1845). Roding, P. F, 1798, Museum Boltenianum sive catalogus ci- meliorum e tribus regnis naturae quae collegerat Joa, Fried Bolten. Pars Secunda Johan, Christi, Trappii, Hamburg, viii, 199 p Salisbury, R, 1981, Little known miters of Hawaii II. Ha- waiian Shell News 29(2):6, I color plate. Smith, E. A. 1876. A list of marine shells, chiefly from the Solomon Islands, with descriptions of several new species. Journal of the Linnean Society of London. Zoology. 12: 535-562, pi. 30. Turner, H. 1989. llncommon and new mitriform gastropods from the Indo-Pacific. Part 1. Swiss Federal Research In- stitute for Forest, Snow and Landscape, Birmensdorf. 35 p., 8 color plates. THE NAUTILUS 105(4):152-158, 1991 Page 152 Parasitic Mites (Acari: Unionicolidae) of Fresh-water Mussels (Bivalvia: Unionidae) in the Duck and Stones Rivers in Central Tennessee Malcolm F. Vidrine Louisiana State University at Eunice P.O Box 1129, Eunice, LA 70535 USA James L. Wilson Tennessee State University Tenth and Charlotte, Nashville, TN 37203 USA ABSTRACT The Duck and Stones Rivers in central Tennessee were sampled for fresh-water mussels in 1956 and 1962. Twenty-nine species of Stones River mussels were examined for parasitic mite in- festations Twenty mussel species were infested by a total of 14 species of mites (Unionicola). Of the 552 mussel specimens examined, 321 (58.2%) were infested with one or more mite species. In the Duck River, 20 species of mussels were examined for parasitic mite infestations. Thirteen mussel species were infested by a total of 11 species of mites (Unionicola). Of the 637 mussel specimens examined, 381 (59.8%) were infested with one or more mite species. Mussel-mite associations are listed. These data provide a unique insight into a native fauna that may now be irrecoverably damaged, at least in the im- pounded areas of Stones River. Key words: Unionidae; mussels; Unionicolidae; Unionicola; mites; Stones River; Duck River; Tennessee. INTRODUCTION Fresh-water mussels (Bivalvia: Unionidae) form a ma- jor part of the macrobenthos in streams and lakes of North America. Although these mussels are commonly parasitized by aquatic mites (Acari: Unionicolidae), few published studies of geographical co-occurrence of the host and parasite species exist. Classical works, especially those of Wolcott (1899) and Marshall (1933), provide a few host records and no comprehensive mite species in- cidence records for the stations sampled. In the first con- cise work on the Tennessee mussel mites, Mitchell and Wilson (1965) describe 5 new species of mites from 5 mussel species in the Duck River. Before the 1980s, the taxonomy of unionicolid mites was very poorly under- stood; recent works (Vidrine 1980, 1985a, 1986d, 1987, 1988), however, propose 53 new species and a revision of the supraspecific systematics. Vidrine (1989), Vidrine and Vidrine (1987), and Vidrine, Bouchon and Poirrier (1986) discussed incidence of mussel-mite associations in Arkansas, Louisiana, Mississippi, and Texas. Earlier at- tempts to describe mussel-mite community structure bv Dobson (1966), Calnan (1976) and \idrinc (1974) were hampered by the lack of recognition and definitive de- scriptions of many of the mite species encountered. This paper describes the mussel-mite associations from collections made by JLW (junior author) in the Stones River (Cumberland River drainage) and the Duck River (lower Tennessee River drainage) in central Tennessee, during 1956 and 1962. These data provide a unique insight into a native fauna of the Stones River (impound- ment in 1968: J. Percy Priest Lake) and the Duck River (impoundment in 1976: the Normandy Project). MATERIALS AND METHODS During the summers of 1956 and 1962, mussels were collected from the following 8 stations in the Stones Riv- er: 1 . Stones River at lower end of Charlton Ford off Couch- ville Pike, Davidson Co., Tennessee. 25 August 1956 (now under backwater of J. Percy Priest Lake). 2. Stones River ca. 200 m above Couch\ille Pike Bridge, Davidson Co., Tennessee. 18 June, 16 and 29 July 1962 (now under backwater of J. Percy Priest Lake). 3. Stones River ca. 100 m above bridge on Stewart's Ferry Road, southeast of Donelson, Davidson Co., Tennessee. 14 August 1962 (now under backwater of J. Percy Priest Lake). 4. Stones River ca. 1.0 km below "Old Jefferson ' Bridge ca. 4.5 km east of Smyrna and 1.3 km north of Hwy. 6306, Rutherford Co., Tennessee. 17 .August 1962 (now under backwater of J. Percy Priest Lake). 5. Stones River ca. 30 m below junction of East Fork and West Fork Stones Rivers and 1.3 km upstream from "Old Jefferson" bridge and ca. 4.5 km east of Sm\rna and 1.3 km north of Hwy. 6306, Rutherford Co., Tennessee. 20 August 1962 (now under back- water of J. Perc\' Priest Lake). 6. East Fork Stones River below bridge below dam at Walterhill on Hwy. 231, ca. 0.8 km north of Mur- freesboro, Rutherford Co., Tennessee. 20 August 1962 M, F. Vidrine and J. L. Wilson, 1991 Page 153 (above impoundment and free flowing; near station E3 of Schmidt (1982)). 7. East Fork Stones River below bridge on Hw). 6189, ca. 4.5 km northeast of Readyville (on Hwy. 705), Rutherford Co., Tennessee. 23 August 1962 (above impoundment and free flowing; near station E14 of Schmidt (1982)). 8. East Fork Stones River ca. 30 m below mill dam (still operating) and ca. 1.3 km east of Lascassas and ca. 3.0 km south of Rte. Tenn. 96, Rutherford Co., Ten- nessee. 23 August 1962 (above impoundment and free flowing; near station E6 of Schmidt (1982)). Mussels were also collected from the following 10 sta- tions in the Duck River: 1. Duck River below bridge on Hwy. 41 north of Man- chester, Coffee Co., Tennessee. 31 July 1962 (above backwater of Normandy Reservoir and free flowing: J. Jenkinson, personal communication, 1990). 2. Duck River below bridge on Hw\'. 231 in Shelby ville, Bedford Co., Tennessee. 26 July 1962 (free flowing, ca. 45 km below the dam of Normandy Reservoir, and minimally impacted by cold water discharge from the impoundment: J. Jenkinson, personal com- munication, 1990). 3. Duck River below bridge on Hw\. 31a south of Chapel Hill, Marshall Co., Tennessee. 5 July 1962 and 26 July 1962 (free flowing: J. Jenkinson, personal communication, 1990). 4. Duck River ca. 200 m below dam (Lillard Mill — an old hydroelectric dam) and 0.7 km N of Hwy. 4277 at Milltown, Marshall Co., Tennessee. 25 June and July 1962 and 26 August 1956 (free flowing and with a large mussel community: J. Jenkinson, personal communication, 1990). 5. Duck River ca. 100 m below old mill dam below bridge on Hwy. 431, Maury-Marshall Co. line, Ten- nessee. 27 July and 29 August 1962 (free flowing and with a large mussel community, old dam now washed out: J. Jenkinson, personal communication, 1990). 6. Duck River above bridge on Rte. Tenn. 7 at north- western city limits of Columbia, Maury Co., Ten- nessee. 2 and 29 August 1962 (free flowing with a large mussel community: J. Jenkinson, personal com- munication, 1990). 7. Duck River below bridge at Williamsport on Rte. Tenn. 50, Maury Co., Tennessee. 2 .\ugust 1962 (free flowing but with few or no mussels: J. Jenkinson, personal communication, 1990). 8. Duck River below Wm. Erwin McEwen Bridge at Rte. Tenn. 50, Maury-Hickman Co. line, Tennessee. 23 July 1962 (free flowing but with few or no mussels: J. Jenkinson, personal communication, 1990). 9. Duck River ca. 100 m downstream from bridge at western city limits of Centerville on Rte. Tenn. 50, Hickman Co., Tennessee. 23 July and 6 August 1962. Obviously polluted by sewage in 1962 (still free flow- ing and polluted by sewage and industrial wastes. with few or no mussels: J. Jenkinson, personal com- munication, 1990). 10. Duck River below bridge on Rte. Tenn. 50, ca. 5 km west of Centerville, Hickman Co., Tennessee. 6 Au- gust 1962 (still free flowing but with few or no mus- sels: J. Jenkinson, personal communication, 1990). .\ total of 557 mussels from the Stones River, and 637 mussels from the Duck Ri\ er was examined by dissection, for adult and n\ mphal mites by JLW. All but 5 of these mussels were identified by Henry van der Schalie and Ralph Sinclair. The unidentified mussels (all from Stones River) are excluded from consideration here. The mites of each individual mussel were stored in separate, la- belled vials of Koenike's fluid. The mites were identified by MF\' in 1987. Mussel names have been updated to conform with the concepts of Burch (1975) and Davis and Fuller (1981). The mussels were deposited in the collections of the Museum of Zoology at the University of Michigan, Ann Arbor. RESULTS Fourteen species of Unionicola were found in 20 of 29 species of fresh-water mussels from the Stones River. Of the 552 mussels examined, 321 (58.2%) were infested with one or more species of Unionicola: 268 had one species, 40 had two species, and 13 had three species. Results of these examinations are summarized in table 1. The mussel diversity and abundance conform gener- ally with those reported by Wilson and Clark (1914) and Schmidt (1982). Eleven species of Unionicola were found in 13 of 20 species of fresh-water mussels from the Duck River. Of the 637 mussels examined, 381 (59.8%) were infested with one or more species of Unionicola: 372 had one species, 7 had two species, and 2 had three species. Results of these examinations are summarized in table 2. The mussel diversity and abundance conform generalK- with those reported bv Isom and Yokelv (1968), H. van der Schalie (1973), Ahlstedt (1986), and Jenkinson (1988). Mussel-mite associations can be classified as resident, transient, or vagrant (Vidrine 1989), as adapted from Treat's (1975) definitions for Lepidoptera-mite associa- tions. All 15 species of Unionicola reported here are naturally resident mites in the encountered mussel hosts, as inferred from associations encountered in other studies (Vidrine 1985b, 1986a,b,c, 1987, 1989). The few vagrant associations encountered here are naturally incidental and/or inevitable artifacts of sample handling. The following associations were encountered in the Stones River (site of natural infestation within branchial chamber of hosts is provided within parentheses) (% in- fection of a specific mussel species by a specific mite species is provided in parentheses): 1. Unionicola dimocki Vidrine 1986b (gills) (83.1%) and U. smithae Vidrine 1986a (mantle and foot) (6.8%) were found in Lasmigona costata (Rafinesque Page 154 THE NAUTILUS, Vol. 105, No. 4 Table Sumniarv of miisscltiiitf associations in the Stones River, Tennessee, based upon collections in 19.56 and 1962 Mussels Mites Number of mussels examined (in bold) Number of mussels infested by parasitic mites Stations Total Anodonta grandis Say 1829 I'nionicold smilhai' X'idrine 1986a Lasmigona costala (Rafinesque 1820) U. dimocki X'idrine 1986b V. smithac X'idrine 1986a Amblema plicata Say 1817 U. amandita Mitchell and XVilson 1965 U. tupara Mitchell and XVilson 1965 U. serrata (XX'olcott 1898) Amblemn gignntea (Barnes 1823) ('. tupara Mitchell and XX'ilson 1965 Fusconnin flnva (Rafinesque 1820) ['. parheri X itirine 1987 Elliptio dilntata Rafinesque 1820 V. scrrala (Wolcott 1898) U. tupara Mitchell and XVilson 1965 E. crassidens (Lamarck 1819) Cyclonaias tuberculata (Rafinesque 1820) U. sakantaha Mitchell and Wilson 1965 U. abnormipes (X\'olcott 1898) U. causeyae X'idrine 1985b U. serrata (XVolcott 1898) U. vamana Mitchell and XVilson 1965 Quadrula pustulosa (Lea 18.31) ['. causeyae X'idrine 1985b Orthonymus cylindrica (Say 1817) I', sakantaka Mitchell and XX'ilson 1965 Tritogonia verrucosa (Rafinesque 1820) I', vamana Mitchell and XX'ilson 1965 Actinonaias carinata (Barnes 1823) U. hoesei X'idrine 1986c v. abnornnpes (XVolcott 1898) U. serrata (Wolcott 1898) A. pectorosa (Conrad 1834) U. hoesei X'idrine 1986c U. serrata (Wolcott 1898) U. abnormipes (Wolcott 1898) Lampsilis orbiculata (Hildrelh 1928) ['. hoesei X'idrine 1986c r. abnormipes (XVolcott 1898) L. ovata (Say 1817) U. hoesei X'idrine 1986c v. almormipes (XVolcott 1898) U. serrata (Wolcott 1898) L. fnsrioln Rafinesque 1820 Ohnvnrin subrotiinda (Rafinesque 1820) Truncitia trunratn Rafinesque 1820 T. donatifurntis (Lea 1828) Plagioln lenior (Lea 1840) /'. interruptii (Rafinesqu<- 1820) I'otaniiliis purpitralus (l.aniurek 1819) P. alatus (Say 1817) (/. hoesei X'idrine 1986c U.fulleri Vidrine 1986c U. australindistincta Vidrine 1985b V. serrata (Wolcott 1898) II 8 26 2 17 12 13 5 3 3 1 17 10 2 2 4 3 1 1 3 1 37 2 2 35 18 2 6 5 1 19 1 1 1 1 6 6 13 10 5 2 15 10 4 4 1 14 14 4 — 10 8 1 18 15 6 — — 1 15 — — 1 2 3 34 13 U 1 4 — 4 — 1 — 1 1 55 51 i 22 15 2 — — 118 98 8 151 15 53 11 23 17 1 1 35 1 2 2 26 24 2 6 2 1 22 1 1 1 7 6 16 12 5 3 33 23 8 4 4 1 2 26 22 6 3 4 1 1 1 1 4 1 16 11 4 5 1 M. F. Vidrine and J. L. Wilson, 1991 Page 155 Table Continued Mussels Mitfs Number of mussels exaniiiied (in bold) Number ot mussels infested b\' parasitic mites Stations ["otal Ligumia recta (Lamarck 1819) U. serrata (VVolcott 1898) [/. ahnormipes (Wolcott 1898) Villosa taeniata (Conrad 1834) ('. hoesei Vidrine 1986c Medionidus conradicus (Lea 1834) ['. hciulrixi N'idrine 1987 Leptoden fragilis (Rafinesque 1820) ['. hoesei Vidrine 1986c Ptychobranchus fasciolare (Rafinesque 1820) U. hoesei Vidrine 1986c C. can.seycw Vidrine 1985b Obliquaria reflexa Rafinesque 1820 Totals ol' mussels 2 2 3 3 11 1 146 1 — 1 — 9 3 2 — J9 ()2 86 6 1 1 12 2 3 3 2 2 33 8 1 1 1820); a single vagrant specimen of V . smithae was found in Anodonta grandis Sa> 1829. 2. Unionicola amandita Mitchell and Wilson 1965 (gills) was found only in Amblema plicata Say 1817 (10.0%). 3. Unionicola tupara Mitchell and Wilson 1965 (gills) was found in A. plicata (35.1%) and Amblema gi- gantea (Barnes 1823) (73.9%). 4. Unionicola sakantaka Mitchell and Wilson 1965 (gills) was found in Cyclonaias tubercidata (Rafin- esque 1820) (92.3%) and Orthonynms cylindrica (Say 1817) (100.0%). 5. Unionicola vamana Mitchell and Wilson 1965 (gills) was found in Tritogonia verrucosa (Rafinesque 1820) (85.7%); a vagrant specimen was found in C. tuber- cidata. 6. Unionicola hoesei Vidrine 1986c (gills), a common parasite of 15 genera in the Lampsilini (Unionidae: Ambleminae) in North America (Vidrine 1986c), was found in Lampsilis ovata (Say 1817) (84.6%), Lamp- silis orbiculata (Hildreth 1828) (25.0%), Villosa taeniata (Conrad 1834) (41.7%), Actinonaias cari- nata (Barnes 1823) (75.0%), Actinonaias pectorosa (Conrad 1834) (69.7%), Leptodea fragitis (Rafin- esque 1820) (100.0%), Ptychobranchus fasciolare (Rafinesque 1820) (24.2%) and Potamilus alatus (Say 1817) (68.8%) (all lampsilinines) in the Stones River. 7. Unionicola fidleri Vidrine 1986c (gills) (25.0%) and Unionicola australindistincta Vidrine 1985b (man- tle and foot) (31.3%) were found only in P. alatus. 8. Unionicola hendrixi Vidrine 1987 (gills) was found only in Medionidus conradicus (Lea 1834) (100.0%). 9. Unionicola parkeri N'idrine 1987 (gills) was found only in Fusconaia flava (Rafinesque 1820) (100.0%). 10. Unionicola causeijae Vidrine 1985b (mantle and foot) was found in C. tubercidata (23.1%), P. fasciolare (3.0%), and Quadrula pustulosa (Lea 1831) (4.5%). 11. Unionicola ahnormipes (Wolcott 1898) (mantle and foot) was found in A. carinata (31.3%). A. pectorosa (12.1%), L. ovata (23.1%), L. orbiculata (50.0%) and Liqumia recta (Lamarck 1819) (16.7%); a vagrant specimen was found in C. tubercidata. 12. Unionicola serrata (Wolcott 1898) (labial palps) was found in A. carinata (18.8%), A. pectorosa (24.2%), A. plicata (7.3%), C. tuberculata {1.1%), Elliptio di- latata Rafinesque 1820 (2.9%), L. ovata (11.5%), L. recta (16.7%), and P. alatus (6.3%). The following associations were encountered in the Duck River (site of natural infestation within branchial chamber of hosts is provided w ithin parentheses) (% in- fection of a specific mussel species by a specific mite species is provided in parentheses): 1. Unionicola dimocki (gills) (89.5%) and U. smithae (mantle and foot) (12.3%) were found only in L. cos- tata. 2. Unionicola amandita (gills) was found only in A. plicata (48.7%). 3. Unionicola tupara (gills) was found in A. plicata (0.3%) and A. gigantea (84.2%). 4. Unionicola sakantaka (gills) was found in C. tuber- culata (9O.7%0 and O. cylindrica (100.0%); a vagrant specimen was found in T. verrucosa. 5. Unionicola vamana (gills) was found in T. verrucosa (83.0%); vagrant specimens were found in C. tuber- cidata (1) and Q. quadrula (Rafinesque 1820) (5). 6. Unionicola vikitra Mitchell and Wilson 1965 (gills) was found in Q. quadrula (47.6%) and C^. pustulosa (100.0%); a vagrant specimen was found in C. tuber- culata. 7. Unionicola hoesei (gills) was found in Lampsilis fas- ciola Rafinesque 1820 (50.0%), Villosa fabalis (Lea 1831) (81.3%), A. pectorosa (62.5%), L.fragilis (8.3%), and P. alatus (30.0%) (all lampsilinines). Page 156 THE NAUTILUS, Vol. 105, No. 4 Table 2. Siiminar> of mussel-mite associations in the Duck Ri\er, Teiuiessee, based upon collections in 19.56 and !9fi2 Number of niu§§eU examined (in biild) Number of mussels infested h\ parasitic mites Stations Mussels Mites 12 3 4 .5 fi 7 8 9 10 Total Anodonia grandis Say 1829 — — — — — I — — — — 1 Alasniidonta marginatn Say I8I*J — — — — 2 — — — — — 2 Lasrnigona complanntn {Barne> 1823) — — — — — o — — — — a L. cosrafa (Rafinesque 1820) — II 18 14 14 _____ 57 Uniotncola dimochi Mdrine l9H6h — 8 18 11 14 _____ 51 ['. smithae \idrine 1986a — 14 1 ]_____ 7 Strophitus undiilatus (Say 1817) — — — — — — — — 3 — 3 .4m6/pma/»/ic«fn Say 18i7 — 3 29 170 74 42 — — — — 318 r. «"m;i(/i7« Mitcliell and WiLson 1965 — 2 17 90 45 i _ _ _ _ 155 C. (K/wro Mitchell and Wilson 1965 _____ i____ i .4. gigantea (Barnes 1823) _ _ _ 5 10 4 — — — — 19 C;. (i/paru Mitchell and Wilson 1965 _ _ _ 4 1(1 2 — — — — 16 Cyc/onaias fubercu/afa (Rafinesque 1820) — 1 28 13 9 I 2 — — — 54 U. sakanlaka Mitchell and Wilson 1965 _ 1 27 12 9 — — — — — 49 U. vikitra Mitchell and Wilson 1965 __!_______ 1 U. varnana Mitchell and Wilson 1965 — — — — — 1 — — — — 1 Tritogonia verrucosa (Rafinesque 1820) — — 19 20 6 5 — — 3 — 53 v. varnana Mitchell and Wilson 1965 — — 19 17 6 2 — — — — 44 U. sakantaka Mitchell and Wilson 1965 ___ ]______ i Orthonymus cylindrica (Say 1817) — — 2 — — — — — — — 2 U. ,sa/ca;i(a/l-a Mitchell and Wilson 1965 __2 — — — — — — — 2 Quadrula quadrtila (Rafinesque 1820) — — — — — 60 — 1 2 — 63 C. (iAi7ra Mitchell and Wilson 1965 _ _ _ _ _ 30 _ _ _ _ 30 ['. varnana Mitchell luul Wilson 1965 — — — — — 5 — — — — 5 Q. pustulosa (Lea 18SI) ___ 1_ i____ 2 U. vikitra Mitchell and Wilson 19(i5 ___ i_ i____ 0 Elliptio crassidens (Lamarck 1819) — — — — — 1 — — 2 — 3 Lampsilis ovatn (Say 1817) — — — — — — — — 1 — 1 L. teres (Rafinesque 1820) _____ 2 — — — — 2 L. fasciola Rafinesque 1820 6 — — — — — — — — - — 6 I', huesei V'idrine 1986c 3 — — — — — — — — — 3 ViUosafabalis (Lea lii3\) 16 — — — — _____ 16 r. /ioesei \idrine 1986c 13 — — — — _____ 13 Actinonaias pectorosa (Conrad 1834) 5 — 1 1 1 — — — — — 8 U. hoesei V'idrine 19S6c 4 — 1 — — — — — — — 5 Lepfodea/ragi/is (Rafinesque 1820) ____ 1 2 — — 9— 12 V. hoesei V'idrine 1986c ____ i_____ 1 Potnmi7us a/afus (Say 1817) ___ 3 i___5110 C. /irw.vn \idrine 1986c — — — 3 ______ 3 U./ii/Zfri \idrine 1986c ___ 2 i_____ 3 V. australindistincta V'idrine 1985b — — — 1 — — — — — — 1 U. serrata (Wolcott 1898) — — — ]______ 1 Totals of mussels 27 15 97 227 118 124 2 1 25 1 637 8. Unionicola fulleri (gills) (30.0%), U. serrata (labial palps) (10.0%), and U. australindistincta (mantle and foot) (10.0%) were found only in P. alatus. The following mussels were not found parasitized by mites in either or both of the rivers studied: Alasniidonta marginata Say 1819, Lasmigona complanata (Barnes 1823), Strophitus undulatus (Say 1817), Elliptio cras- sidens (Lamarck 1819), Lampsilis teres (Rafinesque 1820), Obovaria subrotunda (Rafinesque 1820), Trun- cilla truncata Rafinesque 1820, Truncilla donaciformis (Lea 1828), Plagiola lenior (Lea 1840), Plagiola inter- rupta (Rafinesque 1820), Potamilus purpuratus (La- marck 1819), and Obliquaria reflexa Rafinesque 1820. DISCUSSION Intra-host regulation of a mite-parasite load, perhaps through rivalry of sexual competitors, as described by Mitchell (1965), was apparent in several mite species encountered here. In the Stones River, Unionicola aman- M. F. Vidrine and J. L. Wilson. 1991 Page 157 dita, U. dimocki, U. hoesei, U. sakantaka. U. vamana, V. hendrixi and V. parkeri t>picall\- occurred in small, characteristicaih discrete numbers within each host: a single male, and one or two females usually infested the gills of their mussel host. One to four individuals of V. tupara and L'. fulleri (all males, all females, or combi- nations of both sexes) infested the gills of their hosts. One or two females of U. smithae were usually found on the mantle or foot of infested hosts, while males were absent, probabK due to seasonal reasons. One to four specimens of the gill mites V. tupara and I', fulleri were found in each host. One or two indi\ iduals of the less host-specific labial palp parasite, U. serrata, were found in 7 genera of mussels. One to five individuals of U. abnormipes, U. causeijae and U. australindistincta, which infest the mantle and foot of their hosts, were found in a variety of amblemine hosts. The large number of individuals (range 1-65, but most often l)oi U. abnormipes infesting individual hosts may represent an absence of the strict territorialit) apparent in the above associations. In the Duck River. Unionicola amandita, U. dimocki, V. hoesei, U. sakantaka. U. vamana and V. vikitra typ- ically occurred in small numbers within each host: a single male, and one or two females infesting the gills of their individual hosts. One to nineteen. usualK 5. individuals of U. tupara and V. fulleri infested the gills of their hosts. One to five females of U. smithae were found on the mantle or foot in each infested host, while males were not found. Onl\ a single male of U. serrata, a labial palp mite, was found. A single Potamilus alatus had one male and two females of V. australindistincta, a mantle and foot parasite. In both the Stones River and the Duck River. V . aus- tralindistincta was found in P. alatus. These are new- host and localit\' records for L'. australindistincta, which has been previously known mainly from other species of the genus Potamilus and Leptodea fragilis in .\rkansas, Louisiana, Mississippi, and Te.xas (Mdrine 1986b). The closely related mite species, L'. indistincta (Wolcott 1898). is the usual parasite of P. alatus, both in the upper Mis- sissippi and upper Tennessee river drainages (Clinch and Powell Rivers) (N'idrine, 1980; Wolcott. 1899). The ad- dition of P. alatus to the host list for L'. australindistincta neither confirms nor refutes Mdrine's (1985b) contention that U. australindistincta is a southern form of U. in- distincta. In this sampling. onl\ a single, unparasitized P. purpuratus (a common host of U. australindistincta south of Tennessee) was found in the Stones River. The data in tables 1 and 2 illustrate the incidence of specific mite taxa in specific mussel groups (subfamilies Anodontinae and Ambleminae). The anodontine mussels, for example, are infested by distinctly different mantle and gill mites than are amblemine mussels. This phe- nomenon was noted by Vidrine and Bereza (1978) and reinforced by Vidrine (1980, 1986a,b,c,d, 1987. 1989). These data suggest that Unionicola tend to be found in the more abundant mussel species in a habitat. .^ critical threshold of host population size may be an oblig- atory prerequisite for a population of a specific, extant mite species to be sustained locally. Small and unpar- asitized populations of certain mussel species may be naturally below a threshold assuring sur\ival of mites capable of parasitizing them, or artificially so as a result of human impact, .although found parasitized elsewhere, the following species were locally Unionicola-iree in this study: E. crassidens was found parasitized in neither the Duck or Stones Rivers; L. fasciola, O. subrotunda, T. truncata, T. donaciformis, P. purpuratus, and O. refiexa were not found parasitized in the Stones River; A. gran- dis, A. marginata. L. complanata, S. undulatus, L. ova- ta, and L. teres were not found parasitized in the Duck River. Whether or not this local phenomenon is due to naturalK small or to damaged populations of these mus- sels, is unknown. Some mussel species ma\- have never achieved critical population sizes that could realize the very evolution of a mite into a parasitic relationship with them. The t%p- ically small populations of mussels in the genus Plagiola may exemplify this phenomenon, and may be culpable in hastening the extinction of several species of Plagiola in the last centur>-, little doubt as a result of human impact. Plagiola were never found parasitized by Unio- nicola in the field or in museum holdings thus far bv MFV, and no published records are extant. The data derived from these collections are historicalK important because all mussels that were found were col- lected, providing a rough estimate of the relative species abundance and incidence in these damaged and/or im- pacted faunas. Also, these data are enriched by a mite- mussel parasitization profile, providing another dimen- sion to the systematics and natural history of these faunas, and an insight into a diverse and differentially specialized host-parasite complex. The Cumberlandian mussel fauna and much of the lower Tennessee River fauna, now large- ly decimated down to their few remaining refugia, his- torically represented the most diverse svmpatric fresh- water mussel fauna known on Earth. Insights into the mite-mussel coevolutionar\' strategies and into the deli- cate population control mechanisms that are necessar\ to maintain such a diverse community are now largely unattainable from these mussel communities, as the nec- essary sampling of mussels is untenable in the few viable populations, or impossible in those extirpated by im- poundment. Recent mussel studies (Ahlstedt 1986; Jen- kinson 1988; Schmidt 1982) do indicate that diverse com- munities still exist in portions of both the Duck and the Stones Rivers. The presented baseline data sets will, we hope, provide some corroborative elements to future studies of the now-relict Cumberlandian and other mus- sel faunas and their parasites and perhaps engender in- terest from ecologists who seek to explain the intricacies that drive the complex biotic relationships within benthic invertebrate communities. ACKNOWLEDGEMENTS We sincereK thank Daniel J. Bereza for his assistance in the preparation of this paper. John J. Jenkinson, Ten- nessee X'alley Authority, provided valuable information on the present conditions of the rivers. Page 158 THE NAUTILUS, Vol. 105, No. 4 LITERATURE CITED Ahlstedt, S. A. 1986. Cumberland mollusk conservation pro- gram. Activity 1: mussel distribution surveys. Tennessee Valley Authority, Norris, TN. 125 p. Burch, J. B. 1975. Freshwater Unionacean clams (Mollusca: Pelecypoda) of North .America. Malacological Publica- tions, Hamburg, MI, 204 p. Calnan, T. R. 1976. The systematics and distribution of meta- zoan parasites in the Unionidae from Navasota River, Tex- as. M.S. thesis, Texas A and M University, College Station, TX, viii -I- 112 p. Davis, G. M. and S. L. H. Fuller. 1981. Genetic relationships among Recent Unionacea (Bivalvia) of North America. Malacologia 20(2):217-253. Dobson, R. 1966. A survey of the parasitic Unionicolidae (Arachnida; Acarina) of the Apalachicolan faunal region of the southern United States. M.S. thesis, Florida State University, Tallahassee, FL, vii + 99 p. Isom, B. G. and P. Yokely Jr. 1968. The mussel fauna of Duck River in Tennessee, 1965. American Midland Naturalist 80(l):34-42. Jenkinson, J, ], 1988. Resurvey of freshwater mussel stocks in the Duck River, Tennessee Tennessee Valley Authority — River Basin Operations — Water Resources, Knoxville, TN, iii + 28 p. Marshall, R. 1933 Preliminary list of the Hydracarina of Wisconsin. Part III. Transactions of the Wisconsin Acad- emy Science 40(3):37-61. Mitchell, R, 1965, Population regulation of a water mite par- asitic on unionid mussels. Journal of Parasitology 51:990- 996. Mitchell, R. and J. L. Wilson. 1965. New species of water mites from Tennessee unionid mussels. Journal of the Ten- nessee Academy of Sciences 40(3):104-106. Schmidt, J. E. 1982. The freshwater mussels of the Stones River above J. Percy Priest Reservoir, Tennessee. M.S. thesis. Tennessee Technological University, Nashville, TN, viii -I- 65 p. Treat, A. E. 1975. Mites of moths and butterflies. Cornell University Press, Ithaca, NY, 362 p van der Schalie, H. 1973. The moUusks of the Duck River drainage in central Tennessee. Sterkiana 52:45-55. Vidrine, M. F. 1974. Aspidogastrid trematode and acarine parasites of freshwater clams in south central and south- western Louisiana. M.S. thesis. Department of Zoology and Physiology, Louisiana State University, Baton Rouge, LA, viii + 125 p. Vidrine, M. F. 1980. Systematics and coevolution of unionico- lid water-mites and their unionid mussel hosts in the east- ern United States. Ph.D. dissertation. Department of Bi- ology, University of Southwestern Louisiana, Lafayette, LA, xvii -I- 661 p. Vidrine, M. F. 1985a. Berezatax, new subgenus (Acari: Unio- nicolidae: Unionicolinae: Unionicola). in subtropical and tropical North America, with a re-evaluation of the higher taxonomic groups of related American unionicoline mites. International Journal of Acarology 1U2):133-141. N'idrine, M. F. 1985b. Six new species in the subgenus Poly- atax (Acari: Unionicolidae: Unionicola ) from North .Amer- ica, with a re-evaluation of related taxa International Jour- nal of Acarology ll(4):273-287. Vidrine, M. F. 1986a. Anodontinatax, new subgenus (.\cari: Unionicolidae: Unionicola), in holarctic fresh- water mus- sels (Unionidae: Anodontinae), with a re-evaluation of re- lated taxa. International Journal of Acarology 12(2):95- 104. Vidrine, M. F. 1986b, Five new species in the subgenus Par- asitatax (Acari: Unionicolidae: Unionicola) from North America and Asia, with a re-evaluation of related species. International Journal of Acarology 12(3):141-153. Vidrine, M. F, 1986c. Ten new species in the subgenus Union- icolides (Acari: Unionicolidae: Unionicola) from North and South America, with a re-evaluation of related species. International Journal of Acarology 12(3):123-140, Vidrine, M. F. 1986d, Revision of the Unionicolinae (Acari: Unionicolidae). International Journal of Acarologv 12(4): 233-243. Vidrine, M. F. 1987. Ten new species of water mites (Acari: Unionicolidae: Unionicola: Unionicolides) in North American fresh-water mussels (Unionacea: Unionidae; Ambleminae). International Journal of Acarology 13(4); 251-259. Vidrine, M. F. 1988. Five new subgenera of Unionicola (Ac- ari: Unionicolidae). Proceedings of the Louisiana .Academy of Sciences 51:31-35. Vidrine, M. F. 1989. A summary of the mollusk-mite asso- ciations of Louisiana and adjacent waters. The Louisiana Environmental Professional 6(l):30-63. Vidrine, M. F. and D. J. Bereza. 1978. Some considerations and implications of host-specificity studies of unionicolid mite parasites on the systematics of some groups of North American unionacean fresh-water mussels. Bulletin of the American Malacological Union, Inc. for 1977:85-86. Vidrine, M. F. and M F Vidrine II. 1987 Macroinvertebrate benthic communit> structure in Louisiana Irrigation Ca- nal. The Louisiana Environmental Professional 4(1):66- 74. Vidrine, M. F, R. J. Bouchon, and M. A. Poirrier. 1986. New North American records of water mites (Acari: Unionicoli- dae: Unionicola: Uitionicola and Pentatax). International Journal of Acarology 12(4):229-232. Wilson, C. B. and H. W. Clark. 1914. The mussels of the Cumberland River and its tributaries. Bureau of Fisheries Documents 781:1-63, Wolcott, R. 1898. New American species of the genus Atax (Fabr.) Bruz. Zoological Bulletin 1:279-285. Wolcott, R. 1899. On the North .American species of the genus Atax (Fabr.) Bruz. Transactions of the .American Micro- scopical Society 20:193-259. THE NAUTILUS 105(4):159-164. 1991 Volutes from Sava de Malha Bank: The Saga of Lijria siirinamensis and a New Species Page 159 Philippe Bouchet Museum national d'Histoire naturelle 55 rue Buffon 75005 Paris, France Patrice Bail 2 square LaFontaine 75016 Paris, France ABSTRACT The supposedly fulgorariine volute Dallivoluta surinamensis Okutani, 1982 is shown to have been described from an erro- neous type locality in the Caribbean. It actually originates from Saya de Malha Bank, in the Indian Ocean, where it is probably endemic. Dallivoluta Okutani, 1982 is regarded as a synonym of Lyria Gray, 1847. It is further suggested that Murex suri- namensis Okutani, 1982 was also erroneousK- localized and actually originates from the same submarine Indian Ocean bank. Lyria doutei n.sp. is described from Saya de Malha, and compared with other species in the group of L. lyraeformis. Endemism on Saya de Malha is discussed in connection with the history and isolation of the bank. Key words: Indian Ocean; Mascarene Plateau; Caribbean; Vol- utidae; endemism. INTRODUCTION In the last few months, collectors in Western Europe have obtained from Russian sources specimens of two remarkable volute species trawled by Soviet research vessels on Saya de Malha Bank, in the Indian Ocean. Nothing similar had been described from the Indo-Pa- cific, and the Sa\ a de Malha shells appeared to represent two new species of Lyria. when we noticed the identitv' of one of them with Dallivoluta surinamensis Okutani, 1982. We present below the reasons why we believe that D. surinamensis was erroneously localized in the original description, we synonymize Dallivoluta with Lyria, and we describe the second Lyria as a new species. The species Dallivoluta surinamensis Okutani, 1982: The volute Dallivoluta surinamensis was described as a new genus and species, based on a single dead shell (figure 1 ) and fragments collected by Japan Marine Fish- ery Resources Research Center (JAMARC), supposedly off the coast of Surinam in December 1978 (Okutani, 1982:115). The type locality was given as 11°03'N, 61°01'W, depth 128 m. Okutani (in Takeda & Okutani, 1983:290) provided an abridged description and color photographs of the holotype. No further original infor- mation on Dallivoluta surinamensis has been published since the original description. Okutani (1982) compared Dallivoluta with Tractolira Dall, 1869 and especially Fulgoraria Schumacher, 1817, which is probably the reason why Poppe (1987:14) placed Dallivoluta in the subfamily Fuigorariinae. The type locality: The t\ pe locality indicated by Oku- tani (1982) is not situated off Surinam, as the specific name would imply. The geographical coordinates indi- cate a position 700 km to the northwest of Surinam, between the islands of Trinidad and Tobago, off Vene- zuela. The depth (128 m) is in the range of depths found around these islands. The type locality of Murex suri- namensis Okutani, 1982, described in the same paper, is 10°46'N, 61°32'W, 94 m. These coordinates indicate a position on the island of Trinidad, not in the seas around it. In fact, neither of these localities falls within the area encompassed by the JAMARC study area off Surinam and French Guiana (Takeda & Okutani, 1983:9, 11), and there are grounds to suspect that they are erroneous. Yokes (1990a:4, footnote; 1990b:6) had expressed doubts on the type locaHty of Murex surinamensis, and sug- gested that "opportunity for mi.xed locality data is cer- tainly present" (Vokes, 1990a). Identity of Dallivoluta surinamensis with the Saya de IVIaIha volute: The volutes obtained from Soviet sources belong to two different species of Lyria. Of one of these, we have seen two complete shells, empty but very fresh and probably live taken, said to have been collected by Soviet research vessels on Saya de Malha: one is in the Doute collection (Germany); the other is in the second author s collection (figure 2). In addition, we have seen photographs of a third specimen from another private collection. Comparison of this material with Okutani's (1982, 1983) description and figures (this paper figure 1) leaves no doubt as to their identity. The spiral cords are more distinct on the spire whorls of the newly collected specimen, but this is undoubtedK due to the better pres- ervation of this shell. We therefore conclude that the type locality is erro- neous. When the geographical coordinates of the type locality are read with South instead of North, and East instead of West {i.e.. 11°03'S, 61°01'E, instead of 1 1°03'N, Page 160 THE NAUTILUS, Vol. 105, No. 4 Figure 1. Lyria surinamensis (Okulani, 1982). Holotype (from Okutani, in Takeda & Okutani, 1983). Height 105 mm. 61°0rW), the position given is precisely correct for Saya de Malha Bank, in the Indian Ocean! We also suggest that the type locality of Murex surinamensis is similarly inaccurate, and should be read 10°46'S, 61°32'E, also on Saya de Malha Bank. This would confirm the observation by Ponder and Yokes (1988;12) and Yokes (1990a:4, 1990b:6) that there are no species of Murex s.s. in the New World. It would thus appear that the Surinam material pro- vided to Dr. Okutani by JAMARC had unfortunately been contaminated by specimens trawled from the In- dian Ocean. Dallivoluta surinamensis and Murex surinamensis will add to the list of species with unfortunate geographical specific names, such as the Caribbean Cassis madagas- cariensis Lamarck, 1822, originally believed to come from Madagascar, or Punclurella agulhasae Clarke, 1961, the type locality of which was erroneously believed to be within the limits of Agulhas Basin (see Barnard, 1963: 296). A similar case of mistaken geographical coordinates concerns the magellanic Marginella warrenii Marrat, 1876, described from 50°23'N, 64°04' W (Gulf of St. Law- rence), an error for 50''23'S, 64°04'W (between the Falk- lands and Cape Horn) (Bavay, 1913). Dallivoluta, a synonym of Lyria: Okutani was struck by the general resemblance of D surinamensis with the NW Pacific genus Fulgoraria and regarded the non- tilted protoconch, whitish columellar callus and color pattern to differ from the character states observed in Figure 2. Lyria surinamensis. Specimen collected on Sa\a de Malha Bank by Soviet vessels (Bail collection). Height 87 mm. Fulgoraria. However, we find D. surinamensis most closely related to the E. African Lyria lyraeformis (Swainson, 1821) and regard the generic characters enu- merated by Okutani to be of specific importance only. The shell characters of Dallivoluta surinamensis fall within the range observed in the genus Lyria. as restrict- ed by Bouchet (1990), and we regard Dallivoluta Oku- tani, 1982 as a synonym of Lyria Gray, 1847. Description of Lyria doutei: The second species of Lyr- ia obtained from Soviet sources is known to us from a single shell, originally in the Doute collection. At least one other specimen was collected. It is in a private col- lection in Europe or North America, and was available for examination. Lyria doutei n.sp. (figures 4, 5) Type material: Holot>pe in (Museum national d'His- toirc iiaturclle (MNHN), leg. H. Doute. Type locality: Saya de Malha Bank, 10°30'S, 62°25'E, 135 m. p. Bouchet and P. Bail, 1991 Page 161 Description: Shell (figure 4) large, fusiform, solid, con- sisting of 1.5 protoconch and 6 teleoconch whorls. Pro- toconch (figure 5) bulbous, \\ith an indication of a cal- carella on the initial part, remaining part with convex smooth whorls. Protoconch/teleoconch transition sharp. Teleoconch whorls convex, with appressed, but very slightly channeled suture. Sculpture consisting of or- thocline, gently sigmoid a.xial ribs; no spiral sculpture. Fourteen axial ribs on first teleoconch whorl, gradually increasing to 24 on body whorl. Last rib forming a thick labial varix behind outer lip. ,\perture ovate, outer lip smooth, inner lip with thin glaze over body whorl and two groups of plaits; 8 plaits in columellar region, de- creasing in strength from base to apex of shell; after a smooth zone, 3 weaker plaits in parietal region. Siphonal canal short, broad, open. Ground color creamy beige with a more complex pattern of yellowish brown to brown flamules and lines arranged in spiral bands. Spire whorls with two darker bands, adapically and abapically, sep- arated by lighter band on mid part of whorl. Adapical band with 6 brown flamules usualh extending over one rib and part of the interspaces adjacent to it; brown flamules on abapical band with more indistinct contour Medial light band with two interrupted brown spiral lines. Body whorl with one light band, and one dark basal band in addition to pattern described above. Ap- erture creamy beige. Dimensions: Height 83.5 mm, breadth 30.0 mm, ap- erture 39 mm. Remarks: Lyria doiitei is named after Mr. Harald Doute (Bad Sackingen, Germany), a keen collector specializing in volutes who first recognized it as a new species. Mr. Doute generously provided the holotype, which he had acquired from Mr. Igor Bondarev (Sebastopol, USSR). Four Indian Ocean species have a general shell mor- phology and/or color pattern that resemble that of L. doutei. These are: L. lyraeformis, L. leslieboschae Em- erson & Sage, 1986, L. cloveriana Weaver, 1963 and L. surinamensis. Lyria lyraeformis differs by having fewer axial ribs on the body whorl: it has 16 ribs in southern populations (Mombasa region, southern Kenya), and 20-23 ribs in northern populations (northern Kenya and Somalia). The color pattern of L. lyraeformis is extremely stable over its entire range: there are three uninterrupted brown bands on the body whorl, each lined with a dark brown (sometimes interrupted) thread; these brown bands are separated by two lighter spiral bands, each of them car- rying 4-7 brown lines. Lyria lyraeformis also has a some- what larger protoconch (height 4.9-5.4 mm vs 4.0 mm in doutei), and when more specimens of L. doutei are known it may appear that L. lyraeformis reaches a sig- nificantly larger adult size: Weaver & DuPont (1970) record specimens of 145 mm, but we have seen adults of 180 mm. Lyria leslieboschae is known only from the four shells on which the original description was based (Emerson & Sage, 1986). It is recognized by the distinct anal canal Figure 3. Lyria surinamensis. Protoconch of specimen in fig- ure 2 Scale line 2 mm. formed by its outer lip. Its color is of the lyraeformis type, but with a more diffused pattern. This species dif- fers from both L. lyraeformis and L. doutei by having dark axial flamules interrupting the lighter spiral bands. Lyria cloveriana is more distantK related. It has an even larger protoconch (height 6.4-7.1 mm); the axial ribs become obsolete on the abapical part of the body whorl; and the color pattern consists of evenly spaced reddish brown spiral lines, without distinct darker spiral bands as in the other species discussed abo\e. Finall>, L. surinamensis differs from L. doutei by having spiral cords in addition to the axial ribs. Lyria surinamensis may also have fewer (19) ribs on body whorl, and the color pattern consists only of poorly de- fined orange-brown spiral bands on lighter beige back- ground. Mode of development and biogeographical remarks: The bulbous protoconch of Lyria surinamensis (figure 3) and L. doutei (figure 5) are comparable in size with those of L. lyraeformis and L. cloveriana Weaver, 1963. As in all other modern volutes, it indicates non-planc- totrophic development, and the large size further sug- gests intracapsular metamorphosis and hatching as a crawling juvenile (Bouchet & Poppe, 1988). This type of development is incompatible with long distance passive transport of larvae and/or juveniles and, considering the isolation of Saya de Malha Bank (figure 6), suggests that L. surinamensis and L. doutei are endemic to this bank. Saya de Malha Bank is a large (ca. 4,000 km-) flat plateau arising abruptly from the Indian Ocean sea-floor, from depths in excess of 4,000 m. Its shallowest parts reach as shallow as 10 m, with the large summital area within the 200 m isobath. Together with the Seychelles Page 162 THE NAUTILUS, Vol. 105, No. 4 Figure 4. Lijria doutei n.sp. Holotype MNHN. Height 83.5 Rank to the northwest and Nazareth and Cargados Ca- rajos Banks to the south, it forms the Mascarene Plateau. The Seychelles Bank is composed of late Precambrian granitic rocks and is sometimes called the Seychelles Microcontinent. On the other hand, Saya de Malha, Naz- areth and Cargados C^arajos Banks have been formed by the Reunion hotspot, Saya de Malha being the oldest in the arc. It dates back to the Paleocene, while the car- bonate rocks on Nazareth Bank are not older than Eocene (Backman et ai, 1988). Saya de Malha and Nazareth are separated by a sill shallower than 500 m and are only 200-250 km apart. Saya de Malha is separated from the Figure 5. Lijria doutei n. sp. Protoconch of holotype. Scale line 2 mm. Seychelles Bank by depths greater than 2,000 m and a distance of about 650 km. Distances to other Lyria-in- habited continental shelves are: Saya de Vlalha-Mada- gascar, 1,200 km; Saya de Malha-Kenya, 2,000 km; Saya de Malha-Sri Lanka, 2,700 km. For shelf-living volutes with large, crawl-away juve- niles, this means that Saya de Malha and Nazareth Banks are very isolated structures that are not easily colonized. However, because they have never been connected to a land mass since Lyria appeared in the Eocene, only the dispersal of demersal lecithotrophic larvae can account for colonization of such isolated oceanic banks. It is there- fore remarkable that Lyria surinamensis has one of the largest protoconchs in the genus, and that its concholog- ical affinities appear to be with species geographically remote: L. lyraeformis (Kenya), L. leslieboschae Em- erson & Sage, 1986 (Arabian Peninsula), L. cloveriana Weaver, 1963 (Sri Lanka). We hypothesize that the fol- lowing scenario may have taken place: 1. Saya de Malha is colonized by an ancestral Lyria species with smaller, lecithotrophic, demersal larvae, having a broad Indian Ocean distribution; 2. The species evolves intracapsular metamorphosis with large crawl-away ju\eniles (this is a general trend in volute evolution and has been noted, e.g., by Hansen (1983) in American Paleogene volutes), demersal dispersal is lost; 3. Geographical isolation en- hances genetic isolation, and morphological divergence accumulates through time into different species. Lyria anna (Lesson, 1835) is another volute endemic to this part of the Indian Ocean. The origin of this species has long remained mysterious (Weaver & Dupont 1970), but there are indications that it probabK originates from Mauritius (Richards, 1990), nearb) areas NE of Mauritius in 170-230 m (Anonymous, 1984) and/or Cinq Rrandons p. Bouchet and P. Bail, 1991 Page 163 50« I SOMALI BASIN io» 20' 60» I SEYCHELLES \ SAYA DE MALHA MASCARENE BASIN CARGADOS CARAJOS \r Mauritius Rodi'iguez Reunion Figure 6. Simplified bathymetric map of the southwestern Indian Ocean banks, showing the locaHties mentioned in the text Stippled area, 0-100 m; thin intermediate line, 200 m isobath; thick line, 500 m isobath. Scale line 500 kilometers. shoals on the Cargados Carajos Bank of the Mascarene Plateau (unverified information from local Mauritius col- lectors and dealers). Lijria anna is conchologically not closely related to L. siirinamensis and indicates that the colonization scenario above has occurred several times in the family Volutidae. We predict that more species of Volutidae will be found on banks of the Mascarene Plateau, including the Seychelles Bank. Finally, it is noteworthy that, beside Lyria siirina- mensis and Murex surinamensis, Conus primus Rockel & Korn, 1990 was very recently described from Saya de Malha Bank, at depths of 80-98 m. It also has a paucis- piral protoconch indicating non-planktotrophic larval development. LITERATURE CITED Anonymous. 1984. Lyria (Harpeola) anna live dredged. La Conchiglia 184/185:22. Backman, J. et al. 1988. Introduction. Proceedings of the Ocean Drilling Program, Initial Reports 115:5-15. Barnard, K. H. 1963. Contributions to the knowledge of South African marine Mollusca. Part IV. Ann. S. Afr. Mus. 47(2): 201-360. Bavay, A, 1913. (no title). Journal of Conchologv 14(4):98- 99. Bouchet, P. 1990. Systematics of Plicoliva with description of a new subfamily (Gastropoda; Volutoidea). Archiv fiir Molluskenkunde 120(1/3):1-10. Bouchet, P. and G. Poppe. 1988. Deep water volutes from the New Caledonian region, with a discussion on bioge- ography. Venus 47(1): 15-32. Hansen, T. 1983. Modes of larval development and rates of speciation in early Tertiary Neogastropods. Science 204; 501-502. Okutani, T. 1982. .\ new genus and five new species of gas- tropods trawled from off Surinam. Venus 41(2):109-120. Ponder, W. F. and E. H. Vokes. 1988. A revision of the Indo- West Pacific fossil and Recent species of Murex s.s. and Page 164 THE NAUTILUS, Vol. 105, No. 4 Haustelhtm (Mollusca: Gastropoda: Muricidae). Records of the .Australian Museum, suppl. 8:1-160. Poppe, G. 1987. New Volutes since 1970, part 1. La Con- chiglia 218/219:8-22. Richards, A. 1990. Mauritius, He de France. American Con- chologist 18(4):8-10. Rockel, D. and W. Korn. 1990. Conus species from the West- ern Indian Ocean, dredged by Soviet biologists. Acta Con- chyliorum 2:45-49. Takeda, M. and T. Okutani. 1983. Crustaceans and moUusks trawled off Suriname and French Guiana. JAMARC, To- kyo. 356 p. Vokes, E. H. 1990a. What ever happened to dear old Murex? .American Conchologist 18(l):3-7. Vokes, E H 1990b Cenozoic Muricidae of the Western At- lantic region. Part VIII Murex s.s . Haustelhtm. Chico- reus, and Hexaplex. .Additions and corrections. Tulane Studies in Geology and Paleontology 23(1-3): 1-96. Weaver, C. S. and J. E. DuPont. 1970. Living Volutes. A monograph of the Recent Volutidae of the world. Dela- ware Museum of Natural History, Greenville. 375 p. THE NAUTILUS 105(4):165. 1991 Page 165 Mistaken Localities for Some Shells "from Surinam" Takashi Okutani Tok\o l'ni\ersit\ of Fisheries 4-5-7, Konan, Miiiato-ku Tokvo 108. Japan Dr. Philippe Bouchet, of the Museum national d'Histoire naturelle, Paris, and Dr. William K. Emerson, of the American Museum of Natural History, New York, co- incidentulK alerted me that the type locality reported for DallivoUita surinamensis was incorrect. They had each recently obtained specimens from the Indian Ocean that were identical to D. surinamensis. Both suggested that the correct locality was not ir03'N, 61°01"W but rather ITO.S'S, 61°0rE. In publishing a survey report on shrimp trawl grounds based on samples provided b\ the Japan Marine Fisheries Research Center (JAMARC)' (Okutani, 19S3b), I discov- ered several undescribed species of gastropods and bi- valves in the material that had been sent to me for study. A total of ten new species and one new genus were described based on this material (Okutani, 1982, 1983a). After being alerted b\ Drs. Bouchet and Emerson to the problem with the locality data, I inquired of JAMARC as to the origins of these samples. JAMARC had conducted a bottom-fish survey on the Saya de Malha Bank in the Indian Ocean in December of 1978, and a survey of shrimp grounds off Surinam and French Guiana from June of 1979 to March of 1983. Although different vessels were used for each of these surveys, the frozen samples were stored in the same re- frigerated warehouse when the vessels returned to port. Each frozen block was labeled with precise data as to date, position, depth, haul number, etc. Among the large series of samples, the designations N, S, E, W, were sometimes omitted from the labels, although latitude and longitude were always given. I was not aware of the contamination of the samples as the numerical values of the latitudes and longitudes were so close to each other. In reviewing the records, I found that species described on the basis of material collected prior to 1979 were all from Indian Ocean samples. These ta.xa, together with their correct type localities, are: Murex (Murex) surinamensis Okutani, 1982 Type locality: I0°46'S, 6r32'E, in 94 m. Dallivohita surinamensis Okutani, 1982 (now Lyria suri- namensis, see Bouchet and Bail, 1991) Type locality: 11°03'S, 6I°01'E, in 128 m. Pteria pictura Okutani, 1983a Type locality: 10°34'S, 61°35'E, in 97 m. Spondylus jamarci Okutani, 1983a Type locality: 11°03'S, 6r01'E, in 128 m. The following ta.xa are from the continental shelf or slope off Surinam or French Guiana as originally pub- lished: Plicifusus jamarci Okutani, 1982 Buccinofusus surinamensis Okutani, 1982 Volutomitra bayeri Okutani, 1982 Barbarofusus chocolatus Okutani, 1983a Sarepta surinamensis Okutani, 1983a Portlandia nigromaculata Okutani, 1983a It is hoped that this note will clarify the doubts posed by Ponder and Yokes (1988), Yokes (1990) and Bouchet and Bail (1991). LITERATURE CITED Bouchet, P. and P. Bail. 1991 Mistaken geographical coor- dinates: the saga of Lyria surinamensis and a new species. The Nautilus 105(4):159-164. Okutani, T. 1982. A new genus and five new species of gas- tropods trawled from off Surinam. Venus 41(2)109-120, Okutani, T 1983a. Four new species of bivalves and a new- gastropod trawled from off Surinam. Venus 42(1 );17-2.5. Okutani, T. 1983b, Moliusks, In. Takeda, T, and T, Okutani, Crustaceans and Moliusks traw led off Suriname and French Guiana. Japan Marine Fisher\ Resources Center, Tokyo, p. 189-343. Ponder, W. F. and E. H. Yokes. 1988. A revision of the Indo- Paciflc fossil and Recent species of Murex ss. and Haus- tellum (Mollusca: Gastropoda: Muricidae). Records of the Australian Museum. Supplement 8: 1-160. \dkes, E. H. 1990 What ever happened to dear old Murex? American C:onchoiogist 18(l):3-7. THE NAUTILUS 105(4):166-172, 1991 Page 166 New Species of Gaza, Mirachelus, Calliotropis, and Echinogurges (Gastropoda: Trochidae) from the Northwestern Atlantic Ocean James F. Quinn, Jr. Florida Marine Research Institute Department of Natural Resources 100 Fighlh Avenue, SE St. Petersburg, FL 33701-5095, USA ABSTRACT Five new species of Trochidae from the northwestern Atlantic Ocean are described. Gaza olivacea, from the northern coast of South America, is similar to C. superha (Dall, 1S81) but differs in shell shape and characters of the outer lip and um- bilical septum. Mirachelus acanlhus. known only from Ber- muda, most resembles A/, clinocnemus Quinn, 1979, but differs in shell shape and sculpture. Calliotropis globosa, from off the Yucatan Peninsula, Mexico, Jamaica, and the northeastern coast of South ,\merica from Venezuela to Suriname, is most similar to C. actinophora (Dall, 1890) but differs in relative shell height and umbilical diameter and in sculpture. Calliotropis dentata, known only by the holotype from off Venezuela, is most similar to C. persculpta (Sowerby, 1903) from off South Africa but has a proportionately higher shell and stronger, coarser sculpture. Echinogurges tuherciilatus. from off eastern Florida, the Straits of Florida, and the northern Bahamas, is most similar to £. clavatus (Watson, 1879) but is distinguished by sculptural de- tails. Key words: Trochidae; Gaza. Mirachelus; Calliotropis; Echi- nogurges; systematics; new species. INTRODUCTION Many undescribed species were discovered during the preparation of two geographically limited monographs of western Atlantic Trochidae (Quinn, 1979, in press a). This paper presents accounts of five new species of the subfamilies Margaritinae and Eucyclinae (following the cla.ssification of Hickman & McLean, 1990). New species of the subfamilies Solariellinae and Calliostomatinae are presented elsewhere (Quinn, 1991; in press a,b; in prep- aration). Institutional abbreviations used in this paper are as follows: AMNH — American Museum of Natural History; ANSP — Academy of Natural Sciences of Philadelphia; DMNH — Delaware Museum of Natural History, Wil- mington; UF — Florida Museum of Natural History, Uni- versity of Florida, Gainesville; FSBC I — Florida Marine Research Institute, Department of Natural Resources, St. Petersburg; LACM — Natural History Museum of Los Angeles County, Los Angeles; MCZ — Museum of Com- parative Zoology, Harvard University, Cambridge; TAMU— Texas A&M University, College Station; UMML — Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami; USNM — National Museum of Natural History, Smithsonian Institution, Washington. SYSTEMATICS Family Trochidae Rafinesque, 1815 Subfamily Margaritinae Stoliczka, 1868 Tribe Gazini Hickman and McLean, 1990 Genus Gaza Watson, 1879 Type species (monotypy): Gaza daedala Watson, 1879. Gaza olivacea new species (figures 1-3) Gaza superha: Okutani, 1983:240. Gaza sp.: Hickman and McLean, 1990: fig. 51 A. Description: Shell large, attaining 38. 1 mm height, 44.9 mm width, broadly conical; whorl profiles weakly con- vex, with broadly rounded peripheries; umbilicate with umbilical septum; drab-olive to light brown, nacreous under thin outer porcelaneous layer. Protoconch usually missing. Teleoconch whorls about 6, weakly shouldered subsuturally, weakly convex with broadly rounded pe- ripheries, smooth except for microscopic spiral striae. Base almost flat, smooth except for striae like those above. Umbilicus wide, about 25% maximum whorl diameter; wall smooth, parallel to shell axis. Aperture ovate; outer lip oblique and reflected into slightly thickened, nacreous rim, tangential to periphery of previous whorl; inner lip reflected and expanded to form nacreous septum cov- ering umbilicus; columella weakh' concave in immature specimens, sigmoid in mature specimens. J. F. Quinii, 1991 Page 167 ^ ^. Figures 1-3. Caza olivacea new species, Hoiotype, I'SNM 752369, height 32.9 mm, width 39,8 mm, from N of Cabo de la Vela, Peninsula de la Guajira, Colombia, Oregon Station 5690, 12°30'N, 72°08'W, 470 m. Type material: Holotvpe, USNM 752369; 22 paratypes, USNM 859424; Oregon Station 5690, 12°30'N, 72°08'W, 470 m; 40-foot otter trawl; 10 October 1965.— 17 para- types, USNM 752390; 1 paratvpe, UMML 30.8365; 1 paratype, FSBC I 39513; Oregon Station 4301, 7°35'N, 54°13'W, 366 m; 65-foot otter trawl; 24 March 1963. Type locality: North of Cabo de la Vela, Peninsula de la Guajira, Colombia, Oregon Station 5690, 12°30'N, 72°08'W, 470 m. Other material examined: 14 specimens, USNM 752391 (+ 61 uncatalogued USNM, and duplicates in other mu- seums); Oregon Station 2026, 7°10'N, 53°07'W, 366 m; 40-foot otter trawl; 9 November 1957. — 15 specimens, USNM 752389 (-1- 200-300 uncatalogued USNM, and duplicates in other museums); Oregon Station 2005, 7°34'N, 54°50'W, 366 m; 40-foot otter trawl; 6 November 1957.— 1 specimen, USNM 752383; Oregon Station 1982, 10°00'N, 59°59'W, 457 m; 40-foot otter trawl; 3 Novem- ber 1957—8 specimens, USNM 752388; Oregon Station 5028, ir.30'N, 60°46'W, 366-439 m; 40-foot otter trawl; 22 September 1964—2 specimens, USNM 752393; Or- egon Station 5930, 15°38'N, 61°07'W, 808 m; otter trawl; 5 March 1966—19 specimens, USNM 752384 (-1- 54 uncatalogued USNM, and duplicates in other museums); Oregon Station 2351, ir31'N, 62°24'W, 338-366 m; 40- foot otter trawl; 23 September 1958. — 17 specimens, USNM 752377; Oregon Station 2780, ll''36'N, 62°52'W, 393-421 m; 40-foot otter trawl; 20 April 1960—15 spec- imens, USNM 752366; Oregon Station 4420, 11°46'N, eg'n'W, 421 m; 40-foot otter trawl; 4 October 1963 — 7 specimens, USNM 752372; Oregon Station 5692, 12°31'N, 71°58'W, 375 m; 40-foot otter trawl; 10 October 1965. — 4 specimens, TAMU 4-1737; 2 specimens, TAMU 4-0388; Alaminos Station 70A10-40, 12°40'N, 72°00'W, 622-658 m; 57-foot trawl; 18 Julv 1970—9 specimens, USNM 752373; Oregon Station 4923, 12°21'N, 72°40'W, 439-448 m; 40-foot otter trawl; 2 June 1964—13 spec- imens, UMML 30.8364; John Elliott Pilkburij Station P- 781, 11°30.1'N,73°26.5'W, 567-531 m; 10-foot otter trawl; 30 July 1968—16 specimens, USNM 752376; Oregon U Station 11252, 11°25'N, 73°56'W, 435 m; 71-foot otter trawl; 10 November 1970—3 specimens, USNM 752374; Oregon II Station 11251, ir23'N, 74°16'W, 457 m; 71- foot otter trawl; 10 November 1970. — 7 specimens, USNM 752364; Oregon II Station 10266, 11°04'N, 74°25'W, 27 m; 71-foot otter trawl; 2 December 1968. — 12 specimens, USNM 752365; Oregon II Station 11248, 11°18'N, 74°44' W, 600 m; 71-foot otter trawl; 9 November 1970.— 9 specimens, USNM 752363 (-f 100 uncatalogued USNM, and duplicates in other museums); Oregon II Station 10260, 11°03'N, 7.5°18'W, 366 m; 71-foot otter trawl; 2 December 1970. — 5 specimens, USNM 752375; Oregon Station 5722, 9°.36'N, 76°22' W, 512 m; 65-foot otter trawl; 16 October 1965—1 specimen, USNM 94991; Albatross Station 2143, 9°30'45"N, 76°25'30"W, 283 m; small beam trawl; 23 March 1884—4 specimens, USNM 752.392; Oregon Station .3584, 9°13'N, 81°30'W, 366 m; 40-foot otter trawl; 25 May 1962—20 USNM lots with localities within range indicated above. Remarks: Shells of Gaza olivacea are most similar to those of Gaza superba (Dall, 1881) but differ in being more broadly conical and in having more flattened whorl profiles and much weaker spiral striae; the outer lip is not angulate at the junction with previous whorl (figures 2, 3); and the nacreous septum covers the entire umbili- cus. With the exception of the spiral striae and umbilical septum, shells of G. olivacea differ from those of the Galapagan G. rathbuni Dall, 1890, by the same char- acters that differentiate G. olivacea and G. superba. Gaza olivacea is known from Golfo de los Mosquitos, Panama, eastward along the northern coast of South America to Suriname and French Guiana. Although the depth range for this species is 283-808 m (x = 433 m, n = 38), 87% of all collections were made in depths of 350-550 m; these analyses ignored records of 15 fms (27 m; Oregon II Station 10266) and 25 fms (46 m; Oregon II Station 11275) because they are most likeK' errors for 150 fms and 250 fms, respectively. These depths are shallower than those for G. superba (range = 360-1,006 m; x = Page 168 THE NAUTILUS, Vol. 105, No. 4 Figures 4-6. Mirachelus acanthus new species. 4. Paratype, DMNH 187590, height 3.45 mm, width 2.95 mm, trom oH Bermud.i. .3. Same specimen, aperture, 27 x . 6. Same specimen, obHque view of protoconch and first whorl, 108 x. 556 m, n = 78), for which 86% of all collections have been made between 440 m and 680 m (Quinn, in press a). The geographic ranges of G. olivacea and G. superba overlap slightly only in Panama in the west, and off Grenada and Suriname in the east. A third western At- lantic species, G. fischeri Dall, 1889, has a geographic range that encompasses the ranges of both G. olivacea and G. superba, but occurs in depths deeper than either of those species (80% of lots from 550-825 m, x = 692 m; n = 33), although G. fischeri and G. superba have sometimes been collected together (Quinn, in press a). Like shells of G. superba, almost all shells of G. olivacea have a small, neat hole replacing the protoconch and leading into the umbilicus, and a narrow channel at the junction of the umbilical rim and outer lip (figure 3), both caused by a commensal polychaete worm that lived in a mud tube within the umbilicus (Quinn, in press a). Subfamily Eucyclinae Koken, 1897 Tribe Ch'ilodontini Wenz, 1938 Genus Mirachelus Woodring, 1928 Type species (original designation): Calliostoma corbis Dall, 1889. Mirachelus acanthus new species (figures 4-6) Description: Shell small for genus, attaining 3.70 mm height, 2.95 mm width, conical, nonumbilicate, white, nacreous under thin outer porcelaneous layer. Proto- conch about 325 /irn maximum diameter, about one whorl. Teleoconch whorls about 5.4; first 1.5-2 whorls with strong axial lamellae, remaining strong but becoming more rounded on subsequent whorls, spaced about 0.5 mm apart on fifth whorl; very fine, oblique threads covering whorl surface; 2 spiral cords appearing near end of sec- ond whorl, abapical cord stronger and forming periph- ery; weak angulation or spiral cord sometimes present at suture; strong, conical tubercles at intersections of axial and spiral sculpture. Rase convex, with 4-5 strong spiral cords; spiral cords undulate or weakly beaded abaxially, more distinctly beaded adaxially. Aperture oblique, ovate, thickened within, with 8-9 weak ridges on thickening; columella short, thickened, weakly inflated medially. Type material: Holotvpe, DMNH 187589; 3 paratypes. DMNH 187590; 1 paratype, USNM 860246; Lightbourn- Guest Northstar Expedition Station 582, off Castle Har- bour, Bermuda, 100 m. Type locality: Off Castle Harbour, Bermuda, in 100 m. Remarks: Shells of this species are most similar to those of Mirachelus clinocnemus Quinn, 1979, but differ in having weaker but sharper spiral cords, the peripheral one stronger and more projecting than the one just adapi- cal; in having the axial lamellae sharper and more wideK spaced (0.5 mm apart in M. acanthus, 0.25 mm apart in M. clinocnemus); and in having more prominent, sharply conical tubercles on the peripheral and supra- peripheral spiral cords. The shells of M. acanthus are also slightly broader than those of M. clinocnemus. Tribe Calliotropini Hickman and McLean, 1990 Genus Calliotropis Seguenza, 1903 Type species (original designation): Trochus ottoi Phi- lippi, 1844. Calliotropis glabosa new species (figures 7, 8) Description: Shell of medium size for genus, attaining 9.7 mm height, 8.7 mm width, turbinate with inflated whorls, umbilicate, white, nacreous under thin outer por- celaneous layer. Protoconch large, about 525-550 ^m maximum diameter, 1 whorl. Teleoconch whorls 6.25; first 1.5 whorls with thin axial riblets, rapidly becoming weaker on next whorl, finally becoming restricted to subsutural area on last w horl; pair of spiral threads ap- pearing on first whorl, one subsutural, one just below midwhorl and forming whorl periphery, both becoming slightly stronger on sub.sequent whorls; third spiral thread appearing on third v\ horl between suture and primary subsutural spiral thread, gradually becoming slightly J. F. Quinn, 1991 Page 169 / r- > r V y \ > m Figures 7, 8. Calliotropis globosa new species. Holot) pe, USNM 859419, height 9.45 mm, width S.70 mm, from S of Jamaica, 17°21.4'N, 77°.34.S'W, 805-1,089 m. stronger than primary subsutural spiral thread on sub- sequent whorls; fourth spiral thread similar to peripheral thread, appearing on last whorl, coincident with suture on spire whorls. Base convex, with 4 spiral cords; out- ermost cord almost smooth, cords becoming progressively more strongly tuberculate adaxially; innermost 2 cords most closely spaced; transverse rugae becoming pro- gressively stronger adaxially. Umbilicus deep, walls ax- ialK rugose and almost vertical Aperture subovate, lips thin; columella sigmoid, thin, with rather strong, rounded tooth medially in some specimens. Type material: Holotype, USNM 859419; 2 paratypes, USNM 859420; 2 paratypes, ANSP 383289; 2 paratypes, FSBC I 39515; 2 paratypes, UP 169956; 2 paratypes, MCZ 302452; 2 paratypes, AMNH 232160; 21 paratypes, UMML 30.8358; John Elliott Pillsbury Station P-i262, 17°21.4'N, 77°34.8'W, 805-1,089 m; 10-foot otter trawl; 15 July 1970. Type locality: South of Jamaica, John Elliott Pillsbury Station P-1262, 17°21.4'N, 77°34.8'W, 805-1,089 m. Other material examined: 17 specimens, UMML 30.8355; John Elliott Pillsbury Station P-604, 18°58'N, 87°28'W, 970-988 m; box dredge; 17 March 1968.— 27 specimens, UMML 30.8356; John Elliott Pillsbury Sta- tion P-605; 18°50.1'N, 87°31.5'W, 695-773 m; 10-foot otter trawl; 17 March 1968. — 10 specimens, UMML 30.5703; 11 specimens, UMML 30.8361; John Elliott Pillsbury Station P-607, 18°30'N, 87°37'W, 715-787 m; 10-foot otter trawl; 17-18 March 1968.— 3 specimens, MCZ 294845; Blake Station XVIII, 18°20'30"N, 87°16'40"W, 1,097 m; 1880.— 4 specimens, UMML 30.8360; John Elliott Pillsbury Station P-1255, 17°18'N, 78°32'W, 622-823 m; 10-foot otter trawl; 14 July 1970.— 2 specimens, UMML 30.8359; John Elliott Pillsbury Sta- tion P-1261, 17''13'N, 77°50'W, 595-824 m; 10-foot otter trawl; 15 July 1970—1 specimen, MCZ 135174; Ape, USNM 859419, height 9.4 mm, width 9.2 mm, from off Isla Orchilla, Venezuela, ll''47.8'N, 66°06.8'W, 1,052-1,067 m. coast near Banco Chinchorro, from Cay Sal Bank and Cuba, from south of Jamaica, and along the northern South American coast from Venezuela to Suriname, mostly in depths of about 700-1,100 m (range = 595- 1,829m). Calliotropis dentata new species (figures 9-11) Description: Shell of medium size for genus, attaining at least 9.4 mm height, 9.2 mm width, conical with shouldered whorls, peripherally carinate, strongly sculp- tured, white, nacreous under outer porcelaneous layer. Protoconch and much of first whorl missing. First re- maining whorl with narrow axial riblets and weak sub- sutural and peripheral spiral angulations. Second whorl largely decorticated but showing strengthening angula- tions. Subsutural angulation becoming spiral row of strong, conical, axially elongate tubercles rising above level of suture; peripheral angulation forming carina set with closely set, cog-like tubercles connected and crossed by fine spiral thread. Final whorl with second carina, similar to peripheral carina but slightly weaker, abapical to pe- ripheral carina and coincident with suture line. Whorl surface between spiral carinae flat, with fine, collabral growth lines. Base convex, with 3 strong, tuberculate spiral cords; whorl surface between innermost 2 spiral cords with weak collabral rugae that generally connect tubercles of cords. I'mbilicus about .30% maximum shell diameter, furuiel-shaped, walls with axial rugae and sin- gle weak spiral thread. Lips of aperture and columella broken. Type material: Holotype, USNM 859422; John Elliott Pillslmrij Station P-741, 11°47.8'N, 66°06.8'W, 1,052- 1,067 m; 40-foot otter trawl; 23 July 1968. Type locality: Off Isla Orchilla, Venezuela, John Elliott Pilhbury Station P-741, ir47.8'N, 66°06.8'W, 1,052- 1,067 m. Remarks: The shell of Calliotropis dentata is easily distinguished from those of all other western Atlantic Calliotropis by having the peripheral and subperipheral spiral cords with closely set, cog-like beads instead of conical tubercles. The shell of C. dentata is very similar in sculpture to that of C. persculpta (Sowerby, 1903) (see Kaicher, 1990: card 5695) from South .Africa but differs in being higher spired and in having stronger basal spiral cords and a narrower umbilicus. Genus Echinogurges Quinn, 1979 Type species (original designation): Trochus (Mar- garita) clavatus Watson, 1879. Echinogurges tubercidatus new species (figures 12-14) Echinogurges rhysus: Quinn, 1979:21, 22, figs. 39, 40. [Misidentification; not Trochus (Margarita) rhysus Wat- son, 1879.] Description: Shell small, attaining 4.6 mm height, 3.9 mm width, conical with shouldered whorls, strongly sculptured, umbilicate, white, nacreous under thin outer porcelaneous layer. Protoconch about 375 fim maximum diameter, bulbous, 1 whorl. Teleoconch whorls 5.5, tu- bular, carinate, first whorl with strong, thin axial riblets that become weak collabral ridges on subsequent whorls; pair of spiral ridges appearing on second whorl, one subsutural, one at midwhorl forming carinate whorl pe- riphery; third tuberculate spiral ridge present on body whorl, coincident with suture on spire whorls; strong, conical tubercles at intersections of axial and spiral sculp- ture. Base convex, rounding into umbilicus, with 4 smoothish to finely tuberculate spiral cords. L'mbilicus narrow, pore-like, wall with axial rugae. Aperture sub- circular, lips thin, inner lip weaklv reflected into um- bilicus; columella concave, thin, rounding smoothly into outer lip. Type material: Holotvpe, USNM 330606; 4 paratypes, USNM 859426; Albatross Station 2654, 27°57'3'0"N, 77°27'30"W, 1,207 m; large beam trawl; 2 May 1886.— 13 paratypes, MCZ 302536; Atlantis Station 2993, J. F. Quinn, 1991 Page 171 v^ ^ Figures 12-14. Echinogurges tuberculatus new species. 12, 13. Paratype, UMML 30 8095, height 4,0 mm, width 3.1 mm, from southern Straits of Florida, Cerda Station G-1106, 24°02'N, 81°30'W, 1,706-1,723 m. 14. Protoconch of paratype (USNM 859426) from NE of Little Bahama Bank, Albatross Station 2654, 27°57'30"N, 77°27'30"W, 1,207 m, 97 x , 33°19'N, 80°44'W, 1,061 m; 15 March 1938.— 4 para- types, USNM 859426; Albatross Station 2415, 30°44'N, 79°26'W, 805 m; large beam trawl; 1 April 1885.— 2 paratvpes, UMML 30.8054; Gerda Station G-368, 24''03'N, 8riO'W, 961-1,016 m; 16-foot otter trawl; 15 September 1964.— 1 paratype, UMML 30.8041; Gerda Station G-130, 23°59'N, 81°10'W. 1,021 m; 10-foot otter trawl; 21 June 1963—1 paratype, UMML 30.8163; Ger- da Station G-859, 23°54'N, 8r57'W, 1,161-1,200 m; 10- foot otter trawl; 29 August 1967. — 2 paratypes, USNM 438284; 10 miles N of Cuba, 1,427 m; J. B. Henderson coll. — 2 paratypes, MCZ 7584; Blake Station, Yucatan Channel, 1,171 m. Type locality: Northeast of Little Bahama Bank, Al- batross Station 2654, 27°57'30"N, 77°27'30"W, 1,207 m. Other material examined: 1 specimen, UMML 30.8095; Gerda Station G-1106, 24°02'N, 81°30'W, 1,706-1.723 m; 10-foot otter trawl; 29 April 1969. — 4 specimens, MCZ 135026; Atlantis Station 2988, 23°15'N, 79°57'W, 695 m; 14 March 1938. Remarks: Shells of Echinogurges tuberculatus are most similar to those of E. clavatus (Watson, 1879) but differ in having only a single peripheral spiral ridge; weaker, more widely spaced axial sculpture; and fewer basal spi- ral cords. Specimens of E. tuberculatus were misiden- tified as £. rhysus (Watson, 1879) by Quinn (1979). Sub- sequent examination of the types of Trochus (Margarita) rhysus Watson, 1879, revealed that T. (M.) rhysus is actually a species of Calliotropis Seguenza, 1903 (Quinn, in press a). Echinogurges tuberculatus is known only from off the east coast of Florida, the Bahama Islands, and the Straits of Florida in depths of 805-1,723 m. ACKNOWLEDGMENTS I thank the late Gilbert L. Voss (Rosenstiel School of Marine and Atmospheric Science, University of Miami) for access to the collections under his care and for per- mission to describe the new species in that material. Richard S. Houbrick and the late Joseph Rosewater (both USNM), and Riidiger Bieler (formerly DMNH) kindly allowed me to examine types and other material. Russell Jensen (DMNH) brought the Bermuda specimens to my attention. Marjorie Myers, Llyn French, and Judy Leiby provided valuable word-processing and editorial assis- tance. Thomas H. Perkins and William G. Lyons (both Florida Marine Research Institute, Department of Nat- ural Resources), and two anonymous reviewers provided valuable critiques of previous drafts of this paper. LITERATURE CITED Dall, W. H. 1881. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mex- ico, and in the Caribbean Sea, 1877-79, by the United States Coast Survey steamer "Blake," Lieutenant-Com- mander C. D Sigsbee, L'.S.N., and Commander J. R Bart- lett, U.S.N., commanding. W. Preliminars report on the Mollusca. Bulletin of the Museum of Comparative Zool- ogy, Harvard University 9(2):33-144. Dall, W H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U. S. Coast Survey steamer "Blake , Lieut -Commander C D Sigsbee, U S N , and Commander J R Bartlett, US N , commanding. XXI.X. Report on the Mollusca Part 2, Gas- tropoda and Scaphopoda Bulletin of the Museum of Com- parative Zoology, Harvard I'niversity 18:1-492. Dall, W. H. 1890. Scientific results of explorations by the U. S. Fish Commission steamer "Albatross '. VII. Preliminary report on the collection of Mollusca and Brachiopoda ob- tained in 1887-88. Proceedings of the United States Na- tional Museum 12(773):219-362. Hickman, C. S. and J. H. McLean. 1990. Systematic revision and suprageneric classification of trochacean gastropods. Natural History Museum of Los .Angeles Count)-, Science Series, No. 35, vi -I- 169 p. Kaicher, S. D. 1990. Card catalogue of world-wide shells Pack #56 — Trochidae \'. PrivateK published, St. Peters- burg, FL. Okutani,"T. 1983. Mollusks In: Takeida, M and T Okutani Crustaceans and mollusks trawled off Suriname and French Guiana. Japan Marine Fisherv Resource Research Outer, Tokyo, p. 187-354. Page 172 THE NAUTILUS, Vol. 105, No. 4 Philippi, H. .\. 1844. Enunicratio molhist-orum Siciliae cum viventium turn tellure tertiaria fossiliuni. quae in itinirt- suo observavit, Vol 2 Halis Sa.\oiuini, iv + 303 p. Quinn, J. F., Jr. 1979. Hinlogical rt-sults of the University of Miami deep-.sea e.xpcditioiis 130 The systematics and zo- ogeograph\ of tlie gastropod family Trochidae collected in the Straits of Florida and its approaches. Malacologia 19(1); 1-62. Quinn, J. F., Jr. 1991. Larucllilwchus. a new genus of So- lariellinae (Gastropoda: Trochidae), with descriptions of six new species from the western .Atlantic Ocean The Nautilus 105(3):81-91. Quinn, J. F., Jr. In press a The Trochidae of the Gulf of Mexico (Prosobranchia: .'Vrcliaeogastropoda), Memoirs of the Hourglass CruLses. Quinn. J. !■' , Jr In press b. New species of Soliiriclla (Gas- tropoda: Trochidae) from the uestern .\tlantic Ocean The Nautilus. Seguenza, G. 1903. Molluschi poco noti dei terreni terziarii di Messina Societa Geologica Italiana, Bolletino 21:455- 464. Sowerby, (i. H , 111. 1903 Mollusca of South .Africa Marine Investigations in South .\frica 2:213-232. Watson, R. B 1879. Mollusca of H.M.S. Challenger Expe- dition. III. Trochidae, viz. the genera Seguenzia. Basilissa, Gaza, and Bernhix. Journal of the Linnean Society of Lon- don, Zoology 14:586-605. Woodring, W. P. 1928. Miocene moUusks from Bowden, Ja- maica, II. Gastropods. Carnegie Institution of Washington, Publication 385:1-564, THE NAUTILUS 105(4):173-174, 1991 Page 173 Acroloxus coloraderisis (Henderson), a Rare North American Freshwater Limpet Andrew J. Paul Hugh F. Clifford Deparlment of Zoology Universit) of Alberta, Edmonton, .'Alberta, Canada T6G 2E9 The freshwater limpet family Acroloxidae (Basomma- tophora) contains seven species, all in the genus Acro- loxus. One, A. laciistris (Linnaeus), is common in lentic habitats of much of Europe and parts of Asia. Five spe- cies, A. improvistts (Polinski), A. macedonicus (Had- zisce), A. kobelti (Dybowski), A. sibiricum (Gerstfeldt), and A. froscheli (Dybowski), appear to be endemic to a few ancient lakes (e.g.. Lake Ochrid and Lake Baikal) of Yugoslavia and U.S.S.R. (Clarke, 1973). Only one spe- cies, A. coloradensis (Henderson), is found in North America. It is readily distinguished from other fresh- water limpets by its spine-like apex, which is directed posteriorly towards the left margin (figure 1). Acroloxus coloradensis has hitherto been reported from only seven North American localities. Four of the records are from Rocky Mountain lakes: Colorado (Walker, 1925), Alberta (Moziey, 1930), Montana (Russell and Brunson, 1967), and British Columbia (Clarke, 1981). The other three records are from ponds in Ontario and Quebec (Clarke, 1970). We recently collected Acroloxus coloradensis speci- mens from one location of the Beaver River in Alberta, Canada. Voucher specimens are deposited in the Ca- nadian Museum of Nature, Ottawa, Ontario, Canada, catalog number #92900. This is the first report of A. coloradensis collected from a stream and apparently the first report of any of the seven Acroloxus species from a lotic habitat. The Beaver River runs through the southern limit of mixed boreal forest and the northern limit of agricultural activity in northeastern Alberta and is a trib- utary of the Churchill River, which empties into Hudson Bay.' We sampled various sites of the Beaver River, but Acroloxus coloradensis was collected at only one loca- tion: where the Beaver River is crossed bv a small road bridge south of Highway 55 (54°32'N, lli°52'W). Spec- imens were collected to a depth of about 0.8 meters. Samples were collected between 30 May and 2 August 1989 by kick samples taken with a dip net (mesh size: 500 fim) and by inspecting substratum taken from the river. The width of the Beaver River at the collecting site was 17.5 m; water temperatures ranged from 17 °C on 30 May 1989 to 25 °C on 20 July 1989. There were no riffle regions in the vicinity, and current velocity was undetectable using a Price AA current meter. Free-float- ing and submergent macrophytes were common at the sampling site. The river's substratum immediateK up- stream and downstream of the collecting site (the bridge) consisted of mud and plant detritus with rooted mac- rophytes near the banks of the river. However, A. colora- densis was collected only where the bridge crossed the stream, and here the substratum consisted of gravel, peb- Figure 1 . Scanning electron micrographs of Acroloxus colora- densis from the Beaver River, .'\lberta, Canada, 1. Side view. 2. Top view. Page 174 THE NAUTILUS, Vol. 105, No. 4 bles and large cobbles; presuinabK the large substrata was introduced when the bridge was constructed. Other gastropods collected from this location were Valvata sin- cera helicoidea Dall; Menetiis cooperi Raker, Prome- netus exacuous exacuous (Say), Armiger crista (Linnae- us), Lijmnaea stagnaiis jugularis (Say), Stagnicola spp., Helisoma spp., and Gtjraulus spp. Bryce (1970) postulated that Acroloxits coloradensis, based on its known distribution at tliat time, was a cold- water stenotherm, limited to Rocky Mountain lakes. More recent records of the limpet in Ontario and Quebec (Clarke, 1970), Colorado (VVu 1989), and our finding indicate that A. coloradensis is not restricted to Rocky Mountain lakes. The large temperature variations (17°- 25 °C) experienced over the summer by A. coloradensis in the Beaver River indicate that it is not a cold-water stenothermal species. Clarke (1973) and Burch (1982) suggest A. coloraden- sis might be a rare species with a relic distribution. Its discovery in the Beaver River is not inconsistent with this idea, and we can only speculate on its occurrence in the Beaver River of Alberta, Canada. Possibly A. colora- densis will eventually be recorded from Saskatchewan and Manitoba, indicating a wide but rare distribution across Canada on both sides of and in the Cordillera. The southerly populations of A. coloradensis, e.g., in Montana and Colorado, might now be restricted to in and near the Cordillera. ACKNOWLEDGEMENTS We would like to thank Dr. Arthur H. Clarke, Eco- search, Inc., Portland, Texas, for verifying the identifi- cation of A. coloradensis. Karen A. Saffran (University of Alberta) provided technical assistance; Ronald Koss (University of .Alberta) took the scanning electron mi- crographs. Financial assistance was provided to .^JP and HFC by the Natural Sciences and Engineering Research Council of Canada. LITERATURE CITED Bryce, G. W. 1970. Rediscovery of the limpet Acroloxus coloradensis (Basommatophora: .\croloxidae) in Colorado. The Nautilus 83:105-108. Burch, J B 1982. Freshwater snails (MoUusca: Gastropoda) of North America. United States Environmental Protection Agency, Cincinnati, OH, 294 p. Clarke, A, H. 1970. On Acroloxus coloradensis (Henderson) (Gastropoda, Basommatophora) in Eastern Canada. Pub- lications in Zoology, No, 2, National Museums of Canada, 13 p. Clarke, A. H. 1973. The freshwater molluscs of the Canadian Interior Basin. Malacologia 13; 1-509, Clarke, A, H, 1981, The freshwater molluscs of Canada. Na- tional Museum of Natural Sciences/National Museums of Canada, Ottawa, Ontario, 446 p. Mozley, A, 1930. Reports of the Jasper Park Lakes investi- gations, 1925-26. The Mollusca of Jasper Park. Transac- tions of the Royal Society of Edinburgh 56:647-669. Russell R, H, and R. B. Brunson, 1967. Acroloxus coloradensis from Montana, The Nautilus 81:33, Walker, B, 1925. New species of North American .\ncylidae and Lancidae, Occasional Papers of the Museum of Zo- ology, University of Michigan 165:1-13, Wu, S.-K. 1989. Colorado freshwater mollusks. Natural His- tory Inventory of Colorado, No. 11:1-117. w, t ■J 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. Manuscripts: Each original manuscript and accompa- nying illustrations should be submitted in triplicate. Text must be typed on one side of 8'/2 x H inch white paper, double spaced throughout (including literature cited, ta- bles and figure captions), with at least 1 inch of margin on all sides. 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