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Walle VELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California Volume 20 July 1, 1977 to April 1, 1978 fi Nn AeA cL 4 a ih eR A) Vol. 20; No. 4 TABLE or CONTENTS A color variant of Conus bulbus Reeve SING CKGERY ee hee ceetta ecto thee ecenterirseertetnatenzeer 180 Activity of the gastropod mollusk Olivella biplicata in response to a natural light/dark cycle LD NGean oy) AYA 124 8 60 05 010 XSI een eter con eee 137 Additional molluscan records from Bahia de Los Angeles, Baja Califonia Norte Forrest L. PoorMAN & LeRoy H. PoorMAnN ........... 369 Air drying Giemsa technique for gastropod chromosomes IRR J RUNSY Noy al Cr Ore DSi le rr o 386 A new sea floor oasis Lie, NY RAYS I 6 ToT Nise rete en tee ete ee 179 A new species of Anachis (Gastropoda : Columbellidae) from the Eastern Pacific TR AW VV ETIT NE Vso rea ee tala riatilS aden, 205 A new species of chiton from the Aleutian Islands (Mol- lusca : Polyplacophora ) PAIN ONTO |e ERRETRA, Geren etecteatcstnstseitetcrertinesnesnresrait 27 A new species of Coryphella (Nudibranchia : Flabellin- idae) from Santa Barbara, California RosBert K. Cowen & DAVID R. LAUR ures 292 A new species of Humboldtiana (Helminthoglyptidae) from Coahuila, Mexico RicHarp W. FuLLINGTON & Ear G. ZIMMERMAN 134 A new species of Lithophaga (Bivalvia) from the Great Barrier Reef, Australia IRCAR TED WIKDERNOANIN fasetet nace ncrctccccnntactcnenetlemae 151 A new species of Serpulorbis (Gastropoda : Vermetidae) from South Africa IROGERY Nes EDU GEES) eccencrttratcemeert eran een: 288 A new species of Subcancilla (Gastropoda : Mitridae) from the Gulf of California RAC VVSELIEI NES prec eee tecc stom eters eet ea es 52 A new species tentatively referred to Antonietta, Antont- etta janthina, from Japan (Nudibranchia : Eolido- idea : Facelinidae) KIKUTARO BABA & IWAO HAMATANL o.eccsncscresnencsnsiee 9 A note on changes in marine intertidal fungus taxonomy IDAVIDUING LINDBERG jose tr neta ee een ek 399 Another cephalopod from northern California (Mollusca: Cephalopoda) ROBERT Res AL MADGE test tere Merete sie Asta 299 A rectification of a statement regarding the Lamarckian collection in the book “Murex shells of the world” by George E. Radwin & Anthony D’Attilio ANN TELON Va DVATTILIO Nance cet cates, 399 THE VELIGER Page III A sand-dwelling Elysia from Guam (Opisthobranchia: Sacoglossa ) Cs CARESON@&g Ep) a blORE, fe enennccensecs 14 Behavioral studies of gastropod feeding. ADIAN GET ERIN phe eesti iterate eunets VOL INS 58 Chiton fauna of the Galapagos Islands ALLYN G. SmiTH & ANTONIO J. FERREIRA occu 82 Cholinergic and serotonergic control of buccal muscle in Aplysia K. R. Wess, J. CoHEN & I. KUPFERMANN ns 60 Clonal variation in the parthenogenetic snail Campeloma decisa (Viviparidae) R. K. SELANDER, E. D. Parker, Jr. & R. A. BRowNE 349 Cypraea: a list of the species. III JERRYS DONOHUE ere et eae Nene nna teten eae 159 Deshayes types in the National Museum, Paris PEWALAR BRAT CHER py WitorRe Neat tat Me ere dens Direct development in the intertidal gastropod Batillaria zonalis (Bruguiére, 1792) Sytvia B. YAMADA & CHANDRA S. SANKURATHRI 179 Effects of cornstarch and dextrose on oysters KENNETH W. TurGEON & DExTER S. HAVEN ........ 352 Feeding in gastropod mollusks: Behavioral and neuro- physiological substrates — A symposium HENTAI (Go EST DE RIN gress ees retreats 54 Feeding in Helisoma trivoluis GoRGS: KANEKOVWE)9 4) Dig CATER cies cscs decctiecteees 57 Feeding in Navanax inermis, neurophysiological aspects D. C. Spray, M. E. Spma & M. V. L. BENNETT ..... 59 Flight responses of two species of intertidal gastropods (Prosobranchia : Trochidae) to sympatric predatory gastropods from Barbados DantEt L. HorFMAN & PAUL J. WELDON ....-.0 361 Freeze-etching studies of pulmonate spermatozoa NAA) Ws 9159 UNS) Oy 6, eee cic ee ee ere 71 Further field notes on the behavior of Aplysia dactylomela BUST OBAC EG se escola ochc ces rests ssteoanianttten 359 Glans carpenteri vs. Glans subquadrata: The rules con- cerning renamed transient secondary homonyms ESUGE NERV COAN freee area aan prensa ries 63 Hiatella solida (Sowerby, 1834) (Mollusca : Hiatellidae) on Concholepas concholepas (Bruguiére, 1789) and other substrates Cartos GALLARDO S. & CECILIA OSORIO R, eres 274 Homing in Urosalpinx cinerea in response to prey effluent and tidal periodicity AVA TD VL RIA Te Teeter erates ctatnitscarteettsomreensee 30 Page IV Integration of electron scan and light imagery in study of molluscan radulae CAROLE) St EIIGK MAN eet eee rea saeesgenccecremmeteamses I Internal variables controlling food consumptionin Aplysia AW SUSS WEIN ote cere scenes tors tener eae aa 56 Mechanisms underlying “‘singleness of action” in the feed- ing behavior of Pleurobranchaea californica (Mac- Farland, 1966) JEFFREY L. RAM & WILLIAM J. DAVIS usec 55 Neurobehavioral studies on feeding in the terrestrial slug Ariolimax californicus. DS ME SENSE MAN coc. ds5. essen cinch mina ance 56 Neuronal basis of Pleurobranchaea feeding MELODY ViS)SIEGLERGsrcs atin sn cn eat ecnnceeh eines 59 New range extensions for chitons (Amphineura : Poly- placophora ) GEORGE) AY PUAN SEL MAN erasers narerecesrseerercerecsetes 62 Nomenclatural notes on Hinnites gigantea (Gray) BARRY ROTH & EUGENE V. COAN unscsssessssessistsstineinen 297 Notes on a California hybrid Haliotis (Gastropoda : Hali- otidae ) IROBERT Reps WATEMADG Beer erent nt arena 37 Notes on sea hares of South Texas (Gastropoda : Opistho- branchia ) Nep E. StRENTH & JAMES E. BLANKENSHIP ........... 98 Notes on the opisthobranch fauna of south San Francisco Bay Davip W. BEHRENS & MERRITT TUEL cccscscsssssssensene 33 Notes on the spawning and egg capsules of two proso- branch gastropods: Nassarius tiarula (Kiener, 1841) and Solenosteira macrospira (Berry, 1957) Roy: SSsHOUSTON) 22cm, 367 Observations on feeding, chemoreception, and toxins in two species of Epitonium SIGRID’ SALO} cect csscetoceeoceda eum anatca te ane age 168 Observations on the copulatory behavior of Littorina rud- is (Maton) and Littorina nigrolineata (Gastropoda: Prosobranchia ) Di G. RAFFABLET |2.)iiiieiciatinak nant cack ean eaten 75 Predatory behavior in Navanax inermis IMJ MURRAY, cacaiacacnitataet one ce ce oe ee ae 55 Prey preferences of carnivorous intertidal snails in the Florida Keys Russet E. INGHAM & JAMES A. ZISCHKE ........... 49 Publication dates of Bergh’s 1879 papers describing A- merican chromodorids ROBERT BURN acs ccaceataser aree eee en as 2098 Reevaluation and new description of the genus Bittium (Cerithiidae ) RICHARD 3S: FLOW BRICK 22 rcateatissetstcreisstvineuseraaionncnias 101 THE VELIGER Vol. 20; No. 4 Reproductive biology of Colus stimpsoni (Prosobranchia: Buccinidae). I. Male genital system Davin: Le WEST i. $282. ec ccaants ese 266 Size and age-specific predation by Lunatia heros (Say, 1822) on the surf clam Spisula solidissima (Dillwyn, 1817) off western Long Island, New York Davip R. FRANZ siccetececccctunidesaca lac eee 144 Some aspects of spatfall of the New Zealand rock oyster during 1974 P. DINAMANI & P’Ag LENZ 320.0. 7 Some emergent problems in neural control of molluscan feeding, a chairman’s summary M.V. Li BENNETT ooo 60 Soviet contributions to malacology in 1976 Morris K. JACOBSON & KENNETH J. BOSS oeomennun 390 Spawning and early life history of Murex pomum Gmelin, 1791 EuNA A. Moore & FINN SANDER oecssssssnssststsnssnsnsne 251 Substrate angle, movement and orientation of two sym- patric species of limpets, Collzsella digitalis and Col- lisella scabra. LINDA'S. COLLINS ct. cic.censneen sean eee Subtidal abalone populations in an area inhabited by sea otters Joun Cooper, Mark WIELAND & ANSON HINES 163 Tambja and Roboastra (Mollusca : Opisthobranchia) from the Gulf of California and the Galapagos Is- lands WESLEY M. FARMER ..3i5cciageu eee ee 375 The Chromodoridinae nudibranchs from the Pacific coast of America. — Part I. Investigative methods and supra-specific taxonomy HANS: BERTSOH, csnscetns cee nnn ee 107 The Chromodoridinae nudibranchs from the Pacific coast of America. — Part II. The genus Chromodoris Hans: BERTSGH ws.sct cde eet eee cee ee ee 307 The date of publication of Anton’s “Verzeichnif{ der Conchylien.” WALTER O. CERNOHORSKY ocscsssesssssssnenssinetatseennes 299 The development of conspecific interactions in juvenile Aplysia dactylomela Rang, 1828: an observational study I, Izyja LEDERHENDLER 2 oe een eee 173 - The effect of temperature on the distribution and biomass of Mytilus edulis in the Alamitos Bay area C. Ropert FELDMETH & MEREDYTH ALPERT. ........... 39 The effects of season on visual and photographic assess- ment of subtidal geoduck clam (Panope generosa Gould) populations LYNN (GOODWIN? 5 ssasceninteaoee euntemanermnmtectsnan ates 155 Vol. 20; No. 4 THE VELIGER Page V The family Columbellidae in the western Atlantic. Part Ila. — The Pyreninae GEORGE aE RINAD WAIN, seecttteccr rece ctsnessnrtetne essere 119 The family Columbellidae in the western Atlantic. Part IIb. — The Pyreninae (continued) (GEORGE tia sECAD WIN Iecrerieec ees eter rsceegasantasaieisccses 328 The identity of Conus fulmineus Gmelin A) RUIN EIN GER eet rceec tot ctecters aieizontesccceoticesclseetntsnss 180 The role of the editorial referee P Nem NARA EIN] cesticte satecsetc tre cc ener temece beer sisssnreeceenseer 387 The Samoan land snail genus Ostodes (Mollusca : Proso- branchia : Poteriidae) SLIZABETH-LOUISE GIRARDI occ cccscccectscirsesisitsnnnssnenen 1g! The systematic, adaptive and physiological significance of proteolytic enzyme distribution in bivalves ROBERT) Grabs PRET tea eteiten Rateeae ie Be Ue Ne ean 260 The type specimens of Ammonites hoffmanni Gabb and Melchiorites indigenes Anderson (Cretaceous : Am- monoidea ) Micuaer A. Murpuy & Peter U. Roppa ............... 78 Two new giant epitoniids (Mollusca : Gastropoda) from West Africa PHILIPPE BOUCHET & SIMON TILLIER o..ecccscscssue 345 What’s the difference? Telegraphic style versus normal style NOB LVES RAMEE EN) 2 sere ce sch tee ich ects asec teat 187 Zonation of marine gastropods on a rocky intertidal shore in the Admiralty Gulf, Western Australia, with em- phasis on the genus Nerita BREDWETHANH WIELESY occ ce terinntc ts ncaachasscomnete 279 AUTHOR INDEX PND ITC Osi VWVARRE Nip) Siegen ee eee (188) Apert, MEREDYTH see FELDMETH, C. ROBERT & — Basa, KirkuTarO & Iwao HaMATANI 9 BEHRENS, Davip W. & MERRITT TUEL 33 J oVORNTIND Se ra Wy [oe VA Wr eae elo ee Ne ee 60 see also Spray, D. C., M. E. Sprra & — IBER TS CET PEL AN Sytee e tetr nL Mec etn bok anata, 107, 307 BLANKENSHIP, JAMES E. see STRENTH, NED E. & — Boss, KENNETH J. see JacoBson, Morris K. & — BouCHET, PHILIPPE & SIMON TILLIER scscsssssscssssuceeins 345 BRYA CEDP Re DW TTEA GH pei cre ocr Orla sane Ov CO caoeccrseee 399 Browne, Rosert A. see SELANDER, RoserT K., E. Da- vis PARKER, Jr. & — IBURINADR OBER Tyee ert cette a sin har a ed tans Carison, C. H. & P. J. Horr GERNOHORSK Ya) WALTER LO stecasceisansoaineeitsiecniniavines 299 GOAN SIRIUGENE: Wie cette ce scr cai kesh 63 see also Rotu, BARRY & — CouEN, J. see Weiss, K., R. -, & I. KUPFERMANN COLTIN SH PUIN DAS ait tce tee Rr At one, Mee 43 Cooper, JoHN, Mark WIELAND & ANSON HINESs ..... 163 CowEN, RoBeErT K. & DAVID R. LAUR oreosesesissnesntinn 292 Das, C.C. see Prasap, R. « — VAT TIO AN THON Vette ee tert et ol cree eee 399 Davis, Witu1AM J. see RAM, JEFFREY L. & — Dinamanl, P. & P. A. Lenz IDONOHUES) [ERR Yirre corneas ame VARMERS “ WESEEW. Me acpsssasctstisncscsssnccitee-ancreeny ee estate 375 FELDMETH, C. RoBERT & MEREDYTH ALPERT 0 39 ERREIRAY PAN LONIO MN rcs reeae ste erate eee mene sect 27 RAIN Zs DAVID MIR ee ets une ttt on tea niece lit Aner 144 see also SmitH, ALLYN G. & — FuLuincTon, RicHarp W. & Ear G. ZIMMERMAN ..... 134 GaLiarvo S., CARLOS & CECILIA OSORIO R. ueeeeminn 274 GEEPERIN Gay A DANG etn te eee eee tater an arena 54, 58 GIRARDI, ELIZABETH-LOUISE \..ecsssecsnscssinsntnesnnenntisninnnees IQI (GOOD WIN Se LE YININ escent eee trees 155 HamatTanl, Iwao see Basa, KrKUTARO & — PANS EE MAN GEORGE VAS cc .coscccintin leet seeticcccsctuckrgcen ater 62 Haven, Dexter S. see TuRGEON, KENNETH W. & — IG KMAN A CAROLE RSs ea noes eel ree I Hines, ANson_ see Cooper, JOHN, Mark WIELAND & — Horr, P. J. see Cartson, C. H. & — HoFFMAN, Danie L. & PAUL J. WELDON ..uccsssirran 361 FIOUBRICKSURIGHARD] Soeeeie rete 101 FLOUSTONS RONG Soy rer eater eee reese ssererooh co 367 FAW GRE SS ROGER] Nepean se arses acces eles 288 IncHaAM, RussELt E. & JAMES A. ZISCHKE on... 49 Jacosson, Morris K. & KENNETH J. BOSS .eroensonn 390 IANEKON Gri Re SEIS) Be CATER coerce cctesrssneursccntersens 57 Kater, S. B. see KANEKO, C. R. S. & — IREEN AG MY RA Uo otec. cccciciccticintiection 179, 187, (306), 387 LSSTUBYDIUAINTING SN 1S (E seeercerece cree coer ere meer 151 KupFERMANN, Irvinc see Weiss, K., J. CoHEN & — Laur, Davm R. see Cowen, RoserT K. & — MEEDERELVE NDECER pele mlz) fi Age teeter stern ttecrteteeernarreeseerrer eee 173 Lenz, PA. see Drnamant, P. & - NEENIDBER Gs AUD lp Regie renal cere ceermreeremremeer 399 INAS GUST T eV op Mise etre merece tee crests, ateoesresreersenreretrre 71 MCGOWAN, JOHN Ay oesccscssssesssnssssssseenee Moore, Euna A. & FINN SANDER MurpnHy, MicHaer A. & PETER U. RODDA ercenescren 78 ENV ECT RAS oi lee fice teenie teat resneernreeerreerececronres 55 Osorio R., Cecmia see GALLARDO S., CARLOS & — Parker, E. Davis, Jr. see SELANDER, Rosert K., -, & Rosert A. BROWNE Page VI THE VELIGER Vol. 20; No. 4 LPisdb RES, IBY) ING cceteeeemteaacocosy reroria Heron arene acn 137 Spray, D. C., M. E. Spirra & M. V.L. BENNETT ........... 59 Poorman, Forrest L. & Leroy H. POORMAN 0.0.0 369.) {STonrer, Ryeee ee (189), (190), (306) Poorman, Leroy H. see Poorman, Forrest L. & — StrRENTH, Nep E. & JAMES E. BLANKENSHIP. issue 98 PRASAD, R. & C. C. DAS wonnnnnnniennniinniminnnnnann 386 SUSSWEIN,, Ais jicccnuvlon nek nn epee teen iain aaa 56 Pratt, Davip M. ...... rote 39 TALMADGE,VROBERT Re eo ue eee 37, 299 INAD WINE GEORGES Eee ieee nent eee on Ree 119, 328 Tiurr, Simon see BoucHET, PHILIPPE & — RAFFAELLY, D. Ge revesnnnnininnnnnninininnninnnninninnanannnn 15 TOBAGH; |B wean auton aeol eae aalea i ee 359 » 95 TUCKER,, JOHN Ky oie nie cecntemanel arse ee 180 Rem WROBERT GriBae re ee re ee 260 Tuer, Merrirr see BEHRENS, Davi W. & — KOnDAy PamaR WU. See MOIRA MUCH bo B= Turceon, KENNETH W. & DEXTER S. HAVEN mans 352 Ries S EUGENE Vs COAN Ge ae ten Wines, Sunumuer 2, J, Cosy 2. maasen ce SANDER, Finn see Moore, EunaA A. & — SANKURATHRI, CHANDRA see YAMADA, S. BEHRENS & — SELANDER, R. K., E. D. Parker, Jr. & R. A. BROWNE 349 SENSEMAN DMs coo cere c men a maniecn Ponte caus 56 STEGER GS NEBUODY2 ViS occa cuter eet aan etme meer eae 59 SmitH, ALLYN G. «& ANTONIO J. FERREIRA ... 82 SIMEDE VRVA TD Fa Me eet cei cute ciate acme ee ete (69), (187) Sprra, M. E. see Spray, D. C. -, & M. V. L. BENNETT WELDON, PAu J. see HorrMan, Daniet L. & - Wetts) FRED) ETHAN (ik ieee career ance 279 West; Davin Be csstet ihe eee 266 WHITNEY, Ro AG a) e et ee ee eee eee 52, 205 WieLanp, Mark see Cooper, J., —, & A. HINES YamabA, SyLviA BEHRENS & C. S. SANKURATHRI 179 ZIMMERMAN, EarL G. see FULLINGTON, RICHARD W. & — ZISCHKE, JAMES A. see INGHAM, RUSSELL E. & — — 77. VA Ys Mol. THE VELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California VOLUME 20 JuLy 1, 1977 NuMBER I ConTENTS Integration of Electron Scan and Light Imagery in Study of Molluscan Radulae. (1 Plate; 4 Text figures) CRRORERS MEIC KANE mart tata Volley Veoh qiaryreyitel Nise a itlel Jet veils Me te) a! ot A New Species Tentatively Referred to Antonietta, Antonietta janthina, from Japan (Nudibranchia : Eolidoidea : Facelinidae) . (4 Text figures) ISTIGUTAROPDAB Ans IUW/AOLELAMATANT | (is tel ao ie\ isles acon ess ee lo se A Sand-Dwelling Elysia from Guam (Opisthobranchia : Sacoglossa). (5 Text figures) COME CARESONGE EM y ORFs Miriam iaticrs ee) or oie y oie ol Me Wie a) wie Me SAL Some Aspects of Spatfall of the New Zealand Rock Oyster During 1974. (4 Text figures) PRD INAMAN ERMA LOE NZ ashe tes nem iwes cMtreiirs . cede liis ciel Mey tet Vea. del ao A New Species of Chiton from the Aleutian Islands (Mollusca : Polyplacophora). (1 Plate; 1 Text figure) LAN EONION)| ey PERREIR AME iran maramn en tera) leoioll oct ls sipete tebe: lc) (a cle ten cens 9 Oe Homing in Urosalpinx cinerea in Response to Prey Effluent and Tidal Periodicity. (: Text figure) DAVID View ATsre hee Mn mrad af AE SweoN 2 co 6) ela te evecn Gell) vie ce) bey en wt 30 Notes on the Opisthobranch Fauna of South San Francisco Bay. (2 Text figures) DARD EVV A DETIRE NS eONIERRE GD MGUEI Mi Enils lstye (cpg tc vs ole 6 4 eo)! 33 Notes on a California Hybrid Haliotis ore nae (1 Plate) RoBert R. TALMADGE . . . : 5 SS eR gle el we cio eh a tal Se The Effect of Temperature on the Distribution and Biomass of Mytilus edulis in the Alamitos Bay Area. (1 Map) GC VROBERT PELDMETE & )VWEREDVEH APPERT Mo cs 2 sete is es 5) 8. & e939 [Continued on Inside Front Cover] Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SUPERFAMILY, Famuy, Subfamily, Genus, (Subgenus) New Taxa Second Class Postage Pai at Berkeley, Californis ConTENTS — Continued Substrate Angle, Movement and Orientation of Two Sympatric Species of Limpets, Collisella digitalis and Collisella scabra. (4 Text figures) Linpa S. Co.tins . . SLAM Rese Aen c ates Prey Preferences of Carnivorous Intertidal Snails in the Florida Keys. Russe. E. INcHam & James A. ZISCHKE A New Species of Subcancilla (Gastropoda : Mitridae) from the Gulf of California. (1 Plate) R. A. WHITNEY : Bi ene ee : : : Feeding in Gastropod Mollusks: Behavioral and Neurophysiological Substrates - A Symposium. ‘ Predatory Behavior in Navanax inermis. M. J. Murray Mechanisms Underlying “Singleness of Action” in the Feeding Behavior of Pleurobranchaea californica (MacFarland, 1966) Jerrrey L. Ram & Wiuiam J. Davis Internal Variables Controlling Food Consumption in Aplysia. A. SUSSWEIN Neurobehavioral Studies on Feeding in the Terrestrial Slug, Ario- limax californicus. D. M. SENSEMAN Feeding in Helisoma trivoluis. C. R. S. Kanexo « S. B. Karer Behavioral Studies of Gastropod Feeding. ALAN GELPERIN Feeding in Navanax inermis, Neurophysiological Aspects. D. C. Spray, M. E. Sprra & M. V. L. BENNETT Neuronal Basis of Pleurobranchaea Feeding. ME topy V. S. SIEGLER Cholinergic and Serotonergic Control of Buccal Muscle in A plysia.. Kraupiusz R. Weiss, JosHuA CoHEN & IRvING KUPFERMANN Some Emergent Problems in Neural Control of Molluscan Feed- ing, A Chairman’s Summary. M. V. L. BENNETT NOTES & NEWS wis Weamatey lee MES aaa yeqese cern er hea tal aii dig MU ae ents New Range Extensions for Chitons (Amphineura : Polyplacophora). Grorce A. HANSELMAN Glans carpenteri vs. Glans subquadrata: The Rules Concerning Renamed Transient Secondary Homonyms. EucEnE V. Coan BOOKS, PERIODICALS & PAMPHLETS Distributed free to Members of the California Malacozoological Society, Inc. > 43 49 . 52 > 54 . 62 - 69 Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 20: $25.- plus postage ($1.- in U.S.A.; $2.50 to RU.A.S.; $3.50 to all other foreign Countries, including Canada). Single copies this issue $12.-. Postage additional. Send subscription orders to Mrs. J. DEMouTHE SmirH, % Department of Geology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 Address all other correspondence to Dr. R. StoHuER, Editor, Department of Zoology University of California, Berkeley, CA 94720 Vol. 20; No. 1 THE VELIGER Page 1 Integration of Electron Scan and Light Imagery in Study of Molluscan Radulae CAROLE S. HICKMAN Department of Biology, Swarthmore College, Swarthmore, Pennsylvania 19081 and The Academy of Natural Sciences of Philadelphia (1 Plate; 4 Text figures) INTRODUCTION OuR PERCEPTION OF THE NATURAL WORLD is strongly molded both by the nature of our sensory apparatus and the characteristics of the investigative tools we employ, to an extent that is seldom explicitly discussed in science. Too often we assume that intellectual objectivity is the only cornerstone of good science. In particular, we place a great deal of faith in our instrumentation. Optical microscopy (OM), typically utilizing absorp- tion contrast imagery, has long been the standard method of studying the molluscan radula, and most illustrations in the literature consist of camera lucida drawings, com- posite drawings, or reconstructions based on optical ex- amination. Advances in photomicrography make it pos- sible to produce relatively good photographs containing information not conveyed in artistic renditions, although few workers have used this tool in publication. The ad- vantages of scanning electron microscopy (SEM) have been amply demonstrated in the literature, and SoLEM (1972) has predicted that these advantages “will com- bine to make optical examination of radular structure obsolete.” Having experimented with all of the above methods of observation and communication, it is clear to me that, while SEM is the most powerful tool in radular study, (1) different tools of observation (2. e., different imaging processes) show us substantially different things, and (2) no single graphic method of communication can transmit all that our instruments show us. This paper demonstrates the manner in which data from light and electron scan images can be integrated to provide maximum information about radular form and function. First, I review the uses, advantages, and disad- vantages of each. method. Second, I present specific ex- amples of alternative views of radulae from the phasianel- lid Tricolia pullus (Linnaeus, 1758) and the trochid Solariella nuda Dall, 1896. Effective basic techniques for mounting radulae for OM and SEM examination are readily available in the litera- ture (see especially MEEUSE, 1950; SOLEM, 1972) and are not reviewed herein. Black-and-white photomicro- graphs in this paper were taken with an American Op- tical Series 10 Microstar trinocular microscope with Spencer Model 600 Ortho-Iiluminator, AO Photostar Exposure Meter and 35mm camera back. Scanning elec- tron micrographs were taken with the Cambridge Mark II-A and S4-10 instruments at the U.S. National Muse- um of Natural History with a Tectronix oscilloscope camera Model C-27. The following discussion and illustrations are based on morphological and evolutionary studies of radulae of rhipidoglossate marine archaeogastropods (HicKkMAN, 1976, and in preparation). Optical microscopy and scan- ning electron microscopy are combined to document con- vergent radular configurations that include: (1) com- plex within- and between-row basal overlap and inter- lock systems; (2) within-row shaft interlock and cusp interlock systems; (3) a variety of modifications to si- multaneously accomplish close packing of teeth when the radula is collapsed and maximum reach when it is in use; (4) disproportional development of the outermost lateral tooth cusp, in extreme cases with compensatory development of radular asymmetry; (5) systems of pivotal or articulatory plates in the hinge regions between the portions of the radula functioning along different major axes; (6) modifications and specializations of marginal teeth suggestive of as yet undocumented selec- tive deposit-feeding strategies. Page 2 Some of the above patterns are similar to those docu- mented in other groups of mollusks. The original studies of adaptive convergence were conducted by SoLEM (1971, 1972, 1973, 1974) on the radulae of land snails. His re- search elegantly established the utility of SEM in radular studies. Similar surveys are now available for cephalo- pods (SoLEM & RicHARDSON, 1975; SOLEM & RoPER, 1975) and nudibranchs (Bertscu e¢ al., 1973; BERTSCH & FERREIRA, 1974; FERREIRA & BERTSCH, 1975). Prelimi- nary work on the rhipidoglossate radula has revealed a much more complex set of patterns. This is, in part, because the rhipidoglossate radula is mechanically and morphologically the most complex type of molluscan radula. The typical rhipidoglossate radula is unique in possess- ing a large number of long slender marginal teeth, arrayed symmetrically on either side of the flat central portion of the ribbon. The marginal “books” of teeth are erected by outfolding of the ribbon and subsequently collapsed over the central part of the ribbon during the feeding stroke, as the ribbon passes over the tip of the odonto- phorial cartilage (ANKEL, 1938; EIGENBRODT, 1941; FRETTER, 1965; FRETTER & GRAHAM, 1962; NISBET, 1973; GraHAM, 1973). The central part of the ribbon is special- ized primarily for various kinds of food preparation and acts along an axis parallel to the length of the ribbon, while the marginals engage primarily in food gathering and act through a broad arc perpendicular to the length of the ribbon. For both OM and SEM examination it is necessary to perform delicate manipulations on the radula (e. g., fold- ing out or removing parts of the marginal books, separat- ing longitudinal and transverse rows) in order to minim- ize overlap of teeth and observe individual tooth morpho- logies and interrelationships. Because SEM viewing re- quires dry specimens coated with a conductive metal while optical mounts are made in a fluid medium, dif ferent configurations inevitably result from the two basic kinds of preparation. The most important single complementary difference between SEM and OM viewing of the rhipidoglossate tadula is related to the transparent but differentially mineralized and tanned nature of the chitinous teeth and ribbon to which they are attached. Because SEM viewing is restricted to surface features, it is possible to eliminate the confusion of multiple overlapping that occurs in stan- dard OM preparations. On the other hand, OM permits study of completely obscured teeth and parts of teeth that would never appear in SEM preparations as well as enabling observation of internal structure patterns related to readily visible differential mineralization and differen- tial tanning revealed through staining. THE VELIGER Vol. 20; No. 1 LIGHT MICROSCOPY The decrease in depth of field and resolving power that accompany increased magnification in optical systems are among the most serious problems in the traditional method of studying and illustrating radulae, although these are spectacularly minimized in the finest modern laboratory microscopes. Ease of varying illumination pa- rameters, and, in particular, the use of color filters to bring out detail and contrast (Figure 8b), enhance the data gathering potential of such instruments and play a key role in black-and-white photomicrography. Problems of photomicrography related to adaptation of camera optics (SoLEM, 1970) are eliminated in trinocular systems utilizing built in photographic objectives and rapid means of establishing parafocality between visual image and film plane. An additional, often cited, disadvantage of light micro- scopy lies in the fact that the radula must be mounted (“squashed,” in the words of the most severe critics) between 2 pieces of glass and viewed from a single posi- tion. Limited angle of observation may be compensated by dissecting portions of the radula so that individual teeth are mounted in a variety of positions as well as by use of depression slides or coverslip supports so that the radula retains three-dimensionality within the medium. As suggested in the introduction, the optical properties of radulae present both advantages and disadvantages. In the complex rhipidoglossate radula in particular, there is a great deal of overlapping of elements, making it time consuming and difficult to sort out the morphology and interrelationship of individual teeth. In careful opti- cal preparations, however, it is possible to determine re- lationships by focusing at different levels within groups of teeth and individual teeth in a way that is not possible in SEM viewing, which is restricted to surface features. OM is of great importance in detecting structures that can be subsequently explored more effectively with SEM, but which do not appear in preliminary SEM prepara- tions. One of the most important advantages of OM study of the rhipidoglossate radula is the potential amount of preparation time available for mounting in a fluid medi- um. Using arelatively low-viscosity water-soluble medium (e. g., furtox CMCP-10), one can spend an hour or more dissecting and arranging the radula before adding the cover slip. Even some of the more viscous media allow more preparation time than is available while drying a radula for SEM viewing. The more viscous the medium, the more likely that an arrangement will be preserved when the cover slip is added, although interesting and fortuitous configurations and structures may be revealed Vol. 20; No. 1 by turning and rotating of elements that may result. Distortions resulting from shrinkage of the radular mem- brane during drying for SEM viewing may likewise be either detrimental or fortuitous, the important point being that the two methods of preparation inevitably provide different configurations offering different kinds of information. Differences in the flexibility and hardness of different parts of a radula or of individual teeth must evolve in close relationship to the mechanical constraints on radular functioning and the feeding habits and substrates of in- dividual species. Thus OM observations of mineralization and staining patterns are of importance in radular studies. Documentation of mineralization in radular teeth is pri- marily restricted to the more spectacular instances of polyplacophorans and docoglossate archaeogastropods in which salts of iron and silicon increase tooth hardness (Jones et al., 1935; CarEFooT, 1965; TowE & LoweEn- sTAM, 1967; RunHam ef al., 1969), although various minerals may be added during the transformation of the soft first-formed radula into the toughened and hardened active portion of the radula (GaBE & PRENANT, 1957, 1958). Differential tanning, or formation of protein cross- linkages within the primary chitin structure, is indicated by differential staining patterns. For example, tooth bases are often heavily stained, while the slender flexible shafts of long marginal teeth often do not stain at all. There are three major graphic methods of presenting OM observations of radulae: the camera lucida drawing, the composite drawing or reconstruction, and the photo- micrograph. Camera lacida drawings provide faithful outline reproductions of tooth proportions and are a simple rapid means of illustrating a single halfrow of teeth for comparative taxonomic purposes. Such drawings do not incorporate variations within an individual or population, and, as line drawings of complex objects, they often contain a great deal of subjectivity or ambigui- ty, or both. They are most useful for showing cusps and seldom show details of tooth bases. Camera lucida draw- ings that attempt to show three-dimensional structure and overlapping are often difficult to interpret. Composite line drawings and reconstructions also con- tain a strong subjective element, but if camera lucida is used to establish proportions, such drawings may be extremely useful as summaries of observations at a varie- ty of magnifications, of different portions of a single rad- ula, or of different radulae. Such reconstructions can in- corporate data from electron scan images as well as light images. Photomicrography is most useful for producing a “true” image of the radula. Light micrographs are not particu- larly informative, however, if there is too much loss of THE VELIGER Page 3 resolution and depth of field in the image. Few authors have published light micrographs of radulae, but see McLean, 1971, for a series of informative light micro- graphs of turrid radulae. Micrographs at lower magni- fications are particularly efficient for documenting basic taxonomic characters (number of longitudinal rows of teeth, numbers and kinds of transverse elements, gross morphology, etc.). Micrographs are also an important means of documenting more detailed information and discoveries, and it should not matter that part of an image is blurred if the structure of specific interest is clearly re- solved. Indications of differential strength and hardness in teeth resulting from tanning and mineralization are best recorded through light microscopy. Alternative optical methods to standard absorption contrast may also be useful in radular study. Maes (oral communication, 1977) has experimented with low angle incident illumination to produce some remarkably three- dimensional photomicrographs of glycerine jelly mounts of rachiglossate radulae (see Orr, 1956; Maes, 1967). Phase contrast microscopy is extremely useful for pro- ducing in-focus contrast in unstained preparations (as opposed to the phase contrast that can be achieved in ordinary light microscopy by defocusing). Mitts (1977) has published some excellent phase contrast photomicro- graphs of toxoglossate radulae. Nomarski optics may also prove useful for examining some aspects of surface con- tour and should be explored as Nomarski systems become more widely available in this country. SCANNING ELECTRON MICROSCOPY Scanning electron microscopy has largely replaced light microscopy as the preferred tool for radula study during the past decade. Its most important advantages lie in broadening the potential scope of investigations, particu- larly at magnifications beyond the resolving power of light microscope optics. In functional morphological studies, SEM has become an indispensible tool, not only in its ability to explore microtopography from various angles, but also in the inherent ability of the incident electron probe to penetrate and explore narrow fissures and deep cavities that cannot be exposed by light. Research on functional morphology of the molluscan radula has focused.on the obvious food-preparing and food-gathering operations of the teeth and their relation- ship to design. These aspects are ideally examined from different angles with SEM and require some knowledge of the radula in functioning position. Less attention has been paid to the more routinely mechanical economics of Page 4 THE VELIGER Vol. 20; No. 1 occupying space. SoLeM (1971, 1974) and SoLEM & Roper (1975) have recognized the need to fold teeth to- gether efficiently to prevent interference with food passage and have identified several adaptations in land snails and cephalopods (grooving, curved fit surfaces) that function in tooth compaction. In the complex rhipidoglossate radula, occupation of space might be more profitably ex- plored in terms of designs that simultaneously satisfy maximum coverage of space by erected marginal teeth (as a function of tooth lengths and degrees of arc cover- ed) during sweeping activities and minimum occupation of space when collapsed. Interlocking and complex lock- and-key fits between tooth bases, shafts and cusps, viewed heretofore only as functioning in stress support, also ap- pear to play an important role in the economics of space occupation, particularly in the complex rhipidoglossate radula of marine archaeogastropods. These relationships are most effectively investigated by contrasting SEM data from preparations with marginal teeth erected with OM data from preparations with teeth in overlapping and tightly folded position. Although the great depth of field and high resolution of the electron scan image are impressive, aesthetically satisfying, and “state-of-the-art” in appearance, SEM images do not automatically contribute anything useful to comparative taxonomic research and can be detriment- al and confusing if they have been produced from differ- ent viewing angles and cannot be compared. Mounting between 2 pieces of glass for optical microscopy produces one of the most standard methods for comparing radulae of closely related congeners or for studies of variation patterns within populations, species, or along the length of a single radula. If SEM is used in illustrating strictly taxonomic studies, comparability of images should be maintained by standardizing the viewing angle and mounting the radula flat. Electron micrographs serve as the primary method of recording data during SEM viewing and are the usual means of illustrating published studies. Composite draw- ings based on a series of SEM images could be used to greater effect. Stereoscopic paired micrographs are in- valuable in interpreting SEM imagery. Low magnifica- tion stereoscopic pairs of entire preparations are partic- ularly helpful for planning efficient viewing strategies. THE RADULA OF Tricolia pullus The major features of the radula of the phasianellid Tricolia pullus are illustrated in Figures 7 to 4, 11, and 12. I have chosen this radula for illustration for two rea- sons: (1) it has been known for nearly a century and has been figured by previous authors (TROSCHEL, 1878: pit. 18, fig. 10; Prnssry, 1888: plt. 61, fig. 2; RoBERTSON, 1958: plt. 138, fig. 3) and (2) it contains several features that are difficult to interpret but potentially important in understanding basic functioning patterns of the rhipidoglossate radula. Drawings of the Tricolia pullus radula based on light microscope examination (Figure 10) establish the basic pattern of dentition and overlap of an oval rachidian flanked on either side by 5 strongly cusped, progressively narrower laterals with expanded overlapping bases and series of numerous marginals. The inner marginals have Explanation of Figures 1 to 9 Tricolia pullus (Linnaeus, 1758). USNM 179033 Figure 1: Central portion of whole mount. OM X 100 Figure 2: Central and inner lateral teeth, illustrating transverse basal overlap system. SEM X 570 Figure 3: Detail of inner marginal cusp and rectangular “latero- marginal plate” with adjacent marginal teeth removed. OM X 300 Figure 4: Isolated inner marginal tooth, demonstrating that “lat- eromarginal plate” is the tooth base. Note that cusp was broken during manipulation. OM X 300 Solariella nuda Dall, 1896. USNM 209300 Figure 5: Detail of first lateral tooth from which cusp is broken, revealing basal interlock with central tooth (lower right) and basal interlock with second lateral tooth (upper left). SEM X 760 Figure 6: Detail of lateromarginal plates where marginal teeth have been removed. OM X 160 Figure 7: Detail of lateromarginal plates with first marginal teeth folded out, other marginals removed. OM X 160 Figure 8: Central portion of folded whole mount with a. normal illumination and b. red filter to enhance contrast. OM X 115 Figure 9: Low angle side view showing a. erected marginal teeth; b. lateromarginal plate with thin membranous rudimentary cusp; c. cuspless Jateromarginal plate; d. grooves and ridges on heavy outer lateral tooth; e. depression on rachidian accommodating cusp of following rachidian. SEM X 225 [Hickman] Figures 7 to 9 Tue VeuicER, Vol. 20, No. 1 % Ry ee “ype, Wich phe oat ie = & Vol. 20; No. 1 Figure 10 Line drawing of a part of a half-row of theradulaof Tricolia pullus, based on light microscopy, after RoBERTSON (1955). Figure 11 Reconstruction of part of a half-row of the radula of Tricolia pullus, from light and scanning electron microscope imagery. Inset of lateral teeth shows semi-profile view of cusps. a strong primary cusp and a short secondary cusp along the outer margin. The outer marginals have a single, nar- row, serrate cusp. For comparative taxonomic purposes, drawings of this kind provide excellent documentation from a standard vantage point of characteristic configura- tions or patterns (ROBERTSON, 1958: plt. 138, figs. 3 - 6). Although cusp details are depicted, the drawings are basically two-dimensional outlines that do not attempt interpretation of the thickness of teeth or parts of teeth, the manner in which teeth are attached to the radular ribbon, or the significance of overlapping elements. A photomicrograph of the radula of Tricolia pullus (Figure 1) illustrates the kind of image upon which the line drawing of Figure 10 was based. The photomicro- graph contains a great deal more complexity and addi- tional kinds of information, the most obvious of which are THE VELIGER Page 5 the patterns of multiple overlap of transverse rows and the variation in optical properties of different radular elements. Micrographs such as this and the comparable one for Solariella nuda (Figure 8) provide a rapid means of documenting sources of variation within longitudinal rows, including (1) random variation in actual morpho- logy, as in the number of cusps, resulting from develop- mental differences, (2 )random variation in apparent morphology resulting from viewing a tooth from a slightly different angle, and (3) systematic variation along the length of the ribbon resulting from tooth wear. It is also evident that parts of the absorption contrast image are subject to more than one interpretation. A particularly interesting case of interpretational ambiguity arises from a prominent series of dark-staining rectangu- lar structures between the outer lateral and inner mar- ginal teeth in a whole mount of the Tricolia radula. RoBERTSON (1958) interpreted the structure as a cusp- less tooth and proposed the name “‘lateromarginal plate” for it. Marcus & Marcus (1960) rejected Robertson’s interpretation and suggested that the plate is the base of the innermost marginal tooth. Dissection of the radula to isolate individual teeth confirms the latter view, and photomicrographs of the first lateral tooth with its en- larged rectangular base appear in Figures 3 and 4. Examination of large numbers of rhipidoglossate radu- lae, however, does reveal the presence of a cuspless struc- ture between the lateral and marginal teeth in many archaeogastropods (Hickman, unpubl.), and the name lateromarginal plate is perfectly applicable. It is also interesting to note that there are a number of overlooked illustrations of and independent allusions to these cusp- less structures in earlier literature (e.g., SARs, 1878, Margarites; Puspry, 1889, Trochus, Clanculus; Tro- SCHEL & THIELE, 1891, Moelleria, Puncturella; ScHEP- MAN, 1908, Solariella; Torr, 1914, Emarginula, Lucap- inella, Megatebennus). In many cases the plate is clearly a pivotal or articula- tory structure, either supporting or holding the bases of the marginal books of teeth as they are erected and col- lapsed during feeding. In several trochid genera (e. g., Margarites, Calliotropis, Bathybembix) rudimentary cusps are associated with the plate, suggesting its inde- pendent derivation through progressive enlargement of the base and reduction and loss of the cusp (HicKMAN, in preparation). SEM examination adds significantly to our understand- ing of the Tricolia radula, particularly in interpretation of overlapping structures. Figure 2 reveals a complex and relatively heavy basal stress support system whereby the stress applied to any one tooth may be spread outward to other teeth within the same transverse row. In addition to the basal overlap features, Figure 2 also reveals inter- Page 6 lock features higher on the teeth that are more likely to function in compaction of the radula when it is not in use. Note the potential fit of the thickened convex ridge on the shaft of the third lateral into the corresponding pocket beneath the cusp of the second lateral in Figure 2. A reconstruction of the Tricolia radula (Figure 11), based on a series of 7 SEM images from a variety of angles and magnifications, including 2 stereo pairs, also shows a complex system of interlocking of the inner mar- ginal cusps in which the secondary outer cusp fits into a groove on the flattened top of the neighboring primary cusp. THE RADULA OF Solariella nuda The radula of Solariella nuda has not been figured pre- viously. Radulae have been illustrated by line drawings for at least 15 species of Solariella, however (Sars, 1878; TroscHEL, 1878; MarTENS & THIELE, 1903; SCHEPMAN, 1908; ODHNER, 1912; POWELL, 1951; GALKIN, 1955; McLean, 1964), and some of the major features of the solarielline radula, as heretofore understood, are illus- trated in a line drawing of Solariella delicata Dall (Fig- ure 12). Recurrent characteristic features of the line drawings in the literature and features cited in accompanying texts, all based on optical examination, include (1) the short- ness of the radula, which is sometimes less than twice as long as it is wide; (2) the small number of marginal teeth \/ ~ (approximately 10 per half-row); (3) the prominent dip at the center of each transverse row; (4) the prominent, coarsely serrated cusp of the rachidian; and (5) the ap- parent restriction of serrations to the outer margins of the first and second lateral teeth. Other features are ambiguous or variably interpreted from one drawing to another, particularly with respect to basal overlapping and the shapes of the third, fourth, and fifth lateral teeth, if these are recognized as being present. The reason for the ambiguity lies in the fact that the marginal teeth in the folded solarielline radula multiply overlap and obscure all but the rachidian and inner 2 lateral teeth in an unmanipulated preparation. This is illustrated by a photomicrograph of the folded radula of Solariella nuda (Figure 8). When the marginal books of teeth are folded out or carefully removed, a broad-based third lateral tooth with a serrate, long, medially-directed cusp is revealed, as well as a narrow but heavily based and extraordinarily long, cusped fourth lateral tooth. In addition there is a longitudinal series of rectangular lateromarginal plates THE VELIGER Vol. 20; No. 1 Figure 1 2 Line drawing of a half-row of teeth from the radula of Solarella delicata, redrawn from GaLKIN (1955) and retaining ambiguity of line from his drawing. Figure 13 Reconstruction of the basic solarielline dentition pattern as exem- plified by Solariella nuda, from light and scanning electron micro- scope imagery. A diagrammatic representation of the basal interlock system appears beneath the main part of the reconstruction. (Figures 6, 7) against which the marginal books articu- late. I have found similar series of lateromarginal plates in 7 other solarielline species, although ScHEPMAN (1908: plt. 9, figs. 8, g) is the only author to have recorded them. A SEM preparation in which the marginal teeth have been removed for viewing from the side (Figure 9) pre- sents an alternative view of the lateromarginal plates of Solariella nuda. Note that, although the posterior 2 plates have a small hooked projection on their inner margins, Vol. 20; No. 1 they are uncusped; while the next anterior plate has a thin “cusp” that fits compactly beneath the folded, large, heavy blade of the fourth lateral tooth of the next-most anterior row. By making an additional OM preparation of the radula of Solariella nuda, it became apparent that the “cusp,” when present on the lateromarginal plate, is a flexible membranous structure that does not resist stress and can be bent easily through an arc of 180° without any move- ment of the fixed lateromarginal plate. Although the vestigial cusp looks “solid” in the SEM image, the wrink- ling and filamentous projections from the trailing edge suggest its extreme thinness in contrast to the heavy outer lateral. From both SEM and OM as well as physical manipulation of the radula with minuten insect pins, it appears that the marginal books are supported, probably through fine muscular adjustments (NIsBET, 1973), in erect position against the thick outer edge of the lateromarginal plate as they nearly are at the arrow in Figure 9. An interesting feature revealed by SEM that does not appear in OM preparations is the grooving in the upper surfaces of the fourth lateral teeth. In folded position these grooves accommodate closer, more economical pack- ing of the marginal books. Each of the broad thin mar- ginal tooth blades is convex on its inner surface and con- cave on its outer surface. This design simultaneously facil- itates close packing of the collapsed marginal books of teeth into the folded lateral tooth complex and confers greater strength to each of the thin marginal blades than it would have if it were perfectly flat. Also note how the depression at the top of each rachidian accommodates the top of the cusp of the overlapping rachidian and base immediately posterior. As in the Tricolia radula, there is a complex basal interaction system in the solarielline radula. Unlike the simple Tricolia overlap system, however, the solarielline system involves complex interlocking. It cannot be studied effectively using SEM because the broad cusps complete- ly obscure the tooth bases from nearly all angles of viewing when the teeth are in interlocked position. For- tuitous breakage of an inner lateral cusp confirms the nature of the interlocks between the first lateral and the rachidian on one side and the second lateral on the other (Figure 5). A reconstruction of the radula of Solariella nuda (Fig- ure 13), based on a combination of light images and SEM images, including stereoscopic pairs, is offered to illustrate some of the major features of the solarielline radula. THE VELIGER Page 7 SUMMARY Although SEM provides many new possibilities for un- derstanding morphology and function of the molluscan radula, it does not replace OM as a source of data. Dif- ferences in specimen preparation as well as differences in imaging processes lead to different kinds of data, so that it is particularly instructive to use both in radula studies. The resulting information, likewise, can be graphically transmitted most effectively by combining photomicro- graphs, scanning electron micrographs, camera lucida outline drawings, and composite diagrams and reconstruc- tions. Applied to the complex rhipidoglossate radula of ma- rine archaeogastropods, exemplified herein by Tricolia pullus and Solariella nuda, these methods are elucidating a broader and more complex range of structural patterns than has been documented in other mollusk groups. ACKNOWLEDGMENTS This work was initiated as a Visiting Investigator in the Department of Paleobiology at the National Museum of Natural History, Smithsonian Institution. I am grateful to Thomas R. Waller of the National Museum for arrang- ing for the use of the scanning electron microscope and to Susann Braden for her helpful technical assistance. Dis- cussions with Thomas R. Waller, Alan Solem (Field Mu- seum), and Arthur Cohen (Washington State University) have contributed to development of effective SEM tech- niques. James H. McLean (Los Angeles County Muse- um of Natural History) has provided helpful suggestions for preparation of permanent mounts for OM and kindly read an earlier draft of the manuscript. Virginia Orr Maes, Academy of Natural Sciences of Philadelphia, also read the manuscript and suggested several specific tech- niques for use in optical microscopy from her own radula studies. Literature Cited ANKEL, WuLF Emmo 1938. Erwerb und Aufnahme der Nahrung bei den Gastropoden. Verh. Deutsch. Zool. Gesell. 11: 223 - 295 Bertscu, Hans & ANTONIO J. FERREIRA 1974. Four new species of nudibranchs from tropical west America. The Veliger 16 (4): 343 - 353; 7 plts. (1 April 1974) Page 8 THE VELIGER Vol. 20; No. 1 BertscH, Hans, ANTonio J. Ferrera, Westey M. FarMER & THOMAS L. Hayes 1973. The genera Chromodoris and Felimida (Nudibranchia : Chromo- dorididae) in tropical west America: distributional data, description of a new species, and scanning electron microscopic studies of radulae. The Veliger 15 (4): 287 - 294; 3 plts.; 3 text figs. (1 April 1973) Careroot, Tuomas H. 1965. Magnetite in the radula of the Polyplacophora. col. Soc. London 36: 203 - 212 E1cEnBropT, H. 1941. Untersuchungen tiber die Funktion der Radula einiger Schnek- ken. Zeitschr. fiir Morphol. und Okol. Tiere 37: 735 - 791 Ferrera, ANToNIo J. « Hans BertscH 1975. Anatomical and distributional observations of some opistho- branchs from the Panamic faunal province. The Veliger 17 (4): 323 - 330; 3 plts.; 1 text fig. (1 April 1975) FRETTER, VERA 1965. Functional studies of the anatomy of some neritid prosobranchs, Journ. Zool. London 147: 46-74; 11 text figs. Fretrer, VERA & ALasTaR GRAHAM 1962. British prosobranch molluscs, their functional anatomy and eco- logy. London, Ray Soc. xvit 755 pp.; 317 text figs. Gazz, M. « M. PRENANT 1957. Particularités histochimiques du ruban radulaire et des dents de la radula chez quelques mollusques. Bull. Soc. Zool. France 82: 195 - 196 1958. Particularités histochimiques de l'appareil radulaire chez quel- ques mollusques. Ann. Histochem. 3: 95 - 112 Gauxin, I. U. 1955. Monograph of the mollusks of the family Trochidae of the far eastern seas of the U.S.S.R. Opredeliteli po faune SSSR 57; 131 pp. (in Russian) GraHamM, ALASTAIR 1973. | The anatomical basis of function in the buccal mass of proso- branch and amphineuran molluscs. Journ. Zool. London 169: 317 - 348; 6 figs Hickman, Caro.e STENTZ 1976. Form, function, and evolution in the archaeogastropod radula. Geol. Soc. Amer. Abstr. with Programs 8 (6): 917-918 (Abstract) Jones, E. I., R. A. McCancg « L. R. B. SHAcKLETON 1935. The role of iron in the structure of the radular teeth of certain molluscs, Journ. Exper. Biol. 12: 59 McLzan, Jamzs Hammtton 1964. New species of Recent and fossil West American aspidobranch gastropods. The Veliger 7 (2): 129-133; pit. 24; 1 text fig. (1 October 1964) 1971. A revised classification of the family Turridae, with the proposal of new subfamilies, genera, and subgenera from the Eastern Pacific. The Veliger 14(1): 114-130; 4 plis. (1 July 1971) Mags, Vircinia Orr 1967. ‘Radulae of two species of Pleuroploca (Fasciolariidae) from the Indo-Pacific. The Nautilus 81 (2): 48-54; 1 pit. Marcus, Ernst & EVELINE Du Bois-REYMOND Marcus 1960. On Tricolia affinis cruenta. Bol. Fac. Fil., Cien. Letr. Univ. SZo Paulo 260: 171-211; 6 text figs. Martens, Epuarp [Cart] von & JOHANNES THIELE 1903. Die beschalten Gastropoden der Deutschen Tiefsee-Expedition 1898-1899. B. - Anatomisch-systematische Untersuchungen einiger Gastropoden. Deutsch. Tiefsee-Exped. 7: 149+ 182 Meeusg, A. D. J. 1950. Rapid methods for obtaining permanent mounts of radulae. Basteria 14 (1&2): 28-43; plts. 4-6; 3 text figs. (1 May 1950) Proc. Mala- Mitts, Patricia M. 1977. | Radular tooth structure in three species of Terebridae (Mollus- ca : Toxoglossa). The Veliger 19 (3): 259-265; 3 plts.; 1 text fig. (1 January 1977) Niszet, R. H. 1973, | The role of the buccal mass in the trochid. Soc. London 40 (6): 435-468; 21 text figs. Opuner, Nits Hyatmar 1912. Northern and arctic invertebrates in the collection of the Swedish State Museum. V. Prosobranchia, 1. Diotocardia. Kgl. Svens. Vetensk.-Akad. Handl. 48 (1): 1-93; plts. 1-7 Orr, VIRGINIA 1956. The south African gastropod genus Burnupena (Buccinidae). Proc. Acad. Nat. Sci. Philadelphia 108: 249 - 263; plts. 19 - 20 Pirssry, Henry Aucustus 1888. Manual of Conchology. Philadelphia 1 (10); 323 pp.; 69 plts. 1889. The radula in rhipidoglossate mollusks. Proc. Acad. Nat. Sci. Philadelphia 44: 36 - 37 Powerit, ArTHUR WiLL1AM BADEN 1951. Antarcticand subantarctic Mollusca: Pelecypodaand Gastropoda. Discovery Reprt. 26: 47 - 196; plts. 5- 10 Rosertson, RoperT 1958. The family Phasianellidae in the western Atlantic. 3 (37): 245 - 283; plts. 136 - 148 Runuam, Norman W, PR. Tuornton, D. A. SHAW # R. C. WayTE 1969. The mineralization and hardness of the radular teeth of the nae Patella vulgata L. Zeitschr. Zellforsch. 99: 608 - 626; 9 text Proc. Malacol. Johnsonia Sars, Gzorcz Ossian 1878. Bidrag til Kundskaben om Norges Arktiske fauna: I. Mollusca Regionis Arcticae Norvegiae pp. i-xii+ 1-466; 34 + xviii pits. Christiania ScHEpMAN, MattHEus Marinus 1908. Prosobranchia of the Siboga Expedition, Pt. I. Rhipidoglossa and Docoglossa. Siboga Exped. 28 (49a): 1-107; plts. 1-9 Sotem, ALAN 1970. Malacological applications of scanning electron microscopy I. Introduction and shell surface features. The Veliger 12 (4): 394 to 400; plts. 58-60 (1 April 1970) 1971, Structure and function of teeth in unrelated camivorous snails. Year Book of the American Philosophical Society, pp. 347 - 349. 1972. | Malacological applications of scanning electron microscopy II. Radular structure and functioning. The Veliger 14 (4): 327 - 336; 6 plts.; 1 text fig. (1 April 1972) 1973. | Convergence in pulmonate radulae. The Veliger 15 (8): 165 - 171; 4 pits. (1 January 1973) 1974. Patterns of radular tooth structure in carnivorous land snails. The Veliger 17 (2): 81-88; 7 plts. (1 October 1974) Sorem, ALAN & Eucene S. RicHARDSON 1975. | Paleocadmus, a nautiloid cephalopod radula from the Pennsyl- vanian Francis Creek Shale of Illinois. |The Veliger 17 (3): 233 - 242; 5 plts.; 1 text fig. (1 January 1975) Sotem, Avan a Crype FE. Roper 1975. Structures of Recent cephalopod radulae. 18 (2): 127 - 133; 4 plts. Torr, Craupe M. 1914. | Radulae of some South Autralian Gasteropoda. Soc. So. Austral. 38: 362 - 368; plts. 19 - 20 Tower, KennetH M. « H. A. Lowenstam 1967. Ultrastructure and development of iron mineralization in the radular teeth of Cryptochiton stelleri. Journ. Ultrastruct. Res. 17: 1-13 TroscHEL, Franz HERMANN 1878. Das Gebiss der Schnecken. 2 (5): 118-216; plts. 17-20 TROSCHEL, Franz HERMANN & JOHANNES THIELE 1891. Das Gebiss der Schnecken 2 (7): 294 - 334; plts. 25-28 The Veliger (1 October 1975) Trans. Roy. Vol. 20; No. 1 THE VELIGER Page 9 A New Species Tentatively Referred to Antonietta, Antonietta janthina, from Japan (Nudibranchia : Eolidoidea : Facelinidae) BY KIKUTARO BABA Shigigaoka 35, Minami-11-jyo, Sango-cho, Ikoma-gun, Nara-ken, Japan AND IWAO HAMATANI Tennoji Senior High School of the Osaka Kyoiku University, Tennoji, Osaka, Japan (4 Text figures) THE FIRST SPECIMEN of the species which forms the subject of this paper was collected by Baba in March 1937, when he lived in the Amakusa Marine Biological Laboratory, Kyushu University. The animal, though seemingly distinctive in the vermilion rhinophores and purple branchiae, could not at that time be identified. In later years more specimens were collected from some other stations of Japan which enabled him to continue a taxonomical study of the species. Meanwhile, Hamatani who was the collector of one of these latter specimens came to join with Baba in order to conduct the study of internal organs of the species from serially sectioned mate- rial. A new taxon, janthina, was finally established by the two authors and it was tentatively referred to Antonietta Schmekel, 1966 from the Mediterranean Sea, mainly be- cause of the patterns of liver branchings. Antonietta janthina Baba « Hamatani, spec. nov. (Japanese name: Murasaki-mino-umiushi) (Figures 1 to 4) Distribution: Pacific coast of middle and southern Jap- an: Hayama, Sagami Bay; Seto, Kii; and Tomioka, Amakusa. Japan Sea coast of middle Japan: Ushitsu, Toyama Bay. Holotype: Collected by Hamatani from the shore of Yuzaki, vicinity of the Seto Marine Biological Laboratory, 15 August 1962. After taking a picture of the living ani- mal by Baba, it was fixed and prepared in serial hori- zontal sections. Additional data referred to are as follows: One speci- men collected and figured by the Biological Laboratory, Imperial Household, from shallow water of Samejima near Hayama, Sagami Bay, 19 March 1948. This was used especially for demonstrating jaws and radula. One specimen collected and figured by Baba from the outer shore of Magarizaki, vicinity of the Amakusa Marine Bio- logical Laboratory, 2 March 1937. The animal was found feeding on a light brown hydroid colony (cf. Hydractinia epiconcha) which covered the surface of a snail shell housing a hermit crab. Two specimens collected and fig- ured by Mr. Abe and other members of the Takaoka Biological Club, from the shore of Hime, near Ushitsu, Toyama Bay, 10 August 1971. DESCRIPTION External Form: The holotype measures 10mm in length. The general body form is fundamentally as usual in the family Facelinidae. That is, the body is slender, the oral tentacles are elongated, the rhinophores are shorter, Page 10 THE VELIGER a es ET Si b d Figure 1 Antoniesta janthina Baba & Hamatani, spec. nov. from Seto, Kii, Japan Living animal from dorsal side, length (Ac) 10mm a — orange yellow head b = vermilion rhinophore c — purple diverticulum d — pale yellow cap Vol. 20; No. 1 | ID — eR QYNS SSN & : Sy \ S Os HES iq?) aKS oe SS Fy 7) yy Zr SS Ra QQ SESS Cap. = Z Figure 2 LLY Antonietta janthina Baba x Hamatani, spec. nov. from Seto, Kii, Japan Digestive system; salivary glands not shown a — right liver d — anus b — genital orifices e — left posterior liver c — nephroproct f - left anterior liver Vol. 20; No. 1 and the foot forms tentaculiform corners anteriorly. The rhinophores themselves are smooth on the surface. The tail is short and tapering behind. The liver system is more similar in constitution to Antonietta (see SCHMEKEL, 1966), and Palisa (see Ep- MuNDs, 1964) than to Learchis (see Basa, 1969) which is synonymized by BurN & NaraYANAN (1970) with Caloria. The right liver (and the partner on the left side) is formed of 5 simple oblique branches, and the arrange- ment of branchial papillae on them is shown as 3, 5, 5, 6, and 8, successively. The left posterior liver is differ- entiated on each side into 6 horseshoes followed by 4 short oblique rows. The cleioproctic anus lies in the middle of V THE VELIGER Page 11 the 1" right horseshoe which contains 5 papillae in each of two legs. The papillae decrease in number in the suc- ceeding horseshoes. They are simply elongated fusiform. The nephroproct is interhepatic. The genital orifices are found below the middle of the right liver branches. Coloration: The median part of the head is tinged with orange yellow. The oral tentacles are also orange yellow though this color is more or less accentuated to- wards the tip. The rhinophores are prominently vermil- ion-tinted except at their midlength where this color tends to disappear. The vein (= liver diverticulum) of the papillae is purple throughout its length, and not marked Figure 3 Antonietta janthina Baba x Hamatani, spec. nov. from Hayama, Sagami Bay, Japan A: Right jaw with the jagged edge enlarged (X 60) B-D: Teeth in different views (X 520) Page 12 a SD, SEE prErRA Tei i) Rit, YAuebeo ieee b Figure 4 Antonietta janthina Baba « Hamatani, spec. nov. from Seto, Kii, Japan Genital system from dorsal side (X 40) b — outer oviduct c — spermatocyst d — inner oviduct (distal part not defined) e — hermaphrodite duct f — ampulla g — muscular part of vas deferens h — accessory female gland mass i — prostatic part of vas deferens a — penis distally with a melanin black spot as shown in Caloria (see HAEFELFINGER, 1960) and Learchis (see Basa, 1969). The cap of the papillae is tinted pale yellow. The ground color of the integument (back, sides and sole) is fleshy white. The pericardial prominence is opaque white. Also there are irregular mottlings of opaque white on the median line of the dorsum. The anterior margin of the foot is slightly orange yellow. The tail is colorless. THE VELIGER Vol. 20; No. 1 Internal Anatomy: ‘The mouth parts were studied on a specimen from Sagami Bay. As in Learchis and Antont- etta, the jaws have no dorsal indentation (in Calorza these are indented dorsally). The jaw edge is jagged now, but details of denticulation could not be defined exactly (in Antonietta the jaw edge is said to be smooth). The radu- lar formula is 35 X 0-1-0. The central tooth is typically cuspidate asin that of Antonietta,and bears 8-11 denticles on either side of the highly produced median cusp. As in Learchis and Antonietta, the penis of the holotype is conical, unarmed, and without an accessory penial gland. The distal portion of the vas deferens forms a prostate. In the holotype there occurs a single bursa (—spermatocyst) attached to the outer oviduct (in Learchis there is a single bursa, and in Antonietta it is shown that there are 2 bursae on the female duct; see also MiLtEr, 1974). DISCUSSION It seems still difficult to identify the genera (and species) of the family Facelinidae satisfactorily (see EDMUNDS, 1970 and Mirter, 1974). Here the new species janthina was referred to Antonietta Schmekel, 1966 (Type: A. lu- teorufa from Naples) merely for the horseshoe-shaped composition of the left posterior liver branches, for the non-indented jaws, and for the unarmed penis not accom- panied by an accessory gland. Actually there may be seen some minor differences existing between the genital systems of the 2 species, A. luteorufa and A. janthina. According to Burn «& NarayANAN (1970) the south- west Pacific species Learchis indica Bergh, 1896 (see Baza, 1969) is synonymous with the Indian form Eolis militaris Alder & Hancock, 1864 (see also Miter, 1974); furthermore, the genus Learchis Bergh, 1896 constitutes a junior synonym of Caloria Trinchese, 1888 known from the Mediterranean Sea (Type: C. maculata; see HAEFEL- FINGER, 1960). Then, Caloria (= Learchis) is disting- uished from Antonietta by the left posterior liver which is formed of clusters (not horseshoes) of branches. The species Learchis poica Marcus & Marcus, 1960 (from Miami) and Palisa papillata Edmunds, 1964 (from Jamaica and Miami) are similar to Antonietta in the type of the liver system, but they are distinguished from the latter, respectively, as follows: Learchis poica is provided with an accessory gland to the penis, and Palisa papillata has the rhinophores covered with small papillae on their posterior surface. A thorough re-examination of the interrelationship between the above-mentioned genera (and species) is, therefore, to be expected in the future. Vol. 20; No. 1 THE VELIGER Page 13 SUMMARY 1. Anew species, Antonietta janthina, is suggested from Japan. 2. Externally this new species is especially distinctive in the vermilion rhinophores and purple diverticula of the branchial papillae. 3. Anatomical accounts regarding the jaws, radula, liver system and genitalia of this new species are given, together with some taxonomical comments. 4. This new species is assigned to the Mediterranean genus mainly for the horseshoe-shaped composition of the left posterior liver branches. The penis is unarmed, and there is no accessory gland to this organ. ACKNOWLEDGMENTS One of the authors (Baba) wishes to express his appre- ciation to the Chief of the Biological Laboratory, Imperial Household, and to Mr. Takeo Abe of the Takaoka Bio- logical Club, for giving him the opportunity of identify- ing specimens collected by them. Literature Cited Basa, KikuTARO 1969. Records of Learchis indica Bergh, 1896 from Japan and Hawaii (Nudibranchia : Eolidoidea). Publ. Seto Mar. Biol. Lab. 16 (6): 399 - 403; plt. 27 Burn, Ropert « K. R. NARAYANAN 1970. Taxonomic notes on Eolis militaris Alder and Hancock, 1864 (Opisthobranchia, Eolidacea). Journ. Malac. Soc. Austral. 2 (1): 83 - 86 EpmuNps, Matcorm 1964. Eolid Mollusca from Jamaica, with descriptions of two new genera and three new species. Bull. Mar. Sci. Gulf & Caribb. 14 (1): 1-32; 16 text figs. (March 1964) 1970. Opisthobranchiate Mollusca from Tanzania. II. Eolidacea (Cu- thonidae, Piseinotecidae and Facelinidae). Proc. Malac. Soc. Lond. 39 (1): 15-57; figs. 1-24 HAEFELFINGER, Hans-Rupo.F 1960. Neue und wenig bekannte Opisthobranchier der Gattungen Tra- pania und Caloria aus der Bucht von Villefranche-sur-Mer (A.-M.). Rey. Suisse Zool. 67 (2): 226 - 238; 8 text figs. Marcus, Eve.Line pu Bots-REYMOND & ERNST Marcus 1960. Opisthobranchs from American Atlantic warm waters. Bull. Mar. Sci. Gulf & Caribb. 10 (2): 129-203; 97 text figs. Miiier, MicHaeEL CuHarLes 1974. Aeolid nudibranchs (Gastropoda: Opisthobranchia) of the fam- ily Glaucidae from New Zealand waters. Zool. Journ. Linn. Soc. 54 (1): 31-61; plt. 1; 10 text figs. ScHMEKEL, RENATE LUISE 1966. Zwei neue Facelinidae aus dem Golf von Neapel: Facelina (A.) fusca n. sp. und Antonietta luteorufa n. sp., n. gen. (Gastr. Opistho- branchia). Pubbl. Staz. Zool. Napoli 35 (1): 29-46; 6 text figs. Page 14 THE VELIGER Vol. 20; No. 1 A Sand-Dwelling Elysia from Guam (Opisthobranchia : Sacoglossa ) C. H. CARLSON ano P. J. HOFF The Marine Laboratory, University of Guam (5 Text figures) In Octoser, 1971, the authors were searching for opisthobranchs in a large bed of the green alga Caulerpa racemosa (Forskal) J. Agardh when 2 specimens of a new species of Elysia were found. Since that time, 27 additional specimens have been collected in the same area inside Cocos Lagoon, Merizo, Guam, Mariana Islands. Elysia arena Carlson « Hoff, spec. nov. Description: The animals are from 5.0 to 32.0mm long. A 23.0mm animal was 5.0mm wide at the widest point with parapodia erect. With parapodia completely opened, a 160mm preserved animal is 14.0mm wide; almost as wide as it is long and it is rectangular rather than tri- angular. The animal (Figure 1) is very heavy bodied, covered with conical projections on the head, rhinophores, and Figure 1 Elysia arena. Dorsal view of living animal a — anal opening e — eye m. g. p. — male genital pore * Contribution No. 99 from the University of Guam Marine Laboratory parapodia. The largest projections are about midway down the parapodia. The parapodia are high, irregular in height, undulating on edges, held fairly close but not tight and at times have several openings — frequently with an opening at the highest point. There is no division between the sole and the parapodia, but there is a slight ventral transverse furrow where the head joins the foot. The angles of the foot are prominent. The head is very high, almost square from a dorsal view, sometimes appearing bilobed anteriorly. The rhino- phores are set close together and are heavy, blunt, and auriculate. The eye spots (Figure 1) are clearly visible. The pericardial prominence (Figure 2) is fairly small considering the size of the animal, and extends posterior- ly in a large, long tube (Figure 2). There is a very thin tube (Figure 2) situated on top of the larger tube, and the renal pore (Figure 2) is easily visible on the right side of the pericardial prominence. There are numerous T] rei NAS ( \ NS \ : p.f Figure 2 View of Elysia arena with parapodia open P — pericardium p. £ — parapodial folds v.— venous network r — renal pore Vol. 20; No. 1 veins (Figure 2) extending outward from the reno-peri- cardial complex which have some branching. This venous network is similar to that of other Elysia species on the internal dorsal area, but on the internal lateral surface of the parapodia the veins terminate in extremely large, heavy parapodial folds (Figures 2 and 3) extending al- most to the edge of the parapodia. These large folds, reminiscent of lamellae, are the most distinctive aspect p.f S ; Figure 3 Horizontal section showing parapodial folds Sectioned animal is 3.2 mm wide with parapodia erect p:- a. — outer wall of parapodia p. f. - parapodial folds of the animal. On an animal that was 14.0mm wide when preserved, the longest folds were 4.5mm in length, thus equaling about one third of the animal’s width with parapodia open. The anus and genital pore (Figure 1) are lateral, opening in a single aperture at the furrow just anterior to the right parapodium. The penial opening (Figure 1) is under the right rhinophore. The mouth has a slight “T” shape. The pharynx (Fig- ure 4) is extremely large with the dorsal part covered with heavy annular muscles. The radula is anterior and ends straight in the ascus (Figure 4) with no loose pile of discarded teeth. The radula in a 14.0mm specimen had 17 teeth in the descending series and 7 in the ascending. That of a 23.8mm specimen had 7 teeth ascending and 23 in the descending series. In the latter specimen the tooth in use was 0.31mm in length, the newest tooth 0.35mm, and the oldest 0.02mm. The teeth (Figure 5) have very fine serrations on the cutting edge. Under a microscope the serrations are visible at 400 power but not at 100. In the living animal the head is light green, or some- times tan with a greenish tinge, and has widely scattered black dots and some creamish tan mottling. The projec- tions are white. Minute dots of chocolate brown or black form a triangular pattern immediately anterior to the THE VELIGER Page 15 aos 7~ as a 1.0mm | Figure 4 Pharynx of Elysia arena as — ascus m - annular muscles oe — oesophagus rhinophores. Very dark brown dots appear at the sides of the head below the eye spots and extend to the anterior edge of the parapodia. The rhinophores are tan with white projections and white spots, and have brown dots which are more predominant on outer edges toward the tip. The parapodia are tan, sometimes having an orangish tinge with scattered fine brown flecks and white projec- tions. The brown flecks are denser at the anterior of the parapodia and toward the lower central part of the para- podia; some faint white patches are over the tan. The intensity of the brown pigmentation may vary from ani- 0.3 mm =] 0.025 mm Figure 5 Teeth of Elysia arena Page 16 mal to animal with the exception of the area below the eye spots which so far has been consistently dark. The foot is yellowish green, with the posterior somewhat orange on a few animals, and has a faint white mottling. The large folds on the inner side of the parapodia are salmon to orangish tan. The internal dorsal surface is translucent pale green with cream colored ovotestes show- ing through on either side of the tube leading from the pericardial prominence. Elysia arena lays an orange egg mass with the ribbon being about 1.6mm wide and each 0.25mm capsule con- taining a single egg. Habitat: Elysia arena has only been found in one area of Cocos Lagoon in southern Guam west of a small sand island near Cocos Island. This area has a thick sand sub- strate with very little coral and a water depth of from 0.5 to 1.5m. The animal is directly associated with two species of Caulerpa: C. racemosa and C. cupressoides. Elysia arena differs in its habits from other caulerpan associated Elysza in that it is found just under the surface of the sand around the rhizoids of the algae rather than crawling on the thalli. As is true of many other sand dwelling opisthobranchs, £. arena surrounds itself with a layer of mucus when under the sand. The first animals were found by pulling up the algae and shaking them. Subsequently, animals have been found by searching a shallow area through the sand around the rhizoids of the algae. Occasionally an animal has been seen because of its habit of leaving a slight opening in the sand above the parapodia. Elysia arena has never been found in great numbers in the collecting area and since 1971, only 29 specimens have been examined: 12 in October, 1971; 1 in November, 1971; 5 in October, 1973; 1 in May, 1975; 4 in June, 1975; 3 in July, 1975; and 3 in November, 1975. Remarks: Elysia arena differs from all previously de- scribed Elysia by the presence of the large parapodial folds. Comparison can be made with 2 Elysia that have previously been described as being heavily papillate with conical projections over most of the body — Elysia cauze THE VELIGER Vol. 20; No. 1 Marcus, 1957, and E. papillosa Verrill, 1901 (Marcus & Marcus, 1967: 27 - 28; figs. 22 - 25), both from the West Atlantic. Elysia cauze and E. arena are very similar. From both dissection and serial sections it appears that the genital system of both animals is practically identical. The size of the pharynx with the large annular muscles, the oeso- phagus (Figure 4) opening posteriorly from the pharynx, the radula extending straight into the ascus, and the shape and size of the teeth are all comparable. Besides the presence of the folds inside the parapodia, E. arena dif- fers from E. cauze in the relative size of the parapodia. A 16.0mm preserved E. arena was 14.0mm wide with para- podia unfolded, and a 27.0mm preserved E. cauze was 14.0mm wide when unfolded. Elysia papillosa with a coarsely denticulate radula differs from the finely serrate radulae of both E. cauze and E. arena. The tube extend- ing from the reno-pericardial complex is short in E. papil- losa and extends most of the length of the animal in E. cauze and E. arena. In the Indo-Pacific, a collection of 7 specimens from Hawaii has been described as Elysia aff. cauze (Kay, 1964: 195-196; plt. 9, fig. 3; fig. 2). This form is covered with fine papillae, has a smooth radula, and lacks a transverse furrow. It seems to have little similarity to E. arena and probably represents a distinct species. The specific name is derived from Elysia arena’s habit of burying itself in the sand. Type specimens are deposited in the Bernice P. Bishop Museum, Honolulu, Hawaii. Literature Cited Kay, Euizaszte@ ALison 1964. A new species of Berthelinia and its associated sacoglossans in the Hawaiian Islands. Proc. malacol. Sec. London 36 (3): 191 to 197; plt. 9; text figs. 1, 2 Marcus, Ernat 1957. On Opisthobranchia from Brazil. II. London, Zool. 43 (292): 390 - 486; 246 text figs. Marcus, Eveuine pu Bors-ReyMonp # Ernst Marcus 1967. American opisthobranch mollusks. Studies in tropical oceano- graphy (Univ. Miami Inst. Marine Sci., Miami, Florida), no. 6: vilit+ 256 pp.; figs: 1-155 + 1-95 (22 December 1967) Journ. Linn. Soc. (November 1957) Vol. 20; No. 1 THE VELIGER Page 17 Some Aspects of Spatfall of the New Zealand Rock Oyster During 1974 P DINAMANI anp P. A. LENZ Fisheries Research Division, Ministry of Agriculture and Fisheries Wellington, New Zealand (4 Text figures) INTRODUCTION Manuranci EsTuARY in New Zealand (Lat. 38°30’S; Long. 174°43’E) is an important spat source of the New Zealand rock oyster, Saccostrea glomerata (Gould, 1850) [= Crassostrea glomerata (Gould, 1850)}, being the only area where commercial-scale spatfall had been consistently recorded. Commercial seed collection began in 1968, and Curtin (1971, 1973) has provided spat collection records from 1970 through 1972. Spatfall usu- ally begins early in January and extends into April, with one or two peak settlements occurring around February. The larvae settle in the intertidal region through a 1m vertical range about mean low water level (personal obser- vation). Collectors are usually set out at MLWN about the end of December, and are removed to growing areas at the end of April. The spatting area in Mahurangi is less than 5ha and catches fluctuate. For these reasons methods of increasing production are being investigated: one method is to time the setting out of collectors and the handling so as to get abundant catches of spat and to avoid heavy settlement of fouling organisms that interfere with survival and growth of spat. The summer of 1974 was a successful breeding season which made it possible to study the precise seasonal history of spatfall, patterns of settlement on collectors and other practical problems and procedures in seed collec- tion that are not well understood. The results of these studies are presented and discussed in this paper. MATERIALS anp METHODS Commercial spat collectors were used for all observations for monitoring spat settlement. A standard collector (Fig- ure 1) consists of 30 fibrolite (asbestos-cement) slats, each 1205 X0.6cm, arranged in a bundle of 10 layers and 3 columns, with 5cm between the columns; each layer is separated by 2 wooden spacers, about 12mm thick. Collectors were set out in racks in Huawai Bay in the Mahurangi estuary in the middle of January, and every 4 weeks thereafter 1 collector was examined. A set of collectors was exposed to monitor spatfall between middle of December 1973 and mid-January 1974. For analysis, 1 slat was taken from each layer (usually a slat from the middle column) and the total number of spat was counted on its upper and lower surface, and their sizes measured. The size of the spat refers to its greatest width (THompson, 1968) measured inmm. Where spat density was very high, it was necessary to restrict counting and measuring to one-half of each slat, par- ticularly of the lower surface. In some collectors, with uneven spat distribution, one more slat was examined from each layer to obtain a better estimate of the number of spat/slat. These collectors, examined at 4-weekly in- tervals gave cumulative spat counts (less mortality) for the period January through May 1974. We also made tests to determine the effects of spat col- lector manipulations practised by some farmers. For ex- ample, it is well known that rock oyster settlement is heavier on lower than on upper horizontal surfaces of Fisheries Research Publication No. 288 Figure 1 Arrangement of slats, (a) side view, (b) end view, in a commercial bundle of spat collector; T: top half and B: bottom half of bundle; (c) portion of slat, with U: upper, and L: lower surface the slats, and some farmers overturn their collectors half- way through the spatfall season in the hope of securing an even distribution of spat on both sides of the slat. Our tests were made at the same site and began in February: every month one experimental collector was overturned and at the end of the spatfall season the spat distribution on these collectors was compared with normally handled collectors. Some farmers raise the level of their collectors after the peak spatfall season hoping thereby to avoid settle- ment of competitive organisms like mussels and thus im- prove spat survival. We tested this by shifting one collec- tor in February, March and April from the rack where it received its catch of spat to another rack at the same site that was 30cm higher. At the end of the season spat counts were made on test collectors and compared with counts on collectors that were normally handled. The results observed have been presented in four ways: for each period of cultch exposure, 1) the total count of spat per each collector has been estimated, 2) size frequency distributions (in 5mm groups) of spat within the collector have been analysed, 3) counts of spat settlement on upper and lower surfaces have been pre- sented separately to show preferences, and 4) the slats in each collector have been grouped according to whether they came from the top or bottom 5 layers of the collector (see Figure 1). The terms “upper” and “lower” (within quotation marks) surfaces in the following description refer to new orientations, consequent on collectors being overturned through 180°, but which actually represent the original lower and upper surfaces respectively. THE VELIGER Vol. 20; No. 1 OBSERVATIONS Monitoring collectors exposed from mid-December to mid-January (4 weeks) showed sparse settlement (ap- proximately 3 spat/slat of sizes 1 to 1.5mm), and it is therefore probable that no successful spawning or settle- ment occurred during this period. Most of the seasonal settlement took place after 15 January and Table 1 shows results of spat settlement on experimental collectors that were exposed for approximately 1, 2, 3 and 4 months from mid-January to mid-May 1974. Since collectors were examined only about the middle of each month, precise times of spatfall could not be determined simply from counts of spat. However when considered with size- frequency distribution (Figure 2), and what is known 100 U 80 n= 296 n= 883 n = 136 60 40 z Oo 2 20 Qo = 5 pales full ty & Size group 1 123 1234 12345 : a 2) 20 40 60 80 n= 2862 n= 3038 n=6666 n= 2214 too4 L (a) (b) (c) (d) Figure 2 Size distribution and number of spat on collectors set at the normal level (MLWN) on 19 January 1974, and examined at monthly intervals in (a) February, (b) March, (c) April and (d) May. Spat settled on upper (U) and lower (L) surfaces are shown separately. Size groups: 1 = <5mm, 2 = 5-10mm, 3 = 10-15 mm, 4 = 15- 20mm, 5 = > 20mm Vol. 20; No. 1 THE VELIGER Page 19 about spat growth (normal growth rate is nearly 1mm per week during the summer months — own unpublished data) it is obvious that settlement had been spread out through the 1974 season, with a fairly heavy set in Feb- ruary and again another set after the middle of March. A mean settlement rate of 255 spat/slat was recorded on 19 February when the first collectors were examined (Table 1). The spat averaged 2mm in width suggesting that the first commercial settlement probably began at the end of January/early February. Apparently no heavy spatfalis occurred between February and the middle of March, because collectors examined on 18 March indi- cate a reduction in number of spat from 255 to 238/slat (Table 1), perhaps from natural mortality, and only 12.9% of these were < 2mm wide (Figure 2). However, in the next period, from mid-March to mid-April, there is a marked increase to 417 spat/slat, and nearly 30% of these spat were < 2mm in size, obviously the result of a heavy spatfall during this period. During the second spatfall of the season, the settlement was also heavy in the top half of the collectors. This brought about an increase in the top/bottom ratio of spat from 0.67 for 18 March to 1.09 on 17 April (Table 1). This shows an increase from 37% to 57% of the total number of the spat settling on the top layers of the collector. Table 1 Density of spat settlkement on normal collectors examined at monthly intervals, January through May 1974. Each collector of go slats has been split into two halves, T: top and B:bottom, of 5 layers and 15 slats each. The number of spat on upper(U) and lower (L) surface of each slat has been shown separately. Period Portion No. of No.of Mean No. Spat Total No. Total No. Mean No. Mean spat and Date of slats spat ofspat/ Density ofspaton vo of spat/ ofspat/ _ density/ examined Collector Surface examined measured surface (/cm2) surfaces Total collector slat slat (cm?) Jan-Feb 19 Feb T Ib 7 1083 154.7 0.269 2321 30.3 a U 7 0 0 0 B iL 5 1779 355.8 0.619 5337 69.7 B U 5 0 0 0 1 7658 255.3 0.222 Jan-March 18 March plz L 7 1248 178.3 0.310 2674 37.4 Ty U 7 93 13.3 0.023 200 2.8 B L 7 1790 255.7 0.445 3836 53.7 B U 7 203 29.0 0.050 435 6.1 7145 238 0.207 Jan-April 17 April oy L 10 3902 390.2 0.679 5853 46.7 T U 10 457 45.7 0.079 685 5.5 B Ib 8 2764 345.5 0.601 5182 41.4 B U 8 426 53.2 0.092 798 6.4 12519 417.3 0.363 Jan-May 17 May ay L 5 1258 251.6 0.437 3774 48.6 ay U 5 60 12 0.021 180 2.3 Bi L 4 956 239 0.416 3585 46.2 B U 5 76 15.2 0.026 228 2.9 7767 258.9 0.225 1First spatfall. 2No spatfall. 3Second spatfall. ‘Bottom slats heavily fouled. Page 20 SETTLEMENT PATTERNS in EXPERIMENTAL COLLECTORS (a) Overturned Collectors On 19 February no spat were found on the upper sur- faces of slats in collectors that had been exposed at the normal level for 4 weeks from 19 January. Consequently, when these collectors were overturned on 19 February, what then became their lower surfaces had probably no spat in them. All the spat subsequently recorded on these “lower” surfaces must therefore have settled after 19 Feb- ruary (Table 2). This is borne out by the fact that 61.1% of the spat on the “lower” surface were <.5mm, and only 17% were >10mm (Figure 3). In the controls that were examined at the same time (Figure 2), only 3.2% were <5mm, and 81.1% were >10mm. Corre- spondingly, in experimental collectors, spat counts on “upper” surfaces actually represent settlement that took place on lower surfaces before collectors were overturned. By the middle of March, spat had begun to settle naturally on the upper surfaces of slats as well, reaching a density of 0.036 spat/cm? (sizes <5mm) in normal collectors. The lower surfaces of these collectors showed a spat density of 0.377 spat/cm?. Collectors that were overturned in March showed a higher spat density (0.136 spat/cm?) on “lower” surface but spat were mostly of sizes < 5mm. The “upper” surfaces of overturned collec- tors showed densities of 0.390 spat /cm?, and spat were THE VELIGER Vol. 20; No. 1 80 U 60 n= 1637] |n= 780 n=qar 40 ) tl. Size group 12345 1234 wy n= 2244 n= 2346 N= 2214 n= 136 » fo} Spat number (percent) n = 1928 (a) (b) (c) (d) Figure 3 Size distribution and number of spat on upper and lower surfaces of experimental collectors which were overturned at monthly inter- vals in (a) February, (b) March and (c) April, compared with (d) a normal unturned bundle. All collector bundles removed from racks on 17 May 1974. Other legend as in Figure 1 Table 2 Density of spat settlement on experimental collectors that were overturned in February, March and April 1974. L’ and U ‘refer to original lower and upper surfaces respectively, and therefore denote present U and L surfaces. Similarly T’ and B- are original positions. Date Portion No. of No.of Mean No. Spat Total No. Total No. Mean No. Mean spat Re- of slats spat ofspat/ Density of spat on % of spat/ of spat/ —_density/ oriented Collector Surface examined measured surface (/cm?) surfaces Total collector slat slat (cm?) 19 Feb B 1 5 678 135.6 0.236 2034 19.0 B’ U 5 729 145.8 0.254 2187 20.4 Ts L 5 1250 250 0.435 3750 35.1 alae U" 5 908 181.6 0.316 2724 25.5 10695 356.5 0.310 18 March B Ib 5 1196 239.2 0.416 3588 39.5 B’ U 5 445 89 0.155 1335 14.7 1 L 5 1048 209.6 0.364 3144 34.6 aT U B) 335 67 0.116 1005 iil tt 9072 302.4 0.263 17 April B 1W 5 1164 232.8 0.405 3492 42.1 B’ U 5 158 31.6 0.055 474 5.7 Als Iu 5 1182 236.4 0.411 3546 42.7 a0 U’ 5 263 52.6 0.091 789 9.5 8301 276.7 0.241 Vol. 20; No. 1 THE VELIGER Page 21 of larger size when compared to normal March collec- tors; however both spat modal size and densities (0.421 spat/cm2) were lower than corresponding surfaces of normal collectors examined in May (Figure 3). (b) Collectors Raised to Higher Levels In collectors that were raised 30cm on 19 February, the overall spat density was 0.221/cm? of available sur- face, with 0.368 spat/cm? on lower surface and 0.074 spat /cm? on upper surface (Table 3). The overall spat density was thus very similar to those of normal (control) collectors, which had 0.225 spat/cm?, though the distri- bution of spat was denser on lower surfaces (0.426 spat/cm2) and lighter on upper surfaces (0.023 spat/cm?) of normal collectors. When compared to overturned col- lectors of February, the raised collectors had slightly higher spat densities (-+-0.033/cm?) on lower surfaces but much lower densities (—0.211/cm?) on upper sur- faces (cf. Tables 2 and 3). However, size frequency distributions (Figure 4) show marked differences on sizes of spat, especially on upper surfaces. Collectors that were raised 30cm in March showed an overall density of 0.231 spat/cm?2, a slight increase over those raised in February. The spat were also denser on lower (0.404/cm?) and lighter on upper (0.057/cm2) sur- faces, but were of smaller size. A similar difference was observed when these experimental collectors were com- pared with normal collectors which had also a larger modal size of spat. 80 n=428 n= 330 n = 274 n = 136 60 40 i Al od Size group 1 2 3.45 1234 1234 12345 f 1 80 Spat number (percent) n = 1924 o = 2096 n = 2328 n= 2214 (a) (b) (c) (d) Figure 4 Size distribution and number of spat on upper and lower surfaces of experimental collectors which were raised 30 cm to a higher rack at monthly intervals in (a) February, (b) March and (c) April, compared with (d) a normal level bundle. All collector bundles removed from racks on 17 May 1974. Other legend as in Figure 1 Table 3 Density of spat settlement on experimental collectors that were raised 30 cm from normal level in February, March and April 1974. Legend as in Table 1. Portion No. of No.of | Mean No. Spat Total No. Total No. Mean No. Mean spat Date of slats spat ofspat/ Density — spat on % of spat/ of spat/ density Shifted Collector Surface examined measured surface (/cm?) surfaces Total collector slat (/cm?) 19 Feb T IL, 5 1144 228.8 0.398 3432 44.9 al U 5 172 34.4 0.060 516 6.7 B} L 4 780 195 0.339 2925 38.3 B U 5 256 51.2 0.089 768 10.1 7641 254.7 0.221 18 March T L 5 1186 237.2 0.412 3558 44.7 I U 5 130 20 0.045 390 4.9 B 1G, 4 910 227.5 0.396 3412 42.9 B U 5 200 40 0.069 600 7.5 7960 265.3 0.231 17 April aly L 5 1166 233.2 0.388 3498 44.3 aT U 5 158 31.6 0.055 474 6.0 B L 5 1162 232.4 0.404 3486 44.2 B U 4 116 29 0.050 435 5.5 7893 263.1 0.229 1Bottom slats heavily fouled. Page 22 THE VELIGER Vol. 20; No. 1 Collectors that were raised 30cm in April were nearly identical to those raised in March as regards overall spat density (0.229/cm2), as well as distribution on lower (0.405/cm?) and upper (0.052/cm?) surfaces. They were also similar to experimental collectors overturned in April as regards spat density and modal size. When com- pared to normal collectors (controls) spat size differences were observed, with bigger size spat in larger numbers in the control collectors (cf. Figures 2 and 4). SETTLEMENT on UPPER vs. LOWER SURFACES In normal collectors placed on the racks on 19 January and examined on 19 February, the entire spat settlement was on the lower surfaces of slats; but in normal collec- tors examined towards the end of the spatting season (17 May) 5.25% of spat had settled on upper surfaces of slats. However, collectors examined on 17 April showed a slightly higher rate of settlement (11.86%) on upper surfaces as a result of a second spatfall but these spat had probably failed to survive. This is indicated by the reduc- tion of spat densities on upper surfaces of slats, from 0.085/cm2 in April to 0.023/cm? in May (Figure 2 and Table 1). Analyses of size frequencies reveal that smaller sized spat (~5mm) were greatly reduced in numbers between April and May. Spat distribution was different in collectors that were experimentally treated: those overturned early in the spatting season (February) had somewhat similar spat densities on both surfaces, 0.335 /cm?2 on lower and 0.285 / cm? on upper surfaces, but their size composition showed marked differences (Figure 3); smaller sized spat were predominant on upper surfaces, those < 5mm constitu- ting 61.1% of the spat on the upper surface against 5% on the lower surface. Thereafter, the numbers that settled on upper surfaces of overturned collectors gradually de- creased (Table 2 and Figure 3), so that at the end of the spatting season spat densities were similar to correspond- ing surfaces of normal collectors. In collectors that were raised to higher levels, spat distribution and densities did not vary greatly from month to month, but those shifted in February had slightly fewer spat on their lower surface. This was probably due to the fact that raising the level of the collectors early in the season had caused interruption of spat settlement, and failure of the spat to settle or survive at the new levels. However, spat settlement on upper surfaces of all raised collectors was more than double the density of normal level collectors. SETTLEMENT PATTERNS WITHIN THE COLLECTOR Initial settlement was predominantly in the bottom half of the collectors (Table 1), accounting for 69.7% of the Spat counted on normal collectors in February; however, as the season progressed, the relative number of spat settling in the top half increased, from 40.2% on 18 March to 52.2% on 17 April and, at the end of the spat- ting season, spat numbers were nearly evenly distributed with 50.9% at the top and 49.1% in the bottom half. As a result of differences in spatfall times on the two halves of the collectors, size differences were noticed: thus in the bottom half spat were predominantly (>77%) of larger size (15mm and above) and there were no spat smaller than 5mm; in the top half nearly 30% of the spat were < 10mm and only 53% were > 15mm. In experimental collectors (Tables 2 and 3) the same trends were seen, the degree of difference being related to the time at which the collectors were re-oriented. Thus collectors overturned in February showed greater spat density and settlement on slats which were originally in the top half, but small sized spat (<( 5mm) occurred in greater number in the original bottom half of the collec- tor. However, collectors overturned in March showed lesser spat density and settlement in the original top half, while collectors overturned in April showed slightly higher numbers in the same half. The percentage of small sized spat in the original bottom half of the collector was some- what higher than in the layers of the top half in both March and April collectors. In collectors that were shifted to higher level in different months, top halves of collectors showed on each occasion higher percentage of spat, which were made of small sized spat (< 10mm) particularly in those shifted in March and April. SETTLEMENT or COMPETITIVE ORGANISMS. (a) Other Oysters Spat of two other oyster species are sometimes found on collectors laid out for rock oysters (DinaMANt, 1971). These were an unidentified flat oyster, Ostrea sp., and the Pacific oyster, Crassostrea gigas (Thunberg, 1793), which are somewhat difficult to distinguish from one another at early spat stages. Flat oyster spat were rare in 1974, only 8 - 10 spat/bundle were noted on 19 February, and none were found during the three subsequent monthly samp- Vol. 20; No. 1 lings. However, Pacific oyster spatfall was heavier than in previous seasons (DiNAMANI, 1974b): Table 4 gives de- Table 4 Pacific oyster spat recorded on collectors examined on 17 May 1974. THE VELIGER No. of spat Type of upper lower Total spat collector surface surface /collector 1. Normal collector 27 69 96 2. Overturned collectors: Reoriented Feb. 741 159! 213 Reoriented Mar. 361 130! 166 Reoriented Apr. 66! 67! 133 3. Collectors raised 30 cm: Raised Feb. 42 55 97 Raised Mar. 24 50 74 Raised Apr. 28 36 64 1Refers to present upper and lower surface. tails of the Pacific oyster settlement on collector samples examined during the season. It may be observed that the Pacific oyster settled in densities of 96 spat/bundle of normally handled collectors, 70% of them being on the lower surfaces of slats. Maximum settlement (213 spat/ bundle) was observed in collectors that were overturned on 19 February. The distribution of these spat indicates that most settled on lower surfaces of slats between 19 February and 18 March, just before and soon after the collectors were overturned: this is revealed by the fact that the settlement of the Pacific oysters gradually de- creased through March and April, and also because col- lectors overturned in March and April caught more Pacif- ic oyster spat than the normally handled collectors and by the fact that numbers of spat on upper and lower sur- faces of overturned collectors in April were almost equal. (b) Barnacles Settlement of the barnacle, Elminius modestus, on spat collectors was light in 1974 compared with other years. Normally handled collectors exposed from mid-January to 19 February showed barnacle settlement densities be- tween 0.15/cm? in the top half, to 0.75/cm?2 in the bottom half. Collectors exposed for longer periods showed moder- ate to heavy barnacle settlements (15 to 20/cm?). Over- turned collectors also showed moderate to heavy settle- Page 23 ment, but collectors raised to higher level showed an even denser settlement rate; this was particularly true of col- lectors that were raised in March and April, which had a heavy settlement of > 25 barnacles/cm? as well as a dense deposit of silt on the upper surfaces of the slats. GrEENWAY (1969) has recorded denser barnacle settle- ment at higher levels in Huawai Bay. (c) Polyzoans A cheilostomatous polyzoan, Watersipora cucullata, was found to settle in isolated small groups (1 to 4 polyp- ides each) in all collectors examined from March on- wards, and was most numerous on middle layer slats of collectors overturned in March. It normally appears on the shaded surfaces of slats and grows over settled spat in the form of a thick mat. DISCUSSION Seasonal differences in intensity of rock oyster settlement may be attributed to variable environmental factors such as water temperature which affects spawning, where- as patterns of settlement are usually thought of as depend- ent on behavioral characteristics of larval oysters. The first major settlement of rock oysters occurred about the end of January in 1974 but settlement has varied from year to year; our observations (unpublished data) show that it took place in middle and late February in 1973, and after the middle of January in 1972. A second settle- ment maximum usually occurs in February or March, depending upon the timing and duration of spawning (Dinamani, 1974a). However spatfalls have been ob- served as late as the end of April or early May in years when the first spawning has been delayed. Observations on spawning, as indicated by plankton monitoring of oyster larvae, and on spatfalls, as recorded in commercial collectors, showed the following pattern: oyster larvae (80 - 150 um in length) appeared in Huawai Bay during the 3 week of December 1973, and were common up to the end of the month. Appreciable num- bers (average 40 - 60 larvae/500/ sample) were not found until after the middle of January. These observations checked well with spatfalls on trial collectors exposed from mid-December to mid-January, which showed only 3 spat/slat all < 1.5mm in size. Late stage larvae were found from the last week of January, and the first spat on our monitoring collectors were observed about the middle of February. Thus the evidence points to a major spatfall (settlement rate of ca.0.250 spat/slat) during the first week of February. The length of larval life has been reckoned Page 24 to be about 18 to 24 days (Dinamani, 1973) at a tem- perature of 21 - 22°C. From the end of February to the middle of March, no regular larval monitoring was possible in the Mahurangi area, but experimental collectors examined in the middle of March showed only a few spat of < 2mm (about 12% of the total), and the mean number of spat/slat on these collectors was also lower than that of February. This probably meant that there were few larvae in the water and very little settlement occurred during that period. However, plankton samples taken from 18 March on- wards revealed late-stage larvae, which suggested that a second peak in larval numbers probably occurred in March, and gave rise to a second spatfall. The large num- ber of small spat observed in the collectors examined in April points to this. Thus Mahurangi ordinarily has a long breeding season (GrEENWAY, 1969), with probably two spatfall maxima from January to April. In the Australian rock oyster, Tuompson (1950) recorded two maxima during the spring (November) and autumn (March). We have how- ever no conclusive evidence of spring spawning in the New Zealand rock oyster (DinAMANI, 1974a). Saccostrea glomerata larvae like those of many other species of oyster settle more heavily on lower than upper surfaces of collectors (NELson, 1927; Hopkins, 1935; ScHAEFER, 1937; CoLE & KNIGHT-JoNES, 1939; SrEBING, 1950; THompson, 1950; Mepcor, 1955). This charac- teristic is well demonstrated by our data for collectors that were experimentally overturned at various times in 1974 (Table 2, Figure 3), the subsequent settlement being largely confined to the ‘new’ lower surface. There was also a tendency at the beginning of the settlement season for the spat to settle in the bottom half of the collector bundle as demonstrated in data listed in Table 1. How- ever towards the end of the settlement season, roughly equal numbers of spat are found in the top and bottom halves of the collector bundles. This was shown by the greater percentage of small-sized spat settling on the top half of the bundle after February. This may indicate a tendency, under natural conditions, for late-season settle- ment to occur higher in the intertidal zone than early season settlements. There is also a tendency for the settlement to be almost exclusively on the lower surfaces early in the season, but to occur in small numbers (8 to 10% of the total) on the upper surfaces late in the season. This could be taken as evidence that oyster larvae avoid settling on lower sur- faces that are already heavily populated by their own species. KnicHT-JONES & STEVENSON (1951) and WISELY (1959) have observed that settling larvae tend to avoid THE VELIGER Vol. 20; No. 1 surfaces where recently settled individuals of their own kind have attained a certain density. The term ‘gregari- ousness’ has been used to indicate the selection by settling larvae of surfaces associated with organisms of the same species, though, as pointed out by Bayne (1969), it also ‘Gmplies a response by the larvae to the previously settled individuals of the same species.” This response could therefore either favour aggregation, or result in an avoid- ance reaction, depending upon the density of the individu- als on a surface at any time. In the rock oyster the results of settlement on commercial collectors indicate that larvae probably begin to settle on upper surfaces of slats when spat densities on lower surfaces become high. In this type of collector, as the distance between the lower sur- face of one slat and an adjoining upper surface of another is only 12mm, gregarious response could be a factor. In a series of experiments on the settlement of Crassostrea virginica (Gmelin, 1791), SHAw (1967) reported that more larvae settled on upper surfaces when slats were about 25mm apart, but only on lower surfaces when the slats were 100mm apart. Analyses using ‘t-test’? values for differences in mean spat density between normal and experimental collectors show significant values only for collectors overturned in February (¢ = 2.147, P= 0.05) and March (t = 2.290, P —0.05), compared to normal collectors exposed for the same period. No significant differences were observed in mean spat density between normal collectors and raised collectors. However, it is obvious that in spite of increases in spat density in collectors overturned earlier in the sea- son, spat sizes in these are generally smaller than in normal collectors (cf. Figures 2 and 3). This is also true of collectors raised to higher level. Therefore, reorienta- tion of cultch material, either by overturning or raising during the spatting season, most probably affects spat growth. It remains to be seen whether the more even dispersion of spat on both surfaces of slats (a feature of overturned collectors) may, in later seasons, have the advantages of lesser overcrowding and hence better growth. This has to be weighed against the higher settle- ment rate of competitive species such as the Pacific oyster in the same collectors. No obvious benefits, ¢. g., higher survival of spat or abatement of barnacle settlement, were seen in collectors shifted to higher level. On the other hand, there were signs of higher silt deposition, which normally smothers small sized spat. Spat which settle later in the season generally have a poorer survival rate, as is evidenced here by the considerable drop in percentage of small size spat between April and May in normal collectors, as also in those overturned in April (see also GREENWAY, 1969). Vol. 20; No. 1 CONCLUSIONS 1. Rock oyster commercial seed operations should pre- ferably be initiated to coincide with the period of the first major spawning. This would ensure clean cultch surfaces for maximal settlement, better growth and survival of spat, with minimal fouling by organisms such as Elminius modestus, which have several spawning peaks and could settle at the same level. The extended oyster breeding season from January to March and the occurrence of at least two spawning maxima provide some safety factor for farming operations. 2. Plankton monitoring programmes which provide in- formation on spawning activity and larval abundance might be of use to the seed industry, to enable the growers to catch the first set and to help them plan ahead. As pointed out by LoosaNnorF & ENcLE (1940), information on spawning time, intensity of setting and the probable rate of survival of spat under different conditions, would lead to better planned oyster farming. A monitoring pro- gramme was in fact tested during the 1975 season, and with refinement it may permit forecasts of dates and sizes of spatfall. 3. The study has also indicated that spatfall on the present type of collectors follows a seasonal pattern: initially spat settle almost exclusively on lower surfaces of slats, in the lower halves of bundles that are positioned close to MLWN;; as the season progresses, spat may settle on both upper and lower surfaces of collector slats, in both top and bottom halves of collector bundles, and at higher levels in the intertidal zone than at the beginning of the spatfall season. Thus, at the end of a good season all slats in the bundle have a somewhat even distribution of spat. 4, The study also reveals that spat which settle early in the season survive better and are larger at the end of the season than spat which settle later in the season. It may therefore be worthwhile to maintain a monitoring pro- gramme of planktonic populations of oyster larvae to fore- cast the first sets of the year. 5. The study shows that turning collectors up-side-down during the early part of the spatting season may increase their catch of spat: e. g., collectors overturned in Febru- ary took 37% more spat than normally handled (un- turned) collectors, and collectors overturned in March took 17% more spat. In both cases, however, the increase counts were largely due to the greater number of small- sized late-settling spat. In contrast, the overturned col- lectors have fewer spat of sizes over 10mm compared to the normal collector. THE VELIGER Page 25 6. Another factor is the possibility of settlement of other organisms such as the Pacific oyster on the ‘new’ sur- face; collectors overturned in February had more than double the number of Pacific oysters of normal collectors, and collectors overturned in March had more polypides of Watersipora. The incidence of other competitive species and the time of their settlement may have a marked effect on rock oyster spat, particularly their growth. 7. In good breeding seasons, with at least two spatfalls between January and March, normal settlement patterns provide adequate spat densities and growth in commercial type collectors. Whether there is need for special handling of collectors is an open question. SUMMARY Settlement of the New Zealand rock oyster, Saccostrea glomerata (Gould) [= Crassostrea glomerata (Gould) } on bundles of slat-type commercial collectors in the Ma- hurangi estuary in New Zealand, showed two peaks of spatfall between January and March 1974 in a definite pattern: the first sets were predominantly on the lower surface of slats and on the lower layers of slats in the bundles. As the season progressed sets were still largely on lower surfaces but spread more uniformly through the several layers of collector slats. By overturning the bundles at intervals during the season, significantly higher (P = 0.05) mean density of spat was obtained if carried out in the earlier part of the season, but brought about a re- duction in maximum spat size, and also a greater settle- mentment of competitive species such as the Pacific oyster. Moving collectors at intervals to higher tidal level did not affect spatfall appreciably but increased silting on collector surfaces. ACKNOWLEDGMENTS We are grateful to Dr Carl Medcof, St. Andrews, Canada, for his valuable suggestions and criticism, and to Messrs G. Duncan Waugh and Carel Vooren of the Fisheries Research Division, New Zealand, for helpful comments on the typescript. Literature Cited Bayne, B. L. 1969. The gregarious behaviour of the larvae of Ostrea edulis L. at settlement. Journ. Mar. Biol. Assoc. U. K. 49: 327 - 356 Page 26 THE VELIGER Vol. 20; No. 1 Core, Hersert Ausrey & E. W, Knicut-Jones 1939. Some observations and experiments on the settling behaviour of larvae of Ostrea edulis. Journ. cons. perm. int. Explor. Mer. 14: 85 - 105 Curtin, L. 1971. Marine Department Rock Oyster Catching Programme 1970-71. Fish. Tech. Reprt. 74: 8 pp. N. Z. Mar. Dept. 1973. Rock Oyster Spat Catching Programme 1971-72. Reprt. 110: 48 pp. N. Z. Ministr. Agric. & Fish. DINAMANI, PARAMESWARA 1971. Identification of oyster species competing with rock oysters for settlement space. Fish. Res. Div. Inf. Leafl. 1: 9 pp.; 6 figs. 1973. Embryonic and larval development in the New Zealand rock Oyster, Crassostrea glomerata (Gould, 1850). The Veliger 15 (4): 295 - 299; 3 plts.; 2 text figs.; 1 map (1 April 1973) 1974a. Reproductive cycle and gonadal changes in the New Zealand Rock Oyster, Crassostrea glomerata. N. Z. Journ. Mar. Freshwat. Res. 8: 39 - 65 1974b. Pacific oyster may pose a threat to rock oyster. Catch ’74 6: 5-9. Fish. Manag. Div., Min. Agric. & Fish., New Zealand GREENWAY, J. PC. 1969. The monitoring of rock oyster setlement at Mahurangi, Auck- land 1968-69. Fish. Tech. Reprt. 48: 21 pp. N. Z. Marine Dept. Hopxins, AuBreEY EDWIN 1935. Attachment of larvae of the Olympia oyster, Ostrea lurida, to plane surfaces. Ecology 16 (1): 82-87 Knicut-Jonzs, E. W. & J. PR STEVENSON 1951. Gregariousness during settlement in the barnacle, Elminius mo- destus Darwin. Journ. Mar. Biol. Assoc. U. K. 29: 281 - 297 Fish, Tech. Loosanorr, Victor Lyon « James B. ENGLE 1940. Spawning and setting of oysters in Long Island Sound in 1937, and discussion of the method for predicting the intensity and time of oyster setting. Bull. U. S. Fish. 49: 217 - 255; 11 text figs.; 18 tabl. Mepcor, J. C. 1955. Day and night characteristics of spatfall and of behavior of oyster larvae. Journ. Fish. Res. Brd. Canada 12 (2): 270-286 NELSON, THURLOW CHRISTIAN 1927. | Report of Department of Biology for the year ending June 30, 1926. New Jersey Agric. Exper. Sta. 103 - 113 ScHAEFER, MILNER B, 1937. Attachment of the larvae of Ostrea gigas, the Japanese oyster, to plane surfaces. Ecology 18 (4): 523 - 527 Suaw, WIii1aM N. 1967. | Seasonal fouling and oyster setting on asbestos plates in Broad Creek, Talbot County, Maryland, 1963 - 65. Chesapeake Sci. 8 (4): 228 - 236 SreBLinG, F, W. 1950. Influence of seasoning and position of oyster shells on oyster set- ting. Proc. nation. Shellfish Assoc. 41: 57-61 TuHompson, J. M. 1950. The effect of the orientation of cultch material on the setting of the Sydney rock oyster. Austral, Journ. mar. freshwat. Res. 1: 139 - 154 1968. A note on oyster dimensions. Proc. Sympos. Mollusca held at Cochin, Jan. 1968. Mar. Biol. Assoc. India Wisexy, B. 1959. Observations on the settling behaviour of larvae of the tubeworm Spirorbis borealis Daudin (Polychaeta). Austral. Journ. mar. fresh- wat. Res. 11: 55 - 72 Vol. 20; No. 1 THE VELIGER Page 27 A New Species of Chiton from the Aleutian Islands (Mollusca : Polyplacophora ) ANTONIO J. FERREIRA ! 2060 Clarmar Way, San Jose, California 95128 (1 Plate; 1 Text figure) Tue ALEUTIAN ISLANDS, extending westward some 2000 km from the Alaskan peninsula towards the peninsula ot Kamchatka, are an integral part of the Aleutian Province which, variously defined (ScHENCK & KEEN, 1936; Ex- MAN, 1953; BRIGGS, 1974), stands between the eastern Pacific Cold Temperate Region to the south, and the icy waters of the Arctic Region to the north. For mollusks in general, the endemism of the Aleutian Province has been estimated at 24% (VALENTINE, 1966). Still, its chiton fauna is poorly known. This paper reports on a new species of chiton which appears to be endemic to the Aleutian Islands. POLYPLACOPHORA de Blainville, 1816 NEOLORICATA Bergenhayn, 1955 IscHNOCHITONINAE Bergenhayn, 1930 IscHNOCHITONIDAE Dall, 1889 Ischnochiton Gray, 1847 Ischnochiton allyni Ferreira, spec. nov. (Figures 1 to 4 and 5) Diagnosis: Chiton of moderate size, uniform rusty- brown color. End valves with 15 - 20 radial ribs, often bifurcated, crowned by minute tubercles, and separated by well defined sulci. Lateral areas with 3 - 4 similar ribs. Central areas uniformly pitted for a net-like effect. Girdle covered with imbricating, relatively large, mam- milated, faintly striated scales. Articulamentum of inter- mediate valves with 2 - 3 slits per side. « Research Associate, Department of Invertebrate Zoology, Cali- fornia Academy of Sciences, San Francisco, CA 94118 Description - Holotype: Oval shaped with quasi par- allel sides, circular in front and in back. Dried, but fully extended, it measures (including the girdle) 19mm in length, 12mm in width, and 4.5mm in height. Width/ length ratio = 0.63. Jugal angle about 103°. The teg- mentum and girdle are a uniform rusty-brown color. The tegmentum is microgranulose throughout. The anterior valve displays some 22 radial ribs, most of which bifur- cate resulting in about 36 ribs when counted at the valve’s periphery. The radial ribs tend to have a distinct tri- angular outline in cross-section; they are crested by a row of minute tubercles (about 0.05mm in diameter), often poorly defined, close together, sometimes fused. The radial ribs are neatly separated by a sulcus. The posterior valve has a well defined but not prominent mucro; the post mucro area is plane and shows about 16 radial ribs in every respect similar to those in the anterior valve. The central areas of the intermediate valves are grossly but uniformly pitted resulting in a net-like ap- pearance. The lateral areas are well defined and moder- ately raised; they exhibit 3 - 4 radial ribs with the same characteristics of those in the end valves. Girdle not band- ed, about 2mm in width, covered with loosely imbricated scales. The girdle scales (Figures 3, 4) are strongly con- vex, weakly striated, often reaching 250m in length; their dorsal edge tends to point inwardly, and is usually crowned by a striated mammillus. Towards the periphery, the girdle scales are much smaller, and columnar in shape. The gills, about 28 on each side, extend from about 2mm in front of the anus to about 3mm behind the anterior edge of the foot. The articulamentum is white. Insertion teeth are sharp and straight edged. The slit formula is 16-2/3-14. Eaves are small and subspongeous. The sutural laminae are sharp and semi-oval continuing without notch or demarcation with a thin sinusal lamina which protrudes 0.1 - 0.2mm in front of the anterior edge of the tegmentum. The sinus is well developed but moder- Page 28 THE VELIGER Vol. 20; No. 1 ate in size. The radula (Figure 5) is 9.5mm long and Paratypes: 23mm, and 36mm in length; same color as contains about 45 rows of teeth. The uncinate plate holotype. (major lateral) is unicuspid. Type Locality: North side of Constantine Harbor, Am- chitka Island (51°30’N; 179°00’W), Aleutian Islands, Alaska. The 3 specimens, here designated as holotypes and paratypes, were collected by L. Barr eé al., with SCUBA, on a rock substrate, in 17 - 27m (50 - 80 feet) of water on June 12, 1973; they were made available through the generosity of J. M. Barnes, Brigham Young University, Provo, Utah. Type Material: The holotype, partly disarticulated (CASIZ Type Series No. 683; Type Slide No. 496), and the two paratypes (CASIZ, Type Series Nos. 684 & 685), together with color slides of the specimens (CASIZ, Col- or Slides Series, Nos. 2949 - 2951) are deposited in the California Academy of Sciences, San Francisco. Remarks: The question of a subgeneric assignment for Ischnochiton allyni cannot be readily decided at this time. Similarities in tegmental sculpture, girdle scales and articulamentum suggest that J. allyni may have a not Zoleyriss too remote kinship with J. trifidus (Carpenter, 1864) from the adjacent eastern Pacific Cold Temperate Region, and for which Berry (1919) erected the monotypic subgenus Tripoplax. However, I. allyni with its Lepidozona-like features and radsioid valves seems to have an even greater affinity with a group of species described from the Sea ves of Okhotsk and the northern part of the Sea of Japan, oe in the genera Gurjanovillia and Lepidozona. The close examination of specimens graciously donated by Dr. B. Sirenko, University of Leningrad, U.S.S.R., suggests a phylogentic tie between J. allyni and Gurjanovillia alb- rechti (Schrenck, 1867), G. lindberghi Jakovleva, 1952, aa L. multigranosa Sirenko, 1975, L. thiele: Sirenko, 1975, and L. 7ma Sirenko, 1975. The synonymization of the genus Gurjanovillia Jakov- leva, 1952 (Type species: Chiton albrechti Schrenck, 1867, by OD) under Lepzdozona Pilsbry, 1892 (Type species: Chiton mertensii Millendorff, 1847, by OD), as previously proposed (SmirH, 1960; FERREIRA, 1974; SI- Figure 5 RENKO, 1975) is incorrect: The species listed by Jakov- Ischnochiton allyni Ferreira, spec. nov. leva in the genus Gurjanovillia are radsioid, 7.é., 2-slitted, Holotype - radular teeth [Camera lucida drawing] and therefore not members of the genus Lepidozona Explanation of Figures z to 4 Figure 1: Ischnochiton allyni Ferreira, spec nov. Paratype; length Figure 3: Ischnochiton allyni Ferreira, spec. nov. Holotype — girdle 36 mm [Allyn G. Smith, photograph] scales [SEM micrograph by Hans Bertsch] approx. X 62 Figure 2: Ischnochiton allyni Ferreira, spec. nov. Paratype, close- Figure 4: Ischnochiton allyni Ferreira, spec. nov. Holotype — girdle up of valves i and ii [Allyn G. Smith, photograph] scales [SEM micrograph by Hans Bertsch] approx. X 186 Tue VeE.iocErR, Vol. 20, No. 1 [FrrRruma] Figures 7 to 4 Tesiae oe ° * 2 > ‘ ” , . ; AP j tied \ f q F f . n Rae ‘ ~ te . 4 v 1 Py i Yh sedi a3 ‘| | f fk 0 : ‘ | > i oP at 7 | } o eae | ae a | so Ah = ; uy 5 fm 1 it é j . ‘ ae Gai A ~ eons f . fons a lie if iL : , P 2 i : y =| rs 7 } , ' is i ¥ 1} : 5 “y n {| 7 J { Ba § { t | % ; i : i i ‘ e eee 5 He 5 } rok j i f 2 ? i ae is $ Z E hae : it 4 1 a i ‘ \! j ‘ ; s f ; i h MI \ : : < 3 i ) 5 bf \ j ‘ \ i ; Fa 2 : 3 1 F ; j i i 4 * | : ne ‘ Te ; ne : 1. | - - = - uy é 1 h Find a if e 1 5 ] 2 : \ ' ‘ y i} =e tA es | i j i aan { Up i ea 7 J | 7 ay =f I 5 i we a wr q . i , ; ; ~ : ‘ } : r i i ; ‘v, , * i yn : ‘ . uf N ; ioe : : r as , i 2 Ree ae ot wer A :: ; a , ¥ Aver =| uy ’ Ps ? Vol. 20; No. 1 THE VELIGER Page 29 nnn ne rere eS ernst oaeecneeeneeeer? which (sensu FERREIRA, 1974) contains 1-slitted species exclusively. Thus, the taxon Gurjanovillia must be taken out of the synonymy of Lepidozona and considered anew. The likely kinship between Ischnochiton allyni and the above mentioned northwestern Pacific species suggests the possibility of grouping them at the subgeneric or even generic level. However, the problem is complicated by the fact that the genus Ischnochiton Gray, 1847 (Type species: Chiton crispus Reeve, 1847, by SD, Kaas, 1974) is much in need of revision, and the definitions and differ- ential diagnosis of its all too numerous subgenera remain unsettled. So, for the moment, whether J. allyni would be better assigned to the now available genus Gurjanovillia, or to one of the ill-defined subgenera of Ischnochiton must remain an open question pending further investiga- tion. Under the circumstances, and until a clearer per- spective of these taxa can be achieved, it seems appropri- ate to retain J. allyni in the genus Ischnochiton with no subgeneric assignment. The apparent phylogenetic relationship between Isch- nochiton allyni, seemingly endemic to the Aleutian Pro- vince, and the above mentioned group of species from the Sea of Okhotsk Province suggests that a common ancestor might have inhabited the northernmost part of the Pacific in a possibly continuous distribution between the two great coastal arches. Conceivably, J. allyni (and perhaps also J. trifidus) stemmed from the eastern part of that original stock, and unto the present has remained separated from counterparts in the northwestern Pacific by the deep waters of the North Aleutian Basin. The species is here called allyni after a great man, Allyn Goodwin Smith, who guided my steps and inspired much of my work. ACKNOWLEDGMENTS Appreciation is here expressed to Lou Barr, Auke Bay Fisheries Laboratory, Alaska, who collected the specimens and provided some additional data; to Dr. James Barnes, Department of Zoology, Brigham Young University, Pro- vo, Utah, who generously donated the specimens; to Dr. B. Sirenko, Academy of Sciences of USSR, Leningrad, USSR, and Dr. Iwao Taki, Hiroshima University, and Dr. Kohman K. Arakawa, Hiroshima Fisheries Experi- mental Station, Hiroshima, Japan, who have enriched our knowledge of the north western Pacific chiton fauna with the donation of many specimens; to Hans Bertsch, Donner Laboratory, University of California, Berkeley, for the SEM micrographs; and to Dustin Chivers and Dave R. Lindberg, Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, for their critical reading of the manuscript and assistance in sever- al other phases of this work. Literature Cited Berry, SAMUEL STILLMAN 1919, | Notes on west American Chitons, II. (4) 9(1): 1- 36; plts. 1-8 Briccs, Joun C. 1974. Marine Zoogeography. 475 pp. EKMAN, SVEN 1953. Zoogeography of the sea. xiv+417 pp.; 49 tables; 121 text figs. FERREIRA, ANTONIO J. 1974, The genus Lepidozona in the Panamic Province, with the de- scription of two new species (Mollusca : Polyplacophora). The Veliger 17 (2): 162-180; 6 plts. (1 October 1974) JAKovLEvA, A. M. 1952. Shell bearing mollusks (Loricata) of the seas of USSR. Acad. Sci. USSR, no. 45, Keys to the Fauna of USSR, 127 pp. (Jerusalem transl. 1965) Kaas, P. 1974, Notes on Loricata, 6 - 7. ScuHeEnck, H. G. a A. Myra KEEN 1936. Marine molluscan provinces of western North America. Proc. Amer. Philos. Soc. 76 (6): 921-938; figs. 1-6; table SrrENko, B, I. 1975. On the taxonomy of the genus Lepidozona Pilsbry. Mar. Biol. No. 3, Acad. Sci. USSR, pp. 13 - 28 (in Russian) SmitH, AttyN Goopwin 1960. Amphineura. In Treatise on Invertebrate Paleontology, ed. Moore, Part I, Mollusca 1, pp. 41 - 76; figs. 31-45 VALENTINE, JAMES WILLIAM 1966. Numerical analysis of marine molluscan ranges on the extra- tropical northeastern Pacific shelf. Limnol. & Oceanogr. 11 (2): 198 - 211 (2 April 1966) Proc. California Acad. Sci., (16 June, 1919) McGraw-Hill Book Co., New York London: Sidgwick & Jackson, Basteria 38: 93 - 97 Page 30 THE VELIGER Vol. 20; No. 1 Homing in Urosalpinx cinerea in Response to Prey Effluent and Tidal Periodicity DAVID M. PRATT Graduate School of Oceanography, University of Rhode Island, Kingston, Rhode Island 02881, USA {1 Text figure) INTRODUCTION Ar tHe Unrversiry oF Roope Istanp’s Narragansett Bay Campus, two parallel breakwaters extend at right angles from a sandy beach out into the bay, forming a basin for small boats. They are made of jumbled angular blocks of undressed stone up to 1m%, are heavily popu- lated with Balanus balanoides (Linnaeus, 1758) and, from June to September with oyster drills, Urosalpinx cinerea (Say, 1822), feeding apparently exclusively on the barnacles. The floor of the basin is flat, its sediment a coarse sand. Oyster drills are never found on the sandy shoreline between the breakwaters, and during the sum- mer the two snail populations are discrete. The situation raised the following questions: 1) if snails from the two populations were deposited at an intermediate point, would they tend to home to the breakwaters from which they were taken? and 2) how quickly would they return? MATERIAL anp METHODS On July 8, 1975, 1625 oyster drills were taken from the inner side of the southern breakwater, which is 35m long, and 1928 from the inner side of the northern breakwater, 41m long. They were spray painted with fast drying enamels: the southern snails yellow, the northern snails white; and within 5 hours of capture all were returned to the middle of the boat basin where at Mean Low Water the water is 1.6m deep and the breakwaters are 20m apart at water level. The assistance of Brian Melzian in this day’s task is gratefully acknowledged. For the next 75 days the breakwaters were searched daily at or near low tide, allocating time and effort in approximate proportion to the length of each breakwater. Marked snails were collected and taken to a distant area for release. RESULTS anp DISCUSSION The results are summarized in Table 1. The snails’ ten- dency to return to the breakwater of their origin was quantified in a normal approximation to a test of pro- portions, which yielded the highly significant value of 5.235 (tooo. == 3.291). The probability of no homing response, 7. €. random dispersal, is therefore less than one in a thousand. What directional clue they used is not known. Table 1 Number of oyster drills marked and recaptured North South Breakwater Breakwater Total marked 1928 1625 3553 recaptured on break- water of origin 350 234 584 recaptured on oppo- site breakwater 165 254 419 total recaptured 515 488 1003 Strong as this statistical result is, the homing response was complicated by an attraction to the north breakwater that affected not only the snails originating there (68% of recaptures) but also those from the south breakwater (52% of recaptures). This may be explained as follows. Vol. 20; No. 1 Balanus balanoides exerts a powerful olfactory distant attraction for this oyster drill population (Pratt, 1974). There are more barnacles on the north breakwater, which is not only longer than the south breakwater but also is thickly populated with barnacles for more of its length. In addition, the tide along this shore ebbs southward with visible velocity but on the flood no northward cur- rent is perceptible. The presumed greater volume of barnacle effluent and rate of flow from the north may combine to explain why slightly more snails from the south breakwater were recaptured on the north break- water than on the south. The temporal pattern of the snails’ return to the break- waters can be seen in Figure 1, which shows the numbers of marked snails recaptured daily (all results combined). Thirteen days elapsed before the first returning snails accomplished the 10m trek along the bottom from the 60 50 40 30 Snails Recaptured 20 ide Level Low THE VELIGER Page 31 release point. Their return occurred in waves, of which at least 3 are clearly shown and a fourth is suggested. These waves came during periods of neap tides as can be seen in the correspondence of daily returns with the daily means of the predicted semidiurnal low tides. The lower edge of the band of barnacles is at about 5cm above Mean Low Water. During spring tides, these prey of the snails are exposed to air for varying periods twice a day. Only during neap tides are they continuously im- mersed and infusing the water with their chemical at- tractant. This may account for the biweekly pulsed re- turn of the oyster drills to the breakwaters. The oyster drill populations of the breakwaters, num- bering several thousands in July and August, declined sharply in September. Water temperatures ranged be- tween 20 and 23°C during July and August except for readings of 24-26°C for August 2-5; the September September Figure 1 A. Daily recaptures of marked snails on the breakwaters B. Daily averages of low tide levels (cm) Page 32 THE VELIGER Vol. 20; No. 1 temperatures declined from 19.9 to 18.0°C. Oyster drills normally disappear from the breakwaters in October, presumably to overwinter on the bottom of the boat basin (CarrIkKER, 1954, 1955). The return of marked snails to the breakwaters during the September neap tides was counter to, and largely obscured by, the seasonal down- ward mass movement. Movements on the order of a meter and an hour or two, which return the animal to its approximate starting point, are known in a number of prosobranch and pul- monate snails, their precision and adaptive advantage varying with the species’ way of life. Individual peri- winkles, Littorina littorea (Linnaeus, 1758), make short feeding excursions during a tidal cycle in a U-shaped course that maintains their vertical position on the shore- line (NEWELL, 1958). Limpets, whose homing has been studied for nearly 150 years, home with great precision to their individual scars on the rock surface where they can withstand the molar action of waves (MacGrniTIE & MacGrinirtiz, 1949). In spite of considerable experi- mentation (Cook, 1969; Coox et al., 1969) their navi- gational systems remain obscure. FUNKE (1968) lists 34 species of marine gastropods (29 of them limpets) and 10 species of land snails but none from fresh water that are known to have homing abilities. The homing tendency of Urosalpinx cinerea is on larger spatial and temporal scales than those described for other snails. Quantities and concentrations of prey ade- quate for the summer’s foraging of a population of oyster drills are highly localized. The drill’s habit of overwinter- ing at some distance from its food supply, coupled with its limited powers of locomotion, would be a serious drawback in a random springtime search for a summer feeding ground. These disadvantages are overcome by any tendency to migrate in an appropriate direction. Upward movement from the deeper overwintering grounds will provide enough direction in some situations but in others will misguide the snail. Here the chances for success will be improved if the geotactic response is supplemented by some other clew that has previously rewarded the snail. The oyster drills’ decided preference for the home breakwater suggests such an additional directional impulse. Literature Cited Carrikzr, MELBOURNE ROMAINE 1954. Seasonal vertical movements of oyster drills (Urosalpinx cinerea). Proc. natn. Shellfish Assoc. 45: 190 - 198 1955. Critical review of the biology and control of the oyster drills Urosalpinx and Eupleura. Spec. Scient. Reprt. U.S. Fish & Wild- life Serv. (Fish.) 148: 1 - 150 Cook, A., O. S. Bamrorp, J. D. B. FREEMAN « D. J. TEDEMAN 1969. A study of the homing habit of the limpet. Anim. Behav. 17 (2): 330-339; 9 figs.; 1 table Coox, Susan BLacKFrorpD 1969. Experiments on homing in the limpet Siphonaria normalis. Anim. Behay. 17 (4): 679-682; 1 fig. Funke, WERNER 1968. Heimfindevermégen und Ortstreue bei Patella L. (Gastropoda, Prosobranchia). Oecologia 2: 19 - 142; 25 figs.; 9 tables MacGinitiz, Georce Eser & Nettie MacGInirTIz 1949. Natural history of marine animals. New York, N. Y. xiit+473 pp.; 286 text figs. Pratt, Davip Mariotti 1974. Attraction to prey and stimulus to attack in the predatory gastro- pod Urosalpinx cinerea. Mar. Biol. 27 (1): 37-45; 2 figs. 3 tables McGraw-Hill Book Co., Vol. 20; No. 1 THE VELIGER Page 33 Notes on the Opisthobranch Fauna of South San Francisco Bay DAVID W. BEHRENS 416 Lilac Drive, Los Osos, California 93402 MERRITT TUEL Marine Ecological Institute, Redwood City, California 94063 (2 Text figures) THE REPORTED OCCURRENCES of opisthobranch mollusks in San Francisco Bay have previously been restricted to the northern and central portions of the estuary (APLIN, 1967; GosLINER & WILLIAMS, 1970; BEHRENS, 1971a, 1971b; HoLLEMAN, 1972; GopparD, 1973). This report deals with the opisthobranch fauna of the southern reaches of the bay (Figure 1). Included are a range ex- tension, two occurrences new to San Francisco Bay, one occurrence new to the Pacific coast, and description of an atypical color pattern. Biological samples were taken periodically in South San Francisco Bay over a period of 2$ years (February 1972 to October 1974). Collection was by means of an 18 foot (5.4m) otter trawl sampling at a depth of 3-5m from the R/V Inland Seas (operated by the education program of the Marine Ecological Institute, Redwood City). Hand collected samples were also taken off the boat floats at marinas within the Port of Redwood City, Red- wood City, California (Spring through Autumn 1974). Opisthobranchs were collected at Pete’s Harbor at the foot of Whipple Avenue and at the Port of Redwood City, end of Harbor Boulevard. DISTRIBUTION anp OCCURRENCE SACOGLOSSA 1. Elysia hedgpethi Marcus, 1961 April, July, August 1972 west of Redwood Creek, otter trawl, silty clay bot- tom; north of Dumbarton Bridge, otter trawl, silty clay bottom; September 1974 Port of Redwood City, intertidal, silty clay and rip- rap; NUDIBRANCHIA Doridacea 2. Diaulula sandiegensis (Cooper, 1862) April 1972 southwest of Bay Farm Island, otter trawl, bottom silty clay with shell material; 3. Rostanga pulchra MacFarland, 1905 April 1972 southwest of Bay Farm Island, otter trawl, bottom silty clay with shell material; Aeolidacea 4. Eubranchus misakiensis Baba, 1960 March 1974 Redwood Creek, otter trawl, silty clay bottom; October 1974 Port of Redwood City, off boat floats; 5. Hermissenda crassicornis (Eschscholtz, 1831) February - August 1972 - 1974 throughout South San Francisco Bay, otter trawl, silty clay bottom; Page 34 THE VELIGER Figure 1 Map of South San Francisco Bay showing distribution of species 1. Elysia hedgpethi 4. Eubranchus misakiensis 2. Diaulula sandiegensis 5. Hermissenda crassicornis 3. Rostanga pulchra 6. Trinchesia sp. Vol. 20; No. 1 Vol. 20; No. 1 October 1974 Pete’s Harbor, off boat floats; 6. Trinchesia sp. September - October 1974 Pete’s Harbor, off boat floats; September 1972 Palo Alto Yacht Club, off boat floats. The only report of Elysta hedgpethi within San Fran- cisco Bay was made by Gopparp (1973). Goddard col- lected 30 specimens from the Richardson Bay mudflat during the summer of 1972. On several occasions Elysia was encountered in trawl samples taken in the South Bay (see Figure 1). Trawls were at a depth of about 3m. The Elysia were always in association with the algal species of Bryopsis, Ulva or Gigartina. Four specimens were collect- ed on September 19, 1974 along the riprap shoreline of the industrial wharf at the Port of Redwood City. All specimens were in association with Ulva sp., and were collected at the minus 30cm tide level. Fresh spiral egg masses were found near each specimen. This report ac- counts for the first occurrence of this species in the South Bay. Diaulula sandiegensis was represented by a single spe- cimen only. This specimen was trawled up offshore of Bay Farm Island. HoLLeEMAN (1972) reported it from this same locality. Its occurrence in the North Bay was report- ed by Gopparp (1973). To the authors’ knowledge, Rostanga pulchra has never before been reported from within San Francisco Bay. On April 26, 1972, 2 specimens of R. pulchra were collected by trawl just southwest of Bay Farm Island. Regretfully, no additional observations or ecological comments can be added for this species at this time. BEHRENS (1971b) first reported the occurrence of Eubranchus misakiensis in the eastern Pacific. That re- port, constituting a disjunct range extension from Japan, established the presence of this species at the San Fran- cisco Municipal Marina. As noted in the original report, E. misakiensis from the boat floats occurred in association with campanularid hydroids. Egg masses were found on the hydroids. Of particular interest is the atypical color pattern shown by the Hermissenda crassicornis collected in this area. Although Biircin (1964) describes color variation in this species, none of the more than 50 specimens col- lected during this study exactly fits any of the previous descriptions. THE VELIGER Page 35 In summarizing the color pattern of this highly vari- able species, Biirgin says, “The body pattern consists of blue lines running along the middle of the body and tail, forming two rhomboid patterns, one behind the rhino- phores, a second outlining the pericardium. White or bluish lines also run along the sides of the body between the groups of cerata. They all converge on the tail. Within the first, and sometimes within the second of the rhom- boid patterns, and on the side of the head there are very conspicuous orange markings.” In the animals we have collected the longitudinal lines on the body are always white, never blue. Indeed, this lack of blue coloration is what called our attention to the apparent color difference. The white lines along the cephalic tentacles and foot corners are as described by Btrcin (1964). However, just anterior to the rhinophores, where these lines nor- mally converge and continue down the dorso-medial sur- face to the tail, the pattern deviates from the description. In the South Bay specimens the white lines become bro- ken and irregular just before convergence dorso-medially. Posteriorly they form a single broken line which disap- pears in the region of the pericardium. Further posteriorly on the dorsal surface, particularly between the ceratal groups, is an assemblage of random white specks. The white stripe reforms in normal fashion once again at the last ceratal group, and continues as such to the tip of the tail (Figure 2). No rhomboid patterns typical of Hermis- senda were observed. The only orange coloration on the body was restricted to the head region. The cerata differed in color pattern from Buircin’s (1964) description only slightly, the most obvious differ- ence being the lack of yellow, orange or blue exterior patches or bands. The cerata were seen to be transparent to yellowish-orange in ground color. They were tipped with the typical white or very light yellow cone. Below the cone the coloration deviated from the described color variations. Pigmentation in this region of the cerata con- sisted of white speckles and blotches arranged in a ver- tical line. The color and shape of the digestive diverticula extending up through the cerata conform with Biirgin’s description. Other than the described deviations in color pattern, the specimens were in full agreement with previous de- scriptions. A radular analysis was conducted to eliminate any questions of misidentification. The results of this ana- lysis confirmed the identification as Hermissenda, and most probably H. crassicornis. During September and October, 1974, a nudibranch referable to Zrinchesia was found living in the South Bay. pericardium white to light yellow transparent to yellow /\ white specks and blotches 1-5 designate groups of cerata digestive diverticulum white tail stripe Figure 2 Drawing of Hermissenda crassicornis left — body showing atypical white striping right — cerata showing color pattern Although this species does not appear to be one of the local coastal resident forms, we have not ascertained whether it is a foreign introduction or an undescribed species. Some 105 specimens were either collected or ob- served with egg masses on dense growths of the naked hydroid, Tubularia crocea (Agassiz, 1862) growing on the THE VELIGER Vol. 20; No. 1 boat floats and boat bottoms at Pete’s Harbor, Port of Redwood City, Redwood City, California. The speci- mens measured from 2 to 28mm in length and displayed a wide variation in ceratal color (from yellow to reddish- brown). Egg masses were numerous. Gary McDonald of the Moss Landing Marine Labora- tories (personal communication 27 November 1974) in- formed us that a similar eolid nudibranch was collected from the Palo Alto Yacht Club docks on 20 September 1972, by Mr. Mark Silberstein. On that occasion, 7 speci- mens measuring from 5 to 10mm in length were collected. All were found on the introduced anemone Haliplanella luciae (Verrill, 1898). ACKNOWLEDGMENTS The authors are extremely grateful for the cooperation and assistance of the staff at the Marine Ecological Institute, Redwood City, California, during this study. Thanks to Mark Kehoe for the estuarine sediment de- scriptions. Thanks are also due to Gary McDonald for his critical comments on the manuscript, and to Miss Joan E. Steinberg for her assistance in the analysis of the radulae of Hermissenda and confirmation of the identi- fication of Trinchesia. Literature Cited Apuin, J. A. 1967. _ Biological survey of San Francisco Bay 1963-1966. Calif. Dept. Fish & Game, Mar. Resources Oper. no. 67-4: 1-131 (15 June 1967) Benrens, Dav W. 1971a. The occurrence of Ancula pacifice MacFarland in San Francisco Bay. The Veliger 13 (3): 297 - 298 (1 January 1971) 1971b. Eubranchus misakiensis Baba, 1960 (Nudibranchia: Eolidacea) in San Francisco Bay. The Veliger 14 (2): 214-215 (1 Oct. ’71) Biron, ULRike F 1965. The color pattern of Hermissenda crassicornis (Eschscholtz, 1831). The Veliger 7 (4): 205-215; 9 text figs. (1 April 1965) Gopparp, Jerr 1973. | Opisthobranchs of San Francisco Bay.. The Tabulata 6 (4): 8-10 (1 October 1973) Gos.iner, TERRENCE M. & Gary C. WILLIAMS 1970. The opisthobranch mollusks of Marin County, California. The Veliger 13 (2): 175-180; 1 map (1 October 1970) Hotizeman, Joun J. 1972. Opisthobranch mollusks dredged in San Francisco Bay during the period 1966 to 1971. The Veliger 15 (1): 59-60 1 map (1 July 1972) Vol. 20; No. 1 THE VELIGER Page 37 Notes on a California Hybrid Hahotis (Gastropoda : Haliotidae) ROBERT R. TALMADGE ! (1 Plate) Eureka, California 95501 SINCE THE 1950s, hybridization between species of Hali- otis has been noted and recognized as such in California species of abalone. In 1971, OwEN, McLEaN « MEYER published a detailed paper on such hybridization. In ad- dition to the descriptions and illustrations of the hybrid specimens, they included many of the collecting locali- ties, most of which were either in southern California, U.S. A., or northern Baja California, Mexico. The north- ernmost station was Piedras Blancas in San Luis Obispo County, California (Lat. 35°40’N). Although no definite records were presented for a cross between H. rufescens Swainson, 1822, and H. k. kamtschatkana Jonas, 1842, it was stated that such a cross was possible in more northern waters, as both species occupied a similar range as far north as Cape Arago, Oregon (Lat. 43°18/30”N). Recently such a cross was found and deposited in the Talmadge Collection, Eureka, California (No. 3357). The specimen was taken in a depth between 6 and gm on a rocky wall of a surge channel at Point Cabrillo, Mendo- cino County, California (Lat. 39°21’N). The specimen measures: length, 114mm; width, 81mm; height of dorsal shell surface, 25mm; height of spire, 32mm. The general appearance of the shell is more elongate than the typical Haliotis rufescens of similar dimensions, more ele- vated than H. rufescens, but not as much as H. kamt- ' Field Associate, Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118 and Curator of Natural History, College of the Redwoods, Eureka, California 95501 schatkana (s.s.) in like-sized shells and with the sculp- ture of a relatively smooth H. kamtschatkana. In colora- tion the shell is a basic reddish, but close examination shows on the first 45mm the “Tapestry Pattern” of red with blue-green maculations is present. The open pores are badly distorted. When the shell was 80mm in length, a major injury occurred along the siphonal angle, obviously damaging not only the shell but the mantle with the shell-secreting glands as well. The 2 open pores do not match either Haliotis rufescens or H. kamischatkana (s.1.), while the closed pores are lower than normal in H. kamtschatkana, but with the shape and number of that species. The interior of the shell is as in H. rufescens. Unfortunately I did not see the soft parts, but was told they were “different” from any of the associated speci- mens. This specimen was obtained from the exposed face of a nearly vertical wall which was covered with a low growth of coralline algae and in association with Haliotis kamtschatkana (s.s.). In the deep crevices at the base of the wall were found normal H. rufescens, and amid the brown algae growing near the surface were found speci- mens of H. walallensis Stearns, 1891. ACKNOWLEDGMENTS I wish to express my appreciation to Dr. John De Martini and Mr. Donald Hethcock of Humboldt State University for making their field data available to me. Page 38 THE VELIGER Vol. 20; No. 1 Literature Cited Owen, Ropericx S. 1961. Hybridization in western American haliotids. Amer. Malac. Union Reprt. for 1961, Bull. 28: 34 (abstract) Owen, Roperick S., James Hamitton McLzan & Ricwarp J. MEYER 1971. | Hybridization in the Eastern Pacific abalone (Haliotis). Bull. Los Angeles Cty. Mus. Nat. Hist. Science 9: 1 - 37; illust. TALMADGE, RoBERT RayMOND 1955. Variation factors in the Haliotis. Min. Conch. Club So. Calif. 148; 4-6 Explanation of Figures 1 to 4 Figure 7: Left: Haliotis rufescens Swainson, 1822. Little River, Figures 2-4: Hybrid of Haliotis rufescens X H. kamtschatkana Mendocino County, California Cabrillo Point, Mendocino County, California. Specimen 3357 Middle: Hybrid of Haliotis rufescens X Haliotis kamt- Figure 2: dorsal view; Figure 3: ventral view schatkana, Cabrillo Point, Mendocino County Figure 4: lateral view Right: Haliotis kamtschatkana Jonas, 1899. Cabrillo Point, All figured specimens are in the Talmadge Collection Mendocino County, California [Tatmance] Figures 1 to 4 gure 1 1 F Figure 4 Tue Veuicrr, Vol. 20, No. 1 Ht | | Palio i | a oa} oh pod ,. a if ‘i er) \ . Zz ‘ Bil ‘ ity i K rn : i i “ Vo tome hed tye 4 Vol. 20; No. 1 THE VELIGER Page 39 The Effect of Temperature on the Distribution and Biomass of Mytilus edulis in the Alamitos Bay Area C. ROBERT FELDMETH ann MEREDYTH ALPERT Joint Science Department, Claremont Colleges, Claremont, California 91711 (z Map) INTRODUCTION THE COSMOPOLITAN BIVALVE, Mytilus edulis Linnaeus, 1758 is abundant in bays and harbors in southern Cali- fornia. The purpose of the present study is to describe the distribution of this species with respect to water tempera- ture in the Alamitos Bay area (Los Angeles County) where 4 large electrical power generating plants utilize bay water for cooling purposes and discharge the heated effluent into the lower San Gabriel River. The power plants increase sea water temperatures in the river up to 10°C above ambient, while other water quality para- meters, such as salinity, pH, and dissolved oxygen con- centration, appear not to be substantially changed. Although Mytilus edulis is abundant in both Alamitos Bay and the lower San Gabriel River, its abundance ap- pears to decrease in the warmer waters near the gener- ating plants. The following study describes to what extent high water temperature affects the presence, size, and total biomass of M. edulis communities in the Alamitos Bay area. MATERIALS anp METHODS Alamitos Bay is a small boat harbor and marina used mainly for recreational puproses. Mytzlus edulis forms a climax community on both floating boat docks and pilings throughout the Bay (Reis, 1964). Water is drawn from Alamitos Bay into electrical power generation plants for cooling purposes and then discharged into the adjacent San Gabriel River. Sampling of Mytilus edulis took place at 3 stations within Alamitos Bay and 3 additional locations on the San Gabriel River. Samples were obtained by scraping all organisms from an area of concrete wall or piling from the high water mark vertically downward to the sub- stratum using a putty knife 7.6cm in width attached to a 3m pole. The water depth at each station was 3m (at mean sea level). Each sample was placed into a large plastic bag and returned fresh to the laboratory. The number of mussels in each sample was counted, the mus- sels were measured to the nearest millimeter and the wet weight of the entire sample was determined. No attempt was made to remove barnacles from the samples, so their weight is included in the results. To determine how much of each sample was shell as opposed to animal (tissue) material, a dry weight and ash free dry weight were determined. Dry weight was determined after 24 hours at 105° C and the ash weight was taken after placing the dried sample in a furnace for 20 minutes at 600° C. The ash free dry weight was cal- culated by subtracting the ash weight from the dry weight and used as biomass of animal material for com- parative purposes. The above procedure was used because in a dense Mytilus edulis community numerous shells of dead mussels are included in a sample. To determine if any weight loss occurred due to CO, evolution of shell CaCO, during the furnace combustion, a pre-weighed sample of CaCO, was placed in the fur- nace with the mussels, and its weight change was mea- sured along with that of the sample. Water quality for the 2 habitats was also analyzed using the following methods: dissolved oxygen concentra- tion was measured using a polarographic oxygen analyzer (International Biophysics Co.), salinity was determined with an American Optical Refractometer (sea water model) as total dissolved salts in parts per thousand, and water clarity was measured with a standard Secchi disc. Page 40 Water temperature was measured with an electronic tele- thermometer (Hydrolab, Mark IV). RESULTS anp DISCUSSION Dissolved oxygen concentration ranged from 8.0 to 10.3 ppm for the warm and cool stations with no apparent correlation to temperature. As waters are discharged from the generators with force, considerable turbulence and aeration probably occur and hence the warmer San Gab- riel River has as much and often more dissolved oxygen as does Alamitos Bay. Salinity ranged from 32.8 to 34.0 %, while water clarity varied from 2.0 to 2.9m (Secchi disc), again with no consistent correlation for the 2 habi- tats. In Alamitos Bay temperature measurements were taken at the collection sites and were 14.8°C on 18 February and 15.8°C on 1 April, 1971 when the study terminated. Electrical eo Plant disch < Alamitos Bay ey ape heated effluent 36D Pacific Ocean Cool water Figure 1 Alamitos Bay - San Gabriel River study site Ambient temperature ocean water is drawn from Alamitos Bay and passed through four fossil fuel burning electrical power generating facilities and then discharged into the San Gabriel River. Three stations were located in the cooler waters of Alamitos Bay and three were located in the adjacent San Gabriel River THE VELIGER Vol. 20; No. 1 Water temperature in the San Gabriel River was con- siderably higher with a temperature gradient occurring between the generating plant outfall and the Pacific Ocean about 3.5km to the west (Figure 1). Site 1 in the San Gabriel River was located nearest to the generating plant (ca. 2km east of the ocean), with site 2 located about 1km to the west and site 3 near the river’s mouth. Temperatures taken near site 2 ranged from 22.7°C on 18 February to 25.0°C on 1 April, 1971. Undoubtedly, summer water temperatures are considerably higher than those measured in this study. The warmer waters of the San Gabriel River appear to have a definite effect on the Mytilus edulis population size and biomass (Table 1). The mean number of mussels collected in the colder Alamitos Bay waters was 321 per sample, while the mean for the San Gabriel River was 22 per sample. Shell length was not significantly different (P < 0.05) for the 2 habi- tats, but overall biomass was considerably greater in the colder habitat. The mean wet weight of Alamitos Bay samples was approximately 4 times greater than of those from the San Gabriel River, while the mean ash free weight (soft animal tissue only) was almost 10 times greater in Alamitos Bay samples (Table 1). There ap- peared to be no significant trend toward increased bio- mass of mussels in the cooler, down-stream stations. Physical and biological environmental factors other than temperature may be responsible for the difference in numbers and biomass in these 2 habitats. However, standard water quality analyses for dissolved oxygen, sal- inity, and water clarity (Secchi disc) showed virtually no differences for the 2 habitats. LANDENBERGER (1967) found that Mytilus edulis distribution was significantly limited by sea star predation at the lower end of their vertical distribution on a pier piling. However, no sea stars were observed at any of the collection sites in the San Gabriel River, yet these predators were quite numer- ous in Alamitos Bay. Cor « Fox (1942) found that a related species, Mytilus californianus Conrad, 1837 which inhabits wave-exposed intertidal areas of southern California, showed a sharp decrease in growth in temperatures above 20°C. How- ever, the mussels collected in this study were of similar size in both the warm and cool habitats, hence growth may not have been significantly affected by the warmed temperature of the San Gabriel River. Perhaps the lower biomass of mussels in the San Gab- riel River is due to the loss of both gametes and larvae by the sudden thermal shock upon transport through the power generation facilities. BARNETT (1972) reports some Vol. 20; No. 1 THE VELIGER Page 41 Table 1 Numbers, size and weights of mussels from Alamitos Bay and the San Gabriel River Total Number Mean Size Wet weight Dry weight Ash free dry weight Site of Mussels (mm) (gm) (gm) (gm) ALAMITOS BAY Station 1 543 26.2 1831.0 1184.0 2 289 32.9 2252.2 1030.7 3a 227 47.7 2310.0 817.6 130.7 3b 225 26.4 2348.6 1152.1 823.1 x = 321 33.3 2185.5 1046.1 226.9 S.D. = 150.9 17.5 238.4 166.0 136.1 S.E. = 75.45 8.8 119.2 83.0 96.2 SAN GABRIEL RIVER Station la 5 38.2 352.4 214.0 9.2 1b 8 33.2 521.0 330.0 8.3 Ic & 33.7 449.7 278.9 37 x = 7 35.0 441.3 278.9 Sai S.D. = 2.45 2.75 85.1 66.9 2.96 SS 1.42 1.59 49.2 38.7 1.71 2a 28 13.4 520.4 318.6 23.2 2b 58 25.8 616.7 350.5 40.0 2c 64 35.0 439.3 551.5 2.5 mean 50 24.73 525.5 406.87 21.9 S.D. 19.28 10.84 88.81 126.26 26.56 S.E. 11.15 6.27 51.335 73.0 15.35 3a 2 24.5 724.2 444.1 33.7 3b 3 28.3 669.4 439.6 30.6 mean 2.5 26.4 696.8 441.85 32.15 S.D. 1.96 2.69 38.75 3.18 2.2 S.E. 1.13 E55 22.4 1.84 1.27 mortality of bivalve larvae, and CARPENTER é¢ al. (1974) found mortality of copepods, upon passage through the cooling system of electrical power generation facilities. Since virtually all the water in the San Gabriel River (ex- cept during occasional heavy winter rains) passes through these power generation stations, and then flows back into the ocean there may be sufficient mortality to significantly lower the population size of Mytilus edulis in the San Gabriel River. ReisH (1964) found that larval settlement of Mytilus edulis in Alamitos Bay occurred in late winter and early spring when water temperatures would be coldest. Also, Moore & REIsH (1969) observed that mature ova of M. edulis were present in Alamitos Bay only in late fall and winter when water temperatures were 13 to 15°C. The almost immediate thermal shock of 25° C water, as ova or larvae are passed through the generation plants, may cause sufficient mortality to explain the small numbers of mussels in the San Gabriel River. ACKNOWLEDGMENTS We wish to thank Pitzer College for providing research funds to carry out this project. Page 42 THE VELIGER Literature Cited Barnett, PR. P 1972. Effects of warm water effluents from power stations on marine life. Proc. Roy. Soc. London, B 180: 497 - 509 Carpenter, E. J., B. B. Peck « S. J. ANDERSON 1974. Survival of copepods passing through a nuclear power station on northeastern Long Island Sound, U.S.A. Mar. Biol. 24: 49-55 Coz, WesLey RoswELy & DENIS L. Fox 1942. Biology of the California sea mussel (Mytilus californianus). I. Influence of temperature, food supply, sex and age on the rate of growth. Journ. Exp. Zool. 90 (1): 1-30 (5 June 1942) LaNnDENBERGER, Donatp E. 1967. A study of predation and predatory behavior in the Paeific star- fish, Pisaster. Ph. D. dissert., Univ. Calif. Santa Barbara, 164 pp. Moorz, Donatp R. & Donatp J. REISH 1969. Studies on the Mytilus edulis community in Alamitos Bay, Cali- fornia. — IV. Seasonal variation in gametes from different regions of the bay. The Veliger 11 (3): 250-255; 5 text figs.; 1 table (1 January 1969) ReisH, Donatp J. 1964a. Studies on the Mytilus edulis community in Alamitos Bay, Cali- fornia: I. Development and destruction of the community. e Veliger 6 (3): 124-131; 1 map; 4 text figs. (1 January 1964) 1964b. Studies on the Mytilus edulis community in Alamitos Bay, Cali- fornia: II. Population variations and discussion of the associated organ- isms, The Veliger 6 (4): 202 - 207; 3 text figs. (1 April 1964) Vol. 20; No. 1 Vol. 20; No. 1 THE VELIGER Page 43 Substrate Angle, Movement and Orientation of Two Sympatric Species of Limpets, Collisella digitalis and Collisella scabra LINDA S. COLLINS University of the Pacific, Pacific Marine Station, Dillon Beach, California 94929 ! (4 Text figures) INTRODUCTION HoMINc BEHAVIOR in limpets has been widely discussed in the literature (WELLS, 1917; VILLEE & Groopy, 1940; Hewatt, 1940; FRANK, 1964; GALBRAITH, 1965; Cralic, 1968; JEsSEE, 1968; Eaton, 1968; Miter, 1968; Mr- LARD, 1968; BREEN, 1971). Homing is generally defined as the consistent returning to exactly the same location with the same orientation (Eaton, Craic, MiLLer, MIuiarp, JesseEE, all op. cit.) Frank (op. cit.) makes a distinc- tion between homing and a home range, or area to which limpets return. The results of homing studies are varied. BREEN (1971) states that discrepancies in observations on homing of Collisella digitalis (Rathke, 1833) in the literature (FRANK, 1964; GALBRAITH, 1965; MILLER, 1968; Mriiarp, 1968) could be due to variations in the methods used to study homing. It is assumed in most studies that the effect of tagging (FRANK, op. cit.), painting (Mrmer, op. cit.; Mittarp, op. cit.; HEwatt, 1940; VILLEE & Groopy, 1940) or filing (WELLS, 1917) or other methods of mark- ing the experimental populations has a negligible effect on the animals. Variations in homing methods are not only of interest, but also such variables as length of ani- mal (JESSEE, 1968); time of tide (DEARNALEY, Det Mar, Parr « PopHAM, 1969); duration of the experiment; tidal level (JESSEE, op. cit.; WHITE, 1968); population density (BREEN, op. cit.) ; species; substrate type (VILLEE & Groopy, op. cit.; HEwarTrT, op. cit.) and angle of slope of substrate (Eaton, 1968). Eaton (op. cit.) reports 12 ' Present address: Department of Life and Health Sciences, Eco- logy and Evolution Section, University of Delaware, Newark, Delaware 19711 of 13 Acmaea limatula (Carpenter, 1864) not homing on a vertical surface, while 9 of 15 homed on a horizontal surface. Homing in C. digitalis has been generally studied on vertical surfaces (FRANK; GaLBraiTH; Miter, all op. cit.). VILLEE & Groopy (op. cit.) and Hewatt (op. cit.) studied homing in C. scabra (Gould, 1846) on herizontal surfaces. In this study I investigate the relationship be- tween angle of substrate and movement of C. digitalis and C’. scabra. Orientation of the 2 limpet species is also studied with respect to angle of substrate. The signifi- cance of all results is discussed with respect to allocation of space and food resources in competition between these 2 species of limpets. METHODS anp MATERIALS The relationship between the angle of slope of substrate, movement and orientation of Collisella digitalis and C. scabra was studied on 17 intertidal rock habitats in Zone 1 (Ricketts, Carvin & HepcpetH, 1968) in a locality at Dillon Beach, California, U.S.A. during October 1974. Each rock was partitioned into quadrats using a 0.25m? grid. Grid corners were marked with paint. The entire sur- face of each rock was covered with as many quadrats as would fit. The angle of each quadrat was measured to the nearest degree using a Brunton pocket transit attached to a board placed flat on each quadrat. Limpets were marked without removal from rocks since this seems to disrupt their behavior (BREEN, 1971). A variation of the marking method of Franx (1965) was used. Adhesive tape tags, 53mm in dimensions, numbered with India ink, were attached to limpets with Dekophane cement, and covered with 2 coats of glue to prevent abrasion of Page 44 numbers. Limpets were tagged by either marking all snails that occurred in a quadrat or by selecting them at random on a rock surface. The size distribution of the limpets used was identical for each species. Limpets varied from 10.0 to 17.0mm in length. The 0.25m? quadrat grid was divided into 25 10cm X1ocm squares with nylon cord. The limpet’s position in the grid was defined by a number and a letter. For each tagged limpet, the following information was obtained: position in the grid, orientation, and size. Orientation was analyzed by a variation of the method used by Miter (1968). In my experiments the orientation of a limpet on a rock surface was recorded in terms of an 8-hour clock toward which the head of the animal was pointing: Animals with their heads straight up were in the 8 o'clock position; those with the heads straight down 4 o'clock, etc. Observations in this study were made on consecutive days at day low tides when limpets were stationary. It was not possible to make observations at high tides, whether at night or during the day. It is not possible to detect those limpets which moved and came back to the same spot and those that did not move at all during the period of observation. Thus, a distinction could not be made between homing and non-homing limpets that move and limpets that do not move at all or very little. RESULTS Movement Experiment: The analysis of the data followed that of BREEN (1971). He believed that a significant deviation of observed mi- gration frequencies from expected Poisson migration fre- quencies would be indicative of homing. The fit of data to a Poisson distribution indicates that the probability of movement is random, the limpets moving independently of each other in any time interval irrespective of whether they have moved previously or not. Data for Collisella scabra (Table 1) show that the observed migration fre- quencies do not deviate significantly from an expected Poisson distribution, yet there were many limpets found in the same spot. Since there is a probability of less than 1 of movement in any time interval, it is expected that some limpets do not move at all, some move once, some twice, etc., the frequency of the number of moves being a Poisson distribution. It is expected that many limpets do not move at all if the probability of movement is low, so that in a time interval many would not be observed moving. As the probability of moving in some time inter- val increases, the number of movements in a time interval THE VELIGER Vol. 20; No. 1 Table 1 Observed migration frequencies for Collisella scabra over time in days. The total number of limpets observed. sample mean of the observed migration frequencies and chi square is given. Number of Days Observed Number of Migrations 3 4 5 6 7 0 42 39 31 23 11 1 15 9 10 3 6 2 7 4 4 3 5 3 0) 3 0 0 4 0 0) 0 5 (0) 0 6 0 Sample mean (u) = 0.45 0.33 0.56 0.31 0.73 Total number of limpets observed = 64 52 48 29 22 x2 = ms ns ns ns ns ns = not significant will be greater and the frequency of zeros (the frequency of no movements) will decrease. I use the sample mean of observed migration frequencies as indicative of the amount of movement in a population. The larger the sample mean, the more limpets will not be observed in the same position on consecutive low tides (Table 2). Table 2 Observed migration frequencies for Collisella digitalis over time in days. The total number of limpets observed, the sample mean of the observed migration frequencies and the chi square is given. Number of Days Observed Number of Migrations 3 4 5 6 7 0 34 29 20 10 2 1 36 19 11 15 9 2 17 32 16 15 33 3 11 24 8 8 4 5 6 6 5 8 6 6 2 Sample mean (uw) = 0.81 i133 1.8 Zrii 2.9 Total number of limpets observed = 87 91 76 62 36 x? = Ds 5 4 3 ns ns = not significant 3 = 0.025 > p > 0.01, significant : p < 0.005, significant | ll Vol. 20; No. 1 The sample mean is graphed against time in Figure 1 for Collisella digitalis and C. scabra. The figure shows that at any time the sample mean of C. digitalis is greater than the sample mean of C. scabra. For C. digitalis, the sample mean increases linearly with time (p < 0.001). Sample Mean I 2 3 4 5 6 7 Number of Days Figure 1 Sample means of observed migration frequency distributions plotted against the number of days observed @: Collisella digitalis O: Collisella scabra The regression equation for Collisella digitalis is Y = 0.50 X -0.71 where r = 0.99 and p < 0.001, df = 3 The regression equation for Collisella scabra is Y = 0.05 X + 0.21 where r = 0.49 ando.2p > 0.01 Page 46 THE VELIGER Vol. 20; No. 1 Table 3 Summary of total number of limpets observed, the sample mean, and chi square of observed migration frequency data over time in days for 0-30°, 35-60°, and 65-90° angles for Collisella digitalis. Number of Days Observed Angle 3 4 5 6 7 0-30° 2 1 1 — — Number of limpets observed 35-60° 44 48 40 44 25 65-90° 37 42 24 10 9 0-30° 0.5 0 0 — — Sample mean 35-60° 0.73 1.3 2.0 2.1 3.0 65-90° 0.89 1.5 2.1 2.9 3.2 0-30° ns ns ns — — ».C 35-60° ns 1 2 ns ns 65-90° ns 1 d ns ns ns = not significant 10.05 > p > 0.025 = significant 2 p < 0.005 = significant Table 4 Summary of total number of limpets observed, sample mean and chi square of observed migration frequencies over time in days for 0-30°, 35-60°, and 65-90° angles for Collisella scabra. Number of Days Observed Angle 3 4 5 6 7 0-30° 50 35 13 20 — Number of limpets observed 35-60° 10 12 12 11 3 65-90° 5 5 1 — — 0-30° 0.30 0.06 1.6 0.60 = Sample mean 35-60° 0 0 0.25 0.27 1.0 65-90° 0.40 0 0 — — 0-30° 5 6 6 ns - x? 35-60° ns ns ns ns ns 65-90° ns ns ns = — ns = not significant > 0.01 > p > 0.005, significant 2 = p < 0.005, significant than C. scabra on 35-60° and 65-g0° angles only. Orientation: Collisella scabra is more stationary than C. digitalis at : oe these angles because the chance that it moves in a time The results of the orientation data are shown in Figure interval is very low in comparison to C. digitalis. 3 and the results for o-30°, 35-60", and 65-go0° Vol. 20; No. 1 THE VELIGER Page 47 40.0 Cellisella digitalis: X* = 290.0; p < 0.005 Collisella scabra: X? = 108.3; p < 0.005 Percentage of Collisella digitalis and Collisella scabra rh TN 30 8 Sa al he “nea ae Clock Positions Figure 3 Percentage of Collisella digitalis and Collisella scabra in a particular clock position. Data are summed over 10 days of observation. The total number of Collisella digitalis observed is 801, that of Collisella scabra is 483. © = Collisella digitalis, @a = Collisella scabra angles are shown in Figure 4. A chi square was calcu- lated comparing observed limpet orientation and expect- ed orientation assuming equal chance of moving in any direction. The results show that Collisella scabra and C. digitalis do not orient evenly in any direction on rocks at low tide. In Figure 3, C. digitalis is found in downward positions 3, 4, 5, and C. scabra in position 1 and 5. This confirms Mitier’s (1968) findings for C. digitalis on vertical rocks (Figure 4). Collisella digitalis does not ap- pear to orient in this manner on 0 - 30° angles. Collisella scabra orients downward on 35 - 60° and 65 - go° angles. DISCUSSION Homing behavior in limpets is possibly an adaptation to reduce desiccation during periods of low tide (Haven, 1970; BREEN, 1972) or prevent dislodgement of the ani- mal during periods of wave splash (TEST, 1945). How- ever, there is no agreement among observers in regard to the time at which limpets move and actually home. Hewatt (1940) reported that specimens of Collisella scabra between 14.0 and 30.0mm move away from their homesites when covered by the tide. Specimens less than 14.0mm in length were observed moving during day low tides. WHITE (1968) reported moving C. scabra when submerged or amidst heavy wave action. VILLEE & Groo- Collisella digitalis: X? = 105.6; p < 0.005; N = 302 Collisella scabra: X? = 83.4; p < 0.005; N = 310 50 65 - 90° 25 Collisella digitalis: X? = 95.7; p <0.005; N = 420 Collisella scabra: X? = 70.1; p < 0.005; N = 98 iy ‘ ‘ Collisella digitalis: X? = 54.8; p < 0.005; N = 19 Collisella scabra: X? = 108.9; p < 0.005; N = 386 Clock Positions Figure 4 Percentage of Collisella digitalis and Collisella scabra in a particular clock position at 0- 30°, 35-60°, and 65-90° angles. Data are summed over 10 days of observation. The total number of limpets and chi square is given for each species. 3 = Collisella digitalis, @@ = Collisella scabra Dy (1940) observed no movements of C. digitalis and C. scabra at low tide, but small members of both species moving during day, high tides. Larger C. scabra tended to stay in one spot during low and high tides. GALBRAITH (1965) observed Lottia gigantea and C. digitalis to re- main stationary when dry and exposed. MILLER (1968) reported moving C. digitalis at day and night high tides. This contradicts BREEN (1971) who reported C. digitalis to be in their shelter sites during the day, high tides. Mm- LER (op. cit.) also observed C. digitalis to be stationary Page 48 THE VELIGER Vol. 20; No. 1 at day and night low tides. I have observed C. digitalis and C. scabra moving during night, low tides. In this study, movement and orientation of Collisella digitalis and C. scabra were correlated with angle of sub- strate. There is evidence that the amount of movement of C. scabra decreases with increase in angle and the amount of movement of C. digitalis increases with increase in angle. Collisella digitalis moves more than C. scabra on 35-60° and 65-g0° angles. Larger sample sizes are needed to prove this definitively. Collisella scabra and C. digitalis do not orient randomly on rocks at low tide. Although no differences in desiccation resistance could be detected between C. digitalis and C. scabra on 3 angles in laboratory experiments (CoLiins, 1976), desiccation resistance may play a part in accounting for differences in movement between the 2 species on various angles of substrate in the field. It is clear that other factors may also be important, such as food resources. Limpet size and abundance appear to be related to the availability of microalgae on a particular slope of substrate (MS in preparation). Differences in the amount of movement be- tween the 2 species may be related to time and distance traveled when foraging for food. STIMson (1975) found that in laboratory tanks C’. digitalis grazed for a greater percentage of time than C. scabra and that in field enclosures, C. scabra grazed less efficiently than C. digitalis, leaving more algae behind. WuiTe (1968) showed that C. scabra from -++1.8m tidal levels have lower metabolic activity and larger glycogen stores than animals from -+0.6m levels. Collisella scabra may be able to physiologically compensate for a lack of mobility which precludes frequent feeding. The movement of limpets and its relation to angle of substrate needs to be studied from the viewpoint of food resource partitioning. Previous studies (STIMSON, 1970; HaveEN, 1973; Lomnicki, 1969) have shown this to be the likely factor to pursue in more detail. ACKNOWLEDGMENTS This work formed part of a thesis presented to the faculty of the Department of Marine Sciences, Pacific Marine Station, University of the Pacific, Stockton, California in partial fulfillment of the requirements for the Degree of Master of Science. I wish to thank Dr. Steven Obreb- ski for his insights and critical advice. Literature Cited Breen, Paut A. 1971. Homing behavior and population regulation in the limpet Ac- maea (Collisella) digitalis. The Veliger 14 (2): 177 - 183; 10 tables (1 October 1971) Breen, Pau A. 1972. Seasonal migration and population regulation in the limpet Acmaea (Collisella) digitalis. The Veliger 15 (2): 133 - 141; 7 text figs; 4 tables (1 October 1972) Coxuins, Linpa S. 1976. Abundance, substrate angle, and desiccation resistance in two sympatric species of limpets. The Veliger 19 (2): 199 - 203; 2 text figs (1 October 1976) Craic, Peter CHRISTIAN 1968. The activity pattern and food habits of the limpet Acmaea pelta. The Veliger 11 (Supplement): 13-19; plt. 1; 5 text figs.; 1 table (15 July 1968) Dearna.ey, R., C. B. Der Mar, M. J. Parr & E. J. PopHam 1969. | When limpets move. Nature 221 (5179): 491 - 492 Eaton, Cartes McKENDREE 1968. The activity and food of the file limpet Acmaea limatula. The Veliger 11 (Supplement): 5-12; 7 figs. (15 July 1968) FRANK, Peter WoLFGANG 1964. On the home range of limpets. Amer. Nat. 98: 99 - 104 1965. The biodemography of an intertidal snail population. Ecolo- gy 46 (6): 831 - 844; 8 figs.; 6 tables GavsralTH, Rosert T. 1965. Homing behavior in the limpets Acmaea digitalis and Lottia gigantea. Amer. Midld. Natural. 74 (1): 245 - 246 Haven, Stoner BLACKMAN 1971. Niche differences in the intertidal limpets Acmaea seabra and Acmaea digitalis (Gastropoda) in Central California. The Veliger 13 (3): 231-248; 10 text figs. (1 January 1971) 1973, Competition for food between the intertidal gastropods Acmaea scabra and Acmaea digitalis. Ecology 54: 143 - 151 Hewatt, W. G. 1940. Observations on the homing limpet, Acmaea scabra. Amer. Midl. Nat. 24: 205 - 208 Jesszz, Wittiam FLoyp 1968. Studies of the homing behavior in the limpet Acmaea scabra (Gould, 1846). The Veliger 11 (Supplement): 52-55; 4 tables (15 July 1968) Lomnicki, ADAM 1969. Individual differences among adult members of a snail popula- tion. Nature 223: 1073 - 1074 Mitrarp, Caro, SPENCER 1968. The clustering behavior of Acmaea digitalis. 11 (Suppl.): 45-51; 4 text figs.; 1 table Miter, Atan CHARLES 1968. Orientation and movement of the limpet Acmaea digitalis on vertical rock surfaces. The Veliger 11 (Supplement): 30-44; 18 text figs.; 4 tables (15 July 1968) RICKETTS, Epwarp F, & JACK CALVIN 1968, Between Pacific Tides. i - xiii +502 pp.; 46 plts. Stanford Univ. Press, Stanford, Calif. Stimson, JoHN SzcoMBE 1970. Territorial behavior of the owl limpet, Lottia gigantea. Eco- logy 51 (1): 113-118 Stimson, Joun Secombe & Ropert BLack 1975. Field experiments on population regulation in intertidal limpets of the genus Acmaea. Oecologia 18: 111 - 120 Test, Avery RANsoME GRANT 1945. Ecology of California Acmaea. Ecology 26 (4): 395 - 405 VitieE, CLraupe ALVIN « THomas Conrap Groopy 1940. The behavior of limpets with reference to their homing instinct. Amer. Midld. Natural. 24 (1): 190-204; 25 figs.; 3 tables We tts, M. M. 1917. Behavior of limpets with regard to homing instinct. Journ. Anim. Behav. 7: 387 - 395 Wuite, T. JEFFERY 1968. Metabolic activity and glycogen stores of two distinct popula- tions of Acmaea scabra. The Veliger 11 (Supplement): 102 - 104 (15 July 1968) The Veliger (15 July 1968) Vol. 20; No. 1 THE VELIGER Page 49 Prey Preferences of Carnivorous Intertidal Snails in the Florida Keys BY RUSSELL E. INGHAM anp JAMES A. ZISCHKE Department of Biology, St. Olaf College, Northfield, Minnesota 55057 INTRODUCTION Srx SPECIES OF CARNIVOROUS GASTROPODS are commonly found in the midtidal region (yellow zone, STEPHENSON & STEPHENSON, 1950) of the intertidal rocky platform in the Florida Keys. These snails occur primarily from the southeastern United States through the Caribbean. Their feeding habits are poorly known. The prey preferences of these snails were studied as a first step toward a better understanding of the intertidal trophic relationships in this area and the effects of the predators on the distribution, zonation and diversity of their prey. The relationships between preference and availability of food and the feeding methods and activity patterns were examined for the following species of carnivorous snails: ‘Thais haemastoma floridana (Conrad, 1837), Th. deltoidea (Lamarck, 1822), Th. rustica (Lamarck, 1822), Morula nodulosa (Adams, 1845), Pisania tincta (Con- rad, 1846) and Leucozonia nassa (Gmelin, 1791). STUDY AREAS anp METHODS Observations were made on sections of limestone beach- rock parallel to the shore at sites located on Pigeon Key (24°42’N; 81°09’ W) and on the south shore of Key Vaca (24°43'N; 81°05’ W). The sections, centered at the mid- tidal level, varied in width from approximately 1.0m to 1.5m, depending upon the slope of the platform. Feeding activity was observed at low tide when the platform surface was exposed. Snails were considered to be actively feeding if they proved difficult to remove when gently rolled onto their sides and if the proboscis was seen retracting from the prey. Removing the predator prevented duplicating the same observation and allowed the predator to select another prey before the next obser- vation period. For each feeding observation the following were recorded: species and size of predator and prey; percent of snails feeding, and method of entry by snail. At each site the same strip of platform was examined during each observation period which lasted about 90 minutes and included over 100 observations of feeding. Population sizes of predators and prey were obtained from random square meter samples taken within the “yellow” zone. RESULTS Food Preferences The relationships between preference for and abun- dance of food for the 5 species of carnivorous snails at Pigeon Key are shown in Table 1. The vermetid snail, Spiroglyphus annulatus (Daudin, 1800), was the most common prey (relative abundance 99.2%) and made up 83.9% (Pisania tincta) to 99.8% (Morula nodulosa) of the diets of the predators. The other species of prey: the tree oysters Isognomon bicolor (Adams, 1845) and I. radiatus (Anton, 1839), the mussel Brachidontes exustus (Linnaeus, 1758) and the barnacle Tetraclita squamosa (Lamarck, 1818) constituted no more than 8.1% of the diet of any predator. Thais deltoidea and Th. rustica had the most varied diets, feeding on all 4 of the common spe- cies of prey. Occasional prey of the Pigeon Key carni- vores were the gastropod Astraea tecta americana (Gme- lin, 1791), Batillaria minima (Gmelin, 1791), Cerithium eburneum (Bruguiére, 1792) and Columbella mercatoria (Linnaeus, 1758). Cannibalism by Thais deltoidea was also observed. On Key Vaca the 3 species of Thais, Th. deltoidea, Th. haemastoma floridana and Th. rustica, were the only predators found (Table 2). The commonest prey of the 3 predators combined was Tetraclita (46%), followed by Isognomon (37.2%) and Brachidontes (14.7%). Spiro- glyphus, the most common prey on Pigeon Key, was not found on Key Vaca. Among the 3 thaids, Th. rustica pre- ferred barnacles (60% of its diet) and Th. haemastoma Page 50 THE VELIGER Vol. 20; No. 1 Table 1 Relationship of frequency of prey in the diets of carnivorous intertidal snails to the abundance of prey species at Pigeon Key, Florida Frequency in diet — % of total prey Prey Relative Thats Thais Morula Leucozonia Pisania Sum of all abundance (%) —_deltoidea rustica nodulosa nassa tincta predators Spiroglyphus annulatus 99.2 92.2 92.2 99.8 97.9 83.9 95.7 Isognomon bicolor and I. radiatus 0.5 5.5 6.0 0.2 0.7 8.1 2.8 Tetraclita squamosa 0.2 0.5 0.6 0 0 0 0.2 Brachidontes exustus 0.1 0.2 0.6 0 0) 0 0.16 Other = 1.6 0.6 0 1.4 8.0 1.14 % observed feeding 49.0 69.6 52.9 51.4 23.6 50.2 Total observations 784 240 946 554 263 2787 Total observed predations 384 167 500 285 62 1398 ate large numbers of mussels (30.8% of its diet). Only Th. deltoidea fed on species (Batillaria minima and Th. rustica) other than the 3 major prey. Feeding Methods and Activity Patterns Although all of the predators studied, except Leuco- zonia nassa, are functional drills, active drilling was con- firmed in only 8 of the 1704 predations observed. Entry into the most common prey, Spiroglyphus, was obtained by inserting the proboscis into the tubular shell and forcing the operculum aside. High wave action caused a pronounced reduction in feeding activity. In one case there was a 100% increase in the number of feedings observed immediately after 2 days of high winds. Feeding activity of all of the predators was higher at night. The percentages of snails of the different species feeding at midday versus midnight at Pigeon Key were as follows: Thais deltoidea 37% and 65%; Th. rustica 38% and 48% ; Leucozonia nassa 46.6% and 64.6% and Mo- rula nodulosa 50.77% and 62.4%. Pisania tincta is almost completely nocturnal; only 5 of 263 (1.9%) individuals observed at Pigeon Key were feeding during daylight hours. Specimens of Thais (7 Th. rustica, 6 Th. haemastoma, and 5 Th. deltoidea) were transplanted from Key Vaca, where no Spiroglyphus occurred, to Pigeon Key to deter- mine their reaction to unfamiliar prey. During a 7 day observation period following transplantation 15 feeding snails were observed, all on Spiroglyphus. Table 2 Relationship of frequency of prey in the diets of carnivorous intertidal snails to the abundance of prey species at Key Vaca, Florida Frequency in diet — % of total prey Prey Relative Thais Thais Thais Sum of all abundance (%) deltoidea haemastoma rustica predators Isognomon bicolor and I. radiatus 27.6 41.9 44.2 34.0 37.2 Tetraclita squamosa 71.1 35.5 25.0 60.0 46.0 Brachidontes exustus 163 16.1 30.8 6.0 14.7 Other , _ 6.5 0 (0) 21 % observed feeding 28.8 38.3 45.6 40.2 Total observations 149 227 386 762 Total predations 43 87 176 306 Vol. 20; No. 1 CONCLUSIONS Thais haemastoma has been reported to feed primarily on mussels, oysters, clams and barnacles (BUTLER, 1953, Rapwin, 1968) but there is no published quantitative information on the food preferences of Th. deltoidea or Th. rustica. Our observations show that the common thaids in the Florida Keys feed primarily on sessile ver- metid snails, when they are available, but that in the absence of vermetids they feed on barnacles, oysters and mussels in direct relation to the relative abundance of the prey species. Pacific coast relatives of Morula nodulosa and Pisania tincta are reported to feed almost exclusively on barnacles (Pang, 1966a). In the Keys these 2 species occur low in the intertidal zone and thus their preference for Spiro- glyphus as prey may be because they do not normally Move up to the levels where barnacles or bivalves are common. Leucozonia nassa (family Fasciolariidae) is not a drill but enters its prey by inserting the proboscis together with mild rasping of the radula and use of the shell margin. Paine (1966b) reports that many small fascio- larids feed on tubicolous worms and WiLicox (1895) ob- served Fasciolaria hunteria (G. Perry, 1811) eating the vermetid snail Petaloconchus nigricans (Dall, 1884). The feeding mechanism of L. nassa probably restricts its diet in the Keys primarily to vermetid snails and tubi- colous polychaetes. Preference of all of the predators for Spiroglyphus as food is most easily explained by the abundance and ease of entry to the prey. However, because of its small size (under 1omm long by 2mm wide) the consumption rate for this prey must be very high. Predators on this species, accordingly, would be expected to spend a greater amount THE VELIGER Page 51 of their time feeding. Indeed, the percent of Thais deltoi- dea and 'Th. rustica observed feeding on Pigeon Key was much higher than on Key Vaca, 49.0% versus 28.8% and 69.9% versus 45.6% respectively. Further, all of the predators on Pigeon Key, except Pisania tincta, fed both day and night. Boring activity by the predators was rarely observed although most possess a drilling mechanism. There is probably little or no drilling involved in entering bar- nacles and vermetid snails which made up a large portion of the prey. ConNELL (1961) has observed that the thaids in California are barnacle specialists and seldom drill. Only the 3 species of Thais were found where Spiro- glyphus was absent. However, the exact relationships be- tween predator and prey distribution, abundance and diversity in the Keys will require further study. Literature Cited Butter, PB A. 1953. The southern oyster drill. 67-75 ConneELt, Joszpx H. 1961. Effect of competition, predation by Thais lapillus, and other fac- tors on natural populations of the barnacle Balanus balanoides. Ecol. Monogr. 31: 61 - 104; 22 text figs. Paing, Ropert TREAT 1966a. Food web complexity and species diversity. 65 - 75 1966b. Function of labial spines, composition of diet, and size of cer- tain marine gastropods. The Veliger 9 (1): 17-24; 2 text figs. (1 July 1966) Proc. nat. Shellfish Assoc. 1953: Amer. Nat. 100: Rapwin, Grorce Epwarp 1968. Comparative radular morphology and feeding habits of muricid gastropods from the Gulf of Mexico. Bull. Mar. Sci. 18: 72 - 85 StspHenson, T. A. & ANN STEPHENSON 1950. Life between tide-marks in North America. 1. The Florida Keys. Journ. Ecol. 38: 354 - 402 Wittcox, J. 1895. The habits of Florida littoral molluscs. The Nautilus 8: 78 - 80 Page 52 THE VELIGER Vol. 20; No. 1 A New Species of Subcancilla (Gastropoda : Mitridae) from the Gulf of California R. A. WHITNEY 953 East Prairie Street, Decatur, Illinois 62521 (1 Plate) A species oF Subcancilla which has not been described previously was received from the Gulf of California. The specimens were obtained from local fishermen who had trawled them off the Sonoran coast of Mexico in 1974 near Punta Colorado. As is generally the case with speci- mens procured from fishermen, the data are quite general, although Punta Colorado is a well known landmark at the west end of Bacochibampo Bay. This bay lies north of the town of Guaymas, Sonora, Mexico. The area is noted for its fine shrimp, and much commercial trawling is done at a depth of 54 to 90m, so it may be assumed the speci- mens came from this depth. Subcancilla welkerorum Whitney, spec. nov. (Figures z to 4) Description: Size medium; biconic; color white with sculpture of raised reddish brown spiral ribs, narrower than interspaces; vague brown stains and blotches on shell; nucleus missing; first 5 postnuclear whorls with 4 spiral ribs per whorl; next 2 whorls with 4 spiral ribs per whorl; numerous small axial striae on shoulders of whorls and between spiral ribs, more prominent on posterior por- tion of whorl; sutural ramp angles at 50° ; reddish brown axial lirae on 4 anterior whorls, unevenly spaced and more prominent on body whorl, giving cancellate appear- ance; aperture approximately }$ the length of the shell with outer lip slightly crenulate; ends of spiral ridges appear emarginate inside aperture on outer edge; colu- mella straight with 3 plications, the anterior plication being quite weak; inside of aperture pale buff; fasciole with spiral ridges; anterior canal broad, slightly twisted; length 29.5mm; diameter 9.4mm; 8 whorls, nucleus truncated. Holotype: Los Angeles County Museum of Natural His- tory, Type Collection No. 1814. Type Locality: The holotype and paratypes were col- lected during 1974 off Guaymas, Sonora, Mexico. Paratypes: Paratypes are deposited in the following collections: R. A. Whitney collection, no. 48; length 32 mm; width 9.75mm; no. 49, length 28.6mm; width 9.2 mm; and the third paratype, length 25.3mm; width 8.1 mm is in the Douglas and Sherry Welker collection, no. 15, Decatur, Illinois. Explanation of Figures 1 to 4 Subcancilla welkerorum Whitney, spec. nov. Figure 1: Holotype. Los Angeles County Museum of Natural History, type no. 1814 XK 2.7 Figure 2: Paratypes; Whitney specimens on left; Welker specimen on right X 2.5 Figure 3: Enlarged view of body whorl. Paratype in Whitney Col- lection x7 Figure 4: Enlarged view of aperture. Paratype in Whitney Col- lection X7 [Witney] Figures 1 to 4 Tue Ve iarr, Vol. 20, No. 1 ‘igure 1 st Vol. 20; No. 1 THE VELIGER Page 53 ead EEEEEEEEEEEEEEEEEEEEE EE EEEREEEERIERERERRERRIRERERARIRRIEE TET n tana Other Material Examined: In addition to the type lot, we have examined one other specimen, Los Angeles Coun- ty Museum Type A. 9350.73 which was dredged near El] Tobari, Sonora, Mexico. Largest Specimen Examined: Whitney Paratype no. 48: length 32mm, width 9.75mm. DISCUSSION There is only minor variation among the specimens ex- amined. The sculpture consists of 4 reddish brown spiral ribs per whorl on a white background. The anterior spiral rib of the body whorl is almost keel-like, accentuating the overall biconic appearance of the shell. The main differ- ence is in the number of axial lirae, and the intensity of the color of the lirae. This is most prominent on the body whorl and causes some specimens to have a cancellate ap- pearance. Subcancilla welkerorum superficially resembles several other species of the genus Subcancilla. In S. eryth- rogramma (Tomlin, 1931) the shell has 2 or 3 spiral ribs per whorl and the axial markings are vague streaks and blotches, whereas in S. welkerorum the axial markings are distinct lirae and give the shell a cancellate appear- ance; S. attenuata (Broderip, 1836) lacks the brown spiral ribs; S. phorminx (Berry, 1969) has a narrower aperture, the spiral ribs are yellowish brown and the shell lacks the brown blotches and axial markings characteristic of S. welkerorum. There are 11 species of Subcancilla listed in Kren (1971) as being found in the Eastern Pacif- ic. However, SpHon (1976) has transferred S. sphoni (Shasky « Campbell, 1964) from the subgenus Strigatella to the genus Subcancilla, and he also has named a new species, S. edithrexae. Thus, with the changes noted and the naming of the new species, S. welkerorum, there are now 14 species of Subcancilla found in the Eastern Pacific. The new species is named in honor of Douglas and Sherry Welker of Decatur, Illinois, in appreciation of their encouragement of the author’s study of Mitridae. The Welkers have also accompanied the author on several shell collecting trips. ACKNOWLEDGMENT I am sincerely grateful for the generous cooperation given me over a period of many months by the individu- als who have contributed so much of their time towards the completion of this paper. I wish to express my thanks to James H. McLean of the Los Angeles County Muse- um for his advice, and to Gale Sphon of the Los Angeles County Museum for arranging a loan of museum speci- mens for comparative studies. I also wish to acknowledge with gratitude the excellent photographs of the holotype and paratypes by Bertram C. Draper, Museum Associate, Los Angeles County Mu- seum of Natural History. Literature Cited Berry, SAMUEL STILLMAN 1969. Notices of new Eastern Pacific Mollusca VIII. col. 1 (26): 159 - 166 BRODERIP, WILLIAM JOHN 1835 - 36. Characters of new genera and species of Mollusca and Con- chifera, collected by Mr. Cuming. Proc. Zool. Soc. London for 1835 [pp. 41-47 — 1 June 1835; pp. 192 - 197 — 8 April 1836] Leafl. Mala- (17 December 1969) 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. ed. 2. Stanford Univ. Press, Stanford, Calif i-xiv+ 1064 pp.; ca. 4000 figs.; 22 color pits. (1 September 1971) Suasxy, Donatp R. « G. Bruce CaMPBELL 1964. New and otherwise interesting mollusks from Guaymas, Sonora, Mexico. The Veliger 7 (2): 114-120; plts. 21, 22; 1 text fig.; 1 map (1 October 1964) SpxHon, Gaz G. 1976. The Mitridae of the Galapagos Islands. 90 (2): 63-64 Tomuin, JonNn Reap tz BrockToNn 1931. On the name Mitra lineata. The Nautilus (1 April 1976) The Nautilus 45 (2): 53-55 (14 October 1931) Page 54 THE VELIGER Vol. 20; No. 1 Feeding in Gastropod Mollusks: Behavioral and Neurophysiological Substrates - A Symposium INTRODUCTION Thursday, August 14, 1975, Whitman Auditorium Marine Biological Laboratory, Woods Hole, Mass. BY SessionI Feeding: Behavioral Analyses ALAN GELPERIN 1:20 Introduction A. GELPERIN Department of Biology, Princeton University 1:40 Navanax Bis EE : 2:00 Pleurobranchaea J. Ram Princeton, New Jersey 08540 2:20 Aplysia A, SuSSWEIN 2:40 Limax and Ariolimax D. SENSEMAN . : 3:00 Helisoma C. KanEKo D f ; ber of tigat URING THE SUMMER of 1975, a number of investigators 3.90 § ia A Gone at the Marine Biological Laboratory in Woods Hole, Massachusetts, were working on various aspects of feeding 3:45 General Discussion in gastropod mollusks. Investigators concerned with both behavioral and neurophysiological aspects of feeding in Session II Feeding: Neurophysiological Analyses the 5 most widely used opisthobranchiate and pulmonate species were present at the laboratory. In order to facili- 7:20 Introduction M. BENNETT tate comparisons between the bodies of data available for 7:40 Navanax D. Spray these various species and promote identification of the 8:00 Pleurobranchaea M. SIEGLER most insightful lines of further work, a one-day sympo- 8:20 Aplysia C. WEIss sium on feeding in gastropod mollusks was organized. To 8:40 Limax and Ariolimax D. SENSEMAN broaden the coverage somewhat, three speakers were im- 9:00 Helisoma C. KaneKo ported from New York and Princeton. The remaining 9:20 Summary M. BENNETT seven speakers were investigators for the summer at the 9:45 General Discussion Marine Biological Laboratory. The presentations and dis- cussion provided a unique opportunity to assess the rela- tive progress being made in the several experimental pro- ABBREVIATIONS grams, behavioral and neurophysiological. It was felt that a brief summary of the proceedings would be of EJP excitatory junctional potential general interest to students of molluscan behavior, evo- EPSP lution and neurobiology. IPSP excitatory postsynaptic potential inhibitory postsynaptic potential Vol. 20; No. 1 Predatory Behavior in Navanax inermis BY M. J. MURRAY Departments of Electrical Engineering and Biology Princeton University, Princeton, New Jersey 08540 Navanax inermis (Cooper, 1862) is carnivorous, and consumes chiefly other opisthobranchs (PAINE, 1963). Predation in Navanax proceeds in three main stages: (1) following the mucous trail left by the prey; (2) recog- nition of the prey upon encounter, signalled by protraction of the pharynx; and (3) engulfing the prey by a sucking action of the pharynx (PaINE, op. cit.; MURRAY, 1974; Spray, this issue, p. 59). ‘Trail following is mediated by two symmetrical sets of chemoreceptors located on either anterior fold (ALF) of the cephalic shield, and pharyngeal protraction is evoked by appropriate stimulation of similar receptors found on the tentacles (Murray, 1974). These two types of receptor, known as phalliform organs, are morpho- logically alike, consisting in unitary bundles of cilia mounted on and retractible into a basal tube. However, their physiologies presumably differ, since those on the ALFs are sensitive to trail mucus, while pharyngeal pro- traction is typically driven only by contact with the prey (Murray, op. cit.). Navanax’ trail following is imperfectly coordinated, and errors occur frequently which result in trail loss. Upon loss, the anterior parts of the body may be swept back and forth in a repeated broad pattern; this search mechanism frequently permits the predator to reestablish tracking. Nonetheless, since Navanax is likely to follow trails in the wrong direction (PAINE, 1963) a certain non-zero error rate can be shown to minimize the time between meals, and so presumably possesses adaptive value. On rare occasions, laboratory specimens will follow the trails of other Navanax, and copulate upon overtaking. This conspecific following behavior appears to proceed similarly to predatory tracking, and presumably involves the same sensory organs — but it is infrequent behavior and has not been studied in detail. Literature Cited Murray, M. J. « E. R. Lewis z 1974. Sensory control of prey-capture in Navanax inermis. The Veliger 17 (2): 156-158; 1 plt.; 1 text fig. (1 October 1974) Paine, RoBERT TREAT 1963. Food recognition and predation on opisthobranchs by Navanax inermis (Gastropoda : Opisthobranchia). The Veliger 6 (1): 1-9; pit. 1; 1 text fig. (1 July 1963) THE VELIGER Page 55 Mechanisms Underlying “‘Singleness of Action” in the Feeding Behavior of Pleurobranchaea californica (MacFarland, 1966) BY JEFFREY L. RAM anp WILLIAM J. DAVIS Thimann Laboratories, University of California, Santa Cruz, California 95064 THE SIMULTANEOUS PRESENTATION of stimuli normally eliciting different behaviors usually results in “singleness of action” (SHERRINGTON, 1948: 325); 7. e., the perform- ing of one behavior to the partial or complete exclusion of other behaviors. This is seen in reference to the feeding behavior of Pleurobranchaea californica (MacFarland, 1966). Righting behavior, oral veil withdrawal, and mating are inhibited during feeding; whereas, animals are reluctant to feed during egg laying (Davis et al., 1974a, 1974b). To investigate the interaction of egg laying and feeding, a hormone which causes egg laying in Pleurobranchaea has been localized and partially purified. Injection of pedal ganglia extracts into large (greater than 450ml) recipient Pleurobranchaea caused egg laying in 14 out of 18 recipients. Other ganglia, if dissected from animals not actually in the act of laying eggs, never caused egg laying (cerebro-pleural ganglion, 0/4 recipients; buccal plus visceral ganglia, o/2 recipients). Fractionation of the supernatant of the pedal ganglia on Sephadex G-50 yields hormone in fractions corresponding to a molecular weight of about 6000 daltons. Injection of this partially purified hormone into large recipients caused both egg laying and a greater than 100 fold increase in the thresh- old for eliciting feeding with a standard homogenate of squid mantles (N= 3). Controls (N= 3) receiving non- hormone containing fractions showed no significant change in feeding threshold. The working hypothesis, to be tested when the hormone has been further purified, is that feeding behavior is inhibited during egg laying by a direct inhibitory action of the egg-laying hormone on the neuronal circuitry underlying feeding. To test whether righting and oral veil withdrawal were inhibited during feeding by the stimulus to feeding or the behavioral act of feeding itself, the effect of a feeding stimulus (squid mantle homogenate, SMH) on Page 56 these behaviors was tested before and after satiation. Oral veil withdrawal was inhibited by SMH before satiation but was unaffected after satiation. In contrast, SMH inhibited righting behavior both before satiation, when animals exhibited feeding behavior in response to SMH, and after satiation, when most animals did not show feeding behavior. This suggests a model in which there is a direct inhibitory chemosensory pathway to the righting circuitry, whereas withdrawal circuitry is inhib- ited by output from the feeding circuitry which is present only during feeding. David Dansky provided technical assistance in the localization of the egg-laying hormone. Feeding experi- ments were done in collaboration with J. M. Pinneo, and G. J. Mpitsos. Literature Cited Davis, Witu1aM J., Gzorce J. Mritsos & J. M. PINNEO 1974a. The behavioral hierarchy of the mollusk Pleurobranchaea. I. The dominant position of the feeding behavior. Journ. Comp. Physiol. 90: 207 - 224 1974b. The behavioral hierarchy of the mollusk Pleurobranchaea. II. Hormonal suppression of feeding associated with egg-laying. Journ. Comp. Physiol. 90: 225 - 243 MacFaranp, Frank Mace 1966. Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci. 6: xvi+546 pp.; 72 pits. (8 April 1966) SHERRINGTON, Sir CHARLES 1948. The integrative action of the nervous system. 443 pp. 85 figs. Cambridge, The University Press i-xxvi+ Internal Variables Controlling Food Consumption in Aplysia BY A. SUSSWEIN Rockefeller University 1230 York Avenue, New York, N. Y. 10021 THE CONSUMMATORY PHASE of feeding in Aplysia consists of a fixed action pattern biting response whose intensity is affected in a graded manner by the intensity of the lip chemostimulus which elicits the response. Parametric features of the biting response which were found to be affected were the amplitude and latency of individual responses, and the interresponse interval of repetitive biting responses. These features are also affected in a graded manner as animals ingest a meal, so that as animals satiate the intensity of the biting response elicited by a given stimulus decreases. The effects of satiation on THE WEEIGER Vol. 20; No. 1 biting may be mimicked by injections of non-nutritive bulk into the anterior gut of animals. This indicates that the effects of satiation are due to mechano-stimuli in the gut. A second internal variable affecting the intensity of the biting response is arousal, i.e., the potentiation of a biting response due to previous exposure to food. Arousal represents an internal process which, in food de- prived animals, requires over 1 minute of constant food stimulation to build to a maximal level. When food stimuli are removed, arousal declines over a period of 30 minutes. Both the rise time and decay time of arousal are affected by satiation, so that in partially satiated ani- mals the time needed to produce arousal is increased and arousal falls off more rapidly. These data are sufficient to account for a number of features of Aplysia feeding that have been observed in field studies. Neurobehavioral Studies on Feeding in the Terrestrial Slug Ariolimax californicus BY D. M. SENSEMAN Monell Chemical Senses Center University of Pennsylvania, 3500 Market Street Philadelphia, Pennsylvania 19104 THE CONSUMMATORY FEEDING response of the terrestrial slug Ariolimax californicus Cooper has been investigated, using a combined behavioral and electrophysiological ap- proach. Behavioral experiments have focused on the fac- tors influencing food intake. It has been found that the hardness of the food determines the rate of ingestion. With increasing hardness there is a concomitant decrease in depth of radular penetration, and therefore in bite volume. Hardness was not found to have any significant effect on bite frequency or chemosensory quality. Advantage was taken of the inverse relationship be- tween hardness and consumption rate to manipulate meal size. Feeding slugs hard artificial diets significantly re- duces the amount of food ingested during a feeding bout. Under such conditions the duration of the consummatory response is a direct function of the concentration of feed- ing stimulant contained in the diet. A regression analysis of these data leads to a simple linear equation that accu- rately predicts mean meal duration, given the concentra- tion of feeding stimulant. This equation is mathematically equivalent to a decay process in which the efficacy of the Vol. 20; No. 1 chemosensory input linearily declines as a function of time. It is postulated that the decline in chemosensory efficacy is the result of sensory adaptation. Feeding slugs an artificial diet of similar chemosensory quality, but of various hardnesses, allows the effects of meal size on the consummatory response to be assessed. Decreasing food hardness results in an increase in con- sumption rate, and therefore an increase in meal size. Increasing the meal size results in a decrease in meal duration. The reduction in mean meal duration is signifi- cantly correlated with the increase in meal size. A simple linear equation derived from the regression analysis of these data allows the reduction in meal duration to be predicted given the amount of food consumed during the meal. It is postulated that bulk distention of the gut is the salient characteristic triggering the post-ingestional inhibitory input. The linear equations describing the decline in chemo- sensory efficacy and the build-up of post-ingestional feed- back can be combined into a “Feeding Equation.” This “Feeding Equation” can predict mean meal duration and mean meal size to a high degree of accuracy given the hardness and chemical composition of the food. Present electrophysiological studies are designed to elucidate the physiological substrate embodied in the “Feeding Equa- tion.” To date, 3 symmetrical populations of primary effector neurons have been identified within the paired buccal ganglia. These effector neurons have been found to receive a number of inputs from pre-synaptic inter- neurons. Of particular interest is a large pair of interneu- rons within the buccal ganglia (7. e., L 500 and R 500). These interneurons make widespread excitatory and in- hibitory connections on primary effector neurons in the buccal ganglia and the metacerebral giant cells in the cerebral ganglia. The behavioral significance of these interneurons remains unknown. Feeding in Helisoma trivolois BY C. R. S. KANEKO anp S. B. KATER Department of Zoology, University of lowa Iowa City, lowa 52242 THE BEHAVIORAL ANALYSES Of feeding in the fresh-water pulmonate Helisoma trivolvis Say, 1816, can be conveni- ently summarized using an ethological framework. Little is known about the chemical composition of sufficient sign stimuli. It is known, however, that Helisoma is omni- THE VELIGER Page 57 vorous, grazing on vegetation and carrion when available. The sign stimulus is perceived by receptors located on the anterior foot and tentacles. The ciliated tentacles channel food substances to the highly sensitive flap area located at the base of each tentacle. The response to the sign stimulus is a klinotaxis. When the concentration of the food substances is high enough anticipatory feeding commences. The behavior itself consists of 5 arbitrarily defined stages: 1. movement of the lips and mandibles which results in opening and closing the mouth; 2. move- ment of the odontophore back and forth; 3. movement of the radula on the odontophore; 4. movement of the buccal mass; and 5. activity of salivary glands. Since feeding behavior has been shown to be genetically pro- grammed (below) and to be independent of external sensory input as a source of timing or ordering, the feed- ing behavior of Helisoma fits the definition of a fixed action pattern. Preliminary experiments have used another approach to analyze the feeding behavior: behavioral genetics. The question is whether there are any naturally occurring gene differences which result in individual differences in feed- ing. The answer is yes. Various geographically isolated Strains were brought into the laboratory and inbred over 8 to 10 generations to maintain gene frequencies at initial levels but to eliminate behavioral differences due to environmental factors. Consistent differences between 4 lines tested for rasping rate lead to the conclusion that the differences are due to gene differences. Heritability was calculated to be 0.41 + 0.14. The interesting quest- ions such as whether or not differences are due to dif. ferences in the nervous system and whether the differences reflect some sort of behavioral adaptation are yet to be approached. The neurophysiological analysis of feeding in Helisoma has moved in a stepwise fashion from the effectors more centrally, that is from the muscles to the neural elements which control the muscles. Neuromuscular synapses in Helisoma are vertebrate-like. That is, there is little peri- pheral integration since the suprathreshold excitatory junction potentials give rise to overshooting action po- tentials and neither inhibitory junctional potentials nor polyneuronal innervation has been observed. Five sym- metric pairs of protraction related motor neurons (PMns) and 8 pairs of retraction related motor neurons (RMns) have been identified and classified. Since there are 19 paired muscles which comprise the buccal mass (7. é., the feeding musculature) and since each known motorneuron innervates only a single muscle, these figures represent a maximum of roughly 2 of the motor neurons involved in moving the buccal mass. Proprioceptive feedback arises from stretch-activated receptors located in the buccal Page 58 mass and feeds back positively to RMns and negatively to PMns with excitatory postsynaptic potentials (EPSPs) and inhibitory postsynaptic potentials (IPSPs). Feed- back regulates the intensity of the power stroke (7. e., retraction phase). The motoneurons can be activated to give cyclically recurring feeding-like bursts of activity from any of a group of neurons which we have termed Cyberchron neu- rons (CNs). A particular CN is inhibitory to PMns and an- other is excitatory to RMns. CNs are electrically coupled via non-rectifying, attenuating, high-frequency filtering electrical junctions. The electrical coupling accounts for the ability to initiate feeding from any CN. Our present hypothesis is that the electrical coupling between CNs is the control point of the behavior. Specifi- cally: 1. the electrical coupling mediates the positive feedback between CNs which results in characteristic explosive massive activity which in turn results in the typical antiphase bursts of activity in Mns; 2. decreases in coupling between CNs appear to mediate termination of activity in CNs and thus terminate feeding. Evidence which is consistent with but does not prove the foregoing hypothesis is of 3 types: 1. Activity of CNs controls the timing of the feeding cycle and can be modulated by arti- ficial polarization of CNs; 2. Scrutinization of activity recorded simultaneously in various CNs is consistent with the theory that positive feedback via electrical synapses is important for burst generation. 3. Burst termination is correlated with inhibition and decrease in coupling both during normal feeding and during termination of feeding as the result of the presentation of aversive stimuli. Since blockage of input to CNs from tke rest of the nervous system (via cerebrobuccal connectives) either reversibly with procaine or by section results in feeding, the CNs are apparently potentially autoactive and are normally turned off by inhibition. The mechanisms re- sponsible for the spacing of bursts in CNs are, however, not clear. Behavioral Studies of Gastropod Feeding BY ALAN GELPERIN Department of Biology, Princeton University Princeton, New Jersey 08540 THERE ARE SEVERAL TYPES of information necessary to a complete understanding of an animal’s feeding beha- vior. These include the following: THE VELIGER Vol. 20; No. 1 1) What cues and type of navigation system guide the animal to the food? 2) How do the mechanics of ingestion operate? 3) What factors determine meal length? 4) What factors determine intermeal spacing? 5) How large is the genetic component of food selection? 6) What aspects of food selection are learned? 7) What internal and external variables determine the switching probabilities between feeding and other behaviors? The various gastropods currently being asked questions such as these exhibit a diversity of lifestyles and habitats. Behavioral questions have been motivated by the desire to physiologically dissect neuronal mechanisms; hence there is little overlap in the behavioral data currently available. The questions outlined above serve mainly to highlight the range of data available for a few species as a guide to further work. One glimmering of mechanistic commonality is evi- dent in that for several of these gastropods the critical internal variable determining meal size is bulk of food eaten, as monitored by presumed gut stretch receptors. Feeding experiments with Limax maximus support this hypothesis in the following way. When individual slugs are offered 2 agar diets simultaneously for 1 hour once each day, they sometimes eat some of each diet during the hour, although more often they eat only one diet or the other. The volume of food ingested during the feed- ing session is the same whether they eat one food or some of both foods, suggesting that feeding is suppressed when the gut contains some critical volume of food. As with insects, the screening of potential foods as to their nutri- tional value is done by external chemoreceptors which have tuned over evolutionary time to those chemical stim- uli associated with nutritious prey. Once the food is ingested, no direct monitor of nutritional value is avail- able. The specificity of food selection varies from a few prey species (Aplysia, Navanax) to many dozen (Limaz). Rapid food-avoidance learning has been demonstrated for Limax (GELPERIN, 1975) and it will be very interest- ing to see if other polyphagous herbivores exhibit this form of plasticity. Literature Cited GELPERIN, ALAN 1975. Rapid food-aversion learning by a terrestrial mollusk. Science 189: 567 - 570 Vol. 20; No. 1 THE VELIGER Page 59 Feeding in Navanax inermis, Neurophysiological Aspects BY D. C. SPRAY, M. E. SPIRA anD M. V. L. BENNETT Department of Neuroscience Albert Einstein College of Medicine, Bronx, N. Y. 10461 Navanax Is CARNIVOROUS. It lacks a radula and instead of rasping away bits of food, it engulfs its prey whole by rapid expansion of its pharynx. When prey is located and touched with tentacular receptors, the pharynx is pro- tracted and the lips are exposed. Appropriate contact with lip mechanoreceptors elicits pharyngeal expansion. Peristaltic swallowing then moves the prey from pharynx to esophagus. Protractor motoneurons, identified by antidromic stim- ulation and evoked EJPs in the muscle receive excitatory input from the cerebral ganglia and lip mechanorecep- tors. Protractor sensory receptors respond to protractor stretch or compression and form excitatory synapses on the electrically coupled pharyngeal expansion motoneu- rons. The bilateral pair of largest expansion motoneu- rons each innervate the entire pharynx, while innervation by the remaining motoneurons is unilateral and partially overlapping in the antero-posterior axis so that the phar- ynx is divided into sequentially expandable regions. Phar- yngeal mechanoreceptors possess small discrete receptive fields and have somata within the buccal ganglia. These stretch receptors form either excitatory or inhibitory syn- aptic contacts with expansion motoneurons, the latter both inhibit and decrease electrical coupling between motoneurons when activated. Lip mechanoreceptors also form excitatory or inhibitory connections with expansion motoneurons. Within each group of sensory receptors there appear to be mutually excitatory interactions. Sev- eral motoneurons innervating circumferential pharyngeal muscles have been identified; their activity is reciprocally related to that of expansion motoneurons. Expansion mo- toneurons inhibit circumferential motoneurons, but the converse does not occur. Our working hypothesis is as follows: Activation of (excitatory) lip mechanoreceptors causes activity within the pool, augmented by mutual excitation, that initiates pharyngeal expansion. Stimula- tion of pharyngeal mechanoreceptors by engulfed prey initiates prolonged activity of this sensory pool that causes swallowing. Thus pools of mutually excitatory sensory neurons act as command nuclei that remain active, well after the initial stimulus has been removed. Neuronal Basis of Pleurobranchaea Feeding BY MELODY V. S. SIEGLER University of California, Santa Cruz, California 95064 FEEDING BEHAVIOR in Pleurobranchaea californica, a carnivorous marine gastropod, consists of rhythmic alter- nating eversion and withdrawal of a muscular proboscis and underlying buccal mass. Movements of the intrinsic buccal mass musculature are controlled by the bilaterally symmetrical fused buccal ganglia. The basic cyclic pat- tern underlying feeding is endogenous to the central nervous system and can be elicited by tonic stimulation to the paired stomatogastric nerves of the buccal ganglia (Davis, SIEGLER & Mpirsos, 1973: 258; SIEGLER, Mprrsos « Davis, 1974: 1173). Coordination of Motor Neurons: Antagonists: Cyclic activity in antagonistic buccal ganglion motor neurons is driven by complementary, alternating barrages of IPSP’s and EPSP’s. These opposing synaptic inputs arise from coupled but apparently largely discrete populations of interneurons, as opposing events are only rarely observed to be 1:1. Synergists: The predominant means of coup- ling between synergistic motor neurons, however, is via alternating barrages of 1:1 EPSP’s and IPSP’s, apparent- ly from common presynaptic sources. Sensory Influences: During cyclic feeding sensory input from the buccal mass is phasic. Motor activity in the buc- cal mass-buccal ganglia preparation was compared with that in the buccal ganglia, in response to identical stoma- togastric nerve stimulation. Upon deafferentation both the frequency of the feeding rhythm and the firing fre- quency of units within eversion and withdrawal bursts were significantly reduced. In addition the largest motor neurons were recruited less frequently. Burst Modulator Cells: A population of neurons with central somata in the buccal ganglia, and axons in peri- pheral roots were identified as “burst modulators.” In the isolated ganglia they show no cyclic synaptic inputs during rhythmic feeding output, and rarely fire. Intra- cellular stimulation to individual neurons can elicit or terminate eversion or withdrawal bursts, in some cases via I:I synaptic events in motor neurons. Burst modula- tor cells are likely identical with central sensory cells previously reported by SrecLer, Mprtsos & Davis, 1974: 1173" Page 60 Literature Cited Davis, Witu1AM J., Metopy V. S. Stzcter & Gzorce J. Mptisos 1973. Distributed neuronal oscillators and efference copy in the feeding system of Pleurobranchaea. Journ. Neurophysiol. 36 (2): 258 to (March 1973) Srec.Er, Metopy V. S., Gzorce J. Mritsos « Witi1AM J. Davis 1974. Motor organization and generation of rhythmic feeding output in buccal ganglion of Pleurobranchaea. Journ. Neurophysiol. 37 (6): 1173 - 1196 (November 1974) Cholinergic and Serotonergic Control of Buccal Muscle in Aplysia BY KLAUDIUSZ R. WEISS, JOSHUA COHEN AnD IRVING KUPFERMANN Departments of Physiology and Psychiatry College of Physicians and Surgeons, Columbia University New York State Psychiatric Institute New York, N. Y. 10032 WE HAVE BEEN ANALYZING feeding in Aplysia as a means of studying the neurophysiological bases of motivational controls of behavior. As a first step we investigated the motor control of the buccal mass — the muscular organ that executes the biting response. We recorded intracel- lularly from buccal motor neurons and muscle fibers. All muscles studied appeared to be polyneuronally inner- vated, receiving 2 to 5 distinct EJPs. It was possible to distinguish characteristic EJPs within identified muscles on the basis of size, and rate and extent of facilitation and post-tetanic potentiation. We have studied in detail the accessory radula closer muscle (ARC muscle). Motor neurons producing the 4 characteristic EJPs seen in the ARC muscle have been identified in the buccal ganglion. The EJPs in this muscle were chemically mediated, most likely by acetylcholine. They were increased in high (Ca**) sea water and decreased in high (Mg) sea water. Bathing the muscle in acetylcholine caused it to contract ; cholinergic blocking agents reduced both evoked contraction and EJP size. Action potentials were not ob- served during contraction of this muscle. On the basis of lesion studies we hypothesized that the cerebral ganglion may be involved in some higher order modulation of feeding behavior, and we investigated the possible role of the metacerebral cells (MCCs), a pair of giant serotonergic cells located in the cerebral gang- hia. ; THE VELIGER Vol. 20; No. 1 We found that the MCCs produce subthreshold syn- aptic potentials in buccal motor neurons. In addition, the MCCs had a peripheral action on buccal muscle. Stimu- lation of a MCC did not produce any EJPs, but resulted in a potentiation of muscle contraction elicited by a motor neuron or by acetylcholine applied to the muscle. The mechanism of MCC potentiation of muscle contrac- tion was studied by recording EJPs during potentiation. The data suggest that potentiation, at least in part, is due to a direct action of the metacerebral cell on the muscle, perhaps on its contractility. Our results suggest the hypothesis that the MCC may he a modulatory neu- ron that can operate centrally as well as peripherally. We are currently investigating whether the MCC plays a role in mediating behavioral arousal associated with the presentation of food stimuli. Some Emergent Problems in Neural Control of Molluscan Feeding, A Chairman’s Summary BY M. V. L. BENNETT Department of Neuroscience Albert Einstein College of Medicine, Bronx, N. Y. 10461 THE MOLLUSKS STUDIED to date provide a spectrum. With respect to initiation of feeding: In Helisoma the buccal ganglia initiate the neural signals for rasping movements whenever they are disconnected from the CNS. In Navanax the buccal ganglia require a positive stimulus to initiate feeding, although pathways through the cerebral ganglia probably are not required, and thus influences from higher centers may also be inhibitory. Pleurobranchaea clearly has neurons in the cerebral gan- glia capable of driving the buccal ganglia to generate the feeding rhythm, but as in Navanax sensory pathways not involving the cerebral ganglia are also adequate to initi- ate the activity. The extent of eephalization of control of feeding requires further clarification. With respect to burst formation by mutually excitatory neurons: In Heh- soma regenerative activity of the electrically coupled and mutually excitatory “cyberchrons” apparently drives feeding. An important issue is how a population of such cells fires bursts of appropriate duration and frequency. Sensory neurons in Navanax apparently also show mutu- Vol. 20; No. 1 THE VELIGER Page 61 ally excitatory interactions, and they may function as command elements when prolonged regenerative activity in the population is initiated by evoked activity in a small subgroup of the population. Burst modulator cells of Pleurobranchaea may operate in the same manner. Again, mechanisms are not clear for control of duration and in- tensity of population activity. As yet, little knowledge is available of how motoneurons operate in different pat- terns for different motor acts. The same muscle is likely to operate in different ways in different stages of feeding or rejection of food. In Navanax synaptically controlled uncoupling is suggestive of allowing synchronous and a- synchronous activity in different reflexes, but convincing demonstration of this physiological role has remained elusive. Feedback from peripheral receptors that initi- ates chains of reflexes and preprogrammed fixed action patterns are theoretical poles of a continuum along which the several species appear distributed. Modulation of be- havior, as by the metacerebral cell in Aplysia, seems to be present as a distinct mode of neural interaction. Yet although activity of this cell may not ordinarily cause feeding, in a near threshold situation it would be expected to do so. It seems likely that a precise differentiation of command, sensory, and modulator elements will not be possible and that the terms will retain usefulness only in so far as the inherent limitations are recognized. Page 62 NOTES & NEWS New Range Extensions for Chitons (Amphineura : Polyplacophora ) BY GEORGE A. HANSELMAN Museum Associate, San Diego Natural History Museum P O. Box 1390, San Diego, California 92112 THE RAPIDITY WITH WHICH developments in marine bio- logy follow extensions of the technological or commercial frontier is often gratifying. In March 1975 Mr. Roger Dick was serving aboard the Tenneco-owned vessel Prospector, which at that time was collecting samples of manganese nodules in the east- ern Pacific in an area near 15° N Lat. and 125° W Long. In the course of dredging for nodules at a depth of 4000 m (2167 fathoms) Mr. Dick was able to collect several small marine specimens - a.shrimp, some specimens of a brittle star species, several snail species, a species of ubiquitous abyssal bivalve, and a small chiton — all of which he placed in alcohol and eventually delivered to Dr. George E. Radwin, Curator of Marine Invertebrates, San Diego Natural History Museum. The chiton proved an unusual find as it appears to be a second recorded specimen of Lepidopleurus (Leptochi- ton) benthus (Haddon, 1886). The original specimen was collected by H. M.S. Challenger from 2300 fathoms (4246m) at position 35°41’N Lat. and 157°42’E Long. The present specimen is curled; the straight length would measure 7-8mm. The cross-section is well-rounded, as noted by Haddon, and the nose plate presents a rounded THE VELIGER Vol. 20; No. 1 rather than a straight profile. The sculpture consists of fine round pustules which are generally irregularly scat- tered, but which tend to form rows aligned longitudinally on the pleural areas and somewhat concentrically on the end plates; this linear formation, however, is neither strong nor consistent. Aside from an understandably shrunken and deformed girdle, the specimen is in good condition. In a similar fashion, the advent of the new paved Baja California highway has proved beneficial. Previously, a land trip to Bahia de Los Angeles (29° N Lat., 133°35/ W Long.) on the Gulf of California was an arduous trek of several days which militated against transport of heavy ~ equipment. The trip can now be made in one day in a heavily loaded standard vehicle. In May 1975, shortly after the opening of the new road, Mr. and Mrs. C. Shy and Mr. & Mrs. L. Poorman transported boats and dredg- ing equipment to Bahia de Los Angeles and dredged the Bahia for 10 days. In addition to the usual Gulf chitons, another species, Lepidopleurus (Xiphiozona) heathi Berry, 1919, was collected, representing a surprisingly large range extension. This chiton, described from off Monte- rey, California in 15 fathoms (28m), has previously been collected only from the type area and south to off Redon- do Beach, California, always in approximately 28m. That geographical range has now been significantly extended to Bahia de Los Angeles, Baja California, Mexico, in the channel between the mainland and Isla Coronado (Isla Smith) at a depth of 28m on a bottom of dead shells and small stones. All specimens taken are absolutely typical in all aspects, and in several cases bear the same touch of lavender coloration that appears on some specimens from Southern California. Literature Cited BERRY, SAMUEL STILLMAN 1919. Preliminary netices of some new West American chitons. Lor- quinia, 2 (6): 44-47, January. [Occasional Papers of the Lorquin Natural History Club of the Southwest Museum, Los Angeles, California] Happon, ALFrep C. 1886. Report of the scientific results of the voyage of H. M.S. “Chal- lenger.” XV. — Zoology, iii Polyplacophora, pp. 1-47; plts. 1-3 Vol. 20; No. 1 THE VELIGER Page 63 Glans carpentert vs. Glans subquadrata: The Rules Concerning Renamed Transient Secondary Homonyms BY EUGENE V. COAN Research Associate Department of Geology, California Academy of Sciences Golden Gate Park, San Francisco, California 94118 RENAMED SECONDARY HOMONYMS once transferred out of their condition of homonymy have always created thorny nomenclatural problems. In such cases, workers have of- ten reverted to the original name, and up to the time of the most recent changes in the International Code of Zoological Nomenclature (1972), there was little guidance on the proper course to be taken in the case of taxa so renamed prior to 1961. The recent changes in the Code, and details now becoming evident as to their application, provide more guidance. In my recent review of the northwest American Car- ditidae (Coan, 1977), I discussed a species of Glans under the name G. subquadrata (Carpenter, 1864), having already forwarded an application to the Inter- national Commission on Zoological Nomenclature to pre- serve this commonly used name as would seem to be required by the new rule on secondary homonymy [Art. 59b(i)]. The Secretary of the Commission, Dr. Richard V. Melville has written to me (7 February 1977) provid- ing information with regard to the application of this provision. The upshot of his letter is that we should use Glans carpenteri (Lamy, 1922), as the species was re- named when Lamy placed most of the Carditidae in the genus Cardita, rather than the older and better known G. subquadrata. The Code provision in question states that “A junior secondary homonym rejected before 1961 is permanently rejected and cannot be restored unless the use of the re- placement name is contrary to existing usage. In that case existing usage is to be maintained and the matter is to be referred to the Commission to designate as the valid name ... whichever name will ... best serve sta- bility and uniformity of nomenclature.” (emphasis mine) In his letter, Dr. Melville points out that the words “contrary to existing usage” are to be interpreted in a rigorous manner, and applications should generally not be submitted to the Commission when “existing usage” is partially divided between the original taxon and its re- naming. (The kind of test given for “unused senior syno- nyms” in Art. 79(b) provides guidance on the usage ques- tion.) Melville also indicates that the Commission is in- clined, in practice, to give weight to usage outside the taxonomic field in exercising its Plenary Powers in this sort of case. Such non-taxonomic usage is generally not available with our less common marine mollusks. Thus, for all practical purposes, a taxon renamed as a secondary homonym prior to 1961 is to be considered permanently renamed, even if most workers have re- verted to the earlier name when the condition of second- ary homonymy ended. Literature Cited Coan, EucEene V. 1977. Preliminary review of the northwest American Carditidae. The Veliger 19 (4): 375-402; 4 plts. (1 April 1977) STANFORD COLLECTIONS GO TO THE CALIFORNIA ACADEMY or SCIENCES The molluscan shell collection, the type specimen collec- tion, and a major part of the invertebrate fossil collection of the Department of Geology, Stanford University, have been transferred on indefinite loan to the Department of Geology, California Academy of Sciences. All enquiries, loan requests, etc., relating to the Stan- ford collections should be directed to the Academy. PETER Roppa ANNUAL MEETING or tHE AMERICAN SOCIETY oF ZOOLOGISTS anp SOCIETY or SYSTEMATIC ZOOLOGY HOSTED sy THE CANADIAN SOCIETY or ZOOLOGISTS THE ANNUAL MEETING of the American Society of Zoo- logists and the Society of Systematic Zoology hosted by the Canadian Society of Zoologists will be held Tuesday Page 64 to Friday, December 27 to 30, 1977 in Toronto, Canada at the Royal York Hotel. The following symposia are planned: Comparative Heart Physiology: Cellular Mechanisms of Control, Bio- logy of Air Breathing Fishes, Developmental and Evolu- tionary Aspects of Brain Endocrine Function, Antigen Recognition, Comparative and Developmental Aspects of Immunity and Disease; Regeneration: Neurotrophic and Morphogenetic Controls, Polyamines in Embryonic Development, Developmental Genetics of the Mexican Axolotl; Skeletal Muscle Tissue: Form and Function; Marine Larvae: Metamorphosis and Settlement; Popula- tion Studies: Are There Any General Rules?; Nectar Feeding in Birds: Models for the Ecology and Behavior of Consumers; Numerical Taxonomy, Evolutionary Me- chanisms and Processes, and Phenetic and Phylogenetic Concepts and Methods of Classification. Contributed paper sessions are planned, and the dead- line for abstracts is September 1, 1977. Accommodations at the luxurious Royal York Hotel are $26.- per evening per room with occupancy of 1 to 4 persons. Advance registration (through December 6) is $20.- regular and $10.- for graduate students. Social events include a Happy Hour in the Exhibit Hall on December 27, a Continental Breakfast in the Ex- hibit Hall, a Wine and Sherry Reception and Luncheon following the ASZ Presidential Address, and three divisi- onal socials on December 28, a Sandwich Luncheon in the Exhibit Hall and an open party on December 29. A family program is planned and will include a one-day trip to Niagara Falls on December 31. Groups of 50 or more will be provided with chartered bus transportation round trip from the Royal York and lunch at the Revolv- ing Dining Room of the Skylon Tower in Niagara Falls for a total of about $12.-. Special group air fares will be available from most major cities. Many require only 10 persons travelling on the same flight to Toronto with separate return arrange- ments. For information contact Ms. Lou Bagnara, Travel Unlimited, 3838 East Fifth Street, Tucson, AZ 85726 (telephone: 602-325-2687) in September or October. Reservations cannot be accepted after November 15. Members of the three participating Societies will re- ceive abstract forms, housing and advance registration forms, and many details of the arrangements as they be- come available. Non-members may obtain such informa- tion from: Ms. Mary Wiley, Business Manager, American Society of Zoologists, Box 2739 California Lutheran Col- lege, Thousand Oaks, CA 91360. THE VELIGER Vol. 20; No. 1 Moving? If your address is changed it will be important to notify us of the new address at least six weeks before the effective date, and not less than six weeks before our regular mailing dates. Because of a number of drastic changes in the regulations affecting second class mailing, there is now a sizeable charge to us on the returned copies as well as for our remailing to the new address. We are forced to ask our members and subscribers for reimbursement of these charges; further, because of increased costs in connection with the new mailing plate, we also must ask for reimbursement of that expense. Effective January 8, 1968 the following charges must be made: change of address — $1.- change of address and re-mailing of a returned issue — $2.75 minimum, but not more than actual costs to us. We must emphasize that these charges cover only our actual expenses and do not include compensation for the extra work involved in re-packing and re-mailing returned copies. In view of the ever increasing difficulties in the postal service, it is essential that members and subscribers not only give us prompt and early notice of address changes, but that proper arrangement for forwarding of our jour- nal be made with the local post office (at the old address). - Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. That the printed date is the actual date of pub- lication under the rules of the International Commission on Zoological Nomenclature is based on the following facts: 1) The journal is delivered to the Post Office on the first day of each quarter, ready for dispatch; 2) at least three copies are mailed either as first class items or by air mail; 3) about 20 copies are delivered in person to the mail boxes or to the offices of members ‘in the Berkeley area; 4) two copies are delivered to the re- ceiving department of the General Library of the Univer- sity of California in Berkeley. Thus our publication is available in the meaning of the Code of the ICZN. The printed publication date, therefore, may be relied upon for purposes of establishing priority of new taxa. Vol. 20; No. 1 CauiFoRNIA MaLacozooLocicaL Society, Inc. announces Backnumbers of THE VELIGER and other publications Volumes 1 through 8: out of print Volume 9: $22.- Volume 10: out of print Volumes 11 and 12: out of print Volume 13: $24.- Volume 14: $28.- Volume 15: $28.- Volume 16: $32.- Volume 17: $34.- Volume 18: $34.- Volume 19: $34.- We now have a limited number of volumes 9, 11, 13, 14 to 18 available bound in full library buckram, black with gold title. These volumes sell as follows: Volume 9 at $27.-; Volumes 11 and 13 at $29.- each; Volumes 14 and 15 at $33.-; Volume 16 at $38; Volumes 17, 18, 19 at $41.75; to this we must add a handling charge of $2.75 per volume for shipment to domestic addresses and $4.75 for shipment to any foreign address. Further, we must collect the appro- priate amount of sales tax on the price of the bound vol- umes sold to California residents. Supplements Supplement to Volume 3: $6.-* plus handling charge [Part 1: Opisthobranch Mollusks of California by Prof. Ernst Marcus; Part 2: The Anaspidea of California by Prof. R. Beeman, and The Thecosomata and Gymnosomata of the Cali- fornia Current by Prof. John A. McGowan] [The two parts are available separately at $3.- each plus a handling charge (see below). If purchased separately, each part is subject to the California State sales tax if mailed to California addresses]. Supplement to Volume 6: out of print. Supplement to Volume 7: available again; see announce- ment below. Supplement to Volume 11: $6.-* plus handling charge. [The Biology of Acmaea by Prof. D. P. Azzorr et al., ed.] Supplement to Volume 14: $6.-* plus handling charge. [The Northwest American Tellinidae by Dr. E. V. Coan] Supplement to Volume 15: $15.-* plus handling charges as follows: $1.50 for addresses in the United States of A- merica; $3.00 for all other addresses. [A systematic Revision of the Recent Cypraeid Family Ovulidae by Crawrorp NerLi Cate] Supplement to Volume 16: $8.-* plus handling charge. [The Panamic-Galapagan Epitoniidae by Mrs. Helen DuShane] THE VELIGER Page 65 Supplement to Volume 17: $3.-* plus handling charge. [Growth Rates, Depth Preference and Ecological Succes- sion of Some Sessile Marine Invertebrates in Monterey Harbor by Dr. E. C. Haderlie] Supplement to Volume 18: $10.-* plus handling charge. [The Biology of Chitons by Robin Burnett et al.]. (Our supply of this supplement is exhausted; however, copies may be available by making application to the Secretary, Hopkins Marine Station, Pacific Grove, Cali- fornia 93950.) Unless otherwise specified, the following guidelines for estimating the handling charges should be used: Mini- mum for U.S. A. $2.-. If an order is for 2 or more items, add $0.40 for each. Minimum for all foreign countries $4.-; add $0.55 for each item in addition to the first one. These charges reflect the latest increases in fees by the U. S. Postal Service. These handling charges must, however, remain subject to change without prior notice, depending on the vagaries of rate-fixing by the postal service. Items marked with * are subject to sales tax in the State of California; residents of that State please add the appropriate amount to their remittances. Prices subject to change without notice. Send orders with remittance to: Mrs. J. DeMouthe Smith, Department of Geology, Cali- fornia Academy of Sciences. Golden Gate Park, San Fran- cisco, California 94118. Please make remittance payable to C. M.S., Inc. in U.S. $, net and free of any fees to the Society. Shipments of material ordered are made once a month. We are forced to adopt this measure because of the con- tinual cut-back in personnel at the U. S. Post Office with the amount of time wasted standing in line having increased to intolerable lengths. Since it requires the same amount of time to mail 20 packages as it takes to mail one, the saving of time by our reduced mailing schedule is obvious. It becomes glaringly obvious that with the increase in postage rates and fees, the service is deteriora- ting at increasing rapidity. Although we spend much of our time complaining, one voice is not enough to bring about a change. Subscription rate to Volume 20 remains the same. We must emphasize that under no condition can we ac- cept subscription orders or membership applications for calendar year periods. If “split volumes” are required, we must charge the individual number costs. Individual issues sell at prices ranging from US$12.- to US$20.-, depending on the cost to us. Page 66 Backnumbers of the current volume will be mailed to new subscribers, as well as to those who renew late, on the first postal working day of the month following receipt of the remittance. The same policy applies to new members. Tue VELIGER is not available on exchange from the Cali- fornia Malacozoological Society, Inc. Requests for re- prints should be addressed directly to the authors con- cerned. We do not maintain stocks of reprints and also cannot undertake to forward requests for reprints to the author(s) concerned. A Glossary of A Thousand-and-One Terms Used in Conchology by Winirrep H. ARNOLD originally published as a supplement to volume 7 of the Veliger has been reprinted and is now available from The Shell Cabinet, Post Office Box 29, Falls Church, Virginia 22046, U. S. A. The cost is US$ 3.50 postpaid if remittance is sent with the order. WE ARE PLEASED to announce that an agreement has been entered into by the California Malacozoological Society, Inc. with Mr. Steven J. Long for the production and sale of microfiche reproductions of all out-of-print editions of the publications of the Society. The microfiches are available as negative films (printed matter ap- pearing white on black background), 105mm x 148mm and can be supplied immediately. The following is a list of items now ready: Volume 1: $1.50 Volume 2: $3.00 Volume 3: $3.00 Volume 4: $4.50 Volume 5: $4.50 Volume 12: $9.00 Supplement to Volume 6: $1.50; to Volume 18: $3.00 California residents please add the appropriate amount for sales tax to the prices indicated. Please, send your order, with check payable to Opistho- branch Newsletter, to Mr. Steven J. Long, P.O. Box 243, Santa Maria, CA 93454. Volume 6: $4.50 Volume 7: $6.00 Volume 8: $6.00 Volume 10: $9.00 Volume 11: $9.00 Volumes and Supplements not listed as available in microfiche form are still available in original edition from the Society at prices indicated elsewhere in the NOTES & NEWS section. THE VELIGER Vol. 20; No. 1 WE CALL THE ATTENTION orf our foreign correspondents to the fact that bank drafts or checks on banks other than American banks are subject to a collection charge and that such remittances cannot be accepted as payment in full, unless sufficient overage is provided. Depending on the American banks on which drafts are made, such charges vary from a flat fee of $1.- to a percentage of the value of the draft, going as high as 33%. Therefore we recommend either International Postal Money Orders or bank drafts on the Berkeley Branch of United California Bank in Berkeley, California. This institution has agreed to honor such drafts without charge. UNESCO coupons are N OT acceptable except as indicated elsewhere in this section. Regarding UNESCO Coupons We are unable to accept UNESCO coupons in payment, except at a charge of $4.25 (to reimburse us for the ex- penses involved in redeeming them) and at $0.95 per $1.00 face value of the coupons (the amount that we will receive in exchange for the coupons). We regret that these char- ges must be passed on to our correspondents; however, our subscription rates and other charges are so low that we are absolutely unable to absorb additional expenses. Supplements Many of our members desire to receive all supplements published by the Society. Since heretofore we have sent supplements only on separate order, some members have missed the chance of obtaining their copies through over- sight or because of absence from home. It has been sug- gested to us that we should accept “standing orders” from individuals to include all supplements published in the future. After careful consideration we have agreed to the proposal. We will accept written requests from individuals to place their names on our list to receive all future sup- plements upon publication; we will enclose our invoice at the same time. The members’ only obligation will be to pay promptly upon receipt of the invoice. Requests to be placed on this special mailing list should be sent to Mrs. J. DeMouthe Smith, Manager, C. M.S., Department of Geology, California Academy of Sciences, Golden Gate Park, San Francisco, CA(lifornia) 94118. However, until further notice, we are suspending the pub- lication of supplements until it will be reasonably certain that we will not be forced to spend many hours in tracing Vol. 20; No. 1 of lost insured or registered parcels and entering claims for indemnification. The special mailing list of members and subscribers who have entered an “including all sup- plements” will be preserved because of our innate opti- mism that sometime within our lifetime the postal services throughout the world will return to the former excellent and reliable performance. Claims for defective or missing pages must reach us within 60 days from the publication date. We will not respond to claims of missing issues made less than 30 days by domestic addressees, or less than 60 days by foreign addressees after the publication date of our journal issues. This refusal is necessary as we have received an increasing number of “claims” as much as 6 months before the claimed issue was to be published. We wish to conserve our energy and the cost of postage and stationery for more productive purposes. We are willing to accept requests for expediting our journal via AIR MAIL; however, in that case we must ask for an additional payment of US$8.00 in all cases where the Veliger goes to domestic addresses, and a depos- it of US$18.00 for all foreign addresses (including PUAS). Of course, we will carry forward as a credit toward the postage charges of the following year any amount over the actually required postage charges. At present we are charged a minimum fee of $12.50 on each order for new addressograph plates. For this rea- son we hold off on our order until 6 weeks before mailing time, the very last moment possible. If, for any reason, a member or subscriber is unable to notify us in time and also is unable to make the proper arrangement with the Post Office for forwarding our journal, we will accept a notice of change of address, accompanied by the proper fee and a typed new address on a gummed label as late as 10 days before mailing time. We regret that we are absolutely unable to accept orders for changes of address on any other basis. In view of the probable further cur- tailment in the services provided by the Postal Service, we expect that before long we may have to increase these time intervals. CALIFORNIA MALACOZOOLOGICAL SOCIETY, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the THE VELIGER Page 67 Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. Donors may designate the Fund to which their contribution is to be credited: Operating Fund (available for current production) ; Savings Fund (available only for specified purposes, such as publication of especially long and signi- ficant papers); Endowment Fund (the income from which is available. The principal is irrevocably dedicated to scientific and educational purposes). Unassigned dona- tions will be used according to greatest need. Contributions to the C. M.S., Inc. are deductible by donors as provided in section 170 of the Internal Revenue Code (for Federal income tax purposes). Bequests, lega- cies, gifts, devices are deductible for Federal estate and gift tax purposes under section 2055, 2106, and 2522 of the Code. The Treasurer of the C. M. S., Inc. will issue suitable receipts which may be used by Donors to substan- tiate their respective tax deductions. Membership open to individuals only - no institutional or society memberships. Please send for membership ap- plication forms to the Manager or the Editor. Membership renewals are due on or before April 15 each year. If renewal payments are made after April 15 but before March 15 of the following year, there will be a re-instatement fee of $1.-. Members whose dues pay- ments (including the re-instatement fee) have not been received by the latter date, will be dropped from the rolls of the Society. They may rejoin by paying a new initiation fee. The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. Endowment Fund In the face of continuous rises in the costs of printing and labor, the income from the Endowment Fund would materially aid in avoiding the need for repeated upward adjustments of the membership dues of the Society. It is the stated aim of the Society to disseminate new infor- mation in the field of malacology and conchology as widely as possible at the lowest cost possible. At a Regular Membership meeting of the Society in No- vember 1968 a policy was adopted which, it is hoped, will assist in building up the Endowment Fund of the Society. An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 000.- or more, an entire volume will be dedicated to the memory of the decedent. Page 68 New Postage Rates Although the Postal Service continues to deteriorate con- sistently, the postal rates are increased by 30 to 100%. We are, therefore, forced to increase the handling charges as well as the postage charges on the subscription. The following rates will be in effect on all new subscriptions and subscription renewals as of December 28, 1975: $1.- on subscriptions and memberships in the U. S. A.; $2.50 on memberships and subscriptions to PUAS coun- tries (Mexico, Central and South America and Spain) ; $3.50 to all other foreign countries, including Canada. We wish to stress that we are NOT INCREASING either membership dues or subscription rates, in spite of increased printing costs. But at the same time, we wish to call the attention to our Endowment Fund, the income from which enables us, in part, to keep these charges at the established levels. Contributions (tax deductible in the U.S.A.) are always welcome. It has been announced by the Postal Service that no increase in postage rates would be asked during the cur- rent year; it was not stressed sufficiently that this applies only to the so-called first class mail. Second class mail rates are “phased,” that is, they are scheduled to be in- creased each year until the rates are sufficiently high to pay the actual cost of handling that type of mail. While it is true that the rates have. been very low, it should be borne in mind that the original intent of the special low rates was to aid in the dissemination of knowledge. This philosophy has, seemingly, been abandoned. The regulations pertaining to second class mailing re- quire “pre-sorting” of the mail which involves a large amount of time, especially if the total number of pieces is too small to warrant the employment of computeriza- tion. This requirement seems justified as long as the rates for second class matter remain substantially below those for first class matter. However, our members should be aware of the fact that postal regulations rule that second class matter can not be forwarded three months after an address change, even though the addressee guarantees forwarding postage (in contrast, first class mail, at least for the time being, is forwarded for one year and that without charge!). Thus, issues mailed to the “old” address will be returned to the publisher if return postage is guaranteed at a rate that is considerably higher; we have been charged as much as $1.45 for such returned copies. There is also a charge of 25¢ for a postal notification of the new address. It must be obvious that we cannot keep absorbing such extra expenses and keep membership dues and subscription rate at the current’ low rate. We must THE VELIGER Vol. 20; No. 1 ask for the wholehearted cooperation of all concerned to help us to hold the line against increases. Also, if a copy is returned we will, as in the past, advise the member of this fact and indicate the total costs incurred for which we must seek reimbursement. If this reimbursement is not made, we cannot continue to send future issues to the delinquent member. Membership will have to be considered as terminated and can be re-instated only upon payment of all arrears. We regret that this apparently hard rule is necessary, but we wish to continue publishing the Veliger — which will not be possible if these rules are not observed. REGARDING POSTAL SERVICE Complaints regarding late arrival of our journal are in- creasing in number, steadily, continually. However, we very conscientiously dispatch our journal on the printed publication dates. What happens after deposition at the Post Office is, of course, beyond our control. From some of our members we have been able to construct a sort of probable delivery schedule. In general, within California, 8 days is usual; outside of California, the time lapse in- creases with the distance; the East Coast can consider a lapse of “only” two weeks as rapid service; 4 to 5 weeks are not uncommon. Foreign countries may count on a minimum of one month, six weeks being the more usual time requirement and over two months not rare! To Prospective Authors Postal Service seems to have deteriorated in many other countries as well as in the United States of America. Since we will absolutely not publish a paper unless the galley proofs have been corrected and returned by the authors, the slow surface mail service (a minimum of 6 weeks from European countries, 8 to 12 weeks from India and Africa) may make a delay in publication inevitable. We strongly urge that authors who have submitted papers to the Veli- ger make all necessary arrangements for expeditious read- ing of the proofs when received (we mail all proofs by air mail) and their prompt return by air mail also. Since we conscientiously reply to all letters we actually receive, and since we experience a constant loss in insured and registered mail pieces, we have come to the conclusion that if a correspondent does not receive an answer from us, this is due to the loss of either the inquiry or the reply. We have adopted the habit of repeating our inquiries if we do not receive a reply within a reasonable time, that is 6 weeks longer than fairly normal postal service might Vol. 20; No. 1 be expected to accomplish the routine work. But we can not reply if we have never received the inquiry. Because of some distressing experiences with the Postal Service in recent years, we now urge authors who wish to submit manuscripts to our journal to mail them as insured parcels, with insurance high enough to cover the complete replacement costs. Authors must be prepared to document these costs. If the replacement costs exceed $200.-, the manuscript should be sent by registered mail with additional insurance coverage (the maximum limit of insurance on parcel post is, at present, $200.-). We are unable to advise prospective authors in foreign countries and would urge them to make the necessary inquiries at their local post offices. We wish to remind prospective authors that we have announced some time ago that we will not acknowledge the receipt of a manuscript unless a self-addressed stamped envelope is enclosed (two International Postal Reply Coupons are required from addresses outside the U.S. A.). If correspondence is needed pertaining to a manu- script, we must expect prompt replies. If a manuscript is withdrawn by the author, sufficient postage for return by certified mail within the U.S.A. and by registered mail to other countries must be provided. We regret that we must insist on these conditions; however, the exorbitant in- creases in postal charges leave us no other choice. BOOKS, PERIODICALS, PAMPHLETS The Biology of the Mollusca Second Edition, by R. D. PurcHon. Volume 57 of Inter- national Series of Monographs in Pure and Applied Bio- logy; Division: Zoology. General Editor: G. A. Kerkut. Pergamon Press, Oxford, New York, Toronto, Sydney, Paris, Frankfurt, xxv+560 pp., 1977, $35.-. The publication of a second edition of Dr. Purchon’s valuable “Biology of the Mollusca” (first published in 1968 as vol. 40 of the above-mentioned series) should be an event of great importance to students of mollusks. Unfortunately, this so-called “second edition” is but a reprinting of the first edition, with insertions of a very few recent developments. A careful inspection of the chapter bibliographies reveals no more than 4 new citations, which is a small sample of the papers on THE VELIGER Page 69 mollusks that have appeared since 1966, when the author wrote his Preface to the First Edition. As the author can- didly states in the preface to this “second edition,” the new material comprises only: the time-factor and cyc- lical patterns of feeding and digestion in bivalves (inserted on pp. 233 - 235); a discussion of the distribution of snails on certain Pacific islands, based largely on a paper by Peake (1969) (see p. 344 and the graph which re- places a map in figure 114); the carnivorous habit in the Verticordiidae (Bivalvia) as elucidated by ALLEN & TurRNER (1974) and its bearing on the affinities and origin of the septibranchs; and some minor corrections. The new discussion of the Verticordiidae leads to an interesting reconsideration of the sub-class Lamellibran- chia, which is now considered to include both Verticordi- idae and Septibranchia as carnivorous specializations (see pp. 105 and 109 - 111, and altered figures 39 and 47). Since the changes entered into the second edition are insertions “pasted in,” leaving the body of the text un- changed, there have been left a number of minor incon- sistencies, particularly relating to the re-institution of the sub-class Lamellibranchia, in which Purchon now includes septibranchs (p. 111, fig. 47). In the first edition, Purchon explicitly rejected the name Lamellibranchia in favor of Polysyringia (filter-feeding bivalves), stating (pp. 101- 102) that the majority of bivalves “‘...constitute the Polysyringia (no other name is available for these bivalves —although the term Lamellibranchia is suitably descrip- tive it is undesirable since it has long been used as the name of the whole class).” But now, even though the inclusion of the septibranchs renders Lamellibranchia even less suitable, the latter term is restored as the sub- class designation (pp. 105 and 111). This could be mild- ly confusing to students. Figures 39 and 47 eliminate Septibranchia as a sub-class, but the usage remains in all other parts of the book including the scheme of classifi- cation in Table 16 of Appendix B. Likewise, the term Polysyringia continues to be used in the unchanged part of the text. The lack of any thoroughgoing revision has left certain features of the first edition that might well have been improved. ‘The meaning of the terms Gastrotriteia, Gast- ropempta, and Gastrotetartika (in figure 47 but with no explanation in text) remains unexplained, and the re- vised figure even omits reference to the source. The important discussions of the “four minor classes” and “three major classes” which might logically have come early in such a book remain as Appendices A and B. What these criticisms add up to is not a reflection on the valuable first edition of 1968, but to the fact that the publishers have restricted the “‘second edition” to a paste- in job, doubtless to avoid the expense of resetting of the Page 70 THE VELIGER work, and this has not permitted the author to do justice to a decade of advance in knowledge of mollusks. Con- sidering that the total new material could have been inserted as an Appendix, or published as a pamphlet for about 35 cents, it is regrettable to see a hardbound “second edition” for 35 dollars. Owners of the first edition need only spend an hour making notes in a library to update their old copies. Is it too much to hope that Pergamon Press will not in future dignify slightly modified reprints by calling them new “editions”? Ralph I. Smith Department of Zoology University of California Berkeley, California 94720 Pseudothecosomata, Gymnosomata and Heteropoda (Gastropoda) by S. vAN DER SPOEL. 484 pages; text figures, tables, and charts. Bohn, Scheltema & Holkema, P. O. Box 13079, Utrecht, The Netherlands. $60.00. 1976 I FOUND THIS a difficult book to review. Not so much due to its complexity but rather because van der Spoel’s views of biogeography, ecology, behavior, speciation, and of what constitutes a species and how to go about deciding are so much at variance with my own. In situations like this it is difficult to maintain the appropriate objectivity. The book is meant to be a companion volume to “Eu- thecosomata, a Group with Remarkable Developmental Stages” published in 1967, and therefore (in spite of its title) contains a section on “Additions to the Eutheco- somata,” where “newly distinguished taxa,” “concised synonymy and short diagnoses” are presented along with many illustrations of the shells of the species (?) or for- mae (?) of these animals. But the main body of the book concerns the groups in the title. It is organized into four main sections: an intro- duction with thoughts on phylogeny and speciation, tax- onomy, distribution, and a list of type material. An appen- dix includes diagrams for the “dispersal” over something called “faunal centers,” a large number of maps and some nice illustrations of the animals. Vol. 20; No. 1 Van der Spoel recognizes what he calls formae in the shell bearing species of these groups. As near as I can make out these represent shell shapes that differ from other specimens of the same species. He names the for- mae and presents descriptions and keys, in some cases, to the formae. Each form is, presumably, based on a number of specimens that tend to look more or less alike but yet different from other formae. The formae have distribu- tional “centers” and hence a biogeography. While I think he believes that formae tend to intergrade there is little mention of intergradation nor are there numerical or graphical attempts to show how this might occur in space or time. My own impression is that there is a lot of variability in shell shape within some species in both space and time. The scale(s) on which this occurs is so large that our sampling is pretty much inadequate for the pur- poses of generalizing. For example, I found the charts of distributional centers and diagrams of dispersal to be not very convincing. This problem of scale of pattern and scale of sampling seems to me to have strongly affected the biogeographic charts of species and formae as well. Van der Spoel does not really tell us the number, distribution and type of samples on which these are based. Further, he does not tell us what the blank areas of the charts mean; are the species-formae really absent, were there adequate sam- ples but no specimens, were no samples taken there? The charts of the Pacific are very different than my own and I am naturally curious as to why this might be so. I found the synonymies, lists of type specimens, and bibliography to be very thorough and I think that these will be quite helpful to many persons wishing to study these animals. The descriptions of the species are almost entirely qualitative and while helpful are not nearly as useful as the illustrations of the animals. I don’t know whether or not to recommend purchase of this book. It is very expensive, and if the reader can afford to pay for an idiosyncratic view of the taxonomy, biogeography, speciation and behavior of these animals, in order to obtain what appears to be a good list of synonyms, a large bibliography and some nice illustra- tions, is a judgement beyond my capabilities. Joun A. McGowan, Scripps Institution of Oceanography La Jolla, California THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributional, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS"; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES” will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 812” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 51/2” by 81/2”), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr. Donatp P. Assott, Professor of Biology Hopkins Marine Station of Stanford University Dr. Warren O. Appicott, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Associate Professor of Paleontology, Stan- ford University Dr. Jerry Donouue, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatr Duruam, Professor of Paleontology University of California, Berkeley, California Dr. Capet Hann, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. JoeL W. HepcpetH, Adjunct Professor Pacific Marine Station, University of the Pacific Dillon Beach, Marin County, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Victor Loosanorr, Professor of Marine Biology Pacific Marine Station of the University of the Pacific EDITOR-IN-CHIEF Dr. Rupo.r STOHLER, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Associate Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. Frank A. Prre.xa, Professor of Zoology University of California, Berkeley, California Dr. Rozert Rozertson, Pilsbry Chair of Malacology Department of Malacology Academy of Natural Sciences of Philadelphia Dr. Peter U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. JupirH TERRY SMITH, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Raupu I. Situ, Professor of Zoology University of California, Berkeley, California Dr. CHar.es R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. THompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. Jean M. Cate Rancho Santa Fe, California 77.05 el | WILLIAM H. DALP M45 // veces Fees THE VELIGER A Qvarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California VOLUME 20 OcToBER 1, 1977 NuMBER 2 ConTENTS Freeze-Etching Studies of Pulmonate Spermatozoa. & fee WE MEAXWELE Mocs fet wk ks Sete ceca harmon zt Observations on the Copulatory Behaviour of Littorina rudis es and Littorina nigrolineata Gray epee a lee we Text oe D.G. RaFFAELLI . . Sen rane 1} The Type Specimens of Ammonites yen Gabb and Melchiorites ie Anderson (Cretaceous : Ammonoidea). : a Micuaet A. Murpuy s Peter U. Roppa . . . mF ae WES AOS | Chiton Fauna of the Galapagos Islands. (4 Pate) Attyn G. Smit & ANTONIO J. FERRERA . . SANCIE Pen abeny ot) Wemng OS Notes on Sea Hares of South Texas (Gastropoda : Opistobranchin. Nep E. StrENTH & JamMEsS E. BLANKENSHIP .. . 5 Se ade RICE 7) Reevaluation and New eee of the Genus Bittium n (Cais). os Plate) Ricuarp S. Housprick . . . Sven iu USUIOR The Chromodoridinae Nudibranchs from the Pacific Coast of America. — Part I. Investigative Methods and rie peer oe Text la Hans BertscH .. . ‘ : . 107 The Family Columbellidae in the Western Atlantic. Part IIa. -— The baeuee (4. Plates) GeorcE E. RaDbwIN. . . . Soi tee wae ncn RO A New Species of Humboldtiana Papinaiitisetyetdbe) from Coahuila, Mexico. (1 Plate; 1 Text figure) RicHarp W. FuLLINcToN « Eart G. ZIMMERMAN . . . . 134 Activity of the Gastropod Mollusk Olivella biplicata in Rete to a Nie Light/Dark Cycle. (3 Text figures) TD AVIN RV OR HICEIPS 2 neces imei eter (Osea. oh ors aie tie 8) PS) es ee ie, QD [Continued on Inside Front Cover] Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SUPERFAMILY, Famity, Subfamily, Genus, (Subgenus) New Taxa Secend Clam Postage Paid at Berkaley, California ConTENTS — Continued Size and Age-Specific Predation by Lunatia heros (Say, 1822) on the Surf Clam © Spisula solidissima (Dillwyn, 1817) off Western Long Island, New York. (5 Text figures) Davw R. Franz A New Species of Lee Tey from the Great Barrier Reef, Australia. (2 Plates) Kart H. KiEEMANN : aed, The Effects of Season on Visual and Se ea Assessment of Subtidal Geoduck Clam (Panope generosa se cimieas (1 Plate) Lynn Goopwin . 3 aie Ask batting Cypraea: A List of the Species. III. « 1 Text ro) Jerry DonoHUE Subtidal Abalone Populations in an Area Inhabited = Sea Otters. (2 Text figures) Joun Cooper, Marx WiELAnpD « ANSON HINES Observations on Feeding, Chemoreception, and Toxins in Two ged of Epitonium. (1 Plate; 1 Text ae Sicri Sao aes The Development of Conspecific Interactions in Juvenile Aplysia dactylomela Rang, 1828: An Observational aa (3 Text figures) I. Izya LEDERHENDLER PY GINO Thon Ae UE NOTES & NEWS Direct Development in the Intertidal Gastropod Batillaria zonalis (Bruguiére, 1792). (1 Text figure) SytviA BEHRENS YAMADA & CHANDRA S. SANKURATHRI A New Sea-Floor Oasis. A. Myra KEEN A Color Variant of Conus bulbus Reeve. Joun K. Tucker The Identity of Conus fulmineus Gmelin. _(1 Text figure) Joun K. TUCKER INFORMATION DESK eGR pheccdne toda Wea Seelih s,SAN a aolaca ha Gece i What’s the Difference? ‘Telegraphic Style versus Normal Style. A. Myra KEEN BOOKS, PERIODICALS & PAMPHLETS - 144 . 151 . 168 - 173 - 179 . 187 . 187 Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 20: $25.- plus postage ($1.- in U.S.A.; $2.50 to RU.A.S.; $3.50 to all other foreign Countries, including Canada). Single copies this issue $16.-. Postage extra. Send subscription orders to California Malacozoological Society, Inc. 1584 Milvia Street, Berkeley, CA 94709, U.S.A. Address all other correspondence to Dr. R. StoHLER, Editor, Department of Zoology University of California, Berkeley, CA 94720 Vol. 20; No. 2 THE VELIGER Page 71 Freeze-Etching Studies of Pulmonate Spermatozoa W. L. MAXWELL Anatomy Department, University of the West Indies, Mona, Kingston 7, Jamaica (4 Plates) INTRODUCTION THE LITERATURE DESCRIBING the ultrastructure of pulmo- nate molluscan spermatozoa has been reviewed by THompson (1973), BacceTTi & AFZELIUS (1976). All of these descriptions have, however, been limited to ex- amination of thin sections necessitating lengthy examina- tion to appreciate the complex helical forms of these gametes. Despite THompson’s (op. cit.) demonstration of the usefulness of the freeze-etching technique as a means of rapid appreciation of complex three-dimension- al forms in molluscan spermatozoa there has been no ex- haustive freeze-etching study. Freeze-etching has the ad- vantage that replicas may be produced of membrane sur- faces and three-dimensional structures over considerable areas. The most rapid appreciation of the whole cell form may be achieved with the scanning electron microscope MaxwELL (1975) but this technique lacks the resolution to study internal subcellular components. Freeze-etching allows greater resolution, and fractures reveal the three- dimensional organisation of subcellular components. The technique was applied to studies of mature spermatozoa taken from the vas deferens of a basommatophoran and a stylommatophoran pulmonate. METHODS Mature spermatozoa of Arion hortensis Férussac, 1819 and Lymnea peregra (Miller, 1774) were treated in the Balzers 360M freeze-etching unit (THoMPSoN, 1971) and the replicas were examined with an AEI 6B or 801 electron microscope. RESULTS Lymnea peregra. The mature spermatozoon has already been examined with the SEM (MaxweE Lt, 1975) and differs little from the generalized basommatophoran form described by THOMPSON (1973). The head is short and consists of 7 spiral ridges arranged one behind another. Freeze-etching provides much more detailed information about the inter- relation of these ridges which prove not to lie parallel to one another but are compressed at the base of the head in the region surrounding the implantation fossa (Figure 1). Here the width of the helical ridges is reduced from 0.25 wm to 0.16 um. The helices are discrete and the nuc- lear surface is smooth except at the anterior tip in the region beneath the digitiform, granular acrosome where there are a large number of small but discrete spiral ridges 56nm in width (Figure 2). These ridges are overlaid by the apical acrosome within which little substructure is visible. The nuclear chromatin forms a basal collar around the shallow implantation fossa which contains the anterior tip of the axoneme of the sperm tail. In regions where the plasmalemma is stripped away and the axoneme has been fractured a fine sculpturing is visible on the base of the nucleus (Figure 3). This sculpturing consists of a series of parallel ridges of variable number (12 - 18) arranged in a more or less radial manner and showing discrete points of dislocation such that adjacent groups of ridges lie at an obtuse angle to their neighbours. It is unclear whether the sculpturing is a reflection of nuclear packing or an impression left on the base of the nucleus by the closely apposed top of the mitochondrial derivative which was removed in the preparation of the replica. Page 72 THE VELIGER Vol. 20; No. 2 There are 7 helices in the anterior portion of the mito- chondrial derivative with a sinistral pitch of 4.5 um when viewed from the head posteriad (MaxwELL, 1975). When the plasmalemma is stripped away the helical organisa- tion of the paracrystalline mitochondrial derivative is revealed. While thin sections demonstrate a complex structure (ANDRE, 1962), they give no indication of the helical distribution of subunits. Freeze-etched replicas have now demonstrated that the gnm globular subunits of the paracrystalline material are arranged with a con- sistent transverse translocation such that they lie in heli- cally oriented parallel rows running around the flagellum (Figure 2). In the anterior portion of the sperm tail this orientation parallels that of the 7 underlying mitochond- rial helices but in the posterior region where only 2 heli- ces are present there is a difference in helical pitch be- tween the 2 helices and the overlying paracrystalline material. The 2 helices have a pitch of 3 um (MaAxweELL, 1975) and it is apparent (Figure 4) that the paracrystal- line material has a longer pitch. It may be suggested that the paracrystalline material has a constant helical sub- structure throughout the sperm length and that the pitch of the mitochondrial helices is independent of this. Also visible in surface fractures are series of more or less regular swellings between the 2 mitochondrial helices of the posterior portion of the sperm tail. In fractures where the outer paracrystalline material is also removed, these swellings can be seen to be due to series of regularly ar- Explanation of Figures 7 to 3 Lymnea peregra Figure 1: Longitudinal view of a mature sperm head. The granular acrosome covers the apex and the helices become squashed towards the base. X 120000 Figure 2: The 56nm spiral ridges beneath the acrosome are ex- posed, A small part of the acrosome is still attached at the right side. X 95 000 Figure 3: A surface view of the base of the nucleus. Remnants of the axoneme are visible in the implantation fossa, groups of parallel lines are visible around its periphery. There is 2 marked nuclear collar around the implantation fossa. X 85000 ranged blebs located within the derivative of the mito- chondrial cristae lying between the helices. The blebs are disposed in a helical manner around the flagellum and are about 0.05 um in diameter, spaced at intervals of 0.1 um (Figure 5). The mitochondrial helices contain glycogen (ANDERSON & PERSONNE, 1970) and freeze-etching de- monstrates a granular form with particles 20 - 30nm in diameter. When both mitochondrial paracrystalline layers are removed during the fracture of the frozen specimen the fibres of the axoneme may be exposed directly (Figure 6). Closely apposed longitudinal structures with very little surface ornamentation are displayed, but no doublet structure or regularly arranged projections are visible. Thin sections indicate that the axonemal doublets are spaced at 17.5 - 20nm with the dynein arms lying between them (WaRNER, 1974). A portion of the outer mito- chondrial membrane separates the paracrystalline mate- rial and the axoneme in pulmonate spermatozoa and the replicas may thus be interpreted as images of the surface of the outer mitochondrial membrane overlying the axo- nemal components. The separation of the doublets and the regularly ar- ranged projections of the dynein arms are visible in parts of the sperm tail lacking any mitochondrial derivative, Explanation of Figures 4 to 6 Lymnea peregra Figure 4: A surface view of the paracrystalline material with small patches of plasmalemma still attached. The difference in pitch of the helical organization of this material and the 2 major helices is displayed. X 27000 Figure 5: Longitudinal fractures of 2 sperm tails illustrating the internal mitochondrial structure. The 2 glycogen filled major heli- ces and the regular arrangement between the 2 mitochondrial membranes are displayed. In the right hand specimen the tip of part of the axoneme is exposed. X 26000 Figure 6: When all the paracrystalline mitochondrial derivative has been removed, the parallel disposition of the axoneme fibres lying beneath part of the outer mitochondrial membrane is de- monstrated, The fibres are discrete but little substructure can be determined, X 75 000 [MaxwELtL] Figures 1 to 3 Tue VELIGER, Vol. 20, No. 2 [MaxwELL] Figures 4 to 6 THE VELIGER, Vol. 20, No. 2 A *ft his a be Se MA OY $a" ,) ra.) y Wraith Mase Meme Dag oes Pt tectind arth chirp cena Ke civ as ES don ~— 1pPhen Lite tye Re : TEE lose SAA Rass ~ RAV SSS EM Ss Vol. 20; No. 2 THE VELIGER Page 73 as occurs in the most posterior portion of the Lymnea peregra sperm tail where only a glycogen deposit sur- rounds the axoneme forming a tube 0.5 um in diameter (Figure 7). Arion hortensis Freeze-etching confirms the trihelical form of the ma- ture spermatozoon (MaxwELL, 1975) but allows a greater insight into the spatial inter-relationships between the spermatozoan organelles. The 3 helices of the head run parallel, differing from the posteriad compression de- scribed in Lymnea. The helices are 0.35-0.4um wide and spaced at intervals of 0.1-0.15 mm, providing a much coarser structure. It should also be noted that the helices are dextral in orientation when viewed from the front of the head. This is clearly demonstrated in replicas of the mitochondrial region also. There is no surface sculpturing on the Arion sperm head, either beneath the acrosome or in the region of the implantation fossa (Figure 8). However, there is a discrete collar of material surrounding the axoneme in this latter region formed by a thin membranous structure around the implantation fos- sa (Figure 9) but it is unclear whether this is a remnant of the nuclear membrane. The implantation fossa is circular in section. Transverse fractures of the flagellum demonstrate that the axoneme is closely enveloped by the paracrystalline mitochondrial derivative indicating that the separation of the 2 structures often observed in thin section may, to some extent, be an artefact. In the anterior portion of the flagellum the paracrystalline derivative forms a regular circular periaxonemal sleeve in which 8 - 9 layers of ma- terial may be counted (Figure zo). Three structures are attached at the periphery of the paracrystalline sleeve, one large ‘major’ helix and 2 ‘minor’ helices externally limited by a double layer of paracrystalline material. Transverse fractures indicate a very regular substructure within the paracrystalline derivative which is masked in thin sections where several layers of subunits occur within the thickness of one section. SEM observations show that 724m behind the sperm head the trihelical organisation (Figure 11) is lost and replaced by a single, large helix. This is confirmed in freeze-etched replicas which emphasize the extent of the glycogen deposit in this region where it lies around the axoneme to cover half of its transverse circumference (Figure 13). The prominent major helix of the anterior part of the flagellum increases in size and becomes flat- tened around the axoneme to impose an ellipsoid cross- section. Freeze-etching clearly confirms the helical dis- position of this glycogen deposit when examined in a longitudinal fracture (Figure 12). Both the glycogen deposit and the axoneme are enclosed by portions of the outer mitochondrial membrane resulting in a smooth contour. Two layers of paracrystalline material may be observed, the thinner surrounding the axoneme, the other limiting the whole spermatozoon and enveloping the glycogen deposit. As in Lymnea, the outer mitochondrial membrane is closely apposed to the periphery of the axoneme (Figure 15). The 20nm spacing of the doublets is very clear but the precise doublet nature and location of the dynein arms are masked. DISCUSSION Application of the freeze-etching technique provides in- formation additional to that obtained with the SEM or thin sectioned studies of pulmonate spermatozoa. The form of the head, and any interspecific differences, can be appreciated in one favourable fracture rather than by a laborious reconstruction using thin sections. Freeze-etch- ing also allows the detection of small surface structures or sculpturing which are undetectable using the SEM or thin sections. In this study it provides evidence of a speci- alised region at the base of the acrosome in Lymnea peregra. New information is also obtained about the helical disposition of the mitochondrial derivative and its con- tents. This report includes the first demonstration of a Page 74 THE VELIGER Vol. 20; No. 2 helical arrangement of the subunits of the paracrystalline mitochondrial derivative, and indicates that the helical disposition of these subunits and the mitochondrial helices of Anpré (1962) may not be directly related. Also re- ported is the regular and close packing of the paracrystal- line subunits when observed in transverse fracture and which is masked in thin sections due to the fact that a number of layers of subunits will occur in the thickness of a single section. A previously unreported organisation of the derivatives of the cristae of spermatid mitochondria is demonstrated by freeze-etching where thin section stu- dies had indicated a low order of organisation. In the posterior part of the Lymnea spermatozoon the deriva- tives of the cristae form a double layered structure with regularly spaced concretions of material placed between them and arranged in a helical manner around the flagel- lum. No such regular organisation has been described pre- viously although the pustulose structure of the Helix pomatia spermatozoon (THompson, 1973) might be comparable. Explanation of Figures 7 to 17 Lymnea peregra Figure 7: Part of 2 glycogen filled end-pieces demonstrating the granular deposit around the axoneme exposed in the centre of the tube. Regular projections can be observed on the exposed axonemal fibres. X 80000 Arion hortensis Figure 8: A mature sperm head illustrating the parallel orienta- tion of the 3 helices and the lack of nuclear surface structure. A small membrane remnant is present at the base of the head, and the periaxonemal collar is visible. The axoneme extends backwards inside the paracrystalline sheath. X 30000 Figure 9: The base of the sperm head showing the central location of the implantation fossa containing a remnant of the axoneme and the smooth surface of the nuclear base. X 70000 Figure ro: A transverse fracture of the anterior part of a sperm tail showing the regular organisation of the paracrystalline material and the peripheral location of the 3 mitochondrial helices. X 85 000 Figure 11: External surface of a mature spermatozoon showing the trihelical structure beneath the plasmamembrane. X 48000 ACKNOWLEDGMENTS I am grateful to Professor H. E. Hinton, F R. S., Zoology Department, University of Bristol, England, and Dr H. Waters, Ministry of Overseas Development Fatal Leaf Yellowing Research Project, University of the West In- dies, Jamaica, for allowing me to use their electron micro- scopes. I am grateful to the University of the West Indies for study leave to allow me to complete this project. Literature Cited ANDERSON, WINSTON A. & PAUL PERSONNE 1970. The localisation of glycogen in the spermatozoa of various in- vertebrate and vertebrate species. Journ. Cell Biol. 44: 29-51 ANDRE, JEAN 1962. Contribution 4 la connaissance du chondriome — Etude de ses modifications ultrastructurales pendant la spermatogenése. Journ. Ultrastruct. Res. Suppl. 3: 1 - 185 Baccetti, Baccio & Bjorn AFZELIUS 1976. The biology of the sperm cell. Monographs in developmental biology 10: 1-254; 19 figs.; 26 plts.; 1 table. Basel, S. Karger Maxwe Lt, Wixuiam L. 1975. Scanning electron microscope studies of pulmonate spermatozoa. The Veliger 18 (1): 31-33; 3 plts. (1 July 1975) THoMPsON, THOMAS EVERETT 1971. Application to molluscan ultrastructure research of the Balzers 360 M freeze-etching plant. The Veliger 13 (4): 367 - 368; 1 plt. (1 April 1971) 1973. | Euthyneuran and other molluscan spermatozoa. Malacologia 14: 167 - 206; 16 plts; 11 figs.; 2 tables Warner, FRED 1974. The fine structure of the ciliary and flagellar axoneme. in: Michael A. Sleigh, ed. Cilia and Flagella. Acad. Press London & New York, i- xi+500 pp.; illust. Explanation of Figures 72 to 74 Arion hortensis Figure 12: The content of the single helix in the posterior part of the sperm tail is exposed illustrating the axoneme, the 2 layers of paracrystalline material and the spiral glycogen deposit. | 55000 Figure 13: A transverse fracture of the lower part of a sperm tail showing the ellipsoid transverse form and the extent to which the glycogen deposit surrounds the axoneme. X 100000 Figure 14: A longitudinal fracture to expose the axoneme fibres enveloped by part of the outer mitochondrial membrane. The fibres and their separation are discrete. The paracrystalline material is very closely applied to the axoneme surface. ® 80 000 [MaxwELL] Figures 7 to 11 Tue VELIGER, Vol. 20, No. 2 ate SAR eee ete RR ate “ F Y a: on € - ‘ ) ‘ x i 4 ( te ri Cah f a han a 4 t “| We " fom? a aN , f ‘ p 1 n F i r 2 sz i i i ‘ 4 a . \ ; 5 l = He y i} - | i i ‘ ane i t | = Ny i] ) } i! | i | . Z ! i rs i} E 1 in x Ea ar i “ } cam 4 i : t J Sto E y , ‘Giant i 4 ce to i aj J f Bs. [MaxwELt] Figures 12 to 14 THE VELIGER, Vol. 20, No. 2 x = % See er ae ee = Ties \ Sane t a4 Me Im i % ee es ? 2e~ 3 TNS — ee » =e pu Le . ‘ > ; ~ 23 4 Lva ae oS Lens baie eco, eh aL RY && ’ % ANY 2 3 i | BG a 7 , 3 = | i i 5 , . 5 7 ! . ANY \ at j ‘ i w ti 1 i F ] $ 4 b 3 5 \ i “e 2 | ; al 7 & | ; it it ih H \ I x a pus ree = / ce z ' 5 i B -_! Sean A i : J af 4 Ww “ au a 4 ae * 7 - Tae 7 rie : 4 : ~ , a Vol. 20; No. 2 THE VELIGER Page 75 Observations on the Copulatory Behavior of Littorina rudis (Maton) and Littorina mgrolineata Gray (Gastropoda : Prosobranchia ) BY D. G. RAFFAELLI! Department of Zoology, University College of North Wales, Bangor, Gwynedd, United Kingdom (4 Text figures) INTRODUCTION HicH INTERTDAL MARINE snails are active mainly under conditions of high relative humidity, such as occur when the tide recedes and on cool nights (Berry, 1961; own ob- servations). When the time available for movement is limited, as occurs in the supralittoral zone, copulation by males with individuals other than females of their own species will be disadvantageous. Nevertheless, casual ob- servations on the copulatory behaviour of the Littorina saxatilis species complex (for a discussion of this group see HELLER, 1975; RAFFAELLI, in prep.) suggested that interspecific copulations were not uncommon. An investi- gation of the copulatory behaviour of 2 species within the species-complex L. rudis (Maton, 1797) and L. nigro- lineata Gray, 1839 was carried out in Anglesey, North Wales in order to evaluate the extent of copulation by males with individuals other than conspecific females. METHODS Observations were made at Trwyn y Penrhyn, Anglesey (Figure 1), a south-east facing, semi-sheltered (3 on Lewis’ exposure scale; Lewis, 1964), fucoid dominated boulder shore. On 25" May, 1973, 5" and 15 May, 1975 copula- ting pairs of Littorina rudis and L. nigrolineata were col- lected as the tide receded when animal activity was great- est. Sample sizes are shown in Table 1. Pairs were con- sidered to be copulating only if the penis of one individual was inserted inside the mantle cavity of the other. This was easily seen if the copulating pair was turned apertures uppermost and gently separated. Shell heights (Figure 2) * Present address: Portobello Marine Laboratories, PR O. Box 8, Portobello, New Zealand were measured to the nearest 0.1mm and snails were dissected to determine their sex. 10km Figure 1 Location of Study Site RESULTS Approximately half the copulations in which Littorina rudis males were the ‘active partners’, 7. e., their penes were inserted inside the mantle cavity of the partner, involved L. rudis females and there was a similar trend Page 76 Figure 2 Definition of Shell Height for copulations in which L. nigrolineata males were the active partners (Table 1). Copulations by male Littorina rudis and L. nigrolineata which did not involve conspecific females were with males of their own species or with females and males of other species. There was no relationship between the size of male and female Littorina rudis and L. nigrolineata involved in intraspecific copulations (Figures 3 and 4). DISCUSSION Berry (1956) reported copulations between males of Littorina rudis and L. littorea juveniles. Although BErry THE VELIGER Vol. 20; No. 2 14 ° [e) ° 13 ° on o (O° ie Hist fo) Ss 0° & ° HT oe % a ° 2 10 O/mubROLY PAO Ic ° rz re} (o} (eo) 2 ° 5 9 ipa! : ° re) S 7 8 9 male shell height (mm) 10 II 2) ETS Figure 3 Relationship between size of male and size of female Littorina rudis involved in intra-specific copulations (1956, 1961) described his specimens as L. saxatilis, sub- sequent investigation of his population at Whitstable, Kent has shown it to be entirely L. rudis (RAFFAELLI, un- published, 1975). Copulations by males with males and females of other species and with conspecific males have Table | Frequencies of male x female intraspecific copulations of Littorina rudis and Littorina nigrolineata at Trwyn y Penrhyn. Number of copulations Date by Littorina rudis males 25.5.73 15 5.5.75 27 14.5.75 67 Number of copulations Date by Littorina nigrolineata males 25.5.75 9 5.5.75 15 14.5.75 12 Number of intraspecific male X female copulations 8 14 35 Number of intraspecific male X female copulations 3 7 8 Vol. 20; No. 2 THE VELIGER Page 77 female shell height (mm) male shell height (mm) Figure 4 Relationship between size of male and size of female Littorina nigrolineata involved in intra-specific copulations been recorded in Littorina planaxis Philippi, 1847 (Gp- son, 1964), L. scabra (Linnaeus, 1758), L. pintado Wood, 1828 and L. picta Philippi, 1846 (STRUHSAKER, 1966), L. scabra and L. irrorata Say, 1822 (GALLAGHER & RED, 1974). I have also observed male L. mariae Sacchi & Rastelli, 1966 in copulation with L. rudis females at Trwyn y Penrhyn and L. rudis males copulating with L. neritoides (Linnaeus, 1758) at Trearddur Bay, Anglesey. There is considerable evidence that Littorina species follow mucus trails made by conspecific individuals and that pheromones are involved as specific attractants (Gp- SON, 1964; DINTER « MANos, 1972; WELLS & BUCKLEY, 1972; DinTER, 1974; Hatt, 1974). In these studies there was no direct evidence of sex pheromones in the mucus trails. The present paper has shown that males of L. rudis and L. nigrolineata cannot differentiate between conspe- cific males and females, at least in the early stages of copulation, supporting the contention that female Lit- torina do not attract males with specific sex pheromones. Although the copulatory behaviour of Littorina rudis and L. nigrolineata appears to be inefficient, males copu- lating with individuals other than conspecific females may not release spermatozoa. In other studies male-male copu- lations were observed to last for less time than male-fe- male copulations, but these observations were made under laboratory conditions and their relevance to the field situation is not known. Nevertheless, when time available for mating and feeding is limited, e. g., at high shore levels, the time and energy wasted by copulating with the wrong partner may be significant and adversely affect the reproductive success of individuals living at these shore levels. SUMMARY The incidence of copulations by male Littorina rudis and L. nigrolineata with individuals other than conspecific females was found to be high. There was no relationship between size of male and female individuals involved in intra-specific copulations. It is considered improbable that mucus trails secreted by Littorina species contain sex pheromones. The inability of males to discriminate between con- specific females and other individuals may significantly lower reproductive success and fitness at higher shore levels. Literature Cited Berry, A. J. 1956. Some factors affecting the distribution of Littorina saxatilis. Ph. D. thesis, Univ. London 1961. Some factors affecting the distribution of Littorina saxatilis. Journ. Anim. Ecol. 30: 27-45 DinTER, INGRID 1974. Pheromonal behavior in the marine snail Littorina littorea Lin- naeus, The Veliger 17 (1): 37-39 (1 July 1974) Dinter, INcrip & Peter J. Manos 1972. | Evidence for a pheromone in the marine periwinkle, Littorina littorea Linnaeus. The Veliger 15 (1): 45-47; 1 text fig. (1 July 1972) Ga.iacHe_r, S. B. e G. K. Rep 1974. Reproductive behaviour and early development in Littorina scabra angulifera and Littorina irrorata (Gastropoda: Prosobranchia) in the Tampa Bay region of Florida. Malac. Rev. 7 (2): 105-125 (30 September 1974) Gisson, Daniet G., III 1964. Mating behavior in Littorina planaxis Philippi (Gastropoda: Prosobranchiata). The Veliger 7 (2): 134-139; 7 text figs. (1 October 1964) Ha.z, Joun R. 1974. Intraspecific trail following in the marsh periwinkle Littorina trrorata Say. The Veliger 16 (1): 72-75; 1 plt.; 2 text figs. (1 July 1974) Heer, J. 1975. The taxonomy of some British Littorina species, with notes on their reproduction (Mollusca: Prosobranchia). Zool. Journ. Linn. Soc. 56 (2): 131-151 Lewis, JouNn RoBert 1964. The ecology of rocky shores. London, English Univ. Press Ltd. RAFFAELLI, D. G. 1975. The determinants of zonation patterns of Littorina nerttoides and the Littorina saxatilis species-complex. Ph. D. thesis, Univ. Wales STRUHSAKER, J. S. 1966. Breeding, spawning, spawning periodicity and early develop- ment in the Hawaiian Littorina: L. pintado (Wood), L. picta (Philip- pi) and L. scabra (Linné). Proc. malac. Soc. London 37: 137-166 WELLts, Mary Jane « S. K. L. Buckiey 1972. Snails and trails. Anim. Behav. 20 (2): 345-355 v + 323 pp.; 40 plts.; 85 text figs. Page 78 THE VELIGER Vol. 20; No. 2 The Type Specimens of Ammonites hoffmanni Gabb and Melchiorites indigenes Anderson (Cretaceous : Ammonoidea ) MICHAEL A. MURPHY Department of Earth Sciences, University of California, Riverside, CA(lifornia) 92521 PETER U. RODDA California Academy of Sciences, San Francisco, CA (lifornia) 94118 (2 Plates) THE AUTHORS OF EARLY PUBLICATIONS describing and naming California fossils rarely designated primary type specimens and the existing syntypes for some of these taxa may include specimens referable to 2 or more species. This may reflect the nature of the original collections or the subsequent history of the collections. Much type ma- terial has been lost, scattered or mixed with other speci- mens. In order to achieve nomenclatural stability and taxonomic clarity for these early-described species it has been essential to redescribe existing type material and to designate type specimens. The purpose of this paper is to stabilize the nomenclature of 2 species of desmoceratid ammonoids, Puzosia hoffmanni (Gabb, 1864) and Mel- chiorites indigenes Anderson, 1938, from the Lower Cre- taceous of northern California. In proposing a solution to the problem described below we have tried to disturb the present species concept as little as possible. The concepts of these desmoceratid spe- cies currently held by most workers in the field are based primarily on ANDERSON (1938), because that paper is generally available and because it contains photographic plates which are superior to the hand-drawn plates in the earlier, scarcer papers. ANDERSON (1902: 93; 1938: 184, 187) pointed out that the syntypes of Ammonites hoffmannz include several distinct species and none closely resembled the figures in Gabb (1864). Anderson also recognized the necessity for selecting a type specimen for Ammonites hoffmanni Gabb, and he designated a specimen from the California Academy of Sciences (C. A. S. Geology Type No. 8809) as the lectotype (ANDERSON, 1938: 187; plt. 45, figs. 1, 2). This selection of a lectotype is invalid because the specimen was not chosen from existing syntypes of A. hoff- manni. A simple solution to this problem would be to choose, from among the syntypes, a new lectotype that belongs to the same species as Anderson’s invalid lecto- type. Unfortunately, none of the syntypes of A. hoffmannit belongs to the same species as Anderson’s lectotype. We exclude Anderson’s lectotype from A. hoffmanni; at pres- ent it belongs to an undescribed taxon. The confused status of A. hoffmanni Gabb affects areas far from Cali- fornia. RENz (1972: 710), following ANDERSON’s (1938) diagnosis, has applied the name Puzosia hoffmanni (Gabb) to a specimen from the Lower Cretaceous (Albi- an) of Venezuela, an action that reinforces the need to clarify this taxon. The primary types, designated herein, for both Am- monites hoffmanni and Melchiorites indigenes are chosen from 10 syntypes of Ammonites hoffmanni at the Muse- um of Paleontology, University of California, Berkeley (U.C.M.P.). Four of the syntypes are from the Voy Collection, and these are identified as Ammonites hoff- manni on the old labels with the specimens. One of these, U. C. M. P. No. 12091, also has a small label attached to the specimen on which is printed “Collection Pioche.” The Voy and Pioche Collections were used by Gabb and others in the description of California fossils. One other syntype, No. 12094, has an old label pasted to it on which is printed “Ammonites Hoffmanni.” Five additional syn- type do not have old labels. All of these specimens are be- 2 inn Page 78b THE VELIGER Vol. 20; No. 2 Table 1 Measurements of syntypes of Ammonites hoffmanni Gabb, and of other cited specimens. Figures in parentheses express measurements as percentages of shell diameters. Dashes indicate crushed or incomplete specimens from which full sets of measurements could not be made. Shell diameter Whorl height Whorl width Umbilical diameter (mm) (mm) (mm) (mm) Puzosia hoffmanni UCMP 12094 110 42 (38) 45 (41) 39 (35) UCMP 14154 57 22 (39) 292 (39) 20 (35) UCMP 14839 43 18 (42) 16 (37) 12 (28) UCMP 14921 43 18 (42) 17 (40) 14 (33) Puzosia subquadrata UCMP 12092 68 31 (46) 27 (40) 18 (26) UCMP 14155 57 26 (46) 22 (39) 16 (28) CASG 55399 69 30 (43) 97 (39) 18 (26) Melchiorites indigenes UCMP 14156 — 23 (—) 19 (—) 16 (—) UCMP 14157 84 34 (40) —(-) 27 (32) CASG 8846 78 31 (40) 25 (32) 25 (32) Melchiorites shastensis UCMP 14158 — 20 (—) 17 (—) 13 (—) CASG 8812 80 36 (45) —(-) 16 (20) CASG 8813 48 22 (46) 16 (33) 13 (27) Melchiorites sy. UCMP 12091 80) 35 (44) 29 (36) 29 (28) Lvloceras argonautarum UCMP 14153 50) 21 (42) 19 (38) 91 (42) Vol. 20; No. 2 lieved to be from collections made by, or used by, the Second Geological Survey of California (Whitney Sur- vey), and all probably were collected from the Cotton- wood Creek district, or adjacent areas, of western Shasta County, California. The 10 syntypes in the U.C.M.P collection were graciously loaned to us by Joseph Peck. In addition, we have used the paratype of Melchiorites shastensis, hypotypes of Melchiorites indigenes and Puzo- sta subquadrata, and other specimens from the collection of the Department of Geology, California Academy of Sciences (C. A. S.G.). The existing syntypes of Ammonites hoffmanni include the species discussed below. Measurements of cited speci- mens are summarized in Table 1. Puzosia hoffmanni (Gabb, 1864) (Figures 1 to 5) Ammonites hoffmanni Gabb, 1864: 65; plt. 11, figs. 13-134; plt. 12, fig. 13b Desmoceras dillevi Anderson, 1902: 97; plt. 4, figs. 116-117; pit. ro, fig. 192 Puzosia dilleri (Anderson). — ANDERSON, 1938: 185; plt. 42, figs. 1-3; plt. 46, fig. 1 Puzosia subquadrata (Anderson). — ANDERSON, 1938: 186 (in part) ; plt. 45, fig. 4 One syntype of Ammonites hoffmanni Gabb, 1864, U. C. M. P. No. 12094, is also the holotype of Desmoceras dil- leri Anderson, 1902 [= Puzosia dilleri (Anderson) ]. AN- DERSON (1938: 185) reports that he found the holotype of D. dilleri on the east fork of Huling Creek, Shasta County, but this cannot be true as an old label bearing the name Ammonites hoffmanni is glued to the specimen which is surely from Gabb’s original lot. The sandstone matrix of the specimen suggests that it could have come from Horsetown, Shasta County, the sole locality for A. hoffmanni mentioned by Gasp (1864: 65). ANDERSON (1938: plt. 42, figs. 2, 3) figured 2 additional specimens from his collection (C. A. S.G. Type Nos. 8829, 8830) as plesiotypes. Three other syntypes of A. hoffmanni (U. C. M. P. Nos. 14154, 14839, and 14921) are clearly speci- fically identical to these plesiotypes of Puzosia dilleri (Anderson). Thus 4 syntypes of Ammonites hoffmanni are identified with Puzosia dilleri (Anderson). In order to preserve present nomenclature as much as possible we choose the holotype of Desmoceras dilleri Anderson, 1902 [= Pu- zosia dilleri (Anderson) ], U.C.M.P. No. 12094, (Fig- THE VELIGER Page 79 ures 7, 2) as the lectotype for Ammonites hoffmanni Gabb, 1864 [= Puzosia hoffmanni (Gabb)]. Although this specimen differs considerably from the original figure of Gabb, it has the matrix that is common at the type locality, and it is clearly one of the syntypes because of the old label pasted to it. In addition, other specimens that are conspecific with it have been found at the same stratigraphic level as the principal fossil-bearing strata at Horsetown (Brewericeras hulenense zone) (Murpuy, 1956; Murpuy, Roppa & Morton, 1969). In making this choice we suppress the name Desmoceras dillert An- derson, 1902. The 3 other syntypes, U.C. M.P.Nos. 14154, 14839, 14921 (Figures 3 to 5) become paralectotypes of Puzosia hoffmanni (Gabb, 1864). Puzosia subquadrata (Anderson, 1902) (Figures 6 to 10) Desmoceras subquadrata Anderson, 1902: 96; plt. 4, figs. 118, 119; plt. 10, fig. 193 Puzosia subquadrata (Anderson). — ANDERSON, 1938: 186; pit. 45, figs. 3, 5 (not fig. 4 [=P hoffmanni (Gabb, 1864) ]) One syntype of Ammonites hoffmanni, U.C.M.P. No. 14155 (Figure 6), is clearly a specimen of Puzosia sub- quadrata (Anderson) with the characteristic smooth to striate shell and with pronounced constrictions of the internal mold that do not appear on the external shell. The holotype of Desmoceras subquadrata ANDERSON (1902: 96; plt. 4, fig. 118; 1938: 186; plt. 45, fig. 3), U. C. M. P.No. 12092, which was probably collected by Anderson from Huling Creek, is refigured here (Figures 7, 8). For additional comparison we have illustrated a well preserved specimen of P subquadrata, C.A.S.G. No. 55399 (Figures g, 10). Melchiorites indigenes Anderson, 1938 (Figures rz to 14) Desmoceras hoffmanni Gabb. - ANDERSON, 1902: 94 (in part); plt. 5, figs. 120-121 Melchiorites indigenes Anderson, 1938: 184; plt. 67, fig. 3; plt. 68, fig. 2 According to ANDERSON (1938: 184), the holotype of Melchiorites indigenes was in the collections of the Uni- versity of California, Berkeley, and it was the same speci- men he had figured earlier (1902, plt. 5, figs. 120, 121) Page 80 as Desmoceras hoffmanni. His 1902 drawing seems to be of a single specimen rather than a composite or idealized figure. Unfortunately he did not refigure it photograph- ically in 1938. No specimen with the same shell fracture pattern and of the same size as the figured specimen has been located in the U. C. M. P. collections and the holo- type is considered lost. Possibly the holotype of M. zndi- genes was a syntype of Ammonites hoffmanni. Two ex- isting syntypes of A. hoffmanni (U.C.M.P Nos. 14157 and 14156) resemble M. indigenes. Specimen number 14157 (Figures 17, 12) closely resembles the original figure of the holotype of M. indigenes (ANDERSON, 1902: plt. 5, fig. 120), and it is from the same general locality. This specimen also closely resembles Anderson’s plesio- type of M. indigenes (C. A. S. G. Type No. 8846) figured in 1938 (plt. 67, fig. 3), and refigured here (Figure 13). We therefore designate U. C. M. P No. 14157 as the neotype of Melchiorites indigenes Anderson, 1938. Specimen number 14156 (Figure 14) is similar to specimen 14157, but it is smaller and more poorly pre- served. Explanation of Figures 1 to 10 Puzosia hoffmanni (Gabb, 1864) Figure 1: Lectotype, U.C.M. P. 12094. No locality data. Lateral view. Diameter 110mm Figure 2: Lectotype, U. C. M. P 12094. Ventral view Figure 3: Paralectotype, U.C.M.P 14839. Cottonwood [Creek], Shasta County. (Voy Collection-. Lateral view. Diameter 43mm Figure 4: Paralectotype, U. C. M. P. 14154. (?) Cottonwood Creek, Shasta County. Lateral view. Diameter 57mm Figure 5: Paralectotype. U.C.M.P. 14921. (?) Hulen [= Huling] Creek, Shasta County. Lateral view. Diameter 43 mm Puzosia subquadrata (Anderson, 1902) Figure 6: Hypotype, U.C.M.P 14155. (?) Hulen [= Huling] Creek, Shasta County, Lateral view. Diameter 57mm Figure 7: Holotype, U.C.M.P 12092. Hulen [= Huling] Creek, Shasta County. Ventral view. Figure 8: Holotype, U.C. M. P. 12092. Lateral view. Diameter 68 mm Figure 9: Hypotype, C.A.S.G. 55399. East Fork of Hulen [= Huling] Creek, Shasta County. Lateral view. Diameter 69mm Figure 10: Hypotype, C. A. S. G. 55399. Ventral view. Note: For U.C. M. P specimens the locality data cited in the figure explanations were copied from the labels with the specimens. THE VELIGER Vol. 20; No. 2 Melchiorites sp. (Figure 75) Desmoceras hoffmanni Gabb. -— ANDERSON, 1902: 94 (in part); plt. ro, fig. 203. — ANDERSON, 1938: 187 (in synonymy, reference to ANDERSON, 1902: plt. 10, fig. 203) Melchiorites indigenes Anderson, 1938: 183; fig. 3 (8) Another syntype of Ammonites hoffmanni, U.C.M. P. No. 12091, resembles Melchiorites indigenes, but it is not clearly identifiable with that taxon. Compared with the neotype of M. indigenes it is better preserved, larger, has a greater whorl width, a deeper umbilicus with a steeper umbilical wall, and more sigmoidal constrictions which are more deeply impressed on the internal mold. The suture line of this specimen was illustrated by ANDERSON (1902: plt. 10, fig. 203), and identified as Desmoceras hoffmanni. The same figure was reprinted in reduced size by ANDERSON (1938: 183; fig. 3, No. 8) and identi- fied as M. indigenes, but he also includes the 1902 figure in the synonymy of his revised Puzosia hoffmanni (1938: Explanation of Figures 17 to 19 Melchiorites indigenes Anderson, 1938 Figure 11: Neotype, U. C, M. P 14157. Cottonwood [Creek], Shas- ta County (Voy Collection). Ventral view Figure 12: Neotype, U.C. M. P. 14157. Lateral view. Diameter 84 mm Figure 13: Hypotype, C. A.S.G. Type Collection 8846. Alderson Creek, Shasta County. Lateral view. Diameter 78mm Figure 14: Hypotype, U.C.M.P 14156. Cottonwood [Creek], Shasta County. (Voy Collection). Lateral view. Dia- meter (incomplete) 68mm Melchiorites sp. Figure 15: Melchiorites sp. Hypotype, U.C.M.P. 12091. Cotton- wood [Creek], Shasta County. (Voy Collection, ex Pioche Collection): Lateral view. Diameter 80mm Melchiorites shastensis Anderson, 1938 Figure 16: Paratype, C.A.S.G. Type Collection, 8813. Alderson Creek, Shasta County. Lateral view. Diameter 48mm Figure 17: Hypotype, U. C. M. P. 14158. Cottonwood Creek, Shas- ta County. Lateral view. Diameter (crushed) 65mm Lytoceras argonautarum Anderson, 1902 Figure 18: Hypotype, U. C. M. P. 14153. Cottonwood Creek, Shas- ta County. Lateral view. Diameter 50mm Figure 19: Hypotype, U.C. M. P. 14153. Ventral view Note: For U.C. M. P specimens the locality data cited in the figure explanations were copied from the labels with the specimens. Tue VELIcER, Vol. 20, No. 2 [Murpuy « Roppa] Figures 1 to 10 Figure 4 Figure 6 Figure 9 Figure 10 Tue VeuicEr, Vol. 20, No. 2 [Murpuy « Roppa] Figures 17 to 19 Figure 16 Figure 17 Vol. 20; No. 2 187). At present we cannot assign this specimen to a named species. Melchiorites shastensis Anderson, 1938 (Figures 16, 17) Melchiorites shastensis Anderson, 1938: 182; plt. 40, figs. 1, 2 The holotype and paratype of Melchiorites shastensis AN- DERSON (1938: 182; plt. 40, figs. 1, 2) are C.A.S.G. Type Nos. 8812 and 8813 respectively. The paratype of M. shastensis, refigured here (Figure 16), is nearly identical with one syntype of Ammonites hoffmanni, U.C.M.P. No. 14158 (Figure 17), and we exclude this specimen from Ammonites hoffmanni Gabb. Lytoceras argonautarum Anderson, 1902 (Figures 18, 19) Lytoceras argonautarum Anderson, 1902: 85; plt. 7, figs. 154, 155 Lytoceras (Argonauticeras) argonautarum Anderson. — AN- DERSON, 1938: 149; plt. 17, fig. 3; plt. 19, figs. 1, 2 The remaining syntype of Ammonites hoffmanni, U.C. M. P.No. 14153, is a small specimen of the genus Lyto- ceras, doubtless L. argonautarum Anderson (1902) be- cause of the very rapid expansion of the whorl and the subquadrate whorl section. THE VELIGER Page 81 SUMMARY Ten syntypes of Ammonites hoffmanni Gabb, 1864 [= Puzosia hoffmanni (Gabb) } are at the University of Cali- fornia, Berkeley, Museum of Paleontology, and No. 12094 is selected as the lectotype. This specimen is also the holo- type of Desmoceras dilleri Anderson, 1902, and this spe- cies becomes a junior objective synonym of A. hoffmanni Gabb. Three other syntypes belong to A. hoffmanni. Two syntypes belong to Melchiorites indigenes Anderson, 1938, and No. 14157 is chosen as the neotype of this taxon. The other 4 syntypes belong to 4 taxa: Puzosia subquadrata (Anderson, 1902), Melchiorites shastensis Anderson, 1938, Melchiorites sp., and Lytoceras argonautarum Ander- son, 1902. Literature Cited ANDERSON, FRANK MARION 1902. Cretaceous deposits of the Pacific Coast. Sci. (3, Geology) 2 (1): 1-154; 12 plts. 1938. | Lower Cretaceous deposits in California and Oregon. Geol. Soc. Amer., Spec. Pap. 16: 339 pp.; 84 plts. Gass, WiLLt1AM More 1864. Description of the Cretaceous fossils. ontology 1 (4): 55-243; plts. 9-32 Murpny, M. A. 1956. Lower Cretaceous stratigraphic units of northern California. Amer. Assoc. Petr. Geol. Bull. 40: 2098 - 2119 Murpeny, M. A., Peter U. Roppa & D. M. Morton 1969. Geology of the Ono quadrangle, Shasta and Tehama Counties, California, Calif. Div. Min. Geol., Bull. 192: 28 pp.; 1 plt. Proc. Calif. Acad. Geol. Surv. Calif., Pale- Renz, OTTo 1972. Die Gattungen Puzosia Bayle, Bhimaites Matsumoto und Des- moceras Zittel (Ammonoidea) im Oberen Albien Venezuelas. Ec- log. Geol. Helv. 65: 701 - 725; 10 plts. Page 82 THE VELIGER Vol. 20; No. 2 Chiton Fauna of the Galapagos Islands BY ALLYN G. SMITH anp ANTONIO J. FERREIRA ! (4 Plates) FOREWORD ON HIS DEATH, AucustT 18, 1976, Allyn Goodwin Smith left a nearly completed manuscript on the chiton fauna of the Galapagos Islands with the expressed request that I would complete his work and see it published. In humble spirit I strove to carry out his wish. Although in the process alterations were made upon the original manuscript, most of such alterations merely reflect the consensus we had reached in the course of discussions over the contents of the paper. Thus, Allyn’s work on the Gala- pagos chitons is preserved virtually unchanged. The credit is Allyn’s, the responsibility mine. A.J.E INTRODUCTION THE FIRST COLLECTIONS of Galapagos chitons for scien- tific purposes were made early in the nineteenth century. The large endemic species, Chiton sulcatus, was described in 1815 by Wood and C. goodall: by Broderip in 1832, the latter obtained by the famous British collec- tor and conchologist, Hugh Cuming, who probably found Acanthochitona hirudiniformis there also. Other species now known to make up the Galapagos chiton fauna are not often collected, or are small and easily overlooked. The purpose of this report is to assemble such informa- tion on Galapagos chitons as has appeared in the litera- ture, supported by specimens in the collection of the Cali- fornia Academy of Sciences and in other institutions, and by personal observations. The principal sources of records and specimens have been the major scientific expeditions « Mailing address: 2060 Clarmar Way, San Jose, California 95128 to the Islands, the earlier ones having been summarized by Stearns (1893). Since that date collections have been made during the Hopkins-Stanford Expedition in 1898- 1899, the California Academy of Sciences Expedition in 1905-1906, the Cambridge Expedition to the Suez Canal in 1924, the Norwegian Zoological Expedition in 1925, the Gifford Pinchot Expedition in 1929, the G. Allan Hancock Expedition of the California Academy of Scien- ces in 1931-1932, the California Academy of Sciences Expedition under the auspices of Templeton Crocker in 1932, the William K. Vanderbilt South Pacific Expedi- tion in 1933, the George Vanderbilt South Pacific Expedi- tion in 1937, the United States Presidential Cruise in 1938, and the Galapagos International Scientific Project in 1964. As one of the privileged participants in the Galapagos International Scientific Project (GISP), the senior author (AGS) had the opportunity to collect chitons as well as land and other marine invertebrates and to make field observations during the period January-March, 1964, while this Expedition was headquartered at the Charles Darwin Research Station on Academy Bay, Isla Santa Cruz. Most collections of chitons were made in the inter- tidal zone supplemented by a limited amount of dredging in 30m or less. Personal collecting was assisted and aug- mented by several other GISP scientists and by two en- thusiastic local conchologists living in Academy Bay Village — Mrs. Fritz (Carmen) Angermeyer and Mme. André (Jacqueline) De Roy — to whom special thanks are due. The archipelago of the Galapagos (Archipiélago de Colén) consists of 12 large and several hundred small islands on the equator, some 1000km west of Ecuador, to which country they belong. Potassium-Argon dating of the geologically oldest flows exposed indicate that the Galapagos Islands have a probable maximum age of 3,000 000 years (BaiLey, 1976). Vol. 20; No. 2 THE VELIGER Page 83 Table 1 Family, Genus, and Species Habitat LEPIDOPLEURIDAE Leptochiton albermarlensis A. G. Smith & Ferreira, spec. nov. IsCHNOCHITONIDAE Ischnochiton petaloides (Gould, 1846) CHAETOPLEURIDAE Chaetopleura cl. C. mixta (Dall, 1919) Calloplax duncanus (Dall, 1919) CALLISTOPLACIDAE Callistochiton carmenae A. G. Smith & Ferreira. spec. nov. CHITONIDAE Chiton goodalli Broderip, 1832 Chiton sulcatus Wood, 1815 Tonicia forbesti arnheimi Dall, 1903 ACANTHOCHITONIDAE Acanthochitona hirudiniformis (Sowerby, 1832) Acanthochitona jacquelinae A. G. Smith & Ferreira, spec. nov. Acanthochitona cf. A. avicula (Carpenter, 1866) Dredged, 20 m; endemic? Intertidal zone to 100 m Intertidal zone Intertidal zone Intertidal zone; endemic? Intertidal zone; endemic Intertidal zone; endemic Intertidal zone to 50 m: endemic Intertidal zone Dredged. to 50 m; endemic Intertidal zone The chiton fauna of the Galapagos Islands is not large; nor is it particularly notable except for the occurrence of 2 large, spectacular species of Chiton, C. sulcatus and C. goodalli. Other Galapagos chiton species are considerably smaller when adult, and are not especially abundant. Table 1 lists the species making up the Galapagos chiton fauna; to-date, it includes only 11 species in 8 genera, representing 6 families. It seems worth noting that possibly as many as 7 of the total 11 species, or 64%, are endemic to the Galapagos Islands although it is probable that this percentage will be somewhat reduced with more thorough collecting. In spite of the apparent high degree of en- demism, however, the Galapagos chiton fauna is essen- tially Panamic in composition. The influence of the cold, northward flowing Humboldt Current, which impinges on the Galapagos Islands, does not seem to have affected the chitons inasmuch as no species commonly occurring on the coasts of Pert and Chile are found. SYSTEMATIC TREATMENT The following account lists the known species of chitons inhabiting the Galapagos Islands, describes 3 considered to be new to science, brings together all published records, and includes comments that may be helpful in adding to the knowledge of each species. The list includes 7 species that have been reported from the Islands, which reflect either erroneous records, misidentifications, or species of such rare occurrence that they cannot be accorded per- manent status in the chiton fauna of the archipelago. POLYPLACOPHORA de Blainville, 1816 NEOLORICATA Bergenhayn, 1955 LEPIDOPLEURDAE Pilsbry, 1892 Only one representative of this family has been collected in near shore waters. In addition, one species occurs in very deep water off-shore, although further dredging is likely to produce others. Leptochiton Gray, 1847 Leptochiton albemarlensis A. G. Smith « Ferreira, spec. nov. (Figures 7, 2) Diagnosis: Animal of moderate size, round-backed and cream colored. Over-all decoration of closely spaced, fine granulations on the dorsal side of the valves. Girdle nar- row and spiculose. Page 84 Description of the Holotype: Holotype a whole animal, preserved dry, the valves all somewhat broken but show- ing the characters of the species satisfactorily. Color of both valves and girdle a light cream. Valves round- backed, the head valve rounded and somewhat more than semicircular; the intermediate valves have straight, un- mucronate posterior margins; the lateral areas hardly raised and set off from the central areas; the tail valve semicircular with a prominent, rounded, centrally-placed mucro. Dorsal sculpture consists of a series of very fine, slightly elongated, unconnected granulations arranged in close diagonal rows. There are 2-3 shallow grooves close to the side margins of the valves and parallel to them, which are somewhat better developed on valves ii and vii. The girdle has short, closely set, somewhat ap- pressed, slightly curved or straight, glass-like, blunt-point- ed spicules. Dimension are: length, 9.8mm; width, 5.4 mm; height, 1.5mm. The holotype is deposited in the California Academy of Sciences Geology Department (C ASG) Type Collection, no. 58247. Type Locality: Dredged in 20m, Tagus Cove, Isla Isa- bela (Albemarle Island), Galapagos Islands, 27 January 1968 by André and Jacqueline De Roy. A second dis- articulated specimen with all valves badly broken was dredged with the holotype. It is here designated as a paratype (CASG Type Collection no. 58248). Remarks: This new Leptochiton compares most closely in general size and shape with L. rugatus (Pilsbry, 1892) from central and southern California and from the Gulf of California. However, the latter species has the granular sculpture of the central areas on the intermediate valves arranged in connected, longitudinal rows, and the girdle is decorated with tiny sand-like granules with no spicules. Because of its spiculose girdle, L. albemarlensis may be related to L. nexus Carpenter, 1864, from moderately deep water, 37- 146m, off Catalina Island, California, which has some girdle spicules along with narrow, striated scales. However, we are not sufficiently acquainted with THE VELIGER Vol. 20; No. 2 the detailed characters of L. nexus, a poorly known south- ern California species, to do more than suggest that a possible relationship may exist. Leptochiton opacus (Dall, 1908) (Figures 3, 4) Lepidopleurus opacus Dat, 1908: 354-355 This rare and little-known chiton was dredged by the USS Albatross of the United States Fish Commission be- tween the Galapagos Islands and the Peruvian coast in 2005 fathoms (3670m), ooze, bottom temperature 35.4° F (approx. 2°C) (USFC Sta. 4647). It is illustrated here for the first time. The type lot consists of 2 specimens, preserved dry, that appear to be adults, deposited in the U.S. National Museum of Natural History (USNM 110664). One, considered to be the holotype, is represent- ed by only 7 disarticulated valves, the valve vii being missing. Because of its deep bathyal habitat, L. opacus cannot be included properly in the chiton fauna of the Islands, and it is noticed here only because its locality is in the general vicinity. IscHNOCHITONDAE Dall, 1889 Ischnochitonids are uncommon in the Galapagos Islands. The species most often collected is quite small. Lacking are the larger Panamic species in the genus Stenoplax that occur in the Gulf of California and southward, such as S. limaciformis (Sowerby, 1832) and S. magdalenensis (Hinds, 1843), and in the genus Lepidozona, such as L. clathrata (Reeve, 1847). The genus Lepidozona does not seem to be represented in the Galapagos fauna; specimens previously alluded to as possibly belonging to the genus (FERREIRA, 1974: 175) prove to be, on closer investiga- tion, representatives of the genus Callistochiton. Explanation of Figures 1, 2, 5, 13, 14 Figure 1: Leptochiton albemarlensis A.G. Smith « Ferreira, spec. nov. Holotype (CASG Type Collection, No. 58247). Albemarle Island (Isla Isabela), dredged in 20m, Tagus Cove. Dorsal view; length, 9.8mm (AJF photograph) Figure 2: Leptochiton albemarlensis A.G. Smith « Ferreira, spec. noy. Same as in Figure 1. Close-up of left side-slope to show detail of sculpture of valves and girdle. . (AJF photograph) Figure 5: Ischnochiton (Rhodoplax) petaloides (Gould, 1846). Lot #3 (CASG 40497). Academy Bay, Isla Santa Cruz, intertidally. Largest specimen, 8.7mm long (AJF photograph) Figure 13: Chiton goodalli Broderip, 1832. Academy Bay, Isla Santa Cruz. Dorsal view; length, 104.5mm (CASG 39264). From CASIZ Color Slide No. 1367 (AGS photograph) Figure 14: Chiton sulcatus Wood, 1815. Conway Bay, Isla Santa Cruz. Crocker Galapagos Expedition, 1932, coll. Leo G. Hertlein (CASIZ Collection). From CASIZ Color Slide No. 1369 (AGS phot.) Tue VELIGER, Vol. 20, No. 2 [SmirH & Frrremal] Figures 7, 2, 5, 13, 14 Vol. 20; No. 2 Ischnochiton Gray, 1847 Ischnochiton (Rhodoplax) petaloides (Gould, 1846) (Figure 5) Chiton petaloides Gou.p, 1846: 6 Radsiella petaloides. Tuorre (in Kren), 1971: 869, no. & fig. 19 Ischnochiton mariposa DaLL, 1919: 506-507 In the Galapagos Islands, this small chiton is known to attain an adult length of about 11mm. It is found sparse- ly in the intertidal zone, on the underside of rocks, to a depth of at least 100m. Specimens may be unicolored, or mottled and striped in various light to dark combinations of color; some may have the characteristic electric-blue small spots, but zebra-striped color forms have not yet turned up among the specimens of Ischnochiton petalo- ides collected in the Galapagos Islands. Ischnochiton petaloides has a unique range (A. G. SmirH, 1977). It is known from Hawaii, type locality (Maili, Oahu Id., Hawaiian Islands, leg. Kay Gudnason, low intertidal zone, 1974, CASIZ Colln.; lagoon opposite Hawaii Kai, Oahu, and E of Wailupe, Oahu, leg. H. Bertsch, Dec. 1976-Jan. 1977, AJF Colln.), the outer coast of Baja California from Punta Abreojos (26°42'N; 113°34.5’ W, in 8-10m, leg. J. H. McLean, R/V Search- er sta. 1, January 27, 1971, LACM 71-3) southwards to Cabo San Lucas, and throughout the Gulf of California. It is present, although rarely collected in Central America (El Velero, Nicaragua, intertidally, /eg. A. J. Ferreira, January 21-22, 1974, AJF 132-133; Bahia Herradura, Puntarenas Province, Costa Rica, 10-20m, Jeg. J.H. McLean, R/V Searcher sta. 447, March 9-10, 1972, LA CM 75-52; Tonasi, Las Santar, Bucaru Playa, Panama, leg. E. Bergeron, April 23, 1967, LACM B-23), and in the northern part of South America (El Rubio « Punta Mero, Tumbes Province, Peri, intertidally, leg. J. H. McLean, D. Shasky, and Pefia, April 16, 1972, LACM 72-85; Lobos de Afuera Islands, Peri, January 17, 1935, LACM- AHF 391-35; Salinas, Ecuador, 2°12’S; 80°58’ W, inter- tidally, leg. J. H. McLean, 5-6 March, 1970, LACM 70-9). In the Galapagos Islands, 14 lots of I. petaloides were collected by A. G. Smith and others at Isla Santa Cruz, Isla Genovesa, Isla Isabela, Isla San Cristobal, Isla San Salvador, and Isla Rabida. During the Califor- nia Academy 1932 Expedition, J. petaloides was collect- ed at Conway Bay, N side of Isla Santa Cruz, by Leo G. Hertlein (CASG 27232). Ischnochiton petaloides is the only chiton so far known that is common to both the Indo- THE VELIGER Page 85 West Pacific (Hawaiian Islands), and the tropical West American provinces; this observation is all the more note- worthy when only a few other mollusks, all gastropods, have been reported to span the faunistic barrier between the two zoogeographic zones (ROBERTSON, 1975). Except for the relatively smaller size of the specimens examined, and somewhat less variability in colors, Ischno- chiton petaloides from the Galapagos Islands does not seem to differ from other populations of the same species in the Gulf of California, Central America and Hawaii. Remarks: The synonymy of [schnochiton mariposa Dall, 1919 was established through the examination and side- by-side comparison of the syntype series of I. mariposa (USNM 58865) and the holotype of Chiton petaloides Gould, 1846 (USNM 12922) available on loan through the kindness of Dr. Joseph Rosewater. Within the genus Ischnochiton the systematic position of the species is, together with I. eucosmius Dall, 1919, in the subgenus Rhodoplax Thiele, 1893 (Type species, Chiton squamu- losus C. B. Adams, 1845 [= I. striolatus (Gray, 1828) ] by SD, herein) established to include relatively small, weakly-sculptured species with striated girdle scales and a tricuspid major lateral radular tooth. It should be noted that THorPE in KEEN (1971: 869) had already placed Ischnochiton mariposa in the synony- my of J. petaloides, but with a generic change to Radsiella Pilsbry (including Rhodoplax Thiele, 1893), which can- not be justified inasmuch as Radsiella Pilsbry (type species Ischnochiton tridentatus Pilsbry, 1893, by OD) was Clearly erected to include “ ... Ischnochitons (with) insertion-plates of the intermediate valves having two or several slits.” (PitsBry, op. cit. 14: 140), while Thorpe, following THIELE (1929) used the name to signify “American species” which, without supporting evidence, he considered distinct from “Old World species” of Isch- nochiton. CHAETOPLEURDAE Plate, 1899 Chaetopleura Shuttleworth, 1853 Chaetopleura cf. C. mixta (Dall, 1919) (Figures 6, 7) Tonicia mixta DALL, 1919: 515 The 11 specimens of this small Chaetopleura, referred provisionally to C. mixta, were collected in the low inter- tidal zone, under rocks, at 4 different stations in Academy Page 86 Bay, Isla Santa Cruz, by Allyn G. Smith and Jacqueline De Roy. The specimens range in length up to a maximum of gmm. They all have the typical pustulose sculpture occurring on all species in the genus. The mucro of the tail valve is median in position, which places this form of Chaetopleura in the typical subgenus. The lateral areas of the intermediate valves are prominent and decorated with 3-4, more or less anastomosing, diagonal rows of rounded, projecting pustules. The central areas have less prominent, closely-set, longitudinal pustular rows. The head valve has 15-18 anastomosing rows of pustules; on the tail valve the pustules are fewer and randomly placed. The girdle is covered with exceedingly small, chaffy, scale-like pro- cesses and with numerous, short, curved, glassy spicules; there is a well-developed foot-fringe. Color is a nonde- script brownish-green with some lighter maculations. One of the specimens, measuring 6.4mm in length, has a slit formula 8-—1-10. Remarks: All 11 specimens of the Galapagos Chaeto- pleura appear to be close to C. mixta from the Gulf of California, although smaller in average size. Also, the pustules decorating the lateral areas of the valves, and the central areas as well, tend to be slightly shorter in length, a little greater in diameter, and more regularly spaced than they are on C. mixta. Thus, if these differences do not turn out to be a criterion of age, or of individual vari- ation within the species based on a study of a larger series of specimens than those now available, this small Gala- pagos Chaetopleura may be in need of a new name. One might have expected to find Chaetopleura lurida (Sowerby, 1832) or a chiton equivalent to this hairy- girdled species in the Galapagos group. But, although abundant in the Panamic region from the Gulf of Cali- fornia to at least Panama, so far it has not been reported from these islands. THE VELIGER Vol. 20; No. 2 Calloplax Thiele, 1909 Calloplax duncanus (Dall, 1919) (Figures 8, 9) Callistochiton duncanus DALL, 1919: 512-513 -— THORPE (in KEEN), 1971: 873 This small, yellowish-white chiton was described by Dall from specimens collected on Duncan Island (Isla Pinzén) by members of the USS Albatross. The holotype is in the United States National Museum (USNM 218772). Dun- can Island must be regarded, therefore, as the type locality In addition to the holotype, available through the cour- tesy of Dr. Joseph Rosewater, series of specimens were studied from collecting at Isla Pinzén (CASG 45412), Isla Santa Cruz (ANSP 243647; CASG 40364, 40493, 40497; CASIZ Colln.), Isla Baltra (CASG 42196), Isla Santa Fé (CASG 40365), Isla San Salvador (CASG 42194), Isla Fernandina (CASG 42200), Isla Pinta (CASIZ Colln.). Remarks: Maximum adult size of Calloplax duncanus ordinarily does not exceed 12-13 mm. In sculptural details there is a fair amount of individual variation. In some specimens the central areas of the intermediate valves have a longitudinal series of threads only; in others, usually younger animals, there is a weaker series of transverse threads dividing the interspaces between the axials into squarish pits. Some also have over-all finely granular micro-sculpture. The normal number of annulate ribs on the head valve is 9, and on the tail valve 6 as indicated by Dall in his original description. However, these ribs tend to bifurcate (as they do also on the lateral areas of the intermediate valves) with the size and hence the age Explanation of Figures 3, 4, 8, 9, 12, 22 Figure 3: Leptochiton opacus (Dall, 1908). Holotype (USNM 110664). Dredged in 3667m off the coast of Peri. Side view of curled specimen. From CASIZ Color slide No. 1581 (AGS photog.) - Figure 4: Leptochiton opacus (Dall, 1908). Paratype. Dorsal view of valves i, iv, v(?), and viii; width of valve i, 9.8mm. From CASIZ Color Slide No. 1582 (AGS photograph) Figure 8: Calloplax duncanus (Dall, 1919). Isla Santa Fé. Dorsal view; length, 10.0mm (CASG 40365). From CASIZ Color Slide No. 1394 (AGS photograph) Figure 9: Calloplax duncanus (Dall, 1919). Holotype, (USNM 218772). Duncan Island (Isla Pinz6én). Dorsal view of valves i, ii, ili, vii (?), and viii; width of valve i, 3.5mm. From CASIZ Color Slide No. 1349 (AGS photograph) Figure 12: Placiphorella blainvillii (Broderip, 1832). Dredged off Cocos Island, Costa Rica, in 66 fathoms (120.7m) by the USS Albatross (USNM 122968). Dorsal view; length, 29mm; width, 27mm. From CASIZ Color Slide No. 1579 (AGS photograph) Figure 22: Acanthochitona spec. ? “Seymour Bay” [= S. Seymour Channel?], Isla Santa Cruz. Dorsal view of disarticulated valves i, No. 1351 (AGS photograph) Tue VELIGER, Vol. 20, No. 2 [SmiTH & FERREIRA] Figures 3, 4, 8, 9, 12, 22 a ee ony Vol. 20; No. 2 THE VELIGER Page 87 of the animals, resulting in the rib counts at the valve margins as high as 13 or 14 on both end valves in some instances. While the usual color of the dorsal surfaces of the valves is cream-white, a number of specimens have grayish-green mottlings, and in these the girdles have alternating bands of a similar color. Still fewer animals have light red-brown mottlings on the valves. The “velvety girdle densely covered with minute whit- ish spicules” mentioned by Dall in his original description is true of all specimens studied, especially those preserved in alcohol. The girdle “scales” are extremely minute and chaffy, the striation on them being visible only under magnification of more than 400 X. This is not the typical girdle decoration on species in the genus Callistochiton, which calls for: “Girdle poreless, densely, clothed with minute striated or smooth scales.” (Pirssry, 1893, 14: 260). In Callistochiton species the closely-packed, imbri- cating scales are “minute” but not so minute as to require high magnification to be visible. In addition, the girdle of C. duncanus shows often a fringe of spicules about 100 pm long, and occasional similar spicules interspersed amidst the girdle scales. These spicules are apparently very fragile, and will easily fall off in a dry specimen. Thus, in view of the girdle characteristics, C. duncanus cannot beassigned properly tothe genus Callistochitons.s., and appears more nearly to fit the requirements of the genus Calloplax and hence may be viewed as the Panamic regional counterpart of the Caribbean species, C. janeir- ensis (Gray, 1828). A disarticulated specimen of Calloplax duncanus shows important details: the interior is white; the sutural lami- nae are wide, separated by a rather narrow and weakly denticulated sinus; the teeth are somewhat thickened out- side at the slits which correspond in number and position to the external ribs. Slit formula 7-1-9. The eaves are rather wide, roughened, and grooved. These character- istics of the articulamentum are so much in line with those found in members of the genus Callistochiton as to sug- gest that, for their similarities, Calloplax and Callisto- chiton may eventually be considered to belong to the same family Callistoplacidae. Calloplax janeirensis (Gray, 1828) Lepidopleurus janeirensis (Gray, 1828). WimMMER, 1879: 506 Chaetopleura janeirensis. STEARNS, 1893: 449 This is an Atlantic species, reported by Wimmer from the Galapagos Islands where its occurrence is questionable. We suspect Wimmer’s specimens should be referred now to Calloplax duncanus (Dall, 1919). CaLLisToPpLacwae Pilsbry, 1893 Callistochiton Carpenter in Dall, 1879 Callistochiton carmenae A.G.Smith « Ferreira, spec. nov. (Figures 10, 11) Callistochiton shuttleworthianus Pilsbry, 1893. BERGENHAYN, 1937: 284-285, figs. 3f-3¢ (girdle scales only) Diagnosis: Animal small, yellowish-white. Head valve with 12 low, annulate ribs. Lateral areas of intermediate valves with 2 strong radial ribs; central areas with a pitted appearance. Tail valve with a low mucro, slightly anterior, and ro annular ribs. Girdle clothed with small, imbricating, oval scales. Description of the Holotype: The holotype is a whole animal, preserved in alcohol, with valves i, ii, and viii dis- articulated. Dimensions are 7.8mm in length, and 4.2 mm in width (including girdle). The color is yellowish- white uniformly. The valves are somewhat high-arched, the side slopes gently rounded to a moderately acute jugum. The head valve is semicircular with 12 low, rounded, more or less equally spaced, nodulose, annulate ribs, the posterior ones being strongest and more nodulose. Lateral areas of intermediate valves have 2 strong, radi- al ribs, the posterior ones heavily nodulose, crenulating the posterior margins of the valves. Sculpture of the cen- tral areasconsistsof about equally and closely spaced diag- onal rows of lirae extending from the valve apices and crossed at about right angles by a second series of lirae, forming rounded depressions in the interspaces and giving the central areas a pitted appearance. These lirations become less strong as they cross over the jugal ridges, there being no marked jugal areas. Tail valve has a low rounded mucro positioned slightly anterior to the center of the valve tegmentum, bounded posteriorly by a small semicircular smooth area, and anteriorly by an area sculp- tured the same as on the central areas of the intermediate valves. The tail valve margin is raised into about 10 short, heavy, somewhat pustulate annular ribs. Internally the valves are white, with a configuration typical for the genus. The slit formula is 12 — 1 — 11. The insertion teeth are slightly thickened at the edges, and weakly festooned, corresponding in position to the ribs in the tegmentum. The girdle is clothed with small, diagonally-placed, over- lapping, oval scales, which, under magnification of 200 X, show 13 or 14 slightly granulose, transverse Page 88 THE VELIGER Vol. 20; No. 2 striae at their tops, while on their sides there is a series of irregularly arranged, rather widely-placed pus- tules. Girdle width, 0.4mm. The holotype was collected at Academy Bay, Isla Santa Cruz (Indefatigable Island), leg. A.G. Smith, 15 Feb- ruary 1964 (GISP Sta. G-59), along with a small series of Calloplax duncanus. The holotype is deposited in the CASIZ Type Series no. 696. Two dry paratypes are deposited at CASG Type Series no. 58248. Five additional paratypes from the same locality, collected by Carmen Angermeyer, are preserved in alcohol and deposited in the private collection of Glenn and Laura Burghardt, Oakdale, California. Remarks: Based on the 8 available specimens, animals of this new species vary in length up to 8.7mm. There is a slight variation in the number of ribs on the head valves, which may range from 10 to 12, and on the tail valves ranging from 8 to Io. The sculpture on the intermediate and tail valves of Callistochiton carmenae differentiates it from other de- scribed species in the genus. Callistochiton carmenae be- longs in the group characterized by having the central areas diagonally ribbed which results in the pitted appear- ance of the central areas in the intermediate valves; it adds, thus, a fourth species to the group that already in- cludes the West American C. pulchellus (Gray, 1828), the Caribbean C. shuttleworthianus Pilsbry, 1893, and the Australian C’. antiquus (Reeve, 1847). The specimen from Isla Floreana reported by BERGEN- HAYN (1937) as Callistochiton shuttleworthianus Pilsbry, 1893, was located, preserved in alcohol, at the Zoological Museum of Oslo, Norway (No. D 363). The specimen was borrowed for study through the courtesy of Dr. Tor A. Bakke, Curator at the Museum. The specimen was ac- companied by a label which read: “Zool. Mus. Oslo nr, D-363 / Callistochiton shuttleworthianus, Pilsbr, / Sted: Floreana, i, strandem / 7-9-1925 / Galapagos — Explg. Bergenhayn dt.” The specimen is very small (4.5mm), completely disarticulated. Slit formula 11-1-9, The girdleis preserved entire. On the basis of the conchological characters observed, there can be no hesitancy in refer- ring Bergenhayns’ Isla Floreana specimen to Callistochiton carmenae. Callistochiton carmenae honors the malacological con- tributions of Carmen Angermeyer, of Academy Bay, Isla Santa Cruz, who, with her husband, has added much to the knowledge of the invertebrate fauna of the Galapagos Islands through assiduous shore collecting and dredging using their charter vessel, the M/V Nixe. Callistochiton gabbi Pilsbry, 1893 Callistochiton gabbi Pilsbry, 1893. BERGENHAYN, 1937: 282 to 284; figs. 3a-3e (girdle scales only) No examples of this relatively common Gulf of California Callistochiton were collected in the Galapagos Islands in 1964; nor has it been reported since. In all probability, Bergenhayn’s record (one 7mm specimen collected at Isla Floreana) is a misidentification for Calloplax duncanus. Mopa.opaéeE Dall, 1889 Placiphorella Carpenter in Dall, 1879 Placiphorella blainvillu (Broderip, 1832) (Figure 12) Chiton blainvillii BropERIP (in BRODERIP & SOWERBY), 1832: 27 Placiphorella blainvillii (Broderip, 1832). Datu, 1909: 246 The type locality is in 17 fathoms (31m), Inner Lobos Island, Peri, a “few specimens ... while dredging,” col- lected by Hugh Cuming. The United States National Mu- seum mollusk collection contains no specimen from the Galapagos Islands, although it does have one (USNM Explanation of Figures 17 to 21 Figure 17: Acanthochitona hirudiniformis (Sowerby, 1832). Aca- demy Bay, Isla Santa Cruz, intertidally. Dorsal view; length, 13.8 mm (CASIZ Collection) (AJF photograph) Figure 18: Acanthochitona jacquelinae A.G.Smith « Ferreira, spec. nov. Holotype (CASIZ Type Collection No. 967). Dredged in 40 - 50m, southern end of Academy Bay, Isla Santa Cruz. Dorsal view; length, 8.3mm (AJF photograph) Figure 19: Acanthochitona jaquelinae A.G.Smith « Ferreira, spec. nov. Same specimen as Figure 18. Side view (AJF photogr.) Figure 20: Acanthochitona cf. A. avicula (Carpenter, 1866). Sul- livan Bay, Isla San Salvador. Dorsal view of a cobalt-blue adult; length, 18.2mm (CASG 42193) (AJF photograph) Figure 21: Acanthochitona cf. A. avicula (Carpenter, 1866). Same as Figure 20. Side view (AJF photograph) Tue VELIcER, Vol. 20, No. 2 [Smirn «& Ferreira] Figures 17 - 27 i ry hava! oth 5 MPH T Vol. 20; No. 2 122968) dredged by the USS Albatross in 120m, off Cocos Island, Costa Rica (USFC Sta. 3368) (Figure 12). Dall’s record for Placiphorella blainvilli, from “Galapa- gos, Cocos and Lobos Islands,” appears as the only one published. Since there are no museum lots to substantiate the presence of P blainvillii in the Galapagos group, the record remains in need of confirmation. Cuitoniwaer Rafinesque, 1815 Chiton Linnaeus, 1758 Chiton (Radsia) goodalli Broderip, 1832 (Figure 73) Chiton goodalli BRopERIP (in BRODERIP & SOWERBY), 1832: 25 — Sowersy, 1833-1834: 3, 9; plt. 42, fig.34; plt. 43, fig. 40 — ReeEve, 1847: spec.no.8; plt.2, fig. 8 — Car- PENTER, 1857a: 180 — ParETEL, 1888: 612 — STEARNS, 1893: 404, 449 — PitsBry & VANATTA, 1902: 552 — Dat, 1909: 247 — SCHWENGEL, 1938: 2 — THORPE (in KEEN), 1971: 864; fig. 4 Lophyrus goodalli. CaRPENTER, 1857a: 317, 360 — WIM- MER, 1879: 505 Chiton (Lophyrus) goodalli. E. A. SmirH, 1877: 71 Chiton (Radsia) chierchiae NiERsTRASZ, 1908: 158-163; plt. 3, figs. 15-18, 21-28, 30-31 Chiton (Radsia) goodallii. Pispry, 1893: 14: 191; plt. 28, figs. 5-8; plt. 29, fig. 9 - BarrscH & REHDER, 1939: 17 — LELoup, 1955: 1-10; figs. 2a-2c (girdle scales) This very large, smooth, blackish species heretofore has been considered as being endemic in the Galapagos Is- lands although there is a recent record of its occurrence on the South American mainland at Santa Elena, Ecuador (Metivier, 1969: 587). Among the Recent Polyplaco- phora its adult length of 11 to 12.5cm is exceeded only by the Australian Acanthozostera gemmata (Blainville, 1825), which has been reported as long as 15cm, and the giant Cryptochiton stelleri (Middendorff, 1847), which ranges up to 30cm. Chiton goodalli was described by Broderip from specimens supplied by the British concho- logist Hugh Cuming, who obtained it during one of his voyages around the world in search of rare shells. Brod- erip recorded it from Isla San Salvador (James Island), which can be accepted as the type locality. Other records are from Isla Isabella (Pilsbry & Vanatta), Isla Floreana (Bartsch « Rehder), and from Isla San Cristébal (Schwen- gel). During the California Academy of Sciences 1905-06 THE VELIGER Page 89 Expedition, W. H. Ochsner collected the species at Isla Isabela (CASG 39557), and at Isla Espafiola (CASG 39556). In 1964, the senior author encountered Chiton goodalli fairly commonly in the intertidal zone in the vicinity of Academy Bay on Isla Santa Cruz. The animals were ob- served to congregate in groups of a few to a dozen or more in narrow cracks in the lava. At no time were they seen on top of the rocks, which seems to be the normal habitat of the large Chiton stokesii Broderip, 1832, on Cocos Island, Costa Rica. Local residents in the vicinity of Academy Bay often prize the large foot of C. goodalli as a delicacy; for this reason it was not unusual to find adult specimens drying in the open complete with shells and girdle but with the foot removed. In a series of adults specimens collected near Nelson’s on Academy Bay in 1964 by the senior author, the largest specimen preserved in alcohol in a flattened condition (CASIZ Collection) measures 123mm in length, 78mm in width, and 22mm in height. Although the dorsal surface in specimens of Chiton goodalli normally is quite smooth, some shells exhibit a tendency toward faint, subobsolete ribbing on all valves. This is a sculptural feature, however, that obviously is well within the limits of individual variation for the species. Fossil Record: Two medium-sized tail valves and an intermediate valve of Chiton goodalli were collected in a Pleistocene deposit on Isla San Salvador by the late Leo G. Hertlein during the California Academy’s 1931-1932 G. Allan Hancock Galapagos Expedition. This represents the first fossil record for the species. These valves are deposited in the California Academy of Sciences (CASG 27255). Chiton (Radsia) sulcatus Wood, 1815 (Figure 14) Chiton sulcatus Woop, 1815: 16; plt. 3, fig. 1 -— SowERBy, 1833-1834: 2, 9; plt. 39, fig. 12 - Reeve, 1847, sp. no. 15; plt. 3, fig. 15 - STEARNS, 1893: 404, 408 — Pizs- BRY & VANATTA, 1902: 552 -— DALL, 1909: 247 — ScHWENGEL, 1938: 12 -— THORPE (in KeEN), 1971: 864, fig.6 - Merevier, 1969: 586 Lophyrus sulcatus, CARPENTER, 18574: 317, 360 — WIMMER, 1879: 506 Chiton (Lophyrus) sulcatus. E. A. Smitu, 1877: 71 Chiton (Radsia) sulcatus. Pirspry, 1893: 14: 191-192; plt. 28, figs. 1-4; — STEARNS, 1893: 449 -— BOONE, 1933: 202-203; pit. 127 - BartTscH & REHDER, 1933: 17 Page 90 THE VELIGER Vol. 20; No. 2 This handsome, heavily-sculptured, blue-black chiton has been collected only in the Galapagos Islands. It was de- scribed originally by Wood, who said: “This rugged shell is said to inhabit the south seas,” a statement now known to be incorrect. It has been reported from Isla Isabela (Bartsch & Rehder; Pilsbry « Vanatta), Isla Floreana (E. A. Smith; Stearns), Isla San Cristébal (Schwengel), Isla Espafiola (Reeve; Stearns; Wimmer; Boone), Isla Santa Cruz (Stearns), Isla San Salvador (Reeve), Isla Fernandina (Tomlin), and Isla Baltra (Metivier). During the California Academy’s 1905-1906 Expedition, W. H. Ochsner collected it on several of these islands, and added Isla Pinzén (CASG). Isla San Salvador (James Island) may be accepted as the type locality for Chiton sulcatus based on Reeve’s first localized island record. In 1964, the senior author found Chiton sulcatus in Santa Cruz Island where it was the most abundant spe- cies. Generally the shells were clean, without algal or other growths on them. There was some variability no- ticed in the sculpturing on adult specimens. The largest specimen in the California Academy of Sciences’ collection (CASG 39556) was taken on Isla Espafiola by Ochsner; it measures 95mm in length, 50mm in width, and 21mm in height. Most adult specimens are smaller than this. Chiton sulcatus is active; it has a powerful muscula- ture, clinging to any hard substrate with great tenacity. Once removed, it curls up immediately and will not flatten out readily in a bucket of sea water as many other chitons do. Hence good specimens preserved in a flattened con- dition are the exception rather than the rule. Fossil Record: A single, medium-sized tail valve of Chiton sulcatus was collected in a Pleistocene beach de- posit during the California Academy’s 1931-1932 Expedi- tion by Dr. Hertlein (CASG 27250) and represents the first fossil record of the species. This fossil valve, and the 2 of C. goodalli previously mentioned, were not included by Hertlein & Strong in their account of the marine Pleis- Explanation of Figures Figure 6: Chaetopleura cf. C. mixta (Dall, 1919). Specimen from Academy Bay, Isla Santa Cruz, low intertidal zone (CASG 40290). Length, 7.3mm (AJF photograph) Figure 7: Chaetopleura cf. C. mixta (Dall, 1919). Close-up of lateral area and girdle (AJF photograph) Figure 10: Callistochiton carmenae A. G. Smith & Ferreira, spec. nov. Paratype (CASG Type Collection No. 58248). Academy Bay, Isla Santa Cruz. Dorsal view; length, 9.2mm (AJF photograph) tocene mollusks from the Galapagos Islands published in 1939. Chiton latus Sowerby, 1825 Chiton latus SowERBY, 1825 [not Chiton latus Lowe, 1825] — Boone, 1933: 200; plt. 125, fig. B Boone reported “one large specimen,” collected at Gard- ner Bay, Hood Island, Galapagos Islands, by the Ara, February 4, 1928. Mr. William E. Old of the American Museum of Natural History, New York, has seen the Boone specimen and reports it (in litt.) to be Chiton stokesu Broderip, 1832, quite probably from Cocos Island, Costa Rica, the erroneous record evidently due to a mixing of labels. This species was not collected on Hood Island (Isla Espafiola) by members of the California Academy’s 1905-1906 Expedition and has not been re- ported otherwise from the Galapagos Islands. According to Pitspry (1893, 14: 160-161), Chiton latus Sowerby, 1825, is a synonym of C’. magnificus Des- hayes, 1827, which he cites only from Valparaiso, Chile. Thus it is evident that this Chilean species is not a con- stituent of the Galapagos chiton fauna. Tonicia Gray, 1847 Tonicia forbesii Carpenter, 1857 Tonicia crenulata (Broderip, 1832) [not Chiton crenulatus Risso, 1826]. ToMuIn, 1927: 154 Tonicia forbesti CARPENTER, 1857b: 193 The first Galapagos Island record for this species is by Tomlin who reported it from Isla Floreana under the pre- occupied name of Tonicia crenulata. Although the species 6, 7, 10, 11, 15, 16 Figure 11: Callistochiton carmenae A. G. Smith « Ferreira, spec. nov. Paratype, Burghardt Collection. Academy Bay, Isla Santa Cruz. Close-up of lateral and central areas (AJF photograph) Figure 15: Tonicia forbesti arnheimi Dall, 1903. Adult specimen from Isla Baltra. Dorsal view; length, 23.3mm (CASG 40356) (AJF photograph) Figure 16: Tonicia forbesii arnheimi Dall, 1903. Same. Close-up of lateral areas to show “eyes” (AJF photograph) [Smitu & Ferrera] Figures 6, 7, 10, 11, 15, 16 THE VELIGER, Vol. 20, No. 2 pai “ing? tote g | 1 Vol. 20; No. 2 THE VELIGER Page 91 is relatively common in the Panamic Province from Maz- atlan southward, Tonicias are not common in the Gala- pagos Islands. A number of specimens was collected during the Galapagos International Scientific Project of 1964 and subsequently, both in the low intertidal zone and by dredging. These have been compared with speci- mens taken at Mazatlan, Barra de Navidad, and Acapul- co, México, and also with material from Panama (Taboga Island) and Nicaragua (San Juan del Sur). This com- parison has led to the conclusion that the differences in sculpture and size, as well as color, between Galapagos specimens and those from the mainland, plus the geo- graphical isolation of the Galapagos Islands, warrants the systematic treatment of the Islands’ population as a separate geographical subspecies of 7: forbesi. Fortunate- ly, a name for this subspecies is already available in T. arnheimi Dall, 1903; consequently the Galapagos Toni- cias related to T: forbesii will be treated under that name. Tonicia forbes arnheimi Dall, 1903 (Figures 15, 16) Tonicia arnheimi DaALL, 1903: 37-38 -— THORPE (in KEEN), 1971: 864 Tonicia arnheimi is based on a single specimen (USNM 170297), examined on a loan obtained through the cour- tesy of Dr. Joseph Rosewater. The specimen was dredged in “Noyes Cove” in 20 fathoms (36m) by Capt. W. P. Noyes, master of the Julia E. Whalen, which carried Messrs. Snodgrass and Heller to the Galapagos Islands under the name of the Hopkins-Stanford Expedition, 1898-1899. The exact location of Noyes Cove is not known but as the Expedition spent some time in Tagus Cove, Isla Isabela, probably it is close by. The holotype of T: arn- heimi was presented to the U. S. National Museum by Mr. J. S. Arnheim of San Francisco for whom it was named. Dall described Tonicia arnheimi by comparison with T. crenulata (= T. forbesit) to which he believed it to be closely related. He made a particular point of “the brilli- ancy in the eye-spots, each situated in a deep, minute pit,” which have “a metallic silvery lustre” compared with the black eye spots of T: forbesii. This indeed is a striking character in the holotype and on several of the specimens collected recently. This, however, turns out not to be a constant character in the Galapagos specimens of T. for- bes arnheimi. Adult specimens of Tonicia forbesii arnheimi average smaller in size than the mainland T. forbesii. Also, the Galapagos race has smoother and less prominently incised sculpture on the dorsal surfaces of the valves, and tends to be darker red or pinkish in color. The irregular longi- tudinal ridges on the central areas of some of the Gala- pagos specimens are more numerous and closely-set; those from deeper water have smoother sculpture than those living in the low intertidal zone; some have a smooth jugal area, lacking the pair of well-incised grooves on either side of it that is characteristic of mainland shells. Specimens dredged in Academy Bay (lots 13 and 18), and one dredged off Jervis Island (lot 14) agree closely with the holotype of T: arnheimi. Acanthopleura Guilding, 1830 Acanthopleura echinatum (Barnes, 1824) Chiton echinatus BARNES, 1824: 71; plt. 3, figs. 4a, 4b Acanthochiton spinifera (Frembly, 1827). STEARNS, 1893: 449 (= “C[hiton] aculeatus Barnes’). Acanthopleura (Corephium) echinata. Dat, 1909: 180, 248 Acanthopleura (Mesotomura) echinatum. Pirssry, 1893: 14: 218-219; plt. 47, figs. 6-17 Stearns reported this large South American chiton from the Galapagos Islands based on a specimen in the U. S. National Museum under the name Acanthochiton spini- fera (USNM 59575). It is a good-sized adult specimen measuring over 100mm in length, with the upper surface much encrusted. Dall included the Galapagos Islands in citing its range, which extends from Paita, Peru, to Val- paraiso, Chile, on the mainland. Pilsbry also includes the Galapagos group for this species based on specimens in the collections of the Academy of Natural Sciences of Philadelphia. This conspicuous chiton, with a girdle set with sparse, strong, spike-like spines, can hardly be mistaken for any other species. It was not collected by Snodgrass and Hel- ler during the Hopkins-Stanford Expedition of 1898-1899, nor by Ochsner during the 1905-1906 California Acade- my Expedition. No specimens were collected in Galapagos in 1964 by the senior author, or the local collectors. Why the species has not been collected in the Galapagos Islands in recent years is a mystery. Inclusion of Acanthopleura echinatum in the permanent chiton fauna of the Gala- pagos Islands is therefore withheld pending its possible re-discovery. Page 92 Enoplochiton Gray, 1847 Enoplochiton niger (Barnes, 1824) Chiton niger BARNES, 1824: 71; plt. 3, fig. 3 Tonicia ? coquimbensis (Frembly, 1827). STEARNS, 1893: 449 Enoplocniton niger (Barnes, 1824). Prrspry, 1893:14:252- 253; pit. 52, figs. 22-29 Stearns’ published record is based on a single adult speci- men (USNM 59576). It is the only known one from the Galapagos Islands. The species is a large one with a unique type girdle set with massive, widely-spaced, deep- ly-imbedded scales. It cannot be mistaken for any other described chiton. Pilsbry records it from Pert and Chile, but not from the Galapagos Islands. The record needs confirming. AcANTHOCHITONWDAE Pilsbry, 1893 Acanthochitona Gray, 1821 Acanthochitona hirudiniformis (Sowerby, 1832) (Figure 17) Chiton hirudiniformis SowERBY, (in BRODERIP & SOWERBY), 1832: 59 — SOWERBY, 1533-1834: 7, 10; plt. 41, fig. 23; pit. 174, fig. 142 -— ReEEvE, 1847: spec. no. 54, plt. 10, fig. 54 — CARPENTER, 1857a: 181, 318, 360 Acanthochites hirudiniformis. WIMMER, 1879: 506 — Pus- BRY, 1893: 15: 27; plt. 2, figs. 49, 56 -— Pirssry & VaNATTA, 1902: 552 — DaLt, 1909: 246 ? Tonicia hirudiniformis. STEARNS, 1893: 449 Acanthochitona hirudiniformis. THoRPE (in KEEN), 1971: 866, 868, fig. 13 (p. 867) This species was described by Sowerby from Ancén, Lobos Island, and Paita, Peri, on the mainland of South Ameri- ca, and from Chatham Island (Isla San Cristébal), Gala- pagos Islands. His account, which is quite brief, is re- peated by Pilsbry in the Manual of Conchology (1893). Pilsbry had specimens from Isla Pinzon, Isla Fernandina, and from Tagus Cove, Isla Isabela, the latter collected by Snodgrass during the 1898-1899 Hopkins-Stanford Ex- pedition; these specimens are in the collection of the De- partment of Mollusks, Academy of Natural Sciences of Philadelphia. A sizeable series was collected by W. H. Ochsner in Iguana Cove, Isla Isabela (CASG 37082), and on Isla Pinzén (CASG 37268) during the California Academy’s 1905-1906 Expedition. Members of the GISP THE VELIGER Vol. 20; No. 2 found it on Isla Santa Cruz, Isla Genovesa, and Isla Pinta in 1964. On Isla Santa Cruz, in 1964, Acanthochitona hirudini- formis was found fairly commonly in the general vicinity of Academy Bay from the middle to the low intertidal zone. The species occurs in some abundance on many (and very likely on all) of the islands of the Galapagos Archi- pelago. The animal is rather small and dark-colored, blending perfectly with the color of the lava blocks on and under which it lives; it escapes notice easily because of the habit of nestling in small depressions. It was observed that young specimens around Academy Bay were subject to considerable predation from large gastropods, includ- ing Thais planospira, Th. melones, Acanthina muricata, Purpura columellaris and P. pansa. Extended Description: Galapagos specimens of Acan- thochitona hirudiniformis agree in general with Sowerby’s original but brief diagnosis. The valves are arched, not carinated, the intermediate valves only slightly mucro- nate. The exposed portion of the intermediate valves is subtriangular, usually appearing black when unworn but sometimes with flecks of lighter color. The head valve is semicircular, rounded, and generally has a straight poste- rior margin. On the intermediate valves and the tail valve the wedge-shaped jugal areas are raised and sharply de- fined, with a sculpture of transverse growth-lines, faint longitudinal lirae, and occasional remnants of granular sculpture; this is scarcely visible in perfect specimens but more distinct when the dorsal surface is a little worn. The lateropleural areas have an even sculpture of small, close- ly-spaced, ovate, flat-topped granules. The tail valve has a small, nearly circular termentum, the anterior part being a little contracted with the mucro positioned slightly be- hind the middle. The interior of the valves is blue, more or less purplish at their centers. The insertion plates are long; in the head valve it is about half as long as the tegmen- tum; in the tail valve it is nearly vertical and deeply sinus- ed in an end view. The slit formula is 5—1~-2.The girdle is thick and wide, sepia or brownish in color, and densely covered with very short delicate spicules, giving it a vel- vety appearance; there is a dense fringe of longer spicules around the periphery. The 18 sutural tufts are colored like the girdle but appear lighter in tone under good lighting; those at the sutures of the intermediate valves are about 5mm long while those around the head valve are some- what shorter, Remarks: There has been some uncertainty concerning the proper identification of Acanthochitona hirudinifor- mis, It isnot possible to say with assurance whether Sower- by’s figure 142 on plate 2 of the Conchological IIlustra- Vol. 20; No. 2 tions of 1833-1834 (the better of his 2 illustrations) re- presents a Peruvian or a Galapagan specimen. Because of this uncertainty, Pirspry (in MS) considered Isla San Cristébal (Chatham Island) to be the type locali- ty, a suggestion we propose be accepted. This action will establish the Galapagos Islands as the type locality, rather than one of the Peruvian localities (Ancén, Lobos Island, or Paita). In the California Academy collection there is a series of 3 small, dry, curled specimens of an Acanthochitona collected by Don L. Frizzell at Paita, Peru, in 1938. Since they came from one of the original localities for A. hzru- diniformis cited by Sowerby, a comparison with Galapa- gos material has some significance. This shows several conchological differences. The specimens from Paita are smaller in size; the oval, flat-topped granules on the latero-pleural areas are slightly larger and are much less numerous, not being as closely spaced. Also, the jugal areas are smaller in extent, especially on valves iv to vii, inclusive, being much narrower. The sutural tufts are whitish rather than sepia or brownish-olive as on Gala- Pagos specimens. From the above evidence, which admittedly is insuffi- cient, we suspect the specimens from Paita to represent a species of Acanthochitona different from the one com- mon to the Galapagos Islands. Should this prove to be the case after study of more and better material from Peri, another name can be allocated to the species from the mainland of South America, which can be selected from A. coquimbensis Leloup, 1941, A. peruvianus Leloup, 1941, or A. tabogensis A. G. Smith, 1961. It should be noted that THorPE (7n KEEN, 1971) places all 3 of these species in the synonymy of A. hirudiniformis without pro- viding any substantiating evidence for these allocations. Because adequate material from Peri and elsewhere on the mainland south of Panama does not yet seem to be available, no authoritative judgment can be made at this time. In arriving at a final conclusion, there are several earlier-collected specimens that should be taken into ac- count. As an example, in the mollusk collection of the Academy of Natural Sciences of Philadelphia there are 2 specimens of Acanthochitona from Peri (ANSP 35788) received from Dr. T: B. Wilson. The larger specimen meas- ures 36 X 18mm. They were in the old collection of the Philadelphia Academy under the name Chiton hirudini- formis and were purchased in London some time between 1846 and 1849, probably from Hugh Cuming. These spe- cimens differ from the Galapagos A. hirudiniformis in several sculptural characters and in the girdle decoration, which has stout spines scattered among the finer girdle spicules. They are not mentioned by Pilsbry in the Manual THE VELIGER Page 93 of Conchology. It is possible that Sowerby’s figure 142 in the Conchological Illustrations represents this form. The United States National Museum of Natural His- tory has 3 lots of small Acanthochitonas from South America, 2 of them labeled Acanthochitona hirudiniform- is. The first (USNM 5804) came from Orange Harbor, Patagonia, collected during the United States Exploring Expedition, and probably identified by PP.Carpenter. It is a dry, curled specimen in poor condition, the girdle ornamentation being almost completely gone. The second (USNM 19284), identified by Carpenter according to the museum label, came from Valparaiso, Chile, and was also taken during the U.S. Exploring Expedition. It is dry and curled; the valves, which are disarticulated, are much worn and chipped. The third specimen (USNM 218733) is unidentified as to species and came from between Cape Pillar and Cape Horn, collected by Stokes. It is complete and measures 17 X 8.5mm. The tegmentum is light gray to whitish with dark zig-zag markings around the margins of the valves. Sculpture on the latero-pleural areas con- sists of closely-spaced, small, round granules. So far as can be determined, all of these South American specimens of Acanthochitona differ in several respects from A. hiru- diniformis of the Galapagos Islands and probably repre- sent other species. The Stokes specimen from near Cape Horn possibly can be referred to A. stygma (Rochebrune, 1889), an allocation that is problematical as we have not seen authentic specimens of Rochebrune’s species. Acanthochitona jacquelinae A. G. Smith « Ferreira spec. nov. (Figures 78, 79) Diagnosis: ‘This is a small species taken so far only by dredging in 40 to 50m in the southern end of Academy Bay, Isla Santa Cruz, and off Isla San Salvador. Most specimens are reddish in tone although a few are mottled or white. It is characterized mainly by the wide-spreading spicular tufts, which are unusually prominent for such a small sized chiton. This feature is the principal one that distinguishes it from other Acanthochitonas described from the Eastern Pacific. Description: Animals very small for the genus, generally not exceeding 10mm in length, elongate-ovate, the valves arched but not carinate, intermediate valves mucronate. Color more often mottled with white and reddish tones, with many colored an over-all light orange-red or terra- cotta red; a few specimens are pure white. The girdles are colored the same as the valves and are decorated with the usual spicular tufts or pore-bunches. These tufts Page 94 THE VELIGER Vol. 20; No. 2 are large and wide-spreading for the size of the animal, the spicules being 1.0 to 1.5mm long, yellowish in tone except those which are white on pure white specimens. The girdles are decorated, in addition, with extremely fine, upward-trending and inward-curving, pointed spic- ules, sometimes quite numerous close to the side-margins of the valves and surrounding the bases of the pore- bunches. Below these, the girdles have only a few smaller, scattered spicules but they terminate in a well-developed foot-fringe of closely-packed, straight, pointed spicules about half as long as those making up the spicular tufts. The jugal areas of the intermediate valves are quadri- lateral, cut into 7-10 longitudinal, somewhat beaded lirae by much narrower incised grooves. Latero-pleural areas of intermediate and end valves sculptured with a series of small, round or suboval, slightly convex or flat- topped granules, closely spaced in quincunx rather than in straight rows. The mucro of the tail valve is rounded and centrally placed. Internally, the valves are orange-red (except in pure white specimens). The combined insertion plates and sutural laminae are not especially wide in relation to the tegmentum area; they are smooth-surfaced, and have the usual 5-1-2 slit formula. The slits are deep, near parallel-sided, but are not thickened or buttressed at their sides. The anterior sinus of the tail valve is about 4 its entire width. Type Locality: In 40-50m (21.3-26.5 fms.) on a broken coralline bottom off Isla Coamano (Jensen Is- land) at the southern end of Academy Bay, Isla Santa Cruz, dredged by André and Jacqueline De Roy. A total of 67 specimens was collected in this area, g on Septem- ber 29, and 58 on December 28, 1966. Type Material: ‘The holotype is an orange specimen from the type locality (CASIZ Type Series no. 967), measuring 8.3mm in length, 4.2mm in width, and 1.5 mm in height. The remainder of the type lot (66 specimens) is hereby designated as paratypes, representative examples of which will be deposited in various museums, and in the private collection of Mme. Jacqueline De Roy. Other Material: Another specimen, preserved in alco- hol, measuring 7.9mm in length, was dredged in 9 - 18m (5-10fms.) in Academy Bay on a hard coralline-mud bottom by D. P. Abbott, J. L. Barnard, J. W. Durham and A. G, Smith, 24 February 1964. It was found nestling in a small chunk of hard coralline alga. The basic color of this specimen is white with sea-green markings on valves ill, iv, vii, and viii. The girdle is marbled with reddish brown. The spicular pore bunches are grass-green at their bases, blending into yellowish-white towards their tips. The foot-fringe is colored alternately with lavender and white. A single, tiny, whitish specimen was dredged in 55m (30.1 fms.) in Academy Bay near the type locality by the De Roys, 26 April 1967 (CASG 40495). A small curled specimen, preserved dry, was dredged in 25m off Isla San Salvador by the De Roys, 27 March 1967 (CASG 40323) ; it has the same reddish color as most of the Acad- emy Bay specimens. A single, pinkish specimen dredged in 20-40m (11-22fms.) off Isla Rabida by the De Roys, 22 March 1967, preserved dry (CASG 40492). Remarks: This lovely little chiton is distinct and easily separable from any specimens of Acanthochitona from the Eastern Pacific known to us. The adequate series at hand, which ranges in length from a little over 1mm to hardly more than romm, is quite constant in sculptural and other details, although the color varies from mottled reddish and white to over-all light red tones. In the total of 71 specimens available, 39 (557%) are mottled red and white, 25 (35%) are over-all reddish, and 7 (10%) are pure white or yellowish. Acanthochitona jacquelinae is most closely related to A. arragonites (Carpenter, 1857) from the Gulf of Cali- fornia; both are about the same size. However, it differs from the latter in the following particulars: (1) the jugal tract is finely ribbed longitudinally compared with the smooth jugal tract of A. arragonites; (2) the pus- tules on the latero-pleural areas are smaller and rounder than the more oval shaped pustules in A. arragonites, and arranged in quincunx rather than in straight rows; (3) the basic girle spicules generally are sparse and very small whereas in A. arragonites they are more numerous and interspersed with spicules of varying sizes. Acanthochitona jacquelinae differs from A. avicula (Carpenter, 1866) in: (1) itsmuch smaller size; (2) much brighter colors, often in tones of red to yellow or white, but not blue (com- mon in A. avicula); (3) much smaller and rounder pus- tules on the latero-pleural areas; (4) thinner and shorter basic girdle spicules, and, in contrast, much longer (for the size of the animal) spicular pore-bunches resulting in a bushier appearance. Acanthochitona jacquelinae is named for Mme. Jac- queline De Roy of Academy Bay, Galapagos Islands, who was instrumental in collecting the type lot and who has been contributing much to the knowledge and under- standing of the molluscan fauna of the Galapagos Islands. Vol. 20; No. 2 Acanthochitona cf. A. avicula (Carpenter, 1866) (Figures 20, 21) Acanthochites avicula CARPENTER, 1866: 211 Ten specimens of a still different Acanthochitona have been collected by the De Roys on or in the vicinity of 3 of the larger islands of the Galapagos Archipelago, and 2 of the smaller ones, either in the low intertidal zone, or dredged to a depth of 30m. The specimens examined are in 6 lots as follows: (1) 2 specimens in alcohol (CASIZ Colln.), collected in Aca- demy Bay, Isla Santa Cruz, 1966; 14mm and 12.1mm long; (2) one specimen, dry (CASG 40494) dredged in 10 - 30m off Beagle Island, 20 March, 1967; length, 7.5 mm; color, whitish with longitudinally arranged light- brown, zig-zag markings on the side-slopes of the valves; pore-bunches of spicules are bluish-green while the other girdle spicules are lavender in tone; (3) 2 specimens, dry (CASG 42194), about 7.5mm in length, collected at low tide in Sullivan Bay, Isla San Salvador, 23 Novem- ber, 1967; color mottled light green; girdle spicules lavender, pore-bunches colorless; (4) 2 specimens, dry (CASG 42198), dredged in tom, Tagus Cove, Isla Isabela, 22 January, 1968; one whitish, the o- ther dark slate-grey; (5) 1 specimen, 8.5mm _ long, dry (CASG 42199) dredged in 8m off Isla Sombrero Chino, 1 February, 1969; mottled whitish and light brown; pore-bunches, lavender; (6) 2 specimens, dry (CASG 42193), collected 2 February 1969, in low inter- tidal zone, Sullivan Bay, Isla San Salvador; the largest measures 18.2mm in length, 8.2mm in width, and 2.9mm in height (Figures 20, 21); the spicules in the pore- bunches as well as those covering the girdle are a brilliant iridescent cobalt-blue. Remarks: All of the specimens at hand for study are small in size (with only one exception which is 18.2mm long), and apparently juveniles. The dorsal sculpture, especially the occurrence of closely-spaced, fine, longi- tudinal lirae on the well marked jugal areas agrees with a similar sculpture character on specimens of Acantho- chitona avicula in the California Academy’s collections from Mission Bay, San Diego County, California, and from several localities in the Gulf of California, and Scammons Lagoon, Baja California, Mexico. However, the Galapagos specimens do differ from the A. avicula of northern waters in the very small size of the spicules covering the girdle; in specimens of A. avicula collected in their known range from San Diego to the Gulf of Cali- fornia, the girdle is covered with relatively long and thick spicules, almost the size of the spicules at the fringe, THE VELIGER Page 95 while the examined specimens from the Galapagos Is- lands show no such a character inasmuch as the girdle is covered with spicules which are quite small and thin. Another possible difference between the northern and the Galapagos populations was found in the examination of the radula: in specimens of A. avicula from San Diego and the Gulf of California the radula’s median tooth is as wide in the front as (or even narrower than) in the back, whereas the radula of a specimen from the Gala- pagos Islands (11mm long) has a median tooth which is much wider in the front than in the back. It is interesting to note that in these 2 characters the Galapagos specimens approach A. hirudiniformis more than A. avicula. How- ever, A. avicula, not previously recorded south of the Gulf of California, is given to considerable intraspecific variation (in color, size and shape of the pustules in the latero-pleural areas, degree of striation in the jugal areas, size and color of the pore-bunches spicules), and the dis- tinctions observed in the Galapagos specimens, particular- ly in view of their being juveniles, do not appear large enough to contradict the notion of conspecificity. Thus, until additional adult specimens of this particular species of Acanthochitona can be collected and compared with other described species, it seems best to refer those at hand to A. avicula (Carpenter, 1866) subject to a subsequent review. Acanthochitona species? (Figure 22) In the collection of the Academy of Sciences of Phila- delphia is a series of valves (head, tail, and 3 intermedi- ates) of an Acanthochitona collected in Seymour Bay, Isla Santa Cruz, under a stone, at low tide, by H. A. Pils- bry, 22 June 1929 (ANSP 153484). The girdle of the specimen is missing. Pilsbry recognized it as a new species, and provided a description which is still in MS at the Philadelphia Academy. According to Pilsbry’s descrip- tion, the species would differ from other Eastern Pacific species in the broad shape of the intermediate valves, the coarser granulations [pustules] of the dorsal surfaces, and the minute girdle spicules, so small that the girdle ap- peared smooth under a hand lens. The specimen was borrowed for study and illustration through the courtesy of Dr. R. Tucker Abbott and Mrs. Virginia Orr Maes. Because it is incomplete, and because no other specimens like it have turned up, the above ac- count is included to call attention to the possibility of still another undescribed species of Acanthochitona oc- curring in the chiton fauna of the Galapagos Islands. Page 96 ACKNOWLEDGMENTS For cooperation in the loan of key specimens, apprecia- tion is expressed to Dr. Joseph Rosewater of the Division of Mollusks, United States National Museum of Natural History, to Dr. R. Tucker Abbott of the Delaware Muse- um of Natural History, to Mrs. Virginia Orr Maes of the Academy of Natural Sciences of Philadelphia, Depart- ment of Mollusks, to Dr. George E. Radwin of the San Diego Museum of Natural History, and to Mr. and Mrs. Glenn Burghardt of Oakdale, California. Mr. Spencer R. Thorpe of El Cerrito, California, provided comments and suggestions during the course of the study, and Mr. Maurice Giles, the California Academy’s scientific photo- grapher supplied his expert work in many of the black- and-white illustrations used, Special appreciation is due to Mr. Barry Roth, Department of Geology, California Academy of Sciences, for valuable assistance in many phases of this work and the critical reading of the manu- script. Literature Cited Baitey, KimBerLy 1976. Potassium-argon ages from the Galapagos Islands. 192: 465 - 467 Barnes, D. W. 1824. Description of five species of Chiton. 69-72; plt. 3, figs. 1-4 Bartscu, Paut & Haratp ALFRED REHDER 1939. Mollusks collected on the Presidential Cruise of 1938. Smithson. Misc. Coll. 98 (10): 1-18; 5 plts. (13 June 1939) BERGENHAYN, J. R. M. 1937. Polyplacophoren von den Galapagos Inseln. Norveg. Exp. to Galapagos Is., 1925, conduct. by Alf Wollebaek. Meddel. Zool. Mus. Oslo, no. 49, contr. XIV (Nytt Mag. Naturvidens. 76): 273 - 286; figs, 1-3 Boone, Lez 1933, Scientific results of the yachts Eagle and Ara, 1921-1928, Wil- liam K. Vanderbilt commanding. Coelenterata, Echinodermata, and Mollusca. Bull. Vanderb. Mar. Mus. 4: 1-217; 133 pits. (30 April 1933) Broperip, WILLIAM JoHN & GEeorGE BRETTINGHAM SOWERBY 15st 1832-1833. Characters of new species of Mollusca and Conchifera, col- lected by Mr. Cuming. Proc. Zool. Soc. London for 1832: 25 - 33 (21 April 1832); 50-61 (5 June 1832); 104-108 (31 July 1832); 124-126 (14 August 1832); 173-179 (14 January 1833); 194-202 (13 March 1833) CarPeNnTER, PHitip PEARSALL 1857a. Report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Brit. Assoc. Adv. Sci. Rprt. 26 (for 1856): 159-368; plts. 6-9 + 4 pp. (pre 22 April) [dating: Carpenter, 1857c: iv] 1857b. Catalogue of the Reigen collection of Mazatlan Mollusca, in the British Museum. London, Brit. Mus.: i-iv+ix-xvi+1-552 (1 August 1857) 1866. Descriptions of new marine shells from the coast of California. Pt. Ill. Proc. Calif. Acad. Sci., (1) 3: pp. 207 - 224 (February, 1866) Dati, Witit1am HEALEY 1903. Two new mollusks from the west coast of America. The Nautilus 17 (4): 37 - 38 (12 August 1903) 1908. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U. S. Fish Commission steamer “Albatross,” during 1891, Lieut.-Com- Science (30 April 1976) Amer. Journ. Sci. 7: THE VELIGER Vol. 20; No. 2 mander Z. L. Tanner, U. S. N., commanding. XXXVII. Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U. S. Fish Commission steamer “Albatross,” from October, 1904, to March, 1905, Lieut.-Commander L. M. Garrett, U. S. N., commanding. XIV. The Mollusca and Brachio- poda, Bull. Mus. Comp. Zool. 43 (6): 205 - 487; pits, 1-22 ie (22 October 1908) 1909. Report on a collection of shells from Peru, with a summary of the littoral marine Mollusca of the Peruvian zoological province. Proc. U. S. Nat. Mus. 37 (1704): 147-294; plts. 20-28 (24 Nov.) 1919. Descriptions of new species of chitons from the Pacific coast of America. Proc. U. S. Nat. Mus. 55 (2283): 499-516 (7 June’19) Dersuayes, Gérarp PauL 1827. Oscabrion. In: Bory de Saint-Vincent et al., Dictionnaire classique d’histoire naturelle. vol. 12: 446 - 457. Paris Goutp, Aucustus ADDISON 1846. Expedition shells: described for the work of the United States Exploring Expedition, commanded by Charles Wilkes, U.S.N., during the years 1838-1842. Proc. Boston Soc. Nat. Hist. 2: 142 - 152 [reprinted in Otia Conchologica, 1862] (July 1846) HerTEIN, Leo Gzorcz & ARCHIBALD MoCLureE STRONG 1939. Marine Pleistocene mollusks from the Galapagos Islands. Proc. Calif: Acad. Sci. (4) 23 (24): 367 - 380; plt. 32 (20 July ’39) Keen, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. Stanford Univ. Press, Stanford, Calif. i - xiv+ 1064 pp.; ca. 4000 figs.; 22 color plts. (1 September 1971) Le.oup, EucENE 1941. A propos de quelques Acanthochitons peu connus ou nouveaux, III. Région pacifique, cétes américaines. Bull. Mus. roy. Hist. Nat. Belg. 17 (61): 1-9; plt. 1; 5 text figs. (October 1941) 1954. Chaetopleura unilineata n. sp., chiton de l’Equateur. Inst. roy. Sci. nat. Belg., Bull. 30 (40): 1-6; 2 text figs. (December 1954) 1955. Chiton (Radsia) Goodallii Broderip, 1832 = Chiton (Radsia) chierchiae Nierstrasz, 1908. Bull. Inst. roy. sci. nat. Belg. 31 (23): 1-10; 2 text figs. (June 1955) MetTivier, BERNARD 1969. Mollusques marins récoltés par M. R. Hoffstetter sur les cétes de ’Equateur et aux Iles Galapagos. Chitonidae et Fissurellidae. Bull. Mus. Nat. Hist. Nat. 40(2) (3): 585-596 (31 January 1969) Nigerstrasz, Huco Friepricw 1908. Remarks on the Chitonidae. Vereen. 10: 141-172; plt. 3 PagTEL, FRIEDRICH 1887-1894. Catalog der Conchylien-Sammlung, Berlin. vol. 1, fasc. 1-8: 1-639+16 (1887-1888); 2, fasc. 9-14: 1-505+12 (1888 to 1890); 3, fasc. 15-18: 1-256+40 (1890 - 1891) Pitssrr, Henry Aucustus 1892-1894. Polyplacophora. Manuel of Conchology 14: i - xxxiv+ 1 - 350; 68 plts.; 4 text figs. (25 July 1892 to 1 July 1893); 15: 1 - 133; plts. 1 - 17 (16 November 1893 to 19 March 1894) Philadelphia Pitsspry, Henry Aucustus & EpwArp GuiREY VANATTA 1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898- 1899. XIII. Marine Mollusca. Proc. Wash. Acad. Sci. 4: 549 - 560; pit. 35 (30 September 1902) REEVE, LOVELL AUGUSTUS 1847-1848. Monograph of the genus Chiton. 4; plts. 1-28 (figs. 1- 194) Rosertson, Robert 1976. Heliacus trochoides: An Indo-West-Pacific architectonicid newly found in the Eastern Pacific (mainland Ecuador). The Veliger 19 (1): 13-18; 1 plt.; 1 map; 1 table (1 July 1966) ScHWENGEL, JEANNE S. 1938. Zoological results of the George Vanderbilt South Pacific Expe- tion, 1937, Part 1, Galapagos Mollusca. Proc. Acad. Nat. Sci. Philadelphia 90: 1-3 (13 May 1938) SmitH, ALLyN GoopWIN 1977. Rectification of west coast chiton nomenclature (Mollusca : Polyplacophora). The Veliger 19 (3): 215-258 (1 January ’77) SmitH, Epcar ALBERT 1877. Mollusca No. IV: An account of the zoological collections made during the visit of H.M.S. ‘Petrel’ to the Galapagos Islands, communicated by Albert Gunther. Proc. Zool. Soc. London for 1877: 69-73; pit. 11 (6 February 1877) Sowersy, GEoRCE BRETTINGHAM !8t 1825. A catalogue of shells contained in the collection of the late Earl of Tankerville. pp. i-vii, 1-92; app. i-xxxiv; 9 color plts. London (August 1827) Tidjschr. Nederl. dierkund. Conchologica Iconica, London (Febr. 1847 to Jan. 1848) Vol. 20; No. 2 THE VELIGER Page 97 Sowersy, Georce BRETTINGHAM 2nd 1841 [?1840] The conchological illustrations. A catalogue of the Recent species of Chitones. pp. 1-10; plts. 38-45 (figs. 1-55); plts. 159 to 176 (figs. 56-155). London STzARNS, Ropert EDWARDS CARTER 1893. Report on the mollusk-fauna of the Galapagos Islands, with descriptions of new species. Sci. Reslts. Explor. by the U.S. Fish Comm. Steamer Albatross — XXV. Proc. U. S. Nat. Mus. 16 (942): 353-450; pits. 51, 52; map Tomuin, JoHN Reap Le BrockKTON 1927. Zoological results of the Cambridge Expedition to the Suez Canal, 1924, XVII. Report cn the Mollusca (Amphineura, Gastro- poda, Scaphopoda, Pelecypoda). Trans. Zool. Soc. London 22 (3): 291-320 (July 1927) WimM_ER, AUGUST 1879. Zur Conchylien-Fauna der Galapagos-Inseln. Sitzungsber. k.-k. Akad. Wissensch. (Wien), Math.-naturw. Klasse 80 (5): (I): 465- -514 Page 98 THE VELIGER Vol. 20; No. 2 Notes on Sea Hares of South Texas (Gastropoda : Opisthobranchia ) BY NED E. STRENTH anv JAMES E. BLANKENSHIP Marine Biomedical Institute, Galveston, TX 77550 THOSE SPECIES OF SEA HARES reported from the South Texas Coast are the source of extensive misidentification and widespread nomenclatural confusion. Synonyms as well as inaccurate descriptions are common in recent liter- ature (BREUER, 1962; ANDREWS, 1971; ABBOTT, 1974; and Neck, 1976). Major scientific works such as those by Eates (1960) and Marcus (1972) have apparently been overlooked in many instances. The purpose of this paper is to facilitate ease of identification and eliminate nomenclatural confusion with respect to those species cur- rently recognized from South Texas. The results of this study are based on field work conducted in the vicinity of South Padre Island, Cameron County, during the previous 3-year period. It should be noted that frequency of sightings and abundance of specimens of the various species is sporadic and seemingly unpredictable. The following 5 species of sea hares are currently reported from South Texas. The 4 species of Aplysza belong to the subgenus Varria. Primary synonyms are listed for each species. In several instances new combinations are given due to associated change in the generic name of the taxon. Aplysia Linnaeus, 1767 Aplysia brasiliana Rang, 1828 Synonymy: Aplysia livida D’Orzicny, 1837: 206 Aplysia cailleti DEsHAYES, 1857: 140 (see EALES, 1960: 297) Aplysia guadeloupensis SowERBY, 1869: plt. 5 (see EALEs, 1960: 297) Aplysia willcoxi He1Lprin, 1886: 364; - 1887: 130 Tethys willcoxi var. perviridis Puspry, 1895: 81 Tethys floridensis Pirspry, 1895: 82 Remarks: ‘This is the common mottled sea hare exten- sively referred to as Aplysia willcoxi in both the popular and scientific literature. It is currently placed in syn- onymy with A. brasiliana (STRENTH & BLANKENSHIP, in press). Known locally as an “ink-fish’ (WHITTEN, Ro- SENE & HEDGPETH, 1950) or “sea cow,” its first report from the Texas Coast is in all likelihood that by REED (1941). This species is an excellent swimmer. Occurrence: Although it is most abundant from late spring to August, it has been collected at least once in each month over a 2-year period. While this species is generally found to inhabit the bay, it has also been found washed ashore on the beach front, found attached to rocks of the jetties along the Brazos Santiago Pass, and reported from offshore reefs by TUNNELL & CHANEY (1970) along with A. morio (as A. floridensis). Aplysia dactylomela Rang, 1828 Synonymy: Aplysia protea Rane, 1828: 56 Aplysia schrammiui DESHAYES, 1857: 140 Aplysia guadeloupensis SOWERBY, 1869; plt. 5 (see EazEs, 1960: 307) Aplysia aequorea HEILPRIN, 1888: 325 Tethys dactylomela var. aequorea Heilprin. PirsBry, 1895: 77 Tethys panamensis Pirssry, 1895: 88 Aplysia megaptera VERRILL, 1899: 545 Remarks: Commonly known as the spotted or ringed sea hare, this species is distributed worldwide in warm seas. Only those synonyms with specific reference to the Gulf of Mexico or West Indies are given above. Refer to Eases (1960: 307) for a complete listing of worldwide synonyms. Occurrence: Uncommon, usually found exposed in grass beds at low tide only during late fall and winter. Occa- sionally found in small groups of 2 or 3, seldom in groups of up to 20. Vol20INo; 2 THE VELIGER Page 99 Aplysia morio Verrill, 1901 Synonymy: Tethys (Aplysia) morio VERRILL, 1901: 25 Tethys modesta THIELE, 1910: 124 Remarks: This is the common “sooty” sea hare reported as Aplysia floridensis by BREUER (1962), TUNNELL & Cuaney (1970), ANDREWS (1971), and Neck (1976). Living specimens are dark purple to black in coloration. This species, like Aplysia brasiliana, is an excellent swim- mer. Occurrence: Sporadically common during summer months of certain years as noted by Breuer (1962). During the course of this study it was noted to be com- mon only during late summer of 1974. Aplysia donca Marcus « Marcus, 1960 Remarks: This species was described from a single spe- cimen which was “rather small’ and “has probably not attained the maximum size of the species” (Marcus & Marcus, 1960). This species differs from Aplysia morio with respect to penis morphology, shell foramen and rad- ula formula (Marcus « Marcus, 1960: 253): Due to its similar coloration, possible individual variation of juvenile specimens, and unreliability of both shell foramen (EALES, 1960: 381) and radula formulae (Marcus, 1972: 848; STRENTH & BLANKENSHIP, in press) as taxo- nomic characters within the genus, a comparative exam- ination of a juvenile series of A. morio from Texas with the type of A. donca would appear desirable. Occurrence: Known only from the single type-speci- men collected at Mustang Island, Port Aransas, Texas. Bursatella Blainville, 1817 Bursatella leachu pleu (Rang, 1828) Synonymy: Aplysia pleii Rane, 1828: 70 Bursatella ple Rang. Gray, 1850: 98 Notarchus pleii, Rang. Morcu, 1863: 25 Notarchus (Aclesia) ple (Rang). THIELE, 1910: 124 Remarks: Known as the ragged sea hare, the form com- mon to the Gulf of Mexico is just one of a number of subspecies (EALES & ENGEL, 1935) exhibiting a worldwide circumtropical distribution. Occurrence: Although occasionally found throughout the year, this species appears most abundant during late fall and winter. Observations in South Texas appear simi- lar to those by Henry (1952) for this species in Florida. ACKNOWLEDGMENT Appreciation is extended to Dr. L. O. Sorensen for assistance in field studies as well as facilities support at the Pan American University Marine Laboratory on South Padre Island, Texas. Literature Cited AssotT, RoBerT 1 ucKER 1974. American seashells. Van Nostrand Reinhold Co., New York, 2nd ed.; 663 pp.; 24 color plts.; numerous figs. ANDREWS, JEAN 1971. Sea shells of the Texas coast. xvii+298 pp.; illust. BLAINVILLE, HENRI Martz Ducrotay DE 1817. Dictionnaire des Sciences Naturelles, 5, Suppl. Paris & Stras- bourg, p. 138 BREUER, JosEPH P 1962. An ecological survey of the Lower Laguna Madre of Texas, 1953 - 1959. Publ. Inst. Mar. Sci. Univ. Texas 8: 153 - 183 DesHayes, GERARD PAUL 1857. Note sur differents mollusques de la Guadeloupe, envoyés par M. Schramm. Journ. de Conchyl. 6: 137 - 143 EAvEs, NELLIE B, 1960. Revision of the world species of Aplysia (Gastropoda, Opistho- branchia). Bull. Brit. Mus. (Nat. Hist.), Zool. 5 (10): 267 - 404 (January 1960) Univ. Texas Press, Austin, Eaves, Nevuie B. & HENDRIK ENGEL 1935, The genus Bursatella de Blainville. London 21 (5): 279-303; 31 plts.; 10 text figs. Gray, Maria E. 1850. The figures of molluscous animals selected from various authors for the use of students. 4; 97 - 98 HEILPRIN, ANGELO 1886. A new species of Aplysia. Proc. Acad. Nat. Sci, Phila,; 364 1887. Exploration on the west coast of Florida, and in the Okeechobee Wilderness, with special reference to the geology and zoology of the Floridian Peninsula. Trans, Wagner Free Inst. Sci. 1: 1 - 134; 19 plts. (May 1887) 1888, Contributions to the natural history of the Bermuda Islands. Proc. Acad. Nat. Sci. Philadelphia 1888: 302 - 328; 3 pits. Henry, Laura M. 1952. Observations on the sea hare Bursatella leachit pleit Rang. Florida State Univ. Stud. no. 7: 8-14 Linnazus, CaROLUS 1767. Systema naturae per regna tria naturae ... reformata 1 [Regnum animale] (2): 533 - 1327 rentii Salvii) Marcus, EvELINE pu Bois ReyMonpD 1972. On the Anaspidea (Gastropoda: Opisthobranchia) of the warm waters of the western Atlantic. Bull. Mar. Sci. 22 (4): 841 - 874; 75 text figs. (December 1972) Marcus, Evetine pu Bois-ReyMOND & ErRNsT Marcus 1960. Some opisthobranchs from the northwestern Gulf of Mexico. Publ. Inst. Mar. Sci. Univ. Texas 6 (1959): 251 - 264 Morcxu, Otto AnprEeAs Lowson 1863. Contributions A la faune malacologique des Antilles danoises. Journ. Conchyl. 3e ser., 3 (11):21 - 43 (1 January 1863) Proc. Malacol. Soc. (13 July 1935) editio duodecima, Stockholm (Lau- Page 100 THE VELIGER Vol. 20; No. 2 Neck, Raymonp W. 1976. Recent records of sea hares (Gastropoda : Opisthobranchia) from South Texas. The Veliger 19 (1): 107 (1 July 1976) Orsicny, AtcipE DESSALINES D’ 1837. | Voyage dans Amérique Méridionale ... exécuté pandant les années 1826, . .. et 1833 ... 5; xliii + 1-758; 85 plts. [in Atlas] Paris (A. Bertrand) Pitspry, Henry AuGusTus 1895-1896. vol. 16 in Tryon’s Manual of Conchology. Philadelphia. pp. vii+262; 75 plts. Ranc, SANDER 1828. Histoire naturelle des Aplysiens, premiére famille de lordre des Tectibranches In: Férussac: Hist. nat. gen. et partic. des mollusques. Paris, 1827: 1-84; 24 plts. Reep, Crype T. 1941. Marine life in Texas waters. Anson Jones Press, Houston, 88 pages SoweErBy, GEorcE BRETTINGHAM 1869. Aplysia In: A. L. Reeve, Conchologia Iconica 17: plates only: 1-10 StrENTH, Nep E. & James E. BLANKENSHIP In press. On the valid name of the common Texas and Florida species of Aplysia (Gastropoda, Opisthobranchia) . Bull. Mar. Sci. 28 (2) THIELE, JOHANNES 1910. Molluskenfauna Westindiens. Zool. Jahrb. Suppl. 11: 109 to 132 TuUNNELL, JoHN W. Jr. & ALLAN H. CHANEY 1970. A checklist of the mollusks of Seven and One-half Fathom Reef, northwestern Gulf of Mexico. Contr. Mar. Sci. Univ. Texas 15: 193 - 203 VERRILL, ADDISON EMERY 1899. The nudibranchs and naked tectibranchs of the Bermudas. Trans. Conn. Acad. Arts Sci. for the years 1899-1900, 10, prt. 2: 545 - 550; pit. LXVI 1901. Additions to the fauna of the Bermudas, from the Yale Expedi- tion of 1901, with notes on other species. Trans. Conn. Acad. Arts Sci. 11: 15 - 62 Warten, H. L., H. EF Rosene & Jozt W. Hepopetu 1950. The invertebrate fauna of Texas coast jetties: A preliminary survey. Publ. Inst. Mar. Sci. Univ. Texas 1 (2): 53-87 Vol. 20; No. 2 THE VELIGER Page 101 Reevaluation and New Description of the Genus Bittium (Cerithiidae) RICHARD S. HOUBRICK Division of Mollusks, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560 (1 Plate) INTRODUCTION Tue cENus Bittium Gray, 1847 was proposed in manu- script form by Leach for a classification of British mollus- ca and was subsequently published by Gray (1847a). Leach’s list referred Bittium and many other diverse genera to the Purpuridae and under the 65" entry, listed Bittium reticulatum, Murex tuberculare, M. adversum, M. elegantissimum, and M. spenceri, consecutively. Be- sides Bittium reticulaium, the other species listed by Leach comprise 2 genera, Triphora Blainville, 1828 and Cerithi- opsis Forbes & Hanley, 1853. No description of the genus was given nor was a type-species designated. Three months later, Gray (1847b: 154) cited only Brttzwm reticulatum (Costa, 1778), which becomes the type-species by subsequent designation (Figure 1). To my knowledge, the earliest diagnosis of the genus is that written by H.s A. Apams (1854): “Operculigerous lobe with rudimentary expansions on each side and fur- nished with a roundish, lanceolate cirrhus (Lovén). Oper- culum subcircular, of four volutions. Shell turreted, many- whorled, granular, often with irregular varices; aperture with a slight canal in front, not produced or recurved; inner lip simple; outer lip acute, not reflexed or expand- ed.” Note that this definition mentions the presence of an anterior canal, a point to which I shall return later. Subsequent classic monographic treatments of the Cerithi- idae failed to elaborate on the genus and did not specify further characters. HISTORICAL REVIEW AND STATUS or GENUS Bittium There appears to have been some confusion among early workers about the exact placement and rank of Brttium. SowerBy (1855) included Bitttum in the introductory title of his monograph of Cerithium but did not cite it elsewhere in the text; moreover, he referred Bittium reti- culatum to his “Cerithiopsis group.” Reeve (1865) did not even mention the name Bitttum. Trron (1887) gave it generic recognition but offered no diagnosis. CHENU (1859) appears to have copied the definition of H. & A. ApamMs (1854) but included Bittzwm in the family Potam- ididae where it was assigned as a subgenus of Potamidines (sic) [= Potamides Brongniart, 1810], JEFFREYS (1867) lumped Bitttwm under Cerithium. Dati (1892) later wrote a diagnosis of the genus Brttium that was obviously formulated from his knowledge of Recent and fossil North American species. He mentioned that the last whorl is » usually contracted and overhung by its predecessor. This is typical in some individuals of American species, but there is a great deal of variation within specific popula- tions. Dat (of. cit.) also noted the close resemblance of Bitttum to the genera Diastoma Deshayes, 1850, Sand- bergeria Bosquet, 1861, Alaba Adams « Adams, 1853, and Styliferina A. Adams, 1860 [= Diala A. Adams, 1861] and added that the limits of these genera seemed to be quite artificial. An obvious feature of Brttiwm reticulatum, type-species of Bittiwm, is the presence of a short, shallow, but distinct Page 102 anterior canal. This was mentioned by H. « A. ADAMS (1854) in their description of the genus. The presence of an anterior canal is noted in most subsequent treatments of Bitttwm (CHENU, 1859; Tryon, 1887; Bucguoy, DauTZENBERG & Dotrus, 1884; FiscHER, 1877; DALL, 1892; CossMANN, 1906). Most modern authors also refer to a short anterior canal in their definitions of Bittzwm (Wooprinc, 1928; OLSSON & HARBISON, 1953; McLean, 1969; ABBoTT, 1974) although Keen (1971) regarded it as “ a slight anterior notch, not a canal.” Bittium species are frequently referred to other genera. Keen included Bittiwm in the subfamliy Cerithiopsinae but the radulae, larvae and anatomy of Cerithiopsis spe- cies are far-removed from those of Bittiwm and they should not be considered a closely related group (see FRETTER, 1951; FRETTER & PILKINGTON, 1970; JOHANS- SON, 1956). Recently, Asporr (1974) referred several American species of Bittiwm to the genus Diastoma Des- hayes, 1850. His placement of these species was followed by EMERSON & JACOBSON (1976). DALL (1889) was the first worker to confuse American members of the genus Bitttwm with Diastoma Deshayes, 1850 when he referred the common eastern species, Bittzwm varium Pfeiffer, 1840, to Diastoma. Dat (1889) noted that DESHAYES (1850) considered Diastoma to be a rissoid but the former regarded Diastoma as a subgroup of Bitttwm and added that Alaba Adams « Adams, 1853 was also very similar to Bittium. Although Diastoma was placed in the Cerithi- idae by FiscHEer (1884) and CossMANN (1889), it is now regarded as a separate family, Diastomatidae (see Lupsrook, 1971). The genus Diastoma embraces a group of relatively large snails and is best known from the Eo- cene of the Paris Basin. DEsHaAyEs (1850) designated Melania costellata Lamarck, 1804 as the type-species and later (1864, Description des Animaux san Vertébres, 2: 413-414) described it in detail. This species is large (about 40mm in length) and the lower portion of the outer lip is a smooth semicircle with no evidence of an anterior canal or even a shallow flared depression. Excel- lent illustrations of this type-species, cited as Melania costellata Lamarck, appear in Favre (1918: plt. 2, figs. 26-30). LupBrook (1971) has discussed the nomencla- tural problems of Diastoma in more detail and has pointed out that small cerithiids such as those described from Southern California as Diastoma fastigiata (Carpenter, 1864), D. oldroydae Bartsch, 1911, and D.stearns: Bartsch, IQI1, and an east coast species, D. virginica Henderson & Bartsch, 1914 [= Bittium varium Pfeiffer] differ from THE VELIGER Vol. 20; No. 2 true Diastoma by their small size, different sculpture and in the possession of a short, shallow anterior canal. The only living species of Diastoma, D. melanioides (Reeve, 1849) is found in a very restricted area in south-western Australia and off Eyre Peninsula in South Australia (Lup- BROOK, 1971). This moderately sized species was figured by REEveE (1849, 5: plt. 1, fig. 3) under his section on Mesalia Gray, 1840 and is very unlike the small Bittzum species of authors. I have been unable to examine a spe- cimen of Diastoma melanioides but CoTTon (1932) stated that it bears close relationship to fossil species of Diastoma even though the upper portion of the outer lip is not as far detached from the body whorl as is the lip in D. costellata. It is thus clear that the referral of Ameri- can species of Bitttum to Diastoma by Dati (1889), BarTscH (1911), KEEN (1971), ABBorT (1974) and EMERSON & JACOBSON (1976) should be considered er- roneous. The subgenus Bittinella Dall, 1924 was proposed to ac- commodate Bittium hiloense Pilsbry & Vanatta, 1908, from Hawaii. I suggest that this species and Cerithium zebrum Kiener, 1841, referred to Bitttum by CERNOHORS- Ky (1972), are merely very small members of the genus Cerithium Bruguiére. Both species have thick, heavy shells, short, deep anterior canals and thick, crenulated outer lips that are different from most other species of Bittium. GRANT & GALE (1931: 731) suggested that many sub- divisions of Bittiwm were probably artificial and should be discarded as soon as a natural classification was worked out. My studies on living populations of cerithiids and familiarity with numerous fossil groups and species have convinced me that the adaptive radiation of this large group has led to frequent convergence in shell form, sculp- ture and radular morphology. Most of the supraspecific taxa proposed for Bittitwm are parochial in conception and scope, are based on specific rather than generic char- acters and convey misleading or little phylogenetic infor- mation. In the interest of pragmatism and taxonomic par- simony I am suggesting that a number of generic and subgeneric taxa be abandoned or synonymized. All spe- cies formerly referred to these taxa should be considered as Bittiwm species until the entire group is monographed and subgeneric taxa can be properly evaluated on the basis of more than shell characters. In order to promote taxonomic stability and to clarify the problems discussed above I herein present a synonymy and redescription of the genus Bittium. Vol. 20; No. 2 THE VELIGER Page 103 SYNONYMY Cerithiinae Fleming, 1828 Bitttum Leach in Gray, 1847 Bitttum 1847 (Oct.), LEacH in Gray, Ann. Mag. Nat. Hist., 20: 270. Type-species, Murex reticulatus (Montagu, 1803) (= Strombiformis reticulatus Costa, 1778), by SD Gray, 1847 (Nov.), Proc. Zool. Soc. London part 15 (178): 154 Cerithiolum 1869, T1sErt, Boll. Malacol. Ital. 2: 263. Type- species, Cerithiolum (Strombiformis) reticulatum (Da Costa, 1778), by OD Semibittium 1869, Cossmann, Ann. Soc. malac. Belg., Mem. 31: 29. Type-species, Cerithium cancellatum Lamarck, 1804, by OD Bittiolum 1906, CossMANN, Essais Paléoconch. 7: 139. Type- species Bittium podagrinum Dall, 1892, by OD Stylidium 1907, Datu in Bartscu, Proc. U. S. Nat. Mus. 33 (1564) : 178. Type-species, Bittium (Stylidium) eschrichti (Middendorff, 1849), by OD Lirobittium 1911, BartscH, Proc. U. S. Nat. Mus. 40 (1826) : 384-385. Type-species, Bittium (Lirobittium) catalinensis Bartsch, 1907, by OD Monobittium 1917, MontTeERosaTo, Boll. Soc. Zool. Ital. (3) 4: 20. Type-species, Manobittium latreillei (Payraudeau, 1826), by M Inobittium 1917, MonteRosato, Boll. Soc. Zool. Ital. (3) 4: 20. Type-species, Inobittium lacteum (Philippi, 1836) by M Zebittium 1927, Fintay, Trans. New Zeal Inst. 57: 381. Type- species Zebittium exile (Hutton, 1873), by OD Cacozeliana 1928, StranD, Arch. Naturgesch. Berlin, 92: 66. nom. nov. per Cacozelia Iredale, 1924, Proc. Linn. Soc. N.S. W. 49: 246 [non Grote, 1878]. Type-species, Ceri- thium lacertina Gould, 1861, by OD Bittiscalia 1937, Fintay & Marwick, Paleo. Bull. New Zeal. 15: 44. Type-species, Bittiscalia simplex (Marshall, 1917), by OD Eubittium 1937, Cotton, S. Austral. Natural. 18: 2 [= Para- cerithium Cotton, 1932 (non CossMANN, 1902)]. Type- species, Eubzttium lawleyanum (Crosse, 1863), by OD Brachybittium 1962, WeIsBorp, Bull. Amer. Paleont. 17 (193): 175-176. Type-species, Bittitwm (Brachybittium) carabobo- ense, Weisbord, 1962, by OD Dahlakia 1971, Biccs, Journ. Conchol. 27: 221. Type-species, Dahlakia leilae Biggs, 1971 (= Cerithium proteum Jousseaume, 1930), by OD DESCRIPTION Shell Shell small, turreted and elongate, consisting of many moderately inflated or angular whorls (6-10). Sculpture normally reticulate, of varying combinations of spiral cords and axial riblets, frequently beaded at intersections. Usually 3 spiral cords per whorl. Former varices usually present at irregular intervals. Suture distinct, straight. Protoconch about 24 whorls, smooth, but sometimes with several spiral lirations. Frequently, teleoconch strongly reticulate. Aperture ovate with short, shallow, anterior siphonal canal that is not reflected backwards. Weak anal sinus present. Columella concave and with slight callus. Outer lip thin, smooth and rounded, occasionally flaired at the base on some individuals. Base of last whorl sculp- tured with 5-6 spiral cords. Last whorl occasionally con- tracted in some individuals. Periostracum thin. Animal Moderate sized foot with ciliated duct on right side terminating in knob-like ovipositor. Operculum thin, cor- neous, ovate and paucispiral with an excentric nucleus. Snout moderately large, flattened dorso-ventrally. Eyes at bases of tentacles. Mantle cavity deep with long mono- pectinate ctenidium extending its length, ending at ante- rior siphon. Osphradium bipectinate. Hypobranchial gland well-developed, large. Intestine wide, rectum distal to end of gonoduct. Feces cylindrical with mucus filament at one end. Pallial gonoducts in both sexes open, con- sisting of 2 laminae joined dorsally to mantle roof. Males aphallic, producing spermatophores about 1mm in length containing eupyrene and apyrene spermatozoa. Eupyrene spermatozoa with nucleus about one-twentieth the total length of sperm and with spiral keel on flagellum. Female pallial oviduct a glandular groove between 2 deep lami- nae joined at dorsal wall and open ventrally. In medial lamina, an open sperm-collecting gutter runs into a ciliated tube that opens into a “bursa” at the proximal end of the medial lamina. Bursa is anterior to 2 seminal receptacles, the 1‘ of which serves as a receptacle for the spermatophore, in the left wall of the duct and the 2" along the free edge of the wall. Nervous system character- ized by very large nerve ring and large cerebral ganglia. Jaws cuticular, conical and rough surfaced. Stomach with style-sac and large crystalline style. Radula Radula taenioglossate, ribbon long, about 50 rows. Ra- chidian tooth squarish, concave at top with one large central cusp flanked by 3 denticles on each side. Base of rachidian concave with 2 short lateral projections. Lateral tooth rhomboidal, serrated on top with 2 small entocones, large pointed mesocone and 4 small ectocones, respec- tively. Base of lateral tooth with 2 extensions, 1 at each side; outer basal projection broad, long and extending Page 104 laterally. Marginal teeth long, curved, spatulate and ser- rated at tips. Inner marginal tooth serrated with 4 ento- cones, large pointed mesocone and 3 sharp ectocones. Outer marginal tooth serrated with 4 to 6 entocones, outer surface smooth. Eggs and Larvae Eggs tiny, deposited in gelatinous string that may be coiled counterclockwise into a spiral from 3 to 25mm in length or irregularly folded on itself. Egg mass containing several hundred small, opaque eggs, each about 60-70 um in diameter. Larvae planktotrophic with bilobed, colorless velum; right velar lobe somewhat larger than left. Larval foot colorless. Larval shell about 24 whorls, smooth except for median spiral ridge on 2" whorl. Sutures and colu- mella reddish brown. Outer lip with prominent tongue- shaped beak. REMARKS The description given above is based upon Bittiwm reti- culatum, B. varium, and several other Birttiwm species from throughout the geographic range of the genus. I have attempted to describe the genus as a whole and not just the type-species. I have also endeavored to formulate a holomorphic description, using characters drawn from all phases of development. A thorough treatment of the anatomy of Bittium reti- culatum may be found in JoHANSSON (1947; 1948; 1953). Marcus & Marcus (1963) wrote an extensive paper on the anatomy of Bittzwm varium in which they also described and figured gametes, egg mass and larvae. The spermatozoa of B. reticulatum have been figured by FRANZEN (1955: 411; figs. 31-32) and FRETTFR & GrRa- HAM (1962: 339; fig. 176, 4). Marcus & Marcus (1963) indicated that the sperm of B. varium are very similar to those of B. reticulatum. For further information on the anatomy of Bittzwm and illustrations of internal anatomy see FRETTER & GRAHAM (op. cit.). Excellent illustrations and treatments of Bittcwm eggs and larvae may be found in THorson (1946), LeBour (1937), FRETTER & PILKING- TON (1970) and RicHTER & THoRSON (1975). The last paper contains good scanning electron micrographs of the larval shell of Buttium species and a photograph of a living veliger larva. The reproductive anatomy of Brttium reticulatum and B. varium, allopatric species that differ in shell size, shape and sculpture, shows that the structures of these conserva- tive tracts are virtually identical between the 2 species and indicates that they are indeed members of the same THE VELIGER Vol. 20; No. 2 genus. There is not a shred of anatomical evidence to support the need of a separate genus to accommodate Bit- ttwm varium nor is its referral to Diastoma justified. Al- though there is no information about the anatomy of Indo-Pacific and Eastern Pacific species of Bittiwm, it is not unlikely that they are all similar in their anatomy and habits. A small species from the Red Sea that was previ- ously placed in the Potamididae and for which a new genus was created, Dahlakia Biggs, 1971, has been found to be a true cerithiid and is now assigned to Bittiwm (Housrick, 1977b, in press). The family Cerithiidae is an old, slowly evolving group of prosobranchs (G. M. Davis, pers. comm.) that has not changed significantly since the Tertiary. Although some groups are now extinct, many living species of cerithiids have records going back to the Eocene (see Housrick, 1977). CoSSMANN (1906: 137) has recorded Bittium as far back as the Paleocene. Fossil Bittiwm species are conchologically very similar to Recent members. Bittium is a genus in the family Cerithiidae and is prob- ably very close in affinity to Cerithium Bruguiére, 1789. Familial or subfamilial status for the Bittiwm group is not warranted because there are no characters, beyond those of size and a shorter canal, to justify such a great taxonomic difference. Examination of living populations of Bittitwm in Florida, Belize, and Enewetak Atoll and anatomical studies reveal only minor differences between these animals and those of the genus Certthium Brugui- ere. Cerithiids are highly responsive to microhabitat in their shell shape and sculpture, and even anterior canals may be longer or shorter, depending upon the habitat. Inflation of whorls and spire length are highly correlated with different substrates such as sand, algae, rubble or rock. Populations of a given species frequently exhibit striking ecoclines. Shells of Bittizwm species are remarkably polymorphic in shape and sculpture, and sub- genera such as Lirobittium Bartsch, 1911, Stylidium Dall, 1907, and Semibitttum Cossmann, 1869 have no taxo- nomic value because the characters separating them (see BarTScH, 1911: 384) may be found within a single popu- lation of one species and are thus not even good criteria for specific determination. Brttiwm is a genus with a cos- mopolitan distribution of species and much of the con- fusion regarding generic and subgeneric placement of many species has resulted from parochial views of the genus. Bittiwm species have a cosmopolitan distribution and supraspecific taxa proposed solely for regional forms while ignoring other geographic areas and the full range of morphologies in the group are of little value (see Figures 1 to 5). In conclusion, subgeneric and generic taxa proposed for Bittium are based solely upon shell form and sculpture THE VELIGER, Vol. 20, Figure 7 No. 2 [Housricx] Figures 1 to 5 Figure 2 Figure 3 Figure 4 Bitttum species showing variation in shell form and sculpture as found throughout the genus Figure 1: Bittium reticulaum (Costa, 1778), type-species of the genus, from Cartagena, Spain (12X4mm) Figure 2: Bittium interfossa (Carpenter, 1864) from Catalina Is- land, California (7.4.X 2.5mm) Figure 3: Bittium quadrifilatum (Carpenter, 1864) from Mission Bay, California (10X5.2mm) Figure 4: Bittium varium (Pfeiffer, 1840) from Sarasota, Florida (5.8 X 1.9mm) Figure 5: Bittium varium (Pfeiffer, 1840) from Chincoteague, Virginia, formerly known as B. virginicum, Henderson & Bartsch, 1914 (9.9% 2.9mm) Figure 5 aa Vol. 20; No. 2 THE VELIGER Page 105 of regional species groups. In some cases, these sculptural characters are not of sufficient weight to justify specific differences and as a rule, polymorphism has been over- looked. Many of the taxa are provincial in scope and their authors either ignored or have been unaware of Bittiwm species in other parts of the world. In all cases, the criteria and concepts used in the formulation of these taxa ignore anatomy, eggs, larvae, early developmental stages, the fossil record and evolutionary convergence. For these reasons I think it is best to disregard the generic and sub- generic taxa that appear in the above synonymy and to regard all species previously referred to these taxa as Bittium species. ACKNOWLEDGMENTS I wish to thank Dr. Joseph Rosewater of the National Museum of Natural History, Smithsonian Institution, for critically reading this manuscript. I also thank Photo- graphic Services of the Smithsonian Institution for the photographs used in this paper. Literature Cited AssBotT, RoBeRT TUCKER 1974. American seashells Van Nostrand Reinhold Co., New York, 2nd ed.; 663 pp.; 24 color plts.; numerous figs. Apams, ARTHUR 1860. On some new genera and species of Mollusca from Japan. Ann. Mag. Nat. Hist. (3) 6: 331 - 337 (November 1860) 1861. On some new genera and species of Mollusca from the North of China and Japan. Ann. Mag. Nat. Hist. (3) 8: 135 - 142; 299 - 308 (September and October 1861) Apams, Henry & ArtHUR ADAMS 1853-1858. The genera of Recent Mollusca arranged according to their organization. 3 vols., illust. London. 1: xi+484 pp.; 2: 661 pp.; 3 (atlas): 138 plts. [collation, 2: 661] BartscHu, PAu 1907. in: Ralph Arnold. New and characteristic species of fossil mol- lusks from the oil-bearing Tertiary formations of Santa Barbara Coun- ty, California. Smithson. Misc. Coll. 50 (4): 419-447; plts. 50 - 58 (13 December 1907) 1911. The Recent and fossil mollusks of the genus Bittium from the West coast of America. Proc. U.S. Nat. Mus. 40 (1826): 383 - 414; pits. 51-58 (12 May 1911) Biccs, H. E. J. 1971. On a proposed new genus of cerithid Mollusca from the Dahlak Islands, Red Sea. Journ. Conchol. 27: 221 - 223; plt. 7 BLAINVILLE, HENRI Marie Ducroray DE 1828. Triphora. Dictionnaire des sciences naturelles: Paris, Le Normant; vol. 55, 344 pp. Bosguet, J. 1861. Notice sur le genre Sandbergia, genre nouveau de mollusques gastropodes de la famille des Cerithiopsidae. Mém. pour servir a la descrip. géolog. de la Néerlande 3: 53 - 57; 1 plt. (1861) BRONGNIART, ALEXANDRE 1810. Sur des terrains qui paroissent avoir été formés sous l’eau douce. Ann. Mus. Hist. Nat. Paris 15 (89-90): 357-405; plts. 22 - 23 BRuGUIERE, JEAN GUILLAUME 1789-1792. Encyclopédie Méthodique, Histoire naturelle des vers. Paris. 1(1): 1-344; 2 (2):345- 757 (1792) Bucguoy, E., Paut DautzenBerc & Gustave Doiirus 1884 (1882-1886). Les mollusques marins du Roussilon. 1 (5): 1-570, atlas 66 plts. Gastropodes. CarpenTER, Puitip PEARSALL 1865. Diagnoses of new forms of Mollusca, from the West coast of North America, first collected by Col. E. Jewett. Ann. Mag. Nat. Hist. (3) 15: 177-182 (March, 1865); 394-399 (May 1865) CrRNOHORSKy, WALTER OLIVER 1972. Marine shells of the Pacific, Vol. 2. 411 pp.; 68 plts.; 28 text figs. Pacif. Publ. Ltd., Sydney CHENU, JEAN CHARLES 1859. Manuel de conchyliologie et de paléontologie conchyliologique, Paris. vii + 508 pp.; 3707 figs. Cossmann, ALEXANDRE EpouarRD Maurice 1889. Catalogue illustré des coquilles fossiles de l’Eocéne des environs de Paris faisant fuite aux travaux Paléontologiques de G. P. Deshayes. Ann. Soc. Malac. Belgique 24: 7-385; 12 plts. 1896. Catalogue illustré des coquilles fossiles de l’Eocéne des environs de Paris faisant fuite aux travaux Paléontologiques de G. P Deshayes. Ann. Soc. Malac. Belgique 31: 1 - 94; 3 plts. 1902. Note sur l’Infraelias de la Vendée. in C. Chartron & M. Cossmann. Bull. Geol. Soc. France (4) 2: 163 - 203; plts. 3 - 4 1906. Essais de paléoconchologie comparée. Paris. vol. 7, 248 pp.; 14 plts. pa Costa, E. M. 1776. Elements of conchology: or an introduction to the knowledge of shells. London vii+vit+318 pp; 7 plts. Cotton, BERNARD CHARLES 1932. Notes on Australian Mollusca, with descriptions of new genera and new species. Rec. S. Austral. Mus. 4 (4): 537-547; 9 figs. (31 July 1932) 1937. Nomenclatural note. So. Austral. Natural. 18 (1,2): 2 Crosse, H. 1863. Description d’espéces nouvelles d’Australie. Journ. de Conchyl. 11: 84-90 DALL, WILLIAM HEALEY 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Carib- bean Sea (1879-80), by the U.S. Coast Survey steamer “Blake,” (etc.) XXIX. Report on the Mollusca—Part II. Gastropoda and Scapho- poda. Bull. Mus. Comp. Zool. Harvard 18: 1-492; plts. 1-40 (January to June 1889) 1892. Contributions to the Tertiary fauna of Florida, with especial ref erence to the Miocene silex-beds of Tampa and the Pliocene beds of the Caloosahatchie River. Tertiary mollusks of Florida. Part II. Streptodont and other gastropods, concluded. Trans. Wagner Free Inst. Sci. Philadelphia 3 (2): 201-473; plts. 13-22; 1map (December 1892) 1907. New marine mollusks from the West coast of America. Proc. U.S. Nat. Mus. 33 (1564): 177 - 183 (23 October 1907) 1924. Notes on molluscan nomenclature. Proc. Biol. Soc. Wash- ington 37: 87 - 90 (21 February 1924) DesuHayes, GERARD PAUL 1850. Traité élémentaire de conchyliologie. Paris. vol. 1 (2): 129 to 824 1864. Descriptions des animaux sans vertébres. Paris. vol. 2, text: Mollusques, Acéphalés monomyaires et Brachiopodes. Mollusques Céphalés, premiére partie, 968 pp.; atlas, 64 pllts. Emerson, WILLIAM KeitH & Morris KARLMANN JACOBSON 1976. The American Museum of Natural History guide to shells, land, freshwater, and marine, from Nova Scotia to Florida. xvilit+ 482 pp.; 47 plts. New York, A. A. Knopf Inc. Favre, JULES 1918. Catalogue illustré de la collection Lamarck, premiére partie. Mollusques fossiles. Geneva. 21 plts. Fintay, Harotp JoHN 1927. Additions to the Recent molluscan fauna of New Zealand, No. 2. Trans. New Zeal. Inst. 57: 485 - 487; plts. 24-26 (19 Jan. ’27) Fin.tay, Harotp JoHN & J. Marwick 1937. The Wangaloan and associated Mollusca faunas of Kaitangata- Green Island Subdivision. New Zeal. Geol. Surv., Paleont. Bull. No. 15: 1- 140; 17 plts. FiscHer, PAuL 1887 (1880-1887). Manuel de conchyliologie et de paléontologie con- chyliologique. Paris. xxiv+1369 pp.; 23 plts. FLEMING, JOHN 1828. The philosophy of zoology or a general view of the structure, functions and classifications of animals, etc. Edinburgh, 2 vols. Forses, Epwarp & SYLVANUS HaNLEY 1853. A history of the British Mollusca and their shells. x + 616 pp. vol. 3: Page 106 FRANZEN, AKE 1956. Comparative morphological investigation into spermiogenesis among Mollusca. Zool. Bidr. Uppsala 30: 399-456; 2 plts. 146 text figs. FRETTER, VERA 1951. Observations on the life history and functional morphology of Cerithiopsis tubercularis (Montagu) and Triphora perversa (L.). Journ. Mar. Biol. Assoc. U. K. 29: 567 - 586 FrETTER, VERA & ALASTAIR GRAHAM 1962. British prosobranch molluscs, their functional anatomy and eco- logy. London, Ray Soc. xvi+755 pp.; 316 figs. FretTTerR, VERA & MarcareT C. PILKINGTON 1970. Prosobranchia. Veliger larvae of Taenioglossa and Stenoglossa. Conseil Internat. Explor. mer. Fich. Ident. Zooplank., sheets 129 - 132; 2-26; 35 figs. Gou.p, Augustus ADDISON 1861. Descriptions of shells collected in the North Pacific Exploring Expedition under Captains Ringold and Rodgers. Proc. Boston Soc. Nat. Hist. 7: 385 - 389 (January 1861) Grant, Utysses Simpson, IV « Hoyt Ropney GALE 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and adjacent regions. San Diego Soc. Nat. Hist. Mem. 1; 1- 1036; 15 text figs.; plts. 1 - 32 (3 November 1931) Gray, Joun Epwarp 1840. Synopsis of the contents of the British Museum mollusks. London (expanded 42nd Ed.) pp. 86-89; 106- 156 1847a. The classification of the British Mollusca by N. E. Leach, M.D. Ann. Mag. Nat. Hist. 20: 267 - 273 (September 1847) 1847b. A list of the genera of Recent Mollusca, their synonyma and types. Proc, Zool. Soc. London (for 1847) 15 [part 15] (178): 129 - 219 (November 1847) Grote, A. R. 1878. On the pyralid genus Epiaschia of Clemens, and allied forms. Proc. Boston Soc. Nat. Hist. 19 (1): 262 - 265 (5 December 1877) HENDERSON, JoHN B., Jr. & Paut BarTscH 1914. Littoral marine mollusks of Chincoteague Island, Virginia. Proc. U.S. Nat. Mus. 47 (2055): 411-421; plts. 13-14 (29 Oct. ’14) Housrick, RicHArp STEPHEN 1977a. The family Cerithiidae in the Indo-Pacific, Part 1. The genera Rhinoclavis, Pseudovertagus, Longicerithium and Clavocerithium. Indo-Pacific Mollusca (in press) 1977b. Redescription of Bittium proteum (Jousseaume, 1930) with comments on its generic placement. The Nautilus (in press) Hutton, FW. 1873. Catalogue of the marine Mollusca of New Zealand. Colon. Mus. and Geol. Surv. Dept. 1873: xv+116 pp.; 1 plt. IREDALE, Tom 1924. Results from Roy Bell’s molluscan collections. N.S. W. 49 (3): 179-278; plts. 33 - 36 JEFFREYS, JOHN Gwyn 1867 (1862-1869). British Conchology. JOHANNSON, JOHAN Proc. Linn, Soc. London yol., 4, 487 pp.; 8 plts. 1947, Uber den offenen Uterus bei einigen Monotocardiern ohne Kopulationsorgan. Zool. Bidr. Uppsala 25: 102-110 1953. On the genital organs of some mesogastropods: Cerithium vul- gatum Brug., Triphora perversa (L.) and Melanella (Eulima) inter- media (Cantr.). Contributions to the phylogeny of the pallial gono- ducts of the Prosobranchia. Zool. Bidr. Uppsala 30: 1-23; 1 plit.; 10 figs. 1956. On the anatomy of Tympanotonus fuscatus (L.), including a survey of open pallial oviducts of the Cerithiacea. Atlantidae Rept. No. 4: 149-166; 23 figs.; 1 plt. JoussEAumME, FEéLtx PIERRE 1930. Cerithiidae de la Mer Rouge. 270 - 296; 3 figs. Keen, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. Ed. 2. Stanford Univ. Press, Stanford, Calif. i- xiv +1064 pp.; ca. 4000 figs.; 22 color plts. (21 September 1971) Kiener, Louis Cuaries 1841-1842. Spécies général et iconographie des coquilles vivantes. Genre Cérite. Paris, vol. 5, 104 pp.; 32 plts. Lamarck, Jean-Baptiste PieRRE ANTOINE DE MONET DE 1804. Suite des mémoires sur les fossiles des environs de Paris. Ann. Mus. Hist. Nat. 3: 436 - 441 Journ. de Conchyl. 74: THE VELIGER Vol. 20; No. 2 Lesour, Marie V. 1937. The eggs and larvae of the British prosobranchs with special reference to those living in the plankton. Journ. Mar. Biol. Assoc. U.K. 22: 105 - 166; 4 figs. Lupproox, Netty Hooper 1971. Large gastropods of the families Diastomatidae and Cerithiidae (Mollusca: Gastropoda) in southern Australia. Trans. Roy. Soc. South. Austral. 95 (1): 29-42; 6 plts.; 1 fig. McLgan, JAMEs HamILToNn 1969. Marine shells of Southern California. Los Angeles County Mus. Nat. Hist., Sci. Ser. 24, Zool. 11: 104 pp.; illust. (October 1969) Marcus, ErnsT & EvELINE DU Bois-REYMOND Marcus 1963. Mesogastropoden von der Kiiste von Sao Paulo. Abhandl. Math.-Naturwissensch. Klasse Akad. Wiss. & Lit. Mainz 1: 1-105; 86 figs. MarsHALt, WILLIAM B. 1917. The Wangaloa beds. 49: 450 - 460; plts. 34-37 MIppDENDORFF, ALEXANDER THEODOR VON 1849, Die Meeresmollusken Russlands in ihren Beziehungen zur zoo- logischen und physikalischen Geographie, 8, 1849. Bull. Acad. Sci. St. Petersburg 3 (5): 1-21 MontTacu, GEORGE 1803. Testacea Britannica; or natural history of British seashells, ma- rine, land, and fresh water, parts 1 and 2, 1803. London, i-xxxvii+ 1-606 pp.; plts. 1-30 (Suppl. 1810) MonTEROSATO, MARCHESE 1917. Molluschi viventi e quaternarii raccolti lungo le coste della Tripolitania. Boll. Soc. Zool. Ital. (3) 4: 1-28; 2 plts Ousson, Axe, ADoLPH & ANNE HARBISON 1953. Pliocene Mollusca of southern Florida with special reference to those from North St. Petersburg ... with special chapters on Turridae by William G. Fargo and Vitrinellidae and freshwater mollusks by Henry A. Pilsbry. Monogr. 8, Acad. Nat. Sci. Philadelphia, pp. iv-+1-457; plts. 1-65 (6 November 1953) PAYRAUDEAU, B. C. : 1826. Catalogue descriptif et méthodique des annelides et mollusques de I’lle de Corse, Acad. Sci. séance 12 Mars 1827, Paris 1826, 218 pp. PFEIFFER, L, 1840. Ubersicht der im Januar, Februar und Marz, 1839 auf Cuba gesammelten Mollusken. Arch, Naturgesch. 6 (1): 250-261 Puitiprl, RupoLF AMANDUS ohio he 1836, Enumeratio molluscorum Siciliae. vol. 1, Berolini, 1836, i- xiv +268 pp.; 12 plts. Pirsspry, HENry AucustTus & EDWARD Guirey VANATTA 1908. Descriptions of new Hawaiian marine shells. 22 (6): 56-58; 3 figs. REEVE, LovELL AUGUSTUS ‘ 1849. Conchologica Iconica, London. vol. 5, the genera Mesalia and Eglisia. plt. 1 (June 1849) 1865. Conchologica Iconica. London. vol. 15, the genus Cerithium. 20 pits. + index (December 1865) RicHTER, GoTTHARD & GUNNAR THORSON 1975. Pelagische Prosobranchier-Larven des Golfes von Neapel. Ophelia 13: 109 - 185; 20 plts. (March 1975) SowERBY, GEORGE BRETTINGHAM Trans. New Zeal. Inst. Wellington The Nautilus (12 October 1908) 1855. Thesaurus Conchyliorum, or monographs of genera of shells. London, vol. 2, Cerithium, pp. 847 - 899; plts. 176-186 +plt. 12 (supplementary) STRAND, EmBRICK ; 1928. Miscellania nomenclatorica zoologica et palaeontologica. Arch. Naturgesch. Berlin 92: 30-75 Tuorson, GUNNAR ; : 1946. Reproduction and larval development of Danish marine bottom invertebrates with special reference to the planktonic larvae in the Sound (@resund). Medd. Kom. Danm. Fisk. Hav. Plankt. 4: 1-523; illust. Tryon, GzEorcE WASHINGTON, Jr. 1887. Manual of Conchology, ser. 1, 9: 1 - 488; 71 pits. Acad. Nat. Sci. WEIsBorD, NorMAN EDWARD 1962. Late Cenozoic gastropods from northern Venezuela. Bull. Amer. Paleo. 42 (193): 1-672; plts. 1-48 (5 March 1962) Wooprinc, WENDELL PHILLIPS A 1928. Miocene Mollusca from Bowden, Jamaica, Part 2, gastropods and discussion of results. Carnegie Inst. TS oan Philadelphia, Vol. 20; No. 2 THE VELIGER Page 107 The Chromodoridinae Nudibranchs from the Pacific Coast of America. - Part I. Investigative Methods and Supra-Specific Taxonomy HANS BERTSCH '! Donner Laboratory and Department of Zoology, University of California, Berkeley, California 94720 (3 Text figures) INTRODUCTION CHROMODORID NUDIBRANCHS are a worldwide tropical group, noted for their distinctive and beautiful color pat- terns. The group has been studied extensively, with nu- merous papers in part dealing with members of the Chromodorididae. This contribution is the first in-depth treatment of all the known species of the Chromodoridi- nae from the Pacific coast of America. Previous studies of this group from American Pacific waters have been based on a very small number of dissected specimens or only a few species (e.g., MacFarLanp, 1966; Marcus & Mar- cus, 1967; Bertscu, 1970). Modern reviews of the Chro- modorididae from other geographic provinces (Indo-Pacif- ic: RUDMAN, 1973; Australia: THOMPSON, 1972; Hawaii: Kay & Youne, 1969) are also based upon small numbers of dissected specimens of each species. By obtaining specimens from several major museums, and making many collecting trips to Mexico and Panama, I have obtained a large quantity of specimens, which I dissected and examined critically. Visual and numerical data obtained from these dissections were used to revise systematically the genera and species of the American Pacific coast Chromodoridinae. Principles governing the use of the radula in opistho- branch systematics have been proposed by Bertrscu (1976 a). This study applies Bertsch’s methods of exam- ining opisthobranch radular variation (multiple meristic counts and measurements, ontogenetic patterns, and mor- phological studies using light microscopy and scanning * Present address: Department of Biology, Chaminade University of Honolulu, 3140 Waialae Avenue, Honolulu, Hawaii 96816 electron microscopy) to a large group of phylogenetically related nudibranchs. Radular morphology and meristic qualities, and the external coloration are diagnostic char- acters for each of the 15 species of the Chromodoridinae from the Pacific coast of America. The following abbreviations refer to the collections from which nudibranch specimens were obtained: USNM - United States National Museum of Natural History (Smithsonian Institution), Washington, CAS - California Academy of Sciences, San Fran- cisco, California LACM - Los Angeles County Museum of Natural Histo- ry, Los Angeles, California HB — Hans Bertsch MATERIALS anp METHODS Scanning Electron Microscopy In the past few years, the scanning electron microscope has become a significant source of visual data for mala- cologists studying the ultrastructure, taxonomy, and func- tional morphology of various mollusks. It has been used to depict veligers (ROBERTSON, 1971b), adult shells (e. g., SoLEM, 1970; THomas & BINGHAM, 1972; MARGOLIS & CaRVER, 1974), chiton girdle scales (FERREIRA, 1974, 1976), body soft parts (ARcADI « HopcKINn, 1973; Mur- RAY & LEWIS, 1974), spermatozoa (MAXWELL, 1975), fossil eggs (Tompa, 1976), a fossil radula (SoLEM & RicHARDSON, 1975), and radulae of living mollusks (e. g., SOLEM, 1972). Page 108 Because of the SEM’s clear resolution in the 300-10 000 magnification range and its 3-dimensional imaging of biological samples, it is ideal for studying the total mor- phology of opisthobranch radulae, the details of individu- al teeth, and interrelationships between movable parts. THompson & Hinton (1968) published the first scanning electron micrographs of opisthobranch radulae. BertscH (1970) first used the SEM to illustrate the radula in a new species description of an opisthobranch. Since then, stereo pair micrographs have been published (BerTscu et al., 1973), other workers have begun using SEM to illustrate opisthobranch radulae (e.g., MULLINER & SpHON, 1974; BoucHET, 1975), and scanning electron micrographs have become an established method for pre- senting information about opisthobranch radulae. The principles of operation of the SEM and its bio- logical applications are extensively documented. An entry to this literature can be gained by consulting the papers by Haves (1972, 1973) and EVERHART & Hayes (1972). The recent increase of published scanning electron micrographs necessitates a few precautionary words about their interpretation. Certain artifacts (avoidable or not) are inherent in the preparation and viewing processes of SEM. Criark & GLacov (1976) have discussed 3 types of artifacts: accretion of extraneous materials, distortion of real cell and tissue surface details during processing, and distortion during viewing (including “charging,” cf. Paw- LEY, 1972). There are 2 additional artifacts that can occur by un- critical examination of SEMs: the “2-headed cow” effect (T. L. Hayes, personal communication), and the “other side of the coin.” The first artifact involves the interpre- tation of different planes that are visible in a micrograph. People can easily separate objects seen in the everyday macroscopic world as occurring at different depths in our field of vision (we know that the 2-headed cow is actually 2 cows standing together in opposite directions), but the same phenomenon in the microscope may pass unrecog- nized and be described as an ultrastructural unit. It is the advantage of the SEM — providing great depth of field with high resolution — which causes this possibility of planar reversibility. Analysis of overlapping and shad- ing on the micrograph will often eliminate this problem, but sometimes it can be resolved only by stereopair or by rotating the specimen go° so that the planes in question are aligned parallel, rather than perpendicular, to our field of vision. The second artifact also stems from the imaging proper- ties of the SEM. Image formation occurs by a time se- quencing of points on the specimen surface with points on the cathode ray tube. This contrasts with the light and transmission electron microscopes’ transparent technique o THE VELIGER Vol. 20; No. 2 of point-to-point spatial correspondence between the spe- cimen and its image. Because the SEM operates by tem- poral sequencing across a non-transparent object, the re- searcher does not know what is on the obverse side of the object being examined. The observer is looking at rather rather than through the specimen. Prevention of unduly assuming bilaterality requires tilting or rotating the spec- imen, reorienting the specimen on the mounting plat- form, or (if the details are large and the specimen thin enough) viewing the specimen with a light microscope. Specimen Preparation In this study, I dissected and examined over 270 radulae. By using such a large sample size, by comparing light and scanning electron microscopy, by taking stereo-pair micro- graphs, and by frequent tilting and rotation of puzzling specimens in the SEM chamber, descriptions of the radu- lar teeth were carried through a rigorous checking to insure their accuracy and usefulness to other researchers. Radular preparation involved dissecting the buccal mass, dissolving the tissues surrounding the radula with a cold bleach (sodium hypochlorite) solution (cf. Linp- BERG, 1977), and then allowing the radula to rinse in a 70% ethanol solution. For light microscopy, the cleaned radula was placed flat on a standard microscope slide, allowed to air dry, and then sealed under a cover slip with Permount mounting fluid. Radulae that were to be examined with SEM were dis- sected and cleaned in the same way. After rinsing, each radula was mounted on a petrographic slide without a cover slip, air dried, and then covered with a thin layer of gold by vacuum evaporation to improve specimen conduction and reduce charging. The radular ribbon was bent or torn in places so that lateral views of the teeth could be obtained from the SEM. Any shrinkage or bend- ing caused by the air drying did not significantly distort the rigid teeth shapes; moreover, it often allowed lateral and basal examination of the teeth. Specimens were viewed with a Japan Electron Optics Laboratory Co., Ltd., Scanning Type Electron Microscope Model JSM, at either 10 or 25 kV accelerating voltage. After examina- tion with the SEM, each radula was covered with a cover slip and observed with a light microscope. The gold layer did not interfere with the transparency of the radula to visible light, but enhanced the resolution by acting as a surface stain. Mounting the radula on the smaller-sized petrographic slide permits successive viewing of the specimen by SEM and light microscopy. The petrographic slide is small enough to fit on the specimen holder inside the SEM vacu- um chamber, and is transparent to light. Similar tech- Vol. 20; No. 2 niques for comparative imaging of the same specimen have been used previously by McDonatp & Hayes (1969) and BarBER (1972). Numerical Analysis of Radular Variation Statistical studies of dentition form an important corpus in the understanding of mammalian taxonomy and verte- brate community structure. Examining large numbers of mammal specimens, authors of new species and revisers have given particular attention to numbers of teeth, means of sizes, and other measurements and counts (the litera- ture is extensive, and only a few citations are necessary; cf. MoosER & DaALQueEst, 1975, or VILLA-R., 1966). Re- cent papers by Grant (1967), RouGHGARDEN (1974), and Tamsitr (1967) have estimated trophic relation- ships of vertebrate communities by analysis of the com- parative morphology of trophic structures (bird bills, liz- ard jaws, and bat jaws). FINDLEY (1976) included mea- surements of feeding structures in his investigation of the phenetic structure of tropical and temperate bat commu- nities. By contrast, statistical treatment of molluscan teeth has been relied upon much less. Although knowledge of the radula has advanced considerably since Aristotle’s brief descriptions of the cephalopod radula (“a minute organ of a fleshy nature, and this it uses as a tongue, for no other tongue does it possess” - Ross, 1952: 50) and that of the “spiral-shaped testaceans” (‘““Some of these creatures have a mouth and teeth, as the snail; teeth sharp, and small, and delicate” - Ross, op. cit.: 55), most treatments of the radula have emphasized just mor- phology, and not ontogenetic changes nor inter- and intra- specific size variation. TROSCHEL’s works (1856-1863, 1866-1893) are a not- able exception, since he gave the sizes of most of his ulustrated radulae and even indicated that larger speci- mens can have correspondingly greater-sized teeth (“Die Breite der Mittelplatte des abgebildeten Exemplars ist 0.085 Millim.; an einem grosseren Exemplare messe ich 0.11 Millim.” - vol. I: 111). In recent years, a number of articles have appeared discussing mathematical variations in radular teeth. Of special note is a series of papers by SCHILDER & SCHILDER (1961 a, 1961 b, 1963) and ScuILpER, ScHILDER & Hous- TON (1964), in which numerous cowrie radulae are ex- amined by regression analyses and mean sizes to deter- mine sexual dimorphism and correlations between shell and radular sizes. ROBERTSON (1971 a), reviewing reports of sexually dimorphic radulae among the Muricidae and Buccinidae, discusses meristic and morphological sexual dimorphism and population differences in the radulae THE VELIGER Page 109 of a phasianellid; he speculates on the possible evolution- ary significance of the dimorphism and illustrates the male and female radulae with scanning electron micro- graphs. BANDEL (1974) and BorKowskI (1975) studied the variability of Caribbean Littorinidae radulae. Finally, Broom (1976) correlated radular tooth curvature (a- mount of concavity of the inner [sic] margin of the tooth) with sponge prey items of dorid nudibranchs. Variation in size and number of opisthobranch radular rows and teeth has been considered taxonomically impor- tant for over a century. The use of these characters has been primarily typological ( in the sense of DoBZHANSKy, 1970: 126, and Mayr, 1970: 3-5), resulting in the de- scriptions of intraspecific variations as new species. Some workers have recognized intraspecific and ontogenetic var- iation: e.g., ENGEL & HUMMELINCK (1936: 38) state that the number of teeth per half-row in Dolabrifera do- labrifera (Rang,1828) increases with the size of the ani- mal, and Gonor (1961:91) records a similar observation for the tooth rows of A plysiopsis smithi (Marcus, 1961). Regression analyses of radular size and numerical vari- ables have been applied only recently to opisthobranchs. BEEMAN (1963) graphed the number of lateral teeth vs. total body length of Phyllaplysia taylori Dall, 1900, and THOMPSON (1958) graphed the estimated age in months vs. number of rows and size of the lateral teeth of Adala- ria proxima (Alder « Hancock, 1854) ; neither calculated the regression lines of his plots, but a visual examination of their graphs indicates that their correlations probably are Statistically significant. The first use of regression analysis formulae to prove statistically that the radular sizes and numerical varia- tions of certain opisthobranch species are ontogenetic dif ferences in size was by BertTscH (1976a). The method has also been applied by FERREIRA (1977) to Tropha maculata MacFarland, 1905, and T: catalinae (Cooper, 1863), and by BertscuH « Meyer (in prep.) to Disco- doris heathi MacFarland, 1905, and D. mavis Marcus & Marcus, 1967. These studies have shown that for certain opisthobranch groups, the size of the radula, and the num- bers of rows of teeth and teeth per half-row increase with the age (size) of the animal. BErTscH (1976 a) predicts within which opisthobranch taxa these variations will occur, and gives suggestions to the use of statistical ana- lyses. In this work, regression analyses are used in conjunction with light and scanning electron microscopy and examina- tion of whole animal morphology to elucidate the taxo- nomic relationships of a large assemblage of species. Such a multidimensional approach bases the choice of taxo- nomically important characters on significance and reli- ability. For instance, dependence on color patterns is usu- Page 110 ally reliable at the species level for chromodorid nudi- branchs, but one must take into account changes in this pattern between juveniles and adults (HAEFELFINGER, 1959; Ros, 1974: plt. 2) and adult variation. Grouping species on the basis of coloration (RUDMAN, 1973) is useful for comparative purposes, but does not indicate generic relationships. It is necessary to use multiple para- meters for taxonomic evaluations, so that the animals are viewed as biological, genetic, and ecological entities sub- ject to natural variation. In this study, all measurements and calculations were made at least twice to improve their accuracy. The rows of teeth were counted on both halves of the radula, and the larger number was used for the calculations. Several rows of teeth were counted on each radula, to determine the maximum number of teeth per half-row. All counts of number of teeth per halfrow indicate the maximum number found in each radula. Variations in the number of teeth per half-row exist within a radula because of tooth growth posteriorly and tooth wear anteriorly (BERTScH, 1976 a: 119; table 2; fig. 5). Using the maximum number found in the tooth rows eliminates the ontogenetic biases. Length and width maxima were measured on flattened, slide-mounted radulae. The formulae for means, standard deviations, regression analyses, confidence intervals and tests of regression coefficients are presented in Stimpson, Roe « LEwontTIn (1960: 84, 213-230, 238-241) ; tables of significance probabilities are in the same book (pp. 422, 426). Radular Teeth Definitions of Structural Terms Descriptions of radulae have included a wide variety of terms with inconsistent usage. To help standardize radular descriptions, Figure 1 illustrates the morphological terms used in this paper for the various parts of a chromodorid radular tooth. The base (Figure 1, a) of the tooth joins the radular ribbon; the functional, rasping part grows upward as the erect shaft (Figure 1, b). The shaft curves postero-dorsal- ly, and terminates in a point. Careful discrimination of the different sides of the teeth requires orienting the teeth in the resting stage when the radula is not extended. Along the cephalo-caudal axis of the animal’s body, the forward, convex surface of the shaft is anterior. The concave side behind the shaft is posterior (termed “inner margin” by Boom, 1976). The lateral, left and right, sides of the shaft are more properly THE VELIGER Vol. 20; No. 2 posterior > posterior > a Zon 1B 1A ya ——e posterior —> posterior > mG Figure 1 1D Illustration of structural parts of Chromodoridinae radular teeth A. Unicuspid tooth with posterior denticulation; a — base; b — erect shaft; c — cusp B. Unicuspid tooth with lateral denticulation; ¢ — cusp C. Acuspid tooth (after BERTScH, 1976b) D. Bicuspid tooth; d — main (or primary) cusp; e — secondary cusp designated inner and outer faces. The inner face is closest to the rachidian or mid-line of the radula. Denticles usually occur on the posterior (Figure 1 A) or lateral (Figure 1 B) surfaces of the teeth. When the denticles are significantly smaller than the point on the erect shaft, the point is termed a cusp (Figure 1, c). A unicuspid tooth terminates in one large cusp (Figures 1 A, 1B), and a bicuspid tooth terminates in 2 cusps larger than the denticles (Figure 1 D). The most distal cusp of a bicuspid tooth is termed the main (or primary) cusp (Figure 1, d), and the more proximal cusp (and also closer to the denticles) is the secondary cusp (Figure 1, e). The shaft of an acuspid tooth (Figure 1 C) termi- Vol. 20; No. 2 nates in a point smaller than, or of equal size with, the succeeding denticles, and is also called a denticle. Teeth with cusps have a serrate (posterior or lateral) denticu- Jation (Figures 1 A, 1 B, 1 D) and acuspid teeth exhibit pectinate denticulation (Figure 1 C). TAXONOMIC COVERAGE Supra-Generic Taxa GASTROPODA Subclass Opisthobranchia Superorder Nudibranchia HOLOHEPATICA Doridoidea CHROMODORIDIDAE A family of cryptobranch dorid nudibranchs; usually with bright colors and smooth dorsal skin texture (at times the dorsum may have small, low, round tubercles), with la- mellate rhinophores and gills surrounding anus. Recep- taculum seminis attached to vagina or bursa copulatrix. Radular teeth numerous; erect shaft thicker along antero- posterior axis than laterally; denticles usually present; shaft of tooth may end distally as unicuspid, bicuspid, or acuspid. Cadlininae Characters of the family, but usually colored cream or yellow, and dorsal skin surface covered with small, low, round tubercles, or smooth. Rachidian tooth well devel- oped, having a solid erect shaft with pronounced denticu- lation laterally; lateral teeth unicuspid. Ecologically are predominantly members of temperate, cooler water fau- nal provinces. Chromodoridinae Characters of the family; almost always brightly colored (when seen in laboratory aquaria, intertidally, or at greater depths with artificial lighting). Rachidian tooth not well developed; erect shaft of a small triangular shape, THE VELIGER Page 111 or recurved, thin and narrow; without prominent lateral denticulation. Erect shafts of lateral teeth unicuspid, bi- cuspid, or acuspid; with lateral or posterior denticles or occasionally smooth. The familial and subfamilial divisions of the crypto- branch dorid nudibranchs have not been agreed upon consistently in recent literature. ODHNER (in FRANC, 1968: 866-867) gives full family status to Echinochilidae (with 2 subfamilies, Echinochilinae and Lissodoridinae) and Chromodorididae (Echinochilidae is based on the reject- ed genus Echinochila Morch, 1869; the proper name is Cadlinidae; cf ICZN Opinion 812, 1967). Ros (1975: 320-321) uses Cadlinidae and Glossodorididae. Kay & Younc (1969) and Marcus « Marcus (1967) treat Chromodoridinae as a subfamily of Dorididae. The Mar- cuses (op. cit.) discuss Cadlina Bergh, 1878b, but do not separate the group on a supra-generic level. SCHMEKEL (1972) gives familial status to Chromodorididae and Dorididae. Lance (in KEEN, 1971: 821-823) recognizes Chromodoridinae and Cadlininae as subfamilies of Chro- modorididae (but then erroneously places the Discodorid- inae and Inudinae within this same family). The most natural grouping is to separate Dorididae from Chromodorididae at the family level, and retain Chromodoridinae and Cadlininae. There is a suite of characteristics proper to each family and shared by the respective genera and species: a) Body Form and Texture. The Chromodoridinae all have a smooth dorsal surface; some of the Cadlininae show a slightly more rugose texture, with low, rounded tubercles, but members of this group (e. g., Cadlina eve- linae Marcus, 1958) have a smooth dorsum. The Dorid- idae tend to have much larger tubercles, and a rougher texture of the dorsum. This difference is not absolute, but in general is a fairly reliable external separation between the families. It is the least important phylogenetic char- acteristic because some Chromodorididae show a rougher body texture and because smooth body texture occurs in a number of unrelated dorid forms: e. g., Aphelodoris antillensis Bergh, 1879b, and Dendrodoris krebsit (Morch, 1863). b) Reproductive System. The triaulic cryptobranch dorid reproductive system exhibits little functional and morphological variation (GHISELIN, 1966: 348-349). However, despite this overall homogeneity, the structural arrangement of parts of the female reproductive organs shows 2 patterns, a chromodorid arrangement and a dorid arrangement. These are fairly significant variations; they are not a loss of parts nor a relative difference in size, but Page 112 oA 2B \ 2C Figure 2 Schematic comparative drawings of Chromodorididae and Dorididae reproductive systems A. Chromodorididae reproductive system (after SCHMEKEL, 1972: fig. 50). B. Chromodorididae reproductive system, showing (clockwise from the bottom) vagina, insemination duct, receptaculum seminis and bursa copulatrix attached closely. C. Dorididae reproductive system (after SCHMEKEL, 1972: fig. 57) a gestalt configuration. In Chromodorididae (Figure 2 A) the receptaculum seminis is attached to the distal portion of the vagina (near the bursa copulatrix), or to the bursa copulatrix; it is not attached to the central portion of the insemination duct. At times the attachment points of the vagina, receptaculum seminis, and insemination duct are all close together on the bursa copulatrix (Figure 2 B). The typical dorid arrangement (Figure 2 C) has the re- ceptaculum seminis attached near the middle of the in- semination duct. This configuration is uniform throughout the Chromo- dorididae, and there are only a few exceptions among the other dorids. From tropical west American waters, Dori- opsilla janaina Marcus « Marcus, 1967, has a chromodor- id-type reproductive system (Marcus & Marcus, 1967: 97; fig. 125). It is immediately distinguished from a chromodorid because it is a porostome and it has a rough, highly tubercled dorsum. In 2 Mediterranean species, Doris verrucosa Cuvier, 1804 (authorship and date fide THE VELIGER Vol. 20; No. 2 SCHMEKEL, 1968) and Doris ocelligera (Bergh, 1881), the reproductive systems’ arrangement approaches that of the Chromodorididae. The receptaculum seminis is at- tached to the insemination duct near the latter’s attach- ment with the vagina (ScHMEKEL, 1968: figs. 2, 4,8); a dorsum covered with large tubercles and the shape of the radular teeth clearly separate these species from the Chro- modorididae. It is of special note, showing that these are rare exceptions to the general rule, that related species in these genera have the typical Dorididae reproductive sys- tem arrangement, with the receptaculum seminis clearly attached to the insemination duct: cf. the drawing of Dori- opsilla nigromaculata (Cockerell, in Cockerell & Eliot, 1905) by Marcus & Marcus (1967: 206; fig. 62C) and of Doris marmorata Risso, 1818, by SCcHMEKEL (1968: figs. 6d and 8). c) Radula. The shape of the lateral teeth is the most characteristic feature of the Chromodorididae. Each tooth (Figure 1) has a thin, knifelike, erect shaft, usually with denticles. By contrast, the teeth of Dorididae (cf, Marcus & Marcus, 1967: 81, 83, 86, etc.) are much more hook or sickle shaped, and broader laterally than are those of Chromodorididae. In addition, within the Chromodorid- idae, subfamily and generic separations are based on ma- jor differences of radular morphology. By using jointly the 3 characteristics of body form and texture, reproductive system, and radula, the members of a genus can be placed reliably within (or excluded from) the Chromodorididae. For example, BERcH (1891) established the Chromodorididae as encompassing 5 gen- era, including Aphelodoris Bergh, 1879b. Recent usage justifiably excludes Aphelodoris from the Chromodoridi- dae; although the animals have a smooth dorsum, their reproductive system and radular teeth are not chromo- dorid. Genera of the Chromodoridinae OpHNER (in Franc, 1968: 867) included 3 genera in Chromodorididae which are highly problematic. All 3 (Ceratodoris J. E. Gray, in M.E. Gray, 1850; Gruvelia Risbec, 1928, and Otinodoris White, 1948) need further study and adequate descriptions of their radulae and re- productive systems before determining their taxonomic placement. Rosodoris Pruvot-Fol, 1954, was originally placed within the Glossodorididae; it is probably not a Chromodorididae, but its familial relationships are un- certain without additional material. Ceratosoma Adams & Reeve, 1850, is probably a member of the Chromodorid- idae, but examination of specimens is required to estab- lish its subfamilial status. Vol. 20; No. 2 The following 8 genera are what I recognize as com- prising the Chromodoridinae, with the distinguishing characteristics of their radulae: Chromodoris Alder & Hancock, 1855 (xvil) (type genus of the family and subfamily) Synonym: Felimida Marcus, 1971 (see THOMPSON, 1972: 398, for additional synonymy). Type Species: Doris magnifica Quoy & Gaimard, 1832 (= D. quadricolor Riippell « Leuckart, 1831). Radular teeth unicuspid, with serrate denticulation; innermost lateral tooth with denticles on both sides of cusp; strongly recurved erect shafts of approximately 2"4 through 8" lateral teeth with denticles on lateral face (some species may have denticles on both sides of the cusps of teeth 1-8) ; erect shafts of teeth lengthen toward the middle of the half-row, with denticles on posterior surface. Outermost lateral teeth become smaller, and retain denti- culation. Chromolaichma Bertsch, gen. nov. Type Species: Casella sedna Marcus & Marcus, 1967. Also included: Chromodoris dalli Bergh, 1879¢; Chromo- doris punctilucens Bergh, 1890a; possibly Chromodoris youngbleuthi Kay & Young, 1969. Radular rows at least 2-3 times greater than maximum number of teeth per half-row; width : length ratio greater than 1:3; small rachidian tooth usually present; radular teeth unicuspid, with lateral serrate denticulation; outer- most laterals smooth (with larger radulae, there is a greater number of smooth outer teeth). Etymology: ‘Colorful spear-point,” derived from 76 xe@ua: (color) and 4 aixuy (the point of a spear), re- ferring to the bright colors of the living animals and the elongate, spear-like radula; the letter “J” inserted for euphony; generic name feminine. Mexichromis Bertsch, gen. nov. Type Species: Chromodoris antonii Bertsch, 1976b. Also included: Chromodoris porterae Cockerell, 1901; Chromodoris tura Marcus & Marcus, 1967. Radular teeth acuspid; pectinate denticulation. Etymology: “Mexican sea-fish,” derived from chromis (sea-fish) ; generic name masculine. THE VELIGER Page 113 Thorunna Bergh, 1877 (plate 58, figures 30-32) Synonym: Noumea Risbec (1928: 165). Type Species: Thorunna furtiva Bergh, 1877. Radular formula n-1:0°1-n; first innermost lateral tooth with an extremely enlarged base. Although recent references (e. g., ODHNER in FRANC, 1968: 867; RUSSELL, 1971: 112) cite 1878 as the date of authorship, Bergh published the new names Thorunna furtiva in 1877, accompanied by 3 illustrations. Hence the correct date is 1877. Babaina Odhner, 1968 OpDHNER in Franc, 1968: 867 (not Rotter, 1972, which = Babakina Roller, 1973). Type Species: 143-144). Also included: Babaina arbuta (Burn, 1961), comb. nov. (originally Glossodoris), and Babaina daniellae (Kay & Young, 1969), comb. nov. (originally Hypselodoris). Radular teeth have elongate, narrow, erect shafts; bifid (tooth shaft bifurcates distally, but does not have the prominent anterior primary cusp of Hypselodoris) ; with- out denticulation. Glossodoris florens Baba, 1949 (pp. 53; Casella H. « A. Adams, 1854 (Dp: O27) Type Species: Casella gouldii H. s A. Adams, 1854, = Doris atromarginata Cuvier, 1804. Living animal with a stiff, smooth mantle margin that is lobed. Radular teeth unicuspid, lateral teeth hook-shaped with denticles on the outer face; rachidian tooth absent. Princi- pal diagnosis of Casella rests on the texture of the notum and the notal margin. Hypselodoris Stimpson, 1855 (pp. 388-389 ) Type Species: Goniodoris obscura Stimpson 1855. Radula usually without a median tooth; lateral teeth bi- cuspid; inner lateral teeth with denticles on outer face; most of the lateral teeth with posterior denticles on the erect shaft below the secondary cusp. Page 114 Felimare Marcus & Marcus, 1967 (p. 62) Type Species: Felimare bayeri Marcus « Marcus, 1967. Radula “with a rachidian plate nearly as high as the neighboring teeth, provided with a smooth cusp.” The single species in this currently monotypic genus has bi- cuspid lateral teeth; the inner lateral teeth are without denticles, but the outer laterals are denticulate on the posterior surface of the erect shaft. A number of Hypselo- doris species possess a similar denticulation pattern. Fur- ther study of these species in contrast with those that have denticles on all the lateral teeth, may warrant a re-eva- luation of the importance of the enlarged rachidian tooth, and a generic separation based on patterns of denticu- lation. These major gestalt patterns of the radulae and their teeth separate the Chromodoridinae into 8 genera. Pre- cise feeding studies are needed to elucidate the function- ing differences between these radular patterns. BLoom’s (1976) work correlated the degree of bending of radular teeth with division of sponge food resources. Functional morphology studies examining broad patterns of Chromo- doridinae radular shapes will show additional separation of feeding resource utilization. For instance, studies of feeding specificity should indicate that members of the Chromodoridinae with narrow, long radulae (e. g., Chro- molaichma) will utilize a different sponge prey item than those which have a proportionately wider and shorter radula. Synopsis of Chromodoridinae Species from the Pacific Coast of America For comparative purposes, these species are all illustrated in Figure 3. Chromodoris: Chromodoris baumanni Bertsch, 1970 Chromodoris sp. (to be named in Part IT) Chromodoris marislae Bertsch, in Bertsch et al., 1973 Chromodoris mcfarlandi Cockerell, 1901 Chromodoris norrist Farmer, 1963 Chromodoris sphoni (Marcus, 1971) Chromolaichma: Chromolaichma sedna (Marcus « Marcus, 1967) Chromolaichma dalli (Bergh, 1879c) THE VELIGER Vol. 20; No. 2 Synonyms: Chromodoris banks: Farmer, 1963 Chromodoris banksi sonora Marcus & Marcus, 1967 Mexichromis: Mexichromis antonii (Bertsch, 1976b) Mexichromis porterae (Cockerell, 1901) Mexichromis tura (Marcus & Marcus, 1967) (species name ending not changed because of uncertain etymology) Hypselodoris: Hypselodoris agassizi (Bergh, 1894) Synonym: Chromodoris aegialia Bergh, 1904 Hypselodoris californiensis (Bergh, 1879c) Synonyms: Chromodoris glauca Bergh, 1879b (nomen oblitum) Chromodoris universitatis Cockerell, 1gOl Hypselodoris sp. (to be named in Part IV) Hypselodoris lapislazuli (Bertsch & Ferreira, 1974) (Chromodoris juvenca Bergh, 1898, from Chile, is Cad- lina juvenca, and hence not a Chromodoridinae. ) Detailed analyses of each species will appear in Parts II, III, and IV. Because of length considerations, this study? will appear as 4 separate articles in this journal. 2 Submitted as a dissertation in partial fulfillment of the require- ments for the Ph. D. degree in zoology, University of California, Berkeley. (on facing page —>) Figure 3 Drawings of living Chromodoridinae from the American Pacific coast A. Chromodoris baumanni (after Marcus « Marcus, 1967: 172) B. Chromodoris sp. (after color transparencies by Alex Kerstitch) C. Chromodoris marislae (after BertscH et. al., 1973: 294) D. Chromodoris mcfarlandi (after MacFarLanp, 1966: plt. 22) E. Chromodoris norrisi (after Marcus & Marcus, 1967: 171) E. Chromodoris sphoni (after Marcus, 1971: 356) G. Chromolaichma sedna (after Marcus « Marcus, 1967: 179) H. Chromolaichma dalli (after Marcus « Marcus, 1967: 174) I. Mexichromis antonu (after BERTSCH, 1976b: fig. 1) J. Mexichromis porterae (after MacFaruanp, 1966: plt. 24) K. Mexichromis tura (after Marcus « Marcus, 1967: 52) L. Hypselodoris agassizii (after Marcus « Marcus, 1967: 177, fig. 31) M. Hypselodoris californiensis (after MACFARLAND, 1966: plt. 24) N. Hypselodoris sp. (after Marcus « Marcus, 1967: 177, fig. 30) O. Hypselodoris lapislazuli (after BERTSCH & FERREIRA, 1974: fig. 1) THE VELIGER Page 115 No. 2 ° b) Vol. 20 Page 116 Literature Cited ADAMS, ARTHUR & LoveELL AuGusTUS REEVE 1850 [1848-1850]. Mollusca. In: A. Apams, ed., The zoology of the voyage of H. M. S. Samarang, under the command of Captain Sir Ed- ward Belcher, ... during the years 1843-1846. London (Reeve, Ben- ham, Reeve) x+87 pp.; 24 plts. [pp. 1-24 (November 1848); pp. 25-44 (May 1850); pp. 45-87 (August 1850)] [collation: SHERBORN, 1922: cxi] ApaMs, Henry & ARTHUR ADAMS 1854-1858. The genera of Recent Mollusca; arranged according to their organization. 2: 1-661. Van Voorst, London (Sept. 1854 - Nov. 1858) ALDER, JosHUA & ALBANY Hancock 1854. IX. Notice of some new species of British Nudibranchiata. Ann. Mag. Nat. Hist. (2) 14: 102 - 105 (August 1854) 1855. A monograph of the British nudibranchiate Mollusca, with figures of all species. London, Ray Soc., Family 1, plts. 21a, 27; fam. 2, plts. 1, 2; fam. 3, plts. 38a, 45-48; Appendix pp. I - XXVIII ArcabD!I, JOHN A. & NorMAN HopckIN 1973. Scanning electron micrographic study of the dorsal integument of the land slug Lehmannia poirieri (Mabille, 1883). The Veliger 15 (4): 338 - 339; 2 plts. (1 April 1973) BasBa, KixuTAarO 1949, Opisthobranchia of Sagami Bay, collected by his Majesty the Emperor of Japan. Iwanami Shoten, Tokyo: 4+2+194+7 pp.; 50 plts.; 161 text figs. (September 1949) BanbDEL, Kiaus 1974, Studies on Littorinidae from the Atlantic. 17 (2): 92-114; 5 plts.; 22 text figs. Barser, V. C. 1972. Preparative techniques for the successive examination of biologi- cal specimens by light microscopy, SEM and TEM. Scanning Electron Microscopy/1972 (II): 321-326; 8 text figs. Beeman, Rosert Davip 1963. Variation and synonymy of Phyllaplysia in the northeastern Pacific (Mollusca : Opisthobranchia). The Veliger 6 (1): 43-47; 5 text figs. (1 July 1963) BerocH, Lupwic SopHus Rupo.r 1877, Malacologische Untersuchungen. Jn: C. Semper, Reisen im Archipel der Philippinen 2 (12): 495 - 546; plts. 58-61 (15 Dec. ’77) 18782. Malacologische Untersuchungen. In: C. Semper, Reisen im Archipel der Philippinen 2 (13): 547-601; plts. 62-65 (8 July ’78) 1878b. Malacologische Untersuchungen. Jn: C. Semper, Reisen im Ar- chipel der Philippinen 2 (14): 603 - 645, I- LI; plts. 66 - 68 (23 Dec.) 1879a. Neue Nacktschnecken der Siidsee, malacologische Untersuchun- The Veliger (1 October 1974) gen, IV. Journ, Mus. Godeffroy 5 (14): 1-50; plts, 1-5 1879b. | Neue Chromodoriden: Malakoz. Blatter, N. F 1: 87-116; pit. 3 1879c. On the nudibranchiate gasteropod Mollusca of the North Pacific Ocean, with special reference to those of Alaska, prt. 1. Proc. Acad. Nat. Sci. Philadelphia, 31: 71 - 132; plts, 1-8 (10 May 1879) 1881. | Malacologische Untersuchungen. Nudibranchien. Nachtrage und Erganzungen. In: C. Semper, Reisen im Archipel der Philippinen 2 (Supplement 2): 79-128; plts. G-L 1890a. Report on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78), and in the Carib- bean Sea (1879-80), by the U.S.Coast Survey Steamer “Blake,” Lieut.-Commander C. D, Sigsbee, U.S. N., and Commander J. R. Bart- lett, U.S.N., commanding. XXXII. Report on the nudibranchs. Bull. Mus. Comp. Zoology Harvard 19 (3): 155 - 181; 3 plts. (March 1890) 1891. Die cryptobranchiaten Dorididen. Zool. Jahrb., Abt. Syst. 6 (1): 103 - 144 (20 October 1891) 1894. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U. S. Fish Commission steamer “Albatross,” during 1891, Lieut. Com- mander Z. L. Tanner, U. S. N., commanding. XIII. Die Opistho- branchien. Bull. Mus. Comp. Zool. Harvard Univ. 25 (10): 125 to 233; plts. 1-12 (October 1894) 1898. Die Opisthobranchier der Sammlung Pilate. Zool. Jahrb. (Suppl.) 4 (3): 481-582; plts. 28 - 33 (15 December 1898) 1904. Malacologische Untersuchungen: Nudibranchiata. In: C. Sem- per, Reisen im Archipel der Philippinen, Wissenschaftliche Resultate 9 (6):(1): 1-55; pits. 1-4 (2 February 1904) THE VELIGER Vol. 20; No. 2 BertscH, Hans 1970. Opisthobranchs from Isla San Francisco, Gulf of California, with the description of a new species. Contrib. Sci., Santa Barbara Mus. Nat. Hist 2: 1-16; 13 text figs. (1 December 1970) 1976a. Intraspecific and ontogenetic radular variation in opisthobranch systematics (Mollusca: Gastropoda). System. Zool. 25 (2): 117-122 7 text figs. (9 July 1976) 1976b. A new species of Chromodoris (Opisthobranchia : Nudibran- chia) from tropical west America. The Veliger 19 (2): 156 - 158; 1 plt.; 2 text figs. (1 October 1976) BertscH, Hans & ANTONIO J. FERREIRA 1974. Four new species of nudibranchs from tropical west America. The Veliger 16 (4): 343 - 353; 7 pits. (1 April 1974) BrertscH, Hans, ANTONIO J. FERREIRA, WESLEY M. FARMER & THOMAS L. Hayes 1973. The genera Chromodoris and Felimida (Nudibranchia : Chromo- dorididae) in tropical west America: distributional data, description of a new species, and scanning electron microscopic studies of radulae. The Veliger 15 (4): 287-294; 3 plts.; 3 text figs. (1 April 1973) Bertscu, Hans & KaniauLcono B. MEYER In prep. Opisthobranchs of the Pacific coast of Panama, with additional data on some Californian and Atlantic species. Bioom, STEPHEN A. 1976. | Morphological correlations between dorid nudibranch preda- tors and sponge prey. The Veliger 18 (3): 289-301; 1 text fig. (1 January 1976) Borxkowsk1, THOMAS V. 1975. Variability among Caribbean Littorinidae. (4): 369-377; 4 text figs. BoucHET, PHILIPPE 1975. Opisthobranches de profondeur de !’Ocean Atlantique. 1. -- The Veliger 17 (1 April 1975) (Cephalaspidea. Cah. Biol. Marine 16 (3): 317-365; 4 plts.; 19 text figs. Burn, RoBert 1961. A new doridid nudibranch from Torquay, Victoria. The Veliger 4 (2): 55-56; plt. 15 Criark, Joun M. & SEYMouR GLAcov 1976. Evaluation and publication of scanning electron micrographs. Science 192 (4246): 1360 - 1361 (25 June 1976) CockERELL, THEODORE Dru ALLISON 1901. Pigments of nudibranchiate Mollusca. (1 October 1961) Nature 65 (1674): 79 - 80 (28 November 1901) CockERELL, THEODORE Dru ALLISON & CHareEs N. E. ELior 1905. Notes on a collection of California nudibranchs. Journ. Malacol. 12 (3): 31-53; plts. 7, 8 Cooper, JAMES GRAHAM 1863, On new or rare Mollusca inhabiting the coast of California — No. II Proc. Calif. Acad. Sci. 3: 56-60; fig. 14 (September 1863) Cuvier, Gzorces Liorpotp CHréTIEN FrEDERIC DAGOBERT 1804. Mémoire sur le genre Doris. Ann. Mus. Nation. Hist. Nat. Paris 4: 447-473; plts. 73-74 (August 1804) (not seen, fide Russet, 1971] Dat, WILLIAM HEALEY 1900, On a genus (Phyllaplysia) new to the Pacific coast. The Nautilus 14 (8): 91-92 (6 December 1900) DoszHANsky, THEODOSIUS 1970. | Genetics of the evolutionary process. New York ix + 505 pp.; illus. ENGEL, HENpRIK & P WaGENAAR HUMMELINCK 1936. Uber westindische Aplysiidae und Verwandten anderer Gebiete. Capita Zoologica 8 (1): 1-76; 43 text figs. EveRHART, THomas E. & THomas L. Hayes 1972. The scanning electron microscope. Sci. Amer. 226 (1): 54-69; 28 text figs. (January 1972) FARMER, WESLEY MERRILL 1963. | Two new opisthobranch mollusks from Baja California. ‘Trans. San Diego Soc. Nat. Hist. 13 (6): 81-84; 1 plt.; 1 text fig. (27 September 1963) (29 September 1905) Columbia Univ. Press, FERREIRA, ANTONIO J. 1974. The genus Lepidozona in the Panamic Province, with the de- scription of two new species (Mollusca : Polyplacophora). The Veliger 17 (2): 162-180; 6 plts. (1 October 1974) 1976. A new species of Callistochiton in the Caribbean. The Nautilus 90 (1): 46-49; 6 text figs. (30 January 1976) 1977. A review of the genus Triopha (Mollusca : Nudibranchia). The Veliger 19 (4): 387 - 402; 3 plts.; 10 text figs. (1 April 1977) Vol. 20; No. 2 FINDLEY, JAMEs S. 1976. ‘The structure of bat communities. 129 - 139; 3 text figs. Franc, ANDRE 1968. Sous-classe des opisthobranches. Mollusques gasteropodes et Amer. Natur. 110 (971): (27 February 1976) scaphopodes. In: P. Grassé, Traité de zoologie: anatomie, systém- atique, biologie. 5 (3): 608-893; plts. 10-11; text figs. 310 to 441 Paris, Masson & Cie. Guisetin, MicHaE, TENANT 1966. Reproductive function and the phylogeny of opisthobranch gast- ropods, Malacologia 3 (3): 327-378; 7 text figs. (31 May 1966) Gonor, JEFFERSON JOHN 1961. Observations on the biology of Hermaeina smithi, a sacoglossan opisthobranch from the west coast of North America. The Veliger 4 (2): 85-98; 13 text figs. (1 October 1961) Grant, PR. 1967. Bill length variability in birds of the Tres Marias Islands, Mexi- co. Canad, Journ. Zool. 45 (5) 805 - 815 (14 September 1967) Gray, Maria EMMA 1850. Figures of molluscous animals, selected from various authors. Vol. 4, iv + 219 pp., Longman, Brown, Green & Longmans, London (August 1850) HagFEvrincer, Hans-Rupotr 1959. Remarques sur le developpement du dessin de quelques glosso- doridiens (Mollusques opisthobranches). Rev. Suisse Zool. 66 (2): 309-315; 8 text figs, (August 1959) Hayegs, Tuomas L. 1972. Invariance under transformation: a useful concept in the inter- pretation of SEM images. Scanning Electron Microscopy/1972 (1): 57-64; 9 text figs. (April 1972) 1973. Scanning electron microscope techniques in biology. In: J. F Koehler (ed.), Advanced techniques in biological electron micro- scopy. Springer Verlag, New York, 153-214; 10 text figs. Kay, EvtzaBETH ALIson & Davin K. Youno 1969. The Doridacea (Opisthobranchia; Mollusca) of the Hawaiian Islands. Pacif. Sci, 25 (2): 172 - 231; 82 text figs. (April 1969) Kezn, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. ed. 2. Stanford Univ. Press, Stanford, Calif. i-xiv+ 1064 pp.; ca. 4000 figs.; 22 color plts. (1 September 1971) Linpserc, Davin R. 1977, Artifacts incurred by the treatment of acmaeid radulae with al- kalies. The Veliger 19 (4): 453-454; 1 text fig. (1 April 1977) MacFarranp, Frank Mace 1905. A preliminary account of the Dorididae of Monterey Bay, Cali- fornia. Proc. Biol. Soc. Wash. 18; 35 - 54 (2 February 1905) 1966. Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif, Acad. Sci. 6: xvit546 pp.; 72 plts. (8 April 1966) Marcus, ERNsT 1958, On western Atlantic opisthobranchiate gastropods, Amer, Mus. Novit. 1906: 1-82; 111 text figs. (13 August 1958) 1961. Opisthobranch mollusks from California, The Veliger 3 (Supplmt. I): 1-85; plts, 1-10 (1 February 1961) Marcus, Eve.ine pu BoiseReYMonD 1971. On some euthyneuran gastropods from the Indian and Pacific Oceans, Proc. Malacol. Soc. London 39 (5): 355-369; 20 text figs. (August 1971) Marouvs, Evz.ing pu Bois-ReyMonp & Ernst MaRaus 1967. American opisthobranch mollusks. Studies in tropical oceano- graphy (Univ. Miami Inst. Marine Sci., Miami, Florida), no. 6: viii+ 256 pp.; figs. 1-155 + 1-95 (22 December 1967) Maroouis, STANLEY V. & Rospert E, CARVER 1974, | Microstructure of chalky deposits found in shells of the oyster, Crassostrea virginica. The Nautilus 88 (2): 62-65; 1 text fig. (April 1974) MaxwE Lt, WILLIAM L, 1975, Scanning electron microscope studies of pulmonate spermato- zoa. The Veliger 18 (1): 31-33; 3 plts. (1 July 1975) Mayr, ERNsT 1970. Populations, species and evolution. bridge, Mass. xv + 453 pp.; 51 text figs. McDonatp, Larry W. & Tuomas L. Hayes 1969. Correlation of scanning electron microscope and light micro- scope images of individual cells in human blood and blood clots. Exper. Molecul. Pathol. 10 (2): 186-198; 6 text figs. (April 1969) Belknap Press, Cam- THE VELIGER Page 117 Morcu, Otto AnprEAS LAWSON 1863. Contributions 4 la faune malacologique des Antilles danoises. Journ. de Conchyl. (3) 3 (11): 21-43 (1 January 1863) 1869. Catalogue des mollusques du Spitzberg recueillies par le Dr. H. Kroyer pendant le voyage de la corvette “La Recherche” en Juin 1838. Ann. Soc. Malacol. Belg. 4: 7 - 32 Mooser, Oswatpo & WALTER W. DALQuEsT 1975. Pleistocene mammals from Aguascalientes, Central Mexico. Journ. Mammal. 56 (4): 781-820; 12 text figs. (18 November 1975) Muuuner, Davin K. « Gaz G. SPHON 1974. A new Platydoris (Gastropoda: Nudibranchia) from the Gal4- pagos Islands, Trans. San Diego Soc. Nat. Hist. 17 (15): 209 - 216; 5 text figs, (12 April 1974) Murray, M. J. & E. R. Lewis 1974, Sensory control of prey-capture in Navanax inermis. The Veliger 17 (2): 156-158; 1 plt.; 1 text fig. PAWLEY, James B. 1972. Charging artifacts in the scanning electron microscope. Scanning Electron Microscopy / 1972 (I): 153 - 160; 7 text figs. (April 1972) (1 October 1974) Pruvot-Fo., ALIcE 1954. Etude d’une petite collection d’opisthobranches d’océanie fran- Gaise. Journ, de Conchyl. 94 (1): 3-30; 37 text figs. (April ’54) Quoy, Jean René ConsTAnTIN & JosePpH PAUL GAIMARD 1832-1834, Voyage de la corvette l’Astrolabe exécuté par ordre du Roi, pendant les années 1826-1829, sous le commandement de H. J. Dumont d’Urville; Mollusques, Zoologie. Paris, 2(1): 1-686; Atlas pits. 1-26 (1833) [not seen; fide RusseExi, 1971] Rana, SANDER 1828. Histoire naturelle des aplysiens. 83 pp.; 25 plts. RISBEC, J. 1928. Contribution a l'étude des nudibranches Néo-Calédoniens, Faune colon. frang. 2 (1): 1-328; plts. A-D and 1-12; 98 text figs. Risso, JoseEpH ANTOINE 1818. Mémoire sur quelques gastéropodes nouveaux, nudibranches et tectibranches observés dans la mer de Nice. Journ. Phys. Chim. d’Hist. Nat. 87: 368-377 (November 1818) RoBERTSON, ROBERT 1971a, Sexually dimorphic archaeogastropods and radulae. Amer. Malac, Union, Ann. Rprt. for 1970: 75-78; 2 text figs. (18 February 1971) 1971b. Scanning electron microscopy of planktonic larval marine ga- stropod shells. The Veliger 14 (1): 1-12; 9 plts. (1 July 1971) Rotuer, Ricwarp A, 1972. Three new species of eolid nudibranchs from the west coast of North America (Gastropoda : Opisthobranchia). The Veliger 14 (4): 416-423; 28 text figs. (1 April 1972) 1973. | Babakina, new name for Babaina Roller, 1972, preoccupied. The Veliger 16 (1): 117-118 (1 July 1973) Ros, JoANDOMENEG 1974, Competéncia i evolucié en espécies veines de gasterépodes marins, Collog. Soc. Catal. Biolog. VII. Evoluci6: 101-121; 2 plts.; 1 text fig. : 1975. | Opistobranquios (Gastropoda: Euthyneura) del litoral ibérico, Investig. Pesq. (Barcelona) 39 (2): 269-372; 4 plts.; 6 text figs. (October 1975) Firmin Didot, Paris 7 + Ross, W. D. (ed.) 1952, The works of Aristotle. cago, III. 699 pp. ROUGHGARDEN, JONATHAN 1974, Niche width: biogeographic patterns among Anolis lizard popu- Vol. II, Encyclop. Brit., Inc. Chi- lations. Amer. Natur. 108 (962): 429-442; 1 text fig. (30 August 1974) Rupman, W. B. 1973. | Chromodorid opisthobranch Mollusca from the Indo-West Pa- cific. Zool. Journ, Linn. Soc. 52 (3): 175-199; 2 plts.; 7 text figs. (4 July 1973) Ruprett, WILHELM Peter EDUARD SIMON & FRIEDRICH SIGISMUND LEUCKART 1831 (for 1828). Neue wirbellose Thiere des Rothen Meeres. In: Eduard Riippell, Atlas zu der Reise im nérdlichen Afrika 5: 1-47 + 3 unnumb. pp.; plts, 1-12 RussE.._, Henry DRUMMOND 1971. Index Nudibranchia. Delaware iv+141 pp. Delaware Mus. Nat. Hist., Greenville, (1 July 1971) Page 118 THE VELIGER Vol. 20; No. 2 ScHILDER, FRANZ ALFRED & MARIA SCHILDER 1961a. Sexual differences in cowries. Proc. Malacol. Soc. London 34 (4): 207 - 209 (April 1961) 1961b. Zur Kenntnis der Cypraeidae. 3. Lange und Reihenzahl der Radula. Arch. Molluskenk. 90 (1/3) 33-42; 2 diagr. (17 July) 1963. Statistical studies on cowrie radulae. The Veliger 5 (3): 106-111; 2 text figs. (1 January 1963) ScHILDER, FRANZ ALFRED, MariA SCHILDER & GARFIELD HousTon 1964. The cowrie fauna of Penrith Island (Mollusca : Gastropoda). The Veliger 6(3): 155-161; 1 text fig. (1 January 1964) ScHMEKEL, LUISE 1968. Die Gattung Doris (Gastr. Nudibranchia) im Golf von Neapel. Pubbl. Staz. Zool. Napoli 36 (2): 167-187; 8 text figs. (19 August) 1972. Anatomie der Genitalorgane von Nudibranchiern (Gastropoda Euthyneura). Pubbl. Staz. Zool. Napoli 38(1): 120-217; 67 text figs. (1 October 1972) Simpson, Georce GayLorp, ANNE RoE & RicHarp C. LEWonTIN 1960. Quantitative Zoology, rev. edit. Harcourt, Brace & World Inc., New York vii + 440 pp.; 64 text figs. SoLtem, ALAN 1970. Malacological applications of scanning electron microscopy I. Introduction and shell surface features. The Veliger 12 (4): 394 to 400; plts. 58-60 (1 April 1970) 1972. Malacological applications of scanning electron microscopy II. Radular structure and functioning. The Veliger 14 (4): 327 - 336; 6 plts.; 1 text fig. (1 April 1972) Sotem, ALAN & EuGENE S. RICHARDSON 1975. Paleocadmus, a nautiloid cephalopod radula from the Pennsyl- vanian Francis Creek Shale of Illinois. The Veliger 17 (3): 233 - 242; 5 plts.; 1 text fig. (1 January 1975) STIMPSON, WILLIAM 1855. Descriptions of some new marine Invertebrata. Nat. Sci. Philadelphia 7 (10): 385 - 395 Proc. Acad. (July 1855) TamsitT, J. R. 1967. Niche and species diversity in neotropical bats. Nature 213 (5078): 784 - 786 (25 February 1967) Tuomas, Ronatp FRANK & FRASIER O. BINGHAM 1972. Shell structure in Spirula spirula (Cephalopoda). The Nau- tilus 86 (2-4): 63-66; 8 text figs. (November 1972) THompson, THomAS EVERETT 1958. Observations on the radula of Adalaria proxima (A. & H.) (Ga- stropoda Opisthobranchia). Proc. Malacol. Soc. London 33 (2): 49-56; plt. 6; 6 text figs. (August 1958) 1972. Chromodorid nudibranchs from eastern Australia (Gastropoda, Opisthobranchia). Journ. Zool. Soc. London 166 (3): 391 - 409; 4 plts.; 4 text figs. (13 March 1972) THompson, THomas Everett & H. E. Hinton 1968. Stereoscan electron microscope observations on opisthobranch radulae and shell-sculpture. Bijdr. tot de dierk. 38: 91 - 92; 4 plts. Tompa, ALEX 1976. Fossil eggs of the land snail genus Vallonia (Pulmonata: Valloni- idae). The Nautilus 90 (1): 5-7; 4 text figs. (30 January 1976) TroscHEL, FRaNz HERMANN 1856-1863. Das Gebiss der Schnecken zur Begriindung einer natiirlichen Classification. Nicolaische Verlagsbuchhandl. Berlin 1: viii + 252 + 20 unnumb. pp.; 20 plts. TROSCHEL, FRaNz HERMANN & JOHANNES THIELE 1866-1893. Das Gebiss der Schnecken zur Begriindung einer naturlichen Classification. Nicolaische Verlagsbuchhand]. Berlin 2: viii + 409 + 32 unnumb. pp.; 32 plts. Vitta-R., BERNARDO 1966. Los murcielagos de Mexico. Instit. Biolog. Univ. Nac. Autonom. Mexico, Mexico, D. F xv + 491 pp.; 98 maps; 171 text figs. frontisp. (23 December 1966) Waite, KaTHLEEN M. 1948. On a collection of marine molluscs from Ceylon. Proc Malacol. Soc. London 27 (5): 199-205; 11 text figs. (May 1948) Vol. 20; No. 2 THE VELIGER Page 119 The Family Columbellidae in the Western Atlantic Part Ila. - The Pyreninae BY GEORGE E. RADWIN Natural History Museum, San Diego, California 92112 (4 Plates) INTRODUCTION As NOTED IN THE FIRST article of this series (RADWIN, 1977), the family Columbellidae is a large cosmopolitan family of small buccinacean gastropods which, since an apparent Eocene origin, have undergone considerable ad- aptive radiation. The family may be divided into 2 un- equal sized subfamilies on the bases of radular dentition and, to a lesser extent, shell morphology (RADWIN, op. cit.). In Part I most of what is known concerning the phylogeny, ecology, and zoogeography of the family was reported, together with a systematic treatment of the gen- era and species assignable to the subfamily Columbellinae, the smaller of the 2 columbellid subfamilies. The present contribution continues the systematic treatment, covering the genera Anachis H. « A. Adams, 1853; Costoanachis Sacco, 1890; Metulella Gabb, 1873; Nassarina Dall, 1889; Parvanachis Radwin, 1968; and Steironepion Pils- bry & Lowe, 1932 of the subfamily Pyreninae. A total of at least 22 western Atlantic species is assignable to these genera. Another portion, Part IIb, is projected, which will cover the remaining g genera and their complement of at least 17 species in the region. ACKNOWLEDGMENTS The research upon which this paper is based was done in partial fulfillment of the requirements for the degree of Doctor of Philosophy at George Washington University, Washington, D. C. I would like to acknowledge my appreciation to Dr. Harald A. Rehder under whose direction this research was done; to Drs. Joseph Rosewater, Joseph P. E. Morri- son, and Wendell P. Woodring, who contributed critical suggestions. Dr. Norman Tebble, formerly of the British Museum (Natural History) and Mr. Jack Scott and the staff of the Photographic Laboratory, National Museum of Natural History, Washington, D.C., supplied photo- graphs of specimens. Mr. Anthony D’Attilio, San Diego Natural History Museum re-illustrated the radulae. Thanks are also due to Drs. Kenneth J. Boss, Museum of Comparative Zoology, William K. Emerson, American Museum of Natural History, and R. Tucker Abbott, (formerly of the Academy of Natural Sciences, Philadel- phia), for permitting me to examine the mollusk collec- tions of their respective institutions. In addition, I offer thanks to all individuals, too many to enumerate, who have helped me obtain additional specimens for study. The present research was carried out under a Smithsonian Predoctoral Internship. Standard abbreviations for museums may be found in part I of this series (The Veliger 19 (4): 409 - 410). Pyreninae Suter, 1913 Shell small to moderately large (2 - 40mm), fusiform or subfusiform; spire generally high and acute, spire whorls flat to convex in profile, suture varying from very shallow (Mitrella), to impressed (Amphissa) or incised (Suturo- glypta) ; body whorl varying from large and broad (Cos- mioconcha) to small and slender (Stetronepion), apert- ural lip generally denticulate on inner surface, columella either denticulate or smooth; siphonal canal short (Aeso- pus) to moderately long (Metulella). Sculpture variable, including axial elements, with or without spiral elements of variable strength. Each radular row with a flat, subrec- tangular median tooth, not as broad as typical for the Columbellinae, and devoid of cusps and denticles (Fig- ures A3, A4). Median tooth flanked on each side by a single sigmoid, bi- or tricuspid lateral tooth, narrower than that typical for the Columbellinae. Type genus: Pyrene Roding, 1798. Page 120 Anachis H. & A. Adams, 1853 Columbella (Anachis) H. & A. Avams, 1853. Gen. Rec. Moll.: 184 Anachis H. & A. Adams. Mércu, 1852. Cat. Conch. Yoldi: 254 Type species by SD (Tarte, 1868: 13), Columbella scala- rina Sowerby, 1832 (Figure 24). Shell moderate-sized to large (10 - 25mm in length), and broadly fusiform; spire moderately high and acute, whorls convex, suture distinct, not generally impressed; body whorl } the total shell length, heavy, shouldered, aperture wide, siphonal canal short; sculpture consisting of prom- inent axial ribs. Color pattern of 1 or 2 spiral bands of white on brown or the reverse. Each radular row consists of a single flat, subrectangular median tooth flanked on each side by a sigmoid, distally bicuspid lateral tooth (Figure 29). Remarks: ‘Tate’s selection of Columbella scalarina as the type species of Anachis limited the concept of the genus (s. 5.) to a small number of relatively large species with strong axial ribbing and a prominently shouldered body whorl. Many species previously assigned to Anachis are more properly assigned to Costoanachis Sacco, 1890, as they are smaller, their axial ribbing is neither as strong nor as consistent and they lack the thickened, shouldered body whorl. Anachis lyrata (Sowerby, 1832) (Figure 1) Columbella lyrata SoweErRBy, 1832. Proc. Zool. Soc. London 2: 114 (Panama Bay, Panamé; lectotype, BM(NH) 1966320) Colombella veleda Ductos, 1846. Illust. Conchyl., plt. 7, figs. 19-20 (type locality here designated as Boa Viagem, Bra- zil; representation of lectotype, Ductos, 1846, Illust. Conch. 4: plt. 7, figs. 19-20) Shell moderately large (14 - 20mm in length), and heavy; spire high and acute, whorls slightly shouldered, and THE VELIGER Vol. 20; No. 2 somewhat convex, suture impressed ; body whorl large and broad, aperture moderately wide, apertural lip thickened and denticulate within, columella straight and denticu- late, siphonal canal very short, anal groove present; sculp- ture of strong axial ribs; color ivory with 1 or 2 spiral rows of brown spots on the axial ribs. Each radular row con- sists of a flat, subrectangular median tooth, flanked on each side by a single, bicuspid lateral tooth (Figure jo). Remarks: Originally described from the Panamic-Pacif- ic province of Central and South America, specimens of this species from the Atlantic coast of South America are indistinguishable from Pacific examples. This is the only Recent species of Anachis s. s. occurring in the western Atlantic. Empty and worn shells of A. terp- sichore, an Indo-Pacific species have been collected in the Caribbean. These appear to be adventitious ballast rec- ords. No fossil record has been found for this species, although its Recent amphi-American distribution would suggest a substantial ancestry. Western Atlantic Range: Cuba and Central America to Santa Catharina, Brazil (also inhabits the tropical eastern Pacific). Costoanachis Sacco, 1890 Costoanachis Sacco, 1890 (in BELLARDI) Mem. Reale Accad. Sci. Torino 6: 57 (type species by SD [Pace, 1902, 5 (1/2): 43] Columbella (Anachis) turrita Sacco, 1890, not Sower- by, 1832 (= Costoanachis saccostata nom. nov.). (see Figure 2) Shell small to moderate in size (4 - 15mm in length), and fusiform; spire moderately high, acute, whorls flat to convex, suture shallow to incised or impressed ; body whorl equal to or less than 4 total shell length, aperture moder- ately wide, outer apertural lip generally denticulate with- in; columella straight, weakly denticulate, siphonal canal short and straight; sculpture of prominent axial ribs, in some cases limited to the body whorl, commonly with sub- Explanation of Figures 1 to 14 Figure 1: Anachis lyrata (Sowerby, 1832) Figure 2: Anachis turrita Sacco, 1890 (Costoanachis saccostata nom. nov.) Figure 3: Costoanachis avara (Say, 1822) Figure 4: Costoanachis hotessieriana (Orbigny, 1842) Figure 5: Costoanachis lafresnayi (Fischer & Bernardi, 1856) Figure 6: Costoanachis lafresnayi (Fischer & Bernardi, 1856) Figure 7: Costoanachis floridana (Rehder, 1939) Figure 8: Costoanachis hotessieriana (Orbigny, 1842) Figure 9: Costoanachis hotessieriana (Orbigny, 1842) Figure 10: Costoanachis sparsa (Reeve, 1859) Figure 11: Costoanachis scutulata (Reeve, 1859) Figure 12: Costoanachis catenata (Sowerby, 1844) Figure 13: Costoanachis sertularium (Orbigny, 1839) Figure 14: Costoanachis semiplicata (Stearns, 1873) THE VE icER, Vol. 20, No. 2 [Rapwin] Figures 7 to 74 Figure 5 Figure 14 Figure 10 Figure rr RES LG “ Ba —) i ‘ t ee ' = iY. 7 7 + Pa by iy — ie fa 4 ‘ - Dea r i = 1 ad i _ i hes ? 4 f \ n ‘ a a ¢ & i : tas Vol. 20; No. 2 microscopic spiral grooves between them. Color variable. Each radular row consists of a flat, subrectangular medi- an tooth flanked on each side by a single bi- or tricuspid Jateral tooth. These are shorter and more strongly bent than those of Anachis s. s. Remarks: Sacco, in his extremely brief description (“all whorls axially ribbed (except for C. corrugata Brocchi)”’), apparently did not intend to limit this name to only those species with axial sculpture on every whorl. With the expansion of the original description to include partially ribbed forms, many species previously assigned to Anachis but differing from the type species of that gen- us may now be assigned to Costoanachis (e.g., “A.” avara, “A.’sertularium, “A.” catenata). GarDNER (1948) and Oxtsson « HARBISON (1953) tacitly accepted this wider interpretation of Costoanachis without offering a verbal expansion of the original defi- nition, These authors placed Anachis obesa here, an as- signment which appears to be incorrect. Parvanachis Rad- win, 1968 was erected for the group of very small, squat, ribbed, Anachis-like forms with ventricose apertural lips, typefied by A. obesa. A new name for Columbella (Anachis) turrita Sacco, 1890 is needed, as this name is preoccupied by Columbella turrita Sowerby, 1832 (at present assigned to Strombina). Costoanachis saccostata Radwin, nom. nov. is introduced herein as a replacement. Costoanachis avara (Say, 1822) _ (Figure 3) Colombella avara Say, 1822. Journ. Acad. Nat. Sci. Phila. 2: 230 (type locality restricted by ScHELTEMA (1968) to “Florida”; lectotype ANSP 16887) Colombella cleta Ductos, 1846 (in CHEnu) Illust. Conchyl. pit. 15, figs. 13, 14 (type locality not specified; repre- sentation of lectotype, Ductos, 1846, 4: plt. 15, figs. 13, 14) Anachis avara (Say). Davi, 1889b. Bull. U.S. Nat. Mus. 37: 116 Shell moderate in size (10- 20mm in length); spire slightly greater than 4 shell length, whorls convex, suture shallow; body whorl cylindrical, moderately broad, aper- ture moderately wide, apertural lip slightly thickened, weakly denticulate interiorly, columella straight, nondent- iculate; siphonal canal short and straight; sculpture of widely spaced ribs on the body whorl only, all other whorls smooth. Color pattern of irregular brown blotches on a white background. Each row consists of a flat, rect- angular median tooth flanked on each side by a single sigmoid, bicuspid lateral tooth (Figure 37). THE VELIGER Page 121 Remarks: ‘This species is certainly one of the most com- mon shallow-water columbellids on the U.S. Atlantic coast. It is often confused with Costoanachis lafresnayt (= C. translirata) , another east coast columbellid. Costo- anachis avara is easily distinguished from C. lafresnayi by its lack of all sculpture on all whorls but the last one and by the convexity of its whorls; C. lafresnayi has flat-sided completely sculptured spire whorls. Costoanachis avara has been reported from as far south as Brazil and Argentina (Prspry, 1898; EYERDAM, 1950). These reports probably refer to C. sertulariarum Orbigny, a related South American species lacking spire sculpture. Costoanachis avara differs from C. sertulariarum in its larger apical angle, its more convex whorls, and its more impressed suture. No fossil examples of this species have been reported. Range: The Gulf of Maine to Miami, Florida on the eastern coast of the United States of America. Costoanachis catenata (Sowerby, 1844) (Figure 12) Columbella catenata SowERBY, 1844. Proc. Zool. Soc. London 12: 52 (here restricted to Montego Bay, Jamaica; repre- sentation of lectotype, SowERBy, 1847, sp. 94, fig 171) The location of the holotype is unknown. Originally part of the Cuming Collection, the type is not to be found at the British Museum (Natural History) where the Cuming Collection was deposited. Some small parts of the Cuming Collection were sold prior to its acquisition by the BM(NH). This type was apparently in one of those parts. Shell small ( 4- 7mm in length) ; spire slightly over 4 total shell length, whorls convex, distinctly shouldered, with impressed suture; body whorl cylindrical, apertural lip strongly denticulate within, columella strongly denti- culate, bent at its anterior end, siphonal canal short and bent; sculpture of strong, sharply cut axial ribs with weak intercostal spiral grooves and a single strong subsutural spiral groove; color white with spiral bands of brown or orange “chain-links.” Protoconch 2 swollen, translucent- white whorls. The radula of Costoanachis catenata was not examined, as none of the specimens examined had been collected alive. Remarks: This species has been confused with Costo- anachis sparsa and C. scutulata where their ranges over- lap in Bermuda. However, the strongly denticulate aper- tural lip and columella, the strongly bent columella and siphonal canal, the striking patterns of color and sculp- Page 122 ture, and a rather distinctive bulbous protoconch serve to distinguish C. catenata from these 2 congeners. No fossil specimens of this species have been reported. Range: Bermuda and southern Florida to Colén, Pan- ama and Bahia, Brazil. Costoanachis fenneli Radwin, 1968 (Figure 15) Costoanachis fenneli Rapwin, 1968. Proc. Biol Soc. Wash. 81: 147 (Sacco Sao Francisco, Guanabara, Brazil; holotype, USNM 539122) Shell small (5-7mm in length); spire acute, slightly more than ¢ total shell length, spire whorls slightly con- vex, suture weakly impressed; body whorl expanded, angular, asymmetrical, left side slightly more expanded than right, outer apertural lip denticulate within, anal sinus strong, columella straight, weakly denticulate; sculp- ture of strong axial ribs, crossed by fine shallow, spiral grooves; yellowish-white ground color with anastomosing brown blotches; protoconch of almost 3 full, glassy brown whorls. Each radular row consists of a flat subrectangular median tooth flanked on each side by a single sigmoid, bicuspid lateral tooth (Figure 32). Remarks: This species is known from a single lot of 35 specimens, collected alive by Dr. Doris Cochrane near Nictheroy, Guanabara State, Brazil in April 1935. Its large, angulate body whorl, prominent axial and distinct- ive spiral sculpture, color pattern and multiwhor! proto- conch distinguish this species from all others. No fossil examples have been reported. Known to me only from the type locality, Sacco Sao Francisco, Guanabara, Brazil. Costoanachis floridana (Rehder, 1939) (Figure 7) Anachis floridana REHDER, 1939. The Nautilus 53 (1): 20-21; THE VELIGER Vol. 20; No. 2 plt. 6 (near Cape Canaveral, Brevard County, Florida; holotype, USNM 473202) Shell small (6 - 12mm in length); spire total shell length, whorls slightly convex, suture shallow; body whorl cylind- rical and moderately wide, apertural lip slightly thick- ened, distinctly denticulate within, columella straight, weakly denticulate; sculpture of low, widely spaced axial ribs on only the last 2 whorls, disappearing below the periphery. There are also numerous microscopic spiral lines over the entire adult shell; color pale yellow with irregular blotches of purplish brown. Each radular row consists of a rectangular median tooth flanked on each side by a single sigmoid tricuspid lateral tooth. The 3 cusps of the lateral teeth are very long and strongly curved (Figure 35). Remarks: Considered a variety of Costoanachis avara by some, C. floridana actually appears more closely re- lated, on the basis of the radula and certain shell charac- ters, to C. sertulariarum. It may be separated from that species on the basis of geographic distribution (southern Florida vs. Brazil), its smaller size, more convex spire pro- file, slight differences in lateral radular dentition and presence of microscopic shell sculpture. REHDER (1939) in his original description of this species states that it differs from Costoanachis avara in “lacking the spiral grooves of that species.” The holotype of C. floridana has distinct spiral sculpture. Although the cata- log number and dimensions in the original description agree with those of the labeled holotype, Rehder states that the holotype was collected in Brevard County near Cape Canaveral, Florida. The locality listed with the holotype in the USNM type collection is near S. Jetty, St. John’s Bar, Mayport, Duval Co., Fla. This confusion apparently arose because of a mixup in labeling. A lot of Parvanachis obesa with a label giving the published type locality of C. floridana has been found in the USNM collection. No fossil examples of this species have been reported. Range: Off Beaufort, North Carolina to Dade County, Explanation of Figures 15 to 28 Figure 15: Costoanachis fenneli Radwin, 1968 Figure 16: Steironepion monilifera (Sowerby, 1844) Figure 17: Steironepion minor (C.B. Adams, 1845) Figure 18: Anachis scalarina Figure 19: Metulella columbellata (Dall, 1889) Figure 20: Nassarina bushii (Dall, 1889) Figure 21: Nassarina metabrunnea Dall & Simpson, 1901 Figure 22: Parvanachis rhodae (Radwin, 1968) Figure 23: Nassarina glypta (Bush, 1885) Figure 24: Parvanachis isabellei (Orbigny, 1839) Figure 25: Parvanachis isabellei (Orbigny, 1839) Figure 26: Parvanachis obesa (C. B. Adams, 1845) Figure 27: Parvanachis ostreicola (Sowerby, 1885) Figure 28: Parvanachis melvillei (Strebel, 1905) Tue VEtIcER, Vol. 20, No, 2 [Rapwin] Figures 15 to 28 Figure 15 Figure 16 Figure 21 ie Figure 23 Figure 22 Figure 27 # Figure 25a Figure 25 Figure 26 Figure 2 Vol. 20; No. 2 Florida, and from Carancahua Bay, Texas to Brownsville, Texas. Costoanachis lafresnayi (Fischer & Bernardi, 1856) (Figures 5, 6) Columbella lafresnayi FisHER & BERNARDI, 1856. Journ. de Conchyl. 5: 357; plt. 12, figs. 4, 5 (Guadeloupe; holo- type, CJC) Columbella ocellata REEVE, 1859 (not GMELIN, 1791). Conch. Icon., Columbella, pit. 37 , fig. 237 (type locality not specified; holotype, BM[NH] Columbella translirata RavENEL, 1861. Proc. Acad. Nat. Sci. Philadelphia 3: 41-42 (off Charleston, South Carolina; holotype apparently destroyed) Columbella (Anachis) avara Kosext, 1897 (not Say, 1822). (in) Martini & CHEMNITZ, Conchyl. Cabinet 3-Id: 62; pit. 8, figs. 8, 9 Moderately large (10-16mm in length); spire high (more than $ total shell length), whorls flat-sided, suture moderately impressed; body whorl fusoid; aperture mod- erately broad, apertural lip thickened, denticulate interi- orly, columella straight and strongly denticulate, siphonal canal short to moderately long, slightly bent; sculpture of numerous prominent axial ribs crossed by weaker spiral grooves; color from straw yellow to chestnut brown with a spiral subsutural row of white spots. Each radular row consists of a flat, subrectangular median tooth flanked on each side by a sigmoid, bicuspid lateral tooth (Figure 33). Remarks: ‘The type of Columbella translirata was lost, along with the remainder of Ravenel’s types, which were reportedly destroyed during the burning of Atlanta, Geor- gia in the U.S. Civil War. This species exhibits a cline in shell characters coinci- dent with apparent tropical submergence from north to south along the Atlantic coast of the United States. In the north, where Costoanachis lafresnayi may be collected in moderately shallow water, the shell is relatively broad with a moderately acute apical angle and a very short siphonal canal. Southward, this species inhabits progres- sively deeper water, to 36m or more. Here the shell is lighter-colored and more slender, with a more acute api- cal angle and a somewhat longer siphonal canal. Examples of this species are known from the Pliocene of Florida and are otherwise known only from the Recent. Range: Grand Manan Island, New Brunswick, Canada to Key West, Florida and Yucatan, Mexico; occasional specimens from Barbados, West Indies. THE VELIGER Page 123 Costoanachis hotessieriana (Orbigny, 1842) (Figures 4, 8, 9) Columbella hotessieriana Orsicny, 1842. in Sacra, Hist. Phys. Polit. Nat. Cuba. Atlas, plt. XXI, figs. 37-39 (Guade- loupe; lectotype, BM[NH] 1854.10.4.359; paralectotype BM[NH] 1854.10.4.359/1) Columbella guildingi SowErBy, 1844. Proc. Zool. Soc. London 12: 53 (St. Vincent, B. W. I.; holotype, BM[NH] 1966. 448) Columbella hotessieri OrBIGNY, 1845. (in SacRA), Hist. Fis. Polit. Nat. Cuba. Text, p. 234 Anachis megintyi Usticxe, 1959. Checklist mar. shells St. Croix, p. 67; pit. III, fig. 15 (St. Croix, Virgin Islands; holotype, Usticke Coll.) Shell small (6.0- 7.5mm in length); spire moderately high (about 4 total shell length) and acute, whorls con- vex, suture impressed; body whorl subcylindrical, aper- tural lip moderately thickened, denticulate interiorly, columella straight, weakly denticulate, siphonal canal short to moderate in length, slightly bent, anal sinus barely discernible; sculpture of axial ribs crossed by shal- low, closely-spaced spiral grooves which become crowded just below the suture; color pattern varying from choco- late brown to light tan with a spiral band of white spots of varying width just above the suture. Each radular row consists of a flat, subrectangular median tooth flanked on each side by a sigmoid, tricuspid lateral tooth (Figure 34). Remarks: First named Columbella hotessieriana by Orpicny (1842), this species had its name emended to C. hotessieri without explanation by the same author (1845) in subsequent editions of the same work. Costoanachis scutulata (Reeve, 1859) (Figure 17) Columbella scutulata REEve, 1859. Conch. Icon. Columbella. pl. XXX, spec. 191 (here restricted to Hamilton Harbor, Bermuda; holotype, BM[NH] 1966.48.6) Columbella (Seminella) catenata Tryon, 1883 (not Sowerby, 1844). Man. Conch. 5: 179; pl. 58, figs. 51-55 Shell moderate in size (7 - 10mm in length) ; spire slight- ly less than 4 shell length, whorls slightly convex, shoul- dered, suture impressed; body whorl cylindrical, aperture moderately broad, apertural lip denticulate interiorly, col- umella slightly bent, weakly denticulate, siphonal canal very short, anal sinus distinct; sculpture cancellate on first 2 postnuclear whorls, spiral sculpture almost completely Page 124 THE VELIGER Vol. 20; No. 2 obsolete on later whorls where low axial ribs predominate. A single spiral subsutural groove persists to the most re- cently formed whorl; ground color from light brown to purple black with spiral rows of irregular white blotches below and above the suture. Each radular row consists of a flat, subrectangular median tooth flanked on each side by a bi- or tricuspid sigmoid lateral tooth (Figure 36). Remarks: The validity of this species has been in doubt ever since Reeve’s original description as a result of his poor figure and his failure to give a type locality. There is little doubt in my mind, after having examined a photo- graph of the holotype in the British Museum (Natural History), that this name must be applied to a small, some- what variable species from Bermuda. Although often considered a variety of Anachis sparsa Reeve, 1859, a species with a much greater geographic range, Costoanachis scutulata differs from that species in the general appearance of its aperture (especially its distinct anal sinus), its lower, less prominent sculpture, its unusual subsutural groove, its distinctive color pattern, and in minor radular distinctions. Furthermore, A. sparsa and C. scutulata, although occurring together in several Bermudan localities, have remained distinct, thus appar- ently ruling out the possibility of interbreeding. No fossil examples of this species have been reported. Range: Apparently endemic to the Bermuda Islands. Costoanachis semiplicata (Stearns, 1873) (Figure 14) Anachis semiplicata STEARNS, 1873. Proc. Acad. Nat. Sci. Philadelphia 25: 344 (Tampa Bay, Florida; lectotype, US NM 54275) Anachis avara Tryon, 1883 (not Say, 1822). Man. Conch. 5: 159; plt. 55, fig. 68 only Shell small to moderate in size (8- 15mm in length) ; spire slightly more than 4 shell length, whorls almost flat- sided, suture shallow; body whorl narrow, aperture mod- erately wide, apertural lip slightly thickened, distinctly denticulate within, columella straight, weakly denticulate; sculpture of a few widely spaced low axial ribs limited to the body whorl, all spire whorls smooth, colored light green or yellow-grey. Each radular row consists of a flat, rectangular median tooth flanked on each side by a sig- moid, distally bicuspid lateral tooth (Figure 37). Remarks: ‘This species has been considered a variety or subspecies of Anachis avara, a species that occurs on the east coast of the United States as far south as Matecumbe Key, Florida, whereas Costoanachis semiplicata occurs in the Gulf of Mexico and on the west coast of Florida south to Cape Romano. Several shell differences and certain minor radular distinctions serve to differentiate these 2 species. Costoanachis semiplicata has a longer, narrower shell form, with fewer, more widely spaced axial ribs than those of A. avara. Another form of C. semiplicata from the Gulf of Mexico has many, more closely spaced axial ribs. This form has often been called A. similis Ravenel, 1861, a name of dubious identity as it was never figured by its author, and its holotype, along with the rest of Ravenel’s types was presumably destroyed during the U. S. Civil War. No fossil examples of this species have been reported. Range: Boca Grande, southwestern Florida, along the shores of the Gulf of Mexico to Progreso, Yucatan, Mexico. Costoanachis sertulariarum (Orbigny, 1839) (Figure 73) Buccinum sertulariarum Orxicny, 1839. Voy. Amér. -Merid., Atlas, plt. 61, figs. 13-17 (La Baie de San Blas, Patagonie; holotype, BM[NH] 1854.12.4.451) Columbella sertulariarum Orpicny, 1841. Voy. Amér. Mérid., text: 431 Columbella avara Koxet, 1874 (not Say, 1822). Nachr. Malac. Gesell.: 59 Columbella brasiliana Martens, 1897. Arch. Naturgesch. 1 (2): 171-172; plt. 16, fig. 10 (Desterro, Brasil; holo- type, BM) Moderate in size (9- 12mm in length); spire about 4 shell length, acute, whorls flat-sided, suture shallow; body whorl cylindrical, aperture moderately wide, interior of Explanation of Figures 29 to 37 Figure 29: Anachis scalarina — radular dentition Figure 30: Anachis lyrata — radular dentition Figure 31: Costoanachis avara — radular dentition Figure 32: Costoanachis fennel — radular dentition Figure 33: Costoanachis lafresnayi — radular dentition Figure 34: Costoanachis hotessieriana — radular dentition Figure 35: Costoanachis floridana — radular dentition Figure 36: Costoanachis scutulata — radular dentition Figure 37: Costoanachis semiplicata — radular dentition Tue VE IcER, Vol. 20, No. 2 [Rapwin] Figures 29 to 37 Figure 30 Figure 29 Figure 31 Figure 33 Figure 35 Figure 36 Figure 37 Vol. 20; No. 2 THE VELIGER Page 125 apertural lip denticulate, columella non-denticulate, slightly bent anteriorly; sculpture of sharp axial ribs with microscopic spiral scratches between them limited to the body whorl. Color offwhite with random fine brown punctations and larger purple-brown blotches. Each radu- lar row consists of a flat subrectangular median tooth, flanked on each side by a sigmoid lateral tooth with 2 sharp distal cusps and a more rounded proximal projec- tion (Figure 39). Remarks: The only specimen of Costoanachis sertulari- arum from the Orbigny collection in the British Museum (Natural History) is considered the holotype. It must be noted here that the dimensions of the BM(NH) speci- men are significantly greater than those given by Orbig- ny and the type locality is considerably further south than any locality record in the collections examined. Misidentification of this species is almost certainly re- sponsible for the periodic assertions that Anachis avara Say occurs on the Atlantic coast of south-central South America (KoBeExtT, 1874; Prrspry, 1898; EYERDAM, 1950; Paropiz, 1962). Although these species are similar in general form, the spire whorls of Costoanachis sertulari- arum are less convex, its axial ribs are more poorly de- fined, its columella is not denticulate as in A. avara, and there are minor differences in color pattern and radular morphology. Marcus & Marcus (1962) have described and figured the anatomical features of Anachis brasiliana von Mar- tens, a synonym of Costoanachis sertulariarum. According to them Orbigny’s species has a wider distribution, a larger shell and slightly different body coloration. Al- though these assertions may have a bearing on the dis- crepancy between Orbigny’s type and other specimens we have seen, the fact is that we have seen no other speci- mens that agree in size and locale with the type. Un- fortunately no living specimens of this species have been available for examination. There appears to be a close relationship between this species and Costoanachis floridana (Rehder, 1939). The primary differences are that C. floridana is, on the aver- age, significantly smaller and has a broader apical angle and heavier body whorl, both of which impart a stouter appearance to it. The ranges of the 2 (C. floridana — east coast of the United States from North Carolina to southern Florida and on the Texas coast; C. sertulariar- um -— northern Brazil to southern Argentina) do not overlap to my knowledge. No fossil examples of this species have been reported. Range: East coast of South America from Rio Grande do Norte, Brazil to Mar del Plata, Argentina. Costoanachis sparsa (Reeve, 1859) (Figure 70) Columbella sparsa REEVE, 1859. Conch. Icon. Columbella, plt. 31, figs. 200a, 200b (here restricted to St. Thomas, Virgin Islands; holotype, BM[NH] 1966484) Columbella (Seminella) catenata Tryon, 1883. (not Sowerby, 1844). Man. Conch. 5: 179; plt. 58, figs. 51-55 Shell moderately large (8 - 11mm in length) ; spire mod- erately high (4 shell length), acute, whorls slightly con- vex, shouldered suture moderately deep; body whorl fus- oid, aperture narrow, apertural lip slightly thickened, denticulate interiorly, columella straight, denticulate, si- phonal canal short, slightly bent, anal sinus present; sculp- ture of prominent axial ribs with broad spiral grooves strongest between the ribs; color variable, generally with alternating evenly-spaced squares of orange-brown on a white background. Each radular row consists of a flat, rectangular median tooth, flanked on each side by a sig- moid lateral tooth with 2 sharp distal cusps and one blunt proximal projection (Figure 38). Remarks: This is the most variable of all the species of Costoanachis, with a typical form as described above and other forms varying in shell proportion and color pattern. The sculpture and shouldering of each whorl are, never- theless, constant throughout the species. Although confused by many authors with Costoanachis catenata and C. scutulata, these 3 species are quite dis- tinct. Costoanachis catenata has a unique chain-link color pattern and a bulbous, translucent-white protoconch; C. scutulata differs consistently from the others in the weak- ness and limited extent of its sculpture and its prominent anal sinus; the ribs of C. scutulata are low, sinuous, often disappearing below the middle of the body whorl. The color pattern of C. scutulata also differs markedly from that of C. sparsa. Spiral grooves notably present in the latter species are absent in C. scutulata. No fossil examples of this species have been reported. Range: Bermuda and Lantana, Florida to Venezuela and (possibly) southern Brazil. Metulella Gabb, 1873 Metulella Gass, 1873. Proc. Acad. Nat. Sci. Philadelphia 24: 270 (type species by OD, Metulella fusiformis Gabb, 1873) Shell moderately large to large (10 - 25mm in length) ; spire high, acute, whorls flat-sided, suture squarely in- cised; body whorl fusoid, aperture narrow, strongly con- Page 126 stricted anteriorly, outer lip slightly thickened, strongly denticulate on its inner surface, columella straight, weakly callused and weakly denticulate, length of siphonal canal moderate to long; sculpture of strong, closely-spaced axial ribs, crossed by moderately strong spiral cords. Color white to yellow-white. Radular dentition unknown. Remarks: This genus is almost extinct as the number of species has declined from a high of 4 or 5 in the Pliocene to a single living species today. This living representative of an otherwise extinct genus, Metulella columbellata (Dall, 1889a), should present a unique opportunity to establish the relationship of Metulella to other fossil and Recent columbellid genera. Unfortunately, no live-col- lected specimens of the species are known. Metulella columbellata (Dall, 1889) (Figure 19) Metulella (Nassarina) columbellata Dau, 1889a. Bull. Mus. Comp. Zool. 18: 182 (off Cape Catoche, Yucatan. Mexi- co; holotype, USNM 93019) Shell moderately large (10 - 15mm in length) ; spire high, acute, whorls almost flat-sided, suture incised; body whorl fusoid, aperture narrow, strongly constricted anteriorly, outer lip slightly thickened, strongly denticulate on its inner surface, columella straight, weakly denticulate with a thin callus, siphonal canal moderately long; sculpture of strong, closely-spaced axial ribs crossed by moderately strong spiral cords; color white or yellow-white. Radular dentition unknown. Remarks: This rare species, the only Recent species of Metulella, has been encountered more frequently as more deepwater dredging is carried out in the Gulf of Mexico. Unfortunately, the position of Metulella must await the description of its anatomy and radular dentition. The earliest fossil Metulellas are known from the Mio- cene of the Dominican Republic (Metulella venusta THE VELIGER Vol. 20; No. 2 Sowerby, M. williamgabbi Maury, and M. fusiformis Gabb). Range: Gulf of Mexico (specimens have been seen from Cape Catoche, Yucatan, Mexico — type locality — and off Tampa Bay, Florida. Nassarina Dall, 1889 Nassaria (Nassarina) Dauz, 188g9a. Bull. Mus. Comp. Zool. 18: 182 (type species by OD, Nassarina bushi Dall, 1889a) Pyramimitra (Nassarina) CoSsMANN, 1901. Essais Paléoconch. Comp. 4: 128 Nassarina Dall. Wooprine, 1928. Carnegie Inst. Wash. Publ. 385: 279 Shell small (3 - 1omm in length), fusiform; spire moder- ately high, acute, whorls convex, suture impressed; body whorl approximately 4 of shell length, outer apertural lip denticulate on its inner surface, aperture long, narrow, constricted anteriorly to form a narrow, bent siphonal canal, columella slightly bent anteriorly, its thin, de- tached callus forming a prominent keel; sculpture of numerous swollen axial ribs crossed by strong spiral cords. Each radular row consists of a flat, subrectangular medi- an tooth flanked on each side by a sigmoid, distally bi- cuspid lateral tooth. The cusps of the laterals are shorter and less bent than those in many other genera. Remarks: ‘The shell characters by which this genus may be differentiated from other columbellid genera are: 1) possession of both strong spiral and strong axial sculp- ture, 2) the constricted and elongate siphonal canal, and 3) the keel formed by a thin detached parietal callus. The genus Zanassarina Pilsbry & Lowe, 1932, includes g small eastern Pacific nominal species. It was originally erected as a subgenus of Nassarina Dall. No currently known Recent western Atlantic columbellid species are assignable here. Its affinities seem to be closer to Costo- anachis and Parvanachis than to Nassarina. Explanation of Figures 38 to 47 Figure 38: Costoanachis sparsa — radular dentition Figure 39: Costoanachis sertulariarum — radular dentition Figure 40: Nassarina bushii — radular dentition Figure 41: Nassarina glypta — radular dentition Figure 42: Nassarina metabrunnea — radular dentition Figure 43: Parvanachis isabelle: — radular dentition Figure 44: Parvanachis obesa — radular dentition Figure 45: Parvanachis ostreicola — radular dentition Figure 46: Steironepion minor — radular dentition Figure 47: Steironepion monilifera — radular dentition THE VELIcER, Vol. 20, No. 2 [Rapwin] Figures 38 to 47 Figure 38 Figure 40 Figure 43 Figure 45 Figure 46 <- ? : 25 ae A = f é \ Fy Z = ‘ r 5 i ‘ { in ie { \ a ¢ ~ a j ( 2 7 oo > a ~ y te = 3 aegis 4 ‘ * ' a ee j * Bf + sf - — . ry , * 4 f f 7 . ci = << = SSIS ~ a ——e = = 2 man sian ann 10 footcandles = 108 lumen per m’) from approximately 06:15 to 20:15 (Pacific Daylight Time). After a 24-hour acclimation period in the observation chamber, the snails were observed every hour for 5 days. Since active snails were almost always on the surface of the sand and buried snails rarely moved, the number of snails on the surface was used to quantify the activity of the Olivella population. Figure 1 is a summary of the numbers of Olivella recorded on the surface at one-hour intervals during the 5 days. The most striking feature shown by Figure 1 is that the Olivella were on the surface and active primarily at night. Fully 88% of the observations of animals on the sur- face were recorded during the hours of darkness (21:00 through 06:00). Consistently, the number of animals on the surface began to increase at twilight and rapidly reached a maximum, usually within 2 hours of darkness. Similarly, each night the number of animals began to decrease long before the light of dawn. While activity was 50 40 8 § A 3° g 6 i=] Z 20 z 10 fa) THE VELIGER Vol. 20; No. 2 clearly greatest during the night, it should be noted that some activity was also recorded during each 14-hour day- light period. Since the numbers of Olivella on the surface began to decrease well before dawn and to increase somewhat before complete darkness, it was desirable to directly test the snails’ responses to light and to dark. One hundred Olwella, which had been in the observation chamber for the 2 previous days, were exposed to complete darkness at 14:00 by covering the chamber with aluminum foil. ‘Two hours later, the cover was removed exposing the animals to ambient, late afternoon lighting conditions (go footcandles, ~ 972 lumens per m?). The results of this experiment are reported in Table 1. It can be seen from Table 1 that before darkening the chamber only 1 of the 100 Olivella was on the surface. However, after 2 hours of darkness artificially imposed during the day, 53 of the Olivella had emerged and were on the surface. Re-exposure to daylight produced an immediate burial response. Within 2 minutes, over half of the animals had completely buried themselves, and, within 8 minutes, all of them had disappeared under the sand. Light, and the absence of light, clearly has a direct influence on the activity of Olivella. 8 12 16 20 24 4 8 12 16 20 24 4 8 12 16 20 a4 4 8 12 16 20 24 4 8 12 16 20 24 4 8 Time of Day Figure 1 Numbers of Olivella biplicata on the surface (N=57) at 1-hour intervals through 5 complete light/dark cycles. Snails were main- tained outdoors and were exposed to daylight (> 100 Lux or > 10 footcandles) from approximately 06:15 to 20:15 (Pacific Daylight Time). A snail was recorded as being on the surface if 4% or more of its shell length was visible. Vol. 20; No. 2 Table 1 Responses of Olivella biplicata to darkness artificially imposed during the day (N = 100), and responses of snails on the surface to daylight (90 footcandles) THE VELIGER Number on Operation Time Surface Chamber darkened 14:00 1 Chamber exposed to daylight 16:00 53 16:02 20 16:04 8 16:06 3 16:08 0 16:10 0 16:20 0 Activity Patterns of Individuals The general activity pattern of the Olivella population (Figure 1) was relatively constant from one 24-hour peri- od to the next. In contrast, the activity patterns of individ- ual snails were extremely variable, and the records of 10 individual Olivella are presented in Figure 2 to illustrate this diversity. The median number of times that asnail was recorded on the surface during the 120 hours of obser- vation was 24 times. The activities of many individuals, Page 139 however, varied widely from this median value. For ex- ample, snail #7 was recorded on the surface 53 times (7. e., during almost half of the observations), whereas snail #56 was observed on the surface a total of only 3 times during the 5 days. The percentage of an individual’s activity that occurred in the dark or in the light was also variable. Although most of the snails were active primarily or exclusively at night (e.g., nails #10, 23, 25, 29, 32, 46, and 56), several of the snails had activity patterns that were characterized by substantial percentages of daytime activity (e. g., snails #7, 28, and 31). Even among the snails that were active primarily at night, the specifics of the activity patterns varied considerably from individ- ual to individual. For example, snail + 29 was not seen on the surface at any time during 2 of the 5 nights of ob- servation, but it was very active during the other 3 nights. In contrast, snail #46 was out on the surface every night, but it stayed out each night for only a short time. All of the Olivella were observed on the surface at some time during the 5 days. While observations at 1-hour intervals gave an ade- quate picture of an individual’s general activity cycle, an hour was a long time compared to the time actually needed for an Olivella to bury itself and re-emerge. Was it likely that an animal observed on the surface at 22:00 and again at 23:00 had spent the entire hour on the sur- face? Or had it perhaps buried itself and re-emerged several times during the hour? To answer these questions, Time of Day mm A eh Oo fe i ey BH ne a? #10 #3 #25 #28 #29 #31 # 32 #46 #56 Activity of Individual Snails m me mm OO 1 Te) oy O te yh Cy EP GT ey fee gts} vy § 10 3 6b O (9 He eye © Time of Day Figure 2 Activity records of 10 Olivella biplicata selected to illustrate the diversity of individual activity patterns. Observations were made at 1-hour intervals for 5 complete light/dark cycles. Darkened circles indicate the times that each individual was observed on the surface. Page 140 the 57 numbered Olivella were observed continuously for one hour (23:00 to 24:00) with data being recorded every 5 minutes. The results were clear. Of the 23 indi- viduals that were on the surface both at the beginning of the observation period (t = omin.) and at the end (t= 6omin.), 20 were on the surface for the entire hour. Of the 35 individuals that appeared on the surface at least once during the hour, only 3 individuals were observed on the surface, then not seen, and later observed again on the surface; furthermore, 2 of these 3 were not seen for only 1 observation, suggesting that they might have been on the surface but overlooked. Almost all of the snails that were observed on the surface were active and moving. In addition to showing that an individual observed on the surface for 2 consecutive hourly observations was like- ly to have been on the surface throughout the hour, these short-interval observations provided an important control for possible effects of the red light used to make all of the observations. Table 2 reports the total numbers of Olivella on the surface at 5-minute intervals during the hour of observations. There was no significant difference between the numbers of Olivella on the surface at the beginning and at the end of the hour (x?=0.32; p >0.5). A slight decrease was observed; however, this decline was consistent with the general activity cycle of Olivella (Fig- ure 1) in which activity often appeared to decline between 23:00 and 24:00. If there was a response to the red light, it was clearly unlike the strong burial response to day- light that was reported in Table 1. While it is never pos- sible to completely prove “no effect,” the conclusion ap- Table 2 Numbers of Olivella biplicata (N = 57) on the surface recorded at 5-minute intervals during one hour of continuous observation (23:00 to 24:00) Time Number of snails (Min. after 23:00) on surface 0 31 5 31 10 31 15 32 20 32 25 33 30 30 35 31 40 30 45 29 50 29 55 27 60 27 THE VELIGER Vol. 20; No. 2 pears justified that the red light did not significantly affect the basic activity pattern of Olivella. Relationships Between Shell Length, Sex, and Activity Olivella were collected for the preceding experiments without regard to sex or shell length. After the experi- ments, individuals were measured and sexed. Shell length (apex to siphonal canal) was determined with vernier calipers. The lengths ranged from 18.2 to 27.9mm (X = 22.7; SD = 2.4). Sex was determined by using the pres- ence or absence of a penis as the test criterion. Using this criterion, the sample of 57 Olivella contained 38 females and 19 males. Eleven of the males had functional penes; 8 had rudimentary penes, probably resulting from trema- tode infections. All animals designated as females were examined under the dissecting microscope to make certain that a penis rudiment had not been overlooked. The total number of times an individual was observed on the sur- face was taken as a relative measure of its overall activity. These relative activity measures ranged from 3 to 57 times on the surface during the 5-day experiment (X = 25.4; SD = 12.4). In Figure 3, shell lengths were plotted against the num- ber of times a snail was observed on the surface (7. e., activity). Small snails were generally more active, and activity gradually decreased with increasing shell length. The negative correlation between shell length and activity was highly significant when both males and females were included in the analysis (corr. coeff. =- 0.52; d. f= 55; p 0.5). The sample linear regression for all of the data (males plus females) was expressed by the equation Y = 25.2 -0.1X. Males and females were plotted with different symbols in Figure 3, and it was immediately apparent that males in this sample were considerably larger than females. Males had shell lengths averaging 25.3mm (N= 19; SD = 1.5), whereas the average shell length of a female was 21.4mm (N=38; SD=—1.5). The difference be- tween the sample means was highly significant (t-test; Pp <0.001). Since females were generally smaller than males, and since smaller snails were more active, it was expected that females would be more active than males. This was true. Females were observed on the surface an average of 29.5 times during the 5 days (SD = 11.6) compared to only 17.0 times (SD = 8.9) for males. The difference between Vol. 20; No. 2 Shell Length (mm) THE VELIGER Page 141 Number of Times on Surface Figure 3 Relationship between shell length and the total number of times a snail was observed on the surface during the 5-day experiment (t. €., activity). Males (CE) and females (A) are indicated separately. The correlation is significantly negative when both males and females are included in the analysis (corr. coeff. = -0.52; d.f. = 55; p< 0.001). The sample linear regression is expressed by Y = 25,2-0.1X, the sample means was again highly significant (t-test; pDwarps (op. cit.) emphasized activity taking place during day- light hours; in contrast, the present study emphasized the light/dark cycle of activity, and consequently, activity taking place at night. Edwards’ experiments were also con- ducted indoors with lighting provided by a nearby win- dow. He reported that for a short time each morning, the animals were exposed to direct sunlight through this window, and for the rest of the day, they were in shadow. Considering the day as a whole, he concluded that large snails were generally more active than small snails; he added, however, that during the short exposure to direct sunlight, smaller animals were more active and were found more often on the surface than were larger snails. Edwards’ results in bright light are in agreement with the conclusions of the present study, which was conducted under relatively bright, outdoor conditions. While the present study on Olivella concentrated spe- cifically on the behavioral effects of a light/dark cycle, many factors in addition to light clearly influence this snail’s activity. For intertidal snails, the tidal cycle is prob- ably the most important of these. Olivella which are ex- posed by the receding tide immediately bury themselves in the sand (STOHLER, 1969) ; they remain dormant until the water returns, regardless of the time of day or night (Phillips, unpubl. data). STOHLER (op. cit.) also reports that similar responses to tidal conditions may occur even when the animals remain covered by several decimeters of water. The presence of food also influences activity; if a piece of crab or mussel meat is placed on the surface of the sand during the day, many of the buried Olivella will suddenly emerge and begin searching for the food (Phil- lips, unpubl. data). Mating condition may influence activ- ity; sometimes during experiments, a male would emerge from the sand to begin courting a particular female that passed by on the surface. Many more factors, such as the Vol. 20; No. 2 THE VELIGER Page 143 amount of turbulence and the presence of potential pred- ators, are probably also important. So, while the light/ dark cycle clearly exerts a major influence on the behavior of Olivella, the activity patterns of this snail are complex and intricately related to a variety of factors. SUMMARY 1. The activities of individually marked Olivella biplicata were monitored every hour for 5 complete light/dark cycles. The animals were maintained in a plexiglass trough that was supplied with running seawater and located outdoors. 2. The Olivella were active primarily at night. During the day, most of the snails remained buried in the sand and rarely moved; at twilight, they began to emerge from the sand, and, within 2 hours of darkness, maximum numbers of snails were active on the surface. 3. While the general activity cycle of the Olzvella popu- lation was well-defined and predictable, the activity patterns of the individuals that made up the population were quite variable. A portion of this variability was related to differences in size and sex. 4, Females in this sample were considerably smaller than males (t-test; p < 0.001). 5. Small snails were generally more active than large snails (correlation coefficient —-0.52; d.fi =55; p <0.001). 6. Females were more active than males (t-test; p< 0.001). 7. Field observations during the day and night confirmed the existence of Olivella’s activity cycle. ACKNOWLEDGMENTS All of this work was performed at the Bodega Marine Laboratory of the University of California. I wish to thank the Director, Dr. Cadet Hand, for making these facilities available. In addition, I conducted some pilot experiments on the activity of Olivella biplicata several years ago at the Hopkins Marine Station of Stanford University. Although none of these pilot experiments is reported in this paper, some of my ideas originated there. I wish, therefore, to acknowledge Dr. Donald P. Abbott and the Director, Dr. John H. Phillips, for their past assistance and support. The field observations of subtidal Olivella were made by Mr. Harvey D. Van Veldhuizen. Literature Cited Cottam, CLARENCE 1939. Food habits of North American diving ducks. Agric., Tech. Bull. 643: 139 pp. Epwarps, Darras Craic 1968. Reproduction in Olivella biplicata. The Veliger 10 (4): 297 to 304; plt. 44; 3 text figs. (1 April 1968) 1969a. Predators on Olivella biplicata, including a species-specific pred- ator avoidance response. The Veliger 11 (4): 326-333; pit 51; 1 text fig. (1 April 1969) 1969b. Zonation by size as an adaptation for intertidal life in Olivella biplicata. Amer. Zool. 9: 399 - 417 GrINNELL, JoszpH, Harorp C. Bryant & Tracy I. STorER 1918. The game birds of California. Univ. Calif: Press, Berkeley: x+642 pp.; illust. Iverson, Ernest W. 1972. New hosts and bathymetric range extension for Colobomatus embiotocae (Crustacea, Copepoda). Calif. Fish & Game 58: 323 - 325 JoxuNson, RALPH GORDON 1971. Animal-sediment relations in shallow water benthic communi- ties. Marine Geol. 11: 93 - 104 Keen, A. Myra 1937. An abridged checklist and bibliography of west North American marine mollusca. Stanford Univ. Press, Stanford, Calif: 87 pp.; U.S. Dept. 3 text figs. (29 September 1937) Martin, A. C. & EF M. UnLeR 1939, Food of game ducks in the United States and Canada. U. S. Dept. Agric. Tech. Bull. 634: 156 pp. ReeDeErR, WILLIAM G. 1951. Stomach analysis of a group of shorebirds. 43-45 SToHLER, RupoLF 1959, Studies on mollusk populations: IV. The Nautilus 73: 65 - 72 1960. Studies on mollusk populations: IV. The Nautilus 73: 95 - 103 1962. Preliminary report on growth studies in Olivella biplicata. The Veliger 4 (3): 150-151; plt. 36 (1 January 1962) 1969. Growth study in Olivella biplicata (Sowerby, 1825). The Veliger 11 (3): 259-267; 1 map; 1 text fig. (1 January 1969) Turner, Cuares H., Bart E. Esert & Ropert R. Given 1969. Man-made reef ecology. Calif. Fish & Game, Fish Bull. 146; 1-221; 74 text figs. ZeELL, CLARACE PLumMB Bock 1955. The morphology and general histology of the reproductive sys- tem of Olivella biplicata (Sowerby), with a brief description of mating behavior. M.A. thesis, Univ. Calif, Berkeley. Condor 35: Page 144 THE VELIGER Vol. 20; No. 2 Size and Age-Specific Predation by Lunatia heros (Say, 1822) on the Surf Clam Spzsula solidissima (Dillwyn, 1817) off Western Long Island, New York BY DAVID R. FRANZ Biology Department, Brooklyn College, Brooklyn, New York I1I210 (5 Text figures) INTRODUCTION THE POTENTIAL IMPORTANCE Of the naticid gastropod Lu- natia heros (Say, 1822) as an important predator on the surf clam Spisula solidissima (Dillwyn, 1817) is recog- nized (Ropes, CHAMBERLIN & MERRILL, 1969), but there are little quantitative data on rates of predation or specific predator-prey relationships. Spisula is a commercially val- uable species and the absence of this information is detri- mental to the protection and management of surf clam populations. Part of the problem lies in the difficulties in the quantitative sampling of Spisula populations and in the low density of S‘pisula in natural habitats. The com- mercial hydraulic dredge, which is the standard sampling device for surf clams, does not capture the small individu- als, which are washed through the 2-inch (50mm) rings. In this report, data’are presented on the size and age structure of Spisula eaten by Lunatia heros. The problem of sampling small individuals was partially avoided by the analysis of beached animals and valves. Disadvantages of this approach include: a) no information on the den- sity of living Spisula is possible; b) the possibility exists that size/age structure of beach shells may not reflect the population structure offshore due to differential rates of accumulation and destruction; c) the possibility that beach shells originate from areas far distant and therefore do not reflect the age structure of the adjacent nearshore population. The major advantages of the approach taken in this study are the easy availability of size classes of Spisula which are not usually captured with commercial gear, and the possibility of collecting large numbers of bored Spisula valves which are generally not available in dredged samples. Because of the location and character of the study area — the extreme SW end of Long Island — some of the dis- advantages noted above are minimized. Information on the size structure of the adjacent nearshore populations of Spisula is available from a recent NMFS survey of inshore surf clam resources carried out by the author in 1974/1975. From these and other studies of the inner New York Bight, it is evident that the great majority of recent valves washed onto the Rockaway Beach are de- rived from the extensive populations of small surf clams which occur within 14 to 5km offshore. ACKNOWLEDGMENTS I wish to acknowledge the courtesy of Dr. Jack Pearce, NMFS, Sandy Hook, New Jersey Laboratory for allowing me to examine and measure a collection of surf clams from Twin Gun Beach, New Jersey. METHODS anp MATERIALS A. Study Area: Beach collections of recently washed-up clams were made at Rockaway Beach approximately 14km east of the Rockaway Point Jetty marking the extreme western tip of Long Island (Figure 1). The Rockaway Peninsula, which at this point is part of the Gateway National Rec- reation Area, is bounded on the south and west by the lower reaches of New York harbor. Extensive populations of Spisula occur directly offshore (south) and in the in- Vol. 20; No. 2 THE VELIGER Page 145 Jamaica Bay Rockaway Beach Collection Site ATLANTIC OCEAN Figure 1 Map of the Study Area Station 462, located 800m south of the Rockaway Peninsula, was sampled in August 1974 as part of an NMFS-sponsored survey of the inshore resource of surf clams off Long Island. shore waters to the east (off the Long Island coast) but B. Analysis of Bored Spisula: not to the west. These clams support a small bait clam fishery centered in nearby Brooklyn. This particular Although the boring snail Polinices duplicatus (Say, stretch of beach is famous for periodical, massive wash- 1822) occurs occasionally in dredge hauls, no living speci- ups of Spzsula following winter storms (JacoT, 1920). mens have been observed during 3 years of observations In fact, living clams are almost always available in small on this beach. On the other hand, Lunatia heros is ex- numbers on this beach. tremely abundant offshore and is frequently washed up Page 146 alive at all seasons. Moreover, the occurrence of Lunatia egg masses in the Spring indicates the abundance of this species nearshore. There is no doubt that this species is primarily responsible for the predation discussed below. Quantitative collections of bored Spisula valves were made in May, 1976. Every bored valve was collected at low tide along an approximately room stretch of beach, between the upper drift line and the water’s edge. All valves showing bore holes were collected and returned to the laboratory for analysis. On the same date, 4 1 m® quad- rats were sampled near the low water mark to ascertain the size-frequency distribution of all Spisula valves regard- less of the cause of death. The shell length corresponding to each observable growth ridge was measured for the 135 bored valves col- lected in May (see above). This procedure was repeated on 102 paired valves (7. e., one valve of an attached pair) in June, 1976. A similar analysis was carried out on 46 paired valves collected at the same location in July and August, 1976, with the exception that the latter sample was selected on the basis of the clarity of growth ridges, 7. é., valves were selected in which all growth ridges were discernible. RESULTS A. Size Distribution of Rockaway Beach Clams: West of Jones Inlet, inshore surf clams are generally smaller than those occurring further east (FRANZ, 1976). Figure 2A shows the length-frequency distribution of clams at station 463 off Rockaway Beach as determined in the National Marine Fisheries Service survey of Au- gust, 1974. Figure 2B shows the size distribution of the Rockaway Beach collections of both bored and unbored Spisula in May, 1976. There is general agreement in the size range and distribution of unbored beach clams and offshore clams from the NMFS survey with the exception of the absence of clams less than 85 mm in the latter. This is probably due primarily to the inability of the commer- cial hydraulic clam dredge which was used in the 1974 survey to retain smaller clams. Clams present on the beach in May are considered to be a reasonable sample of the adjacent inshore (800m) population. Figure 2B also indicates that over 80% of the bored valves are less than 80mm, Although some bored valves were observed up to about 140mm, B. Possible Age Distribution: In order to determine the age distribution of the bored clams, it is necessary to estimate the age of clams from THE VELIGER Vol. 20; No. 2 A 407 Station 462 - 800m IK | off Rockaway Beach / 30+ August 1974 | \ N=242 | 20- | \ | | \ | / | 25 -10- \ | | i\ | \ \ lisse! elo aa?) | 20- O- ees PRN / i \Rockaway Beach 5 B | | ; 8157 / Bored . \ Collection < 7 Ses \ | ee 1976 ry aie 2 ae ED 35) DD 75) 195) (0015) RSS Length, mm Figure 2 A. The size-frequency distribution of adult clams collected at station 462 off Long Island, August 1974 B. Size-frequency distribution of bored and unbored Spisula valves from Rockaway Beach, New York, collected in May, 1976 size-frequency data. At present, the growth rates of Spisu- la are uncertain. Growth curves of S. solidissima have been proposed by various workers (BELDING, 1910; WEST- MAN, 1946; YANCEY & WELCH, 1968; LoEscH, 1975) based on the analysis of presumed annual growth ridges on the shell. However, growth estimates of one-year ani- mals in all of the above studies are significantly larger than those observed on the bored Rockaway Beach shells Table 1 Growth Statistics— Rockaway Beach Clams Ridge 0.05 Confidence No, N Mean (mm) s.d, Interval 1 46 20.22 2,449 0.730 2 45 37,00 5,543 1,652 3 46 50,56 6.684 1,992 4 46 68.19 6.177 1,842 5 46 81.46 7.092 2.114 6 44 92.89 8.627 3.988 7 31 104.64 7.026 3.582 8 5 115.0 3.937 4,722 ee _ NN ——e-a—SaS._——eeeee Vol. 20; No. 2 7 \N=31 N=44 / 5 Nee 30 4 \N=46 Shell length, mm Figure 3 Frequency distribution of shell lengths associated with growth ridges from a suite of paired valves collected on Rockaway Beach in July and August 1976 me WELCH, 1963a LogscH, 1975 Rockaway Beach - 1976 20 Hey See SO 7-819 Ridge Number THE VELIGER Page 147 in the present study. Therefore, a separate analysis of the growth ridges of the Rockaway shells was carried out. It was possible to observe and measure the first 8 growth ridges with relative ease. These data are presented in Table 1 and the frequency distribution of each ridge group (one through eight) is shown in Figure 3. These data are compared in Figure 4 with a growth curve proposed by A. WEtcH (1963) for a New Jersey population and cor- rected recently by Lorscu (1975). It is clear from these data that the observed growth rates of the Rockaway clams are significantly lower than the rates observed in the other population. Using the above information on size and presumed age, the probable age of both bored and unbored samples was determined. This indicates (Figure 5) that maximum pre- dation occurred among 3 and 4 year clams, and dropped precipitously in clams older than 5 years. @ Bored, N=135 © Unbored N=166 Percent S a —d x CAT EZR B dog} Ge (7°78 Ridge Number Figure 5 Age distribution of bored and unbored Rockaway Beach valves of Spisula (< adjacent column) Figure 4 Growth curves corresponding to ridge numbers 1-8 based on analysis of paired Rockaway Beach clams (lower curve). Vertical bars represent 95% confidence intervals of the mean. Upper curve represents comparable portion of a growth curve for offshore New Jersey clams based on WELCH (1963) as modified by Lorscu (1975) Page 148 DISCUSSION Evidence presented above supports the conclusion that the Rockaway Beach collection of surf clam valves are largely contributed from the adjacent inshore population of living Spisula. However, the size distribution of bored valves does not necessarily reflect the true size distribution of prey since there is a chance that very small bored valves are underrepresented on the beach. It is clear that predation by Lunatia heros is most in- tense on smaller clams (less than 80mm), and that this component of the population is less than 5 years old. How- ever, it is significant that older and larger clams are not completely immune to attack since bored valves of up to 160mm were occasionally observed. As emphasized by Epwarps (1974, 1976) the existence of an upper critical prey size, beyond which the probability of predation is small, may have been an important component in the evolved reproductive strategies of the bivalve prey of naticid predators. Since it is likely that reproductive output of bivalves such as Spisula increases exponentially with increasing body size, the reproductive value of larger individuals, i.¢., those which are by virtue of their size relatively immune to attack, is likely to be significantly greater than the reproductive value of smaller animals. This is a factor for consideration in management decisions regarding the establishment of size limits for harvesting surf clams. It may make better sense to harvest populations of predominantly smaller clams -— for which man must compete any- how with Lunatia - and conserve populations of predominantly larger clams which are less subject to predation. Inshore populations of Spisula along Long Island differ in size structure. West of Jones Inlet, clams are generally smaller, and the component of very large clams is absent. East of Jones Inlet, the reverse is true, 7. e., populations are composed of very large clams with relatively few indi- viduals less than 100mm (FRANz, 1976). There are sever- al alternate explanations for this including differences in settlement and survival of juveniles, differences in fishing intensity, etc. To these must be added the possible role of Lunatia. Since predation pressure by Lunatia is maximal on small clams, populations of predominantly small to medium clams (less than 1oomm) may suffer greater THE VELIGER Vol. 20; No. 2 predation than populations composed of predominantly larger animals. It is reasonable to assume that predator density would also be higher in areas, such as western Long Island, characterized by abundant numbers of smaller clams. If this is true, the absence of larger clams may simply reflect the small proportion of individuals surviving to the critical size of predation immunity. Like- wise, predator “carrying capacity” should be lower, off eastern Long Island, where most clams are relatively immune to Lunatia predation because of size. This re- duction in predator density should increase the probabili- ty that small clams can avoid predation and survive to the critical size. The importance of Lunatia as a consumer of surf clam production remains to be determined. In his recent ele- gant study of the ecology of Polinices duplicatus, based on field studies at all seasons, E>nwarps (1976) has reported that about 100 small Mya arenaria are consumed per individual predator per year. If we assume that L. heros harvests a comparable quantity of Spisula, each Lunatia would consume a quantity of clams which, had they reached commercially marketable size, would have been equivalent to about 7ol (2 bu.) of clams per year. Whether this is a reasonable estimate of the potential role of Luna- tia heros, and if so, whether this level of predation would constitute a significant proportion of Spisula production are problematic. However, Spisula shares with many tem- perate bivalves the characteristic of periodic recruitment. Although spawning occurs annually (Ropgs, 1968), sig- nificant recruitment does not. The average time span between successful recruitments of S‘pisula is not known but may be as much as 4 or 5 years or perhaps longer. Assuming that Lunatia harvest roughly 5% of prey standing stock per annum (e. g., see GREEN, 1968) and that successful recruitment occurs one year in 5, mortality due to Lunatia alone would approach 20% during the intervening years of minimal recruitment. During this time period, of course, other sources of mortality including fishing mortality would continue to occur. If fishing mor- tality continues at a constant rate during this same time interval, 7. e., 5% per annum, then standing stocks could decline as much as 40% due to the combined Lunatia and human predation. Under these conditions, the viability of remaining stocks could be endangered. Review of the size frequency distribution of surf clams from the mid-Atlantic region between 1965 and 1974 Vol. 20; No. 2 (RopEs, 1975) indicated that the components of small to medium clams (less than 125mm) comprise a small pro- portion of total clams in all areas surveyed (22% and 11% respectively). Since these are the clams most sus- ceptible to predation, it may be that predation is a prima- ry factor in accounting for skewed size distributions of Spisula populations. The apparently lower growth rate of inshore Rockaway clams relative to other populations, if it can be confirmed by more direct growth studies, is interesting in that it seems to support a suggestion by H. H. Haskin (personal communication) that inshore New Jersey clams are smal- ler, even stunted, compared with populations from deeper water. WESTMAN (1946) also noted that clams from off Long Beach (western Long Island) tended to exhibit sig- nificantly slower growth than populations further east (Jones Beach). The reasons for such differences remain to be discovered but could be related to increased sedi- ment instability and turbulence in shallow waters. The size of clams at the end of their first growing season (corresponding to ridge No. 1) ranged in this study from about 12 to 28mm with a mean of 20mm, a value fairly close to that reported in other studies. The lowest reported valueis 10mmat Boothbay Harbor, Maine (Yancey & WELCH, 1968). A sample of small Sprsula collected in the NMFS survey of inner New York Bight in February 1969 (station 48: PEARCE, 1972) averaged 18.1mm. Likewise, a large homogeneous sample of small Spisula from “Twin Gun Beach” near Sandy Hook, New Jersey, collected in November 1969, ranged from g - 25 mm with a modal! size of 17.5mm. Ropes & MERRILL (1970) reported a dense population of surf clams in Oc- tober 1964 averaging 21.1mm at Wallops Island, Virgin- ia, which averaged 22.5mm by 20 November of that year. Thus, data from a variety of sources indicate that O-class clams in their first season will attain an average size of 10- 25mm with the higher averages occurring further south, By the end of the first year, however, the Wallops Island clams noted above attained a length of 44mm (Ropes & MERRILL, op. cit.) which agrees also with Welch’s data as modified by Lorscu (1975). As seen in Figure 4, however, Rockaway clams at 1 year (7.e., ata size about halfway between ridge nos. 1 and 2) are still less than 30mm. Thus, the low growth rates of the Rocka- way inshore clams may reflect a lag in the initiation and rate of new growth in the Spring of their second growing season. THE VELIGER Page 149 SUMMARY A comparison of the size distribution of Spisula valves with bore holes of Lunatia heros with unbored valves in- dicated that Lunatia predation is largely limited to clams less than 80mm. An analysis of growth ridges on paired, beach-collected Rockaway clam valves indicated an esti- mated growth rate which is significantly lower than values predicted from other proposed curves available in the literature. Application of the growth information to the collections of bored Spisula valves showed that Lunatia tends to select clams under 5 years of age. These data suggest that although the adjacent subtidal populations of Spisula are susceptible to Lunatia predation, popula- lations of predominantly larger clams (> 100mm) oc- curring further east on Long Island are substantially immune to Lunatia attack. This suggests the possibility that size-specific predation by Lunatia may operate to maintain the smaller size structure of western Long Is- land populations of Spisula. Literature Cited Be.pinc, Davp L. 1910. The growth and habits of the sea clam (Mactra solidissima). Reprt. Comm. Fish, Game Mass. 1909, Publ. Doc. 25: 26 - 41 Epwarps, Davin Craic 1974. Preferred prey of Polinices duplicatus in Cape Cod inlets. Bull. Amer. Malacol. Union for 1974: 17 - 20; 1 text fig.; 3 tables. in press. Feeding and growth rates of Polinices duplicatus preying on Mya arenaria at Barnstable Harbor, Massachusetts. (MS under review, provided by author) Franz, Davin RicHaRD 1976. Distribution and abundance of inshore populations of the surf clam (Spisula solidissima) in the inner New York Bight off Long Island. Amer. Soc. Limnol. Oceanogr. Spec. Symp. 2, Middle Atlantic Cont. Shelf and the New York Bight: 404 - 413 GreEN, Rocer H. 1968. Mortality and stability in a low diversity sub-tropical intertidal community. Ecology 49: 848 - 854 JacoTt, ARTHUR PAUL 1919. | Some marine Mollusca about New York City. The Nautilus 32 (3): 90-94 (14 January 1919) Logscu, Joserx G. 1975. Invetitory of surf clams in near shore waters from Cape Hen- lopen to the False Cape area. Reprt. No. 4. Unpubl. Rprt. Fish. Managmt. Branch, State-Feder. Relat. Div., NMFS, Gloucester, Mass. Pearce, JoHN B. 1972. The effects of waste disposal in the New York Bight. Final Reprt. Sect. 2: Benthic studies. NMFS, Sandy Hook Labor. Unpubl. Rprt. Ropes, JoHN W. 1968. The reproductive cycle of the surf clam, Spisula solidissima, in offshore New Jersey. Biol. Bull. 135: 349 - 365 1975. Allowable surf clam harvest estimates — 1974. Unpubi. MS, NMFS, Oxford, Maryland, 10 pp. Ropes, JoHN W, J. Lockwoop CHAMBERLIN & ARTHUR S. MERRILL 1969. Surf clam fishery pp. 119-125 in: Frank E. Firth, ed.: The Encyclopedia of Marine Resources, Van Nostrand Reinhold Co. Page 150 THE VELIGER Vol. 20; No. 2 Ropes, Joun W. & ArtHuR S. MERRILL 1970. Marking surf clams. Proc. Natl. Shellfish. Assoc. 60: 99 - 106 WE cH, WatTeER R. 1963. Unpubl. Reprt. cited in Yancey & Wetcu, 1968 and LozscaH, 1975 (see below and above, respectively) Westman, James R. & Mitton H. Bmwe.ti 1946. The surf clam. Economics and biology of a New York marine resource. Unpubl. MS in files of NMFS, Oxford, Maryland. Yancey, Ropert & WALTER R. WELCH 1968. The Atlantic coast surf clam — with a partial bibliography. U.S. Fish & Wildlife Serv., Circ. 288, 13 pp. Vol. 20; No. 2 THE VELIGER Page 151 A New Species of Lithophaga (Bivalvia) from the Great Barrier Reef, Australia KARL H. KLEEMANN I. Zoologisches Institut der Universitat Wien, Austria (2 Plates) INTRODUCTION Durine 1974 I stupiep the ecology of burrowing bivalves (terminology after CarRIKER & SMITH, 1969: 1011) at the Great Barrier Reef (GBR), where I found 5 Litho- phaga species restricted to live coral (cf. OTTER, 1937; IREDALE, 1939). One of them is an undescribed species. The species are: Lithophaga lessepsiana Vaillant, 1865 Lithophaga lima Lamy, 1919 Lithophaga hanleyana sensu Gohar & Soliman, 1963 (non Reeve, 1857) Lithophaga simplex Iredale, 1939 Lithophaga kuehnelti Kleemann, spec. nov. DESCRIPTION or THE NEW SPECIES MYTILOIDA MYTILACEA MYyTILIDAE Lithophaginae Lithophaga Roding, 1798 (Leiosolenus) Carpenter, 1857 Lithophaga (Leiosolenus) kuehnelti Kleemann, spec. nov. (Figures rc, 2) Named in honor of Wilhelm Kihnelt with regard to his “Bohrmuschelstudien” I, II, and III (1930, 1933, and 1942, respectively). Holotype: BM(NH) Reg. No. 1976135; size: length, 28.4mm; height 9.0mm; breadth 8.6mm (Figure 1c). Paratypes: NHMW 80635 (dry), NHMW 80636 (for- malin), Vienna, Austria. Type Locality: Heron Island (Lat. 23°27’'S; Long. 151°55 E), Capricorn Group, GBR, Queensland, Austra- lia. Description: Shell small, thin, fragile, translucent, and smooth. Periostracum pale-yellowish, covered by more than one type of calcareous matter. The soft paste-like, non-adhesive deposits are on the dorsal part of the shell as well as on the corresponding part of the burrow. On the ventral part of the burrow, which is free of calcareous deposits, in a few cases a very fine longitudinal ridge can be observed, where the byssus is attached (this species does not rotate). Ventral margin straight, dorsal margin almost parallel; umbonal parts not quite at the anterior end, causing a slight depression in the lateral outline, otherwise both ends of shell rounded. The characteristic feature is the incrustation of the posterior ends of the valves, which is adhesive, rather hard, smooth, thickening towards the ends, and slightly protruding beyond them, where it generally terminates in a conspicuous manner, similar to concave lips of a mouth (Figure 2). Due to the individual position of the orifice on the coral surface and the angle of the burrow, these lips are modified and rarely equal on both valves of a specimen. Habitat: Lithophaga kuehnelti occurs in Acropora (Iso- pora) palifera (Lamarck) (Figure 3) at various sites off Heron Island and One Tree Island, both in the Capri- com Group, GBR. Host corals from well exposed sites (in terms of water movement) may be crowded by L. kuehnelti, without showing any harm (cf Woop-Jongs, Igt0: 127). An encrusting A. palifera (25X15 Xgcm) Page 152 from One Tree Island contained 150 specimens as well as 34 barnacles. Distribution: At the GBR the species is common in the subtropical part at Heron Island, One Tree Island, and Wistari Reef, all 3 in the Capricorn Group, and less com- mon to rare at the tropical Lizard Island. The local distri- bution pattern seems to be related to the water movement, but not to the available surface area of the host coral (KLEEMANN, in prep.). In the bivalve collection of the BM(NH) there is a sample of Acropora (Isopora) pali- fera collected by J. D. Taylor from Addu Atoll, Maldive Islands in May 1975 with 12 specimens of Lithophaga kuehnelti still in their burrows and 4 loose specimens (BM[NH] Reg. No. 1976134). In the coral collection of the British Museum (Natural History), the following samples contain L. kuehnelti: BM(NH) Reg. No. 1892. 6.8.50 and 51, both A. palifera from Capricorn Islands, GBR; BM(NH) Reg. No. 1887.1.29.5, A. palifera from New Guinea; BM(NH) Reg. No. 1884.12.11.17, A. pali- fera from Shortland Island, Solomon Islands. Other sam- ples of Acropora-Isopora were found infested by Litho- phaga, but without breaking up the corals I could not tell with certainty if the burrower was indeed Lithophaga kuehnelti. COMPARISON wir otuer SPECIES WITH WHICH IT COULD BE CONFUSED With the conspicuous posterior incrustation present, Lith- ophaga kuehnelti can easily be distinguished from other small Lithophaga species, such as L. lessepsiana Vaillant, 1865 (Figures zd, e; 4 a-d), with which it sometimes occurs in Acropora palifera. Lithophaga lessepsiana, described from Stylophora pistillata by VAILLANT (1865: Explanation Figure 1: a Lithophaga hanleyana (Reeve) sensu Gowar « SOLI- MAN (1963); b Lithophaga simplex Iredale; c Lithophaga kuehn- elti Kleemann, spec. nov., holotype (BM[NH] Reg. No. 1976135) ; d Lithophaga lessepsiana Vaillant (host coral Stylophora pistillata, Red Sea), part of BM[NH] Reg. No. 1871.9.8.53; ¢ Lithophaga lessepsiana Vaillant (host coral Heteropsammia michelini, Lizard Island, GBR). Scale in mm. THE VELIGER Vol. 20; No. 2 124) can be regarded as the first record of Lithophaga from live coral, although not actually stated as such. Lithophaga kuehnelti can be distinguished from L. han- leyana Gohar & Soliman, 1963 by the differences in the patterns of their posterior incrustations (Figures 2, 4, 5). Without its incrustation, L. kuehnelti is very similar to L. simplex Iredale, 1939; this species was determined with reference to Australian Museum Reg. No. C. 105 340 (part), as neither the type nor the paratypes could be investigated. The description and figure of L. simplex (IREDALE, 1939: 421; plt. 6, fig. 25) are inadequate and I consider it worth while to give more details and figures. Lithophaga (Leiosolenus) simplex Iredale, 1939 (Figures rb, 6a - 6d, 7) = L. cumingiana Otter, 1937, not Reeve, but only as described from living Favia (BM[NH] Reg. No. 1952.1.29.845-854; one specimen of this lot is shown in Figure 6a). Description: Shell thin, fragile, translucent, smooth, with faint growth lines, Periostracum pale-yellowish (col- our of the living animal yellowish fluorescent green, ex- cept the chalky layers). No real incrustation, but fine, thin, very smooth chalky layer, sometimes covering the whole surface, rarely completely absent, but more often showing bare areas; no tip or prolongation of the posterior end (Figure 8). Collected specimens reaching 32.4mm in length. The shell has a straight ventral margin, the dor- sal margin is generally angulated (Figures 6a to 6c), but is sometimes almost parallel to the former (Figure 6d). The position of the dorsal angulation is in most cases slightly anterior, rarely posterior to the middle of the of Figures 7 to 4 Figure 2: The conspicuous incrustation of the posterior end of Lithophaga kuehneltt. Scale in mm. Figure 3: Acropora (Isopora) palifera (Lamarck) in situ, showing orifices of burrows. Host coral of Lithophaga kuehnelti and rarely of Lithophaga lessepsiana. X1.5. Figure 4: Four specimens of Lithophaga lessepsiana; c and d are the same as in Figure 1d and re. Scale in mm. All photographs, except Figure 3, by P. Richens, courtesy of the Trustees of the British Museum (Natural History) THE VELIGER, Vol. 20, No. 2 [KLEEMANN] Figures 1 to 4 Seat Vol. 20; No. 2 shell. Siphons with dark pigmentation, edges of the mantle lighter coloured. Habitat: Lithophaga simplex was found in Porites and Symphillia (IREDALE, 1939); in Favia (OTTER, 1937) ; in Porites (BM[NH] Reg. No. 1976136). I found most of L. simplex burrowing in Favia (F pallida). Other host corals are Lobophyllia, Symphyllia, and Gontastrea reti- formis. The highest population density was observed in a living Favia (18 X 12 X 8cm), inhabited by 63 speci- mens. Generally infestation is much less, but Favia seems to be the preferred host at Lizard Island, GBR. The in- fested Porites samples I examined contained only L. han- leyana Gohar & Soliman (KLEEMANN & RosEN, in prep.) (Figures ra, 4, 5). Distribution: Low Island, GBR (OTTER, 1937; IRE- DALE, 1939) ; Palfrey Island (near Lizard Island) in Fa- via; Lizard Island, GBR, in Favia pallida, Symphillia, and Lobophyllia; Wistari Reef, Capricorn Group, GBR, in Lobophyllia; One Tree Island, Capricorn Group, GBR, in Goniastrea retiformis. In the bivalve collection of the British Museum (Natural History) there is one specimen, BM(NH) Reg. No. 1976136, collected by J. D. Taylor in December 1971 at Pungutiayu, Shimoni, Kenya (Ad- miralty Chart 998), which in my opinion is a L. simplex. DISCUSSION All Lithophaga species show considerable variation in shape and outline (Figures 4, 7; LYNGE, 1909: 138 (42) ; KLEEMANN, 1974: fig. 3, Lithophaga lithophaga Schalen- formen). Sometimes it is obviously influenced by the thick- ness of the substratum, e. g.. L. kuehnelti from branched upgrowing forms of Acropora palifera are more elongated than those from encrusting colonies of the same host coral. There is also a considerable difference in the relationship between length of the burrow and width of the orifice (KLEEMANN, in prep.). The geographical distribution of most subtropical and tropical Lithophaga species from dead as well as live coral is fairly wide, but there seems to be a rather distinct difference between Atlantic and Pacific (including the Red Sea) forms. For the encrusted Lithophaga species, in my opinion, the pattern of the calcareous deposits, especially of the posterior end, is the best characteristic for specific deter- THE VELIGER Page 153 mination (SoLiMAN, 1969: 889). Even there we should not be too dogmatic, for observations in the field and many samples have shown that it is impossible to find identical stereotyped copies of a pattern, as would be ex- pected from artificial products. Organisms have individ- ual responses to environmental conditions. The stated differences are not just variations of one species, corre- sponding to the host coral infested, for Lithophaga species appear with the same pattern in different hosts. Further- more, although rarely observed, one coral can be in- fested by more than one Lithophaga species, e. g., L. kuehnelti and L. lessepsiana in Acropora palifera, as stated above. GoHAR & SOLIMAN (1963: 67) stated “a specificity of the boring species to the coral attacked ob- viously exists,” and Sortiman (1969: 887) “... and furthermore to a particular coral species.” This seems to be the case only for L. kuehnelti, but not for any other Lithophaga species (KLEEMANN & RosEN, in prep.). Some host corals have been given only generic assign- ments; this list will undoubtedly be extended by further investigations. A full discussion of coral-living Lithophaga species and their hosts, with specific determination, is in preparation (KLEEMANN & RosEN, in prep.). SUMMARY Five Lithophaga species from the GBR, Australia, are restricted to live host corals. Lithophaga kuehnelti in- fests Acropora (Isopora) palifera (Lamarck) at subtropical and tropical sites of the GBR, and was also found on Addu Atoll, Maldive Islands. Lithophaga kuehnelti is compared with L. simplex Iredale, 1939, for which are presented an extended description, including figures, and more biological data. Lithophaga species are recorded from a number of host corals, and in rare cases two Litho- phaga species occurred in the same host. ACKNOWLEDGMENTS I am very grateful to the Australian Department of Education for a nine month award; to the “Fonds zur Forderung der wissenschaftlichen Forschung in Oster- reich” (Antrag No. 2123); to the Royal Society and the Austrian Academy of Sciences. I am indebted to the staff of the British Museum (Natural History), Section Mol- lusca. Page 154 THE VELIGER Literature Cited CarrRIKER, MELBOURNE Romaine # EpMuND Hosart Smit 1969. Comparative calcibiocavitology: summary and conclusions. Amer. Zool. 9 (3): 1011-1020; 1! table (August 1969) Gouar, H. A. F & Gamit N. Soiiman 1963. On three mytilid species boring in living corals. Publ. mar. biol. Sta. Al-Ghardaga; Red Sea (no. 12): 65-98; 28 plts.; 18 figs. IREDALE, Tom 1939. Mollusca, Part I. Great Barrier Reef Exped. 1928-29, Sci. Reprt. 5 (6): 209 - 425; 7 plts. KLEEMANN, Kart H. 1974, | Raumkonkurrenz bei Atzmuscheln. Mar. Biol. 26: 361-364 (in prep.) Distribution pattern of some Lithophaga (Bivalvia) from the Great Barrier Reef, Australia. KLEEMANN, Karu H. & Brian R. Rosen (in prep.) Burrowing bivalves and their host corals from the Great Barrier Reef, Australia. KiHNELT, WILHELM 1930. Bohrmuschelstudien I. Paldobiologica 3: 1-91 1933. | Bohrmuschelstudien IT. Paldobiologica 5: 371 - 408 1942. Bohrmuschelstudien III. Palaobiologica 7: 428 - 447 Lamy, Epouvarp 1919. Les lithodomes de la Mer Rouge. Bull. Mus. Nat. Hist. Nat. Paris 25; 252-257; 344 - 350 Lynce, H. 1909. The Danish expedition to Siam, 1899-1900. IV. Marine Lamelli- branchiata. K. dansk. Vidensk. Selsk. Skr., Raek. 7, 5 (3): 97 - 299; 5 plts.; 1 map Orter, G. W. 1937. Rock-destroying organisms in relation to coral reefs. Great Barrier Reef Exped. 1928-29, Sci. Reprt. 1 (12): 323 - 352; plts. 1-4 Reeve, Lovett AucustTus 1857-1858. Conchologica Iconica. Lithodomus. (plt. 1: March, 1858; pits. 2, 3, 4: October 1857; plt. 5: January 1858) Sotiman, Gamiv N. 1969. Ecological aspects of some coral-boring gastropods and bivalves of the northwestern Red Sea. Amer. Zool. 9 (3): 887 - 894; 5 figs. Woop-Jones, F 1910. Coral and Atolls. London 23: xxiii+392 pp.; 27 plts.; 1 map Explanation of Figures 5 to 8 Vol. 20; No. 2 Figure 5: Posterior end of a preserved specimen of Lithophaga hanleyana (Reeve) sensu Gouar & SOLIMAN (1963) in lateral view. Scale in mm. Figure 6: The conspicuous incrustation of the posterior end of Lithophaga hanleyana (Reeve) sensu GoHaR & Sotiman (1963). Scale in mm. Figure 7: Four specimens of Lithophaga simplex Iredale, showing variation in their outline; a part of BM[NH] Reg. No. 1952.1.29. 845-854; 6 same specimen as in Figure 1b. Scale in mm. Figure 8: Posterior end of Lithophaga simplex Iredale; same spe- cimen as in Figures 1b and 7b. Scale in mm. All photographs by P. Richens, courtesy of the Trustees of the British Museum (Natural History) Tue VELIGER, Vol. 20, No. 2 [KLEEMANN] Figures 5 to 8 te a ie yt Sa Vol. 20; No. 2 THE VELIGER Page 155 The Effects of Season on Visual and Photographic Assessment of Subtidal Geoduck Clam (Panope generosa Gould) Populations LYNN GOODWIN Washington State Department of Fisheries, Shellfish Laboratory, Brinnon, Washington 98320 (1 Plate) INTRODUCTION Tue WASHINGTON STATE DEPARTMENT of Fisheries rou- tinely conducts surveys of clam stocks in Puget Sound for management purposes by divers equipped with SCUBA. Geoduck populations [Panope generosa (Gould, 1850) | are estimated by visual counts of “shows” (either siphons or marks in the substrate made by siphons) along meas- ured transect lines (Goopwin, 1973). Earlier surveys demonstrated that the portion of the geoduck population detected by divers varied widely from 26% to 87% (Goopwin, 1973). Variability in “showing” has been reported in other clam species (FLOWERS, 1973). Geoducks live permanently buried in the substrate, the average burrow being 52cm deep in Hood Canal, Wash- ington (ANDERSEN, 1971). When the siphons are extended up to or above the substrate surface and the clams are actively pumping water, they are readily visible to divers. At other times the siphons may be withdrawn below the surface and the siphon holes filled with sand, mud and detritus, leaving no indication of the clam buried below. Frequently marks in the substrate are observed and the presence of geoducks can be verified by divers probing the depression with their fingers. The siphons are large, up to 8cm across and have a characteristic texture. The objective of the present study was to more precisely define how the percentage of the geoduck population detected by divers changes seasonally in 2 small subtidal plots. The information developed in this study is used to correct diver survey data to provide more accurate population estimates. The study area was a delta formed at the mouth of Big Beef Creek in Hood Canal, Washington. The delta has a gentle slope with sand and mud substrate. The study plots were established in a high density geoduck bed at the minus g.1m level (calculated from zero tide). MATERIAL anp METHODS Geoduck abundance within the 2 plots (each 45.7m 1.8 m) was assessed by divers placing small wire stakes next to each siphon observed. This process was repeated on each visit until no unstaked siphons were present. Considerable effort was expended to insure that all geoducks detectable were staked. The total number staked represented the actual geoduck population within the plots. All wire stakes were then removed and the plots allowed to return to normal. Monthly counts of geoduck siphons were then made from January to December 1974 using our standard tran- sect method and the percentage of the total estimated population observed during each monthly visit calculated. These observations were carried out by 3 divers who alter- nated between the plots to reduce the chance of bias from an individual diver remembering the location of certain geoducks within the plots. In addition, 4 small 0.46m X 0.46m plots were photo- graphed monthly to further document the seasonal change in “shows.” The plots were carefully approached and photographed manually using a Nikonos 35mm camera with ectachrome film. Tripod-mounted cameras proved unsatisfactory because the slightest disturbance of the bottom caused the clams to retract their siphons. McEr- LEAN & Howarp (1971) found that mechanical disturb- ance of the bottom affected “shows” of Eastern soft-shell clams (Mya arenaria). RESULTS anp DISCUSSION The percentage of the geoduck populations detected in the 2 plots varied from a low of 5% in January to a high of 59.8% in May, and averaged 38.0% in Plot A and 36.8% Page 156 THE VELIGER Vol. 20; No. 2 Table 1 Number of geoducks and percentage of estimated populations observed in monthly visual counts in two plots near Big Beef Creek, Washington Plot A Plot B Date (estimated geoduck population = 316) (estimated geoduck population = 358) {a= 1974 Number observed % observed Number observed % observed sa a EE —————————— Jan. Jan. 33 10.4 18 5.0 Feb. 101 32.0 117 32.7 March 142 44.9 169 47.2 April 133 42.1 170 47.5 May 172 54.4 214 59.8 June 168 53.2 178 49.7 July 105 33.2 106 29.6 Aug. 146 46.2 150 41.9 Sept. 144 45.6 174 48.6 Oct. 160 50.6 134 37.4 Nov 195 30.1 114 31.8 Dec 4] 13.0 37 10.3 Mean 38.0 36.8 in Plot B over the entire year (Table 1). Student’s T test was conducted on various combinations of the data using the arc sine transformation as suggested by SoKaL & Rou LF, 1969. Winter data (November-February) from Plot A were compared with winter data from Plot B and no significant differences were found (t = 0.25; d. f = 3). Plot A summer data (March-October) were also tested against summer data from Plot B and no significant dif- ferences were found (t = 0.27; d. f= 7). When data from Plots A and B were pooled and winter data tested against those from the summer, the differences were highly sig- nificant (t—=6.01; t.f=-11) demonstrating that the siphons were more readily detected by divers in the sum- mer than in the winter. Data from the photographic plots also demonstrated that geoducks were more easily observed in the summer months than in the winter (Table 2). The seasonal changes are shown in Figures ra- 1d. None of the 5 geoducks present in the 0.45m < 0.45m plot can be seen in the February (winter condition) photograph. The May photograph (summer condition) clearly shows 5 clams with their siphons extended. Four can be seen in the July photograph; 3 of these are not distinct. In the November photograph, only 1 siphon mark can be seen. Several small-scale surveys have been conducted in Puget Sound with underwater television and geoducks were found in water as deep as 60m. Geoducks living at these depths apparently behave in a similar manner as Explanation of Figures 1 to 4 Plot 1: Geoduck Population = 5 Figure 1: Figure 2: (pumping water) Figure 3: February 1974; no geoducks showing May 1974; all 5 geoducks showing, siphons open July 1974; 4 of 5 siphons showing, algae covered geoduck in lower left hand corner Figure 4: November 1974; 1 out of 5 geoducks showing as a slight mark, siphons are withdrawn, algae cover al- most gone Tue VELIGER, Vol. 20, No. 2 [Goopwin] Figures 1 to 4 Figure 4 ae 2 =~. A an = ue 4 E 2 i my 4 5 vo iv =f if ~ = @ = J A i at i a fT PS ‘ = a ee i é 7 : ; * t 2 ‘a - = r s 5 ’ < 4 0.10) (Table 1). All individuals observed during this study were crevice- inhabiting, and Haliotis walallensis were subjectively judged to be found in shallower cracks than H. rufescens. The size-frequency distribution of Haliotis rufescens showed no distinguishable modes over a broad range of sizes, While H. walallensis showed a single, distinct mode at 7.5cm (Figure 2). These frequency distributions are not significantly different from those found by Lowry «& Pearse (1973) (Kolomogorov-Smirnov test, p > 0.20) (see Figure 2). Lowrie & PEARSE (1973) Vol. 20; No. 2 THE VELIGER Page 165 Table 1 Densities of Abalones (X + 1 s.d. per m2) Haliotis spp. Haliotis rufescens Haliotis walallensis Lowry and Pearse (1973) (N = 45 1 X10m swath counts) 0.21 + 0.34 0.15 0.07 This paper (1976) (N = 25 10m? circular quadrats) 0.16 + 0.20 0.08 + 0.14 0.08 + 0.12 Lowrie & Pearse (1973) 12 12 8 8 4 4 ca) ° < 12 14 20 3 E re Haliotis walallensis Haliotis rufescens 5 This paper (1976) : 16 6 2 I 12 4 fo) 2 4 8 10 12 14 20 Shell Length (cm) Figure 2 Size-Fequency distributions of Haliotis walallensis and Haliotis rufescens off Hopkins Marine Station in the present study (summer, 1976) and in Lowry « Pearse (1973; data collected in November, 1972). The distributions from the two dates are not significantly different (Kolomogorov-Smirnov test, p > 0.20). Page 166 DISCUSSION The densities of Haliotis rufescens and H. walallensis have remained quite constant over the last 4 years since they were measured by Lowry & Pearse (1973). The slight differences in densities between Lowry & PEARSE (op. cit.) and the present paper may be a result of the differ- ences in sampling methods. Lowry & Pearse used belt transects (total sampling area = 3 560m”), while we used random quadrats in a smaller study area. One of their transects had densities (westline = 0.17 per m*) virtually identical to those in the present study, while the other transect densities were somewhat higher (east- line = 0.35 per m’), resulting in a slightly higher overall mean density (0.21 per m?). This stability, coupled with the observation that abalones in the study area continue to be found only in crevices, supports the contention that the crevice-dwelling populations have established a refuge from sea otter predation. Sea otters are, however, con- tinuing to feed on abalones, as well as a wide variety of other invertebrates in Hopkins Marine Life Refuge (Dan Costa, personal communication). Other predators in the area, including sea stars (Pycnopodia helianthoides (Brandt, 1835); Pisaster giganteus (Stimpson, 1857) ; and Orthasterias koehleri (de Loriol, 1897) ), octopuses (Octopus spp.), rock crabs (Cancer antennarius Stimp- son, 1856), and cabezone (Scorpaenichthys marmoratus Ayres, 1854), are known to feed on abalone, and all of these predators have ready access to the crevices inhabit- ed by abalone (Cox, 1962; FepEer, 1963; O’CoNNELL, 1953; personal observations). However, abalones have e- volved behavioral responses (clamping down, “running,” shell twisting, secretion of mucus, etc.) which minimize their capture (FEDER, op. cit.; MonTGOMERY, 1967; per- sonal observations). During other studies in the area, ab- alones smaller (<< 3cm) than most of those observed in this study have been found on open, exposed surfaces (Lowry & PEARSE, 1973; Hines & Pearse, unpublished ob- servations). It would be interesting to know if only those recruits which settle in crevices survive, or if abalones settle everywhere and later move into crevices. Our ob- servations agree with Lowry & Pearse (op. cit.) that H. walallensis is found in smaller, that is shallower, cracks than H. rufescens. It would also be interesting to know if this difference in crack distribution is the result of compe- tition between the 2 species, differences in crack prefer- ence, or perhaps simply a reflection of small abalones (both H. walallensis and smaller H. rufescens) tending to occur in smaller cracks. THE VELIGER Vol. 20; No. 2 The size-frequency distribution of Haliotis rufescens found in this study and that of Lowry & PEARSE (1973) lack any distinct modes over a wide size range. This is expected from the year-round spawning of this species (BooLootian et al., 1962; Younc & DEMarmTINI, 1970). However, the size-frequency distribution of H. walallensis in the present study is also identical to that found 4 years ago by Lowry & Pearse (op. cit.) with a single, distinct mode at 7.5cm. This unchanged size-frequency distribu- tion does not support the hypothesis that the H. walallens- is observed by Lowry «& Pearse represented a single age- class resulting from a single earlier successful settlement. Rather, as with H. rufescens, H. walallensis at Hopkins Marine Life Refuge appears to be in a stable size (age) distribution. LorKa (1922) showed that a population with constant survivorship and birth rates will reach and maintain a stable age distribution. The stable size (age) distribution of H. walallensis and H. rufescens thus sug- gests constant survivorship and recruitment rates for these populations, with recruitment into each size (age) class balanced by losses due to mortality and growth (aging). Finally, it is interesting to note that only one abalone measured in the Lowry & PEARSE (of. cit.) study and none in the present study was larger than legal size (7 inches or 170mm). ACKNOWLEDGMENTS We gratefully acknowledge John S. Pearse for his support and constructive reading of the manuscript. Donald P. Abbott gave generously of the facilities of Hopkins Marine Station, Stanford University. Dan Miller of Cali- fornia Department of Fish and Game provided the aerial photograph used for Figure 1. This work was supported by Sea Grant NOAA 04-6-1584402 to John S. Pearse. Literature Cited Bootootian, Ricwarp A., A. FARMANFARMAIAN & A. C. GIESE 1962. On the reproductive cycle and breeding habits of two western species of Haliotis. Biol. Bull. 122: 183 - 193 Cox, Kezirn W. 1962. California abalones, family Haliotidae. Fish. Bull. 118; Calif. Dept. Fish and Game, Sacramento, Calif; 133 pp. FEepEerR, Howarp MitTcHELL 1963. Gastropod defensive responses and their effectiveness in reducing predation by starfishes. Ecology 44: 505 - 512 Lotxa, ALFRED J. 1922. The stability of the normal age distribution. Acad. Sci. U. S. 8: 339 - 345 Lowry, L. FE « J. S. Pearse 1973. Abalones and sea urchins in an area inhabited by sea otters. Mar. Biol. 23: 213 - 219 Proc. Nat. Vol. 20; No. 2 THE VELIGER McLEAn, JAMES HAMILTON 1962. Sublittoral ecology of kelp beds of the open coast areas near Carmel, California. Biol. Bull. 122: 95 - 114 Montcomery, Davip H. 1967. Responses of two haliotid gastropods (Mollusca), Haliotis assim- ilis and Haliotis rufescens, to forcipulate asteroids (Echinodermata), Pycnopodia helianthoides and Pisaster ochraceus. The Veliger 9 (4): 359-368; plts. 50-51; 2 text figs. (1 April 1967) O’Conne ti, CuHarRLeEs P 1953. The life history of the cabezon. Scorpaenichthys marmoratus (Ayres). Calif. Dept. Fish & Game Fish Bull. 93 (1): 1-76 Youne, James S, & Joun D. DEMartin1 1970. The reproductive cycle, gonadal histology, and gametogenesis of the red abalone, Haliotis rufescens (Swainson), Calif Fish & Game 56: 298 - 309 Page 167 Page 168 THE VELIGER Vol. 20; No. 2 Observations on Feeding, Chemoreception and Toxins in Two Species of Epitonium BY SIGRID SALO! Hopkins Marine Station of Stanford University, Pacific Grove, California 93950 (1 Plate; 1 Text figure) INTRODUCTION Epitonium tinctum (Carpenter, 1864) and Epitonium indianorum (Carpenter, 1864) are small marine meso- gastropods often associated with sea anemones. THoR- SON (1957: 57) reported commonly finding E. tinctum at the base of the anemone Anthopleura xanthogrammica (Brandt, 1835). HoctBEerc (1971: 23) described E. tinctum feeding on the tentacles of A. elegantissima (Brandt, 1835). I have made further observations of E. tinctum and E. indianorum feeding on the tentacles and verrucae of 5 anemone species and have observed that Epitonium can apparently locate anemones by using a chemosensory method. Epitonium species produce a purple dye from the hypobranchial gland area, as do related snails. I have seen this dye staining the foot of an Epitonium. Perhaps partly due to speculation by David (cf: Witson & WIL- SON, 1956: 292) that the purple dye of Ianthina jan- thina (Linnaeus) anaesthetizes the Velella velella (Lin- naeus) on which it feeds, RoBERTSON (1963: 51) sug- gests that the dye produced by Epitonium species may be anaesthetic. I report here experiments that support this suggestion. MATERIALS anp METHODS Epitonium tinctum were collected intertidally at Mussel Point, Monterey Bay, California. Epitonium indianorum were collected at depths from 9 to 24m off Mussel Point and at the Pinnacles, near Pescadero Point, Monterey County, California. All collections were made in May 1975- * Present address: Box 455, Pateros, Washington 98846 For experiments on chemosensory perception of ane- mones, two tanks (approximately 7/ capacity each), one containing an anemone and one without, were connected by rubber tubing to a glass ‘Y’ tube. One branch of the Y received water from the anemone’s tank, the other from the tank without an anemone. Sea water flowed through the system at a constant rate of about 5ml/ per minute in each branch. Epitonium were placed in the junction of the branches of the Y and their positions recorded after 15 minutes. Five sets of experiments were run in which the motion of E. tinctum and E. indianorum was compared for the anemones Anthopleura elegantissima, A. xanthogrammica, Metridium senile (Verrill, 1865), Tealia lofotensis (Gosse, 1858), and T. crassicornis (Gosse, 1858). All anemones but T: crassicornis had a wet weight of about 30g; the smallest T. crassicornis available weighed about 60g. Five E. tinctum and 5 E. indianorum were used in each set of experiments; each was given 2 runs, for a total of 10 readings for each anemone for each Epitonium species. In a control experiment, Epitonium were tested when neither tank contained an anemone. In testing for a possible toxin or anesthetic, 2 types of experiments were carried out. First, to compare the be- havior of an Epitonium extract to that of previously ex- amined toxins, the ability of Epitonium extract to extin- guish the compound action potential of the sciatic nerve of the leopard frog, Rana pipiens, was tested. The nerve was placed over a set of silver stimulating electrodes and 2 sets of recording electrodes, one on either side of a plastic well which contained the suspected toxin. Supra- maximal stimuli were delivered by a Grass SD-5 stimu- lator. The resulting compound action potential was ampli- fied by a Grass P-8 pre-amplifier and displayed on a Tektronix 502-A dual beam oscilloscope. The ratio be- tween the peaks from the 2 sets of recording electrodes was noted before, and at intervals after the toxin was added, and the oscilloscope image was photographed Vol. 20; No. 2 before and after the toxin had taken effect (cf. Kao & FuHRMAN, 1967: 27). The effect of whole Epitonium, and of the pallial area of Epitonium, each crushed in amphibian Ringers solution, and of Epitonium crushed and then dialysed through Spectrapor tubing (#3. Ap- prox. MW cut-off 3500) against 2000ml of amphibian Ringers solution for 12-24 hrs was examined. Both E. tinctum and E. indianorum were used. The second set of experiments tested the reaction of an anemone tentacle to administration of Epitonium ex- tract. The tip of a severed Tealia crassicornis tentacle was tied to a Grass FT-03 force transducer, and the base of the tentacle was attached to a flared piece of capillary tubing over a syringe needle. A short length of rubber tubing connected the needle to the syringe body. Injec- tions of Epitonium crushed in sea water, of tryptamine HCl, and of sea water (as a control) were made into the tubing, and the tentacle’s response to the injections and to tactile stimulus was recorded on a Grass poly- graph. RESULTS Feeding Behavior Generally, Epitonium tinctum were found on Antho- pleura elegantissima and A. xanthogrammica, and E. indianorum were found on Tealia crassicornis and T. lo- fotensis, though there were numerous exceptions. I have found Epitonium on the column of an anemone and at the base. However, neither species of Epitonium is a THE VELIGER Page 169 permanently attached parasite; I have seen E. indianor- um 30cm away from the nearest anemone, and both species frequently crawled about in the tank. When feeding, an Epitonium may position itself on the column, on shell fragments on the column, if these are present, or beside the base of an anemone. The E ‘pitonium everts and extends up its long proboscis, which it bran- dishes until it encounters the tip of a tentacle. When this occurs, the proboscis is slipped over the tentacle, which can be seen inside since the proboscis is quite translucent. One or two minutes later, the anemone contracts in the area of the attacked tentacle, and the proboscis, with the tentacle still visible inside, is retracted. Epitonium will also feed in the same manner on the verrucae of ane- mones, especially the larger species such as Tealia crassi- cornis. Epitonium tinctum tends to feed on Anthopleura sp. and E. indianorum on Tealia sp. Neither Epitonium crawls onto the column of Metridium and this anemone is rarely attacked. Chemosensory Perception Results of experiments on movement of Epitonium with respect to anemones to test chemoreception are listed in Table 1. Of the 4 responses in the table, one is a positive response: motion toward the anemone. Two are avoidances: away from the anemone up the other branch of the Y, or down the Y. The fourth, lack of res- ponse, could be due to a variety of factors, possibly in- cluding an inability to sense the anemone in that test. Table 1 Chemosensory Perception of Anemones by two Species of Epitonium Epitonium tinctum or Epitonium indianorum were placed at the junction of branches of a Y tube. Sea water from a tank containing one of five species of anemones flowed through one branch of the Y, while sea water from a tank without animals flowed through the other branch. Movement was recorded toward the anemones, away from the anemones up the other branch of the Y, down the Y away from both tanks, or no movement if less than 0.5 cm. The values marked with ! indicate significant differences from controls with P less than 0.001. Epitonium tinctum Epitonium indianorum Toward Away Down Nomovement} Toward Away Down No movement Anthopleura elegantissima 91 0 0 1 J 1 4 4 Anthopleura xanthogrammica 7 2 0 1 0 4 oI 2 Metridium senile 1 2 5 2 1 4 3 2 Tealha crassicornis 3 10 0 3 ii! 1 1 3 Tealia lofotensis 3 2 2 3 5 2 2 1 Control 2 0 7 5 4 3 5 3 Page 170 THE VELIGER Vol. 20; No. 2 To see if the positive response of movement toward an anemone was significantly different from movement in control experiments without anemones, I made a x2 comparison. For the ‘expected’ value, I took the greater incidence of motion (as a percentage) up either branch of the Y in the absence of an anemone. I compared this value to the incidence of motion toward the anemone for each Epitonium species to each anemone species. Using this method, the statistically significant results (P < 0.001) are the increased motion of E. tinctum toward both Anthopleura species and of E. indianorum toward Tealia crassicornis. P is 0.05 for the increase in motion of E. indianorum toward T. lofotensis. All other combina- tions have P > 0.1 for increased motion toward the ane- mone. Epitonium were apparently able to sense whether the water around them had contained an anemone of the genus with which I had usually found them. Since their information came from the water itself, I assume some chemical derived from the anemone served as the means of location. Epitonium Toxin Extracts of Epitonium were found to inhibit the ampli- tude of the compound action potential of the sciatic nerve of Rana pipiens (Figure 1). In this case, the ratio of the spike recorded from a position on the nerve after the toxin well to the spike recorded from before the well decreased from 0.97 in the control to 0.50 after 1 hr, or a decrease of about 48%. The second spike also has been broadened as some of the faster conducting axons were extinguished. Similar results were obtained in other experiments. Epi- tonium extracts had been dialysed for 24 hrs against a total of 2000 mi Ringers solution produced results simi- lar to that shown in Figure 7. Thus the active substance is either a large molecule or is bound to a large molecule. In another experiment, the pallial area of Epitonium crushed in amphibian Ringers solution, decreased the action potential of the nerve by about 60% in 1 hr. Under the same conditions, the remainder of the body had no effect. Isolated tentacles from Tealia crassicornis exhibited spontaneous contractions and could be stimulated me- chanically or by tryptamine (cf. Ross, 1960). Extracts of Epitonium added to the sea water in which the ane- mone tentacle was suspended reduced or abolished he spontaneous contractions as well as the response to trypt- amine and mechanical stimuli. The results of a represent- ative experiment are shown in Figure 2. In one experi- ment extracts of Epitonium abolished spontaneous con- tractions for 18 min., after which the rhythm was re- established. The results suggest that Epitoniwm toxin in- hibits some part of the neuromuscular system of the anemone, but further work should be done to localize the site of action. DISCUSSION Experimental evidence is presented here for the sug- gestion made by others (Witson & Witson, 1956; RoBERTSON, 1963) that Epitonium produces a toxin or anesthetic that paralyzes the anemones on which it feeds, The toxin appears to be either produced or stored in the pallial area, possibly the hypobranchial gland. If the toxin is present in the hypobranchial gland, it may, like the purple dye produced in the gland, sometimes be present on the Epitonium’s foot, where it could anesthetize the anemone on which the snail crawls. In this case, Epitoni- um would itself be expected to be resistant to the anes- thetic. The toxin may instead be injected when the snail feeds. RoBERTSON (1963: 51) reports that some Epztoni- um possess hollow thorns in the proboscis, presumably to inject a toxin. However, using light microscopy, I was unable to find thorns or a duct which would transport the toxin from the pallial area up the proboscis on Epi- tonium indianorum. Explanation of Figure z Oscilloscope recordings of the compound action potential of frog sciatic nerve in the arrangement shown in the diagram. The super- natant from 39mg Epitonium indianorum crushed in 1ml Ringer solution was placed in the cup marked EPI. The spike recorded from the site upstream from the toxin remained essentially un- changed in amplitude, while that recorded from the site downstream from the toxin decreased Tue VELIGER, Vol. 20, No. 2 Control [Sato] Figure r Vol. 20; No. 2 THE VELIGER Page 171 4 5 + -_+—___+—__+——_ ++ fo) minutes 5 f fh} Figure 2 Tracings from polygraph records of contractions of an isolated tentacle from Tealia crassicornis. 1. Spontaneous contractions. (stretch) . 4. Addition of extract of Epitonium. 5: SUMMARY Epitonium tinctum and E. indianorum are two meso- gastropods that feed predominantly on the tentacles of certain sea anemones. In the field, the former was gener- ally found on Anthopleura species, and the latter on Tealia species. Experiments in the laboratory showed that they may locate specific genera of anemones using a chemosensory method. Both Epitonium species were found to contain a toxin in the pallial area which is either a large molecule or is bound to a large molecule. The toxin inhibits the compound action potential of frog nerve and may also inhibit the contraction of a tentacle from Tealia crassicornis. NoTE ADDED IN PROOF After this article was accepted for publication a report by Carey Rescu Smiru appeared in this Journal (19 (4): 331 - 340) confirming chemical recognition of Anthopleu- 2. Contraction in response to tryptamine. and subsequent arrows: repeated mechanical stimuli (stretch). 3. Contraction in response to mechanical stimulus ra elegantissima and A. xanthogrammica by Epitonium tinctum (Frederick Fuhrman). ACKNOWLEDGMENTS I appreciate the great amount of help I received from Dr. Frederick and Geraldine Fuhrman, of Hopkins Ma- rine Station. I would also like to thank Dr, Myra Keen for use of the Stanford shell collection, and Dr. Nathan Howe for many helpful suggestions. Supported in part by NIH Grant No. GM-16031. Literature Cited Hocuserc, Frep G., Jr. 1971. | Functional morphology and ultrastructure of the proboscis com- plex in Epitonium tinctum (Gastropoda: Ptenoglossa). Abstr. & Proc. West. Soc. Malacol. 4th Ann. Meet. Asilomar, Calif: 22 - 23 (27 December 1971) Page 172 THE VELIGER Kao, C. Y. « EF A. FunRMAN 1967. Differentiation of the action of tetrodotoxin and saxitoxin. Toxicon 5: 25 - 34 RoserTson, RoBEert 1963. Wentletraps (Epitoniidae) feeding on sea anemones and corals. Proc. Malacol. Soc. London $5; 51 - 63 Ross, D. M. 1960. The effects of ions and drugs on neuro-muscular preparations of anemones. Journ. Exper. Biol. 37: 760 - 763 Tuorson, GUNNAR 1957. Parasitism in the marine gastropod family Scalidae. Vidensk. Medd. Dansk Naturh. Foren. 119: 55 - 58 Wison, D. P « M. A. Witson 1956. A contribution to the biology of Janthina janthina. Journ. Mar. Biol. Assoc. U. K. 35: 291 - 305 Vol. 20; No. 2 Vol. 20; No. 2 THE VELIGER Page 173 The Development of Conspecific Interactions in Juvenile Aplysia dactylomela Rang, 1828: An Observational Study I. IZJA LEDERHENDLER '* Department of Animal Behavior, American Museum of Natural History Central Park West at 79” Street, New York, NY 10024 (3 Text figures) INTRODUCTION THE PURPOSE OF THIS STUDY was to describe develop- mental changes in the interactions of “juvenile” Aplysia dactylomela Rang, 1828. It became possible with the dis- covery of an apparently “juvenile” population in the Plajita Rosada collection on the southwest coast of Puerto Rico during April and May 1975 (see Figure 1). This was an unusual research opportunity in that the capabili- ty of raising sea hares through metamorphosis has been restricted to only a few laboratories in which behavioral development was, to date, not investigated. The methods and procedures used were peculiar to an opportunity such as this, in that they depended on the continued suc- cess of finding “juveniles.” Aplysia californica Cooper, 1863 has been raised through metamorphosis and some of the morphological and behavioral changes related to metamorphosis have been described (KrIEGSTEIN, 1976; KRiEGSTEIN et al., 1974). Further progress in the establishment of labora- tory cultures of Aplysia has been reported by HapFiExp, (1975) and STRENTH & BLANKENSHIP (1976). Recently studies of morphological development have become avail- able for some related opisthobranchs (THompson, 1958, 1962, 1967; Tarpy, 1970; Bonar « HapFIELD, 1974), but there are no published studies of behavioral develop- ment in the opisthobranchs. * Present address: Laboratory of Biophysics, National Institutes of Health, Marine Biological Laboratory, Woods Hole, MA 02543 The first occurrence of copulation in Aplysia seems to be related to size. Although no data were provided, in a brief report Newsy (1972) stated that copulation was never observed in animals less than 40g body weight. SmitH & CareFroot (1967) collected small sea hares (1g-8g) and kept these animals together as a group. Copulation was not observed until the 16% day in the laboratory. The body weights at the time of copulation were not reported; however, copulation was observed at about the time the gonads became mature. Thus, it ap- pears that the reproductive behavior develops some time after metamorphosis. Morphological studies of the bag cells support the no- tion that the reproductive system as a whole may develop after metamorphosis. The bag cells are a neurohormonal group of the abdominal ganglion which contribute to egg- laying (ArcH, 1976). The cells are few in number and may be non-secretory in small animals (2g). In sea hares larger than 50g, however, secretory granules are present and the number of cells in the cluster increases markedly (Frazier, et al., 1967). Thus, some neuroendo- crine aspects of egg-laying become functional around the stage when sea hares begin to copulate. Although the data are scanty, it appears from the above studies that copulation and size might be related. In this study the same sea hares were observed in regular repeated pairings. Systematic observations of their be- havior and weight measurements were made. Page 174 THE VELIGER Vol. 20; No. 2 02’00” S Arriah Htia Romie Zone : & s~ Magueyes O Sr Isla Sy Guayacan t * Laboratory of the Department of Marine Sciences University of Puerto Rico ---- Plajita Rosada Zone — Enrique Zone ie Collado Zone 0b LS a meters 150 300 600 900 1824 Figure 1 METHODS anp PROCEDURES A. Subjects and Maintenance A sample of 56 sea hares was collected off the southwest coast of Puerto Rico at Plajita Rosada. This is a rocky intertidal beach marked by the presence of Laurencia paptlosa and sargassum (for details of collection see LEDERHENDLER, 1977). The sea hares were maintained in an indoor laboratory in 1/-capacity aquaria with running sea water and con- stant daylight illumination. Food was always available and consisted of Acanthophora spicifera which was thor- oughly rinsed before being placed in the tanks. Wet weights of the animals were measured every other day on a Mettler balance. To carry out observations of the interindividual behavi- or, the members of a pair were matched for size. As a result, the smallest animals were paired at the time of collection, to permit a maximum amount of observation during a long period of growth and development. : : : I Februa Five pairs were chosen for observation. These were all pees a) A el = N OH c i=) oO we NO April Number of Animals “DH OOF OI ew NOP OV March from the April and May collections (Figure 2). A failure ; ; 5 2 January in the sea water supply cut the period of observation I short, resulting in making only 15 to 22 days available ; A : : : : O- 16- 3I- 46- 61- 76- 9I- 106- ra1- 136- 161- in which a total of 22 paired observations were made. 15. go 45) 65. 9B Go) RvG5 © 20a euGOMENIE Weight (g) B. Apparatus and Procedure Figure 2 Observations occurred between 5:00 p.m. and 7:30 Pom. under a 60-watt fluorescent bulb. The animals were Frequency Distribution of Initial Weights of Aplysia dactylomela paired by their size. Two pairs were observed for 30 Collected at Plajita Rosada, Puerto Rico (N = 56) Vol. 20; No. 2 THE VELIGER Page 175 minutes every 6 days; 2 pairs were observed every other day. The 5" pair was observed as individuals on experi- mental day 1 and then every 6 days (Table 1). A circular bowl 30cm in diameter and 10.2cm high was filled with fresh sea water to a depth of 8cm. The animals were taken from their home tanks and weighed. The observa- tion began immediately upon the introduction of the second animal. Table 1 Characteristics of Five Pairs of Pre-Copulatory Aplysia dactylomela Observed Repeatedly in Different Frequencies Initial Number of _ Interval between weight (g) pairings pairings Pair 1 Partner A 14 4 6 days Partner B 16 Pair 2 Partner A 25 3 6 days Partner B 26 Pair 3 Partner A 13 2 6 days? Partner B 16 Pair 4 Partner A 6 7 2 days Partner B 11 Pair 5 Partner A 21 6 2 days Partner B 24 *Unpaired on first experimental day. The frequency and duration of contact and copulation as well as more detailed aspects of the interactions were recorded. Contact was defined as any part of an individual touching any other part of another individual. A sea hare initiated contact if it approached the other and touched it with any part of the body. Approach was defined as any locomotion which decreased the interindividual dis- tance while oriented toward the second individual. Reci- procal contact occurred when both animals initiated the contact. Copulation was said to occur upon the intromis- sion of one animal’s penis into the common genital open- ing of the partner. Under the observational conditions described above, this could be determined exactly. If a pair was still in contact or copulating by the end of the observation, spotchecks every 5 minutes continued until the animals separated. Table 2 Patterns of Contact and Copulation of “Juvenile” Aplvsia dactylomela Paired Every Iwo Davs Pairing 4 ) 6 Pair 4 Number of Contacts Initiated 0 0 (0) 3 1 4 0 By Partner A Number of Contacts Initiated 1 3 By Partner B Number of Contacts Initiated 0 1 1 0) 2 wd 2 Reciprocally Pair 5 Number of Contacts Initiated 2 0 0 2 0 0 = By Partner A Number of Contacts Initiated 0 (0) 0 0 0 0 = By Partner B Number of Contacts Initiated l 0 1 Reciprocally ihe} fo) Xe) No) oo Sperm Recipient — — B B B C. Sample Characteristics Figure 2 shows the distribution of weights for the popu- lation from which the experimental subjects were se- lected. The median weight of the entire group was 56g. The experimental sea hares were collected in April and May when most of the population (15 out of 28) were below 60g. The median weight of the experimental ani- mals was 16g. Only 1 pair of the 56 animals from Plajita Rosada was found copulating and no other animals were found in contact even though most were collected near each other, within approximately 75m of coastline. RESULTS Figure 3a - 3e shows the growth curves and duration of contact and copulation with repeated pairings for each pair in the study. As these data indicate, copulation oc- curred in only 1 pair (5). There appears to be a tendency for longer contact interaction with increasing size where- Page 176 THE VELIGER Vol. 20; No. 2 120) pair 1 1800 110 Y, 1650 100 1500 go 1350 80 1200 Oo % 70 1050 § = ce = 60 goo 8 = 50 759 @ 40 600 30 450 20 300 10 150 (0) == 10} 5/3 5/9 5/16 5/22 Dates of Observation Figure 3a Total duration (sec) of contact (solid bar) and weight changes of immature Alysia dactylomela found at Plajita Rosada, Puerto Rico. Copulation did not occur (92s) uonema 5/25 5/31 6/6 Dates of Observation Figure 3b Total duration (sec) of contact (solid bar) and weight changes of Aplysia dactylomela found at Plajita Rosada, Puerto Rico. Copulation did not occur ee are 1207 Pair 3 1800 110 1650 100 1500 go 1350 80 1200 1, 6 70 1050 5 % 60 goo 5 = 50 750 os 40 600 30 450 20 300 10 150 to) fo) 5/25 5/31 6/6 Dates of Observation Figure 3c Total duration (sec) of contact (solid bar) and weight changes of Aplysia dactylomela found at Plajita Rosada, Puerto Rico. Copulation did not occur 1207 Pair 4 1800 110 1650 100 1500 90 1350 80 1200 2p 1050 yy ~ 60 goo fi z E “2 50 750 3 = 40 600 8 30 450 20 300 10 150 Co) Co) 135 ot Se faiel ys 6/6 Dates of Observation Figure 3d Total duration (sec) of contact (solid bar) and weight changes of Aplysia dactylomela found at Plajita Rosada, Puerto Rico. Copulation did not occur Vol. 20; No. 2 90 80 270 cH 9 = 60 ey Z 2° 40 ‘g ey oO (o} 20 5/ 26 o! 6 ; /289! 3 6/1 /3 65 Dates of Observation was lotal Duration of Contact [— 1 Tootal Duration of Copulation o—o Partner A e—e Partner B Figure 3e Total duration (sec) of contact (solid bar), copulation (open bar) and weight changes of Aplysia dactylomela found at Plajita Rosada, Puerto Rico ever there is a consistent tendency to contact at all. Pairs 4 (Figure 3d) and 5 (Figure 3e) each had 7 and 6 observations respectively. This permitted somewhat more detailed statements about the interactions. As Table 2 shows, pairs 4 and 5 were different in their patterns of contact. In pair 4, a single contact, initiated by B, oc- curred early during the initial pairing and lasted for 35 seconds. The animals weighed 11g and 20g respectively during this observation. During the 1“ 3 pairings, 2 con- tacts were initiated mutually; 6 other contacts were ini- tiated only by Partner B (1 in the 1, 3 in the 2°, and 2 in the 3" pairing). In the 4" observation B initiated 6 out of g contacts. In the 5", B initiated 2 contacts, A initiated 1, and 2 were mutually initiated. In the 6" pairing, out of 8 separate contacts, B initiated only 2, 4 were initiated by A, and 2 were mutual. But in the 7" pairing B initiated 3 out 5 contacts where 2 were mutually initiated. Thus, out of 27 separate contacts which were not initiated by both animals (total of 8), B initiated 19 and A initiated 8. In pair 5, contact occurred during the 1“ observation when the pair weighed 28g and 34g respectively. There THE VELIGER Page 177 was no contact during the 2" pairing. Considering all 6 observations, 6 contacts were initiated reciprocally and 4 were not. These 4 were all initiated by partner A, who was also the sperm donor in 3 of the 4 observation periods where copulation occurred. Partner B became the sperm donor only after A had assumed that role 3 times. The animals contacted shortly after the 3 pairing and began to copulate after 90 seconds of the observation period had elapsed. At this time, they weighed 53g and 58g respec- tively. Copulation occurred in each of the next 3 pairings. The 4 copulations lasted for 43, 12, 21 and 49 minutes respectively. It is noteworthy that when partner B as- sumed the sperm donor role the duration increased to a level equivalent to the first copulation when A was the sperm donor. DISCUSSION The Aplysia observed in this study were assumed to be pre-copulatory because of the following: 1. Of 56 animals collected from the same location, only one pair was found copulating in the field; these 2 individuals weighed 47g and 55¢ respectively. In addition, none of the animals found in the field was in contact, although many were near each other. 2. Most of the sea hares did not copulate with repeated pairings in the laboratory. 3. None of the animals laid eggs while they were in the laboratory. Together with their small size when collected, these observations suggest that they were not reproduc- tively mature. Although only 1 of the 5 pairs copulated, it shared some characteristics with those that did not copulate, such as change in weight, duration of contacts, etc. However, it is useful to examine this pair in detail. For example, Frazier et al. (1967) found that the bag cells of A. cali- fornica begin to contain neurosecretory granules when animals reached 50g in weight. Copulation occurred after each animal of pair 5 passed the 50g mark in body weight, indicating that this may be a necessary but not sufficient condition for copulation to occur, as all pairs reached that weight before observations ended. In the one pair which copulated, the animal which was the sperm recipient was heavier at the start (B). Follow- ing the first copulation there was a spurt in its growth rate in comparison with its partner (A’s), and their congruent pre-copulatory pattern of weight change. If the spurt in weight proves reliably associated with copulation, the on- set of egg-laying (which may depend on bag-cell hor- mones) could be related to copulation and the contacts which precede it. Page 178 THE VELIGER Vol. 20; No. 2 LEDEHENDLER et al. (1976) found that in reproduc- tively mature Aplysia dactylomela animals which behave more consistently in the sperm recipient role copulate with more partners than the consistent sperm donors. They suggest that this may be based on a feedback system related to amount of sperm received. In the observa- tions recorded here, role reversal occurred after 3 copu- lations in which the same roles were assumed. This is consistent with the function of such a feedback system. These observations are considered preliminary but indi- cate directions for further research on the possible inter- dependence of social stimulation and physiology in the development of reproductive processes in this species. ACKNOWLEDGMENTS I wish to thank W. P. Aspey, E. S. Hodgson, E. Tobach, H. R. Topoff and H. P. Zeigler for their helpful criticisms in the preparation of this paper. I also wish to thank Kate Herriges for her assistance in all phases of the re- search. The work was supported in part by the Depart- ment of Animal Behavior of The American Museum of Natural History, the Research Foundation of The City University of New York, Training Grant MH-13051 and NIMH Grant No. 1RO3-MH24275-0. Literature Cited ArcH, S. 1976. Neuroendocrine regulation of egg-laying in Aplysia californica. Amer. Zool. 16: 167 - 175 Bonar, Dare B. a M. G. Haprizip 1974. | Metamorphosis of the marine gastropod Phestilla sibogae Bergh (Nudibranchia: Aeolidacea). I. Light and electron microscopic an- alysis of larval and metamorphic stages. Journ. Exp. Mar. Biol. Ecol. 16: 227 - 255 Frazier, W. T, E. R. Kanner, I. KuprerMann, R. Waziri & R. E. CocczsHaii 1967. Morphological and functional properties of identified neurons in the abdominal ganglion of Aplysia californica. Journ. Neurophysiol. $0: 1288 - 1357 Haprie.p, M. G. 1975. Continuous laboratory culture of two Aplysia species nears suc: cess. Labor. Anim. 4 (3): 17 Kriecstein, A. R. 1976. Stages in post-hatching development of Aplysia californica. Journ. Exp. Zool. 199: 275 - 288 Krugostezin, A. R., V. Castertucci « E. R. KANDEL 1974. Metamorphosis of Aplysia californica in laboratory culture. Proc. Nat. Acad. Sci. 171 (9): 3654 - 3658 LEDERHENDLER, I. Izja 1977. Reproductive behavior of Aplysia dactylomela Rang, 1828 (Ga- stropoda : Opisthobranchia). Ph. D. Thesis, City Univ. New York LEDERHENDLER, I. Izyja, A. Lopzz & ETHEL, TosacH submitted Patterns of copulation in the simultaneous hermaphrodite Aplysia dactylomela Rang, 1828 (Gastropoda: Opisthobranchia). Newsy, N. A. 1972. Aspects of the behavior of Aplysia dactylomela Rang, 1828. Hawaii. Inst. Mar. Biol. Tech. Rep. 18: 19 Smiru, S. T. « T. H. Cargroor 1967. Induced maturation of gonads in Aplysia punctata Cuvier. Nature 215: 652 - 653 StrentH, Nep E. « James E. BLanKgNSHIP 1976. | Laboratory culture and metamorphosis of larval Aplysia brasili- ana Rang (Gastropoda, Opisthobranchia). Bull. Amer. Malacol. Union, Inc., in press Tarpy, Jean 1970. Contributions a l’6tude des métamorphoses chez les nudibranches. Ann. Sci. Nat., Paris 12: 299 - 370 THOMPSON, THOMAS EVERETT 1958. The natural history, embryology, larval biology and post-larval development of Adalaria proxima (Alder and Hancock) (Gastropoda: Opisthobranchia). Phil. Trans. Roy. Soc. London, B. 242: 1-58 1962. Studies on the ontogeny of Tritonia hombergi Cuvier (Gastropoda: Opisthobranchia). Phil. Trans. Roy. Soc. London B 245: 171 - 218 (4 October 1962) 1967. Direct development in a nudibranch, Cadlina Iaevis, with a dis- cussion of developmental processes in Opisthobranchia. Journ. Marine Biol. Assoc. U. K. 47: 1-22; 8 figs. Vol. 20; No. 2 NOTES & NEWS Direct Development in the Intertidal Gastropod Batillaria zonalis (Bruguiére, 1792) BY SYLVIA BEHRENS YAMADA AND CHANDRA S. SANKURATHRI Department of the Environment, Fisheries and Marine Service, Pacific Biological Station Nanaimo, British Columbia, Canada VoR 5K6 (1 Text figure) Tue Mup snaiL Batillaria zonalis (Bruguiére, 1792), a native of Japan, was introduced to North America with the Japanese oyster, Crassostrea gigas (Thunberg, 1793) (Quays, 1964). Well established populations of B. zon- alis can be fourd on mud flats from California to British Columbia (MacDona _p, 1969) wherever Japanese oysters were planted, but not in other suitable habitats. Nothing is known about the breeding habits and early life history of this mollusk. On the basis of B. zonalis’ discrete distri- 4 0.4mm Figure 1 Egg Capsule of Batillaria zonalis (Bruguiére, 1792) THE VELIGER Page 179 bution, QuayLE (op. cit.) suggested that it may have either a direct development or a short pelagic life. Adult Batillaria zonalis were collected from Fanny Bay (49°18’N Lat.; 124°48’ W Long.) on June 25, 1976 and maintained in filtered sea water at 22°C. Seven days after the collection B. zonalis released sedentary egg cap- sules measuring 0.4mm wide and 0.6 - 0.7mm long. De- tritus and mud adhere to egg capsules thus enabling them to anchor to the sediment orothersubstrates. Each capsule contained 1 zygote measuring 0.13mm in diameter. One end of the capsule is notched and the other nippled (Fig- ure 1). Three days after the deposition of egg capsules embryos were moving in 67 out of 70 capsules. Larval shells were observed bv day 7 and on day 11 juvenile snails crawled out of their capsules. The juvenile snails were 0.3mm long and appeared to be benthic and not equipped for pelagic life. The absence of a pelagic dispersal stage might be a controlling factor in preventing B. zonalis from spreading to other suitable mud flats. We thank Dr. Dan B. Quayle for reading the manu- script. Literature Cited Macpona_p, KeitH B. 1969. Quantitative studies of salt marsh faunas from the North Ameri- can Pacific coast. Eco]. Monogr. 39 (1): 33-60 14 text figs.; 12 tables (Winter 1969) Quay e, DanieL BrancHu 1964. Distribution of introduced marine Mollusca in British Columbia waters. Journ. Fish. Res. Brd. Canada 21 (5): 1155-1181; 10 text figs. (10 November 1964) A New Sea-Floor Oasis BY A. MYRA KEEN 2241 Hanover Street, Palo Alto, California 94306 RECENT EXPLORATION of the sea-bottom along the Gala- pagos rift zone has revealed a rich fauna. This find is second only to the discovery in the 1950s of Neopilina west of Costa Rica. At depths of around 2500m (1400 fathoms), hot springs along the rift somehow make pos- sible a bottom fauna of spectacular size and abundance. In February, 1977 an expedition funded by the Interna- tional Decade of Ocean Exploration (National Science Foundation), jointly sponsored by Oregon State Univer- Page 180 sity, Woods Hole, Massachusetts Institute of Technology, and Scripps Institution, with participation by others, such as Prof. Tjeerd van Andel of Stanford University, made several dives in a submersible craft, the “Alvin,” to study the hot springs. The bottom fauna surrounding the springs came as a surprise, and only a few specimens could be recovered with the equipment on the “Alvin.” Photographs made during the dives reveal crabs, tube worms, limpet-like mollusks, abundant large clams and mussels, and others. Dr. van Andel reported seeing large gastropods up to }cm long grazing on a black slime on the rocks. All of the invertebrates seen were of phenomenal size. One Calypto- gena shell that was recovered intact measures more than 25cm in length, and the 2 valves weigh 525g (18 oz.). Bottom temperature around the springs is 10° C, where- as the surrounding water is about 2° C. Fish were abun- dant, also, and sessile organisms such as hydroids, bryo- zoans, crinoids, and sea anemones were observed. Water from the springs is of normal salinity, low in oxygen, rich in manganese and iron, and markedly radioactive. This find may give us a whole new direction in our efforts to explain faunal movements of the past. It also underscores the need for further study to reveal how such abundant life can be nourished in an oxygen-poor envi- ronment so far away from the photic zone. The area is about 320km (200 miles) ENE of the Galapagos Islands, where 3 large plates of the earth’s crust are gradually moving apart (hence the term “rift zone”). A Color Variant of Conus bulbus Reeve BY JOHN K. TUCKER Department of Biological Sciences Illinois State University, Normal, IL 61761 (3 Text figures) ALTHOUGH MANY SPECIES of Conus are known to be highly variable in color pattern, the West African C. bulbus does not seem to be particularly noted for color variants. All specimens that I have seen have a pattern of longitudinal bars with some degree of variation in the nature of the bars. Recently I purchased a specimen of C. bulbus with a highly unusual and apparently previous- ly unreported color pattern. While it has the character- THE VELIGER Vol. 20; No. 2 istic shape and spire of C. bulbus, the longitudinal bars are broken up into small dashes. At first glance, the dashes appear to be arranged in spiral lines. Closer examination shows that traces of the longitudinal bars may still be seen near the anterior end of the shell. I do not know how common this color variant is but it is the first I have seen among the more than 50 C. bulbus I have had an opportunity to examine. Figure 1 Conus bulbus, usual phenotype from Luciras Bay, Angola Figure 2 Conus bulbus, usual phenotype from Lobito Harbor, Angola Figure 3 Conus bulbus, color variant from Lobito Harbor, Angola All Figures X1.5 The Identity of Conus fulmineus Gmelin BY JOHN K. TUCKER Department of Biological Sciences Illinois State University, Normal, IL 61761 (1 Text figure) Kouwn (1966) REVIEWED the nomenclatural history of Conus fulmineus Gmelin, 1791. He (Kohn) designated plt. 58, fig. 644 of Martini (1773) as the lectotype of this species. KoHN (1966, 1968) noted that C. fulmineus, Vol. 20; No. 2 THE VELIGER Page 181 although considered a provisionally valid species, is not represented by any specimens that he knew of. Recently, shell dealers and collectors have been identifying a West Pacific cone (figured by Hinton, 1972: plt. 34, fig. 13) as C. fulmineus. This identification is certainly erroneous. The figure of the lectotype of C. fulmineus (KouHN, 1966; 1968) shows a cone with an elevated, sharp pointed, sca- lariform spire. The body of the shell is marked by numer- ous interrupted spiral lines which are at intervals covered by longitudinal brown bars. The specimens erroneously identified as C. fulmineus lack interrupted spiral lines. They also have 2 irregular brown bands, one of which covers the posterior half of the body, the other on the anterior half leaving white near midbody. They do not have longitudinal bars that run the length of the whorl. The identity of this species is currently under study and at present I know of no name that has been applied to it. Figure 1 In the last few years, several specimens have come to my attention which appear to be conspecific with the lectotype of Conus fulmineus. A specimen at present in my private reference collection is shown in Figure 1. The color pattern and the elevated, scalariform spire closely resemble the lectotype and general proportions are also similar. The only difference of importance is that the shoulder is more angular on this specimen than the shoul- der of the lectotype. This difference could well be due to artistic inaccuracy, or possibly the lectotype was beach worn. The close resemblance in other major features out- weighs this difference and specimens similar to the one shown in Figure 1 should be considered C. fulmineus Gmelin. Synonyms of C. fulmineus include C. fulgurans Hwass, 1792 which was also based on the Martini figure (Koun, 1968), C. selectus A. Adams, 1855 from Malacca, C. lentiginosus Reeve, 1844 and C. optabilis A. Adams, 1853. The latter 2 species were described from shells lacking locality data. The geographic range of C. fulmine- us is uncertain, but specimens that I have seen have been from the Indian Ocean in the vicinity of the Indian sub- continent. The specimen figured here was trawled from an unknown depth by fishermen near Madras, India. Literature Cited Hinton, A. 1972. Shells of New Guinea and the central Indo-Pacific. 44 plts.; 2 text figs. Jacaranda Press, Brisbane Konn, ALan Jacops 1966. Type specimens and identity of the described species of Conus. IIJ. The species described by Gmelin and Blumenbach in 1791. Journ. Linn. Soc. (Zool.) 46: 73 - 100 1968. Type specimens and identity of the described species of Conus. IV. The species described by Hwass, Bruguiére and Olivi in 1792. 94 pp.; Journ. Linn. Soc. (Zool.) 47: 431 - 503 Martini, Frieprich Heinrich WILHELM ; 1773. Neues systematisches Conchylien-Cabinet. 2. Niirnberg. W.S. M. AT THE TENTH ANNUAL meeting of the Western Society of Malacologists the following slate of officers was elected to serve during the fiscal year 1977/1978: President: Dr. Peter D’Eliscu First Vice-President : Barry Roth Second Vice-President: Dr. Vida Kenk Salle Crittenden Carol Skoglund Sally Bennett Patrick LaFollette The Eleventh Annual Meeting will be held at Santa Clara University, Santa Clara, California 95053 from Wednesday, June 28 to Saturday, July 1, 1978. The Society elected Mr. Emery Perkins Chace, the old- est living, active conchologist in the United States, and Dr. Wendell Oliver Gregg to Honorary Life Membership. Secretary: Treasurer: Members-at-Large: Page 182 Another Generous Contribution from the San Diego Shell Club AFTER OUR JULY ISSUE had come back from the printer, we received a very generous donation to our Endowment Fund from the San Diego Shell Club. It was, therefore, impossible for us to express our thanks to the Club and its members until now. As in the past number of years, the continued generosity of our friends helps us to keep the membership dues at their low level in spite of the uninter- rupted steady rise in costs of paper and printing. Thus, our thanks to the Club is not only on behalf of the journal itself but also on behalf of the many members of the California Malacozoological Society. Moving? If your address is changed it will be important to notify us of the new address at least six weeks before the effective date, and not less than six weeks before our regular mailing dates. Because of a number of drastic changes in the regulations affecting second class mailing, there is now a sizeable charge to us on the returned copies as well as for our remailing to the new address. We are forced to ask our members and subscribers for reimbursement of these charges; further, because of increased costs in connection with the new mailing plate, we also must ask for reimbursement of that expense. Effective January 8, 1968 the following charges must be made: change of address - $1.- change of address and re-mailing of a returned issue — $2.75 minimum, but not more than actual costs to us. We must emphasize that these charges cover only our actual expenses and do not include compensation for the extra work involved in re-packing and re-mailing returned copies. In view of the ever increasing difficulties in the postal service, it is essential that members and subscribers not only give us prompt and early notice of address changes, but that proper arrangement for forwarding of our jour- nal be made with the local post office (at the old address). THE VELIGER Vol. 20; No. 2 New Postage Rates The U. S. Postal Service has increased second class mail rates, effective on July 6, 1977. We will, however, not in- crease our charges for mailing The Veliger, but must insist that we are reimbursed in all cases for returned copies and for the expenses involved in remailing such copies to a new address. It is very important for our members to realize: a) the postal service will not forward any mail other than first class for more than go days, even though forwarding postage may be guaranteed by the addressee; and _b) it is totally impossible for us to make changes in addresses in less than 6 weeks. We must make an address change even if only one digit in the ZIP code is changed, and the cost to us is the same as for a completely new address. Under no circumstances are we able to supply free re- placement copies of issues that fail to reach their proper destination. However, we will ship by insured mail re- placement copies at half the announced single copy rate of the particular issue plus postage. We have developed a triple check system so that, if we say that a copy has been mailed, we are absolutely certain that we delivered that copy to the post office in Berkeley and on the date we indicate. From our experience with the loss of insured mail, we are tempted to suggest that subscribers figure on a 10% reserve fund for the purchase of replacement copies. The only alternative remaining would be for us to increase subscription rates and membership dues by at least 10%. This, however, does not seem quite fair to us as some of our subscribers in almost 20 years have never failed to receive their copies. On July 6 the rates for book parcels and the library rate have been increased. This necessitates that we must in- crease the postage charges on back volumes, supplements and individual back numbers. The charges stated must be increased by 20¢ for one item and by 8¢ for each addi- tional item. Regarding UNESCO Coupons We are unable to accept UNESCO coupons in payment, except at a charge of $4.25 (to reimburse us for the ex- penses involved in redeeming them) and at $0.95 per $1.00 face value of the coupons (the amount that we will receive in exchange for the coupons). We regret that these char- ges must be passed on to our correspondents; however, our subscription rates and other charges are so low that we are absolutely unable to absorb additional expenses. Vol. 20; No. 2 CaiForNIA MALAcozooLocicaL Society, Inc. announces Backnumbers of THE VELIGER and other publications Volumes 1 through 8: out of print Volume 9: $22.- Volume 10: out of print Volumes 11 and 12: out of print Volume 13: $24.- Volume 14: $28.- Volume 15: $28.- Volume 16: $32.- Volume 17: $34.- Volume 18: $34.- Volume 19: $34.- We now have a limited number of volumes 9, 11, 13, 14 to 18 available bound in full library buckram, black with gold title. These volumes sell as follows: Volume 9 at $27.-; Volumes 11 and 13 at $29.- each; Volumes 14 and 15 at $33.-; Volume 16 at $38; Volumes 17, 18, 19 at $41.75; to this we must add a handling charge of $2.75 per volume for shipment to domestic addresses and $4.75 for shipment to any foreign address. Further, we must collect the appro- priate amount of sales tax on the price of the bound vol- umes sold to California residents. Supplements Supplement to Volume 3: $6.-* plus handling charge [Part 1: Opisthobranch Mollusks of California by Prof. Emst Marcus; Part 2: The Anaspidea of California by Prof. R. Beeman, and The Thecosomata and Gymnosomata of the Cali- fornia Current by Prof. John A. McGowan] [The two parts are available separately at $3.- each plus a handling charge (see below). If purchased separately, each part is subject to the California State sales tax if mailed to California addresses]. Supplement to Volume 6: out of print. Supplement to Volume 7: available again; see announce- ment below. Supplement to Volume 11: $6.-* plus handling charge. [The Biology of Acmaea by Prof. D. P. Assott et al., ed.] Supplement to Volume 14: $6.-* plus handling charge. [The Northwest American Tellinidae by Dr. E. V. Coan] Supplement to Volume 15: $15.-* plus handling charges as follows: $1.50 for addresses in the United States of A- merica; $3.00 for all other addresses. 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These charges reflect the latest increases in fees by the U. S. Postal Service. These handling charges must, however, remain subject to change without prior notice, depending on the vagaries of rate-fixing by the postal service. Items marked with * are subject to sales tax in the State of California; residents of that State please add the appropriate amount to their remittances. Prices subject to change without notice. Send orders with remittance to: Mrs. J. DeMouthe Smith, Department of Geology, Cali- fornia Academy of Sciences, Golden Gate Park, San Fran- cisco, California 94118. Please make remittance payable to C. M.S., Inc. in U.S. $, net and free of any fees to the Society. Shipments of material ordered are made once a month. We are forced to adopt this measure because of the con- tinual cut-back in personnel at the U. S. Post Office with the amount of time wasted standing in line having increased to intolerable lengths. Since it requires the same amount of time to mail 20 packages as it takes to mail one, the saving of time by our reduced mailing schedule is obvious. It becomes glaringly obvious that with the increase in postage rates and fees, the service is deteriora- ting at increasing rapidity. Although we spend much of our time complaining, one voice is not enough to bring about a change. Subscription rate to Volume 20 remains the same. We must emphasize that under no condition can we ac- cept subscription orders or membership applications for calendar year periods. If “split volumes” are required, we must charge the individual number costs. Individual issues sell at prices ranging from US$12.- to US$20.-, depending on the cost to us. Page 184 Backnumbers of the current volume will be mailed to new subscribers, as well as to those who renew late, on the first postal working day of the month following receipt of the remittance. The same policy applies to new members. Tue VELIGER is not available on exchange from the Cali- fornia Malacozoological Society, Inc. Requests for re- prints should be addressed directly to the authors con- cemed. We do not maintain stocks of reprints and also cannot undertake to forward requests for reprints to the author(s) concerned. A Glossary of A Thousand-and-One Terms Used in Conchology by WiniFrep H. ARNOLD originally published as a supplement to volume 7 of the Veliger, has been reprinted and is now available from The Shell Cabinet, Post Office Box 29, Falls Church, Virginia 22046, U. S. A. The cost is US$ 3.50 postpaid if remittance is sent with the order. WE ARE PLEASED to announce that an agreement has been entered into by the California Malacozoological Society, Inc. with Mr. Steven J. Long for the production and sale of microfiche reproductions of all out-of-print editions of the publications of the Society. 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Volumes and Supplements not listed as available in microfiche form are still available in original edition from the Society at prices indicated elsewhere in the NOTES & NEWS section. : THE VELIGER Vol. 20; No. 2 WE CALL THE ATTENTION oF our foreign correspondents to the fact that bank drafts or checks on banks other than American banks are subject to a collection charge and that such remittances cannot be accepted as payment in full, unless sufficient overage is provided. Depending on the American banks on which drafts are made, such charges vary from a flat fee of $1.- to a percentage of the value of the draft, going as high as 33%. Therefore we recommend either International Postal Money Orders or bank drafts on the Berkeley Branch of United California Bank in Berkeley, California. This institution has agreed to honor such drafts without charge. UNESCO coupons are N OT acceptable except as indicated elsewhere in this section. Supplements Many of our members desire to receive all supplements published by the Society. Since heretofore we have sent supplements only on separate order, some members have missed the chance of obtaining their copies through over- sight or because of absence from home. It has been sug- gested to us that we should accept “standing orders” from individuals to include all supplements published in the future. After careful consideration we have agreed to the proposal. We will accept written requests from individuals to place their names on our list to receive all future sup- plements upon publication; we will enclose our invoice at the same time. The members’ only obligation will be to pay promptly upon receipt of the invoice. Requests to be placed on this special mailing list should be sent to Mrs. J. DeMouthe Smith, Manager, C.M.S&., Department of Geology, California Academy of Sciences, Golden Gate Park, San Francisco, CA(lifornia) 94118. However, until further notice, we are suspending the pub- lication of supplements until it will be reasonably certain that we will not be forced to spend many hours in tracing of lost insured or registered parcels and entering claims for indemnification. The special mailing list of members and subscribers who have entered an “including all sup- plements” will be preserved because of our innate opti- mism that sometime within our lifetime the postal services throughout the world will return to the former excellent and reliable performance. Claims for defective or missing pages must reach us within 60 days from the publication date. We will not Vol. 20; No. 2 THE VELIGER Page 185 respond to claims of missing issues made less than 30 days by domestic addressees, or less than 60 days by foreign addressees after the publication date of our journal issues. This refusal is necessary as we have received an increasing number of “claims” as much as 6 months before the claimed issue was to be published. We wish to conserve our energy and the cost of postage and stationery for more productive purposes. We are willing to accept requests for expediting our journal via AIR MAIL; however, in that case we must ask for an additional payment of US$8.00 in all cases where the Veliger goes to domestic addresses, and a depos- it of US$18.00 for all foreign addresses (including PUAS). Of course, we will carry forward as a credit toward the postage charges of the following year any amount over the actually required postage charges. At present we are charged a minimum fee of $12.50 on each order for new addressograph plates. For this rea- son we hold off on our order until 6 weeks before mailing time, the very last moment possible. If, for any reason, a member or subscriber is unable to notify us in time and also is unable to make the proper arrangement with the Post Office for forwarding our journal, we will accept a notice of change of address, accompanied by the proper fee and a typed new address on a gummed label as late as 10 days before mailing time. We regret that we are absolutely unable to accept orders for changes of address on any other basis. In view of the probable further cur- tailment in the services provided by the Postal Service, we expect that before long we may have to increase these time intervals. CALIFORNIA MALACOZOOLOGICAL SOCIETY, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. 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Please send for membership ap- plication forms to the Manager or the Editor. Membership renewals are due on or before April 15 each year. If renewal payments are made after April 15 but before March 15 of the following year, there will be a re-instatement fee of $1.-. Members whose dues pay- ments (including the re-instatement fee) have not been received by the latter date, will be dropped from the rolls of the Society. They may rejoin by paying a new initiation fee. The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. Endowment Fund In the face of continuous rises in the costs of printing and labor, the income from the Endowment Fund would materially aid in avoiding the need for repeated upward adjustments of the membership dues of the Society. It is the stated aim of the Society to disseminate new infor- mation in the field of malacology and conchology as widely as possible at the lowest cost possible. At a Regular Membership meeting of the Society in No- vember 1968 a policy was adopted which, it is hoped, will assist in building up the Endowment Fund of the Society. An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 000.- or more, an entire volume will be dedicated to the memory of the decedent. REGARDING POSTAL SERVICE Complaints regarding late arrival of our journal are in- creasing in number, steadily, continually. However, we very conscientiously dispatch our journal on the printed publication dates. What happens after deposition at the Post Office is, of course, beyond our control. From some of our members we have been able to construct a sort of probable delivery schedule. In general, within California, Page 186 8 days is usual; outside of California, the time lapse in- creases with the distance; the East Coast can consider a lapse of “only” two weeks as rapid service; 4 to 5 weeks are not uncommon. Foreign countries may count on a minimum of one month, six weeks being the more usual time requirement and over two months not rare! To Prospective Authors Postal Service seems to have deteriorated in many other countries as well as in the United States of America. Since we will absolutely not publish a paper unless the galley proofs have been corrected and returned by the authors, the slow surface mail service (a minimum of 6 weeks from European countries, 8 to 12 weeks from India and Africa) may make a delay in publication inevitable. We strongly urge that authors who have submitted papers to the Veli- ger make all necessary arrangements for expeditious read- ing of the proofs when received (we mail all proofs by air mail) and their prompt return by air mail also. Since we conscientiously reply to all letters we actually receive, and since we experience a constant loss in insured and registered mail pieces, we have come to the conclusion that if a correspondent does not receive an answer from us, this is due to the loss of either the inquiry or the reply. We have adopted the habit of repeating our inquiries if we do not receive a reply within a reasonable time, that is 6 weeks longer than fairly normal postal service might be expected to accomplish the routine work. But we can not reply if we have never received the inquiry. Because of some distressing experiences with the Postal Service in recent years, we now urge authors who wish to submit manuscripts to our journal to mail them as insured parcels, with insurance high enough to cover the complete replacement costs. Authors must be prepared to document these costs. If the replacement costs exceed $200.-, the manuscript should be sent by registered mail with additional insurance coverage (the maximum limit of insurance on parcel post is, at present, $200.-). We are unable to advise prospective authors in foreign countries and would urge them to make the necessary inquiries at their local post offices. We wish to remind prospective authors that we have announced some time ago that we will not acknowledge the receipt of a manuscript unless a self-addressed stamped envelope is enclosed (two International Postal Reply Coupons are required from addresses outside the U.S. THE VELIGER Vol. 20; No. 2 A.). If correspondence is needed pertaining to a manu- script, we must expect prompt replies. If a manuscript is withdrawn by the author, sufficient postage for return by certified mail within the U.S.A. and by registered mail to other countries must be provided. We regret that we must insist on these conditions; however, the exorbitant in- creases in postal charges leave us no other choice. Some recent experiences induce us to emphasize that manuscripts must be in final form when they are sub- mitted to us. Corrections in galley proofs, other than errors of editor or typographer, must and will be charged to the author. Such changes may be apparently very simple, yet may require extensive resetting of many lines or even entire paragraphs. Also we wish to stress that the require- ment that all matter be double spaced, in easily legible form (not using exhausted typewriter ribbons!) applies to all portions of the manuscript — including figure explana- tions and the “Literature Cited” section. It may seem inappropriate to mention here, but again recent experience indicates the advisability of doing so: when writing to us, make absolutely certain that the cor- rect amount of postage is affixed and that a correct return address is given. The postal service will not forward mail pieces with insufficient postage and, if no return address is given, the piece will go to the “dead letter” office, in other words, it is destroyed. Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. That the printed date is the actual date of pub- lication under the rules of the International Commission on Zoological Nomenclature is based on the following facts: 1) The journal is delivered to the Post Office on the first day of each quarter, ready for dispatch; 2) at least three copies are mailed either as first class items or by air mail; 3) about 20 copies are delivered in person to the mail boxes or to the offices of members in the Berkeley area; 4) two copies are delivered to the re- ceiving department of the General Library of the Univer- sity of California in Berkeley. Thus our publication is available in the meaning of the Code of the ICZN. The printed publication date, therefore, may be relied upon for purposes of establishing priority of new taxa. Vol. 20; No. 2 INFORMATION DESK What’s the Difference? Telegraphic Style versus Normal Style BY A. MYRA KEEN 2241 Hanover Street, Palo Alto, California 94306 IN SYSTEMATIC WorK telegraphic style is generally adop- ted for the descriptions of taxa, yet few style manuals give discussions on how and when to use it. A compact or condensed manner of writing, telegraphic style omits verbs or uses them only as present participles, and articles of- ten are omitted also. Sentences tend to run together, sep- arated by commas and semicolons rather than periods. The subject of each phrase or sentence is stated early, usually as the first word. This is an appropriate style for descriptions of species and for tabular matter that fol- lows -— statements of type locality, distribution, type repositories, etc. In adopting the style one needs to re- member to be consistent within the paragraph and not switch from telegraphic to normal until one is ready to go on in normal style to conclude the section or the paper. As a sample and as evidence of how much space can be saved by use of telegraphic style, here is a somewhat abridged and emended description from a recent issue of The Veliger, followed by a transcription into normal style: Shell large, elongate, strong; protoconch of about 2 smooth, convex whorls, followed by 9 gently convex whorls forming a high, pointed teleoconch; suture slightly indented; sculpture consisting of heavy axial ribs (12 on penultimate whorl), and microscopic lines of growth throughout, bent forward at suture, with a few fine spirals at base ... Paraphrase in normal style: The shell is large, elongate, and strong. The proto- conch has about two smooth, convex whorls, followed by nine gently convex whorls that make up a high, pointed teleoconch. The suture is slightly indented. The sculpture consists of heavy axial ribs, of which THE VELIGER Page 187 there are twelve on the penultimate whorl. Micro- scopic lines of growth are present over the entire shell and are bent forward at the suture. A few fine spiral threads are present at the base ... BOOKS, PERIODICALS, PAMPHLETS Living Marine Molluscs by C. M. Yonce & T. E. Tompson. Collins, London, 288 pages, 16 plates (8 in color), 162 text figures, cloth- bound, $13.95, 1976. This is a richly interesting and valuable book for any invertebrate zoologist, marine biologist, or professional or amateur malacologist. Co-authored by T. E. Thompson, who contributed two excellent chapterson opisthobranchs, it must be regarded as largely the work of C. M. Yonge, who furnished the remaining 16 chapters and epilogue. The focus on living marine molluscs and their adaptive radiation has been achieved, in part, by a clear limitation of the aspects of molluscan biology that are covered in detail. Thus, there is little on internal anatomy and the physiological functioning of circulatory, respiratory, di- gestive, and excretory systems; the complexities of stom- ach morphology and the details of ctenidial structure in bivalves are not spelled out (they have been well covered by other authors) ; there is little discussion of systematics and phylogeny; there is, of course, no discussion of the highly successful land and fresh-water pulmonates; and at first glance the book might appear to be strongly de- voted to British molluscs. Yonge and Thompson view the Mollusca from the outside, or from within the mantle cavity, where the former is so much at home. What has resulted from their selective omissions is a finely balanced account of the astonishing adaptive radiation of the Mol- lusca, by authors who have a feel for the subject. In short, they give an unsurpassed picture of what molluscs are all about, and the varied ways in which they make a living. British examples are used where appropriate, but the authors, notably Yonge, draw heavily upon experience on the California coast, the Australian Barrier Reef, and other regions. The book is written in clear, simple lan- guage, no more technical than necessary, but that does not imply that it is intended for the beginner. Without some Page 188 THE VELIGER Vol. 20; No. 2 basic knowledge of molluscs, one could be overwhelmed in the accounts of so many and diverse adaptations. The breadth of the book will make it appeal to any malaco- zoologist, since it presents so well-balanced an overview of the evolutionary achievements of so diverse a phylum. There are a few minor errors: the reference on p. 84 to fig. 3 should be to fig. 4; on p. 178 reference to fig. 160 should be to fig. 162; and on p. 125 a reference to Lima- pontia depressa (fig. 65) sends one to a picture of Alderia modesta. That the motion-detecting eyes of scallops, with their unique reflecting-telescope focusing, should be said to be “similar” (p. 257) to the box-camera eyes of ceph- alopods is a bit too much of an oversimplification. But only a dour New England reviewer would call attention to the typographical error (p. 211) that designates the familiar quohog or little-neck or hard clam (Mercenaria mercenaria) as a soft-shelled clam (this name being used at the Hub of the Universe for Mya arenaria, also long-necked clam, which passes in Britain as the sand gaper). The book is well-illustrated and indexed. It is by all means highly recommended, to be read and enjoyed. Ralph I. Smith Department of Zoology University of California Berkeley, California 94720 Checklist and Bibliography of the Tertiary and Quaternary Mollusca of Japan 1950 - 1974 by Koicuimro Masupa & Hirosui Nona. 494 pages, 4 text figures. Saito Ho-On Kai, 20-1, Honcho-2 chome, Sendai, Japan. 15 December 1976 This comprehensive volume complements the 1952 Hatai & Nisiyama checklist of Japanese Tertiary mollusks de- scribed and illustrated through 1949. It is an alphabetic listing of nearly 5000 fossil mollusks described or illus- trated during the 25 year period 1950 - 1974. Mollusks from Tertiary and Quaternary formations are listed separately by taxonomic order and by geologic peri- od. The format is similar to that of the well-known catalogue of Tertiary mollusks of California by A. M. Keen and Herdis Bentson published in 1944. Masuda and Noda have indicated taxonomic revisions and corrections, where appropriate, following the author’s original cita- tion and an indication of geologic age, formation, and geographic location. A bibliography of papers dealing with Cenozoic mollusks of Japan accompanies the check- list; part of the bibliography covers papers on fossil mol- lusks from neighboring countries, another deals with topi- cal studies such as biogeochemistry, absolute age dating, and shell structure. This excellent report, together with the earlier com- panion volume, will be especially valuable to paleontolo- gists studying the systematics and biogeography of mollus- can faunas of the circum-North Pacific area, as well as to neontologists interested in the origin and migrational history of modern molluscan faunas of this region. Warren O. Addicott Directory of Palaeontologists of the World Third Edition, 1976 by Epxram Gerry. 3” edition, 1976.Universitetsforlaget, P.O. Box 307, Blindern, Oslo 3, Norway. 1976. US$g.50. The third edition published under the auspices of the International Palaeontological Association, includes names and addresses of some 6000 paleontologists throughout the world. It represents a much more com- plete compilation than the 1968 edition which included only about half as many names. Following the alphabetic listing of paleontologists, is a listing of specialists by taxonomic specialization. This section includes several specialty subheadings under each phylum or plant group. Finally, there is a geographic listing, by countries and provinces, states or cities, of institutions at which pale- ontologists work. Warren O. Addicott The Biogeography and Numerical Taxonomy of the Oegopsid Squid Family Ommastrephidae in the Pacific Ocean by Joun H. Wormutu. Bulletin of the Scripps Institution of Oceanography vol. 23: 1 - 90; 22 text figs.; 15 tables. $5.00 from Univ. Calif. Press, Berkeley or Los Angeles. [2223 Fulton Street, Berkeley, CA 94720]. 22 Nov. 1976 A partly confused and contradictory taxonomic ar- rangement within the squid family Ommastrephidae is studied on the basis of as large a quantity of material as could be assembled and also on the basis of numerous characters that could be ‘measured’ accurately in one way or another. The result of this approach was a clari- fication of many obscure relationships and a partial recti- fication of taxonomic problems, as well as the recognition Vol. 20; No. 2 THE VELIGER Page 189 that more work along the same lines will be needed for an eventual total resolution of the problems of interspec- ific relationships. R. Stohler Das grosse Buch der Meeres Muscheln by S. Peter Dance. Translated and re-worked by Rupo von CosEL. 304 pages; 1520 color photographs and 73 pen and ink drawings. Eugen Ulmer, P. O. Box 1032, 7000 Stuttgart 1, West Germany. DM 88.- (ca. $40.-) In recent years the frequency of publication of books in the German language on mollusks has increased; not only the monographic works such as “Das Tierreich’ and the indispensable work of Thiele-Zilch, but a number of books aimed at the collector have appeared. On a recent visit to Switzerland we noticed in the show windows of book stores several books of this latter category by German authors. A still more recent development is the appear- ance of translations of books by American and English authors. Some of these are straight translations, taking over all the errors of the original works. Others, however, show that considerable effort has been expended in re- viewing the original work during the task of translation and errors have been eliminated as far as possible. This is the case in the present book. The systematic portion of the book is preceded by a preface from the pen of the well-known malacozoologist Wulff Emmo Ankel, an introduction including brief chap- ters on systematics and nomenclature, morphology, bio- logy, on collecting and preparing shells, an historical review and a plea for conservation. A key with excellent explanatory figures completes this 34 page introduction. The 250 pages of the systematic portion are followed by an appendix containing accounts of species not included in the original work, a glossary and a bibliography. This part is organized into separate lists of works relating to the different geographical regions and contains some brief, but trenchant comments. The index concludes the volume. It seems probable that this volume will be of value to many collectors who have at least a working knowledge of the German language as the descriptions and comments are written in a relatively uninvolved style. There is not a single sentence that can be called overly long — in con- trast to the German style of about 50 or 60 years ago, when it seems that a sentence of less than a page-length was considered poor style. R. Stohler Type Specimens of Invertebrates (excluding Insects) Held at the Royal Scottish Museum, Edinburgh by G. Smatpon, D. Heppett and K. R. Watt. Royal Scottish Museum Information Series, Natural History 4: 118 pp. August 1976 This well organized, large-sized (21 X 30cm) publication includes 21 pages devoted to the mollusks. The following quotes from the introduction to the list are of great im- portance because they indicate the scope of the list as well as what further discoveries in the holdings of the Museum may be made: “As a number of hitherto unrecognized type-specimens came to light as a direct result of investigation for the purposes of this Catalogue it is expected that further re- search will reveal the status of other similar material in certain of the collections. This is particularly likely in the case of early collections such as those of L Dufresne, E Forbes, R K Greville and W Nicol ... In general, para- types have not been included ... The ‘Scotia’ collections ... contain a number of potential types awaiting descrip- tion. ... In two cases as yet unpublished lectotype desig- nations are included while a third lectotype (for Cerithium angustissimum Forbes) is designated herein.” There are very valuable annotations for many of the taxa listed. It seems to us that this particular list might well serve as a sample for similar contributions from other museums. R. Stohler Simposio sui molluschi terrestri e dulcicoli dell’Italia Settentrionale Proceedings of the Societa Malacologica Italiana, Gruppo Naturalistico Mantovano. 103 pp.; numerous illustrations. Eleven papers presented at the symposium on terrestrial and freshwater mollusks are presented; of these 10 are in Italian and one in English. All articles have at least 2 summaries in languages other than Italian. One of these summaries in each case is in English. This will facilitate the use of this interesting collection of papers presented at the symposium held at Mantua on May 10 and 11, 1975. Interested workers should inquire of Dr. Dario Franchini, 37, Via Cremona, 46100, Mantova, Italy about cost and availability of this volume. R. Stohler Page 190 The Western Society of Malacologists, Annual Report Pacific Grove, California, June 23-26, 1976. Volume 9 71 pages; 4 photographs, 12 October 1976 Abstracts and full texts of papers presented at the 9" annual meeting, as well as some brief resumés of business transactions are contained in this publication. It is dis- tributed to members of the Society; available at $5.- to non-members from Dr. Eugene Coan, 891 San Jude Ave., Palo Alto, CA(lifornia) 94306. R. Stohler Catalogo de los Moluscos Terrestres Cubanos del Genero Cerion (Mollusca-Pulmonata-Ceriidae) (con una Bibliografia General) — Catalogo de la Fauna Cubana - XXXVII by Micuet L. Jaume Garcia. Ciencias Biologicas (4) 51: 46 pp. April 1975 Of the total of 46 pages, 24 are devoted to a taxonomic, ecologic and zoogeographical discussion of the 147 species and subspecies, and keys to the 3 subgenera. The remain- der of the pamphlet is taken up by an apparently com- plete bibliography of the family in the West Indies. R. Stohler Muscheln und Schnecken des Meeres by R. Tucker Assott. German translation by Dr. Heinz Schréder; illustrations by George and Marita Sandstrom. Bunte Delphin-Biicherei Nr. 28. 160 pp.; numerous color illustrations; price unknown. 1976. This pocketbook is essentially the same as the one pub- lished a few years earlier in the Golden Press series. The illustrations are, for the most part, well done. However, in contrast to the American version of this work, there are no translations of the scientific names, creating so-called THE VELIGER Vol. 20; No. 2 ‘common names’, but the few German names that do ap- pear in the booklet are genuine vernacular names. This booklet will be of use to general collectors whose command of the German language is better than that of the English language. R. Stohler Uber Ammoniten des Schwabischen Juras by BERNHARD ZIEGLER. pp. 3 to 35; § plates; 22 text figs. Die entrollten Ammoniten des Schwabischen Juras by Gerwarp Drett. pp. 36 to 43; 1 plate; 4 text figs. both papers in Stuttgarter Beitrage zur Naturkunde, ser. C, Nr. 4. 1975 These two beautifully illustrated and clearly written ar- ticles are obviously intended for the interested layman as well as for the specialist, be the latter paleontologist or geologist. R. Stohler Pacific Marine Life by Cartes J. DeLuca and Diana Macintyre DeLuca. Charles E. Tuttle Company, Rutland, Vermont 05701. 66 pages; 1 map; numerous text figs. $2.75. 30 Aug. 1976 This small pocket book has the ambitious subtitle: A Survey of Pacific Ocean Invertebrates. Pages 29 to 44 are devoted to the mollusks. Within this limited space the very large phylum of the Mollusca can be treated only most sketchily and this the authors accomplish by apparently leaning heavily on Spencer Tinker’s book “Pacific Sea Shells.” The approach, however, is much more in a popu- lar vein than is Mr. Tinker’s work. In the hands of young children this small book will do no damage, probably. R. Stohler THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributional, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS"; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES’ will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 812” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 51/2” by 81/2”), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr, Donatp P. Aszott, Professor of Biology Hopkins Marine Station of Stanford University Dr. Warren O. Appicott, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Associate Professor of Paleontology, Stan- ford University Dr. Jerry DonouuE, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatr Duruam, Professor of Paleontology University of California, Berkeley, California Dr. Caper Hann, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. JoeL W. Hepcpetu, Adjunct Professor Pacific Marine Station, University of the Pacific Dillon Beach, Marin County, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Vicror Loosanorr, Professor of Marine Biology Pacific Marine Station of the University of the Pacific EDITOR-IN-CHIEF Dr. Rupo.tr STOHLER, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Associate Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. Franx A. Prrevxa, Professor of Zoology University of California, Berkeley, California Dr. Rosert Rozertson, Pilsbry Chair of Malacology Department of Malacology Academy of Natural Sciences of Philadelphia Dr. Peter U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. JupirH Terry Situ, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Raupu I. Smitu, Professor of Zoology University of California, Berkeley, California Dr. CuHar.es R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. THompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. Jean M. Care Rancho Santa Fe, California 674.05 V43 Mell, NILLIA é ANE Ba ‘ ~*~ A rm F SECTIONAL LIBRARY Co) ee PCT YA |} on RACE Et ft od Pedi *. DIVIS 21N41 , OF VIVLLUGAS THE VELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California VOLUME 20 JANuaARY 1, 1978 NUMBER 3 CoNntTENTS The Samoan Land Snail Genus Ostodes (Mollusca : Prosobranchia : Poteriidae). (2 Plates; 36 Text figures) ELIZABETH-LouIsE GrrarpI oi Miele 0h: es I Epace he ies a NA Spawning and Early Life History of Murex pomum Gmelin, 1791. (2 Plates; 2 Text figures) Euna A. Moore & Finn SANDER eterna ra Veritas: uN MIE Wet ht se The Systematic, Adaptive and Physiological Significance of Proteolytic Enzyme Distribution in Bivalves. Rosert G. B. Rep . Reproductive Biology of Colus stimpsoni (Prosobranchia : Buccinidae). I. Male Genital System. (5 Plates; 2 Text figures) Davip L. West Ua imEs Sueno ee lel lon har inane ays Malate" ra ay Pages Hiatella solida (Sowerby, 1834) (Mollusca : Hiatellidae) on Concholepas concho- lepas (Bruguiére, 1789) and other substrates. (3 Text figures) Cartos Ga.iarpo §. & Cecitia Osorio R. Zonation of Marine Gastropods on a Rocky Intertidal Shore in the Admiralty Gulf, Western Australia, with Emphasis on the Genus Nerita. (3 Text figures) Frep EtHan WELLS Gr Gi gt aes as ae RCS ee ame ER a A New Species of Serpulorbis ee Vermetidae) from South Africa. (4 Text figures) Rocer N. HucuHes . Ne Maem ae AMES 2 MoS cdi ait cal Sihian?' esse A New Species of Coryphella (Nudibranchia : Flabellinidae) from Santa Barbara, California. (2 Plates) Rosert K. Cowen « Dav R. Laur . [Continued on Inside Front Cover] . IQ! 7 251 . 260 . 266 - 274 - 279 . 288 . 292 Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SUPERFAMILY, FamiLy, Subfamily, Genus, (Subgenus) New Taxa Second Class Postage Paid at Berkeley, California ConTENTs — Continued A New Species of Anachis (Gastropoda : Columbellidae) from the Eastern Pacific. (2 Text figures) R.A; WYENEY 90 eS acs elt cee ea eae os een og ie eo mmr mre 2 Oy NOTES &: NEWS) 200 27th ye ance teloer elec Glee es ee aac sgakon Dene Mot ns Meet emmne 2 Op Nomenclatural Note on Hinnites giganteus (Gray). Barry RoTH & Eucene V. Coan Publication Dates of Bercu’s 1879 Papers Describing American Chromo- dorids. Rosert BuRN The Date of Publication of ANTON’s “‘Verzeichnif® der Conchylien” WALTER O. CERNOHORSKY Another Cephalopod from Northern California (Mollusca : Cephalopoda) RosBert R. TALMADGE BOOKS, PERIODICALS &°PAMPHEEMS! (3/5 ya) jn nn on meen Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 20: $25.- plus postage ($1.- in U.S.A.; $2.50 to RU.A.S.; $3.50 to all other foreign Countries, including Canada). Single copies this issue $16.-. Postage extra. Send subscription orders to California Malacozoological Society, Inc. 1584 Milvia Street, Berkeley, CA 94709, U.S. A. Address all other correspondence to Dr. R. STOHLER, Editor, Department of Zoology University of California, Berkeley, CA 94720 Vol. 20; No. 3 THE VELIGER Page 191 The Samoan Land Snail Genus Ostodes (Mollusca : Prosobranchia : Poteriidae) ELIZABETH-LOUISE GIRARDI Department of Zoology, Field Museum of Natural History Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605 (2 Plates; 36 Text figures) TABLE or CONTENTS WAU CLeaeeFeMN 6 9 Bed 6 6) 650 a do ole o OM AS Spatial ey Sele) “oR 6 os Jone ols AD I. Introduction . A 191 1. Distribution on iene Se tenlin.G yaw Wu eeltae them rep vae neue YI II. Historical Review of Gyelophorid Classification 5 192 QuSyMpathysacwser ne eres eins sere | by kenge eal enol III. Previous Studies of the Genus Ostodes . 193 37 Relativeyabundance/e ascii mien f-n-24g IV. Material Studied 194 B. Mee SA sedis 30 1, 6 wo. oF 6 LLG V. Methods 194 . Effects of environmental) differehices Staring, ban renee TG} VI. Systematic Review . ; 195 2. Effects of sympatry . . . . We br ae ou AO A. Variations in structural features ‘ 195 3. Species self-recognition mechanion go oe wld ‘og BAG 1. Shell form and ornamentation 195 C. Patterns of eve 6 gy Hb a Ain So deal gn fo en 2G) 2. Opercula 197 IX. Summary . . ee nchnclelllias (Mod POL lately tnterinaty daly ace aL) 3. Radulae . 201 X. Literature Cited . . . . 250 4. Gross anatomy . 203 XI. Appendix: Abbreviations Wied in eetomicat ipaires a. General 203 b. Digestive system 203 c. Male reproductive system 203 I. INTRODUCTION d. Female reproductive system 207 5. Sexual dimorphism 2i1 : : ; ; if, Greases pecans - as THIS REPORT Is MODIFIED from a dissertation submitted in 1. The genus Ostodes Gould, 1860 . 214 1973, in partial fulfillment of the requirements for the 2. Key to the species Bi 214 degree of Doctor of Philosophy, to the Graduate School 3. Criteria for species recognition 215 of Northwestern University, Evanston, Illinois. It revises 4. Species diagnoses and distributional data . 216 the species of the land snail genus Ostodes Gould, 1862, a. Ostodes plicatus (Gould, 1848) 216 which is found only in Western Samoa and American b. Ostodes gassiesi (Souverbie, 1858) . . 217 . . : a ee Samoa. Three new species, Ostodes reticulatus, O. exas- c. Ostodes reticulatus Girardi, spec. nov. . 220 : , d. Ostodes strigatus (Gould, 1848) . Aap peratus, and O. lanero, are described, and the previously e. Ostodes savaii Clench, 1949 . 224 compound species O. plicatus (Gould, 1848) is redefined. f. Ostodes exasperatus Girardi, spec. nov. . 227 Investigation showed that there are species recognition g. Ostodes llanero Girardi, spec. nov.. 228 differences in terminal structures of the genitalia. While h. Ostodes upolensis (Mousson, 1865) 230 _ the presence of such differences is well known in pulmon- te OOD CARS (SIGE, TED) 233 ate land snails, this is, to my knowledge, the first report of J. Ostodes cookei Clench, 1949 . 234 ; . : this phenomenon in land prosobranchs. k. Ostodes tiara (Gould, 1848) . 235 : siete L. Ostodes garretti Clench, 1949 236 The work is based upon materials in the mollusk col- C. Ts generic affinities of Ostodes 237 lection of the Field Museum of Natural History, Chicago. 1. Gonatorhaphe spp. 237 The methodology used required having shells and soft parts 2. Gassiesia sp. . . 238 associated at the start of data capture. Material in the Mu- 3. Mexcyclotus panamensis (Da Costa, 1908) 238 seum of Comparative Zoology, Harvard University, and AD ecu an Neca ene : rise Bernice P. Bishop Museum is stored with the shells in VII. Descriptive Geography 241 : . Nipper f ane one cabinet and the soft parts in another; therefore loans im Gea - 242 of anatomical materials from those institutions were not Civegetationy mrs oma eiite requested. Page 192 In the absence of personal field work, no information on niche differentiation can be presented. Some data on the anatomy of extralimital Pacific island taxa and one Neotropical taxon are presented for establishing outgroup comparisons. The members of my committee, Dr. Frank A. Brown, Jr., Dr. Olin Rulon, and Dr. Alan Solem have given advice and counsel, and have been most patient with a very pro- tracted period of study. Drs. Kenneth J. Boss, of the Mu- seum of Comparative Zoology, Harvard University; Frank Climo, Dominion Museum, Wellington, New Zealand; Joseph Rosewater, United States National Museum; Bert- ram Woodland and Carol Jones, Field Museum of Nat- ural History; Shi-Kuei Wu, University of Colorado, Boul- der; and Frederick Schramm, Eastern Illinois University, Charleton, loaned specimens or gave advice. Dr. Winston Ponder, Australian Museum, Sydney, allowed me to ex- amine the holotype of Cyclostoma brazieri Cox, 1870. Mrs. Carole W. Christman bore patiently with my in- experienced dissections, and taught me a great deal in the process of producing the illustrations. Her artistic skill was available through the generosity of my father, Fred- erick K. Leisch, whose timely gift to the Field Museum was used in part to support Mrs. Christman’s work. Mr. Fred Huysmans and Mr. John Bayalis of Field Museum of Natural History prepared the photographic prints used in this paper. Elizabeth Liebman drew some of the quan- titative diagrams. I am indebted to Dr. Solem for the SEM photographs of the radulae, to Dorothy Karall and Claire Kryczka for mounting figures, and to Jayne Freshour for manuscript typing. Miss Evelyn Patterson and Mr. Vic Snodgrass helped with translations from the Latin. Barbara Walden assisted in labeling figures and cataloguing. Finally, I wish to thank my husband, Joseph B. Girardi, who refused to let me quit when I got discouraged, helped with German translations, and put up with sloppy house- wifery while I studied; and my daughters, Stephanie and Susan, who bore cheerfully with their part-time mother. II. HISTORICAL REVIEW or CYCLOPHORID CLASSIFICATION The family Cyclophoridae was first established by Gray (1847: 181-182). It included a miscellany of genera pre- viously included in Cyclostomacea of earlier authors such as ANTON (1839: 52-54). Over the next few years, addi- tional genera were added by various authors, the place- ment being based entirely on shell characters. PFEIFFER (1852: 14) formally grouped the Cyclophoridae and the Pomatiasidae into the superfamily Cyclostomacea. He di- THE VELIGER Vol. 20; No. 3 vided the Cyclophoridae into 4 groups that were roughly equivalent to modern subfamilies, which he named Cy- clotea, Diplommatinea, Cyclophorea, and Pupinea. FiscHER (1885: 739) separated the Cyclophoridae from the Cyclostomatidae on animal and radular characters. KosELT (1902: 336, 484, 231) recognized 3 additional subfamilies — Alycaeinae, Craspedopomatinae, and Neo- cyclotinae, giving great importance to zoogeographic fac- tors. THIELE (1929: 94—113) essentially utilized Kobelt’s system. The first substantial innovation was by TIELECKE (1940). His work was based on consideration of male and female reproductive systems, central nervous system, and organs in the pallial cavity, particularly the kidney and circulatory system. He felt strongly that anatomical dif- ferences were more fundamental than similarities in shell and radular characters, and that Thiele’s system was arti- ficial. On the basis of his findings he re-grouped the cyclo- phorids into a superfamily with 5 families: Cyclophoridae (with 2 subfamilies); Maizaniidae (which THIELE, 1929, had considered to be a subgenus of Ostodes); Poteriidae (with all the New World and Pacific Island helicoid trop- ical genera, including Ostodes); Pupinidae (with 2 sub- families); and Cochlostomatidae, also with 2 subfamilies. Because he derived information from 3 separate organ- systems, Tielecke’s work is the most extensive and the soundest yet done on this group. Torre, BARTscH, & Morrison (1942) used only shell and opercular characters in classifying the American Cy- clophoridae into 4 subfamilies: Megalomastominae (elon- gate shells with corneous operculum); Diplommatininae (pupoidal shell); Amphicyclotinae (helicoid shells with corneous operculum); and Aperostominae (helicoid shells with calcareous operculum). Morrison (1955) described the external male repro- ductive systems of several American cyclophorids, and grouped all the genera into 2 families: the Amphicyclo- tidae with an enclosed, tubular vas deferens; and the Neo- cyclotidae, with an open seminal groove ascending the penis. TuHompson (1967, 1969) used the concept of a single large family, Cyclophoridae, recognizing the subfamilies Megalomastominae, Neocyclotinae, and Crocidopominae in Neotropical taxa, based on several characters in both male and female reproductive anatomy. My own work has been mostly at the species level. I agree with Sotem (1959: 182 and unpublished MS.) that the change from open seminal groove to closed vas de- ferens is by itself insufficient grounds for separation of families. I accept Tielecke’s system and consider the cy- clophorids as a superfamily containing 5 families, as fol- lows (TIELECKE, 1940: 365, 366): Cyclophoridae, with Vol. 20; No. 3 subfamilies Cyclophorinae and Spirostomatinae; Mai- zaniidae; Poteriidae; Pupinidae, with subfamilies Pupi- nellinae and Pupininae; and Cochlostomatidae, with subfamilies Diplommatininae and Cochlostomatinae. The family Poteriidae has representatives in Central America, South America, the West Indies and the South Pacific. It is characterized anatomically in the male by the location of the penis on the dorsal mid-line of the head, behind the tentacles, while in the female the oviduct, seminal receptacle, and bursa copulatrix all enter the uterus via a common genital duct. Tielecke dissected four of the genera: Poteria Gray, 1850, from Jamaica; Amphi- cyclotus Crosse & Fischer, 1879, from Central America; Aperostoma Troschel, 1847 (as Cyrtotoma Morch, 1852) from Mexico; and Ostodes Gould, 1862 from Samoa. Tuompson (1967, 1969) has dissected additional taxa, although coming to very different systematic conclusions. III. PREVIOUS STUDIES OF THE GENUS Ostodes Early work was purely descriptive, as expeditions and traders brought back material for study. The 1840’s to 1870's were the height of land snail descriptive work, with full-time efforts devoted simply to naming the flood of new materials. Because the association of some names with actual populations has presented problems, detailed item- ization of early descriptions is necessary. Between 1838 and 1842, the United States Exploring Expedition, under the command of Charles Wilkes, vis- ited various South Pacific Islands, including Upolu, West- ern Samoa. Shells were collected principally by J. P. Cout- houy, assisted by C. D. Pickering (anthropologist), Joseph Drayton (artist), and W. D. Brackenridge (botanist). Upon the Expedition’s return, the mollusk collection was shunted from one depository to another, badly mishan- dled, and finally turned over (in part) to Augustus A. Gould for study and description (JoHNson, 1964: 6-11). Gould read his descriptions of Cyclostoma tiara, C. stri- gatum, and C. plicatum in 1847. His descriptions were published the following year (GouLp, 1848), and subse- quently expanded in the text of the U. S. Exploring Ex- pedition (Gouxp, 1852). The Expedition’s Atlas of Shells finally appeared several years later (GouLD, 1860), with the three species of Ostodes illustrated on plate 8. Reciuz (1851: 213; plt. 6, figs. 10, 11) described and figured Cyclostoma apiae, collected by M. Charbonnier, from Apia, Upolu. PFEIFFER (1854: 301 — 302; plt. 40, figs. 13, 14) described and figured Cyclostoma pulverulentum from Upolu, and in the same year HOMBRON & JACQUINOT (1854: 50, plt. 12, figs. 25-28) described and figured Cy- THE VELIGER Page 193 clostoma albida from Samoa, without further locality. SOUVERBIE (1858: 294; plt. 8, fig. 6) published a descrip- tion and illustration of Cyclostoma gassiesi from an un- known locality. GouLp (1862: 283) proposed the generic name Ostodes for the group of 7 Samoan species previously included in Pfeiffer’s Cyclophorus, section 6. The cited differences of these 7 species from those remaining in Cyclophorus were the elongated shell form, simple aperture, deep, spiral umbilicus, and closer resemblance to the assimineid genus Omphalotropis. Mousson (1865: 180) described Cyclophorus upolensis, collected on Upolu by M. Graeffe. Four years later the same author (Mousson, 1869: 351; plt. 14, fig. 9) de- scribed Cyclophorus (Ostodes) adjunctus from Tutuila, Samoa. Cox (1870: 85) described Cyclostoma brazieri, col- lected by Brazier from under decaying logs on the moun- tains of Upolu, Navigator’s Islands (Samoa). I have ex- amined the holotype of C. brazieri, (Australian Museum C64837) and find it to be definitely not an Ostodes, nor even a poteriid; it probably belongs to the Assimineidae. GarrETT (1887: 124-153) published the only geo- graphical survey of Samoan land snails, drawing on his own experiences and collections in the field. KoseEtt (1902: 153) summarized the genus and listed 16 Polynesian species. THIELE (1929: 99) combined Os- todes with an African genus, Maizania, and figured a radula from the latter group. Only one of the above authors gave data on other than shell, operculum, and radula. Gou.p (1852: 103) reported that the animal of Ostodes plicatus had a long muzzle like a proboscis, long tapering tentacles, and distinct eyes. Its foot was ‘‘pale ochreous, upper part of head, neck, and sides a faint red.” TIELECKE (1940) examined the anatomy, at least in part, of cyclophorids belonging to 44 species and 19 ge- nera. Most of the material came from the Sunda-Archipel Expedition of Dr. Rensch, and some from the collection of Dr. Degner in the Zoologisches Museum, Hamburg. Specimens identified as Ostodes strigatus were available from Apia, Upolu. Their true identity is uncertain, since O. strigatus, although originally described from Upolu, is now known only from Tutuila, American Samoa. Tiel- ecke examined the male and female reproductive tracts, the central nervous system, and the pallial cavity, partic- ularly in reference to the kidney and circulatory system. CLENCH (1949: 4—29) revised the Pacific Island cyclo- phorids in the collection of the Bernice P. Bishop Mu- seum, Honolulu. He worked only with shell, operculum, and radula, and did not study the soft anatomy. Clench described and figured 3 new species (Ostodes cookei, O. savaii, and O. garretti), and refigured 5 previously known Page 194 species. Earlier authors, such as PEASE (1871: 475), had synonymized O. plicatus and O. strigatus, but CLENCH (1949: 14) stated that they were distinct and that Gould ‘was in error in assigning this species [O. strigatus] to Upolu rather than to Tutuila.”” Clench synonymized Cy- clostoma gassiesi with Ostodes plicatus. I have seen some of the material studied by Clench. Of the 15 shells in one lot identified as O. plicatus (MCZ 140507), two are O. plicatus, the rest are O. gassiesi. Since anatomical data led to my recognition of their distinctness, this confusion could only be expected. From extralimital areas, Clench described a number of species and genera. These are men- tioned in the section below on other Pacific Island poteriid genera. IV. MATERIAL STUDIED Most of this study was based on some 900 live-collected snails in the collection of Field Museum of Natural His- tory, Chicago (hereafter FMNH). These specimens were collected on Upolu and Savaii, Western Samoa, by Dr. Alan Solem and Mr. Laurie Price in October through December 1965, and by Mr. Price on Tutuila in March and April 1975. This field work was supported by Na- tional Science Foundation Grant GB-6779 to the Field Museum in support of research by Dr. Solem, and by Of- fice of Endangered Species contract 14-16-0008-873 to Field Museum to survey endangered land snails of Amer- ican Samoa. The animals had been drowned in water to which chloral hydrate had been added as a relaxant, and then preserved in 70% ethyl alcohol. Shells and soft parts were stored together. Specimens of poteriids from New Caledonia and the New Hebrides in the Field Museum collection were dis- sected for comparative purposes, as were specimens of Mexcyclotus panamensis, from Panama. The latter are figured for outgroup comparison. Some shell material was borrowed from the Museum of Comparative Zoology (hereafter MCZ), Harvard Univer- sity, and from the United States National Museum (here- after USNM), Washington, to check on published identi- fications and to verify some of my own identifications, V. METHODS For each specimen whose sex could be determined, 10 shell parameters were recorded. Height, diameter, um- bilical width, and horizontal diameter of the aperture were measured with a vernier caliper or (for very small specimens) an ocular micrometer. Whorl count, and both THE VELIGER Vol. 20; No. 3 dorsal and ventral sculptural elements were counted with a binocular microscope at 10X magnification, spire angle was estimated by holding the shell against a printed pro- tractor, and thickness of the parietal callus was estimated on a scale of one to three (one indicating a very thin callus, and three indicating a parietal callus as thick as the outer rim of the aperture). The character of the umbilical mar- gin was scored as being smoothly rounded, sharply angled, or bordered by a rim composed of the innermost ventral spiral lira. Ratios of shell height to diameter, and of shell diameter to umbilical width were calculated for each specimen. Empty shells, and snails that had been so severely con- tracted during preservation that they could not be ex- tracted for sexing, were not measured in detail. Where many specimens in a lot were severely contracted, some shells were broken after measurement to allow sexing. Specimens were accepted as adult that had started a changed pattern in growth, indicated by the following 3 features: (1) thickening of the parietal callus; (2) a change in the sculptural pattern on the terminal portion of the body whorl; and (3) an increase in the rate of decoiling resulting in the last portion of the body whorl being inset under the periphery of the penultimate whorl. In many snails the onset of reproductive activity is marked by just these alterations in growth patterns, when the energy budget of the individual shifts from individual growth to production of the next generation. Characteristic ex- ternal genital features of both male and female Ostodes develop while the shell is still very small, although the internal reproductive systems remain small and undevel- oped. Neither material nor time was available for deter- mining exactly when sperm and egg production begins; much larger samples, preserved for histological examina- tion, would be required to establish the degree of correla- tion between shell growth changes and onset of reproduc- tive activity. In the absence of such material, specimens showing the growth changes were classed as mature, while those without the changes were considered juvenile. Only adult shells are cited in the species diagnoses, and means and standard deviations of populations are based on adult materials only. For males, the penis was measured with an ocular mi- crometer, the lengths of the thick proximal trunk and the tapering distal thread were recorded, and the ratio of thread to trunk was calculated. For female snails, shapes and orientations of anus and vaginal orifice profiles were noted. Internal reproductive anatomy was studied by dissect- ing, where possible, at least three males and three females of each species. Drawings were rendered by close collabor- ation of artist and investigator, with the specimen being Vol. 20; No. 3 THE VELIGER Page 195 dissected in progressive stages so that the artist could de- pict internal structures and fine details. The method of preservation used precluded histological studies. Opercula were removed and studied for approximately 25% of the specimens. Examples of each opercular type were studied with both ordinary and polarized light, then embedded in paraffin and sectioned. The presence of cal- cium deposit was tested for with dilute HCl. Radulae of Ostodes gassiesi, O. reticulatus, O. plicatus and O. Ilanero, as well as Gonatorhaphe sp. and Gassiesia sp., were prepared and photographed by Dr. Alan Solem with a scanning electron microscope as part of a coopera- tive research program between Field Museum of Natural History and the American Dental Association. Prepara- tion technique was that of SoLEm (1972). VI. SYSTEMATIC REVIEW A. Variations in Structural Features The basic systems used in species separation and classi- fication are: (1) the shell; (2) the operculum; (3) the radula; and (4) the gross anatomy. The patterns of varia- tion within Ostodes are here reviewed in order of their initial use in systematics. 1. SHELL FORM anp ORNAMENTATION The basic shell form in Ostodes is turbinate. The form varies from species to species, with O. gassiesi and O. pli- catus adults being narrowly turbinate (taller than wide), O. savaii being of approximately equal height and width, and the other species being more or less broadly turbinate, with O. garrett: having the least elevation of the spire. Juveniles of all species, even O. gassiesi and O. plicatus, are broadly turbinate, wider than high (Figure 1). The periphery of the body whorl in Ostodes tiara, O. garretti, O. adjunctus, and the ‘‘smooth” form of O. upol- ensis varies from moderately to sharply carinate. Adults of all the other species have round-shouldered body whorls, even when the early whorls were carinate. Ostodes gassiesi, O. plicatus, O. savait, O. tiara, and O. garrettt have deep, narrow umbilici (mean D/U ratio 3.31). O. strigatus, O. exasperatus, O. reticulatus, O. ad- junctus, O. upolensis, and O. llanero have wide open um- bilici, with a mean D/U ratio of 2.76. The width of the umbilicus is affected by 2 factors: the rate of whorl trans- lation (rate of decoiling) and the size of the generating curve, as measured by the diameter of the aperture (RAUP & STANLEY, 1971: 158). In two species with equal-sized apertures, that which decoils faster will have the smaller Figure 1 Shell outlines of Ostodes gassiest b - adult a - juvenile umbilicus. In two species with equal rates of whorl trans- lation, the one with the larger generating curve will have the smaller umbilicus. Of the 2 factors, the rate of decoil- ing has the greater influence in Ostodes. Table 1 shows Page 196 THE VELIGER Vol. 20; No. 3 the species of Ostodes arranged, from the top down, in order of increasing size of umbilicus: O. gassiesi with the smallest umbilicus at the top, O. llanero and the “stepped” form of O. upolensis at the bottom. (Table | compares juveniles of O. Ilanero with adults of the other species; Figure 23 compares the same specimens of O. Tanero with juveniles of the other species. The relative position of O. llanero with respect to the other species is the same. It therefore seems reasonable to assume that, if adults of O. llanero had been available for examination, they would still have fallen at the bottom of Table 1.) This arrangement also puts the species generally in order of rate of decoiling, fastest at the top and slowest at the bottom, and in approximate order of size of aperture, largest at the top, smallest at the bottom. If the data for pairs of similar species, such as O. tiara and O. garretti, or O. savaii and O. exasperatus, be compared, it will be seen that that species of the pair which decoils faster will have a smaller umbilicus and usually also a larger aperture. The pair for which the above statement does not hold true is the 2 forms of Ostodes upolensis. The stepped form decoils much faster than the smooth form (Figure 24) and should therefore logically have a smaller umbilicus. In- stead, its umbilicus is larger than that of the smooth form, which would lead one to suspect it must therefore have a much smaller aperture. Not so; the stepped form has a larger aperture than the smooth form. The reasons for this apparent violation of the rules governing shell mor- phology can be seen in the silhouette of the smooth form (Figure 24b). Unlike most other species of Ostodes and the stepped form of O. upolensis, this form retains a pro- nounced peripheral carina which persists into adulthood. The carina protrudes sideways and increases the diameter of the shell in proportion to its umbilicus. Also, the inset of the terminal portion of the body whorl under the pe- nultimate whorl is proportionally much greater in the smooth form of O. upolensis than in the other species of Ostodes, so that the umbilicus is impinged on and made smaller. The aperture ranges in shape from round to very slightly compressed, and is appressed to the penultimate whorl. The outer apertural lip is simple, with no flange or ornamentation. The thickness of the parietal callus varies with the species and the age of the individual. The spire and early whorls are smooth in all species of Ostodes. Although the smoothness might be due to wear, I think it is more likely the true primary condition of the shell because (a) it is seen even in very small, unworn juveniles and (b) the sculpture shows a distinct pattern of beginning from a smooth surface, as discussed below. Except for Ostodes cookei, which is smooth, Ostodes shells are ornamented with spiral lirae, radial plicae, or both, according to the species. All species (except O. cooket) have spiral lirae, although they are quite unob- trusive in O. plicatus. The first spiral lira begins low down on the whorl, just above the suture, at about the beginning of the third whorl. The second spiral thread starts a little further along the whorl, and a little higher up, and so forth, until the full complement of spiral lirae is present by about the start of the fourth whorl. The spiral lirae are usually evenly distributed on the whorl, except in O. gassiesi, where they are clustered on the lower portion. The spirals continue in juvenile shells right up to the aperture; in adults, they fade out a few millimeters Table 1 Mean Diameters of Umbilicus and Aperture in Relation to Shell Diameter in Ostodes Species N D/U D/A H/D Spire angle (°) Ostodes gassiest 126 3.42 + .04 2.40 + .01 1.07 = .01 80 + 0.5, tiara 15 Beale) Se, IY 2.40 + .03 0.79 = .O1 107 + 0.9 plicatus 57 Ba Se {0} Dei =: {02 1.06 + .01 Gil ae Jit savaui 103 3.18 + .04 2.43 + .01 0.98 = .01 86 + 0.7 garretti 4 3.18 2.51 0.78 108 adjunctus 17 3.02 = .09 2.74 = .05 0.75 = .O1 92 + 1.9 exasperatus 24 2.98 + .09 PS se (0 0.92 + .01 90 + 1.6 upolensis (smooth) 10 2.91 + .19 ADO se Pl 0.76 += .05 98 + 4.0 strigatus - 84 2.89 = .02 Dis] a= {Oil Q:91 == 701 86 + 0.8 reticulatus 12 2.78 += .04 5) ==, 108) 0.93 + .02 OP se Il.3} upolensis (stepped) 12 2.37 = .08 Pe) 3= S08) 0.78 + .O1 98 + 2.6 llanero 3 2.37 2.61 0.70 110 Vol. 20; No. 3 before the aperture, at about the point where the body whorl is indented under the penultimate whorl. This probably marks the point at which reproductive maturity is reached. The spirals are continued on the lower palatal surface in O. reticulatus, O. exasperatus, O. llanero, and O. adjunctus. There may be traces of spirals on the ventral surface in O. upolensis, O. savait, O. tiara, and O. garretti, but the lower surface of O. gassiesi and O. strigatus is al- most always smooth, while that of O. plicatus shows only radial plicae. Radial plicae are an important element in the orna- mentation of Ostodes gassiesi and O. reticulatus, and are the major sculptural element of O. plicatus. In O. pli- catus they are close-spaced, moderately broad and in high relief. They continue over the shoulder of the body whorl, across the ventral surface and down into the umbilicus. At the edge of the umbilicus, the radial plicae may some- times be pinched into an acute angle, but they do not form a true rim around the umbilicus in the same way that a spiral lira would. The radial plicae of O. gassiest are somewhat broader and shallower than those of O. pli- catus; they hardly ever continue onto the lower palatal surface of the shell. The radial plicae of O. reticulatus are somewhat more widely spaced and a little narrower, so that there are definite valleys between them. The cross- ing of these valleys by the overlying spiral lirae creates the reticulated surface which gives the shell its name; it is par- ticularly noticeable on the ventral surface of the shell. Radial plicae are but poorly defined in O. strigatus, O. savaii, and O. upolensis, and are absent in O. exasperatus, O. llanero, O. adjunctus, O. garretti, and adults of O. tiara. The periostracum in Ostodes varies from light yellow to moderately dark brown. It is adherent in O. gassies: and O. tiara, but is deciduous in the other species. Possibly the most deciduous periostracum is that of O. plicatus. The color of most Ostodes shells is white, off-white or cream-color. The only exceptions are O. gassiesi, O. ad- junctus, and O. plicatus, which (sometimes, half the time, and often, respectively) have pink or red spires. If a pink spire of O. plicatus be broken, and the broken edge ex- amined under the microscope, it will be seen that the shell is composed of three layers: the inner and outer layers are a translucent white, but the middle layer is a very intense rose-red to salmon color. It looks pinkish from the outside of the shell because of the white layer covering it. In O. adjunctus, the color is deeper and the inner white layer is absent. Shells of Ostodes are subject to wear in varying degrees. The factor of wear is important because a badly worn shell is very difficult to place with certainty within a spe- cies, especially if it is empty so that anatomical informa- THE VELIGER Page 197 tion is lacking. Of all the Ostodes, O. reticulatus and O. upolensis seem to show the greatest resistance to wear, while O. tiara and O. garretti are most apt to be found badly worn. Mousson (1865: 179) says of O. tiara that even fresh specimens are almost always badly worn, with not only the periostracum missing, but most of the sculp- tural elements also gone, so that the shell presents a bony- looking, matte surface. This may reflect space niche differ- ences in microhabitat selection, but no data on this aspect are available. 2. OPERCULA (Figures 2, 3) CLENCH (1949) described the opercula of Ostodes adjune- tus, O. cookei, O. plicatus, O. strigatus, O. tiara, and O. upolensis, and figured but did not describe that of O. savaii. He mentioned only one type of operculum for each species. The only differences between species cited were concerned with the degree of central depression and the character of the nucleus. During this study, 198 Ostodes opercula were studied in detail. This sampling consisted of adult examples of both sexes from all species except O. llanero, for which only juveniles were available. Although all opercula were found to be circular, corneous, and transparent, there were 6 variations of this pattern. They are: Type A-1: laminate, thin (Figure 2) Of fairly uniform thickness throughout except for very thin and fragile outermost edge, which resembles cello- phane in texture and transparency. Slight thickening sometimes found just inside this ‘cellophane edge.” No spiral structure can be seen with either ordinary or polar- ized light. Central portion of operculum often shows in- clusions resembling air-bubbles or fragments of mucus. Disconnected fragments of spiral lamellae occasionally seen around outer margin. Thin horizontal layering seen in cross-section. Specimen illustrated, 5.1 mm in diameter. Type A-2: laminate, thick (Figure 2) Cross section shows approximately same number of horizontal layers as in type A-1, but each layer is much thicker. Operculum thicker in center and near outer rim than in circum-central area. Upper surface flat or very slightly concave. No spiral structure apparent in central part, which shows inclusions as in type A-1. Three or 4 spiral lamellae may be seen around outer edges. No ‘‘cel- lophane edge.’ Lower surface shows large central papilla, formed by a dipping-down of the horizontal layers, and Page 198 THE VELIGER Vol. 20; No. 3 Figure 2 Opercular Types A-1, A- 2, and B many faint, fine, closely-spaced radial lines, like brush- marks in wet paint. With transmitted polarized light, many faint, closely-spaced spiral lines can be detected, apparently within the body of the operculum, as they do not show on either surface in any kind of reflected light. Specimen illustrated, 5.9mm in diameter. Type B: oligogyrous (Figure 2) Extremely thin throughout; sometimes flat, sometimes cup-shaped, with flat central portion and edges turned up all around; in Ostodes strigatus, may be so acutely concave as to approach a cone-shape. Few (3—5) broad, irregular spirals from center to edge. Nucleus often somewhat off- Vol. 20; No. 3 THE VELIGER Page 199 Figure 3 Opercular Types C-1, C-2, and C-3 center. Strongest features of upper surface are many Ir- regularly placed, raised, radial cords. They show up quite well under ordinary light, and are of extreme brightness with transmitted polarized light. Cords occasionally em- phasized by parallel (i.e., radial) inclusions of opaque, white, non-calcareous material, possibly mucus. Under- side irregular in contour and without elements of true radial or spiral relief. Only very faint traces of horizontal layering seen in cross-section. Specimen illustrated, 3.7 mm in diameter. Type C-1: polygyrous, thin (Figure 3) Thin and flexible, of fairly uniform thickness through- out, except for “cellophane edge.’’ No central papilla on undersurface. Flat or concave, depending on contracture of specimen. Upper surface shows many narrow, tightly organized, clearly defined spirals, lying flat and slightly overlapping. Many of these opercula, especially those found in Ostodes plicatus and O. upolensis, show a silvery, iridescent sheen on the upper surface, which obscures the spirals under ordinary light. With transmitted polarized Page 200 light, the sheen disappears and the spirals are revealed. Specimen illustrated, 2.3mm in diameter. Type C-2: polygyrous, thick, flat (Figure 3) Basically flat, with slightly concave central portion; nearly all examples about 4 times as thick near outer mar- gin as through center. Many spiral lamellae, elevated at about 45° angle, surrounding smooth, depressed, central portion. Lamellae thick at base, with outer edges very thin and flexible, curving over inter-lamellar air-space, to touch next outermost lamella and producing a smooth outer surface like that of type C-1. Specimen illustrated, 2.7mm in diameter. Type C-3: polygyrous, thick, concave (Figure 3) Of more uniform thickness from center to edge than type C-2, showing horizontal layering in base more clearly. Most have pronounced central papilla; all very deeply concave. Upper surface resembles type C-1 in having many narrow, clearly defined, tightly organized spirals. Inner structure closer to type C-2 in that spiral lamellae stand up from their bases. Lamellae differ from those of type C-2 in 3 ways: they are generally stouter and heavier; they are differently shaped, being thicker at the upper surface than at the base, and having no thin, overhanging edge; and the spaces between lamellae are not air-spaces, but are filled with an opaque white material, which is not calcareous; it may be mucus. Specimen illustrated 6mm in diameter. Discussion: The 6 opercular types are not sex-linked, and are not species-specific. Table 2 lists the distribution THE VELIGER Vol. 20; No. 3 of opercular types by species. Three species, Ostodes llanero, O. reticulatus, and O. upolensis, are known to have only one type of operculum. However, in each case, the opercular type concerned is also found in at least 3 other species. Larger samples of these 3 species probably would include additional opercular types. One type of operculum, type C-3, has been found only in O. tiara, which also shows type A-2. I would expect to find addi- tional examples of type C-3 in O. garretti, if more indi- viduals of that species were available for examination. Type G-3 is a large, heavy operculum, which would accord as well with the large, heavy shell of O. garretti, as do the already-found types A-2 and C-2. From available evidence, I conclude that in Ostodes opercular type is not, as has been previously indicated, a diagnostic character at the species level. The only correlation found between opercular type and habitat is in the 6 specimens of Ostodes savaii which had type C-2 opercula. These 6 snails came from a collection station (Station 32, Savaii) having a more pro- nounced dry season than the other stations on the same island. No species other than O. savaii were found there. Of 7 specimens examined for opercula from this lot, 6 had type C-2; the 7th, as well as all other specimens of O. savaii, from more uniformly humid places, had thinner opercula (types A-1, B and C-1). A thick operculum would of course convey a selective advantage in a dry season, offering rather more protection against desiccation. Even this correlation breaks down, however, on Upolu. Of 21 collection stations on that island, there is a cluster of 5 (Stations 5, 7, 14, 16, and 17) that has the same climate Table 2 Distribution of Opercular Types by Species Species A-1 A-2 B Ostodes adjunctus 1 exasperatus 7 2 4 garretli 1 2 gassiest 19 9 llanero plicatus 8 reticulatus 8 savait 14 3 strigatus 7 3 15 lara : 3 upolensis Totals 64 10 32 C-1 C-2 C-3 Totals 14 15 3 ih 23 1 4 7 35 3 3 11 19 8 7 6 30 16 2 43 5 8 10 10 68 19 5 198 Vol. 20; No. 3 as Station 32 on Savaii. Of 17 snails examined for opercula from these 5 stations, 15 were found to have thin opercula. One individual from Station 14 had a thick operculum that appeared to be of type C-1, but with many added layers. There were no upstanding lamellae, as in types C-2 and C-3, there was no central papilla, as in type A-2, and the layers themselves were no thicker than usual for type C-1; there were simply more of them. One individual from Station 17 had a similarly thickened operculum, re- sembling type A-1, but with many more layers than nor- mal. The “‘dry stations’ on Upolu are in somewhat denser forest than the one ‘‘dry station” on Savaii; perhaps there is enough difference in micro-habitat to decrease the selection pressure for thick types of operculum. As to the very peculiar opercula designated type B, I think it is unlikely that they are replacements for opercula accidentally torn off: FRETTER & GRAHAM (1962: 82) de- scribe a replacement operculum as being thickest in the middle and tapering towards the edges, and without ‘‘dif- ferentiation of structure apart from being laminated,” which description does not fit type B at all. It is possible that type B opercula are the product of some disease proc- ess or injury to the snail, although no gross abnormalities of shell or body were observed in the snails from which they were taken. Of 43 animals of Ostodes strigatus ex- amined for opercula, 15 had type B opercula, and of these, 8 were so deeply concave as to be cone-shaped, with the greatest diameter of the operculum much less than the diameter of the aperture. The snails with these peculiar opercula looked perfectly normal, except that the back of the foot, beneath the operculum, was deeply hollowed out to accept the pointed end of the cone. One such snail, a female, was very carefully dissected, in search of any anatomical abnormality; none was found, except for the hollow place in the foot. Two of the cone-shaped opercula were greatly thickened by a multiplication of layers. Two other very thick opercula were found in O. strigatus; one was type B (ordinary flat form), and the other was type A-1. The 6 opercular types described above have been ob- served. The reasons for their existence remain to be ex- plained. It would seem that protection against desiccation or predation could be achieved as effectively by a simple multiplication of layers as by the more elaborate methods of thickening each layer as in type A-2 or creating up- standing lamellae, as in types C-2 and C-3. I have no explanation of these differences. THE VELIGER Page 201 3. RADULAE (Figures 37-48) Cyclophorid snails have a taenioglossate radula, with one lateral tooth and two marginal teeth on either side of the central tooth. Photographs taken with a scanning electron microscope (SEM) show slight differences between genera and species which have not heretofore been noticed. For this study, SEM photographs were taken of the radulae from Ostodes gassiest, O. plicatus, O. reticulatus, and O. lanero, plus examples of Gonatorhaphe sp. and Gassiesia sp. for comparison. The results are summarized below. Central (rachidian) tooth: In all species examined, the central tooth is tricuspid. The base of the tooth is greatly thickened and deeply hollowed-out from below, with a smaller hollow on each side. When the radula is in the folded position, the central teeth lie in their longitudinal row without touching each other, the curved top of one resting in the space made available by the hollowing of the base of the next anterior tooth (Figure 47). The shape of the central tooth varies slightly among the species studied. In Ostodes gassiesi it is peg-like, appearing of equal width at the level of the cusps, in the middle of the shank, and at the base. The other 3 species of Ostodes show central teeth that are slightly flared at the base, and O. reticulatus and O. llanero also show a slight narrowing above the flare. In Gonatorhaphe there is a pronounced narrowing above a moderately flared base. Gasstesia shows no narrowing, but a very strong flare is present. Lateral tooth: ‘The lateral teeth are about twice the length of the central teeth. When the teeth are not in use, the top of each lateral tooth rests against the shank of the tooth next anterior to it, at about the mid-point of the shank (Figures 37, 41). In the 4 species of Ostodes that were examined, the lateral tooth has 4 cusps, of which the most medial is the smallest and the third from the inside is by far the largest. All Ostodes lateral teeth have long, slender shanks. In O. gassiesi and O. reticulatus, there is a medially projecting flange that seems to join the tooth to the radular membrane. The flange is less clearly seen in the photographs of O. plicatus and O. llanero, but it is shown in Thiele’s drawing of the radula of Maizania preussi (THIELE, 1929: 99; fig. 76). In Gonatorhaphe sp. the lateral tooth still has 4 cusps, but the medial one is much reduced. The shank is long and slender, and there is no flange. In Gasszesia sp. the long shank has a flange, Page 202 Ostodes gassiest BQ Ostodes plicatus baa Ostodes reticulatus io See Ostodes llanero aa cae Gonatorhaphe sp. Se Gassiesia sp. CTs ALP Cn CTf THE VELIGER Vol. 20; No. 3 but the tooth has only 3 cusps — the most medial cusp has disappeared altogether. Marginal teeth: The viewing angles are such that the inner marginal tooth cannot be seen consistently and clearly enough for comparative discussion. The outer mar- ginal tooth is seen clearly, however, and shows more vari- ation than any of the other radular teeth. The tooth al- ways has 3 (some views of Ostodes gassiesi seem to show 4) cusps. The cusps are pointed, except in O. plicatus, where the outermost one is truncated. Below the outermost cusp, the outline of the tooth is first deeply indented and then protrudes into a sharply angled “shoulder.” In the folded position, the outer edge of the inner marginal tooth is cradled in the indentation of the outer marginal tooth (see Figures 38 and 46). Below the ‘“‘shoulder” a shank of moderate length connects the tooth to the radular mem- brane. In the lower extremity of the shank there is a fora- men through which the radular membrane passes. This arrangement permits the tooth to grasp the edge of the membrane firmly, but with the potential for a great deal of movement, due to the size and shape of the foramen. The angles of the indentation and ‘‘shoulder,’’ as well as the character of the foramen, vary from species to species, and are summarized in Table 3. The functional signifi- cance of the differences in the foramen is unknown. Discussion: Figure 4 shows 3 teeth from one row of the radula of each of the 6 species studied, presented together for ease of comparison. The differences between the 4 species of Ostodes are so slight that I do not feel they could be used as a diagnostic character. The teeth of Gonotorha- phe seem to be a sort of mixture: the central tooth re- sembles that of O. reticulatus, the lateral tooth resembles the lateral tooth of O. llanero, and the outer marginal tooth is very similar to the corresponding tooth of O. gas- siesi. I would hesitate to try to differentiate between Gono- torhaphe and Ostodes on the basis of their teeth. The teeth of Gassiesia, however, are very different from those of the other 2 genera. The strongly flaring central tooth; the lateral tooth with only 3, not 4, cusps; and the very broad, low foramen in the outer marginal tooth set this (< adjacent column) Figure 4 Comparison of Radular Teeth of Six Poteriid Species BH - basal hollow CTf£ —- frontal view of central tooth CTs - side view of central tooth FL - flange FO - foramen LT -— lateral tooth OMT -— outer marginal tooth RM -— radular membrane Vol. 20; No. 3 THE VELIGER Page 203 Table 3 Character of Outer Marginal Tooth Species Angle of indentation Angle of shoulder Character of foramen Ostodes gassiesi 30°— 45° 60° —90° broad, squarish Ostodes plicatus 75° 90° broad, low Ostodes reticulatus 60° 90° broad, low Ostodes llanero 45° 90° tall and narrow Gonatorhaphe sp. 45° 90° very tall, pointed top Gassiesia sp. 60° 90° very broad, low radula apart from the others quite clearly. These radular differences are sufficiently clear-cut to be a useful diag- nostic character. 4, GROSS ANATOMY a. General: Uppermost part of visceral hump occupied entirely by testis in male; in female, ovary lies on colu- mellar side, digestive gland on outer side. Middle portion of visceral hump occupied entirely by digestive system in both sexes, except for passage of gonoduct on columellar aspect. Lowest part of visceral mass in both sexes has re- productive organs on columellar side, digestive system on outer side. Position of organs in pallial cavity as follows: on right side, gonoduct runs below rectum from mantle line nearly to anterior opening of cavity. On left, kidney lies between layers of mantle, extending forward to about middle of pallial cavity. Pallial renal orifice not located. Pericardial cavity, with heart, nestles against lower middle portion of kidney. Hypobranchial gland J-shaped, beginning to left of mid-line, passing transversely behind kidney, then turning to run anteriorly between upper renal margin and rectum. Point of origin of transverse portion, and length and thickness of anterior portion of hypobranchial gland differ in the several species, and will be discussed individually. Anterior portion of mantle thin, fragile, vascular, except for thickened leading edge. b. Digestive system: In female, digestive gland reaches apex of visceral mass, surrounding ovary for approxi- mately first whorl, except on columellar side. In male, di- gestive gland begins at lower edge of testis. Stomach em- bedded in outer side of digestive gland and visible through integument. In both sexes, stomach and digestive gland together occupy entire lumen of whorl, in middle section of visceral hump, except for passage of gonoduct and esophagus on columellar side. Massed loops of intestine occupy outer half of anterior visceral hump, with most distal segment crossing from left to right just behind mantle cavity. Intestine continues forward as rectum be- tween layers of mantle on right side. Anal orifice just in- side leading edge of mantle. In males, anal orifice points straight forward. In females, orientation of anal orifice and degree of hypertrophy of its edges are species-specific. c. Male reproductive system (Figure 5): Testis fills en- tire lumen of first 1-2 whorls, depending on age of indi- vidual. Color ranges from uniform creamy white through tan and brown to very dark grey. Younger individuals have smaller and lighter-colored testes. Color caused by pigment granules which appear on surfaces of testicular tissue lobules, and in interlobular spaces. In young adults they are light brown, small in size, and scattered sparsely but fairly evenly, giving whole organ a tan color, rather than the cream of the juvenile. As snail ages, granules increase in size, number, and intensity of color, causing whole organ to appear darker and darker. In an occasional individual, the granules will cluster into scattered black blotches; sometimes granules are also found on the inner surface of the encapsulating membrane. The pigment granules, whether clustered or single, are not usually firmly adherent to testicular tissue, but may be picked off fairly easily, revealing tissue itself, still creamy white underneath. Testicular tissue consists of a number of multiple- branched trees of digitiform alveoli, arranged in a single row on the outer side of a single collecting tube, which runs down the columellar side of the whorl. Each major alveolar trunk or duct divides into 2 equal main branches, which in turn divide and re-divide fairly symmetrically. Outermost tips of the terminal alveoli give surface of the organ a lobulated appearance. There are at least 3 major alveolar ducts near anterior end of testis, with an unde- termined number of progressively smaller ducts running back toward the posterior tip of the organ. Even in deeply pigmented testes, the collecting tube, and in some cases, the major ducts, are without pigment granules and thus appear white. Collecting tube runs forward as seminal vesicle on col- umellar side of visceral mass. In central visceral section, it lies lateral to the esophagus and ventral to the stomach. Page 204 THE VELIGER Vol. 20; No. 3 Figure 5 Elements of the male reproductive system in Ostodes "a — generalized composite male, view from above b — testis external appearance; c — testis, internal arrangement, dia- grammatic; d — cross-section of prostate; e - diagram showing folded structure of prostate, seen from underside; f — diagram showing prostate unfolded, seen from underside [for explanation of abbreviations see Appendix on foldout] Vol. 20; No. 3 After passing proximal tip of prostate, it turns sharply upon itself and runs back to enter ventral surface of prostate slightly anterior to its proximal end. Prostate creamy white, visible through integument on columellar side of body. Structure of prostate resembles a sandwich, sealed around the edges, and folded in half longitudinally, slightly off center. This structure gives a cross-section with 2 layers of bread (prostatic tissue) in the middle, the sandwich filling (lumen of the prostate) lying in a U-shape, and another layer of bread (prostatic tissue) around the outside of the U (see Figures 5d, he, 5f). Most proximal portion of prostate, which lies beneath distal end of digestive gland, is flattened dorso-ventrally, more anterior portions oval to round. Medial upper aspect of prostate touching intestinal loops. Prostate enters pallial cavity on right side, within mantle wall, below rectum and in angle formed by junc- ture of mantle and body. Prostate terminates in a blunt tip at about mid-point of pallial cavity. In some specimens, the prostatic tip is truncated. In others, there is a constric- tion of the medial half of the organ, a short distance be- hind the tip, resulting in the presence of a small pouch or sac on the anterior medial corner of the prostate. The presence of this sac is in part a function of age, in part of species. Juveniles almost never have sacs; adults are more likely to have sacs in some species than in others. Table 4 shows the frequency of occurrence of sacs, and also of a small, fragile duct, of varying origin and termination, which appears to be a “‘safety-valve” for venting excess sperm (see FRETTER & GRAHAM, 1962: 345, 348). There is no correlation between presence or absence of a sac and Table 4 Prostatic Characters (mixed ages) THE VELIGER Presence of Presence of Species Sac “Safety-Valve” Ostodes adjunctus Oof 4 Oof 4 exasperatus 2of 5 lof 5 garretti 2of 2 lof 2 fassiest 11 of 15 2 of 14 llanero Oof 1 Oof 1 plicatus lof 5 4of 5 reticulatus 3o0f 3 Joi. 3 savant 10 of 13 5 of 14 strigatus Oof 6 Oof 6 tiara (“smooth”) lof 5 4of 5 tiara (“bumpy”) lof 1 lof 1 upolensis 3of 4 4of 4 Page 205 presence or absence of a “‘safety-valve.” If a ‘‘safety-valve” occurs in a specimen having a sac, the most usual origin of the duct is the dorsum of the sac. In a snail without a sac, the most usual origin of the duct is the medial anterior corner of the truncated prostate. In either case, the duct usually runs laterally and anteriorly across the base of the vas deferens, and terminates beneath and slightly behind the anal orifice. Variations from these patterns are men- tioned in the species descriptions. Vas deferens originates from anterior lateral corner of prostate. It is a closed tube running anteriorly and me- dially beneath the skin to the base of the penis, where it passes upwards, lying at first near the surface, but pene- trating progressively more deeply, to reach the center of the penis about half-way up the trunk. TIELEcKE (1940: 332) states that males of Ostodes strigatus have an open sperm groove. I have not seen the specimens on which he based his statement, and so do not know what species he actually had before him. The 64 males of O. strigatus available to me agree with Gould’s original description and figure for O. strigatus (GouLp, 1848: 204—205; 1852: 102—103; 1860: plt. 8, figs. 117, 117a, 117b), and they all have closed vasa deferentia. Penis on cephalic mid-line, just posterior to tentacles. Base usually thick, somewhat rugose, probably partly con- tractile but not introversible. The ovoid to rounded trunk narrows gradually to abruptly into a thin distal thread, which may be long or short in proportion to the trunk. Some specimens have a sub-terminal bulb or swelling on the thread; there is no correlation between presence or absence of a bulb and length of thread. Penis usually car- ried folded sharply back from its base, the whole length of the trunk lying flat along the neck and body inside the mantle cavity. There is another sharp fold at the base of the distal thread, which lies folded against the trunk, the tip pointing forward. In the case of a very long thread, or a thread with a sub-terminal bulb, the tip sometimes may be found folded under. The ratio of thread-length (in- cluding taper) to trunk-length is diagnostic at the species level. Ratios for adults of the several species are found in Table 5. For discussion purposes, a specimen is considered to have a short thread if the ratio of thread to trunk is 1/1.6 or more. This ratio was chosen as the dividing point because, on a graph showing the thread to trunk ratio of every available specimen of Ostodes gassies: (a consistently short-threaded species) and O. plicatus (a consistently long-threaded species), 95% of O. gassiesi specimens had ratios of 1/1.7 or more, while 97% of O. plicatus speci- mens had ratios of 1/1.5 or less. Although the lengths of thread and trunk differ from species to species, the total length of the penis is fairly uniform, at least among the mid-sized species (Figure 6). Page 206 Table THE VELIGER Vol. 20; No. 3 5 Penial Characters Bulb present Ratio of thread length to trunk length (adult) on thread Short thread Long thread Species (mixed ages) Range Ostodes adjunctus 0 of 11 1/1.7-1/1.5 9 1/0.5-1/1.5 1/0.97 exasperatus (Upolu) lof 4 0 — — 4 1/1.05-1/1.5 1/1.22 (Savaii) 4of 6 4 1/2.5-1/7.3 1/4.68 2 both 1/1.5 1/1.5 garretti Oof 2 0 — - 2 1/1.4-1/1.6 1/1.5 gassiest (all) 7 of 12 53 1/1.7-1/11 1/4.73 3 1/0.86-1/1.43 1/1.12 (Upolu) 5of 7 36 1/1.7-1/10 1/3.8 2 1/0.86-1/1.43 1/1.13 (Savaii) 2o0f 5 17 1/2.7-1/11 1/6.7 1 — 1/1.1 llanero Oof 1 0 = = 1 — approx. 1/1.2 plicatus Oof 5 1 — 1/1.7 31 1/0.25-1/1.5 1/0.98 reticulatus Oof 3 0 — — 5 1/0.46-1/1.25 1/0.91 savant (all) 11 of 23 43 1/1.7-1/10 1/3.66 7 Wl -1/1.5 1/1.3 (Upolu) 2o0f 5 14 1/2.0-1/4.5 1/3.0 2 1/1.38-1/1.5 1/1.44 (Savaii) 9 of 17 29 1/1.7-1/10 1/3.97 5 Wl -1/1.5 1/1.24 strigatus 0 of 43 0 — _ 43 1/0.31-1/1.20 1/0.60 tiara (“smooth”) Oof 6 0 — — 6 1/0.54-1/1.5 1/0.93 (“bumpy”) 2of 2 2 thread/collar/trunk 0 — = 1/0.22/1.67 1/0.2/1.8 1/0.22/1.85 upolensis Oof 2 1 — 1/2.2 | 6 1/0.54-1/1.5 1/1.15 In both Ostodes gassiesi and O. savaii, short-threaded specimens from Savaii have shorter threads than compar- able individuals from Upolu. I have been unable to find any reason for this difference. There are no important conchological differences between snails from the two is- lands, nor are there significant differences between long- thread and short-thread snails of the same species. In O. exasperatus, all males from Upolu are long-threaded, while the Savaii population has 67% with short threads. The shells from the two populations are not significantly different, and the female reproductive systems are virtu- ally identical. Explanation of Figures 37 to 42 Figure 37: Ostodes gassiesi, central tooth and tetracuspid lateral teeth X 477 Figure 38: Ostodes gassiesi, radula in folded position, showing edge of radular membrane and foramina in outer marginal teeth X 509 Figure 39: Ostodes plicatus, central tooth, showing hollowed-out base; tetracuspid lateral teeth and inner marginal teeth x 478 Figure 40: Ostodes plisatus, edge of radular membrane and outer marginal teeth with foramina X 360 Figure 41: Ostodes reticulatus, central teeth, tetracuspid lateral teeth, and inner marginal teeth X< 368 Figure 42: Ostodes reticulatus, outer marginal teeth, showing fora- mina 1115 Tue VE.IcER, Vol. 20, No. 3 [Grrarp1] Figures 37 to 42 Vol. 20; No. 3 Ostodes tiara presents special problems. Sixteen male specimens, 9 adults and 7 juveniles, from 5 different lo- calities, were examined. One was a freak, having a penis with three threads. Ten, 6 adults and 4 juveniles, had standard long-thread penes. These (Figure 7a) are referred to in the tables as ‘‘smooth.”” Some males with “smooth” penes were found at each of the 5 localities. At Station 8, on Upolu, in addition to 3 males with ‘‘smooth’”’ penes, 2 adults and 3 juveniles were found with a very different 10 HG) pee ee = 5 +p = —+— “Ger =S2= G P S R S&t 12 — 1/9 gr 2 = 8 1/7 5° E E J oO I 2 6 se] Ue) z. E: 3 3 4 1/3 2 mL tt G PS RS&t Gi PS" Rast Figure 6 Lengths of penis thread, penis trunk, and whole penis, and ratios of thread length to trunk length, in five mid-sized species of Ostodes G — Ostodes gassiesi; P - O. plicatus; R — O. reticulatus S - O.savai; St — O. strigatus — _ vertical line — range of measurements; horizontal line — mean; hollow box — two standard errors of the mean on either side of the mean THE VELIGER Page 207 type of penis, referred to as “bumpy.” The trunk is sur- mounted by a white bump, resembling a knuckle. Beyond the bump isa constricted portion, or collar, which may be slightly reddish, and which is followed by a short swollen area which tapers to a short thread with a sub-terminal bulb (Figure 7b). Males with this type of penis have um- bilici averaging 15.4% larger than those of “smooth” penis males. The females from Station 8 also had larger umbilici than females from the other stations. No other statistically significant conchological differences were found, nor did the female reproductive systems from Station 8 have any unusual features. d. Female reproductive system (Figures 8, a, b; 9): Ovary begins slightly anterior to apex of visceral hump and occupies columellar side for approximately one whorl. Ovarian tissue composed of a mass of white granules of irregularly spherical contour, held together by an encap- sulating membrane, but not otherwise macroscopically organized. There is no system of collecting tubules equiv- alent to that of the male testis. The ovary tapers anteri- orly to a blunt point, with the encapsulating membrane continuing forward as the oviduct. Oviduct lies on col- umellar side of body, lateral to esophagus and ventral to stomach and digestive gland, running anteriorly nearly to mantle line, then reflexing sharply upon itself and con- tinuing up to approximately the level of the uterine fundus; reflexes again and runs downward for a short distance. Slightly below level of uterine fundus, oviduct becomes progressively thicker and begins to fold upon itself accor- dion-fashion in a single plane. There are from 3 to 6 fold- ings, each slightly deeper than the preceding one. The folds nearest the columella are laid down even with each other, with the increasing depth of the folds producing a triangular mass, lying ventral to the uterus, its long axis parallel to the long axis of the visceral hump. The folds are held together by an encapsulating membrane which is transparent and covered with brown speckles, which are most numerous in the valleys between folds. I believe that this area of folded oviduct serves as a seminal receptacle (Figure ga). Distal to the seminal receptacle, the now quite thick oviduct is joined, on its lateral aspect, by an equally thick duct which leads away from the columella to a large, heavily pigmented, bulbous structure, which I believe to be a bursa copulatrix. It lies partly beneath the uterus but mostly ventral to the mass of intestinal loops. If dis- tended, it can often be seen through the integument of the visceral mass, just behind the kidney at the mantle line. If not distended, it resembles a collapsed balloon, lying beneath the intestine. The pigment particles lie Page 208 THE VELIGER Vol. 20; No. 3 a EE ele ee Figure 7 Penis types in Ostodes tiara and Ostodes garrett: a — O. tiara, “smooth”; b - O. tiara, “bumpy”; c — O. garretti [for explanation of abbreviations see Appendix on foldout] Vol. 20; No. 3 THE VELIGER Page 209 a IEEE Frage 209 CDP Figure 8 Female reproductive system in Osfodes tiara and Ostodes garrett: a — O. tiara; b — O. tiara, ovary; c — O. garretti; d — O. garretti, entrance of common genital duct into uterus [for explanation of abbreviations see Appendix on foldout] Page 210 THE VELIGER ECPM Figure 9 Generalized details of female reproductive system in Ostodes a — cutaway views of bursa copulatrix and.seminal receptacle b — diagram showing elements of reproductive system in true anatomical relationship c — cross-section of uterus [for explanation of abbreviations see Appendix on foldout] Vol. 20; No. 3 Vol. 20; No. 3 partly on the covering membrane, and partly on the outer surface of the wall itself. The walls are thick and soft, and appear to be constructed of fibers laid vertically to the lumen of the organ. The inner surface is soft and creamy white. The duct protrudes part way into the lumen, which is sometimes found filled with a pearly white amorphous mass. Previous workers do not agree concerning the probable function and nomenclature of the folded oviducal area and pigmented sac. TIELECKE (1940: 331 — 332) called the folded area of Ostodes strigatus a seminal receptacle and the pigmented sac a bursa copulatrix. THoMPson (1969: 40-41), working on Central American cyclophorids, called the folded area an albumin gland and the pig- mented sac a seminal receptacle. Neither gives histological details to support his conclusions. Work on Pomattas elegans (CREEK, 1951) and Acme fusca (CREEK, 1953) did involve histological examination of the entire genital tract, extensive tissue staining studies, and laboratory observations of breeding and egg laying. In Pomattas elegans there is a widened muscular section of the renal oviduct which functions as a seminal recep- tacle, its walls being covered with large numbers of orien- tated sperm at all seasons of the year (CREEK, 1951: 606). There is a sac at the posterior end of the pallial oviduct, which although homologous with the seminal receptacle of many other gastropods, functions as a bursa copulatrix (CREEK, op. cit.: 636). This bursa opens directly into the mantle cavity, and the penis is actually inserted into this sac at copulation (CREEK, op. cit.: 606). In Acme fusca, however, the bursa opens to the mantle cavity by means of a duct, and the penis probably does not actually enter it during copulation (CREEK, 1953: 234). As in Pomatias, the anterior portion of the renal oviduct is convoluted and slightly swollen. This area, homologous to the seminal receptacle of Pomatias, probably functions in Acme as a fertilization chamber, as it has been found to contain in- gesting cells for the disposal of excess sperm. The structures in Ostodes are very similar to those found in Acme. Both genera show a thickening of the an- terior renal oviduct; both have a sac-like structure located near the upper end of the uterus; both have a duct leading from the mantle cavity to the sac, the renal oviduct, and the uterus. I do not know exactly where fertilization takes place, but the pigmented sac in Ostodes certainly seems to function as a bursa copulatrix, and I therefore agree with Tielecke and call the thickened area of the renal oviduct the seminal receptacle. Distal to the entrance of the duct from the bursa copu- latrix, the oviduct, now a common genital duct, again be- comes thin and fragile. The length of the common duct, from the junction of the ducts from bursa and receptacle THE VELIGER Page 211 to the mantle line, varies greatly from species to species (see descriptions). The common duct enters the pallial cavity and continues downward along the medial margin of the uterus, to which it is bound by a thin sheet of tissue. There is a small hole, the copulatory pore, in the medial aspect of the genital duct. This is located between the pos- terior margin of the mantle cavity and the point of entry of the genital duct into the uterus, its position varying with the species. The copulatory pore is the posterior ter- mination of a channel which is formed by an overlayment of the uterus onto the right side of the mantle cavity floor. The medial margin of the channel is formed by a very slight ridge in the body tissues, and when the mantle is in its normal position (not laid back as for dissection) the channel would function as a closed tube. There is no grossly visible specialization of the tissues forming the channel floor. Whether this channel serves to guide the penis to the copulatory pore, or as a passageway for sperm deposited at its anterior end, must remain a matter for speculation until behavioral studies can be made. Uterus lies along columellar side of body, partly within visceral hump beneath most anterior portion of digestive gland, and partly at right side of pallial cavity, below rec- tum. Uterus bi-lobed, ventral lobe occupying the middle half to three-fourths of dorsal lobe length. Dorsal lobe overlies upper end of ventral lobe, folding upon itself to form entire uterine fundus, and continuing forward, past anterior end of ventral lobe, to form entire vaginal por- tion of uterus. In cross-section (Figure gc) it is seen that the two lobes are entirely separate, except for a proximal end-to-end anastomosis, so that the lumen of the uterus is one continuous passage from the anterior end of the ventral lobe, up around the fundus, and down the length of the dorsal lobe to the vaginal orifice. The texture of the two lobes is quite different. The ventral lobe is com- posed of firm, densely packed, fine-grained tissue which is often pink in color, especially near the anterior end. The tissues of the dorsal lobe are creamy white, and ap- pear softer, more friable, and more loosely organized. Dorsal lobe passes forward, beneath the rectum, toward the front of the pallial cavity. Vaginal orifice near anus. The shapes, sizes, and orientations of these 2 openings are species-specific, and will be discussed in the species de- scriptions. 5. SEXUAL DIMORPHISM For this study, there were available 13 populations of Ostodes which contained both adult males and adult fe- males. Ten of the 12 species were included in the 13 pop- ulations: O. plicatus, O. reticulatus, O. upolensis, O. Page 212 height (mm) THE VELIGER Vol. 20; No. 3 Figure 10a Sexual dimorphism in height in ten species of Ostodes (adults) A — Ostodes adjunctus; E - O. exasperatus, Upolu population; e — O. exasperatus, Savaii population; G - O. gassiesi, Upolu population; g — O. gassiesi, Savaii population; Gt — O. garretti; P — O. plicatus; R - O. reticulatus; S - O. savaii, Upolu popu- tiara, O. garretti, O. adjunctus, and O. strigatus were rep- resented by one population each, while O. gassiesi, O. savati, and O. exasperatus were represented by one popu- lation from Upolu and another from Savaii. Only juvenile shells of O. llanero were available, and only one empty shell of O. cookei was seen. lation; s — O. savaii, Savaii population; St — O. strigatus; T — O. tiara; U — O. upolensis. Vertical line — range of measurements; horizontal line — mean; box — two standard errors of mean on either side of mean; hollow box — male; solid box — female When the shell measurements of males and females of the above-named 13 populations are examined, the usual pattern is seen to be that females are both taller and wider than males of approximately the same whorl-count (Fig- ures 10a, b). The only population in which this situation is reversed is that of Ostodes exasperatus on Upolu; at a very Vol. 20; No. 3 THE VELIGER Page 213 diameter (mm) 22 _ N late yu a a 8 Figure 10b Sexual dimorphism in diameter in ten species of Ostodes (adults) [for explanation of symbols see Figure 10a] Figure 11 Percentage differences in mean height and diameter between males and females in 13 populations of Ostodes. Species designations as in Figure 10a [__] difference in diameter 1 difference in height Page 214 slightly lower whorl-count, males average 0.75 mm higher and 0.69mm wider than females. The Savaii population of O. exasperatus follows the usual pattern of larger females. Male-female differences in height are slightly greater than differences in diameter in 10 of the 13 populations. The differences are usually rather small, nearly three- fourths of them being 9% or less, with the remainder widely scattered between 12% and 29%. Figure 11 shows the distribution of percentage differences. It is interesting that the greatest differences occur in the smallest and larg- est species; in little Ostodes upolensis, females average 18% taller and 15% wider than males; in O. adjunctus, the differences are 15% and 12%, respectively. In the largest species, O. tiara, females are 12% taller and 15% wider. The greatest differences of all occur in the second- largest species, O. garretti, its females are 21% taller and 29% wider than males at very nearly the same whorl count (XW =5.22 female, 5.19 male). B. Systematic Accounts 1. THe GENUS Ostodes Gould, 1862 Gov Lp, 1862, Proc. Boston Soc. Nat. Hist., 8: 283; KoBELT & VON MOLLENDoRFF, 1897, Deutsche Malak. Ges., Nachr., 29: 112; Kose tt, 1902, Das Tierreich, 16: 153, fig. 32 (O. plicatus); THiELe, 1929, Handb. syst. Weicht., 1: 99; TIELEcKE, 1940, Arch. f. Naturgesch., N. F., 9: 331, 332, 350, 361, 362, 364, figs. 7, 8, 19 (O. strigatus [?], male and female genitalia and nervous system); CLENcH, 1949, Bull. B. P. Bishop Mus., 196: 9-10. Type species: Cyclostoma strigatum Gould, 1848, by OD. Shell narrowly to broadly turbinate, spire elevated. Thin and fragile to thick, heavy, and strong. Small (3.2 mm high X 2.8mm wide) to large (12.5mm X 22.5 mm). Females average larger than males at same whorl-count, except Upolu population of Ostodes exasperatus. Four to 6 whorls, spire angle 70° — 120°. Aperture round, ho- lostomatous, appressed to whorl above; outer lip simple, parietal callus thin to thick. Umbilicus deep, narrow to wide; may be smoothly rounded, sharply angled, or bor- dered by rim. Apical whorls smooth. Later whorls show spiral lirae, radial plicae, or both, except O. cookei, which is smooth. Terminal portion of body whorl of adults in- dented under penultimate whorl, except O. adjunctus. Periostracum thin, yellowish to dark brown, adherent or deciduous. Color cream to dirty white, sometimes with pink to red spire. Operculum round, corneous, transpar- ent, thin to thick, with many or few volutions, or laminate. THE VELIGER Vol. 20; No. 3 Penis external, on dorsal midline of head behind ten- tacles, with closed vas deferens and terminal thread. Ovi- duct, seminal receptacle, and bursa copulatrix discharge into uterus via common genital duct, wherein is copula- tory pore. 2. KEY to THE SPECIES la Adult shell smaller than 7.5mm high by 8mm WIdE) vec ee citaon nae Mee 2 b Adult shellilargerithantaboveler to eee 4 27a Shell¥smoothy a. 5540 oatae eee O. cookei b Shell xwith distinct) sculpture!) eee eee 3 3a Strong spiral sculpture on ventral surface, umbilicus with bordering rim; from Teutuila::h3,) 54: c aerate too ke eee O. adjunctus b Ventral spirals weak or absent, umbilicus without arim; from Upolu or Savail ....... O. upolensis 4a Shell as high as or higher than wide ............ 5 by Shelliwiderthan high 2 eee ee 7 5a Shell with very prominent radial plicae........ 6 b Radial plicae weak or absent, spiral lirae SUPON Qs ois eyelid ce eee te ee eee O. savait 6a Radial plicae very prominent, spiral lirae evenly distributed on whorls, ventral surface with radial plicae; females with rectum and vagina bent downward, males with long-threaded [es o\ cian cts tecenadaitin tino aidclo 6. o'6 © bof. © O. plicatus b Radial plicae broad and low, spiral lirae concentrated on lower half of whorl, ventral surface smooth; females with straight anus and vagina, males with short-threaded penes O. gassiesi ~I » Radial plicae and spiral lirae of approximately equal prominence, making a reticulated SUTLAGE hs -ha oeet a eae ec Pree O. reticulatus b) Spirallirae predominant sec eee eee 8 SiauspineanelelessythamylO Os i vce ener ree 9 b) Spirelangle more thanvl\00 esr een teen 10 9a Ventral surface smooth; from Tutuila . .O. strigatus b Ventral surface with spiral sculpture; from OO Oe SANA setccccosccecne O. exasperatus 10a Shell smaller than 10.5 mm high by 13 mm WAGE! i. elute cgay SRR nV aw tstedeuenias ar O. llanero by Shellilangemthanyaboxeiy jee eee 1] 11 a Posterior part of hypobranchial gland thick and heavyeetcoms ©) pOlty seta ties iene O. tiara b Posterior part of hypobranchial gland much reduced; from Savall .............. O. garretti Vol. 20; No. 3 THE VELIGER Page 215 3. CRITERIA ror SPECIES RECOGNITION In order for 2 taxa to be recognized as separate species, they must display significant differences, not only in the shell, but also in the anatomy. The greater the number of differences in unrelated systems, and the greater the mag- nitude of the differences, the greater the probability that the 2 taxa really are different. To illustrate, let us consider 3 specific cases: one in which a few important differences made the species decision easy; one in which a number of smaller differences made the separation harder; and one in which the differences were deemed insufficient to warrant separation of taxa. The first case is that of Ostodes plicatus vs. O. gassiest. Both taxa have shells of approximately the same size and shape — higher than wide, and with narrow umbilici. Neither has any sculptural element that is totally lacking in the other; the conchological differences lie in the dis- tribution of the spiral lirae, and the distribution and rela- tive prominence of the radial plicae. The shell differences, though present and consistent, would not be sufficient ground for separating the 2 taxa, were it not for anatomi- cal differences of much greater magnitude. In O. plicatus, the males almost always have penes with long, bulbless threads, and the females almost always have a distinctive downward bend in the terminal portions of rectum and vagina. In O. gassiesi, the female openings point directly forward, and the males have short-threaded penes, often with sub-terminal bulbs. The relatively minor differences in sculpture, plus the major differences in genitalia, give ample evidence for the designation of these 2 taxa as separate species. The second case, that of Ostodes savati vs. O. exaspera- tus is less clear-cut. Individuals of O. exasperatus differ conchologically from those of O. savaii in height, width, umbilical diameter, and contour —- all elements which are influenced by the single factor of rate of decoiling — and in having more whorls for their size than does O. savau (Figure 12). They also differ in sculpture, in that radial plicae are totally lacking in O. exasperatus, while they are present, though unobtrusive, in O. savazi. In ad- dition, O. exasperatus shows stronger spiral lirae on its ventral surface than does O. savaii. There are also differ- ences in the anatomy: in the female, the proportions of the proximal and distal portions of the pallial common duct are different in the 2 taxa, and the male shows dif- ferences in proportions of penis thread to trunk, and in the slenderness or thickness of the penial trunk. Here are four areas of slight difference — shell shape, shell sculp- ture, male anatomy, and female anatomy — which to- gether are sufficient to warrant separation of the 2 species. diameter (mm) 14 5.00 5.25 5-50 5°75 6.00 whorls Figure 12 Relationship of whorl-count to diameter in Ostodes savait and Ostodes exasperatus A - Ostodes savai, male A - Ostodes exasperatus, male © - Ostodes savati, female @ — O. exasperatus, female In the third example, the two forms of Ostodes upolen- sis were not deemed different enough to warrant their separation. As in the preceding example the differing ap- pearances of the 2 types of shell are caused primarily by a single factor — the rate of decoiling. The size, the con- tour, and the width of the umbilicus are all determined in part by the rate of decoiling, and hence differences be- tween them must be considered as having only a single cause. The anatomical differences observed in the 2 forms are in size only — not in shape or location — and are so Page 216 slight as to be statistically insignificant. There are no sig- nificant differences in sculpture. The single important difference between the forms — the rate of decoiling — was not thought sufficient to warrant separation of the taxon into 2 species, as it was unaccompanied by any significant differences in sculpture or anatomy. 4. SPECIES DIAGNOSES AND DISTRIBUTIONAL DATA a. Ostodes plicatus (Gould, 1848) (Figures 13a, b; 14b; 15¢, d, e) Cyclostoma plicatum Goutp, 1848, Proc. Boston Soc. Nat. Hist., 2: 205 - Upolu; Gourp, 1852, U. S. Expl. Exped., 12: 103-104: Gouxp, 1860, Atlas of Shells, U. S. Expl. Exped., plt. 8, figs. 118, 118a, 118b. Cyclostoma apiae Récxuz, 1851, Journ. de Conch., 2: 213, 214; pit. 6, figs. 10, 11 - Apia, Upolu. Cyclophorus aptae (REcLUuz), Gray, 1852, Cat. Phan., pp. 57-58 — Pacific Islands, Upolu; Preirrer, 1852, Mon. Pneum., 1: 83 — Upolu. Cyclostoma pulverulentum PuitipPi, 1854, in PFEIFFER, Conch. Cab., 1 [19(1)]: 301 - 302, plt. 40, figs. 13, 14 - Upolu. Cyclophorus plicatus (Gould), REEve, 1862, Conch. Icon., XIII: sp. 58, plt. 14, fig. 58. Cyclophorus strigatus (Gould), Mousson, 1865, Journ. de Conch., 13: 179-180 - Upolu and Manua, Western Samoa — partly. Cyclophorus (Ostodes) strigatus (Gould), Mousson, 1869, Journ. de Conch., 17: 350-351, Upolu, Savaii, Tutu- ila — partly. Ostodes strigatus (Gould), Garrett, 1887, Proc. Acad. Nat. Sci. Philadelphia, 1887: 147-148 - Upolu, Tutuila, Savaii - partly. Ostodes plicatus (Gould), Kosett, 1902, Das Tierreich, 16: 156, fig. 32 (p. 153) - Samoa (Upolu); CLENcH, 1949, Bull. B. P. Bishop Mus., 196: 15-17 — partly. Diagnosis: Shell narrowly turbinate, height of adults 9.0-11.6mm {10.45mm}', diameter 8.8-10.6mm {9.8 mm} with 54-6 whorls {58}. Whorls with rounded shoulders, suture slightly incised. Usually (85%) higher than wide; H/D ratio 0.92—1.19 {1.06}. (Juveniles, es- pecially males, often wider than high.) Spire angle 70° — 100° {81°}. Umbilicus deep, narrow, margin either smoothly rounded or pinched into an acute angle, but usually without a bordering rim. D/U ratio 2.69-4.46 {3.37}. Aperture round, holostomatous, slightly appressed to whorl above. Parietal callus of adults approximately as * Data in braces { } represent the means of the preceding data THE VELIGER Vol. 20; No. 3 thick as outer lip. Early whorls smooth. Body whorl with 4—12 {7.1} fine, evenly spaced spiral threads on upper palatal surface, 6-14 {9.3} spiral threads on lower palatal surface, crossing 15-27 {21.0} broad, strong, close-set radial plicae on upper surface, of which 13-23 {17.4} continue across rounded ventral surface and enter umbili- cus. Color often salmon to rose pink on apex and spire, otherwise cream-color; body whorl nearly always cream- color. Periostracum thin, transparent, yellowish-brown, deciduous. Opercular types, A-1 and C-1. Hypobranchial gland with moderately heavy posterior (transverse) portion, beginning well over to left side; an- terior portion much reduced. Males have long-threaded penes without bulbs. Prostatic sac seldom present, “‘safety- valve’’ nearly always present; ‘‘valve” larger, sturdier, more obvious than in any other species. Female internal common duct very long (1.15 X diameter of bursa copu- latrix). Pallial common duct very short, with copulatory pore immediately below mantle line, and entry of com- mon duct into uterus immediately below pore, via slightly elevated oval papilla at 30° angle from long axis of uterus. Anus shows slight hypertrophy of upper margin. Vaginal orifice simply a round hole on the distal end of the uterus, equal in diameter to diameter of uterus. Terminal por- tions of both rectum and vagina bent sharply downward, at an angle approaching 90° in 80% of specimens seen. This bend is absolutely diagnostic of this species, as it occurs in no other. In the 20% without the bend, the anus points straight ahead, the uterus tapers to a point below the anus, and the vaginal orifice is a small hole on the ventral surface of the uterus. Comparative remarks: Differing emphasis and distribu- tion of sculptural elements on the body whorls are the chief conchological differences between Ostodes plicatus (Figure 13) and the most similar species, O. gassiest (CLENcu, 1949: fig. 7a). Some of the material used by Clench in his work on O. plicatus was lent to me and proved to contain examples of both O. plicatus and O. gassiest. In O. plicatus, the spiral threads are rather un- obtrusive and evenly distributed over the entire height of the body whorl; in O. gassiesi they are clustered on the lower half, and are stronger. The radial plicae are much stronger in O. plicatus; they are narrower and much more numerous, and continue across the lower surface of the shell, which is smooth, or marked only by very faint spirals in O. gassiesi. The strong radial plicae also serve to dif- ferentiate O. plicatus from O. strigatus, O. savait, O. Wanero, and O. exasperatus, in all of which the radials are very weak or absent. In addition, the latter 4 species are usually as wide as or wider than tall, whereas O. pli- catus is usually taller than wide. The sculpture of O. plicatus also differs considerably from the cross-hatched Vol. 20; No. 3 THE VELIGER Page 217 Figure 13 Ostodes plicatus a — Shell seen from side b — shell seen from below surface of O. reticulatus, which is a slightly smaller shell, wider than tall, with a much wider umbilicus. Ostodes plicatus is, of course, larger than O. upolensis, O. adjunc- tus, and O. cookei, and smaller than O. garretti and O. tiara. In anatomy, although the reduced anterior portion of the hypobranchial gland is fairly distinctive, Ostodes plt- catus is set off from other species of similar size chiefly by the reproductive systems. Ostodes plicatus males have longer penial threads, on the average, than do males of any species of comparable size except O. strigatus and O. reticulatus. In the female reproductive systems, the down- ward bend of the terminal portions of rectum and vagina is unique to O. plicatus; in the unusual specimen in which the downward bend is lacking, the individual could still be distinguished from O. reticulatus and O. strigatus by the proportionate lengths of the common ducts. The in- terior common duct in O. plicatus is slightly longer than the diameter of its bursa copulatrix (as is that of O. striga- tus), while that of O. reticulatus is less than one-half as long as its bursa is wide; the pallial common duct is very short in O. plicatus, but neazly 4 times the diameter of the copulatory pore in O. reticulatus, and 15 times the pore- diameter in O. strigatus. Ostodes plicatus and O. gassiesi approach each other very closely in conchological characters. An empty shell, especially a worn shell, might well be impossible to assign with certainty to either species. The anatomy, however, is quite distinctive, and there should be no difficulty in identifying any whole specimen. Range: Upolu, Western Samoa. Northeastern foothills, central uplands, and southern lowlands near coast. Alti- tude range 15m—750m. Material: Upolu: Station 2, 1.2km above Afiamalu seismographic station, at 720m elevation, on right side of road in dense forest (6 specimens, FANH 159173); Station 8, 1.2km above Afiamalu seismographic station, at 720m elevation, on both sides of road in mixed to good forest (7 specimens, FANH 170546, 159183, 159166); Station 9, Tafatafa, at 15 m elevation, in heavy forest (28 specimens, FMNH 152941, 159171): Station 18, at foot of Mount Solaua, at 180-240 m elevation, under Ficus tree in banana patch at edge of forest (59 specimens, FMNH 152894, 152842, 159163, 153107); Station 19, rim of Lake Lanuot’o crater, at 750m elevation (1 specimen, FMNH 152836); Station 39, top of range above Solaua, at 600m elevation, narrow ridge in heavy forest (6 specimens, FMNH 152588). Totals: 107 specimens: 33 adult males, 24 adult females, 5 juvenile males, 4 juvenile females, 5 accidentally broken, 17 not sexed, 19 empty. b. Ostodes gassiesi (Souverbie, 1858) (Figures la, b; 14a; 15a, b) Cyclostoma gassiesi Souverbie, 1858, Journ. de Conch., 7 294, plt. 8, figs. 6 a, 6 b. Locality unknown. Ostodes gassiesi (Souverbie), Kosett, 1902, Das Tierreich, 16: 155 — Polynesia. Page 218 Ostodes plicatus (Gould), CLencn, 1949, Bull. B. P. Bishop Mus., 196: 15-17, fig. 7 a — partly. Diagnosis: Shell narrowly turbinate, height of adults 9.2-14.6mm {11.3mm}, diameter 9.0-12.2mm {10.6 mm}, with 5$ to 64 whorls {5§}. Whorls with rounded shoulders, suture slightly incised, ventral surface rounded. Usually (87%) higher than wide; H/D ratio 0.91-1.31 {1.07}. Spire angle 70°-100° {80°}. Umbilicus deep, narrow, margin usually smoothly rounded and without bordering rim. D/U ratio 2.75—4.61 {3.42}. Aperture round, holostomatous, slightly appressed to whorl above, parietal callus of adults not quite as thick as outer lip of aperture. Early whorls usually smooth. Body whorl with 3-9 {5.6} strong, narrow, spiral threads clustered on lower half of upper palatal surface, crossing 14-23 {17.6} broad radial plicae. At maturity, spiral threads fade out completely, radial plicae become broader, flatter, further apart and less definite, underlying numerous fine radial growth lines. Lower palatal surface usually smooth: oc- casionally shows faint traces of spiral lirae without relief. Rarely shows pink on apex and spire; generally cream- color; periostracum thin, brown, adherent. Opercular types A-1, B, C-1. Juveniles differ from above in that they are usually wider than high (H/D ratio 0.79-1.07, mean 0.95), the body whorl has a sharply angled periphery, and the ven- tral surface is almost flat. Compare Figures 1a and 1b. Hypobranchial gland large and prominent posteriorly, beginning well over to left side. Anterior portion tapers smoothly and is quite short, ending opposite middle of kidney. Males have short-threaded penes, often with sub- terminal bulb. Prostatic sac present frequently, “‘safety- valve” seldom. If “safety-valve’’ is present, it does not cross over the vas deferens to terminate behind and below anus, as usual, but is either very short, terminating on top of the vas, or if longer, runs down the medial aspect of the vas and terminates about midway between the anterior end of the prostate and the base of the penis. Female in- ternal common duct short, only about ¢ as long as bursa copulatrix is wide. Pallial common duct also short; copu- latory pore down from mantle line only by its own diam- eter, with entry of common duct into uterus 1.5 pore diameters below pore. Common duct enters uterus via an elongate oval papilla, slightly elevated, its long axis at right angles to long axis of uterus. Anus a simple tube, pointing directly forward without hypertrophy of lips. Vaginal orifice a broad triangular slit, apex inward, its length 1.6 times the width of the uterus at the inner end of orifice. Comparative remarks: For the conchological differences between Ostodes gassiesi and the species most similar to THE VELIGER Vol. 20; No. 3 PTR STBP PTH 2mm l Figure 14 Males of (a) Ostodes gassiesi and (b) Ostodes plicatus [for explanation of abbreviations see Appendix on foldout] it, O. plicatus, see comparative remarks under the latter species. Ostodes gassiesi differs from O. savaii in being nar- rower at a higher whorl count as well as in proportion to Vol. 20; No. 3 THE VELIGER Page 219 DG Figure 15 Female reproductive system of Ostodes gassiesi and O. plicatus a — Ostodes gassiesi; b — O. gassiesi, entrance of common genital duct into uterus; c — O. plicatus; d - O. plicatus, ovary; e — O. plicatus, entrance of common genital duct into uterus [for explanation of abbreviations see Appendix on foldout] its height, and by its strong radial sculpture, which 1s lacking in O. savaii. Ostodes gassiesi is also taller and nar- rower than O. reticulatus, and taller than O. strigatus, with a more acutely angled spire and a much narrower umbilicus than either of the latter species. As mentioned under Ostodes plicatus, that species and O. gassiesi are quite distinct anatomically. Ostodes gassiesi is closer anatomically to O. savaii than to any other of the middle-sized species, but is still distinct. In the male, both species are short-threaded, but O. gassiesi hardly ever has a ‘‘safety-valve,”” while that structure is present in more than 4 of the males of O. savaii. In the female reproduc- tive system, both sections of the common duct are pro- portionately shorter in O. gassiesi than in O. savaii; the anal tip in O. gasszesi is a simple tube without hypertro- phy, whereas in O. savaii there is considerable hypertro- phy of the upper margin; and in O. gassiesi the upper lip of the vaginal orifice is longer than the lower lip, while the reverse is true in O. savaiv. Range: Upolu, southern and northeastern lowlands, western, central and northeastern foothills, central up- lands. Altitude range, 15m—750m. Savaii, eastern and southeastern lowlands, southern and southeastern foot- hills. Altitude range, 75m -—450m. Material: Upolu: Station 2, 1.2km above Afiamalu seismographic station, right of road, at 720m elevation, in disturbed upland forest (10 specimens, FMNH 152705); Station 5, 1.6km N of Tanumalala at 300m elevation, in a thinned logging area with some bananas (9 specimens, FMNH 152765); Station 6, 1.6km NW of Mt. Siga’ele, at 555m-570m elevation, in forest patch in gully-pasture area (8 specimens, FMNH 152927); Station 7, foot of SE peak Tafua-Upolu at 450m elevation, in forest part of transition zone above taro patch and newly cleared land (30 specimens, FMNH 152784, 152912); Station 8, 1.2km above Afiamalu seismographic station, a wide area on both sides of road, including Station 2, 720 m elevation in disturbed upland forest (2 speci- mens, FMNH 170545, 152690); Station 9, Tafatafa, at 15m elevation in heavy lowland forest (1 specimen, FMNH 152938); Station 10, Togitogiga, at 15m elevation, in heavy lowland forest (1 specimen, FMNH 170535); Station 13, summit of pass to Fagaloa Bay, at 225 m elevation under a large mango tree in mixed foothill forest (1 specimen, FMNH 152666): Station 14, lower slope of SE peak Tafua-Upolu, at 390m-420m elevation in disturbed to good foothill forest (9 specimens, FMNH 152753); Station 16, SE peak Tafua-Upolu at 480 m elevation, in undisturbed foothill forest (8 specimens, FANH 159175); Station 17, top ridge SE peak Tafua-Upolu at 600m elevation in disturbed foothill forest (2 specimens, FMNH 152774); Station 18, foot of Mt. Solaua, between 180 m- 240m elevation, under a large Ficus tree in a banana patch at edge of lowland forest (3 specimens, FMNH 170551, 166215); Station 19, rim of Lake Lanuto’o crater at 750m elevation, in heavy upland forest (18 specimens, FMNH_ 152809); Station 20, Lake Lanuto’o-Tapatapao trail, at 540 m elevation, in heavy foothill forest (41 specimens, FMNH 152795); Station 23, gully to foot of north side Mt. Siga’ele at 600 m — 645 m elevation in mixed to good foothill forest (8 specimens, FMNH 152606); Station 24, same gully as Station 23, but at 690m elevation, in good foothill forest (1 specimen, FMNH 152905); Station 26, Afiamalu-Lake Lanuto’o track at 735m-750m elevation, in good to excellent upland forest (36 specimens, FMNH 152670, 152719, 152721). Page 220 Savaii: Station 28, Vai'a'ata, near Vailoa, at 270m elevation, on new road in tall open lowland bush (31 specimens, FMNH 152572, 152649); Station 30, about 8km NW Vailoa at 180m elevation, in heavy lowland forest (10 speci- mens, FMNH 170539); Station 36, about 12.8 km NW Vailoa at 270m elevation in heavy foothill forest (8 specimens, FMNH 152634, 152642); Station 37, about 6.4km W of Gatavai in dry stream bed at about 450 m elevation for 9.6 km in heavy foothill forest (6 specimens, FMNH 170541); Station 38, 800m inland, 8km E of Vailoa on road to Salelologa wharf, at less than 75 m elevation (1 specimen, FMNH 152631). Totals: 244 specimens (188 from Upolu, 56 from Savaii); 86 adult females, 71 adult males, 19 juvenile females, 16 juvenile males, 28 not sexed, 24 empty. c. Ostodes reticulatus Girardi, spec. nov. (Figures 16a; 17a, b; 18a, b) Diagnosis: Shell turbinate, height of adults 8.0—10.8 mm {9.2mm}, diameter 9.1—11.2mm {10.0mm} with 54-58 whorls {5%}. Whorls with rounded shoulders, dropping to flat, vertical area below periphery. Usually (92%) wider than high: H/D ratio 0.82—1.04 {0.93}. Spire angle 90° - 100° {92°}. Umbilicus deep, very wide, often bordered by a rim formed by the innermost cord of spiral sculpture. D/U ratio 2.53 — 3.09 {2.78}. Aperture round, holostomatous, slightly appressed to whorl above. Parietal callus of adults usually not quite as thick as outer lip. Spire smooth; sculpture usually badly worn on early whorls of adults. Body whorl with 4-8 {4.5} evenly spaced raised spiral cords on upper palatal surface, 6-11 {7.6} spiral cords on lower palatal surface, crossing 19-29 {23.0} radial cords on upper surface, of which 18-24 {20.6} continue on ventral surface. Both radial and spiral cords are of approximately the same height and width, and their crossing gives a reticulated surface effect, leav- ing square or oblong hollows, wider and taller than the width of the cords, between them. Color creamy white. Periostracum thin, light amber-brown; wears off raised cords but remains in hollows between them, where it is covered by accumulated environmental debris. Opercular type A-1. Hypobranchial gland with reduced posterior portion; anterior portion long and dense, but quite slender. Males have long-threaded penes without bulbs. Both prostatic sac and ‘‘safety-valve” present. Female internal common duct moderately short — 0.43 times the diameter of the bursa copulatrix. Length of pallial common duct equal to approximately 4 times diameter of copulatory pore, with pore located slightly above mid-point of duct. Com- mon duct enters uterus via an elevated papilla, narrowly heart-shaped with apex up, at right angles to long axis of uterus. Anus shows slight hypertrophy of upper margin. Vaginal orifice an oval hole on ventral aspect of uterus, THE VELIGER Vol. 20; No. 3 its horizontal dimension being equal to ¢ the uterine diameter at posterior tip of orifice. Comparative remarks: Ostodes reticulatus resembles O. strigatus more closely than it does any other of the mid- sized species of Ostodes. (See comparative remarks under PTH PTR Figure 16 Males of (a) Ostodes reticulatus and (b) Ostodes strigatus [for explanation of abbreviations see Appendix on foldout] Vol. 20; No. 3 THE VELIGER Page 221 O. strigatus.) As regards the other species in the genus, the from O. savait. Ostodes savait lacks the radial threads that reticulated surface and the wide umbilicus are the most contribute to the reticulated surface of O. reticulatus, has notable conchological characters separating O. reticulatus a narrower umbilicus, and is in addition a somewhat larger shell, and one that tends to have almost equal vertical and horizontal measurements, instead of being definitely wider than high, as is O. reticulatus. Both O. gassiesi and O. plicatus are usually taller than they are wide, and have Figure 17 Female reproductive systems of Ostodes reticulatus and O. strigatus a — O. reticulatus;b — O. reticulatus, entrance of common genital Figure 18 duct into uterus; c - O. strigatus; d — O. strigatus, entrance ot common genital duct into uterus Ostodes reticulatus (holotype) [for explanation of abbreviations see Appendix on foldout] a — Shell seen from side b — shell seen from below Page 222 narrow umbilici. Ostodes reticulatus differs from O. exas- peratus and O. Ianero not only in its reticulated surface, but also in the shape of its whorls, which descend from the spire almost in a series of steps, whereas in the latter two species, the outline of the shell is a smooth, almost unin- terrupted, convex curve. Anatomically, Ostodes reticulatus seems more closely related to O. strigatus and O. plicatus than to the other middlesized species, but there should be no difficulty in differentiating among the three. For anatomical differ- ences between O. reticulatus and O. strigatus, see under O. strigatus. The hypobranchial gland of O. plicatus is heavy posteriorly and reduced anteriorly, while that of O. reticulatus is reduced posteriorly but quite prominent an- teriorly. Males of both species have similar penes (long threaded, without bulbs), but O. reticulatus is more apt to have a prostatic sac than is O. plicatus, and the ‘‘safety- valve” of the latter species is much more noticeable. In the female reproductive systems, O. reticulatus never has the downward bend of the terminal portions of rectum and vagina that characterizes O. plicatus, and the propor- tionate lengths of the common ducts are different, the internal duct being the longer in O. plicatus, the pallial duct being the longer in O. reticulatus. Description of holotype: An adult female specimen, height, 9.3 mm; diameter, 10.2 mm; spire angle, 90°; H/D ratio, 0.91. Whorls 5%, decoiling at even moderate rate until near aperature, when rate increases so last 5mm of body whorl are indented under periphery of penultimate whorl. Whorls with rounded shoulders but with narrow, flat, vertical area at lower margin of each, between lowest spiral thread and suture below. Nuclear whorl smooth; juvenile whorls somewhat worn. Upper palatal surface of body whorl with 25 radial cords, overlain by 4 spiral cords. Lower palatal surface with 19 radial and 6 spiral cords, of which the innermost forms a rim around the umbilicus. Intersecting radial and spiral cords produce a reticulated surface. Last 5 mm of body whorl with close- set radial lines of gerontic growth replacing major sculp- ture. Aperture round, holostomatous, diameter 3.9mm, slightly appressed to whorl above; parietal callus as thick as outer lip. Umbilicus wide, deep, bounded by rim; D/U ratio 2.76. Deepest part of umbilicus filled with environ- mental debris. Shell white, translucent; periostracum ex- tremely thin, pale amber color, very closely adherent; worn off raised portions of surface, present under organic debris in hollows. Operculum lost. Collected by A. Solem and L. Price, on November 4, 1965, at Station 16, SE peak Tafua-Upolu, in undisturbed foothill forest at 480m elevation, Upolu, W. Samoa. FMNH 170532. THE VELIGER Vol. 20; No. 3 Range: Upolu, W. Samoa, central foothills and uplands, northeastern foothills. Altitude range, 180-750 m. Material: Upolu: Station 5, 1.6km NW of Tanumalala, at 300m elevation, in thinned logging area with some bananas (2 specimens, FMNH 152897); Station 6, 1.6km NW Mt. Siga’ele, at 555-570 m elevation, in forest patch in gully-pasture area (2 specimens, FMNH 152928, 159186); Station 8, 1.2km above Afiamalu seismographic station, at 720m elevation, on both sides of road in mixed to good forest (2 specimens, FMNH 159167, 159182); Station 16, SE peak Tafua-Upolu, at 480m elevation, in undisturbed forest (7 speci- mens, FMNH 152757 (paratypes), 170532 (holotype)); Station 17, top ridge of SE peak Tafua-Upolu, at 600m elevation, in disturbed forest (1 specimen, FMNH 159181); Station 18, foot of Mt. Solaua, at 180-240 m elevation, under a Ficus tree in banana patch at edge of forest (2 specimens, FMNH 159180, 166213); Station 23, gully to foot of N side Mt. Siga’ele, at 600-645 m eleva- tion, in mixed to good forest (2 specimens, FMNH 159169); Station 24, N side Mt. Siga’ele, at 690 m elevation, in good forest (6 specimens, FMNH 159162); Station 26, Afiamalu-Lake Lanuto’o track at 735-750m elevation, in good to excellent forest (1 specimen, FMNH 159178). Total: 25 specimens; 7 adult males, 6 adult females, 5 juvenile males, 1 juvenile female, 3 not sexed, 3 empty. d. Ostodes strigatus (Gould, 1848) (Figures 16b, 17c¢, d) Cyclostoma strigatum Goutp, 1848, Proc. Boston Soc. Nat. Hist., 2: 204-205 - Upolu; Goutp, 1852, U. S. Expl. Exped., 12: 102-103; PFrEirFEr, 1853, Conch. Cab. 1 {19(1)]: 302-303, plt. 40, figs. 15, 16 — Upolu; Goutp, 1860, Atlas of Shells, U. S. Expl. Exped., plt. 8, figs. 117, 117a, 117b. Cyclophorus strigatus (Gould) Gray, 1852, Cat. Phan. p. 58 — Upolu; PFEIFFER, 1852, Mon. Pneum. I: 83-84 — Upolu; Reeve, 1862, Conch. Icon., 13: sp. 77, plt. 17, fig. 77; Mousson, 1865, Journ. de Conch., 13: 179-180 — Upolu and Manua — partly. Cyclostoma albida HOMBRON & JACQUINOT, 1852-1854, Voy- age au Pole Sud, 5 [4(2)]: 50-51, plt. 12, figs. 25-28 - Samoa. Cyclophorus (Ostodes) strigatus (Gould) Mousson, 1896, Journ. de Conch., 17: 350-351 - Upolu, Savaii, and Tutuila — partly. Ostodes albidus (Hombron and Jacquinot) KoseEtrT, 1902, Das Tierreich, 16: 153 — Samoa. Ostodes strigatus (Gould) Garrett, 1887, Proc. Acad. Nat. Sci. Philadelphia, 1887: 147-148 - Upolu, Tutuila, and Savaii — partly; Koper, 1902, Das Tierreich 16: 156 - Samoa (Upolu); CLEeNcH, 1949, Bull. B. P. Bishop Mus., 196: 13, figs. 4a, 4b — Tutuila: Laulii Valley at 30-150 m elevation; Fagatoga at 30-270 m elevation; foot of Mt. Tau at 24m elevation; Logatala Ridge at 60 m elevation; Leone at 45 m elevation; Leone-Aolaoa Trail at 270-360 m elevation: Fagasa-Maupasaga Trail at 150 m elevation. Diagnosis: Shell turbinate, height of adults 7.3—10.85 mm {9.32 mm}, diameter 8.6—11.6 mm {10.82 mm}, with 4§ to 54 whorls {53}. Sides of upper whorls somewhat flattened; body whorl with rounded shoulder. Suture in- Vol. 20; No. 3 cised on approximately first 2 whorls; on later whorls, lowest spiral lira tends to overlie suture up to the start of the whorl-inset which marks maturity. Usually (93%) wider than high: H/D ratio 0.73—1.10 {0.91}. Spire angle 75°—100° {86°}. Ventral surface rounded. Umbilicus wide, deep, bordered by a protruding rim which is seen to be hollow in juveniles but becomes solid in adults; D/U ratio of adults 2.38-3.52 {2.89}. Aperture circular to (occasionally) sub-ovate with vertical diameter the greater; holostomatous, gently appressed to whorl above. Parietal callus approximately as thick as outer lip. Early whorls smooth. Body whorl with 3—7 {4.8} strong spiral cords, with which, in 42% of the shells examined, are in- terspersed from one to three slender spiral threads. In 37% of the juvenile shells examined, but only 19% of the adults, the spiral threads are beaded in such a way as to give the appearance of radial plicae crossing the spirals. There may be from 19-51 {adult: 30.7} such beads on the spiral cords of the body whorl, but only very rarely is there any trace of true raised radial sculptural elements between the spiral cords. In those few shells with true radial sculpture, the radial elements extend only from the suture down through the highest one or two spiral cords; they seldom reach the whorl shoulder, and never extend onto the ventral surface. Ventral surface usually (86%) smooth in adults. Remaining adults and about half the juveniles examined showed from 8-12 {8.9} ventral spirals, ranging from moderately prominent threads to barely perceptible lines without any relief at all. Since many more juveniles than adults showed some ventral spiral sculpture, its absence in adults may be due to wear. The absence of radial ventral sculpture, however, is a true shell character, not due to wear. Only one shell of 153 examined showed even the faintest trace of any radial sculptural elements on the ventral surface. Color light tan to cream-color, with an occasional pinkish spire. Periostra- cum thin, amber-colored; wears off raised spiral cords, but sometimes remains in valleys between cords, where it be- comes covered by adherent environmental debris, giving the shell a spiral-striped appearance. Opercular types A-1, A-2, B, C-1, C-2, with half of the type B opercula (20% of the total number) being very concave (see p. 201). Hypobranchial gland faint, tenuous; begins slightly to left of midline, turns and runs forward only to a point opposite the middle of the kidney. Males have penis with very long thread without bulb. In one male of 43 exam- ined, penis was just to right of cephalic midline; penis on midline in all other specimens. Contrary to previous re- ports, all specimens seen had a closed vas deferens, not an open sperm groove. Vas deferens seems to be quite a long tube; it can be seen to be very convoluted as it runs beneath the integument from the prostate across to the THE VELIGER Page 223 penis. No ‘‘safety-valve’’ present, no prostatic sac; anterior end of prostate simply squared-off, without any constric- tion. Anterior medial corner makesasort of flat flap, under which vas deferens crosses to penial base. Length of female internal common duct approximately equal to diameter of bursa copulatrix. Pallial common duct long; copulatory pore just below mantle line; entry of duct into uterus down 15 pore diameters. Common duct enters uterus via simple hole, partly hidden beneath soft, transverse fold of tissue, and opening into a sort of gutter or channel, which runs forward a short distance before fading into surround- ing tissue. Anus with only very slight hypertrophy. Va- ginal orifice a broad triangular slit, apex pointing inward, length approximately 1.3 times width of uterus at inner end of orifice. Comparative remarks: Ostodes strigatus resembles O. reticulatus very closely. Although O. strigatus tends to have a lower whorl-count for its size than O. reticulatus, and a slightly more acutely-angled spire, shell sizes and proportions in the two species are almost identical. ‘There are, however, considerable differences in sculpture. The spiral cords in O. reticulatus are evenly spaced and of uni- form strength, and are crossed by radial cords of the same spacing and strength. Both spiral and radial elements con- tinue strongly on the ventral surface of the shell. In O. strigatus, the spiral cords tend to be unevenly spaced and interspersed with much weaker threads. True radial sculpture is seldom found on the upper shell surface, and never on the ventral surface, which is usually entirely smooth. The anatomical differences between Ostodes strigatus and O. reticulatus are much greater than the concholog- ical differences. The hypobranchial gland of O. strigatus is much smaller and fainter. In the female, both internal and pallial common ducts are longer in O. strigatus, and the vaginal orifice is differently shaped and much larger than it is in O. reticulatus. Males of both species have long-threaded, bulbless penes, but those of O. strigatus are longer (mean ratio, thread to trunk, 1/0.60 + 0.03, compared to 1/0.91 + 0.15) and a Student's ¢t Test of the two ratios yields a t of 3.065 with 46 degrees of freedom, which indicates a probability of less than 0.01 that O. strigatus and O. reticulatus are the same. Although Goutp (1848, 1852, 1860) clearly differen- tiates between Ostodes strigatus and O. plicatus, later au- thors, notably Mousson (1865, 1869) confuse the two. Ostodes plicatus is taller than it is wide, with radial sculp- ture predominant; O. strigatus is wider than it is tall, with almost exclusively spiral sculpture. Students’ t Tests com- paring male and female heights and diameters of O. stvri- gatus and O. plicatus indicate a probability of less than 0.001 that these two species are the same. Page 224 Similar ¢ tests of heights and diameters between Ostodes strigatus and the 3 other mid-sized species of Ostodes indi- cate that there is less than 0.01 probability of identity be- tween O. strigatus and O. gassiesi, and less than 0.001 probability of identity between O. strigatus and O. savaii; in addition, O. strigatus has fewer whorls and a wider umbilicus than either O. gassiesi or O. savaii. The prob- ability of identity between O. strigatus and O. exasperatus is approximately 0.015 as regards shell height and diam- eter, with O. strigatus a smaller shell with fewer whorls, that is much less likely to have spiral sculpture ventrally. Anatomically, both O. gassiesi and O. savaii usually have short-threaded penes, often with sub-terminal bulbs; O strigatus has a very long-threaded, bulbless penis. The chief anatomical difference between O. strigatus and O. exasperatus is in the female genitalia, with the copulatory pore of O. strigatus much closer to the mantle line, and the pallial common duct about 3 times as long as that of O. exasperatus. Comparison of juvenile shells of Ostodes strigatus with the available juveniles of O. llanero reveals that O. stri- gatus is taller, with a more acute spire angle and a nar- rower umbilicus. Ostodes strigatus is, of course, larger than O. upolensis, O. adjunctus, and O. cookei, and smaller than O. tiara and O. garrett. Range: Tutuila, American Samoa; western portion of island: center and south-east edge of central plateau, ex- treme southern coast, mountain slope near (just south of) Pago Pago. Altitude range, 60—390m. Materials: Tutuila. Station AS-6; central plateau above Aolaoufou, 2] km SW Pago Pago, 360-390 m elevation (74 specimens, FMNH 481051/N). Sta- tion AS-10; middle slopes NE side Matafao Peak, reservoir track, behind Pago Pago, in dense forest (33 specimens, FMNH 181083/N). Station AS-19; upper slopes Olotele Mt., edge of central plateau, at 270-330m elevation, SW of Pago Pago (26 specimens, FMNH 181151/N). Station AS-20; seaward slopes Fagatele Crater, 19 km SW Pago Pago, at 60 m elevation (20 specimens, FANH 181157/N). Totals: 153 specimens: 44 adult males, 40 adult females, 20 juvenile males, 17 juvenile females, 32 not sexed. e. Ostodes savaii Clench, 1949 (Figures 19a; 20a, b, c) Ostodes savai CLencu, 1949, Bull. B. P. Bishop Mus., 196: 14, 15, figs. 5, 6 — Savaii: Salailua, at 90-180 m eleva- tion; Matavanu, +900m; Siuvao-Auala (1.6-6.4 km inland) at 150-600 m elevation. Diagnosis: Shell turbinate, height of adults 7.6-13.6 mm {10.7mm}, diameter 8.2—13.0mm {11.0mm} with 5$—5% whorls {54}. Whorls with rounded shoulders, su- ture well defined. Tends to be approximately equal in THE VELIGER Vol. 20; No. 3 height and width, H/D ratio 0.79-1.19 {0.98}. Spire angle 70°-100° {86°}. Umbilicus deep, narrow; inner edge of final whorl often sharply angled at umbilical bor- der, but seldom having a rim. D/U ratio 2.32—4.40 {3.18}. Aperture round, holostomatous, slightly appressed to whorl above. Parietal callus not as thick as outer lip of aperture. Early whorls usually smooth. Body whorl with 4—12 {5.8} fine but strong spiral threads on upper palatal surface, crossing 11-26 {19.9} broad, rather low and poorly-defined radial plicae. Lower palatal surface with 4-17 {10.8} spiral threads or lines, usually not as prom- inent as those on upper surface, often just lines, without any relief. Radial plicae hardly indicated at all on lower surface. Color creamy; periostracum thin, brown, decid- uous. Opercular types A-1, B, C-1, C-2. Hypobranchial gland with quite prominent posterior portion: anterior portion reduced in width and thickness, but runs almost % of way from mantle line to anterior mantle margin. Males mostly (86.5%) with short-threaded penes; about half have sub-terminal bulbs. Most have pro- static sacs, but only about 4 have “‘safety-valves.” Female anterior common duct quite long, being slightly longer than bursa copulatrix is wide. Pallial common duct of moderate length, copulatory pore being anterior to mantle line by its own diameter, with entrance into uterus 4 pore-diameters below pore. Entrance of common duct into uterus via an elevated, round papilla. Anus shows considerable hypertrophy of upper margin, only slight hypertrophy of lower margin. Vaginal orifice a narrow triangular slit, apex in, its length equal to 2.1 X diameter of uterus at inner end of orifice. Comparative remarks: Ostodes savati resembles O. exas- peratus very closely; the differences and similarities be- tween these two species are discussed under comparative remarks for O. exasperatus. Ostodes savait differs from O. gassiest and O. plicatus in proportion, being wider at a lower whorl count as well as in proportion to its height. In addition, both O. gassiesi and O. plicatus have strong radial sculpture which is lacking in O. savaii. Ostodes Savaii is a larger shell, with more whorls and a narrower umbilicus, than O. strigatus. Ostodes reticulatus is a smaller shell than O. savati, and has a reticulated anges quite unlike that of the larger species. The anatomical differences between Ostodes savaii and O. exasperatus, O. strigatus and O. gassiesi are discussed under the latter 3 species. Range: Upolu, Western Samoa: northeastern, southern, and extreme southeastern lowlands; western, west-central, central and north-central foothills; central uplands. Alti- tude range, 3+750m. Savaii, Western Samoa: eastern, southeastern and southwestern lowlands; western, south- Vol. 20; No. 3 THE VELIGER Page 225 Figure 19 Males of Ostodes savaii, Ostodes llanero and Ostodes exasperatus a — O. savaii; b —- O. Ilanero; c — O. exasperatus, male from Upolu; d — O. exasperatus, type of penis found in Sava population [for explanation of abbreviations see Appendix on foldout] Page 226 THE VELIGER Vol. 20; No. 3 Figure 20 Female reproductive system of Ostodes savaii, Ostodes llanero and Ostodes exasperatus a — O. savaii; b - O. savaii, ovary; c - O. savaii, entrance of common genital duct into uterus; d - O. lanero; e — O. Ilanero, entrance of common genital duct into uterus; f — O. exasperatus; g — O. exasperatus, entrance of common genital duct into uterus, and position of copulatory pore in both populations [for explanation of abbreviations see Appendix on foldout] Explanation of Figures 43 to 48 Figure 43: Ostodes llanero, central, lateral, and inner marginal teeth X 736 Figure 44: Ostodes llanero, outer marginal teeth, showing fora- mina x 786 Figure 45: Gonatorhaphe sp., radula torn, showing base of central tooth and shank of lateral tooth X 235 Figure 46: Gonatorhaphe sp., radula partly closed, showing edge of membrane and foramina in outer marginal teeth X 344 Figure 47: Gassiesia sp., central tooth and tricuspid lateral teeth X 548 Figure 48: Gassiesia sp., radula partly closed, showing outer mar- ginal teeth X 235 Tue VE.icER, Vol. 20, No. 3 [Grrarpi] Figures 43 to 48 Vol. 20; No. 3 DiREMVEEIGER Page 227 ern and southeastern foothills. Altitude range, 150- 450m. Material: Upolu: Station 2, 1200 m above Afiamalu seismographic station, on right of road at 750m elevation in disturbed upland forest (5 specimens, FMNH 159174); Station 6, 1.6km NW of Mt. Siga’ele at 555 m elevation, in a forest patch in a gully-pasture area (1 specimen, FMNH 159187); Station 7, foot SE peak Tafua-Upolu at 360m elevation in forest part of transition zone above taro patch and newly cleared land (3 specimens, FMNH 159177); Sta- tion 8, 1.2km above Afiamalu seismographic station, in wide area on both sides of road, including Station 2, at 720 m elevation in mixed to good upland forest (12 specimens, FMNH 170538, 152875, 159168); Station 9, Tafatafa, in heavy lowland forest at 15m elevation (3 specimens, FMNH 170537, 170533, 159172); Station 10, Togitogiga, two logging roads toward ocean, at 15m elevation, in thinned lowland forest (1 specimen, FMNH 170543); Station 16, SE peak Tafua-Upolu at 480m elevation in undisturbed foothill forest (1 specimen, FMNH 159176); Station 18, foot of Mt. Solaua between 180-240 m elevation, under a large Ficus tree in banana patch at edge of forest (6 speci- mens, FMNH 153020, 159164, 159165, 166214); Station 20, Lake Lanuto’o Tapatapao trail, at 540 m elevation in heavy foothill forest (1 specimen, FANB 170534); Station 24, N side of Mt. Siga’ele at 690 m elevation, in good foothill forest (1 specimen, FMNH 170542); Station 25, cliffs at Tuiolemu at 3- 150m elevation in mixed to good lowland forest (8 specimens, FMNH 153108, 153432); Station 26, Afiamalu-L. Lanuto’o track at 735-750m elevation in good to excellent upland forest (9 specimens, FMNH 170544, 170547, 170536, 159178, 159179, 159184, 159185); Station 40, summit of Mt. Vaea, near tomb of Robert Louis Stevenson, at 420m elevation in much disturbed foothill forest (26 specimens, FMNH 152582). Savaii: Station 28, Vai’a’ata, near Vailoa, at 270m elevation, on a new road in tall open lowland bush (19 specimens, FMNH 170548, 170540); Station 30, about 8km NW of Vailoa at 180m elevation in heavy lowland forest (17 specimens, FMNH 153001, 153008); Station 31, 2km in from Salilua, 41.6km W of Vailoa, at 150 m elevation, in heavy lowland forest (38 specimens, FM{NH 152741, 152791); Station 32, up a survey track at 450 m elevation, approximately 8 km inland from Asau in open foothill forest (26 specimens, FMNH 152526); Station 36, about 12.8km NW of Vailoa, at 270m elevation, in heavy foothill forest (1 specimen, FMNH 170549); Station 37, about 1.6km W of Gatavai, at about 450m elevation, in dry stream bed for 10km in heavy foothill forest (10 specimens, FMNH 152560). Totals: 188 specimens (111 from Savaii, 77 from Upolu); 58 adult males, 52 adult females, 11 juvenile males, 16 juvenile females; 22 not sexed, 18 empty. f. Ostodes exasperatus Girardi, spec. nov. (Figures 19c, d; 20f, g; 21a, b) Diagnosis: Shell turbinate, height of adults 8.6—12.9 mm {10.4mm}, diameter 9.8—13.4mm {11.3 mm}, with 54—64 whorls {53}. Silhouette smoothly convex, suture not incised. Usually (88%) wider than high; H/D ratio 0.81—1.00 {0.92}. Spire angle 80°—105° {90°}. Umbili- cus wide, deep, usually (67%) bounded by a rim formed by innermost spiral cord, D/U ratio 2.36—3.85 {2.98}. Aperture sub-circular, slightly compressed vertically, very slightly appressed to whorl above. Parietal callus not as thick as outer lip. Apex and early whorls usually smooth. Body whorl with 4—7 {5.5} strong spiral cords on upper palatal surface, 6-20 {12.0} spiral cords on lower palatal surface. Only one shell of 24 adults exam- ined showed any trace of radial sculpture except fine growth lines. Color creamy white. Periostracum thin, brown, deciduous. Opercular types A-1, A-2, B, C-1, C-2. Hypobranchial gland narrow and elongate; runs nearly to front margin of mantle cavity. Males from Upolu have long-threaded penes without bulbs; those from Savaii mostly have short-threaded penes with bulbs. In both pop- ulations prostatic sacs and ‘‘safety-valves’” are unusual. Female internal common duct very long (approximately equal to the diameter of the bursa copulatrix). Pallial common duct also long; copulatory pore anterior to man- tle line by 3.5 times its own diameter. In females from Upolu, entrance of common duct into uterus is immedi- ately below copulatory pore; in Savaii females, the en- trance is approximately 2 pore diameters below the pore itself. This is the only difference in the female systems between the 2 populations. Entrance of common duct into uterus is via an elevated round papilla. Anus points di- rectly forward, shows considerable hypertrophy of upper margin, slight hypertrophy lower margin. Vaginal orifice a narrowly triangular slit, apex inward, on the medio- ventral aspect of the uterus. Length of vaginal orifice approximately 1.6 times uterine diameter at inner end of orifice. Comparative remarks: Ostodes exasperatus is very sim- ilar to both O. savaii and O. llanero. For the differences between O. exasperatus and O. llanero, see comparative remarks under the latter species. The principal concho- logical difference between O. exasperatus and O. savau is the contour of the shell. The spire of O. savai: descends with a progressively increasing rate of decoiling; although the early whorls present a fairly even contour, there is a definite “‘step’’ effect between the penultimate whorl and the body whorl, with an almost vertical drop from the periphery of the penultimate whorl to the suture below. In O. exasperatus, the rate of decoiling is slower and more even, with the periphery of each whorl slightly overlying the suture. In addition, O. exasperatus averages very slightly shorter and wider than O. savaiz, its umbilicus is slightly wider in proportion to the diameter of the shell, and it has a slightly higher whorl count for its size than does O. savaii. For differences between O. exasperatus and O. strigatus, see under the latter species. Anatomically, Ostodes exasperatus differs considerably from O. llanero, as discussed under that species. The dif- ferences between O. exasperatus and O. savazi are smaller, but consistent. In the female, the difference is in the length and proportions of the pallial common duct. In O. exasperatus, the portion of the pallial common duct above the copulatory pore is considerably longer than that below the pore; in O. savaz?, the reverse is true. In the male, O. savaii is much more apt to have a prostatic sac than is O. exasperatus, and slightly more apt to have a ‘‘safety-valve.’’ The penial trunk of O. savazi is slender Page 228 Figure. 21 Ostodes exasperatus (holotype) a — Shell seen from side b — shell seen from below in proportion to its length; that of O. exasperatus is thick and heavy. On Upolu, where the 2 species may be sym- patric, 86% of O. savaii males have short-threaded penes; THE VELIGER Vol. 20; No. 3 all O. exasperatus males are long-threaded. On Savaii, where the 2 species are definitely allopatric, they are both mostly short-threaded. Description of holotype: A young adult male, height 10.2mm, diameter 11.5mm, spire angle 85°, H/D ratio 0.88. Whorls 53, decoiling slowly until the last 4mm of body whorl, when rate increases so that terminal portion of whorl is slightly inset. Turbinate, with slightly in- dented suture, for first 2$ whorls. Thereafter suture not indented; periphery of each whorl over-lies whorl below very closely, producing a smoothly convex silhouette. First 23 whorls smooth; succeeding whorls progressively more sculptured. Body whorl with 6 strong, narrow, raised spiral cords on upper palatal surface, 15 somewhat less emphatic spiral cords on lower palatal surface. Innermost spiral cord of ventral surface forms rim of umbilicus, which is open to nuclear whorl. D/U ratio, 2.74. Major sculpture replaced by fine radial growth lines on last 4mm of body whorl. Aperture sub-circular, diameter 4.5mm, vertically compressed and slightly appressed to whorl above. Parietal callus thin. Creamy white with brown, deciduous periostracum. Operculum type C-2. Collected by L. Price on November 20, 1965 at Station 34, approximately 8 km SE of Asau along main road, then inland about 8 km along a track to about 540 m elevation, in light upland forest, Savaii, W. Samoa. FMNH 170530. Range: Savaiil, Western Samoa, northwestern uplands. Upolu, W. Samoa, locality unknown. Material: Upolu; Station unknown (12 specimens, FMNH 153820). Savaii: Station 34, approximately 8km SE of Asau along main road, then inland about 8km along a track to about 540 m elevation, in light upland forest (17 specimens, FMNH 170531). Totals: 29 specimens, (12 from Upolu, 17 from Savaii); 14 adult females, 2 juvenile females, 10 adult males, 3 juvenile males. Remarks: The 2 populations of this species, from the 2 islands, are not exactly alike. They are definitely closer to each other than either one is to anything else, therefore they are kept together. As has been mentioned above, the Upolu population of Ostodes exasperatus reverses the usual condition of sexual dimorphism in the genus by having males that are bigger than females; the Savaii pop- ulation conforms to the more normal state of having bigger females. g. Ostodes llanero Girardi, spec. nov. (Figures 22a, b; 19b; 20d, e) Diagnosis: Shell broadly turbinate; height of juveniles 6.6-6.9mm {6.7mm}, diameter 9.0—9.8mm {9.44mm} Vol. 20; No. 3 THE VELIGER Page 229 with 5—5¢ whorls. Whorls very slightly convex, almost flat-sided, suture covered by lower margin of whorl above. Body whorl with rounded shoulder. Always wider than high; H/D ratio 0.64-0.74 {0.70}. Spire angle 100°- 120° {110°}. Umbilicus very wide, deep, bounded by rim, D/U ratio 1.99-2.61 {2.37}. Aperture sub-circular, slightly compressed, appressed to whorl above, and with extension into hollow peri-umbilical rim. Parietal callus very thin. Apex smooth, early whorls usually worn; body whorl with 5—7 {6.0} close-set spiral lirae on upper pala- tal surface, 12-15 {13.3} finer spiral traces on lower pal- atal surface. These strong spirals crossed by numerous exceedingly fine radial threads. Shell thin, fragile, white, almost transparent; periostracum very thin, brown, decid- uous. Opercular type C-2. Hypobranchial gland not well developed in these ju- venile specimens. Transverse (posterior) portion begins only slightly to left of midline of mantle; anterior portion also very short. Male has long-threaded penis without bulb; no prostatic sac, no “‘safety-valve.”’ Female internal common duct moderately short, 0.47 times the diameter of the bursa copulatrix. Pallial common duct very short, with copulatory pore anterior to mantle line by approxi- mately twice its own diameter, and entry of common duct into uterus immediately below copulatory pore, via a ver- tical slit, slightly wider at top than at bottom, at right angles to long axis of uterus. No papilla, no hypertrophy of edges of slit. Anus shows very slight hypertrophy of both upper and lower edges. Vaginal orifice a very long slit in medial aspect of uterus. Length of slit 4—4.5 times diameter of uterus at inner end of slit. Comparative remarks: Ostodes llanero is a sibling spe- cies of O. exasperatus. The only differences between ju- venile shells of the 2 species (no adults of O. llanero were seen) are the spire angle, which averages 21° wider in O. Hanero, and the D/U ratio, which averages 0.57 smaller. (Table 6) Otherwise the shells are virtually identical. The Figure 22 anatomy, however, is quite different. Males of O. exasper- atus from Savaii tend to have short-threaded penes with Ostodes llanero (holotype) sub-terminal bulbs; the single male seen of O. llanero a — Shell seen from side b — shell seen from below had a long thread without a bulb. The greatest difference Table 6 Differences between Ostodes llanero and Ostodes exasperatus (juveniles, mixed sex) ——————S—S—=—=——==s Spire angle D/U ratio Range Range 5 2.52 - 3.30 2.94 3 1.99 - 2.61 2.37 Ostodes exasperatus Ostodes llanero 2 100° - 120° 110° Page 230 between the two species is in the female reproductive sys- tem. Each section of the common duct in O. exasperatus is approximately twice the length of the corresponding section in O. llanero, each measured in relation to its own bursa copulatrix and copulatory pore. The vaginal orifice in O. Uanero is 4—4.5 times the width of the uterus at the inner end of the orifice, whereas in O. exasperatus, the vaginal orifice is only 1.6 times the uterine width. Also, in O. llanero, the upper lip of the vagina extends farther forward than does the lower lip, while in O. exasperatus, the lower lip is the longer of the two. Although its smoothly convex outline, smaller size, and lack of strong radial sculpture set O. llanero apart from the other mid-sized species of Ostodes, the most striking difference is the very wide umbilicus. Although there is some overlap of ranges, at 2.37, the mean D/U ratio of O. Ilanero is considerably lower than that of any other mid-sized species. (Figure 23) 5 4 D/U o , be 2 species IG 1 SiGe iE ip N 9 5 Be) oy 6 ya 8 Figure 23 Comparison of D/U ratios of mixed-sex juveniles of 7 mid-sized species of Ostodes E - O. exasperatus; G — O. garretti; L — O. llanero; P — O. plicatus; R — O. reticulatus; S — O. savaii; St — O. strigatus Vertical line — range of measurements; horizontal line -— mean; box — two standard errors on either side of mean THE VELIGER Vol. 20; No. 3 Description of holotype: A juvenile female, height 6.6 mm, diameter 9.78mm, very broadly turbinate, spire angle 120°, H/D ratio 0.67. Five whorls, decoiling grad- ually. Almost flat-sided; first 44 whorls show very slight rounding of upper portion, then slight concavity, then very sharp periphery which overlies upper portion of lower whorl; sharp periphery changes on body whorl to rounded shoulder at aperture. First 3 whorls smooth. Fourth whorl partly smooth. Upper palatal surface of body whorl with 7 shallow spiral lirae crossing a multitude of very fine radial lines. On lower palatal surface, 13 flat spiral lines, not at all raised, cross many very fine radial lines. Even where shell is badly worn, traces of the spiral lines remain. Sculpture continues unchanged up to aper- ture. Aperture sub-circular, slightly compressed verti- cally, slightly appressed to penultimate whorl, slight ex- tension at lower, inner quadrant. Parietal callus very thin. Umbilicus very wide, bounded by rim, open to nuclear whorl. D/U ratio 2.50. Thin, fragile, creamy white, with few remnants of thin, brown, deciduous periostracum. Operculum type C-2. Collected by L. Price on November 20, 1965, at Station 34, approximately 8 km SE of Asau along main road, then inland about 8km along a track to about 540m, in light upland forest; Savaii, W. Samoa. FMNH 152991. Range: Savaii, W. Samoa: northwest foothills and up- lands. Altitude range 540-600 m. Material: Station 33, up a survey track at 600m elevation, approximately 8km inland from Asau in heavy primary foothill forest (2 specimens, FMNH 152997); Station 34, approximately 8km SE of Asau along main road, then inland about 8km along a track to 540m elevation in light upland forest (1 specimen, (holotype) FMNH 152991). Total: 3 specimens; 2 juvenile females, 1 juvenile male. h. Ostodes upolensis (Mousson, 1865) (Figures 24a, b; 25a; 26a, b) Cyclophorus upolensis Mousson, 1865, Journ. de Conch., 13: 180, 181 — Upolu, Western Samoa. Cyclophorus (Ostodes) upolensis (Mousson), 1869, Journ. de Conch., 17: 352 — Upolu. Ostodes upolensis (Mousson), GARRETT, 1887, Proc. Acad. Nat. Sci. Philadelphia, 1887: 148 - Upolu; Kosext, 1902, Das Tierreich, 16: 157 - Samoa (Upolu); CLENCH, 1949, Bull. B. P. Bishop Mus., 196: 12, 13; figs. 3 c, 28 e — Upolu: Latuafara, at 63.8m elevation; Mt. Vaea; Lake Lanuto’o at 720m elevation; Maldolelei, at 450m elevation; Sinaele, at 420m elevation; Tiavi, at 570- 660 m elevation. Savaii: Salailua, at 300-600 m eleva- tion. Vol. 20; No. 3 Diagnosis: Shell broadly turbinate, small; height of adults 3.2-7.0mm {4.9mm}, diameter 2.8—8.6mm {6.2 mm}, with 44 to 54 whorls {4}. Some have whorls with rounded shoulders, suture incised; others have smoothly sloping shoulders, very sharp keels, suture not incised. Nearly always wider than high; H/D ratio 0.66-1.14 {0.77}. Spire angle, 85°-110° {98°}. Umbilicus very wide, deep, nearly always sharply margined, but without a bordering rim; D/U ratio 1.62—3.77 {2.61}. Aperture round to subcircular, parietal callus not as thick as outer rim. Sculpture of strong spiral lirae and, in about half the adults seen, unobtrusive radial plicae. Body whorl with 5-8 {6.3} spiral lirae on upper palatal surface, crossing 15-25 {18.0} radial plicae; in about 4 of adults seen, lower palatal surface showed 2-10 {6.0} spiral lirae only, often faint. Cream color, with thin, brown, deciduous periostracum. Opercular type, C-1. Hypobranchial gland begins at left posterior margin of mantle cavity, runs across to slightly past mid-line, turns and runs forward only a short distance. Both anterior and posterior portions of moderate thickness. Male penis usu- ally long-threaded, without sub-terminal bulb. Usually has both prostatic sac and “‘safety-valve.”’ Female internal common duct very short, only %o as long as bursa copula- trix is wide. Pallial common duct quite long; copulatory pore down from mantle line 3.5 pore diameters, distance from pore to entry into uterus equal to 6.5 X diameter of copulatory pore. Duct enters uterus via an L-shaped slit, the stem of the L at right-angles to long axis of uterus, the leg of the L pointing forward and having hypertrophied edges. Anus shows considerable hypertrophy of lips, es- pecially on the upper margin. Vaginal orifice a slit on the medio-ventral aspect of uterus, its length approximately equal to uterine width at inner edge of orifice. Comparative remarks: Ostodes wpolensis is one of 3 small species in this genus. It occurs in 2 forms, the ordi- nary, or “stepped” form (Figure 24a) illustrated by CrLencu (1949: 11; fig. 3c), and a ‘‘smooth” form (Figure 24b). The “smooth” form was found only on Upolu, with juveniles at Station 2, and both adults and juveniles at Station 18. No “‘stepped” shells were found at those sta- uons. Shells of the “smooth’’ form are slightly smaller than those of the ‘‘stepped”’ form and have a narrower umbilicus. Although males of both forms have long- threaded penes, those of the “smooth” form are very slightly longer, and the vaginal orifice of the females is very slightly smaller. The difference in shape of the 2 forms is caused by a difference in rate of decoiling. The rate is quite slow in the “smooth” form, with each whorl placed well up on the preceding one, and the peripheral carina is extended over the suture, thus producing a THE VELIGER Page 231 Figure 24 Ostodes upolensis shell outline: a — “stepped form -- b — “smooth” form smoothly conical silhouette. In the ‘‘stepped’’ form, the rate of decoiling is much faster, each whorl being placed further down the whorl preceding it, and the peripheral carina being almost vertically below the next higher spiral lira, so that the shoulders are rounded. The narrower um- bilicus in the smooth form is caused by a very pronounced insetting of the terminal portion of the body whorl under the penultimate whorl (see p. 196). Neither the difference in shape nor the very slight anatomical differences are considered sufficient to warrant separation of the 2 forms. Page 232 THE VELIGER Vol. 20; No. 3 Neither form of Ostodes upolensis is likely to be con- be confused with O. adjunctus, from Tutuila. Clench’s fused with another of the small species, O. cookei. Ostodes figure of O. adjunctus (CLENCH, 1949: 11, fig. 3b) shows upolensis is quite strongly sculptured, whereas O. cookei a shell midway in shape between the two forms of O, is the only species in the genus to present a smooth shell surface. The smooth form of O. upolensis, however, might Sooo Figure 26 Figure 25 Female reproductive systems of Ostodes upolensis and O. adjunctus . a — O. upolensis; b — O. upolensis, entrance of common genital Males of (a) Ostodes upolensis and (b) Ostodes adjunctus duct into uterus; c - O. adjunctus [for explanation of abbreviations see Appendix on foldout] [for explanation of abbreviations see Appendix on foldout] Vol. 20; No. 3 THE VELIGER Page 233 upolensis. Mousson’s original figure, however (Mousson, 1869: plt. 14, fig. 9) shows a very smoothly conical shell. The 40 shells of O. adjunctus available to me are also smoothly conical. Comparison of the “smooth” form of O. upolensis with the shells of O. adjunctus immediately shows the differences between the two. At the same or a lower whorl count, O. adjunctus is larger than O. upolen- sis (Figure 27). The umbilicus of O. adjunctus always has height (mm) 7 diameter (mm) 9 whorls whorls a b Figure 27 Relationship of whorl-count to size in Ostodes adjunctus and O. upolensis (both forms together) a — height/whorls; b — diameter/whorls A - Ostodes adjunctus, male @ - O. adjunctus, female A - Ostodes upolensis, male © - O. upolensis, female a bordering rim, whereas that of O. upolensis is rimless. Ostodes adjunctus has very strong spiral sculpture on the lower palatal surface, and is entirely lacking in radial sculpture, while in O. upolensis ventral sculpture is faint or absent and radials, though unobtrusive, are often pres- ent. Ostodes adjunctus is the only species of Ostodes in which shells that were otherwise clearly adult did not have the terminal portion of the body whorl indented beneath the penultimate whorl; in O. upolensis, especially in the smooth form, the indentation is very pronounced. Finally, fully half the shells of O. adjwnctus examined had a bright red spire, a character never seen in O. upolensis. Anatomically, the hypobranchial gland of Ostodes upolensis is much larger, especially the posterior portion, than that of O. adjunctus. Males of O. adjunctus entirely lack the prostatic sac and ‘“‘safety-valve” so often present in O. upolensis. In the female, the pallial portion of the common genital duct is only half as long in O. upolensis as it is in O. adjunctus, and the vaginal orifice is only 3 as long. Range: Upolu, Western Samoa: southern and north- eastern lowlands, central foothills and uplands. Altitude range, 15—750m. Savaii, Western Samoa: southwestern lowlands at 150m elevation. Material: Savaii: Station 31, 2.2km in from Salilua, 42km west of Vailoa, at 150m elevation, in heavy lowland forest (1 specimen, FMNH 152621); Upolu: Station 2, 1.2km above Afiamalu seismographic station, on right of road, at 720m elevation, in disturbed upland forest (4 specimens, FMNH 152710); Station 6, 1.6km N of Mt. Siga’ele, at 555-570m elevation, in a forest patch in a gully-pasture area (1 specimen, FMNH 152924); Station 8, 1.2km above Afiamalu seismographic station, a wide area, including Station 2, on both sides of the road, at 720m elevation, in mixed to good upland forest (5 specimens, FMNH 152686, 166212); Station 10, Togitogiga, two log- ging roads toward ocean at 15m elevation in thinned lowland forest (3 speci- mens, FMNH 152953); Station 18, foot of Mt. Solaua, between 180-240m elevation, under a large Ficus tree in a banana patch at edge of forest (45 specimens, FMNH 152850, 153026, 153093, 153104, 153169); Station 23, gully to foot of N side of Mt. Siga’ele, at 600-645 m elevation, in mixed to good foothill forest (18 specimens, FMNH 152605, 159170); Station 26, Afiamalu- Lake Lanuto’o track, at 735-750m elevation, in good to excellent upland forest (1 specimen, FMNH 166211). Totals: 77 specimens; 11 adult females, 11 adult males, 21 juvenile females, 24 juvenile males, 8 not sexed, 2 empty. 1. Ostodes adjunctus (Mousson, 1869) (Figures 25b, 26c) Cyclophorus (Ostodes) adjunctus Mousson, 1869, Journ. de Conch., 17: 351-352; plt. 14, fig. 9 - Tutuila. Ostodes adjunctus (Mousson) GarreETT, 1887, Proc. Acad. Nat. Sci. Philadelphia, 1887: 148 - Tutuila; Kosett, . 1902, Das Tierreich 16: 153 - Samoa (Tutuila); CLENcH, 1949, Bull. B. P. Bishop Mus., 196: 11-12; fig. Page 234 3b -— Tutuila: Fagasa and nearby valley; Pago Pago at 60-150 m elevation; Amalau Bay, at 30m elevation; NW slope on Mt. Pioa, at 180-240 m elevation; trail between Olofau and Amouli, at 90-120m elevation; Aua-Afono Trail, at 300 m elevation; Amouli, half-way between Pago Pago and Alofau. Diagnosis: Shell turbinate; height of adults 4.25—6.5 mm {5.25mm}, diameter 5.75—8.5mm {7.07 mm} with 4—4% whorls {44}. Upper whorls have rounded shoulder, body whorl has angled shoulder, often with moderate keel. Suture incised. Only 4 adults of 17 examined showed any trace of indentation of terminal portion of body whorl, although the sculptural change marking the onset of ma- turity was very clear. Always wider than high; H/D ratio 0.67 — 0.83 {0.75}. Spire angle 80° — 110° {92.5°}. Ventral surface flattened. Umbilicus wide and deep, margined with upstanding rim made by innermost ventral spiral lira; D/U ratio 2.50 —3.89 {3.02}. Aperture almost square in juveniles to round in adults, appressed to whorl above. Parietal wall usually not as thick as outer lip. Early whorls smooth; body whorl with 5 —6 strong spiral lirae on upper surface, 6-8 {6.5} spirals on ventral surface. Ventral spi- rals nearest umbilicus show the most relief; outer spirals less raised, may be just lines without relief; this may be due to wear. Shell has no radial sculptural elements of any kind except fine, close-set radial growth lines, which dominate shell surface at onset of maturity, when spiral lirae stop very abruptly. Color cream to dirty white except that slightly more than half the specimens seen had intense raspberry red spires for first 1-34 whorls {22}. Periostra- cum deciduous, pale yellow-brown, wears off raised spiral lirae quickly. Operculum types B, C-l. Hypobranchial gland very small, extending only short distance to left of midline, and running forward hardly at all. Of 11 males examined, 2 had the penis to the right of the cephalic midline; the other 9 had the penis on the cephalic mid-line. Penis of some specimens seems unusu- ally large in proportion to head. Penis usually long- threaded; no sub-terminal bulb, no prostatic sac, no “safety-valve.” Prostate squared-off at anterior end, medial corner forming a flap under which vas deferens crosses to penial base. Vas deferens a short, straight tube, not long and convoluted. Female internal common duct very short, only 0.2 times diameter of bursa copulatrix. Pallial com- mon duct quite long: copulatory pore down 2 pore diam- eters from mantle line, entrance of duct into uterus 16 pore-diameters below pore. Interior aspect of entrance of genital duct into uterus not seen. Anus points straight forward, with considerable hypertrophy all around edges. Vaginal orifice a long, narrow slit on medio-ventral aspect of uterus, its length 1.4 times diameter of uterus at inner end of orifice. THE VELIGER Vol. 20; No. 3 Comparative remarks: Ostodes adjunctus resembles O. upolensis more closely than it does any other species of Ostodes; see comparative remarks under O. upolensis. The sculptured shells of O. adjunctus could not possibly be confused with the smooth shells of O. cookei, and of course all the other species of Ostodes are too large to be confused with O. adjunctus. Range: Tutuila, American Samoa: eastern portion of island: south-east coast and ridge of central mountains. Altitude range, 60-330 m. Material: Station AS-1; steep hillside, banks of Visa Stream, 11km E of Pago Pago at 60m elevation (6 specimens, FMNH 181005/6). Station AS-2; steep ridge slope, at 90m elevation, Siliataligalu Point, 11.7km E of Pago Pago (27 specimens, FMNH 181011/N). Station AS-7; main ridge above Fagaitua, 18.4km E of Pago Pago, at 210-240m elevation (4 specimens, FMNH 181059/4). Station AS-9; crest of main ridge, left side of Aua-Afono track, 4.8km E of Pago Pago, at 300- 330m elevation (4 specimens, FMNH 181078/4). Totals: 40 specimens: 11 adult males, 5 adult females, 8 juvenile males, 12 juvenile females, 4 empty. j. Ostodes cookei Clench, 1949 Ostodes cooket CLENcH, 1949, Bull. B. P. Bishop Mus., 196: 10-11; fig. 3 a - Upolu, Western Samoa: Tiavi, at 630 m elevation. Diagnosis: Adult shell, turbinate, small, height 5.6mm, diameter 7.5mm, with 4§ whorls. Whorls with rounded shoulders, suture incised. Last 4 of body whorl inset under penultimate whorl. Wider than high, H/D ratio 0.74. Spire angle 110°. Aperture round, holostomatous, parietal wall as thick as outer lip. Umbilicus very wide, deep, open to nuclear whorl, D/U ratio 2.34, bounded by a rim that protrudes over the umbilicus but is not raised from the lower palatal surface of the shell. Shell cream color with very thin yellowish iridescent periostracum which is peel- ing off in spots and is missing from most of the spire and the last $ of body whorl. Where periostracum is missing, shell shows pitting and wear. Shell appears entirely smooth to naked eye. Magnification of 16X reveals texture of many fine radial growth lines, crossed on upper palatal surface of body whorl only by 7 spiral lines, which are just lines, not raised at all from shell surface. Spiral lines stop and growth lines coarsen for last $ whorl. Operculum not seen. Anatomy not seen. Comparative remarks: Ostodes cookei approaches the other two small Ostodes, O. upolensis and O. adjunctus in size and general configuration, but is smooth while the other two species are strongly sculptured. Although O. tiara and O. garretti may produce a secondarily smooth Vol. 20; No. 3 surface due to wear, the difference in size makes confusion of either with O. cookei impossible. Range: Tiavi, Upolu, Western Samoa, at 630m eleva- tion. Material: Tiavi, Upolu, at 630m elevation (1 specimen, MCZ 140504, para- type). k. Ostodes tiara (Gould, 1848) (Figures 7a, b; 8a, b) Cyclostoma tiara, GouLp, 1848, Proc. Boston Soc. Nat. Hist., 2: 204 - Upolu, Western Samoa; Goutp, 1852, U. S. Explor. Exped., 12: 101; Goutp, 1860, U. S. Explor. Exped., Atlas of Shells, plt. 8, figs. 116, 116 a. Cyclophorus tiara (Gould), Gray, 1852, Cat. Phan., p. 58 - Upolu; PFEIFFER, 1852, Mon. Pneum., 1: 84 — Upolu; REEVE, 1862, Conch. Icon., XIII: sp. 76, plt. 16, fig. 76 — Upolu; Mousson, 1865, Journ. de Conch., 13: 179 - Upolu to 1,000 m elevation. Cyclophorus (Ostodes) tiara (Gould), Mousson, 1869, Journ. de Conch., 17: 350 - Upolu. Ostodes tiara (Gould), Garretr, 1887, Proc. Acad. Nat. Sci. Philadelphia, 1887: 146-147 - Upolu; KoseEtt, 1902, Das Tierreich, 16: 156, 157; CLeENcH, 1949, Bull. B. P. Bishop Mus., 196: 17, 18; fig. 7b - Upolu, near Tiavi, at 630 m elevation. Diagnosis: Shell broadly turbinate, very large; height of adults 12.5-16.2mm {14.3mm}, diameter 16.0—22.5 mm {18.2mm}, with 4§—5§ whorls {4%}. Whorls moder- ately carinate, suture not incised. Always wider than high; H/D ratio 0.71—0.86 {0.79}. Spire angle 100°-110° {107°}. Umbilicus usually wide, deep, margin smoothly rounded without bordering rim. D/U ratio 2.58-4.13 {3.37}. Aperture round, holostomatous, slightly ap- pressed. to whorl above, parietal callus of adults almost as thick as outer rim of aperture. Sculpture usually worn off early whorls. Body whorl with 9-20 {15.4} spiral lirae on upper palatal surface, 5-11 {8.0} spiral cords on lower palatal surface. On juvenile shells, 14-23 {16.8} radial threads cross the spiral lirae on the upper palatal surface; radial threads may persist in a few adults, but most adults show no radial sculpture except fine growth lines. Color whitish. Periostracum thin, brown, adherent. Opercular types A-2 and C-3. Hypobranchial gland very thick and heavy posteriorly; anterior portion narrow but dense nearly to front of man- tle cavity. Males have two different types of penis, as dis- cussed under anatomy of reproductive system of genus. Of 9 adult males measured, 6 had “‘smooth” penes — long threaded, without bulb (Figure 7a) while 2 had “bumpy” THE VELIGER Page 235 23, diameter (mm) 21 13 4.50 5-00 5.50 6.00 whorls Figure 28 Relationship of whorl-count to diameter in Ostodes tiara and Ostodes garretti A — Ostodes tiara, male A - Ostodes garretti, male © - Ostodes tiara, female @ — Ostodes garretti, female penes — short threaded, with “knuckle,” “collar,” and bulb (Figure 7b). (Of juvenile males examined, 4 had “smooth’’ penes, while 3 had “bumpy” penes.) Both types of penis can be associated with a “‘safety-valve,” but a pro- static sac is rare in an animal with a ‘‘smooth”’ penis. Fe- male internal common duct short, its length equal to only half the diameter of the bursa copulatrix. Pallial common duct very long; copulatory pore anterior to mantle line by twice its own diameter, and distal portion of duct equal to 26 times the diameter of the copulatory pore. Unlike all other species, Ostodes tiara has the entrance of the common duct into the uterus on the posterior (ventral) surface of that organ, near the lateral margin (all other species have the entrance on the medial margin of the Page 236 uterus). Several attempts to expose the interior papilla of the common duct for drawing failed, and its shape and orientation remain unknown. Anus shows considerable hypertrophy of edges all around. Vaginal orifice a rela- tively small triangular slit, apex inward, its length only 4 the diameter of the uterus at inner end of orifice. Comparative remarks: Ostodes tiara is more similar to O. garrett: than to any other species. For similarities and differences between the two, see comparative remarks under O. garrettt. Range: Upolu, Western Samoa: western, west-central, and north-eastern foothills; central uplands. Altitude range, 180—750m. Material: Upolu: Station 1, 1.6km above Afiamalu seismographic station, to left of road, at 720 m elevation, in heavy upland forest to edge of sago-palm swamp (1 specimen, FMNH 152804); Station 2, 1.2km above Afiamalu seismo- graphic station, to right of road, at 720m elevation, in disturbed upland forest (4 specimens, FMNH 152700, 153088); Station 8, 1.2km above Afiamalu seismographic station, wide area on both sides of road, including Station 2, at 720m elevation, in mixed to good upland forest (19 specimens, FMNH 152663, 152877); Station 16, SE peak Tafua-Upolu, at 480m elevation, in un- disturbed foothill forest (1 specimen, FMNH 152760); Station 18, foot of Mt. Solaua, between 180-240m elevation, under a large Ficus tree in banana patch at edge of forest (1 specimen, FMNH 153175); Station 19, rim of Lake Lanuto’o crater, at 750m elevation, in heavy upland forest (4 specimens, FMNH 152812, 152835); Station 26, Afiamalu-Lake Lanuto’o track, at 735— 750m elevation, in good to excellent upland forest (11 specimens, FANH 152672, 152676, 152718, 166210); no locality given (1 specimen, FMNH 153819). Totals: 42 specimens; 9 adult males, 10 adult females, 7 juvenile males, 5 juvenile females, 5 not sexed, 6 empty. 1. Ostodes garretti Clench, 1949 (Figures 7c; 8c, d) Ostodes garrettt CLENCH, 1949, Bull. B. P. Bishop Mus., 196: 18; fig. 7 c — Savaii: Siavao-Auola, at 150-600 m ele- vation; Salailua, from sea level to + [sic] 780 m. Diagnosis: Shell broadly turbinate, very large; height of adults 10.8-14.8mm {12.5mm}, diameter 13.2—19.2 mm {16.3mm)} with 5—54 whorls {54}. Whorls carinate, suture only slightly incised. Always wider than high; H/D ratio 0.69—0.82 {0.78}. Spire angle 100°-115° {108°}. Umbilicus deep, wide; sometimes smoothly rounded, sometimes with bordering rim. D/U ratio 2.9—3.5 {3.19}. Aperture round, holostomatous, slightly appressed to whorl above, parietal callus of adults as thick as outer rim of aperture. Sculpture worn off early whorls. Body whorl with 7—15 {10.0} raised spiral cords on upper palatal sur- face, 11-13 {12.0} less prominent spiral cords on lower palatal surface. No radial sculpture except fine lines of THE VELIGER Vol. 20; No. 3 growth. Color whitish, periostracum very thin, deciduous, brownish-yellow. Opercular types A-1, A-2, C-2. Hypobranchial gland surprisingly small for so large a snail. Posterior portion very short, extending hardly at a:i to left of midline. Anterior portion runs well forward but is very narrow. Male penis with long thread without sub-terminal bulb; prostatic sac present; “‘safety-valve” may or may not be present. Female internal common duct very short, its length equal to only 4 diameter of bursa copulatrix. Copulatory pore very close to mantle-line, but distal portion of pallial common duct very long — eight times diameter of copulatory pore. Entry of common duct into uterus is via a round elevated papilla inside medial aspect of uterus. Anus a simple tube without hypertrophy of lips, pointing forward. Vaginal orifice a long triangular slit, apex inward, its length equal to 2.6 times diameter of uterus at inner end of orifice. Comparative remarks: Ostodes garretti is closely related to O. tiara. Ostodes garrett: averages slightly smaller both in absolute measurement and in diameter for a given whorl count (Figure 28). Its umbilicus is very slightly wider in proportion to the diameter of the shell. The anatomical differences are considerably greater than those of the shells. Ostodes garretti has a much smaller hypo- branchial gland. Its males have long-threaded penes with- out bulbs, whereas O. tiara males either have long- threaded penes without bulbs, or short-threaded, “bumpy” penes with bulbs. The greatest differences be- tween the 2 species are found in the female reproductive system. Although both distal pallial common ducts are long, that of O. tiara is proportionately more than 3 times as long as that of O. garrett:, and terminates on the ventral aspect of the uterus near the lateral margin, while O. garretti’s common duct enters the uterus in the usual way on the medial margin. Although O. garretts females have far less anal hypertrophy than do females of O. tiara, their vaginal orifices are proportionately more than 3 times as _ large as those of the latter species. Range: Savaii, Western Samoa; northwest uplands. Material: Station 34, approximately 8 km SE of Asau along main road, then inland about 8km along a track to 540m elevation in light upland fores¢ (8 specimens, FANH 152986). Totals: 8 specimens; 2 adult males, 2 adult females, 4 empty shells. Remarks: Ostodes garretti has a more pronounced sex- ual dimorphism in size than any other member of the genus. At almost exactly the same whorl count, males averaged 21% shorter and 29% narrower than females. Vol. 20; No. 3 C. The Generic Affinities of Ostodes Tielecke’s Poteriidae includes all of the helicoid cyclo- phorid genera from Central America, South America, the West Indies, and the South Pacific. Of the Pacific genera, Ostodes is the only Polynesian taxon, and it is restricted to Western and American Samoa (CLENCH, 1949: 3, 4). Names and distributions of the other nominate Pacific genera are: Gassiesia Clench, 1949 from New Caledonia and the Loyalty Islands; Dublonia Clench, 1949, Paramia Clench, 1949, and Kondorhaphe Clench, 1949 from the Caroline Islands; Gonatorhaphe MOllendorff, 1898 from the New Hebrides and Fiji; and Fijiopoma Clench, 1949 from Fiji. All of these genera were established on the basis of shell and opercular differences. Except for a few radular notes by CLENCH (1949: 48) and data in SoLtem (1959: 182-185; plt. 6, figs. 9, 10) on Gonatorhaphe genitalia, no anatomical information has been recorded. During this study, radulae from Gassiesia and Gonatorhaphe were examined with the aid of a scanning electron microscope (pp. 201-203), and their shells, anatomy, and opercula were examined rather briefly, as presented below. I have not seen any anatomical material from the Caroline Islands or from Fiji. The Neotropical genera placed by THompson (1969) in the subfamilies Neocyclotinae and Crocidopominae are included by Tielecke in the Poteriidae. Examples of a representative species from that group, Mexcyclotus pan- amensis (Da Costa, 1903), were dissected so that their re- productive systems could be compared with those of Os- todes. The results of those dissections are also presented below, along with a brief discussion of the shell and oper- culum. The radula of Mexcyclotus was not seen. Specimens of too few genera have been dissected to permit any statements concerning phylogeny within the Poteriidae. 1. Gonatorhaphe spp. Shell broadly turbinate, wider than high. Dorsal surface shows fine, close-set spiral lirae; some individuals also show broad, rounded, radial plicae. The ventral surface is smooth in the specimens from Espiritu Santo, and there is usually no umbilical rim. The spiral lirae are retained on the ventral surface of the specimens from Aoba, and the umbilicus has a rim. Aperture round, lip simple. Oper- culum with 2 layers. Lower layer smooth, shiny on under (attachment) side, thick centrally, diminishing to the van- ishing point toward the edges. Upper layer of thin, trans- parent corneous material with an iridescent sheen for first THE VELIGER Page 237 1.5—2.5 volutions, then becomes much thicker and heav- ily impregnated with calcium, the volutions lying imme- diately next to each other, their upper surfaces all on same plane. Result is a smooth-surfaced, thick, polygyrous, cal- cified operculum, with a small, non-calcified, depressed central area. Radula similar to that of Ostodes. Male reproductive system: ‘Testis occupies entire vol- ume of approximately first whorl; similar to that of Os- todes — branched digitiform alveoli, arranged in a single plane, feed into one collecting duct on the columellar side. Collecting duct runs forward as seminal vesicle. Prostatic lumen not seen. Anterior tip of prostate truncated, with- out sac or ‘‘safety-valve.”’ Vas deferens a closed tube. Penis quite variable. In specimens from Espiritu Santo, penis is standard short-thread type, without sub-terminal bulb; approximate thread-to-trunk ratio, 1 /2.8. In speci- mens from Aoba, thread is shortened into a terminal ap- pendage, triangular in cross-section and with sharp edges, usually found folded back acutely against trunk. Mean ratio, appendage to trunk, 1/3.8. Two specimens were seen in which the appendage was longer, with less acute edges, and looked more like a true thread; both had ratios of appendage to trunk of 1/2. One individual was seen which had both a closed vas deferens and an open groove on the posterior side of the penis. The groove did not run from prostate to penis, but began above penial base, ran up posterior side of penis, and ended just before distal end of penis folded over to become terminal appendage. The groove ran in a straight line, with the loosely coiled vas deferens running along beside it, plainly visible through the integument. Female reproductive system: Ovary occupies columellar side of visceral mass for approximately first whorl. It is cylindrical in shape, composed of loose white granules, not macroscopically organized, enclosed in an encapsulating membrane which tapers anteriorly and becomes the oviduct. Oviduct passes down columellar side of body almost to mantle line, then reflexes upon itself, passes upward, and makes another 180° turn into seminal receptacle. Oviduct throughout is thicker than the oviduct of Ostodes, and upon entry into seminal receptacle, it appears to be coiled, rather than folded, within the encapsulating membrane. Bursa copulatrix similar to that of Ostodes, its duct join- ing that from seminal receptacle in same way to become common genital duct. Common genital duct heavier than that of Ostodes; runs anteriorly along medial margin of uterus. Copulatory pore hidden by a flap of tissue extend- ing from medial margin of uterus to floor of mantle cavity and forming a sort of roof over upper part of infra-uterine Page 238 channel, which is quite short. Entry of common duct into uterus is via an elevated papilla shaped like an inverted heart. Uterus bi-lobed, as in Ostodes. Vaginal orifice a triangular slit on medio-ventral aspect of uterus, just below anus, which shows moderate hypertrophy of upper lip. Material: Baukaharijitoa above Dunduy, at 450-600 m elevation, Aoba, New Hebrides, FMNH 109428; Tasmalune, Espiritu Santo, New Hebrides, FMNH 109424. 2. Gassiesia sp. Shell broadly turbinate, wider than high. Dorsal surface with spiral lirae, larger, wider-spaced, and more promi- nent than those of Gonatorhaphe. No radial plicae. Spi- rals continue on ventral surface, but umbilicus has no rim. Aperture round, lip simple. Operculum is similar to that of Ostodes: corneous, transparent with an iridescent sheen, polygyrous. Radula is enough different from that of Ostodes to be useful in differential diagnosis (pp. 202-203). Male reproductive system: Testis occupies whole vol- ume of approximately first whorl. Composed of branch- ing, digitiform alveoli, arranged along outer aspect of collecting tubule on columellar side of whorl. Alveoli much finer, more closely packed, with fewer branches than in Ostodes. Collecting tubule becomes seminal ves- icle, runs forward to enter prostate. Whole proximal tip of prostate turned under, with seminal vesicle entering at very tip. Prostatic lumen in turned-under portion is Y- shaped, stem of Y pointing medially, arms pointing later- ally. Anterior to the turn in the main part of the prostate, the stem of the Y disappears, and the lumen is U-shaped, as in Ostodes. Anterior end of prostate shows small sac, but no “‘safety-valve.” Vas deferens a closed tube. Penis long-threaded, thread/trunk ratio 1/1. In adult speci- mens, thread appears to have begun as a flattened ribbon which twisted upon itself repeatedly, so that its former edges show as two white lines, spiraling down from ‘begin- ning of taper to very tip of thread. Thread of juveniles more rounded, lacks the white lines. Female reproductive system: Ovary as in Ostodes: a ta- pering cylinder filled with loose white granules, enclosed in encapsulating membrane which continues forward as oviduct. Seminal receptacle as in Ostodes, with oviduct folded in one plane within encapsulating membrane. Bursa copulatrix very fragile, very thin-walled. Common duct as in Ostodes, but hidden for most of its length by a thin membrane running from medial aspect of uterus to mantle cavity floor, turning most of infra-uterine channel THE VELIGER Vol. 20; No. 3 into a sort of tunnel. Copulatory pore not located. Uterus with two lobes as in Ostodes. Anus shows great hyper- trophy of edges, especially of upper edge. Vaginal orifice a fairly large triangular slit, apex in, on medio-ventral aspect of the uterus. Material: Station N.C. 15, near Thiem, N. E. New Caledonia, (juveniles, FMNH 159215): Station N.C. 28, W side of main range, 4 up, 6.4km E of Ouegoa, N.E. New Caledonia. (adults, FANH 159358). Both collected by Laurie Price, 1967. 3. Mexcyclotus panamensis (Da Costa, 1903) (Figures 29, 30) Shell broadly turbinate, wider than high. Surface smooth, showing only very fine radial growth lines. Perio- stracum adherent, thin, yellow to yellow-brown. Aperture round, lip simple. Operculum with 2 layers, as in Gona- toraphe, but of more complicated structure. Under layer smooth, shiny, thick in center, thinning towards rim, from which it protrudes as ‘cellophane edge.” Upper layer corneous, transparent, with iridescent sheen for approxi- mately first three volutions; then become heavily im- pregnated with calcium. Each subsequent volution is L- shaped, with the central half lying flat, the outer half standing up almost vertically from the surface. Flat half of calcified layer is overlaid by thin layer of iridescent, corneous material, extending upward from the under layer. Result is a small central corneous area, surrounded by upstanding, calcified lamellae, which are separated from each other by flat areas with a layer of calcium under a layer of corneous material. Environmental debris col- lects in interlamellar areas, where it seems to buttress the very brittle vertical lamellae against breakage. Radula not seen. Male reproductive system (Figure 29): Testis fills ap- proximately first whorl of visceral hump. Internal struc- ture not observed. Seminal vesicle runs down columellar side of body, enters prostate directly, without reflexing upon itself as it does in Ostodes. Prostatic lumen approximately L or T shaped, the short arm or arms lying medio-dorsally, and the main part of the lumen running transversely. Prostate terminatés in narrow, tapered tip, nearer to the base of the penis than in Ostodes. There is neither a prostatic sac nor any “‘safety- valve.” Instead of a closed vas deferens, there is an open seminal groove, running from tip of prostate to base of penis. Penis on cephalic midline, well behind tentacles; short, thick, rugose, tapering abruptly into a slender neck surmounted by a narrowly-ovate, pointed terminal ap- pendage. The seminal groove runs up the posterior sur- Vol. 20; No. 3 THE VELIGER Page 239 2mm Figure 29 Mexcyclotus panamensis, male a — viewed from above; b — testis; c — cross-section of prostate [for explanation of abbreviations see Appendix on foldout] face of the penis and across the neck, then opens into a deep, narrow, sharp-edged, spoon-like depression on the upper surface of the appendage. No important differences in proportion of terminal appendage to length of trunk were observed in the individuals examined. Female reproductive system (Figure 30): Two forms of ovary have been observed in this species. One resembles the ovary of Ostodes: It occupies the columellar side of the visceral hump for approximately one whorl, and con- sists of white granular tissue, held together by an encap- sulating membrane. It is shaped like a slender sausage, tapering at the ends, with the encapsulating membrane continuing anteriorly as the oviduct. Although ovaries with the second form occupy the same position, they are differently shaped: on the side away from the columella, 4 to 5 short, heavy, clavate alveoli protrude into the tis- sues of the digestive gland. The alveoli are arranged one behind the other, in a single row. Although one or two may be bifurcated near the outer end, the alveoli are not otherwise branched. This form of ovary seems to be made of the same sort of tissue as the smooth, sausage-like form; if the encapsulating membrane is torn, the white ovarian granules fall out loosely. As in Ostodes, the oviduct runs down the columellar side of the body, becoming somewhat thickened and con- voluted, then forms a seminal receptacle. A small sac-like structure, the bursa copulatrix, lies nearby, but is not directly connected to the seminal receptacle, and there is no common genital duct. Instead, both the receptacle and the bursa have long, slender ducts which open separately into a diamond-shaped vulval area, which is located at the proximal end of a very short infrauterine channel. The duct from the bursa copulatrix opens directly into the upper corner of the vulval area, while the duct from the seminal receptacle crosses behind the bursal duct and Page 240 LUT THE VELIGER Nolo makes a right-angled turn before entering the vulva distal to the entrance of the bursal duct. The floor of the diamond-shaped area is thrown into deep, longitudinal rugae. Its medial margin is a promi- nent rounded ridge on the right side of the mantle-cavity floor; its lateral margin, and probably roof, is a thin flap of tissue extending medially from the lower portion of the medial aspect of the uterus. At the distal end of the vulvar diamond is a very small hole, hidden in and prob- ably capable of being closed by the vulvar rugae, which opens into the uterus. Two types of uterine structure have been observed. The first is similar to that seen in Ostodes (Figure gc): there are two lobes, arranged in tandem, with the ventral lobe tan in color and of rather denser tissue than the dorsal lobe, which is creamy white and soft. The second type (Figure goc) has a U-shaped lumen, which is continuous throughout the length of the organ. No bi-lobed structure is present, nor is there any differentiation in tissue texture or color in different parts of the uterus. There seems to be no correlation between type of ovary and type of uterus, both types of ovary having been found with each type of uterus. Although it is possible that the clavate ovary is a development of advancing age (the smooth types seen were smaller than the clavate examples), it is difficult to see how the uterine type could change during the life- time of the individual. The placement of the anal orifice and vaginal opening in Mexcyclotus is similar to the general arrangement in Ostodes, except that in the Central American snail, the hypertrophy of the anal orifice is horseshoe-shaped, open end up, and the vaginal orifice is partly obscured by a fold of tissue extending posteriorly and downward from the anterior tip of the upper vaginal lip. Material: La Barca, Finca Lerida, at 1695 m elevation, Boquete, Chiriqui, Panama, FMNH 84611, 84621. (< adjacent column) Figure 30 Mexcyclotus panamensis, female a — reproductive system, with bilobed uterus; b — greatly enlarged view of vulvar area; c — cross-section of uterus with U-shaped lumen [for explanation of abbreviations see Appendix on foldout] Vol. 20; No. 3 4. DISCUSSION Gonatorhaphe and Gassiesia are obviously closely re- lated to Ostodes. The shells of the 3 genera are very simi- lar, using the same basic sculptural elements in varying combinations and with varying degrees of emphasis to produce shells of different appearance. The method is the same as that used between species in Ostodes, but the dif- ferences are greater between genera than between species. In regard to operculum and radula, Gassiesia has a transparent, corneous operculum, very like Ostodes’. Gas- stesia’s teeth, however, differ enough from those of Os- todes to be a useful diagnostic character. The radula of Gonatorhaphe is very similar to that of Ostodes, but the operculum, with its upper calcareous layer overlying the comeous under-layer, is quite different from the simple, corneous Ostodes operculum. Although the anatomical differences between the 3 genera are greater than the differences between species of Ostodes, the differences all involve small details. In the male, Gonatorhaphe’s internal and prostatic arrange- ments are almost identical to those of Ostodes, the only differences being in the thread or terminal appendage of the penis. In Gassiesia, the testicular alveoli are smaller, more numerous, and less branched than are those of Os- todes. The upper end of the prostate is slightly different, and the penial thread is twisted. The 3 genera are alike, however, in having a closed vas deferens, and all have the penis on the dorsal mid-line of the head, behind the ten- tacles, the prescribed Poteriid position. The situation is similar in the female systems. The questions of whether the oviduct folds or coils within the seminal receptacle; whether the wall of the bursa copula- trix is thick or thin; whether the infra-uterine channel is open or roofed-over, long or short — are all essentially minor points. All 3 genera have the oviduct, seminal re- ceptacle, and bursa copulatrix opening into the uterus through a common duct — in accordance with Tielecke’s definition of the Poteriidae. Mexcyclotus is more strongly differentiated from Os- todes in shell, operculum, and anatomy, than Gonatorha- phe or Gassiesia. The shell is smooth and unsculptured; only O. cookei of the twelve species of Ostodes is smooth. Mexcyclotus has a heavily calcified operculum, in contrast to the corneous operculum of Ostodes. In the male, the testis is smaller in Mexcyclotus than in Ostodes; the prostatic lumen is L- or T-shaped, rather than U-shaped; there is an open seminal groove rather than a closed vas deferens, and a spoon-like penial append- age rather than a smooth thread. Nevertheless, the penis is in the normal midcephalic position for the Poteriidae, and the cited differences do not seem to me to be sufficient THE VELIGER Page 241 cause for removing Mexcyclotus from Tielecke’s family. The differences between Mexcyclotus and Ostodes are much greater in the female reproductive system than in the male system. In Ostodes, there is a common duct serv- ing the oviduct, seminal receptacle, and bursa copulatrix; in Mexcyclotus, the ovary and seminal receptacle enter the vulval diamond through one duct, while the bursa copulatrix enters through another. Can this be reconciled with Tielecke’s definition of the family based on a single common duct? I think so. The basic requirement is that the products from the 3 organs enter the uterus together, and they do — they meet and mingle among the rugae of the vulval area and drain, together, through the single hole at the bottom of the vulva, into the uterus. This is a larger difference than exists between Ostodes and Gas- stesia or Gonatorhaphe, but the functional pattern is the same. For comparison, let us consider the situation in 2 of the other families. In the CycLoPpHorIDAE, either the ovi- duct, receptacle and bursa all enter the uterus separately (Cyclophorus), or the oviduct enters through the recep- tacle, and the bursa enters separately (Spirostoma). In the male, the penis is located behind the right tentacle, and there is an open sperm groove. In the MAIZzANIIDAE, the penis has an open sperm groove and an accessory flagel- lum, and is located behind and at the same height as the right tentacle. The seminal receptacle drains into the ovi- duct, and the oviduct and bursa each have a very long, individual duct leading to the uterus. The differences between these families and the PoTERIIDAE are much greater than the differences among the genera within the Poteriidae which I have examined. The differences I have seen between Ostodes, Gonatorhaphe, Gassiesia, and Mex- cyclotus are certainly ample to differentiate them as gen- era, but they are not important enough to warrant separa- tion at the family level. VII. DESCRIPTIVE GEOGRAPHY A. Location Samoa consists of a group of islands at the westernmost extremity of Polynesia, lying between 11° and 15° S lati- tude, and longitudes 169° to 173° W. In Western Samoa, two small islands, Apolima and Manono, lie between two major islands, Upolu and Savaii. Savaii is the largest of the islands, being 50km long by 28.8km wide, with a maximum altitude of 1784m. Upolu, lying southeast of Savaii, is also 50 km long, but only 17.3 km wide, reaching Page 242 an altitude of 1170m. Tutuila, the largest island of Amer- ican Samoa, lies SE of Upolu. Tutuila is much smaller than Upolu, having an area of only 133.1 km. Matafao Peak, on Tutuila, reaches an elevation of 643m. Manua lies E of Tutuila, and is smaller; Rose Island lies E of Manua, and is smaller yet. Swain Island lies N and slightly W of Tutuila. The main Samoan islands are the tips of submerged volcanos, and volcanic activity has been ob- served as recently as 1911 (ScHRoTH, 1971: 291). B. Climate The climate of Samoa is hot and wet. The southeast tradewinds flow along both sides of the high central long1- tudinal axis of the islands, dropping their moisture fairly evenly on the north and south sides. Only the northwest tips of the islands can be said to have an actual dry season. For the rest of the area, the mean annual rainfall ranges from 3124cm to more than 500cm. The rainfall increases with increasing altitude, the central uplands receiving more moisture than the coastal lowlands. The tempera- ture falls an average of 1.5°C for every 300m increase in altitude (WricHT, 1963: 30), with the mean annual temperature ranging from 25.5° C in the coastal lowlands to 15.5 in the central highlands of Savaii. C. Vegetation The lowlands, from sea-level to about 225m, used to be covered with a forest whose canopy reached 30-—39m high. Scattered remnants of this forest remain at present. The foothill forests, extending from approximately 225 -— 540 m elevation, have a canopy height approximately the same as the lowland forests. They have more tree ferns, and the trunks of the canopy trees are adorned with mosses and lichens. The upland forests, from 540-1200 m, have still more tree ferns and ground ferns, more mosses and lichens, and an ample representation of monocotyledon- ous trees and epiphytes, including orchids and perching lilies. There is a 24—74cm layer of leaf litter on the ground. An extensive list of the various species of trees, shrubs, and vines found in these classes of forest may be found in WricuT (1963: 35 — 38). THE VELIGER Vol. 20; No. 3 VIII. ZOOGEOGRAPHY A. Spatial 1. DISTRIBUTION on ISLANDS During their field work in 1965, Dr. Solem and Mr. Price collected on both major islands of Western Samoa. The material available to me from Savaii came from a wide crescent including the western, southern, and eastern sections of the island. From Upolu, there was material from both north and south coasts, the easternmost tip, and the central and west-central uplands and foothills. The altitude range was from sea-level forest to 750m. The material from Upolu contained individuals belong- ing to seven species: Ostodes gassiesi, O. plicatus, O. re- ticulatus, O. savaii, O. exasperatus, O. upolensis, and O. tiara. No examples of O. cookez were present in this col- lection. From Savaii came individuals belonging to six species: O. gassiesi, O. savaii, O. exasperatus, O. garretti, O. llanero, and O. upolensis, of which only one specimen was found. In 1975, Mr. Price collected on Tutuila, Amer- ican Samoa, and sent back examples of O. strigatus and O. adjunctus from almost the whole length of the main. central ridge, as well as the southern coast, taken at alti- tudes of 60m to 390m. Four species of Ostodes from Western Samoa occur on both islands: O. gassiest, O. savaii, O. exasperatus, and O. upolensis. Ostodes plicatus, O. reticulatus and O. tiara (as well as O. cookez) have been found only on Upolu, while O. garretti and O. llanero are known only from Savaii. Although O. strigatus has been reported from Western Samoa by older authors, it is now known only from Tutuila, as is O. adjunctus. 2. SYMPATRY Over both islands of Western Samoa, the distributional pattern is one of sympatry. Of 30 collecting stations, only 9 yielded but one species. Of these 9 stations, 4 had only one or two specimens, and hence yield no significant data. In regard to the other 5 single-species stations, the 4 on Upolu yielded from 6 to 26 shells each, while Station 32, Savali, gave 25 examples of Ostodes savaii. Each of the other 21 stations gave at least 2 species; while no station on Savaii had more than 3 species in residence, Stations 2 and 26 on Upolu each had 5 species, and Stations 8 and Vol. 20; No. 3 18 each had 6 species. Station 18 was the best sampled and most intensively collected of any station visited. In contrast, the situation on Tutuila seems to be one of complete allopatry. Stations 1, 2, 7, and 9, all on the eastern portion of the island, had only Ostodes adjunctus, while Stations 6, 10, 19, and 20, all on the western portion of the island, had only O. strigatus. Upolu THE VELIGER Page 243 3. RELATIVE ABUNDANCE A total of 914 specimens were present in the material available to me. Of these, 526 came from Upolu, 195 were from Savaii, and 193 were from Tutuila. These numbers should not be taken as indicative of the true populations of the islands, however; Savaii is much larger than Upolu, but had less than half as many collecting stations, visited in a much shorter time. Tutuila was quite thoroughly collected. As stated above, 2 species were found on Tu- tuila, 7 species were found on Upolu, and 6 on Savaii, Figure 31 Abundance of Ostodes in Western Samoa E - O. exasperatus; G — O. gassiesi; Gt — O. garretti; L - O. llanero; P - O. plicatus; R — O. reticulatus; S - O. savau; T - O. tiara; U - O. upolensts Page 244 THE VELIGER Vol. 20; No. 3 with 4 species being found on both Upolu and Savaii. These species were not collected in equal numbers, how- ever. Taking all islands together, the most abundantly collected species is Ostodes gassiesi, with 244 specimens, followed by O. savaii (188), O. strigatus (153), and O. plli- catus (106). The 3 species from Western Samoa were taken in different proportions on the two islands. On Upolu, 1.8 times as many specimens of O. gassiesi (188) were col- lected as of the next most abundant species, O. plicatus G 10 P S U T See Station 2 T 19 S P 12 U 7 G R 5 2 2 Station 8 Station -26 (106); O. savaii follows with 77 specimens. On Savaii, there are almost exactly twice as many O. savaii (111) as O. gassiesi (56), and O. plicatus does not occur at all. Fig- ure 31 shows the varying abundance of the several species from both islands. Within a given species in Western Samoa, the pattern of distribution seems to include both clustering and scat- tering. Each of the species which was found in any con- siderable number (more than 25 specimens) showed one to 4 stations having perhaps a dozen or more specimens, with the other individuals of the species divided, singly or in small groups, among a large number of different stations. For example, the 77 specimens of Ostodes savait taken from Upolu came from 13 different stations, only 2 of which yielded 12 or more specimens, and 6 of which gave 3 shells (of this species) or less. As mentioned above, 4 stations on Upolu had 5 or 6 species living together. The relative abundance of species Figure 32 Composition of sympatric populations of Ostodes on Upolu (Species designations as in Figure 31] 45 Station 18 Vol. 20; No. 3 at these stations is shown in Figure 32. Station 2 (5 species, 29 individuals) is an area on the right side of a road, 1.2 km above Afiamalu seismographic station, at 720m ele- vation, in disturbed upland forest. Station 8 (6 species, 47 individuals) includes Station 2 in a wider area of mixed to good upland forest on both sides of the road. The mean annual rainfall for these two stations is 4374—500cm. Station 26 (5 species, 58 individuals) is on the Afiamalu- Lake Lanuto’o track, at 735—750m elevation, in good to excellent upland forest. It receives more than 500cm of rain annually. Station 18 is at the foot of Mt. Solaua, between 180 and 240m elevation, under one large Ficus tree in a banana patch at the edge of the forest. The mean annual rainfall is 375-4374cm. From this station, 116 individuals were obtained, belonging to 6 species. Dr. Solem has told me that this fig tree was an unusually rich collecting area, and was the only station at which he and Mr. Price made an attempt to “get everything.” It is interesting to note that at Stations 26 and 18, one or 2 species are dominant, with the other 4 represented by only a few individuals. At stations 2 and 8 the species are more evenly represented. The latter 2 stations repre- sent larger sampled areas which probably included more microhabitats than the first 2 stations which were, respec- tively, a small remnant forest patch and a single huge fig tree. B. Ecological 1. EFFECTSor ENVIRONMENTAL DIFFERENCES The principal environmental influence on land proso- branchs is moisture supply. Rainfall in Samoa is oro- graphic, with higher altitudes receiving greater amounts of rainfall than lower elevations. Degree of moisture re- tention is controlled primarily by the vegetation cover, with heavy forest retaining moisture at ground level longer than open forest or cleared areas. Ostodes was col- lected only in areas with heavy tree cover (Solem, personal communication). Although specimens were taken in areas having different soil types, these stations also differed in rainfall regimes, and the relative importance of soil type and moisture supply could not be assessed. It is not known whether Ostodes obtains calcium through its plant food or by making use of free calcium in the soil. The influence of a dry season on opercular type has already been mentioned (pp. 200-201). As stated there, specimens of Ostodes savaii from Station 32, Savaii (which THE VELIGER Page 245 receives a mean annual rainfall of 312.5-375 cm and has a more pronounced dry season than other stations) had thick opercula of a different type than individuals of the same species from areas which receive more rain, more evenly distributed through the year. That correlation did not hold for Upolu. Wet area-dry area differences have also been found in the shells and in the male anatomy, and these latter differences do hold for both Upolu and Savail. Specimens of Ostodes savaii from Station 32, Savaii, when compared with all other shells of the same species from the same island, prove to be smaller at the same whorl count (Figure 33). They are from 0.6 to 1.7mm shorter, and up to 0.8mm _ narrower. Additionally, al- though the males still have short-threaded penes, the threads are not quite as short as those of snails from more regularly moist areas. Males from Station 32 have a mean thread-to-trunk ratio of 1/3.2; short-threaded males from wetter areas have a mean ratio of 1/4.6 (Table 7). The height (mm) 5.00 5.25 5:50 5-75 6.00 Figure 33 Effect of moisture on size in Ostodes savaii on the island of Savaii A — males, Station 32 (dry area) @ -— females, Station 32 A —- males, all other stations © - females, all other stations Page 246 THE VELIGER Vol. 20; No. 3 Table 7 Effect of a Drv Season on Penial Thread-to- Trunk Ratio in Short- Threaded Males Species Island Station N Range Mean = S.E.M. Ostodes savatt Savaii 32 (dry) 10 1/1.7-1/ 4.5 1,3.18 £0.28 all other 19 1/1.7 - 1/10.0 14.38 = 0.62 Ostodes gassiest Upolu dry cluster 10 1/2.3-1/ 4.0 1/3.07 = 0.17 all other 26 1/1.7 - 1/10.0 1/4.11 £0.39 total length of the penis is the same in both wet and ‘“‘dry”’ areas; only the proportion of thread to trunk changes with the climate. On Upolu, there is a cluster of collection stations (Sta- tions 5, 7, 14, 16, 17) which has the same climate as Station 32 on Savaii. Specimens of Ostodes gasstesi from the “‘dry cluster,” when compared with individuals of the same species from areas without a dry season, tend to be smaller at the same whorl count (Figure 34). Males from these whorls A 6.00 ® 0 O 4 Oey a OHO) fo) fe) ©e, 5:75 “s@@a @e00°4 0 4 04 eo Onna fe) diameter (mm) Figure 34. Effect of moisture on size in Ostodes gassiesi on Upolu A -— males from “dry cluster” A — males, all other stations @ - females from “dry cluster” © - females, all other stations stations, like the ‘dry area” males from Savaii, have less- short penial threads (mean ratio 1/3) than do males from wetter areas (mean ratio 1/4) (Table 7). As important as the amount and regularity of rainfall, is what happens to the rain after it hits the ground. If a large amount falls, but runs off or evaporates quickly, it has much less value than a retained lesser amount. There- fore, the amount of vegetation — the density of the can- opy and understory — is very important; the denser the cover, the more moisture will be retained, and the more favorable conditions will be for snails. This principle can be illustrated by a comparison of adult female Ostodes gassiesi from Station 2, Upolu, with adult female O. gas- siesi from Station 19, Upolu. Both stations are in the up- land forest, and both have the same type of climate, re- ceiving from 4374—500cm of rain per year, with no dry season. They differ, however, in that Station 19 has just average cover, while that of Station 2 is unusually lush. The effect of the increased cover is to make the moisture supply not necessarily more abundant, but less fluctuating in terms of ground-level near-saturation. The results are very apparent; at the same whorl-count, females from Station 2 are 9.3% taller and 8.7% wider than females from Station 19. Table 8 shows the results of a Student’s t Test, which demonstrates that the size difference is really significant, and not just an artifact of collecting. 2. EFFECTS or SYMPATRY Sample size and distribution were such that comparisons of sympatric with allopatric populations were extremely difficult. In species with one good-sized allopatric popu- lation, individuals sympatric with other species were usu- ally so scattered that meaningful statistical analysis was impossible. Indeed, there was only one instance in which I had both sympatric and allopatric populations of work- able size: Ostodes plicatus from Upolu. Ostodes plicatus lives by itself at Station 39, and shares Station 18 with 5 other species. Comparison of 5 adults from Station 39 with 32 adults from Station 18 is shown in Figure 35; the Vol. 20; No. 3 THE VELIGER Page 247 Table 8 Correlation Between Shell Size and Density of Vegetation in Adult Female Ostodes gassiesi from Upolu Station 2:N=8 dense cover Mean + S.E.M. 1225532105338 11.89 + 0.093 5.616 + 0.055 Height (mm) Diameter (mm) Whorls Station 19:N=9 ate average cover with Mean = S.E.M. 15 df. Significance 11.38 + 0.159 2.283 > .98 10.86 + 0.118 6.764 = .999 5.619 + 0.055 snails from the allopatric population are all larger than those from the sympatric population. Perusal of fragmen- tary data on other species suggests that “larger when al- lopatric’” may be the general rule in Ostodes, but it is impossible to be sure. As to the reason for the larger size of individuals in an allopatric population, it may be that Ostodes require a trace element that is in short supply, but no specific information is available. 12 diameter (mm) It 10 5:25 5:59 5:75 6.00 whorls Figure 35 Effect of sympatry on size in Ostodes plicatus on Upolu A - males from Station 39 (allopatric) ; Station 39 © - females from A - males from Station 18 (sympatric) ; Station 18 @ — females from The only anatomical difference that may be related to sympatry has already been mentioned (on p. 228). The penial thread of Ostodes exasperatus is long in the Upolu population, which may be sympatric with O. savait, a short-threaded species. On Savaii, where the two species are known only from allopatric populations, O. exaspera- tus has a short penial thread. No other instances of dif- ferences in thread length between sympatric and allopat- ric populations were noted, and no consistent differences in the female reproductive system were found. 3. SPECIES SELF-RECOGNITION MECHANISM Species-specific recognition features in the terminal geni- tal organs are well-known in pulmonate land-snails. ‘The differences in sympatric European helices have been known for more than a century (ScHmipT, 1855), and ex- tensive differences in Pacific Basin endodontids exist (SoLEM, 1976), but this phenomenon has not, to my knowledge, been reported in terrestrial prosobranchs. TIELECKE (1940), Morrison (1955), and THOMPSON (1967, 1969) studied the reproductive anatomy of Pacific and Middle American cyclophorids, but were concerned with generic and family relationships. Rarely did they deal with sympatric taxa, and they did not mention any possible species-recognition characters. As has been mentioned previously (p. 242, Figures 31- 32), the prevailing distributional pattern of Ostodes in Western Samoa is one of sympatry. One of the purposes of this investigation was to discover, if possible, the species isolating mechanism that enables as many as 6 different but closely related species to live together in a small area without losing their identities.While the actual isolating factors are unknown, I believe that the most important factor in species recognition by the snails is the structural differences found in the pallial reproductive organs: the penial thread-to-trunk ratio in the male, and the shape and orientation of the anus and vaginal orifice as well as the location of the copulatory pore in the female. Figure Page 248 LT ST THE VELIGER Vol. 20; No. 3 Figure 36 Arrangement of pallial genital organs and hypobranchial gland in female Ostodes, with corresponding penis types, by species A — Ostodes adjunctus: penis as shown, 82%; none with bulbs E — Ostodes exasperatus: LT (long thread) penis - 100% Upolu males, 33% Savaii males, 25% with bulbs; ST (short thread) penis - 67% Savaii males, 67% with bulbs G -— Ostodes gassiesi: penis shown, 95%; 58% with bulb Gt — Ostodes garretti: as shown, both specimens, no bulbs L - Ostodes llanero: one specimen, no bulb P — Ostodes plicatus: b — vagina and anus bent medially, 80%; 36 shows diagrams of the typical arrangement of the man- tle cavity in the female of each species, along with the appropriate penial type. The hypobranchial glands are included in the diagrams; although the secretion of this s — vagina and anus straight, 20%; penis as shown, 97%, no bulbs R - Ostodes reticulatus: penis as shown, 100%, no bulbs S — Ostodes savaii: ST (short thread) penis, 86%; LT (long thread), 14%; 50% have bulb St — Ostodes strigatus: penis as shown, 100%, no bulbs T — Ostodes tiara: s -“smooth” penis, 67%, none with bulb; b - “bumpy” penis, 33%, all with bulbs U — Ostodes upolensis: penis as shown, 83%, no bulbs gland has not been demonstrated to have any sexual sig- nificance, its configuration is species-specific, and may possibly be an additional factor in the species-recognition mechanism. Vol. 20; No. 3 From the diagrams, it can be seen that the onus of rec- ognition falls sometimes on the male, sometimes on the female. A female of Ostodes plicatus, for example, could easily distinguish between a male of her own species (long thread without a bulb) and a male of O. gassiesi (short thread, probably with a bulb). She might, however, have difficulty in distinguishing a male of O. plicatus from a male of O. reticulatus (both with long, bulbless threads). The male in question, however, could easily distinguish between a female of O. plicatus (copulatory pore at cx- treme rear of mantle cavity, vaginal orifice a largish, round hole, both vagina and rectum probably bent medi- ally at a sharp angle) and a female of O. reticulatus (cop- ulatory pore well down from rear wall of the mantle cav- ity, vaginal orifice a smallish oval hole, vagina and rectum pointing straight ahead). Although matings between dif- ferent species of Ostodes are probably physically possible, I doubt that they occur; if they begin, I doubt that they proceed beyond the exploratory stage. ‘The configurations of the terminal genitalia are so distinctively species-spe- cific that mis-mated pairs undoubtedly quickly recognize unlikeness and desist. C. Patterns of Diversity ‘Twelve species are now recognized in the genus Os- todes. ‘Vheir recognition was made possible by a correla- tion of shell characters with anatomical features. Although the snails probably differentiate like from unlike on the basis of the pallial anatomy alone, human observers must consider both shell and anatomy in identifying to the species level. Size, shape and proportions of the shell as well as the occurrence and relative prominence of sculp- tural clements will serve to identify O. adjunctus, O. upolensts, and O. cookez, but the identities of the other species are very difficult to establish positively without reference to the anatomy. ‘There are several sets of sibling species in the genus: O. plicatus and O. gassiest; O. tiara and O. garretti; O. strigatus and O. reticulatus; O. savaii and O. exasperatus; and O. exasperatus and O. llanero, at least as juveniles. In each of these sets, the conchologi- cal differences, while present and consistent, are relatively small-scale, and may well be obliterated in an old, worn shell. ‘Phe differences in the pallial anatomy are of much greater magnitude, and should always be relied on in cases where identification by shell alone is somewhat doubtful. If a shell is empty as well as badly worn, it may well be impossible to decide to which species of a sibling pair it belongs though the pairs may be differentiated on propor- tions alone. ‘The material available to me had certain drawbacks. Firstly, 1 did not collect it myself, it was not collected THE VELIGER Page 249 randomly, and only partial coverage of the islands’ area was attempted. Because of these limitations, I am unable to make definitive statements concerning the relative abundance of the various species. The distributional in- formation presented here must be considered partial and tentative until a great deal more field work can be done, planned in such a way as to minimize collecting bias. Secondly, the material was preserved in alcohol, and was not seen by me until it had been so preserved for over a year. I was therefore unable to attempt any histological work or the analysis of the stomach contents. Until I can obtain material properly preserved for histological exam- ination, the designation of certain female structures as “seminal receptacle” or “‘bursa copulatrix’’ must be con- sidered tentative. Until I can observe the living snails feeding or examine the stomach contents of fresh speci- mens, or both, any niche differentiation based on food selection (or indeed any other behavioral characteristic) must remain unknown. This study of Ostodes has uncovered a certain amount of new information about this one genus within the Po- teriidae. Only a very limited amount of material from other poterid genera was available to me. Without access to a great deal more material for outgroup comparisons, it would be premature to attempt to make any statements concerning the directions or patterns of evolutionary change in the family. IX. SUMMARY The land snail genus Ostodes in Samoa contains 12 spe- cies: 8 are found on Upolu, 6 on Savaii, and 2 on Tutuila, with 4 species occurring on both Upolu and Savaii. Of the 12 species, 3 were unknown prior to this study. Ihey are O. reticulatus, from Upolu; O. llanero, {rom Savaii; and O. exasperatus, found on both Upolu and Savail. Most species in this genus cannot be identified by study of the shell alone; examination of the pallial anatomy is necessary for the separation of sibling species. Although there are easily observable differences between species in the hypobranchial gland, the chicf characters that serve to separate species are found in the pallial reproductive systems of both sexes. In the male, the proportionate lengths of the distal thread and trunk of the penis vary with the species. In the female, the shapes, positions, and relationships of the anus, vaginal orifice, and copulatory pore are species-specific. he species self-recognition mechanism is thought to be based on these differences in genitalia. The anatomical characters confirm ‘Tielecke’s placement of Ostodes in the Poteriidae. Opercular characters are useless as a diagnostic tool at the species level, as the 6 types of operculum found in XI. APPENDIX: Abbreviations Used in Anatomical Figures A anus : AG albumin gland BCP _ bursa copulatrix CDO common duct opening CDP —_ common duct papilla CDS common duct slit CG capsule gland COL = “collar” of penis CP copulatory pore CPS copulatory pore, Savaii population CPU copulatory pore, Upolu population DBCP duct of bursa copulatrix DG digestive gland DRCS _ duct of seminal receptacle ECPM encapsulating membrane GO genital opening — HBG __hypobranchial gland ICGD internal common genital duct TUC infra-uterine channel KN “knuckle” of penis LUT lumen of uterus ME mantle edge MI. mantle line OD oviduct OV ovary PCGD pallial common genital duct PR prostate PRS prostatic sac PTA terminal appendage of penis PTH __ penis thread PTR penis trunk R rectum RCS seminal receptacle SG seminal groove > 2 ST stomach STBP _ sub-terminal bulb of penis SV “safety valve” TE testis AP uterus VA vulval area VD vas deferens VES seminal vesicle vO vaginal orifice Page 250 THE VELIGER Vol. 20; No. 3 Ostodes are not species-specific. There are no major dif- ferences in radular structure between species, but a num- ber of small differences have come to light through use of a scanning electron microscope. Sexual dimorphism is confined to size, with females averaging larger than males of equal age. No significant differences between sexes as to shell shape or sculpture were observed. The approximate age of a specimen can be judged by changes in the terminal portion of the body whorl involving sculpture, rate of decoiling, and thick- ness of the parietal callus. The size of the shell, and in one case the character of the operculum, are influenced by both the amount and regularity of rainfall, and the density of the covering vegetation which serves to retain moisture. X. LITERATURE CITED Anton, H. E. 1839. Verzeichniss der Conchylien. Ciencu, WILLIAM JAMES 1949. Cyclophoridae and Pupinidae of Caroline, Fijian, and Samoan Halle, 110 pp. Islands. Bull. B. P Bishop Mus. 196: 1-52; figs. 1- 28 Cox, James C. 1870. Descriptions of seventeen new species of iand shells from the south-sea islands, in the cabinet of Mr. John Brazier of Sydney. Proc. Zool. Soc. London 1870: 81 - 85 (June 1870) Creek, GwENDOLINE A. 1951. The reproductive system and embryology of the snail Pomatias elegans (Miiller). Proc. Zool. Soc. London 121: 599 - 640; 18 figs. 1953. The morphology of Acme fusca (Montagu) with special reference to the genital system. Proc. London Malacol. Soc. 29: 228 - 240; 4 text figs. FiscHEr, Paut 1880-1887. Manuel de conchyliologie. 1138 text figs. Fretter, Vera « ALASTAIR GRAHAM 1962. British prosobranch molluscs, their functional anatomy and eco- logy. London, Ray Soc. xvi+ 755 pp.; 317 text figs. Garrett, ANDREW 1887. The terrestrial Mollusca inhabiting the Samoa or Navigator Is- lands. Proc. Acad. Nat. Sci. Philadelphia 1887: 124 - 153 GouLp, Aucustus AppISoN 1848. Species of Cyclostoma from the collection of the Exploring Ex- pedition. Proc. Boston Soc. Nat. Hist. 2: 204 - 206 1852-1856. U.S. Exploring Expedition. 12: Mollusca and shells. xv+510 pp. Paris, 1369 pp.; 23 plts. 1860. Atlas. Mollusca and shells. [for the above] C. Sherman, Philadelphia: 16 pp.; 52 plts. 1862. Descriptions of new genera and species of shells. Proc. Bos- ton Soc. Nat. Hist. 8: 280 - 284 Gray, Joun Epwarp 1847. A list of the genera of Recent Mollusca, their synonyma and (February 1862) types. Proc. Zool. Soc. London (for 1847) [prt. 15] (178): 129 - 219 (November 1847) 1852. Catalogue of Phaneropneumona in the collection of the British Museum. 324 pp. Homsron, & Honoré JAcQuiNnoT 1842-1854. Voyage au Péle Sud. Jounson, RicHarp IRwIN 1964. The Recent Mollusca of Augustus Addison Gould. Mus. Bull. 239: 182 pp.; 45 plts. 5: Molluscs, 131 pp.; 29 plts. U. S. Nat. (28 July 1964) Kose.t, WILHELM 1902. Cyclophoridae. map Kosett, WILHELM & OTTO voN MOLLENDORFF 1897. Catalog der gegenwartig lebend bekannten Pneumonopomen, Nachr. Deutsch. Malak. Gesellsch. 29(7 & 8): 105-120 Morrison, JosepH PauL ELDRED 1955. Notes on American cyclophorid land snails, with two new names, eight new species, three new genera, and the family Amphiclotidae, separated on animal characters. Journ. Wash. Acad. Sci. 45 (5): 149 - 162; 31 figs. Mousson, A. 1865. Coquilles terrestres et fluviatiles de quelques iles de l’océan pacifique, recueillies par M. le Dr. E. Graeffe. Journ. de Con- chyl. 13: 164-209; plt. XIV 1869. Faune malacologique terrestre et fluviatile des tles Samoa, pub- liée d’aprés les envois de M. le Dr. E. Graeffe. Journ. de Conchyl. 17: 323 - 390; plts. XIV-XV Pease, WILLIAM HarPER 1871. | Catalogue of the land-shells inhabiting Polynesia, with remarks on their synonymy, distribution, and variation, and descriptions of new genera and species. Proc. Zool. Soc. London 1871: 449 - 477 (August 1871) In: Das Tierreich 16: 662 pp.; 110 figs.; 1 PFEIFFER, LUDWIG 1852. Monographia pneumonopomorum viventium 1: xvi+439 pp. London, Cassellis 1854. Die gedeckelten Lungenschnecken (Helicinaceae et Cyclostoma- ceae). In: Chemnitz, Conchylien Cabinet 1[19(1)]: 400 pp.; 50 pits. Raup, Davm « STEvEN M. STANLEY 1971. Principles of paleontology. Co. 388 pp.; 200 figs. Réctuz, Constant A. 1851. Description de quelques coquilles nouvelles. chyl. 2: 194-216; plts. 5, 6 Reeve, Lover. Avaustus 1862. Conchologia Iconica XIII. Section: Cyclophorus; plts. XIV, XVI, XVII Scummvt, A. 1855. Der Geschlechtsapparat der Stylommatophoren in taxonomischer San Francisco, W. H. Freeman & Journ. de Con- Hinsicht. Abhandl. naturwissensch. Ver. f. Sachsen & Thiringen in Halle 1: 52 ScurotH, C. L. 1971. Soil sequences of Western Samoa. Pacif. Sci. 25 (3): 291-300; 2 figs. So_emM, ALAN 1959. Systematics and zoogeography of the land and fresh-water Mol- lusca of the New Hebrides. Fieldiana: Zoology 43: 359 pp.; 38 text figs.; 34 plts. 1972. Malacological applications of scanning electron microscopy. II. Radular structure and functioning. The Veliger 14 (4): 327 - 336; 6 plts.; 1 text fig. (1 April 1972) 1976. Endodontoid land snails from Pacific Islands. Part I. Family Endodontidae. Field Mus. Spec. Publ. Zool. xii+508 pp.; 208 figs. Souversiz, M. 1858. Description d’espéces nouvelles. Journ. de Conchyl. 7: 289 - 296; pit. VIII THIELE, JOHANNES 1929[-1931]. Handbuch der systematischen Weichtierkunde Jena, Gustav Fischer, 1154 pp.; 893 text figs. Tuompson, Frep G. 1967. A new cyclophorid land snail from the West Indies (Prosobran- chia), and the discussion of a new subfamily. Proc. Biol. Soc. Wash. 80: 13-18; figs. 1-7 1969. Some Mexican and Central American land snails of the family Cyclophoridae. Zoologica 54 (2): 35-78; plts. 1-7; figs. 1-14 TiELECKE, Hans 1940. Anatomie, Phylogenie und Tiergeographie der Cyclophoriden. Arch. f. Naturgesch. N. EF 9 (3): 317-371; 24 figs. Torre, CARLOos DE LA, Pau BartscH & JosEPpH PauL ELDRED MorRISON 1942. The cyclophorid operculate land mollusks of America. Bull. U.S. Nat. Mus. 181: 306 pp.; 42 plts. Waicut, A.C. S. 1963. Soils and land use of Western Samoa. New Zeal. Dept. Sci. & Industr. Res., Soil Bur. Bull 22: 188 pp.; 41 figs.; 2 maps Vol. 20; No. 3 THE VELIGER Page 251 Spawning and Early Life History of Murex pomum Gmelin, 1791 EUNA A. MOORE Department of Biology, University of the West Indies, Barbados FINN SANDER Bellairs Research Institute, St. James, Barbados (2 Plates; 2 Text figures) INTRODUCTION THE LITERATURE ON reproduction and larval develop- ment of tropical marine prosobranchs has recently been greatly expanded. Of this group, the genus Murex has been investigated by KNupsEN (1950), NATARAJAN (1957), CERNoHorSKy (1965), Fioront (1966), GoHar & EISAWY (1967), D’Asaro (1970), SpicHT, BrKELAND & Lyons (1974), and BaNDEL (1976). Observations on the spawning behavior and egg capsules of M. pomum have been made by D’Asaro (of. cit.) and BANDEL (of. cit.), but as these formed parts of general surveys of many prosobranchs from the South Florida-Bahamas region and the South Caribbean Sea respectively, detailed accounts, especially of the development of the hatchlings, were not given. This study, inasmuch as it was possible to monitor closely the early development of M. pomum from egg masses de- posited by adults in the laboratory, will extend existing information on the species. Specifically, the use of two separate sample groups of M. pomum differing in size and geographical origin (St. Kitts and St. Vincent, West Indies) served to establish what is the norm for this species, and also helped to provide comparative infor- mation on spawning as well as quantitative and qualita- tive data on the capsules, eggs and young hatchlings. ACKNOWLEDGMENTS We wish to thank Dr. Tom M. Spight and Dr. Jorgen Knudsen for offering relevant and up-to-date material from the literature. We are also very grateful to Dr. Carol Lalli for reading the original manuscript and providing useful criticism. The work was supported jointly by a subvention from the University of the West Indies to the senior author and a grant in aid of research from the National Research Council of Canada to the junior author. DESCRIPTION, DISTRIBUTION AND HABITAT Murex pomum is among the larger of the Muricidae. HumpuHrey (1975) gave measurements of adults of 2 to 4.5 inches (ca. 5-11.5cm), but our largest specimen was just over 12.5cm long. The shell is heavy and strong with a rough surface, but shell quality varies impressively from one adult to the next and with the age of the newest varix. The colour of the shells is usually a combination of cream, tan, and dark brown, and occasionally tints of purple. The interior aperture is polished, and most often is coloured yellow to ivory, but may be light pink, and least often has all these shades inter-mixed with purple tints. There is usually one dark spot on the upper end of the parietal wall and 2 or 3 on the outer lip. According to Appott (1958) and HUMPHREY (1975), it has a distribution ranging from South Carolina in the southeastern United States to Trinidad in the West Indies. It has been collected from Biscayne Bay in Florida (D’Asaro, 1970), and reported as quite common in Cuba and Puerto Rico (ABsBorTtT, op. cit.), as common in the Bahamas (ZEILLER, 1974) and off the coast of Colombia (BANDEL, 1976), and as the most common Murex snail in Jamaica (HUMPHREY, op. cit.). It has been found by the junior author to be quite abundant in St. Kitts and St. Vincent and off the coast of Venezuela, but for some reason it is very scarce around Barbados. Only a single Page 252 THE VELIGER Vol. 20; No. 3 specimen was taken from numerous dives over years of shell collecting in Barbados. For this study Murex pomum was obtained from Frigate Bay, St. Kitts, at a depth of approximately 4.5 to 6m. The animals inhabited an area of mixed rubble and sand, in which Thallassia testudinum grass flourished, and were invariably seen in groups of 2-4 in shallow depressions. In St. Vincent, //. pomum, which was gen- erally larger than the average St. Kitts specimens, was collected off Young’s Island in 4.5 - 6m of water, also in an area of rubble and sand. However, unlike the St. Kitts specimens, these were always seen buried in the sand with little or no part of the shell protruding above the sub- stratum. Curiously, although HumpnHrey (1975) has described M. pomum as located in every imaginable en- vironment, this species was obtained at only one site in St. Vincent and not in numerous other bays of compa- rable bottom. The species has been described both as a carnivore (BANDEL, 1976; and others) and a browser (ZEILLER, 1974). The specimens under study exhibited both feeding patterns. They were routinely fed on the flesh of Strombus pugilis, alternated with that of the sea urchins, T77i- pneustes esculentus and Diadema antillarum. MATERIALS anp METHODS Eighteen adult specimens of Murex pomum, captured at St. Vincent on the 29" of February, 1976, were transferred to the Bellairs Research Institute, Barbados, where they were kept in a concrete aquarium of 228L' capacity with running sea water. Ambient temperatures averaged 27.1°C. Within 4 weeks of arrival a large mass of egg capsules and a few smaller clusters were deposited by some of the animals. Since the major part of this event occurred over a week-end, when they were not under close obser- vation, there was no certainty which of the animals were involved in the process. Just over a week later a speci- men was observed to have deposited a few capsules. It was immediately marked with a numbered tag attached by nylon fibre and kept under close watch from 8:00 p. m. to 10:00p.m. each day. It was the only specimen seen associated with the increasing egg mass and was not ob- served to leave the mass until 6:30p. m. on the 3" day. Thus, not only was the identity of the egg layer certain, but also the time taken for deposition was determined to within minutes. Figure 1 is a photograph of the egg- * “The Veliger” has adopted the SI metric prefixes and abbrevia- tions exclusively, according to which L stands for liter. depositing female and the egg-capsule mass. The latter is hereinafter referred to as sample A. The egg-capsule mass was removed from the concrete wall of the tank and after careful blotting, was weighed and partitioned. The largest portion was put into a nylon basket of mesh size 0.239mm, which was then suspended in the water table and kept under observation. The second portion was frozen for later biochemical and histological studies, and the third was used for estimation of the total number of egg-capsules, number of eggs per capsule, and for dry weight determinations. In May, another batch of 20 Murex pomum was cap- tured in waters off St. Kitts, labelled, and housed in the same water table. No egg-capsules had been deposited up to 10:00p. m. on June 17", but by the following morn- ing there was a mass of oothecae on the upright wall of the tank. Five specimens were observed associated with this egg-mass, but 4 of these moved off at various times during that day and only one continued to be associated with the growing mass until 5:oop. m. on the 21". Thus in both cases a single female was responsible for deposit- ing at least the majority of the egg-capsules. This second capsule mass, now designated sample B, was treated like the first. Daily samples of 5-6 capsules were removed from the egg-mass and preserved for later study of development. Almost daily observations were made on the development of the hatchlings. From the 21* day onward, samples of the larvae were transferred to nursery baskets, sometimes still in the capsules and in other instances without cap- sules. In this way it was possible to determine at what stage they were most likely to survive removal from the egg capsules. The surrounding fluid in the nursery basket was regularly sampled for veligers. The sizes of egg capsules, larvae and adults were deter- mined and a size factor for the latter was calculated from length and width (L X W). RESULTS Adult Sizes The adult animals from St. Kitts and St. Vincent clear- ly constituted two distinct size populations. Average length and width for the St. Kitts snails were 6.6cm (5.3-7-9cm) and 4.5cm (3.6-5.4cm) respectively, while the same measurements for the St. Vincent speci- mens were 10.4cm (8.5 -12.5cm) and 6.5cm (5.8 - 7.9 cm). The single female (from St. Vincent) observed in the deposition of sample A egg mass measured 12.3cm in length and 6.95cm in width and weighed 191.50g Vol. 20; No. 3 THE VELIGER Page 253 after egg deposition. Of the 5 specimens from St. Kitts associated with capsule mass B, average measurements were: length, 7.1cm; width, 4.8cm; and weight, 62.70g. The female that deposited for the longest time was 7.7cm in length, 5.4cm in width, and weighed 76.86¢. Egg Deposition All the egg-capsule clusters were deposited just on the waterline in the aquarium. Areas free of fouling seemed to have been preferred, but the animals themselves also seemed to do some clearing away of tubiculous worms. The capsules were very firmly cemented to the upright surface of the tank and to each previous layer by their basal points. The time taken for deposition by the first single female was 54 hours. It is not known whether this female had contributed to any of the other clusters de- posited earlier or whether it had spawned earlier in the year but did not do so again up to October of that year. The same observations hold for the second set of deposi- tors. In this instance, one female was spawning continu- ously over a period of approximately 87 hours. The total wet weight of sample A capsule mass was 129.90g and that of sample B, 53.49g. The number of capsules esti- mated from counting an approximate 4 subsample was 1 862 for sample A and 1 662 for sample B. Capsules from sample A contained 45-110 eggs and those from B, 25 - 45. The capsules were filled with additional albumin- ous fluid and lined by a thin albumen membrane. Description of Egg Cases The egg cases of Murex pomum are roughly tongue- shaped with the convex side uppermost. This side is patterned with striations of variable thickness. The stria- tions are mostly in the vertical plane but those towards the shoulders branch and anastomose to make a network. The pattern is variable. The concave side has striations of much thinner fibres. An exit-window of much more even consistency is located in the apical half of this side (Figure 2). The egg capsules of sample A averaged 7.8+0.5mm at the base and 7.5+0.5mm in height. For sample B, width and height measurements averaged 5.30.4 and 5.40.5mm respectively. There was little difference in thickness, capsules in sample A averaging 2.0+0.2mm and in sample B, 1.8+0.1mm. The oothecae are leathery in texture and strength and of a creamy off-white colour when deposited, but develop a more yellow colour with age. They retain this colour long after the eggs have hatched. Several times during the late pre- and early- hatching stages, the oo-tests became infested and covered with black fungus. For treatment of this infection, 2mg of streptomycin and 2mg of sulphadiazine were dissolved in 1L of millipore-filtered sea water, and the capsules were immersed therein for 8 hours. But, curiously, after hatching was completed, the portions of egg capsules kept in sea water did not develop any fungal growth or discolorations. Pre-Hatching Development Development was not synchronized throughout the egg- mass, nor within the same capsule. This was evidenced by the fact that, although deposition of egg capsules was accomplished within 54 and 87 hours for samples A and B respectively, hatching continued over a 12 day period in both instances. Furthermore, individual capsules removed from the mass during any of the hatching days contained Figure 2 Murex pomum egg capsules taken from an original drawing by the late Dr. Gunnar Tuorson, by courtesy of Dr. Jorgen Knudsen Page 254 young of different sizes and showing different increments of growth. Nevertheless, it was possible to follow the de- velopmental sequence quite easily. The qualitative data on larval stages, hatching times, and later development showed 100% comparability. In the most advanced larvae, the foot was apparent by the 18" day after spawning had ended. The velum was formed and so was the pulsatile larval heart. The frequen- cy of the heart beat seemed to be related to velar activity, ceasing for several seconds when the velum was with- drawn. Eye-spots were barely visible. The gut was ob- served extending from the stomodeum, with the gut diver- ticulum, seen as a coarsely granular mass, jutting towards the posterior end of the larva, which was by then encased in a membrane-like shell. Between the 19™ and 21* day torsion took place. New shell material was deposited and the larva then assumed definitive shape. The eyes had been formed, and the gut diverticulum had increased in size. By the 224 day the shell had 24 whorls, was gritty on the outside and quite brittle with a smooth outline on the outer lip. In the St. Kitts specimens, however, the last 4 whorl was marked by a prominent mid-whorl shoulder. At the anterior end of the larva the 4-lobed velum, foot and operculum showed further growth. Orange and brown pigmentation had begun in the foot and opercu- lum. There was also brown pigment in the eyes which were by then supported on short stalks. Tentacles were also formed. The pulsating larval heart could be seen through the still transparent shell just below the outward curvature for the siphon which was not well formed. The gills, however, were present and could be seen coiling backwards from the stomodeum. The mid-gut gland ex- tended into the apical whorl but did not fill it. At this stage the larval velum was very active, and the larvae moved about within the capsules. Some exit-windows of the capsules opened but most of the veligers remained inside. When flushed from the capsules, they swam quite strongly up and down the water column. The foot was by then quite well developed and pigmented, and the larva used it to a limited extent. Larvae, that were trans- THE VELIGER Vol. 20; No. 3 ferred to nursery baskets at this stage, did not survive. The causes of death were not known, but were not related to shell damage inflicted during handling as most shells remained intact. By the 23" day the shell began to develop colour, and a layer of new material was laid down on the outer lip (Figure 3). In the St. Vincent specimens the tubercle marking the beginning of the mid-whorl shoulder was then apparent. The eyestalks had grown to extend beyond the shell. The velum had increased in size — when fully extended each lobe on a sample A specimen measured 1.25mm long. Between the 24" and 26" day the foot became more active and was preferred as a means of loco- motion. Pigmentation intensified, particularly in the shell, and so internal structures were less discernible, except for the mid-gut gland which then occupied more of the 2™ whorl. Shell growth continued at the rate of one layer per day, the layers averaging 324m in width. Larvae from any capsule showed from 2-6 layers on the outer lip. As a result of this variation, their lengths varied in the same way as the lengths of the hatchlings given in Table 1 and shown in Figure 4. At this stage the highest mortality in nurslings from all causes of death (discussed later in this text) was among the lesser developed speci- mens. Table 1 Size variations (in mm) in twenty hatchlings from three connecting egg capsules taken from sample B. L and W denote length and width respectively. — L Ww L WwW 1.4 1.2 1.6 1.2 1.4 Ne 1.6 1.2 1.4 1.2 1.6 1.2 1.4 1.2 1.6 1.2 1.5 1.1 1.6 1.2 15 1.2 1.6 1.3 1.5 1.2 1.6 1.3 1.5 1.2 17 1.2 1.5 1.2 1.8 1.3 1.5 1.2 1.9 1.4 Explanation of Figures 7, 3, 4 Figure 7: Egg capsule mass A and depositor Figure 3: Murex pomum pre-hatching larvae about 24 days after egg deposition showing a day’s increment of shell growth and - developing foot Figure 4: Difference in size of Murex pomum hatchlings taken from the same egg-capsule Tue VELIGER, Vol. 20, No. 3 [Moore & SANDER] Figures 1, 3, 4 Figure 3 Figure 4 Vol. 20; No. 3 There seemed to be a gradual recession in use of the velum, and by the 28" day there was no outward sign of it. When turned over, the protoconchs all righted themselves by extending the foot until contact was made with the surface -— none extended the velum although it had not yet degenerated and could still be seen when the shell was broken. Hatching From observations on the transfers to nursery baskets and daily sampling of the water from the main hatchery basket, it was determined that successful hatching oc- curred from the 26" day onwards. None of the transfers made before that day survived, nor did any of the veligers that emerged into the main basket, even though this could be considered a well-protected environment. By then the larvae had 4-6 extra layers on the outer lip, the apical whorl was almost filled by the hepatopancreas, and the shell was heavily pigmented. Hatchlings varied in size as shown in Table 1 and Figure 4. Hatching continued up to the 38" day, the visually-determined peak being be- tween the 32™ and 33™ days. By the 40" day all veligers remaining in capsules, although well developed, showed no sign of life. This happened mainly in the cases where the windows remained closed, but also in instances where the windows were opened. Necrotic larvae were quickly devoured by saprozoic protozoa, leaving only shells. Most hatching protoconchs crawled out on foot, but many, particularly the early hatchlings, did so when both foot and velum were equally active. Some of them trav- elled relatively great distances in the first few hours after hatching ; many of them crawled up above the water line and perished as a result. Presumably this was an instinctive dispersal response and in their natural environ- ment the problem of desiccation would be of a lesser degree. ‘Table 2 gives a summary of relevant pre-hatching and hatching data for the 2 samples studied. THE VELIGER Page 255 Post-Hatching Development Since hatching continued up to the 38" or 39” day, some later hatchlings undoubtedly were more developed than the earlier ones and so passed some of the so-called post-hatching stages inside the capsule. Early in this peri- od there was marked extension of the eye-stalks and the proboscis became quite prominent. The foot elongated and presumably development and differentiation of in- ternal organs continued — the most obvious of which was the hepatopancreas which continued to extend to- wards the apical whorl. Colour intensified, further occlud- ing internal structures. Visible growth was by the daily increment of shell material at the outer lip of the aper- ture (Figure 5). When 7 - 8 layers had been laid down (2. e., 3 - 5 days after hatching), crenellation of the outer lip began (Figure 6). At the g - 10 layer stage, multiple layers were put down at the same time and same level, somewhat like a set of flounces. This formed the first varix as shown in Figure 7. The lip was continued from the innermost of these at the same rate of one layer per day. The day’s increment could always be seen as the thinnest translucent layer at the edge, since older layers were progressively thickened by new material added from the inside. The second varix was formed similarly to the first at the 12" layer of the new extension, 7. e., roughly 18 days after hatching. The third extension involved 14 layers. This possibly marked the real end of the larval stage, since in the 4" growth stage shell deposition was more complicated. The shell material was much thicker and the surface was rougher and corrugated both vertically and transversely, markedly different from the earlier nu- clear whorls. Fluting at the lip edge was more pronounced so the number of corrugations at the edge increased grad- ually (cf. Figures 6 and 7). This occurred 30 - 32 days after hatching. Because of the intensity of colour and the difficulty in handling the by then very active protoconchs, it was not certain whether growth continued by the same daily increments, but growth measurements done at wider Table 2 Summarized data on the respective spawn from the St. Vincent and St. Kitts animals. H, W and T denote height. width and thickness respectively. Collection Average size measurements Calculated Average Average Average site of capsules average net number of eggs number of eggs Percentage number of of in mm (N = 20) weights of per capsule hatching per of eggs nurse eggs samples H W T capsules in g. (N = 20) capsule (N = 40) hatching per embryo St. Vincent 7.5 7.8 2.0 0.070 78.7 18.3 23.3 3.3 St. Kitts 5.4 5.3 1.8 0.032 34.4 7.9 23.0 3.3 Page 256 time intervals on a single individual showed that the 32 jum per day input was closely approximated (‘Table 3). Table 3 Growth measurements in mm for a single hatchling from sample B. THE VELIGER Date Length Width 20/7/76 1.45 1.20 26/7/76 1.55 1.20 4/8/76 1.80 1.30 10/8/76 1.95 1.35 27/8/76 2.40 1.46 10/9/76 2.90 1.87 14/10/76 3.30 1.90 28/10/76 3.30 1.90 15/11/76 3.30 1.90 By the end of the 10” post-hatching week the snails had each completed one new whorl and were then 3} whorls big. By far the greater portion of the last whorl was of the thicker corrugated shell type. The 3 last meas- urements in Table 3 show that no growth took place for a month after this stage had been reached. Presumably the protoconchs had by then adopted the growth pattern of the adults; this involves massive shell growth over a relatively short period of time with increasingly large “resting” intervals. BANDEL (1976) has described how older well-fed animals will bury themselves in the sand, and put on a new chamber in about a week. Our experi- ence confirms this and shows further that, if interrupted in the process, the murex will carry around the half-com- pleted chamber for weeks until it finds conditions suitable to resume shell building. This seems to involve some factor other than the availability of sand or mud. Survival of Young There was a very high mortality in the hatchlings. Of an estimated 16268 veligers in Sample A, 99% either Vol. 20; No. 3 failed to emerge or died before reaching the age of 3 months, and that in a protected environment. It has al- ready been pointed out that all naturally and artificially hatched veligers died, some from shell damage; many of the veligers and later stages died because their shells were perforated and viscera eaten out by boring worms. Many of the young snails died apparently from dehydration upon crawling above the water level, but others died from no readily discernible cause. The same results were obtained with sample B. The best rearing results were obtained from those cul- tures that contained portions of the egg-capsule mass and turtle grass, Thallasia testudinum. Coincidentally, rela- tively large populations of cyclopoid copepods also devel- oped in the same cultures, and the protoconchs, as they crawled over grass and egg capsules, no doubt fed on the crawling nauplius stages as well as nematode worms and bryozoan larvae found on the turtle grass. Later-stage juveniles were fed on the same diet as the adults. DISCUSSION High mortality is usually concomitant with high fecundi- ty in animals. Despite the results obtained from rearing in a medium with turtle grass and egg capsules, it is most unlikely that lack of food played an important part in mortality among the young. SpicHT (1976) has pointed out that few hatchlings starve to death (citing his ex- perience of keeping many young Thazs lamellosa alive without food for a month or more). Physical stresses and predation are more of a threat to the newly hatched snail than is starvation. Shell damage, dehydration and shell boring by worms naturally qualify as stress factors along with others that would occur in the natural environment. SpiGHT (1975) has calculated that a newly hatched T- lamellosa has a 1 - 2% chance to survive its first 3 months. A snail reaching 3 months has a 35% chance to reach age 1, and older snails have a 40 - 60% chance to survive through subsequent years. With this rapid increase in sur- vival rates a 2mm snail should have much better pro- spects than a newly hatched 1mm snail. Our 1% sur- vival of young tallies with Spight’s 1 - 2% for T: lamellosa. Explanation of Figures 5 to 7 Figure 5: A Murex pomum protoconch showing several daily increments of growth and also the mid-whorl ridge Figure 6: Murex pomum protoconch showing crenellation of lip as well as pigmented foot, gills and portions of the gut Figure 7: Murex pomum shell of 14 week old protoconch showing nuclear whorls and subsequent increments of growth Tue VELIGER, Vol. 20, No. 3 [Moore « SANDER] Figures 5 to 7 Figure 6 Figure 7 Figure 5 Vol. 20; No. 3 It is noticeable from Table 3 that growth in Murex pom- um is relatively fast in the early post-hatching stage, that the 2mm size was reached within one month of hatching, and the 3mm stage in less than one month thereafter. Considering Spight’s calculated increased chances of sur- vival in the open, almost 100% of those reaching the 3 month stage should reach maturity in a protected labora- tory environment. The principal factor upon which depend the early life histories of marine prosobranchs is the type of develop- ment. THorRSON (1940, 1946) has shown that the per- centage of species with a pelagic larval development in- creases when going from the Arctic seas towards the equator. For example, “in the high Arctic seas (East Greenland) about 95% of all marine species of bottom invertebrates seem to develop without a pelagic phase” (THorson, 1950), whereas in New Caledonia 57% of the prosobranchs displayed pelagic development, in the Persi- an Gulf 75%, and off Bermuda about g0% (THoRSON, 1940; LeBour, 1945). On the other hand, KNupDsSEN (1950) determined that 69% of 32 species from tropical West Africa depended on non-pelagic development, while D’Asaro (1970) reported that exactly half of the proso- branchs which he studied from South Florida and the Bahamas relied, too, on this direct form of development. In his later studies, THorSON (1950, 1952) also demon- strated the correlation of a relatively small number of eggs with non-pelagic development in marine inverte- brates. Most of the species with non-pelagic larval devel- opment investigated by Thorson produced less than 1000 eggs per female per breeding season. In contrast, the 2 egg-masses from St. Vincent and St. Kitts were estimated to contain 146539 and 57173 eggs respectively. In each instance, at least the majority of the eggs were contributed by a single snail. D’Asaro (1970), though, found far fewer eggs (g 700) per mass for M. pomum indicating, thereby, a wide variation in individual fecundity in this species. It is tempting to offer the explanation that around small Caribbean islands with narrow shelves, such as St. Vin- cent and St. Kitts, direct development constitutes a means by which snails guarantee that the larval stock is not all dispersed offshore. However, inshore eddy systems acting as retaining mechanisms generally ensure that for such islands at least a portion of the pelagic larvae of benthic invertebrates remain inshore sufficiently long for the planktotrophic larvae to complete development to the crawling stage before settling over a propitious substra- tum. A species which most likely benefits from such re- taining systems is the Rooster Tail Conch, Strombus gal- THE VELIGER Page 257 lus, which in St. Kitts co-exists with M. pomum at Frigate Bay, yet is known to have a pelagic development (D’Asa- RO, 1970). It is, however, more probable that elimination of the pelagic stage is a conservative mechanism adopted in order to lessen the higher mortality rate from preda- tion associated with the temporary planktonic existence. This option possibly allows for the co-existence of M. pomum with S. gallus, inasmuch as it would reduce com- petition between the young of M. pomum and other benthic snails such as S. gallus, especially if the latter also produces large numbers of larvae and breeds through- out the year. Noconclusions regarding spawning seasoncan be drawn from the spawning periods (April and June) of Murex pomum from the present study. Although they agree with the results of D’Asaro (1970) who collected spawn from this species from March through May in subtropical Bis- cayne Bay, Florida, it is likely that this snail is reproduc- tively active all year off the tropical Caribbean islands. In fact, BANDEL (1976) has observed spawning activity by M. pomum at Santa Marta, Colombia, throughout the year. D’Asaro’s observations probably are linked to season- al climatic changes which are more apparent at higher latitudes. Judging from all 3 studies, it appears that communal spawning is characteristic of the species, and that indi- vidual acts of egg mass deposition precipitate spawning in other ripe animals alerted through chemoreception. It is probable that in instances where spawning appears to be limited to a single snail, no other ripe animals are present. The hatching time of 26 to 38 days observed for both the St. Vincent and St. Kitts samples, and the close agreement with the results of BaNDEL (1976), who reported a range of 24 to 39 days, probably indicate a norm for the species of the latter range — at least under tropical con- ditions. D’Asaro (1970) did not give hatching time, but insofar as the accumulated data indicate spawning time to bea latitudinal effect, incubation and hatching times could also be functions of temperature for this species. Table 2 shows that the larger female(s) from St. Vincent deposited larger capsules than the smaller one(s) from St. Kitts. This agrees with the findings of Hancock (1959) and McKenzie (1961) for the muricids Urosal- pinx cinerea and Eupleura caudata respectively. Also, capsules of the larger snail(s) contained the greater num- ber of eggs — a simple relationship noted long ago when ConK.in (1897, 1898), in his now classic studies on the embryology of Crepidula, reported that the number of eggs produced was roughly proportional to the size of the body. Average capsule measurements (H, W and T) of Page 258 samples A and B are comparable to those reported by D’Asaro (1970) (7.5, 6.0 and 2.5mm) and BANDEL (1976) (3-4, 3-5-4 and 1.5mm) for Murex pomum. Approximately the same degree of variation in number of eggs hatching per capsule exists. D’Asaro listed an average of 13 and Bandel a range of 6-15, compared to an average of 18 and 8 for our samples A and B respectively. These findings suggest a rough correlation between cap- sule size and number of eggs hatching - a relationship previously noted by GaLiarDo (1973), SPiIGHT, BiRKE- LAND & Lyons (1974) and CasTILLa & CancINo (1976) for other muricids. There is a remarkable agreement in the percentage of eggs hatching in the 2 egg-masses, and, by extrapolation (on the assumption that the remainder constitute such), the average number of nurse eggs per embryo (Table 2). Neither D’Asaro (1970) nor BANDEL (1976) reported aver- age number of nurse eggs for Murex pomum, but the pres- ent finding (3.3) is relatively low when compared to those given for other species of Murex, which range from 5.9 for M. torrefactus (CERNOHORSKY, 1965) to 91.4 for M. quadrifrons (KNUDSEN, 1950). However, as is known (Hyman, 1967), albumen is engulfed as food by develop- ing encapsulated prosobranchs and some embryos in a capsule often devour the others. SpicHT (1976) has pointed out that, in contrast to the standard size reached when an embryo has its entire yolk supply enclosed with- in its egg membrane (Fioront, 1966), the prosobranch embryo that feeds on nurse eggs will often reach a hatch- ing size much larger or smaller than the mean for its species. This variable hatching size, which may be dis- advantageous if there is an optimum hatching size (SmrrH & FRETWELL, 1974), is demonstrated for a random sample of M. pomum hatchlings (Table 1 and Figure 4) and may, according to SpicHT (op. cit.), be caused by unequal nurse egg supply amongst egg capsules; equal nurse egg supply but unequal embryo numbers amongst egg cap- sules; and unequal sharing of existing nurse eggs by embryos. In conclusion, some of the summarized data have been arranged in Table 4 in order to demonstrate the similar numerical ratios between comparable sets of data for A and B. It is apparent that the ratios of slightly more than 2:1 obtained for all the reproductive parameters reflect the same order of difference in size. Hence, larger body size ultimately translated into higher fecundity, agreeing with the results of Spicht « EMLEN (1977) who noted that clutch size in Thais lamellosa and T. emarginata was directly proportional to body size. Size differences may be attributed to any one or combination of several intrinsic and extrinsic factors. First, it may be a simple function of THE VELIGER Vol. 20; No. 3 Table 4 Table of ratios of comparative reproduction data. A and B represent the known major contributors to egg-mass deposition by the St. Vincent and St. Kitts animals respectively. H, W and T denote average height, width at base and thickness respectively (in mm). A B Ratio Net weight of animals (g) 191.5 76.86 2.49 Size factor of capsules 117.0 51.50 2.27 (H X W XT) Calculated average wet 0.070 0.032 2.18 weight of capsules (g) Average no. of eggs/capsule 78.7 34.4 2.28 Average no. of hatchlings/ 18.3 7.9 2.32 capsule age, in which case the value of the reproductive effort increases with age of the adult since the animals are interoparous. It is possible that, at least for this species, growth does take place after initial spawning at maturity although SpicuT, BIRKELAND & Lyons (1974) have pointed out that most muricid adults do not grow. Hyman (1967), on the other hand, has stated that while some proso- branchs cease to grow after attaining sexual maturity, most do continue to grow, albeit at a diminishing rate, for several years, spawning annually. Second, it may re- flect a difference in nutrient supply at the 2 sites, especial- ly since it is known that size at maturity may vary with availability of food (Spicut et al., op. cit.). Third, we may be dealing with 2 geographically separated groups, each with its own size potential intrinsically induced. Fourth, the sample groups possibly comprise 2 distinct species. This possibility, however, can be! ruled out by visual inspection of the animals. In addition, wet weights and shell dimensions, represented by individual size fac- tors (L X W), are highly correlated, and, although the 2 sets of species characteristics data form separate clus- ters on a graph (Figure 8), the least squares of regression line fitted to the total data has a high correlation coeffi- cient of 0.9g. Statistically, at least, the 2 groups obviously belong to the same universe (species) and size differences are not indicative of any taxonomic differences. To sum- marize, any one or combination of the first 3 factors may apply since in each case individuals in the 2 groups would describe just such a graph as given in Figure 8. Vol. 20; No. 3 THE VELIGER Page 259 D’Asaro, Cuarzes N. 1970. Egg capsules of prosobranch mollusks from South Florida and the Bahamas and notes on spawning in the laboratory. Bull. mar. Sci. 20: 414 - 440; 7 figs.; 2 photogr. Fioront, Pio 1966. Zur Morphologie und Embryogenese des Darmtraktes und der transitorischen Organe bei Prosobranchiern (Mollusca, Gastropoda). Rev. Suisse Zool. 73: 621 - 876; 113 figs. 260 200 Ga.tarpo, C. SS 1973. Desarollo intra capsular de Concholepas concholepas (Brugu- SS iére). Publ. Ocas. 16. Museo Nac. Hist. Nat., Santiago, Chile: 1 - 16 Br Gouwar, H. A. F « A. M. E1rsawy ‘oe 1967. The egg masses and development of five rachiglossan proso- > branchs (from the Red Sea). Publ. Mar. Biol. Sta. Al-Ghardaque, Egypt 14: 216 - 266 3 Hancock, Donatp A. = 1959. The biology and control of the American whelk tingle, Urosal- pinx cinerea (Say) on English oyster beds. Fish. Invest., London Ser. 2, 22: 1-66; 1 fig. Humpugrey, M. 1975. Seashells of the West Indies. William Collins & Sons, Ltd., Glasgow, 351 pp., 32 plts.; 20 figs. Hyman, Lisprz HENRIETTA 1967. The Invertebrates: vol. 6: Mollusca I. New York, McGraw- Hill Book Co. vii+ 792 pp.; 249 figs. KNUDSEN, JORGEN 1950. Egg capsules and development of some marine prosobranchs from tropical West Africa. Atlantide Reprt. 1: 85 - 130; 31 figs.; 7 phot. 8 oO 10 50 100 Lgsour, Mary V. ¥ 1945. The eggs and larvae of some prosobranchs from Bermuda. Size Factor (LXW) . Proc. Zool. Soc. London 114 (4): 462-489; 43 text figs. MacKenzz, C. L., Jr. 1961. Growth and reproduction of the oyster-drill Eupleura caudata " in the York River, Virginia. Ecology 42: 317-338 ; 7 figs.; 5 Figure 8 photographs : 5 3 , Natarajan, A. V. The relationship between wet weight and size of Murex pomum 1957. Studies on the egg masses and larval development of some proso- branchs from the Gulf of Mannar and Palk Bay. Proc. Indian Acad. Sci. B. 46: 170 - 228; 96 figs.; 30 photogr. Smit, C. C. eS. D. FretTweii 1974. | The optimal balance between size and number of offspring. Amer. Natural. 108: 499 - 506; 2 figs. 5 . Spicht, Tom M. Literature Cited 1975. Ona snail’s chance of becoming a year old. Oikos 26: 9 - 14 1976. Hatching size and the distribution of nurse eggs among proso- branch embryos. Biol. Bull. 150: 491 - 499; 3 figs. Aspott, Roszrt Tucker Sriont, Tom M., C. Brxeranp # A. Lyons 1958. The marine mollusks of Grand Cayman Island, British West 1974. Life histories of large and small murexes (Prosobranchia: Muri- Indies. Acad. Nat. Sci. Philadelphia, Monogr. 11: i- viiit+1 - 138; cidae). Mar. Biol. 24: 229-242; 4 figs. pits. 1-5; 11 maps; 7 text figs. (31 December 1958) SricuT, Tom M. « J. EMLEN BanveL, Kraus 1977. Clutch sizes of two marine snails with a changing food supply. 1976. Morphologie der Gelege und Gkologische Beobachtungen an Ecol. Monogr (in press): 5 figs. Muriciden (Gastropoda) aus der siidlichen Karibischen See. Verh. Tuorson, GUNNAR Naturf. Gesellsch. Basel 85: 1 - 32; 20 figs. 1940. Studies on the egg masses and larval development of arctic Castitza, J. C. & J. Cancrno prosobranchs. Medd. Grenland 100: 1-71 1976. Spawning behaviour and egg capsules of Concholepas concho- 1946. Reproduction and larval development of Danish marine bottom lepas (Mollusca: Gastropoda: Muricidae). Mar. Biol. $7; 255 - 263; invertebrates, with special reference to the planktonic larvae in the 7 figs. Sound (Gresund). Medd. Komm. Danm. Fisk. Havundersgg., Cgrnonorsxy, WALTER OLIver Copenhagen, Ser. Plankton 4 (1): 1-523; 199 figs. 1965. The radula, egg capsules and young of Murex (Chicoreus) torre- 1950. Reproductive and larval ecology of marine bottom invertebrates. factus Sowerby (Mollusca : Gastropoda). The Veliger 8 (4): Biol. Rev. 25 (1): 1-45; 6 text figs. (January 1950) “231-233; 6 text figs. (1 April 1965) 1952. Zur jetzigen Lage der marinen Bodentiere. — Okologie. Conk, E. G. Verh. deutsch. Zool. Gesellsch. 1951; Zool. Anz., Suppl. 16: 276 - 327 1897. The embryology of Crepidula. Journ. Morphol. 13: 1 - 226 Zeitier, W. ; 1898. Environmental and sexual dimorphism in Crepidula. Proc. 1974. Tropical marine invertebrates of southern Florida and the Acad. Nat. Sci. Philadelphia 1898; 435 - 444 Bahama Islands. John Wiley & Sons, New York, 132 pp.; 244 photos. Page 260 THE VELIGER Vol. 20; No. 3 The Systematic, Adaptive and Physiological Significance of Proteolytic Enzyme Distribution in Bivalves ROBERT G. B. REID Department of Biology, University of Victoria, Victoria, British Columbia, Canada INTRODUCTION THE DISTRIBUTION OF digestive enzymes in animals is believed to be adaptive, and related to the constituents of the diet (YoNGE, 1937). Most bivalves have a low pro- tein diet. Consequently, high levels of proteolysis would not be expected, and there has been relatively little inter- est in protein digestion in this class of molluscs. YONGE (1926) was satisfied that in Ostrea edulis Linnaeus, 1758 protein digestion was an intracellular process in the amoe- bocytes and digestive cells of the digestive diverticula, and he contended that gastric protein digestion would be in- compatible with proteinaceous crystalline styles (YONGE, 1930). Therefore some controversy was aroused when Mansour (1946) and Mansour & ZAKI (1946) argued that the digestive diverticula of bivalves were secretory and that animal food could be digested in the stomach. However, the low levels of protein digestion in the stom- achs of Tridacna elongata Lamarck, 1819, and Pinctada vulgaris Linnaeus, 1758 detected by MaANsour-BEK (1946) seemed insufficient to justify Mansour’s claim. BALLANTINE & Morton (1956) noted the presence of gastric proteinases in Lasaea rubra (Montagu, 1803), and suggested that they were released from “excretory spheres” from the digestive diverticula. OwEN (1956) examined Nucula sulcata Bronn, 1831 and found signifi- cant gastric protein digestion, but no diverticular activity. Differences in the distribution of proteolytic enzymes of two herbivorous, suspension-feeding bivalves, Lima hians (Gmelin, 1791) and Mya arenaria Linnaeus, 1758 indi- cated an adaptive relationship between enzyme distribu- tion and gastric morphology (Rem, 1966). This led to an investigation of animals belonging to different taxa pro- posed by PurcHon (1963) on the basis of stomach mor- phology. This study (Rem, 1968) provided some support for the original hypothesis concerning the relationship between stomach form and digestion. KozLovSKAYA & Vaskovsky (1970) surveyed a variety of marine inver- tebrates, including 14 species of bivalves, for alkaline proteases. For the bivalves only the digestive diverticula were assayed. A range of results from zero activity to low activity were obtained for the bivalves, very low in com- parison to carnivorous invertebrates. Since the survey dealt only with intracellular conditions it shed no light on the relationship between gastric morphology and digestion. However, a study of 7 species of Macoma (Tellinacea) by Rem & RaAucHERT (1972) indicated that the suggested relationship between gastric morphology and protein digestion might be spurious. Within the genus Macoma a range of proteolytic levels was noted, though all the species have similar stomach structures. This prob- lem was compounded when B. Morton (1970, 1973) proposed that bivalves undergo rhythmic and possibly endogenous cycles of ingestion, secretion, digestion and absorption. Such cycles might involve proteases, and thus account for differences observed within the genus Macoma. In carnivorous septibranchs there is high proteolytic activity which is related to the high-protein animal food (Rep, 1977). SYNOPSIS or PREVIOUS OBSERVATIONS To aid the comprehension of differences and similarities in protein digestion in bivalves, some of the relevant data are summarized below. The animals are grouped by family and gastric morphological type (PURCHON, 1956, 1957, 1958, 1960). Data are given in the following se- quence: habit; level of gastric proteolysis; gastric en- zymes if known; pH optima of gastric juice level of diver- ticular proteolysis; pH optima of diverticular extracts; diverticular enzymes if known. The stomach and the digestive diverticula are the main sites of protein di- gestion (Rem, 1966). Gastric proteolytic activity is expressed in units per mL of stomach fluid and di- verticular activity in units per mg fat-free dry weight of tissue. These are spectrophotometric units, each equi- valent to an absorbance change of 0.001 per minute at 280nm at 37° C. Absorbance increase is due to the release of amino acids as a result of proteolysis. Free amino acids present in extracts before enzyme assay are measured and Vol. 20; No. 3 THE VELIGER Page 261 deducted from final assay results. Soluble proteins which would also affect absorbance readings are removed by trichloracetic acid precipitation. Fuller technical details may be found in accounts by Rem (1966) and Rem « RAUCHERT (1970, 1972, 1976). In the following synopsis “low” gastric activity signifies 0-100 units per mL; “intermediate” gastric activity signifies 100 - 300 units per mL; “high” gastric activity signifies more than 300 units per mL. “Low” diverticular activity signifies o - 50 units per mg; “intermediate” diverticular activity signifies 50-150 units per mg; “high” diverticular activity signi- fies more than 150 units per mg. Family Nucutwae, Stomach Type I Acila castrensis (Hinds, 1843): deposit-feeder; inter- mediate gastric proteolysis; peaks at pH 3, 6, 7.5; diver- ticular condition unknown. (unpublished observations of author) Family NucuLtanmaeg, Stomach Type I Yoldia thraciaeformis Storer, 1838: deposit-feeder, inter- mediate gastric proteolysis; peaks at pH 3.5, 5.5, 7-5; diverticular condition unknown (RED, 1977). Family Cuspmarmpag, Stomach Type II Cardiomya planetica Dall, 1908: carnivorous; high gast- ric proteolysis; peaks at pH 3, 6, 7.5; enzymes cathepsin B and trypsin; diverticular condition unknown (RED, 1977). Family Mytitmag, Stomach Type III Mytilus californianus Conrad, 1837; suspension-feeder; low gastric proteolysis; peaks at pH 6, 7.5; trypsin; low diverticular proteolysis; peaks at pH 3, 6, 8 trypsin (un- published observations of author). Family Ostremar, Stomach Type III Crassostrea gigas (Thunberg, 1793): suspension-feeder; low gastric proteolysis; intermediate diverticular proteo- lysis; peaks at pH 3, 6, 7.5 (unpublished observations of author). Family Pectinmar, Stomach Type IV Chlamys hericius (Gould, 1850) : suspension-feeder; low gastric proteolysis; trypsin; low diverticular proteolysis; peaks at pH 2.5, 6, 7.5; chymotrypsin (Rem & RAUCHERT, 1970). Family Limmae, Stomach Type IV Lima hians (Gmelin, 1791) : suspension-feeder; low gast- ric and diverticular proteolysis; diverticular peaks at pH 5.5 and 8 (Rem, 1966). Family Unionwag, Stomach Type IV Anodonta kennerlyi Lea, 1860: suspension-feeder ; tryptic gastric and chymotryptic diverticular enzymes (unpub- lished observations of author). Family VENERIDAE, Stomach Type V Saxidomus giganteus Deshayes, 1839: suspension-feeder; low gastric proteolysis; peak at pH 5.2; low to intermedi- ate diverticular activity; peaks at pH 3, 6, 8; trypsin (Rew, 1977 and unpublished observations). Family Mactrwar, Stomach Type V Tresus capax (Gould, 1850) : suspension-feeder ; low gast- ric proteolysis; peaks at pH 2, 5, 8; gastric cathepsin B and trypsin; intermediate diverticular proteolysis; peaks at pH 2.5, 6, 8; diverticular cathepsin B and D and chy- motrypsin (Rem & RAUCHERT, 1976). Family TELLINmAE, Stomach Type V Macoma secta (Conrad, 1837): large particle deposit- feeder; intermediate gastric proteolysis; peak at pH 6; gastric trypsin; intermediate diverticular proteolysis; peaks at 2.5, 5.5, 8; diverticular chymotrypsin (RED & RAUCHERT, 1972). Macoma inquinata (Deshayes, 1854), suspension-feeder; intermediate gastric proteolysis; peaks at pH 5.5, 7; inter- mediate to high diverticular proteolysis; peaks at pH 3, 5, 7-5 (Rem & RAUCHERT, 1972). Macoma lipara Dall, 1916: deposit-feeder; low gastric proteolysis; peaks at pH 7; low to intermediate diverticu- lar proteolysis (RED & RAUCHERT, 1972). Family Mymae, Stomach Type V Mya arenaria Linnaeus, 1758; suspension-feeder; inter- mediate gastric proteolysis; peaks at pH 5.5, 7.2; cath- epsin B and trypsin; intermediate diverticular activity; peaks at pH 2.5, 3.5, 5.5, 8; diverticular cathepsin B, tryp- sin (RosEN, 1949; KozLovsKAyYA & VASKOVSKY, 1970; Rem, 1966 and unpublished observations) . Before discussion of these data some qualifications must be noted: in some of the earlier works only qualitative Page 262 THE VELIGER Vol. 20; No. 3 observations were made. In most cases, enzyme assays are incomplete; consequently, the identification of certain en- zymes in some examples does not imply that those enzymes are absent from other animals investigated. The studies on Chlamys hericius and Tresus capax indicated the pres- ence of carboxypeptidases and aminopeptidase in both the acid and alkaline pH range, as well as possible cathep- sin A and cathepsin C activity. These exopeptidases are probably of universal distribution. The results of Koz- LOVSKAYA & VASKOvSKy (1970) are largely excluded from the above due to the different expressions of unit activity employed by them. It is of note that 7 of their 14 bivalve species showed no alkaline endopeptidase activity. Mya arenaria showed the most activity. In addition to the data summarized above, I have made observations concerning variations in proteolytic activity within spe- cies. In Macoma secta, samples taken in May for 2 con- secutive years were found to have significantly different proteolytic levels, and gastric protein digestion in Maco- ma inquinata was found to vary significantly (RED & RAUCHERT, 1972). Further observations on variations in Tresus capax are presented in the following report. PRELIMINARY OBSERVATIONS on THE DIGESTIVE PHYSIOLOGY oF Tresus capax (Gould, 1850) IN RELATION to TIDAL RHYTHMS As noted above in the introduction, considerable interest has been generated in rhythmic physiological activity in bivalve alimentary tracts, by the work of Morton (1970, 1973). Gastric protein digestion in Macoma inquinata was found to vary significantly, and these variations were attributed to feeding behaviour, relative to tidal condi- tions (Rem & RAUCHERT, 1972). Proteolytic levels and the relative activities of particular proteases would be direct indicators of rhythmicity. Accordingly a pilot study of Tresus capax was carried out, to investigate gast- ric volume, gastric pH and proteolytic activity, and di- gestive diverticular activity. Tresus capax was chosen for its large size and because its proteolytic enzymes are better characterized than those of other bivalves (RED & RAUCHERT, 1976). Specimens were obtained from the intertidal region at Cherry Point, in the vicinity of Satellite Channel, British Columbia. This is a locality with a mainly diurnal tide; 7. e., where at the spring tidal period the animals in their natural habitat are uncovered once in 24 hours. Specimens of similar shell size were placed in wire baskets and re- buried in their natural habitat. The baskets were buoyed, so that they could be collected by boat at high water. After a 48 acclimatization period, during which it was verified that animals were able to use their siphons effec- tively, sampling began, was carried out at 3 hour inter- vals for 24 hours. During the 24 hours of sampling the experimental animals were exposed by the tide for 6 hours and submerged for 18 hours. Gastric pH in the samples was found to range from pH 6.2 to 7.2. Exposed animals were in the lower part of the range, submerged animals in the higher part of the range. Gastric volume was lowest (1 mL) in exposed animals, and highest (up to 8 mL) in animals which had been submerged for 8 hours or more. Gastric proteolysis was low, but constant in all samples. Diverticular proteo- lysis appeared to be low in exposed animals and inter- mediate in submerged animals. Sample sizes were too small to provide statistically reliable conclusions beyond the above. There did not appear to be any drop in gast- ric volume and pH during the 18 hour submergence, which might have corresponded with the minor ebb, which did not expose the specimens. These observations are discussed under the heading “Protein digestion and rhythmic physiological events in bivalves” in the following general discussion. DISCUSSION General Features of Protein Digestion in Bivalves In spite of the cursory nature of some of the observations reviewed above, it may be concluded that all bivalves are capable of digesting protein. Though comprehensive en- zymological studies are few it may be inferred that most bivalves have a complement of most of the proteases which are found throughout the animal kingdom. Partic- ular categories of proteases cannot be indentified with par- ticular habits, gastric morphology, nor systematic status. In general, the pH optima of gastric enzymes correspond with the pH of the stomach, 7.e., in the pH 6 range (Morton, 1970; MaTHERS, 1973; RED, 1977). In the digestive diverticula low pH (2 - 3.5), intermediate pH (4-6.5) and high pH (7-8) activity peaks occur uni- versally. In Tresus capax the enzymes cathepsin D, cathep- sin B and chymotrypsin are active in the low, intermediate and high pH regions respectively. This may hold true for other bivalves. Protein Digestion and Gastric Morphology It might be generalized that the stomach types such as I and II, which are structurally the simplest, are the ones which have the highest levels of proteolytic activity (e. g., Vol. 20; No. 3 Cardiomya, Yoldia). It does not follow that there is an adaptive relationship, since it has been shown that there are significant differences in proteolytic levels within the type V group. The simplicity of the type I and II stom- achs is probably more directly related to the triturative process than to the enzymatic process. No definitive adap- tive relationship between protein digestion and gastric morphology can be stated at present. Protein Digestion and Habit The suspension-feeding bivalves which have been studied exhibit a range of proteolytic activity from low to inter- mediate in terms of the present study. The question of what constitutes the diet of suspension- and deposit-feed- ers has been discussed by Ponto (1969) and Rem (1971). The protein content of the phytoplankton food of suspen- sion feeders is variable, but does not exceed 30% by dry weight (STRICKLAND, 1965). It is generally believed that deposit-feeders depend largely on the microorganisms as- sociated with the detrital and inorganic particulate mate- rial of their diet (NEWELL, 1965). That the protein con- tent of the food of deposit-feeders would warrant differ- ent levels of proteolytic enzymes from those found in sus- pension-feeders is debatable. However, it may be that gastric proteinases in some deposit-feeders play a signif- icant role in separating potential food particles from substrate particles which have no food value, prior to endocytosis, as was suggested for Macoma secta (RED & RauCHERT, 1972). While this may also be the case for the deposit-feeding protobranchs it does not apply to other deposit-feeding members of the genus Macoma. Only in the carnivorous septibranchs is there a positive relationship between diet and protein digestion. The family Cuspidariidae is the only group so far investigated which shows high levels of proteolysis. The diet consists of small copepods, ostracods, chaetognaths, polychaetes, etc. (PELSENEER, 1891; YONGE, 1928; KNUDSEN, 1970; BERNARD, 1974), and the food is taken live (REM « REM, 1974). The major gastric proteolytic enzymes are trypsin and cathepsin B (Rem, 1977); 7.¢., the same enzymes which are found in traces of the stomach of other bivalves. Thus, only a minor enzymological adaptation is involved, namely an increase in the synthesis and secretion rates, achievable by a simple gene duplication. In Cardiomya and Cuspidaria species the crystalline style is truncated, suggesting some attack by the gastric enzymes. THE VELIGER Page 263 The Systematic Significance of Proteolytic Enzymes There are a number of ways in which research into proteo- lytic enzymes might reveal their systematic significance. The first is the study of the primary structure of proteases of a given category. Although this would provide useful phylogenetic information about the bivalves and their affinities with other molluscs this expensive and time-con- suming technique has not been applied. The second ap- proach is the comparative study of protease zymograms; 1.é@., the visualized patterns of electrophoretically sepa- rated enzyme extracts. The applicability of electrophoretic data to zoological systematics has been evaluated by AvisE (1974). While a number of enzyme categories have been tested for their genetic or general systematic significance in bivalves (e. g., KOEHN & MitTTon, 1972), and some of these studies have included categories of proteolytic en- zyme (LEVINTON, 1975; LEVINTON & FUNDILLER, 1975) the information to date is insufficient to suggest clear systematic relationships. As AvisE (op. cit.) points out, there are two schools of thought concerning the evolu- tionary significance of allozymes: the first is the selectionist view that each molecule hasbeen tested by natural selection and is of positive value to the organism or population; the other view being the neutralist one which claims that some allozymatic variations have no particular superiori- ty to others, but, since they are no less valuable, they re- main as neutral mutations. Malacologists generally take the traditional selectionist view. However, if there is any truth in the neutralist view, then there is a possibility that some enzymological phenomena have no adaptive significance, but nevertheless have systematic significance. For example, in some members of the Pectinacea there is relatively high activity in the electrophoretically fast-run- ing chymotrypsin. This might be a systematic phenom- enon which lacks adaptive significance. Protein Digestion and Rhythmic Physiological Events in Bivalves J. E. Morton (1956) outlined phases of alimentary activity in Lasaea rubra. These were absorptive, intra- cellular digestive and excretory phases. The release of gastric proteinases was believed to be part of the ex- cretory phase. B.S. Morton (1970) observed similar phases in Cardium edule, and inferred that these were digestive rhythms which correspond to tidal rhythms. He proposed that such rhythmic digestive phenomena were universal among bivalves, even those unaffected by the Page 264 direct influence of the tide (Morton, 1973). LANGTON « GaspoTt (1974) noted rhythmic changes in style pH, style protein, and amylase activity in Ostrea edulis, and concluded that these changes were modulated by the tide. However, they noted that the rhythms were lost following a period of continuous immersion. The pertinent literature has been more extensively reviewed by OWEN (1974). In my study of Tresus capax it was observed that gast- ric volume was at its lowest while animals were exposed and highest about 8 hours after submergence by the rising tide. Gastric pH was lowest in exposed animals and high- est in submerged animals. The minor ebb, which did not expose the animals, had no obvious physiological effect. It is particularly significant that the gastric proteolytic levels remained constant. This indicates an active release of enzymes as the animal feeds and the gastric volume increases. Otherwise the proteolytic activity would de- crease with dilution. The gastric pH changes are similar to those observed by LancTon & Gaszott (1974) for oysters. The sequence of events outlined above may accord with Morton’s hypo- theses concerning rhythmic digestive phenomena in bi- valves (MorToN, 1973). However, the question of whether these events are endogenous, exogenous and modulated by the tide, or possibly synchronized by experimental arti- fact, remains open. The Evolution of Protein Digestion in Bivalves The range of categories of proteolytic enzymes present in bivalves is no different from that found in other inverte- brates. In animals which have been closely examined most of the known invertebrate proteases have been found. The evolution of the endopeptidases is known to have been highly conservative in biochemical terms, and the view that animals, as they undergo adaptive radia- tion, encounter new diets and consequently evolve “new” digestive enzymes is fallacious with regard to the prote- ases. The range of peptide groupings in the most primitive protein substrate is the same as in the most highly evolved protein. Consequently, in the course of evolution, the need for a specifically novel category of protease is rarely necessary, though changes in the physiological en- vironment might favour variants of pre-existing categories of enzyme. While qualitative changes have not occurred in enzyme category, some innovative uses have been made by the bivalves of pre-existing enzymes, particularly in the use of cathepsin B for digestive purposes. In most organisms the lysosomes have a cathepsin complement responsible for autolysis, autophagy and general protein turnover. As indicated by seasonal changes in the relative levels of THE VELIGER Vol. 20; No. 3 cathepsin in Tresus capax (REM & RAUCHERT, 1976), cathepsin B has been turned to a digestive role and in the carnivorous septibranchs is employed for extracellular gastric digestion. A large quantitative change has been made by the septibranchs in response to their high pro- tein diet. If the septibranchs were regarded as a primitive group of bivalves, and considering the protobranch con- dition, it might be concluded that the relatively high proteolytic capacity of the stomach is characteristic of primitive bivalves. However, considering the relative sim- plicity of the quantitative evolutionary step and the ab- sence of qualitative differences it is impossible to justify such a conclusion, and, in general, in the absence of in- formation on primary structure, proteases and the process of protein digestion are poor indicators of the direction of bivalve evolution. SUMMARY 1. There are no systematically significant qualitative dif ferences in the distribution of proteolytic enzymes in bivalves. Members of the Pectinacea have prominent, electrophoretically fast-running chymotryptic en- zymes which may be of quantitative systematic signifi- cance. Electrophoretic zymograms of proteases may prove to be useful in determining systematic relation- ships. 2. No definitive adaptive relationships can be traced be- tween protein digestion and gastric morphology. Her- bivorous suspension-feeding bivalves have low proteo- lytic capacities. Carnivorous septibranchs have high proteolytic capacities. Deposit-feeding bivalves range from low to intermediate proteolytic capacities. The adaptive significance of the latter case is not clear. 3. Gastric volume, gastric pH, gastric protease secretion and digestive diverticular proteolytic activity appear to vary in relation to tidal events. It is not known if these are exogenous or endogenous rhythms. 4. The evolution of protein digestion in bivalves has been biochemically conservative. Proteases found in bivalves occur universally in the animal kingdom. Since quan- titative changes are relatively simple evolutionary events, and since nothing is known of the primary structure of bivalve proteases, proteases and the process of protein digestion are poor indicators of phyloge- netic relationships. Literature Cited Avisz, Joun C. 1974. Systematic value of electrophoretic data. 465 - 481 Syst. Zool. 23: (September 1974) Vol. 20; No. 3 THE VELIGER Page 265 BALLenTINE, DortHy & JoHN Epwarp Morton 1956. Filtering, feeding, and digestion in the lamellibranch Lasaea rubra. Journ. mar. Biol. Assoc. U. K. 35: 241 - 274 BerNnarp, Frank R. 1974. Septibranchs of the Eastern Pacific (Bivalvia Anomalodesmata). Allan Hancock Monogr. Mar. Biol. 8: 1 - 279 (28 February 1974) KNnupseEn, Jorcen 1970. The systematics and biology of abyssal and hadal Bivalvia. Galathea Rprt. 11: 241 pp.; 20 plts.; 132 text figs. (6 Nov. 1970) Koeun, Ricwarp K. « J. B. Mitton 1972. Population genetics of marine pelecypoda. I. Ecological hetero- geneity and evolutionary strategy at an enzyme locus. Amer. Natur. 106: 47 - 56 Koztovsakay, E. P « V. E. VAsKovsKy 1970. A comparative study of proteinases of marine invertebrates. Comp. Biochem. Physiol. 34: 137 - 142 LancTon, Ricuarp W. « P. A. GasBotT 1974. The tidal rhythm of extracellular digestion, and the response to feeding in Ostrea edulis. Marine Biol. 24: 181 - 187 LEvINTON, JEFFREY S. 1975. Levels of genetic polymorphism at two enzyme encoding loci in eight species of the genus Macoma (Mollusca: Bivalvia). Marine Biol. 33: 41 - 47 LEvINTON, JEFFREY S. & Doreen L. FuNDILLER 1975. An ecological and physiological approach to the study of bio- chemical polymorphisms. Proc. 9th Europ. mar. biol. Symp. pp. 165 - 178 Mansour, K. 1946. Food and digestive processes of the lamellibranchs. 157: 482 Mansour, K. « EG. Zax1 1946. The digestive diverticula of Unio prasidens as organs of secretion. Proc. Egypt. Acad. Sci. 2: 38 - 44 Mansour-Bex, J. J. 1946. The digestive enzymes of Tridacna elongata Lam. and Pinctada vulgaris L. Proc. Egypt. Acad. Sci. 1: 13 - 20 Matuers, Nicer F 1974. Digestion and pH variation in two species of oysters. Proc. Malacol. Soc. London 41: 37 - 40 Morton, Brian S. 1970. The tidal rhythm and rhythm of feeding and digestion in Cardi- um edule. Journ. mar. Biol. Assoc. U. K. 50: 499 - 512 1973. A new theory of feeding and digestion in the filter-feeding La- mellibranchia. Malacologia 14: 63 - 79 (23 January 1973) Morton, JoHN EDWARD 1956. The tidal rhythm and action of the digestive system of the lamelli- branch Lasaea rubra. Journ mar. Biol. Assoc. U. K. 35: 583 - 586 Owen, GarETH 1956. Observations on the stomach and digestive diverticula of the Lamellibranchia. II. The Nuculidae. Quart. Journ. micr. Sci. 97: 541 - 567 1974. Feeding and digestion in the Bivalvia. Biochem. 5: 1-35 PevsenzEr, Paut 1891. Contribution a l'étude des lamellibranches. Arch. Biol. 11: 147 - 312 (August 1891) (15 January 1970) Nature (13 April 1946) Adv. Comp. Physiol. Poxto, Ross H. 1969. Confusion concerning deposit-feeding in the Tellinacea. Proc. malacol. Soc. London 38: 361 - 364 PurcHon, RicHarD DENISON 1956. The stomach in the Protobranchia and Septibranchia (Lamelli- branchia). Proc. Zool. Soc. London 127: 511-525 (Dec. 1956) 1957. The stomach in the Filibranchia and Pseudolamellibranchia. Proc. Zool. Soc. London 129: 27 - 60 (September 1957) 1958. The stomach in the Eulamellibranchia; Stomach Type IV. Proc. Zool. Soc. London 135: 431 - 489 (December 1958) 1960. The stomach in the Eulamellibranchia; stomach types IV and V. Proc. Zool. Soc. London 135 (3): 431-489; 19 text figs. (20 October 1960) 1963. Phylogenetic classification of the Bivalvia, with special regard to the Septibranchia. Proc. malacol. Soc. London 35: 71 - 80 1971. Digestion in the filter-feeding bivalves — a new concept. Proc. malacol. Soc. London 39: 253 - 262 Rew, Rosert G. B. 1966. Digestive tract enzymes in the bivalves Lima hians Gmelin, and Mya arenaria L. Comp. Biochem. Physiol. 17: 417 - 433 1968. The distribution of digestive tract enzymes in lamellibranchiate bivalves. Comp. Biochem. Physiol. 24:727-744 (15 Sept. 1968) 1971. Criteria for categorizing feeding types in bivalves. The Veliger 13 (4): 358-359 (1 April 1971) 1977. Gastric protein digestion in the carnivorous septibranch Cardio- mya planetica Dall; with comparative notes on deposit and suspension- feeding bivalves. Comp. Biochem. Physiol. 56A: 573 - 575 (1 January 1977) Rem, Rosert G. B. e Katuy RAUCHERT 1970. Proteolytic enzymes in the bivalve mollusc Chlamys hericius Gould. Comp. Biochem. Physiol 35: 689 - 695 (15 July 1970) 1972. Protein digestion in members of the genus Macoma (Mollusca: Bivalvia). Comp. Biochem. Physiol. 41A: 887 - 895 (1 Dec. 1972) 1976. Catheptic endopeptidases and protein digestion in the horse clam Tresus capax (Gould). Comp. Biochem. Physiol. 54B: 467 to 472 (15 December 1976) Rem, Rosert G. B. # ALison REID 1969. Feeding processes in members of the genus Macoma (Mollusca: Bivalvia). Canad. Journ. Zool. 47: 649-657. (November 1969) 1974. The carnivorous habit of members of the septibranch genus Cuspidaria (Mollusca: Bivalvia). Sarsia 56: 47-56 (8 July 1974) Rostn, Broer 1949. Proteases in the digestive gland of lamellibranchs. Archiv Demi. 1: 205 - 211 StrIcKLAND, Joun D. H. 1965. Production of organic matter in the primary steps of the marine food chain. In: Chemical Oceanography 1: 477 - 610. Acad. Press, London, New York Yonez, Cuartes Maurice 1928. Structure and function of the organs of feeding and digestion in the septibranchs, Cuspidaria and Poromya. Phil. Trans. Roy. Soc. Ser. B. 26: 221 - 263 (30 June 1927) 1930. The crystalline style of the Mollusca and a carnivorous habit cannot normally co-exist. Nature 125 (3151): 444-445 (22 March 1930) 1937. Evolution and adaptation in the digestive system of the Metazoa. Biol. Revs. Cambridge Phil. Soc. 12: 87-115 Page 266 THE VELIGER Vol. 20; No. 3 Reproductive Biology of Colus stimpsoni (Prosobranchia : Buccinidae) I. Male Genital System ' BY DAVID L. WEST? (5 Plates; 2 Text figures) INTRODUCTION Tue Buccinmaer is A large family, generally occurring in temperate latitudes, and many of its members undergo larval development within the egg capsules, emerging as juveniles (RADWIN & CHAMBERLIN, 1973; PONDER, 1974). A few studies have dealt with the reproductive biology of this family (Dakin, 1912; FRETTER, 1941; Houston, 1976) and the larval stages and egg capsules have been described for a few species (PoRTMANN, 1926, 1927; THORSON, 1935, 1940; FRETTER & GRAHAM, 1962; WEST, 1973)- Previous studies on the reproductive system have indi- cated a similarity in the organization of these structures (DAKIN, 1912; FRETTER, 1941; FRETTER & GRAHAM, 1962; Houston, 1976), but these studies mainly concern the female genital system. The male system, particularly in the genus Colus, has been neglected. It is of value to study the male genital system in the genus Colus for comparative functional and phylogenetic relationships. The present study concerns the male reproductive sys- tem in Colus stimpsoni (Morch, 1867). MATERIALS anp METHODS Numbers of Colus stimpsoni were collected at Cobscook State Park, Edmunds, Maine, and at Eastport, Maine, and maintained in running seawater aquaria at the Marine Science Institute, Northeastern University, Nahant, Mas- sachusetts. In the laboratory, C. stimpsoni was supplied Littorina littorea (Linnaeus, 1758) and, occasionally, Polinices triseriata (Say, 1826), Lunatia heros (Say, 1822), and Thats lapillus (Linnaeus, 1758) as food items (RIsER, 1969; WesT, 1973). * Contribution No. 56 from the Marine Science Institute, North- eastern University, Nahant, Massachussetts * Present address: Center for Pathobiology, University of Cali- fornia, Irvine, California 92717 For light microscope studies, tissues were excised from freshly opened, unrelaxed animals and fixed for 1-24 hours in the following: (a) Hollande-Bouin’s, (b) Bou- in’s, (c) alcohol-formalin-acetic acid, (d) buffered for- malin (pH 7.4), (e) seawater-Bouin’s (WALKER & Mac- Grecor, 1968), or (f) buffered glutaraldehyde (pH 7.4). Following fixation, tissues were dehydrated through a graded series of ethanol and embedded in polyester wax (STEEDMAN, 1960). Sections were cut 3 - 10 um in thick- hess and mounted on albuminized slides. Sections were stained with the following: (a) Heidenhain’s iron hem- atoxylin, (b) Pollack’s rapid trichrome, (c) Heidenhain’s azan, or (d) GaBe’s (1968) modification of Gomori’s tri- chrome. LEHMAN’s (1965) polychrome was used for lo- calization of various molecular groups. For cytochemical tests, tissues were excised from freshly opened, unrelaxed animals and fixed at either 5 - 6° C or room temperature in the following: (a) buffered for- malin (pH 7.4), (b) Bouin’s or (c) buffered glutaralde- hyde (pH 7.4). Following fixation, the tissues were de- hydrated in a graded series of ethanol or acetone and embedded in polyester wax or in paraffin (45°C M. P). Sections were cut 4 - 6m in thickness and mounted on unalbuminized slides. The wax was removed from the sections with absolute acetone and the sections were air dried. The periodic-acid-Schiff (PAS) technique or Best’s carmine was used to test for the presence of polysaccha- rides. The presence of glycogen in PAS positive sections was determined with salivary amylase controls incubated at 37° C for 2 hours. To test for acid mucopolysaccharides, the alcian blue (pH 1.0 and 2.3) or colloidal iron tech- niques were employed. The acid haematein technique was used to test for the presence of phospholipids. Localiza- tion of DNA and RNA was by the Feulgen reaction or by the methyl green-pyronin Y method. Protein groups were stamed by LEHMan’s (1965) polychrome method or by mercuric bromphenol blue. Acid and alkaline phosphat- ases were localized by Gomori’s lead nitrate technique (Humason, 1967) or by theazo-coupling method (PEARSE, Vol. 20; No. 3 1968). Localization of glucose-6-phosphatase was by the azo-coupling method (PrarsE, op. cit.). No difference in enzyme activity was found between tissues fixed in buf fered glutaraldehyde or buffered formalin, nor was en- zyme activity inhibited by embedding in polyester wax. For electron microscopy, tissues were excised from freshly opened, unrelaxed animals, and fixed for 1 hour in cold (5-6°C) primary fixative. The primary fixative was prepared just before use and contained: 3% glutaral- dehyde, 1% formaldehyde, made from paraformaldehyde (Karnovsky, 1965), 0.1 M phosphate buffer (pH 7.4), 2-3% NaCl, and 2-4% sucrose. Dimethylsulfoxide (0.2%) was added to aid penetration. Following primary fixation, the tissue was washed for 1 hour in cold 0.1 M phosphate buffer (pH 7.4) which contained 6-8% sucrose. The tissue was post-fixed at room temperature with 1% OsO, in 0.1 M phosphate buffer (pH 7.4). Fol- lowing post-fixation, the tissue was washed for $ hour in cold o.1 M phosphate buffer (pH 7.4) and dehydrated in a graded series of cold acetone or ethanol. The tissue was allowed to come to room temperature in 95% acetone or ethanol, and final dehydration was at room tempera- ture. The tissue was embedded in Epon or an Epon-Aral- dite mixture (BURKE & GEISELMAN, 1971; GEISELMAN & BurKE, 1973). Thin sections were double stained with aqueous uranyl acetate (Watson, 1958) and lead citrate (REYNOLDS, 1963), and examined with a Zeiss EM 9-S electron microscope. RESULTS General Morphology The testis is yellow to orange-yellow in color and spreads over the digestive gland in the ultimate and in part of the penultimate whorls of the shell (Figures 1a, 1b). In large males it may cover as much as } of the digestive gland. The seminiferous tubules are generally separated from the digestive gland but they occasionally intrude between the tubules of the latter. The seminiferous tubules join to form a single genital duct which passes along the columellar side of the visceral mass, across the right side of the body and exits through the penis. The penis is flat- tened laterally and issituated just behind the right tentacle (Figure 1a). It varies from 2 to 5cm in length and is directed posteriorly when not projected. The posterior portion of the genital duct, the vas defer- ens, lies superficially on the visceral mass. It is convoluted and the thin walls contain a few muscle fibers. The poste- rior 3 of the vas deferens are filled with spermatozoa throughout the year and form the seminal vesicle (Figures THE VELIGER Page 267 1a, tb). The anterior 4 is the renal vas deferens (Figure 1b) which is separated from the seminal vesicle by a small sphincter and straightens at the posterior extremity of the mantle cavity. It opens into the glandular pallial portion Figure 1 Male Colus stimpsoni: a-— whole animal with shell removed. b — dissection with mantle cut along right edge and reflected to the left side of the body; proboscis removed from its cavity and the floor of the cavity cut to expose the salivary gland and coiled prostate PR -— prostate RV - renal vas deferens SV — seminal vesicle TE -— testis PE — penis Sg - salivary gland Page 268 THE VELIGER Vol. 20; No. 3 of the genital duct. At the junction of the renal vas defer- ens and pallial portion, the duct makes an S-shaped turn and is slightly constricted (Figure 2). Near this constric- tion, at the posterior limit of the mantle cavity, the renal vas deferens gives off a small diverticulum which passes posteroventrally and opens into the mantle cavity (Figure 2). No functional gonopericardial duct nor any remnant of this duct is found in the male. The pallial portion of the genital duct, beginning near the posterior limits of the mantle cavity, is muscular and glandular along its entire length and functions as the pros- tate. The prostate passes just beneath the body wall to the base of the penis where it turns medially and passes into the body. Along the body, the prostate projects as a prom- inent ridge near the mantle floor (Figure 1b). Within the body, the prostate coils among the lobes of the right salivary gland (Figure 2). It passes outward through the body wall adjacent to its point of entrance and continues through the penis to open at the tip of the penis (Figure 2). Within the penis, the duct is centrally located. Histology Testis: ‘The testis and visceral mass are covered by a single layer of cuboidal epithelium, the pallial epithelium. Subjacent to the pallial epithelium is a thin basal lamina (0.15 -0.50um thick) and a layer of loose connective tissues and muscle fibers. Seminiferous tubules (Figure 3) end blindly at the surface of the testis and are generally oriented perpendicular to the spiral axis of the shell. The seminiferous tubulesmeasure 100-800 umin diameter and are delineated by a layer of connective tissue and a thin Figure 2 Male reproductive system: line posterior to diverticulum (D) represents the posterior limits of mantle cavity PE — penis PR - prostate RV -— renal vas deferens Sg — salivary gland SV — seminal vesicle TE -— testis (not drawn to scale) basal lamina. The basal lamina is composed of a compact layer of fibers and varies from 0.4 to 0.6 um in thickness. Beneath the connective tissue on the periphery of the seminiferous tubule, a layer of lipid-rich accessory (basal) cells surrounds the spermatogenic cells. These accessory cells are similar to the basal cells in Gontebasis laqueta (Say) described by Woopwarp (1935). The layer of . basal cells is irregular in thickness, varying from one small cell in starved animals to one large cell or 2 - 3 smaller cells in well-fed animals, and is occasionally interrupted by spermatogenic cells. Explanation of Figures 3 to 6 Figure 3: Cross section of testis showing seminiferous tubules Figure 4: Basal cell within seminiferous tubule. B — basal lamina; L - lipid; N — nucleus Figure 5: Type I accessory cell. I - inclusion; N — nucleus; R — rough endoplasmic reticulum Figure 6: Type II accessory cell. D — dense granules within nucleoplasm; N - nucleus I — inclusion; Explanation of Figures 7 to 11 Figure 7: Type II atcessory cell with fields of small tubules (*) and small vesicles with electron-dense granules (arrows). Figure 8: High magnification of small tubules (arrows) shown in Figure 7. Figure 9: High magnification of vesicles with electron-dense granules (arrows) shown in Figure 7. R — rough endoplasmic reticulum Figure 10: Type II accessory cell with flagella (arrows) and dense inclusion (1). Figure 11: Cross section through seminal vesicle (S) and renal vas deferens (V). THE VELIGER, Vol. 20, No. 3 [West] Figures 3 to 6 [West] Figures 7 to 11 Tue Veicer, Vol. 20, No. 3 Vol. 20; No. 3 THE VELIGER Page 269 seme The cytoplasm of the basal cells (Figure 4) contains glycogen granules and is dominated by lipid droplets which are dissolved during standard histological proce- dures but are retained with electron microscope tech- niques. Basal cells are irregular in shape, 10 - 25m in length, and contain few organelles. Mitochondria are cylindrical, measuring 2-3 m by 0.2 - 0.4m. Occasi- onally residual bodies are observed in the cytoplasm. The nucleus (7 - 15pm in length) is lobed and contains 1 or occasionally 2 nucleoli. The chromatin is distributed around the periphery of the nucleus and small clumps of chromatin are scattered throughout the nucleoplasm. The nuclear envelope is perforated by a few pores. The nucle- olus is heterogeneous with a light, granular medulla and a dense, granular cortex. Spermatogenic cells and accessory cells fill the semini- ferous tubules subjacent to the layer of basal cells. In sections stained with Heidenhain’s azan, accessory cells contain an intensely staining material and are easily dis- tinguished from spermatogenic cells. Two types of acces- sory cells can be distinguished by their nuclear morpholo- gy. They are here designated as type I and type II. The type I accessory cell has a single nucleus (Figure 5) ; whereas the type II cell is multinucleated (Figure 6). From squash preparations and sectioned material, both types of accessory cells appear to be polyploid. Accessory cells occur in groups (4 - 24 cells per group) within the tubules. Each group consists of one type of accessory cell and is usually near the periphery of a tubule. The groups are irregularly distributed along the length of a tubule and are not associated with any particular stage of sper- matogenesis. Generally, type II cells are more abundant than type I cells. The type I cell (Figure 5) is irregular in shape, meas- uring 7 - 12m in length and is mostly filled with mem- brane-limited inclusions which stain intensely red with Heidenhain’s azan. The nucleus measures 5 -8pm in diameter and the chromatin is condensed into large clumps which are distributed around the periphery of the nucleus and within the granular nucleoplasm. The nucle- us is lobed and is surrounded by the membrane-bounded inclusion. The small amount of cytoplasm contains numer- ous free ribosomes and a few small mitochondria. The cisternae of the rough endoplasmic reticulum are en- larged and filled with an electron-dense, homogeneous matrix. The contents of the cisternae accumulate and form a large inclusion which occupies nearly the entire cell volume. Type II cells contain an intensely staining material. However, the material stains lavender or blue with Hei- denhain’s azan. The type II cell (Figure 6) contains 2 relatively large nuclei (4-6m in diameter) and 3-6 smaller nuclei ( 1- 3 um in diameter). The chromatin is condensed into a thick patchwork with small amounts of granular nucleoplasm. The nucleoplasm contains numer- ous electron-dense granules which vary from 0.05 to 0.3 zm in diameter (Figure 6). Generally, the nuclei are em- bedded in the large inclusion. The cytoplasm contains numerous free ribosomes and the Golgi complex. The cisternae of the rough endoplasmic reticulum are discon- tinuous and are filled with a homogeneous electron-dense substance which is similar in appearance to the material in the inclusion. Localized areas of small tubules which contain a homogeneous material occur within the cyto- plasm (Figure 7). These short tubules measure between 0.06 and 0.15 um in diameter (Figure 8). The cytoplasm also contains membrane-bounded vesicles which contain electron-dense, rod-shaped granules (Figure 9). These vesicles measure 0.2 - 0.4 4m in diameter and the granules measure about 0.05 zm in diameter. Occasionally, type II cells have flagella which vary in number from 1 to 12 and also large, electron-dense inclusions which are similar to large residual bodies or pycnotic nuclei (Figure 10). These flagella and inclusions may be from ingested germ cells. Seminal Vesicle and Renal Vas Deferens: ‘The convo- luted posterior portion of the single genital duct is em- bedded in connective tissue and is separated from the digestive gland. The loops of the duct are generally per- pendicular to the columellar axis and these loops are separated from one another by loose connective tissue. The duct varies from 100 to 800m in diameter and is surrounded by a thin layer of muscle fibers (2-5 um thick) which are generally circularly arranged. The loops of the duct overlap one another such that portions of the renal vas deferens are found in the seminal vesicle region (Figure 17). The seminal vesicle and the renal vas deferens are lined with a columnar epithelium of varying height. In the seminal vesicle, the epithelial cells (Figure 72) rest on a substantial basal lamina (0.6-1.1 um thick). The epithelium varies between 10 and 20pm in thickness and is composed of 2 cell types. One type (ciliated cell) has long cilia and simple, occasionally branched, micro- villi on the apical surface (Figure 12). The second type (microvillar cell) has only long, anastomosing microvilli on the distal surface (Figure 12). The cytoplasm of both cell types contains numerous small mitochondria (about 0.34m in diameter) and several large residual bodies. Numerous Golgi lamellae are distributed throughout the cytoplasm. The microvillar cell contains numerous small, membrane-bounded vesicles which are filled with fine, electron-dense granules. The rough endoplasmic reticu- Page 270 lum of the microvillar cell is more abundant than that of the ciliated cell. These cisternae are discontinuous and filled with a homogeneous matrix. The epithelium of the renal vas deferens (Figure 13) is thicker (15 - 30m) than that of the seminal vesicle. The undulating basal lamina is also slightly thicker (0.9 to 1.2m). Microvillar cells are absent but the ciliated cells are similar to those of the seminal vesicle. The diverticulum (Figure 14), which branches from the renal vas deferens, is histologically similar to the renal vas deferens and is about 1mm long. This short duct is ciliated and opens into the mantle cavity. Prostate: The anterior portion of the genital duct passes along the body wall just beneath the epidermis and is histologically similar along its entire length. It is yellow- ish in color and functions as the prostate (Figure 15). The epithelium is 15 - 50m in height and is composed of ciliated columnar and secretory cells which rest on a thin basal lamina. The ciliated cells (Figure 76) of the epithelium extensively interdigitate with each other and their apical surfaces havenumerousciliaand pseudopodial extensions. The cytoplasm contains numerous rod-shaped mitochondria, and is less electron-dense than the neigh- boring secretory cells. The cytoplasm also contains large clumps of glycogen granules. The nuclei of ciliated cells are lobed and generally elongated in the long axis of the cells. The secretory cells of the epithelium appear similar in shape to “goblet” cells and are filled with numerous mem- brane-bounded secretory vesicles (Figure 17). These ve- sicles are irregular in outline and are filled with electron- dense granules. Lipid droplets abut against these gran- ules and, generally, are hemispherical in shape. The cyto- plasm of secretory cells is dominated by the secretory vesicles, but it also contains rough endoplasmic reticulum THE VELIGER Vol. 20; No. 3 and a well developed Golgi complex. The cisternae of the rough endoplasmic reticulum are filled with an electron- dense matrix. The nucleus is irregular in outline and generally contains 2 nucleoli. A layer of circular muscle (0.8 - 1.2 um in thickness) lies beneath the epithelium and is interrupted at intervals by secretory cells which are more prominent during peri- ods when males are reproductively active (Figure 15). The secretory cells are oriented radially within the muscle layer. However, there is no apparent orientation to the secretory cells on the periphery of the duct. The secretory cells are of 2 types and are here desig- nated as type 1 and type 2. Judging from electron micro- graphs, type 1 and type 2 secretory cells appear to under- go a maturation sequence. In early type 1 cells (Figure 18) the cytoplasm is filled with rough endoplasmic reti- culum and the Golgi complex. The cisternae of the rough endoplasmic reticulum are discontinuous and dilated. These cisternae are filled with a homogeneous matrix which is slightly less electron-dense than the surrounding cytoplasm. In the middle stage of type 1 cell maturation (Figure 78) the rough endoplasmic reticulum becomes vesiculated and the cisternae are filled with a dispersed granular material. The lamellae of the Golgi complex are elongated and flattened. Membrane-bounded vesicles which contain an electron-dense material appear in the cytoplasm. The mature type 1 secretory cell (Figure 19) contains numerous membrane-bounded vesicles. These vesicles contain a heterogeneous, electron-dense matrix which surrounds a small area of electron-dense granules. These small granules are clumped together and this clump is separated from the surrounding matrix by an elec- tron-lucent area. Microtubules occupy much of the re- maining cytoplasm of mature type 1 cells and are aligned in the long axis of the cell. Explanation of Figures 12, 13 Figure 12: Epithelium of seminal vesicle. C - ciliated cells; M - microvillar cells Figure 13: Epithelium of renal vas deferens. Explanation of Figures 14 to 17 Figure 14: Cross section through posterior region of mantle cavity (M) and diverticulum (D) V — renal vas deferens Figure 15: Cross section through prostate. B — body wall; arrows — secretory regions Figure 16: Ciliated cell of prostate epithelium. Figure 17: Epithelium of prostate. arrows — interdigitating cells arrow — lipid droplet; g — clump of glycogen particles within ciliated cell; S — secretory cell Tue VE uicER, Vol. 20, No. 3 [West] Figures 12, 13 > fs 2 = > ' ; i at a =i in a 7 ; ~ ee - io, ed se i ) = mings - = = Ge 2 R r re ‘ = : ee {7 2 > . Pi z \ o 5 = : = sa se (S: i : - , rt “y < : i =| e = ne ie ae F “e wi ie : <) : ‘ a ; C = = i é ; = = 5 =F 3 ie i ; 5 a 2 : a } : = oy = ~ M m , Bs ty A a % = i : : ~ : : can , oi 3 ¢ > ; 5 f i y " is Ps i ; iy * » 4 ~ ” 7 t = ne Lot ? 4 >. 3 Pam 4 fi ‘ Y = : G 3 : ; 7 =e = " ~~ ~ “ 72> ; ? aa = > ed a t tm ws } ' 7 J > — = : = : . i x \ ; S i « i ; : ———— ee ee = ————————————————————————S Se DA Na a > shee —— oe Tue VELIGER, Vol. 20, No. 3 OF ete J [West] Figures 14 to 17 Vol. 20; No. 3 THE VELIGER Page 271 Type 2 cells also appear to undergo a maturation se- quence similar to type 1 cells. Young type 2 cells (Figure 20) are filled with small membrane-bounded vesicles which contain a granular material. The mitochondria are rod-shaped and small. Elements of the Golgi complex are scattered among the vesicles. Mature type 2 cells (Figure 27) are filled with larger vesicles which contain a flocculum and a few vesicles containing a heterogeneous material. The Golgi complex of older type 2 cells is com- posed of flattened lamellae and the cisternae are filled with an electron-dense material and small vesicles are associated with the tips of the Golgi lamellae. Cytochemistry Testis: Results from polyester wax sections stained with Lehman’s polychrome indicate the specific locali- zation of a number of macromolecular groups. The differ- ential staining suggests the following: spermatogonia and primary spermatocytes are rich in RNA;; the vacuoles of type I and type II accessory cells contain an acid or neutral protein; the basal cells contain a polysaccharide. Basal cells contain granules which stain with PAS and Best’s carmine. The basal cells are also positive for glu- cose-6-phosphatase tests. The inclusions of type I accessory cells stain positively with bromphenol blue. Type I cells with large inclusions retain a reddish color with this dye, suggesting the protein is of an acidic nature. Type I ac- cessory cells concentrate Sudan black, but react negatively for phospholipids. These results suggest that the inclusion contains a lipid moiety and the contents of the inclusion may be a lipoprotein. In young type I cells, a few granules which stain with PAS are present. Type I cells react nega- tively to both acid and alkaline phosphatase cytochemist- ry. Type II accessory cells give similar results to type I cells. The inclusion of the type II cell probably contains a lipoprotein which is not a phospholipid and the protein moiety is probably acidic in nature. Type II cells also give a positive reaction for acid phosphatase, but negative for alkaline phosphatase. Type II cells are negative to the PAS technique. Table 1 summarizes the results of cyto- chemical tests for specific molecular groups. Seminal Vesicle and Prostate: The results obtained from sections stained with Lehman’s polychrome tech- nique suggest the presence of mucopolysaccharides and proteins in the secretory cells of the seminal vesicle and prostate. Specific cytochemical tests suggest that cells of the seminal vesicle contain glycogen and sulfonated, acid- ic mucopolysaccharides. The type I secretory cells of the prostate contain mucopolysaccharides which are probably sulfonated and acidic in nature. Type 2 secretory cells are rich in sulfonated acidic mucopolysaccharides. Type 1 cells also stain positively for acidic or neutral protein which suggests that the secretory products of the type 1 cells are acidic proteinaceous mucopolysaccharides. Table I gives the results of specific cytochemical tests of the seminal vesicle and prostate. Table 1 Results of cytochemical techniques of the testis, seminal vesicle and prostate Testis (Accessory Cells) Technique Basal Cell Alcian Blue (pH 1.0) a a Alcian Blue (pH 2.3) = = Acid Haematein ce nal Best’s Carmine alent Se Bromphenol Blue st aF PAS aR APOE AP Sudan Black = ar Thionin ae = Acid Phosphatase a ay Alkaline Phosphatase i = Glucose-6-Phosphatase AP cs Type I Type II Seminal Vesicle Prostate = ++ +++ — ++ ++ - - + + + ++ — ++ ++ + a + — + +++ + + + = de de +, ++, +++, increasing degrees of positive staining intensity; —, negative reaction; + questionable reaction Page 272 DISCUSSION The reproductive system of Colus stimpsoni conforms to the fundamental plan described for other neogastropods (Dakin, 1912; FRETTER, 1941; FRETTER & GRAHAM, 1962; PonpER, 1974; Houston, 1971,1976). In the male genital system of neogastropods, the genital duct may be divided into a posterior thin-walled portion which con- nects to the gonad, and a glandular, pallial portion which begins at the posterior limits of the mantle cavity. The thin-walled portion generally consists of a coiled, upper vas deferens which may be modified to form a sperm- storing seminal vesicle, and a lower, straight portion, the renal vas deferens (FRETTER, op. cit.; PONDER, op. cit.). In some neogastropod species, areas of the seminal vesicle may also function to ingest sperm (FRETTER, op. cit.; SMITH, 1967). The renal vas deferens opens into the pallial portion which is lined with prostatic tissue. The prostatic tissue may be confined to a localized area (the prostate gland) or may be found along the entire duct. In Colus stimpsoni both upper and lower portions of the vas deferens are convoluted. However, the renal vas deferens is not looped to the extent of the seminal vesicle. Also, sperm-ingesting areas within the seminal vesicle were not observed. Prostatic tissue in C. stimpsoni is found along the entire pallial portion of the genital duct and also lines the duct within the penis. The unusual con- dition of the prostate entering the body and coiling among the lobes of the salivary gland has not been reported for other neogastropods. In Colus stimpsoni, as in most animals, the spermato- genic tissue is confined to compartments which are sep- arated from somatic tissue. These compartments, the sem- iniferous tubules, contain accessory cells in addition to the germ cells. The accessory cells in C. stimpsoni are of 3 types: basal cells, type I and type II cells. Basal cells are generally restricted to the periphery of the semini- ferous tubule and evidence from light and electron micro- scopy suggests that these cells serve a nutritional function. The increases in the number and size of basal cells in relation to an increase in the amount of food available to the snail suggest that these cells may store nutriments, THE VELIGER Vol. 20; No. 3 probably in the form of lipids or fatty acids. Basal cells may provide nutriments for the developing germ cells. The type I and type II accessory cells appear to be endopolyploid. In type I cells, the nuclear material is confined to 1 or occasionally 2 nuclei; in type II cells the chromatin is divided into several nuclei of unequal sizes. Cytochemical and ultrastructural evidence indicates that these accessory cells synthesize proteins and store these products. The type II cell may also ingest abnormal or degenerating germ cells. This function is suggested by the presence of multiple flagella within these cells and the cytoplasmic inclusions which are reminiscent of residual bodies with pycnotic nuclei. The type II cells also show acid phosphatase activity. Type I and type II accessory cells in Colus sttmpsoni may function in providing large amounts of proteins for gamete development. Endopolyploid cells have been reported in various tis- sues of gastropods and in the ovotestis of pulmonates (BaBRAKZAI & MILLER, 1974). Babrakzai « Miller postu- lated that endopolyploid cells provide high rates of pro- tein synthesis for rapid use by developing gametes in the ovotestis of pulmonates. Results from the present study are consistent with this postulation. SUMMARY The male genital duct in Colus stimpsoni consists of a thin-walled posterior portion and a glandular, pallial portion. The posterior portion is convoluted and consists of the seminal vesicle and the renal vas deferens. In C. stimpsoni, the renal vas deferens is convoluted, unlike some other neogastropods. The pallial portion of the genital duct is lined with prostatic tissue along its entire length and functions as the prostate. The prostate passes into the body at the base of the penis and coils among the lobes of the right salivary gland. It then passes back out of the body near its point of entrance and enters the penis. The prostate is centrally located within the penis. At the junction of the renal vas deferens and prostate, a short diverticulum arises from the renal vas deferens and Explanation of Figures 78 to 21 Figure 78: Early stage (1) and middle stage (2) of maturation of type 1 secretory cell. R - rough endoplasmic reticulum; V -— vesicle Figure 19: Mature type 1 secretory cells of prostate. Arrow — microtubules; M - muscle; S — secretory cell Figure 20: Early maturation stage of type 2 secretory cell (2) with heterogeneous inclusion (*). (1) — type 1 secretory cell Figure 21: Mature type 2 secretory cell. Arrows — Golgi complex; * — heterogeneous inclusion Tue VELIGER, Vol. 20, No. 3 [West] Figures 18 to 21 ee Vol. 20; No. 3 THE VELIGER Page 273 ————————————— aad passes posteroventrally, opening into the posterior limits of the mantle cavity. The testis of Colus stimpsoni consists of numerous semi- niferous tubules which end blindly at the surface of the testis and are perpendicular to the spiral axis of the shell. The seminiferous tubules are filled with accessory cells and spermatogenic cells. Basal cells occur on the peri- phery of the seminiferous tubules and are filled with lipid droplets. It is suggested that these cells function in a nutritive manner. Type I and type II accessory cells occur in groups of 4 - 24 cells and these groups of cells are irregularly scattered within the seminiferous tubules. These accessory cells are endopolyploid and contain large amounts of protein. It is suggested that these cells function to provide large amounts of proteins for gamete develop- ment and phagocytose abnormally developing gametes. ACKNOWLEDGMENTS I would like to thank Drs. N. W. Riser and M. P. Morse for their advice and encouragement throughout this study. This study was submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at Northeastern University, Boston, Massachusetts, and was supported in part by HEW Grant No. RR 07143. Literature Cited Baprakzal, NooORULLAH & WALTER BERNARD MILLER 1974. Endopolyploid cells in pulmonate gastropods. 14 (3): 1264 Burge, C. N. « C. Wayne GEISELMAN 1971. Exact anhydride epoxy percentages for electron microscope em- bedding (Epon). Journ. Ultrastruct. Res. 36: 119 - 126 Daxin, WILLIAM JoHN 1912. Buccinum. Liverp. Mar. Biol. Comm. mem. on typ. British marine plts. & anim. 20: 123 pp. FretrTer, VERA 1941. The genital ducts of some British stenoglossan prosobranchs. Journ. mar. Biol. Assoc. U. K. 25: 173 - 211 Faetrer, Vera & AtasTam GRAHAM 1962. British prosobranch molluscs, their functional anatomy and eco- logy. London, Ray Soc. xvi+755 pp.; 316 figs. Gane, M. 5 1968. Techniques histologiques. Masson & Cie., Eds., Paris 1113 pp. Amer. Zool. GeIsELMAN, C. Wayne « C. N. Burke 1973. Exact anhydride:epoxy percentages for araldite and araldite- epon embedding. Journ. Ultrastruct. Res. 43: 220 - 227 Houston, Roy S. 1971. Reproductive biology of Thais emarginata (Deshayes, 1839) and Thats canaliculata (Duclos, 1832). The Veliger 13 (4): 348 - 357; 1 plt.; 5 text figs. (1 April 1971) 1976. The structure and function of neogastropod reproductive systems: with special reference to Columbella fuscata Sowerby, 1832. The Veliger 19 (1): 27-46; 1 plt.; 10 text figs. (1 July 1976) Humason, GretcHEN LYON 1967. Animal tissue techniques. 2nd ed. San Francisco, ix + 569 pp.; illus. Karnovsky, Morris J. 1965. A formaldehyde-glutaraldehyde fixation of high osmolarity for use in electron microscopy. Journ. Cell Biol. 27: 137A Leuman, H. E. 1965. A cytochemical “screening” method for demonstrating general molecular groups in single-tissue sections. II. Polychrome stain. Journ. Elisha Mitchell Sci. Soc. 81: 89 - 90 Pzarsz, ANTHONY Guy EvERSON 1968. Histochemistry. Theoretical and applied. Wilkins Co., Baltimore, 2 vols. Ponper, WINSTON F. 1974. The origin and evolution of the Neogastropoda. gia 12 (2): 295-338; 6 text figs. PorTMANN, ADOLF 1926. Le réle du spermatozoide atypique dans la formation des oeufs nourriciers de Buccinum undatum L. Arch. Zool. exp. gén. 65 (Notes et revues): 103 - 124 1927. Die Nahreierbildung durch atypische Spermien bei Buccinum undatum L. Zeitschr. Zellforsch. 5: 230 - 243 Rapwin, Georcz Epwarp & J. L. CHAMBERLIN 1973. Patterns of larval development in stenoglossan gastropods. Trans. San Diego Soc. Nat. Hist. 17: 107 - 118 Reyrno ps, E. S. 1963. The use of lead citrate at high pH as an electron-opaque stain in electron microscopy. Journ. Cell Biol. 17: 208 - 212 Riser, NaTHAN W. 1969. Feeding behavior of some New England marine gastropods. The Nautilus 82: 112-113 SmitH, Epmunp Hosart 1967. The reproductive system of the British Turridae (Gastropoda : Toxoglossa). The Veliger 10 (2): 176-187; plt. 18; 16 text figs. (1 October 1967) StzepMaN, H. E 1960. Section cutting in microscopy. ford, 172 pp. TuHorson, GUNNAR 1935. Studies on the egg-capsules and development of Arctic marine prosobranchs. Medd. on Grenland 100 (5). 1940. Notes on the egg-capsules of some North-Atlantic prosobranchs of the genus Troschelia, Chrysodomus, Volutopsis, Sipho and Trophon. Vidensk. Medd. fra Dansk. naturh. Foren 104: 251 - 265 Wavxer, Muriet « H. C. MacGrecor 1968. Spermatogenesis and the structure of the mature sperm in Nu- cella lapillus (L.). Journ. Cell Sci. 3: 95 -104 Watson, M. L. 1958. Staining of tissue sections for electron microscopy with heavy metals. Journ. Biophys. Biochem. Cytol. 4: 475 - 478 West, Davo L. 1973. Notes on the development of Colus stimpsoni (Prosobranchia: W. H. Freeman and Company, 3rd ed., Williams & Malacolo- (11 March 1974) Blackwell Scient. Publ. Ox- Buccinidae). The Nautilus 87 (1): 1-4 Woopwarp, T. M. 1935. Spermic dimorphism in Goniobasis laqueata (Say). Journ. Morph. 57: 1 - 29 Page 274 THE VELIGER Vol. 20; No. 3 Hiatella solida (Sowerby, 1834) (Mollusca : Hiatellidae) on Concholepas concholepas (Bruguiére, 1789) and other Substrates BY CARLOS GALLARDO S. Instituto de Zoologia, Universidad Austral de Chile. Casilla 567. Valdivia, Chile AND CECILIA OSORIO R. Laboratorio de Hidrobiologia, Departamento de Biologia, Facultad de Ciencias. Casilla 147. Santiago, Chile (3 Text figures) INTRODUCTION THE ABILITY TO PERFORATE hard substrates has evolved independently in different groups of bivalves. In species of the genus Hiatella this specialization was probably assumed after initial life as byssally attached nestlers (YonceE, 1964). The primitive byssiferous nestler char- acteristic is still present in Hzatella gallicana (Lamarck, 1818) and in H. arctica (Linnaeus, 1767), according to Hunter (1949). Members of a single species may reveal other fixation behavior determined by the substrate. On a hard but creviced rock surface, the spat nestle, using the byssus, and those settling on a smooth rock surface of a soft, homogenous rock will move about until they find a crevice wherein to bore. HUNTER (of. cit.) points out that the byssus is absent in boring individuals. According to Narcui (1973) and to data compiled by CarcELLes (1944), Hiatella solida uses both behavioral patterns to place itself securely in the substrate. In Brazil, Narcui (1973) reports that individuals use the byssus to fix themselves to the sea-squirt Polyandrocarpa zorritens- ts, among tubes of the polychaete Phragmatopoma lapid- osa, or under clusters of the bryozoan Zoobotryon pellu- cidum. They have also been found boring or nestling in the intertidal zone. According to CarcELLEs (of. cit.), in Ar- gentina the species lives on rocky sea-beds in inter- or sub- tidal zones, boring the sandstone or incrusted in bivalve or gastropod shells. There is very little in Chilean literature on habitat or relationship of the different substrates. SooTt-RYEN (1959) states that Hiatella occurs on hard sea-beds from the intertidal zone to depths of 70m. MarincovicH (1973) found it nestling in crevices of intertidal rocks as well as on holdfasts of the kelp Lessonia nigrescens Bory, 1825. Its geographic range is rather extensive, from South Ecuador to Cape Horn,and in the Atlantic its northern limit is the southern part of Brazil. This paper provides information on occurrence in southern Chile, especially the association with the gastropod Concholepas. MATERIAL anp METHODS Two collections of adult Concholepas concholepas (Bru- guiére, 1789) (Table 1) as well as sporadic samplings of Fissurella nigra (Lesson, 1830) and the tunicate Pyura chilensis (Molina, 1782) were made. Samples were ob- tained by scuba-diving in the south bank of the entrance to Corral Bay (39°51’S; 73°27’ W). Specimens of Concholepas were kept in aquaria, where the presence of Hiatella solida was revealed by their pro- jecting siphons. To remove the bivalve, the shells of the Concholepas were gradually broken up. Notes were taken on the size of the bivalves, their location, and their rela- tion with the accompanying epibiontic community. Ma- croscopic sections of Concholepas shells showed the shape of the excavations and the position of the Hiatella within. Vol. 20; No. 3 Other substrates on which Hiatella solida occurs were also studied: Megabalanus psittacus (Molina, 1782), bought at the market in Valdivia on October 26, 1975; intertidal samples of Phragmatopoma virginu Kinberg, 1867, and holdfasts of the kelp Lessonia nigrescens ob- tained in Mehuin Bay (39°25’S; 73°10’ W) on April 15, 1976. Descriptions by Oitsson (1961) and DeLi (1964) were used for the identification of the Hiatella, the identi- fication being later confirmed by Dr. Myra Keen. RESULTS Hiatella solida was found as an incrustive epibiont in the calcareous shells of Concholepas concholepas from Corral Bay (Table 1); on Fissurella nigra from among the kelp Durvillaea antarctica (Chamisso) (obtained with scuba- diving by Col. C. Moreno on May 5, 1975), and on Mega- balanus psittacus. Hiatella solida A. — Incrustive manifestations of the bivalve as seen from the inside of a Fissurella nigra shell B — Ventral view of Hiatella solida THE VELIGER Page 275 On a Fissurella nigra specimen measuring 11.6cm in length, gcm in width, 4 Hiatella specimens occurred. Three of them were on different parts of the shell, with an oblique orientation to the surface. Their lengths were 11.5, 6.4, and 6.2mm, respectively. The 4" specimen was gmm long and was next to the apical foramen in a mound formed by the internal layers of the shell (Figure 1A). Both the periostracum and the middle layers of the Fissur- ella were seriously damaged by several tubiculous poly- chaete tunnels. A specimen of Hiatella measuring 27 X 14mm was found to have incrusted the base of a Mega- balanus shell. In sublittoral samples of the colonial tunicate Pyura chilensis in Corral Bay, the Hiatella nestled in the inter- stices between individuals. It forms part of the Pyura community, along with other bivalves such as Hormomya granulata (Hanley, 1843), Lyonsia fretalis Dall, 1915, Aulacomya ater (Molina, 1782), and Mytilus chilensis Hupe, 1854, cited by ZamoraNo & Moreno (1975). However, Hiatella was absent in intertidal samples of Pyura chilensis, Phragmatopoma virginii, and Lessonia nigrescens from Mehuin Bay. EPIBIOSIS on Conchole pas The frequency with which Hiatella solida acts as an in- crustive epibiont on Concholepas is variable, as can be seen in Table 1. It may also be noted that on a single shell both juveniles — the smallest measuring just over 2mm — and adults of different sizes may occur. The largest adult specimen studied was 17mm long. The bivalves lie in their burrows approximately perpendicular to the surface, with the posterior end directed toward the external open- ing (Figure 2A). Some individuals assume a more ob- lique position, thus attaining greater length without per- forating the inner layer (Figure 2B). However, no matter which is the orientation, larger specimens may reach the inner layer. At such points, the shell material takes on a yellowish-brown color and is fibrous and brittle. In high density areas individuals are very close together and 2 or more burrows may come to communicate with each other as individuals grow. In a 9cm’ area, 10 medi- um and large sized individuals were counted. One speci- men was byssally attached to its excavation by a single thread. HuNTER (1949) observed in Hiatella gallicana and H. arctica, a single byssus thread is used, but rarely by borers, to alter their position in the burrow. There seems to be no direct relationship between the frequency of bivalve occurrence on Concholepas and the size of the gastropod shell. Mostly, presence on the shell is closely related to the developmental stage of the epi- Page 276 Figure 2 Hiatella solida A - Concholepas concholepas shell section showing an adult bivalve incrusted perpendicularly B - Section showing the bivalve in an oblique orientation biontic community (see Table 1). Specimens of Hiatella solida are frequently found on shells with large numbers of tubicolous epibiontic polychaetes, calcareous seaweeds, and empty dead barnacle shells, or among abundant phoronids. These seem to be the most important in help- ing the Hiatella to bore successfuly, especially on the less protected faces of the substrate. Several juvenile and medium sized bivalves with siphons projected were found inside empty shells of Balanus flosculus Darwin, 1854, and B. laevis Bruguiére, 1789 (Figure 3). This indicates that the bivalve first entered the shell, then became a burrower. Being small, it had been capable of going through the narrow opening. A similar habitat was re- THE VELIGER Vol. 20; No. 3 ported for Hiatella by HunTER (1949) — non-boring adults byssally attached inside empty shells of large bar- nacles. Some parts of the gastropod shell were eroded either by mechanical or by abrasive agents, leaving the Hiatella partially exposed or even completely dislodging it from the shelter (Figure 3). Incrustive juvenile bivalves may often be found at the umbo or in adjacent areas Figure 3 Hiatella solida Surface view of Concholepas concholepas shell with Hiatella solida and other incrustive epibionts. Young specimens of Hiatella solida are observed inside empty dead barnacle shells eroded by tubiculous polychaetes of the family Spionidae or by abundant Phoronis ovalis Wright, 1856, as epibionts. The incrustive worms therefore play an important part in preparing an adequate substrate for Hiatella to bore in. Gastropod shells with only a few epibionts do not af ford the bivalves adequate conditions for quick penetra- tion. Then the Hiatellas are restricted to the crevice on the left side of the Concholepas shell, under the umbo, where attachment is usually by a byssus. Hiatellas using this shelter, especially large individuals, bore deep bur- rows, possibly to avoid exposure as they grow. The cor- rosion carried on by other burrowers greatly aids Hzatella with its boring. Some other filtering epibionts also colon- ize the area, the most important being Verruca laevigata (Sowerby, 1827), bryozoans, and some sea-squirts. Vol. 20; No. 3 THE VELIGER Page 277 Table 1 Incidence of Hiatella solida and other epibionts on Concholepas concholepas shells from Corral Bay (11/10/74 - 1/4/76) Epibionts Diameter of Hiatella Polychaete Spirorbis — Phoronts Balanus Verruca Balanus Chthamalus Calcareous Concholepas solida tunnels sp. ovalis Bryozoa _flosculus laevigata laevis scabrosus algae concholepas 63 mm 3 ++ ~ - ++ — ++ — ~ - 96 i i x a x er = = a =a 98 3 ap ar x = = = = = = = 99 3 aPar = = = + = = = = 100 - x x = » = ~ - - - 102 = ar = FS. rz a a + = ay 102 o rE = a = ear = = = x 103 > x = = = a > = = 103 3 ats ate = = + = — = = 104 16 ++ seat — = ++ = - — ++ 105 4 ++ - - _ ++ - — ++ ++ 105 = + = = = a = = = = 107 5 = = ++ a ++ = = — = 108 = = at = = + = = = = 110 9 Staats = — — = ++ — = — 110 i = = = Par = aR ar = = = 112 1 amar an ates = = = rar = = 112 = + = = = = x x = = 120 1 ++ - — - ++ - ++ - ~ 123 4 re = ++ = = = = = = 125 28 ++ — ~ ~ ++ - — ~ ++ ++ = abundant; + = scarce; X = presence. DISCUSSION Hatella solida is a nestler, using the byssus to fix itself among colonies of the tunicate Pyura chilensis, in inter- tidal crevices, in Lessonia nigrescens holdfasts in northern Chile (Marincovicu, 1971), or in the sea-squirts Poly- androcarpa zorritensis and Phragmatopoma lapidosa on the coasts of Brazil (NaRcHI, 1973). The ability to per- forate certain substrates and gastropod shells, mentioned briefly by CarcELLes (1944), is confirmed here by obser- vations on Concholepas concholepas and Fissurella nigra. It can bore not only in the calcareous shells of mollusks but also in those of other invertebrates, such as Megaba- lanus psittacus. The nestling or boring behavior is determined not only by the hardness of the substrate but also by whether there are adequate shelters — cracks or crevices — in which to nestle. These shelters seem to be required for attachment on hard, impenetrable substrates or in intertidal rocks (MarrncovicH, 1973). Thus, the species never prospers in impenetrable hard substrates with homogeneous areas lacking shelters. These conclusions also seem to apply for Hiatella gallicana and H. arctica. Substrates with plenty of safe shelters for permanent nestling would be those offered by such sea-squirt colonies as Phragmatopoma, Lessonia nigrescens holdfasts, and prob- ably some others. Concholepas affords Hiatella solida shelter and crevices on a transitory basis. Neither the chink under the umbo of the Concholepas or the empty cirriped shells attached on it offer Hiatella a safe shelter on which it can grow without becoming exposed. How- ever, these shelters are vital in helping the bivalve to bore into the shell successfully. Empty barnacle shells play an essential role. Moreover, abrasion and burrows of other incrustive epibionts such as polychaetes and Phoronis ovalis seem to be prerequisites for successful boring into the substrate. The calcareous shells of Megabalanus psittacus appar- ently offer the Hiatella an unsafe substrate for byssal fixation alone. Besides being easy prey for possible preda- tors while so attached, the bivalves must also undergo the same abrasion as the Concholepas, especially when the Page 278 THE VELIGER Vol. 20; No. 3 UUM ———————— ee gastropod enters into cracks and crevices of the hard substrate. The fact that the valves of Hiatella solida close partially, leaving the ventral side exposed, seems to indi- cate that the species is adapted to life only inside shelters in the substrate. As YoncE (1962, 1964) postulated, the boring habit of Hiatella in calcareous shells as well as in other sub- strates can be considered to have developed subsequently to the nestling habit and byssal fixation in benthonic shelters. Adopting the boring habit seems to be an impor- tant step toward the colonization of penetrable substrates that are open and homogeneous and therefore not suit- able for long-time nestling. Assessment of the damage that incrustive epibionts cause on Concholepas shells is scanty. Only ARENAS (1972) gives information on the effect that Phoronis ovalis has on the gastropod. It has been observed that large specimens of Hzatella solida can penetrate even the inner layer. However, no manifestations of layer thicken- ing were apparent at the affected points, though this occurs with massive incrustations of Ph. ovalis. The effect of H. solida, though damaging, is local and secondary, almost always following incrustations of polychaetes and Phoronis. A similar situation seems to occur in Fissurella nigra. In one specimen, shown in Figure 1A, the mound around the incrustive bivalve may possibly be a defense used when the penetrator endangers vital functions. No such response was apparent in other areas of the shells that were affected by H. solida. In systematic studies of Hiatella solida Soot-RyEN (1959) distinguishes two forms and Dett (1964) ob- served variability in form and shape in specimens from the East Coast of South America and the Magallanes region. This variability, which has probably led to the series of names for the species, could be caused by the types of surface on which they settle. In studies carried out on epibionts in Concholepas con- cholepas from Caleta Leandro (36°39'S; 73°05’W) Hia- tella solida was not found (Lozapa et al., 1976). ACKNOWLEDGMENTS The authors acknowledge the help of C. Jara, C. Moreno, W Steffen and J. Zamorano in obtaining the samples. We also wish to thank Professor N. Bahamonde and Dr. R. Schlatter for reviewing the paper and for their helpful suggestions, Literature Cited Arenas, J. 1972. Phoronis ovalis Wright in Chile. 7: 113 - 115 Carce.ies, ALBERTO R. 1944. Cat4logo de los moluscos marinos de Puerto Quequén (Répub- lica Argentina). Rev. Mus. de la Plata, Zooligia (n.s.) $: 233 to 309; pits. 1-15 Det, R. K. 1964. Antarctic and subantarctic Mollusca: Amphineura, Scapho- poda and Bivalvia. Discovery Reprts., Cambridge Univ. Press, Cambridge) 33: 222 - 226; pit. VII, 11 Hunter, W. R. 1949. The structure and behaviour of Hiatella gallicana (Lamarck) and H. arctica (L.) with special reference to the boring, habit. Proc. Roy. Soc. Edinb., Sect. B, 63 (3): 271 - 289 Lozapa, E., M. T. Lopez & R. Desquevroux 1976. Aspectos ecolégicos de poblaciones chilenas de loco Concholepas concholepas (Bruguiére, 1789) (Moll. Gast. Muricidae). Biol. Pesq. Chile (8): 5-29 Marincovicn, Lourz N., Jr. 1973. Intertidal mollusks of Iquique, Chile. Mus. Nat. Hist., Sci. Bull. 16: 1-49 Narcui, WALTER 1973. On the functional morphology of Hiatella solida (Hiatellidae: Bivalvia) . Mar. Biol. 19: 332 - 337 Oxsson, AxeL ADOLF T1961. Mollusks of the tropical eastern Pacific particularly from the southern half of the Panamic-Pacific faunal province (Panama to Peru). Ithaca, New York (Paleo. Res. Inst.) 574 pp.; 86 plts. (10 March 1961) Soot-Ryen, TRON 1959. Pelecypoda. Repts. Lund Univ. Chile Exped. (1948-49). 35: 1 - 86; plts. 1-4; 6 text figs. Yonoz, CHartes Maurice 1962. On the primitive significance of the byssus in the Bivalvia and its effects in evolution. Journ. Mar. Biol. Assoc. U. K. 42: 112 - 125 1964. Physiology of Mollusca, Vol. 1. Acad. Press Inc., New York. ZAMORANO, J. & C. Moreno 1975. Comunidades benténicas del] sublitoral rocoso de Bahia de Cor- ral. I. Area minima de muestreo y descripcién cuantitativa de la Asocia- cién de Pyura chilensis Molina. Medio Ambiente 1: 58 - 66 Stud. Neotrop. Fauna Los Angeles County Vol. 20; No. 3 THE VELIGER Page 279 Zonation of Marine Gastropods on a Rocky Intertidal Shore in the Admiralty Gulf, Western Australia, with Emphasis on the Genus Nerita FRED ETHAN WELLS Western Australian Museum, Perth 6000, Western Australia (3 Text figures) INTRODUCTION THE ZONATION OF MARINE MOLLUSCS along the intertidal Shorelines of the world has been intensively investigated in many areas since the broad perspectives of the Stephen- sons were published (STEPHENSON & STEPHENSON, 1949). A number of more recent studies have investigated the zonation of marine molluscs on the eastern and southern coasts of Australia (ENDEAN et al., 1956; UNDERWOOD, 1972) or have examined particular features of the biology of intertidal molluscs occurring on those shores (CoLE- MAN, 1976; UNDERWOOD, 1975, 1976). Development of our understanding ‘of the marine intertidal communities of Western Australia has been severely handicapped by the size of the coastline, about 6500km, and the small number of marine biologists in the state. The marine inter- tidal communities of the Perth area have been examined (Hopcxin, Marsu «& SmitTH, 1955; Marsu « Hopckin, 1962; PrmEAux, 1976) but there is almost no published information available on any aspect of the molluscan communities of the north coast of Western Australia east of Derby. The coastline of that area has been largely inaccessible from land because of a lack of roads and access from the sea has been prevented by the lack of available ship-time. Over the last several years the West- ern Australian Museum has conducted several field pro- grams in the northeastern corner of the state and it was felt that sufficient information was now available to pin- point a number of areas of particular interest which could be examined on a major expedition. The Western Austral- ian Museum in conjunction with the Field Museum of Natural History, Chicago, USA, conducted an intensive survey of the land, freshwater and marine fauna of the Admiralty Gulf and adjacent Mitchell Plateau from 16 October to 6 November 1976. The teams had a variety of interests: mammals, birds and reptiles, fish, insects, mites, and molluscs. This paper is the first report on the work done on molluscs during the expedition. THE STUDY AREA The Admiralty Gulf (Figure 1) is a large marine embay- ment with an area of approximately 1700km* on the extreme northeast coast of Western Australia about 300km W of the Northern Territory border. The waters of the Gulf are essentially marine but minor freshwater inflows are received from the Mitchell and Lawley Rivers; lim- ited estuarine areas are associated with each river. The complex system of tides which occurs in the Gulf makes the area an interesting one for the examination of the patterns of zonation of marine intertidal mollusc species. The tidal range is extensive, with a maximum range of as much as 8.8m on spring tides, from —1.7m on the shore to +7.1m. The average tidal amplitude is 4.4m. The tides are predominantly semidiurnal, but diurnal tides occur during the changeover between neap and spring tide periods. When the changeover occurs the tidal levels remain static for several hours at a level of about +2.6m. ‘Two major intertidal habitats occur in the Admiralty Gulf. A myriad of small bays is found, each with extensive mangrove development at the midtide levels. In many of the bays well developed mudflats occur landward of the Page 280 Admiralty Gulf ! Western Australia THE VELIGER Vol. 20; No. 3 ut # ° Admiralty Gulf <3 Sri ° Se a es eats eee 5 Figure 1 Map of Australia Showing the Position of the Admiralty Gulf. The Insert shows the Admiralty Gulf itself and the Location of Walsh Point in the Gulf. L, Lawley River; M, Mitchell River; MI, Malcolm Island W, Walsh Point mangroves. These areas are baked dry by the tropical sun during the neap tides but are submerged intermittently during spring tides. Few molluscs occur on or in the land- ward mudflats. Extensive mudflats which project seaward of the mangroves for as much as a kilometer are exposed by low spring tides. A variety of molluscs is found on the surface and in the mud of these flats. The headlands be- tween the bays are rocky shores in the intertidal zones; the rock is of either sandstone or basalt, depending on the particular location in the Admiralty Gulf. Three marine programs were conducted in the Admiralty Gulf by the WAM Mollusc Department. The results of a faunal sur- vey of the Gulf and the nearby Institut Islands and an intensive examination of the molluscs of the mangrove swamp at Port Warrender in the Gulf will be published elsewhere. The present paper examines the zonation of _epifaunal marine gastropods at Walsh Point in the Ad- miralty Gulf. Walsh Point is a small basaltic peninsula 2.7km long (Figure 1) located at latitude 14°35’S and a longitude of 125°50’E. The seaward tip of Walsh Point, where the samples reported here were collected, is almost complete- ly rocky in the intertidal areas. The shore of the high inter- tidal zone is composed of small rocks 5 to 13cm in dia- meter which are several layers deep. These rocks are easily moved about by even moderate wave action. The rocks are larger at the midtide level, about 10 to 30cm in diameter, and are only 2 or 3 deep. Base rock isexposed insome areas of the midtide zone. At the lower intertidal areas the basalt rocks are 30cm or larger in diameter and are separated Vol. 20; No. 3 from each other by up to 30cm of sand; very few rocks adjoin each other in the low intertidal zone. At a level of about 0.5m the shore changes rapidly into a sandy beach in which rocks are rare. Walsh Point is located well within the Admiralty Gulf and is thus protected from the wave action of the open sea. The water is extremely turbid at low tide even on calm days, with a visibility of 15cm or less. As the tide rises, visibility increases to about gocm. MATERIALS anp METHODS Four transects approximately 10m apart were made along the rocky shore of Walsh Point from the 0.5m tide level to the 7.0m level. The tide level was determined by comparison of the water levels at Walsh Point with tide charts for nearby Malcolm Island. A 1mX1m quadrat was searched at every other meter along the transect and all gastropods found were removed and preserved in 10% formalin. All specimens were returned to the museum lab- oratories where they were identified and counted. Shell length of all individuals of Nerita was measured with vernier calipers. The resulting data were grouped into 0.5m shore levels. Voucher specimens of all species col- lected have been deposited in the Western Australian Museum where they have catalogue numbrs WAM 1256- 76 to 1274-76. RESULTS A total of 3375 individuals belonging to 18 species of gas- tropod molluscs was collected in the study. The density of each species and the standard deviation in each zone is given in Table 1. Three zones were demonstrated by the species: species which had their maximum level of abun- dance at the lowest tide level, then tapered off as the height on the shore increased; species with a maximum density near the midtide level whose densities decreased both above and below the midtide region; and species most abundant in the upper tidal levels whose density decreased below the hightide zone. The species compris- ing each zone will be discussed in turn. Of the 18 species collected 10 had their maximum abundance in or near the lowest zone studied. Although a majority of species was restricted to this lowest tidal area, the numbers of individuals of each species were generally very low. The entire group was represented by 507 individuals, which was 15.0% of all gastropods col- lected. Included in this group is a number of minor spe- cies which were represented by 25 or fewer individuals: THE VELIGER Page 281 Cantharus erythrostoma, Cantharidus strigatus, Colum- bella duclosiana, Haliotis varia, Onchidium daemelli, Pattelloida saccharina, Montfortula variegata, and Tro- chus lineatus. The most abundant species in the low inter- tidal zone was Morula margariticola. This species had its maximum density in the lowest two zones (Figure 2), from 0.5 to 1.5m on the shore, where it had an average density of 13.7/m?. The density of M. margariticola de- creased rapidly above this level, with only 3 individuals being found above the 2.5m mark. The other abundant species in the lowest group, Thais kienert (Figure 2), had a wider distribution which ranged from 0.5 to 4.5m. This species was much less abundant than M. margariti- cola and had a maximum density of only 3.5/m? in the 1.5 to 2.0m zone. The second group of 6 species had their highest densi- ties in the middle tidal ranges and were less abundant at both higher and lower levels on the shore. The midtide group was numerically dominant on the shore and ac- counted for 76.0% of all specimens collected. Four minor species are included in this group: Monodonta labio, Morula granulata, a species tentatively identi- fied as Cuma gradata, and an unidentified thaiad species. The most interesting feature of the midtide group was the dominance of 2 species of the genus Nerita: N. reticulata and N. undata. Nerita reticulata (Figure 2) was the most abundant species collected in the study, being represented by 1300 individuals or 38.9% of all gastropods collected. Nerta undata (Figure 2) was second with 1090 individuals (32.3% of the total). Both species had a wide vertical range on the shore that encompassed virtually all of the zones studied. Nerita reticulata had low numbers of in- dividuals below the 1.5m mark and increased gradually in density to the 3.0m level, after which the numbers dropped steadily. While it had essentially the same verti- cal range as N. reticulata, the zonation pattern of N. un- data was very different (Figure 2). Low numbers of 4.8 and less per m* were encountered both below 2.5m and above 5.5m. Instead of a gradual increase toward the midtide level as was observed in N. reticulata, the num- ber of individuals of N. undata was relatively constant at high levels of 16.0 to 20.3/m* between the 2.5 and 5.5m levels. The maximum abundance of N. reticulata occurred in the 2.5 to 3.0m level, one step below the high- est density of N. undata. While the mean heights of the populations of the 2 species were different, N. reticulata had a mean of 3.0+2.5m and N. undata was 3.9+ 2.6m, the differences were not statistically significant (t-test, 0.05 level). Only 2 species had their maximum abundances in the upper intertidal levels and together they constituted only Page 282 THE VELIGER _ Vol. 20; No. 3 Table 1 Density (No./m*) of Marine Gastropods in the Intertidal Zone of Walsh Point, Admiralty Gulf, Western Australia, With one Standard Deviation from the Mean Indicated for all Zones in which more than Four Individuals of a Species were Found. The Level of Maximum Density of a Species is enclosed by a Square Height above datum (m) MESS as saa. 3.5-4.0 4.0-4.5 4.5-5.0 | 5.0-5.5 5.5-6.0 6.0-6.5 6.5-7.0 Species 0.5-1.0 1.0-1.5 1.5-2.0 |] 2.0-2.5 2.5-3.0 3.0-3.5 Lower intertidal species 0.1 0.1 0.1 0.1 Cantharidus strigatus Cantharus erythrostoma Linnaeus, 1758 Columbella duclosiana Haliotis varia Linnaeus, 1758 Montfortula variegata 0.2 0.3 Adams, 1852 Morula margariticola 3.4 2.4 0.3 0.1 Broderip, 1832 acg}8) |) seule Onchidium daemelli | 0.5 Semper, 1882 Patelloida saccharina 0.1 0.1 0.1 0.1 (Linnaeus, 1758) Thais kieneri 0.9 1.7 1.5 1.4 1.4 0.1 0.3 Deshayes, 1844 as 12 +1.6 acl.) +13 +0.7 Trochus lineatus Lamarck, 1822 Middle intertidal species cf. Cuma gradata 0.3 0.1 0.5 0.8 0.8 0.4 1.0 +0.6 | +04 +111 +23) +2.7 Monodonta labio 0.1 0.3 0.4 1.8 2.5 0.8 0.3 Linnaeus, 1758 +15 +1.4 +1.7 +1.4 Morula granulata j Duclos, 1832 Nerita reticulata 5.0 7.6 12.8 13.4 35.9 32.5 20.8 11.7 3.8 0.4 0.3 Karsten, 1789 +144 +94 +143) +£9.6 +143) +13.2 +9.0 +5.0 +0.4 Nerita undata 0.5 1.3 4.8 17.9 19.5 19.3 17.6 16.3 3.4 15 Linnaeus, 1758 ac) ails)! || aeia\(0) +61 +60 +92 | +89 +30 +19 Thaiad Species 1 0.3 0.6 0.4 1.0 0.3 +0.6 +0.5 +0.9 Upper intertidal species Nerita polita 0.3 1.1 0.6 1.5 0.6 Linnaeus, 1758 +0.5 +2.2 +0.5 Planaxts sulcatus 0.3 1.1 3.5 5.8 1.4 (Born, 1780) +26 +3.6 +84 +3.0 29.5 11.0 3.5 +185 +109 +3. 47.6 +1.8 38.1 +18.4 Total density 23.3 24.7 23.7 24.9 61.1 58.7 £11.7 £108 +17.5 | 10.1 {+11.3 | 25.8 Vol. 20; No. 3 THE VELIGER Page 283 30 30 Morula margariticola Nerita polita 15 ‘ fC) I 2 3 4 5 6 7 3° Thais kieneri 3° Planaxis sulcatus 15 = fo) & 6 4 5 6 7 ° I 2 3 4 5 6 7 Z i fas fa} Nerita reticulata Density 30 60 15 go e o OF Mmm Z esr kal) 6 yon) OF” 7 ORMU I Dee acquire tin) (65 Gag 30 Nerita undata 80 Diversity 15 40 ) ) to) I 2 3 4 5 6 7 0 I 2 3 4 5 6 7 Height (m) Figure 2 Zonation of Marine Gastropod Species at Walsh Point, Admiralty Gulf; Western Australia, Showing the Density of Each Species at the Various Tidal Heights. Densities Shown are Means and the Standard Deviation for Each Mean is Indicated Page 284 9.0% of all gastropods collected. Both Nerita polita (Fig- ure 2) and Planaxis sulcatus (Figure 2) were found from 3.5 to 6.5m on the shore and had their maximum con- centrations in the zone of 5.0 to 5.5m. Planaxis sulcatus was the dominant member of this group, being 4 times as abundant as N. polita. The total density of gastropods (Figure 2) was mod- erate in the lower intertidal levels, ranging from 24.3/m” at the 0.5 to 1.0m level to 24.9/m’ at the 2.0 to 2.5m interval. Density in the next shore zone more than doubled to 61.1/m?, largely due to increases in the densities of Nerita undata and N. reticulata. The fact that the total abundance of gastropods declined steadily above the 3.0m mark is a further indication of the numerical dominance of these 2 midtide species. A Stmpson’s (1949) index of diversity was calculated for each of the intertidal zones. Maximum diversity, where each individual belongs to a different species, is indicated by a value of 0 on the Simp- son index; minimum diversity, where all individuals are of the same species, has a value of 1. The diversity index closely paralleled the total gastropod density (Figure 2). The midtide region of highest density was also the area of highest diversity. Diversity was lower both above and below the midtide region, as was the total density. The index is designed to prevent the total number of individu- als studied from influencing the index of diversity. Thus the higher diversity in the midtide region is real and is not simply an artifact of the higher number of individ- uals which occurred there. One of the most interesting features that can be ex- amined with a study of this type is the partitioning of the environment by groups of closely related species in a sym- patric association with each other. Two such groups were identified in this investigation. Four species of thai- ad: Cuma gradata (tentative identification), Morula granulata, M. margariticola, and an unidentified species, occurred in the middle and lower intertidal regions. Despite the fact that they are all carnivorous species in the same subfamily, the thaiads belong to 3 different genera and are not closely related taxonomically. Together they comprised only 16.1% of all gastropods collected. More interesting are the 3 species of the herbivorous genus Nerita which feed by rasping algal material off the sur- faces of the rocks on which they live. In contrast to the thaiads the nerites all belong to the same genus and nu- merically dominated the study, accounting for 72.9% of all gastropods collected. A number of studies (e. g. Epwarps, 1969) of inter- tidal gastropods have demonstrated that smaller individu- als of a species tend to be concentrated on the lower shore areas and larger specimens tend to be located higher in the intertidal zone. Lower intertidal areas are covered THE VELIGER Vol. 20; No. 3 by seawater for a greater length of time during each tidal cycle than the areas higher up, providing a more gentle environment for a marine species. Juvenile individ- uals are generally more susceptible to variations in condi- 20 16 A 12 A & 8 4 Nerita undata 4 oO ° 1 2 $1. 4-0. 05 ee ame 7 Level (m) \ . . 20 Nerita polita 5 bo S oe = eee teeny igre Finca aN 5 6 7 i Level (m) Figure 3 Mean Lengths of the Three Nerita Species Collected at Walsh Point, Admiralty Gulf, Western Australia. For Each Tidal height a Standard Deviation of the Mean is Indicated for Every Sample of More than Five Individuals Vol. 20; No. 3 tions than are the adults, which usually have greater envi- ronmental tolerances. Thus the mean sizes of each of the 3 Nerita species would be expected to increase as the area sampled moved up the shoreline, but this was not the case (Figure 3). The mean sizes of each species were stable throughout the tide levels and no demonstrable trends were evident, indicating that juveniles were not in fact found lower on the shore than adults, but there was a distinct tendency for very large individuals of each spe- cies to be concentrated in the upper meter of the species’ range. The species themselves demonstrated a clear in- crease in size with tidal height (Table 2). The average length of all N. reticulata measured was 9.4+0.7mm. This species was the lowest on the shore with a mean tidal height of 3.0+2.5m. The intermediate N. undata at 3.9 +2.6m averaged 13.0+2.1mm, 1.4 times as large as N. reticulata. Nerita polita, which was highest on the shore at 5.2+0.2m, averaged 16.141.6mm, 1.2 times as large as N. undata. The index of diversity devised by Simpson (1949) was modified by Mortsira (1959) as the basis of an index of overlap; the appropriateness of the overlap index has been discussed by Horn (1966). The overlap index de- veloped by Morisrra (op. cit.) has been used to deter- mine the degrees of overlap between the populations of the 3 Nerita species on the shore of Walsh Point (Table 2). The 2 species which occur at opposite ends of the shore, N. reticulata and N. polita, had the lowest overlap, 0.07. Nerita undata occupied an intermediate position on the shore and had a high overlap with both N. reticulata below and N. polita above. DISCUSSION The diversity of gastropods was highest in the midtide areas and tapered off towards both the higher and lower THE VELIGER Page 285 portions of the shore. One would expect diversity to be highest in the lowest intertidal areas which are covered by seawater for the longest periods during each tidal cycle. REIMER (1976), for example, studied the intertidal fauna on rocky shores on the Pacific coast of Panama. Of 220 species identified, 42% were found exclusively in the lower intertidal zone, 17% were limited to the midlittoral and only 6% were exclusive to the upper shore. The remaining species had distributions that overlapped at least 2 zones. The lower intertidal zone had the highest diversity and the splash zone the lowest. The unexpectedly low diversity in the lower intertidal zone of the Admiralty Gulf can be easily explained in terms of the beach structure. The rocks of the midtide region are large enough that they are not rolled about by waves except during storms. With one rock lying on another there is an abundance of crev- ices and nooks and crannies for the snails to retreat into during the emersion at low tide. In the lower portion of the intertidal zone the rocks are spaced about 30cm apart, eliminating the availability of refuges in the gaps between adjacent rocks. The index of diversity is based on the quantitative samples in which only 18 of the most common gastropod species were found. Intense collection in the area for the report on the molluscan fauna of the Admiralty Gulf has shown that at least 75 gastropod spe- cies occur at Walsh Point. The dominance of the 3 Nerita species was overwhelm- ing: N. reticulata constituted 38.9% of all gastropods col- lected and N. undata 32.3%. Nerita polita was a minor species at 1.8%. The dominance of the 3 congeneric spe- cies raises the question of how they partition the available resources and which factors are potentially limiting. The population of N. polita is partially segregated spatially from the other 2 species. Nerita polita is an upper inter- tidal species with a mean population level of 5.2+0.2m, above the levels of N. reticulata and N. undata. Table 2 Characteristics of the Populations of Nerita at Walsh Point, Admiralty Gulf; Western Australia Shell length Niche overlap Shore height Minimum Maximum Mean+1SD | Minimum Maximum Mean+1SD (m) (mm) (mm) Nerita reticulata 4 12 Nerita undata 3 22 Nerita polita 11 22 9.4+0.7 13.0+2.1 16.1+1.6 Species 3.0+2.5 3.9+2.6 5.2+0.2 0.07 Nerita reticulata— Nerita polita 0.29 Nerita undata— Nerita polita 0.35 Nerita reticulata— Nerita undata Page 286 The index of overlap between Nerita polita and N. reticulata was very low, only 0.07. The overlap between N. polita and N. undata, which occupies the intermediate position on the shore is much higher, 0.29, indicating a con- siderable overlap. Populations of the 2 dominant nerites, N. reticulata and N. undata, are also somewhat segregated vertically, with N. undata being higher on the shore (3.9 +2.6m) than N. reticulata (3.0--2.5m), but the over- Jap between populations was substantial. Both occurred at virtually every tide level sampled. The maximum density of N. undata was in the 3.0 to 3.5m tide level, one zone above the maximum abundance of N. reticulata. The index of niche overlap was high, 0.35, reflecting the sub- stantial mixing of the populations. Most studies of density regulation of intertidal gastro- pods have been done on limpets. FRANK (1965) showed that the density of the limpet Acmaea digitalis Rathke, 1833, was limited by an interaction of space and food re- sources. This appears to be a general condition in limpets. Unperwoop (1975; 1976) found density regulation in Nerita atramentosa Reeve, 1855, to be a function of food competition. Significant natural mortality was only found in the adults of the population. Underwood suggested that if the density of N. atramentosa was increased above the critical level by excessive numbers of juveniles settling on the shore the growth rates of the juveniles would be slowed and adult mortality would increase until the popu- lation had returned to normal levels. Nerita atramentosa feeds on diatoms and algal spores which are thought to be replenished on each high tide (UNDERWOOD, 1975; 1976). The exact food of the 3 Nerita species of the Admi- ralty Gulf is unknown, but their radular structure is simi- lar to that of N. atramentosaand it is reasonable to suggest that food could also be a limiting factor for the Admiralty Gulf species. Food competition would be lessened by the differences in mean tidal height of the 3 species, which means that each is concentrating its feeding on a differ- ent shore level. Another possible factor in reducing com- petition is differences in the behavior of the species. This possibility has not been examined in the present study. Coteman (1976) reported on the activity patterns of 3 species of Nerita: N. albicilla Linnaeus, 1758; N. plicata Linnaeus, 1758, and N. polita. All 3 actively move about while immersed at high tide and during the initial stages of emersion as the tide falls. When the rocks dry during low tide, the snails retreat into sheltered areas and become quiescent. While the pattern of behavior of the 3 species is similar, the selection of sheltered areas varies between spe- cies: N. plicata congregates in crevices and hollows; N. al- bicilla remains in damp, shaded crevices; and N. polita tends to burrow into the sand. If similar behavioral differ- THE VELIGER Vol. 20; No. 3 ences are operating in the species found in the Admiralty Gulf they could influence the areas grazed by each species. A number of nerite species occurs along the rocky shores of the Western Australian coastline (WILSON & GILLETT, 1974). Nerita atramentosa is a cold water spe- cies which is widely distributed along the south coast and extends northward along the west coast as far as Point Cloates. Nerita albicilla, N. plicata, N. polita, N. reticu- lata, and N. undata are all tropical species which have a range extending across the entire tropical north coast of the state and range southwards on the west coast. The numerous possible interactions between these 6 species offer a wide range of interesting avenues of investigation. Possible behavioral differences and mechanisms of density regulation have already been mentioned. Varying com- binations of the nerites occur at various localities along the northwestern coasts of the state. It would be interesting to investigate the populations of several species for evi- dence of character displacement. FENCHEL (1975) has shown that character displacement occurs in the gastro- pod genus Hydrobia where mean individual sizes of the species change when 2 or more species are present. A similar factor could be operating in Nerita: N. undata averaged 1.4 times the mean shell length of N. reticulata, and the average N. polita was 1.2 times as large as N. undata. The sizes of all 3 species in the Admiralty Gulf were well below the maximum sizes reported by both CERNOHORSKY (1972) and Wison & GmteTT (1974). A second possible mechanism of character displacement is a change in the mean shore level of a nerite species when it occurs with one or more other nerites. Particularly interesting in this regard is N. atramentosa which is the only nerite on most shores, where the population is cen- tered in the low intertidal zone (PrmEAUx, 1976). It would be interesting to see if the population of N. afra- mentosa is forced to lower shore levels by competitive interaction with other nerites in areas where it is sym- patric with other species of Nerita. ACKNOWLEDGMENTS An expedition such as this cannot succeed without assist- ance of all kinds from a wide variety of people too numer- ous to mention here. The Western Australian Museum participation was made possible by a special grant from the Treasury Department of the State of Western Austral- ia to the Western Australian Museum. The area studied by the expedition is on a mining lease owned by the ALUMAX Mining Company. ALUMAX provided in- valuable assistance to the expedition, particularly through Vol. 20; No. 3 their camp manager, Mr. Morris Marshall. Dr. B. R. Wilson headed the expedition and was largely responsible for its success. Mrs. L. M. Marsh read the manuscript and provided valuable criticism. Literature Cited Czrnonorsry, WALTER OLIVER 1972. | Marine shells of the Pacific, vol. 2, 411 pp.; 68 pits. Pacif. Publ. Sydney, Austral. Coxreman, Noe. 1976. Aerial respiration of nerites from the north-east coast of Austral- ja. Austral. Journ. Mar. Freshwater Res. 27: 455 - 466 Epwarps, Daias Craic 1969. Zonation by size as an adaptation for intertidal life in Olivella biplicata. Amer. Zool. 9: 399 - 417 ENpEAN, Rosert W., W. STEPHENSON & R. KENNY 1956. The ecology and distribution of intertidal organisms on certain islands off the Queensland coast. Austral. Journ. Mar. Freshwater Res, 7: 317 - 342 FgencHEL, Tom 1975. | Character displacement and coexistence in mud snails (Hydro- biidae). Oecologia (Berlin) 20: 19-32 Frang, Peter WoLFcANo 1969. | Growth rates of some gastropod molluscs on the coral reef at Heron Island. Oecologia (Berlin) 2: 232 - 250 Hopokrin, Ernest P, Loisette M. Marsg « G. G. Suir 1955. Rottnest Island: ‘The Rottnest Biological Station and recent scientific research. 19. The littoral environment of Rottnest Island. Journ. Roy. Soc. West. Austral. 42: 85 - 88 THE VELIGER Page 287 Horn, Henry S. 1966. Measurement of “overlap” in comparative ecological studies. Amer. Nat. 100: 419 - 424 MarsH, Loisetre M. # Ernest P Hopoxkin 1962. A survey of the fauna and flora of rocky shores of Carnac Ie- land, Western Australia. West. Austral. Nat. 8: 62 - 72 Morysrra, M. 1959. Measuring of interspecific association and similarity between communities. Mem. fac. Sci., Kyushu Univ. Ser. E (Biol.) 3: 65-80 PripEAuXx, PETER 1976. The role of competition in the structure of communities of proso- branch gastropods at Strickland Bay, Rottnest Island. B. Sci. Thesis, Univ. West. Austral., 47 pp. Remar, A. A. 1976. Description of a Tetraclita stalactifera panamensis community on a rocky intertidal Pacific shore of Panama. Mar. Biol. 35 (3): 225 - 238 Simpson, E. H. 1949. Measurement of diversity. STEPHENSON, T. A. & ANNE STEPHENSON 1949. The universal features of zonation between tide-marks on rocky coasts. Journ. Ecol. 37: 289 - 305 Unpgrwoop, ANTHONY J. 1972. | Tide-model analysis of the zonation of intertidal prosobranchs. II. Four species of trochids (Gastropoda; Prosobranchia). Journ. exp. mar. Biol. Ecol. 9: 257 - 277 1975. | Comparative studies on the biology of Nerita atramentosa Reeve, Bembicium nanum (Lamarck) and Cellana tramoserica (Sowerby) (Gastropoda; Prosobranchia) in S. E. Australia. Journ. exp. mar. Biol. Ecol. 18: 153 - 172 1976. Food competition between age-classes in the intertidal neritace- an Nerita atramentosa Reeve (Gastropoda; Prosobranchia). Journ. exp. mar. Biol. Ecol. 23: 145 - 154 Wison, Barry R. & Kerra Gitett 1974. Australian Shells. 168 pp.; illus. Sydney, Australia (A. H. & A. W. Reed) Nature, London 163: 688 Page 288 THE VELIGER Vol. 20; No. 3 A New Species of Serpulorbis (Gastropoda : Vermetidae) from South Africa BY ROGER N. HUGHES Department of Zoology, University College of North Wales, Bangor, Gwynedd, LL57 2UW, U. K. (4 Text figures) INTRODUCTION In DECEMBER 1975, a new species of Serpulorbis was dis- covered living on stones within the mouth of the Umnga- zana estuary near Port St. Johns, Transkei and in large rock pools on the open coast outside the estuary. The species is named Serpulorbis (Serpulorbis) aureus after the golden yellow colour of the exposed parts of the living animal. The taxonomy is based on KEEN (1961). DESCRIPTION Taxonomy Genus Serpulorbis Sassi, 1827 Subgenus (Serpulorbis) Keen, 1961 Serpulorbis (Serpulorbis) aureus Hughes, spec. nov. Morphology SHELL The adult shell (teleoconch) is cemented to the substra- tum forming a dextral spiral with loose concentric whorls. The whorls are approximately triangular in cross-section with the flat base cemented to the substratum. Their surface is textured with low axial ridges which are sig- moid in lateral aspect (Figure 1a). A few individuals possess faint longitudinal ridges. The outer whorl is ex- tended into a vertical feeding tube circular in cross- section. Scars of old feeding tubes are evident at every quarter or half whorl. The holotype is 2.5cm across its outer whorl, the feeding tube is 1.5cm long with an in- ternal diameter of 4mm. Most adults are approximately of this size. The protoconch is about 1mm long, mostly clear but with a brown tinge to the dorsal lip or hood of the aper- ture. The apical whorl is almost as large as the basal whorl (Figure 2a). ADULT BODY The animal reaches a length of about 4cm. The shapes and relative proportions of the various organs are shown in Figure 3a. Brooding females have a deep dorsal cleft in themantle (Figure 3b) toaccommodate the egg capsules which hang from the roof of the shell. The head, foot and exposed part of the mantle have a golden yellow back- ground colour which fades after fixation. The mantle edge is coloured with alternate bands of golden yellow and dark brown. In many individuals the foot and dorsal parts of the head are dark brown. EGG CAPSULES anp EMBRYOS A brooding female contained 20 egg capsules attached to the roof of the shell in 2 staggered ranks (Figure 2c) from the base of the feeding tube to half way along the outer whorl. Older capsules each contained about 30 ad- vanced embryos. Younger capsules contained eggs with a distinctive pale green yolk. The green colour is lost on fixation. The advanced embryos had well developed pro- toconchs and were clearly destined for benthic life after hatching. RADULA The radula, shown in Figure 4a, has a row formula of 2°1:‘1°'1°2. The rachidian tooth has convex sides while the inner marginal tooth has 2 or 3 well developed lateral Cusps on its outer edge and 2 lateral cusps on its inner edge. Vol. 20; No. 3 THE VELIGER Page 289 Figure 2 (a) Protoconchs from left lateral aspect (b) Protoconchs from dorsal aspect In both figures the larger protoconch is of Serpulorbis natalensis and the smaller protoconch is of Serpulorbis aureus (c) Serpulorbis aureus egg cases attached to the roof of the adult shell Fi ee DIAGNOSIS Serpulorbi. hol from | J (oy Serre “(by Faas ae oe sae Genus Serpulorbis: lack of operculum in adult; bright (c) Serpulorbis natalensis pigmentation of exposed body. Figure 3 Serpulorbis aureus (a) The mantle slit has been opened further to expose the contents of the mantle cavity (b) The intact female showing the dorsal slit in the mantle through which the egg capsules hang cm — columellar muscle ct — cephalic tentacle f — foot g - gill me — mantle edge ms — mantle slit o — osphradium Pp — proboscis r — rectum vm — visceral mass Subgenus (Serpulorbis): shell not tightly planorboid throughout life. Species aureus: adult shell with concentric whorls and vertical feeding tube; protoconch clear, apical whorl al- most as large as basal whorl; rachidian tooth with con- vex sides, inner marginal tooth with at least 2 lateral THE VELIGER Vol. 20; No. 3 Figure 4 Radular teeth of (a) Serpulorbis aureus (b) Serpulorbis natalensis R -rachidian 1 - lateral 2 -— inner marginal (marginal 1) 3 — outer marginal (marginal 2) 2" — lateral aspect of inner marginal showing cusps cusps on each side; exposed body with golden yellow background colour; embryos with pale green yolk. TYPE MATERIAL British Museum [Natural History], Department of Zoo- logy. Holotype 1976/W/1. Embryos 1976/W/2. Late embryos 1976/W/3. TYPE LOCALITY On stones at side of main channel, southern shore, mouth Vol. 20; No. 3 THE VELIGER Page 291 Table 1 Serpulorbis aureus Serpulorbis natalensis Body up to 4 cm long. Foot, head and mantle with golden yellow background colour. Yolk pale green. Sides of rachidian tooth convex. Inner marginal tooth with more than one cusp on each edge. Protoconch mostly clear, apical whorl almost as large as basal whorl. Teleoconch with dextral, usually concentric spirals triangular in cross-section, vertical feeding tube circular in cross-section. Feeding tube scars evident. Sculptured with axial ridges sigmoid in lateral aspect. Body up to 6 cm long. Background colour orange. Yolk pale yellow. Sides of rachidian tooth concave. Inner marginal tooth with one or no cusp on each edge. Protoconch pale brown, apical whorl much smaller than basal whorl. Teleoconch vermiform or with loose laterally displaced spirals circular in cross-section. No feeding tube. Sculptured with longitudinal ridges and fine axial striae. of Umngazana estuary near Port St. Johns, Transkei, South Africa, 32°S; 294° E. HABITAT Serpulorbis aureus forms loose aggregations on the tops of large silt-covered stones (HUGHES, in press a, fig. 2) at the edge of the main tidal current in the Umngazana estuary. They are not uncovered at low tide. Isolated individuals were also found in large tide pools at mid to high tide level outside the estuary. The feeding tube raises the head away from the silty substratum and thus avoids excessive clogging of the mucous net which is used for feeding. The shell whorls of the estuarine individuals were covered with short algae matted with silt. DISCUSSION It is possible that Serpulorbis aureus has been confused previously with S. natalensis (Morch, 1862) which is an abundant species in more southerly waters of the Cape Province (HuGHEs, in press b). Anatomical differences between the two species are summarized in Table 1, Figures 1a, 1b, 2a, 2b and 4a, 4b. The anatomy of the soft parts shows little variation within the genus (Morton, 1951). Serpulorbis aureus and S. natalensis differ in mi- crohabitat, S. aureus colonising the upper surfaces of stones in turbid water whereas S. natalensis usually colon- ises the under surfaces of stones where silting is less heavy. Feeding tubes raising the head away from the substratum are not found in S. natalensis (personal observations). Serpulorbis aureus probably replaces S. natalensis as an ecological equivalent in the warmer waters of the South African Indian Ocean just as Dendropoma tholia Keen & Morton, 1960 replaces D. corallinaceum (Tomlin, 1939) (HucHESs, in press b). The coiled shell with vertical feeding tube of Serpul- orbis aureus Closely resembles that of S. squamigerus (Carpenter, 1857) of Californian waters, but the latter is distinguished by its geographical range and by its pre- dominantly black colour with orange pigmentation around the foot and lateral regions of the head (Hap- FIELD, 1970). ACKNOWLEDGMENTS I thank George Branch for inviting me on the expedition to Umngazana and for pointing out the colony of Ser- pulorbis aureus in front of our camp site. Literature Cited Haprizetp, Micwasgt G. 1970. Observations on the anatomy and biology of two California ver- metid gastropods. The Veliger 12 (3): 301-309; plt. 45; 5 text figs. (1 January 1970) Huocues, Rooer N. in press a. Colonialism in Vermetidae. In: Biology and systematics of colonial organisms. Spec. Vol. Syst. Assoc. in press b. The biology of Dendropoma corallinaceum and Serpulorbis natalensis, two South African vermetid gastropods. ZooJ. Journ. Linn. Soc. Keen, A. Myra 1961. A proposed reclassification of the gastropod family Vermetidae. Bull. Brit. Mus. (Nat. Hist.) (Zool.) 7: 183 - 213 Page 292 THE VELIGER Vol. 20; No. 3 A New Species of Coryphella (Nudibranchia : Flabellinidae) from Santa Barbara, California BY ROBERT K. COWEN Moss Landing Marine Laboratories, P O. Box 223, Moss Landing, California 95039 AND DAVID R. LAUR Department of Biology, University of California, Santa Barbara, Goleta, California 93016 (2 Plates) On Marcu 25, 1973, 2 nudibranch specimens were col- lected by David Laur while diving in 20m of water, 1.6 km off Arroyo Burro County Beach, Santa Barbara, Cali- fornia. At first they appeared to be color varieties of Cory- phella iodinea (Cooper, 1863) ; however, upon closer ex- amination they were found to have many unique traits. During the ensuing 2 years, several more individuals were found. Preliminary investigations carried out by Robert Cowen yielded promising results, thereby initiating the following study. On April 16, 1975, 25 specimens were collected in 18 to 24m of water, 1.3 to 1.7km offshore Arroyo Burro County Beach by Craig Fusaro, Shane Anderson and the the senior author. Comparison of these specimens with Coryphella iodinea were made using starch-gel electro- phoresis, scanning electron micrographs of the radula, and morphometric counts. From our results (given in the discussion) we propose the following taxon. FLABELLINDAE Bergh, 1890 Coryphella Gray, 1850 Coryphella sabulicola Cowen & Laur, spec. nov. Description: Body typical flabellinid shape, iaterally compressed, elongate, tapering posteriorly. Body length, excluding oral tentacles, 46 to 55mm, average length 50.5 mm. Length of oral tentacles 17 to 23mm, average length 19.25 mm. Body pale bluish-purple in color, foot bordered with white (Figure 1). Basal third of oral tentacles same color as body, distal two-thirds white. Base of cerata same as body color, remainder pale orange. Rhinophores red- brown in color. Cerata in 8 to 10 groups per side, each arising from single ridge-like processes. Anterior-most group separated Explanation of Figures 7, 6 Figure 1: Coryphella sabulicola Cowen & Laur, spec. nov.; photo- graph by Dave Laur Figure 6: Example of starch-gel electrophoresis results. Starting point indicated by arrow. Cathode-attracted proteins (e. g., present in Coryphella iodinea), are below the starting point. Odd numbered bands are C. iodinea, even numbered bands are C. sabulicola. THE VELIGER, Vol. 20, No. 3 [Cowen « Laur] Figures 7, 6 Figure 1 —_ = —= == == ——-s ——= —. —— S eenamnad Seamed a eS ad cd a eS eh aa 1 2 3 4 5 6 7 8 9 10 11 12 Vol. 20; No. 3 from remainder by cardiac projection. Number of cerata per group decreases from anterior to posterior. Total num- ber of cerata per side 67 to 82, mode 75. Rhinophores perfoliate with mode of 50 plates. Radular formula 16 (1:1:1). Rachidians triangular in shape with deep U-shaped base, bearing 11 to 13 denticles per side (Figure 2). Median denticles about twice width of side denticles. Laterals roughly triangular in shape with strong central cusp and 13 to 16 equal den- ticles on inner margin; inner margin 2 length of outer margin (Figures 3, 4). Laterals set at angle to rachidians; laterals of rows 10 to 16 folded over rachidians and within sheath. Type Locality: 24m depth, approximately 1.4km SSW of Arroyo Burro County Beach, Santa Barbara, Califor- nia (34°24’N; 119°44’W). Type Disposition: Holotype - Invertebrate Zoology Type Collection No. 700, California Academy of Sciences, San Francisco, California. Paratypes: submitted to the California Acad- emy of Sciences for distribution to other institutions. Habitat: Known from sand and silt bottom with no rock relief, in 15 to 29m of water, off Arroyo Burro Coun- ty Beach. Several specimens also have been dredged 3.2 to 4.8km off Santa Barbara Harbor from a depth of 50m. Discussion: Coryphella sabulicola differs from C. iodin- ea by its paler colors and white foot border and its habitat preference of deep sand and silt bottom rather than shal- THE VELIGER Page 293 lower rocky relief or pier piling preference of C. iodinea. Our study yielded further differences. A morphometric comparison of 8 specimens of each species was made. The cerata were counted on the left side, and additional counts and measurements were taken which included: total length, oral tentacle length, number of cerata groups, length of longest and shortest cerata. Means and variances were calculated and a Student’s t-test was applied (Table 1). The difference in cerata number between the 2 species, relative to length, was significant at the 0.001 level. None of the other morpho- metric measurements proved significant. The heads of starved animals were used to test protein differences by starch-gel electrophoresis. Twelve animals of each species were tested, utilizing a general protein stain. In 11 Coryphella iodinea tested, there was a cathode attracted protein which was absent in all 12 C. sabuli- cola (Figure 6). A study of the radulae of 3 animals of each species by scanning electron microscope did not reveal any ob-. vious differences, yet some trends are evident. In Cory- phella sabulicola the rachidians have 11 to 13 denticles per side, while in C. iodinea the number of denticles ranges from 12 to 14 per side. The range of denticles on the lateral teeth, though more variable, again was smaller in C. sabulicola than in C. iodinea, 13 to 16 and 13 to 18, respectively (Figures 2 to 5). The results of the above study, in conjunction with the color and habitat differences, strongly support the desig- nation of Coryphella sabulicola_as a species distinct and Table 1 Cerata count results. Cerata counted on left side only. Body length (mm) Number of Cerata Animal Coryphella Coryphella Coryphella Coryphella Number sabulicola todinea sabulicola iodinea 1 52 53 81 110 2 50 55 74 96 3 46 50 81 91 4 46 46 67 94 5 55 49 73 78 6 53 51 82 102 7 52 51 71 103 8 50 56 74 106 xX 50.5 51.4 75.4 97.5 SZ 29.4 102.3 a t = 11.82 p 0.001 THE VELIGER Page 294 separate from C. iodinea. Additional work on the eco- logical differences between these 2 species of Coryphella should yield further support for this contention. Coryphella sabulicola is named with reference to its sand-dwelling habit. ACKNOWLEDGMENTS We would like to thank Dr. Demorest Davenport and Dr. FE G. Hochberg for critically reading this manuscript; Lou Haldorson and Dr. Jack King for their help with the electrophoresis; Dr. Preston Cloud for the use of his lab- oratory’s scanning electron microscope and David Pierce for operating it. Chris Kitting helped with the prelimi- nary study. Cathy Engle’s help and interest is also ap- preciated. Explanation of Figures 2 to 5 Figures 2 to 4: The radula of Coryphella sabulicola Cowen & Laur, spec. nov.; scanning electron micrographs by David Pierce Figure 2: First and second rachidians showing base and denticles X 400 Figure 3: Center of radula showing position of laterals relative to rachidians X 260 Figure 4: Rachidians and laterals showing denticles X 260 Figure 5: The radula of Coryphella iodinea (Cooper, 1863). Center of radula showing denticles on laterals and rachidians X 260 Vol. 20; No. 3 Tue VELIcER, Vol. 20, No. 3 [Cowen « Laur] Figures 2 to 5 Vol. 20; No. 3 THE VELIGER Page 295 A New Species of Anachis (Gastropoda : Columbellidae) from the Eastern Pacific R. A. WHITNEY 2140 North Main Street, Decatur, Illinois 62526 (2 Text figures) I HAVE EXAMINED numerous specimens of Anachis over the past year, and have encountered a species of Anachis from the Eastern Pacific region which has not been previ- ously described. This species is seen with some frequency in several of the larger institutional collections as well as in private collections. Through the interest and generosity of the Los Angeles County Museum of Natural History, I have been able to examine a number of this Anachis species, and locality records as well as geographical range can be reported here. These are recorded here immediately following the listing of paratypes. I have assigned the new species to the genus Anachis, and, in order not to add confusion to the taxonomic problems in the family Columbellidae, I have avoided the use of other supraspecific taxa, leaving this to a future investigator who might make the necessary anatomical studies to complete a monograph on the Co- lumbellidae of the Eastern Pacific. Anachis lillianae Whitney, spec. nov. (Figures 1, 2) Description: Size small; color white with light to deep brown markings; whorls somewhat convex with body whorl inflated; axial sculpture predominant over entire shell; nucleus of 2} glassy whorls; 6 post-nuclear whorls; axial ribs wider than interspaces with ribs on penultimate and earlier whorls having keel-like ridge running down center of each rib; keeled appearance not so much in evidence on body whorl as ribs become less pronounced and more rounded; intercostal striae between ribs imping- ing on ribs but not crossing center of ribs; 15 ribs on body whorl; basal portion of body whorl has spiral cording; parietal wall of aperture smooth; outer lip of aperture with 6 weak denticles inside lip; deep-brown band ante- rior to suture forms white subsutural band; deep-brown band broken, giving irregular zig-zag appearance; base of shell marked with irregular cloudings, blotches, and streaks of light brown; anterior canal short; length 7mm; diameter 3mm; 6 whorls plus nucleus. Holotype, Los Angeles County Museum of Natural History Type No. 1852. Dorsal view xX 1g Holotype: Los Angeles County Museum of Natural History, Type Collection No. 1852. Type Locality: All specimens of the type material were collected at Playa Alicia near San Felipe, Baja Califor- nia, Mexico, under rocks at low tide, March 1976. Paratypes: Los Angeles County Museum of Natural History Type collection no. 1847; Academy of Natural Sciences, Philadelphia, Type Collection no. 344375; Douglas and Sherry Welker collection no. 6; R. A. Whit- ney collection no. 37. Page 296 Figure 2 Holotype, Los Angeles County Museum of Natural History Type No. 1852. Ventral view X 1g Other Material Examined: In addition to the type lot — Los Angeles County Museum of Natural History no. 67- 17 - Libertad, Sonora, Mexico; no. 11833 -San Blas, Nayarit, Mexico; no. 11834 - Bahia Adair, Sonora, Mexi- co; no. 63-56 - Puerto Penasco, Sonora, Mexico; no. 30- 74 - Mazatlan, Mexico; no. 59-9 - Topolobampo Bay, Mexico; all intertidal. Also no. 72-58 - 30-75 ft. (9 - 22.5 m), rocky, small islets off Punta Quepos, Puntarenas Province, Costa Rica, 9°52’43” N; 84°09/41” W. Discussion: There is little variation among the speci- mens examined of this species. The shell has a striking color pattern and varies in the number and intensity of brown blotches that comprise the subsutural bands. The variation in color markings is especially marked at the base of the body whorl, where the brown markings may appear as streaks, dots, or cloudings. The number of axial ribs on the body whorl varies from 14 to 17. Anachis lillianae bears some superficial resemblance to other species which should be considered in making identifications. A very closely related species is A. adelinae (Tryon, 1883). This species differs from A. lillianae in THE VELIGER Vol. 20; No. 3 having a distinct checkerboard pattern of brown spots on the base of the body whorl, and in the spire being more attenuated. Anachis albonodosa (Carpenter, 1857) is light brown with white spots showing at the ends of the axial ribs; it is a somewhat smaller shell, with the body whorl being less inflated than A. lillianae. Anachis dalli Bartsch, 1931 is pale yellow with 2 zones of interrupted brown spots and 1 of white; A. pygmaea (Sowerby, 1832) has a distinct pattern of 2 or 3 rows of brown spots; A. diminuta (C. B. Adams, 1852) could possibly be mistaken for an immature A. lillianae, but the brown shell of A. diminuta is 4mm or less in size. The new species is named in honor of Mrs. Lillian Whitney of Decatur, Illinois, in appreciation of her en- couragement of, and assistance in, the author’s study of Columbellidae. ACKNOWLEDGMENT I am sincerely grateful for the generous cooperation given me over a period of many months by the individuals who have contributed so much of their time towards the com- pletion of this paper. I wish to express my thanks to Dr. James H. McLean of the Los Angeles County Museum of Natural History for his advice, and to Mr. Gale Sphon, also of the Los Angeles County Museum, for arranging a loan of museum specimens for comparative studies. I also wish to acknowledge the excellent photographs of the holotype by Mr. Bertram C. Draper, Museum Associ- ate, Los Angeles County Museum of Natural History. Literature Cited Apvams, Cmarizes Baker 1852. Catalogue of shells tala ee at Panama with notes on synonymy, station and habitat. Ann. Lyc. Nat. Hist. New York 5: 229 - 296 (June) ; 297-549 (July) eae (1852) Catalogue of shells col- lected at Panama, with notes on their synonymy, station, and geo- graphical distribution. New York (Craighead): viii+334 pp.] [see aleo FT ouea! 1956] Bartscz, Paur 1931. Descriptions of new marine. mollusgs from Panama, with a figure of the genotype of Engina. Proc. U. S. Nat. Mus. 79 (2881): 1-10; 1 plt. (1 August 1931) Carpenter, PHitip PEARSALL 1857. Catalogue of the collection of Mazatlan shells in the British Museum, collected by Frederick Reigen, ... London (British Museum). iv+ix-xvi+552 pp. (Preface by J. E. Gray) (1 August). Warrington (Oberlin) edition: viii+xii+552 pp., with author’s preface, published simultaneously [dating: Carpenter, 1872: xi; Irepave, 1916; SHer- BORN, 1934] [reprinted by Paleo. Res. Inst., Ithaca, New York, 1967] SoweErsy, GEORGE BRETTINCHAM 1832-1833. Characters of new species of Mollusca and Conchifera, col- lected by Hugh Cuming. Proc. Zool. Soc. London (for 1832): 25-48 (21 April 1832); 49-76 (5 June); 77-108 (31 July); 109 to 120 (14 August); 173-179 (14 January 1833); 194-202 (13 March 1833). [some of the species were described by W. J. Broderip] Tryon, Gzorcze WASHINGTON, Jr. 1883. Manual of Conchology. Philadelphia, vols. 1-17 Vol. 20; No. 3 NOTES & NEWS Nomenclatural Notes on Hinnites giganteus (Gray) BY BARRY ROTH! anp EUGENE V. COAN THE COMMON ROCK SCALLOP, or “purple-hinged scallop,” of the Pacific Coast appears in recent literature under two names: Hinnites giganteus (Gray, 1825) and Hinnites multirugosus (Gale, 1928). GALE (1928: 92) introduced the latter because he believed that Gray’s name and other possible substitutes were homonyms and therefore unavail- able. Other authors (ADAM, 1960; HERTLEIN & GRANT, 1972) have given reasons for preferring use of the older name, H. giganteus. The valid name of this taxon depends on two related factors: (a) the availability, or not, of Gray’s name, and _ (b) the nature and validity of Gale’s rejection of it — both points to be evaluated in light of the ICZN rules governing such cases. From a review of the relevant literature, we have come to the following conclusions: 1. Ga.e’s (1928) stated reasons for proposing “Pecten (Chlamys) multirugosus’ —a new taxon, not a simple renaming — and his remark that Lima gigantea Gray was “stillborn” are erroneous. Lima gigantea Gray (1825: 139), as the species was first named, is the earliest use of this combination of generic and specific names. It is not a primary homonym of Plagiostoma giganteum J. Sower- by, 1814, whether or not Lima Bruguiére, 1797, and Pla- giostoma J. Sowerby, 1814, are considered synonymous genera of the Limidae. Grau’s (1959) claim that Plagio- stoma was proposed as a subgenus of Lima is incorrect. Moreover, Gray (1826) removed his gigantea from Lima and placed it in “Hinnita” (an invalid emendation of Hinnites Defrance, 1821), and the species has not been reallocated to Lima since that time. We have located no citation of Sowerby’s giganteum in the genus Lima prior to those by DesHayEs (1831, 1832), although the possible synonymy of Lima and Plagiostoma was being debated as early as 1823 (G. B. Sowersy, 1823). Modern workers consider the two genera separable. ' Department of Geology, California Academy of Sciences, San Francisco, California 94118 THE VELIGER Page 297 As Apam (1960) noted, Gray’s and Sowerby’s names were not secondary homonyms at the time of Gatez’s (1928) publication. It is evident that Gale himself did not believe the two species to be congeneric, since he cited Sowerby’s giganteum as a Lima but described multi- rugosus as a species of Pecten. Under ICZN rules [Art. 59(b) (2)], Gray’s name required no replacement then and requires none today. 2. Jay (1835) and many subsequent authors placed Gray’s gigantea in the genus Pecten Miller, 1776. While several other pectinid taxa share the specific epithet “gi- ganteus,” the earliest of these is Pecten giganteus Miin- ster in Goldfuss, 1833. The gigantea of Gray (1825) is therefore not a junior secondary homonym in Pecten. 3. Gate (1928) specifically stated that his Pecten (Chla- mys) multirugosus was proposed as a new species, and not the simple renaming of a homonym: “Pecten multiru- gosus is virtually a new name for the common Pliocene to Recent West Coast species formerly known as Pecten (Hinnites) giganteus (Gray); but in order to avoid any questions about the location or identity of the original types, the species is described as new and a new type is cited” (GALE, 1928: 92; emphasis supplied). In contrast, “Pecten (Chlamys) multirugosus var. crassiplicatus,” a renaming of the homonymous Hinnites crassa Conrad, 1857, by Gale in the same paper, was unequivocally pro- posed as a replacement name. 4. In summary, Gray’s (1825) name gigantea was available when proposed and has remained available throughout its nomenclatural history. GALE’s (1928) re- jection of it was invalid because he did not contend that the two species-group taxa, “Lima” gigantea Gray and Plagiostoma giganteum Sowerby, were congeneric. Pecten (Chlamys) multirugosus Gale, 1928, is therefore a junior synonym of Hinnites giganteus (Gray, 1825), and the latter is the valid name for the Pacific Coast rock scallop. Literature Cited Apam, WILLIAM 1960. A propos de Chlamys (F. acc. Hinnites) abscondita (P. Fischer, 1898) de la céte occidentale de l’Afrique.. Bull. Inst. Roy. Sci. Nat. Belg. 36 (20): 1-10; 2 plts. (February 1960) DeswHayes, GERARD PAUL 1831. Description des coquilles caractéristiques des terrains. Paris EG. Levrault. vii+264 pp.; 14 plts. 1832 (1830-1832). Encyclopédie méthodique, histoire naturelle des vers. Vol. 2, prt. 2. Paris, Agasse. 594 pp. [pp. 1-144 publ. 6 February 1830; pp. 145-594, containing Lima, publ. 29 Sept. 1832; fide SHerporn, C. D. « B. B. Woopwaarp, 1906, Ann. Mag. Nat. Hist. (7) 17: 579] Gare, Hoyt RopNry 1928. West coast species of Hinnites. Trans. San Diego Soc. Nat. (29 February 1928) Hist. 5 (9): 91-94 Allan Hancock Pac. Ex- Grau, GitBERT 1959. Pectinidae of the eastern Pacific. (25 September 1959) ped. 23: i- viii, 1- 308; pits. 1-57 Page 298 Gray, JonN Epwarp 1825. A list and description of some species of shells not taken notice of by Lamarck. Ann. Philos. (n.s.) 9: (2) (art. 9): 134-140; 1 text fig. (February 1825) 1826. On a Recent species of the genus Hinnita of DeFrance, and some observations on the shells of the Monomyaires of Lamarck. Ann. Philos. (n.s.) 12 (2) (art. 4): 103-106 Hertuzin, Leo Gzorocgz « Utysszs Simpson Grant, 1V 1972. The geology and paleontology of the marine Pliocene of San Diego, California. Part 2B: Paleontology, Pelecypoda. Mem. San Diego Soc. Nat. Hist. 2: 135-409; frontisp.; plts. 27-57; text figs. 7-13 (21 July 1972) (August 1826) Jay, Joun CLarKson 1835. Catalogue of Recent shells in the cabinet of John C. Jay. New York, D. Fanshaw. 55 pp. Sowersy, Grorce BRETTINGHAM !5t 1823 (1821-1834). The genera of Recent and fossil shells. 2 vols. Lon- don. 262 plts. & accompanying text (unpag.)[Part 17, incl. genus Lima, publ. about 20 August 1823, fide SHerporn, C. D., 1894, Ann. Mag. Nat. Hist. (6) 13 (76): 371) SowerBy, JAMES DE CARLE 1814 (1812-1815). The mineral conchology of Great Britain ... Vol. 1. London. 236 pp., 102 plts. [Part 14, pp. 169-178, plts. 74-78, publ. 1 December 1814, fide Syxes, E. R., 1906, Proc. Malacol. Soc. London 7 (3): 191) Publication Dates of Bergh’s 1879 Papers Describing American Chromodorids BY ROBERT BURN Honorary Associate, National Museum of Victoria Melbourne, Victoria, Australia 3000 THREE PAPERS CONTAINING descriptions of American chromodorid opisthobranchs were published by Rudolph BERGH in the year 1879. In an attempt to establish an order of priority for original descriptions, synonymies and future revisions, the publication dates of the 3 papers have been determined. According to internal evidence, 7. e., the numbering in the lists of Chromodoris species in the papers, Bergh wrote his papers in the sequence (1) Neue Nacktschnecken der Siidsee, (2) On the nudibranchiate gasteropod Mol- lusca of the North Pacific Ocean, with special reference to those of Alaska, and (3) Neue Chromodoriden. The papers are treated here in this order. (1) The 4" and last part of Bergh’s series “Neue Nacktschnecken der Siidsee” was published in part 14 of volume 5 of Journal des Museum Godeffroy. In reply to an enquiry to the Zoological Library, British Museum (Natural History), Miss A. Lucas informed me that their copy has “a list of other publications of the publisher THE VELIGER Vol. 20; No. 3 (L. Friederichsen & Co.), on the back cover, at the end of which are the words ‘Hamburg, im Februar 1879’. The other piece of evidence is the words ‘Zool. Dept. 28/3/79’ written faintly in pencil on the front cover — presumably the date of receipt here” (in litt. 19 June 1970). Thus it can be shown that the date of publication was probably not earlier than the printer’s date of February 1879, and certainly prior to 28 March 1879, the date of receipt at the B. M. (N. H.). In the absence of evidence to the contrary, it is possible to stipulate the last day of February, i. e., 28 February 1879, as the date of publica- tion. (2) Bergh’s “On the nudibranchiate gasteropod Mol- lusca of the North Pacific Ocean, with special reference to those of Alaska. Part I” appeared in 2 simultaneously published journals (W. H. Dat, in Bercn, 1879d: 125) ; RussEL1, 1968: 141). A printer’s date, 10 May 1879, appears at the foot of the first page of Part I. In the certi- fication notice in the title pages for the Proceedings of the Academy of Natural Sciences of Philadelphia, 1879, the then editor and recording secretary, Edward J. Nolan, reports that the pages containing Bergh’s article were presented at the meeting of the Academy on 13 May 1879. The closeness of these 2 dates, plus the certification Notice, suggests that it is reasonable to accept 10 May 1879 as the publication date. (3) There being no internal evidence to qualify the date of publication for Bergh’s “Neue Chromodoriden” more specifically than 1879, under I.C.Z.N. Article 21(b) the publication date would be 31 December 1879. However, external evidence indicates a much earlier date. In the ‘Literaturbericht’ of Nachrichtsblatt der deut- schen Malakozoologischen Gesellschaft 1879, the “Mala- kozoologische Blatter, herausgegeben von S. Clessin, Neue Folge, Bd. 1, Lfg. 1, Mit 3 Tafeln’ is reviewed article by article on pages 41 - 42. Page 42 has the following entry: ‘p. 77. Bergh, Dr. R., Notizen tiber Pleurophyllidia lo- véni.’ The last lines of Bergh’s Pleurophyllidia paper are printed on the same page (p. 87) as the title and first paragraphs of his “Neue Chromodoriden” (yet for some reason there is no entry in the ‘Literaturbericht’ for this latter paper). Plate 3, which belongs to “Neue Chromo- doriden”, is the 3" plate of the ‘Mit 3 Tafeln’ mentioned in the ‘Literaturbericht.’ It is thus demonstrable that these 2 papers by Bergh were published at the same time. Dates of publication of the parts of Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft 1879 are found in the printer’s notices at the end of each part. Page 48, the last page of ‘No. 2 & 3 Februar - Marz Vol. 20; No. 3 1879’, is dated ‘Marz 1879’. Under combination of I. C. Z.N. Articles 21(b) (i) and 21(f), the date of publica- tion for “Neue Chromodoriden” must be regarded as 31 March 1879. Literature Cited Beron, Lupwic Soruus Rupo.r 18792. Neue Nacktschnecken der Siidsee, malacologische Untersuchun- gen. IV. Journ. Mus. Godeffroy 5 (14): 1-50; pits. 1-5 (28 February 1879) Malakozool. Blatt., N. (31 March 1879) Malakozool. Blatt., N. FE 1: 87 - 116; pit. 3 -(31 March 1879) 1879d. On the nudibranchiate gasteropod Mollusca of the North Pacific Ocean, with special reference to those of Alaska. Part I. Proc. Acad, Nat Sci. Philadelphia 31: 71 - 132; plts. 1-8 (10 May 1879) 1879e. On the nudibranchiate gasteropod Mollusca of the North Pacific Ocean, with special reference to those of Alaska. Part I. Sci. Results Explor. Alaska 1; 127 - 188; plts. 1-8 (10 May 1879) RussgiL, Henry Drummonp 1968. Chromodoris californiensis and C. calensis. 81: (4): 140-141 1879b. Notizen tiber Pleurophyllidia lovéni. F 1: 77-87; plt. 2 1879c. Neue Chromodoriden. The Nautilus (April 1968) The Date of Publication of Anton’s “Verzeichniss der Conchylien” BY WALTER O. CERNOHORSKY Auckland Institute and Museum, Auckland, New Zealand ANTON’s IMPORTANT CONCHOLOGICAL work “Verzeichnif der Conchylien ...”, which besides descriptions of new taxa also contains generic type-species designations, has always been deemed to have been published in 1839, despite the note on page 110 “Im Druck beendigt den 9. Oktober 1838”, meaning that by that date the printing was completed. Irrefutable proof that Anton’s work already appeared in 1838 can be found in TRoscHEL (1839) who states: “Unter dem Titel: VerzeichniB der Conchylien, welche sich in der Sammlung von Hermann Eduard Anton be- finden, herausgegeben von dem Besitzer, Halle b. Eduard Anton 1839, erschien in der Mitte des Jahres 1838 ein Buch, das schon bei seiner Geburt sich um ein Jahr jiinger machte.” TROSCHEL (of. cit.), who every year reviewed mollus- can literature published during the previous year for the “Archiv fiir Naturgeschichte”, clearly states that Anton’s work appeared already in the middle of 1838 and that the THE VELIGER Page 299 book “made itself one year younger at the time of its birth”. The date 1839 on the title page is thus certainly an error and the date of 9. Oktober 1838 on page 110 a probable error which does not coincide with a middle of the year publication date. Both these dates may represent tentative completion dates. The acceptance of 1838 as the publication date for Anton’s “Verzeichni® der Conchylien ...” will change only a few specific name priorities and no major taxon- omic changes are expected. Literature Cited Anton, Hermann EpuARD 1838. Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Eduard Anton befinden. Halle, pp. i- xvi; 1-110 TroscHEL, Franz HERMANN 1839. V. Mollusca. Arch. f. Naturgesch. 5 (2): 201-241 Another Cephalopod from Northern California (Mollusca : Cephalopoda ) BY ROBERT R. TALMADGE ! Eureka, California 95501 Late mv January 1977, the crew and Captain James Riley of the drag boat Jna noted a rather unusual “squid” on the deck of the vessel after making a daylight tow in 330 fathoms (603m) on a soft (green) mud substrate, The tow was made at approximately 40°15’N, north of Eureka, California. The specimen was salvaged from amid the fish on deck and preserved on shipboard in a 10% solution of formalin. The specimen came to my hands and I deposited it in the “wet collection”, Department of In- vertebrate Zoology, California Academy of Sciences, San Francisco, California. Although this squid appeared to represent a species new to this part of the northern California coast, a possib- ly related species or variety has been taken a few times ' Field Associate, Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118 and Curator of Natural History, College of the Redwoods, Eureka, California 95501 Page 300 previously, Both species belong to the genus Calliteuthis. One species with an elongate body has been called C. hoylei (Pfeiffer, 1900), while the short-bodied form has been referred to C. meleagroteuthis Chun, 1910. Voss (1963) places the two taxa in synonymy. This note is presented in order to alert students of the Cephalopoda as to the whereabouts of this specimen. Based on my own observations, both species are uncom- mon in collections. Literature Cited Voss, GILBERT LINCOLN 1 Cephalopods of the Philippine Islands. Mus. 234: i-v+1- 180; 35 text figs. Bull. U. S&S Nat. W.S. M. THE 1978 MEETING of the Western Society of Malaco- logists will be held at the University of Santa Clara, near San Jose, California, June 28 through July 1. Presentation of papers concerning mollusks and other invertebrates will be complemented by field trips, work shops, and dis- plays. A symposium concerning current studies of Ameri- can land mollusks is also scheduled. A second symposium of research dealing with freshwater mollusks is probable. Information about the meeting is available from Dr. Peter Neal D’Eliscu, President, W.S.M., Department of Bio- logy, University of Santa Clara, Santa Clara, CA 95053. PN Ze: THE AMERICAN Society of Zoologists, Society of System- atic Zoology and the American Microscopical Society will meet Wednesday, December 27 through Saturday, De- cember 30, 1978 at the Hotel John Marshall in Richmond, Virginia. A call for contributed papers will be issued in April, 1978. For more information write to: Ms. Mary Wiley, American Society of Zoologists, Box 2739 Califor- nia Lutheran College, Thousand Oaks, CA 91360. THE VELIGER Vol. 20; No. 3 Late News Rate Increases: Because of continued inflationary pressures, it has become necessary to increase subcription rate and membership dues effective with volume 21. The subscription rate will be U. S. $30.00 plus postage, while membership dues will be U.S. $15.00 plus postage charges. For the time being, that is, until the U.S. Postal Service further increases postal rates, our charges will remain as follows: $1.00 for all domestic addresses, $2.50 for addresses in the so-called P. U.A.S. countries, and $3.50 in all other countries, in- cluding Canada. A more detailed announcement will be included with the invoices sent to subscribers. Subscrip- tion agencies have already been informed, those in foreign countries by air mail. Sale of C. M. S. 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WE ARE PLEASED to announce that an agreement has been entered into by the California Malacozoological Society, Inc. with Mr. Steven J. Long for the production and sale of microfiche reproductions of all out-of-print editions of the publications of the Society. The microfiches are available as negative films (printed matter ap- pearing white on black background), 105mm x 148mm and can be supplied immediately. The following is a list of items now ready: Volume 1: $1.50 Volume 6: $4.50 Volume 2: $3.00 Volume 7: $6.00 Volume 3: $3.00 Volume 8: $6.00 Volume 4: $4.50 Volume 10: $9.00 Volume 5: $4.50 Volume 11: $9.00 Volume 12: $9.00 Supplement to Volume 6: $1.50; to Volume 18: $3.00 California residents please add the appropriate amount for sales tax to the prices indicated. Page 302 Please, send your order, with check payable to Opistho- branch Newsletter, to Mr. Steven J. Long, P. O. Box 243, Santa Maria, CA 93454. 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CALIFORNIA Matacozootocicat Society, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. Donors may designate the Fund to which their contribution is to be credited: Operating Fund (available for current production) ; Savings Fund (available only for specified purposes, such as publication of especially long and signi- ficant papers); Endowment Fund (the income from which is available. The principal is irrevocably dedicated to scientific and educational purposes). Unassigned dona- tions will be used according to greatest need. Contributions to the C. M.S., Inc. are deductible by donors as provided in section 170 of the Internal Revenue Code (for Federal income tax purposes). 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The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. Endowment Fund In the face of continuous rises in the costs of printing and labor, the income from the Endowment Fund would materially aid in avoiding the need for repeated upward adjustments of the membership dues of the Society. It is the stated aim of the Society to disseminate new infor- mation in the field of malacology and conchology as widely as possible at the lowest cost possible. At a Regular Membership meeting of the Society in No- vember 1968 a policy was adopted which, it is hoped, will assist in building up the Endowment Fund of the Society. An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 000.- or more, an entire volume will be dedicated to the memory of the decedent. REGARDING POSTAL SERVICE Complaints regarding late arrival of our journal are in- creasing in number, steadily, continually. However, we very conscientiously dispatch our journal on the printed publication dates. What happens after deposition at the Post Office is, of course, beyond our control. From some of our members we have been able to construct a sort of probable delivery schedule. In general, within California, Page 304 8 days is usual; outside of California, the time lapse in- creases with the distance; the East Coast can consider a lapse of “only” two weeks as rapid service; 4 to 5 weeks are not uncommon. Foreign countries may count on a minimum of one month, six weeks being the more usual time requirement and over two months not rare! 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This, however, does not seem quite fair to us as some of our subscribers in almost 20 years have never failed to receive their copies. On July 6 the rates for book parcels and the library rate have been increased. This necessitates that we must in- crease the postage charges on back volumes, supplements and individual back numbers. The charges stated must be THE VELIGER Vol. 20; No. 3 increased by 20¢ for one item and by 8¢ for each addi- tional item. It has been announced by the Postal Service that no increase in postage rates would be asked during the cur- rent year; it was not stressed sufficiently that this applies only to the so-called first class mail. Second class mail rates are “phased,” that is, they are scheduled to be in- creased each year until the rates are sufficiently high to pay the actual cost of handling that type of mail. 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But at the same time, we wish to call the attention to our Endowment Fund, the income from which enables us, in part, to keep these charges at the established levels. Contributions (tax deductible in the U.S. A.) are always welcome. Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. That the printed date is the actual date of pub- lication under the rules of the International Commission on Zoological Nomenclature is based on the following facts: 1) The journal is delivered to the Post Office on the first day of each quarter, ready for dispatch; 2) at least three copies are mailed either as first class items or by air mail; 3) about 20 copies are delivered in person to the mail boxes or to the offices of members in the Berkeley area; 4) two copies are delivered to the re- ceiving department of the General Library of the Univer- sity of California in Berkeley. Thus our publication is available in the meaning of the Code of the ICZN. The printed publication date, therefore, may be relied upon for purposes of establishing priority of new taxa. To Prospective Authors Postal Service seems to have deteriorated in many other countries as well as in the United States of America. Since we will absolutely not publish a paper unless the galley proofs have been corrected and returned by the authors, the slow surface mail service (a minimum of 6 weeks from European countries, 8 to 12 weeks from India and Africa) may make a delay in publication inevitable. We strongly urge that authors who have submitted papers to the Veli- ger make all necessary arrangements for expeditious read- ing of the proofs when received (we mail all proofs by air mail) and their prompt return by air mail also. Since we conscientiously reply to all letters we actually receive, and since we experience a constant loss in insured THE VELIGER Page 305 and registered mail pieces, we have come to the conclusion that if a correspondent does not receive an answer from us, this is due to the loss of either the inquiry or the reply. 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Such changes may be apparently very simple, yet may require extensive resetting of many lines or even entire paragraphs. Also we wish to stress that the require- ment that all matter be double spaced, in easily legible form (not using exhausted typewriter ribbons!) applies to all portions of the manuscript — including figure explana- tions and the “Literature Cited” section. It may seem inappropriate to mention here, but again recent experience indicates the advisability of doing so: when writing to us, make absolutely certain that the cor- rect amount of postage is affixed and that a correct return address is given. The postal service will not forward mail pieces with insufficient postage and, if no return address is given, the piece will go to the “dead letter” office, in other words, it is destroyed. Page 306 BOOKS, PERIODICALS, PAMPHLETS Shells of New Zealand by A. W. B. Powe t. Fifth revised edition. Whitcoulls Publisher, Christchurch, New Zealand. 154 pp., 45 plts. (2 in color) $12.00. April 1977 This latest edition of a classic work lists a total of 2256 molluscan species, both marine and non-marine, that have been reported in New Zealand, and it illustrates 555 of them. Except for updating, the general format of the work is unchanged; papers cited in the bibliography again are cross-referenced into the body of the list, and this has been completed up through 1974. Some new illustrations are added, as well as new paragraphs in the introductory text, which includes useful notes on collecting, on classification, and on the significance of molluscan studies in under- standing both the geologic past and present-day distribu- tion patterns of marine life. The bibliography runs to 14 pages; the check list, with its mine of references, to 56. The book is hardbound, and the dust-jacket carries a small portrait of Dr. Powell, who has been on the staff of the Auckland Museum for more than 56 years. Al- though he formally retired in 1968 as Director, he con- tinues work as Honorary Research Associate. This book is yet another in a long list of significant contributions to malacology that have come from his pen. A. Myra Keen Sea Shells from Cape Verde Islands by Luis P Burnay & ANTONIO A. Montero. Lisbon, 88 pp., 2 text figures, 2 maps, 54 photographs on coated paper. January 1977 Available from Seashell Treasures, P O. Box 730, Oak- hurst, CA 93644 at US$5.00 This paperback book, rather charmingly written though not without an occasional grammatical or spelling error THE VELIGER Vol. 20; No. 3 (the authors deserve commendation for their erudite use of the English language which is far better than that of some American writers!) is organized into two parts, the first of which contains chapters on collecting and preserv- ing techniques; but of special interest are the introductory remarks on the collecting area. The second part contains a systematically arranged account of the species found by the authors during 2 successive collecting trips in 1975 and 1976. A bibliography of 122 titles and a 2-page index complete the volume. As can be expected when collecting is extended to an area that has not been readily accessible in the past, specimens are obtained that do not completely agree in every detail with the otherwise well-known representa- tives of a particular taxon. Such differing forms are too frequently described as new species with little supporting or comparative material being adduced. It is therefore extremely refreshing to find that these authors, while acknowledging having found such different forms, have specifically refrained from describing and naming them and thus they have avoided joining the ranks of the ex- treme splitters. We congratulate them on this very wise choice. R. Stohler Wonders of Starfish by Morris K. JacoBson & WiLL1AM K. Emerson. Dodd, Mead & Company. New York, N. Y. 10016. 80 pp., nu- merous, though unnumbered, photographs and drawings. $4.95 5 July 1977 Although the hardbound booklet is intended for the juve- nile reader (from 10 years up), it is nevertheless scientifi- cally accurate. It is a reliable introduction to a very in- teresting and highly specialized group of marine animals that even the most unobservant casual visitor to the sea shore cannot fail to notice. And young children with their unsatiable curiosity will find almost all the answers in this book. R. Stohler THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributional, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS"; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES” will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 812” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 51/2” by 81/2”), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr. Donatp P. Azssort, Professor of Biology Hopkins Marine Station of Stanford University Dr. Warren O. Anpicott, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Associate Professor of Paleontology, Stan- ford University Dr. Jerry DononveE, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatr Duruam, Professor of Paleontology University of California, Berkeley, California Dr. Cavet Hann, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. Joe, W. HepcpetH, Adjunct Professor Pacific Marine Station, University of the Pacific Dillon Beach, Marin County, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Victor Loosanorr, Professor of Marine Biology Pacific Marine Station of the University of the Pacific EDITOR-IN-CHIEF Dr. RupotF STOHLER, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Associate Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. Frank A. Prrexa, Professor of Zoology University of California, Berkeley, California Dr. Rosert Rosertson, Pilsbry Chair of Malacology Department of Malacology Academy of Natural Sciences of Philadelphia Dr. Peter U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. JupirH Terry Smiru, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Ratpy I. Situ, Professor of Zoology University of California, Berkeley, California Dr. Cuarces R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. THompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. Jean M. Cate Rancho Santa Fe, California A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California VOLUME 20 . APRIL 1, 1978 NUMBER 4 CoNnTENTS The Chromodoridinae Nudibranchs from the Pacific Coast of America. — Part II. The Genus Chromodoris. (3 Plates; Text figures 4 - 15) EVAN SHDERTSC Hise Sir MBE Ieuan Fev ON Toh NAN reh tale Ve Neder Ve) var ncGOY The Family Columbellidae in the Western Atlantic. Part IIb. — The Pyreninae (Continued). (4 Plates: 1 Text figure) GrORGEB Es RAD YIN ae umirun ley miet eerie. Weare eC gelesen as BB Two New Giant Epitoniids (Mollusca : Gastropoda) from West Africa. (1 Plate; 1 Text figure) PHILIPPE, BOUCHE, & SIMON MDILEIER) «al 6) s/s) sf ee e's ew ss 345 Clonal Variation in the Parthenogenetic Snail Campeloma decisa (Viviparidae). Ropert K. SELANDER, E. Davis ParKEr, Jr.,& RoBERT A. BROWNE .. . . « 349 Effects of Cornstarch and Dextrose on Oysters. KENNETH W. TurGEoN & Dexter S. HAVEN . . . . . 1. 1 ss ee es 352 Further Field Notes on the Behavior of Aplysia dactylomela. ERWOBACH Mis mae mice Lt, el hier Rate ay ae ONS ee Ste al a 2: we BRO Flight Responses of Two Species of Intertidal Gastropods (Prosobranchia : Troch- idae) to Sympatric Predatory Gastropods from Barbados. DANIEL HOFEMAN) & (PAUL) J, WELDON «1-05 <1. - 6) 5 se ye eo GOT Notes on the Spawning and Egg Capsules of Two Prosobranch Gastropods: Nassari- us tiarula (Kiener, 1841) and Solenosteira macrospira (Berry, 1957). (2 Text figures) ROVE HOUSTON IC Mn re ee ee eis se lee ter eels cee «G07 Additional Molluscan Records from Bahia de los Angeles, Baja California Norte. Forrest L. PoorMAN & Leroy H. PoorMAN . . . «©.» + + @ @ = + + 369 [Continued on Inside Front Cover] aaa SSS SSSI Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SUPERFAMILY, FAmiLy, Subfamily, Genus, (Subgenus) New Taxa Second Class Postage Paid at Berkeley, California ee ee ee ee ee ConTENTS — Continued Tambja and Roboastra (Mollusca : Opisthobranchia) from the Gulf of California and the Galapagos Islands. (20 Text figures) Westry M. Parmer) 6020 514062) eis sg se te etek ee aE res METHODS & TECHNIQUES ... . 2) ANOeS Heloise ene Air Drying Giemsa Technique for Chien Chee (1 Plate) R. Prasap « C. C. Das INFORMATION DESK. ... . rh ipkies A Ry RRM teat te oem 2 er Re aren ra The Role of the Editorial Rete A. Myra KEEN NOTES & NEWS .. . . er asters: Woteene nce 6! GYO\O) Soviet Contributions to Matacoleayt in ee Morris K. JACOBSON & KENNETH J. Boss A Note on Changes in Marine Intertidal Fungus Taxonomy. Davin R. LINDBERG A Rectification of a Statement Regarding the Lamarckian Collection in the Book “Murex Shells of the World” by George E. Radwin and Anthony D Attilio. AnTHoNy D’ATTILIO Deshayes Types in the National Museum, Paris. Twila BRATCHER Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 21: $30.- plus postage ($1.- in U.S. A.; $2.50 to P.U.A.S.; $3.50 to all other foreign Countries, including Canada). Single copies this issue $16.-. Postage extra. Send subscription orders to California Malacozoological Society, Inc. 1584 Milvia Street, Berkeley, CA 94709, U.S.A. Address all other correspondence to Dr. R. Stou.er, Editor, Department of Zoology University of California, Berkeley, GA 94720 Vol. 20; No. 4 THE VELIGER Page 307 The Chromodoridinae Nudibranchs from the Pacific Coast of America. - Part II. The Genus Chromodoris HANS BERTSCH '! Donner Laboratory and Department of Zoology, University of California, Berkeley, California 94720 (3 Plates; Text figures 4 - 15) THIs IS THE SECOND article in a 4-part series monograph- ing the Chromodoridinae from the Eastern Pacific. Fig- ures and tables are numbered consecutively through all 4 articles. Part I (BertscH, 1977) consisted of Materials and Methods and supra-specific chromodorid taxonomy. In Materials and Methods, I examined the use of the radula in opisthobranch taxonomy by: 1) analyzing both the contribution of scanning electron microscopy (SEM) to radular studies (cf. also a recent paper by HICKMAN, 1977, which appeared too late for inclusion in Part I) and methods of specimen preparation for successive viewing by SEM and light microscopy; 2) discussing the numer- ical analysis of radular variation (see also BEertscu, 1976a);and, 3) defining the structural terms applicable to radular teeth. The taxonomic coverage of Part I was at the levels of the family, subfamily and genus. Chromodoris Alder & Hancock, 1855 Chromodoris baumanni Bertsch, 1970 (Figures 3-A, 4, and 10-12) References and Synonymy: Chromodoris norrisi (not Farmer, 1963). Marcus & Marcus, 1967: 170-173; fig. 24; Material 3 Chromodoris sp. BERTSCH, 1971: 16 Chromodoris baumanni Bertsch, 1970: 8 - 12; figs. 3- 13 SPHON & MULLINER, 1972: 150-151. SPHON, 1972b: 59. BertscH, FERREIRA, FARMER & Hayes, 1973: 289, 292. BERTSCH, 1973: 108, 110. KEEN & COAN, 1975: 43. BERTSCH, 1976b: 157 ' Present address: Biological Sciences, Chaminade University of Honolulu, Honolulu, Hawaii 96816 Editor’s Note: For Figures 3-A, 3-B, etc. see The Veliger 90 (2): 115 Material Examined and Distribution: Baja California, Mexico: 1) 1 specimen, Isla Carmen; leg. C. Gage, April 1974 (i- dentified from a color transparency) 2) 3 specimens, 1km north of Isla Monserrate; leg. A. J. Ferreira, 16 June 1974 (HB 377 A- C) 3) 1 specimen, La Paz area; leg. E. Janss, Jr., April 1972 (HB 398; LACM A-9555) 4) 1 specimen, 2 - 3m subtidal, Bahia Carisalito, 4km N of Bahia Las Cruces; leg. H. Bertsch, 23 July 1972 (HB 5) 5) 1 specimen. Isla Espiritu Santo; leg. W. M. Farmer, 29 June 1964 (identified from a color transparency) 6) 1 specimen, Isla Espiritu Santo; leg. E. Janss, Jr., April 1974 (HB 401; LACM A-9555) 7) 1 specimen, Isla Cerralvo; leg. W. M. Farmer, May 1962 (identified from a color transparency) 8) 2 specimens, intertidal to 6m subtidal, Cabo Pulmo; leg. C. Gage et al., 25-26 May 1971 (HB 386 A-B; LA CM) Mainland Mexico: 9) 1 specimen, 20m subtidal Guaymas, Sonora; leg. A. Kerstitch, 12 March 1966 (HB 321; USNM 753559) 10) 2 specimens, intertidal, Sayulita, Nayarit; leg. G. G. Sphon, January 1970 (HB 397 A-B; LACM 70-4) II) 3 specimens, intertidal, Punta Mita, Nayarit; leg. F a R.Poorman, 2 January 1976 (HB 416 A-C; LACM A- 8477) South America: 12) 1 specimen, Academy Bay, Santa Cruz Island, Galapagos Islands; leg. A. G. Smith, February 1964 (identified from a color transparency) 13) 1 specimen, intertidal, Academy Bay, Santa Cruz Is- land, Galapagos Islands, leg. Ameripagos Expedition, March 1971 (HB 400; LACM 71-44) Type Locality: The type locality of Chromodoris baumanani is Isla San Francisco, Baja California; previous collecting records of this species include the Gulf of California, southern Mexi- Page 308 co, Central America, and the Galapagos Islands. Its north- ernmost reported occurrence is Bahia San Carlos, Sonora, Mexico (BerTscu et al., 1973). External Morphology and Coloration: Chromodoris baumanni reaches at least 62mm in total length. Notal and foot background color is white; in the notal dorso-median region this background color is light yellow. Numerous small red-purple dots cover the notum and the posterior and lateral surfaces of the foot. An inter- rupted band of orange (or light red) dots and streaks surrounds the lateral edges of the notum and foot. The rhinophores are white, with a red-purple coloration dis- tally; the extreme tip is white. The gills are white, with a purplish hue on the distal portion (cf. the color illustra- tions in BerTScH, 1970: figs. 3 - 6). Radula: Sizes of each radula examined, the respective width: length ratio, number of tooth rows and maximum number of teeth per halfrow are given in Table 1. The combined radular formula (including literature references) is 47 - 84 rows with 2g - 68 teeth per half-row; rachidian tooth ab- sent. Least squares regression line analysis shows that the number of rows of teeth and the maximum number of THE VELIGER Vol. 20; No. 4 teeth per half-row are positively correlated (Figure 10-A). The coefficient of correlation (r) is 0.949 (n= 16, P < 0.001). The regression line is described by the equa- tion Y =-11.09+0.922 X. The number of rows is positively correlated with the length of the radula (Figure 11-A). The regression line formula is ¥Y = 37.96+ 11.228 X; r = 0.922, n = 14, and P < 0.001. The maximum number of teeth per half-row is depend- ent upon the width of the radula (Figure 12-A). The equation, Y = 24.52+19.674 X, describes the regression line and the coefficient of correlation equals 0.8808, n = 14, P < 0.001. Figure 4-A presents an outline sketch of a flat-mounted radula. The innermost lateral tooth (Figure 4-B) has 3 - 4 denticles on the inner face, and 2 - 6 (usually 3-5) den- ticles on the outer face. The inner lateral teeth (approxi- mately the first 5 - 10) are short, with a strongly recurved shaft and a proportionately long base; the cusp is much longer than the succeeding denticles (Figures 4-C - 4-G). Throughout the middle of the half-row, the teeth (Figures 4-H, 4-1) have a longer, straighter shaft, with 6 - 9 den- ticles on the posterior surface. The denticles are larger in size relative to the cusp. The length of the cusps (measured on a straight line, from the notch joining the first denticle with the cusp, Table 1 Radular variation in Chromodoris baumannt Width: Number Maximum number Specimen Length Width length of tooth of teeth per (11B numbers) (in mm) (in mm) ratio rows half-row i = = = 82 65 3 i Ee uk 84 64 5 1.99 3} ena 7 53 4) 321 3.33 1.82 iLsil fot 85 68 377 A 2.99 1.66 1:1.8 74 55 377 B 2.8] 1.58 1:1.78 70 55 SMC. 3.29 1.86 Nev7/ 67 54 386 A 3.76 1.58 1:2.38 80 7 386 B 3.04 1.92 1:1.89 78 55 397 A 1.27 ().57 12193 59 34 397 B 0.85 0.46 1:1.85 47 31 398 2 26 Hib} 1:2 64 55 401 1.52 0.75 1:2.03 49 37 4I6A 1.18 0.53 I e@ 28} 55 42 416 B ().92 0.36 1:2.56 48 29 416C 1.58 0.85 1:1.86 55 39 x 2.94 1.16 1:1.99 65.625 49.438 s 1.045 0.572 0.254 13.657 13.261 "Marcus & Marcus, 1967; 7Bertscn, 1970) Vol. 20; No. 4 to the tip of the cusp) averaged 0.oo86mm (range, 0.005 - 0.013mm, n= 26) for the inner 15 lateral teeth and random teeth from the middle of the half-row. The mean ratio of the length of the first denticle to the length of the cusp is 1:1.79 (range, 1.2 - 3.25; S=0.457; n= 27). The outermost lateral teeth (Figures 4-J, 4-K) be- 4-A 4-B 4-C 4-D ~) 4-E 4-F 4-G ~ 4-H ail 4-J THE VELIGER Page 309 come smaller, with cusp and denticles greatly reduced in size. Developing teeth (Figures 4-L-4-O) show earliest development of the base and shaft, with minute denticles. Denticles begin growth as small points on the future pos- terior surface of the tooth. After initial formation, growth 2 4-M Figure 4 Radular teeth of Chromodoris baumanni A — HB 398; outline sketch of entire radula B - HB 5; innermost lateral tooth, approximately row 17, right side of radula (RSR) ; length between marks (LBM) 0.059mm C — HB 5; postero-lateral view of 2™4 lateral tooth, row 40, RSR; LBM, 0.053 mm D - HB 398; distal portion of shaft, 24 lateral tooth, row 55, left side of radula (LSR) E — HB 368; distal portion of shaft, 3™ lateral tooth, row 55, LSR F — HB 398; distal portion of shaft, 7'* lateral tooth, row 55, LSR G — HB 386 A; distal portion of shaft, 11‘ lateral tooth, row 37, RSR H —- HB 398; distal portion of shaft, 26 lateral tooth, row 21, RSR I — HB 5; tooth from middle of half-row, approximately row 42, LSR; LBM, 0.081 mm J - HB 5; second outermost tooth, approximately row 42, LSR; LBM, 0.079mm K — HB 5; outermost tooth, approximately row 40, LSR; LBM 0.055 mm L — HB 308; developing 1*t lateral tooth, row 64, RSR; LBM 0.03 mm M - HB 3098; developing 5" lateral tooth, row 64, RSR; LBM, 0.039 mm N -— HB 398; developing 10" lateral tooth, row 64, RSR; LBM, 0.071 mm O - HB 398; developing 36% lateral tooth, row 64, RSR; LBM, approximately 0.12mm P — Monaxon sponge spicule (with recurved ends) found under the shafts of lateral teeth in Chromodoris baumanni (HB 377-C) radula; LBM, approximately 0.24mm Page 310 proceeds by thickening and enlarging of the various tooth parts. In the original description of this species, BERTSCH (1970: figs. 7-10) presented scanning electron micro- graphs of the radular teeth. Discussion: Chromodoris baumanni is closely related to C. norrist. Differences between the 2 species will be analyzed under C. norrist. Chromodoris baumanni also shows similarities with the Hawaiian C. lilacina (Gould, 1852), but the colorations and radular morphologies of these species fall outside each other’s range of variation. The rhinophores and gills of C. lilacina are orange-yellow or straw-colored (Kay & Younc, 1969: 202), distinct from the reddish- purple hues of C. baumanni;C.lilacina has proportionate- ly fewer teeth per half-row (relative to the number of tooth rows) than does C. baumanni. Feeding habits of Chromodoris baumanni are not known; a sponge spicule (Figure 4-P) was lodged beneath the tooth shafts of one radula. The cusp and denticle morphology of the radular teeth are primarily adapted for scraping across sponge tissue, but the teeth can also hook the curved ends of such C-shaped spicules and extract more tissue adjacent to the spicule. Chromodoris galexorum Bertsch, spec. nov. (Figures 3-B, 5, 10-12, 33 - 36) Material Examined and Distribution: Gulf of California, Mexico: 1) Holotype. 17m subtidal, under a ledge, Isla San Pedro Martir, Sonora, Mexico (approximately 28°22’N; 112° 20’W); leg. A. Kerstitch, 16 June 1976 (HB 467 B). This dissected specimen and its mounted radula have been deposited in the collections of the Los Angeles County Museum of Natural History, LACM Type Series, No. 1848. 2) Paratypes. 2 specimens, 17m subtidal, under a ledge, Isla San Pedro Martir; leg. A. Kerstitch, 16 June 1976 (HB 467 A, D). Water temperature at depth collected, 22.3°C I specimen, 12m subtidal, in a dark cave, Isla San Pedro Nolasco, Sonora, Mexico (approximately 27°58’N; 111° 22’W) ; leg. A. Kerstitch, 22 June 1976 (HB 467C) 2 specimens, subtidal, Guaymas, Sonora, Mexico; seen by A. Kerstitch, April 1972 (identified from a color trans- parency ) 5) 1 specimen, La Paz area, Baja California; leg. E. Janss, Jr., April 1974 (HB 256) &> a The known occurrence of this new species is from sub- tidal localities in the central and southern Gulf of Cali- fornia. Isla San Pedro Martir is the type locality. Mr. A. THE VELIGER Vol. 20; No. 4 Kerstitch (personal communication) states it is fairly common subtidally around Guaymas in the springtime. He collected an additional specimen of Chromodoris galexorum at Isla San Pedro Nolasco, in 15m of water, on 16 March 1977. External Morphology and Coloration: The lengths of 5 preserved specimens were 15, 16, 17, 22, and 24mm. The number of gills (usually more numer- ous in larger specimens} were 7, 10, and 16 in 3 specimens; rhinophore lamellae varied from 18 - 24. The body background color is white. Scarlet spots oc- cur on the dorsum, many of them being immediately sur- rounded by a chrome yellow ring. At times the yellow coloration is present also as small splotches within the scarlet. The scarlet spots are largest down the midline of the dorsum. One specimen also had a transverse series of 3 larger spots (with overlapping edges) halfway between the rhinophores and the gills. The notal border is rimmed dorsally with a solid chrome yellow band. The lateral and posterior surfaces of the foot are white, with small scarlet dots scattered throughout; the sides of the foot can have 4-5 loose rows of these scarlet maculations. Except for yellow rings surrounding 2 or 3 scarlet spots on the midline of the postero-dorsal foot surface, there is no yellow color- ation on the foot. The rhinophores and gills are scarlet, darker distally. Whitish glands occur on the underside of the notal over- hang. There are 6 - 7 per side of the body, and each con- sists of an ovalish structure that has 4-5 finger-like ex- tensions protruding towards the animal’s body. Radula: Meristic data of 5 radulae are found in Table 2. The range of variation of the radular formula is 56-59 (47- 57°1°47-57). The number of teeth per halfrow is only slightly less than the number of rows. There were too few specimens available to perform regression analyses on radular characteristics; the points of the radular para- meters are graphed in Figures 10 - 12. The radular teeth have the typical unicuspid Chromo- doris shape: inner laterals have shorter shafts with den- ticles on the outer face (Figure 34), followed by longer shafted teeth in the middle of the half-row that show denticulation on the posterior surface (Figures 35, 36). A triangular rachidian tooth is present (Figures 5-A, 33), with an elevated central cusp. The innermost lateral tooth (Figures 5-A, 33) has 3-4 denticles on the inner face, 5-6 on the outer face. Denticles increase in number on each tooth towards the middle of the halfrow, where there are usually 15 - 18 denticles on the posterior surface. Scanning electron micrographs cannot be used to count Vol. 20; No. 4 THE VELIGER Page 311 Table 2 Radular variation in Chromodoris galexorum Width: Number Maximum number Specimen Length Width length of tooth of teeth per (HB numbers) (in mm) (in mm) ratio rows half-row 256 — LEST — -- 53 467 A 3.2] 1.64 1:1.96 57 52 467 B 2.87 1.56 1:1.84 56 47 467 C 3.11 1.92 1:1.62 59 49 467 D 3.88 2.24 ieila33 59 57 X 3.27 , as 1:1.79 57.75 51.6 s ().433 0.34 0.146 1.5 3.847 i, = Figure 5 Radular teeth of Chromodoris galexorum Bertsch, spec. nov. A — HB 256; rachidian tooth and 1* lateral tooth drawn in relative position to each other, row 11, LSR; LBM, 0.071 mm;; rachid- ian 0.028mm long B — HB 256; lateral tooth 52, row 11, LSR; LBM, 0.071 mm denticles routinely, since overlapping teeth in the field of view obscure parts of teeth behind them. Denticular counts must be verified with the light microscope. Lengths of tooth cusps ranged from 0.012 - 0.024mm, with a mean of 0.019mm (n= 21). The cusp of each tooth averages 5.05 times as long as the first denticle (n = 21; range is 3 - 7; s== 1.073). The outermost 4-5 teeth approach the first denticle length : cusp length ratio of 1:1; the cusp, denticles, and shaft length become markedly shorter (Figure 5-B). It is of special note (with reference to the later dis- cussion of Chromodoris sphoni) that in the last 8 radular rows of specimen HB 467 D, the inner 3 - 6 lateral teeth have a slight denticulation or ridging of 1 - 5 denticles on the inner face of the distal shaft area. These are in addi- tion to the normal positioning of denticles on the outer face of the tooth shaft. Discussion: Chromodoris galexorum needs to be compared with Panamic chromodorids that exhibit a white notal back- ground. It is immediately distinguishable from those spe- cies with dominant blue coloration. Chromodoris bau- manni does not have a solid yellow band bordering the notum edge, but it has a broken orange or red band; moreover, it does not have bright yellow enclosing its red- violet spots. The spots of C. baumanni are also smaller and more regularly circular than the big, irregular red blot- ches of C. galexorum.Chromodoris marislae has yellow- orange circlets (some with open centers, others solid), with white surrounding the larger markings. Chromodoris norrisi has evenly circular markings of red and yellow, with a broken orange band around the notum. Chromo- laichma dalli has black dots on its notum, and Chromo- laichma sedna has a completely white notum with solid yellow and red bands around the periphery of the mantle. Two Australian species appear similar to Chromodoris galexorum, but can be readily separated on the basis of coloration and radular morphology. The ranges of varia- tion of the 3 species do not overlap. Chromodoris daphne (Angas, 1864) lacks yellow borders to the scarlet macula- tions, and has both yellow and red color bands surround- ing the notum; it has proportionately shorter teeth cusps Page 312 THE VELIGER Vol. 20; No. 4 and fewer denticles on the shafts of the teeth than C. galexorum (the teeth of C. daphne are illustrated in THOMPSON, 1972: figs. 3i, 3j, 3k). Chromodoris splendi- da (Angas, 1864) also lacks yellow rimming the scarlet maculations; in large specimens, the scarlet spots are highly irregular blotches, often fusing into a thick reticu- lating pattern. The teeth of C. splendida have fewer den- ticles (cf. THompson, op. cit.: plt. 2d, figs. 2k - 20) than C. galexorum; the outer teeth, not reduced in size, have thin erect shafts devoid of denticles (THompson, op. cit.: pit. 2 c). Thompson also presents color drawings of living C. daphne and C. splendida. Etymology: The specific name galexorum is chosen as an acronym of Gale and Alex, to honor Mr. Gale Sphon (Los Angeles County Museum of Natural History) and Mr. Alex Ker- stitch (Tucson, Arizona), who provided me with speci- mens of this new species. Chromodoris marislae Bertsch, in BERTSCH, FERREIRA, FARMER & HAYES, 1973 (Figures 3-C, 6, 13 - 15) References: Bertscu, Ferreira, FARMER & Hayes, 1973: 289 - 292; figs. 1-11. BERTSCH & FERREIRA, 1974: 344. KEEN & Coan, 1975: 45 Material Examined and Distribution: Baja California, Mexico: I) 1 specimen, Guaymas, Sonora; leg. A. Kerstitch, April 1972 (identified from a color transparency) 2) 1 specimen, Los Islotes; leg. A. J. Ferreira, September 1971 (HB 370) ; 3) 5 specimens, La Paz area; leg. E. Janss, Jr., April 1972 (HB 34 A-E) The known distribution of Chromodoris marislae had been limited previously to localities along the Baja Cali- fornia Gulf coast, between Isla Santa Catalina (type lo- cality) and La Paz. The Guaymas record is the first report from the coast of mainland Mexico, and constitutes a northward range extension of over 230km. The specimens from La Paz are part of Lot 4 of the original description. External Morphology and Coloration: Living Chromodoris marislae reach 80mm in total length (BerTscH et al., 1973). Body color is an ofEwhite, with 2 or 3 irregular rows of orange spots encircling the periphery of the notum; centrad to these rows is a roughly circular arrangement of larger orange ringlets, which are often surrounded or marked centrally with a pure white coloration. The bases of the rhinophores have almost a translucent quality, and the distal end is light brown; there is a prominent median white longitudinal septum on both its anterior and posterior faces. The gills are also light brown, with pure white on the center of each branching of the gills (color photographs of C. marislae are in BERTSCH et al., op. cit.: figs. 1, 2). Radula: The radular formula varies from 59-82 (53-70'1'53-70). Meristic data for 8 radulae are presented in Table 3. The number of rows and the maximum number of teeth per half-row are positively correlated (Figure 13) ; Y = 20.56+0.6039 X; r—=o0.9005, n= 8, P< 0.01. Table 3 Radular variation in Chromodoris marislae Specimen Length Width (HB numbers) (in mm) (in mm) 34A 5.07 2.99 34B 4.14 2.26 34C 3.9 2.3 34D 3.66 2.02 34 E 4.71 2.16 370 4.59 2.3 Holotype’ _ - Paratype 2° - - x 4.35 2.34 s 0.535 0.3366 SBERTSCH e/ al, 1973 Width: Number Maximum number length of tooth of teeth per ratio rows half-row 1:1.69 76 70 1:1.83 60 57 NENEZ/ 58 54 1:1.81 60 59 1:2.18 82 68 1:1.99 77 67 _ 59 59 — 62 53 1:1.87 66.75 60.875 0.188 9.8 6.578 Vol. 20; No. 4 A positive correlation exists between the length of the radula and the number of rows (Figure 14). The regres- sion line formula is Y = -6.27+17.28 X, r= 0.8691 (10501 0.1). The parameters are simply plotted in Figure 15. The rachidian tooth (Figure 6-A) has a long base, and a bifurcated, erect cusp. There are 3 - 4 denticles on both the inner and outer faces of the innermost lateral tooth (Figure 6-B). The denticles increase in number from the center of the radula to the middle of each row, and then decrease towards the outermost teeth. For example, row 41 (left side of radula, HB 34 B) has 3 denticles on the outer face of each of the first 13 lateral teeth, 4-5 den- ticles on the posterior surfaces of the next 10 teeth, 5 - 7 on each of the succeeding 20, and 5 - 3 denticles on each of the outermost 14 teeth. The innermost laterals are not smooth (contrary to the statement in BerTscH et al., 1973: 290), but are denticled on the outer face. Figure 8 of Bertsch et al. is a view of the inner face of an inner- most lateral, and denticles are not visible because of the non-transparency of teeth in scanning electron micro- graphs. The teeth of Chromodoris marislae are large, with solid cusps and fairly large denticles. The cusp lengths of the inner 36 teeth of row 20 (HB 34 B) varied from 0.02 -0.032mm (X==0.027mm). The reduced size of the outermost lateral teeth is accompanied by reduced cusp and denticle size. The outer 10 teeth had cusp lengths of 0.02 - 0.o1mm (X =0.016mm). In this same row, the cusps averaged 2.26 times longer than the den- ticles (range, 1.25 - 3.2, S== 0.449, n= 46). Quite a few of the lateral teeth show a unique specific denticulation pattern (Figures 6-C to 6-H). The first 2 denticles below the cusp are united from a common base, giving the appearance of bifurcating. Although this is not universal, it appeared often enough on each of the radulae examined to be considered diagnostic. This con- dition was illustrated by Bertscu et al. (1973: figs. 5 A, B, D, E) but they did not comment upon its occurrence. Scanning electron micrographs of Chromodoris maris- lae radular teeth are in Bertscu et al., 1973. Discussion: The current records of Chromodoris marislae are all from subtidal locations; increased SCUBA research in the Gulf of California should yield more specimens. Chromodoris mcfarlandi Cockerell, 1g01 (Figures 3-D, 7, 13 - 15, 37 - 40) References and Synonymy: Chromodoris mcfarlandi CocKERELL, 1901: 79 - 80. COCKER- ELL, 1902: 20- 21. BerTscH, 1976b: 157 - 158 Page 314 Glossodoris macfarlandi (Cockerell). O’DoNoGHUE, 1926: 212. O’DonocHUE, 1927: 8g - go, 116; plt. 2, figs. 33 - 37. Pruvot-For, 1951a: 120. Pruvot-Fo1, 1951b: 152. Lance, 1961: 66. PAINE, 1963: 4, 7, 8. FARMER « Co1- LIER, 1963: 62. STEINBERG, 1963:69. MAcFarLanp, 1966: 153-157; plt. 22, figs. 1-5; plt. 34, figs. 1-11. SpHoN & Lance, 1968: 79. RickeTTS & CaLvin, 1968: 119, 514. McBeru, 1970 (not seen; fide BLoom, 1976). ABBOTT, 1974: 354, fig. 4239. BLoom, 1976: 292 - 294 Chromodoris macfarlandi Cockerell. CockERELL & ELIOT, 1905: 36. MacFaranpb, 1906: 129. CocKERELL, 1908: 106. JOHNSON & SNOOK, 1927: 494; plt. 11, fig. 2. Marcus & Marcus, 1967: 178. Ro ier « Lone, 1969: 425,424. LANcE, 1969: 37. ROLLER, 1970a: 371. Rot- LER, 1970b: 482. Lonc, 1970: 19. McBeEtTH, 19714: 28. KEEN, 1971: 822; SCHMEKEL, 1972: 194. SPHON, 1972: 59. Berrscu et. al., 1973: 287. SMITH & Cari- TON, 1975: 528, 538; plt. 121, fig. 6. KEEN « Coan, 1975: 43 (# 2330) The correct date of authorship, as O’ DONOGHUE (1927: 89) showed, is 1901. The description published 28 Novem- ber 1901 has priority over Cockerell’s intended new spe- cies description published June 1902. The spelling of the specific name requires comment. Cockerell’s original spelling was a misspelling of Mac- Farland’s name. The species name occurs once in his 1901 paper and once in his 1902 article, both times as Chromodoris mcfarlandi. In acknowledging the patro- nym, CocKERELL (1902: 21) wrote, “Named after Prof. FM. McFarland [sic] of Stanford University, who has done some excellent work on the nudibranchs of Pacific Grove, California.”’ Cockerell was clearly in error on the proper spelling of Dr. MacFarland’s name. The rules of zoological nomenclature provide that the original spelling of a species name is to be retained as the correct spelling unless it contravenes certain manda- tory provisions or there is clear evidence in the original publication that an inadvertent error (lapsus calami, copy- ist’s or printer’s error) has occurred (Mayr, 1969: 312 to 313; 355 to 356). The evidence is to the contrary. Both of Cockerell’s papers misspelled MacFarland (including the patronymic designation), and there are no multiple spellings in the original 1901 paper (nor in the 1902 paper). Repetition of the error in 2 different papers sub- mitted to 2 separate journals, makes it highly unlikely that a slip of the pen or a printer’s error occurred. There is no clear evidence of an inadvertent error (in the sense of the Code), and no contravention of mandatory pro- visions. Therefore, the spelling of C. macfarlandi is an unjustified emendation and a junior objective synonym of the original C. mcfarlandi. THE VELIGER Vol. 20; No. 4 Material Examined and Distribution: California and Offshore Islands: 1) 1 specimen, subtidal, off Monterey breakwater; leg. R. Ames, January 1963 (HB 458; CAS) 2) 2specimens, Monterey Bay; leg. F M. MacFarland, De- cember 1908-January 1909 (HB 453 A-B; CAS) 3) 1 specimen, 22m _ subtidal, off Del Monte, Monterey Bay; leg. S. S. Berry (no date) (CAS) 4) 2 specimens, Pacific Grove; leg. F M. MacFarland, June 1908 (HB 454 A-B; CAS) 5) 1 specimen, Point Pinos, Monterey; Jeg. R. Page, 12 July 1941 (HB 455; CAS) 6) 1 specimen, 24m subtidal, north end San Jose Creek Beach (36°32’N; 121°56’W); leg. J. McLean, 9 July 1971 (HB 393; LACM 60-24) 7) 1 specimen, intertidal, White’s Point, Palos Verdes Pen- insula (33°43’N; 118°18’W); leg. G. G. Sphon, 9 De- cember 1969 (HB 396; LACM 69-37) 8) 2 specimens, Newport Bay; leg. G. E. MacGinitie, June 1948 (HB 457 A-B; CAS). These are the specimens drawn on plt. 22, figs. 1-5, of MacFarLanp, 1966 9) 2 specimens, Corona del Mar; leg. G. E. MacGinitie, 25 June 1948 (HB 456; CAS) 10) 1 specimen, La Jolla; leg. T.D. A. Cockerell, 1902 (HB 452; CAS) 11) 1 specimen, Isthmus, Santa Catalina Island; leg. A. J. Ferreira, 9 July 1975 (HB 361) 12) 1 specimen, intertidal, Catalina Harbor, Santa Catalina Island; leg. G. G. Sphon, 7 March 1970 (HB 392; LACM 0-8 13) : ae 23 - 29m subtidal, upper reef of Farnsworth Bank, Santa Catalina Island (33°21’N; 118°31’ W) ; leg. C. Turmer, 1-2 June 1970 (HB 394; LACM 70-74) 14) 1 specimen, 27 - 34m subtidal, northwest of Pyramid Head, San Clemente Island; leg. C. Swift, 1 July 1971 (HB 395; LACM A 9325) Mexico: 15) 3 specimens, 21 - 24m subtidal, Isla Coronado (31°48" N; 116°48’W); leg. A. J. Ferreira, 28 September 1973 (HB 253) 16) 1 specimen, intertidal to 11m_ subtidal, Man-of-War Cove, Bahia Magdalena (24°37.5’N; 112°7.5’ W) ; leg. J. McLean and P LaFollette, 31 October 1971 The original material used by Cockerell was collected from La Jolla and San Pedro. The reported range of Chromodoris mcfarlandi has been from Monterey to the Isla Cedros area, Baja Cali- fornia (KEEN, 1971: 822). The specimen collected at Bahia Magdalena represents a southern range extension of over 480km. Despite this new record at the northern boundary of the Panamic faunal province (tropical West America), KEEN & Coan (1975: 43) were correct in con- Vol. 20; No. 4 THE VELIGER Page 315 sidering C. mcfarlandi a member of the temperate and cooler water Californian and southern Oregonian marine provinces (sensu VALENTINE, 1966, and 1973: 351 - 356). This new record of C. mcfarlandi, while representing a true “range extension,” is not indicative of the normal occurrence of this species. It is a thermally anomalous record (ZINSMEISTER, 1974) on the extreme periphery of the species’ range. Chromodoris mcfarlandi has also been reported from San Luis Obispo County, Santa Barbara County, Laguna Beach, Santa Catalina Island (California), and Isla Co- ronado (Mexico). ; Lot 14 represents a new subtidal bathymetric range. External Morphology and Coloration: Living Chromodoris mcfarlandi are reported to reach 50 - 60mm in length (JoHNSON & SNOOK, 1927: 494), but more commonly will vary in length up to 35mm (Mac- FaRLAND, 1966: 156). Overall body color a brilliant reddish-violet. A yellow line runs down the center of the notum, from just anteri- or to the rhinophores to the forward edge of the gill pocket; an additional yellow line begins postero-laterally to each rhinophore, runs lengthwise along the animal’s body, and joins behind the gills. A longitudinal yellow line is on the dorso-posterior foot surface. The notum is rimmed by narrow yellow and white bands (color photo- graph in Lance, 1969: 37). MacFarianp (1966: 155) also described slight yellow markings on the posterior sur- face of the foot and on the notum posterior to the gills. Radula: Although Chromodoris mcfarlandi was named over 75 years ago, there are published descriptions of only 2 radulae. O’DoNoGHUE (1927: 90) reported about 62 rows of teeth, with about 50 teeth per half-row, and Mac- FaRLAND (1966: 155) gave a radular formula of 62 (47- 50°1°47-50). These reports indicate little of the vari- Table 4 Radular variation in Chromodorts mcfarlandi Width: Number Maximum number Specimen Length Width length of tooth of teeth per (HB numbers) (in mm) (in mm) ratio rows half-row 4 — = = 62 50 2 = = = 62 50 253 A 1.5] 0.48 1:3.15 46 32 253 B 1.43 0.51 1:2.8 4] 27 BS} (C, 1.11 0.69 1:1.61 4) 32 361 1.66 0.63 1:2.63 41 30 392 0.87 0.32 1:2.72 4] 23 393 2.95 1.56 1:1.89 76 49 394 75S) 1.03 1:2.48 57 50 395 1.76 0.73 1:2.41 44 32 396 0.78 0.32 1:2.44 36 16 452 1.72 0.83 1:2.07 50 34 453 A 1.818 0.808 1:2.25 51 38 453 B 1.66 0.67 1:2.48 56 = 454A 1.39 0.65 1:2.14 46 32 454 B 1.68 0.73 253, 48 31 456 A 21 1.01 1:2.08 52 36 456 B 1.94 1.05 1:1.85 51 34 457 A 1.66 1.01 1:1.64 43 36 457 B 1.64 0.909 1:1.8 60 35 458 2.1 1.05 1:2 52 43 XK 17 0.789 1:2.249 50.286 33.89 s 0.515 0.297 0.412 9.482 8.123 4O’DONOGHUE, 1927; 5MACFARLAND, 1966 Page 316 ability of the radular teeth counts. Table 4 gives the sizes, rows and teeth counts for 21 radulae. The combined radu- lar formula is 36-76 (16-50:1- 16-50). MacFarland and O’Donoghue examined specimens at the upper size range of the species. Their data were used by Boom (1976: ON fe 7D Figure 7 Chromodoris mcfarlandi radular teeth and sponge spicules found among the teeth A — HB 393; rachidian tooth; LBM, 0.02mm B — HB 394; lateral view of distal portion of 5** tooth from outer edge of radula (prominent cusp does not occur on the outer- most teeth) ; approximately row 31, RSR C — HB 394; 3" tooth from outer edge of radula; row 25, LSR D — HB 414 A; sponge spicule (0.071 mm long) found on radula E — HB 455; sponge spicule (monaxon, recurved ends with acces- sory points) found on radula; LBM, 0.04 mm THE VELIGER Vol. 20; No. 4 292) in his significant study correlating predator-prey morphology of dorid nudibranchs and sponges. His “radu- lar mean” column, however, should not be used uncritical- ly. The mean of C. mcfarlandi is given as 62 (49°0° 49). The 49 is based on only the numbers (47, 50, 50) given by O’Donoghue and MacFarland, and contrasts sharply with the mean number of maximum teeth per half-row (33-89) that I calculated based on many more speci- mens. Chromodoris mcfarlandi possesses a rachidian tooth. The expected positive correlation exists between the number of tooth rows and the maximum number of teeth per half-row (Figure 13). The regression line formula is Y = -4.47+0.799X; r = 0.8421, n = 20, P < 0.001. The number of tooth rows is dependent on radular length (Figure 14). The regression line formula is Y = 24.12+ 14.65 X; r = 0.8283, n = 19, P < 0.001. The maximum number of teeth per half-row is depend- ent on the radular width (Figure 15). The regression line formula is Y = 15.51+23.11 X; r=0.8664, n= 78, P < 0.001. O’DonocHUE (1927: 90) and MacFar.tanp (1966: 154-155) have described the morphology of the teeth. The rachidian is small and triangular-shaped (Figure 7-A). The innermost lateral teeth have 3 - 4 main den- ticles on the inner face of the shaft, 3 - 6 small ridge-like denticles on the inner side of the cusp, and 4 - 5 denticles on the outer face of the shaft (Figure 37). Lateral teeth in the middle of each half-row have a prominent cusp and numerous denticles (10- 16) on the posterior surface of the shaft (Figure 38). The outer lateral teeth (Figures 7-B, 7-C, 39) become smaller, with a greatly reduced cusp. Developing lateral teeth (Figure go) are thin and nar- row, with weak cusps and needle-like denticles. Discussion: Sponge spicules (Figures 7-D, 7-E) were found under the teeth in 2 radulae. BLoom (1976: 294) reports that Explanation of Figures 33 to 38 Scanning Electron Micrographs of the Radular Teeth of Chromodoris galexorum and Chromodoris mcfarlandi Figure 33: Chromodoris galexorum; rachidian and innermost lat- eral teeth (specimen HB 256) Xx 775 Figure 34: Chromodoris galexorum; inner lateral teeth (HB 256) xX 1300 Figure 35: Chromodoris galexorum; lateral teeth from middle and outer portions of halfrow (HB 256) xX a75 Figure 36: Chromodoris galexorum; cusps and distal portion of teeth from middle of half-row (HB 256) X 800 Figure 37: Chromodoris mcfarlandi; innermost lateral teeth (HB 253 C) X 1975 Figure 38: Chromodoris mcfarlandi; lateral teeth from middle of half-row (HB 253 C) X 1650 [BertscH] Figures 33 to 38 Tue VELIGER, Vol. 20, No. 4 Figure 33 Figure 38 Figure 37 Vol. 20; No. 4 Chromodoris mcfarlandi feeds on sponges from the genera Gellius and Haliclona. Chromodoris mcfarlandi is a beautiful nudibranch, im- mediately identifiable by its coloration and the large num- bers of denticles on the lateral teeth in the middle of the half-rows. It is a fitting tribute (even though the name is misspelled) to Prof. MacFarland and his magnificent detailed work on Californian nudibranchs, Chromodoris norrisi Farmer, 1963 (Figures 3-E, 8, 13 - 15, 41 - 46) References and Synonymy: Chromodoris norrisi FARMER, 1963: 81 - 84; plt. 1 a; text figs. 1 a-e. Marcus « Marcus, 1967: 170-173; figs. 21 - 23 (Material 1 only); 237-238. FARMER, 1968: 24-25, VII. Ev. Marcus « Er. Marcus, 1970: 198. BERTSCH, 1970: 8, 12; fig. 3. BertscH, 1971: 16. KEEN, 1971: 822, fig. 2331; plt. 20, fig. 4. Farmer, 1971: 109. Sphon, 1972b: 59. Berrscu et al., 1973: 292-293. BERTSCH, 1973: 108. Brusca, 1973: 174. BERTSCH, 1975: 105. BERTSCH, 1976b: 157 Glossodoris norrisi (Farmer). ABBOTT, 1974: 355; fig. 4246 Material Examined and Distribution: Baja California, Gulf Coast: 1) 1 specimen, Bahia de Los Angeles; leg. J. Lance, April 1968 (identified from a color transparency) 2) 1 specimen, subtidal, Notri (13km S of Loreto) ; leg. H. Bertsch, M. Ghiselin, & J. Allen, 4 July 1974 (HB 95) 3) 2 specimens, intertidal, Juncalito (19km S of Loreto) ; leg. H. Bertsch and B. Rose, 24 December 1973 (HB 40 A-B) 4) 5 specimens, subtidal 2-3m, Nopolo and Juncalito; leg. H. Bertsch, M. Ghiselin, & J. Allen, 27 June 1974 (HB 85 A-E) 5) 1 specimen, subtidal 1.5 - 3m, N end of Isla Santa Cruz; leg. D. Chivers, 26 June 1964 (HB 448; CAS) 6) 3 specimens, 14m subtidal, S end of Isla San Diego; leg. E. Janss, Jr., April 1974 (LACM) 7) 5 specimens, intertidal on mangrove roots, Isla San Jose; leg. G. G. Sphon, 2 April 1974 (LACM) 8) 1 specimen, Caleta San Evaristo, Isla de San Francisco; leg. G. G. Sphon, 1 April 1974 (LACM) 9) 1 specimen, 6m subtidal, Piedra del Saltito, 6km S of Puerto Mejia, La Paz; leg. A. J. Ferreira, 12 June 1974 (HB 376) 10) 7 specimens, 1 - 3m subtidal, Bahia Carisalito (4km N of Las Cruces) ; leg. H. Bertsch, T. Cooke, & G. Stellern, 26 July 1972 (HB 15 A-G) 11) 4 specimens, Las Cruces; leg. H. Bertsch, 1-18 July 1969 (HB 68 - 71) 12) 4 specimens, Las Cruces; leg. H. Bertsch, 1 July 1974 (HB 92 A-B, 93, 94) THE VELIGER Page 317 13) 6 specimens, 1.5-10m subtidal, N of Punta Gorda, 8 km S of Las Cruces; leg. H. Bertsch, 22 July 1972 (HB 14 A-F) 14) 1 specimen, N end of Isla Cerralvo; leg. H. Bertsch, 29 July 1969 (HB 72) 15) 1 specimen, subtidal, SW Isla Cerralvo; leg. H. Bertsch, 25 July 1972 (HB 19) Mainland Mexico, Gulf Coast: 16) 1 specimen, intertidal, Puerto Penasco, Sonora; leg. H. Bertsch, 24 December 1975 (HB 344) 17) 2 specimens, intertidal, Puerto Penasco, Sonora; leg. H. Bertsch, 26 and 29 December 1975 (HB 350 A-B) 18) 3 specimens, rocky intertidal, Guaymas, Sonora; leg. A. Kerstitch, 30 July 1966 (HB 323 A-C; USNM 753561) 19) 2specimens, Puerto Pefiasco and Guaymas, Sonora; leg. P Pickens & M. A. Hill, 7 August 1964 and 28 June 1965 (HB 322 A-B; USNM) The type locality of Chromodoris norrisiis Isla Cerralvo, The animal has been collected from the outer coast of Baja California and numerous localities within the Gulf of California (summarized in Bertscu et al., 1973: 292 to 293) ; the known range along the Gulf coast of Baja Cali- fornia is from Bahia San Luis Gonzaga to SW Isla Cerral- vo; it has been reported from Puerto Penasco to Guaymas along Mainland Mexico. Lot 6 represents a new bathymetric range for this species. External Morphology and Coloration: Large specimens of living Chromodoris nornst will reach 61mm (FarMer, 1963: 83). Between 1 - 19 July 1969, 22 - 26 July 1972, and 27 June -4 July 1974, 28 specimens collected near Loreto and Las Cruces varied from 21-50mm long alive (X = 33.5, s== 8.66). Six specimens collected during winter and early spring (24 December 1973, Loreto area; 24 December 1975, Puerto Pefiasco; April 1968, Bahia de Los Angeles) varied from 4-12mm total length alive (X = 8.7, s= 2.66). There is a significant difference (t 6.867, P <0.001) be- tween these 2 seasonal groups, that hints at possible pop- ulation cycles within the Gulf of California. As water warms in spring and summer, the animals grow to repro- ductive size, copulate and lay eggs. Settling of larvae may occur twice (in early summer and early autumn), with rapid summer growth and F, reproducing late summer. Those hatching in autumn metamorphose and are the smaller individuals found during winter. A year-round study (with biweekly or monthly sampling) is needed to test this hypothesis. Body background color is white. Some specimens show a violet coloration in the middle of the dorsum that is sub-epidermal in origin (FARMER, 1963: 81). The notum and lateral and posterior surfaces of the foot have numer- Page 318 THE VELIGER. Vol. 20; No. 4 Table 5 Radular variation in Chromodoris norrist Width: Number Maximum number Specimen Length Width length of tooth of teeth per (HB numbers) (in mm) (in mm) ratio rows half-row © — — — 65 44 a = = _ 68 48 4A 3.39 1.74 1:1.95 68 58 14B 3.29 1.58 1:2.08 69 52 14C 3.84 1.86 £ 1:2.06 71 58 14E 3.58 1.88 1:1.9 74 63 14 F 3.696 1.96 1:1.89 71 60 15 A 3.11 1.62 1:1.92 76 58 15B 3.8 1.8 1:2.) 75 64 15C 3.15 1.76 1:1.79 70 56 15 D 4.02 2.22 1:1.81 80 62 ISDE 2.83 1.82 1:1.55 67 62 IDF 3.37 2.0 1:1.69 68 59 15G 3.47 1.74 1:1.99 68 56 19 = os — 65 54 40 A 0.76 0.299 1:2.54 36 25 40 B 1.13 0.57 1:1.98 50 40 68 3.92 2.08 1:1.88 71 55 69 4.1 2.1 1:1.95 73 61 70 4.55 2.32 1:1.96 85 67 7\ 3.66 1.76 1:2.08 65 62 72 3.9 1.9 1:2.05 76 62 85 A 4.6 2.16 1:2.13 86 71 85 B 3.72 1.96 1:1.9 67 52 85 C 3.96 Dy 1:1.8 79 63 85 D Sh 2.08 1:1.78 69 55 85 E 2.95 1.53 1:2.06 64 51 92 A 4.06 2.99 1:1.83 79 67 92 B 4.24 23 1:1.84 76 69 93 3.9 2.02 1:1.93 68 58 94 3.76 2.22 1:1.69 74 56 95 4.44 1.94 1:2.29 83 56 322 A 2.08 1.35 1:1.54 69 55 322 B 1.15 0.77 1:1.49 49 38 323A 3.37 1.9 1:1.77 80 66 323 B 3.64 2.08 1:1.75 97 72 323 C 3.01 1.86 1:1.62 77 58 344 ].07 0.57 1:1.88 48 39 350 A 2.02 0.87 1:2.32 5] 37 350 B 1.74 1.01 1:1.72 53 40 x 3.26 1.73 1:1.905 69.5 55.725 s 1.015 0.525 0.217 11.59 10.233 ®Marcus & Marcus, 1967; 7FARMER, 1963 Vol. 20; No. 4 (, pt 8-C fi 8-B 8-A — 8-E 8-F 8-G 8-I THE VELIGER Page 319 8-R 8-S Figure 8 Radular teeth of Chromodoris norrisi A — HB 4oB; outline sketch of entire radula prior to flat-mounting B — HB 14 A; dorso-lateral view of 2"4 tooth, row 55, RSR C — HB 14 A; 3" lateral tooth, row 55, RSR D — HB 14 A; 5* lateral tooth, row 55, RSR; adjacent inner tooth overlaps the lateral basal flange E — HB 14 A; outer face of 4** lateral tooth, row 60, LSR F — HB 14 F; 2"4 tooth, row 64, RSR G — HB 69; inner face of 5‘* tooth, row 66, RSR H — HB 15 B; dorso-lateral view of 5" lateral tooth, row 74, RSR; LBM, 0.057mm I — HB 323 B; distal portion of shaft, tooth 28, approximately row 30 J - HB 15 B; distal portion of shaft, tooth 26, row 69, LSR K — HB 323 B; distal portion of shaft, 5' tooth from outer edge of radula, approximately row 32 ous small red dots; the notum also has about 4 as many yellow dots. A bright orange broken band encircles the edge of the notum. There is a wide range of variation in the number and size of the dots from very many small ones to fewer, larger spots (BERTSCH et al., 1973: 293). L — HB 323 C; approximately tooth 13, row 28 M - HB 323 CG; 10" tooth from outer edge of radula, row 28 N - HB 323 C; 5‘ tooth from outer edge of radula, row 28 O - HB 14 A; distal shaft of developing 1° lateral tooth, last row, LSR (outer face with heavier denticulation) P — HB 14 A; distal shaft of developing 3™ lateral tooth, last row, LSR Q - HB 14 A; distal shaft of developing 4" lateral tooth, last row, LSR R — HB 14 A; shaft of developing lateral tooth, approximately 4 the distance from radular outer edge to center of radula; last row, RSR S — HB 14 B; shaft of developing 27 lateral tooth, drawn to same scale as Figure 8 R; last row, RSR Color photographs appear in FARMER (1963; plt. 1a) and KEEN (1971: plt. 20, fig. 4). Radula: FARMER (1963) and Marcus & Marcus (1967) have Page 320 published radular formulae of Chromodoris norris. Al- though Farmer gives a range of variation for 6 specimens, he only gives the holotype formula separately. This is the only count from Farmer’s paper that can be analyzed statistically. Table 5 presents the individual radular sizes and counts of the 2 published radular formulae and the 38 radulae I examined. The new combined radular formula is 36-111 (25-72° 1° 25-72). Least squares regression analysis shows that the number of tooth rows and maximum number of teeth per half row are positively correlated (Figure 13). The equation, Y —-0.147+0.804 X, describes the regression line. The coefficient of correlation is 0.9106 (P < 0.001, n= 40). The number of tooth rows is positively correlated with the radular length (Figure 14). The regression line formu- la is Y —36.84+10.07X%, and r=0.8513 (P < 0.001, N= a7) The radular width and maximum number of teeth per half-row (Figure 15) are positively correlated, with r= 0.8965. The formula describing the regression line is Wes NE SoG] DX (JO KCOWOIONN, 10 === B77) Figure 8-A is an outline sketch of the entire radula; the smaller posterior portion has not been flattened. The innermost lateral tooth has approximately 5 inner and 5 outer denticles. Inner lateral teeth have a thick, squat shaft, with 4-9 denticles on the outer face (Figures 41 and 42). Toward the middle region of the halfrow, the teeth become longer with more erect shafts. The denticles change from thick lateral structures to pointed prongs (7 - 14 in number) on the posterior surface of each tooth, and the cusp becomes quite prominent (Figures 8I - 8L). Towards the outer margin the teeth are reduced in size, and cusps and denticles become smaller (Figures 8M, 8N, 43). Denticles on the outer and posterior surfaces averaged 8.19 across a half-row (n = 32; of the 66 teeth in the half-row, only half were lying in such a way as to allow denticle counts). The length of the cusps across a half-row averaged 0.0154mm (range, 0.006 - 0.022mm, n = 53). The mean ratio of the length of the first denticle to the length of the THE VELIGER Vol. 20; No. 4 cusp is 1 :4.517 (range from 1.17 on the extreme endsof the halfrow to 11 in the central region of the halfrow; s= 254.20 ——1530\e The radulae of 18 specimens (47% of the Chromodoris norrisi radulae dissected for this study) had a double- denticulation pattern on the first 2-9 innermost teeth (Figures 8B - 8H). In addition to denticles on the outer face of the shaft, these teeth had 2-8 denticles on the inner face of the cusp (K =5.1, n= 45). This is the same as the double-denticulation pattern described for the first 4 lateral teeth of C. sphoni. Chromodoris norrist specimens 14 A, E, F, 15 B, had an average of 6.7 double- denticled teeth (n = 39) on each half-row in the posterior 16, 6, 7, and 6 rows respectively. This pattern was not visible in the anterior half of the radulae. Patterns of tooth growth are shown in Figures 44 and 8-O to 8-S (progressing from the innermost laterals to teeth nearer the middle of the half-row). The early devel- opment of the central lateral teeth is from thin shafts with just the hint of posterior denticulation (Figure 45) to stronger shafts with needle-like denticles (Figure 46). The scanning electron micrographs (Figures 41 - 46) offer an immediate visual comparison of developing and fully-formed teeth. Discussion: Chromodoris norrisi is readily separated from the re- lated C. baumanni. Visual examination of their radulae reveals immediate perceptual differences. The cusps of the middle lateral teeth of C. norrisi are longer and more pro- nounced than the C. bawmanni cusps (compare Figures 8-I to 8-K with Figures 4-H, 4-1). This gestalt impression can be proved statistically. The length ratio of the first denticle:cusp is significantly different (longer) for C. norrisi than C. baumanni (t=5.78, PBM[NH]) Columbella schrammi Petit de la Saussaye, 1853. Journ. de Conchyl. 4: 364; plt.12, figs. 3, 4(Point- a-Pitre, Guadéloupe, F W.I.; holotype Mus. Rouen) Nitidella elegans Dall, 1871. Amer. Journ. Conchol. 7 (2): 122 (Panama - adventitious; holotype, USNM 182582) Shell small (6- 9mm); spire high (about 2 total shell length), acute; whorls flat-sided, suture very shallow; body whorl sub-cylindrical, aperture moderately broad, interior of apertural lip denticulate, columella straight, weakly denticulate, siphonal canal short, slightly bent; sculpture entirely absent; color white with chestnut brown longitudinal bars. Each radular row consists of a flat, sub- rectangular median tooth flanked on each side by a sig- moid bicuspid lateral tooth with a proximal rounded pro- jection (Figure 42). Remarks: Although said by Sowerby to be a part of the Cuming Collection, the type of this species was probably given or sold to a private party prior to the acquisition of the Cuming Collection by the British Museum (Natural History). According to Dr. Norman Tebble, former head of the Mollusca Section, it is not in the British Museum (N THE VELIGER Vol. 20; No. 4 H), nor has it ever been catalogued in their collection. Its present whereabouts is unknown. A table of size range for Mitrella dichroa shows a cline with regard to shell length. The specimens from the north- ern end of its range are small (6.0 - 6.4mm) and exhibit white and brown punctations overlying the typical zebra- striped color pattern. This form has beencalled M. parvula and M. argus. Specimens from the southern end of the species’ range are appreciably larger (7.0- 8.0mm) and generally have no overlaid pattern of punctations. This form has been called M. dichroa. These color patterns can be shown to intergrade gradually if specimens from local- ities throughout the geographical range are available. No fossil examples of this species are known to me. Range: Mitrella dichroa ranges from Palm Beach Coun- ty, Florida, to the Lesser Antilles and from Vera Cruz, Mexico, to southern Brazil. Mitrella ocellata (Gmelin, 1791) (Figure 14) Voluta ocellata Gmelin, 1791. Syst. Nat. Ed. 13: 3455 (type locality here designated as Nassau, New Providence Island, Bahama Islands; representation of lectotype, MarTINI & CHEMNITZ, Neues Syst. Conch.-Cab. 4: plt. 150, fig. 1409) Buccinum cribrarium Lamarck, 1822. Hist. Nat. Anim. s. Vert. 7: 274 (type locality, Les mers de Java; holotype, GM) Columbella guttata Sowerby, 1832. Proc. Zool. Soc. London f. 1832: 118 (type locality, Panama, Pacific; holotype, BM[NH]) Shell moderate in size (11-13mm), fusiform; spire ? of total shell length, acute when complete (shell is usu- ally decollate), whorls flat-sided, suture shallow; body whorl cylindrical, aperture moderately broad, apertural lip slightly thickened, denticulate on its inner surface, columella straight, smooth, siphonal canal very short, slight anal canal present; sculpture lacking, color usually consisting of white punctations on a black background. Other color variations include a completely yellow-white Explanation of Figures 43 to 48 Radular Dentition Figure 43: Mitrella ocellata (Gmelin, 1791) Figure 44: Suturoglypta albella (C.B. Adams, 1850) Figure 45: Suturoglypta pretri (Duclos, 1846) Figure 46: Suturoglypta iontha (Ravenel, 1861) Figure 47: Strombina pumilio (Reeve, 1859) Figure 48: Strombina recurva (Sowerby, 1832) Tue VELIcER, Vol. 20, No. 4 [Rapwin] Figures 43 to 48 ' \ 1 yh 5 t . i AS Dy Vol. 20; No. 4 shell and one in which the background is yellow or light brown with darker spots. Each radular row consists of a flat, subrectangular rachidian tooth flanked on each side by a sigmoid, bicuspid lateral tooth (Figure 43). Remarks: According to DaNcE (1967), the figures re- ferred to by Gmelin were based on specimens in the Spengler Collection and these must be considered the types of Gmelin’s species. The specimen from which Gmelin’s Voluta ocellata was described and upon which the figure, cited by Gmelin, in Martini « Chemnitz was based, is said to be in the Spengler Collection (Copen- hagen Museum). Often assigned to Nitidella, REHDER (1962) has correctly assigned this species to Mitrella. In addition to shell characters, major radular differences exist between Mitrella ocellata and Nitidella nitida, the type species of Nitidella. Most striking is the disparity in the width and degree of flexure of the lateral radular teeth. The shell of Mitrella ocellata is extremely variable with regard to color, color pattern, and overall propor- tions throughout its very wide geographic range. A slen- der form occurring in parts of the Panamic-Pacific re- gion is known as M. delicata Sowerby, and the dark, stub- by form from the coast of East Africa is known as M. ocellata nomadica (Melvill). There is also a very dark, stout form from the Galapagos Islands which is known as Nitidella guttata baileyi Bartsch & Rehder, 1939. It ap- pears to be quite similar to certain populations of M. ocel- lata on the basis of shell characters but has a markedly different lateral radular tooth and appears to be a com- pletely distinct species. The only fossil examples of this species are known from the Pleistocene of Venezuela. Wooprinc (1928) de- scribed a form from the Miocene of Jamaica as Mitrella ocellata bowdenensis and noted that it is so similar in shell characteristics to the living form that, perhaps, it should not be separated even subspecifically. Range: This species, in one or another of its forms, is known from tropical areas almost entirely around the world. In the eastern Pacific it ranges from the central Gulf of California to Panama and in the western Atlantic it ranges from Palm Beach County, Florida and the north- Bahama Islands to Trinidad and Panama. In addition, I have seen scattered Museum records from the eastern and western coasts of Africa. Strombina Morch, 1852 Strombina Morch, 1852. Cat. Conch. Yoldi: 85 (type species by SD (CossMann, 1901: 241) Colum- bella lanceolata Sowerby, 1832) (see Figure 18) THE VELIGER Page 339 Strombocolumbus Cossmann, 1901. Essais Paléo- conch. Comp. 4: 241, objective synonym of Strom- bina Morch, 1852 Shell large (20 - 40mm) and fusiform; spire moderately high, almost 4 total shell length), acute, whorls convex, shouldered, suture impressed; body whorl large, angulate, aperture long, narrow, apertural lip usually thickened with strong denticulate plates on its inner surface, colu- mella bent, often with a strongly denticulate callus, siphonal canal short to moderate in length, bent; sculp- ture generally consisting of a row of strong axial knobs below the suture which become ephemeral anteriorly; color variable with whites, browns, and purples predom- inating. Each radular row consists of a flat, subrectangu- lar median tooth flanked on each side by a long, sigmoid lateral tooth similar in shape to those of Anachis (see Fig- ure 48 for extrapolation to Strombina lanceolata). Remarks: Considering the wealth of fossil species of this genus in the western Atlantic, the survival of only a single, rarely encountered species suggests that Strombina is ap- proaching extinction in that region. Indeed, this genus, once widely distributed, seems to be flourishing today only in the Panamic Province of the eastern Pacific. In this it is not alone. A number of molluscan genera, represented sparsely or not at all in other regions, have undergone remarkable adaptive radiation in this region. Strombina pumilio (Reeve, 1859) (Figure 19) Columbella pumilio Reeve, 1859. Conch. Icon. 11 (Columbella) : species 147; plt. 24, fig. 147 (type locality Cumana, Venezuela; holotype ?>BM[NH]) Strombina terquemi Jousseaume, 1876. Mém. Soc. Zool. France 1: 265 - 266; plt. 5, figs. 1, 2 (type locality East Africa [erroneously]; holotype not located ) Strombina caboblanquensis Weisbord, 1962. Bull. A- mer. Paleo. 42 (193): 323 (type locality, Venezu- ela — Pleistocene; holotype, Paleo. Res. Inst., I- thaca, N. Y.) Shell moderately large (15- 20mm), stromboid; spire high, acute, whorls moderately convex, suture impressed, body whorl large (2 of total shell length), ventricose, shouldered, aperture long and narrow, apertural lip strongly thickened, bearing a long denticulate plate on its inner surface, columella straight, non-denticulate, with a slight callus, whose raised edge extends along the anterior % of the aperture, siphonal canal short, bent; shell surface smooth; sculpture of axial riblets ending in round knobs Page 340 at the shoulder of the body whorl and, in some instances, on the spire whorls, spire rarely with spiral threads; the periostracum is thin, brown, parchment-like. Shell color is chestnut-brown with irregular white blotches. Each rad- ular row consists of a flat, subrectangular rachidian tooth flanked on each side by a sigmoid, bicuspid lateral tooth (Figure 47). Remarks: The earliest fossil example of this species 1s reported from the Pliocene of Venezuela (WEISBoRD, 1962). Range: ‘This species ranges from Jamaica to the north- ern coast of South America. Suturoglypta Radwin, 1968 Anachis (Suturoglypta) Radwin, 1968. Proc. Biol. Soc.Wash. 81: 145 (type species by OD, Colombel- la pretri Duclos, 1846) (see Figure 22) Shell small (4- 8mm), fusiform; spire high, whorls flat- sided, suture squarely incised; body whorl moderately small, aperture narrow, constricted anteriorly, apertural lip swollen, denticulate interiorly, columella slightly bent, non-denticulate, siphonal canal moderate to long; sculp- ture predominantly axial, ribs distinctly raised, almost square in cross-section; color wax yellow to white with pale brown axial flammules and, in one species, entirely translucent red-brown. Each radular row consists of a flat, subrectangular rachidian tooth flanked on each side by a sigmoid bicuspid lateral tooth (Figure 45). Remarks: Species in this group show great variability in the number and strength of the axial ribs. The 3 spe- cies included here may be distinguished by slight shell differences and consistent radular divergences. The most diagnostic features are the distinctly fusoid shell form, the strong, square-cut axial ribs, and the squarely incised suture. Suturoglypta albella (C. B. Adams, 1850) (Figure 27) Pleurotoma albella C. B. Adams, 1850a. Contrib. to Conch. 1 (4): 63 (type locality, Jamaica; holotype, MCZ 186006) Shell small ( 4- 6mm) ; spire moderately high (about 2 of total shell length) and acute, whorls slightly convex, more so toward the body whorl, suture deep, squarely THE VELIGER Vol. 20; No. 4 chiseled; body whorl subcylindrical, aperture narrow, a- pertural lip slightly thickened, denticulate on its inner sur- face, columella bent, weakly denticulate, siphonal canal moderately long, somewhat bent, distinct anal sinus pre- sent; sculpture of square-cut prominent axial ribs with no spiral elements, color pure white, in some instances with tust-brown blotches or even entirely red-brown. Each radular row consists of a flat, subrectangular median tooth flanked on each side by a sigmoid lateral tooth. Each of the lateral teeth has 2 sharp distal cusps and 1 more rounded proximal lobe (Figure 44). Remarks: Assigned originally to the turrid genus Pleuro- toma by C. B. Adams because of its size and marked anal sinus, this species is the smallest, least common species in Suturoglypta. Its distribution is apparently restricted to the Bahama Islands and the Greater Antilles. Suturo- glypta albella differs from S. iontha and S. pretri by its smaller size, its more convex spire profile, its geographic distribution, its consistently shorter siphonal canal, and minor radular differences. No fossil examples of this species are known to me. Range: This species ranges from the central Bahamas Islands to Grand Cayman Island. Suturoglypta iontha (Ravenel, 1861) (Figure 23) Columbella iontha Ravenel, 1861. Proc. Acad. Nat. Sci. Phila. 3: 41 - 42 (type locality off Charleston, South Carolina; holotype apparently destroyed) Anachis acuta Stearns, 1873. Proc. Acad. Nat. Sci. Phila. for 1873: 344-347 (type locality, Tampa Bay, Florida; holotype, USNM 53779) Shell moderately small (6-10mm); spire high (2 of total shell length), acute, whorls flat-sided; body whorl broad, fusoid, aperture moderately narrow, apertural lip thickened, flaring, denticulate interiorly, columella weak- ly denticulate, strongly bent, distinct anal sinus present; sculpture consists of strong, square-cut axial ribs, varying in number and spacing; color wax-yellow, in many cases with spiral strokes of chestnut-brown. Each radular row has a flat subrectangular median plate, flanked on each side by a sigmoid, bicuspid lateral tooth (Figure 46). Remarks: Although Dance (1967) claims that the Ra- venel Collection is in the mollusk collection of the Charles- ton (South Carolina) Museum, according to Mrs. Amelia Scheltema, who visited the museum in 1965, the collec- Vol. 20; No. 4 tion is not there (A. Scheltema, personal communication). Ravenel’s entire collection was apparently moved from Charleston to Atlanta to avoid the Union Army’s depre- dations, only to be burned with the remainder of that city. Apparently, the type of Suturoglypta iontha was de- stroyed at that time. This species differs from Suturoglypta pretri, its closest relative, in having a broader and more foreshortened body whorl, a shorter, more bent, siphonal canal, differing de- tails of axial ribbing, a distinctly more northern geograph- ical distribution, and minor radular distinctions. No fossil examples of this species are known to me. Range: This species ranges from off Cape Hatteras, North Carolina to Key West, Florida and to South Padre Island, Texas. Suturoglypta pretri (Duclos, 1846) (Figure 22) Colombella pretrii Duclos, 1846. (in Chenu) Illust. Conchyl. 4: plt. 16, figs. 7, 8 (type locality here designated as Charlotte Amalie Harbor, St. Tho- mas; representation of lectotype, Ductos, 1846, 4: pit. 16, figs. 7, 8). Columbella mangelioides Reeve, 1859. Conch. Icon. 11 (Columbella): plt. 31, fig. 197 (type locality, West Indies; holotype, BM[NH]) Anachis samanensis Dall, 1889. Bull. Mus. Comp. Zool. 18: 188 (type locality, Samana Bay, Domini- can Republic; holotype, USNM 54285) Columbella pretrei Kobelt, 1897. Martin & Chemn. Conchyl.-Cab. 3-1d: 252, 319; plt. 34, fig. 4; plt. 41, fig. 7 (error for C. pretri) Shell moderately small (6-gmm), spire high (about 2 total shell length), acute, whorls flat-sided, suture square- ly incised and moderately deep; body whorl fusoid, aper- ture moderately narrow, apertural lip thickened, flaring, denticulate within, columella denticulate, bent, covered with a callus, siphonal canal moderately long, bent; anal sinus present. Sculpture of strong, square-cut axial ribs, color wax-yellow with brown spiral markings. Each radu- lar row consists of a flat, subrectangular median tooth flanked on each side by a sigmoid lateral tooth. Each lateral has 2 sharp distal cusps and a rounded proximal lobe (Figure 45). Remarks: Suturoglypta pretri, the type species of the genus, differs from S. iontha in having fewer and more broadly spaced axial ribs, a less inflated body whorl, less THE VELIGER Page 341 bent siphonal canal, a smaller apical angle, and a less thickened apertural lip. There are also minor radular differences. Suturoglypta pretri differs from S. albella in its consistently larger size, higher, more acute spire, flatter spire profile, and greater length and degree of curvature of its siphonal canal. Populations of this species from the western end of Cuba and from Yucatan are very variable with regard to sculpture. Lots have been examined in which sculpture ranged from typical, strong, square-cut axial ribbing to weak, sinuous axial ribs to an almost complete absence of sculpture with complete intergradation between these forms. The earliest fossils of this species have been reported from the Pleistocene of Panama. Range: ‘This species ranges from southeastern Florida and eastern Mexico to Barbados and Curagao. OruHerR REeporTED WESTERN ATLANTIC SPECIES Several other columbellid species have been described from the Magellanic province of South America. Efforts to obtain specimens of these species have been almost entirely unsuccessful. The most extensive report on the mollusks of this area was made by STREBEL (1905) in which 3 columbellid species were originally described and figured. These are the earlier treated Parvanachis meluvil- lec (Figure 29), P paessleri (Figure 28), and PR decorata (including the subspecies P decorata inornata (Figure 27)). The first 2 species were mistakenly placed in the subgenus Seminella Pease. Another species, Columbella rubra Martens, 1881 (Figure 26), also inhabits the Mag- ellanic province. Although no specimens were available for examination, the figure of this species accompanying the original description by Strebel suggests an affinity with the genus Mitrella Risso. The adventitious occurrence of gastropod shells is no novelty in the history of malacology. During the period of history when the wooden sailing vessel was the sole form of transoceanic transportation, ship-bottom ballast in the form of beach sand and rocks was often necessary. Other material, such as beach-drifted shells, was often included and the entire load of ballast was then unloaded at another port of call where the vessel replaced the bal- last with cargo. Such a ballast origin may be suspected in the case of 2 supposedly western Atlantic species. The most obvious instance concerns Anachis ter psichore Sowerby, 1824. The type locality is western Atlantic and a number of worn Page 342 shell specimens from several West Indian localities are in the collection of the Division of Mollusks, National Mus- seum of Natural History. In none of these cases were the specimens collected alive. A large number of live-collected and freshly dead specimens from the east coast of India has been examined. This appears to be the true habitat of the species. A number of spurious Caribbean records of A. terpsichore has also been based on misidentification of A. lyrata Sowerby, 1833, another large species of Anachis that superficially resembles A. terpsichore. It differs from the latter species in having more regular spiral brown color bands and more convex whorls; those of A. terpst- chore are flat at the periphery. Anachis lyrata occurs along the Pacific coast of Central and South America and on the Atlantic coast of Brazil. In addition, a few worn, empty shells have been collected in the Caribbean. These may also be examples of adventitious distribution. Several empty shells of Costoanachis varia, a large spe- cies found throughout most of the Panamic province, have been collected at a single locality (Wounta Haulover, Nic- aragua) on the Caribbean coast of Central America. An apparently close relative of C. varia, C. asphaltoda Mau- ry, was common in Trinidad and elsewhere in the Carib- bean during the Pliocene, but there is no clear indication that C. varia or C. asphaltoda occurs today in the Carib- bean. The shells from the above locality are moderately fresh and may have been carried across the narrow part of Nicaragua by traders or other travelers, or they may be additional ballast-shell records. Until this species is found living in the Caribbean, its status as a bona fide western Atlantic species will remain doubtful. The fate of the type specimens of the species described by Edmund Ravenel provides an ironic commentary on the Civil War period. SHERBORN (1940) located the above types at the Charleston Museum of Natural Histo- ry. Authorities of the CNHM have been unable to locate this collection. As accurately as can be determined, during the Civil War the Ravenel collection, including the types, was moved from Charleston to protect it from Union troops then thought to be threatening the city. The col- lection was apparently moved to Atlanta, Georgia, shortly before Sherman’s troops burned that city. This circum- stance does not present a major difficulty in most cases. In one case, however, Ravenel’s description is so unclear, ambiguous, and generally inadequate that there is doubt as to the species’ identity. This is Columbella similis Ra- venel, 1861. There are 2 apparently undescribed Costo- anachis species or subspecies on the Atlantic and Gulf coasts of the United States. One is probably an unusual form of C’. avara and the other may be the C. similis of THE VELIGER Vol. 20; No. 4 Ravenel. It is unfortunate that with no extant type, an inadequate description, and no published figure, the as- signment of this name to any biological entity is impossible. Literature Cited ADAMS, CHARLES BAKER 1850a. Notes on the synonymy of certain marine shells. Contrib. to Conch. 1 (4): 54-55 (January 1850) 1850b. Descriptions of supposed new species of marine shells which in- habit Jamaica. Contrib. to Conch. 1 (4): 56-68 (January 1850) 1851. Description of new species and varieties of shells which inhabit Jamaica. Contrib. to Conch. 8: 129 - 140 (March 1851) ADAMS, HENRY & ARTHUR A. ADAMS 1853-1858. The genera of Recent Mollusca arranged according to their organization. 3 vols., John van Voorst, London vol. 1: i- xl + 1-484 Ancas, GEeorGE FRENCH 1867. Descriptions of thirty-two new species of marine shells from the coast of New South Wales. Proc. Zool. Soc. London 35: 110 - 117; pit. 13 (24 January 1867) BartscH, Paut & Haratp ALFRED REHDER 1939. Mollusks collected on the Presidential Cruise of 1938. Smithson. Misc. Coll. 98 (10): 1-18; 5 plts. (13 June 1939) BiaInviLLe, Henri Marie Ducrotay DE 1829. Faune Frangaise Mollusca. Levrault, Paris: 1-320; 41 pits. Broccui, Giovanni BaTTISTA 1814. Conchologia fossile subappenina con osservazioni geologiche sugli Appennini e suolo adiacente. Milano; 2 vols.; 712 pp.; 16 plts. Brown, Amos P & Henry AucustTus PILSBRY 1913. Two collections of Pleistocene fossils from the Isthmus of Pana- ma. Proc. Acad. Nat. Sci. Phila. 65 (1913): 493-500; figs. 1-3 (19 August 1913) Buceuoy, E., Pau, DauTzeNBEeRG & GusTAveE DoLiFus 1884 (1882-1886). Les mol!usques marins du Rousillon. 1 (5): 1-570, atlas 66 plts. Gastropodes. Cater, H. 1864. Description d’une espéce nouvelle. Journ. de Conchyl. 12: 279; plt. 2 (1 July 1864) CaNnTRAINE, FRANGOIS JOSEPH 1835. Notice contenant les diagnoses ou descriptions succinctes de quel- ques espéces nouvelles de mollusques. Lettres Bruxelles, II (11): 380-401 CasTELLANOS, ZULMA DE & DELICIA FERNANDEZ 1967. Nuevos aportes a los Pyrenidae de Argentina (Mollusca, Gastro- poda). Neotropica 13 (41): 49-52; 1 fig. (1 August 1967) CossMANN, ALEXANDER EDouARD MAurRICE 1901. Essais de paléoconchologie comparée. Paris, 4: 1 - 293; plts. 1-10 (October 1901) Cox, Lesiie REGINALD 1927. Mollusca in Report on the paleontology of the Zanzibar Pro- tectorate. pp. 13-102; plts. 3-19 (September 1927) DALL, WILLIAM HEALEY 1871. Descriptions of sixty new forms of mollusks from the west coast of North America and the North Pacific Ocean, with notes on others already described. Amer. Journ. Conchol. 7 (2): 93-160; plts. 13 - 16 (2 November 1871) 1881. Preliminary report on the Mollusca. In: Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico, and in the Caribbean Sea, 1877-79, by the United States Coast Survey steamer “Blake” ... Bull. Mus. Comp. Zool. 9 (2): 33-144 (July to December 1881) 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Carib- bean Sea (1879-80), by the U.S. Coast Survey steamer “Blake,” (etc.) XXIX. Report on the Mollusca.—Part II. Gastropoda and Scapho- Bull. Acad. Roy. Sci. Belles poda. Bull. Mus. Comp. Zool. Harvard 18: 1-492; plts. 1-40 (January to June 1889) 1900. Some names which must now be discarded. The Nautilus (4): 44-45 (1 August 1900) Vol. 20; No. 4 Dax, Wittiam HeEAtey 1912, New California Mollusca. The Nautilus 25 (11): 127-129 (8 March 1912) Proc. U.S. (11 June 1913) 1913. Diagnoses of new shells from the Pacific Ocean. Nat. Mus. 45 (2002): 587 - 597 1916. Notes on west American Columbellidae. The Nautilus 30 (3): 25-29 (14 July 1916) 1919. New shells from the northwest coast. Proc. Biol. Soc. Washington 32: 249 - 251 (31 December 1919) 1924. Notes on molluscan nomenclature. Proc. Biol. Soc. Wash. 37: 249 - 252 (21 February 1924) 1927. Small shells from dredgings off the southeast coast of the United States by the United States Fisheries steamer ‘“‘Albatross” in 1885 and 1886. Proc. U. S. Nat. Mus. 70 (2667) (art. 18): 1-134 (20 April 1927) Dati, Witiiam Hearey & CHARLES TorREY SIMPSON 1901. The Mollusca of Porto Rico. U.S. Fish Comm. Bull. 20 (1): 351 - 524; plts. 53 - 58 (November 1901) Dance, STANLEY PETER 1967. Shell collecting, an illustrated history. Berkeley & Los Angeles, pp. 1 - 344; plts. 1-35 De Kay, James ELtSwortH 1843. Natural History of New York, prt. 1: Zoology of New-York, or the New-York fauna; ..., prt. 5: Mollusca. Albany (Carroll & Cook) [viii] +271 pp.; 40 plts. [1844 ?] Duc os, P. L. in JEAN CHARLES CHENU 1846 [tn Chenu] [Illustrations conchyliologiques. Paris, 4: plts. 1 - 27 Dunxker, WILHELM BERNHARD RuDOLPH HADRIAN Univ. Calif. Press, 1847. Diagnoses Buccinorum quorundam novorum. Zeitschr. f. Malakozool. 4: 59 - 64 (April 1847) 1853. 1853. Diagnoses molluscorum novorum. Zeitschr. f. Malakoz. 10: 110-112 GisteL, JOHANNES NerpomuK FrANz XAVIER VON 1848. Naturgeschichte des Thierreichs fiir hoéhere Schulen. Stutt- gart, pp. i- xvi, 1 - 216; plts. 1 - 32 Gme.in, JoHANN FriepRicH 1791. Caroli Linnaei systema naturae per regna tria naturae ... Edi- tio decima tertia, aucta, reformata, Vermes Testacea. Leipzig 1 (6): 3021-3910 (pre May 1791) Gou.p, Aucustus ADDISON 1841. Report on the Invertebrata of Massachusetts, comprising the Mollusca, Annelida, and Radiata. Cambridge, Mass., vit+373 pp-; 15 plts. (comprising 213 figs.) 1851. [Descriptions of new shells from California, collected by Maj. William Rich and Lieut. T. P Green, U.S.N.] Proc. Boston Soc. Nat. Hist. 4: 87 - 93 (November 1851) 1859-1861. Descriptions of shells collected in the North Pacific Explor- ing Expedition under Captains Ringgold and Rodgers. Proc. Bos- ton Soc. Nat. Hist. 7: pp. 382-384 (December 1860); pp. 385 - 389 (January 1861) [dating: Richard I. Johnson: The Recent Mollusca of Augustus Addison Gould. U.S. Nat. Mus. Bull. 239. 1061] Grant, Utysses Simpson, IV a Hoyt Ropney GaLz 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and adjacent regions. San Diego Soc. Nat. Hist. Mem. 1: 1- 1036; 15 text figs.; plts. 1 - 32 (3 November 1931) Gray, Joun Epwarp 1847. A list of the genera of Recent Mollusca, their synonyma and types. Proc. Zool. Soc. London (for 1847) 15 [part 15] (178): 129-219 (November 1847) Guppy, Rosert JoHN LECHMERE 1867. On the Tertiary fossils of the West Indies ... Assoc. Trinidad 3: 145-176 JEFFREYS, JOHN Gwyn 1867. British Conchology pits. 1-8 JoussEAuME, FE.Ix PierRE 1876. Description des mollusques recueillis dans la Mer Rouge et le Golfe d’Aden. Mém. Soc. Zool. France 1:165 - 223 Kose.t, WILHELM 1887. in Martini & Cuemnitz, Conchylien-Cabinet. Columbella. von Bauer & Raspe, Niirnberg, 3-ld: 1-344; plts. 1-44 Lamarck, JEAN BapTisTE PIERRE ANTOINE DE MONET DE 1822. Histoire naturelle des animaux sans vertébres. 7 [Mollusques] Paris 1-711. Linnaeus, CAROLUS 1758. Systema naturae per regna tria naturae. Ed. 10, reformata. Hol- miae 1 (1): i-iii+1 - 824 Maury, CarLotta JOAQUINA 1917. Santo Domingo type section and fossils. Part 1, Mollusca. Bull. Amer. Paleontol. 5 (29): 165-415; plts. 1-39 (number 29 also paged separately: 1 - 251); 39 plts (March-April 1917) (July 1853) Proc. Sci. Van Voorst, London, 4: 1 - 486; THE VELIGER Page 343 Mevviti, James Cosmo « Ropert STANDEN 1901. The Mollusca of the Persian Gulf, Gulf of Oman and Arabian Seal aie Proc. Zool. Soc. Londn,1901 (2): 327 - 460; pits. 21 - 24 (18 June 1901) Mo ter, Hans Peter CHRISTIAN 1842. Index molluscorum groenlandiae. 76 - 97. Copenhagen Morcn, Otto ANDREAS Lowson 1852. Catalogus conchyliorum quae reliquit D. A. d’Aguirra et Gadea Comes de Yoldi. Klein, Hafniae, prt. 1: 1 - 170 1859. Note sur les dents linguales du genre Columbella. Conchyl. 7: 254 - 262 Otsson, AxeL ADoLF & ANNE HARBISON 1953. Pliocene Mollusca of southern Florida with special reference to those from North St. Petersburg ... with special chapters on Turridae by William G. Fargo and Vitrinellidae and fresh-water mollusks by Henry A. Pilsbry. Monogr. 8, Acad. Nat. Sci. Philadelphia: i- v+ 1-457; plts. 1-65 (6 November 1953) Ovsson, AxeEL ADOLF & THomas Lapuz McGinty 1958. Recent marine mollusks from the Caribbean coast of Panama with the descriptions of some new genera and species. Bull. Amer. Paleontol. 39 (177): 1-58; plts. 1-5 (10 December 1958) Orsicny, AtcipeE Cuaries Victor DESSALINES D’ 1842. in Sagra, Histoire physique, politique, et naturelle de Vile de Cuba. Atlas plts. 1 - 28. Bertrand, Paris. 1853. in Sagra, Histoire physique, politique, et naturelle de l’ile de Cuba. Spanish text. Bertrand, Paris. Pace, STEPHEN 1902. Contributions to the study of the Columbellidae. No. 1 Proc. Malacol. Soc. London 5 (1/2): 1-154 (23 April 1902) Petit DE LA SAUSSAYE, SAUVEUR 1853. Descriptions de coquilles nouvelles. 364; plt. 12, figs. 3, 4 PHILIPPI, RUDOLF AMANDUS 1846. Diagnosen neuer Conchylien Arten. April 1846: 49-55 Prrssry, Henry Auoustus # Herbert Nerson Lowe 1932. West Mexican and Central American mollusks collected by H. N. Lowe, 1929-31. Proc. Acad. Nat. Sci. Philadelphia 84: 33 - 144; pits. 1-17; 7 text figs.; 2 photogr. (21 May 1932) Powys, WiLLIAM LyTTLETON 1835. Undescribed shells contained in Mr Cuming’s collection ... Proc. Zool. Soc. London 3: 93 - 96 (25 September 1835) Rapwin, Georce EpwarpD 1968. New taxa of western Atlantic Columbellidae (Gastropoda, Pro- sobranchia). Proc. Biol. Soc. Wash. 81: 143-150 (30 April 1968) 1977. | The family Columbellidae in the western Atlantic. The Veliger 19 (4): 403-417; 1 plt.; 7 text figs. (1 April 1977) Ravene., EDMUND Naturhist. Tidsskr. 4: Journ. de Journ. de Conchyl. 4: (November 1853) Zeitschr. f. Malakoz. 1859. Descriptions of three new species of univalves, Recent and fossil. Proc. Elliott Soc. 1859: 280 - 282 1861. Descriptions of new Recent shells from the coast of South Caro- lina. Proc. Acad. Nat. Sci. Philadelphia 13: 41-42 (31 March) Regve, Lovett AucustTus 1846-1847. Conchologia Iconica vol. 3, Buccinum. plts. 1 - 14 (plts. 1- 12, December 1846; plts. 13, 14, February 1847) Reeve, London 1858-1859. Conchologia Iconica vol. 11, Columbella plts. 1-37 (plt. 1, April 1859; plts. 2-8, January 1858; plts. 9-12, April 1858; plts. 13 to 18, October 1858; plts. 19-23, November 1858; plts. 24-25, Feb- ruary 1859; plts. 26-37, April 1859) Reeve, London 1859. Conchologia Iconica, vol. 11, Meta. plt. 1 (May 1859) Reeve, London 1860. Conchologia Iconica. vol. 12, Terebra. plts. 1-27 (plt. 1, June, 1860; plts. 2-3, February 1860; plts. 4- 10, March 1860; plts. 11 - 15, May, 1860; plts. 16-27, June 1860) REHDER, HARALD ALFRED 1962. Contribucion al conocimiento de los moluscos marinos de archi- pelago de Los Roques y la QOrchila. Mem. Soc. Cien. Nat. La Salle 22 (62): 116-132 (May-August 1962) Risso, JOSEPH ANTOINE 1826. Histoire naturelle des principales productions de l’Europe méri- dionale et particulitrement de celles des environs de Nice et des Alpes maritimes. 4: i- viit 1 - 439; plts. 1-12 Sars, GEorcE OssIAN 1878. Bidrag til Kundskaben om Norges Arktiske fauna: I. Mollusca Regionis Arcticae Norvegiae pp. i-xii+ 1-466; 34 + xviii pits. Christiania Say, THOMAS 1822. Descriptions of marine shells recently discovered on the coast of the United States. Journ. Acad. Nat. Sci. Phila. 2: 207-221 (July 1822) Page 344 Scaccu!, ARCANGELO 1836. Catalogus conchyliorum regni Neapolitani quae usque adhuc reperit A. Scacchi. Napoli, pp. 1 - 18; plt. 1 SHERBORN, CHARLES Davies 1940. Where is the pp. 1-148 Sowersy, GzeorcE BRETTINGHAM Ist 1822-1824. The genera of Recent and fossil snails. Sowerby, London (no page numbers) (#9, Columbella, 30 September 1822) 1832. On new species of Columbella collected by Mr. H. Cuming. Proc. Comm. Sci. Corr. Zool. Soc. London 2: 113 - 120 (14 August 1832) collection? Cambridge, England, Sowersy, Georcz BRETTINGHAM 2nd 1844. Description of some new species of Columbella in the collection of H. Cuming, Esq. Proc. Zool. Soc. London 12: 48 - 53 (September 1844) StTzaRNS, Rospert Epwarps CARTER 1873. Descriptions of new marine shells from the west coast of Florida. Proc. Acad. Nat. Sci. Philadelphia 25: 344-347 (28 October 1873) Stimpson, WILLIAM 1851. Shells of New England. A revision of the synonymy of testaceous mollusks of New England ... pp. 1-56; plt. 1 StreEBEL, HERMANN 1905. Beitrage zur Kenntnis der Molluskenkunde der Magalhaen-Pro- vinz. Zool. Jahrb. 22 (6): 575-666; plts. 21-24 SWAINSON, WILLIAM 1840. A treatise on malacology; or the natural classification of shells and shellfish. London, Longman et al., i- viiit 1-419; 130 text figs. (May 1840) Tryon, Gzorcz WasHINGTON, Jr. 1883. Manual of conchology. vol. 5, Columbellidae. Tryon, Phila- delphia, pp. 100-276; plts. 42-63 (March-October 1883 — see Van- atta, 1927, The Nautilus 40 (3): 96-99) VERRILL, ADDISON EMERY 1882. Catalogue of marine Mollusca added to the fauna of New Eng- land during the past ten years. Trans. Conn. Acad. Arts Sci. 5 (2): 447 - 488 THE VELIGER Vol. 20; No. 4 Watson, Rosert Booc 1882. Mollusca of H. M.S. Challenger Expedition. Soc. London, Zoology 16 (93): 324 - 343 WEIsBORD, NorMAN EDWARD 1962. Late Cenozoic gastropods from northern Venezuela. Bull Amer. Paleo. 42 (193): 1-672; plts. 1-48 (5 March 1962) Wooprinc, WENDELL PHILLIPS 1928. Miocene Mollusca from Bowden, Jamaica, Part 2, gastropods and discussion of results. Carnegie Inst. Wash., 564 pp.; 40 plts. (November 1928) 1964. Geology and paleontology of the Canal Zone and adjoining parts of Panama. Description of Tertiary mollusks (Gastropoda: Columbelli- dae to Volutidae). U. S. Geol. Survey Prof. Paper 306C: 241 - 297; plts. 39 - 47 (27 February 1964) Journ. Linn. (12 June 1882) Epitor’s Note: This is the second time in 20 years of publishing The Veliger that an author died before proofs could be submitted to him. Responsibility for proof read- ing was shared in the present instance by Mr. Anthony D’Attilio, a former co-worker of Dr. Radwin, and an unnamed volunteer worker at the San Diego Museum of Natural History; Mrs. Jean M. Cate; Mrs. R. Stohler, and the editor. Any typographical errors remaining are the sole responsibility of the editor. Factual errors or errors of judgment and interpretation, if any, are the responsibility of the late Dr. George E. Radwin as is the credit for the work completed. Vol. 20; No. 4 THE VELIGER Page 345 Two New Giant Epitoniids (Mollusca : Gastropoda ) from West Africa PHILIPPE BOUCHET anv SIMON TILLIER Laboratoire de Biologie des .Invertébrés Marins et de Malacologie Muséum National d’Histoire Naturelle, 55, rue Buffon, 75005 Paris (1 Plate; 1 Text figure) IN 1971, Two sPEciEs of Epitoniids of large size were found washed ashore after a storm near Rufisque (Sene- gal). One of them is identical with a gastropod trawled in 1964 off Pointe-Noire, Congo, in 300m and accessioned by MNHIN in 1969. The second species has subsequently been found by various persons in West Africa. Surprising- ly enough, the 2 species, both longer than 50mm, appear to be undescribed. They are to be added to the list of spe- cies described in recent years from the coast of Senegal (BoucHET, 1975; Rosso, 1976, 1977; BoucHET & Nick- LES, 1976) and show how little we know about even the larger species inhabiting the Senegalese shelf. We shall first describe the species and then present a short synopsis of the family Eprronmpae in West Africa. Amaea africana Bouchet & Tillier, spec. nov. (Figures 7,256; 7; 8) Type Material: Holotype (shell only) in MNHN (Kerg- roach coll., Lozet leg.). Type Locality: Coastline near Rufisque, Senegal. Material Examined: One adult shell, Bay of Gorée, Séné- gal, MNHN (Marche-Marchad, coll.) ; one juvenile shell, Cavally, Ivory Coast, 50 - 55m, MNHN (Rancurel and Marchal, coll.) ; one adult shell from Senegal, without precise data, MNHN (Lozet leg.) ; 4 shells, off Grand Bassam, Ivory Coast, MNHN and IFAN (Le Loeuff, coll.) ; east of Sassandra, Ivory Coast, 56m, 1 shell; Grand Bereby, Ivory Coast, 56m, 1 live-taken specimen; south of Cavally River, Ivory Coast, 44m, 1 live-taken speci- men: all MNHN (Le Loeuff leg.) ; 1 adult shell, Dakar area, coll. M. Pin. Description: Holotype: shell slender, rather thin, frag- ile, composed of 13 whorls (the larval shell and about 1.5 postlarval whorls lacking) ; colour brownish orange with a narrow whitish subsutural band; columellar zone thick- ened, white; no umbilicus. The sculpture is cancellate, composed of axial and spiral lines. On the “first” (see supra) postlarval whorl, there is a single, sharp, spiral cord forming a sort of keel visible down to the 5" whorl. On the second whorl 2 spiral lines appear under the keel; at the beginning of the 4” whorl a 3" one appears under these 2, together with 2 other spiral lines above the keel. Along the 4" whorl, secondary spiral lines appear above and between the main lines. The number of lines increases in the following whorls: on the body-whorl a dozen main spiral lines can be counted, with 3 to 10 secondary ones between 2 adjacent main lines. The axial sculpture is formed by thin lamellae cross- ing over the spiral sculpture; these lamellae are straight and are bent only frontwards in the subsutural zone. On the body-whorl they show a tendency to merge together and we count 65 of them. The growth lines are hardly visible. The basal disc is lined by a rather strong spiral cord; below this, the numerous spiral threads are thin and more regular than above. Dimensions of the Shell: Height, 48mm, diameter, 18 mm. Other Specimens: The juvenile shell also lacks the apex but probably only the protoconch has fallen off; this shell has 1.5 more upper whorls than the holotype. The keel is present up to the top; the mode of formation of the spiral sculpture is the same as in the holotype. All other speci- mens correspond well with the holotype with slight vari- Page 346 ation in colour and number and breadth of the spiral threads of the last whorls. At a diameter similar to that of the body-whorl of the holotype, the axial lamella-num- ber varies from 50 to about 80. The longest shell (M. Pin, coll.) is 55mm high, with a diameter of 21 mmand 11.5 whorls (first whorls missing). Radula: 53 X (46-50) -0:0-0: (46-50): operculum illustrated in Figure 7. Figure 8; ar Ly, eves nA | o \ { j i / (eal SSO EY ( (( | i J WW) GN \ ‘ 5° 46 19 17 15 12 I Figure 8 Amaea africana Bouchet « Tillier, spec. nov. Halfrow of the radula of a specimen from Grand Bereby, Ivory Coast, 56m Remarks: There is no doubt that this is the species fig- ured by CaricaTi (1975: 235; plt. 4, figs. 1 - 2) as Amaea cf. mitchelli (Dall) ; Caricati mentioned West Africa only as the probable origin. Indeed, A. africana is not closely similar to any other West African species and is closer to A. mitchelli Dall, 1896 than to any other. We have com- pared A. africana with the plates of CLencH « TURNER (1950: plt. 106, figs. 5-7), ANDREWS (1971: 84), and with actual specimens of A. mitchelli from the coast of Texas, including the type (USNM 465611). The Ameri- can species is more solid and has a coarser sculpture; it is white with a median brown band and with a similar band below the basal keel. Amaea africana also comes near A. brunneopicta (Dall, 1896) from the Panamic Province, but the latter is more slender and thinner. We do not know any fossil from Europe to which it can THE VELIGER Vol. 20; No. 4 be compared, but a few American fossils could be con- sidered ancestors of the africana-mitchelli-brunneopicta group of species: — Epitonium eleutherium Pilsbry & Olsson, 1941, from the Pliocene of Ecuador is very close to Amaea brunneo- picta; — Ferminoscala pseudoleroyi (Maury, 1925), from the Miocene of Jamaica, as Tluswatcd by Wooprinc (1928: 402; plt. 32, figs. 3-4; ?== Scalina gardnerae Olsson, 1967: Miocene of Florida) differs by its slenderness and stronger spiral sculpture. Amaea guineense Bouchet & Tillier, spec. nov. (Figures 3, 4, 5) Type Material: Holotype and one paratype (shells only); in MNHN (Kergroach coll., Lozet leg.). Type Locality: Coastline near Rufisque, Senegal. Material Examined: One shell, off Pointe-Noire, Con- go, 300m (Crosnier, Orstom leg.). Description: Holotype: shell slender, solid, of a uniform creamy colour. There are 8 whorls, but the protoconch and several postlarval whorls have been broken off at a diameter of 5mm. The whorls are twice as broad as high and the suture is deep. The columella is remarkably thick- ened for an Amaea. There is no umbilicus. The sculpture is cancellate, composed of axial and spiral lines. The spiral sculpture is formed by 9 main threads, visible from the first postlarval whorl still pres- ent. From the 3™ preserved whorl, secondary threads ap- pear between them: their number (1 to 3 between 2 adjacent main spiral threads) and importance are not constant. In the subsutural zone, there are 4 to 6 second- ary spiral threads; they are oblique and cannot be fol- lowed from one side of any axial lamella to the other. The axial lamellae are developed to roughly the same degree as the main spiral threads and they deviate a little back- wards when they cross over the latter. Near the upper Explanation of Figures 1 to 7 Amaea africana Bouchet « Tillier, spec. nov. Figure 1: Holotype, ventral view; length 48mm; diameter 18mm Figure 2: Juvenile specimen, Cavally, Ivory Coast; ventral view; length 1omm; diameter 4mm Figure 6: Holotype, view of the basal disc; diameter 18mm Figure 7: Operculum of a specimen: from south of Cavally River, Ivory Coast, 44m Amaea guineense Bouchet «& Tillier, spec. nov. Figure 3: Paratype, ventral view; length 39.5mm; diameter 15.5 mm Figure 4: Holotype, ventral view; length 64mm; diameter 26mm Figure 5: Paratype, view of the basal disc; diameter 15.5 mm (Photographs by A. Foubert) Tue Ve icrr, Vol. 20, No. 4 [BoucHeT & TiLieR] Figures 1 to 7 = Vol. 20; No. 4 suture they bend frontwards and show a winglike projec- tion; the lamellae of 2 consecutive whorls do not join. As we approach the peristome the lamellae grow thicker and several of them may join to form varices: this is why the outer lip looks so thick. Because of these varices, the number of lamellae on the body-whorl is from 61 along the basal keel to 50 in the subsutural zone. From the 3! whorl onwards, thin growth lines can be seen between the lamellae: there is a clear intersection with the second- ary spiral threads, not with the main ones. The basal disc is sculptured by many spiral threads, axial lamellae and growth lines. Dimensions of the Shell: Height, 64mm; diameter, 26 mm. Other Specimens: The specimen from Congo is a 61 mm broken shell, on which a sea-anemone has grown. From the holotype, it differs mainly by showing a greater tendency to form varices so that only 30 axial lamellae can be counted on the body-whorl. The paratype is a younger shell with the protoconch and at most 1 postlarval whorl broken. The 9 main spiral threads are present at the very top; secondary ones appear in the subsutural zone on the 4 whorl and between the main threads on the 5" whorl. The upper whorls are regularly convex in outline. Remarks: We do not know any Atlantic species close to Amaea guineense; only the Panamic species A. tehuan- arum DuShane « McLean, 1968 is similar but it has keeled upper whorls and as a result has a different pattern in sequence of appearance of spiral lines. Apparently, there is no fossil in the American Ceno- zoic, but several in Europe which might be considered an- cestors : ~ Amaea subreticula (d’Orbigny, 1852), from the middle and upper Miocene, does not show any growth lines be- tween axial lamellae (after the description of GLIBERT, 1952: 44); it has lower whorls and the main spiral threads are fewer. — Amaea pellati (de Raincourt « Munier-Chalmas, 1863), from the Biarritz Oligocene (= Acrilla amoena var. eo- subcancellata Sacco: see Boussac, 1911: 83) is closer to A. subreticula than to A. guineense. — Amaea phoenix (de Boury, 1912) from the lower Mio- cene of Dax and Corsica has a rather similar sculpture, but is more slender and has a more oblique suture. — Amaea elegantissima (Deshayes, 1861) from the Grig- non Lutetian is also close to A. phoenix and A. guineense, but can be separated by a sculpture with fewer axial lamellae. Amaea guineense then appears to be the living form of a lineage starting in the Paris Eocene, through the Mio- cene of southern France to present-day Gulf of Guinea. THE VELIGER Page 347 Unfortunately, we have too little information on the up- per Cenozoic of northwestern Africa to be able to identify additional elements of this lineage. The family EprronnpaeE in West Africa Over 20 Epitoniids have already been mentioned from tropical West Africa; however, many are known only from the original description and we have no idea on the dis- tribution of most species. Also their biology is virtually unknown as far as feeding and larval development are concerned. To be added to the species already described, we have seen several undescribed ones, especially in the de Boury Collection and among recently collected mate- rial: when the family is better known, there will probably be more than 40 species from the shelf of tropical West Africa. It should also be remembered that the fauna of the slope is almost completely unknown. LIST OF THE WEST AFRICAN SPECIES Opalia crenata (Linnaeus, 1758): a Mediterranean spe- cies known from Mossamédés (DAUTZENBERG, 1913) and Sao Thomé (ToMLIN & SHACKLEFORD, 1914) Opalia gaini (deBoury in Lamy, 1923) from the Bissagos archipelago and Gaboon; Opalia hellenica (Forbes, 1844) Epitonium pulchellum (Bivona, 1832) Epitonium grossicostatum (Nyst, 1873) Epitonium candidissimum (Monterosato, 1877) Amaea smithi (Watson, 1897 non Tryon, 1887) — all mentioned from Sao Thomé by ToMLIn & SHACKLE- FORD (1914) together with Epitonium commutatum (Monterosato, 1877), occurring from the Mediterranean to Angola (Lamy, 1907; 1908; NosreE, 1909; Hmatco, 1910; DAUTZENBERG, 1913). ‘Two manuscript names, credited to de Boury, have been published by TomuLIN & SHACKLEFORD (1914): Epitoni- um atlanticum and E. tenuipunctatum. These specimens are present in the de Boury collection but are apparently nomina nuda. SmiTH (1871) has described 2 new species from Why- dah (or Ouidah), Dahomey: Scala miranda Smith, 1871, changed to Scalaria smithi by Tron (1887) because he considered it to be preoc- cupied by Crossea miranda A. Adams; Scala bairdi Smith, 1871. In 1890, the same author mentioned 7 more species from Saint Helena, 5 of them being new: Scalaria confusa Smith, 1890 Scalaria mellisii Smith, 1890 Scalaria sanctaehelenae Smith, 1890 Scalaria atomus Smith, 1890 Scalaria commoda Smith, 1890 Scalaria fragilis Hanley, 1842 Page 348 Scalaria cf. multistriata Say, 1825, the 2 last mentioned being already known from the West Indies. Other species: Scalaria pachygyra Locard, 1896, dredged by the “Tra- vailleur” in the Cape Verde Islands in 100 - 318m; Scalaria cochlea Sowerby, 1844 from the coasts of Angola (SowERBY, 1847; DUNKER, 1853); Scalaria fusca Sowerby, 1844, from Sierra Leone (SowER- By, 1847) ; Epitonium tenuicostatum von Martens, 1882, dredged by the “Gazelle” in Cape Verde Islands. Finally, the following 3 species have been described from Gorée, Senegal, but never illustrated, by von Mattzan, (1885). Scalaria boettgeri von Maltzan, 1885 Scalaria trochiformis von Maltzan, 1885 Scalaria senegalensis von Maltzan, 1885. ACKNOWLEDGMENTS We would like to thank Mr. J. B. Lozet for the interesting material put at our disposal, and Dr. P. Le Loeuff and Mr. Pin for additional specimens. We also thank Dr. Joseph Rosewater for the loan of Dall’s type and other shells of Amaea mitchelli. Literature Cited ANDREWS, JEAN 1971. Sea shells of the Texas coast. xvii+298 pp.; illust. Boucuet, PHILIPPE 1975. Nudibranches nouveaux des cétes du Sénégal. 25 (1A): 119-132; 1 plt. Boucuert, Puitippe & Maurice NICKLES 1976. Mactra diolensis, bivalve nouveau du Parc national de basse Univ. Texas Press, Austin, Vie et Milieu Casamance. Bull. IFAN 38(1A): 57-61 Boury, EDGAR DE 1912. Description de Scalidae nouveaux ou peu connus. Journ. de Conchyl. 60: 267 - 322; plts. 7-8 Boussac, JEAN 1911. Etudes stratigraphiques et paléontologiques sur le Nummulitique de Biarritz. Ann. Hébert, Paris 5: 1-95; plts. 1-24 Caricati, AUGUSTO 1973. Reperti inconsueti dalle coste occidentali africane. lie 9 (11-12): 235 - 238; plt. 4 CrencH, WiLiIAM James & RutH Dixon TurNER 1950b. The genera Sthenorytis, Cirsotrema, Acirsa, Opalia and Amaea in the western Atlantic. Johnsonia 2 (29): 221-248; plts. 96 to Conchig- 107 (30 September 1950) Dai, WiLit1Am HEALEY 1896. On some new species of Scala. The Nautilus 9 (10): 111-112 DAUTZENBERG, PHILIPPE 1913. Mission Gruvel sur la cote occidentale d’Afrique (1909-1910): Mollusques marins. Ann. Inst. Océanogr. 5 (3): 1-111; plts. 1-3 DesuHayes, GERARD PAUL 1860-1864 (livraisons) Description des animaux sans vertébres découverts dans le bassin de Paris. Vol. 2, Mollusques Acéphalés Monomyaires et Brachiopodes — Mollusques Céphalés, premiére partie. Paris (Baillére): pp. 1 - 968; atlas plts. 1 - 107 DunKER, WILHELM 1853. Index molluscorum quae in itinere ad Guineam inferiorem col- legit Georgius Tam Med. Dr. Cassell (T. Fischer): 1 - 74; plts. 1 - 10 (3 February 1896) THE VELIGER Vol. 20; No. 4 DuSuHane, HELEN 1974. The Panamic-Galapagan Epitoniidae. plement): 1-84; 15 plts.; 5 text figs.; 1 map DuSwanz, HELEN & JAaMEs Hamitton McLean 1968. Three new epitoniid gastropods from the Panamic province. Contrib. Sci. Los Angeles County Mus. Nat. Hist. 145: 1-6; 6 figs. (14 June 1968) The Veliger 16 (Sup- (31 May 1974) G.LiBERT, MAxIME 1952. | Faune malacologique du miocéne de la Belgique, II, Gastéro- podes. Mém. Inst. Roy. Belg. Sci. nat. 121: 1 - 197; plts. 1- 10 Hmatco, Joaquim 1910. Moluscos de la Guinea espafnola. nat. 1 (29): 507-524 Lamy, EpouarD 1907. Liste des coquilles marines recueillies par M. Ch. Gravier 4 Vile San Thome (1906). Bull. Mus. Natn. Hist. nat. 13: 145 - 154 1908. Liste des coquilles recueillies par M. A. Chevalier sur la céte occidentale africaine (1900-1907). Bull. Mus. Natn. Hist. nat. 14: 285 - 289 1923. Mission du Comte Jean de Polignac et de M. Louis Gain (cam- pagne du Sylvana 1913): Mollusques testacés. | Compt. Rend. Congr. Soc. Sav. 1922, sciences: 24 - 37 Locarp, ARNOULD 1896. Expéditions scientifiques du Travailleur et du Talisman: Mol- lusques testacés. 1. Paris (Masson): 1-516; plts. 1 - 22 MatLtTzan, HERMANN VON 1885. Neue Gastropoden vom Senegal. Gesellsch. 17: 25 - 30 Martens, Epuarp Cari von 1882. Vorzeigung zweier neuer Arten von Meer-Conchylien von der Expedition S. M. Sch. Gazelle. Sitz. Ber. Gesellsch. Naturf. Freunde, Berlin 1882: 107 Nosre, AuGusto 1909. Matériaux pour I’étude de la faune malacologique des possessions portugaises de l’Afrique occidentale. Bull. Soc. Port. Sci. Nat. 3 (suppl. 2): 1-108 Oxsson, AxEL ADOLF 1967. Some Tertiary molluscs from South Florida and the Caribbean. Paleont. Res. Inst. Ithaca, New York. pp. 1 - 61; 9 plts. (10 March 1967) Pirspry, Henry Aucustus & AxeLt ADoLF OLSSON 1941. A Pliocene fauna from Western Ecuador. Sci. Philadelphia 93: 1 - 79; plts. 1-19 RarncourT, JEAN BAPTISTE CHARLES PROSPER DE & ERNEST PHILIPPE AucusTEe MuNIER-CHALMAS 1863. Description d’un nouveau genre et de nouvelles espéces fossiles du bassin de Paris et de Biarritz. Journ. de Conchyl. 11: 194 - 203; Mem. R. Soc. Esp. Hist. Nachr. Deutsch. Malakol. Proc. Acad. Nat. pits. 7-8 Rosso, JEAN-CLAUDE 1976. Psammotreta (Florimetis) elouardi, nov. spec. des cotes du Séné- gal. Conchiglie 12 (1-2): 57-60 1977. Un Volutocorbis des ctes du Sénégal: V. nicklest sp. nov. (Gast- ropoda Prosobranchia, Volutidae). Bull. IFAN 38 A (1): 62 - 68; 1 text fig. SmitH, Epcar ALBERT 1871. A list of species of shells from west Africa, with descriptions of those hitherto undescribed. Proc. Zool. Soc. London 1871: 727 - 739; pit. 75 1890. Report on the marine molluscan fauna of the Island of St. Helena. Proc. Zool. Soc. London for 1890: 247 - 317; plts. 21 - 24 (August 1890) Sowersy, GzorczE BretrincHamM 2nd 1844, Descriptions of new species of Scalaria, collected by Hugh Cum- ing, Esq. Proc. Zool. Soc. London prt. 12: 1 - 38 (July 1844) [1842] 1847 - 1848. Thesaurus conchyliorum, or monograph of genera of shells, edited by G. B. Sowerby, Jr., completed by G. B. Sowerby 3rd. London, vol. 1, prt. 4: 83-146; plts. 32-40 (11 April 1844) Tomuin, Joun Reap Le BrockTon & Lewis J. SHACKLEFORD P 1914. The marine Mollusca of Sao Thomé, 1. Journ. Conchol. 14: 239 - 256 Tryon, Gzorce WASHINGTON, Jr. 1887. Manual of Conchology, vol. 9: 1 - 488; plts. 1-71. Philadelphia Watson, Rosert Booa 1897. On the marine Mollusca of Madeira; with description of thirty- five new species, an index-list of all the known sea-dwelling species of that island. Journ. Linn. Soc. London 26: 233 - 329; 2 plts. Wooprinc, WENDELL PHILLIPS 1928. Miocene mollusks from Bowden, Jamaica: Part II: Gastropoda and discussion of results. Carnegie Inst. Wash. publ. 385: 1 - 564; pits. 1-40; 3 text figs. (28 November 1928) Vol. 20; No. 4 THE VELIGER Page 349 Clonal Variation in the Parthenogenetic Snail Campeloma decisa (Viviparidae) ROBERT K. SELANDER, E. DAVIS PARKER, Jr. anD ROBERT A. BROWNE Center for Evolution and Paleobiology and Department of Biology, University of Rochester Rochester, New York 14627 and Department of Biology, Syracuse University, Syracuse, New York 13210 INTRODUCTION RECENT sTupiEs OF clonal diversity in parthenogenetic animals, as revealed by electrophoretic analysis of allo- zymic variation, have contributed to an understanding of processes involved in the evolution of non-Mendelian genetic systems (review in Lox, 1976; PARKER & SE- LANDER, 1976; PARKER et al., 1977). We here report on clonal diversity and genic heterozygosity in several pop- ulations of the parthenogenetic aquatic snail Campeloma decisa (Say, 1822) in central New York. MATERIALS anp METHODS Samples of Campeloma were collected by hand or by baiting with meat (ALLISON, 1942) at 5 localities in New York: Song Lake,near Syracuse, Onondaga County ; west side of Jamesville Reservoir at Craner Oil Depot, Jamesville, Onondaga County; north end of Honeoye Lake, Livingston County; Delaware River at Walton, Delaware County; and Dryden Lake, near spillway east of Dryden, Tompkins County. Individual snails were processed for electrophoresis according to the techniques of SELANDER & Hupson (1976). Allozymic variation at 13 enzyme systems encoded by 21 structural gene loci was assayed by horizontal starch-gel electrophoresis (tech- niques described by SELANDER et al., 1971). The follow- ing enzyme loci were scored: 3 leucyl-alanine peptidases (Pep—1, —2, —3), 2 leucine aminopeptidases (Lap—r, -2), 2 esterases (Est—1,—2), 2 phosphoglucomutases (Pgm-r,-2), phosphoglucose isomerase (Pg:), mannose phosphate iso- merase (M pi), B-glucuronidase (8-Glu), 2 superoxide dis- mutases (Sod —1, -2), a-glycerophosphate dehydrogenase (a-Gpd), glucose-6-phosphate dehydrogenase (G6pd), 2 malate dehydrogenases (Mdh-1, -2), 2 isocitrate dehydro- genases (Idh—-1, -2), and 6-phosphogluconate dehydro- genase (6-Pgd). Electromorphs (corresponding to allelic variants) at variable loci were numbered according to their relative mobility, the most common being designated 100 in all cases. Average individual heterozygosity (H) was determined by direct count of heterozgotes. RESULTS anp DISCUSSION Nine of the 21 loci assayed were polymorphic in the 5 samples of Campeloma examined (Table 1). Two clones Table 1 Genetic diversity in clones of Campeloma decisa Genotype Locus! Clone I? Clone II? Pep-2 100/100 100/95 Lap-1 100/100 100/90 Lap-2 null 100/100 Est-1 100/100 100/90 Pgm-1 110/100 100/100 Pgm-2 100/100 110/100 Sod-1 100/100 105/100 a-Gpd 100/100 100/90 Idh-1 100/90 110/100 1The following loci were monomorphic for the same allele in both clones: Pep-1, Pep-3, Est-2, Pgi, Mpi, B-Glu, Sod-2, G6pd, Mdh-1, Mdh-2, Idh-2, and 6-Pe¢d. 2Clone I: Song Lake, N = 30 specimens; Walton, N = 11; Dryden Lake, N = 38. 3Clone II: Jamesville Reservoir, N = 30; Honeoye Lake, N = 1. Page 350 (designated I and II) can be distinguished, and these differ at each of the 9 variable loci. The nature of the difference is such that only at one locus (Lap—2) do the clones not share an allele. The clones are monomorphic and indistinguishable at the remaining 12 loci. Thus, the clones differ genotypically at 43% of the loci examined, but share alleles at 95% of the loci. The clones differ in level of individual heterozygosity, with Clone I having 2 of 21 loci fixed in heterozygous con- dition (H =9.5%), and Clone II having 7 of 21 loci fixed as heterozygotes (H 33.3%). The occurrence of fixed heterozygosity in one or both clones at 9 loci strongly sug- gests that Campeloma decisa has an apomictic system of egg maturation in which recombination does not occur (Uz- ZELL, 1970; Nur, 1971), since, in the absence of heterotic selection, recombination in parthenogenetic lineages should lead to homozygosity (ASHER, 1970). MatTox’s (1937) cytological work on the related parthenogenetic form C. rufum was interpreted by SUOMALAINEN (1950) as evidence that odgenesis is apomictic. Without knowledge of the nature and extent of genetic diversity in the bisexual populations of Campeloma from which the parthenogens were derived, we can only specu- late on the mode of origin of the clones. Because hetero- zygosity (and clonal diversity) is expected to increase in apomictic parthenogenetic lineages as a consequence of the accumulation of mutations (WurITE, 1973; LoKKI, 1976), it might be suggested that Clone II is older than Clone I. However, an alternative hypothesis is that the clones are equivalent in age but were derived from sexual individuals (belonging to the same or different popula- tions) differing markedly in heterozygosity. A third pos- sibility, which we regard as the least likely, is that the clones diverged genetically following the origin of a single parthenogenetic lineage from an individual of an ancestral sexual population. Parthenogenetic forms of Campeloma apparently are confined to northeastern and north-central North Ameri- ca, while sexual species occur south of a line from Ken- tucky to Illinois (Matrox, 1938; PoLLISTER & POLLISTER, 1940; Husricut, 1943; VAN DER SCHALIE, 1965; AN- DERSON, 1966). This pattern of distribution suggests the possibility that the parthenogens evolved and colonized northern areas at the time of recession of the last Pleisto- cene glacier. A similar interpretation has been advanced by SUOMALAINEN (1962) to account for patterns of dis- tribution of parthenogenetic insects in Europe. Our findings for Campeloma are consistent with the results of previous work demonstrating that parthenoge- netic “species” generally are clonally diverse (SuomMAL- AINEN & SaurRA, 1973; Loxxi et al., 1975; PARKER & SELANDER, 1976; PARKER et al., 1977). Each of the 4 pop- THE VELIGER Vol. 20; No. 4 ulations for which we have reasonably adequate samples apparently consists of individuals belonging to one clone. Because populations of Campeloma are widely spaced and isolated in central New York, migration probably is in- frequent, and colonization normally may involve the trans- port of only one or a few individuals. Hence, the uniclonal composition of populations may merely reflect the founder effect. However, the possibility that interclonal competi- tion is involved should also be considered. For a number of parthenogenetic “species” and strongly selfing species, stud- ies already reported or currently in progress in our labora- tory suggest that local areas are inhabited by a small num- ber of clones or strains that are genetically very divergent and differ in habitat distribution. For example, populations of the self-fertilizing land snail Rumina decollata in south- ern France are composed of 2 strains differing at 50% of their loci and showing “preferences” for relatively xeric or humid microhabitats (SELANDER & HupsoNn, 1976). Clones of the parthenogentic aquatic snail Potamopyrgus jenkinst (WaRWICK, 1952; WINTERBOURN, 1970) differ at about half their loci (SELANDER & JONES, in prepara- tion). Similarly, many populations of the parthenogenet- ic earthworm Octolasion tyrtaeum in central New York consist of 2 clones differing at 40% of their loci (JouNn JAENIKE, in preparation). This pattern suggests that lim- iting similarity (MacArRTHUR & LEvINS, 1967; May & MacArTHDvR, 1972) is involved in determining the extent of interclonal and interstrain diversity in local populations of parthenogenetic and selfing organisms. Our findings for Campeloma are consistent with the genetic aspect of this hypothesis, but the number of populations sampled is too small to provide evidence regarding possible clonal differ- ences in habitat utilization. The hypothesis that clones or selfing strains can coexist only if they are rather divergent in genetically-determined adaptive traits related to dif- ferential niche utilization (or if they are minimally distinct genetically and, hence, ecologically equivalent or nearly so) is, of course, simply an extension of current theories relating to interspecific competition and resource par- titioning. It can be tested through intensive study of the genetic and ecological relationships of parthenogenetic organisms such as Campeloma. SUMMARY Populations of the parthenogenetic aquatic snail Campe- loma decisa in central New York consist of one of two clones differing genotypically at 43% of their structural gene loci and sharing alleles at 95% of the loci. The occur- rence of fixed heterozygosity at 9 loci in one or both clones suggests a non-recombinational system of egg ma- Vol. 20; No. 4 THE VELIGER Page 351 turation. Individual heterozygosity was 9.5% in one clone and 33.3% in the other. It is suggested that both the founder effect and ecological limiting similarity potenti- ally are important factors determining the clonal struc- ture of parthenogenetic “species.” ACKNOWLEDGMENTS Research supported by NSF grant BMS 75-18891, NIH grant Ro1-GM-22126, and the Theodore Roosevelt Fund of the American Museum of Natural History. E. D. Parker is a Predoctoral Trainee on NIH grant T32-GM-41112. Literature Cited At.ison, L. N. 1942. Trapping snails of the genus Campeloma. Science 95: 131 to 132 ANDERSON, B. E. 1966. Studies on the molluscan genus Campeloma Rafinesque, 1819. Sterkiana 23: 9-18 AsuHer, J. H. 1970. Parthenogenesis and genetic variability. II. One-locus models for various diploid populations. Genetics 66: 369 - 391 Husricnt, Les.iz 1943. Notes on the sex ratios in Campeloma. 138 - 139 Loxx1, J. 1976. Genetic polymorphism and evolution in parthenogenetic animals. VII. The amount of heterozygosity in diploid populations. Hered- itas 83: 57 - 64 Loxxl, J., E. SUOMALAINEN, A. Saura & P. LANKINEN 1975. Genetic polymorphism and evolution in parthenogenetic animals. II. Diploid and polyploid Solenobia triquetrella (Lepidoptera: Psychi- dae). Genetics 79: 513 - 525 MacArtuHur, Rosert H. # RicHarp Levins 1967. The limiting similarity, convergence, and divergence of coexist- ing species. Amer. Natural. 101: 377 - 385 The Nautilus 56 (4) : Mattox, Norman T. 1937. Oogenesis of Campeloma rufum, a parthenogenetic snail. Zeitschr. Zellf. Mikrosk. Anat. 27: 455 - 464 1938. Morphology of Campeloma rufum, a parthenogenetic snail. Journ. Morphol. 62: 243 - 261 May, R. & R. H. MacArtHurR 1972. Niche overlap as a function of environmental variability. Proc. Nat. Acad. Sci. USA 69: 1109 - 1113 Nor, U. 1971. Parthenogenesis in coccids (Homoptera). Amer. Zool. 11: 301 - 308 Parker, E. D., Jr. & R. K. SELANDER 1976. The organization of genetic diversity in the parthenogenetic lizard Cnemidophorus tesselatus. Genetics 84: 791 - 805 Parker, E. D., Jr.. R. K. Seranver, R. O. Hupson « L. J. Lester 1977. Genetic diversity in colonizing parthenogenetic cockroaches. Evolution, in press Poxuister, A. W. & P F. Po.iistTer 1940. Distribution of males in the genus Campeloma, with a note on the chromosome numbers in Viviparidae. Anat. Res. 78: 128 (abst.) SELANDER, Rosert K. « R. O. Hupson 1976. Animal population structure under close inbreeding: The land snail Rumina in southern France. Amer. Natural. 110: 695 - 718 Sevanper, R. K., M. H. Situ, S. Y. Yana, W. E. JoHNson « J. B. Gentry 1971. Biochemical polymorphism and systematics in the genus Pero- myscus. I. Variation in the old-field mouse (Peromyscus polionotus). Stud. Genet. VI, Univ. Texas Publ. 7103: 49 - 90 SUOMALAINEN, E. 1950. Parthenogenesis in animals. Adv. Genetics 3: 193 - 253 1962. The significance of parthenogenesis in the evolution of insects. Ann. Rev. Entomol. 7: 349 - 366 SUOMALAINEN, E. & A. SAURA 1973. Genetic polymorphism and evolution in parthenogenetic animals. I. Polyploid Curculionidae. Genetics 74: 489 - 508 UzzzLL, T. 1970. Meiotic mechanisms of naturally occurring unisexual verte- brates Amer. Natural. 104: 433 - 445 VAN DER SCHALIE, HENRY 1965. Observations on the sex of Campeloma (Gastropoda: Vivipari- dae). Occas. Pap. Mus. Zool., Univ. Michigan No. 641: 1-15 Warwick, T. 1952. Strains in the mollusc Potamopyrgus jenkinst (Smith). Na- ture 169: 551-552 Wuite, M. J. D. 1973. Animai cytology and evolution. 3™ ed. Cambridge Univ. Press, Cambridge, England WINTERBOURN, M. 1970. | The New Zealand species of Potamopyrgus (Gastropoda: Hydro- biidae). Malacologia 10: 283 - 321 (10 July 1971) Page 352 THE VELIGER Vol. 20; No. 4 Effects of Cornstarch and Dextrose on Oysters ' BY KENNETH W. TURGEON anp DEXTER S. HAVEN Virginia Institute of Marine Science, Gloucester Point, Virginia 23062 INTRODUCTION THE GROWTH AND FATTENING of the American oyster, Crassostrea virginica (Gmelin, 1791), has long been of interest to oyster biologists since the market value of oys- ters is directly related to the quantity and quality of “meat” obtained per bushel (== 35.2 L) of harvested oys- ters. Much of the early work done on the growth and fattening of oysters attempted to demonstrate the superior nutritional value of specific planktonic organisms and detritus (Moore & PopE, 1910; MARTIN, 1923, 19274, 1927b, 1928; Gavarp, 1927; NELSON, 1947). MITCHELL (1917) was the first to study the effects of carbohydrates of known composition on oysters. He reported that oysters held in standing seawater containing 0.25% glucose had higher glycogen levels than control oysters. YONGE (1928) showed that oysters were capable of removing dissolved carbohydrates from seawater. GILLESPIE, INGLE & Ha- VENS (1964) demonstrated that oysters receiving only dextrose at 30mg/L lived an average of 68.2 days longer than starved oysters. NELSON (1934) was the first to investigate effects of particulate carbohydrates on oysters. Although his report did not present quantitative data, he stated that only cornstarch was “successful.” More recent investigators have studied the effects of particulate and dissolved carbo- hydrates on the glycogen content, tissue weight and shell size of oysters. HAVEN (1965) demonstrated that oysters receiving wheatflour or cornstarch at concentrations of 2mg/L would, at certain seasons, have dry tissue weights significantly greater than controls receiving only flows of natural river waters. His results of supplemental feeding with dextrose were not definitive, and dextrose concentra- tions as high as 34 mg/L were required to produce a sig- nificant influence on dry tissue weight. These studies were corroborated by Gm.espiE, INGLE & Havens (1966). Kuwatari & Nisuu (1967) showed that rice powder ' Contribution No. 828 from the Virginia Institute of Marine Science, Gloucester Point, Virginia 23062 added to the diet of the pearl oyster resulted in a higher tissue weight. In a more recent study, CASTELL & TRDER (1974) demonstrated the nutritional value of carbohy- drates, lipids and other dietary substances of known com- position in the diet of oysters. Previous authors have suggested that stored glycogen is a major factor influencing the size and quality of oyster tissue and have shown that the level of stored glycogen on a dry weight basis varies seasonally with a minimum of about 3% in late summer and a maximum of about 24% in early spring and late fall (ENcGLE, 1950; Hopxins, Mackin & MENZEL, 1953). As a consequence of this seasonal change in glycogen, effects of supplements added at various seasons might vary. There were three purposes to this study. The first was to define the minimum quantity of particulate carbohy- drate (in the form of partially hydrolyzed cornstarch) necessary to produce a measurable increase in glycogen content, tissue weight, shell height, underwater shell weight and total volume of oysters. The second was to determine if the effect of starch varied with season. The third was to eliminate some of the confusion concerning the uptake and utilization of dextrose. That is, does sup- plemental feeding with low levels of dextrose result in statistically significant increases in glycogen content, tis- sue weight, shell height, underwater shell weight and total volume. Also, are these increases, if they occur, statisti- cally comparable to those produced by cornstarch. In order to answer these questions oysters were offered dex- trose in two forms: 1) asa solution; and 2) mixed with clay. The rationale behind mixing dextrose with clay was that soluble sugars are known to be adsorbed on clay particles similar to those present in marine waters (BapErR, 1962). It was theorized, therefore, that if oysters were offered clay particles coated with dextrose they would ingest the coated particle and the dextrose would be stripped from the particle and assimilated. The diges- tive diverticula of oysters are known to be quite acidic (GALTSOFF, 1964), a situation which would favor this process. Vol. 20; No. 4 THE VELIGER Page 353 MATERIALS anp METHODS Four experiments were conducted during the summer and fall of 1966 and the spring of 1967 using different concen- trations of cornstarch, dextrose and clay. The types of supplements added, size and treatment of experimental groups, sources of oysters, size, meat measurements and statistical treatment of data are presented in this section and in Table 1. Oysters for all studies were obtained from Horsehead Bar in the James River, Virginia, an area free of known oyster diseases (Andrews, personal communication). Ex- perimental oysters were selected for uniformity (40 to. 45 mm in shell height), freed of attached fouling, randomly separated into groups of 20 and numbered with an enamel paint. Each group of 20 oysters became a “unit” which was subject to various experimental conditions. One day prior to the start of an experiment, the follow- ing measurements were taken on individual oysters in all groups: 1) underwater shell weight to the nearest 0.01 g by the method of ANDREWS (1961); 2) shell height to the nearest 0.1 mm with vernier calipers; and 3) total volume to the nearest 0.1cc by water displacement. In addition, one group of 20 oysters was sacrificed on the first day of each experiment to determine the following meas- urements: 1) wet tissue weights to the nearest 0.01; and 2) glycogen levels estimated to the nearest 0.01% of the wet tissue weight. Wet tissue weights were recorded after the tissues had drained for 1 minute on a fine mesh, metal grid. Glycogen was extracted by the method out- lined by CALDERWOOD & ARMSTRONG (1941) with the ex- ception that the precipitated glycogen, rather than being collected on filter paper, was centrifuged down and the supernatant discarded (Armstrong, unpublished). Glyco- Table 1 Specific Details of Experiments I, II, III and IV Showing Dates, Water Temperatures, Salinities and Treatments. Temp. Range Salinity Range Exp. and and No. Date (mean) C (mean) % Treatment I 7/ 7/66-13/ 8/66 27.9-29.4 20.4-22.4 1. Cornst.—2 mg/L (25.6) (21.6) 2. Dext.—2 mg/L 3. Mont.—2 mg/L Max. 4. Dext. & Mont.—2 mg/L each Summer 5. Control —no supp. Temp. 6. Initial —sacrifice II 27/ 7/66- 11/10/66 27.2-17.2 21.3-24.2 1. Const.—2 mg/L (22.1) (23.1) 2. Const.—5 mg/L 3. Dext.—2 mg/L Decreasing 4. Mont.—2 mg/L Fall 5. Dext. & Mont.—2 mg/L each Temp. 6. Control —no supp. 7. Initial —sacrifice III 21/10/66- 6/12/66 18.2- 5.6 22.0-23.3 1. Cornst.—2 mg/L (12.6) (20.9) 2. Cornst.—5 mg/L 3. Dext.—2 mg/L To 4. Control—no supp. Lowest 5. Held in tray, York Rv. Fall Temp. 6. Initial —sacrifice IV 1/ 4/67-29/ 6/67 12.8-25.6 20.7-17.9 1. Cornst.—0.25 mg/L (18.2) (19.1) 2. Cornst.—0.50 mg/L 3. Dext.—5 mg/L Increasing 4. Dext.—5 mg/L & Spring Mont.—2 mg/L Temp. 5. Control —no supp. 6. Held in tray, York Rv. ~ . Initial — sacrifice Page 354 THE VELIGER Vol. 20; No. 4 gen levels were determined colorimetrically by the method of Kemp «& Kits vAN HEIjNIGEN (1954). Analysis of Variance testing (SoKAL & RoHLF, 1969) showed that the initial mean underwater shell weights, shell heights and total volumes among the oyster groups in each experi- ment were statistically similar (7. e., differences among the oyster groups were non-significant at the 5% significance level). Consequently it was assumed that, for each experi- ment, the sacrificed and experimental groups had statis- tically similar initial mean wet tissue weights and glycogen levels. Oysters were held in Plexiglas troughs under running York River water. Troughs measured 36cm long by 19cm wide by 6cm high and consisted of 5 compartments (4 oysters per compartment) and a baffle to insure thorough mixing of water and supplement. Oysters were oriented with bills facing the current and the right valve (flat side) up. Position of oysters in respect to the inflow was changed daily on a random basis. Troughs and oysters were cleaned of feces and pseudofeces every other day. York River water was pumped to a large overhead trough in the laboratory, flowed into a conducting column of Plexiglas, through Tygon tubing to flow meters and then to the oyster-holding troughs through Tygon tubing. Water flow through the flow meters was checked twice a day and readjusted to the proper flow if necessary. In Experiments I, II and III, the water flow was 1 L/min. ; in Experiment IV the water flow was 0.5 L/min. Water temperature and salinity were not regulated but were the same as that of the York River. The dietary supplements were hydrolyzed cornstarch, dextrose in solution, and dextrose mixed with the clay mineral, montmorillonite. Montmorillonite alone was run as a “control” for the montmorillonite and dextrose mix- tures in Experiments I and II but not in Experiment IV; the dextrose and montmorillonite mixture was not used in Experiment III. Individual supplements were prepared by mixing aliquots of either cornstarch, dextrose, mont- morillonite or dextrose and montmorillonite mixed with 3 ooomL of tapwater in 4 ooomL Erlenmeyer flasks. The flasks were fitted with 2-holed rubber stoppers. One hole contained a short piece of glass tubing plugged with cot- ton which functioned as a vent. The second hole contained a glass tube which extended to near the bottom of the flask and served as a delivery tube. Before use, supple- ments in their flasks, rubber stoppers and glass tubing were autoclaved for 10 minutes at 10 pounds pressure (115° C). This treatment sterilized the supplements and suppressed bacterial growth in the holding flasks. It also converted part of the cornstarch into amylose and amylo- pectin through hydrolysis of the (# ) 1-6 linkages. Supplements were delivered to the experimental hold- ing troughs through Tygon tubing attached to the glass delivery tube. Flows were regulated by peristaltic pumps and checked twice daily. Particulate supplements were kept in suspension by means of magnetic bars and stirrers. The contents of each flask lasted about 3 days and as the flasks became empty, they were replaced by full, pre- sterilized, reserve flasks. Addition of supplements to the oyster groups was stopped 24 hours prior to the end of each experiment, and oysters received only York River water during this period so that any undigested supplement might be elim- inated from their digestive tracts. Individual oysters were then cleaned of particulate debris and measured for final underwater shell weight, shell height, total volume, wet tissue weight and glycogen level by the previously de- scribed methods. For each experiment, statistical comparisons between initial and final measurements of each oyster group were conducted to determine if significant changes in the vari- ables of interest occurred during the course of the experi- ment. Comparisons between initial and final mean under- water Shell weights, shell heights and total volumes were facilitated by paired “t”tests (SokAL & ROHLF, 1969). Comparisons between mean wet tissue weights and glyco- gen contents of the sacrificed group to the final values of the control and test groups were carried out by single classification Analysis of Variance and the appropriate F-tests (SOKAL & ROHLF, op. cit.). Statistical comparisons between the final mean measurements of the control group and those of the test groups were also conducted using single classification Analysis of Variance and the appro- priate F-tests. All statistical tests were evaluated at the 5% significance level. All oyster groups were held in the laboratory with the exception of Experiments III and IV where one group per experiment was maintained in a wire tray suspended in the York River to compare “natural changes” in vari- ables to those of the laboratory control group. RESULTS EXPERIMENT I This study was conducted during mid-summer when water temperatures were maximal and after most spawn- ing had occurred. Oyster groups receiving the cornstarch and dextrose supplements and the dextrose and montmorillonite mix- ture had final glycogen levels significantly higher than Vol. 20; No. 4 THE VELIGER Page 355 Table 2 Experiment I: initial and final measurements on oysters (7 July 66-13 Aug. 66). Group Glycogen Wet tissue Shell height Underwater shell weight Total volume designation content weight mm g cc To g Initial Final Initial Final Initial Final Initial 0.51 1.42 39.6 4.88 _ 7.01 _ Control 0.47 1.50 42.5 45.5! 5.33 7.21) 7.51 9.13! Cornst.—2 mg/L 1.06! ? ae: 40.1 45.0! 5.54 8.50! 7.08 O75} Dext.—2 mg/L Welle e 1.31 38.9 44.0! 5.45 7.81} 7.05 9.19! Mont. —2 mg/L 0.63 1.27 37.8 41.8} 5.18 7.35} 7.11 8.96! Dext. & Mont. — 0.87! 2 1.40 415 46.1} 5.46 Tod (e371 9155} 2 mg/L each 'Final mean value is significantly different from the initial mean value, a = 0.05. Final mean value is significantly different from that of the control, a = 0.05. their initial mean levels (Table 2). The control group and the group receiving montmorillonite alone showed no sig- nificant change from initial levels. Only the group receiv- ing cornstarch increased significantly in mean wet tissue weight. Final mean underwater shell weight, shell height and total volume of each oyster group were significantly greater than the initial mean values. Comparison among final mean measurements showed that the oyster groups receiving cornstarch, dextrose and the dextrose and montmorillonite mixture had final gly- cogen levels significantly higher than that of the control group. Levels were respectively 2.2, 2.5, and 1.9 times greater than the control level of 0.47%. Final mean levels of the other variables of all test groups, however, were not significantly different from those of the control group. EXPERIMENT II This experiment was conducted during a period of fall- ing water temperatures when oysters in nature normally begin to accumulate glycogen and spawning has ceased. All oyster groups exhibited significant increases in all variables measured during this experiment (Table 3). Un- fortunately, changes in glycogen levels could not be deter- mined due to the failure of the freezer in which the ini- tially sacrificed tissues were held. Only the cornstarch and the dextrose in solution sup- plements produced final mean glycogen levels significantly higher than that of the control group. Final mean glyco- gen levels of the groups receiving cornstarch at 2 and 5 mg/L showed gains which were respectively 13.6 and 15.6 Table 3 Experiment II: initial and final measurements on oysters (27 August 1966-11 Sept. 66) Group Glycogen Wet tissue ] Shell height Underwater shell weight Total volume designation content weight mm g cc To gz Initial Final Initial Final Initial Final Initial — 1.37 40.9 6.27 — 8.15 ~ Control 0.68 1.88} 41.5 47.8} 6.08 8.40! 8.31 10.45! Cornst.—2 mg/L 9.26? PAOD Ta2 42.0 50.1! 6.15 9.99! 8.19 11.96! Cornst.—5 mg/L 10.59? 3.24} 2 41.2 49.2! 6.28 9.70! 8.56 11.99! Dext.—2 mg/L 1.62? 1.95} 41.7 48.9! 6.21 9.06! 8.08 11.01} Mont.—2 mg/L 1.08 1.67! 41.8 46.1! 648 8.83! 8.14 10.33} Dext. & Mont. — 1.23 2.00! 42.1 47.5} 6.46 9.23} 8.31 11.02} 2 mg/L each Final mean value is significantly different from the initial mean value, a = 0.05. *Final mean value is significantly different from that of the control, a = 0.05. Page 356 THE VELIGER Vol. 20; No. 4 times greater than the control level (0.68%). The group receiving dextrose in solution was only 2.4 times greater than the control level. The significantly higher glycogen levels in the cornstarch-fed groups were accompanied by significantly heavier mean wet tissue weights; this was not so for the dextrose-fed group. As in Experiment I, none of the supplements had a significant influence on shell height, underwater shell weight and total volume. EXPERIMENT III This study was conducted in late fall when all oyster spawning is over and oysters in nature have accumulated glycogen to maximal storage levels (based on the yearly cycle of glycogen accumulation). Final mean measurements, with the exception of shell height, of all laboratory groups were significantly greater than initial measurements (Table 4). Only the group receiving cornstarch at 5mg/L exhibited a significant in- crease in shell height. The York River group did not in- crease significantly in any of the variables measured. At the end of the study, all supplements yielded final mean glycogen levels significantly higher than the control level of 2.8%. The groups receiving the cornstarch supple- ments had levels 3.2 (2mg/L) and 4.1 (5mg/L) times greater than that of the control group. The mean glycogen level of the dextrose-fed group was 1.6 times greater than the control level. However, only the cornstarch supple- ments resulted in final mean wet tissue weights significant- ly greater than that of the control group. The York River group had a final mean glycogen level and wet tissue weight which were significantly less than those of the control group, but did not differ significantly from the control group in the other variables measured. Again, supplements had no significant influence on shell height, underwater shell weight and total volume. EXPERIMENT IV This study was conducted in early spring during a period of rising water temperatures. Early spring is prior to spawning and is normally a period of glycogen accumu- lation after winter depletion in populations of oysters oc- curring in nature. All oyster groups, including the York River group, in- creased significantly in all variables measured during this experiment (Table 5). The cornstarch supplements pro- duced final mean glycogen levels significantly higher than that of the laboratory control group. These levels were ap- proximately 1.5 times greater than the control level of 3.08%. Dextrose in both supplemental forms had no significant influence on glycogen levels. None of the sup- plements had a significant influence on mean wet tissue weights, shell heights, underwater shell weights and total volume. The oyster group held in the York River had final mean glycogen level significantly less than that of the control group, but did not significantly differ from the control group in the other variables measured. DISCUSSION It is assumed in this study, that significant differences be- tween test and control oysters in final measurements were primarily due to the addition of supplements and not to adverse laboratory conditions. This assumption is sup- Table 4 Experiment III: initial and final measurements on oysters (21 Oct. 66-6 Dec. 66). Total volume cc Underwater shell weight & Group Glycogen Wet tissue Shell height designation content weight mm Yo g Initial Final Initial 1.84 York River 2.08? 1.80? Control 2.84) 2.31} Cornst.—2 mg/L 9.131 2 3.10! 2 Cornst.—5 mg/L 11.48! 2 3.09! 2 Dext.—2 mg/L 4.64) 2 2.49} Initial Final Initial Final 1Final mean value is significantly different from the initial mean value, a = 0.05. *Final mean value is significantly different from that of the control, a = 0.05. Vol. 20; No. 4 THE VELIGER Page 357 Table 5 Experiment IV: initial and final measurements on oysters (1 April 67-19 June 67). Group Glycogen Wet tissue Shell height Underwater shell weight Total volume designation content weight mm g cc ones To g Initial Final Initial Final Initial Final a Eee ee ee eee eee eee = Ee Initial 1.67 1.91 46.1 | 6.88 = 9.84 — York River DERM 2 2.94) 44.4 47.8} 6.82 10.30! 9.76 12.24) Control 3.08} 2.46} 44.2 49.3} 6.98 9.47} 9.44 11.32} Cornst.—0.25 mg/L 4.58! ? 2.33! 43.8 50.6! 6.98 9.59} 9.30 11.60} Cornst.—0.50 mg/L 4.73! ? 2.83! 45.8 52.3} 6.68 9.47} 9.86 12.33} Dext.—5 mg/L 3.48! 2.69! 45.2 500m 6.96 9.91} 9.52 12.09! Dext.—5 mg/L & 3.02! 2.49! 44.8 50.8! 7.08 10.10! 9.45 12.02! Mont.—2 mg/L | ‘Final mean value is significantly different from the initial mean value, a = 0.05. 2Final mean value is significantly different from that of the control, a = 0.05. ported by Experiments III and IV in which control oys- ters held in the laboratory showed increases equal to or exceeding those of oysters held in the York River. Our studies showed that effects of carbohydrate sup- plements on glycogen levels varied with season. Seasonal changes in glycogen content are typical of oysters occur- ring naturally in Chesapeake Bay (GaLTsorFr, CHIPMAN, ENGLE & CaLDERWOOD, 1947; ENGLE, 1950). These changes were reflected in glycogen levels among the con- trol groups held in the laboratory; percentage change and absolute levels were lower in summer and early fall than they were in late fall and early spring. This agrees with known aspects with the oyster’s spawning cycle. Fall and spring are periods of glycogen storage, while stored glyco- gen is utilized in the formation of sexual products during the summer months. Effects of cornstarch and dextrose in the fall and spring on glycogen levels paralleled this natural cycle. The oysters receiving cornstarch during these periods of glycogen storage had substantially higher glycogen levels than those normally found in oysters oc- curring in the natural environment. Consequently, utili- zation of carbohydrates as feeding supplements must be considered in relation to the natural glycogen cycle. Comparison of the cornstarch and dextrose results clearly shows that cornstarch was the better supplemental food. Its effect was most noticeable in the fall. At this time concentrations of cornstarch ranging from 2 to 5mg/L greatly influenced both glycogen levels and wet tissue weights, but dextrose influenced only glycogen levels and to a much lesser extent. HavEN (1965) and GmLLEsPrE et al. (1966) concluded from their results that dextrose was limited in value as a carbohydrate supplement for oysters. It is noted that cornstarch at 2 and 5mg/L in both fall experiments resulted in similar levels of percentage glyco- gen (approximately 9 to 11%). This suggests that corn- starch concentrations much over 2mg/L could not be assimilated into stored glycogen. The effect of mixture of dextrose and montmorillonite was, in absolute terms, minimal, the only positive influ- ence on glycogen levels occurring in the summer experi- ment. Assuming that the dextrose was adsorbed on the clay, it is postulated that the oysters were not capable of stripping the adsorbed dextrose off the clay particles. Another possibility is that the uptake route of glucose is through bacteria and that bacteria can not strip the dextrose off the clay particles. Cornstarch at the low concentrations of 0.25 and 0.50 mg/L in the spring experiments influenced glycogen lev- els, yields being only about one-half of those of the 2 and 5mg/L concentrations in the fall experiments. This point is emphasized since it shows that during spring, cornstarch concentrations of only one-tenth of those used during the preceding fall yielded one-half of the glycogen yielded by the higher concentrations. It is also emphasized that during the spring study, water flows were only one-half of those used in the preceding study. The lack of influence of cornstarch on shell height, underwater shell weight and total volume at any season agrees with the data of Haven (1965). GmLLesprE et al. (1966) concluded that cornstarch influenced various pa- rameters of shell size as well as glycogen levels and tissue weight and volume. However, the lack of appropriate statistical comparisons in the presentation of their data makes their contention difficult to support. It is concluded that the influence of cornstarch as a supplement for oys- ters is limited to glycogen content and tissue size with the major influence being on glycogen levels. Thus, supple- ments which influence glycogen levels to a high degree Page 358 THE VELIGER Vol. 20; No. 4 can be expected to have a similar influence on tissue size. This relation supports the statement of MrrcHELL (1917) that investigations leading to increased meat yields must consider supplements which influence glycogen formation. Haven (1965) pointed out that supplemental feeding with cornstarch offered promise as a cultural technique for increasing meat production. Recently the method has been used along with an algal supplement on a routine basis to condition oysters prior to their spawning in a hatchery (Dupuy «& Rivkin, 1972). It is quite probable that the seasonal changes in meat quality of oysters observed by Haven (1962) are associ- ated with the natural fluctuation of a particulate carbo- hydrate in estuarine waters. Possible sources of this sub- stance might be algal cells or detrital material originating from the breakdown of grasses such as Spartina or Zostera. Literature Cited Anprews, J. D. 1961. Measurement of shell growth of oysters by weighing in water. Nat’l. Shellfish. Assoc. Conv. Add. 1961: 1-11 ARMSTRONG, A. R. Undated. The determination of glycogen in oyster meat. Rapid method using the centrifuge. The College of William and Mary; unpubl. 1 p. Baper, R. G. 1962. Some experimental studies with organic compounds and min- erals, pp. 42-57 in Nelson Marshall (ed.) Symposium on the environ- mental chemistry of marine sediments. Narr. Mar. Lab., Occas. Publ. No. 1 CaLpERwWoop, H. N. # A. R. ARMSTRONG 1941. Determination of glycogen in oysters. Chem. 24: 154 - 165 CasTELL, J. D. & D. J. TRwer 1974. Preliminary feed trials using artificial diets to study the nutri- tional requirements of oysters (Crassostrea virginica). Journ. Fish. Res. Brd. Canada 31: 95 - 99 Dupuy, J. L. & S. Rivkin 1972. The development of laboratory techniques for the production of cultch free spat of the oyster Crassostrea virginica. Ches. Sci. 13: 45 -52 ENGLE, James B. 1950. The condition of oysters as measured by the carbohydrate cycle, the condition factor and the percent weight. Nat'l. Shellfish. Assoc. Conv. Add. 150; 20 - 25 Journ. Assoc. Off. Agr. Ga.tsorr, Paut S1mon 1964. The American oyster Crassostrea virginica Gmelin. Fishery Bull. U. S. Fish & Wildlife Serv. 64: 1 - 480; 400 figs.; 46 tables Gattsorr, Paut Simon, W. A. Curpman, Jr., James B. ENGLE & H. N. Catperwoop 1947. Ecological and physiological studies of the effect of sulfate pulp mill wastes on oysters in the York River, Virginia. U.S. Fish. Wildl. Serv. Fish. Bull. 51: 58 - 186 Gavarp, D. 1927. De quoi se nourissent les huitres? Castiglione 2: 238 - 254 Giutespiz, L., R. M. Incite & W. K. Havens, Jr. 1964. Glucose nutrition and longevity in oysters. Fla. Acad. Sci. 27: 279 - 288 1966. Nutritional studies with adult oysters, Crassostrea virginica (Gmelin). Fla. Brd. Conserv., Tech. Ser. No. 51, 26 pp. Haven, D. S. 1962. Seasonal cycle of condition index of oysters in the York and Rappahannock rivers. Proc. Nat'l. Shellfish. Assoc. 51: 42 - 66 1965. Supplemental feeding of oysters with starch. Ches. Sci. 6: 43-51 Hopkins, S. H., J. G. Mackin & R. W. MENZEL 1953. The annual cycle of reproduction, growth and fattening in Louisiana oysters. Nat'l. Shellfish. Assoc. Conv. Add. 1953: 39 - 50 Kemp, A. & A. J. M. Kits van HIEJNINGEN 1954. A colorimetric micromethod for the determination of glycogen Bull. Stat. Agron. Péche Quart. Journ. in tissues. Biochem. Journ. 56: 646 - 648 Kuwatanl, Y. & T. Nisao 1967. Effect of the amount of rice powder supplied as the diet on the growth of Japanese pear] oyster in tank culture. Nat. Research Lab. Bull. 12: 1409-1431 Martin, G. W. 1923. Food of the oyster. Contributions from the Hull Botanical Labor- atory 303. Bot. Gaz. 75: 143 - 169 1972a. Utilization of food by young oysters. Sta. Bull. 442: 12 pp. 1927b. Enteromorpha and the food of oysters. New Jersey Agr. Expt. Science, 66: 662 1928. Experimental feeding of oysters Ecology 9: 49-55 MitcHeE 1, P. H. 1917. Nutrition of oysters: glycogen formation and storage. U.S. Bur. Fish. Bull. 35: 151 - 162 Moore, H. F « T. E. B. Pore 1910. | Oyster culture experiments and investigations in Louisiana. Bur. Fish. Doc. No. 731: 52 pp. NELson, THURLOW CHRISTIAN 1934. Studies of the food and feeding of oysters. Annual Reprts. New Jersey Agr. Exp. Sta. 1933: 19 - 21 1947. Some contributions from the land in determining conditions of life in the sea. Ecol. Monogr. 17: 337 - 346 Soxa., Ropert R. @ FE James Rowr 1969. Biometry. xxi+776 pp.; illust. San Francisco, Calif. (W. H. Freeman and Co.) Yonoce, CHARLES MaAurRIcE 1928. The absorption of glucose by Ostrea edulis. Assoc. U. K. 15: 643 - 653 45th and 46th Journ. Mar. Biol. Vol. 20; No. 4 THE VELIGER Page 359 Further Field Notes on the Behavior of Aplysia dactylomela BY E. TOBACH Department of Animal Behavior, American Museum of Natural History, New York, New York 10024 ON A RECENT COLLECTING TRIP to the waters off La Par- guera, Puerto Rico (21 V through 24 V 1977) 3 observa- tions offer data supportive of previous reports of the be- havior of Aplysia dactylomela Rang, 1828 (Tosacu, Gop & ZIEGLER, 1965; LEDERHENDLER, BELL & Tosacu, 1975; LEDERHENDLER, 1977; LEDERHENDLER & TOBACH, 1977) and Aplysia californica Cooper, 1863 (KUPFER- MANN & CaREw, 1974; AUDESIRK, 1976). Because of weather conditions, it was only possible to collect on one afternoon (21 V) and 3 mornings (22 V, 23 V and 24 V). We searched 2 sites thoroughly on each occasion except as noted below: a reef lagoon called La Gata. The leeward side of La Gata is a shallow, sandy- bottomed area with coral rubble and rock bordering on a shallow water bed of Thalassia. Ulva lactuca and Acanth- ophora spicifera are found in abundance in the Thalassia, but in the rocky area close to the reef. Mangroves border the area in the north. The Enrique site is an extensive shallow water bed of Thalassia, with sandy bottoms, in which A. spicifera, U. lactuca, and Laurencia obtusa are found in profusion. The site is bordered by mangroves on the leeward side. The 3 species of algae are readily eaten by Aplysia dactylomela in the laboratory and they are found feeding on them in these waters (LEDERHENDLER, 1977). On the first afternoon, we tagged 8 animals in the La Gata lagoon in an area approximately 30 meters square. We also tagged 8 animals in an area approximately 40m by 20m in the Enrique area. Only 1 pair of these animals was found copulating. There was a significant difference between the weights of the 2 populations (Mann-Whitney “U” test, p= 0.002). The La Gata sea hares ranged in weight from 25g to 220g (median — 150g), while those found at Enrique ranged from 165g to 450g (median = 350g). The 2 copulating animals were found at La Gata; the sperm donor weighed 420g; the sperm receiver weighed 410g. On the subsequent 3 mornings we found the following: 1. All 8 animals tagged at La Gata were recaptured. Only 2 were recaptured at Enrique. The recaptured pair were copulating; the sperm receiver was the same animal that was the sperm receiver the first afternoon; the sperm donor was a different animal and weighed 450g. 2. An additional 20 animals were captured. On the first morning, 2 of 10 animals were copulating; on the second morning, the ratio was 2/7; on the third day, 6 of 8 were found copulating (see Table 1). Table 1 ‘Reproductive Behavior of Aplysia dactylomela La Parguera, Puerto Rico (1977) Enrique Reef La Gata Reef Not Copulating Copulating Not Copulating Copulating PM: 5/21 2 6 0 8 AM: 5/22 0 0 2 10 5/23 2 1 0 4 5/24 6 2 — —/ 'Not visited because of weather X? = 3.08. d.f. = 1, 0.10>p>0.05 3. At the Enrique area on the third morning, the 8 animals collected were all in a sandy area approximately 10 meters square which was not as rich as surrounding areas in Ulva lactuca and Acanthophora spicifera. No Laurencia was seen. This area had been searched during the previous afternoon and morning visits. We did not find any copulating animals in that area or in the nearby areas where the algae were thick on those occasions, or on the third morning. The recapture of animals in the same area over a 3-day period is not unusual for Aplysta dactylomela as we had previously been able to recapture large numbers of ani- Page 360 THE VELIGER Vol. 20; No. 4 mals over a 12-day period in Bimini (LEDERHENDLER, BELL & ToBAcH, 1975). The difference in numbers of animals found copulating during the morning and after- noon conforms with previous findings (see LEDERHENDLER et al., 1975; LEDERHENDLER & ToBACH, 1977). The gathering of copulating animals in a small area is remi- niscent of the observations made by KUPFERMANN & Ca- REW (1974) and AuDEsIRK (1976) of Aplysia californica. ACKNOWLEDGMENTS I wish to thank Antonio Lopez and Tanina Rostain for their help in making these observations and Dr. I. Leder- hendler for his comments on the manuscript. Literature Cited AuDESIRK, T. 1976. The role of seasonal periodicity, chemical communication and chemoreceptive organs in reproduction in Aplysia californica Cooper. Ph. D. thesis, Univ. South. Calif, Los Angeles, California KuPFERMANN, I. & T. C. Carzw 1974. Behavior patterns of Aplysia californica in its natural environ- ment. Behav. Biol. 12: 317 - 337 LEDERHENDLER, I. Izja 1977. Reproductive behavior of Aplysta dactylomela Rang, 1828 (Ga stropoda : Opisthobranchia). Ph. D. Thesis, City Univ. New York LEDERHENDLER, I. Izya, L. Bert & ETHEL ToBACH 1975. Preliminary observations of the behavior of Aplysia dactylomela (Rang, 1828) in Bimini waters. The Veliger 17 (4): 347 - 353 1 text fig. (1 April 1975) LEDERHENDLER, I. IzjaA & ETHEL ToBACH 1977. Reproductive roles in a simultaneous hermaphrodite, Aplysia dactylomela. Nature 270: 238 - 239 TosacH, ETHEL, PeTER Go_p & Amy ZIEGLER 1965. Preliminary observations on the inking behavior of Aplysia (Varria). The Veliger 8 (1): 16-18 (1 July 1965) Vol. 20; No. 4 THE VELIGER Page 361 Flight Responses of Two Intertidal Gastropods (Prosobranchia : Trochidae) to Sympatric Predatory Gastropods from Barbados DANIEL L. HOFFMAN anv PAUL J. WELDON Department of Biology, Bucknell University, Lewisburg, PA 17837 INTRODUCTION THE ROCKY INTERTIDAL ZONE of Barbados is exceedingly rich in species of gastropods. Lewis (1960) records over 20 different species of prosobranchs inhabiting the narrow littoral zone. At the present time there is an inverse rela- tionship between the number of species and the published reports concerning any aspects of their natural history and behavior. While in residence at the Bellairs Institute of McGill University on Barbados during January, 1977, we studied the escape responses of 2 species of top shells (Prosobranchia, Trochidae), Tegula excavata (Lamarck, 1822), the Green-based Tegula, and Cittartum pica (Lin- naeus, 1758), the West Indian Top Shell. These herbivo- rous gastropods are ideal for studying escape behavior in that they are available in sufficient numbers and that they share their rocky habitat with at least 4 species of carniv- orous gastropods (Prosobranchia, Thaidida@: Purpura pa- tula (Linnaeus, 1758), the wide-mouthed Purpura; Thais haemastoma floridana (Conrad, 1837), the Florida Rock Shell; Thais rustica (Lamarck, 1822), the Rustic Rock Shell; and Thais deltoidea (Lamarck, 1822), the Deltoid Rock Shell. Although temperate water species of top shells show well defined flight responses to asteroids (FEDER, 1963; YARNALL, 1964), the tropical top shells have been reported to show flight behavior to the presence of thaid- id gastropods (Crark, 1958). In a region of the Carib- bean that is depauperate in predatory asteroids (Lewis, op. cit.), the only carnivores that could be implicated as gastropod predators are the thaidids. Although the tem- perate species of Thais (now Nucella, ApBott, 1974) have been identified as predators of acorn barnacles (Con- NELL, 1961, 1970), the number of acorn barnacles that were noted in the rocky intertidal zone of Barbados is extremely sparse and would preclude them from being a major food resource by the gastropods. Through a study of the behavior of the 2 species of trochid gastropods presented with possible predators, we wish to demonstrate the flight responses and shed light on possible trophic relationships among this assemblage of intertidal gastropods. DESCRIPTION or SPECIES Observations on all species of gastropods were made both in the field and in the laboratory during January, 1977. Field data were obtained and collections for laboratory experiments were made at 2 small indentations on the northeast exposed rocky coast of Barbados, River Bay (13°19’N; 59°36’ W) and Little Bay (13°18’N; 59°35’ W). In the laboratory the carnivorous species were kept separate from the herbivorous forms using 2 separate con- crete holding tanks, both of which were supplied with running sea water, the temperature of which varied from 25.5 to 28°C during the period of the experiments. No attempt was made to provide any of the snails with food while in the laboratory, although the carnivorous forms were found to prey on each other. The Herbivorous Gastropods Tegula excavata is a common intertidal gastropod that ranges throughout the rocky shoreline of the Caribbean (AsBott, 1974). On Barbados the species is found in moderately large numbers under rocks and small boulders at the mean low water level, just in or below the “pink zone” of Lewis (1960). There is little published informa- tion concerning the biology of this species. Approximately 150 specimens were collected at River Bay and ranged in shell axis length from 0.5 to 1.2cm. From the size range for this species by WARMKE & ABBoTT (1962), both juve- nile and adult specimens were used in our experiments. Page 362 THE VELIGER Vol. 20; No. 4 Cittarium pica, an economically important species, ranges throughout the rocky intertidal zone of the Carib- bean, but it appears only as fossils in Florida and Bermu- da, although it apparently died out in these 2 areas in comparatively recent times (CLENCH & ABBOTT, 1943). Lewis (1960) reports that the Juvenile stages occur in the “pink zone” above mean low water at Barbados. The adults are found from slightly above the water level to several decimeters below, usually within 60cm of the sur- face (RANDALL, 1964). Many of the natives of Barbados supplement their diet with these large “whelks,” often attaining diameters of over 10cm. Almost all the adult animals that we were able to find were located in grooves and pits on the underside of large rocks and boulders below the mean water level at River Bay; whereas the juvenile animals were collected on the surface of exposed ledges at low tide in the “pink zone” above the mean low water mark at Little Bay. Approximately 100 specimens of C. pica were collected and ranged in shell axis length from 1.0 to 4.8cm, mean length 1.9+ 2.94 S. D. The Carnivorous Gastropods Purpura patula ranges in the Caribbean from Palm Beach, Florida south to Trinidad (CLENcH, 1947). Lewis states that the Barbados population is most abundant from mean sea level to a few decimeters above mean high water, regions Lewis has named the “black zone” and the “green zone.” We have found many specimens of P. patula on the surface of exposed rocks as well as hidden in pits and depressions. Man appears to be the only real preda- tor of this high intertidal gastropod, as many of the natives along the east coast collect them for food. According to Lewis (1960) the diet of P patula consists primarily of the chiton Acanthopleura granulata (Gmelin, 1791) and the barnacle, Tetraclita squamosa Darwin, 1854. Thais haemastoma floridana again has a rather broad distribution from North Carolina, south through the West Indies and along the Central American Coast to Trinidad (CLENCH, 1947). On Barbados, Th. haemastoma flori- dana is an open coast inhabitant found in the “pink zone” of Lewis from below mean low water to mean sea level. Many of the specimens that we collected had shell lengths in excess of 4.0cm. Lewis (1960) again reports that “T: floridana is carnivorous and feeds upon barnacles, chitons and other molluscs.” Also found in the same “pink zone,” but ranging below mean sea level, Thais rustica can be readily confused with the preceding species, although adults rarely exceed 3.0 cm in length (AsBort, 1974). At River Bay we observed many juvenile specimens between 0.4 and 1.2cm below the mean low water mark in the surf zone of Lewis, be- neath the algal canopy or among the clusters of the ver- metid, Spiroglyphus irregularis (d’Orbigny, 1842). This species ranges from South Florida through the Caribbean south to the coast of Brazil (CLENcH, 1947). Lewis makes no mention of this species in his monograph, and little is known of its diet. Thais deltoidea is found still further down in the rocky intertidal zone, generally below the low water level, al- though it does range upward to the lower portion of the “pink zone.” ABBoTr (1974) reports that this species ranges from Jupiter Inlet, Florida, and Bermuda through- out the Caribbean southward to Brazil, and that it is “an abundant species where intertidal rocks are exposed to the ocean surf.” At River Bay and Little Bay, most of the specimens of Th. deltoidea were quite large, shell length 3.0 to 4.5cm, and heavily encrusted with coralline algae. Little is known of its feeding preferences. EXPERIMENTAL METHODS The following experiments were designed to test whether the 2 species of top shells would demonstrate escape re- sponses to specific carnivorous gastropods, and to deter- mine whether such behavior was induced by contact or distance chemoreception. Six to 10 specimens of either Tegula excavata or Cittarium pica were placed in glass finger bowls (top diameter 11.5cm), each of which con- tained approximately 150mL of sea water. The T: excava- ta ranged in shell length from 0.5 to 1.2cm, whereas the C. pica were all juvenile animals with shell length varying from 1.0 to 2.6cm (RANDALL, 1964). The experiments were designed to collect both descriptive and quantita- tive data by counting the number of top shells to leave the water in an arbitrarily determined time of 10 minutes after adding either the predator or an aliquot of sea water that contained the predator’s “scent.” In order to stand- ardize the procedures the following steps were taken: 1) After being placed in the bowls, the snails were al- lowed to come to rest, usually 20 minutes before adding the carnivorous snails, 2) the experiments were per- formed between 0800 and 1100 Barbados time under the subdued lighting conditions of the laboratory, 3) an herbivorous gastropod, Nerita versicolor (Gmelin, 1791) was added to each of the control bowls in order to mimic the predator’s presence by moving about the bowl acting as a potential disruptive agent, 4) to test for distance chemoreception, approximately 25mL of sea water taken from a container that held one carnivorous snail per 100omL of sea water was slowly added to a finger bowl that contained top shells. The control for this experiment was adding an equal volume of sea water from the intake sea water valve to a Vol. 20; No. 4 bow] with snails. When it appeared necessary, the data were tested for significance using the X? test for 2 inde- pendent samples (S1EGEL, 1956). After some initial observations, it became apparent that there may be a differential size response of Cittarium pica to predatory gastropods. To test this hypothesis, the fol- lowing experiment was performed: 6 small, juvenile C. pica (1.0 to 2.6cm) and 6 large specimens (3.5 to 4.8cm) were placed into each of four 2L bowls that were half filled with sea water. After all locomotory activity had ceased in the bowls, usually 15 minutes, a specimen of Thais deltoidea (4.0 to 4.2cm) was placed into each of 2 bowls, and a specimen of Nerita versicolor (3.0 to 3.5cm) was added to each of the remaining 2 bowls as controls. The experiment was designed to measure the number of snails, and the size of these snails, to evacuate the bowls in IO minutes. RESULTS 1. Laboratory Experiments Flight responses were elicited in Tegula excavata and Cittarium pica juveniles by 3 of 4 sympatric gastropod species that were tested in the laboratory (Table 1). A THE VELIGER Page 363 strong response was elicited when contact was made be- tween the top shells and the carnivores. This was most ap- parent with T. excavata, for actual contact was not neces- sary to elicit a strong response in C. pica. The significance of contact to elicit a strong response in T. excavata can be seen in Table 1. The top shells that remain in the bowls, even after adding the carnivores, is almost half the total number. These top shells were not brought into direct contact with the carnivores, a case especially true for Th. deltoidea which is far less active than Th. haemastoma floridana. Generally the flight behaviors of both species were identical. The initial response was a slight elevation of the shell to expose the head, followed by a rapid flail- ing of the cephalic and epipodial tentacles. The top shells then exhibited increased locomotory activity, which usu- ally resulted in the evacuation of the bowl of water. The behavior of Te. excavata differed from that of C. pica ju- veniles in that contact with such carnivorous species as Th. haemastoma floridana and Th. deltoidea often elicited a rapid, jerky torsion or twisting of the shell through an arc of 180°. This was most apparent when contact was made between the propodium of the carnivore and the soft parts or shell of the top shell. No such shell torsion or twisting was demonstrated by C. pica juveniles. On 3 separate occasions during the experiments, a specimen of Table 1 Size of # of topshells # to leave # topshells # to leave Chi- Test Snail in exptl. bowls expu. bowls in control control bowls Square in cm at We in 10 min. bowls at T, in 10 min. Value Tegula excavata 0.5-1.2 Thais haemastoma floridana 3.0-4.2 75 62 73 0 — Thais deltoidea 2.5-4.0 90 4] 90 1 _ Thats rustica 2.0-3.0 40 2] 40 ] — Purpura patula 2.5-3.2 40 1 40 2 _ Th. floridana water 32 ) 32 0 3.471 T. deltoidea water 32 11 32 2 6.178! T. rustica water 32 8 32 it 4.655! Cittarium pica (juveniles) 1.0-2.6 Thais haemastoma floridana 3.0-4.2 40 36 40 0 — Thais deltoidea 2544.2 40 38 40 0 — Thats rustica 2.0-3.0 24 16 24 0 — Purpura patula 2.8-3.3 24 0 24 0 _ T. h. floridana water 40 35 40 0 _ T. deltoidea water 32 31 32 0 _ T. rustica water 24 3 24 1 — 1The difference between the experimental and control counts is significant (p = 0.05) when chi-square is greater than 3.840. Page 364 THE VELIGER Vol. 20; No. 4 Te. excavata was captured, inverted and eaten by a speci- men of Th. haematoma floridana. This occurred when the path to escape was blocked by other top shells in the bowl. As well as exhibiting a strong response to contact, C2tta- rium pica juveniles react strongly to water that contained the “scent” of both Thais haemastoma floridana and Th. deltoidea (Table 1). When Thais water was added to a bowl of quiescent top shells, the snails became agitated and began to move about the bowls to the extent that most evacuated the bowls en masse within the first 3 minutes. On the other hand, Tegula excavata react less strongly to such water samples. Although there was ten- tacular flagellation, movement in the bowls tended to be slower and less direct. Even though a statistically sig- nificant number did vacate the bowls (Table 1), they did not do so simultaneously, and after 10 minutes those that did remain in the bowls had once more become quiescent. There appeared to be a relationship between the age or size of Cittarium pica and its ability to respond to a predator (Table 2). Only the smaller, juvenile animals, Table 2 Differential size response of Cittarium pica to Thais deltoidea Experimental —_ Control Group Group Number of Large Topshells (3.5-4.8 0 0 cm) to vacate bowls in ten min- utes or less (n = 12 per group) Number of Small Topshells (1.0-2.6 1] 0 cm) to vacate bowls in ten min- utes or less (n = 12 per group) with shell lengths between 1.0 and 2.6cm, would respond to the presence of Thais deltoidea. The response was spec- tacular for in most cases the small top shells would evacu- ate the bowl not only within 3 minutes after adding the predator, but in unison. Although no motion was observed in the larger top shells with shell lengths between 3.5 and 4.8cm, the cephalic tentacles did show slight flailing. When bowls that contained similar numbers and sizes of C. pica were moved out of doors in bright light or direct sunlight, all the top shells became highly agitated before the addition of the predator, although they would not eva- cuate the bowls. 2. Field Observations During our residence on Barbados, field trips were made on a daily basis with one purpose being to observe possible feeding behavior of the carnivorous gastropods. Barbados is characterized by mixed semidiurnal tides. All observations were made during the lowest low tide se- quence at either Little Bay or River Bay. Purpura patula was observed feeding on 13 separate occasions on the striped nerite, Nerita tessellata (Gmelin, 1791). In all cases the carnivores were small specimens, averaging about 2.4cm in length. The nerites had been flipped over and the purple shell was covering the ventral surface with its large foot. On 3 separate occasions we noted larger specimens of P patula (average size, 4.0cm) feeding on the chiton, Acanthochiton granulata. The chi- tons had also been flipped over and the purple shell had initially drilled (eaten) a hole through the center of the large foot to get at the viscera. Thais haemastoma floridana and Th. rustica were not observed feeding in the field, although the former was a most opportunistic carnivore, even cannibal, in the labor- atory holding tank. We did note a Jarge number of Tegula excavata shells that were either empty or inhabited by small hermit crabs at River Bay. Generally we would find large specimens of both Th. haemastoma floridana and Th. rustica, in excess of 2.5cm in association with Te. ex- cavata. So they may be guilty of preying on them, guilty at least by association. We did observe a Thais deltoidea (shell length 4.5cm) in the process of consuming a carved star shell, Astraea caelata (Gmelin, 1791) with a shell length of 3.0cm. We were unable to collect large numbers of star shells to test for flight behavior, but in 2 cases we did observe, under the laboratory conditions previously described, the shell twisting phenomenon when this species of star shell was presented to Th. deltoidea. Although we did not make an actual count, we ob- served numerous large empty Cittarium pica shells scat- tered in the intertidal zone of River Bay. Many of these shells were broken at the apical end, exposing the dorsal visceral chamber. The only large carnivore besides man that was in evidence on the beach were the Sally Lightfoot crabs, Grapsus grapsus. Although they easily avoided capture and thus made direct measurements impossible, many of the males appeared to have carapace widths in excess of 15cm. Vol. 20; No. 4 DISCUSSION Tegula excavata and juvenile Cittarium pica exhibit flight behavior that is typical of trochid gastropods (see ANSELL, 1969 for a review). The differences between these 2 species appear to be in the sensory cues used to elicit the response, the complexity of the response, and most im- portantly, the age or size of the responding snail. The responses of Tegula excavata to carnivorous gast- ropods are mediated primarily through contact and in this regard differ from the responses of C. pica juveniles where predator detection appears to involve dis- tance chemoreception. Clearly, there is an advantage con- ferred to a snail able to detect predators from a distance since contact with some carnivorous gastropods may be fatal; for example, contact with cone species where a toxin is injected by means of radular teeth in the eversible proboscis (KoHN, 1959). In the course of our experiments, we have observed that T: excavata, a snail that responds primarily to contact, was captured and eaten by Thais haemastoma floridana on 3 separate occasions, whereas under similar experimental conditions no C. pica were ever captured. Shell twisting in Tegula excavata may be an additional adaptation to contact induced flight behavior as it would serve to extricate the top shell from the grasp of the pre- hensible foot of a carnivorous gastropod. Shell twisting was initially reported in Australian trochid gastropods, induced by thaidid snails (CLarK, 1958). Apparently it is also quite commonly induced by molluscivorous echino- derms and gastropods in such genera as Tegula (FEDER, 1963), Gibbula and, in addition to the more vigorous shell rolling movements, Calliostoma (FEDER, 1967). The abil- ity to twist the shell around the long axis of the visceral mass does cross familial boundaries in that we have ob- served it in the turbinids, Astraea caelata and Turbo castanea (Gmelin, 1791), although in this latter species, the response was elicited by handling and not by a spe- cific predator. The apparent dichotomy in response between adult and juvenile Cittarium pica is noteworthy. Differential size responsiveness has been reported only once. MONTGOMERY (1967) reports that larger specimens of Haliotis rufes- cens Swainson, 1822 and H. assimilis Dall, 1878, are un- responsive to predatory asteroids, while smaller specimens exhibit shell twisting, mantle covering, and mucus ex- pulsion. However, the possible significance of this phe- nomenon is not discussed. Failure of the large specimens to respond can have several explanations, not mutually exclusive: 1) the THE VELIGER Page 365 larger snails may move into a zone unoccupied by preda- tors, flight to which would confer an advantage; 2) the snails may attain a size relative to their predators at which they would be less susceptible to predatory tactics; 3) especially where contact is important, larger speci- mens may not receive stimulation sufficient to elicit a flight response; or 4) ina case where shell twisting is part of the flight repertoire, the shell may reach a bulk at which it would be energetically unfavorable to shell twist. In the case of Cittarium pica, the adults appear to migrate downward, entering a zone where different preda- tors are encountered. RANDALL (1964) notes that one of the large top shells was found in the arms of a small octo- pus, and that octopods readily feed on live top shells. In addition, Randall has discovered top shells or their oper- cula in fish stomachs. Whatever the significance, we feel that it is important that future investigations address themselves to the possibility of size being a relatively im- portant factor in the behavior of mollusks. The 2 species of Barbadian trochids exhibit a strong response to those 3 species of Thais that share their habitat in the mid to lower intertidal zones. Although RANDALL (1964) observed a 3.5cm specimen of Purpura patula feeding on a 3.7cm specimen of Cittarium pica, neither species demonstrated a response to the presence or contact of P patula, which from field observations appears to prey on other species of mollusks. Purpura patula is a high intertidal carnivore that may only on rare occasions be found in the lower zone inhabited by the 2 species of top shells. This situation may be similar to that of Acmaea scabra (Gould, 1846) which does not show an escape response to Pisaster ochraceus, with which it overlaps only at the lower extreme of its range in the intertidal zone, yet ranks high on a percentage basis of food for the asteroid (FEDER, 1963). We feel that the relatively strong response of both Tegula excavata and juvenile Cittarium pica to Thais haemastoma floridana, Th. deltoidea and Th. rusti- ca has ramifications in the predator-prey interactions of these species. Whether or not top shells comprise the major portion of the thaidids’ diet and the extent to which the flight responses of potential prey are involved, remain unanswered until more ecological data are available. SUMMARY Two species of mid to lower intertidal top shells, Tegula excavata and Cittartum pica, exhibit flight behavior to the presence of 3 sympatric thaidid gastropods. The former species tends to respond strongly to contact with predatory Page 366 THE VELIGER Vol. 20; No. 4 forms; whereas the latter primarily responds to distance chemoreception. Neither of the snail species responds to a high intertidal predatory gastropod, Purpura patula. Also, there are 2 major differences in the behavior of the 2 top shell species in that only Tegula excavata shows the shell twisting response to contact and it is only the small or juvenile Cittarium pica that have a flight reaction to the predators, the adults being relatively unresponsive. ACKNOWLEDGMENTS We wish to thank Dr. Finn Sander, Director of the Bell- airs Research Institute of McGill University for facilities and warm hospitality afforded us while in Barbados. Also Dr. Joseph Rosewater, Division of Malacology, the U. S. National Museum for identifying several of the species of gastropods reported in this study. And finally Debbie Barnard, Joe McGinley and Jeff Churchill for their much appreciated assistance in the field and laboratory. Literature Cited Assotr, Rosexrt Tuckzr 1974. American seashells. Van Nostrand Reinhold C., New York, 2zd ed.,; 663 pp.; 24 color plts.; numerous figs. ANSELL, ALan Davip 1969. Defensive adaptations to predation in the mollusca. Symp. Mar. Biol. Assoc. India 487 - 512 Crarg, W. C. 1958. Escape responses of herbivorous gastropods when stimulated by carnivorous gastropods. Nature 4602: 137 - 138 CiencH, WILLIAM JAMES 1947. The genera Purpura and Thais in the Western Atlantic. Johnsonia 2 (23): 61-92 Cenc, Wituiam JAMES & RoperT TuCKER ABBOTT 1943. The genera Gaza and Livonia in the Western Atlantic. John- sonia 12: 1-12 Connett, Joserx H. 1961. Effects of competition, predation by Thais lapillus, and other fac- tors on natural populations of the barnacle Balanus balanoides. Ecol. Monogr. 31 (1): 61-104; 22 text figs.; 17 tables 1970. A predator-prey system in the marine intertidal region. I. Bale nus glandula and several predatory species of Thats. EcoL Monogr. 40 (1): 49-78; 9 figs.; 17 tables Fgeprer, Howarp MITrcHELL 1963. Gastropod defense responses and their effectiveness in reducing predation by starfishes. Ecology 44 (3): 505-512 1967. | Organisms responsive to predatory sea-stars. 371 - 394 Kon, Aan Jacoss 1959. The ecology of Conus in Hawaii. 47-90; 30 figs.; 19 tables Lewis, Joun B. 1960. The fauna of rocky shores of Barbados, West Indies. Journ. Zool. 38: 391 - 435; 20 figs. Montcomery, Davy H. 1967. Responses of two haliotid gastropods (Mollusca), Haliotis assimi- lis and Haliotis rufescens, to the forcipulate asteroids (Echinodermata), Pycnopodia helianthoides and Pisaster ochraceus. The Veliger 9 (4): 359 - 368; plts. 51, 52; 2 text figs. (1 April 1967) Sarsia 29: Ecol. Monogr. 29 (1): Canad. RanpDALL, HELEN A. 1964. A study of the growth and other aspects of the biology of the West Indian Topshell, Cittarium pica (Linnaeus). Bull. Mar. Sci. Gulf and Caribb. 14 (3): 424-443; 10 figs.; 1 table SIEGEL, SIDNEY F. 1956. | Nonparametric statistics for the behavioral sciences. McGraw-Hill, New York WarmMxe, GerMAINE L. & RospertT Tucker Assott 1961. Caribbean seashells; a guide to the marine mollusks of Puerto Rico and other West Indian islands, Bermuda and the lower Florida Keys. Livingston Publ. Co., Narberth, Pa. x+345 pp.; 44 plts.; 34 text figs. YARNALL, JoHBN L. 1964. The responses of Tegula funebralis to starfishes and snails ((Mollusca: Gastropoda). The Veliger 6 (Supplement) : 56 - 58; 2 tables (15 November 1964) 312 pp. Vol. 20; No. 4 THE VELIGER Page 367 Notes on the Spawning and Egg Capsules of Two Prosobranch Gastropods: Nassarius tiarula (Kiener, 1841) and Solenosteira macrospira (Berry, 1957) ROY S. HOUSTON Department of Biology, Loyola Marymount University, Los Angeles, California 90045 (2 Text figures) Botu Nassarius tiarula (Kiener, 1841) and Solenosteira macrospira (Berry, 1957) occur intertidally on mudflats throughout the Gulf of California. According to KEEN (1971), S. macrospira is endemic to the Gulf, but N. tia- rula ranges as far south as Panama. Collections and ob- servations for this study were made at Cholla Bay, 11 km north of Puerto Penasco in the northern Gulf of Cali- fornia. Spawning can be observed from April through the last part of June for Nassarius tiarula. Females deposit pale yellow egg capsules in clusters of 15 to 30 on empty shells lying just beneath the surface of the mud. The capsules are vase-shaped and have 4 sides. The front and back are flat with slightly convex sides (Figure 1a). Each has an opening at the top fitted with a mucous plug. The bottom of the capsule tapers to a long, narrow stalk, which is attached to an adhesive disk. The entire capsule measures Figure 1 Egg capsules: a-Nassarius tiarula; b-Solenosteira macrospira 13mm high and about 3mm wide. Each capsule contains about 50 eggs which float in an albuminous fluid. During dissection of gravid females, egg capsules were released from the genital opening. First a small trans- parent bubble appeared which proved to be the basal region of the capsule. As the bubble was squeezed out, the opening was distended to about 3 times its normal dia- meter. Shortly after, the eggs were released and appeared initially as a single string surrounded by albuminous fluid. Prior to release of the capsule, the eggs clumped into a central mass. The top of the capsule with the mucous plug intact was the last to leave the nidamental opening. This newly formed capsule was soft and transparent. It was then transported along a ciliated groove to the foot. Histological sections reveal the 5 layers that compose the capsule wall (Figure 2a). An innermost mucous layer (mu) is surrounded by a coat of circular fibers (ci). Peripheral to this is a loose network of fibers interspersed by lacunae (mpl), which are filled with a mucoid sub- stance. The 4" layer is another stratum of circular fibers. The final layer is a tenuous mucous coat surrounding the entire capsule. Solenosteira macrospira begins spawning in March and continues through the first week of June. This is an inter- esting species because the female deposits the capsules on the shell of a living male buried beneath the mud. On many occasions the shell of the male was completely covered by the capsules. GEMMELL (1973) first reported this behavior in populations of S. macrospira near San Felipe, Baja California. The capsules are transparent and the reddish-brown eggs can be observed inside floating in the albuminous fluid. The flask-shaped capsules stand about 7mm high and are attached to the shell by a long, slender stalk similar to that of Nassarius tiarula (Figure 1b). In each Page 368 THE VELIGER Vol. 20; No. 4 Figure 2 Egg Capsule Histology a- Nassarius tiarula b - Solenosteira macrospira ci — circular protein fibers mp — muco-protein mpl — muco-protein fibers interspersed by lacunae mpc — muco-protein fibers mu — mucus capsule there are up to 500 eggs averaging 200m in diameter. The wall of the egg capsule consists of 3 layers (Figure 2b). The inner region is a muco-protein layer (mp), not a mucous one as in Nassarius tiarula. The middle layer is a thick, fibrous muco-protein coat (mpc). This is covered by an outer layer consisting of circular fibers (ci). Sim- ilar conditions were observed by FRETTER (1941) for egg capsules of Nucella lapillus (Linnaeus, 1758). ANKEL (1937) suggested that the egg capsules of N. lapillus are composed of 2 substances, the protein and conchiolin intermixed with mucus or a mucoid substance. This has been substantiated in histochemical studies by BAYNE (1968), in which acid mucopolysaccharides and conchi- olin were found to be the major components of the capsule wall. The long slender stalks allow the egg capsules to extend above the mud where the developing embryos are ex- posed to ample water circulation. Since competition for hard substrates is severe, females of Solenosteira macro- Spira have overcome this problem by depositing egg cap- sules on the shells of their mates. ACKNOWLEDGMENT I am grateful to Dr. Carl Kadner for reviewing the manuscript. Literature Cited ANngEEL, WuLF Emmo 1937. Der feinere Bau des Kokons der Purpurschnecke Nucella lapil- lus (L.) und seine Bedeutung fiir das Laichleben. Verh. deutsch. Zool. Gesellsch. 39: 77 - 86 BayNE, CHRISTOPHER JEFFREY 1968. Histochemical studies on the egg capsules of eight gastropod molluscs. Proc. Malacol. Soc. London 38: 199-211 FReETTER, VERA 1941. The genital ducts of some British stenoglossan prosobranchs. Journ. Mar. Biol. Assoc. U. K. 25: 173 - 211 GesMELL, Joyce 1973. Field observations on gastropod breeding and egg laying. The Festivus 4 (5): 32-34 Keen, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. Stanford Univ. Press, Stanford, Calif. i- xiv+ 1066 pp.; ca. 4000 figs.; 22 color pits. (1 September 1971) Vol. 20; No. 4 THE VELIGER Page 369 Additional Molluscan Records from Bahia de Los Angeles, Baja California Norte FORREST L. POORMAN anp LEROY H. POORMAN'! Los Angeles County Museum of Natural History (1 Map) Bauia DE Los ANGELEs is located on the east side of the Baja California Peninsula about 440km south of the Unit- ed States Border (29°00’N; 113°30’'W). The area covers approximately 63 square kilometers and is protected on the east side by about 15 small islands, with Isla Smith as the largest. The literature records a number of expeditions which collected in this part of the Gulf of California. The earli- est of these was the Albatross in 1911. Although several of the expeditions did collect in the bahia, most attention was given to dredging around Isla Angel de la Guarda, more than 30km offshore. The reports of the molluscan fauna of the bahia were scarcely significant, the largest number of species reported being 28 by STEINBECK & RICKETTS (1941). Two more recent papers have been published in the Transactions of the San Diego Society of Natural History, reporting on the mollusks collected at Bahia de Los An- geles. McLEAN (1961) reported on the mollusks collected from shore and beach drift. This paper included records from his collection and 4 other sources. Coan (1968) reported benthic species from the inner part of the bahia. Collecting for that report was done with an orange peel grab and followed a grid over the southern part of the bahia where the bottom is primarily silty mud. Accompanied by Carl and Laura Shy of Westminster, California, we have made 3 collecting trips of about 10 days each to this area in recent years. Base camp was at Punta la Gringa, a small point of land about 14km by road north of the village. Using small boats and hand- operated dredging equipment, we made approximately 240 drags in depths of 20- 30m. The maximum depth recorded in the bahia is 4om (BARNARD & Grapy, 1968). The bottom was primarily broken rock and shell but in- * 15300 Magnolia Street, Westminster, CA(lifornia) 92683 cluded samplings of other substrates as indicated on the map. Each trip was associated with periods of extreme low tides, so we did extensive intertidal collecting. Some of this was done along the rocky reefs between Punta la Gringa and the sand spit north of the village. Much time was spent along the rocky shore several kilometers north of camp and along the west side of Isla Smith. A number of species of marine algae and sponges cover the rocky intertidal areas and extend out to about 10m. Many of the specimens obtained at the bahia have been significantly larger than the average sizes indicated in KEEN (1971), and have been registered with the Lost Operculum Club, a project of the Conchological Club of Southern California. RoDEN « Groves (1959) report, “An area of low temperatures around Isla Angel de la Guarda is found throughout the year.” This is the second largest island in the Gulf of California and parallels the coast about 32km from the bahia. The channel between reaches a depth of more than 600 fathoms (1096m). Strong tidal currents force cold water from these depths into the bahia. The maximum temperature in the bahia was reported to be 29.8° C in August, 1962, and the min- imum temperature was reported to be 15° C in February, 1962 (BaRNaRD & Grapy, 1968). The unusually cold water together with the abundance of algae, sponges, coral, and other food sources may contribute to an ex- planation of why so many “giants of the species” are found here. A total of 510 different identified mollusks was included in the papers by McLean and Coan. Only 67 species were reported by both authors. This paper adds 160 identified species to their total, which increases the recorded mol- luscan fauna for Bahia de los Angeles by 36% for a total of 603 species. Page 370 THE VELIGER Vol. 20; No. 4 The following list includes those mollusks collected by the Shys and the Poormans and which were not reported in either of the other 2 papers. Identifying numbers from KEEN (1971) precede the names of each species listed. Bahia de Los Angeles,, Baja California Norte, Mexico 1. broken rock and shell _—3.. silty mud 2. fine sand and shell 4. fine living coral 5. valves of Pecten vogdest Dredging or intertidal records are indicated by D or S after each species name. Significant range extensions are also included. Several undescribed opisthobranchs and chitons were taken and are now in manuscript. Our appreciation is extended to Gale Sphon, Los An- geles County Museum of Natural History, for his help in identifying opisthobranch species, and to George Hansel- man, San Diego, for help with the chitons. PELECYPODA ARCIDAE 77 Anadara (Anadara) adamsi Olsson, 1961 D (Panama) g2 Anadara (Rasia) formosa (Sowerby, 1833) D (Cedros Isld.) GLyYCYMERDIDAE 117 Glycymeris (Tucetona) strigilata (Sowerby, 1833) D MytTILDAE 119 Brachidontes adamsianus (Dunker, 1857) S 122 Brachidontes semilaevis (Menke, 1849) S PEcTINIDAE 183 Chlamys lowei (Hertlein, 1935) D 187 Cyclopecten exquisitus Grau, 1959 D 194 Delectopecten zacae (Hertlein, 1935) D PLICATULIDAE 207 Plicatula inezana Durham, 1950 S _ (Southern Gulf of Cali- fornia) 208 Plicatula penicillata Carpenter, 1857 D (Southern Gulf of California) SPONDYLIDAE 213 Spondylus ursipes Berry, 1959 D LuciINDAE 271 Lucina (Here) excavata Carpenter, 1857 D 277 Lucina (Pleurolucina) leucocymoides (Lowe, 1935) D 278 Lucina (Pleurolucina) undatoides Hertlein « Strong, 1945 D UNGULINDAE 293 Diplodonta suprema Olsson, 1961 S (Panama) 296 Phlyctiderma (Phlyctiderma) discrepans (Carpenter, 1857) D SPORTELLIDAE 343 Basterotia (Basterotella) hertleini Durham, 1950 S Vol. 20; No. 4 CHAMIDAE 348 Chama frondosa Broderip, 1835 D 352 Chama venosa Reeve, 1847 D 357 Pseudochama janus (Reeve, 1847) D CARDIDAE 373 Lophocardium annettae (Dall, 1889) D VENERIDAE 381 Ventricolaria isocardia (Verrill, 1870) D 401 Pitar (Pitar) helenae Olsson, 1961 D 414 Pitar (Lamelliconcha) frizzelli Hertlein « Strong, 1948 D (Southern Gulf of California) 424 Megapitaria aurantiaca (Sowerby, 1831) D 456 Chione (Chionopsis) purpurissata Dall, 1902 D ~ MactTrDAE 490 Mactra (Mactrotoma) nasuta Gould, 1851 D 497 Anatina cyprinus (Wood, 1828) D TELLINDAE 517 Tellina (Angulus) meropsis Dall, 1900 D 546 Tellina (Phyllodina) pristiphora Dall, 1900 D 551 Tellina (Tellinella) cumingu Hanley, 1844 D PSAMMOBIDAE 603 Gari (Gobraeus) helenae Olsson, 1961 D SOLECURTIDAE 614 Solecurtus guaymasensis Lowe, 1935 D (Guaymas, Sonora) SEMELIDAE 626 Semele californica (Reeve, 1853 ex A. Adams, MS) S (Cabo San Lucas) 634 Semele junonia (Verrill, 1870) D 636 Semele lenticularis (Sowerby, 1833) DS (Barra de Navidad, Jalisco) 647 Semele simplicissima Pilsbry « Lowe, 1932 D My Dare 673 Sphenia fragilis (H. & A. Adams, 1854) D CorBULIDAE 687 Corbula (Juliacorbula) ira Dall, 1908 D (Panama) 692 Corbula (Varicorbula) speciosa Reeve, 1843 D PANDORIDAE 733 Pandora (Pandora) uncifera Pilsbry « Lowe, 1932 D 738 Pandora (Pandorella) cornuta C.B. Adams, 1852 D_ (Pana- ma) 739 Pandora (Pandorella) granulata Dall, 1915 D THE VELIGER Page 371 LYoNSDDAE 747 Entodesma (Phlycticoncha) lucasanum (Bartsch « Rehder, 1939) D THRACIDAE 764 Cyathodonta dubiosa Dall, 1915 D VERTICORDIDAE 789 Verticordia (Verticordia) ornata (Orbigny, 1846) D GASTROPODA FISSURELLDAE 13 Rimula mexicana Berry, 1969 D 22 Diodora pusilla Berry, 1959 S 23 Diodora saturnalis (Carpenter, 1864) D 43 Leurolepas roseola McLean, 1970 D TROCHDAE 86 Calliostoma mcleani Shasky « Campbell, 1964 D 97 Tegula (Agathistoma) corteziana McLean, 1970 S SKENEIDAE 124 Parviturbo stearnsti (Dall, 1918) D LIoTIDAE 130 Arene (Arene) lurida (Dall, 1913) D 134 Arene (Marevalvata) balboai (Strong & Hertlein, 1939) D 135 Arene (Otollonia) fricki (Crosse, 1865) D 138 Macrarene lepidotera McLean, 1970 D_ (Socorro Island) TURBINIDAE 149 Turbo (Marmarostoma) squamiger Reeve, 1843 D LiTTORINIDAE 179 Littorina aberrans Philippi, 1846 S VITRINELLIDAE 393 Teinostoma (Teinostoma) politum A. Adams, 1851 S_ (Ec- uador) ARCHITECTONICIDAE 429 Heliacus caelatus (Hinds, 1844) D Page 372 THE VELIGER Vol. 20; No. 4 441 442 445 447 TURRITELLIDAE Turritella mariana Dall, 1908 D (Guaymas, Sonora) Turritella nodulosa King & Broderip, 1832 D (Guaymas, Sonora) Turnitella rubescens Reeve, 1849 D Vermicularia frisbeyae McLean, 1970 DS catita, Jalisco) (Bahia de Tena- CERITHODAE 558 Diastoma chrysalloidea Bartsch, 1911 S 633 634 636 678 686 797 752 797 817 B22 828 866 893 972 988 1019 1020 1021 1058 1063 1093 EprronmDAE Epitonium (Cirsotrema) togatum Hertlein « Strong, 1951 D Epitonium (Cirsotrema) vulpinum (Hinds, 1844) D Epitonium (Hirtoscala) reflexum (Carpenter, 1856) S Opalia (Dentiscala) crenatoides (Carpenter, 1864) D Opalia (Nodiscala) spongiosa Carpenter, 1864 D EULIMDAE Eulima townsendi (Bartsch, 1917) D Niso (Niso) splendidula (Sowerby, 1834) D VANIKORIDAE Vanikoro aperta (Carpenter, 1864) D CALYPTRAEIDAE Crepidula striolata Menke, 1851 D Crucibulum (Crucibulum) lignartum (Broderip, 1834) D Crucibulum (Dispotaea) concameratum Reeve, 1859 D NATICIDAE Natica (Natica) scethra Dall, 1908 D (Panama) Sinum sanctijohannis (Pilsbry « Lowe, 1932) D (Nicaragua) CoLUBRARIDAE Colubraria (Colubraria) siphonata (Reeve, 1844) D MuRICIDAE Murexiella humilis (Broderip, 1833) D Aspella (Trialatella) cunninghamae (Berry, 1964) D Attiliosa carmen (Lowe, 1935) D : Attiliosa incompta (Berry, 1960) D Pterotyphis (Tripterotyphis) lowei (Pilsbry, 1931) D CoRALLIOPHILIDAE Coralliophila (Pseudomurex) costata (Blainville, 1832) S THADDAE Morula (Morunella) lugubris (C. B. Adams, 1852) D 1099 1124 1252 1261 1265 1273 1293 1304 1363 1379 1382 1510 1531 1535 1538 1954 1579 1599 1615 1637 1641 1643 1648 1656 1663 1676 1688 1691 1699 1701 1702 1723 1727 1777 1791 1810 1813 1839 BuccinDAz Caducifer (Monostiolum) biliratus (Reeve, 1846) D Engina fusiformis Stearns, 1894 D CoLUMBELLIDAE Nassarina (Steironepion) tincta (Carpenter, 1864) D Parametaria dupontii (Kiener, 1849-50) S Strombina (Strombina) angularis (Sowerby, 1832) D Strombina (Strombina) gibberula (Sowerby, 1832) D NASSARIDAE Nassarius cerritensis (Arnold, 1903) D Nassarius insculptus (Carpenter, 1864) D OLIVIDAE Oliva (Oliva) polpasta Duclos, 1833 DS Olivella (Olivella) gracilis (Broderip « Sowerby, 1829) D Olivella (Olivella) sphoni Burch « Campbell, 1963 D ConDAE Conus (Lithoconus) archon Broderip, 1833 D TEREBRIDAE Terebra crenifera Deshayes, 1859 D Terebra elata Hinds, 1844 D Terebra glauca Hinds, 1844 D Terebra ornata Gray, 1834 D TURRIDAE Calliclava alcmene (Dall, 1919) D Kylix paziana (Dall, 1919) D Drillia (Drillia) acapulcana (Lowe, 1935) D Splendrillia bratcherae McLean & Poorman, 1971 D Bellaspira acclivicosta McLean « Poorman, 1970 D Bellaspira melea Dall, 1919 D Polystira oxytropis (Sowerby, 1834) D Knefastia dalli Bartsch, 1944 S Pyrgospira obeliscus (Reeve, 1843) D Crassispira (Crassispira) maura (Sowerby, 1834) D Crassispira (Crassispirella) rustica (Sowerby, 1834) D Crassispira (Dallspira) bifurca (E. A. Smith, 1888) S$ Crassispira (Striospira) kluthi E. K. Jordan, 1936 D Crassispira (Striospira) tepocana Dall, 1919 D Crassispira (Striospira) xanti Hertlein « Strong, 1951 D Carinodrillia dichroa Pilsbry x Lowe, 1932 D Strictispira ericana (Hertlein « Strong, 1951) D Nannodiella fraternalis (Dall, 1919) D Kurtziella (Granoturris) antipyrgus (Pilsbry « Lowe, 1932) D Kurtzia arteaga (Dall « Bartsch, 1910) D Kurtzia granulatissima (Mérch, 1860) D Daphnella mazatlanica Pilsbry & Lowe, 1932 S Vol. 20; No. 4 ACTEONIDAE 2229 Acteon panamensis Dall, 1908 D (Guaymas, Sonora) ATYIDAE 2245 Haminoea virescens (Sowerby, 1833) S AGLAJIDAE 2254 Chelidonura inermis (Cooper, 1863) S APLYSIDAE 2296 Aplysia (Neaplysia) californica Cooper, 1863 S —— Phyllaplysia padina Williams « Gosliner, 1973 S PLEUROBRANCHDAE 2306 Pleurobranchus areolatus (Morch, 1863) S Berthellina citrina (Ruppell « Leuckart, 1828) S DorIDIDAE 2327 Conualevia mizuna Marcus & Marcus, 1967 S CHROMODORIDIDAE 2328 Chromodoris banksi Farmer, 1963 S POLYCERATIDAE 2350 Polycera alabe Collier « Farmer, 1964 S 2352 Polycera hedgpethi Marcus, 1964 S 2353 Laila cockerelli MacFarland, 1905 DS GyYMNODORIDIDAE 2354 Nembrotha eliora Marcus & Marcus, 1967 DS DENDRODORIDIDAE 2359 Dendrodoris krebsii (Mérch, 1863) S — Dendrodoris fulvua (MacFarland, 1905) S 2360 Doriopsilla albopunctata (Cooper, 1863) S 2362 Doriopsilla rowena Marcus & Marcus, 1967 S TETHYIDAE 2370 Melibe leonina (Gould, 1852) S ARMINIDAE 2373 Armina californica (Cooper, 1862) DS 2374 Histiomena convolvula (Lance, 1962) D FLABELLINIDAE 2382 Coryphella iodinea (Cooper, 1863) D AEOLIDIDAE 2388 Spurilla chromosoma Cockerell « Eliot, 1905 S THE VELIGER Page 373 ONCHIDIDAE 2395 Onchidella binneyi Stearns, 1893 S MELAMPIDAE 2399 Melampus (Melampus) mousleyi Berry, 1964 S POLYPLACOPHORA ISCHNOCHITONIDAE 16a Radsiella eucosmius (Dall, 1919) D 23x Stenoplax circumsenta Berry, 1956 D 32 Lepidozona crockeri (Willett in Hertlein « Strong, 1951) D 35 Lepidozona formosa Ferreira, 1974 D 37 Lepidozona serrata (Carpenter, 1864) D MopPALIODAE 48 Dendrochiton lirulatus Berry, 1963 D 50 Placiphorella velata Dall, 1879 S SCAPHOPODA DENTALIDAE 7 Dentalium (Graptacme) sectum Deshayes, 1826 D 11 Dentalium (Tesseracme) tesseragonum Sowerby, 1832 D SIPHONODENTALIDAE 17 Siphonodentalium quadrifissatum (Pilsbry & Sharp, 1898) D Literature Cited Barnarp, J. Laurens & J. R. Grapy 1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. I. General Account. Trans. San Diego Soc. Nat. Hist. 15 (6): 51-66 Coan, Euceng Victor 1968. A biological survey of Bahfa de Los Angeles, Gulf of California, Mexico. III. Benthic Mollusca. Trans. San Diego Soc. Nat. Hist. 15 (8): 107-132; 2 text figs. (25 September 1968) Draper, Bertram C. 1973. Lost Operculum Club list of champions. Conch. Club South. California. 1975. Supplement to “Lost Operculum Club list of champions.” Conch. Club South. California. DuSuane, HELen 1974. The Panamic-Galapagan Epitoniidae. plement): 1-84; 15 plts.; 5 text figs.; 1 map DuSwane, HELEN & Roy PooRMAN 1967. A checklist of mollusks for Guaymas, Sonora, Mexico. The Veliger 9 (4): 413-441; 1 map (1 April 1967) Keen, A. Myrna 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. Stanford Univ. Press, Stanford, Calif. i- xiv+ 1066 pp.; ca. 4000 figs.; 22 color plts. (1 September 1971) The Veliger 16 (Sup- (31 May 1974) Page 374 THE VELIGER Vol. 20; No. 4 Keen A. Myra & EucENE Victor Coan 1975. Sea shells of tropical West America: additions and corrections to 1975. West. Soc. Malacol. Occas. Paper 1: 66 pp.; 2 text figs. (22 June 1975) McLean, James HamILTON 1961. Marine mollusks from Los Angeles Bay, Gulf of California. Trans. San Diego Soc. Nat. Hist. 12 (28): 449-476; figs. 1-3 (15 August 1961) Ropen, Gunnar I. & G. W. Groves 1959. Recent oceanographic investigations in the Gulf of California. Journ. Mar. Res. 18: 10 - 35 STEINBECK, JoHN & Epwarp F RicketTTs 1941. Sea of Cortez. Viking Press, New York: x+598 pp.; 40 plts. TowNnseEND, Cuares H. 1916. Voyage of the “Albatross” to the Gulf of California in 1911. Bull. Am. Mus. Nat. Hist. 35: 399 - 476 Vol. 20; No. 4 THE VELIGER Page 375 Tambja and Roboastra (Mollusca : Opisthobranchia ) from the Gulf of California and the Galapagos Islands BY WESLEY M. FARMER 1327 East Donner Drive, Tempe, Arizona 85282 (20 Text figures) EvELINE AND Ernst Marcus and James R. Lance pub- lished a description of Nembrotha eliora Marcus & Mar- cus, 1967, and Nembrotha hubbsi Lance, 1968, respec- tively. The papers describe the same animal species as shown by photographs, illustrations, and descriptions. Fur- ther work now shows that Marcus & Marcus’ N. eliora was described from 2 lots of material. Their “material one”’ is similar to Lance’s species and therefore N. hubbsi is considered a junior synonym of N. eliora. The Marcuses have 2 species in their description. Their “‘material two” is of a new species of Tambja. Lance’s N. hubbsi also in- volves 2 different species. His description of N. hubbsi is correct, except the radula figure is that of a Roboastra. The members of the genus Nembrotha as used by Mar- cus & Marcus have denticles or serrations on the leading edge of the teeth. In Tambja, the leading edge of the rachidian is smooth. NUDIBRANCHIA Doridoidea PHANEROBRANCHIA NONSUCTORIA GyYMNODORIDDAE Tambja Burn, 1962 Type species: Nembrotha (?) verconis BasEpow « HEDLEY, 1905; 29: 146, 158, 159; plt. 2, figs. 1-3 “Rachidian is rectangular with notched or smooth up- per margin, lateral tooth has a bifid or simple cusp with 3 - 7 marginal plates. The buccal collar is strong; a labial armature is absent. The small prostate gland is confined to a glandular section of the vas deferens.” (BurN, 1967: 214). Tambja eliora (Marcus & Marcus, 1967) (Figures 1 - 3, 18) Nembrotha eliora Marcus & Marcus, 1967: 194 - 196; figs. 57, 58; — FarMER, 1968: 26, 27 (fig.); - 1970a: 16; — 1970b: 73, 75, 78 (figs. 6, 8), 79, 84; — 1970c: 5, 12, figs.; -KEEN, 1971: 827, 828; plt. XX, fig.3; — ABBoTt, 1974: 358; - SKOGLUND, 1974: 5 Nembrotha hubbsi Lance, 1968: 6-8; plt. 1, fig. 6 Syntypic material (eliora) : the holotype number is USNM 678410 and radular slide no. 915.6. One specimen and its radula are represented. The type of N. hubbst is an intact specimen, CASIZ type series no. 315. The radula illustrated is of a Roboastra from another specimen no longer intact. Type Locality: Puerto Lobos, Sonora, Mexico. Distribution: Angel de la Guarda Island, Guaymas, Es- condido Bay, Carmen Island, Monserrate Island, San Figure 1 Tambja eliora (Marcus & Marcus, 1967) Page 376 THE VELIGER Vol. 20; No. 4 Diego Island, Punta Entrada, Espiritu Santo Island, La Paz, Cerralvo Island, and Magdalena Bay, Mexico. Description: The radula, based on a minimum of speci- mens, is typically that of a Tambja with the formula 15X5-4°1°1°1°4-5 (Figure 2). No labial armature Os Figure 2 Radula of Zambja eliora (Marcus « Marcus, 1967) is evident. The animal can attain a length of 50mm and is limaciform. The foot color is gray; the edge of the foot is blue. Three marked turquoise blue stripes on the dorsum are bordered by black. The rhinophores have 22 leaves. The gills are unipinnate and nearly black in color; the axis of the gills is marked with turquoise, black, and yellow ochre. The morphology of the head is characteristic in this animal where the cephalic shield dips into a “V” or “U” shape (Figure 1). The cephalic tentacles are broad and flat. A dorsal orientation of the reproductive system shows many of the parts in surface view. Carefully teasing the organs apart (Figure 3), the atrium is found to be en- larged. The hermaphrodite duct leads into the large vag- inal gland by way of an ampulla. A thin sac-like structure is connected to the nidamental duct as is the adjacent round spermatotheca. Behavior: Tambja eliora swims by undulating the body from side to side (FARMER, 1970b). This is an escape response to the advances of the predatory slug, Roboastra tigris Farmer (spec. nov., herein) (Figure 18). Other possible reasons for swimming are not known. Discussion: The “material one” of Marcus & Marcus, 1967, is compatible with the above description. Their “material two” is also part of the original material, but it represents another species, which is described below as Tambja abdere. Lance’s work with Nembrotha hubbsi was nearly con- current with that of Marcus & Marcus, although each was unaware of the other’s involvement. Figure 3 Gonads of Tambja eliora (Marcus & Marcus, 1967) 1 — capsule membrane 2 — penis 3 — atrium 4 — prostate 5 — hermaphrodite duct 6 — ampulla 7 — oviduct 8 — nidamental duct 9 — spermatocyst 10 — spermatotheca 11 — Sac 12 — Ampulla 13 — vaginal gland 15 — gonad in dorsal orientation Specimen distribution of Tambja eliora and the number of specimens include: CASIZ 003565 San Pedro Island, 32 km N of Guaymas, Mexico, in 24-27m of water. Leg. Bill Van Zandt, (2 specimens). USNM 709790 San Pedro Island, 32 km N of Guaymas, Mexico, in 24-27 m of water. Leg. Bill Van Zandt, (1 specimen). LACMNH 71-14 3-15m, east side of Punta Entrada at Sail Rock (N en- trance to Magdalena Bay), Baja California, Mexico (24° 32.4’ N, 112° 04’ W). Leg. James H. McLean, go- 31 January, 1971. (1 specimen). LACMNH 16904 La Paz area, Baja California, Mexico, Leg. Edwin Janss. May, 1972 (3 specimens). LACMNH A.g942 Near Escondido Bay, Carmen Island, Baja California, Mexico. Leg. Carl Gage and Chuck Fisher, March 28, 1974. (1 specimen), LACMNH A.9555 Isla Monserrate, Baja California, Mexico. Leg. Edwin Janss, April, 1974. (2 specimens). LACMNH A.9555 13.5 m, S end of Isla San Diego, Baja California, Mexico. Leg. Edwin Janss, April, 1974. (6 specimens and 1 Tambja abdere Farmer). SDMNH 63051 Isla Monserrate, Baja California, Mexico, April 13, 1969. Leg. Bill Van Zandt. (1 specimen). DMNH 92182 San Pedro Island, 32 km N of Guaymas, Mexico in 24-27 m of water, April, 1969. Leg. Bill Van Zandt. (1 specimen). BMNH 197936 San Pedro Island, 32km N of Guaymas, Mexico in 24-27m of water, April 13, 1969. Leg. Bill Van Zandt. WMF # 650. (1 specimen). WMF 651 San Pedro Island, g2km N of Guaymas, Mexico. 24-27m of water. Leg. Bill Van Zandt, April 13, 1969. (12 specimens). Vol. 20; No. 4 Tambja abdere Farmer, spec. nov. (Figures 4 - 6) Nembrotha eliora Marcus « Marcus, 1967: 195 (original “material two” only) Nembrotha sp. FarMER, 1970a: 16 Holotype: CASIZ No. 687. An egg mass is also included. Paratypes: USNM 709791; LACM 1717; SDSNH 63052; DMNH 92183; and BM[NH] 997838. Type Locality: La Paz, Baja California, Mexico. Ed- win Janss, Jr., coll. May 1972. Distribution: Punta Lobos, Guaymas, Danzante Island, San Diego Island, San Francisco Island, and La Paz, Mexico. Figure 4 Tambja abdere Farmer, spec. nov. Description: The 80mm long type specimen is limaci- form (Figure 4). The sole of the foot is brown and the foot margin is ochre. The general color is yellow ochre with 4 distinct areas of turquoise, 2 on each side and 2 dorsal. These areas extend the length of the body, and within them are irregular patches of yellow ochre. Be- tween the turquoise and yellow ochre is black. The head shield is horizontal across the front. Two flat cephalic flaps under the head shield of yellow ochre have turquoise between them. The blue-black rhinophores are retractile into highly set shields. The eye spots are prominent and blue-black. The gills (one unipinnate and 2 bipinnate) are dark and are set on a yellow ochre axis anterior to the anus. The radular formula, based on a minimum of prep- arations (Figure 5) is 174:'1-°1°1-4. The labial arma- ture is weak. Dorsal observation of the reproductive system shows surface areas and placement of the organs before they are teased apart. The penis is a narrow structure proxi- mally. Distally the spring-like prostate narrows to a very thin tube at its apex (Figure 6) and descends through the center of the coils, merging into an ampulla. The hermaphrodite duct leads into the vaginal gland by way THE VELIGER Page 377 of an enlarged, convoluted ampulla. A small round sper- matocyst is connected to the nidamental duct at the site of Figure 5 Radula of Tambja abdere Farmer, spec. nov. Figure 6 Gonads of Tambja abdere Farmer, spec. nov. 1 — capsule membrane 2 — penis 3 — vas deferens 4 — prostate 5 — ampulla 6 — hermaphrodite duct 7 - ampulla 8 — vaginal gland 9 — nidamental duct 10 — spermatocyst I1 — insemination duct 12 — spermatotheca 14 — aperture of outer oviduct 13 — vagina 15 — dorsal orientation of gonad Page 378 the insemination duct. The long insemination duct con- nects at a small bulbous portion of the bi-bulbar sperma- totheca. Also connected to this small bulbar area of the spermatotheca at the opposite side is the lengthy vagina. Behavior: Tambja abdere secretes great amounts of a substance from numerous goblet cells found in the skin. The reaction, triggered by the advances of Roboastra, is depicted in Figure 19. Discussion: This species represents the “number two” material of Marcus & Marcus, 1967. The authors did not illustrate the species, and it was presented only in the text. However, the text revealed the difference between the 2 materials. This difference was seen in Dr. Pickens’ photographs as well as in the text. Tambja eliora has blue- black rhinophoral sheaths and dark foot margins; T: ab- dere has yellow or yellow ochre sheaths and “bright yel- low” foot margins. Specimen distribution of Tambja abdere and the num- ber of specimens included are: CASIZ 687 La Paz area, Baja California, Mexico. Leg. Edwin Janss. Micro- scope slide of radula No. 497 specimen egg mass included. (1 specimen). USNM 709791 La Paz area, Baja California, Mexico, May, 1972. Leg. Edwin Janss. (1 specimen). LACM 1717 La Paz area, Baja California, Mexico, May, 1972. Leg. Edwin Janss. (1 specimen). LACM A.9555 Subtidal, Isla de San Francisco, Baja California, Mexico. Leg. Edwin Janss, February, 1974. (Specimen label incorrectly labeled Ro- boastra. 3 specimens). SDSNH 63052 La Paz area, Baja California, Mexico, May, 1972. Leg. Edwin Janss. (1 specimen). DMNH 92183 La Paz area, Baja California, Mexico, May, 1972. Leg. Edwin Janss. (1 specimen). BMNH 197438 San Pedro Island, 32km N of Guaymas, Mexico in 24-27m of water, April 13, 1969. Leg. Bill Van Zandt. (1 specimen). WMF 461 La Paz, Baja California, Mexico, 19 April, 1964. Leg. Richard Adcock. (1 specimen). Etymology: The specific name abdere, which is Latin and Greek, meaning “‘away from” and “skin,” is selected to call attention to the ability of the animal to produce copious amounts of defensive secretions. Tambja fusca Farmer, spec nov. (Figures 7 - 9) Holotype: CASIZ No. 688. It consists of a dissected ani- mal and a radula slide. Paratypes: USNM 709792; LACM 1719; SDSNH 63052; DMNH 92184; and BM[NH] 197435. Type Locality: Isla Monserrate, Baja California, Mexi- co; Edwin Janss, Jr. coll. April 1964. Distribution: Concepcion Bay, La Paz, and Cabo San Lucas, Baja California, Mexico. THE VELIGER Vol. 20; No. 4 Figure 7 Tambja fusca Farmer, spec nov. Description: The 40 - 50mm long animal is limaciform (Figure 7). The sole margin is ochre and is bordered by a black line. The general color is a dark yellow ochre with areas of turquoise of an irregular pattern. Between the ochre and turquoise is a thin line of black. The retrac- tile rhinophores are ruddy with dark tips. They have 19 leaves. One unipinnate and 2 bipinnate gills are set about half way on the back of the animal. The gills are dark tipped with slightly lighter areas attached to the ochre axis of the gill supports. The anal papilla is located just posterior of and next to the gills. The eye spots are dark in color, easily recognizable, and set posterior of and medi- al to the rhinophores. The radular formula, based on a minimum of prepara- tions, is 15 X4-3°1°1°1°3-4 (Figure 8). A thin labial armature Is present. Figure 8 Radula of Tambja fusca Farmer, spec. nov. Viewing the reproductive system dorsally before the capsule membrane is removed, many tubules and a wide surface of the vaginal gland are observed. Careful teasing of the system reveals the spermatotheca to be bi-bulbar (Figure 9) with a relatively short insemination duct. The spermatocyst is attached to the insemination duct, but not directly to the vaginal gland. The spring-like coils of the vas deferens and prostate surround the thin ascending Vol. 20; No. 4 4 Figure 9 Gonads of Tambja fusca Farmer, spec. nov. I — capsule membrane 4 — prostate 7 - ampulla 10 — spermatocyst 13 — vagina 3, — vas deferens 6 — hermaphrodite duct 8 — vaginal gland 9 — nidamental duct 11 — insemination duct 12 — spermatotheca 14 — aperture of outer oviduct 15 — gonad in dorsal orientation 2 — penis 5 — ampulla tube from an ampulla. The hermaphrodite duct leads into the vaginal gland through a thicker ampulla. Discussion: The lateral teeth variations are from 3 - 4 in Tambja fusca, and only 4 laterals in T: abdere. The first tooth appears to be more robust in T: fusca than in T: abdere. Specimen distribution of Tambja fusca and the number of specimens identified include: CASIZ 688 Isla Monserrate, Baja California, Mexico. Leg. Edwin Janss, April, 1974. Radula on microscope slide No. 498. (1 specimen). USNM 709792 Isla Monserrate, Baja California, Mexico, April, 1974. Leg. Edwin Janss. (1 specimen). LACM 1719 Isla Monserrate, Baja California, Mexico. April, 1974. Leg. Edwin Janss. (1 specimen). LACM A-9555 Gulf of California. Leg. Edwin Janss, 1972. (Not a Roboastra as per LACM label, 1 specimen). SDSNH 68053 Isla Monserrate, Baja California, Mexico. April, 1974. Leg. Edwin Janss. (1 specimen). THE VELIGER Page 379 DMNH 92184 Isla Monserrate, Baja California, Mexico. April, 1974. Leg. Edwin Janss. (1 specimen). BMNH 197435 La Paz area, Baja California, Mexico. April, 1972. Leg. Edwin Janas. (1 specimen). WMF 649 San Pedro Island, 3gkm N of Guaymas, Mexico, April 13, 1969. Leg. Bill Van Zandt. (4 specimens). Etymology: The specific name fusca, the feminine form of the Latin adjective fuscus (meaning dark, blackish) is selected to emphasize the relatively dark coloration of Tambja fusca as compared to the lighter color of T. abdere. Tambja mullineri Farmer, spec. nov. (Figures 10 - 12) Holotype: CASIZ No. 689, a dissected specimen and a microscope radula slide. Paratypes: LACM 1718; SDSNH 63052; DMNH 92185; BM[NH] 197437. Type Locality: Academy Bay, Santa Cruz Island, Gala- pagos Islands, Ecuador. David Mulliner and Gale G. Sphon, coll. 18 March, 1971. Distribution: James Island; Jerris Island; Academy Bay, Santa Cruz Island; NW Onslow Island, and Flore- ana, Galapagos Islands. Figure 10 Tambja mullineri Farmer, spec. nov. Description: The 15mm long animal is limaciform (Figure 10). The foot of the living animal is like the general background color of ochre-turquoise grading into a lighter shade. Broad turquoise stripes are bordered by a thin line of navy blue. The colors are noted from a photo- graph by David K. Mulliner. The one unpinnate and 2 bipinnate gills are navy blue with a light ochre-turquoise trunk supporting the gill fil- aments. A wide turquoise stripe extends from the gill, Page 380 THE VELIGER Vol. 20; No. 4 anteriorly between the rhinophores, to the cephalic rim. How the stripe ends anteriorly is not known since it has not been observed in relaxed living material or photo- Figure 11 Radula of Tambja mullineri Farmer, spec. nov. Figure 12 Gonads of Tambja mullineri Farmer, spec. nov. 1 — capsule membrane 2 - penis 3 - atrium 4 — prostate 5 — hermaphrodite duct 6 — ampulla 7 — spermatocyst 8 — spermatotheca 9g — vaginal gland 10 — nidamental duct 11 — ampulla 12 - vagina 13 — gonad in dorsal orientation graphed from the front. Two dark areas are behind the rhinophores adjacent to the medial stripe. The rhinophores are navy-blue with ochre-turquoise leaves. The rhinophoral sheaths are thick and stout in appearance. The blunt navy-blue tipped rhinophores have 12 or 13 leaves; they are retractile. The eye spots are moderately dark. The reproductive system, viewed dorsally with the cap- sule membrane intact, shows a large atrium extending to the far side. At this point, the prostate arises with a loop descending into the inner part of the mass. The hermaph- rodite duct enters the mass posteriorly about 4 the distance from the inside of the snail. Teasing apart of the repro- ductive system (Figure 12) reveals a large atrium and coiled prostate of uniform diameter connected to an am- pulla. This ampulla rests on and is connected to the vagi- nal gland. The hermaphrodite duct connects to a large ampulla. A smail round spermatotheca of equal size and shape and the spermatocyst join to this ampulla by a common duct. This ampulla’s connection to the rest of the system has yet to be uncovered. The radular formula (Figure 11) from a few prepara- tions is 15X4°1-°1:1°4. The labial armature is moder- ately stout. Discussion: Tambja mullineri is similar to T. eliora in appearance. The frontal veil suggests a uniform sweep across the animal, unlike that of T: eliora which dips down the front of the head. The blue stripes are wider in T. mullineri than in T. eliora. The first lateral tooth in T. mullineri has a hook, but it is absent in T: eliora. The distribution of specimens of Tambja mullineri and the number of specimens identified include: CASIZ 689 Academy Bay, Santa Cruz Island, Galapagos Islands, Ecuador, on a reef in 22-30m of water. Leg. Los Angeles County Museum of Natural History # 18171, August 26, 1974. Radula on microscope slide, No. 499 (1 specimen). LACM 1718 James Bay, James Island, Galapagos Island, Galapagos Islands, March, 1971. Leg. Los Angeles County Museum of Natural History. (1 specimen). LACM 1718 Jerris Island, Galapagos Islands, March, 1971. Leg. Los Angeles County Museum, David K. Mulliner and Gale Sphon. (1 specimen), LACM 16905 On nullipores, North West of Isla Onslow, Floreana Island, Galapagos Islands, Ecuador, in 20m. Leg. Gerard M. Wellington, January 13, 1975. Dissected gonad. (2 specimens 13 mm long each). SDSNH 63056 Jerris Island, Galapagos Islands, March, 1971, in 10.5m of water. (1 specimen). BMNH 197437 Academy Bay, Santa Cruz Island, Galdpagos Islands, Ecuador, in 22 — go m of water on reef, March 18, 1971. Leg. Los Angeles County Museum of Natural History. (1 specimen). Etymology: Tambja mullineri is named for David K. Mulliner, a member of the Ameripagos Expedition, whose photographs have been useful in this study. Vol. 20; No. 4 Roboustra Bergh, 1877 Type species: Roboastra gracilis Bergh, 1877: 458; plt. 56; fig. 12 “Oral tentacles are strongly developed as dorsolater- ally grooved cylindrical projections equal in length to the rhinophores. Rachidian teeth are reduced to three prongs of well defined denticles. The lateral tooth is deeply bifid. Marginal plates are reduced and linear, 1 - 3 in number. The buccal collar is very weak. A labial armature is ab- sent. A small prostate gland is confined to a coiled glandu- lar section of the vas deferens.” (BURN, 1967: 215). Roboastra tigris Farmer, spec. nov. (Figures 13 - 19) Roboastra sp. FARMER, 1970a: 16 Holotype: CASIZ No. 690; it is entire. A microscope slide of the radula from another animal is with the speci- men. Paratypes: CASIZ No. 691; LACM 1720; SDSNH 63054; DMNH 92186; and BM[NH] 1974309. Type Locality: San Pedro Island, 32km N of Guaymas, Sonora, Mexico, in 24 - 27m. Distribution: Los Angeles Bay, Guaymas, Monserrate Island, San Diego Island, and La Paz, Mexico. Figure 13 Roboastra tigris Farmer, spec. nov. Description: The 200 to 300mm long animal is limaci- form (Figure 13). The sole is blue-black; the foot margin is blue. The basic color is yellow ochre with 5 black stripes bordered by bluish green. The sides may have an interrup- ted stripe. The retractile rhinophores are blue-black, sur- rounded by a rhinophore sheath, also blue-black. The THE VELIGER Page 381 cephalic tentacles are placed anterior laterally and folded with the rib of the fold downward. The eye spots are generally bluish in color and well defined. The anterior cephalic shield is marked by a slight ridge. The radular formula, based on a few microscope slide preparations, is 35X4-3°1°1°1'3-4. The radula is typically that of a Roboastra. The rachidian tooth has lateral prongs. The lateral teeth vary in number from 3 or 4 and are without prongs (Figure 14). Lance (1968) ols Figure 14 Radula of Roboastra tigris Farmer, spec. nov. figures Roboastra radula for Nembrotha hubbsi or Tamb- ja (Nembrotha) eliora. One should also keep in mind that there is another Roboastra species living in the Gulf of California. The oral area is cobalt blue and everts to surround its prey. Two cephalic tentacles are internal, proximal to the mouth. The radula is distal to the mouth in the buccal cavity (Figure 17). The body is primarily a hydrocoel, which plays an im- portant role in the swift eversion of the buccal envelope used to capture prey (Figures 18, 19). The digestive system is a centrally located tube in the coelomic cavity. There is a digestive gland at the junction of the esophagus and stomach. The intestine has its in- sertion antero-dorsally to the stomach. The cephalic ganglia produce 4 neurons of large dia- meter involving the external cephalic tentacles and the rhinophores. The genital pores are located on the right side and are cobalt blue. Viewing the encapsulated reproductive sys- tem from the dorsal side, the hermaphrodite duct and am- pulla overlying the vaginal gland can be seen. The round spermatotheca, vas deferens and prostate are also seen at the surface. The teased reproductive system (Figure 15) shows a uniform diameter vas deferens and slightly larger diameter prostate connected to a thin tube, the oviduct. The ampulla from the hermaphrodite duct is also con- nected to the oviduct. The elongate spermatocyst and round spermatotheca are situated over the large vaginal gland. An ampulla is attached to a Y-shaped tube leading Page 382 THE VELIGER Vol. 20; No. 4 Figure 15 Gonads of Roboastra tigris Farmer, spec. nov. 1 — capsule membrane 4 — prostate 7 -— ampulla 10 — ampulla 3 — vas deferens 5 — spermatotheca 6 — spermatocyst 8 — hermaphrodite duct 9 — vaginal gland 11 — oviduct 12 — gonad in dorsal orientation 2 — penis from the spermatocyst and spermatotheca. The penis is armed with hooks (Figure 16). The 5 unipinnate gills are blue-black; the anus is centrally located. An underwater photograph by Boris Innocenti clearly indicates that Roboastra tigris attains a length of 300mm. Figure 16 Penial hooks of Roboastra tigris Farmer, spec. nov. The large size of this predatory slug might be linked to its feeding mechanism. A slug of such large size has mostly fluid within the coelomic cavity. This reservoir of fluid can be used to operate the rapid eversion of the buccal envelope to capture the rapidly swimming Tambja eliora. Figure 17 Roboastra tigris Farmer, spec. nov. Illustrations of moribund specimen (a, e, f) h — oral tentacle, swellings indicate mode of peristalsis along the length of the oral tentacle m — mouth ot — oral tentacle be — buccal envelope r — radula In the type species description no mention is made of the feeding apparatus seen in Roboastra tigris. The oral tentacles are strongly developed as dorso-laterally grooved cylindrical projections. Roboastra tigris has these projec- tions; however, they cannot be called oral tentacles; they might rightly be called cephalic projections since the oral tentacles are inside the buccal area behind and above the Vol. 20; No. 4 THE VELIGER Page 383 mouth. The oral tentacles are not merely static organs but undergo peristalsis as illustrated in Figure 17h. Figure 18 Sketches from selected motion picture frames of Roboastra tigris capturing Tambja eliora Behavior: Roboastra tigris is photographically recorded as attempting to feed on a large specimen of Tambja ab- dere (Figure 19). Motion pictures of one feeding on T. eliora are partially illustrated here. It is the first known nudibranch as a predator of another nudibranch. It is known to live in a rocky environment and to attain a size of 300mm. Chelidonura inermis (Cooper, 1862), a cephalaspidian, also preys upon sea slugs. Figure 19 Roboastra tigris attempting to capture Tambja abdere A color photograph illustrating the unusual feeding be- havior of Roboastra tigris is reproduced in Sea Frontiers as Nembrotha sp. The distribution of specimens of Roboastra tigris and the number of specimens identified are as follows: CASIZ 690 San Pedro Island, 32km N of Guaymas, Mexico in 24-27m of water. April 13, 1969. Leg. Bill Van Zandt. WMF 650. (1 specimen). CASIZ 691 La Paz, Baja California, Mexico, 1964. Leg. Dick Adcock. WMF 392. A specimen and microscope slide of radula No. 500. USNM 708793 La Paz, Baja California, Mexico. May, 1972. Leg. Edwin Janss. (1 specimen). LACM 1720 San Pedro Island, 32 km N of Guaymas, Mexico in 24-27 m of water, April 13, 1969. Leg. Bill Van Zandt. WMF 650. (1 specimen). LACM 16906 La Paz, Baja California, Mexico. Leg. Edwin Janss. April, 1972. (1 specimen). LACM A.9555 S End of Isla San Diego, Baja California, Mexico, in 13.5 m. Leg. Edwin Janss, April, 1974. LACM A.9555 Isla Monserrate, Baja California, Mexico. Leg. Edwin Janss, April, 1974. (1 specimen). SDSNH 63054 San Pedro Island, 32km N of Guaymas, Mexico, in 24-27m of water, April 13, 1969. Leg. Bill Van Zandt. WMF 650. (1 specimen). DMNH 92186 La Paz area, Baja California, Mexico. May, 1972. Leg. Edwin Janss. (1 specimen). BMNH 197439 San Pedro Island, 32 km N of Guaymas, Mexico in 24-27m of water, April 13, 1969. Leg. Bill Van Zandt. WMF 650. (1 specimen). WMF goz2 La Paz, Baja California, Mexico. 1964. Leg. Richard Adcock. (1 specimen). WMF 650 San Pedro Island, 32km N of Guaymas, Mexico. 24-27m of water. April 13, 1969. Leg. Bill Van Zandt. (1 specimen). Etymology: Roboastra tigris is so named for its stripes and carnivorous appetite similar to the tiger. Page 384 THE VELIGER Vol. 20; No. 4 Roboastra sp. (Figure 20) Edwin Janss snapped a photograph of Roboastra sp. near La Paz in June 1972 (Figure 20). It shows an animal similar to Roboastra tigris, but differing in detail of color and patterns, suggesting that another species of Robo- astra may be present in the Gulf of California. Unfortu- nately, the specimen was devoured by one of its Robo- astra companions in transit from the field to the labora- tory. A strong resemblance in dorsally grooved oral ten- tacles (or cephalic tentacles) is noted between the “new” species and Roboastra. However, it differs by having orange-tipped rhinophores, white gills, and many more stripes on the body. Figure 20 Roboastra sp. Color Notes: The limaciform animal is about 60mm long. The edge of the foot is blue-black; the body is marked with numerous stripes of yellow ochre and bluish- black. Some of the stripes are lighter in shade. The eye spots are white. The cephalic basal folds are blue-black. The base of the rhinophores is blue-black with a similar core; the leaves are bright orange. The rhinophores are tipped in blue-black; the gills are blue-black distally and white proximally. There are 5 unipinnate gills. The mouth area could not be seen in the photographs. Abbreviations used are: BM[NH] - British Museum [Natural History] CASIZ — California Academy of Sciences, Invertebrate Zoology DMNH - Delaware Museum of Natural History LACM -— Los Angeles County Museum of Natural His- tory : SDSNH - San Diego Society of Natural History USNM - United States National Museum WMF - Wesley M. Farmer collection. Key to the Eastern Pacific Species of Roboastra and Tambja 1 Mouth anterior and centrally located in the head ........emnm 2 — Mouth located antero-ventrally near the f00t ...msmmnnsmnennen 3 2 Rhinophores blue-black, black eae on body few, bordered by light blue .. is Roboastra tigris — Rhinophores zai, pees on n body numerous . . Roboastra sp. 3 Frontal veil horizontal ou... dicate 4 — Frontal veil dips into Hy” ¢ or “eu shane wie ates Tea dieaa 4 Turquoise blue areas with “islands” of yellow OCHIE ......-csese 5 — Turquoise blue stripes with “islands” of yellow ochre; both stripe colors of equal Width .....ccnnne Lambja mullinen Yellow ochre dark, turquoise blue areas with “islands” of yel- low ochre appearing “irregular” with variable interface be- tween ochre and turquoise .......-ccseenene . Tambja abdere — Yellow ochre light, turquoise blue areas with ‘ “islands” of yel- low ochre appearing “organized” with uniform appearing in- terface between ochre and turquoise ......ccnessn Tambja fusca o ACKNOWLEDGMENTS The author appreciates the help and support of James Lance for his contributions of literature; and Bill Van Zandt and Boris Innocenti for initial collections of 3 species of sea slugs and photographs. David K. Mulliner and Edwin Janss are responsible for the fine photographs of sea slugs, specimens, and data. Robert Burn and Eveline Marcus offered many excellent comments on procedure. Type material was sent for examination by Walter J. Byas and Clyde FE E. Roper of the Smithsonian Institution; the late Allyn G. Smith of the California Academy of Scien- ces; and the Santa Barbara Museum of Natural History. The manuscript was read by several friends who wish to remain anonymous; they offered editorial suggestions for the manuscript. The combined efforts of these people helped to bring this work to completion. The author described the species; his wife, Michaelene Farmer, worked with the historical aspects of the manu- script. Literature Cited Axssott, RoBpert TuCKER 1974. American seashells Van Nostrand Reinhold Co., New York, 2nd ed.; 663 pp.; 24 color plts.; numerous figs. Baszepow, HerBerT & CHartes HEDLEY 1905, South Australian nudibranchs, and an enumeration of the known Australian species. Trans. Roy. Soc. So. Austral. 29: 146 - 159; plt. 2 Vol. 20; No. 4 THE VELIGER Berce, Lupwio SopHus RupoLr 1877. Malacologische Untersuchungen. In: C. Semper, Reisen im Archipel der Philippinen. Wissensch. Result. 2 (11): 429-494; plts. 54-57 Burn, Rosert 1962. Descriptions of Victorian nudibranchiate Mollusca, with a com- prehensive review of the Eolidacea. Mem. Nation. Mus. Melbourne 25: 95-128; text figs. 1 - 26 (May 1962) 1967. Notes on an overlooked nudibranch genus, Roboastra Bergh, 1877, and two allied genera (Mollusca : Gastropoda). Austral. Zool. 14 (2): 212-221; figs. 1-6 Cooper, James GraHAM 1862. Some genera and species of California Mollusca. Proc. Calif. Acad. Nat. Sci. 2: 202 - 207 FaRMER, WESLEY MERRILL 1968. Tidepool animals from the Gulf of California. Wesword Co., Tempe, Arizona, illust. 1970a. A swimmer, the secretor and the Blue Tiger (three nudibranchs from the Gulf of California). The Echo 2: 16 1970b. Swimming gastropods (Opisthobranchia and Prosobranchia). The Veliger 13 (1): 73 - 89; 20 text figs. (1 July 1970) 1970c. Skin secretions in three species of opisthobranchs and one pul- monate from the Gulf of California (Mollusca: Gastropoda). Master of Sci. thesis, Ariz. State Univ. Keen, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. Stanford Univ. Press, Stanford, Calif i- xiv+ 1066 pp.; ca. 4000 figs.; 22 color plts. (1 September 1971) KerstTitcH, ALEX 1977. Photograph in Sea Frontiers 2g (6) (Nov. - Dec. 1977) Lance, JAMes RoBERT 1968. New Panamic nudibranchs (Gastropoda: Opisthobranchia) from the Gulf of California. Trans. San Diego Soc. Nat. Hist. 15 (2): 3-13; 2 plts.; 11 figs. (8 January 1968) Maacus, Eveing pu Bois-Reymonp # Eranst Marcus 1967. American opisthobranch mollusks. Stud. Trop. Oceanogr. Miami, 6: vilit+256 pp.; figs. 1 - 155+1-95 (December 1967) SkocLunp, Carot CHRISTINE 1974. Intertidal marine Mollusca of Cape Tepoca, Sonora, Mexico. Master Sci. thesis, Ariz. State Univ. Taytor, DwicHT Witiarp e Norman F. Soni 1962. An outline of gastropod classification. Malacologia 1 (1): 7-32; 1 text fig. (14 November 1962) Page 385 Page 386 THE VELIGER Vol. 20; No. 4 METHODS & TECHNIQUES Air Drying Giemsa Technique for Gastropod Chromosomes BY R. PRASAD anp C. C. DAS P. G. Department of Zoology, Berhampur University, Berhampur - 7, Orissa, India (1 Plate) INTRODUCTION PERHAPS NOT MORE than 0.5% of the known species of molluscs are known karyologically. Again, most of the chromosomal studies in this group have been undertaken rather recently (PATTERSON, 1971). Unfortunately, the available information on the morphology of somatic metaphase chromosomes and behaviour of germinal chro- mosomes during meiosis lacks details in many of the in- vestigated species and thus needs reinvestigation. Recently several new techniques for preparations of meiotic and mitotic chromosomes of Mollusca have been described (Burcu, 1968; PaTTERSON, op. cit.; STERN, 1975a), but most of these consume much time and labour. The present authors wish to describe here a technique for gastropod chromosomes which is quite simple and takes much less time. MATERIALS anp METHODS Ovo-testes of a land snail, Euplecta subdecussata (Pfeif- fer), a pulmonate, constitute the material for the present study. The specimens received an injection of 0.1 mL of 0.025% colchicine solution directly through the apex of the shell. Two hours after the injection the shell was removed, the ovo-testes were taken out and placed in distilled water at room temperature for 45 minutes. Incu- bation for less than 45 minutes yielded poor spreading of chromosomes. These ovo-testes were minced and flushed in a 0.9% sodium citrate solution to obtain a thin emul- sion which was centrifuged for 5 - 10 minutes. The tissue precipitate was fixed in freshly prepared aceto-methanol mixture (1 : 3). The latter was thoroughly flushed and after 10 minutes the fixative was replaced with fresh fixa- tive. After 45 minutes of final fixation the suspended material was spread on chilled slides (stored in 50% al- cohol), which were exposed to flame and then air dried. On being dried for 24 hours or more the slides were stained with Giemsa stain (stock solution diluted 1 : 10) for 1 hour, the excess of stain being removed by washing in tap water. The dried slides were then ready for micro- scopic examination. DISCUSSION The aceto-orcein squash technique used by STERN (1975a, 1975b) for the study of chromosomes from gonads and embryonic tissue of molluscs was undoubtedly an improve- ment over the techniques for the chromosome cytology of Mollusca previously employed by Hustep «& BurcH (1946) since the latter required an elaborate fixing and sectioning procedure. And yet this technique failed to re- solve the precise morphology, chiasma frequency, etc. of chromosomes. Sometime earlier, BuRcH (1968) has described a tissue culture technique for karyotype analysis of pulmonate land snails. However, as noted by PATTERSON (1971), tissue culture is not only time-consuming but is very ex- pensive and requires sterile conditions. The present authors, using the air-drying technique, ob- tained excellent preparations of meiotic chromosomes. The chromosomes were very well spread and thus ren- dered the study of their morphology, chiasma frequency, etc., extremely easy. The diploid number of Euplecta sub- decussata, as revealed by the gonial metaphases, was found to be 54 (Figure 1). The chromosomes were rod- Explanation of Figures 1 to 3 Chromosomes of Euplecta decussata (Pfeiffer) Figure 1: Gonial metaphase Figure 2: Pachytene stage Figure 3: Diakinesis stage THE VELIGER, Vol. 20, No. 4 [Prasap « Das] Figures 7 to 3 Figure 3 Chromosomes of Euplecta decussata (Pfeiffer) Figure 1: Gonial metaphase Figure 2: Pachytene stage Figure 3: Diakinesis stage Vol. 20; No. 4 shaped and none of them could be considered a “marker chromosome” in the complement. The early stages of mei- osis represent a growth phase (Figure 2) and the haploid number of 27 bivalents could be confirmed from diakine- sis and subsequent stages (Figure 3). Most of the bivalents at diakinesis stage contained a single interstitial chiasma each and had undergone extreme condensation and ter- minalization at metaphase I. A heteromorphic or hetero- pycnotic sex-element could not be marked in any of the stages. ACKNOWLEDGMENTS The authors wish to thank the Director, Zoological Survey of India, Calcutta, for his kind help in the identification of the specimens. Literature Cited BurcH, Joun Bayazp 1968. A tissue culture technique for caryotype analyses of pulmonate land snails, § The Venus 27: 20 - 27; 6 figs. (August 1968) Hustep, Laptey a Paut RaNpoLtpH BurcH 1946. The chromosomes of polygyrid snails. 410 - 429; 34 figs. PATTERSON, CHARLOTTE MORGAN 1971. A karyotype technique using freshwater snail embryos. Rey. 4: 27; 1 fig. Stzrn, EDWARD Marc 1975a.__A technique for the preparation of gastropod chromosomes. The Veliger 17 (3): 296-298; 2 text figs. (1 January 1975) 1975b. The chromosomes of Viviparus subpurpureus (Say). Mala- col. Rev. 8: 107 - 108; 5 text figs. Amer. Natural. 80: Malaool. INFORMATION DESK What’s the Difference? The Role of the Editorial Referee BY A. MYRA KEEN 2241 Hanover Street, Palo Alto, California 94306 WE WHO SET OUT to publish our work are rarely aware of the number of people who will have handled the script THE VELIGER Page 387 before it appears in print. We negotiate with the editor, but there are many back-stage workers, such as typeset- ters, proof-readers, layout designers, and printers. Also there are the referees who form the most faceless category of all, and to whom the editor soon relays the paper. (I use the term “referee” in the sense of “consultant,” not in the sports sense of “umpire”; an alternative term, “re- viewer,” I prefer to reserve for one who writes a critique after publication, not before.) My purpose here is to give young authors some tips on how the process of pub- lication can be expedited, because understanding the tasks of the editor and the referees may enable the writer to forestall much of their criticism, thus saving both time and the fraying of nerves. The referee is usually someone who has special know- ledge of the field of the paper and who thus can advise the editor on its merits as well as its faults, catch inaccura- cies of text or outright errors, point out inadequate cover- age of literature, and in general provide an impartial judgment of value as a contribution to science. The editor, of course, has the responsibility of seeing that the paper then is put into optimum condition before it goes to press. With what finesse he can muster, he relays the referee’s criticisms (if any) to the author and works with the author in getting the paper into acceptable form. Somewhere along the line a decision must come from among four alternatives: to accept the paper in its submitted form, to take on the task of correcting minor errors, to send it back to the author for further work, or to reject it entirely. No one feels easy about making such decisions alone, so that here the referee can be of service in reinforcing the editorial judgment. Often more than one referee exam- ines the paper. This is a way modern journals have for maintaining publication standards. Because of the frank appraisals the referees must make, they normally are happy to remain nameless and unacknowledged, and the breaching of their protective anonymity can lead to no little embarrassment. Many editors, having had experience with typescripts of less-than-polished quality, supply their referees with a list of suggestions for evaluating papers, designed to help watch for common faults. One such list starts with the title (is it clear, concise, informative?) ; then it asks about the general organization (is it logical, with well- chosen subheads?) ; the text itself (is the sentence struc- ture free of ambiguities, vagueness, indefinite references, superfluous words and phrases?) ; the illustrative matter (are tables consistent and essential?) ; literature cited (is the reference list adequate but not over-long, and is it properly cued in the text?) ; figures (are they essential, Page 388 of a quality to reproduce well, and are the captions clear?) ; footnotes (are they properly indicated in the text, and are they appropriately placed?) ; lastly, the abstract, if there is one (is it a concise summary?). How does the referee work? Well, hope greets each new script — maybe this will be the near-perfect one. It’s a good omen if the word “Acknowledgments” is correct- ly spelled. Next comes a glance to see whether words and phrases like those in Table 1 are sprinkled too liberally through the first pages (if so, that’s a bad omen). Most of these terms are correct if rightly used, but unwary Table 1 Words and phrases that frequently occur in poor writing as to in case of so that but which instance standpoint different than interesting the fact that etc. located very found to be often was found writers overwork them to the point of obscuring meaning. The referee becomes hypercritical when too many of them interfere with the author’s line of thought, and reading becomes a struggle through the verbal shrubbery. Next, the referee begins to watch for some of the common terms that authors frequently use incorrectly, Table 2 Words and phrases frequently misused appears is not an exact synonym of seem and should be reserved for the meaning, “becomes visible” around is not a synonym of about curious is only in part a synonym of strange or unusual due to is not an adequate substitute for because of presently refers to the future, not to now quite means entirely or completely; it is not a comparative term which and that are not interchangeable (“which is non- defining and should be used only after a comma” [see Fow Er (1926) }) while and since are time terms, inappropriate as synonyms of whereas and although THE VELIGER Vol. 20; No. 4 such as those listed in Table 2. Misuse of any of these signals a lack of sensitivity to meanings. Some phrases even betray a failure to think clearly. For example, can anything be quzte unique? Or an idea be utterly basic or have first priority? Are there any other than true facts? Yet, authors have committed these and similar infelicities in manuscripts ! Matters of syntax: should not have to concern the ref- eree, but they do. The dangling participle, the unclear antecedent, the indefinite “it,” and erratic shifts in tense or number all may slide in and make for hard reading. Such redundancies as, “measuring xmm in length” are a distraction. Logical defects may trip up the writer whose mind is more on content than on form. KEsLine (1958) quotes several examples of fuzzy writing: “The ends of the spines are 2mm. long.” “The average length is about 25mm. but it varies greatly.” “Certain crabs spend all their lives in empty gastropod shells.” These, then, are some of the pitfalls that the referee hopes the author will have avoided, so that there would be no need to alert the editor about awkward usages. Malacologists (and for that matter biologists in gen- eral) have some peculiarities of style that need examina- tion. Here is a gem culled from a manual on invertebrates that manages five faults in three short sentences [emphasis and commentary are mine]: “A large slender species ! found? in mud at low water. It 3 sometimes grows to nearly two feet in length. Only the curious* head is shown, which bears > four short curved hooks. ...” Commentary: !) A species is a concept, drawn up from the study of specimens. Only the specimens, then, really can have the morphological characteristics of size, shape, color, texture, etc.; the species itself has only relatedness to other forms and distribution in time and space. 2) Here the animal described occurs in mud; finding it has nothing to do with its ecological niche. 3) “Jt” is an indefinite reference word, presumably applying to “‘species,” but if so, the species cannot grow in any such fashion; the animal does the growing. *) Whether the animal was curious (2. é., inquisitive) or not we really cannot judge; the illustration of it shows a bulbous head studded near the end with four recurved hooks. 5) Bear is a word that FowLer (1926) classes as formal or pretentious; its syn- onym carry is preferable. Formal words tend to be poly- syllabic and much longer than their plainer synonyms; here, though, the formal word is actually the shorter. An- other formal word that appeals to many malacologists is possess. A clam, as I used to point out to my students, may have siphons, but — I asked — does it really “possess,” 2. é€., “own” them? Vol. 20; No. 4 A confusion between verb forms and adjectives creeps into many a species description with the use of such terms “imbricated,” “angulated,” and the like. The -ed ending implies a process. Thus “elongated” may be justifiable, for growth is a process that can increase length. Mainly, however, the use of these terms implies the adjectival or descriptive aspect — elongate, angulate, imbricate, etc. The word “intertidal” often appears as a noun, which grates on the sensitive ear; one should say either “inter- tidal zone” or “‘intertidally” to be etymologically correct. Another word often misused in descriptions is promi- nent, which may indeed mean evident or conspicuous. However, it really denotes something that juts out or is elevated. An umbilicus may be well-marked, but it can hardly be prominent. Paleontologists and stratigraphers are prone to err in use of the word interval when they imply it to be the actual rock stratum — a “sandstone interval.” An inter- val is a hiatus or gap, not a physical entity. Telegraphic style is almost universally adopted in writing of species descriptions. Achieving consistency in its use is not always easy, however. I have discussed this in a recent note (KEEN, 1977). How, then, should the aspiring author go about prep- aration of a paper? The first need is a good outline, to ar- range the ideas in a logical sequence, to clarify the scope, and to put the study in its proper setting. Before starting the actual composition one should take time to read or re-read any of several available style manuals (a few are listed in Table 3) or some of the useful short papers cited under “References.” That is, one should give oneself a refresher course on good writing, after which it is only human to feel a creeping paralysis, what with all the do’s and don’ts that must be remembered, like the victim in this bit of verse (author unknown) : A centipede was happy quite Until a frog in fun Said, “Pray, which leg goes after which?” This raised her mind to such a pitch She lay distracted in the ditch, Considering how to run! According to a Chinese proverb, the longest journey begins with a single step. Therefore, the wise course at this point is to exert some will-power, to sit down, and to start writing. Let the words flow — visualizing it as a conversation with the prospective reader — until the first draft has been knocked into a semblance of shape. Scan it and make whatever corrections seem immediately needed, based on the previous reading. For the next step, THE VELIGER Page 389 Table 3 Style Manuals Geowrtting, a Guide to Writing, Editing, and Printing in Earth Science. American Geological Inst., Falls Church, VA. 24 ed., 1974. 80 pp. Manual of Style for Authors, Editors, and Copywriters. Univ. Chicago Press. 12" ed., 1969. 546 pp. Style Manual for Biological Journals. American Inst. Biological Sciences, Washington, D. C. 3" ed. 1972. 97 pp. two courses are open, and it doesn’t matter which comes first: either find someone who will give the work a critical reading and point out weaknesses; or put it aside for a while before the critic sees it. McCarTNEY (1954) tells us that the Roman teacher Quintilian advised his stu- dents to let a manuscript rest until it would seem, upon re-reading, the work of another. Our modern pace rarely permits such refrigeration, but a pause of a few days or weeks can give enough perspective to enable a more criti- cal review of one’s own work, which then should be read for order, for clarity of presentation, for correctness of wording and spelling, and for consistency between parts. Reading it aloud may help, especially if one has a friend do the reading and can listen, as the French zoologist Buffon did — for passages where the reader halts or fails to catch the meaning at once. Buffon revised or rewrote these. Soliciting criticism before publication is wise in- surance against adverse reviews that could come later. Having taken these precautions the author makes the paper really ready to go to the editor. Lest anyone reading this feels that I am more prone to give advice than to act upon it, I would point out that this entire paper has been framed with almost no revert- ing to use of the passive voice or to those words “perhaps” and “apparently” that have hitherto been a part of my writing kit. The passive voice is a useful device, at times, especially for avoiding the forthright “I” or the wordy “the writer,” and “perhaps” may be a way of softening didactic statements. However, while doing for myself the advance reading that I had recommended, I noticed such frequent injunctions against these mannerisms — for example, StrucK (1954) says that “every passive should be considered sick until proved healthy” — that I resolved to write one paper without using them. I found it possible to change my habits, but not easy. In fact, I had to omit one sentence: in introducing the gem that manages five faults in three sentences I wanted to recount the tale of Page 390 THE VELIGER Vol. 20; No. 4 the politician who is said to have established an all-time record of three errors in only two words: “’Them’s them.” But I could see no way to do it, because I could not positively document the story. I also had the intention, when I sat down to write, to stress the modern lack of drill in school training (what child nowadays learns to parse or diagram a sentence?). Those who are not taught to distinguish between counter- feit and genuine verbal coins, who find half-a-dozen repe- titions of “you know” quite acceptable in a spoken sen- tence, can hardly be expected to be skillful at writing. But then, in my reading, I found that Lane (1935) was quoting an editor who as long ago as 1915 deplored the then poor quality of English training in the schools! It may be that anyone, with determination, can learn to write; but writing, like any other craft, requires practice. That practice must be coupled with a constant awareness: we learn only when we begin to notice our own mistakes. ACKNOWLEDGMENTS I am grateful to Sandy Gardner and Eugene Coan for having read and made constructive criticisms of the first draft of this paper. Their suggestions led to improvements in the final draft. Suggested References Fow er, H. W. 1926. A dictionary of modern English usage. Revised, 1965, by Sir Ernest Gowers. 725 pp. Keen, A. Myra 1977. What’s the difference? Telegraphic style versus normal style. The Veliger 20 (2): 187 (1 October 1977) Kesuina, R. V. Oxford Univ. Press. 1958. Crimes in scientific writing. Turtox News 36(12): 274-276 (Reprinted, 1973, by Mus. Paleontol., Univ. Michigan, Ann Arbor, Mich. ) Lang, B. H. Suggestions to authors of papers submitted for publication by the United States Geological Survey. U.S. Govt. Printing Office, Washington, D. C., 4th ed., 126 pp. (later revised but now out of print; available in libraries) McCartney, E. S. 1954. Does writing make an exact man? Science 119 (3095): 525 - 528 (23 April 1954) Struck, H. R. 1954. Recommended diet for padded writing. Science 119 (3095) : 522 - 525 (23 April 1954) NOTES & NEWS Soviet Contributions to Malacology in 1976 BY MORRIS K. JACOBSON AND KENNETH J. BOSS Museum of Comparative Zoology, Harvard University Cambridge, Massachusetts 02138 WE ONCE AGAIN provide a listing of malacological publi- cations which appeared in the Soviet Union and were abstracted in the Referativnyy Zhurnal during the year 1976. This annual resumé has appeared previously (see The Veliger 19 (4): 440 for last year’s report and refer- ence to earlier ones). The editors of the Referativnyy Zhurnal have altered the arrangement of the categories in their format, and we have followed their procedures herein. Published malacological research in the Soviet Union shows no marked departure in quantity or quality during 1976. Several important papers appeared concerning the phylogeny, and thus, the evolutionary biology and system- atics of the Mollusca during the last year. Most note- worthy is probably Shileiko’s study of the excretory appa- ratus of the stylommatophoran Pulmonata. Since the higher categories in the taxonomy of this group are defined largely in reference to the configuration of visceral organs, especially the ureter, any evidence which questions the stability or phyletic importance of these structures demands attention. Shileiko points out that the sigm- urethrous condition found among the higher pulmonates may have been independently derived more than once from the more primitive, simple orthurethrous stocks. Minichev and Starobogatov propose a rather question- able system for the phylogeny of the Mollusca, deriving the phylum from Dinophilus-like annelids; such a scheme seems somewhat dated when very strong evidence indi- cates direct molluscan-turbellarian affinities. Staroboga- tov also wrote a review of the phyletic relationship among the Lymnaeidae while Zuev and his colleagues discussed the systematics of a number of cephalopod genera. Sirenko Vol. 20; No. 4 THE VELIGER Page 391 also provided some phyletic re-arrangements within the Polyplacophora. Several new taxa were introduced and many of these in turn are doubtful, being established not only on typo- logical criteria but rather weak biological evidence. Sirenko introduced a new subfamily, several new genera and a number of new species of chitons. Among terrestrial pulmonates, Sklyar described a new Deroceras from the Crimea and Shileiko established a couple of new sub- genera and several new species of lauriine pupillids as well as two further new subgenera of orculids. Krivosheina added 5 new species of Euglesa (Sphaeriidae) from mon- tane lakes in the Altai while Timms studied and ‘revised’ the finger-nail clams of an Estonian lake-reservoir system, introducing 3 more new Euglesa. Nominally the remark- able malacofauna of Lake Baikal has been increased by the introduction of about 4 new subgenera and 13 new species of freshwater gastropods, both pulmonate and prosobranch (see the paper by Bekman and Starobogatov). However, several other studies tended more fully to appreciate species as biological entities with their con- comitant genetic, populational, and behavioral charac- teristics: Alimov on Sphaerium, Chukhchin on hydrobiids, Mikulich and Biryulina on Crenomytilus, Khokhutkin on polytypy in terrestrial gastropods, and Bogatov on varia- bility in Lymnaea stagnalis. The molluscan faunas of zoologically poorly known geo- graphical regions were studied in some detail: Arutyunova on terrestrial mollusks in the Crimea, Izzatullaev on Cent- ral Asian and Tadzhikistani land snails, Gul’bin on shallow water marine gastropods of the Kurile Islands, and Uva- lieva and Mukhitdinov on Kazakhstan. Several investiga- tions dealt with physiological problems attendant upon the adaptation to waters of varying salinities, especially in the White and Barents Seas. Researches which reviewed or discussed other regional faunal assemblages include Govberg on the White Sea and Ignat’ev et al. on the Okhotsk Sea and the Sea of Japan. Roginskaya investigated the Opisthobranchia of Sosnovets Island in the White Sea, while Zuev et al. stud- ied the populations of various cephalopods in the Atlantic Ocean. Certain researches centered on the autoecology of ma- rine and freshwater forms: Shustov on Bythinia leachi, Klebovich on Hydrobia ulvae, Spiridonov on Dreissena, Berger on Littorina, Potafeev on Lymnaea truncatula and Zhuravleva and Prazdnikov on Mytilus edulis. Biochemical analyses stressed adaptations to new en- vironments: Petkevich et al. on Mya arenaria in the Black Sea, while several papers were concerned with physio- logical parameters, most notably those concerning the re- productive biology of squids (Shevtsov) and scallops (Mal’tsev). Several studies reported the occurrences of mollusks introduced into the territory of the USSR; namely Mya arenaria in the Black Sea (Petkevich et al.), Melanoides tuberculatus pamirensis in the Pamirs (Akhrorov and Churshina) and Physodon integrum in Tadzhikistan (Izzatullaev). We have provided below a listing of abbreviations and acronyms which we have utilized in this compilation. AN — Akademiya nauk (Academy of Science) Biol. Morya — Biologiya Morya (Marine Biology) Biol. Morya Resp. Mezhved. sb. — (Interrepublic Marine Biology Collection) BSV__— Biol. Shel’fa Tezisy dokl. Vses. Konf. Vladivostok (Scientific theses on ocean shelf biology from the All Union Conference in Vladivostok) ES — English summary. GZ — Gidrobiol. Zhurnal (Hydrobiological Journal) IANT - Izvestiya Akad. Nauk Tadzh. SSR. Otd. biol. (Bulletin of the Academy of Sciences of Tadzhikistan, Depart- ment of Biological Sciences) IFML - Issled. fauny morei, Leningrad, Nauka (Studies of ma- rine fauna, Leningrad Science Press) ITNIIRKO —- Izv. Tikho-okeansk. nauch.-issled instituta rybn. khoziaistva i okeanogr. (Bulletin of the Pacific Scien- tific Research Institute of Fisheries and Oceanography) LMA _ - Lietuvos TSR mosklu Akademija DARBAI (Trudy A- kademiya Nauk Litovskoi SSSR). Vilna. (Works of the Academy of Sciences of the Lithuanian Republic of the USSR) NDVS -—- Nauch. Dokl. Vyssh. Shkol. Biol. Nauk. (Scientific Re- ports of the Higher Educational School for Biological Sciences) TLOE - Trudy Leningradskogo Obshchestva Estestvoispytatelei (Works of the Society of Leningrad Naturalists) TRO - Trudy Instituta Okeanologii. Akademiya Nauk SSSR. (Works of the Institute of Oceanology, Academy of Science, USSR) TVNIIMRXO -— Trudy Vsesoiuznogo Nauchno-Issledovatel’skogo Instituta morskogo rybnogo Khozyaistva i okeanografii. (Articles of the All-Union Research Institute of Marine Fisheries and Oceanography) Vestn. Zool. — Vestnik Zoologii (Zoological News) VLGU -— Vestnik Leningr. Gosudarstvennogo Universiteta (News from Leningrad State University) ZOB = - Zhur. Obshch. Biol. (Journal of General Biology) ZZ — Zoologicheskii Zhurnal. (Zoological Journal) We thank Mrs. Mary Jo Dent for her careful typing of the manuscript. Page 392 THE VELIGER Vol. 20; No. 4 GENERAL Gottxov, A. N. 1976. Regularities in the growth and variability of some mol- lusks. Gidrobiol. issled. samoochishcheniya vodoemov (Hydro- biological investigations of self-cleaning reservoirs), Leningrad, Pp. 97 - 118 [Variability is reflected in phenotypical modifications correlated with environmental influences, of which temperature is of prime importance] Govsers, L. I. 1975. The formation of the mollusk fauna of the White Sea. BSV, pp. 28 - 29 [Four faunistic complexes are distinguished, beginning in pre-boreal times and corresponding to 4 stages of development. The malaco- fauna derives from the Barents Sea] Ionat’Ev, A. V, E. V. Krasnov « V. E. SHEIGus 1975. The determination [of the effects] of temperature on growth of mollusks in the Sea of Japan and the Sea of Okhotsk by the isotopic oxygen method. BSV, pp. 62 - 63 [Analyses of isotopic oxygen in the calcite of the shells of 7 species of bivalves were made] IZZATULLAEV, Z. 1975. Little known land mollusks (Mollusca, Gastropoda) of the fauna of Central Asia. IANT, No. 2, pp. 39 - 45 [A description of the shell and partial genital anatomy, as well as the ecology, taxonomy, and geographic distribution are represented for 4 species of terrestrial mollusks] Kozurmna, L. M. 1976. Cellular stability in certain mollusks to changes in salinity. Biol. Morya, No. 1, pp. 36 - 40 (ES) [The stability of ciliated epithelium and isolated muscle of marine mollusks was studied. The damaging effect of low salinity depends on its low tonicity. Addition of saccharose to the saline solution con- siderably increases the duration of tissue sensitivity] KrivosHeEIna, L. V. 1976. On the molluscan fauna of the mountain lakes of South Altai. KazSSR Fylym Akad. Khabarlary Izv. AN KazSSR (Bulletin of the Khabarlary Academy of Science of Kazakh- stan) (Biol.) No. 3, pp. 26 - 32 [5 new species of Euglesa (Sphaeriidae) are proposed. Remarks on the distribution of mollusks and the reduced diversity along the gradient, plains-foothills-montane lakes, are provided] Minicuev, Yu. S. & Ya. I. StaRoBOGATOV 1976. On the phylogeny of mollusks. TLOE 84 (1): 37-42 [The hypothesis is proposed that the mollusks were derived from a Dinophilus-like ancestor. A new system of mollusks is proposed: 2 basic branches appeared at different stages of evolution, 1) mol- lusks with an internal cavity (coelom?) — _ the subtype Cochleo- phora; and, 2) mollusks without such an internal cavity — the subtype Peltophora. The latter diverged into 2 lineages: the Bival- via and the Loricata, from which the Aplacophora developed. The Cephalopoda and Monoplacophora arose independently within the limits of the Cochleophora, and the latter, in turn, led to the Gastro- poda] Srarosocatov, YA. I. 1975. 5 All-Union Conference on the study of mollusks. Sym- posium: ‘New methods of studying mollusks.’ ZZ 54 (9): 1423 [56 reports were given on the biology of mollusks] Voros’Ev, V. I. 1975. Dynamics of trace elements in the mollusks of the Volga Delta. Nekotoriye probl. ekol. zhivotnikh Nizhn. Povolzh’ya i sev. Kavkaza (Some problems of the ecology of the fauna of the lower Volga and northern Caucasus), Volgograd, pp. 3 - 13 [Aluminum, cobalt, manganese, nickel, molybdenum, zinc, lead, vanadium, titanium and silver were detected both in gastropods and bivalves. These elements were also noted in the sediments and waters of reservoirs] ZHGENTI, E. M. 1976. The Lutetiidae of the Middle Miocene, their evolution and stratigraphic significance. Metsniereba Tbilisi, 156 pp. [The systematics, phylogeny, ecology, extinction and geochrono- logical distribution of the family are reviewed] APLACOPHORA anp POLYPLACOPHORA Smenxo, B. I. 1975. On the systematics of the chiton genus Lepidozona Pils- bry. Biol. Morya, No. 3, pp. 13 - 28 (ES) [Seven species of Lepidozona inhabit the seas of the USSR: data are provided on their distribution (including Asiatic and Ameri- can waters), ecology and biology. Gurjanouvillia is synonymous with Lepidozona, and the species G. derjugini with L. kobjakovae. New taxa are L. thielei, L. multigranosa, L. ima and L. kobjakovae kamtchatkana] 1975. A new chiton subfamily — Juvenichitoninae (Ischnochi- tonidae) from the northwest Pacific. ZZ 54 (10): 1442-1451 (ES) [A new subfamily with 3 new genera (Juvenichiton, Micichiton and Nanichiton) and 5 new species is erected primarily on the basis of the radula’s having only 11 to 13 teeth per transverse row rather than, as the author states, the usual 17. Sirenko terms this putative reduction in the number of teeth: hypomorphosis] 1976. The systematics of the chiton genus Spongioradsia Pils- bry, 1893 (Ischnochitonina, Ischnochitonidae). Vestn. Zool. No. 2, pp. 50-55 (ES) [The status of the genus is verified and a diagnosis given. Spongio- radsia includes 3 species distributed in the Pacific, including a new one from the shores of the Kuril Islands] Vol. 20; No. 4 THE VELIGER Page 393 GASTROPODA, GENERAL AxuHrorov, FE. « N. M. CuyursHINA 1976. The tropical mollusk Melanoides tuberculatus pamirensis Lindh. in the Pamirs. ZZ 55 (5): 767-768 (ES) [In 1973 and 1974, large numbers of this snail, described from ex- cavated material, were found living in Dzhaushangoz Springs in the Pamirs] Beer, S. A., A. V. Lirsuits, L. K. Mas.ova & V. D. Zavorkin 1976. Localized distribution and ecology of Bythinia inflata in the northern Tomsk Region. Report 1: The influence of abiotic factors on the distribution of Bithynia. Med. parazitol. 1 para- zitar bolezni (Medical parasitology and parasitic diseases) 45 (1): 74-81 (ES) [Abiotic, hydrological factors affect the occurrence of B. inflata which was found locally at 54 of 439 stations throughout the flood plain of the Ob River] Gavrtov, S. I. « I. I. DEsyatiK 1975. The gastropods of Naroch’ Lake. Vestn. Belorus. un-ta (Reports of the White Russian University), ser. 2, No. 3, pp. GO) Bo [28 species are listed, most of which are littoral; greatest population densities are found in lightly silted substrates with rich vegetation] Guv’stn, V. V. 1975. The distribution and biogeographical composition of the gastropod fauna of the higher portions of the Middle Kurile Shelf (Urup and Simushir). Biol. Morya, No. 4, pp. 46 - 50 (ES) [The complex of hydrological conditions, especially water tempera- ture, influences most the vertical and horizontal distribution of the 81 species of snails] Uva.teva, K. K. 1976. The land mollusks of Kazakhstan and the problem of their further study. Chteniya Pamyati akad. E. N. Pavloskogo Dokl. prochit. na 8-om (1973) i g-om (1974) ezhegodn. chteniyakh. Nauka (Lectures of the Memorial Academy of E. N. Pavlov. Reports delivered at the 8t4 (1973) and gt (1974) Annual Lectures (Science). Alma-Ata, pp. 25 - 35 [Discussed are the roles of land mollusks in agriculture, in geology, and in the dispersal of helminths] PROSOBRANCHIA BEKMAN, M. Yu. & Ya. I. STAROBOGATOV 1975. Deep water Baikal mollusks and their related forms. Tr. Limnol. in-ta. sib. Otd. Acad. Nauk. SSSR (Works of the Lim- nological Inst. of the Siberian branch of the AN SSSR) 18 (38): 92-111 [6 new deep water species (from up to 200m) and 7 new shallow water species are described from Baikal. 'The latter, while showing clear relationship with the deep water forms, nevertheless point to the formation of an abyssal Baikal fauna. Intraspecific subdivisions of the genera Benedictia and Choanomphalus are discussed. Within the limits of the latter, 4 new subgenera are established. A revision of the systematics of the Baikal species of the subgenus Gyraulus of the genus Anisus shows that the only species of this genus in the “sors” zone (shallow peripheral lagoons) is A. (G.) ignotellus with 2 subspecies. It is also observed that among the Baikal deep water species there are certain ones not present in shallow water forms] BercER, V. YA. 1976. On the adaptation to decreasing salinity of several littoral White Sea mollusks. IFML, AN SSSR 17 (25): 59-111 [The molecular, cytological, and organismic reactions to changing salinity by Littorina littorea, L. obtusata, L. saxatilis and Hydrobia ulvae were studied, especially in differing ecological settings] 1976. A comparison of the reactions of Barents Sea and White Sea Littorina to variations in environmental salinity and a dis- cussion of the criteria of physiological races. IFML, AN SSSR 17 (25): 112-123 {In regard to respiratory and hematological activity, L. obtusata and L. littorea transferred from one sea to another adapted pheno- typically to the different salinity regimes] 1976. Seasonal changes in the sensitivity of White Sea Littorina obtusata (L.) to salinity in its habitat. IFML, AN SSSR 17 (25): 155-159 Bercer, V. Ya. & N. N. CHERNYSHEVA 1975. A comparative study of the reaction of White Sea and Barents Sea Littorina to altered salinities. Ekologiya 5: 49 - 53 [Samples from populations of L. littorea and L. obtusata eventually acclimatize to salinity regimes of respective seas] Cuuxkucuin, V. D. 1976. The systematic placement and ecology of Hydrobiidae of the Black Sea. Biol. morya Resp. mezhved. sb. vyp. 36, pp. 65-75 [Black Sea hydrobiids are closely related to H. ventrosa of the At- lantic. Several ecological varieties appear and are correlated with such factors as depth distribution, growth, sexual dimorphism and parasitical gigantism. There are species with pelagic larvae (H. acuta) and those with direct development (H. ventrosa and H. pusilla). Although H. acuta has the greatest geographical distribu- tion, H. ventrosa exhibits a wider salinity tolerance] 1976. The life cycle and growth rate of Hydrobia acuta (Drap.) and H. ventrosa (Mont.) in the Black Sea. Biol. morya Resp. mezhved. sb., vyp. 37, pp. 85-90 [Both species have a one-year life cycle. From numerous diminutive eggs, benthic larvae develop directly in H. ventrosa while they become pelagic in H. acuta] Ku_esovicu, V. V. x A. PR KoNDRATENKOV 1976. The influence of hydrolized albumen on the adaptability of Hydrobia ulvae to reduced salinities. IFML, AN SSSR 17 (25): 132-135 1976. Influence of catabolic products of some marine organisms Page 394 THE VELIGER Vol. 20; No. 4 on the saline adaptability of Hydrobia uluae. IFML, AN SSSR 17 (25): 136-141 [The generation of metabolic products of low molecular weight facilitates adaptability to different salinity regimes] KHLEBOVICH, V. V. & O. Yu. MIKHAILOVA 1975. The influence of periodic changes in salinity on the activ- ity of Hydrobia ulvae. ZZ 54 (10): 1452-1456 (ES) [Experimental evidence illustrates the acclimatization of this eury- haline snail to different and variable salinity regimes (6 - 24%) ] KonpraTENnKov, A. P 1976. Analysis of inter-populational differences of Hydrobia ulvae using gradual acclimatization. IFML, AN SSSR 17 (25): 124 - 131 [Between populations, clear differences in salinity tolerances are shown] KorzuHuev, P. A. « I. O. ALYAKRINSKAYA 1976. Biochemical characteristics of the hemolymph of some hemocyanin-bearing gastropods. Dokl. AN SSSR (Reports, AN USSR) 228 (1): 239-241 [The quantitative contents of hemocyanin in the hemolymph of Littorina littorea and Viviparus contectus were examined] PotisHcHuK, V. V.« I. B. Lyurin 1976. The occurrence of Potamopyrgus jenkinsi (Smith, 1889) in the Pripyatskii Forest. Dokl. AN SSSR (Reports, AN USSR) (B) No. 4, pp. 367 - 369 (ES) [P. jenkinsi, usually an inhabitant of zones near the ocean, was col- lected in the Pripyatsky River] RaDOMAN, PAVLE 1975. Species formation in the genus Belgrandiella and related genera in the Balkans. Glasnik Prir. Muzeya Beogradu (Pub- lic Natural History Museum of Belgrade) (B) 30: 29-69 (Serbian, ES) [A new monotypic genus (Sarajana) and 13 new species of the 3 o- ther genera in the subfamily Horatiinae, family Orientaliidae, are introduced. The name Frauenfeldia Clessin (non Eger) is replaced by Graziana (novum). Some remarks on the distribution of the forms are provided] Suustoy, A. I. 1975. Characteristics of the biology of Bithynia leachi (Shep.) in the reservoirs of Central Kazakhstan. Inst. Zool. AN KazS- SR, Alma-Ata, pp. 131-141 [Snails are active from the end of April to early October. Data on their abundance and some of their environmental tolerances are provided] OPISTHOBRANCHIA Kyarazova, A. D. « V. V. Rostova 1976. A study in the changes of synthesis of albumens and RNA in the tissues of the White Sea Coryphella rufibranchialis in re- duced environmental salinity. IFML, AN SSSR 17 (25): 142 - 154 [Synthesis is reduced in hyposaline conditions and sharply restored with the return to normal salinities] Rocinsxaya, I, S. 1976. Opisthobranchia of Sosnovets Island (White Sea). ZZ 55 (1): 23-28 (ES) [Three littoral opisthobranchs, along with data of their ecology and reproductive biology, are recorded from Sosnovets: the sacoglossan Acteonia cocksi and the nudibranchs, Onchidoris fusca and Ancula cristata] PULMONATA, LAND ArutTyunova, L. D. 1975. Notes on some land mollusks along the southern shore of Crimea. Biol. zh. Armenia (Biological Journal of Armenia) 28 (10): 104-109 (Armenian Summary) [8 species of stylommatophorans are noted; descriptions of genital morphology and anatomical variability are provided] BraTcuik, R. Ya. 1976. A method for the rapid fixation of land snails. ZZ 55 (7): 1078 - 1079 (ES) Harris, S. V. 1976. ‘The vertical stratification of the distribution of land mol- lusks on the Iraqi slope of the Persian Mountains and in the Skaly Hills of Alberta, Canada. Vysokogorn. Geoekologiya (High Mountain Ecology), Moscow, pp. 101 - 102 IZZATULLAEV, Z. 1975. Some notes on the biology of slugs (Mollusca; Pulmonata) which are harmful to agriculture in Tadzhikistan. IANT, No. 4, Pp. 22 - 24 [Deroceras caucasicum and Parmacella levanderi are especially dele- terious to agriculture in the autumn and spring, respectively] 1975. Characteristics of the distribution of land mollusks in the Gissarskii Mountain Ridge and contiguous regions of Tadzhiki- stan. Zool. sb. ch. 1 (Zool. Coll. Pt. 1), Dushanbe “Donish”, Pp. 212-224 [The malacofauna falls into 5 groups: 15 widely spread species, 7 species from the mountain areas of Europe and Eastern Asia, 27 Central Asian endemics, 6 tropical species or endemics with tropi- cal relationships, and 3 introduced species] 1975. A study of the molluscan fauna in the environs of Khorog City. Zool. sb. ch. 2 (Zool. Coll. Prt. 2), Dushanbe “Donish”, PP. 5-7 [On the plains of the Gunt, Shakhdara, and Pyandzh Rivers, 9 species of land mollusks occur. Nesovitrea petronella is reported for the first time and 3 species of freshwater pulmonates are also re- corded] Vol. 20; No. 4 THE VELIGER Page 395 Kuoxkuutkxm, I. M. 1975. Polymorphisms and [population] structure of land snail taxa in the molluscan fauna of the USSR. Trudy in-ta ekol. rast. zhivotnikh. Uralsk nauch. tsentr AN SSSR (Transactions of the Institute of floral and faunal Ecology, Ural Sci. Center, Acad. Sci. USSR) 96, pp. 116- 138 [Polytypic land mollusks have large numbers of subspecies] 1976. On the physiological differentiation of populations of the molluscan genus Bradybaena. NDVS, No. 7, pp. 54-57 [No differences were detected in carbon dioxide respiration between Bradybaena fruticum and various morphs of B. lantzt] KuoxkuutTn, I. M. & A. I. LAzAREVA 1975. Polymorphism and imitative coloring in populations of land mollusks. ZOB 36 (6): 863 - 869 (ES) [Polymorphism occurs more frequently than usually appreciated. A complex of phenotypic variants in 3 helicid species in the Caucasus is discussed] Lyurin, I. B. 1975. Seasonal changes in several morphological and physio- logical features in mollusks from the Loess of the Ukraine. Prirod. obstanovka i fauny proshlogo (Natural Conditions and Fauna of the Past) 9, Nauk dumka, Kiev, pp. 57 - 67 [An investigation of the tempo of evolution in land and freshwater mollusks of the Loess shows that changes in temperature to extreme- ‘ly high or low values led to a reduction in the range of phenotypic variations] Mouxurtoinov, A. B. 1975. Zoogeographical characteristics of the land molluscan fauna of northern Tadzhikistan. IANT, No. 4, pp. 18-21 (Tadzhik Summary) [42 species, some endemic, are listed from this mountainous region of Central Asia. Notes on local distribution in the Gissaro-Darvas- kii Region of the Afghano-Turkestani frontier are presented] Namatov, T. N. 1976. Respiration of some land mollusks at various tempera- tures. Perekrestn. adaptatsii k prirod. faktoram sredy (Corre- lation of adaptations to natural environmental factors), Frunze, Ilim, pp. 59 - 65 [Metabolic rate depends on body weight. Respiratory rates are lower at 20° C than at 10° C] Ostroumova, N. K. 1976. The fine structures of neural fibres terminating in. the pericardium of the cardiac muscles of Achatina. Fiziol. i bio- khimiya mediator. protsessov. (Physiology and Biochemical Mediating Processes), Moskva, pp. 99 - 100 PakHoruKkova, L. V. 1976. A quantitative assay of feeding in Deroceras agrestis and D. reticulatum. ZZ 55 (1): 29 - 33 (ES) [Feeding and assimilation were studied in these 2 species of slugs. Bioenergetically, D. agrestis was more efficient. As total weight of the animals increases, the amount of food consumed decreases] Runxowa, G. G., V.N. Maxsimov, L. A. Kovau’cHuk & I. M. KHokuyurKIN 1975. Extra-mitochondrial and mitochondrial oxidative path- ways in sinistral and dextral morphs of the snail, Bradybaena lantzi, at various temperatures. Trudy Inst. Ekol. rast. i zhivot- nikh, Uralsk, nauch. tsentr AN SSSR (Transactions of the Inst. of floral and faunal Ecology, Ural Sci. Center, Acad. Sci. USSR) 97: 113-116 1975. Thyroxin sensitivity of endogenous oxidase in homogen- ates of 2 morphs of Bradybaena lantzi. Trudy Inst. Ekol. rast. i zhivotnikh, Uralsk, nauch. tsentr AN SSSR (Transactions of the Institute of floral and faunal Ecology, Ural Sci. Center, Acad. Sci. USSR) 97: 117-120 [Although some distinctions were shown, no consistent differences occur between sinistral and dextral snails] Sumeixko, A. A. 1975. The mollusks of the subfamily Lauriinae in the USSR (Pulmonata, Pupillidae). ZZ 54 (12): 1767-1782 (ES) [The Caucasian Lauriinae show considerable conchological and anatomical variation, especially in the structure of the male sexual apparatus. In the Caucasus the Lauriinae include the genus Euxino- lauria with 4 subgenera: Matschachelia (new subgenus), Caucasi- pupa, Euxtnolaurta, and Neolauria (new subgenus), and the single species Lauria cylindracea in the genus Lauria. The subfamily Arg- ninae has the single Carpathian species Orna bielzi. 3 new species of the genus Euxinolauria are also described] 1976. Characteristics of the organization and systematics of the family Orculidae (Gastropoda). NDVS, No. 4, pp. 47-58 [In conchological characteristics and in genital anatomy, the family is one of the more unique groups of Paleoarctic Orthurethra. The complex of shell characters indicates the primitiveness of the group while the structure of the penial appendix testifies to an advanced degree of specialization. The family is composed of 2 subfamilies, the Orculinae and the Pagodulininae, the former with 3 genera, the latter with 1. Two new subgenera are introduced: Mesorculella (of Orculella) and Crystallifera (of Pagodulina) | 1976. Evolutionary pathways and the phylogenetic significance of the excretory apparatus of the Pulmonata. ZZ 55(2): 215-225 (ES) [Really limited to the Stylommatophora, this study points out that from the primitive orthurethrous condition, wherein the ureter consists of open ciliated grooves, several advance stages in the evo- lution of the excretory apparatus can be recognized: Mesurethra (shortened kidney); Sigmurethra (ureters closed); Heterurethra (lung shorter and wider). The Orthurethra represent a phylo- genetically primitive unit whereas the other presently recognized taxonomic grouping may consist of heterogeneous components in- dependently evolved] Sxtyar, I. Ya. 1975. A new slug of the genus Deroceras (Gastropoda, Lima- cidae) from the Crimea. Vestn. zoologii, No. 6, pp. 79 - 82 (ES) Page 396 [Distinguished chiefly by its penial structure, D. ramosum is de- scribed as new from the forest zone on the Crimean peninsula] PULMONATA, AQUATIC Bocartoy, V. V. 1975. Variability of Lymnaea stagnalis L. in small reservoirs. Vopr. ekol. zhivotnykh (Problems in animal ecology) 2, Kalin- in, pp. 3 - 10 [Populations of snails in 6 small reservoirs in the Kalinin Region were studied. The shape of the shell is influenced by the OLA (Os- cillating Level of Amplitude) of the reservoir. As the OLA increases, the shell increases in length; this effect is especially marked from May 1 to September, the period of the major blooms of macro- phytes upon which the snails feed] IZZATULLAEV, Z. 1975. On the discovery of the freshwater mollusk Physodon in- tegrum (Haldeman) (Mollusca, Basommatophora, Physidae) in Tadzhikistan. Zool. sb. ch. 2 (Zool. Coll. Prt. 2), Dushanbe “Donish”, pp. 8-12 [This is the first recorded occurrence of the species in Central Asia in the Plain of Vakhshskya; the shell and the genital anatomy are illustrated] Kamarown, N.N. 1976. The structure and cellular organization of the osphradium of Lymnaea stagnalis L. Arkhiv anatomii gistologii, i embrio- logii 71 (8): 87-90 (ES) [The osphradium consists of epithelial canals composed of 3 types of cells (secretory, filamentous, and epithelial) connected to basal ganglia which provide fine sensory innervations] Krustov, N. D. 1975. On an analysis of contemporary methods of molluscan systematics and the limit of their applications in the case of the lymnaeids. Vopr. Biol. i sistematiki zhivotnikh Smolensk. i sop- redel’n obl. (Problems of biology and faunal systematics of Smolensk and the surrounding region), Smolensk, pp. 12 - 28 Keruc ov, N. D., S. D. Drozpova « T. N. MAaKAROVA 1975. An ecological and morphological study of Lymnaea cor- vus (Gmelin, 1778) and L. corviformis (Bourgignat in Ser- vain, 1881). Vopr. biol. i sistematiki zhivotnikh Smolensk. i sopredel’n obl. (Problems of biology and faunal systematics of Smolensk and the surrounding region), Smolensk, pp. 29 - 35 Lersson, N. L. « L. T. Frotova 1975. Localization of cell division in the intestinal epithelium of marine animals. 1. Crenomytilus grayanus. Biol. Morya, No. 5: 15-22 (ES) [Autoradiographic methods using thymidine-Hs noted no special localization of mitoses along the gut of the mussel, a condition un- like that found in higher vertebrates] Marorova, V. G. & I. V. CHERNOVA 1976. Selective sensitivity to traces of acetylcholine in separate neurons in the large parietal and visceral ganglia of pond snails. THE VELIGER Vol. 20; No. 4 Fiziol. i Biokhimiya mediator. protsessov. (Physiology and Biochemical Mediating Processes), Moskva, pp. 85 - 86 PorarEzev, N. E. 1975. Some data on the ecology of Lymnaea truncatula in the Kursk region. Ekol. i eksperim. parazitol. (Ecology and ex- perimental parasitology), Leningrad Univ., 1: 154 - 160 (ES) [L. truncatula is an intermediate host of Fasciola hepatica. Its seasonal and diurnal activity was studied; 2 main biotopes, based on hydrological conditions, were recognized and infected snails were limited to one of the biotopes] Porapina, N. V. 1976. A study of the amebocytes in the blood of Lymnaea stag- nalis by autoradiography. Materialy III. Nauch. Konf. molo- dykh uchenykh-morfologov Moskvy (Materials of the 3° sci- entific conference of young student morphologists) , First Mos- cow Med. Inst., Mosk. Univ., pp. 34 - 37 STaRosocaTov, YA. I. 1976. ‘The systematics and phylogeny of the Lymnaeidae (Gast- ropoda, Pulmonata, Basommatophora). Probl. zoologii, Lenin- grad, Nauka, pp. 79 - 81 BIVALVIA Autmov, A. EF 1976. Notes on the variability of some mollusks of the family Sphaeriidae. Gidrobiol. issled. samoochishcheniya vodoemov (Hydrobiological investigations of self-cleaning reservoirs), Leningrad, pp. 119 - 128 [Variability in the shells of Sphaertum corneum, S. scaldianum, S. solidum and Sphaeristrum rivicola was examined. As might be ex- pected, stenotopic species exhibited little variation while eurytopic species, such as S. corneum, exhibited the greatest variability] BeLocrupov, E. A. «& V. N. Mat’TsEV 1975. The spawning of scallops in Pos’et Bay. ITNIIRKO 96: 273-278 (ES) [Spawning takes place earlier in shallow water coves than in the open parts of the bay. Spawning continues for about 10 days in each different portion of the bay and lasts about 1.5 months in the bay as a whole] Bercer, V. Z. 1976. The effect of several physiologically active materials on the adaptive reaction of ciliated epithelial cells on the ctenidia of mussels in changing saline conditions. Tsitologiya 18 (8) : 981 - 984 (ES) Ecereva, I. V. 1976. Novelties in the fauna of the Kuibyshev Reservoir. Ryb. khozyaistvo (Fisheries), No. 3, p. 29 [From 1966 to 1970 1615000 specimens of Monodacna colorata from Taganrogsky Bay were introduced, and in September 1975, only 12 specimens, measuring 4 to 22mm, were recovered] Vol. 20; No. 4 GorsBarENKO, S. A. & A. V. IcNAT’EV 1975. Changes in isotopic oxygen concentration in shells [in relation] to temperatures during growth of deep-water bivalves in the Japan Sea. BSV, pp. 32 - 33 Gurina, V. I. 1976. An autoradiographic study of the synthesis of albumen and RNA in the intestinal epithelium of White Sea mussels in relation to differing degrees of salinity. VLGU, No. 3, pp. 59 - 63 (ES) Ivanrtsiv, V. V. 1975. Features of the distribution of the Unionidae in Kremen- chusky Reservoir. Vestn. zoologii. No. 6, pp. 82 - 84 Karanov, A. I. 1975. General conformity of variability in marine bivalves. BSV, PP. 74-75 1975. On the interpretation of the logarithmic spiral in connec- tion with the analysis of the variability and growth of bivalve mollusks. ZZ 54 (10): 1457-1467 (ES) [Equations are provided which describe the dynamics of growth components in bivalves] KHARCHENKO, T. A. 1975. The Dniepr sea-roach as a factor in the control of dreis- senids in the canals. Biol. samoochishchenie i formir. kachestva vody (Biological methods for producing cleaner water). Nauka, pp. 73 - 74 [Since the sea-roach feeds on Dreissena polymorpha and D. bug- ensis, introduction of this fish into the Kakhovsky reservoir and else- where would help in the control of the bivalve] Kopotova, O. P « B. M. LocviNENKO 1976. Electrophoresis of muscle albumens of freshwater union- ids. NDVS, No. 4, pp. 142-144 [Differences were detected in the 2 different contractile elements of the adductor muscle] Korcuacin, V. P. « E. V. Krasnov 1975. A comparative study of the amino acid contents of albu- men of the shells and byssus of several species of mussels in Peter the Great Bay (Sea of Japan). BSV, pp. 87 - 88 [Mytilus edulis, Modiolus modiolus, and Crenomytilus grayanus were studied; the amino acid contents of the periostracum, the bys- sus and the ligament are species specific] Kurakovsku, E. E. . 1976. The influence of reduced salinity on the neuro-secretory system of Mytilus edulis (L.). IFML, AN SSSR 17 (25): 160 - 166 [In the cerebral ganglia, 3 morphologically and topographically distinct types of neurosecretory cells occur. Although neurosecretion is altered under reduced salinity, our data are insufficient to quanti- fy the response] Kuz’movicy, L. G., I. V. SHust e I. M. Kostnix 1976. Histological characteristics of the gonads of Anodonta piscinalis Nilss. NDVS, No. 6, pp. 63 - 68 THE VELIGER Page 397 [Histochemical analysis of lipids, neutral fats, and polysaccharides of unparasitized sex cells] Mav’tsev, V. N. 1975. Some regularity in the settling of Pecten larvae in collec- tors in Pos’et Bay. ITNIIRKO 96: 179-282 (ES) [The extent, date, and density of spat by depths are given for Peter the Great Bay; data on the dependence of the date of settling on the temperature of the water are provided] Mixuticu, L. V. « M. G. BrrruLina 1975. On the problem of the species-concept for Crenomytilus grayanus (Dunker). Trudy Tikhookean. in-ta (Transactions of the Pacific Ocean Institute) 9: 114 - 118 [The length/width index varies between 1.4 and 2.7 in populations from various localities and different ecological stations. These differ- ences in the form of the shell reflect the range of intraspecific varia- tion, and all individuals with different indices form a single species] Naumov, A. D. 1976. Variability of Portlandia arctica (Gray) in the complex of [environmental] characteristics of the White Sea. Probl. zoo- logii, Leningrad, Nauka, pp. 67 - 69 1976. The adaptation of Portlandia arctica var. portlandica (Taxodonta) of the White Sea to waters of lower salinity. ZZ 55 (3): 449-453 (ES) [Differences were noted in the physiological activity of individuals from shallow and deep water; it is suggested that in the White Sea there are 2 races, with the deep water one being euryhaline] PetKevicn, T. A., R. P Kanpyux « I. A. STEPANYUK 1975. On the biochemistry of the mollusk Mya arenaria (L.), newly discovered in the Black Sea. GZ, 11 (5): 101 - 106 Raxoy, V. A. 1975. Changes in the shell shape of Swiftopecten swifti during growth. ITNIIRKO 96: 302-304 (ES) Sanina, L. V. 1975. Preliminary estimates of the nutrient requirements for filter-feeding mollusks in the northern Caspian. TVNIIMRXO 107: 43-47 (ES) SKIRKYAVICHENE, Z. Yu. 1975. Amino acid content of the soft parts of dreissenids in Kur- shyu-Mares Bay. 2. Free amino acids at various times of the year. LMA (B), No. 3 (71): 127-133 (ES) [Chromatographs from dreissenid tissue taken in 1967 and 1968 showed a content of 21 free amino acids; seasonal variations were noted] Spmmonov, Yu. I. 1975. ‘The growth structure of a population and persistence of life of Dreissena in the Volgograd reservoir. Tr. kompleks. eks- peditsii Saratov. un-ta po izuch. Volgogr. i Saratov vodokhrani- lishch. (Transactions of the Combined Expedition of Saratov University to Study Volgograd and Saratov Water Reservoirs) 5, Pp. 84-86 Page 398 Tapi¢, ANTE 1975. Some Unio and Anodonta species in various habitats. Glasnik. Prir. Muzyea Beogradu (Public Natural History Mu- seum of Belgrade) (B), 30: 103-118 (Serbian, German S.) [Studies of 10 species of Anodonta and 2 species of Unio show that shell morphology is correlated with environmental conditions] Timm, Vir 1976. On the Pisidiidae of the lakes of the Chudsky-Pskov Re- gion (Lake Peipsi-Pihkva). Izv. AN EstSSR, Biologiya (Bulle- tin of the Academy of Science of Estonia, Biology) 25 (1): 37-52 (ES) [30 species of 6 genera were collected from 1964 to 1974: Sphaeri- um, 1; Amesoda, 1; Musculium, 1; Pisidium, 2; Euglesa, 22; Neo- pisidium, 3. 26 species were collected for the first time in reservoirs, 2 of which are new to Estonia, and 3 species are new to science (E. petpsi, E. pihkva, and N. stelfoxt). Although the majority of species occurs in all the reservoirs, none appears in large numbers. Data are provided on the depth distribution and bottom preferences of Pisidium] ZHURAVLEVA, N. G. « E. V. PRAZDNIKOV 1975. The ecology, reproduction, and development of the mussel Mytilus edulis in the Barents Sea. BSV, pp. 52 - 53 CEPHALOPODA Brryuxova, Yu. E. « G. V. ZuEv 1976. The asymmetrical eyes of the Histioteuthidae in connec- tion with their ecology. Biol. morya. Resp. mezhved. sb., vyp. 38, pp. 63 - 67 [It is concluded that the larger eye functions on the surface, the smaller in the depths] Ngsis, K. N. 1975. Evolution of adaptive forms in Recent cephalopods. TRO 101: 124-142 (ES) [The morphological adaptations to diverse habitats (i.e. bentho- pelagic, nectobenthic, nektonic, benthic and planktonic) are con- sidered] 1976. Stimulation of bioluminescence in cephalopods. Okeano- logiya 16 (1):150-154 (ES) [A weak solution of hydrogen peroxide intensifies the luciferin-luci- ferase reaction and was used on ship-board to stimulate the photo- phores in several species of epipelagic squids] SucHepkin, V. Ya., G. E. SHuL’MAN & T. G. SIGAEVA 1976. Features of the lipid content in Mediterranean squids in various ecologies. GZ 12 (3): 76-79 THE VELIGER Vol. 20; No. 4 Suevrsoy, G. A. 1975. Spawning in the Pacific Ocean squid, Todarodes pacificus Steenstrup, in the southern part of the Sea of Okhotsk. ITNII RKO o6: 121-127 (ES) [A population appears in the Okhotsk in autumn; length-weight measurements are given for males and females] Zuev, G. V. 1975. Some notes on the intraspecific grouping of Ommastre- phes pteropus (Cephalopoda, Mollusca) in the tropical At- lantic. Biol. issled. v. tropich. zonye okeana (Biological studies of the tropical zone of the ocean), Kiev, Nauk dumka, pp. 68 - 76 [In this zone, the epipelagic Ommastrephes pteropus forms 3 bio- logically specific populations: Canarian, mid-passage and equa- torial] ZueEv, G. V., K. N. Nesis & Cu. M. NicMATULLIN 1975. The systematics and evolution of the squid genera Omma- strephes and Symplectoteuthis (Cephalopoda, Ommastreph- idae). ZZ 54 (10): 1468-1479 (ES) [The authors claim that photophores are the most significant taxo- bases in deducing the phylogenetic relationships of several omma- strephid squids. As is general in teuthology, most of the genera are monotypic. Some paleontological speculation based on the modern distribution of the species as well as a phylogenetic dendrogram are provided] 1976. The distribution of the genera Ommastrephes d’Orbigny 1835, Stenoteuthis Verrill 1880, and Toderodes Steenstrup, 1880 (Cephalopoda, Oegopsidae) in the Atlantic Ocean. Biol. Mosk. ob-va ispyt. prirody (Bull. Moscow Naturalists Soc.) 81 (4): 53 - 63 (ES) [From field observations and the literature, it is concluded that S. pteropus consists of no fewer than 9 partially sympatric popula- tions mainly equatorial, that O. bartrami is subtropical, and that 7: Sagittatus occurs in subtropical and boreal waters of the northwest- ern Atlantic. Another species of Todarodes, T. angolensis, is of a more southerly distribution in sub-Antarctic waters] Zuev, G. V. & Cu. M. NicMATULLIN 1975. The spatial structure of the distribution of the ocean squid Ommastrephes pteropus Steenstrup in the Eastern At- lantic. Biol. issled. v tropich. zonye okeana (Biological studies of the tropical zone of the ocean), Kiev, Nauk dumka, pp. 56 - 67 [The Atlantic distribution of the species includes the entire tropical zone with northern limits in Africa at 20° to 22° N in winter and in summer to 30° or 32°N. The southern limit is 20° -23°S throughout the year. Productivity depends on variations of the vertical circulation of the water, and the population density of the squid increases in the winter] Vol. 20; No. 4 THE VELIGER Page 399 A Note on Changes in Marine Intertidal Fungus Taxonomy BY DAVID R. LINDBERG Department of Invertebrate Zoology California Academy of Sciences, Golden Gate Park San Francisco, California 94118 WITH EVER INCREASING LITERATURE and specialization in the natural sciences it is becoming increasingly difficult for workers to keep abreast of developments within their respective fields. It is even more difficult to be aware of the developments and changes within areas or with organ- isms only tangentially associated with one’s specialty. So while a worker may publish a modern up-to-date mono- graph or revision of his or her specialty, he or she may un- knowingly include incorrect information on other organ- isms in the same text. I recently came across one of these problems in my work and since it involves an organism familiar to many malacologists, cirripedologists, and inter- tidal ecologists, I am calling attention to the taxonomic status of this species. The marine fungus Didymella conchae Bonar, 1936, infects mollusks, barnacles, and other calcium carbonate secreting organisms, and is a major biological modifier of color patterns and morphologies of these organisms. This fungus is easily identified by the spongy appearance of the substratum and the black fruiting bodies visible at low magnification at the surface of the infected area. However, the name D. conchae is a junior synonym of Pharcidia balani (Winter) Bauch, 1936 (J. J. Kohlmeyer, in litt. July, 1977) and the usage of D. conchae is incor- rect. Dr. Kohlmeyer further advises that this species could also be treated as a lichen as it sometimes forms associa- tions with blue-green algae. The lichen name A7thopyre- nia sublitoralis (Leighton) Arnold, 1891 is available, but junior to P. balani. However, the genus Arthopyrenia is in need of revision and an earlier available lichen name may be found, again changing the name of the shell-infecting organism. Until that time, Dr. Kohlmeyer considers P balani the valid name for this fungus. For further infor- mation on marine fungi see KoHLMEYER & KOHLMEYER (1964 - 1969). Literature Cited Koxuumeyer, J. & E. KOHLMEYER 1964-1969. Icones Fungorum Maris. Germany. 7 vols. J. Cramer, Weinheim « Lehre, A Rectification of a Statement Regarding the Lamarckian Collection in the Book ‘“‘Murex Shells of the World” by George E. Radwin and Anthony D’Attilio BY ANTHONY D’ATTILIO SHORTLY AFTER THE UNTIMELY death of Dr. George E. Radwin, senior author of “Murex Shells of the World” (Stanford University Press, Stanford, California, U.S. A., 1976, 284 pp.) an unfortunate situation has come to light. On page 58, the following sentence appears: “Cernohorsky’s assertion of having located this and other Lamarckian types is open to question, for Dr. Binder, of the Muséum d’Histoire Naturelle, Geneva, has stated (in litteris) to both of us separately that it is essentially im- possible to determine the identity of Lamarck’s types.” The basis of this statement is unknown, and the state- ment is completely erroneous. The junior author never corresponded with Dr. Binder on this or any other matter. Furthermore, in inventorying Dr. Radwin’s professional papers following his death, no correspondence relating to the above matter was found. Because the junior author failed to notice thisstrange statement during proof-reading of the book, it was unfortunately printed. As Dr. Radwin is deceased, the unusual circumstances responsible for the above matter may never be known. However, it is hoped that this notice will help clarify the regrettable situation for all, especially the most affec- ted parties, Walter O. Cernohorsky and Dr. E. Binder. Deshayes Types in the National Museum, Paris BY TWILA BRATCHER IN A PERSONAL COMMUNICATION from Dr. Philippe Bouchet, he states that the Deshayes types, formerly in the Ecole des Mines, Paris, have been accessioned by the Muséum Nationale d’Histoire Naturelle, Laboratoire de Biologie des Invertébrés Marins et de Malacologie, 55 Rue de Buffon, 75005 Paris, France. Page 400 A very Generous Contribution from the Conchological Club of Southern California A FEW DAYS AFTER our January issue had gone to press, a very generous donation to the Veliger Endowment Fund was received from the officers and members of the Con- chological Club of Southern California. We express not only the thanks of the C. M. S., Inc., but of malacologists in general since all benefit from this gift, at least indirectly. As we have had occasion to point out on previous occa- sions, the income from the Endowment Fund helps the Society to keep the level of the dues as well as the sub- scription rate for the journal from rising as rapidly as the pressures of inflation would otherwise make necessary. W.S. M. Tue WESTERN SocteTy oF MALACOLocIsTS gave their 1977 Honor Award to Dr. Rudolf Stohler in appreciation for his many years as Editor of The Veliger, and for his continued support of the field of Malacology. Helen DuShane, President, 1977 Western Society of Malacologists Important If the address sheet of this issue is PINK, it is to indicate that your dues remittance had not arrived at the time the mailing was prepared (7. e. by March 1, 1978). We wish to take this opportunity to remind our Members that a reinstatement fee of one dollar becomes due if member- ship renewals have not been received by C.M.S., Inc. by April 15, 1978. However, in view of the unreliability of the postal service, members should not be alarmed by this notice as their remittances may be received between the first of March and the date of mailing this issue on April 1. From overseas addresses we must allow a minimum of 6 THE VELIGER Vol. 20; No. 4 weeks for surface mail. On the other hand, it is possible that the envelope and dues notice enclosed between pages 299 and 300 of the January issue have escaped your atten- tion. If so, now is the time to use them to avoid interrup- tion in the delivery of this periodical. Sale of C. M. S. Publications: Effective January 1, 1978, all back volumes still in print, both paper covered and cloth bound, will be available only from Mr. Arthur C. West, P. O. Box 730, Oakhurst, CA(lifornia) 93644, at the prices indicated in our Notes and News section, plus postage and, where applicable, California State Sales Tax. The same will apply to the Supplements that are still in print, except for supplements to vol. 7 (Glossary) and 15 (Ovulidae), which are sold by The Shell Cabinet, P. O. Box 29, Falls Church, VI (rginia) 22046; and supplement to volume 18 (Chitons) which is available from Hopkins Marine Station, Pacific Grove, CA (lifornia) 93950. Single Copies of “The Veliger”: We have on hand some individual copies of earlier issues of our journal and are preparing a list of the various issues available with the prices. Some issues are present in only one or two copies, while others may be present in 10 or more copies. As we are anxious to make room, we will offer these numbers at an exceptionally low price. This list will be presented in a forthcoming issue in the Notes and News section. Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. That the printed date is the actual date of pub- lication under the rules of the International Commission on Zoological Nomenclature is based on the following facts: 1) The journal is delivered to the Post Office on the first day of each quarter, ready for dispatch; 2) at least three copies are mailed either as first class items or by air mail; 3) about 20 copies are delivered in person to the mail boxes or to the offices of members in the Berkeley area; 4) two copies are delivered to the re- ceiving department of the General Library of the Univer- sity of California in Berkeley. Thus our publication is available in the meaning of the Code of the ICZN. The printed publication date, therefore, may be relied upon for purposes of establishing priority of new taxa. Vol. 20; No. 4 CaLiFoRNIA MaracozootocicaL Socrety, Inc. announces Backnumbers of THE VELIGER and other publications Volumes 1 through 8 and 10 through 12 are out of print. Volume 9: $22.- — Volume 13: $24.- — Volume 14: $28.- Volume 15: $28.- Volume 16: $32.- Volumes 17, 18 and 19: $34.- each. We have a limited number of volumes g, 11, 13, 14 to 19 available bound in full library buckram, black with gold title. These volumes sell as follows: 9 - $27.-; 11 and 13 - $29.- each; 14 and 15 - $33.- each; 16 - $38.-; 17, 18 and 19 - $41.75 each. Supplements Supplement to Volume 3: $6.- [Part 1: Opisthobranch Mollusks of California by Prof. Ernst Marcus; Part 2: The Anaspidea of California by Prof. R. Beeman, and The Thecosomata and Gymnosomata of the Cali- fornia Current by Prof. John A. McGowan] [The two parts are available separately at $3.- each] Supplement to Volume 6: out of print. Supplement to Volume 7: available again; see announce- ment below. Supplement to Volume 11: $6.-. [The Biology of Acmaea by Prof. D. P. Assortrt et al., ed.] Supplement to Volume 14: $6.-. [The Northwest American Tellinidae by Dr. E. V. Coan] Supplement to Volume 15: Our stock is exhausted, but copies are still available from The Shell Cabinet, P. O. Box 29, Falls Church, Virginia 22046. [A systematic Revision of the Recent Cypraeid Family Ovulidae by Crawrorp Nem. Cate] Supplement to Volume 16: $8.-. [The Panamic-Galapagan Epitoniidae by Mrs. Helen DuShane] Supplement to Volume 17: Our stock of this supplement is exhausted. Copies may be obtained by applying to Dr. E. C. Haderlie, U. S. Naval Post-Graduate School, Mon- terey, CA(lifornia) 93940. [Growth Rates, Depth Preference and Ecological Succes- sion of Some Sessile Marine Invertebrates in Monterey Harbor by Dr. E. C. Haderlie] Supplement to Volume 18: $1o.-. [The Biology of Chitons by Robin Burnett e¢ al.]. (Our supply of this supplement is exhausted; however, copies may be available by making application to the THE VELIGER Page 401 Secretary, Hopkins Marine Station, Pacific Grove, Cali- fornia 93950.) Prices subject to change without notice. Subscription rate to Volume 21 is $30.- plus postage. We must emphasize that under no condition can we ac- cept subscription orders or membership applications for calendar year periods. If “split volumes” are required, we must charge the individual number costs. Individual issues sell at prices ranging from US$12.- to US$24.-, depending on the cost to us. Backnumbers of the current volume will be mailed to new subscribers, as well as to those who renew late, on the first postal working day of the month following receipt of the remittance. The same policy applies to new members. THE VELIGER is not available on exchange from the Cali- fornia Malacozoological Society, Inc. Requests for re- prints should be addressed directly to the authors con- cerned. We do not maintain stocks of reprints and also cannot undertake to forward requests for reprints to the author(s) concerned. A Glossary of A Thousand-and-One Terms Used in Conchology by Winirrep H. ARNOLD originally published as a supplement to volume 7 of the Veliger has been reprinted and is now available from» The Shell Cabinet, Post Office Box 29, Falls Church, Virginia 22046, U. S. A. The cost is US$ 3.50 postpaid if remittance is sent with the order. WE ARE PLEASED to announce that an agreement has been entered into by the California Malacozoological Society, Inc. with Mr. Steven J. Long for the production and sale of microfiche reproductions of all out-of-print editions of the publications of the Society. The microfiches are available as negative films (printed matter ap- pearing white on black background), 105mm x 148mm and can be supplied immediately. The following is a list of items now ready: Volume 1: $1.50 Volume 2: $3.00 Volume 3: $3.00 Volume 6: $4.50 Volume 7: $6.00 Volume 8: $6.00 Volume 4: $4.50 Volume 10: $9.00 Volume 5: $4.50 Volume 11: $9.00 Volume 12: $9.00 Supplement to Volume 6: $1.50; to Volume 18: $3.00 California residents please add the appropriate amount for sales tax to the prices indicated. Page 402 Please, send your order, with check payable to Opistho- branch Newsletter, to Mr. Steven J. Long, P.O. Box 243, Santa Maria, CA 93454. Volumes and Supplements not listed as available in microfiche form are still available in original edition from Mr. Arthur C. West, P.O. Box 730, Oakhurst, CA(lifornia) 93644. Orders should be sent directly to Mr. West. WE CALL THE ATTENTION orf our foreign correspondents to the fact that bank drafts or checks on banks other than American banks are subject to a collection charge and that such remittances cannot be accepted as payment in full, unless sufficient overage is provided. Depending on the American banks on which drafts are made, such charges vary from a flat fee of $1.- to a percentage of the value of the draft, going as high as 33%. Therefore we recommend either International Postal Money Orders or bank drafts on the Berkeley Branch of United California Bank in Berkeley, California. This institution has agreed to honor such drafts without charge. UNESCO coupons are NOT acceptable except as indicated elsewhere in this section. Supplements Many of our members desire to receive all supplements published by the Society. Since heretofore we have sent supplements only on separate order, some members have missed the chance of obtaining their copies through over- sight or because of absence from home. It has been sug- gested to us that we should accept “standing orders” from individuals to include all supplements published in the future. After careful consideration we have agreed to the proposal. We will accept written requests from individuals to place their names on our list to receive all future sup- plements upon publication; we will enclose our invoice at the same time. The members’ only obligation will be to pay promptly upon receipt of the invoice. Requests to be placed on this special mailing list should be sent to Dr. George V. Shkurkin, Manager, 1332 Spruce Street, California 94709 However, until further notice, we are suspending the pub- lication of supplements until it will be reasonably certain that we will not be forced to spend many hours in tracing of lost insured or registered parcels and entering claims for indemnification. The special mailing list of members and subscribers who have entered: an “including all sup- THE VELIGER Vol. 20; No. 4 plements” will be preserved because of our innate opti- mism that sometime within our lifetime the postal services throughout the world will return to the former excellent and reliable performance. Claims for defective or missing pages must reach us within 60 days from the publication date. We will not respond to claims of missing issues made less than 30 days by domestic addressees, or less than 60 days by foreign addressees after the publication date of our journal issues. This refusal is necessary as we have received an increasing number of “claims” as much as 6 months before the claimed issue was to be published. We wish to conserve our energy and the cost of postage and stationery for more productive purposes. We are willing to accept requests for expediting our journal via AIR MAIL; however, in that case we must ask for an additional payment of US$8.00 in all cases where the Veliger goes to domestic addresses, and a depos- it of US$18.00 for all foreign addresses (including PUAS). Of course, we will carry forward as a credit toward the postage charges of the following year any amount over the actually required postage charges. Moving? If your address is changed it will be important to notify us of the new address at least six weeks before the effective date, and not less than six weeks before our regular mailing dates. Because of a number of drastic changes in the regulations affecting second class mailing, there is now a sizeable charge to us on the returned copies as well as for our remailing to the new address. We are forced to ask our members and subscribers for reimbursement of these charges; further, because of increased costs in connection with the new mailing plate, we also must ask for reimbursement of that expense. Effective January 8, 1968 the following charges must be made: change of address -— $1.- change of address and re-mailing of a returned issue — $2.75 minimum, but not more than actual cost to us. Vol. 20; No. 4 THE VELIGER Page 403 We must emphasize that these charges cover only our actual expenses and do not include compensation for the extra work involved in re-packing and re-mailing returned copies. At present we are charged a minimum fee of $12.50 on each order for new addressograph plates. For this rea- son we hold off on our order until 6 weeks before mailing time, the very last moment possible. If, for any reason, a member or subscriber is unable to notify us in time and also is unable to make the proper arrangement with the Post Office for forwarding our journal, we will accept a notice of change of address, accompanied by the proper fee and a typed new address on a gummed label as late as 10 days before mailing time. We regret that we are absolutely unable to accept orders for changes of address on any other basis. In view of the probable further cur- tailment in the services provided by the Postal Service, we expect that before long we may have to increase these time intervals. CALIFORNIA MALACOZOOLOGICAL Society, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. Donors may designate the Fund to which their contribution is to be credited: Operating Fund (available for current production) ; Savings Fund (available only for specified purposes, such as publication of especially long and signi- ficant papers); Endowment Fund (the income from which is available. The principal is irrevocably dedicated to scientific and educational purposes). Unassigned dona- tions will be used according to greatest need. Contributions to the C. M.S., Inc. are deductible by donors as provided in section 170 of the Internal Revenue Code (for Federal income tax purposes). Bequests, lega- cies, gifts, devices are deductible for Federal estate and gift tax purposes under section 2055, 2106, and 2522 of the Code. The Treasurer of the C. M. S., Inc. will issue suitable receipts which may be used by Donors to substan- tiate their respective tax deductions. Membership open to individuals only - no institutional or society memberships. Please send for membership ap- plication forms to the Manager or the Editor. Membership renewals are due on or before April 15 each year. If renewal payments are made after April 15 but before March 15 of the following year, there will be a re-instatement fee of $1.-. Members whose dues pay- ments (including the re-instatement fee) have not been received by the latter date, will be dropped from the rolls of the Society. They may rejoin by paying a new initiation fee. The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. Endowment Fund In the face of continuous rises in the costs of printing and labor, the income from the Endowment Fund would materially aid in avoiding the need for repeated upward adjustments of the membership dues of the Society. It is the stated aim of the Society to disseminate new infor- mation in the field of malacology and conchology as widely as possible at the lowest cost possible. At a Regular Membership meeting of the Society in No- vember 1968 a policy was adopted which, it is hoped, will assist in building up the Endowment Fund of the Society. An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 000.- or more, an entire volume will be dedicated to the memory of the decedent. REGARDING POSTAL SERVICE Complaints regarding late arrival of our journal are in- creasing in number, steadily, continually. However, we very conscientiously dispatch our journal on the printed publication dates. What happens after deposition at the Post Office is, of course, beyond our control. From some of our members we have been able to construct a sort of probable delivery schedule. In general, within California, 8 days is usual; outside of California, the time lapse in- creases with the distance; the East Coast can consider a lapse of “only” two weeks as rapid service; 4 to 5 weeks are not uncommon. Foreign countries may count on a minimum of one month, six weeks being the more usual time requirement and over two months not rare! In view of the ever increasing difficulties in the postal service, it is essential that members and subscribers not only give us prompt and early notice of address changes, but that proper arrangement for forwarding of our jour- nal be made with the local post office (at the old address). Page 404 New Postage Rates The U. S. Postal Service has increased second class mail rates, effective on July 6, 1977. We will, however, not in- crease our charges for mailing The Veliger, but must insist that we are reimbursed in all cases for returned copies and for the expenses involved in remailing such copies to a new address. It is very important for our members to realize: a) the postal service will not forward any mail other than first class for more than go days, even though forwarding postage may be guaranteed by the addressee; and _b) it is totally impossible for us to make changes in addresses in less than 6 weeks. We must make an address change even if only one digit in the ZIP code is changed, and the cost to us is the same as for a completely new address. Under no circumstances are we able to supply free re- placement copies of issues that fail to reach their proper destination. However, we will ship by insured mail re- placement copies at half the announced single copy rate of the particular issue plus postage. We have developed a triple check system so that, if we say that a copy has been mailed, we are absolutely certain that we delivered that copy to the post office in Berkeley and on the date we indicate. From our experience with the loss of insured mail, we are tempted to suggest that subscribers figure on a 10% reserve fund for the purchase of replacement copies. The only alternative remaining would be for us to increase subscription rates and membership dues by at least 10%. This, however, does not seem quite fair to us as some of our subscribers in almost 20 years have never failed to receive their copies. On July 6 the rates for book parcels and the library rate have been increased. This necessitates that we must in- crease the postage charges on back volumes, supplements and individual back numbers. The charges stated must be increased by 20¢ for one item and by 8¢ for each addi- tional item. It has been announced by the Postal Service that no increase in postage rates would be asked during the cur- rent year; it was not stressed sufficiently that this applies only to the so-called first class mail. Second class mail rates are “phased,” that is, they are scheduled to be in- creased each year until the rates are sufficiently high to pay the actual cost of handling that type of mail. While it is true that the rates have been very low, it should be borne in mind that the original intent of the special low rates was to aid in the dissemination of knowledge. This philosophy has, seemingly, been abandoned. THE VELIGER Vol. 20; No. 4 The regulations pertaining to second class mailing re- quire “pre-sorting” of the mail which involves a large amount of time, especially if the total number of pieces is too small to warrant the employment of computeriza- tion. This requirement seems justified as long as the rates for second class matter remain substantially below those for first class matter. However, our members should be aware of the fact that postal regulations rule that second class matter can not be forwarded three months after an address change, even though the addressee guarantees forwarding postage (in contrast, first class mail, at least for the time being, is forwarded for one year and that without charge!) . Thus, issues mailed to the “old” address will be returned to the publisher if return postage is guaranteed at a rate that is considerably higher; we have been charged as much as $1.45 for such returned copies. There is also a charge of 25¢ for a postal notification of the new address. It must be obvious that we cannot keep absorbing such extra expenses and keep membership dues and subscription rate at the current low rate. We must ask for the wholehearted cooperation of all concerned to help us to hold the line against increases. Also, if a copy is returned we will, as in the past, advise the member of this fact and indicate the total costs incurred for which we must seek reimbursement. If this reimbursement is not made, we cannot continue to send future issues to the delinquent member. Membership will have to be considered as terminated and can be re-instated only upon payment of all arrears. We regret that this apparently hard rule is necessary, but we wish to continue publishing the Veliger — which will not be possible if these rules are not observed. Although the Postal Service continues to deteriorate con- sistently, the postal rates are increased by 30 to 100%. We are, therefore, forced to increase the handling charges as well as the postage charges on the subscription. The following rates will be in effect on all new subscriptions and subscription renewals as of December 28, 1975: $1.- on subscriptions and memberships in the U. S. A.; $2.50 on memberships and subscriptions to PUAS coun- tries (Mexico, Central and South America and Spain) ; $3.50 to all other foreign countries, including Canada. We wish to stress that we are NOT INCREASING either membership dues or subscription rates, in spite of increased printing costs. But at the same time, we wish to call the attention to our Endowment Fund, the income from which enables us, in part, to keep these charges at the established levels. Contributions (tax deductible in the U.S. A.) are always welcome. Vol. 20; No. 4 THE VELIGER Page 405 To Prospective Authors Postal Service seems to have deteriorated in many other countries as well as in the United States of America. Since we will absolutely not publish a paper unless the galley proofs have been corrected and returned by the authors, the slow surface mail service (a minimum of 6 weeks from European countries, 8 to 12 weeks from India and Africa) may make a delay in publication inevitable. We strongly urge that authors who have submitted papers to the Veli- ger make all necessary arrangements for expeditious read- ing of the proofs when received (we mail all proofs by air mail) and their prompt return by air mail also. Since we conscientiously reply to all letters we actually receive, and since we experience a constant loss in insured and registered mail pieces, we have come to the conclusion that if a correspondent does not receive an answer from us, this is due to the loss of either the inquiry or the reply. We have adopted the habit of repeating our inquiries if we do not receive a reply within a reasonable time, that is 6 weeks longer than fairly normal postal service might be expected to accomplish the routine work. But we can not reply if we have never received the inquiry. Because of some distressing experiences with the Postal Service in recent years, we now urge authors who wish to submit manuscripts to our journal to mail them as insured parcels, with insurance high enough to cover the complete replacement costs. Authors must be prepared to document these costs. If the replacement costs exceed $200.-, the manuscript should be sent by registered mail with additional insurance coverage (the maximum limit of insurance on parcel post is, at present, $200.-). We are unable to advise prospective authors in foreign countries and would urge them to make the necessary inquiries at their local post offices. We wish to remind prospective authors that we have announced some time ago that we will not acknowledge the receipt of a manuscript unless a self-addressed stamped envelope is enclosed (two International Postal Reply Coupons are required from addresses outside the U.S. A.). If correspondence is needed pertaining to a manu- script, we must expect prompt replies. If a manuscript is withdrawn by the author, sufficient postage for return by certified mail within the U.S.A. and by registered mail to other countries must be provided. We regret that we must insist on these conditions; however, the exorbitant in- creases in postal charges leave us no other choice. Some recent experiences induce us to emphasize that manuscripts must be in final form when they are sub- mitted to us. Corrections in galley proofs, other than errors of editor or typographer, must and will be charged to the author. Such changes may be apparently very simple, yet may require extensive resetting of many lines or even entire paragraphs. Also we wish to stress that the require- ment that all matter be double spaced, in easily legible form (not using exhausted typewriter ribbons!) applies to all portions of the manuscript — including figure explana- tions and the “Literature Cited” section. It may seem inappropriate to mention here, but again recent experience indicates the advisability of doing so: when writing to us, make absolutely certain that the cor- rect amount of postage is affixed and that a correct return address is given. The postal service will not forward mail pieces with insufficient postage and, if no return address is given, the piece will go to the “dead letter” office, in other words, it is destroyed. Regarding UNESGO Coupons We are unable to accept UNESCO coupons in payment, except at a charge of $4.25 (to reimburse us for the ex- penses involved in redeeming them) and at $0.95 per $1.- face value of the coupons (the amount that we will receive in exchange for the coupons). 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