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THE

VELIGER

A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

a Ee rSupyhEMENT peoT KECE/VED WHEW YolumE wyes BouwD ~ 3/19] 87

Volume 21

July 1, 1978 to April 1, 1979

Mars «hp syery Coe aN Ga

Vol. 21; No. 4

TABLE or CONTENTS

Abnormal callus development in Nautilus pompulius Royat H. Mapes, TERRENCE J. FREST & STEVEN

JOSSIOVSNS. cesveceertereine cermin erat aero coer 442

Abnormality of shell plates in three chitons from New England

DEA HEP SBICAIN GER Stern sects scene etnersavatiirestsareasjereeteeee 274

Accumulation of “C-labelled algal exudate by Mytilus californianus Conrad and Mytilus edulis Linnaeus. An aspect of interspecific competition P. V. FANKBONER, WM. M. BriayLock & M. E. DE AB RG Hip er ere eel ne Saha nratesste eosin esetanetantedcae 276 A comparative study of the structure, development and morphological relationships of chambered cephalo- pod shells KLAUS BANDEL & SIGURD V. BOLETZKY @cscsessensnins 313 Additional notes on Spurilla alba (Risbec, 1928) (Mollus- ca : Opisthobranchia ) GAVE EGE SPRHON (ise ce Seccscctecttna sit pcctescbathntasbetiatsonba 305 Additional notes on the food of some California nudi- branchs with a summary of known food habits of California species Gary R. McDona.p « JAMES W. NYBAKKEN ..... 110 Aestivating giant African snail population in South Anda- man during 1973, 1974 and 1975 G. P. Gupta, S. 8. S. Gautam, S. R. ABBAS &

IPP) SRIVASTAVA. ceo ceccascmaancsuesatnciatestcnenn 135 A fossil Haliotis from the Galapagos Islands ANN ATT DU REPAM oh cecececsecsescccaseeetntttntsesnctiesrsuscnesita 369 A new Panamic Mitrella (Mollusca : Gastropoda) WILLIAM PItr & ROY KOR on. ccsssesssscsnencsseneensecemeene 467

A new species record for Mytilopsis sallet (Récluz) in central America (Mollusca : Pelecypoda) Dan C. Mareu & Rosert E. BERREND .............. 144 Antipredator behavior in Octopus dofleint (Wiilker) B. Hartwick, G. THoRARINSSON & L. TULLOCH 263 A range extension of Anachis lillianae

18s Je SNA = 04 07 Dy Cote eer en mn Use a EN 402 California’s Cretaceous Haliotis BJ WVATI DURA SE se esc cetesnretericemermnnsenietas 373

Checklist of marine mollusks at Coyote Point Park, San Francisco Bay, California : INVAR YO KG? WICKSTEN: foo ccdic cree tntencsnnetenarneniiers 127 Chlamydoconcha orcutti Dall: a review and distribution of a little-known bivalve

NJANCES HE: CARETON( sneer seins nee eee ae, 375 Cypraea goodalliu Sowerby, 1832 on Fanning Island FLU Gite: BRADNER 2st ancien, eure a on 306

Description of a previously misidentified species of Epi- tonium (Gastropoda : Epitoniidae) ELE REINS DUS EVAN Es creer cise rece nea cca erases 379

THE VELIGER

Page iii

Distributional patterns of juvenile Mytilus edulis and My- tilus californianus RETERYOs PETRATIIS (ioe 2 see cieeiasosa. wees peae ere cece: 288 Egg capsule and young of the gastropod Beringius beringii (Middendorff) (Neptuneidae) RICHARD A. MACINTOSH. ....-ccsscsnsscuesesesitisseeneissitintsse 439 Evidence for an additional Littorina species and a sum- mary of the reproductive biology of Littorina from California SDALBOT | MURRAY ann pects rte cere erorna aee 469 First record of Okenia impexa Marcus, 1957 from the western Atlantic in the Mediterranean IL WISE SCH MERE D thi, aincinsioniss.nsesnrsenneient se ister » 355 Flight responses of three congeneric species of intertidal gastropods (Prosobranchia : Neritidae) to sym- patric predatory gastropods from Barbados Danret L. HorrMan, WiLuiaM C. Homan, Jay SWANSON & PAUL WELDON. ....cscsssscsssssetnsseses 293 Food preferences, food availability and food resource par- titioning in two sympatric species of cephalaspid- ean opisthobranchs Davm SHONMAN & JAMES W. NYBAKKEN ...........0.0 120 Growth in the keyhole limpet Fissurella crassa Lamarck

(Mollusca : Archaeogastropoda) in northern

Chile MAR TAM BRETOS ccstdicestscossst sti tobsccscccecs tterstosstesncsonsitirensine 268 History of the Pliocene molluscan fauna of northern Japan BRAIN KEK MER oer ee colette 227

Infection of Ostrea lurida and Mytilus edulis by the para- sitic copepod Mytilicola orientalis in San Francisco Bay, California WILLIAM BRADLEY & A. E. SIEBERT, JI. sessssssesnsine 131 Laboratory cultivation of Haminoea solitaria (Say, 1822) and Elysia chlorotica (Gould, 1870) June FE Harrican & DanteL L. ALKON .........0 299 Laboratory culture, metamorphosis and development of Aplysia brasiliana Rang, 1828 (Gastropoda : Opisthobranchia ) Nep E. STRENTH & JAMES E. BLANKENSHIP .......... 99 Late Neogene succession of molluscan fauna on the Pacific coast of Southwestern Japan, with reference to planktonic foraminiferal sequence

RYUICHI TSUCHI & MASAKO IBARAKEI oe nsesscscstssninie 216

Late Oligocene through Pleistocene molluscan faunas in the Gulf of Alaska Region

IRYGHVARD) | Cie ALAS ON ssterercresromcctrecnsteresccsarstrevicevarecrcrias 171

Malagarion paenelimax gen. nov., spec. nov., a new slug- like helicarionid from Madagascar (Pulmonata:

Helicarionidae) PS) 0 (ODS B10 6) B18 a eer eve Re 361 Mating behavior of Octopus joubini Robson NJENINIRER UAL) MATHER oscissccstsceteseticouscscctectstecencorsancns 265

Page iv

Nematodes in the alimentary canal of terrestrial slugs Dario TE. CAPPUCGE, ris ccccbe ese tetaseees 306 Neogene molluscan faunas in the Japanese Islands: An ecologic and zoogeographic synthesis

ISTVOTAIKA, CHIN ZI MO pee i ee Me eye teneetoace 155 Neogene Pectinidae of the northern Pacific KOICHIRG MASUDA Tee ie es earn 197

Neptunea (Gastropoda : Buccinacea) in the Neogene of the North Pacific and adjacent Bering Sea CriprorD)/ Mi NELSON) ee ee aeraeats 203 Notes on the cephalopods of Menterey Bay, California, with new records for the area MEO ERIC ANDERSON ( set schetdi rcs ncssenecttatsrsetuentatctcn acts 255

Notes on the reproductive strategies of the south African .

vermetid gastropods Dendropoma corallinaceum and Serpulorbis natalensis

ROGER Ni HUGHES | eee eee cansantscsnesnaen 423 Notes on the winter epiphragm of Pupoides albilabris M. CHRISTOPHER BARNHART. ecsssssssssssmtussessenenanses 400

On the relevance of smail gastropod shells to competing hermit crab species

DENIS WANG & DAVID A. JILLSON.ecsssssssessssnssnnemss 488

Papers on Neogene mollusks of the North Pacific Margin: an introduction

WARREN, ©) ADDIGOTI 22.0 oir cnscosescsneonsntbesenssacttseanes 153

Premises of Neogene correlation in the northern part of the circum-Pacific

VUREB: GreADENKOV, 2p. cieecsaanssnnnsnintacnstamasiremcass 225 Protoconch of ovoviviparous volutes of West Africa

PRWITA)/BRATCHERY 20g bce tu cates acastacelt celine 144 Possible predation on Nautilus pompilius

Joun K. Tucker & ROYAL H. MAPES oes 95

Rearing experiments on the California market squid Loligo opalescens Berry, 1911 RoceEr T. Hanon, RaymMonp FE Hrxon, WiLuiaM H. Huet & WON TACK YANG ...necseceennen 428 Recognition of Cyclocardia ovata (Riabinina, 1952) in the eastern Pacific BUGENE/| Vii COAN) eli a oh Aan Ss ee ae 488 Reproductive biology of Colus stimpsoni (Prosobranchia : Buccinidae). II. Spermiogenesis DAVID IL. WEST ie cat a es ccs Oe I Reproductive biology of Colus stimpsoni (Prosobranchia : Buccinidae). III. Female genital system Davin iE.) WEST aii teeee ee Ler ce ns 432 Reviews of biology of commercially important squids in Japanese and adjacent waters I. Symplectoteu- this oualaniensis (Lesson)

TAKASHI OKUTANI & IH-HSIU TUNG ose 87 |

THE VELIGER

Vol. 21; No. 4

Review of the bivalve genus Pholadomya from the Terti- ary of California and the description of two new species

WILLIAM J. ZINSMEISTER cessesscsesersssueueneseenuetntutntnenee 232

Selection and turnover of coelenterate nematocysts in

some aeolid nudibranchs Rosert MarsHALt Day & Larry G. Harris ......... 104

Sexual characteristics of Margaritifera margaritifera

(Linnaeus) populations in central New England

Dovuetas. G. SMirm 220. eee 381 Soviet contributions to malacology in 1977 KENNETH J. Boss & Morris K. JACOBSON otescnene 490

Studies on the Mytilus edulis community in Alamitos Bay, California: VII. The influence of water-soluble petroleum hydrocarbons on byssal thread formation

Rosert Scott Carr & DONALD J. REISH ........00- 283

Temporal changes in a tropical rocky shore snail commu- nity

TOM Mi. SPIGEIT ciceeccscecitenenorsncetternnreenec scence 137

The Chromodoridinae nudibranchs from the Pacific coast of America. Part III. The genera Chromolaich- ma and Mexichromis

HANS BERTSCE (0 ssecccointetctenecueit ene ae 70

The Chromodoridinae nudibranchs from the Pacific coast

of America. Part IV. The genus Hypselodoris FIANS. BERTSOHD oo. scccctosctunustssssanis sec nee eee 236

The digestive diverticula of Alderia modesta and Elysia chlorotica (Opisthobranchia : Sacoglossa)

Desra A. Graves, M. A. Gisson & J. S. BLEAKNEY Peat amr MUMIA Pe ue eA aT AME MEN eres 415

The effects of an ectoparasitic gastropod, Caledoniella montrouzieri, upon molting and reproduction of a stomatopod crustacean, Gonodactylus viridis

MAR JORIE LINDQUIST REAKA osnsssssesenneciescteenesenee 251

The epibiota of Arca zebra and Arca imbricata: a com-

munity analysis THOMAS B. SCANLAND. \.essssessmrssnensnsnsnesisnetuenenseneetin 475

The genus Callistochiton Dall, 1879 (Mollusca : Poly- placophora) in the eastern Pacific, with the de- scription of a new species

ANTONIO, J: FERREIRA, (cece: lcscnecesentae serena 444

The genus Lepidozona (Mollusca : Polyplacophora) in the temperate eastern Pacific, Baja California to Alaska, with the description of a new species

ANTONIO: J.) PERRETRA Son ete) es aeons 19

The population dynamics of two sympatric species of Macoma (Mollusca : Bivalvia)

Joun Gipson Rar TT coe eae ene 384

Vol. 21; No. 4

The ultrastructure and evolutionary significance of the cerebral ocelli of Mytilus edulis, the bay mussel Mark Davm Rosen, Cuartes R. STASEK & Coin O. ETE REA S Qi seater cote deebcccs sient hsde a scetoeseesiatees 10 Ultrastructure cf the mosaicostracal layer in the shell of the bivalve Mytilus edulis MELBOURNE R. CARRIKER essssscscsssssiistssneenessninint 4II Viability of sperm in two land snails, Achatina fulica Bowdich and Macrochlamys indica Godwin-Austin

Ste AUT &p Kel Gs GHOSE face cantina tomes 486 Wharf piling fauna and flora in Monterey Harbor, Cali- fornia Eucene C. HaDERLIE & WINFIELD Donat, III ..... 45 Winter reproduction in the gastropod Nassarius trivittatus SPAIN ANSBEDE CED NIKO este tee cee ecesheteoncicttorasscesatetstessat 207 World-wide biostratigraphic correlation based on turritel- lid phylogeny IVAN TO BIO DATCA i cServcsecasc setscetenevisdlvetonectanstecectestnit 189

AUTHOR INDEX

Assas, 8. R. see Gupta, G. P. et al.

PADDICOTT) | WARREN Ooo cccsscscresrtecsesnecesnsetesnestoe 153, (408) ALKon, Dantet L. see HARRIGAN, JUNE FE & PATTISON MRIGHIARD: Cig, tenn ce ctte cre ots tee tc cceareteaeetecmusisincee 171 DONIDER SONG | EsRUG tices cts estsvntissncnseasssnetesrteatns 255 ARONOFF, STEVEN M. see Mapes, Royat H. et al. BANDEL, KLAUS & SIGURD V. BOLETZRY ...-seccerenssseeuee 313 BARNHART, M. CHRISTOPHER .ecscsscssssssssussssssiststtnensusnt 400 BERREND, Rospert E. see Mareu, Dan C. & IBERTSCHAGETANS ini recreate anes Paves 70, 236

BLANKENSHIP, JAMES E. see STRENTH, Neb E. & BLayLock, WiLLIAM M. see FANKBONER, P. V, et al. BLEAKNEY, J. S. see Graves, Desra A. et al. BOLETZKY, SIGURD v. see BANDEL, KLAUS &

Boss, KENNETH JAY & Morris K. JACOBSON ........c:u0 490 BRADLEY, WILLIAM & A. E. SIEBERT, JI. osssssecssnnsnsnesee 131 BRADNER FLU GH ys ciccess tte ct tes tac eee aera aN cates 306 BRAT CHER CD WITA sche ath t atin etn nm eaves eens 144 IBRETOSN PART AN cect hl Bee Sehr Goto an lohan ae 268 CAPPU GCI) DARIO! et [Tse cee etree erect ne ad 306 CARLTON, JAMES To wecscscsnseeeteen cae em 375 Carr, Rosert Scott & Donatp J. ReisH ea ee 283 C@ARRIKER) MELBOURNE Re eee cenit eeracettiaens All GETINZET: PISTYOTAICA petra ene ee eae re ecb 155 GOAN IUGENE Vi oon encicnnc ere en ode wee. 488 Day, Robert MarsHALt & Larry G. Harris ........... 104

DE Burcu, M. E. see FANKBONER, P V, et al.

THE VELIGER

Page v

Donat III, Winrretp see Hapverui, Evcene C. «

DuR TAME SWAT tedcucnunimemen noe, 369, 373 DU SHANE i IEEE NG cote sare sscnspeten iio incebioatte ihiideocaboacte 379 FANKBONER, P. V, Wm. M. Biaytock « M. E. pe Burcu

sere teh rrr cece ne te See 276

FERREIRA, ANTONIO Jo escssssssnsscsne FREST, TERRENCE J. see Mapes, Roya H. al. Gautam, S. 8. S. see Gupta, G. P et al.

GuoseE, K.C. see Raut, S. K. &

Greson, M. A. see Graves, Desra A. et al.

GUADEN OV: V\URT! Based tena tecritar ttn cnaresnontante merits 225 Graves, Desra A., M. A. Grson & J. S. BLEAKNEY 415 Gupta, G. P, S. S. S. Gautam, S. R. Appas & PD.

SRIVASTAVA tratistesrcoeracaraia seracos crane anomie nent 135 HaperLeE, Evcene C. & WinFiELp Donat III .......... 45 Hanon, Rocer T, RaymMonp F Hixon, Wituam H.

HU LET & WON TACK YANG uncctcsicssssrsnseseitienns 428 HARRIGAN, JUNE F & DANIEL ALKON oeescsscsntiesitsntceie 299

Harris, Larry G. see Day, Ropert MARSHALL & Hartwick, B., G. THorartnsson & L. TuLtocH 263 Hermans, Cotin O. see Rosen, Marc Davw et al. FICK MAN, CAROLE) Si jue toniscasetias sane tance (506) Hrxon, RaymMonp F. see Hanton, Rocer T. et al. HoFFrMan, Danrt L., Wittiam C. Homan, Jay Swan-

SON: @ DAUE J.) WELDON) cise sets cherotesccranrcs 293 Homan, WiiuaMm C. see HorrMan, Danret L. al. BAW GHES ROGER Nic accesses cesta atteaatie ome tier re 423

Huet, Wiiuiam H. see HAnton, Rocer T. al. IBARAKI, Masako see TSucHI, RyvicHI & Jacosson, Morris K. see Boss, KENNETH J. e— Jmtson, Davm A. see Wane, DENIS &

KSEE NAME MV RADI artnet caer SU Yak ea a coho IKSTEEMIE RAV BIRAIN Ko Ed sy oercscen cise trcrstse ter ct see veascencersare Kout, Roy see Prrt, WILLIAM & Kotaka, TAMIO Lancer, Paut D.

IMPACIIN TOS HY, RICHARD (AG $cc stccectectectsotecns dt ecneetcercnes 439 Mapes, Royat H., TERRENCE FREST & STEVEN M. PNRONOEE. rtecs et teee ertsdeac ecco fncteoteasoatssbbatcacosaee 442 Mapes, Royat H. see Tucker, JoHn K. & Mare. Dan C. & ROBERT E. BERREND. .csoscsscscsssnces 144 MasupaA, KéicHiR6 IMA TELE Ss JENIN TR ER) Psy eso srncnserteresc sin eeeaseatsetrsattericceeies 265 McDonatp, Gary R. & James W. NYBAKKEN ......... 110 INET R RA eCAT BO Tig teeta eset astncstlacoestencantynprgnaaieanvesic 469

INEE'SON@ 1 G:LTEFORD VMs encores rte senrceorestecocssnentes 203 NYBAKKEN, JAMES W. see McDona.p, Gary R. & see also SHONMAN, Davp &

OKUTANI, TAKASHI & IH-HSIU TUNG ounce 87 PECHENTR: PANG Ai orc stertsctasseu ccssaatastaarsuratvanounnciebiaaraniss 207 IPETRIATIISS PEE DER Serer ett eee or eee 288

Page vi

Print, WinliAMie Roy KOHL oer 467 RAr Mt, JOHN GIBSON) cineca nce calematees 384 Raum, S) Kee Koy GHOSE nee tee eens 486 REAKA, MARJORIE LINDQUIST nesessssssssusstuseseenteneesnctnessses 251

ReisH, Donatp J. see Carr, Rospert Scotr & Rosen, Marc Davm, CuHartes R. STASEK & Coin O.

EVER MANS 3 5c10 cid cadre cestattns mteaonrnnnont noone Co) ROTH, BARRY. o.sssssnsssineen (408) SCANEAND; THOMAS) Bi 2 oe o2 a Soscsnsnneneneamenn oe 475 SCHMEKEL, UWNGUISE (otic a niacin mamta unmet teneeen 355 SHONMAN, Davm & JAMES W, NYBAKKEN ooscsssassneee 120 SIEBERT, Jr., A. E. see BRADLEY, WILLIAM & SMITH; DOUGEAS) Gioia wad eM coher ea 381 SPHON, GALE G. ......... we 305) SPIGHT, TON Mi tiiirea tei ee chateau oe 137

Srivastava, PD. see Gupta, G. P. é¢ al. STASEK, CHARLES R. see RosEN, Marc Davo et al.

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Vol. 21; No. 4

STOHLER, Re woes ae (150), (152), (409), (410) STRENTH, Neb E. & JAMES E. BLANKENSHIP... 99 Swanson, Jay see Horrman, Dani L. et al. THorRARINSSON, G. see Hartwick, B. et al.

Ticuier; SIMON Wu 361 TSUCHI, RYUICHI & MASAKO IBARAKLT oosssssssssssssssssessneseee 216 TuckeER, JOHN K. & ROYAL H. MAPES ...cssecnee 95 Tuuiocu, L. see Hartwick, B. al.

Tune, In-Hsrtu see OKUTANI, TAKASHI &

WANG, DENIS & DAVID A. JILLSON oecscsassncsnssneinensintne 488 WELDON, Paut J. see HorrMan, Dantet L. al. WEST): DAVID Ts) 2 iiiieitssdeusecmeapameante cease I, 432 WHITNEY, R. A.

WICKSTEN, MARY. Koo iccnccsunscccnnccnimieneneten ee 127 Yanc, Won Tack see HANLon, Rocer T. é al. ZINSMEISTER, WILLIAM J. <..ccnccsccenoneee eee 232

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THE

VELIGER

A Qvorterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California f VOLUME 21 JuLy 1, 1978 NuMBER 1

CoNTENTS

Reproductive Biology of Colus stimpsoni (Prosobranchia : Buccinidae). II. Spermio-

genesis. (3 Plates) H DAVID BIG VVES TamC ere Acne Niue a Ie elite oe fetes 6) ) lal lg VE The Ultrastructure and Evolutionary Significance of the Cerebral Ocelli of Mytilus edulis, the Bay Mussel. (4 Plates; 3 Text figures) Marc Davm Rosen, CHaries R. STaSEK & Couin O. HERMANS . « » « «© ~ IO

The Genus Lepidozona (Mollusca : Polyplacophora) in the Temperate Eastern Pacific, Baja California to Alaska, with the Description of a New Species.

(5 Plates; 3 Text figures)

ANTONIO J. FERREIRA eae siaiee Lei esis ceuaad ies atl Cae Mele oy Soler hepa (98 19 Wharf Piling Fauna and Flora in Monterey Harbor, California. (1 Plate; 10 Text figures) Eucene C. Haperym & WinrietD DonaTIII . . . . . 1 eee ee ee AS The Chromodoridinae Nudibranchs from the Pacific Coast of America. Part ITI. The Genera Chromolaichma and Mexichromis. (2 Plates; Text figures 16 - 25) RIANSEDERTSCH ret atic! aaa. a. edocs lee ae ele «ee en 6, 90 Reviews of Biology of Commercially Important Squids in Japanese and Adjacent Waters I. Symplectoteuthis oualaniensis (Lesson) (7 Text figures) PAKASHHOKUTANIS LH-FISIU: GUNG =e (5 mela Me el ee ee ew 87 Possible Predation on Nautilus pompilius. (1 Plate; 1 Text figure) OHNIKG TUCKERIa ROYATVHGMAPES) 005. 2s. ee i ew | 85 Laboratory Culture, Metamorphosis and Development of Aplysia brasiliana Rang, 1828 (Gastropoda : Opisthobranchia). (1 Text figure) Nep E. STRENTH & JAMES E. BLANKENSHIP . . . . - © «© © «© © «© © « 99

[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

Second Class Postage Paid at Berkeley, California

ConTENTS Continued

Selection and Turnover of Coelenterate Nematocysts in Some Aeolid Nudibranchs. Rosert MarsHa.i Day « Larry G. Harris ... . 6 BG Valo Go. NOM

Additional Notes on the Food of Some California Nudibranchs with a Summary of Known Food Habits of California Species.

Gary R. McDonatp & James W.NypaKKEN . . . . . ~ - + + «© 6 © « 110 Food Preferences, Food Availability and Food Resource Partitioning in Two Sym- patric Species of Cephalaspidean Opisthobranchs. (1 Text figure) Davi SHONMAN & JaMES W.NYBAKKEN. . . . . 7 + 7 + + ee ee E20 Checklist of Marine Mollusks at Coyote Point Park, San Francisco Bay, California (1 Text figure) Mary K. WicksTEN. . . ge aa HS cr a , Pe mi 6), fou 027)

Infection of Ostrea lurida and cont edulis by the Parasitic Chew Mytilcola orientalis in San Francisco Bay, California.

Wirviam Brapiey A. EB. Sresert, Jr) 26 ee ee Aestivating Giant African Snail Population in South Andaman During 1973, 1974 and 1975. G. P. Gupta, S. S. S. Gautam, S. R. ABBAS & P D. SRIVASTAVA . . «. «© © «© © 135 Temporal Changes in a Tropical Rocky Shore Snaii Community. (1 Text figure) Tom M. 'SPIGHT i aie 8 ee een Be a ok. CaS Yoon UNC earn ST NOTES & NEWS) 32) 3. PS ie earen eG Met AIG or ui. Udidt A New Species Record for Renee sallei (Récluz) in Central America (Mollusca : Pelecypoda). Dan C. Maret & RoBert E. BERREND Protoconch of Ovoviviparous Volutes of West Africa. (1 Text figure) TwiLa BRATCHER BOOKS, PERIODICALS & PAMPHLETS ......... » «= « «= « 150

Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc.

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Vol. 21; No. 1

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Reproductive Biology of Colus stimpsoni

(Prosobranchia : Buccinidae)

II. Spermiogenesis '

DAVID L. WEST?

(3 Plates)

INTRODUCTION

MOoLLUSCAN SPERMATOZOA, like most sperm, are highly specialized for motility and the other activities involved in fertilization of the ova. Rerzius (1906), in his splendid monograph, described and illustrated the male gametes for a number of different gastropods. His survey indicates that there is considerable heterogeneity in size and shape of gastropod spermatozoa. FRANZEN (1955) investigated a great variety of mollusks and concluded that the ob- served morphological modifications are related to the dif- ferences in the media into which the spermatozoa are dis- charged and fertilization occurs. He (FRANZEN, 1955, 1970) suggested that in those molluscan groups which have external fertilization, the spermatozoa are less mor- phologically specialized (primitive sperm type) than in those groups with internal fertilization (modified types of spermatozoa).

Most neogastropod spermatozoa, particularly those in- volved in fertilization, are thread-like in shape and are considered as a modified type which is associated with internal fertilization (FRANZEN, 1955, 1970; FAWCETT, 1970). Many investigators have contributed to the under- standing of the structure, genesis, and functioning of subcellular components of molluscan spermatozoa (for more extensive literature, see Tuzet, 1930; Dupovy, 1964; RoosEN-RuNGE, 1969; THOMPSON, 1973; ANDER- SON & PERSONNE, 1970). However, many questions con- cerning these aspects of spermatozoa still remain.

An interesting phenomenon occurring in prosobranchs is sperm dimorphism; that is 2 or more morphologically distinct types of spermatozoa simultaneously produced by an individual male. Since von StrBoLp (1836) described this phenomenon, many workers have investigated the

* Contribution No. 58 from the Marine Science Institute, North- eastern University, Nahant, Massachusetts

* Present address: Center for Pathobiology, University of Cali- fornia, Irvine, Irvine, California 92717

genesis and function of these spermatozoa (PoRTMANN, 1930; TuzET, 1930; Dupouy, 1964; Tocurmoto, 1967; NIsHIWAKI, 1964), and a variety of terms has been used for their designation. MEvEs (1903) termed those sperma- tozoa which contain the normal complement of chroma- tin, “eupyrene”; those spermatozoa which have a smaller quantity of chromatin, “oligopyrene”; and those which have no chromatin, “apyrene.” KuSCHAKEWITSCH (1913) later shortened the terminology to “typical” for the nor- mal, eupyrene spermatozoa and “‘atypical” for all abnor- mal, oligopyrene and apyrene spermatozoa.

Atypical prosobranch spermatozoa exhibit various shapes and sizes, ranging from small worm-like forms to the giant, multiflagellated spermiozeugma. NISHIWAKI (1964) classified atypical spermatozoa into 8 morphologi- cal categories. TocHimoto (1967) cytochemically ex- amined a number of examples representing these catego- ries and added one additional type of atypical spermato- zoa, the “‘free cell” of the Littorinidae.

A few workers (ANKEL, 1924; PORTMANN, 1926, 1927, 1930, 1931; WoopwarD, 1940; BULNHEIM, 1962; REIN- KE, 1914) have dealt with the function of atypical sperma- tozoa, and some have suggested that they may serve a nurse-cell function (Woopwarb, op. cit.). Others (Port- MAN, opp. cit.; Dupouy, 1964) have proposed that the atypical spermatozoa play a role in the determination of nutritive eggs (2. e., those ova which are deposited in a single egg capsule but abort in early embryogenesis and serve as food for the developing young). However, not all prosobranchs that exhibit sperm dimorphism utilize nut- ritive eggs, nor do all those prosobranchs that utilize nutri- tive eggs exhibit sperm dimorphism (HyMav, 1967; FRET- TER & GRAHAM, 1962). PoRTMANN (1931) suggested that in those species which do not exhibit a morphological sperm dimorphism but utilize nutritive eggs in the course of their development, for example Thais lapillus (Linnaeus, 1758), have a “physiological” sperm dimorphism. This dimorphism is a result of chromosome elimination in a portion of the gametes during meiosis (oligopyrene sperm)

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Vol. 21; No. 1

and fertilization of ova by these spermatozoa results in abortive embryos which then serve as a food source. STAIGER (1950, 1951) and THoMpPSON (1973) indicate that atypical spermatozoa are incapable of participating in egg-penetration or amphimixis.

Members of the neogastropod genus Colus utilize nu- tritive eggs (THORSON, 1935, 1940; RADWIN & CHAMBER- LIN, 1973; WEST, 1973) but little is known about the reproductive biology of this genus. The present study re- ports some ultrastructural and cytochemical findings on spermiogenesis in a species of Colus to further the know- ledge of this genus and to add to the body of information concerning molluscan spermatozoa. ;

MATERIALS anp METHODS

Mature Colus stimpsoni (Morch, 1867) were collected in- tertidally at Cobscook State Park, Edmunds, Maine, and at Eastport, Maine, and maintained in running sea water aquaria at the Marine Science Institute, Northeastern U- niversity, Nahant, Massachusetts. For light microscopy tes- tes were excised from freshly opened, unrelaxed snails and fixed for 1-24 hours in the following: (a) Hollande- Bouin’s; (b) Bouin’s; (c) sea water Bouin’s (WALKER & MacGrecor, 1968); (d) buffered formalin (pH 7.4), or (e) buffered glutaraldehyde (pH 7.4). Following fixation, tissues were dehydrated through a graded series of ethan- ols or acetones and embedded in polyester wax (STEED- MAN, 1960). Sections were cut 4 - 6m in thickness and stained with the following: (a) Heidenhain’s azan, (b) Heidenhain’s iron hematoxylin, (c) GaBE’s (1968) modification of Gomori’s trichrome, (d) the Feulgen technique, or (e) the periodic acid-Schiff (PAS) tech- nique (Humason, 1967). LEHMAN’s (1965) polychrome was used for localizing various molecular groups. Permanent sperm smears were prepared by diluting sperm from the seminal vesicle with sea water and placing a few drops of this suspension on glass slides. The slides were inverted over formalin vapors for 1-2 hours and subsequently washed briefly with distilled water. The Feul- gen reaction (HuMAsoN, 1967) was used to localize the

sperm head, and Altmann’s aniline-fuchsin technique (HuMASON, op. cit.) was employed to localize the middle piece. The PAS technique was used to determine the presence and distribution of carbohydrates, particularly glycogen. Observations on living sperm and spermatogenic cells from freshly prepared testis smears were made with Nomarski differential-interference optics.

Nuclear proteins were stained with alkaline (pH 8.3) fast green (HuMason, 1967), alkaline (pH 8.3) eosin Y (Biocu « Hew, 1960) or bromphenol blue adjusted to pH 2.3 with acetic acid (BLocH « Hew, op. cit.). Prior to staining, nucleic acids were removed from tissue sec- tions with trichloroacetic acid at 99°C for 30 minutes or with a saturated aqueous solution of picric acid at 60°C for 24 hours (BLocu & Hew, op. cit.). These acid hydrol- ysis treatments usually resulted in the sections detaching from the slides. Consequently, slides were coated with a thin coat of celloidin following removal of wax from the sections. Protein end groups were blocked by deamination (after removal of DNA by acid hydrolysis) with Van Slyke’s reagent (HuMASON, op. cit.; BLocH « HEw, op. cit.) or by acetylation (HuMASON, op. cit.).

For electron microscopy, testes and segments of seminal vesicles were excised from freshly opened, unrelaxed snails and treated in accordance with the methods described in the first paper of this series (WeEsT, 1978).

OBSERVATIONS

The cellular elements and the organization of the male reproductive system were considered in detail in the first paper of this series (WEsT, 1978), and only the salient features of the genital system will be considered here. The testis lies on the distal portion of the visceral mass and consists of numerous seminiferous tubules (100 - 800 um in diameter) which are separated from each other by a thin stroma. These tubules join to form the single vas deferens which passes along the columellar side of the body. The posterior portion of the vas deferens functions as the seminal vesicle and is filled with spermatozoa throughout the year.

Explanation of Figures 1 to 5

Figure 1: Spermatogonia (S) clustered near basal cell (B) X 5000 Figure 2: Primary spermatocyte. Arrows centrioles; M mito- chondria; G Golgi complex X 8500 Figure 3: Early spermatid showing polar nucleoplasm (P) with small granules (arrows) and small patches of nucleoplasm X 11 goo Figure 4: Early spermatid at early flagellar tube formation. Arrows

small tubules within flagellar tube; B - layer of condensed chro- matin; C centriole; P - polar nucleoplasmic cone X 18 600 Figure 5: Crescent-shaped spermatid. A early acrosome with attached band of electron-dense material; B layer of condensed chromatin; C centriole; M mitochondrion; P flagellar tube projection; T tubules within flagellar tube X 18 600

[West] Figures 1 to 5

Tue VeE.icErR, Vol. 21, No. 1

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On the periphery of each seminiferous tubule, beneath the stroma, a layer of lipid-rich basal cells surrounds 2 types of accessory cells and the spermatogenic cells. Sper- matogenic cells are generally clustered in small groups which are in the same maturation phase, and these groups of cells are irregularly distributed within the tubules. Spermatozoa generally fill the lumen of the tubule, but they also occur scattered between spermatogonia and spermatocytes.

Spermatogonia and Spermatocytes

Spermatogonia are difficult to distinguish from early spermatocytes. However, they stain intensely lavender with Lehman’s polychrome stain and have irregularly shaped nuclei (Figure 1). Groups of spermatogonia (8 to 24 cells per group) generally occur near the periphery of seminiferous tubules, lying just beneath the layer of basal cells (Figure 1). Gonial cells measure 8 - 11 ym in diameter, and the nucleus (7 - 9 um in diameter) occupies nearly the entire cell volume. The chromatin is unevenly distributed, giving the nucleus a patchy appearance, and the nuclear envelope is perforated by numerous, small pores. The single nucleolus is homogeneous and eccentri- cally located. Distributed within the moderately electron- dense nucleoplasm are numerous small granules. The scanty cytoplasm contains a few relatively long cisternae of rough endoplasmic reticulum and numerous, free ribo- somes.

At the onset of spermatogenesis, spermatogonia increase in size to about 15 4m in diameter, giving rise to primary spermatocytes. Primary spermatocytes (Figure 2) have a small amount of cytoplasm surrounding the subspherical nucleus (11 - 13m in diameter) and, in addition to the elements found in spermatogonia, the cytoplasm contains several juxta-nuclear mitochondria. Situated near these mitochondria are 2 centrioles and a prominent Golgi complex (Figure 2). The chromatin is condensed and synaptonemal complexes which are concomitant with syn- apsed chromosomes (PARCHMAN & RorH, 1971) are frequently observed. First and second meiotic divisions apparently occur in rapid succession, and secondary sper- matocytes are rarely seen.

Spermiogenesis

Nucleus: Following the second meiotic division, the nuclear material of the early spermatid is coarsely granu- Jar and irregularly distributed throughout the nucleus, leaving small, irregular patches of granular nucleoplasm. Occasionally, cytokinesis is not complete and 2 or 4 early spermatids are joined by cytoplasmic bridges. However, later spermatids are not joined. As spermiogenesis pro-

ceeds, the nucleus increases in size, to about 7m in diameter, and is irregular in outline. The coarsely granular chromatin is irregularly distributed within the nucleus, leaving a large area of nucleoplasm (polar nucleoplasm) near the nuclear envelope and several, small patches ir-