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A Quarterly published by

CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

Volume 15

July 1, 1972 to April 1, 1973

vs

ee

Lae enaninia ao te Sas Te

Vol. 15; No. 4

TABLE or CONTENTS

Additional notes on cephalopods from northern California TROT IRL, TALEMUOCH, cocceaseeebsee non uesn eme nme secon ecter 348 Additional notes on some Pacific coast Mollusca geo- graphical, ecological, and chronological IROBER ie RGM lA TE MIAD GE) He eees natin pacacre getter 232 Additions to the opisthobranch mollusk fauna of Marin County, California, with notes on their natural history TERRENCE M. Gosuiner & Gary C. WILLIAMS ....... 352 A device for collecting free-swimming bivalve larvae from laboratory aquaria SEIN CIB TEBARGEReaerenta melee caterer cearencce: 256 Aerial and aquatic respiratory responses to temperature variations in Acmaea digitalis and Acmaea fenestrata SHaron Rose Doran & Donatp S. McKENZIE... 38 A field study on the clustering and movement behavior of the limpet Acmaea digitalis WIVERISWW es WALL OU GHIBY cco aces niacetnantee 223 A new species of Conus from Taiwan Epwarp JAMeEs PeTucH & GrorcE MENDENHALL 96 A new species of monoplacophoran from the abyssal North Pacific RANK MI MIROKOP te cell ici necien eeu Aeunaniede etre 91 A note on the anatomy of the circumesophageal ganglion complex of several doridacean nudibranchs ASD ONAUDREARTRIDGE |) Dalene sche ss. dtats mentisitents 349 Apertural barriers in Pacific island snails of the families Endodontidae and Charopidae PAINTS OTEE IMG) fee tate na ine IN Me ee SU 300 A preliminary list of known opisthobranchs from the Galapagos Islands collected by the Ameripagos Ex- pedition

GaLe G. SpHON & Davip K. MULLINER 2.0.0 147.

A Recent Oreohelix (Gastropoda : Pulmonata) from Baja California Sur, Mexico NWATSEPR VB SIMIIE DER js oe Ba lc 332 A report on cephalopods collected by Stanford Oceano- graphic Expedition 20 to the eastern tropical Pacific Ocean September to November, 1968 W. Gorpvon Fietps & VERONICA A. GAULEY 00.0. is) A systematic note on Ocenebra poulsoni Carpenter, 1865

(Gisopexena) 1B) IVA AONT eect noarteeeceatrecrenoneas eee ceeede 35 Authorship of a taxon RFU GENIE WV ACO AN see tie i ie ee teal Rea de, J abate VW 64 Biology of Okenia ascidicola spec. nov. (Gastropoda : Nudibranchia ) IMIESEATRIGIAWNIOR SE e2c0 cece clea cnce! a meme 2c He Of

Comments on the authorship of some subfamilial names in the Turridae (Mollusca : Gastropoda) WALTER (Os CERNOHORSKYq cuenta cowed hank en 127

THE VELIGER

Page III

Comments on VoKEs’ paper MIGEVAET MD SELUBBARD! seamen tsnccnsincunias 360 Convergence in pulmonate radulae PNTDANT SOLE Mee treet ae cea M ara tin aati mir ath ais 165 Dentaliid taxa referred to the Siphonodentaliidae (Mol- lusca : Scaphopoda) with a description of a new species OEDND Ne pINRARUTER giant ah marin oinn ence uae 21 Description of a new species of the genus Latiaxis Swain- son, 1840 from the South Mozambique Channel, Indian Ocean (Gastropoda : Coralliophilidae) INDASAO) IAZU MAG arum aia cetera neni teutedeantitee Zon Embryonic and larval development in the New Zealand rock oyster, Crassostrea glomerata (Gould, 1850)

IPD UNV ACCA TG es meetnE Ua talaga een mae sede 295

Embryonic development of the camaenid snail, Varohadra yeppoonensis

ALEX S. TOMPA & JAMES N. CATHER ocsssssustseusnen 193

Evidence for a pheromone in the marine periwinkle, Lit- torina littorea Linnaeus INGRID DINTER & PETER J. MANOS urssmecscsnsnninnin 45 External description of a living Aranucus bifidus (Odh- ner, 1936) (Opisthobranchia : Dendronotacea ) Cue CARWSON | &:P JOBE p A enue eC 172 First recorded occurrence of Littorina tessellata Philippi, 1847, from the shores of North America ERASIERV OLA BING HA Man oma etnias a0 rc en nem 158 Food-preference of the nudibranch Aeéolidia papillosa, and the effect of the defenses of the prey on predation VIR GINTAGIEMIVVATERG eee Neto enn a nee ain 174 Further comments on deepwater Volutidae from off south- . east Africa, with descriptions of two new species of Volutocorbis ELAR ATED PAC INEETDER ean area Mien nese 11 Geographic distribution of Pinna rugosa Sowerby, 1835 (Mollusca : Bivalvia), and its occurrence on Clipper- ton Island IBUISAL VAT HSI OAL VAT: aro itanern tn. Ueh keay its Ga cher 43 Mantle changes in the pearl oyster Pinctada maxima in- duced by the pea crab Pinnotheres villosulus

(REVORIGHD ix veer e sie oie se Nats ie ay ba 330 Microarmature and barriers in the aperture of land snails ITAA S SOULE Mga ee ME EEC ae NI ee a ae 81

Morum dennisoni Reeve (Gastropoda : Cassidae) and Strombus costatus Gmelin (Gastropoda : Strombi- dae) collected off the North Carolina coast

EDVARD ROLE DU CE teueeas ni valli cone tone. ie asian sass 51

New name for Pyramidella (Triptychus) olssoni Bartsch,

1926 IPANIES PC CORG ANNI eet eraectenre eter eta 258

Page IV

New records of Pleistocene marine Mollusca from Pacific Beach, San Diego, California MES PBISTHIOPE& 1 So) ey DISIELO Pyetteereee ster tetre meats 6 Nomenclatural notes on West Coast Odostomia (Gastro- poda : Pyramidellacea)

JAMES POU CORGAN Heese titra sie themeertae ane eeeas 357 Note on secondary homonymy Enieysll VOKES) ne rnc se eue ee ae eee eee 102

Notes on abyssal gastropods of the eastern Pacific, with descriptions of three new species BRANKGRROKOP fico erate one ete niatie tr tote ate 15 Notes on Cypraea cinerea Gmelin and Cyphoma gibbos- um (Linnaeus) from the Caribbean Sea, and descrip- tion of their spawn

TKGEAUSHMBAND EIA pote ween re ate nain Miser Reape ne ie 335 Notes on two endemic South African Cypraea 1, INL, IXanepyoieny Ge ID, WWE ANIESDINT ccomereettcommecrenomonmnenooadite 125

Observation of the glochidium, metamorphosis, and juve--

nile of Anodonta californiensis Lea, 1857 BETERYNMD EIS CU ee oe DER Oe lc Noreeetare 57 Observations of the feeding habits of Tochuina tetraquetra (Pallas) (Gastropoda : Tritoniidae) Mary K. WicksTEN & JoHN D. De Martin 1....... 195 Observations on growth, feeding, reproduction, and devel- opment in the opisthobranch, Fiona pinnata (Esch- scholtz) JOHNGIE THORERIAN (ie eal ei te Naru aten eters 142 Observations on Hexabranchus from the Australian Great Barrier Reef (Gastropoda : Opisthobranchia) Aaa MIO MPS ONG ace aii nee i tea rete eee 1 Observations on removal of spines by muricid gastropods during shell growth IMIBISR OW RIN EM Res C/ARRTKSER eee tenet 69 Odostomia minutissima Dall & Bartsch, 1909, a synonym of Odostomia raymondi Dall & Bartsch, 1909 PAMESPXUCORG AN ee See ete au tie oat ane rm amen 359 Opisthobranch mollusks dredged in San Francisco Bay during the period 1966 to 1971 JorNny HOLEEMAN neo ee eee 59 Oxidation of C'‘-glucose by the aestivating snail Pila glob- osa (Swainson ) S. RacHupatui Rami Reppy « R. RAMAMURTHI... 355 Polydora and related genera as borers in mollusk shells and other calcareous substrates (Polychaeta : Spion- idac ) JaMEs A. BLAKE & JOHN W. EVANS cocci 235 Preliminary note on hermaphroditism and embryonic stages in Diplodon variabilis

CEMA GuuziwANEDE, LASCARY ) eee nena 213 Preserving terrestrial slugs by freeze-drying PLB CROWET Mie wren. alee ent ah geen ama Ee 254

THE VELIGER

Vol. 15; No. 4

Range extensions of Conualevia alba Collier & Farmer, 1964 ANTONIO. J.. FERREIRA joss dis.cte nines one eee 53 Range extensions of several littorinids (Mollusca : Gastro- poda) in Florida FRASIER’ ©) BINGHAM) (20) cecsiecncnncuiticn eee 250 Review of the bathyal gastropod genus Phanerolepida (Homalopomatinae) and description of a new spe- cies from the Oregon Oligocene Carone’S) Hick MAN fin. ceeecde ee 107 Scanning electron micrographic study of the dorsal integ- ument of the land slug Lehmannia poirieri (Mabille, 1883 ) Joun A. Arcani & NORMAN HODGKIN ...secsnnee 338 Seasonal migration and population regulation in the lim- pet Acmaea (Collisella) digitalis PAUL A} BREEN 2). Gacotteee eee 133 Selective predation and prey location in the sea slug Navanax inermis Grec M. Bian & ROGER R. SEAPY ciccsssssanennnen 119 Settlement, growth rates and depth preference of the ship- worm Bankia setacea (Tryon) in Monterey Bay

BG) HApERME & J.C; MELIOR =. nena 265 Some notes on the genus Teramachia (Volutidae : Callio- tectinae ) HarRALp: Az) REHDER: (iteci ch ccncacie cee ee 7 Some opisthobranchs (Mollusca : Gastropoda) from Oregon GALE G.) SPHON (1.0.00, Soe ae 153

Some records on West American Cenozoic gastropods of the genus A foria

MADOKHT JAVIDPOUR) een 196 Soviet contributions to malacology in 1971 KENNETH J. Boss & Morris K. JACOBSON owen 362

Spawning and development of the eggs, in the laboratory, of Illex coindetu (Mollusca : Cephalopoda)

S. v. BoLetzKy, L. Rowe & L. AROLES ween 257 Status of Obeliscus clavulus A. Adams, 1854 James UX; \CORGANS. ee eee ee 359 Stenoplax circumsenta Berry, 1956, in the Gulf of Cali- fornia ANTONIO YU BERREIRAG HI Gee a DS

Systematics and distribution of western Atlantic Ervilia (Pelecypoda : Mesodesmatidae) with notes on the living Evvilia subcancellata

JOHN Ds DAVIS) 2.2 use ee 307

Thecacera picta spec. nov. from Suruga Bay, Japan (Nudi-

branchia : Doridoidea : Polyceridae)

KiKUTARO! BABA ina. cee eee eee ne 88 The ecology and behavior of Nautilus pompilius in the Philippines

Norint Haven

Vol. 15; No. 4

The genera Chromodoris and Felimida (Nudibranchia : Chromodorididae) in Tropical West America: dis- tributional data, description of a new species, and scanning electron microscopic studies of radulae

Hans Bertscu, ANToNIo J. FERREIRA, WESLEY M. FARMER & THOMAS L. HAYES cites 287

The genus Chelidonura from the Marianas Islands (O-

pisthobranchia : Cephalaspidea )

CHER CARUSON & P. Je HOFF! 22.20 cca eetnntennee 20

The intertidal behavior of the bean clam, Donax gouldii Dall, 1921

PI TONOAS WETS ER WAN se sect sececcsae sentra sedettach tacsiebacienistine 206

The mucus holdfast of Littorina irrorata and its relation- ship to relative humidity and salinity

ERSTE RG @ se BUN GWA eee eee eee 48 The northwest American Semelidae AU CINE RAN 5 CONN ee eres NUE eae te ae lle 314

The occurrence of Polycera zosterae O'Donoghue, 1924 in the Bodega Bay region, California, with notes on its natural history (Gastropoda : Nudibranchia)

TERRENCE M. GosLINeR & Gary C. WILLIAMS ..... 252

Thermal and salinity effects on ciliary activity of excised

gill tissue from bivalves of North and South Carolina PNIGAN AEs SIELO EWA KOERO oe nee meteors, 215

The role of wave impact and desiccation on the distribu-

tion of Littorina sitkana Philippi, 1845 RSYPTLIYUN, LBV BTS DeQISIN IS Gaia stl ene 129

The use of the foot and the captacula in the feeding of Dentalium (Mollusca : Scaphopoda )

MO WISB ER GATINIE AM [ieee ent tem tenant 29

Two additions to the opisthobranch fauna of the southern Gulf of California

IRAN SBE ERTES Cgc ie rene ee a est eee erste a 103 Unusual egg-deposit by a cuttlefish So Wo THONISIA SSC: 6 atten crcaneeemene sense rere ete ean 61 Zoogeography and ecology of seven species of Panamic- Pacific Scaphopoda VIE S Ee CAD IEING Sect teeter creme rae nme uO eet 340

AUTHOR INDEX

AIKEN, D. W.see KiLpurn, R. N.

AppicoTt, WARREN O. ....... (GO), (CIO), (OR), (A4e3) ARCADI, JOHN A. & NORMAN HODGKIN cece 338 ARoLes, L. see Boterzxy, S. v., L. Rowe & Azuma, Masao Basa, KiKuUTARO BANDEL, KLAus REED RIES et OVA VITAN oe test sat Mee ee teat ecole et lenge ae 129 IB DRALS CHPRILAIN| Stee am Mersie) SPU seek Ore sean e el ease tec patet ieee 103

THE VELIGER

Page V

Bertscu, Hans, ANTONIO J. FERREIRA, WESLEY M. Far-

MER NSM FEONAS Maa ELAVE SI muenne seen nnrne 287 IBINGEVAMEMERASTE RU ©) sient eet tated cee 48, 158, 250 BISHOP AVI meg Sane DISH OD sures ntin css cmsneitnrecienrctcis Briar, Greco M. & ROGER R. SEAPY usec BLAKE, JAMES A. & JoHN W. Evans BOGE DZKVAN Sony lo eer te cr pena nner tee Asai i ate ete 61 BoLeTzky, S. v., L. ROWE & L. AROLES .......00 Boss, KENNETH J. & Morris K. JAcoBSON Books, periodicals and pamphlets ........ 66, 161, 262, 370 IBREE N/R EAU Geert een recat eer Oe pen se hee 133 ChrrsonpiG He Pls HOnE eet neces 20, 172 CAR RTE ROS VUBIEB OURINIE ge eee en een nnn nee 69 CaTHER, JAMES N. see ALEZ S. TomPaA & CERINOHORSKYA WN ATETERM © ieee teen ee ae ene 127 (COANAMENU GENIE WV eee ee 64, 314 CORGANANJANMES XG a: enue nie tet DX, By BOY) GROWER TET EM My ae Ao oie ec eer une ener nessa neues 254 DAVIS in] OUND ern eee 307 DAE TIScUAPEEDERUN Geers sree ee 57 De MartIN1, JoHN D. see WicksTEN, Mary K. &

TTB) uN ANN OAS gee ec ee ee ie sen San oeeere re eee ta 295 Dinter, INGRIp & PETER J. MANOS fooler 45 VTP oR NRE VOR Gree ei mete teal conti peels sue on ttc: 330 Doran, SHARON Rose & Donatp S. McKENZIE ........ 38 BCKEL BARGER WIGEVINGA Mire tenis een santa tattin satiate 256 Evans, JoHN W.see BLAKE, JAMES A. &

Farmer, WesLey M. see BertscH, Hans, and others BERREIR AW AINILONIOM Janes tet renee crt tintacnis BS, DD

see also BertscH, Hans, —, and others

FieLps, W. Gorpon & VERONICA A. GAULEY uisiccccon 113 GISUNE Veg OUIS UL ta) cee ener ene Len tinrtstoteere ces 29

GauLey, Veronica A. see Fietps, W. GORDON & GosLINER, TERRENCE M. & Gary C. WiLuiAMsS 252, 352

VADER UTE ies Cre ups CoulVERBDOR MPa tents cense 265 TES VASA SIN INU ODSDENIT cay eect ee ee at eerec pointer ocr i) Hayes, Tuomas L. see BertscH, Hans and others

FAUT @ MAIN GA RO TEE DSi cece eterna eran nats raare caters 107

Hopcxin, Norman see Arcap!, JOHN A. & Horr, P J. see Carson, C. H. « TLOMMEN CANE OEUNGM sree eases eesti recerectrmcer 59, 142

HUBBARD, MICHAEL D. ...ccccceee

TRAIN Sg INET OMOAS WITS fete Metter ree tte stcecat ot tear sceatoen eats 206 Jacopson, Morris K. see Boss, KENNETH J. & ANADEGURMIMIADOKGE iy een nnmncitcm rename sirarnteer atte: 196 IGE EIN GW ASS Vi RAG deere mnnun tei ects (163) Kitpurn, R. N. & D. W. AIKEN 125 TSGEYATEN OF EON Sat KODE OTs ING Hes eraser terete ere eer 21 Manos, Peter J. see DiInTER, INGRID &

INGOIPAD TIE Nia VEN S te ee cect ie kat nar etter e ane nantes seta 340

McKenzig, Donatp S. see Doran, SHARON ROSE &

Page VI

ME Ltor, J. C. see HApERLIE, E. C. & MENDENHALL, GEorGE see PeTUucH, Epwarp J. &

INITEDERE PO WiATSTE REIS se tee ete on cee needs mecca ere caea 332 INTORSEN Vey Ac CHA es eee erent natn e entesele re nena 97 MuLuiner, Davin K. see SpHON, GALE G. & INGtesmandenewsme tence te Oil, WEY, Ba, Si) PARTRIDGE spe) O NATED al] geese eee nee eee are 349 PASGCAR, CELIA GLUZMAN DE oecsssssssssssnscsnsssssenssusetisssssnssiteens 213 PETUGH, EDWARD) iter eet ee Cait tee menace tae rsat ane oil Petucu, Epwarp J. & GeEorGE MENDENHALL ............ 96 IRVAD WWII GEORGE Eyes tie ea oc sores online eer ae ae DRL 35 RAMAMURTHI, R. see Reppy, S. R. R. &

Reppy, S. RAGHUPATHI Rami & R. RAMAMURTH ....... 355 IREHDER: IARAT DAK 28 isa yor ieee ehoM mnt east Ul ROKOP) PRANK iif. svc nd eeeen ee ree ear tae 5,

Rowe, L. see BoLetzxy, S. v., & L. AROLES

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Vol. 15; No. 4

SATEVAT BS 6c ES SATE VA Taner aiar ret rene 43 Seapy, Rocer R. see Grec M. Biar & SHOEMAKER, ALAN Hi iio) dea nusadacintin eee 215 SOE Ms GAGA Niji teers leer na 81, 165, 300 SPHON, Gate, G., decoction nena 153 SPHON, Gate G. & Davin K. MULLINER occ 147 STOHLER, R. (66), (67), (68), (162), (163), (164), BME tak (263), (264), (370) Ean ow) SWAT ADC ER BROBERT | Rem enrer te aerate ean 232, 348 IGHOMPSON IE, Bis 2 lea ae ee 1 Tompa, ALEX S. & JAMES N. CATHER ouesccssneninssneien 193 VOKES, EMItey: He. o.scccncsccicsisndsenccient ee ee 102 WATERS, VIRGINIA, L.. .c:c.c:ccrecngoscsanceicnae eee ee 174 WicksTEN, Mary K. & Joun D. De Martin 1.......... 195 WILLIAMS, Gary C. see GOSLINER, TERRENCE M & ........

WILLOUGHBY; JAMES) Wi ciiceccecencnsneca cee eee 223

SECTIONAI DIVISION OF

A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME 15 July 1, 1972 “NuMBER 1

CONTENTS

Observations on Hexabranchus from the Australian Great Barrier Reef (Gastropoda : Opisthobranchia) . (2 Text figures) T. E. THomMPson 66 oat

New Records of Pleistocene Manag Melee hoa Pacific Resch: San Diego, Cali-

fornia. M. J. Bishop « S.J. BisHop . . 6) On) GL RE ROM ON ROUMSSe PO NORE SMD trOeieL Some Notes on the Genus Teramachia ete : a (1 Plate) Haracp A. REHDER . 3 ASie Oat cen wey / Further Comments on Dees oleae Genk off Sent Aftica, with Descriptions of Two New ae of Volutocorbis : es Haratp A. REHDER . . fs : ei eel Notes on Abyssal Gastropods Fe the ee Pacific, mth De npcons # Three Ne Species (2 Plates)

Frank J. Roxop A ; . 15 The Genus Chelidonura Boe ae eae tases (Gemichanc: Gon:

aspidea) . C.H.Cartsone PJ.Horr. . . . : - 20 Dentaliid Taxa Referred to the Sr sionaden aitdse etnies See AG a Description of a New ee (3 Text aay Joun N. KraEuTER sp tatNe . QI The Use of the Foot and the Capenune in ie Featns of Deetaleaa (uote. Scaphopoda). : 5 Text ae Louis F Gainey, Jr. . . : era Se Me eeler sa Mer icetineiiier Pe enre 2O A Systematic Note on Ocenebra ates Gos Ais (6 Text figures) GrorcE E.RapwIN . ... . MM Me Meee 1 ol fos AS yeh Wop eg) Aah) tom, ei ee OS

[Continued on Inside Front Cover]

Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 15: $18.- Domestic; $19.50 in the Americas; $20.- in all other Foreign Countries Single copies this issue $10.-. Postage extra.

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WILLIAM H. DALE!

LIBRARY MOLLUSKS

Contents Continued

Aerial and Aquatic Respiratory Responses to Temperature Variations in Acmaea digitalis and Acmaea fenestrata. (2 Text ae SHaron Rose Doran & Donato S. McKenzr.. . Se Bene Geographic Distribution of Pinna rugosa Sowerby, 1835 (Mollusca : Bivalvia), and its Occurrence on Clipperton Island. B. SALVAT & Fo SAL VAT 2 (opie eri Bro on Soetoro ile vat aN ool cata atn Vann Evidence for a Pheromone in the Marine Periwinkle, Littorina littorea Linnaeus. (1 Text figure)

Incrip DinTER & PETER J. Manos Sic : elias iesiUle

The Mucus Holdfast of Littorina irrorata and its Resco to nee Humidity and Salinity. (7 Text figures)

Frasier ©: BINGHAMG 9 (Eo 3. Appa ura ne eat eit hoor Darter te

Morum dennisoni Reeve (Gastropoda : Cassidae) and Strombus costatus Gmelin (Gastropoda : Strombidae) Collected Off the North Carolina Coast. (1 Plate) EDWARD J. /PETUGHD 1551/5) 62) 2) 7) er ia er tot notin OMe otaet Ina nt ae eRe

Range Extensions of Conualevia alba Collier « Farmer, eee ANTONIO J. FERREIRA .

Stenoplax circumsenta Berry, 058 in ee Gulf a California. (1 Plate) ANTONIO J. FERREIRA . . . ail och oui Uauehl ee ey anoal ach Salts tie aN Cog Observation of the Glochidium, hceseaene and Juvenile of Anodonta califor- niensis Lea, 1857. e Plate; 1 Text ae Peter N. D’Euiscu. . . Seren im ners Opisthobranch Mollusks mee in ers PANES Bay Duane ae ‘Period 1966 to 1971. (1 Text figure) orn) }-ELOLeE MAN) ac) meneame sit ial oe nei. Sarai atu eehestatth neh quelle geval tr olalKe NOTES & NEWS. . . og Selgin by Seri ea eagle [eo ean a ane Unusual egg-deposit Ge a cuttlefish. (1 Text figure) S. v. BoLeTzky INFORMATION DESK .. . . at. (ek | Sel eeu Ra

What’s the Difference? —- Mutiontne of a Tagen’ EucEene V. Coan BOOKS, PERIODICALS & PAMPEMEEAS ioe cline. toy alin on) say me re arenmeniey ne

. 38

° 43

» 45

. 48

- 51

- 53

- 59

- 57

» 59 . 61

. 64

. 66

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

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Observations on Hexabranchus

from the Australian Great Barrier Reef

(Gastropoda : Opisthobranchia)

T. E. THOMPSON

Zoology Department, University of Bristol, United Kingdom

(2 Text figures)

INTRODUCTION

THE SWIMMING DISPLAY of Hexabranchus is one of the most splendid sights a malacologist may hope for. Despite the interest this has aroused there are great gaps in our knowledge of the genus. The normal diet is unknown, for instance, and the taxonomic situation is very confused, with dozens of proposed species. In order to help towards the solution of these problems, the present paper contains a description of the external features and anatomy of Australian specimens, together with notes on the gut contents and the swimming behaviour. These have been compared with observations on Hexabranchus by Morton (1964: Australia), GoHar & Sotiman (1963: Red Sea), ViceNTE (1963: Madagascar) and Epmunps (1968: Tanzania ). A suggested synonymy concludes the paper.

Miss J. A. Langhorne prepared the drawings for pub- lication.

SOURCE or MATERIAL

Alive: One specimen, length 21 cm, creeping over coral sand near the reef crest at Heron Island, Capricorn Group, Australian Great Barrier Reef, June 1968. The specimen weighed 334 gr, and the pH of the skin was neutral.

Preserved: Several specimens in the collections of the Queensland Museum and of Miss Isobel Bennett of the University of Sydney.

Other Material: Kodachromes from life of Great Barrier

Reef individuals, kindly given by Mr. D. Henderson and by Dr. L. Harris.

DESCRIPTION

A. External Features (Figures 1 and 2)

The basic colour of the body was rich dark red, with

irregular pale mottling. The rhinophoral tentacles were red-brown; the rims of the rudimentary pallial sheaths were bright red, with pale radiating markings. The pedal sole was paler in colour than the rest of the body. The oral tentacles took the form of a pair of flattened lobes (Figure 2C), with frilled edges. The gills, 7 in number, were ample and tripinnate, each being retractile into a separate pallial pit (7.¢e. phanerobranchiate). The gills twitched continuously. Within the ring of gills lay the conspicuous white anal papilla and the nephroproct (Fig- ure 2B).

The colour pattern described above is that which can be seen in the creeping animal. If alarmed, however, the mantle edges are spread out and the animal takes on a vivid aspect, in marked contrast to the well camouflaged resting stage. The unrolling of the mantle margins exhib- its the white edges and brilliant purple sub-marginal zones, observable in both dorsal and ventral views. In the dorsal view can be seen an additional feature, where bril- liant red radial streaks penetrate the purple zones. There are also approximately bilaterally symmetrical slightly embossed patches of pink surrounding bright red spots; these are partially concealed when the animal is creeping (Figure 2A).

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B. Swimming Behaviour

In swimming, the enrolled mantle skirt is spread out, dilated, and strong locomotor waves are propagated rear- wards from the anterior pallial margin. The waves are synchronous on the two sides of the body, and each wave takes about 4 seconds to travel its course. ‘Two waves are usually in progress along the sides of the body at any moment. At the same time, the body undergoes great dorso-ventral flexions, a full cycle of activity occupying about 4 seconds, Swimming may continue for many min- utes, and it is probable that this is more than a simple escape reaction. Certainly, a brilliant display of colour accompanies the unrolling of the mantle edge, and it may be that swimming here is the behavioural component of an aposematic or warning display. While swimming, the lateral edges of the foot are brought together in order to conceal the sole and the rhinophores are held back against the dorsal mantle.

C. Anatomy

The central nervous system was orange-pink in colour and pustulose like that of a pleurobranchomorph. The blood-gland (of uncertain function) was large and dark red, overlying the pallial nerves behind the nerve ring. A conspicuous pear-shaped gastric caecum was present. The penis was extremely large, 3 cm or more in preservative. The penial sheath was helically coiled around the narrow vas deferens. The penis was unarmed.

The jaws were dark brown, lying one on each side of the oral canal. They exhibited antero-posterior wrinkles and were composed of minute rods of chitin. The radular for- mula was 48 X 91 -0-91. This is somewhat different from Red Sea specimens (40 X 60:0-60 to 60 X 78:0-78, according to GoHar « Soimman, 1963). The hooked simple teeth lacked subsidiary denticulations of any kind.

D. Diet

The hindgut was filled with fragments of greenish material which contained minute calcareous spicules. Dr. P. Kott kindly identified these remains as the ascidian Didemnum moseleyi Herdman. This is the first record of

(< on facing page)

Figure 1

Hexabranchus sanguineus

A Swimming vigorously B_ The same, creeping over coral debris, Heron Island, Great Barrier Reef

such a diet for Hexabranchus. Other authors have identi- fied the gut contents as sponges (Younc, 1969), or as foraminiferans, worm-tubes, and gastropod and echino- derm shells (EaLEs, 1938).

DISCUSSION

Many species of Hexabranchus have been proposed, but as long ago as 1909, Exior stated that “It may be doubted whether the species of Hexabranchus are for the most part more than colour varieties”. In the following year, this distinguished authority again made it clear that he doubted the validity of nearly all the claimed specific divisions (Euiot, 1910). Since that time, OsTERGAARD (1955) and Marcus & Marcus (1962) have added new proposed species of Hexabranchus but without seriously attempting to distinguish their specimens from previous described species. Indeed, apart from a minor difference in radular morphology, the only important feature distin- guishing H. morsomus Marcus & Marcus, 1962, from older described forms is the fact that it came from the Virgin Islands in the Atlantic Ocean, whereas other rec- ords had been exclusively from the Indo-Pacific. Only in GoHar & SOLIMAN’s (1963) paper is there an attempt to describe and illustrate the variations which may occur in the colour patterns within a particular sea-area. Their results support Eliot’s view that there may be only one valid species of Hexabranchus. Eymunps (1968) claimed to have detected differences in the swimming behaviour of Tanzanian specimens compared with published accounts of Australian material (Morton, 1964), but his illus- trations do not support this contention. His plate 1 (1) shows two waves on the mantle margin on each side, despite his assertion in the text of his paper that only one is present in Tanzanian material. There are thus no sub- stantial differences either in behaviour or colouration in Hexabranchus recorded from localities all over the Indo- Pacific Basin. The published records probably all refer to a single species. The name Doris sanguinea Riippell & Leuckart, 1828, appears to have priority.

CONCLUSION

List of Principal Synonyms of Hexabranchus sanguineus (Riuppell & Leuckart) Doris sanguinea Ruppell « Leuckart, 1828 (Red Sea)

Hexabranchus proetextus Ehrenberg, 1831 (Red Sea) Doris flammulatus Quoy « Gaimard, 1832 (“fle des Amis”)

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Doris marginatus Quoy & Gaimard, 1832 (China, Madagascar, Red Sea, Ryukyu Islands, Japan, Caroline Islands, Mar- shall Islands, Australia [Great Barrier Reef to Abrolhos Islands], New Caledonia, Hawaii, Tonga, British Solomon Islands, Fiji, Ceylon)

Doris cardinalis Gould, 1852 (Hawaii)

Doris sandwichensis Eydoux « Souleyet, 1852 (Hawaii)

Hexabranchus pulchellus Pease, 1860 (Hawaii)

Hexabranchus pellucidulus Abraham, 1876 (unknown locality)

Hexabranchus suezensis Abraham, 1876 (Red Sea)

Hexabranchus aneiteumensis Abraham, 1877 (New Hebrides)

Hexabranchus mauritianus Abraham, 1877 (Mauritius)

Hexabranchus orbicularis Abraham, 1877 (Mauritius)

Hexabranchus aneitus Bergh, 1878 (Philippines)

Albania formosa Collingwood, 1881 (Formosa)

Hexabranchus imperialis Kent, 1897 (Abrolhos Islands)

Hexabranchus lacer Bergh, 1900 (not Cuvier, 1804) (unknown locality)

Hexabranchus plicatus Hagg, 1901 (unknown locality)

Hexabranchus digitatus Eliot, 1906 (Maldive Islands)

Hexabranchus tinkeri Ostergaard, 1955 (Hawaii)

Hexabranchus aureomarginatus Ostergaard, 1955 (Hawaii)

(<on facing page)

Figure 2

Hexabranchus sanguineus

A Creeping posture B_ Spread out on a table, dorsal view C. The same, ventral view D-G_ Various views of the vigorous swimming movements

Literature Cited

Eates, NEvue B. 1938. A systematic and anatomic account of the Opisthobranchia. Sci. Rep. John Murray Exped. 5: 77 - 122 Epmunps, MALcoLm 1968. On the swimming and defensive response of Hexabranchus mar- ginatus (Mollusca, Nudibranchia). Journ. Linn. Soc. (Zool.) 47: 425 - 429 Exviot, Cuarzes N. E. 1909. Notes on a collection of nudibranchs from Ceylon. Spolia Zeylanica 6: 78-95 1910. Nudibranchs collected by Mr. Stanley Gardiner from the Indian Ocean in H. M.S. Sealark. Trans. Linn. Soc. London (Zool.) 13: 411 - 438 Gonar, H. A. EF «& G. N. Sotiman 1963. The biology and development of Hexabranchus sanguineus (Rupp. and Leuck.) (Gastropoda, Nudibranchiata). Publ. Mar. Biol. Sta. Al-Ghardaqa 12: 219 - 247 Marcus, Ernst & EvELINE DU Bois REYMOND Marcus 1962. Opisthobranchs from Florida and the Virgin Islands. Bull. Mar. Sci. Gulf & Carib. 12 (3): 450 - 488; 28 text figs. Morton, JoHN EDwarD

1964. Locomotion. In Physiol. Mollusca 1: 383 - 423 OsTERGAARD, JENS MATHIAS 1955. Some opisthobranchiate Mollusca from Hawaii.