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POeLIGER

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

Volume II

July 1, 1968 to April 1, 1969

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Vol. 11; No. 4

THE VELIGER

Page III

TABLE OF CONTENTS

A color variation of Aldisa sanguinea.

RICHARD AY ROEBER Ml Vers sissicdeo eee ee aie cosne eens 280
A cowrie mutant from the Gulf of Thailand.
FRANZ ALFRED SCHILDER sw... ee ee eee eee eee 34

A list of types of the family Volutidae held by the National
Museum of Victoria.

BREAN BP pS MIUBEWM ly pcre left alnrcteie cralsustes ote te ale sale sve 39

A method of color preservation in opisthobranch mollusks.

(CORDONT AC ROBILLIARD! a). ciescts sate cle ieie oi citlets 288
A method of tagging mollusks underwater.
IRIGHARD J ROSENTHAL alee ccs eee ee 0 289

An additional record for Cypraea teres in the Galapagos
Islands.
WituiaM K. Emerson & WituiaM E. OL, Jr. 98
An annotated list of opisthobranchs from San Luis Obispo
County, California.

Ricuarp A. ROLLER & STEVEN J. LONG... 424
An emendation.
RAW EORDINS GATED) fice ieiald dole ental eletelete 80
An immunological study of pelecypod taxonomy.
BMBIARKEVINGFISHER) «ni suc. ceca desis saves’ 434

A new genus and two new species of Typhinae from the
Panamic Province (Gastropoda: Muricidae).
_ Inline, IDS) Ge ocodanosuasoorGooedS 343
An investigation of the commensals of Cryptochiton stel-
ler: (MipEenporrF, 1847) in the Monterey Peninsula
area, California.

SIEEVELINM IME SIRE Run Mnrtenareiiereiariersiebecicnciae 121
A new Neptunea from the Pacific Northwest.
PANTOTE VANE Gr IMELTRET Cu asl yeiatopousileiicers. oiete: sisi olore laude 117

A new record of Corambella steinbergae LANCE, 1962.
PRERRENGEMGOSEINER (0) y.'efs it srctvieie vie.’ vin levee sis 147
A new species of enopleuthid squid, Abraliopsis (Wata-
senia) felis, from the California Current.
Joun A. McGowan & TakasHi OKUTANI aor! Le
A new species of Murexsul (Gastropoda: Muricidae) from
the Galapagos Islands.
WiturAM K. EMerson & ANTHONY D’ATTILIO 324
A new species of Puncturella (Cranopsis) from the north-
eastern Pacific.
I. McT. Cowan & JAMES H. McLean... 105
A new species of Strombina from the Galapagos Islands.
Wiiu1AM K. Emerson & ANTHONY D’ATTILIO 195
A note on feeding and excretion in bivalves.
Em TINA NUAINT igi salsa far aiiltas also eTavar a ua railele aya 198
A note on the range of Gastropteron pacificum (Opistho-
branchia : Cephalaspidea).
Hans BertscH
An overlooked subgenus and species from Panama.
A. Myra KEEN

A preliminary survey of mollusks for Consag Rock and
adjacent areas, Gulf of California, Mexico.
HELEN DUSHANE & ELLEN BRENNAN... 351
Archidoris odhneri (MacFartanp, 1966) comb. nov.,
with some comments on the species of the genus on
the Pacific coast of North America.
ROBERTHBURNs) Hic a cassie: ava veieiee cea NEO 90
A record of the Indo-Pacific cone, Conus ebraeus, in Gua-
temala.
WIE LTANAKGrE:MERSON). sie ay stitiemciereete 33
A review of the living leptonacean bivalves of the genus
Aligena.
FVAROLDNWAHEVARRY ys. -hojerereiets > cre oidintes Shel ieee 164
A survey of the littoral marine mollusks of the Caribbean
coast of Costa Rica.
JosepH RicHarp Housrick
Banding patterns in Haliotis - II. Some behavioral con-
siderations and the effect of diet on shell coloration
for Haliotis rufescens, Haliotis corrugata, Haliotis so-
rensent, and Haliotis assimilis.
DAVIDyS AOUSENT 9 i ahi) recs creeeioeiets te mheretees 135
Distribution of organic bromine compounds in Aplysia
californica CoopEr, 1863.
IEINDSAV@ReAVVINIKEERY® are ia-rdeicisisii reine erie 268
Effects of turbidity-producing substances in sea water on
eggs and larvae of three genera of bivalve mollusks.
Harry C. Davis & HERBERT Hipui............... 316
Familial placement of Hanetia JoussEAUME, 1880 (Mu-
ricidae) and Solenosteira Dati, 1890 (Buccinidae).

WIEETA MG KS EMERSON) Ue: ojorercie) shores srecaiejeieteiee 1
Feeding behavior of Corambella steinbergae.
WANES WMICBEDH® 6 loeesroiccc'siapo)ata sie suckers 145

Five new species of Mitridae from the Indian and Pacific
oceans.
AAI GMI CATE 50 Beracavsc ots eicle crstwetel 0 ¢s-ceysterctonone te 85
Further observations on the West American Marginellidae
with the description of two new species.

Barry RotH & EUGENE COAN ...............- 62
Gross anatomy and classification of the commensal gastro-
pod, Caledoniella montrouzieri SOUVERBIE, 1869.

JOSEPEPROSEWATERWM gyoiiaerya crt aacioirclelere ire 345
Growth characteristics of Acmaea persona ESCHSCHOLTZ.
RONBISE NINYip Mateo cise ctotnsiieloictens: ceemekepel termed 336
Growth study in Olivella biplicata (SowErRBy, 1825).
RupoLF STOHLER
Haliotis pourtalesii Dati, 1881 from Florida waters.
@HARTESH) AGUICE Ws fcetreis.n = (ey. deed 140
Invertebrates taken in six year trawl study in Santa Moni-
ca Bay.

JOHN G CARTISTES| fa |. scr <a aan 237
Laternula living on the Pacific Coast?
ACEMIYRAGIGEEN®, \'steiScoresossnejeostielevoroheetee serene 439

Page IV

List of type specimens of Terebridae in the British Muse-
um (Natural History).

WALTER OLIVER CERNOHORSKY _........... «ss 210

Littorina littorea in California (San Francisco and Trin-
idad Bays).

J/NMGSS (CARTONS =p bo oad vbo pono anO CODE DOOOS 283

Maturation of gonads of oysters, Crassostrea virginica, of
different geographical areas subjected to relatively
low temperatures.

WHGHOR IL, ILOOYNNOR s000000d000000000000 153
Mediargo, a new Tertiary genus in the family Cymatiidae.
UDITED SSMEERRY Sie selon oagued Aarne 42
Molluscan faunas of Pacific Coast salt marshes and tidal
creeks.
[Xaaeist 15}, IMUANGDONYNED) = = can00accccc000000000 399
New northern limit for the limpet, Acmaea digitalis.
WitrianivAN [ESSER © .pciii loins ere rnis oeneree 144
New records of nudibranchs from New Jersey.
Rosert E. LovELAND, GorDON HENDLER & GARY
INE WiKIRKe 9 0s dassteh itis aircon ava ores: 418

Nomenclatural changes for the new species assigned to
Cratena by MacFartanp, 1966.
RIGHARDPAG ROLUER 2 Litt on irene rea 421
Notes on Cryptochiton stelleri (MippENpDorFF, 1846).

G. E. Mac Ginitie & Nettie MacGInIrTle heh 59
Notes on the food of Conus dalli Strarns, 1873.
AMES WRINVYBAKICEN) ~ Ste} sietoie eortettctiemnaetate 50

Notes on the habitat and anatomy of Jouannetia quillingi
from North Carolina coastal waters.
IDOUGTASUAUAWOLEES! 9 ¥.ic0 1. <br srr eert ete 126
Observations on aquarium specimens of Oliva sayana
RAVENEL.

AxeL A. Otsson & Mrs. L. E.Crovo ........ 31
On an octopod from Placentia Bay, Newfoundland.
Freperick A. ALtpricHe C.C.Lu .......... 70

Pelecypod-sediment association in Tomales Bay, California.
DON MAURERS otic Se shinee eee 243
Predators on Olwella biplicata, including a species-specific
predator avoidance response.
DHERAIGSEDWARDS) 4. otieinotaemmns Heme 326
“Ptychodon” misoolensis ADAM & VAN BENTHEM JUTTING,
1939, a new Guinea strobilopsid land snail and review
of the genus Enteroplax. ;
INTEAN) SOLEM S220, cbc shee nape tateis sau aevenere ec 24
Quantitative relationships between gill number, respira-
tory surface, and cavity shape in chitons.
Keay. MisJOEUNSON:. <j Shiagalil tetas Seaeraniteeenene ate 272
Quantitative studies on cowries (Cypraeidae) of the
Allan Hancock Foundation collections.
GERALD J. Bakus

THE VELIGER

Vol. 11; No. 4

Recognition of an Eastern Pacific Macoma in the Coral-
line Crag of England and its biogeographic signifi-

cance.
EUGENE. V. COAN)) ota ce eee 277
Rediscovery of Terebra trochlea DESHAYES.
Twita BRATCHER® | .oalace cee ene eee eee 334

Removal of pea crabs from live oysters by using Sevin.
Jay D. ANDREws, Donna TurGEON & MariAN HREHA

aids sia, aueltiea 6 baie Ore Od oe eee 141

Revision of “Introduced mollusks of western North Amer-
1Gaeae

JAMES CARLTON © i3..0.00)) Jee eee 284

Seasonal gonadal changes in two bivalve mollusks in To-
males Bay, California.
VERNON KENNETH LEONARD, Jr. ......--.00- 382
Seasonal observations on diet, and stored glycogen and
lipids in the horse clam, Tresus capax (Gouxp, 1850).

Rosert'G. BEREID | onc ok eee 378
Sexual dimorphism in Tegula funebralis.
PETER W. FRANK oo aecene coc un ce 440

Some observations on the ecology and behavior of Luca-
pinella callomarginata.
RicHARD L, MILLER 5.220. 50 eee 130
Spawning and development in the trochid gastropod Eu-
chelus gemmatus (Goutp, 1841) in the Hawaiian

Islands.

Tuomas M. Duch 2320505: ieee 415
Spawning notes, II. Mitra dolorosa.

Fay H. WOLFSON oie che slo's eclorlel eee 281
Spawning notes, III. Strombina maculosa.

Fay H. WOLFSON "ais. coho ieice el eee 440
Spawning notes, IV. Cerithium stercusmuscarum.

Fay H."WOLFSON? oitissrq sia cna eee 44]

Spawning of the American oyster Crassostrea virginica,
at extreme pH levels.
ANTHONY CALABRESE & Harry C. Davis Bip one)
Studies on the Mytilus edulis community in Alamitos Bay,
California. IV. Seasonal variation in gametes from
different regions of the bay.

Donatp R. Moore & DonaAtp J. REISH_........... 250
Studies on populations of the cowrie Erronea errones
(LINNAEUS).
FRANZ ALFRED SCHILDER & MARIA SCHILDER 4 os)
Tellina ulloana, a new species from Magdalena Bay, Mexi-
co.
Lro.G. HERTLEIN’ © (FO: ero oa 80

The ecology of Macoma inconspicua (BRODERIP & SOWER-
By, 1829) in Central San Francisco Bay. Part I. The
vertical distribution of the Macoma community.

Marityn T. VASSALLO

Vol. 11; No. 4

The egg capsules of Jenneria pustulata (LicutFoor, 1786),
with notes on spawning in the laboratory.
GCrARTES IN DPAISARO) sleet eine errr 182
The egg masses and veligers of southern California saco-
glossan opisthobranchs.
IRIGEFARDSWaAGREENE, | ciclele ela ateyeeietale nie sole « 100
The interrelationships of certain boreal and arctic species
of Yoldia MOuuErR, 1842.
I. McT. Cowan
The Littorina ziczac species complex.
Tuomas V. Borkowski & MariLyNNn R. BorKowsKI
VN Micrel sien cfooeon cconetet ays tenet ex cicss 410
The Muricidae of Fiji. - Part II. Subfamily Thaidinae.
WALTER OLIVER CERNOHORSKY ~~... ....- sss 293
The rediscovery of Voluta (Lyria) grangeri SoweRBy **,
1900.
CLIFTON STOKES WEAVER & JOHN ELEUTHERE DuPont
MERE apbsLb SEIEO ao tae noe een 36
The role of behavioral traits in influencing the distribu-
tion of two species of sea mussel, Mytilus edulis and
Mytilus californianus.
JPM EEETARGER fo ire Siat tet ciwialeevers Set oesecat 45
The shell structure and mineralogy of Chama pellucida
BRODERIP.
Joun Davm Taytor & WiLuIAM JAMES KENNEDY
5 6S dota tid dtn. Ste aD CIE aie ani 391
‘The taxonomic significance and theoretical origin of sur-
face patterns on a newly discovered bivalve shell
layer, the Mosaicostracum.

(Conc Isl, BAIGNSION, ~~ 5 coos odoucucob0bodee 185
The type of Tegula funebralis (A. ApAms, 1855).

RU DOEISTORICERS 97sec. ois occ cele wie ose meneueveni ee « 408
Two new cypraeid species in the genus Erronea.

CRAWEORDINS@ ATE) (anaes ciomeeeaelse - 256
Two new records of Cratena abronia.

STEVE N@ RON Glue siete etct-rs taiee. ay ci oncyas es ae ehthuel « 281

Two new species of the genus Volva R6pinc, 1798 (Ovu-
lidae FLEMING, 1828).
CRAWFORD NEMU(@ATE =a )jrec iis eicaei- oie 6 <1oiae 364
Use of the propodium as a swimming organ in an ancil-
lid (Gastropoda: Olividae).
BARR Wa RG AVVAESONG) 9-2 «i tistoie’e; els! ays lode efoynieaees re 340
Volutocorbis andFusivoluta, two genera of deepwater Vo-
lutidae from South Africa.

ARAL D YAR INEFIDER GS sie eie cic sie sisters ave eine ie 200
What is Macoma truncaria DALL?

PAW GEN IPaN CONN on canner citsis orecicuslin neva recsiuecc 281
Zoogeographical studies on living cowries.

RAN ZATEREDISCHIEDER] Sl cece sone 367

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Page V

AUTHOR INDEX
AT DRICHs EREDERICKSAG eG. C2 LU) ee eee 70
ANbrREWS, Jay D., D. Turczon & M. HreHa gop a
BAKUS;)GERAUD Ie]... 2 sisre- systole eienciz ei esata eererire 93
IBERTSCHAEIANSH! > ys tysrecicln cos aca ciak ne et 431

Borkowsk1, MariLyNN — see BorKowskI, THOMAS V. &

BorkowskI, THomas V. & M. R. BorKowskI .. 410
IBOSSSISENNETH 2) saree hess @ecieeies oes (150), (445)
IBRATGHER MEWIEAL | sarc cusses seni ete eae 334
BRENNAN, ELLEN — see DUSHANE, HELEN &
IBURINMSROBERT sp erascaresyoeaa eeeraitrs cue seus encletes rere 90
CALABRESE, ANTHONY & Harry C. Davis... 235
@ARTISER JOH NGG) Il GenecalGas ng is a ccnsian sine 235i
CARTON, WPS MES Wa | toyecistsie gets ie erenarens orev ateisvonve 283, 284
CATE CRAWEORDPNEIDT I) ae etieieciee sce 80, 256, 364
GATHER PEAING@ MINT 9 Boo rates ck isratec ch onotste os youll cteleraueinee 85
CERNOHORSKY, WALTER OLIVER _............ 210, 293
COANE UGENE: Ver ol aaancian on tenho ee 277, 281
see also: Rotu, BARRY &
COWANPIIRUM Gis rai, 3h cease ae tote cine eiuonsereans 51
Cowan, I. McT.« James H. McLean __.......... 105
Crovo, Mrs. L. E. see: Otsson, AXEL A. &
IDZNSAR Os CETARTE ESB Ne mums icuseys wey acyeieteiccceseeteusion tele 182
D’Artitio, ANTHONY see: EMERSON, WILLIAM K. &
Davis, Harry C.& HERBERT Hmu ............. 316
see also: CALABRESE, ANTHONY &
IDEN PAIVOAIND st] Pn Tis lanes on MSIE ERG rr eG Cena Goin aie 198
D UGE RE ONAGH MEE eerercrie sctovcecianveun eet te socie 415
DuPont, JoHN ELEUTHERE see: WEAVER, CLIFTON S. &
DU SEAN SE RISEN Pte peters e orelene, sicvarciain Sieve: pause ee 343
DuSHANE, HELEN & ELLEN BRENNAN ......... B5il
LBD WARDS, 1D), CRN cooooncosoonvogccoconcdc 326
EMERSON VMI TAO Ke titers ere 1, 33, (444)
Emerson, WILLIAM K. « A. D’ATTILIO . 195, 324
EmeErSoN, WiLtiAM K. & W.E. OLD, Jr. ...... 98
ISISIEDPRAPIGARIKSISVINE 9 | ce Sis tevalereosseie see aide aitvs eee soso 434
IESRVANN AK ORR VV Sie ite a. or eels erica ai reyecoyesreseeh evmvekere taterens 440
GOSTINERMOEERRENCHEa erence se ceiene aie 147
GREENE VRIGEVARD) WW alanine secs oa sien vere ccs eicineeiels. 100
(Cone, CrUNBuIS |, - Wooenoodcancsvovouusoscns 140
lslaawmmenom, Gionen, 15 ~§ soosoounesocsocne0dooc 185
ls leineie, Io Iolo, “epocepeccnpggouscnedonbasu 45
ARR YZ g VAR OLD: Wo aiitsers a cre ocala oreuereceryctereceue ens 164
HENDLER, GorpDon, see: LovELAND, RoBErT E. &
THERTEEINS LEORGH o inrierc leis ss «rena cteiare 80, (149)
Hinu, Hersert, see: Davis, Harry C. &
Housrick, JosEpH RICHARD ............ 2.2.44. 4

Hrena, Marian, see: ANDREWS, J. D., D. TURGEON &

[psisisisies WW HOIVEIOS Geo abeonecapdaocoteceuac 144
JOEINISONMKGAY MIN | fens cce cre lesion terior cron 272
ICEEN WAWeMiVRAS © 9 perv creniesieameerecte (83), 439

KENNEDY, WILLIAM J., see: TAYLor, JOHN Davip &

Page VI

IKENINY, RON) oJ) giesssatiea ghecetay tale aitacweriaagedseuses 336

LEONARD, VERNON KENNETH, Jr. ...... «200s 382

ILONGHS DEVENS) Eo Sei cetacean er 281
see also: RoLLER, RicHarp A. &

IL@OGANOme, WiemOR IL; ‘oc agbvoscucdd00b00000s 153

LovELAND, Rosert E.,G. HENDLER & G. NEwxirK 418
Lu, C. C. see: ALDRICH, FREDERICK A. &

IMiAcmoRaTm, Xmas 1B Salo do oo dauoacceac0000 399
MacGinitig, G. E. & Nett MacGIniTiE ......... 59
MacGiniriz, Nettie, see: MacGrnrriz, G. E. «&

INPSWURERS DON Me ccs cie s,s 8 eacaneisee civesueucrs ceoraen ie 243
Itoh syereae, IOS WWE) Gogenddoobancdscodcdc0 du: 145

McGowan, JouN A. & TAKASHI OKUTANI pees 40/2
McLean, JAMEs H., see: Cowan, I. McT &

Miuer, RicHarp L.
Moore, Donatp R. & Donatp J. REISH
Newkirk, Gary, see: LovELAND, R. E., G. HENDLER &

INKZDAIRON, |/IMIIS WWE se oneboGocconoG0Kg0000 50
OxuTAnl, TAKASHI, see McGowan, JoHN A. &

Op, WittiaM E., Jr., see EMERSON, WILLIAM K. &
@OESEN MDA VEDA: is ea ener epe-ares ote euevey ogsveueereys 135
Otsson, AxeL A. & Mrs. L. E.Crovo _......... 31
IREEDER EARATDIAG. (018s cerhere ere cysts merce euereaete 200
REM WROBERTEG#Bae she. ce cieeusnise tire cctaenon 378

ReisH, DonaLp J., see Moore, Donatp R. &

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Vol. 11; No. 4

ROBIELTARDs GORDON TAU Eee eee Eee ere 289
ROLLER; -RIGHARD) Ay =). 22 odes omen 280, 421
Router, RicHarp A. & STEVEN J. LonG _.......... 424

- ROSENTHAL, RICHARD J..05 {ya ec oe yee 288
ROSEWATER, JOSEPH. 7) jiecitiei cise eo sees ere 345
RotuH, Barry & EUGENE V.CoAN _............««- 62
SCHIEDER ERANZVATERED eee erie 34, 367

SCHILDER, FRANZ ALFRED & MARIA SCHILDER poo IO)
ScHILDER, Marts, see: SCHILDER, FRANZ ALFRED &

SmirH, Attyn G9 7 visas: a eee eee 117
SMITH; BRYAN Js.) sG050 siete clei ee eee 39
SOLEM, ALAN = 2° sche a bheteeiele aps st oeeeeneds ete 24

STOHLER, RUDOLF (84); (lol); (156) Zao aie
(292), 408, (444)
TayLor, JoHN Davin « W. J. KENNEDY
TERRY, JUDITH Sd sists ois. bree eee 42
TurcEoNn, Donna, see: ANDREWS, J. D., —, & M. HREHA

WASSALLO; MARILYN DT; 9 3 acla nce aGeneeee 223
WEAVER, CLIFTON S.& JOHN E. DuPont... 36
WEBSTER, STEVEN K. | ni. sate os eee eee 121
Witson, BARRY Ri sued ace oa oe eee 340
WINKEER, Linpsay R. 1 adc anlecen one ones 268
Wore; DouGiAS S.o. is has sete ase See 126

Wo trFson, Fay H.

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME II Jury 1, 1968 NUMBER I

CoNnTENTS

Familial Placement of Hanetia JoussEAUME, 1880 (Muricidae) and Solenosteira
DALL, 1890 (Buccinidae) (Plate 1; 2 Text figures)
WiLuiaM K. EMERSON

Lael

A Survey of the Littoral Marine Mollusks of the Caribbean Coast of Costa Rica
(1 Text figure; 1 Map)

JosEpH RicHARD Hovusrick

4
“Ptychodon” misoolensis ADAM & VAN BENTHEM JUTTING, 1939, A New Guinea
Strobilopsid Land Snail and Review of the Genus Enteroplax (1 Text
figure; 1 Map)
PNWANS OLE Meemmmrermne min Wire Wirt iul ona tan Ne sh oua itis (eb cat a) iene cl ey taille), 2 D4
Observations on Aquarium Specimens of Oliva sayana RAVENEL (Plate 2)
AXEL ACL OUSSONMEMVITSs ecm Een GROVOW GE) 62%.) be) eta) lceeminhe el ie ley ieee. a QT
A Record of the Indo-Pacific Cone, Conus ebraeus, in Guatemala
VBE 1S ERTIRSOR, go 6 igh Bo wot D ouoy la G86" fo) top eaaeo lara acienes
A Cowrie Mutant from the Gulf of Thailand (1 Text figure)
BRAN ZWALFEREDI SCHILDERVY aricte. um noe) s. eueil te qe ayios jl, ei Siel 6 8 GA
The Rediscovery of Voluta (Lyria) grangeri SowERBY 3°, 1900 ~=«(Plate 3; 1 Map)
Cuirton Stokes WEAVER & JOHN ELEUTHERE DUPONT. . . ..... . « 36
A List of Types of the Family Volutidae Held by the National Museum of Victoria
BRIAN Pp SMUD Eee rst ie rey dol ene MRC TaUn REN lhe) es dey asi lle’ ie hts. 6) <3) 3Q
Mediargo, a New Tertiary Genus in the Family Cymatiidae (Plate 4)
ODI See UERR Ya umes Mii Cnc memm Ee aie FS se else) ae ae A

[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 11: $18.- Domestic; $18.80 in the Americas; $19.20 in all other Foreign Countries
Single copies this issue: $8.-. Postage extra
Send subscription orders to Mrs. JEAN M. Cate, 12719 San Vicente Boulevard,

Los Angeles, California 90049. Address all other correspondence to Dr. R. STOHLER, Editor,

Department of Zoology, University of California, Berkeley, California 94720
Second Class Postage paid at Berkeley, California

ConTENTs — Continued

The Role of Behavioral Traits in Influencing the Distribution of Two Species of
Sea Mussel, Mytilus edulis and Mytilus californianus (3 Text figures)

J. R. E. Harcer

Notes on the Food of Conus dalli STEARNS, 1873
James W. NyBAKKEN ,
The Interrelationships of Certain Boreal and Arctic Species of Yoldia MOLLER, 1842
(Plate 5; 9 Tables)
I. McT. Cowan

Notes on Cryptochiton stellert (M1ppENDoRFF, 1846) (Plate 6)
G. E. McGinirie « Nettie McGIniTIzE

Further Observations on the West American Marginellidae With the Description of
Two New Species (Plate 7; 2 Text figures; 1 Map)
Barry RotH & EucGENE Coan

On an Octopod from Placentia Bay, Newfoundland (Plate 8)
FREDERICK A. ALpricH « C. C. Lu

A New Species of Enoploteuthid Squid, Abraliopsis (Watasenia) felis, from the

California Current (Plates 9 and 10; 1 Map)
Joun A. McGowan & TakKAsHI OKUTANI
NOTES & NEWS Be ee Gea ee MO NED soils aL RAC A cc
Tellina ulloana, A New Species from Magdalena Bay, Baja California,
Mexico Leo G. HERTLEIN
An Emendation Crawrorp N. Cate

BOOKS, PERIODICALS & PAMPHLETS

. 80

. 83

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

Vol. 11; No. 1

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Page 1

Familial Placement of Hanetia JoussEAUME, 1880 (Muricidae)

and Solenosteira DALL, 1890 (Buccinidae)

WILLIAM K. EMERSON

Department of Living Invertebrates
American Museum of Natural History
Seventy-ninth Street and Central Park West, New York, New York 10024

(Plate 1; 2 Text figures)

INTRODUCTION

THE GENERIC NAME Hanetia was proposed by Jous-
SEAUME (1880, p. 335) with Murex haneti Petir (1856,
p. 90, plt. 2, figs. 7, 8) the type species by tautonymy
and monotypy. Unfortunately, Petir’s taxon was briefly
described and poorly illustrated on the basis of specimens
stated to have been collected by Hanet Cléry from the
‘coast near Rio de Janeiro, Brazil. As Wooprinc (1964,
p. 256) has recently stated, despite the questionable iden-
tity of Petit’s Murex haneti and the lack of confirmation
of the Brazilian locality, Hanetia JoussEAuME, 1880, has
been employed in recent years for the distinctive group
of New World fossil and living buccinids for which DaLi
(1890, p. 122) had later proposed the generic name
Solenosteira, with the Panamic species, Pyrula anomala
Reeve, 1847, the type species by original designation.
This group of cantharid-like gastropods is well repre-
sented by numerous named fossils, ranging in age from
late Miocene to late Pleistocene, that are known from
Florida, Panama, Ecuador, Peru, and the west coast of
Colombia, Costa Rica and Mexico. Several Recent species
survive in the warm waters of the eastern Pacific, ranging
from Mexico to Peru (see Keen, 1960, pp. 400 - 402).
Evidence is presented in the present study to demon-
strate that Hanetia JoussEAUME, 1880, is referable to the
Muricidae, and therefore, is not a senior synonym of
Solenosteira Datu, 1890, which is referable to the Buc-
cinidae.

DISCUSSION

Although Petit’s original figures of Murex haneti, the
type species of Hanetia, suggest an inflated, strongly

ribbed specimen of “Pyrula” anomala Reeve, 1847 (Plate
1, Figures 1 - 3), similar specimens were not subsequently
reported from Brazil, or elsewhere in the western Atlan-
tic. Thus the identity and the western Atlantic occurrence
of Petit’s taxon could not be confirmed. MaxweELi SMITH
(1939, p. 16; plt. 12, fig. 7), however, illustrated a beach
worn Brazilian shell as “Tritonalia haneti Petit,’ which
does not compare favorably with Petit’s illustrations.
According to Wooprinc (1964, p. 256) this specimen
was collected by von Ihering and was deposited in the
U.S. National Museum (No. 150767). Lance DE Mor-
RETES (1949, p. 94) subsequently recorded “Uvosalpinx
haneti (Petir)” from Rio de Janeiro, citing Prtit’s
original locality, and he reported this taxon from Santa
Catarina Island, Florianopolis, Brazil, citing specimens
collected by von Ihering.

At my request, Dr. Bernard Salvat of the Muséum
National d’Histoire Naturelle, Paris, kindly provided me
with photographs of the type of Murex haneti Petit
from the collection of his institution (Plate 1, Figures 4
to 6). The holotype was found to be referable to speci-
mens identified as “Urosalpinx haneti Petrr” that were
collected by Dr. Bernard Tursch in 3 fathoms off Rio de
Janeiro, Brazil (Plate 1, Figures 7, 8) and were depos-
ited without soft parts in the collection of the American
Museum. Through the courtesy of Mr. Masao Azuma of
Nishinomiya, Japan, a drawing of the radula of a small
specimen labeled “Murex haneti Petr’ from Governa-
dor Island off Rio de Janeiro, Brazil (ex—U. S. N. M. No.
364129) was provided me (Text figure 1). The radula,
although from an immature specimen, is that of a muri-
cid, whereas the radula of “Pyrula” anomala (REEVE,
1847) is that of a buccinid (Text figure 2).

THE VELIGER

Vol. 11; No. 1

Figure 1

Radular rachidian tooth and lateral teeth of “Murex” haneti

Petit, 1856, greatly enlarged; from a juvenile specimen collected

by Dr. Waldo L. Schmitt at Governador Island, Brazil, ex U.S. N.
M. No. 364129 (A. M.N.H.No. 142443)

On the basis of shell morphology, and opercular and
radular characters, Murex haneti Petit, 1856, is, there-
fore, referable to the Muricidae, subfamily Tritonaliinae
[= Ocenebrinae]. Hanetia JouSSEAUME, 1880, is avail-
able for Murex haneti and possibly for other Urosalpinx-
like species with similar radular and opercular charac-
ters that are placed at the present time in Urosalpinx
Stimpson, 1865, or in other genera. Urosalpinx rushiu
Pitssry (1897, p. 297), a taxon described from Mal-
donado Bay, Uruguay without an illustration, may be a
junior synonym of Murex haneti Petit, 1856.

Solenosteira Dax, 1890, thus can be retained, as it is
the earliest available name for the Cantharus-like Buc-

Figure 2

Radular rachidian tooth and lateral teeth of Solenosteira anomala

(REEVE, 1847), greatly enlarged; from a specimen collected by

Ben and Ruth Purdy at Puerto Pefiasco, Sonora, Mexico (A. M.N.

H. No. 141801). Drawings courtesy of Mr. Azuma; redrafted by
A. D’Attilio

Explanation

Figures 1-2, 4-8: “Murex” haneti Petit, 1865. Figures 1, 2:
copy of Perrit’s (1865, plt. 2, figs. 7, 8) original figures; Xt.
Figures 4, 5, 6: holotype in Muséum National d’Histoire Naturelle,
Paris; photograph through courtesy of Dr. B. Salvat; <1. Figures

cinidae of the New World that had been erroneously
assigned to Hanetia JoussEAUME, 1880. The generic
name Fusinosteira Ousson (1932, p. 179), type species,
by original designation: Purpura fusiformis BLAINVILLE,
1832, of the Panamic faunal province, also is available
for angulated species of Solenosteira, should these prove
to be a biologically valid group.

ACKNOWLEDGMENTS

In addition to Dr. Bernard Salvat and Mr. Masao Azuma,
I am greatly indebted to the following friends and col-
leagues for courtesies of various kinds: Mr. Anthony
D’Attilio, Mrs. Dorothy Germer, Dr. Miguel A. Klap-
penbach, Mr. Henry R. Matthews, Mr. William E. Old,
Jr., Mr. & Mrs. Ben Purdy, Dr. Joseph Rosewater, Dr.
Donald R. Shasky, and Dr. Bernard Tursch.

LITERATURE CITED

BLAINVILLE, HENR1 Marie DucrotTay DE
1832. Disposition méthodique des espéces récentes et fossiles
des genres Pourpre, Ricinule, Licorne et Concholépas.
Nouv. Ann. Mus. Hist. Nat., Paris 1: 189 - 263; plts. 9-12
Dati, WiLtiAM HEALEY
1890. Contributions to the Tertiary fauna of Florida.
Pt. 1, Pulmonate, opisthobranchiate and orthodont gastropods.
Trans. Wagner Free Inst. Sci., 3: 1-200; plts. 1 - 12.
JousSEAUME, FELIX PIERRE
1880. Division méthodique de la famille des purpuridés.
Le Naturaliste, Paris, 1 (42): 335 - 336
Keen, A. Myra
1960. Sea shells of tropical west America, 2%? printing.
Stanford, Calif. (Stanford Univ. Press)
Lance DE MorreETES, FREDERICO
1949. Ensaio de catalogo dos molluscos do Brasil. Arq.
Mus. Paranaense 7: 5-216 Curitiba, Parana, Brasil
Oxsson, AxEL ADOLF
1932. Contributions to the Tertiary paleontology of Northern
Peru: Part 5, the Peruvian Miocene. Bull. Amer. Paleo.
19 (68) : 1-272; plts. 1 - 24 (30 June 1932)
PETIT DE LA SAUSSAYE, S.

1856. Description de coquilles nouvelles.
Paris, 5 (ser. 2, 1): 87-92; plt. 2

Journ. Conchyl.

of Plate 1

7, 8: mature specimen from 3 fathoms off Rio de Janeiro, Brazil,
Dr. B. Tursch coll. (A. M.N.H.No. 129117); X2.

Figure 3: Solenosteira anomala (REEVE, 1847), copy of REEVE’s

(1847, plt. 3, figure 12) original figure of “Pyrula” anomala; X1

TuHeE VELIGcER, Vol. 11, No. 1 [EMERSON] Plate 1

s

Figure 1 Figure 2 Figure 3

Figure 4 Figure 5 Figure 6

Figure 7 Figure 8

WolkeilesINow! THE VELIGER Page 3

Pi_tspry, HENRY AUGUSTUS

1897. | New species of mollusks from Uruguay. Proc.
Acad. Nat. Sci. Philadelphia 49: 290 - 298; plts. 6, 7
Reeve, Lovett AucustTus
1847. | Conchologia iconica; or illustrations of the shells of
molluscous animals. London, Monograph of the genus
Pyrula. 4: 9 plts. + text
SmirH, MAXxwELL
1939. An illustrated catalogue of the Recent species of the
rock shells. Lantana, Florida: 83 pp.; 21 plts.
Stimpson, WILLIAM
1865. On certain genera and families of zoophagous gastero-
pods [sic]. Amer. Journ. Conch. I (1): 55-64; plts. 8-9
(25 February 1865)
Wooprinc, WENDELL PHILLIPS
1964. | Geology and paleontology of Canal Zone and adjoining

parts of Panama. U.S. Geol. Surv. Prof: Paper 306-C:
238 - 297; plts. 39 - 47

Page 4

THE VELIGER

Vol. 11; No. 1

A Survey of the Littoral Marine Mollusks

of the Caribbean Coast of Costa Rica

BY

JOSEPH RICHARD HOUBRICK

Division of Science and Mathematics , St. Leo College, St. Leo, Florida 33574

(1 Text figure; 1 Map)

INTRODUCTION

THE COMPOSITION OF the marine molluscan fauna of the
western Caribbean is relatively unknown. For a period
of six weeks the writer was able to study and collect
marine mollusks in two different areasof Limon Province,
Costa Rica. The purpose of this work and of the re-
search that followed was to investigate the composition
and range of the fauna in order to compile a list of the
Recent species, to observe their ecological distribution,
and to provide some basis for future study in this little-
known section of the Caribbean region.

Although the period was barely sufficient to allow for
an exhaustive survey of the mollusks and the entire range
of the variety of their habitats, a good portion of the
littoral or shallow water fauna is believed to be recorded.
However, more complete and sophisticated collecting
methods in these areas would undoubtedly extend the
list of existing species and provide further pertinent
information.

The marine mollusks of the Caribbean coast of Costa
Rica have received little or no attention from recent
workers. Indeed, the whole of the Caribbean coast of
Central America has not received adequate scientific in-
vestigation and is poorly known in relation to other areas
of the Caribbean Province.

With the prospect of a future sea-level canal being
constructed somewhere in the Central American isthmus
an investigation of the marine fauna of both the Carib-
bean and Pacific sides is desirable. A sea-level canal
would make possible a limited mixing of the Caribbean
and Panamic faunas for the first time since the late
Miocene.

The area near Puerto Limon presented the most favor-
able base for living, transportation facilities, and collect-
ing. Consequently, the major portion of this paper deals
with the marine mollusks from this vicinity.

ACKNOWLEDGMENTS

It is a pleasure to acknowledge at this time Dr. Donald R.
Moore of the Institute of Marine Science, University of
Miami, whose help, interest and suggestions throughout
this project have been invaluable. I am also indebted to
Drs. William Clench and Kenneth Boss who kindly read
the manuscript and offered suggestions. Gratitude is also
due to Dr. Axel A. Olsson and Mr. Ted LaRoe who
assisted in the identification of some of the fauna.
Finally, I am indebted to the Organization for Tropical
Studies for the opportunity to collect in some of the
areas of Costa Rica and to St. Leo College for both
financing the field work and providing leave time for the
research involved.

PREVIOUS WORK
on WESTERN CARIBBEAN MOLLUSKS

The literature dealing with the marine mollusks of the
Caribbean coast of Central America is rather sparse. This
important portion of the Caribbean Province is perhaps
the most poorly known by way of actual collection and
study. Otsson & McGinty (1958) believed that the
principal reason is the scarcity of convenient collecting
stations along the Caribbean coast as compared to the
abundance and ready accessibility of such grounds on the
Pacific side.

The earliest literature on the Recent fauna deals with
the mosquito coast of Nicaragua and consists of several
short articles and notes published by FLuck (1900-1901;
1905). The next reference to the area occurs in 1958
with the work of OLtsson & McGinty in Panama. This
seems to be the only substantial report we have about
this area. The paper consists of a faunal list along with
the description of some new genera and species. Recently,

Vole It-sNo; 1

Baxus (1968) has published a comparative study of
littoral zonation of the gastropods on the Pacific and
Atlantic coasts of Costa Rica in which a brief account
of the Limon area is given with a list of the mollusks
collected at Portete. With the exception of the above
paper and the distributional records listed in Johnsonia,
nothing more seems to be recorded of the marine mol-
lusks of the Caribbean coast of Costa Rica.

Two publications dealing with the southwestern Carib-
bean fauna are those by Coomans (1958) and REHDER
(1962), the former on the gastropods of the Netherlands
Antilles and the latter on the Los Roques mollusca.

The fossil mollusca of the western Caribbean have re-
ceived more attention by workers. Gasp (1881) and Oxs-
son (1922) have reported on the Pliocene and Miocene
molluscan faunas of eastern Costa Rica. WooprINc (1957)
has investigated the Canal Zone and Weisporp (1961;
1964) discussed the late Cenozoic mollusca of northern
Venezuela and provided an excellent bibliography of the
Caribbean fauna.

In conclusion, a survey of the literature reveals that
knowledge of the Recent western Caribbean fauna is
surprisingly incomplete and spotty. Further research is
needed to gain a full picture of this sector of the Carib-
bean Province.

REGIONAL GEOGRAPHY
AND GEOLOGY

The Caribbean slope of Costa Rica is composed of
Cenozoic strata which dip away from the central moun-
tains to the sea. The eastern coastline is slightly concave,
lacking large bays or inlets. The Atlantic coastal plane
occupies nearly one-third of Costa Rica and is almost
flat. Offshore, the narrow shelf which is rarely more than
10 miles wide, slopes to a depth of 600 feet.

The climate of this region is tropical with a heavy
rainfall. Most of the northern coast of the province of
Limon is low and covered with dense forest interlaced
with rivers, creeks, brackish estuaries, and swampy areas
of Raffia palms. The beaches are broad and of a gray
volcanic sand. An excellent description of this coastline
is given by Carr (1956).

According to Otsson (1922), there was a complete
inter-oceanic connection in this region during the lower
Miocene, and nearly all of the Costa Rican isthmus was
then beneath the sea. At the close of this epoch, eastern
Costa Rica was again emergent for a short period. During
the upper Miocene the whole of Costa Rica as well as
Panama were in an uplifting process. This elevation de-
stroyed the inter-oceanic connections and culminated in

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Page 5

the early Pliocene. Thus, during the whole of the Terti-
ary, Costa Rica underwent great changes due to periods
of submergence and uplift of its land masses.

In contrast to the northern coastline, the central Limén
peninsula is of a coralline limestone formation. OLsson
(1922) and ScHucHeErT (1935) both believed that this
outcrop belongs to the Gatun formation. This area is very
rich in fossiliferous marls, and Gasp (1881; 1881 a)
described about 70 species of fossil mollusks from the
Limon area. O_sson (op. cit.) described 334 species
from the Gatun formation.

FAUNAL ORIGINS anp RELATIONSHIPS

The warm tropical waters of the western Caribbean
support a molluscan fauna similar to that of the West
Indies and the Florida Keys, and are a portion of the
zoogeographical area known as the Caribbean Province.
WarMKE & AppoTT (1961) presented a map showing
the extent of this province. Conditions are fairly uniform,
allowing many species to exist throughout the province,
and the molluscan fauna is impressive. AppotT (1962)
remarked that the larger West Indian islands have over
1200 species of marine mollusks, while the smaller iso-
lated coral islands, poor in food, have only about 350
kinds of mollusks. Moore (personal communication)
found a depauperate fauna in the Albuquerque Cays off
Nicaragua. Otsson « McGinty (1958) recorded about
543 species and subspecies from the Panama coast. We
report 250 species herein from Costa Rica. The lower
numbers of species from the western Caribbean may
indicate a poorer fauna or reflect the inadequate work
done in this region.

Many species of the Caribbean Province have close
affinities to those of the Panamic Province. C. B. ADAMS
(1852) was the first to point out the similarities and to
draw up a list of analogous species between the two
provinces. Dati (1912), in his study of the fossil shells
from eastern Panama and Costa Rica, noted that these
mollusks were mostly of common Caribbean living forms,
but found an interesting admixture of species now living
on the Pacific coast. Later, ScHUCHERT (1935) remarked
that the Miocene shell faunas of Central America were an-
cestral to the living ones of the Caribbean and to some in
the Pacific. Otsson (1922) stated that “. . . thirteen
percent of the Gatun fauna is identical or closely re-
lated to the recent species the majority of which are
found living along the Caribbean coast, certain others only
on the Pacific side, and a few common to both.” He
found 6 species to be specifically identical with forms in
the Pacific and 18 others that showed close relationships.

Page 6

There were probably two natural seaways crossing Cent-
ral America during the Miocene, the main one across
Costa Rica and a second through the Darien of Panama.
This permitted free circulation of water between Atlantic
and Pacific oceans and it was in these warm seas that the
Miocene faunas developed which were ancestral in part
to the bulk of modern Caribbean and Panamic faunas.
Keen (1958) observed that the sizeable number of
parallel or analogous species on either side testify both
to the slowness of change in the isolated stocks and to
their past relationships. Orsson & McGinty (1958)
and Orsson (1961) pointed out that the fauna has re-
mained little modified in the Pacific section while the
Caribbean has undergone a greater change through a
major extinction of many of the older Miocene groups
due to the drainage of large tracts of its shallower parts
during mid-Pleistocene glaciation in the northern hemi-
sphere. Other groups later invaded this area from the

PACIFIC

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Vol. 11; No. 1

north and east, and present day Caribbean mollusks,
when compared to the rich Miocene fauna, appear to be
impoverished.

COLLECTING LOCALITIES

The two areas discussed in this paper are located in the
province of Limén, Costa Rica. A map of the area is
given in Figure 1. The station most thoroughly worked
was that of Portete, a small fishing settlement on the
shores of a lagoon located a few miles to the north of
Puerto Limén. This is in the central portion of Costa
Rica’s Caribbean coastline, more precisely at Lat. 10°N,
Long. 83°02’W. The whole area consists of a projection
of the land into the Caribbean, the coastline being of
a coralline limestone formation and somewhat elevated.
It is surrounded by moist tropical forest with a network

Figure 1

Vol. 11; No. 1

B3°4' 30"

83°0'00"

83°1'00"

83°1'30"

83°2'00"

STATION No. |

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83° 4'00"

——— @ THALASSIA BEDS

Page 7

8523' 30:

(@) ROCKY AREAS
@) TIDAL CREEK ZONE
@) MUD-SAND BOTTOM

PORTETE

Figure 2

of brackish water lagoons and Raffia palm swamps.

Collections and observations at Portete were made
during the dry season in March of 1965 for a period of
3 days. The same station was more thoroughly collected
for a period of one month during July of 1966, the rainy
season.

The second station, Barra del Colorado, will be treated
only briefly because it was worked but for a few days. It
is located in the northeast portion of Costa Rica at the
mouth of the Rio Colorado, Lat. 10°40’ N, Long.
63-35) W.

Collecting was confined to the shallow water or littoral
fauna in both stations and included beach specimens and
drift material. A face mask was used in the shallow
areas and on the reefs. Algae were sorted and picked
through for the smaller species and drift material was
selected and carefully sorted with the aid of a stereo-

microscope for the smaller micromollusks. The Thalassia
beds and sandy bottoms were sifted with + inch mesh
strainers. When possible, specimens were obtained from
local fishermen. No dredging was undertaken, but this
would undoubtedly have increased the faunal list by
adding more burrowers and pelecypods.

COLLECTING STATIONS

STATION 1: PORTETE

This area (Map, Figure 2) consists primarily of rocky
shoreline, surfwashed coves and a somewhat protected
lagoon. A small freshwater creek empties into the lagoon
and to the north and south of this station there are rivers
emptying their muddy contents into the sea. Rainfall

Page 8

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Vol. 11; No. 1

can be heavy during the wet season and is undoubtedly
a factor in the salinity of the shallow portions of the
coast at this time.

Unfortunately, quantitative data for many of the eco-
logical factors of this area are not known. However, the
author’s short experience in the area produced some
qualitative information which indicates some factors of
possible ecological significance and will serve as a basis
for future investigation.

No record of the variations of sea temperatures was
compiled, but temperatures appear to be typical of
the shallow waters of the Caribbean. Values of the oce-
anic quadrant 10- 15° N, 80-85° W have been used.
In this quadrant, sea surface temperatures average 27.3°
C with a range of mean monthly values of 2° (D.R.
Moore, personal communication). Very shallow areas
become much warmer during the day due to the effect
of the sun. The same is true of the local effects of
cooling due to the heavy rains in the wet season. Varia-
tions, however, do not appear to be.of any significance
in the distribution of the common mollusks.

Due to the prevailing trade winds (E — NE) there is
a consistent wave action. Periods of severe storms cause
wave action to become sufficiently intense to damage the
plant and coral growth and to destroy certain ecological
environments. In sandy areas the author observed mol-
lusks torn from their substrate and cast up on the
beaches by heavy surf action. Wave action seems fairly
consistent around this station.

Tides at Portete are diurnal, having a range of 1.2 feet
and a mean tide level of 0.3 feet. There are no noticeable
strong local currents at this station, but currents related
to the wind nearly always flow W -— SW.

The ranges of salinity do not seem to be of a magni-
tude to be detrimental to coral growth and the typical
reef-dwelling fauna. The expected forms of mollusks were
found except along the margin of the lagoon area into
which the freshwater creek emptied. A water sample from
this station was taken and analysis of it showed a salinity of
34.083 ppt. Heavy rains can apparently change the salin-
ity in the more shallow portions along the shore and
increase the magnitude of the nearby river’s discharge.

Turbidity is influenced by the constant wave action.
There is a moderate amount of sedimentation contributed
by the rivers and creeks near this region. The portion of
the lagoon directly affected by the discharge of the creek
is devoid of the usual fauna. During periods of heavy
rain it appears to discharge considerable quantities of
sand and mud, creating a sand bar near its mouth.

Bottom conditions in Portete do not vary greatly. The
one exception is the lagoon area where substrate con-
ditions were extensively controlled and modified by flora
and fauna and were quite varied.

Substrate conditions at Portete were mostly of 3 types:
mud-sand bottom, ‘Thalassia-sand flats, and rock-coral
bottom. Intermediates and mixtures of these types were
present, some areas apparently in transition from one
type of bottom to another, but in general the variations
of substrates were strikingly defined. For the purpose of
study the writer divides the station of Portete into 3
general areas of substrate mentioned above.

A. Mud-Sand Bottom

The mud-sand bottom has a fine film of algal growth
covering the sediment and is found only in the center of
the lagoon; it extends back toward the shore near the
creek. The freshwater creek and its mouth are included
in this type of bottom. Few mollusks are found on this
substrate, but Strombus pugilis is abundant. Water usually
covers the mud-sand bottom to a depth of 5 feet or more.
No portions of this type bottom seem to be exposed at
low tide.

B. Thalassia-Sand Bottom

The Thalassia flats are all intertidal in nature and prove
to be the most rewarding areas in the number of species
encountered. The most extensive Thalassia beds are con-
fined to the south shores of the lagoon. They consist of
beds of Thalassia in the deeper portions which slope
upward, forming shallow flats of mixed Thalassia, Cau-
lerpa, Halimeda and Porites. The water is seldom more
than a foot deep here and with low tide there are periods
of incomplete exposure. Along the shore this area appears
sandy with large rocks and occasional Thalassia.

C. Rock and Coral Bottom

Rock bottom environments are the most common and are
usually covered with marine algae and corals. They are
exposed to the wave action and in some places heavily
pounded by surf. In areas of this substrate definite zones
characterized by certain flora and fauna are noticed. A
zone as defined by Ropricuez (1959) is a “band or
region in the shore defined by physical conditions.” Zo-
nation was, in many instances, reflected by the types of
mollusks found. It is most apparent in areas which are
affected by the tides. Substrate, wave action, and the
distribution of bottom types are lesser contributing fac-
tors in zonation. Lewis (1960) found that tides, with the
varying degrees of exposure they impose, are the primary
cause of zonation. This certainly appears to be the case
at Portete as zonation is remarkably constant in most
rocky areas. Mollusks of these zones show varying de-
grees of adaptation to tidal factors. That this is due to
their larval stages is evident, for the success of these
mollusks is not only due to their ability to maintain them-

Vol. 11; No. 1

selves as adults, but should be considered as the sum of
successes of the various stages in their life histories.

For purposes of convenience, rocky areas are divided
into 5 zones which seemed to predominate at Portete.
This partially follows the zonation system of ARNow,
St. Cra, & ARNow, 1963.

1. Splash Zone — the area above the mean high wa-
ter levels, including splash pools, areas affected by
spray and beach debris.

2. Intertidal Zone — the area between mean high
and mean low water levels. An area of loose rocks and
stones, sandy-bottom pools and shallow water and fre-
quently completely exposed by tidal action.

3. Rock-Ledge Zone — the subtidal shelf that
fringes the coast. It is transversed by cracks filled with
holes and niches. Heavy growths of Padina, Hypnea,
and Acanthopora and other algae are predominant
here. This zone is narrow in most parts of Portete;
however, in one area the rocky ledges are quite ex-
tensive.

4. Reef Zone — this zone is characterized by coral
growth and deeper water. At the edge of this zone
and along the outer edge of the rocky shelf zone are
heavy growths of Zoanthus. Further out Siderastraea
and Acropora coral are common. The hydrocoral
Milleporina is abundant in this zone. Wave action is
generally heavy.

5. Tidal Creek Zone — an unusual cleft cut through
the promontory which formed one side of the lagoon.
This promontory is heavily pounded by waves on its
outer, southern portion. The water thus enters a cleft
in the clifflike sides and forms a salt water creek
which eventually enters the lagoon and is subject to
tidal action. It is a protected zone, always in the shade,
and in some places the creek is almost 15 feet deep.
The sea urchin Diadema is common on the bottom.
Shallow areas are covered with a fine filamentous green
alga.

STATION 2: BARRA DEL COLORADO

An area of wide sandy beaches bisected by the estuary of
the muddy Rio Colorado, the Barra del Colorado is bor-
dered by wet, tropical rain forest. The beaches are broad
and slope gently into the turbid sea. This area is quite
similar to that of Tortuguero described by Carr (1956).

No ecological data were recorded, but substrate con-
ditions along the beaches are of the sand-bottom type.
No collecting was done in the estuarine areas, but KELso
(1965) found Polymesoda triangula, Mytilopsis zeteki,

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Page 9

and Neritina reclivata to be common in the Tortuguero
estuary further south.

This station afforded a number of species and habitat
records of mollusks not encountered at Portete.

SUMMARY or SURVEY
AND CONCLUSIONS

A checklist of the species collected in Limén Province
whose presence in the Western Caribbean was previously
unrecorded in the literature follows:
Arene tricarinata (STEARNS, 1872)
Astraea tuber (Linnaeus, 1758)
Tegula viridula (GmEuIN, 1791)
Puperita tristis (OrpicNy, 1842)
Parviturboides interruptus (C.B. Apams, 1850)
Solariorbis infracarinata Gass, 1881
Heliacus bisulcatus Orsicny, 1845
Heliacus infundibuliformis (GMELIN, 1791)
Petaloconchus irregularis (OrpicNy, 1842)
Caecum jucundum Foun, 1867
Caecum ryssotitum Foun, 1867
Caecum cycloferum Foun, 1867
Cerithium auricoma SCHWENGEL, 1940
Cerithiopsis pupa DALL & Stmpson, 1901
Cerithiopsis latum (C. B. Apams, 1850)
Cerithiopsis bicolor (C. B. Apams, 1845)
Triphora nigrocincta (C. B. ApaMs, 1839)
Triphora pulchellum (C. B. Apams, 1850)
Balcis intermedia CANTRAINE, 1835
Balcis conoidea Kurtz & Simpson, 1851
Eulima auricincta Apsott, 1958
Stilifer subulatus BRODERIP & SOWERBY
Risomurex roseus Reeve, 1856
Anachis pulchella Sowersy, 1844
Bailya parva (C.B. Apams, 1850)
Colubraria lanceolata MENKE, 1828
Nassarius nanus UstickeE, 1959
Pusia gemmata (Sowersy, 1874)
Marginella denticulata Conrap, 1830
Bullata ovuliformis (Orxicny, 1842)
Hyalina tenuilabra Tomutn, 1917
Crassispira fuscescens (REEVE, 1843)
Leptadrillia splendida Bartscu, 1934
Mangelia fusca (C.B.Apams, 1845)
Pyrogocythara coxi Farco, 1953
Retusa candei Orxgicny, 1841
Aplysia brasiliana Rane, 1828
Aplysia dactylomela Rane, 1828
Cingulina babylonia (C. B. ApAms, 1845)
Miralda havanensis Pitspry & AGUAYO

Page 10

Lobiger souverbit FiscHER, 1856

Tralia ovula (BrucutzrE, 1789)

Arca imbricata BrucutzRE, 1789

Pteria colymbus Ropine, 1798
Crassinella martinicensis (OrBicNy, 1842)
Condylocardia bernardi Daux, 1903
Mytilopsis domingensis (REcLUz, 1852)
Diplodonta notata Dat & Simpson, 1901
Codakia costata Orpicny, 1842
Acanthochitona rhodeus Pitspry, 1893
Octopus vulgaris Lamarck, 1798

Of the Recent mollusks found in eastern Costa Rica,
a total of nearly 250 species were collected consisting of
181 species of gastropods, 35 pelecypods, 3 scaphopods,
8 chitons, and 2 cephalopods. Table 1 illustrates their
composition.

Table 1

Composition of Molluscan Fauna

Fam. Genera Spec.

Prosobranchia 43 88 161
GASTROPODA _ Opisthobranchia 9 12 16

Pulmonata 1 3 3
PELECYPODA 19 30 35
AMPHINEURA 3 5 8
SCAPHOPODA 2 2 2
CEPHALOPODA 2

Twelve families are represented by only one species.
The families best represented are the Vitrinellidae with 8
genera and the Cerithiidae with 5. The best represented
genera are Rissoina with 9 species, Caecum and Cerithi-
opsis with 6 species each, and Nerita, Epitonium, and
Triphora, each represented by 4 species. Sixteen species
of Opisthobranchs remained unidentified. Thirty-five spe-
cies of bivalves were collected of which the largest shares
were of the families Arcidae and Tellinidae.

Table 2

Ecological Distribution
of Limén Province Marine Mollusks

1p G A st
Sand bottom species 11 0 0 3
Mud-sand bottom species 3 3 0 0
Thalassia-sand bottom species 7 38 0 0
Rock-coral bottom species 8 56 7 0

1 P = Pelecypoda;
S = Scaphopoda

G = Gastropoda; A = Amphineura

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2 2

Vol. 11; No. 1

A summary of the general ecological distribution of
mollusks collected alive follows in Table 2.

The mollusks collected in this survey have been depos-
ited in the following places: Division of Mollusks, U.S.
National Museum; Museum of Comparative Zoology,
Harvard University; Museum of the Institute of Marine
Science, University of Miami; and in the writer’s col-
lection.

SYSTEMATICS

The following systematic listing includes all identified
mollusks collected by the author in Limon Province, Costa
Rica. Some unidentified opisthobranchs and many micro-
mollusks have been omitted from the list. Although nearly
210 species were determined, this in no way implies a
complete survey.

In this checklist an attempt is made to record the
general distribution of each species throughout the Car-
ibbean Province. The stations and ecological areas where
the species were collected then follows, together with
additional information of interest. Many species were
not found alive and are indicated as beach drift or beach
shells.

The classification used in this paper is based upon
that of WarMKE & ApzorTT, 1961. The general geograph-
ical ranges of mollusks were taken principally from the
monographs in Johnsonia, from publications by WARMKE
& AppoTT (1961) and Aspott (1954; 1958).

Stations, areas and zones appear after the general range
information and are assigned symbols as follows:

P — Portete
m mud-sand bottom
t Thalassia-sand bottom
r rock-coral bottom

splash zone
intertidal zone
rockledge zone
reef zone

tidal creek zone

Sac lll
1

= Barra del Colorado

29 66

The terms “abundant,” “common,” etc. indicate the
relative occurrence of the species:

abundant — 100 or more
common — 15 to 100
uncommon — 6 to 15
rare — 5 or fewer

Species with doubtful determination are indicated by

a question mark in parenthesis: (?).

Vol. 11; No. 1

GASTROPODA

ScISSURELLIDAE

Scissurella Orsicny, 1823

Scissurella sp., P, beach drift. Rare.

FISSURELLIDAE
Hemitoma Swainson, 1840

Hemitoma octoradiata (GmeuIN, 1791). Range: south
Florida and the Caribbean to Brazil. P, r, 2. Found
during March in holes and pits on rocks. Uncommon.

Diodora Gray, 1821

Diodora listeri (OrsicNy, 1853). Range: southeast Flor-
ida and the southeast Caribbean. P, r, 2, 3. Uncommon
on rocks.

Diodora arcuata (SowerBy, 1862). Range: Florida, the
Bahamas and the Caribbean. P, t. Uncommon under
stones and dead Porites coral. Common in beach drift.

Fissurella BRuGUIERE, 1798

Fissurella nodosa (Born, 1778). Range: lower Florida
Keys, southern Mexico and the Caribbean. P, r, 3. Com-
monly covered with calcareous algae. Common.

Fissurella angusta (GmeEtIN, 1791). Range: the lower
Florida Keys, the Bahamas, British Guiana and the east
coast of Central America. P, r, 2, 3. Found living in
depressions on rocks and exposed flats. Common.

Fissurella barbadensis (GMELIN, 1791). Range: southeast
Florida, Bermuda, West Indies, and the Caribbean coast
of Central America. P,r, 2, 3. Common on rocks and
ledges.

ACMAEIDAE

Acmaea ESCHSCHOLTZ, 1830

Acmaea antillarum (Sowersy, 1831). Range: south half
of Florida to the West Indies and the Caribbean. P, r, 2,
3. Found in cup-like depressions on rocks and ledges ex-
posed to heavy surf. Common.

TROCHIDAE

Cittarium Puuiuipri, 1847

Cittarium pica (LinNAEuS, 1758). Range: Bahamas to
the West Indies and eastern Central America; north coast
of South America. P, r, 3. Commonly found adhering to
alga-covered rocks in surf-pounded places; younger spe-
cimens were common in the intertidal zone during the
month of March, but were absent in the summer.

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Page 11

Tegula Lesson, 1832

Tegula fasciata (Born, 1778). Range: southeastern Flor-
ida and the Caribbean. P, t. Found commonly on Thalas-
sia blades.

Tegula excavata (LAMARCK, 1822). Range: Caribbean.
P, r, 2. Found only during March under rocks or in
sheltered places. Common.

Tegula viridula (GmeE.in, 1791). Range: Central Amer-
ica and the northern coast of South America to Brazil.
P, t. Found in Thalassia beds. Uncommon.

Parviturbo Pitspry & McGinty, 1945
Parviturbo sp. P. Beach drift. Common.

"TURBINIDAE

Arene H. « A. Apams, 1857
Arene tricarinata (STEARNS, 1872). Range: southeast

U. S. to the West Indies and eastern Central America.
P, t. Uncommon. Common in beach drift.

Astraea Ropinc, 1798

Astraea caelata (GMELIN, 1791). Range: southeast Flor-
ida, the West Indies and the Caribbean. P, r. Shells only.
Fiuck, 1905, found it abundant in Nicaragua.

Astraea tuber (LINNAEuS, 1758). Range: southeast Flor-
ida, the West Indies and the Caribbean. P, r. One dead
specimen.

PHASIANELLIDAE
Tricolia Risso, 1826

Tricolia affinis cruenta Ropertson, 1958. Range: West-
ern Gulf of Mexico and the Caribbean to Brazil. P, t.
Common; abundant in beach drift.

NERITIDAE

Nerita Linnaeus, 1758

Nerita peloronta LiINNAEUS, 1758. Range: eastern Flor-
ida, Bermuda and the West Indies to Trinidad, and also
eastern Central America. P, r, 1. Found only during
March. Uncommon on large rocks.

Nerita versicolor GMELIN, 1791. Range: Florida to Texas,
Bermuda and the Caribbean to Brazil. P, r, 2. Abundant
on wet rocks in shaded areas.

Nerita fulgurans GMEuIN, 1791. Range: eastern coast of
Florida, Bermuda, the Caribbean and Brazil. P, r, 2.
Common under rocks in sheltered places. RUSSELL, 1941,
stated that this species is found in waters of lower salinity
but this writer found it commonly distributed along the
rocky shores of the Limon peninsula.

Page 12

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Vol. 11; No. 1

Puperita Gray, 1857

Puperita tristis (Orpicny, 1842). Range: Antilles and
eastern Central America. P, t. Abundant only in lagoons
on stones, dead coral, and debris.

Neritina Lamarck, 1816

Neritina virginea (LinNAEuS, 1758). Range: Florida to
Texas, Bermuda and the Caribbean to Brazil. P,m. Found
only near mouth of fresh water creek. Few live speci-
mens taken. Common in beach drift.

Neritina meleagris LAMARCK, 1822. Range: British Hon-
duras to Columbia and the Antilles to Brazil. P, t.
Found on rocks and debris with Puperita tristis. Common.

Neritina piratica Russe1, 1940. Range: Central Amer-
ica, the Antilles, the north coast of South America to
Brazil. P, m. Common in freshwater creek on submerged
logs and leaves. Intergrades in color pattern ranged from
typical N. piratica markings to N. recliata and N. zebra
types, but Russett (1941) claimed there is no inter-
grading of species. FLuck (1905) found this species to
be common in the lagoons and channels peculiar to the
mosquito coast of Nicaragua.

Smaragdia Issei, 1869

Smaragdia viridis viridemaris Maurey, 1917. Range:
southeast Florida, Bermuda, Veracruz to Yucatan, Mex-
ico, and the Caribbean. P, t. Common.

LITTORINIDAE

Lirrorina Férussac, 1822

Littorina ziczac (GMELIN, 1791). Range: southern Flor-
ida to Texas, Bermuda and the Caribbean, the Pacific
side of Panama near the canal and South America to
Uruguay. P, r, 1, 2. Abundant in splash zone.

Littorina lineolata Orzicny, 1842. Range: southern Flor-
ida to Texas and the Caribbean. P, r, 1, 2. ABport, 1964,
has pointed out the close resemblance between this species
and L. ziczac. Littorina lineolata was usually found in
wetter areas; it is smaller and of a deeper color. Abun-
dant in intertidal zone.

Littorina nebulosa (LAMARcK, 1822). Range: the Gulf
of Mexico and the Caribbean. P, r, 1. Found on rocks,
trees, and debris. Common.

Nodilittorina von Martens, 1897
Nodilittorina tuberculata (MENkE, 1828). Range: south-
ern Florida, Bermuda and the Caribbean. P, r, 1. Com-
mon in holes and pitted surfaces on rocks.

Tectarius VALENCIENNES, 1833

Tectarius muricatus (LINNAEUS, 1758). Range: lower

Florida Keys, Bermuda and the Caribbean. P, r, 1.
Common on rocks and vegetation.

TRUNCATELLIDAE

Truncatella Risso, 1826

Truncatella pulchella bilabiata Preirrer, 1839. Range:
southeast U.S. and the Caribbean. P, r, 1. Common in
rotting vegetation under rocks and debris.

RISSOIDAE

Rissoina Orpicny, 1840

Rissoina chesneli (MicHaup, 1830). Range: southeast
U.S. and the West Indies. P, beach drift. Common.

Rissoina decussata (Montacu, 1803). Range: North
Carolina to the Lesser Antilles and Central America. P,
beach drift. Uncommon.

Rissoina aberrans (C. B. ApAMs, 1850). Range: Gulf of
Mexico and the Caribbean. P, beach drift. Uncommon.

Rissoina toroensis (Otsson & McGinty, 1958). Range:
Central America. P, beach drift. Uncommon.

Rissoina cancellata Putuprt, 1847. Range: southeast
Florida and the West Indies. P, beach drift. Uncommon.

Rissoina sp. P, beach drift. Common.

Rissoina sp. P, beach drift. Common.

Zebina H. & A. Apams, 1855

Zebina browniana (OrsicNy, 1842). Range: Carolinas
to the West Indies. P, t. Under stones and debris in
Thalassia beds; beach drift. Abundant.

Alvania Risso, 1826

Alvania auberiana (Orpicny, 1842). Range: West In-
dies. P, beach drift. Uncommon.

Alvania chiriquiensis Otsson & McGinty, 1958. Range:
Central America. P, beach drift. Rare.

VITRINELLIDAE
Vitrinella C. B. Apams, 1852

Vitrinella elegans Otsson & McGinty, 1958. Range:
Central America. P, beach drift. Rare.

Vitrinella heliocoidea C. B. ApAMs, 1850. Range: south-
cast U.S., the West Indies, and the east coast of Central
America. P, beach drift. Uncommon.

Cyclostremiscus Pitspry & McGinty,
1945

Cyclostremiscus jeannae Pitspry & McGinty, 1945.
Range: Central America. P, beach drift. Uncommon.

Cyclostremiscus schrammu (FiscHER, 1857). Range:

Volatt- Nort

Guadeloupe and Central America. P, beach drift. Un-
common.

Cyclostremiscus ornatus Otsson & McGinty, 1958.
Range: Florida and the Caribbean. P, beach drift. Un-
common.

Teinostoma H.&A.Apams, 1854

Teinostoma megastoma (C.B.Apams, 1850). Range:
Carolinas to the Caribbean. P, beach drift. Common.

Parviturboides Pirspry & McGinty,
1950

Parviturboides interruptus (C.B. ApaMs, 1858). Range:
South Carolina to Gulf of Mexico and the Caribbean.
P, beach drift. Common.

Solariorbis ConrapD, 1865

Solariorbis blakei REHDER, 1944. (?) Range: South Car-
olina to the Gulf of Mexico and the Caribbean. P,
beach drift. Rare.

Solariorbis infracarinata Gasp, 1881. Range: North Car-
olina to the Gulf of Mexico and the Caribbean. P, beach
drift. Uncommon.

Solariorbis sp. P, beach drift. Rare.
Vitrinorbis Pitspry & Oxsson, 1952

Vitrinorbis elegans Otsson & McGinty, 1958. Range:
Central America. P, beach drift. Rare.

Circulus JEFFREYS, 1865

Circulus semisculptus (Otsson & McGinty, 1958).
Range: southern Florida and Central America. P, beach
drift. Uncommon.

Macromphalina CossmMann, 1888
Macromphalina sp. P, beach drift. Rare.

ARCHITECTONICIDAE
Heliacus Orpicny, 1842

Heliacus bisulcatus Orpicny, 1845. Range: southeast U.
S. and the Caribbean. P, beach drift. Uncommon.

Heliacus infundibuliformis (GmEuIN, 1791). (?) Range:
Caribbean. P, beach drift. Rare.

VERMETIDAE

Petaloconchus H.C. Lea, 1843

Petaloconchus irregularis (OrBIGNY, 1842). Range:
southern Florida and the Caribbean. P, r, 2. Cemented
on rocks. Common.

Stephopoma Mo6rcu, 1860
Stephopoma myrakeenae Otsson & McGinty, 1958.

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Page 13

Range: Central America. P, beach drift. Uncommon.

CAECIDAE

Caecum FLEminc, 1813

Caecum pulchellum Stimpson, 1851. Range: eastern U.
S. and the Caribbean. P, beach drift. Common.

Caecum jucundum Foun, 1867. Range: Caribbean. P,
beach drift. Abundant.

Caecum ryssotitum Foun, 1867. Range: Florida and the
Caribbean to Brazil. P, beach drift. Abundant.

Caecum clava Foun, 1867. Range: Caribbean. P, beach
drift. Rare.

Caecum cycloferum Foun, 1867. Range: Central Amer-
ica. P, beach drift. Rare.

Caecum sp. P, beach drift. Rare.

PLANAXIDAE

Planaxis LAMARCK, 1822

Planaxis lineatus (pA Costa, 1778). Range: lower Flori-
da Keys and the Caribbean. P, r, 2. Abundant beneath
rocks.

Planaxis nucleus (BrucutERE, 1789). Range: southeast
Florida and the Caribbean. P, r, 2. Abundant on rocks.

MopuLIDAE

Modulus Gray, 1842
Modulus modulus (LinnaEus, 1758). Range: Florida
to Texas, Bermuda, and the Caribbean to Brazil. P, t.
Screened in Thalassia beds. Uncommon.
Modulus carchedonius (LAMaRcK, 1822). (?) Range:

Greater Antilles to the north coast of South America and
Central America. P, beach drift. Rare.

CERITHIDAE

Cerithium BrucuikreE, 1789
Cerithium variabile C. B. Abas, 1845. Range: southern
Florida to Texas and the Caribbean. P, t, r, 2. Abundant
in July; uncommon in March.
Cerithium eburneum BruculkRE, 1792. Range: south-
east Florida, the Bahamas and the Greater Antilles, to
Central America. P, r, 5; t. Abundant on rocks; uncom-
mon in Thalassia area.
Cerithium auricoma SCHWENGEL, 1940. Range: Florida
and the Caribbean. P, beach. Uncommon.

Bitttum Gray, 1847

Bittium varium (PFEIFFER, 1840). Range: Maryland to
Florida, Texas, Mexico and the Caribbean. P, t. Under

Page 14

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Vol. 11; No. 1

rocks and debris in Thalassia beds. Uncommon; common
in beach drift.
Alaba H. & A. ApAMs, 1853

Alaba incerta (Orxicny, 1842). Range: Bermuda, Ba-
hamas, southeast Florida and the Caribbean. P, t.
Screened in deeper portions of Thalassia beds. Also found
in beach drift. Uncommon.

Cerithiopsis ForBes & Haney, 1851

Cerithiopsis green (C.B. ApAMs, 1839). Range: Cape
Cod to Florida and the Caribbean. P, t. Common alive
under stones in Thalassia beds. Abundant in beach drift.
Cerithiopsis regulosum (C. B. ApaMs, 1850). Range:
Caribbean. P, beach drift. Uncommon.

Cerithiopsis latum (C. B. ApAms, 1850). Range: Greater
Antilles and Central America. P, beach drift. Uncommon.

Cerithiopsis pupa DALL & Stimpson, 1901. Range:
Greater Antilles and Central America. P, beach drift.
Rare.

Cerithiopsis emersoni (C. B. ADAMS, 1838). Range: Mas-
sachusetts to the Caribbean. P, r, 5. Rare.
Cerithiopsis bicolor (C. B. ApAMs, 1845). P, r, 5.
Screened in green algae on rocks. Rare.

Seila A. ApAMs, 1861

Seila adamsi (H.C. Lea, 1845). Range: Massachusetts
to Florida, Texas and the Caribbean. P, t. Common under
stones and also in beach drift.

‘TRIPHORIDAE

Triphora BLAINvILLE, 1828

Triphora turristhomae Horten, 1802. Range: Carib-
bean. P, t. Also in beach drift. Uncommon.

Triphora nigrocincta (C.B.Apams, 1839). Range:
Massachusetts to Florida, Texas and the Caribbean. P,
t. Uncommon.

Triphora ornata DesHAyES, 1832. Range: Florida and
the Caribbean. P, beach drift. Rare.

Triphora pulchellum C.B. Apams, 1850. Range: Car-
ibbean. P, beach drift. Uncommon.

Triphora sp. P, beach drift. Uncommon.

EPITONODAE
Epitonium Roopine, 1798

Epitonium krebsi (Mércu, 1874). Range: southern Flor-
ida to Lesser Antilles and Central America. P, beach
drift. Uncommon.

Epitonium albidum (Orpicny, 1842). Range: south
Florida to Argentina. P, t. Rare.

Epitonium lamellosum (LamMarck, 1822). Range: south-
ern Florida and the Caribbean. Also Europe. P, beach
drift. Uncommon.

Epitonium candeanum (Orzicny, 1842). Range: south-
ern Florida to the Barbados and Central America. P,
uncommon in beach drift.

EULIMIDAE

Balcis Leacu, 1847

Balcis intermedia (CANTRAINE, 1835). Range: New Jer-
sey to the Caribbean. Also Europe. P, t. Uncommon.

Balcis conoidea Kurtz & Stimpson, 1851. Range: Flori-
da, the Gulf of Mexico and the Caribbean. P, uncommon
in beach drift.

Balcis sp. P, beach drift. Uncommon.
Balcis sp. P, beach drift. Uncommon.
Eulima Risso, 1826

Eulima auricincta Appott, 1958. Range: southern U.S.
to Greater Antilles and Central America. P, uncommon
in beach drift.

STILIFERIDAE
Stilifer Bropertp, 1832

Stilifer subulatus BRopERIP & SowerBy, 1832. Range:
Caribbean. P, rare in beach drift.

Athleenia Bartscu, 1946
Athleenia burryi Bartscu, 1946. Range: southern Flori-

da and eastern Costa Rica. P, rare in beach drift.

HIpPonicIDAE

Cheilea MoverEr, 1793

Cheilea equestris (LinNAEuS, 1758). Range: southeast
Florida and the Caribbean. P, common in beach drift.

Hipponix DeFRANCE, 1819

Hipponix antiquatus (LinNAEuS, 1767). Range: south-
east Florida and the Caribbean. P, common in beach
drift.

Hipponix subrufus subrufus (LaMaRcK, 1822). (?) Range:
southeast Florida and the Caribbean. P, common in
beach drift.

FosSSARIDAE
Fossarus Putri, 1841

Fossarus orbignyi FiscHer, 1864. Range: Caribbean. P,
common in beach drift.

Fossarus sp. P, beach drift. Uncommon.

Wooler Nor

STROMBIDAE

Strombus LinNAEus, 1758

Strombus gigas LinNAEUS, 1758. Range: southeast Flor-
ida, Bermuda and the Caribbean. P. Only a few broken
pieces of this species were found.

Strombus pugilis LinnaEus, 1758. Range: southeast
Florida through the Caribbean to Brazil. P, m. This spe-
cies was found in deeper water in the center of the
lagoon on a bottom covered with sediment and a fine
algal mat. It was abundant in July. The specimens appear
to be typical S. pugilis and do not have the character-
istics of S. pugilis nicaraguensis as described by FLuck
in 1905.

Strombus raninus GMELIN, 1791. Range: southeast Flor-
ida and the Caribbean. P, t. Uncommon.
ERATOIDAE
Trivia BRopERIP, 1837
Trivia leucosphaera ScHILDER, 1931. Range: southeast U.
S. and the Caribbean. P, beach drift. Uncommon.
CyPRAEDAE
Cypraea LINNAEUS, 1758

Cypraea zebra LINNAEUS, 1758. Range: southeast Flori-
‘da and the Caribbean. P, r, 2. Found with egg masses in
March. Uncommon under large rocks.

Cypraea cinerea GMELIN, 1791. Range: southeast Flori-
da and the Caribbean. P, beach drift. Common.
NATICIDAE

Polinices MontrFort, 1810

Polinices lacteus (GutLpinc, 1834). Range: southeastern
U.S. and the Caribbean. P, beach drift. Uncommon.
Fiuck (1905) found it abundant in Nicaragua.

CASSIDIDAE

Cypraecassis STUTCHBURY, 1837

Cypraecassis testiculus (LinNaEus, 1758). Range: south-
east Florida, Bermuda and from South Mexico and the
Caribbean to Brazil. P. Uncommon.

CyMATIDAE

Charonia GIsTEL, 1848

Charonia variegata (LAMARCK, 1816). Range: southeast
Florida and the West Indies. P, r, 3, 4. Usually covered
with thick coralline algal growth. Common.

Cymatium Ropine, 1798
Cymatium pileare (LinnagEus, 1758). Range: southeast-

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Page 15

ern U.S. to the Caribbean. P, beach drift. Common.

BuURSIDAE

Bursa Ropinc, 1798

Bursa cubaniana (Orpicny, 1842). Range: southeast
Florida and the Caribbean. P, r, 2. Uncommon under
large rocks in March.

ToNNIDAE

Tonna BRuNNIcH, 1772

Tonna maculosa (Dittwyn, 1817). Range: southeast
Florida and the Caribbean to Brazil. P, beach fragments.
Uncommon.

MurIcIDAE

Drupa Rép1ne, 1798

Drupa nodulosa (C. B. ApaMs, 1845). Range: southern
Florida and the Caribbean. P, r, 2; t. Common under
rocks.

Risomurex Otsson & McGinty,
1958

Risomurex roseus (REEVE, 1856). Range: throughout the
Caribbean. P, r, 2. Uncommon under rocks.
Risomurex muricoides (C.B. Apams, 1845). Range:
Caribbean. P, beach drift. Uncommon.

Purpura BRUGUIERE, 1789

Purpura patula (LinNAEuS, 1758). Range: southeast
Florida and the Caribbean. P, r, 2. Found in depressions
on wave-dashed rocks. Lewis (1960) stated that this
species feeds on Acanthopleura granulata, which is abun-
dant in this zone. Common. Larger specimens eroded.

Thais Ropinc, 1798

Thais rustica (LaMaRcK, 1822). Range: southeast Flori-
da, Bermuda and the Caribbean. P, r, 2, 3. Common.

Thais deltoidea (LamMaRcK, 1822). Range: Florida, Ber-
muda and the Caribbean. P, r, 2, 3. Found on rocks
covered with Hypnea and Acanthophora. Common.

Aspella Moércu, 1877

Aspella paupercula (C.B.Apams, 1850). Range: Car-
ibbean. P, t. Uncommon under rocks and dead Porites
coral.

MacILIDAE
Coralliophila H.& A. Avams, 1853

Coralliophila caribaea Assott, 1958. Range: southeast
Florida and the Caribbean. P, t, r, 4. Uncommon.

Page 16

CoLUMBELLIDAE
Columbella Lamarck, 1799

Columbella mercatoria (LINNAEUS, 1758). Range:
southeast Florida and the Caribbean. P, t. Uncommon on
Thalassia blades.

Anachis H.« A. ApAMs, 1853

Anachis catenata (SowrrBy, 1844). Range: Bermuda
and the Caribbean. P, r, 3, 4. Found in colonies under
rocks and coral heads. Uncommon.

Anachis obesa (C. B. Apams, 1845). Range: Virginia to
Florida, the Gulf of Mexico and the Caribbean. P, r,
2, t. Common under stones.

Anachis pulchella (Sowersy, 1844). Range: Florida
Keys and the Caribbean. P, t. Rare.
Anachis sp. P, beach drift. Rare.

Nitidella Swainson, 1840
Nitidella ocellata (GMELIN, 1791). Range: lower Flori-
da Keys, Bermuda and the Caribbean. P, beach drift.
Uncommon.

Psarostola REHDER, 1943

Psarostola monilifera (SowERBY, 1894). Range: Florida
and the Caribbean. P, uncommon in beach drift.

BucciINnIDAE
Bailya M. Smitu, 1944

Bailya parva (C. B. Apas, 1850). Range: Bahamas and
the Caribbean. P, t, r, 4. Found under dead Porites coral
and in company with Aspella paupercula under coral
heads.

Bailya intricata (DALL, 1883). (?) Range: southern Flor-
ida and the Caribbean. P, t. Uncommon under stones
and rubble.

Antillophos Wooprinc, 1928
Antillophos candei (Orxicny, 1853). (?) Range: south-
east U.S. and the Caribbean. P, t. Uncommon under
dead Porites coral.

Engina Gray, 1839

Engina turbinella (KieNneER, 1836). Range: lower Florida
Keys and the Caribbean. P, t. Found under stones in
sheltered areas. Common.

Engina sp. P, beach drift. Rare.
Colubraria ScHUMACHER, 1817

Colubraria lanceolata MENKE, 1828. Range: southeast
U.S. and the Caribbean. P, beach drift. Uncommon.

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Vol. 11; No. 1

Pisania Brvona-BERNARDI, 1832
Pisania pusio (LINNAEUS, 1758). Range: southeast Flor-
ida and the Caribbean. P, r, 4. Uncommon.

Cantharus Roépinc, 1798

Cantharus auritulus (Linx, 1807). Range: southeast
Florida, south through the Caribbean to Brazil. P, beach
specimens. Rare.

NASSARIDAE

Nassarius DumériL, 1806
Nassarius vibex (Say, 1822). Range: Cape Cod to Flori-
da, the Gulf States and the Caribbean. P, t. Found on
Thalassia in deeper water.
Nassarius albus (Say, 1826). Range: southeastern U.S.
to the Caribbean. P, beach drift. Uncommon.

Nassarius nanus Usticke, 1959. Range: Caribbean. P, t.
Uncommon.

FASCIOLARUDAE
Leucozonia Gray, 1847

Leucozonia nassa (GMELIN, 1791). Range: Florida to
Texas and the Caribbean. P, r, 2, 3. Common under rocks
covered with coralline algae.

Leucozonia ocellata (GmeEtIN, 1791). Range: Florida
and the Caribbean. P, beach specimen.

XANCIDAE

Vasum Roopine, 1798

Vasum muricatum (Born, 1778). Range: southern Flori-
da and the Caribbean. P, t, r, 4. Common on rocks and
debris.

OLIVIDAE

Oliva Brucutére, 1789

Oliva reticularis LAMARCK, 1811. Range. southeast Flor-
ida, the Bahamas and the Caribbean. P; B. Beach frag-
ments. Rare.

Agaronia Gray, 1839

Agaronia testacea (LaMarcK, 1811). Range: Central
America and in the Panamic Province from the Gulf of
California to Peru. P, t. FLuck (1905) found this spe-
cies to be abundant in Nicaragua and observed the living
animals. Orsson (1958) also listed it from Panama.
Tryon (1883) found it to be indistinguishable from A.
hiatula (Gmeutn, 1791) from West Africa and added
that both species exhibited a wide variation in both color
and form. Uncommon, dead.

Vol. 11; No. 1

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Page 17

Olivella Swainson, 1831

Olivella minuta Linx, 1897. Range: Caribbean. P, beach
drift. Uncommon. This may be the subspecies O. minuta
marmosa since Otsson & McGinty, 1958, list it as
occurring nearby in Panama.

MITRIDAE

Mitra Lamarck, 1799

Mitra nodulosa (GMELIN, 1791). Range: southeast U.S.
and the Caribbean. P, r, 4. Uncommon under coral heads.

Mitra barbadensis (GMELIN, 1791). Range: from south-
east Florida to the Caribbean. P, uncommon.

Mitra olssoni McGinty, 1955. (?) Range: southeastern
Florida and Central America. P, beach drift. Rare.

Pusia Swainson, 1840
Pusia gemmata (Sowersy, 1874). Range: Florida and
the Caribbean. P, r, 5. Common under rocks and on
algae.

MARGINELLIDAE

Marginella Lamarck, 1799
Marginella denticulata Conrav, 1830. Range: southeast
U.S. to the Caribbean. P, beach drift. Common.

Bullata JoussEAuME, 1875

Bullata ovuliformis (Orpicny, 1842). Range: the south-
east U.S. and the Caribbean. P, uncommon in beach
drift.

Persicula SCHUMACHER, 1817

Persicula adamsiana weberi Otsson & McGinty, 1958.
Range: Central American coast. P, rare in beach drift.

Persicula lavalleana (Orxicny, 1842). Range: southern

Florida and the Caribbean. P, uncommon in beach drift.
Hyalina ScHUMACHER, 1817

Hyalina avena KieneEr, 1834. Range: southeast U.S. and

the Caribbean. P, uncommon in beach drift.

Hyalina tenuilabra Tomuin, 1917. Range: Florida to the
Caribbean. P, uncommon in beach drift.

CoNnIDAE

Conus LinnarEus, 1758

Conus regius GMELIN, 1791. Range: Florida to southern
Mexico, the Bahamas and the Caribbean south to Brazil.
P, r, 4. No living specimens were taken.

Conus mus Hwass, 1792. Range: southern Florida to the
Caribbean. Burcu (1960) records this species as occur-
ring in the eastern Pacific at Panama Bay. P, r, 2, 3.

Common on exposed rocky shelves covered with Hypnea
and Acanthophora.

TURRIDAE

Crassispira Swainson, 1840
Crassispira fuscescens (Reeve, 1843). Range: Florida
Keys to the Caribbean. P, r, 5. Common in shady, pro-
tected areas.

Leptadrillia Woonrinec, 1928
Leptadrillia splendida Bartscu, 1934. Range: Carib-
bean. P, beach drift. Uncommon.

Mangelia Risso, 1826
Mangelia fusca (C.B. Apams, 1845). Range: Caribbe-
an. P, t. Common under rocks, on Thalassia leaves, and
in beach drift.

Pyrgocythara Wooprinec, 1928

Pyrgocythara coxi Farco, 1953. (?) Range: Florida, the
greater Antilles and Central America. P, beach drift. Un-
common.

ONCHIDIDAE

Onchidella Gray, 1850
Onchidella sp. P, r, 1, 2. Abundant on damp rocks.

ACTEONIDAE

Acteon Montrort, 1810
Acteon finlayi McGinty, 1955. Range: southeast Florida
and Cuba to Central America. P, beach drift. Rare.

BuLLIDAE
Bulla Linnaeus, 1758

Bulla striata Brucutére, 1792. Range: west coast of
Florida to Texas and the Caribbean. P. Common as
beach specimens.

ATYIDAE

Atys Montrort, 1810

Atys caribaea (Orpicny, 1841). (?) Range: from
southeast Florida to the Caribbean. P, r, 5. Found on
green algae and rocks in shady, protected areas. Common.

RETUSIDAE
Retusa Brown, 1827

Retusa candet Orpicny, 1841. Range: Caribbean. P,
uncommon in beach drift.

Retusa oxytatus Born, 1885. Range: southeast U.S. and
the Caribbean. P, uncommon in beach drift.

Page 18

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Vol. 11; No. 1

ACTEOCINIDAE

Acteocina Gray, 1847
Acteocina inconspicua Otsson & McGinty, 1958.
Range: Central America. P, uncommon in beach drift.

APLYSIDAE
Aplysia LinnaEus, 1767

Aplysia brasiliana RaNnc, 1828. Range: Atlantic Ocean
from New Jersey to St. Helena. P, r, 2, 3. Found in inter-
tidal and rock-shelf zones. Common on rocks covered
with Hypnea and Acanthophora.

Aplysia sp. P, t. Animals difficult to detect due to light
green color and darker green reticulations. Foot white,
lightly mottled with purple. Uncommon.

Aplysia dactylomela Rane (?) 1828. Range: world-wide
in warm seas. P, r, 2. Intertidal zone. Common on rocks
covered with red algae.

PyRAMIDELLIDAE

Cingulina A. Apams, 1860

Cingulina babylonia (C. B. Apams, 1845). Range: Ber-
muda and the Caribbean. P, beach drift. Common.
Miralda A. Avams, 1863

Miralda havanensis Pitspry & Acuayo, 1933. Range:
southern Florida to the Caribbean. P, beach drift. Com-
mon.

Miralda abbott: Otsson & McGinty, 1958. Range:
Central America. P, beach drift. Uncommon.

Odostomia FLEMING, 1817
Odostomia sp. P, beach drift. Common.

Odostomia sp. P, beach drift. Uncommon.
Oxy NOEIDAE

Oxynoe RAFINESQUE, 1819

Oxynoe antillarum Morcu, 1863. Range: southeast Flor-
ida to the Caribbean. P, t. Animals brilliant green and
quite active. Common on Caulerpa.

Lobiger Kroun, 1847
Lobiger souverbu FiscuHer, 1856. Range: southeast Flor-
ida, the Caribbean to Brazil. P, t. Uncommon on Cauler-
pa with Oxynoe.
ELLOBODAE

Pedipes MituuFexp, 1818
Pedipes mirabilis MiHLFELD, 1818. Range: Florida to

the Caribbean. P, r, 1. Common under damp rocks and
rotting vegetation in colonies.
Tralia Gray, 1840

Traha ovula (Brucutire, 1789). Range: Bermuda,
Florida, West Indies and the Caribbean coast of Central
America. P, r, 1. Common under damp rocks and rotting
seaweed.

Melampus Montrort, 1810

Melampus coffeus (Linnaeus, 1758). Range: Florida to
the West Indies and Caribbean coast of Central America.
P, r, 1. Abundant under rocks and debris, feeding on
rotting vegetation.

PELECYPODA

ARCIDAE

Arca Linnagus, 1758
Arca imbricata Brucut&zrE, 1789. Range: North Caro-
lina to the Caribbean. P, beach drift. Common.
Barbatia Gray, 1847
Barbatia domingensis (LaMarcK, 1819). Range: south-
east U.S., the Lesser Antilles and Central America. P, r,
2. Common in large groups under coral and rocks.
Arcopsis VON KoENEN, 1885

Arcopsis adamsi (E.A.SmituH, 1888). Range: North
Carolina, the Caribbean to Brazil. P, r, 2, t. Abundant
under rocks, in large groups.

Anadara DesHayeEs, 1830

Anadara ovalis (BrucutiRrE, 1789). Range: Cape Cod
to the Gulf States and the Caribbean. P, beach speci-
mens. Uncommon.

IsoGNOMONIDAE

Isognomon LicHTFooT, 1786

Isognomon alatus (GMELIN, 1791). Range: southern
Florida to the Caribbean. P, r, 2. Found in large clusters
attached in sheltered crevices on rocks. Common.

Isognomon radiatus ANTON, 1839. Range: southeast
Florida to the Caribbean. P, r, 2. Attached in fissures on
rocks and usually closer to shore than J. alatus. Common.

PTERMDAE

Pteria Scorou, 1777
Pteria colymbus Répinc, 1798. Range: southeast U.S. to

Vol. 11; No. 1

THE VELIGER

Page 19

the Caribbean. P, r, 2. Uncommon. Attached to rocks in
sheltered areas. Found only in March.

SPONDYLIDAE

Spondylus LinnaEus, 1758
Spondylus americanus HERMANN, 1781. Range: Florida
and the Caribbean. P, beach specimens.

LIMIDAE

Lima Brucuiére, 1797
Lima scabra (Born, 1778). Range: southeast Florida
and the Caribbean. P, beach specimens.
CRASSATELLIDAE
Crassinella Guppy, 1874

Crassinella martinicensis (OrpicNy, 1842). Range: Car-
ibbean. P, beach drift. Common.
Crassinella guadalupensis (Orpicny, 1842). Range: Car-
ibbean. P, beach drift. Common.

CoNDYLOCARDIDAE

Condylocardia BERNARD, 1896
Condylocardia bernardi (Dat, 1903). Range: Caribbe-
an. P, beach drift. Common.
DREISSENIDAE
Mytilopsis Conrap, 1857
Mytilopsis domingensis (REcLUz, 1852). Range: Car-
ibbean. P, m. Found near fresh water creek attached to
stones. Common.
DIPLODONTIDAE
Diplodonta Bronn, 1831

Diplodonta notata Dati & Simpson, 1901. Range: Flor-
ida to the Greater Antilles and Central America. P, beach
drift. Uncommon.

LucINnIDAE

Phacoides Gray, 1847

Phacoides pectinatus (GMELIN, 1791). Range: southeast
U.S., Texas and the Caribbean. P, t. Common in sand
under Thalassia.

Anodontia Linx, 1807

Anodontia alba Link, 1807. Range: southeast U.S. and
the Caribbean. P, t. Uncommon. Anodontia edentuloides
is the Panamic analogue.

Codakia Scorou, 1777
Codakia orbicularis (Linnagus, 1758). Range: Florida

to Texas and the Caribbean. P, t. Common in sand under
Thalassia beds. Codakia distinguenda is a closely related
species in the Pacific.

Codakia orbiculata (Montacu, 1808). Range: south-
east U.S. and the Caribbean. P, t. Uncommon, dead
valves.

Codakia costata Orpicny, 1842. Range: southeast U. S.
and the Caribbean. P, uncommon in beach drift.

CHAMIDAE

Chama Linnagus, 1758

Chama macerophylla GmMe.tn, 1791. Range: southeast
U.S. and the Caribbean. P, r, 2, 3. Common attached to
rocks in protected places.

Chama congregata Conrab, 1833. Range: southeast U.
S. and the Caribbean. P, t. Common as dead valves in
beach drift.

VENERIDAE

Chione MUuH.Fep, 1811

Chione cancellata (LinNAEus, 1767). Range: southeast
U.S. to Texas and the Caribbean. P, t. Only immature
specimens taken in sand under Thalassia. Rare.

Tivela Link, 1807

Twela mactroides (Born, 1778). Range: Caribbean

south to Brazil. P; B. Common as single valves on beaches.

The Panamic analogue is T: byronensis (Gray, 1838).
Pitar Romer, 1857

Pitar dione (LinNaEus, 1758). Range: Texas and the
Caribbean. B, common as beach specimens. Pitar lupana-
ria (Lesson, 1830) is a closely related species in the
Pacific.

TELLINIDAE

Tellina Linnarus, 1758

Tellina alternata Say, 1822. Range: southeast U.S., the
Gulf of Mexico and the Caribbean. B, on the beach with
soft parts. Shell completely pink. Uncommon.

Tellina lineata Turton, 1819. (?) Range: Florida and
the Caribbean. P, t. Uncommon.
Tellina sp. P, beach drift.

Arcopagia Brown, 1827

Arcopagia fausta (PULTENEY, 1799). Range: southeast
U.S. and the Caribbean. P, t.

Strigila Turton, 1822

Strigilla carnaria (LINNAEUS, 1758). Range: southeast
U.S. and the Caribbean. B, beach specimens. Abundant.

Page 20

Vol. 11; No. 1

THE VELIGER

SEMELIDAE

Semele ScHuUMACHER, 1817

Semele purpurascens (GMELIN, 1791). Range: southeast
U.S. and the Caribbean. P, beach specimens. Uncommon.

Semele nuculoides (Conran, 1841). Range: southeast U.
S. and the Caribbean. P, beach specimens. Uncommon.

DONACIDAE

Donax Linnaeus, 1758
Donax denticulatus LinNAEus, 1758. Range: Caribbean.
B. Some specimens unusually large. Common, burrowing
at the edge of the surf.

Iphigenia SCHUMACHER, 1817

Iphigenia brasiliensis (LAMARCK, 1818). Range: south-
ern Florida and the Caribbean. P, beach specimens. Un-
common. FLtuck (1905) found it plentiful in Nicaragua.
The Panamic analogue is J. altior (SoweRBy, 1833).

MactTrRIDAE

Mactra Linnaeus, 1767
Mactra alata SpENGLER, 1802. Range: Caribbean to
Brazil. Also occurs in the Pacific. P; B, beach specimens.
Abundant. This species is eaten by the natives.

Mulinia Gray, 1837
Mulinia cleryana Orsicny, 1846. Range: Caribbean to
Brazil. B, beach specimens. Common.

CorRBULIDAE

Corbula Brucuikre, 1792

Corbula sp. P, beach drift. Uncommon.
PHOLADIDAE

Martesia BLAINvILLE, 1824

Martesia striata (LinNAEvS, 1767). Range: west Florida
to Texas and the Caribbean. Also occurs in the Pacific.
P, in driftwood.

Pholas Linnaeus, 1758

Pholas campechiensis GMELIN, 1791. Range: North Caro-
lina to Brazil, Caribbean, Senegal to Liberia. B, beach
specimens. Common. This species is the analogue of P
chiloensis Mouina, 1782 of the Pacific.

PERIPLOMATIDAE

Periploma SCHUMACHER, 1817

Periploma inaequivalvis ScHuMACHER, 1817. Range:
Caribbean. B, beach specimens. Uncommon.

AMPHINEURA

CrYPTOPLACIDEA

Acanthochitona Gray, 1821

Acanthochitona rhodeus Pitspry, 1893. Range: Carib-
bean. P, r, 2. Uncommon under rocks and in dark places.

Acanthochitona sp. P, r, 2. Rare under rocks.

ISCHNOCHITONIDAE

Ischnochiton Gray, 1847

Ischnochiton pectinatus (SowerBy, 1849). Range: Car-
ibbean. P, r, 2. Common under rocks and in dark places.

Ischnochiton sp. P, r, 2. Rare under rocks.
Callistochiton Dau, 1882.
Callistochiton sp. P, r, 2. Rare under rocks.

CHITONIDAE

Chiton Linnaeus, 1758

Chiton tuberculatus LinNaEus, 1758. Range: Caribbe-
an. P, r, 2. Common in groups under wet rocks. Glides
quickly away from light when disturbed.

Chiton marmoratus GMELIN, 1791. Range: Caribbean.
P, r, 2. Uncommon. This species is quite active and shows
a negative phototropic reaction.

Acanthopleura Guitpine, 1829

Acanthopleura granulata (GmeE.in, 1791). Range:
southern Florida and the Caribbean. P, r, 1, 2. The
most abundant chiton and seen everywhere in groups.

SCAPHOPODA

SIPHONODENTALIIDAE
Cadulus Puiippi, 1844
Cadulus sp. P, beach drift.
DENTALIDAE
Dentalium Linnagus, 1758
Dentalium sp. P, beach drift.
Dentalium sp. P, beach drift.

CEPHALOPODA

SPIRULIDAE

Spirula Lamarck, 1799

Spirula spirula (LinNAEuS, 1758). Range: worldwide.
B, beach specimens.

Vol. 11; No. 1

THE VELIGER

Page 21

OcTOPODIDAE
Octopus Lamarck, 1798

Octopus vulgaris Lamarck, 1798. Range: Connecticut
to Florida, the Caribbean and Europe. P, t, r, 2, 3, 4.
Common.

LITERATURE CITED

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John-

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Golden Press, New York.

440; 1 fig.
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BEQUAERT, JOSEPH CHARLES
1942. Cerithidea and Batillaria in the western Atlantic.
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1943. The genus Littorina in the western Atlantic.
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Boss, KENNETH JAY
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John-

127 - 142
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Buscu, KATHERINE JEANNETTE
1899. _ Descriptions of new species of Turbonilla of the western

Atlantic fauna.
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Carr, ARCHIE
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CLeNcH, WILLIAM JAMES
1942. The genus Conus in the western Atlantic.
sonia 1 (6): 1-40; 15 plts.
1947, The genera Purpura and Thais in the western Atlantic.
Johnsonia 2 (23): 61-91; plts. 32-40
1964. The genera Pedipes and Laemodonta in the western
Atlantic. Johnsonia 4 (42): 117-127; plts. 76-79
Ciencu, WitiiaM James & Ropert Tucker ABBorT
1941. The genus Strombus in the western Atlantic.
sonia 1 (1): 1-15
1942. The genera Tectarius and Echininus in the western
Atlantic. Johnsonia 1 (4): 1-4
1943. The genera Cypraecassis, Morum, Sconsia, and Dolium
in the Western Atlantic. Johnsonia 1 (9): 8 pp.; 4 plts.
1943. The genera Gaza and Livona in the western Atlantic.
Johnsonia 1 (12): 1-9
CLeNcH, WILLIAM James « L. C. Smiru
1944. The family Cardiidae in the western Atlantic.
Johnsonia 1 (13): 1-32
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1956. The genus Truncatella in the western Atlantic.
Johnsonia 2 (25): 149 - 164; plts. 65 - 73
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Johnsonia 2 (30): 249 - 288; plts. 108 - 130
1956. The genera Epitonium (part I1), Depressiscala, Cylind-
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Johnsonia 2 (31): 289 - 356; plts. 131-177
1957. The family Cymatiidae in the western Atlantic.
Johnsonia 3 (36): 189-244 plts. 110-135
Coomans, H. E.
1958. A survey of the littoral gastropoda of the Netherlands
Antilles and other Caribbean islands. Stud. Fauna Cura-
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Dai, WiLtt1aMmM HEALEY
1886. Reports on the results of dredging, in the Gulf of
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Mollusca, pt. 1, Brachiopoda and Pelecypoda. Bull. Mus.
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Alfred A. Knopf, Inc. New

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John-

Page 22

THE VELIGER

Vol. 11; No. 1

FarFANTE, ISABEL PEREZ
1943. The genera Fissurella, Lucapina, and Lucapinella in the
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(7 August 1943)
1943. The genus Diodora in the western Atlantic. John-
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FiscHer, Paut
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Trochidae (Fin). 8. Journ. de Conchyliol. 6: 284 - 288
Fiucx, WILLIAM HENRY

1900: Shell collecting on the mosquito coast. Nautilus
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1901. Correspondence [from Nicaragua] Nautilus 15 (4):
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1905. Shell collecting on the mosquito coast. Nautilus
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19 (7): 78-80; 1906 - 20(1): 1-4
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Garcia-Cusas, A.
1963. Systematica y distribucion de los micromoluscos reci-
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Inst. Geol. Mex. 67 (4): 1-55
Ho ie, Paut Aucust & CLARENCE FE. DINEEN
1959. Studies on the genus Melampus (Pulmonata).
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Jaume, Micuet L.
1946. Moluscos marinos litorales del Cabo Catoche, Yucatan,
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1958. Sea shells of tropical West America; marine mollusks
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Ketso, DonaLp P.
1965. A contribution to the ecology of a tropical estuary.
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Kress, H. J.
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1960. The fauna of rocky shores of Barbados, West Indies.
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McGinty, THomas LaDUE
1955. | New marine mollusks from Florida.
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1941. The oysters of the western Atlantic.
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1964. The family Vitrinellidae in South Florida and the
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1922. The Miocene of northern Costa Rica.
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1963. | Zoogeography and ecology of some macro-invertebrates,
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1946. The type specimens of C.BApam’s Jamaican species
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Vol. 11; No. 1

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1941. The Recent mollusks of the family Neritidae of the

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1935. Historical geology of the Antillean — Caribbean region,
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1962. Three Caribbean atolls: Turmeffe Islands, Lighthouse
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1954. The family Pholadidae in the Western Atlantic and
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1959. A check list of the marine shells of St. Croix, U. S. Virgin
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Page 23

WarMKE, GERMAINE L. & RoBERT TUCKER ABBOTT

1961. Caribbean seashells; a guide to the marine mollusks of
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1926. Notes on marine mollusks from the Yucatan Peninsula,
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(5 March 1962)

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1928. | Miocene mollusks from Bowden, Jamaica: Part II:
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3 text figs. (28 November 1928)
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1957. | Geology and paleontology of the Canal Zone and ad-
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Woopwarp, S. P.

1875. A manual of the Mollusca. Lockwood & Co.,
London; 486 pp.; 272 figs.; 24 plts.; 1 map

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Vol. 11; No. 1

“Ptychodon’’ misoolensis ADAM & VAN BENTHEM JUTTING, 1939

a New Guinea Strobilopsid Land Snail

and Review of the Genus Enteroplax

BY

ALAN SOLEM

Field Museum of Natural History, Chicago, Illinois

(1 Text figure; 1 Map)

SINCE PUBLISHING A REVIEW of Pacific Island land snail
distributions (SoLeM, 1959a), I have written several pa-
pers (SoLEM, 1959b, 1964a, 1964b) concerning the
family position of puzzling genera whose accepted fam-
ily assignment produced geographically discordant data.
During preparation of major monographic reviews on
the Pacific Island, Australian, Melanesian and New Zea-
land endodontid land snails, many additional changes in
family position have been discovered. The reclassification
of Ptychodon misoolensis ADAM & VAN BENTHEM JUT-
TING, 1939, as Enteroplax misoolensis is presented sepa-
rately because of the great zoogeographic interest in-
herent in this change.

Basic data on the Strobilopsidae have been summarized
by Pitspry (1927 - 1931, pp. 1 - 63 and 1948, pp. 848 to
868). Subsequent descriptions by Ho « LEonarp (1961)
of fossil Strobilops from the Great Plains and of the
Bermuda S$. (Discostrobilops) pilsbryi and the West
Mexican S. (D.) sinaloa by Morrison (1953) do not
alter the basic patterns.

The family is common in the Eocene to Pliocene
fossil beds of Western Europe, but is absent from the
European Pleistocene and Recent faunas. Several species
live in the United States and Canada south of about 52°
N Latitude and generally east of the 100™ meridian.
Mid-Pliocene fossils are known from the High Plains re-
gions of Kansas and Oklahoma (Taytor, 1960). Rec-
ords are sparse for Northern Mexico. Strobilops aenea
mexicana Pitspry (1927, pp. 32-33) is known from
near Monterrey, Nuevo Leon and Necaxa, Puebla (also
see Pirssry, 1953, p. 165); S. stnaloa Morrison, 1953,
probably from Sinaloa; S. hubbardi (A.D. Brown, 1861)
from San Luis Potosi, Mexico (Pitssry, 1927, p. 49);
S. strebeli (Pretrrer, 1862) and S. veracruzensis Pits-

BRY, 1927 (pp. 33-37) from Vera Cruz; and S. hannai
Pitssry, 1931, from Socorro Island, West Mexico. Stro-
bilops strebeli guatemalensis H1iNKLEy, 1927 and S.
salvini (TRIsTAN, 1863) are known from Guatemala. There
are three peripheral South American records — S.
hellert (Dati, 1900) from the Galapagos; S. brasiliana
Frep Baker, 1913 from Para, Brazil; and S. morsei
(Dati, 1885) from Puerto Cabello, Venezuela. Strobi-
lops piratica Pitssry, 1930, from Old Providence Island,
S. wenziana Pitspry, 1930, from Grand Cayman and
forms of S. hubbard: (A.D. Brown) from Bermuda,
Cuba and Jamaica comprise the West Indian records.
Data on the above are summarized by Pirspry (1927
to 1931).

Two Asian groups complete the known taxa. The
section Eostrobilops Pitspry, 1927, consists of 4 species
— Strobilops nipponica Pitssry, 1927 from Yonezawa,
Uzen, Japan; S. hirasei Prrssry, 1927 from Cheju, Quel-
part Island, Korea; S$. coreana Pitspry, 1927, from
Pyong-Yang, North Korea; and §. diodontina HEUupDE,
1885 from Tchen-k’eou, China (probably is Ch’éng-k’ou,
northeast Ssuch’uan Province = Szechwan, at about
108° 47’ long., 31° 56’ lat.).

Enteroplax Gupr, 1899, previously was thought con-
fined to the Philippine Islands. There are 5 named taxa
representing 4 species, with available data on the pre-
viously included species summarized by Pirspry (1927
to 1931, pp. 50-56). Strobilops quadrast (MOoELLEN-
porFF, 1893) from Northern Luzon and S. polyptychia
(MoELLENDoRFF, 1887) from Cebu Island are easily
recognizable as distinct species. Strobilops trochospira
(MoELLENDoRFF, 1887) from Mt. Licos, Cebu Island
is quite similar to E. misoolensis. FausTINO (1930, p. 116)
listed these as “Plectopylis’ in the “Helicidae” together

Vol. 11; No. 1

with the endodontid Stenopylis coarctata (MOELLEN-
porFF, 1894). GupeE (1899, p. 149) separated Enteroplax
as a section, but indicated it probably was of generic
level separation. A review of this taxon is presented
below.

Enteroplax Gupr, 1899

Low conic to discoidal shells with prominent radial ribs
above threaded periphery and in umbilicus, sculpture
absent on body whorl below periphery. Edge of parietal
callus strongly elevated, upper parietal lamella fusing
with it anteriorly. Parietal lamellae 2, extending posteri-
orly about 4 whorl, upper high and blade-like, smooth
or serrate on expanded upper edge, 2*” much lower and
usually slightly recessed, rarely with a short and deeply
recessed interparietal lamella, sometimes a callus con-
necting the inner ends of the parietals. No columellar
lamella. Palatal wall with 3 to 10 short to long lamellae
recessed about $ to 4 whorl behind aperture, connected
posteriorly by a transverse callus. Anatomy unknown.
Type Species: Plectopylis quadrasi MoELLENDORFF,
1893, by OD.

Originally considered as related to the corillid genus
Plectopylis, their strobilopsid nature was recognized by
Pitsspry, 1908. References to the early literature are con-
tained in Pirssry (1927 - 1931, pp. 50-56). They are
not repeated here in the list of references, although in-
cluded in the synonymies. While Pirspry (1948, pp.
848 - 868) continued to treat the family Strobilopsidae
as monotypic on the generic level, ZitcH (1959, p. 178)
raised Enteroplax to generic level. I concur with this
decision, since there is a greater morphologic gap be-
tween the shell of Enteroplax and those classified as
subgenera or sections of Strobilops, than between any of
the latter. The elevation of the parietal callus and devel-
opment of a threaded edge to the periphery are not
extraordinary changes, but their presence in Enteroplax
and absence from all other groups of Strobilops, s.1. is
sufficient for generic recognition.

The 3 Philippine Island species were previously re-
ported from Luzon, Siquijor, Bohol and Cebu Islands.
A new record from Mindanao is added below. ‘The Misool
Island Ptychodon misoolensis ADAM & VAN BENTHEM
Juttinc, 1939, proved to be a fourth species. Specimens
of Enteroplax quadrasi and E. polyptychia were avail-
able at Field Museum of Natural History (hereafter
FMNH). Through the kindness of Dr. Adolf Zilch,
Natur Museum Senckenberg, Frankfurt am Main
(hereafter SMF), it was possible to examine specimens
of E. trochospira. Study of the Misool species was possible
through the cooperation of Mrs. W.S.S. van Benthem

THE VELIGER

Page 25

Jutting (Zoologisch Museum, Amsterdam, ZMA) and
Dr. W. Adam (Institut Royal des Sciences Naturelles de
Belgique, IRB). This work was supported by National
Science Foundation grants G-16419, GB-3384 and GB-
6779. For preparation of the illustrations I am indebted
to Miss Margaret Moran. Mrs. Sandra Rendleman, Mrs.
Lynda Hanke and Mrs. Rita Mecko assisted in various
aspects of this study.

Measurements of the specimens are summarized in
Table 1 and enable separation of the entities. All mea-
surements were made with an ocular micrometer to the
nearest 0.035 mm. It soon became obvious that the char-
acters used to separate species — number of palatal
lamellae, presence or absence of an interparietal lamella,
presence or absence of a transverse palatal callus, and
degree of serration on the parietals -— were highly
variable and, in certain cases, that species definitions were
based on aberrant conditions. Only 26 adult examples
were available. A few statements concerning similarities
of populations are possible, but considerably more ma-
terial will be required before the exact relationships of
Enteroplax quadrasi, E. trochospira and E. misoolensis
are clarified. The latter species apparently lacks serra-
tions on the parietal lamellae, but the two former have
the same apertural dentition and are separable only on
size and overlapping shape differences that may be
bridged when more populations are sampled.

Outline figures of the Philippine Island taxa, drawn
with the aid of a camera lucida on a Wild M-5 micro-
scope, are presented in Figure 1 to indicate the rather
subtle shape differences separating Enteroplax quadrasi
and E. trochospira. All indications of radial sculpture
have been omitted from the drawings, since there is no
interspecific variation in sculptural characters and addi-
tion of this feature would have been very time-consuming.

Enteroplax quadrasi (MoELLENDOoRFF, 1893)
(Figures 1b, 1c)

Plectopylis quadrast MoELLENDORFF, 1893, Nachr. deut. Ma-
lak. Gesell. 25 (11-12): 172-173 — Siamsiam, north-
ern Luzon, Philippine Islds.; Gupre, 1897, Science Gossip
4 (39): 71; figs. 54a - 54e; MoELLENDorFF, 1898, Abhdl.
naturf. Ges. Gorlitz 22: 122; Faustino, 1930, Philippine
Journ. Sci. 42 (1): 116

Plectopylis quadrasi subsp. boholensis MoELLENDORFF, 1898,
Abhdl. naturf. Ges. Gorlitz 22: 123 — Bohol, Philippine
Islands (nude name)

Helix (Plectopylis) quadrast (MoELLENDORFF), HIpDALco,
1891, Mem. Real. Acad. Cienc. Madrid 14: 167 —
Buguey, Sitio Siam-Siam en Claveria Sitios Dimacapac
y Cabayo en Palanen, Camino de Ambubuc, Prov. Caga-
yan, Luzon (not the cited figs. on plt. 156, figs. 9, 10)

Page 26 THE VELIGER Vol. 11; No. 1
Table 1
Local Variation in Enteroplax
Number
of
Specimens Ribs Height Diameter H/D Ratio Whorls D/U Ratio
Enteroplax
quadrasi
Luzon
FMNH 48337-8 87.344.19 2.13+0.036 3.58 0.068 0.596 + 0.0038 3- 3.66 =0.085
(82-97) (2.04—2.20) (3.42-3.75) (0.588-0.606) (5$-6) (3.45-3.86)
Bohol
FMNH 48341-2 88.5 = 2.04 2.16£0.044 3.66 £0.058 0.590 + 0.0062 64 - 3.35 +0.097
(86-91) (2.07-2.20) (3.55-3.75) (0.583-0.598) (53-64) (3.17-3.50)
Mindanao
FMNH 54943 107 2.40 3.68 0.652 64 4.31
trochospira
Cebu
SMF 9287/6 8 96.8 £1.57 2.32 £0.030 4.140.026 0.561 £0.0070 S§+ 3.63 + 0.050
SMF 118092/2 (90-101) (2.20-2.43) (4.05-4.24) (0.535-0.597) (55-52 (3.41-3.88)
misoolensis
Misool
ZMA, IRB 6 79.8+£8.29 2.63 0.055 4.180.066 0.629 +0.0144 64 4.22 + 0.068
(67-104) (2.43-2.80) (3.88-4.34) (0.573-0.664 ) (5$-63) (4.00-4.45)
polyptychia
Cebu
FMNH 48339-40 4 115 1.48 +0.036 4.03£0.028 0.368+0.0109 53+ 2.83+0.031
(1.38-1.55) (3.98-4.08) (0.339-0.388) (54-54) (2.75-2.88)

Plectopylis trochospira var. boholensis Gupr, 1898, Science
Gossip 4 (46): 285, fig. 74 - Bohol, Philippine Islds.

Strobilops (Enteroplax) quadrasi (MOELLENDORFF), WENZ,
1916, Nachr. deutsch. Malak. Gesell. 48 (4): 189; Pits-
BRY, 1931, Man. Conch. (2) 28: 55-56; plt. 11, figs. 11-14

Strobilops (Enteroplax) quadrasi var. brunnescens “MoE L-
LENDORFF, WENzZ, 1916, Nachr. deutsch. Malak. Ge-
sell. 48 (4): 189 -— nude name

Strobilops (Enteroplax) boholensis (GupE), Pirssry, 1931,
Man. Conch, (2) 28: 54-55; plt. 11, figs. 1-4, 7-10

Range: Luzon, Bohol and Mindanao, Philippine Islands.

Material: LUZON - Tayabas (1 specimen, FMNH
48388) ; Palanan (3 specimens, FMNH 48337). BOHOL
— Batuan (1 specimen, FMNH 48341); Vilar (2 speci-
mens, FMNH 48342). MINDANAO -— east slope of
Mt. McKinley, Davao at 7000 feet elevation (1 specimen,
FMNH 54943).

Remarks: Separation into two species seems to have
resulted from chance inspection of aberrant specimens.
The Luzon Island Enteroplax quadrasi was reported by
MoELLENporrFF as having only 3 palatals, but 3 of the 4
specimens seen by me had 4 palatals and only one had 3
palatals. Similarly, the form described as boholensis was

recorded to have an interparietal lamella and 5 palatals,
but all material that I saw lacked the interparietal and
had only 4 palatals. The size and shape of the Luzon and
Bohol specimens are identical except in whorl count and
a Statistically insignificant difference in D/U ratio (Table
1). Without more differences than revealed by study of
these specimens, separation of these populations cannot
be maintained. Reported differences in parietal serration
are a factor of wear after death and correlate with worn
versus unworn external sculpture.

A single shell collected on Mindanao (Figure 1c)
has been referred here provisionally. Although identical
in size to the other Enteroplax quadrasi samples, the shell
is obviously higher and with a narrower umbilicus. It has
typical parietals and 4 palatals. Whether this is another
atypical specimen, or is representative of populations
that are consistently higher and with narrower umbilici
cannot be known without further collections. In the latter
case, subspecific separation may be justifiable.

Enteroplax quadrasi differs from both E. trochospira
and E. misoolensis in size. Other differences are bridged
by variation between populations and probably have no
great significance.

Vol. 11; No. 1

THE VELIGER

Page 27

Enteroplax trochospira (MorLLENDoRFF, 1887)
(Figure 1a)

Plectopylis trochospira MoELLENDORFF, 1887, Jahrb. deutsch.
Malak. Gesell. 14 (3): 273-274; plt. 8, figs. 9a-9c —
Mt. Licos, Cebu, Philippine Islds.; MoELLENDorRFF, 1890,
Ber. Senckenb. naturf. Gesell. 1890: 221; MorELLENDoRFF,
1897, Abhdl. naturf. Ges. Gérlitz 22: 123; Gung, 1898,
Science Gossip 4 (46): 285; figs. 73a-73e; FaustTINo,
1930, Philippine Journ. Sci. 42 (1): 116

Helix (Plectopylis) trochospira (MoELLENDoRFF), H1patco,
1890, Mem. Real. Acad. Cienc. Madrid 14: 118, 167
(1891) — Sitio Cambaque en Vilar y Sierra Bullones,
Bohol, Philippine Islds. (not cited figs. on plt. 156)

Strobilops (Enteroplax) trochospira (MoELLENDORFF), WENZ,
1916, Nachr. deutsch. Malak. Gesell. 48 (4): 189; Pis-
BRY, 1931, Man. Conch. (2) 28: 52-53; plt. 11, figs. 5a-5c,
6a - 6e

Range: Cebu, Philippine Islands.

SS

Material: CEBU - Mt. Licos (8 specimens, SMF 9287
/6 paratypes, SMF 118092/2).

Remarks: Differences in spire shape and elevation (see
Figures 1a to 1c) between Enteroplax quadrasi and E.
trochospira are subtle. Extremes of both species show a
slight overlap. There is a distinct size difference (‘Table 1)
with the height and diameter not overlapping, although the
whorl counts are nearly identical. Examined specimens
of E. trochospira showed 4 (5 specimens) or 5 (1 speci-
men) palatals that were connected posteriorly by a trans-
verse callus and showed no structural differences from
the lamellae seen in FE. quadrasi. With some hesitation I
am accepting them as distinct species.

In size, Enteroplax trochospira and E. misoolensis are
identical, but the latter is distinctly more elevated, has a
narrower umbilicus, the parietal lamellae are not ser-
rated above, and averages 4 whorl more.

Figure 1

Shells of

a: Enteroplax trochospira (MoELLENDoRFF), Mt. Licos, Cebu, Philippine Islands; SMF 9287/6; Paratype.

b, c: Enteroplax

quadrasi (MoELLENDORFF), b: Palanan, Luzon, Philippine Islands. FMNH 48337; c: Mt. McKinley, Davao, Mindanao, Philip-
pine Islands; FMNH 54943; d: Enteroplax polyptychia (MoELLENDorFF), Mt. Licos, Cebu, Philippine Islands; FMNH 48339.
Probable paratype. Scale line equals 1 mm

Page 28

THE VELIGER

Vol. 11; No. 1

Enteroplax polyptychia (MoELLENDORFF, 1887)
(Figure 1d)

Plectopylis polyptychia MorLLENDoRFF, 1887, Jahrb. deut.
Malak. Gesell. 14 (3): 272-273; plt. 8, figs. 8a-8c —
Mt. Licos, Cebu, Philippine Islds.; MoELLENDORFF,
1890, Ber. Senckenb. naturf. Gesell. 1890: 221; Moert-
LENDoRFF, 1897, Abhdl. Naturf. Ges. Gorlitz 22: 122 —-
Siquijor, Philippine Islds.; Gupe, 1898, Science Gossip 4
(40): 102; figs. 55a-d; Faustino, 1930, Philippine Journ.
Sci. 42 (1): 116

Helix (Plectopylis) polyptychia (MoELLENDORFF), HiDALco,
1890, Mem. Real. Acad. Cienc. Madrid 14: 118, 167
(1891) -— not the cited figures on plt. 156

Strobilops (Enteroplax) polyptychia (MoELLENDORFF) ,
Wenz, 1916, Nachr. deut. Malak. Gesell. 48 (4): 188
to 189; Pmussry, 1931, Man. Conch. (2) 28: 52; plt. 12,
figs. 14-16

Range: Cebu and Siquijor, Philippine Islands.

Material: CEBU - Mt. Licos (3 specimens, FMNH
48339, probable paratypes) ; Barili (1 specimen, FMNH
48340).

Remarks: No Siquijor material was seen, but I have no
reason to question MoELLENDorFF’s identification. In its
greatly depressed shape and wider umbilicus, Enteroplax
polyptychia is immediately separable from the other spe-
cies. Although described as having about 9-10 minute
palatal lamellae, 2 of the 3 examples checked had the
typical 3 or 4 palatals found in other Enteroplax. Although
obviously distinctive, the depressed spire and wider um-
bilicus are linked characters caused by the change in
coiling pattern. Since the tooth pattern as originally
described is known to be aberrant, this species is now
shown to differ only in the single linked pair of changes.

Enteroplax misoolensis (ADAM & VAN BENTHEM JUTTING,

1939)

Ptychodon (Nesophila) misoolensis ADAM & VAN BENTHEM
Jurtinc, 1939, Bull. Mus. Roy. Hist. Nat. Belg. 15 (17) :
1-3; fig. 1 — forest 10 km north of Lilinta, Misool Is.,
in Asplenium; ApAM & LELoup, 1939, Mem. Mus. Roy.
Hist. Nat. Belg. (hors ser.) 2 (20): 16; plt. 2, fig. 9

Ptychodon misoolensis ADAM & VAN BENTHEM JUTTING, 1958,
Nova Guinea 9 (2): 327

Nesophila misoolensis (ADAM & VAN BENTHEM JUTTING), VAN
BENTHEM Jurtine, 1964, Nova Guinea 26: 11

Diagnosis: Shell large, diameter 3.89-4.35mm (mean
4.18mm) with 5% - 63 tightly coiled whorls. Apex and spire
strongly elevated, slightly rounded above, H/D ratio
0.573 - 0.664 (mean 0.629). Umbilicus broadly “U”-
shaped, barely decoiling, contained 4.00-4.45 times
(mean 4.22) in the diameter, with somewhat shouldered

margins. Upper surface of shell with irregularly protrac-
tive radial ribs, 67 - 104 (mean 79.8) on the last whorl,
whose interstices are less than twice their width. Major
ribs absent from below periphery of body whorl but re-
appearing within umbilicus. Periphery of body whorl
protrudingly and narrowly keeled. Aperture ovate, in-
clined about 25° from the shell axis. Parietal lamellae 2,
extending well beyond line of vision: upper high, blade-
like, attaining its greatest height at anterior end, becom-
ing bifurcated and forming raised edge to parietal callus;
lower less than 4 the height of the upper, stopping short
of parietal callus edge. Palatal wall with 4-5 short la-
mellae recessed */:; to 4 whorl within aperture.

The smooth parietal lamellae, moderately narrower
umbilicus, slightly more elevated spire and higher whorl
count separate Enteroplax misoolensis from the Philip-
pine E. trochospira. Enteroplax quadrasi is perhaps most
similar in shape, but differs in its smaller size and serrated
parietal lamellae.

Description: Shell large, with 64 tightly coiled whorls.
Apex and spire strongly and almost evenly elevated,
slightly rounded above, H/D ratio 0.621. Apical whorls
13, sculpture of vaguely radially oriented granulosities.
Remaining whorls with prominent, irregularly shaped,
protractive radial ribs, about 67 on the body whorl, which
are much less prominent near the suture and absent from
the body whorl below the narrow periphery, but reap-
pear within the umbilicus. Microsculpture of vaguely
reticulated radial and spiral riblets, often obscured by
changes in the major growth wrinkles. Umbilicus broadly
“U-shaped, regularly decoiling, contained 4.26 times in
the diameter. Sutures relatively shallow, whorls moder-
ately rounded above, somewhat flattened laterally below
periphery. Aperture ovate, lip strongly thickened and re-
flected, inclined about 25° from shell axis. Parietal wall
with heavy callus and 2 lamellae extending posteriorly
beyond line of vision: upper parietal high, bladelike,
sharply descending near anterior margin, then bifurcat-
ing to form a raised edge to parietal callus; lower parietal
much reduced in height, ridgelike, extending not quite
to edge of parietal callus. Columellar wall without lamel-
lae. Palatal wall with 4 lamellae visible from aperture, a
5™ seen through the shell, recessed */:s to + whorl within
aperture. Uppermost tooth thin, low, very short; 2°” much
higher, slightly expanded above and longer; 3°’ even
more prominent; 4™ slightly reduced in height and
length; 5™ trace apparently even shorter. Height of
holotype 2.70mm, diameter 4.35 mm.

Holotype: New Guinea: Misool Island. Collected Feb-
ruary 26, 1929 in forest 10 km north of Lilinta, on
Asplenium. Musée Royal d’Histoire Naturelle de Bel-
gique no. 9223.

Vol. 11; No. 1 _THE VELIGER Page 29

QUELPART®

WV £ostrobilops

* Enteroplax

v

* Chungking

Figure 2

Distribution of Enteroplax (gg) and Strobilops (Eostrobilops) (‘p)

Page 30

THE VELIGER

Vol: 1 Nomi

Paratypes: ZMA and IRB.

Range: Misool Island. Known from near Lilinta, Waima
and Fakal.

Material: MISOOL - 10km north of Lilinta (4 spe-
cimens, ZMA, IRB) ; Waima (3 specimens, ZMA) ; Fa-
kal (1 specimen, ZMA).

Remarks: Although identical to Enteroplax trochospira
in size, the absence of any serrations on the upper
parietal, distinctly narrower umbilicus, proportionately
higher spire, and presence of a half whorl more at the
same diameters separate E. misoolensis. Only 4 examples
had the aperture clear enough or the body whorl sufh-
ciently transparent to enable counting the palatal lamel-
lae — one had 5 palatals and 3 had 4 palatals.

ZOOGEOGRAPHY

Prior to the discovery that Enteroplax was present on
Misool Island, the distribution of Asian Strobilopsidae
appeared anomalous. Species of the section Eostrobilops
were known from China, Korea and Japan (see Map)
with the Philippine Islands containing an endemic sub-
genus, Enteroplax. Viewed from a North Pole perspec-
tive, as in Prrspry (1927 - 1931, p. 15, fig. 2), a deriva-
tion of the Philippine Enteroplax from the north, using
the deus ex machina of accidental transport on the
feathers of birds would seem logical and reasonable.
Extension of the known distribution to Misool Island is
highly significant in that it brings the probable derivation
of Enteroplax into the typical pattern of 1) movement
from southeast Asia into Indonesia; 2) reaching the
Philippines and Misool; and 3) followed by replacement
in the Indonesian area leaving the isolated populations
in Misool and the Philippines. Almost certainly popula-
tions will be found in New Guinea.

Specimens of Eostrobilops lack the raised parietal cal-
lus and the peripheral thread, plus having the parietal
lamellae with superior serrated nodes (see Pitspry,
1927 - 1931, pl. 10, fig. 14). Enteroplax and Eostrobilops
are not closely related to each other and their geographic
juxtaposition has no systematic significance.

LITERATURE CITED

Apam, WILLIAM & T. vAN BENTHEM JUTTING
1939. Une nouvelle espéce de Ptychodon de VArchipel In-
dien. Bull. Musée roy. d’hist. nat. Belg. 15 (17): 1-3;
fig. 1 (April 1939)
FaustTINo, LEopotpo A.
1930. Summary of Philippine land shells.
Sci. 42 (1): 85-198
Gupe, GrErarp K.
1898. | Armature of helicoid landshells.

Philipp. Journ.
(May 1930)

Sci. Gossip 4 (46):

284 - 285 (March 1898)
1899. Armature of helicoid landshells. Sci. Gossip 6 (66):
174-177 (November 1899)

Ho, Tonc-Yun « ARTHUR Byron LEONARD
1961. Two new strobilopsids from the Pleistocene of the high
plains. Nautilus 75 (2): 43-49; plt. 6; 5 figs.
(4 October 1961)
Morrison, JosEPH PauL ELDRED
1953. Two new American species of Strobilops.
67 (2): 53-55; plt. 6
Pirspry, HENry AuGUSTUS
1908. | Notes on the genus Strobilops.

Nautilus
(11 November 1953)

Nautilus 22 (8):

78 - 80 (11 December 1908)
1927 - 1931. Manual of conchology (2) 28 (1-2): 1-96; plts.
1-12

1948. Land Mollusca of North America (north of Mexico).
Acad. Nat. Sci. Philadelphia Monogr. 3, 2 (2): i-xlvii +
521-1113; figs. 282 - 585 (19 March 1948)

1953. Inland Mollusca of northern Mexico. II. Urocoptidae,
Pupillidae, Strobilopsidae, Valloniidae, and Cionellidae.
Proc. Acad. Nat. Sci. Philadelphia 105: 133 - 167; plts. 3-10; 3

figs. (9 December 1953)
SoLeM, ALAN
1959a. Systematics and zoogeography of the land and fresh-

water Mollusca of the New Hebrides.
43: 359 pp.; 34 plts.; 38 figs.; 18 tables (15 Oct. 1959)
1959b. On the family position of some Palau, New Guinea,
and Queensland land snails. Arch. f. Molluskenk. 88: 151
to 158; plts. 12-13; 2 figs. (21 December 1959)
1964a. Amimopina, an Australian enid land snail. The
Veliger 6 (3): 115 - 120; 4 text figs. (1 January 1964)
1964b Foxidonta, a Solomon Island trochomorphid land snail.
The Veliger 6 (3) : 120- 123; 5 text figs. (1 January 1964)
TayLor, DwicHtT WILLARD
1960. Late Cenozoic molluscan faunas from the high plains.
U.S. Geol. Surv. Prof. Pap. 337: 94 pp.; 4 plts.; 2 text figs.;
19 tables
ZiLcH, ADOLF
1959. Handbuch der Palaozoologie. 2 (1):
Berlin, Germany (Gebr. Borntraeger)

Fieldiana: Zoology

1-200; 701 figs.

Vol. 11; No. 1

THE VELIGER

Page 31

Observations on Aquarium Specimens of Oliva sayana RAVENEL

AXEL A. OLSSON

1906 Ferdinand Street, Coral Gables, Florida

AND

Bonoe

Mrs. L. E. CROVO

2915 S W 102” Avenue, Miami, Florida 33165

(Plate 2)

INTRODUCTION

Oliva sayana 1S THE COMMON, large Oliva in Florida
occurring along both coasts, and in some localities quite
plentifully although like Olivella nivea Gmeuin, 1791,
fast disappearing in others. It is an excellent aquarium
snail with interesting habits and will thrive for many
years with minimum attention. Essential is a fairly large
or spacious tank of sea water, provided with a deep sand
layer, the water kept fresh and well aerated and on
occasions changed when cleaning is required. Normally,
the snail lies buried in the sand out of sight, the siphonal
tube generally protruding above, straight as a stick, or
swinging back and forth as if to keep track of what may
be taking place above. At meal time, the snails will
emerge obediently and accept a great variety of animal
food, such as pieces of fish, shrimp, steak, and best of
all, other live mollusks such as Nassarius, Donax and
Laevicardium. Olives have long been known to be both
scavengers as well as active predators on other mollusks
but the actual manner of food capture remained imper-
fectly known. The following notes record some of our
observations along this line made over a period of several
years.

SNAIL WATCHING

In July 1967, our pastime of Snail Watching became
even more exciting when several of our aquarium in-
mates began to produce egg capsules in a seemingly
endless procession, a performance repeated at intervals
for about three months. Although the egg capsules of
Olivella and its veligers have been described in detail by
Marcus in Brazil, those of Oliva itself remained unre-

corded at far as we have been able to learn. We are
offering a few illustrations based on photographs; those
of the veligers are experimental and were taken under
rather high power. Since a live veliger is a very active
little creature, passing in and out of the limited field of
view and at different levels, instantaneous exposures were
taken and only partial focus secured.

FOOD HABITS

Several years ago the senior author observed Agaronia
preying on Olivella on a beach near Manta, Ecuador.
After capture of an Olivella, the body of the Agaronia
would swell up into a rounded ball suggesting that the
smaller animal had been directly swallowed. Gently
squeezing the body of the Agaronia, the Olivella would
pop out, and placed in a dish of sea water, would quickly
recover. As these observations were made in the field,
incidental to more pressing duties, it is possible that a
mistaken interpretation had been made as suggested by
Marcus (1959). That small mollusks, such as Ervilia,
Semelina, Caecum, as well as ostracods and foraminifera,
are habitually swallowed by several species of Olivella
(O. nivea (GMELIN, 1791), O. gracilis (BRoDERIP & Sow-
ERBY, 1829), etc.) is proved, since their shells are com-
monly found intact and empty in the digestive tract
on dissection. Logically, Oliva should be able to do
the same with even larger forms; but additional obser-
vations on this subject are required. Even if the food
pieces are large, the following procedure is normal. If
pieces of fish, shrimp, or steak be placed in the tank, the
Olives soon become aware of them and will begin to
emerge and start on a round of investigation. When the
food morsel is discovered, it is quickly seized and pushed

Page 32

back under the foot and infolded in its hinder section as
if tucked away in a pocket, which swells up into a
rounded ball, the food item hidden away so completely
that no part of it is visible externally. The Olive then
retreats below sand level, going down head-first. The
same procedure is used with live mollusks, such as Donax
and Laevicardium, to which latter Oliva sayana is espe-
cially partial. A L. vitellinum (Reeve, 1844), if placed in
the tank, seems to become aware of a lurking danger and
will flip about energetically, especially if pursued by an
Olive. Large Laevicardium, nearly the size of the Olive,
may be taken, infolded in the foot, and carried under-
ground. Clams with a rough surface, such as Chione can-
cellata (LinNAEus, 1767), are consistently rejected.

EGG CAPSULES ann VELIGERS

On July 13, 1967, the junior author noticed that one
large Oliva had emerged and seemed to be acting rather
peculiarly, the lobes of its mantle loose and flabby. At
the same time, the surface of the sand appeared as if
covered with small, crystal-clear balls, which like minia-
ture balloons swayed and shifted about with the least
agitation of the tank water. These balls proved to be
transparent egg capsules containing minute white eggs,
the capsules varying in shape from nearly spherical to
oblong and about 1.7mm in greatest length and about
1.5mm in diameter. Each capsule held from 20 to 50
eggs, the eggs at first quite clear, darkening to nearly
black as incubation set in. The skin of the capsule is
quite tough, flexible when touched with a sharp needle.
A group of capsules set apart in a dish of sea water in-
cubated in from 3 to 5 days, the hatched embryos devel-
oping rapidly into nearly full-sized veligers which could
be seen spinning around within the crowded space of
the capsule for a day or more. In about the seventh day,
the capsule would puncture at one end, the exit generally
so small that only a single veliger could escape at a
time, the others pushing on behind. From one capsule
closely watched, the average time for the exit of a
veliger was one minute and the entire capsule of 38

THE VELIGER

Volchlss Nos!

embryos to empty required a little over half an hour. In
several other capsules, the last few veligers, unable to
find the tiny exit, perished within. Once out, the veligers
were extremely active for about two days, the cilia on
their vela vibrating vigorously in a continuous wave-
like pattern, producing the visual impression of rapidly
rotating blades. Observed from above, the veligers looked
like small helicopters with rapidly spinning rotor blades,
or like a man on a bicycle, the shell hanging below. Oc-
casionally the velar lobes would fold inward, and the
embryo would sink to the bottom, sometimes to rest, or
the embryo would begin to spin around actively as if
to test, or to bore into, the substrate, for further meta-
morphosis, then to rise again for another tour about. At
first, the velar flaps were clear but as they grew a little
larger 4 brown spots appeared. As noticed by Marcus
(1959) for the veligers of Olivella verreauxii (Ducros,
1857), those of Oliva sayana RAvENEL, 1834 subsisted
entirely on the yolk sacs within; these at first were dark
and became gradually lighter in color. None of our
veligers were able to undergo further metamorphosis,
the substrate evidently being unfavorable.

SUMMARY

The egg capsules of Oliva sayana are free, hence subject
to dispersal by currents; they differ from those of Oli-
vella verreauxu, which are attached (Marcus, 1959).
Whether this distinction is generic remains for future
research to determine.

Our observations show that many details of molluscan
life may be obtained by patient snail watching of aquar-
ium specimens. Species of Oliva lend themselves ad-
mirably for this purpose through their ready adaptability
to aquarium life, and they possess many interesting
habits.

LITERATURE CITED

Marcus, ERNST
1959. Studies on Olividae.
Zool. 22: 99 - 188

Bol. Fac. Filos. Cienc. S. Paulo,

Explanation of Plate 2

Figure 1:
Figure 2:

Oliva scouting and sensing shrimp.
Oliva going down with shrimp infolded in the foot.

Figure 3: Oliva held in hand showing sole of foot, the propodium
down and the food package infolded in the hinder section of the

foot.

Figure 4: Egg capsule, punctured and partly emptied. One veliger
free outside, another part way out. ca. X 35.

Figures 5, 6:
operculum, etc.

Free veligers. ca. X 160. Note small foot, shell,

Tue VELIGER, Vol. 11, No. 1 [OLtson « Crovo] Plate 2

Figure 2

Figure 4

Te

Vol. 11; No. 1

THE VELIGER

Page 33

A Record of the Indo-Pacific Cone, Conus ebraeus, in Guatemala

BY

WILLIAM K. EMERSON

Department of Living Invertebrates
American Museum of Natural History

Seventy-ninth Street and Central Park West,

Conus ebraeus LINNAEUS, 1758, A WIDE-RANGING, Shallow
water, Indo-Pacific species, was first known to occur in
the eastern Pacific at Clipperton Island and in the Gala-
pagos Islands (Hertiein, 1937). In 1953, the late Ted
Dranga found this species living on the mainland of the
west coast of Central America. He reported (in litt.)
taking a living specimen ... “‘in a crevice on an
extensive area of hard rock wave bench covered with
short sea weed >” on the Pacific coast of Guana-
caste Province, Costa Rica (HERTLEIN & Emerson, 1953,
p. 351). This record was subsequently cited by KEEN,
1958, p. 480; Hanna, 1963, p. 61; and Emerson, 1967,
p. 89 (recorded erroneously as “Conus chaldeus R6-
DING”). Housrick (1968) reported collecting two ad-
ditional living specimens in the intertidal zone near
Playas del Coco, Guanacaste Province, Costa Rica, in
1965. According to Dr. Kenneth J. Boss (in litt.), one
of these specimens, cat. no. 256447, is deposited in the
Museum of Comparative Zoology, Harvard University
and it is a typical specimen of Conus ebraeus.

The American Museum of Natural History recently
received from Mrs. Jane Zager of West Orange, New
Jersey, a specimen of Conus ebraeus that was stated to
have been collected by G. Farris on flats exposed by a
minus tide at San José, Escuintla Department, Guate-
mala, in 1947. This specimen, cat. no. 114575, which
measures 24.5mm in length, is nearly identical with one
figured by HerTLEIN (1937, plt. 1, fig. 2) from Clipper-
ton Island.

These records indicate that this species may be expec-
ted to occur in suitable habitats along the coast of the
West Americas within the Panamic faunal province.
Collectors should make a special effort to look for this

New York, New York 10024

species and other Indo-Pacific faunal elements when col-
lecting in the coastal waters of this region. At the present
time most of the Indo-Pacific species occurring in the
tropical eastern Pacific are known only from the oceanic
islands, Clipperton, Revillagigedo, Cocos, and the Gala-
pagos, off the west American coast (EMERSON, 1967).

LITERATURE CITED

Emerson, WILLIAM KEITH
1967. Indo-Pacific faunal elements in the tropical eastern
Pacific, with special reference to the mollusks. Venus,
Japan. Journ. Malac. 25 (3/4): 85-93; 1 map (July 1967)

Hanna, G Datias
1963. | West American mollusks of the genus Conus; II.
Calif. Acad. Sci. Occ. Pap. 35: 1-103; plts. 1-11
(28 January 1963)
HERTLEIN, LEO GEORGE
1937. A note on some species of marine mollusks occurring in
both Polynesia and the western Americas. Proc. Amer.
Philos. Soc. 78 (2) : 303 - 312; 1 plt.; 1 map
(December 1937)

Hertvein, LEo Georce « WILLIAM K. EMERSON
1953. Mollusks from Clipperton Island (eastern Pacific) with
the description of a new species of gastropod. Trans. San
Diego Soc. Nat. Hist. 11 (13): 345 - 364; plts. 26, 27
(22 July 1953)
Hovusrick, JosepH R.

1968. New record of Conus ebraeus in Costa Rica.

Veliger 10 (3): 292.
Keen, A. Myra

1958. Sea shells of tropical West America; marine mollusks

from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)

The
(1 January 1968)

Page 34

THE VELIGER

Vol. 11; No. 1

A Cowrie Mutant from the Gulf of Thailand

BY

FRANZ ALFRED SCHILDER

University of Halle, German Democratic Republic

(1 Text figure)

MUTANTS ARE HEREDITARY VARIANTS which differ from
typical specimens by one or a few striking characters;
there are no, or only rare, intermediates. The mutants live
among the not-varied individuals of the same species
and do not show any environmental specialization. They
show a distinct center of frequency and become gradually
scarcer in populations living around this locality. True
mutants differ from morphes (ScHILDER, 1966, p. 185)
by their area of distribution being still very restricted, if
compared with that of the typical species.

In cowries such local mutants evidently are very rare, as
only three variants can be classified in this way: Ovatipsa
chinensis tortirostris (SowERBy, 1906) from South Africa
(see ScHILDER, 1966, p. 186), Lyncina carneola titan ScuiL-
DER & SCHILDER, 1962 from southern Kenya (established as
a distinct species), and Erronea errones azurea SCHILDER,

1968, from Broome.

Recently a fourth case has been disclosed by several
series of cowries presented to me chiefly by Mr. J. Orr
and Mr. Franz B. Steiner: Mauritia arabica LINNAEUS,
1758, which formerly was thought to be represented by
a single race in the Gulf of Thailand (ScuiLprEr, 1965, p.
26), has proved to consist of two well separable variants
along the east coast of the interior part of this Gulf.

The primary Mauritia arabica is mostly small, oblong to
ovate, and rather depressed; in shells from the northern
Gulf of Thailand the length usually (i. e. in 67% of
shells approaching the mean) varies from 43 to 55mm,
the usual breadth from 60 to 62% of length, and the
usual height from 48 to 51% of length.

These shells belong to the subspecies Mauritia arabica asi-
atica SCHILDER & ScHILDER, 1939 (ranging from Japan to
the Gulf of Thailand) if this “race” really can be separated
from the Malayan typical M. a. arabica by its lips being

more acuminate chiefly at the rear.

The mutant is larger and broader, as its usual length
varies from 53 to 66mm and its breadth from 64 to 69%
so that the means differ in a significant way; besides, the
outline of the base is rather deltoidal (instead of ellip-
tical) and the dorsum is humped, viz. relatively higher
(usually 52 to 55% of length, the mean being 54 instead
of 50%) and its top is displaced to the rear of the shell so
that the shells recall Trona stercoraria (LinNaEus, 1758)
in shape ( Figure 1). The brown dorsal longitudinal striae
are evidently far less interrupted by pale lacunae than it is
in the other races of Mauritia arabica.

This mutant should be called Mauritia arabica gibba
CoENn, 1949 (see ScuiLpER, 1964, p. 104) in spite of the
inaccurate type locality “China Sea”: for this region also
includes the Gulf of Thailand, and Corn’s indications of
habitat were mostly uncertain or even false.

The extremely broad and callous Mauritia arabica dilatata
Coen, 1949 (see loc. cit.) becomes a synonym of M. a. gibba
as it applies to an individual variant of which several speci-
mens have been collected in the populations of the more
frequent and less extreme M. a. gibba.

The center of distribution of Mauritia arabica gibba
evidently lies in the islands of the Ko Sichang group
(about 70km SE of Bangkok), especially in Ko Taitamun
(see map in Figure 1) ; here and in the Ko Khrok group
(opposite of Pattaya) only has M. a. gibba been collected
as yet, but no M. a. arabica. On the beaches of the
mainland, however, from Pattaya to east of Rayong both
M. a. arabica and M. a. gibba have been collected, e. g.
in Ban Pe (Bang Pae) and in Ban Klaeng Lang, where
the specimens of the species M. arabica include 75% and
68% respectively M. a. gibba. Farther east, i. e. at Laem
Sing and in the Ko Chang group no M. a. gibba have
been collected at all, as it is, according to Mr. J. Orr, the
case also in the whole west coast of the Gulf of Thailand
(Hua Hin, Chumporn, Songkla).

Vol. 11; No. 1

Bankok
°

Ko Sichang fe

g
Ko Khrok-*

Ko Khram? |

Gulf of Thailand

THE VELIGER

Page 35

Figure 1

Map of the northeastern Gulf of Thailand. — Right: Profile and
basal view of Mauritia arabica (dotted line) and of its mutant
gibba (solid line).

The largest shells of Mauritia arabica gibba, varying from
67 to 79mm, have been collected in a Japanese wreck sunk
in the Ko Sichang group during World War II: the dorsum
of all specimens is rich chestnut showing the usual dark
brown striae; this discoloration varying in intensity shell by
shell evidently has been caused by the rusty water in which
these animals lived; the base, however, is rather pale as usual.

Therefore Mauritia arabica gibba is restricted to the
east coast of the northernmost part of the Gulf of Thai-
land from the Ko Sichang group to Ban Klaeng Lang.
The center of the morphologically well separable mutant
is in the Ko Sichang group in the innermost part of the
Gulf of Thailand, from where it spread about 100km
to the southeast.

Note — The subspecific name gibba of Coen, 1949 is not
preoccupied by Cypraea gibba GMELIN, 1791, because CoEN
established the name for a members of the genus Mauritia.
Lumpers, using the single generic name Cypraea for all
cowries, will be obliged to establish a new name for the
secondary homonym: for the only synonym of gibba, called
Mauritia (Arabica) arabica dilatata by Corn, 1949 (see above)

would become also a secondary homonym preoccupied by
Cypraea spurca var. dilatata MonTEROSATO, 1897.

LITERATURE CITED

ScHILDER, FRANz ALFRED
1964. The cowries established by Cozn in 1949 (Mollusca:
Gastropoda) . The Veliger 7 (2): 103-107 (1 Oct.’64)
1965. The cowrie fauna of Thailand. The Veliger
8 (1): 23-28; 1 map (1 July 1965)

1966. Personal views on taxonomy. The Veliger 8 (3):
181-189; 1 text fig. (1 January 1966)

1968. An interesting mutation in cowries. Hawaiian
Shell News (in press)

SCHILDER, FRANZ ALFRED, & MARIA SCHILDER
1938 - 1939. Prodrome of a monograph of living Cypraeidae.
Proc. Malacol. Soc. London, 23 (3): 119-180; (1939) 23
(4) :-181 - 231; 1 text fig.; 9 maps.
1962. Zur Kenntnis der Cypraeidae: 5. Eine neue Riesenform
aus Ostafrika. Arch. Molluskenk. 91 (4/6): 207 - 212;
1 fig. (30 December 1962)

Page 36

THE VELIGER

Vol. 11; No. 1

The Rediscovery of Voluta (Lyria) grangert SOWERBY 3"°, 1900

CLIFTON STOKES WEAVER

Hawaiian Malacological Society, 2777 Kalakaua Avenue, Honolulu, Hawaii 96815

AND

JOHN ELEUTHERE puPONT

Delaware Museum of Natural History, Greenville, Delaware 19807

(Plate 3; 1 Map)

In JuLy oF 1967 we received from Mr. and Mrs. C. N.
Cate of Los Angeles an unidentified shell that appeared
to be a new volutid species in the genus Lyria Gray, 1847.
It had been trawled by pearling luggers during August,
1949 in 5 fathoms of water a few miles north-northwest
of Port Hedland, central Western Australia. The arrival
of a second specimen from the Cates in October of the
same year from the same locality seemed to confirm our
opinion. However, further research showed that in 1900
Sowersy 3” described and figured Voluta (Lyria) gran-
geri, in the Annals and Magazine of Natural History, ser.
7, vol. 5, p. 440, plt. 11, fig. 2, but he gave no locality data.
His figure and description closely resembled the shells in
question.

When photographs of the holotype of Voluta (Lyria)
grangeri arrived from the British Museum (Natural His-
tory) in London and we compared them with the Cates’
specimens, there remained little doubt that all 3 shells
were conspecific.

Since 1900 Lyria grangeri seems to have been missing
from molluscan literature. We therefore take pleasure in
reestablishing grangeri as a valid specics in the genus Ly-
ria and in designating a type locality for it.

Lyriinae Pitspry «& Oxtsson, 1954
1954. Lyriinae Pitspry & Oxsson, Bull. American Paleont.
35 (152): 285, 286
Type Genus: Lyria Gray, 1847.

Lyria Gray, 1847

1847, Lyria Gray, Proc. Zool. Soc. London 15: 141

Type Species: Voluta nucleus LAMarcK, 1811; Recent,
Australia, by OD.

(Lyria) s. s.

Lyria (Lyria) grangeri (SowERBY 3", 1900)
(Plate 3, Figures 1 to 6)

1900. Voluta (Lyria) grangeri SowErBy 3°, Ann. Mag. Nat.
Hist. (7) 5: 440, plt. 11, fig. 2 (no locality given).
Type: Holotype, British Museum (Natural History),
London, No. 1900.5.22.83.

Type Locality: Here designated a few miles north-
northwest of Port Hedland, central Western Austral-
ia (see Map).

Explanation of Plate 3

Ventral and Dorsal Aspects of Lyria (Lyria) grangeri (Sowrrsy 3", 1900)

Figures 1, 2: Holotype, ex British Museum (Natural History),
No. 1900.5.22.83; no locality data on original label; photo-
graphs courtesy BM (NH).

Figures 3, 4: Homeotype No. 1, cx C. N. Cate Coll.; trawled by
a pearling lugger several miles NNW of Port Hedland, central

Figures 5, 6:

Western Australia in 5 fathoms; photographs by Clifton S.
Weaver.

Homeotype No. 2, ex C. N. Cate Coll.; same locality
data as for the preceding specimen; photographs by Clifton
S. Weaver.

TuHeE VELIGER, Vol. 11, No. 1 [WEAVER & DUPONT] Plate 3

7.)

Figure 1 Figure 2

Figure 3 Figure 4

Figure 5

Vol. 11; No. 1 THE VELIGER

INDIAN OCEAN

DAMPIER
ARCHIPELAGO

BEZOUT

at" Port Hedland
ISLAND~.% / en

NICHOL
Det, BAY ,

e” Cossack

LONG
ISLAND

A: Range limits of

NORTH WEST CAPE /

as presently known.

PT. CLOATES

a ee

or CAPRICORN — — — ae

5 Carnarvon

| SHARK *
\ BAY

A

HOUTMAN
ABROLHOS IS.

ie Geraldton

CAPE BOSSUT &

of
By :

Lyria (Lyria) grangeri (Sowerby II, !900) |

| Broome

La Grange

20°

30°

Page 38 THE VELIGER Vol. 11; No. 1
Table 1
Measurements (in millimeters) and Collecting Data for Lyria (Lyria) grangenn (SowERBY 3*°, 1900)
Maximum Collecting
Height Diameter Aperture Depth in Growth
Specimen Length Locality Fathoms Collection Stage
Holotype 40.0 21.0 26.5 Unknown Unknown BM (NH) No. 1900.5.22.83 Adult
Homeotype No. 1 41.3 24.0 27.2 A few miles NNW of Port 5 Cc. N. Cate Adult
Hedland, central Western
Australia
Homeotype No. 2 42.8 24.6 30.0 A few miles NNW of Port 5 Cc. N. Cate Adult

Hedland, central Western

Australia

BM (NH) = British Museum (Natural History)

Range: Known only from type locality.
Habitat: In 5 fathoms. Substrate unknown.
Dimensions: See Table 1.

Shell Description: Shell of medium size for genus, sub-
globose, light in weight. Spire low, convex; apex blunt.
Protoconch small, turbinate, of 2 smooth whorls. Teleo-
conch of 5 swollen whorls indented at sutures. Sculpture
consists of low narrow axial ribs extending from suture
to suture, these ribs becoming broader and finally obso-
lete on adult body whorl, about 19 or 20 such ribs on
penultimate whorl of shell measuring 42.8 mm in length.
Aperture wide, expanding anteriorly; interior a yellowish-
cream color. Columella curved inward, with 3 anterior
plaits, followed by numerous fine lirae which disappear
about midpoint on columella. A small tooth-like projec-
tion appears at posterior end of columella below junction
of outer lip and body whorl. Outer lip thickened exter-
nally to form a parallel raised ridge about 7mm wide.
Siphonal notch wide, shallow; fasciole well defined. Base
color creamy-white overlaid with chestnut-brown blot-
ches forming 2 broad bands on body whorl, | at periphery
and 1 above anterior tip. Fine brown lines, about 2mm
apart, encircle body whorl and intrude just inside outer
lip.

Animal and Radula: Unknown.

Remarks: We have removed Lyria grangeri from Voluta
and placed it in the genus Lyria for obvious reasons. The
species has much in common with Lyria (Lyria) deliciosa
(Montrouzirr, 1859) including the small, smooth, tur-
binate protoconch, axially ribbed early whorls, and the
arrangement of columellar plaits and lirae. The single
tooth-like projection on the posterior end of the columella
also resembles that found in L. (L.) mitraeformis (La-
MARCK, 1811) and on several other species of Lyria s. s.

Although the distribution of Lyria (Lyria) mitrae-
formis extends as far west as Albany, south Western

Australia, no other Lyria, to our knowledge, inhabits
waters off the central or north Western Australian coasts.
We know of the existence of only 3 specimens of
Lyria (Lyria) grangeri: The holotype is in the British
Museum (Natural History), London, and 2 are in the
collection of Mr. and Mrs. Crawford N. Cate; the pres-
ent whereabouts of a possible fourth specimen, mentioned
by Sowersy 3"? in his original description, is unknown.

ACKNOWLEDGMENTS

We wish to thank our friends, Mr. and Mrs. C. N. Cate
for allowing us to see the 2 specimens which triggered
this investigation. We are also grateful to Dr. Norman
Tebble and the photographic staff at the British Museum
(Natural History), London, who kindly sent the type
photographs which appear in this paper.

LITERATURE CITED

Gray, JoHN Epwarp
1847. A list of the genera of Recent Mollusca, their synonyma
and types. Proc. Zool. Soc. London (for 1847) 17 [part 15]
(178): 129 - 219 (November 1847)
Lamarck, JEAN-BapTisTE PIERRE ANTOINE DE MONET DE
1811. De la détermination des espéces des mollusques testa-
cés. Ann. Mus. Nat. d’Hist. Nat. 17: 62-69 (April 1811)
Monrrouzigr, R. P.
1859. Descriptions d’espéces nouvelles de l’Archipel Calé-
donien. Journ. de Conchyl. (2) 7: 375 (June 1859)
Pirspry, HENry Aucustus « AxEL ADOLF OLSson
1954. Systems of the Volutidae.
35 (152): 271 - 306; plts. 25 - 28

SowERBY, GEORGE BRETTINGHAM (3rd of name)

Bull. American Paleont.
(7 September 1954)

1900. New species of mollusca of the genera Voluta, Conus,
Siphonalia, and Euthria. Ann. Mag. Nat. Hist. (7) 5:
440; plt. 11, fig. 2 (May 1900)

Vol. 11; No. 1

THE VELIGER

Page 39

A List of Types of the Family Volutidae

Held by the National Museum of Victoria

BRIAN J. SMITH

Curator of Invertebrates
National Museum of Victoria, Australia

BoTH PRIMARY AND SECONDARY TYPES of the family
Volutidae held by the National Museum of Victoria are
listed under the original name. The present status in each
case is included according to the Second Provisional Spe-
cies List of Living Volutidae, by C.S. WEAVER (1964).

Cymbiola randalli Stoxes, 1961.
Holotype.

Reference: Stokes, A. J., 1961. A new species of Volute
from Undine Reef, Queensland Cymbiola randalli sp. nov.
R. Soc. S. Aust. Malac. Sec. Publ. 16: 3, fig.

Type Locality: 3 miles from Undine Reef, North Queens-
land on sandy bottom. Collected by Mr. A. J. Randall.
Present Status: Synonym of Cymbiolacca wisemani
(Brazier, 1870)

National Museum of Victoria Reg. No. F 22758
Remarks: Shell only.

Livonia mamilla leucostoma Maysiom, 1951.
Holotype.

Reference: Maysiom, A. 1951. Deep-sea shells from New
South Wales. Aust. Zool., 11 (4): 283; plt. 26, fig. 3.
Type Locality: Below 80 fms. around Gabo Island.
Present Status: Synonym of Livonia mamilla (SowERBY,
1844).

National Museum of Victoria Reg. No. F 19280
Remarks: Shell only.

Voluta hamillei Crosse, 1869.
Holotype.

Reference: Crossr, H., 1869. Diagnose d’une espéce
nouvelle de Voluta. Journ. de Conch., Paris. 17: 278.
Crosse, H., 1870. Description d’espéces nouvelles.
Journ. de Conch., Paris. 18: 97 - 99; plt. 1, fig. 5 and plt.
2, fig. 4.
Type Locality: Japan. In the original description (1869)
the type locality was quoted as Solomon Islands. This
was corrected to Japan in the next year.

Present Status: Fulgoraria hamillei (Crossr, 1869).
Valid Species.

National Museum of Victoria Reg. No. F 26773.
Remarks: Shell only. Specimen purchased from Mr. R.
Damon in 1875 for 15 pounds.

Voluta roadnightae McCoy, 1881.
Holotype.

Reference: McCoy, F, 1881. Description of new Volute
from South Coast of Australia. Ann. Mag. Nat. Hist.
Fifth. Ser. 8: 88 - 89; plt. 7, figs. 1 and 2.

Type Locality: Southern Coast of Victoria. Type speci-
men found on the beach, Lake’s Entrance, Gippsland,
Victoria. Collected by Mrs. Roadnight.

Present Status: Livonia roadnightae (McCoy, 1881).
Valid species.

National Museum of Victoria Reg. No. F 662
Remarks: Shell only. The type specimen was first noticed
by the eminent botanist, Baron von Mueller when on a
visit to the Gippsland Lakes district where it was serving
as a prop to keep his bedroom window open at his hotel.

Voluta (Amoria) spenceriana GatuFF, 1908
Holotype.

Reference: Gaturr, J. H., 1908. Description of Voluta
(Amoria) spenceriana, sp. nov. from North Queensland.
Victorian Nat. 25 (5), fig.

Type Locality: North Queensland. Collected by J. F
Bailey.

Present Status: Amoria spenceriana (GaTurFF, 1908).
Valid species.

National Museum of Victoria Reg. No. F 455
Remarks: Shell only.

Voluta thatcheri McCoy, 1868.

Holotype.

Reference: McCoy, F, 1868. On a new Volute. Ann. Mag.
Nat. Hist. Fourth Ser., 1: 54; plt. 2, fig. 1.

Page 40

Type Locality: From the type description habitat un-
known. Dr. Cox (1872) states that the type specimen
was purchased by Mr. C. R. Thatcher in a pawnbroker’s
shop in Collins Street, Melbourne. Specimens have since
been taken alive from Bampton Reefs, north of New
Caledonia.

Present Status: Notovoluta thatcheri (McCoy, 1868).
Valid species.

National Museum of Victoria Reg. No. F 22764
Remarks: Shell only.

Aulica kellneri TREDALE, 1957.
Paratypes

Reference: IREDALE, T., 1957. Another Australian Volute.
Journ. Proc. R. Zool. Soc. N. S.W. 1955-1956 [1957]: 91
to 92; plt.

Type Locality: Eastern Arnhem Land, North Australia.
Locality on paratypes — Blythe River Gulf, Northern
Territory.

Present Status: Synonym of Aulica flavicans (GMELIN,

1791).

National Museum of Victoria Reg. No. F 18192

Remarks: 2 shells.

Cymbiolacca cracenta McMicwae1, 1963.
Paratype

Reference: McMicuaet, D.F, 1963. Descriptions of
two new species of the genus Cymbiolacca IREDALE (Gas-
tropoda: Volutidae). Journ. malac. Soc. Austral. 7: 33 - 41
Type Locality: Dredged in 17 - 20 fathoms 15 miles SE
of Cape Green (south of Townsville), Queensland. Col-
lector probably T. Nielsen.

Present Status: Cymbiolacca cracenta McMicuatk1, 1963.
Valid species.

National Museum of Victoria Reg. No. F 23822
Remarks: One shell.

Cymbiolacca complexa nielsent McMicHak1, 1959.
Paratype

Reference: McMicuac1, D. FE, 1959. Marine Mollusca of
Eastern Australia. 1. The genus C'ymbiolacca IREDALE
(Family Volutidae). Aust. Zool. 12 (4): 374 - 383; plt.
44,

Type Locality: 30 miles off Burnett Heads, Hervey Bay,
Queensland, 10 fathoms (locality of the N. M. V. paratype
given as off Bundaberg, Queensland).

Present Status: Cymbiolacca complexa nielsent McMt-
CHAEL, 1959. Valid subspecies.

National Museum of Victoria Reg. No. F 18259
Remarks: Specimen (shell and animal) in spirit; also a
radula slide from the same specimen with the same Reg.
No. This is the only type radula slide of this subspecies.

THE VELIGER

Vol. 11; No. 1

Cymbiolacca peristicta McMicHae1, 1963.
Paratypes

Reference: McMicwaet, D.F, 1963. Descriptions of
two new species of the genus Cymbiolacca IREDALE (Gas-
tropoda: Volutidae). Journ. malac. Soc. Aust. 7: 33 - 41;
plt. 6.

Type Locality: Big Sandy Cay, Swain Reefs; dredged 2
fathoms. Collector Mr. T: Nielsen.

Present Status: Cymbiolacca peristicta’ McMicHAEL,
1963. Valid species.

National Museum of Victoria Reg. No. F 22757
Remarks: Two shells.

Nannamoria parabola Garrarb, 1960.
Paratype

Reference: Garrarp, T: A., 1960. A new species of Nann-
amoria (Mollusca: Volutidae) from Southern Queens-
land. Journ. malac. Soc. Aust. 4: 2 - 13; plt. 1.

Type Locality: 125 fathoms off Moreton Island, Queens-
land.

Present Status: Nannamoria parabola GarRARD, 1960.
Valid species.

National Museum of Victoria Reg. No. F 21107
Remarks: One shell.

Pseudocymbiola provocationis McMicwHaet, 1961
Paratype

Reference: McMicuaet, D. F, 1961. New Species and
new records of marine Mollusca from Australia. Journ.
malac. Soc. Aust. 5: 51-57; pit. 4, figs. 9 - 10.
Type Locality: Off Port Kembla, N.S. W.
Present Status: Pseudocymbiola provocationis McMicu-
AEL, 1961. Valid species .
National Museum of Victoria Reg. No. F 23821
Remarks: Shell only.

Voluta (Amoria) gatliffi SowErBy, 1910.
Cotypes

Reference: Sowersy, G. B., 1910. Description of a new
Volute. Ann. Mag. Nat. Hist. Ser. 8, 6: 611.
Type Locality: Port Keats, Northern Territory, Australia.
Present Status: Synonym of Amoria damoni (Gray,
1864).
National Museum of Victoria Reg. No. F 620 & F 5066
Remarks: Shells only. — There may possibly be a third
cotype, Reg. No. F 626, but this cannot be found.

Voluta macgillivrayi Cox, 1873.

Paratype

Reference: Cox, J.C., 1873. Distribution of Australian
Volutes. Second edition.

Vol. 11; No. 1

Type Locality: Woodlark Island. New Guinea.
Present Status: Synonym of Aulicina norrisii (GRAY,
1838).

National Museum of Victoria Reg. No. F 5067
Remarks: Shell only.

Voluta mamilla Gray, 1844
Figured Specimen
Reference: Gaturr, J.H. « C.J. Gasrier, 1909, First
record of the animal of Voluta mamilla, Gray; with re-
marks thereon. Victorian Nat. 26 (8): 117-118; plts. 2
& 3, figs. 1-5.
Type Locality: 15 miles SW of Gabo Island, Victoria, in

THE VELIGER

Page 41

30 fathoms.

Present Status: Livonia mamilla (Gray, 1844). Valid
species.

National Museum of Victoria Reg. No. F 805
Remarks: Radula slide cannot be found. Shell only. Shell
and animal of this species all figured.

Also in the collections of the National Museum of
Victoria are photographs of a paratype of Mesericusa
stokesi Cotton, 1961.

Reference: Cotton, B. C., 1961. A new species of Volute,
Mesericusa stokesi sp. nov. from South Australia. R. Soc.
S. Aust. Malac. Sec. Publ. 16: 1; fig.

Page 42

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Vol. 11; No. 1

Mediargo, a New Tertiary Genus in the Family Cymatiidae

BY

JUDITH S. TERRY

Department of Geology, Stanford University, Stanford, California 94306

(Plate 4)

INTRODUCTION

In THE COURSE OF RESEARCH on Recent and fossil species
of the cymatiid genera Argobuccinum, Fusitriton, and
Priene, the writer observed (TERRY, 1968) a number
of Pliocene specimens that possess characters of all three
genera but could not be assigned to any one. Juveniles
resemble Recent species of Gyrineum Linx, 1807, and
late juveniles are easily confused with those of Fusttriton
oregonensis (REDFIELD, 1848). Young adults are similar
to Recent species of Argobuccinum HERRMANNSEN, 1846,
while gerontic forms such as the holotype of Gyrineum
lewisti Carson, 1926 and the mature specimen figured
herein are distinct from all other cymatiids. Specimens of
different growth stages have been compared, and 2 species
can be recognized: Mediargo mediocris (Dax, 1909) of
middle or late Miocene to Pliocene age, and M. mathew-
sonii (Gass, 1866), which ranges from the middle Oligo-
cene to middle Miocene. Only the former, the type
species, will be considered here.

Mediargo TERRY, gen. nov.

Type Species: Gyrineum mediocre DAL, 1909
Specimens referred to Mediargo by the writer have

been classified most commonly in Ranella, Bursa and
Gyrineum. Although characters, such as lateral varices
and anterior pillar folds, are common to all four, the
new genus is distinguished by a combination of these
and other morphologic features that place it closer to
Argobuccinum and Fusitriton.

Diagnostic generic characters that are seen in all but
the largest gerontic specimens include the following: two
nearly continuous lateral varices on each volution, high
spire, rounded whorls having tabulate shoulders, mod-
erate to long anterior canal, anal notch oriented at an
angle to the axis of coiling (as in Fusitriton, in contrast
to the apically directed notch in Argobuccinum), trans-
verse pillar folds over most of the columella, and a denti-
culate or plicate outer lip. Axial and spiral costae may
be marked or obsolete and are commonly present on
juvenile whorls but conspicuously lacking in the later
stages. Mediargo differs from Argobuccinum in its tabular
whorls and anal notch, from Fusitriton in its lateral
varices and plicate aperture, and from Gyrineum in its
outline, tabulate shoulders, ovate aperture, marked anal
notch, and large size.

The name Mediargo is feminine in gender. It combines
the root of Gyrineum mediocre, here designated as the
type species, and the prefix of Argobuccinum, the genus
to which it is morphologically most similar.

Explanation of Plate 4

Figure 1: Mediargo mediocris (Dat, 1909). Holotype U.S.N.
M. No. 153900, Coos Bay, Oregon. Pliocene. Holotype of Gyrine-
um mediocre Dati, 1909. 4.4cm high.

Figure 2: Mediargo mediocris (Datt), LACMIP No. 2155.96,
Palos Verdes Hills, California. “Pleistocene.” [reworked Pliocene.]
4.2cm high.

Figure 3: Mediargo mediocris (DaLL), Univ. Oregon F 2638,
Bandon, Oregon. Mio-Pliocene. 4.2cm high. Compare variation
in sculpture between specimens in Figures 2 and 3.

Figure 4: Mediargo mediocris (Dat), CAS No. 11689, from a
well in San Diego, California. Pliocene. 4.5 cm high.

Figure 5: Mediargo mediocris (DatL), Paratype, USNM No.
645876, Coos Bay, Oregon. Miocene or Pliocene. (Former number
was 153900.) 6cm high.

Figure 6: Mediargo mediocris (Datt), LSJU No. 138, Santa
Maria District, California. Pliocene. [Paratype of Gyrineum
lewisti GARSON, 1926.] 6.2 cm high.

Figure 7: Mediargo mediocris (DatL), UCMP No. 10105, Coos
County, Oregon. Pliocene. Note plicate aperture. 4.6cm high.
Figures 8, 9: Mediargo mediocris (Dat), LSJU No. 31625,
Kettleman Hills, Fresno County, California. Pliocene. Mature spe-
cimen. 11.3 cm high.

Tue VELIGER, Vol. 11, No. 1 [Terry] Plate 4

Figure 3

Figure 6 Figure 7

Figure 9

Vol. 11; No. 1

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Page 43

Mediargo mediocris (Dati, 1909)

(Plate 4, Figures 1 to 9)

Gyrineum mediocre Dati, 1909, U.S. Geol. Surv. Prof. Pa-

per 59: 54-55; plt. 7, fig. 6
Weaver, 1943, Univ. Wash. Publ. Geol. 5 (II) : 423 - 424;

(III): plt. 83, fig. 13

Gyrineum lewisii Carson, 1926, Bull. So. Calif: Acad. Sci.
25: 53 - 54; plt. 2, figs. 1, 2

Ranella (Priene) mediocris (DaLL): Grant & Gate, 1931,
Mem. San Diego Soc. Nat. Hist. 1: 736

Ranella (Priene) lewisti (CARSON): Grant & Gate, 1931,
Mem. San Diego Soc. Nat. Hist. 1: 736

“Gyrineum” mediocre lewisiti CARSON: Wooprinc & BraM-
LETTE, 1950, U.S. Geol. Surv. Prof: Paper 222: 73 - 74;
pit. 12, figs. 13, 15; plt. 13, figs. 23, 24, 26, 27

Type Information: The holotype, USNM 1539000, and
paratype, USNM 645876, were collected by Mr. B.H.
Camman from the vicinity of Fossil Point, Coos Bay,
Oregon (Lat. 43°23’ N, Long. 124°12’ W). They are in
the type collection of the Division of Invertebrate Pale-
ontology at the Smithsonian Institution, Washington, D.
C. Both are worn, incomplete, half grown forms, the
holotype measuring 44mm, the paratype 60mm in
height. Dati (1909) considered them Miocene in age
but later workers refer them to the Pliocene. The holo-
type is certainly Pliocene and the paratype may be either
Pliocene or reworked from Miocene beds near Fossil
Point.

Descriptive Notes: Mature adult specimens have at
least 8 whorls and may be 136mm or more in height.
Two conspicuous, discontinuous lateral varices occur on
each whorl, the alignment being more perfect in juvenile
forms. Shoulders are tabulate and whorls rounded and
inflated. Shells are high spired, a character that is not
evident in adults with broken apices. Differences in
sculpture are seen in progressive growth stages, which
also show considerable intraspecific variability. Numerous
axial ribs bearing nodes at the junctions with bifurcated
spiral costae are generally present on juvenile whorls but
obscure to totally absent in adults. Most Pliocene adult
specimens are characterized by sharply incised spiral
grooves. Although pronounced in material from tar seeps
in the Santa Maria District, California, incised spirals are
not produced by a particular mode of preservation.

The aperture is ovate and modified by a marked anal
notch near the posterior parietal callous deposit. The
slightly flexed pillar and outer lip bear plications that
are present in juveniles and half grown adults, but
lacking in large gerontic forms in which the outer lip
is flared.

Juveniles of Mediargo mediocris have been confused
with those of Fusitriton oregonensis, although spiral

sculpture differs slightly and E oregonensis lacks pillar
furrows and regularly spaced varices. The division be-
tween Mediargo mediocris (Datt) and M. mathewsonii
(Gasp) is drawn between strongly grooved Pliocene and
late Miocene specimens of the former and older, more
coarsely sculptured forms of the latter, none of which
attains the large size of M. mediocris adults.

Distribution in Time and Space: Range: middle (?) or
late Miocene to late Pliocene, from the Olympic
Peninsula, Washington to San Diego, California.

Stratigraphic units from which the species has been
collected include the Quillayute Formation (Pliocene)
of Washington, Astoria Formation (Miocene) and Em-
pire Formation (Pliocene) of Oregon, and the Falor,

Wildcat, Merced, Etchegoin, Fernando and Niguel For-

mations (all Pliocene) of California.

ACKNOWLEDGMENTS

Specimens in addition to those in the collections at
Stanford University were made available by Drs. Thomas
Waller and Druid Wilson of the U.S. National Museum,
Dr. Leo G. Hertlein of the California Academy of
Sciences, Mr. Joseph Peck of the University of California
at Berkeley, Mr. George Kanakoff, Mr. Edward Wilson,
and Mr. Louie Marincovich of the Los Angeles County
Museum of Natural History, Dr. Warren Addicott and
Mr. J. G. Vedder of the U.S. Geological Survey, Menlo
Park, and Mrs. Carole Hickman, Department of Geology,
University of Oregon. Their kind generosity in providing
loans and locality information is gratefully acknowledged.
The manuscript was read by Dr. A. Myra Keen, whose
suggestions and criticisms are most appreciated.

LITERATURE CITED

Carson, C. M.
1926. New molluscan species from the California Pliocene.
So. Calif. Acad. Sci. Bull. 25: 49 - 62; plts. 1-4
Dati, WILLIAM HEALEY
1909. Contributions to the Tertiary paleontology of the Pa-
cific coast. I. The Miocene of Astoria and Coos Bay, Oregon.
U.S. Geol. Surv. Prof. Paper 59: 1 - 278; plts. 1 - 23
Gass, Witt1am More
1866. Paleontology of California, v. II, Cretaceous and
299 pp; 36 plts. (1869)

Grant, Utysses S., IV. « Hoyr Ropney GALE

Tertiary fossils.

1931. Catalogue of the marine Pliocene and Pleistocene
Mollusca of California and adjacent regions. Mem. San
Diego Soc. Nat. Hist. 1: 1- 1036; 15 text figs.; plts. 1 - 32

(3 November 1931)

Page 44 THE VELIGER

Terry, JupiTH S.

1968. Taxonomy and distribution in space and time of the
marine gastropod genera Argobuccinum, Fusitriton, and Priene
(Family Cymatiidae). Unpubl. Ph.D. thesis, Stanford
University, 203 pp.; 11 plts.

WEaveR, C. E.

1943. Paleontology of the marine Tertiary formations of
Oregon and Washington. Univ. Washington Publ. Geol.
5 (2): Mollusca, pp. 275 - 562; (3): plts.; faunal tables, bibl.,
faunal localities, index, pp. 563 - 790; 104 plts.

Wooprinc, WENDELL PHILLIPS & Mitton NuNN BRAMLETTE

1950. Geology and paleontology of the Santa Maria District,
California. U.S. Geol. Surv. Prof. Paper 222: 1 - 185; plts.
1-23

Vol. 11; No. 1

Vol. 11; No. 1

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Page 45

The Role of Behavioral Traits in Influencing the Distribution

of Two Species of Sea Mussel,

Mytilus edulis and Mytilus californianus

J. R. E. HARGER

Department of Biological Sciences
University of California at Santa Barbara, Santa Barbara, California '

(3 Text figures)

INTRODUCTION

IN THE COURSE of an investigation into the nature of
competitive interactions between two species of sea mus-
sels (Mytilus edulis Linnatus, 1758 and M. californi-
anus Conrap, 1837) on the coast of Southern Califor-
nia, I discovered that in addition to certain morphol-
logical characters which served to differentiate the two
species, a difference in behavior was also apparent. Know-
ledge of this behavior enabled me to account for a
difference in the distribution of the two species in large
mussel clumps occurring intertidally on pier pilings. The
pier concerned (property of Signal Oil & Gas Company)
is located at Ellwood, some 14 miles west of the city of
Santa Barbara, California, on an open sandy shore. It
is constructed from steel girders and is about half a mile
long; the pilings support clumps of mussels in the inter-
tidal region, which range in size from a few individuals
up to giant masses some 17 feet (5.2m) in circumference
with a vertical extent of around 9 feet (2.7m).

A. FIELD INVESTIGATION

METHODS

I investigated the distribution of the two species of
mussels within the mussel clumps in the following man-
ner. Before removing samples of mussels from the clumps
on the pilings, I sprayed the outside animals thoroughly
with white enamel paint. This provided an objective

1 Present address: Department of Zoology, University of British
Columbia, Vancouver, B. C., Canada

70
60 60
50 50
40 40
30 30
a 20 20
5 10 10
go Co)
= Oo
= ° 210 oO 13
20 20
° °
0 Co)
j Size (cm) “Y
Figure 1

Size distribution of Mytilus edulis and Mytilus californianus
from the inside and outside of a mussel clump at Ellwood Pier.
The total sample represents approximately one third (by volume)

of a clump measuring 7 feet (2.13m) in cricumference.

Diagram 1: Mytilus edulis from outside the clump. Sample size is
262, mean is 4.6cm

Diagram 2: Mytilus edulis from inside the clump. Sample size is
72, mean is 3.5cm

Diagram 3: Mytilus californianus from inside the clump. Sample
size is 175, mean is 3.10cm

Diagram 4: Mytilus californianus from outside the clump. Sample
size is 53, mean is 7.82 cm
The class interval for each histogram is 1.0cm.

Page 46

THE VELIGER Vol. 11; No. 1

Table 1

Numbers of Mytilus edulis and Mytilus californianus from the
inside and outside portions of a sample taken from a mussel clump
containing both species. The sample represents approximately one
third (by volume) of a clump measuring 7 feet (2.13m) in

circumference.
Outside Inside
Mytilus californianus Mytilus edulis Mytilus californianus Mytilus edulis
Top 68 117 317 72
Middle 168 555 687 302
Bottom 73 262 343 114

method for determining whether a particular mussel came
from inside or outside a cluster. Mussels completely or
partly covered with paint were recorded as “outside”
mussels and the rest as “inside” mussels (hereafter the
term “inside” mussels refers to animals selected in this
manner). The mussels were detached by taking a straight
hoe and slicing the inside individuals away from a piling
face, from the top to the bottom of the aggregation. In
this way, a sample of 4 to 4 of the complete clump was
removed at one time. I then divided the slab of mussels
into 3 equal portions: top, middle, and bottom. Individ-
ual mussels of each species then were classified into 6
categories.

Cs ee

iType Mytilus californianus or Mytilus edulis
Position Outside or Inside

Depth Top, Middle or Bottom

RESULTS or FIELD INVESTIGATION

Table 1 records the number and positions of mussels
within 4 of a clump measuring 7 feet (2.1m) in circum-
ference. In mixed aggregations most of the Mytilus
edulis occur on the outside of the clumps. Mytilus edulis
juveniles (up to 2.5cm) tend to be found on the outside
of the clumps while M. californianus juveniles tend to
occur within the body of the clump. Figure 1 illustrates
this distribution.

B. LABORATORY INVESTIGATION

METHODS ann RESULTS

For the sake of convenience I will include experiments
carried out in the field under the above heading (ex-
periments of this nature were carried out at Ellwood
Pier and in the Santa Barbara Harbor).

In the course of studying competition among juvenile
mussels (1 -2cm in length), I assembled cages contain-
ing Mytilus edulis mixed evenly with M. californianus
(100 individuals of each species). These cages were sus-
pended beneath Ellwood Pier below low water level.

At the first inspection of this experiment (after an
immersion interval of 1 month) evidence implying a
difference in behavior between the two species was re-
vealed. Most of the Mytilus edulis in the cages had
moved to the outside of the enclosed clumps, leaving the
M. californianus in the center. Such a difference in
behavior would plainly be of advantage for M. edulis
in a competitive situation since it would seem reasonable
to assume that mussels on the outside of a clump would
be the first to capture food particles from the surrounding
water and, in addition, would not be pressed upon, and
therefore interfered with, by their neighbors. This im-
plied difference in behavior suggested a mechanism
which would account for the disparate clump distribu-
tion shown by the two mussel species. It seemed reason-
able to suppose that any M. edulis individuals settling
within the matrix of a cluster would move to the outside,
attach firmly, and then grow to maturity.

To test the possibility of such a difference in behavior
between the two species I took small individuals (1 - 2 cm)
of both species into the laboratory, placed them (sepa-
rately) about 5cm below the surface of road gravel (0.5
to 0.7/5 cm diameter) within perforated plastic containers
and immersed these containers in running sea water.
Figure 2 records the cumulative numbers of mussels ap-
pearing on the surface of the gravel with the passing of
time. Mytilus edulis obviously crawls out from the gravel
at amuch higher rate of speed than does M. californianus.

When individuals of Mytilus edulis were placed 5cm
below the surface of a gravel filled plastic tube (1.5
inches [3.81cm] in diameter) which was sealed at one
end, they always crawled through the gravel, towards and
out of the open end of the tube, regardless of whether

Vol. 11; No. 1

THE VELIGER Page 47

=
°
°

~I
o

oa
fo}

N
on

°o

°
re)
RSS
EN

[e2)
sr
»
ve)

n
=
.)
(o)

148

Number of Mussels Appearing on Surface

Time (hours)

Figure 2

Comparison between the rate of “crawling out” by Mytilus edulis
and Mytilus californianus when both species are initially buried
under 5cm of road gravel.

Two replicate runs for each species are recorded. Solid symbols
represent Mytilus californianus, open symbols represent Mytilus
edulis, The total number of animals used for each trial was 100.

the opening faced directly upwards, horizontally or at
any intermediary angle.

If small mussels (1 -2cm) are buried in gravel at a
depth greater than 13cm, very few of them are able to
crawl to the surface, presumably because the weight of
the stones is too great for them to move against. This
probably helps to explain why the mussels crawled to-
wards the open end of the tube rather than the closed
end.

Initially I had thought that small Mytilus edulis might
settle within the matrix of the clump of mussels and then
crawl to the outside using light as a possible orientation
cue. To test this hypothesis I placed 3 cages containing
small mussels at a depth of 5cm below the gravel sur-
face, inside light-tight boxes through which ran sea water
The first cage was illuminated from the top by a direc-
tional light beam, the second was illuminated from the
side and the third was kept completely dark. All mussels
crawled to the top of the cages and no difference in
“crawl-out” rate could be detected among the 3 treat-
ments. This indicated that it was not a positive reaction
to light which caused Mytilus edulis to orient themselves
on the outside of the mussel clumps.

This “crawling-out” reaction, then, seems to be evoked
in response to some tactile sensation induced by the pres-
sure of objects resting against the mussels. This hypo-
thesis is supported by the fact that when small Mytilus
edulis are placed separately within the interstices of a
cairn of “fist-sized” stones in such a manner that they
press neither against themselves nor against the stones,
they do not crawl out (except for an occasional individ-
ual), but remain fixed in one place.

The only other mussel to occur in large numbers on the
coast of Southern California is Septifer bifurcatus (Con-
RAD, 1837), a rather small animal (maximum size 3.5 to
4.5cm), found only on exposed shores. The crawling be-
havior of Septifer is quite different from that of Mytilus
edulis or M. californianus. When Septifer is placed a few
centimeters below a gravel surface it dos not move from
its position. When placed on top of a bed of gravel or on
a pitted surface, Septifer reacts by drawing itself down
into any depression which it may encounter.

Septifer is generally found in the mid-intertidal area
where it nestles in nooks and crannies. Where all 3 species
of mussels occur together, Septifer is to be found right
next to the rock face. Mytilus californianus is next, on top
of Septifer, and finally, M. edulis occurs on the outside
of such a “three species” clump.

If one considers the two extremes of exposure to wave
impact to which Mytilus edulis and M. californianus are
separately adapted, the behavioral differences can be
correlated with a presumed advantage for each species.
In quiet waters, where M. edulis predominates, mussel
clumps tend to become saturated with fine mud which
settles out from the still water. Table 2 records the mean
dry weight of silt taken at bi-monthly intervals from
cages immersed in Santa Barbara Harbor (heavy silting),
and Ellwood Pier (less silt). Plainly it would be advan-
tageous for any individual to be able to crawl above this
mud and so avoid being buried.

Table 2

Comparison of the dry weight of silt taken from cages of mussels
sct at Santa Barbara Harbor and at Ellwood Pier (below low
water), at bi-monthly intervals.

N Ye S N Y )
1 4 72.13 + 43.38 4 6.13 +3.18
2 2 34.80 + 22.0 2 6.88 +6.03
3 2 113.40 + 55.80 2 8.40 +1.20
4 4 54.45 ar 114) 8) 3 3.51 2°13
5 2 89.75 = 4.5 2 7.04 +£5.98
N = sample size Y = mean S = standard deviation

[In this connection it may be noted that when
small mussels of both species are grown together
within cages suspended in the Santa Barbara Harbor,
Mytilus edulis crawls to the outside of the clumps,
leaving M. californianus on the inside to be eventually
smothered by the accumulating silt.]

On the exposed coast where Mytilus californianus pre-
dominates, it would perhaps be advantageous to this
animal to attach itself quite firmly to the substratum

Page 48

TEM EIGER

Vol. 11; No. 1

since any individual crawling to the outside of a clump
would presumably run the danger of being washed off. In
fact, pure Af. californianus clumps tend to be very
tightly bound up with byssal threads and it is almost
impossible to tear chunks of such clumps loose with
one’s bare hands. In contrast to this, when M. edulis
forms pure species clusters (i. e., on pilings within har-
bors), it is possible to tear these clumps apart with little
effort. It is as if individuals of M. edulis sabotage the
structure of the clumps of which they are a part by con-
tinually attempting to crawl out on top of each other.

An alternative reason for the marked crawling behav-
ior of Mytilus edulis might be that this is a “behavioral
character displacement” which has been evolved through
competition with M. californianus (i.e., individuals of
M. edulis which become trapped within a clump are soon
crushed and only those on the outside of a clump may
survive).

To throw some light on the latter hypothesis, I took
some Mytilus edulis from the East Coast of the U.S.A.
(Cape Ann, Massachusetts) where M. californianus does
not occur, and compared their “crawl-out” behavior with

25

~ x)
° nn °

Number of Mussels Appearing on Surface
o

QO 6 1

18 24 30 36 42 48 54 60 66 72

‘Time (hours)

Figure 3

Comparison between the rate of “crawling out” shown by Mytilus
edulis from the West Coast of North America (Santa Barbara,
California) and Mytilus edulis from the East Coast of North
America (Cape Ann, Massachusetts), when both samples are
initially buried under 5cm of road gravel.
Three replicate runs for each sample are recorded, solid symbols
representing West Coast Mytilus edulis, open symbols representing
East Coast Mytilus edulis, with total number of animals used for
each trial 25 and the water temperature at 18° C.

that of the West Coast (Santa Barbara) M. edulis. The
East Coast mussels crawled out from the gravel at a
much faster rate than the West Coast mussels (Figure 3).
This experiment was repeated in water at temperatures
of 12°C and 18°C (the latter was the temperature of
Santa Barbara ocean water) with similar results. Since
M. edulis occurs by itself on the East Coast, it would
seem that this behavior might have developed as an adap-
tation to the physical environment rather than as a
response to interspecies competition. If this were so, one
could then view “crawling out” behavior as a preadap-
tation for competitive interaction with M. californianus.

DISCUSSION

Crawling behavior of Mytilus edulis has been noted by
European workers, i. e., Fietp (1922) and Maas GEEs-
TERANUS (1942). FieLp (op. cit.) described the course
of several small mussels as they worked their way up an
aquarium wall. Movement takes place in the following
manner: the mussel extends its foot and at the same
time produces a byssal thread which it then fastens by
means of the foot-groove and associated glands at the
tip of the foot, then releasing its old thread(s) and pulling
itself onto the new one. The process is then repeated
until journeying is done. This characteristic behavioral
trait of M, edulis is in itself rather insignificant but it
is enough to give the animal a considerable advantage
when competing with M. californianus (HarceEr, 1967).
Of course, the position on the outside of a mussel clump
is not entirely without danger, as the likelihood of being
swept away by waves or taken by some predator is
greatly increased (HarRGER, op. cit., LANDENBERGER,
1967).

It is possible that it is this “crawling out” behavior
which has enabled Mytilus edulis to become so wide-
spread throughout the world (see Stussrnes, 1954, for
an account of the distribution). Certainly M. edulis is
spread widely throughout the Northern and Southern
hemispheres and, further, it is known to successfully col-
onize both rocky shores and muddy bottomed bays
(Lewis, 1953).

It would indeed be interesting to know whether any
other species of mussel is as adept at this crawling be-
havior as Mytilus edulis.

SUMMARY

I have shown that a difference in behavior exists between
Mytilus edulis and M. californianus in that the former

Vol. 11; No. 1

THE VELIGER

Page 49

species tends to crawl more rapidly out from under
objects which press against it, than the latter. A third
mussel, Septifer bifurcatus, does not show this behavior
but rather tends to pull itself down against the substratum.

The crawling behavior ensures that Mytilus edulis be-
comes arranged on the outside of mixed species (M. cali-
fornianus and M. edulis) mussel clumps and so enjoys
an initial competitive advantage over M. californianus.
This behavior also insures that M. edulis can keep above
the surface of mud when it colonizes soft muddy areas
(bays, etc.).

ACKNOWLEDGMENTS

This work forms part of a Ph. D. dissertation submitted
at the University of California at Santa Barbara. I wish
to thank Dr. J.H.Connell for collections of Mytilus
edulis made at Cape Ann, Massachusetts, for guidance
and for criticism and above all for allowing me to develop
ideas under his guidance. I wish also to thank Dr. D. E.
Landenberger and Mr. J. Stimson for constant and
helpful criticism, and Dr. E. W. Fager for his contribution
in the form of a pointed question which led to my dis-

covering that mussels could crawl sideways as well as
upwards.

iw
SAN

secu

CT
tS

Finally I would like to thank the Signal Oil and Gas

Company for making their pier at Ellwood, California,
available for ecological research.

LITERATURE CITED

Fie.p, Irvine A.

1922. Biology and economic value of the sea mussel (Mytilus
edulis). Bull. U.S. Bur. Fish. 38: 127 - 259
Harcer, JouN R. E.
1967. Population studies on Mytilus communities. Ph. D.

dissertation, Univ. Calif. at Santa Barbara
LANDENBERGER, DONALD E.
1967. Studies on predation and predatory behavior in Paci-

fic starfish (Pisaster). Ph. D. dissertation, Univ. Calif.
at Santa Barbara

Lewis, J. R.

1953. The ecology of rocky shores around Anglesey. Proc.
Zool. Soc. London 123: 481 - 549

Maas GEESTERANUS, R.A.

1942. On the formation of banks by Mytilus edulis (L.).
Arch. Neerl. Zool. 6: 283 - 326

Stussincs, H. G.

1954. Biology of the common mussel in relation to fouling
problems. Research, London 7: 222 - 229

Page 50

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Vol. 11; No. 1

Notes on the Food of Conus dalli Stearns, 1873

JAMES W. NYBAKKEN

Moss Landing Marine Laboratories, Box 223, Moss Landing, California 95039

Conus dalli STEARNS, 1873 HAS LONG BEEN considered to
be the West American representative of the C. textile
group and indeed its color pattern is such that it is
difficult to distinguish from its Indo-Pacific congener
(Hanna, 1963).

Koun (1959) has reported that Conus textile Lin-
NAEUS, 1758 is a mollusk eater, and it is therefore of
interest to know whether or not C. dalli shares this die-
tary preference. To my knowledge the food of this species
has not been reported in the literature, a gap which may
be due in part to the fact that it is not commonly collected
alive. This note reports on observations of C. dalli feeding
on mollusks.

On November 13, 1967, while I was participating in
Stanford University’s Te Vega Expedition 16 ', four living
specimens of Conus dalli were collected at Cape San
Lucas, Baja California, Mexico, by members of the ex-
pedition. These four specimens were all collected in a
small area of loose rock on a sand substrate. The depth
varied from 5 to 8 feet. All four specimens were found
buried in sand under rocks. Specimens of C. diadema
Sowerby, 1834, C. tiaratus BropErip, 1833, and C. nux
BroperiP, 1833, were collected at the same time.

I kept the four specimens of Conus dalli alive in a
small aquarium on the ship. Three days after they were
collected I attempted to feed them by introducing a
specimen of the gastropod genus Acanthina (probably
A. tyrianthina Berry, 1957) into the aquarium. After
about half an hour, during which time the potential
prey made several passes around the aquarium very near
to, and sometimes over, the C. dalli specimens, the lar-
gest cone began to show interest and extruded its pro-
boscis. The proboscis was moved around to the posterior
part of the prey, and the animal was stung on the
postero-lateral portion of the foot. The Acanthina imme-
diately retracted into its shell. The C. dalli again probed
the aperture area with its proboscis, but I could not
determine whether the prey was stung again. The C.
dalli then rolled the Acanthina over with the front of its
foot so that the aperture faced upward toward the cone.
The front part of the foot remained in place over the

* Operated under NSF Grants GB 6870 and GB 6871

siphonal notch, and again the proboscis made several
additional probings of the aperture.

Following the probings, the buccal area expanded over
the aperture of the Acanthina, obscuring any further ob-
servation. This condition prevailed for 27 minutes. At the
end of this period the cone withdrew its buccal area but
remained in position over the shell of the Acanthina for
another 4 minutes before moving off.

A check of the shell of the Acanthina immediately
after the cone had moved off revealed that it was com-
pletely empty, the whole animal having been consumed
by the cone.

I subsequently brought all four specimens of Conus
dalli back to my aquaria at Moss Landing, where they
were placed together with several other West American
Conus species.

A few days after introducing the Conus dalli into the
aquarium I noticed a large number of empty shells of
C. nux. This sudden increase in mortality of a cone that
had theretofore lived well in aquaria led me to suspect
that C’. dalli was preying on C. nux. This suspicion was
confirmed when I placed one specimen of each of the
two cone species in an isolation aquarium. The initial
attack was not observed, but the position of the C. dalh
while feeding on the C’. nux was similar to that observed
for Acanthina.

The Conus dalli showed a marked preference for C.
nux over other Conus species. Conus dalli did not, for
example, attack C. diadema or C. tiaratus of a size similar
to that of C. nux even though they were left together in
aquaria for several weeks. Whether C’. nux is the normal
food of C. dalli in the natural environment is not known,
but both species were collected together so it would not
seem unreasonable to suspect this.

LITERATURE CITED

Hanna, G Datias
1963. | West American mollusks of the genus Conus; II. Calif.
Acad. Sci. Occ. Papers 35:1 - 103; plts 1-11 (28 Jan. 1963)

Koun, ALan J.
1959. The ecology of Conus in Hawaii.
29: 47 - 90.

Ecol. Monogr.

Vol. 11; No. 1

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Page 51

The Interrelationships of Certain Boreal and Arctic Species

of Yoldia MOLLER, 1842

BY

I. McT: COWAN

University of British Columbia, Vancouver, British Columbia, Canada

(Plate 5; 9 Tables)

MEMBERS OF THE PROTOBRANCH genus Yoldia are among
the most widely distributed Lamellibranchia in northern
oceans. They inhabit mud bottoms to depths of 100
fathoms + and may be the most abundant mollusk on
them. Their abundance in the stomachs of bottom-feeding
fishes suggests that they are an important element in the
benthic food chains.

Within this genus there is a group of species charac-
terized by the simultancous presence of an elongate form
with tapered, sub-acute posterior extremity, an approxi-
mately central umbo, slender rather than bulbous form,
a highly polished cuticle devoid of sculpture or surface
pattern of any kind, and with an unstriated resilifer.

Several names have been applied to members of this

group:

Yoldia amygdalea VALENCIENNES, 1846, described
from specimens obtained on the cruise of the
‘Venus’ at Kamchatka.

Yoldia hyperborea (LovEN) Toreti, 1859, was de-
scribed on the basis of specimens from Spitzbergen.

Yoldia limatula gardneri OupRoyp, 1935, from Gar-
den Bay, Pender Harbour, British Columbia.

Yoldia limatula (Say, 1831) was based upon speci-
mens taken in fish stomachs in Massachusetts
waters.

Yoldia norvegica. DAUTZENBERG & FISHER, 1912
regarded the specimens from the coast of Norway
as distinctive and applied this name, without des-
ignating a holotype.

Yoldia hyperborea limatuloides OCKELMANN, 1954,
is the most recent name to appear. The type
locality is Fossfjord, N. W. Iceland, but the form
is stated to occur also from the Lofoten Islands to
eastern Finmark.

All have been described from shell characters alone
and in only one instance (OcKELMANN) has an attempt

been made to express the observed differences quantita-
tively. He used 3 shell ratios in an attempt to distinguish
Yoldia limatula, Y. sapotilla and Y. hyperborea and found
them to be of little use.

One of the problems of developing quantitative ex-
pression of difference in the form of the lamellibranch
shell is that in most species it grows throughout life. In
many species also the shell changes in proportions with
growth and thus renders dimensional ratios meaningless.
This leaves the possibility that in any comparison of shell
dimension between different populations the bias due to
age or size may be great.

In the present study I have attempted to avoid this
problem by regressing pairs of measurements so as to
develop an expression of the relative growth of the dif-
ferent parts of the shell in each apparently distinct popu-
lation. Analysis of covariance was used in making mathe-
matical comparison between the equivalent regression
coefficients of different populations.

In order to develop these criteria my series for meas-
uring were selected to contain the widest possible size
range from each locality.

METHODS

Samples from individual geographic locations were given
close visual inspection followed, in some cases, by trial
comparison of ratios. Where no or almost no differences
were apparent, the samples were pooled into composite
samples suspected of representing definable entities of
possible taxonomic value. Greatest length of the shell
has been used as the basis of each regression. While the
slope of the regression line is certainly a_ biologically
meaningful expression, the position of the intercept on
the Y axis is probably no less a characteristic of the
population. Its biological meaning, however, is less ap-
parent.

Page 52

All statistical procedures were undertaken on a digital
computer.

MEASUREMENTS

All measurements were taken with a sliding caliper

equipped with a dial vernier. The measurements taken

and the symbols employed to designate them on the ana-
lytical tables are as follows:

X = The greatest length of a single valve taken in a
straight line. Left and right valves were measured in
each instance where they were available and intact.

Y1 = The greatest height of the valve from ventral mar-
gin to peak of the umbo taken at right angles to the
dorsal plane.

Y2 = The length of the pallial sinus measured from the
posterior tip of the shell.

Y3 = The anterior length of the valve taken from highest
point on the umbo to the most distant point on the
anterior margin.

Y4 = The posterior length measured similarly.

Y5 = The number of teeth on the posterior side of the

umbo.

Y6 = The number of teeth on the anterior side of the
umbo,

Y5 + Y6 = thus the total number of teeth on the hinge
line.

The small teeth close to the resilifer were difficult to
count on certain large specimens but every effort was
made to expose them and count them accurately.

POPULATIONS SELECTED
FOR COMPARISON

The selection of populations to be compared statistically
is a Critical step in the process of exploring for significant
differences. I know of no substitute for careful visual
inspection of the field samples supported by trial com-
parison of measurements and this has been the basis of
selection used here. It was felt to be important to include
for comparison those populations that had been previ-
ously described as differing from others, along with those
that the present study revealed. Thus the following stocks
have been treated separately.

1. The form inhabiting the high Arctic seas around
the world.

2. The population of Kamchatka, the southwest Ber-
ing Sea and the Sea of Okhotsk.

3. The population from Iceland and the coast of Nor-
way.

4, The population inhabiting the western Atlantic
from Nova Scotia to Massachusetts.

THE VELIGER

Vol. 11; No. 1

5. A small collection from depths of 155 - 285 fathoms
in Delgado Canyon, off Cape Mendocino, California.
This population appears to be discontinuous from others,
both geographically and by depth occupied.

6. The specimens from the coast of Alaska south of
the Alaska Peninsula.

7. The many localities represented in Georgia Strait,
British Columbia.

8. A sample from Malcolm Island, Queen Charlotte
Strait, British Columbia.

9. A sample from Masset Inlet, Queen Charlotte Is-
lands, British Columbia.

At the same time some local variation seemed ap-
parent along the coast of British Columbia where barriers
to free dispersion seemed to be few. To explore the nature
of variation under these circumstances, 3 populations
(7, 8, 9) were included in comparison with specimens
from the contiguous areas along the Alaskan coast.

COMPARISONS

Tables 1 to 6 present the results of covariance analysis
derived from the regressions obtained from the 9 popu-
lations. For purpose of interpretation I have used .01 as
the level of significance to which systematic meaning can
be attached.

Let us first examine the several samples drawn from
the relatively continuous north-south distribution along
the coast of British Columbia and southeastern Alaska
using the centrally placed Masset Inlet population as a
basis of comparison (Table 4). Gross inspection sug-
gests that the specimens from the Strait of Georgia are
uniformly deeper for their length than those from the
other areas. The only exception from this general con-
dition is found in the small series that served as the type
lot for Yoldia limatula gardneri OupRroyp. These are pro-
portionately shallower, i.e. more slender than the Strait
of Georgia form but can be matched among series from
the outer coast of British Columbia and Alaska.

It will be seen that the Queen Charlotte Islands stock
is identical with the Malcolm Island stock in all slopes of
the regressions and differs in only one intercept feature
— that is the posterior length. In comparison with the
Georgia Strait stock the regression of width on length is
different, confirming the general impression referred to
above. Two of the intercept figures are different, the
pallial sinus scar and the anterior teeth.

The Queen Charlotte Island sample is identical with
the Alaskan sample in all features except length — width
relationship. It differs from the California sample in two
intercept features only.

Vol. 11; No. 1 THE VELIGER Page 53

Table 1
Probability of Significance of Covariance Analysis of Slopes of Yoldia “norvegica”
Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+ Y6/x
Y. hyperborea + — — — + + '-
Y. amygdalea (K) ' — 1 — .05- .1 — — _
Y. amygdalea, Alaska — .1-.2 — 05 — _— —
Y. limatula, Atlantic — — — .O1- 025 05-.1 01-025 .025

Probability of Significance of Covariance Analysis of Intercepts of Yoldia “norvegica”

Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+Y6/x
Y. hyperborea + + aE =F + + +
Y. amygdalea (K) .05- .1 — + .01- .025 + .005- .01 a
Y. amygdalea, Alaska + + .001- .005 005 .025-.01 .025-.01 01
Y. limatula, Atlantic .1- .2 — + qF 01 + +

* Kamchatka stock
+indicates a probability of accidental occurrence < .001
— indicates a probability of accidental occurrence > .1

Table 2
Probability of Significance of Covariance Analysis of Slopes of Yoldia hyperborea
“Compared with Yi/x Y2/x -Y3/x.—sY4/x.~=s«éY'Bs/x = Y6/x = Y6/x
Y. amygdalea (K) + + 3 .025- .05 + + ar
Y. amygdalea, Alaska + .001 — 1-2 + + oF
Y. limatula, Atlantic + 01 .001 .001- .005 + at +

Probability of Significance of Covariance Analysis of Intercepts of Yoldia hyperborea

Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+ Y6/x
Y. amygdalea (K) + oF oP + + + +
¥. amygdalea, Alaska + at .01- .005 + + + +
Y. limatula, Atlantic + + 1-2 + + + +
Table 3
Probability of Significance of Covariance Analysis of Slopes of Yoldia amygdalea (K)
Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+ Y6/x
Y. amygdalea, Alaska _— — — — — — —_—
Y. amygdalea, Georgia Str. — — — — — .05 al
Y. amygdalea, California _ = — — .005- .01 at +

Probability of Significance of Covariance Analysis of Intercepts of Yoldia amygdalea (K)

Compared with Y1/x Y2/x NS} / x Y4/x Y5/x Y6/x Y5+ Y6/x
Y. amygdalea, Alaska + ain + ate .025- .05 2 .05- .1
Y. amygdalea, Georgia Str. ar .005- .01 + + .05- .1 ar .001- .005
Y. amygdalea, Humboldt — + + .005- .01 all ar 1-2
Thus along the 1000+ miles of coastline from the tential comparisons for any 2 of the sub-populations
Alaskan Peninsula to the coast of California there is a that gave rise to the samples. These represent a bathy-
great uniformity in the measurable features of this Yoldia, metric range from 10 fathoms to 200 fathoms. The con-

with significant differences in no more than 2 of 14 po- clusion reached is that the Pacific Coast is inhabited by

Page 54 THE VELIGER Vol. 11; No. 1
Table 4
Probability of Significance of Covariance Analysis of Slopes of Yoldia amygdalea, Masset

Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+ Y6/x

Y. amygdalea, Malcolm Isld. 1-2 — 1-2 1-2 — — —

¥. amygdalea, Georgia Str. + .05- .1 1-2 .05- .1 — — —

Y. amygdalea, Alaska .001-.05 .1-.2 — — — — —

Y, amygdalea, California .01- .02 — — — — 1-.15 2

Y. amygdalea (K) .005- .01 — — — — — —

Y. norvegica, Norway-Iceld. .025- .05 .005- .01

Probability of Significance of Covariance Analysis of Intercepts of Yoldia amygdalea, Masset

Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+ Y6/x °
¥. amygdalea, Malcolm Isld. _ —_— .1- .2 001-.005 .025-.05 .05-.1 .025- .05
Y. amygdalea, Georgia Str. .025- .5 + = = 1 001-.005 .01- .025
Y. amygdalea, Alaska _— _ .025- .05 .1-.2 1-2 .05- .1 .05- .1
Y. amygdalea, California + — l .001- .005 ne — —
Y. amygdalea (K) + al + ar 2 .05- .1 ol
Y. norvegica, Norway-Iceld. .001- .005 + .001-.005 .001-.005 .025-.05 .025 .025- .05
Table 5

Probability of Significance of Covariance Analysis of Slopes of Yoldia amygdalea (Bering “Intergrades”’)
Compared with Y1/x Y2/x Y3/x Y4/x Y5/x Y6/x Y5+ Y6/x
Y¥. amygdalea (K) .01- .025 05 05 — .O1 .001 .001
Y. hyperborea _— 1-2 — — .025- .05 l .05

Probability of Significance of Covariance Analysis of Intercepts of Yoldia amygdalea (Bering “Intergrades”)

Compared with Y1/x Y2/x Y3/x
¥. amygdalea (K) aP .005- .01 +
¥. hyperborea + .05 ap

a remarkably uniform stock of this species of Yoldia,
displaying only minor geographic variation in form and
in relative growth features. The most obvious of these
variants is that occupying the heavily silted bottoms of
the Strait of Georgia and Puget Sound.

Table 6 summarizes the differences appearing in the
comparisons of the 14 potential differences in numerical
attributes that could occur between each of the stocks
analysed. If these are arranged to reveal geographic. re-
placement, the Pacific Coast stock differs from that of
the Okhotsk-Kamchatka areas in a total of 5 of the

Y4/x Y5/x Y6/x Y5+Y6/x
.05 -1- .2 = a
oF aR + oP

indices, Kamchatka from the Arctic stock in 12, the Arc-
tic from Norway-Iceland in 11, and from the American
Atlantic coast stock in 13. The number of differences
appearing between the Norway-Iceland form and the
American Atlantic form is 5.

The Arctic form therefore differs from all others with
which it is in contact in 11 or more of the 14 features
studied here, whereas none of the other stocks, even those
separated by the entire extent of the Arctic Seas, and
presently occurring in the North Atlantic and North Pa-
cific, differ in more than 5 features when one uses the .01

Explanation of Plate 5

Figure 1: Yoldia amygdalea, Sakhalin Island, NW Pacific. X 1.2

Figure 2: Yoldia amygdalea, Masset, Queen Charlotte Island,
British Columbia x1
Figure 3: Yoldia amygdalea, Strait of Georgia, British Columbia
X 1.6

Figure 4: Yoldia amygdalea, California Coast X 1.2
Figure 5: Yoldia amygdalea, Iceland xX 1.6
Figure 6: Yoldia hyperborea, Point Barrow, Alaska xX 1.5
Figure 7: Yoldia hyperborea, Disco, Greenland xX 15
Figure 8: Yoldia limatula, Eel Pond, Massachusetts xX 2

THE VELIGER, Vol. 11, No. 1 [Cowan] Plate 5

Figure 7

Vol. 11; No. 1

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Page 55

Table 6

Numbers of Features Distinguishing Stocks of Yoldia

Comparison Number of Significant Differences
Slope (max. 7) Interc. (max. 7) Totals (max. 14)
(a) (b) (a) (b). (a) (b)
Significant Significant
at.01 at.05 at.01 at.05
1. Masset : Malcolm Isld. 0 0 1 3 1 3
2. Masset : Georgia Str. 1 1 1 4 2 5
3. Masset : Alaska 1 1 0 1 1 2
4. Masset : California 0 1 2 2 2 3
5. Masset : Kamchatka 1 1 4 4 5 5
6. Masset : Norway-Iceland 1 2 3 6 4 8
7. Arctic : Pacific Coast 5 5 7 7 12 12
8. Arctic : Amer. Atlantic 7 7 6 6 13 13
9. Arctic : Norway-Iceland 4 4 7 7 11 11
10. Arctic : Kamchatka 5 6 7 7 12 13
11. Kamchatka : Norway-Iceld. 0 1 4 5 4 6
12. Norway-Iceland :
American Atlantic 0 3 5 5 5 8

level of probability as the criterion of significance. The
.05 level of significance does not alter the situation re-
specting the Arctic population. It is clearly a very differ-
ent stock. This level, however, reveals the Georgia Strait
stock as having more distinctive features than the other
samples drawn from the NE Pacific. It also increases the
apparent difference between the Norwegian and the A-
merican Atlantic stocks.

Returning to the differences at the .01 level (total “a”
of Table 6), and comparing geographically contiguous
stocks, it is possible to suggest that 3 clearly differentiable
stocks exist: (1) that inhabiting the NE Pacific Ocean,
the Sea of Okhotsk, the coast of Kamchatka and the
southern Bering Sea; (2) the Arctic Seas of both hemi-
spheres; and (3) the shallow waters of the North Atlantic
Ocean including Iceland and the coast of Norway. The
American Atlantic stock differs from Arctic and Pacific
stocks to a similar degree.

The data derived from the covariance analysis of
regressions and intercepts reveals the numbers of criteria
by which the several stocks differ from one another but
tell us little about the nature or direction of the differ-
ences. These details can be derived from inspection of the
specimens and from the conventional measurements.
They do not alter the conclusions to be reached from the
present analysis.

Specimens originally from Iceland and the coast of
Norway are remarkably similar in general appearance to
the specimens taken along the Pacific coast of North
America. In fact, the characteristics by which they each
differ from the Arctic population (Tables 1-4) are in

general the same. One might ask, therefore, are they
indeed the same organisms separated into the North At-
lantic and North Pacific stocks by an inhospitable area
of Arctic environment occupied by another form.

A direct comparison between the North Pacific and
North Atlantic stocks reveals that, at the .01 level of
significance, they do not differ in any of the 7 regressions
(Tables 1 & 4), but do so in 4-6 of the intercepts, de-
pending upon which Pacific Coast sample is used in the
comparison. Clearly then, the differences between the
stocks of the two oceans is greater than the differences
between any two of the samples from the Pacific coast
of North America.

The relationship between the North Atlantic stock and
that of the American Atlantic is also interesting. As
OckKELMANN (1954) pointed out, there are distinct and
consistent differences in the shape of the shell between
these two stocks. The American Atlantic specimens lack
the anterior and posterior indentations in the ventral
margin of the shell. There are differences also in the
shape of the siphonal muscle scar. My data suggest two
features of shell proportion and dentition (p < .025)
and 5 intercept characteristics that distinguish the two
Atlantic stocks.

The general form of the North Atlantic specimens
suggests that they might be the outcome of intergrada-
tion between the Arctic form and that of the American
Atlantic. Their geographic situation and the absence of
any substantial evidence of intergradation along the
coasts of Newfoundland and Labrador leave us with no
evidence to support such an explanation. I have seen no

Page 56

THE VELIGER

Vol. 11; No. 1

series of Yoldia from Newfoundland and Labrador, so
future collections may require a re-examination of the
relationships between the 3 forms encroaching upon the
North Atlantic.

TAXONOMIC CONSIDERATIONS

It has been shown that there are few differences between
several stocks of Yoldia separated widely along the Pacific
coast of North America. It has also been shown that
these do not differ substantially from the form inhabiting
the Sea of Okhotsk and the Kamchatka coast. The first
available name for this stock is Yoldia amygdalea VALEN-
CIENNES, 1846.

The name gardneri O.pRoyp, 1935, applied to a sup-
posed subspecies of Yoldia limatula (now Y. amygdalea)
in the northern part of the Strait of Georgia, British
Columbia, appears to be without foundation. It referred
to unusually slender examples that occur among the
normal population.

Yoldia hyperborea (Loven) Toreti, 1859 is the ac-
cepted name applicable to the distinctive and remark-
ably uniform stock of Arctic Seas.

In my opinion, the characteristics of the stock from the
North American coast of the Atlantic, taken with the
absence of certain evidence of intergradation with Yoldia
hyperborea to the north argue for recognizing this as a
separate species under the name Y. limatula (Say, 1831),
that has been used for this form for more than a century.

OcKELMANN (1954) regarded the form of the North
Atlantic as a subspecies of Yoldia hyperborea and applied
the name limatuloides to it. It has now been shown that
this population differs as profoundly from Y. hyperborea
as any of the other stocks compared. The weight of evi-
dence now suggests that this stock be regarded as con-
specific with Y. amygdalea and that this species, as is
the case with Y. myalis, is now divided into Atlantic and
Pacific stocks differing but little from one another. If
a subspecies designation be regarded as useful, the name
Y. norvegica DAUTZENBERG & FiscHeER, 1912, though not
described with a designated type specimen, was clearly
applied to the form of the Norwegian coast and has
priority over limatuloides of OckELMANN. This author
regarded Y. norvegica as a synonym of Y. hyperborea, but
the geographic reference by DAUTZENBERG & FISCHER
seems to deny this. In this instance there seems to be no
need for a separate designation for this stock.

It is pertinent to ask whether there is any strong
evidence of intergradation between stocks I have defined
as possessing characteristics suggesting clear cut differ-
ences. Thirteen specimens from the eastern Bering Sea

(St. Matthews Island, Nunivak Island, N of Unimak
Island, near Pribiloff Islands) originate from an area
where intergradation between Yoldia hyperborea and Y.
amygdalea might be expected to occur. They do indeed
show characteristics of form apparently intermediate in
some degree. Table 5 presents the probabilities of signi-
ficance of covariance analyses comparing the possible
intergrades with both the putative parent stocks. While
Y. amygdalea differs from Y. hyperborea in 12 criteria, it
differs from the supposed intermediates in 5 and Y. hyper-
borea differs from them in 6. The differences are in the
expected direction.

On the strength of these comparisons, and others in-
volving unmeasurable aspects of shell form there seems
to be evidence that these specimens are indeed inter-
mediates between the Arctic and North Pacific-Kam-
chatka forms. The question then arises as to the inter-
pretation to be placed upon this intergradation. It could
be regarded as evidence that complete introgression is
possible and that the two populations should be regarded
as subspecific expressions of a single wide ranging species.

In the genus Yoldia, Y. myalis and Y.thraciaeformis both
occur in twin populations occupying the North Pacific
and North Atlantic and separated by the Holarctic re-
gion in which they do not now occur. In this study
evidence is produced to reveal Y. amygdalea as similarly
distributed. The first two species named have no surviving
close relative in Arctic waters. One can conclude, there-
fore, that they previously occupied Arctic waters but have
been exterminated in them so recently that little or no
morphological difference has developed between the relict
populations. The failure of the Arctic stocks may well
have occurred during the closure of Bering Straits,
with the considerable alteration in the circulation of
Arctic waters and possibly of local mid water habitats.
A possible alternative explanation would involve conti-
nuity of distribution through tropical waters at a time
when the two Americas were separate. This seems less
likely.

In the case of Yoldia amygdalea rather than extermi-
nation, evolutionary change gave rise to a form adapted
to arctic conditions (Y. hyperborea). The evidence of the
samples now available from the southern Bering Sea sug-
gest that genetic isolation between the two species is
incomplete. It seems to me, however, that the existence
of the North Atlantic stock of Y. amygdalea, the very
extensive range of this species in the Pacific, the extent
of the area occupied by Y. hyperborea in the Arctic, and
the absence of any evidence of intergradation between
it and the Atlantic Y. amygdalea all point to species status
as best expressing the relationship between Y. hyperborea
and Y. amygdalea despite the evidence of some introgres-
sion over a narrow area of contact in the Bering Sea.

Vol. 11; No. 1 THE VELIGER Page 57

Table 7

Regression Formulae for Yoldia amygdalea

Yoldia amygdalea (British Columbia & Alaska)

N = 76 x/Y1 x/Y2 x/Y3 x/Y¥4_x/Y5._x/Y6 Y3/Y4 Y5/Y6

Correlation between 9896 .9896 .9959 .9949 .9016 .8821 .9867 .8821
Slope of Regression 4085 8=.5248 = 5252) 5331 = 9567 «1.2466 1.0025 1.2466
Intercept 1.0274 -.2126 .6928 -.6860 8.7841 12.4484 -1.2110 12.4485

Yoldia amygdalea (Kamchatka)

N = 39 x/Y1 x/Y2 x/Y3 x/Y4 x/Y5 x/Y6 Y3/Y4 Y5/Y6

Correlation between 9749 9575 .9765 .9815 .8773 .3167 .9316 .8672
Slope of Regression 4239 =.5359 5146 = 5418 = .3908)=— 2977 Ss 9759 1.1894
Intercept 1.5869 -1.7885 2.3090 -1.7059 8.5078 19.0575 —2.4483 3.5084

Yoldia amygdalea (Iceland & Norway)

N = 22 x/Y1 x/¥2 x/Y¥3 x/Y4_x/Y5.x/Y6 Y3/Y4 Y5/Y6

Correlation between 9855 .9601 .9795 .9842 .7486 .8453 .9547 9224
Slope of Regression 4250 .4476 .5329 4819 .4164 5777 8566 1.1334
Intercept 1.2303 1.0776 .7688 .7797 10.0510 11.8195 .8456 3.4409

Yoldia amygdalea (California)

N = 27 x/Y¥1 x/Y2 x/Y3 x/Y4_x/Y5. x/Y6 Y3/Y4 Y5/Y6

Correlation between 9675 .9048 .9594 .9898 3466 .1850 .9482 .5634
Slope of Regression 4488 5059 5030 5469 1551 .2714 9992 .5672
Intercept 5167 = .7929 1.4659 -1.4475 1.9588 18.6113 -.9333 16.8931

Table 8

Regression Formulae for Yoldia hyperborea

N = 107 x/Y1 x/Y2 x/Y3 x/Y¥4 x/Y5 x/Y6 Y3/Y4 Y5/Y6

Correlation between 9935 .9906 .9835 .9848 8445 .8384 9465 .9582
Slope of Regression 4898 4444 5686 .4882 .3867 .4336 .8114 1.0750
Intercept 0602 =.3732 -1.4579 2.2097 10.2074 13.9943 4.1348 3.1618

Table 9

Regression Formulae for Yoldia limatula

N = 48 x/Y1 x/Y2_x/Y3._x/Y4__x/Y5.-x/Y6 Y3/Y4 Y5/Y6

Correlation between 9964 9965 .9990 9985 9102 9037 9967 .9674
Slope of Regression 4430 4692 5229 5268 .2799 .3431 1.0046 1.1942
Intercept 4979 §=.4674 ~=—.4598 »9=—.0071 12.3751 15.0194 -.1296 5069

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Vol. 11; No. 1

Résumé of Species

Yoldia amygdalea VALENCIENNES, 1846 (Plate 5, Figures 1 - 5)
Type locality: Kamchatka.

Original description: VALENCIENNES, A. Mollusques (in) ABEL
Du Petit-THouars. 1846. Voyage autour du monde de la
Frégate Venus. plt. 23, fig. 6.

Selected Synonymy:
Y. amygdala [sic] REEvE, 1873. Conchologia Iconica 18: plt. 1,
figs. 3a, 3b.
Y. limatula (pars) Dati, 1921. U.S. Nat. Mus. Bull. 112: 13
(and elsewhere)
(pars) La Rocegue, A., 1953. Nat. Mus. Canada 1953: 25;
Otproyp, I.S., 1924. Stanford Univ. Publ. Geol. Sci.
Y. gardneri Otproyp, 1935. Nautilus 49: 14 (Type locality:
Garden Bay, Pender Harbour, Vancouver Island, B.C.)
Y. norvegica DAUTZENBERG & FiIscHER, 1912. Résultats des
campagnes scientifique ... par Albert I, Prince Souverain
de Monaco. Fasc. 37: 403 (Type locality: Norway).
¥. hyperborea limatuloides OckELMANN, 1954. Medd. om
Gronland 107 (7): 11 (Type locality: Fossfjord. N. W.
Iceland).

Distribution: This is the species from the Okhotsk Sea and Kam-
chatka and from the Pacific Coast of North America, south of the
Bering Sea into Northern California. The Atlantic stock occupies
both coasts but comes north to Iceland but not Greenland. South
further in the west than in the east. Specimens examined from:
USSR: Avancha Bay; Plover Bay; Kamchatka coast at 55°57’N,
162°20’E; Okhotsk Sea at 52°43’N, 155°41’E; Aniva Bay,
South Sakhalin Island.
NE Pacific: Alaska: Redfish Bay, Stikine Delta; St. Matthews
Is.; Nunivak Is.; N of Unimak Is.; near Pribiloff Islands.
British Columbia: Masset, Queen Charlotte Island; Malcolm
Island; Smith Inlet; Beaver Harbour; Okeover Arm; Tofino
Inlet; Barclay Sound; Georgia Strait (many localities) .
Washington: off San Juan Island.
California: Delgado Canyon off Buck Creek, Mendocino County.
I have seen no specimens from Oregon or from the outer coast
of Washington.
Iceland: Fossfjord.
Norway: Lofoten Islands; Bergen; Ramsfjord; Vadso.
Western Atlantic: 40°29’N, 70°46’W; Bedford Basin; N of

Sable Island, Nova Scotia; Bradilla in Gulf of St. Lawrence;
Grand Manan Island; off Halifax, Nova Scotia.

Yoldia hyperborea (LovEN) TorE., 1859. (Plate 5, Figures 6, 7)
Type locality: Spitzbergen.

Original description: Torexi, O. 1859. Bidrag till Spitzenbergens
Molluskenfauna. Stockholm, pp. 149-150; plt. 2, figs.6.a,6b.

Selected Synonymy:

Yoldia arctica M. Sars, 1851. Nyt Maf. f Naturv., 6 (2).
Kristiania.

Nucula sapotilla REEVE, 1855 (non Goutp, 1841) (in) BEx-
cHER, E., The last of Arctic voyages. vol. 2, London.

Yoldia limatula G.O. Sars, 1878. Mollusca regionis arcticae
Norvegiae, Christiania.

Yoldia glacialis Gray, 1828, La RocguE, 1953. Nat. Mus. Can-
ada, Bull. 129.

Distribution: Believed to be circumboreal in the Arctic basin, but
I have seen no specimens, nor published records of occurrence,
from the North American Arctic east of Point Barrow and west of
Foxe Basin; nor from the Siberian Arctic east of Novaya Zemlya.

Specimens examined from:
Spitzbergen (several localities); Novaya Zemlya; Disco, Green-
land; West Coast Greenland; Foxe Basin; Hudson Bay; and
many stations in Canadian Eastern Arctic; Point Barrow,
Alaska. OcKELMANN (1954) cites specimens from the Murman
coast and the Kara Sea.

Yoldia limatula Say, 1831. (Plate 5, Figure 8)

Type locality: Nahant, Massachusetts.

Original description: Say, R. 1831. American Conchology, prt.
2, pit. 12, 3 figs.

Distribution: The North American coast from Nova Scotia to
North Carolina. Specimens examined from: :
Halifax, Nova Scotia; Sand Island, New Brunswick; Casco and
Frenchman’s Bay, Maine; Woods Hole, Massachusetts; New-
port, Rhode Island; Rariton Bay, New Jersey; Cape Hatteras,
North Carolina.
I have seen two specimens from close to Halifax that suggest
that intergradation may occur between Yoldia limatula and Y.
amygdalea. However, the evidence is too meager to interpret.

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Page 59

Notes on Cryptochiton stelleri (MIDDENDORFF, 1846)

G. E. MacGINITIE

NETTIE MacGINITIE

Route 1, Box 93A, Friday Harbor, Washington 98250

(Plate 6)

For QUITE A NUMBER OF YEARS, we have been intrigued
by the fact that the giant chiton Cryptochiton stelleri is
almost invariably free of epibiotic growths. As we have
emphasized (MacGinitie & MacGinitie, 1968), there
are nearly always exceptions to any generalization, so it
is not surprising that the defense mechanism of the
chiton is occasionally damaged or fails to function. On
rare occasions, a barnacle, a tuft of seaweed, or a tunicate
may be found growing on the giant chiton. These ex-
ceptions are no doubt due to some injury sustained from
a rolling rock or floating debris or other abrasive causes.

The present study was begun in an attempt to deter-
mine the means by which Cryptochiton stelleri keeps its
surface free of epibiotic growth. In order to do this, it
was first necessary to learn something of the food and
habits of the animal so that a number of them could be
maintained in aquaria in as healthy and normal condi-
tions as possible.

FOOD anp HABITS

This chiton species is found from low tidal exposures
down to a depth of 50 or 60 feet or possibly deeper. Its
range is from San Nicolas Island to Alaska and across
to Japan. It is most abundant in the northern part of
its range. Because its food consists mainly of Ulva,
Monastrema, Iridia, or other thin-fronded algae, it may
be found where such growth is abundant. We have kept
specimens in our laboratory for more than a year and
tried them with different types of seaweed, but they
show a decided preference for Ulva and Monostrema.
We have withheld food until they were hungry enough to
eat, very sparingly, some heavier types of seaweed, even
including the stipe of Nereocystis. However, they are
able to go for months without food of any kind. Examin-

ation of the fecal material reveals that the heavier
types of seaweed pass through with little apparent di-
gestion. Even some of the preferred sea weeds, such as
Ulva, pass through in larger pieces than one would
expect.

These giant chitons are extremely slow-moving crea-
tures and move very short distances, say a few inches, in
a night when browsing. Those we have kept in aquaria
seem to be strictly nocturnal. Very rarely does one move
about during the day. Two that we have kept in an
aquarium at temperatures ranging from 3°C to 5 or 6°C
ceased all visible activity while the temperature remained
this low. As most of the seaweeds are annuals, we doubt
that even in their natural environment these chitons feed
to any extent during the winter months.

Although the chitons of this species reject plants or
animals that might adhere to their dorsal surface, they do
have commensals. A scale worm Arctonoe vittata (GRUBE,
1855) and a pinnotherid crab Opisthopus transversus
RaTHBUN, 1893, are often found in the gill chamber,
usually singly, but occasionally together.

GROWTH anpb AGE

Cryptochiton stellert is the largest chiton in the world.
Specimens 330mm (13 inches) long have been recorded.
The largest specimen we used was 240mm long by 115
mm wide. It was about 32 mm high at the highest point.
The smallest was 95mm by 60mm.

This species grows very slowly. From a count of
growth lines on the dorsal valves, it appears that any
Cryptochiton stelleri that is 7 or 8 inches long or longer
is at least 20 years old. On most of the valves, the growth
lines die out and become obscure about two thirds of the
way in from the outer edge. We have counted the growth

Page 60

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Vol. 11; No. 1

lines on many valves that showed up to 18 or 20 lines
and we concluded that such specimens were more than
25 years old. We have not checked the reproduction care-
fully, but the lines of growth and habits of the animal
indicate that it spawns but once a year. Small juvenile
specimens are seldom found. This scarcity of young is
typical of marine invertebrates that live a long time.

MECHANISM ror KEEPING CLEAN

We believe we have found the answer to how Cryptochi-
ton stellert keeps so free of epibiotic growth. To under-
stand how this is accomplished, one must know something
about the structure of the dorsal integument. The entire
dorsal surface of the animal is covered with tufts of
fine spicules of calcium carbonate (Plate 6, Figures 2
and 3). Sometimes these tufts, as well as the integument,
are a dark brick red over the entire surface but in some
specimens the surface is mottled by patches of grayish
tufts of spicules. The tufts of spicules terminate abruptly
at the edge of the dorsal surface where the integument
turns under to form the under side. The number of
spicules per tuft varies from about 25 to 365 or over. On
a specimen 217mm long, the number of tufts per square
centimeter was fairly uniform, averaging 118 tufts. These
tufts of spicules may stand erect in tight vertical columns
or they may spread out more or less in the form of
rosettes. The individual spicules are 1 mm long or some-
what longer. They are individually controlled, so that
some of the spicules in a tuft may stand erect while the
others spread out.

A tuft of spicules originates as a bulbous structure
within the integument, with a small opening to the sur-
face, and gradually evaginates until it becomes only a
depression on the surface with a full bloom of spicules
(Plate 6, Figure 1). The bulbous structure is lined with
cells, each of which secretes a spicule. When a tuft of
spicules is at the surface and functional, each spicule is
controlled by a cell at the base.

Our first reaction was to suppose that these tufts spread
into rosettes to prevent the settling of larvae on the
integument, but quite the reverse is true. The introduction
of fine planktonic material to the surface of the chiton
causes the tufts of spicules that are spread in rosettes to
become erected into columns. Further study revealed that
these columns of spicules were supporting a sheet of
mucus.

We have never found a dye that will stain secreted
mucus. It is possible to stain preserved mucous cells, but
not fresh mucus itself. It is impossible to demonstrate
mucous layers by any means except by the introduction
of finely divided material. To show the presence of
mucus, we have nearly always used detritus (MacGInITIE
& MacGiniTig, 1968, p. 11) because it does not in any
Way irritate the secreting animal.

Stained sections of the integument show that mucous
glands are abundant and open to the surface in the areas
between the tufts of spicules. Though there are no tufts
of spicules on the under side, mucous secreting cells con-
tinue to be present in that portion of the integument that
turns under at the edge. With secretions from the mucous
glands, Cryptochiton is rapidly able to cover its dorsal
surface with a sheet of mucus from 1 to 2mm thick. As
is well known, when mucus comes in contact with water,
it quickly swells. As has been shown before (MacGrnitTIEz,
1945), mucus, at least as it is used by several invertebrate
animals (MacGinitie & MacGinitiz, 1968, pp. 186, 203,
336), will prevent the passage of anything larger than
40 A. Although mucus allows water to flow through it
almost unimpeded, it will prevent the passage through it
of any larvae, algal spores, bacteria, or viruses — or even
molecules with a molecular weight of 160000 or larger.

By placing some detritus or fine planktonic material on
the surface of Cryptochiton, the presence of mucus is
easily observed. We were unable to determine whether
the stimulus of a settling larva causes secretion of mucus
only in that local area or whether the mucus covers more
of the surface of the chiton. Evidence points to a pro-
gressive weakening of the mucous secretion in specimens

Explanation of Plate 6

Cryptochiton stelleri (MippENDoRFF, 1846)

Figure 1: Section through the dorsal integument; 60. The irreg-
ular diagonal line is the upper surface of the integument. Lower
left: developing tuft of spicules. Upper center: mucus secreting
glands. Upper right: section through depression where a tuft of
spicules is attached. At the base of each spicule is a cell that

regulates the position of the spicule.

Figure 2: View of portion of dorsal surface, showing tufts of
spicules; X 2.

Figure 3: Same as Figure 2; <4. Note that the tufts of spicules
are spread into rosettes.

[MacGinitie « MacGinirtie] Plate 6

Tue VELIGER, Vol. 11, No. 1

Vol. 11; No. 1

of Cryptochiton the longer they are kept in aquaria. In
specimens that have been kept in tanks 3 or 4 months or
longer, we have found as many as 6 different organisms
among the spicules, although they were not attached to
the integument.

One of the functions of the spicules is to hold the sheet
of mucus in place. The invasion of the surface by foreign
material stimulates the animal to secrete and discharge
mucus and draw the spicules together into columns. How-
ever, most of the time, the tufts are spread in the form of
rosettes (Plate 6, Figure 3), which undoubtedly is a sort
of passive way of excluding larger larvae and tiny ani-
mals from its surface. There is no indication that a toxic
substance is secreted, for larvae that are introduced onto
the sheet of mucus do not respond as they would to a
toxic material. They continue to struggle for a much
longer time than they would if affected by a toxin. The
surface of the sheet of mucus gradually disintegrates and
the entrapped organic material is carried away by cur-
rents in the water.

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Page 61

ACKNOWLEDGMENTS

We are grateful to the personnel of the Friday Harbor
Laboratories of the University of Washington for several
courtesies. Through Dr. Paul Illg, we were able to use for
study and photographic purposes the excellent slides of
the integument (Plate 6, Figure 1) of Cryptochiton stel-
leri made by Deborah L. Christensen, microtechnician at
the University of Washington, Seattle. We also wish to
thank SCUBA divers Rich Bettger and Jerry Brown of
Everett, Washington, for obtaining specimens for us.

LITERATURE CITED

MacGrnitiE, GEoRGE EBER
1945. The size of the mesh openings in mucous feeding nets
of marine animals. Biol. Bull. 88 (2): 107-111
MacGinitie, Georce Esper & Nettie MaAcGInitie
1968. Natural history of marine animals. 2%? ed. xii+523
pp.; 286 text figs. McGraw-Hill Book Co., New York

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Vol. 11; No. 1

Further Observations on the West American Marginellidae

With the Descriptions of Two New Species

BY

BARRY ROTH

1217 Waller Street, San Francisco, California 94117

AND

EUGENE COAN

Department of Biological Sciences, Stanford University, Stanford, California 94305

(Plate 7; 2 Text figures; 1 Map)

IN oUR ACCOUNT oF the west American Marginellidae
(Coan & Rorn, 1966), we stated that with further work
on the family, especially the examination of more speci-
mens, our ideas would probably change. In addition, we
felt certain that the article would arouse interest among
workers, both to study the material they had on hand
and to collect additional specimens. Both anticipations
have proved correct. We have accumulated comments
adding to and correcting our published account. It seems
appropriate to make the new information available at
this time.

1. Marginella (Prunum) sapotilla Htnps, 1844
(Plate 7, Figures 1, 2)
Coan & Rortn, 1966: 279 - 280; plt. 48, figs. 1-3; text fig. 1

Photographs of the lectotype of Marginella evax Lt,
1930, are presented on the Plate. The synonymy of M.
evax and M. (P.) sapotilla remains certain. Because L1’s
photograph was poor and because his article is not readily
available, illustrations of this lectotype are included here.

In addition we are able to refine the distributional
information. The species has been found from Bahia
Honda, Panama, to Bella Vista, Panama, and as far off
the mainland as Isla Pedro Gonzales in the Bay of
Panama. Ecuadorian records remain uncertain. PARKER
(1964) reported it from the Gulf of California. His spe-
cimen is in the Museum of Comparative Zoology at
Harvard University and proves to be a young Olivella.

2. Marginella (Prunum) albuminosa Daur, 1919
(Plate 7, Figures 3, 4)
Coan & RotH, 1966: 281 - 282; plt. 48, fig. 11

Records and correspondence in the Division of Mol-
lusks at the United States National Museum indicate that
Dr. Alfred Dugés sent the unique specimen, along with
other material, to Dall in 1895, and that Dall initially
identified it as Marginella (Prunum) curta SOWERBY,
1832. There is some indication that Dugés’ shipment may
have been poorly packed and have arrived at the USNM
somewhat mixed. Thus there is additional reason to
doubt the occurrence of this species in the west American

fauna. Correspondence in the USNM files suggests that

Dugés may also have sent some of his material to the
Paris Museum, but a search there prompted by our in-
quiry turned up nothing. We have not yet located other
specimens of this species from any province in any muse-
um.

STANTON (1966) reported a specimen of Marginella
(Prunum) from the Castaic Formation (Upper Miocene)
of Los Angeles County, California, as “Marginella cf.
M. albuminosa Dau.” This is a new geologic record for
the family and genus in northwest America. STANTON’s
species will probably require a name of its own in light
of the continuing doubt about Dugés’‘type specimen.

Two better photographs of the holotype of M. ig nee
albuminosa are provided here.

Vol. 11; No. 1

3. Persicula porcellana (GMELIN, 1791)
(Plate 7, Figures 5, 6)

Coan & Roru, 1966: 282 - 283 (pars) ; plt. 48, figs. 12 - 15;
non ibid.: plt. 48, figs. 16-17

In our earlier article we designated one of the speci-
mens therein illustrated ( plt. 48, figs. 14, 15) “lecto-
type” of Marginella tessellata LaMaRcK, 1822 - a
mistake, since, as the only specimen in the Lamarckian
collection, it is assuredly the holotype. At the same
time we designated the identical specimen neotype of
GmeE.in’s Voluta porcellana — the specimen figured by
Cuemnirz (1788: plt. 150, figs. 1419, 1420), on which
GMELIN based his species, having been lost. We hoped to
stabilize nomenclature by this step, which we took after
consultation with several other malacologists. At the time
of that writing we had examined many west American
specimens, but no Caribbean ones. Now we have seen,
from the Caribbean, specimens that look like LAMarcK’s
type and still others which closely resemble the original
Cuemnirz figures. Some species of Persicula are mor-
phologically consistent; others show considerable varia-
tion even within local populations (Coan « Rot, 1966:
pp. 284 - 285). We suspect that there is only one, some-
what variable species in the Caribbean. In light of the
Caribbean specimens, there is much less doubt what form
Chemnitz had; in the absence of serious doubt, our
neotype designation seems superfluous. Since it helps
stabilize the nomenclature, however, and since its retrac-
tion could cause needless additional confusion, we intend
to let the designation stand.

Lamarck’s holotype of Marginella tessellata is illus-
trated here with two better photographs. It is without
locality data, but most likely came from the Caribbean,
considering the early date of Lamarck’s publication.
Voluta porcellana was incorrectly stated by Gmelin to
have come from the Indian Ocean.

While the Atlantic species retains the name Persicula
porcellana, the Pacific species, isolated from it since the
Pliocene by the Isthmus of Panama, requires a new name
as follows:

Persicula accola RotH & Coan, spec. nov.
(Plate 7, Figures 7,8 - Holotype)

Persicula porcellana (GMELIN), CoaAN &« RotH, 1966: 282 -
283 (pars); plt. 48, figs. 16-17; non Gmeuin, 1791:
3449 (species 139)

Description of Holotype: Shell of moderate size, solid;
elongate-ovate, narrower anteriorly; pale yellowish-tan,
with about 10 spiral rows of dark reddish-brown, more
or less rectangular blotches which show a tendency to-
ward doubling with maturity; entirely covered with a

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Page 63

thin glaze of translucent whitish enamel; outer lip thick-
ened, finely denticulate, white, tinged with brown along
outer edge; inner lip covered by a pad of white callus;
spire low, covered with clear enamel, circled by a solid
brown band; aperture even, slightly wider anteriorly,
white within; anterior canal deep, oblique; columella
with 7 folds, second fold from anterior end widest, most
anterior fold at base of columella.
Dimensions of Holotype: Length 13 mm; width 8.2 mm.
Paratypes: Of the 3 paratypes, 2 are worn mature
specimens and 1 is a live-collected, sub-mature speci-
men with a sharp, uncalloused outer lip. All 3 differ very
slightly from the holotype in arrangement of the rows of
brown blotches.
Dimensions of Paratypes:

1: Length 12.5 mm; width 7.5 mm (live)

2: Length 13.3 mm; width 8.2 mm

3: Length 12.2 mm; width 7.2 mm
Type Locality: Isla Coiba, Panama (about 7°30’ N by
81°45’ W) ; collected by A. Mendez.
Discussion: The pattern of rows of rectangular blotches
distinguishes Persicula accola from all other west Ameri-
can species of the genus. A key to the west American
species (in which this species is identified as “Persicula
porcellana’) appears in Coan & Rotu, 1966, p. 278.

This species is morphologically very similar to its Carib-
bean analogue, Persicula porcellana (GmeEun). Although
museums contain only a few lots of each species from
which to make comparisons, some minor differences have
been noted. First, the west American species is broadest
much posterior to the middle of the body whorl, while
the Caribbean one tends to be more nearly ellipsoidal.
Persicula porcellana tends also to be more obese. Second,
as in the Chemnitz figures mentioned above, P. porcellana
may have smaller spots, especially in the northern part
of its range — Honduras, Panama, Venezuela. Brazilian
specimens have larger spots. Persicula accola has, in gen-
eral, larger, squarer spots. Third, the posterior end of
the outer lip of the Caribbean species tends to be pro-
duced slightly more than that of the west American
form.
Etymology: The specific name derives from the Latin
noun for “neighbor.”
Material Examined and Range: This species has evi-
dently been collected only 4 times, and 2 of these lots
have been divided among several museums. The recorded
localities are Isla Coiba, Isla Jicaron, and Bahia Montijo,
all in roughly the same area of Panama.
Deposition of Types: Holotype: USNM 513647; Para-
types (3): USNM 665526.

4. Persicula bandera CoaAN & Rotu, 1965

Coan & Rotn, 1965: 67 - 69; pit. 12, figs. 1-5
Coan & Ror, 1966: 285; plt. 50, figs. 38, 39

Page 64

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Vol. 11; No. 1

When first describing Persicula bandera, we compared
it to the closely related P hilli (SmitH, 1950), but not
to any species from the Atlantic. We have recently ob-
tained a photograph of the holotype of Marginella
multilineata SowerBy, 1846 from the British Museum
(Natural History) (Plate 7, Figures 9, 10); that species
is a Persicula, very similar to P. bandera.

Marginella multilineata was described from a speci-
men in the Cuming collection, picked up by a Mr.
Dyson at “Belieze [sic], Bay of Honduras.” Belize, Brit-
ish Honduras, and the Gulf of Honduras are on the
Atlantic coast of Central America; and other Dyson-
collected material from “Honduras” has proven to be
Atlantic, so there is no reason to doubt the Caribbean
locality. REEvE (1865) compared M. multilineata to M.
interrupta LAMARCK, 1822 (= Persicula interruptoline-
ata [MEGERLE von MUHLFELD, 1816]), a variable Carib-
bean and west African species; and Tomutn (1917) in-
correctly synonymized the two.

It seems plausible that, as with so many other mar-
ginellids, analogous Atlantic and Pacific species are in-
volved. If it ever were to be shown conclusively that
Dyson’s shell came from the Pacific side of Central
America, the name Persicula bandera would become a
synonym of PR multilineata (SowERBy, 1846).

5. Persicula hilli (Smitu, 1950)
(Plate 7, Figures 11, 12)
Coan & Rotu, 1966: 285; plt. 50, figs. 40, 41

At the time of our earlier review, we were unable to
locate the type specimens of this species. Through the

Explanation

Figures 1, 2: Marginella (Prunum) sapotilla Hinps, 1844. Lecto-
type of Marginella evax Li, Columbia Univ. Paleo. Coll. 22118;
Panama Bay, 2.1 (ventral and dorsal views)

Figures 3, 4: Marginella (Prunum) albuminosa Dat, 1919.
Holotype, USNM 10168, “West Mexico,” 1.5 (ventral and
dorsal views)

Figures 5, 6: Persicula porcellana (GMELIN, 1791). Neotype of
Voluta porcellana Gmeuin and holotype of Marginella tessellata
Lamarck, Mus. Hist. Nat. Geneva, no locality given; < 2.1 (vent-
ral and dorsal views). Photographs courtesy of Dr. E. Binder
Figures 7, 8: Persicula accola RoruH & Coan, spec. nov. Holotype,
USNM 513647, Isla Coiba, Panama, X 3 (ventral and dorsal views)
Figures 9, 10: Persicula multilineata (SowERBy, 1846). Holotype,
British Museum (Natural History), Cuming Coll., “Belieze, Bay
of Honduras,” X 3 (ventral and dorsal views). Photographs cour-
tesy of and © by British Museum (Natural History)

Figures 11, 12: Persicula hilli (Smitu, 1950). Lectotype, Univ.
Alabama, Maxwell Smith Coll. 15374, Bahia Chamela, Jalisco,
Mexico, X 2.6 (ventral and dorsal views)

courtesy of Dr. Herbert T: Boschung, we have been
loaned the type lot from the Maxwell Smith collection in
the Museum of Natural History at the University of
Alabama at Tuscaloosa. The type lot consists of 4 speci-
mens. SmiTH (1950: plt. 4, fig. 6) figured 2 of these. We
hereby designate as lectotype the specimen we have
illustrated here — the specimen which SmirH figured in
ventral view — leaving 3 paralectotypes. The lectotype
measures 14.7mm in length and 9.7mm in width.
Smitu’s figured paralectotype is 14.5mm long and 9.2
mm wide.

6. Volvarina sp., cf. V. taeniolata MOrcu, 1860

Among uncatalogued material at Stanford University,
we have seen one immature specimen belonging to the
genus Volvarina, collected at Salinas, Ecuador, in 1951
by Dr. Donald L. Frizzell. This specimen extends the
known west American range of the genus to South
America. Adult specimens will have to be studied to
determine whether this is Morch’s species.

7. Cystiscus politulus (Dati, 1919)
(Plate 7, Figures 13, 14; Map)

Coan & Rotu, 1966: 290-291; plt. 51, fig. 64

Hyalina myrmecoon Datu, 1919: 308

Cystiscus myrmecoon (Dati), Coan & Roru, 1966: 291; plt.
51, fig. 65

Examination of additional material now leads us to
synonymize these two species. Since both of Datt’s
names date from the same article, we are acting as “first

of Plate 7

Figures 13, 14: Cystiscus politulus (Dat, 1919). Hypotype, US
NM 268953, Bahia Magdalena, Baja California Sur, Mexico, 25m,
<9 (ventral and dorsal views)

Figure 15: Cystiscus jewettu (CARPENTER, 1857). Hypotype, SU
PTC 9943, Punta Abreojos, Baja California Sur, Mexico, x 8.8
(ventral view)

Figure 16: Cystiscus jewettu. Hypotype, CASGTC 13107, Point
Pinos, Monterey County, California, X 8.8 (ventral view)

Figures 17, 18: Cystiscus palantirulus RoTH & COoAN, spec. nov.
Holotype, AMNH 128732, Los Frailes Bay, Baja California Sur,
Mexico, 37- 73m, X 8.8 (ventral and dorsal views)

Figure 19: Kogomea polita (CARPENTER, 1857). Lectotype, Brit-
ish Museum (Natural History) 57.6.4.2108/1, Mazatlan, Sinaloa,
Mexico, X 35 (ventral view). Photograph courtesy of and © by
British Museum (Natural History)

Figure 20: Cypraeolina margaritula (CARPENTER, 1857). Lecto-
type, British Museum (Natural History) 57.6.4.2109/1, Mazatlan,
Sinaloa, Mexico, X 16.8 (ventral view). Photograph courtesy of
and © by British Museum (Natural History)

TuHeE VELIGER, Vol. 11, No. 1 [RotH &« Coan] Plate 7

Figure 3 Figure 4 Figure 5

Figure 9

Figure 11 Figure 12 Figure 13 Figure 14 Figure 15

Figure 16 Figure 17 Figure 18 Figure 19 Figure 20

Vol. 11; No. 1 THE VELIGER Page 65

LEGEND
Rangeof Cysts plus: — ———E—————_——>
Range of Cystiscus jewettii: —a_=» aa=> a=

Range of Cystiscus palantirulu’: —e a= @

Isla
Guadalupe

MEXICO

Punta Isla

Pacific Ocean Be Poe
“aN Monserrate’: ”

Ranges of the West American Species of Cystiscus

Page 66

revisers” in the sense of Article 24a of the International
Code of Zoological Nomenclature. The holotype of Hya-
lina myrmecoon is simply an elongate specimen of C’ystzs-
cus politulus.

Additional range records include the southern Gulf of
California (Map). The Los Angeles County Museum of
Natural History collection includes specimens recently
collected by Dr. James H. McLean at Cabo San Lucas,
Pulmo Reef, and Isla Cerralvo, Baja California Sur. We
have tentatively determined as Cystiscus politulus a poor
specimen from Bahia de las Banderas, mainland Mexico,
so collectors should keep watch for the species in that
area as well.

A specimen from Bahia Magdalena on the outer coast
of Baja California Sur is here illustrated and discussed
in connection with the description of a new species.

8. Cystiscus jewettti (CARPENTER, 1857a)
(Plate 7, Figures 15, 16; Text figure 1; Map)
Coan & Rortn, 1966: 291; plt. 51, figs. 66 - 68

From its previously reported distribution (Monterey,
California, to Isla San Martin and Isla Guadalupe, Baja
California) we can now extend the known range south-
ward to Punta Abreojos, Baja California Sur. Note that
this is still considerably north of the range of the next
species. We are illustrating a specimen from Punta Abre-
ojos and one from the northern end of the range, Point
Pinos, Monterey County, California, for comparison with
the new species described below.

Cystiscus jewetti is common in the intertidal area at
Pacific Grove, Monterey County, California, where speci-
mens were collected and observed in August, 1967 (Text
figure 1). Throughout the period of observation, the

THE VELIGER

Vol. 11; No. 1

mantle was never extruded over the top of the shell of
C. jewettii in the manner of Cypraeolina margaritula in
the same dish.

9. Cystiscus palantirulus RoTH & CoAN, spec. nov.
(Plate 7, Figures 17, 18 [Holotype]; Map)

Cystiscus sp., Coan & Roru, 1966: 291 - 292; plt. 51, figs.
69 - 70

We have now seen a sufficient number of specimens of
this form to propose a name for it.
Description of Holotype: Shell pear-shaped, broad and
evenly rounded posteriorly, elongate and narrow anteri-
orly; white, smooth, highly polished; spire very low,
covered by irregular callousing; outer lip extending high
on body whorl, forming a 90° arc slightly anterior to the
spire, most strongly thickened by callus posteriorly; a-
pertural margin (columellar area) forming a long S-curve,
concave anteriorly, very bulbous posteriorly, scarcely
thickened by callus, with 2 large folds at base of colu-
mella and 4 smaller ones posterior to them, evenly
decreasing in size, extending very slightly onto face of
body whorl; aperture moderately wide anteriorly; ante-
rior margin of aperture rounded; anterior end slightly
flaring and twisted toward columella.
Dimensions of Holotype: Length 3.5 mm; width 2.1 mm.
Dimensions of Paratypes:

1. Length 3.1 mm; width 1.7 mm

2: Length 3.2 mm; width 1.7 mm
Paratypes: The two paratypes differ from the holotype in
the following details: the apex of Paratype 1 projects as
a minute nubbin; the apex of Paratype 2 is worn to a
smooth dome; in curving to meet the spire, the outer
lip of Paratype 1 changes direction by less than 90°.

Figure 1

Cystiscus jewettii (CARPENTER, 1857)

Living specimen, Pacific Grove, Monterey County, California.

Intertidal

xX 14

Vol. 11; No. 1

THE VELIGER

Page 67

Type Locality: Station 89 of the Puritan-American Mu-
seum of Natural History Expedition to West Mexico: Los
Frailes Bay, Baja California Sur (Gulf of California),
23°21’ N by 109°25’ W, 20-40 fathoms, fine sand, 19
April 1957, taken with the Puritan dredge.

Deposition of Types: Mollusk collection, American Mu-
seum of Natural History, New York, No. 128732 (Holo-
type) and AMNH No. 77942 (Paratypes).

Range: Specimens of C'ystiscus palantirulus in other mu-
seum and private collections indicate a range limited to
the southwestern part of the Gulf of California, from
off Isla Monserrate to Cabo San Lucas, Baja California
Sur (see Map), with one possible specimen from Aca-
pulco, Guerrero. Collection records range from 6 to 80m
depth, plus 2 shore-collected dead shells. In addition to
the type lot, we have examined the following material:

Los Angeles County Museum of Natural History:

Invert. Zool. Loc. 66-12, Cabo San Lucas, diving in 6 - 24m
near the pinnacle - 3 specimens

Invert. Zool. Loc. 66-17, dredged in 18m between El Tule
and Punta Palmilla, Baja California Sur - 35 specimens

Invert. Zool. Loc. 66-23, Between Punta Ventana and Isla
Cerralvo, Gulf of California, 18- 27m - 12 specimens

California Academy of Sciences:
No. 24062, Bahia San Lucas - 2 specimens

San Diego Museum of Natural History:
No. 33654, Isla Espiritu Santo, Gulf of California - 1 speci-

men, juvenile

Locality L-2155, Isla Espiritu Santo - 3 specimens

Stanford University Paleontological Type Collection:
No. 9849, 10 miles north of Isla Espiritu Santo - one speci-
men (figured by Coan « Rorn, 1966)
Collection of Dr. Donald R. Shasky, Redlands, California:
37 - 73m off Isla Monserrate, Gulf of California

Collection of Dr.S. Stillman Berry, Redlands, California:

No. 10074, Acapulco, Guerrero - one specimen cf. C. pa-
lantirulus, but not recently re-examined

Discussion and Comparisons: A comparison of Cystiscus
palantirulus with the other 2 west American species of
Cystiscus is presented in Table 1.

Neither Cystiscus jewettii nor C. palantirulus has yet
been taken in the area between Punta Abreojos and Cabo
San Lucas (see Map). This apparent gap of about 400
coastal miles between the ranges supports the notion of
C. palantirulus as a specifically distinct population. In
addition, although C. jewettii exhibits consistent differ-
ences in form between the northern and the southern
ends of its range (Plate 7, Figures 15, 16; and Table 1),
and, although there is some individual variation between
specimens from a single locality, there is no tendency for
its populations to take on the distinctive shape of C.
palantirulus.

In areas where their ranges overlap, Cystiscus politulus
(Plate 7, Figures 13, 14) can be distinguished from C.
jewettu or C. palantirulus by its narrower, more elongate
aspect, and by its generally smaller size.

Etymology: The palantiri, from which the specific name
is derived, are magical globes of crystal mentioned in
the fiction of J. R. R. Tolkien.

Table 1

Cystiscus politulus

Size: Small for genus; 2.9mm (average of Medium to large for genus; 4.8mm
(average of 2 northern specimens)

10 from 6 localities)

Cystiscus jewetti

Cystiscus palantirulus

Medium for genus 3.2mm (average
of 27 from 9 localities)

3.1mm (average of 4 from Pta.

Abreojos)
Shape: Elongate; not shouldered; length to
width ratio 1.8 (average of 10
from 6 localities)
Texture:

Callousing: Thin

Spire: Low to slightly elevated; not callus-
covered
Outer Lip: Slightly thickened at maturity
Aperture: Anterior end rounded, even
Columella:
Range:
(see Map) Gulf of California; intertidal to

60m 50m

Ovate; shouldered; length/width ra-
tio 1.5 (average of 6 from 3 local.)

Much thickened at maturity

Anterior end effuse, even

Pear-shaped; shouldered; length/width
ratio 1.6 (average of 27 from 9
localities)

All species shiny, translucent
Often thick, esp. on northern specim. Thin

Low to well elevated (in south. spec.) ; Low; sometimes mamillate
northern specimens callus-covered

Slightly thickened

Anterior end effuse, twisted to left in
dorsal view

Much the same in all 3 species; also variable from specimen to specimen

Santa Barbara, California, to southern Monterey, California, to Pta. Abreojos, Isla Monserrate to Cabo San Lucas,
Baja California Sur; intertidal to

Baja California Sur; 6 to 80m

Page 68

THE VELIGER

Vol. 11; No. 1

10. Kogomea polita (CARPENTER, 1857b)
(Plate 7, Figure 19)’
Coan & Roru, 1966: 293; plt. 51, fig. 75; text fig. 3

We are illustrating the previously designated lectotype
of this species by a photograph for the first time.

11. Cypraeolina margaritula (CARPENTER, 1857b)
(Plate 7, Figure 20; Text figure 2)
Coan & Rotu, 1966: 294 - 295; plt. 51, fig. 77; text figs. 4, 5

A photograph of the lectotype is provided.

One of a number of specimens collected in August,
1967, at Pacific Grove, Monterey County, California,
where the species is abundant intertidally, is illustrated
(Text figure 2). Its mantle was black, with red patches
and minute spots of pale blue, and a narrow white mar-
gin. The rest of the animal was translucent tan, closely
spotted with pale lemon-yellow.

Figure 2

Cypraeolina margaritula (CARPENTER, 1857)

Living specimen, Pacific Grove, Monterey County, California.
Intertidal X 14

ACKNOWLEDGMENTS

The authors wish to thank the following persons for
providing data and photographs, for offering recommend-
ations, and for permitting the loan of specimens used in
this study: Dr. S. Stillman Berry, Dr. Eugéne Binder, Dr.
Herbert T: Boschung, Dr. Kenneth J. Boss, Mr. Robert C.
Bullock, Mr. S. Peter Dance, Dr. William K. Emerson,
Dr. Leo G. Hertlein, Dr. John Imbrie, Sister Marion
Johnson, Dr. A. Myra Keen, Dr. Jorgen Knudsen, Dr.
James H. McLean, Dr. Joseph Rosewater, Dr. Donald R.
Shasky, Dr. Norman Tebble, Dr. Edward C. Wilson.

LITERATURE CITED

CarPENTER, Puitip PEARSALL
1857a. see GouLD & CARPENTER, 1857
1857b. Catalogue of the collection of Mazatlan shells in the
British Museum collected by Frederick Reigen ... London
(British Museum) pp. i-iv + ix-xvi + 1-552 (June 1857)

CuHEmnitz, JoHann HiEROoNyMUS
1788. | Neues systematisches Conchylien Cabinet 10.
Niirnberg (G. N. Raspe) pp. [1-22] + 1-376; plts. 137 - 173

Coan, Eucene Victor & Barry RoTH
1965. A new species of Persicula from West Mexico. The
Veliger 8 (2) : 67 - 69; plt. 12; 1 map (1 October 1965)

1966. |The West American Marginellidae. The Veliger 8 (4) :
276 - 299; plts. 48-51; 5 text figs. (1 April 1966)

Dai, WiLt1AM HEALEY
1919. Descriptions of new species of Mollusca from the North
Pacific Ocean in the collection of the United States National
Museum. Proc. U.S. Nat. Mus. 56 (2295): 293 - 371
(30 August 1919)

GMELIN, JoHANN FREDERICH
1791. Systema naturae per regna tria naturae
decima tertia, aucta, reformata 1 (6): 3021 - 3910.

Goutp, Aucustus Appison & Puitip PEARSALL CARPENTER
1857. _ Descriptions of shells from the Gulf of California, and
the Pacific coasts of Mexico and California. Part 2. Proc.
Zool. Soc. London 26 [for 1856] [part 24] (313): 198-208
(7 January 1857)

Editio
Lipsia

Hinps, Ricuarp BrinsLey
1844. Descriptions of Marginellae collected during the voyage
of H. M.S. Sulphur, and from the collections of Mr. Cuming.
Proc. Zool. Soc. London 14 (134) [for 1844]: 72-77
(September 1844)

LaMaRcK, JEAN-BaptisTE PrerrRE ANTOINE DE MONET DE
1822. Histoire naturelle des animaux sans vertébres, 7 [Mol-
lusques]. Paris (“chez l’auteur, au jardin du Roi”) pp. 1-711
(August 1822)
Li, Co1n CHanc
1930. The Miocene and Recent Mollusca of Panama Bay.
Bull. Geol. Soc. China 9 (3): 249-296; plts. 1-8; 1 map
(October 1930)

Vol. 11; No. 1 THE VELIGER Page 69

MEGERLE von MUuLrFELD, JOHANN Karu
1816. Beschreibung einiger neuen Conchylien. Gesellsch.
naturforsch. Freunde Berlin: Mag. f. neuest. Entdeck. i. d.
ges. Natupk. 8 (1) [article 1]: 3-11; plts. 1-2

Morcu, Otro AnprEas Lowson
1860. Beitrage zur Molluskenfauna Central-Amerikas.
Malak. Blatter 7: 66-96 (July 1860); 97- 106 (Aug. 1860)

Parker, Rosert H.

1964. | Zoogeography and ecology of some macro-invertebrates,
particularly molluscs, in the Gulf of California and the conti-
nental slope off Mexico. Vidensk. Medd. Dansk naturh.
Foren 126: 1 - 178; 15 plts.; 29 text figs.; 7 tables

REEvE, Lovett Aucustus
1864-1865. Conchologia iconica: or illustrations of molluscous
animals 15 (Monograph of the genus Marginella) : plts. 1 - 27
plts.2-13 —- August 1864
pits. 1,14-27 + Index and Errata — January 1865
SmitH, MAaxwELu
1950. | New Mexican and Panamic shells. Nautilus 64 (2):

60-61; plt. 4 (27 October 1950)
SoweErsy, GrorcE BrRETTINGHAM (157 of name)

1832. | Characters of new species of Mollusca and Conchifera
collected by Mr. Cuming. Proc. Zool. Soc. London 2 [for
1832] [part 2] (17): 50-61 (5 June 1832) and
(18): 104-108 (31 July 1832)

SoweErsBy, GeorcE BRETTINGHAM (2%? of name)
1846. Descriptions of new species of Marginella. Proc.

Zool. Soc. London 16 [for 1846] [part 14] (164): 95-97
(November 1846)
Stanton, Roser J., Jr.
1966. Megafauna of the Upper Miocene Castaic Formation,

Los Angeles County, California. Journ. Paleont. 40 (1):

21-40; plts. 5-7; 2 text figs. (January 1966)
Tomuin, JoHN ReaD LE BRocKTON

1917. A systematic list of the Marginellidae. Proc.

Malacol. Soc. London 12 (5) : 242 - 306 (22 August 1917)

Page 70

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Vol. 11; No. 1

On an Octopod from Placentia Bay, Newfoundland

BY

v

FREDERICK A. ALDRICH

AND

Cc. GC. LU

Marine Sciences Research Laboratory

Memorial University of Newfoundland, St. John’s, Newfoundland

(Plate 8)

IN CONTRAST TO THE ABUNDANCE of squid, octopods
appear to be rare in Newfoundland waters. On October
28, 1967, a local fisherman obtained an octopod from
water 60 fathoms deep, one mile west from Grass Island,
Placentia Bay. Due to the rarity of octopods in this area,
he brought the specimen to the Marine Sciences Research
Laboratory on October 30, where it was turned over to
the authors for study. Due to lack of proper treatment,
the specimen was damaged and its surface somewhat
peeled.

The animal is identified as Benthoctopus piscatorum
(VERRILL, 1879) (Plate 8, Figure 1). The body is globu-
lar; its surface smooth and purplish brown in color. Arms
are long and slender, tapering to the tips, and suckers are
small. The measurements and indices are given in
iablemls

Benthoctopus piscatorum was first described by VER-
RILL in 1879 as Octopus piscatorum. The present genus
was established by Grimpe in 1921, with B. piscatorum as
the type species. Specimens from fairly wide distributions,
both vertical and horizontal, have been reported, i. e., Le
Have Bank, Nova Scotia in 120 fathoms; Grand Bank in
200 fathoms; 39° 26’N, 70°02’ W, in 1362 fathoms
(VERRILL, 1880 - 1881) ; 78° 02’ N, 9° 25’E, in 416 fath-
oms (ApPELLOF, 1893); 66°41’ N, 6°59’E, in 350

fathoms (AppELLOF, 1893) ; 60° 40’ N, 4° 50’ W, in 563
fathoms (RussELL, 1909); 61° 27’ N, 1° 47’ W, in 681
fathoms (British Museum [quoted from Rosson, 1932]) ;
51° 15’ N, 11° 47’ W, in 707 - 710 fathoms (Massy, 1907);
Faroe Channel in 540-608 fathoms (Hoyts, 1886).
Muus (1962) postulated that this species “might be
expected” in the waters of Davis Strait west of Green-
land. It is the first time that this species has been reported
from the inshore waters of Newfoundland.

The specimen is a female, with a very large ovary,
measuring 35 by 42 mm (Plate 8, Figure 2). The ovarian
egg measures 18 by 6 mm. The spermathecae are globular,
black and large, measuring 18 by 10mm The values
quoted in Table 1 are considerably different from those
presented by Rosson (1932) for Benthoctopus piscator-
um. The discrepancies are undoubtedly due to the preser-
vation of our specimen.

The rarity of the octopod in Newfoundland waters
may only be apparent and further collection by use of
proper gear would increase our knowledge of the local
octopodan fauna.

We wish to express our appreciation to Dr. G. L. Voss
of the Institute of Marine Sciences of the University of
Miami for sending us specimens from his collections for
purpose of comparison. —

Explanation of Plate 8

Figure 1:
Figure 2:

Benthoctopus piscatorum (VERRILL, 1879), dorsal view of specimen from Placentia Bay, Newfoundland.

Eggs (in situ) of described specimen of Benthoctopus piscatorum (VERRILL, 1879).

EPR avions Vol an NO. i [ALpRicH & Lu] Plate 8
HE VELIGER, Vol. 11, No.

SINddNS AYOLVHORV) NvIaYNYD

Figure 1

Figure 2

Vol. 11; No. 1

THE VELIGER Page 71

Body proportions and indices of the examined specimen
of Benthoctopus piscatorum (VERRILL, 1879)

Table 1

(following Voss, 1956 and 1963).

Body Proportions (length in mm)

Web Depth:

Number of gills:

Indices (%)

TL,
ML
VML
MW
HW

AUP

MWI
HWI
MAL
WDI
ALI
SI
SLI
SWI

Left 10, Right 8

362
89
58
45
31
25
12

260

203

212

179

56
52
62
45
26

50.68
34.57
34.15
23.85
71.82
5.4
28.15
12.84

LITERATURE CITED

APPELLOF, A.

1893. Teuthologische Beitrage III. Bergens Mus. Aarb.

No. 1
GrimpPeE, GEORG

1921. Teuthologische Mitteilungen VII. Systematische Uber-

sicht der Nordseecephalopoden. Zool. Anz. 52: 296 - 304
Hoye, WiLii1AM E.

1886. Report on the Cephalopoda collected by H.M.S.
Challenger during the years 1873 - 76. Rept. Sci. Res.
Voy. Challenger 1873-76. Zool. 16 (44): 1-245; 33 plts.;
10 figs. in text

Massy, ANNE L.

1907. Preliminary notices of new and remarkable cephalo-
pods from the S. W. coast of Ireland. Ann. Mag. Nat. Hist.
(Ser. 7) 20: 377 - 384

Muus, B. J.

1962. Cephalopoda. Meddelelser om Gronland 81 (5):

1 - 23
Rosson, Guy Cosurn

1932. A monograph of the Recent Cephalopoda. Part 2:
The Octopoda, excluding the Octopodinae. Brit. Mus.
2: 1-359; 6 plts.; 79 text figs.

RussELL, E. S.

1909. _—‘ Preliminary notice of the Cephalopoda collected by the
fishery cruiser Goldseeker, 1903 - 1908. Ann. Mag. Nat.
Hist. (Ser. 8) 3: 446 - 453

VERRILL, ADDISON E.

1879. Notice of recent additions to the marine fauna of the
eastern coast of North America, No. III. Amer. Journ.
Sci. (Ser. 3) 17: 239 - 243

1880-1881. The cephalopods of the northeastern coast of
America. Part II. The smaller cephalopods, including the
“squid” and the octopi, with other allied forms. Trans.
Connecticut Acad. Sci. 5: 259-446; 2 plts.

Voss, GiLBerT L.

1956. A review of the cephalopods of the Gulf of Mexico.
Bull. Mar. Sci. Gulf and Carib. 6 (2): 85-178; 18 figs.

1963. | Cephalopods of the Philippine Islands. U.S. Nat.
Mus. Bull. 234: 1-180; 4 plts.; 36 text figs.

Page 72

THE VELIGER

Vol. 11; No. 1:

A New Species of Enoploteuthid Squid,

Abraliopsis (Watasenia) felis, from the California Current

JOHN A. McGOWAN

Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California 92037

TAKASHI OKUTANI

Tokai Regional Fisheries Research Laboratory, Tokyo

(Plates 9 and 10; 1 Map)

INTRODUCTION

IN AN EXAMINATION of the extensive collection of pelagic
cephalopods at the Scripps Institution of Oceanography
a number of specimens of apparently undescribed spe-
cies was found. One group of these specimens was so
numerous and the series so complete that we have
decided to describe it separately from the rest of the
collection. A further reason for describing this species
now is that it was discovered in a study of the distribu-
tion and abundance of the larval squid of the California
Current that this species ranked first in abundance
(OxuTani & McGowan, in press). Including larvae,
juveniles and adults, we have examined about 1850 in-
dividuals of this new species.

ACKNOWLEDGMENTS

The specimens reported on here were collected by many
individuals of the staff of the Scripps Institution of
Oceanography. Twenty specimens were obtained from
Dr. William Pearcy of Oregon State University. Mr.
Charles B. Miller made many of the measurements and
Mr. Kenneth H. Isaacs sorted many of the specimens.
This work was supported by the Marine Life Research
Program, the Scripps Institution of Oceanography’s part
of the California Cooperative Oceanic Fisheries Investi-
gations, which are sponsored by the Marine Research
Committee of the State of California and by the Nation-
al Science Foundation Grant GB 2861.

Abraliopsis (Watasenia) felis McGowan & OXKUTANI,
spec. nov.

(Plates 9 and 10)

Synonyms: None.

Diagnosis: An Abraliopsis with a double row of tenta-
cular hooks, the ventral row of which consists of 3 or
occasionally 4 large hooks and the dorsal row of 3 or
occasionally 4 small hooks. The fixing apparatus on the
carpus is compact and consists of 4 suckers and 7 knobs.
The right ventral arm and occasionally both the right
and left ventral arm of the male is hectocotylized. The
hectocotylus consists of two semilunar membranes on the
lateral edges of the arm just proximal to the 3 large
photophores of the arm tip. These membranes are offset
from one another, the inner one being more proximal
than the outer. The ventral surface of the mantle with
many small photophores arranged randomly but with a
narrow median bare area running the entire length of
the mantle. The body is elongate and fusiform with the
width being less than 4, but more than 4 the length.

Description: The body is muscular and firm. The mantle
shape is fusiform and somewhat elongated. Based on
measurements of 94 adult specimens, the width is less
than + of the dorsal mantle length but more than + of
the dorsal mantle length. The broadest part of the mantle
is at the free margin which flares out slightly, The dorsal
margin of the mantle is triangularly lobed and projects
slightly at the mid-point. On the ventral side there are

THE VELIGER, Vol. 11, No. 1 [McGowan « Oxutani] Plate g

Abraliopsis (Watasenia) felis McGowaNn & OKUTANI, 0. sp.

Holotype, Male

De GURL hE | Sie DEA OR a ne a
cnt Seer mal yep lb ai i dam ey engi merge ym—me lis ws arcs tna mmc ele te eco

“ iB f ¥

5 ;

mye

Vol. 11; No. 1

THE VELIGER

Page 73

slight angular projections at the position of both funnel
cartilages. Between these two projections the margin is
slightly concave (Plate 9).

The fins are broadly sagittate with the posterior margin
being slightly concave and the anterior margin somewhat
convex. The width of both fins, taken together, is about
86% of the mantle length (22 adult specimens measured,
see Table 1). The free margins of the fins are thinner
than the remaining parts.

The head is rather large and is as wide as the mantle
and approximately 4 as long as the mantle length. The
eye opening has a distinct sinus on the anterior edge at
about the position of the third arm. There are 3 nuchal
folds on either side of the head, the ventral 2 of which
are semi-lunar in shape and the dorsal one somewhat
squared and elongated (Plate 10, Figure 1). The central
fold bears an “olfactory organ. ”

The neck is constricted and distinct and has a shallow
funnel excavation. The funnel is conical and short. The
funnel organs consist of a dorsal V-shaped pad and two
ventral oblong pads (Plate 10, Figure 3). There is a
transverse fold anterior to the dorsal funnel organ. The
funnel cartilages are about 3 times as long as wide and

have a slightly curved but simple groove (Plate 10, Fig-
ure 1). The nuchal cartilage is straight, spatulate and bi-
laterally symmetrical. It is widened a little on both ends.
The arms are nearly equal in length with an arm
formula 4,3, 2,1 or 4, 2,3,1. The dorsal arms have a
low membranous keel on the distal +. The oral face of the
dorsal arms has from 12 to 17 hooks arranged in a
zigzag row (Table 1, based on 12 specimens). Distal to
the hooks are 10 to 18 suckers, also alternating in a zigzag
row. The outer margin of the hook-bearing face of these
arms has a low swimming membrane supported by nu-
merous muscular filaments. The second arms are similar
to the dorsals except that they tend to have a slightly
greater number of hooks. The third arms have well-de-
veloped aboral keels extending along the entire length of
each arm. The armature is similar to that of the second
but with fewer distal suckers and somewhat fewer hooks
(Plate 10, Figure 7). The ventral arms have aboral keels
their entire length. They have from 7 to 14 hooks in a
zigzag row but no distal suckers. Generally the right vent-
ral arm is hectocotylized but in an examination of 39
males, 2 of them had both the right and the left ventral
arms hectocotylized. The hectocotylus is a pair of semi-

Table 1

Measurements of Types

Holotype

Total length (including tentacle) 84.8

mm

Mantle length, mm 36.7

Mantle width, mm 10.6

Fin length at attachment, mm 20.5

Width across the fins, mm 26.2

Width of the head (inter-ocular 10.6
length), mm

Head length (from anterior end 12.7
of nuchal cartilage to the prox-
imal end of the dorsal arms),
mm

Eye (transverse length X longi- 5.5 X 5.5
tudinal length of eye opening),
mm
Dorsal arm: Right 18.9
length, mm Left 18.1
Second arm: Right 20.9
length, mm Left 20.2
Third arm: Right 19.2
length, mm Left 19.3
Ventral arm: Right 21.5
length, mm Left 21.8
Tentacle: Right 37.0

length, mm Left 32.2

Paratype 1 Paratype 2 Paratype 3
(= Allotype)

105.5 - 80.0
417 33.5 39.2
12.4 10.2 11.8
25.0 16.0 21.6
33.8 26.1 29.8
11.1 11.5 9.5
11.0 10.1 12.0

5.9 X 4.5 4.1 X 3.3 4.3 X 3.5
17.8 12.5 18.0
18.5 13.0 16.0
21.8 15.8 21.0
21.8 14.2 20.2
22.8 16.6 =
22.4 15.0 -
27.0 - 24.0
27.0 - 21.0
42.7 - 345
41.0 - =

Page 74

lunar membranes offset from one another with the inner
membrane being somewhat more distal than the outer
(Plate 10, Figures 6, 6a, 6b). Between these crests is
a deep groove.

The tentacle is as long as the mantle and has a gradu-
ally tapered stem which is flattened on the oral surface.
The club (or manus) is not expanded and occupies */«
to */; of the tentacle length. There is no membrane on
the outer side of the club. The armature of the proximal
portion of the club consists primarily of hooks arranged
in two rows. In the ventral row there are 3 or occasionally
4 large hooks and muscular pads are sometimes present
between these. The dorsal row consists of 3 and occasion-
ally 4 small hooks which are usually about half the size
of the large hooks (Plate 10, Figures 5a and 5b). There
are very small suckers and sometimes pads between these
hooks. The distal part of the club has 13 transverse rows
of 4 suckers each. The horny rings of these suckers have
5 to 6 rather blunt teeth on the margin (Plate 10, Fig-
ure 5a). The carpus bears a fixing apparatus which con-
sists of 4 suckers arranged in a quadrate and 6 to 7
small papillae or knobs (Plate 10, Figure 5).

The mantle has many photophores, particularly on its
ventral surface. These range in size from 150 to 250u.
There is an almost random arrangement of these, but
a narrow longitudinal strip without photophores exists
on the mid-ventrum. The funnel has about 60 photo-
phores, particularly on the ventral side, but also on the
dorsum. The ventral surface of the head also has photo-
phores on its integument, but these are arranged in no
consistent order except about 28 surrounding the peri-
phery of the eyes (Plate 9). The eyeball has 2 large
(600) and 3 small (400) photophores along the vent
ral periphery. The third arms have a row of 9 to 11
photophores on the ventral surface. The ventral arms
bear 3 rows of photophores of which the dorsal-most
consists of 5 small ones on the keel. The other 2 rows are
along both edges of the aboral surface (Plate 9). Very
near the distal tip of the ventral arms are 3 very large
ellipsoid luminous organs about 0.8mm in length (Plate

THE VELIGER

Vol. 11; No. 1

10, Figure 6). Distal to these are 4 small photophores.
These large luminous organs are a characteristic feature
of the genus.

The buccal membrane has 8 supporting muscles and
its inner surface is heavily papillated (Plate 10, Figure 2).

The radula is quite delicate and has 7 unicuspid rha-
chidian teeth in a row and no accessory plates (Plate 10,
Figure 4). The gladius is penniform with a maximum
width about $ of the length. The rhachis has a sharp
keel which appears as a dark streak on the dorsum of
the mantle. The spermatophores are about 5.5mm in
length and 0.16mm wide. The sperm cord is thick and
occupies about 3 of the length. The distal portion of
this is opaque and without structure, posterior to this it
coils 2$ to 3 times and continues to a striated portion
(Plate 10, Figure 8).
Types:

Holotype (male): Station 87.90 of CalCOFI Cruise

25 (31°59’N, 122°24’W), May 1951.

Paratype No. 1 (female): (32°39.3’N, 117°37.2’ W),

August, 1953.

Paratype No. 2 (male): (32°49’N, 117°43’W),

August 1953.

Paratype No. 3 (male): St. 93.90 of CalCOFI Cruise

5704 (30°49.5'N. 121°32’W), April 1957.

Deposition of Types:

Holotype: U.S. National Museum, one male, USN
M 678792

Paratype 1: U.S. National Museum, one female, US
NM 678793

Paratype 2: California Acad. Sci. Dept. Invert. Zool.,
one male, CAL Acad. Invert. Type Coll. No. 310

Paratype 3: Scripps Institution of Oceanography,
Marine Invertebrate Collections, one male.

DISCUSSION

There are at least 3 other species of the genus in the
North Pacific. As is frequently the case with oceanic in-
vertebrates, which are difficult to collect, many of the

Explanation of Plate 10

Abraliopsis (Watasenia) felis McGowan & OKUTANI, Nn. sp.

Figure 1: Latero-ventral view of the cephalic part (Paratype No.
3, male) showing funnel cartilage, olfactory lobe, nuchal folds and
the funnel adductors. The photophores on the eyeball are shown
by removing a part of the eye lid. The photophores on the integu-
ment are not drawn.

Figure 2: Buccal membrane (Paratype No. 3). Ventro-anterior view.
Figure 3: Funnel organ (Paratype No. 3). The funnel is cut open.
Figure 4: Three transverse rows of the radula (Paratype No. 3).
Figure 5: Club and carpus of the tentacle (Holotype).

Figure 5a: A large sucker of the tentacle (Paratype No. 3).

Figure 5b: A large hook of the tentacle (Paratype No. 3).

Figure 6: Right ventral arm of the male (Paratype No.2).

Figure 6a: The distal tip of the right hectocotylized arm
(Holotype).

Figure 6b: The distal tip of the left hectocotylized arm (Holotype).

Figure 7: Left third lateral arm (Holotype).

Figure 8: A spermatophore taken from the spermatophore sac

(Paratype No. 3).

THE VELIGER, Vol. 11, No. 1

[McGowan & Oxutanl] Plate 10

ge abs beetle wnt

Vol. 11; No. 1

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Page 75

descriptions and reports of these species are confusing
and contradictory because they are based on immature
or damaged specimens. Further, the problem of discrim-
inating between these species is difficult because the
authors did not report the ranges over which important
meristic, taxonomic characters varied. It has been neces-
sary, therefore, to re-examine these and other characters
of two closely “related” species, Abraliopsis (Watasenia)
scintillans BERRY, 1911 and Abraliopsis affinis (PFEFFER,
1912). The former is closely related to our new species
in a morphological sense and the latter has a contiguous
and occasionally overlapping geographical distribution.

Abraliopsis scintillans from Japanese waters was de-
scribed by Berry in 1911 and 1912. However, earlier
WatasE (1905) had reported a new species of the genus
from Japan but did not describe it. IsHrkawa (1913) in
reviewing the characteristics of this species decided that
Berry’s and Watase’s squids were identical and created
the new genus Watasenia (= Watasea IsHtKawa, 1913)
for it. This was done on the basis that Berry’s A. scintillans
had only 2 large tentacular hooks instead of the 3 large
and 3 small ones of A. morisii (VERANY, 1837) or 5
large and 5 small ones of A. pfefferi Jounin, 1896. A
further distinction of the new genus was based on the
fact that the right ventral arm was hectocotylized with 2
small semi-lunar membranes near the tip. This, according
to ISHIKAWA, set it off from A. morisii and presumably
other members of the genus because Coun (1910) after
synonymizing most other species of the genus with A.
morisi illustrates a spectacularly enlarged swimming web
(or “Schutzsaum’’) on the left ventral arm which he calls
the hectocotylus (plt. 6, fig. 1 and plt. 10, fig. 1). How-
ever, in at least one of the species he synonymized
(namely A. hoylei of Hoye, 1904, which Prerrer, 1912
described as a new species, A. affinis, see below) this large
swimming web is not the hectocotylus. The hectocotylus
of A. hoylei (of Hoye, 1904, not of Prerrer, 1884 or
1912 but rather = A. affinis Prerrer, 1912) is on the
right ventral arm and is somewhat similar to those of
A. scintillans Berry, 1911 and A. felis, spec. nov. Further,
Voss (1960) mentions having a specimen of A. morisii
with the right ventral arm hectocotylized. The type of
hectocotylization is not mentioned. Therefore one of
the two characters IsHiKAwa uses to distinguish his new
genus Watasenia is common in the genus Abraliopsis and
cannot be used to set off Watasenia uniquely. The other
character used by him, the presence of only 2 hooks on
the tentacle club rather than the 6 to 10 found in the
other members of the genus seems to us to be insufficient
for the creation of a new genus. At best, Watasenia can
be considered only a subgenus of Abraliopsis JousIN,
1896. We so consider it here.

The differences between Abraliopsis scintillans and A.
felis are listed in Table 1. It is evident that A. felis has
a different body shape and mantle length to fin width
ratio. It has more hooks on the manus of the tentacles
and the fixing apparatus of the tentacle carpus differs.
The ventral mantle photophores are arranged somewhat
differently and the arms have, generally, more hooks
but fewer distal suckers than in A. scintillans. Most of
the differences between these species are quantitative
rather than qualitative; however, they are consistent and
significant and may be used for a rather easy visual
separation of specimens. It is obvious that the two species
are Closely related and should be considered members of
the subgenus Watasenia.

South of the area occupied by Abraliopsis felis a second
species of the genus, A. affinis (PFEFFER, 1912), is found
in abundance (see Map). The first report of the presence
of this species in the eastern tropical Pacific was that of
Hoye in 1904 who illustrated it and identified it as
Abraliopsis hoyle: Jounin, 1896 (= Enoploteuthis hoylei
Prerrer, 1884). However, PFEFFER (1912), without ap-
parently seeing any specimens and using Hoy te’s descrip-
tion and figures as a basis, claimed that Hoy.e’s speci-
mens were not the A. hoyle: that he had described in
1884, but rather a new species, Abralia (= Abraliopsis)
affinis, which he proceeded to describe. PFEFFER further
re-described his A. hoyle: in more detail and it is clear
that Hoy e’s specimens do not fit this description. In
spite of this, Rosson (1948) reports that 8 specimens
(one male) were collected in the vicinity of the Galapagos
and Cocos Islands by the Arcturus Expedition. Rosson
chose to call these specimens A. hoylei PFEFFER. But he
did not illustrate the specimens, gave no counts of meris-
tic characters and apparently has confused the swimming
web of the left ventral arm with the hectocotylus. Fur-
ther, his single male specimen was “damaged” and “not
very well preserved” (op. cit., p. 118). It would seem,
therefore, that Abraliopsis affinis (PFEFFER, 1912) must
be considered a valid species.

We have in our collection at Scripps Institution of
Oceanography several hundred specimens of an Abrali-
opsis species from the eastern tropical Pacific that fit
both Hoy.e’s (1904) illustrations of “A. hoyle’ and
PreFFER’s (1912) description of A. affinis very well.
There is no doubt that they are the A. affinis of PFEFFER,
1912, but because his original descriptions are incomp-
lete, additional details need to be added. For example,
the left ventral arm is as Hoye shows it and resembles
strongly the left ventral arm of A. morisu as figured by
Cuun (1910) ; that is, with a greatly enlarged swimming
web. Cuun called this swimming web “the hectocotylus.”
While this is a good secondary sexual characteristic, being

No. 1

°
’

Vol. 11

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Page 76

Character

Tentacle
armature

Fixing
apparatus
Hectocotylus

Ventral
mantle
photophores

Arms, hooks

Arms
distal
suckers

Body shape

Mantle length (mm)
Fin width (mm)
Mantle length (mm)
Volume cm®

Eye ball
photophores

Geographic
range

_———————— ee ee a a a el a a ee ee See ee

Abraliopsis felis, n. sp.

Table 2

A Comparison of Several Species

Abraliopsis scintillans BERRY

3, or occasionally 4 large hooks 2 large hooks and no small

and 3 or occasionally 4 small
hooks in a double row. No
membrane on club.

4 suckers and 7 knobs,
compact.

Right or occasionally left and
right ventral arms. Two small
semilunar crests, offset.

A “random” arrangement but
with a narrow median bare area
entire length of mantle.

Ist L. range = 15-17, * 16
ist R. range = 12-17, x 15.4
2° L. range = 16-18, x 17
2” R. range = 16-19, x 17.4

3P0L. range = 15-18, x 15.9
322 R. range = 13-17, x 16
4m]. range = 10-14, x 11.8

4m R.range = 7-14, x 114
Ist L. range = 13-18, x 15
Is™R. range = 10-18, x 13
2° L, range = 13-18, x 15
2° R. range = 12-20, x 16
3 L. range = 4-18, x 12.3
3) R.range = 6-18, x 12.1
Elongate, fusiform

Width <4 length

Range = 1.0-1.3 3

x = 1.11
Range = 0.74 - 3.66
x = 1.28

2 large

3 small

California Current

hooks. Club with semilunar
membrane on outer side.

3 or 4 suckers and 3 knobs,
somewhat diffuse.

Right ventral arm. Two small
semilunar crests, offset.

An almost even arrangement
with median bare area entire
length of mantle.

1st L. range = 9 -12, x 10.8
1s™R. range = 8-12, x 10.5
2xo L. range = 11 - 14, x 13.1
2° R. range = 11-14, x 12.7
38> L. range = 10- 14, x 12.2
32 R. range = 12-13, x 12.5
4m™L.range = 9-11, x 10.2
4m R. range = 9-12, x 10.4
1st L. range = 26 - 34, x 29
1s*R. range = 26 - 34, x 29
2x L. range = 27 - 31, x 28
2 R. range = 28 - 34, x 31
38> L. range = 24 - 34, x 28.6
3" R. range = 28 - 34, x 29.5
Elongate, fusiform

Width <3 length.

Range = 1.2- 1.4

x = 1.28
Range = 0.57 - 0.86
x = 0.67

2 large

3 small

Coastal waters of Japan,
Korea and Okhotsk Sea

Abraliopsis affinis
(PFEFFER, 1912) '

3 large and 4 small hooks in
a double row. Club with semi-
lunar membrane on outer side.

3 to 5 suckers and knobs,
compact.

Right ventral arm with 3
small semilunar crests, offset.
Left ventral arm with greatly
enlarged swimming web 3 X
width of arm.

Arranged in longitudinal rows
with wide median bare area
terminating on distal 4 of
mantle in bare circular patch.
1s L. range = 14 - 25, x 18.2
ISR. range = 15-27, x 19
2x? LL. range = 16 - 26, x 20.6
2x? R. range = 16 - 26, x 20.4
3° L. range = 16 - 24, x 19.8
3°? R. range = 13 - 24, x 19.7
4m]. range = 14-47, x 31
4™ R, range = 17 - 40, x 24
1s L. range = 10 - 24, x 16

IR. range = 8-22, x 15
2° L. range = 6-26, x 11
2° R. range = 3-20, x 11
372L. range = 6-14,x 98
3P°R. range = 7-14, x 10.8

Short, semifusiform
Width <4 length

Range = 0.96 - 1.1

x = 0.98
Range = 0.90 - 2.8
x = 1.50

2 large

3 small

Eastern tropical Pacific

Abraliopsis hoylet
(PFEFFER, 1884) *

4 large and 4 small hooks in
a double row.

4 or 5 suckers and 7 knobs.

Not described.

Somewhat irregularly arranged
but tend to be in longitudinal
rows. Narrow median bare
area on posterior % of mantle.
1* 20

xv 19-21
3 19-21
4™ 19.91

“97 distal to hooked area”

“Short semifusiform”
Width <4 length

0.91, single specimen

1 large
4 small
A single specimen, “presumably
from the Mascarenes,” Indian
Ocean.

Vol. 11; No. 1

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Page 77

found only in the males, it is not the hectocotylus. The
right ventral arm of this species is hectocotylized just as
is the right ventral arm of A. scintillans and A. felis.
This hectocotylization (in A. affinis) takes the form of
3 small semilunar membranes near the tip of the right
ventral arm just posterior to the 3 large swellings that
contain the light organs. These membranes are offset
from one another, 2 of them are on the inner and 1 on
the outer lateral edges of the oral surface of the arm.
The most proximal of these is somewhat larger than the
other 2. It may be seen in Table 1 that the number of
hooks on this arm is fewer than on the left ventral arm.
The females of A. affinis do not, of course, have a hecto-
cotylus nor do they possess the greatly enlarged swimming
web. The web of the left ventral arm of the female is
similar to the male’s but very much smaller, being only
about $ as wide as the arm width. That the specimens
examined were, as a matter of fact, females of A. affinis
was determined by examining them for the presence of
ovaries and by comparing the other taxonomic charac-
to those of the male. A particularly useful character for
this purpose is the arrangement of the ventral mantle
photophores.

These ventral mantle photophores were illustrated well
by Hoyte (1904, plt. 10, fig. 1), but he did not discuss
the uniqueness of their arrangement nor did PFEFFER
in his description of A. affinis. In the more than 100
specimens examined by us, both the males and females
of this species have rather indistinct longitudinal rows
of photophores on their ventral mantle surface but with
a very broad medial area lacking such photophores. This
bare area begins at the anterior edge of the mantle and
terminates on the distal 4 of the mantle in a bare, circu-
lar patch somewhat greater in diameter than the width
of the medial bare area.

A fourth species of this genus is present in our collec-
tions from the waters off southern Baja California. It
resembles greatly CHuN’s (1910) description and illus-
trations (particularly plt. X, fig. 1) of Abraliopsis morisu
(VerRANyY, 1837). We cannot, however, formally identify
it as such because the hectocotylus of our specimens is
similar to that of A. scintillans, A. affinis and A. felis,

<— Explanation to Table 2

1 This is based primarily on an examination of 136 specimens in
our collections. The data presented here do not disagree with
PFEFFER’s description but merely enlarge upon it.

2 Based entirely on Prerrer, 1912.

3 It is obvious that the range over which this character varies
overlaps that of A. scintillans; however, by the Mann-Whitney
“U” test the differences between the means for the two species is
significant, p < 0.001 (Tate « CLELLAND, 1959, p. 89).

that is, small semilunar membranes near the tip of the
right ventral arm. The left ventral arm of these Baja
California specimens have the enlarged swimming web
as illustrated by Coun in A. morisii and which he called
the hectocotylus. Until the Atlantic and Indian Ocean
species of this genus (A. morisii, A. pfefferi and A. hoylei)
are re-examined and the nature of the hectocotylization
of the ventral arms clarified, no further discussion of
this fourth North Pacific species seems warranted.
Natural History: The identifying characteristics, distri-
bution and abundance of the larvae and juveniles of
Abraliopsis felis spec. nov. have been studied (OKUTANI
& McGowan, in press). The larvae of this species were
found to be the most abundant ones in the California
Current. They are more than five times as numerous as
the larvae of Loligo opalescens Berry, 1911. Seasonal
studies on the occurrence of the larvae show them to be
both most abundant and most frequent during the months
of June and July and although some larvae are found all
year, this probably indicates that the peak spawning ac-
tivity is in the early summer. They are more abundant
offshore (i.e., greater than 50 nautical miles from the
coast) than inshore and more abundant in the central
sector of the California Current (i.e., between Point
Conception [34°30’N], California and Punta Eugenia
[27°51’N], Baja California, Mexico). Based on an ex-
amination of 91 adult specimens the male to female ratio
male
female ue:

Thus far this species has been found only in the Cali-

fornia Current (see Map).

LITERATURE CITED

Berry, SAMUEL STILLMAN
1911. Note on a new Abraliopsis from Japan (A. scintillans,
n. sp.) Nautilus 25: 93 - 94
1912. A catalogue of Japanese Cephalopoda. Proc. Acad.
Nat. Sci. Philadelphia 64: 380 - 444; 4 figs.; 5 plts.

Cuun, Cari
1910. | Die Cephalopoden. I. Teil: Oegopsida. Wissen-
schaftl. Ergebn. deutsch. Tiefsee-Exp. Valdivia, 18: 1-402;
32 figs.; 61 plts.

Hoye, WILLIAM Evans
1904. Reports on the scientific results of the expedition to the
eastern tropical Pacific Albatross V. Reports on the
Cephalopoda. Bull. Mus. Comp. Zool. Harvard 43 (1):
legis iO) jks
Isuikawa, C.
1913. | Einige Bemerkungen iiber den leuchtenden Tintenfisch,
Watasenia n. gen., (Abraliopsis der Autoren) scintillans Brrry,

aus Japan. Zool. Anz. 43: 162 - 172

Page 78 THE VELIGER Vol. 11; No. 1

130° 120° 110° 100° SOg 80° 70°

Abraliopsis felis ©

Abraliopsis affinis oO

40°

30°

20°

10°

02

20°

FROM HO CHART 1500

130° (20° 110° 100° SO 80° 70°

Localities at which specimens of Abraliopsis felis and A. affinis
have been collected. The solid circles indicate A. felis, the open
circles A. affinis.

Vol. 11; No. 1

THE VELIGER

Page 79

Jounin, Louts
1896. Observations sur divers céphalopodes. Abraliopsis pfef-
fert (nov. gen. et nov. spec.) Bull. Soc. scient. et méd.

Ouert 5: 19-35

OxuTANI, TAKASHI & JoHN A. McGowan
In press. Systematics, distribution and abundance of plank-
tonic decapod (Cephalopoda) larvae from the California
Current, April 1954- March 1957. Bull. Scripps Inst.
Oceanogr.

PFEFFER, GEorG J.
1884. Die Cephalopoden des Hamburger Naturhistorischen
Museums. Abhandl. Naturw. Ver. Hamburg 8: 1 - 30
1912. Die Cephalopoden der Plankton-Expedition. Ergebn.
Plankton-Exped.. Humboldt-Stift. 2: 1-815; 48 plts.

Rosson, Guy Cosurn
1948. The Cephalopoda Decapoda of the Arcturus Oceano-
graphic Expedition, 1925. Zoologica 33 (3): 115 - 132

Sasaki, MapoKa
1929. A monograph of the dibranchiate cephalopods of the
Japanese and adjacent waters. Journ. Coll. Agr. Hokka-
ido Imp. Univ. 20: 1 - 357; 158 figs.; 30 plts.
Tate, M. W.& R. C. CLELLAND

1957. Non parametric and shortcut statistics. Interst.
Printers & Publ., Danville, Ill.
VERANY, J. B.
1837. | Mémoire sur six nouvelles espéces des céphalopodes

trouvée dans le Méditerranée 4 Nice. Mem. Acad. Sci.

Torino, Ser. 2, 1: 91 - 98

Voss, GILBERT L.
1960. Bermudan cephalopods.
to 446

WaTASE, S.

1905. Hotaru-ika no Hakkoki (“On the luminous organs
of the firefly squid’). Dobutu-Gaku-Zassi. 17: 119 - 123

Fieldiana, Zool. 39: 419

Page 80

NOTES & NEWS

Tellina ulloana

A New Species from Magdalena Bay,
Baja California, Mexico

BY
LEO G. HERTLEIN

Tellina declivis SowERBy, 1868 (species 261, plt. 44, fig.
261) was described without information as to the locality
from which it came. Tellina declivis ConraD, 1834 (p.
131) was described from the late Tertiary of Yorktown,
Virginia. This made necessary a new name for T. declivis
Sowersy. Datt (1900, p. 301) on the basis of the orig-
inal figure, judged the form to be a West American
species, and in this he has been followed by other authors.
We proposed a replacement name for the homonym as
Tellina (Merisca) proclivis HERTLEIN & StRoNG (1949,
p. 83; plt. 1, figs. 6, 7, 14), with an illustration of a West
American specimen.

Recently Dr. Kenneth J. Boss examined Sowerby’s
holotype and found that it is not a West American form.
An illustration of that specimen shown to me recently
by Eugene Coan bears out Boss’ observation.

Under the International Code of Zoological Nomen-
clature (Art. 72d), the replacement name applies to
Sowerby’s species and not to the West American form
that had been erroneously identified as Tellina declivis.
Therefore, Tellina ulloana HERTLEIN is here proposed
as a new species, based upon type specimen 9226 (Calif.
Acad. Sci. Dept. Geol. Type Coll.) , from Magdalena Bay,
Baja California, with the description, illustrations and
other details as given by HERTLEIN & StronG for T: pro-
clivis. This species has been reported as ranging south to
Panama.

Both Boss and Coan called my attention to the similar-
ity between the West American species here described as
Tellina ulloana and the Caribbean species T: martinicensis
D’Orpicny, 1842 (see WaRMKE & AssorT, 1961, p. 196;
plt. 40, fig. i).

Dr. Boss informed me (written communication Octo-
ber 20, 1967) that the type specimen of Tellina declivis
(T: prochvis) appears to be macomoid, perhaps from the
Caribbean but possibly elsewhere. LyncE (1909, p. 193)
cited “Tellina (Arcopagia?) declivis SowErBy” from the
Gulf of Thailand (Gulf of Siam) and from Singapore

THE VELIGER

Vol. 11; No. 1

and stated, “My specimens exactly agree with SowERBY’s
description and figure.”

This species is named for Francisco de Ulloa, the first
navigator to reach the head of the Gulf of California in
September, 1539.

LITERATURE CITED

Conrap, TimoTHy ABBOTT

1834. | Observations on the Tertiary and more recent forma-
tions of a portion of the southern states. App. Descriptions
of new Tertiary fossils from the southern states (pp. 130 - 157).
Journ. Acad. Nat. Sci. Philadelphia 7(1): 116-157 (post
September, according to Sherborn)

Dai, WILLIAM HEALEY

1900. Synopsis of the family Tellinidae and of the North
American species. Proc. U.S. Nat. Mus., 23 (1210): 285
to 326; plts. 2 - 4 (14 November 1900)

HErRTLEIN, LEo GeorcE & ARCHIBALD McCLurRE STRONG

1949. Eastern Pacific expeditions of the New York Zoolog-
ical Society. XL. Mollusks from the west coast of Mexico and
Central America. Part 7. Zoologica 34 (2): 63 - 97; 1 plt.

(10 August 1949)
Lynce, H.

1909. The Danish Expedition to Siam 1899-1900. IV. Ma-
rine Lamellibranchiata. Kongl. Danske Vidensk. Selsk.
Skrift., Raekke 7, Naturvidensk. Mathem. Afd. 3: 97 - 299;
pits. 1-5; 1 map

SOWERBY, GEORGE BRETTINGHAM (2%? of name)

1866 - 1869. Conchologia Iconica; or, illustrations of the shells
of molluscous animals (London), vol. 17: Teéllina, plts.
1-58 with expl. (July 1866 - April 1869)

WarMKE, GERMAINE L. & RoBERT TUCKER ABBOTT

1961. | Caribbean seashells. Livingston Publ. Co., Nar-
berth, Pennsylv. pp. i- x, 1 - 346; plts. 1-44; maps 1-19; 34
text figs.

AN EMENDATION

BY
CRAWFORD N. CATE

12719 San Vicente Boulevard, Los Angeles,
California 90049

It has come to my attention that a recent paper on West
Australian cowries (CaTE, 1968) contains some inaccu-
racies and errors which I wish to correct. They are as
follows:

Entry 18, p. 227: Cypraea (Lyncina) leviathan gedlingae
Cate, 1968. — The diagnosis of this new subspecies was
omitted. The shell of C. (L.) leviathan gedlingae differs
from that of C. (L.) I. leviathan ScHILDER & SCHILDER,

Vol. 11; No. 1

1937 by being narrower, more cylindrical, less pyriform;
more thickly, more solidly formed; by apparently being a
smaller form (it will require additional live collected ma-
terial to determine this) ; and by apparently being geo-
graphically isolated from the presently known range of
the nominate subspecies. Since publication of the above
mentioned article another specimen of C. (L.) I. ged-
lingae has been received; it was collected by Molly Ged-
ling in 1962 on the beach just S of Vlaming Head
Light, North West Cape.
Entry 19, p. 227: The data should read as follows:

(Go 12/3) 10/07 30) 25)
Entry 32, pp. 220 and 228; The designations for the
species in the text and on plate 29, fig.42 are incorrect.
They should be corrected to read: Erosaria (Erosaria)
lamarcku lamarckit (Gray, 1825). For the reference on
p. 220 read : Zool. Journ. 1: 506
Entry 37, p. 29: The data should read as follows:

POR ION 8:5 199 17)
Entry 61, p. 232: The assertion is made that the “species
is clearly separable from Bistolida s. stolida because of
the total absence of lateral marks at each quarter of the
shell.” Bistolida brevidentata does possess lateral marks
on the angle of the shell margins; however, what I in-
tended to convey is that these markings do not normally
become broader, extending up the sides of the shell, often
coming into direct contact with the large central, chest-
nut-brown color blotch as in B. s. stolida. This particular
shell pattern is almost never seen in B. brevidentata.

The indications of magnification for figs. 26 (plate 26)
and 33 (plate 27) should be corrected to x 13.

LITERATURE CITED

Cate, Crawrorp NEILL
1968. | West Australian cowries. — a second, revised, and ex-
panded report. The Veliger 10 (3) : 212 - 232; plts. 21 - 34;
5 maps. (1 January 1968)

CORRECTION OF AN OMISSION

The title of the paper by Dr. E. C. Haderlie, starting on
page 327 of our April 1968 issue should read the same as
in the Table of Contents, i. e.
Marine Fouling and Boring Organisms
in Monterey Harbor.

The two words “and Boring” were omitted quite acci-
dentally; in citing the work these two words should, of
course, be included. Your Editor apologizes for this very
regrettable oversight on his part.

THE VELIGER

Page 81

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Page 82

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On May 15, 1968 we published the second part of the
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The Biology of Acmaea

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THE VELIGER

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[Part 1: Opisthobranch Mollusks of California
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BOOKS, PERIODICALS, PAMPHLETS

Marine Botany: An Introduction

by E. Yate Dawson. Holt, Rinehart and Winston, Inc.
New York. 1966. pp. xii+371; illust. $10.95

The malacologist will find this a useful reference work
if for no other reason than that it aids in the recognition
of the seaweeds among which mollusks occur. It is more
than an identification manual, however, for there are
chapters on the marine environment, bacteria, fungi,
phytoplankton, the several groups of algae, their mor-
phology, physiology, ecology and distribution. The three
final chapters are on the history of the study of seaweeds,
utilization of marine algae, and collecting procedures.
Two appendices bring together several lists of useful
information, such as the research facilities of the United
States where marine collections are housed.

We may be glad that Yale Dawson was able to see this
book into print before his tragic drowning. It is a book
that one can read without feeling bogged-down by special
terminology, yet the technical terminology is there when
needed. Numerous line drawings and halftone pictures
are well chosen as illustrations.

MK

VENUS The Japanese Journal of Malacology

Vol. 25, nos. 3 and 4, July 1967. | Some papers from the
Eleventh Pacific Science Congress, Tokyo, 1966

Indo-Pacific faunal elements in the Tropical Eastern Pa-
cific, with special reference to the mollusks. —
WiLuiaM K. EMERSON
The composition and relationships of marine molluscan
fauna of the Hawaiian Islands. — E. ALison Kay
Cenozoic history of Indo-Pacific and other warm-water
elements in the marine Mollusca of New Zealand. —-
CuHar_es A. FLEMING
Mass production of molluscs by means of rearing larvae
in tanks. — Takeo IMAI
Cytological relationships of some Pacific Gastropods. -
J. B. Burcu
Characteristics and origin of archibenthal molluscan
fauna on the Pacific Coast of Honshu, Japan. —
T. OKUTANI
MK

Page 84

Natural History of Marine Animals.

by G. E. MacGinitie and Nettie MacGiniriz.

McGraw-Hill Book Company, New York, 1968. pp. xii
+ 523; frontisp. + 286 figs. in text (line drawing and
halftone illustrations) . Second Edition.

The authors are well-known naturalists who have de-
voted together more than one human life span to the
loving study of the marine animals. It seems almost
superfluous to say anything about this second edition, at
least as far as the many users of the first edition are con-
cerned, except, perhaps, to mention that various portions
of the book have been rewritten to include the results of
more recent findings. However, as is the earlier edition,
so is this one eminently readable, though there is no
avoiding of technical terminology where needed. As the
authors have intended, the book is useful to a beginning
nature-lover as well as to the teacher of marine biology.
In these days when molecular biology is the vogue - we
almost said, the fashion - it is good to see a book again
that deals with the entire living organism in its many
facets. It is unnecessary to enumerate the various chap-
ters into which the book is divided; the appendices,
occupying 40 or so pages, are, however, worth a special
mention.

RS

THE VELIGER

Vol. 11; No. 1

American Opisthobranch Mollusks

by Evetine Marcus and Ernst Marcus.

Studies in Tropical Oceanography No. 6. Institute of
Marine Sciences, University of Miami, Florida. De-
cember 1967. pp. viii + 256; 1 color plate; 155 + 92
line drawings in text. Clothbound, $7.50.

The book is divided into two sections. The first portion
deals with 62 species of mollusks from Florida, the West
Indies, the Pacific side of the Panama Canal Zone, and
from Brazil. The second portion deals with 47 species of
mollusks, mostly, if not exclusively, from the northern
Gulf of California.

In all, 40 new taxa are described on the species and
subspecies level and 2 on the generic level; 3 replacement
names are introduced.

The authors have devoted many extremely productive
years to the study of mollusks in general and to opistho-
branchs in particular. The present work reflects the rich
background in the knowledge possessed by the authors.
In view of the modern trend in secondary schooling to
neglect the teaching of the so-called classical languages,
it might be regretted that the authors steadfastly refuse
to give the etymology of the names they choose or coin.
Aside from this slight flaw, the work appears to come up
to the usual standards of these well known workers. It
is unequivocally an important contribution to the know-
ledge of a beautiful group of animals.

RS

THE VELIGER is open to original papers pertaining to any problem concerned
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P ELIGER

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME II OcrToseErR 1, 1968 NUMBER 2
CoNnTENTS
Five New Species of Mitridae from the Indian and Pacific Oceans. (Plate 11)
aEANN Ge VIC CAE ete eh eae Sie cess ieee ae) ah Gerda Go ee 6 ee ee lo ne OG

Archidoris odhneri (MacFar.Lanp, 1966) comb. nov., With some Comments on the
Species of the Genus on the Pacific Coast of North America.

INOBER TOURING 6 usta tte Many roe ot eels )ig) cel) ai Gioaee ou este Sota elutes: OO

Quantitative Studies on Cowries (Cypraeidae) of the Allan Hancock Foundation

Collections.
GERALD A) BAIKUS Hen Muerte iat cn ooy a i See ee ee on ms Gator 2 OS
An Additional Record for Cypraea teres in the Galapagos Islands. (Plate 12)
WiInLIAM KeeEMERSON & WILLIAM, FE. Oup, fro. 2 4. 5 a 98

The Ege Masses and Veligers of Southern California Sacoglossan Opisthobranchs.
(6 ‘Text figures)

IRIGHARDMEWa GREENE Gi (4050 ie cack aos 2 ea rae hol ee weil ces eu TOO

A New Species of Puncturella (Cranopsis) from the Northeastern Pacific.
(Plate 13; 1 Text figure)

Ie WMicikiCowANn, <ujAMes Ely MCLEAN] 95 {0 90002) 0 a) 2 es es LOH
Studies on Populations of the Cowrie Erronea errones (LINNAEUS). 5 Maps.

Franz ALFRED SCHILDER & Maria SCHILDER . . . .. . . + + «© « « « 109
A New Neptunea from the Pacific Northwest. (Plate 14)

PATEUENING (Gg OMUBED oe oe > loci ou a eee, ee eles pee ce a gee ee te A

An Investigation of the Commensals of Cryptochiton stelle: (MipDENDORFF, 1847)
in the Monterey Peninsula Area, California. (5 Text figures)

CEE WE NMI VIER SIDER rete ae ane ee ase noe en oui ae) reek oc oe TOE

[Continued on Inside Front Cover]

A a sn se
Distributed free to Members of the California Malacozoological Society, Inc.
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Second Class Postage paid at Berkeley, California

ConTENTs — Continued

Notes on the Habitat and Anatomy of Jouannetia quillingi from North Carolina
Coastal Waters. (Plate 15; 1 Text figure)

Doueras\. Ay Wore! | tees. ee ee ee TE

Some Observations on the Ecology and Behavior of Lucapinella callomarginata
(Plate 16; 1 Text figure)

RICHARD T./MILLER) 2s 15 a) ay, yom anetbeccmo hota Bunton mmol eae a ETO
Banding Patterns in Haliotis — II Some Behavioral Considerations and the
Effect of Diet on Shell Coloration for Haliotis rufescens, Haliotis corrugata,
Haliotis sorensent, and Haliotis assimilis. (Plate 17)
DAVID A.SOUSEN! (0 sy cel Sole) ts eee ae) Ome fen ea CS
Haliotis pourtalesii Dati, 1881 from Florida Waters. (Plate 18)
CHARLES J. GUuIGE ) a S50) Ce ey a te rege en

Removal of Pea Crabs from Live Oysters by Using Sevin.

Jay D. ANprews, Donna TurcEON & Marian HREHA . ...... . . IAI
New Northern Limit for the Limpet, Acmaea digitalis

WitiiaAm FE JESSEE/| (00.0 oo) kis ere) coll sete cele cote tc a state teers SLE
Feeding Behavior of Corambella steinbergae.

James W. MeBETH 50) .. #- ci) 207 eyo) salen ees sino tet Ue CO
NODES S2NEWS) 0c cer Eee oy (ie C17

A New Record of Corambella steinbergae LANcE, 1962.
TERRENCE GOSLINER

BOOKS? PERIODICALS & PAMPHEE MS yi mentrsimcn ae ene) tease etme LAC)

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

Vol. 11; No. 2

THE VELIGER

Page 85

Five New Species of Mitridae

from the Indian and Pacific Oceans

JEAN M. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plate 11)

INTRODUCTION

DuRING THE PAST SEVERAL YEARS a number of unknown
mitrids from widespread localities have come to my at-
tention from several sources. At least five of these seem
to be unquestionably new species. Three of them were
first encountered during my visit to Western Australia in
1966; the fourth was a single specimen from Easter Island
given to me perhaps a dozen years ago by Raymond L.
Summers of Petaluma, California; and the final species,
from Coron, Philippine Islands, was recently sent to me
for identification by Fernando G. Dayrit of Manila. All
five will be described here.

ACKNOWLEDGMENT

I am grateful for the valuable assistance of all of the
individuals and institutions mentioned herein. In addi-
tion, I should like to thank Dr. Takeo Susuki for his
expert help in processing my film, and Crawford Cate
for many less tangible kinds of aid.

1. In October 1966, Alexander Gilbertson of Geraldton,
Western Australia showed me the collection of miscel-
laneous shells he has put together during several years of
lobster fishing and dredging in the vicinity of the Abrol-
hos Islands, some 50 miles offshore from his home base.
Among these shells I noticed a species of mitrid I had
never seen before; later, though it was his only specimen,
Mr. Gilbertson very generously presented it to me for my
own collection. About a week later, while working in the
mitrid collection at the Western Australian Museum upon
the invitation of its Curator of Molluscs, Dr. Barry
Wilson, I found an almost identical shell from a different

locality some 250 miles to the south of the Abrolhos
Islands. To the best of my knowledge these are the only
two specimens of this species now known.

Pterygia gilbertsoni J. Cate, spec. nov.

(Plate 11, Figures 1a - 1d)

Shell large, solid, heavy, cylindrically ovate, ventricose,
light tan in color, mottled and vaguely axially striped
with irregular white “flames”. Sutures moderately im-
pressed and lightly crenulated; spire slightly less than
half the length of the shell. Whorls convex, numbering
8, plus 14 nuclear whorls; apex slightly eroded in both
known specimens. Surface of earliest whorls decussate
with numerous axial grooves and about 3 smooth, shallow
spiral grooves; body whorl with 3 distinct spiral grooves
at shoulder and about 8 additional irregularly-spaced,
very shallowly impressed spiral grooves on body whorl to
base of shell; in the paratype these are somewhat stronger
than in the holotype, and appear as darker brown,
faintly incised lines. Aperture about twice as long as spire,
straight; labrum in the holotype thin (immature); in the
paratype thick, simple and with barely perceptible callus-
nodes at end-points of incised spiral lines of body whorl.
Columella, aperture and lip porcelain-white. Columella
calloused, with 5 to 6 prominent oblique folds; anterior
canal short, straight.

Discussion: Pterygia gilbertsoni is somewhat similar to
P. hayashu (Kira, 1959), a rare form from Japan. It dif
fers from that species by its lack of axial spots and punc-
tate spiral sculpture throughout, both of these characters
being prominent in P hayashu. Pterygia gilbertsoni dif-
fers from P nucea (GMELIN, 1791) by its more produced
spire, its faintly incised spiral sculpture which is lacking
in P. nucea, its straight lip (flaring in P nucea), and the

Page 86

apparent lack of a periostracum, which is blackish and
fairly heavy in P nucea. Pterygia gilbertson differs from
P. dactylus (LinNaAkEus, 1758) in its more produced spire,
its fewer columellar folds and its less flaring lip.
Holotype: No. 1129-67, Western Australian Museum,
Perth, Western Australia. Length: 46.4mm; Width: 18.9
mm; Length of Aperture: 28.7mm. Collected 5 miles
west of NW end of Rottnest Island in 194 fms, on sand
and dead coral substrate. Leg. Hawo, 5/7/60.

Paratype: No. 12379 in coll. Jean M. Cate, Los An-
geles, California, U.S. A. Length: 51.6mm; Width: 20.4
mm; Length of Aperture: 32.7mm. Dredged in 24 fms
on coral bottom, 4 mile NE of Eaglesnest Island, Easter
Group, Abrolhos Islands, Western Australia, July 1963.
Leg. Alexander Sutton Gilbertson, a crayfisherman re-
siding at Geraldton; the new species is named in his
honor.

Type Locality: Off Rottnest Island, near Fremantle,
Western Australia in 19 fms (Lat. 32°00’S; Long.
115°30’ W). The known range at present is from Rott-
nest Island to the Easter Group of the Abrolhos Islands,
a span of 4°32’,

2. By an interesting coincidence, the other new species
from Western Australia also seems to belong to the com-
paratively small genus Pterygia Ropinc, 1798. It is re-
presented by a unique specimen now in the collection of
the Western Australian Museum at Perth.

Pterygia barrywilsoni J. CATE, spec. nov.

(Plate 11, Figures 2a, 2b)

Shell large, heavy, solid; cylindrically ovate, ventricose.
Sutures moderately impressed, spire less than half the
length of shell. Whorls convex, numbering 7 plus 1
nuclear whorl which is somewhat worn. All whorls
marked by closely punctate spiral grooves, about 21 on
the body whorl and 7 on the penultimate whorl. Colu-
mella whitish, with 4 prominent rust-colored oblique
folds. Color of shell off-white, with prominent brownish-
gray to yellowish irregular flame-like axial markings;
base of shell rust colored. Two fairly narrow white trans-
verse zones appear at about mid-point of dorsum, though
these are not so clearly defined on ventral side.

Discussion: Pterygia barrywilsoni somewhat resembles P
gilbertsoni, but has a rougher, rather chalky texture
whereas P. gilbertsoni appears smooth and shining. Ptery-
gia barrywilsoni has a distinctive surface ornament, a
much more developed punctate sculpture, and a vivid
coloration not present in other Pterygia species. The
original museum label indicates that this specimen had

THE VELIGER

Vol. 11; No. 2

been misidentified as Scabricola sphaerulata (= S. papi-
lio (Linx, 1808)), an error brought about evidently
through the combination of color and pattern which are
remotely similar to that species; however, P barrywilsoni
quite definitely seems to fall within the subfamily Cylind-
romitrinae and in the genus Pterygia because of its short
spire and general shell shape. The soft parts and radula
are unknown.

Holotype: No. 334-66 in the Western Australian Muse-
um, Perth, Western Australia. Length: 38.9mm; Width:
15.5mm; Length of Aperture: 23.0mm. Collected at
Nightcliffe, Darwin, Northern Territory, Australia. Leg.
Jo Cunningham, 1962.

Type Locality: Darwin, Northern Territory, Australia
(bets, WA AOV Ss Ibo, IxO°SS) 18))).

This new species has been named for Dr. Barry R.
Wilson, Curator of Molluscs at the Western Australian
Museum, in recognition of his pioneering activities in the
field of malacology on the west coast of Australia.

3. ‘The third new species to come to my attention in the
Western Australian Museum Collection is not from Aus-
tralian waters, but from the Sulu Sea area near Borneo,
where it was collected by the Pele-Sulu Expedition of
1964. Specimens from that expedition were shared with
the museum at Perth because its Curator of Molluscs,
Dr. Barry Wilson, accompanied the expedition as a collec-
tor. The type lot consists of 14 shells, which will be
shared with other appropriate museums of Dr. Wilson’s
choice.

Vexillum sitangkaianum J. CaTE, spec. nov.

(Plate 11, Figures 3a, 3b)

Shell long, slender, fusiform, somewhat turriculate; spire
shorter than body whorl. Protoconch deviated, pauci-
spiral, transparent glassy brown; teleoconch consisting of
10 flatly convex abutting whorls plus 14 nuclear whorls.
Axial sculpture of prominent smooth collabral costae
(10 to 11) on penultimate whorl which tend to become
obsolete near outer lip; costae not regularly aligned at
sutures. Spiral ornament of equidistant shallowly incised
grooves, 3 on spire whorls, about 15 on body whorl,
faintly granulose at neck. Aperture straight, siphonal
canal short and slightly recurved. Labrum relatively thin,
simple, flattened in the middle, constricted at base, nu-
merous faint irregular lirae within. Columella straight,
with 4 strong oblique posterior folds and one weak fold
anteriorly; peristome continuous. Siphonal fasciole weak-
ly produced, helicocone nonumbilicate.

Wolo! 1 Not2

Shell color pure white throughout except protoconch,
which is glassy brown. Protoconch present only in Para-
type 7, eroded in others. Weak, colorless periostracum
present in some specimens. Animal and radula unknown.
Discussion: Vexillum sitangkaianum most closely resem-
bles V. vulpecula (Linnageus, 1758), but differs from
that species in the following ways: it has a more slender
and tapering spire, more shouldered whorls, a more con-
stricted base, a flattened and more constricted outer lip,
fewer axial costae and a total absence of color or surface
ornament.

Holotype: No. 1230-67 in the Western Australian Mu-
seum, Perth, Western Australia. Collected by the Pele-
Sulu Expedition in 9 to 13 fms, South Lagoon, Sitangkai,
Sibutu Island in the southernmost part of the Sulu
Archipelago, northwestern Celebes Sea, March 25, 1964.

MEASUREMENTS or tHe TYPE LOT

(in millimeters )

Maximum Length of

Height Diameter Aperture
Holotype 34.2 13.3 20.4
Paratype 1 31.6 11.8 IV/E2,
Paratype 2 Bol) oF 14.1
Paratype 3 25.6 V8) 14.6
Paratype 4 DSS) OM 15.8
Paratype 5 24.3 9.4 14.5
Paratype 6 Bae 8.5 13.6
Paratype 7' ZR Oat 14.3
Paratype 8 Zeal 8.9 13.8
Paratype 9 23.0 ie 13.0
Paratype 10 22.3 10.1 159
Paratype 11 20.4 Us eZ
Paratype 12 20.0 8.9 12.1
Paratype 13 19.6 7.3 Hild

‘ with protoconch

Type Locality: 9 to 13 fms, South Lagoon, Sitangkai
(Lat. 4°50’ N; Long. 119°50’ E). The range of the spe-
cies is unknown, as the type lot represents the only speci-
mens known to me at this time.

The specific name sitangkaianum is derived from the
name of the type locality.

4. The fourth new species under discussion was collected
at Easter Island in the southeast Pacific Ocean by Father
Sebastian Englert in 1955; he sent a number of specimens
to Raymond Summers, who divided them among several

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Page 87

collections. One shell was sent to me, 4 went to the Cali-
fornia Academy of Sciences, and it is believed that
additional specimens were sent to the U.S. National
Museum, the American Museum of Natural History,
and possibly to other museums as well; no records were
kept of where they were sent. The 5 specimens now in
California (those at the California Academy of Sciences
and in my own collection) will be enumerated as the type
lot; I have not seen the shells sent elsewhere by Mr.
Summers.

Strigatella rapanuiensis J. CATE, spec. nov.

(Plate 11, Figures 4a, 4b)

Shell small, cylindrically ovate, nearly smooth; whorls
slightly convex, numbering 5 including the body whorl,
nucleus lacking; spire less than half the length of shell;
sutures impressed; labrum thick, smooth, slightly reflec-
ted in the middle, flaring at base; columella with 44
oblique folds; anterior canal very short, straight; color
dark yellowish-tan with approximately equidistant deep
chocolate-brown, minutely punctate spiral stripes; one
pale ochre band appears in center of spire-whorls and
2 near base of shell; interior of aperture and columella
glossy white. A periostracum may be present in living
examples. Animal and radula unknown.

Discussion: Strigatella rapanuiensis superficially re-
sembles Mitra vexillum REEveE, 1844 because of its similar
size and the general color and pattern. It differs, how-

ever, in that the stripes are not deeply incised as in M.

vexillum, instead being smooth, though minutely punc-
tate; S. rapanuiensis has a lower spire, a strigatelliform
outline, and a glazed white aperture, all of which are
lacking in M. vexillum.

The species is tentatively assigned to the genus Striga-
tella on the basis of its reflected lip, and its general
outline and color which conform to the typical characters
of that genus; however, further research is needed to
determine whether this placement is correct. Among the
Strigatellas it most closely resembles S. tristis (BRODERIP,
1836) from the Panamic Province, but the presence of
the striped pattern and a lack of a whitish band below
the sutures, among other characters, mark it as clearly
different from S. trastis.

Holotype: No. CAS 13103 in the California Academy
of Sciences Geology Department Type Collection, San
Francisco, California. Leg. Father Sebastian Englert,
1955.

Paratypes: Paratypes 1 to 3 in the California Academy
of Sciences Geology Department Type Collection (CAS
13104, 13105, 13106), San Francisco, California. Para-

Page 88

type 4 in the collection of Jean M. Cate, Los Angeles,
California (No. 12380).

MEASUREMENTS or tHe TYPE LOT

(in millimeters)

Maximum Length of

Height Diameter Aperture
Holotype 25.0 1179 15.7
Paratype 1 22.1 10.4 15)67/
Paratype 2 Al 10.8 14.4
Paratype 3 21.4 10.8 14.4
Paratype 4 24.3 11S 15.6

Type Locality: Easter Island (Rapa Nui), south Paci-
fic Ocean (Lat. 29°00’S; Long. 109°30’ W). The spe-
cific name rapanuiensis is derived from the local name
for the type locality. The range of the species is unknown,
as the type lot constitutes the only specimens of record
at present.

Addendum: My conviction that the single specimen I
received nearly a dozen years ago was a valid new species
now seems to be borne out by the discovery of at least 9
additional specimens (including the 4 at the California
Academy of Sciences). I have recently verified that the
American Museum of Natural History has at least one
specimen (W. E. Old, Jr., personal communication).
After the species description was in manuscript I received
4 additional shells as a further gift from Mr. Summers;
these had been collected by Father Englert (presumably
at the same locality) in 1965 — ten years later than the
original lot. I have not included the measurements of
these specimens with those of the type lot, but note that
they all fall within the size range listed above. It is
likely, from Mr. Summers’ recollections, that specimens
of Strigatella rapanuiensis exist in the collection at the
U.S. National Museum and possibly others as well.

5. The final new species to be described is a small,
colorful shell from the southern Philippines; it is repre-
sented by two nearly identical specimens.

THE VELIGER

Vol: i= INow2

Vexillum coronense J. CATE, spec. nov.

(Plate 11, Figures 5a, 5b)

Shell small, slender, fusiform, somewhat turriculate; spire
about the same height as the body whorl. Protoconch
lacking in both known specimens; teleoconch consists of
8 flatly convex whorls. Axial sculpture of 10 prominent
costae (11 to 12 on penultimate whorl) which are not
regularly aligned at sutures. Sutures impressed; spiral
ornament consists of numerous equidistant shallowly in-
cised transverse grooves (15 on body whorl) in intercostal
interstices, becoming nearly obsolete over ribs. Aperture
straight, siphonal canal short and slightly recurved. Neck
of shell, starting at first posterior columellar fold, is
coarsely ribbed, granulose. Labrum thin, simple, con-
stricted at base, with numerous faint irregular lirae
within. Columella straight, with 3 strong oblique folds
posteriorly and 1 very weak fold anteriorly. Siphonal
fasciole weakly produced, helicocone nonumbilicate.

Shell color bright, deep orange except neck and colu-

mella which are both porcelain-white. First 4 whorls
bear narrow white spiral bands; final 2 to 3 whorls are
surrounded by a fainter orange band at periphery. Animal
and radula unknown.
Discussion: Vexillum coronense is so markedly different
from any other species known to me that it was difficult
to select a species for diagnostic comparison. Almost ar-
bitrarily I have chosen V. moana J. Cate, 1963 because
of its similarity in coloration and construction at the base
of the shell; the sharply defined rugose white base is
striking in both species. Otherwise, however, V. coronense
and V. moana differ markedly, V. coronense being more
slenderly tapering and having an altogether different
sculpture, surface ornament and color.

In overall shape Vexillum coronense more closely re-
sembles V. intertaeniatum (SowerBy, 1874), but that
species lacks the interstitial spiral sculpture, is not con-
stricted at the base, has a greater number of axial costae
and bears a different type of surface ornament. The color
of V. coronense recalls the brilliant orange of V. taenia-
tum (Lamarck, 1811), but is perhaps a shade or two
deeper and richer in color than V. taeniatum.

Explanation of Plate 11

Figures 1 a, 1b: Holotype, Plerygia gilbertsoni, spec. nov. (ca. X1)
Figures 1c, 1d: Paratype, Pterygia gilbertsoni, spec. nov. (X 1)
Figures 2a, 2b: Holotype, Pterygia barrywilsoni, spec. nov. ( X 14)

Figure 3.a,3b: Holotype, Vexillum sitangkaianum, spec. nov. (X 14)
Figures 4a, 4b: Holotype, Strigatella rapanuiensis, spec. nov. (X 2)
Figures 5a, 5b: Holotype, Vexillum coronense, spec. nov. (X 3)

Tue VELIcER, Vol. 11, No. 2 [J. Care] Plate 11

Figure 1c

Figure 2b

photographs by Jean M. Cate

Wolk ll: No. 2 THE VELIGER Page 89
Holotype: No. 13112 in the California Academy of ean ee FREDERICH ant

; : “ : ystema naturae per regna tria naturae itio
peromces Gec doey Department Type Calicenion, San Fran decima tertia, aucta, reformata 1 (6): 3021 - 3910 Lipsia.
cisco, California. Length: 16.7mm; Width: 6.0mm; ein, TRS
Length of Aperture: 8.3mm. 1959. Descriptions on Siphonalia trochulus tokaiensis, n. sub-
Paratype: No. 12382 in collection of Jean M. Cate, sp., and Mitra hayashu, n. sp. Venus 20 (4): 339 - 342;
Los Angeles, California. Masts ew

: 4 - Phil Lamarck, JEAN-BAPTISTE PIERRE ANTOINE DE MONET DE

Type Locality: Coron Island, Calamian Group, Phil- 1811. Sur la détermination des espéces parmi les animaux sans

ippine Islands (Lat. 12°10’ N; Long. 120°13’ E).
The specific name coronense is derived from the name
of the type locality.

LITERATURE CITED

Broverip, WILLIAM JOHN
1836. Genus Mitra (Lamarck & Swainson).
Soc. London for 1835, part 3: 192 - 198
Care, JEAN McCreery
1963. Revision of Dati’s Hawaiian Mitrids with descriptions
of three new species (Mollusca: Gastropoda) . The Veliger
6 (1): 23-43; plts. 5-8, 1 map (1 July 1963)

Proc. Zool.
(8 April 1836)

vertébres, et particulierement parmi les mollusques testacés.
Ann. Mus. d’Hist. Nat. Paris (Mitra) 17: 195 - 222 (July 1811)
Link, Hertnricu FRIEDRICH
1807. Beschreibung der Naturaliensammlung der Universitat
zu Rostock, 2%¢ - 3'¢ Abth., pp. 82 - 160
(29 March-17 May, 1807)
LINNAEUS, CAaROLUS
1758. | Systema naturae per regna tria naturae. Ed. 10, refor-
mata. Holmiae 1 (1): 1-824
REEVE, LoveLL AUGUSTUS
1844-1845. Conchologia Iconica. Monograph of the genus Mitra.
10: plts. 1 - 39
SoweErBy, GEorcE BRETTINGHAM
1874. Thesaurus Conchyliorum.
Mitra. 4: 1 - 46; plts. 1 - 28.

Monograph of the genus

Page 90

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Vol. 11; No. 2

Archidoris odhnert (MacFartanp, 1966) comb. nov.,

With some Comments on the Species of the Genus

on the Pacific Coast of North America

BY

ROBERT BURN

Honorary Associate in Conchology
National Museum of Victoria, Melbourne, Victoria

IN HIS MAGNIFICENT memorial volume on the Opistho-
branchia of the Pacific Coast of North America (1966),
the late Frank Mace MacFaruanp introduced many
new species with minutely detailed descriptions and beau-
tifully executed plates, both in colour and pen and wash.
The validity and synonymy of the new species is a
desideratum for students of the opisthobranch molluscs
the world over and remains an important task for com-
petent Pacific Coast researchers. In some instances, the
generic placement of species is open to question. The
present contribution, though very short, presents the case
for the generic transfer of one species.

Within the large family Dorididae, often the only cri-
teria for generic separation are found in differences of
the reproductive systems. Thus, genera have been founded
upon the armature or its absence in the male and female
ducts, the presence of a prostate gland whether discrete,
a mere dilation or its absence, and the formation of the
spermatheca and spermatocyst in their modes of attach-
ment to the vagina and uterine duct and in their rela-
tionship to one another. In the subfamily Doridinae
(OpHNER, 1939: 26, 27; = Doridinae plus Archidori-
dinae of OpHNer, 1926: 54), the external and pharyngeal
differences between some genera are so slight that only
careful examination of the reproductive organs can in-
dicate the true generic position. Thus in the following
text, considerable emphasis is placed on the various parts
of the reproductive organs.

Austrodoris odhneri MacFaruanp, 1966 (173 - 179;
plt. 26; plt. 29, fig. 14; plt. 36, figs. 1-19) is a large
apparently rare dorid from the Monterey Bay region of
California. MacFarLanp (pp. 171-173) gave a generic
definition, noted that his species was the first of the
genus from the northern hemisphere, and tabled the
valid and reputed species of the genus. The writer has
recently examined 3 species of Austrodoris from Austral-

ian Antarctica as well as 2 species of Archidoris from
New Zealand and Heard Island (an Australian depend-
ency in the subantarctic Indian Ocean), during which
time a survey of the literature of these 2 genera was
made.

Austrodoris ODHNER, 1926 (p. 55) is defined as having
(1) the winding vas deferens of uniform diameter with-
out prostate gland or prostatic section, and enclosed with-
in a tough leathery sheath for its whole length, (2) no
penis or penial armature, and (3) the spermatheca and
spermatocyst in vaginal combination. Almost all know-
ledge of Austrodoris is contained in two papers (ODHNER,
1926, 1934) where good figures of the reproductive sys-
tems of various species are given.

Study of MacFaranp’s figure (1966: plt. 36, fig 7)
shows that Austrodoris odhneri has (1) the vas deferens
at first wider and coiled, then much narrower and winding
tortuously within a long broad sheath of fibrous tissue,
(2) the former terminating in a low wide penial papilla,
and (3) the spermatheca and spermatocyst in semiserial
combination.

As these reproductive differences are of generic value,
Austrodoris odhneri cannot be maintained in Austrodoris.

However, this species does closely resemble Archidoris
wellingtonensis (ABRAHAM, 1877, p. 259) from New
Zealand in which (1) the vas deferens is at first narrow
and neatly coiled in a glomerate mass, then wider and
straighter as it passes through the long broad sheath of
fibrous tissue, (2) there is a low wide penial papilla, and
(3) the spermatheca and spermatocyst are in semiserial
combination. Hence the writer believes Austrodoris odh-
nert and Archidoris wellingtonensis to be congeneric and
the former to belong to Archidoris BeRcH, 1878, where it
will be known by the new combination Archidoris odhnern
(MacFar.anp, 1966).

Vol. 11; No. 2

On the other hand, the reproductive system of Archi-
doris montereyensis (Cooper, 1862) (MacFarLanp, 1966,
p. 181), a common species of the Pacific Coast from
Alaska to San Diego (STEINBERG, 1963: 70), differs con-
siderably from A. odhneri in that there is a very distinct-
ive digitiform penis and the spermatheca and spermato-
cyst, while still semiserially combined, lie very close
together. These reproductive characteristics are also pres-
ent in specimens of A. kerguelensis Bercu, 1884 (p. 85)
from Heard Island. It seems therefore that a re-appraisal
of generic and subgeneric units is necessary for these
species of Archidoris, but this must await comparative
examinations with specimens of the type species of the
genus, Archidoris tuberculata (Cuvier, 1804).

Furthermore, it should be pointed out that Marcus,
1961 (p. 16) appears to have confused Archidoris odh-
neri and Archidoris montereyensis. His figure of the re-
productive system (plt. 3, fig. 55) suggests by the spacing
of the spermatheca and spermatocyst and the absence of
a distinct penis that he examined small specimens of
Archidoris odhneri.

Two other species of Archidoris are reported from the
Pacific Coast of North America. The first is A. tubercu-
lata (Cuvier, 1804) to which there are but two references.
In the “Albatross” report on the dredgings along and off
the American west coasts, BercH (1894: 158) gives a
brief description of some Atlantic specimens of A. tuber-
culata, and in a separate paragraph (p. 159) records a
single specimen from off La Paz, Baja California (24°11’
N, 109°55’ W) in 10 fathoms. It was somewhat like A.
montereyensis in colour and shape of the radular teeth
though these were rather more numerous in number of
rows and teeth per half row (formula 56x 84084).
This specimen was 21 mm long, 14mm broad and 9mm
high. BERGH was quite familiar with A. tuberculata from
European and eastern Atlantic waters with its distinctive
patterning of larger tubercles set among more numerous
smaller tubercles (ALDER & Hancock, 1854; Family 1,
pit. 3, figs. 1-2, 6). As this specimen had a radular for-
mula within the range of that species, he no doubt
considered it identical. O’DoNoGHUE (1926, p. 207) re-
corded this specimen as Archidoris britannica (JOHNSTON,
1838).

Shortly afterwards, BeErcH (1900: 221) recorded a
smaller specimen from Bare Island (between Vancouver
Island and the Canadian mainland) which he also identi-
fied with Archidoris tuberculata. It was only 13mm long,
8mm broad and 4mm high with the radular formula
29 x 37-037. In this specimen the number of teeth per
row is half that of European specimens, hence the iden-
tification must be regarded as rather uncertain.

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Page 91

Zoogeographically, it is possible that Archidoris tuber-
culata should occur on both coasts of North America.
Already there are several nudibranch species with this
distribution, viz. Aeolidia papillosa (LinNAEus, 1761),
Onchidoris bilamellata (LinNaEuS, 1767), Dendronotus
frondosus (Ascantus, 1774) (Marcus, 1961: 56-57).
These 3 species also occur in Hokkaido, northern Japan
(Basa, 1957) and A. tuberculata is recorded from far
eastern Russian seas (VOLODSCHENKO, 1941: 60; 1955:
Ja), Itskd8 Foyle Gao, sitex, Se

ABRAHAM'S specimens from Vancouver Island, listed as
Doris tuberculata (1877: 198), were examined by O’Do-
NOGHUE (1926: 206, footnote) who found them to be
identical with Archidoris montereyensis.

The last species of Archidoris is A. nyctea BERGH,
1900 (p. 222) from Bare Island. It is known only from a
single 50mm long specimen with small (2mm diameter)
and smaller rounded tubercles and 8 branchiae. The
radula of 37 x 70:0:70 is both very close in formula and
shape of teeth to A. montercyensis. Similarly, the repro-
ductive organs with the vas deferens coiled into a twisted
mass and the semiserial spermatheca and spermatocyst
each with a long duct, are very close to A. montereyensis
(MacFartanp, 1966: 182; plt. 37, figs. 9, 10). BercH
appears not to have examined A. montereyensis in detail
(1878: 624; 1879: 107) except for the radula, hence
when confronted with reasonably fresh material in which
a spurious rhachidian tooth occurred, he preferred to
create a new species instead of referring it to the former.
Until it can be shown otherwise, the writer believes that
A. nyctea should be maintained among the synonymy of
A. montereyensis. O DONOGHUE (1921: 154; 1926: 206)
does not mention A. nyctea as a separate species nor list
it among the synonymy of any other species.

To summarize, there are four points:

1. Study of various figures of reproductive systems show
that Austrodoris odhneri MacFarLanp, 1966 is unten-
able in that genus and must be transferred to Archidoris
in the new combination Archidoris odhneri (MaAcFar-
LAND, 1966).

2. Three species of Archidoris occur on the Pacific Coast
of North America: A. montereyensis (Cooper, 1862)
with low bluntly conical small tubercles of uniform size
is widespread and common; A. odhneri (MAcFaRLAND,
1966) with low large and small tubercles is rarer and
probably often confused with the first; and A. tuberculata
(Cuvier, 1804) with larger tubercles set in a field of
smaller tubercles is reported from 2 internally differing
specimens.

3. Archidoris nyctea BercH, 1900 is most probably a
junior synonym of A. montereyensis; its only distinction
is a spurious rhachidian in the radula.

Page 92

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Vol. 11; No. 2

4. Archidoris tuberculata from Baja California and Bare
Island needs to be re-discovered and compared directly
with European specimens. The Bare Island specimen may
be only a small A. montereyensis with somewhat reduced
radula; that from Baja California may represent an extra-
limital southern form of the same species in which the
radula has evolved a greater number of teeth.

LITERATURE CITED

ABRAHAM, P.
1877. Revision of the anthobranchiate nudibranchiate Mol-
lusca. Proc. Zool. Soc. London 1877: 196 - 269; plts. 27
to 30 (August 1877)

ALDER, JosHuA, « ALBANY HaNcock
1845 - 1855. A monograph of the British nudibranchiate mollusca
1845. Ray. Soc., pts. 1-7.

Basa, K1KUTARO
1957. A revised list of the species of Opisthobranchia from
the northern part of Japan. Journ. Fac. Sci. Hokkaido
Univ. 6, Zool. 13 (1-4): 8- 14
Bercu, Lupwic SopHus RUDOLF
1878. | Malacologische Untersuchungen. In C. SEMPER,
Reisen im Archipel der Philippinen 2 (14): 603-645; plts.
66 - 68

1879. On the nudibranchiate gastropod mollusca of the north
Pacific ocean, with special reference to those of Alaska. Proc.
Acad. Nat. Sci. Philadelphia, pt. 1: 71-132; plts. 1-8.

1884. | Report on the Nudibranchiata.
Challenger Exped. 10: 1 - 154; plts. 1 - 14

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 “Alba-
tross’, during 1891. XIII. Die Opisthobranchien. _—_ Bull. Mus.

Rep. Sci. Results

Comp. Zool., Harvard Univ. 25 (10): 125 - 233; plts. 1-12
(October 1894)

1900. Ergebnisse einer Reise nach dem Pacific. Zool.
Jahrb., Abt. Syst. 13: 207 - 246; plts. 19-21

MacFarianp, Frank Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Marcus, ErNstT
1961. | Opisthobranch mollusks from California. The
Veliger 3 (Supplement, pt. I) : 1-85; plts. 1-10. (Feb. 1, 1961)

Opuner, Nits HyjALMAR
1926. Die Opisthobranchien. in: Further Zoological Re-
searches of the Swedish Antarctic Expedition 1901-1903, 2

(1): 1-100; plts. 1-3.

1934. | The Nudibranchiata. In British Antarctic (“Terra
Nova”) Expedition, 1910. Nat. Hist. Rep. Zool. 7 (5):
229-310; plts. 1-3. London.

O’ DONOGHUE, CHARLES HENRY

1921. Nudibranchiate mollusca from the Vancouver Island
region. ‘Trans. Roy. Canad. Inst. 13 (1): 147-209; plts. 7-11
1926. A list of the nudibranchiate mollusca recorded from the

Pacific coast of North America, with notes on their distribution.
Trans. Roy. Canad. Inst. 15 (2): 199 - 247.

STEINBERG, JOAN EmILy
1963. Notes on the opisthobranchs of the West Coast of North
America - IV. A distributional list of opisthobranchs from Point
Conception to Vancouver Island. The Veliger 6 (2) : 68 to

73 (1 October 1963)
VoLopcHENKO, N. I.
1941. New nudibranchiate molluscs from seas of the far east

of the U.S.S.R.
53 - 68; plts. 1-4

1955. Atlas of the invertebrates of the eastern seas of Russia.
Russian Acad. Sci. Moscow and Leningr. (Opisthobranchia:
181-185) 240 pp.; 66 plts.

Invest. Far East. Seas U.S.S.R. 1:

Vol. 11; No. 2

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Page 93

Quantitative Studies on the Cowries (Cypraeidae)

of the Allan Hancock Foundation Collections

BY

GERALD J. BAKUS

Allan Hancock Foundation, University of Southern California, Los Angeles, California 90007 '

THE CypRAEIDAE OF THE ALLAN Hancock Foundation
Collections consist of shells and preserved specimens taken
during the Hancock Pacific and Atlantic Expeditions (see
Fraser, 1943; Gartu, 1945). The remainder of the col-
lection includes cowrie shells collected from the Indo-
Pacific by various persons. Because information obtained
on Hancock Expediton-collected Cypraeidae is detailed,
quantitative studies were made on these specimens. Most
cowries included in this paper are presently on loan to
the Los Angeles County Museum of Natural History
under the care of Dr. James H. McLean, Curator of
Invertebrate Zoology. For convenience, the order of pres-
entation of information is as follows: 1) Panamic Re-
gion; 2) Caribbean Region; and 3) Indo-Pacific Re-
gion. Continental and insular geographical distribution
of Hancock Panamic and Caribbean Cypraeidae is dis-
cussed alphabetically by species. Quantitative information
is summarized in ‘Table 1, using standard statistical tests
found in Bamey (1959) and Simpson et al. (1963).
Hancock station number, latitude and longitude are in-
dicated in parentheses. Distributions, habitats, and shell
measurements are based on mature specimens unless
otherwise indicated. Hancock Indo-Pacific species are
listed alphabetically. Raw data and detailed records of
Hancock Cypraeidae are on file in the Hancock Foun-
dation. The living Cypracidae of the western hemisphere,
including data on certain meristic characters, are dis-
cussed in INGRAM (1951). A more recent summary of
Panamic cowrie distribution is given by Keen (1958)
and Emerson & Orp (1963), and that of Caribbean
cowrie distribution by WARMKE & Apspott (1961). Mol-
lusks of the Galapagos Archipelago are covered in part
by HERTLEIN & Strone (1955) and the Galapagos Cyp-
raeidae by INcRAM (1948).

' Allan Hancock Foundation Contribution No. 313

PANAMIC CYPRAEIDAE

1. Cypraea (Erosaria) albuginosa Gray, 1825

Localities: San Francisquito Bay, Gulf of California
(AHF 531-36; 28°25'55” N and 112°53’30” W) south
to Braithwaite Bay, Socorro Island, Mexico (AHF 128
-34; 18°42’45” N and 110°56’50” W) ; Bahia Honda,
Panama (AHF 247-34; 7°43’32” N and 81°32’19” W)
south to Port Utria, Colombia (AHF 419-35; 5°59’10”
N and 77°21’20” W) and Sulivan Bay, James Island,
Galapagos Archipelago (AHF 796-38; 00°17/00” S
and 90°35/13” W).

Specimens examined: 26 adults and 1 immature; 1 adult
specimen is preserved (AHF 2601-54).

Remarks: Data from the Hancock Collections support
KeeEn’s (1958) statement that this species has a dis-
continuous distribution, i. e., between the mid-Gulf of
California and Socorro Island (Hancock data) and
Manzanillo, Mexico (McLean, pers. communic.) and
from Panama to Ecuador and the Galapagos. There is
no significant difference in adult shell lengths between
the northern and southern populations (‘Student’s’ f-
teste 0105)

2. Cypraea (Zonaria) annettae Dat, 1909

Localities: Tepoca Bay, Sonora, Gulf of California (AHF
1077-40; 30°15’45” N and 112°53’20” W) and Puerto
Refugio, Angel de la Guarda, Gulf of California (AH
F 1049-40; 29°32’47” N and 113°34’35” W) south to
Agua Verde Bay, Gulf of California (AHF 655-37;
25°31’00” N and 111°01’45”W) and San Gabriel
Bay, Espiritu Santo Island, Gulf of California (AHF
634-37; 24°25'25” N and 110°20'55” W).

Specimens examined: 46 adults and 17 immatures.

3. Cypraea (Zonaria) arabicula LAMarRck, 1811

Localities: Isabel Island, Gulf of California (AHF 124-
33; 21°51’30” N and 105°53’35” W) and west of is-
lets off Navidad Head, Tenacatita Bay, Mexico (AHF

Page 94

275-34; 19°12’50” N and 104°49’48” W) south to La
Plata Island, Ecuador (AHF 22-33; 01°16’00” S and
81°05’10” W) and west of Manta, Ecuador (AHF
403-35; 00°56’43” S and 80°44’43” W).

Specimens examined: 38 adults and 24 immatures; 1
adult specimen is preserved (AHF 2596-54).

4. Cypraea (Macrocypraea) cervinetta KieNER, 1843

Localities: Isabel Island, South of Mazatlan, Mexico
(AHF 278-34; 21°51’30” N and 105°53’35” W) and
Tenacatita Bay, Mexico (AHF 272-34; 19°16’38” N
and 104°50’35” W) south to the Galapagos Archipel-
ago (6 stations) and west of Manta, Ecuador (AHF
403-35; 00°56’43” S and 80°44’43” W).

Specimens examined: 29 adults and 58 immatures.

5. Cypraea (Luria) isabellamexicana STEARNS, 1893

Localities: Secas Islands, Panama (AHF 867-38; 07°57’
10” N and 82°00’45” W) south to Sulivan Bay, James
Island, Galapagos Archipelago (AHF 796-38; 00°17’
00” S and 90°35’13” W).

Specimens examined: 4 adults.

Remarks: SHasxy (1961) reported collecting 3 living
specimens of this species near Guaymas, Sonora, Mex-
ico. McLean (personal communication) says that it
occurs with relative frequency around Cape San Lucas,
Baja California.

6. Cypraea (Zonaria) nigropunctata Gray, 1828

Localities: Jicarita Island, Panama (AHF 243-34; 07°
12/50” N and 81°48’05” W) south to the Galapagos
Archipelago (15 stations).

Specimens examined: 26 adults and 18 immatures.

7. Cypraea (Zonaria) robertsi Hwatco, 1906

Localities: Puerto Culebra, Costa Rica (AHF 115-33;
10°25’20” N and 85°40’20” W) south to Port Utria,
Colombia (AHF 413-35; 5°59/10” N and 77°21/20” W)
and off North Point, Gorgona Island, Colombia (AHF
409-35; 3°02’00” N and 78°10’30” W).

Specimens examined: 15 adults and 3 immatures.

8. Cypraea (Zonaria) spadicea Swainson, 1823

Localities: South of San Miguel Island, California (AHF
894-38; 34°01’00” N and 120°24’00” W) and Portu-
guese Bend, California (AHF 1446-42; 33°44’10” N
118°22’06” W) south to mouth of Rio Santo Tomas,
outer Baja California (AHF 1595-47; 32° N and 117°
W) and Melpomene Cove, Guadalupe Island (AHF
1920-49; 28°51'03” N and 118°17’43” W).

Specimens examined: 47 adults and 2 immatures.

Remarks: Smitu (1962) reported the collection of 20
specimens off Cypress Point, Monterey County, Cali-
fornia, at a depth of 80 feet (24m).

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Vol. 11; No..2

9. Cypraea (Blasicrura) teres GmMeuin, 1791

Localities: Secas Islands, Panama (AHF 447-35; 07°57’
10” N and 82°00’45” W) south to Bahia Honda, Pan-
ama (AHF 247-34; 07°43’32” N and 81°32/19” W).

Specimens examined: 2 adults.

Remarks: The specimen taken at Bahia Honda, Panama,
represents the first record of this Indo-Pacific species
from the Pacific American coast, so far as is known.
The first report from the Galapagos Archipelago
(beach, Puerto Grande, San Salvador Island) was that
of EMERSON & Oxp (1965).

ATLANTIC CYPRAEIDAE

1. Cypraea (Luria) cinerea GmeEun, 1791

Localities: Punta Arenas, Joyuda, Puerto Rico (Mattox
92); Punta Basora, Aruba, Netherlands West Indies
(A 16-39); and Buccoo Reef, Tabago Island, British
West Indies ( A41-39).

Specimens examined: 9 adults; 5 of these are preserved
(A 41-39).

2. Cypraca (Erosaria) spurca LinNAEuS, 1758

Localities: Outside Caledonia Bay, Panama (A 10-39),
Cabo Mala Pascua, Puerto Rico (Mattox 137), and
southwest of San Nicolaas Bay, Aruba, Netherlands
West Indies (A 18-39).

Specimens examined: 7 adults; 1 specimen is preserved
(A 18-39).

3. Cypraca (Macrocypraea) zebra LinNAEus, 1758
Localities: Rincon, Puerto Rico (Mattox 88).
Specimens examined: 1 adult and 1 immature.

INDO-PACIFIC CYPRAEIDAE

The number of shells examined for each species is placed
in parenthesis.

1. Cypraca (Monetaria) annulus Linnaeus, 1758 (503)
2. Cypraca (Mauritia) arabica Linnaeus, 1758 (35)
3. Cypraca (Palmadusta) artuffeli JoussEAUME, 1876 (2)
4. Cypraca (Erosaria) caputserpentis LinNAEUS, 1758 (49)
5. Cypraca (Cypraea) carneola Linnaezus, 1758 (3)

6. Cypraca (Erronea) caurica LinNAEUuS, 1758 (2)

7. Cypraca (Bistolida) coxeni Cox, 1873 (4)

8. Cypraca (Mauriti) depressa Gray, 1824 (8)

8. Cypraca (Mauritia) depressa Gray, 1824 (8)

9. Cypraca (Erosaria) erosa LINNAEUS, 1758 (20)

10. Cypraea (Erronea) errones LinnaEus, 1758 (11)
11. Cypraca (Erronea) felina GMEun, 1791 (1)

Vol. 11; No. 2

12. Cypraea (Palmadusta) gracilis GasKotn, 1848 (14)

13. Cypraea (Erosaria) helvola Linnaeus, 1758 (14)

14. Cypraea (Bistolida) hirundo Linnagus, 1758 (5)

15. Cypraca (Luria) isabella LINNAEUS, 1758 (12)

16. Cypraea (Erosaria) labrolineata Gasxotn, 1848 (5)

17. Cypraea (Cypraea) lynx Linnaeus, 1758 (18)

18. Cypraea (Palmadusta) lutea Gmeuin, 1791 (1)

19. Cypraea (Mauritia) maculifera ScuiLpER, 1932 (7)

20. Cypraca (Erosaria) miliaris GmMeuin, 1791 (5)

21. Cypraea (Monetaria) moneta LinnaEus, 1758 (492)

22. Cypraea (Erronea) onyx LINNAEUS, 1758 (7)

23. Cypraea (Bistolida) pallidula Gasxotn, 1849 (3)

24. Cypraea (Erosaria) poraria LINNAEUS, 1758 (1)

25. Cypraea (Bistolida) quadrimaculata Gray, 1824 (1)

26. Cypraea (Mauritia) scurra GMELIN, 1791 (1)

27. Cypraea (Talparia) talpa Linnarus, 1758 (1)

28. Cypraea (Cypraea) tigris LinnaEus, 1758 (5)

29. Cypraca (Cypraea) vitellus LINNAEUS, 1758 (7)

30. Cypraea (Erronea) walkeri Sowersy, 1832 (2)

31. Pustularia (Pustularia) cicercula (Linnagus, 1758)
(1)

32. Pustularia (Pustularia) globulus (Linnagus, 1758)
(1)

33. Staphylaea (Staphylaea) limacina (Lamarck, 1810)

(3)

34, Staphylaea (Staphylaea) nucleus (LinNAEus, 1758)
(7)

35. Staphylaea (Staphylaea) staphylaea (Linnaeus, 1758)
(2)

DISCUSSION

The discontinuous distribution of Cypraea albuginosa has
been narrowed, but reasons for the discontinuity are ob-
scure. It is assumed that geographical isolation, if gen-
uine, may have taken place in relatively recent geological
history because the meristic and ornamental features of
shells in the two populations are similar. Geographical
isolation may have originated during Pleistocene periods
of wandering isotherms (see BANDy, 1967). ScHILDER &
ScuHiLpEr (1938) consider the two populations to be
different races, but this decision is questionable according
to IncraM (1951) and quantitative data from the present
author.

Cypraea teres is reported from the Pacific coast of
the Americas for the first time, so far as is known. From
present data on habitat affinity and information on Cyp-
raeidae from eastern Pacific Islands (EMERSON & OLD,
1965), it is suggested that this Indo-Pacific species may
live on the Pacific coast of Central America in very small
numbers because of the absence, with few exceptions, of

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Page 95

coral reefs. EMerson (1967) noted that 47% of the
Clipperton Island (atoll) molluscan fauna has Indo-Pa-
cific affinities and concluded that the present impover-
ishment of coral reef habitat would appear to be the
primary factor that limits establishment of Indo-Pacific
mollusks in the Panamic region. I agree with EMERSON
and would add that lack of coral reefs would also imply
possible lack of specific foods preferred by Indo-Pacific
mollusks. Moreover, complex spatial heterogeneity (e. g.,
coral reefs) would appear to be an important factor in
the maintenance of high species diversity since it provides
numerous physical “niches” and this would offer the po-
tential for frequent biological interactions. Some of the
major reasons why Panamic animals have not been suc-
cessful in invading the Indo-Pacific region include pat-
terns of ocean currents, brief larval periods, competition
(Briccs, 1967), and in the case of Panamic cowries, their
affinity for non-coral hard substrates (see below).

An analysis of Table 1 provides certain insights into
cowrie populations. Caution should be taken since the
number of stations and specimens is somewhat low and
approximately equal effort and time in collecting cowries
were not allotted to each of the habitats. All Panamic
cowries were collected more frequently on rock than on
coral and several species showed no significant affinity for
coral over sand. This suggests that most Panamic cowries
may not be well adapted to the coral reef habitat, some-
thing quite different from that of many Indo-Pacific
Cypraeidae. Moreover, those Panamic species with close
Indo-Pacific affinities (i. e., Cypraea isabellamexicana and
C. teres) may have the most restricted habitat preferences
but the data are insufficient to support this hypothesis.
Cypraea albuginosa, C’. robertsi and C. spadicea show the
greatest affinity for marine plants. Whether this is indi-
rectly related to feeding specificity is unknown. Cypraea
albuginosa and C. spadicea show the greatest depth distri-
bution and C’. cervinetta the least, since the latter species
was collected at 28 shore stations from a total of 29
stations. Mean adult shell lengths are greatest in C.
cervinetta and least in C’. albuginosa and C. robertsi.
Shell lengths of the vicariate (very closely related) species
C’. cervinetta and its Caribbean counterpart C’. zebra are
similar although sufficient data are lacking for the latter
species. Confidence in the position of mean shell length
is greatest in C. annettae and least in C. cervinetta. All
species (with data calculated) show a somewhat high
degree of variation in shell length with C. roberts: the
most variable and C’. cervinetta the least variable for
their respective sizes. The statistics present here, with the
exception of certain coefficients of variation, compare
favorably with those of ScHILDER & SCHILDER (1966)
for the same species. Data from the present study serve to

Page 96 THE VELIGER Vol. 11; No. 2
Table 1
Habitat, depth, and shell length of cowries (Cypraea)
from the Pacific and Atlantic Hancock Expeditions
gs
Bey og £
a os s = g
Sea So uae
: oe g 03 *) as) o po
Species Se a2 eI § § pa
ral 40 4 ae) a ae} =| ae} s
Cypraea
albuginosa _ 22 35% 18% 4% 21% - 4% - 11% 7%
annettae 28 55% 14% 10% 7% 10% 7% — - =
arabicula 20 73% 9% 4% 14% = - - - =
cervinetta 29 67% 7% 13% 10% - 3% - - =
cinerea 3 2stations — - lstation — - - - =
isabellamexicana 3 2stations — - lstation — - - - -
nigropunctata 17 83% 11% - 6% - = = = a
robertsi 10 62% - 8% 7% - - - 23% =
spadicea 27 55% 7% 21% - - - 7% = 10%
Spurca * 3 - - - Il station station -— - - =
teres 2 - - - 2stations — - - - =
zebra * 1 - - - - - - - = =

provide information necessary for partial interpretation
of population structure. What is needed are specific data
on age and sex distribution, natality per female, mortality,
and considerably greater emphasis on ecological aspects
such as specific habitat affinity and feeding habits. Be-
cause cowries are thought to be rather specialized carni-
vores, feeding on ascidians (GRAHAM, 1955), detailed
knowledge of food and habitat should provide a logical
basis for successful discovery leading to thorough bio-
logical studies.

ACKNOWLEDGMENT

Greatly appreciated are the initial identifications of
Hancock cowries by the late Dr. Norman T. Mattox,
Allan Hancock Foundation, and especially the recent
identifications made by Mr. Crawford N. Cate, Los
Angeles, California. The manuscript was kindly read by
Dr. John S, Garth, Allan Hancock Foundation, and Dr.
James H. McLean, Los Angeles County Museum of
Natural History. Of considerable help were the references
provided by and cooperation given by Dr. McLean.

LITERATURE CITED

BariLey, Norman T. J.
1959. Statistical methods in biology.
London, England (English Univ. Press)
Banpy, OrvILLE L.
1967. Problems of Tertiary foraminiferal and radiolarian zona-
tion, circum-Pacific area. pp. 95-102 in: Harat, K. (ed.)

i-ix + 200 pp.

Tertiary correlations and climatic changes in the Pacific. Symp.
No. 25 of the 11™ Pacific Sci. Congr. (1966). Sasaki Publ. Co.,
Sendai, Japan, pp. 1 - 102
Briccs, JoHN C.
1967. Dispersal of tropical marine shore animals: coriolis
Nature 216 (5113): 350

parameters or competition?
Emerson, WILLIAM KEITH

1967. Indo-Pacific faunal elements in the tropical eastern
Pacific, with special reference to the mollusks. Venus 25
(3-4) : 85 - 93

Emerson, WiLuiAm KeituH & WILLIAM Erwoop OL, Jr.

1963. Results of the Puritan-American Museum of Natural
History expedition to western Mexico. 17. The recent mollusks:
Gastropoda, Cypraeacea. Amer. Mus. Novitates no. 2136,
32 pp; 18 text figs. New York, N. Y.

1965. | New molluscan records for the Galapagos Islands.
Nautilus 78 (4): 116 - 120

Fraser, C, McLean

1943. | General account of the scientific work of the Velcro
III in the eastern Pacific, 1931-41. Part III. A ten-year list
of the Velero III collecting stations. Allan Hancock Pac.
Exped. 1 (3): 259-431. Univ. South. Calif: Press, Los Angeles.
Calif.

GarTH, JOHN S.

1945. | Geographical account and station records of Velero III
in Atlantic waters in 1939. Allan Hancock Atlantic Exped.
Rept. 1: 1 - 106

GrauHam, ALASTAIR

1955. Molluscan diets.

144 -159.
HErTLEIN, LEo Georce & ARCHIBALD McC.iure StroNnG

1955. Marine mollusks collected at the Galapagos Islands

during the voyage of the Velero III, 1931-32. pp. 111-145

Proc. Malac. Soc. London 31:

Vol. 11; No. 2

THE VELIGER

Page 97

Table 1

Habitat, depth, and shell length of cowries (Cypraea)
from the Pacific and Atlantic Hancock Expeditions

ce €
£8 i E fe ,
Se e Oo +l — SI °
ao a¢ 3 Ps g £ ‘
ap SES 2 se Sy a5
Se See BE be De Be
>? GB » Qs S28 ao Se
isi =o =U = 6 = oO 38 =
ae goa oe oe 5 BC 45
ad Ce Seed ie 28 Il 23
pn ae esie 38 ee 3%
om ee oS Gs] aS} (G) 36
me} in eo co an au ao ao
Cy praea 22+0.14 22etie39 16.41
albuginosa 0-18-93 to 274 15-22-31 3.61
[13] [26] 37+0.65 37 +£1.04 12.03
annettae 0-5-38 25-37-47 4.45
[20] [46] 23 +0.79 23 + 1.20 16.43
arabicula 0- 3-46 to 64 17-23-29 3.78
[18] [38] 69+0.90 69+2.72 10.86
cervinetta O- 1-46 39-69-89 7.49
[28] [29] 2 - - -
cinerea shore 21-26-34 -
(9 - : =
isabellamexicana shore 29-37-43 -
[3] [4] 27 +0.88 27+1.75 16.89
" nigropunctata 0- 4-28 to 37 22-27-37 4.56
[14] [26] 22 +0.28 22+2.16 19.41
robertst Q- 4-37 17-22-28 4.27
[8] [15] 45 +0.89 45+1.70 13.22
spadicca 0- 7-91to 93 30-45-57 5.95
[21] 3 [47] - - -
Spurca 0 to 44 17-19-25 -
(7) S E >
teres shallow water 23 and 26 -
(I - - -
zebra littoral 68
[14 - - -

ee 00 Sa——>~"a>«>*>TRa22"— va——=

" Caribbean specics
2 AHF 48-33 (immature shell) measures 79 mm in length

In: Hatmos, D. (ed.) Essays in the natural sciences in honor
of Captain Allan Hancock. Univ. South. Calif. Press, Los
Angeles, Calif., pp. 1 - 345
IncRAM, WILLIAM Marcus
1948. The cypracid fauna of the Galapagos Islands. Proc.
Calif. Acad. Sci. 26 (7): 135-145
1951. The living Cypraeidae of the western hemisphere.
Bull. Amer. Paleont. 33 (136): 1-55
Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
i-xi+624 pp.; 10
Stanford Univ. Press, Stan-

from Lower California to Colombia.
colored plts.; 1700 text figs.

ford. Calif. (5 December 1958)
Scuitper, Franz ALFRED & MARIA SCHILDER
1966. The size of ninety-five thousand cowries. ‘The Veliger

8 (4): 208-215 (1 April 1966)

3 Two shells (AHF 2049-51) were collected from a mud bottom
at a depth of 604m in San Pedro Channel, southern California
4 An immature shell (Mattox 88) mcasures 74mm in length

SCHILDER, Marta, & FRANZ ALFRED SCHILDER
1938 - 1939. Prodrome of a monograph on living Cypraeidae.
Proc. Malacol. Soc. London, 23(3- 4): 119 - 231.
Suasky, Donatp R.
1961. Range extension for Cypraea (Luria) isabellamexicana
Stearns, 1893. The Vecliger 3 (4): 111-112 (1 Apr. ’61)
Simpson, Grorcr GAYLorp, ANNE Rowe & RicHarp C. LEwontIN
1963. Quantitative Zoology. i-vii + 440 pp. New York,
N. Y. (Harcourt, Brace & World)
Smirn, ALLyN GoopWwIN
1962. California brown cowrie in central California. The
Veliger 4 (4): 215 (1 April 1962)
WarMkr, GERMAINE L. « RoBpertT TucKER ABBOTT
1961. Caribbean seashells; a guide to the marine mollusks of
Puerto Rico and other West Indian islands, Bermuda and the
lower Florida Keys. x+346 pp.; 44 plts.; 34 text figs.
Narberth, Pennsylvania (Livingston Publ. Co.)

Page 98

THE VELIGER

Vol. 11; No. 2

An Additional Record for Cypraea teres in the Galapagos Islands

BY

WILLIAM K. EMERSON

AND

WILLIAM E. OLD, Jr.

Department of Living Invertebrates
American Museum of Natural History

Seventy-ninth Street and Central Park West, New York, New York 10024

(Plate 12)

IN A PREVIOUS PAPER on the marine mollusks of the
Galapagos Islands (EMERSON & OLD, 1965), we reported
the first records for the Indo-Pacific species, Cypraea (Ta-
lostolida) teres GMELIN, 1791, in these islands. One large,
nearly complete, beach specimen (A.M.N.H. No. 110483)
had been obtained at Puerto Grande, Isla San Cristobal
(not Isla San Salvador, as we erroneously stated). A
second dead specimen, which was mentioned in our
report, is now known to have been found by Mrs. Jacque-
line De Roy in a fathom of water in Academy Bay, Isla
Santa Cruz.

A third specimen has since been found by Mrs. Carmen
Angermeyer. It is a glossy, fresh, dead-collected specimen
obtained at Sombrero, Isla San Salvador at a depth of
2 fathoms (Plate 12, Figures 7 to 9). This specimen,
which is in the Angermeyer collection, is the largest ex-
ample of this species that we have seen, having a length
of 50mm and a height of 22.7 mm. The specimen from
Puerto Grande, although not complete, measures 46.2 mm
in length and 20.7 mm in height. The largest example of
this species recorded by ScHILDER & SCHILDER (1964, p. 8)
is a specimen in the British Museum (Natural History)
measuring 45 mm in length. No locality was cited for it.

Dr. E. Alison Kay kindly examined our Galapagan
specimens and she confirmed our identification. She com-

mented (7n litteris) that the specimen from Isla San Sal-
vador was the largest that she had seen with the possible
exception of a large fragment representing the dorsal part
of a shell that was dredged off Kihei Lagoon, Maui,
Hawaiian Islands, which is in the collection of Dr. C. M.
Burgess. According to Dr. Kay, the fragment fits over the
dorsum of the largest Galapagan specimen, and she con-
cludes that the Hawaiian specimen must have been nearly
50mm in length. She pointed out, however, that all
other specimens she has seen from the Hawaiian Archi-
pelago, including Midway Island, do not exceed a maxi-
mum of 35mm in length. We have found the larger
Galapagan specimen to be about 10 mm greater in length
than the largest specimen with data in the 50 lots from
throughout the Indo-Pacific region contained in the col-
lection of the American Museum.

In this widely ranging Indo- Pacific species, the shells
exhibit two distinct forms that apparently represent sexu-
al dimorphism. In the larger inflated form, presumably
the female, a maximum length of about 36 mm is most
commonly encountered. The smaller, less globose shells
of the apparent males, rarely attain a maximum length
greater than 30 mm. A similar dimorphic condition exists
in the population occurring at Clipperton Island, the only
other record for this species in the eastern Pacific. The

Explanation of Plate 12

Figures 1 to 3: Cypraea (Talostolida) teres pellucens MELVILL, 1888,
Secas Island, Panama, 7°57’10” N, 82°00'45”W, shallow water,
coral, 1935 (A. H.E 447-35; Frazer, 1943) ; K 1B

Figures 4 to 6: Cypraea (Talostolida) teres pellucens MELVILL, 1888,
off Fort Kamehameha, Oahu, Hawaii, shallow water, C. M. Bur-
gess coll. (A. M.N.H. 91898, ex-Burgess) ; X 1.3

Figures 7 to 9: Cypraea (Talostolida) teres GMELIN, 1791 (s. l.),
off Sombrero, Isla San Salvador [James Island], Galapagos Islands,

in 1 fm, Angermeyer collection;

XI

THE VELIGER, Vol. 11, No. 2 [EMERSON & OLD] Plate 12

Figure 4 Figure 5

Figure 7 Figure 8 Figure 9

Seige
7“

es if :

>
any)
- - =
: 4
mt =
f
o
oe
is
D
.
i f
a
\ ey
, ©
VW
aia
“i
“ 7

Vol. 11; No. 2

THE VELIGER

Page 99

largest specimen we have seen from this oceanic island is
35mm in length, and these specimens are typical of
examples from populations occurring in the Hawaiian
Archipelago and Central Pacific that have been recog-
nized as a subspecies, Cypraea (Talostolida) teres pellu-
cens MeEtvitt, 1888. A specimen from Oahu, Hawaii
(A. M.N.H. No. 91898; ex Burgess) is illustrated for
comparison (see Plate 12, Figures 4 to 6).

As one might expect giant specimens to occur in iso-
lated, peripheral areas of a widely ranging species, we
believe, on the basis of our limited Galapagan sample,
that it is prudent to consider these insular specimens a
large growth form of Cypraca teres s.1. that does not
merit subspecific recognition. Additional collecting may
demonstrate, however, that the Galapagan population is
unique.

ADDENDUM

After the text for this manuscript was completed, Bakus
(1968, p. 94, this issue) cited specimens of Cypraea
teres from west Panama, the first known records for this
taxon on the continental shelf of the New World. These
specimens, one from Secas Island and one from Bahia
Honda in the Gulf of Panama, were obtained by the
Allan Hancock Pacific Expeditions more than 30 years
ago.
Through the courtesy of Drs. Gerald J. Bakus and
James H. McLean these specimens were examined by us,
and they were found to be the small, narrow form, pre-
sumably males. The specimens appear to be referable to
the subspecies Cypraca (Talostolida) teres pellucens MEL-
VILL, which is known to occur in the eastern Pacific at
Clipperton Island. The Panamanian specimens were col-
lected alive in association with stony corals, apparently the
same habitat occupied by this species at Clipperton Is-
land (HERTLEIN & ALtison, 1960). The larger of the 2
Panamanian specimens, which is nearly an inch less in
length than the largest of the Galapagan specimens, is
here illustrated (Plate 12, Figures 1 to 3). The smaller
specimen from Panama (A.H.F. Station 247-34; Fra-
SER, 1943), measures 22.7 mm in length, 12.3 mm in
width, and 9.5 mm in height; it is a more mature indi-
vidual than the larger, figured specimen.

The discovery of this Indo-Pacific species in the Gulf
of Panama, although belatedly recorded, serves to de-
monstrate once again our limited knowledge of the
faunas associated with the isolated coral reefs occurring
in the southern part of the Panamic Province (EMERSON,
1967). Additional specimens of this species complex are
required from the eastern Pacific in order to compare

them with populations from the Hawaiian Archipelago
and the western Polynesian islands before the taxonomic
significance of the large Galapagan specimens can be
determined more critically.

ACKNOWLEDGMENTS

In addition to Mesdames Carmen Angermeyer and
Jacqueline De Roy of Academy Bay, Isla Santa Cruz,
Galapagos Islands, we are indebted to the following
individuals for courtesies of various kinds: Dr. E. Alison
Kay of the University of Hawaii, Honolulu and Dr. C.
M. Burgess of Honolulu, Hawaii; Dr. James H. McLean
of the Los Angeles County Museum of Natural History,
Los Angeles, and Dr. Gerald J. Bakus of the Allan Han-
cock Foundation, University of Southern California, Los
Angeles, California.

LITERATURE CITED

Bakus, GERALD JOSEPH
1968. Quantitative studies on cowries (Cypraeidae) of the
Allan Hancock Foundation Collections. The Veliger,
11 (2): 93-96 (1 October 1968)
EMERSON, WILLIAM KEITH
1967. Indo-Pacific faunal elements in the tropical eastern
Pacific, with special reference to the mollusks. Venus 25
(3, 4): 85-93; 1 fig.
Emerson, WILLIAM KeiTrH & WILLIAM ERwoop OLDp, Jr.
1965. | New molluscan records for the Galapagos Islands.
The Nautilus 78 (4): 116 - 120
Fraser, C. McLEAN
1943. General account of the scientific work of the Velero III
in the eastern Pacific, 1931-41. Part III. A ten-year list of
the Velero III collecting stations (charts 1-115). Allan
Hancock Pacif. Exped. 1 (3): 259 - 431. Univ. So. Calif. Press,
Los Angeles, California

GMELIN, JOHANN FRIEDRICH
1791. Caroli Linnaei systema naturae per regna tria naturae.
Ed. 13, aucta, reformata, Vermes Testacea Lipsiae 1 (6).
HERTLEIN, LEo GeorcE & Epwin C. ALLISON
1960. Species of the genus Cypraea from Clipperton Island.
The Veliger 2 (4): 94-95; plt. 22 (1 April 1960)
MELviILL, James Cosmo

1888. A survey of the genus Cypraea (Linn.), its nomencla-
ture, geographical distribution, and distinctive affinities; with
descriptions of two new species and several varietics. Mem.
& Proc. Manchester Literary & Philosoph. Soc., Ser. 4, 1:
184 - 252; plts. 1, 2

ScHILDER, MartA & FRANZ ALFRED SCHILDER

Hawaiian

(October 1964)

1964. Minima and Maxima in cowry shells.

Shell News 12 (12): 6-8

Page 100

THE VELIGER

Vol. 11; No. 2

The Egg Masses and Veligers

of Southern California Sacoglossan Opisthobranchs

RICHARD W. GREENE

Department of Zoology, University of California, Los Angeles, California 90024

(6 Text figures)

INTRODUCTION

Hurst (1967) HAS DEMONSTRATED the importance of
egg masses and veliger larvae to the systematics of north-
east Pacific opisthobranchs. The present paper deals with
the order Sacoglossa as an addition to the work of Hurst,
and as a contribution to our knowledge of this group.
Horst (op. cit.) has described the egg mass and veliger
of Olea hansineensis AcERSBoRG, 1923 (Sacoglossa:
Oleidae). A brief description of the egg mass of Her-
maeina smithi Marcus, 1961 has been given by Gonor
(1961), and Lance (1962) has given a short description
of egg masses of Stiliger fuscovittata LANCE, 1962 (both
Sacoglossa: Hermaeidae).

The animals included in the present study are Elysia
hedgpetht Marcus, 1961 (Elysiidae), Hermaeca dend-
ritica ALDER & Hancock, 1846 and Hermaeina smithi
(Hermaeidae). All three species were collected intertid-
ally in Los Angeles County, California. In order to make
the data more useful, the methods used by Hurst (1967)
for description of egg masses and veligers have been
applied.

EGG MASSES

All the egg masses described in this paper belong to the
Type B of Hurst (1967). They are more or less cylindri-
cal in cross-section through the individual egg bands.
When attached to a flat substrate, the masses possess a
very thin jelly-free layer, while those found tangled among
filamentous algae (i. e., Chaetomorpha) have no appar-
ent jelly-free layer.

Elysia hedgpethi (Text figure 1; Tables 1 and 2):

The egg mass of Elysia hedgpethi (Figure 1) is in the
form of a counter-clockwise spiral. The mass is invariably
attached along its entire length to the substrate. In the
field the eggs are laid among the fronds of Codium
fragile Harior, 1889, the alga upon which the animal
lives and feeds. Individual masses of eggs may range
between 4 and 6mm in diameter. The egg band itself
measures between 1 and 2 mm in width depending upon

the size of the spawning animal. The terminal portion of
the egg band sometimes encloses several capsules con-
taining no ova. A single ovum is found in each capsule.
The entire egg mass is white in color, and the eggs within
the band appear randomly distributed in space.

Figure 1

Egg mass of Elysia hedgpethi

The pattern of spawning and structure of the egg mass
of Elysta hedgpethi closely resembles that of E. maoria
Powe LL, 1937 from New Zealand (Rem, 1964).

Hermaca dendritica (Text figure 2; Tables 1 and 2):

This species lays an egg mass very similar to that of
Elysia hedgpetht. The mass is in the form of a counter-
clockwise spiral and is found in the field attached to the
fronds of Codium fragile. When kept in captivity, Her-
maea may lay thread-like masses along the glass sides of
the container. The egg mass of H. dendritica is white
and is attached to the substrate along its entire length
(Figure 2). The eggs are deposited in a spiral within the
egg mass which serves to distinguish the egg mass of H.
dendritica from that of E. hedgpethi, which is found on
the same alga species.

Vol. 11; No. 2

THE VELIGER

Page 101

Table 1

Characteristics of the Egg Masses of Sacoglossans

= on a

G, 8 &q Bs 5

os § 2 o 4

u af 3 5 8

ts} wn 9 Ss on n

qo) iS bo s a a a

(@) = 3 <x a > Q

& on

io)

: :

& 2 as) e
Ga 8 5s Le} ww
o = : g Sell Pom mies 1a a a 8 = a &
2 ° i=} 2 o o Oo a & 3 ae} (eS)
egg g ee oe fee

Ss VA & © @ 2 ee =e <x

Elysia hedgpetht x x XxX xe eX 14 10
Hermaea dentritica X >< ils 7-8 5
Hermaeina smithi X xX <i GHC <n nlimx 5-6 4

' see description

Hermaeina smithi (Text figures 3a, 3b; Tables 1 and 2) :

The egg masses of Hermacina smithi take at least two
different forms (Figures 3a and 3b). Gonor (1961) has
described masses taking the shape of a “C” with a total
length of about 20 mm. These are the most common type
found in aquaria when animals are kept in captivity.
In the field, however, the masses are deposited as tangled
strings among the filaments of Chaetomorpha aerea
Kirzinc, 1849. Tangled strings are also frequently found
on the sides of glass aquaria containing H. smithi. These
masses are a variation on the plano-spiral mass (Figures
1 and 2) and differ in that they are attached to the sub-
strate only at intermittent points along their length
(Figure 3b). The C-shaped masses are found either
floating on the surface film of the aquarium or are

Figure 2

Egg mass of Hermaca dendritica

Figure 3

Egg masses of Hermaeina smithi

attached along their entire length to the glass surface.
The egg capsules appear to be arranged in a spiral pattern
inside the egg mass.

In addition to the two shapes of egg masses observed,
there are also two color types. Gonor (1961), working
with specimens from San Juan Island, Washington, has
described the eggs as being “lemon-yellow” when spawned,
becoming paler with further development. In the present
study of Hermaeina smithi collected in southern Califor-
nia, both yellow and white egg masses were deposited by

Page 102

THE VELIGER

Vol. 11; No. 2

Table 2

Egg Capsule Dimensions

Elysia hedgpethi
Hermaea dendritica

Hermaeina smithi

Gon 80pn 100,

adults and were observed through hatching. The yellow
masses became visibly paler with development, while
the white egg masses remained white.

VELIGER SHELLS

The veliger shells of the three species of sacoglossans
studied are of the uninflated type, or Type I. Hurst

Figure 4
Veliger Shell of Elysia hedgpethi
a — right side b — left side c — ventral d — dorsal

@ minimum
maximum @

> average of greatest diameter

< average of least diameter

(1967) discussed at length the difficulties of orienting
veliger shells for measurement. Table 1 gives data on
development of the egg masses, and Table 3 gives meas-
urement data and length: width: depth ratios for the
veliger shells.

Elysia hedgpetht (Text figure 4; Tables 1 and 3):

The veliger shell of Elysta hedgpethi resembles that of
both other species in the present study in that the only
apparent sculpture consists of minute pits over the entire
surface of the shell which are visible only under 150
magnification. The lip around the aperture is somewhat
variable in that the anterior portion may project beyond
the rest of the lip (Figure 4a) or may be even with it.
In general, the gross shape of the shell seems to be of
major importance when comparing it with other species
(i. e., Figures 5 and 6).

Hermaea dendritica (Text figure 5; Tables 1 and 3):

Of the species considered in the present study, Her-
maca dendritica has the roundest of the shells examined.
In addition, Figure 5 shows that the posterior portion of
the shell is narrower than the anterior portion, and that
there is little evidence of coiling externally (Figure 5a).
Sculpture consists of small pits as in the other two species.

Hermacina smithi (Text figure 6; Tables 1 and 3):

The apertural lip on the veliger is commonly flared,
though not all shells examined had this appearance (Fig-
ure 6). Sculpturing is manifested once again by small
pits over the surface of the veliger shell. Of the three
species included in the present study, Hermaeina smithi
shows the greatest degree of coiling on the right side.

DISCUSSION

The usefulness of data on opisthobranch egg masses and
veliger larvae has been pointed out by Hurst (1967)
along with the problems of obtaining such data. THomp-
son (1961) discussed the importance of veliger shells in
the classification of sacoglossans.

Vol. 11; No. 2

THE VELIGER

Page 103

Table 3

Veliger Shell Dimensions

a

qe)

3

B

g

me

(e) Length Width Depth Ratio L:W:D
Elysia hedgpethi 10 105. +206 66.1p +11.2 Usp, =) Bpil 1e5f2)-3 ih 3 ily)
Hermaea dendritica 10 97n +15.0 65.54 + 8.2 Wiley, se Mil 1.48: 1: 1.09
Hermaeina smithi 10 1092 + 1.7 69.54 + 3.6 ey, se Gsil eay7/,S ilo jlale

Figure 5
Veliger Shell of Hermaca dendritica
a — right side b — left side c — ventral d —- dorsal

The egg masses of sacoglossans may be distinguished
not only by their gross form and positioning of egg cap-
sules, but also to a great extent by where they are found
in the field. Sacoglossan opisthobranchs are found in
specific habitats such as fronds of algae and generally
leave their eggs on the algal substrate. Thus, eggs of
Elysia hedgpethi and Hermaea dendritica are found on
the fronds of Codium fragile, while egg masses of Her-

maeina smithi are found among the filaments of Chaeto-
morpha acrea, or, as reported by Gonor (1961), among
the various algae in the Enteromorpha mat. LANCE
(1962) shows an egg mass of Stiliger fuscovittata attached
to Polysiphonia pacifica HoLtenBerc, 1942, the alga
upon which the animal feeds.
It is interesting to note that the three species of saco-
glossans considered all maintain a symbiotic relationship

Figure 6

Veliger Shell of Hermaeina smithi

b — left side c — ventral d - dorsal

a — right side

Page 104

THE VELIGER

Vol. 11; No. 2

with the chloroplasts of their algal substrate (GREENE,
in preparation). The chloroplasts are obtained during
feeding and are retained within the cells of the digestive
diverticula in a functional condition. The egg masses have
been examined for evidence of transmission of chloro-
plasts from one generation to the next, and the results are
negative. The chloroplast symbionts are apparently not
obtained until some time after settling and metamorphosis
of the veligers.

The veliger shells of the species described here are
distinguishable by their gross form and dimensions. The
shells of all three species are small compared with most
of those described by Hurst (1967). The egg masses
and veliger shells described here correspond well with
those described for sacoglossans in other parts of the
Pacific Ocean (OsTERGAARD, 1950; Rem, 1964).

ACKNOWLEDGMENT

The author wishes to express gratitude to Dr. Leonard
Muscatine of the Department of Zoology, University of
California at Los Angeles for his critical reading of the
manuscript and to James R. Lance for confirmation of
identifications of species used in the study.

LITERATURE CITED

Gonor, JEFFERSON J.
1961. Observations on the biology of Hermaeina smithi, a
sacoglossan opisthobranch from the west coast of North Amer-
ica. The Veliger 4 (2): 85-98; 13 text figs. (1 Oct. ’61)
Hurst, ANNE
1967. The egg masses and veligers of thirty Northeast Pacific
opisthobranchs. The Veliger 9 (3) : 255 - 288; plts. 26 - 38;
31 text figs. (1 January 1967)
LaNncE, JAMES RoBERT
1962. <A new Stiliger and a new Corambella (Mollusca:
Opisthobranchia) from the Northwestern Pacific. The
Veliger 5 (1): 33-38; plt. 6; 10 text figs.
OsTERGAARD, JENS MaTuHIAs
1950. Spawning and development of some Hawaiian marine
gastropods. Pac. Sci. 4 (2): 75-115.
Rew, J. D.
1964. The reproduction of the sacoglossan opisthobranch
Elysia maoria. Proc. Zool. Soc. London 143 (3) : 365 - 393
Tuompson, THoMas E.
1961. The importance of the larval shell in the classification of

the Sacoglossa and the Acoela (Gastropoda, Opisthobranchia) .
Proc. Malacol. Soc. London 34 (5): 233 - 239

Vol. 11; No. 2

THE VELIGER

Page 105

A New Species of Puncturella (Cranopsis )

from the Northeastern Pacific

I. Mcl COWAN

Department of Zoology, University of British Columbia, Vancouver, Canada

AND

JAMES H. McLEAN

Los Angeles County Museum of Natural History
goo Exposition Boulevard, Los Angeles, California 90007

(Plate 13; 1 Text figure)

West AMERICAN SPECIES of the genus Puncturella Lowe,
1827, have been treated in a dissertation by McLean
(1966). We have recently become aware of an unde-
scribed species not covered by McLEAn (op. cit.) and
have decided to present a detailed description before the
review of the genus is to be published by him.

The subgenus Cranopsis A. Apams, 1860 (type species,
Cranopsis pelex A. Apams, 1860, Japan), has previously
been characterized as applying to species of Puncturella
having the fissure placed in nearly central position on the
anterior face of the shell. McLEAN (op. cit.) showed
that a more useful criterion is the presence of a major
double rib, extending from the anterior terminus of the
fissure to the margin of the shell. The two halves of this
rib are connected by a channel showing a distinct line
of suture, visible under magnification. In the soft parts
of the animal the roof of the mantle cavity is split from
the mantle margin to the fissure. The suture on the
anterior rib evidently results from the shell being pro-
duced by the split mantle margin. In Puncturella s. str.
the mantle roof is continuous and is perforated only in
contact with the fissure of the shell.

West American species heretofore placed in Puncturella
s. str. but having the above diagnostic features of Punc-
turella (Cranopsis) are: Puncturella (C.) major Da.t,
1891; P (C.) cucullata (Goutp, 1846); and P (C.)
multistriata DauL, 1914,

Puncturella (Cranopsis) decorata Cowan & McLean,
spec. nov.

(Plate 13, Figures 1 to 5; 1 Text figure)

Description of Holotype: Shell of moderate size for
the genus; basal outline ovoid, sides nearly parallel, ante-
rior end slightly narrower than the posterior. Width to
Iength ratio 0.74. Anterior outline slightly convex; poste-
rior slightly concave; apex approximately central, strong-
ly down-curved and slightly deflected to the left. Fissure
long and narrow with a constricted lower portion. Radial
sculpture strong, composed of primary and secondary
ribs regularly placed and clearly demarked; primary ribs
originating on the apex; secondary ribs originating 2 to
3 mm below; 5 mm below the apex these number 13 ribs
(7 primary and 6 sccondary) in a 5 mm horizontal dis-
tance; secondary ribs precisely cquidistant from each
primary and nearly equal in strength to the primary ribs
at the shell margin. Tertiary ribs originating 5 to 8mm
below the apex, but not reaching the size of the primary
and secondary ribs at the margin. Ribs at margin broader
than interspaces. The primary rib extending below the
fissure is a double rib having a well defined sutural line
extending to the margin. This compound rib is slightly
deflected toward the right and is broader but not as raised
as are the adjacent primary and sccondary ribs. Ribs
elegantly beaded, deriving from regular horizontal rid-

Page 106

THE VELIGER

Vol. 11; No. 2

ges associated with the lines of growth, extending across
the sulci as well as over the radial ridges. Numerous
minute whitish punctations scattered in the channels be-
tween ribs; entire external surface speckled with minute
chestnut flakes of what appears to be a cuticle, their
abundance giving a brown appearance over the white
shell. Inner surface of shell glossy and translucent, trans-
mitting traces of the radial sculpture. Septum in the
form of an open arc, slanted forward. Internal groove
from fissure to margin clearly defined; margin of shell
crenulate, corresponding to extension of the ribbing.
Dimensions: Long. 19.7 mm; lat. 14.5 mm; alt. 10.3 mm.
Type Material: Holotype, National Museum of Can-
ada (NMC), cat. no. 45745 (Plate 13, Figure 1).

Type Locality: Off west coast, Queen Charlotte Island,
British Columbia, 53°21.3’ N latitude; 133°04.1’ W
longitude, at a depth of 106 fathoms (193 m). Collected
by Mr. Frank Bernard aboard Fisheries Research Board
of Canada vessel G. B. Reed, Bernard station 67-46, 11
August 1967. Seven additional paratypes of smaller size
were taken in the same haul. Two are deposited in the
type collection of the Los Angeles County Museum of
Natural History (LACM), cat. no. 1175; one in the
United States National Museum (USNM), cat. no.
678542; one in the Paleontological type collection of the
California Academy of Sciences (CAS), cat. no. 13102;
one in the Stanford University Paleontological type col-
lection (SUPTC), cat. no. 9961; and two in the Cowan
collection, cat. no. 7283a-b.

Distribution: Localities for additional specimens identi-
fied as belonging to this species are as follows:

1) Six specimens, Bjorka Island, near Sitka, Alaska,
56°49’ N; 135°50’W; 110-117 fms.; G. McT: Cowan at
Fisheries Research Board of Canada Station 66-2-26, 5
September 1966; LACM 6764 a (Plate 13, Figure 2) ;
Cowan coll. 6764b-f;

2) Five specimens, off Cape James, Hope Island, Queen
Charlotte Strait, B.C., 85-95 fms.; Cowan sta. 748,
Cowan and McLean, 22 May 1963; Cowan coll. 4649a-d;
LACM 4649e;

3) Three specimens, south side Matole Canyon, Califor-
nia, 300 - 100 fms.; N. B. Scofield sta. B. 17, 11 October
1950; CAS loc. 33179 (Plate 13, Figure 4) ;

4) One specimen off west end of San Nicolas Island,
California, 30-50 fms., Templeton Crocker Expedition,
27 August 1932; CAS loc. 27603 (Plate 13, Figure 5);
5) One specimen (juvenile), between Cortez and Tanner
Banks, California, 80 fms.; Louis Zermatten, April 1965;
S.S. Berry coll., Redlands, California, cat. no. 33354
(Plate 13, Figure 3) ;

6) One specimen, Cortez Bank, California, 60 fms., USFC
sta. 2911, USNM 130419.

Table 1
Specimen Length Breadth Height
No. (in millimeters)
NMC (Fig. 1) 45745 19.7 14.5 10.3
LACM 1175a 11.2 8.4 5.5
LACM 1175b 11.7 8.5 5.4
SUPTC 9961 13.7 9.4 6.4
CAS 13102 13.5 10.4 6.7
USNM 678542 11.4 8.1 5.9
Cowan 7283a 12.8 9.3 5.4
Cowan 7283b 9.0 6.5 4)
LACM (Fig. 2) 6764a 13.3 10.0 5.8
Cowan 6764b 12.8 9.0 5.0
Cowan 6764c 17.3 12.0 8.0
Cowan 6764d 11.0 79) 5.2
Cowan 6764e 13.0 10.8 6.0
Cowan 6764f 9.9 Tell 4.3
Cowan 4649a 15.0 12.3 7.8
Cowan 4649b 15.3 11.0 7.8
Cowan 4649c 10.7 8.4 5.6
Cowan 4649d 10.8 — 5.2
LACM 4649e 10.5 URE 5.7
CAS (Fig. 4) 33179a 23.4 17.8 12.8
CAS 33179b 22.9 15.2 10.0
CAS 33179c — 14,7 8.4
CAS (Fig. 5) 72603 18.7 13.5 8.5
SSB (Fig. 3) 33354 6.5 4.5 2.7
Ratio of parameters to length 75% 48%

314.1 236.7 159.3

Explanation of Plate 13

Figures 1 to 5: Puncturella (Cranopsis) decorata Cowan & McLEAN

spec. nov.

Figure 1: Holotype, National Muscum of Canada, cat. no. 45745.
Off west coast, Queen Charlotte Island, British Columbia, 106 fms.
Long. 19.7mm; lat. 14.5 mm; alt. 10.3 mm; x2
Figure 2: Off Bjorka Island, near Sitka, Alaska, 110-117 fms.
LACM 6764 a. Long. 13.3mm; lat. 10.0mm; alt. 5.8mm; X 2
Figure 3: Between Cortez and Tanner Banks, California, 80 fms.
SSB 3354. Long. 6.5 mm; lat. 4.5 mm; alt. 2.7 mm; x 5

Figure 4: South side of Matole Canyon, California, 300 - 100 fms.
CAS 33179 a. Long. 23.4mm; lat. 17.8mm; alt. 12.8 mm; X 2
Figure 5: Off San Nicolas Island, California, 30-50 fms. CAS
27603. Long. 18.7mm; lat. 13.5 mm; alt. 8.5mm; x 2
Figures 6 to 7: Puncturella (Cranopsis) multistriata DALL, 1914.
Figure 6: Cadboro Bay, Victoria, British Columbia. LACM A.375.
Long. 16.6mm; lat. 12.4mm; alt. 8.4mm; xX 2

Figure 7: Puget Sound, Washington, dredged. LACM A.8487.
Long. 25.0mm; lat. 20.0mm; alt. 15.7mm; X2

[Gowan & McLean] Plate 13

THE VELIGER, Vol. 11, No. 2

Vol. 11; No. 2

Dimensions: Table 1 gives the external dimensions of
24 specimens of Puncturella decorata. Width to length
ratio is indicated as 75%, height to length ratio 48%. A
corresponding series of 26 P multistriata has the width to
length ratio 82%, differing to an insignificant degree
but this species is taller, with a height to length ratio of
62%. Regressions of width against length and height
against length have been determined for samples of both
species. The formulae for the width to length regression
of PR decorata are W —0.771 L—0.7, height to length
H = 0.482 L+0.07. Equivalent values for P multistriata
are W — 0.758 L—0.51 and H = 0.716 L- 1.753.

The significance of differences between the correspond-

ing regressions of the two species have been examined by
the method suggested by Stmpson, Roe & LEWONTIN
(1960). The probability of significance of the differences
between the length-width relationship of the two species
is 0.8 to 0.9. The corresponding probability for the length-
height relationship is 0.05 to 0.02.
Discussion: The soft parts are not differentiable on the
basis of material on hand from other members of the
subgenus. The mantle is split from the margin to the fis-
sure. The radula of a paratype specimen (LACM 1175a)
has been examined (Text figure 1). The rachidian and 4
adjacent lateral teeth have slightly overhanging edges,
while the fifth lateral tooth has a large inturned central
cusp and a smaller basal cusp. A lateromarginal plate is
present and the marginals are numerous. The radula is
similar to that of other west American species of
Puncturella.

Figure 1

Radula of Puncturella (Cranopsis) decorata Cowan & McLEAN

The fissure extends nearly half the length of the ante-
rior face of the shell in juvenile specimens (Plate 13,
Figure 3), but this area decreases to about one-third to
one-fourth of this dimension in mature shells.

THE VELIGER

Page 107

The specimens examined are remarkably uniform in
sculptural detail. The structure of the internal area ad-
jacent to the septum is variable. A few specimens exam-
ined (Plate 13, Figure 2) show slight traces of the butt-
ress structure posterior to the septum, typical of Punct-
urella (Puncturella) galeata (Gouutp, 1846). In the
subgenus C’ranopsis this is also a characteristic feature of
P. major Datt, 1891. Its development in P decorata is
considerably less than in either of these species.

The species is closely related to Puncturella multistriata
Dat, 1914 (Plate 13, Figures 6, 7), which differs chiefly
in that the secondary ribs do not attain the size or prom-
inence of the secondary ribs in P decorata. Puncturella
multistriata sometimes bears distinct series of pits between
the vertical ridges of the external sculpture. These can
be impressed into the sides of the ridges scalloping their
sides, and suggesting beading. However, none bear the
regular horizontal ridges, crossing the primary and scc-
ondary ribs, as well as the intervening sulci, that give
rise to the characteristic beaded sculpture of P decorata.
The new species differs also in the position of the apex;
it is centrally placed in P decorata, and about one-third
the length of the shell from the anterior margin in P
multistriata. The upper portion of the fissure is consist-
ently broader than that of P multistriata, whereas in the
latter species it is only slightly broader than the lower
portion. Specimens of P multistriata have not been found
with the buttresses to the septum shown by a few speci-
mens of P. decorata. On the basis of specimens examined,
P. multistriata reaches a much larger size (to 32 mm long),
whereas the largest P decorata seen is 23 mm long (Plate
13, Figure 4). In the new species the shell appears to be
thinner and more fragile in specimens of similar size.

Different geographic and bathymetric patterns of
distribution are indicated for Puncturella decorata and
P. multistriata. Puncturella multistriata has been taken in
shallow depths from the Aleutian Islands and ranges south
to the Puget Sound area, where it has been dredged in
depths ranging to 50 fathoms. No verified records of the
species are known from south of Puget Sound. DaLw’s
(1921) record of the species “south to San Diego,” was
based on USNM 211927, not a P (Cranopsis); and the
“Cortez Bank” record (USNM 130419) is PR decorata.
The known range of P decorata is therefore from Sitka,
Alaska, south to Cortez Bank, California. The two specics
have thus far not been taken at the same collecting
station.

Page 108

THE VELIGER

ACKNOWLEDGMENTS

We are grateful to Mr. Frank Bernard of the Fisheries
Research Board of Canada for making the type material
available for our use. Dr. Leo Hertlein of the California
Academy of Sciences and Dr. S. Stillman Berry of Red-
lands, California kindly arranged the loan of specimens.
Photographs are by Mr. Armando Solis, Museum Photo-
grapher. The radular drawing was prepared by Cathy
Pearse.

LITERATURE CITED

Dai, WILtiAM HEALEY
1921. Summary of the marine shellbearing mollusks of the
northwest coast of America, from San Diego, California, to
the Polar Sea... . Bull. U.S. Nat. Mus. 112: 1 - 217; plts.
1-22 (24 February 1921)
McLean, James H.
1966. | West American prosobranch Gastropoda: superfamilies
Patellacea, Pleurotomariacea, and Fissurellacea. i- x+ 255 pp.;
pits. 1-7. Ph. D. thesis, Stanford Univ. (June 1966)

Simpson, Georce Gaytorp, ANNE Ror « RicHARD C. LEWOoNTIN
1960. Quantitative Zoology, rev. ed. viit440 pp.; illus.
New York, Harcourt, Brace & Co.

Vol. 11; No. 2

Vol. 11; No. 2

THE VELIGER

Page 109

Studies on Populations of the Cowrie Erronea errones (Linnaeus)

BY

FRANZ ALFRED SCHILDER

AND

MARIA SCHILDER

University of Halle, German Democratic Republic

(5 Maps; 6 Tables)

Erronea errones (LINNAEUS, 1758) IS A RATHER common

cowrie species distributed throughout the Indo-Pacific

Ocean from Ceylon to Samoa and from Japan to the

Exmouth Gulf and Sydney (EF A. Scuitper, 1965, p.

181). The individual variation in many characters of

the shell is considerable; nevertheless the differences in

the means or medians of different populations often can
be demonstrated to be mathematically significant.

Recent examination of far more than 4000 specimens (cf.

M. Scuitper, 1967a, p. 373) has indicated that the geograph-

ical races defined 30 years ago (F A. & M. Scuixper, 1938, p.

152) can hardly be maintained as taxonomic units. However,

our studies demonstrated the existence of an unspotted sym-

patric mutant living around Broome in Western Australia
(EA. Scuitper, 1968a, 1968b).

In the present paper we intend to investigate the cor-
relation between the medians of 5 selected characters in
133 natural populations coming from all regions of the
inhabited Indo-Pacific Ocean, and the geographical dis-
tribution of these medians. These investigations quanti-
tatively far exceed former studies on the same subject
published 7 years ago (FA. & M. Scuiper, 1961), though
localities from which we studied one or very ‘ew shells
only have been completely disregarded.

CHARACTERS

The following 5 characters have been selected for this
study, as their records are most complete in our card
registry of all personally examined cowrie shells:

L
BL

length of the shell in millimeters;

maximum breadth in % of length, mostly indi-
cating also the degrce of callosity of the margins
and of the base;

dorsal central blotch;

anterior terminal spots; usually the size of the
right (labial) spot has been considered only, the
few cases excepted in which the left (columellar)
spot is larger than the right one;

color of the base and the unspotted margins; the
bluish grey zonate inner lip in rather young spe-
cimens has been disregarded.

According to previous papers (e. g. SCHILDER, SCHIL-
DER & Houston, 1964, pp. 158-159, table 2) we have
classified cach character in 6 classes; Table 1 shows the
meaning of the figures | to 6 in the following paragraphs.

DB
AS

| ll

BC

The convexity of the base, the narrowness of the aperture,
and the visibility of the dorsal zones (SCHILDER « Houston,

Table 1
Class 1 2 3 4 5 6
L 17-19 20-21 22-23 24-25 26-28 29-35
BL 53-54 55 56 57 58 59-61
DB absent obsolete small rather large large very large
AS absent obsolete small rather large large very large
BC white almost white yellowish white pale yellow rich yellow orange

Page 110 THE VELIGER Vol. 11; No. 2
1964, table 2) have been omitted because these characters Ln DDENAGEEG
have not been registered in many populations, especially sev-
eral decades ago; in our earlier paper on Erronea errones we 15d Darwin: Lee Pt. L. Watson 30 56 5 1 3
have considered the characters DB and AS only (EA. « M. 5b Sunday Isld. (Perth) 28 58 5 2 2
Scuiper, 1961). Broome (Schelechoff) 24 59 3 1 4

A dash (-) in a column indicates that the character has not Broome (Uetz) 27:59 5 1 3
been recorded in a sufficient way. * Willie Creek Kalnins 22 59 5 1 3
* Quandong Kalnins 22058 ome las

* Gantheaume Pt. Kalnins 25 58 5 1 4

* Black Ledge Kalnins 26 57 4 1 3

POPULATIONS 15e Port Hedland (Schelechoff) 22 57 4 1 4

Port Samson B. Wilson 28 57 4 1 3

Table 2 contains the 133 populations considered for the Dampier Arch. Davena Exp. 22 56 4 1 2

present study, as they comprise a satisfactory number of 5th Lord Howe Islds. (Melbourne) 28 57 4 2 3

adult specimens. The 8 columns indicate: 47s Avoca: Norah Head W. Krause 2557

1. the designation of the region and area according to 2d fs Wiealgoallee W. Krause 24 6s

; * Moreton Bay Schelechoff 32 57 4 1 4

RA. SCHILDER, 1965, BE. 174-1755, ‘ Caloundra Schelechoff 32 56 5 2 4

2. the locality; an asterisk (*) indicates populations Miapl aise Lee & Matcott, 24m 5 TiCmOmne
comprising at least 40 adult specimens; 47c Lady Elliott Isld. “61” Schelechoff 24 59 3 1 2

3. the collector, or if in parentheses, the collection or Lady Elliott Isld. “65” — Schelechoff 26 56 3 4 3
museum in which the population has been preserved ; Mast Head Isld. (Melbourne) 25 56 4 1 2
series collected by the same collector at the same “ (Che hee Isic Clongonn DS De
place but at different times have been pooled to a oe ae aan 5 e 2 ; :
single population; Fitzroy Reef Coucom 28 56 5 1 1

4. median length (L) in millimeters; Humpy Isld. Coucom A 35 BH i 3

5. median relative breadth (BL) in % of length; * Middle Isld. Coucom BS 3 it a

6. median size of dorsal blotch (DB) : * Pumpkin Isld. Coucom 26 56 5 2 3

7. median size of the anterior spots (AS) ; and ~ Noa Lsappel Isld. Couto 26 56 5 1 3

8. median color of the base (BC), each expressed in Con ae recon 20 Da eee

; ; Piccaninny Pt. Coucom a) Sf 3 3 4

classes 1 to 6 as explained in Table 1. Double Heads Cree 35 55 4 3 3
Yeppoon (before 1954) (Summers) 30 56 4 2 38

Yeppoon (after 1954) Rita Mada 30 57 5 3 4

Table 2 Seaforth Matcott 29 60 4 4 4

Mackay Matcott 29 57 4 1 4

= SHIT DIGREERG Mackay: Shell Pt. Griffiths 28 58 5 3 3

Finlayson Griffiths 26 54 4 1 3

13m Tuticorin Winckworth 32 68 3 2 s Shute Harbour W. Krause me) S 3 2 B
13c Ceylon Eulenburg 2 BF to gS 4 * Proserpine: Dingo.Bay (Schelechoff) 23 57 4 3 4
Ceylon (Steinfurth) 28 56 4 5 4 * Penrith Isld. Houston wy BW BS i 3
Ceylon Stoliczka i, BB) 8 BH 4 * Scawfell Isld. Houston 20 56 5 1 3
Krusadai Winckworth pa iS) Dab) * Hayman Isld. Uetz 22 3) § i §

* Trincomali Winckworth mw Sf 1 § 47q Bowen (Summers) wm as i 8 8

14t Phuket Isld. Brandt i By By ab ae Port Denison (Tomlin) Byyn Oey 4 il @
14m Medan Hiner 18 55 - 5 — Port Denison G. W. Young 30) 04 alee

14a Andaman Islds. A. W. King me BN) ae a Cairns (Schelechoff) 22 59 4 4 5

Andaman Islds. Sandys 7 3 BB 4 Green Isld. Price lf” 8} 3 i il

Aves Isld. Winckworth es f4 2585 Buchans Pt. Price A) 3 8 B ZB
Interview Isld. Winckworth Ue OY Ge il UI 46c |New Caledonia Bougier 23 56 3 3 4
Nankauri Winckworth 2 Me = lie 2 ied: New Caledonia Durand 28 59 4 4 1

* Port Blair Winckworth Ol Sf & 2s * New Caledonia Rossiter Pee) 4 2 8

14s Pulo Weh Buitendijk 21°55 - 4 - Anse Vata Cernohorsky 20 58 3 3 3
Nias Isld. Schroder 21 58 4 4 2 Mondoure: — shore Cernohorsky 20 55 3 5 4
Padang de Priester 20 56 1 4 4 Mondoure: outer reef | Cernohorsky IS) Ge Ge a
Oosthaven de Priester 2, DON = 157723 46f Cuvu Cernohorsky 21 56 2 4 4

14) Labuan de Priester I) os) 4 * Vuda Pt. Cernohorsky 21 57 3 3 4
* Tyilaut Eureun de Priester 2 Ws) OD 4b B * Vatia Wharf Cernohorsky 20 55 3 4 3

Tjilatjap de Priester 23 58 - = - Manava Isld. Cernohorsky 19 56 2 1 3

—

Vol. 11; No. 2

Table 2 [Continued]

L_ BL DB AS BC

Caboni Cernohorsky 21 57 3 3 4

Nananu-i-Ra Cernohorsky 19 56 4 3 3

Vitilevu Bay Cernohorsky 18 56 3 3 4

Lodoni Cernohorsky 18 57 3 3 4

Bau Isld. Sixten Bock 20 56 2 2 3

Namuka Sixten Bock 22 57 1 3 2

46t Vavau Cordeira AS “Sy il) 4

Haapai Cordeira IQ) BO il 4b as

41s * Malaita: Ata’a van der Riet 25 55 4 3 3

41b New Britain: Vuatom O. Meyer Oil Bo Boe

New Britain: Ulamona J. Schneider I Ye 45

Admiralty Islds. (Melbourne) 22 56 4 1 3

41g Roon Isld. Deelder Bil G3 4g

Manokwari Jochim IS) a) Ge Ge

48m Sorong Barton 20 56 4 4 2

Sangir Islds. de Priester 18 55 - 4 -

Halmahera Bernstein 25 57 - - -

Menado de Priester 24 55 - 5 -

Busak (NW Celebes) (Leiden) 22a 5O) Ane ONS

* Amboina Hoedt Dp Bia Gh 8) 8)

Amboina Koller DON m=" 20) ee) 2

Amboina Ledru 23a de lta

48a Kaimana (West Irian) Ahlers Ail GH) a Bh

48t North Timor Wienecke 22 Daa
Bali de Priester 24 55 -— 4
48c Tijger Islds. Verdonk 20 55 - 5

Macassar Semper 3) 9 al ase
Macassar Toxopeus = ~ 42

* Gulf of Madjene van Nisse 200-56) 4) On —

48} “Indische Zee” (evi- (Leiden) Pil Gia), se ae

dently 1 locality)

Madura Jochim 7). Sy) BD)

Batavia Bay de Priester I SB 4 3 @

Batavia Bay id. (Dautzbrg.) 23 - 1 3 -

Edam Isld. de Priester 19 56 - 4 -

48s Singapore Doria 7 a4 DD

Singapore Semper 7%) SB) B ae =

Pulo Sakra Winckworth 26 5/744 4

48¢ Ko Si-Chang Orr, Steiner 24 54°95 4 4

* West of Ban Pe Brandt 2p Bo) 1B) Ge

* East of Ban Pe Brandt 2g BB) BS) 4a 3}

48p Zamboanga Semper OD = B a B

Bohol: Ubay Semper OP 88 4 2

Bohol: Panglao Semper A) 3 2 We

Cebu Ringe 18° 56 3 4 2

Samar Jagor 2 Gil BBQ 4s

Batangas Tucker 18 56 4-4 4

Palawan Clover I) OF B a 3

48y Pulo Condor Bavay 10-63 3 2 8

49r Okinawa: Kue C. Young 21564: 93, 3.

49s Tosa Azuma 306 4a 3

42p Palau Islds. (Godeffroy) Die 2d

Palau Islds. Semper Dil Be By iW 2

Yap Isld. Volkens 2 Ol i 8 @

42c Truk Isld. M. Hill 2 83 to Be

THE VELIGER

Page 111

FREQUENCY or MEDIANS

The frequency of the 6 classes distinguished in the 5
recorded characters is illustrated by Table 3.

Table 3
oe ee ono
L 1G) EB 2 RE ay iG
BL OB By oe) Ip
DB 12 55249489 132 =) =
AS 3) il Be
BC Goo 250632, ee

The extreme classes 1 and 6 are well represented in
L and BL because they comprise more millimeters or %
than the central classes; in the characters of color, how-
ever, the extreme class 6 is never represented as median.
In L, BL, and BC the maximum figure is in class 3,
whereas in the two characters of markings (DB and AS)
there are evidently 2 maxima of each in class 1 (markings
mostly absent) and 4 (markings mostly rather large) ;
the intermediate classes 2 and 3 are less frequent.

CORRELATION or CHARACTERS

The correlation of the medians of various pairs of char-
acters indicated in Table 2 may be illustrated by the 6
squares of Table 4.

(see page 111 for Table 4)

There is no distinct correlation between any pair of
the characters tabulated in Table 4. The average greater
breadth of populations of medium classes of length than
of extreme classes must be regarded as random; there is
no correlation in L: DB nor in BL: BC; the slightly
negative correlation in DB : AS and in DB : BC also can-
not be proved mathematically, as well as the slightly posi-
tive correlation in AS: BC. In any case, there is no
gencral parallelism in accumulation of pigment in the 3
characters DB, AS, and BC (see also F A. & M. ScHILDER,
1961, p. 304).

INDIVIDUAL VARIATION

The individual variation within extreme populations may
be illustrated by Table 5; the figures indicate the number
of specimens belonging to each class of the character.

Page 112 THE VELIGER Vol. 11; No. 2
Table 4
length length breadth
| 2 8 4} 12 Sh PAS coe 56) I. 2 3) 45 aeaG
hc a Ce 6 - - = = = = 6 - - = = = =
Bo. QA Bs 4 fF cot 5) al a5) 38. ROM OR aed a” 1 8
oi Ana ANOS cy em aed 2 29 6 6 9 6 SS 4 1° 3 9 1
S Bi Shy a9 O23 Rs. 3 Bi Ge CP De ae Bes at 3 ya Bly 2 6 8
te VLC RG e VG) feo ay Sey One Oe a Oe ee S82 3 5 5 Ouse
flee ele, IS ee “Ste itle et 32 z lees Qk 5s Oke she al fo-= = (2) 2a
dorsal blotch dorsal blotch anterior spots
12S) A ous <6 2 35 4 eo 6 sania ) (6)
ie PR Re ages 8 6 =) Seen 6+ en
Sho 2 2m er 6 6 8 fi os 426 ge 5So= 2 1 20m
S 4 4 2 7 13 8 - 8 A A 2 Tl VB = S) 4 6 49 9 4 =
6g 8 42 so 8 8 Q = Ru i reel Ge = —@ Se 9 10 “Iie
Shug ee 2 De AC APB NOUS, 8 2 3° 6 8 10 G = S 20-8 4 47
iin ellie t3,, Seal a t= Pe eam GB = 1 S&S -+ - iio
Table 5 relative breadth (BL) show great local differences which
may be influenced by environmental conditions of the
Ginn habitat. Besides, there is a distinct negative correlation
Chama, Lees eT o wa G WGETo between length and relative breadth in these 4 popula-
L* Vatia Wharf 13 Oh 13 42 = Q PD tions:
Moreton Bay - - -=- = 2 44 6
BL" Malaita: Ata’a 82 45 32 17 5 2 2
Willie Creek 2 2 7 13 2 © G Table 6
DB Trincomali 33 240 13 @ 2B il 1
Hayman Isld. = = 8 ly 3 4 5 i ae bo. Ge
AS Black Ledge SPA CIA Nn Un hubstheanteys 4 es
Trincomali 1 4 10 19 32 14. 5 L Willie Creek 8 24 27 22, 13 1
BG One Tree Isld. 2-7) @ = = = 2B Quandong It 14 29° 16 ie
Port Blair = = 6 29 51 21 5 Gantheaume Pt. - 16 15 27 23 2
Black Ledge = 1 13 1) 8 7
BL Willie Creek 2 4 7 13 20 41
™ The individual extremes in the measurable characters are Quandong - 2 8 16 20 28
L= 15 to 39mm and BL=5S0 to 65% in the 133 popu- Gantheaume Pt. — 1 15 18 21 29
lations. Black Ledge 3 7 10 21 24 14
DB Willie Creek 6 4 14 21 38 12
Quandong a 6) LD} OI IIS)
These figures establish that there are really significant Gantheaume Pt. 1 2 3 15 @y NZ
differences between populations. On the other hand, ad- Black Ledge Fk 7) we iO
jacent populations show great resemblance in many char- AS Willie Creek 33 9) Te
acters, even if they live under different ecological condi- ‘Quancong a =
5 ji ; A , Gantheaume Pt. 49-5) Will 3'4) 6a
tions. This fact may be illustrated in Table 6 by 4 popu- law Iuedes a 8 Gee
lations from an area of 41 miles around Broome: the BC. Willie Creek 9 12 57 21 2 1
distance between the 4 localities Willie Creek (95 shells), Quandong 2 13 34 14 10 1
Qvandong (74 shells), Gantheaume Point (85 shells), Gantheaume Pt. 3 il 2 @ 19 1
and Black Ledge (80 shells) is about 15, 15, and 11 miles Black Ledge 1 13 4% 20 4 1

respectively (according to Mr. A. Kalnins) ; Gantheaume
Point is a rocky coast, Black Ledge is a muddy reef (FA.
ScuiLper, 1968 a, 1968b).

Table 6 shows that the distribution of the 3 characters
in color (DB, AS, BC) is very similar in these 4 popula-
lations, as it may be caused by a common gene pool in
the whole area of 41 miles, whereas the size (L) and

2 The medians are printed in italics.

Sexual differences exist in the relative breadth (BL) only,
as according to previous studies the femalesof Erronea errones
are about one class broader than the males, whereas the
other characters show no sexual differences (SCHILDER &
Houston, 1964, tables 3 and 4).

Vol. 11; No. 2

THE VELIGER

Page 113

LOCAL POLYMORPHISM

The only real mutant concerns the sky-blue absolutely
unspotted variant living in the populations 26 miles
around Broome, but never collected elsewhere (FA.
ScHILDER, 1968a); it lives with usual specimens of
Erronea errones without producing intermediates, and
constitutes 1 to 6% in all populations around Broome.
This morphe has been called azurea ScHILDER (1968b).
All other extreme varieties, e. g. the subrostrate melan-
istic shells (New Caledonia, Pumpkin Island, One ‘Tree
Island, etc) or those with the labial anterior spot pro-
duced along the margin or even rare shells with one or
two posterior terminal spots must be regarded as extreme
individual aberrations connected with the normal speci-
mens by many intermediates. Shells showing the dorsum
suffused with a layer of unspotted pale enamel should be
regarded as pathological, as this accessory layer mostly
includes particles of mud or even parasites, and the usual
markings are shining through; they occur scattered in

Map 1:

many populations, but seem to accumulate in some popu-
lations living under special conditions.

GEOGRAPHICAL DISTRIBUTION
or CHARACTERS

The Maps 1 to 5 illustrate the geographical distribution
of the median classes listed in Table 2; the 6 classes of
each character have been designated by different symbols
progressive in darkness from 1 to 6. Careful examination
of these maps will show the following general trends
which do not exclude occasional exceptions in one or a
few populations.

Map 1: Length (L). The smallest Erronea errones
live along an approximately equatorial zone from the
Andaman Islands to Java, New Guinea, and Fiji; around
this zone such populations are mixed with populations of
larger specimens, viz. in the West (Ceylon), in the North
(north of the line Singapore-Celebes-Palau), and the
South (West Australia, East Australia to New Caledonia

Length

Page 114 THE VELIGER Vol. 11: No. 2

Map 2: Breadth

Map 3: Dorsal Blotch

Wolsll >No. 2 THE VELIGER Page 115

Map 4: Anterior Spots

Map 5: Basal Color

Page 116

and Tonga): this arrangement recalls that observed in
other cowrie species too (KE A. ScurLper, 1961, 1962).
The largest populations live along the Queensland coast
from Mackay to Moreton Bay, while the shells collected at
the offshore islands become smaller by increasing distance
(see also F A. & M. Scuitper, 1964; M. « F A. ScHILper,
1967).

Map 2: Breadth (BL). Mostly narrow populations
live in the area between Bali, Solomon Islands, Japan, and
Thailand; the western populations ranging from Ceylon
to Java are mostly broader, as well as the southern popu-
lations from West Australia to Fiji. It is curious that the
shells coming from the Keppel Bay Islands are narrower
than those from the opposite mainland coast as well as
those from the Capricorn Islands farther away from the
shore.

Map 3: Dorsal blotch (DB). In the great central area
from the Andaman Islands to New Caledonia popula-
tions with rather large blotches are mixed with those
with reduced blotches; in the farthest West (Ceylon)
and the farthest East (Fiji, Tonga) the dorsal blotch is
mostly small to absent, as it seems to be in Micronesia
also, whereas in the South (i.e. both regions of Aus-
tralia) the blotches usually are large.

Map 4: Anterior spots (AS). In West Australia the
terminal spots are mostly absent, and in East Australia
they are small, as well as from the Admiralty Islands to
Fiji and Micronesia; in the central zone, i.e. in Tonga
and New Caledonia as well as from western New Guinea
to Ceylon, these spots are usually well developed to large,
though sporadic populations with reduced spots may
occur, especially in the Andaman Islands.

Map 5: Basal color (BC). In the populations living
in a central zone from Java to the Solomon Islands the
base is whitish; this zone may extend northward to Thai-
land and Japan with few darker populations scattered
among many pale ones. In the South from West Australia
to Fiji and in the West from Ceylon to Singapore yellow
populations are prevalent.

One will observe that in all characters discussed there
are widely regional differences in the predominant classes.
One could surmise that there is usually a rather equatorial
central zone differing from peripheral regions especially
in the West and South, and less markedly also in the East
and North: so one could summarize the predominant
characters as follows:

Central Zone Peripheral Regions

L small large

BL narrow broad

DB various N, W, E: small
S: large

AS large reduced

BC whitish yellowish

However, there is no parallelism between the bound-

THE VELIGER

Vol. 11; No. 2

aries of the regions, because the border lines of the pre-
dominant classes of each character cross each other in
various ways. Therefore no distinct geographical races
can be established by the sum of several characters (EF A.
& M. Scuiwp_er, 1961, p. 305).

Nevertheless, in some restricted areas all or most popu-
lations agree in several characters which fact may be
explained by a common gene pool influenced by selection
in similar environments.

Besides, one will observe that in small islands off-shore,
as Dampier Archipelago, Capricorn Islands, Whitsunday
Islands, Green Island, Mondoure outer reef, Manava
Island, Edam Island, the shells are smaller, narrower,
with less developed anterior spots and paler base than in
the adjacent coastal populations.

LITERATURE CITED

SCHILDER, FRANZ ALFRED
1961. _A statistical study in cowries: the size of Mauritia ara-
bica (LINNAEUS). The Veliger 4 (1): 15-17; 2 text figs.
(1 July 1961)

1962. ‘The size of Cypraca tigris LINNAEUS. The Cowry
1 (3): 43-44 (1 February 1962)
1965. The geographical distribution of cowries. The

Veliger 7 (3): 171-183; figs. 1-3

1968a. An interesting mutation in cowries.
News (n.s.) 98: 5; 1 fig.

(1 January 1965)

Hawaiian Shell
(February 1968)

1968b. Zur Kenntnis der Cypraeidae: 13. Eine Mutation von
Erronea errones (LINNAEUS). Arch. Molluskenk. 97:
(in press)

SCHILDER, FRANZ ALFRED, & MARIA SCHILDER
1938. Prodrome of a monograph on living Cypraeidae.
Proc. Malacol. Soc. London 23 (3/4): 119 - 231; 1 fig.; 9 maps
(15 November 1938 and 15 March, 1939)

1961. Zur Variabilitat der Zeichnungselemente bei Porzellan-
schnecken (Cypraeidae) . Zool. Anz. 167 (7/8) : 303 - 309;
2 diagrams

1964. Latitudinal differences in size of East Australian cowries.
The Cowry 1 (7): 100 - 101 (1 November 1964)
ScHILDER, FRANZ ALFRED, Maria SCHILDER & GARFIELD Houston
1964. The cowrie fauna of Penrith Island. The Veliger
6 (3): 155 - 161; 4 tables; 1 text fig. (1 January 1964)
ScHILDER, Maria
1967. Length, breadth, and dentition in living cowries. The
Veliger 9 (4) : 369 - 376; 1 diagram (1 April 1967)
ScHILDER, Marta & FRANZ ALFRED SCHILDER

1967. Studies on East Australian cowries.
10 (2): 103 - 110; 9 tables

The Veliger
(1 October 1967)

Vol. 11; No. 2

THE VELIGER

Page 117

A New WNeptunea from the Pacific Northwest

ALLYN G. SMITH

Associate Curator
Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118

(Plate 14)

For MANY YEARS the operations of commercial trawlers
along the west coast of North America, particularly off
central and northern California and the Pacific North-
west, have been restricted to moderate depths generally
not exceeding 100 fathoms and usually in the range of
50 to 75 fathoms. In the last ten years, however, due to
offshore fisheries investigations by the California Division
of Fish and Game and similar organizations, excellent
table fish, such as sole and sablefish (black cod), have
been discovered in commercial quantities in much deeper
water.

As a result, trawler captains have been lengthening
their lines and making successful hauls on new fishing
grounds at depths ranging from 100 fathoms to 400 fath-
oms with the possibility of increasing the range to as
deep as 500 fathoms. These deeper-water trawl hauls
started early in 1948, when two otter-trawl captains
worked their nets in 200 fathoms off the northern Cali-
fornia coast and continued to get good results in the 185
to 215 fathom range (Hotmperc, 1948; ScorieLp, 1948).

To conchologists fortunate enough to enlist the co-
operation of trawl fishermen to save some of the deeper-
water mollusks brought up in their nets instead of throw-
ing them back overboard as they usually do, some of
the deeper hauls have turned out to be veritable bonan-
zas. The larger carnivorous mollusks, such as Neptunea
and Beringius, have been turning up more frequently
and this has been true also for many smaller species.

Among other mollusks obtained from trawler captains
in the last several years by Mr. Everett C. Stiles of
Bellingham, Washington, is a new species of Neptunea
taken sparingly in an area from Cape Flattery, Washing-
ton, north to Cape Scott at the northern end of Vancouver
Island, British Columbia.

Neptunea stilesi A. G. Smitu, spec. nov.

(Plate 14, Figures 1 to 7)

General Diagnosis: This Neptunea is of medium size
for the genus and is distinguished mainly by its much

shorter spire compared with other Northwest Pacific
species, such as Neptunea pribiloffensis (Dati, 1919),
N. phoenicea (Dati, 1891), and N. lyrata (GMELIN,
1791). In general shape it is closer to N. beringiana
(Mippenvorrr, 1848), which has a smaller, somewhat
heavier shell and a geographical range much farther to
the north. It has a capacious bodywhorl with an unflared
lip when adult, and a short canal. In color it is off-white
to beige or yellowish and occasionally a reddish-brown.
Major sculpture consists of widely-spaced, moderate to
heavy, rounded spiral chords or ribs, which are usually
darker in color than the ground-color of the shell.

Description of the Holotype: Shell from an adult living
specimen with operculum, of short fusiform shape with
a relatively short spire, a large, evenly-rounded, tumid
bodywhorl that terminates in a capacious aperture, and
a short canal bent slightly to the left and rather sharply
to the rear. Divergence of spire (apical angle) approx-
imately 87°. Nuclear whorls damaged at the tip, about
2 remaining, nearly straight-sided, with a small channel
near the suture and a low chord below it, otherwise
smooth. Postnuclear whorls 44, rounded, and decorated
with a series of evenly-spaced, low, rounded, spiral ribs
of which there are 14 on the bodywhorl and 3 each on
the preceding postnuclear whorls, with a fourth showing
near the suture at the base of the penultimate whorl;
uppermost rib weak, the remaining are stronger. The
spacing of the spiral ribs becomes gradually wider with
the growth of the shell, being 5 to 9 mm apart near their
terminations at the peristome. Spaces between the spiral
ribs slightly concave and decorated with from 2 to 4
fine spiral chords. Transverse sculpture consists of many,
closely-spaced, rather rough lines of growth, overridden
by the fine intercalated spiral chords; it is much
roughened on the canal. Sutures distinct and slightly
channeled; below them the tops of the whorls are slightly
flexed, forming a shallow, encircling channel. Outer lip
fairly thick, blunt-edged, unflared, somewhat crenulated
by the major spiral chords, with a small notch at its
anterior terminus with the bodywhorl. Inner lip a smooth
wash of callus only. Columella sinuate, terminating on

Page 118

THE VELIGER

Vol) Hie Now2

the canal in a well-marked siphonal fasciole that extends
to the tip of the canal. Color of shell cream-white, the
major spiral chords a contrasting red-brown. Aperture
white, porcellaneous. Operculum heavy, normal for the
genus. Periostracum lacking.

Measurements are: length, 93.9; maximum diameter,
64.9; length of aperture (including canal), 71.5; width
of aperture, 31.8; length of canal, about 20 mm.

Type Locality and Range: Because of lack of accurate
data, no specific type locality can be pinpointed. Most
specimens received for study were trawled in depths of
approximately 100 to 125 fathoms in the area bounded,
in general, on the south by La Perouse Bank, 40 miles
west of Cape Flattery, Washington, and on the north by
Cape Scott at the northern tip of Vancouver Island,
British Columbia. A single specimen was dredged in 34
fathoms, Hakai Pass, British Columbia, by Dr. I. McT:
Cowan of the University of British Columbia (UBC No.
1540). Another halfgrown shell was also taken by Dr.
Cowan in about 85 fathoms, Queen Charlotte Sound,
just north of Cape Scott. An old, dead adult specimen in
the Stanford University Collection (Department of Geo-
logy) was dredged in 110 fathoms, Virago Sound, British
Columbia.

Disposition of Specimens: Holotype deposited in the
California Academy of Sciences, Geology Type Collec-
tion, no. 13124.

Twenty-eight paratype shells have been placed in
several institutions, including the California Academy
of Sciences, Stanford University, the University of
British Columbia, Los Angeles County Museum of
Natural History, United States National Museum, and
the Academy of Sciences of the U.S. S. R. (Leningrad) ;
and in the private collection of Everett C. Stiles. Pre-
served animals of 6 paratypes have been deposited in
the California Academy of Sciences, Invertebrate Zoo-
logy Type series, nos. 371 to 376, inclusive.

Remarks: This fine new species of Neptunea is dedi-
cated to Mr. Everett C. Stiles of Bellingham, Washing-
ton, whose diligence in enlisting the interest and cooper-

ation of several Pacific Northwest trawler captains has
resulted in most of the specimens that have been available
to date. Although it is found in the same general area
and at about the same depths as the heavily-ribbed Ber-
ingius eyerdami A. G. SmitH, 1959, it has been taken less
frequently and must be rated as a relatively rare species.

In the type lot of 29 shells, 23 were obtained alive, 17
with opercula, although in most instances the animals
had disintegrated and could not be saved. There is con-
siderable variation in this series of shells in size, sculp-
ture and color. Size limits are shown by the following
measurements:

Dimension Largest Smallest
Length, over-all 116.1mm 69.5 mm
Maximum diameter 72.3mm 48.6mm
Length of aperture and canal 78.2mm 544mm
Width of aperture 35.3mm 26.4mm
Length of canal 26.0 mm 8.3 mm
Number of postnuclear whorls 6 De
Apical angle 96° Je

The most striking sculptural feature is the presence on
typical specimens of prominent spiral ribbing of a red-
brown color against a cream-white background. This
feature is not at all consistent, however. Twenty-eight of
the 29 shells can be ranked as follows in terms of the
presence and prominence of the spiral ribs or chords:

Ribs obsolete or weak 6
Ribs of medium strength J
Ribs fairly prominent 15

The shell texture of full-grown adults is only moder-
ately heavy; in younger specimens the shell is quite thin
and partially translucent. Older shells have a heavy wash
of callus on the inner lip, which makes the peritreme
complete. There is no tendency for the outer lip to flare
with age or senility. While a small anterior notch, though
present, is not especially prominent in the holotype, it

Explanation of Plate 14

Figure 1: Neptunea stilesi A.G. Smiru, spec. nov. Holotype, Calif.
Acad. Sci. Geol. Type coll. no. 13124. Length: 93.9mm; maximum
diameter: 64.9mm; apical angle:83°. Apertural view.

Figure 2: Same, dorsal view.

Figure 3: Enlarged view of the nuclear tip of a subadult paratype
from 85 fms, Queen Charlotte Sound, British Columbia. Length
(nuclear whorls only) : 5.5mm; maximum diameter: about 3mm;
number of nuclear whorls: 24.

Figure 4: Paratype. Brownish color-form with subobsolete spiral
sculpture. Length: 94.3mm; maximum diameter: 65.8mm; apical
angle: 77°.

Figure 5: Paratype with well-developed spiral ribs. Length: 106.0
mm; maximum diameter: 71.7mm; apical angle: 92°.

Figure 6: Paratype with somewhat weaker spiral ribs having fine
intercalaries between them, stronger on the body-whorl. Length:
106.4mmi; maximum diameter: 69.1 mm; apical angle: 78°.

Figure 7: Radula of paratype animal, Calif. Acad. Sci. Inv. Zool.
Type Series no. 371, slide no. 490. Center section, mounted width:

I.I mm.

THE VELIGER, Vol. 11, No. 2 [A. G. Smiry] Plate 14

ie ie 7 ;
| of 7 Hay
aa Po

Vol. 11; No. 2

is quite deep in other shells and can be considered as a
normal character for the species.

Two shells in the series have complete nuclear whorls.
An enlarged view of one of these is shown in Plate 14,
Figure 3. It consists of about 2} smooth whorls with a
length of 5.5 mm and a width of about 3 mm. The tip is
dome-shaped and folded over on to the second nuclear
whorl, which has slightly diverging sides and minutely
channeled sutures. This tip broke off during measurement
in spite of careful handling; fortunately it could be
repaired satisfactorily. This incident developed the fact
that while the tip itself was hollow spirally, the be-
ginning of the first postnuclear whorl had already been
plugged with shelly material as a protective step prepar-
atory to the loss of the nuclear tip from erosion or other
causes. This illustrates why so few Neptunea shells of
this and other species are collected with their nuclear
tips intact.

The normal color-phase is represented in the holotype
(Plate 14, Figures 1, 2), which has relatively prominent
spiral ribs. Another color-phase is dark red-brown, oc-
curring on 4 shells. Plate 14, Figure 4 shows a smoother
form of the latter color although another similar, red-
brown shell has relatively heavy spiral ribs marked with
a still darker color. A shell with the heaviest, most
prominent spiral ribs is shown on Plate 14 in Figure 5,
and another with several fine intercalary spirals between
the ribs on the upper part of the bodywhorl is shown in
Figure 6. The shell apertures are generally pure white
inside; however, two have the columella tinted a deep
orange-brown with a lighter wash inside the outer lip
and in another the entire aperture is colored lavender-
pink, edged with white in the vicinity of the peristome.
Animal and Radula: Of the 6 specimens with preserved
animals, 2 were males and 4 were females. One of each
sex had shells with obsolete ribbing; one male and 3
females had shells with more or less prominent spiral
ribs. From this, and from the size and general configura-
tion of the shells, the possible occurrence of any sexual
dimorphism in this species is not evident.

The animals (in alcohol) are yellowish-cream color,
with occasional black streaks and maculations on the
head and foot.

The radula (Plate 14, Figure 7) is typically neptuneid.
The central tooth is tricuspid, the central cusp being
slightly larger than the other two. The major laterals are
also tricuspid, the innermost pair being close together
and separated by a narrow V-shaped interval, the inner-
most cusp being the larger of the two; this pair is
separated from the third, narrowly elongate, outer cusp

THE VELIGER

Page 119

by a broadly U-shaped interval. Radulae from other
animals are not basically different from the one illustrated
except for the minor differences that might be expected
to occur from age, wear, or individual variation.

Although the available information is somewhat in-

definite at present, the species apparently lives on a
fairly soft mud bottom. Its egg-capsules have not been
identified as yet.
Relationships: Neptunea stilesi has no really close rela-
tives. Its short-spired aspect, tumid bodywhorl, and the
sculptural characters distinguishes it specifically from
other described Neptunea, Recent and fossil. Comparison
with living species has been made with the work of
Go.ikov (1962, 1963), who has discussed and illustrated
those from northern seas in considerable detail; compari-
son with fossil species was made on the basis of unpub-
lished work of Dr. F Stearns MacNeil, formerly of the
United States Geological Survey.

Neptunea stilesi approaches the shape and in some
instances the type of spiral ribbing of N. beringiana
(Mippenporrr, 1848) but the latter has a somewhat
smaller-sized, heavier shell and a range extending to the
north of the Aleutian chain (Goutkov, 1963, fig. 95).

Its relationship with other Neptunea species occurring off

the Pacific Northwest coast is less close as all of them
have long-spired shells. Of these, N. lyrata (GMELIN,
1791) has heavy spiral keels and a range extending
southward into southeast Alaska and northern British
Columbia; N. phoenicia (Dat, 1891), from British Co-
lumbia and Puget Sound, has a dark-brown shell covered
with an olivaceous periostracum and a sculpture of
rather fine spirals devoid of prominent ribs; N. pribi-
loffensis (Dat, 1919), from the Gulf of Alaska to British
Columbia, has more rounded whorls, much deeper su-
tures, and more closely-spaced, less prominent spiral
sculpture; N. amianta (Dati, 1889) has a smaller,
thinner shell, finely lirated spirally but sometimes with
fairly strong spiral keels, and a much deeper habitat
ranging from 400 to 1000 fathoms; N. smirnia (Datu,
1919), which ranges south off Humboldt Bay, California,
has a totally different, larger, smooth, brownish shell
with deeper sutures; N. zthia (DALL, 1891) is of the same
type as N. smirnia but is much smaller and has been
recorded so far with certainty only in Monterey Bay,
California, in 200 to 400 fathoms.

Possibly Neptunea stiles: could be considered to be an
offshoot from N. beringiana stock that has developed in
a new, more southerly region under changed ecological
conditions but this can hardly be more than pure specu-
lation.

Page 120 THE VELIGER Vol. 11; No. 2

ACKNOWLEDGMENTS

Thanks are due to the following associates in the Cali-
fornia Academy of Sciences: Mr. Dustin D. Chivers for
his excellent microscope slide preparations of radulae;
Dr. Victor Zullo for photographing the radula slide used
on Plate 14; and to Mr. Maurice Giles for photographing
the shells and the preparation of the black-and-white
prints used for illustrations.

LITERATURE CITED

Go.tkov, A. N.

1962. New species of gastropods of the genus Neptunea Bot-
TEN from the far-eastern seas of the U.S.S.R. (Gastropoda,
Prosobranchia) . Acad. Sci. USSR, Trudy, Zool. Inst. 30:
1-10; 2 plts.; 2 maps Moscow/Leningrad [in Russian]

1963. The gastropod mollusks of the genus Neptunea BoLTEN.
Acad. Sci. U.S.S.R, Zool. Inst., n.s. no. 85 (Fauna of the U.
S.S.R., Mollusks, vol. 5, issue 1): 1-217; plts. 1-28; figs.

1-97 in text Moscow /Leningrad [in Russian]
HoLMBERG, Epwin K.
1948. Deep dredging by Eureka otter trawlers. Calif.
Fish & Game 34 (4): 218-219 (October 1948)
MacNetm., FE STEaRNS
1957. Cenozoic megafossils of northern Alaska. U. S.

Geol. Surv. Prof: Paper 294-C: 99 - 126; plts. 11-17

ScoFIE.p, W. L.
1948. Trawling gear in California. Calif. Div. Fish &
Game, Fish Bull. 72: 1-60; figs. 1 - 24

Vol. 11; No. 2

THE VELIGER

Page 121

An Investigation of the Commensals

of Cryptochiton stelleri (MIDDENDORFF, 1846)

in the Monterey Peninsula Area, California

BY

STEVEN K. WEBSTER

Department of Biological Sciences, Stanford University, Stanford, California 94305

(5 Text figures)

INTRODUCTION

THE GIANT CHITON or THE Pacific Coast, Cryptochiton
steller. (Mippenporrr, 1847), occasionally attains a
length of 35cm, and often harbors commensals in its
pallial grooves. A pea crab, Opisthopus transversus
(RaTHBUN, 1893), and a polynoid worm, Arctonoe vit-
tata (GruBE, 1855), are frequently found in the pallial
grooves of C. stelleri in the Monterey Peninsula area.
These grooves, each with a row of about 70 ctenidia,
are bordered on the inside by the dorso-lateral edges of
the foot and on the outside by the girdle margins.

While Arctonoe vittata may be found free-living under
rocks, Opisthopus transversus is always found as a com-
mensal. Both O. transversus and A. vittata are found as
commensals in other hosts. For A. vittata some of these
are Solaster, Dermasterias, Pisaster, Henricia, Crossaster
(asteroids); Stichopus (a holothurian); Diodora, Acmaea,
and Puncturella (gastropods). Opisthopus transversus is
also found with Megathura, Aplysia, and Astraea (gas-
tropods) ; Mytilus, Tresus, and the siphons of pholads
(pelecypods) ; and Stichopus (HARTMAN & Retsu, 1950;
Davenport, 1950; Ricketts & Cavin, 1962; BEonpé£,
1968). A more complete list of hosts for O. transversus
can be obtained in BEonpé (1968).

The primary objective of this investigation was to
determine the incidence and distribution of the com-
mensals on Cryptochiton in the Monterey Peninsula area.
In addition, observations were made concerning the nat-
ural history of Cryptochiton, and experiments on the role
of diffusible attractants to commensals produced by
Cryptochiton were conducted.

FIELD METHODS

Two subtidal sites were chosen within which dives were
made using SCUBA equipment. The two stations were:
Cabrillo Point, Monterey Bay (121°54’10” W; 36°37’30”
N) and San Jose Beach (121°55’30” W; 36°30’50” N)
on more exposed coast 2 miles S of Carmel. Both sites
contained granite boulders surrounded by granite gravel
and shell fragments. Boulders extended to a depth of
20m at Cabrillo Point and to over 50m at San Jose
Beach. These sites were chosen for their accessibility and
because they represented contrasting conditions of ex-
posure. While the largest number of dives was made
between October, 1966 and September, 1967 (20 at each
site), several dives were conducted in June and July,
1965. The data from both periods have been included in
the discussion. A single dive was made in August, 1965,
on the shale beds 4 mile offshore at Del Monte Beach,
Monterey Bay.

Dives were of about 45 minutes’ duration and involved
the use of either rectangular or zig-zag swimming pat-
terns in order to avoid repeated encounters with the same
chitons. Each chiton was removed from the substrate and
turned over to expose the pallial groove. When the chiton
began to curl and close the pallial grooves, a finger was
inserted in the anterior end of the groove and moved pos-
teriorly, exposing the groove and commensals. Specimens
which curled so tightly as to preclude thorough examina-
tion were not counted. Notes on the numbers of Crypto-
chiton and commensals observed were recorded on an
underwater slate. The incidence of juvenile Arctonoe (less
than 3cm) was recorded separately. Additional obser-

Page 122

vations (physical conditions, behavior, etc.) were record-
ed shortly after each dive.

RESULTS or FIELD WORK

Although chitons were observed most frequently on
granite boulders, they were occasionally seen on gravel
within a meter or so of large boulders. In addition, C’ryp-
tochiton was found in both stations on Diapatra ornata
(Moore, 1911), a polychaete tube-worm, apparently to
browse on red algae found in association with the worm
colonies. Cryptochiton stelleri in both sites was distributed
more or less uniformly at depths of about 4 to 40m. An
occasional aggregation of 3 or 4 specimens was observed.

Most of the chitons were between 20 and 35cm
long. Young specimens of 12 to 20cm were common in
some areas. Smaller individuals were rare but are more
common _intertidally. Cryptochitons were observed
browsing red and brown algae common to these rocky
subtidal areas. Gigartina, Iridaea, small Laminaria,
Ulva, and Macrocystis are common foods (TucKER &
Giese, 1962). I also found them feeding on Plocamium
and various coralline red algae. Although RickETTS &
Carvin (1962) mention that feeding occurs primarily at
night intertidally, numerous individuals were observed in
both subtidal stations feeding actively during daylight
hours.

1) Incidence of Commensals on Cryptochiton
(Figures 1 to 4)

While Arctonoe occurred on Cryptochiton at both San
Jose Beach and Cabrillo Point, Opisthopus was found
only at Cabrillo Point and on the shale beds opposite Del

100

3 Cabrillo Point 1966-1967
FI San Jose Beach 1966-1967
: 80 Cabrillo Point 1965

8 “6 San Jose Beach 1965

fj S 97

“= $ 60

mire

3 40

S

3 20

ws

Figure 1

Numbers adjacent to points on the graph represent the numbers of
observations from which the % of incidence of commensals on
Cryptochiton was calculated.

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Vol. 11; No. 2

Monte Beach. It appears, therefore, that Opisthopus is
confined to Monterey Bay.

The incidence of Arctonoe (Figure 1) reached a max-
imum of nearly 60% at both stations between October
and December. The minimum incidence of 22 to 25%
occurred in both stations between May and August.
Juvenile Arctonoe were most abundant in August and
September (Figure 2), just preceding the period of
highest adult incidence. These data indicate that a high
yearly turnover rate may be occurring within the Arc-
tonoe population.

iS)
on

Cabrillo Point 1966-1967
San Jose Beach 1966-1967

20

% of C. stelleri with commensal
A. vittata juveniles (<3 cm long)
a

S -O'N D JF M "Aes
Months

Figure 2

Numbers adjacent to points on the graph represent the numbers of
observations from which the % of incidence of commensals on
Cryptochiton was calculated.

The incidence of Opisthopus (Figure 3) fluctuated
somewhat throughout the year but was considerably
more stable than that of Arctonoe. The incidence of
Opisthopus at the Cabrillo Point station averaged be-
tween 20 and 40% throughout the year. The incidence
of Opisthopus in the shale beds during August, however,
was 90%. The occurrence of more than one commensal
Opisthopus on a single chiton was common in this area.
Two chitons harbored 2 Opisthopus each (both in the
same pallial groove), and one chiton contained 3 (2 in
one groove, one in the other). The high incidence of
Opisthopus might be a result of the abundance of rock-
boring clams (Pholadidae), whose siphons may provide
space for large numbers of Opzsthopus. Encounters be-
tween Opisthopus and potential host chitons seem more
likely in this area than at Cabrillo Point where potential
Opisthopus hosts are less dense.

While C’ryptochiton was often found to harbor 2 or
even 3 Opisthopus at one time, chitons were never found
to harbor more than one Arctonoe. Several Cryptochitons
observed at Cabrillo Point contained both Arctonoe and

Vol. 11; No. 2

100

Cabrillo Point 1966-1967
Cabrillo Point 1965

80

% of C. stelleri with commensal
O. transversus

‘Sm One Nae Die) hae Me Av Many ay vA
Months

Figure 3
Numbers adjacent to points on the graph represent the numbers of
observations from which the % of incidence of commensals on
Cryptochiton was calculated.

Opisthopus, one of each in opposite pallial grooves.
These combinations were most abundant (18$% inci-
dence) in December, coinciding with the period of
maximum occurrence of Arctonoe. The incidence of
Opisthopus alone is average (31.6%) during December.
This appears to be one of the few instances where more
than one species of commensal is found at the same
time on one host.

25

a Cabrillo Point 1966-1967
& 3
=
aS
os

Q
sO
es
‘2 8
8
3 &
us
wN
[o} 2
IN

SHOMNED IN|

1 ME AN OME AN
Months

Figure 4

Numbers adjacent to points on the graph represent the numbers of
observations from which the % of incidence of commensals on
Cryptochiton was calculated.

2) Position and Orientation of Commensals

Arctonoe were usually found wholly within and toward
the anterior end of the pallial groove, facing anteriorly.

THE VELIGER

Page 123

The worms were usually clinging to the dorso-lateral
surface of the grooves or to the ventral surfaces of the
ctenidia. In either case, the worm is upside-down in
relation to the Cryptochiton and the substrate. Occasion-
ally, Arctonoe was found clinging to the ventral surface
of the foot of Cryptochiton, or partially within the
groove and partially on the foot. In such cases the worm
also faces anteriorly and is upside-down.

Opisthopus was always found within the pallial groove,
any place throughout its length; they were either upside-
down or right-side-up, but always faced the midline of
the host.

The crabs were often found to have varying amounts
of mucus adhering to them, which presumably originates
from the pallial mucous tracts on the inner walls of the
pallial grooves. Mucus was rarely found adhering to
Arctonoe.

DISCUSSION

By labeling the relationships between Arctonoe, Opistho-
pus and C’ryptochiton as commensal ones, it is assumed
that the commensals derive some benefit from the host
at its expense, without harming it. It is evident that the
pallial grooves of the Cryptochiton provide shelter for
the commensals that is well protected from wave surge,
predators, and desiccation (during rare periods of inter-
tidal exposure). The respiratory currents in the pallial
grooves also provide the commensal with a nearly con-
tinuous supply of sea water containing plankton and bits
of detritus, a ready supply of food. Gut analyses of
Arctonoe indicated that they are detritus feeders. That
they always face the inhalant stream and are never
observed crawling on the substrate would seem to support
this interpretation. The frequently observed association
of Opisthopus with the pallial mucus might indicate
that the mucus and its contents serve as food for the crab.

Both Arctonoe and Opisthopus produce planktonic
larvae. It is possible that the initial contact of these com-
mensals with Cryptochiton occurs when they are in the
plankton and that they are carried into the pallial grooves
via the inhalant respiratory current. Settling and meta-
morphosis might then occur within the pallial grooves.
It is also possible that both commensals enter the C'rypto-
chiton as metamorphosed juveniles or as adults. An un-
disturbed Cryptochiton lifts the margin of the girdle
both anteriorly and posteriorly to form the inhalant and
exhalant openings to the pallial grooves. These raised
portions of the girdle would provide easy access for com-
mensals. The smallest of the commensal organisms ob-
served in Cryptochiton were already adults. Additional
study of this question is warranted.

Page 124

LABORATORY STUDIES

Introduction and Methods

The presence of substances which serve as physiological
attractants of the host for its commensals has been dem-
onstrated by Davenport (1950) and DaveNpoRT &
Hicxockx (1951) in the Arctonoe fragilis — Evasterias
association. Using an apparatus similar to the one used
by these authors, the Arctonoe vittata — Cryptochiton
stelleri relationship was investigated for the presence or
absence of such an attractant. A Y-shaped tube was con-
structed of 1/16 inch plexiglass, with inside dimensions
of 24 by 24cm. The stem was 20cm long and the two
arms were 10 cm each. Each arm of the Y-tube was con-
nected to two plywood tanks each with a volume of one
cubic foot. Pinchcocks were installed to control the flow
of water from the tanks to each arm of the Y-tube such
that water could be directed from either tank to either
arm of the tube (Figure 5). Each tank contained sea

\Y . By
“Point of Choice”

Ye Cork “A”

Pinch Clamp

ty & Stopcock

Figure 5

water with a temperature between 15.8 and 17.0° C.
Sea water was added to maintain the two tanks at equal
levels. Between trials, sea water was run through all
tubing. The flow characteristics of the Y-tube were

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Vol. 11; No. 2

checked with dye, indicating that a definite side-to-side
concentration gradient occurred at the “point of choice”
where water from each arm of the Y-tube converged.

A Cryptochiton without commensals was placed in one
tank of sea water, while the second tank contained sea
water only. The appropriate pinchcocks were opened to
allow for flow from the two tanks into opposite arms of
the Y-tube. When the stem of the Y-tube was filled,
cork “A” was removed and a commensal inserted. The
cork was then replaced. The speed of flow was adjusted
so as not to wash the commensal backward. Flow of
water through the arms of the Y-tube was alternated
from one trial to the next.

If after 10 minutes the commensal failed to enter
either arm of the Y-tube, a failure was recorded but not
counted. Entrance of the commensal into the plain sea
water arm was counted as “negative.” Movement into
the arm containing water from the tank with the Crypto-
chiton was considered “positive.”

RESULTS
Arctonoe Opisthopus

Number of trials 45 45
Number of failures 5 1
Number and % of

movements into

the Cryptochiton arm 34 (85%) 33 (75%)
Number and % of

movements into

sea water arm 6 (15%) Il (25%)
P >0.01 0.05 to 0.01

DISCUSSION

Two other experiments were performed, for which data
are not presented in detail.

Both Opisthopus transversus and Arctonoe vittata, or
A. vittata - A. pulchra (JouNson, 1897) “intergrades”
inhabit the cloacal chamber of Stichopus californicus
(Strmeson, 1857), a holothurian common in both sta-
tions (Davenport, 1950). An experiment was run in
which Stichopus (free of commensals) was substituted
for the Cryptochiton and tests made on Arctonoe and
Opisthopus collected from the pallial grooves of Crypto-
chiton. In 31 trials, a considerably larger number of
“failures” occurred with Stichopus than with Crypto-
chiton in the previous experiment, and the runs into the
two arms of the Y-tube approached randomness.

Vol. 11; No. 2

‘THE VELIGER

Page 125

An experiment was conducted with Stichopus califor-
nicus in one tank and Cryptochiton stelleri in the other.
Arctonoe vittata and Opisthopus transversus collected
from Cryptochiton were tested. Thirty trials were made
with each commensal. The results were essentially identi-
cal with those of the first experiment, with both com-
mensals showing a preference for the arm containing
water from the tank with Cryptochiton.

Although the above data are based on too few trials
to be conclusive, I believe the presence of a diffusible
chemical serving as an attractant produced by Crypto-
chiton stelleri is indicated. It seems possible that Sticho-
pus produces a diffusible attractant which operates as
such only on commensals previously conditioned to it
in their development. Such an adaptation might be
entirely independent of genetic factors (DAVENPoRT,
1950).

ACKNOWLEDGMENTS

The author wishes to express his appreciation to Dr.
Donaid P. Abbott, Dr. Myra Keen, and Eugene V. Coan
for their helpful suggestions in the preparation of the
manuscript.

The services of George W. Ackerman and Don G.

Tennyson as diving companions are also gratefully
acknowledged.

LITERATURE CITED

BEonbE, ANTHON CRAIG
1968. Aplysia vaccaria, a new host for the pinnotherid crab,
Opisthopus transversus. The Veliger 10 (4): 375 - 378; 2
text figs. (1 April 1968)
DavENPORT, DEMOREST
1950. Studies in the physiology of commensalism. 1. The poly-
noid genus, Arctonoe. Biol. Bull. 98 (2): 81-93
Davenport, DEMorEsT & JoHN FE. Hicxock
1952. Studies in the physiology of commensalism. 2. The poly-
noid genera Arctonoe and Halosydna. Biol. Bull. 100 (2):
71 - 83
Hartman, Orca & Donatp J. ReIsH
1950. The marine annelids of Oregon. Oregon State
Monographs, Studies in Zoology 6: vit 64 pp.; 1 plt.; 3 maps
Oregon State College, Corvallis (August 1950)
Ricketts, Epwarp FE « Jack Cavin
1962. Between Pacific tides. 34 ed., rev. by J. W. Hepcretu
xill+502 pp.; illust. Stanford Univ. Press, Stanford, Calif.
Tucker, JouHn S. & ArrHuR C. GiEsE

1962. The reproductive cycles of Cryptochiton stelleri (Mi-
DENDORFF). Journ. Exper. Zool. 150 (1): 33 - 43

Page 126

Walls WIEUIGER

Vol. 11; No. 2

Notes on the Habitat and Anatomy of Fouannetia quillingi

from North Carolina Coastal Waters

DOUGLAS A. WOLFE

Bureau of Commercial Fisheries, Radiobiological Laboratory
Beaufort, North Carolina 28516

(Plate 15; 1 Textfigure)

Jouannetia quillingi 1S A RARE BIVALVE which I have
recently discovered burrowed into pieces of marl and
sandstone submerged in coastal waters near Beaufort,
North Carolina. Before describing Jowannetia (Pholadop-
sis) quillingt (Pholadidae; Jouannetiinae) as a new spe-
cies, TURNER (1955) examined (among others) 2 speci-
mens collected in 1885 on the Albatross cruises southeast
of Cape Fear, North Carolina. The specimens came from
depths of 17 and 18 fathoms, but no records were avail-
able on whether the specimens were living or dead, or on
the substratum in which they were found. Specimens of
J. quillingi are rare, and the species has not been reported
from elsewhere off North Carolina, even though several
faunal surveys have been conducted in the area (CERA-
ME-VIVAS & Gray, 1966; Menzies et al., 1966; WELLS,
WELLS, & Gray, 1964; Pearse & WiLuiaMs, 1951). The
soft parts of this species were heretofore unknown.
Throughout 1966 and in early 1967, several boats from
Beaufort, North Carolina, regularly trawled for calico
scallops in 10 to 15 fathoms just south of Beaufort Inlet
and west of Cape Lookout (approximately 34°20-32’ N;
76°35-55' W). The bottom in this area consists mainly of
sand and shell fragments, but large pieces of Trent marl,
coquina and coral (Siderastea elegans) were encountered
occasionally in the trawls. I periodically met the trawlers
at the dock to collect specimens from the catch as it was
unloaded. On January 23, 1967, I found several large
pieces of Trent marl which had been dumped beside the
dock after the previous day’s work. Nine dead, but other-
wise intact, specimens of Jowannetia quillingi were re-
trieved from their tight burrows in one piece (about 10
in. x 7 in. x 24 in.); 2 others in the same rock were dis-
carded because they were badly damaged. No more spe-
cimens were found in other similar pieces of rock in the
same pile. The clams had been out of the water at least
27 hours, but no more than 36 hours; the tissues were

still quite moist, but were probably in early stages of
decomposition. Seven of the 9 specimens were preserved
in 40% methanol containing 0.1 molar Na-CO;; the
others were cleaned and dried, one in situ in its burrow
(No. 1253; Plate 15, Figure 1 a). Other specimens (about
20 in all) were collected from burrows in similar rocks
trawled from the same area on January 17, March 2,
March 3, and May 12, 1967 (No. 1544; Plate 15, Figure
1b), but only one of these (No. 1366; March 2, 1967)
contained the soft parts; it was preserved as described
above. Several small immature specimens were also in-
cluded in the March 2 sample. A representative selection
of immature and adult specimens, including the one
illustrated in Plate 15, Figure 1 b, now is in the collection
at the Museum of Comparative Zoology, Harvard Uni-
versity.

HABITAT anp ECOLOGY

Substratum: All specimens of Jouannetia quillingi were
burrowed into the tops or sides of pieces of soft rock
obtained from depths of 10 to 15 fathoms. The rock usu-
ally was marl (limestone) from the Trent Formation,
which outcrops at several points along the continental
margin of the Carolinas (PEARSE & WitLams, 1951). In
one such piece of marl, recent burrows and shells of J.
quillingt were found among a concentration of lower
Miocene fossil casts of Venus gardneri Kettum. On
March 3, 1967, J. quilling: were found in a softer piece
of rock (Plate 15, Figure 2), which seemingly consisted
of cemented sand grains, more nearly characteristic of
the formations of coquina in this area (WELLS & RicH-
ARDS, 1962; Menzigs et al., 1966). The shell of J. quil-
lingi completely fills the lower part of the burrow, and
the callum of the left valve in the adult is loosely cemen-
ted against the bottom and sides of the burrow so that

Vol. 11; No. 2

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Page 127

only the right valve is free to move (Plate 15, Figure 1).
The callum, which is so extensive in the non-burrowing
adult, is completely lacking in young burrowing speci-
mens. Jouannetia, like other callum-building pholads
(Turner, 1954), probably extends its foot through the
anterior gape of the shell while burrowing, but when
burrowing ceases, the callum is built and the foot is

resorbed. The siphons must be extended a few millimeters
behind the siphonoplax to reach the narrow (2 to 4mm
diameter) aperture of the burrow (Plate 15, Figure 2).
The entrances to the burrows are extremely inconspicuous
from the outside; the posterior end of the siphonoplax
of the entrenched mollusk is generally not visible through
the aperture. Jouannetia quillingi’s apparent preference

Figure 1

Internal morphology of Jouannetia quilling:

A: Specimen collected January 23, 1967, with left valve removed.
C: Different

B: Same specimen, with left mantle laid back.

specimen from same sample, left mantle laid back.
collected March 2, left mantle laid back.

D: Specimen

Explanations of numbers (after TuRNER, 1962)

. Anterior adductor muscle

. Thickened anterior margin of mantle
Mantle

Shell

. Posterior adductor muscle

. Excurrent siphon

. Siphonoplax

. Siphonal retractor muscle

. Incurrent siphon

KHSOoOomrgaopre

—a—

12. Labial palps

13. Foot

14. Visceral mass (abdominal mass of FiscHeEr, 1860, 1862)
15. Inner demibranch of gill

16. Outer demibranch of gill

18. Closed mantle of adult

19. Edge of calcareous callum

20. Brownish digestive tissue

21. Retracted siphons

22. Cut edge of mantle

Page 128

for substrata of soft rock is characteristic of other species
of the genus, although the holotype of J. quillingi was
reportedly found insubmerged and rotted wood (TuRNER,
955)

Associated Mollusks: Principal mollusks found in con-
junction with Jouannetia quillingi on and in similar
pieces of rock taken in hauls made for calico scallops
are listed below:

GASTROPODA

Vermicularia spirata Puiipri, 1836
Vermicularia knorri (DESHAYES, 1843)

PELECYPODA

Lithophaga bisulcata (OrBicNy, 1842)
Lithophaga aristata (Dittwyn, 1817)
Gastrochaena hians GmMeE.in, 1791
Gastrochaena ovata SOWERBY, 1834
Petricola typica (Jonas, 1844)
Hiatella arctica (LINNAEUS, 1767)
Pododesmus rudis (BropErip, 1834)
Diplodonta punctata (Say, 1822)
Chama macerophylla GmMeun, 1791
Chama congregata Conrab, 1833
Pseudochama radians (LAMaRCK, 1819)
Arca imbricata BruGuizRE, 1789

ANATOMY oF Jouannetia quillingi

Three preserved specimens were dissected: two from the
January sample and the one collected on March 2. The
tissues of the former specimens were very soft and dis-
integrated upon extensive probing, and the tissues in the
latter were distinguished much more readily. General
internal morphology of all 3 specimens is shown in Text
figure 1. For convenience, the numbers identifying ana-
tomical features are the same, where possible, as those
used by TurNeErR (1962).

Upon removing the left valve with its overlapping cal-
lum, one sees only the adductor muscles and the closed
mantle of the organism (Text figure 1 A). The mantle is
thickened along the anterior margin and also posteriorly
where it joins the siphonal retractor muscles. In the 3

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Vol. 11; No. 2

specimens dissected, the siphons were fixed in different
positions, ranging from fully extended (Text figure 1 D)
to completely retracted (Text figure 1 B). The siphons are
of equal length and are covered with a rather heavy
brown periostracal sheath.

When the mantle is opened and laid aside (Text fig-
ures 1 A to 1 D), the most prominent anatomical feature
is the large, white, smooth, rounded body of the retracted
siphons (No. 21).When the siphons are fully retracted
(Text figure 1 A, 1B), they occupy most of the space en-
closed by the mantle and thereby displace the visceral
mass anteriorly and dorsally. The visceral mass is differ-
entiated into a cream-colored ventral portion (No. 14) and
a more dorsal brownish digestive gland which was green-
ish in the March 2 specimen (No. 20).The paired labial
palps are long and strap-like and lie on either side of the
slit-like mouth on the antero-dorsal surface of the cream-
colored portion of the visceral mass. The labial palps were
overlooked during the dissection of the first 2 specimens
because of the condition of the tissues, but I have included
them in their proper position in Text figures 1 B and 1 C.
The 2 paired demibranchs of the gill are situated mainly
dorsal and posterior to the visceral mass and the retrac-
ted siphons.

The anatomy of Jouannetia quilling: differs somewhat
from that of J. cumingi and J. globulosa as illustrated by
Fiscuer, 1860, 1862). [FiscHEr’s original drawings are
reproduced in part by TurNeER, 1962.] FiscHER pictures
the siphons of both J. cwmingi and J. globulosa as being
very much shorter than those of J. quillingi, so that the
mantle cavity is occupied by only the visceral mass, labial
palps, and gills. FiscHeR also does not mention any dif
ferentiation of the visceral mass, a feature quite notice-
able in my 3 dissections of J. quilling:. According to Tur-
NER (1954, 1962), a detailed description of the anatomy
of J. cuming: SowErsy is given also in a private publica-
tion by Eccrr (1887), but I have been unable to examine
this reference.

SUMMARY

Jouannetia quillingi Turner, 1955, first found off the
coast of North Carolina during the Albatross cruises of

Explanation of Plate 15

Jouannetia quillingi from 10 to 15 fathoms off Beaufort,
North Carolina

Figure 1 a: Dorsal view of specimen in situ in shallow burrow in
Trent marl. The original entrance of the burrow was destroyed
during removal of excess rock.

Figure 1 b: View of left valve of deeply embedded specimen. The

upper one-half of the burrow has been removed.

Figure 2: Typical burrow of Jouannetia quillingi in soft cemented
sandstone, sectioned to show the shape and proportions of the
burrow.

Tue VELIGER, Vol. 11, No. 2 [Wo FE] Plate 15

Figure 1b

es

te.

——>» 1966.

Vol. 11; No. 2

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Page 129

1885, has been rediscovered south of Beaufort Inlet in
10 to 15 fathoms. The species lives burrowed into soft
marl and sandstone. It seems likely that the rarity ascribed
to the species arises at least partly from the seclusion
afforded by its inconspicuous burrows in submerged rock.
The general internal morphology of the species, hereto-
fore unknown, has been described.

ACKNOWLEDGMENTS

The Radiobiological Laboratory is supported by a co-
operative agreement between the Atomic Energy Com-
mission and the Bureau of Commercial Fisheries. Photo-
graphs are by Twyla Miner and Curtis Lewis of this
laboratory. I thank Dr. Ruth Turner for her comments
and suggestions on the manuscript.

LITERATURE CITED
\
CrraME-Vivas, M. J. « I. E. Gray
The distributional pattern of benthic invertebrates of
the Continental Shelf off North Carolina. Ecology 47 (2):
— > 260 - 270 ; ng ; A)
Eccerr, E.
1887. Jouannetia cumingii Sow.
baden, pp. 1 - 69; plts. 1-3

Privately published, Wies-

FiscHer, PAUL
1860. Etudes sur les pholades.
337-351; plt. 15

1862. Note sur l’animal du Jouannetia cumingi, suivie de la
description de deux espéces nouvelles du méme genre. Journ.
de Conchyl. 10: 371 - 377; plt. 15

Menzigs, R. J., O.H. Pirxey, B. W. BLackweLpEr, D. DExTER,
P Hutine, & L. M. McCLosKeEy
1966. A submerged reef off North Carolina.
ges. Hydrobiol. 51 (3): 393 - 431
Pearse, A.S. & L. G. WILLIAMS
1951. The biota of the reefs off the Carolinas.
Mitchell Sci. Soc. 67: 133 - 161
Turner, RutH Dixon
1954. The family Pholadidae in the Western Atlantic and
Eastern Pacific. Part I — Pholadinae. Johnsonia 3 (33):
1-63; plts. 1 - 34
1955. The family Pholadidae in the Western Atlantic and
Eastern Pacific. Part II - Martesiinae, Jouannetiinae and
Xylophaginae. Johnsonia 3 (34): 65-160; plts. 35 - 93.

1962. Nettastomella japonica Yokoyama in North America
and notes on the Pholadidae. Occas. Pap. Mollusks Harvard
Univ. 2 (28): 289 - 308

WELLS, H. W. & H. G. RicHarps

1962. Invertebrate fauna of coquina from the Cape Hatteras

region. Journ. Paleont. 36 (3): 586 - 591

WELLS, H. W, M. J. WELts « I. E. Gray
1964. The calico scallop community in North Carolina.
Bull. Marine Sci. Gulf Carib. 14 (4): 561 - 593

Journ. de Conchyl. 8:

Int. Rev. d.

Journ. Elisha

Page 130

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Vol. 11; No. 2

Some Observations on the Ecology and Behavior

of Lucapinella callomarginata

RICHARD L. MILLER

Department of Zoology, Oregon State University, Corvallis, Oregon 97331

and the Kerckhoff Marine Laboratory, Corona del Mar, California 92625 '

(Plate 16; 1 Text figure)

INTRODUCTION

McLean (1967) 1n HIS REVISION of the West American
species of Lucapinella reported Lucapinella callomarg-
inata (Dati, 1871) from the intertidal: “on the under-
side of rocks and on pilings near aggregations of Mytilus
edulis in bays and channels in southern California. It has
not been collected in the sub-littoral zone.” No report
other than this exists as to the habitat of this species, nor
is there any description of the general biology of this
organism. During the late summer of 1967, while dredg-
ing in the main channel leading to upper Newport Bay,
California, I found 2 L. callomarginata in a radically
different niche — the mud-flat sponge Tetilla mutabilis
DE LAUBENFELS, 1930. Subsequent collections and labo-
ratory observations have served to confirm and extend
this finding.

COLLECTIONS
AND PRELIMINARY OBSERVATIONS

Dredging runs, made parallel to the northern shore of
the channel about 300 yards east of the Route 101 bridge
in about 10 to 15 feet of water, yielded a large number
of Tetilla and 2 limpets were found tightly embedded in
one of them. The importance of this find was not realized
at the time, so no further search was made for limpets.
The 2 limpets were brought back to the laboratory for
identification and kept isolated in a running sea-watcr
aquarium for 2 weeks before a fresh sponge was pro-
vided. The animals moved under the sponge and burrowed
up and into it.

‘ Current address: Department of Biology, Temple University,

Philadelphia, Pennsylvania 19122

A second collection was made some weeks later at the
same location by raking sponges by hand and a total of
9 limpets was recovered from 15 sponges. In most cases,
if one limpet was found in a sponge, others would be
present. Three limpets were placed on one of the sponges
brought back to the laboratory at this time and it was
used for the photographs in Plate 16.

A third collection was made at a small marina just east
of the bridge, where, at low tide, sponges could be raked
by hand in from 3 to 5 feet of water. Both Tetilla and a
massive, branched, yellow sponge (tentatively identified
as a species of Halichondria) were found, but there were
only 3 limpets in about 30 specimens of Tetilla. All of
these were found in one sponge. This particular Tetilla
was highly irregular in form with 4 large oscula, whereas
the vast majority of the others was more regular, with
a single central osculum (Plate 16, Figure 5). The Hali-
chondria (?) contained no limpets.

A fourth collection was made from the pilings under
the bridge in an attempt to recover animals in the habitat
described by McLean. A single limpet was found here
away from any sponge. This animal was brought back to
the laboratory alive for analysis of the contents of the
fecal pellets.

EXTERNAL MORPHOLOGY

oF Lucapinella callomarginata

Figures 1 to 4, Plate 16, are photographs taken of a
representative specimen of Lucapinella callomarginata.
Figure 1 is a dorsal view, showing the highly sculptured
shell, the fringed mantle, the excurrent siphon set off by
an anterior and posterior set of 6 or more large, fleshy
papillae, and the massive posterior portion of the foot.
Figure 2 shows the same animal from the side. The ante-

Vol. 11; No. 2

rior end is to the right in the figure. The fringed mantle
extends just over the edge of the shell and down over the
foot for a distance of 2 to 4mm. The portion of the
mantle shielding the head is considerably broader than
the portion that extends over the foot. Each of the
mantle papillae on the upper portion of the flap appears
to line up with a shell ridge. Many times two anterior
portions of the mantle are held up, forming fan-shaped
openings on either side of the head at about the position
of the tentacles. This can be seen from above in the
upper lefthand animal in Figure 4, Plate 16 and is prob-
ably used as a method of increasing the incurrent water
flow. The excurrent siphon is extendable to at least 5mm
above the shell surface at the aperture. An animal with
its siphon extended is shown in Plate 16, Figure 6. The
shell of this animal has been described (McLean, 1967).
Figure 3, Plate 16, shows an animal resting in the lower
corner of an aquarium. The lower mantle flap has been
lifted, displaying the proboscis, head tentacle, and a line
of epipodial tentacles decreasing in size posteriorly. The
obvious fold in the foot is not a permanent structure.
Figure 4, Plate 16, shows 6 animals found during one of
the collecting trips and demonstrates the variability in
shell and body color and the differences in spot patterns
on the surface of the foot. In general, the over-all color
of the flesh is an orange-brown to orange-red. The larger
scattered spots on the foot range from darker shades of
brown to black. The brown or reddish color tended to
fade when the animals were kept away from sponges.

FEEDING BEHAVIOR

General observations were made of the feeding behavior
of Lucapinella callomarginata. The animals appear to
attack Tetilla anywhere along the sides but seem to prefer
the area around the base where the sponge is attached to
the mud (Plate 16, Figure 6). The limpet grazes on the
surface and forms a cavity which is eventually enlarged
to encompass the entire foot of the animal (Plate 16,
Figure 8). The limpet crawls forward, burying itself in
the sponge, evidently eating its way through. It may be
covered by subsequent growth of the sponge around and
over it (Plate 16, Figures 5, 6). As the limpet moves for-
ward in the sponge, it creates a groove, one of which was
4 inches long. In 2 cases, limpets were seen to enter the
sponge by way of the large osculum and commence
feeding while in it. No experiments were done to see if
the limpets were attracted to the sponge.

The fecal pellets of limpets grazing on Tetilla were col-
lected and examined. They contained sponge spicules and
unidentifiable debris. Several of the animals were then
starved for a few days and presented a number of sponges

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Page 131

in isolation to determine if Tetilla was the sole food. In
each situation fecal pellets were collected and a search
was made for sponge spicules. The types found in the
feces were compared with the spicules obtained from the
sponge on which the animal was feeding. Text figure 1
summarizes the results when limpcts were exposed to the
sponges Tetilla mutabilis, Hymeniacidon synapium DE
LAUBENFELS, 1930 (Plate 16, Figure 10) and the speci-
men of Halichondria (?) (Plate 16, Figure 9) mentioned
earlier. As the figure shows, the limpets attacked all 3
sponges, and in each case the spicules in the feces, though
somewhat broken up, matched those of the sponge endo-
some. Also shown (G) are the spicules found in the fecal
pellets of the one limpet taken from the pilings. This
assemblage, not found in any of the 3 sponges tested, is
probably a conglomerate. The tylostyle is probably from
a sponge in the family Clionidae while the prodiaene may
be from the ectosome of Tetilla (pE LAUBENFELS, 1932).
Indications are, then, that sponges of the order Demo-
spongia are eaten and the limpets do not restrict their
diet to a single genus. It is also evident that the limpets
can do severe damage to the sponges they attack (Plate 16,
Figure 9).

Since the limpets are often found in areas away from
sponges (MacGinitie, personal communication), it is pos-
sible that these animals are grazers as well as predators.
During a series of observations of isolated limpets it was
noticed that portions of the surfaces of the aquarium
had been cleaned of the coating of debris. Eventually,
one limpet was observed through the glass wall of an
aquarium to be grazing on the surface. In contrast to
Tegula or Haliotis, the stroke of the radula was very
short, which might be an indication that it is designed
more for tearing than for scraping. Confirming evidence
of feeding on algae was obtained from the feces of the
limpet found on the bridge piling. Fragments of plant
material were seen along with the spicules. No diatom
tests were noted, though there was much unidentifiable
debris present.

Although the animals may remain in one Tetilla for a
considerable period of time, they must eventually exhaust
the resources of their prey and have to find another
sponge. Therefore, observations were made on the behav-
ior of limpets placed on a soft, sandy-mud substrate.
Surprisingly, the animals had little difficulty, making
almost as rapid progress as they do on glass.

Initially, the anterior portion of the foot is splayed out
in front of the animal, forcing itself into the mud. At
the same time, the proboscis is extended forward as a
narrow tube, sliding on the upper surface of the foot.
When it reaches its maximum extension, the tip expands
and remains expanded as the proboscis is drawn back

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THE VELIGER

Vol. 11; No. 2

on
fo)

Figure 1
(<— adjacent column)

(not to scale)

A. Tornote from the endosome of Tetilla mutabilis. The longest
spicule seen measured about 33mm in length while the smallest
measured 8mm.

B. Oxea from the endosome of the yellow sponge thought to be
Halichondria. The largest measured 3.2mm in length, 0.8mm in
width.

C. Style from the endosome of Hymeniacidon synapium (2 views).
The largest measured 0.75 mm in length, 165, in width.

D. Tornote from the fecal pellets of a specimen of Lucapinella cal-
lomarginata fed only on Tetilla mutabilis. This was the only type of
spicule found.

E. Oxea from the fecal pellets of a specimen of Lucapinella callo-
marginata fed only on Halichondria (?). This was the only type of
spicule found.

F. Style from the fecal pellets of a specimen of Lucapinella callo-
marginata fed only on Hymeniacidon synapium. This was the only
type of spicule found.

G. Representative assemblage of spicules found in the fecal pellets
of a specimen of Lucapinella callomarginata collected on a piling
in Newport Bay, California. Shown are (from left to right) a
prodiaene, a tylostyle (0.15mm length X 4ou width) and an oxea
(100p length X 5 width).

toward the head. The animal then glides forward. Poster-
iorly, the foot is spread out to at least twice the length
of those shown in Plate 16, Figure 4, and the active
secretion of mucus is in evidence as a trail of compacted
mud behind the animal. In this manner, the limpet
actively burrows along, though it was not seen to move
into the mud above the level of the shell. The mantle
flap is maximally lifted in the head tentacle areas and
these openings may be pressed posteriorly or even oblit-
erated by the mud piling up in front of and beside the
animal. The epipodial tentacles are maximally extended.
Small masses of sediment particles taken into the mantle
cavity are ejected periodically through the excurrent
siphon, which is also maximally extended.

Explanation of Plate 16

Figure 1: A specimen of Lucapinella callomarginata from above.
The smallest scale division is 1 mm.

Figure 2: The same animal in side view. Same scale as Figure 1.
Figure 3: The same animal from below. The scale is in millimeters.
Figure 4:
Figure 5: A large specimen of Tetilla mutabilis in the “regular
massive’ form. The dish on which the sponge rests measures six
inches in diameter. The arrow points to the position of a limpet.
Two more limpets are in the sponge but hidden from view. Scale
equals 25mm (Figures 6 to 10 on same scale).

Figure 6: The same sponge in side view. The long root attachment
system is clearly shown. The arrow indicates the position of the

Six specimens of Lucapinella callomarginata.

limpet. The shell aperture and siphon of the animal are visible.
Figure 7: The same view with sponge overgrowth cleared away.
Figure 8: The limpet has been pulled out of the sponge. The
large cavity left by the foot can be seen, as well as a thin layer of
sponge still attached to the foot (arrow). The extended tentacles
of the limpet can also be made out.

Figure g: The same sponge after three weeks. Four possible centers
of regeneration have been created. The two finger-like structures in
the foreground are portions of the yellow sponge thought to be a
species of Halichondria.

Figure 10: A large specimen of Hymeniacidon synapium collected
in four feet of water at low tide.

THE VELIGER, Vol. 11, No. 2 [Mixter] Plate 16

Vol. 11; No. 2

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Some observations were also made of the behavior of
the animals during spawning. Recently collected animals
often spawned within 3 hours after being brought into
the laboratory. Some of the animals which had been in
the laboratory for a few weeks could be induced to spawn
by placing them in a six-inch finger bowl. Evidently the
rise in water temperature would trigger spawning. In one
case, a number of animals was observed to spawn en
masse in a 5-gallon capacity aquarium containing a
healthy Tetilla on which they had been feeding. All of
the animals appeared on the surface of the sponge close
to the osculum prior to spawning. Each animal showed a
definite contraction of the foot and uniform compression
of the shell down against the foot every 3 to 5 minutes.
This was accompanied by a burst of gametes from the
excurrent siphon. Spawning lasted approximately 2 hours
after which the limpets returned to the interior of the
sponge. Some of the fertilized eggs were collected. They
were encased in 2 jelly layers, an inner dense one about
100 uw thick, and an outer, less dense one, about 200 u
thick. The egg itself was a dull yellow color and 132 p in
diameter. Cleavage was not observed.

DISCUSSION

A number of interesting questions is raised by the finding
that Lucapinella callomarginata feeds on the sponge Te-
tilla, as well as on other Demospongia. Since no observa-
tions were made on the detailed behavior of the limpets
as they enter the sponges, the question remains as to the
actual use of the foot during this process. It does provide
a strong grip on the sponge, as attested to by the dif
ficulty of pulling the limpets out of the sponges. In
many cases, in fact, removal of the limpet would result
in tearing away a layer of sponge that remained attached
to the foot (Plate 16, Figure 8). Whether it could provide
some leverage to aid the limpet in penetrating the sponge
remains to be seen. The use of the foot and proboscis
during locomotion in mud is interesting in this regard.

There is also the distinct possibility that the limpets
rarely, if ever, completely destroy a sponge. Since natural
growth rates may far exceed those in the laboratory
the sponges in nature might well be growing faster than
the limpets can eat them. Sponges are notorious for their
sensitivity to laboratory conditions. Although Tetilla
seemed to do reasonably well, there is always the possi-
bility that some subtle deterioration was taking place that
would make the sponges in the laboratory more suscep-
tible to predation by the limpets and thus tend to
amplify whatever effects the limpets might have in
nature.

DE LAUBENFELS (1932) recorded two shapes for Tetilla
mutabilis, “pedunculate-clavate” and “irregularly mas-
sive.” In general, the “clavate” form has a relatively
symmetrical shape with a central osculum and is rather
small. The largest size observed by p—E LAUBENFELS was
22mm. This is probably the juvenile Tetilla. I have no-
ticed that the “massive” form, on the other hand, com-
monly appears in two shapes, one “irregular” and the
other “regular.” The “irregular” has a low, almost flat
outline and bears a number (3 or more, usually) of
oscula, about 3 inches apart. This form is much larger
than the “pedunculate clavate,” and may be over 12 inches
by 9 inches by 3-5 inches when full of water. The “regu-
lar massive” form resembles a much enlarged “clavate”
form although its outline is more nearly spherical (Plate
16, Figures 5, 6). It bears a central osculum, from which
extend, like blunt spokes, 3 or 4 irregular lobes of con-
siderable thickness and it may be up to 6 inches or more
in diameter. Limpets were rarely found in this form,
and the one pictured in Plate 16, Figures 5, 6, 7, and 8
was introduced.

It is possible that the “irregular massive” form is
produced from the “regular massive” by the predations
of the limpets. Plate 16, Figure 9 shows the results of
predation by 3 limpets on the “regular massive” sponge
shown in Figure 5 aftera period of 3 weeks. This collapsed
form could easily give rise to the “irregular massive”
form if the limpets left the sponge and regeneration
occurred; the lobes of the “regular” form that remained
giving rise to secondary centers of organization each with
its own osculum.

Distribution of Tetilla is erratic in Newport Bay, al-
though it is a common species. The few collections of
limpets indicate that their distribution may be even more
erratic. The largest number taken were in an area where
great numbers of Tetilla occurred but too few collections
were made in other areas to see if there was any corre-
spondence. Locality records in the Los Angeles County
Museum of Natural History collections reveal that large
numbers of Lucapinella have been taken on the bridge
pilings in the past, confirming a lack of restriction to
sponges. I have noticed that Tetilla seems to disappear
during the late spring or early summer and Dr. Charles
Brokaw confirmed this (personal communication). Prof.
G. E. MacGinitie stated (personal communication) that
the sponge was seen during the months of October, Jan-
uary, February, and April. pp LauBENFELS (1932) com-
mented that although the sponges were extraordinarily
plentiful in November, he could find none in June of
that same year. Possibly there is a true seasonal bloom of
Tetilla in Newport Bay, but it is doubtful that the distri-

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THE VELIGER

Vol. 11; No. 2

bution of the limpet can be correlated with it, considering
the prevalence of other edible sponges elsewhere in the
bay.

Little is known of the feeding behavior of most Fis-
surellidae except the common assumption that the rocky
intertidal reef-dwellers are surface grazers, foraging on
the layers of diatoms and encrusting algae on hard sur-
faces. Dr. Jefferson Gonor has observed (personal com-
munication), however, that the limpet Diodora aspera
(EscHscHoLtz, 1833) will seek out and consume en-
crusting bryozoans in preference to algae. FRETTER &
Granam (1962) report that D. apertura (Montacu, 1807)
utilizes similar food sources. Sponges are common food
sources of many opisthobranchs and some, such as
Rostanga pulchra MacFartanp, 1905 may have diets
restricted to a single species (Coox, 1962). In some cases,
the mollusks are definitely attracted by their food (Coox,
op. cit.; Lone, 1967; Owen, 1966). No tests were run
to determine the attraction of Lucapinella to Tetilla.
However, the possibility that the limpets do aggregate in
a sponge once it is being attacked cannot be overlooked,
since in most collections more than one limpet was usually
found in a sponge. Observations on the feeding behavior
of Aeolidia indicate that damaged prey becomes more
attractive to other Aeolidia (STEHOUWER, 1952). In
Melongena, the first feeding organism rather than the
prey becomes attractive to other Melongena, which in
turn attack the prey (TuRNeR, 1959). Attraction mech-
anisms like these are probably of advantage for spawning
or mating purposes, and would certainly be of advantage
to Lucapinella.

Turner, Rutu Dixon

ACKNOWLEDGMENTS

I should like to thank Dr. Charles Graham at the Kerck-
hoff Laboratory for his hospitality during my stay at the

jaboratory and Dr. Jefferson J. Gonor for reading the
‘manuscript.

LITERATURE CITED

Cook, EMILy F.

1962. _A study of food choice of two opisthobranchs, Rostanga
pulchra McFartanp and Archidoris montereyensis (Coo-
PER). The Veliger 4 (4): 194-196; 4 text figs.

(1 April 1962)
DELAUBENFELS, Max WALKER

1932. The marine and fresh-water sponges of California.
Proc. U.S. Nat. Mus. 82: 1 - 140

FRETTER, VERA & ALASTAIR GRAHAM
1962. British prosobranch molluscs, their functional anatomy
and ecology. London, Ray Soc. xvi+755 pp.; 317 figs.
Lone, Epwarp
1967. The associates of four species of marine sponges of
Washington and Oregon. Thesis, Oregon State Univ.

McLean, JAMES HAMILTON
1967. | West American species of Lucapinella.
9 (3): 349-352; plt. 49; 3 text figs.
Owen, GaRETH

1966. Feeding. In: Physiology of Mollusca, ed. K. M. WiLBuR
&« C.M. Yonce. II: 1-51

STEHOUWER, E. C.
1952. The preference of the slug Aeolidia papillosa (L.)

for the sea anemone Metridium senile (L.). Arch.
Neéerl. Zool. 10: 161 - 170.

The Veliger
(1 January 1967)

1959. Notes on the feeding of the slug Melongena corona.

The Nautilus 73: 11 - 13

Vol. 11; No. 2

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Page 135

Banding Patterns in Haliotis - II

Some Behavioral Considerations and the Effect of Diet

on Shell Coloration for Haliotis rufescens, Haliotis corrugata,

Haliotis sorensen, and Haltotis assimilis

DAVID A. OLSEN

Zoology Department, University of Hawaii, Honolulu, Hawaii 96822 '

(Plate 17; 1 Table)

INTRODUCTION

IN A PREVIOUS PAPER (OLSEN, 1968) I attempted to
develop a technique for using the sequence of colored
bands in the shells of Haliotis rufescens Swainson, 1822
to trace the seasonal variation in the abundance of Nereo-
cystis luetkeana (MERTENS) PosTELs & RuPRECHT, 1840,
the primary component in the diet of the abalones in
Morro Bay (Cox, 1962). Nereocystis is a large brown
alga that floats well off the bottom. Commercial fishermen
make it available to the abalones by cutting the stipes
while diving and with the propellers of their boats. I
mentioned that this technique could be used to study
algal abundance in the abalone’s home area and would
also be of use to investigators interested in abalone growth
rates. In addition, it would be useful in studying seasonal
variations in feeding intensity as reported by INo (1943)
and later tied in to seasonal temperature fluctuations by
SAKAI (1962a). Abalone growth rate is economically
important since Haliotis is a widespread genus with some
50 or more species (SORENSEN, 1949) and is important
in a number of countries including Japan, Australia,
‘South Africa, Mexico, and the United States of America.
Forster (1967) discusses the importance of growth rate
data in fisheries investigations.

Before the information contained in the banding pat-
terns can be used by sublittoral ecologists certain behav-

1 This work was completed while the author was working on the
Master of Arts program in the Department of Biological Sci-
ences at the University of California at Santa Barbara. The com-
puter analysis was supported by University of California Com-
mittee on Research grant for computer research no. 5997.

ioral information must be considered. The most important
aspect to be considered is whether the animals graze the
macroalgae in their immediate vicinity or trap food par-
ticles which are carried to them by currents and surge.
Any attempt to use the banding patterns to study the algal
abundance in the abalone’s home area would certainly
be unsuccessful if the animals were eating food that was
carried to them by the current from a distant point.

Haliotis feeding behavior has been reported for a num-
ber of species. Cox (1962) says that the red abalone, H.
rufescens, feeds by extending its epipodium and capturing
pieces of macroalgae that float by in the drift. SivcLair
(1963) described the grazing of H. iris kept in aquaria.
She also described populations of abalones living in the
New Zealand tidepools where there were no algae avail-
able for grazing. She hypothesized that the abalones left
these pools at night in order to feed, but it seems possible
that they feed by trapping algal debris washed into these
pools.

STEPHENSON (1924) describes the grazing of captive
Haliotis tuberculata LINNAEUS, 1758 in England, but also
mentions inducing animals in nature to feed by holding
a piece of Fucus near the mouth. From these and other
studies it appears that although abalones will graze, the
normal method of feeding is by capturing particles of
macroalgae in the drift.

Cox (1962) reported annual growth of Haliotis rufes-
cens of up to 48mm but says that it is irregular and
varies greatly. LEIGHTON & BooLootiANn (1963) found the
annual growth rate of H. cracherodu Lracu, 1817 to
be 20mm. In England, Forster (1967) found that H.
tuberculata grew 15 to 16mm per year. SaKar (1962 b)

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found that H. discus hannai Ino, 1953 grew an average
of 20mm per year over a 5-year period. Commercial
fishermen report annual growth of up to 2 inches (51 mm)
but feel that 4 inch (13 mm) is more likely the average
figure.

It is the purpose of this paper to supply the behavioral
information that is necessary before this shell banding in-
formation can be used for dating purposes. In addition,
I have presented the results of a series of feeding experi-
ments which were an extension of the diet and shell color
studies of Ino (1952), Saxar (1962a) and LEIGHTON
(1961, 1963). A preliminary attempt at taking the yearly
patterns of shell colors and utilizing them to group the
sample into age classes is also reported.

MATERIALS ann METHODS

Growth was measured by taking the shell band data
described in the previous paper and totalling the distance
from one red winter band to (but not including) the next.
This was done with a simple program on an IBM 360-50
computer.

To study the effects of changes in diet upon shell color,
5 species of abalones were collected by SCUBA divers,
brought into the laboratory, placed in 8-liter plastic pans,
and maintained at 14 to 20° C in running sea water. I
used small individuals because of space limitations and
because earlier workers reported a rapid early growth
rate. The number of individuals, species, and the size
range were as follows: 8 Haliotis rufescens (64 to 153mm),
10 H. sorensent Bartscu, 1940 (64 to 127mm), 4 H.
assimilis Dati, 1878 (51 to 102mm), 3 H. corrugata
Gray, 1828 (19 to 140mm), and 1 H. cracherodu (33
mm). The animals were fed by placing a liberal quantity
of the algae into the tank and leaving it there until it
was eaten or until it began to deteriorate. The tanks were
cleaned regularly to prevent growth of other species of
algae which would contaminate the experiment. The ani-
mals were fed one species of alga until there was an
obvious area of new shell growth. At this time they were
transferred to another condition. They were fed Macro-
cystis as a brown alga, Gelidium for red and Entero-
morpha for green alga.

Behavioral information came from the abalones used
in the shell color studies, from a series of simple condi-

tioning experiments run predominantly on red abalones,
and from 2 years spent as a commercial abalone fisher-
man. The behavioral observations were conducted in the
following manner: after collection, the abalones were
placed in tanks with running sea water; the tanks were
4x4x1 foot in dimension. The abalones were left alone
for 3 or 4 days to acclimate themselves. The experiment
itself consisted of touching the epipodial, cephalic, and
respiratory tentacles with Macrocystis, probes, and in one
series, juice from ground Macrocystis. Any change in the
behavior of the animal was then recorded. Forty Haliotis
rufescens were used in these experiments.

RESULTS

Annual Growth: There were 361 annual band series
(from one red, or winter band, to but not including the
next red band) observed in the 45 shells. This gives an
average age of the animals used in this study of 8.02
years. There was too much variation in the width of
the year bands to group them into well defined year
classes. The width of these bands represents one year’s
growth and the average was 17.13 mm (S?= 256). The
width of these annual bands ranged from 2mm to
146 mm.

Shell Color and Diet: The results of the feeding - shell
color study are listed below and in Table 1 by species. The
colors given are comparable to colors listed in MAERZ &
Paut (1950) ; the designations in the atlas are given in
parenthesis with each color.

Haliotis sorenseni: Red algae in the diet in every case
resulted in a deep red color (L-7, p. 36) while brown
algae resulted in green-white (G-9, p. 77) or brown-white
(1-3, p. 47). The shell color of the Sorensen abalones that
were fed green algae was the same green-white as when
the animals were fed brown algae. None of the abalones
formed blue-grey shell, a color frequently encountered in
specimens collected from waters deeper than 15 m.

Haliotis rufescens: New shell growth was red (L-7, p.
36) on a red algal diet. When fed brown algae, animals
with shells under 85 mm formed new shell of a pale
blue-green color (E-4, p. 77), while the larger animals
had shells of brown-white (A-1, p. 47).

Haliotis corrugata: New shell growth of Haliotis cor-
rugata was a brown-red (J-12, p. 36) in every case when

Explanation of Plate 17

Figures a to c: Feeding behavior of Haliotis cracherodit.
a: Epipodial tentacles are extended into the water in search of food.
b: Contact is made with a piece of Macrocystis and the animal
extends the foot to capture it. After the food is captured, it
is pulled under the foot (c) before the animal commences eating.

Figure d: Defensive behavior of Haliotis rufescens.
The epipodial tentacles are extended while the shell is clamped
down close to the substrate.
e — epipodial tentacle; f —- foot; m — piece of Macrocystis frond;
r — respiratory tentacle

THE VELIGER, Vol. 11, No. 2 [OtsEN] Plate 17

Figure 1 b

Figure 1 d

Vol. 11; No. 2

THE VELIGER

Page 137

Table 1
Haliotis Shell Color Under Various Algal Diets

Species (sample size) Red algae

Haliotis assimilis (4)

Brown Algae Green Algae Diatoms

red, blue, and white mottled turquoise and white mottled

pale brown orange

Haliotis corrugata (3) — brown-red

turquoise, white

Haltotis cracherodit (1) reddish-brown ' blue-black blue-green 2
blue-green !
Halhiotts rufescens (8) red white pale green 3

blue-green 3
pale green 3

olive green 3

Haliotis sorensent (10) red pale green pale green
purple ? pale brown

Haliotis discus hannai brown 45 bluish green 4 bluish green? bluish green 4
yellowish brown 5 green 5

Haliotis gigantea 5 brown or yellowish-brown green

Haliotis steboldu 5 brown or yellowish-brown

brown or yellowish-brown

' LEIGHTON, 1963

2 LEIGHTON, personal communication
3 LEIGHTON, 1961

4 Saxat, 19624

5 INO, 1952

fed red algae. When they were fed brown algae 2 of
them (one 19 mm and the other 140mm) laid down a
vivid turquoise shell (1-3, p. 83) and the other (85 mm)
laid down a pure white shell.

Haliotis cracherodiu: When fed brown algae this aba-
lone laid down the deep blue-black shell described by
LeIicHTon, 1963.

Haliotis assimilis: Haliotis assimilis was the most vari-

able of the species studied. When fed brown algae indi-
viduals with a predominantly blue shell laid down a
turquoise and white mottled shell; one individual with a
large proportion of red mixed with the blue background
laid down a brown and white mottled shell, and one small
individual (51 mm) with an orange shell continued to
lay down the same color when put on a brown algal diet.
When fed red algae the 2 animals with blue and white
mottled shells laid down shells which were red and white
and blue mottled. No results were obtained for red algae
from the other H. assimilis.
Behavior: The first behavioral action was noticed when
the abalones were brought in from the field. At this time
they would draw the shell down as close as they could to
the substrate and extend their epipodial tentacles as is
shown in Plate 17, Figure 1 d. They would wave these
tentacles rapidly through the water. If the animal was
disturbed further, it would retract the tentacles and clamp
the shell down to the substrate.

Feeding behavior was similar to that described by Cox,
1962. The animal would raise its shell and extend the
cephalic and epipodial tentacles into the water (Plate
17, Figure 1a). When a piece of alga touched one of
these tentacles (Plate 17, Figure 1 b), the abalone turned
and extended the foot in the direction of the contact. At
the same time tentacular waving would increase and con-
sequently there was an increased number of contacts be-
tween the tentacles and the food. Once the foot made
contact with the alga, it pulled the piece under the shell
and the animal began eating (Plate 17, Figure Ic).
Some abalones held food in this position for 12 hours
without eating it. This feeding behavior occurred in all
of the species used in the diet - shell color experiments
and invariably in the 40 Haliotis rufescens studied. Feed-
ing could be stimulated by a variety of stimuli, among
them water current and light. A successful attempt at
conditioning red abalones was made using light as the
stimulus and will be reported in a later paper.

Small abalones (40 mm and under) probably do feed
primarily by grazing since they are found under rocks
and in cracks where drift algac are seldom available. In
abalone cultivation efforts they are frequently fed crustose
calcareous algae which are found on rocks. Abalones can
distinguish the difference between food particles and non-
food particles with their tentacles but not with their foot.
If a glass probe, a finger or any other non-food substance

Page 138

was presented to the tentacles, it would be rejected unless
it had been coated with the juice of some food item. If
the same substance was presented directly to the foot it
would be passed under the foot and the animal would
begin to feed upon it with the radula. The epipodial
tentacles, followed by the cephalic and respiratory ten-
tacles, were the most sensitive to stimulation. All of the
species would graze if no other food was presented.

DISCUSSION

The growth rate of 17.13 mm annual shell increase for
Haliotis rufescens is in remarkably good agreement with
the yearly figures for other species of Haliotis as reported
in the literature. The fact that these shells could not be
grouped into year classes can be attributed to the fact
that they were all greater than 7? inches (196mm) in
length since the California Game Laws forbid the taking
of red abalones under this size by commercial fishermen.
Year classes might have become evident if there had been
a greater size range. The individual variation in growth
rate first noted by Cox, 1962, occurred in 5 abalones used
in a simple conditioning study. They were fed nearly iden-
tical amounts of Macrocystis daily for 160 days and grew
4.7, 6.7, 7.6, 12.2, and 16.8 mm respectively.

Shell color of Halotis sorenseni, H. rufescens, H. cor-
rugata, and to some extent, H. cracherodu varies with
the diet. Red algal diets are recorded in the shells by red
shell color. It would be interesting to raise these species
on a diet of blue-green algae to see if the similarity in red
and blue-green photosynthetic pigments would result in
the same red shell color. Anything but red algal diets
results in a variable color which is usually a turquoise
blue or off-white. Haliotis assimilis presents a more comp-
licated situation and the results of this study indicate
that although diet plays a role in shell color, there may be
some genetic control over the capabilities of the indi-
vidual abalone. The genetics of this species would be
extremely interesting since there appear to be several
easily distinguishable phenotypes.

In using the banding patterns of a haliotid species as
an indicator of algal abundance, care should be taken to
choose a species where the diet effect is readily apparent
and not masked by the genetic makeup of the animal.

Many characteristics of the feeding behavior are sim-
ilar to the escape behavior in Haliotis rufescens and H.
assimilis described by MontTcoMEry (1967). It is curious
that he has not mentioned the defensive behavior de-
scribed in this study. This defensive behavior, with the
shell retracted and the tentacles extended to their maxi-
mum, appears to allow the abalone to test the environ-

THE VELIGER

Vol. 11; No. 2

ment for danger while exposing itself to a minimum of
that danger. The marked similarity between the defen-
sive behavior and the feeding behavior demonstrates how
an animal with a limited behavioral repertoire meets the
demands of a complex environment.

Abalones held in captivity would graze only when no
other food was presented. Normally they fed by finding
pieces of algae with their cephalic and epipodial ten-
tacles and capturing these algae with their foot. They
then pulled a piece under the foot before eating. This
is apparently the normal method of feeding as evidenced
by the following observations from the commercial fishery.
Abalones are normally found clamped on a “scar” on the
rocks, a scar which they have grazed to the bare rock.
The algae around this scar show no signs of grazing. The
second observation is that commercial abalone fishermen
normally find a large proportion of their catch with a
piece of Macrocystis clamped to the bottom of the foot,
even in areas above and below the range of this kelp spe-
cies. These pieces could only have been carried to these
areas by surge and current. Commercial fishermen also
report frequently seeing the abalones raised off the bot-
tom, apparently feeding, during the day, although most
workers feel that abalones are most active at night. These
drift algae are sufficient to support populations of aba-
lones since pockets of them are found in healthy condition
in areas devoid of algae. It would also account for Cox’s
(1962) figure of 540 feet as the lower limit of Haliotis
rufescens distribution, although this level is several hun-
dred feet below the photic zone given by Dawson (1966).
More likely this figure is a misprint of CurTNER’s (1919)
figure of 240 feet.

This feeding behavior probably resulted in the annual
cycle of Nereocystis being recorded in red abalone shells
from Morro Bay, California (OLsEN, 1968). The pre-
servation of the cycle can probably be regarded as an
artifact of the heavy local fishery. The fishermen cause
many stipes to sink to the bottom where the abalones can
get them.

This study has presented some information on the effect
of diet on shell color for 3 previously unreported species.
This information, along with the observations of feeding
behavior, should serve to make the technique of using
shell banding patterns in ecological studies more available
to investigators from a variety of locales.

SUMMARY

The annual cycle of the brown alga Nereocystis luetkeana
has been recorded as a regular pattern of colored bands
in the shells of the red abalone, Haliotis rufescens. A

Vol. 11; No. 2

dating system based on this information could be useful
to sublittoral ecologists if certain assumptions about ab-
alone feeding behavior are satisfied. Abalones feed by
trapping particles of macroalgae in the drift instead of
by grazing. The effects of variation of the algal diet on
the shell color of abalones has been discussed by several
authors and is expanded here to include several heretofore
unreported species. Growth rates of 45 H. rufescens have
been determined by an indirect method and compared to
other rates reported in the literature. Haliotis growth
rates are markedly similar between species.
ACKNOWLEDGMENTS
I would like to thank Barbara Johanson for help in pro-
gramming, Larry Liddle for moral support and help in
caring for the abalones, Chuck Sites for furnishing the
abalone shells from Morro Bay, and my wife Susan for
typing and other assistance.

LITERATURE CITED

Cox, Keitu
1962. California abalones, family Haliotidae.
& Game Comm., Fish Bull. 118: 1 - 133

Curtn_r, W. W.
1919. Observation of the growth and habits of the red and
black abalones. Stanford Univ. Master’s thesis, 21 pp.

Calif. Fish

Dawson, E. YALE
1966. Marine botany.
New York, N. Y.

Forster, G. R.

1967. The growth of Haliotis tuberculata: results of tagging
experiments in Guernsey 1963-65. Journ. Marine Biol.
Assoc. U. K. 47: 287 - 300

Ino, TAKASHI

1943. Ingestion and growth of abalone.

Sci. Fisheries 2: 171 - 174

1952. _ Biological studies on the propagation of Japanese aba-
lone (genus Haliotis). Tokai Regional Fish. Res. Lab. Bull.,

Holt, Rinehart & Winston, Inc.

Bull. Japan. Soc.

THE VELIGER

Page 139

Tokyo, 1953.—English transl., mimeogr. Stanford Univ.,
Calif., pp. 1-114.

LEIGHTON, Davin L.
1961. | Observations on the effect of diet on shell coloration in
the red abalone, Haliotis rufescens SwAInson. The Veliger
4 (1): 29-32; plt. 6 (1 July 1961)
LeicHTon, Davin L. « R. BooLooTIAN
1963. Diet and growth in the black abalone Haliotis crachero-
dit. Ecology 44: 227 - 238

Maerz, Atoys J. « M. Rea PauL
1950. A dictionary of color (2nd. ed.). | McGraw-Hill Book
Co., Inc., New York, pp. 1 - 208; 56 composite color plates.

Montcome_ry, Davin H.

1967. Responses of two haliotid gastropods (Mollusca), Hali-
otis assimilis and Haliotis rufescens, to the forcipulate asteroids
(Echinodermata) , Pycnopodia helianthoides and Pisaster ochra-
ceus. The Veliger 9 (4) : 359 - 368; plts. 50, 51; 2 text figs.

(1 April 1967)
OtseEn, Davin

1968. Banding patterns of Haliotis rufescens as indicators of
botanical and animal succession. Biol. Bull. 134 (1) :
139 - 141

Saxkal, S.

1962a. Ecological studies on the abalone Haliotis discus hannat
Ino — I. Experimental studies on the food habit. Bull.
Japan. Soc. Sci. Fisheries 28: 766 - 779

1962b. Ecological studies on the abalone Haliotis discus hannai

Ino — IV. Studies on the growth.
Fisheries 28: 899 - 908

Srncrair, M.
1963. Studies on the Paua, Haliotis iris MARTYN in the Wel-
lington district, 1945-46. Victoria Univ. of Wellington,
New Zealand, Zool. Publ. 25: 1 - 16

SoRENSEN, ANDREW W.
1949. Some observations on Haliotidae (abalones) and their
world distribution. The Nautilus 62 (4): 138 - 142

Bull. Japan. Soc. Sci.

STEPHENSON, T. A.
1924. Notes on Haliotis tuberculata.
Assoc. U. K. 13: 480 - 495

Journ. Marine Biol.

Page 140

THE VELIGER

Voll 1 Now2

Haliotis pourtalest:i DALL, 1881 from Florida Waters

CHARLES J. GUICE

Fishery Biologist
Bureau of Commercial Fisheries Biological Laboratory, Galveston, Texas 77550 '

(Plate 18)

THE RARE ABALONE, Haliotis pourtalesu DA.u, 1881, pre-
viously has been collected in the Gulf of Mexico on the
steep continental slope bordering the Yucatan and Flor-
ida Straits (Foster, 1946; Harry, 1966). Also, PARKER
(1960) illustrated a fragmented shell of H. pourtalesu
taken from a calcareous bank in the northwestern Gulf.
On April 21, 1967, personnel of the U.S. Fish and
Wildlife Service, Bureau of Commercial Fisheries dredged
a specimen from 45 fathoms on the continental shelf of
Florida at Latitude 24°56.8’ N, Longitude 83°40’ W. The
specimen was taken about 100 nautical miles west-north-
west of the location reported by Foster (1946) and 20
fathoms shallower than any previous record. Although
about 200 bottom grabs were taken along 8 transects from
5 to 1 800 fathoms between St. Petersburg and Key West,
Florida, Haliotis pourtalesi occurred in only one sample.
The specimen is illustrated in Plate 18. Shell dimen-
sions are: Height, 8mm; maximum diameter, 30 mm;
and minimum diameter, 21 mm. There are 28 pore tu-
bercles, 6 of which are open. The color of the shell is

' Contribution No. 268, Bureau of Commercial Fisheries, Bio-
logical Laboratory, Galveston, Texas.

brick red, with irregular splotches of tan scattered be-
tween the line of pores and the columellar border. The
apex has a well-defined protoconch, pore tubercles, spiral
and shoulder threads.

The specimen is deposited in the museum of the
Bureau of Commercial Fisheries Biological Laboratory,
Galveston, Texas. Identification was verified by Dr.
Harold Harry, Texas A&M University.

LITERATURE CITED

FosteEr, RicHarp W.
1946. The family Haliotidae in the Western Atlantic.
Johnsonia 2 (21): 36 - 40
Harry, Haroitp WILLIAM
1966. Haliotis pourtalesii Darr, 1881, from Yucatan. The
Veliger 8 (4): 207 - 208; plt. 30 (1 April 1966)

Parker, Rosert H.

1960. Ecology and distributional patterns of marine macro-
invertebrates, northern Gulf of Mexico. In: Recent sediments,
northwestern Gulf of Mexico, ed. F R.SHEpARD, E B. PHLEGER
& T.H. Van ANDEL, pp. 302 - 337. Amer. Assoc. Petrol.
Geol., Tulsa, Oklahoma

Explanation of Plate 18

Dorsal View of Haliotis pourtalesit Dati, 1881

[GuicE] Plate 18

THE VELIGER, Vol. 11, No. 2

ate

3

Vol. 11; No. 2

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Page 141

Removal of Pea Crabs from Live Oysters by Using Sevin”

JAY D. ANDREWS
DONNA TURGEON

MARIAN HREHA

Virginia Institute of Marine Science, Gloucester Point, Virginia 23062

Minchima nelsoni, A HAPLOSPORIDIAN PARASITE, known
by the anonym MSX, caused a severe epizootic of oysters
in Chesapeake Bay beginning in 1959 (ANpDREwsS, 1967).
Old surviving oysters were dredged for breeding in 1964
but occurrence of pea crabs (Pinnotheres ostreum) was
high. Poor condition, attributed in part to the parasitic
crabs, made spawning and stripping of sexual products
difficult. In the fall of 1964, progeny of these survivors
being monitored for resistance to MSX also acquired
serious infestations of pea crabs. Although early growth
appeared normal, lots held at VIMS pier contained
stunted oysters at 2 years of age in 1966. These stunted
oysters often contained large pea crabs. A safe method of
removing pea crabs from live oysters was needed to elim-
inate another variable in our field tests of oysters for
disease resistance.

In 1966, Miss Hreha, a NSF summer research student,
was assigned the task of finding a method for removal of
pea crabs. After brief attempts with low-salinity waters,
we turned to Sevin, a widely used pesticide for insects
and crustaceans. Our objective was short-term exposure
to kill the crabs rather than continuous exposure at low
concentrations commonly used in tolerance studies. A
method was developed which removes pea crabs without
harming oysters.

The choice of Sevin was based on garden experience
and literature which indicated low toxicity for mammals
and quick detoxification. Also, we found later that Loos-
ANOFF (1961, 1965) had casually reported control of
Pinnotheres ostreum on oyster beds treated with Sevin
and chlorinated benzenes. Probably Sevin was the effec-

® Registered trademark, Union Carbide Corp., New York, N. Y.
* Contribution No. 281 from the Virginia Institute of Marine

Science

tive agent and he suggested use of this insecticide for
pea crabs in 1961.

MATERIALS ann METHODS

For treatment, oysters were crowded in shallow pans (25
in a pan 18 by 24 by 4 inches) holding about 10 1 of
water. Technical Sevin (95%) in powder form was added
to freshly pumped ambient sea water by stirring. Very
low solubility of Sevin made concentrations to which
oysters were exposed quite subjective. Periods of expo-
sure and concentration of pesticide were gradually re-
duced as elimination of crabs continued successfully.
Periods of time for expulsion were measured from the
beginning of treatment. After exposure, oysters were held
in standing or running water until sacrificed or returned
to the York River. Wooden vats were used for treating
trays of experimental oysters.

RESULTS

Summer Studies: Oysters 2 years or more of age were
treated. Only mature pea crabs were present. Treat-
ments were for 24 hours with 100 mg/l of Sevin followed
by continued observation in trays of running sea water.
Concern that oysters might not pump led to use of
freshly-drawn salt water and addition of cultured algae.
Five lots of old oysters from one tray of dredged
survivors of MSX were treated. Some crabs became ap-
parent in 4 hours after Sevin was first added and most
were out or protruding from oysters within 48 hours
(Table 1). All samples had over 50% prevalence of crabs.
Oysters extruding crabs were marked for later observa-
tions of gill and palp damage. Damage by mature crabs
to gill lamellae was always quite evident (see STAUBER,

Page 142

1945). No additional crabs were found when oysters
were opened after 72 hours. All crabs were dead.

Table 1

Expulsion’ of Pea Crabs by Old Mobjack Bay Oysters
exposed 24 hours to 100 mg/I of Sevin
(2 to 12 August 1966)

No. of Number of pea crabs expelled in

oysters 24 hrs. 48 hrs. 72 hrs. Total
25 9 2 3 14
25 9 5 1 15
25 12 1 1 14
27 12 7 1 20
27 10 3 2 15

Total:

1292 52 18 8 78

' Defined as first appearance of pea crab along valve margins.
2 Pea crabs in 60% of oysters

Mature crabs were slowly extruded (often visible sev-
eral hours on shell margins) after compression by valve
action or decay. Most crabs were pushed out the ventral
margin opposite their position posterior to the palps in
the oysters. Legs usually appeared first but sometimes the
orange-red egg sponges were the earliest parts visible.

Smaller oysters (2-year-olds) yielded 26 crabs from 97

oysters, of which 4 came out on the fourth and fifth days.
The crabs ranged from 4 to 15 mm but only 2 were less
than 9 mm. These oysters were not sacrificed because they
were progeny being monitored for resistance to MSX.
The treated oysters, returned to trays in the York River,
exhibited no differences from controls in growth and mor-
tality during the subsequent year. Other groups gave
similar results, with all crabs dead or dying even when
found inside oysters.
Winter Experiments: The winter experiments focused
on small crabs in dormant oysters whereas the summer
studies involved large crabs and active oysters. In the
winter of 1966/1967, attention was given to removal of
immature crabs after a rather heavy late-summer infes-
tation of oysters at VIMS pier. Our objectives were to
determine the effects of temperature, dosage of Sevin,
length of treatment, and method and timing of ejection
of small crabs. Crabs of the 1966 year-class (prehard to
stage III and 1.6 to 5.2mm in carapace width) were
easily distinguished from older mature ones (stage V and
8 to 15 mm). Descriptions of small crab stages are given
by StauBer (1945) and CuristeENSEN & McDermott
(1958).

THE VELIGER

Vol. 11; No. 2

We were concerned about effects of dosage and length
of exposure time on oysters in winter because running
water could not be used effectively. Oysters were inactive
at ambient winter temperatures of about 5°C and warm-
ing to induce pumping was presumed to be necessary.

In a series of experiments, dosage was reduced from
100 mg/I to 50, 25, and 10 without change in effective-
ness and timing of expulsion. Furthermore, treatment
with Sevin was reduced from 48 hours (crabs were ejected
more slowly in winter) to 12 hours with complete elimi-
nation of crabs thereafter. At first pan water was warmed
to room temperature and cultured food added before
treating oysters. However, oysters placed in ambient
river water with Sevin added and allowed to warm for 12
hours at room temperature pumped sufficiently to kill
and expel small crabs within 48 hours (water changed to
remove Sevin at 12 hours). Small crabs (<( 5mm) were
ejected intact (sometimes moribund) beginning about 12
hours after initial treatment and usually removal was
completed in 48 hours. Large crabs (> 8 mm) took sev-
eral days to be extruded, depending upon size of oysters
and rate of decay of pea crabs.

In all experiments, oysters were held 48 hours or
longer and pea crabs were collected frequently. It was
not always possible to determine which oysters ejected
small pea crabs. All oysters were opened and examined
for pea crabs and gill injury. Almost without exception
and regardless of size of crabs, conspicuous and charac-
teristic gill damage was observed. Data on sexes, stages
and sizes of pea crabs were collected.

Table 2
Prevalence of Pea Crabs in VIMS Pier Oysters

No. of No. of % pre-

Date oysters pea crabs valence
Jan. 13-16, 1967 33 9 27.3
Jan. 26 - 31 30 22 73.3
Feb. 1-6 60 35 58.3
Feb. 9- 12 40 21 52.2
Feb. 18 - 20 25 12 48.0
Feb. 21 - 24 50 33 66.0
Feb. 28 - Mar. 2 40 16 40.0
May 31 - June 7 30 7 23.3
June 26 25 5 20.0
Total 278 146 52.5

Prevalences in 1964 year-class oysters attached to
pilings at VIMS pier are given for the winter of 1967
(Table 2). Oysters were treated with various dosages of
Sevin for 12 to 48 hours and were opened from 2 to

Vol. 11; No. 2

THE VELIGER

Page 143

several days after first treatment. The small crabs were
nearly always out in 48 hours but large crabs were found
dead or dying in opened oysters. All crabs were expelled
if oysters were held a week or more, even at ambient
(winter) water temperatures. Only 11 of 146 pea crabs
in Table 2 were stage V crabs (6 to 12mm). The rest
were 1966 year-class pea crabs (prehard to stage III).
The decrease in prevalence between winter and June
samples is probably due in part to disappearance of
males which were last found in the May 31 lot (see
CHRISTENSEN & McDermott, 1958). Possibly tiny crabs
were overlooked in the first sample, thereby accounting
for the low prevalence, or oysters from a higher level on
pilings may have been used.

RECOMMENDED TREATMENT

Our purpose was to find a program of treatment which
would remove pea crabs from oysters without injury to
the hosts. We sought a simple and quick method for ex-
perimental purposes only. Oysters pumped freely in Se-
vin concentrations up to 100 mg/l. Use of Sevin in a fine
hydrophobic powder state raised questions about the con-
centration in solution. Lowe (1967) states that the re-
ported solubility of Sevin in water is less than 99 mg/l.
Hence, all of our experiments were conducted within the
probable solubility range of Sevin although not all flakes
were dissolved, particularly at 100 mg/1 despite limited
stirring.

In October 1967 and April 1968, experiments were
performed using Sevin powder and Sevin dissolved in 50
ml of acetone at 1 and 10 mg/l. Only mature pea crabs
of the 1966 year-class were available in oysters because
infestation by pea crabs failed completely in 1967 in lower
Chesapeake Bay. Acetone appears to hasten the appear-
ance of pea crabs being extruded between the valves of
oysters but both treatments were equally effective in
eventual removal of the parasites. For example, Brown
Shoal oysters with a prevalence of 13 pea crabs in 25
oysters, were treated with 10 mg/l of powdered Sevin on
Tuesday. By Friday 1 pea crab was out and another
visible. Then the oysters were placed in the York River
to stimulate pumping and were opened Monday. Seven-
teen of 26 oysters showed typical gill erosion and only 1
decaying pea crab was found. Mature pea crabs cannot
physically pass between the valves of oysters until they
are mashed or rotted into a collapsed state.

For simplicity we recommend treatment for 24 hours
with 10 mg/l of powdered Sevin in ambient sea water.
Oysters should then be placed in natural waters where
vigorous pumping will occur and pea crabs be expelled.

DISCUSSION

Pea crabs are killed when the oyster hosts are exposed to
10 mg/1 technical Sevin in freshly drawn sea water for
24 hours. After this exposure, oysters may be returned to
ambient conditions with a high probability that pea crabs
will be eliminated. Salinity and temperature levels are
not crucial provided the oysters pump. In winter, standing
sea water should be allowed to warm from ambient tem-
peratures during treatment. Attempts to accelerate
pumping with artificial food were ineffective and un-
necessary.

Sevin is toxic to pea crabs at concentrations of less
than 10 mg/] as it is to other arthropods and fish (Lowe,
1967). Pea crabs in oysters were killed with 1 mg/1 Sevin
in acetone in one experiment. Heavier dosage seems to
provide quicker response in terms of appearance of large
pea crabs being ejected from oysters. For our purpose,
a dosage of 10 mg/I for limited periods seems reasonable
in view of rapid hydrolysis of Sevin. Although 12 hours
has been adequate for killing crabs in oysters, 24 hours
provides some margin for oysters which may be slow to
resume pumping, and longer exposure does no harm.
No attempt has been made to determine critical toxicity
levels for oysters or crabs because chronic exposure is
not desirable or necessary. No consideration was given to
testing oysters for Sevin or its derivatives since the method
is intended only for experimental purposes.

LITERATURE CITED

AnprEws, Jay D. « J. L. Woop
1967. | Oyster mortality studies in Virginia. VI. History and
distribution of Minchinia nelsoni, a pathogen of oysters, in
Virginia. Chesapeake Sci. 8 (1): 1-13

CHRISTENSEN, AaGE MOLLER & JOHN J. McDERMOTT
1958. Life-history and biology of the oyster crab, Pinnotheres
ostreum Say. Biol. Bull. 114 (2): 146-179

LoosanorF, Victor L.
1961. | Recent advances in the control of shellfish predators
and competitors. Proc. Gulf Caribb. Fish. Inst. 13 (1960):
113 - 128

1965. Pesticides in sea water and the possibilities of their
use in mariculture, pp. 135-145. In Research in Pesticides.
Academ. Press, New York, N. Y.

Lowe, Jack L.
1967. Effects of prolonged exposure to Sevin on an estuarine
fish, Leiostomus xanthurus LACEPEDE. Bull. Environ. Con-
tamin. and Toxicol. 2 (3): 147-155

Srauser, L. A.
1945. Pinnotheres ostreum, parasitic on the American oyster,
Ostrea (Gryphaea) virginica. Biol. Bull. 88: 269 - 291

Page 144

THE VELIGER

Vol. 11; No. 2

New Northern Limit for the Limpet, Acmaea digitalis

WILLIAM F. JESSEE

School of Medicine, University of California, San Diego
La Jolla, California 92037

DurING THE RECENT Bering Sea Expedition of the re-
search vessel Alpha Helix, there was some brief oppor-
tunity to examine the intertidal fauna of the southern
coast of the Bering Sea. The intertidal zone in this region
presents an extremely hostile environment in comparison
with either the strictly terrestrial or the strictly marine
zones. During the winter, intertidal invertebrates are ex-
posed twice daily to submergence in water of temper-
atures from +2° to —1.5° C, followed by exposure to
air in which temperatures of — 10° C or lower are quite
common. Survival and propagation under such conditions
is indeed remarkable.

In mid-March, 1968, 6 specimens of the limpet Acmaea
digitalis EscHScHOLTZ, 1833 were collected from the
rocks along the shore of the enclosed bay at Dutch Har-
bor, on the Bering Sea side of Unalaska Island. The
area of collection was at a latitude of about 53°50’ N.
Air temperatures ranged from a daily high in the mid-
thirties (Fahrenheit) to an early morning low of +15°
to +20° FE The animals collected were unremarkable in
any aspect of their morphology, and were definitely iden-
tified as specimens of A. digitalis. The largest specimen
measured 25 mm in length, 18 mm in width, and 15 mm
in height.

As far as can be determined from the literature, this is
the first instance in which Acmaea digitalis has been re-
ported from the Bering Sea. Test (1945, 1946) cites
Acmaea as being a Pacific genus with the northern limit
of distribution of A. digitalis being the Aleutians. Earlier
works (Datt, 1871, 1878) appear somewhat contradic-
tory in their descriptions of the distribution of this species,
but in no instance does reference appear to its occurrence
on the shores of the Bering Sea. Dati (1871) states
that the northern limit appears to be Cape Spencer (near
Juneau), a region of much milder climate than Unalaska
Island. Attempts to locate specimens of Acmaea species
further north, through colleagues in Nome and on the
island of Nunivak, were unsuccessful. Although their re-
ports are most likely correct, personal confirmation of

the absence of Acmaea in these areas is not available at
this time.

That an intertidal species should have such a wide
distribution as does Acmaea digitalis seems truly amazing.
At the southern extreme of its distribution, near San
Diego, California, individuals may be exposed to tem-
peratures as high as 35° to 40° C during summer low
tide periods, while at Dutch Harbor the animals are
subject to winter low tide temperatures considerably be-
low the freezing point of sea water. It is hoped that
studies on temperature tolerances, desiccation, and freez-
ing resistance in this species can soon be undertaken.

ACKNOWLEDGMENTS

The author wishes to express his gratitude to Professor
Per FE Scholander, director of the Physiological Research
Laboratory of Scripps Institution of Oceanography, for
the opportunity to participate in his phase of the Bering
Sea Expedition of the R/V Alpha Helix. This expedition
was made possible by grant GB-7173 from the National
Science Foundation to Professor Scholander.

LITERATURE CITED

Dati, WiLL1AM HEALEY
1871. | On the limpets; with special reference to the species of
the West Coast of America, and to a more natural classifica-
tion of the group. Amer. Journ. Conch. 6: 227 - 282

1878. Report on the limpets and chitons of the Alaskan and
Arctic regions. Proc. U.S. Nat. Mus. 1: 281 - 344

Test, Avery Ransom (Grant)
1945. Ecology of California Acmaea.
395 - 405

1946. Speciation in limpets of the genus Acmaea. Con-
trib. Lab. Vertebrate Biol., Univ. Michigan, Ann Arbor. No.
31: 1-24

Ecology 26 (4):

Vol. 11; No. 2

THE VELIGER

Page 145

Feeding Behavior of Corambella steinbergae

BY

JAMES W. McBETH

Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California 92037

INTRODUCTION

ONE OF THE MOST CoMMON nudibranchs found in the
Friday Harbor region of the San Juan Islands, Washing-
ton, is Corambella steinbergae LANcer, 1962. In the sum-
mer of 1967 I had the opportunity to study the feeding
behavior of this dorid in relation to the bryozoan, Mem-
branipora sp., upon which it is exclusively found and
whose color and pattern it mimics perfectly. A search of
the literature yielded nothing concerning the interaction
between these organisms, but it is generally agreed that
C. steinbergae teeds upon this ectoproct.

Studies on the feeding of the Corambidae in general
are also very sparse. Corambe pacifica MacFaRLAND &
O’DonocuuE, 1929 has been reported to feed on Mem-
branipora villosa Hincxks, but no detail is given (Mac-
FarLAnp & O’DonocHUE, op. cit.). Similarly, a recent
paper on the biology of Doridella obscura VerriL1, 1870
mentions only that this nudibranch is “always found in
association with and feeding on encrusting Bryoza”
Membranipora crustulenta PaLvas in this case (FRANZ,
1967).

The purpose of this study was to find out if Corambella
Steinbergae does, in fact, feed upon Membranipora colo-
nies, and if so, to describe the feeding behavior.

OBSERVATIONS

Initial laboratory observations with a dissecting micro-
scope revealed that Corambella steinbergae definitely
does feed upon Membranipora colonies. The dorid ap-
proaches a fresh zooecium by pressing its mouth against
the smooth surface of the frontal membrane. The lips
are then protruded to form a water-tight attachment to
the membrane. The radula begins to move in short bursts,
from its normal position under the crop, out to the frontal
membrane, where a breach is made. The effective rasping
stroke is the posterior to anterior one, during which the
large lateral spines point forward.

When a slit has been made in the outer, tough, dorsal
covering of the zooid, the soft part is sucked out. Sucking
is achieved by pulsating dilations of the buccal pump,

rhythmically assisted by the lips, which press on the zooid
membrane, compressing the contents of the cell. Action
of the radula is continued and the more solid parts of
the zooid body are carried up on the radular teeth, al-
though the radula was never seen to extend into the
body cavity of the zooecium. As soon as the zooecium is
emptied of living material, the radula is retracted and
the nudibranch pulls its head away. The lips continue to
stick to the frontal membrane for a brief moment until
the suction is released. This lip protrusion can be artifi-
cially induced by forcing the head up while the animal
is feeding. When a zooecium has been cleaned out, the
nudibranch moves to another one and repeats the process.

Studies were also made on the potential feeding behav-
ior of Corambella steinbergae on other genera of encrust-
ing ectoprocts, but in no case was feeding observed.

DISCUSSION

The feeding behavior of the Corambidae offers a rela-
tively untapped field of research. Detailed studies could
be accomplished by growing Membranipora colonies on
glass plates in order for the feeding process to be observed
through the underside of the plate. Other interesting
problems related to feeding behavior could include a com-
parison of the bryozoan growth rate to the nudibranch
grazing rate. Tentative studies show that the colonies of
Membranipora could support many more individuals of
Corambella than they do, despite the presence of other
predators, such as Corambe pacifica. Many colonies of
Membranipora were found to be free of Corambella
steinbergae, while others of the same size had as many as
20 individuals on them. It would, therefore, be of great
interest to make a detailed, local distributional study of
these organisms, as well as a study of movement and
post-metamorphic settling.

Another study that demands attention is concerned
with the relationship between Corambella steinbergae
and Corambe pacifica. These two species have very sim-
ilar physical and behavioral characteristics and are often
found on the same Membranipora colony. It would be
of great ecological interest to compare the biology of

Page 146

THE VELIGER

these two nudibranchs to expose the full extent of their
similarity.

SUMMARY

Direct observations show that the nudibranch Corambella
steinbergae preys on the colonial bryozoan Membrani-
pora sp. Its mode of feeding is a combination of radular
and sucking action. Further possible studies on the bio-
logy of Corambella steinbergae are mentioned.

ACKNOWLEDGMENTS

I am grateful for the opportunity afforded me to do
this study at the Friday Harbor Laboratories, University
of Washington, through the aid of N.S.E Marine Sci-
ences Training grant 11-5055. I would like to express
appreciation to Dr. J. A.C. Nicol and Dr. D.L. Ray
for their help throughout the course of the study; to J.
R. Lance for his valuable initial suggestions; and to Dr.
D.L. Fox for his criticisms of the manuscript.

LITERATURE CITED

Franz, Davip R.
1967. On the taxonomy and biology of the dorid nudibranch
Doridella obscura. The Nautilus 80: 73 - 79

Lance, JAMES ROBERT
1962. A new Stiliger and a new Corambella (Mollusca: Opis-
thobranchia) from the northwestern Pacific. The Veliger
5 (1): 33-38; plt. 6; figs. 1-10 (1 July 1962)
MacFarianpb, FRANK Mace « CuarLes Henry O’DoNOGHUE
1929. A new species of Corambe from the Pacific Coast of
North America. Proc. Calif. Acad. Sci. ser. 4, 18: 1 - 27

Vol..11; No.:2

Vol. 11; No. 2

NOTES & NEWS

A New Record of
Corambella steinbergae LANCE, 1962

BY
TERRENCE GOSLINER

859 Butterfield Road, San Anselmo, California 94960
On May 16, 1968 I collected at San Francisco Yacht
Harbor (Marina) one specimen of Corambella stein-
bergae LANcE, 1962. It measured 8 mm in length and was
found on the Membranipora membranacea (LINNAEUS,
1767) which was encrusted on Laminaria. The range of
this dorid extended from San Diego to British Columbia.
The next area north of Monterey, where C. steinbergae
was known to occur, was Puget Sound, Washington. Thus,
the occurrence of this species in San Francisco Bay estab-
lishes an important intermediary location.

The specimen was identical with the original specimen
described by Lance and was very close to Corambella
bolini MacFartanp, 1966. The specimen has smooth
rhinophores with a clear transparent dorsum marked
with narrow lanes that closely resemble Membranipora.

My thanks to Miss Joan Steinberg who enlightened
me as to the significance of the presence of this animal
in San Francisco Bay and also for her help in the identi-
fication of the specimen.

LITERATURE CITED

Lance, JAMES RoBERT
1961. A distributional list of Southern California opistho-
branchs. The Veliger 4 (2): 64-69 (1 October 1961)

1962. A new Stiliger and a new Corambella (Mollusca: Opis-
thobranchia) from the northeastern Pacific. The Veliger
5 (1): 33-38; plt. 6; figs. 1- 10 (1 July 1962)

W. S. M.

THE SECOND ANNUAL MEETING of the Western Society
of Malacologists will be held at the conference grounds
at Asilomar State Park, Pacific Grove, California June
18 to 21, 1969. Scientific papers, symposia on related
problems, and exhibits will be presented in various fields
related to the study of malacology and invertebrate zoo-

logy.

THE VELIGER

Page 147

Officers for the coming year who were elected at the
1968 conference are as follows: President, Dr. William
K. Emerson, American Museum of Natural History;
First Vice President, Dr. A. Myra Keen, Stanford Uni-
versity; Second Vice President, Mr. Eugene Coan, Stan-
ford University; Secretary, Mrs. Paul O. Hughes, Los
Alamitos; Treasurer, Mrs. Leroy Poorman, Pasadena;
Members-at-Large, Dr. Judith Terry, Palo Alto, Cali-
fornia and Miss Betsy Harrison, Honolulu, Hawaii.

All persons interested in malacology and conchology
are cordially invited to attend and participate in the
coming conference. Excellent accommodations in varying
price ranges (American Plan) will be available for those
making their reservations early.

For information on the conference or on membership
in the Society please address the Secretary, Mrs. Paul O.
Hughes, 12871 Foster Road, Los Alamitos, California
90720.

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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
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We are forced to ask our members and subscribers for
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Effective January 8, 1968 the following charges must be
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We must emphasize that these charges cover only our
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returned copies.

Page 148

Important Notices

Because of the changed rules affecting second class
mail matter, we will no longer be able to include the
customary reminders in our January issue, nor can we
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Manuscripts received up to February 14 each year will
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On May 15, 1968 we published the second part of the
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Part 2 will be available at $3.- plus a handling charge
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their journal bound. To these readers we extend our
apologies.

We are pleased to announce the publication on July
15, 1968 of a Supplement to Volume 11, entitled:

The Biology of Acmaea

edited by D. P. Abbott, D. Epel, I. A. Abbott and R.
Stohler. This supplement is a group of 20 papers dealing
with various aspects of the biology of several different
species of limpets. It comprises 112 pages, 7 halftone

THE VELIGER

Vol. 11; Now2

plates and numerous text figures, charts and tables. This
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THE VELIGER

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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
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BOOKS, PERIODICALS, PAMPHLETS

The Deep-Sea Bivalvia.

The John Murray Expedition 1933-34, Scientific Reports,
Vol. XI, No. 3, Publ. No. 657, pp. 237 - 343, pls. 1 - 3,
figs. 1-38 in text, May 19, 1967. Published by the
Trustees of the British Museum (Natural History).

by J. KNupsEN.

Forty-four species of deep-sea bivalves from the Indo-
west Pacific region are described and illustrated in this
paper. Of these 39 were dredged by the John Murray
Expedition and 5, chiefly in the Zoological Museum col-
lection in Copenhagen, were collected by other expedi-
tions.

The species are distributed into 16 families and 24
genera. Families embracing the largest number of species
are the Pectinidae (9) and the Cuspidariidae (8). Six
species are described as new: Tindaria murrayi, Spinula
filatovae, Amussium sewelli, Pitar sewelli, Xylophaga
murrayt and Lyonsia murrayt.

Most of the specimens were recovered from depths
below 400m. The minimum and maximum depths re-
ported for stations from which the assemblage was ob-
tained were 196 and 3872 m respectively. Remarks on
the systematics, observations on the soft parts, and repro-
ductive development of the various species are included
as well as distribution, bottom content and temperature.

Four, possibly 6, are widely distributed abyssal species.
Four were from the Red Sea, but 3 of these also live
outside that area. Thirty-four are bathyal of which 10
range to East Africa and to Japanese waters. The author
of the paper mentions that the bathyal bivalves suggest
the existence of an Indo-West Pacific fauna having the

Page 150

same range as the adjacent shallow water fauna but
distinct from the bathyal bivalves of New Zealand,
Australia, and South Africa.

Twenty species were found to be carnivorous. Seven of
these, whose stomach content revealed small crustaceans
(copepods, ostracods) and foraminifera, belong in the
Pectinidae; 13 other species belong to the Septibranchia.

An interesting distribution is given for Amygdalum
politum (VERRILL & SMITH) (pp. 269-272, text figs. 14
and 38 [map]), from Japan to the Red Sea, east and west
Africa, western Europe and in the western Atlantic off
eastern United States.

A species of special interest to west American workers
is that cited as C'ryptodon bisecta (Conran, 1849), (pp.
284, 325, pl. 2, figs. 7, 8), from the south Arabian coast
and from the Banda Sea. Conrad’s species was originally
described from strata of middle Miocene age at Astoria,
Oregon. It has been reported as a fossil at other localities
in western North America, western Panama, Japan,
Kamtschatka, Spitzbergen, and Colombia. In the synony-
my of C. bisecta in the present paper are included both
C. investigatoris E. A. SmirH, 1895, originally described
from the Indian Ocean, and Conchocele disjuncta Gass,
1866, originally described from strata of late Pliocene
or carly Pleistocene age at San Pedro, California.

Most west American authors currently consider the
Recent species cited in the literature as Thyasira bisecta
living in west American waters to be referable to Thya-
sira disjuncta (Gass) (Conchoccle disjuncta Gass, 1866),
an assignment discussed by TEGLAND (Nautilus, vol. 41,
pp. 129-130, 1928). The shell of this species differs from
that of T. bisecta in that the anterior end is steeply trun-
cated rather than rounded and projecting. Thyasira dis-
juncta also has been reported living in Japanese waters
and recently Boss (Bull. Mar. Sci., vol. 17, no. 2, pp. 386
to 388, 1967) reported it living in Caribbean waters in
the Gulf of Darien off Colombia at depths of 421-641 m.
In view of the variation and wide distribution reported
by various authors for these large forms of Thyasira,
further study of their identity and range is desirable.

A survey of the deep-sea bivalves of the R.I. M.S.
Investigator and a pertinent list of references are in-
cluded in this interesting contribution to knowledge of
deep-sea Pelecypoda.

LGH

Lavori della Societa Malacologica Italiana

Volumes 1 to 3. Milano, 1964-1966. $12.00, Dr. Fernando
Ghisotti, Via Grotto, 9, Milano.

This collection represents the rebirth of an Italian mala-
cological journal, replacing the long defunct Giornale

THE VELIGER

Vol. 11; No. 2

di Malacologia (Pavia, 1853-1854), Bullettino Malaco-
logico Italiano (Pisa, 1868-1874), and Bullettino della
Societa Malacologica Italiana (Pisa, 1875-1895). Fol-
lowing the establishment of the European Malacological
Union, the Italian contingent, led by Sacchi (Naples)
and Ghisotti (Milan) formed a society in 1963.

The first two volumes consist of reprints of articles
published in other Italian scientific journals, the most
notable paper is by Sacchi on the origin and evolution
of the molluscan fauna of the southern Apennines which
originally appeared in 1963 in the Annual of the Insti-
tute and Museum of the University of Naples.

The first convention of the society took place in Sep-
tember 1966, and volume 3 of the Lavori includes the
original articles, some with descriptions of new species,
which were presented at this meeting. One of the best
papers was given by the French authority Lamotte. His
summary of the factors contributing to the polymorphism
in color and banding in populations of Cepaea nemoralis
generated considerable interest. Phenotypic differences in
color and banding are of selective significance as shown
in the results of a number of experiments, and environ-
mental conditions, such as temperature and proximity to
the sea are related to the frequency of the banding;
unbanded snails are more resistant to environmental
extremes.

A molecular approach to the study of molluscan
phylogeny was postulated by Parisi; he recounted the
value of immunological experiments in ascertaining re-
lationships between species and summarized information
available on the distribution and occurrence of molluscan

_ blood pigments. Girod and Pezzoli presented an exhaust-

ively documented account of the ecology and distribution
of two species of the hydrobiid Bythinella in Lombardy.

The first convention was dedicated to Giorgio Jan who,
with Guiseppe de Cristofori, was responsible for the
founding of the Milan Museum of Natural History. -
Conci presents a short history of that institution along
with a list of Jan’s published works. For American mala-
cologists who have had references to the works of Stop-
pani or Brocchi, it is unfortunate to discover that the
Stoppani collection was destroyed during the second
World War. However, much of Brocchi’s material was
saved and critically redescribed by C. Rossi Ronchetti
(1952-1955. I tipi della “Conchiologia fossile subapen-
nina” di G. Brocchi. Riv. Ital. Paleont. Stratigr., Mem.
V, Milano, 343 pp., 185 figs.).

Of the remaining articles, the more important include
Mariani and Ravera on the trophic ecology of Physa
acuta; Ravera on mollusks as experimental subjects in
radiobiology; Torchio on stranding in cephalopods; and
Carrada et al. on the biogeography of the mollusks of

Vol. 11; No. 2

THE VELIGER

Page 151

Sardinia. Most of the articles have summaries in English,
German or French.

Attention should be called to the “Schede Malacolo-
giche del Mediterraneo,” edited for the Societa by Dr.
Ghisotti (price for the first 11 Schede, 3 175 Lire, $5.00;
see review by S. Peter Dance, 1968, Journ. Conch., 26
(4): 281-282).

K. J. Boss

The Systematic Relationship of Pomatiopsis lapidaria
and Oncomelania hupensis formosana

by G. M. Davis.
pp. 1-143; 32 plts.; 15 text figs.

Malacologia, vol. 6, nos. 1-2,
1967.

The comparative anatomy and the potential for hybridi-
zation of the two members of related hydrobiid genera
were studied, as well as the electrophoretic properties and
laboratory ecology in an attempt to ascertain differences
which might be of value to systematics.

On the basis of their anatomy Pomatiopsis and Onco-
melania are judged to be distinct genera in the subfamily
Pomatiopsinac. This is well documented in the paper.

Disc electrophoretic studies revealed the fact that the
members of the two genera have specific patterns of
protein components, those of the Oncomelanias being
dense and faster moving, while those of the species of
Pomatiopsis are less dense and not fast moving.

Members of Oncomelania did well in laboratory cult-
ure, while the species of Pomatiopsis studied did not
adapt well to laboratory culture. Young of Oncomelania
grew at the rate of 0.65 mm per week with low mortality,
whereas the young of Pomatiopsis grew at a rate of less
than 0.14 mm per week, with a mortality of over 30%
in 2 months,

Documentation of the findings is supported by numer-
ous well executed drawings and graphs. The pertinent
literature seems to be thoroughly reviewed.

RS

Summary of North American Blancan
Nonmarine Mollusks

by Dwicut W. Taytor. Malacologia, vol. 4, no. 1,
pp. 1-172; plts. 1-8; 18 text figs.

The author reviews all known North American nonmarine
mollusks of late Pliocene and early Pleistocene (Blancan)
age. Lists of last appearances of genera and families
are given. About 50 or so Blancan assemblages together
with 10 to 15 older or younger faunas (these included

for convenience of discussion) are summarized under 57
local geographic headings.

On pp. 132 and 133 taxonomic changes are listed;
among these are: 1 new genus (Savaginius) ; 3 new sub-
genera (Calibasis, Oreobasis and Idabasis, all subgenera
of Juga) ; a new species (Planorbella wilsoni) and a new
family (Pliopholygidae in the Viviparacea).

New names are proposed as follows: Juga chrysopylica
for “Goniobasis rodeoensis’ of Hanna, 1923, not of
Crark, 1915; and Radix intermontana for Lymnaea ida-
hoensis YEN, 1946 non HEeNnperson, 1931.

There are 54 taxonomic changes in classification in-
cluding 20 species being synonymized and the remaining
changes involving transfers of species to other genera, re-
storation of a family, and other clarifications of status.
About 20 pages are taken up by the list of references.

RS

Notice des Titres et Travaux Scientifiques (1941-1966)

by Denise Monain.
pp. 1-78; 7 figures.

Apparently privately printed;
Paris, July 1967.

This brochure contains a complete list of papers, both
published as well as unpublished ones, of Dr. Mongin.
With each title is given a brief abstract. There is a list
of the new taxa proposed by the author, 7.¢., 3 new
genera and 23 new spccies; this list is followed by a
detailed list of the collections containing the fossils studicd
by this author. In all, 50 papers are listed, comprising
from 2 or 3 pages to about 500 pages. These papers are
the result of 25 years of active scientific work by Miss
Mongin.

Of special interest to the reviewer is the statement
by the author of her philosophy which guided her in all
her work. She states that in the course of her studies of
fossil mollusks her views were always oriented toward a
biological and ecological interpretation of the collected
faunas and a reconstruction of the bathymetric conditions
of the marine deposits and from these the paleography of
the epoch under study.

RS

Les Mollusques du Bathonien Saumatre
du Moyen-Atlas

by Denise Moncin. Notes et Mémoires du Service
géologique du Maroc, no. 200; pp. 35-95; plts. 1-5, 1967.

New spccies described are:Isognomon marocanus; Eo-
miodon gardcti; Neomiodon skouraensis; and Terebrella

Page 152

THE VELIGER

Vol. 11; No. 2

falloti. The treatment of the species is in the best taxo-
nomic tradition and all species discussed — not only the
new ones — are beautifully illustrated.

RS

Der Schwimmvorgang bei
Gasteropteron rubrum (Rafinesque 1814)

by H.R. HArreLrincer & A. Kress. Revue Suisse
de Zoologie, vol. 74, no. 3: pp. 547-554; 4 text figs.

Gastcropteron rubrum, an opisthobranch, occurring in
the Mediterranean and in the Atlantic, lives primarily on
slimy bottoms in depths of 50 to 80m, but occasionally
is encountered on rocky bottoms. Although this species
has been observed at times in great numbers in the vicin-
ity of Banyuls-sur-Mer, the authors were able to obtain
but a single specimen over a period of 30 months during
which they sought it intensively. The specimen observed
and photographed was obtained while dredging over a
coralline bottom in about 50m depth on May 30, 1964.
The swimming activity was filmed at 24 frames/second.

It was observed that the parapodia are used in a man-
ner comparable to the fluttering of butterflies. They per-
form a few strong motions, lasting about 30 to 40 seconds,
after which time the motions may become weaker or stop
altogether for a restperiod of 20 to 30 seconds. During
this quiescent period the animal slowly sinks to the bot-
tom. The parapodia are extended sideways like wings..
Such swimming could be induced by disturbing the
resting animal; however, the animal also could start the
swimming spontancously.

The swimming of Gasteropteron was compared with
that of other opisthobranchs and it was noted that it
is similar to that of several listed species, but differs from
that of Notarchus punctatus. That species seems to swim
by the principle of repulsion.

RS

Die embryonale Ausgestaltung
der frithen Mitteldarmanlage von Octopus vulgaris Lam,

by Sicurp v. BoLeTzky. Revue Suisse de Zoologie,
vol. 74, no. 3: pp. 555-562; 5 text figs.

In this paper the development from stage VII to stage
XII (stage designation according to Narr, 1921) is pre-
sented. It was noted that the formation of the mid-gut
complex in Octopus is essentially the same as it is in
decapods.

RS

Beobachtungen tber den Feinbau des Schulps
von Sepia officinalis ,

by Icnaz KALtn. Revue Suisse de Zoologie, vol. 74,
no. 3: pp. 596-602; 3 text figs.; 1 plt.

The author demonstrates that the supporting clements
in the swelling (“Wulst”) of the cuttlebone of Sepia
officinalis are not pillars, but rather ramified bands which
are not closed in themselves. They differ in length and in
their meandering form in the dorsal and ventral portions
of the chambers. The crossing of wide and narrow me-
andering bands on the dorsal and ventral sides of the
septa are of utmost importance in the supporting task
in the cuttlebone. Where the bands meander widely, the
connection of the bands is strong, whereas where they
meander narrowly, the connection is weak. These obser-
vations are interpreted as pointing to the possibility that
the cuttlebone is not an absolutely rigid hydrostatic sys-
tem but that it may possess some elasticity.

RS

Statement of Ownership, Management, etc.
of “The Veliger,” published quarterly, on the first day of July, October, January,
and April, at Berkeley, California, as required by the Act of August 24, 1912.
Publisher: California Malacozoological Socicty, Inc. Editor: Rudolf Stohler. Owner:
California Malacozoological Soicety, Inc., a non-profit, educational corporation.
Bondholders, mortgagees, and security holders: none.

(signed)

R. SToH eR, Editor.

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, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.

It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.

Short articles containing descriptions of new species or other 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 geo-
graphical longitudes and latitudes added.

Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS’; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A. M.U.,
U. M. E., W.S. M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION 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 invited. The column “BOOKS, PERIODICALS, and 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, not
exceeding 81,” by 11”, at least double spaced and accompanied by a clear carbon
or photo 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 accommodate the pamphlet (which
measures 51,” by 81”), with double first class postage, should be sent with the
request to the Editor.

EDITORIAL BOARD

Dr. Donatp P. Assort, Professor of Biology

Hopkins Marine Station of Stanford University

Dr. Jerry DonouueE, 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

Dr. E. W. Facer, Professor of Biology

Scripps Institution of Oceanography, La Jolla
University of California at San Diego

Dr. Caner Hann, Professor of Zoology and
Director, Bodega Marine Laboratory
University of California, Berkeley

Dr. G Darras Hanna, Curator

Department of Geology

California Academy of Sciences, San Francisco
Dr. Jorn W. Hepcpetn, Resident Director
Marine Science Laboratory, Oregon State University
Newport, Oregon

Dr. Lro G. HERTLEIN,

Curator of Invertebrate Paleontology
California Academy of Sciences, San Francisco

EDITOR-IN-CHIEF

Dr. RupotF SToH LER, Research Zoologist
University of California, Berkeley

Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology

Stanford University, Stanford, California

Dr. Victor Loosanorr, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr. Journ McGowan, Associate Professor of
Oceanography

Scripps Institution of Oceanography, La Jolla
University of California at San Diego

Dr. Frank A. Piretxa, Professor of Zoology
University of California, Berkeley

Mr. Attyn G. Smiru, Associate Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

Dr. Ratpu I. Smiru, Professor of Zoology
University of California, Berkeley

Dr. Cuartes R. STasEK, Associate Professor
of Zoology

Florida State University, Tallahassee, Florida

Dr. Donatp M. Witson, Professor of Biology
Department of Biological Sciences

Stanford University, Stanford, California

ASSOCIATE EDITOR

Mrs. JEAN M. CaTE
Los Angeles, California

THE

VELIGER

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME II JANUARY I, 1969 NUMBER 3

CoNnTENTS

Maturation of Gonads of Oysters, Crassostrea virginica, of Different Geographical
Areas Subjected to Relatively Low Temperatures (Plates 19 to 2
Victor L. LoosaANorF . .-. , Pan) 6G
A Review of the Living Llentanaeean, Bivalves of the Gens Atenas
(40 Text figures)

EIAROLD Wa Harry) > ..\ eee sete gOS
The Egg Capsules of Jenneria Pastulata ee arroow 1786); with ‘Notes on
Spawning in the Laboratory (1 Text figure; 1 Table)
CuHartes N. D’Asaro . . . : Sicoub yee (a)

The Taxonomic Significance and Theoketical Gaen af ‘Surace pattems ona
Newly Discovered Bivalve Shell Layer, the Mosaicostracum
(Plates 26 to 38; 3 Tables)

Georce H. Hami_ton At SAE Fe eRe ue aa Gey) loe POG
A New Species of Strombina from the @alapaccs Helarids (Plate 39)

WILTIAMUK EMERSON S7ANTHONY, DYATEILIO) (0) 5) ee es TOR
A Note on Feeding and Excretion in Bivalves

P DinaMANI . . . 198

Volutocorbis and F aivotlta: Two Gan of ‘Deepwater Woluudae ibe South Atica
(Plates 40 to 43; 1 Table)

Haratp A. REHDER . . . + - 200
List of Type Specimens of Terebridaes in tHe British iim ‘(Natural Ehistony)
WALTER O. CERNOHORSKY .. . 250

The Ecology of Macoma inconspicua (Eeopenm & SeUreLoY. 1829) i (Central
San Francisco Bay. Part I. The Vertical Distribution of the Macoma

Community (Plate 44; 6 Text oe 2 Tables)
Marityn T. VASSALLO , . . ; mores e223
Spawning of the American Oper Gore virginica, at Bictreme we Levels
(Plate 45)
ANTHONY! CALABRESE & HARRY, ©. DAVISuseiseeies Se a EE O35

[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 12: $18. Domestic; $19.- in the Americas; 19.50 in all other Foreign Countries
Single copies this issue $12.-. Postage extra.

Send subscription orders to Mrs. JEAN M. Cate, 12719 San Vicente Boulevard,

Los Angeles, California 90049. Address all other correspondence to Dr. R. STOHLER, Editor,
Department of Zoology, University of California, Berkeley, California 94720
Second Class Postage paid at Berkeley, California

ConTENTs — Continued

Invertebrates Taken in Six Year Trawl Study in Santa Monica Bay

Joun/ Gy CARMISEE sti ae Sry Fy chiara Uh WN 26%]
Pelecypod — Sediment Association in Tomales Bay, Colina (1 eee 3 Tables)
Don Maurer . . . . 243

Studies on the Mytilus oa Communion in Atartos iBese leaieamel - IV, Sexsonal
Variation in Gametes from Different Regions of the Bay
(5 Text figures; 1 Table)

Donavp R.- Moore & DoNALD J. REISH = 72 =) 2 2 Se oO
Two New Cypraeid Species in the Genus Erronea uaa 46)

CrawFrorpD N.CaTteE. . . : . 256
Growth Study in Olivella eRaete (oman, ness) G Manan I Text foure)

RupoLtF STOHLER . . . 259

Distribution of Organic Bromine Compounds: in A Dyes dalifoniea Coma 1863
(1 Text figure)

Linpsay R. WINKLER . . . . 268
Quantitative Relationships Between Gill Number Respiratory Seb, and Chit
Shape in Chitons (6 Text figures)
Kay M. JoHNson . . 5 Bye

Recognition of an Eastern Bacibe Messina in he Comiline icine fea England and
its Biogeographic Significance

EUGENE V.'GOAN 3s 0 ot D ieee Sake AO 2 EI ee ee oa 07
NOTES & NEWS .. . . eee eid re Saat an EEE Duke Wig 6: PISO
A Color Variation of Aldisa sanguinea (1 Text figure)
RicHarp A. ROLLER
Two New Records of Cratena abronia STEVEN J. Lonc
What is Macoma truncaria Datu? (1 Text figure) EucENE V. Coan
Spawning Notes, II. — Mitra dolorosa (3 Text figures)

Fay H. WoLFson
Littorina littorea in California (San Francisco and Trinidad Bays)
James CarLToNn
Revision of “Introduced Mollusks of Western North America”
JaMEs CARLTON
METHODS & TECHNIQUES .. . MT ee MERT CS fer es. 8 Ge)
A Method of Tagging Mollusks Wnderwater, (1 Text figure)
RicHarp J. RosENTHAL
A Method of Color Preservation in Opisthobranch Mollusks
Gorpon A. RoBILLIARD

BOOKS; PERIODICALS) &PAMPHIEEAS) 202 sete eee tee Cea 2 Oy

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

WoleeliliNon3

THE VELIGER

Page 153

Maturation of Gonads of Oysters, Crassostrea virginica,

of Different Geographical Areas

Subjected to Relatively Low Temperatures '

BY

VICTOR L. LOOSANOFF

Pacific Marine Station, University of the Pacific, Dillon Beach, California 94929?

Bureau of Commercial Fisheries, Biological Laboratory, Milford, Connecticut

and

06460

(Plates 19 to 25)

INTRODUCTION

THE QUESTION OF PHYSIOLOGICAL VARIANTS in,natural
populations of a species has interested biologists for a long
time. The problem was well summarized by PrRossER
(1955) who stated that, regardless of the general interest,
the variations in physiological characters of races, or sub-
species, have rarely been studied systematically. This lack
of research is unfortunate because a comparison of phys-
iological adaptations and requirements of closely related
forms should contribute much to the understanding of
intraspecific relations. Prosser, nevertheless, named sev-
eral criteria used in distinguishing subspecies, and indi-
cated that the response to temperature was one to be
considered. My article deals with several aspects of gon-
adal development of oysters, Crassostrea virginica (GME-
LIN, 1791), of widely separated geographical areas,
subjected to several different, relatively low temperatures.

The existence of physiologically different groups within
the general population of Crassostrea virginica was first
suggested by Coe (1934) during his studies of alternation
of sexuality of these oysters. Long before Coer’s statement
was made, however, practical biologists noticed certain
differences among the oysters of different areas of the
Atlantic Coast. For example, Wetts (1925), who dis-
cussed this matter at some length, stated that an oyster
planter of New England or New York would prefer

' This study was supported in part by National Science Foundation
Grant No. GB-5250

2Present address: 17 Los Cerros Drive, Greenbrae, California
94904, U.S.A.

oysters originating in certain locations: “Today a bushel
of seed from the famous Bridgeport bed will command
double the price of Delaware or Chesapeake seed” (p.
19). Several years later, in studying spawning of Long
Island Sound oysters LoosANOFF & ENGLE (1942) con-
cluded that this population was not homogencous in its
spawning behavior but consisted of different groups, per-
haps races or subspecies, some of which required a
higher temperature than others for completing matura-
tion of their gonads and initiation of spawning. This
opinion was shared by STauBER (1947, 1950) who, using
information available in the literature on spawning of
oysters of different areas, came to the conclusion that
there are probably several physiologically different races
within the general population of C. virginica of our
Atlantic Coast.

The question remained speculative until LOOSANOFF &
Nomeyjko (1951) presented the first evidence on the dif
ference in temperature requirements for gonad matura-
tion in several groups of Crassostrea virginica. These
authors, working with oysters that originated in Massa-
chusetts, Connecticut, New Jersey, and Virginia, demon-
strated that, even though all the mollusks were of the
same species, the temperature requirements for gonad
development and successful spawning of the northern
groups were lower than for groups living in warmer,
southern waters. Preliminary examination of data based
on new and much more extensive studies, in which
samples again represented populations of different areas
of our oyster producing belt from the Gulf of Mexico to
Cape Cod were used, supported the original conclusion
(LoosanorrF, 1958a). Moreover, it is now generally ac-

Page 154

cepted that there are physiological subspecies in Ostrea
edulis Linnazus, 1758 (Korrinca, 1957) and Crass-
ostrea gigas (THUNBERG, 1793) (Imai & SaKat, 1961)
which differ in their responses to environmental factors.
Recently, Harman (1964) working with two groups of
oysters, one indigenous to Long Island Sound and the
other to James River, Virginia, offered additional evi-
dence, based on chromatographic studies of intraspecific
serological differences in C’. virginica. A year later the
same author (HiLttman, 1965), using oysters representa-
tive of Long Island Sound, Delaware Bay, several sections
of Chesapeake Bay, Virginia, and Louisiana, further
demonstrated inherent metabolic differences among the
populations.

MATERIALS anp METHODS

This article is based principally on the results of experi-
ments performed and material collected while I was at
the Bureau of Commercial Fisheries Laboratory, Milford,
Connecticut. Additional material has been collected, how-
ever, and practically all microscopic examination and
study of about 1500 histological preparations of gonads
of oysters, as well as the analysis of the data, have been
completed more recently, since I became associated with
the University of the Pacific Marine Biological Station
at Dillon Beach, California.

Some of the oysters used in these studies came from
Long Island Sound. Others were shipped from New
Jersey, Virginia, North Carolina, and Florida. All oysters

were mature and were 3 to 6 inches long. Except for the ~

lot from Florida (which was sent on November 5), they
were shipped to Milford during the second half of Sep-
tember or early in October after, or near the end of, the
spawning period.

The sources of the different lots of oysters were as
follows: Long Island Sound, dredged in Milford Bay at
a depth of about 30 feet; Miah Maull, New Jersey; the
lower part of York River, Virginia (where the salinity is
about 21 ppt); the vicinity of the State Laboratory near
Bears Bluff, South Carolina; and Apalachicola Bay,
Florida.

Upon arrival the oysters were suspended in wire trays
in Milford Harbor to recover from shipment and to
complete the extremely complex physiological processes
taking place in their bodies at the end of the spawning pe-
riod. These processes include the resorption of gonads and
accumulation of glycogen and other reserve materials
before the oysters begin hibernation (Loosanorr, 1942).
They remained in the Harbor until the middle of Janu-
ary and then were transferred to the laboratory and

THE VELIGER

Vol. 11; No. 3

placed in trays through which sea water of different
temperatures was running at a constant rate.

Approximately 48 hours after the oysters were placed
in Milford Harbor samples of gonads of all groups were
taken for histological studies. All samples came from the
same anatomical portion of the oyster, namely, the right
side behind the line passing through the stomach on a
level with the lower edge of the palps. In this way ana-
tomical uniformity of the samples was assured. The
blocks of gonadal tissue were carefully removed with a
sharp razor blade to avoid pressure that could distort the
tissue and produce various artifacts. The tissue was pre-
served in Bouin’s solution and later processed by stand-
ard histological methods, sectioned at 5, and stained
with iron-hematoxylin and eosin.

REACTIONS or OYSTERS
To DIFFERENT TEMPERATURES

PRELIMINARY CONDITIONING
AND EXPERIMENTAL ARRANGEMENTS

Examination of the Long Island Sound oysters, made
approximately 48 hours after they were placed in Mil-
ford Harbor, showed that they were nearing completion
of, or had completed spawning, and many had virtually
undifferentiated gonads. In this group, resorption, which
follows spawning, had been completed and the oysters
were accumulating glycogen and other reserve materials
(Plate 19, Figure 1). A few individuals, nevertheless, still
contained small quantities of spawn. Gross examination
showed that the meats were healthy and all oysters had
crystalline styles and food in their stomachs.

The New Jersey oysters were not as far advanced as
the Long Island Sound group; they had more individuals
in the late stages of gonad resorption (Plate 19, Figure 2).
The majority of these oysters, nevertheless, had already
completed resorption and were approaching winter con-
dition. All oysters contained large quantities of glycogen,
and crystalline styles and large quantities of food were
found in their stomachs.

The Virginia oysters were, as a rule, in the final stages
of gonad resorption or already displayed winter-like gon-
ads, although a few individuals of both sexes still con-
tained different quantities of undischarged sexual prod-
ucts. Many oysters contained large quantities of glycogen
(Plate 19, Figure 3). This observation suggests that a
lack of this material could not be the reason for the
failure of Virginia oysters to develop gonads later in the
season when they were subjected to the conditioning

THE VELIGER, Vol. 11, No. 3 [LoosanorF] Plate 19

Ly al ay , :
VRS Y
* ¢ «J

pS

Figure 1: Gonad of a typical Long Island Sound oyster late in
September after completion of spawning. The thin gonadal layer
is confined between the body wall of the oyster and the tubules of
the digestive diverticula. The small, undifferentiated gonadal fol-
licles are surrounded by large cells of connective tissue containing
glycogen. Later in the season, when the oysters have accumulated
more reserve material, the glycogen-gonadal layer will become much
thicker. (X 125)

Figure 2: Gonad of New Jersey female oyster late in September
still containing some eggs that will soon be either discharged or
resorbed. (X 125)

Figure 3: Gonad of Virginia oyster late in September. The small,
newly formed gonadal follicles are surrounded by large masses of
connective tissue. (X 125)

Figure 4: Gonad of South Carolina male oyster late in September
still containing a large quantity of undischarged spermatozoa.

(% 125)

Y,
i eh Y
t
i
lh
if =
T ¥ 1

THE VELIGER, Vol. 11, No. 3 [LoosanorFF] Plate 20

mf mo}
lee 3 I 4 Lt
tw ¥, 5 A :
? ae: ’
ee NOE
beh. ENE

Figure 6

Figure 5: Gonad of Florida oyster collected on November 15.
Gonadal follicles are virtually absent and a large portion of the
entire area is occupied by leucocytes. Part of genital duct is
visible. (X 125)

Figure 6: Gonad of Virginia oyster collected on January 15 con-
taining small, undifferentiated follicles and large quantities of
glycogen-laden tissue. (X 125)

Figure 7: Gonad of South Carolina oyster collected on January
15. (X 125)

Figure 8: Gonad of one of the most advanced Long Island Sound
female oysters conditioned for 45 days at 12° C. (X 125)

4 i ‘
i i
FEL i; ,
, é
a )
y
i A,
7
i ‘
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B diy
" 7 7
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Vol. 11; No. 3

THE VELIGER

Page 155

methods employed at Milford Laboratory (Loosanorr,
1945; LoosanorF & Davis, 1950).

The South Carolina group was distinctly different from
the 3 others described. Nearly all oysters still contained
large quantities of spawn, and some appeared entirely
unspawned (Plate 19, Figure 4). All were feeding well;
their stomachs contained large quantities of food and all
had large, well developed crystalline styles.

The first examination of the Florida oysters showed
that they were in a much poorer condition than any of
the previously described groups. They were “watery”,
and contained little glycogen or gonadal tissue (Plate 20,
Figure 5). They had almost no gonadal follicles and the
area between the digestive diverticula and the body wall
was often filled with leucocytes or phagocytic cells. The
presence of large numbers of these cells was characteristic
of this group of oysters. Regardless of their poor condi-
tion, these oysters were feeding, although their crystalline
styles were thin and their stomachs usually contained
much less food than was found in the oysters of the
other groups.

All groups remained suspended in Milford Harbor un-
til January 15, at which time the bottom temperature
was 3° C. Then, the oysters were brought into the labor-
atory, cleaned, and placed in trays of running water the
temperature of which was about 5° C. The next day the
temperature was increased to about 8° C and the oysters
began to receive, as supplementary food, a mixture of
phytoplankton grown in a tank of about 3000 liter capac-
ity (LooSANOFF & ENGLE, 1942 b).

All groups of oysters brought into the laboratory on
January 15 were examined to ascertain their pre-experi-
mental physical condition. The Long Island Sound oysters
were in excellent condition, containing large quantities
of glycogen; their gonads were in typical winter stage.
The New Jersey oysters were in the same condition but
contained even larger quantities of glycogen. The Vir-
ginia oysters were also in typical winter stage with un-
differentiated gonadal follicles. None carried unspawned
cells and, as a rule, all individuals had sufficient glyco-
gen for normal development of gonads (Plate 20, Figure
6).

Although the South Carolina oysters still contained
much spawn in September, by January 15 it was either
entirely discharged or (more probably) resorbed in all
individuals. Therefore, at the beginning of the condition-
ing experiments these oysters were in about the same
physiological state as those of the 3 northern groups,
since these South Carolina oysters also contained suffi-

cient quantities of glycogen for development of gonads
(Plate 20, Figure 7).

The Florida oysters differed radically from all other
groups because, with the exception of a few individuals
whose glycogen reserve was probably adequate (although
considerably less than that of the other groups), the
majority displayed a very thin glycogen-gonadal layer
in which a few winter-like follicles were imbedded. Large
numbers of blood cells were present throughout the bodies
of these oysters, much the same as is shown in Figure 5,
Plate 20. No individuals were found, however, with un-
spawned eggs or spermatozoa.

The general plan of the experiments was to hold some
oysters of each geographic group at temperatures of 12°,
15°, 18°, 21°, and 24°C. = 1°C for different periods to
determine the number of days required for maturation
of their gonads, including the stage when the oysters
become so ripe that they can be induced to spawn
(LoosanorrF, 1945; Loosanorr & Davis, 1963). In all
trays the water temperature was increased gradually to
the desired level to avoid possible physiological shock to
the oysters. Positions of the trays were randomized and
all trays received the same quantity of water and plank-
ton food. Enough oysters were left in reserve for repeti-
tion of the experiments, if necessary, or for studies of the
behavior of the oysters at higher temperatures. The pres-
ent article discusses the observations made at compara-
tively low temperatures, namely, 12°, 15°, and 18°C
=e Ile (@,

In evaluating the condition of the oysters of the dif
ferent groups, especially those of Florida, it was antici-
pated that many might be heavily infested with parasites,
such as Bucephalus, or disease-causing forms, such as
Dermocystidium. Because these organisms may adversely
affect gonadal development, it was decided to base final
conclusions only on those oysters which, upon microscopic
examination, showed no easily recognizable parasites. It
was realized, nevertheless, that even this precaution to
achieve a fair approach in estimating the gonadal devel-
opment was not entirely reliable because the numerous
microorganisms that may affect oysters are not fully
known. Nevertheless, the decision to base conclusions on
only those oysters that appeared healthy helped, undoubt-
edly, to estimate more fairly the condition of the experi-
mental animals.

Two criteria were used to evaluate the rate of gonad
development. The first consisted in determining the num-
ber of days required, at a given temperature, for 50%
of the oysters in the sample to develop physiologically
ripe spermatozoa or fertilizable eggs. Attainment of this
state was ascertained by examination of gonadal material
from each oyster as it was dissected. This material was
suspended in a small quantity of sea water and examined

Page 156

THE VELIGER

Vol. 11; No. 3

under a microscope to determine whether it contained
eggs or sperm. Gametes of the opposite sex were then
added to determine if those of the examined oyster were
physiologically ripe. The sex cells for this test were always
taken from the oysters conditioned at a temperature of
about 24° C and known to be entirely ripe.

The second criterion, applied chiefly to higher tem-
peratures not discussed in this article, was to determine
the length of the conditioning period necessary at each
temperature before 50% of the oysters in a sample could
be induced to spawn. This test was accomplished by plac-
ing each oyster in an individual container and then stimu-
lating it to spawn by a rapid increase of water tempera-
ture to about 28° C or 30°C, and by simultaneous addi-
tion of a suspension of gonadal material taken from ripe
oysters. Normally, each sample, especially those taken for
final examination, consisted of 50 individuals. These
groups will be discussed in geographic order, the northern
oysters being considered first.

OBSERVATIONS AT 12° C

On March 4, after 45 days of conditioning at 12° C, the
Long Island Sound oysters were again measured and
examined. All showed new growth and some, whose total
length was near 100 mm at the beginning of the experi-
ment, had grown as much as 10mm. The meats were
good and the thickness of the gonadal layer varied from
0.5 mm to 1.0 mm. All stomachs contained food and crys-
talline styles were present.

None of the oysters of this group spawned when sub-
jected to thermal and chemical stimulation. A later histo-
logical examination of the gonadal tissue showed that
none were ripe and that many still had typical winter
gonads characterized by small follicles containing virtu-
ally undifferentiated cells. Nevertheless, some advanced
females possessed ovocytes measuring up to 30, although
most of the cells in the follicles were considerably smaller
(Plate 20, Figure 8). After a total of 50 days of condi-
tioning at 12°C a larger number of females had ovo-
cytes measuring between 25 and 30p, and in some males
gametogencsis had progressed to the stage of formation
of spermatids or, possibly, even a few spermatozoa. Even
though the thickness of the gonadal layer of the most
developed oysters was 1.5 mm, none could be induced to
spawn.

The next group of Long Island oysters was examined
after being kept for 61 days at 12°. They were subjected
to spawning stimuli, by slowly raising the temperature of
the water and holding it near 30° C for 2 hours, and by
the addition of spermand egg suspensions. None spawned.

Histological examination of the gonad showed that the
oysters varied greatly in degree of ripeness. Some females
contained many eggs measuring 45, but the majority
had unripe gonads in which the largest ovocytes were
only about 15 in diameter. Spermatozoa were found in
several males (Plate 21, Figure 9). None of the ripe eggs
of the normal females, to which sperm of the 12° C males
was added, were fertilized, however, even though the
sperm remained active for at least 2 hours. It was also
noted that most of the spermatozoa suspended in the
water were still connected by the heads in groups of 2 or
4, probably because they were not entirely ripe. Some of
the eggs taken from the most advanced females were
fertilized by sperm that came from males conditioned at
a temperature of 24°C. Fertilization occurred but none
of the eggs developed into straight-hinge larvae.

The first spawning of the male oysters conditioned at
a temperature of 12°C was induced after 68 days.
Spawning began when the temperature in the spawning
dish reached 28°C. Discharged spermatozoa were ap-
parently normal because they fertilized the eggs of fe-
males conditioned at higher temperatures; the eggs de-
veloped into normal straight-hinged larvae. The male
spawned for 22 minutes, discharging a large quantity of
spermatozoa.

A female was induced to spawn after 78 days. Initially,
I attempted to induce spawning in this group merely by
adding sperm and egg suspensions to the 12° C water in
which the oysters were conditioned. When none of the
oysters responded after 1 hour, the temperature was rap-
idly increased; when it reached 23°C, 15 minutes later,
the first female responded. It was the only female that
spawned in the group of 49 oysters. The eggs released
were fertilized with normal sperm. Many embryos de-
veloped into abnormal larvae, which never progressed
beyond the trochophore, but some reached the normal
straight-hinge stage. Thus, experimental evidence was ob-
tained that certain Long Island Sound oysters can be
conditioned to ripeness at a temperature as low as 12°C.

Histological examination of the gonads of the oysters
used in the spawning experiment after 78 days of condi-
tioning showed that about 80% contained either large
numbers of apparently normal spermatozoa or some fer-
tilizable eggs (Plate 21, Figure 10) in contrast to the
smaller groups that still had essentially winter gonads
(Plate 21, Figure 11). This difference may indicate, as
already suggested by LoosANorr & ENGLE (1942a), that
the population of Long Island Sound is not genetically
homogeneous.

The New Jersey oysters and the more southern groups
were kept under the same experimental conditions as

THE VELIGER, Vol. 11, No. 3 [LoosanoFF] Plate 21

a

, xy)

Figure 12

Figure 9: Gonadal follicle of Long Island Sound male oyster kept
for 61 days at 12°C. The follicle contains ripe or nearly mpe
spermatozoa. (X 575)

Figure 10: Gonad of Long Island Sound female oyster conditioned
for 78 days at 12° C. This individual, one of the most advanced of
the group, contains some ripe eggs. (X 125)

Figure 11: Gonad of Long Island Sound oyster showing almost no
development after conditioning for 78 days at 12°C. The small
follicles, just beginning to differentiate, are surrounded by large
quantities of connective tissue. (X 125)

Figure 12: Gonad of one of the most advanced New Jersey
oysters conditioned at 12° C for 67 days. Note small, undeveloped
follicles and large quantities of connective tissue. Genital duct is
visible. (X 125)

Tue VE.icrEr, Vol. 11, No. 3

ed

is
“ 4

Figure 15 ; | Figure 16

Figure 13: Gonad of New Jersey female oyster conditioned at
12° C for 78 days. (X 125)

Figure 14: Gonad of one of the most advanced Virginia oysters
conditioned for 78 days at 12° C. (X 125)

Figure 15: Gonad of South Carolina oyster conditioned for 78
days at 12°C. (X 125)

Figure 16: Gonad of Florida oyster after 78 days of conditioning
at 12°C. (X 125)

[LoosaNnorF] Plate 22

y
S
o
+ 7
%,
-
oa
2»
\
: a
c =!
ond
“ \
i by
12

Vol. 11; No. 3

those described for Long Island Sound oysters, but the
New Jersey group was not subjected to spawning stimuli
until the 67" day of conditioning because microscopic
examination of the gonadal material before that day
always showed the absence of ripe cells. Regardless of
slow ripening, the New Jersey oysters were healthy, their
meats containing a large quantity of glycogen. In some
the thickness of the gonadal-glycogen layer exceeded
1.5 mm. Practically all showed new shell growth; often
more than 10mm had formed since the beginning of
conditioning. Food and crystalline style were usually
present.

After 67 days of conditioning the oysters were subjec-
ted to a temperature between 25° C and 30° C for over
24 hours and to regular chemical stimulation, but none
spawned or made the rhythmic shell motions character-
istic for females which are ripe or nearly ripe. Examina-
tion of the gonadal material revealed the presence of
neither spermatozoa nor recognizable ovocytes. Histolog-
ical examination showed that even the most advanced
individuals still possessed typical undeveloped winter gon-
ads (Plate 21, Figure 12).

The New Jersey oysters were again subjected to spawn-
ing stimulation after 78 days of conditioning. None re-
sponded after being kept at a temperature of 25°C to
30° C for about 24 hours. A few ovocytes measuring as
much as 25 in diameter were present in about 20% of
the oysters (Plate 22, Figure 13). Strangely enough, not
a single ripe male was found. None of the ovocytes taken
from the most advanced females could be fertilized. Clear-
ly, the New Jersey oysters were not yet ripe. The advent
of spring was making it so difficult to keep the oysters at
12° C that the experiment was terminated at this time.

The Virginia oysters still possessed winter gonads even
after 78 days of conditioning at 12° C. The 50 individuals
that constituted the sample included only 2 with a few
ovocytes, the largest less than 25 in diameter (Plate 22,
Figure 14). The males of this group were also obviously
too immature to be able to spawn. Again, the experiment
had to be ended because of the approach of spring.

After 61 days of conditioning the South Carolina
oysters differed from those of the first 3 groups by being
much poorer. This condition was characterized by the
small quantities of glycogen and by the thinness of the
gonadal layer which in the South Carolina oysters was
almost undetectable; in most it was between 0.0 and
0.5mm. Morcover, these oysters grew much more slowly
than the northern oysters; the best showed an increase in
length of only about 5mm, and at least half of the
oysters showed no new growth. After 6/7 days at this
temperature the conditions remained practically the

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Page 157

same; there was no increase in gonadal thickness and the
oysters remained unripe.

The same situation prevailed on the 78" day of con-
ditioning when the experiment was terminated. The
sexes still could not be easily ascertained and the thickness
of the gonadal layer remained at 0.5 mm or less (Plate
22, Figure 15). The final gross examination showed that
the meats remained watery until the end. These oysters
grew poorly or not at all. Approximately 75% had no
crystalline style or food in their stomachs. Since, as is
shown later, the South Carolina oysters conditioned at
15° C and 18° C contained crystalline styles and had food
in their stomachs, it may be assumed that these were
absent in the oysters kept at 12° C because the tempera-
ture was too low for normal feeding.

The Florida oysters resembled those of South Caro-
lina in appearance and behavior. Throughout the con-
ditioning period of 78 days they remained watery and
poor and displayed a thin gonadal layer of only about
0.5mm or, more commonly, even less. They also grew
much more slowly than the oysters from the 3 northern
areas but, nevertheless, somewhat better than those from
South Carolina. At the end of the experiment no recog-
nizable sex cells could be found in gonadal smears and, as
is shown in Figure 16 (Plate 22), even the best specimens
were in retarded condition. As in the South Carolina
group, more than 3 of the Florida oysters had no crystal-
line styles or food in their stomachs. Obviously, the Flor-
ida oysters could not reach ripeness at 12°C even after
about 24 months of conditioning.

OBSERVATIONS AT 15°C

Previous observations on the behavior of Crassostrea vtr-
ginica at low temperatures have shown that Long Island
Sound oysters can be conditioned to ripen and induced
to spawn at a temperature of 15°C (LoosANOFF «&
Davis, 1952). Nevertheless, these oysters were included
in this experiment primarily to serve as the control for
the more southern groups.

When, after 45 days of conditioning, the Long Island
Sound oysters were exposed to spawning stimuli about
60% of them responded. Histological examination
showed, however, that although many males spawned, a
large portion of the sex cells in their gonadal follicles
was still unripe (Plate 23, Figure 17). Females with well-
developed but not fully ripe gonads were also common
(Plate 23, Figure 18). Abnormalities, principally mani-
fested by the presence in the gonadal area of a large
number of phagocytes, were observed in some oysters.

Page 158

The spawning experiment was repeated with Long
Island Sound oysters conditioned for 50 days. About 70%
spawned, many females discharged several million eggs
each. These eggs, fertilized with the spermatozoa of
males also conditioned at 15° C, developed into normal
larvae. Gross examination of the oysters showed that
they were in good condition, having gonads that aver-
aged slightly more than 1 mm in thickness. The follicles,
nevertheless, still contained large numbers of immature
sex. cells.

Soon after the above-described experiment a natural,
unprovoked mass spawning occurred in all 15°C trays
containing Long Island. Sound oysters. Since we already
knew that these oysters can spawn at 15°C, it was not
entirely unexpected. Because these oysters had a long
history of conditioning at a relatively low temperature,
however, it was decided to use them in auxiliary experi-
ments in which an attempt was made to induce spawning
by chemical stimulation alone at a temperature as low as
12° C. In the first experiment 15 oysters conditioned for
59 days at a temperature of 15° C were placed in water
of 12°C. After being held at this temperature for 2
hours a suspension of ripe sex products was added to the
water. None of the oysters of either sex responded to
this stimulation even after a 2-hour contact with the sex
cells.

In another experiment a second group of 15 oysters,
conditioned at 15° C for 68 days, was placed in water of
12°C and treated as were the oysters used in the first
experiment. After being kept at 12° C for 1 hour and 10
minutes none spawned or showed any response to stimu-
lation. Then, the temperature was slowly increased; when
it attained 13.2°C one male began to spawn copiously.
This, to the best of my knowledge, is the lowest tempera-
ture at which the American oyster, C'rassostrea virginica,
has ever been seen to spawn. Possibly, other oysters of
this same group would have spawned also if a natural
spawning had not occurred in the trays containing this
group only 3 days before this experiment.

The New Jersey oysters conditioned at 15°C were,
throughout the experiment, in excellent physical con-
dition, containing a larger quantity of glycogen than any
other group. The oysters also grew well and, after 45
days of conditioning, their new shell growth averaged
about 5mm. Microscopic examination of smears of the
gonadal material, and histological studies showed that
all except 2 of the oysters of this group possessed virtu-
ally undifferentiated winter gonads. Even the most ad-
vanced males were in the early stages of gametogenesis
and still had only small gonadal follicles (Plate 23, Figure
19). None of the oysters spawned upon stimulation.

THE VELIGER

Vol. 11; No. 3

About 2 weeks later, after 60 days of conditioning,
the oysters were examined again. Most still possessed
undeveloped gonads, but one female contained a few fer-
tilizable eggs. Only a few ripe spermatozoa could be de-
tected in the most advanced male, and nearly all of the
cells still remained in the earlier stages of spermato-
genesis.

After 72 days of conditioning 20% of the New Jersey
oysters could be recognized as females in early stages of
ovogenesis. In only 1 of 25 oysters could some of the
stripped eggs be fertilized and, in general, the oysters in
this group were unripe; most still displayed almost winter-
like gonads (Plate 23, Figure 20). The experiment was
terminated on that date. The remaining oysters were in
excellent condition. Until the very end of the experiment
the oysters fed vigorously, discharging large quantities of
normal true feces.

None of the Virginia oysters conditioned at 15°C
showed much progress at the end of 45 days. After 60
days, however, one male contained a few motile sperma-
tozoa although most of them were connected by their
heads, indicating that they were still not ripe. The rest
displayed virtually winter gonads. After 70 days of con-
ditioning none of the 25 oysters examined possessed
either ripe sperm or fertilizable eggs. Histological prepa-
rations of gonads of these oysters showed that at the
end of the conditioning period, which extended for 72
days, even the most advanced males were still far from
being ripe; the follicles were undeveloped and small
(Plate 24, Figure 21).

It is probably significant that, contrary to conditions
observed in the Virginia oysters kept at 12° C, about 75%
of those subjected to 15°C contained well-developed
crystalline styles and large quantities of food. This dif
ference may indicate, as previously mentioned, that the
group of Virginia oysters, with which we dealt, could not
feed efficiently at 12° C, whereas the temperature of 15°
was high enough for them to feed actively.

At the end of the experiment the meats of the Virginia
oysters were still good and contained large quantities of
glycogen. Approximately 40% of the oysters did not show
any new shell growth, however, a condition which was
contrary to that recorded for the 2 northern groups where
the oysters showed considerable increase in shell length.

The South Carolina oysters differed radically in their
general condition from the oysters of the 3 more northern
areas. Samples of 50 oysters examined on the 60™ and
72° days of conditioning showed that only about 20% of
the individuals were in good condition, possessing a glyco-
gen-gonad layer about 1 mm thick; 26% were medium
poor (gonadal layer of only 0.5mm), and 54% were

Tue VELIcER, Vol. 11, No. 3 [LoosanorrF] Plate 23

Figure 18

> ery e
hats he Fs.
4d
r 4 s

+ i : Ae ey
pe we 4
- wy ie yb ’ ‘ y ‘ LS - ” Ag
4 iy Fie 3 ah Se a 2 ~ a ey!

Figure 19 Figure 20

Figure 17: Gonad of partially spawned Long Island Sound male
oyster conditioned for 45 days at 15° C. (X 125)

Figure 18: Gonad of Long Island Sound female oyster condi-
tioned for 45 days at 15° C. Although the oyster is approaching
ripeness, a majority of the cells in its follicles are still relatively
small. (X 125)

Figure 19: Gonad of New Jersey male oyster conditioned at 15° C
for 45 days. (X 125)

Figure 20: Gonad of New Jersey female oyster in early stages of
ovogenesis, conditioned for 72 days at 15° C. (X 125)

Vol. 11; No. 3

THE VELIGER

Page 159

watery (gonadal layer so thin that it could not be ac-
curately measured). Only about 38% of the oysters con-
tained food in their stomachs and possessed crystalline
styles; the rest had neither styles nor food.

All the South Carolina oysters still displayed undevel-
oped gonads after 60 days of conditioning. Even after 72
days only 1 male of the entire sample contained some
sperm and only 2 females had recognizable small ovo-
cytes. Because the rest of the oysters in the group were
clearly immature (Plate 24, Figure 22), no stimulation
of spawning was attempted.

Of the entire Florida group of 50 oysters conditioned
at 15°C for 72 days only 8 had medium good meats; the
rest were either poor or in the still lower category known
as “watery”. At the end of the experiment only 2 indi-
viduals had a gonadal layer about 1 mm thick; 8 had a
layer of approximately 0.5mm, and the remaining 40
oysters had no definite gonad.

The thinness of the gonadal layer was characteristic
of Florida oysters. Moreover, even this layer was usually
invaded by a large number of leucocytes. In general, the
gonads were so poorly developed that at the end of the
long conditioning period the sex of most of the oysters
could not be easily determined even in the histological
preparations (Plate 24, Figure 23). Strangely enough,
these oysters with poor meat grew well. Only 4 of 50
oysters showed no increase in length, and several showed
an increment of more than 10 mm. Thirty percent con-
tained food and crystalline style, but some individuals
lacked a crystalline style although some food was found
in their stomachs.

OBSERVATIONS AT 18°C

The temperature of the third series of observations is the
one at which Long Island Sound oysters, living under
natural conditions, can fully develop their gonads and
spawn (LoosANoFF & ENGLE, 1940). It was not known
at the time these experiments were begun, however, how
the oysters of other geographic areas would behave at
this temperature. Moreover, it was of interest to compare
the behavior of Long Island Sound oysters at this tem-
perature with that at 12°C and 15°C.

Examination of the Long Island Sound oysters after
only 15 days of conditioning at 18° C showed that their
gonads were sufficiently developed to contain easily recog-
nizable ovocytes or spermatids although, in general, the
oysters were still unripe. After 21 days, however, occa-
sional females already contained some mature, fertiliz-
able eggs and many males had physiologically ripe sper-
matozoa. Three days later the majority of the oysters

contained either ripe sperm or eggs. Regardless of the
presence of some ripe cells, the gonadal follicles of the
Long Island Sound oysters were not fully developed at
that time (Plate 24, Figure 24). Nevertheless, some oys-
ters spawned when subjected to our usual method of
stimulation.

After 32 days at 18°C 75% of the oysters in the
sample were ripe enough to spawn, while approximately
15% still contained immature, winter-like gonads.

The New Jersey oysters exposed to 18°C differed
radically from the Long Island Sound group of which 75%
could be induced to spawn at the end of 32 days. Almost
none of the New Jersey oysters, even the most advanced,
contained easily recognizable sex cells (Plate 25, Figure
25) in 32 days. Strangely enough, this retarded condition
prevailed even though the oysters contained much glyco-
gen, fed well, and appeared healthy.

A few fertilizable eggs or ripe spermatozoa were found
in several New Jersey oysters after 51 days of conditiop-
ing, but none of either sex could be induced to spawn.
After 71 days the majority still displayed virtually undif-
ferentiated gonads but, strangely enough, about 10% of
them could be induced to spawn. Histological studies of
the tissues of the unripe oysters clearly showed that,
regardless of their retarded gonadal development at the
end of the conditioning period, they were healthy and
contained large quantities of glycogen (Plate 25, Figure
26).

All Virginia oysters exposed to 18°C for 30 days pos-
sessed undifferentiated winter-like gonads. Nevertheless,
the generally good appearance of the oysters excludes the
possibility that gonad development was retarded because
of poor physical condition. Oysters examined after 51
and 58 days of conditioning also contained neither active
sperm nor fertilizable eggs, and most displayed winter-
like gonads surrounded by large quantities of glycogen-
laden connective tissue cells. None could be induced to
spawn. Even after 71 days at 18° C, when the experiment
was discontinued, the most advanced females were still
in the early stages of ovogenesis, containing follicles in
which the largest ovocytes were only about 10m or 15p
(Plate 25, Figure 27). No spermatozoa were found and
none of the oysters composing the sample responded to
spawning stimulation.

Histological studies of the tissues preserved at the end
of the experiment showed that large portions of the oys-
ters’ bodies, frequently including the gonadal area, con-
tained unusually large numbers of leucocyte-type cells.
The exact nature of this phenomenon is not understood
but even if it represented a pathological condition, it
did not interfere with feeding; practically all of the oys-

Page 160

THE VELIGER

Vol. 11; No. 3

ters contained food in their stomachs and possessed large,
normal crystalline styles.

Our lack of success in conditioning Virginia oysters
for spawning during the winter was later shared by other
investigators. For example, in a letter to me dated March
30, 1955, Dr. Jay Andrews of Gloucester Point, Virginia,
wrote: “Could you send us a dozen oysters conditioned
for spawning? We have been trying for two months to
prepare Virginia oysters without success.” He was sent
several dozen Long Island Sound oysters which were
easily induced to spawn by our usual method in Dr. And-
rews’ laboratory. During a recent conversation with Dr.
Andrews, in January 1968, I was told that they still can-
not condition their oysters for normal spawning, and
that to culture the larvae of these oysters they still have
to depend on stripping the gonads to obtain some ripe
eggs that are later fertilized artificially.

Since the Virginia oysters kept at 18°C failed to de-
velop ripe gonads at this temperature even after 71 days,
it was anticipated that the South Carolina oysters, a more
southern group, would probably react in the same way.
This expectation was strengthened by the previously re-
ported observation that these oysters when kept at 15° C
did not develop fertilizable eggs or sperm, had poor,
watery meats and, typically, showed a near absence of
glycogen in their tissues. Examination of the South Caro-
lina oysters conditioned at 18°C clearly showed, how-
ever, that these expectations were not justified because
the meats of these oysters were considerably better than
those of the 15°C group. This observation strongly sug-
gests that a difference of 3°C within this temperature
range is highly important ecologically and physiologi-
cally because the South Carolina oysters feed and as-
similate food much more efficiently at 18° C than at
15° C. Possibly 15° C is near the minimal temperature
for the active feeding of this geographic group. Prob-
ably a temperature of about 18° C for South Carolina
oysters can be compared with the temperature level
of approximately 10° C for Long Island Sound oysters,
below which the water-pumping rate is significantly lower
than between 12°C and 15°C (Loosanorr, 1958b). It
should be extremely interesting to verify this possibility
by repeating the experiments on rate of pumping of
Long Island Sound oysters on the Virginia, South Caro-
lina, and Florida populations.

After 30 days at 18° C the South Carolina oysters still
possessed undeveloped follicles. After 51 and 58 days at
this temperature, however, some of them, unlike the Vir-
‘ginia group kept at the same temperature, showed well-
developed gonads. Although none could be induced to
spawn, some males had a few spermatozoa, and some fe-

males had a few ripe eggs that could be fertilized
(Plate 25, Figure 28). Strangely enough, nearly 50% of
the oysters in this sample were still entirely undeveloped.
They were not the poorest individuals; on the contrary,
many of them showed a layer of connective tissue, con-
taining glycogen, that was in excess of 1 mm.

Oysters of both sexes were induced to spawn after 71
days of conditioning. The spawners represented only
about 15% of the sample, but ripe sperm and eggs were
found in other individuals that could not be induced to
spawn. Again, about half the oysters were still unripe,
displaying winter-like gonads.

Samples from the most southern, or Florida, group of
oysters exposed to a temperature of 18° C for a period of
71 days were examined macroscopically and microscop-
ically on the 30™, 51%, and 71 days. All of them were
poor, watery, and contained little glycogen. None showed
even partially developed gonads and neither spermatozoa
nor recognizable ovocytes could be seen in histological
preparations. A common condition in these oysters was
the presence of masses of blood cells in the area normally
occupied by gonadal follicles. In general, these observa-
tions showed conclusively that it was impossible under
the conditions of the experiment to ripen the Florida
oysters at 18°C even after 71 days of conditioning.

DISCUSSION anp SUMMARY

This article is the first comprehensive report on the prog-
ress of gametogenesis of different populations of Crass-
ostrea virginica subjected to low temperatures. It shows
that groups of oysters from Long Island Sound, New Jer-
sey, Virginia, South Carolina, and Florida, kept in Mil-
ford Harbor, Connecticut, for about 3 months and then
subjected to a long conditioning period at temperatures
of 12°C, 15°C, or 18°C, displayed sharp differences in
the stage attained in the development of their gonads.
Some of the oysters that originated in Long Island Sound
were able to ripen even at 12°C. After 68 days of con-
ditioning at this temperature about 65% of these oysters
contained either active spermatozoa or mature eggs.
Spawning of one male was induced after this period and
10 days later a female also spawned. Some Long Island
Sound oysters, therefore, can be brought to ripe condi-
tion by being kept at a temperature of only 12°C.
The New Jersey oysters and those of the more southern
groups differed radically from the Long Island Sound
oysters by being unable to carry on active gametogenesis
at this temperature. As a rule, the gonads in most of these
oysters were so poorly developed at the end of 78 days

Tue VELIcER, Vol. 11, No. 3 [LoosanorF] Plate 24

Figure 23 Figure 24

Figure 21: Gonad of Virginia oyster conditioned for 72 days at
15° C. (X 125)

Figure 22: Gonad of South Carolina oyster conditioned for 72
days at 15° C. (X 125)

Figure 23: Gonad of Florida oyster conditioned for 72 days at
iio (Oh, (X 125)

Figure 24: Gonad of Long Island Sound female oyster condi-
tioned for 24 days at 18° C. (X 125)

Vol. 11; No. 3

that the sexes could not be easily distinguished even by
microscopic examination of gonadal smears.

The differences in the ability of the oysters of the
different groups to ripen at the same temperatures were
also clearly demonstrated at a temperature of 15°C.
Again, the Long Island Sound oysters ripened compara-
tively well under this condition, giving about 60% spawn-
ers after a conditioning period of 45 days. Even after 72
days of conditioning, however, only 20% of the New Jer-
sey oysters could be recognized as males or females and
the majority possessed winter-like gonads. Only a single
oyster had some eggs sufficiently developed to be fertiliz-
able. The same condition existed with minor modifica-
tions in the Virginia and South Carolina groups, and the
Florida oysters were so poorly developed that no definite
gonad was detectable in most.

When exposed to a temperature of 18°C the Long
Island Sound oysters again stood apart from the other
groups. After 21 days of conditioning the majority of
these oysters already contained ripe gametes. When these
oysters were stimulated by exposure to higher tempera-
ture, after only 24 days of conditioning, many spawned.
On the other hand, none of the New Jersey group could
be induced to spawn after 51 days of conditioning, and
even after 71 days of conditioning at 18°C, most of the
oysters in this group still contained virtually undifferenti-
ated gonads, even though some could be induced to
spawn. The Virginia oysters could not develop ripe gam-
etes after being subjected to 18°C for 71 days. Strangely
enough, after the same period of conditioning about 15%
of the South Carolina oysters were ripe and could be
induced to spawn, although about half of the individuals
possessed such undifferentiated gonads that the sex could
not be determined.

The ability of a considerable percentage of South
Carolina oysters to develop gonads at 18° C, although
none of the Virginia group could, interrupted the other-
wise orderly progression, from north to south, of the tem-
perature requirements for gonad development of different
geographical populations of Crassostrea virginica. An
explanation for the unexpected behavior of this preco-
cious fraction of the South Carolina population is not
available at present. Moreover, these precocious few seem
incongruous, considering that about 50% of the group
failed to respond to conditioning at 18° C for more than a
2-month period. The possibility is suggested that the
South Carolina oysters, even though, according to the
sender, collected from the same area, were apparently of
different genctic composition and thus resemble the ap-
parent mixture of several distinct genotypes in the oyster
population of Long Island Sound (Loosanorr, 1965).

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Page 161

Nevertheless, causes other than genetic may be responsible
for such differences in the gametogenic activities of the
South Carolina oysters.

Obviously, such atypical fractions of the general popu-
lations, as in the samples of South Carolina oysters, make
it increasingly clear that, in addition to histological and
chromatographic studies of the oysters, to which refer-
ences have already been made, it may be necessary to
employ genetic studies to clarify that rather difficult
problem. While I was still the director of Milford Labor-
atory a program for genetic studies of bivalves was estab-
lished, and at present experiments of this nature are being
conducted by Dr. Arlene C. Longwell. According to Dr.
Longwell, with whom I corresponded, “There is sufficient
justification for the interesting speculation that genetic
differences play a role in the differences in gametogenic
activities of the South Carolina oysters exposed to the
same temperatures which failed to elicit a response from
the other southern groups studied.”

The design of these experiments may be criticized on
the grounds that the differences which I found among the
groups of oysters of different regions may be due to the
fact that the preconditioning period, extending roughly
from the second half of September until the middle of
January, was too short to allow the oysters to become ac-
climated to the new environment. The matter of acclima-
tion in mollusks, which was so ably discussed by SEGAL
(1961), is still poorly understood, however. It can prob-
ably be considerably clarified by conducting a series of
experiments with successive generations of laboratory-
reared oysters originating from parents of different geo-
graphical areas to determine whether successive genera-
tions of these oysters still fail to spawn in Long Island
Sound. We had planned a series of these experiments in
the past but, unfortunately, a heavy mortality of south-
ern oysters in Milford Harbor during winters (especially
Florida seed oysters of which almost 100% died) pre-
vented completion of the research. Nevertheless, our
preliminary studies definitely established that oysters
that originally came as seed from different geographical
regions and spent approximately 2 years in Milford Har-
bor did not become acclimated even after that long period
(LoosaNorF & Nomejxko, 1951).

Additional observations of a similar nature were made
on 150 New Jersey oysters shipped to me as seed by Dr.
Thurlow Nelson in the fall of 1948. These oysters, which
came from the Delaware River near Cape May, were
attached to Mactra shells and were easily identifiable.
They grew well and during the first winter showed no
mortality whatsoever — indeed were the best of the 6
groups with which we were working at that time, excel-

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THE VELIGER

Vol. 11; No. 3

ling even the Long Island Sound oysters which showed a
mortality of 9%. In the fall of 1949 the average length
of these oysters was 77.7 mm; in the fall of 1950 it was
92.8 mm; and in the late summer of 1951 it exceeded
12 cm. When these large, healthy oysters were examined
in the fall of 1951 over 30% still contained nearly un-
discharged gonads, and many others were in only parti-
ally spawned condition. Thus, even after 3 years of exist-
ence in Long Island Sound the New Jersey oysters, or at
least a large portion of them, were unable to spawn.

Still another evidence of lack of acclimation in oysters,
which has already been briefly reported (LOOSANOFF &
Nomeyjko, 1951), was found in a large shipment of
native oysters from Seaside of Virginia, near Chinco-
teague. These oysters were planted by F: Mansfield & Sons
Company in comparatively shallow water in Long Island
Sound south of West Haven. Our observations showed
that these Virginia oysters, even after 3 years existence in
the new environment, failed to discharge their gonads.
These oysters, which, by general appearance and other
peculiarities, were easily distinguished from those of Long
Island Sound, went into hibernation each fall still re-
taining all or at least large quantities of undischarged
spawn, thus clearly demonstrating that they had not ac-
climated to the new set of conditions.

One more group of oysters transplanted to Long Is-
land Sound that were unable to discharge their spawn in
the new environment was that of several carloads of
Crassostrea virginica shipped by EM. Flower «& Sons
Oyster Company from the lower part of the Hudson
River, the area between Hastings-on-Hudson and Tarry-
town, and planted near the mouth of the Housatonic
River, which enters Long Island Sound near Milford. We
examined these oysters for more than 3 years and found
that they also went into hibernation with undischarged
gonads. This last transplantation is of special interest
because it clearly demonstrates that distinctly different
oyster populations, as far as their spawning requirements
are concerned, may exist comparatively close to each
other since the distance between the Housatonic River
and the area of the Hudson River from which the oysters
came is only about 70 miles.

Perhaps the most striking evidence of the inability of
some pelecypods to acclimate to the new conditions was
offered by Davis (1955). He has shown conclusively in
his experiments with Ostrea lurida CarPENTER, 1864,
that the 6" generation of these oysters transplanted from
Puget Sound to Long Island Sound could not survive New
England winters, despite some selection, through the death
of the most susceptible each year, before the remainder
were brought into the laboratory during the coldest
period.

We may close this discussion by concluding that there
are distinct populations of Crassostrea virginica that re-
quire different temperature regimes for completion of
gametogenesis and spawning. ‘This conclusion is based on
the experiments described in this article, which were
devoted to studies of the comparative progress of gam-
etogenesis of oysters of different geographical areas kept
in Milford Harbor for approximately 3 months and then
subjected to conditioning at relatively low temperatures
ranging from 12°C to 18° C. These observations support
our earlier conclusion (LoosANoFF & NomEjxko, 1951)
that Netson’s (1928) assumption that the breeding
temperature is the same for C’rassostrea virginica over all
parts of its range is incorrect. Moreover, the present ex-
periments clearly indicate the existence of several phyiso-
logical variants within the general populations of C.
virginica.

ACKNOWLEDGMENTS

I thank Dr. Harold Haskins of Rutgers University, Dr.
Jay Andrews of Virginia Fisheries Laboratory, Dr. G. R.
Lunz of Bears Bluff Laboratories, and Dr. Robert Ingle
of ‘Tallahassee, Florida, for sending me the samples of
oysters from their respective areas; Mr. Harry Davis of
the Bureau of Commercial Fisheries Biological Labora-
tory at Milford, Connecticut, for assisting in some of the
studies; Miss Rita Riccia of the same laboratory for her
assistance in the preparation of this manuscript, and Mr.
Andrew Driscoll of the Pacific Marine Station, for pre-
paring the photo-micrographs. I also thank the National
Science Foundation for giving me the opportunity to
complete these studies.

LITERATURE CITED

Cor, WESLEY ROSWELL
1934. Alternation of sexuality in oysters.
68 (716) : 236 - 251

Davis, Harry Cari

Amer. Natur.

1955. Mortality of Olympia oysters at low temperatures.
Biol. Bull. 109 (3) : 404 - 406

Hituman, Rosert E.

1964. | Chromatographic evidence of intraspecific genetic dif-
ferences in the Eastern oyster, Crassostrea virginica. Syst.
Zool. 13 (1): 12-18

1965. | Chromatographic studies of allopatric populations of
the Eastern oyster Crassostrea virginica.

6 (2): 115-116

Chesapeake Sci.

Tue VELIGER, Vol. 11, No. 3 [LoosanorF] Plate 25

’
j a) bbe Ye! (74 7%
ig BI SUPT Pome me
Pid vas BLY e ae Re ”

Figure 26

Figure 25:Gonad of New Jersey oyster conditioned for 32 days at
18° C. (X 125)

Figure 26: Gonad of unripe New Jersey oyster conditioned for
71 days at 18° C. (X 125)

Figure 27: Gonad of Virginia oyster conditioned for 71 days at
18° C. (X 125)

Figure 28: Gonad of South Carolina oyster conditioned for 58
days at 18° C. Some ripe eggs were found in this individual.

(X 125)

Vol. 11; No. 3

THE VELIGER

Page 163

Imat, Takeo « Seucut SAKAI

1961. Study of breeding of Japanese oyster, Crassostrea gigas.
Tohoku Journ. Agr. Res. 12 (2): 125-171
Korrinca, PETER
1957. _\Water temperature and breeding throughout the geo-
graphical range of Ostrea edulis. Ann. Biol. 33 (1-2):
1-17
LoosanorrF, Victor Lyon

1942. Seasonal gonadal changes in the adult oysters, Ostrea
virginica, of Long Island Sound. Biol. Bull. 82 (2): 195

to 206

1945. Precocious gonad development in oysters induced in
midwinter by high temperature. Science 102 (2640) :
124 - 125

1958a. Challenging problems in shellfish biology In: Per-

spectives in marine biology, Part IV (1956), A. A. Buzzatti-
‘Traverso (Ed.), Univ. Calif. Press, Berkeley & Los Angeles, pp.
483 - 495

1958b.
peratures.

Some aspects of behavior of oysters at different tem-

Biol. Bull. 114 (1): 57-70

1965. | Gonad development and discharge of spawn in oysters
of Long Island Sound. Biol. Bull. 129 (3) : 546 - 561

Loosanorr, Vicror Lyon « Harry Cari Davis
1950. Conditioning V. mercenaria for spawning in winter and
Biol. Bull. 98 (1) :

breeding its larvae in the laboratory.

~ 60-65

1952. ‘Temperature requirements for maturation of gonads of
Biol. Bull. 103 (1): 80-96

1963. Rearing of bivalve mollusks Jn: Advances in Marine
Biology, EF S. Russet (ed.), Acad. Press, London 1: 1 - 136

northern oysters.

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 in-
tensity and time of oyster setting. Bul. U.S. Bur. Fish.
49: 217 - 255

1942a. Accumulation and discharge of spawn by oysters living
at different depths. Biol. Bull. 82 (3): 413 - 422

1942b. Use of complete fertilizers in cultivation of microor-
ganisms. Science 95 (2471): 487 - 488

Loosanorr, Victor Lyon « GHartes A. NomMEJKo

1951. Existence of physiologically-different races of oysters,
Crassostrea virginica. Biol. Bull. 101 (2): 151 - 156
Netson, THURLOW CHRISTIAN
1928. __ Relation of spawning of the oyster to temperature.
Ecology 9 (2): 145 - 154
Prosser, Citrrorp Lapp
1955. Physiological variation in animals. Biol. Rev.

30 (3): 229 - 262
Seca, Earu

1961. Acclimation in molluscs.
235 - 244; 5 figs.; 4 tables

Srauser, LEsuie A.

Amer. Zoologist 1 (2) :

1947. On possible physiological species in the oyster, Ostrea
virginica. Anat. Rec. 99 (4): 614

1950. The problem of physiological species with special ref-
erence to oysters and oyster drills. | Ecology 31(1): 109-118

WEL Ls, WILLIAM FE

1925. Oyster investigations. Conserv. Comm., State of
New York, Fourteenth annual report for the year 1924: 3 - 22

Page 164

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Vol. 11; No. 3

A Review of the Living Leptonacean Bivalves

of the Genus Aligena

HAROLD W. HARRY

Texas A&M Marine Laboratory, Galveston, Texas 77550, and Rice University, Houston, Texas 77001

(40 Text figures)

IDENTIFYING A SPECIES OF Aligena from Galveston, which
proved to be undescribed, led to the present review. I
have reappraised all Recent species named in the genus
or referred to it. Possibly other species will soon be found
to belong here, for many of the descriptions and illustra-
tions of leptonid bivalves in the literature are tantalizingly
vague with regards to critical characters of the shell, and
their generic allocations are questionable.

I am grateful to Mr. Cornelius Mock and Mr. Charles
Guice of the U.S. Bureau of Commercial Fisheries, Gal-
veston, for aid in field work and much useful material.
Dr. J.P. E. Morrison, Dr. Harald Rehder and Dr. Joseph
Rosewater, as well as the whole staff of the Department
of Mollusks of the U.S. National Museum were very
generous in aiding my studies there. This study was sup-
ported in part by National Science Foundation Grant

GB 2753.

Aligena H.C. Lea, 1846

The genus Aligena H. C. Lea, 1843, when proposed, con-
tained but two species of fossil bivalves from the Miocene
beds of Petersburg, Virginia, A. striata and A. laevis, both
nude names with neither designated as type. Lea de-
scribed these species in 1846. In his synopsis of the Lep-
tonacea of North America, Dati (1899) designated
“Abra aequata Conrab’ as type of the genus. The fol-
lowing year he published a monographic account of
Aligena (Dati, 1900: pp. 1175 - 1177), in which he noted
that Conran’s species was originally named Amphidesma
aequata Conrap, 1843, and that this is a senior synonym
of Aligena striata H.C. Lea, 1846. Kelliopsis VERRILL &
Busn, 1898, type by monotypy Montacuta elevata StimP-
son, 1851, was considered a subjective synonym of

Aligena by Dat. (1899, p. 895; 1890, p. 1175). In this

he is probably justified, for the type species of Kelliopsis
differs from the type of Aligena chiefly in being smaller,
more quadrate, and in having weaker sculpturing. I have
not had the opportunity to examine specimens of 2 other
nominal genera based on Tertiary fossils of Europe,
which Dati (1900) cites as congeneric (Laubriereia
CossMann, 1887) or closely related (Spaniodon REUuss,
1867).

Several additional living species have been added to
Aligena in the twentieth century. Aligena cerritensis AR-
NoLD, 1903 was described from a Pleistocene deposit in
southern California and has since been found living, chief
ly south of there. Aligena borniana Datt, 1908 from
deep water in the mid-Pacific is dubiously a member of
the genus, and A. pisum Datt, 1908, from the Strait of
Magellan, is generically misplaced. Aligena cokert DALL,
1909 from Peru is a member of the genus, as is A. nucea
Dat, 1913 from the Gulf of California.

TueLe (1935) noted that Aligena differed so slightly
from Montacuta that he was only doubtfully willing to
recognize it as a subgenus therein. He pointed out that
most living Aligena were on the shores of the American
continents, but he also suggested, correctly, that Mont-
acuta salamensis JAECKEL & THIELE, 1931 (in THIELE &
JAeckeEL, 1931) from the east coast of Africa might be
an Aligena,

Two more species have recently been added to the list
from the eastern Pacific, Aligena redondoensis T: BurcH,
1941, and A. borealis Cowan, 1964. For reasons noted
below, these probably belong elsewhere.

The nondescript bivalves which are usually cited in the
superfamily Erycinacea or Leptonacea are so poorly
known that the allocation of genera to families is a
complex question, which cannot yet be fully decided. For
the present we may place Aligena in the Montacutidae,
and define the genus as follows.

Vol. 11; No. 3

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Page 165

DEFINITION or THE GENUS Aligena

Shell equivalve, small, 1 cm or less in maximum dimen-
sion; color always uniformly white; shape various, but
usually the height and length are about the same; umbos
touching, slightly prosogyrous, located midway the length
or behind it, or rarely before it. There is no lunule,
escutcheon or corcelet. The dorsal margins do not project
across the midline. Outer surface sculptured only with
growth lines, which may be very faint, or moderately
pronounced, regular in size and spacing or irregular in
those characters. Adductor scars are about equal in size,
subquadrate to suboval; the pedal retractor scars join
the adductor scars above (or are separated from them
slightly, perhaps depending on stage of growth). Pallial
line simple, without sinus, joining the ventral end of the
adductor muscle scars, usually with ragged margins, at
least on the upper side, and often discontinuous in one or
several places. Valve margin thin and smooth. Umbonal
cavity deep, the hinge plate very narrow, usually with a
finger-like tooth below the umbo in each valve, of which
the right onc is slightly larger than the left. Lateral teeth
are always absent. Ligament consisting of a thin, exter-
nal part (tensilium) on the very margin of the shell
following its curvature and extending before and behind
the umbos, and a larger, straight, elongate internal part
(resilium) which is entirely separated from the tensilium.
The resilium extends in a straight line from immediately
below the umbos for a moderate distance backward, and
is therefore entirely opistodetic. Its attachments, or re-
silifers, are usually buttressed by a moderate thickening of
the shell, or chondrophore, often terminated behind in a
sharp angle. The resilifers diverge outward and down-
ward from the umbos, and thus the resilium is narrow in
front, broader behind, with the form of an elongate tri-
angle in ventral view. The ventral surface of the resilium
may be calcified, forming a lithodesma. The protoconch is
oval, smooth, large (about 0.3 to 0.4mm long) and
remarkably persistent, being plainly evident in most
large shells which are not worn. The periostracum is
thin or moderately thick, light tan, usually with a smooth
outer surface. It is only moderately persistent in most
species.

In some species there is a vague, shallow sulcus on the
disc of the valve, running from the umbo to the mid-
ventral margin, and a corresponding sinus in the latter.
Such species are slightly reniform in profile. This sulcus
and marginal sinus are found in several other gencra of
the Leptonacea.

The species of Aligena as presently restricted are ma-
rine, mostly living from the lower tide level to shallow
depths of perhaps 40 fathoms along the temperate and
tropical shores chiefly of the American continents. One
species of the Indo-Pacific fauna extends to greater
depths. None are known from the eastern Atlantic or
Antarctic faunas.

Although a few species of Leptonacea may be free
living, most seem to be commensal with burrowing in-
vertebrates, chiefly crustacea, polychaete worms and echi-
noderms. Nothing definite is known about the habits of
most species of Aligena, but A. cokeri and possibly A.
texasiana are probably polychaete associates.

TYPE SPECIES oF THE GENUS Aligena

Aligena aequata (Conran, 1843)
(Figures 1, 2, and 3)

Amphidesma aequata Conran, 1843. Proc. Acad. Sci. Phila-
delphia 1:307. Not figured. Type locality: St. Mary’s Co.,
Maryland, and Wilmington, North Carolina. (Miocene)

Aligena striata H: C. Lea, 1846, Trans. Amer. Philosoph. Soc.,
28° Ser. 9: 238; pl. 34, fig. 13. Type locality: Tertiary beds
of Petersburg, Virginia. (Miocene, fide DAL)

Aligena acquata Conrap, 1843. Dati, 1900, Trans. Wagner
Free Inst. Sci. 3 (5): 1175; pl. 24, figs. 8, 8a, 8b (of vol.
3, no. 4).

The following description is based on a single left valve
which is lot No. 144173 of the U.S.N. M. Paleontologi-
cal Collection. It is labeled “St. Mary’s River, Md.,
Miocene.”

Shell subtriangular, dorsal margins only slightly con-
vex, of about equal length, sloping abruptly at about the
same angle. Ends and ventral margin evenly rounded.
Umbo moderately inflated, protoconch large (but too
indistinct to measure), located midway the length of

Figure 1

Aligena aequata Conrap. Hinge of the specimen, USNM 144173,
from the Miocene of Maryland, which Dall studied

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TRE WERIGER

Vol. 11; No. 3

the valve, and slightly turned forward. Curvature of the
disc even, slightly flattened over the middle, but with no
trace of a sulcus, nor any appreciable expansion of the
anterior slope. No lunule, escutcheon or corselet. Sculp-

Figure 2

Interior of valve of same shell as Figure 1. Length 9.93 mm

ture of numerous, closely spaced growth rests, of un-
equal size, but periodically enlarged to form small con-
centric ridges. Valve thin for its size, inner margin smooth.
Hinge plate narrow, with a single finger-like tooth below
the umbo in this, the left valve. The tooth projects well
across the midplane. The resilial and tensilial parts of
the ligament may have been completely separated, with
the latter running along the very margin, where a
nymphal roughening is evident. The inner part attached
along a resilifer which begins below the umbo, and
diverges ventrally and laterally. The chondrophore is very
thickened, ending in a rounded point too far from the
median plane to be a tooth. Length (extrapolating for
the broken end) 9.93 mm; height 8.78 mm; semi-dia-
meter 2.84 mm.

Another lot, U.S.N.M. 112378, labeled “Aligena
aequata Conrap from the Pliocene, Waccamaw River,
South Carolina”, consists of a left and a right valve much

Figure 3

Exterior of valve of same shell as Figure 1

better preserved than the lot from the Maryland locality.
These valves are perhaps the ones figured by Dat, 1898,
pl. 24, figs. 8, 8a, 8b. The ridges on the specimens are
regular, but not quite as strong as he has shown them.
They have no differentiated nymphal attachment for the
tensilium, and the hinge plate is narrower than that of
the valve from the Maryland Miocene.

To evaluate the relationship of Aligena to Montacuta
Turton, 1822, specimens of M. substriata MontTacu
from the Jeffreys collection at the National Museum were
studied. This species, originally named Mya substriata
Montacu, 1808, is the type species of Montacuta Tur-
TON, 1822 by subsequent designation of Gray, 1847. I
have not seen Montagu’s work, but according to ForBEsS
& Haney (1850, 2: 77), the species was published in the
supplement, which appeared in 1808 (fide Parmer, 1958,
js DZ).

Montacuta substriata (Montacu, 1808)
(Figures 4 to 7)

The shell is minute (2.97 mm long, 2.84 mm high, 0.79 mm
semidiameter), thin, obliquely ovate, with the anterior
end longer than the posterior. The umbos are scarcely in-
flated, touching, and essentially orthogyrous. The proto-
conch is persistent, small and oval (0.125 mm long).
Equivalve, with the disc moderately and regularly in-
flated, somewhat polished, with a few widely spaced

Figure 4

Montacuta substriata (Montacu). A specimen from Zetland in the
Jeffreys collection (USNM 170517). Length 2.97 mm

growth rests. Major sculpture is of 3 to 10 widely spaced,
faint radial lines, slightly elevated, most prominent on
the middle third of the disk and toward the ventral
margin. Valve margin smooth and sharp; pallial line
wide, without a sinus and with even margins, joining the

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adductor muscle scars at their lower ends. Adductor scars
about equal in size, irregularly elongate, and with dorsal
projections which are probably the scars of the pedal
retractor muscles. Hinge short, with a single tooth in
each valve. The teeth are rounded, elongated ridges be-
ginning under the umbos and extending forward. That
of the right valve is larger, and the tooth at the left

Figure 5

Interior of same valve as in Figure 4

valve fits in a shallow, poorly defined socket in front of
and above the right one. A tensilial part of the ligament,
if present, must be small and without noticeable attach-
ment on the shell. The resilium is a flat ribbon, its attach-
ments beginning at the umbo and sloping downward and
laterally. The mid-ventral part of the resilium isa calcified
strip.

Thus, the resilium is very similar to that of Aligena,
which it also resembles in lacking lateral teeth, lunule and
escutcheon. But these negative characters are outweighed
by the differences between this species and Aligena
aequata: Montacuta substriata is a much smaller shell,
with small protoconch and umbos orthogyrous rather

Figure 6

Interior of left valve of same shell shown in Figure 4

than prosogyrous. Growth striae are so poorly developed
that the surface appears polished, whereas all species in-
cluded in Aligena as here defined and limited, have
moderate to prominent growth striae, giving the shell a
silky texture, or even expanded to form concentric ribs.
The pallial line in Montacuta is smooth and continuous,
but the dorsal margin is ragged in all species of Aligena
in which I have been able to study it. The radial sculpture
of Montacuta is lacking in Aligena, and there is no finger-
like projection of the teeth which are nearly always
present in Aligena.

Another species of the eastern Atlantic generally in-
cluded in Montacuta, M. ferruginosa (Montacu, 1808),
has finger-like cardinal teeth projecting normal to the
median plane of the shell, as well as low lamelliform
bases of these teeth, continuing forward along the hinge
plate. This may be why Tutete (1935) included Aligena
in Montacuta. Montacuta ferruginosa is an elongate shell,
with smooth surface, smooth margined pallial line, small
size and orthogyrous protoconch, thus showing similari-
ties to M. substriata. Its resilium is somewhat shorter
than in M. substriata, the attachments diverging down-
ward and outward more abruptly, so that it approaches
the transverse ligament of Mysella. The differences in
anatomy of the soft parts between Montacuta substriata

Figure 7

Ventral view of the hinge line of a shell of Montacuta substriata

from the same lot as Figure 4, which had been fortuitously broken

in such a way that the interlocking of the teeth and the resilium,
with lithodesma, can be seen

and M. ferruginosa as detailed by OLpFIELD (1961) are,
when considered in conjunction with differences in the
shell, sufficient to suggest that the two species may ulti-
mately be placed in different genera.

The loss of teeth may be a secondary character within
the leptonid bivalves, which has occurred several times
independently. At least one true Aligena, A. cokeri, is
consistently edentate. We may suppose that ancestral pop-
ulations may have had true lateral teeth (1.e. no pro-
jections of one valve margin below the other) and a
cardinal tooth in each valve which had an elongated base
extending forward under the hinge margin, with a digiti-
form projection normal to it at its umbonal end. The
lateral teeth were lost in some later populations (including
Montacuta and Aligena), the digitiform process lost in

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Vol. 11; No. 3

M. substriata (but retained in M. ferruginosa), the elon-
gate base lost in most Aligena, but not A. coker: and
A. (?) borniana.

Montacuta substriata and M. ferruginosa are the only
two species recognized in that genus by WINCKworRTH
(1932) in his list of British Marine Mollusca. A third
species was included with doubt. According to ForBES &
Hantey (1850), M. ferruginosa is known only from Brit-
ish seas, and lives subtidally to at least 30 fathoms.
Montacuta substriata was known to them from Britain
and Scandinavia, but not southward. They report it from
depths of 5 to 140 fathoms. Orprietp (1961) gave an
extensive anatomical account of both species, and noted
that they are commensals on sea urchins. Insofar as com-
parison is possible, there are few significant differences
in the anatomy of the soft parts of M. substriata and
Aligena texasiana (see below).

SPECIES or THE WESTERN ATLANTIC

Aligena elevata (Stimpson, 1851)

(Figures 8 and 9)

Montacuta bidentata “Turton 1822” of Goutp, 1841, In-
vert. Mass. Ed. 1, p. 59, not of Turron (whose species
is Mya bidentata Montacu, 1803, now placed in
Mysella) .

Montacuta elevata Stimpson, 1851. Shells of New England,
p. 16 (merely renamed the species misidentified by
GouLp).

Kelliopsis elevata Stimpson, 1851. VerRRILL & BusuH, 1898,
Proc. U.S. Nat. Mus., 20: 784; pl. 93, figs. 2-4; pl. 94,
figs. 7, 8.

Aligena elevata Stimpson, 1851. Dati, 1899, Proc. U.S.
Nat. Mus. 21: 879, 884; 1900, Trans. Wagner Free Inst.
Sob, 3) (@))e Jhl77

The shell is small (length 5.84 mm, height 4.75 mm, dia-
meter 3.34 mm), equivalve, moderately inflated, oval to
subtriangular, with the umbos moderately prominent,
slightly closer to the hind end, and distinctly prosogyrous.
The dorsal and ventral margins are only slightly convex,
the front and hind margins more strongly rounded. The
disc is evenly convex, with no median sulcus. Sculpture of
fine, somewhat irregular growth lines, giving the shell a
silky texture. There is a very thin, light tan periostracum.
The protoconch is oval, about 0.4 mm long.

The hinge line is narrow, with a prominent finger-like
cardinal tooth, projecting normal to the median plane.
The resilifer is typical of the genus, and a tensilium at-
taches to the very margin of the shell over it. The dorsal
margin of the broad, simple pallial line is ragged.

Figure 8

Aligena elevata (Stimpson), interior of a right valve from Chelsea
Beach, Massachusetts (USNM 159288). Length 5.81 mm

Figure 9

Exterior of same valve as Figure 8

This species is represented in the National collection
from 16 lots ranging from Massachusetts to Long Island,
from the shore to 10 fathoms.

Aligena texasiana Harry, spec. nov.
(Figures 10 to 14)
Shell small (length 4.81 mm, height 3.75 mm, diameter

2.50 mm), white, chalky, moderately inflated; equivalve,
sculptured only with numerous, irregularly spaced, fine

Vol. 11; No. 3

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growth lines. Subtriangular to suboval or subquadrate,
almost equilateral, the beaks being slightly closer to the
hind end, but distinctly turned forward. The dorsal mar-
gins are very slightly convex, the hind one somewhat
shorter than the front one. Both the front and hind ends
of the shell are evenly rounded, and of nearly the same

Figure to

Aligena texasiana Harry, spec. nov. Holotype. Length 4.81 mm

convexity. They continue as a smooth curve into the
ventral margin, which is almost straight, but shows a
slight sinus midway its length, at the end of the sulcus.
The region of the disc between the umbo and antero-
ventral part of the margin is somewhat more inflated
than the rest of the disc. From the umbo to the middle
of the ventral margin is a shallow, vague sulcus, only
prominent on the lower half of larger shells. The proto-
conch is prominent, oval, about 0.348 mm in diameter.

Figure 11

Aligena texasiana Harry, spec. nov. Holotype

The umbos touch, and there is no lunule or escutcheon.
The valves are thin, with smooth inner surfaces and mar-
gins. Adductor scars are prominent, suboval, and about
equal in size. Retractor scars are adjacent or confluent
(depending on growth) with them dorsally. The pallial
line is prominent, without sinus, and broken into a series
of subtriangular marks probably representing discrete
muscle bundles. The resilium is flattened, short, attached
by its long edges to each valve, along a slight ridge just
under the dorsal valve margins and behind the umbos.
The resilial attachments descend and diverge posteriorly.
The medial third of the ventral surface of the resilium is
calcified along its axis. There is a single tooth in each
valve, rounded and projecting slightly forward across the
midline. The one in the right valve is larger, and passes
in front of the left tooth. A thin, light tan periostracum
on fresh shells gives them a satiny luster.

Figure 12

Aligena texasiana Harry, spec. nov. Paratype, Length 5.13 mm

Type Locality: East end of West Bay, Galveston, Texas.
The holotype is USNM no. 679103. This species has been
found only in West Bay and Lower Galveston Bay at
Galveston. It was not found in more than 60 samples
ranging from 3 to 18 fathoms offshore. Within the bays,
it occurs from the lowest tide level to 7 feet, which is
about the deepest water of the bays, outside of the passes
and dredged channels. It seems to be only local within the
bays, and is not common. Frequently polychaetes are
abundant in the samples containing it, suggesting it may
have some mutualistic relationship with them, but details
are lacking. It also occurs with Mysella planulata, with
which it is easily confused. The latter is more flattened,
has opistogyrous umbos with a smaller protoconch, and
is without a sulcus on the disc. PARKER (1959, pp. 2128-

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WBE VEEIGER

Vol. 11; No. 3.

2129) evidently confused them, labeling a figure of Alz-
gena texasiana with the name “Mysella planulata” (l.c.,
figs. 21a, 21b). His specimens were from bays farther
south on the Texas coast, but no exact localities are in-
dicated. This species also occurs in Barataria Bay, Louisi-
ana, where I recorded it as Aligena sp. (Harry, 1942).

From Aligena elevata, this species differs in being gen-
erally smaller, more inflated, and with a median sulcus on
the disc. Aligena texasiana is perhaps closest to what I
have called A. nucea Datu from the west coast of Mexi-
co, but the latter is not as inflated and the sulcus is less
prominent. The Texas species is smaller than A. cokeri,
with the sculpture and sulcus not as prominent, and
cardinal teeth seem always to be present.

ANATOMICAL NOTES

In the laboratory, Aligena texasiana is moderately active,
extending the foot and pulling its body, resting on its
side, up to it. It crawls up the vertical wall of the glass
dish, and up large shell fragments, attaching itself very
loosely by a minute byssal thread or two. In the active
animal, the siphonal opening is not evident, and my study
did not extend to the direction of water currents. The
mantle margin is exposed along the edge of the shell
only to show a row of minute, closely spaced papillae,
scarcely longer than their diameter, uniformly distributed
along the edge of both mantle lobes between the adductor
muscles.

Figure 13

Aligena texasiana Harry. Postventral view of an animal from
which the shell has been decalcified

Figure 14

Aligena texasiana Harry, spec. nov.
Diagram of the superficial anatomy of a specimen from which the
shell has been removed

Symbols used in Figures 13 and 14:

aad anterior adductor muscle
ar anterior pedal retractor muscle

d diaphragm

f foot

fl fused lamellae of mantle margin

g gill
im dorsal margin of ascending gill lamella
k_ kidney

Ip labial palp
mr retractor muscle of mantle margin
np nascent periostracum
P periostracum
pa posterior adductor muscle
pr posterior pedal retractor muscle
s siphonal opening (excurrent)
vg visceral ganglion
vh ventricle of the heart

The flesh of the living animal internally is snow white,
except that the liver is colored light brown, and the kidney
is faint tan. A byssal gland is not evident in the living
animal, but there is a groove along the keeled ventral
margin of the foot. The foot is somewhat finger-shaped,
although short and rounded when retracted.

Studies on specimens preserved in 70% isopropyl al-
cohol were made after the shell was dissolved in a very
weak hydrochloric acid (2%) solution in alcohol of
70% strength. The outer, or true, periostracum is thin,
transparent, light tan colored, with a roughened outer
surface in harmony with the growth ridges of the shell.
The free edge of this layer of periostracum is held firmly
in the periostracal groove between the outer and middle
lamellae of the mantle margin. Since the latter has re-

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tracted some distance from the shell margin, there is an
abrupt turning of the outer periostracal sheet, along the
line of the shell margin, and a wide strip of nascent peri-
ostracum between it and the mantle margin.

An inner periostracal sheet seems to adhere closely to
the outer surface of the mantle in specimens deshelled in
acid, but it was probably separated from the mantle by
the innermost layer of calcareous material, the hypostra-
cum. I therefore propose for this sheet of periostracum
the name mesostracum. In Aligena texasiana it is smooth,
tough, pliable, thin, and very faint tan colored. It seems
to extend to the very margin of the mantle.

The margins of the two mantle lobes are fused for a
short distance below the posterior adductor muscle. The
fusion is in the form of a thin, transverse sheet, with a
large round hole at its upper end. This hole has a smooth
margin and is the excurrent siphonal opening. The papil-
lae extend along the sides of the sheet, and the two rows
of them join above the siphonal opening.

The mantle margin is thickened, but the rest of the
mantle is very thin, almost transparent. Discrete bundles
of retractor muscles can be seen in it, extending upward
for a short distance, and originating on the valves of the
shell along the pallial line.

-The gills are represented only by a single huge subtri-
angular demibranch on each side. There is no muscular
suspensory septum. Each demibranch has about 50 fila-
ments, the longest being anterior, and the others shorten-
ing in length progressively toward the hind end. The
lamellae are not pleated. They are eulamellibranchian,
with firm inter-filamentary junctions orderly arranged
horizontally to give the gill a latticed appearance (not
shown in Figure 14). The free margins of the demi-
branchs are grooved. The anterior filament of each is
attached along the visceral mass throughout its length.
The ascending lamellae are somewhat narrower than the
outer, descending ones. They attach along the visceral
mass in the front part of the gill, and behind it to each
other. The hind tips of the gills do not reach quite to the
siphonal opening, but there is a triangular, horizontal
sheet of tissue extending from the hind tip of the gills to
the fusion of the mantle lobes just below the opening.
Thus, the mantle cavity is completely separated into in-
current and excurrent chambers by a structurally com-
plete diaphragm.

The labial palps are minute triangles, which overlap
only the first few filaments of the gill. The ventricle of
the heart surrounds the rectum, the kidney is between
the pericardium and the posterior adductor muscle.

No internal detail of the viscera has been determined.
On the surface of the visceral mass, there were a few
low projections, which could as easily be interpreted as

phenomena of contraction rather than as projections of
the liver and gonads. The latter condition has been de-
scribed in other leptonids (OLpFIELD, 1961).

A specimen collected in early spring had several hund-
red early embryos in the gills, but there was no evidence
of embryos in specimens examined in June and July.
OLDFIELD (of. cit.) describes a sequential shift in sexu-
ality of some British leptonids.

SPECIES or tHe EASTERN PACIFIC

: Aligena cerritensis ARNOLD, 1903
(Figures 15 to 17) .

Aligena cerritensis ARNOLD, 1903. Mem. Calif. Acad. Sci. 3:
138; pl. 13, fig. 3. Type locality: Los Cerritos (Signal
Hill) near Long Beach, Los Angeles County, California.
(Pleistocene)

The following description is based on the holotype USNM
162529, a single right valve (Figure 15).

Figure 15

Aligena cerritensis ARNOLD.

Holotype, USNM 162529. Length 9.03 mm

Shell large for the genus (length 9.03 mm, height 8.39
mm, semidiameter 2.58mm), obliquely ovoid, the pos-
terior dorsal margin sloping downward at a greater angle |
with the horizontal than the anterior dorsal one. The
shell is unusually thick, and the inside of the disc, above
the pallial line, has an irregular surface. The chevron-
shaped origins of the pallial retractor muscle give the
upper part of the pallial line an irregular appearance.
These are actually engraved on the thickening of the disc.
The adductor scars seem to be large and slightly kidney-

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Vol. 11; No. 3

shaped. The resilifer is as in Aligena, and the shell ridge
along it is large. There is a single, finger-like tooth in
front of the ligament in this, the right valve, projecting
normal to the midplane and across it. Outer surface
slightly flattened over the middle of the disc, smooth,
with only a few irregular growth rests on the lower
third of the shell. No lunule or escutcheon.

This species is living on the coast of Lower California
and on the mainland opposite. Some of the 14 lots of

Figure 16

Aligena cerntensis ARNOLD
A Recent shell from “off Lower Calif? USNM 212588 to show the
steep slope of the postdorsal margin. Length 5.63 mm

this species in the National Museum have depth data,
showing it occurs in from } to 5 fathoms, and in fine
grey or black sand. It may occur with Aligena nucea.
Fresh shells are rarely as thickened as the holotype. They
often have a periostracum which is much thicker than on
any other species of Aligena, and which may persist only
over the lower third of the shell. It is brownish grey in

Figure 17

Aligena cerritensis ARNOLD
A Recent shell from Magdalena Bay, Lower California (USNM
217809) showing papillate periostracum over lower third of valve.
Length 5.00 mm

some shells, light tan on others, and usually has small
triangular projections from its surface. Figure 17 is
drawn from a Recent shell having such a cuticle. The
protoconch is large. The shape is variable, and small
shells are difficult to separate from A. nucea. The beak
cavities are not as deep as in that species, there is never
a sulcus on the disc, and the ventral margin is always
slightly convex, never having a sinus. The umbos are
always well toward the hind end of the shell.

Aligena obliqua Harry, spec. nov.

(Figures 18 to 20)

Shell small (length 6.19 mm; height 6.19 mm, at right
angle to length; diameter 2.38 mm; protoconch oval,
0.284 mm long), white, covered with a light tan, very
thin periostracum. Moderately and regularly inflated,
obliquely oval, with the margins evenly rounded. Equi-
valve, inequilateral, with the umbos about a third of the

Figure 18

Aligena obliqua Harry, spec. nov.
Holotype, USNM 532795. Length 6.19 mm

length from the hind end. Umbos only moderately in-
flated, turned slightly forward, touching, with persistent
protoconchs. No lunule or escutcheon. Sculptured only
with growth striae, faint, close together, and mostly reg-
ular, with occasional (3 on holotype) growth rests more
prominently defined. The growth striae give the surface
a silky texture. Valves of moderate and even thickness,
margin smooth and sharp, adductor muscle scars very
elongate oval, with the foot retractor scars confluent with
their dorsal ends. Beak cavities deep, hinge very narrow.
A single prominent finger-like tooth arises just in front
of the umbo in each valve, projecting well across the
midline. The tooth of the right valve is larger than that

Wola ll: Nov 3

Figure 19

Aligena obliqua Harry, spec. nov.
Holotype, interior of valve

of the left, more directly under the umbo, and it passes
behind the left when the valves are joined. Ligament with
a thin, external part on the very margin of the shell, and
of the same length as the resilial part. The latter begins
under the umbo, its attachment slopes downward and
laterally, for not quite half the distance from the umbo
to the top of the posterior adductor scar. The ventral
surface of the resilium is strongly calcified. The shell is
thickened slightly along the resilial attachment.

Holotype, USNM no. 532795, is from Bacochibampo
Bay, Sonora, Mexico (Orcutt). This lot also contains 2
paratypes with joined valves and 10 disjoined valves. All
are in fresh condition, showing little evidence of weath-
ering.

The distinctive shape of this species easily separates it
from the other 3 living on the west coast. It is very similar

Figure 20

Aligcena obliqua Harry, spec. nov.
Dorsal view of a paratype from same lot as that of Figure 18

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Page 173

to Aligena minor Dat (1900, p. 1177, pl. 44, fig. 8), a
Miocene species from North Carolina, which differs
chiefly in being smaller, and not as oblique.

The National Museum has 6 lots of this species, from
Guaymas (Sonora) and Mazatlan (Sinaloa) on the main-
coast of the Gulf of California. Depths are indicated on
the labels as 4 to 14 fathoms, and substrate type of
medium-coarse grey sand.

Aligena nucea Dat, 1913

(Figures 21 to 24)

Aligena nucea Dati, 1913. Proc. U.S. Nat. Mus., 45: 597.
Not figured. Type locality: Gulf of California. Burcn,
1941, Nautilus 55: pl. 4, fig. 3 (photograph of holotype).
Otsson, 1961. Panamic-Pacific Pelecypoda, p. 234; pl.

The single left valve, which is the holotype of this species
(USNM no. 267149), is probably a more extreme variant
of the populations which I think constitute this species
than is ordinarily found in populations of Aligena. A
description of the holotype (Figure 21) is as follows:

Figure 21

Aligena nucea Datu. Holotype, USNM 267149. Length 3.88 mm

Shell small (length 3.88 mm, height 3.19 mm, semidia-
meter 1.31 mm), the umbo is very slightly behind the
midpoint of the length, slightly prosogyrous, and evident-
ly touched the one opposite. Protoconch large, persistent

as

Figure 22

Aligena nucea Datu. Dorsal view of holotype

but indistinct. Resilifer typical of Aligena, but unusually
large and prominent. There is a single finger-like tooth
just in front of the umbo. Sculpture of growth lines only,
more prominent on the lower third of the valve. Disc

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Page 174

Figure 23

Aligena nucca Datv. Interior of valve from Conception Bay, Gulf
of California (USNM 558735). Length 5.63 mm

curvature uniform, with no flattening or sulcus. Ventral
margin evenly rounded, with the post-ventral part slightly
flattened. No lunule or escutcheon.

It differs from Aligena cerritensis of the same length
in having a more swollen umbo, which is more centrally
placed. Aligena obliqua of this size is already markedly
oblique. From A. cokeri it differs in being dentate. Several
lots of specimens have accumulated in the National Mu-
seum since Dall named this species, which, though poorly
represented by the holotype, seem nonctheless to be con-
specific with it.

Figures 23 and 24 are drawn from a valve of USNM
558735 from Conception Bay, Gulf of California. The
shell is about equally rounded at both ends, with the
ventral margin flattened, or more often with a slight sinus
in the middle. There is usually a vague, very shallow
and broad sulcus on the lower half of the middle of
larger valves, and the anterior slope of the disc sometimes
is slightly inflated. The texture is silky, formed by very
fine, closely spaced and somewhat irregular growth lines.
The cardinal tooth is always present. The umbos vary in
position, may be slightly in front or slightly behind the
midpoint of the length.

Figure 24

Aligena nucea Datu. Exterior of same valve as in Figure 23

Vol. 11; No. 3

Eleven lots of this species at the National Museum
are all from the Gulf of California, with depths indicated
as 1 to 7 fathoms, and substrate as fine grey or black

sand. Otsson (1961) notes it extends south to Nicaragua.

Aligena cokert Datx, 1909
(Figures 25, 26)

Aligena cokeri Datu, 1909. Proc. U.S. Nat. Mus. 37: 155-156;
pl. 28, figs. 5, 6. Type locality: “Attached to worm tubes
thrown upon the beach of the lagoon at Capon, Peru.”
Oxsson, 1961. Panamic-Pacific Pelecypoda, p. 234; pl.
33, figs. 6, 6a, 6b.

Shell small (Holotype, USNM 207759, length 7.74 mm,
height 6.45 mm, diameter 5.68 mm), very inflated, ends
of about equal curvature, rounded, ventral margin only
slightly curved, and with a rounded shallow notch in the
middle, where the sulcus ends. Beaks strongly prosogyrous,
slightly in front of the midpoint of the length. Proto-
conch persistent and large (about 0.38 mm long, slightly
oval). A vaguely defined, moderately wide furrow runs
from the umbo to the midpoint of the ventral margin, is

Figure 25

Aligena cokeri Dati. Holotype, USNM 207759. Length 7.74 mm

scarcely evident on the umbones, becomes more pro-
nounced on the lower 3 of the shell. Growth lines very
prominent, standing up as small, sharp ridges, but some-
what irregular, though closely spaced; the shells tend to
be peculiarly maleolated in some places, with vague, ir-
regular depressions and elevations. In other lots than the
type, the growth-line sculpture may be very regular or
nearly absent, and the depressions and elevations absent,
or variously prominent. In some specimens from the
northern part of the range, the growth lines are absent,
the bumps and hollows so pronounced and numerous that
one might almost proclaim it a different species, were
there no connecting intergrades. The form may become

Vol. 11; No. 3

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Page 175

subtriangular in these, and the sulcus on the disc be
absent. The resilifer is typical of the genus; in some para-
type shells with both valves connected, a tensilium is evi-
dent, but this is scarcely more than a slight thickening of
the periostracum, joining the valves over the resilium.
The finger-like tooth of Aligena is absent in this species,

Figure 26

Aligena cokeri Datu. Hinge of right valve of a syntype

which has a slight, rounded ridge running forward from
the umbos along the hinge. The ridge of the left valve is
smaller than that of the right one. Neither ridge projects
across the median plane.

The edentulous condition was noted by Dati (1909)
and Orsson (1961, pl. 33, fig. 6) has illustrated it with
a good photograph.

Besides the type lot, there are several others at the
National Museum ranging from the Gulf of California to
Panama. Some labels indicate depths of 1 to 4 fathoms.

SPECIES or tHE WESTERN PACIFIC
AND INDIAN OCEANS

Aligena salamensis (JAECKEL & THIELE, 1931)

(Figures 27 to 29)

Montacuta salamensis JAECKEL & THIELE, 1931. Wiss. Ergeb.
d. deutsch. Tiefsee- Expedit. 21 (1): 226; pl. 4, fig. 98.
Type locality: Dar es Salam, 50 meters deep.

Ture.e (1935) noted that this species from the coast of
East Africa might belong to Aligena. Several lots in the
National Museum from southeastern Asia seem to fit it,
insofar as comparison with the meager description and
figure of JAECKEL & THIELE (in THIELE & JAECKEL,
1931) allow comparison.

Figure 27

Aligena salamensis JAECKEL & THIELE.
China Sea, 230 fathoms, off Pratas Island. USNM 297075.
Length 9.42 mm

Shell small (length 9.42 mm, height 8.39 mm, semidia-
meter 3.57 mm), very inflated, subquadrate in profile,
with margins evenly rounded. Beaks about midway the
length, turned forward, touching, the protoconch oval,
large, smooth, and persistent. Ventral margin slightly
sinuous, where the sulcus of the disc meets it. The sulcus
is prominent only over the lower half of larger shells,
and the anterior slope of the shell is slightly more inflated
than the rest of the disc. Sculpture of closely placed
growth lines of uniform size, which become coarser on
the lower half of the shell.

Figure 28

Aligena salamensis JAECKEL & THIELE.
Dorsal view of valve drawn in Figure 29

Valve thin, margin smooth and sharp. The hinge has a
large fingerform tooth projecting below the umbo; the
resilial attachment slopes downward and outward from
the umbo, and is buttressed by a thickening of the shell
along it, ending in a prominent angle at the hind end of
the resilium.

Page 176

Young specimens are evenly inflated and almost circu-
lar. They lack a median sulcus, and might be thought
another species, were it not for connecting intergrades.
The National Museum has 4 lots from the China Seas, off
Pratas Island, ranging from 88 to 230 fathoms, and 7 lots
from the Philippines, ranging from 10 to 37 fathoms. A

Figure 29

Aligena salamensis JAECKEL & THIELE.
A valve from 32 fathoms S.E, of Bantayan Island, Philippines
(USNM 293038). Length 4.69 mm. This was drawn at twice the
magnification of Figure 27

single lot from the Buton Strait, Celebes, is from 37
fathoms also. Most lots have only one or two valves, the
maximum being six. All of the smaller shells are from the
greater depths, but there are sufficient intergrades to
demonstrate they are conspecific with the larger ones.
These were collected in 1909 by the research vessel
“Albatross.”

This species seems to penetrate to greater depths than
others in the genus, but its shell is very characteristic of
Aligena in all respects.

SPECIES INCERTIS SEDIS

Aligena (?) borniana Datt, 1908
(Figures 30, 31)

Aligena borniana Dat, 1908. Bull. Mus. Comp. Zool. 43 (6) :
413; pl. 10, fig. 2. Type locality: Pacific Ocean, 16°32’ S
latitude, 119°59’ W longitude, 2012 fathoms.

Shell large for the genus (length 14.2 mm, height 10.3 mm,
semidiameter 3.6mm; protoconch oval, about 0.44 mm
long), elongate oval, the front and hind margins evenly

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Vol. 11; No. 3

Figure 30

Aligena (?) borniana Day. Holotype, USNM 110585.
Length 14.2 mm

rounded and of nearly the same arc, the ventral margin
almost straight. Valves inflated, of the same size, beaks
about + the length from the hind end, prosogyrous and
touching each other.

Very closely spaced growth lines and a light tan peri-
ostracum give the outer surface a silky texture. Growth
lines are more prominent near the beaks than elsewhere.
No lunule or escutcheon, but there is a narrow, vaguely
defined sulcus on the posterior slope on the upper third
of the shell only. The middle of the disc is slightly flat-
tened, but shows no trace of a sulcus (contrary to Dall’s
description). No evidence of a tensilium on the disjoined

Figure 31

Aligena (°) borniana Datu. Hinge of holotype
with resilium attached to left valve

Vol. 11; No. 3

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Page 177

valves of the single available specimen, the holotype.
The resilium is typical of Aligena, but seems not to have
a calcified strip. The hinge line in front of the umbo is
slightly thickened, more so in the right than in the left
valve, but there is no finger-like projection, and neither
thickening extends across the midplane. The adductor
scars and pallial line could not be distinguished.

The relatively compressed appearance of this species,
its large size, the faint sulcus on the posterior slope, as
well as the great depth at which it occurs, are characters
which, if relatively insignificant, argue against this species
being an Aligena. The lack of a projecting tooth typical
of most species of Aligena is not significant, since A.
cokeri has no such tooth. This might be placed in Mont-
acuta, except for the large size and the faint posterior
sulcus. The latter suggests a corcelet, and thus the Thya-
siridae, but the shape is unlike that group.

SPECIES REMOVED From Aligena

Three species have been described as Aligena which
belong elsewhere. Two of these are probably Thyasiridae:
Aligena redondoensis T. Burcu, 1941 and A. piswm Dat,
1908. The third, A. borealis Cowan, 1962 may belong
to that family. The three are sufficiently distinct from
each other to merit being placed in separate genera.

The smaller species of Thyasiridae may be covered
with iron encrustation, a character which has not been
found in Aligena, although it is known to occur in Mont-
acuta ferruginosa (see OLpFIELD, 1961). The thyasirids,
at least the smaller ones, are generally found at greater
depths than most species of Aligena, and occur in cooler
waters when living in shallow depths. A lunule, escutcheon
and corcelet (see CarTErR, 1967, for a discussion of these
characters) may occur in various combinations and mani-
fest to varying degrees in the Thyasiridae, but they are
absent in Aligena.

Axinulus VERRILL & Busu, 1898

Axinulus (?) pisum (Datt, 1908)
(Figures 32, 33)

Aligena pisum Datt, 1908. Bull. Mus. Comp. Zool. 43 (6) :
413. Not figured. Type locality: “Magellan Strait, in 61
fathoms.”

Only the holotype of this species, a slightly worn left
valve, is known. In a large series at the National Museum
of unidentified shells dredged near the Philippines by the
“Albatross,” there are a few valves which are very similar,

if not identical (USNM 300962, Lagonoy, G., E. Luzon,
Philippines, 569 fms.). The following description is based
on the holotype.

Figure 32

Axinulus (?) pisum DALL.
Holotype, USNM 110715. Length 2.43 mm

Figure 33

Axinulus (?) pisum Datu. Holotype, interior view

Shell small (length 2.43 mm, height 2.69 mm, semi-
diameter 0.82 mm), subcircular, white and opaque with
a very light tan, thin periostracum to which no dirt is
adhering. Umbo moderately inflated, and slightly proso-

Page 178

gyrous. The protoconch is scarcely evident, but seems to
be slightly oval, about 0.156 mm long. The curvature of
the disc is regular, with no lunule, escutcheon, or corce-
let evident externally. Surface almost smooth, showing
only a few very faint growth rests. The valve is thin, its
edge smooth, Adductor scars and pallial line not evident.
There is a faint but distinct ridge, ill-defined but fairly
broad, extending from the umbonal cavity near the pos-
terior dorsal margin, very suggestive of a corcelet. Hinge
plate narrow, with no evidence of a tensilium, but with
a resilifer beginning just under the umbo and sloping
downward and outward for a moderate length. A single
irregular tooth is immediately below and in front of the
umbo. The hind part of the tooth may be a broken,
finger-like projection, but it appears to have several
rounded bosses, one of which is on the top side. A
triangular extension forward of this tooth has a moder-
ately sharp ventral border, along which are several
incised lines sloping upward and backward.

Tomburchus Harry, gen. nov. in Thyasiridae

Type Species: Aligena redondoensis T. Burcu, 1941.

Shells small, oval, equivalve, inequilateral, with the
umbos nearer the hind end, and prosogyrous. An escutch-
eon is well defined by a sharp angle on the posterior slope,
extending from the umbo to the hind end of the shell.
No lateral teeth, but the postdorsal slope of the left
valve extends under the margin of the right one. The left
valve margin is also deflected under and just before the
umbo, as a tooth-like lamella with bifid margin, passing
under a small, rounded, tooth-like extension of the mar-
gin of the right valve. Ligament superficial, but covered
by a turned up margin of the shell, extending along the
adumbonal half of the escutcheon. The genus is named
in honor of Dr. Tom Burch, of a family whose several
members have greatly stimulated and furthered the study
of malacology.

Tomburchus redondoensis (1. Burcu, 1941)
(Figures 34 to 37)

Aligena redondoensis T. Burcn, 1941. Nautilus 55: 50 - 51; pl.
4, figs. 5, 6, 7. Type locality: 75 fathoms off Redondo
Beach, California, about latitude 33°38’50” W, longitude
118°26’30” N.

The paratypes of this species cited by T Burcu (1941)
as sent to the National Museum were misplaced and un-
available to me when I studied Aligena there in 1968.
Two lots of the National collection identified by Dr. A.

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Vol. 11; Nor3

M. Keen and cited by T: Burch were studied instead. One
of these, USNM 211882 (48 fms., off Santa Rosa Id.,
Calif.) has only a single very worn valve, but is possibly
this species. The other, USNM 331216a (129 fms., off La
Jolla, Calif), has two specimens which fit very closely
the figures and description of Aligena redondoensis. The
following description is based on that lot.

Figure 34

Tomburchus redondoensis (T. Burcu)
From 128 fathoms “off La Jolla, Calif” (USNM 331216a).
Hinge of left valve. Length 3.10mm

Shell small (length 3.10 mm, height 2.53 mm, semidia-
meter 1.14 mm), oval, equivalve, inequilateral, with the
umbos slightly closer to the hind end, moderately inflated
and prosogyrous. Protoconch persistent, oval, about 0.158
mm long. Valves white, moderately inflated, sculptured
only by fine growth lines, with a thin periostracum to
which mud and iron deposits adhere tenaciously in a few
places. The margin of the shell is formed by one contin-
uous curve, somewhat more flattened anterodorsally and
along the ventral margin, but closed by the more flattened
segment of the postdorsal margin, which meets the vent-
ral margin in a rounded angle, about halfway between
the top and the bottom of the shell.

(i

Figure 35

Tomburchus redondoensis (T. BurcH)
Top view of same valve as in Figure 34. The dashed line is the
limit of the postdorsal margin inserted below that of the right valve

No lunule is differentiated. A narrow, lenticular es-
cutcheon is well defined by a sharp margin and deeply
excavated surface, which is pinched up into a keel along
the umbonal half of the shell margin. There is no corce-
let. The ligament seems to consist of one piece, with the

tensilium and resilium possibly fused. It is about } as

Vol. 11; No. 3

long as the escutcheon, and attached to the inner side of
the pinched up part of the shell margins. It is effectively
concealed by this keel. There are no lateral teeth as
such, but in the left valve the posterior half of the dorsal
margin within the escutcheon is extended across the me-

Figure 36

Tomburchus redondoensis (T. BurcH)
Right valve of same shell as in Figure 34

dian plane, and bent slightly downward. It fits under the
margin of the right valve, which does not cross the me-
dian plane there. The margin of the left valve is also
extended just before and under the umbo, as a flattened,
bifid tooth. Seen from above, this “tooth” is narrow,
elongate along the shell margin, with its free margin
smoothly rounded. In the right valve there is a small,
hemispherical extension of the shell margin, which does

Figure 37

Tomburchus redondoensis (T. Burcw)
Top view of a shell with valves joined, from same lot as Figure 34.
Darkened areas are detritus

not extend across the median plane, but fits in the notch
of the tooth of the left valve. The adductor scars and
pallial line could not be studied.

A few lots of unidentified shells at the National Mu-
seum, dredged near the Philippines by the “Albatross” in
1909, evidently are this species or a closely related one, but

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Page 179

the present study could not be extended to include them.
Another member of Tomburchus may be Axinus dubius
DAUTZENBERG & FiscHER, 1897 (10: 215 - 216, pl. 6, figs.
18 to 21; renamed Thyasira dubia (DAUTZENBERG &
FiscHer by DauTzenseErG, 1927: 312 - 313, pl. 8, figs.
35 to 38) from the Azores, but I have seen no authentic
specimens.

“Lucina” ferruginosa Forses, 1843, as described and
figured by Forpes « HANLEy (1850, 2: 60 - 62, pl. 34,
fig. 1) from Britain may also be of this genus. WincK-
worTH (1932, p. 242) puts Forses’ species in Thyasira.
VERRILL & BusH (1898, p. 793, pl. 87, figs. 7, 8) cite
what may be the same species as “C'ryptodon (Axinulus)
ferruginosus (ForBEs)”, but they give no description and
the figure has insufficient detail.

Odontogena Cowan, 1964

Odontogena borealis (Cowan, 1964)
(Figures 38 to 40)

Aligena (Odontogena) borealis Cowan, 1964. Veliger 7 (2):
108 - 109; pl. 20, figs. 1, 2. Type locality: Georgia Strait,
British Columbia, Canada, 190 fathoms.

The following description is based on the single paratype,
disjoined valves of one shell, which is USNM 657130.
Shell small (length 2.56 mm, height 2.37 mm, semi-
diameter 0.63 mm), subcircular, resembling a minute
Lucina, with slightly convex, long post-dorsal margin.
The moderately convex shorter anterior margin meets the
umbo in such a way as to give the dorsal margin a notched
appearance. No lunule, escutcheon or corcelet. Inflated,
and covered with a ferruginous deposit which had been
mostly removed, to show a faint tan surface of silky
texture with a few irregular growth lines as the only

Figure 38

Odontogena borealis (Cowan). Paratype, USNM 657130
Length 2.56mm

Page 180

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Vol. 11; No. 3

sculpture. Beaks touching, slightly turned forward, about
midway the length of the shell. Valves thin, the margin
acute and smooth. Adductor scars and pallial line could
not be made out. Hinge line very narrow, poorly mani-
fest; ligament entirely internal, elongate, flattened, at-
tached along its side to the valves, beginning at the umbos
and extending a short distance behind; the attachments
do not diverge posteriorly. Teeth two in each valve; these

Figure 39

Odontogena borealis (Cowan) .
Right valve of same shell as in Figure 38

are large masses, poorly defined, as are the sockets into
which they fit. One tooth in cach valve is just under and
in front of the umbo, the other is behind the ligament.
Those in the right valve are bifid and receive in the vague
groove thus formed a short rounded projection from the
teeth of the left valve. The ligament does not appear to
have a calcareous midstrip, but this could be a matter of
age.

Figure 40
Odontogena borealis (Cowan).
Top view of shell in Figures 38 and 39

This species resembles Aligena of the east coast in type
of ligament, general shape, texture and size of shell, and
absence of lunule and escutcheon. The type of teeth,
however, removes it from that genus, as characters in this
group go, and the subgenus Cowan erected for it may be
used unless an earlier name is found.

LITERATURE CITED

ARNOLD, RALPH

1903. The paleontology and stratigraphy of the marine Plio-
cene and Pleistocene of San Pedro, California. Mem. Calif.
Acad. Sci. 3: 1-420; plts. 1-37

Burcu, THomas ADAMS

1941. A survey of the West American Aligenas with a descrip-

tion of a new species. The Nautilus 55 (2) : 48-51; plt. 4
Carter, Rosert M.

1967. On the nature and definition of the lunule, escutcheon

and corcelet in the Bivalvia. Proc. Malacol. Soc. London
37: 243 - 263

Conrap, TimotHy ABBOTT

1843. Descriptions of a new genus, and of twenty-nine new
Miocene and one Eocene fossil shells of the United States.
Proc. Acad. Nat. Sci. Philadelphia 1: 305 - 311

Cowan, Ian McTaccart
1964. A new species of the lamellibranch genus Aligena from
western Canada. The Veliger 7 (2): 108-109; plt. 20
(1 October 1964)
Dati, WitutAM HEALEY
1890-1903. Contributions to the Tertiary fauna of Florida.
Trans. Wagner Free Inst. Sci. 3 (1-6): 1654 pp.; 60 plts. [part
4, 1898, and part 5, 1900, contain the data relevant to the
present study]

1899. Synopsis of the Recent and Tertiary Leptonacea of
North America and the West Indies. Proc. U.S. Nat. Mus.
21: 873 - 897; plts. 87, 88

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 XIV. The
Mollusca and Brachiopoda. Bull. Mus. Comp. Zool., Har-
vard; 43 (6): 205 - 487; plts. 1-22 (October 1908)

1909. Report on a collection of shells from Peru, with a sum-
mary of the littoral marine Mollusca of the Peruvian zoological
province. Proc. U.S.N.M. 37 (1704): 147 - 294; plts. 20
to 28 (24 November 1909)

1913. | Diagnoses of new shells from the Pacific Ocean.
Proc. U.S. Nat. Mus. 45: 587 - 597

DauTzENBERG, PHILIPPE

1927. Mollusques provenant des campagnes scientifiques du
Prince Albert 1*® de Monaco dans lOcéan Atlantique et dans
le Golfe de Gascogne. Rés. Camp. Scient. Prince de Mo-
naco Fasc. 72: 1 - 400; plts. 1-9

DauTZENBERG, PHILIPPE & HENRI FISCHER

1897. Dragages éffectués par L’Hirondelle et par La Prin-
cesse-Alice. Mém. Soc. Zool. France 10: 139 - 234; plts.
3-7

Forses, Enwarp & SYLVANUS HANLEY

1848-1853. A history of British Mollusca and their shells. Van

Vorst, London; 4 vols.; illustr.

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Goutp, Aucustus ADDISON Otsson, AxEL ADOLF
1841. | Report on the Invertebrata of Massachusetts, comprising 1961. Mollusks of the tropical eastern Pacific particularly

the Mollusca, Crustacea, Annelida, and Radiata. Cambridge
(printed for the legislature): i-ili+ 1-373; 15 plts.
Gray, JoHn Epwarp
1847. _A list of the genera of Recent Mollusca, their synonyma

and types. Proc. Zool. Soc. London (for 1847) 17 [part 15]
(178) : 129 - 219 (November 1847)
Harry, Harotp WILLIAM
1942. List of Mollusca from Grand Isle, Louisiana, recorded
from the Louisiana State University Marine Laboratory, 1929-
1941. Occ. Pap. Marine Lab., Louisiana State Univ. 1:
1-15
Lea, Henry C.

1843. Descriptions of some new fossil shells from the Tertiary
of Virginia. Proc. Amer. Philos. Soc. 3: 162 - 165

1846. Descriptions of some new fossil shells from the Tertiary
of Petersburg, Virginia. Trans. Amer. Philos. Soc. 9:
229 - 274; plts. 34 - 37

Monracu, GrorcE

1803. Testacea Britanica or natural history of British shells,
marine, land and fresh-water, including the most minute:
systematically arranged and embellished with figures.
Romsey (J.S. Hollis). pp. i-xxxvii + 1-606; plts. 1-16
Supplement (1808): 183 pp.; plts. 17 - 30 [not seen]

OLpFIELD, E.

1961. The functional morphology of Kellia suborbicularis
(Monracu), Montacuta ferruginosa (Montacu) and M. sub-
striata (Monrtacu). Proc. Malacol. Soc. London 34 (5):
255 - 295

from the southern half of the Panamic-Pacific faunal pro-
vince (Panama to Peru). Panamic-Pacific Pelecypoda.
Paleont. Res. Instit. Ithaca, N. Y.: 574 pp.; 86 plts.
(10 March 1961)
PaLMER, KATHERINE VAN WINKLE
1958. ‘Type specimens of marine mollusca described by P. P
Carpenter from the West Coast (San Diego to British Colum-
bia). Memoir 76, Geol. Soc. Amer. i- viii + 1-376; plts.
1-35. New York, N. Y. (8 December 1958)
Parker, Ropert H.
1959. Macro-invertebrate assemblages of Central Texas coastal
bays and Laguna Madre. Bull. Amer. Assoc. Petrol Geol.
43 (9): 2100 - 2166
Stimpson, WILLIAM

1851. Shells of New England.
Boston, 56 pp.; 2 plts.

Phillips, Sampson & Co.,

THIELE, JOHANNES
1931-1935. Handbuch der systematischen Weichtierkundc.
Jena, pp. 1 - 1154; 893 text figs.
THIELE, JOHANNES & SIEGFRIED JAEGKEL
1931. | Muscheln der Deutschen Tiefsee Expedition. Wiss.
Ergebn. d. deutsch. Tiefs. Expedit. “Valdivia”, 1898 - 1899
21 (1): 161 - 268; pits. 6-10 [pp. 1-110; plts. 1-5, reprint
numbers]
Turton, WILLIAM

1822, Conchylia dithyra insularum Britannicarum. MA
Nuttall; 279 pp.; 20 plts.

VERRILL, Appison & KATHARINE JEANNETTE BusH

1898. Revision of the deep-water Mollusca of the Atlantic
coast of North America, with descriptions of new genera and

species.

WINCKworTH, RONALD

1932. The British marine Mollusca.

19 (7): 211-252

Proc. U.S. Nat. Mus. 20: 775 - 901; plts. 71 - 97

Journ. Conch. London

Page 182

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Vol. 11; No. 3

The Egg Capsules of Jenneria pustulata (Licutroor, 1786)

with Notes on Spawning in the Laboratory

CHARLES N. D’ASARO

Institute of Marine Sciences, University of Miami', Miami, Florida 33149

(1 Text figure; 1 Table)

INTRODUCTION

Jenneria pustulata (LicHtTroot, 1786) is a brightly
colored cypraeacean distributed in shallow water from
‘the southern end of the Gulf of California to Ecuador
(Keen, 1958). It is a hardy species normally found in
association with the stony corals upon which it feeds.
The relationship of the species to other members of the
superfamily is not well established. KEEN (of. cit.)
believes the shape of the radular teeth and other ana-
tomical characters can be homologized with those of
Trivia; however, SCHILDER (1936) places J. pustulata in
the Ovulidae (Amphiperatidae). Data on the structure
of the egg capsule and the larval stages can add to the
understanding of phylogeny in this species.

METHODS

Adult specimens of Jenneria pustulata were collected
by Dr. E M. Bayer of the Institute of Marine Sciences,
University of Miami, from Venado Island in the Bay of
Panama in August, 1965. Of this material, a pair (female
18mm, male 13mm) were transported alive to the In-
stitute, where they were maintained for three years. The
specimens were kept in an aquarium with running sea
water at ambient temperatures. Assorted coelenterates
were also kept in the same aquarium. Stony corals, in-
cluding Porites sp., Phyllangia americana MiLNe-Ep-
warps & Haine, 1850 and Siderastrea siderea ELLIS &
SoLANDER, 1786 were used as food. There was a marked
preference for Phyllangia over the the other species; how-
ever, all were eventually accepted. During the period of
observation, which extended from January, 1967 to May,

‘ Contribution No. 955 from the Institute of Marine Sciences,
University of Miami. This investigation was conducted under
the auspices of the U.S. Public Health Service (GM-125-41-02).

1968, the animals were not disturbed and had a con-
stant supply of food.

The egg capsules and embryos were preserved in 10%
sea water formalin. All drawings were made from pre-
served material with the aid of a camera lucida.

BREEDING HABITS ©

Spawning began when the water temperature exceeded
24° C for 3 to 4 weeks (Table 1). The breeding season
at Miami extended from April to December. During this
period, 20 egg masses were produced. The number of
capsules deposited per spawning ranged from 14 to 233
with an average of 125. A total of 2495 capsules was
produced. Copulation took place from about 4 days to
minutes prior to the beginning of spawning. The female
moved onto the walls of the aquarium away from the
sediment before beginning to spawn. No capsules were
found on the corals. During the annual period, this pro-
cess was repeated approximately twice a month.

Before selecting a site for oviposition, the female ex-
amines the walls of the aquarium for several hours. After
selection, an area slightly larger than the diameter of a
capsule is cleaned of periphyton and other surface de-
posits by the radula for about 8 minutes. Then the pro-
podium is folded over the cleared area, and a capsule is
passed from the oviduct down a ciliated groove on the
right side. Transport involves about one minute, The
propodial muscles knead the capsule while it is held in
place. Glandular cells secrete a layer of adhesive which
imparts the final shape. Cleaning begins again about 2
minutes before the capsule is completely attached. The
whole cycle involves approximately 20 minutes. At this
rate to produce the larger egg masses, oviposition must
continue for at least 3 days. Brooding, as noted among
related cypraeaceans by OsTerGAARD (1950), does not
occur.

Vol. 11; No. 3

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Page 183

Table 1

The Spawning of Jenneria pustulata at Miami, Florida

Number of
Capsules

Date per Mass Bay Temperature (° C)
January, 1967 0 21.51
February 0 21.5!
March 0 22.0!
April - 25.0!
April 26 120 26.5
April 30 14 25.0
May 21 ye) 28.0
May 31 101 28.5
June 19 72 28.0
June 26 85 30.0
July 4 50 2 29.5
July 7 81? 29.0
July 20 99 3 30.0
July 22 643 30.0
July 28 79 30.5
August 7 192 29.0
September 3 169 30.0
September 18 111 29.5
September 25 110 29.0
October 22 205 25.5
October 27 ala 26.5
November 9 233 25.0
November 20 202 24.0
December 5 152 23.0
January 1968 0 Zs)
February 0 20.0:
March 0 21.5%
April - DAs
April 179 26.0
™ = average; 7 = communal mass; 3 = communal mass

There is some indication that communal spawning
takes place in the natural habitat. In two instances, the
parent returned to an egg mass several days after the
initial spawning to deposit capsules (Table 1). In areas
with dense populations, it is possible that several females
will use the same mass. Communal structures can be
identified during development by noting that the contents
of the older oothecae have a uniformly darker pigment.

THE EGG MASS

Each cluster of oothecae forms a very irregular oval
with a long dimension varying between 5 mm and 30 mm.
The pustulate capsules are always placed in a single
layer. When overlapping between adjacent structures
occurs, only a small portion of the edge of each is in-

volved (Figures 1 a, 1b). The escape aperture is never
covered. Placement of the oothecae in the mass does not
vary. The side with the aperture is always placed between
2 capsules in a preceding row. The arrangement of the
capsules in long arcs reflects the movements of the parent
during spawning.

Imm

Figure 1

The Egg Capsules of Jenneria pustulata

a: dorsal view of 6 capsules from a typical egg mass. One capsule
contains embryos.
b: lateral view of 3 capsules.

Oothecae of Jenneria pustulata are transparent, color-
less, pustulate structures which are somewhat variable in
shape. The basal outline of each capsule is roughly the
shape of an obtuse triangle with rounded angles (Fig-
ure 1 a). Occasional capsules, ovate or round in outline,
are the result of the parent’s ability to mold the capsules
into a compact mass. An oval suture in the membranes
of the upper surface close to the obtuse angle marks the
escape aperture. Indistinct sutures extending laterally
from the aperture reflect the bilobed structure of the
oviduct (FRETTER & GRAHAM, 1962). The average di-
mensions of the capsules are: length — 2.3mm; width
— 1.7mm; and height — 0.5mm. Individual oothecae

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Vol. 11; No. 3

contain from 65 to 107 embryos and average about 90.
The number of embryos per mass varied between 1 200
and 21 000 with an average of 11 200. The total seasonal
production for the female was 224 500 embryos.

Development is indirect, with the formation of a
long-term planktotrophic veliger. The pinkish-brown
pigment which appears in the capsules after about 4 days
is the result of the development of a pigmented proto-
conch. Hatching begins after 13 or 14 days and is com-
pleted in 24 to 72 hours.

DISCUSSION

The egg masses of cypraeaceans have general characters
which are very uniform within a given genus; however,
in some families (as presented by KEEN, 1958) consid-
erable variation does occur. ‘These variations are recon-
ciled in part by suggestions to separate the groups with an
echinospira larva (see FRETTER & GRAHAM, 1962). A
number of cypraeids were examined by Lo Branco (1899),
VayYSSIERE (1927), and OsTERGAARD (1950). A survey
of these investigations and personal observations on the
spawn of Cypraca spurca acicularis GMELIN, 1791, C.
cervus LINNAEUS, 1771, and C. zebra Linnagus, 1758,
have shown that the structure of the egg capsules in this
family is very uniform. As noted by OsTERGAARD (of. cit.),
brood protection is also a characteristic of the family.
Erronea errones (LINNAEUS, 1758) (Narvarajan, 1958)
has a layered egg cluster quite similar to that of Cypraca
and also exhibits brood protection. The capsules of Trivia
arctica (Monracu, 1803) and T: monacha (pa Costa,
1778), which were studied by Lesour (1932 a), differ
greatly from those of their suggested close relative, Jen-
neria pustulata, in that they are vasiform and are im-
bedded individually in compound ascidians. Both species
have a characteristic echinospira which does not occur
in J. pustulata. In the Ovulidae, however, Simnia patula
(PENNANT, 1777) (LeBour, 1932 b) has pustulate cap-
sules arranged in a single layer like J. pustulata. Similar
structure and placement is also found in another ovulid,

Cyphoma gibbosum (LinNAEuS, 1758) (personal ob-
servation ) .

In conclusion, it can be stated that the structural ar-
rangement and placement of capsules by Jenneria pus-
tulata is most similar to the processes occurring in the
Ovulidae and that a close relationship is implied.

LITERATURE CITED

FRETTER, VERA, & ALASTAIR GRAHAM
1962. British prosobranch molluscs, their functional anatomy
and ecology. London, Ray Soc. xvi + 755 pp.; 316 figs.
Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i- xi + 624 pp.; illus.
Stanford Univ. Press, Stanford, Calif. (5 December 1958)
Lesour, Marie V.
1932a. The British species of Trivia: T: arctica and T: monacha.
Journ. Mar. Biol. Assoc. U.K. 18 (2): 477 - 484
1932b. The larval stages of Simnia patula.
Biol. Assoc. U.K. 18 (1): 107-115

Lo Branco, S.

Journ. Mar.

1899. Notizie biologiche riguardante specialmente il periodo
di maturita sessuale degli animali del golfo di Napoli.
Mitt. Zool. Stat. Neapel 13: 448 - 573

NatarajAN, A. V.

1957. Studies on the egg masses and larval development of
some prosobranchs from the Gulf of Mannar and the Palk Bay.
Proc. Indian Acad. Sci. B. 46: 170 - 228

OSsTERGAARD, JENS Matias
1950. Spawning and development of some Hawaiian marine
gastropods. Pacific Sci. 4: 75 - 115
ScHILDER, FRANZ ALFRED
1936. | Anatomical characters of the Cypraeacea which confirm
the conchological classification. Proc. Malacol. Soc. London,
22 (2): 75-112; 2 plts.
VAYSSIERE, ALBERT JEAN BapTisTE Marie

1927. | Recherches zoologiques et anatomiques sur les mollusques
de la famille des Cypraeidés. 2™° partie. Ann. Mus. Hist.
Nat. Marseille, Zool., 21: 133 - 184; plts. 24 - 28

Vol il INows

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Page 185

The Taxonomic Significance and Theoretical Origin

of Surface Patterns on a Newly Discovered Bivalve Shell Layer,

the Mosaicostracum

GEORGE H. HAMILTON

Department of Paleobiology, Division of Invertebrate Paleontology, Smithsonian Institution

Washington, D.C. 20560

(Plates 26 to 38)

INTRODUCTION

THE STRUCTURE OF BIVALVE SHELLS has been studied
with the light microscope at magnifications which allow
for a point resolution of 0.2 micron. However, structures
present in the shell which are consistent at the species
level have not been resolved. The body of knowledge
gained so far through use of the light microscope has
shown that shell structure is taxonomically consistent to
varying degrees at the ordinal level (NEWELL, 1965) and
in some cases (i. e. Ostreidae) at generic through familial
levels (GUNTER, 1950; CarPENTER, 1848; OBERLING,
1955, 1964; Boccitp, 1930).

The two major advantages that the electron microscope
technique offers are resolutions in the 10 A range and
replicas that permit observation of finely structured sur-
faces. Electron microscopic examination and an under-
standing of calcification in the bivalve shell is still in
preliminary phases. The body of knowledge is quite large
but only a few of the major problems have been resolved
regarding the mechanism of crystal growth and related
organic components. Significant new discoveries and con-
cepts of shell formation are still being made. An under-
standing of the genetic control of the mode and product
of calcification may furnish valuable new tools to biology
and paleontology in the fields of systematics, ecology,
phylogeny and evolution.

Electron microscopic examination of the calcified outer
surface in fossil and living bivalve shells has resulted in
significant new discoveries concerning bivalve shell struc-
ture. These potentially have an important bearing on
taxonomy and the interpretation of phylogenetic relation-
ships.

A discrete, previously unknown calcareous layer, the
mosaicostracum (Plates 26 to 38), between the ecto-

stracum and periostracum of numerous, if not all Bival-
via, and at least in some gastropods, has been defined and
studied in detail on the Tellinidae. In this family, the mo-
saicostracum is unique in that it displays patterns which
are easily recognized and are definitive for each species
(Plates 26 to 35). The most cursory examination of areas
extraneous of the mosaicostracum shows many additional
features of the shell useful in species recognition and pos-
sibly new approaches to taxonomic description based on
shell structure. In view of the large amount of data that
has been obtained, this study is restricted to the mosaic-
ostracum for the present inasmuch as it is apparently the
most useful of the valve structures for taxonomic and
phylogenetic studies. Other structures were observed in
the ectostracum and mesostracum and will be considered
in subsequent studies.

Species recognition from the shell, whether it be whole
or fragmented, presents a basic problem to the paleonto-
logist and neontologist. The main purpose of this study
is to examine the possibilities of species determination
based on electronmicroscopic shell structures and to estab-
lish control for the investigation of similar shell structures
on fossil bivalves. Preliminary investigations indicate the
preservation of the mosaicostracum in bivalves at least as
old as the Late Cretaceous (Campanian).

This study establishes a technique whereby the mala-
cologist and paleontologist engaged in the study of bi-
valves can solve problems inherent in the macromorpho-
logical approach to taxonomy with the additional infor-
mation gained by observations with the electron micro-
scope at the ultramicromorphological level. The mosaic-
ostracum and the whole study of microstructure adds a
potent new tool to the systematic and phylogenetic study
of the Bivalvia.

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Vol. 11; No. 3

With the application of modern technology to classical
problems in systematics some limit will emerge at which
the evidence for species identification is reached. Those
faunal elements which form mineralized components
have given the systematist a most useful device to decipher
the intricate code of evolution. If these mineralized com-
ponents are studied at the ultramicromorphological level,
then the limit of investigation should be the smallest set
of organized structures that consistently, at the species
level, can aid in a rigorous evaluation of patterns useful
to the systematist. The patterns found on the mosaicos-
tracum fit into this realm of evidence.

ACKNOWLEDGMENTS

The author gratefully acknowledges Dr. K. M. Towe of
the Smithsonian Institution for the many stimulating con-
versations and his liberal attitude toward the use of the
electron microscope laboratory; Dr. E. G. Kauffman of
the Smithsonian Institution for his constant encourage-
ment, criticism and constructive advice; Dr. K. J. Boss
of the Museum of Comparative Zoology, Harvard Uni-
versity for his assistance in the taxonomy of the Tellini-
dae; Dr. Joseph Rosewater of the Smithsonian Institution
for allowing the author to use specimens of the Tellini-
dae in the Division of Mollusks collection; Dr. P. E. Hare
of the Geophysical Laboratory, Washington, D. C.; Mr.
David Massie of the U.S. Geological Survey for his in-
valuable aid in many of the problems connected with
the photography; Mr. Larry B. Isham of the Smithsonian
Institution for his help in preparing the plates; Mr. J. M.
Jackson, now a graduate student at Yale University, for
suggesting the Tellinidae as a group most fitting for the
problem and for many discussions of the biology of
modern bivalves.

Explanation of Plate 26

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. xX 15000
Figure 1: Tellina (Tellina) radiata LinnaEus, 1758;
USNM No. 598232 .
Figure 2: Tellina (Moerella) salmonea (CARPENTER, 1864)
USNM No. 108639

Explanation of Plate 27

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. % 15 000
Figure 3: Tellina (Laciolina) laevigata LINNAEUS, 1758
USNM No. 83334
Figure 4: Tellina (Laciolina) magna SpENGLER, 1798
USNM No. 36174

METHODS

The mosaicostracum was detected when replicas of the
outer shell surface of Brachidontes recurvus (RAFIN-
ESQUE, 1820) were being examined in an attempt to iso-
late the area of the valve where calcite exists. Initial
observations of the outer surface revealed a type of
calcification that was not continuous with the underlying
shell structures. The nonconformity with the underlying
ectostracum and the unique appearance of this layer
incited further studies on other species of the family
Mytilidae. It appeared that a systematic study of the
layer may have implications on the problems of inter-
preting shell structure as well as a possible tool for the
recognition of shell structure differences at the species
level.

The family Tellinidae was chosen for a systematic
study of the patterns on the mosaicostracum for the
following reasons:

1. They are well represented in Recent and fossil bivalve
assemblages.

2. They have world-wide distribution. Many species have
broad geographic ranges spanning several biogeo-
graphic provinces.

3. They are euryhaline, have a depth tolerance from the
littoral zone to hundreds of feet and inhabit numer-
ous marine environments (YONGE, 1949).

4. Large, relatively complete collections are in the Smith-
sonian Institution. These had been utilized in a recent
revision of the Tellinidae by Dr. K. J. Boss, Harvard
University, and could therefore be relied upon to be
taxonomically up-to-date and in good order. Boss’s
systematics were employed in this study.

5. The species are infaunal and are commonly well pre-
served in the fossil record. The Tellinidae are there-
fore a group useful to the geologist in interpreting

Explanation of Plate 28

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. 15 000
Figure 5: Tellina (Phyllodina) squamifera DESHAYES, 1855
USNM No. 461776
Figure 6: Tellina (Phyllodina) persica DALL & SIMPSON, 1901
USNM No. 161780

Explanation of Plate 29

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. x 15000
Figure 7: Tellina (Merisca) cristallina SPENGLER, 1798
USNM No. 530395
Figure 8: Tellina (Merisca) aequistriata Say, 1824
USNM No. 194658

Tue VELIGER, Vol. 11, No. 3 [Hamitton] Plate 26

Figure 2

THE VELIcER, Vol. 11, No. 3

ne

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, Man Ys re
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- | furans

ef

ur
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. e, ~

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[Hamitton] Plate 27

THE VELIGER, Vol. 11, No. 3 [Hamitton] Plate 28

THE VELIGER, Vol. 11, No. 3 [Hamiton] Plate 29

Vol. 11; No. 3

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Page 187

the species-substrate relationships not only from the
paleoecologic point of view but also for correlation
of ancient sediments.

The specimens of Recent Tellinidae studied were ob-
tained from the United States National Museum, Division
of Mollusks collection. Fossils from USNM Department
of Paleobiology were classified according to the recent
work of Dr. K. J. Boss. The technique was basically that
of Dr. K. M. Towe with modifications (Tower & CIFELLI,
1967).

The specimens were soaked in a 5% solution of sodium
hypochlorite for 30 to 60 days to remove the periostracum
and an organic layer layer covering the mosaicostracum.
The organic layer on different species required varying
periods of time to dissolve. The smaller species were
treated whole and the larger species were cored with an
0.25 inch diamond circular saw. The cores were treated
separately from the whole shell. This technique prevented
the complete destruction of the shell which could then be
returned to the collection. The smaller species were
crushed between two pieces of lens tissue while the cores
were either broken in half or used whole. Following the
bleach treatment the specimens were rinsed and soni-
cated in de-ionized water, dried at room temperature
and shadowed with carbon and platinum simultaneously
at an angle of 45°. The coated specimens were decalcified
in Na EDTA at a pH of 7.5 and rinsed for 30 minutes in
HCl and NaClO with intermediate rinses in distilled
water. The cleaned replicas, supported on copper grids
covered with a parlodion film, were examined with an
RCA EMU 2D electron microscope. The initial magni-
fication of the micrographs was 2500 diameters which
were recorded on 210 glass lantern slide plates and
developed in Microdol-X to produce a fine-grained im-
age. The micrographs were reversed and dodged onto
Kodak Fine Grain Positive film with a Log Etronics con-
tact printer in the full dodge position and then photo-
graphically enlarged 6 diameters to give a final magni-
fication of 15 000 diameters. A comparison of the pat-
terns at magnifications of less than and more than 15 000
diameters proved unsatisfactory. Judgment as to opti-
mum enlargement was determined by a comparison of
the structures that form the patterns. Observation of the
shell surface patterns was always executed at this mag-
nification so that an objective evaluation could be made
of the information on each specimen.

Some of the smaller species could be replicated whole
so that the entire surface of the valve was examined in
the microscope without breaking up the continuity of
the patterns over the surface. The replicas made from the
cores were broken up into two or three pieces and then
examined separately. No difference could be detected be-

tween replicas made from the cores or the crushed
specimens.

DESCRIPTION
oF THE MOSAICOSTRACUM

The mosaicostracum, a calcareous veneer having an
average thickness of 0.3, is defined as a layer between
the periostracum and the ectostracum (Plate 36) and is
continuous over the entire outer surface of the valve of
bivalve mollusks. The thickness of this layer is uniform
and apparently reflects the microscopic surface of the
underlying ectostracum. This relief can be observed on
some species (Plate 29, Figure 8; Plate 35, Figure 19).
The remaining structures which can be seen only with the
electron microscope form the most important features of
the mosaicostracum and are unique to this layer. They
are of the order of 0.5 and less in size, generally. Species
of the family Tellinidae have megascopic shell surface
architectures ranging from very smooth to one of coarse
concentric ridges. However, the mosaicostracum is con-
tinuous over the most pronounced ornamentation (Plate
36). Examination of replicas made of the entire surface
of the valve confirms this situation for all of the species
studied. There are variations in the distribution of the
patterns, or the mosaicostracum may be lacking as a
result of abrasion or minor defects due to handling of the
specimens prior to this study (see Plate 38). The mosa-
icostracum may be obscured by the occurrence of ran-
dom features such as larger than average crystals or
organic material insoluble in the bleach.

From the features of the mosaicostracum it is possible
to group the patterns into major categories: crystalline,
pustular, linear, mosaic, and planar.

Crystalline: Surface displays structures with geometric
forms recognized as elements of crystal morphology
which can be coarse, medium or fine (Plate 28, Fig-
ure 6).

Pustular: Blisterlike elevations of coarse, medium or
fine pustules which may or may not have a crystalline
appearance (Plate 29, Figure 8).

Linear: Basically linear elements superimposed on a pus-
tular, crystalline or planar surface (Plate 35, Figure 19).

Mosaic: Planar surface broken by incisions or anastom-
osing sutures and grooves (Plate 30, Figure 10).

Planar: Basically flat surface with a generally smooth
appearance (Plate 32, Figure 13).

A definitive description of each category with the in-
clusion of variations resulting from a compounding of
smaller features will necessarily imply that there are

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Vol. 11; No. 3

similarities between patterns. The categories are based on
observations of the mosaicostracum at 15 000 diameters.
Differences in magnification would change the informa-
tion available for interpretation and thereby alter the
position of the pattern in the categories. These catego-
ries are tentative, proposed here only as an aid to illus-
trate the differences between patterns. Future studies will
undoubtedly expose new categories.

In order to preserve the simplicity of approach to
species recognition by observation of the patterns a de-
scription will not be made of each species pattern. They
are illustrated on Plates 26 to 35. The difference between
patterns of species in the same subgenus is apparent, for
example, in Plates 26 to 31, 33, and 35. In some rare
instances the difference between patterns of species is
subtle and an observation of greater areas of the shell
surface is necessary before the pattern for the particular
species can be recognized. Experience has shown that an
absolute minimum of 3 separate valves or valve frag-
ments of each species must be examined before confidence
of new pattern recognition is achieved. Decisions were
made on those patterns that appeared dominant or
unique. An evaluation of the percent distribution of the
pattern indicative of each species over the surface of the
valve is shown in Table 1. This evaluation was made
from examination of the entire outer surface of smaller
valves of each species. Those species with patterns ex-
hibited on less than 50% of the mosaicostracum show
other structures which are helpful in species identification
but do not fit into any recognizable systematic relation at
the present time. Future studies may prove these struc-
tures to be of greater use.

THEORY or THE ORIGIN
oF THE MOSAICOSTRACUM

The scheme for the formation of the mineralized com-
ponents in the invertebrates, especially those having car-
bonate hard parts, is not fully understood and no known
system of reactions can express even the simplest of

Explanation of Plate 30

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. % 15 000
Figure 9: Tellina (Merisca) martinicensis D’ORBIGNY, 1842
USNM No. 42897
Figure 10: Téllina (Eurytellina) angulosa GMELIN, 1791
USNM No. 33740

structures that occur. There is a general consensus that
2 phases are present in the formation of the mineral com-
ponents of the skeletal material: the mineral or inorganic
phase and the organic phase. The mineral phase, which
in the Tellinidae can be restricted to CaCOs, is formed in
conjunction with various responses of the organism
either to a time sequenced mechanism such as circadian
cell activity or cell sequences of some shorter period, or
to changing conditions of the environment not only of
the immediate area of the zone of mineralization but
also to an interaction of the organism with the gross
environment (Orton, 1928). There is the possibility
that the metabolism of the bivalve reflects the catabolism
and assimilation of food with a correlation in shell struc-
tures (Fox & Cor, 1943). The rate of growth of the valve
has been shown to be related to salinity and temperature
(MatoneE & Dopp, 1967). Probably the most significant
influence on the type of CaCO; formed is the presence
of organic material (KiTANo & Hoop, 1965). It is on
this last point that the theory of the origin of the mosa-
icostracum is based.

The role of the mantle tissue in the secretion of the
various layers of the shell has been investigated and the
secretion of the periostracum by the inner epithelium of
the outer mantle lobe has been confirmed (BEEDHAM,
1958; BEVELANDER & NAKAHARA, 1967). The formation
of the mosaicostracum occurs after secretion of the peri-
ostracum and prior to secretion of the ectostracum and
is related to calcification either on or in an organic sub-
strate forming the innermost surface of the periostracum
and the outermost surface of the ectostracum. The com-
position of this organic substrate is not known but it is
within the bounds of the present theory that it is some
combination of protein and carbohydrate. If the protein
is recognized as being the active compound and the car-
bohydrate as the passive compound a reasonable relation
between the two in terms of their role in the formation
of shell material can be visualized. The source of both
compounds is the mantle. Within each mantle cell there
is the necessary machinery to synthesize proteins and their
synthesis is dependent on the proper stimulus from the

Explanation of Plate 31

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. X 15000
Figure 11: Tellina (Eurytellina) alternata (Say, 1822)
USNM No. 40536
Figure 12: Tellina (Eurytellina) punicea Born, 1778
USNM No. 122226

Tue VELIGER, Vol. 11, No. 3 [HamiLton] Plate 30

Tue VELIcER, Vol. 11, No. 3 [HamiLTon] Plate 31

Figure 12

Vol. 11; No. 3

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Page 189

cell nucleus by the DNA. The synthesis of proteins is not
completely understood, so only a brief outline will be
given on which to base a particular aspect of the theory.

The cytoplasm of the cells making up the mantle-epi-
thelia contains ribosomes which are a combination of
RNA and protein. These ribosomes are utilized in the
synthesis of proteins when they react to messenger RNA
which is patterned after the DNA. When the proper or
specific information is transferred from the DNA by the
RNA to the ribosome, individual amino acids are assim-
ilated in some sequence to form a chain-like structure or
protein. The protein separates from the ribosome and
assumes some role in the organism. It is important to bear
in mind that the DNA contains the genetic information.
Although many types of protein are produced in the cell,
some of these (if not just one) are instrumental in the
calcification process in the shell and have certain genetic
properties reflected in their structure. We could consider
these properties as being the smallest unit of species
indication where species are based on the shell character-
istics. The protein is then secreted from the mantle and
arrives at the shell surface or, as is the case with the
mosaicostracum, at the surface of the periostracum distal
of the periostracal groove and immediately adjacent to
the ectostracum (Plate 38, Figure 31). This transport of
protein implies that the mantle is in intimate contact
with the shell and the periostracum or that there is some
mechanism that drives the proteins across the extrapallial
fluid. It is most likely a combination of both, depending
on when the secretion occurs. There is a detectable differ-
ence in electric potential across the mantle so the
mechanism could be electrophoretic (Wi1pur, 1964).
The passive or carbohydrate compounds which may
have a role in the process of calcification are derived
from mucus cells adjacent to the mantle epithelial cells.
They are found to be abundant outside the pallial line
and particularly in the outer mantle fold and secrete a
mucus-like compound best described as a mucoprotein
or neutral polysaccharide (BEEDHAM, 1958). This mucus
may play an active role in the transport of calcium or
calcium salts and is necessary for lubrication of the
mantle-shell interface. Essentially an environment con-
ducive to shell formation is created by the mantle in the
zone between the outer lobe and the periostracum. What
the source and transport mechanism of the calcium is
and how it enters into a reaction with the organic phase
is a subject of great speculation in current research. The
shell material is forming and this aspect can be inter-
preted on the basis of the interaction of the specific
amino acids making up the shell protein, accepting the
presence of the carbohydrate, and examining the patterns
of calcification. It has been demonstrated that some pro-

teins can give evidence as to their stage of evolution by
the number and sequence of amino acids in the peptide
chain (AcHER, 1966). It has been proposed that there
are active and passive sites in the protein-carbohydrate
complex and that these active sites coincide with or are
more prone to formation of the mineral components that
form the shell. Theoretically these active sites are a speci-
fic amino acid or amino acid combination such as aspar-
tic acid or glutamic acid or both (PE. Hare, personal
communication). Their position on the protein molecule
responsible for the calcifying process is determined by the
genetic nature at the species level of the material forming
the patterns observed on the outer surface of the shell or
the mosaicostracum.

The difference in size between the structures seen on
the mosaicostracum and the individual protein fibers
which contribute to the formation of the patterns is on
the order of many hundred times. The protein fibers or-
ganize into a three-dimensional network before calcifi-
cation begins in a configuration governed by the chemistry

Table 1

Estimated Percent Distribution of Mosaicostracum Showing
Distinctive Pattern

Tellina (Tellina) radiata 95%
Tellina (Moerella) salmonea 95%
Tellina (Laciolina) laevigata 50%
Tellina (Laciolina) magna 25%
Tellina (Phyllodina) squamifera 25%
Tellina (Phyllodina) persica 25%
Tellina (Merisca) cristallina 95%
Tellina (Merisca) aequistriata 50%
Tellina (Merisca) martinicensis 25%
Tellina (Eurytellina) angulosa 75%
Tellina (Eurytellina) alternata 75%
Tellina (Eurytellina) punicca 95%
Tellina (Arcopagia) fausta 25%
Tellina (Acorylus) gouldu 75%
Tellina (Scissula) similis 95%
Tellina (Scissula) iris 95%
Tellina (Angulus) versicolor 95%
Tellina (Angulus) carpenteri 75%
Tellina (Tellinella) listeri 50%
Tellina (Tellinella) virgata 25%

of the protein-polysaccharide mix. Again, this network
symmetry is determined by the genetic nature of the
calcifying proteins and polysaccharides. ‘There are prob-
ably networks which are common to many genera and
families of bivalves but the specific characteristics are
seen only in the subunit distribution which forms the

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Vol. 11; No. 3

overall pattern. These subunits are generated at the pre-
viously mentioned specific sites on the protein molecule.
The remanent of such a calcifying network is illustrated
(Plate 38, Figure 34).

DISCUSSION

The previous sections have given evidence for the presence
of a structure unique to each species of the Tellinidae
studied by a technique with the electron microscopee.
These structures form patterns which are believed to be
controlled by the genetic nature of the calcifying sub-
strate of organic material. Furthermore, evidence has
shown that the mosaicostracum is preserved in fossil ma-
terial. One specimen of Tellina (Arcopagia) gabbi Garp-
NER, 1916, from the Upper Cretaceous, Monmouth for-
mation, Brightseat, Prince Georges County, Maryland,
and one specimen of Tellina sp. from the Upper Creta-
ceous, Ripley formation, Quitman County, Georgia were
examined. Both specimens showed a well preserved mo-
saicostracum. This is the result of a durable layer of the
periostracum covering the mosaicostracum. The necessary
elements for a phylogenetic study of the mosaicostracum
have been proposed concurrent with the theory that the
portion of the periostracum covering the mosaicostracum
will prevent diagenesis to a degree that overall morpho-
logy of the patterns will not be altered in the geological
record. ‘Tests on fossil bivalves through time and space
where differences in environment of local diagenesis could
be detected as well as variation in time of the structure
of the patterns is the next step in confirming the time
sequenced substitution of different amino acids acting to
produce new and hopefully related patterns on the mosa-
icostracum. Such studies may not only elucidate the phy-
logeny of calcified tissues but also the type of calcification
that can be considered as being primitive and advanced.

Explanation of Plate 32

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. * 15 000
Figure 13: Tellina (Arcopagia) fausta PULTENEY, 1799
USNM No. 461708
Figure 14: Tellina (Acorylus) gouldii HANLEY, 1846
USNM No. 383373

Explanation of Plate 33

Plan Views from Pt/C Single Stage Replicas of the Surface of the

Mosaicostracum. 15.000
Figure 15: Tellina (Scissula) similis SowERBY, 1806
No Number

Figure 16: Tellina (Scissula) iris Say, 1822 USNM No. 193052

If these studies are made in conjunction with amino
acid analyses of the shell proteins, new evidence may be
gained to show more precisely the interrelationships of
species, genera, families, etc., of bivalves. In the event
these studies show that the mosaicostracum is subject
to some degree of alteration in the geologic record, the
usefulness of the techniques may be lost for the pale-
ontologist but it will still give the neontologist a new tool
to study the bivalves. At the outset an extensive study of
modern bivalves may be more in order before going to the
fossil record so that possible revisions of related species
and genera may be confirmed or revised. Moreover, the
phylogeny of modern bivalves may be better understood
when the interrelationship of calcified patterns and gen-
etically controlled organic calcifying substrates is fully
understood.

A more qualified inference of the space relationships
of the same species of Tellinidae is facilitated by recog-
nizing geographic sites for the collection of specimens that
best exhibit the designated pattern for the species. The
patterns illustrated were selected from specimens which
display the essential characteristics of the species based
on the categorical system proposed in the previous sec-
tion. The USNM collection numbers for the illustrated
species are to be regarded as designating the locality
where specimens of the species can be found which best
show the pattern. This is intended to designate a type
locality and population and not to designate a type
specimen.

PHYLOGENETIC IMPLICATIONS
oF THE PATTERNS

At the outset it was hoped that the patterns seen on each
different species would be so distinct that possible taxo-
nomic confusion resulting from a similarity between the

Explanation of Plate 34

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. > 15 000
Figure 17: Tellina (Angulus) versicolor (CozzENs MS) DeKay
USNM No. 462008
Figure 18: Tellina (Angulus) carpenteri DALL, 1900
USNM No. 219470

Explanation of Plate 35

Plan Views from Pt/C Single Stage Replicas of the Surface of the
Mosaicostracum. X 15000
Figure 19: Tellina (Tellinella) listeri Rovinc, 1798
USNM No. 530399
Figure 20: Tellina (Teéllinella) virgata LINNAEUS, 1758
USNM No. 17632

THE VELIGER, Vol. 11, No. 3 [HamiLTon] Plate 32

SESE TY
es KG SSS SI

Figure 14

[HamiLTon] Plate 33

THE VELIGER, Vol. 11, No. 3

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THE VELIGER, Vol. 11, No. 3 [HamiLTon] Plate 34

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THE VELIGER, Vol. 11, No. 3 [HamiLton] Plate 35

Vol. 11; No. 3

THE VELIGER

Page 191

patterns would be avoided. The study demonstrated,
however, that closely related, but not precisely similar
patterns characterized many species. In some cases, spe-
cies with very similar patterns represented the subgenera
defined on macromorphological grounds. In a few cases
species of different subgenera had similar patterns. These
features and the apparent developmental sequence of
patterns imply that the patterns are related phylogene-
tically.

The pattern groups were erected without a conscious
pre-existing scheme in mind and only after attempts to
establish the phylogenetic relationships of the patterns
did it seem plausible to propose how the patterns are
related and what the implications of such an arrangement
might be. Some of the pattern similarities agree with
Boss’s (1966) divisions at the subgeneric level and some
are distinct to such a degree that it would be a simple
matter to regroup the species into different existing sub-
genera or to identify closely related subgenera. The cri-
terion of any phylogeny is the progression from similar
characteristic to similar characteristic until the two dis-
tinct end members are separated by a progression of
interrelated features. The degree of complexity exhibited
by the mosaicostracum patterns serves as another basis
for evaluating phylogenetic relationships within the Tel-
linidae. The basis of this phylogenetic scheme will be the
degree of complexity that the patterns display. The de-
gree of pattern complexity is divided into 3 major divi-
sions: the simple group, the compound group, and the
complex group. The schematic representation is shown in
Table 2.

The simple pattern groups which are comprised of the
mosaic, pustulose, crystalline, linear, and planar patterns
will represent the basic pattern groups upon which to
erect a hierarchy. These 5 groups are divided into 3
sets. (A), which is the mosaic sct, contains the species
Tellina alternata T. angulosa, T: punicea, T. carpenter,
and T: magna. (B), which is the crystalline and pustulose
set, contains the species T: cristallina, T: martinicensis, T.
iris, T: persica, T. aequistriata, and T: similis. (C), which
is the planar and lincar sct, contains the species T- listeri
and T: virgata. (A) subset, which is the compound group
pustulose-mosaic, contains the species T: radiata and T.
versicolor. (B) subset, which is the compound group
crystalline-pustulose, contains the species T: salmonea.
(C) subset, which is the compound group crystalline-
linear, contains the species T: squamifera and T. gouldit.
(D), the last subset, is the complex group crystalline-
linear — crystalline-pustulose — pustulose-lincar, contains
one species, T: laevigata.

Table 2

COMPOUND

SIMPLE

COMPLEX

Laciolina

Eurytellina
A Set Eurytellina B
Eurytellina & A Subset
angela Angulus
Tellina

Merisca

D Subset

Scissula

Pustulose

Moerella

Scissula

Merisca

Merisca
Phyllodina

Laciolina

C Subset

Crystalline
Pustulose-Linear

Phyllodina

Acorylus
Tellinella

Crystalline-Linear |Crystalline-Pustulose | Pustulose- Mosaic

C Set
Tellinclla

Arcopagia

Starting with set (A) it may be concluded that Tellina
(Laciolina) magna, T: (Eurytellina) angulosa, T. (E.)
alternata, T. (E.) punicea, and T. (Angulus) carpenteri
are Closely related. Tellina (Angulus) versicolor represents
the transition species at the subgeneric level from the
simple mosaic group to the compound pustulose-mosaic
group and is also closely related to T: (Tellina) radiata.
In set (B), 7: (Merisca) acquistriata and T. (Scissula)
similis are closely related as are T. (S.) iris, T. (Merisca)
martinicensis, T. (Af.) cristallina and T. (Phyllodina) per-
sica. These species have no transition species at the sub-
gencric level to subset (B), which is represented by T.:
(Moercella) salmonea. Note that T: salmonea is a Pacific
species. This could imply that some other unstudied spe-
cies in the subgenus Moerella may have a transition pat-
tern or that no transition pattern exists at the subgeneric
level. Tellina (Phyllodina) persica is the transition species

Page 192

THE VELIGER

Vol. 11; No. 3

to subset (C) and is related to T: (P.) squamifera, which
is closely related to T: (Acorylus) gouldii. In set (C)
T. (Tellinella) listert is not as closely related to T. (T.) vir-
gata and T. (Arcopagia) gouldii as it is to the species in
subset (C). In set (C) there is no transition pattern at
the subgeneric level and only an allusion of the rclation-
ship to subset (C) is seen with the linear pattern on T:
(Tellinella) listeri. The species T: fausta and T. virgata
seem to be more isolated than any of the other species
in the simple pattern group. This could imply that the
planar pattern is the most primitive in terms of its posi-
tion in the phylogenetic hierarchy.

The compound group contains 5 species, each of which
may be related at the subgeneric level to 2 common
ancestors: Tellina (Phyllodina) squamifera and T. (Angu-
lus) versicolor.

Subset (D), which contains one species, Tellina (Lacio-
lina) laevigata, is the end member of the hierarchy. It
probably represents the most advanced species of the 20
studied. Boss (1966) remarks that Laciolina represent-

atives have not been discovered in the fossil record of the
Western Atlantic. It is noteworthy that most of the other
subgenera studied do have fossil representatives.

The overall validity of the phylogeny can be justified
by the number of species that share the same pattern. At
the subgeneric level there are 3 species in Eurytellina
confined to set (A), 3 species in Merisca and 2 in Scissula
confined to set (B), and 2 species in Tellinella confined to
set (C). The departure from this degree of similarity is
better born out by the interrelationships of the patterns
in the subsets.

The fact that simple patterns are combined with each
other to form the compound patterns, and that the com-
pound patterns are combined to form the complex pat-
terns leads to the question as to how this sequence of
combinations can be used to determine the phylogenetic
relationship of the species and subgenera. Three assump-
tions can be made: (1) that the simple patterns are
primary and the complex advanced phylogenetically and
therefore the complex pattern represents a recombina-

Explanation of Plate 36

Figure 21: Oblique view, single stage Pt/C stage replica from
contact of mosaicostracum with ectostracum fractured edge. Note
mosaicostracum overhanging at middle and far right. Tellina (Eu-
rytellina) angulosa. X 15000

Figure 22: Oblique view, single stage Pt/C replica of concentric
growth ridge showing continuity of mosaicostracum over ridge and
two areas where the ridge is fractured revealing the underlying
ectostracum and the mosaicostracum overhanging at the lower left
and upper right. Tellina (Eurytellina) alternata. % 5000

Figure 23: Oblique view, single stage Pt/C replica from contact of
mosaicostracum with ectostracum at fractured edge. Note angular
break of mosaicostracum lacking an overhang as in Figures 21, 22,
and 24. Tellina (Scissula) similis. 15.000

Figure 24: Oblique view, single stage Pt/C replica from contact of
mosaicostracum with ectostracum fractured edge. Note mosaicost-
racum overhanging the contact. Tellina (Eurytellina) alternata.

X 15 000

Explanation of Plate 37 Explanation of Plate 38

Figures 25, 26, 27: Ontogenetic series with range in shell size by Figure 31: Diagrammatic sketch to illustrate a typical mantle-shell

length 9mm, 13mm, 21 mm. Plan view, single stage Pt/C replica
of mosaicostracum. Tellina (Merisca) aequistriata. USNM No.
40737. % 5000

Figure 28: Plan view, single stage Pt/C replica of mosaicostracum
from Tellina (Merisca) aequistriata. USNM No. 194658.

XX 5 000

Figure 29: Plan view, single stage Pt/C replica of mosaicostracum
from Tellina (Merisca) aequistriata. USNM No. 461671.

*X 5 000

Figure 30: Plan view, single stage Pt/C replica of mosaicostracum
from Tellina (Merisca) aequistriata. USNM No. 46309.

XX 5 000 ?

relationship and probable area (A) where the mosaicostracum is
first formed. Not to scale. After BerpHamM, 1958.

Figures 32 and 33: Plan view, single stage Pt/C replica of con-
centric growth ridge (GR) with mosaicostracum to either side of
ridge and absent, probably due to abrasion, on top of ridge. Note
organic material clinging to replica in Figure 33 along lower left
edge of ridge. XX 5000

Figure 34: Plan view, sing’e stage Pt/C replica of reticulated
fibrous matrix which may be a bleach-insoluble calcifying network
between the major portion of the periostracum and the mosaicost-
racum. Observed on all species with same general appearance.
Tellina (Tellina) radiata. | 15000

THE VELIGER, Vol. 11, No. 3 [HamiLTon] Plate 36

EEE,
4 Tree

Sh J oe ; a Fe
“ a. “EN ae SS
Figure 23 Figure 24

THE VELIGER, Vol. 11, No. 3 [HamiLTon] Plate 37

Figure 27 Figure 30

THE VELIGER, Vol. 11, No. 3

periostracal

: roove
periostracum s

7,

as Je AC WM “a

44 ee we) poral

mucous

cells epithelium

of outer fold
Figure 31

Fi igure 33

[HamILTon] Plate 38

Vol. 11; No. 3 THE VELIGER Page 193

Table 3
Number in Specimens
Growth Series Examined
USNM Left Right Left Right
Subgenus and Species Number Locality Valve Valve
Tellina

(Tellina) radiata Linnazus, 1758 598232 Varadero Beach, Cuba 3 2 4 15
(Tellina) radiata LinnagEus, 1758 621563 Bermuda 1
(Tellina) radiata Linnagus, 1758 589833 Bimini, Bahamas 6
(Tellina) radiata Linnarus, 1758 360265 San Salvador 2 1
(Tellina) radiata Linnagus, 1758 27175 Cedar Keys, Florida 1 3
(Tellina) radiata Linnarus, 1758 440497 Haiti 2 1
(Tellina) radiata Linnazus, 1758 161443 Miami, Florida 2 1
(Tellina) radiata Linnarus, 1758 122222 La Guayra, Venezuela 2 1
(Tellina) radiata Linnazus, 1758 168854 Sanibel Island, Florida 2 1
(Moerella) salmonea (CARPENTER, 1864) 108634 Kiska Island, Alaska 2
(Moerella) salmonea (CarPENTER, 1864) 108649 Saint Paul, Alaska 3
(Moerella) salmonea (CARPENTER, 1864) 13688 Monterey, California 1 2
(Laciolina) laevigata Linnarus, 1758 83354 Bermuda 1 2
(Laciolina) magna SPENGLER, 1798 36174 Tampa Bay, Florida 1
(Laciolina) magna SPENGLER, 1798 57151 East Coast, Florida 1
(Laciolina) magna SpENGLER, 1798 426098 Cuba 1
(Laciolina) magna SpencuiER, 1798 621627 Bermuda 1
(Phyllodina) squamifera Desuayes, 1855 461776 Miami, Florida 2 1
(Phyllodina) persica Day « Simpson, 1901 161780 Puerto Rico 2 1
(Merisca) cristallina SPENGLER, 1798 530395 Saint Thomas, Virgin Islands 1 2
(Merisca) aequistriata Say, 1824 461671 Santa Lucia, Cuba 3
(Merisca) aequistriata Say, 1824 465309 Cape Hatteras, N. C. 1
(Merisca) aequistriata Say, 1824 194658 West Coast of Florida 4 1
(Merisca) martinicensis p’'OrBicNy, 1842 42897 Saint Dominique 2 1
(Eurytellina) angulosa Gmeuin, 1791 33740 Saint Croix 1
(Eurytellina) alternata Say, 1822 40536 Cape Hatteras, N.C. 3 4
(Eurytellina) alternata Say, 1822 35836 Cape Hatteras, N.C. 3 3
(Eurytellina) alternata Say, 1822 36173 Charleston Bay, S.C. 2 4
(Eurytellina) alternata Say, 1822 461622 Texas 1 1
(Eurytellina) alternata Say, 1822 36608 Beaufort, N.C. 1 1
(Eurytellina) alternata Say, 1822 464192 West Florida 1 1
(Eurytellina) alternata Say, 1822 46866 Florida 2 1
(Eurytellina) punicea Born, 1778 122226 La Guayra, Venezuela 3
(Arcopagia) fausta PuLTENEY, 1799 461708 Cabanas Bay, Cuba 1 2
(Arcopagia) fausta PuLTENEY, 1799 56583 Vera Cruz, Mexico 2 1
(Arcopagia) fausta PuLTENEY, 1799 53639 Key West, Florida 2 3
(Arcopagia) fausta PuLTENEY, 1799 461709 Kings, Jamaica 1 2
(Acorylus) gouldii HANLEy, 1846 500913 Barbados 3 1
(Acorylus) gouldii HANiEy, 1846 383373 Jeremie, Haiti 1
(Scissula) similis Sowersy, 1806 no No. Florida 3
(Scissula) iris Say, 1822 429015 Cape Hatteras Pt., N. C. 2
(Scissula) iris Say, 1822 193052 Cameron, Louisana 1
(Scissula) iris Say, 1822 426111 Saint Augustine, Florida 2 1
(Scissula) iris Say, 1822 128435 South Carolina 1 4
(Scissula) iris Say, 1822 461670 Bimini, Bahamas 1 1
(Angulus) versicolor (CozzEns MS) DeKay 40566 Cape Hatteras, N. C. 5 3
(Angulus) versicolor (GozzENs MS) DeKay 462123 Key West, Florida 3 2
(Angulus) versicolor (CozzEns MS) DreKay 462098 Santa Lucia, Cuba 3 2
(Angulus) carpenteri Dati, 1900 219470 Chirikoff Island, Alaska 2 1
(Tellinella) listeri ROpinc, 1798 461634 Miami, Florida 1
(Tellinella) listeri Rovinc, 1798 27238 Tortugas, Florida 1 1
(Tellinella) listeri ROpinc, 1798 426079 Havana, Cuba 1
(Tellinella) listeri ROpinc, 1798 83334 Bermuda 2 1
(Tellinella) listert Ropinc, 1798 441522 Montego Bay, Jamaica 1
(Tellinella) listeri Ropinc, 1798 393926 El Rogue Island, Venezuela 1
(Tellinella) listeri ROpinc, 1798 426323 Haiti 1 1
(Tellinella) listeri Rovinc, 1798 530399 Saint Thomas, Virgin Islands 6 4

po

(Tellinella) virgata LinnaEus, 1758 17632 Singapore, China 2

Page 194

TimEMVIELIGER

Vol. 11; No. 3

tion of genetic material from simple stocks; (2) that the
compound groups represent links between simple pattern
groups in which only one pattern mutated and introduced
a new pattern; (3) that the complex and compound
lineages are ancestral to simple lineages which represent
‘genetic isolation of the simple pattern groups contained
in the complex group.

SUMMARY

A newly discovered calcareous shell layer, the mosaicost-
racum, forms a veneer between the ectostracum and peri-
ostracum of bivalve shells and externally displays distinct
surface patterns when examined with the electron micro-
scope at magnifications of 2500 & and greater. These
patterns appear to be taxonomically consistent at the
species level and can generally be arranged in morpho-
groups, some of which are compatible with existing sub-
generic categories.

The family Tellinidae was chosen to test the constancy
of the patterns at the generic, subgeneric and species
levels under 5 parameters relevant to each tested species:
1. variation in the adult population; 2. variation between
the right and left valves; 3. ontogenetic variation; 4. ge-
ographic variation; 5. variation over the area of a single
valve with tests to compare the patterns between pig-
mented zones and unpigmented zones. Representative
species for each subgenus of the Tellinidae (Boss, 1966)
of the Western Atlantic and a few from the Pacific were
tested. Two fossil Tellins were examined from the Late
Cretaceous of Georgia and Maryland.

The species examined consistently displayed the same
patterns regardless of the tested parameters and could be
identified, on shell fragments alone, by this method. Pat-
terns of species within a single subgenus are sometimes
closely related, and the general patterns of genera and
subgenera are distinct to varying degrees, suggesting that
the ultrastructure of the mosaicostracum is a potential
tool for specific identification, and for the study of phylo-
geny and evolution in living and fossil bivalves. The prob-
able origin of this layer is discussed.

LITERATURE CITED

AcuHer, R.
1966. Evolutionary aspects of the structure of proteins.
Angewandte Chemie 5 (9): 798 - 806
BEEDHAM, G. E.

1958. Observations on the mantle of the Lamellibranchia.
Quart. Journ. Micro. Sci. 99 (2), 3° ser., no. 46: 181 - 197;
plts. 1, 2

BEvELANDER, G. « H. NAKAHARA
1967. An electron microscope study of the periostracum of
Macrocallista maculata. Calcified Tissue Res. 1 (1): 55
to 67; 6 figs.
Boccixp, G. B.
1930. The shell structure of the mollusks. D. Kg. Danske
Vidensk. Selsk. Skr., Naturv. Math. Afd., 9 R II 2: 231 - 325;
10 figs.; plts. 1-15
Boss, KENNETH JAY
1966. The subfamily Tellininae in the western Atlantic. The
genus Tellina (part 1). Johnsonia 4 (45): 217 - 272; plts.
127 - 142
CarPENTER, W.
1848. | On the microscopic structure of shells.
meeting Brit. Assoc. Adv. Sci., pp. 1 - 24; 20 plts.

Reprt. 147

Fox, Dennis LLEWELLYN & WESLEY ROSWELL COE
1943. Biology of the California sea mussel Mytilus californi-
anus. Journ. Exper. Zool. 93 (2): 205 - 249
GuNTER, GORDON
1950. The generic status of living oysters and the scientific
name of the common American species. Amer. Midl. Nat.
43 (2): 438 - 449
Kirano, Y. & D. W. Hoop
1965. The influence of organic material on the polymorphic
crystallization of calcium carbonate. Geochim. et Cosmo-
chim. 29: 29 - 41

Matong, P.G. « J. R. Dopp

1967. ‘Temperature and salinity effects on calcification rate in
Mytilus edulis and its paleoecological implications. Lim-
nol. & Oceanogr. 12 (3): 432 - 436

NeweE Lt, N. D.

1965. Classification of the Bivalvia.
2206; 3 figs.; 1 tab.

OBERLING, J. J.

Amer. Mus. Novit.

1955. Shell structure of West American Pelecypoda.
Wash. Acad. Sci. 45 (4): 128 - 130

1964. | Observations on some structural features of the pelecy-
pod shell. Mitteil. Naturforsch. Gesellsch. Bern, new ser.,
20: 1 - 63; 6 plts.; 59 figs.

Orton, Joun H.

Journ.

1928. On rhythmic periods in shell-growth in Ostrea edulis
with a note on flattening. Journ. Marine Biol. Assoc. U.

K. 15 (2): 365 - 427
Towe, K. M. « R. CIreLur
1967. Wall ultrastructure in the calcareous Foraminifera:
crystallographic aspects and a model for calcification. | Journ.
Paleont. 41 (3): 742 - 762; plts. 87 - 99

Wivzur, K. M.
1964. Shell formation and regeneration In K.M. WILBUR
« C.M.Yonce (eds.) Physiology of Mollusca; Academic
Press, New York, N. Y., 1: 243 - 282

YoncE, CHARLES MAurRICE
1949. On the structure and adaptations of the Tellinacea,
deposit-feeding Eulamellibranchia. Phil. Trans. Roy. Soc.
London, ser. B, Biol. Sci. 234 (609): 29-76

Wolyplli-sINow3

THE VELIGER

Page 195

A New Species of Strombina from the Galapagos Islands

BY

WILLIAM K. EMERSON

AND

ANTHONY D’ATTILIO

Department of Living Invertebrates,

American Museum of Natural History
Seventy-ninth Street and Central Park West, New York, New York 10024

(Plate 39)

AMONG SEVERAL NEW or otherwise interesting mollusks
received from Mrs. Jacqueline DeRoy of Academy Bay,
Santa Cruz Island, The Galapagos Islands, are two spe-
cimens of an apparently new species of Strombina. We
take pleasure in describing this species in honor of Mrs.
DeRoy.

Strombina (Cotonopsis) deroyae EMERSON & D’ATTILIO,

spec. nov.

Diagnosis: This species may be distinguished from
Strombina (Cotonopsis) esmeraldensis Otsson, 1964, a
precursor in the Ecuadorian Neogene, by having a much
larger shell (attaining nearly twice the length of the
fossil), with a less robust outline, a proportionately larger
and more strongly recurved siphonal canal, and with finer
spiral threads on the base of the body whorl.

Description: The shell is thin though large for the genus,
surface smooth, except for first 4 post-nepionic whorls;
spire extended; body whorl swollen, ending in a long
recurved canal. The holotype is a well-preserved, but
dead-collected specimen, lacking soft parts. The shell has
10 whorls, the nepionic whorls having been lost. The
individual whorls of the spire are slightly convex; the
suture is well-defined but simple; each of the 4 whorls
of the spire are sculptured with about 11 axial ribs
which extend the length of each whorl to give the suture
a slight undulation; the ribs are about equal in size to
the interspaces and gradually become obsolete on the fifth
whorl. A few widely spaced spiral striae are present on
the body whorl above the aperture; the lower half of

the body whorl is sculptured with numerous, closely set,
rounded, spiral striae that continue in the same manner
to the base of the siphonal canal. The length of the
aperture and canal is about one half the height of the
shell; the aperture is narrowly elliptical; the inner lip
sinuously turns into the canal; the parietal region is
poorly defined except below, along the siphonal canal;
anteriorly a strong cord arises from within, on the inner
lip, and curves around to join the labial edge to form
an anal canal; in addition, the labrum is thickened ante-
riorly within by a large rounded knob-like plica furthering
the formation of the anal canal; above this knob there
are 2 smaller plicae. The lip edge is thickened into an
axial cord above and diminishes below into a thin edge
along the siphonal canal; within the labrum there are
present below the knob forming the anal canal 12 to 14
lirations extending within the aperture for a short dis-
tance; the lirations are weakest anteriorly. The strongly
recurved siphonal canal is widely open.

The opercular and radular characters are not known,
as the holotype and paratype were both empty shells
when dredged.

The color on the dorsum of the shell consists of a
medium shade of brown broken up into a pattern ar-
ranged in axial, lightning-like streaks and daubs of brown
over white in a broad band on the spire; the lighter area
is directly below the suture; on the body whorl an addi-
tional lighter band is found starting anteriorly near the
anal canal and divides the pattern on the body whorl
more or less into 2 more lightly and 2 more heavily
maculated areas.

Measurements: Holotype, 49mm in length (early
whorls missing), 26.7 mm in width (Plate 39, Figures 1

Page 196

to 3). Paratype, 46 mm in length (early whorls missing),
25.4 mm in width (Plate 39, Figures 4 to 7).

Type Locality: Southeast of Academy Bay, Santa Cruz
Island, dredged in 102 fathoms, April 26, 1967, by the
DeRoys.

Type Repositories: Holotype, American Museum of
Natural History no. 146277; paratype, collection of Mrs.
DeRoy.

Remarks: Except for the recently re-discovered Carib-
bean species, Strombina pumilio (REEVE, 1859) (WEIs-
BORD, 1962; Wooprinc, 1964), representatives of this
New World genus are now confined to the warm waters
of the eastern Pacific Ocean. We here add an additional
species to the approximately 24 known from the Panamic
faunal province (see KEEN, 1958, for a list of the living
species). The genus flourished in the tropical western
Atlantic and eastern Pacific during early to late Miocene
time, ranging from the southeastern United States, the
West Indies, to Venezuela, and from Costa Rica to
Ecuador. In the Pliocene, its distribution became more
limited, with species known only from Florida, Trinidad,
Venezuela, Ecuador, and westem Panama. With the
closure of the trans-American seaways in the late Terti-
ary, only the above mentioned species is known to have
survived in the western Atlantic. WrIsBorp (1962, pp.
323 - 329) lists about 50 nominal species of Strombina
reported as fossils from deposits ranging in age from early
Miocene to late Pliocene. Of this number, only 4 of the
records refer to living species.

The new species appears to represent a living repre-
sentative of a small group of the genus that lacks a
strombinoid hump on the body whorl. This group was
afforded subgeneric recognition by Orsson (1942, p.
227), who proposed the name Cotonopsis, with Strombina
(C.) panacostaricensis OLsson (1942, pp. 227, 228; plt.
23, fig. 5) the type of the subgenus. The type species is
a fossil from the Pliocene Charco Azul formation of the
Panamanian Burica Peninsula. According to Oxsson
(1942, p. 227), a second, unnamed species of this group
also occurs in these deposits. A third species, S. (C.)
esmeraldensis OLsson (1964, p. 148; plt. 28, figs. 3, 3a),
which more closely resembles the present species, occurs
in the Mio-Pliocene Esmeraldas formation at Quebrada
Camarones, Ecuador. The Ecuadorian fossil is much

THE VELIGER

Vol. 11; No. 3

smaller than the new species, being only 25.9mm in
length, and it possesses a stouter appearance.

Of the known living species of Strombina (sensu lato)
that lack a strombinoid hump, the shell of the new Gala-
pagan species is reminiscent of Strombina (?Cotonopsis)
edentula Dat, 1908, but the resemblance may be solely
superficial. DaL’s holotype, which has not been illus-
trated, has a shorter (34 mm in length) and stouter shell
with a proportionately shorter and less recurved siphonal
canal (teste A. A. Olsson, in litteris). DaLu’s type has
each of the first 4 post-nepionic whorls ornamented with
14 to 15 axial ribs, whereas the new species has about 11
ribs on each of these whorls. Strombina turrita (SowER-
By, 1832), which is a living ally of S. edentula, also lacks
denticulations or lirations of any sort on the inside of the
outer lip. SowErBy’s taxon has a much smaller, more
slender shell than that of the new species; and unlike
the new species and S. edentula, it lacks axial ribs on the
early whorls.

ACKNOWLEDGMENTS

In addition to Mrs. DeRoy, we are indebted to Dr. George
E. Radwin of the San Diego Museum of Natural History,
Mr. Axel A. Olsson of Coral Gables, Florida, Dr. James
H. McLean of the Los Angeles Museum of Natural
History, Dr. Leo George Hertlein of the California Aca-
demy of Sciences, Dr. Joseph Rosewater of the U.S.
National Museum, and Mr. William E. Old, Jr. of the
American Museum of Natural History, for various cour-
tesies. Mr. Olsson and Dr. Radwin kindly read a draft
of the manuscript.

LITERATURE CITED

Dari, WitutiAM HEaALey

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. . XIV. The
Mollusca and Brachiopoda. Bull. Mus. Comp. Zool.,
Harvard 43 (6): 205-487; 22 plts.

KEEN, A. Myra

1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i- xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)

Explanation of Plate 39

Strombina (Cotonopsis) deroyae EMERSON & D’ArTILio, spec. nov.

Figures 1 to 3: Holotype, A.M.N.H. no. 146277;
Figures 1-2: X1.5; Figure 3: early whorls greatly enlarged

to show axial ribs

Figures 4 to 7: Paratype, DeRoy Collection; 1.5

TuHE VELIGER, Vol. 11, No. 3 [EMERSON & D’ArTILI0] Plate 39

Figure 1 Figure 2

Figure 4 Figure 5 Figure 7

Vol. 11; No. 3 THE VELIGER

Oxsson, AXEL A.
1942. ‘Tertiary and Quaternary fossils from the Burica Penin-
sula of Panama and Costa Rica. Bull. Amer. Paleo. 27 (106) :
153 - 258; plts. 14 - 25
1964. | Neogene mollusks from northwestern Ecuador. Ithaca,
New York (Paleo. Res. Inst.) pp. 1 - 256; plts. 1 - 38 (28 Oct.)

REEvE, LovELL Aucustus
1859, | Monograph of the genus Columbella. In Conchologia
Iconica, London, vol. 2; 37 plts. and text.

SoweErsBy, GEorcE BRETTINGHAM (1ST of name)
1832. | Characters of new species of Mollusca and Conchifera
collected by Mr. Cuming. Proc. Zool. Soc. London for
1832, pt. 2 (19): 113 - 120 (14 August 1832)

WEIsBorD, NorRMAN EDWARD
1962. Late Cenozoic gastropods from Northern Venezuela.
Bull. Amer. Paleo. 42 (193): 1-672; plts. 1-48
(5 March 1962)

Wooprinc, WENDELL PHILLIPS

1964. Geology and paleontology of Canal Zone and adjoining
parts of Panama. U.S. Geol. Surv. Prof. Paper 306-C:
238 - 297; plts. 39-47

Page 197

Page 198

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Vol. 11; No. 3

A Note on Feeding and Excretion in Bivalves

BY

P DINAMANI

Department of Marine Biology, University of Kerala

RECENT stTupIEs on the faecal pellets of bivalve molluscs
(Kornicker, 1962; ARAKAwA, 1963, 1965) and on their
rectum (JEGLA & GREENBERG, 1968) have drawn atten-
tion to yet another aspect of variation in the morphology
and functioning of the gut of bivalves. There is, however,
a facet of functional variation which I believe might be
relevant to these studies and which came to my notice
while I was conducting some feeding experiments on bi-
valves (Mytilus, Cardium), using different unialgal cul-
tures. Some of the main points of these observations are:

a) the form and type of faecal ribbon ejected from
the animal was found to depend upon the time ingested
material was retained within the gut;

b) this was found to vary with the type of ‘food’
material used (or available to the animal), as well as
the rate of its passage through the gut;

c) with higher concentration of food material in the
medium, strong rejectory mechanisms in the stomach
caused ingested matter to be rejected and expelled rap-
idly as faeces, at times within 20 minutes of intake;

d) faecal ribbons were initially composed wholly of
unutilized ‘food’ (in my experiments, live algal cells)
loosely bound with quantities of mucus;

e) subsequent faecal ribbons were usually of a dif
ferent texture, colour and form; and those expelled
after an interval of 12 to 20 hours after intake were
markedly different in these aspects from the first-formed
ribbons; the quantity of mucus in the faeces usually
proved to be an indicator of retention-time within the
gut: [in Mytilus for example, faecal ribbons varied in
cross-section from shapeless to oval to cylindrical mass,
with a thick or thin or a separated keel-like part; and
in colour from greenish to brownish shades;

f ) coarser material tended to be localized in faecal
ribbons and these probably marked the sites of grooves
in the gut.

Therefore, in classifying faecal ribbons on the basis of
their physical characteristics, one has to take into account
this variability which may be directly related to the
amount and type of food available to the animal. Korn-
ICKER (1962) has in fact recorded for filibranchs and
eulamellibranchs a wide range of faccal types from ‘un-

sculptured’ to ‘oval’ to ‘shapeless’ masses. ALLEN (1961)
and van WEEL (1961) have reported how faecal material
may vary in colour varying with the time within the gut.
Observations by LoosaANorF & ENGLE (1947) and Haven
& ALAMO (1966) may also be cited in this connection
since these authors have drawn attention to negative cor-
relation between faeces and total seston and to greater
faecal production at lowest levels of seston in oysters.
These reveal sources of variability in faecal production
according to the amount and type of food available to
continuous feeders such as the bivalves.

Another consideration is that though the term ‘faeces’
generally means all matter rejected from the gut, usually
the larger sense that it specifies refuse matter after
digestion is brought in by connotation. This is, for ex-
ample, implied in the observations by JEGLA & GREEN-
BERG (1968), who regard the “preparation and propul-
sion of fecal matter as an important and mandatory
function of the rectum” in the bivalves also. The time
of retention within the hindgut of ingested matter, as
well as the rate of its passage through other regions of
the gut are factors to be established for a more functional
interpretation of these structures in the bivalves. There
appears in fact to be a greater need for observations at
different times in natural populations in the field in
order to resolve many questions of bivalve nutrition.

ACKNOWLEDGMENTS

I acknowledge with pleasure the kindness of the Director
and Staff of the Marine Laboratory, Plymouth, where
this work was carried out while I was in receipt of the
Royal Society and Nuffield Foundation Commonwealth
Bursary.

LITERATURE CITED

ALLEN, J. A.

1962. Preliminary experiments on the feeding and excretion
of bivalves using P32. Journ. Mar. Biol. Assoc. U.K.
42: 609 - 623

Vol. 11; No. 3 THE VELIGER Page 199

ARAKAWA, KonHMAN Y.
1963. Studies on the molluscan faeces (1). Publ. Seto Mar.
Biol. Lab. 11: 185 - 208
1965. Studies on the molluscan faeces (II). Publ. Seto Mar.
Biol. Lab. 13: 1-21; plts. 1-6
Haven, D.S. « R. ALamo
1966. Aspects of biodeposition by oysters and other inverte-
brate filter feeders. Limnol. & Oceanogr. 11 (4:) 487 - 498
Kornicker, Louis S.

1962. Evolutionary trends among mollusk fecal pellets.
Journ. Paleontol. 36: 829 - 834
Jecia, THomas Cyrit « MiciarL, Joun GREENBERG
1968. Structure of the bivalve rectum. — I. Morphology.
The Veliger 10 (3) : 253 - 263; plts. 36-40; 1 text fig.
(1 January 1968)
Loosanorr, Victor L. « James B. ENGLE
1947. Effect of different concentrations of microorganisms on

the feeding of oysters (O. virginica). U.S. Wildlife Serv.
Fishery Bull. 51. 42: 31 - 57

vAN WEEL, P. B.

1961. The comparative physiology of digestion in molluscs.
Amer. Zoologist 1: 245 - 252

Page 200

THE VELIGER

Vol ial INoms

Volutocorbts and Fusivoluta,

Two Genera of Deepwater Volutidae from South Africa

HARALD A. REHDER

Senior Zoologist, Division of Mollusks

Smithsonian Institution, Washington, D. C. 20560

(Plates 40 to 43)

THE SUBMISSION SOME YEARS AGO by Mr. Clifton S.
Weaver of Lanikai, Hawaii, and Mrs. Helen Boswell of
Valhalla, Transvaal, South Africa, of specimens of deep-
water species of volutes from South Africa for identifica-
tion induced me to make a study of the species assigned
to the genera Volutocorbis and Fusivoluta. To these indi-
viduals and to Mr. John E. duPont, Delaware Museum
of Natural History, I give my thanks for the loan of
important material. In addition, I am grateful to Dr.
John A. Grindley, late of the South African Museum of
Natural History, and to Mary Louise Penrith, Head of
the Department of Marine Biology of that institution,
for the loan of specimens studied by Barnard and for
information regarding this material. Dr. R. Tucker Ab-
bott of the Academy of Natural Sciences, Philadelphia,
and Dr. Ruth D. Turner of the Museum of Comparative
Zoology, Harvard University, have assisted this project
by the loan of specimens.

In referring to specimens from these various institu-
tions, I have used the following abbreviations:

ANSP Academy of Natural Sciences, Philadelphia
DMNH _ Delaware Museum of Natural History
MCZ Museum of Comparative Zoology, Harvard
University
SAM _ South African Museum, Cape Town
USNM_ United States National Museum

VOLUTIDAE FLEMING, 1822

Athletinae Prtspry & Oxisson, 1954
Volutocorbis Datu, 1890
Volutocorbis Dax, 1890, p. 75; BarNarb, 1959, p. 24

This name was originally proposed by Dall as a subgenus
of Volutilithes Swarnson, 1831, a taxon of which he

considered the type to be Voluta spinosa LaMARCK
[= Voluta spinosa (LinnaEus, 1767)] of the Paris Basin
Eocene, but which is now recognized to have as type
species Voluta muricina LaAMaARCK, 1803, also of the
Paris Basin Eocene.

At that time, Dall recognized only four species as
belonging to Volutocorbis: the type species Voluta lim-
opsis Conrap, 1860, Paleocene of Alabama and Texas,
V. lima Sowersy, 1823, from the Barton Beds of the
English Upper Eocene, V. digitalina Lamarck, 1811, of
the Paris Basin Upper Eocene, and the Recent V. abyssi-
cola ADAMS & Reeve, 1848.

CossMann (1899, p. 138), in reviewing the family
Volutidae, considered Volutocorbis to be of only sectional
value, placing it under Volutilithes s. str. To Dall’s list of
species he added two species from the Paleocene: V.
muricata ForBES, 1846, from India and V. radula Fores,
1846, from India and Brazil. The first is not a Voluto-
corbis, but the Indian specimens of radula appear to
belong to a member of this group, while those from
Brazil belong elsewhere in the Volutidae.

To the Eocene species he added Volutocorbis crenuli-
fera Bayan, 1870 (V. crenulata Lamarck, 1803, not
Gein, 1791) from the Lutetian (Middle Eocene) of
the Paris Basin.

The Oligocene species that he cites as belonging here is
not a Volutocorbis but possibly a species of Athleta.

The first worker to consider Volutocorbis a distinct
genus was R. Bullen Newton in 1906, although other
workers still considered it a subgenus or section of Ath-
leta. Since 1931, however, the group has quite generally
been given generic rank.

Vol. 11; No. 3

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Page 201

The following list, undoubtedly incomplete, comprises the fossil species that appear to belong
in Volutocorbis

Table |

Group, Stage,

Age or Formation Location
Volutocorbis

limopsis (Conrap, 1860) Paleocene Midway Alabama, Texas
texana PLUMMER, 1933 Paleocene Midway Texas
kerenensis PLuMMER, 1933 Paleocene Midway Texas
daviest (VREDENBURGH, 1923) Paleocene Ranikot W. Pakistan
burtonit (VREDENBURGH, 1923) U. Paleocene _ U. Ranikot W. Pakistan
eugeniae (VREDENBURGH, 1923) U. Paleocene — U. Ranikot W. Pakistan
‘victoriae (VREDENBURGH, 1923) U. Paleocene __U. Ranikot W. Pakistan
indica DouviLLE, 1929 U. Paleocene U. Ranikot W. Pakistan
sykest (p’ArcHiaAc « Hare, 1853) U. Paleocene U. Ranikot W. Pakistan
soriensis EAMES, 1952 L. Eocene M. Laki W. Pakistan
harnaiensis Cox, 1931 L. Eocene M. Laki W. Pakistan
grauert (OPPENHEIM, 1915) L. Eocene Togo
olssoni PLUMMER, 1933 L. Eocene L. or M. Wilcox Texas
stenzeli PLUMMER, 1933 L. M. Eocene L. Claiborne Texas
crenulifera (BAyAN, 1870) M. Eocene Lutetian France
pyrenaica (ViILLALTA CoMELta, 1956) M. Eocene Spain
pakistanica Eames, 1952 M. Eocene Khirthar W. Pakistan
celebesensis (DotiFus, 1915) 4 M. Eocene ? Lutetian Celebes
darchiaci (Datton, 1908) U. Eocene Burma
digitalina (Lamarck, 1811) U. Eocene Bartonian France, England
icket (Martin, 1914) U. Eocene Nanggoelan Java

4 This species was originally assigned to the Oligocene by DoLiFus.

The single specimen undoubtedly came from nearby beds of

Eocene age (see Martin, 1917)

Certain other species originally allocated to Volutocor-
bis belong to other genera. Thus, meridionalis OLsson,
1944, from the Upper Cretaceous of Peru and oregon-
ensis TURNER, 1928, from the Eocene of Washington are
not members of Volutocorbis. Volutocorbis exornata
Petuo, 1906, from the Upper Cretaceous of Hungary is
close to Lyria. Volutospina multispinosa Newton, 1922,
from the Upper Eocene of Nigeria was placed by EAMES
(1957, p. 46) in Volutocorbis, but this species also does
not belong here. Many of the species placed by PLum-
MER (1933, pp. 381-384) in Volutocorbis belong to
Athleta.

At the present time, four living species, all occurring
in the waters off the South African Coast, have been
assigned to this genus. These are discussed below along
with the description of two new species.

Burnett SmitrH (1906) has shown effectively that
Volutocorbis limopsis Conrap of the Midway Paleocene
is the ancestral form of a phylogenetic line that evolved
into the Oligocene species that are now placed in Athlcta

(Volutispina). A similar phylogenetic evolution can un-
doubtedly be traced out among some of the European
species of this complex.

In the following systematic review of the Recent spe-
cies of the genus, I have figured two of the fossil species
to demonstrate their resemblance to some of the Recent
species (Plate 42, Figures 26 - 30).

The genus Volutocorbis is represented only in the Pale-
ocene, Eocene, and Recent, and the considerable gap in
time between the fossil species and those found living
today, amounting to some forty million years, led me to
try to find some tangible differences between the two
groups of species. Although some species such as V.
abyssicola have evolved characteristics quite divergent
from the fossil forms, other recent species resemble the
fossil species closely in shape and sculpture; for instance,
V. gilchristt SowErBY looks very much like V. digitalina
(Lamarck ). I am, therefore, constrained to retain both
Recent and fossil species in the one genus.

The geographic distribution of this genus in the Pale-

Page 202

THE VELIGER

Vol. 11; No. 3

ocene and Eocene coincides very closely with the extent
of the Tethys Sea, with the living forms now restricted to
the deeper waters off South and Southeast Africa.

The protoconch in almost all shells of Recent species
that I have seen has been so eroded that the outer shell
layer is gone, leaving what appears to be the inner calcitic
shell layer. Our single specimen of Volutocorbis gilchristi
SowErRBY does show part of the complete protoconch,
and from an examination of the latter, we see that it is
short and obtuse, consisting of 1¢ smooth glossy whorls
(Plate 40, Figure 1). The thin, outer glossy layer in this
specimen has become chipped away in places revealing
the dull calcitic layer underneath.

The protoconch of Volutocorbis limopsis (CoNRAD)
has a more slender smooth nucleus of slightly over 2
whorls, but specimens of V. digitalina (LAMARCK) from
the Bartonian Upper Eocene of England show a more
broadly conical protoconch of about 24 whorls (Plate 40,
Figure 2), resembling that found in the Recent V. dispar-
lis REHDER, new species.

Volutocorbis abyssicola (ADAMS & REEvE, 1848)
(Plate 40, Figure 3; Plate 41, Figures 10 to 13)

Voluta abyssicola ADAMS & REEvE, 1848, p. 25, pl. 7, fig. 6

Volutilithes abyssicola (ADAMS & REEVE), Woopwarp, 1900,
p. 121; Sowersy, 1902, p. 97, pl. 2. fig. 6

Volutocorbis abyssicola (ADAMS & REEVE), BARNARD, 1959, p.
25, figs. 7a, 9a

Distribution: In 85 - 300 fathoms off the South African
coast from off Hondeklipbaai, Namaqualand, western
South Africa, to the southern edge of Agulhas Bank.
Remarks: A distinctive species, reaching on occasions a
relatively large size (97 mm), it is characterized by its
elongate-ovate shape and strong reticulation of axial and
spiral sculpture, particularly on the penultimate whorl;
the body whorl occasionally shows a greater prominence
of the spiral sculpture. The eroded apical whorls suggest
a broadly conical protoconch of 2 whorls or less (Plate
40, Figure 3).

Barnarp (1959, pp. 26-27) included specimens of
Volutocorbis lutosa Kocu and V. boswellae REHDER (de-
scribed in this paper) in his concept of V. abyssicola. The
characters differentiating the latter from V. lutosa and V.
boswellae will be discussed under these species.

This species varies considerably in size and somewhat
in shape, as well as in the strength and number of the
columellar plaits. A comparison of Figures 11 and 12
shows a marked difference in breadth of shell and length
of spine. I have seen two specimens (DMNH 19306 and
Helen Boswell Collection) from 300 fathoms off Agulhas
Bank that are somewhat heavier, with a shorter broader
spine, a thickened, rather straight outer lip which is
distinctly angulate near its juncture with the body whorl
(Plate 41, Figure 13).

Volutocorbis lutosa Kocu, 1948
(Plate 40, Figure 6; Plate 41, Figures 14, 15)

Volutocorbis lutosa Kocu, 1948, pp. 5-6, pl. 2
Volutocorbis abyssicola var. lutosa Kocu, BARNARD, 1959, p. 27

Distribution: In 40 - 100 fathoms off the western coast
of South Africa, from 14 miles almost due south of Cape
of Good Hope to the mouth of the Orange River.
Remarks: I am retaining this as a species distinct from
Volutocorbis abyssicola because of its very striking char-
acters: broad-oval shape, shorter body whorl, more or
less distinctly shouldered below the suture, and fewer
and more irregular folds on the columella. This species
seems to inhabit somewhat shallower water than V.
abyssicola does, and specimens are frequently more or
less encased in a hard, reddish-brown clay. It has to
date been found only on the western side of South Africa
and off the western slope of the Agulhas Bank.

The sculpture on the penultimate and antepenultimate
whorls is more distinctly and distantly clathrate than on
the body whorl, and the axial sculpture on the last whorl,
when present in unworn specimens, consists of more or
less distant ribs diminishing in strength towards the base.

The rather slender, elevated protoconch apparently
consists of about 14 whorls (Plate 40, Figure 6).

Volutocorbis boswellae REHDER, spec. nov.

(Plate 40, Figure 5; Plate 41, Figures 16 to 19)

Diagnosis: Shell glossy, generally smaller than Voluto-
corbis abyssicola ADAMS & REEVE, with strong axial ribs,
and with the spiral sculpture restricted to the subsutural

Explanation of Plate 40

Figure 1: Volutocorbis gilchristi (SowERBY), USNM 652796

Figure 2: Volutocorbis digitalina (Lamarck), USNM 645874

Figure 3: Volutocorbis abyssicola (ADAMS & REEVE),
USNM 631849

Figure 4: Volutocorbis disparilis Reuprer, SAM A3335

Figure 5: Volutocorbis boswellae Rruper, SAM A2011
Figure 6: Volutocorbis lutosa Kocu, SAM A2012

Figure 7: Fusivoluta blaizci (BARNARD), SAM A3433
Figure 8: Fusivoluta clarket Rruprr, DMNH 12833
Figure 9: Fusivoluta barnardi Renper, DMNH 10751

Magnification: x 3.75

THE VELIGER, Vol. 11, No. 3 [REHDER] Plate 40

Vol. 11; No. 3

area and the lower half of the last whorl.

Distribution: In 80 - 300 fathoms off South Africa, from
Saldanha Bay to Cape Seal.

Description: Shell clongate-ovate in shape, glossy, light
yellowish-brown to dark orange-ycllow in color'; some
specimens, particularly immature ones, show sevcral
rows of squarish maculations of moderate reddish-orange
color. Nuclear whorls 2 - 2+, bulbous, subcylindric, with
large initial whorl. Postnuclear whorls moderately con-
vex, strongly ribbed, with a pronounced shoulder at the
suture. Ribs bluntly and broadly angulate, gradually
noticeably flattening out towards the base of the body
whorl; ribs spinosely nodulate at sutural shoulder, some-
times causing whorl to appear channeled at suture; ribs
below subsutural row of nodules are depressed followed
by a pointed nodulation and another very shallow depres-
sion; the result is that whorls in their upper part appear
to be encircled by 2 rows of rather sharp nodules sepa-
rated by a shallow channel, the upper one being stronger.
Upper half of whorl below nodulation without any spiral
sculpture, lower half of body whorl marked by broad,
shallow grooves very gradually increasing in strength an-
teriorly; in some specimens these grooves are faintly
noticeable in upper portion of body whorl. Aperture
narrow, broadly acuminate at posterior end, broad, with
a shallow sinus at anterior end. Outer lip in adult speci-
mens with an external, whitish, rather heavy callus, some-
what thickened and faintly denticulate within. Columella
with 9 to 11 spiral folds increasing in strength anteriorly;
immature specimens usually show from 4 to 9 thin folds.
A thin whitish glaze, more or less sharply delimited,
covers last whorl in columellar area.

Holotype: In 300 fathoms off Mosselbaai (Mossel Bay),
South Africa (USNM 683585).

Measurements (in mm)

height diameter locality
Holotype 59.1 25.0 off Mosselbaai
Paratype 34.35 16.5 off Cape Town
Paratype 36.35 Wed off Cape Town
Paratype ses) 15.3) off Cape Town
Paratype 34.5 15.4 off Cape Town
Paratype 36.3 17.0 off Cape Town
Paratype 28.9 15.3 off Cape Town

Eight other paratype specimens in the collection of
the South African Museum from 4 stations off San Sebas-

' Color terminology in this paper from the ISCC-NBS Mcthod
(KELLY « Jupp, 1965)

THE VELIGER

Page 203

tian Bay to Cape Seal in 85 - 90 fathoms vary in length
from 24.6 to 43.5 mm.

One paratype in the Helen Boswell collection from off

Saint Sebastian Bay in 80 - 100 fathoms measures 28.3 mm
in height and 11.2 mm in diameter (Plate 41, Figures
15, 16). Another paratype (DMNH 19308) from the
same locality measures 24.7 mm in height, and 10.9 mm
in diameter.
Remarks: I have been able to examine some of the
material that K. H. Barnard used in his discussion of this
group of species (BARNARD, 1959, pp. 26 - 27). The spe-
cimens with reduced spiral sculpture to which he refers
belong to this species and not to Volutocorbis abyssicola.
It should be pointed out that V. abyssicola and V. bos-
wellae, as well as the following species, V. disparilis, have
all been collected together at one station, A3335 in 80 to
100 fathoms.

The differences between Volutocorbis abyssicola and V.
boswellae are well marked, and in the material I have
examined I have found no intermediate forms. Not only
is the spiral sculpture greatly reduced in V. boswellae,
but the axial ribs are much fewer in number, numbering
36 in one specimen of V. boswellac while in a comparable
specimen of V. abyssicola the ribs number 60. The sub-
sutural nodes are stronger in V. boswellae, which makes
the furrow below much more pronounced. The surface in
all fresh specimens I have seen of V. abyssicola is dull,
whereas V. boswellae has a glossy surface.

Volutocorbis disparilis REHDER, spec. nov.

(Plate 40, Figure 4; Plate 41, Figures 20 to 22)

Diagnosis: Close to Volutocorbis boswellae but smaller,
more inflated, with whorls of the spire proportionately
broader, the subsutural serics of projections not as
pointed but more rounded, and the protoconch broadly
conical with about 2} whorls and a small pointed apex.
Distribution: Between Cape Town and Mosselbaai,
South Africa in 80 - 300 fathoms.

Description: Shell glossy when fresh, dark grayish-yellow
in color, rather broadly ovate. Protoconch consisting of
24 to 24 whorls, broadly trochoid, with pointed apex
and small initial whorl. Postnuclear whorls moderately
convex, regularly ribbed, sharply shouldered at the shoul-
der where the ribs are marked by an angulate laterally
compressed node; below the first series of nodes is a broad
spiral groove, followed by another series of obtuse nodes
on the ribs, and below that by another broad depression;
as in Volutocorbis boswellae the latter 2 spiral grooves
are caused by the nodes on the ribs and are not marked

Page 204

on the body of the whorl. Central part of body whorl
similarly sculptured by ribs and spiral rows of nodules,
but showing more or less distinct but narrower spiral
grooves; lower half with distinct but broader spiral
grooves, rendering ribs closely nodulose. Aperture narrow
with shallow sinus at anterior end. Outer lip in adults
with a rather thin external callus, and somewhat thick-
ened and faintly denticulate within; parietal wall with
callus on lower half, bearing 7 to 8 spiral folds, the ante-
rior one largest; immature specimens show only 3 slender
folds on anterior part of parietal wall.

Holotype: In 80 - 100 fathoms, 36°40’ S, 21°26’ E (125
miles off Saint Sebastian Bay), Station A3335, southern
slope of Agulhas Bank, South Africa (SAM A3335).
Paratypes: One specimen in South African Museum
from Station A3335; two specimens (USNM 683651)
from 300 fathoms, off Agulhas Bank; three specimens
(DMNH 10760) also from 300 fathoms, Agulhas Bank.

Measurements (in mm)

height diameter
Holotype SAM A3335 ZOE 14.0
Paratype SAM 3335 30.7 14.6
Paratype USNM 683651 36.4 16.0
Paratype USNM 683651 32.85 ont
Paratype DMNH 10760 3D 17.4
Paratype DMNH 10760 30.9 13.5
Paratype DMNH 10760 30.2 14.0

Remarks: The combination of characters that separate
this species from both Volutocorbis abyssicola and V. bos-
wellae are such that I am constrained to consider it
distinct, and because all three have been brought up in
one dredge haul and are clearly distinguishable without
any cross-over of characters I cannot but consider them
to be three specics.

THE VELIGER

Vol. 11; No. 3

Volutocorbis gilchristi (SowERByY, 1902)

(Plate 40, Figure 1; Plate 42, Figures 23, 24)

Volutilithes gilchristi Sowersy, 1902, p. 99, pl. 2, fig. 5
Volutocorbis gilchristi (SowERBY), BARNARD, 1959, p. 28

Distribution: In 160-200 fathoms off the coast of
Natal, from Durban to northern Zululand.

Remarks: This distinctive species is characterized by
its small size (25.2 to 30mm long), strong axial sculp-
ture, and deeply channeled suture. The specimen figured
(USNM 652796), taken from a specimen of Xenophora
pallidula Reeve, 1843 trawled in 160-180 fathoms off
Zululand, fortunately had the nuclear whorls fairly well
preserved.

The protoconch (Plate 40, Figure 1) is globosely low-
spired, consisting of 12 large smooth, glossy whorls; the
thin outer shell layer is broken away in places, revealing
the dull shell layer below that is seen in most specimens
of Volutocorbis. The inner edge of the outer lip is weakly
denticulate as BARNARD (1959, p. 28) mentions, bearing
a series of low, rounded, somewhat distant nodules. The
columellar folds in this specimen number seven.
Measurements: Height, 25.2mm; width,
USNM 652796.

11.8 mm.

Volutocorbis epigona (Martens, 1904)
(Plate 42, Figure 25)

Voluta epigona Martens, 1904, pp. 106 - 107, fig.
Volutocorbis cpigona (MaRTENS), BARNARD, 1959, p. 28

Distribution: In 224 fathoms off Dar es Salaam, Tan-
zania.
Remarks: The three specimens and one fragment on

which Martens based his description are apparently the
only known representatives of this species. For the sake
of completeness, I have reproduced the drawing that

Explanation of Plate 41

Figure 10: Volutocorbis abyssicola (ADAMS & Reeve), Coll. Helen

Boswell (x 1)

Figure 11: Volutocorbis abyssicola (ADAMS & ReEvE), ANSP
204528 (x 1)

Figure 12: Volutocorbis abyssicola (ADAMS & REEVE), USNM
612629 (x 1)

Figure 13: Volutocorbis abyssicola (ADAMS & REEVE), DMNH
(x 1)

Figure 14: Volutocorbis lutosa KocH, USNM 592439 (x 1)

Figure 15: Volutocorbis lutosa Kocu, USNM 612628 (x 1)

Figures 16 & 17: Volutocorbis boswellae REHDER, spec. nov.,
Holotype USNM 683585 (x 1)
Figures 18 & 19: Volutocorbis boswellae REHDER, spec. nov.
Paratype, Collection Helen Boswell (x 1.7)
Figure 20: Volutocorbis disparilis REHDER, spec. nov., Paratype

DMNH

Figure 21: Volutocorbis disparilis REHDER, spec. nov., Paratype
DMNH

Figure 22: Volutocorbis disparilis REHDER, spec. nov., Holotype
SAM A3335 (x 1.75)

Tue VELIcER, Vol. 11, No. 3 [REHDER] Plate 41

Figure 10

Figure 14

Figure 22

Figure 18 Figure 19 Figure 20 Figure 21

Vol. 11; No. 3

THE VELIGER

Page 205

accompanies the original description. This species differs
from all other living species in the strong subspinose
sculpture and the internal liration of the outer lip.

The sculpture of Volutocorbis epigona bears consider-
able resemblance to the type species V. limopsis (Conrap)
from the Paleocene of Alabama and Texas, except that
V. limopsis bears strong spiral furrows within the aperture
behind the outer lip. Volutocorbis epigona has an even
closer resemblance to V. icket Martin, 1914, of the
Upper Eocene of Java, in general shape and in the pres-
ence of internal furrows (or ridges) on the inside of
the outer lip.

Volutocorbis limopsis (Conrap, 1860)
(Plate 42, Figures 26, 30)

Volutilithes limopsis Conran, 1860, p. 292, pl. 7, fig. 24
Volutilithes (Volutocorbis) limopsis Conrap, Dat, 1890, p. 75
Volutocorbis limopsis (Conrap), Newton, 1906, p. 10

Distribution: Alabama and Texas, in the Midway group

of the Paleocene.

Remarks: ‘This species has two folds near the base of

the columella with occasionally a smaller one in between.
‘The specimens figured (USNM 137044) were collected

at Matthews Landing, Wilcox County, Alabama.

Volutocorbis digitalina (LAMARcK, 1811)
(Plate 40, Figure 2; Plate 42, Figures 27 to 29)

Buccinum scabriculum SOLANDER in BRANDER, 1766, p. 33, pl.
5, fig. 71 (not Linnarus, 1758)

Voluta digitalina Lamarck, 1811, p. 77; DEsHayEs, 1824, p.
693, pl. 93, figs. 1 - 2

Voluta lima Sowrrsy, 1823, p. 136, pl. 398, fig. 2

Distribution: France and England, in the Bartonian
Stage of the Upper Eocene.
Remarks: This species resembles in general shape Volu-
tocorbis gilchristt Sowrrsy but has strongly cancellate
sculpture and possesses three or four folds on the lower
part of the columella.

The specimens figured are from the Paris Basin (Plate
42, Figures 24, 25, USNM 644670) and from the Isle of
Wight, England (Plate 42, Figure 26, USNM 644671).

Fusivoluta von Martens, 1902
Fusivoluta von Martens, 1902, p. 237; Barnarp, 1959, p. 29

I consider this genus to comprise 7 deepwater species
which I discuss below. They occur from off South Africa

to off Kenya, East Africa. Fusivoluta elegans BARNARD
(1959, p. 32), described from off East London, South
Africa, does not appear to be a Fusivoluta and may
belong in the Fusinidae.

The type of Fusivoluta is Fusivoluta anomala (von
MarTENS), designated by M. SmitH (1942, p. 17).

Fusivoluta pyrrhostoma Watson, 1882
(Plate 42, Figure 31)

Fusus (Sipho) pyrrhostoma Watson, 1882, p. 374; WATSON,
1886, p. 208, pl. 12, fig. 2

Neptuneopsis pyrrhostoma (Watson) Sowersy, 1903, p.
226, pl. 3, fig. 1

Fusivoluta pyrrhostoma (Watson), Martens, 1904, p. 32,
pl. 3, fig. 15; BaARNarp, 1959, p. 29, fig. 7 (b, c), 9 (b, c)

Distribution: In 39-200 fathoms off the coast of
South Africa, from Cape Saint Blaize to Saldanha Bay.
Remarks: Large examples of this species were named
forma major by BARNarD (1959, p. 30). From an exam-
ination of 12 specimens of this large form, ranging in
length from 55.9 mm to 83 mm, I can see no reason for
giving these specimens a distinctive name, even of infra-
subspecific rank.

Fusivoluta capensis (THIELE, 1925)
(Plate 42, Figure 32)

Glypteuthria (?) capensis TuiELE, 1925, p. 179, pl. 19, fig. 27

Glypteuthria capensis Tomutn, 1932, p. 165, fig. 6 (not
THIELE)

Glypteuthria sculpturata Tomuin, 1945, p. 135

Fusivoluta capensis (THIELE), BARNARD, 1957, p. 210; Bar-
NARD, 1959, pp. 30 - 31

Fusivoluta capensis (THIELE), WEAVER, 1965, p. 7, figs. 5, 6

Distribution: In 250 - 560 fathoms from off Cape Point
to off Danger Point, South Africa.

Remarks: A distinct species because of its less fusiform
shape and stronger spiral sculpture.

Measurements: Height 30.4mm; diameter 13.0mm
(DMNH 10668) figured. Height 30.0mm; diameter
125mm (DMNH 10123).

Fusivoluta anomala (Martens, 1902)
(Plate 42, Figure 33)
Voluta (Fusivoluta) anomala Martens, 1902, p. 237

Fusivoluta anomala (Martens), 1904, pp. 107 - 108, pl. 3,
fig. 14

Page 206

Distribution: In 257 - 480 fathoms from north of Zan-
zibar Island to off Takaunga, Kenya.

Remarks: I have seen no specimens of this species,
which is isolated geographically from the other species of
the genus. The whorls are more convex and angulate,
and the anterior canal seems to be more strongly twisted.

Fusivoluta decussata BARNARD, 1959
(Plate 42, Figure 34)
Fusivoluta decussata BARNARD, 1959, p. 31, fig. 8c

Distribution: In 310 fathoms, 15 miles off mouth of
Buffalo River, East London, South Africa.

Remarks: This species was assigned to Fusivoluta with
some doubt by Barnard, and because of its general form
and nature of the nuclear whorls, it may provisionally
remain in this genus. I have not seen specimens of this
species.

Fusivoluta blaizei (BARNARD, 1959)
(Plate 40, Figure 7; Plate 43, Figures 35, 36)

Fulgoraria blaizei BARNARD, 1959, p. 28, fig. 8 (b)
Fusivoluta blaizei (BARNARD), WEAVER, 1963, p. 1; WEAVER,
1964, p. 2 (in part)

Distribution: In 105 - 125 fathoms on southern slope of
Agulhas Bank off Cape Saint Blaize, South Africa.

Remarks: Barnard undoubtedly placed this species in
Fulgoraria largely because of the presence of a fold at
the base of the columella. Undoubtedly, however, he
intended to compare his new species with the Japanese
species now placed in Psephaea Crosse, 1871, and not
with Fulgoraria, which has a large, bulbous protoconch,
and 8 to 12 folds on the columella. From Psephaea, how-
ever, this species also differs in the nature of protoconch,
the weakness of the columellar fold, and in the propor-
tionately shorter aperture and greater length of the spire.

THE VELIGER

Vol. 11; No. 3

Until more adult specimens are found, I follow Weaver
in assigning this species to Fusivoluta. The presence of
a weak columellar fold may be a juvenile character.

I have not seen a specimen, but through the kind
cooperation of Mr. John E. duPont, I am reproducing
photographs taken by him of the holotype in the South
African Museum.

Fusivoluta clarket REHDER, spec. nov.

(Plate 40, Figure 8; Plate 43, Figures 37 to 39)

Diagnosis: Shell rather large, solid, fusiform, of a
yellowish flesh color; protoconch of 2 whorls, first 4
whorl low, submersed, rest of protoconch with axially
oriented nodules; early postnuclear whorls more or less
angulate, with prominent axial ribs, which become obso-
lete on the fourth to sixth postnuclear whorl; postnuclear
whorls with fine, sharp, subequal, spiral cords.
Distribution: In 240-300 fathoms off Lourengo Mar-
ques District, Mozambique.

Description: Shell dull when fresh, color grayish-yellow-
ish pink, covered with a thin, adherent pale brownish-
pink periostracum; elongate-fusiform in shape. Proto-
conch of about 2 nuclear whorls, usually rather worn,
first nuclear whorl low, submersed, subsequent nuclear
whorls made angulate by axial peripheral nodules (14
in last nuclear whorl of holotype) ; first postnuclear whorl
shows commencement of spiral cords which gradually
increase in strength and number, crossing the strong axial
ribs, and becoming rather distant; spiral cords number 18
on antepenultimate whorl; on penultimate and ultimate
whorl weak threads are often present in the interspaces;
axial ribs become weak on fifth postnuclear whorl and
disappear on the penultimate and ultimate whorls. Outer
lip effuse, occasionally somewhat thickened, and with a
broad shallow sinus just above the periphery; internally
the outer lip is somewhat thickened, rarely very strongly
so, color moderate yellowish-pink in contrast to the pale
to light yellowish-pink color of the rest of aperture.

Explanation of Plate 42

Figures 23 & 24: Volutocorbis gilchristi (SoOwERBY) ,
USNM 652796 (x 2)
Figure 25: Volutocorbis epigona (MartTENS), copy of original
figure (x 1.4)
Figure 26: Volutocorbis limopsis (Conrap), USNM 137044 (x 2)
Figures 27 & 28: Volutocorbis digitalina (Lamarck),
USNM 644670 = (x 2)
Figure 29: Volutocorbis digitalina (LAMaRcK), USNM 644671
(x 2)

Figure 30: Volutocorbis limopsis (GonraD), USNM 137044 (x 2)

Figure 31: Fusivoluta pyrrhostoma (Watson), USNM 612611
(x 1)

Figure 32: Fusivoluta capensis (Turete), DMNH 10123 (x 1.7)

Figure 33: Fusivoluta anomala (MarTENS), copy of original figure
(x 1)

Figure 34: Fusivoluta decusssata BARNARD, copy of original figure

(x 3)

“re

THE VELIGER, Vol. 11, No. 3 [REHDER] Plate 42

Figure 23

Figure 29 Figure 30

Figure 31 Figure 32 Figure 33

Vol. 11; No. 3

Aperture elongate-acuminate; anterior canal straight,
rather short, moderately broad. Columella slightly sinu-
ous, without trace of folds, and with a sharply limited
yellowish-pink glaze over the parietal wall. Operculum
half the size of aperture, elongately ovate, somewhat
unguiculate; nucleus terminal, dark brown.

Specimens Examined: Holotype (DNMH 12833) and
two paratypes (MCZ 264225, USNM 686370): off Joao
Belo, District Gaza, Mozambique in 240 fathoms; two
paratypes (DMNH 19305): 80km E of Inhaca Island,
District Lourengo Marques, Mozambique; paratype
(DMNH 19307): 92km E of Inhaca Island.

Measurements (in mm)

aperture
length width length

Holotype 96.7 SHES 47.6
Paratype 98.3 B28) AOE
Paratype 80.1 28.1 40.6
Paratype 79.8 Diol 39.0
Paratype 75.6 26.9 40.2
Paratype (239) 25 ol) 38.7

locality

Off Joao Belo
Off Joao Belo *
Off Joao Belo

Remarks: ‘This species is closest in general size and shape
to Fuswvoluta barnardi Reuoer, differing markedly, how-
ever, in the stronger sculpture, different protoconch, and
in the color of the shell. Fusivoluta blaizei differs from F
clarke in the lack of spiral sculpture on the later whorls
(except for lirae on the base of the body whorl) and in
possessing a smooth bulbous protoconch of 24 whorls
(Plate 42, Figure 31).

The two paratypes from off Joao Belo, sent originally
by C.S. Weaver to Dr. Ruth Turner, contained the soft
parts, which, however, were poorly preserved. Dr. Turner
was able, nevertheless, to obtain some anatomical infor-
mation, which she has kindly permitted me to include
in this paper, as follows:

The siphon had a large left lobe but no real right
lobe. The eyes are minute, black, and situated at the
base of the broad flat tentacles. The anatomy of the
digestive tract is very close to that figured for the
Volutinac (CLENcH & Turner, 1964, p. 136, pl. 82).
The racemose salivary glands had nearly disinte-
grated, and the tubular glands were so free of them
that it is possible that their condition resembles that
in the Volutinac. It is, however, equally possible that
they were only loosely connected with the racemose
glands as in the Zidoninae, and became “easily un-
wound” (CLenci & TuRNER, 1964, p. 134) due to

THE VELIGER

92km E of Inhaca
80 km E of Inhaca
80km E of Inhaca

Page 207

the disintegration of the soft parts. Leiblein’s gland
was as in the Volutinae with an enlarged distal end.
Both specimens were males, with very large intro-
mittant organs.

The radula resembles that described and figured for
Fusivoluta pyrrhostoma (Watson, 1882) (BaRNARD,
1959, p. 30, fig. 9b, 9c) except that the basal edge is
more broadly emarginate, similar to the radula of Callio-
tectum vernicosum (Dau, 1890) (Pirspry & OLsson,
IQ ES, Jol, Bp oe, 1G).

Fusivoluta barnardi Revver, spec. nov.
(Plate 40, Figure 9; Plate 43, Figures 40 to 43)

Fulgoraria blaizei BARNARD, 1963, p. 163 (not BARNARD, 1959)

Diagnosis: Shell rather large, solid, fusiform, with a
somewhat bulbous, smooth protoconch of few whorls,
early postnuclear whorls angulate, axially ribbed, and
with fine spiral threads, which on the later whorls become
very fine and obscure, giving a smooth appearance to
penultimate and ultimate whorls. Outer lip somewhat
effuse.

Distribution: Off the Natal coast, South Africa, gener-
ally in the vicinity of the mouth of the Tugela River, in
120 - 180 fathoms.

Description: Shell rather large, 105 - 117 mm in length,
elongate fusiform, dull white in color when fresh, dead
shells orange-yellow (stained by iron oxide?). Protoconch
large, bulbous, consisting of 1 to 14 smooth whorls; the 8
postnuclear whorls sculptured as follows: first whorl with
slightly angulate periphery marked by axially elongate
nodules, which are at first crowded, but become gradually
more separated; following whorls begin to show spiral
cords, at first below the periphery, and then appearing
above and gradually becoming more numerous by inter-
calation. These fine spiral cords cross axial ribs of which
the first few are obscure and somewhat distantly sepa-
rated, but then become more pronounced, especially at
the periphery, making these whorls subangulate. At about
the beginning of the fourth whorl the axial ribs begin
diminishing in strength and the whorls become moderate-
ly and evenly convex; with the fifth whorl ribs become
very faint; on the last two whorls they have completely
disappeared, the surface being marked only by irregular
growth lines crossing the crowded, fine, unequal, spiral
threads and rendering the latter somewhat irregular; on
the penultimate whorl of the holotype these fine spiral
threads number 62. ‘The body whorl terminates in a
moderately short attenuate canal, and the outer lip flares

Page 208

THE VELIGER

Vol. 11; No. 3

out somewhat. The aperture is rather elongate, the colu-
mella straight, without folds, and the parietal wall is
usually covered by a thin glaze.

Specimens Examined:

Holotype (DMNH 10751): Off Natal coast, trawled in
120 fathoms, June 1962.

Paratypes: (USNM 686300) off North Natal to South
Zululand coast, trawled in 120 fathoms; (DMNH 10750):
off mouth Tugela River, South Zululand, in 160 - 180
fathoms, July 1960.

Measurements (in mm)

apert.

length width length?

Holotype (DMNH 10751) Moe 34.6 S28)
Paratype (USNM 686300) O34 32:3, 45.2
Paratype (DMNH 10750) WO." B90 45.9
Remarks: ‘This species is closest to Fusivoluta clarkei

but can be readily separated from it by the bulbous pro-
toconch, the finer, more numerous spiral threads, be-
coming obscure on the body whorl, the less prominent
axial ribs, and the apparently different color of fresh
shells. Fusivoluta clarkei occupies a more northerly geo-
graphic range than does F barnardi.

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Explanation of Plate 43

Figures 35 & 36: Fusivoluta blaizei (BARNARD), Holotype
SAM A3433 (x 2)

Figure 37: Fusivoluta clarkei REupER, spec. nov., Holotype
DMNH 12833 (x 1)

Figures 38 & 39: Fusivoluta clarkei Revver, spec. nov., Paratype
DMNH 19307 (x 1)

Figures 40 & 41: Fusivoluta barnardi Rewer, spec. nov., Holotype
DMNH 10751 (x 1)

Figures 42 & 43: Fusivoluta barnardi Reuper, spec. nov., Paratype

USNM 686300 (x 1)

Tue VELicER, Vol. 11, No. 3 [ReHpER] Plate 43

Figure 35 Figure 36

Figure 39

Figure 42 Figure 43

Vol. 11; No. 3

THE VELIGER

Page 209

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Part XIV. Journ. Linn. Soc. London, Zoology 16: 372-392

1886. Report on the Scaphopoda and Gastropoda collected by
H.M.S. Challenger during the years 1873 - 1876. Chall.
Reprts. Zoology, vol. 15, 756 pp.; 50 plts.

Journ. Conch.

WEAVER, CLIFTON STOKES
1963. Provisional species list of living Volutidae (Gastropoda)
8 pp. Honolulu, Hawaii

1964. Second provisional list of living Volutidae.
Honolulu, Hawaii

1965. _Volute problems.
8 text figs.

10 pp.
Hawaiian Shell News 13 (3): 7;

Woopwarpb, Martin FouNTAIN

1900. Note on the anatomy of Voluta ancilla (Sov.), Nep-
tuneopsis gilchristi Spy., and Volutilithes abyssicola (Ap. &
Rve.). Proc. Malacol. Soc. London 4: 117-125 plt. x

Page 210

THE VELIGER

Vol. 11; No. 3

List of Type Specimens of Terebridae

in the British Museum (Natural History)

BY

WALTER OLIVER CERNOHORSKY

Vatukoula, Fiji Islands

and Division of Mollusks, Smithsonian Institution, U.S. National Museum, Washington, D.C. 20560

DuRING A RECENT VIsIT to the British Museum (Natural
History), London, I have examined and recorded the
dimensions of all Terebridae types preserved in the type-
collection of the British Museum. All the types present
were photographed in colour, with the exception of 4
Carpenter types from Mazatlan, which have been dealt
with recently by Keen (1968).

It has been found in some instances that specimens
marked in the collection as types or syntypes were in fact
neither. Specimens have been selected as “types” which
do not correspond with either the original author’s stated
dimensions or figure, if such was appended; usually
larger or better preserved specimens have been selected
as types possibly by Smith, Tomlin or some other worker.
Tomuin (1944) suggested that Cuming may have sub-
stituted better specimens at a later date and subsequent
to the description of the species. My own observations
show that such specimens have indeed been added at a
later date and were glued to the same tablet as the orig-
inal type or type series.

The isolation of unequivocal types depends firstly on
the undoubted physical and possible presence of such
types in the British Museum. If Deshayes for instance
described a species from his own collection, we can ex-
pect the holotype to be in the Ecole des Mines in Paris
and not in the Cuming collection in London. Secondly,
the types present must correspond as closely as possible
with either the original description, the stated dimensions
or the original figure. Only those types which meet the
foregoing requirements have been listed as “holotypes.”
In a case where 2 or more specimens in a syntype series
agree in dimensions with a fraction of a millimeter with
the original measurements given for the “type,” an ap-
propriate lectotype has been selected. Such holotypes and
lectotypes should be interpreted purely as units of refer-

ence representative of a species and not as units of
natural populations.

Hinps (1844) stated his size in “lin.” (= lines) for
one particular specimen of a species he described. Since
British naturalists used the “inch,” “pollex” or “mm’ as
units of measure for mollusks, it is presumed that Hinps
(loc. cit.) adopted the French “ligne” of continental
authors, and popular with French malacologists at that
time, due to the restoration period re-introduction. The
French “ligne” is equivalent to 2.25 mm, and using this
conversion factor, type specimens present in the type col-
lection match the given dimensions far better than the
one-tenth of an inch conversion factor adopted by KEEN
(1966).

Some of the type specimens present in the type col-
lection have been marked with an “x” in either the aper-
ture of the shell or on the tablet next to the supposed
holotype. It should be pointed out that some specimens
have been wrongly marked as the type in a few instances.
Specimens described by E. A. Smit have a small spot of
red wax attached to the type, in some cases in addition to
an “x” mark in the aperture or on the tablet; we can
presume that these have been marked in this way by
Smith himself.

ACKNOWLEDGMENTS

I would like to express my appreciation to Dr. N. Tebble,
British Museum (Natural History), London, for having
made the collection available for examination, and for
the help and assistance extended by himself and his staff.

This work was supported (in part) by a National Re-
search Council postdoctoral Research Associateship with
the Smithsonian Institution, U. S. National Museum,
Washington, D. C.

Wole lili Nox3

THE VELIGER Page 211

LIST or SPECIES Pana (3 le mnn))ies Var bs eraciuionle4 \(—
25.4 mm). Holotype no. 74.11.10.17: 42.5 mm; var.
a: 31.0 mm; var. b: 27.6 mm.

africana, Terebra — Gray in GRIFFITH & PipcEON,

1834, Moll. Reg. Anim. Cuv., plt. 23, fig. 5. Hab. ?

Species are arranged in alphabetical order for quick ref-
erence. The following abbreviations for publications and 8.
journals have been adopted in this paper:

Coll. J. E. Gray. Size of figure: 68.0 mm. Holotype

= i f Natural
Ao NL INe IRL ate eee see Sree no. 74.10.12.12:67.4mm (marked x inside aperture) ;
LGU Beet Canine ee syntype is 61.6mm. = Terebra variegata Gray, 1834.
‘i a : Se ae Gmauisione pare 9. albida, Terebra — Gray, 1834, P.Z.S.L., p. 63. Hab. ?

M.P.M.L.P.S. = Memoirs and Proceedings of the
Manchester Literary and Philo-

Coll. J. E. Gray. Size: 13” (=44.5 mm); Var. a.

Coll. J. E. Gray. Size: 14” (=31.7 mm). Holotype
no, 74.11.10.4: 34.3 mm.

Soltteall Seorstoy 10. albocincta, Terebra (Myurella) — CARPENTER, 1857.
PM.S.L. -= Proceedings of the Malacological SECISEEN; re Hine Volts NB 228) jolt, 98) ig, 00.
Soccer Uondon = Terebra variegata Gray, 1834.
: ; ; 11. albomarginata, Terebra — DesHayes, 1859, P.Z.S.L.,
Pe Za Sli = Proceedings of the Zoological Soci- 3 : : E
ayy of loadin: p. 314. Hab. L’Australie. Coll. Cuming. Size: 44x
8mm. Holotype: 44.5 mm (marked x inside aper-
. aciculina, Terebra — Reeve, 1860, Conch. Icon., plt. urte) ; this shell is yellowish-orange in colour with
23, fig. 121 b (non LamMarcx, 1822). Hab. ? Coll. a white sutural band, obsolete axial plicae and 4
Taylor. Size of REEve’s figure: 38.0 mm. Holotype: spiral rows of deep pits on each whorl. There is
No. 74.12.11.298: 36.0 mm. another specimen in the same box measuring 39.6
. acuminata, Terebra — REEvE, 1860, Conch. Icon., plt. mm, and the loose label says “Coll. Dr. Gray” — no.
26, sp. 143 (non Borson, 1820; nec GRATELOUP, 74.10.12.14. This Gray specimen is identical in sculp-
1834). Hab. ? Coll. Cuming. Size of figure: 28.5 mm. ture with the holotype of Terebra albomarginata,
Holotype: 29.5 mm (marked x near apex of shell on except that it is uniformly dark brown in colour, and
tablet). = Hastula exacuminata Sacco, 1891. is the T: cumingit of authors, not of DrsHayes,
3. acuta, Terebra — Desuayes, 1857, J.C. P, 6: 100; 1857.
plt. 4, figs. 4, 5. Hab.: les mers de Chine. Size: 57 12. albozonata, Terebra — E. A. Smiru, 1875, A. M.N.
7mm (1857); 977mm (1859). Coll. Cuming. H., 15: 415 and 1877, A.M.N.H., 19: 226. Hab.
Supposed holotype: 46.9 mm (marked x inside aper- Matoza Harbour, Japan (Commander St. John).
ture). The dimension of 97 mm given by Deshayes Coll. J. G. Jeffreys. Size: 25><7 mm. Holotype, no.
in 1859 is certainly an error for 57 mm which was 73.8.6.9: 24.7 mm (spot of red wax on labial lip) ;
the largest specimen among the specimens examined paratype: 8.0mm.
by Deshayes. The specimen present in the collection 13. ambrosia, Terebra — Metvit, 1912, P.M. S. L., 10:
is appreciably smaller. = Terebra anilis (RODING, 250, pit. 11, fig. 10. Hab. Mekran coast, Charbar.
1798). Size: 1644 mm. Holotype no. 1912.9.17.38: 15.9
4. adamsi, Terebra — E. A. Smitu, 1873, A. M.N.H., mm; Paratype no. 1912.9.17.39: 12.4 mm. — Tereb-
11: 264. Hab. Japan (A. Apams). Coll. Cuming. ra exigua DEsHAYES, 1859.
Size: 366} mm. Holotype: 35.5 mm; paratypes: 14. amoena, Terebra — Desuayes, 1859, P. Z. S. L., p.
33.6 mm and 21.8 mm. 297. Hab. Les mers de la Chine. Coll. Cuming. Size:
5. adansoni, Terebra — Desuayes, 1859, P.Z.S.L., p. 246mm. Holotype: 24.2 mm.
291. Hab. Sénégal. Coll. Cuming and Deshayes. Size: 15. andamanica, Terebra — MeELviILL & SyKEs, 1898, P
398 mm. There are 3 syntypes in the type collec- M.S. L., 3: 41, plt. 3, fig. 3. Hab. Andaman Islands.
tion: 32.6 mm, 32.5 mm and 31.8 mm, but these are Size: 479 mm and 35 <7 mm. Only holotype pres-
appreciably smaller than the stated type, which is ent, no. 1895.4.30.2: 47.9 mm (marked x inside aper-
possibly in the Deshayes collection in the Ecole des ture). — Terebra pertusa Born, 1778.
Mines in Paris. — Terebra micans Hinps, 1844. 16. anomala, Terebra — Gray, 1834, P Z. S. Ibn p. 62.
6. addita, Terebra — Desuayes, 1859, PZ. S.L., p. 293. Hab. ? Coll. J. E. Gray. Size: 12” (= 44.5 mm).
Hab. La Terre de Van Diemen. Coll. Cuming. Size: Holotype: 42.6mm (marked x near apex of the
337 mm. Holotype: 32.3 mm. shell on tablet).
7. affinis, Terebra — Gray, 1834, P.Z.S.L., p. 60. Hab. ? 17. antarctica, Abretia — E. A. Smiru, 1873, A. M.N. lel

11: 270. Hab. Antarctic region (“New Zealand” on

Page 212

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Vol. 11; No. 3

18.

20.

Zi

22.

23.

24.

2).

26.

tablet). Coll. Justice Gillies. Size: 14><44 mm. Holo-
type no. 80.7.2.5: 13.3mm; plus 5 paratypes pres-
ent in collection. = Terebra tristis DESHAYES, 1859.
apicina, Terebra — Desuayes, 1859, P. Z. S. L., p.
284. Hab. Singapore. Coll. Cuming. Size: 225mm;
Holotype: 22.7 mm; syntypes 21.2mm and 18.8mm.

. apicitincta, Terebra — SowerBy, 1901, P. M.S. L., 4:

5, pit. 1, fig. 14. Hab. The Kowie. Size: 12.53 mm.
Holotype no. 1900.5.22.42: 12.6 mm (marked x near
apex of shell on tablet) ; paratypes no. 1900.5.22.43-
44: 12.6 and 11.8 mm, respectively.
arabella, Terebra — Tureve, 1925, Wiss. Erg. Deut.
Thee, lesqp, Walleliy,, 7/8 Sl, jak, AS), ise, De. Isley,
Sumatra. Ex coll. Berlin Museum. Three paratypes
no. 1948.12.10.5-7.
archimedis, Terebra — DESHAYES, 1859, P. Z. S. L., p.
314. Hab. ? Size: 316mm. Holotype: 32.7 mm;
syntypes: 39.6mm (this specimen is erroneously
marked as the type with an x inside the aperture)
and 20.5 mm. This is the Terebra funiculata of au-
thors, not of Hinps, 1844.
areolata, Terebra — ADAMS & REEVE, 1850, Voy. Sam-
arang, p. 30, plt. 10, fig. 23 (non Vertagus areolatus
Link, 1807). Hab. China Seas (“Fiji Islands” on
label). Size of figure: 36.0 mm. Three syntypes pres-
ent, rather questionable ones at that, of which the
largest measures 24.0 mm and is marked with x in-
side the aperture and near the apex of the shell on
the tablet. These 3 specimens are the same species as
Terebra kilburni R. D. Burcu, 1965.
argus, Terebra — Hinps, 1844, P.Z.S.L., p. 160 (nom.
nov. pro Terebra nebulosa KieNER, 1838). Hab.
Tahiti, Society Islands, Cuming; and Nukuhiva,
Marquesas, Belcher. Size: none given. Lectotype no.
1968253: 88.7 mm; paralectotypes 81.6mm and 74.1
mm.
armullata, Terebra — Hinps, 1844, P. Z.S. L., p. 154.
Hab. Between Panama and the Bay of Magdalena
in Lower California, also Galapagos. Coll. Belcher
and Cuming. Size: 22 lin. (= 49.5 mm). Holotype:
49.2 mm; syntypes: 43.3 mm and 43.1 mm.
aspera, Terebra — Hinps, 1844, P.Z.S.L., p. 154 (non
Bosc, 1801). Hab. Panama, Monte Christi, St. Ele-
na, west coast of America (‘Punta St. Elena, West
Colombia” on label). Coll. Cuming. Size: 23 lin.
(= 51.8mm). Holotype: 54.5 mm (marked x in-
side aperture) ; syntypes: 46.8 mm and 37.9 mm.
assimulis, Acus (Abretia) — Ancas, 1867, P.Z.S. L., p.
111, plt. 13, fig. 8. Hab. Dredged in Port Jackson.
Coll. Angas. Size: 54> 14 lines (= 12.4X3.4mm).
Holotype no. 70.10.26.53: 11.7 mm; syntype: 10.2
mm. = Terebra fictilis Htnps, 1844.

Pall

28.

2OF

30.

Ble

32)

33)

34.

35.

36.

Sie

38.

australis, Terebra — E. A. Smiru, 1873, A. M.N. H.,
11: 264. Hab. Swan River, and Paterson’s Bay, Tor-
res Strait, North Australia (J. R. Elsey). Size: 55
124 mm. Holotype: 54.4 mm.

bacillus, Terebra — DEsSHAYES, 1859, P. Z. S. L., p.
285. Hab. files Sandwich. Coll. Cuming. Size: 235
mm. Holotype: 22.9 mm; syntypes: 22.0mm, 19.4
mm, 17.4 mm, and 17.0 mm.

ballina, Duplicaria — HEpiey, 1915, Proc. Linn. Soc.
N.S. W, 39: 729, plt. 84, fig. 86. Hab. Trial Bay, N.
S. W. Coll. Hedley. Paratype: 19.6 mm.
bathyrhaphe, Terebra (Myurella) — E. A. Situ,
1875, A.M.N.H., 15: 415 and 1877, A. M.N.H.,
19: 226. Hab. Gulf of Yedo, Japan (Commander
St. John). Coll. J. G. Jeffreys. Size: 255mm. Holo-
type no. 73.8.6.10: 24.8mm (spot of red wax on
columella) ; syntypes: 21.7 mm, 20.0 mm, 17.0 mm,
and 10.3 mm.

belcheri, Myurella — E. A. Smitu, 1873, A. M.N. H.,
11: 267 (non Terebra belcherit Puiuipri, 1851). Hab.
Guayaquil, Ecuador. Coll. Belcher. Size: 398 mm.
Holotype no. 44.6.7.77: 38.0 mm. = Terebra guay-
aquilensis E. A. Smiru, 1880.

bernardu, Terebra — DesHayes, 1857, J. C. P, 6: 84,
pit. 4, fig. 10. Hab. Les cétes orientales de l’Australie.
(Moreton Bay on label). Coll. Cuming. Size: 58
14mm. Holotype: 57.4 mm (marked x inside aper-
ture) ; syntypes: 52.8 mm and 49.9 mm.

bicolor, Acus (Abretia) — ANGAs, 1867, P. Z. S. L., p.
111, plt. 13, fig. 7. Hab. Dredged in Middle Har-
bour, Port Jackson. Coll. Angas. Size: 82 lines
(= 1844 mm). Six syntypes: 15.0 mm, 15.0 mm,
14.3 mm, 13.4 mm, 12.8 mm, and 11.6 mm.

bifrons, Terebra — Hinps, 1844, P. Z. 8. L., p. 155.
Hab. Japan; leg. Dr. Siebold. Coll. Cuming. Size:
23 lines (= 51.8 mm). Holotype no. 1968237: 51.0
mm (marked x inside aperture). There is another
specimen present in the type collection 45.3 mm in
length, which is not a type, as Hinps (1844) men-
tioned the specimen he described to be unique.
bipartita, Terebra — DrEsHaAyeEs, 1859, P Z. S. L.,
p. 284 (non Sowersy, 1850). Hab. les Sandwich.
Coll. Cuming. Size: 225 mm. Holotype: 22.3 mm;
syntype: 19.1 mm. = Hastula albula (MENkE, 1843).
blanda, Terebra — DesHAyeEs, 1859, P. Z. S. L., p.
298. Hab. Les mers du Japon. Size: 308mm.
Holotype: 29.9 mm.

booleyi, Terebra crenulata var. — MELVILL & SYKES,
1898, PR M.S. L., 3: 44, plt. 3, fig. 5. Hab. Andaman
Islands. Size: not given. Holotype: 68.7mm. =
Terebra crenulata (LinNAEUuS, 1758).
bourguignati, Terebra — DesHAYES, 1859, P.Z. S. L.,

Wolm il INoes3,

39.

40.

Al.

42.

44.

4n3).

46.

ae

48.

ars),

p. 288. Hab. Les mers de la Chine. Coll. Cuming
and Deshayes. (“coll. Fortune” on label). Size: 19
4mm. Paratype only present, measuring 14.5 mm;
holotype probably in Ecole des Mines in Paris. =
Terebra plumbea Quoy & Gamarp, 1833.
brasiliensis, Abretia — E. A. Smiru, 1873, A. M.N.
H., 11: 271. Hab. Botafogo Bay, Rio de Janeiro
(Macgillivray, H. M.S. Rattlesnake). Size: 113
mm. Holotype no. 60.5.2.28: 10.3 mm; syntypes:
9.0 mm and 7.5mm.

brazieri, Terebra (Hastula) — Ancas, 1871, P.Z. S.L.,
p. 16, plt. 1, fig. 15. Hab. Brisbane Water, New South
Wales; Brazier. Size: 1” 2 lines 3 lines (—29.9
6.8mm). Holotype no. 71.7.5.15: 27.8mm; syn-
types: 26.2 mm, 25.8 mm, and 23.2 mm.

brevicula, Terebra — DESHAYES, 1859, P. Z.S. L., p.
296. Hab. La Terre de Van Diemen. Coll. Cuming.
Size: 37><8 mm. Two specimens present: 25.6 mm
(erroneously marked x inside aperture) and 21.6 mm,
but neither corresponds with Deshayes’ given size.
bruguicri, Terebra — DesHayes, 1859, P. Z. S. L., p.
297 (nom. nov. pro Terebra hindsii DEsHAyEs, 1857),
Hab. La Chine. Coll. Cuming. Size: 429 mm.
Holotype: 39.8mm (marked x inside aperture) ;
syntype: 37.5mm, — Terebra conspersa Hutnps,
1844.

. buccinulum, Terebra — DesHayes, 1857, J. C. P, 6:

92. pit. 5, fig. 12. Hab. lAustralie, céte orientale.
Coll. Cuming. Size: 37><10mm. Lectotype: 37.1
mm; syntypes: 36.9mm, 36.0mm, and 36.0 mm.
This is the same species as Bullia turrita Gray, 1839.
caelata, Terebra — ADAMS & REEvE, 1850, Voy. Sam-
arang, p. 30, plt. 10, fig. 22. Hab, China Seas. Size
of figure: 40.6 mm. Holotype: 41.0 mm. — Terebra
fenestrata Hinps, 1844.

caledonica, Terebra — Sowersy, 1909, PR M.S. L., 8:
198, text fig. Hab. Isle of Pines, New Caledonia.
Size: 47><11 mm. Holotype no. 1909.10.19.103: 46.8
mm.

caliginosa, Terebra — DesuHayeEs, 1859, P. Z. S. L., p.
287. Hab. Isles Philippines. Coll. Cuming. Size: 30
“6mm. Holotype: 29.7 mm.

capensis, Myurella — E. A. Smiru, 1873, A. M. N. H.,
11: 269. Hab. Port Elizabeth, Cape of Good Hope.
Size: 19><5 mm. Holotype: 17.0 mm; syntypes: 14.1
mm, 12.7 mm 11.6 mm, 9.2 mm, and 9.0 mm.

casta, Terebra — Hinps, 1844, P.Z.S. L., p. 156. Hab.
Tlo-Ilo, Island of Panay, Philippines. Size: 13 lin.
(= 29.2mm). Holotype no. 1968242: 29.0 mm;
syntypes: 38.2 mm, 33.3mm, 32.5 mm, and 30.3 mm.
= Hastula alba (MENxkE, 1843).

celidonota, Terebra — MELVILL & Sykes, 1898, P.M.
S.L., 3: 42, plt. 3, fig. 2, Hab. Andaman Islands.

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50.

58),

OAs

ie

60.

Page 213

Size: 153mm. Holotype no. 98.4.30.5: 15.0 mm;
paratype no. 98.4.30.6: 13.2 mm. = Hastula strigi-
lata (LINNAEUS, 1758).

cernohorskyi, Hastula (Punctoterebra)—R.D. Burcu,
1965, The Veliger 7: 244, plt. 31, fig. 3. Hab. Nata-
dola, Fiji Islands. Coll. W. O. Cernohorsky. Para-
type no. 1965142: 59.8 mm. Rather similar to Has-
tula laurina (Hinps, 1844).

. cinctella, Terebra — Desuayes, 1859, P. Z. S. L., p.

305. Hab. L’embouchure de l’Indus. Coll. Cuming.
Size: 296 mm. Holotype: 27.8 mm; syntypes: 31.0
mm (erroneously marked x inside aperture), and
26.9 mm.

. circinata, Terebra — Desuayes, 1857, J. C. P, 6: 99,

plt. 4, figs. 6, 7. Hab. les mers de Chine. Coll. Cu-
ming. Size: 42 <6 mm. Holotype: 41.3 mm (marked
x inside aperture). == Tercbra anilis (ROpING, 1798).
circumcincta, Terebra — DesHayeEs, 1857, J. C. P, 6:
77, pit. 3, fig. 9. Hab. la mer Rouge (Red Sea and
Port Curtis on label). Coll. Cuming (ex Stutchbury).
Size: 408 mm. Holotype: 38.0 mm (marked x in-
side aperture) ; probable syntype: 50.1 mm.
cognata, Terebra (Myurella) — E. A. Situ, 1877, A.
M. N. H., 19: 229. Hab. Persian Gulf (Colonel
Pelly). Size: 1434mm. Coll. R. MacAndrew.
Holotype: 73.7.5.7: 13.3mm (spot of red wax on
whorl above aperture and marked x near apex of
shell on tablet) ; syntypes: 6.1 mm, and 5.6 mm.

5. columellaris, Terebra — Hinps, 1844, P. Z. S. L., p.

151. Hab. ? (“Tahiti?” on label). Coll. Cuming.
Size: 19 lin. (= 42.8mm). Holotype no. 1968228:
43.0 mm; syntypes: 44.0 mm (erroneously marked x
in aperture), and 40.8 mm.

. concolor, Terebra — E. A. Smiru, 1873, A. M. N. H.,

11: 265. Hab. ? Size: 226mm. Holotype no. 1858.
5.12.252: 22.0mm. This appears to be a worn speci-
men of Duplicaria jukesu (DESHAYES, 1857).
consobrina, Terebra — Desuayes, 1857, J. C. P, 6:
72, pit. 3, fig. 3. Hab. la mer Rouge (“Massau, Red
Sea” on label). Coll. Cuming. Size: 93> 14mm
(9312mm in 1859). Holotype: 89.3 mm; _syn-
types 88.5 mm, and 84.0 mm.

. consors, Terebra — Hinps, 1844, P Z. 8. L., p. 150.

Hab. Tahiti, Socicty Islands. Coll. Cuming. Size:
31 lin. (= 69.8 mm). Holotype no. 1968173: 70.9

mm; syntypes: 53.2 mm, and 46.8 mm.

. conspersa, Terebra — Hinps, 1844, P. Z. S. L., p. 153.

Hab. Catbalonga, island of Samar, Philippines. Coll.
Cuming. Size: 10 lin. (= 22.5mm). Holotype:
23.5 mm; syntype: 19.5 mm.

continua, Terebra — Desutayes, 1859, P. Z. S. L., p.
286. Hab. ? (“Japan” on label). Coll. Cuming and
Deshayes. Size: 317mm. Holotype: 32.5 mm,

Page 214

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Vol. 11; No. 3

61.

62.

63.

64.

65.

67.

66.

68.

69.

contracta, Myurella — E. A. Smiru, 1873, A. M. N.
H., 11: 268. Hab. ? Coll. Belcher. Size: 173%
mm. Holotype: 16.00mm._.
copula, Terebra — Hinps, 1844, P Z.S.L., p. 151.
Hab. Guinea. Coll. Cuming. Size: 17 lin. (= 38.3
mm). Holotype no. 1968227: 39.4mm; syntypes:
33.8 mm, and 29.8 mm.
crenifera, Terebra — DesuayeEs, 1859, P. Z. S. L., p.
298. Hab. les mers de la Chine. Coll. Cuming. Size:
306mm. Holotype: 29.8 mm. This species is oc-
casionally synonymized with T: cingulifera LAMaARCK,
1822, but it is not that species; the locality itself is
cispect. The type of T: crenifera bears a great resem-
blance to T: ligyrus Prtspry & Lowe, 1932: the
vxial ribs are well defined and slender and form
slender nodes at the sutures; the interstices are very
broad and there are traces of an orange-brown
colour on the shell and in the interstices of the sut-
ural nodes.
cumingu, Terebra — DESHAYES, 1857, J. C. P, 6: 66,
pit. 3, fig. 1. Hab. les mers de Chine. Coll. Cuming.
Size: According to DesHayes (1857), he examined
3 examples, the largest of which measured 95x12
mm. Holotype: 93.0mm; only one syntype: 58.3 mm.
cuspidata, Terebra — Hinps, 1844, P. Z. S. L., p. 157
(non Bosc, 1801). Hab. Cape Coast, Africa. Coll.
Cuming. Size: 13 lin. (= 29.3 mm). Holotype no.
1968247: 29.4mm; syntypes: 29.9mm, and 27.7
mm. = Hastula traillu (DESHAYEsS, 1859).
delicatula, Terebra — Preston, ? Hab. Martinique
on label. Holotype no. 1911.8.22.273: 9.0 mm. I was
not able to trace the reference for this species.
decorata, Terebra — DesuHayEs, 1857, J. C. P, 6: 75,
pit. 4, fig. 13. Hab. Pidang (ile Sumatra). Capt.
Martin. Coll. Cuming. Size: 286mm. Holotype:
27.0 mm. There is another specimen in the type col-
lection, 44.0 mm in length, which was collected by
Lt. Col. Wilmer at Aden, and is not a type. (= “T.
tessellata Gray, 1834”, on label).
dillwyni, Terebra — DEsHAYES, 1859, P. Z. S. L., p.
279. Hab. les mers du Japon. Coll. Cuming and
Deshayes. Size: 409mm. Holotype: 38.9 mm;
juvenile syntype: 12.6 mm.
diversa, Terebra (Abretia) — E.A.Smiru, 1901, J.C.
L., 10: 115, plt. 1, fig. 6. Hab. Umzinto, Natal (H.
Burnup). (On label is written “Turton”, not Burnup).
Size: 275mm. Questionable syntypes: 22.5 mm,
17.6 mm, and 17.6 mm. These syntypes appear to be
the same species as Hastula parva (Bairp in BRENCH-
LEY, 1873), except that on the latter species the sut-
ural brown spots are set closer.

70.

71.

2

73:

74.

78).

76.

die

78.

9).

80.

edgaru, Terebra — Metvi1, 1898, M.P M.L. PS.,
42: 8, plt. 2, fig. 12. Hab. Karachi. Size: 19<44mm.
Holotype no. 98.7.5.32: 19.0 mm — juvenile speci-
men.
elata, Terebra — Hinps, 1844, P. Z. S. L., p. 156. Hab.
Bay of Montijo, west coast of America. Coll. Cum-
ing. Size: 12 lin. (= 27.0 mm). Holotype no. 1968-
240: 26.6 mm; syntypes: 24.9 mm, and 24.3 mm.
evoluta, Terebra — DesHayes, 1859, P Z. S. L., p.
292. Hab. Japon. Coll. Cuming. Size: 5011 mm.
Probable holotype: 56.2mm; syntypes: 29.6 mm,
22.3 mm, and 21.7 mm.
exigua, Terebra — DresHayeEs, 1859, P. Z. S. L., p.
301. Hab. La céte orientale d’Australie. Coll. Cum-
ing. Size: 1934 mm. Holotype: 20.8 mm; there is
another 14.6 mm specimen in the type collection ex
Wilmer from the Andaman Islands, which is not a
type.
fatua, Terebra — Hinps, 1844, P. Z.S. L., p. 150. Hab.
St. Christopher, West Indies (Mr. Miller, 1799).
Coll. Cuming. Size: 34 lin. (= 76.5 mm). Holotype
no. 1968175: 74.9 mm (marked x inside aperture) ;
syntype: 62.7 mm.
fenestrata, Terebra — Hinps, 1844, P Z.S. L., p. 153.
Hab. San Nicholas, Island of Zebu, Philippines.
Coll. Cuming. Size: 15 lin. (=33.7 mm). Holotype
no. 1968233: 32.7 mm; syntypes: 31.3 mm and 30.0
mm.
festiva, Terebra — Desuayes, 1857, J. C. P, 6: 74, plt.
3, fig. 4. Hab. le Sénégal. Coll. Cuming. Size: 35 <8
mm (398 mm in 1859). Holotype: 35.5 mm; syn-
types: 36.7mm (wrongly marked x in aperture),
and 32.8mm. = Terebra senegalensis LAMARCK,
1822.
fictilis, Terebra — Hinps in Sowersy, 1844, Thes.
Conch. 1: 183, plt. 45, figs. 109, 110. Hab. Australia.
Coll. Cuming. Size of figure: 144 mm. Holotype:
14.8 mm; syntypes: 15.9 mm, and 12.6 mm.
fijiensis, Myurella — E. A. Smiru, 1873, A. M.N. H.,
11: 266. Hab. Ovalau, Fiji Islands (J. McGillivray
on label). Size: 214mm. Holotype no. 1856.10.
27.8: 19.8 mm.
fimbriata, Terebra — DesHayeEs, 1857, J. C. P, 6: 71,
plt. 5, fig. 1. Hab. ? (Coll. Cuming and Deshayes in
1859). Size: 8819 mm. Paratype: 71.7 mm (holo-
type probably in Ecole des Mines in Paris). =
Terebra crenulata (LinnaEus, 1758).
flava, Terebra — Gray, 1834, P. Z.S. L., p. 60. Hab. ?
Size: 1” (= 25.4 mm). Holotype: 23.4 mm (marked
x inside aperture). = Terebra lutescens E. A. Smiru,
1873.

Vol. 11; No. 3

81.

82.

83.

84.

85.

86.

87.

88.

89.

90.

flavescens, Terebra — Desuayes, 1859, P. Z.S. L., p.
299. Hab. Les files Sandwich. Coll. Cuming (“Mr.
Pease” on label). Size: 45><9 mm. Holotype: 44.6
mm.
formosa, Terebra —- Desuayes, 1857, J. C. P, 6: 65,
plt. 3, fig. 6. Hab. la mer de Panama. Coll. Cuming.
Size: 7213 mm. Holotype: 70.9 mm.
fortunei, Terebra — Desuayes, 1857, J. C. P, 6: 79,
pit. 4, fig. 1. Hab. les mers de Chine. Coll. Cuming.
Size: 699mm. Holotype: 68.1 mm.
funiculata, Terebra — Hinps, 1844, P. Z. S. L., p. 153.
Hab. ? (“Marquesas” on label). Coll. Belcher and
Cuming. Size: 23 lin. (= 51.7 mm). Holotype: 50.3
mm (marked x inside aperture) ; syntype: 46.5 mm.
(Terebra langfordi Pitspry, 1921 is a synonym).
fuscobasis, Terebra (Myurella) — E. A. Smiru, 1877,
A. M. N. H., 19: 227. Hab. Persian Gulf (Colonel
Pelly). Coll. R. MacAndrew. Size: 1133mm.
Holotype: 11.3 mm (spot of red wax on whorl above
aperture) ; syntype: 9.2 mm.
fuscocincta, Terebra (Myurella) — E. A. Smiru, 1877,
A. M.N. H., 19: 228. Hab. Persian Gulf (Colonel
Pelly). Coll. R. MacAndrew. Size: 8><2 mm. Holo-
type no. 73.7.5.3: 7.7 mm (spot of red wax on whorl
above aperture); syntypes: 6.3mm, 5.3mm, 4.9mm,
and 3.8 mm.
geminata, Terebra — Desuayes, 1859, P. Z. S. L., p.
296. Hab. Cap Natal. Coll. Cuming. Size: 307mm.
Holotype: 30.3mm (marked x inside aperture) ;
syntype: 264mm. = Terebra spectabilis Hinps,
1844.
glabra, Terebra — Desuayes, 1857, J. C. P, 6: 101,
pit. 5, fig. 13. Hab. les mers de Chine. Coll. Des-
hayes. Size: 7013mm. The specimen present in
the Cuming Collection and measuring 43.3 mm in
length is not a type; the holotype is probably in the
Ecole des Mines in Paris. The specimen present is a
worn Terebra consors Hinps, 1844.
glauca, Terebra — Hinps, 1844, P. Z. S. L., p. 155.
Hab. ? Coll. Cuming. Unique! Size: 14 lin. (= 314
mm). Probable holotype: 27.8mm. There is another
specimen present in the type collection, measuring
18.3 mm and from the Seychelles; since Cuming’s
specimen was said to be unique, the Seychelles speci-
men is a later addition.
gotoensis, Terebra — E. A. Smitu, 1879, P. Z.S.L., p.
183, plt. 19, figs. 1, la. Hab. Goto Island, Japan.
Coll. J. G. Jeffreys. Size: 255mm (fig. 1); var-
iety: 2942 mm (fig. la). Holotype no. 78.11.7.18:
25.1 mm (spot of red wax on whorl above aper-
ture) ; paratype of variety 29.3 mm; a third syntype:
16.4mm.= Hastula melanacme (E. A. Smiru,1877).

THE VELIGER

Oe

92.

93}

94.

95.

96.

Di.

98.

99.

Page 215

gouldu, Terebra — DesHayeEs, 1857, J. C. P, 6: 89,
pl. 5, fig. 2. Hab. les iles Sandwich. Coll. Cuming
and Deshayes. Size: 70 18mm. Paratypes: 61.6 mm
(wrongly marked x inside aperture), and 59.2 mm.
Holotype is probably in the Ecole des Mines in Paris.
gracilis, Terebra — Retve, 1860, Conch. Icon., plt. 24,
spec. 131 (non Lea, 1833; nec Gray, 1834). Hab.
Africa. Size of figure: 21.6 mm. Holotype no. 74.10
29.2: 22.0mm. (“Coll. J. E. Gray” on loose label).
According to SmirH (1877), T. gracilis REEVE is not
the T. gracilis of Gray; he points out that T: gracilis
Gray was mounted on the same tablet as T. gracilis
REEVE. This would account for the “Coll. J. E. Gray”
on the loose label. On the reverse side of the tablet
on which the specimen of T: gracilis REEVE is present-
ly mounted, is the following note: “this specimen is
figured by Reeve as the type of gracilis. For the true
type see another tablet.” No such tablet with Gray’s
T. gracilis could be traced. REEvE’s T. gracilis is a
small specimen of T: spectabilis Hinps, 1844, with
fewer ribs per whorl.
granulosa, Myurella — E. A. Smitu, 1873, A. M. N.
H., 11: 268 (non Terebra granulosa Lamarck, 1822).
Hab. Japan (A. Adams). Coll. Cuming. Size: 266
mm. Holotype: 24.6 mm (marked x next to specimen
on tablet) ; syntypes: 21.8 mm, and 18.2 mm.
= Terebra pustulosa E. A. Smiru, 1879.
gray, Terebra — E. A. Smiru, 1877, A. M. N. H.,
19: 227 (nom. nov. pro Terebra gracilis Gray, 1834).
No types present in the type collection.
guayaquilensis, Terebra — E. A. Smiru, 1880, P. Z. S.
L., p. 481 (nom. nov. pro Myurella belcheri E. A.
SmitH, 1873 — non Terebra belchert Puiuprt, 1851).
For types see under Myurella belcheri E. A. SmitH;
this appears to be an earlier name for Terebra monti-
joensis Pitspry « Lowe, 1932.
helichrysum, Terebra — MELvILL & STANDEN, 1903,
A. M. N. H., 12: 310, plt. 22, fig. 14. Hab. Persian
Gulf, Mussandan. Size: 245mm. Holotype no.
1903.12.15.117: 244mm; paratype no. 1903.12.15.
118: 15.3 mm.
hindsii, Terebra (Myurella) — Carpenter, 1857 —
see KEEN, 1968, p. 428, plt. 58, figs. 71a-b.
bra variegata Gray, 1834.
hindsi, Terebra — DESHAYES, 1857, J. C. P, 6: 81, plt.
5, fig. 5 (non CarPENTER, 1857). Hab. les mers de
Chine. Coll. Cuming. Size: 429 mm. Only one spe-
cimen present in the type collection which measures
27.6 mm; it is a worn T. conspersa Hinps, 1844.
incolor, Terebra — Desuayes, 1859, P. Z. S. L., p.
283. Hab. fles Philippines. Coll. Cuming. Size: 34>
8mm. Holotype: 33.4mm. =Hastula alba (MEN-
KE, 1834).

——Wene=

Page 216

100.

101.

102.

103.

104.

105.

106.

107.

108.

109.

110.

incomparabilis, Terebra — DesuayeEs, 1859, P. Z.S.
L., p. 307. Hab. Panama. Coll. Cuming. Size: 85
13 mm. Holotype: 84.6 mm; there is a slightly shoul-
dered ridge in the centre of each whorl and the 2
spiral rows of spots are equal sized. = Terebra
lingualis Hinps, 1844.

inconstans, Terebra — Hinps, 1844, P. Z. S. L., p.
156. Hab. Sandwich Islands. Coll. Cuming. Size:
16 lin. (= 36.0 mm). Lectotype no. 1968243: 33.2
mm; syntypes: 33.1 mm, and 32.6 mm.

insalli, Terebra (Triplostephanus) — BRATCHER &
Burcu, 1967, The Veliger, 10: 7, plt. 2, figs. 1-3.
Gulf of Akabar (sic). Paratype: 52.4 7.9 mm.
insignis, Terebra — DESHAYES, 1857, J. C. P, 6: 70,
plt. 3, fig. 2. Hab. Panama. Size: 7815 mm. Holo-
type: 77.0mm; another specimen, possibly added
later, 118.7 mm. = T: lingualis Hinps, 1844.
interlineata, Terebra — DESHAYES, 1859, P. Z. S. L.,
p. 277. Hab. Les files Sandwich. Coll. Cuming. Size:
60.0 mm. Holotype: 61.2 mm, = juvenile specimen
of Terebra crenulata (LinNaAEus, 1758).
intertincta, Terebra — Hinps, 1844, P. Z. S. L., p.
155. Hab. Gambia. Coll. Cuming and Saul. Size:
20 lin. (= 45.0 mm). Holotype no. 1968235: 44.1
mm; syntypes: 42.7 mm, and 40.4 mm (this specimen
is erroneously marked x inside aperture). The local-
ity indication is obviously erroneous; the species re-
sembles in form, colour and sculpture the species T:
armillata Hinps, 1844. The figure of T: intertincta
in Keen (1958) is dissimilar to the type, which has
vertically concave whorls.

japonica, Terebra — E. A. Smitu, 1873, A. M.N. H.,
11: 265. Hab. Japan (A. Apams). Size: 327 mm.
Holotype: 31.4mm (marked x inside aperture) ;
syntypes: 29.3 mm, and 26. 8mm.

jeffreysu, Terebra — E. A. Situ, 1879, P. Z. S.L., p.
184, plt. 19, fig. 2. Hab. Station 20 & 21, Japan.
Size: 255 mm. Holotype no. 78.10.16.4: 25.0 mm
(spot of red was on body whorl); syntypes: 22.5 mm,
21.0mm, and 13.2 mm.

jenningsi, Terebra (Triplostephanus) — R. D. Burcu,
1965, The Veliger, 7: 248, plt. 31, fig. 9. Hab. Namo-
tu Island, Fiji Islands. Coll. W. Cernohorsky. Para-
type no. 19655143: 47.8 mm.

jukesu, Terebra — DesuayeEs, 1857, J. C. P, 6: 95,
pit. 6, fig. 9. Hab. Port Essington. Coll. Cuming.
Size: 338 mm. Holotype: 32.8 mm; syntype: 34.2
mm (erroneously marked x inside aperture).
kienert, Terebra — Desuayes, 1859, P. Z. S. L., pp.
292, 294. Hab. La Terre de Van Diemen. (‘“‘Tas-
mania” on label). Coll. Cuming and Paris Museum.

THE VELIGER

I.

2,

113.

116,

13),

116.

il

118.

118),

120.

Walle

122.

Vol. 11; No. 3

Size: 226 mm. Holotype: 21.2 mm; syntype: 19.1
mm.

kilburni, Terebra (Decorthastula)- R. D. Burcu,
1965, The Veliger, 7: 249, plt. 31, fig. 8. Hab. Off
Wading Island, Fiji Islands. Coll. W. Cernohorsky.
Paratype no. 1965141: 16.3 mm.

knockert, Terebra — E. A. Smiru, 1871, P. Z. S. L.,
p. 730, plt. 75, fig. 7. Hab. Whydah [West Africa].
Size: 204 mm. Lectotype no. 70.1.3.47: 18.9 mm;
syntypes: 18.6mm, 14.9mm, 14.5mm, and 13.8mm.
= Hastula modesta (DEsHAYEs, 1859).

knorri, Terebra — Gray, 1834, P. Z. S. L., p. 59.
Hab. ? Size:25 <4” (= 634 12.7 mm). Holotype:
62.7mm. = T. chlorata LaMarck, 1822.

lactea, Terebra — DesHayes, 1859, P. Z. S. L., p.
285. Hab. Isles Sandwich (“Vasigapatam, Dr. Traill”
on label). Coll. Cuming and Deshayes. Size: 214
mm. Probable holotype: 19.6mm; syntypes: 18.2
mm, and 16.0mm. =Hastula bacillus (DESHAYES,
1859).

laevigata, Terebra — Gray, 1834, P. Z. S. L., p. 61.
Hab. ? Size: 13” (= 31.7 mm). Holotype no. 74.11.
10.9: 28.0 mm; syntype: 21.7 mm.

laevis, Terebra — Gray, 1834, P. Z. S. L., p. 61. Hab.
? Size: 14” (= 31.7mm). Probable holotype no.
74.11.10.10: 34.3 mm. == T. cingulifera LAMarcK,
1822.

larvae formis, Terebra — Hinps, 1844, P. Z. S. L., p.
155. Hab. St. Elena, Monte Christi, west coast of
America (“Punta St. Elena, West Colombia” on
label). Coll. Cuming. Size: 23 lin. (= 51.7 mm).
Holotype no. 1968239: 50.7 mm; syntypes: 37.3 mm,
and 31.7 mm.

laurina, Terebra — Hinps, 1844, P. Z. S. L., p. 152.
Hab. Western Africa. Coll. Cuming. Size: 32 lin.
(= 72.0mm). Holotype no. 1968229: 67.6mm;
syntype: 63.0 mm.

lepida, Terebra — Hinps, 1844, P. Z. S. L., p. 158.
Hab. Guinea. Coll. Cuming. Size: 10 lin. (= 224 mm).
Holotype: 23.0mm; syntypes: 244mm, and 21.9
mm. = Hastula strigilata (LinnaEus, 1758).
ligata, Terebra — Hinps, 1844, P. Z. S. L., p. 153.
Hab. Marquesas. Coll. E. Belcher. Size: 154 lin.
(= 34.9mm). Probable holotype: 31.1mm (juv.
specimen).

lightfooti, Terebra (Abretia) — E. A. Smiru, 1899,
J Geo 24 7p plts op atie- alae labaalablem ban,
(Lightfoot). Coll. J. H. Ponsonby. Size: 194 <6 mm.
Holotype no. 99.9.9.2: 19.5 mm.

ligneola, Terebra — REEve, 1860, Conch. Icon., plt.
7, spec. 25 (nom. nov. pro T: castanea Hinps, 1844

Vol. 11; No. 3

THE VELIGER

Page 217

123.

He

125.

126.

Di

128.

129.

130.

UBil.

132.

—non Krener, 1838). Hab. China; Fortune. Coll.
Cuming. Size of figure: 63.2 mm. Holotype: 63.0
mm; syntypes: 62.4 mm, and 57.2mm. = T: badia
Desuayes, 1859.

lima, Terebra -DEsuayes, 1857, J. C. P, 6: 69, plt.
4, fig. 2. Hab. les mers de Chine. Size: 7811 mm.
Holotype: 764 mm.

lingualis, Terebra — Hinps, 1844, P. Z. S. L., p. 153.
Hab. Gulf of Papagayo, Bay of Montejo, west coast
of America. Coll. Cuming. Size: 30 lin. (= 674mm).
Holotype no.1968232: 68.1 mm; syntypes: 73.3 mm,
and 57.2 mm. The specimen figured by CAMPBELL
(1962) as the “type” of T: lingualis was the 73.3
mm syntype.

livida, Terebra — REEVE, 1860, Conch. Icon., plt. 22,
spec. 116. Hab. Philippine Islands. Coll. Taylor. Size
of figure: 41.8mm. Holotype no. 74.12.11.299:
38.4 mm.

loisae, Terebra — E. A. SmituH, 1903, P. M. S. L., 5:
360, plt. 15, fig. 1. Hab. Umkomaas, Natal (Burnup).
Coll. J. H. Ponsonby. Size: 306mm. Holotype
no. 1901.9.22.81: 28.4mm; there are another 2
specimens present in the type collection which are
not type material.

longiscata, Terebra — DesuHAYES, 1859, P. Z. S. L., p.
294. Hab. Les Isles Philippines. Coll. Cuming. Size:
299 mm. There is one specimen in the type collec-
tion which is most probably the holotype and meas-
ures 24.4><4.8 mm. Deshayes’ stated dimensions are
obviously an error, since the given width in relation
to the total length is out of proportion for such a
slender species of Terebra.

luctuosa, Terebra — Hinps, 1844, P. Z. S. L., p. 157.
Hab. Gulf of Nicoya, Puerto Portrero, west coast of
America. Coll. Cuming and Belcher. Size: 17 lin.
(= 38.3 mm). Holotype no. 1968246: 38.4 mm; syn-
types: 40.6mm, and 33.0mm. The types are not
smooth, but have fine slender axial ribs.

lutescens, Terebra — E. A. Smiru, 1873, A. M.N. H.,
11: 264 (nom. nov. pro T: flava Gray, 1834). For
types see under Terebra flava Gray, 1834.
macandrewi, Terebra (Myurella) — E. A. Smitu,
1877, A. M. N. H.,19: 228. Hab. Persian Gulf (Colo-
nel Pelly). Coll. R. MacAndrew. Size: 1344 mm.
Holotype no. 73.7.5.5: 13.4mm (spot of red wax
on whorl above aperture) ; syntype: 11.2 mm.
macgillivrayi, Myurella — E. A. Smiru, 1873, A. M.
N. H., 11: 267. Hab. Brunei Island, south coast of
New Guinea (J. MacGillivray, H. M. S. Rattle-
snake). Size: 225mm. Holotype no. 56.12.8.93:
21.8 mm.

mamillata, Terebra (Myurella) — Watson, 1886,

133.

134.

135%

136.

IS

138.

140.

141.

Voy. Challenger, Zool., 15: 381, plt. 16, fig. 1. Hab.
Station 204A or B, Philippines. Size: 1.35><0.25
inches (= 34.3 mm). Holotype no. 87.2.9.1159: 33.7
mm; paratype no. 87.2.9.1160: 29.6 mm,
marginata, Terebra — Desuayes. 1857, J. C. P, 6:
86, plt. 4, fig. 8. Hab. L’embouchure de la Gambie.
Coll. Cuming. Size: 348 mm. Holotype: 34.0 mm
(marked x inside aperture) ; syntype: 29.6 mm. The
locality indication of Gambia appears to be errone-
ous, since the types are coarsely sculptured T. armil-
lata Hinps, 1844.

mariesi, Terebra — E.. A. Situ, 1880, P. Z. S. L., p.
480, plt. 48, fig. 5. Hab. Japan (Mr. Maries). Size:
457 mm. Holotype no. 1880.3.1.94: 43.0 mm (spot
of red wax on whorl above aperture) ; syntype: 40.6
mm.

marmorata, Terebra — DesHayes, 1859, P. Z. S. L.,
p. 279. Hab. Moreton Bay (there is also a label
present bearing the locality “Port Curtis, ex Stutch-
bury”). Coll. Cuming. Size: 418 mm. Holotype:
4034 mm.

melanacme, Terebra — E. A. Smiru, 1875, A. M.N.
H., 15: 415, and 1877, A. M. N. H., 19: 224. Hab.
Cape Sima, Japan (Commander St. John). Coll. J.
G. Jeffreys. Size: 174mm (juvenile specimen).
Holotype no. 73.8.6.11: 17.8mm (spot of red wax
on second whorl above aperture). There are two
other specimens in the type collection, no. 1900.2.13.
58.7: 23.2 mm, and 11.8 mm, which are later addi-
tions since SmirH (1877) had only 1 juvenile speci-
men on hand at the time of description.

micans, Terebra — Hinps, 1844, P. Z. S. L., p. 157.
Hab. ? (“Cape Palmas, West Africa” on label). Coll.
Cuming. Size: 13 lin. (= 29.3mm). Holotype no.
1968248: 30.4mm; syntypes: 23.1 mm, and 21.8mm.
miranda, Myurella — E. A. Smitu, 1873, A. M.N.
H., 11: 267. Hab. Malacca. Coll. Cuming. Holotype:
24.7 mm; syntype: 20.8 mm. The locality is suspect,
since the types appear to be faded specimens of the
West American radula-aspera group of species.

9. modesta, Terebra — DEsSHAYES, 1859, P. Z. S. L., p.

288. Hab. L’embouchure de l’Indus. Coll. Cuming.
Size: 224mm. Holotype: 21.6 mm; syntype: 20.6
mm (on label is written “== T: matheroniana DEs-
HAYES, 1859).

morbida, Terebra — Rerve, 1860, Conch. Icon., plt.
24, spec. 133. Hab. ? Coll. Taylor. Size of figure:
13.8mm. Holotype no. 79.2.26.25: 14.1 mm.
nana, Terebra — DesuayeEs, 1859, P Z.S.L., p. 291.
Hab. L’embouchure de l’Indus. Coll. Cuming. Size:
1024 mm. Holotype: 9.9 mm.

Page 218

1

143.

144.

145,

146.

WAT):

148.

149,

150.

151.

—_—

152.

nassoides, Terebra — Hinps, 1844, P. Z. S. L., p. 158.
Hab. ? Coll. Cuming. Size: 6 lin. (= 13.5 mm).
Lectotype no. 1968251: 13.8mm; syntypes: 15.5
mm, and 13.9 mm. This appears to be the same spe-
cies as T: bridgest Dat, 1908.

nimbosa, Terebra — Hinps, 1844, P. Z.S.L., p. 151.
Hab. ? Coll. Cuming. Size: 25 lin. (= 56.3 mm).
Lectotype: 58.8 mm; syntypes: 64.9mm, and 58.7
mm (worn and faded), and 52.66mm. = Hastula
hectica (LinnaEus, 1758).

nitida, Terebra — Hinps, 1844, P. Z. S. L., p. 152.
Hab. Marquesas. Coll. Belcher. Size: 10 lin. (= 224
mm). Holotype no. 1844.6.7.85: 20.5 mm.
nodularis, Terebra — DEsHAYES, 1859, P. Z.S.L., p.
295. Hab. Les Isles Sandwich. Coll. Cuming & Des-
hayes (“ex Pease” on label). Size: 356 mm. Holo-
type: 346 mm; syntypes: 37.1 mm, and 33.0 mm.
obesa, Terebra — Hinps, 1844, P Z. S. L., p. 158.
Hab. ? Coll. Cuming; unique! Size: 6 lin. (= 13.5
mm). Holotype no. 1968250: 13.6 mm. The labrum
of the aperture is lirate in the type.

pallida, Terebra — Desuayes, 1857, J. C. P, 6: 87,
plt. 4, fig. 3. Hab. Les isles Marquises. Coll. Cuming.
Size: 72<11mm. Holotype: 71.8mm; syntypes:
59.2 mm (erroneously marked with x inside aper-
ture), and 51.9mm. The holotype is the same spe-
cies as T’ punctatostriata Gray, 1834; the two smaller
syntypes are T: cingulifera LAMarckK, 1822.

parva, Terebra (Leiostoma) Bairp in BRENCHLEY,
1873, Cruise H. M. S. Curacao, p. 435, plt. 37, figs.
5, 6. Hab. New Caledonia. Size: 8-11 lin. & 2 lin.
(= 18-24.8mm 4.5 mm). Lectotype no. 56: 22.8
mm (marked with an x on the tablet near the apex
of the shell) ; paralectotype no. 65: 18.4 mm.
paucistriata, Myurella — E. A. Smirn, 1873, A. M.
N. H., 11: 269. Hab. Ovalau, Fiji Islands (J. Mac-
Gillivray, H. M. S. Rattlesnake). Size: 19x4 mm.
Holotype no. 1856.11.3.17: 18.6 mm.

peasu, Terebra — Desuayes, 1859, P. Z. S. L., p.
302. Hab. Les Iles Sandwich (“ex Pease” on label).
Size: 45 <9 mm. Holotype: 45.6 mm.

pellyi, Terebra — E. A. Smitu, 1877, A. M. N. H.,
19: 226. Hab. Persian Gulf (Colonel Pelly). Coll.
R. MacAndrew. Size: 133 mm. Holotype no. 73.7.
5.1: 12.4 mm (spot of red wax on whorl above aper-
ture and marked with x under specimen on tablet).
There are additionally 7 syntypes in the type collec-
tion, all smaller than the holotype.

penicillata, Terebra — Hinps, 1844, P. Z. S. L., p.
157. Hab. Seychelles. Coll. Belcher & Cuming (“ex
Rousseau” on label). Size: 17 lin. (= 38.3 mm).
Holotype no. 1968244: 40.0 mm; syntypes: 34.4 mm,
and 26.4 mm.

THE VELIGER

153.

WS),

155.

156.

Hore

158.

159.

160.

161.

162.

163.

Vol. 11; No. 3

persica, Terebra — E. A. Smitu, 1877, A. M. N. H.,
19: 225. Hab. Persian Gulf (Colonel Pelly). Coll. R.
MacAndrew. Size: 123mm. Holotype no. 73.7.
5.4: 11.4mm; syntypes: 10.0 mm, 9.0mm, and 8.1
mm.

petiveriana, Terebra — DesHayeEs, 1857, J. C. P, 6:
85, plt. 5, fig. 10. Hab. Panama. Coll. Cuming. Size:
4210 mm. Holotype: 43.0 mm (marked with x in-
side aperture); syntype: 409mm. = T: radula
Hinps, 1844.

philippiana, Terebra — DesHayEs, 1859, P. Z. S. L.,
p. 289. Hab. isles Marquises? Size: 824mm. Lec-
totype: 9.3 mm. Two specimens have been previous-
ly attached to the tablet on either side of the lecto-
type, but these are missing now. == Hastula albula
(MENKE, 1843).

polygonia, Terebra — Reeve, 1860, Conch. Icon., plt.
27, spec. 154. Hab. ? Coll. Taylor. Size of figure:
10.6mm. Holotype no. 74.12.11.302: 10.7 mm.
polygyrata, Terebra — DesHAYES, 1859, P. Z. S. L.,
p. 301. Hab. Les fles Philippines. Coll. Cuming.
Size: 133mm. Holotype: 12.6 mm.

praclonga, Terebra — DEsSHAYES, 1859, P. Z. S. L., p.
315. Hab. Port Curtis. Coll. Cuming. Size: 939
mm. Holotype: 92.4mm; syntype: 102.6mm (er-
roneously selected as the “type” in the collection).
= T. triseriata Gray, 1834.

pretiosa, Terebra — ReEve, 1842, P. Z. S. L., p. 200
& 1842, Conch. Syst., 2: plt. 274, fig. 2. Hab. ?
(“China” on label). Coll. Stainforth. Size: 59/16
+ poll. (= 141.317.4 mm). Obtained by Cum-
ing; unique! Holotype: 140 mm; another specimen,
136.6mm long, is present in the type collection
which is not a type, as the specimen was said to be
unique at the time of description.

pulchella, Terebra — Desuayes, 1857, J. C. P, 6:
94, plt. 5, fig. 4 (non Epitonium pulchellum Noopt,
1819). Hab. les mers de la Chine. Coll. Cuming. Size:
40 <8 mm. Holotype: 39.2 mm; syntype: 41.2 mm
(erroneously marked x inside aperture).

pulchra, Terebra — Hinps, 1844, P. Z. S. L., p. 151.
Hab. Marquesas. Coll. Belcher. Size: 11 lin. (= 24.7
mm). Holotype: 24.7/mm. = T. cerithina La-
MARCK, 1822 (juvenile specimen).

punctatostriata, Terebra — Gray, 1834, P.Z.S.L., p.
61. Hab. ? Coll. J. E. Gray. Size: 22” (= 69.9 mm).
Holotype no, 74.11.10.1: 70.2mm. This species is
not conspecific with T. cingulifera Lamarck, 1822,
but is an earlier name for T: pallida Desuayes, 1857.
pura, Terebra — Desuayss, 1857, J. C. P, 6: 82, plt.
5, fig. 8. Hab. Zanzibar. Coll. Cuming. Size: Not
given in 1857; (69>11mm in 1859). Holotype:
64.7 mm (marked x inside aperture, also on whorl

Vol. 11; No. 3

above aperture and near apex of the shell on the
tablet). Syntypes: 51.5 mm, and 45.8 mm.

. pustulosa, Terebra — E. A. Smiru, 1879, P. Z. S. L.,
p. 186 (nom. nov. pro T. granulosa E. A. Situ,
1873 — non Lamarck, 1822). For types see under
T. granulosa E. A. Smitu.

. radula, Terebra — Hinps, 1844, P. Z. S. L., p. 155
(non GraveNnHorsT, 1807). Hab. Puerto Potrero,
west coast of America. Coll. Cuming. Size: 19 lin.
(= 42.7mm). Holotype no. 1968236: 40.8 mm.
= T. panamensis Dati, 1908 (fide CAMPBELL in
Keen, 1968), but 7: glauca Hinps, 1844 is a small
specimen of T: radula. Terebra aspera Hinps, 1844,
is not T. variegata Gray, 1834, as suggested by KEEN
(1968), but is rather similar to T: radula; it differs
in having 3 prominent spiral rows of coarse nodules
on each whorl which do not have the tendency to
form axial ribs; this, however, may be a variable
factor.

. reevei, Terebra — DesHayeEs, 1857, J. C. P, 6: 88,
plt. 4, fig. 14. Hab. les iles Moluques. Coll. Cuming.
Size: 5211 mm (9211 m min 1859). Holotype:
50.9 mm; syntypes: 50.0 mm, and 45.5 mm.
= Duplicaria duplicata (Linnarus, 1758).

. regina, Terebra — Desuayes, 1857, J. C. P, 6: 67,
plt. 3, fig. 7. Hab. le Sénégal (“Loanda” on label).
Coll. Cuming. Size: 78*13mm (8913 mm in
1859). Holotype: 76.3mm. = T? corrugata, La-
MARCK, 1822.

. remanalva, Terebra — Me viii, 1910, A. M. N. H.,
6: 12, plt. 2, fig. 21. Hab. Persian Gulf, Bundo Ab-
bas and Bushire. Size: 31><7 mm. Holotype no.
1911.6.21.11: 33.8 mm (marked x inside aperture) ;
paratype no. 1911.6.21.12: 229mm. = a worn
and faded T: spectabilis Hinps, 1844?

. robusta, Terebra — Hinps, 1844, P. Z.S. L., p. 149.
Hab. Panama, Gulf of Nicoya, Gulf of Papagayo
and San Blas, west coast of America. Coll. Belcher
& Cuming. Size: 57 lin. (= 128.2 mm). Lectotype
no. 1968171: 117.4mm; syntypes: 136.7 mm, and
88.0 mm.

. roseata, Terebra — ADAMS & REEVE, 1850, Voy. Sam-
arang, Zool., p. 30, plt. 10, fig. 24. Hab. Sooloo Is-
lands. Size of figure: 29.5 mm. Holotype: 29.2 mm;

GES VELIGER

—

Page 219

variegata Gray, 1834.

. rufopunctata, Terebra (Hastula) — E. A. Smitu,

1877, A. M.N. H., 19: 229. Hab. ? Coll. J. E. Gray.
Size: 225 mm. Holotype no. 74.10.12.3: 21.6 mm
(spot of red wax on whorl above aperture) ; syntypes:
19.9mm, 19.6mm, and 17.4 mm. = Hastula
trailli (DesHayEs, 1859).

. Sallaeana, Terebra — Desuayes, 1859, P. Z. S. L.,

p. 287. Hab. Mexico (Sallé). Coll. Cuming. Size:
245mm. Holotype: 2355 mm; syntypes: 25.3
mm, and 23.2mm. = Hastula cinerea (Born,
1778).

. serotina, Terebra — AvdAMS & Reeve, 1850, Voy.

Samarang, Zool., p. 30, plt. 10, fig. 20. Hab. Japan
Island, Nagasaki Bay (“ Isle of Luzon” on label).
Size of figure: 45.0mm. Holotype: 48.3 mm; syn-
types: 55.8mm, and 52.2mm. The syntypes may
possibly be later additions from the Philippines.
= T. anilis (R6ptnc, 1798).

. severa, Terebra — MEtvitt, 1897, M. P M. L. B.S.,

41 (7): 9, plt. 6, fig. 8. Hab. Mekran coast. Size:
143.4 mm. Holotype no. 97.7.30.103: 14.8 mm.

. sicyodes, Terebra nitida var. — MELVILL & SYKES,

1898, P. M.S. L., 3:.43, plt. 3, fig. 8. Hab. Andaman
Islands. Size: 37><6mm. Holotype no. 98.4.30.4:
35.0mm. = T! nitida Hinps, 1844.

. similis, Terebra — F.. A. Smitu, 1873, A. M. N. H.,

11: 265. Hab. ? (“Madras” on label). Coll. Cuming.
Size: 226mm. Questionable holotype: 27.4 mm.
(== “T. concolor E. A. Smirn” on label).

. solida, Terebra — Desuayes, 1857, J. C. P, 6: 78,

pit. 3, fig. 11. Hab. le Japon. Coll. Cuming. Size:
308mm. Holotype: 29.9 mm.

. sowerbyana, Terebra — Desuayes, 1857, J. C. P, 6:

93, plt. 3, fig. 8. Hab. la mer de Gambie (“River
Gambia, Mr. Deal” on label). Coll. Cuming. Size:
Not given in 1857, 5612 mm in 1859. Holotype:
56.3 mm.

. Specillata, Tercbra — Hinps, 1844, P.Z.S.L., p. 155.

Hab. San Blas (“San Blas, Mexico” on label). Coll.
Belcher. Size: 20 lin. (== 45.0mm). The only speci-
men present in the type collection is a Cuming speci-
men no. 1844.6.7.84: 39.3 mm, and is most prob-
ably a paratype.

syntypes: 25.2 mm, 21.0 mm; 20.0 mm, and 6.8 mm 182. speciosa, Terebra — Desnayes, 1859, P. Z. S. L., p.
(juvenile specimen). 279 (non BEAN in TuHorPeE, 1844). Hab. ? (“West

171. rudis, Terebra — Gray, 1834, P Z. S. L., p. 60. Africa” on label). Coll. Cuming & Deshayes. Size:
Hab. ? Coll. J. E. Gray. Size: 14” (= 38.1 mm). 397 mm. Holotype: 39.8X7.6mm. = T. sene-
Holotype no. 74.11.10.11: 36.8mm. = T. dislocata galensis LAMARCK, 1822.

Say, 1822. 183. spectabilis, Terebra — Hinps, 1844, P. Z. S. L., p.

172. rufocinerea, Terebra (Myurella) — CARPENTER, 1857 150. Hab. Guinea and Sumatra. Coll. Cuming (“ex

= Gee Iasi, IGOR, jo, 4243, poll, SS, ie, fo, = IE E. Layard” on label). Size: 134 lin. (= 30.4 mm).

Page 220

THE VELIGER

Vol. 11; No. 3

184.

185.

186.

187.

188.

189.

190.

191.

—

192.

193.

Questionable syntypes: 49.7 mm, 38.0 mm, and 35.8
mm. (“= T. gracilis Gray, 1834” on label).
straminea, Terebra — Gray, 1834, P. Z.S. L., p. 62.
Hab. ? Coll. J. E. Gray. Size: 23” (= 63.5 mm).
Holotype: 63.0mm (marked x inside aperture).
There are another 2 larger specimens in the type
collection ex coll. Cuming from the Philippine Is-
lands, which are not type material. = worn and
faded T. anilis (Rop1nc, 1798).

striata, Terebra — Gray, 1834, P. Z.S. L., p. 60 (non

Basterot, 1825; nec Quoy « Garmarp, 1833). Hab.

? Coll. J. E. Gray. Size: 14” (= 38.1 mm). Holo-
type no. 74.10.11.3: 35.0 mm (juvenile specimen).
= T. babylonia Lamarck, 1822.

stylata, Terebra — Hinps, 1844, P. Z. S. L., p. 152.
Hab. Japan, Philippine Islands (“Cagayan, Island
of Mindanao” on label). Coll. Cuming. Size: 21 lin.
(= 47.3mm). Holotype no. 1968230: 48.8 mm;
syntypes: 49.3mm, and 38.0 mm.

subnodosa, Terebra (Myurella) — CarPENTER, 1857
— see KEEN, 1968, p. 428, plt. 58, fig. 72, = T.
intertincta Hinps, 1844.

subtextilis, Terebra — E. A. Smiru, 1879, P. Z. S. L.,
p. 185, plt. 19, fig. 3. Hab. Station 21; Japan. Coll.
J. G. Jeffreys. Size: 376mm. Holotype no. 78.10.
16.28: 35.0mm. = T! textilis Hinps, 1844.
succinea, Terebra — Hinps, 1844, P. Z. S. L., p. 149.
Hab. ? Coll. Cuming. Size: 54 lin. (= 121.5 mm).
Probable holotype no. 1968172: 113.5 mm (marked
x inside aperture); syntypes: 100.6mm, and 78.8
mm (juvenile specimen).

sumatrana, Terebra strigilata — THIELE, 1925, Wiss.
Erg. Deut. Tief. Exp. Valdivia, 17: 344, plt. 29, fig. 20.
Hab. Sumatra. Ex coll. Berlin Museum. Paratypes
no. 1948.12.10.3-4: 18.7 mm, and 12.8 mm.

= Hastula strigilata (LInNAEUs, 1758).

suspensa, Terebra — EK. A. Situ, 1904, Journ. Ma-
lac. 11: 30, plt. 2, fig. 12. Hab. Port Elizabeth (“‘Pt.
Alfred, Cape Colony; ex Turton” on label). Size:
204.5 mm. Holotype no. 90.3.12.19.858: 19.8 mm.
Syntypes no. 90.3.12.19.859-862: 18.0 mm, 17.5 mm,
16.3 mm, and 16.1 mm.

swainsoni, Terebra — DESHAYES, 1859, P. Z. S. L., p.
299. Hab. Les Isles Sandwich. Coll. Cuming (“ex
Pease” on label). Size: 305 mm. Holotype: 30.0
6.0 mm; syntypes: 30.2 mm, and 26.5 mm.
tantilla, Myurella — E. A. Smitu, 1873, A. M. N. H.,
11: 270. Hab. Japan (A. Adams). Coll. Cuming.
Size: 63>2$mm. Holotype no. 73.7.5.2: 7.5mm
(marked with an x above the apex of the shell on the
tablet). There are another 3 specimens in the type
collection, ex coll. R. MacAndrew, mounted on the

194.

NE,

196.

197,

198.

IS),

200.

201.

—

202.

203.

same tablet on the extreme right, but these are not
type material.

taylort, Terebra — Reeve, 1860, Conch. Icon., plt.
23, spec. 124. Hab. Torres Strait, Australia. Coll.
Taylor. Size of figure: 27.2 mm (? enlarged). Prob-
able holotype no. 74.12.11.300: 19.3 mm.

tenera, Terebra — Hinps, 1844, P Z. S. L., p. 158.
Hab. Straits of Malacca; Ceylon. Coll. Belcher.
Size: 4 lin. (= 9.0 mm). Three syntypes present in-
stead of two, no. 44.6.7.86-67: 6.2 mm, 5.9 mm, and
5.8 mm; one of the three specimens is not a type.
textilis, Terebra — Hinps, 1844, P. Z. S. L., p. 156.
Hab. Sorsogon, Bay of Manila, Philippines; Straits
of Macassar. Coll. Cuming & Belcher. Size: 114 lin.
(= 25.9mm). Holotype: 25.7 mm (spot of red
wax on whorl above aperture and x marked inside
aperture) ; syntypes: 22.4mm, and 22.3mm. An-
other 15.2 mm specimen ex coll. Belcher is a different
species.

thyraea, Terebra (Euryta) — Me.vi.i, 1897, M. P.
L. P.S., 41 (7): 10, plt. 6, fig. 13. Hab. Karachi and
Mekran coasts. Size: 124 mm. Holotype no. 97.7.
30.89: 11.9 mm.
tiarella, Terebra — DEsHaAYES, 1857, J. C. P, 6: 91,
plt. 5, fig. 7. Hab. Natal. Coll. Cuming. Size: 328
mm. Holotype: 31.6mm; syntype: 31.1mm. The
locality indication is erroneous, as the types appear
to be conspecific with T: fitchi Berry, 1958.
torquata, Terebra — ADAMS & REEVE, 1850, Voy.
Samarang, Zool., p. 30, plt. 10, fig. 13. Hab. China
Sea (“Nagasaki Bay, Japan” on label). Coll. Cum-
ing. Size of figure: 45.0mm. Holotype: 45.5 mm;
syntype: 52.1 mm. There are another 2 specimens in
the type collection, 34.1 mm, and 22.9 mm in length,
ex coll. J. G. Jeffreys, which are probably later ad-
ditions; the locality of “Japan” on the label prob-
ably applies to the Jeffreys specimens.

traillu, Terebra — DEesHayes, 1859, P. Z. S. L., p.
285. Hab. Vasigapatam, Océan Indien (“Madras
Presidency, India” on label). Coll. Cuming. Size:
2343 mm. Lectotype: 23.84.6 mm; syntypes:
24.9 mm, 24.2 mm, and 22.4 mm.

tricincta, Terebra — E. A. Smiru, 1877, A. M.N. H.,
19: 225. Hab. Persian Gulf (Colonel Pelly). Coll.
R. MacAndrew. Size: 1123mm. Holotype no.
73.7.5.6: 10.9 mm; syntype: 10.8 mm.

triseriata, Terebra — Gray, 1834, P. Z. S. L., p. 62.
Hab. ? Size: 12’ (= 44.5 mm). Holotype no. 74.9.
9.1: 46.7 mm.

trismacaria, Terebra — MEtviLu, 1917, J. C. L., 15:
188, text fig. Hab. Mekran coast, probably off Char-
bar. Size: 184mm. Holotype no. 1925.3.12.14:

Vol. 11; No. 3 THE VELIGER Page 221
170mm. = T. polygyrata DesHayeEs, 1859. CAMPBELL in KEEN, 1968), but T: larvaeformis
204. tristis, Terebra — DESHAYES, 1859, P. Z.S. L., p. 306. Hinps, 1844 is the same species.

205.

206.

207.

208.

209.

210.

Hab. les mers du Japon (= error! the species occurs
in New Zealand waters). Coll. Cuming. Size: 199
mm. Holotype: 17.7 mm; another specimen, 14.8 mm
in length, is a J. E. Gray specimen. Deshayes’ stated
width of 9 mm is obviously an error for 6mm.
trochlea, Terebra — DesHayeEs, 1857, J. C. P, 6: 89,
plt. 5, fig. 6. Hab. Zanzibar. Coll. Cuming. Size:
6513 mm (6913 mm in 1859). Holotype: 65.3
mm (marked x inside aperture) ; syntype: 50.5 mm
(juvenile specimen).

tuberculosa, Terebra — Hinps, 1844, P. Z. S. L., p.
154. Hab. Panama, Gulf of Papagayo and San Blas.
Coll. Belcher. Size: 24 lin. (= 54.0 mm). Holotype
no. 1844.6.7.82: 55.7 mm (marked x inside aperture) ;
syntypes (ex coll. Cuming, Panama): 46.4 mm, and
42.6mm. The holotype of T: tuberculosa is not like
the shell figured for the species by CAMPBELL (1964)
on plt. 17, fig. 7, but rather similar to T. cracilenta
Li, 1930, as figured by the same author on plt. 17,
fig. 9. The penultimate whorl has 4 spiral rows of
small nodules and the body whorl 7 rows up to the
peripheral ridge.

turrita, Myurella — E. A. Smiru, 1873, A. M.N. H.,
11: 266. Hab. Torres Strait. Coll. Cuming. Size: 26
44 mm. Holotype: 23.0mm. = T: exigua Des-
HAYES, 1859.

undulata, Terebra — Gray, 1834, P. Z. S. L., p. 60.
Habs Size; 13/7 (= 38mm); Coll, J, Ey Gray.
Lectotype no. 74.11.10.5: 33.6mm; syntypes: 33.3
mm, and 32.7 mm.

ustulata, Terebra — DesHayeEs, 1857, J. C. P, 6: 97,
plt. 3, fig. 12. Hab. la Terre de Van Diemen. Coll.
Cuming. Size: 3510mm. Holotype: 35.0 10.7
mm (marked x inside aperture) ; syntypes: 36.9 mm,
and 31.3 mm. There are another 3 specimens in the
type collection which are not type material but have
been purchased at a later date.

varicosa, Terebra — Hinps, 1844, P. Z. S. L., p. 152
(non Buccinum varicosum GMELIN, 1791; nec Tereb-
ra varicosa Bosc, 1801). Hab. Gulf of Papagayo,
west coast of Central America. Coll. Belcher. Size:
11 lin. (= 248mm). Holotype no. 1844.6.7.88:
25.0 mm; questionable Cuming syntype: 19.8 mm.
It is suspected that sometime during labelling the
smaller Cuming specimen has been interchanged
with the larger Belcher specimen; from the dimen-
sions supplied by Hinps, it is evident that the larger
holotype originated from the Belcher collection.
= T. brunneocincta Pirspry & Lowe, 1932 (fide

211. variegata, Terebra — Gray, 1834, P. Z. S. L., p. 61.
Hab. ? Coll. J. E. Gray. Size: 24” (= 63.5 mm).
Holotype: 61.4mm (marked x inside aperture).
There are 2 other specimens in the type collection
(“Guaymas, Gulf of California, ex Mr. Babb?” on
label) 78.0mm and 61.7 mm in length, which are
later additions and not type material.

venilia, Terebra — TENtISoN-Woops, 1880, Proc.
Timn. Soc. N.S. W,.4: 23, plt. 4; figs! 2) 2a: Hab:
Sow and Pigs reef, Port Jackson. Size: 4274 mm.
Holotype no. 96.11.30.28: 41.1 mm.

venosa, Terebra — Hinps, 1844, P. Z. S. L., p. 157.
Hab. ? Coll. Cuming. Size: 16 lin. (= 36.0mm).
Holotype no. 1968245: 35.0 mm; syntypes: 34.9 mm
and 27.0 mm. = Hastula penicillata (Hinps,
1844).

virginea, Terebra — DESHAYES, 1857, J. C. P, 6: 83,
pit. 4, fig. 12. Hab. Zanzibar. Coll. Cuming. Size:
5211mm. Holotype: 52.3 mm. —= worn and
faded T: consors Hinps, 1844.

walkeri, Terebra — E. A. Smiru, 1899, P. M. S. L.,
3: 312, fig. 1. Hab. Holothuria Banks, North-West
Australia; J. J. Walker. Size: 266mm. Holotype
no. 91.11.21.326: 26.2 mm (juvenile specimen).
wilkinsi, Strioterebrum (Partecosta) — DANCE &
Eames, 1966, P. M.S. L., 37: 42, plt. 3, fig. 4 & pit.
5, fig. 5. Hab. Fao, Persian Gulf. Coll. W. D. Cum-
ming. Size: 9.22.8 mm. Holotype no. 1893.12.15.
74: 9.2 mm; plus 6 paratypes nos. 1893.12.15.75-80.
= Terebra fuscobasis E. A. Smitu, 1877.

Tomutn (1944) did not locate the type specimens of
the following species: Terebra archimedis Desuayes, T:
buccinulum Desuayes and T: lactea DESHAYEs; these
types were in the type collection when I examined them.
However, I did not find the type of T: histrio DESHAYES,
1857, which Tomuin (loc. cit.) reported as being in
Cuming’s collection.

For 14 species Tomutn reported a single type in each
case but I found 2 or more syntypes present in the type
collection. It is obvious that in the short span of time
since ToMLIN reported on DEesHAyeEs’ Terebra types, the
type collection has undergone changes, and more types
are now present in the type collection than there were
in 1944. It is probable that additional types have been
isolated and removed from the general collection. Apart
from bona fide holotypes and lectotypes, remaining syn-
types are E. A. Smith’s, Melvill, and Melvill and Sykes
material excluded, rather questionable paratype material.

PAE

213.

Blab,

“Alle

216.

Page 222

LITERATURE CITED

CAMPBELL, G. BRUCE
1963. Rediscovery of Terebra formosa DrsHayes, 1857.
The Veliger 5 (3): 101-103; plts. 12, 13.
(1 January 1963)
1964. | New terebrid species from the eastern Pacific (Mollusca:
Gastropoda). The Veliger 6 (3): 132-138; plt. 17
(1 January 1964)

DEsHAYES, GERARD PAUL

1857. Descriptions d’espéces nouvelles du genr: Terebra.
Journ. de Conchyl. 6: 65 - 102; plts. 3-5
1859. A general review of the genus Jerebra, and a description

Proc. Zool. Soc. London, 270 - 321

of new species.

5 pits.
Gray, JoHN Epwarp
1854. | Untitled. [Terebra] _ Proc. Zool. Soc. London, Part II
(1834) : 50 - 63 (50 - 56: 26 Sept.; 57-63: 25 Nov. 1834)

Hinps, Ricrarp BrRINSLEY

1844. | Description of new shells, collected during the voyage
of the Sulphur, and in Mr. Cuming’s late visit to the Philip-
pines. Proc. Zool. Soc. London for 1843, pt. 11: 149 - 168
(June 1844)

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Vol. 11; No. 3

Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks

from Lower California to Colombia. i- xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)

1966. | West American mollusk types in the British Museum
(Natural History), II. Species described by R. B, Hinps.
The Veliger 8 (4) : 265 - 275; plts. 46, 47; 6 text figs.
(1 April 1966)
1968. West American mollusk types at the British Museum
(Natural History). - IV. Carpenter's Mazatlan collection.
The Veliger 10 (4): 389-439; plts. 55-59; 171 text figs.
(1 April 1968)
REEvE, LovELL AuGcustus
1860. | Conchologia Iconica. Monograph of the genus Terebra.
13: plts. 1-27 (May - June 1860)
SmitH, Epcar ALBERT
1873. | Remarks on a few species belonging to the family Tere-
bridae, and descriptions of several new forms in the collection
of the British Museum. Ann. Mag. Nat. Hist., (ser. 4),
11: 262 - 271

1877. Descriptions of new species of Conidae and Terebridae.
Ann. Mag. Nat. Hist., (ser. 4), 19: 222 - 231

ToMuin, Joun Reap LE BrockroNn

1944. Deshayes’ review of Terebra.
104 - 108

Journ. Conch. 22 (5):
(30 November 1944)

Vol. 11; No. 3

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Page 223

The Ecology of Macoma inconspicua (Broprrip & SOWERBY, 1829)

in Central San Francisco Bay.

Part I. The Vertical Distribution of the Macoma Community

BY

MARILYN T: VASSALLO

Department of Biological Sciences

California State College at Hayward, Hayward, California 94542 '

(Plate 44; 6 Text figures; 2 Tables)

INTRODUCTION

THE PRESENT STuDy of the distribution of Macoma
inconspicua (BRODERIP & SOWERBY, 1829) (== Macoma
balthica LinNAEus, 1758) community of central San
Francisco Bay is of significance as no systematic subtidal
or intertidal studies have been made in the expanse of
shallow water (less than 5m in depth) covering 48%
of the central bay.

The purpose of this research was fourfold: (1) to
delineate the vertical distribution of the fauna, (2) to
describe the physical structure of the community within
the mud, (3) to observe the interactions among the in-
vertebrate fauna, and (4) to compare this community
to parallel communities investigated by other authors.
The second aspect of the study will be discussed in a later
paper.

In this paper the name Macoma inconspicua is treated
synonymously with M. balthica as in previous studies on
this species in the San Francisco Bay region.

LOCATION or tHe STUDY

San Francisco Bay is located in the central portion of
California. For purpose of discussion, it is convenient to
divide San Francisco Bay into 3 sections: northern, cent-
ral and southern. Because the United States Coast and

' Present address: Department of Biological Sciences, University
of Victoria, Victoria, B. C., Canada

Geodetic Survey does not recognize these divisions, pre-
vious authors have used conflicting boundaries.

Although the U.S. Coast and Geodetic Survey has not
defined north, central, and south San Francisco Bay as
such, its charting system divides the northern region
from the central portion in maps 5531 and 5532. Central
San Francisco Bay is here defined as bounded on the north
by a line extending from Hunters Point to Bay Farm
Island and on the south by the San Mateo Bridge. These
boundaries were used by Gram (1966) in his compre-
hensive study of the marine geology of central San
Francisco Bay.

Benthic samples analyzed in this study were taken from
an intertidal mud flat bordering the ponds of the Oliver
Brothers’ Salt Company on the east side of central San
Francisco Bay one half mile north of the San Mateo
Bridge (Text figure 1).

PREVIOUS ECOLOGICAL STUDIES

Previous ecological studies have, for the most part, been
restricted to subtidal work by investigators who oper-
ated a dredge from a large boat or ship. The U.S.S.
“Albatross”, commissioned by the U.S. Bureau of Fish-
eries to make a biological survey, cruised San Francisco
Bay from January 1912 to April 1913 (Packarp, 1918a).
A report of the physical conditions from Suisun Bay to
San Mateo Point based on data taken during this cruise
was made by SuMNER et al. in 1914. This work was
preceded by the limited geological investigations made

Page 224

from the Russian ship “Rurik” in 1816 (in Gram, 1966)
and the biological investigations of H. M.S. “Blossom”
in 1826 (RosEwatTEeR, 1968).

The report by SuMNeER et al. (1914) was the basis of
ecological studies made on the molluscan fauna by E. L.
Packarp. His first report was a qualitative study based
on the fauna obtained from “several common types of
dredges” (1918a). The second study was quantitative
based on material taken with an orange peel dredge, the
first time such a dredge was used in biological studies
(Packarp, 1918b).

Another early investigation of the physical factors of
the bay was that of Miter et al. in 1928.

Extensive work in the northern part of the bay has
been carried out by Fruice (1954a, 1954b, 1958) and M.
L. Jones (1961). In his work on the distribution of in-
vertebrates in the northern bay Finice used both Ekman
and Petersen dredges. Jones collected his samples off
Point Richmond with a core sampler.

An investigation of water sediment quality and pollu-
tional characteristics of the bay was made by McCarty
(1962). This study was followed by a faunal study in
which samples were taken with an orange peel dredge in
the north and central bay (Storrs et al., 1965). Most
recently, samples from 6 stations in the north and central
bay were taken, again with an orange peel dredge, by
Apuin (1967).

PHYSICAL CHARACTERISTICS
OF THE ENVIRONMENT

TIDES:

On the Pacific coast two high tides of unequal magni-
tude and two low tides also of different heights occur
during each lunar day. San Francisco Bay communicates
to the ocean through the narrow Golden Gate. The
narrow opening creates a lag in the time of low and high
tides between the entrance to the bay and the southern
reaches of the bay. At the San Mateo Bridge low tide is
1 hour 14 minutes later and 0.1 ft higher than at San
Francisco (Fort Point). Miter (1928) and Gram (1966)
have emphasized that the time and height of the tide is
influenced by the prevailing winds. The wind is generally
from the northwest and 46% of the time is between 4
and 10 knots (Conomos, 1963).

WAVES:

Due to the shallow character of the central bay, the
prevailing winds produce an important amount of wave
activity. Gram (1966) calculates that in east central San

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Vol. 11; No. 3

Francisco Bay “the water depths are such that waves two
feet or less in height would readily disturb the bottom
material, thereby allowing the materials once deposited
to again become suspended and probably transported.”

The term wave front seems more appropriate than
surf as a description of the advancing edge of the tide
on a mud flat. The wave front of the incoming and out-
going tides suspends and carries sediment with it.

CURRENTS anp SEDIMENTS:

Ripples in a substrate indicate the presence of a cur-
rent. On the Oliver Brothers’ mud flat ripples are formed
on the mud closest to shore and tend to diminish towards
the lower part of the mud flat (Plate 44, Figures 1 to 3).

The upper zone is most frequently subjected to sedi-
ment suspension by the wave front. SANDERS (1956) and
Kitcuinec et al. (1952) note that suspended sediments
will tend to be deposited where the current which carries
them decreases in speed. ‘The largest particles are first to
settle out and then, as the current decreases, smaller and
smaller particles will settle.

On the present study site the combination of current
(as reflected by ripples) and wave action produces a
gradient in which coarse sediments are left in the upper
part of the mud flat and the finest settle in the lower part
of the flat.

METHODS anp MATERIALS

LINE TRANSECTS:

The relationship between tide level and distribution
was determined by taking core samples along 3 line tran-
sects. Because the slope of the mud was very slight (es-
pecially below the 1.5 ft tide level), a tidal difference of
one tenth of a foot would bare 30m to 70m of mud.
Exposure in this broad band of mud was the same, so
samples were taken at 36.6 m intervals.

The first line of 22 samples was taken from May 10
to June 3, 1967. This line was perpendicular to the shore-
line and extended up into a small creek which drains the
surrounding marshland.

A line parallel to the first and 180m south was sampled
the following month from June 10 to June 22. The
backshore of this area consisted of a rock and concrete
levee. Twenty-one samples were taken.

From July 1 to July 12 a third set of 20 samples was
taken along a line north of the first line lying at a 30°
angle to it. The backshore consisted of marshland (Text
figure 1).

Vol. 11; No. 3 THE VELIGER Page 225

———

100 meters

Ocean

{Hayward Air

*: Terminal

Pacific

70) ky US We iG WS
. ao 6

° ° ° °

43.42 41 40 39EIB 37) 736)9(35134 133) 32) 1311 30) 29) 28 517) 716

Figure 1

Number and location of samples taken on the Oliver Brothers’ mud flat. The scale on the figure refers only to the east-west dimension
of the tide flat which is somewhat foreshortened in the north-south direction. The map is based on an aerial photograph taken from an
oblique (south to north) angle.

Page 226

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Vol. 11; No. 3

CONSTRUCTION or tHe SAMPLERS:

Two parameters were utilized to measure the fauna of
the mud flat. The organisms in the mud were measured
as numbers of individuals per core (in cubic centimeters).
Those living on the mud surface were measured as num-
bers of individuals per quadrat (in square centimeters).

The core sampler consisted of an aluminum casing
which fit snugly over a halfgallon milk carton with its
base removed. The cross section of a half-gallon milk
carton is 9.7 cm by 9.7 cm; the area 94.09 cm?. This is
less than 6% short of a sampling area of 0.01 m?.

To obtain a sample of mud the casing with the carton
in it was pushed into the mud and brought up with a
shovel. The carton was prevented from collapsing by ap-
plying strips of masking tape to the inside of the carton
which were then extended through the battom of the
carton to the outside of the casing.

TIDE

2.6 1.0

Nassarius
obsoletus

Macoma

icons picua

Tharyx
parvus

Flosmaris
grandis

Ampelisca
miller

Mediomastus
californiensis

Gemma

gemma
Neanthes
succinea

Mya

arenaria

Marphysa
sanguinea

Hemigrapsus
oregonensis

Ostrea
1 lurida

0 100 200

Previous authors have sampled to varying depths for
Macoma inconspicua. FRASER (1932) went to a depth of
7.5 cm. No reason was given for this limitation, nor did
Rees (1940) explain why he sampled to a depth of
18cm. BEANLAND (1940) took only the top 2.5 cm be-
cause: “Previous work in the area seemed to indicate
that the surface inch contained a representative fauna.”
Brapy (1943) sampled to a depth of 25cm _ since the
organisms did not penetrate to a greater depth.

Preliminary work on the Oliver Brothers’ mud flat
revealed a dense layer of clay about 20 cm beneath the
surface. No animals penetrated into the clay; therefore,
a sampling depth of 21 cm was chosen.

Counting the numerous thread-like polychaetes was
facilitated by subsampling. At one corner of each sample
a 3cm by 3cm by 21 cm section was marked off. The
subsample was sieved through a 0.297 mm mesh under
running water. The upper 2 cm were also sieved through

LEVEL

ee ee ee ee ee

300 400 50cm 600

DISTANCE FROM SHORE IN METERS

Figure 2

A comparison of the 3 transects. Heavy line, transect 1; light line, transect 2; dashed line, transect 3. Single light line indicates distri-
bution of fauna not taken in the sampler in significant numbers.

Vol. 11; No. 3

THE VELIGER

Page 227

a 0.297 mesh. A geologic sieve of 0.5mm was used to
sort the remainder of the sample.

SAMPLING ror Nassarius obsoletus (Say, 1822):

The mud snail Nassarius obsoletus (Say, 1822) was
extremely abundant on the mud surface. The milk carton
sampler took an adequate volume of mud to obtain
significant numbers of the infauna; however, even at
optimum densities of N. obsoletus the surface area of
94 cm? could potentially take only 3 or 4 snails per sample.
In addition, the mud snail’s tendency to converge on a
given area when a choice morsel of food was found also
indicated a larger surface area should be taken.

An aluminum frame 50cm by 50cm by 5cm was
placed on the mud. The milk carton sample was taken in
one corner and then the snails within the frame were
picked up by hand.

Nassarius
obsoletus

Macoma Mya

inconspicua arenaria

WWE ile Wie te

(0)
Up

Tharyx
parvus

RESULTS

DISTRIBUTION or COMMON SPECIES:

The distribution according to tide level was the same
for the 3 transects (Text figure 2). The difference in
backshores did not appear to affect the faunal distribu-
tion except in the case of transect 1. The backshore of
this transect consisted of a creek bed and the mud
therefore extended beyond the marshland and the rock
levee of the other transects.

The upper zone from the 2.5 ft tide level to 1.3 ft was
characterized by Nassarius obsoletus, Macoma inconspi-
cua and Tharyx parvus BERKELEY, 1929. Specimens of
Streblospio benedictt WeBsTER, 1789 and Flosmaris gran-
dis HAND & BUSHNELL, 1967 were occasionally observed
in this region.

Neanthes Mediomastus

succinea

Marphysa
sanguinea

Ambelisca

milleri californiensis

ee dd
we ee tee eee eee e

Figure 3

Per cent of each population at a given tide level.

Page 228

From 1.2 ft to 0.9 ft several additional species ap-
peared in the samples. This area appeared to be the
upper limit for Mediomastus californiensis HARTMAN,
1944: Neanthes succinea (FREY & Leuckart, 1847) ;
Marphysa sanguinea (Montacu, 1804); Mya arenaria
Linnaeus, 1758; and Ampelisca milleri BARNARD, 1954.
In addition, small numbers of Gemma gemma (ToTTEN,
1834); Hemigrapsus oregonensis (DANA, 1851) and a
goby were recovered. This area corresponds to one of the
critical tide levels described by Doty (1946). At the
levels of 3.5 (LLHW), 3.0 (HHLW) and 1.0 (LHLW)
there is a sudden increase in duration of a single
exposure.

At the end of the second transect Ostrea lurida Car-
PENTER, 1864 was abundant on the surface of the mud.
Specimens of the slipper shell Crepidula convexa Say,
1822 were found on the oysters. Specimens of the ane-
mone Diadumene leucolena (VERRILL, 1866), two pos-
sessing catch tentacles (VASSALLO, in press), were also
found on the oysters.

Text figure 3 represents the per cent of individuals in
8 common populations found at a given tide level.

Nassarius obsoletus

A total of 348 mud snails were taken in 43 quarter
meter square samples. The maximum number taken in a
single sample was 36. There was a gradual decrease in
the numbers from the optimum region at 2.4 ft where
13% of the population was found.

Macoma inconspicua

Two hundred and fifty - six specimens of Macoma in-
conspicua were taken in 63 milk carton samples. The clam
was most abundant at the 2.6 ft tide level. A maximum
of 13 per core was found in the optimum region where
17.5% of the population occurred.

Mya arenaria

Seventy specimens of Mya arenaria were taken in the
core sampler. The upper limit was 1.3 ft. The optimal
region was ().9 ft. No specimens were taken beyond the
0.1ft level. No more than 3 specimens were recovered in
a single core.

Neanthes succinea

A total of 11 specimens of this polychaete was taken
with the core sampler. The maximum number per core
was 2. Neanthes succinea was evenly distributed over its
range. The upper limit found during this survey was 1.0 ft.

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Vol. 11; No. 3

Marphysa sanguinea

Thirteen specimens of Marphysa sanguinea were ob-
tained with the core sampler. No more than 1 specimen
was found per core. The upper limit was 0.8 ft.

Mediomastus californiensis

Subsampling was utilized in the recovery of Medio-
mastus californiensis from the milk carton core. Sixty-
three specimens of this polychaete were taken. The upper
limit of the polychaete was 1.3 ft. There was an increase
from 1 per subsample to 3 per subsample in the optimum
area at 0.9 ft.

Tharyx parvus

The cirratulid polychaete was found from the shoreline
out to a distance of 100 m. Limitation to the upper zone
is apparently not the direct effect of exposure per se since
Jones (1961) found it in high numbers at a station 3 ft
below MLLW.

Ampelisca milleri

Counts of this amphipod were made along one tran-
sect because of the difficulty they presented to counting.
Two hundred and fifty-five amphipods were taken in the
subsamples from transect 2. The upper limit was about
1.2 ft where 3 amphipods per subsample were found. The
numbers increased to the 0.7 ft level where a maximum
of 29 per subsample were found. This is a conservative
count. Sieving through a 0.297 mm mesh did allow some
amphipods to pass through. When the mud was screened
over trays, small amphipods passed through and within 2
days had built new tubes.

DISTRIBUTION or tHE DOMINANT SPECIES:

In a previous section on the sediments of the mud flat
it was noted that ripples in the upper zone indicated a
current was present along the shoreline. The current as
well as the wave front of the incoming tide suspended
sediment. Larger particles settled out first and lighter
ones settled as the current decreased.

At a distance of 150m from shore the ripples became
less distinct until they were no longer present 300m
from shore. The area between 150m and 300m was
intermediate in strength of current and in size and
amount of particles deposited on the surface. This zone
between the area experiencing strong currents and the
lower zone where no current was apparent might be con-

Tue VELIcER, Vol. 11, No. 3 [VAssaLLo] Plate 44

“y

Figure 3 Figure 4

Figure 1: Detail of well defined ripples in the upper zone. Six
inch rule for scale.

Figure 2: Detail of intermediate zone. Note faint ripples.

Figure 3: Lower zone of the mud flat. Ripples absent. Small pro-
jections from the mud are tubes of Ampelisca milleri. The snails
covered with detritus and diatoms are Nassarius obsoletus.

Figure 4: Indentations characteristic of Busycotypus canaliculatus
found in the lower zone of the mud flat.

Vol. 11; No. 3

°
/o
rl Macoma inconspicua

a] Other
Ampelisca milleri

THE VELIGER

Page 229

SLIGHT
CURRENT

INTERMEDIATE

CURRENT
STRONG
uw
x CURRENT
=
<
72)
a
wi
a
(2)
<
>
(a)
e
a
z
uw
o}
fe)
Zz

0 100 200

300 400 500

DISTANCE FROM SHORE IN METERS

Figure 4

The dominant species of the community changes with distance
from shore.

sidered an ecotone. The existence of an ecotone on an
intertidal mud flat was suggested by DExTER (1947).

In the upper zone of the mud flat from 2.6 ft to 1.3 ft
Macoma inconspicua formed 55% of all the individuals
in the community and the amphipod Ampelisca milleri
represented 1% of the individuals. In the intermediate
zone the per cent of M. inconspicua dropped to an aver-
age of 16% and the amphipod increased to 50%.

In the lowest zone from 0.8 ft to —0.1 ft Ampelisca mil-
leri was the dominant species forming 90% of the com-
munity. The change of dominance is shown in Text figure
4. At the extreme ends of the figure two communities
are evident.

DISCUSSION

Tue Macoma COMMUNITY:

Macoma inconspicua occurs in subtidal (Forp, 1923;
TuHorson, 1957) as well as intertidal muds. Brapy
(1943), Spooner & Moore (1940) and Fraser (1932)
found no definite trend with respect to tide level. ALLEN
(1954) reports M. inconspicua in mud as well as mud-
sand. HotmMe (1949) found it in soil ranging from moist
sand to mud. Studies by BEANLAND (1940), ReEs (1940)
and Fraser (1932) indicate that soil grade in itself does
not appear to be the determining factor in the distribu-
tion of M. inconspicua.

Page 230

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Vol. 11; No. 3

BEANLAND suggests that the abundance of Macoma
inconspicua in estuaries depends on two factors: (a)
quality and quantity of available food supply correlated
with type of soil; and (b) available feeding time corre-
lated with distance below high tide mark.

Yonce (1949) found this species to be a detritus feeder.
In the field as well as in the laboratory Macoma incon-

1.2 Transect 1
Y= 1:0798 —- .0023 x

75 150 225 300 375 450

Transect 2
= 13838 — .0036x

8

=

“3

Dy

=

S

8

Ss

8

§

is)

S

s

75 150 225 300 375 450

O

Zz

) 1.2 é Transect 3

rs Y = 1:0796 —- .0022
—

75 150 225 300 375 450

DISTANCE FROM SHORE IN METERS

Figure 5
Regressions of the number of Macoma taken on the
Oliver Brothers’ mud flat

Spicua was observed sweeping its siphon over the surface
of the mud. In the course of the sweeping it would some-
times crook the end of the siphon into the mud as if
to loosen the surface of the mud. The inhalant siphons
extended several centimeters out of the mud. Although
they were long enough to extend towards the water above
them, the siphons remained sweeping at the detritus-
water interface.

McNutty et al. (1962) found detritus feeders most
abundant in fine soils. On the Oliver Brothers’ mud flat
the greatest amount of detritus occurs in the lower zone.
This area also has the greatest amount of feeding time,
yet, contrary to BEANLAND’s suggestion, Macoma incon-
spicua is most abundant in the upper zone where there
is less detritus and less feeding time.

In summary, the literature indicates that there does
not appear to be direct relationship between distribution
and exposure or particle size. The present study does not
support BEANLAND’s suggestion that detritus and avail-
ability of feeding time are important factors.

THE Ampelisca COMMUNITY:

Ampelisca milleri is found subtidally to a record depth
of 65 fathoms (Barnarp, 1967). In San Francisco Bay,
Jones (1961) found it to be abundant at a station 3 ft
below Mean Lower Low Water.

Jones (op. cit.) noted that the sediment was coarse
at the station where Ampelisca was abundant and finer
where it was present in smaller numbers. No other infor-
mation on the ecology of Ampelisca milleri was found.
Eneguist (1950) noted that the ampeliscid Haploops
cannot build its tubes in pure clay or coarse sand. The
tubes of A. milleri on the Oliver Brothers’ mud flat
were made of fine sediment. The need for tube construc-
tion could restrict the species to finer soils.

All of the ampeliscids studied by ENEQuisT were found
to be detritus feeders. In the present study Ampelisca
was observed in the laboratory to continuously swecp
detritus with its long antennae and move particles toward
its mouth — indicating this species may also be a detritus
feeder.

In the intermediate and lower zones of the mud flat
water does not drain completely from the mud flat. This
residual water could serve to protect Ampelisca from
desiccation. Hart (1930) suggests that such a layer of
water may protect Corophium volutator from desicca-
tion. Like Ampelisca milleri, C. volutator feeds on detri-
tus and lives at the mud surface in semi-permanent tubes
(Hart, op. cit.).

BEANLAND (1940) noted that when Corophium leaves
its tube, it is at the mercy of the currents. Ampelisca

Vol. 11; No. 3

THE VELIGER

Page 231

was observed caught in the movements of outgoing tides
on the Oliver Brothers’ mud flat. BEANLAND suggested
that shelter may be an important factor affecting the
distribution of Corophium. She noted that, where Coro-
phium was found in previous studies, it occurred in
sheltered arcas where the amphipod was protected from
currents which could sweep it away. GoopHarT (1941)
records that it is absent from steep banks where it
might be washed away by the current.

Transect 2

Y = 1-3700 + .0008 X

2.1

LOG NO. Ampelisca | miller

DISTANCE FROM SHORE IN METERS

Figure 6

Regression of the number of Ampelisca taken on
Oliver Brothers’ mud flat

In the Dovey Estuary, BEANLAND found that as shelter
decreases from shore to low water the numbers of Coro-
phium decrease. On the Oliver Brothers’ mud flat the
presence of ripples in the upper tide level and their ab-
sence in the lower level indicates that “shelter” in the
sense of less overturn of mud by the current increases
with a decrease in tide level. The upper tide level is also
less sheltered in the sense it is more frequently exposed to
the sediment stirring influence of the incoming and
outgoing tide.

In summary, the accumulation of detritus and the
presence of shelter seem to be important in affording
favorable conditions to the maintenance of the Ampelisca
community in the lower zone of the mud flat.

It could be argued that the distribution (abundance
in the upper flat) of Macoma inconspicua is due to phys-
iological differences in the San Francisco Bay population.

However, north of this study area in upper San Francisco
Bay, Fitice (1958) reports that M. inconspicua prefers
the intertidal and upper 5 subtidal feet.

Predation by Busycotypus canaliculatus (LINNAEUS,
1758) may be a factor limiting Macoma. Large inden-
tations made by the whelk have been found in the lower
zone of the mud flat (Plate 44, Figure 4). No living spe-
cimens or shells have been dug from the mud or have
been found on or near the Oliver Brothers’ mud flat.
However, hundreds of individuals of this species are
found on the mud flats in west central bay and fishermen
stranded by low tide about a mile from the east shore
have observed the whelk on the mud surface.

The fact that Busycotypus has never been directly ob-
served on the Oliver Brothers’ mud flat and the presence
of relatively few indentations (in contrast to abundant
numbers occurring on exposed flats of west central bay)
suggests that Busycotypus may not be a major limiting
factor.

Both in San Francisco Bay and the Dovey Estuary
(BEANLAND, 1940) numbers of detritus feeding amphi-
pods populate the mud flats. In the Dovey Estuary the
amphipod community dominates the upper tide levels
(BEANLAND, op. cit.). As the level decreases, the numbers
of amphipods decrease and the clam Macoma incon-
Spicua appears and gradually replaces the amphipod as
the dominant species. On the Oliver Brothers’ mud flat
it is M. inconspicua that dominates the upper tide level
and is replaced by an amphipod community at the lower

Table 1

Faunal Change with Distance from High Tide Mark
on the Oliver Brothers’ Mud Flat

Sample No. Ampelisca miller Macoma inconspicua
23 - 9
24 — 12
25 - 13
26 9 7
27 24 4
28 24 10
29 36 a
30 33 6
31 45 3
32 60 1
33 57 1
34 81 =
35 78 =
36 78 =
37 81 =
38 75 =
39 75 =

Ragves232

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Vol, Mig Noss

level. Tables 1 and 2 give the results of the present study
and that of BEANLAND’s study.In both areas the sample
number represents increasing distance from shore. It
should be noted that BEANLAND’s sampling area was one
meter square.

Table 2

Faunal Change with Distance from High Tide Mark,
BEANLAND, 1940

Sample No. Corophium volutator Macoma inconspicua
1 = =
2 445 1
3 24 -
4 191 1
5 44 11
6 2 2
7 844 6
8 455 2
9 82 1

10 Di2 2
11 20 1
12 29 -
13 11932 16
14 1 11
15 5 7
16 24 15
17 2 19
18 1 8
19 - 401
20 = 195
21 - 369
22 - 416
23 - 657
24 - 58
25 - 4
26 - -

2 Probably a mixture of Corophium volutator and C. arenarium

The Tables and the regressions in Text figures 5 and 6
show an inverse relationship between the amphipod and
Macoma inconspicua. The correlation itself does not
necessarily imply a direct effect of one variable on an-
other. However, the feeding habits of M. inconspicua
and the amphipods suggest that this correlation may be
significant.

That Ampelisca lies on its dorsum sweeping its anten-
nae over the surface, has already been mentioned. If,
incleed, this is an indication that it is a detritus feeder as
were all of the ampcliscids studied by ENEguist (1950),

then the tremendous numbers of A. miller: (roughly
8000 individuals per m’) are significant competitors with
Macoma inconspicua for this food.

In addition to competition for food, the sweeping habit
could conceivably affect the spat fall. SEGERSTRALE
(1957) reported that in an area of the Baltic, where the
amphipod Pontoporeia affinis is abundant, the Macoma
inconspicua population has failed over a number of
years. He suggested that the amphipods may eat young
Macoma as they sink to the bottom after the planktonic
stage.

KaANNEWoRFF (1965) found that Ampelisca macro-
cephala feeds on detritus in the field. In his laboratory,
however, the amphipod was observed to create currents
in which Mytilus larvae, copepods and Artemia salina
were carried to the gnathopods and then transferred to
the mouth.

Pontoporeia and Ampelisca macrocephala are about
4 times larger than A. millert. While A. millert may not
be large enough to prey on the spat, the sweeping motion
of the antennae could prove a physical barrier to the
spat.

At Icast 3 possibilities of competition between Macoma
and Ampelisca millert exist: (1) the use of detritus as
food, (2) predation by the amphipod on the spat, and
(3) physical interference with spat fall.

SUMMARY

1. The distribution of invertebrates on and in a mud flat
of central San Francisco Bay was studied.

2. Near the 1.0 ft level, one of the critical tide levels
cited by Doty (1946), the number of animals in the mud
flat increased.

3. Two communities were found on the mud flat. Maco-
ma inconspicua was the most abundant organism in the
upper zone. This community gradually gave way to one
dominated by the amphipod Ampelisca millert.

4. The distribution of Macoma inconspicua appears to
be determined by currents which affect sediment settle-
ment and by degree of shelter. Sediment and shelter
favor the amphipod which competitively excludes Maco-
ma inconspicua.

ACKNOWLEDGMENTS

For their advice and encouragement I would like to
thank the members of my committee: Dr. James Ny-
bakken, Dr. Howard L. Cogswell and Dr. Robert Main.

Vole its No:3

I am grateful to Dr. Olga Hartman for her help with
the identification of the polychaetes. To Dr. J. L. Barnard
I owe thanks for his lesson in the identification of an
amphipod.

I would also like to thank the Oliver Brothers’ Salt
Company for the use of its tide lands.

LITERATURE CITED

ALLEN, J. FRANCES

1954. ‘The influence of bottom sediments on the distribution
of five species of bivalves in the Little Annemessex River,
Chesapeake Bay. Nautilus 68 (2): 56-65

Apuin, J. A.

1967. Biological survey of San Francisco Bay 1963 - 1966.
Mar. Resources Oper. Ref. No. 67-4, Mar. Operat. Lab.,
Menlo Park, Calif:

BarRnarD, J. LAURENS

1967. | New species and records of Pacific Ampeliscidae (Crus-

tacea : Amphipoda). Proc. U.S. Nat. Mus. 121: 1-20
BEANLAND, FE. Louise

1940. Sand and mud communities in the Dovey estuary.

Journ. Mar. Biol. Assoc. U. K. 24: 589 - 611
Conomos, T. Joun

1963. Geologic aspects of the Recent sediments of South San
Francisco Bay. Master of Sci. Thesis, San Jose State
Coll., San Jose, Calif.

Dexter, Ratpu H.

1947. The marine communities of a tidal inlet. Ecolog.
Monogr. 17: 236 - 295
Doty, M.S.

1946. Critical tide factors that are correlated with the vertical
distribution of marine algae and other organisms along the
Pacific coast. Ecology 27: 315 - 328

ENEQuIST, PAUL

1950. Studies on the soft-bottom amphipods of the Skagerak.

Zool. Bidr. Uppsala 28: 297 - 492
Firice, Francis P.

1954a. An ecological survey of the Castro Creek area in San
Pablo Bay. Wasmann Journ. Biol. 12: 1 - 24

1954b. A study of some factors affecting the bottom fauna of

a portion of the San Francisco Bay estuary.
Journ. Biol. 12: 257 - 293

Wasmann

1958. Invertebrates from the estuarine portion of San Fran-
cisco Bay and some factors influencing their distribution.
Wasmann Journ. Biol. 16: 159 - 211

Forp, E.

1923. Animal communities of the level sea-bottom in the
waters adjacent to Plymouth. Journ. Mar. Biol. Assoc. U.
K. 13: 164 - 224

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Page 233

Fraser, JAMES H.

1932. Observations on the fauna and constituents of an
estuarine mud in a polluted area. Journ. Mar. Biol. Assoc.

U.K.
Goopuart, C. B.
1941. The ecology of the amphipods in a small estuary in
Hampshire. Journ. Anim. Ecol. 10: 306 - 322
Gram, RALPH

1966. The marine geology of the Recent sediments of central
San Francisco Bay. Master of Sci. Thesis, San Jose State
Coll., San Jose, Calif.

Hart, T. J.
1930. Preliminary notes on the bionomics of the amphipod
Corophium volutator. Journ. Mar. Biol. Assoc. U.K. 16:
761 - 789
Houmeg, N. A.

1949. The fauna of sand and mud banks near the mouth of
the Exe estuary. Journ. Mar. Biol. Assoc. U.K. 28: 189
to 237

Jones, MerepitH L.
1961. A quantitative evaluation of the benthic fauna off
Point Richmond, California. Univ. Calif. Publ. Zool. 67:
219 - 320

KANNEWORFF, EBBE
1965. — Life cycle, food and growth of the amphipod Am pelis-
ca macrocephala LitjEBorc from the Oresund. Ophelia
2: 305 - 318
Kitcuine, J. A., Sytvia J. Litty, SHeita M. Lopce, J. F SLoane,
R. BassInpALeE & FE J. Esiinc

1952. The ecology of Lough Ine rapids with special reference
to water currents. III. The effect of current on other environ-
mental conditions. Journ. Ecol. 40: 179 - 202

McCarty, JAMEs C., R.A. WacGNER & ERMAN A. PEARSON

1962. An investigation of water and sediment quality and pol-
lutional characteristics of three areas in San Francisco Bay,
1960 - 61. Berkeley: Sanit. Eng. Res. Lab., Univ. Calif.

McNutty, J. KNEELAND, Ropert C. Work & Hirary B. Moore

1962. Some relationships between the infauna of the level
bottom and the sediment in south Florida. Bull. Mar. Sci.
Gulf and Carib. 12: 322 - 332

Miier, Rospert C., Witttam D. RaAMacE « Epcar L. LAziEer

1928. A study of physical and chemical conditions in San
Francisco Bay, especially in relation to the tides. Univ.
Calif. Publ. Zool. 31: 201 - 267

PacKarp, Earv L.
1918a. Molluscan fauna from San Francisco Bay. Univ.
Calif. Publ. Zool. 14: 199 - 452
1918b. A quantitative analysis of the molluscan fauna of San
Francisco Bay. Univ. Calif. Publ. Zool. 18: 299 - 336
Rees, Couin B.

1940. A preliminary study of the ecology of a mud flat.
Journ. Mar. Biol. Assoc. U.K. 24: 189 - 199

Page 234

ROsEWATER, JOSEPH
1968. Itinerary of the voyage of H.M.S. Blossom, 1825 to
1828. The Veliger 10(4) : 350 - 352 (1 April 1968)
SANDERS, Howarp L.
1958. Benthic studies in Buzzards Bay. I. Animal sediment
relationships. Limnol. & Oceanogr. 3: 243 - 258
SEGERSTRALE, SVEN G,

1957. Baltic Sea.
ecology. J. W. Hedgpeth (ed.)
1: 751 - 800

SPoonerR,.G. M. « Hitary B. Moore

In Treatise on marine ecology and paleo-
Geol. Soc. Amer. Mem. 67,

1940. The ecology of the Tamar estuary. VI. An account of
the macrofauna of the intertidal muds. Journ. Mar. Biol.
Assoc. U.K. 24: 283 - 330

Srorrs, Puitip N., Ropert BE. SELLEcK & ERMAN A. PEARSON

1964. A comprehensive study of San Francisco Bay 1962-1963.
Berkeley: Sanit. Eng. Res. Lab., Univ. Calif.

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Vol. 11; No. 3

Sumner, Francis B., Gzorce D. LoupERBAcK, WALDO SCHMITT &
Epwarp C. JoHNSTON
1914. A report upon the physical conditions in San Francisco
Bay based upon the operations of the United States Fisheries
Steamer “Albatross” during the years 1912 and 1913.
Univ. Calif. Publ. Zool. 14: 1 - 198

THorRSON, GUNNAR
1957. | Bottom communities. In Treatise on marine ecology
and paleoecology. J. W. Hedgpeth (ed.) Geol. Soc. Amer.
Mem. 67, 1: 461 - 534
VASSALLO, MariLyn T.
in press. A report on the sea anemone Diadumene leucolena
(VERRILL)
YoncE, CuHarLes Maurice

1949. On the structure and adaptations of the Tellinacea,
deposit-feeding Eulamellibranchia. Phil. Trans. Roy. Soc
London, B. 234: 29 - 76

Vol. 11; No. 3

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Page 235

Spawning of the American Oyster, Crassostrea virginica,

at Extreme pH Levels

BY

ANTHONY CALABRESE

AND

HARRY C. DAVIS

Bureau of Commercial Fisheries, Biological Laboratory, Milford, Connecticut 06460

(Plate 45)

Tue AMERICAN OYSTER, Crassostrea virginica (GMELIN,
1791), is the most important commercial mollusk of the
Atlantic Coast and the Gulf of Mexico. Of the various
interacting biological, physical, and chemical factors that
affect oysters in these waters, pH has received less atten-
tion than any other major factor. PryTHERCH (1928),
who measured pH at several stations in Milford Harbor
and the Milford area of Long Island Sound, found that
it ranged from 7.2 to 8.4 during the day. He observed
that oysters in Milford Harbor spawned at pH 7.8 and
8.2 and concluded that low pH inhibited oyster spawning
and that oysters in Milford Harbor spawned at high tide
because this was the only tidal stage at which the pH was
between 7.8 and 8.2.

In laboratory experiments LoosANOFF & ‘TOMMERS
(1947) determined that adult oysters kept in water ad-
justed to pH 4.25 remained open, on an average, 76% of
the time, but pumped only 10% as much water as did
control oysters kept in water of pH 7.75. Oysters kept
at pH 6.75 and 7.00 initially pumped more vigorously
than the controls, but the rate of pumping later decreased
to less than that of the controls.

Although the pH of sea water usually ranges from
7.5 to 8.5, the pH in tidepools, bays, and estuaries may
decrease to 7.0 or lower due to dilution and production
of H.-S (Sverprup et al, 1942). These inshore areas con-
stitute a major portion of the habitat of oysters, and
Davis & CaLaBrese (1964) suggested that these regions
may also be exceedingly important as nursery grounds for
larval stages. Since oyster larvae must, at times, encoun-
ter a wide range in pH in their natural habitat, it is
possible that success or failure of recruitment in some
areas may be determined by variations in pH.

In previous studies CALABRESE & Davis (1966) found
that normal development from the fertilized egg to the
48-hour, straight-hinge larval stage took place within the
pH range from 6.75 to 8.75. In experiments with oyster
larvae, which were terminated after 12 days at experi-
mental pH levels, more than 68% of the larvae survived
within the pH range from 6.25 to 8.75, and the lowest
pH limit for survival was 6.00. The optimum pH for
larval growth was from 8.25 to 8.50, although growth
was good from 6.75 to 8.75. At pH 9.00 to 9.50 the per-
centage of eggs that developed normally, the percentage
of larvae that survived, and the percentage increase in
mean length decreased rapidly.

The present experiments were initiated to determine
the minimum and maximum pH at which oysters will
spawn and to determine the length of time that eggs and
sperm remain viable at these extreme pH levels. The
methods used at this laboratory for conditioning oysters
and obtaining gametes have been described previously
by LoosanorF & Davis (1963). In these experiments we
placed oysters in normal laboratory sea water (pH ap-
proximately 7.8) and sea water adjusted to the desired
pH level with either HCl or NaOH. We attempted to
induce spawning by either of two methods: (1) by
raising the temperature of the sea water to approximately
29° C and simultaneously adding to the sea water small
quantities of sperm suspension made from gonadal mate-
rial of ripe male oysters (combined thermal and chemi-
cal stimulation), and (2) by thermal stimulation only
(Plate 45).

We concluded from these experiments that the mini-
mum and maximum pH levels at which American oysters
will spawn are 6.0 and 10.0, respectively. A summary of

Page 236

the data collected at these pH levels is presented in
Table 1.

Table 1

Spawning of American oysters at extreme pH levels
as compared to spawning at normal pH

Group 1 Group 2
= a
2 as a are
S Es Sa Bees
3
a 2S aie
Number of oysters used 155 114 118 100
Number spawned 23. « 58 28 46
Percentage spawned 14.8 50.9 23.7 46.0
Number of males spawned 21 38 19 27
Number of females spawned 2! 20 9 19

' pH when spawning was observed was 6.4

As indicated in Table 1, the percentage of oysters that
spawned at pH 6.0 and 10.0 was considerably lower than
the percentage that spawned at normal pH of sea water.
In all tests male oysters spawned more readily than fe-
males and at pH 6.0 it was most difficult to induce females
to spawn.

To determine the viability of eggs and sperm from
oysters spawned at pH 6.0 and 10.0 we induced spawn-
ing by thermal stimulation only. As male or female gam-
etes were released, we attempted fertilization with male
or female gametes released from spawners at normal pH.
Eggs or sperm released at pH 6.0 or 10.0 were removed
after 15, 30, 45, 60, 90, 120, 150, 180, 240, and 360
minutes of exposure to those pH levels and we tried ferti-
lization with normal eggs or sperm to test viability. Al-
though the data are insufficient to determine precisely
how long eggs and sperm can tolerate these extreme pH
levels, it is obvious that oyster eggs and sperm released at
pH 6.0 and 10.0 lose their viability rapidly, and are no
longer viable after 2 to 4 hours. GaLttsorr (1964) re-
ported that fertilizability of oyster eggs kept under nor-
mal sea-water conditions decreased to about 20% in 10
hours, and that only a few eggs fertilized after 10 and
24 hours cleaved normally. He also stated that sperm,
kept at room tempcrature in a dilute suspension, lost their
fertilizing ability within 4 to 5 hours. In consideration of
this finding we tried fertilization with 15- and 210-min-
ute-old gametes from a male and female that spawned at
normal pH. Development to the straight-hinge stage with
the 15-minute-old gametes was 100% ; with the 210- min-
ute-old gametes development was only 77%. At this time

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Vol. 11; No. 3

we also tried fertilization with 15- and 210-minute-old
gametes from a male that spawned at pH 6.0 and the
same female that spawned at normal pH. Development
to the straight-hinge stage with the 15-minute-old gam-
etes was 100%; with the 210-minute-old gametes devel-
opment was only 48%. The lowered viability of eggs and
sperm in these experiments was, therefore, due to a com-
bination of pH and aging.

The spawning season of the American oyster in Long
Island Sound extends from late June to early September
(LoosanorF, 1965). During this time excessive changes
of pH caused by siltation or pollution could cause a
failure of recruitment of oysters into the population.

SUMMARY

The minimum and maximum pH levels at which the
American oyster, Crassostrea virginica, will spawn are
6.0 and 10.0, respectively. Oyster eggs and sperm released
at pH 6.0 and 10.0 lose their viability rapidly within 2
to 4 hours. Lowered viability was due, however, to a
combination of pH and aging.

LITERATURE CITED

CaLaABRESE, ANTHONY & Harry Cari Davis
1966. _ The pH tolerance of embryos and larvae of Mercenaria
mercenaria and Crassostrea virginica. Biol. Bull. 131 (3):

427 - 436 (December 1966)
Davis, Harry Cart & ANTHONY CALABRESE
1964. Combined effects of temperature and salinity on de-

velopment of eggs and growth of larvae of M. mercenaria
and C. virginica. Fish. Bull. U.S. Fish and Wildlife Serv.
63: 643 - 655

Ga.tsorF, PauL SIMoNn

1964. The American oyster Crassostrea virginica GMELIN.
Fish. Bull. U.S. Fish and Wildlife Serv. 64: 480 pp.

LoosaNorF, Victor Lyon

1965. | Gonad development and discharge of spawn in oysters

of Long Island Sound. Biol. Bull. 129 (3): 546 - 561
(December 1965)
LoosanorF, Victor Lyon « Harry Cart Davis

1963. Rearing of bivalve mollusks In: Advances in Marine

Biology, EF S. Russet (ed.), Acad. Press, London 1: 1 - 136
Loosanorr, Victor Lyon « FRANCES DorEetTa TOMMERS

1947. Effect of low pH upon rate of water pumping of

oysters, Ostrea virginica. Anat. Record 99: 112 - 113
PryTHERCH, HERBERT F.

1928. Investigation of the physical conditions controlling
spawning of oysters and the occurrence, distribution, and
setting of oyster larvae in Milford Harbor, Connecticut.
Bull. U.S. Bur. Fish. 44: 429 - 503

Sverprup, H. U., Martin W. JoHNson & RicHarp H. FLeminc

1942. The Oceans, their physics, chemistry and general biol-

ogy. Prentice-Hall, Inc., New York, 1087 pp.

THE VELIGER, Vol. 11, No. 3 [CALABRESE] Plate 45

Inducing spawning of oysters by immersing dishes of sea water
containing oysters in warm water on spawning table and
adding sperm suspension

Vol. 11; No. 3

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Page 237

Invertebrates Taken in Six Year Trawl Study

in Santa Monica Bay

JOHN G. CARLISLE, Jr.

Marine Resources Operations, California Department of Fish and Game, California State Visherics Laboratory

Terminal Island, California 90731

INTRODUCTION

In 1958 THE BuREAU OF SANITATION, City of Los Ange-
les and the California Department of Fish and Game
entered into an informal agreement as part of a surveil-
lance program for Santa Monica Bay. The purpose of
this study was to evaluate the effects on the marine en-
vironment, as measured by bottomfish catches, of waste
discharges from the Hyperion Treatment Plant, the large
scale sewage waste disposal plant for the City of Los
Angeles. This plant discharged effluent varying from 261
to 283 million gallons of waste a day during the 6 years
of the study. In addition, solids discharged into the ocean
by the sludge line ranged from 130 to 156 tons per day.
The effluent line is 5 miles long, the sludge line 7.

The biological work was done by Department of Fish
and Game biologists; the City of Los Angeles provided
the boat, equipment, and crew. In the past, all attempts
to prove any long-term and subtle influence of major
waste discharges on an open coastal biotope have failed
(Lupwic & Onopera, 1964). A large amount of data
was gathered during 6 years of trawling from 1958
through 1963. The analysis of the data pertaining to
bottomfish populations is the subject of another report
(CaruisLeE, in press). The present report deals with the
large quantity of invertebrates, primarily mollusks, taken
incidental to the catch of bottomfish.

METHODS

Trawling was conducted from the Prowler, a 65-foot
converted salmon troller used in the Santa Monica Bay
monitoring program. Trawling equipment included a
winch deriving its power from the main engine, and an
A-frame on the fantail through which a single cable led
over a block to the trawl net. A 25-foot cable bridle was

attached to a pair of otter boards fixed to short lines on
the wings of the net. The boards were 12- by 18-inch
steel-reinforced plywood weighted heavily along the bot-
tom. The net was a 24-foot semi-balloon or tri-net with a
body of 14-inch mesh, number 18 twine. The liner of the
bag was of 4-inch mesh synthetic material.

Thirty-nine regular stations in depths of 60 to 600 fect
were sampled every 3 months. While difficulties in sched-
uling the boat and crew made it impossible to adhere to
the quarterly program perfectly, it was followed as close-
ly as possible.

Each trawl lasted 10 minutes, clocked from the time
the net was on the bottom (and fishing) until the winch
was started and net retrieving began. All fishes were
returned to the California State Fisheries Laboratory for
further study. Invertebrates were counted and returned
to the water or saved for later identification. During the
6 years, 705 net hauls were made.

It was impossible to assess the catch of invertebrates
in the same manner as the bottomfish because the net
was designed to catch fishes and not invertebrates. How-
ever, the large incidental catch of invertebrates was of
considerable interest since it provided information on
population densities, distribution in depth and ecological
associations. No attempt was made to evaluate the cat-
ches as proof of pollution or non-pollution of the area.
Only live individuals were tallied; dead shells were abun-
dant in many hauls but they were not noted since their
distribution could have been influenced by currents or
other factors.

ACKNOWLEDGMENTS

This research was begun and supervised for the first 5
years by John L. Baxter before I took over the project.
The work accomplished and subsequent help on many

Page 238

occasions are gratefully acknowledged. John E. Fitch
was extremely helpful throughout the entire study.

Sincere thanks are due to many in a truly cooperative
project, especially in carrying out the field work. The
following, in particular, must be included: Gordon Chap-
man, William Craig, William Donnelly, Ronald Little,
Dennis Peterson, and Donald Zumwalt. Norman J.
Abramson also gave fully of his time with advice and
help.

I would like to take this opportunity to thank the
following personnel of The Hyperion Treatment Plant
and the Los Angeles Bureau of Sanitation for their con-
sideration through the years: Eugene Nelson, Charles
Gunnerson, Homer Rheinschmidt, Charles Imel, John
Stanton, and Joseph Arai.

Many thanks are also due those who helped in the
task of invertebrate identification, particularly: S. Still-
man Berry, James H. McLean, John Garth, Olga Hart-
man, and Fred Ziesenhenne.

Without all this help the project would have been
truly impossible. One of the pleasures inherent in the
completion of a research project is the opportunity to
express gratitude to all those who made it possible.

LIST or INVERTEBRATES
TAKEN sy TRAWL INCIDENTAL to
FISH CATCHES, 1958 To 1963

Frequency and Depth of Occurrence

PORIFERA

Occasional Depth range: 192 - 426 feet

COELENTERATA

Stylatula sp. sea pens

Frequent Depth range: 60 - 600 feet
Renilla sp. sea pansy

Frequent Depth range: 60 - 104 feet
Sea anemones

Frequent Depth range: 180 - 600 feet
Gorgonians

Occasional Depth range: 120 - 570 feet

ECHIUROIDEA

Listriolobus pelodes VisuErr, 1946
Occasional Depth range: 108 - 354 feet

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Vol. 11; No. 3

SIPUNCULOIDEA
Occasional Depth range: 174 - 192 feet
ANNELIDA
Polychaeta
Aphrodita refulgida Moore, 1910 sea mouse
Frequent Depth range: 60 - 600 feet
Phyllochaetopterus sp. seca mouse
Occasional Depth range: 60 - 300 feet
Hyalinoccia juvenalis Moore, 1911
Occasional Depth range: 186 - 300 feet

Pectinaria californicnsis HARTMAN, 1941
Frequent, extremely large numbers at times
Depth range: 360 - 564 feet

ARTHROPODA

Crustacea
Cancer anthony Stimpson, 1856 yellow crab
Frequent Depth range: 60 - 430 feet
Cancer gracilis DANA, 1852 slender crab
Frequent Depth range: 60 - 570 feet
Loxorhynchus crispatus Stimpson, 1875 decorator crab
Frequent Depth: 60 feet
Loxorhynchus grandis Stimpson, 1857 ___ spider crab
Frequent Depth range: 60 - 126 feet
Pyromaia tuberculata (LockincTon, 1876) spider crab
Frequent Depth range: 60 - 120 feet
Hetcrocrypta occidentalis (DANA, 1854) elbow crab
Occasional Depth range: 60 - 300 feet
Mursia gaudichaudi’ (MtLNE-Epwarps, 1837)

Frequent Depth range: 120 - 600 feet
Randallia ornata (RANDALL, 1839)

Frequent Depth range: 60 - 540 feet
Portunus xantusu (Stimpson, 1860)

Occasional Depth: 60 feet

Portunus sp.
Occasional
Pinnixa longipes (LockincTon, 1877)
Occasional Depth range: 120 - 420 feet
Pleuroncodes planipes Stimpson, 1860

Depth: 60 feet

Frequent Depth range: 174 - 180 feet
Paralithodes sp.
Occasional Depth not recorded

Lopholithodes foraminatus (Stimpson, 1859)
Occasional Depth range: 516 - 540 feet
Panulirus interruptus (RANDALL, 1839)
Occasional Depth: 60 feet

Vol. 11; No. 3

Galathea californiensis BENEDICT, 1904
Occasional Depth range: 276 - 378 feet
Crago nigromaculata (LockincTon, 1877)

Frequent Depth range: 60 - 570 feet
Crago alaskensis (RaTHBUN, 1902)
Occasional Depth not recorded

Sicyonia ingentis BURKENROAD, 1938

Frequent Depth range: 120 - 600 feet
Pandalus jordani RatuBun, 1902

Frequent Depth range: 564 - 600 feet
Spirontocaris sp.

Frequent Depth range: 60 - 600 feet

Penaeus californicus Kincstey, 1878
Occasional Depth not recorded
Pseudosquilla bigelowi (Scumirt, 1940)

Occasional Depth range: 60 - 222 feet
MOLLUSCA
Cephalopoda
Rossia pacifica Berry, 1911 squid
Frequent Depth range: 120 - 600 feet
Loligo opalescens Berry, 1911 squid
Frequent Depth range: 60 - 600 feet
Octopus spp. octopus
Frequent Depth range: 60 - 600 feet
Tectibranchiata sea hares
Frequent Depth range: 60 - 600 feet
Nudibranchiata sea slug
Frequent Depth range: 60 - 600 feet
Gastropoda

Diodora aspera (EscHSCHOLTZ, 1833)

Occasional Depth range: 180 - 540 feet
Puncturella cucullata (Goutp, 1846)
Occasional Depth range: 240 - 300 feet

Puncturella coopert CARPENTER, 1864

Occasional Depth range: 216 - 466 feet
Crepidula sp. slipper shell
Occasional Depth range: 60 - 246 feet

Acteocina intermedia WILLETT, 1928

Frequent Depth range: 120 - 600 feet
Philene alba Matrox, 1958

Occasional Depth range: 120 - 240 feet
Turcica caffea Gass, 1865

Rare Depth range: 192 - 198 feet
Cidarina cidaris ADAMS, 1864

Rare . Depth range: 252 - 360 feet
Calliostoma tricolor Gasp, 1865

Frequent Depth range: 60 - 558 feet

Calliostoma annulatum (LicutFoot, 1786)
Occasional Depth range: 192 - 300 feet

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Page 239

Calliostoma gloriosum Datu, 1871

Rare Depth: 80 feet
Calliostoma turbinum Dat, 1895

Rare Depth: 80 feet
Calliostoma variegatum CARPENTER, 1864

Rare Depth range: 252 - 324 feet

Solariella peramabilis CARPENTER, 1864

Frequent Depth range: 180 - 300 feet
Sinum scopulosum (Conran, 1849)

Rare Depth range: 60 - 600 feet
Acteon painei Dau, 1903

Rare Depth range: 252 - 300 feet
Acteon punctocaelatus (CARPENTER, 1864)

Rare Depth range: 210 - 240 feet
Polinices altus (Dat, 1909)

Rare Depth: 120 feet
Polinices draconis (Dati, 1903)

Occasional Depth range: 60 - 600 feet
Polinices lewisti (Gouxp, 1847)

Occasional Depth range: 60 - 126 feet

Polinices recluzianus (DESHAYES, 1839)

Frequent Depth range: 120 - 180 feet
Cypraca spadicca Swainson, 1823

Rare Depth: not recorded
Trivia rittert RayMonp, 1903

Rare Depth range: 276 - 300 feet
Turbonilla sp.

Occasional Depth range: 60 - 600 feet
Balcis micans (CARPENTER, 1864)

Occasional Depth range: 276 - 300 feet
Eunaticina oldroydi (Dat, 1897)

Frequent Depth range: 570 - 588 feet
Eulima sp.

Occasional Depth range: 120 - 588 feet

Epitonium bellistriatum (CARPENTER, 1864)

Rare Depth range: 54- 60 feet
Epitonium sp.

Rare Depth: 588 feet
Opalia wroblewsky: Morcu, 1852

Rare Depth range: 276 - 288 feet
Turritella coopert CARPENTER, 1864

Rare Depth: 192 feet
Erato sp.

Occasional Depth range: 120 - 228 feet
Neosimnia catalinensis Berry, 1916

Frequent Depth range: 192 - 450 feet

Bittium interfossa (CARPENTER, 1864)

Rare Depth: 588 feet
Bittium subplanatum Bartscu, 1911

Rare Depth: 180 feet
Bittium sp.

Frequent Depth range: 180 - 600 feet

Page 240

Terebra pedroana Datu, 1908

Occasional Depth range: 60 - 120 feet
Nassarius cooperi (ForBES, 1850)

Rare Depth: 180 feet
Nassarius fossatus (Goutp, 1850)

Occasional Depth: 60 feet

Nassarius insculptus (CARPENTER, 1864)

Frequent Depth range: 180 - 600 feet
Nassarius mendicus (Goutp, 1850)

Occasional Depth: 60 feet
Nassarius perpinguis (Hinps, 1844)

Occasional Depth range: 60 - 192 feet
Olivella baetica CARPENTER, 1864

Occasional Depth range: 60 - 300 feet
Olivella pycna Berry, 1935

Occasional Depth: 60 feet
Cancellaria coopert Gass, 1865

Frequent Depth range: 330 - 588 feet

Cancellaria crawfordiana Dat, 1891

Frequent Depth range: 432 - 600 feet
Mitra idae Mrivint, 1898

Frequent Depth range: 192 - 300 feet
Mitrella carinata (Hinps, 1844)

Rare Depth: 120 feet
Mitrella gausapata (GouLp, 1850)

Frequent Depth range: 180 - 420 feet
Mitrella tuberosa (CARPENTER, 1864)

Rare Depth: 60 feet
Amphissa bicolor Dau, 1892
Rare Depth range: 414 - 420 feet

Amphissa undata (CARPENTER, 1864)

Occasional Depth range: 180 - 540 feet
Shaskyus festivus (Hinps, 1844)

Frequent Depth range: 60 - 564 feet
Fusinus barbarensis (Trask, 1855)

Occasional Depth range: 180 - 228 feet

Pteropurpura macropterus DESHAYES, 1839

Rare Depth range: 66 - 180 feet
Pteropurpura vokesaec EmMERSon, 1964

Occasional Depth range: 60- 72 feet
Trophonopsis bentleyi (Dati, 1908)

Frequent Depth range: 420 - 600 feet
Trophonopsis lasius (Dati, 1919)

Rare Depth range: 240 - 300 feet
Trophonopsis scitulus (Day, 1891)

Rare Depth range: 216 - 300 feet
Trophonopsis triangulatus (CARPENTER, 1864)

Rare Depth range: 180 - 600 feet
Ocenebra barbarensis (Gass, 1865)

Rare Depth: 252 feet
Kelletia kelletu (Forsrs, 1850)

Frequent Depth range: 60 - 564 feet

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Vol. 11; No. 3

Daphnella clathrata Gass, 1865

Rare Depth range: 180 - 432 feet
Forreria belchert (Hinps, 1843)

Rare Depth: 60 feet
Bursa californica (Hinps, 1843)

Frequent Depth range: 60 - 387 feet
Conus californicus Hinps, 1844

Frequent Depth range: 60 - 192 feet

Megasurcula carpenteriana (Gass, 1865)
Frequent Depth range: 60 - 600 feet
Megasurcula stearnsiana (Raymonp, 1904)

Occasional Depth range: 216 - 360 feet
Antiplanes litus Dau, 1919

Rare Depth: 588 feet
Antiplanes perversa (Gass, 1865)

Rare Depth: 588 feet
Antiplanes sp.

Rare Depth range: 432 - 444 feet
Burchia redondoensis (T. Burcu, 1938)

Occasional Depth: 60 feet
Elaeocyma empyrosia (DA, 1899)

Occasional Depth range: 180 - 420 feet

Mangelia arteaga DALL & Bartscu, 1910

Occasional Depth range: 180 - 420 feet
Mangelia beta Datt, 1919

Rare Depth range: 336 - 456 feet
Clathurella crystallina Gass, 1865

Rare Depth range: 576 - 600 feet
Ophiodermella halcyonis (Daux, 1908)

Rare Depth: 120 feet
Pelecypoda
Acila castrensis Hinps, 1843

Frequent Depth range: 360 - 600 feet
Nucula tenuis Montacu, 1808

Occasional Depth range: 180 - 588 feet

Nuculana hamata (CARPENTER, 1864)

Occasional Depth range: 186 - 600 feet
Nuculana acuta (Conrap, 1832)

Occasional Depth range: 408 - 600 feet
Yoldia ensifera Dati, 1897

Frequent Depth range: 336 - 600 feet

Acquipecten latiauratus (ConraAp, 1837)

Occasional Depth range: 60 - 456 feet
Chlamys hastatus SowERBy, 1843
Occasional Depth range: 216 - 228 feet

Lima hemphilli HERTLEIN & STRONG, 1946

Occasional Depth range: 60 - 234 feet
Pecten diegensis Dati, 1898

Frequent Depth range: 60 - 408 feet
Cardita ventricosa GouLp, 1850

Rare Depth: 558 feet

Vol. 11; No. 3

Cyclopecten vancouverensis (WHITEAVES, 1886)

Rare Depth range: 60 - 248 feet
Delectopecten sp.

Occasional Depth range: 180 - 564 feet
Anomia peruviana D’OrBIGNY, 1846 ;

Occasional Depth range: 324 - 330 feet

Amygdalum pallidulum (Dau, 1916)

Frequent Depth range: 180 - 600 feet
Modiolus neglectus Soot-RYEN, 1955

Rare Depth: 180 feet
Solamen columbianum (Datu, 1897)

Occasional Depth: 180 feet
Pandora filosa (CARPENTER, 1864)

Occasional Depth range: 186 - 420 feet
Solemya panamensis Dau, 1908

Occasional Depth range: 252 - 588 feet
Hiatella arctica (Linnarvus, 1767)

Occasional Depth range: 180 - 372 feet
Lyonsia californica Conrap, 1837

Occasional Depth range: 180 - 192 feet
Thracia trapezoides Conran, 1849

Rare Depth range: 330 - 480 feet
Thyasira barbarensis Dati, 1889

Occasional Depth range: 180 - 600 feet
Solen rosaceus CARPENTER, 1864

Occasional Depth: 60 feet
Pscudochama exogyra (Conran, 1837)

Occasional Depth range: 186 - 324 feet
Pseudochama granti StRoNG, 1934

Occasional Depth range: 198 - 306 feet
Cuspidaria apodema Dat1, 1916

Frequent Depth range: 180 - 588 feet
Cuspidaria planetica DALL, 1908

Frequent Depth range: 234 - 600 feet
Cuspidaria californica Dati, 1886

Rare Depth: 336 feet

Nemocardium centifilosum (CARPENTER, 1864)

Frequent Depth range: 120 - 444 feet
Trachycardium quadragenarium (Conran, 1837)

Occasional Depth range: 60 - 192 feet
Lucinoma annulata (ReEve, 1850)

Occasional Depth range: 384 - 420 feet
Lucinoma tenuisculpta (CARPENTER, 1864)

Occasional Depth range: 316 - 318 feet
Compsomyax subdiaphana (CarPENTER, 1864)

Frequent Depth range: 120 - 600 feet
Cooperella subdiaphana (CARPENTER, 1864)

Rare Depth: 120 feet
Gari edentula (Gass, 1868)

Rare Depth: 120 feet
Macoma carlottensis WritrEAvEs, 1880

Frequent Depth range: 342 - 600 feet

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Page 241

Macoma sp.
Rare Depth range: 330 - 480 feet
Tellina arenica HERTLEIN & STRONG, 1949
Occasional Depth range: 300 - 588 feet
Amphineura
Lepidozona retiporosa (CARPENTER, 1864)
Occasional Depth range: 180 - 600 feet
Scaphopoda
Dentalium rectius CARPENTER, 1864

Frequent Depth range: 360 - 600 feet
Dentalium vallicolens RayMonp, 1904

Rare Depth range: 384 - 420 feet
BRACHIOPODA

Glottidia albida (Hinps, 1844)
Rare Depth range: 120 - 174 feet
Terebratalia occidentalis (Dai, 1871)

Occasional Depth range: 216 - 378 fect
Laqueus californicus (Cocu, 1848)
Rare Depth range: 384 - 432 feet
ECHINODERMATA
Asteroidea
Astropecten sp.
Frequent Depth range: 60 - 600 feet

Large numbers at times
Mediaster aequalis Stimpson, 1857

Frequent Depth range: 120 - 432 feet
Astrometis sp.
Occasional Depth: 300 feet

Pisaster brevispinus (Stimpson, 1857)

Frequent Depth range: 60 - 420 feet
Luidia foliolata GruBe, 1866
Occasional Depth range: 60 - 564 feet

Rathbunaster californicus FisHer, 1906

Occasional Depth range: 60 - 240 feet
Ophiuroidea

Frequent Depth range: 114 - 600 feet
Echinoidea

Allocentrotus fragilis Jackson, 1912
Frequent Depth range: 219 - 570 feet
Large numbers at times
Lytechinus anamesus H. L.Criarx, 1912
Frequent Depth range: 174 - 436 fect
Large numbers at times
Strongylocentrotus sp.
Occasional Depth range: 114 - 252 fect
Brisaster townsendi (Acassiz, 1898)
Frequent Depth range: 60 - 540 feet
Large numbers at times

Page 242 THE VELIGER Vol. 11; No. 3

Crinoidea
Florometra perplexa A. H.Ciark, 1907
Frequent Depth range: 60 - 387 feet
Holothuroidea
Stichopus spp.
Frequent Depth range: 60 - 600 feet

Large numbers at times

TUNICATA

Salpa spp.
Frequent Depth range: 60 - 378 fect
Large numbers at times

LITERATURE CITED

CaruisLE, Joun G. Jr.
in press. Results of a six-year trawl study in an area of heavy
waste discharge: Santa Monica Bay, California.
Lupwic, Harvey F « BEN ONopERA

1964. Scientific parameters of marine waste discharge.
pp. 37-49 In: BE. A. Pearson (ed.), Advances in water pollu-
tion research, vol. 3. Pergamon Press, New York, N. Y.; 431 pp.

Vol. 11; No. 3

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Page 243

Pelecypod-Sediment Association in Tomales Bay, California

BY

DON MAURER

Shellfish Laboratory, University of Delaware, Lewes, Delawarc

19958 '

(1 Map; 3 Tables)

INTRODUCTION

THIS STUDY WAS UNDERTAKEN to determine whether
there was an association between distribution, abundance
and size of pelecypods and sediment particle size. Re-
search was conducted during the summers of 1961
through 1963 at the Pacific Marine station in Tomales
Bay, located in Marin County, California between 38°
1400” N Latitude, 122°58’35’" W Longitude and 38°05’
30” N Latitude, 122°49’40’” W Longitude. The species
under study, Tellina buttont Dati, 1900, T: salmonea
(CarPENTER, 1864), Mysella tumida (CarPENTER, 1864)
Lyonsia californica Conrab, 1837, and Transennella tan-
tilla (GouLp, 1852) were selected because they are nu-
merous, widely distributed and occur in a variety of
sediment types in Tomales Bay. Most research on bivalve-
sediment relationships has been chiefly concerned with
species commonly collected by dredges. Such collections
mix different sediment types and tend to confuse natural
pelecypod-substrate associations. The present study dif-
fers from earlier investigations in that samples were col-
lected with a quantitative grab which assures a collection
of bivalves from the sediment in which they live, an
important prerequisite for the determination of pelecy-
pod-sediment associations.

The purpose of the paper is to report the results of
an investigation on bivalve-sediment relationships in
Tomales Bay. Results indicate that distribution and
abundance of all species except Transennella tantilla
appear to be strongly influenced by sediment type. The
average size of the tellinids and Mysella tumida is statis-
tically associated with sediment particle size, whereas a
corresponding significant association did not exist for
Lyonsia californica and Tr. tantilla.

ALLEN (1963) provides the most recent comprehensive
review of studies concerning distribution, abundance and
growth of bivalves related to substrate influences. Associ-

' Contribution No. 55, University of Delaware Marine Laboratories

ation of tellinid species with sediment type has been
reported in California by several authors, Packarp (1918)
observed the occurrence of Tellina buttoni and T. salmo-
nea in mud and sand and of the latter in gravel, and
sand-shell substrates of San Francisco Bay. Reisu (1961)
found T: buttonz in gray clay of a boat harbor in south-
ern California. Ricketts & Carvin (1962) presented a
sketch map which showed a Tellina sand facies in Toma-
les Bay and Jones (1964) stated that species of Tellina
were common benthic mollusks living in sand bottoms off
southern California. As regards the other species, OLp-
royD (1924) and Keep (1935) indicated that Transen-
nella tantilla lives in fine sand and Lyonsia californica
occurs in muds of bays. ReisH (1961) included L. cali-
fornica in a species list of mollusks that were found in
clay. However, these studies were primarily concerned
with reporting the occurrence and relative number of
these species in a general sediment type rather than de-
fining any quantitative aspects of bivalve-sediment re-
lationships. Furthermore, no studies are available on
these species concerning their size related to sediment

type.

MATERIALS anno METHODS

During the months of June, July and August of 1961
through 1963, 142 pelecypod-sediment samples were ob-
tained aboard the R/V Bios Pacifica with a 0.1 square
meter van Veen grab. At each collecting site, a sediment
sample was taken, and the position, depth of water,
volume of grab and a brief description of flora and
fauna were recorded. Samples were washed aboard
through a screen of 1.0mm mesh and the residue was
placed in jars with 5 to 10% formalin. Pelecypods were
sorted, identified, and counted in the laboratory.
Measurements were taken on 3 297 individuals. Maxi-
mum anterior-posterior dimension was selected to repre-
sent the size of clams. This measurement is convenient

Page 244

Sand

; Lawsons Flat
Point iw:

Pacific Ocean

TOMALES BAY
CALIFORNIA

0 5000

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Vol. 11; No. 3

% = Tomales Bay

LEGEND

CS Coarse Sand
MS Medium Sand
SF Fine Sand
SC Silt - Clay

¢ Sample Site

f Double Point

Figure 1

Index Map of Tomales Bay, California.
Sediment Distribution and Sample Sites.

and has been commonly used as an indication of size
(KrisTENSEN, 1957). A vernier caliper was used to
measure Lyonsia californica and larger specimens of tel-
linids. An ocular micrometer and microscope reticle were
used on Mysella tumida, Transennella tantilla and smaller
tellinids. Measurements with all devices had an opera-
tional error of 0.02 mm.

Sediments were analyzed by standard sieve analyses
Pipette analyses were required for silt and clay samples.
A measure of the 20th, 50th, and 80th sediment per-
centiles, the percentage of silt and clay, the sorting coef
ficient (80th minus 20th divided by 50th) were deter-
mined. Among 142 sediment samples 22%, 59%, 4%,
and 14%, were mud, fine, medium, and coarse sand

Vol. 11; No. 3

respectively (Table 1). Sediment types are defined in
this table.

Jounson (1965) has summarized the hydrographic
conditions of the bay. These factors together with the
substrate are important features in the ecology of bi-
valves. It is often difficult to recognize the effect or to
isolate the importance of one ecological factor to the
exclusion of others. In order to assess the effect of sedi-
ment type on pelecypod ecology in Tomales Bay an
intensive study of animal-sediment relationships was also
conducted in White Gulch. White Gulch is a small cove
4.8 km from the mouth of the bay (Map, Figure 1). The
cove is naturally suited for such studies in that it contains
various sediment types and has a relatively narrow range
of hydrographic features compared to the wider range
of conditions throughout the bay. Mertes (1962) has
shown that subtidal conditions in White Gulch are homo-
geneous with regard to water temperature and salinity.
If pelecypod-sediment associations are relatively uni-

Table |

Abundance of Five Species of Bivalves in
Different Sediment Types

Se of Be Se
GB gS oy a ¢ ie
gle Be ce Se &
Seo ef Bo He .
56 LS gO a
AY =K A oA
— —<
Number of
Sediment Samples 31 85 6 20 142
Percent of
Sediment Samples 22 59 4 14
Tellina buttont
Number of Clams 42 1049 53 9 1153
Percent of Clams 4 91 5 0.8
Tellina salmonea
Number of Clams - 207 78 422 707
Percent of Clams ~ 29 11 60
Mysella tumida
Number of Clams 6 308 — - 314
Percent of Clams 2 98 - -
Transennella tantilla
Number of Clams 592 160 33 170 955
Percent of Clams 62 17 3 18
Lyonsia californica
Number of Clams 102 66 = = 1638
Percent of Clams 61 39 = =
Total: 3297

The measurements which define sediment type refer to median
particle size.

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Page 245

form, then the ecological significance of sediment can be
extended to similar associations found in the bay at large.
A more detailed account of methods can be found in
Maurer (1964).

DISTRIBUTION ann ABUNDANCE

In Tomales Bay Tellina buttoni has been collected from
depths of 0.6 to 18.9m and lives mainly in depths of
4.6 to 10.6m. Locally 7: buttont occurs in the northern
part of the bay from Sand Point to Pelican Point. South
of Pelican Point it has been taken occasionally from
holes and channels that lie along the western margin of
the bay. This tellinid lives in sediment that varies from
mud through coarse sand and, in rare cases, pebbles. It
occurs most frequently and in greatest numbers in fine
sediment. Ninety-one percent of JT. buttont were found
in fine sand (Table 1) with 5 to 30% silt and clay. On
the northeastern margin of White Gulch JT. buttonz lives
in sediment that contains as much as 50% silt and clay.
Fine sediment here is due to the gradual decay of eel
grass and trapping of particulate matter by clusters of
Zostera.

Tellina salmonea dwells most frequently in depths be-
tween 3.0 and 7.9m and it has the same depth range as
T. buttoni. Although the salmon tellinid has not been
collected intertidally, subtidal sampling and presence of
many shells washed ashore on a beach north of Sand
Point indicate that a large population lives in or just
beyond the surf zone. Tellina salmonea ranges from the
open ocean south to Pelican Point. Distribution south of
Pelican Point agrees with that of 7? buttoni. Tellina sal-
monea may inhabit fine sand; however, it occurs most
frequently and in greatest numbers in coarse sediment
with less than 5% silt and clay. Sixty percent of T. sal-
monea were found in coarse sand with 29% and 11%
in fine and medium sand respectively (Table 1).

Range of depth of Mysella tumida is from the inter-
tidal zone to 10.6m though this species more often in-
habits depths from 1.2 to 9.1m. Mysella tumida ranges
from ‘Toms Point to south of Pelican Point. South of
Pelican Point distribution seems to be restricted to the
same submarine topography that contains the tellinids.
In this southern portion within the central axis of the
bay, M. tumida is replaced by M. ferruginosa (Dati,
1916.) The former species lives in mud and fine sand and
occasionally in medium sand, but typically inhabits fine
sand with 10 to 30% silt and clay. Ninety-eight percent
of M. tumida were taken from fine sand (Table 1). On
the other hand, M. ferruginosa lives characteristically in
mud with 70 to 100% silt and clay and was infrequently

Page 246

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Vol. 11; No. 3

Table 2

Pelecypod and Sediment Measurements

Median Particle Size

Average Clam Size
in Relation to

Spccies Clam Size in mm mm or % Silt & Clay Median Particle Size

a © = o Sts oe = g
5 > 3 > 54 5 &
4 < a < Au 7) (e)
Tellina buttoni 1.9-18.5 6.5 0.063 — 0.830 0.165 38 < 0.154 39)
38 > 0.154 Wee,
Tellina salmonea 2.0 - 18.7 9.4 0.113 — 2.090 0.634 19 < 0.370 6.0
19 > 0.370 11.7
Mysella tumida 16- 38 2.7 0.052 - 0.190 0.132 18 < 0.122 2.8
18 > 0.122 2.6
Transennella tantilla 15- 9.4 4.3 100% to pebbles 43.1% 18 < OMI 4.4
18 > 0.177 4.3

Lyonsia californica 4.5 — 38.5 13.7 10.7% — 98.4% 86.5% 19 - -

found in sand. In White Gulch M. tumida is numerous in
Zostera beds and occurs often with Tellina buttoni.

Lyonsia californica was collected from depths of 1.2 to
13,7 m and is taken more commonly from 3.0 to 9.1m. It
ranges from just north of White Gulch to south of Double
Point. This bivalve has been obtained almost exclusively
from mud and fine sand, and when it occurs in White
Gulch, it is collected from very fine sand. It lives essen-
tially in sediment with 80 to 100% silt and clay and is
characteristic of the southern two-thirds of the bay or
of more northern localities where high concentrations
of silt and clay accumulate in protected coves. Sixty-one
percent and 39% of L. californica were collected from
mud and fine sand respectively (Table 1). Among the 5
species it shows the most restricted occurrence in fine
sediment.

Transennella tantilla ranges from intertidal locations to
the deepest (19.9 m) holes in the bay. The species may
occur intertidally in great numbers at Lawsons Flat and
White Gulch, although its occurrence is sporadic and
seems to fluctuate like populations of Donax. Subtidally
it generally inhabits depths from 3.0 to 9.1 m. Contrary
to the distribution and abundance of the other 4 species
Tr. tantilla is one of the most ubiquitous clams in the bay.
It was found recently in a rocky intertidal area on the
ocean side of Tomales Point and its range extends south
of Double Point within the bay. This venerid occurs in
channels and shoals from Hog Island northwards to Law-
sons Flat and throughout the White Gulch area. Its great-
est numbers and frequency are seen in mud. Sixty-two
percent of Tr. tantilla were gathered from mud, while
18%, 17%, and 3% were collected from coarse sand, fine

sand, and medium sand respectively (Table 1). In White
Gulch, Tr. tantilla occurs throughout the range of sedi-
ment types.

SIZE ANALYSIS

In addition to distribution and abundance another pele-
cypod-sediment relationship was recognized. From Table
2 it can be inferred that there is some association between
the size of tellinids and sediment particle size. For
example, the average size of 19 samples of Tellina sal-
monea from sediment with a median size less than 0.370
mm is 6.0mm, whereas the average size of 19 samples
from sediment with median size greater than 0.370mm
is 11.7 mm.

To determine whether these relationships were signifi-
cant, a hypothesis was posed and tested statistically. ‘The
hypothesis, which defines an association between a de-
pendent and independent variable, was that pelecypod
size is associated with sediment particle size. Since bivalve
and sediment sizes may not assume a bivariate normal
distribution a nonparametric statistic, the Kendall Cor-
relation Coefficient, was applied. HrepcpetH (1957)
cautioned “that many of the data of ecology are non-
parametric and hence not amenable to parametric tests,...’
According to SteceL (1956) the Kendall Correlation
Coefficient does not make any assumptions concerning
the distributional function. For each sample an average
bivalve size was correlated with a sediment measurement
(20th, 50th, and 80th sediment percentiles, sorting co-
efficient, percent of silt and clay) and depth of water.
From this a correlation coefficient was obtained and its

Vol. 11; No. 3

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Page 247

probability was determined. Level of significance is 0.05
for all analyses.

If size-frequency distributions are bimodal or polymod-
al and standard deviations are high, average size can be
misleading as an estimate of central tendency. Histograms
of bivalve size indicated that, in most samples, variance
was relatively low and that there were only a few irregu-

lar distributions. As a check on the use of average size
the median pelecypod size of some samples was correlated
with a sediment particle measure. Results were compa-
rable to those analyses in which the average clam size was
used. However, average size was preferred because:

(1) samples were generally large enough not to be un-
duly influenced by extreme values; (2) average size

Table 3

Summary of Statistical Analyses

Sediment Percentiles

20th
Tellina buttoni
1 White Gulch
Correlation Coefficient 0.394 0.394
Probability 0.038*
Samples (Specimens) 62(828)
2 Tomales Bay
Correlation Coefficient 0.667 0.538
Probability 0.001+
Samples (Specimens) 13(241)
Tellina salmonea
3 White Gulch
Correlation Coefficient 0.286 0.786
Probability 0.199-
Samples (Specimens) 16(144)
4 Tomales Bay
Correlation Coefficient 0.636 0.673
Probability 0.003+*
Samples (Specimens) 22 (563)
Transennella tantilla
5 White Gulch
Correlation Coefficient 0.238 0.095
Probability 0.281-

Samples (Specimens)
6 Tomales Bay

Correlation Coefficient 0.143 0.107
Probability 0.360-

Samples (Specimens) 8 (266)

Mysella tumida

7 White Gulch

Correlation Coefficient -0.697
Probability 0.001+

Samples (Specimens) 36 (297)

Lyonsia californica

8 Tomales Bay
Correlation Coefficient
Probability

Samples (Specimens)

0.038*
IBID

0.005*
IBID

0.003*

0.002*
IBID IBID IBID IBID IBID

0.430-
13(97) IBID

0.406-

-0.515
0.010*

a
o)
on Fu
Ss =] Ss
cee Sah vide
80th ao gz =A
0.424 0.348 -0.576 0.394
0.027* 0.058 0.005 ~—-0.038+
IBID IBID 58(870) 62(828)
0.461 -0.460 -0.187 0.026
0.014" 0.026 0.176- 0.452-
IBID IBID 14(283) 13(241)
0.643 0.571 -0.429 0.357
0.016 0.031* 0.089- _—0.138-

IBID IBID IBID IBID

0.636 0.418
0.003* 0.037*

—0.163 0.079
0.245- 0.500-

0.333 0.333 0.143 0.238
0.1917 0.191- 0.386- 0.281-
IBID IBID IBID IBID

0.286 -0.071 0.030 0.152

0.199 0.452- 0.448 0.248-
IBID IBID 23(858) 23(858)
0.273 0.303 0.227. 0.500
O03 O08 — OSI OOM
IBID IBID 37(314) 36(297)
0.398 0.444

0.090- —_0.060-

19(168) 19(168)

+ Statistically significant at 0.05
- Not statistically significant at 0.05

Page 248

THE VELIGER

Vol. 11; No. 3

of large numbers of specimens is easier to calculate than
median or mode which require additional manipulation
of raw data to define; (3) small samples contain in-
sufficient entries to form clusters and so the mode is
indeterminate. In such cases the mode can not be used
and unless the average is applied, information may be
lost.

For Tellina buttoni, T. salmonea and Transennella tan-
tilla the null hypothesis is that there is no association
between increasing pelecypod size and increasing sedi-
ment particle size, whereas the null hypothesis for Mysella
tumida and Lyonsia californica is that there is no associa-
tion between decreasing pelecypod size and increasing
sediment particle size.

Table 3 indicates that the average size of Tellina but-
toni in the bay and White Gulch is, in general, signifi-
cantly associated with sediment particle size. In White
Gulch the highest correlation is the 80th sediment per-
centile and the highest correlation in the bay is the 20th
sediment percentile. For T: salmonea a significant associ-
ation was found in the bay and White Gulch and its
highest correlation is the 50th sediment percentile in both
areas. The average size of Mysella tumida is also signifi-
cantly associated with particle size. This species was not
collected in sufficient numbers throughout the bay to be
compared statistically with samples from White Gulch
and therefore analyses were only made for samples from
the latter area (Table 3). In contrast to the tellinids, the
correlation of M. tumida is significant in a negative di-
rection. With Transennella tantilla the average size is not
significantly associated with particle size. Finally, since
Lyonsia californica occurs almost exclusively in fine sedi-
ment its average size was correlated only with percent of
silt and clay. The correlation coefficient was not signifi-
cant.

DISCUSSION

In the northern part of the bay, Sand Point to White
Gulch, sediment type can be characterized as a sandy
substrate, with medium and coarse sand in the channels
and off headlands, and fine sand in protected coves and
shoals. The stretch from White Gulch to Pelican Point
represents a transitional area as sand size becomes finer
and increases in organic material content. In the transi-
tion area the occurrence of all five species overlaps.
Pelican Point affords an excellent physiographic boundary
for distribution patterns as it marks the beginning of
the “mud line” or local sedimentary threshold in Tomales
Bay. South of this point the substrate contains 70 to 100%

silt and clay. Small patches of coarse sand and pebbles
may occur off headlands, but these are usually confined
to the western shore. Occasional beds of fragmented
Ostrea lurida shells are present between Marshall and
Double Point. Based on intensive sediment sampling it
was found that fine sand actually extends in tongues
slightly farther south than Pelican Point on both sides of
the bay (personal communication, Dr. C. C. Daetwyler).

The distribution of the species under study and the
majority of subtidal pelecypods in the bay responds in
a very sensitive fashion to the mud line. For example,
the tellinids and Mysella tumida range essentially from
the northern part through the transitional area and stop
abruptly at the mud line. Tellina buttoni occurs in a
wider range of sediment than T. salmonea. The former is
best represented in fine sand with some organic material,
while the latter finds its maximum development in clean,
coarse sand. A few specimens of Lyonsia californica were
found in White Gulch, yet it is almost exclusively restric-
ted to sediment (70 to 100% silt and clay) south of
Pelican Point. Distribution and abundance of Transen-
nella tantilla is an exception as it appears to be uninflu-
enced by sediment type.

Concerning bivalve size and sediment size only the
tellinids and Mysella tumida show any significant associ-
ations. In general, as sediment particle size increases, size
of the tellinids living in those sediments increases. This
association is so marked that it can be observed in the
field for both tellinids. Not so with M. tumida, for the
difference in size between this mollusk in finer and coarser
sediment is too small for detection in the field. Until
statistical analyses were made it was unknown that the
size of M. tumida decreases as particle size increases.

It should be emphasized for at least the tellinids and
Transennella tantilla that sediment associations in White
Gulch are consistent with those found throughout the bay.
If, as previously suggested, pelecypod-sediment associa-
tions exist where hydrographic conditions are relatively
uniform, then the ecological importance of sediment to
similar bivalve associations in the bay at large can be
confidently asserted. In passing, the largest forms and
greatest numbers of Tellina salmonea occur north of
Sand Point in pebbles and shell-sand and are commonly
twice the size of individuals in medium and coarse sand
south of the point. Here the influence of the ocean is
unmistakable. Nevertheless, samples collected in close
proximity to that area were shown statistically to be
drawn from two different populations. Particle size was
the striking environmental factor that differed. Although
there were insufficient numbers of Mysella tumida and
Lyonsia californica to compare sediment associations from

Vol. 11; No. 3

THE VELIGER

Page 249

White Gulch to the bay, both species were obviously
best developed in a specific sediment type, fine sand for
the former and mud for the latter.

Work by Swan (1952), Pratr (1953), Pratr &
CamMpBELL (1956), Weis (1957), and Pearce (1965)
with other bivalves support the view that there can be
causal relationship between bivalve distribution, abun-
dance, size and sediment type. Maurer (1967a) was
concerned with the biological significance of sediment to
pelecypods. Working with the same species except Lyon-
sia californica, Maurer, (1967 a, 1967 b, 1967 c)_ pro-
vided experimental evidence to indicate that the effect
of turbidity and sediment as a food source were impor-
tant factors influencing bivalve-sediment associations.
The writer submits that sediment type may be a limiting
factor in determining the distribution and abundance of
Tellina buttoni, T: salmonea, Mysella tumida, and L.
californica in Tomales Bay. Moreover, the size of the
tellinids and Mysella tumida appears to be influenced by
sediment. No explanation for the significant negative
correlation coefficient obtained with M. tumida is offered
at the present, and no noteworthy bivalve-sediment asso-
ciations were recognized with Tr. tantilla. Although the
effect of some important life-cycle aspects (life span,
reproductive periods, size of settling larvae, larval sub-
strate preference) remain to be studied in order to more
completely evaluate the proposed sediment relationships,
the writer asserts that the broad outline and definition
of the pelecypod-sediment associations are valid.

LITERATURE CITED

ALLEN, JoHN A.

1963. Ecology and functional morphology of molluscs, in
Harotp Barnes (ed.), Oceanography and marine biology, an
annual review. 1: 253 - 288. | George Allen and Unwin, Ltd.,
London

HepcpPETH, JoEL W.

1957. | Concepts of marine ecology. Jn: Treatise on marine
ecology and paleoecology. J. W. Hedgpeth, ed. Geol. Soc.
Amer. Mem. 67 (1): 29-52

JouNson, Ratpu G.

1965. _Pelecypod death assemblages in Tomales Bay, Califor-

nia. Journ. Paleon. 39 (1): 80-85
Jones, G. FE

1964. The distribution and abundance of subtidal benthic
Mollusca on the mainland shelf of southern California.
Malacologia 2 (1): 43-68

Keen, A. Myra

1963. Marine molluscan genera of western North America:
An illustrated key. Stanford Univ. Press; 126 pp.; text
figs. not separately numbcred (14 February 1963)

KEEP, JosIAH
1935, | West coast shells. Rev. by Josuua L. Baty, Jr.
Stanford Univ. Press, Stanford, Calif. pp. 1 - 350
KRISTENSEN, INGVAR
1957. Differences in density and growth in a cockle population
in the Dutch Wadden Sea. Arch. Neerl. Zool. 12: 351 - 454
Maurer, Don
1964. Pelecypod size and sediment size.
tion, Univ. of Chicago, Library, pp. 1 - 98

Ph. D. disserta-

1967 a. Filtering experiments on marine pelecypods from Toma-
les Bay, California The Veliger 9 (3): 305 - 309
(1 January 1967)
1967b. Burial experiments on marine pelecypods from Tomales
Bay, California. The Veliger 9 (4): 376 - 381
(1 April 1967)

1967. Mode of feeding and diet, and synthesis of studies on
marine pelecypods from Tomales Bay, California. The
Veliger 10 (1): 72-76 (1 July 1967)

MertTEs, Davip

1962. Summary of various salinity determinations from To-
males Bay during the periods from Dec. 3, 1960 to Jan. 27,
1962. Unpubl. Rep. Pacific Marine Station

OLprRoyp, IpA SHEPARD

1924. ‘The marine shells of the west coast of North America.

Stanford Univ. Publ. Geol. Sci., 1: 1 - 247; plts. 1-57
Packarp, Ear L.

1918. A quantitative analysis of the molluscan fauna of San
Francisco Bay. Univ. Calif. Publ. Zool. 18 (13): 236 to
299

Pearce, Jack B.

1965. On the distribution of Tresus nuttalli and Tresus capax
(Pelecypoda: Mactridae) in the waters of Puget Sound and the
San Juan Archipelago. The Veliger 7 (3): 166-170; plt.

27; 1 text fig. (1 January 1965)
Pratt, Davin M.
1953. | Abundance and growth of Venus mercenaria and Callo-

cardia morrhuana in relation to the character of bottom sedi-
ments. Journ. Mar. Res. 12: 60 - 74
Pratt, Davin M. « D. A. CAMPBELL
1956. Environmental factors affecting Venus mercenaria.
Limnolog. « Oceanogr. 1: 2 - 17
ReisH, Donatp J.
1961. A study of a benthic fauna in a recently constructed
boat harbor in southern California. Ecology 42: 84 - 91.
Ricketts, Epwarp FE « Jack CaLviIN
1962. Between Pacific tides. 34 ed., rev. by J. W. HepcPeTH
xili+502 pp.; illust. Stanford Univ. Press, Stanford, Calif.
SIEGEL, SIDNEY FE
1956. Nonparametric statistics for the behavioral sciences.
McGraw-Hill series in psychology, New York, pp. 1 - 312
Swan, Emery FREDERICK
1952. ‘The growth of the clam Mya arenaria as affected by the

substratum. Ecology 33 (4): 530 - 534
WELLs, Harry W.
1957. Abundance of the hard clam Mercenaria mercenaria in

relation to environmental factors. Ecology 38: 123 - 128

Page 250

THE VELIGER

Vol. 11; No. 3

Studies on the Mytilus edulis Community

in Alamitos Bay, California. - IV.

Seasonal Variation

in Gametes from Different Regions in the Bay

DONALD R. MOORE

DONALD J. REISH

Department of Biology, California State College at Long Beach, California 90804

(5 Text figures; 1 Table)

INTRODUCTION

THE BAY MUSSEL, Mytilus edulis LINNAEUS, 1758, is
distributed throughout Alamitos Bay, California (Figure
1). It is especially abundant wherever floating boat
docks occur, but in the upper reaches of the bay, notably
Colorado Lagoon and Cerritos Channel, the number of
specimens decreases. The chlorinity of the bay is uniform
throughout except immediately following infrequent win-
ter rains (STONE & ReisH, 1965). Since formation of
the gametes and subsequent fertilization and development
is essentially for repopulation, and since M. edulis is
distributed throughout the bay, the authors were curious
whether or not the mussels will produce gametes in all
areas of their distribution in the bay, and, if so, would
they be produced at the same or at different times of the
year. In other words, is the reproductive population in a
limited area of the bay surrounded by a vegetative, non-
reproductive population of /. edulis? Gonadal tissue was
examined periodically from 4 areas of the bay to de-
termine whether or not seasonal differences occurred
within an area and from area to area.

The seasonal reproduction of Mytilus edulis has been
studied in the past by the analysis of gonads (FIELp,
1922), CuipperFieLD (1953), Sucrura (1959), and by

seasonal attachment of larvae by ENGLE & LOOSANOFF
(1944), GraHamM & Gay (1945), CHipperFIELD (1953),
and Loosanorr & Davis (1963), and ReisH (1964a).
CHIPPERFIELD indicated sexually mature mussels occurred
from mid-April to the end of May in British waters.
Gamete maturation began when the water temperatures
reached 7° C and spawning commenced while the tem-
peratures were rising from 9}° to 124° C. In warmer
Japanese waters, SuciurA (1959) noted gamete matura-
tion occurred when the water temperature dropped to
18° C in the fall. Larval settlement took place in late
spring and early summer in Connecticut (ENGLE &
LoosanorF, 1944), in central California (GRAHAM &
Gay, 1945), and in England (CHippEN¥FIELD, 1953).
However, in Alamitos Bay larval settlement occurred in
late winter and early spring when the water temperatures
were not below 13° C (Reisu, 1964a).

Since the previous studies on seasonal reproduction in
Mytilus edulis have been carried out in more northern
latitudes, the purpose of this study was to determine
whether or not a seasonal variation in the maturity of
the ova and sperm occurred in Alamitos Bay. An addi-
tional purpose of this study was to investigate whether
or not variations in the degree of intensity of reproduc-
tion existed in populations from different areas of the
bay.

Vol. 11; No. 3

MATERIAL anp METHODS

Monthly samples of Mytilus edulis were collected from
4 localities within Alamitos Bay, Long Beach, California,
from November, 1964 through July, 1966 (Figure 1).
Subsurface water temperatures were taken at the time of
collection. Water samples were taken for chlorinity deter-
minations during the first year. Since this value varies
only during periods of rainfall (Stone & ReisH, 1965),

Figure 1

Map of Alamitos Bay, California, showing Station Locations

the chlorinity data were not included herein because they
were of little significance to this report. Samples were
taken from Stations 1 and 2 at the same time and at
Stations 3 and 4 one or two weeks later.

Stations 1, 3, and 4 represented extremes in the distri-
bution of the population where sufficient numbers of
specimens were present. Station 2 is in an intermediate
location where earlier studies on Mytilus edulis were
conducted (Reis, 1964a, 1964b). Occasional specimens
occur along the rock jetty at the channel entrance and
up the channel from Station 3, but there was not a
sufficient number of specimens to permit analysis. All
mussels were collected from floating boat docks; thus,
the specimens were always covered with water.

The specimens were brought back to the laboratory to
ascertain the degree of gamete development. Smears

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Page 251

were made of gonadal tissues, and the classification sys-
tem of CuippERFIELD (1953) was followed as:

Stage 0. Sex unknown (indeterminate); no follicles
present.

Stage 1. Follicles present; sperm mother cells and sper-
matids present; a few small oocytes scen.

Stage 2. Sperm present, but not motile; ova not fully
developed, wrinkled, and would not become spher-
ical when placed in sea water; no fertilization.

Stage 3. Sperm motile; ova fully developed and be-
came spherical when placed in sea water; ferti-
lization.

Recent spent stage: A few residual gametes left in
follicles.

Ten individuals of each sex were selected for micro-
scopic examination of gonadal smears. Mature males
could be distinguished by the cream or yellowish-cream
colored gonads, and mature females by their apricot or
red-brown colored gonads. The colors of the gonads
were light orange or light yellow in stages 1 and 2 of
both sexes. Gonadal smears were made of each speci-
men and the stage of development was noted. A separate
count of indeterminates or recently spent mussels was
made.

An average stage of gamete development was calcu-
lated for each male and female each month by totalling
the figures for the different stages observed and dividing
the number of individuals examined. The number of
indeterminates was totalled separately.

DATA

The data for the seasonal variation in male and female
gamete development over the 20-month period of obser-
vation are presented in Table 1 and Figures 2 to 5. The
range and average stage of gametogenesis for each sex
are presented by date for the 4 stations in Table 1. The
data are summarized graphically by station for each sex
for each period of observation as the average stage of
gamete development. These averages are based on 10
specimens of each sex except when a large number of
indeterminates was encountered. The number of inde-
terminates and the water temperature are also included
in each graph.

The data show that some degree of gametogenesis
occurred in both sexes to a greater or lesser extent at all
stations throughout the entire period of observation from
November 1964 through July 1966. However, the degree
of gametogenesis fluctuated seasonally in both males and
females. The higher levels of gamete development were
observed during the late fall and early winter months.

Page 252

Mussels with mature gametes decreased in numbers from
March to the end of July each year. The greatest num-
ber of indeterminates was observed from May to October.

SEASONAL VARIATION
IN MALE GAMETOGENESIS

Mature sperm were present in some specimens at all 4
stations throughout the period of observation except at
Station 2 in March and April 1965 and at station 4 in
July 1966 (Table 1). However, a seasonal variation in
the average stage of gamete development occurred at all
stations, and, furthermore, variations in the degree of
gamete maturity from station to station were noted (Fig-
ures 2 to 5).

A total of 68 samples was taken from all stations
during the 20 months of study; of these 14 (or 21%)

THE VELIGER

Vol Gi Nors

of the samples of males were composed entirely of
mussels at stage 3 of gamete development. Six of these
14 samples were observed at Station 1, 3 each at Stations
2 and 3, and 2 at Station 4. Only 3 samples, or 4%, of
the 68 samples did not have any individuals at stage 3 of
gamete development. The overall average stage of male
gamete development was 2.6, 2.5, 2.5, and 2.4 for Sta-
tions | to 4, respectively. The length of time during which
the average gamete stage of development was high (over
2.5) and longer was at Stations 1 and 2.

An inverse relationship between the male gamete stage
and water temperature was observed at all stations (Fig-
ures 2 to 5). The low point in the average stage of male
gamete development occurred from the months of March
to August when the water temperature rose from 17° C
to 24°C. The high point in the average stage of male
gamete development occurred from October to February
when the water temperature dropped from 24° to 13° C.

Table 1
Seasonal Changes in the Stages of Gamete Development in Males and Females of Mytilus edulis Based on a Numerical Scale
of | to 3!
Date Station 1 Station 2 Station 3 Station 4
Males _- Females Males Females Males Females Males Females
oO o o o o o
Bos) Css ee SB 88 See
[a4 < ew < fa <= < a4 <G (24 < C4 4 <
1964
November 18, 24 3 gO oe Io 2-3 28 1.8 1-3 23 1-2 18 2-3 24 1-3 18
December 2, 9 3 gd it To 2-3 28 1.1 2-3 28 1-3 1.7 Se od alc
1965
January 6, 13 3 30 toe 12 1-3 26 1.6 3 sg) wae iy Bos BS) i io
February 3, 10 3 30 1-2 IO Bos Pall 1.2 1-3 28 1-3 1.3 QoS By i io
March 3, 17 loQ Ze ioB iil 1-2 1.5 1F2 1-13) 23 21 1-3 21 1 £41.
April 7, 21 oS WS ot | tO 1-2 1.5 1.0 lo 2H te 13 log) 42 Wok ial
May 5, 12 1e=130 223) nol eee 1-3 25 1.0 1-3 23 1-2 1.4 2-3 24 1 1.0
June 2 to3 2B 1 10 1-3 21 1-2 11
July 12, 19 oe) ZA9 los ilo 1-3 2.0 12 lo Bi eZ 3} los 2A ils2 i133
August 2, 16 log ZO to2 ily 1-3 2.3 1.6 1-3) 2-2) et
September 9, 15 ls3 BS Ws Ws 2-3 2.9 1.3 loS 28 We2Q 18 lof 28 1209 ip
October 6, 13 3 30 toe 13 3. -S0 1.2 3 30 1-2 14 2-3 26 1-2 14
November 10 2283 28) WB) id 2-3 2.9 1.4 3 30 1-2 15
December 1, 8 3 gO oD i 2-3 29 1.4 2-3 29 1-3 19 2-3 28 1-2 1.1
1966
February 2 2-3 29 1-3 18 2-3 2:9 2.4 3 30 1-3 18 1-3 21 1-2 12
March 17, 22 2o8 B83 NeoQ ill 3 3 1.6 2-3 29 1-3 16 2-3 2.7 1-2 1.2
May 3, 31 1o8 2G iloQ ill 3 3.0 1.3 1-3 20 1-3 23 1-3 26 1 1.0
July 12 1-3 18 1 1.0 1-3 2.2 162 oe eo Gl 1.0 1-2 14 1-2 13

Based on analysis of 10 specimens of each sex whenever possible.
The analysis of less than 10 specimens occurred frequently
during the summcr months when the number of indeterminates

was higher.

Vol. 11; No. 3

ee S|

1

Number of Indeterminates

iS)

i) =

& .

2

2 hg aa

ral 0 es

i=)

& 13 T T T T aT a T a T T

1964 1965 1966
Figure 2

The seasonal variation in the average stage of male and female
gamete development, the number of indeterminates, and water
temperature at Station 1

SEASONAL VARIATION
IN FEMALE GAMETOGENESIS

Mature ova were present in some of the specimens in
only 12 of the 68 samples analyzed and 6 of these were
from Station 3 (Table 1). These mature ova were ob-

5 male
es)
S «— female
7)
2 6
OF 3
iS Je
3S FS S
OL, 0,6
vo
612
° indeterminate Se,
O és —~e ~ lie
Ore T | (aaa ea a y £
iS e
2| Z
=) 5 °
a {19
ot
o. e
& 13 T T T —
L1964 1965 1966
Figure 3

The seasonal variation in the average stage of male and female
gamete development, the number of indeterminates, and water
temperature at Station 2

THE VELIGER

Page 253

served during some of the months between November
and March except one instance in September 1965 at
Station 4 and one in May 1966 at Station 4. A seasonal
variation in the average stage of gamete development
was noted at all stations during the 20-month observation-
al period. While a considerable variation occurred from
station to station, especially regarding the extent of the
period of higher averages, the average stage of female
gamete development was highest from November through
February and lowest from about March through Sep-
tember (Figures 2 to 5).

Of the 68 samples analyzed for the stage of female
gamete development 12 (or 18%) had specimens only in
stage | or the follicle stage. Stage 1 was always observed
in some of the specimens in a sample except at Station 2
in February 1966. The overall stage of female gamete
development was 1.2, 1.4, 1.5, and 1.2 for stations 1 to 4,
respectively. The length of time during which the average
gamete stage of development was highest (over 1.7) was
longer at Station 3 and shortest at Station 4.

Gamete Stage

lop ce)
Serene)

Number of Indeterminates

3
OL (0)
@ = er
123
al e
ome 5
Bl
See ee —
1964 1965 1966

Figure 4

The seasonal variation in the average stage of male and female
gamete development, the number of indeterminates, and water
temperature at Station 3

NUMBER
or INDETERMINATE INDIVIDUALS

The fluctuation in the number of indeterminates observed
correlated inversely with the seasonal aspect of male and
female gamete development. The number of indeter-
minates increased between April and August when the
average level of gamete development was decreasing. The

Page 254

total number of indeterminates observed during the study
was 31, 13, 43, and 36 at Stations 1 to 4, respectively.

o

s

n

2 b

& 3

& a

io)
:
<
5
6

7 E

& 123 E

2 <J % 2 Z

3

é i une

o

e

oO

a 7o6) leas Ta 1965 1966

Figure 5

The seasonal variation in the average stage of male and female
gamete development, the number of indeterminates, and water
temperature at Station 4

DISCUSSION

The occurrence of mature gametes in Mytilus edulis
reached a peak at all stations in the fall and early winter
months in Alamitos Bay when the water temperatures
decreased from 20° to 21° C in September to 15° C in
December. Some stages of gametogenesis were observed
in both sexes at all stations during the period of 20
months. More advanced stages were observed for longer
periods of time in males than in females (Figures 2 to
5). These observations were similar to those reported by
Sucrura (1959) for M. edulis in Japanese waters. Sexu-
ally mature mussels were abundant in October and No-
vember when the water temperature was 18° C and least
abundant during August when temperatures reached
284° C. Both of these observations are different to what
was observed by CuippERFIELD (1953) in colder British
waters. Gametes matured when the water temperatures
rose above 7° C and spawning occurred from 2 to 6
weeks from mid-April through May when temperatures
rose from 94° to 124° C. After spawning, CHIPPERFIELD
noted a period of 2 to 3 months when all mussels were
indeterminate.

Apparently, in the colder British waters the develop-
ment of gametes and spawning are limited by water

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Vol. 11; No. 3

temperatures to a greater extent than in the more tem-
perate waters of Japan and southern California. The
period of time in which mature gametes are present is
considerably longer in Japanese and southern California
waters. Recovery from spawning with the subsequent re-
development of the next generation of gametes occurs
faster in the Japanese and southern Californian popula-
tions of Mytilus edulis. Longer reproductive seasons were
found in the sea urchin Strongylocentrotus purpuratus
(Stimpson) which were collected from warmer waters
of lower latitudes than from colder waters of higher
latitudes (BooLooTIAN & GiEsE, 1959).

Gonad analyses could not be made from specimens of
Mytilus edulis collected at the extremes of its distribution
within Alamitos Bay because of insufficient numbers being
present. Some differences, indicated by trends, have been
noted at Stations 1 to 4. The overall average stage of
male gamete development was slightly higher at Station
1 than at Station 4 with the other 2 stations intermediate.
The period of time during which motile sperm are found
is longer at Stations 1 and 2 than at Stations 3 and 4
(Figures 2 to 5). Variations in the average stage of
female gamete development were observed but not at the
same stations as for male gametes. The highest overall
averages were noted at Stations 2 and 3 and the lowest
at Stations 1 and 4.

It is apparent from these data that the propagation of
Mytilus edulis in Alamitos Bay is dependent upon the
presence of mature ova. Some males with mature, motile
sperm were observed throughout the year, but mature
ova were observed only during the late fall and winter
months when water temperatures were at their lowest
level (13° to 15° C). The maturation of male gametes
does not seem to be affected by the range of water tempe-
raturesin Alamitos Bay asit isin British waters (CHIPPER-
FIELD, 1953): however, the maturation of the ova ap-
parently is influenced by temperature in Alamitos Bay.
The presence of motile sperm for a longer period of
time than mature ova may be the result of: (1) not all
sperm from a mussel are discharged at any one time,
(2) additional sperm are produced rapidly and many
times a year following a complete spawning, or (3) a
combination of these. The shorter period of time in which
mature ova are present may be the result of: ova are
spawned from one mussel at one time and additional ova
mature slowly and are produced only once a year.

The period of settlement of Mytilus edulis on boat
floats in Alamitos Bay in the late winter to early spring
months substantiates these observations of a late fall to
winter spawning season. The length of larval life is un-

Vol. 11; No. 3

THE VELIGER

Page 255

known for these waters, but CHIPPERFIELD (op. cit.)
calculated it to be about 4 weeks.

As noted above, there are indications that some eco-
logical variations in the average stage of gamete develop-
ment occurs in Alamitos Bay. It would be of interest to
study this variation in a body of water where the popula-
tion of mussels is sufficient in the environmental extremes.
Secondly, Mytilus edulis would be a convenient organ-
ism with which to work in order to study the relationship
of temperature, as a function of latitude, to the length
of the reproductive period.

SUMMARY

1. Gonadal smears of the bay mussel Mytilus edulis were
made at 4 stations at monthly intervals for 20 months in
Alamitos Bay, Long Beach, California, to ascertain
whether or not seasonal variation in gamete maturity
exists.

2. Some degree of gametogenesis occurred throughout
the period of observation. Mature sperm were present
in some mussels throughout the year, but mature ova
were present only from November through May. The
number of indeterminate individuals varied directly
with the water temperature.

3. Some slight variations were noted in the average
stage of gamete development with both sexes collected
from the different stations.

LITERATURE CITED

CHIPPERFIELD, Pup N. J.
1953. Observations on the breeding and settlement of Mytilus
edulis (L.) Journ. Marine Biol. Assoc. U. K. 32: 449 - 476
ENGLE, James B. « Victor Lyon LoosANoFF
1944. On season of attachment of larvae of Mytilus edulis
Linn. Ecology 25: 433 - 440.
Fie.p, Irvine A.
1922. Biology and economic value of the sea mussel (Mytilus
edulis). Bull. U.S. Bur. Fish. 38: 127 - 259
GraHaM, HERBERT WILLIAM & HELEN Gay
1945. Season of attachment and growth of sedentary
marine organisms at Oakland, California. Ecol. 26 (4):
375 - 386
LoosanorrF, Victor Lyon « Harry Cari Davis
1963. Rearing of bivalve mollusks. Jn Advances in marine
biology, vol. 1, FS. Russell (ed.). Academic Press, London,
pp. 1 - 136
Retsu, Donatp J.
1964a. Studies on the Mytilus edulis community in Alamitos Bay,
California: I. Development and destruction of the community.
The Veliger 6 (3): 124-131; 1 map; 4 text figs. (1 Jan. 64)
1964 b. Studies on the Mytilus edulis community in Alamitos Bay,
California: II. Population variations and discussion of the asso-
ciated organisms. The Veliger 6 (4): 202-207 (1 April’64)
Stone, ALFRED N, « Donatp J. Reisu
1965. The effect of fresh-water run-off on a population of
estuarine polychaetous annelids. Bull. So. Calif. Acad. Sci.

64: 111-119
Suciura, Y.
1959. Seasonal change in sexual maturity and sexuality of

Mytilus edulis. Bull. Japan. Soc. Sci. Fish. 25: 1-6

Page 256

THE VELIGER

Vol. 11; No. 3

Two New Cypraeid Species in the Genus Erronea

BY

CRAWFORD N. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plate 46)

ANOTHER COWRIE SPECIES that appears not to have been
previously known has been sent to me by Mr. Fernando
G. Dayrit of Manila. This very interesting animal was
collected at the northern end of Samar Island, eastern
Philippines, in the area of San Antonio.

At first inspection the shell reminds one of the east
Australian Erronea (Gratiadusta) walkeri continens (IrE-
DALE, 1935). However, many of the shell characters and
the dorsal color-layering prove it to be otherwise. The
differences are striking, and there seems to be no other
alternative than to compare it with another east Austral-
ian species, E. (Adusta) xanthodon (SoweErsy, 1832), to
which it seems most closely related. Because this shell
does appear to be new to science I propose to call it

Erronea fernandoi C. N. Cate, spec. nov.

(Plate 46, Figures la to 1d)

Holotype: Shell long, narrow, sub-pyriform, heavy; base
somewhat convexly swollen, narrowly concave anteriorly;
lip narrow; terminals well formed, semi-beaked; margin
perceptibly thickened on left side and rounded; right
margin thickened, angled, upswept and_ shouldered;
both margins flattened anteriorly, flanged, forming a
sharp angle with vertical surface of body whorl; aper-
ture straight, narrow, sides parallel; teeth numerous, very
small, fine, slightly longer on columella, — on outer lip

short, blunt, broader, with wider interstices, — in either
case only very slightly extending onto base; columella
smooth posteriorly, with adaxial edge indistinctly lined
with widely spaced, barely visible, larger, rudimentary
teeth; columellar teeth pronounced at adapical point of
long, narrow, shallow fossula, where base teeth are much
heavier, and cross the fossula in bold relief; terminal ridge
almost straight, abapical left side bifidly grooved into
two parts; rear surface of shell umbilicate, with dark-
brown to brownish black apex faintly visible within.
Primary shell color light grey, overlaid with a thick
mass of irregularly spaced light brown flecks, resulting in
an olive green appearance, which is much lighter in some
specimens; a central wide dark brown blotch, in some
cases a broad band, covering approximately one third of
the dorsum (a central dorsal blotch would be an excep-
tion to this), extends from margin to margin, becoming
lighter as it approaches left side; large dark brown color
blotches on either side at base of abapical terminal col-
lar, with loosely scattered large brown spots in both mar-
gins, nearly becoming obscure on the base; base, teeth,
interstices, and inner surface of terminals white, occa-
sionally very pale beige; columella and upper surface of
terminals pale brown.
Measurements: The holotype of Erronea fernandoz is
19.0mm long, 10.4mm wide, and 8.8mm high, and
shows 22 labial and 18 columellar teeth; the terminal
ridges were excluded from the count.

Explanation of Plate 46

Figures 1a to 1d: Erronca fernandoi C. N. Cate, spec. nov. (X 24)
Figures 2a, 2b: Erronea steineri C. N. Cate, spec. nov. (X 24)

1a, 2a: Dorsal view; 1b, 2b: Ventral view; 1c: Right lateral view; 1d: Left
lateral view of the holotypes.

THE VELIGER, Vol. 11, No. 3 [C.N. Cate] Plate 46

Figure 2 a Figure 2 b

photographs by Jean M. Cate

Vol. 11; No. 3

THE VELIGER

Page 257

Type Locality: The animal was alive when collected in
approximately 10 feet of water at San Antonio, Samar,
Philippines, the type locality for the species. Located in
southwestern Samar, on San Pedro Bay (approximately
11° 03’ N Latitude; 125° 00’ E Longitude), this locali-
ty is noted for many other cowrie species.

Type Repository: The holotype will be deposited in the
California Academy of Sciences Geology Department
Type Collection where it will bear the catalogue number
13158.

I have named this species in honor of Mr. Fernando

G. Dayrit, who has contributed many new species to
science and thereby has added to our knowledge of the
mollusca.
Discussion: The species most closely related to Erronea
fernandoi appears to be E. xanthodon (SowErBy™, 1832),
but E. fernandoi differs from the East Australian form in
having 2 unusual obliquely transverse ridges on the left
anterior ridge; by having shorter, oblique abapical ter-
minal teeth; by having a long, narrow, shallow and
heavily denticulate fossula; by the distinctly sharp upper
edge of the columella; by the margins becoming pinched
and more distinctly flanged anteriorly, which produce
deep indentations on either side of the terminal collar;
by having a large dark brown dorsal blotch; and by
having a white base, white teeth and interstices.

The affinity of these two species to one another is
evident. The outstanding differences in the two species
are distinctly obvious in the unusual white base (occasion-
ally it may vary to a pale beige), and the large dorsal
blotching (a character almost never seen in Erronea xan-
thodon) . However, while there might be some justification
for considering these two cowrie forms as subspecies be-
cause of some similar morphological features, I am pro-
visionally listing this new cypraeid as a full species.

Since having completed the work on the supposedly
unique specimen of Erronea fernandoi, two additional
specimens were given to me for identification by Mr.
William Old, American Museum of Natural History. I
am including them here for comparative purposes. I shall
designate the larger of the two shells hypotype 1, and the
smaller of them hypotype 2 (the 2 new specimens are
not a part of the original type lot from southwestern
Samar, but appear to have been collected at Zamboanga
City area, Mindanao, a point considerably to the south
of the type locality). The data for the 2 shells from
southern Mindanao, using the millimeter scale for length,

width, and height, and counting the teeth on lip and col-
umella are as follows:

hy pPotypemliz i(29 ion alG:Siw 3:00 24 ar 1))

hypotype 2: (22.4 13.3 11.0 21 18)

The Zamboanga specimens, as can be seen, are both
considerably larger shells; in addition it should be pointed
out that the large brown dorsal blotches on these 2 shells
appear as large central islands of color, rather than the
sweeping transverse band of color shown in the holo-
type (Plate 46, Figure 1a). In contrast, this dorsal color
blotching is almost never, if at any time, seen in the East
Australian Erronea xanthodon. I wish to thank Dr. Franz
A. Schilder for confirming the decision I had made that
E. fernando? is, in fact, a new species.

Erronea steineri C. N. Cate, spec. nov.

(Plate 46, Figures 2a, 2b)

Shell of medium size, pyriform, elevated and bulbously
inflated, light weight, strong; base narrow, convex; ter-
minals well formed, barely protruding, with openings ap-
pearing to be small for the size of the shell, especially
abapically; apex umbilicate; sides only slightly thickened,
rounded; teeth on outer lip fairly large, broad, interstices
shallow; base teeth weakly formed, though distinct, cent-
rally long over base, becoming shorter in front; curving
fossula extends half the length of shell, is fairly deep,
denticulate, terminating boldly on inner edge of fossula;
columella broad, smooth; aperture wide, curving, weakly
constricted abapically; shell narrowly, transversely band-
ed with 4 dark lines barely visible through surface, most
distinct on left side. Primary shell color pale ivory-white;
dorsum loosely, though generously, flecked with light
chestnut-brown; sides, terminals, base, teeth, interstices
pale ivory-white; columella pale ivory-white spotted with
8 large pale chestnut-brown spots, well separated, ar-
ranged in vertical pairs; brownish-black apex barely
visible at bottom of umbilical depression.

Measurements: ‘The unique holotype of Evronea steineri
is 22.6mm long, 14.5 mm wide, 12.3 mm high, and has
16 labial and 17 columellar teeth; terminal ridges are not
included in the count.

Type Locality: This animal was collected alive during
April, 1968. in the shallow subtidal waters of Ufa; this
small island is a member of the Russell Island Group,
situated approximately 30 miles northwest of Guadal-

Page 258

THE VELIGER

Vol. 11; No. 3

canal in the southeast British Solomons (approximately
9°08’ S Latitude; 159°00’ W Longitude).

Type Repository: The holotype will be deposited in the
California Academy of Sciences Geology Department
Type collection, where it will bear the catalogue number
13157.

Discussion: Erronea steineri is a fully adult live-collected
shell, possessing many of the visible morphological aspects
of both Erronea hungerfordi (SowErBy™, 1888)and E.
subviridis (REEvE, 1845). Possibly the new species fits
morphologically somewhere in between the two. How-
ever, it seems quite distinct from any other cowrie species.

Erronea steinert differs by having less prominent termin-
als, a conspicuously different shell shape, longer base
teeth, less thickly formed margins and sides, and a differ-
ent and unusual color design on the dorsal shell surface.

I am naming this new cowrie species in honor of Mr.
Franz B. Steiner, Field Associate, Department of Inver-
tebratcs, California Academy of Sciences, San Francisco,
who first realized this cypraeid might be new to science.
I wish to thank Jean Cate for the photographs illustrat-
ing both of these new cowrie species, as well as for other
assistance in arriving at the conclusions contained in this

paper.

Vol. 11; No. 3

THE VELIGER

Page 259

Growth Study in Oliwella biplicata (Sowersy, 1825)

BY

RUDOLF STOHLER

Department of Zoology, University of California, Berkeley, California 94720

(1 Map; 1 Text figure; 11 Tables)

INTRODUCTION

THE STENOGLOSSAN SNAIL, Olivella biplicata (SowERBY,
1825) has a wide range of distribution along the coast
of western North America, extending, according to KEEN
(1937), from Magdalena Bay to southern British Colum-
bia. Within this range I have collected or observed this
species from Estero de Punta Banda in Lower California
to Seaside, Oregon and found it always to be very abun-
dant in the proper habitat.

At Duxbury Reef in Bolinas Bay, Marin County, Cali-
fornia, where I made regular visits at 2-week intervals
for over a year and a half, there are several sand flats
measuring, on an estimate, between 2000 (the smallest)
and 10000 m? (the largest). An attempt was made to
estimate the population of Olivella in the latter area.
One square meter areas were delimited by strips of
stainless steel and the sand was removed to a depth of
4 inches and sifted through a 1 mm mesh screen. From
5 such samples, taken randomly, I arrived at an estimate
of 250000 individuals for this particular area. Estimates
in other localities along the range of my observations
vary from a minimum of 10 to a maximum of 98 indi-
viduals per square meter. Chance encounters of 540 or
so individuals concentrated in an area of less than 4
square decimeters (i.e., less than half a square foot!) at 3
different localities were, of course, disregarded in these
estimates. It is, therefore, justified to call O. biplicata
a common species.

Olivella biplicata seems to thrive equally well in the
intertidal sand flats of exposed stretches of the coast and
in protected areas of harbors, bays, and esteros. This
statement is based on estimates of population densities
in various areas, as discussed above.

As Olivella biplicata reproduces throughout the year,
at least within the area of its occurrence in California
(Epwarps, 1968; SToHLER, 1959/1960; Zeit, 1955)
and in Oregon (Epwarps, op. cit.), estimates of longe-
vity based on “year classes” are impossible. At Duxbury
Reef I noted that these animals upon the return of the

waves after a low tide travel surprisingly rapidly and far,
often several meters in less than half an hour. Under
such conditions it appeared hopeless to undertake a long-
range growth study in an area facing the open ocean,
as the probability of recapturing marked individuals
would be practically zero on the basis of pure chance
alone. However, when I observed the thriving population
of O. biplicata in the Flood Control Channel (FCC here-
inafter) at San Diego, California (see Sketch Map) a
possibility of recovering adequate numbers of marked
individuals appeared reasonably certain. Early efforts
along this line of endeavor have been detailed in StoH-
LER, 1962. Not only was this assumption proved correct,
but additionally several observations became possible
here which could not have been made in the open ocean
without grave danger to personnel.

In the FCC there are channels in the sand which at
extreme low tide are still covered by several feet of wa-
ter; no accurate measurements have been made, but by
the fact that divers standing at the bottom were covered
by about 2 feet of water, an estimate of about 7 to 8
feet depth appeared reasonable. It was reported (SToH-
LER, 1959/1960) that Olivella biplicata remains dor-
mant in the intertidal area during exposure of the habitat
at low tide and that, upon first return of waves after the
turning of the tide, the animals become very active again.
The same behavior is observed even though the sand re-
mains covered by several feet of water in the FCC; thus,
the best success in recovering marked specimens was ob-
tained just after slack tide; this held true whether the
low tide was early in the morning or late in the evening.

Another observation which seemed surprising to me
was that certain individuals were recaptured within a
few feet of the spot where they had been dropped on
previous occasions. To be sure, there was a certain a-
mount of scattering to be noted; however, it is impossible
to state that this scattering is due solely to the active
moving of the animals, or that it may be attributable
wholly or in part to a passive dislocation by external
agents, such as heavy waves or strong currents. It was

Page 260

THE VELIGER

Vol. 11; No. 3

Entrance to Mission Bay N\
Y

> we
Experimental Are:

IRS

Pacific Ocean

Map of the Flood Control Channel in San Diego, California

observed under water that individuals are carried fairly
long distances by the currents if the animals become dis-
lodged. Nevertheless, it must be assumed that some active
travelling does occur as all captured animals, marked
and unmarked alike, were returned after examination
in one small area of less than a square meter. ‘These
artificially created densitics of populations —_ several
thousand individuals per square meter — would prob-
ably cause the individuals to seek less densely populated
feeding grounds.

The FCC was, however, not without disadvantages.
Since it is a flood control channel, at times of extremely
heavy winter rains there is a very heavy run-off of fresh
water, and though fresh water may tend to form a layer
above the sea water it will, nevertheless, tend to kill off
many of the animals living in the upper portions of the
sand exposed or nearly exposed at low tide. Such extreme-

ly heavy rainfalls do not occur often in the San Diego
area and during the term of our observations there only
two such events took place, both times with -— for the
purposes of this study -— catastrophic results.
Another disadvantage of the FCC for the study is its
accessibility to the pleasure fishermen and their families.
The children pick up the rather attractive shells of Oli-
vella biplicata and carry them away. However, the worst
case was that of a “shell jewelry” collector who scooped
up many thousands of Olivella animals exactly in the ex-
perimental area. These untoward occurrences are re-
flected in the tables, to be discussed below, by the small
numbers of animals gathered on various occasions.
The entire study was carried out over a period of 9
years, from July 27, 1959 to July 13, 1968. The area was
visited every 3 months and strenuous efforts were made
to recover as many marked individuals as possible. ‘This

Vole INow3

required the participation of several to many individuals
and it is a great pleasure to acknowledge the enthusiastic
participation of many persons; it would hardly be fair to
single out any individual for special thanks, except per-
haps a few who participated at every “Olivella Dive’;
among these are Mrs. Fay Wolfson, Mr. and Mrs. R.
Dilworth and, later, practically all the members of the
San Diego Shell Club and at one time or another, mem-
bers of various SCUBA diving clubs from the San Diego
region. Without this help, it would have been impossible
to carry out this study since one individual could not
possibly hope to collect, even under the best conditions,
the number of individuals obtained on the different oc-
casions. To all these persons go my sincere thanks.

MATERIAL ann METHODS

The species used for the investigation reported here is
Olivella biplicata (SoweERBy, 1825). As described earlier
(STOHLER, 1962), the animals were collected first at

1960

(226)
16.0 - 18.0 mm

VANS
1961
II

(347) (316) (167)
18.1 - 20.0 mm

14.1 - 16.0 mm 16.1 - 18.0 mm

(964)
14.1- 16.0 mm

»)

1967
IX

(1005)
15.5 - 16.0 mm

THE VELIGER

IV

Page 261

Solana Beach, a few miles north of San Diego; later, the
collections were made at Shelter Island in San Diego
Harbor by members of the San Diego Shell Club. Until
measured, selected and marked, the snails were main-
tained in large tanks with running sea water in the ex-
perimental aquarium of Scripps Institution of Oceano-
graphy at La Jolla. The method of measuring and mark-
ing was given in an earlier paper (STOHLER, 1962).

Other methods of marking had been tried, as stated in
the earlier report, and found unsuitable. Tagging with
discs, permitting individual numbering, as described by
Darsy (1964) for Tegula funebralis (A. ApAMs, 1855)
was deemed highly desirable from the point of view that
this method would be capable of yielding accurate re-
sults, but was considered unsatisfactory because the pro-
truding tags would interfere with normal activities of
the marked snails and thereby possibly affect growth rate
and life span.

To obtain simultaneously data for several stages in the
growth series of Olivella, the animals in 1961 were di-

(70)

20.1 - 22.0 mm 22.1 - 23.0 mm

23.1 - 24.0 mm

Numbers in parentheses indicate number of individuals marked for that size class.
Roman numerals refer to the corresponding column in Table 10.

Figure 1

Marks used in different years and for different size classes

Page 262

vided into different size classes and marked diffcrently
(see Figure 1). After the heavy rains of December 1964
this particular series was forcibly terminated. In 1965 I
returned to the marking of a single size class, but to
offset the predation by sea gulls and removal by human
beings I used almost 1000 individuals. When this partic-
ular experiment again was terminated by the heavy rains
of 1966, I selected for the final attempt in 1967 a much
more limited size class, spanning not more than 4mm,
while previously the size classes comprised all individuals
within a 2mm range. This last selection was made pos-
sible by the enormous number of specimens collected by
members of the San Diego Shell Club. It was possible to
select over 1000 individuals in the size range 15.5 to
16.0 mm, with a majority measuring exactly 15.5 mm.
Although I did not keep a count of the total number, I
think close to 25000 animals were collected; from these
the experimental group was selected. Of course, as al-
ways, the unused animals were returned to their original
habitat and the marked ones were deposited in the ex-
perimental area in the FCC.

THE VELIGER

Table 1
oO 43) wr) —
u = i) @
S ye ies ou
oO. oO. So a
2 Se ee
ae a we ay £e
Gey Bi 77 Ss + N
© -Q Ss 6} (S|
g so es me q
g 5... § a S
oC Z 7p) —] a
1960 XI 5 5 {18.6 } {20.25} {19.07}
x6 2 (BE GO” } KO23)
18:6 20.25 19.14
1961 I 28 7 17.8 21.9 20.01
IV 22 4 17.7 20.9 19.1
VII 4 6 17.6 19.7 18.53
xX 14 0
1962 i 6 2 19.3 20.1 19.7
IV 14 3 18.5 25.0 20.83
VII 12 2 19:2 21.0 20.01
x © 0
1963 It iS 1 22.52
IV 6 1 20.9
VII 21 4 20.18 25.55 23.11
MS 5 0)

starting date: 16 VII 1960

226 individuals marked

starting sizes: 16.0 mm to 18.0 mm
minimum increment 2.18 mm to 4.18 mm
maximum increment 9.55 mm to 7,55 mm

Vol. 11; No. 3

RESULTS

The results are summarized in the 11 Tables that follow.

General explanation for the tables: Because of the
range in each size class (2 mm in the case of Tables 1, 2,
3, 4, 5, and 8; 1 mm in that of Tables 6 and 7; and 4mm
in that of Table 8) minimum and maximum increments
given at the foot of each Table are cited with these be-
ginning ranges reflected in the end results. In Table 1,
separate recovery collections were made on 2 successive
days and the results computed separately; these are re-
corded brackets in { } in the Table, with the total from
both days following in the appropriate columns. Because
of the possibility of adversely affecting the marked ani-
mals by keeping them overnight in an aquarium, no
more collections were made on successive days thereafter.

Table 1: Of the 226 individuals marked in 1960 only 4
were recaptured after 3 years. Because of the initial size
range of 2.0 mm an accurate measurement of the total
increment is impossible. However, it is possible to state
that Olivella biplicata grew not less than 2.18 mm nor
more than 9,55 mm in the 3 years. It is noteworthy, how-
ever, that at least one individual had attained almost
the final size as much as 14 years earlier.

Table 2
- Bs =,
5 BS E
a 2 Se Le
2 Se se ie eI
. Ba ae
2 eae aoe d
A Ze Ss =
1961 XxX 14 13 15.4 19.2 Ad
1962 I 6 3 17.8 19.1 18.46
IV 14 7 17.4 20.4 18.72
VII 12 7 17.9 Diol 20.24
xX 6 8 18.8 22.7 20.54
1963 I 13 5 22.8 23.9 23.49
IV 6 8 19.0 25.2 22.53
VII 21 10 19.6 26.8 22.64
x5 4 Di 2) 23.6 22.5
1964
1965
1966 VII 9 1 24.9
1967 Ihy 2 1 27.8

starting date: 3/4 VII 1961

347 individuals marked

starting sizes: 14.1 mm to 16.0 mm
minimum increment 11.8 mm
maximum increment 13.7 mm

Vol. 11; No. 3 THE VELIGER Page 263

Table 2: Of the 347 individuals marked in 1961, one Table 4
was recovered as much as 53 years later and had, by that
time, attained almost the maximum size (31.2 mm) ‘

"al nae
reported for Olivella biplicata (STOHLER, 1960). = 5 5 ‘g
Table 3: This group was of some interest because for e e 28 = 2
almost 2 years no member of this size class was recovered ; fe & ol = = =
yet, when at the end of 4 years one specimen was found, i § Ks 9 z I =
it had grown less in this total time than one other speci- © Be oo I
men had in one year. A 7 & 5 =
1961 X 14 3 20.1 20.8 24.43
Table 3 1962 I 6 2 21.4 22.0 2ilied
IV 14 5 20.7 (ep 21.42
mae ar VII 12 1 2353
e = s S x © 1 24.0
2 8) ol eles 1963 1S | Bit BD Bi Pare.
§ SS ee NY Gi Ge Le ee Ee
‘s Bet oa f=) 9A, VII 21 3 2319 25.8 24.63
5 se Oe ae P Kee re 24.6
S ERTS ee er S 1964 WII 5 1 22.3
fa Za 4 a
i561 X 14 74 198 18.96 starting date: 3/4 VII 1961

: 90.2 167 individuals marked

5)
ae 2 © l starting sizes: 18.1 mm to 20.0 mm
IV 14 2 IS) 21.0 20.45 aes ‘ Sian
minimum increment (last captured individ.) 2.3 mm
WADTeptr2 3 21.0 25.8 22.83 : : Nerene
maximum increment (last captured individ.) 4.2 mm
2S © ‘ Wed Lis) al maximum increment (any individual) 7.7 mm to 5.8
1963 I 13 2 2p DB) 24.3 aa
EV 16 2 235ml 2323 Def
VII 21 5 Dee 24.3 23.42
ieee Table 5
N9G5 ee WALT 24 1 23.6
starting date: 3/4 VII 1961 v 2 3 =
316 individuals marked 2 fe ao) =
0 0 Q. jor ae) =O
starting sizes: 16.1 mm to 18.0mm aS s ao 2 oO
maximum increment (any indiv.) 9.7 mm to 7.8 mm g & Je 5 = =
minimum increment (last captured individ.) 5.5 mm ‘ g % S z 3s Bs
maximum increment (last captured individ.) 7.5 mm c E |x 5p a
S 5 E a ~
a) Zz, Nn — a
Table 4: Again one individual recovered after 3 years 1961 X 14 1 21.6
had grown less than one other individual had grown in 1962 I 6 0
one year. IV 14 1 22a
WANE WP 0
Table 5: This group and the next yielded the poorest XenG 1 25.8
results. Because of a peculiar mark caused by a slip as 1963 I 13 0
the shell was pressed against the marking drill, the indi- ay & 0
vidual animal was recognized; unfortunately, an oppor- VII 21 1 95.9
tunity to obtain exact results was missed in this instance. == = =
I did not anticipate the possibility of recapturing exactly starting date: 3/4 VII 1961
the same individual 4 times at various intervals and ther- 70 individuals marked
fore did not record the individual size of the one where starting sizes: 20.1 mm to 22.0 mm
the shell had slipped. Thus, the animal whose measure- minimum increment 3.9 mm

ments are recorded in ‘Table 5 may havc been as small as maximum increment 5.8 mm

Page 264

20.1 mm when marked but could not have been more
than 21.6 mm. However, it attained almost its maximum
size within the first 18 months.

Table 6: Only one individual of 26 originally marked
was recovered after 14 years. In that time it had not
grown more than 14mm.

Table 6

e 2 OS =

Z 2 oe ee

< s 22 38

9 © E5 a5

= =) om rv & =F

‘S & be fa

© I = x 5p & q

a 5 & o

fa Ze =] =
1961 xX 14 3 22.1 23.7 23.3
1962 li © 1 23.6

starting date: 3/4 VII 1961

26 individuals marked

Starting sizes: 22.1 mm to 23.0 mm
minimum increment 0.6 mm
maximum increment (any individual) 1.6 mm
maximum increment (last captured individ.) 1.5 mm

Table 7: In this group we again find that one individual
attained in one year a greater length than another in 9
more months.

Table 7
@ o° S ra
3 i es) 2
Se ee Be Be
© So Ee Ee
% Be ae
g ¢ de is ;
A 2 &§ 4 s
1961 xX 14 4 24.1 24.4 24.75
1962 I 6 0
IV 14 OAD 95,5 94.7
VII 12 DDG 26.6 25,65
x 6G 1 26.3
1963 I 13 2 | OBO 26.3 95.75
IV 6 1 24.5

THE VELIGER

Vol. 11; No. 3

Table 8: This group was subjected to the effects of one
extremely heavy rainfall runoff and one fairly heavy one.
Yet one individual survived both of these catastrophes.
The lone survivor had added in 3 years a maximum of
11.2 mm to its length.

Table 8
0 i) =
5 3. 5
a Qa So 255)
§ Bu sh ee
S 2 &3 ES
Ge w wy A, “= Oy
} ® o8 2 S
L G& AR bn 5
3 5 g a x
a) ZS 4 a
1965 VII 24 10 14.8 19.8 17.4
Xl, 16 15.3 21.4 18.37
1966 I 15 0
IV 17 3 16.9 20.6 19.17
VII 9 4 16.9 19.6- 18.23
ee 115) 6 18.0 21.6 18.22
1967
1968 VII 13 1 DP)

starting date: 10/12 I 1965

964 individuals marked

starting sizes: 14.1 mm to 16.0 mm
minimum increment 9.2 mm
maximum increment 11.2 mm

Table 9: In some respects this was the most disappointing
experiment. Of the very large number (1005) of experi-

Table 9

Smallest individual
recaptured

Largest individual
recaptured

| Date of recapture
a

—
[op)
nN
—_
“I
1S)
—
Dn
~I
on

1967
1968 I 113)
IV 6 13 170 19.8
VIL 13 16 15.6 20.6

© ~| Number recaptured

starting date: 3/4 VII 1961

109 individuals marked

starting sizes: 23.1 mm to 24.0 mm
minimum increment 0.5 mm to 1.4 mm
maximum increment 3.5mm to 2.6 mm

starting date: 11/13 VII 1967

1005 individuals marked

starting sizes: 15.5 mm to 16.0 mm
minimum increment 0.1 mm
maximum increment 5.1 mm to 4.6 mm

Vol. 11; No. 3 THE VELIGER Page 265
Table 10
Minimum and Maximum sizes At Six Months Intervals
I II III IV V VI VII VIII IX
(226) (347) (316) (167) (70) (26) (109) (964) (1005 )
14.0-16.0 14.1-16.0 16.1-18.0 18.1-20.0 20.1-22.0 22.1-23.0 23.1-24.0 14.1-16.0 15.5-16.0
6 mo 17.6 21.9 17.8 19.1 20.2 21.4 22.0 - 23.6 = 14.6 19.8 =
12 mo 7a Wa gS) Bile Pil Pos 23.3 - ~ 24.7 26.6 — 15.6 20.6
18 mo 19.3 20.1 22.8 23.9 23.1 25.5 21.2 24.7 - _ 25.2 26.3 16.9 19.6
24 mo 19.2 21.0 19.6 26.8 22.2 24.3 23.9 25.8 25.9 = = =
30 mo 22.5 - - = = - - —
36 mo 20.2 22.6 - - 22.3 = = = =
42 mo = = - - - - - 252.
48 mo = — 23.6 = - — =
54 mo = = - - - — =
60 mo = 27.8 - - - — -

The Roman numerals at the heads of the columns refer to the preceding 9 Tables.
Numbers in parentheses indicate the numbers of individuals marked in each size group.
The third line in the column headings indicates the respective size ranges for each class.

mental animals only very small numbers were recovered
on subsequent dives. Still, because of the narrowness of
the initial size range, the results seem to me to have some
value. One individual could not possibly have grown
more than one tenth of one millimeter in a full year,
while another could not have grown more than 5.1 mm.

Table 10: I have attempted to summarize the results
presented in the first 9 Tables, listing the maxima and
minima attained at 6 months intervals. In columns VIII

and IX the heavy line indicates the termination of the
study on July 13, 1968; for all other columns the bottom
line represents the final date.

In this representation the columns II through VII,
including all the size classes of the experiment started in
1961, present an especially interesting picture. Each size
class had last representatives (1 or more) at shorter and
shorter intervals; this could lead one to assume that in
the FCC Olivella biplicata dies when it has attained what

Table 11

Size Increases in Pooled Size Classes

{1537} {316}
(14.0-16.0) (16.1-18.0)
1 yr 17.6-21.7 21.0-25.8
2 ys 19.2-26.8 22.2—24.3
3 ys 20.2-22.6 -
4 ys - 23.6
5 ys 27.8

minimum and maximum increments at yearly intervals (in millimeters)

1 yr 1.6—7.7 3.0 — 9.7
2 ys 3.2-12.8 4.2-8.2
3 ys 4.2-8.6 —

4 ys — 5.6—7.5
5 ys

{167} {70} {109}
S200) (ON 2 220) en (2341-2420)
23.3 = 24.7-26.6
23.9-25.8 25.9
22.3
BGM = 0.7-35
S914) 3.95.8
VRAD

11.8-13.8 =

Page 266

THE VELIGER

Vol iisNows

appears to be its maximum size here, i.e. ca. 26 to 28
mm. Yet on April 22, 1961 an unmarked specimen meas-
uring 29.3 mm was found, indicating that in FCC Olivel-
la might attain as large a size as it does elsewhere and
that the 26 to 28 mm size is not the absolute maximum.
The possible explanation for the disappearance of the
“Jarge” specimens from the FCC may have to be sought
in the selective collecting by the shell jewelers.

Special mention may be made of the specimen listed
in column JI. It had survived 5 full years since the
marking.

Table 11: This Table was compiled by pooling all results
for each of the 5 different size classes used, in an attempt
to uncover possible general trends.

DISCUSSION

Darsy (1964) showed that in Tegula funebralis from
Sunset Bay, Oregon age can be estimated on the basis of
growth lines visible on the underside of the body whorl.
In the collection of the Department of Zoology, Univer-
sity of California in Berkeley, there is a large series of
the same species from many localities in California.
Close examination of these specimens reveals the presence
of the “small” incremental lines but does not show any
of the “larger” lines which Darsy equates to annual
growth lines, except in a few exceptionally large speci-
mens from Duxbury Reef. On the other hand, among an
almost equally large series of T. brunnea (Putippt, 1848)
I observed growth lines of a similar nature on a speci-
men from Tomales Bay, Marin County, California. I
hesitate to interpret the presence of the distinct lines in
Oregon material and the general absence of them in our
California material as a consequence of possibly more
rigorous climatic conditions in Oregon. However, DARByY’s
work did alert me to the possible significance of lines on
the shells of Olivella biplicata. By close examination of
the specimens that I know to have spent more than one
year in the FCC I hoped to discover any significant lines.
But there is no regularity in the lines discernible. The
general shell pattern is almost devoid of macroscopically
visible distinct lines and the few very distinct lines that
may be observed on an occasional individual either have
an irregular course or are so irregularly spaced that it is
hard to conceive of them as “annual” growth lines. The
irregularly shaped lines would seem to me to be an indi-
cation of a repaired lip and the irregularly spaced lines
might be indicative of fluctuations in diet. It cannot be
denied, of course, that the growth lines could give a clue
to the age of the individual. Unfortunately, our overall

knowledge of the biology of O. biplicata at present is in-
sufficient to answer this question.

It seems clear from a study of the various Tables that
different individuals of Olivella biplicata in the FCC grow
at different rates. The greatest increase in size in one year
is a possible 9.7 mm (from 16.1 to 25.8mm), while the
smallest is a possible 0.1 mm (or, conceivably, 0.0 mm).
It would be interesting to know if the differences in
growth rates are due to the sex of the individuals con-
cerned. No attempt had been made to determine the sex
of the individuals to be marked as this would have
required too much time. Dr. D.C. Edwards was kind
enough on the occasion of one of the “Olivella dives” to
show me his method of ascertaining the sex of an indi-
vidual; but even with his considerable experience it took
an average of 10 minutes per specimen.

The most interesting specimen of the entire series is
the one originally marked in July 1961 and recovered in
April 1967. In the intervening 5 years and 9 months this
animal almost doubled in size. Another specimen of al-
most equal interest is the one included in Table 9, for
which a possible increment of 0.1 mm in one full year
was reported.

Epwarps (1968) reports that Olivella biplicata ma-
tures sexually at about the time the shell attains a length
of 16mm. It would seem then, if this holds true for O.
biplicata in the San Diego region, that I picked out, with
one exception, sexually mature individuals for marking.
The consideration for picking the particular sizes was,
however, one of convenience in handling and marking.
The only exception was the group reported in Table 2. In
this case 347 individuals (perhaps) not yet sexually ma-
ture were marked; yet within 3 months they had appar-
ently attained a size at which sexual maturity may be
presumed. It may appear reasonable to allow a lapse of
2 or possibly 3 years, at the most, for O. biplicata to arrive
at a sexually mature stage from the veliger stage. But this
relatively narrow limit need not really apply when the
fact that one individual did not grow more than 0.1 mm
in one year is taken into account. It then becomes possible
to assume with some degree of justification that the sex-
ually mature stage is not reached before 4, 5, or more
years after metamorphosis. When this figure is added
to the actually observed time in the one specimen, i.e.
5 years, an estimate of a total life span, under ordinary
conditions, of 8 to 12 years would not appear exagger-
ated; and under exceptionally favorable conditions per-
haps an even longer life span may be expected. This
does not appear exorbitant in the light of the estimates
of 30 years, more or less, for Tegula funebralis made by
Darsy (1964).

Vol. 11; No. 3

THE VELIGER

Page 267

ACKNOWLEDGMENTS

In addition to the many individuals who have contributed
numerous hours in swimming, diving, or otherwise col-
lecting the experimental animals, I owe gratitude to Dr.
E. W. Fager who repeatedly made all the facilities of his
laboratory available to me for measuring and marking of
the shells as well as for maintaining the animals prior to,
during and immediately after the marking procedure.
The Department of Zoology, University of California,
Berkeley, supported the study throughout the several
years. Dr. Cadet Hand critically read the manuscript
and Mrs, Emily Reid drew the map and the text figure
accompanying this paper.

LITERATURE CITED

Darsy, RicHarp L.
1964. On growth and longevity in Tegula funebralis (Mollus-

ca: Gastropoda) . The Veliger 6, Supplement: 6-7;
pit. 1. (15 November 1964)
Epwarps, DALLas CRAIG
1968. Reproduction in Olivella biplicata. The Veliger
10 (4): 297 - 304; plt. 44; 3 text figs. (1 April 1968)
Keen, A. Myra
1937. An abridged check list and bibliography of west North
American marine Mollusca. Stanford Univ. Press, Stanford,
Calif.; pp. 1 - 88; 2 figs. (29 September 1937)

STOHLER, RUDOLF
1959. Studies on mollusk populations: IV. The Nautilus
73 (2): 65-72; 3 text figs.
1960. Studies on mollusk populations: IV. The Nautilus
73 (3): 95 - 103
1962. Preliminary report on growth studies in Olivella bipli-
cata. The Veliger 4 (3): 150-151; plt. 36 (1 Jan. 1961)
ZELL, CLARACE PLUMB Bock
1955. The morphology and general histology of the repro-
ductive system of Olivella biplicata (SowERByY), with a brief
description of mating behavior. Univ. Calif., Berkeley,
M.A. Thesis

Page 268

THE VELIGER

Vol. 11; No. 3

Distribution of Organic Bromine Compounds

in Aplysia californica Cooper, 1863

LINDSAY R. WINKLER

College of the Desert, Palm Desert, California 92260

(1 Text figure)

INTRODUCTION

THE EXISTENCE OF organic bromine compounds in the
lipids of Aplysia kurodai Baza, 1937 was first noted
during routine studies of marine organisms by TANAKA
& ToryAMA (1959). This report led indirectly to the pres-
ent exploratory study to determine the presence and dis-
tribution of bromine-containing organic compounds in
the California species, Aplysia californica Cooper, 1863.

The work of TANAKA & ToyAMA was introduced to
the non-Japanese speaking world by an article on the
chemical structure of 2 of the compounds by YAMAMURA
& Hirata (1963). In the first sentence of this latter
report the authors state “. . . but it is our experience
that the kind and content of bromo-compounds in A fly-
sia kurodai depends on when and where they were
gathered.” Since bromo-synthesization is rare in animal
metabolism the origin and function of such compounds
might be of interest.

As a sequel to this and other papers (WINKLER, 1959
and 1961; WINKLER & Dawson, 1963) an unsuccessful
attempt to locate a biological precursor for these bromine-
containing organic compounds in the sea weed Ploca-
mium pacificum (KyLIN) was reported by DarLinc &
Coscrove (1966).

Search of the recent botanical literature revealed ar-
ticles by a number of authors who reported the presence
of organic bromine compounds in a number of species
of the red algae. The most recent paper is that of
Craicig & Gruenic (1967), which, together with one by
Hopckin, Craicie, « McInnes (1966) has bibliograph-
ic references to most extant algal papers on the subject.
Among the red algae containing bromine organic com-
pounds reported by these authors were listed members
of the genus Laurencia, previously noted as the major
dietary item of some specimens of Aplysia californica
(WINKLER & Dawson, 1963).

The previously mentioned paper by YAMAMURA & HI-
RATA employed the term ‘aplysin’ in several combinations
to denote the bromine-containing lipids of Aplysia ku-
rodai. This term, however, had previously been employed
to refer to a different, water-soluble toxin found in the
digestive gland of A. californica (WINKLER, 1961 and
1962). As a result of this duplication of terminology, the
term ‘organo-bromines’ is used in the present paper to
avoid confusion.

MATERIALS ann METHODS

The previous Japanese papers reported only on extracts
made from entire, dried animals. However, previous ex-
ploratory work by the present writer has indicated that
the digestive gland was a main locus of lipids in Aplysia.
Preliminary studies of digestive gland lipids made on
celite columns led to the isolation of apolar fractions giv-
ing the Beilstein test for halogens. With this assurance of
halogen content the determination of relative content of
bromine in the various organs was begun using a semi-
quantitative paper chromatographic technique adapted
for the purpose.

Animals were collected at La Jolla and Doheny
Beach, California. Specimens were killed and dissected in
the laboratory under hypothermia and were usually di-
vided into the following parts: digestive gland, digestive
tract (including oral capsule), kidney, glandular mantle
margin, opaline gland, and the foot including total body
wall and dermis. Each organ was minced finely with scis-
sors and placed in the thimble of a flask-type lipid extrac-
tor and after thorough extraction with acetone the residue
was removed, dried, and weighed. After the acetone had
beenremoved from the extract inacurrent of air, the ex-
tract was cleansed of hydrophilic residues, after which it
too was weighed and the resultant 2 weights for each or-

Vol. 11; No. 3

THE VELIGER

Page 269

gan were added together to give the total dry weight. The
extract was then subjected to sodium fusion after which
the sodium salts produced were taken up in water, fil-
tered, and reduced to a quantity equivalent to the gram
weight of the extracted material, or multiples of that
weight if the amount was too small for convenient dilu-
tion or too much NaOH was present as a result of ex-
cess sodium being used during the sodium fusion.

Chromatograms made of 648 inch sheets of What-
man No. | filter paper were spotted along a line drawn
4inch from one of the long edges of the paper with
0.005 ml aliquots of the solution resultant from the so-
dium fusions. The papers were then formed into cylin-
ders, stapled, and stood in self-supporting position in
petri dishes containing the developing solvent. Each was
then covered with a gallon-capacity mayonnaise jar dur-
ing the development period. The developer giving best
separation between bromine and chlorine was pyridine
and water (90:10). After drying on an histological slide
warmer the spots on the chromatograms were made
visible by one of 2 methods. The first method involved a
silver nitrate spray followed by a 20 minute exposure to
fluorescent room lighting. The removal of excess silver
nitrate was accomplished by a wash in a weak nitric
acid solution and a water rinse. The alternate method
involved the use of a spray modified from FrEicy (1958)
made up of equal parts of glacial acetic acid, 30%
hydrogen peroxide and 5% uramine (sodium fluorescein).
After spraying, the chromatograms were placed on the
slide warmer set at maximum temperature of 65° C and
dried. The bright red color of eosin was formed wherever
bromine was present. After thorough airing, such chro-
matograms were made “permanent” by spraying several
times with ‘Krylon’ plastic spray. These chromatograms
are still bright after 9 months. While this method was
very specific for bromine, care was required in order to
prevent running of the water-soluble eosin dye. It was
consequently somewhat less accurate for quantitative
estimation than the less specific silver nitrate method
which depended entirely upon rf value for the final
identification of the spots.

In order to obtain the degree of quantitation required
each chromatogram was dried and the limits of the spots
were outlined with heavy pencil lines. The sheet was then
imprinted with a rubber stamp having } inch quadran-
gular rulings. The number of squares counted within each
spot outline provided a close estimate of the relative area
of each spot.

A standard curve was prepared by spotting chromato-
grams with 0.005 ml of standard dilutions of sodium

bromide, using 4 spots for each dilution. The areas of the
4 spots were averaged and the resulting figures were plot-
ted against quantity of bromine (not NaBr) actually
present. The resulting curve became the standard from
which quantitative estimates of the unknown spots were
determined. Experimental results were read from this
curve in grams of bromine in a 100 gram sample.

EXPERIMENTAL RESULTS

Table 1 summarizes the amounts of bromine found in
the various organs of a single small specimen taken from
La Jolla, California, in February 1966. Figure 1 illustrates

Table 1

Sample Analysis of Bromine Distribution
by the Silver Nitrate Method.

Results represent average values on five chromatographic determin-

ations for each tissue. Averages of four determinations for each

organ using the uramine/eosin method gave similar results but the
variability between runs was greater.

o 2 S
= de
=) cole : © 8 g
3 Ep) eae Fea
Organ Ee eee es Bb «
8 fal 2A o QO. Ss
Digestive gland 4.89 1.64 11.6 12.8
Body wall and dermis 6.65 2.74 0.2 5.48
Anterior digestive tract 1.12 0.20 Neg
Mantle margin 0.18 0.13 Neg.
Opaline gland 0.74 0.13 Neg.
Kidney 0.32 0.08 0.45 0.36
Total dry weight 13.9 gm 4.92 gm
Dermis ' 0.114gm 0.02gm _ Trace
Blood ! 6 ml Trace

' Results from different animals

the percentage of the total bromine found in each organ
in which it was demonstrable as well as the lipid distri-
bution in those organs. Other specimens taken on the
same collection trip and on subsequent trips to Doheny
Beach, California, indicated similar levels of bromine.
The levels given for blood and dermis represent runs on
blood and on skin scrapings from other individuals.
Amounts of extract from these samples were insufficient,
however, for successful quantitation.

Page 270

1 Lip Br
SS —._—-—_— ~~
Digestive Kidney Body Wall
Gland Complex
Figure 1

Graphic representation of the percentage of the total bromine-

organic compounds (Br) and lipids (Lip) in each organ in which

measurable amounts were found. Though the total quantity of

lipids was considerably greater in the body wall complex the

organo-bromides were found in relatively small quantities compared
with the reverse situation in the digestive gland.

DISCUSSION ann CONCLUSION

While bromine was clearly identified by rf value when
using the silver nitrate method, the specific reaction of
bromine with uramine to form the red dye eosin was a
further proof of its identity. The latter reaction appeared
quite spectacular against the yellow background of ur-
amine (sodium fluorescein) and even more pronounced
when viewed under long-wave ultraviolet light. Since no
color reaction was produced with chlorides when this
method was used, it was possible to use the less odorifer-
ous ethanol: water (diluted 90:10) as a developer. This
latter developer produced a nicer, rounder spot but this
was too close to the chloride spot for use when silver
nitrate (which was equally good as a chloride revealer)
was used for visualization. The uramine/eosin method

THE VELIGER

Vol. 11; No. 3

was less satisfactory for quantitative work, however, due
to running of the soluble eosin produced as opposed to
the highly stable spot size resulting from the use of the
silver nitrate precipitation.

The results of this preliminary study indicate that the
digestive gland is the principal site of organic bromine
compounds in the sea hare and that these materials are
probably excreted by the kidney and perhaps also by the
skin which has previously been suggested as an excretory
organ (MacMunn, 1899; WinKLEr, 1959). These sub-
stances may very well be transferred to the foot muscle
by way of the blood to become one of the predation-
deterrents employed by this mollusk which lacks an ex-
ternal shell. Furthermore, it may be inferred that the
brominated organic compounds are probably obtained
from the environment rather than being the result of
synthesis within the animal. The reports of various red
algae containing amounts of brominated organic com-
pounds and the fact that Aplysia californica is known to
include at least one genus of organo-bromine producing
algae in its diet (WINKLER & Dawson, 1963) along with
the concentration of bromo-organic compounds in the
digestive gland, makes the former hypothesis seem more
plausible, especially in view of the original statement
from YAMAMURA & Hirata (1963) quoted in the intro-
duction to this paper. On the other hand, its concentra-
tion in the digestive gland which is recognized as the
normal site of great numbers of enzymatic reactions
may indicate that it is there bromated as one of the
various, little-undertsood, detoxification processes.

The absence of the bromine compounds from the
opaline gland and the mantle margin would seem to
exclude the organo-bromine compounds from a part in
these particular predation-deterrent systems. However,
their presence in the body wall and skin may indicate that
they may function to make Aplysia repulsive to potential
predators.

The number of specimens used in this study is admit-
tedly small but is justified by the fact that the information
obtainable by this method involving the destruction of
the total acetone-extractable lipids is necessarily limited
to what we here report — the distribution of the organic
bromine fractions within the animal. Further study into
the ultimate origin of the brominated compounds and
their variation from animal to animal requires the frac-
tionation and identification of the apolar lipids and the
determination of the relative quantities of the different
bromine containing fractions. Such studies. usiny more
sophisticated equipment and methods, are now in the
planning and development stages.

Vol. 11; No. 3

THE VELIGER

Page 271

SUMMARY

Organs and organ systems of specimens of Aplysia cali-
fornica Cooper, 1863 were dissected out and extracts
were made with acetone. The residues after removal of
the acetone were subjected to sodium fusion. The result-
ant sodium salt was then taken up in water and semi-
quantitatively estimated by paper chromatography. Re-
sults indicated over 90% of the bromine-containing or-
ganic compounds were to be found in the digestive gland
and of the remainder more than # were found in the foot,
body wall and skin complex. Less than 0.3% were found
in the kidney. Trace amounts, unmeasurable by the
methods used, were found in the blood or hemolymph.

LITERATURE CITED

CraiciE, J.S. « D. E. GRuEnic
1967. Bromophenols from red algae.
Daruinc, STEPHEN D. & Ricwarp E. Coscrove
1966. Marine natural products. I. The search for Aplysia
terpenoids in red algae. The Veliger 8 (3): 178 - 180.
(1 January 1966)

Science 157: 1058

FEIcL, Fritz
1958. Spot tests in inorganic analysis. Elsevier Publ. Co.

Amsterdam, 5'* ed.

Hopcxin, J.H.,J.S. Craicre « A.G. McINNEs
1966. The occurrence of 2, 3-dibromobenzyl alcohol 4, 5-di-
sulfate, dipotassium salt in Polysiphonia lanosa. Canad.
Journ. Chem. 44: 74
MacMunv, C. A.
1899. The pigments of Aplysia punctata.
24: 1-10
Tanaka, TATSUO & YOSHIYUKI TOYAMA
1959. ‘Fatty oils of aquatic invertebrates: XXI Fatty oil of
Aplysia kurodat and its sterol components; XXII Unsaponi-
fiable components other than sterols in the oil of the sea hare
Aplysia kurodai. Nippon Kagaku Zasshi 8: 1326 - 1332
(in Japanese with English summary)
WINKLER, LINDSAY ROBERT
1959. A mechanism of color variation operating in the west
coast sea hare, Aplysia californica Cooper, Pacific Sci.
13 (1): 63 - 66; 1 text fig.
1961. Preliminary tests of the toxin extracted from California
sea hares of the genus Aplysia. _ Pacific Sci. 15 (2) : 211 - 214
WINKLER, Linpsay RosBErT & E. YALE Dawson
1963. | Observations and experiments on the food habits of
California sea hares of the genus Aplysia. Pacific Sci. 17 (1):

Journ. Physiol.

102 - 105
WINKLER, LinpsAy Ropert, BERNARD E. TILTON & MERVYN G.
HarDINGE
1962. A cholinergic agent extracted from sea hares. Arch.

Internat. Pharmacodyn. Thérap. 137: 76 - 83; 5 Text figs.
YAMAMURA, S. & YOSHIMASA HiraTA
1963. Structures of aplysin and aplysinol, naturally occurring
bromo-compounds. Tetrahedron 19: 1485 - 1496

Page 272

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Vol. 11; No. 3

Quantitative Relationships Between Gill Number,

Respiratory Surface, and Cavity Shape in Chitons

KAY M. JOHNSON |

(6 Text figures)

INTRODUCTION

CHITONS HAVE A NUMBER of characteristics which sug-
gest that they are a very primitive group of mollusks:
an ovoid shape, a broad, flat foot, shells of only two
layers — the tegmentum and the articulamentum — , a
microphagous, herbivorous mode of feeding, a non-gang-
lionated nervous system, and a trochophore larva which
metamorphoses directly into the adult form. Yet, with
all of these primitive characteristics they are not termed
an “ancestral mollusk” type. That is largely because of
the many gills found in the grooves along the sides of
the foot.

YoncE (1939, 1947) describes an “ancestral mollusk”
with a posterior mantle cavity with only 2 gills. He
believes that the chitons evolved from this ancestor due
to a flattening of the animal with an extension of the
pallial cavity in grooves alongside of the body. The
gills were seemingly forced into multiplying in number
and shortening so as to fit into the grooves.

Until recently YonceE’s hypothesis was not challenged.
With the discovery of Neopilina galathea LEMcHE, 1957,
a doubt as to the complete validity of YoncE’s hypothesis
was raised. Neopilina was obviously a very primitive
mollusk with its ovoid shell, flat foot, and radula. But the
most astonishing fact was that it had a series of gills
along the side of the foot. To LEMcHE (1957) this was
enough to prove that multiplicity of gills was the prim-
itive condition of the molluscan pallial cavity and also
enhanced the idea that mollusks had a segmented ances-
tor. So, from LEMcHE’s hypothesis it could be concluded
that the gills in chitons are arrayed like the primitive
condition and that the number of gills in other groups
of mollusks is a reduction.

Only Hunter & Brown (1965) have tried studying
the gills in chitons in an attempt to show with data just
which theory might be reasonable. HUNTER & BRowN
studied only the relationship between the weight-length

and the number of gills. Their results showed what they
considered too great an asymmetry between the number
of gills on each side of animals of equal weight-length
for there to be any basis for the idea that chitons are
related to a segmented ancestor. And, therefore, the con-
dition of the gills was not even possibly like the primitive
state. In other words, they decided that the gills were a
secondary replication of structures of the ‘ancestral mol-
lusk’ as YoncE had hypothesized it.

Hunter & Brown may have been correct in conclud-
ing that the asymmetry in gill numbers may show a lack
of segmentation. But that does not necessarily mean that
a multiplicity of gills was not a primitive condition of

- the mollusks. There are 3 groups which are very primi-

tive mollusks and do have numerous gills — Neopilina,
Nautilus, and the chitons. So, why could not the primi-
tive condition have been a “mantle cavity as a groove
bounded by the mantle edge and surrounding the head-
foot rather than a posterior cavity”? (FRETTER & Gra-
HAM, 1962).

HunrtTeER & Brown used only 2 species, Chaetopleura
apiculata (Say, 1834) and Lepidochitona cinereus (Lin-
NAEuS, 1767). And they only counted the number of
gills. I have sought to further clarify that picture of the
chiton gill cavity by making additional measurements.

MATERIALS ann METHODS

Various measurements of a chiton’s gill cavity assist in
portraying the gills in relation to the whole animal:
(1) the length of the animal along the foot; (2) lengths
of the gill series on both sides; and (3) the length of
the most posterior gill itself. Also, I counted the number
of gills on both sides of the animal. And by likening the
effective gill surface (that which water must pass through
during respiration) to a triangle I could compute that
surface area by using the length of the last gill and the

Vol. 11; No. 3

length of the gill series in the formula: 4bh = area of
a triangle.

These various measurements can be used for a descrip-
tion of the gill cavities of a group of chitons with a wide
range of types, sizes, and shapes. I used chitons from 2
suborders and 4 families of different adult lengths and
shapes, and of a range of sizes within each species. The
species I used are further described below:

Acanthochitonina
ACANTHOCHITONIDAE

1. Acanthochitona exquisita Pitspry, 1893, is a small
(1.1 to 3.3. cm) oblong chiton.

Lepidopleurina
ISCHNOCHITONIDAE

1. Stenoplax (Stenoradsia) magdalenensis (Hinps, 1844)
is a long (2.1 to 9.0cm), rather narrow chiton.

2. Tonicella lineata (Woop, 1815) is a small chiton (1.8 to
4.4 cm). Its general shape is a gently rounded oblong.

MoPALIDAE

1. Mopalia muscosa (Goutp, 1846) is a large (1.1 to 8.1
cm) oval chiton.

2. Placiphorella velata DAL, 1879 is a small (0.9 to 2.5
cm) broad, oval, flattened chiton.

CHITONIDAE

1. Chiton sulcatus Woop, 1815 is a large (0.5 to 7.5 cm)
chiton which is fairly broad and oval.

All of the specimens were preserved in the collection

at the Department of Invertebrate Zoology, California
Academy of Sciences, San Francisco. So, I tried to look
at a range in sizes of each species of chiton that had
come from one collecting area. In that way I could pos-
sibly be working on a growth series of a species. There-
fore, I chose the following lots:
100 Acanthochitona exquisita from Puerto Refugio, An-
gel de la Guarda Island, Gulf of California; 25 Stenoplax
magdalenensis from Punta Abreojos, Baja California; 92
Tonicella lineata from False Bay, San Juan Island, Puget
Sound, Washington; 20 Placiphorella velata from Cedros
Island, Mexico; 40 Chiton sulcatus from near and in
Academy Bay, Santa Cruz Island, Galapagos Islands.

Mopalia muscosa was the exception. The Academy did
not have a large number of animals from any one place.
Therefore I took groups which had been collected along
the range of this species. Thus, 17 M7. muscosa were from

RE WWEEIGER

Page 273

False Bay, Washington; 8 from near San Simeon, Cali-
fornia; and 10 from one mile north of Camalu Arroyo
near Guerrero, Baja California.

RESULTS

Just as HUNTER & Brown (1965) showed in Chaeto-
pleura and Lepidochitona, there is an asymmetry of the
right and left sides in the chitons studied. It is evident
when one compares the average number of gills per side
with the length of an animal, that the right and left
sides are not similar. Growth seems to be random. This
is easily seen by comparing the gill numbers on either side
of Tonicella lineata in Figures 1 and 2. Thus, it appears
that HUNTER « BRown’s work is correct.

30
es
S 28
a
2
g
3
Zz

26

24

2. 4
3 2 Length (cm)
Figure 1

Graph showing the number of gills found on the right side of
Tonicella lineata of different lengths.

But much more is evident from my data, as may be
seen in Figure 3. Those chitons which eventually reach a
“large” adult size (8 to 9cm) have more gills than do
those which attain only a “small” adult size (1 to 5cm).
The difference is quite obvious when one compares ani-
mals of the same sizes, between 0.5 and 5.0 cm, but of
different adult sizes. Thus, some may have adult stages
larger than 5 cm and some may not. The “large” chitons
are from at least 2 species and 2 different families, yet

of Gill

Page 274

30
3 28.
es
o
2
A 06
24 tt ____—_—_1—
2.3 3 4
Length (cm)
Figure 2

Graph showing the number of gills on the left side of Tonicella
lincata. A comparison with Figure 1 demonstrates the lack of
symmetry between the two sides.

ihey possess on the average from 4 to 10 more gills per
side than do the “small” chitons (which include even
representatives of 2 different suborders). Thus, there is

“Small Chitons
——-— “Large” Chitons

==--=-= Chiton sulcatus

50 ns Set NY
;

I 2 3 4 5
Length (cm)

Figure 3

chitons have more gills than “small” chitons at all lengths

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Vol. 11; No. 3

some major difference between “large” and “small” chi-
ton growth.

Chitons of different adult size have the same effective
gill surface per unit length. This is found by comparing
the gill cavity area with the length as in Figure 4.

ax BY
Oo
&
oO
~~
3
o
~
<
ip
a
2)
)
.06
(0)
1.2 2 3

Length (cm)

Figure 4

Both “large” and “small” chitons have equal gill surface area per
unit length

The difference between “large” and “small’ chitons
is found when one compares the gill series lengths with
the body lengths, and the length of the last gill with
the length of the gill series. It appears that the length of
the gill series of the “large” chitons are somewhat longer
than in the “small’ chitons, even when both groups are
of the same lengths (Figure 5). And the “small” chitons

2.0

“Small Chitons
—— — “Large” Chitons ne /

& es)

Length of Last Gill (cm)
5

1.2 2 3
Length (cm)
Figure 5

“Large” chitons have longer gill series than “small” chitons

Vol. 11; No. 3

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Page 275

achieve equal gill cavity area by lengthening their gills
more than the “large” chitons do (Figure 6). Thus, the
most posterior gill of the “small” chitons are longer than

“Small Chitons
— — — “Large” Chitons

Gill Series Length (cm)

Length (cm)

Figure 6

“Small” chitons have longer gills than “large” chitons

the last gill of “large” chitons of equal length. In other
words, the “small” chitons have shorter, wider gill cavi-
ties than do the “large” chitons.

DISCUSSION

Hunter & Brown (1965) may be correct regarding the
non-segmented nature of an “ancestral mollusk” type. In
fact, my results seem only to support that part of their
arguments. The right and left gills of chitons are never
completely equal in number.

Deciding that the “ancestral mollusk” was not segmen-
ted does not necessarily support YoONGE’s hypothesis of a
two-gilled ancestor. The additional information I have
supplied about the chiton gill cavity adds a new perspec-
tive to the problem. If the many gills in chitons are due
to a forced multiplication of them due to the chitons’
being flattened over evolutionary history, why do “large”
and “small” chitons differ? If each type, “large” and
“small”, was achieved separately, why do such similar
animals, as those found in the same family, have differ-
ently shaped gill cavities? Thus, it seems that YONGE’s
hypothesis fails to account for these differences in gill
cavity shapes.

A simple and more adequate explanation would be
that the multiplicity of gills as seen in Neopilina or the
chitons may be like the primitive condition of the mol-
luscan gills. It seems much more plausible that the two
trends seen in the “large” and “small” chiton gill cavities
were present in the “ancestral mollusk” and have been
maintained by the chitons, than that they were the result
of a couple of stages of flattening of the animals. Thus,
the two gills found in gastropods and lamellibranchs and
the four gills in some cephalopods are most probably the
product of a reduction in number of gills with an en-
largement of the posterior part of the gill cavity. This
reduction in the number of gills would, of course, oblit-
erate the pattern of “small” and “large” animal gill
areas. Therefore, it appears that the chitons have main-
tained a condition found in the “ancestral mollusk” type,
although more gills may have been added, since the
chitons have so many more gills than does the more
primitive Neopilina.

SUMMARY

Chitons of “large” adult size have more gills present on
both sides of the animal at all sizes than do the “small”
adult-size chitons. The difference in number does not
affect the effective gill surface per unit length. The ef
fects of the difference in number on the area of gills are
equalized in the “small” chitons by possession of longer
gills. Therefore, the “large” chitons have long narrow
gill cavities, while the “small’ chitons have short wide
gill cavities.

These two trends in the chitons do not seem to be
adequately explained by YonceE’s hypothesis as to the
origin of the multiplicity of gills in chitons. A more ade-
quate explanation is that the trends were present in some
“ancestral mollusk” type and have been maintained in
the chitons and in Neofilina. All other mollusks have,
therefore, modified this primitive gill cavity.

ACKNOWLEDGMENTS

I would like to thank Dr. Michael T: Ghiselin of the
Department of Zoology, University of California, Ber-
keley, for his kind assistance throughout this project and
for criticizing the manuscript. Also, I would like to
thank Mr. Allyn G. Smith of the California Academy of
Sciences for his help and for allowing me to use the col-
lection of preserved chitons in his care.

Page 276

THE VELIGER

LITERATURE CITED

FRETTER, VERA & ALASTAIR GRAHAM
1962. _ British prosobranch molluscs, their functional anatomy
and ecology. London, Ray Soc. xvi+/55 pp.; 316 figs.
Hunter, W. RusseE.i « STEPHEN C. BROWN
1965. | Ctenidial number in relation to size in certain chitons
with a discussion of its phyletic significance. Biol. Bull.
128 (3): 508 - 521 (June 1965)
LemMcHE, HENNING « Kart GEorG WINGSTRAND
1957. The anatomy of Neopilina galathea LemcuE, 1957
(Mollusca: Tryblidiacea) M. Galathea Rep. 3: 9-71
SmitH, ALLYN GoopwIN
1960. Amphineura Jn Treatise on invertebrate paleontology:
Mollusca I 1. 41 - 84; figs. 31-50 Raymond C. Moore, ed.
Univ. Kansas Press
YONGE, CHARLES MAURICE
1939. On the mantle cavity and its contained organs in the
Loricata (Placophora). Quart. Journ. Micro. Sci. 81 (3) :
367 - 390 (June 1939)
1947. The pallial organs in the aspidobranch gastropoda and
their evolution throughout the Mollusca. Phil. Trans. Roy.
Soc. London, B 232: 443 - 518 (22 April 1947)

Vol. 11; No. 3

Vol. 11; No. 3

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Page 277

Recognition of an Eastern Pacific Macoma

in the Coralline Crag of England

and its Biogeographic Significance

BY

EUGENE V. COAN

Department of Biological Sciences, Stanford University
Stanford, California 94305

EvIDENCE HAS BEEN ACCUMULATING over the last few
years that Bering Strait was open during the late Mio-
cene, closed through most of the Pliocene, then open and
closed several times from the late Pliocene through the
Pleistocene. These submergences allowed the exchange of
marine life, chiefly from the Pacific to the Atlantic O-
ceans (MacNem, 1965; Hopkins, 1967).

Pacific elements in the Coralline Crag of England, of
Astian (Pliocene) age (BADEN-PowELL, 1960), may rep-
resent either late arrivals from the late Miocene submer-
gence (Hopkins, personal communication) or early
arrivals from the late Pliocene submergence (Zullo, per-
sonal communication). The larger influx (in terms of the
number of species) of Pacific elements present in the
Red Crag (Pleistocene) of England and similar deposits
on the mainland of Europe and in Iceland may represent
either an event of submergence in the Bering Strait area
or similar tectonic events in the Canadian archipelago
(Hopkins, 1967).

In connection with a revision of the Eastern Pacific
Tellinacea, I can now report further evidence concerning
the trans-Arctic migrations of mollusks. The boreal spe-
cies of the genus Macoma, with a geological record from
the Eocene to the Recent in the Eastern Pacific (KEEN
& Bentson, 1944), have recently been discussed as not
having reached the Atlantic Ocean until the Pleistocene
influx (DuRHAM & MacNeit in Hopxtins, 1967). I find
that M. obliqua (Sowersy, 1817)', reported from the
Coralline Crag of England (Woop, 1848, 1874; British
Museum [Natural History|], 1963) and from the correl-
ative, Scaldisian strata in Belgium (Gutpert, 1958a, b),
is conspecific with a Recent West American boreal spe-
cies, commonly identified with M. incongrua (von Mar-
TENS, 1865)’.

Macoma lyelli Dati, 1894, described from the late
Miocene or early Pliocene of Marthas Vineyard, Mas-
sachusetts (also Dati, 1900b), and M. cookei GARDNER,
1943, described from the Upper Miocene of Virginia
seem to be closely related.

As Recent and fossil Eastern Pacific specimens of
Macoma obliqua differ significantly from Recent material
from Japan, type locality of M. incongrua, the Western
Pacific form should be regarded as a distinct subspecies
or species. Macoma obliqua has become extinct in the
North Atlantic since the Pleistocene.

' Tellina obliqua J. Sowersy, 1817, non Woop, 1815. The Inter-
national Commisson on Zoological Nomenclature has been pe-
tioned to conserve the name of this well-known Cenozoic fossil
which has only recently been discovered to be a junior homo-
nym of an unimportant junior subjective synonym. Type spe-
cimens of the Sowerby species are in the British Museum
(Natural History), and Stanford University now has specimens
from the Red Crag which have been compared with this type
material.

2 Tellina incongrua voN Martens, 1865. A potential lectotype,
measuring 25.4mm in length, is in the Zool. Mus., Humboldt
Univ., Berlin, no. 7624. Synonyms appear to be Tellina nasuta
truncata MuippENporFF, 1851, non Linnarus, 1767, and T:
nasuta brevior SCHRENCK, 1867. Macoma frigida (HANLEY,
1844), described from Kamchatka, seems to be a closely-related,
but distinct species.

Dati (1900a) suggested that Tellina rotundata SoweErRBy,
1867, might be a synonym of Macoma incongrua. The type
specimen in the BM(NH) proves to be M. balthica (LINNAEus,
1758). He also suggested that M. californiensis Bertin, 1878,
was a synonym. Photographs of the type specimens of the
latter, kindly provided by I’Ecole des Mines, Paris, prove these
to be Macalia bruguiert (Hantey, 1844), mislabeled as to
locality, for the species is Asian.

Page 278

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Vol. 11; No. 3

The synonymy of Macoma obliqua and the West A-
merican M. incongrua of authors, foreshadowed by the
association of the two names in the Pleistocene of Iceland
(Erarsson, Hopkins, & Doet in Hopxins, 1967), sug-
gests that migrations from the Pacific to the Atlantic
during the first Cenozoic submergence of Bering Strait in
the late Miocene may have been more extensive in terms
of the number of species involved than previously
thought (as listed by DuRHAM & MacNetL in Hopkins,
1967). West American paleontologists and marine mol-
luscan systematists will have to take into account this
exchange and the resulting nomenclatural involvements
with European and Atlantic fossil species.

I would also suggest that the isolation created by the
Pliocene land bridge subsequent to the late Miocene sea
passage may partially explain the presence of so many
boreal species of some genera, such as Macoma, in the
North Pacific Basin. For instance, M. middendor ffi Dat,
1884, a related Bering Sea species, with published records
in the Miocene, Pliocene, and Pleistocene of the North
Pacific, may represent the population which remained
in the Pacific Ocean when M. obliqua traveled to and
became isolated in the Atlantic-Arctic in the Pliocene.

A preliminary survey of literature indicates that an-
other, now extinct species of boreal Macoma may have
reached the Atlantic as early as the European Anversian
(Miocene). Taxa that ought to be compared with one
another in order to prove this are M. albaria (Conrap,
1849), M. virginiana (Conrap, 1866) (and its subspe-
cies), and M. elliptica (Broccui, 1814) of GutBERT
(1958a, 1958b) and others.

LITERATURE CITED

BapEN-PoweELL, D. F W.
1960. On the nature of the Coralline Crag.
CH (Bis WB = ey
BErTIN, VICTOR
1878. Révision des tellinidés du Muséum d’Histoire Natu-
relle .... Nouv. Arch. Natl. Mus. Hist. Nat. Paris (2)
1: 201 - 361; plts. 8-9
BritisH Museum (Natura History)
1963. British Caenozoic fossils, 2%” ed.
Mus.) pp. i- vii+1 - 133; plts. 1 - 44
Broccu1, GIovANNI BATTISTA
1814. Conchiologia fossile Subapennina con osservazioni geo-
logiche sugli Apennini e sula suolo adiacente. Milano (Dalla
Stamp. Reale) 1: 1-56 + i-Ixxx + 1-240; 2: 241-712;
plts. 1 - 16
Conrapb, TimotHy ABBOTT
1849. Fossils from northwestern America. In J.D. Dana
“Geology: United States Exploring Expedition, during the
years ... , under the command of Charles Wilkes, ... .”
10: 723 - 728; plts. 17 - 21
1866. Note on the genus Gadus, with the descriptions of some

Geol. Mag.

London (Brit.

new genera and species of American fossil shells. Amer.
Journ. Conch. 2 (1): 75-78 (1 January 1866)
DALL, Wi1LLt1aAM HEALEY
1884. | Report on the Mollusca of the Commander Islands,
Bering Sea, collected by Leonard Stejneger in 1882 and 1883.
(Contributions to the history of the Commander Islands, no.
3). Proc. U.S. Nat. Mus. 7 (22) : 340 - 349; plt. 2
(3 October 1884)
1894. | Notes on the Miocene and Pliocene of Gay Head,
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the Ashley River district, South Carolina. Amer. Journ.
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American species. Proc. U.S. Nat. Mus., 23 (1210): 285
to 326; plts. 2-4 (14 November 1900)
1900b. Contributions of the Tertiary fauna of Florida with
especial reference to the Silex Beds of Tampa and the Plio-
cene beds of the Caloosahatchie River including in many cases
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Garpner, JULIA ANNA
1943. Mollusca from the Miocene and Lower Pliocene of
Virginia and North Carolina. Part I. Pelecypoda. U.S.
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1958a. ‘Tableau stratigraphique des mollusques du Neogene de
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1958b. Pelecypodes du Diestien, du Scaldisien et du Merxe-
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1967. ‘The Bering land bridge.
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1944. Check list of California Tertiary marine Mollusca.
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Stanford, Calif. (Stanford

2 tab. (30 August 1944)
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1758. Systema naturae per regna tria naturae ... editio

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1767. Systema naturae per regna tria naturae ... editio
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MacNem, Francis STEARNS

1965. Evolution and distribution of the genus Mya, and Ter-
tiary migrations of Mollusca. U.S. Geol. Surv. Prof.
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MippEnporFF, ALEXANDER THEODOR (VON)

1851. Mollusken Jn “Dr. A. Th. v. Middendorff’s Reise in
den dussersten Norden und Osten Sibiriens” 2 [Zoologie] (1-
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THE VELIGER Page 279

ScHRENCK, LEoPoLD IvANovICH (voN) VON MartENs, EDUARD
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pits. 12 - 30 (post-October 1867)
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1867[1866 - 1869]. Monograph of the genus Tellina In Con-
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dated.

SoweErRBY, JAMES

1817 [1815 - 1818]. The mineral conchology of Great Britain; or
coloured figures and descriptions of those remains of testaceous
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and depths in the earth. London (privately printed) 2:
1-251; plts. 103 - 203

Woop, SEARLES V.

1848. A monograph of the Crag Mollusca, with descriptions
of shells from the upper Tertiaries of the British Isles 2 (Bi-
valves) . Palaeo. Soc. 9: 217 - 342; plts. 21 - 31

1874. Supplement to the monograph of the Crag Mollusca,
with descriptions of shells from the upper Tertiaries of the
east of England. Palaeo. Soc. 27: 99-231; plts. 8-11
+1 more

Woop, WILLIAM
1815. | General conchology; or, a description of shells, arranged

according to the Linnean system, and illustrated with plates
drawn and coloured from nature, .. . 1. London (John

Booth). [only one volume ever issued]: i-lxit1-7+1- 246;

pits. 1-59, 4* (issued in as-yet undated parts, from 1814 to
1815)

Page 280

THE VELIGER

Vol. 11; No. 3

NOTES & NEWS

A Color Variation of Aldisa sanguinea

BY

RICHARD A. ROLLER
1127 Seaward Street, San Luis Obispo, California 93401

(1 Text figure)

COLLECTION AND OBSERVATION of a fairly large number
of specimens of the nudibranch Aldisa sanguinea (Coo-
PER, 1862) over a 2-year period in San Luis Obispo
County, California, have shown that this species commonly
has varying amounts of yellow pigment on the dorsal
surface. This pigment tends to be arranged in repeated
patterns, frequently in the shape of a “T”.

Color descriptions of Aldisa sanguinea given in the
literature by Baza (1949), Cooper (1862), MAcFARLAND
(1905, 1906, 1966), Marcus (1961), and O’DoNocHUE
(1927) indicate a body color of red, in varying shades,
accompanied generally by two black spots on the median

line of the dorsum. No mention is made in the literature.

of any yellow pigment or distinctive yellow pattern.

Pruvot-Fou (1954, p. 268; fig. 106) pictures a “T”-
shaped marking for A. banyulensis Pruvot-Fo., 1951,
a red species with whitish stripes. BaBa (1949, plt. 25,
fig. 94) shows a figure of Halgerda rubicunda Basa, 1949
with a similar “T-shaped marking, except that the “T”
points in an anterior direction, rather than a posterior
one. The animal is described (op. cit. p. 135) as “Gen-
eral body colour orange-red, a short longitudinal band
of dull yellow medianly between the rhinophores, and
there is another, but transverse, band of the same colour
a short distance in front of the branchiae.”

To date 27 specimens of Aldisa sanguinea (from 11 to
35 mm in length) have been collected or observed alive
by the author. Of these animals, 17 had an obvious
chrome-yellow “T”’-shaped marking on the posterior half
of the dorsum. This marking is made up of a dense
series of fine yellow dots forming an irregular band across
the dorsum, just ahead of the posterior black spot. This
yellow pigment disappears completely from the median
area of the dorsum. A similar yellow band extends from
the branchial area, posteriorly along the median line to
the mantle edge (Figure 1).

Of the 27 animals observed, all but 2 exhibited a
transverse yellow band across the dorsum, and only one

animal had no observable yellow pigment. Many of the
animals were heavily marked with yellow pigment, scat-
tered in varying amounts, over the anterior or posterior
regions of the dorsum, or both, the branchial plumes,
and the anal area inside of the branchial plumes.
Continued collection of specimens with variations from
the usually reported color tends to make this color vari-
ety the expected one for this area. Baba (personal com-
munication) states that he has not seen a single “T”-
shaped marking in Japanese specimens of Aldisa sangui-

Figure 1

Aldisa sanguinea (Cooper, 1862)
Stippling indicates yellow pigment

nea. No collectors that I have contacted have seen this
color variation anywhere except in San Luis Obispo
County; however, a color slide by Mr. Robert Ames of
Oakland, California, shows 2 specimens with transverse
yellow bands. It is hoped that anyone collecting specimens
of A. sanguinea will look for similar markings and pig-
mentation; and I would appreciate any information on
the color pattern of this species.

I acknowledge the assistance of Mr. Steven J. Long
and Mr, Gary McDonald in obtaining specimens. I also
appreciate the aid and advice of Mr. James R. Lance
in preparing the manuscript.

Vol. 11; No. 3

LITERATURE CITED

Basa, K1KUTARO
1949. Opisthobranchia of Sagami Bay, collected by his Majesty
the Emperor of Japan. 4, 2, 194, 7 pp., plts. 1-50 (in
color), 161 text figs. Iwanami Shoten, Tokyo.
Cooper, JAMES GRAHAM
1862. Some genera and species of California Mollusca.
Proc, Calif. Acad. Nat. Sci. 2: 202 - 207
MacFaruanp, FRANK Mace
1905. A preliminary account of the Dorididae of Monterey
Bay, California. Proc. Biol. Soc. Washington, 18: 35 - 54
1906. | Opisthobranchiate mollusca from Monterey Bay, Cali-
fornia, and vicinity. Bull. U.S. Bur. Fish for 1905, 25:
109 - 151; plts. 18 - 31
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Marcus, Ernst
1961. Opisthobranch mollusks from California. The
Veliger 3 (Supplement, pt. I) : 1-85; plts. 1-10. (Feb. 1, 1961)
O’ DONOGHUE, CHARLES HENRY
1927. Notes on a collection of nudibranchs from Laguna
Beach, California. Journ. Entom. Zool. Pomona College
19: 77-119; plts. 1-3
Pruvot-Fox, ALICE
1951. Etudes des nudibranches de la Méditerrannée (2
partie) . Arch. Zool. Expér. Génér. 88: 1-80; 42 figs.;
pits. 1-4
1954. | Mollusques opisthobranches. in Faune de France, 58:
460 pp.; 1 plt.; 173 text figs.; Paris, Paul Lechevalier.

Two New Records of Cratena abronia

BY

STEVEN J. LONG
126 Esparto Avenue, Pismo Beach, California 93449

On June 11, 1968, a single specimen of Cratena abronia
MacFar.anp, 1966 was collected by the author at Dino-
saur Cave in the north end of Pismo Beach, San Luis
Obispo County, California. The specimen was found in
the mid-tide zone on Desmarestia herbacea (TuRNER,
1808). The specimen measured 5mm in length, and
matched well MacFar.anp’s description and illustration
of the species. Mr. Richard A. Roller, of San Luis Obis-
po, California, made a series of color transparencies of
the animal and extracted the radula to permit corrobora-
tion of the identification. The radula has been preserved
in alcohol.

On June 12, 1968, a second Cratena abronia was taken
at the south end of Sunset Palisades, about 2 miles north
of the first collecting point. This specimen was found in
the same tidal region, also on Desmarestia herbacea. This
animal was in poor condition, having lost several cerata.

THE VELIGER

Page 281

It measured 5 mm in length when alive; it was preserved
in alcohol for later study.

Cratena abronia was described from several speci-
mens collected at Point Pinos, Monterey Bay, California
(36°38’ N; 121°55’ W). Pismo Beach is located at 35°09’
N and 120°38’ W. This constitutes a southward range
extension of about 230 miles.

LITERATURE CITED

MacFarianp, Frank Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)

What is Macoma truncaria DALL?

BY
EUGENE V. COAN

Department of Biological Sciences, Stanford University
Stanford, California 94305

(1 Text figure)

WHILE STUDYING TYPE SPECIMENS of West American
tellinaceans at the United States National Museum, I
photographed and sketched the unique holotype of Ma-
coma truncaria Dati, 1916, described from between
Cape Halkett and Garry Creek [as “River’] on the
Arctic coast of Alaska. A brief synonymy is as follows:

Datt, 1916a: 37 [nomen nudum]

Dati, 1916b: 414

Dati, 1921: 48

O.proyp, 1925: 177

Burcu, 1945: 15

Figure 1

“Macoma” truncaria DaLL, 1916, holotype, USNM 210916, X 2.4
Drawing by Mr. Perfecto Mary

Page 282

THE VELIGER

Vol. 11; No. 3

The holotype, USNM 210916, a broken pair with only
the left valve complete, measures 15 mm in length and
proves to be a small, thickened specimen of Thracia
(Crassithracia). It seems closest to T. (C.) beringi Dat,
1915, but the correct allocation would require careful
study by one more familiar with the several circum-Arc-
tic species of the genus than I. A line drawing is here
provided for the use of other workers.

LITERATURE CITED

Burcu, JoHNn Quincy (ed.)
1945. Superfamily Tellinacea.
Calif. no. 43: 3 - 33
Dai, WituiamM HEALEY
1915. A review of some bivalve shells of the group Anatina-
cea from the west coast of America. Proc. U.S. Nat. Mus.
49 (2116): 441-456 (27 November 1915)
1916a. Checklist of the Recent bivalve mollusks (Pelecypoda)
of the northwest coast of America from the Polar Sea to San
Diego, California. Los Angeles, Calif. (Southwest Mus.).
pp. 1 - 44; 1 portrait (28 July 1916)
1916b. Diagnoses of new species of marine bivalve mollusks
from the northwest coast of America in the collection of the
United States National Museum. Proc. U.S. Nat. Mus.
52 (2183): 393 - 417 (27 December 1916)
1921. Summary of the marine shellbearing mollusks of the
northwest coast of America, from San Diego, California, to
the Polar Sea, mostly contained in the collection of the
United States National Museum, Smithson. Inst., U.S.
Nat. Mus. Bull. 112: 1-217; plts. 1-22 (24 February 1921)
O.proyp, Ipa SHEPARD
1925. The marine shells of the west coast of North America
1 [Pelecypoda]. Stanford Univ. Publ., Univ. Ser. Geol. Sci.
1 (1): 1-247; pits. 1-57 (September 1925)

Min. Conch. Club South.
(January 1945)

Spawning Notes, II. - Mitra dolorosa

BY

FAY H. WOLFSON

San Diego Natural History Museum,
San Diego, California 92115

(3 Text figures)

No spAWNING BY Mitra (Strigatella) dolorosa DAtt, 1903
had been noted during field trips to Bahia de los Angeles,
Baja California, Mexico in April, August, September and
December 1967 or in January and April 1968. However,
on 23 July 1967 and again on 25 May 1968, I observed
spawning at approximately mid-tide level during low tide.
In both cases, eggs were being deposited on a small stone
situated beneath larger overhanging rocks. There were 2

Figure 1

Figure 2

Detail of capsule, top view

Figure 3

Detail of capsule, side view

individuals on the stone in July and 5 on the stone in
May, although in each case only a single individual ap-
peared to be in the process of oviposition. Approximately
50 capsules, forming an irregular ellipse, had been ex-
truded in both instances.

The translucent sausage-shaped capsules rest lightly
against each other. Each tapers to a thin stalk which
flares into a basal membrane by which the cluster is
attached to the substrate (Figure 1). The capsules aver-

Voll I Nos 3

age 1.04 < 2.16 mm in size, with a range of from 0.88 x
1.91 mm to 1.17 & 2.35 mm.

An oval inner membrane completely fills the “sau-
sage” and contains approximately 100 opaque, cream-col-
ored eggs, many of which bear a small cap of darkly
pigmented cells. The eggs average 0.147 mm in greatest
diameter and appear to be distributed in random clumps.

Figures 2 and 3 show the raised, thickened structure
situated at or near the top of the capsule. Rupturing of
the thin membrane between the “lips” in the center of
the structure presumably provides an exit at hatching
time.

The figures were drawn by Anne Acevedo of the San
Diego Museum of Natural History.

Littorina littorea in California

(San Francisco and Trinidad Bays)

BY

JAMES CARLTON
521 Mandana Boulevard, Oakland, California 94610

A SINGLE LIVING SPECIMEN of the common Atlantic peri-
winkle, Littorina littorea LinNaEuS, 1758 was collected
by the writer at Berkeley, in San Francisco Bay, Califor-
nia, on July 11, 1968, approximately 300 m north of the
foot of Ashby Avenue, along the East Shore Highway.
The animal was in a rock crevice, and despite an hour’s
search, no additional specimens were located. The pres-
ent specimen measures 15.7 mm in length and 13.8mm
in width,

Littorina littorea is known from the Pacific coast of
North America through the collection of at least 5 speci-
mens in Deception Pass, Puget Sound, Washington, in
1937 (Hanna, 1966). The hypotype illustrated by Han-
NA was compared with the present specimen and its
identification confirmed by Dr. Leo G. Hertlein of the
California Academy of Sciences. All 5 specimens from
Washington bear opercula, indicating live collection.

In addition, there are 2 specimens in the collections
of the California Academy of Sciences (cat. no. 39464)
from Trinidad Bay, California (just north of Humboldt
Bay), collected alive by Dr. Doris K. Niles on July 28,
1942, but not received at the Academy until 1965, too
late for inclusion in Dr. Hanna’s paper. The specimens
measure 27.0mm in length by 21.4 mm in width, and
26.1 mm by 20.4 mm.

THE VELIGER

Page 283

It may be noted here that no specimens of Littorina
littorea from Pacific coast localities have been found in
the Stanford University collection (Dr. A. Myra Keen,
personal communication), nor in the collection of the
Department of Paleontology of the University of Cali-
fornia, Berkeley (examined by the writer through the
courtesy of Dr. Joseph H. Peck).

Thus, living Littorina littorea are now known on the
Pacific coast from Puget Sound, Washington; Trinidad
and San Francisco Bays, California. The collection of
additional specimens from San Francisco Bay will help
to determine whether this mollusk has been able to estab-
lish a reproducing population, or whether the present
collection represents only one of a limited number of
individuals directly transported to the Bay. Dr. Niles
reports (in litt.) that she has been unable recently to
find this species in Trinidad Bay and there are no known
subsequent reports from Puget Sound.

It is interesting to note that the discovery of Littorina
littorea on the Pacific coast has proceeded chronologically
southward. A similar situation has prevailed since the
early part of the nineteenth century on the Atlantic
coast of the United States, where this mollusk has been
recorded as progressively moving southward, from Nova
Scotia (since 1840) to Maryland (in 1959) (WELLS,
1965). Its probable southern limit is correlated with a
21° C mean water isotherm (WELLS, op. cit.). There is
no indication, however, that the Pacific coast occurrences
illustrate a natural southward movement, but rather it
appears that this species was accidentally introduced
into the 3 known localities on as many different occa-
sions. The progressively southern occurrence of L. litto-
rea on the Pacific coast is apparently a coincidence with
the Atlantic coast phenomenon.

The means of introduction of this northern Atlantic
snail into San Francisco Bay, and earlier into Trinidad
Bay, remain problematical. The abundance of exotic
mollusks in the San Francisco Bay region [e. g., the At-
lantic species Modiolus demissus (Dittwyn, 1817), Mya
arenaria LINNAEUS, 1758, Gemma gemma (TorTTEN,
1834), Nassarius obsoletus (Say, 1822), Urosalpinx cin-
ereus (Say, 1822), Busycotypus canaliculatus (LINNAE-
us, 1758), and the Oriental species Tapes japonicus DEs-
HAYES, 1853, and Musculus senhouse: (BENSON, 1842) ],
as well as a number of other invertebrates, supposedly
largely introduced through oyster transport, well illus-
trates the ease with which exotic species have established
themselves locally. Almost all of the accidentally intro-
duced Atlantic mollusks arrived when shipments of the
American oyster, Crassostrea virginica (GMELIN, 1791)
were still being transported on a large scale to the Pacif-

Page 284

ic coast, before 1935 (Hanna, 1966). The possibility
that this rather conspicuous species was introduced with
Crassostrea into San Francisco Bay more than 30 years
ago, and has been overlooked, is not probable, when the
continuous collecting and numerous field expeditions by
local biologists, students, and shell collectors are con-
sidered. The possibility that the large introduced Atlantic
whelk, Busycotypus canaliculatus, was overlooked for a
period of 10 years (between 1938 and 1948) is held im-
probable by SToHLER (1962). However, small shipments
of Crassostrea from the Atlantic coast of the United
States are still being received for placement in San Fran-
cisco Bay, and between 1962 and 1967 plantings of
oyster spat were made 2 or 3 times in the southern por-
tion of the Bay on an experimental basis only (Mr. W.
Dahlstrom, Calif. Dept. Fish and Game, personal com-
munication). It is possible that juvenile Littorina, or Lit-
torina eggs, may have been introduced with these ship-
ments.

A second method by which this snail may have been
introduced involves the Atlantic quahog, Mercenaria
mercenaria (LINNAEUS, 1758). The report by NortH
(1963), cited by Hanna (1966), concerning the planting
6f 2000 Mercenaria in San Francisco Bay in 1963 is in
error (personal communications from Dahlstrom and Dr.
V. Loosanoff), and thus there have apparently been no
official plantings of this bivalve in the Bay (Dahlstrom,
Loosanoff, and J. A. Aplin, personal communications).
However, in January, 1968 several living Mercenara
were discovered at Coyote Point, San Mateo County,
in San Francisco Bay, together with numerous specimens
of the venerid Tapes japonicus by Mr. R. Setzer, a grad-
uate student at San Francisco State College (personal
communications from Setzer and Mr. D. Chivers, Calif.
Acad. Sci.). This small and questionably established
population is apparently the result of an independent ef-
fort by an unknown person, and was without the planting
permission of the California Department of Fish and
Game (Aplin, personal communication). In addition,
local restaurants are continually importing Atlantic qua-
hogs, and may dump spoiled shipments into the Bay. It is
also known that many independent introductions of
Atlantic oysters and clams have been made into the San
Francisco Bay region, and that, additionally, plantings
made officially in other California bays may be brought
into San Francisco Bay by private persons (Loosanoff,
personal communication). It is thus also quite possible
that specimens of Littorina may have been transported
recently from the Atlantic coast with Mercenaria.

The present shell with the operculum has been depos-
ited in the Department of Geology, California Academy

THE VELIGER

Vol. 11; No. 3

of Sciences (catalogue no. 40981); the animal has been

placed in the wet collections of the Department of In-

vertebrate Zoology of the same institution.

ACKNOWLEDGMENTS

For generous assistance with various details and in the
examination of specimens in the collection of the Cali-
fornia Academy of Sciences, I thank Dr. Leo G. Hertlein.
Mr. Dustin Chivers of the Academy kindly aided with the
curating of the specimen and contributed suggestions
and advice as did Mr. John Holleman of Merritt College,
Oakland. Dr. Joseph Peck generously permitted me to
examine the large shell collection of the Department of
Paleontology of the University of California, and Dr.
Myra Keen kindly provided information about the Stan-
ford collections. Dr. Doris K. Niles also furnished addi-
tional information concerning the material from Trini-
dad Bay. I wish to thank also Mr. J. Allan Aplin, Mr.
Walter Dahlstrom, Dr. Victor Loosanoff, and Mr. Robert
Setzer for communicating to me their records of Mer-
cenaria or Crassostrea in San Francisco Bay.

LITERATURE CITED

Hanna, G Datias
1966. Introduced mollusks of western North America.
Occ. Pap. Calif. Acad. Sci. no. 48: 108 pp.; 85 figs.; 4 plts.

(16 February 1966)
Nortu, Dan

1963. Oysters from Europe for San Francisco. San Fran-
cisco Examiner, sect. 1, p. 23 [cited in Hanna, 1966]

(16 June 1963)
STOHLER, RUDOLF

1962. Busycotypus (B.) canaliculatus in San Francisco Bay.
The Veliger 4 (4): 211-212; 1 text fig. (1 April 1962)
WELLs, Harry M.
1965. Maryland records of the gastropod, Littorina littorea,
with a discussion of factors controlling its southern distribution.
Chesapeake Sci. 6 (1): 38-42; 1 fig. (March 1965)

Revision of
“Introduced Mollusks

of Western North America’’

JAMES CARLTON
521 Mandana Boulevard, Oakland, California 94610
A REVISION OF G DaLias HANnna’s recent paper on the

introduced Mollusca of western North America (HANNA,
1966) is currently being undertaken under the supervision

Vol. 11; No. 3

of Allyn G. Smith, Associate Curator, Department of
Invertebrate Zoology, California Academy of Sciences,
San Francisco. Geographic regions covered include Cali-
fornia, Oregon, Washington, and British Columbia. Wor-
kers with any corrections, comments, suggestions, or new
records are encouraged to send such information to Mr.
Smith at the Academy, Golden Gate Park, San Francisco,
California 94118.

LITERATURE CITED

Hanna, G Datias
1966. Introduced mollusks of western North America.
Occ. Pap. Calif. Acad. Sci. no. 48: 108 pp.; 85 figs.; 4 plts.
(16 February 1966)

A.M. U.

AT THE ANNUAL MEETING of the American Malacologi-
cal Union at Corpus Christi, Texas, the following officers
were elected to serve during 1968-1969:

Dr. Joseph Rosewater, President

Dr. Alan G. Solem, Vice-President

Dr. Bruce G. Campbell, Second Vice-President
' Mrs. Margaret C. Teskey, Secretary

Mrs. H. B. Baker, Treasurer

Mr. Morris K. Jacobson, Publications Editor

Councillors-at-Large elected are: Donald R. Moore,
Robert Robertson, Donald R. Shasky, Myra Taylor.

The 1969 annual meeting will be held July 21 to 25 at
Marinette, Wisconsin.

Margaret C. Teskey, Secretary.

A. M. U.

Pacific Division

AT THE ANNUAL MEETING at Asilomar, California, in
June 1968 the following officers were elected to serve for
an indefinite period:

Dr. Bruce G. Campbell, Chairman

Dr. James H. McLean, Vice-Chairman

Mrs. Ruth French, Secretary and Treasurer

No meetings are planned for the near future. The an-
nual assessment of 50 cents per person was rescinded.

The Honor Award, bestowed by the Division from time
to time, was presented to Dr. Leo G. Hertlein of the
California Academy of Sciences for his continued out-
standing contributions to Malacology.

THE VELIGER

Page 285

W. S. M.

AT THE FIRST ANNUAL MEETING at Asilomar, California,
in June 1968, the Society adopted its bylaws. Officers
elected are:

Dr. William K. Emerson, President

Dr. A. Myra Keen, First Vice-President

Mr. Eugene V. Coan, Second Vice-President

Mrs. R. (Forrest) Poorman, Treasurer

Mrs. Bernadine Hughes, Secretary

As Representatives-at-Large were elected: Dr. Judith
Terry, and Miss Betsy Harrison.

At the meeting a contest was held for the best paper
submitted by a college student. A first and a second prize
was awarded to Miss M. T. Vassallo and Mr. S. K. Webs-
ter, respectively.

The second annual meeting will be held from June 18
to 21, 1969, at Asilomar.

U. M. E.

AT THE THIRD Concress of the European Malacologi-
cal Union in Vienna, September 2 to 7, 1968, the follow-
ing officers were elected:

Dr. Eugéne Binder (Geneva), President

Dr. FE E. Loosjes (Wageningen), Vice President

Dr. A. Zilch (Frankfurt), Secretary

Dr. Lothar Forcart (Basle), Treasurer

The next meeting of the Unitas Malacologica Euro-
paea will take place in Geneva, Switzerland in the year
LOWE

Important N otices

Because of the changed rules affecting second class
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attach any flyers to our journal henceforth. We shall,
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has arrived or announcements regarding special publi-
cations in our Notes « News column.

Page 286

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us of the new address at least six weeks before the
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by Prof. Ernst Marcus;
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THE VELIGER

Vol. 11; No. 3

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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
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THE VELIGER

Page 287

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Endowment Fund

At a Regular Membership meeting of the Society in No-
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ABOUT SUPPLEMENTS

Many of our members desire to receive all supplements
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Page 288

THE VELIGER

Vol. 11; No. 3

METHODS & TECHNIQUES

A Method
of Tagging Mollusks Underwater

BY

RICHARD J. ROSENTHAL

Westinghouse Ocean Research Laboratory
San Diego, California 92121

(1 Text figure)

INTRODUCTION

THE TAGGING OF SUBTIDAL marine mollusks has usually
been done at the surface. Most of these tagging studies
have involved a small number of economically important
species. Cox (1962) studied the red abalone, Haliotis
rufescens SwAINSsoN, 1822, along the California coast,
attaching the tags by wire threaded through the respira-
tory holes of the shell. NEwMAn (1966) used a disc tag
fastened by a nickel pin passing through a respiratory
pore to study the movements of the South African aba-
lone, H. midae LinNAEuS, 1758. More recently, FoRSTER
(1967) described tagging the British ormer, H. tubercu-
lata LINNAEUS, 1758, with disc tags using resin cement
as an adhesive. A study being conducted off southern
California by T- C. Tutschulte (personal communication)
on the migratory movements of Haliotis spp. utilizes
time-lapse photography and luminous tags cemented
with epoxy to the shells in situ.

In the first three investigations the animals were col-
lected by divers and the majority of the tagging was
completed above water. Surface tagging may introduce
more variables than are necessary, since the organisms
are removed from their normal environmental situation
and exposed for a brief period of time to unnatural con-
ditions. Placing the animals back in their original habitat
after marking does not guarantee that they will return
to a “normal” pattern of behavior. Tagging at the sur-
face may also be impractical when dealing with subtidal
mollusks which are solidly attached or cemented to the
substrate.

This study was undertaken to perfect a method for
tagging hard-shelled mollusks underwater, with a mini-
mum amount of disturbance to the animals and their
surroundings.

MATERIALS anp METHODS

A commercially available non-toxic epoxy’ was used to
attach the tag to the shell surface. This particular epoxy
was selected because it hardens thoroughly underwater.
The curing time depends on the water temperature and
the thickness of the epoxy used on each shell. Mixing the
resin and catalyst in a 1:1 volume on the surface was
found to be the best procedure, since attempts at mixing
the two components underwater proved unsatisfactory.
The resin-catalyst mixture is grey-white in color, which
may be suitable for mollusks having lightly pigmented
shells. For those with darker shells, a black pigment can
be added to darken the epoxy, thereby lessening the at-
tention of predators or the attraction of curious fishes to
the tagged mollusks.

Engraved disc tags made from gravoply were used to
increase the permanence of the number and ease of read-
ing underwater. The tags and newly mixed epoxy were
taken underwater in separate containers by scuba divers.
When suitable animals were located, the overall shell
length was recorded and the tagging region of the shell
was cleaned with a wire brush. Then the epoxy was ap-
plied to the mollusk shell and a numbered tag was pressed
into the epoxy (Figure 1).

RESULTS anv DISCUSSION

A tagging experiment on Kellet’s Whelk, Kelletia kelletu
(Forses, 1850) was started in December 1967 to acquire
more knowledge about the ecology and life history of
this marine gastropod. A total of 120 K. kelletu have
been tagged in depths of 15 to 20 m of water at 3 differ-
ent localities off San Diego County, California. The tag-
gings have been conducted off of Point La Jolla, Mission
Beach, and along an ecological survey line near Del Mar.
These 3 subtidal areas were selected because they differ
substantially in substrates and species composition.

A great deal of the tagging took place during the
reproductive season (March-June), when large numbers
of these whelks were copulating and laying eggs at the 3
localities. During subsequent dives, copulation and egg
laying was observed in tagged individuals — an indica-
tion that the tagging had not disturbed their behavioral
activities.

The tags remain securely attached to the shells of the
animals located to date (September 1968), 263 days after

* Sea Go-In Poxy Putty — 1324; Permalite Plastics Corporation,
Costa Mesa, California.

Vol. 11; No. 3

THE VELIGER

Page 289

the initial 11 Kelletia kelletui were tagged on December
11, 1967. Success in finding these motile benthonic ani-
mals is largely dependent upon the visibility underwater,
although searching from a permanently emplaced survey
line eases the difficulty in locating them.

Other mollusks have been successfully tagged using
this method: The rock scallop, Hinnites multirugosus
(GaeE, 1928); the red abalone, Haliotis rufescens; and
the wavy top shell, Astraea undosa (Woop, 1828).

Figure 1

The drawing shows a tagged Kelletia kelletii with a numbered disc
embedded in epoxy and cemented to the dorsal region of the shell.

The size of the mollusk does not present a problem,
since the epoxy can be shaped to conform to the charac-
teristics of the outer shell and the disc tags sized accor-
dingly. This reduces any hindrance to the movement of
the animal and the possibility of the epoxy coming in
contact with the animal’s living tissue (Figure 1).

The advantages of this technique appear to be the
permanence of marking, and the fact that tagging can
be completely carried out underwater with a minimum
amount of disturbance to the subjects and their environ-
ment.

ACKNOWLEDGMENTS

I am particularly grateful to Dr. W. D. Clarke for valuable
advice and encouragement, and to Mr. R. E. Bower for
technical assistance in the field.

LITERATURE CITED

Cox, Keitn W.

1962. California abalones, family Haliotidae. Fish Bull. 118,

Calif. Dept. Fish and Game, Sacramento, Calif.
Forster, G.R.

1967. The growth of Haliotis tubcrculata: results of tagging
experiments in Guernsey 1963-65. Journ. Marine Biol.
Assoc. U. K. 47: 287 - 300

Newman, G. G.

1966. Movements of the South African abalone Haliotis

midae. Invest. Rep. Div. Sea Fish. S. Afr. No. 56: 1 - 20
RANDALL, JOHN E.

1964. Contributions to the biology of the queen conch, Strom-
bus gigas, Bull. Mar. Sci. Gulf and Carib. 14 (2): 246
to 293

A Method of Color Preservation
in Opisthobranch Mollusks

BY

GORDON A. ROBILLIARD

Department of Zoology
University of Washington
Seattle, Washington 98105

INTRODUCTION

CoLor AND COLOR PATTERN are useful taxonomic charac-
ters in the opisthobranch mollusks. However, I know of
no published reports describing a method of preserving
color in opisthobranchs. Instead, it is usually assumed
that the colors will fade or be bleached out in formalin,

Page 290

THE VELIGER

Vol. 11; No. 3

alcohol, or other preservatives. Color is usually preserved
in color photographs or field notes or both,

ToyaMa & Mriyosui (1963) and WALLER & EscH-
MEYER (1965) successfully employed an antioxidant,
butylated hydroxytoluene (hereafter referred to as Ionol
C. P. -40), to preserve some colors in fish and a prawn.
WALLER & ESCHMEYER (op. cit.) used 1, 10, and 20 cc
of Ionol C. P.-40 per 4500 cc of 10% formalin to make
the test solutions (0.02%, 0.22%, and 0.44% concentra-
tions respectively). Eighteen months later, they found
that the color of fish in the 0.44% solution was best
preserved, but preservation of colors of fish in other test
solutions was superior to that of fish in untreated formalin
solution.

In this note, I report the results of testing this tech-
nique on opisthobranchs from 3 orders (Cephalaspidea,
Sacoglossa, and Nudibranchia) with emphasis on the
nudibranchs.

MATERIALS ann METHODS

A stable emulsion was formed by vigorously stirring Ionol
C. P. -40 into hot sea water (55° to 65° C) (Shell Tech-
nical Bulletin, IC:67-16). Concentrations of Ionol C, P.
-40 used in the emulsions gave final dilutions of 0.1%
to 0.5% Ionol C. P. -40 by volume. Formalin was added
to this emulsion to give a final concentration of 5%
formalin by volume (hereafter referred to as Ionol C. P.
-40 emulsion). Ionol C. P. -40 is also readily soluble in
alcohol.

Most of the opisthobranchs were relaxed for 1 to 5
minutes, depending on size, with succinylcholine chloride
(BEEMAN, 1968). A few, especially the Cephalaspidea,
were relaxed for 2 to 8 hours in propylene phenoxetol
using 1% by volume of propylene phenoxetol in sea
water (Owen, 1955; OWEN & STEEDMAN, 1958).

Small animals (< 1.5 to 2.0cm long) were then put
directly into Ionol C. P. -40 emulsion. Formalin diffuses
inward rapidly enough to preserve the internal tissues.
Larger animals (>2cm long) were injected with a
small amount of 5% formalin in sea water without Ionol
C. P. -40, to preserve the internal tissues, then put into
the Ionol C. P. -40 emulsion.

The Ionol C. P. -40 emulsion was not injected into the
animal because it eventually forms an oily film in the
body cavities and over the tissues. This is particularly un-
desirable if the animals are to be dissected. In any case,
there are few internal organs in an opisthobranch in
which color is distinctive or taxonomically important.

The method was tested on 1 sacoglossan, Elysia hedg-
pethit Marcus, 1961; 2 cephalaspideans, Haminoea sp.,

and Aglaja diomedea (Brercu, 1893) ; and 30 species of
nudibranchs of which 12 were dorids, 12 were dendro-
notaceans, and 6 were aeolids.

RESULTS

Of the concentrations tested, 0.3% Ionol C. P. -40 emul-
sion resulted in maximal color retention and was used
for most of the specimens.

Stored in bottles in glass-door cabinets, the specimens
in Ionol C. P. -40 emulsion have been exposed to normal
artificial lighting, but protected from direct sunlight, for
up to 2 years. In most cases, the original color has been
retained albeit with varying degrees of fading in some
species. The color of virtually all specimens stored under
similar conditions, but in untreated 5% formalin in sea
water, has faded markedly or disappeared.

Some colors are preserved better and longer than
others. The yellow body color and black spots of Aniso-
doris nobilis (MacFarianp, 1905) and Archidoris mon-
tereyensis (Cooper, 1862) are well preserved as are the
yellow and orange pigments of Triopha carpenter
(Stearns, 1873). The orange on the cerata of Laila
cockerelli MacFar.anp, 1905 is completely preserved
while the orange on Hermissenda crassicornis (EscH-
SCHOLTZ, 1831) faded slightly. Slight fading has occurred
in the orange-red body color of Rostanga pulchra Mac-
FarLAND, 1905. The salmon-pink body color of Tritonia
festiva (STEARNS, 1873) and T: gilberti (MacFarLanp,
1966) and the orange body color of T: (Tochuina) tetra-
quetra (Pauvas, 1788) fade only slightly. The same is
true of the brown blotches of Diaulula sandiegensis
(Cooper, 1862). Many specimens of Dendronotus spp.
have been preserved in Ionol C. P. -40 emulsion and the
colors — white, metallic orange, brown, magenta, yel-
low, mauve, purple, pink, grey, red — have faded slightly
over periods of up to 2 years. The orange body color and
white spots of Dirona aurantia Hurst, 1965 and the
white pigment on the cerata of D. albolineata CocKErR-
ELL & Eviot, 1905 fade somewhat. The dark green body
color of Elysia hedgpethi and the black (or dark purple)
of Aglaja diomedea have faded very little.

In a few cases, the color faded markedly or completely
disappeared. The chocolate brown color of Onchidorts
bilamellata (Linnaeus, 1767) faded to about a third of
its original intensity. In Cadlina marginata MAcFaRLAND,
1905, the yellow pigment disappears completely within
12 to 24 hours. The yellow on the notal papillae of
Acanthodoris hudsoni MacFarLanp, 1905 fades marked-
ly over a week. Structural colors, such as blue on Her-
muissenda crassicornis and other opisthobranchs (Burcin,

Vol. 11; No. 3

THE VELIGER

Page 291

1965, Bircin-Wyss, 1961) usually disappear when the
animal is killed and they cannot be preserved by an
antioxidant.

CONCLUSION

The use of an antioxidant such as Ionol C. P. -40 with a
preservative seems to be a successful and relatively simple
method of preserving most colors in the opisthobranchs.
However, on the basis of this study, it is not possible to
generalize about which colors will be preserved between
different taxa or even within a certain taxon. Further
experimentation with Ionol C. P. -40 will likely demon-
strate a more widespread applicability as well as deline-
ating more precisely which colors will or will not be
preserved.

The optimal concentration proved to be 0.3% Ionol
C. P. -40 by volume in a solution of 5% formalin in sea
water.

Color retention is an obvious advantage to the taxon-
omist in that it allows him to augment the description of
a species. This is especially true in soft-bodied animals
like opisthobranchs where color, though quite variable
intraspecifically, often serves as a guide in species identi-
fication.

ACKNOWLEDGMENTS

I wish to extend my thanks to Brian Case, University of
Manitoba, for first bringing the method to my attention
and to the Shell Chemical Company, Industrial Chemicals
Division (Portland Office) for the sample of Ionol C. P.
-40. I am grateful to Dr. Alan J. Kohn for reading the
manuscript and for his suggestions. This work was sup-
ported by a Special Scholarship from the National Re-
search Council of Canada.

LITERATURE CITED

ANONYMOUS
1967. Ionol CP antioxidant.
BEEMAN, Ropert Davip
1968. Order Anaspidea.
prt. 2) :87- 102; plt. 11; 12 text figs.
Burcin-Wyss, ULRIKE
1961. Die Riickenanhange von Trinchesia coerulea (Mon-

Shell Tech. Bull. IC: 67-16

The Veliger 3 (Supplement,
(1 May 1968)

TAGU). Rev. Suisse Zool. 68 (4): 461 - 582; plt.
Burcin, Urrike F
1965. The color pattern of Hermissenda crassicornis (Escu-
scuoLtz, 1831) (Gastropoda: Opisthobranchia: Nudibranchia)

The Veliger 7 (4): 205-215; 9 text figs. (1 April 1965)
OweEN, GARETH
1955. Use of propylene phenoxctol as a relaxing agent.

Nature 175 (4453) : 434 (5 March 1955)

Owen, GareTtu « H. F. STEEDMAN
1958. Preservation of molluscs.
33 (3): 101 - 103
Toyama, Kenzo « Gota Miyosu1
1963. Prevention from color fading of aquatic animals under
preservation. Journ. Tokyo Univ. Fish. 50 (1): 43-48
WALLER, RicHarp A. & WILLIAM N. EscHMEYER
1965. A method for preserving color in biological specimens.
Bioscience 15 (5): 361

Proc. Malacol. Soc. London

BOOKS, PERIODICALS, PAMPHLETS

Between Pacific Tides

by Epwarp FE Ricketts & Jack Catvin, Fourth edition,
revised by Joe, W. HepcpetH. September 19, 1968. xiv
+614 pp.; 8 color plates; 302 text figures.

Stanford University Press, Stanford, California. $10.-.

This classic work on the ecology and natural history of
plants and animals inhabiting the Pacific shores in the
area between the low and the high tide marks was written,
originally, “for laymen, for beginners ... ” but it has
filled a need far beyond that envisioned by Ed Ricketts.
Because of the thoroughly annotated bibliography, the
book has become an important first source for many
serious investigators and an indispensable text book in
many courses in invertebrate zoology.

The book has gone through three previous editions, the
first two by the original author, and the third edition
revised by Dr. Hedgpeth. Each edition was better than the
previous one, each updated to include the latest develop-
ments in the fields of endeavor covered by the scope of
the work.

This fourth edition, again revised and brought up to
date, is a worthy successor to the other three editions.
Dr. Hedgpeth has brought his critical talents to bear and
his influence can be perceived on practically every page,
although he has managed beautifully to preserve the orig-
inal charm of the book. And while a number of the illus-
trations are the old familiar ones, many have been super-
seded by better pictures illustrating more precisely what
was desired to be called to the reader’s attention. Even
the color plates have been much improved, although they
may perhaps, in a future edition, be printed with a still
better technique which will render the delicate colors of
the intertidal area more life-like.

Page 292

We can only say that this new edition is a must for the
layman who wishes to become better acquainted with
the seashore and its wealth of animals and plants; it is
indispensable for the serious student, be he a beginner or
advanced; and the professional invertebrate zoologist will
find many tidbits of important knowledge in the pages of
this book.

There is, however, one part of the book that was not
possible in the “good old days” of Ed Ricketts, a part
which should be required reading not only for the student
of the seashore, but for politicians and all those persons
who are in a position to make policy and to decide “what
is best for all.” To this portion of the book Dr. Hedgpeth
brings some of his best thoughts and it is evident from
these pages that “conservation” is not a sometime thing
for him. It is a truly thought-provoking chapter and it
alone would be worth the price of the entire book.

RS

Tidepool Animals from the Gulf of California

by Westey M. Farmer. 70 pp.; 4 color plates (with 6
photographs each) ; 170 line drawings. Wesword Comp.,
Post Office Box 15333, San Diego, California 92115.

Price, $6.50. Stiff paper bound.

Since travel into Lower California is increasingly pop-
ular, shell collectors from near (San Diego, for example)
and far (New York and Teaneck, for example) roam over
the sandy beaches and rocky shores of the rich Gulf
of California. The climatic conditions are, during a part
of the year, very agreeable and collecting thus becomes a
pleasant occupation.

For the Pacific coast of North America we have the
book by Ricketts and Calvin, as well as a few other, more
limited books to help the collector in identifying his loot.
Aside from Steinbeck’s “The Sea of Cortez” there is no
popularly written work available for the Gulf. Or, rather,
there was, until now, no such work available.

Mr. Farmer, who was Curator of exhibits at the San
Diego Museum of Natural History, has brought together
in this booklet a brief description accompanied by an ex-
cellent pen-and-ink drawing of somewhat over 100 of
the animals one is likely to encounter in the tidepools.
His color photographs, especially of his favorite nudi-
branchs, are superb. His descriptions are pithy and to the
point.

It may be anticipated that future editions will be ex-
panded to include many species of invertebrates that are

THE VELIGER

Vol. 11; No. 3

relatively frequently encountered, although, perhaps not
by the person casually strolling along the beaches.

RS
Descriptions of New Species of Gastropods
from Clipperton Island
by Leo G. HeErTLEIN & Epwin C. Atuison. Occas.

Papers Calif. Acad. Sci. No. 66; 13 pp.; 13 figs. June 27,
1968.

Nine species of some extremely small (less than 4 mm)
to medium sized (50mm) snails are discussed. Six of
these species are new to science.

RS

Kelp Habitat Improvement Project

by WHEELER J. NortH. 123 pp.; 47 figures; 22 tables.
Published after July 1, 1968 by the W. M. Keck Labora-
tory of Environmental Health Engineering, California
Institute of Technology, Pasadena, California.

This, the fifth annual report, appears to us to be more
encouraging than its predecessors inasmuch as the general
outlook for the re-establishing of the once thriving kelp
beds along the California coast seems more promising.
Much detailed information, of value to the student of
marine ecology, is contained in these pages. The conser-
vationist also will take heart from the perusal of the
report.

RS

Water Quality and Biologic Conditions
South Bay Aqueduct

by C. A. McCuttoueu. State of California Resources
Agency, Department of Water Resources. 180 pp.; nu-
merous tables; 1 multiple fold-out; 15 pages with 2 to 4
photographs.

Of particular interest to the malacologist is the part
of the report that deals with the Asiatic clam, Corbicula.
RS

Age of first Marine Terrace Near Santa Cruz, California

by Witi1aAM C. BrapLey & WARREN O. AppicotTT. Geol.
Soc. America Bull., vol. 79, pp. 1203-1210; 1 table; 1
fold-out insert. September 1968.

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, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.

It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.

Short articles containing descriptions of new species or other 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 geo-
graphical longitudes and latitudes added.

Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS’; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A.M. U.,
U.M.E., W.S.M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION 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 invited. The column “BOOKS, PERIODICALS, and 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, not
exceeding 81,” by 11”, at least double spaced and accompanied by a clear carbon
or photo 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 accommodate the pamphlet (which
measures 51/,” by 81”), with double first class postage, should be sent with the
request to the Editor.

EDITORIAL BOARD

Dr. Donatp P. Assort, Professor of Biology

Hopkins Marine Station of Stanford University

Dr. Jerry DononvueE, 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

Dr. E. W. Facer, Professor of Biology

Scripps Institution of Oceanography, La Jolla
University of California at San Diego

Dr. Caper Hann, Professor of Zoology and
Director, Bodega Marine Laboratory
University of California, Berkeley

Dr. G Datias Hanna, Curator

Department of Geology

California Academy of Sciences, San Francisco
Dr. Jorn W. Hepcretu, Resident Director
Marine Science Laboratory, Oregon State University
Newport, Oregon

Dr. Leo G. HERTLEIN,

Curator of Invertebrate Paleontology

California Academy of Sciences, San Francisco

EDITOR-IN-CHIEF

Dr. Rupotr STroutre, Research Zoologist
University of California, Berkeley

Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology

Stanford University, Stanford, California

Dr. Victor Loosanorr, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr. Joun McGowan, Associate Professor of
Oceanography

Scripps Institution of Oceanography, La Jolla
University of California at San Diego

Dr. Frank A. Pireixa, Professor of Zoology
University of California, Berkeley

Mr. Attyn G. Smiru, Associate Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

Dr. Ratpu I. Siru, Professor of Zoology

University of California, Berkeley

Dr. Cuarces R. STASEK, Associate Professor
of Zoology

Florida State University, Tallahassee, Florida
Dr. Donatp M. Witson, Professor of Biology
Department of Biological Sciences

Stanford University, Stanford, California

ASSOCIATE EDITOR

Mrs. Jean M. Cate
Los Angeles, California

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME II APRIL 1, 1969 NUMBER 4
ConTENTS
The Muricidae of Fiji — Part II. Subfamily Thaidinae. (Plates 47 to 49;
21 Text figures; 1 Map)
WaLTER OLIVER CERNOHORSKY .. . . 203

Effects of Turbidity-Producing Substances) in Shp Waren! on Gas ane anae af
Three Genera of Bivalve Mollusks. (7 Text figures)
Harry C. Davis & HERBERT Hipu ae Bee hae PO RSnRE | maul iia were se ONT
A New Species of Murexsul (Gastropoda: Murieidae) from the Galapagos Islands.
(Plate 50; 1 Text figure)
WitiiaM K. Emerson & ANTHONY D’ArTILIO’. Cw “E324.
Predators on Olivella biplicata, Including a Species- opecinc Pretor ‘Aveiasate
Response. (Plate 51; 1 Text figure)

D. Craic Epwarps . . . Ra INAS Wan A Memes 8) IMSS Eis etc i HI 8)
Rediscovery of Terebra trochlea eee eee (Plate 52)

Twita BRATCHER . . ‘ Bi te se Gita eRe og la eR OA:
Growth Characteristics of Aenea peicons Eee cenon (4 Text figures)

Ron KENNY . + . 336

Use of the Binet € asa Semis: Oran in an Peete (Gastronede: Onatey
(Plate 53; 1 Text figure)

Barry R. Witson . . Br eo eee 3 G40
A New Genus and Two New Species oe lay pings fom ‘Ihe Panamie Province
(Gastropoda: Muricidae) . (Plate 54)
HELEN DUSHANE . . » 343
Gross Anatomy and Giesineation of ie Gommensal: Gascon, \Galedonicla fone:
rouzierl SOUVERBIE, 1869. (Plate 55; 4 Text figures)
JosEPH ROSEWATER Wisc: - 345

A Preliminary Survey of WeaTnes or Cone: Rook abe Ndideent Areas! Gulf of
California, Mexico. (1 Map)
HELEN DUSHANE & ELEN DRENNAN ee) eos ee hs fe se ene) S61

[Continued on Inside Front Cover]

Distributed free to Members of the California Malacozoological Society, Inc.
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Send subscription orders to Mrs. JEAN M. Caté, 12719 San Vicente Boulevard,

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Second Class Postage paid at Berkeley, California

AS SONA

JUN 17 1968

LIBRARIES

ContTENTs — Continued

Two New Species of the Genus Volua Ropine, 1798 (Ovulidae FLEMING, 1828).
(Plate 56)
CrawForp NEILL CaTE : aoe, 2 eae (G8
Zoogeographical Studies on Living Cowes (1 Map)
Franz ALFRED SCHILDER :
Seasonal Observations on Diet, and Stored Glycouent atid pias in dhe Home Cen.
Tresus capax (GouLp, 1850). (Plate 57; 1 Text figure)
Rosert G. B. Rei
Seasonal Gonadal Changes in Two Bivalve Moluckoe in remales, Pay Grifter,
(Plates 58 to 60; 4 Text figures; 1 Map)
\, VERNON KENNETH oe |i OS ee
The Shell Structure and Mineralogy of Ghana pellucida IBeOnER TT (Plates 61
to 64; 6 Text figures)
Joun Davin TayLor « WILLIAM JAMES KENNEDY

Molluscan Faunas of Pacific Coast Salt Marshes and Tidal Gree (1 Map)
Keir B. MacponaLp von che le spor soak eee ae
The Type of Tegula funebralis (A. Apts: "BER (Plate 65)
RupoLF STOHLER ‘ Eee ae wy
The Littorina ziczac Species Compler! (Plate 66; 4 Text figures)

Tuomas V. Borkowski & MarRILyNN R. Bonen apart
Spawning and Development in the Trochid Gastropod Euchelus penunan us (GouLp,
1841) in the Hawaiian Islands. (2 Text figures)
Tuomas M. DucH
New Records of Nudibranchs fron New Tose
Rosert E. LovELAND, Gorpon HENDLER & Gary NEWKIRK ‘
Nomenclatural Changes for the New Species Assigned to Cratena by MING SARaAi,
1966. (2 Text figures)
RicHarp A. ROLLER
An Annotated List of Opisthobranere Gorn on ine Onen (Conny, Galera.
(1 Map)
RicHaArp A. ROLLER & STEVEN J. Lone
A Note on the Range of Gastropteron pacificum (Onismobrenenn Corhalesatilen)

(1 Map)
XS Hans Brertscu ‘ PAM al Oe nd. Gl
‘An Immunological Study of Belecypod conten (3 Text figures)

LARKLYN FISHER

NOTES & NEWS ieee ee Ser oe
An Overlooked Subcenus ond Saists from Barrie A. Myra KEEN
Laternula Living on the Pacific Coast? A. Myra KEEN
Sexual Dimorphism in Tegula funebralis. PETER W. FRANK
Spawning Notes — III. Strombina maculosa. Fay Henry WOLFSON
Spawning Notes — IV. Cerithium stercusmuscarum. Fay Henry

WOLFSON

BOOKS, PERIODICALS & PAMPHLETS

- 364

= 307

5 yi)

. 382

- 391
2 B89)

. 408

. 410

9 43i'5)

. 418

. 421

= 424

» 431

- 434
- 439

- 444

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

Vol. 11; No. 4

THE VELIGER

Page 293

The Muricidae of Fiji

Part II — Subfamily Thaidinae

BY

WALTER OLIVER CERNOHORSKY

Vatukoula, Fiji Islands, and Division of Mollusks, Smithsonian Institution
U.S. National Museum, Washington, D.C. 20560

(Plates 47 to 49; 21 Text figures; 1 Map)

THIS IS THE NINTH PART in the series of faunal mono-
graphs dealing with the marine mollusca of the Fiji
Islands; it is the second part of the Muricidae of Fiji, the
first having been published by the author in 1967.

Species recorded from Fiji have an Indo-Pacific distri-
bution and none of the species is endemic to the Fiji
Islands. For notes on the geography of the Fiji Islands
and other pertinent data see CERNoHOoRSKY, 1964.

TAXONOMY

The systematics of the muricid subfamily Thaidinae is
presently in a state of flux. Several applications are pend-
ing with the International Commission on Zoological
Nomenclature (BRADLEY & PaLMer, 1963; Keen, 1964,
1964a) for either a suppression or validation of thaidine
genera, type species and even family groups. Since rulings
by the ICZN on these applications cannot be anticipated,
pre-application taxonomy has largely been adopted.

Existing taxonomic treatments of the thaidine group
are not only few but highly conflicting in opinion; some
workers retain the thaidine group of species within the
family Muricidae while others assign them to a separate
family Thaididae. Powerit (1962) arranged the Neo-
zelanic species of Thais s. 1. under the family Thaisidae.
IREDALE & McMicuHaev (1962) accorded familial rank
to East Australian Thaidinae, Typhinae and Trophoninae.
Keen (1958) considers West American Trophoninae and
Typhinae to be of muricid stock but separates Thaididae
as a separate family.

Workers who study the shell, animal, ecology, and em-
bryology of thaidine species as a single biological unit
disagree with an elevation of Thaidinae to familial rank
and consider the various muricid groups under discussion
to be of a common phylogenetic origin. Cooke (1919),
TuieEre (1929), ARAKAwa (1962), Wu (1965a), Hane

& Kosuce (1966, 1967), and Orr-Maes (1967) are some
of the workers who retain thaidine genera in the family
Muricidae.

As far as tropical Pacific thaidine species are concerned,
there seems to be little or no zoological reason for a
separation from the Muricidae. In shell morphology,
animal, anatomy, radula, habitat, feeding requirements,
and geographical distribution, Thaidinae are so closely
related to other muricid genera that existing natural af-
finities and common phylogenetic origin can hardly be
concealed. There is no decided gap between the tritonali-
ine genera Urosalpinx Stimpson, 1865 and Muricopsis
Bucguoy, DauTZENBERG & Bouce, 1882, on the one hand
and the Morula-Cronia group in Thaidinae on the other;
here the characters are transitional and rather nebulous.

THE THAIDINE RADULA

A generic grouping based on shell characters alone dis-
agrees with a classification based on radula features.
Cooke (1919) contended that “an examination of the
radula of the various specis of Thais lends little support
to the groupings of that genus, based on the forms of
the shell, to which are given the names of Stramonita,
Tribulus, Polytropa, Thalessa, etc.’ He indicated that
certain groupings are desirable, but rather in a different
direction.

How generic groupings based on shell characters alone
cut right across an arrangement based on radular evi-
dence is best illustrated by Mancinella Linx, 1807. The
two species M. mancinella (Linnazus, 1758) and M.
tuberosa (ROpiNG, 1798) do have a common radula pat-
tern which differs from radula features of some species
of Thalessa H. & A. Apams, 1853 and would support a
generic separation based on shell characters (lirate lab-
rum). Thais aculeata Desuayes, 1844 (= T. hippocas-

Page 294

THE VELIGER

Vol. 11; No. 4

tanum auctt.) is generally assigned to Thalessa on the
basis of shell characters, but belongs to Mancinella on
the basis of radula features. The radula of the type spe-
cies of Thais Ropinc, 1798, i.e. T: nodosa (LINNAEUS,
1758) is of the same pattern as the radula of the Man-
cinella group, according to Cooxe’s description (op. cit.,
as T. neritoides); the variant Thais nodosa var. ascension-
is (Quoy & Gammarp, 1833) [== T. meretricula Rovine,
1798] has, however, the radular pattern of Thalessa. It
is realized that there is a certain amount of variation in
muricid radula pattern within a species, but the differ-
ences between the radula of a species and a varicty as
described by Cooke (op. cit.) exceed the expected range
of variation. Should Cooxe’s results prove to be correct,
and Thais nodosa and the variety T: meretricula are in-
deed conspecific, then the variability of the thaidine radula
is much ercater than anticipated.

The radulae of the Thais-Mancinella-Thalessa group
shade through gradual stages from a simple tricuspid
rhachidian of Afancinella to the claborate pattern of
Thais and Thalessa with bifid side cusps, numerous side
denticles and prominent outer cusps. The radulae of
Drupa Rovinc, Drupella Tuireve, Drupina Dart and
Morula ScuiuMACHER have a sufficiently modified radula

and shell to merit recognition of these groups as full
genera.

ACKNOWLEDGMENTS

I would like to record my thanks to Dr. Harald A. Rehder,
Smithsonian Institution, U.S. National Museum, Wash-
ington, for the research facilities made available by the
Division of Mollusks, and for his valuable discussions and
opinions on nomenclatural problems.

Thanks are due to Dr. E. Binder, Muséum d’Histoire
Naturelle, Geneva; Dr. N. Tebble, British Museum (Nat-
ural History) London and Mr. J. O'Grady, Linnean
Socicty, London, for the opportunity afforded me to
examine the type collections of Muricidae in these Insti-
tutions.

Fiji collectors have generously made available their
collections for this study, and my thanks are due to Mr.
and Mrs. R. I Browne, Nausori; Mr. and Mrs. FE Freitag,
Suva; Mr. E. Gardner, Suva; Mrs. C. Jameson, Lautoka;
and Mr. A. Jennings, Auckland, New Zcaland.

This work was supported (in part) by a National
Research Council postdoctoral Research Associateship
with the Smithsonian Institution, U.S. National Museum,
Washington.

INDEX

( * indicates a homonym or synonym)

aculeata Drsuaves ( Thais) 295 clathrata (Drupa) 298n |) Sailinstrisc WOOD) pesete rae 2098
* aculcata Link 296 | * concatenalus . 306 | * indigoferum
* affine . 298 | * confragosa 311 | * infwmata
* affinis 295 cornus (Drupella) 3 SOV |) CAIRO! cxcimercmnerernarenconcec
* alba Womnron & JAQuINOT 304 | * dactylordes 303 intermedia (Thais) co.
* alba More 310 | * dealbata GYOV.L, || HOMOMPV —coemnseererumronsancn ecerncetin con snienranene
* albolabris . 299 | * digitata 303 * Taurentiana
* alouina .. 296 | * churnca 304 | * lobata
* ambusta . 309 | * echinata . 309 | * major .
anaxercs (Mforula) 307 caregius (Maculotriton) » BID mancilla eae 304
* andrewst 309 | * clata . 304 mancinella (Mancinella) occccoecuennen 296
angulata (Drupella) 306 fixcclla Guiviin (Morula) 5 Ru margariticola (MOrUIA) cececccncunne 312
* angulalum 309 | * fiseclla BustINVILLE . 312 marginatra (Drupa) a. 298
* arachnoides GEXGVGY || UU PRCLCOLAE essteae ee nseerceet ae ne ear dhens 306 | * martiniana
armigera (Thais) 295 | * fusca . 298 | * miticula |
2 @SIWOMO. porcecenernre HO) || fMOGKCOMUGIE croocerteesrmrcom txconnerrriernensooscen 298 MIOTILUP ONG! Pe cninscmicnumdna:t atc.csee nee
aurantiaca (Morula) 311 | * gemmulata 296 morum (Drupa) ..
* bicalenala 308 | * glacialis 296 | * morus
biconica (Morula) - 308 | * glossularia BYOB. || ACISTVONCOR caraos stoinccanstensanseterorrscos
* canccllala 307 granulata (Morula) 308 | * nassoides .
* castanca 297 grossularia (DrUpinad) coe GIO) |) 2 GAGINNCNELS WAVE: canorennomnm
* castancum . 297 | * hederacca : 313 | * nenitordea .
* ceylonica . 308 | * hippocastanum auct. POR || WCRROICHS swrnvanreccomomorseer
* chaidca Orr-Mars 304 | * Iippocastanum Woon .. 301 nodicostata (Morula) ccc
* chrysalis . MOOR Pion day acne ee aaadronesuncmen nna PHOKS) |) PORMYOCRE crrrsrecrrseeccertereeecransrees
* chrysostoma so SX) |] HAG INMTITS WANS cxcirnconncconommscer FON |) POGUE crmunscemaxcammmrenoreiiae
* cingulifera 308 | * hystrix LINNAEUS .. . 299 Ochrostoma (Drupella) voces

Vol. 11; No. 4 THE VELIGER Page 295
"9 ALE OANGNGTORY rae See cee ert rape Pee ey 310 | * rubuscaestwm KIRA wissen QO SMa na Tell ammeter sen ienumnnmmeenean og
parva (MOrula) vrcmesmcnnnnnnennunnnnn 309 | * rubuscaesius HIRASE & TAKI wins PSS} | NS CH AYE) aul hee re a ee 297
S111 CORMe et rae ne eee ee OOH TLD USC LESLUS INODING | aomrenntstiect tee er PHOR) || HAYS HIOUE, cerccerctcerrceeacetreceeeert terete 311
* picta RODING ...... 313 rubusidaeus (Drupa) .. Bon |) * tribulus vec. . 299
* picta SWAINSON 1. 313 rugosa (Drupella) . 306 | * tuberculata . 308
* pseudohippocastanum . 205 serta (Nassa) wu. . 313 tuberosa (Mancinella) w 297
"GO OTOOGDOTE ccmmmrecrtemimeerermemnncremmees SXOS),|| GAC DVORS cocemmmnmmcacommvesnemmnm 6{)! HC WORN) ccarcrrmmronmirerseceemrmaerecn, GO)
PRTC CON CMe rene mere ene One a RULE her Ot | prrae SPE CETILINN \acnite Santee tr etatetus cant nisnciactstonicont BOM MCA TECDILIS i ise rrenctrnrcetse ones comin eee ace 308
PRET TLV SSCL teeter iste erste cen 312 | * sphaeridea ...... « 310 wevill mn (Veoill a) ieee teen ee nenane 313
ricinus LINNAEUS (Drupa) 299 spinosa (Morula) PHS OG a |e ULOLA Ge armen eee sl mene ene tenes es 298
STIGINUS\VIOOD Ee seeteer eres Bog | A stellaris, vnc: meSile |) a Wittenselen 304
San UDIUSGACSIOWEAIRASE)eenatee een YOKE) |) OCHIOUDO: ceerrecertacer eterna ccc BION aCe NS tS yteanece ee en eee teenie renee ies 304.
EGON IAL A te rrersstonen stn sterteeittee tanemtesa teens QQ

NEOGASTROPODA Type Locality: None.
Habitat: On reefs, under coral and basalt rocks, in shal-

MuRICACEA low water. Uncommon.
Distribution: Throughout the Fiji Islands. — Indo-Pacific.
AiroRierD Discussion: Murex hippocastanum Linnarus, 1758 is
Thaidinae not the Thais hippocastanum of authors. The Linnaean

Thais Rovinc, 1798

Thais Ropinc, 1798, Mus. Bolten., p. 54. Type species by SD
(Stewart, 1927) T: lena Ropvtinc, 1798 = Nerita no-
dosa LinNAEus, 1758

Thais aculeata (DESHAYES & MILNE Epwarps, 1844)
(Plate 47, Figure 1)

— Thais hippocastanum auctt. (non Murex hippocastanum
Linnaeus, 1758)

1822. Purpura hippocastanum Lamarck, Anim. s. vert., 7:
258 (non Murex hippocastanum LinnaEvs, 1758)

1836. Purpura hippocastanum Kiener, Spec. gén. icon. coq.
viv., 7: 52; pl. 12, figs. 33, 33a (non Murex hip pocastan-
um LinnAEus, 1758)

1844. Purpura aculeata DEsHAYES & MILNE Epwarps, Anim.
s. vert., 10: 104 (nom. nov. pro P hippocastanum
Krener, 1836)

1919. Thais hippocastanum Lamarck, Cooke, Proc. Malacol.
Soc. London, 13: 95 (descr. radula) (non Murex hippo-
castanum LINNAEUS, 1758)

1929. Purpura (Thalessa) pseudohippocastanum DaAvutTzEN-
BERG, Contrib. Faun. Madag., p. 221 (nom. nov. pro P.
hippocastanum KiENER, 1836)

Shell: Dirty grey in colour, nodules occasionally dark
brown. Sculptured with 4 rows of blunt or spinose nodes
on the body whorl and 1 - 2 rows on earlier whorls; in-
terstices spirally striate. Interior of aperture bluish-white
or light violet, edge of labial lip purplish-brown and
denticulate; columella chocolate-brown, occasionally
banded with white, sculptured anteriorly with 1 - 2 weak
plicae.

L: 24-55mm; W: 66-79%; HA: 63 - 75%

collection of the Linnean Society in London contains 3
specimens in the box for Murex hippocastanum which
represent 2 different species; one specimen is the Man-
cinella tuberosa (ROp1NG, 1798) and 2 specimens are the
Volema galeodes (LAMARCK, 1822). LinNAgus’ taxon
cannot be simply ignored, and the most appropriate spe-
cimens conforming to Linnagus’ diagnosis and figure
citations must be selected to serve as types of Murex
hippocastanum. The following are here selected as types:
Lectotype: length- 53.7 mm; width- 44.7 mm; height
of aperture- 46.8mm; this specimen has “Mur. B”
written on the labrum and “545” on the columella. Para-
lectotype: length- 39.0mm; width- 31.8mm; height
of aperture - 34.2 mm. These 2 specimens are conspeci-
fic with the Pyrula galeodes Lamarck, 1822. The third
specimen (53.4 < 47.2 33.5mm) is the species Gale-
odes tuberosa ROpinc, 1798, and should be excluded
from the type series.

Thais armigera (Linx, 1807)

(Plate 47, Figure 2)

1807. Mancinella armigera Linx, Beschr. Nat. Samml. Univ.
Rostock, p. 115

1846. Purpura affinis ReEve, Conch. Icon., 3, pl. 13, sp. 77

1919. Thais armigera “CHEMNITZ”, Cooxe, Proc. Malacol.
Soc. London, 13: 92, fig. 10 (radula)

Shell: Dirty-white in colour, occasionally ornamented
with brown bands. Body whorl sculptured with 2 - 4
spiral rows of prominent somewhat spinose nodes, earlier
whorls with a single row; interstices spirally corded. Aper-
ture wide, white to yellowish in colour, edge of labial lip

Page 296 THE VELIGER Vol. 11; No. 4

flecked with brown, denticulate; columella white, irreg-
ularly banded with fawn or chocolate-brown, anteriorly
with weak or prominent plicae.

L: 40 - 98.0 mm; W: 65-80%; HA: 56-61%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

Distribution: Throughout the Fiji Islands, — Indo-Pacific.

Thais intermedia (KiENER, 1835)
(Plate 47, Figure 3)

1835. Purpura intermedia KiENER, Spec. gén. icon. coq. viv.,
7: 51; pl. 12, figs. 34, 34a

1919. Thais hippocastanum var. intermedia KIENER, COOKE,
Proc. Malacol. Soc. London, 13: 95, fig. 4 (radula)

Shell: Similar in form, size and sculpture to Thais acule-
ata but differing in apertural features. The aperture is
white, the edge of the labial lip has 4 - 5 squarish choco-
late-brown blotches and 25 - 30 small and sharp exterior
denticles; interior denticles white and round, numbering
from 9 - 13.

L: 30-50 mm; W: 63 - 70%; HA: 57 - 64%

Type Locality: Les cétes du Sénégal, Océan des Grandes
Indes, et la mer Pacifique.

Habitat: On reefs, under coral rocks, in shallow water.
Moderately rare.

Distribution: North and West Viti Levu. — Indo-Pacific.

Mancinella Linx, 1807

Mancinella Linx, 1807, Beschr. Nat. Samml. Univ. Rostock,
3te Abth., p. 115. Type species by T Murex mancinella
Linnaeus, 1758

Mancinella mancinella (LINNAEUS, 1758)
(Plate 47, Figure 4; Text figure 1)
1758. Murex mancinella LinNAEUuS, Syst. Nat., ed. 10: p. 751

1798. Volema alouina R6pinc, Mus. Bolten., p. 58
1798. Volema glacialis R6pinc, Mus. Bolten., p. 58

1807. Mancinella aculeata Linx, Beschr. Nat. Samml. Univ.
Rostock, p. 115

1816. Purpura gemmulata Lamarck, Tabl. Encycl. Méthod.,
p. 2; pl. 397, figs. 3a, 3b

1822. Purpura mancinella Lamarck, Anim. s. vert., 7: 239

Shell: Globose, yellowish to reddish orange in colour,
irregularly flecked with white. Sculptured with 4 spiral
rows of sharp or blunt spines and spiral cords. Aperture
wide, labrum with orange lirae, columella smooth or
irregularly plicate.

Radula: Odontophore 10.9 mm long and 0.24 mm wide
in an individual with a shell 52.7 mm in length; rows

100p
Ee)

Sem

Figure 1
Mancinella mancinella (LINNAEUS)

half row of radular teeth
Fiji Islands

of teeth number 159 (plus 13 nascentes). Rhachidian
with a long central cusp, intermediate denticles lacking,
side-cusps bifid, and laterals simple and curved.

L: 25-60mm; W: 72-80%; HA: 65 - 73%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: E. A. SmitH (1913) in an article on Murex
mancinella LINNAEUS pointed out that the Linnaean
taxon is a composite species, but failed to designate a
type or restrict the usage of M. mancinella. The Linnae-
an collection in London contains 3 specimens in a box
for M. mancinella, which were isolated by HANLEY when

Explanation of Plate 47

Figure 1: Thais aculeata (DESHAYES) « 0.75
Figure 2: Thais armigera (Link) x 0.75

Figure 3: Thais intermedia (KIENER) xX 1.0

Figure 4: Mancinella mancinella (LINNAEUS) xX 1.0
Figure 5: Mancinella tuberosa (Ropinc) x Oo
Figure 6: Drupa clathrata (LAMaARcK) x 1.2

Figure 7: Drupa morum Ropinc X1.2

Figure 8: Drupa ricinus (LinNarus) - White form X15

Figure 8a: Drupa ricinus (LINNAEUS) - Orange spotted form
x 1.5

Figure 9: Drupa marginatra (BLAINVILLE) xX 1.0

Figure 10: Drupa rubusidaeus RopiING x tO

Figure 10a: Drupa rubusidaeus ROpING X 1.0

THE VELIGER, Vol. 11, No. 4 [CeERNOHORSKY] Plate 47

Figure 4

Figure 5 Figure 8

Figure 8 a Figure 9 Figure 10 Figure 10 a

photographs by W. O. CernoHorsky

Vol. 11; No. 4

he examined the collection: 2 specimens are the species
Drupa cornus Roévinc, 1798 and 1 specimen is the Man-
cinella mancinella of authors. LinNAEvs’ original de-
scription applies only to the M. mancinella of authors,
not to the Drupa cornus Ropinc. LinnaEus described
the aperture of Murex mancinella as toothless, but
both specimens of Drupa cornus in the Linnaean collec-
tion have 5-6 prominent labial denticles. To preserve
the usage of Mancinella mancinella of authors, the fol-
lowing specimen is designated here as the lectotype of
Murex mancinella LinnAEus: length- 41.6mm; width
- 30.9 mm; height of aperture - 27.7 mm; the shell has
16 orange labral lirae and represents the species Man-
cinella mancinella of authors.

The other two specimens (43.2 25.8 & 22.4mm
and 35.5 & 23.9 & 22.8mm) have the number “544”
marked on the columella; on the larger specimen the
number “762” has been obliterated. These 2 specimens
are the Drupa cornus ROp1NG and are excluded from the
type series; they were either added by Linnagus after
1758 or were erroneously selected by HANLEy as types
of Murex mancinella.

Mancinella tuberosa (RopiNc, 1798)

(Plate 47, Figure 5; Text figure 2)
1798. Galeodes tuberosa Rovinc, Mus. Bolten., p. 53, No. 679
1798. Drupa trapa Ropinc, Mus. Bolten., p. 56, No. 709
1798. Vasum castaneum Ropinc, Mus. Bolten., p. 57, No. 716
1807. Mancinella castanca Linx, Beschr. Nat. Samml. Univ.
Rostock, 3te Abth., p. 115
1832. Purpura pica BuainvittE, Nouv. Ann. Mus. Hist. Nat.,
1: 213; pl. 9, fig. 9
1907. Purpura pica var. major Couturier, Journ. Conchyl.
55: 142

Shell: Dirty white in colour, nodes flecked with brown.
Sculptured with 2 rows of angulate nodes on the body
whorl and smaller nodules towards the base; interstices
spirally striate. Aperture wide, yellowish in colour, edge
of labial lip with small and numerous denticles and
blackish spots; the labrum has prominent orange lirae,
the columella has small plicae anteriorly. Operculum has
either an offcentral or lateral nucleus.

Radula: Odontophore 15.6 mm long and 0.27 mm wide
in an animal with a shell 49.5 mm in length; rows of
teeth number 196 (plus 13 nascentes). Rhachidians with
a long central cusp, intermediate denticles simple, outer
cusps prominent; laterals simple.

L: 30-60 mm; W: 67-77%; HA: 63-73%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

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Page 297

100u

Figure 2
Mancinella tuberosa (Ropinc)

Fiji Islands
half row of radular teeth

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Mancinella tuberosa is the type species of
Menathais Irepate, 1937. The radula of Mancinella
tuberosa is basically similar to the radula of M. manci-
nella; both species have a lirate labrum. Cooke (1919)
figured a radula of M. mancinella (as Thais gemmulata)
which had a rhachidian with simple side-cusps and no
intermediate denticles whatever, and M. tuberosa (as
Thais pica) with rhachidians equipped with short inter-
mediate denticles. It is obvious that the presence or ab-
sence of intermediate denticles is a variable character.

Drupa Ropine, 1798

Drupa Ropinc, 1798, Mus. Bolten., p. 55. Type species by SD
(RoverETO, 1899) D. morum Ropine, 1798

Discussion: Suter (1913) is generally credited with the
type species designation; however, RovERETo’s (1899)
designation is an earlier one.

The radula of Urosalpinx cinereus (Say, 1824), the
type species of Uvosalpinx Stimpson, 1865, as figured
by Stimpson (1865), does not differ in basic features
from the radula of the Drupa group. It is suggested that
Urosalpinx, on the basis of shell and radula characters,
would be more properly placed in the Thaidinae than in
the Tritonaliinae.

HerTLEIN (1960) differentiates Drupa ROpinc from
Morula ScHUMACHER by the feature of grouped and un-
grouped labial denticles respectively. This feature is not
constant, as both types of denticles can occur in the same
species.

The radula of Drupa species differs from those of
Morula in one essential characteristic: the side-denticles
of the rhachidians in Drupa are bifid to quadrifid, but
are always simple in Morula and the subgenus Cronia
H. « A. ADAMS.

Page 298

THE VELIGER

Vol. 11; No. 4

For a discussion on Canrena Linx, 1807, see under
Drupa morum Ropine.

Drupa clathrata (LAaMarck, 1816)

(Plate 47, Figure 6)

1816. Ricinula clathrata Lamarck, Tabl. Encycl. Mécth., p. 2;
pl. 395, figs. 5a, 5b

1828. Murex hystrix Woop, Ind. Test., pl. 26, fig. 50 (non
Linnaeus, 1758)

1936. Drupa rubuscaesia BottTEN-ROpinc, Hirase, Col. Jap.
shells, p. 79; pl. 110, fig. 10 (non D. rubuscaesius ROp-
Inc, 1798)

1951. Drupa rubuscaesius Ropinc, Hirase & Taxkt, Handb.
illust. shells col., pl. 110, fig. 10 (non Ropine, 1798)

1959. Drupa rubuscaesium Ropinc, 1798, Kira, Col. Ill. shells
Japan, 1: 58; pl. 23, fig. 9 (non D. rubuscaesius Rop-
ING, 1798)

Shell: Light fawn to yellowish in colour, window-like in-
terstices dark brown. Sculptured with about 6 spiral rows
of spinose nodules on the body whorl, earlier whorls with
a single row; interstices spirally corded. Edge of labial
lip orange-brown, ornamented with bifid or trifid dent-
icles; columella violet posteriorly, plicae white, inter-
stices orange-brown. Operculum moderately thin, orange-
brown in colour and with a lateral nucleus.

L: 25-35 mm; W: 80-87%; HA: 72 - 86%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Moderately rare.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Hepitey (1913) initiated the erroneous as-
sumption that Drupa clathrata is synonymous with D.
rubuscaesius, and several authors have adopted this taxo-
nomic treatment. R6prnc referred to Martini (1777),
plate 102, figures 976 and 977 for his species; the figures
cited are a convincing likeness of the orange-flecked form
of Drupa ricinus (LINNAEUS).

Drupa marginatra (BLAINVILLE, 1832)

(Plate 47, Figure 9; Text figure 3)

1832. Purpura marginatra BLAINVILLE, Nouv. Ann. Mus. Hist.
Nat. 1: 218; pl. 10, fig. 1

1853. Purpura infumuta HomBRON & JaQuiNoT, Voy. Pole Sud
Ast. & Zélée, Atlas, pl. 22, figs. 13, 14°

1854. Purpura infumata HomBRON & JAQUINOT, Voy. Pole Sud
Astr. « Zélée, text, p. 85 (emend.)

1862. Ricinula fusca Ktster, Conch. Cab. 2nd ed., 3: 26; pl.
4, fig. 16

1862. Sistrum affine Pease, Proc. Zool. Soc. London, p. 244

1871. Purpura (Sistrum) fusco-nigra DUNKER, Malak. Blatter,
18: 154

1901. Sistrum indigoferum MeEtvitt, Ann. Mag. Nat. Hist.,
7: 551; pl. 9, fig. 1

Shell: Dirty grey to dark brown in colour; sculptured
with coarse and blunt nodules, interstices of nodules ei-
ther grooved or scaly. Aperture wide, dark chocolate to
purplish-brown; labial lip with 4-5 denticles, columella
with obsolete plicae. The interior of the aperture either
bluish-white, violet, or brownish-purple, and lirate. Peri-
ostracum thick, brown in colour.

Radula: Odontophore 3.1 mm long, 0.14 mm wide in an
animal with a shell 17.8 mm long; rows of teeth number
171 (plus 9 nascentes). The central cusp of the rhachi-
dian is simple, intermediate denticles are absent and the

100n
Da ey

Figure 3

Drupa marginatra (BLAINVILLE)

Fiji Islands
half row of radular teeth

side cusps are trifid. Side-denticles number from 4 to 6.
L: 14-26mm; W: 69-75% HA: 67 - 75%

Type Locality: fles Samoa.

Habitat: Under basalt rocks, near the high tide level.
Common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.

Drupa morum Ropinc, 1798

(Plate 47, Figure 7; Text figure 4)

1798. Drupa morum Répinc, Mus. Bolten., p. 55, No. 694

1807. Canrena neritoidca Linx, Beschr. Nat. Samml. Univ.
Rostock, 3te Abth., p. 126 (restricted to Martint, 1777,
3, pl. 101, figs. 972, 973)

1816. Ricinula horrida Lamarck, Tabl. Encycl. Méth., p. 1;
pl. 395, figs. la, 1b

1817. Ricinella violacea ScHUMACHER, Ess. nouv. syst., p. 240

Shell: Creamy-white in colour, ornamented with dark
brown nodes. Sculptured with 5 - 6 spiral rows of nodules
on the body whorl and a single row on earlier whorls;
interstices spirally lirate. Aperture constricted, light or
dark violet, labial lip with 2 groups of multiple denticles
and 2 single denticles, columella plicate anteriorly. Oper-
culum dark brown with a lateral nucleus.

Radula: Odontophore 11.0 mm long and 0.19 mm wide
in an animal with a shell 39.3 mm in length; rows of

Vol. 11; No. 4

; 100n

Figure 4
Drupa morum RODING

Fiji Islands
‘half row of radular teeth

teeth number 275 (plus 16 nascentes). Rhachidian with
a simple central cusp, prominent bifid or trifid side-cusps
and 5 - 7 side-denticles.

L: 25-42mm; W: 82-94%; HA: 80-90%

Type Locality: None.

Habitat: In crevices of reefs and under coral rocks, in
shallow water. Common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: On the basis of Linx’s (1807) cited refer-
ences, his Canrena neritoidea is a composite species. The
cited figures from Martini (1777, pl. 101, figs. 972, 973)
represent the species Drupa morum Roopinec, and have
also been cited by ROpinc (1798) for his species; the
other cited figures from Martini (pl. 102, figs. 976 to
979) are Drupa grossularia Ropinc, 1798 (figs. 978,
979 only). To prevent Canrena Linx, 1807, being uti-
lized as an earlier name for Drupina Dat1, 1923, figures
972 and 973 on plate 101 in Martini (1777) are here
designated as the lectotype representatives of Canrena
neritoidea Link. Canrena neritoidea is the type species
of Canrena Link by monotypy, and thus becomes an
objective synonym of Drupa Répine.

Drupa ricinus (LinnaEus, 1758)

(Plate 47, Figures 8, 8a; Text figures 5, 6)

1758. Murex ricinus LINNAEUS, Syst. Nat., ed. 10, p. 750, No.
464 (orange-spotted form and white form)

1758. Murex hystrix LINNAEUS, Syst. Nat., ed. 10, p. 750, no.
468 (? specim. juv.)

1798. Drupa tribulus Ropinc, Mus. Bolten., p. 55, No. 695

1798. Drupa rubuscaesius Ropinc, Mus. Bolten., p. 55, No.
696 (orange-spotted form)

1816. Ricinula arachnoides Lamarck, Tabl. Encycl. Méth., p.
1; pl. 395, figs. 3a, 3b (orange-spotted form)

1832. Purpura albo-labris BuratnvittE, Nouv. Ann. Mus. Hist.
Nat., 1: 208; pl. 9, fig. 5 (white form)

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Page 299

Shell: White in colour, nodules dark brown. Generally
sculptured with 5 spiral rows of blunt or spinose brown
nodules, earlier whorls with a single row; interstices spi-
rally lirate. Aperture narrow, and either pure white or
flecked with orange; the orange flecks are situated op-

100p

»

Figure 5
Drupa ricinus (LINNAEUS) - white form
Fiji Islands
a. half row of radular teeth
b. part of rhachidian showing variation of cusps
c. penis

posite the nodules on the labial lip, one pair of flecks
borders the anal canal and a single orange streak is
situated on the columellar side of the siphonal canal.
Labial denticles are generally grouped, the columella is
prominently plicate.

Radula: Odontophore 6.2 mm long and 0.11 mm wide in
an animal with a shell 25.0 mm in length; rows of teeth
number 204 (plus 19 nascentes). Rhachidian with a
simple central cusp, side-cusps long or short, bifid to
quadrifid, side denticles number from 2 - 3; laterals are
simple.

L: 18-32mm; W: 84- 107%; HA: 82-94%

Type Locality: O. Asiatico.

Habitat: In crevices of coral rocks, on algae-matted
reef platforms, in shallow water. Common.
Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Authors usually separate the form with a pure
white aperture as Drupa albolabris (BLatNnviLLe) and the
form with an orange-flecked aperture as D. arachnoides

Page 300

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Vol. 11; No. 4

(Lamarck). The Linnaean type-series of Murex ricinus
consists of 3 specimens: 1 specimen is the form with the
white aperture, another specimen is the orange-spotted
form and the third specimen is juvenile. Linnaeus’ speci-

' 100p \

ATR ws

Figure 6
Drupa ricinus (LINNAEUS) - orange spotted form

Fiji Islands
a. half row of radular teeth
b. part of rhachidian showing variation of cusps

fic name is therefore applicable to cither form.

The specific validity of the 2 forms has been open to
question, and a more detailed study of this common spe-
cies was indicated. In the notes which follow, the form
with a pure white aperture will be referred to as the
“white form” and the form with orange flecks as the
“orange-spotted form.”

Seven localities right around Viti Levu, a circumfer-
ence of approximately 320 miles, were sampled, and
these will be referred to as stations A to G.

Station A: Nananu-i-Ra Island, off the North coast of
Viti Levu: Both forms occur in about equal numbers
and share the same habitat.

Station B: Islands of the Mamanuca group: Both forms
occur in about equal numbers and share the same
habitat.

Station C: Manava Island, off the coast of Northwest
Viti Levu: the orange-spotted form is rather more
numerous than the white form, but they share the
same habitat.

Station D: Nadroga reef, extending from Cuvu to Koro-
levu, a distance of approximately 40 miles: Both

forms are represented in about equal numbers and
share the same habitat.

Station E: Namagumagua village, South Viti Levu: Col-
lecting in January 1968 produced only the orange-
spotted form and not a single specimen of the white
form; however, 10 miles further west and one day
later, both forms were found on the same reef, under
the same conditions.

Station F: Rat-Tail Passage, main Suva reef: Here the
orange-spotted form was more numerous than the
white form, but both shared the same habitat.

Station G: Viti Levu Bay, Northeast of Viti Levu: Both
forms occur in about the same numbers and share
the same habitat.

Habitat: Both forms of the species are rare in muddy-
sand localities of Northern Viti Levu coastal reefs; the
species is more common in clean coral sand localities of
Southern coastal reefs. The species occurs in great num-
bers on algae-matted reef platforms and is more frequent
towards the reef-edge. It also lives in cracks and crevices
of reefs without algal matting and is also found on the
underside of coral rocks. Both forms may be found on
the same recf only a few feet apart or under the same
coral rock.

Shell Morphology: Apart from the different colouring of

the aperture, there is no single diagnostic character which

would separate the species; in beach-worn condition the

2 forms cannot be separated.

The majority of specimens of the orange-spotted form
generally have 7 orange spots which are arranged as fol-
lows: 2 spots border the anal canal (1 on each side), 1
spot is situated opposite the first quadrifid group of
labial denticles, another spot opposite the bifid labial
denticle, and a spot each is situated opposite the single
anterior denticles; 1 spot is placed on the columellar side
of the siphonal canal. Occasional specimens show an
orange line on the edge of the columella.

Specimens have been collected which are obvious inter-
grades between the 2 colour-forms: specimens with a
pure white aperture, apart from a thin orange line on the
labial lip; specimens with either 1 or 2 single orange spots
near the anal canal, and specimens with a single orange
spot on the labial lip.

Radula: No constant difference could be observed in ei-

Explanation of Plate 48

Figure 11: Drupina grossularia (ROpING) xX 1.5
Figure 12: Drupella cornus (Ropinc) - Female XK 1.5

Figures 12a, 12b: Drupella cornus (Ropinc) - Males x 1.75
Figure 13: Morula biconica (BLAINVILLE) X 2.0
Figures 14, 14a: Drupella ochrostoma (BLAINVILLE) X .2.0

Figures 15, 15a: Drupella rugosa (Born)
Figure 16: Drupella cf. D. angulata (REEVE)

X 1.75
X 2.25

Figures 16a, 16b: Drupella cf. D. angulata (REEVE) X 2.25
Figures 17, 17a: Morula anaxeres (KIENER) X 2.0
Figure 18: Morula biconica (BLAINVILLE) X 2.0

THE VELIGER, Vol. 11, No. 4 [CeRNOHORSKY] Plate 48

Figure 12 Figure 12 a Figure 12 b Figure 13

Figure 15 Figure 15 a Figure 16

Figure 16 a Figure 16 b Figure 17 Figure 17 a Figure 18

photographs by W. O. CernoHorsky

Wole tl Noy 4

ther the pattern of the rhachidians or laterals, nor in the
ratio of length : width of the odontophore or in the oc-
currence of either bifid, trifid, or quadrifid side-cusps.
Wu (1965) was unable to find any constant differences
in the radula of both forms, but pointed out that one
form had more numerous side-denticles than the other;
this feature I found extremely variable in both forms (see
Text figures).
Sexual Dimorphism of the Shell: None was observed;
the white and orange-spotted forms occur in both males
and females.
Sexual Dimorphism of the Radula: None was observed ;
variations which do exist are erratic in both males and
females of both forms.
Penis: The muriciform penis is common to both colour
forms and is appreciably different from the tubular penis
of Drupella THIELE.
Stomach Pouch: This was found to be either orange-red
or brown in both colour forms of both sexes.
Gut Content: Same in both forms, consisting of coral
fragments, sponge spicules and algal matter.
Living Animal: The short, stubby tentacles may be either
plain brownish-fawn or may have an additional blackish-
brown transverse band on the tentacles in either form.
Both forms have been reported from widely-scattered
Indo-Pacific localities. HErTLEIN (1960) reports both
colour forms from the Galapagos and Clipperton Islands.
Evidence on hand strongly suggests that the two forms
are conspecific. The occurrence of 2 colour forms in a
species is by no means confined to Drupa ricinus, but
can be observed in Strombus gibberulus gibbosus (Rop-
ING, 1798), Duplicaria duplicata (LinNAEus, 1758),
and Conus marmoreus LinNAgEus, 1758, and _ several
other gastropods. This once again confirms the futility
of using shell-colour as a guide for specific diagnosis of
molluscs.

Drupa rubusidaeus Ropine, 1798

(Plate 47, Figures 10, 10 a)

1798. Drupa rubusidaeus Rovinc, Mus. Bolten., p. 55, No. 698

1807. Mancinella hystrix Linx, Beschr. Nat. Samml. Univ.
Rostock, p. 115

1822. Ricinula miticula Lamarck, Anim. s. vert., 7: 231

1828. Murex hippocastanum Wooo, Ind, Test., pl. 26, fig. 53
(non Murex hippocastanum Linnaeus, 1758)

1832. Purpura spathulifera BLAINvILLE, Nouv. Ann. Mus. Hist.
Nat., 1: 212; pl. 9, fig. 8

1862. Ricinula reeveana CrossE, Journ. Conchyl. 10: 47; pl.
1, fig. 3

Shell: Yellowish in colour, generally heavily encrusted
with coral growth. Sculptured with 4-6 spiral rows of

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Page 301

bluntly spinose nodes, earlier whorls with a single row.
Edge of aperture yellow or orange, labial lip rose-violet
and ornamented with 7-9 denticles which may extend
into the aperture and become whitish within. Columella
rose-violet in colour, irregularly flecked with yellow or
orange, and sculptured with plicae anteriorly; the edge
of the columellar callus occasionally has 6 - 7 small den-
ticles. Operculum moderately thin, orange-brown in col-
our and with lateral nucleus.

L: 25-58mm; W: 78 -94%; HA 76 - 87%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: The species Drupa rubusidaeus has a con-
fused nomenclature, and R6p1no’s specific name has oc-
casionally been applied to the species Drupa clathrata
(Lamarck). The confusion probably started with Hep-
LEY (1913) who placed Ricinula clathrata Lamarck, R.
speciosa DUNKER, 1867 and Purpura spathulifera BLAIN-
VILLE in the synonymy of Drupa rubusidaeus Ropinc;
several Japanese workers subsequently adopted HEDLEY’s
synonymy. The identification of D. rubusidaeus depends
on the interpretation of Ropinc’s (1798) cited figures
from Martini (1777, 3: 283, pl. 101, figs. 974, 975). It
has to be admitted that the cited figures are somewhat
ambiguous, and could represent in part either D. clath-
rata (LaMarcK) or D. rubusidaeus ROpiNnc. The general
form, numerous spines, and shape of the juncture of
the aperture would favour D. clathrata, but the colouring
and general aspect of the apertural features strongly
favour D. rubusidaeus. MARTINI (op. cit.) called his spe-
cies “Der Igel,” and referred to Guattierr (1742), pl.
28, fig. N as a reference; it is unfortunate that these
figures represent the species D. ricinus (LinNAEUS). No
denticles are visible in the Martini figures of D. rubusi-
daeus, but they are mentioned by Martini in the descrip-
tion. Although ambiguous, the figures tend to represent
the same species as the Purpura spathulifera BLAINVILLE,
and R6p1Ne’s earlier name has been accepted in this
paper.

DunKeErR (1867) described a Ricinula speciosa (non
Purpura speciosa VALENCIENNES, 1832), but in 1870 (p.
139) he stated that his R. speciosa is a smaller and mature
example of R. reeveana Crosse. The United States Na-
tional Museum collection has numerous specimens of
Drupa speciosa from Polynesia, and it is evident that
this is a valid species confined to Polynesia, and inter-
mediate in characters between D. clathrata and D. rubu-
sidaeus. EMERSON in WEAVER (1966) reported the species
from the Fiji Islands, but it does not occur there. Oos-

Page 302

THE VELIGE

Vol. 11; No. 4

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Vol. 11; No. 4

THE VELIGER

Page 303

TINGH (1931) reports D. hystrix var. speciosa DUNKER
from Sumatra, but this record needs clarification.

The types of Ricinula miticula Lamarck are in the
Muséum d’Histoire Naturelle in Geneva; the holotype
measures 26.3 mm, the paratype 26.0 mm; they are the
species Drupa rubusidaeus Ropinc. The holotype of Rz-
cinula reeveana Cross: is in the British Museum (Natural
History) in London; it measures 48.2 38.8 mm; this spe-
cies which was described from Nouhiva (=Nukuhiva),
Marquesas, is not conspecific with D. speciosa (DUNKER),
but is a large, immature example of D. rubusidaeus ROp-
ING with a bright maroon coloured aperture.

Drupina Dat, 1923

Drupina Dax, 1923, Proc. Acad. Nat. Sci. Phila., 75: 303.
Type species by OD Ricinula digitata Lamarck, 1816
= Drupa grossularia Rovinc, 1798

Discussion: The shell of Drupina is similar to that of
Drupa Rooprne, but the sculpture is more scabrous and
minutely foliated, labial teeth are smaller and grouped,
although they may be paired, and the columella is heav-
ily calloused. The labial digitations are extended.

The radular pattern of Drupina differs appreciably
from any known radula of other thaidine genera, and
fully supports DaLL’s generic separation on morphologi-
cal features of the shell. The genus is at the present time
monotypic and contains an Indo-Pacific species.

Drupina grossularia (Ropinc, 1798)

(Plate 48, Figure 11; Text figure 7)

1798. Drupa grossularia Rovinc, Mus. Bolten., p. 55, No. 700

1816. Ricinula digitata Lamarck, Tabl. Encycl. Méth., p. 2;
pl. 395, figs. 7a, 7b

1817. Ricinella dactyloides ScHUMACHER, Ess. nouv. syst., p.
241

1828. Murex ricinus Woop, Ind. Test., pl. 26, fig. 51 (non
Linnaeus, 1758)

1832. Purpura lobata BLAINvILLE, Nouv. Ann. Mus. Hist. Nat.,
1: 210; pl. 9, fig. 7 (Indian Ocean subspecies)

1850. Purpura laurentiana PETIT DE LA SAUSSAYE, Journ. Con-
chyl., 1: 403; pl. 13, fig. 2 (specim. juv.)

1965. Drupina glossularia (Ropinc), Wu, Bull. Inst. Zool.
Acad. Sin., 4: 99 (invalid emendation)

1967. Drupina glossularia (LinNE), Hase & Kosuce, Std.
Book Japan. shells col., p. 70; pl. 27, fig. 22

Shell: Uniformly creamy-white to yellow in colour. Sculp-
tured with about 5 heavy spiral cords and intermediate
scabrous lirae on the body whorl, and a single row of
obsolete and indistinct nodules. Aperture narrow, orange
in colour, edge of labial lip with 5 digitations which are

actual extensions of the spiral cords; the labial lip has
prominent white nodules and the columella is heavily
calloused and plicate; the edge of the labial callus is
sometimes finely denticulate. Operculum brown in colour,
with a lateral nucleus.

Radula: Odontophore 6.5 mm long and 0.14 mm wide in
an animal with a shell 30.3 mm in length; rows of teeth

100. :

CLA

Figure 7
Drupina grossularia (ROop1NGc)
Fiji Islands
half row of radular teeth

number 271 (plus 17 nascentes). The rhachidian is
equipped with about 13 moderately short cusps of al-
most uniform size, the side-cusps are trifid; laterals small
and simple.

L: 22-36mm; W: 96- 113%; HA: 85-91%

Type Locality: None.

Habitat: On reefs, near reef’s edge, in crevices and under
coral rocks, in shallow water. Moderately common.
Distribution: Throughout the Fiji Islands. — Pacific.
Discussion: Drupina lobata (BLAINVILLE, 1832) is a
subspecies or colour form of D. grossularia with a dark
brown-maroon aperture, and appears to be restricted to
the Indian Ocean.

Drupella Turere, 1925

Drupella THIELE, 1925, Wissensch. Erg. Exp. Valdivia, 17: 171.
Type species by SD (TurEte, 1929) Drupa (Drupella)
ochrostoma (BLAINVILLE, 1832) = Purpura ochrosto-
ma BLAINVILLE, 1832

Discussion: In shell features, species of Drupella re-
semble Morula ScHuMACHER, 1817 and Cronia H.& A.
ApamMs, 1853, or even some species of Maculotriton
Dati, 1904. They are a small compact group whose radu-
la bears no similarity to any other muricid genus, with
the possible exception of Vexilla Swarnson, 1840. Spe-
cies of Vexilla also have a plain or serrated rhachidian;
the laterals, although not as long as those of Drupella,
are more elongate than the typical muricid lateral, and
are also either simple or denticulate (see Cooke, 1919,
and TureLe, 1929). The penis also differs in structure

Page 304

THE VELIGER

Vol. 11; No. 4

from that of Drupa s. str.; it is stout, stubby and tubular,
and has a large oval seminal duct exit.

Species of Drupella are widely distributed in the Indo-
West Pacific. They inhabit a variety of habitats and are
not confined to branching coral (fide Cooke, 1895 and
Demonp, 1957) ; they may be found in branching coral,
in crevices of algae-matted reef platforms and on the
underside of coral blocks which litter Fiji reefs, and
occur on coastal reefs as well as outlying islets.

Drupella cornus (Ropine, 1798)

(Plate 48, Figures 12, 12a, 12b; Text figure 8)

1798. Drupa cornus Ropinc, Mus. Bolten., p. 56, No. 704
(female form)

1832. Purpura nassoidea BLAINvILLE, Nouv. Ann. Mus. Hist.
Nat., 1: 205 (male form)

1832. Purpura elata BLAINvILLE, Nouv. Ann. Mus. Hist. Nat.
1: 207; pl. 11, fig. 1 (female form)

1833. Purpura nassoides Quoy & GatmarD, Voy. Astrolabe,
Zool., 2: 564; pl. 38, figs. 7, 8, 9 (male form)

1839. Purpura (Ricinula) martiniana ANTON, Verz. Conch.,
p. 88

1846. Ricinula spectrum REEvE, Conch. Icon., 3, pl. 3, sp. 19
(female form)

1846. Ricinula dealbata ReEEvE, Conch. Icon., 3, pl. 4, sp. 26

1852. Ricinula eburnea Kuster, Conch. Cab., 2™4 ed., 3: 17;
pl. 3, fig. 9

1853. Purpura alba HomBRON & Jagutnot, Voy. Pole Sud Ast.
« Zélée, Atlas, pl. 22, figs. 30, 31 (specim. juv.)

1918. Drupa vitiensis Prrspry & Bryan, Nautilus 31, pl. 9,
fig. 5 (dark noduled male form)

1921. Sistrum vitiense Prtspry, Proc. Acad, Nat. Sci. Phila.,
72: 319

1967. Drupella mancilla [sic] LinNE, Habe « Kosuce, Std.
Book Japan. shell, p. 70; pl. 27, fig. 20 (female form)

1967. Drupa (Drupella) chaidea (Ductos), Orr-Maes, Proc.
Acad. Nat. Sci. Phila., 119: 129; pl. 11, fig. 6 (male
form)

Shell (Female): White in colour throughout. Sculptured
with 4 spiral rows of prominent and pointed nodes, earlier
whorls with a single row; interstices of nodules with 3 - 5
spiral grooves. Aperture narrow, porcellaneous white,
labial lip with from 5-8 moderately sized denticles,

lower half of columella with 2 -5 irregular short plicae.
Anal notch indistinct, base with a heavy callus.

Shell (Male): Generally smaller than that of females,
but rare specimens may attain the same size. The shell
is more bulbous in appearance and the sculpture is ap-
preciably more discrete. ‘The spiral nodules are more nu-
merous, smaller and blunter, and the shell is spirally
corded, not grooved; the aperture appears wider and
labial denticles smaller.

Radula: Odontophore brown in colour, 8.0 mm long and
0.28 mm wide in an animal with a shell 38.0mm in
length; rows of teeth number 264 (plus 33 nascentes).
Rhachidian with a massive central cusp, moderately
smaller outer cusps, thin and sharp intermediate den-
ticles and 8 - 13 serrated denticles at either side of the
central cusp; laterals are long and thin, ca. 6 times as
broad as the rhachidian, sharply and minutely denticu-
late at the base, distal end curved and pointed, and
either simple or bifid.

Eighteen specimens from various Fijian localities were
examined and no sexual dimorphism of the radula was
evident. Both males and females had the same type of
rhachidian; the curved distal end of the laterals was
simple in most specimens, bifid in others, and 1 male
specimen 22.3 mm in length showed both simple and bi-
fid tips on laterals in the same ribbon.

The inner part of the base of the lateral teeth is
obviously also used in abrading food particles; in the first
one dozen or more rows of teeth, the inner denticles were
completely worn off and came back to normal size as the
rows progressed towards the end with the nascentes.

L: 18-50mm; W: 51-61%; HA: 48-57%

Type Locality: None.

Habitat: On reefs in crevices and under coral rocks, in
shallow water. Common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: ARAKAWA (1958) figured the radulae of
Drupella cornus (RopiNG) and D. fragum (BLAINVILLE),
and reported on the sexual dimorphism of the radula in
the two species. His text figures 1 and 2 of D. fragum are
both immature specimens of D. cornus; the rhachidians
and laterals of the radula of D. cornus depicted by Ara-

Explanation of Plate 49

Figure 19: Morula granulata (Ductos) X 2.0
Figure 20: Morula nodicostata (PEASE) xX 3.0
Figure 21: Morula parva (REEVE) 352
Figures 22, 22a: Morula spinosa (H. « A. ADAMS)
Figures 23, 23a: Morula uva (Rop1nc) xX 1.5

ln?

Figure 28: Vexilla vexillum (GMELIN)
Figure 29: Nassa serta (BRUGUIERE)

Figure 24: Morula (Cronia) aurantiaca (HOMBRON & JAQUINOT)

x 15
Figure 25: Morula (Cronia) fiscella (GMELIN) X 1.2
Figure 26: Morula (Cronia) margariticola (BRoDERIP) xX 15
Figure 27: Maculotriton egregius (REEVE) X15
xX 1.3
xX 1.0

THE VELIGER, Vol. 11, No. 4 [CeRNoHOoRSky] Plate 49

Figure 21 Figure 22

Figure 22 a Figure 23 Figure 23 a

Figure 25 Figure 26 Figure 27 Figure 28 Figure 29

photographs by W. O. CernoHorsky

Vol. 11; No. 4

THE VELIGER

Page 305

KAWA on plate 6, figures 5 and 12 fully agree with the
radula of D. cornus from the Fiji Islands. In the 8 males
and 10 females of D. cornus from Fiji waters examined,
no difference in the pattern of rhachidians between the
sexes was observed; a sexual dimorphism in the shell,
however, was evident. In the juvenile stage (thin labial
lip devoid of denticles) the shells of both sexes are the
same, i. e., they are barrel-shaped with blunt small nod-
ules and distinct spiral cords. In fully mature specimens,
however, all the females examined had a heavy and large
shell with prominent and spinose nodes, while all adult
males retained the barrel-shaped form with blunt nodules
and prominent spiral lirae. Both forms occurred in small
colonies under the same rock and were observed in mating
position.

The reed-like laterals of Drupella cornus are always
serrated for a short distance at the base, whereas they are
smooth in D. rugosa (Born, 1778) and D. ochrostoma
(BLAINVILLE, 1832). No side-denticles on the rhachidians

100p. ,

200u

Figure 8

Drupella cornus (Rop1nc)

Fiji Islands
a. half row of radular teeth
b. basal portion of lateral teeth - enlarged
c. distal end of lateral teeth - enlarged
d. penis

of D. rugosa (=D. fragum of ARAKAWA) were observed
in any of the specimens examined. The distal end of the
laterals may, as pointed out by ARAKAWA (op. cit.) be
either bifid or simple. In view of the discrepancies be-
tween ARAKAWA’s and my own findings, it is suspected
that the differences in radula pattern are either regional
or a third similar species may be involved.

The male form of Drupella cornus has been occasion-
ally listed as Purpura chaidea Ductos, 1832. Ductos’
species, however, is a nassarid-shaped species with prom-
inent and thick axial folds which are decussated by close-
set spiral striae.

The species Ricinula siderea REEVE, 1846, is generally
placed in the synonymy of Drupella cornus (R6pINGc).
REEvE’s 3 syntypes are in the British Museum (Natural
History), London, and from examination of the types it
is evident that Rerve’s Ricinula siderea is a columbellid
species and not a muricid; the holotype measures 12.8 by
6.4 mm.

Drupella ochrostoma (BLAINVILLE, 1832)
(Plate 48, Figures 14, 14a; Text figure 9)

1832. Purpura ochrostoma BiainviLte, Nouv. Ann. Mus. Hist.
Nat., 1: 205

1833. Purpura nassoides yar. Quoy & GarmarD, Voy. Astro-
labe, Zool., 2: 564; pl. 38, figs. 10, 11 (non BLaINvILLE,
1832)

Shell: Whitish to creamy-yellow in colour, nodules oc-
casionally brown. Sculptured with 4 spiral rows of small,
blunt and often brownish nodules (ca. 9-12 per row)
which decrease in size towards the base; the penultimate
whorl with a single or rarely double row of nodules.
Interstices sculptured with 2 regular and scabrous spiral
cords. Aperture orange, labial lip with 5-6 prominent
white denticles; siphonal canal short and calloused.

Radula: Odontophore 7.0 mm long and 0.15 mm wide in

100pn

=

00
200K |

Figure 9
Drupella ochrostoma (BLAINVILLE)

Fiji Islands
half row of radular teeth

Page 306

an animal with a shell 22.5 mm in length; rows of teeth
number 205 (plus 26 nascentes) and early rows are worn.
Rhachidians with a massive central cusp, thick outer
cusps and small intermediate denticles; laterals long,
slender and simple, curved at the distal end, and approx-
imately 9 times as broad as the rhachidians.

L: 17-25mm; W: 58-60%; HA: 56 - 63%

Type Locality: Tonga.

Habitat: On reefs, in crevices and under coral rocks, in
shallow water. Uncommon.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: BLAINvILLE’s (1832) reference to figure 8 is
possibly an error for figure 10, which depicts a specimen
with an orange aperture. Figures 7 - 9 are Purpura nasso-
idea BLAINVILLE, 1832.

Drupella rugosa (Born, 1778)

(Plate 48, Figures 15, 15a; Text figure 10)
1778. Murex rugosus Born, Ind. rer. nat. Caes. Vindob., p. 303
1822. Murex concatenatus Lamarck, Anim. s. vert., 7: 176

1832. Purpura fragum BuainvitLtE, Nouv. Ann. Mus. Hist.
Nat., 1: 203; pl. 9, fig. 4

Shell: White in colour, rarely creamy-yellow, nodules oc-
casionally dark brown. Sculptured with 3 - 4 spiral rows
of prominent nodules, penultimate whorl with a single
row; nodules are connected by scabrous spiral cords, and
interstices are ornamented with 3-5 spiral ridges, base
of shell with 2-3 heavy cords; some specimens show a
distinct varix on the body whorl. Aperture white, labial
lip with 6-10 denticles which are uniformly spaced,
lower half of columella with 3 - 4 short plicae; siphonal
canal short, stained with orange anteriorly.

Young specimens are scabrous, dark orange in colour
with the nodules dark brown; penultimate whorl carries
2 rows of nodules which in the adult stage fuse into a
single row; interstitial spiral cords very weak.

Radula: Odontophore brown in colour, 8.7mm _ long
and 0.23 mm wide in an animal with a shell 23.0 mm in
length; rows of teeth number 166 (plus 21 nascentes).
Rhachidians with a massive central cusp, prominent outer
cusps and with or without an intermediate denticle. Lat-
erals long and about 8 times as broad as the rhachidians,
simple and curved at the distal end.

L: 20-35mm; W: 52-56%; HA: 55 - 60%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water,
Uncommon.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Born (1778) cited a figure from CHEMNITZ
(1780, pl. 124, figs. 1155, 1156) for his species, which is
a rather indifferently executed figure for the species un-

THE VELIGER

Vol. 11; No. 4
100p
200u

Figure 10
Drupella rugosa (Born)

Fiji Islands
half row of radular teeth

der discussion; LAMARCK’s Murex concatenatus was based
on the same Cuemnirz figure. In addition, Lamarck
(1822) cited Lister (1685-92, pl. 954, fig. 5) which is
Morula uva (Ropinc, 1798), while the reference to
Knorr (1769, pl. 26, fig. 2) could just about represent
any similar species. BLAINvVILLE (1832) described his
Purpura fragum as yellowish-white on the outside, with
dark rose nodules and a completely white aperture. Fiji
specimens agree with BLAINVILLE’s diagnosis of a white
aperture, but specimens with a yellowish-orange aperture
do occur in other Pacific localities.

The shells of Drupella cornus and D. rugosa may ap-
pear similar when both are adult and heavily encrusted
with coral growth; juvenile shells of both species, how-
ever, differ quite prominently.

The species Drupella rugosa from Fiji has a different
radula pattern than has been figured by ARAKAWA
(1958, as D. fragum), and no sexual dimorphism of the
radula has been observed in this species.

Drupella c.f. D. angulata (REEVE, 1844)
(Plate 48, Figures 16, 16a, 16b; Text figure 11)

1844. Triton angulatus REEVE, Conch. Icon., 2, pl. 9, sp. 88
Shell: Small and moderately slender, variable in colour,
but generally white or yellow, occasionally ornamented
with spiral rows of black spots or orange-brown trans-
verse zones. Whorls number from 5 - 6, and protoconch
is not distinguishable in adult specimens. Body whorl
sculptured with 1-3 rows of prominent or feeble and
blunt nodules, while the penultimate whorl has either a
single row of nodules or from 10 - 13 angulate and nod-
ulose axial ribs; body whorl has an additional 12 - 20
moderately coarse spiral cords which may override the
nodules. Aperture white, labial lip ornamented with 5 - 6

Vol. 11; No. 4

50

200K

Figure 11
Drupella cf. D. angulata (REEVE)

Fiji Islands
a. half row of radular teeth
b. distal end of lateral teeth

small denticles, columella smooth; anterior canal short
and open, anal canal prominent and almost resembling
a daphnelloid notch.

Radula: Odontophore 4.6 mm long and 0.12 mm wide in
an animal with a shell 12.5 mm in length; rows of teeth
number 120 (plus 8 nascentes) and early rows are worn.
Rhachidians with a long and slender central cusp, inter-
mediate denticles small but slender, side cusps broad;
laterals long and slender, 10 times as broad as the rhachi-
dians, the distal end terminates in a simple curved cusp
which has at the base a smaller accessory cusp.

L: 9- 20mm; W: 49-58%; HA: 52 - 60%
Distribution: Throughout the Fiji Islands. — ?
Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

Discussion: This species, although moderately common in
Fiji, is not represented in the rather comprehensive mol-
luscan collections of the U.S. National Museum. The
species has been tentatively associated with REEvE’s
Triton angulatus as his description and figure appear
fairly compatible with the Fiji species.

Morula ScHuMacHeER, 1817

Morula Scuumacu_Er, 1817, Ess. nouv. syst., pp. 68, 227.
Type species by M Morula papillosa ScHUMACHER,
1817 = Drupa uva Roving, 1798

Discussion: ARAKAWA (1965) established the new genus
Tenguella with Purpura granulata Ductos, 1832 as the
type species; he erected his new genus on the basis of
differences in the number of side-denticles of the rhachid-
ians of the radula in Morula uva (ROpINc) and M.
granulata (Ductos). Wu (1965a) commented on the
variability of the number of side-denticles in the two

THE VELIGER

Page 307

species under discussion and doubted the validity of
Tenguella. The number of side-denticles was found to
vary in both species from 1-3 and the length of these
denticles also was found to be variable; one can see no
merit in retaining this monotypic genus in muricid no-
menclature.

The type species of Morulina Dati, 1923 is Ricinula
mutica LAMARCK, 1822 by original designation, and not
Morulina ceylonica Dax, 1923 as pointed out by ArRa-
KAWA (1965); the latter species is an immature speci-
men of Morula granulata (Ductos, 1832).

The side-cusps of the rhachidians in Morula are always
simple and not bifid to quadrifid as in Drupa.

Morula anaxeres (KIENER, 1835)

(Plate 48, Figures 17, 17a; Text figure 12)

1835. Purpura cancellata KiENER, Spec. gén. icon. coq. viv.,
p- 25; pl. 7, figs. 16, 16a (non Quoy « Garmarp, 1833)
1835. Purpura anaxeres KIENER, Spec. gén. icon. coq. viv., p.

26; pl. 7, figs. 17, 17a
Shell: Variable in colour, dirty grey or brown, nodules
occasionally white; interstices sometimes banded with
brown. Sculptured with spiral rows of nodules and striae.
Labial lip either dark purplish-brown or with occasional
white flecks; labial denticles number from 4-5. Columella
either fully dark purplish-brown or brown, occasionally
streaked with white or light violet, sculptured with 2-4

small and irregular plicae.

Figure 12

50p
SEE.

Morula anaxeres (KIENER)

Fiji Islands
half row of radular teeth

Radula: Odontophore 3.4 mm long and 0.14 mm wide in
an animal with a shell 25.0 mm in length; rows of teeth
number 94 (plus 4 nascentes). Rhachidians with a large
central cusp, smaller side-cusps, small intermediate den-
ticles and 0 - 2 side denticles.

L: 10-28mm; W: 61-79%; HA: 63-70%

Type Locality: L’ile Tycopia, des Nouvelles Hebrides.
Habitat: Under basalt rocks, near high tide level, rarely
under coral rocks in the intertidal zone. Common.
Distribution: Throughout the Fiji Islands. — Indo-Pacific.

Page 308

Morula biconica (BLAINVILLE, 1832)

(Plate 48, Figures 13, 18; Text figures 13, 14)

1832. Purpura biconica BuLatInviILLE, Nouv. Ann. Mus. Hist.
Nat., 1: 203; pl. 9, fig. 1

1846. Ricinula bicatenata REEVE, Conch. Icon., pl. 6, sp. 48

1868. Engina variabilis Peasr, Amer. Journ. Malac., 3: 275;
pl. 23, fig. 9

Shell: Small, whitish to dark grey in colour, pointed at
both ends. Sculptured with blackish axial ribs which are
generally spinose, and white, prominent and_ scabrous

cae

Figure 13
Morula biconica (BLAINVILLE)

Fiji Islands
half row of radular teeth

\ 50H j

TARE oi

Figure 14
Morula biconica (BLAINVILLE)
Fiji Islands

a. half row of radular teeth
b. part of rhachidian showing variation of cusps

spiral cords. Edge of labial lip white and scabrous, aper-
ture angulate and narrow, violet to dark purple in colour;
columella irregularly plicate anteriorly.

Radula: Odontophore 1.16 mm long and 0.06 mm wide
in an animal with a shell 11.5 mm in length; rows of
teeth number 69 (plus 3 nascentes). Rhachidians have a
large central cusp, short or long intermediate denticles,
simple side-cusps and 2 - 3 side denticles; laterals simple
and curved.

L: 8-17 mm; W: 57-65%; HA: 56 - 63%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.

THE VELIGER

Vol. 11; No. 4

Morula granulata (Ductos, 1832)

(Plate 49, Figure 19; Text figures 15, 16)

1832. Purpura granulata Ductos, Ann. Sci, Nat., 26: 9; pl. 2,
fig. 9

1832. Purpura tuberculata BLAINvILLE, Nouv. Ann. Mus.
Hist. Nat., 1: 204; pl. 9, fig. 3

1835. Purpura tuberculata cingulifera KiENER, Spec. gén.
icon coq. viv., pl. 5, fig. 10a

1908. Sistrum chrysalis Sowersy, Proc. Malac. Soc. London,
8: 17; pl. 1, fig. 5

1923. Morulina ceylonica Dau, Proc. Acad. Nat. Sci. Phila.,
75: 305 (specim. juv.)

Shell: Dirty white in colour, ornamented with spiral rows
of brown nodules. Sculptured with 5-6 spiral rows of
squarish but rounded large brown nodules, earlier whorls
with 1 - 2 rows; interstices finely spirally lirate. Aperture
violet, edge of columella and labial lip dark purple-
brown; labial lip with 4 - 5 denticles which are generally
bluish-white, columella with 2-4 sinuous plicae. Oper-
culum with a lateral nucleus.

100u
Ss)

Figure 15
Morula granulata (Ductos)

Fiji Islands
half row of radular teeth

100u,

Figure 16

Morula granulata (Ductos)

Fiji Islands
Rhachidian of radula

Radula: Odontophore 5.6 mm long and 0.14 mm wide in
an animal with a shell 26.0 mm in length; rows of teeth
number 203 (plus 12 nascentes), Rhachidians with a

Vol. 11; No. 4

THE VELIGER

Page 309

large central cusp, smaller side-cusps and small inter-
mediate denticles; the side denticles number from 1 -3,
but one specimen examined almost lacked side denticles
altogether, apart from a weak hump on one side.

L: 15-30mm; W: 62-73%; HA: 50-69%

Type Locality: Nouvelle-Hollande.

Habitat: On reefs, under coral rocks, in shallow water.
Common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.

Morula nodicostata (PEASE, 1868)

(Plate 49, Figure 20; Text figure 17)
1868. Engina nodicostata Peasz, Amer. Journ. Conch., 3:
274; pl. 23, fig. 8
1909. Engina purpureo-cincta Preston, Rec. Ind. Mus., 3: 136

Shell: Small, light violet in colour, nodules white, inter-
stices occasionally dark brown. Sculptured with 3 spiral
rows of white round nodules; the nodules nearest the
suture are rather large and are bisected by 3-4 spiral
cords; earlier whorls with a single row of nodules. Inter-
stices smooth and ornamented with dark brown blotches.
Aperture violet in colour, interior of aperture with a
broad, white median band; labial lip has 6 small denticles,
columella 2-3 short plicae.

50p ;

Figure 17
Morula nodicostata (PEASE)

Fiji Islands
half row of radular teeth

Radula: Odontophore 1.28 mm long and 0.06 mm wide
in an animal with a shell 9.8mm in length; rows of
teeth number 110 (plus 6 nascentes). Rhachidians have
a moderately long central cusp, short intermediate den-
ticles, simple side cusps and 2 - 3 side denticles; laterals
simple and curved.

L: 7-11mm; W: 61-67%; HA: 60-64%

Type Locality: Paumotus.

Habitat: On reefs, under coral rocks, in shallow water.
Uncommon.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Orr-Mars (1967) figured the radula of
Morula nodicostata from Cocos-Keeling Islands (Indian

Ocean) ; the particular specimen examined lacked side-
denticles on the rhachidians of the radula. It is obvious
that the presence or absence of such side-denticles in the
Morula-Cronia group of species is a variable feature,
and has been observed in other species of Morula.

Morula parva (Reeve, 1846)

(Plate 49, Figure 21)

1846. Ricinula parva Reeve, Conch. Icon., pl. 6, sp. 43

1846. Ricinula echinata Reeve, Conch. Icon., pl. 6, sp. 54

1860. Engina nodulifera PEAsE, Proc. Zool. Soc. London, p.
142

1893. Sistrum angulatum Sowersy, Proc. Malac. Soc. Lon-
don, 1: 46; pl. 4, fig. 3

Shell: Small, white in colour, ornamented with white and
orange nodules and spines. Sculptured with a row of
orange nodules at the sutures and prominent white or
orange spinose spiral cords; the penultimate whorl has
one row of orange nodules at the sutures and 2 spinose
spiral cords; interstices finely axially striate. Aperture
narrow and elongate, white in colour, interior of aper-
ture with a dark purplish-brown band; labial lip has 4 - 5
prominent white nodules and the columella 2 - 3 plicae.
L: 6-10 mm; W: 56-62%; HA: 56-60%

Type Locality: Island of Luzon, Philippines.

Habitat: On reefs, under coral rocks, in shallow water.
Distribution: Throughout the Fiji Islands. — Indo-Pacific.

Morula spinosa (H. & A. Apams, 1853)
(Plate 49, Figures 22, 22a)

1846. Ricinula chrysostoma DesHAYES, REEVE, Conch. Icon.,
pl. 2, fig. 12b only (non Desuayes, 1844)

1853. Pentadactylus (Sistrum) spinosus H. « A. Apams, Gen.
rec. Moll., 1: 130 (nom. nov. pro Ricinula chrysosto-
ma REEVE, 1846)

1853. Murex iostomus A. Apams, Proc. Zool. Soc. London,
p. 267

1862. Ricinula chrysostoma DEsHayEs, Kuster, Conch. Cab.,
274 ed., p. 24; pl. 4, fig. 7 (non Desuayes, 1844)

1909. Sistrum andrewsi E. A. Smiru, Proc. Malac. Soc. Lon-
don, 8: 369; 2 figs.

1923. Morula ambusta Datu, Proc. Acad. Nat. Sci. Phila.,
75: 304 (nom. nov. pro Ricinula chrysostoma REEVE,

1846)

Shell; Variable in colour, uniformly whitish, yellow or
grey, occasionally ornamented with brown spiral cords
and spinose nodes. Sculptured with about 9 spinose vari-
ces on the body whorl which are arranged in 3 - 4 spiral
rows; penultimate whorl with a single row of spinose
nodes. Interstices sculptured with 4-10 brown spiral
cords. Aperture elongate and narrow, yellow to orange or

Page 310

violet in colour, labial lip with about 5 small denticles;
columella with 0-5 obsolete short plicae anteriorly.
L: 18-38mm; W: 63-72%; HA: 57-67%
Type Locality: Islands of Bohol and Ticao, Philippines.
Habitat: On reefs, under coral rocks, in shallow water.
Uncommon.
Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: The holotype of Murex iostomus A. ADAMS
from the Philippine Islands, ex coll. Cuming, no. 196572
— 23.0 < 17.8 mm, is in the British Museum (Natural
History) ; it is an immature specimen with dark brown
fronds. The types of Sistrum andrewsi E. A. SmitH from
Christmas Island (Indian Ocean) are in the British Mu-
seum collection; the holotype no. 1909.5.8.62 — 25.4 mm
in length is a fully mature specimen of the species under
discussion.

Hase & Kosuce (1966, pl. 20, fig. 12) figure this spe-

cies as Thaisiella rugosa (Born).

Morula uva (Ropinc, 1798)

(Plate 49, Figures 23, 23a; Text figure 18)

1798. Drupa uva Rovinc, Mus. Bolten., p. 56, No. 703

1816. Ricinula aspera Lamarcx, Tabl. Encycl. Méth., pl. 395,
figs. 4a, 4b

1816. Ricinula nodus Lamarcx, Tabl. Encycl. Méth., p.2;
pl. 395, figs. 6a, 6b

1817. Morula papillosa ScHuMacuER, Ess. nouv. syst., p. 227

1822. Ricinula morus LAMaARCcK, Anim. s. vert., 7: 232

1832. Purpura sphaeridea Ducios, Ann. Sci. Nat., 26: 9;
pl. 2, fig. 10

1852. Ricinula alba Morcn, Cat. Conch. Yoldi, p. 87

1868. Sistrum striatum Pease, Amer. Journ. Conch., 3: 276;
pl. 23, fig. 12

1966. Morula nodilifera [sic] HaBE « Kosuce, Shells world
col., 2: 14; pl. 20, fig. 14 (specim. juv.) (non Purpura
nodulifera MENKE, 1829)

Shell: White in colour, spiral rows of nodules or short
spines blackish-brown. Sculptured with 5 spiral rows of
small, blackish-brown nodules, early whorls with 2 such
rows; interstices of nodules with 1-4 scabrous spiral
cords. In some forms the nodules are spinose, may co-
alesce and form ill-defined blackish-brown axial folds.
Aperture narrow, violet in colour, labial lip with about 4
denticles, basal denticles usually smaller, columella with
2 - 4 short plicae.

Radula: Odontophore 2.1 mm long and 0.09 mm wide in
an animal with a shell 20.6 mm in length; rows of teeth
number 162 (plus 8 nascentes). Rhachidian with a large
central cusp, smaller side-cusps and small intermediate
denticles; side denticles number from 1 - 3.

L: 13-30mm; W: 58-70%; HA: 57-63%

Type Locality: None.

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Vol. 11; No. 4

50H.

Figure 18
Morula uva (Ropinc)

Fiji Islands
one row of radular teeth - lateral shown in different position

Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Drupa uva, Morula papillosa and Ricinula
alba are all based on the same figure from Martini (1777,
pl. 101, fig. 970). Ropinc (1798) cited an additional
reference to Lister (1685-92, pl. 954, fig. 5), which
clearly represents the species Drupa uva, and has been
erroneously cited by Lamarck (1822) for his Murex
concatenatus.

Purpura nodulifera MENKE, 1829, which is the type
species of Oppomorus IrREDALE, 1937, is absolutely un-
identifiable. MENKE (1829) described his species as
white, conically-ovate, axially nodulosely costate, nodes
transversely striate; he did not cite a figure and his de-
scription is applicable to several species of Morula.

The types of Ricinula aspera LAMARCK are in the
Muséum d’Histoire Naturelle in Geneva. The holotype
no. 1107/17/1 — 20.5 mm, is the species Morula uva, as
is the 15.5 mm paratype. There is a third specimen in
the type-collection measuring 21.5 < 14.3 mm, which is
the species Drupa marginatra (BLAINVILLE). This speci-
men must have been added after the description, since
according to Rosalie de Lamarck’s annotation in her
father’s copy of the “Animaux sans vertébres,” Lamarck
possessed only 2 specimens. These two specimens are the
form with the spinose nodes. Lamarck’s holotype of
Ricinula nodus is preserved in the same Museum under
no. 1101/18/2 — 24.7 * 16.4 mm, and also is the spe-

cies Morula uva.

(Cronia) H. & A. Apams, 1853

Cronia H.« A. Apams, 1853, Gen. rec. Moll., 1: 128. Type
species by M Purpura amygdala KiENER, 1835

Discussion: In this subgenus are placed species of Morula
with prominent spiral and sometimes axial sculpture. The

Vol. 11; No. 4

radula of Cronia is said to differ from that of Morula in
the absence of side denticles on the rhachidians of Cro-
nia, While this is true as far as the 2 species of Cronia
from Fiji are concerned, rhachidians devoid of side den-
ticles do occasionally crop up in the Morula group (see
M. granulata Ductos and M. nodicostata Pease — fide
Maes-Orr, 1967).

Morula(Cronia) aurantiaca
(Homsron & Jaguinot, 1853)

(Plate 49, Figure 24)

1853. Purpura aurantiaca HomBron & JaQuiNnor, Voy. Pole.
Sud Ast. & Zélée, pl. 22, figs. 28, 29

1854. Purpura aurantiaca HoMBRON & JAQuINoT, Voy. Pole
Sud. Ast. « Zélée, text, p. 91

Shell: Only one juvenile specimen has so far been re-
corded from the Fiji Islands. The adult shell has strong
and broad axial ribs, close-set spiral striae which are oc-
casionally purplish-brown, about 6 labial denticles and a
few weak columellar plicae and some denticles.

L: ca. 25 mm

Type Locality: L’ile Hogoleu.

Habitat: The single specimen was found under a coral
rock near the reef’s edge, in 2 feet of water.
Distribution: West Viti Levu. — Pacific.

Discussion: A comparison of specimens of Morula amyg-
dala (KiENER) and M. aurantiaca (HoMBRON & JAQUI-
NOT) in the United States National Museum would in-
dicate two extremely similar species. Morula aurantiaca
from Pacific localities seems to be more brightly coloured
with a predominance of orange than are specimens of
M. amygdala from Australian localities.

Morula (Cronia) fiscella (GMELIN, 1791)

(Plate 49, Figure 25; Text figure 19)

1791. Murex fiscellum Gmeuin, Syst. Nat., ed. 13, p. 3552,
No. 160

1845. Murex fiscellum “Cuemnitz,’ Reeve, Conch. Icon.,
pl. 27, fig. 124

1853. Purpura stellaris HoMBRON & JAQUINOT, Voy. Pole Sud
Ast. & Zélée, Atlas, pl. 22, figs. 13, 14

1863. Coralliophila confragosa H.x A.Avams, Proc. Zool.
Soc. London, p. 432

1868. Ststrum triangulatum PEasE, Amer. Journ. Conch., 3:
278; pl. 23, fig. 15

1869. Murex fiscellum Ktster, Conch. Cab., ed. 2, p. 95;
pl. 33, figs. 10, 11

Shell: White or dirty grey, base of depressions in latticed
sculpture brown in fresh specimens. Sculptured with 7 - 8
broad and scabrous axial ribs, which consist of a number

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Page 311

of axially arranged fluted foliations; axial ribs overridden
by 3-4 spiral cords, which are either double or triple
stranded; towards the base are generally 2 rows of
oblique foliated but blunt spines. The axial and spiral
sculpture gives rise to recessed “windows” which are
brown in colour. Aperture light or dark violet, edge of
labial lip slightly fluted and sculptured with 6-7 small
regular denticles, columella with 2-4 small denticles
anteriorly. Operculum with lateral nucleus.

Young specimens have an angulate labial lip near the
juncture of the aperture; the apertural interior is whitish
and ornamented with 1 - 2 broad dark brown bands.

100
————————d

Figure 19
Morula (Cronia) fiscella (GMELIN)

Fiji Islands
half row of radular teeth

Radula: Odontophore 3.03 mm long and 0.19 mm wide
in an animal with a shell 24.5 mm in length; rows of
teeth number 129 (plus 7 nascentes). Rhachidians with
a long central cusp, small intermediate denticles and
simple side cusps; side denticles absent. Central cusp may
be shorter in some specimens than in others; intermediate
denticles are sometimes close-set to the side cusps. Laterals
simple and curved.

L: 18-35mm; W: 63-67%; HA: 68-73%

Type Locality: Insulam Pulo Condore, prope Sinam.
Habitat: On reefs, under coral rocks, in shallow water.
Moderately common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: Morula fiscella (GMEtIN) and M. margariti-
cola (BropErip, 1832) had a very confused nomencla-
tural history and both names have been applied to both
species interchangeably. GmMeELIn’s (1791) description is
equivocal on its own merits, and the interpretation of
the species depends on the cited figures from CHEMNITZ
(1788, pl. 160, figs. 1524, 1525) and on CHEmnirz’s
description. CHEMNITZ (op. cit.) clearly described his
species as strongly clathrate with deeply recessed “win-
dows” (depressions) which are coloured brown; his de-
scription and figure are only applicable to M. fiscella
and not to M. margariticola. The species has been cor-
rectly figured by ReEve (1845).

Page 312

Morula (Cronia) margariticola (BropERtP, 1832)
(Pate 49, Figure 26; Text figure 20)

1832. Murex margariticola BropEriP, Proc. Zool. Soc. Lon-
don, p. 177

1832. Purpura fiscella Lam., BiainviLLE, Nouv. Ann. Mus.
Hist. Nat., 1: 206; pl. 10, fig. 8 (non Murex fiscellum
Ge in, 1791)

1833. Purpura thiarella Quoy & Garmarp, Voy. Astrolabe,
Zool., 2: 571; pl. 39, figs. 4, 5, 6 .

1923. Morula rhyssa Dax, Proc. Acad. Nat. Sci. Phila., 75:
304 (nom. nov. pro Ricinula fiscellum REEVE, 1846)

Shell: Dark brown to almost black in colour, interstices
whitish. Sculptured with 9-12 broad axial ribs on the
body whorl and 11 - 14 ribs on the penultimate whorl;
axial ribs bisected by scabrous spiral cords which form
nodules at the point of intersection. Spiral ridges number
from 13-19 on the body whorl and from 5-9 on the
penultimate whorl; usually 3 - 4 ridges are more promi-
nent than the remainder. Aperture light violet or bluish-
white, labial lip with 5-6 small denticles, columella
with 1-3 elongated denticles anteriorly; parietal wall

100u
—E——————

Figure 20

Morula (Cronia) margariticola (BRoDERIP)

Fiji Islands
a. half row of radular teeth
b. ventro-lateral view of rhachidian

occasionally with a brown blotch.

Radula: Odontophore 6.0 mm long and 0.22 mm wide in

an animal with a shell 26.0 mm in length; rows of teeth

number 161 (plus 6 nascentes). Rhachidians with a long

central cusp, small intermediate denticles and simple side

cusps; side denticles absent. Laterals simple and curved.
The chitinous teeth are set and hinged onto a darker

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Vol. 11; No. 4

coloured base plate (dotted line in Text figure). Early
rows of teeth are so worn that only the elongated base-
plate remains.

L: 18- 37mm; W: 55-63%; HA: 57 - 62%

Type Locality: Oceano Pacifico, Lord Hood’s Island.
Habitat: Under basalt boulders, near the high tide level
and on reefs under coral rocks, in shallow water. Mod-
erately common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: There are 3 syntypes of Murex margariticola
in the British Museum (Natural History) in London.
Broperip (1832) gave the measurements of the holotype
as 13 x °/1” (= 34.9 X 16.8mm), but all 3 syntypes
are smaller than the given size. The largest specimen
measures 30.0 mm and is a fully mature specimen; this
specimen is obviously the holotype. A smaller (25.3 mm)
specimen is immature and has an “x” marked inside the
aperture.

Maculotriton Datu, 1904

Maculotriton Dati, 1904, Smiths. Misc. Coll., 47: 136. Type
species by OD Triton bracteatus Hinps, 1844

Discussion: Species of this group are ranellid in appear-
ance and may, or may not, have a varix on some of
the whorls. The radula (fide THIELE, 1929) is similar to
that of Cronia H. « A. ADAMs.

Maculotriton egregius (REEVE, 1844)

(Plate 49, Figure 27)
1844. Triton egregius REEVE, Conch. Icon., pl. 18, sp. 78

Shell: White to yellowish in colour, occasionally orna-
mented with faint brown spots. Sculptured with 2 promi-
nent varices on the body whorl and 3 coarse angulate
axial ribs between the varices; penultimate whorl has
8-9 axial ribs and varices become indistinct on earlier
whorls. Axial ribs are crossed by spiral cords, some of
which are more prominent than others. Aperture creamy-
white to light yellow in colour, labial lip with 7 - 8 elon-
gated denticles, columella either smooth or with 1 - 2 very
small denticles.

L: 20-30 mm; W: 55-60%; HA: 58 - 62%

Type Locality: Island of Masbate, Philippines.
Habitat: On reefs, under coral rocks, in shallow water.
Rare.

Distribution: North Viti Levu. — Philippines.
Discussion The spiral striae are not brown as described
by REEvE, and the Fiji species is only tentatively associ-
ated with REEveE’s Triton egregius. Triton eximius REEVE,

Vol. 11; No. 4

THE VELIGER

Page 313

1844, is another similar species, but the axial ribs are
much finer and more numerous.

Vexilla Swainson, 1840

Vexilla Swanson, 1840, Treat. Malac., pp. 69, 300; fig. 67.
Type species by M Vexilla picta Swainson, 1840
= Strombus vexillum GMELIN, 1791

Discussion: The radula of Vexilla (fide Cooke, 1919) is
similar to the radula of Drupella THIELE, except that the
laterals are less broad; it also resembles the radula of
Nassa ROpiNnc in some aspects.

Vexilla vexillum (GMELIN, 1791)

(Plate 49, Figure 28)

1791. Strombus vexillum GMELIN, Syst. Nat., ed. 13, p. 3520,
No. 52

1836. Purpura taeniata Powts, Proc. Zool. Soc. London, p. 96

1840. Vexilla picta Swainson, Treat. Malac., p. 300, fig. 67

Shell: This well-known species hardly requires a detailed
description. The shell is dark brown in colour, ornamented
with white transverse bands; the aperture is wide, creamy-
yellow in colour and the labial lip is denticulate.

L: 15- 22mm; W: 54-58%; HA: 78 - 84%

Type Locality: Oceano Indico.

Habitat: On reefs, under coral rocks, in shallow water.
Uncommon.

Distribution: South Viti Levu. - Indo-Pacific.

Nassa Ropinc, 1798

Nassa Ropinec, 1798, Mus. Bolten., p. 132. Type species by
SD (Dati, 1909) Nassa picta Ropinc, 1798 = Buc-
cinum sertum BruGuiERE, 1789

Nassa serta (BRuGUIERE, 1789)

(Plate 49, Figure 29; Text figure 21)

1789. Buccinum sertum BrucutkrRE, Encycl. Meéth., 1: 262

1798. Nassa picta Ropinc, Mus. Bolten., p. 132, No. 1655

1817. Stramonita hederacea SCHUMACHER, Ess. nouv. syst.,
p. 227

1967. Nassa serta (BRUGUIERE), CERNOHORSKY, Mar. shells
Pacif., pl. 60, fig. 443 (spawn)

Shell: Reddish-brown in colour, ornamented with broad
and irregular white or light yellow zones and bands. Sculp-
tured with from 25 - 35 obsolete axial ribs and 20 - 30
nodulose spiral cords; axial ribs generally more promi-
nent on earlier whorls. Aperture wide, creamy-yellow in
colour, edge of labial lip obsoletely denticulate; colu-

mella creamy-white, smooth, apart from an anterior den-
ticle. Operculum dark brown with a lateral nucleus.

100% 100u

AA) AX)

Figure 21

Nassa serta (BRUGUIERE)

Fiji Islands
a. half row of radular teeth
b. rhachidian of radula

Radula: Odontophore is 9.8 mm long and 0.26 mm wide
in an animal with a shell 41.0mm in length; rows of
teeth number 151 (plus 10 nascentes). Rhachidian with
a long or moderately short central cusp, with or without
intermediate denticles and short but massive side cusps;
laterals simple and curved.

L: 30-60mm; W: 49-56%; HA: 69 - 76%

Type Locality: None.

Habitat: On reefs, under coral rocks, in shallow water.
Common.

Distribution: Throughout the Fiji Islands. — Indo-Pacific.
Discussion: The egg capsules of Nassa serta are typically
muriciform and rather similar to those of Chicoreus car-
neolus (Ropinc 1798).

SUMMARY

A total of 28 species of Thaidinae have been recorded

from the Fiji Islands to date; all the species inhabit the

littoral zone. The species recorded from Fiji are distrib-
uted over the following genera:

Thats 3

Mancinella 2

Drupa 5

Drupina 1

Drupella 4

Morula 7

(Cronia) 3

Maculotriton 1

Vexilla 1

Nassa 1

8

Total 28 species

Page 314

THE VELIGER

Vol. 11; No. 4

LITERATURE CITED

ARAKAWA, KOHMAN Y.

1958. | On the remarkable sexual dimorphism of the radula of
Drupella. Venus, Japan. Journ. Malacol., 19 (3/4): 206
to 214; plt. 6; 2 text figs.; tables (February 1958)

1958a. Some notes on the radula of Purpura echinulata La-
MARCK. Venus, Japan. Journ. Malacol., 20: 69-75

(July 1958)

1962. _A study on the radulae of the Japanese Muricidae. (1)
The genera Purpura, Thais, and Mancinella. Venus, Jap-
an. Journ. Malacol., 22 (1): 70- 78; plts. 5, 6 (Aug. 1962)

1964. A study on the radulae of the Japanese Muricidae (2).
The genera Vexilla, Nassa, Rapana, Murex, Chicoreus, and
Homalocantha. Venus, Japan. Journ. Malacol., 22 (4):
355 - 364; pit. 21 (March 1964)

1965. A study on the radulae of the Japanese Muricidae (3).
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lina, Phrygiomurex, Cymia, and Tenguella gen. nov. Ve-
nus, Japan. Journ. Malacol., 24 (2): 113-126; plts. 13, 14

(October 1965)
BLAINVILLE, HENR1 Marie Ducrotay DE

1832. Disposition méthodique des espéces récentes et fossiles

des genres Pourpre, Ricinule, Licorne et Concholépas de M. de

Lamarck. Nouv. Ann. Mus. Hist. Nat., Paris. 1: 189 - 263;
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1778. Index rerum naturalium Musei Caesarei Vindobon-

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Broperip, WILLIAM JOHN
1832. no title. Proc. Zool. Soc. London, pt. 2: 173 - 179
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1780-1795. Neues systematisches Conchylien-Cabinet (conti-
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Bull. zool.

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DEMOND, JOAN

1957. | Micronesian reef-associated gastropods.

11 (3): 275 - 341; plts. 1-4; 42 text figs.
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1957. A historical review of the mollusks of Linnaeus. Part 5.
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Ductos, M.

1832. Description de quelques espéces de Pourpres, servant de
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DuNKER, WILHELM

1858-1870. Novitates conchoiogicae. Mollusca marina. Beschreib-
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GMELIN, JOHANN FREDERICH

1791. Systema naturae per regna tria naturae

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Pacif. Sci.
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Editio
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Page 316

THE VELIGER

Vol. 11; No. 4

Effects of Turbidity-Producing Substances in Sea Water

on Eggs and Larvae of Three Genera of Bivalve Mollusks

BY

HARRY C. DAVIS

AND

HERBERT HIDU!

Bureau of Commercial Fisheries, Biological Laboratory, Milford, Connecticut 06460

(7 Text figures)

INTRODUCTION

THE EFFECT OF SILT or other turbidity-producing sub-
stances on shellfish generally, and particularly on oysters
and their larvae, has long been of interest to biologists
and to shellfish producers. In addition to the obvious
damage to oyster beds silted over in harbor dredging and
in building bridges and roads in estuarine areas, it has
been suspected that turbidity from these sources affected
larval development and setting of oysters. Lunz (1938),
however, found no harmful effects from high turbidities
on adult oysters or on spawning and setting, in his field
study during the dredging of the Intra-Coastal Water-
way of South Carolina, except where the oysters were
actually buried by the spoil.

Only a few studies of the effects of turbidity have been
made under controlled conditions. LoosANoFF & Tom-
MERS (1948) who, in laboratory experiments, studied the
effects of a series of concentrations of turbidity-producing
materials on adult oysters found that 0.1 gm of silt per
liter of sea water reduced the pumping rate of adult
oysters by 57% and that concentrations of 3 and 4 gms
per liter reduced pumping by 94%. They reported gener-
ally similar results with kaolin and chalk, and that 0.5 gm
of Fuller’s earth per liter reduced pumping by 60%.

More recently, Davis (1960) determined quantitatively
the effects of a series of concentrations of several tur-
bidity-producing substances on the percentage of eggs
of the hard clam (Mercenaria (= Venus) mercenaria)
that developed into normal straight-hinge larvae and on

' Present address: University of Maryland, Natural Resources
Institute, Box 38, Solomons, Maryland 20688

the survival and growth of the larvae. In the work
reported here the same techniques were used to determine
the effects of several concentrations of silt, clay, and
Fuller’s earth on embryonic development of the American
oyster (Crassostrea virginica) and on survival and growth
of larvae of the American oyster and the European
oyster (Ostrea edulis).

In addition, a series of experiments was run to com-
pare the effects of different-sized particles of pure silicon
dioxide on embryos and larvae of hard clams and Ameri-
can oysters, in an attempt to determine the effect of
particle size of a suspended material, as opposed to pos-
sible effects of its chemical composition.

MATERIALS ann METHODS

The turbidity-producing materials used — silt, clay (ka-
olin N. E VII Mallinckrodt)?, and Fuller’s earth (dusting
powder, McKesson), — were the same as those used in
the experiments on clam larvae (Davis, 1960). The
silt was collected from the bed of Milford Harbor just
upstream from the laboratory. It was washed through a
325-mesh screen to remove larger particles and debris,
after which it was collected in a Buchner funnel and
washed with distilled water to remove the salts. It was
then dried at 200° C and the dried cake was ground in an
all-ceramic jar mill. The Fuller’s earth was similarly
ground.

Fisher’s flotated silica powder (about 240 mesh) was
used for the experiments to determine the effect of particle

2 Mention of trade names does not imply endorsement of com-
mercial products by the Bureau of Commercial Fisheries.

Vol. 11; No. 4

THE VELIGER

Page 317

size. It was suspended in distilled water and allowed to
settle in a tall column. The settled plug was pushed out
and cut into sections. The sections used were chosen to
give particle sizes within the ranges: <5y, 5 - 25, and
25 - 50m. Particle-size determinations, made by Dr. Keith
E. Chave, Director, Marine Science Center, Lehigh Uni-
versity, are given in Table 1.

Table 1

Percentages by weight of particles of different sizes
composing the materials tested *

Particle Fuller’s
Sizes Silt Kaolin Earth
> 62.5p 8.24 12.06 8.87
62.5 - 15.6u 28.36 6.03 18.16
15.6- 3.9u 39.10 10.72 26.10
3.9- 0.98 9.85 44.87 21.27
< 0.98y 14.54 26.32 25.60
Silicon Dioxide
< Sp 5 - 25p 25 - 50u
125 - 64y — trace 1.8
62.4 - 31.2u — 1.3 17.3
31.2 - 15.6 — 9.3 64.9
15.6- 7.84 trace 84.8 16.0
7.8- 3.9u 4.1 2.4 trace
3.9- 1.95u 56.9 2.3 —_
1.95 - 0.98 18.3 trace —
0.98 - 0.49u 17.2 — —
< 0.49n 3.5 — —

3 Particle size determinations made by Dr. Keith E. Chave, Director,
Marine Science Center, Lehigh University, Bethlehem, Pennsyl-
vania

In our larval cultures appreciable quantities of kaolin
and Fuller’s earth, particularly at low concentrations,
flocculated as particles 100u or more in size. Dr. Chave
reported similar flocculations in his particle size deter-
minations and believed, “the significant percentages of
material greater than 62.54 probably represent floccula-
tion of particles in the water” (personal communication).

Concentrations of all materials are expressed as grams
of dry powder per liter. In making up a suspension, a
weighed quantity of the powdered material was thorough-
ly mixed with sea water and the various experimental
concentrations were prepared by serial dilution.

The methods for obtaining fertilized oyster eggs out of
season have been described, as have methods for deter-
mining the percentage of eggs developing to the straight-
hinge larval stage, and for determining the survival and
rate of growth of larvae (LoosanorF & Davis, 1963).

In all experiments the sea water in the larval cultures
was changed every second day and larvae in all cultures
were fed a mixture of Isochrysis galbana and Mono-
chrysis lutheri daily. The quantity and quality of food
were the same for all cultures within an experiment since
each culture received an equal volume of algal food. The
quantity and quality of food in successive experiments,
however, were not necessarily equal, due to changes in
the algal cultures and sea water. Duplicate cultures were
used in each experiment at each concentration of sus-
pended material. Temperatures of all cultures were main-
tained between 23° C and 25°C and salinity between 26
ANC OAc:

European oyster larvae may start setting 8 days after
release under optimum conditions and are, thereafter, not
available for quantitative sampling. Consequently, samples
for determining percentage survival and rate of growth of
these larvae were taken on the 7" day after the larvae
were released, i.e., after 7 days at experimental condi-
tions. Few American oyster larvae set before the 14"
day even under near-optimum conditions. Samples of
American oyster larvae, therefore, were taken on the 14"
day after fertilization or after 12 days at the experimental
conditions. Some clam larvae may reach setting size and
undergo metamorphosis as early as the 8" day, but even
these early post-setting stages can be suspended and
accurate quantitative samples can be taken as late as the
12" day after fertilization, i.e., after 10 days at the
experimental conditions.

EFFECTS on EMBRYONIC DEVELOPMENT

Silt was more harmful to American oyster embryos
than either clay (kaolin) or Fuller’s earth. As little as
0.188¢/1 of silt had a significant effect on the percentage
of oyster eggs developing to the straight-hinge larval
stage, whereas Fuller’s earth had no significant effect
until concentrations exceeded 1g/1 and kaolin had no sig-
nificant effect until concentrations exceeded 2g/1 (Table
2). Moreover, in only one experiment did any (3%) of
the oyster eggs develop normally in 1g/1 of silt, whereas
the average number developing normally, for all experi-
ments in 4g/l1 was 26% for Fuller’s earth and 76% in
kaolin.

Davis (1960) found that a normal number of clam
eggs developed in silt concentrations as high as 0.75g/I,
but it required only 0.125g/1 of Fuller’s earth or 0.5g/1
of kaolin to cause a significant reduction in the number
of clam eggs developing normally. The experiments re-
ported here show that oyster embryos are less tolerant
of silt than are clam embryos, but oyster embryos are
considerably more tolerant of kaolin and Fuller’s earth

Page 318

than are clam embryos. Results with clam eggs and with
oyster eggs were similar, however, in that no embryos
of either species developed normally at the higher con-
centrations of silt, whereas a significant number of em-
bryos of both species did develop normally in even the
highest concentrations (4g/1) of kaolin and Fuller’s earth.

Table 2

Percentage of clam and American oyster eggs developing

to the straight-hinge larval stage in different concen-

trations of suspended materials. The number of eggs

developing to the straight-hinge stage in control cultures
was considered 100%

Concentration Silt Kaolin Fuller’s Earth
grams/liter Clam4 Oyster Clam* Oyster Clam*+ Oyster
0.0 (controls) 100 100 100 100 100 ~=©100
0.125 95 95 82 100 75 104
0.188 90 78 — — — —
0.250 96 73 82 100 61 103
0.375 93 66 — — — —
0.500 99 31 52 104 41 102
0.750 92 9 — _ — —
1.000 79 3 37 +108 57 98
1.500 65 0 — — — —
2.000 39 0 49 94 50 79
3.000 0 0 — — = —
4.000 0 0 42 76 45 26

Silicon Dioxide Particles

< Su 5 - 25u 25 - 50p

Clam Oyster Clam Oyster Clam Oyster
0.0 (controls) 100 ~=100 100 ~=100 100 ~=100
0.125 106 94 106 =105 114 = 112
0.250 105 109 96 104 98 94
0.500 105 93 98 106 102 100
1.000 OD) 107 945 103 98 111
2.000 85 =114 CP iY 96 109

4.000 69 = 123 103 96 92 ly

4 from Davis, 1960

The experiments with silt, kaolin, and Fuller’s earth
led us to conclude, tentatively, that the larger particles
present in silt and, to a lesser extent, in kaolin and Ful-
ler’s earth were primarily responsible for the deleterious
effects on oyster eggs, whereas the smaller particles, most
numerous in kaolin, were probably responsible for the
effect on clam eggs.

The percentage of oyster eggs developing normally did
not decrease significantly, however, in any of the various
concentrations of silicon dioxide tested, regardless of
particle size. Moreover, only at the highest concentrations

THE VELIGER

Vol. 11; No. 4

of the smallest particles of silicon dioxide did the per-
centage of clam eggs developing normally decrease sig-
nificantly. Obviously, then, the decreases in the percentage
of clam and oyster eggs developing in the higher concen-
trations of silt, kaolin, and Fuller’s earth, and the dif
ferences between their effects arc not wholly the result of
differences in particle size. Although silicon dioxide has
a higher density than the other substances tested and
fewer particles of any given size per unit of weight, it
seems unlikely that this difference alone can account for
the difference in results. Although 4¢/1 of silicon dioxide
gave no reduction in the percentage of oyster eggs
developing normally, only 1g/1 of silt reduced the number
by 97%. Even if the silicon dioxide were 4 times as
dense as the silt, there should be at least as many particles
in 4g of silicon dioxide as in 1g of silt.

EFFECTS on SURVIVAL or LARVAE

European oyster larvae were the most hardy of the 3
species tested in that survival did not decrease significant-
ly in concentrations up to 4g/1 of any of the suspended
material used, except kaolin (Figures la, 2a, 3a). This
result is in striking contrast to that with clam larvae,
which showed a fair survival in 4g/ of silt but no survival
in 1g/1 of either kaolin or Fuller’s earth (Davis, 1960).
American oyster larvae were less tolerant of silt than
either European oyster larvae or clam larvae, for as little
as 0./5g/1 of silt caused a significant decrease in the per-
centage of American oyster larvae surviving. American
oyster larvae, nevertheless, were more tolerant of kaolin
and Fuller’s earth than were clam larvae. Approximately
27 to 34% survived in 4g/1 of either kaolin or Fuller’s
earth.

We noted that many clam larvae, in the presence of
the smaller particles of kaolin and Fuller’s earth, eventu-
tually lost their ability to reject these particles. The lar-
vae then ingested the particles, the stomach became
packed, and the larvae died. We postulated that both
species of oyster larvae were better able to reject these
small particles and, hence, showed a lower mortality than
clam larvae in kaolin and Fuller’s earth.

The experiments with silicon dioxide indicated that it
is primarily the smaller particles that affect survival. Both
hard clam (Figure 4a) and American oyster (Figure 5a)
larvae suffered severe mortality at comparatively low
(0.5g/1) concentrations of the smallest (up to 5u) par-
ticles, whereas neither species suffered significant mortali-
ty at values as great as 4g/1 of the larger particles; even
at 4g/l of the smallest particles of silicon dioxide, how-
ever, mortality was not as great as with lesser concentra-
tions of some of the other materials.

Vol. 11; No. 4

THE VELIGER

Page 319

The effects of kaolin and Fuller’s earth, for example,
particularly at higher concentrations, on survival and
growth of clam larvae were more drastic than would be
expected from the effect of silicon particles of similar size.
Nevertheless, the much greater mortality of clam and
oyster larvae in the presence of the smallest particles of
silicon dioxide, as compared to mortality in the presence
of larger particles, did follow the same trend as with
kaolin and Fuller’s earth. The observation of numerous
oyster larvae with their guts packed with silicon particles
demonstrated that the American oyster larvae could not
reject these small silicon dioxide particles as successfully
as they did the particles of kaolin or Fuller’s earth. Euro-
pean oyster larvae, however, were apparently capable of
rejecting all suspended particles and, hence, suffered
little mortality.

EFFECTS on GROWTH or LARVAE

The effects of suspended material on growth of both
species of oyster larvae were somewhat different from
their effects on clam larvae. For example, silt had no
deleterious effect on growth of clam larvae until con-
centrations reached 1g/l, and growth was not reduced
drastically until silt concentrations were in excess of 2g/I.
Moreover, clam larvae showed evidence of feeding even
in 4g/1 of silt although growth was negligible (Davis,
1960). Larvae of American and European oysters reared
in 0.75g/1 of silt, however, suffered a significant reduction
in growth and those in 2g/] and over did not grow at all
(Figure 1b). At 3g/l and 4g/1 of silt all American oyster
larvae eventually died. Even though most European oyster
larvae survived at all silt concentrations, reduction in
growth of these larvae at concentrations of 1g/1 and less
was more drastic than for clam or American oyster larvae.

Conversely, both species of oyster larvae tolerated ka-
olin (Figure 2b) and Fuller’s earth (Figure 3b) better
than clam larvae did. Growth of clam larvae was drasti-
cally reduced by 0.5g/1 of either kaolin or Fuller’s earth
and all clam larvae were killed by 1g/1 of either material
(Davis, 1960). Growth of American oyster larvae, on
the other hand, was not significantly reduced by 1g/I1 of
kaolin. Moreover, in at least 3 of the 4 experiments these
oyster larvae showed appreciable growth in 2g/1 of kaolin
and growth of European oyster larvae was significantly
affected only in concentrations of 4g/l of kaolin (Figure
2b).

Fuller’s earth had a more deleterious effect than kaolin
on growth of both species of oyster larvae. Nevertheless,
both species showed significant growth and good survival
at 1g/l of Fuller’s earth, but American oyster larvae
showed only fair survival and no appreciable growth at

2g/l. European oyster larvae still showed good survival
and fair growth even at 2g/1 of Fuller’s earth (Figure 3b).

Growth of clam larvae was not seriously affected by
concentrations up to 2g/1 of either the 25-50 or the
5 - 25u particles of silicon dioxide (Figure 4b). Even in
the presence of the smallest particles (<5) growth of
the clam larvae that survived was fairly good. Results with
American oyster larvae (Figure 5b) were similar. The
rate of growth decreased at almost all concentrations of
all particle sizes of silicon dioxide, but the largest particles
interfered least with growth of these larvae. Even in the
presence of the smallest particles that interfered most,
with oyster larvae, as with clam larvae, those that sur-
vived showed appreciable growth.

VARIATIONS
AMONG SUCCESSIVE EXPERIMENTS
AND POSSIBLE CAUSES

Davis & CHANLEY (1956) pointed out that the first evi-
dence of toxicity to larvae, either of toxins produced by
living microorganisms or of synthetic chemical toxins,
was a reduction in the growth rate of larvae. It was sug-
gested by Davis (1960) that the more rapid growth of
clam larvae in the lower concentrations of silt, kaolin, and
Fuller’s earth was due, in part, to chelation or adsorption
of toxic substances present in the sea water or produced
by the algae and bacterial contaminants added as foods.

In the 3 experiments with silt, the effect of silt in con-
centrations of 0.5g/1 and lower was to increase the rate
of growth of the oyster larvae. The rate of growth of
the larvae differed somewhat in the 3 consecutive ex-
periments, but the increases in length, when converted
to percentages of the length increase in control cultures,
were in general agreement and results of the 3 experi-
ments were averaged to give the curve plotted in Figure
1b.

Four experiments were run with kaolin and Fuller’s
earth concurrently but subsequent to the experiments
with silt and, consequently, the food cultures and sea
water were different. Growth of larvae in the control
cultures was poor in Experiments 2 and 3, of the 4 con-
secutive experiments, and the shapes of the curves are
different from those of Experiments 1 and 4 in which
growth of larvae in control cultures was normal (Figures
6, 7). We believe the poor growth of larvae in the control
cultures was due to an excessive amount of toxins pro-
duced by microorganisms present in the sea water or,
possibly, by toxin-producing bacteria contaminating our
algal food cultures.

When the quantity of toxins appeared to be low (Ex-
periments 1, 4) (“normal” for our laboratory cultures)

Page 320

growth was optimum in about 0.5g/1 of kaolin, i. e., 0.5g/1
was sufficient to adsorb the toxins (Figure 6). When the
quantity of toxins appeared to be greater (Experiments
2, 3), about 1g/l of kaolin was required to adsorb the
toxins and give optimum growth, whereas at the lower
concentrations the harmful effects of suspended mate-
rials and toxins almost appeared to be additive (Figure 6).

The data from the experiments with Fuller’s earth gave
less clear-cut but similar results. When the concentration
of toxins was low (Experiments 1, 4) 0.25g to 0.5g/1 was
sufficient to adsorb the toxins, whereas 0.5g/1 or, perhaps,
slightly more was required when the concentration of
toxins was high (Experiments 2, 3, and Figure 7).

Of the materials used, only silt caused any appreciable
change in the pH of our sea water. Even with silt the
maximum change in pH was from 7.5 (normal for our
laboratory sea water) to 6.40. Other experiments have
shown that at pH 6.40 the decrease in the proportion of
clam or oyster eggs developing into normal larvae and in
the survival and growth of larvae of these two genera of
bivalves is less drastic than is caused by silt alone (CaLaB-
RESE & Davis, 1966). A portion of the effect of silt may
be attributable, nevertheless, to its effect on the pH of
sea water.

Other possible causes for the difference in effects of
silt, kaolin, Fuller’s earth, and silicon dioxide include
possible soluble toxic components or, more probably, dif-
ferences in their adsorptive and chelating characteristics.
Silt, a mixture of organic and inorganic materials, would
seem most likely to contain toxic components and highly
effective chelators capable of causing mortality of embry-
os or larvae by over-chelation.

Our experiments indicate that bivalve larvae can toler-
ate turbidities higher than those normally encountered in
natural waters, and that under certain circumstances low
concentrations of turbidity-producing materials may be
beneficial. Nevertheless, higher silt concentrations, such as
those produced by dredging or filling operations, could be
detrimental to bivalve larvae, both as a direct effect of the
particulate matter and, indirectly, as a result of lowered
pH. We also suspect that, in natural waters, disturbing
the bottom may release numerous bacteria, some of which
may be toxic, and organic enrichment sufficient to enable
these bacteria to reproduce rapidly. The effect of dredging,
therefore, may be more deleterious to bivalve larvae than
would be indicated by their tolerance to turbidity-pro-
ducing materials alone.

SUMMARY

1. As little as 0.188g/1 of silt caused a significant de-
crease in the percentage of oyster eggs developing nor-

THE VELIGER

Vol. 11; No. 4

mally, as did 3g/1 of kaolin or 4g/1 of Fuller’s earth.

2. The percentage of American oyster eggs developing
normally was not affected by concentrations of silicon
dioxide of 4g/1, regardless of particle size. Clam eggs
were affected only at 4g/1 of the smallest particles (<5).
3. Survival of European oyster larvae was less affected
by silt, kaolin, and Fuller’s earth than was survival of
larvae of either American oysters or hard clams.

4, The smallest particles (<(5y) of silicon dioxide had
the greatest effect on survival and growth of clam and
oyster larvae. Larger particles (5-25 and 25 - 50m)
had little effect on survival of either species or on growth
of clam larvae. Growth of American oyster larvae de-
creased progressively as the size of silicon dioxide par-
ticles was decreased.

3. Growth of European oyster larvae was less affected
by kaolin and Fuller’s earth than was growth of larvae of
the American oyster or hard clam, but was more strongly
affected by silt than was growth of larvae of either of
the other two species.

6. Bivalve larvae grew faster in low concentrations of
turbidity-producing substances than in clear sea water,
possibly because the suspended particles chelate or ad-
sorb toxins present in larval cultures. The “optimum”
concentration of suspended material probably depends
upon the amount of toxin to be chelated or adsorbed.

LITERATURE CITED

CALABRESE, ANTHONY & Harry Caru Davis
1966. The pH tolerance of embryos and larvae of Mercenaria
Biol. Bull. 131 (3) :

mercenaria and Crassostrea virginica.

427 - 436 (December 1966)
Davis, Harry Cari
1960. Effects of turbidity-producing materials in sea water on

eges and larvae of the clam (Venus (Mercenaria) mercenaria).
Biol. Bull. 118 (1): 48 - 54

Davis, Harry Cari & Paut Epwin CHANLEY
1956. Effects of some dissolved substances on bivalve larvae.
Proc. Nat. Shellfish. Assoc. 46: 59 - 74

LoosanorF, Victor Lyon « Harry Cart Davis
1963. Rearing of bivalve mollusks In: Advances in Marine
Biology, F S. RussEiu (ed.), Acad. Press, London 1: 1 - 136

LoosanorrF, Victor Lyon & Frances DorEetTa TOMMERS
1948. Effect of suspended silt and other substances on the rate
of feeding of oysters. Science 107: 69 - 70

Lunz, Ropert GEORGE
1938. Part I. Oyster culture with reference to dredging oper-
ations in South Carolina, 1 - 135. Part II. The effects of the flood-
ing of the Santee River in April 1936, on oysters in the Cape
Romain area of South Carolina. Rep. U.S. Engineer
Office, South Carolina, 1 - 33

*

Vol. 11; No. 4 THE VELIGER Page 321

‘ * Crassostrea virgini 160 —
PERCE ° ° Crassostrea virginica
(after 12 days exposure) °
: (after 12 days exposure) |
Ctoccooe ° Ostrea edulis 140; :
160+ fe q Me Cnecrooosed * Ostrea edulis
a ae) ays SHORE) \ (after 7 days exposure) |
140+ ier: Mee ee 1207 \ OssesaHe « Mercenaria mercenaria
oe 1960) : wae (Davis, 1960)
i-t) ae’ ~o-0\_
“gral (after 10 ae exposure) § 100" oe (after 10 days exposure) |
o use
1S) “a
g o Ss |
s os |
n
i a 3 : |
ec 3 :
o op
[=¥) . |
SS |
20} js : SAS poamaaa aera A
) OR eis ee MAR ako 4.0
eearsee, L pent, . grams per liter
) 0.5 #%110 1.5 20 2.5 3.0 4.0
grams per liter
Figure 1a Figure 1b
Percentage survival of three genera of bivalve larvae reared in Increase in mean length of three genera of bivalve larvae, grown in
sea water suspensions containing different concentrations of silt. different concentrations of suspended silt, plotted as percentages of
The number of larvae surviving in control cultures the increase in mean length of larvae in control cultures.
was considered 100%.
a [ = e Crassostrea virginica
165 ° ° Crassostrea virginica 160 \(after 12 days exposure)
(after 12 days exposure) Sa is *|Ostrea edulis |
yeni e Ostrea edulis 140} (after 7 days exposure) |
1407 (after 7 days exposure) Ore * Mercenaria mercenaria |
. . i}
e------ e Mercenaria mercenaria Pool (Davis, 1960) |
120} « (Davis, 1960) ° (after 10 days exposure) |
Fea ets eS (after 10 days exposure) © Ee Reo Casboso99n09K2200%% Soe |
Be PROP caerr “, 2 100k \
S 100F**, oe s ; . |
peony oe eS 5
2 \ Np = g \
2 \ eae A © 8or \ °
2 80r \ eye ee \
One Coe « \
[s) \ “0 fel e
a N 85, + 60f \
B fo: sy :
‘ c Bo \
40} \ | - 407 ‘ Ys
\ 20r i"
207 \ \ |
‘ 1. ® —t 4 —tL 1 g
. ® is ere fm fo) i 1.0 Te 2.0 2. Ae) 0
° 05 %11.0 1.5 20 25 3.0 4,0 0.5 5 of ie 3 4
grams per liter gramsip.
Figure 2a Figure 2b
Percentage survival of three genera of bivalve larvae reared in Increase in mean length of three genera of bivalve larvae, grown in
different concentrations of suspended kaolin. The number of larvae different concentrations of suspended kaolin, plotted as percentages

surviving in control cultures was considered 100%. of the increase in mean length of larvae in control cultures.

Page 322 THE VELIGER

Vol. 11; No. 4

° e Crassostrea virginica —
(after 12 days exposure) 140} . ° Crassostrea virginica i
140+ .
Sheen * Ostrea edulis (after 12 days exposure)
(after 7 days exposure) AeA Orreeeees ° Ostrea edulis
120} e-----=- * Mercenaria mercenaria (after 7 days exposure)
(Davis, 1960) a Qzeesnas ¢ Mercenaria mercenaria
ne _ ees 10 days exposure) 2, FENN cn (Davis, 1960)
ts tae 8 \ bce (after 10 days exposure)
encdctelo Vator (= 4 “095,
“4 S 80 | #9
oO ‘ ‘° “8,
3 8ot ~~ g 4
: © Gap |
OF 1
+ 6 L S 1
g 00 : \
o H
2 E,
o 407 \ .
407 \ yo 5)
: \ | 207 me
207 \ SS
‘ (ee ee a a aT. F —— ih Z|
: eee otis eds eae ya fo) 0.5 1.0 1.5 2.0 2.5 3.0 4.0
° 0.5 1.0 im a® Be 3.0 4.0 grams per liter
grams per liter
Figure 3a

Figure 3b
Percentage survival of three genera of bivalve larvae reared in

different concentrations of suspended Fuller’s earth. The number of

Increase in mean length of three genera of bivalve larvae, grown in
larvae surviving in control cultures was considered 100%.

different concentrations of suspended Fuller’s earth, plotted as per-
centages of the increase in mean length of larvae in control cultures.

ial ’ 120
3 7 . BO a |
fey ee ass eee ‘|
z eas [TOO fegx¢=~ g==— == gg — by
|i | g oy : = ===2 nnd
gg 80; : *-----*| & 5 micron particles = 80} Bie Et
Psi ‘ Onanesce *|5- 25 micron particles | 2 a at Peceo2acon3<9.
Sob 8 *——. 25 - 50 micron particles Saf a F
en ; 4 4 |
oO | ‘ | |
=
B, 40; | : 40}
Para | ag gies sear ° Casncbnios * <5 micron particles
eal : 2or »------» 5 - 25 micron particles
0 e 25 - 50 micron particles
° 0.5 1.0 15 2.0 ; 4.0 0 0.5 a 1.5 DD wate
grams per liter grams per liter
Figure 4a

Figure 4b
Percentage survival of clam larvae reared in different concentrations

Increase in mean length of clam larvae grown in different concen-
of suspended silicon dioxide particles of three different particle sizes. trations of suspended silicon dioxide of three different particle sizes.
The number of larvae surviving in control cultures The increase in mean length is plotted as a percentage of the

was considered 100%. —- increase in mean length of larvae in control cultures.

Vol. 11; No. 4 THE VELIGER Page 323
120 ee = 5 = =
100K YN, |
: ee Hea, Kg
3 80}: s 80+ aa DISS S Sea i ime
le ee
= ; 5 =.
@ Gor § e-....-e <5 micron particles es a)
5 siezale: * 5-25 micron particles 2 i
3 gol * * 25 - 50 micron particles S$ 40+ SUNOS 0s Se eee Paco ssoh cabaret A
io A e
nana Canora | e e------@ <5 micron particles
GYAL Sol GN ie ane a cPERREOcao paca Onboentne CosqoocHcoooBHeDeHo@oanen cod | 20- : :
| e------- * 5-25 micron particles |
. * 25-50 micron particles |
FS 05 a Tae eD 4.0 o Of LO mA 20 4.0

grams per liter

Figure 5a

Percentage survival of American oyster larvae reared in different

concentrations of suspended silicon dioxide particles of three differ-

ent sizes. The number of larvae surviving in control cultures was
considered 100%.

190 Leger ae zee eee
: * Experiment 1
Gop =e aoe eane Experiment 12)
-— —.— Experiment 3
170[° % Oneonaooens + Experiment 4

-

fop)

(o)
SI

ay

Y»

/

a ae

mean length in microns
oo
°
=

—— 2 ——
o 05 1.0 2.0 3.0 4.0
CONCENTRATIONS IN GRAMS PER LITER

Figure 6

Effect of different concentrations of kaolin on mean length of
American oyster larvae at 14 days in four separate experiments.
Points plotted are averages for duplicate cultures at each
concentration in each experiment.

grams per liter
Figure 5b

Increase in mean length of American oyster larvae grown in differ-

ent concentrations of suspended silicon dioxide of three different

particle sizes. The increase in mean length is plotted as a percentage
of the increase in mean length of larvae in control cultures.

180 ]

|
|
* Experiment 1 |
« Experiment 2 |
¢ Experiment 3

-* Experiment 4 |

\]\ -
\ Oh

mean length in microns
—s
es

1007,

.

80r Etat 4
|

foe i 1 L I
QO Of 1 2.0 3.0 4.0

CONCENTRATIONS IN GRAMS PER LITER

Figure 7

Effect of different concentrations of suspended Fuller's earth on

mean length of American oyster larvae at 14 days in four separate

experiments. Points plotted are averages for duplicate cultures at
each concentration in each experiment.

Page 324

THE VELIGER

Vol. 11; No. 4

A New Species of Murexsul from the Galapagos Islands

BY

WILLIAM K. EMERSON

AND

ANTHONY D’ATTILIO

Department of Living Invertebrates,

American Museum of Natural History

Seventy-ninth Street and Central Park West, New York, New York

10024

(Plate 50; 1 Text figure)

THROUGH THE KINDNESS of Mrs. Jacqueline DeRoy and
Mrs. Carmen Angermeyer of Academy Bay, Santa Cruz
Island, Galapagos Islands, we are privileged to describe
a new muricid species which we have placed in the genus
Murexsul on the basis of shell morphology and radular-
opercular characters.

Murexsul jacquelinae EMERSON & D’ATTILIO, spec. nov.
Plate 50; Text figure 1

The shell is small, pink, flesh-colored, with numerous, fine,
deeply cut, scabrous, spiral ridges; there are 5 post-nucle-
ar whorls preserved, the nepionic whorls being eroded.
The fusiform shell has a centrally located, medium sized
aperture; the whorls are angled, with an impressed su-
ture, and there are, on the penultimate whorl of the
spire, 8 spiral ridges that are of equal size with the inter-
spaces; these are crossed axially by 8 varices, at the most
elevated point of which the spiral ridges develop into
upturned open spines; both spiral and axial sculpturing
gradually increase in number on each succeeding whorl.
The body whorl has 6 varices crossed by 5 spiral ridges,
which are similar in character to those of the spire,
above the shoulder angle; these are followed by 6 ridges

below the shoulder; the canal has 4 additional ridges,
with some of intermediate and minor development. The
varices, on the elevated portion of the body whorl, have
spiral ridges that form more prominent spines than those
developed on the spire.

The aperture is ovate, vividly colored a rosy-violet; the
inner lip is elevated mostly posteriorly; the crenulated
labrum is sculptured within, reflecting the exterior ridges.
The siphonal canal is long, narrowly opened, and slightly
recurved; it is broad above where it possesses only growth
striae on its flattened surface; below it is narrowly tube-
like; on the left the canal bears the distal portion of
previous canals which are the projecting ends of the
varices of the body whorl.

The operculum is muricid, with a basal nucleus. The
radular characters (Text figure 1) suggest placement of
the new species in the genus Murexsul IREDALE, 1915.
Drawings of the radula of Murex octogonus Quoy &
Gamarp, 1833, the type species of Murexsul, were pre-
viously published by Hutton, (1883; plt. 13, fig. C) and
by Ponper (1968, plt. 1, fig. 1).

Measurements: Holotype, 20.4 mm in length, 11.1 mm
in width; largest paratype, 26.3 mm in length, 15.6 mm
in width.

Explanation of Plate 50

Murexsul jacquelinae EMERSON & D’ATTILIO0, spec. nov.

Figures 1, 2: Holotype (A. M. N.H. No. 147968), 100 m off Tagus
Cove, Isabella Island, Galapagos Islands; 3. Operculum in
aperture.

Figures 3, 4: Paratype (A.M.N.H.No. 147969), 50m off Bar-
rington Island, Galapagos Islands; 3. Note foreign growths
and drill hole.

Figures 5, 6: Paratype (coll. D’Attilio) 30m off Jervis Island,
Galapagos Islands; 3. Labrum broken.

[EMERSON & D’ATTILI0] Plate 50

THE VELIcER, Vol. 11, No. 4

Vol. 11; No. 4

THE VELIGER

Page 325

Type Locality: Off Tagus Cove, Isabella Island, Gala-
pagos Islands in 100 m, January 30, 1968, by the DeRoys,
holotype (Plate 50, Figures 1, 2) and 2 paratypes.

Figure 1

Murexsul jacquelinae Emerson « D’ArtTILIO, spec. nov.

Two lateral teeth and a central tooth from the radula of the
holotype; 300

Remarks: The new Galapagan species may be distin-
guished from its apparent congeners in the eastern Pa-
cific, Murexsul vittatus (Broperip, 1833) and Murexsul
lappa (Broperip, 1833), by their possession of heavier
and coarser sculptured shells that have fewer spiral ridges.
Additionally, the shells of MZ. vittatus may be essentially
white or may be colored with brown or blackish bands,
and the shells of Af. lappa are commonly white with 2
broad, ochre-colored bands. The aperture of the new
Galapagan species is distinctively colored a rosy-violet.

Type Repositories: Holotype, A. M.N. H. No. 147968
and 2 paratypes from the type locality (DeRoy Collection) ;
3 paratypes from off Barrington Island, Galapagos Is-
lands in 50 m, by the DeRoys, December 29, 1967 (1 - A.
M.N.H. No. 147969; 2 - D’Attilio Collection) ; 1 paratype
from off North Barrington Island, Galapagos Islands, in
55 - 64 m, by the Angermeyers, 1967 (ex Angermeyer Col-
lection) ; 1 paratype from off Jervis Island, Galapagos

Islands, in 30 m, by the DeRoys, March 22, 1967 (DeRoy
Collection ).

The largest paratypic specimen represents a more ma-
ture growth stage than that of the holotype (Plate 50,
Figures 1, 2) and a figured paratype (Plate 50, Figures
3, 4). This specimen, here illustrated, Plate 50, Figures
5, 6, which was collected dead, is bleached white, with
the labrum badly broken. It shows the mature character
of the elevated ridges with the long, spiny processes on the
varices.

ACKNOWLEDGMENTS

In addition to the Angermeyers and the DeRoys, we are
indebted to Professor Masao Azuma of Nishinomiya,
Japan, who kindly provided us with the original radular
drawing from which we have drafted the present drawing.
Our colleague, William E, Old, Jr., assisted us in various
ways.

LITERATURE CITED

Broperip, WILLIAM JOHN
1833. | Characters of new species of Mollusca and Conchifera
collected by Mr. Cuming.
1832 (prt. 2): 173-179
Hutton, FREDERICK WOLLASTON
1883. | Notes on some branchiate Gastropoda. Trans.
Proc. New Zealand Inst. 1882, 15: 118-131; plts. 13 - 16
(May 1883)

Proc. Zool. Soc. London for
(14 January 1833)

TrREDALE, Tom

1915. A commentary on Suter’s “Manual of New Zealand

Mollusca.” Trans. Proc. New Zealand Inst. 47: 417 - 497
Ponper, W. F.

1968. Nomenclatural notes on some New Zealand rachiglos-
san gastropods with descriptions of five new species. Rec.
Domin. Mus. 6 (4): 29-47; plts. 1-5

Quoy, JEAN RENE Constant, & JosEpH PAuL GaIMarRD

1833. | Voyage de découvertes de l’Astrolabe exécuté par ordre
du Roi, pendant les années 1826 - 1829, sous le commandement
de M. J. Dumont d’Urvile. Zoologie, Mollusca 2: 321 - 686

Page 326

THE VELIGER

Vol. 11; No. 4

Predators on Olivella biplicata,

Including a Species-Specific Predator Avoidance Response

BY

D. CRAIG EDWARDS !

Department of Zoology, University of Chicago,

and Department of Oceanography, Scripps Institution of Oceanography, La Jolla, California

(Plate 51; 1 Text figure)

IN THE COURSE OF MORE GENERAL STUDIES of Olivella
biplicata (SowerBy, 1825) (Epwarps, 1965, 1968) it
has been possible to assemble for the first time a list of its
predators with some observations on their feeding. The
predators include Octopus sp., Conus californicus Hinps,
1844, Polinices recluzianus (DESHAYES, 1839), Pisaster
brevispinus (Stimpson, 1857), Astropecten armatus Gray,
1840, one or more species of crabs, shore birds, and man.
Olivella was also found to exhibit a specific, vigorous es-
cape response to Pisaster brevispinus, but not to P ochra-
ceus (BRANpbT, 1835) or Astropecten armatus. The new
information on predators extends food lists and will aid
community energetics studies; it should also facilitate fur-
ther studies of the ecology of O. biplicata and could form
the basis of a thorough investigation of the effect of
predation on this long-lived (see below), abundant, inter-
tidal gastropod. The demonstration of a species-specific
avoidance response to a predator adds to the growing
literature on this ecologically important form of chemi-
cal communication.

Field work was done from Newport, Oregon, to Ensen-
ada, Baja California. Most observations were made at
low tide when the animals were exposed or shallowly
immersed; snorkeling was also used at most sites. Labora-
tory studies were done at the Oregon Institute of Marine
Biology, Charleston, Oregon, and at Scripps Institution of
Oceanography, La Jolla, California.

PREDATORS

The most diverse and possibly the chief predators on adult
Olwella biplicata are other mollusks. Numerous laboratory
and field observations by R. T: Paine (personal communi-
cation), N. Fotheringham (personal communication) , and
myself show O. biplicata of all sizes (8.5 to 26.1 mm)

' Present address: Department of Zoology, University of Massa-
chusetts, Amherst, Massachusetts 01002

are eaten by small Octopus bimaculoides PickForD &
McConnaucHeEYy, 1949, or O. bimaculatus VERRILL, 1883,
or both, in southern California. (Olivella size data herein-
after are shell lengths from siphonal canal to apex.) The
octopuses emerge from beneath rocks on sand flats, seize
Olivella, drill a very small hole (ca. 0.5mm maximum
external diameter) near the shell apex (edge of body
whorl or on whorl above; Plate 51, Figure 1), and appar-
ently inject a paralyzing venom (see Pirson & TayLor,
1961). The effect of Octopus predation on natural Oli-
vella populations, however, appears small: very few emp-
ty Olivella shells bear Octopus bore holes (see below),
and the proportion remains small even when shells are
collected near a regularly occupied Octopus burrow.

Several gastropods eat Olivella. In the San Diego,
California, area Conus californicus eats O. biplicata both
in aquaria and in the field (SAUNDERS & WoLFson, 1961);
and, of 56 known observations of prey taken by this pred-
ator in nature, more (10) were of O. biplicata than any
other species (KoHN, 1966). Conus swallows its prey
whole and may prefer small Olivella: two large C. cali-
fornicus (> 25mm) kept in an aquarium provided with
running sea water, a sand substrate, and 20 medium to
large O. biplicata (20- 28mm) for three weeks ate no
Olivella.

Another carnivorous prosobranch, Polinices recluzianus,
widely co-occurs with Olivella biplicata in California and
Baja California, and a small percentage of empty O. bi-
plicata shells from the field regularly bears the character-
istic countersunk bore hole of this predator (Plate 51,
Figure 2). Polinices, which is rarely abundant, feeds
primarily on bivalves; it also accepts gastropods as alter-
native food, but probably takes few olives in nature (see
below). The maximum rate of predation on O. biplicata,
were no other prey available, was determined, using
three Polinices of similar size (means and standard errors:
33.43 + 1.05 mm shell height & 29.50 + 1.15 mm shell
width; 1531 + 234 mgm shell-free dry weight). The
predators were first acclimated to eating Olivella in cap-

Vol. 11; No. 4

tivity and then placed in an aquarium provided with
running sea water, a sand substrate, and ca. 20 prey
ranging in size from 18.3 to 28.4mm. The three Poli-
nices (4593 mgm total shell-free dry weight) ate 13 O.
biplicata (4198 mgm total shell-free dry weight) in 15
days (3-18 November 1965); thus, the Polinices con-
sumed 6% of their own dry weight per day. This feeding
rate, while exceeding the 1.2 to 4.6% rate of some species
of Conus in the tropics (Koun, 1959, and in press), falls
far short of the 15 - 25% of body weight per day rates
given for other predatory gastropods, including Polinices
duplicatus (Say, 1822) and other Conus (THorson,
1958; Koun, in press). Possibly the non-preferred prey
status of Olivella and the season of observations — many
predaceous gastropods feed little, if at all, in winter — or
both affected the feeding rate of P recluzianus.

The feeding behavior of Polinices recluzianus is like
that described for other naticids (Gonor, 1965; FRETTER
& GRAHAM, 1962). Prey are seized with the propodium,
drawn under the predator, and wrapped up in the con-
cave ventral surface of the mesopodium. Under attack,
Olivella moves with strong forward lurches and often for-
ces its way under or around the enclosing mesopodium; es-
capees then continue their lurching progression for 8 - 10
cm. Polinices recluzianus may carry captured prey around
in the mesopodium before burrowing into the sand and
drilling them. Through apparently stereotyped behavior
in the handling of prey, P recluzianus always bores O.
biplicata on the body whorl near the posterior end of the
aperture (Plate 51, Figure 2). Boring the shell seems an
extremely inefficient way to attack Olivella, whose oper-
culum is a thin corneous scale. In fact, of 21 O. biplicata
eaten in aquarium trials, only 4 were completely bored,
14 were incompletely bored, and 3 were unbored. Incom-
pletely bored, empty Olivella shells are also taken in the
field. Although Polinices may occasionally force its prey’s
operculum, the incomplete bore holes suggest another ex-
planation, viz., that O. biplicata suffocates while wrapped
in the predator’s foot and relaxes. Since Olivella suffers no
harm from being deeply withdrawn into its shell for 10
hours or more in response to osmotic stress (EDWARDS,
1965), this answer requires the snails be deprived of
oxygen for lengthy periods. Moribund, but unbored, O.
biplicata can be recovered from P. recluzianus. Since par-
tially drilled, live snails also occur, escape is still possible
after drilling begins. Probably the length of time the prey
is enveloped in the predator’s foot determines the mode
of death.

Although Olivella biplicata often frees itself from Poll-
nices grasp, both the largest (28.4mm) and smallest
(17.0 mm) individuals available in feeding trials were
eaten by the medium-sized P recluzianus: neither was
there any statistically significant difference between the

THE VELIGER

Page 327

sizes of Olivella taken and those surviving, though sample
sizes were inadequate (13 vs. 9 respectively). Other field
and laboratory observations by N. Fotheringham (person-
al communication) and myself show olives of all sizes
(10.4 - 28.4 mm) fall prey to P recluzianus. Occasionally
O. biplicata exhibits strong escape responses upon coming
in contact with Polinices (see below).

Other gastropods may prey on Olivella. Larger olivids,
though absent from the area under study, feed on Olivella
species in Latin America (Otsson, 1956). In the labora-
tory small O. biplicata are drilled (Plate 51, Figure 3)
and eaten by Acanthina spirata (BLAINVILLE, 1832)
(FoTHERINGHAM, 1966). These two species do not nor-
mally occur together in the field, A. spirata preferring
rocky shores; but at more exposed sites Olivella are most
abundant on sand near rocks and pilings where predators
from solid substrates could attack them. Jaton festivus
(Hinps, 1844), Nassarius fossatus (Goutp, 1849), and
the opisthobranch Navanax inermis (Cooprr, 1862) co-
occur with O. biplicata and may eat it. No Jaton bore
holes have been found in Olivella shells, however, and
Nassarwus is mainly a scavenger. Navanax proved unres-
ponsive to Olivella in the field (Edwards) and in the
laboratory (Paine, 1963), and O. biplicata is not found
in Navanax fecal pellets (PatneE, op. cit.).

FoTHERINGHAM (1966) has noted a distinctive bore
hole on the columella of Olivella (Plate 51, Figure 4)
and other gastropod shells. These shells were occupied by
hermit crabs and were collected from rocky intertidal
sites in the San Diego area. I have not found this marking
on the shells of living Olivella or on hermit crab shells
collected from subtidal sand bottom. The bore hole re-
sembles those of gastropods, but may be made by a
barnacle (Tomuinson, 1953) or a flatworm (WoELKE,
1961). The much higher incidence of the bore holes in
worn, encrusted “old” shells than in clean, “new” shells,
though, suggests a non-predatory agent (Fotheringham,
personal communication).

Asteroids commonly prey upon gastropods, and at least
two eat Olivella biplicata. Pisaster brevispinus was occa-
sionally observed taking these snails on the northern Cali-
fornia and Oregon coasts. At Duxbury Reef (37°54’N
Latitude), the only site where the two species co-occurred
abundantly at low tide, ca. 40 seastars were examined;
only one was feeding, taking an Olivella (22.7 mm)
whose shell lip was damaged, possibly by a crab. These
two species also live together near extreme low tide at
Monterey Harbor and subtidally off North Spit at Coos
Bay. Olivella biplicata shows a species-specific avoidance
response to this seastar (see below).

The second asteroid predator on Olivella biplicata,
Astropecten armatus, is more specialized for sand bottom
life. The two species are commonly taken together in

Page 328

southern California and Baja California, usually subtidally
(Scripps Pier) but occasionally intertidally (Estero de
Punta Banda, below Ensenada, Baja California). The
feeding sequence was observed in an aquarium provided
with fresh sand and running seawater. The seastar, ap-
parently using distance chemoreception, quickly “runs”
to and over an O. biplicata placed nearby and begins
forcing its arms into the sand around the prey. While
burying its arms, the seastar may shift laterally in the
direction the snail was moving, or it may appear to
carry or push the prey for a distance. Astropecten does
not extrude its stomach but swallows prey whole. Even
the largest O. biplicata do not gain safety through their
size or strength: a 29.0 mm snail was readily taken and
completely ingested in just over 2 minutes. During in-
gestion the predator’s central disc may swell aborally
and prey Olivella shells can be felt inside. Intraoral di-
gestion may be an adjustment to life in sand or to the loss
of suckers on the podia, or both; in any case, external
examinations in the field will fail to reveal the foods
of Astropecten. Although the shell of the 29.0mm Ol-
vella was still inside the seastar 23 hours after ingestion,
in several aquarium trials an initially unfed Astropecten
regularly ate > 1 adult snail per day over 4 - 5 days; and
MacGinitre & MacGinitic (1949) report 3 or 4 snails
(species unspecified) may be found at one time in an
Astropecten. Clearly Astropecten armatus, to which O.
biplicata shows no definite escape response (see below),
could be an important predator upon O. biplicata.
Various crabs eat snails, either by cracking them open
or by pulling them from their shells (see SHoup, 1968, for
references). Small Cancer magister DANA, 1852, which co-
occurred with Olivella biplicata on several beaches in
Oregon, ate live, though possibly moribund, O. biplicata
in the laboratory. The crab held the snail with the legs
of one side of its body, while reaching deeply into the
shell aperture with the opposite cheliped. Considerable in-
direct field evidence — shells broken into many pieces
(sometimes with all the fragments lying within a few
square centimeters), empty shells with the body whorl
alone broken away (Plate 51, Figure 5), occupied and
empty shells with the lip broken back, and rare moribund
snails with both the shell lip broken away and the foot
torn raggedly -— indicates crab predation on O. biplica-
ta. Suspects include adult Portunus xantusu (Stimpson,
1862), Cancer antennarius Stimpson, 1856, and Cancer
magister, all of which have been taken with O. biplicata.
Exsuine et al. (1964) found Carcinus maenas LINNAEUS,
1758, Portunus puber (Linnaeus, 1767) and Cancer pa-
gurus LINNAEUS, 1758, when kept in cages in the field
with Nucella lapillus (LinNAEus, 1758) (similar size
range as Olivella), smashed and ate the snails as well as

THE VELIGER

Vol. 11; No. 4

removed others from their shells. Further, Portunus puber
observed in the laboratory, used its master chelae to
break away the body whorl of Nucella, producing shell
damage like that observed in Olivella shells (Plate 51,
Figure 5). Similarly, Kon (1959) found xanthid crabs
attacked Conus in the laboratory, breaking the outer lip
of the snail’s shells. These crabs never killed snails, though,
probably because the latter could retreat into older,
thicker portions of the shell. Clearly, then, crabs can and
do produce the type of shell damage observed in Olivella
shells.

Among vertebrates, gulls and other shore birds regular-
ly forage in the sand at low tide, taking snails and other
forms. Olivella is eaten by shore birds in northern Peru
(Korpcke & Korprcxe, 1952), and Stohler (personal com-
munication) and I have observed gulls taking O. biplicata
in California and Oregon, respectively. The gulls, probing
irregularities in the sand, sometimes throw the snails out
on the surface, and then overlook their find. All along
the Pacific coast small Olivella shells can occasionally be
found in the droppings of shore birds’. With the rising
tide, on the other hand, fish, which are major predators
on snails in some areas, move in large numbers over
Olwvella-rich sand flats and may well prey upon the snails,
but this has not been observed. Purple dye substances
irritating to fish are known for some olive shells (AsgorrT,
1954).

Probably the most important depredator on Olwella
biplicata in accessible localities is man. Humans do not
eat olive snails on our Pacific Coast, though they do in
Brazil, where one species is locally called “vitela” (veal)
or “vaquinha”’ (little cow) (Marcus & Marcus, 1959),
and in Peru (Oxsson, 1956). Olwwella biplicata shells
were extensively traded by the American Indians, who
used them as jewelry and money (STEARNS, 1889; HEN-
DERSON, 1930; Batty, 1935; Girrorp & Girrorp, 1942;
and Appott, 1954). Today the animals are collected in
great numbers simply for the beauty of their shells. Shell
dealers take them for sale at tourist stands across the
country; some shells are made into earrings. A few West
Coast residents also use Olivella to “decorate” furniture
and to construct “artistic” shell mosaics. The protective
shell has caused heavy losses for O. biplicata.

Thus, at low tide, Olivella biplicata are attacked by
shore birds, especially gulls, and man; and, as the tide
rises, they are preyed upon by seastars, octopuses, at
least two species of gastropods, and probably by crabs and
fish as well.

2 REEDER (1951) also found Olivella in the stomach of a Marbled
Godwit, Limosa fedoa, in southern California.

Vol. 11; No. 4

SPECIES-SPECIFIC
PREDATOR AVOIDANCE RESPONSE

Various marine gastropods show specific escape reactions
to predatory seastars and gastropods. The literature has
been reviewed regularly (BuLLocK, 1953; Passano, 1957;
Koun, 1961; Feper, 1963, 1967; Marcouin, 1964a; FE-
DER & CHRISTENSEN, 1966). Further work is available
on herbivorous gastropods responding at a distance to
camivorous ones (see KoHN & WATERS, 1966, and refer-
ences therein). The escape responses generally consist of
rapid movement (“running,” “galloping”) away from
the predator, sometimes with the shell being violently
swung about, and occasionally accomplished by means of
curious “leaping”; some naticids, Haliotis rufescens
SwaInson, 1822, and Diodora aspera (ESCHSCHOLTZ,
1833) extend a fold of the foot or mantle over their
shells, preventing seastar tube foot adhesion (Marco.in,
1964a; Montcomery, 1967). The stimulus is chemical
and apparently emanates from the external epithelium
of the predator, especially from the epidermis of tube
feet (FEDER & Lasker, 1964). Contact with a single tube
foot elicits vigorous escape reactions. Effective stimulation
also occurs through distance chemoreception of whole
animals and extracts.

Tests of escape responses in Olivella biplicata consisted
of touching the snail’s foot (propodium and parapodia)
with isolated and intact tube feet from the seastars Pis-
aster ochraceus, P. brevispinus, and Astropecten armatus,
as well as observing outcomes of contacts between whole
animals. Controls consisted of similarly touching snails
with a blunt probe or a glass rod. Olivella biplicata ex-
hibited a vigorous avoidance response to contacts with P
brevispinus tube feet, but no greater reaction to P ochra-
ceus and A. armatus than to control treatments. The
snails failed to give strong responses to contacts with the
aboral surfaces of any of the three seastar species.

The minimum escape reaction of Olivella biplicata
consists of a sharp turning away from the point of contact
with Pisaster brevispinus or one of its tube feet, followed
by top-speed crawling. More often upon contact O. bipli-
cata rears up on the hind portion of its foot, withdrawing
the propodium and throwing the parapodia forward; this
response frequently flips the snails over backwards in a
reverse half-somersault. In the extreme form of the
response the animal, after throwing itself on its back,
pumps the expanded metapodium up-and-down violently,
lifting the snail from the substrate and carrying it away
some 5 - 10 cm in a form of upside-down swimming. This
metapodial swimming response is qualitatively distinct
from any previously reported gastropod escape behavior.
It is effected by holding the parapodia close to the sides

THE VELIGER

Page 329

of the shell, especially at the anterior end, so the vigorous
down beats of the large, horizontally extended metapod-
ium force water down and back, lifting the gastropod and
propelling it forward. After each of the more energetic
responses the snails quickly right themselves and rapidly
crawl a short way (ca. 8 - 12 cm) before resuming a nor-
mal pace. Escape movements, while carrying the animals
away from the point of first contact with a whole seastar,
may carry them up against another ray of the predator;
in this case the avoidance reaction is repeated.

Initially Olivella biplicata gave no noticeable response
to Polinices recluzianus in feeding trials, even crawling
over the predator’s propodia without reaction. Later in
the trials, however, Olivella occasionally gave violent
avoidance responses to contacts with Polinices, including
the “swimming” response. Whether experience with the
predator accounts for the appearance of the sporadic
escape behavior is not known. Olivella also forces its way
from the grasp of Polinices as described above.

Pisaster ochraceus is abundant on rocks and pilings pro-
jecting from beaches, but only occasionally ventures onto
the sand; whereas, P brevispinus, the species Olivella bi-
plicata avoids, is a natural neighbor of and predator on
Olivella (see above). Thus, the escape reaction is only
elicited by the species of Pisaster that O. biplicata is
likely to meet in the field, and the response appears to
have direct survival value in carrying the snail away from
a known predator. Astropecten armatus, on the other
hand, lives with and eats O. biplicata, but elicits no defi-
nite avoidance response from it. Qualitative observations
indicate Astropecten moves so quickly over sand that
escape efforts by O. biplicata, including the “swimming”
response with its slow start, would be futile and could not
be selected for. Several Astropecten species can crawl
10mm/sec (FEDER & CHRISTENSEN, 1966), which ex-
ceeds all but the fastest escape responses of gastropods
(KoHn & Waters, 1966).

Animals exhibiting avoidance responses show varying
abilities to discriminate between possible predators. Two
highly specific responses are pertinent to the case of Oli-
vella. Appropriately for its rocky intertidal habitat and
vulnerabilities, Acmaea pelta EScHSCHOLTZ, 1833, shows
a “running” escape response to Pisaster ochraceus, but not
to PR brevispinus (Marcotin, 1964b) -— the opposite
reactions from those of O. biplicata. In contrast, Dendr-
aster excentricus ESCHSCHOLTZ, 1831, responds like Oli-
vella, avoiding the predatory P brevispinus by burrowing
into the sand, while ignoring Astropecten armatus (Mac-
Ginrrie & MacGinitte, 1949), even though the latter
commonly feeds on sand dollars (E. W. Fager, personal
communication). These differing responses could be used
as bioassays in isolating the specific chemicals eliciting

Page 330

THE VELIGER

Vol. 11; No. 4

avoidance reactions. Whether animals of the various taxa
respond to the same or different chemicals is a question
of particular interest; representatives of different phyla
do respond to certain seastar extracts (FEDER & ARVIDSSON,
1967), and the substances in extracts of different seastars
show chemical similarities (Mack et al., 1968).

Gonor (1966), from a consideration of the escape
responses of various prosobranchs, suggests these behavior
patterns were not evolved de novo, but developed from
pre-existing locomotory and righting mechanisms. In the
case of Olivella biplicata the minimum avoidance res-
ponse of rapid crawling does, but the more vigorous
responses do not, appear to agree with this view. The
“rearing” and “swimming” escape behavior has, however,
other uses than predator avoidance: it is seen when,
under conditions of crowding, one olive is bitten by an-
other, or when a snail is placed in an alien environment,
e. g., a plastic bag. The truly striking thing about Olivella
is the dexterity and versatility with which it uses its
foot — in locomotion, burrowing, handling of live prey,
and in reproductive behavior. Perhaps a more intriguing,
and resolvable, aspect of the development of avoidance
responses concerns not the movements themselves but the
acquisition of, and sensory mechanisms used in, the dis-
crimination of substances from predators.

Various investigators have seen gastropod avoidance
behavior succeed in permitting at least some individuals
to escape predators (see FEDER, 1967; Marco in, 1964a).
Feper (1963) found that snails exhibiting escape respon-
ses to Pisaster ochraceus were not fed upon in proportion
to their abundance and availability, whereas mussels and
barnacles were; the non-responding Acmaea scabra
(GouLtp, 1846), on the other hand, ranks high on the
seastar’s food list even though few live within the latter’s
range. The apparent rarity of captures of Olivella bipli-
cata by Pisaster brevispinus, despite the abundance of
both, was noted earlier. Defensive escape reactions, even
if effective in only a limited percentage of predator con-
tacts, would be of selective advantage to gastropods. In-
deed, that these behavior patterns have evolved demon-
strates both their adaptive value and, in turn, the
effectiveness of asteroids as predators.

POPULATIONAL EFFECTS or PREDATION

The significance of predation for Olivella biplicata
populations can only be speculatively evaluated since
available information is fragmentary. Probably larvac
and newly metamorphosed young, vulnerable to a wider
and different array of predators than adults and to
fluctuations in physical factors as well, suffer heavy losses,
though the evidence is circumstantial. Size frequency
distributions obtained through quantitative sampling of
natural populations at Coos Bay, Oregon, Yaquina Bay,
Oregon, and Duxbury Reef, California (Edwards, unpubl.)
show that, while new young are occasionally abundant,
proportionally few reach reproductive size (over 16 mm).
Olivella populations are often primarily composed of
larger, slow-growing animals, which may live 10 years or
more (P. W. Frank and D. C. Edwards, unpubl.; Storer,
1962, and personal communication). Yet, despite a po-
tentially lengthy reproductive life during which increasing
numbers of progeny are likely produced each year, even
a medium-sized Olivella female (20.6mm) produced
4 236 eggs in a single spawning period (Epwarps, 1968).
Also the species reproduces year-round (EDWARDS, op. cit.).
Thus many young are spawned for even the low, though
continuous, population turnover indicated. Numerous sus-
pension feeders and small predators could, of course, take
heavy tolls on the bottom-swimming larvae (Epwarps,
op.cit.) and newly settled young. In some species the
escape responses of smaller animals to seastars are less
successful than those of larger individuals (FEDER, 1967).
Gulls and Conus also take small, immature Olivella (see
above).

Although a variety of predators has been shown to
attack adult Olivella biplicata, several factors mitigate
their populational effects. First, the snails may evade
predators. A species-specific avoidance response to Pisas-
ter brevispinus was detailed above, and escape reactions
to Polinices recluzianus were noted. In addition, larger
Olivella, which live higher on the shore, tend to remain
buried during bright light conditions (EpDwarps, 1965) ;
and this behavior may aid in the avoidance of visual pred-

Explanation of Plate 51

Figure 1: Olivella biplicata shell (11mm) bored by Octopus sp.
(photograph by Ron Lam)

Figure 2: Olivella biplicata shell (22mm) bored by Polinices re-
cluzianus (photograph by Ron Lam)

Figure 3: Olivella biplicata shell (8mm) bored by Acanthina
spirata (photograph by Ron Lam)

Figure 4: Olivella biplicata shell (15mm) with bore hole of un-
known origin (photograph by Ron Lam)

Figure 5: Olivella biplicata shells (ca. 25mm) with the body

whorl broken back, possibly by crabs

John W. Evans)

(photograph by

THE VELIGER, Vol. 11, No. 4 [Epwarps] Plate 51

i

Figure 4

Vol. 11; No. 4

THE VELIGER

Page 331

ators. Second, not all the predators attack O. biplicata
in any given locality, e. g., Conus californicus and Astro-
pecten armatus occur only in southern California, whereas
Cancer magister and Pisaster brevispinus are more north-
erly species. Further, in the temperate latitudes under
consideration predator food choices may vary seasonally
and feeding may cease altogether in winter. But likely
the main factor reducing predation on adults is their
size: they have simply grown too large to be taken by
small carnivores, e. g., various worms. Indeed the larger
snails themselves prey on polychaetes (Edwards, unpubl.) .
Nonetheless, Polinices, Astropecten, and Octopus eat even
the largest Olivella.

Since empty Olivella shells of different sizes probably
do not disintegrate at very different rates, an unbiased
collection of such shells would provide information on
both size specific mortality rates and predation rates for
predators that leave distinctive marks. Empty shells are
always rare on Olivella beaches, so hermit crabs, Holo-
pagurus pilosus Homes, 1900, were employed as collect-
ors. (A difficulty here is that bored shells, which should
afford less protection, may be selected against by the
crabs.) E. W. Fager kindly provided me with an arbitrary
sample of 130 O. biplicata shells from a large hermit
‘crab aggregation in 18 feet of water at Scripps Pier. The
size-frequency distribution of the shells (Text figure 1)
shows a sharp decline in the abundance of larger ones:
only 12 shells (9%) were of adult size (>16mm).
Although the size structure of the La Jolla population is
not known, the shell data suggest relatively high mortali-
ty among immature snails, but very low mortality among
adults. Only 10 (7.7%) of the 130 Olivella shells gave

30

| fy |

fo} — LL
20 22

6 8 10 I 14 16 18
Shell Length (mm)

ix)
fo)

Number of Shells
3 a

uo

Figure 1

Size-frequency distribution of 130 Olivella biplicata shells from a
Holopagurus pilosus aggregation at Scripps Pier

evidences of particular predators: 4 had Polinices bore
holes, 4 had broken shell lips indicative of crab attacks,
and 2 bore possible Octopus drill holes. Possibly Polinices
and Octopus prefer larger prey, as half their marks fell
among the few adult shells.

With the possible exception of man’s destruction of
snails in some localities, predation on adults is apparently
not important in the regulation of Olivella biplicata pop-
ulations, though losses among the more vulnerable young
may well be crucial. In nearly 3 years of observations
that included diving, only a few scattered cases of preda-
tion were observed. Neither is food availability likely
limiting in this gregarious, omnivorous species (Edwards,
unpubl. ), at least not until great densities (> 130 per m?
on bay beaches, over 700 per m? in lagoons) are attained,
and here again the stress falls primarily on the very young
(Epwarps, 1965). Olivella biplicata populations are often
heavily infected with parasitic trematodes (STEINMETZ,
1951; Duerr, 1965; Epwarps, 1968; Ivan Pratt, personal
communication), and well over 50% of adults may be
unsexed by the infections (Epwarps, 1968). Parasitic
disease definitely reduces natality and could limit popula-
tion sizes. Sporadic “disasters” may also kill many Oli-
vella. Exceptional fresh water flooding and silting took
heavy tolls in the San Diego Flood Control Channel in
November, 1965; and similar threats arise periodically
on the northern coast. Storms and winter sea turbulence
— also tidal waves like that from the 1964 Alaska earth-
quake — produce shifts of beach sand, posing dangers of
deep burial and transport out of suitable habitat. Whole
beaches may be carried away in winter, and one storm
(October, 1963) deposited 1-2m of new sand on a
Coos Bay, Oregon, beach, markedly reducing O. biplicata
densities (Epwarps, 1965). These erratic upsets eliminate
snails, but probably occur too rarely and unpredictably
to regulate densities or to account for the regular size-
frequency structure of most Olivella populations. The
size (age) structure in Olivella populations, charac-
terized by a preponderance of adults, probably results
from high mortality among juveniles (and possibly re-
duced adult fecundities in parasitized populations) and
excellent survivorship and individual longevity among
adults; turnover of the breeding population is likely
limited.

ACKNOWLEDGMENTS

I wish to thank Drs. Peter W. Frank and E. W. Fager for
making available facilities for these studies at the Oregon
Institute of Marine Biology and Scripps Institution of
Oceanography, respectively. I am also indebted to Mr.

Page 332

THE VELIGER

Vol. 11; No. 4

Nick Fotheringham and Drs. E. W. Fager and Robert T.
Paine for contributing useful information and to Drs.
Alan J. Kohn, Peter W. Frank, and Howard M. Feder
for critical comments on the manuscript. This work was
supported by National Science Foundation Predoctoral
and Postdoctoral Fellowships.

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DueErr, FREDERICK G.
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by crabs. Journ. Anim. Ecol. 33: 73 - 82
Epwarps, DaLias Craic
1965. Distribution patterns within natural populations of

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in reducing predation by starfishes.

to 512; 2 figs.
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1967. Studies on a sea-star (Marthasterias glacialis) extract

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1966. Aspects of asteroid biology. pp. 87-127 In: Physio-
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Intersci. Publ. (Wiley), New York.
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1964. Partial purification of a substance from starfish tube feet
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1966. An analysis of predation on gastropods. Unpubl.
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FRETTER, VERA & ALASTAIR GRAHAM
1962. _ British prosobranch molluscs, their functional anatomy
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GirrorD, DELiLa Sara GirFEN & Epwarp WINSLow GIFFORD
1942. Olivella pycna. Nautilus 55 (3): 92-93
Gonor, JEFFERSON JoHN
1965. Predator-prey reactions between two marine prosobranch
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Page 334

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Vol. 11; No. 4

Rediscovery of Terebra cochlea DrsHayes

BY

TWILA BRATCHER

8121 Mulholland Terrace, Hollywood, California 90046

(Plate 52)

In 1857 DESHAYES DESCRIBED A SINGULAR and beautiful
species of Terebra (T. trochlea) in the Journal de Con-
chyliologie, page 89, number 24 and illustrated on plates
5 and 6. This medium large species (holotype 65 mm X
13 mm) of a soft fawn color is decorated with flammules
of creamy white. The type locality was given as Zanzibar.
Two specimens of the type lot from the Cuming collection
are in the type collection of the British Museum (Natural
History). These I examined and photographed. Deshayes
also retained a specimen or specimens of the type lot in
his own collection (Ecole des Mines, Paris). In 1860 REEVE
published an excellent figure of this species in his Mono-
graph of the genus Terebra, in the Proceedings of the
Zoological Society of London (species no. 54), though
his description reverses the colors and lists the shell as
whitish with fawn markings. In 1885 Tryon included it
in the Terebra section of the Manual of Conchology
(vol. 7, p. 16; plt. 12), and in 1888 Partet included it in
the “Catalog der Conchylien Sammlung” (p. 255). He
also included T: cochlea DEsuHayYEs, type locality Zanzi-
bar, which obviously was a misspelling for T: trochlea.
For nearly a hundred years T: trochlea seems to have been
dropped completely from molluscan literature, and since
the type lot this species has not been reported from
Zanzibar or any other locality.

In July 1967 Clifton S. Weaver of Hawaii sent me for
examination a specimen of Terebra which matches the
holotype of 7: trochlea. This specimen was collected at
Nukuhiva in the Marquesas by Richard Sixberry. At the
convention of the American Malacological Union in 1968
at Corpus Christi, Texas, another specimen of this species
was shown on a slide accompanying a paper by Dr. Harald
Rehder of the Uniter States National Museum, though at
that time the Jerebra was unidentified. This latter speci-

men (U.S.N. M. no. 679155) was collected by Richard
Sixberry also at a depth of 20 feet in muddy sand at
Taiohae Bay, Nukuhiva, Marquesas. The specimen is 65.3
mm X 12.8mm.

Though it is possible that this species occurs in the

Zanzibar area, one must assume that, like many other
species of Terebra described in early literature, the type
locality is erroneous unless other specimens are reported
from that area.
Description of the shell: Shell medium large; color fawn
to light reddish brown with flammules of creamy white;
whorls slightly convex and turreted, shouldered anterior
to prominent convex subsutural band which is set off by
well defined suture and broad, deeply impressed subsu-
tural groove which is microscopically punctate in the
early whorls; sculpture of early whorls of indistinct axial
grooves and obsolete flat axial ribs which form small
nodes at their posterior endings on both whorl and sub-
sutural band; in later whorls axial grooves become nu-
merous fine axial striations, but nodes continue to occur
anterior to suture and subsutural groove; body whorl of
medium length; aperture ovate; outer lip thin with color
pattern showing through; columella white, heavy, quite
straight; siphonal fasciole well defined, striate, set off by
a sharp keel; anterior canal very broad, short, recurved.

ACKNOWLEDGMENTS

I wish to thank Dr. Harald A. Rehder of the Smithsonian
Institution for the loan of the specimen of Terebra troch-
lea DesuHayeEs for study and photography. I also wish to
thank Clifton S. Weaver for sending for my examination
the specimen which brought this species to my attention.

Explanation of Plate 52

Terebra trochlea DESHAYES, 1857

Figure 1: Holotype and paratype, British Museum (Natural His-
tory) ; holotype on left.

Figures 2, 3, 4: Specimen no. 679155, in the United States Na-
tional Museum.

THE VELIGER, Vol. 11, No. 4 [BRATCHER] Plate 52

Figure 2

mV)

” ‘

Figure 3 Figure 4

photographs by Twita BRaATCHER

Vol. 11; No. 4 THE VELIGER Page 335

LITERATURE CITED

DesHAYES, GERARD PAUL
1857. | Descriptions d’espéces nouvelles du genre Terebra.
Journ. de Conchyl. 6: 65 - 102; plts. 3-5
PAETEL, FRIEDRICH
1875. Die bisher veréffentlichten Familien- und Gattungs-
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1888. Mit Hinzufiigung der bis jetzt publicierten Recenten
Arten, sowie der ermittelten Synonyma; Fam. Terebridae:
pp. 247 - 255
REEvE, LovELL AuGUSsTUS
1860. | Conchologia Iconica. Monograph of the genus Terebra.
12: plts. 1-27; species 1-155; a descriptive letter press plus
index
13: plts. 1 - 27 (May - June 1860)
Tryon, GEorcE WASHINGTON, Jr.
1885. | Manual of Conchology. Monograph of the family Te-
rebridae. 7: 1-64; plts. 1-12. Philadelphia

Page 336

THE VELIGER

Vol. 11; No. 4

Growth Characteristics of Acmaea persona EscHSCHOLTZ'

RON KENNY

Zoology Department, University College of Townsville, Queensland, Australia

Marine Science Center, Oregon State University, Newport, Oregon 97365

(4 Text figures)

WILBuR & Owen (1964) HAVE REVIEWED the literature
on growth studies of mollusks and Frank (1965a, 1965b)
has discussed some aspects of growth for 3 species of
Acmaea on the Pacific coast of North America.

Acmaea persona ESCHSCHOLTZ, 1833 has a geograph-
ic range from latitude 37°N to latitude 60°N on the
Pacific coast of North America (FrircHMan, 1962). The
vertical range of the species in the intertidal zone ex-
tends approximately from mean sea level to mean higher
high water. Characteristically this limpet is found in

' Supported by U.S. Office of Naval Research, Project Number
NR 104-936, “Marine Ecological Studies.”

areas sheltered from the heaviest wave action, such as the
heads of small bays (KENNy, in MS).

The specimens used in this study were collected at
Boiler Bay and Yaquina Head near Newport, Oregon
(lat. 44° N; long. 124° W) during the summer of 1967.

During the collecting period temperatures in the area
ranged from 10° to 23° C (air), 8.5° to 16°C (sea), and
10.5° to 31°C (rock surface). January sea temperatures
for the area range between 7.7° and 10.3°C (Wyatt,
STILL & Haac, 1965) and the mean range of air tem-
peratures for January is 3.4° to 9.7°C (U.S. Weather
Bureau records).

No particular selection was exercised in collecting these
limpets, but as the collections were made initially for
physiological experiments, they are representative of size
range rather than truly random collections. One hundred
and eighty-nine limpets were collected and the analysis
of annual growth rings is based on records from 166
individuals in which the rings were considered not to
have been affected by erosion. Allometric growth data
are based on measurements of approximately 100 limpets.

Measurements were taken of shell length, breadth and
height (mm), shell weight (gm) and tissue weight (gm)

Figure 1

Dorsal and lateral views of Acmaea persona, length 44.4mm,

Vol. 11; No. 4

after removal from the shell and surface drying with
paper towelling. Shell volume (ml) was measured with
distilled water containing liquid detergent, from gradu-
ated pipettes.

In analysis and graphical presentation all measure-
ments have been related to shell length.

The step-like ridges on the shell surface are caused by
differences in the rate of shell formation. As the result
of observations in the field on a small number of marked
animals, Dr. P. W. Frank has stated (personal communi-
cation) that these ridges are annual growth rings formed
in winter (Figure 1). Further, within each annual step,
fine, regularly arranged, secondary rings may be seen
under magnification.

The measurements of shell length have been arranged
in classes on the basis of number of annual rings and the
size frequencies for these classes are shown in Figure 2.
As the shell edge must be presumed to be forming part
of the next annual ring, the age of the limpet is the annu-
al ring number of years plus part of a year.

The number of animals in the 0 ring class is too small
to warrant detailed analysis, but the samples for other
classes show a mean annual increase in shell length of
6 mm in the second year, 4 mm in the third year, 4 mm in
the fourth year, and 3 mm in the fifth year.

Shell Length in mm

O I II III IV Vv VI
Ring Class

Figure 2

Frequency distribution of shell length related to number of annual
showing annual rings; drawn from glass tracings.
rings, for Acmaea persona.

As the largest specimen collected was 44.4mm in
length, it is probable that the life span is considerably
longer than 6 years.

THE VELIGER

Page 337

Counts of the number of secondary rings occurring on
the surface of an annual growth area (both summer and
winter sections of the step) were made from 55 individu-
als, with no surface erosion, ranging in shell length from
25.3 mm to 35.8 mm and showing 2, 3, or 4 annual rings.
In each case the count was made on the area one step in
from the shell edge, i. e. irrespective of age, for the same
period of time. The number of rings recorded varied
from 114 to 201 with a mean of 149.

The allometric patterns of growth for various shell
dimensions and for the soft tissues are shown in Figures
3 and 4. Growth equations for these relationships are
listed below, where L is shell length in mm, B is shell
breadth in mm, H is shell height in mm, V is shell volume
in ml, W is shell weight in gm, and wt is tissue weight in
gm.

B= 0.85 L- 1.2
log V = 3.27 (log L) — 4.4
log wt = 4(log L) —5.7

30 A
25

20

mls : gms

15

xe 15 20 25 30 35 40
Shell Length in mm

Figure 3

Relationship between shell length and shell breadth (A) and shell
height (B), for Acmaea persona.

The relationships of shell height and shell weight to
shell length are more complex and neither growth pattern
can be described readily by a single mathematical formula
throughout the range of shell lengths. The shell weight
relationship can be expressed as two equations, one for
small shells and a second for larger specimens, with the
change of form taking place at lengths between 25 and
28 mm.

Page 338

log W = 2.7 (log L) — 3.9 (small shells)

log W —5 (log L) —7.2 (large shells)
Similarly the shell height to length equation,

log H = (log L) —0.6

: gms

mls

10 15 20 25 30 35 40
Shell Length in mm

Figure 4

Relationship between shell length and shell volume (A),
shell weight (B), and tissue weight (C), for Acmaea persona.
The vertical scale represents millilitres for the shell volume graph,
and milligrams for the shell weight and tissue weight graphs.

applies only to small shells and for shells longer than
25 mm, a single formula cannot be derived.

Frank (1965a, 1965b), commenting on other species
of Acmaea (A. pelta EscuscHo.tz, 1833, A. digitalis
EscuscHo tz, 1833, A. paradigitalis FrircHMan, 1960),
has noted that due to the wide size range in any age
class, size alone does not permit an estimate of either
age or longevity. A similar statement would apply to
A. persona.

Hyman (1967) has combined the results of studies on
Patella vulgata LinnaEus, 1758 from English coasts by
RussELL & Orton to give size ranges for age classes of
that species. Calculations from these data show P. vulgata
having a 50% increase in length in the second year and a
28% increase in the third year, compared with corre-
sponding figures of 36% and 16% for Acmaea persona.

The suggested probable life span of more than 6 years
is similar to the “5 to 16 years” listed for Patella vulgata
by Hyman (op. cit.). Comrort (1957) gives 15 years
as the life span for Acmaea dorsuosa Gou.tp, 1859 and
Darsy (1964) discussing a gastropod from the Oregon

THE VELIGER

Vol. 11; No. 4

coast, Tegula funebralis (A. ApaMs, 1855), mentions
individual specimens estimated to be 12 and 30 years old.

The graphs for length-breadth and length-height rela-
tionships are similar in form to those presented by HAMAI
(1937) for Patelloida grata (GouLp, 1859). The increase
in height of the shell in older specimens of Acmaea per-
sona can be related to the change of direction of shell
growth in older animals (see lateral view, Figure 1).

“Ledging” ‘has been reported by Moore (1934) in
Patella vulgata. Moore described this as a “temporary
change in the angle of shell growth” and considered it to
be due to slowing of shell formation during the winter.
As noted above, FRANK has suggested a similar explana-
tion for the formation of annual rings in Acmaea persona
shells.

From the available data the overall winter range of
environmental temperatures for Acmaea persona is 3.4°
to 10.3° C and the overall summer range 8.5° to 31° C.

WILBUR & Owen (1964) cite several studies showing
direct relationships between growth rate of mollusks and
environmental temperature. It seems probable that the
annual differences in rate of shell growth in Acmaea per-
gona are primarily related to seasonal differences in en-
vironmental temperatures in the intertidal zone.

The number of secondary rings developed in an annual
growth area suggests the number of short term active
growth periods in one year. The wide range recorded for
Acmaea persona shows considerable individual variability
in number of short term growth periods per annum. For
an intertidal species, such as A. persona, this variability
is probably influenced by several environmental factors
(microclimate, tidal patterns, and wave action) as well
as by physiological factors.

ACKNOWLEDGMENT

The author wishes to thank Dr. Joel W. Hedgpeth for
the laboratory accommodation and facilities made avail-
able at the Oregon State University, Marine Science
Center, Newport.

LITERATURE CITED

Comrort, ALEXANDER

1957. ‘The duration of life in molluscs.
London 52: 219 - 241

Darsy, Ricuarp L.

1964. | On growth and longevity in Tegula funebralis (Mollus-
ca: Gastropoda). The Veliger 6, Supplement: 6 - 7;
pit. 1. (15 November 1964)

Frank, PETER WOLFGANG

1965a. Growth of three species of Acmaea.
7 (3): 201 - 202; 1 fig.; 1 table

1965b.
Ecology 46 (6) :

Proc. Malacol. Soc.

The Veliger
(1 January 1965)

831 - 844; 8 figs.; 6 tables

The biodemography of an intertidal snail population.

—

Vol. 11; No. 4 THE VELIGER Page 339

FRITCHMAN, Harry K.., II
1962. A study of the reproductive cycle in the California
Acmaeidae (Gastropoda). Part IV. The Veliger 4 (3): 134

to 140; plts. 30 - 32 (1 January 1962)
Hamat, I.
1937. | Some notes on relative growth, with special reference to
the growth of limpets. Sci. Repts. Tohoku Univ., 4" ser.,
12: 71-95

Hyman, Lipsy H.
1967. Mollusca, I - vol.6 of The invertebrates. McGraw-
Hill, New York
Kenny, Ron P.
(in MS) Ecology of the limpet genus Acmaea on the Oregon

coast.
Moore, Hitary Brooke
1934. On “ledging” in shells at Port Erin. Proc. Malacol.

Soc. London 21: 213 - 222
Wixeur, K. M. & GARRETH OWEN
1964. Growth. In K. M. Wilbur « C. M. Yonge (ed.) Phys-
iology of Mollusca. Acad. Press, New York
Wyatt, B., R. Stitz, & C. Haac
1965. Surface temperature and salinity observations at Pacif-
ic Northwest shore stations for 1963 and 1964. Dept.
Oceanogr., Oregon State Univ., Data Rept. 21

Page 340

THE VELIGER

Vol. 11; No. 4

Use of the Propodium as a Swimming Organ in an Ancillid

(Gastropoda : Olividae)

BY

BARRY R. WILSON

Western Australian Museum, Perth, Western Australia

(Plate 53; 1 Text figure)

AMONG THE PROSOBRANCH molluscs swimming as the
prime method of locomotion is confined to the pelagic
Heteropoda although there are reports that several mem-
bers of the families Olividae and Naticidae are capable
of brief “flights” of swimming activity.

p’Orpicny (1841, p. 418) observed this type of loco-
motion in Oliva tehuelchana (p’Orxicny, 1841), and
Oxsson (1956, p. 161) mentioned the swimming ability
of species of the genus Olivella. Marcus & Marcus
(1959, p. 103) reported swimming in Olivella verreauxu
(Ducros, 1857). In all these cases the movement was
effected by the lateral wing-like flaps of the metapodium
which characteristically fold over and cover the shell in
members of the Olividae. There are no details in these
reports concerning the duration of the swimming activity
or the circumstances in which it occurred.

ZIEGELMEIER (1958) reported swimming in the Medi-
terranean naticid Polinices josephinus (Risso, 1838). In
this case the propodium is the propulsive organ. The
animal “swims” in two positions. Most frequently it re-
mains in the normal crawling position and the “whip-
ping” of the propodium lifts the front end of the body
off the sand and propels it forward. The posterior part of
the foot remains in contact with the surface of the sand.
Less commonly the animal twists into an upside-down
position and the “whipping” of the propodium lifts the
animal off the substrate and propels it forward a distance
of 3 to 5cm (in a large animal) for each stroke. Each
propodium stroke is from the horizontal position upwards,
almost as far as the anterior wall of the shell, and then
downwards again to the original position, and lasts be-
tween 1 and 2 seconds. ZIEGELMEIER interpreted both
forms of “swimming” behaviour as escape or “flight”
reactions. It occurred when crawling animals were
touched with a stick or when they collided with another
animal.

Similar behaviour by the ancillid Ancillista cingulata
(SowerBy, 1830) was observed on a recent field trip in
the North-West of Western Australia. A single specimen

was collected in fairly coarse sand on an intertidal sand
flat on the western side of Point Cloates (22° 43’S;
113° 40’ E). It was crawling beneath the surface of the
sand and left a long trail, wider than those of Oliva orna-
ta Marrat, 1870, which is common at the same place.
The preserved body and the shell of the specimen are now
in the collection of the Western Australian Museum (cat.
no. 4902-68).

External Anatomy of Ancillista cingulata

Like other olivids, the foot of this specimen of Ancillista
cingulata is divided by curved transverse grooves on both
dorsal and ventral surfaces into a crescent-shaped anterior
part, or propodium, and a posterior part, or metapodium.
(For detailed structure of Oliva, Olivella and Olwancil-
laria see Marcus & Marcus, 1959). The anterior edge of
the propodium does not have a deep horizontal groove as
in Oliva and Olivella but there is a deep medial longitu-
dinal groove from the anterior margin to the posterior
margin on the dorsal surface. At its posterior end the me-
tapodium is deeply forked, a characteristic of the subfam-
ily Ancillarinae (see Otsson, 1956, p. 169). The much
reduced head is completely covered by the anterior ends
of the lateral metapodial flaps, and is represented by only
2 small flap-like tentacles. There are no eyes. Beneath the
right hand tentacle the small mouth is located. There is a
very thin, elongate operculum carried on the dorsal sur-
face of the metapodium medially, below the spire and
concealed by the posterior ends of the lateral metapodial
flaps.

In the fully extended crawling position (Plate 53) the
foot was greatly expanded (approximately 9cm_ long,
6cm broad) and the length of the propodium was a
little less than one quarter of the total foot length, i. e.
rather larger than in other olivids. The lateral flaps of
the metapodium covered the body whorl of the shell,
except when the animal was disturbed, but they gaped
around the spire so that that part of the shell was usually

Tue VELIcER, Vol. 11, No. 4 [Witson] Plate 53

A living specimen of Ancillista cingulata (SowERBY, 1830),
collected at Point Cloates, Western Australia,
in the fully extended crawling position.

Vol. 11; No. 4

THE VELIGER

Page 341

visible. The siphon was carried in an oblique-erect posi-
tion. The body was white, flecked with pale grey and
fawn.

Swimming Behavior of Ancillista cingulata

The animal made no attempt to burrow into a thin
cover of sand which was put in the bottom of the dish
of water. Its usual method of locomotion was rapid
gliding aided by secretion from the anterior edge of the
propodium of a thick mucous sheet which passed over
the dorsal and ventral surfaces of the body. When the
animal was agitated by prods with a pencil it suddenly
threw itself into a series of violent contractions which
carried it to the surface of the water and around the dish
in a jerky, apparently random manner, reminiscent of
the random swimming movement of scallops. The pro-
podium repeatedly flapped backwards from the horizon-
tal plane, first dorsally and then ventrally at regular
intervals of slightly more than one second. Each move-
ment threw the animal forward a distance varying be-

Sp

Figure 1

Diagram illustrating swimming movement by beating
of the propodium.in Ancillista cingulata.
p = propodium; m = metapodium; sp = shell spire; s = siphon.
1. dorsal stroke; 2. intermediate position (and crawling position) ;
3. ventral stroke.

tween 5 and 15 cm. The ventral beat appeared to be the
most effective. Most of the time the animal remained in
the normal horizontal position with the shell uppermost,
but “barrel-rolls” were frequent. This activity continued
for 45 seconds, during which time the propodium flapped
35 times. The animal then sank to the bottom and re-
sumed its crawling activity.

During the next 24 hours the same swimming response
was observed several times, sometimes stimulated by agi-
tation and sometimes occurring spontaneously. Each time
the number of beats and the duration of the swimming
period were about the same as before. Although the prog-
ress of the animal was erratic and confined within the
dish, it is estimated that the total distance travelled in
the open water during each burst of swimming activity
would be about 3 meters.

The following day a thick layer of sand, an inch or so,
was put in the dish and the animal buried itself and there-
after remained motionless with only the siphon tip visible.
When dug out and agitated, it again responded by the
swimming reaction with the same duration as before.

It seems most likely that this response is an escape re-
action. The distance travelled and the erratic progress
would allow the animal to evade most predators if this
is its function. Observations on the locomotory behaviour
of other ancillids which have a similarly broad propodi-
um would be of interest. It is noteworthy that in the two
unrelated families Naticidae and Olividae similar foot
structure, i. e. division into propodium and metapodium,
has been evolved independently, and that in each family
at least one species has made use of the propodium as a
swimming organ.

SUMMARY

A specimen of the ancillid Ancillista cingulata SowERBY,
collected at Point Cloates, Western Australia, was ob-
served to swim by dorso-ventral flapping of its broad
propodium. This behaviour is interpreted as an escape
reaction.

ACKNOWLEDGMENTS

Dr D. J. G. Griffin of The Australian Museum and Miss
Anne Paterson of the Western Australian Museum shared
in this observation. Mrs Shirley Slack-Smith helped in
the literature search for other records of swimming in
prosobranchs and Mrs A. Neumann kindly translated
Ziegelmeier’s paper.

Page 342 THE VELIGER Vol. 11; No. 4

LITERATURE CITED

Marcus, EvELINE DU Bots-REYMOND & ErNstT Marcus

1959a. Studies on Olividae. Univ. Sao Paulo Fac. Filos.

Cienc. e Letras. Bol. Zool. 22: 99 - 188; plts. 1 - 11
Oxtsson, AxEL ADOLF

1956. Studies on the genus Olivella. Proc. Acad. Nat. Sci.

Philadelphia 108: 155 - 225; plts. 8 - 16
p Orsicny, ALCIDE DESSALINES

1834-1847. Voyage dans l’Amérique Méridionale. Mollusques.

Paris, vol. 5, pt. 3, xliii + 758 pp., atlas, 85 plts.
[not seen]
ZIEGELMEIER, E.

1958. Zur Lokomotion bei Naticiden (Gastropoda Prosobran-
chia). Kurze Mitteilung uber Schwimmbewegungen bei Poly-
nices josephinus Risso. Helgolander wissenschaft. Meeres-
untersuch. 6: 202 - 206

Vol. 11; No. 4

THE VELIGER

Page 343

A New Genus and Two New Species of Typhinae

from the Panamic Province

(Gastropoda : Muricidae)

BY

HELEN DuSHANE

Research Assistant, Invertebrate Zoology, Los Angeles County Museum of Natural History

Los Angeles, California 90007

(Plate 54)

IT SEEMS DESIRABLE to put on record the following un-
described forms of Typhinae. One is of a group previously
not known in the Panamic province. In Typhinae, with
a world wide range of distribution from the Eocene to
the Recent, it is unusual to have in one’s possession two
new species to be described, one of them even requiring
a new genus. Since KEEn’s (1944) revision of the Typhi-
nae, new interpretations of typhine morphology have been
projected by FLeminc (1962) and Vetta (1961). How-
ever, until such time as workers have expressed their views
in a proposed revisional work the author prefers to fol-
low the outline presented by KEEN (op. cit.).

Cinclidotyphis DUSHANE, gen. nov.
(Plate 54, Figures 1 to 3)

Varices 4 per whorl; sculpture of numerous axial riblets
and fine, raised spiral cords that are continuous across
the varices and interspaces; tubes folded back, not form-
ing vertical ribs; cancellate sculpture distinctive and un-
like that in all other species of Typhinae.

Type Species: Cinclidotyphis myrae DUSHANE, spec. nov.

Cinclidotyphis myrae DUSHANE, spec. nov.
(Plate 54, Figures 1 to 3)

Shell small, fusiform, color dingy white, protoconch white
(nuclear whorl worn off in holotype), followed by 4
subsequent whorls; shoulder narrow, sloping, not deeply
channeled between whorls; varices 4 per whorl, each a
rounded fold which extends above shoulder to join pre-
ceding varix; tubes folded back, not forming vertical ribs,

each with a suture on the anterior surface which is carried
horizontally onto face of succeeding varix; varices with
prominent spiral and axial sculpture producing a beaded
effect at the junction, about 22 spiral cords on the last
whorl; every other cord at the middle of the body whorl
being smaller; 18 axial ribs on the body whorl at a line
with the top of the aperture; aperture oval elongate and
set off by a raised margin; lip narrow and crenulated;
anterior canal open, narrowing toward the base with a
slight dorsal curve; operculum lacking in type.
Type Material: The holotype is on deposit in the Los
Angeles County Museum of Natural History, Invertebrate
Zoology Type collection, catalogue number 1194.
Type Locality: The holotype was collected by the author
at Tenacatita Bay, Jalisco, Mexico, January 25, 1968,
among rocks near a sand beach; Lat. 19°16’50” N; Long.
104°48’27” W. It is the only specimen known.
Dimensions: Height 13 mm; maximum diameter 6.5 mm.
Discussion: The appearance of this shell in the Panamic
province is all the more remarkable because of the paucity
of world wide material in the nearest related genus,
Siphonochelus. The largest concentration of species of
this genus occurs in the Tertiary of Europe with the
earliest known species S. parisiensis (D’OrBIGNY, 1850)
dating back to the Middle Eocene according to KEEN &
CampBELL (1964). The genus is represented in the Aus-
tralian fauna with 5 species and in the Japanese fauna
with 2, but up to the present time no specimens of this
genus have been reported from the Panamic province.
The Mexican specimen differs from those in the genus
Pterotyphis by having more than 3 tubes per whorl and
in having finer sculpture. It has weak varices as in
Siphonochelus, with 4 tubes per whorl. The cancellate
sculpture is unlike that in all species of Siphonochelus.
The open canal is distinctive. Perhaps it is closed in some

Page 344

part of the growth cycle, but one would have to have
more material to be certain. Another peculiar feature
is the manner in which the tubes seem to have been folded
back, not forming vertical ribs.

This species is named in honor of Dr. A. Myra Keen
whose ready help is always a boon to the worker.

Pterotyphis (Tripterotyphis) arcana DUSHANE, Spec. nov.
(Plate 54, Figures 4 to 6)

Shell small, base and tubes ivory white, body whorl brown
with blotches of brown on earlier whorls, whorls 5 to 6,
nuclear whorls two in number, rounded, smooth; adult
sculpture developed in first postnuclear whorl, the third
with the first tube; varices 3 per whorl, the upper end
of each varix left open as a tube; tube openings elevated
and at tips of spine; outer face of body whorl with 2
strong white spiral ribs riding over the varices; aperture
oval, outer lip with a wide sinuous margin, reflected
against the ends of 6 to 10 spiral ribs, smooth within;
anterior canal sealed except at end; pillar with remnants
of 2 previous canals.

Type Material: The holotype is on deposit in the Los
Angeles County Museum of Natural History, Invertebrate
Zoology Type Collection, catalogue number 1195; 2 para-
types, DuShane collection.

Type Locality: Mazatlan, Sinaloa, Mexico; Lat. 23°11’N,
Long. 106°26’ W. The 3 specimens were collected by the
author on a rocky reef, February 24, 1968.

Two additional specimens from Banderas Bay, collected
in January 1969 by J. DuShane indicate that the range
of this species extends at least as far as 275 miles south of
the type locality.

Dimensions: Holotype, height 15 mm, maximum diameter
8mm, length of aperture 3.2mm; Paratype 1, height 16

THE VELIGER

Vol. 11; No. 4

mm, diameter 8.5mm; Paratype 2, height 14mm, dia-
meter 7mm.
Discussion: Unlike Pterotyphis (Tripterotyphis) fayae
KEEN & CAMPBELL, 1964, which is sculptured with about
22 spiral ribs, specimens of the new taxon have 2 major
cords or ribs on the body whorl. Also, the brown coloration
is less diffused on P (T.) arcana. It differs from P (T.) lowei
Pitssry, 1931 of the West Central American coast, by
being easily separable on the basis of the 2 major ribs
and the remnants of 2 previous canals; P (T:) lowei has 3.
The earliest fossil record seems to be in the Miocene of
Europe. Recent species occur in the Caribbean, Panamic,
and Gulf of California areas.

ACKNOWLEDGMENTS

I am grateful to Dr. A. Myra Keen for her encouragement
in suggesting that specimens of this importance should
be reported, and to Dr. Bruce Campbell for his critical
reading of the manuscript. Photographs are by Mr. Per-
fecto Mary, Stanford University.

LITERATURE CITED

FLemine, C. A.

1962. The genus Pterynotus Swainson (Gastropoda, Family
Muricidae) in New Zealand and Norfolk Island. Trans. Roy.
Soc. New Zealand, Zool. 2 (14): 109-119; 22 figs.

KEEN, A. MYRA

1944. Catalogue and revision of the gastropod subfamily

Typhinae. Journ. Paleontology 18 (1): 50-72; 20 figs.
Keen, A. Myra, & G. BRucE CAMPBELL

1964. ‘Ten new species of Typhinae (Gastropoda: Muricidae).

The Veliger 7 (1): 46-57; plts. 8-11; 3 text figs. (1 July)
VELLA, PAUL

1961. Australasian Typhinae (Gastropoda) with notes on the

subfamily. Palaeontology, 4 (3): 362 - 391; plts. 46 - 47.

Explanation of Plate 54

Figure 1: Cinclidotyphis myrae DUSHANE, gen. nov., spec. nov.
Ventral view of holotype, LACM, Invertebrate Zoology Type Col-
lection, catalog number 1194 (X 4)
Figure 2: Lateral view of the holotype
Figure 3: Apical view of the holotype

Figure 4: Pterotyphis (Tripterotyphis) arcana DUSHANE, spec. nov.
Ventral view of holotype, LACM, Invertebrate Zoology Type Col-
lection, catalog number 1105 (X 4)
Figure 5: Lateral view of the holotype
Figure 6: Apical view of the holotype

THe VELIGER, Vol. 11, No. 4 [DuSHan_E] Plate 54

Figure 1 Figure 2 Figure 3

Figure 4 Figure 5

Vol. 11; No. 4

THE VELIGER

Page 345

Gross Anatomy and Classification of the Commensal Gastropod,

Caledoniella montrouziert SOUVERBIE, 1869

BY

JOSEPH ROSEWATER

Division of Mollusks, U.S. National Museum, Washington, D.C. 20560

(Plate 55; 4 Text figures)

SPECIMENS OF A COMMENSAL GASTROPOD were reported
by Manninc (1968) on two species of Gonodactylus
(Crustacea:Stomatopoda) from the southwest Indian
Ocean. Upon examination these were found to be Cale-
doniella montrouziert SOUVERBIE, 1869, originally described
from New Caledonia, a species whose anatomy was virtu-
ally unknown and whose systematic relationships are poorly
understood. In the present paper an attempt is made to
describe the gross anatomy of this species, comment on
its supposed habits and to suggest for it a familial allo-
cation.

HISTORICAL

The monotypic genus Caledoniella was described briefly
by SouversicE (1869) with C. montrouzieri as its only
contained species. In the following year SOUVERBIE &
Montrouzier (1870) supplemented the original descrip-
tion and published a figure of the shell, commenting that
the animal had been found living on the thoracic append-
ages of Gonodactylus. They made no assignment of the
taxon to a higher category. Tryon (1886) assigned the
species to the subfamily Lamellariinae of his family Nati-
cidae with some doubt because the animal was unknown.
As noted by ALLAN (1936), BAsEpow (1905) described
a second supposed species of C’aledoniella which subse-
quently has been found not to belong to that genus.
Preston (1912) described Epistethe gonodactyli from the
Persian Gulf and Andaman Islands, and it is now con-
sidered to be an absolute synonym of C. montrouzieri.
He gave the first sketchy description of the animal, but

did not assign it to a family. ALLAN (op. cit.) reported
specimens of Caledoniella attached to a Gonodactylus
from Albany Passage, Cape York, Queensland. In all
probability these are C’. montrouzieri (see Text figure 1).

Latest attempts at suprageneric classification of Cale-
doniella are those of THrete (1929) and Wenz (1940).
THIELE placed it in the Superfamily Lamellariacea, sub-
family Lamellariinae and described a radula (see below
under Radula). The assignment quite probably was made
on the basis of shell shape since Caledoniella resembles
Lamellaria, WENz placed it in the Superfamily Pyrami-
dellacea, Family Stiliferidae, probably because of the
commensal relationship with Gonodactylus.

Hoituuis (1951) reviewed the known information
relating to the geographic occurrence of Caledoniella
montrouziert and mentioned the presence of gastropod
egg-capsules on the ventral surface of several of its host
Gonodactylus. The same author (1941) figured similar
egg-capsules and suggested that they were probably those
of Epistethe (= Caledoniella; see Text figure 4).

ACKNOWLEDGMENTS

I express my appreciation to Raymond B. Manning for
calling to my attention the specimens of gastropods upon
which this paper is based and to both him and L. B.
Holthuis for discussing the relationship with Gonodactylus.
I am also grateful to Mrs. Carolyn Bartlett Gast for
preparing the plate illustrating animal and shells of Cale-
doniella montrouzieri. Ideas and interpretations are mine
alone and I accept full responsibility for them.

Page 346

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EXTERNAL ANATOMY

Material:
Shell Tocality
H' W’
1. Male 2.6 4.3 AnjouanId, USNM 679176
(Plate 55, Figs. A - F) Comoro Ids.
2. Female 2.8 5.2 AnjouanId, USNM 679176
(not figured ) Comoro Ids.
3. Male 14 2.2 ‘Tulear, USNM 679177
(Plate 55, Figs. G- I) Madagascar
4. Female 2.8 4.6 ‘Tulear, USNM 678177
(Plate 55, Figs. J - L) Madagascar

H' = height in mm; W? = width in mm

The male animal from Anjouan, Comoro Islands, was
used mainly for the following description and accompa-
nying Plate. The three other available specimens were
compared and found to agree.

Foot: Preston’s (1912) description, although brief, char-
acterized the foot and its epipodial fringe. He described
the foot as an “‘adhesive organ’, and this may be the case
(see Plate 55, Figure C). The ventral surface of the foot
(mesopodium) is disklike, and in one of the specimens
examined was covered with a thin, chitinous “epidermis”
which may be secreted for attachment to the host. Attach-
ment is here not considered to be permanent, as evidence
gained from a specimen examined zn situ indicates these
gastropods move about on the host (see Reproduction).
At the anterior edge of the foot there is a groove which
ascends dorsally to the base of the propodium.
“Epipodial Fringe”: As described by Preston (1912)
there occurs dorsal to the foot-disk, laterally and posteri-
orly, a foliated (in preserved specimens) fringe of tissue
which he termed the “epipodial fringe” (Plate 55, Figures
A to C). This structure appears to be the metapodium.
Its function in living Caledoniella has not been reported,
but may be analogous to that in Natica (FRETTER & Gra-
HAM, 1962, fig. 304) in partially covering the shell. A
somewhat similar metapodial fringe was described and
figured by Quoy « Garmarp (1832, 1833) and by BERcH
(1896) in Vanikoro cancellata (LAMaRcK, 1822) (see
Rosertson, 1962 and Text figures 2, 3).

An operculum was not observed in Caledoniella mont-
rouziert. It is probable that the operculum has been lost
during the evolution of the species in connection with its
adaptation to existence as a commensal.

Propodium: Located antero-medially above the anterior
edge of the foot and between the anterior edges of the
“epipodial fringe” there is a partially bilobed structure

Vol. 11; No. 4

which was dorso-ventrally folded in the preserved animal
(Plate 55, Figures A to C). This is the propodium. When
unfolded (Figure A), it forms a structure which appears
to be highly utilitarian. YoncE (1953) has suggested that
in Hipponix antiquatus (Linnagus, 1767) the propodi-
um may be concerned with egg-capsule attachment. He
did not mention the structure in males of H. antiquatus,
although it may be shown in the figure of a male (see

Figures 1 to 4

Vol. 11; No. 4

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Page 347

Yonce, 1960, fig. 1). It is well developed in both male
and female Caledoniella montrouziert. Since it is prob-
able that only the female takes part in egg-mass deposi-
tion, the presence of the propodium in males may signify
protandry, or the structure may have some additional
function, such as in feeding.

Proboscis: A prominent, muscular and extensible pro-
boscis is present above the propodium with the mouth as
a vertical slit at its extremity.

Radula: A radula was not found, although three of
the animals were dissected in search of that structure.
The only known report of a radula in Caledoniella mont-
rouzieri is that of THrere (1929): “Mittelplatte der
Radula rundlich quadratisch, Schneide breit dreieckig,
jederseits mit einigen Zahnchen, Zwischenplatte maBig
gro, Schneide spitz, mit einer inneren Nebenzacke, die
beiden Seitenplatten mit einfachen hakenformigen Spit-
zen.” Unfortunately the radula was not figured, and
THIELE’s description contains no clue to the relationships
of Caledoniella. 1 am somewhat confused by the absence
of a radula in the specimens examined by me and am
led to question THIELE’s report until it is confirmed.
Tentacles: A pair of fleshy tentacles is located above the
proboscis. Due to preservation it is difficult to be certain
how extensible these may be in the living animal, but
they are relatively long. Tentacles are joined by a con-
necting lobe between their bases. The eyes occur as black-
pigmented spots buried within the semitransparent tissue
of the tentacles and are located somewhat toward the
outer edges of their bases. Similar eyes have been re-
ported in Vanikoro (BEercuH, 1896) and in Natica (FReET-
TER & GRAHAM, 1962).

Penis: The penis, located near the base of the right
tentacle, is long and slender in the two male specimens
examined (see Plate 55, Figures A to C). It is simple
(i. e., unbranched) and semitransparent; there is a
closed gonoduct passing through its tissue.

Mantle and Mantle Cavity: In the illustration (Plate 55,
Figures A to C) the mantle edge is rolled back to reveal
the animal’s tentacles, eyes and penis. The mantle edge
is grayish-white in color and somewhat thickened; behind
the edge the outside of the mantle is darkly pigmented, the
dark color showing through the transparent shell; it is
semitransparent and thin in the region of the ctenidia.
The ctenidia are not excessively developed, nor is the
mantle cavity extremely large.

Shell: There is evidence of sexual dimorphism in the shells
of male and female specimens, especially in size. All are
wider than long. In shells of male Caledoniella montrouz-
teri (Plate 55, Figures D-I) the body whorl is by far the
major constituent and the spire consists of the proto-
conch and only about one subsequent whorl. Shell color

is pale yellowish-white and there is a shiny darker yellow
periostracum which becomes deciduous upon drying.
Sculpture is limited to fine, irregular axial lines of
growth. The columellar lip may be roughly formed and
rather deeply concave. Columellar muscle scar is elon-
gate-oval, large and well defined; it is clearly visible
within the aperture on inner wall of outer lip near pos-
terior junction of outer lip and columella. The name
“columellar muscle scar” in C. montrouzieri is somewhat
a misnomer, as it is not altogether located on the colum-
ella! The muscle probably performs the same function
as in other prosobranchs, that of pulling the animal into
its shell.

In the female shell (Plate 55, Figures J to L) the spire

of the specimen from Tulear is definitely lifted above the
posterior limit of the aperture. There are about 3 whorls
present, but the columella is considerably shorter. A small
columellar muscle scar is visible within the aperture in
a position similar to that in the male specimen. The
shell is very narrowly umbilicate, while in the males
there is no umbilical opening. The shell of the female
from Anjouan is much the same as that of the male,
but larger.
Protoconch: The protoconch in the two male specimens
available for study is unsculptured, smoothly rounded
and hardly raised, consisting of about 14 whorls. There
is a sharp change in sculpture between protoconch and
adult shell, the latter being defined by the beginning of
the fine growth lines (Plate 55, Figures D, G). The
female protoconch differs only in being partly covered
by the adult shell, a phenomenon possibly related to
sexual dimorphism.

GEOGRAPHICAL DISTRIBUTION

Distributional data accumulated to date indicate a
rather extensive range for Caledoniella montrouziert
from the western Indian Ocean through the East Indies
to western Polynesia in the tropical Indo-Pacific. Until
recently the gastropod was reported only on Gonodacty-
lus chiragra or Gonodactylus “species.” MANNING
(1968) has distinguished two additional host species. It
is apparent, therefore, that the relationship is not specific
so far as the species of Gonodactylus is concerned.

Locality Records: Persian Gulf (Preston, 1912; on
Gonodactylus chiragra (Fasricius) ). Comoro Islands:
Anjouan Island (Manninc, 1968; on G. platysoma
Woop-Mason). Madagascar: Tulear (MANNING, 1968;
on G. smithu Pocock). Andaman Islands (Preston,
1912; on G. chiragra). Australia: Albany Passage, Cape
York, Queensland (ALLAN, 1936; Gonodactylus sp.). In-

Page 348

donesia: Amboina (Hottuuts, 1941; G. chiragra). New
Caledonia: Art Island (Souversiz, 1869; SouVERBIE &
Montrouzier, 1870; Gonodactylus sp.). Samoa: Mata-
pao, Savaii (HottHuts, 1951; G. chiragra).

REPRODUCTION

Yonce (1953, 1960) has discussed the problems asso-
ciated with reproduction in a sedentary gastropod. He
demonstrated how in Hipponix antiquatus, a species
which lives permanently attached to hard substrates, and
in which protandric hermaphroditism is the apparent
condition, fertilization is probably effected by individuals
of the male phase extending the long penis to the mantle
cavity and oviduct of an adjacent female. YoNGE sug-
gested that, although sedentary, these gastropods oc-
curred in sufficient density to make feasible such an ar-
rangement for the adequate perpetuation of the species.

Available data indicate that Caledoniella montrouziert
always lives on Gonodactylus; at least none has been
reported in any other habitat. Information concerning
placement of the gastropod on its host seems to indicate
a trend. More than one Caledoniella per Gonodactylus
has been reported in the literature 4 times: (1) by
ALLAN (1936), (2) by HottuHurs (1941, 1951 same
case), and (3, 4) by Mannine (1968, specimens de-
scribed and figured here). In three of these cases the
larger of the two gastropods, here believed to be female,
was located near the ventral posterior end of the crusta-
ceans’ abdomen between the pleopods; the smaller gas-
tropod, believed to be male, was always located near the
ventral posterior end of the thorax, between the perio-
pods. In the fourth case, Mannine (1968) found two
gastropods between the pleopods of G. platysoma. Of
the last two specimens reported, only one, a male (Plate
55, Figures A to F), was transmitted to me for study
initially. It was located on the Gonodactylus on the poste-
rior surface of the first right pleopod, and the place of
attachment was later noted to be marked on the pleopod
by a mucoid deposit with the outline of the foot. The
second specimen was not removed from the Gonodactylus
and was not received for examination by me until much
of the present paper had been written. It caused a
change in certain of the concepts I had formed con-
cerning the permanency of attachment to the host.

The specimen, a female, was located between the
fourth and fifth pleopods on the right side of the ventral
posterior abdomen of Gonodactylus platysoma. It did
not appear to be attached by its foot to the host, but
rather was very firmly grasping a single gill filament of
the Gonodactylus between the lobes of its propodium. It

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Vol. 11; No. 4

is probable that the foot attachment was disturbed
during collection or preservation. The animal probably
was attached by its foot to the broad anterior surface of
the fifth pleopod.

In all cases reported, therefore, of a male and female

occurring together on Gonodactylus, the male shows a
tendency to be located on the posterior thorax or in the
last case, the anterior abdomen. The females have been
found on the posterior abdomen. The following infor-
mation concerning distribution of egg-masses indicates
that it is most probable that the female, at least, moves
about and is not sessile.
Egg-Masses: HottHuis (1941) mentioned and figured
egg-masses, supposedly those of Caledoniella montrouzi-
ert, on the abdominal pleopods of specimens of Gono-
dactylus chiragra from the Moluccas and from Samoa
(see Text figure 4, from Hottuuis, 1941). The egg-
masses are similar to those described and figured by
YonceE (1953, fig. 8) in Hipponix antiquatus except that
C. montrouzieri deposits its masses outside of its own
mantle cavity. Eleven masses are shown by HoLTHutIs
to be attached over an area comprising about 3 abdom-
inal segments.

The specimen of Gonodactylus platysoma from Anjou-
an Island had distributed over its abdominal segments a
total of 35 egg-masses. These were attached to the more
medial gill filaments of all but the first pair of pleopods.
Their distribution was as follows:

Left Right
Pleopod Number 1 0 0
Pleopod Number 2 3 4
Pleopod Number 3 3 d
Pleopod Number 4 6 7
Pleopod Number 5 4 1
Total 16 IG) = 3)

Eggs within the capsules were in all stages of develop-
ment. Some capsules contained material having no defi-
nite form, others contained fully formed young snails
whose shells, consisting of about 1 whorl, resembled the
protoconchs of the adults; several of the capsules were
empty and evidenced terminal apertures through which
the young may have escaped. Considering the advanced
stage of development of the young in several of the
capsules it is almost certain that when hatched from the
capsules they are already at the crawling stage. The
large number of capsules, together with their widespread
placement on the host, would indicate the female moves
about over the surface of the pleopods during their depo-
sition. It is most probable that fertilization takes place
when the female approaches the position of the male in

Page 348a THE VELIGER Vol. 11; No. 4

Through an oversight, the explanations to the Text figures

and the Plate figures were omitted in our April issue of

Volume 11. We apologize to the author, Dr. Joseph
Rosewater, as well as to our readers.

Explanation of the Text figures
(see page 346)

Figure 1: Caledoniella montrouzieri on ventral side of
Gonodactylus sp. from Albany Passage, Cape York,
Queensland; note supposed male on thorax, female on
posterior abdomen (from ALLAN, 1936, plt. 26, fig. 1).
Figures 2 and 3: Vanikoro cancellata LAMARCK; two views
of animal; Figure 2, animal in aperture of shell, showing
circular foot, propodium, epipodial fringe, operculum,
proboscis, tentacles and eyes; Figure 3, animal withdrawn
from shell and viewed from side (from Quoy & GammarD,
Atlas, 1833, plt. 66 (bis), figs. 20, 21).

Figure 4: Egg-masses of Caledoniella montrouzieri on
abdominal appendages of Gonodactylus chiragra (from
Ho.ituurs, 1941, fig. 7).

Explanation of Plate 55

Figures A to F: Caledoniella montrouziert, male from Figure D: Enlarged view of protoconch.

Anjouan Island, Comoros Islands. Figures E and F: Two views of shell.

Figures A to C: Front and side views showing propodium, Figures G to I: Shell of male from Tulear, Madagascar.
foot, “epipodial fringe”, proboscis, tentacles and penis. Figures J to L: Shell of female from same locality.

(scale represents 1 mm; detail squares are 0.75 mm)

a

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[Rosewater] Plate 55

Vol. 11; No. 4

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Page 349

its location anterior on the host, although it has not yet
been proven satisfactorily that the male is completely
sedentary.

As mentioned by YoncE for Hipponix, the propodium,
well developed in Caledoniella montrouzern, probably
plays an active role in egg-mass formation and deposition.

It was mentioned above that the female specimen of
Caledoniella montrouziert which was found in situ on
Gonodactylus platysoma was grasping between the lobes
of its propodium a single gill filament of the stomato-
pod. All of the 35 egg-capsules observed on the stomato-
pod were fastened to it by means of the capsules having
incorporated along one edge | or 2 gill filaments. When
the propodium was examined, its “pouch” was found to
be lined with material very much resembling that which
forms the egg-capsules. These items of circumstantial
evidence leave the strong impression that the propodium
is directly involved in egg-mass formation, that its in-
trinsic secretions form the egg-capsule around eggs trans-
ported from the female gonoduct, and that the lobes of
the propodium also manipulate the Gonodactylus’ gill
filaments and incorporate them into the capsules. The
groove running from the base of the foot to the propodi-
um also appears to enter into this process.

FEEDING

Little real information is available regarding the feeding
habits of Caledoniella montrouzieri except in a negative
sense. Ctenidia do not appear to be developed for ciliary
feeding, nor were a radula or jaws found, contrary to
THIELE’s (1929) description. A hint to the possible food
of this species was extrapolated from observations made
during the examination of the female specimen attached
to Gonodactylus platysoma (see Reproduction). It was
noted that numerous gill filaments were damaged at
their tips and showed evidence of subsequent repair.
Damage may have resulted from other causes or have
been caused during egg-laying, although filaments were
involved other than those to which capsules were at-
tached. It is possible that the gastropod manipulates the
gill filaments with the propodium (a well-developed pro-
podium is also present in males), removes the tips of the
filaments (just how this is done would be difficult to
explain in the apparent absence of jaws or a radula;
perhaps by chemical means), and “sucks” the body fluids
of the host with its well-developed proboscis. If this is
the case, the stomatopod-gastropod relationship should be
termed a host-parasite one rather than a commensal one.
If this theory proves invalid, the species also may feed
on detritus or remnants of the food of its host. Direct

observation of the living animals is needed to settle this
question.

CLASSIFICATION

The previous assignments to family group of Caledoni-
ella montrouziert by Tryon and THIELE (Lamellariinae)
and by Wenz (Stiliferidae) are considered erroneous. In
spite of the lack of a radula in present material, resemb-
lance of the animal features to both Vanikoro and Hip-
ponix, and of egg-laying habits to Hipponix, suggest
placement in the Superfamily Hipponicacea (see TayLor
& SoHL, 1962). As Caledoniella does not appear clearly
to belong in either the Vanikoridae or Hipponicidae,
however, a new family is proposed here:

Family Caledoniellidae, new family

Type Genus: Caledoniella Souversie, 1869, monotypic.
Diagnosis: The family concept is based on the character-
istics of the type genus, in turn essentially those of its
type species Caledoniella montrouzieri.

Further study may uncover evidence which will change
this concept, but at present this placement seems to be
the most logical for the species. The chance that Cale-
doniellidae belongs in Calyptraeacea seems more remote
as animal resemblances are not so great (YonGE, 1938).

SUMMARY

Caledoniella montrouzieri is a reduced-spirally-coiled
mesogastropod which lives on species of the stomatopod
crustacean Gonodactylus throughout the tropical Indo-
Pacific.

The animal is adapted for its commensal existence by
possessing a “sucker-like” foot. A fringe-like metapodium
probably partially covers over the shell in life. A well-
developed propodium below the proboscis serves in the
formation and attachment of egg-masses and probably
is involved in feeding activities.

Placement of male gastropods on terminal thoracic or
anterior abdominal and of females on posterior abdom-
inal segments of Gonodactylus indicates a possible pref
erence of these loci as habitats, although the female, at
least, moves about on the abdomen of the host during
egg-laying and probably feeding activities.

The family Caledoniellidae is proposed for Caledoni-
ella montrouzieri and the group is assigned to the super-
family Hipponicacea with which it shows strongest rela-
tionship.

Page 350

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Vol. 11; No. 4

LITERATURE CITED

ALLAN, JOYCE

1936. Mysticoncha, a new generic name for Caledoniella Ba-
SEDOW, non SOUVERBIE. Rec. Austral. Mus. 19: 391 - 396;
pit. 26

BaseEpow, H.

1905. On naticoid genera Lamellaria and Caledoniella from
South Australia. Trans. Proc. Roy. Soc. South Austral. 29:
181 - 186; plts. 26 - 29

Bercy, Lupwic SopHus RUDOLF

1896. Beitrag zur Kenntnis der Gattungen Narica und Onustus.
Verhandl. k.-k. zool.-bot. Gesellsch. Wien, 1896: 1-14 [re-
print]; plts. 2-3

FRETTER, VERA & ALASTAIR GRAHAM

1962. _ British prosobranch molluscs, their functional anatomy

and ecology London, Ray Soc. xvit+ 755 pp.; 317 figs.
Hottuuts, L. B.

1941. | The Stomatopoda of the Snellius Expedition. Biological
results of the Snellius Expedition, XII. Temminckia,
Journ. Syst. Zool. 6: 241 - 294; figs. 1-9

1951. Note on Caledoniella montrouziert SOUVERBIE, a gastro-
pod mollusc living commensally on stomatopod Crustacea.
Basteria 15: 69-71

Mannine, R. B.

1968. Stomatopod Crustacea from Madagascar. Proc. U.
S. Nat. Mus. 124 (3641): 1-61; figs. 1-16
Preston, H. B.
1912. On a new genus and species of marine parasitic Gastro-

poda from the Indian region. Jn: N. ANNANDALE, Fauna
Symbiotica Indica No. 2. Rec. Indian Mus. 7: 126 - 127;
figs.

Quoy, JEAN RENE ConsTANr & JoSEPH Paut GarmarpD

1832. Voyage de découvertes de l’Astrolabe, Zoology 2 (1): 1
to 320; Atlas (1833): plt. 66 (bis), figs. 20 - 22

Rosertson, RoBERT
1962. Vanikoro Quoy & Gatmarp, 1832 (Mollusca, Gastro-
poda) ; proposed validation under the Plenary Powers. Z.
N. (S) 1524. Bull. Zool. Nomenclat. 19: 332 - 336
SOUVERBIE, S. M.
1869. Diagnoses de mollusques inédits provenant de la Nou-
velle-Calédonie. Journ. Conchyl. 17: 416 - 421
SouversiE, S. M. « R. P MontTROUZIER
1870. _ Descriptions d’espéces nouvelles de I’archipel calédonien.
Journ. Conchyl. 18: 71 - 83; plt. 9
Taytor, DwicHT WILLARD « NorMAN SOHL
1962. An outline of gastropod classification.
1: 7-32
THIELE, JOHANNES
1929. Handbuch der systematischen Weichtierkunde. Erster
Teil, Loricata; Gastropoda. I: Prosobranchia (Vorderkiemer).
pp. 1-376; 470 text figs.
Tryon, Georce WASHINGTON, Jr.
1886. Naticidae, Calyptraeidac, etc. In: Manual of Con-
chology, structural and systematic 8: 1 - 461; plts. 1-79
WENZ, WILHELM
1940. Handbuch der Palaozoologic 6, prt. 1 (6): 721 - 960;
figs. 2084 - 2787
YonGcE, CHARLES MAurRICE

Malacologia

Jena, Gustav Fischer Verlag

1938. Evolution of ciliary feeding in the prosobranchia, with
an account of feeding in Capulus ungaricus.
Biol. Assoc. U. K. 22: 453 - 468; figs. 1-6

1953. | Observations on Hipponix antiquatus (LINNAEUS).
Proc. Calif. Acad. Sci. ser. 4, 28 (1): 1-24; 9 figs.

(15 July 1953)

1960. Further observations on Hipponix antiquatus with notes

Proc. Calif. Acad.

Journ. Marine

on North Pacific pulmonate limpets.
Sci., ser. 4, 31: 111-119; figs. 1-4

——=.

Vol. 11; No. 4

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Page 351

A Preliminary Survey of Mollusks for Consag Rock

and Adjacent Areas, Gulf of California, Mexico

HELEN DuSHANE!

AND

ELLEN BRENNAN 2

(1 Map)

INTRODUCTION

Consac Rock, GuLF oF CALIFORNIA, Mexico is at Lati-
tude 31°07’N, Longitude 114°27’W in the extreme upper
reaches of the Gulf of California, east by north of San
Felipe Bay, offshore approximately 20 miles. As far as
we have been able to determine, no extensive collecting
of mollusks has been done in the waters surrounding Con-
sag Rock. For this reason we consider it desirable to
publish the results of a 3-day trawling expedition in this
and the immediately adjacent areas.

The list of mollusks which follows is based on the
results of a joint trip made by the authors and other indi-
viduals on June 27, 28, and 29, 1968 during which 217
species were collected. The trip was organized primarily
for members of the Conchological Club of Southern Cali-
fornia by Ellen Brennan. The following persons have made
their collections from this trip available to us: Twila
Bratcher, Don Cadien, Billee Dilworth, Joseph DuShane,
William and Joyce Gemmell, Roy Poorman, William E.
Viney, Erwin and Gertrude Wahrenbrock.

The nudibranch fauna of the Panamic province is only
recently becoming better known through the efforts of
FarMER (1963, 1966, 1967), Lance (1961, 1966, 1968),
and Marcus (1967). Therefore, a determined effort was
made by Don Cadien of the Los Angeles County Museum
of Natural History to collect representatives of this group
of animals and to transport them back alive.

Some of the other areas in the Gulf of California have
been rather extensively explored over a period of years

* 15012 El Soneto Drive, Whittier, California 90605
2 9636 La Cima Drive, Whittier, California 90603

by the California Academy of Sciences (1888 - 1921),
the Velero III of the Allan Hancock Foundation (1931 to
1941), and the Puritan - American Museum of Natural
History Expedition (1958). Poorman dredged mollusks
over a period of years (1961 - 1967) in the areas of San
Carlos and Bacochibampo Bays, Guaymas, Sonora, Mexi-
co. The specimens collected have been reported (Du-
SHANE & Poorman, 1967).

PREVIOUS COLLECTING
IN THE VICINITY or CONSAG ROCK

Historically, the Jesuit, Father Fernando Consag was
among the first to make a reconnaissance trip in 1746 to
the extreme northern reaches of the Gulf of California
(ENGELHARDT, 1929: p. 266). The rock named for him
is 289 feet high. A manuscript map by Consag (AppING-
Ton MSS), with notations in his own handwriting, states
that at Santa Isabel, “hasta aqui llegan los placeros de
perlas” (thus far extend the pearl grounds). Santa Isa-
bel was at a point on the east coast of peninsular Lower
California about where Puertecitos is today.

Two records of molluscan collecting in the vicinity of
Consag Rock appear in the literature: (1) The Allan
Hancock Pacific Expeditions of 1937 and 1940 collected
on and around Consag Rock and in San Felipe Bay.
Fraser (1943) listed 6 dredging stations and 2 shore
stations, but the mollusks remain largely unreported.
(2) The 1957 Puritan - American Museum of Natural
History Expedition reported 2 dredging stations 14 miles
SW of Consag Rock. A general account of this expedition
was given by Emerson (1958).

Page 352

OCEANOGRAPHIC CONSIDERATIONS

Very little is known concerning meteorological and oce-
anographic conditions at San Felipe and Consag Rock.
Since the Jesuit missionaries established no mission at
San Felipe we must rely on other, more recent, reports
for information on air and water temperatures. No year
around water temperature data are available (RoDEN
& Groves 1959: p. 11). The air temperature in the
northeastern section of the peninsula shows extremes from
18° F in winter to 128° F in summer (NeEtson, 1922: p.
102), making this the hottest and driest part of Lower
California. Occasionally violent storms come up the Gulf
of California from the south or are driven over the high
Peninsular Range from the Pacific ocean. The most recent
devastating storm occurred at San Felipe on September 1,
1967. Summer rains are capricious and irregular; one
place may be deluged while another close by may not
receive a single drop. These rains are commonly in the
form of cloudbursts, usually in July, August, or September
with resultant heavy runoff into the western edge of the
Gulf of California. Winter rains are also irregular and
sometimes last only a few hours. Some years no winter
rains fall. Average yearly rainfall is about 2.5 inches
(Nelson, 1922: pp. 96, 98, 99).

One cannot discuss oceanographic conditions in this
area without due consideration of the effects of the
Colorado River upon the waters of the Gulf of Califor-
nia. Prior to 1938, when Boulder Dam was completed, the
Colorado River watershed annually discharged billions
of tons of silt into the waters of the Gulf. Since the
tidal current in the Gulf is counterclockwise (BERRY,
1954: p. 24; Ronen, 1958: pp. 24, 33) this detrital
material had been carried from the Colorado River as
far south as San Felipe Point, an isolated volcanic part
of the Peninsular Range on the western shore of the Gulf.
As a result of this deluge of silt there are many pockets
of mud in the northern end of this body of water.
According to Sykes (1937: p. 107), “Driftwood, plainly
of Colorado River origin, has been observed as far to
the southward as the San Luis Islands, along the Lower
California shore (latitude 30° N), and in this case the
transporting agency was probably tidal current rather
than wind.” In the literature no such phenomenon has
been reported on the Sonoran coast. With the comple-
tion of Boulder Dam and the further construction of an
impounding and diversion dam at Parker, the movement
of fresh detrital material is practically stopped (SyKEs,
IQB/e jee, WS)

San Felipe Bay is a shallow depression on the west side
of the Gulf, with a sandy bottom and a tidal range of
about 22 feet (7 m). The sea floor outside the bay slopes

THE VELIGER

Vol. 11; No. 4

very gradually to a depth of 27 fms (48 m) beyond Con-
sag Rock at Lat. 31°12’N, Long. 114°22’W with many
mud holes and sandy ridges present. Extensive evapora-
tion, which increases salinity, occurs on the shallow pro-
tected bays of San Felipe as well as at Adair Bay on the
Sonoran side and Concepcion Bay on the Lower Cali-
fornia side (RopEN & Groves, 1959: p. 16). In the 30
years since the completion of Boulder and Parker Dams
the silt, being more dense than the current-driven sand,
still lies in pockets on the ocean floor. The current U. S.
Navy Oceanographic map (Chart 620) shows these
deposits.

FAUNAL RELATIONSHIPS

The following lists record 217 species of mollusks, of
which 58 are pelecypods, 5 are scaphopods, and 154
are gastropods. One is doubtfully identified (“cf”) and
7 are identified only to genus.

Because, to us, there are no known records of mollusks
collected this far north in the Gulf of California we con-
sider all specimens collected to represent northern exten-
sions of the known range with the exceptions of Callio-
stoma palmeri, Crepidula arenata, Crepidula incurva,
Terebra glauca, Terebra armillata, Nassarius iodes, and
Nassarius moestus which the senior author collected inter-
tidally in November 1967 at El Gulfo, Sonora, Mexico.

Although the molluscan fauna is predominantly Pan-
amic, some members of the Californian province are
represented in the northern Gulf. Specimens of the fol-
lowing species occurring in both the Californian and Pan-
amic provinces are also to be found in trawling at Consag
Rock and in the immediate vicinity of San Felipe Bay:
Nucula linki Dau, Hiatella arctica (LINNAEUS), Aesopus
chrysalloides CARPENTER, and Iselica fenestrata CARPEN-
TER.

The faunal element restricted to the northern and
northwestern shores is less well known but includes: Ac-
maea strongiana, Nomaeopelta dalliana, Cantharus mac-
rospira, Turritella anactor, Terebra berryi, Terebra du-
shanae, Recluzia palmeri and Melampus mousleyi. Speci-
mens of Nassarina pammicra reported by McLean
(1961) from Los Angeles Bay as a range extension
northward from Nicaragua have also been collected at
Gonzaga Bay by DUSHANE & SpHON (1968) as well as
at Puertecitos by DuSHANE (1964). Terebra berry: and
Terebra dushanae seem to have a very limited distribution
on the northwestern shores of the Gulf of California
(type locality for both: Puertecitos). The former has not
been found living at San Felipe but occurs sparingly at
Gonzaga Bay to the south. The latter species occurs inter-

Vol. 11; No. 4

THE VELIGER

nnn nn eee UNE EEUE SEIS SNE SEE URE SEEED UE

tidally uncommonly at Agua Chale, 24 miles south of
San Felipe. Terebra variegata, well known throughout the
Gulf, is missing from the San Felipe fauna, but was
trawled off Agua Chale. Strombina dorsata occurs at
both Puertecitos and Gonzaga Bay but is unrecorded at
Los Angeles Bay. It occurs uncommonly at San Felipe
Bay. Nassarina anitae, described by Campsett (1961)
from Guaymas, occurs uncommonly on clumps of Pteria
sterna in the vicinity of Consag Rock. Specimens collected
are essentially the same as the ones ParKER (1963: p. 167)
reported from 11 to 26 meters in the northern Gulf.

The more unusual extensions of range northward are
represented by specimens including: Nucula linki, Diplo-
thyra curta, Ostrea megodon, Lophocardium cumingi,
Psephidia cymata, Macoma undulata, Aequipecten pal-
meri, Iselica fenestrata, Clathrodrillia adonis, Crassispira
bacchia, Clathurella acapulcana, Vitularia salebrosa, and
Conus tornatus.

Allyn G. Smith (personal communication) reports that
a new species of Fusinus was taken in the area of Consag
Rock. Another species of Fusinus awaits determination
after a comparison of the radula with intertidal speci-
mens of Fusinus ambustus.

An unusual occurrence of an arthropod taken merits
a note: Eupagurus varians BENeEbIcT, together with its
commensal hydractinian Janaria mirabilis StTECHOW
(Treatise Inv. Paleo.: p. 84) was common at stations 7,
8, and 13.

SYSTEMATIC ACCOUNT

The following format is adopted:

1. The order in the checklist, the nomenclature, and the
species numbers are those given by KEEN (1958) with
a few changes as new material was discovered. Re-
ferences to species listed by KEEN may be located in
her bibliography. References to species described since
1958 are included in the present paper.

2. The habitat and relative abundance of species taken
are given. Indications are made for those species not
taken alive.

3. The collecting stations referred to in the list by num-
bers are shown on the accompanying map. All species
reported are from depths of from 3 to 21 fms (5 to
38m).

4. Unusual range extensions are indicated by an asterisk
(*) following the “Keen numbers.” The area from which
the range is extended follows the collector’s initial.

5. The following collectors are designated by initials:

Page 353
Twila Bratcher B
Ellen Brennan Br
Don Cadien Cc
Billee Dilworth D
Joseph « Helen DuShane Du
William & Joyce Gemmell G
Roy Poorman P
William E. Viney Vv
Erwin & Gertrude Wahrenbrock W

The specimens reported are in the private collections of
the individuals named and the Cadien collection is in the
Los Angeles County Museum of Natural History.

ACKNOWLEDGMENTS

We wish to express our gratitude to Dr. A. Myra Keen
and to Roy Poorman for identification of many of the
small species; to Dr. James H. McLean of the Los Angeles
County Museum of Natural History for critical evalua-
tion of the manuscript; to Don Cadien of the same
institution for identification of the nudibranchs.

ECOLOGICAL NOTES
on 16 COLLECTING STATIONS

(see Map)

Over a three-day period mollusks were trawled at the

following 16 stations. After an initial pull at 3 mph, which

proved to be too rapid, all pulls were made at 2 mph, the
minimum speed of the boat. in the notes below, longitude
and latitude are given for the beginning of each pull.

1. Two miles off Punta Diggs, 12 miles S of the town of
San Felipe; Lat. 30°51’12” N, Long. 114°39’ W; course
160°. The nets were down for 40 minutes over sand
bottom at a depth of 5 fm. Fish and sea stars comprised
the bulk of the material trawled. The stomachs of one
species of sea star yielded many mollusks, among them
species of Nuculana, Pitar, Pandora, Olivella and Acte-
ocina.

2. Two miles offshore from Alicia Playa, 18 miles S of the
town of San Felipe; Lat. 30°46’N, Long. 114°40’W;
course 130°. The nets trawled over sand bottom at a
depth of 6 to 7 fm. A great number of sea stars were
included in this haul and their stomachs yielded species
of Crassinella, Trigoniocardia, Nassarius, and Olivella,
along with Chione mariae and juvenile Cosmioconcha
palmeri.

Page BAe THE VELIGER

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Gulf of California

Map showing dredging stations
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Vol. 11; No. 4

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Vol. 11; No. 4

3.

Four rope tangles were set south of Alicia Playa, 18
miles S of the town of San Felipe, Lat. 30°44’30” N,
Long. 114°41’W, in 3 fm of water over rocks. They
were baited with decaying fish contained in nylon bags.
The tangles were left in the water about 3 hours and
when retrieved, the bait was gone. Quick examination
on the spot revealed no mollusks; however, later careful
examination revealed several specimens of Anachis va-
ria, A. milium, and a juvenile Muricanthus nigritus
which had crawled high into the untwisted strands of
the tangle. If a way could be devised to keep the bait
in the tangles, this could prove to be a good collecting
device.

4. Two miles offshore from Agua Chale, 24 miles S of

=

the town of San Felipe; Lat. 30°41’ N, Long. 114°40’06”
W;; course 195°. This pull covered one mile of sand
bottom at a depth of 5 fm paralleling an offshore rock
reef. It yielded fish, crustaceans, and 2 species of sea
stars. Examination of the stomachs of these echino-
derms produced several species of Nuculana, Pandora
brevifrons, Calliostoma palmeri, and Eupleura murici-
formis.

Eight miles E of Agua Chale, 24 miles S of San Fe-
lipe; Lat. 30°41’N, Long. 114°32’W; course 340°.
This one-mile pull over sand and mud pockets at a
depth of 10 fm yielded a large quantity of fish and 2
species of sea stars, their stomachs containing species
of Nuculana, juvenile Trachycardium senticosum, Chi-
one mariae, C. gnidia, and C. pulicaria.

. Two miles SW of Consag Rock, Lat. 31°06’ N, Long.

114°30’30” W; course 095°. On this pull and all sub-
sequent ones, more weight was added to the nets, pro-
ducing better results. The nets trawled over one mile of
sand bottom at a depth of 12 fm, producing quantities
of Hexaplex erythrostomus, Muricanthus nigritus with
eggs. Calyptraea, Crepidula, and Crucibulum were com-
mon attached to dead shells.

Two miles S of Consag Rock, Lat. 31°05’ N, Long.
114°28’30” W; course 100°. On this pull the nets
trawled over ? of a mile of sand bottom at a depth of
11 fm. Among the mollusks trawled were Calliostoma
palmeri, Ficus ventricosa, Murex elenensis, M. recurvi-
rostris lividus, Pteropurpura erinaceoides, Acanthina
tuberculata, and Cancellaria cassidiformis. Basket stars
of the genus C’rinoidea were common as were sponges,
red-brown in color, each growing upon and completely
enveloping a shell.

Eight miles ENE of Consag Rock, Lat. 31°08’30” N,
Long. 114°21’W; course 340°. A one mile pull over
sand bottom at a depth of 18 fm produced many

THE VELIGER Page 355

clumps of living Pteria sterna among the lamellae of
which were Septifer zeteki, Hiatella arctica, Epitonium
keratium, Iselica fenestrata, and Nassarina anitae. Vitu-
laria salebrosa was taken living on dead P. sterna shells,
as was Modiolus capax.

9. Seven miles ENE of Consag Rock, Lat. 31°08’ N, Long.

114°21’W; course 020°. A one mile pull over sand
bottom at 15 fm depth produced approximately the
same mollusk species as were trawled at Station 8, with
the addition of Lioberus splendida and Atrina tuber-
culosa.

10. Seventeen miles NW of Consag Rock, Lat. 31°20’18”
N, Long. 114°41’W. After a one mile pull at a depth
of 14 fm over mud bottom, the nets were full of
Astropecten. Examination of the stomachs of the sea
stars produced such mollusks as Nucula linki, Natica
broderipiana, and Strombina dorsata.

11. Three-fourths of a mile offshore, beginning north of
San Felipe Point and ending south of San Felipe Point;
Lat. 31°02’30” N, Long. 114°48’ W. This pull was 2 of
a mile long, over sand and mud bottom at a depth of
6 fm. The nets yielded a coral-related material con-
taining Diplothyra curta and Lithophaga attenuata
rogersi. A large mass of aborted egg strings was found
to contain a multitude of small shells. Among the gen-
era represented were Nucula, Nuculana, Balcis, Niso,
Epitonium, Cyclostremiscus, Seila, Aesopus, Nassarina,
Anachis, Clavus, Clathrodrillia, and Mangelia.

12. Three miles SW of Consag Rock, Lat. 31°04’30” N,
Long. 114°31’30” W; course 080°. This was a one mile
pull over sand bottom at a depth of 13 fm. The otter
trawl brought up Polinices intemeratus, Hexaplex eryth-
rostomus, Muricanthus nigritus with eggs, and Soleno-
steira capitaneus.

13. Five miles SSE of Consag Rock, Lat. 31°02’12”N,
114°26’24” W; course 070°. A one mile pull over sand
bottom at 11 fm depth produced such mollusks as Ficus
ventricosa, Murex recurvirostris lividus, Pteropurpura
erinaceoides, Cancellaria cassidiformis, and Conus poor-
mani.

14. Eight miles NE of Consag Rock, Lat. 31°12’ N, Long.
114°22’ W; course 080°. A 2 mile pull over sand and
clay bottom at a depth of 21 fm produced essentially
the same species as were trawled at Stations 8 and 9.

15. Ten miles ENE of Consag Rock, Lat. 31°09’12” N,
Long. 114°17’36” W; course 314°. This was a one mile
pull over sand and clay bottom at 16 fm depth. The
trawl was full of clumps of Pteria sterna attached to
which were Ostrea conchaphila, O. megodon, O. pal-

Page 356

mula, Chama sordida, C. buddiana, and Nassarina pam-
micra.

16. Five miles S of Consag Rock, Lat. 31°03’N, Long.
114°31’30” W; course 270°. This one mile pull over
sand bottom at 11 fm depth produced essentially the
same mollusk species as were trawled at Station 13.

3%

10

12

13

87

9la

101

106

107

tts

ING)

123

124

126

PELECYPODA

Nucula declivis Hinps, 1843. Uncommon (10, 11),
6 - 14 fm, mud and sand bottom, in sea star stom-
achs and entangled in aborted egg mass (B, C, D,
Ee IP ))e

Nucula linki Day, 1916. Rare (10), 14 fm, mud
and sand bottom, from sea star stomachs; Pt. Fer-
min, Gulf of California (Du, G).

Nuculana elenensis (SowERBy, 1833). Uncommon
(11), 6 fm, sand and mud bottom, entangled in ab-
orted egg mass (Br, C, Du).

Nuculana impar (Pitssry & Lowe, 1932). Common
(1, 4, 5, 10, 15), 5-16 fm; sand, mud, and clay
bottom; in sea star stomachs (B, Br, C, D, G).
Nuculana laeviradius (Pitspry & Lowe, 1932). Com-
mon (4, 5, 9, 10, 14), 5 - 21 fm; sand, mud, and clay
bottom; in sea star stomachs (B, C, D, Du, G, P).
Septifer zeteki HERTLEIN & STRONG, 1946. Rare (8),
18 fm, sand bottom (C, Du).

Lithophaga attenuata rogerst Berry, 1957. Uncom-
mon (11), 6 fm, in chunks of coral-related material

(C, W).

Modiolus capax (Conrab, 1837). Rare (8), 18 fm,
attached to Pteria sterna (Du).

Lioberus splendida DuNKER, 1857. Uncommon (7,
14), 11-21 fm, sand bottom and on Pteria sterna
(Br, Du, G).

Pteria sterna (Goutp, 1851). Common (8, 9, 14, 15),
15 - 21 fm, sand and clay bottom (B, Br, C, D, Du,
G, P, V, W).

Atrina tuberculosa (SowERBy, 1835). Rare (9), 15
fm, sand bottom (C).

Ostrea conchaphila CarPENTER, 1857. Uncommon
(9, 10, 14, 15), 15 - 21 fm, sand and clay bottom, on
dead Pteria sterna shells (Du).

Ostrea megodon Han ey, 1846. Rare (15), 16 fm,
sand and clay bottom (D).

Ostrea palmula CarPENTER, 1857. Uncommon (9,
10, 14, 15), 15-21 fm, sand and clay bottom, on
Pteria sterna (Du).

Pecten vogdesi ARNOLD, 1906. Uncommon as valves
and dead specimens (8, 13), 11- 18 fm, sand bottom
(Br).

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128

IZ

147

159

231

238

241

251

256

262

263

264

265

282

286

287

296

306

Vol. 11; No. 4

Aequipecten palmeri (Dati, 1897). Common (7,
8, 9, 13, 16), 11-18 fm, sand bottom (Br, C, Du,
G, P).

Aequipecten circularis (SowERBY, 1835). Common
(6, 7, 8, 9, 12, 13, 14, 15, 16), 11-21 fm, sand and
clay bottom; attached to Pteria sterna, gastropods,
and in sea star stomachs (Br, C, Du, G, P).

Plicatula anomioides Kren, 1958. Uncommon (15),
16 fm, sand and clay bottom, on dead Pteria sterna
shells (Du).

Crassinella pacifica (C.B. ADAMs, 1852). Uncom-
mon (2, 11), 6 fm, sand and mud bottom, in sea star
stomachs and entangled in aborted egg mass (B, Br,
C, D, Du, P).

Mysella compressa (Dat, 1913). Rare (9), 15 fm,
sand bottom, in sea star stomachs (P).

Chama buddiana C.B. Apams, 1852. Uncommon
(8), 18 fm, sand bottom, juveniles attached to dead
shells (C, Du).

Chama sordida Broperip, 1835. Common (8, 9,
14, 15), 15-21 fm, sand and clay bottom, attached
to Pteria sterna (Br, C, Du, G).

Trachycardium senticosum (SowErBy, 1833). Com-
mon (5, 16), 10-11 fm, sand bottom, juveniles in
sea star stomachs (Br, C, Du, P).

Trigoniocardia granifera (BRODERIP & SOWERBY,
1829). Uncommon (2, 4), 5 - 7 fm, sand bottom, ju-
veniles in sea star stomachs (C, Du, G, P).

Laevicardium elatum (SoweErsy, 1833). Rare (6),
juvenile, 12 fm, sand bottom (C).

Laevicardium elenense (Sowersy, 1840 [?1841]).
Uncommon (11, 12), 6- 13 fm, sand and mud bot-
tom, juveniles in crevices of coral-related material
and in sea star stomachs (Br, C, G, P).

Lophocardium annettae (DA.u, 1889). Rare (14),
valve only, 21 fm, sand and clay bottom (Br).

Lophocardium cumingu (BropErip, 1833). Rare
(9), valve only, 15 fm, sand bottom (Du).

Transennella tantilla (Goup, 1853). Rare (9), 15
fm, sand bottom, in sea star stomachs (P).

Pitar helenae Ousson, 1961. Uncommon (6, 10),
12-14 fm, sand and mud bottom, in sea star stom-
achs (Br, Du).

Pitar perfragilis Pitspry « Lowe, 1932. Rare (4),
5 fm, sand bottom, in sea star stomachs (P).

Pitar concinnus (SoweErBy, 1835). Common (1, 2,
4,5), 5-10 fm, sand and mud bottom, in sea star
stomachs (C, Du, G, P).

Dosinia ponderosa (Gray, 1838). Uncommon (2,
13), 6-11 fm, sand bottom, juveniles in sea star
stomachs (C, Du, P).

Vol. 11; No. 4

315 Psephidia cymata Datu, 1913. Uncommon (10, 14,
15), 14-21 fm, sand and clay bottom, in sea star
stomachs (C, Du, P).

326 Chione gnidia (BRoDERIP & SOWERBY, 1829). Com-
mon, juveniles (1, 2, 4,5), 5-10 fm, sand and mud
bottom, in sea star stomachs (C, Du, G, P).

327. Chione pulicaria (BroperiP, 1835). Uncommon (2,
3, 5), 5- 10 fm, sand and mud bottom, juveniles in
sea star stomachs (Br, C, Du, P).

331 Chione mariae (p’OrpicNny, 1845). Common (2, 4,
5, 10, 13, 16), 5-14 fm, sand and mud bottom, in
sea star stomachs (Br, C, Du, G, P).

335 Chione picta WittETT, 1944. Rare (2), 6-7 fm,
sand bottom, in sea star stomachs (B).

392 Tellina amianta Da.t, 1900. Common (10, 14, 15),
14 - 21 fm, mud, sand, and clay bottom; from wash-
ings of Pteria sterna and in sea star stomachs (Br, C,
Dia, Lee

425 Macoma undatella (HANLEy, 1844). Uncommon
(10), 14 fm, mud bottom, in sea star stomachs (Br,
Du) GP).

436 Macoma pacis Pitspry & Lowe, 1932. Uncommon
(4), 5 fm, sand bottom, in sea star stomachs (G).

438 Strigilla cicercula (Puttippt, 1846). Rare (9), 15
fm, sand bottom, in sea star stomachs (G).

441 Strigilla lenticula Puiipri, 1846. Rare (7), 11 fm,
sand bottom, in sea star stomach (P).

450 Donax gracilis HANLEY, 1845. Uncommon (4, 5),
5 - 10 fm, sand and mud bottom, in sea star stomachs
(Br, Du, G).

475 Tagelus politus (CARPENTER, 1857). Uncommon
(11), 6 fm, sand and mud bottom, juveniles en-
tangled in aborted egg mass (C, P).

483 Semele guaymasensis (Pirspry & Lowe, 1932). Un-
common (6), 12 fm, sand bottom, in sea star stomachs
(Duy G; Py:

489 Semele pacifica Dati, 1915. Uncommon (7), 11 fm,
sand bottom, in sea star stomachs (Du, G).

507 Abra tepocana Datu, 1915. Rare (5), 10 fm, sand
and mud bottom, in sea star stomachs (G).

527 Corbula nasuta Sowersy, 1833. Common (6 to 10,
12 to 16), 11 - 21 fm, sand, mud, and clay bottom; in
sea star stomachs and in washings from Pteria sterna
(BY Bre; Dp: Du; GP):

539 Gastrochaena ovata SowErRBy, 1834. Rare, valves
only (11), 6 fm, sand and mud bottom, embedded in
coral-related material (Br).

542 Hiatella arctica (LINNAEUS, 1767). Common (7, 8,
9, 13, 14, 15), 11-21 fm, sand and clay bottom,
nestling on outer edges of Pteria sterna and on Muri-
canthus nigritus (Br, C, Du, G, P).

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Page 357

545 Panope globosa DaLt, 1898. Valve only (13), 11 fm,
sand bottom (C).

553 Diplothyra curta (SowERBy, 1834). Common (11),
6 fm, sand and mud bottom, boring in coral-related
material (Br, C, W).

567 Pandora brevifrons SowErBy, 1835. Common (4, 6,
7, 12, 13, 16), 5-13 fm, sand bottom, in sea star
stomachs (Br, C, G).

569 Pandora claviculata CARPENTER, 1855. Uncommon
(10), 14 fm, mud bottom, in sea star stomachs (Du).

576 Pandora granulata Datu, 1915. Uncommon (1, 2),
5 - 7 fm, sand bottom, in sea star stomachs (Du, P).

591* Asthenothaerus villosior CARPENTER, 1864. Uncom-
mon (10), 14 fm, mud bottom, in sea star stomachs;
Cape San Lucas, Lower California (P).

597 Cuspidaria didyma (Hinps, 1843). Rare (14),
valves only, 21 fm, sand and clay bottom, from sea
star stomachs (Du).

SCAPHOPODA

2 Dentalium inversum DersHayEs, 1826. Uncommon
(14), 21 fm, sand and clay bottom, in sea star stom-
achs (B, D).

3 Dentalium oerstedii MOrcuH, 1860. Common (9, 11,
15), 6-10 fm, sand, clay, and mud bottom; in sea
star stomachs and entangled in aborted egg mass (B,
Br, C, D, Du, P).

5 Dentalium sectum DesuayeEs, 1826. Uncommon (4),
5 fm, sand bottom, in sea star stomachs (G).

9 Dentalium quadrangulare Sowersy, 1832. Uncom-
mon (8), 18 fm, sand bottom, in sea star stomach (G).

12 Cadulus panamensis PitsBry & SHARP, 1897. Com-
mon (1, 2, 8, 10, 11), 5-18 fm, sand and mud bot-
tom, entangled in aborted egg mass, in sea star stom-
achs, and in washings from Pteria sterna (B, Br, C,
D, Du, G, P).

GASTROPODA

30 Diodora alta (C. B. Aas, 1852). Rare (8), 18 fm,
sand bottom, attached to dead shell (C).

45 Calliostoma marshalli Lowe, 1935. Common (2, 4,
8, 9, 14, 15), 18 fm, sand and clay bottom, in sea
star stomachs and in clumps of Pteria sterna (B, C,
D, Du, G).

47 Calliostoma palmeri Dat, 1871. Uncommon (7, 8,
13), 11-18 fm, sand bottom, also in sea star stom-
achs (Br, C, Du, P).

Page 358

60 Solariella triplostephanus Dati, 1910. Rare (11), 6

66

87

99

105

106

107

108

fm, sand and mud bottom, entangled in aborted egg
mass (G).
Turbo mazatlanicus Pitspry & Lowe, 1932. Rare
(4), 5 fm, sand bottom, in sea star stomachs (B, D).
Liotia balboai STRONG & HERTLEIN, 1939. Rare (11),
6 fm, sand and mud bottom, entangled in aborted egg
mass (B, D, G).
Liotia stearnsi DALL, 1918. Rare (2), 6-7 fm, sand
bottom, in sea star stomachs (B, D).
Arene rammata (Datu, 1918). Uncommon (4, 5),
5 - 10 fm, sand and mud bottom, in sea star stomachs
(B, D).
Tricolia equilirata CARPENTER, 1857. Uncommon (9),
15 fm, sand bottom, in sea star stomachs (B, D).
Balcis mexicana BartscH, 1917. Rare (1, 11), 5-6
fm, sand and mud bottom, in sea star stomachs and
entangled in aborted egg mass (Br, C, Du).
Niso excolpa BartscH, 1917. Uncommon (11), 6 fm,
sand and mud bottom, entangled in aborted egg mass
(B, C, D, Du).
Epitonium keratium Dat, 1919. Uncommon (8, 11),
living on Pteria sterna shells, 18 fm; entangled in
aborted egg mass, 6 fm, sand and mud bottom (Br,
C, Du).

Epitonium walkerianum HERTLEIN & STRONG, 1951.
Uncommon (2, 11), 5 - 6 fm, sand and mud bottom,
in sea star stomachs and dead in aborted egg mass
(Br, C, G).

Epitonium reflexum (CARPENTER, 1856). Uncom-
mon (11), 6 fm, sand and mud bottom, dead in
aborted egg mass (C, Du).

Epitonium bakhanstranum KEEN, 1962. Uncommon
(11), 6 fm, sand and mud bottom, dead in aborted
egg mass (Br, C, Du).

Epitonium appressicostatum Daur, 1917. Uncom-
mon (11), 6 fm, sand and mud bottom, dead in
aborted egg mass (Du).

Epitonium barbarinum Datu, 1919. Common (11),
6 fm, sand and mud bottom, dead in aborted egg
mass (B, D, P).

Epitonium durhamianum HERTLEIN & STRONG,
1951. Rare (11), 6 fm, sand and mud bottom, dead
in aborted egg mass (B, D, P).

Lacuna succinea Morcu, 1860. Rare (8), 18 fm,
sand bottom, juveniles in washings from Pteria ster-
na; Gulf of Nicoya, Costa Rica (C, Du).
Cyclostremiscus bifrontia CARPENTER, 1857. Rare
(4), 5 fm, sand bottom, in sea star stomach (G).
Cyclostremiscus tricarinatus C.B.Apams, 1852.
Common (11), 6 fm, sand and mud bottom, entangled
in aborted egg mass (B, Br, C, D, Du, P).

THE VELIGER

180

187

233

240

242

245

248

251

Vol. 11; No. 4

Macromphalina sp. Rare (11), 6 fm, sand and mud
bottom, entangled in aborted egg mass (Br).
Delphinoidea hambachi Stronc & HERTLEIN, 1939.
Rare (11), 6 fm, sand and mud bottom, entangled in
aborted egg mass (P).

Teinostoma amplectans CARPENTER, 1857. Rare (11),
6 fm, sand and mud bottom, entangled in aborted
egg mass (C).

Teinostoma ecuadorianum Pitsspry & Oxsson, 1941.
Rare (11), 6 fm, sand and mud bottom, in aborted
egg mass; Punta Blanca, Ecuador (B, D, P).
Vitrinella dali (Bartscu, 1911). Uncommon (7),
11 fm, sand bottom, in sea star stomach (D).
Assiminea sp. Uncommon (15), 16 fm, sand and clay
bottom, on Pteria sterna shells (G).

Turritella anactor BERRY, 1957. Uncommon (7, 13),
11 fm, sand bottom, dead specimens and juveniles in
sea star stomachs (Br, C, Du, G).

Turritella nodulosa Kinc & Broperip, 1832. Uncom-
mon (11), 6 fm, sand and mud bottom, juveniles en-
tangled in egg mass (P).

Metaxia sp. Uncommon (11), 6 fm, sand and mud
bottom, in aborted egg mass (G).

Seila assimilata C. B. Apams, 1852. Common (11),
6 fm, sand and mud bottom, entangled in aborted
egg mass (B, Br, D, P).

Seila sp. Uncommon (11), 6 fm, sand and mud bot-
tom, entangled in aborted egg mass (C, Du).
Triphora hanna: Baker, 1926. Rare (8), 18 fm,
sand bottom, in washings from Pteria sterna (Du).
Iselica fenestrata CARPENTER, 1864. Common (8, 9,
14, 15), sand and clay bottom, on Pteria sterna; San
Diego, California (Br, C, Du, G, P).

Calyptraea mamillaris Bropertp, 1834. Common
(1, 7, 8, 9, 12 to 16), 5 - 21 fm, sand, clay, and mud
bottom; attached to dead shells (Br, Du, P).

Crepidula arenata (Broperip, 1834). Common (7,
8, 13), 11 - 18 fm, sand bottom, on Calliostoma pal-
ment (Du).

Crepidula incurva (Broperip, 1834). Common (7,
8, 13), 11-18 fm, sand bottom, on Calliostoma pal-
meri (Du).

Crepidula onyx SowERBY, 1824. Common on dead
shells (6, 7, 12, 13), 11-13 fm, sand bottom (B, Br,
D, Du, P).

Crepidula striolata MEeNKeE, 1851. Uncommon (7,
11), 6-11 fm, sand and mud bottom, entangled in
aborted egg mass and on dead bivalves (G, P).

Crucibulum personatum KEEN, 1958. Common (11,
13, 16), sand and mud bottom, on coral-related ma-
terial and on dead shells (B, Br, C, D, Du, P).

Vol. 11; No. 4

252 Crucibulum scutellatum (Woop, 1828). Common
(13, 16), 11 fm, sand bottom, on other shells (Br, C,
Duy PE):

254 Crucibulum spinosum (SoweErBy, 1824). Common
(8, 11, 14, 15), 6-21 fm, sand, mud, and clay bot-
tom; juveniles on other shells and entangled in abort-
ed egg mass (Du, G, P).

261 Natica idiopoma Pitssry &« Lowe, 1932. Rare (8),
18 fm, sand bottom, in sea star stomachs (B, P).

263 Natica broderipiana Réciuz, 1844. Uncommon.
(10), 14 fm, mud bottom, in sea star stomachs (Br,
G).

266 Polinices bifasciatus (GriFFITH & PcEoN, 1834 from
Gray MS.). Uncommon (13), 11 fm, sand bottom,
juvenile in sea star stomach (Br).

269 Polinices intemeratus (PxHivippt, 1853). Uncommon
(12), 13 fm, sand bottom (Du).

272 Polinices uber (VALENCIENNES, 1832). Uncommon
(1, 2,4), 5-7 fm, sand and mud bottom, in sea star
stomachs (B, D, G).

274 Polinices reclusianus (DESHAYES, 1839). Common
(1, 4, 5), 5-10 fm, juveniles in sea star stomachs
(B, Br, C, D, Du, G, P).

280 Lamellaria inflata (C. B. ApaMs, 1852). Rare (14),

_ 21 fm, sand and clay bottom, in washings from Pteria
sterna (B).

289 Erato columbella MENxKE, 1847. Uncommon (15),
16 fm, sand and clay bottom with clumps of Pteria
sterna, in sea star stomachs (D, P).

317 Ficus ventricosa (SowERBY, 1825). Uncommon (7,
13, 14), 11 - 21 fm, sand and clay bottom (C, D, Du,
G).

335 Murex elenensis Dati, 1909. Uncommon (7, 16),
11 fm, sand bottom (Du, V).

336a Murex recurvirostris lividus CARPENTER, 1857. Un-
common (7, 13), 11 fm, sand bottom (B, C, D, P).

339 Hexaplex erythrostomus (Swainson, 1831). Com-
mon (6, 7, 8, 12, 13, 14), 11 - 21 fm, sand and clay
bottom (Br, C, Du, P, V, W).

344 Muricanthus nigritus (PHttiprt, 1845). Common (6,
7, 12, 13), 11 - 13 fm, sand bottom (B, Br, C, D, Du,
G, P, V).

348 Pteropurpura erinaceoides (WALENCIENNES, 1832).
Uncommon (7, 13), 11 fm, sand bottom (B, Br, D, V).

364 Eupleura muriciformis (BropEriP, 1833). Common
(4, 11), 5-6 fm, sand and mud bottom, juveniles in
sea star stomachs and entangled in aborted egg mass
(B, Br, C, D, Du, G, P).

370* Vitularia salebrosa (KiNG & Broperip, 1832). Un-
common (8, 9, 14, 15), sand and clay bottom, at-
tached to Pteria sterna; Guaymas, Sonora, Mexico
(Br, C, Du, G).

THE VELIGER

Page 359

409 Acanthina tuberculata (SowrErBy, 1835). Uncom-
mon (7), 11 fm, sand bottom (C, Du).

417 Aesopus sanctus Dati, 1919. Uncommon (11), 6
fm, sand and mud bottom, in aborted egg mass (B,
Br, D, P).

* Aesopus chrysalloides CARPENTER, 1864. Rare (11),
6 fm, sand and mud bottom; San Diego, California
(Du, G).

429 Anachis diminuta (C.B.Apams, 1852). Uncommon
(11), 6 fm, sand and mud bottom, dead in aborted
egg mass (Du, P).

442 Anachis milium (Dat, 1916). Common (3, 11),
taken in rope tangles, 3 fm, rocky bottom; in aborted
egg mass, 6 fm, sand and mud bottom (Br, C, Du).

444 Anachis nigricans (SowErBy, 1844). Uncommon
(11), 6 fm, sand and mud bottom, dead in egg mass
(P).

454 Anachis sanfelipensis Lowe, 1935. Uncommon (11),
6 fm, sand and mud bottom, juveniles dead in egg
mass (Du).

464 Anachis varia (SoweRBy, 1832). Common (3, 7,
11), taken in rope tangle, 3 fm, rocky bottom; in sea
star stomachs and on chunks of coral-related materi-
al, 6 - 11 fm, sand and mud bottom (B, Br, C, D, Du,
GSP)r

470 Cosmioconcha palmeri (Dati, 1913). Common as
juveniles (2, 5, 8, 11), 6-18 fm, sand and mud bot-
tom, in sea star stomachs and in egg mass (B, Br, C,
D, Du, G, P).

490 Nassarina pammicra PitsBry & Lowe, 1932. Rare
(15), 16 fm, sand and clay bottom, on Pteria sterna
(Du).

Nassarina anitae CAMPRELL, 1961. Uncommon (8),
18 fm, sand bottom, in washings from Pteria sterna
(Gs ID wy, IP )e

Nassarina, possibly new spec. Rare (11), 6 fm, sand
and mud bottom, in aborted egg mass (Br, G).

508 Strombina dorsata (SowerBy, 1832). Uncommon
(10, 11, 14), 6- 21 fm, sand, mud, and clay bottom;
in sea star stomachs and dead in egg mass (Br, Du).

512 Strombina gibberula (SoweRsy, 1832). Uncommon
(14), 21 fm, sand and clay bottom with clumps of
Pteria sterna, in sea star stomachs (B, C, D, Du, G).

541 Solenosteira capitaneus (Berry, 1957). Uncommon
(8, 11), 6- 13 fm, sand bottom (Br, P).

543 Solenosteira macrospira (BERRY, 1957). Uncommon
(8, 11), 6- 18 fm, sand bottom, dead specimens and
juveniles entangled in egg mass (C, Du, G).

563 Phos gaudens Hinps, 1844. Common (11, 14), 6
to 21 fm, mud, sand, and clay bottom; in sea star
stomachs and entangled in egg mass (B, Br, C, D,
Du, G, P).

Page 360

577 Nassarius guaymasensis (Prtspry & Lowe, 1932).
Common (11), sand and mud bottom, dead in egg
mass (C, Du, G).

583 Nassarius pagodus (REEvE, 1844). Uncommon (2,
6, 10), 6 - 14 fm, sand and mud bottom (Br, Du, P).

586 Nassarius taeniolatus (Puiipri, 1845). Common (2,
10, 11), 6- 14 fm, sand and mud bottom, in sea star
stomachs and entangled in egg mass (B, Br, C, D,
Du, G, P).

587 Nassarius versicolor (C. B. ApaMs, 1852). Common
(9, 10, 11), 6- 15 fm, sand and mud bottom, in sea
star stomachs and in egg mass (B, Br, C, D, Du, G, P).

591 Nassarius iodes (Dati, 1917). Common (11), 6 fm,
sand and mud bottom, entangled in egg mass (Du,
G, P).

593 Nassarius moestus (Hinps, 1844). Uncommon (1,
2), 5-7 fm, sand bottom, in sea star stomachs (Du, P).
Nassarius howardae Cuace, 1958. Uncommon (11),
6 fm, sand and mud bottom, entangled in egg mass
(Br, P).

610 Fusinus dupetitthouars: (KieNeR, 1846). Uncom-
mon (8), 18 fm, sand bottom, dead (Br, C).

612 Fusinus ambustus (Goutp, 1853). Uncommon (13),
11 fm, sand bottom, dead (G).

614 Fusinus fredbakert Lowe, 1935. One dead specimen
(11), 6 fm, sand bottom, entangled in egg mass (C).
Fusinus, new spec. Uncommon (7, 13), 11 fm, sand
bottom (C, Du).

Fusinus spec. (possibly new, or dredged form of EF
ambustus). Common (7 to 10, 13, 16), 11-16 fm,
sand and mud bottom (B, Br, D, Du, P, V).

625 Oliva spicata (R6p1NG, 1798). Uncommon (11, 14),
6-21 fm, sand bottom (Br, C, Du, G).

627 Oliva undatella LaMarck, 1810. Uncommon (11),
6 fm, sand bottom, dead (C, Du, P).

634a Olivella fletcherae BERRY, 1958. Common (2, 4, 7,
11, 13, 16), 5- 11 fm, sand bottom, in sea star stom-
achs and in egg mass (Br, C, Du, G, P).

Olivella stevent BurcH & CAMPBELL, 1963. Rare
(4), 5 fm, sand bottom, in sea star stomachs (P).

693 Cancellaria cassidiformis SowERBY, 1832. Uncom-
mon (6, 7), 11-12 fm, sand and mud bottom (C,
Du, G).

717 Daphnella bartschi Dati, 1919. Rare (13), 11 fm,
sand bottom, in sea star stomachs (B, D).

741 Clavus roseolus (HERTLEIN & STRONG, 1955). Rare
(7), 11 fm, sand bottom, in sponge (P).

747° Clavus aeolius (Dati, 1919). Uncommon (1, 11),
5 - 6 fm, sand and mud bottom, in sea star stomachs
and in egg mass (B, C, D, Du, G).

748 Clavus aerope (Datu, 1919). Rare (11), 6 fm, sand
and mud bottom, in egg mass (Br).

THE VELIGER

Vol. 11; No. 4

753 Clavus ianthe (Dati, 1919). Common (11), 6 fm,
sand and mud bottom, in egg mass (B, Br, C, D, Du,
G).

758 Clavus pembertoni Lowe, 1935. Uncommon (7), 11
fm, sand bottom, in sponge (P).

765 Clathrodrillia callianira Datu, 1919. Rare (15), 16
fm, sand and clay bottom with clumps of Pteria
Sterna, in sea star stomachs (Br).

767 Clathrodrillia maura (SowerRsy, 1834). Rare (8),
18 fm, sand bottom, in sponge (Br).

769 Clathrodrillia pilsbryi Lowe, 1935. Uncommon (11,
14), 6-21 fm, sand, mud, and clay bottom; in sea
star stomachs and in egg mass (B, Br, D, Du, G).

771 Clathrodrillia adonis (Pirspry & Lowe, 1932). Un-
common (15), 16 fm, sand and clay bottom, from
sea star stomachs (Du).

772 Clathrodrillia alcestis Dat, 1919. Rare (15), 16 fm,
sand and clay bottom, in sea star stomach (P).

774 Clathrodrillia duplicata (Sowersy, 1834). Un-
common (7), 11 fm, sand bottom, in sponge (B, Br,
D).

776 Clathrodrillia halis Dati, 1919. Uncommon (11),
6 fm, sand and mud bottom, entangled in egg mass
(B, D, P).

776a Clathrodrillia halis soror (Pitspry & Lowe, 1932).
Common (6, 7, 11, 13), 6 - 12 fm, sand and mud bot-
tom; in sponge, in sea star stomachs, and in aborted
egg mass (Br, C, Du, G, P).

790 Crassispira bacchia Datu, 1919. Common (6, 11),
6 - 12 fm, sand and mud bottom, in sea star stomachs
and dead in egg mass (B, Br, C, D, Du, P).
Crassispira cf. C. pauxillus Reeve, 1843. Uncommon
(1), 5 fm, sand bottom, in sea star stomachs (Du).

825 Crassispira pluto Pispry « Lowe, 1932. Uncom-
mon (11), 6 fm, sand and mud bottom, dead in egg
mass (Br).

843 Hindsiclava militaris (Reeve, 1843 ex Hinps MS).
Uncommon (7), 11 fm, sand bottom, in sponge (Br).

858 Mangelia aethra (Dati, 1919). Rare (11), 6 fm,
sand and mud bottom, in egg mass (D).

862 Mangelia melita (Dati, 1919). Common (11), 6
fm, sand and mud bottom, in egg mass (B, Br, C, D,
Du, P).

866 Mangelia subdiaphana CarPENTER, 1864. Rare (11),
6 fm, sand and mud bottom, in aborted egg mass
(C, Du).

867 Mangelia antiochroa Pitspry & Lowe, 1932. Com-
mon in sea star stomachs (B, D, G).

868 Mangelia antipyrgus Pitspry & Lowe, 1932. Com-
mon (11), 6 fm, sand and mud bottom, in egg mass
(C, Du, P).

Vol. 11; No. 4

869

875

881

883

909

910

Mangelia cymatias Pitspry « Lowe, 1932. Uncom-
mon (9, 11), 6- 15 fm, sand and mud bottom, in sea
star stomachs and in egg mass (B, Br, D).

Mangelia roperi Dati, 1919. Uncommon (1, 11),
5 - 6 fm, sand and mud bottom, in sea star stomachs
and entangled in egg mass (B, C, D, Du).

Clathurella rigida (Hinps, 1843). Rare (2), 6-7
fm, sand bottom, in sea star stomach (B).
Clathurella trichoides (Dat, 1919). Rare (15), 16
fm, sand and clay bottom (B).

Clathurella acapulcana (Pirspry & Lowe, 1932).
Uncommon (10), 14 fm, mud bottom, in sea star
stomachs (Du, G, P).

Clathurella adria (Dax, 1919). Uncommon (11),
6 fm, sand and mud bottom, in aborted egg mass (B,
D, Du, P).

Pleuroliria artia Berry, 1957. Uncommon (8), 18
fm, sand bottom, dead (G).

Pleuroliria nobilis (Hinps, 1843). Rare (13, 15),
11 - 16 fm, sand and clay bottom, in sponges and in
Pteria sterna clumps (Du, G).

91la Pleuroliria oxytropis albicarinata (SowERBY, 1870).

913

926

928

956

963

966

980

992

Uncommon (7, 13), 11 fm, sand bottom, in sponges
(Br, P).

Pleurolinia picta (REEVE, 1843, ex Beck MS). Un-
common (13), 11 fm, sand bottom, in sponges (Du, P).

Conus perplexus SowErBy, 1857. Rare (8), 18 fm,
sand bottom, juveniles (C).

Conus tornatus SowERBy, 1833, ex BropErip MS.
Uncommon (9), 11 fm, sand bottom (Du).

Conus poormani Berry, 1968. Rare (13), 11 fm,
sand bottom (Br).

Terebra armillata Hinps, 1844. Juveniles common
(11, 14), 6-21 fm, sand, mud, and clay bottom; in
sea star stomachs and in aborted egg mass (B, Br, C,
D, Du).

Terebra glauca Hinps, 1844. Common (6, 11),
6 - 12 fm, sand and mud bottom, juveniles in sea star
stomachs and in egg mass (B, Br, C, D, Du, G).

Terebra ira PitsBry & Lowe, 1932. Uncommon (11),
6 fm, sand and mud bottom, juveniles in aborted egg
mass (B, Br, C, D, Du).

Terebra variegata Gray, 1834. Uncommon (4, 5),
5 - 10 fm, sand and mud bottom, juveniles in sea star
stomachs (G).

Terebra dushanae CAMPBELL, 1964. Rare (11), 6
fm, sand and mud bottom, dead in aborted egg mass
(G).

Acteocina angustior BAKER & Hanna, 1927. Com-
mon (2, 4, 8, 9, 14, 15), 5-18 fm, sand and clay
bottom, in sea star stomachs and in washings from
Pteria sterna (B, Br, C, D, Du).

THE VELIGER

Page 361

994 Acteocina inculta (GoutD & CarPENTER, 1857).

997

Uncommon (11), 6 fm, sand and mud bottom, in
aborted egg mass (G).

Cylichna defuncta Baker & Hanna, 1927. Uncom-
mon (1), 5 fm, sand bottom, in sea star stomachs (C,
Du).

Cylichna fantasma (BAKER « Hanna, 1927). Rare
(11), 6 fm, sand and mud bottom, in aborted egg
mass (C, Du).

Cylichnella sp. Rare (11), 6 fm, sand and mud bot-
tom, in aborted egg mass (G).

1000 Pyramidella adamsi CarPENTER, 1864. Rare, in

sea star stomachs (G).

1003. Pyramidella mazatlanica Dat. & BartscH, 1909.

Uncommon (10), mud bottom, in sea star stomachs
(D).

Odostomia clathratula C. B. ApaMs, 1852. Rare (11),
6 fm, sand and mud bottom, in egg mass (P).
Odostomia corintoensis HERTLEIN & STRONG, 1951.
Uncommon (6), sand bottom, in sea star stomach;
Corinto, Nicaragua (Du).

Turbonilla academica STRONG & HERTLEIN, 1939.
Rare (11), 6 fm, sand and mud bottom, in egg mass;
Bahia Honda, Panama (G).

Turbonilla amortajadensis BAKER, HANNA & STRONG,
1928. Rare (11), 6 fm, sand and mud bottom, in
egg mass (Du).

Turbonilla azteca BAKER, HANNA & STRONG, 1928.
Uncommon (11), 6 fm, sand and mud bottom, in
egg mass (Du).

Turbonilla contrerasiana HERTLEIN & STRONG, 1951.
Rare (11), 6 fm, sand and mud bottom, in egg mass
(Du).

Turbonilla gonzagensis BAKER, HANNA & STRONG,
1928. Rare (11), 6 fm, sand and mud bottom, in egg
mass (Du).

Turbonilla prolongata CarPENTER, 1857. Uncommon
(10, 11), 6-14 fm, mud and sand bottom, in sea
star stomachs and in aborted egg mass (B, C, D, P).
Nembrotha eliora Marcus, 1967. Uncommon (14),
21 fm, sand and clay bottom with clumps of Pterza
sterna (C).

Coryphella cynara Marcus, 1967. Uncommon (14),
21 fm, sand and clay bottom with clumps of Pteria
sterna (C).

Flabellinopsis iodinea (Cooper, 1862). Uncommon
(8, 9, 14, 15), 15 - 21 fm, sand and clay bottom with
clumps of Pteria sterna (C).

Spurilla chromosoma CocKERELL & Euiot, 1905.
Uncommon (8, 9, 14, 15), sand and clay bottom, on
Pteria sterna (C).

Page 362

LITERATURE CITED

ApaMs, CHARLES BAKER
1852. Catalogue of shelis collected at Panama, with notes on
synonymy, station, and habitat. Ann. Lyceum Nat.
Hist. New York 5: 229 - 344 (June) ; 345-548 (July)
Appincton MSS No. 17660 British Museum
BAKER, FREDERICK
1926. Mollusca of the family Triphoridae.
Acad. Sci., ser. 4; 15 (6) :223 - 239; plt. 24
BAKER, FREDERICK & G Dattas HANNA
1927. | Marine Mollusca of the order Opisthobranchiata.
Proc. Calif. Acad. Sci., ser. 4, 16 (5): 123-135; plt. 4
(22 April 1927)
Baker, FREDERICK, G DaLLas HANNA & ARCHIBALD M. STRONG
1928. Some Pyramidellidae from the Gulf of California.
Proc. Calif. Acad. Sci., ser. 4, 17 (7): 205 - 246; plts. 11-12
(29 June 1928)

Proc. Calif.
(26 April 1926)

BartscH, PAauL
1911. The Recent and fossil mollusks of the genus Cerithi-
opsis from the west coast of America. Proc. U.S. Nat.
Mus. 40 (1823) : 327 - 367; plts. 36 - 41 (8 May 1911)
1917. A monograph of West American melanellid mollusks.
Proc. U.S. Nat. Mus. 53 (2207): 295-356; plts. 34-49
(13 August 1917)
Berry, SAMUEL STILLMAN
1958. Is the Colorado River an efficient barrier to mollusk
distribution? Amer. Malacol. Union Bull. 24: 24
1962. Leaflets in Malacology 1 (20): 123-130 (13 Nov.)
1964. Notices of new eastern Pacific Mollusca — VI. Leaflets
in Malacology 1 (24): 147 - 154 (29 July 1964)
1968. Notices of new eastern Pacific Mollusca. -— VII.
Leaflets in Malacology 1 (25): 154 - 158 (26 Sept. 1968)
Burcu, JoHN Quincy « G. Bruce CAMPBELL
1963. Four new Olivella from the Gulf of California.
Nautilus 76 (4): 120-126; 2 plts.; 6 figs. (April 1963)
CampBELL, G. Bruce
1961. Four new Panamic gastropods. The Veliger 4 (1):
25 - 28; plt. 5 (1 July 1961)
1964. New terebrid species from the eastern Pacific (Mollusca:
Gastropoda). The Veliger 6 (3): 132 - 138; plt. 17
(1 January 1964)
CARPENTER, PHILIP PEARSALL
1857. © Catalogue of the collection of Mazatlan shells in the
British Museum collected by Frederick Reigen. London,
British Museum xvi + 552 pp. (June 1857)
1864. | Supplementary report on the present state of our know-
ledge with regard to the Mollusca of the west coast of North
America. Reprt. Brit. Assoc. Adv. Sci. for 1863: 517 - 686
(August 1864)
Cuace, Emery PERKINS
1958. A new mollusk from San Felipe, Baja California.
Trans. San Diego Soc. Nat. Hist. 12: 333 - 334; fig. 1
(16 October 1958)
CockERELL, THEODOR Dru ALISON & CHARLES ELIOT
1905. Notes on a collection of California nudibranchs.
Journ. Malacol. 12 (3): 31-53; plts. 7, 8
Cooper, James GRAHAM
1862. Some genera and species of California Mollusca.
Proc. Calif. Acad. Nat. Sci. 2: 202 - 207

THE VELIGER

Vol. 11; No. 4

DaLL, WILLIAM HEALEY
1918. Descriptions of new species of shells, chiefly from Mag-
dalena Bay, Lower California. Proc. Biol. Soc. Washington
31: 5-8 (27 February 1918)
1919. Descriptions of new species of Mollusca from the North
Pacific Ocean in the collection of the United States National
Museum. Proc. U.S. Nat. Mus. 56 (2295): 293-371
(30 August 1919)
DuSHANE, HELEN
1962. A checklist of mollusks for Puertecitos, Baja California,
Mexico. The Veliger5 (1): 39-50;1map (1 July 1962)
DuSwan_e, HELEN « Roy PoorRMAN
1967. A checklist of mollusks for Guaymas, Sonora, Mexico.
The Veliger 9 (4): 413-441; 1 map (1 April 1967)
DuSwane, HELEN & GaLe G. SPHON
1968. A checklist of intertidal mollusks for Bahia Willard and
the southwestern portion of Bahia San Luis Gonzaga, Baja
California, Mexico. The Veliger 10 (3) : 233 - 246; plt. 35;

1 map. (1 January 1968)
Emerson, WILLIAM KEITH
1958. Results of the Puritan-American Museum of Natural

History expedition to western Mexico. 1. General account.
Amer. Mus. Novit. No. 1894: 1 - 25; 9 figs. (22 July 1958)
ENGELHARDT, Fr. ZEPHYRIN, O. F. M.

1929. The Missions and Missionaries of California, vol. 1
Lower California. Mission Santa Barbara, Santa Barbara,
California, 784 pp.

FarMErR, WESLEY M.

1963. Two new opisthobranch mollusks from Baja California.
Trans. San Diego Soc. Nat. Hist. 13 (6): 81 - 84; plt. 1; fig. 1
1966. Range extension of Berghia amakusana (Basa) to the

East Pacific. The Veliger 9 (2): 251; 1 text fig.
(1 October 1966)
1967. Notes on the Opisthobranchia of Baja California, Mexi-
co, with range extensions - II. The Veliger 9 (3): 340 - 342;
1 text fig. (1 January 1967)
Fraser, C. McLEAN
1943. General account of the scientific work of the Velero III
in the eastern Pacific, 1931-41. Part III. A ten-year list of
the Velero III collecting stations (charts 1 - 115). Allan
Hancock Pacif. Exped. 1 (3) : 259 - 431. Univ. So. Calif: Press,
Los Angeles, California
HeErTLEIN, LEo Grorce & ARCHIBALD McCiure STRONG
1951. | Eastern Pacific expeditions of the New York Zoological
Society. XLIII. Mollusks from the west coast of Mexico and
Central America; PartsI-X. Zoologica, prt. 10; 36: 67 - 120;

pits. 1-11 (20 August 1951)
KEEN, A. Myra
1958. Sea shells of tropical West America; marine mollusks

from Lower California to Colombia.
Stanford, Calif. (Stanford Univ. Press)
1966. | MorcwH’s west central American molluscan types with
proposal of a new name for a species of Semele. Calif.
Acad. Sci. Occ. Papers 59: 1 - 33; 41 figs. (30 June 1966)

LaNncE, JAMES ROBERT

1961. A distributional list of southern California opistho-
branchs. The Veliger 4 (2) : 64 - 69 (1 October 1961)
1966. New distributional records of some northeastern Pacific
Opisthobranchiata (Mollusca: Gastropoda) with descriptions
of two new species. The Veliger 9 (1) : 69- 81; 12 text figs.
(1 July 1966)

i-xi + 624 pp.; illus.

Vol. 11; No. 4

Lance, JAMES ROBERT
1968. New Panamic nudibranchs (Gastropoda: Opisthobran-
chia) from the Gulf of California. Trans. San Diego
Soc. Nat. Hist. 15 (2): 13 pp.; 2 plts.; 5 figs. (8 January)
LeicH, RANDOLPH
1941. Forgotten waters.
New York, N. Y.
McLean, James Howarp
1961. Marine mollusks from Los Angeles Bay, Gulf of Cali-
fornia. Trans. San Diego Soc. Nat. Hist. 12 (28): 449 to
476; figs. 1-3 (15 August 1961)
Marcus, EveLine « Ernst Marcus
1967. American opisthobranch mollusks. Studies in tropical
Oceanography no. 6. Inst. Marine Sci., Univ. Miami,

324 pp. J.B.Lippincott Co.,

Florida; viii+ 256 pp.; 1 plt.; 155 +95 text figs. (December)
Moore, Raymonp Cecm. (editor and director)
1956. Treatise on invertebrate paleontology - (F) Coe-

lenterata — Geol. Soc. ; Univ. Kansas Press; 498 pp.; illus.
Morcu, Orro AnprEAs Lowson
1860. Beitrige zur Molluskenfauna Central-Amerikas.
Malak. Blatter 7: 66-96 (July 1860); 97- 106 (Aug. 1860)
NeEtson, Epwarp WILLIAM
1922. Lower California and its natural resources. Nat.
Acad. Sci. 16. First Mem. Governmt. Printing Office, Wash-
ington, 194 pp. (reprinted 1966)
Oxsson, AxEL ADOLF
1961. Mollusks of the tropical eastern Pacific particularly
from the southern half of the Panamic-Pacific faunal pro-
vince (Panama to Peru). Panamic-Pacific Pelecypoda.
Paleont. Res. Instit. Ithaca, N. Y.: 574 pp.; 86 plts.
(10 March 1961)

Syxes, GopFREY
1937. | The Colorado delta.
no. 19, 193 pp.

U.S. Navy Oceanographic Office

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Page 363

Parker, Rosert H.

1963. Zoogeography and ecology of some macro-invertebrates,
particularly mollusks, in the Gulf of California and the conti-
nental slope off Mexico. Vidensk. Medd. Dansk Naturhist.
For., 126: 1-178; plts. 1-15

Pitspry, Henry Aucustus « Axe, ApoLtF OLsson

1941. A Pliocene fauna from Western Ecuador.

Nat. Sci. Philadelphia 93: 1 - 79; plts. 1 - 19
ReEEvE, Lovett Aucustus

1843. Descriptions by Mr. Lovell Reeve of new species of
shells figured in the “Conchologia systematica.” Proc. Zool.
Soc. London for 1842: 197 - 200 (February 1843)

Ropen, Gunnar I.

1958. | Oceanographic and meteorological aspects of the Gulf

of California. Pacific Sci. 12 (1): 21-45; 19 figs.; 4 tables
(January 1958)

Proc. Acad.
(9 Sept. 1941)

RopEen, Gunnar I. « G. W. Groves

1959. Recent oceanographic investigations in the Gulf of Cali-
fornia. Journ. Mar. Res. Sears Found. 18 (1): 10-35; 16
figs.; 3 tables (30 June 1959)

SLEvin, JoserH R.

1923. Expedition of the California Academy of Sciences to
the Gulf of California in 1921. General Account. Proc.
Calif. Acad. Sci., ser. 4, 12 (6) :55-73; 1 map (June 1923)

Smiru, ALLYN Goopwin

1966. Staghorns and longhorns. Pacif. Discov. 19 (5):

30-31; 5 photographs (September/October 1966)
Srronc, ArcHiBALD McCuure & Leo GrorcE HERTLEIN

1939. Marine mollusks from Panama collected by the Allan
Hancock expedition to the Galapagos Islands, 1931 - 32.
Allan Hancock Publ. Univ. South. Calif. 2 (12): 177 - 245;
plts. 18 - 23 (21 August 1939)

Amer. Geogr. Soc. spec. publ.

Surveys between 1873 and 1875, with additions to 1962 (chart
620) Dept. Navy, Washington, D.C.

Page 364

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Vol. 11; No. 4

Two New Species of the Genus Volva Ropinc, 1798
(Ovulidae FLeminc, 1828)

CRAWFORD N. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plate 56)

THE PURPOSE OF THIS PAPER is to describe two new
species belonging to the genus Volua Ropinc, 1798; it
will list and illustrate the more common Pacific species for
comparative purposes, and will present the currently
accepted synonymy. For further comprehensive detail of
the family Ovulidae FLEmine, 1828, the reader is referred
to ScHILDER, 1932.

CYPRAEACEA Gray, 1824

OvuLwAE FLEMING, 1828

Syn.: AMPHIPERATIDAE SCHILDER, 1927 (em.)

Ovulinae FLEminc, 1828

(Ovulini) FLemine, 1828

Volva Ropine, 1798

Mus. Bolten. 21
Type species: Bulla volua LinNaEus, 1758

Syn.: Radius Montrort, 1810.
Conchyl. Syst., 2: 627
Type species: Bulla volua LinnaEus, 1758

Birostra Swainson, 1840.
Treat. Malacol. 1840: 325
Type species: Bulla volua LinnaEus, 1758

(Phenacovolva) Irepae, 1930.
Mem. Queensld. Mus., 10 (1): 85
Type species: Phenacovolva nectarena IREDALE, 1930

1. Voluva (Phenacovolva) birostris (LINNAEUS, 1758)
Syst. Nat., ed. 10: 725

(Plate 56, Figure 3)

Syn.: Ovula longirostrata SowErRBy ', 1828
Zool. Journ. 1828, 4: 160

The species is known to range into Japanese waters and
throughout the Malayan Archipelago, from the Gulf of
Tonkin, to Vietnam, to Siasi Island, to New Guinea, and
along the east coast of Australia. The species appears to
maintain a rather constant shell morphology, differing
only as to length, and it can be recognized without much
difficulty. From available information, it would appear
to be most abundant on an unnamed species of coral
(local collectors call it Alcyon-type coral). The shells
seem to average about 41mm in length and 7.5mm in
width at the body whorl and outer lip. ‘The specimen illus-
trated here was collected at Cooktown, Queensland, Aus-
tralia, living on the coral mentioned above.

2. Volua (Phenacovolva) brevirostris (ScHUMACHER, 1817)
Ess. Nouv. Syst. Vers Test. 1817: 259

(Plate 56, Figure 5)

Syn.: Ovule birostre Lamarck, 1810
Ann. Mus. Hist. Nat. Paris 16: 113
Ovula rosea A. Apams, 1854
Proc. Zool. Soc. London 22: 130
Ovula recurva A. ApAMS & REEVE, 1848
Voy. Samarang, Moll., 1848: 21; plt. 6, fig. 3
Cyphoma elongatum A. ApamMs, 1864
J. P. Linn. Soc. London 7: 96
Phenacovolva nectarena IREDALE, 1930
Mem. Queensld. Mus., 10 (1): 85; plt. 9, fig. 6

This species has a wide living range throughout the
Pacific region. It is a rather common species along the
East Asian coast and the offshore islands, including the
Philippines, Taiwan, Ryukyu Islands, and the Japanese is-
land chain, as well as the coast of mainland China; other-
wise the distribution of this species is as far east as
Hawaii, then south to the Cook Islands, and southwest
into the Celebes-Sulu Seas. The shell possesses a charac-
teristic shape, with sharply angled shoulders; a singular

THE VELIGER, Vol. 11, No. 4 [C. N. Cate] Plate 56

Figure 1 Figure 2
Volva (Phenacovolva) brunneiterma Volva (Phenacovolva) lahainaensis
C. N. Cate, spec. nov. X 23 C. N. Cate, spec. nov. X 14

Figure 3
Volva brevirostris (ScHUMACHER, 1817) X 2

Biomed Figure 5
Volva (Phenacovolva) philip pinarum
(Sowersy ?"4, 1848) x 24 Volva birostris (LINNAEUS, 1758) X 13

photographs by Jean M. Cate

Vol. 11; No. 4

shade of beige-grey color; a shorter, broader shell, with
shorter, sturdier terminal beaks and with a conspicuous
white transverse band of color across the central dorsal
area. The shell varies in size, with an average length of
29mm and an average width of 11.5mm at the body
whorl and lip. The specimen illustrated was collected in
190 feet of water off Lahaina, Maui, Hawaii, living on
black coral, the same habitat as one of the species de-
scribed as new herein.

3. Volua (Phenacovolva) philip pinarum (Sowersy ™, 1848)
Proc. Zool. Soc. London 1848: 136
(Plate 56, Figure 4)

Syn.: Volva carpentert DuNKER, 1877
Malakol. Blatter 24: 75
Volua maccoyi TENIsoN-Woops, 1878
Trans. Roy. Soc. Victoria 14: 56
Volva haynesi SowErRsy '"', 1889
Journ. Linn. Soc. London 20: 397; plt. 25, figs. 1-2

This is one of the smallest Volva species to be considered
in this report, with the shell averaging about 21.5mm in
length and 5.5mm in width. The species seems to range
from Japan south through the Ryukyus, Taiwan, the Phil-
ippines, the east coast of Australia, and possibly into
South Australia [Volua (Phenacovolva) exsul (IREDALE,
1935) is probably the same as V. philippinarum]. VeERco,
of the South Australian Museum, recorded a specimen
collected in South West Australia as Ovula philippinarum
Sow_Ersy; it had been dredged from 72 fathoms, 40 miles
at sea from Eucla. I have not seen this shell and therefore
cannot ascertain the correctness of this identification at
this time. The Philippine Islands seem to be the center
of greatest abundance for this species. The shell is narrow,
with an outer lip somewhat flaring anteriorly, with fine
transverse dorsal sculpture; it varies in color from pale
beige-white to pink. The specimen illustrated is from Kii,

Japan.

4. Volva (Phenacovolva) lahainaensis C. CATE, spec. nov.
(Plate 56, Figure 2)

Shell narrow, elongately-ovate, subcylindrical, light in
weight though strong; body whorl inflated, narrowing ab-
ruptly toward the terminals; dorsum transversely sculp-
tured with faint, parallel, zigzag embossed lines adapically
and abapically, though lines do not zigzag anteriorly; ter-
minals long, narrow, pinched laterally, both in front and
in back, more so to the front; aperture narrow to the rear,
widening, becoming openly constricted abapically; left
margin rounded, not thickened, right margin on outer lip

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Page 365

thickened, narrowly, sharply, abruptly formed, creating
a sharp angle with the body whorl, thus becoming keeled
into a distinct carina, which extends entire length of
outer lip; fossula primitively formed, subconcave, slightly
more so centrally; columella smooth, glossy, with a bifid
(almost trifid) first funiculum adapically (see Carte,
1964: plt. 19, fig. 3); shell predominantly translucent
orange in color, although the body whorl reflects an
overtone of orange-grey; a distinct, deep, rich orange
line encircles the shell at the margins, broken only by ter-
minal openings, — on the right side enveloping the sharp
marginal angle of the outer lip carina; untoothed, semi-
sharp, adaxial edge of outer lip noticeably lighter in color
in some specimens; interior of shell, fossula, columella,
and interior of terminal channels rich orange color, funic-
ulum a shade lighter.

Type Locality: The specimen was collected alive on black
coral in 190 feet of water off the southwest shore of the
Hawaiian island of Maui; the specific area is referred to
locally as “Lahaina Roads” (20°52’N Lat.; 156°41’ E
Long.), which may generally be stated as being between
Maui and Lanai. SCUBA divers brought the coral to the
surface where this and other specimens were easily re-
moved. It was sent to me for identification by Clifton S.
Weaver of Lanikai, Oahu, Hawaii.

Type Repository: The holotype will be deposited in the
Bernice P. Bishop Museum, Honolulu, Hawaii; it will bear
the Malacology Catalogue no. 217597.

Discussion: The holotype of Volva lahainaensis is 31.2
mm long and 9.1 mm broad at the outer lip, and 7.3 mm
high from ventral to dorsal surface. The shell shows a
gently curving aperture, flaring, widening, then abruptly
constricting abapically. To my knowledge there are be-
tween 5 and 10 specimens known at present; all are from
deep water habitats, and all have been collected from
black coral. At the present time it is not known whether
the species occurs elsewhere in this island chain; future
discovery will determine the extent of its range.

This new Hawaiian species appears morphologically
distinct from other forms of Volva. It differs from another
Hawaiian species, V. brevirostris (see Plate 56, Figure 5),
by its lighter weight, narrower, less sharply angled shoul-
ders, and slimmer overall appearance; by its lack of a
central color band; by the intense, dark orange color;
and by the presence of a deep, rich orange peripheral
line bordering the sides of the shell. Also, V. brevirostris
appears generally wider and shorter.

Perhaps the most similar form would be that of the
color variant of Volva brevirostris mentioned as living on
the coasts of mainland China and Japan, reported as Ovu-
la rosea A. ApaMs, 1854. However, its apparent strong

Page 366

morphological affinity to V. brevirostris despite its pinkish
coloring seems to separate the two forms. The new Maui
species differs from V. rosea by being a narrower, more
delicate shell; by the absence of the angulate shoulder;
by the presence of the distinct funiculum, by the intense
orange coloring, and particularly by the presence of the
encircling orange line.

5. Voluva (Phenacovolva) brunneiterma C. CATE, spec. nov.
(Plate 56, Figure 1)

Shell delicate, though strongly constructed, well formed,
elongately narrow; dorsal surface smooth (except for mi-
nute, barely visible, longitudinal growth lines), untex-
tured; funiculum absent; terminals minutely flaring; ap-
erture exceedingly narrow, opening broadly at constric-
tion of outer lip anteriorly; left margin not thickened,
rounded; right margin without callus, lip flatly thickening
adaxially, then rolling and curving upward within; fos-
sula smooth, not impressed; primary shell color milk-
white; terminal ends dark brown, with a pale brush of
yellow-brown abapically the length of terminal ridge area.
Type Locality: The specimen was found in the subtidal
waters off Siasi Island, Sulu Sea (5°32’N Lat.; 120°52’E
Long.). It was collected in 1960, and was sent to me by
Mr. Fernando G. Dayrit, Manila, Philippines, for identi-
fication.

Type Repository: The holotype, a unique shell, will be
deposited in the California Academy of Sciences Geology
Department Type Collection, where it will bear the number
13161.

Discussion: The holotype of Volva brunneiterma is 20.3
mm long, 4.7 mm broad at the left margin and outer lip,

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Vol. 11; No. 4

and 4.2 mm high. This new species has been confused with
V. philippinarum, which it vaguely resembles; however, it
differs radically by being a narrower shell; by possessing a
nearly closed aperture; by the compact outer lip (not
flaring as in V. philippinarum); by having a smooth
dorsal surface, although there are very faint growth lines
longitudinally (as compared with the horizontally sculp-
tured, evenly spaced lines of the latter species) ; by the
outer lip being peculiarly crimped and constricted ante-
riorly; by the dark brown coloring at the terminal ex-
tremities. Because of the lack of adequate material, the
distribution of this new taxon is at present unknown.

The conspicuous dark brown terminals are a character-
istic color feature in this species, providing a means for
easily separating it from others. Therefore it seemed ap-
propriate to use this distinguishing character as a basis
for its specific name, which is derived from the Latin
word brunneus (dark brown) and the Greek terma (for
the shell terminals) ; hence the name brunneiterma.

I want to thank Jean Cate for the excellent photo-
graphic work, and Dr. Takeo Susuki for the film proces-
sing. I also want to thank Dr. Franz Alfred Schilder for
his considerations of the new species.

LITERATURE CITED

Cate, Crawrorp NEILL ¢
1964. A new species of Primovula from the Philippines (Mol-
lusca: Gastropoda) . The Veliger 7 (2): 102-103; plt. 19
(1 October 1964)
ScHILDER, FRANZ ALFRED

1932. The living species of Amphiperatinae. Proc. Malacol.
Soc. London 20 (1): 46-64; plts. 3-5 (78 figs.)

Vol. 11; No. 4

THE VELIGER

Page 367

Zoogeographical Studies on Living Cowries

FRANZ ALFRED SCHILDER

University of Halle, German Democratic Republic

(1 Map)

Four years Aco I published a systematic list of species
and races of living cowries with exact indications of the
then known geographical distribution (ScHILDER, 1965).
In the intervening 4 years several new cowrie species have
been discovered and enthusiastic collectors have enlarged
the knowledge of occurrence by filling small gaps in evi-
dently continuous distribution as well as by slightly ex-
tending the known boundaries of inhabited areas.

However, in present times zoogeographers must be very
careful in taking localities for granted even if the determina-
tion can be checked; for since World War II soldiers and
tourists often have thrown away shells in areas other than
where they had collected them, seamen emptied ballast tanks
on remote coasts, and dealers and natives indicated wrong
localities to gain higher prices. Therefore the Central Pacific
species collected as beach shells on the West Coast of Amer-
ica have been rejected in the present paper as no living speci-
men has ever been collected here. Even single living speci-
mens cannot exclude the possibility of having been intro-
duced by man (for references see ScHILDER, 1965, p. 171).

The present paper does not consider often uncertain
details but aims to give a survey about the general rules
concerning the distribution of living cowries and to com-
pare similar distributions in species of different descent
and in taxa of different evolutionary status.

Its aim is to encourage similar investigations in other groups
of marine mollusca.

TAXONOMY

Species and subspecies have been treated in the same way
as the classification of these taxa is often dubious and the
rules of distribution seem to be hardly influenced by the
present phase of evolution of species.

Intraspecific taxa have been considered to be of subspecific
rank (prospecies and subspecies) only if they are well sepa-
rable by morphological characters without knowledge of the
habitat of the specimens; infraspecies have been treated only
if their distribution is discontinuous; clines and local mu-
tants mostly have been ignored (see ScHILDER, 1966).

The customary citation of generic names does not make
clear the exact position of the species in Cypraeidae to a
reader not well acquainted with the phylogenetical sys-
tematics of the family. Therefore I prefer to replace the
generic names by 3-digit numbers according to my paper
on the generic classification of cowries (ScHILDER, 1968);
the first figure indicates the subfamily, the second indi-
cates the tribus (infrafamily), the third indicates the (in
1968 not numbered) genus; subgenera have been ignored
so that 4-digit numbers can be avoided.

The figures indicate the following taxa of living Cypraeidae:

4 Bernayinae 62 (Cypraeovulini)
42 (Bernayini) 622 Cypraeovula
421 Bernaya 623 Notocypraea
424 Zoila 624 Unmbilia

426 Siphocypraea 63 (Erroneini)

5 Cypraeinae 631 Erronea

51. (Cypraeini) 632 Notadusta
512 Trona 633 Palmadusta
513. Macrocypraea 634 = Bistolida
514 Mauritia 635 Ovatipsa
515 Talparia 636 = Cribrarula
516 Cypraea 7 Erosariinae

71 ( Pustulariini)

517 Lyncina = :

50 (meen) 711 Pustularia

OME GEO aiaes 714 Propustularia

BOE z d yp 72. (Erosariini)
Case 721 Monetaria

6 _Erroneinae TOD Naan

61 (Zonariin1) 723 ~+Erosaria

612 Schildera 724 Staphylaea

613. Zonaria 725 Nucleolaria

The citing of authors of specific names in the text
might distract the reader’s concentration on the zoogeo-
graphical facts; these names can be found in the ap-
pended list of living cowries.

ZOOGEOGRAPHICAL CLASSIFICATION

There is a surprising symmetry in the natural arrange-
ment of littoral territories with regard to the distribution
of living cowries.

Page 368

THE VELIGER

Vol. 11; No. 4

Hemispheres

The boundaries between the western and eastern hemi-
spheres lie in the eastern Pacific between Easter Island
and the Galapagos Islands, about at Walvis Bay in South
West Africa, and in the Suez Canal. They are never
crossed by any living cowrie species.

For the Indopacific 523 isabella and its West American rep-
resentative 523 mexicana should be regarded as distinct spe-
cies rather than as prospecies, and the pair 523 lurida (Med-
iterranean) and 523 pulchra (Red Sea) is still far less allied
each to the other, as these species belong to different sub-
genera.

Even on the generic level the hemispheral separation
becomes evident as only 3 genera are common to both
hemispheres, viz. 523 (Luria), 612 (Schilderia the Pacif-
ic species of which possibly should be separated gener-
ically), and 723 (Erosaria); 5 genera (425, 512, 513
613, 714) are restricted to the western hemisphere,
whereas the remaining 21 genera live in the eastern
hemisphere only.

On the tribus level the separation of the hemispheres is
much reduced as 6 tribes occur in both hemispheres and only
2 (62, 63) are restricted to the eastern one. All 4 subfamilies
live in both hemispheres.

Latitudinal Zones

In both hemispheres one can distinguish a broad tropical
zone along the equator and two less extended and usually
less warm marginal zones accompanying it in the north
and in the south.

Many tropical cowries extend far beyond the equatorial
zone into remote areas, while only few obviously northern or
southern taxa have also been discovered in scattered locali-
ties situated in the equatorial belt.

In the two southernmost regions inhabited by cowries
live 4 genera all species of which are restricted to these
relatively cold waters.

They are 622 (Cypracovula) in South Africa and 424 (Zoila),
623 (Notocypraea), and 624 (Umbilia) on the south coast of
Australia. In these 4 genera there is a curious splitting into
many allied sympatric species, Umbilia excepted in which the
splitting into many south Australian species partially of unique
features (624 gastroplax McCoy) took place in Tertiary
times, as it was also in Zoila (424 platypyga McCoy, 424
gigas McCoy). Along the other borders of the territory in-
habited by living cowries no similar unusual fauna will be
found: the splitting of 613 (Zonaria) in Senegal and of 612
(Schilderia) in Japan is far less striking.

Longitudinal Districts

Moreover, in either hemisphere 3 longitudinal districts
can be distinguished: a western, a central, and an eastern
district.

Their boundaries are caused by the present coasts of the
continents.

In the western hemisphere the 3 districts are distinctly
separated with regard to the cowrie fauna: there are un-
surmountable boundaries between West America, East
America, and West Africa (the Mediterranean Sea in-
cluded), as no cowrie species crosses the Panama Canal
nor the half way line along the Atlantic.

For the West American 513 cervinetta and the East A-
merican 513 cervus can be well separated at least as pro-
species, and the intermediate status of 723 spurca sanctaehe-
lenae between the East American 723 acicularis and the West
African 723 spurca needs confirmation.

In the eastern hemisphere, however, the 3 longitudinal
districts gradually pass each into the other as many cowrie
species occur far beyond the borders of the western half
of the Indian Ocean, the central district from Japan to
Australia, and the small islands of the Central Pacific.

There are plenty of species ranging from East Africa to
Polynesia, but there are also many species limited to one of
these 3 districts, e. g., 633 diluculum in the West, 633 lutea
in the middle, and 517 schilderorum in the East.

Central and Peripheral Regions

From the combination of latitudinal zones and longitu-
dinal districts a net-like pattern results in which a cent-
ral region is surrounded by at least 9 peripheral regions
in either hemisphere. In the more extended eastern hem-
isphere the central region is far more spacious than the
peripheral regions which look like appendices to the vast
Indopacific region. The 4 corner regions (South Africa,
Red Sea plus Persian Gulf, Tuamotu, Hawaii) are more
important faunistically than the border regions between
them, Japan and the south coast of Australia excepted.

Among the many cowrie species which live across the
tropical Indopacific from East Africa to Polynesia rather few
species inhabit also the 4 corners — Natal, Red Sea, Tuamotu,
Hawaii: 514 mauritiana, 515 talpa, 517 lynx, 517 carneola,
523 isabella, 721 moneta, 723 helvola, 725 nucleus; however,
only 517 carneola, 523 isabella, 721 moneta, and 723 helvola
reach the south coast of South Africa and the most widely
spreading cowrie species 517 carneola reaches also the Per-
sian Gulf, but none of them reaches the south coast of
Australia (the most western parts excepted) nor New Zea-

Vol. 11; No. 4

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Page 369

land. The other widely spread species are absent in at least
one corner region, or replaced there by closely allied, but well
distinguishable species or races.

In some species there is a tendency to produce similar
clines along the whole periphery of the inhabited area.

So, e.g., in 514 arabica (Scumper, 1961) and 516 tigris
the general size increases distinctly from an Indopacific cen-
ter to all border areas, and in 631 errones (SCHILDER & SCHIL-
DER, 1968) several characters show gradual peripheral changes.

CONTINUOUS DISTRIBUTION

The primary mode of distribution of a monophyletic
natural taxon undoubtedly is continuous.

The inclusion of very small areas in which a taxon cannot
live on account of unsuitable environments does not contra-
dict the term ‘continuity of distribution’.

Monotypic Taxa

Continuous distribution can be observed in taxa restrict-
ed to very small areas as well as in species which range
from East Africa to Polynesia without developing mor-
phologically distinguishable races.

Examples of such Indopacific species are 514 mauritiana,
517 vitellus, 633 asellus, 634 teres, 721 moneta, 725 nucleus.
In such species we have tried 30 years ago (ScHILDER &
ScuiLper, 1938/1939) to distinguish widely distributed geo-
graphical races (infraspecies) by the sum of variable minute
characters; the distribution of these minor taxa is continuous
and rather coincides with similar taxa separated by distinct
gaps in discontinuous Indopacific species. However, as we
have examined far more populations of these species for these
30 years, we now doubt the reality of many infraspecies and
we consider them to represent indistinct clines at most.

In some rare species the few known localities seem to
be separated by larger vacant areas, but future research
most probably will fill these gaps.

So, for instance, 633 gracilis and 633 saulae evidently in-
habit the same territory from Japan to Australia and from the
Gulf of Bengal to the westernmost Pacific; within this terri-
tory the common 633 gracilis has been found in 36 “areas”
(according to ScHILDER & ScuiLpeER, 1965, p. 173, paragraph
6), but the rare 633 saulae sofar only in 13.

The size of the inhabited territory of a species varies
from 1 to 80 “areas” as distinguished by ScHILDER (1965,
pp. 173-175, to which the area “54a: Auckland and
North New Zealand, 10-16 centigrades” should be
added).

Many very rare species are known from one area only, and
even 631 vredenburgi of which many hundred beach shells
have been collected in southern Java, just reaches the border
of a second area. The most widely distributed species is the
Indopacific 517 carneola which inhabits at least 80 areas.

Even if the arrangement of coasts and islands would
allow a rather circular expansion of the boundaries of
a species, the real distribution often consists in a rather
narrow band longitudinally or latitudinally.

Rather circular areas are inhabited, e. g., by 723 boivinii,
631 ovum, and 631 errones (all 3 in Malaysia, but with in-
creasing diameter), 514 immanis (western Indian Ocean),
631 bregeriana (Melanesia), and 634 goodallii (Polynesia).
Linear distributions are either longitudinal as in 613 robertsi
in West America and 512 stercoraria in West Africa (both
avoiding the off-shore islands), 631 felina and 723 margina-
lis on the west coast of the Indian Ocean, 631 xanthodon in
East Australia (common on the coast, rare on the off-shore
islands, absent from Lord Howe Islands), 723 guttata on the
west coast of the Pacific, and 725 granulata (cassiaui in-
cluded) in Polynesia; or latitudinal as 426 mus along the
north coast of South America, 723 gangranosa in the north-
ern Indian Ocean, 723 maturata (marielae included) in the
northern Pacific, and 636 fischeri on its southern border; the
distribution of 631 pallida and 723 ocellata could be called
more oblique as these species range from the Persian Gulf to
southern Malaysia.

Polytypic Taxa

Continuous distribution is necessary in species developing
clines especially in less warm areas (ScHILDER, 1966, p.
184).

Thus 723 caputserpentis gradually passes from the tropics
into the South-East-Australian cline caputanguis, 723 erosa
in the same region into the cline pulchella, and 723 helvola
into the South-African cline meridionalis.

But continuous distribution often can be found also in
species that can be divided into 2 or more morphologi-
cally well distinguishable geographical subspecies or even
in pairs of allied species constituting a superspecies
(ScuiLper, 1966, p. 182) ; their territories are separated
along a common boundary line which mostly runs longi-
tudinally.

This common case may be illustrated by pairs of allied
taxa which have been arranged according to their boundary
line being displaced from west to east (ScHILDER, 1965, pp.
173-175). The first named partner occupies the larger ter-
ritory or is obviously the more primitive taxon, the second
may be called peripheral; an asterisk (*) marks pairs of
undoubted specific rank; for the inhabited regions see the
appended list or more accurately in Scuttper, 1965 (pp. 176
to 183):

Page 370

THE VELIGER

Vol. 11; No. 4

* 523 isabella/mexicana

* 613 annettae/spadicea
513 cervinetta/cervus

* 523 cinerea/lurida

723 acicularis/spurca

631 listeri/felina

723 redimita/lamarcku

723 erosa/nebrites

634 stolida/erythracensis

723 marginalis/ocellata

723 miliaris/redimita

631 pallida/vredenburgi

631 cylindrica/sowerbyana

* 494 thersites/friendiu

* 624 hesitata/armeniaca
631 subviridis/dorsalis
631 walken/bregeriana
633 humphreysiu/lutea
517 schilderorum/kuroharai

kk OK Ox

723 marielae/maturata
* 723 caputserpentis/caputdraconis
* 723 scarabaeus/englerti
725 granulata/cassiaui
Also the tripartite species can be added:
514 arabica/immanis/grayana
631 adusta/onyx/melanesiae

However, in some pairs the adjacent territories do not
meet along a common boundary line, but they distinctly
overlap in a narrow zone, in which occasionally hybrids
may be observed:

* 613 zonaria/picta
613 pyrum/petitiana (hybrids? “senegalensis ScHiL-
DER”)
* 516 tigris/pantherina (hybrids)
* 517 vitellus/camelopardalis
* 723 lamarckiu/turdus
723 miliaris/eburnea (hybrids)

Rarely is the continuous area of Indopacific distribu-
tion composed of more than 3 taxa; on the subspecific
level such series of taxonomic units may be classified as
clines, but on a higher level they become real subspecies
or even well separable species which replace each other
along definite boundary lines or slightly overlap at their
borders.

In 523 isabella 3 clines and a border species can be distin-
guished by the black terminal spots increasing from west to
east: * isabella, lekalekana, controversa, and * mexicana.
On the other hand, in the series 723 * lamarcku, redimita,
* miliaris, eburnea the brown pigmentation decreases from
west to east.

The areas of peripheral species often become almost
covered by widely spread central allies so that a very
small corner remains for the sole presence of the former.

So, for instance, 523 pulchra lives also in the Persian Gulf
where the Indopacific 523 isabella is absent, while in the

whole Red Sea both live side by side; 633 artuffeli extends in
Japan farther to the north than its widely distributed ally
633 clandestina; and 634 subteres seems to be an analogous
taxon in the Tuamotu Islands with regard to its Indopacific
ally 634 teres.

In rare extreme cases the area of the peripheral species
has been quite covered by its widely distributed ancestor
species, so that both become sympatric in the whole terri-
tory of the former.

Thus 513 cervus is restricted to the northernmost border
areas of its ancestor 513 zebra, and 515 exusta is restricted to
the southern Red Sea, while the Indopacific 515 talpa crosses
it as far as Sinai.

WAYS to DISCONTINUITY

In some latitudinally or longitudinally tripartite species
the two border taxa are more similar to each other than
to the immediately adjacent central taxon; there is
the question whether the two border taxa are farther
developed in a parallel way than the older central taxon,
or, on the contrary, the latter is the younger taxon arisen
in the central area and pushing back the border halves
of the original taxon. Morphological aspects seem to favor
the latter interpretation.

Such cases may be illustrated latitudinally by 633 gracilis
of which the northern and southern clines japonica and ma-
cula are hardly distinguishable, while the central gracilis is
different at least statistically; the Mediterranean 613 pyrum
and the hardly separable unique angolensis from southern An-
gola are connected by the morphologically distinct tropical
petitiana. Longitudinally the western 631 adusta becomes sep-
arated from the very similar melanesiae by the central onyx
differing in color, at least, and the western 635 variolaria from
the eastern chinensis by the north Indian coloba. On the spe-
cific level one could add 523 isabella between pulchra and
mexicana, both with large terminal spots and brown pigment
on the teeth or margins, respectively; and possibly also 514
eglantina between histrio and maculifera.

The rare cases of several minor taxa of a species sur-
rounding the often overlapping central territory occupied
by another less closely related species may belong to the
same category.

The discontinuous races of 631 pulchella (viz. vayssierei,
pericalles, pulchella, novaebritanniae) and of 723 cernica
(viz. cernica, viridicolor, tomlini, maturata, marielae) seem
to be grouped around the central territory occupied by 631
pyriformis and 723 labrolineata, respectively.

In some few cases of now discontinuous distribution
the at present existing gap can be filled by extinct an-
cestors so that the distribution becomes continuous for
the scientist familiar with both living and fossil species.

Vol. 11; No. 4

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Page 371

ee

The rare Pleistocene 631 semicostata ScH1LpER from Java
seems to connect the living races of pulchella just mentioned,
and the recent pyriformis which is more frequently repre-
sented in the same beds (propyriformis ScHILDER). The now
discontinuous genus 613 (Zonaria) becomes continuous by
considering fossil species, as the living species 613 zonaria
(West Africa) and 613 aequinoctialis (plus annettae, both in
West America) were connected in Miocene times by the East
American 613 raymondrobertsi Pirspry and 613 bowdenen-
sis Prtspry. Possibly one could add on a still higher level the
3 relic species of 421 Bernaya, viz. teulerei (Arabia), fultoni
(South Africa), and catei (West Australia) which live about
in corners of the eastern Tethys region occupied by their
many Eocene ancestors in 421 (Bernaya); even 426 mus
whose Neogene ancestors were widely distributed in North
and South America could be regarded as a fourth corner relic
of the vast distribution of 42 (Bernayini) along the Paleo-
gene Tethys Sea.

DISCONTINUOUS DISTRIBUTION

According to the generally adopted rules of evolution
and dispersion of all living beings there is no primary
discontinuous distribution, as all taxa originated from
single populations inhabiting a small continuous territory.
Therefore the discontinuity is always secondary, in cowries
1. if members of a population have been displaced
a) naturally in larval stages by currents, etc. or
b) accidentally by man with his ships and hydroplanes
to remote areas where the species could gain a
footing; or
2. if the discontinuity is the result of extinction of con-
necting taxa in past geological times.

The discontinuity also may be apparent only if it is caused
by still insufficient knowledge of recent faunas and if there
is some hope to discover geographically connecting taxa es-
pecially in the as yet less explored deep waters.

The distribution of living cowries is rather well known,
though future research undoubtedly will fill many gaps
and correct many boundary lines. So the specialist is the
more surprised by the relatively large number of evidently
real discontinuities in distribution revealed by critical ex-
amination of all available indications of habitat.

Some isolated discoveries of single specimens often said to
have been collected alive in remote areas will be excluded
from the following considerations; they need further confir-
mation, or may rely on specimens accidentally introduced
by man. Such cases include 631 errones and 723 miliaris in
East Africa, 514 maculifera, 517 nivosa, and 631 barclayi in
Lemuria, etc.

There are two prevalent directions of geographical
discontinuity in living cowries:

1. latitudinal: two allied taxa live in the north and in
the south of an equatorial belt in which no related
taxon occurs;

2. longitudinal: two related taxa live in the west and
in the east along the equatorial belt, but they are
separated by a central region uninhabited by any re-
lated taxon.

Of course, latitudinal discontinuities refer only to pairs of
taxa living in about the same longitudinal district. The only
non-equatorial longitudinal discontinuity exists along south-
ern Asia between 631 pericalles (Persia) and 631 pulchella
(China).

Latitudinal Discontinuities

Some pairs of taxa, from allied species to hardly discern-
ible infraspecies, live in opposite areas of the northern
and southern zone without being connected geographic-
ally by a related equatorial taxon. One could call this fact
“bipolarity” in a broad sense, as it probably is caused by
parallelism of the areas concerning temperature. There-
fore one must bear in mind that future research may
supply connecting links in very deep waters of the equa-
torial zone. Such bipolarity has been observed almost
only in 6 (Erroneinae).

An accumulation of such “bipolar” pairs recently has been
discovered to exist in the Japanese and East Australian faunas:
612 langfordi — moretonensis
* 612 hiraset — queenslandica
631 hungerfordi — coucomi
631 piscatorum — subviridis
* 632 katsuae — hartsmithi
723 maturata — tomlini
but it occurs also in the central Pacific (Hawaii - Melanesia) :
634 latior — rashleighana
636 gaskoini — fischeri
as well as in West Africa (Mediterranean - Angola) :
612 achatidea — inopinata
613 pyrum — angolensis (here the gap is filled by the
less closely allied petitiana, see above)
and in West America (Gulf of California - Ecuador) :
613 annettae — aequinoctialis.

In a few pairs a northern or a southern taxon is re-
placed by an allied cowrie in the equatorial zone; how-
ever, as the boundaries of the equatorial partners are
still uncertain on account of their rarity, these cases also
may be interpreted as continuous distribution:

631 pulchella (China) — novaebritanniae (Melanesia)
631 xanthodon (East Australia) — fernandoi (Phil-
ippines)
633 hammondae (Australia) — rayswmmersi (Malay-
sia)
one could add from the past:

Page 372

THE VELIGER

Vol. 11; No. 4

* 424 friendiu (West Australia) — kendengensis ScuiL-
DER (Pleistocene of Java)

Longitudinal Discontinuities

In many widely distributed taxa a distinct discontinuity
between a western and an eastern branch can be observed,
comparable to the north-southern “bipolarity” discussed
above. This “bicentricity”, however, cannot be explained
by climatic reasons, but we must suppose that a continu-
ously distributed species has become extinct in its central
territory so that the geographically separated halves even-
tually could develop in different ways.

In a few cases a slightly less closely allied taxon still fills
the area between the two very similar border taxa or an
extinct central taxon has been discovered as fossil (see above).

In the western hemisphere the longitudinal range of
species is too small so that such longitudinal discontinuity

* 613 aequinoctialis (Ecuador) — zonaria (West Africa) ;
the gap in East America is filled by Tertiary species (see
above) .

In the eastern hemisphere, however, many discontinu-
ities have been discovered by accurate research on trust-
worthy localities. These discontinuities occur chiefly in
widely distributed pairs of rather common species sep-
arating an Indian and a Pacific taxon. The central area
devoid of any member of the pair is not identical in any
of the cases, but it changes from India to Central Mal-
aysia.

In the following lists the vacant area has been named be-
tween arrow heads ( > <) separating the first named west-
ern taxon and its eastern ally.

Such discontinuities occur on the infraspecific level, 1.
e. the two partners cannot be distinguished morphologi-
cally from each other; nevertheless one must expect that
the partners separated for a long time have developed
hereditary differences by different selection of genes,

can be shown only in which possibly could be discovered by statistical methods:

631 amabilis > India < walken
724 interstincta > Andamans-Sumatra <_ limacina
635 variolaria > Ceylon-Borneo <_ chinensis
522 ingens > West and Central Malaysia < testudinaria
711 lemurica > southern Malaysia < children
514 dispersa > central Malaysia < depressa
723 poraria > central Malaysia < scarabaeus ,
631 adusta > Malaysia <_ melanesiae
The differences have been intensified to true subspecies level in:
514 alga > Maldives, India < mappa (incl.
geographica)
633 notata > India < gracilis
633 chrysalis > India, Sumatra < microdon
631 pericalles > India, South Malaysia < pulchella
514 scurra > central Malaysia < indica
Rarely pairs of equal specific rank occur:
* 723 macandrewi > India to Celebes < becku
* 514 histrio > Malaysia, New Guinea < maculifera

To these examples of discontinuous distribution one can add 2
pairs of species, each pair living in restricted parts of the Indo-

pacific:
* 517 broderipu > Lemuria < _nivosa in the
Indian Ocean and
* 723 boivinit > Micronesia

< ostergaardi in the
Pacific

Vol. 11; No. 4

THE VELIGER

Raversio

SYSTEMATIC LIST

These general observations may be followed by a system-
atic list of living species, (p) prospecies, (s) subspecies,
(c) clines, (i) infraspecies, and (m) local mutants (ScHIL-
DER, 1966). The present list shows far less details in
distribution than the list published 4 years ago (ScHILDER,
1965, pp. 176-183), but it isa clearer survey of the
general distribution over the greater regions.

These “regions” have been designated in the former list
(Scumper, 1965) by 2-digit numbers and 3 capital letter-
abbreviations; for augmenting the clarity in a more concen-
trated table I now prefer to return to the 1-capital system
used in a previous paper (ScuiLpER, 1941 with partially
modified regional capitals and limits). As there are 30 regions
at least, some capitals must be used twice; therefore the less

‘numerous regions of the western hemisphere have been print-
ed in italics, and the western species have been marked with

lok

The following alphabetical list of regions contains the
abbreviation (one capital letter), the name of the re-
gion, and its former designation with 2 digits. The Map
shows the boundaries of these regions.

WESTERN HEMISPHERE

A Antillean 81 H St. Helena 85

B Brasilian 86, 73 L Lusitanian 96

C Californian 89, 35 M Mediterranean 95
E Ecuadorian 87 P Panamic 88

F Floridan 82 S Senegal 83

G Guinean 84

EASTERN HEMISPHERE

East African 18
South African 17
Dampierian 15
Erythraean 19
Fijian 46

M Malayan 48

N New Guinean 41

P Persian 12

Q Queenslandian 47
(including 54h = Lord Howe Is.)

PoaHmoOAmMpaAS

Guam (Micronesian) 42 R Rapanuian 45

Hawaiian 43 S Sumatran 14

Indian 13 T Tahitian 44

Japanese 49, 37 V Victorian 58
Z

Zealandian 54
(h excluded)

Lemurian 11

In the list of valid taxa of living cowries the regions
inhabited by each taxon have been enumerated according
to my previous arrangement (ScHILDER, 1965, pp. 173
to 175) as follows:

CPEFABMLSGH
CALEPISDVZQFNMJGTRH

western ((_]):
eastern:

If it appears to be necessary to emphasize the restric-
tion to a part of a region, the nine digits used in my
former paper (ScHILDER, 1965, p. 171) will be used as
exponents:

Sie NW 2 N oa NE
8 — W ™ — central AOE
7 = SW 6—S§ SSE;

The sign || marks greater discontinuities within the taxon.

Page 374
CYPRAEIDAE Gray, 1824
4 Bernayinae ScHILDER, 1927
42 (Bernayini ScuiLpEr, 1927)
421 Bernaya JoussEAUME, 1884
teulerei (CAZENAVETTE, 1846) lp Je
fultoni (SowerBy, 1903) C?
catei SCHILDER, 1963 D’
424 Zoila JouUSSEAUME, 1884
decipiens (Smiru, 1880) D”
venusta (SowErRBy, 1846) D’ V®
thersites (GasKotn, 1849) V'
friendii (Gray, 1831) D’ Vv’
marginata (Gaskotn, 1849) DY WY
rosselli (Cotton, 1948) We
426 Siphocypraca Herrin, 1887
[] mus (Linnaeus, 1758) A
¥) Cypraeinae Gray, 1824
51 (Cypraeini) Gray, 1824
512 Trona JoussEAuME, 1884
L] stercoraria (LinNAEuS, 1758) S*+G

513. Macrocypraea Scuitper, 1930
L] zebra (Linnatus, 1758) FAB
[J cervus (Linnaeus, 1771) je “EC
[] — (p)cervinetta (Krener, 1843) GP IP ja
514 Mauritia TroscHeEL, 1863
valentia (PERRY, 1811) Q? N?
mappa (Linnaeus, 1758) D?Q? FNM J°GT
—(c) geographica SCHILDER & SCHILDER, 1933 S
—(s) alga (Perry, 1811) INE TL, TBP
arabica (LinNaEuS, 1758) ISDQFNMJGT?

—(m) gibba Corn, 1949 M®
— (s) ¢mmanis SCHILDER & SCHILDER, 1939

CP AN IL
grayana SCHILDER, 1930 AO 18, 12 IP
eglantina (Ductos, 1833) S*>DQFNMJ°G
histrio (GmeEtin, 1791) ALISD
maculifera Scuitper, 1932 (L'?)||F'J°GTH
depressa (Gray, 1824) He Neves le Gang

— (i) dispersa SCHILDER & SCHILDER, 1939
AES
mauritiana (LINNAEUuS, 1758)
(Cz) PASIEN ES ISS DEO EeNe MiG alve Et
scurra (GMELIN, 1791) AX Mb, IS

THE VELIGER

—(s) indica (GMELIN, 1791)
QFNM*)°>GTH
515 Talparia TRoscHEL, 1863
talpa (Linnaeus, 1758)
C A LE PS DO? FN Maia
exusta (SOwERBY, 1832) 18
516 Cypraea LinnaEus, 1758
tigris LINNAEUS, 1758
CALE ES DO EN Mi ijcGeeG
— (c) schildertiana Cate, 1961 Te
pantherina SoLANDER, 1786 E
517 Lyncina TroscHEL, 1863

aurantium (GMELIN, 1791) FNMGT
broderipu (SOWERBY, 1832) (Ee (Le?)
nivosa (BRopERIP, 1827) (ie? TESS?
leucodon (Bropverip, 1828) (14?) Ms

argus (LINNAEUS, 1758)
ALIS D OE MeGa
portert (CATE, 1966) Mea
lynx (Linnaeus, 1758)
GAL ELS Di OQ FN MejsGwiar
vitellus (LINNAEUS, 1758)
GCALETSD V2 ZO FIN Maj Caria

camelopardalis (Perry, 1811) E°
reevel (SOWERBY, 1832) Ve
ventriculus (LAMARCK, 1810) EN MM ¢
schilderorum (IrREDALE, 1939) FGTiH
—(s) kurohara (Kuropa & Hasse, 1961) J
sulcidentata (Gray, 1824) H

carneola (LinNAEUS, 1758)
GCALEPIS DZ OE NeMaiiGaiee

—(m) titan ScHILDER & ScHILDER, 1962 A L’

—(m) leviathan (SCHILDER & SCHILDER, 1937) H

52 (Luriini) ScHILper, 1932

522 Chelycypraca SCHILDER, 1927
testudinaria (LINNAEUS, 1758)
DIO_E N MesiiGal
— (i) ingens ScHiLpeR & ScuitpeR, 1938 A'LI°
523 Luria JouSSEAUME, 1884
tessellata (Swainson, 1822) H
pulchra (Gray, 1828) EP
isabella (LInNAEuS, 1758) CALE TI
— (c) lekalekana (Lapp, 1934)
s) 1D) 22 (QO) Ja IN| WME J) © WE
—(c) controversa (Gray, 1824) H

L] mexicana (Stearns, 1893) GP IP? ja?
[_] cinerea (GMELIN, 1791) FAB
[] lurida (Linnaeus, 1758) IGE Ie SS (GE Sel

Vol? 11; Now:

———

Vol. 11; No. 4

6
61

Erroneinae ScHILDER, 1927

(Zonariini) ScHILDER, 1932

612 Schilderia Tomutn, 1930

[] achatidea (Sowersy, 1837) M®
[] -(i) inopinata ScuitpeEr, 1930 G®
langfordi (Kuropa, 1938) J
— (i) moretonensis ScHILDER, 1965 Q'
hiraset (Roperts, 1913) J
queenslandica ScHILDER, 1966 Q'
teramachu (Kuropa, 1938) i

midwayensis (AzUMA & Kurowara, 1967) HH?

613 Zonaria JousSEAUME, 1884

62

L] gambiensis (SHaw, 1909) S4
L_] zonaria (GMELIN, 1791) I We (G
L] picta (Gray, 1824 ) S
[_] sanguinolenta (Gmeun, 1791) St
L] pyrum (Gmeun, 1791) M L S*
L] - (i) angolensis (OpHNER, 1923) G®
L] —(p) petitiana (Crosse, 1872) S* G'
|] annettae (Dat, 1909) Gi
_] — (p) aequinoctialis ScuttpER, 1933 E*
L] spadicea (Swainson, 1823) (OR
L] roberts: (Hmatco, 1906) GE IP 1d:
[_] arabicula (Lamarck, 1810) GRBE
[_] nigropunctata (Gray, 1828) E

(Cypraeovulini) ScriLpEr, 1941

622 Cypraeovula Gray, 1824

fuscorubra (SuHaw, 1909) c*
fuscodentata (Gray, 1825) Ce
cohenae (Burcess, 1965) Cc’
algoensis (Gray, 1825) C®
edentula (Gray, 1825) C
amphithales (MELvILL, 1888) C
capensis (Gray, 1828) C
623 Notocypraca ScHILDER, 1927
pulicaria (Reeve, 1846) Vv’
bicolor (Gasxoin, 1849) ve
— (c) reticulifera (ScutmLpER, 1924) Vv’
— (s) occidentalis IREDALE, 1935 ve
piperita (Gray, 1825) V OQ
—(c) compton (Gray, 1847) ve
—(m) casta SCHILDER & SUMMERS, 1963 \We
declivis (SowERBY, 1870) Vv
angustata (GMELIN, 1791) Vi
624 Umbilia JoussEAUME, 1884
armeniaca (VeERco, 1912) V'
hesitata (IrEDALE, 1916) W253 @x

THE VELIGER

Page 375

(Erroneini) ScHiLpErR, 1927

631 Erronea TroscHEL, 1863

walkeri (SowerBy, 1832) SDOQOMJ@
— (i) amabilis (JoussEAuME, 1881) 1b
—(p) bregeriana (Crosse, 1868) Fo Nt
pyriformis (Gray, 1824) It'sSoDQ’M
pulchella (Swainson, 1823) MES lor
— (i) vayssierei SCHILDER & ScHILDER, 1938 E*
—(s) pericalles (MELVILL & STANDEN, 1904) P®
—(s) novaebritanniae SCHILDER & SCHILDER, 1937

LINE
hungerfordi (SoweErRBy, 1888) J
— (1) coucomi ScHILpER, 1964 Q'
barclayt (REEvE, 1837) @? (2?)
xanthodon (Sowersy, 1832) Q
—(p) fernandoi Cate, 1969 M3
vredenburgi ScHILDER, 1927 S> M®
pallida (Gray, 1828) PE S* M2

— (c) insulicola SCHILDER & SCHILDER, 1938 M®

stohlert CATE & SCHILDER, 1968 M3
subviridis (REEVE, 1835) QF’
— (p) dorsalis ScHILDER & SCHILDER, 1938 D
— (i) piscatorum ScHiLper, 1965 J
onyx (LinNaEus, 1758) N° M J G
—(p) adusta (Lamarck, 1810) A L™® PTI 8

— (i) melanesiae SCHILDER & SCHILDER, 1937
FY N*
—(m) nymphae (Jay, 1850) 14
ovum (GMELIN, 1791) SDQ? NM f
errones (LINNAEUS, 1758)
(A?) | ISDQFNMJGT
— (m) azurea ScuILpER, 1968 D'
cylindrica (Born, 1778) SOQOFNMJG
— (s) sowerbyana ScHILDER, 1932 D
caurica (LinNAEuS, 1758)

CPAC Es Pale SD OpkeNeMes Gaal
felina (GMELIN, 1791) CAL’
—(c) fabula (Krener, 1843) 18? 12
—(p) listeri (Gray, 1824)

eee? SD ES Nie Many «Gah?
—(c) velesia (IREDALE, 1939) ve QO

632 Notadusta ScHILpER, 1935

punctata (LInNAEUuS, 1771)
ALS DY OLE N May (Gan

rabaulensis SCHILDER, 1964 N?
katsuae (Kuropa, 1960) M? J
hartsmithi ScHILDER, 1967 Q°

martini (SCHEPMAN, 1907)
—(p) superstes (SCHILDER, 1930) Fs

Page 376

633 Palmadusta IREDALE, 1930
asellus (LINNAEUS, 1758)
ALISDQFNMJGT
clandestina (LINNAEUS, 1767)
CALEISDOFNMJGT?
artuffeli (JoUSSEAUME, 1876) J@
saulae (Gasko1n, 1843) S° || D Q M3 J G
contaminata (SowERBY, 1832)
c’Li|SsSDOFNM jf
lutea (GmMeEtIN, 1791) (1°?) SD N° M J G°
—(p) humphreysu (Gray, 1825) QF N* G
ziczac (LINNAEUS, 1758)
CALEPISDQFNMJ GT
diluculum (ReEEvE, 1845) (CP AN 18?
— (c) virginalis SCHILDER & SCHILDER, 1938 L
lentiginosa (Gray, 1825) 1sy 1 II
gracilis (Gaskoin, 1849) IS F' NMG

—(c) japonica (ScuitpeEr, 1931) J
—(c) macula (Ancas, 1867) re)
— (s) trescens (SoweErRBy, 1870) D
—(p) notata (Git, 1858) A’ E P
hammondae (IrEDALE, 1939) Q N

— (i) dampierensis (SCHILDER & CERNOHORSKY,
1965 ) D
—(p) raysummersi (ScHILDER, 1960) mM”
fimbniata (GMELIN, 1791)
CALISSDFNMJG
—(c) unifasciata (MicHELs, 1845) TH
minoridens (MEeEtviLz, 1901)
S || QFN MJIGT
— (p) serrulifera SCHILDER & SCHILDER, 1938 T
microdon (Gray, 1828) SOFNM J
—(s) chrysalis (KieneEr, 1843) A’ L?® E#
634 Bistolida Cossmann, 1920
quadrimaculata (Gray, 1824)
SDQFNMJG
coxent (Cox, 1873) Fo Ns
— (s) hesperina (ScHILDER & SUMMERS, 1963)

Nrt 4 M3
interrupta (Gray, 1824) IS M*®
pallidula (Gaskon, 1849) M G
— (c) luchuana (Kuropa, 1960) J
— (i) rhinoceros (SouverBiE, 1865) DQEN
—(c) summers: (ScHILDER, 1958) F*
rashleighana (MELvIL1, 1888) Fe
— (s) latior (MELvILL, 1888) H

teres (GMELIN, 1791)

CALI SDQFNMJGTH
—(p) subteres (WeEINKAUFF, 1881) ae
goodallui (SoweErRBy, 1832) F3 G+ T
kieneri (Hwatco, 1906) Agile Tc

THE VELIGER

Vol. 11; No. 4

—(c) depriestert (ScHILDER, 1933)
SDQFNMJG

— (i) landent (ScHILDER & GriFFITHS, 1962) T°

owentt (SoweErBy, 1837) L

—(s) vasta (SCHILDER & ScutLpER, 1938) CA
hirundo (LINNAEUS, 1758)

A LEISDQFNMJGT
ursellus (GmMEtIN, 1791) SD QFNM J G@

erythraeensis (SOWERBY, 1837) E
stolida (LinNAkEus, 1758)
CALESDOEN MijgeGae
635 Ovatipsa IREDALE, 1931
chinensis (GMELIN, 1791)
SDOFNM*jJGTH
— (i) variolaria (LaMaRcK, 1810) CALE
—(m) tortirostris (SowERBy, 1906) C
—(p) coloba (MeEtvi11, 1888) lee | ISP
636 Cribrarula Stranp, 1929
cribraria (LINNAEUS, 1758)
1G a IOS DY WE O18 NW IME | G

—(c) comma (Perry, 1811) (CP A IL?
cribellum (Gasxkotn, 1849) A’ >
esontropia (Ductos, 1833) Ls
catholicorum (SCHILDER & SCHILDER, 1938)
Oe aNe
gaskoin (REEVE, 1846) H
—(p) fischert VayssizreE, 1910) is (IEP)
cumingu (SOWERBY, 1832) G: T
Erosariinae SCHILDER, 1924
71 (Pustulariini) Scuiper, 1932

711 Pustularia Swainson, 1840
mariae ScHILDER, 1927 Q? F N M3 J’ GT H
globulus (LinnaEus, 1758)
rPSDQFNMJG
— (i) brevirostris SCHILDER & SCHILDER, 1938
AuaE
mauiensis (BuRGESS, 1967) H
cicercula (LINNAEUS, 1758)
ALE|SDQFNMJGT
— (c) margarita (Dittwyn, 1817)
FN MGT
bistrinotata SCHILDER & SCHILDER, 1937
SDQFNMJGTH
childreni (Gray, 1825) 1) IN| WWE J) Ge IP Tel
— (1) lemurica SCHILDER & SCHILDER, 1938
1G (Q72)
714 Propustularia ScuiLpER, 1927

L] surinamensis (Perry, 1811) F° A B?

Vol. 11; No. 4

THE VELIGER

Page 377

72 (Erosariini) ScHriLpErR, 1924

721 Monetaria TroscHEL, 1863
moneta (LINNAEUS, 1758)
CALEPISDQFNMJGTH
annulus (LINNAEUS, 1758)
CALEPISDQFNMJGT
— (c) obvelata (Lamarck, 1810) TS
722 Naria Broperip, 1837

irrorata (Gray, 1828) F? N? M?G T
723 Erosaria TRoscHEL, 1863
dillwyni (ScuiLpvER, 1922) F* (G?) T

beckii (Gaskon, 1836) Q? N M‘J GT? H
macandrewi (SoweErBy, 1870) E P®
labrolineata (Gasxotn, 1849)

Ss DQFNMJGH

—(m?) ogasawarensis SCHILDER, 1944 G’
cernica (SowERBY, 1870) L*
—(s) viridicolor (CaTE, 1962) D
—(p) tomlini Scuitper, 1930 Z™ Q F* (T°?)
—(s) maturata (Kira, 1959) JG
— (i) marielae (Cate, 1960) H
citrina (Gray, 1825) Cc A® (L??)

L3 E° I S M®
S* M

gangranosa (Dittwyn, 1817)
bowini (KiENER, 1843)
ostergaardi (DALL, 1921) H
helvola (Linnaeus, 1758)
ALEISDV.QFNMJGTH

—(c) meridionalis ScHILDER & SCHILDER, 1938 C
caputserpentis (LInNAEUuS, 1758)

GALE ISDZ7> QO FNMJGT
—(c)kenyonae SCHILDER & SCHILDER, 1938 D’ V®

—(c) caputanguis (Putprt, 1849) Q®
— (i) caputophidi ScuiwpeEr, 1927 H
caputdraconis (MELVvILL, 1888) R
[_] albuginosa (Gray, 1825) GP 122 15
[] spurca (Linnaeus, 1758) MLSG
L] - (i)sanctaehelenae ScuiLperR, 1930 H
L] — (p) acicularis (GmeEuin, 1791) FAB
poraria (Linnaeus, 1758) ASI SED

— (i) searabaeus (Bory, 1827)
2ZOFNMJGTH
englerti (SUMMERS & BurceEss, 1965) R

erosa (LinnaEus, 1758)
CALISDZ2OFNMIJGTH
—(c) pulchella Corn, 1949 QO
—(p) nebrites (MELvILL, 1888) IX 1S, Jet 12
ocellata (LinNaEus, 1758) EPs

marginalis (Dittwyn, 1827) C A L? Et P®
miliaris (GMELIN, 1791)

(A?) |S) D Q N™» MJ G

— (p) eburnea (Barnes, 1824) OFN
lamarckii (Gray, 1825) CAL’?
— (s) redimita (MEtvitt, 1888) L**? P+ I S?®
turdus (LAmMarRcK, 1810) FXp 13, Ve
—(c) pardalina (DuNkER, 1852) 1)
— (c) winckworthi ScHILDER & SCHILDER, 1938

Pt
guttata (GMELIN, 1791) F? N M? J G’

724 Staphylaea JoussEauME, 1884
limacina (Lamarck, 1810)
SDV OPE Ne May G:
— (i) interstincta (Woop, 1828) OPN IAI
— (p) semiplota (MicueEts, 1845) H
staphylaea (LinnaEus, 1758)
CALE SDOQOFNMIJGTH
725 Nucleolaria Oyama, 1959
nucleus (LINNAEUS, 1758)
ALEFSDQOFNMJGTH
granulata (PrEasE, 1862) H
— (s) cassiaut (BurcEss, 1965) a te

LITERATURE CITED

ScuiLperR, Franz ALFRED
1941. | Verwandtschaft und Verbreitung der Cypraeacea.
Arch. Molluskenk. 73 (2-3): 57-120; 2 plts.
(15 May 1941)
1961. _A statistical study in cowries: the size of Mauritia ara-
bica (Linnaeus). The Veliger 4 (1): 15-17; 2 text figs,
(1 July 1961)
1965. The geographical distribution of cowries (Mollusca:

Gastropoda) The Veliger 7 (3): 171 - 183; 2 maps
(1 January 1965)
1966. Personal views on taxonomuy. The Veliger 8 (3):
181-189; 1 text fig. (1 January 1966)
1968. The generic classification of cowries. The Veliger

10 (3): 264 - 273
SCHILDER, FRANZ ALFRED, & MARIA SCHILDER
1938 - 1939. | Prodrome of a monograph on living Cypraeidae.
Proc. Malacol. Soc. London 23 (3/4): 119 - 231; 1 fig.; 9 maps
1968. Studies on populations of the cowrie Erronea errones
(LinnaEus). The Veliger 11 (2): 109-116; 5 maps; 6
tables. (1 October 1968)
ScHILpER, Maria
1967. Length, breadth, and dentition in living cowries. The
Veliger 9 (4) : 369 - 376; 1 diagram (1 April 1967)

(1 January 1968)

Page 378

THE VELIGER

Vol. 11; No. 4

Seasonal Observations on Diet,

and Stored Glycogen and Lipids in the Horse Clam,

Tresus capax (GouLD, 1850)

BY

ROBERT G. B. REID

Department of Biology, University of Victoria, Victoria, British Columbia, Canada

(Plate 57; 1 Text figure)

INTRODUCTION

A sTUDY OF TWO BIVALVES, Lima hians (GMELIN, 1791)
and Mya arenaria LINNAEUS, 1758 revealed the presence
of fat globules in the digestive diverticula of these ani-
mals (Rei, 1966). Although glycogen is generally re-
garded as the major food storage compound in the
Bivalvia, the fat contained in the digestive diverticula
represents a significant amount of food energy. It was
of interest therefore to undertake a seasonal study of
the diverticular lipid as it related to lipid and glycogen
storage in the gonad, which is the major organ of food
storage, in the hope that the following questions might be
answered: does the lipid of the digestive diverticula
increase with an increase in food, or is it kept constant
by the transport of excess lipid to the gonad? Does the
diverticular fat represent a final energy store, to be de-
pleted when the gonad glycogen and lipid are exhausted,
or does the level of diverticular fat fall along with the
level of glycogen and fat in the gonad? The seasonal
availability of food was relevant to these problems and a
study of the diet was undertaken along with stored food
observations.

MATERIALS anp METHODS

Specimens of Tresus (=Schizothaerus) capax (GouLp,
1850) were dug at Esquimalt Lagoon, near Victoria, B. C.
The average age of the animals was 4 years. They were
collected at each spring tide over the period of a year,
from July 1966 to June 1967. Adverse tidal and weather
conditions made it occasionally impossible to collect, and
sometimes limited the number of animals collected to two.
The maximum number of animals sampled at any col-

lection time was 6, though the general condition of the
gonads of all the animals dug was observed.

Bearing in mind the admonition of Mansour (1946)
that digestion of some of the diet components in bivalves
may be so rapid that any delay between collecting and
examination of the stomach contents might give a wrong
impression about diet, the stomach contents during the
first few collections were examined on the beach, minutes
after the animals were dug. It soon became obvious that
a delay of many hours made little difference to the quali-
tative aspects of the stomach contents, and subsequent
observations were made in the comfort of the laboratory.
The stomach contents of animals which were dug while
still covered with several feet of water during the ebb of
the tide were also examined to compare actively feeding
animals with those which had been exposed to the air
for some hours. Within 12 hours of collection of the ani-
mals plankton samples were taken in the vicinity of the
Tresus bed to determine the degree of selection exercised
by the animals.

Lipid from the digestive diverticula and gonad was
estimated by the chloroform/methanol extraction method
of ForcH et al. (1957). Glycogen was estimated by the
anthrone method of ViILES & SILVERMAN (1949) after
extraction with 5% trichloracetic acid at 90° C.

The digestive diverticula were usually invested with
gonad tissue and it was difficult to separate the two tissues
completely. This problem was surmounted thus: a previ-
ous histochemical study (Rem, 1968) indicated that gly-
cogen is absent from the digestive diverticula proper. Thus
the glycogen value obtained from the diverticula must
have come from investing gonad. Therefore, applying the
glycogen: lipid ratio obtained from the gonad proper, the
% dry weight of lipid derived from investing gonad

——.

Vol. 11; No. 4

THE VELIGER

Page 379

could be subtracted from the diverticular lipid value to
obtain the true lipid value.

It should be observed here that the histochemical ob-
servation on Tresus diverticular fat included in the author’s
1968 paper was false, due to a technical mishap, but
that the observations on the other species were valid.

RESULTS

Stored Lipid and Glycogen

The results presented in Table 1 are graphed in Text
figure 1, with different scales for average % dry weight
lipid (left hand ordinate) and average % dry weight gly-
cogen (right hand ordinate). Proceeding chronologically

Table 1

Percent dry weight of gonad lipid, diverticular lipid, and
gonad glycogen of Tresus capax, averaged for all
individuals collected.

* — sample lost through technical error]

Date Gonad Diverticular Gonad
Lipid Lipid Glycogen
(corrected value)
July 19 13.0 11.7 83.0
August 15 11.3 13.3 77.3
September 12 15.5 12.5 63.9
October 5 tS oe 63.9
November 16 14.0 9.5 52.4
December 1 15.7 S.0 44.2
December 15 18.0 10.2 33.2
December 28 13.3 10.2 3.2
January 9 13.7 10.6 3.4
January 24 11.8 8.8 10.4
February 7 14.2 8.3 19.5
February 22 13.3 10.6 19.8
March 6 8.1 10.0 2.8
March 23 8.7 6.5 6.8
April 13 12.8 7.2 lef
April 27 9.9 12.1 18.8
May 10 10.4 11.7 16.6
May 24 8.88 10.6 20.6
June 10 12.7 9.9 9.3
June 24 11.5 11.7 60.1

a falling off in glycogen begins about the end of July,
1966, the curve steepens in December and rapidly falls
to a minimum value at the beginning of January. Rising
temporarily during February it falls to a new low which
lasts throughout March and part of April. Then the
glycogen content begins to rise, interrupted by a slight

gonad glycogen

Co oO
o oO

a
°
average % dry weight glycogen

gonad lipid
digestive diverticular lipid 70

-_

oP OO WN
wo
°

tS)
fo}

average % dry weight lipid
a

°

yooAN S) © IW iD yp Pe SNe
Months

Figure 1

Graph of average % dry weight of gonad glycogen and lipid,
and digestive diverticular lipid, in Tresus capax, over the period
July 1966 to June 1967

dip in the beginning of May, to rise finally to the peak
in July.

Both digestive diverticular and gonadal lipid are fairly
similar in their progress. However, during November there
is a slight rise in gonad lipid to reach the highest peak of
the year, while the digestive diverticular lipid falls slightly.
Both organs reach a low in lipid content which coincides
with the second low in glycogen content, recovering in step
with the glycogen halfway through April. In May the
gonad lipid drops along with the glycogen, but the lipid
in the digestive diverticula shows less of a dip at the same
time.

Incidental observations on the general condition of the
gonads of all the animals collected are included in Table
2, along with the diet observations. From these observa-
tions it seems that individual animals vary considerably
during the period December to March in the amount
of stored material (i.e., in degree of fatness). Since an
average of 4 animals only was sampled at each collecting
time over the year it looks as if this number of animals
is too small to make the individual differences insignifi-
cant, and so the first low and the small peak in glycogen
content during February is probably not indicative of the
general trend throughout the population. Thus it is likely
that a larger sample would have the glycogen content of
the gonad falling smoothly from July to March.

Diet

Observations on diet and available phytoplankton are
detailed in Table 2. Typical summer and winter stomach
contents are shown in Plate 57. Naked flagellates are
almost impossible to identify after exposure to the gastric
juice of the clam, since they immediately become dis-
torted and the flagellae are no longer obvious. Wherever

Page 380

THE VELIGER

Vol. 11; No. 4

Table 2

Comparison of stomach contents of Tresus capax with available phytoplankton in the sea; notes on the general
condition of the gonads of all animals collected

Date Stomach Contents Phytoplankton not Ratio of fat:
in Stomachs thin: depleted
gonads
June 20 Many Fragilaria; colonial Navicula; various diatoms; detritus 0 Cpe 75 6 (0)
June 30 Mainly Fragilaria; detritus 0 T2210
July 18 Mainly Melosira 0 6:0:0
August 15 Various diatoms and flagellates 0 AS OO
August 27 Melosira; Meridion; many flagellates peridinians 5:0: 0
September 1 Various diatoms 0 6:0:0
September 12 Meridion; Gomphonema; Melosira; many flagellates peridinians 4:0:0
October 5 Meridion; Gomphonema; Melosira; many flagellates Coscinodiscus 780) 3
November 2 Various diatoms; flagellates Coscinodiscus as 2 3 0
November 16 A few, varied diatoms 0 73-2073 5 (0)
December 5 A few Melosira; flagellates Gyrosigma; Triceratium; Isthmia; 130
Chaetoceras; Coscinodiscus

December 15 Mainly detritus As December 5 Os 23%
December 28 Mainly detritus; a few varied diatoms; flagellates As December 5 Os 23 8
January 8 Mainly detritus; a few varied diatoms; flagellates peridinians; Coscinodiscus OFS eS
January 24 Mainly detritus; a few varied diatoms; flagellates Biddulphia arctica 0:2: 4
February 7 Mainly detritus; a few varied diatoms; flagellates Biddulphia arctica Ose
February 22 Mainly detritus; a few varied diatoms; flagellates 0 Oo a!
March 6 Many Biddulphia aurita 0 Os 032
March 22 Various diatoms Chaetoceras OF aie
April 15 Many varied diatoms 0 gl g 3
April 27 Many small diatoms 0 rece s al
May 10 Many Fragilaria; Melosira 0 0:2:0
May 24 Many Fragilaria; Melosira 0 0:2:0
June 10 Many Fragilaria; Melosira 0 15250
June 24 Many, varied diatoms 0 sy 8 W

possible diatoms are identified to genus and occasionally
to species.

In summary, there are a number of larger diatoms
(with large cell bodies, or made effectively large by pos-
sessing many long spines) such as Coscinodiscus, Chaeto-
ceras, and Isthmia species, and Biddulphia arctica
(BRIGHTWELL, 1853) which commonly occur in the re-
gion of the clam beds, but which the animals do not in-
gest, presumably because of their large size, being selected
out and rejected by the sorting mechanisms of the mantle
cavity. At times there are also large peridinian dino-
flagellates which are available to the clams, and which
would seem to be within the upper limits of the accept-

able particle size (150 u), but which are not ingested,
for reasons unknown. For the most part Tresus ingests all
of the other smaller phytoplankton members, any bloom
of a particular species being reflected by the stomach
contents. Some time in November the amount of the food
in the stomachs was noticeably reduced from the earlier
part of the year, and from December to the end of
February most of the particulate material in the stomach
was inorganic or detrital. As early as March 6 the first
phytoplankton bloom, of Biddulphia aurita (LyNcBYE,
1819) appeared, and the quantity of phytoplankton
found in the stomachs gradually increased into June
where it seemed to level off.

Explanation of Plate 57

Figure 1: Stomach contents of Tresus capax, collected July 1966;
various diatom species; flagellates

Figure 2: Stomach contents of Tresus capax, collected December
1966; several flagellates; diatom frustules; Cristispira

[ReEr] Plate 57

ba
5
7
S
>
a
(<2)
Q
a
>
|
rr
i

Vol. 11; No. 4

THE VELIGER

Page 381

The stomachs of all the animals examined contained a
species of the spirochaete genus Cristispira. These bac-
teria were most abundant during the summer. Currently
work is underway to determine the role of the spiro-
chaetes in the alimentation of Tresus.

DISCUSSION

When this work began the unwarranted assumption was
made that adequate information would be available on
the biology of Tresus. Unfortunately, most of the work on
Tresus has been ecological, and the only relevant work is
that of Swan & Finucane (1953) in which the authors
imply that Tresus capax is a winter-spawning species. Dr.
Quayle of the Biological Station, Nanaimo, in a personal
communication informs me that the condition of the
spat also indicates that T. capax is a winter-spawner. My
lipid and glycogen analysis, together with the observations
of gonad condition are only explicable in terms of T.
capax being an early spawner, but it would have been
more satisfactory to have done a seasonal histological
study of the gonads along with the other aspects of
the work.

_ The preliminary questions which this work set out
to answer are satisfied, with respect to this species. ‘The
amount of fat in the digestive diverticula is relatively
constant, falling from a summer level of around 13%
dry weight to a winter low of about 8%. Thus fat
stored in the digestive diverticula of Tresus can be re-
garded mainly as an energy store which is called upon
only after several months of food scarcity, during which
time the glycogen store of the gonad is rapidly being
depleted. How dynamic is the flow of lipid in and out
of the storage globules is not known, and the transport
mechanism responsible for moving excess fat from the
tubule cells of the digestive diverticula is equally unclear,
though amoebocytes may have a role in this transport
as suggested by YoncE (1926).

The lipid in the gonad fluctuates rather more. Of
particular interest is the peak in mid-December which
probably is related to the build-up of lipid in the forma-
tion of eggs in the gonad. In a study of Crassostrea gigas
(THunsErG, 1793) by Masumoto et al. (1934), reported
in a review of the subject by Gopparp & Martin (1966),
a similar lipid peak was observed during the period of
“gonadal ripening.” This accumulation of gonad lipid
probably occurs at the expense of both gonadal glyco-
gen and diverticular lipid. The drop in gonadal lipid
after mid-December is explicable as the release of eggs
and the continued use of fat as an energy source. This
peak cannot be easily dismissed as an artifact caused by

the smallness of the samples, since the gonadal glycogen
of the animals sampled is rapidly decreasing.

The sudden drop in the level of glycogen occurs partly
because food becomes noticeably scarce in November,
and the glycogen is the major storage product. As well
as the depletion of glycogen for the purpose of fueling
the general metabolism of the animal, extra demands
are being made upon this food store for the production of
eggs and sperm. The re-accumulation of glycogen lags
about one month behind the re-availability of phyto-
plankton; the lag presumably reflects the increased en-
ergy requirements of the animals for growth. After the
middle of April the food available to the clams is in
excess of all the energy requirements, and storage can
rapidly take place.

ACKNOWLEDGMENTS

I am grateful to Miss Margaret Campbell who did most
of the analytical work. This study was supported by a
grant of the National Research Council of Canada.

LITERATURE CITED

Fotcu, Jorpi, M. Lees & STantey G. H. SLoaNE
1957. A simple method for the isolation and purification of
total lipids from animal tissues. Journ. Biol. Chem. 226:
499 - 509
Gopparp, C. K. & A. W. Martin
1966. Carbohydrate metabolism. In: Physiology of the Mol-
lusca, 2: 275 - 308 Acad. Press, New York & London
Mansour, K.
1946. Food and digestive processes of the lamellibranchs.
Nature, London 157: 482
Masumoto, B., M. Masumoto « M. Hrsino
1934. Biochemical studies of Magaki (Ostrea gigas). II. The
seasonal variation in the chemical composition of Ostrea gigas.
Journ. Sci. Hiroshima Univ., Ser. A, 4: 47 - 56
Rem, Rosert G. B.
1966. Digestive tract enzymes in the bivalves Lima hians
(Gmeutn) and Mya arenaria L. Comp. Biochem. Physiol.
17: 417 - 433
1968. The distribution of digestive tract enzymes in lamelli-
branchiate bivalves. Comp. Biochem. Physiol. 24: 727 - 744
Swan, Emery Freperick & JoHN H. FINUCANE
1952. Observations on the genus Schizothaerus.
66 (1): 19-26; plts. 2-4
ViLEs, FREDERICK J., Jr. « LESLIE SILVERMAN
1949. Determination of starch and cellulose with anthrone.
Jour. Anal. Chem. 21: 950 - 953
Yonce, CHARLES MAUuRICE
1926. Structure and physiology of the organs of feeding and
digestion in Ostrea edulis. Journ. Mar. Biol. Assoc. U.K.
14: 295 - 386

Nautilus

Page 382

THE VELIGER

Vol. 11; No. 4

Seasonal Gonadal Changes in Two Bivalve Mollusks

in Tomales Bay, California

BY

VERNON KENNETH LEONARD, Jr.

Pacific Marine Station, Dillon Beach, California 94925

(Plates 58 to 60; 4 Text figures; 1 Map)

INTRODUCTION

Pododesmus cepio (Gray, 1850), A BIVALVE mollusk of
the family Anomiidae, is native to the Pacific coast of
North America and common in Tomales Bay, California.
A small population of the European oyster, Ostrea edulis
Linnaeus, 1758, is being raised experimentally at the
Tomales Bay Oyster Company. The histological study of
the seasonal gonadal changes of these two species, in
Tomales Bay, is described below.

The study of gametogenesis in Ostrea edulis in Tomales
Bay is desired to evaluate the reproductive adaptation of
this species to a new set of ecological conditions. The
inclusion of the native Pododesmus cepio in this study
allows comparison of the gonadal development of these
two species. Such a comparison aids not only in under-
standing the seasonal gonadal changes in both species,
but also in examining the adjustment of the introduced
form, O. edulis, to a new environment.

At the present time the oyster industry in Tomales
Bay imports large numbers of oysters bred in other local-
ities and raises them until they reach marketable size.
These imported oysters, the Japanese [Crassostrea gigas
(THunNBERG, 1793) | and the American oyster [C. virgin-
ica (GMELIN, 1791)], do not normally reproduce in
California waters. The oyster industry in California
would be revolutionized if an exotic oyster, such as Ost-
rea edulis, were able to grow and reproduce in these
waters.

Knowledge of the reproductive cycle in Pododesmus
cepio is valuable not only because it offers a natural com-
parison to the introduced species, Ostrea edulis, but also
because it would then be the first member of the family
Anomiidae whose natural seasonal gonadal changes have
been examined.

There are no references to gonadal studies of Podo-
desmus cepio. However, prior to 1953 (Fircu, 1953) P
cepio was considered to be the same species as a very
closely related form, PR macroschisma (DEesHayes, 1839).
All available literature deals with this northern species.
Kettoce (1915) described the gross morphology and
ciliary currents of the mantle cavity of Monia (= Podo-
desmus) macroschisma. FrizzELt (1930) published a
note on the collection of a specimen of P macroschisma
in a Teredo navalis (LinNaEus, 1758) burrow from
Puget Sound. Brief references were also made to P mac-
roschisma in discussions of the family Anomiidae by At-
KINS (1937) and YonceE (1962). There are no works on
the seasonal gonadal changes in any other member of
the Anomiidae.

In contrast, Ostrea edulis is one of the most thoroughly
studied of all mollusks. Orton (1920, 1927, 1933, 1937),
Core (1936, 1941, 1942), and Korrinca (1940, 1952,
1957) have studied many aspects of the reproductive
cycle and sex change of the European oyster in its native
waters. LoosaNorr (1955) introduced O. edulis to New
England waters in 1949 and later studied the seasonal
gonadal changes of these oysters while they were adjust-

Explanation of Plate 58

Figure 1: Section of gonad of undifferentiated male Pododesmus
cepio. Collected October 1966 X125

Figure 2: Section of gonad of undifferentiated female Pododes-
mus cepio. Collected October 1966 125

Figure 3: Section of gonad of undifferentiated Ostrea edulis.
Collected November 1966 X125

Figure 4: Section of gonad of developing male Pododesmus cepio.
Collected February 1967 X 500

Figure 5: Section of gonad of developing female Pododesmus
cepio. Collected January 1967 X125

Figure 6: Section of gonad of developing Ostrea edulis.
Collected December 1966 X125

THE VELIGER, Vol. 11, No. 4 [LEonarp] Plate 58

Figure 6

Vol. 11; No. 4

ing to the new environmental conditions (LoosANoFF,
1962).

Three hundred European oysters were raised from lar-
vae cultured in the U. S. Bureau of Commercial Fisheries
Laboratory at Milford, Connecticut. With the assistance
of Dr. Victor L. Loosanoff and the courtesy of the State
of California, these oysters were imported to Tomales
Bay in October 1965.

ACKNOWLEDGMENTS

I wish to thank Drs. Victor L. Loosanoff and Edmund H.
Smith for their kind assistance and encouragement during
this study. I also thank Mr. Carl Berg and Mr. Les
Watling for aid in collecting the specimens.

METHODS ann MATERIALS

Sampling Methods

About 25 specimens of Pododesmus cepio were taken
monthly from a natural population in White Gulch, a
small cove in Tomales Bay, California (see Map, Text
figure 1). Samples were collected from October 1966 to
March 1968. With each collection, water temperature,
salinity, and turbidity measurements were taken.

Samples of Ostrea edulis were collected twice a month
at the beginning of the study and during the most active
reproductive period (March to July), then once a month
for the duration of the study. The sampling program ex-
tended from October 1966 to December 1967. As with
the White Gulch samples, temperature, salinity, and
turbidity were measured at each collection.

During the late spring and summer months, attempts
were made to obtain larvae from the spawning oysters.
To collect settling larvae, strings of cleaned oyster shells
were suspended in the water around the racks in which
the oysters were kept.

Laboratory Procedure

Each specimen was measured and the gonad examined
externally for features, such as size and color, which
might indicate its stage of development. The gonadal tis-
tue was prepared for histological study as follows: trans-
verse sections of the gonad, 1 to 2mm thick, were fixed
in Bouin’s fluid, dehydrated in isopropyl alcohol and em-
bedded in paraffin (melting point 52.5° C). Sections, 8u
thick, were stained with Heidenhain’s iron hematoxylin
and counterstained with eosin.

THE VELIGER

Page 383

California

lg Pacific Marine

Ke pu,; Station
White Gulch | | 3 Q
B . ~ ..Tomales Bay
: i EON. “<9 Tomales Bay
é pS TN Eke Onster
2 \ Company
F 3286
KS ee
ieee 2 : i
BS boa: miles

Figure 1
Map of the Study Area

CRITERIA ror ESTABLISHING STAGES

oF DEVELOPMENT
After examination of the gonadal tissue, a number from
1 to 5, representing a particular stage of development,
was given to each specimen. Each of these stages repre-
sented a naturally occurring and characteristic change
in the gonads of the animals during the entire reproduc-
tive cycle.

Stage I

This stage represents the resting or indifferent phase
through which Ostrea edulis and Pododesmus cepio pass

Page 384

following the activity of the previous reproductive period
(Plate 58, Figures 1, 2, and 3). In O. edulis, the gonadal
tissue surrounding the digestive gland is reduced to a
thin, translucent layer less than 1mm thick. The fine
tubules connecting the follicles to the genital ducts can be
seen through this layer. In P cepio, the gonad forms a
layer of tissue partially covering the stomach and con-
tinues anteriorly as an irregular tube attached to the
byssus gland. During this period the gonad is clear, and
the delicate genital tubules can be seen beneath the sur-
face epithelium. The gonad of P cepzio is usually 2-3mm
thick at this time.

In both species the follicles are separated by connective
tissue, often forming isolated pockets of follicular tissue.
Usually the follicles are small and completely void of
sex cells. However, primary germ cells are sometimes
present and are attached to the follicular wall. The
gonads of both species are colorless at this time; it is
difficult to determine their sex.

Stage II

Early gametogenesis in Ostrea edulis and Pododesmus
cepio is characterized by the development of primary
and secondary oogonia, spermatogonia, and, in some
advanced cases, oocytes and spermatids (Plate 58, Fig-
ures 4, 5, and 6). The connective tissue surrounding the
follicles supplies nutrient material (glycogen) for the
developing cells (Loosanorr, 1942). As the sex cells
proliferate, the follicles slowly enlarge and become more
numerous. During early development all stages of gam-
etogenesis can usually be found in one individual. Ostrea
edulis often has male and female sex cells in various stages
of gametogenesis in the same-or adjacent follicles. In
general, however, mature gametes are not present at this
time.

Externally the gonads become opaque as the develop-
ing cells fill the follicles. At this time the gonad of the
female Pododesmus cepio has a faint red-orange color
which becomes more brilliant as the animal ripens.
Throughout their development, the gonads of the male

THE VELIGER

Vol. 11; No. 4

have a creamy white color, similar to that of Ostrea
edults.

The later stages of gametogenesis are characterized
by the rapid increase in numbers of mature eggs and
sperm. In Ostrea edulis, ripe eggs interspersed with oo-
cytes line the follicles. In males and hermaphroditic in-
dividuals, the spermatids and spermatozoa fill the lumen
of the follicle. In this species the sperm characteristically
form spermballs with the tails projecting outwards as
in Plate 59, Figure 9.

As the gonads of Pododesmus cepio ripen, eggs rapidly
fill the follicle, with a corresponding decrease in the
number of cells in earlier stages of oogenesis. Male fol-
licles typically have primary stages of spermatogenesis
in the outer portion of the follicle, with more mature
sex cells occupying the lumen. In P cepio, the sperm are
arranged in branching networks with their tails aligned
toward the center of the follicle.

Externally the gonads of both species become firmer
and larger than during earlier stages. The color also
becomes brighter, especially in female Pododesmus cepio.

Stage III

When fully ripe, both species possess large, character-
istically swollen follicles containing mature gametes (Plate
59, Figures 7, 8, and 9). Developing stages of gameto-
genesis are seldom present at this time. Ripe ova and
spermatozoa are commonly found in the genital ducts
awaiting discharge. Mature eggs in Pododesmus cepio
often have a highly compressed appearance. The eggs are
60 to 75y. in size at this time. In Ostrea edulis, the eggs
have a diameter of 85 to 90m but do not appear com-
pressed.

At this time the gonad of Pododesmus cepio attains
its largest size, sometimes a centimeter or more in diam-
eter. Ostrea edulis, however, never develops a thick gon-
adal layer. In Tomales Bay, as in Boothbay Harbor,
Maine (LoosanorrF, 1962), the gonad of O. edulis is
rarely more than 14 mm thick. The ripe condition in the
European oyster is obvious when the gonad is punctured,
allowing the tightly packed gametes to be released.

Explanation of Plate 59

Figure 7: Section of gonad of ripe male Pododesmus cepio.
Collected June 1967 X125

Figure 8: Section of gonad of ripe female Pododesmus cepio.
Collected June 1967 125

Figure 9: Section of gonad of ripe Ostrea edulis.

Collected May 1967 X125

Figure 10: Section of gonad of spawned male Pododesmus cepio.
Collected August 1967 X125

Figure 11: Section of gonad of spawned female Pododesmus
cepio. Collected August 1967 X125

Figure 12: Section of gonad of spawned Ostrea edulis.
Collected August 1967 X125

ora

Tue VELIGER, Vol. 11, No. 4 [LEonarp] Plate 59

Figure 11 Figure 12

Vol. 11; No. 4

Stage IV

Examination of recently spawned specimens of Podo-
desmus cepio and Ostrea edulis reveals large empty fol-
licles (Plate 59, Figures 10, 11, and 12). Although a
few unspawned gametes are often visible in the lumina
of the follicles, no developing cells are present. In many
cases, partially spawned animals will have empty fol-
licles on the periphery of the gonad but fully ripe fol-
licles closer to the digestive gland. Specimens of O.
edulis commonly have cells of the next sexual phase
undergoing gametogenesis in the evacuated follicles im-
mediately following spawning. For example, a female
phase may be completed early in the summer with a male
phase beginning soon after the spawning of the eggs.

The appearance of the gonads following spawning is
dramatically different from the previous ripe condition.
The gonads are greatly reduced in size, no longer firm,
and usually void of sex cells in areas adjacent to the
genital ducts. This shrunken condition is especially ob-
vious in Pododesmus cepio.

Stage V

Ostrea edulis and Pododesmus cepio pass through a
period of resorption following spawning, in which any
remaining sex cells are absorbed by phagocytes (Plate
60, Figure 13). During this time the follicles of both
species are small; the lumina are filled with phagocytes
and fragments of partially resorbed gametes. Resorption
is a slow process and often extends over several months’
time, thereby overlapping with the development of new
gametes.

GAMETOGENESIS

It is appropriate to begin the description of gameto-
genesis in Pododesmus cepio and Ostrea edulis with the
inactive period (Stage I) following spawning and re-
sorption. In Tomales Bay this period occurred during
October and November, in both 1966 and 1967. The
water temperature was 16° C over the O. edulis popula-
tion and approximately 13°C in White Gulch (Text
figures 2 and 3). The gonads of both species were typical-
ly in the Stage I condition, undifferentiated and empty.
During this stage the oyster usually had some primary
germ cells attached to the follicular wall, while P cepio
was completely barren. The duration of this indifferent
period was similar for P cepio and the oyster (Tables 1
and 2).

Early gametogenesis (Stage II) began late in Novem-
ber 1966 and continued until April and early May 1967.

THE VELIGER

Page 385

Temperature °C
ES

8
OENED EM pea Meare) Ay Ss” OFN De] br M
1966 1967 1968

Figure 2

Water Temperature at White Gulch, Tomales Bay, California,
from October 1966 to March 1968

Although the water temperature in both study areas
dropped to 12.5°C, each species had begun to show
gonadal differentiation by early December 1966. The
European oyster developed more rapidly than did Podo-
desmus cepio. Ripe Ostrea edulis were found as early as
December 1966, while a significant number of ripe P
cepio was not found until April 1967. All stages of
spermatogenesis and oogenesis were found during this
active period. In both species, individuals were found
lagging behind or developing more rapidly than the rest
of the specimens of the sample.

As the water warmed from a low of 9° C in January
to 12- 13°C in March 1967, both species continued to

Temperature °C
=

OENeDI Ee MyArMe J VAlS) OFNEDeI
1966 1967 1968

Figure 3
Water Temperature at Tomales Bay Oyster Company, California,
from October 1966 to January 1968

Page 386 THE VELIGER Vol. 11; No. 4

Table 1

Numbers of Pododesmus cepio in different stages of
gonad development recorded at each collection;
October 1966 to March 1968

See ta ce a aes Mae ae ass
QQ

Hoe eb gd 4 of BF ee 8
By 2 Se eB ef 2 S 8 BS Se eB BB

Stage 8B Bg BSB) 9) ey SBS) ores) coe
Oe es Ss ee 4s 48 85262-0242 46 Sea a
1966 1967 1968

rt A 1 Bi 6 210 12 iO

in, 6 1 ih 1 1 2) WM Ge S Dn i 3 AO Si e316

IL. 3 AA MG Bf 4 10 8 8

IV. iO 22 op 10 4

Visi? VM IG SF Ml 12° 12 3

Note: The approximate duration of each stage is indicated by the
blocks surrounding the numbers in the table above.

Table 2

Numbers of Ostrea edulis in different stages of gonad
development recorded at each collection;
October 1966 to December 1967

Bes ame ze PAS ail sion rs!
OB ea he ae 484 & 6a © 8 2
1966 1967
I. 10 9 1 4 3S)
II. 220) 260) LOM Se t/a 1 1
III. 1 1 3. 2 8 ml 1 7 ie 1 2 vA Byes
IV. 1 Do Bae 385 36 1
V. 1 1 1 tied a | 1 1

Note: The approximate duration of each stage is indicated by the
blocks surrounding the numbers in the table above.

Explanation of Plate 60

Figure 13: Section of gonad of resorbing female Pododesmus Figure 14: Section of gonad of hermaphroditic Pododesmus
cepio. Collected November 1967 125 cepio. Collected November 1967 X125

Tue VELIcER, Vol. 11, No. 4 [LEonarp] Plate 60

Figure 13

x -

Figure 14

Vol. 11; No. 4

THE VELIGER

Page 387

ripen, reaching advanced stages of gametogenesis in
April.

The fully ripe condition (Stage III) was not prevalent
in Ostrea edulis until April 1967, when 30% of the
sample attained this stage. Pododesmus cepio did not
become fully mature until late May or early June 1967,
although ripe individuals were observed in late March.
During April the water temperature in both localities
rose above 15° C, after which the gonads of both species
ripened rapidly.

The climax of the reproductive cycle of the two bi-
valves in Tomales Bay (Stage IV) was reached during
July 1967 (Tables 1 and 2). The duration of the spawn-
ing period in Ostrea edulis was from May to September
1967, while in Pododesmus cepio it extended only from
June to August. The period of most intense spawning
coincided with the warmest water temperatures of the
year, 19° C in White Gulch and 21.5° C at the oyster bed.

Following spawning, each species entered a period of
resorption (Stage V) during September and October of
1966 and 1967. Phagocytic cells were found within the
lumina of the follicles, resorbing unspawned material. In
Pododesmus cepio the phagocytes invade the follicles
from the surrounding connective tissue, while in Ostrea
edulis the phagocytes enter directly from the blood vessels
(LoosanorF, 1962). The resorptive phase provides some
nutritive material for the development of new germ cells.
Resorption is a slow process, both in Tomales Bay and in
Boothbay Harbor, Maine (Loosanorr, 1962), and often
overlaps the early stages of gametogenesis in the next
season, thereby completing the reproductive cycle.

FACTORS AFFECTING GAMETOGENESIS

Temperature

Gametogenesis in Ostrea edulis and Pododesmus cepio
is an uninterrupted process in Tomales Bay. Once begun
in November, the development continued until the gon-
ads were ripe in March or April.

This uninterrupted development is unusual for Ostrea
edulis. OrToN (1933) stated that O. edulis in England
began rapid gonadal development in the early spring fol-
lowing a period of hibernation (December to February).
LoosanorF (1962), in discussing gonadal changes in O.
edulis in Maine, noted that the oyster passed through a
period of dormancy during the winter months and did
not begin gametogenesis until April or May each year
when the water temperature reached 10° C. In the Adri-
atic Sea, O. edulis began gametogenic activity in the early
spring and usually spawned by April following the period
of winter quiescence (PERUSKo, 1967).

Interrupted gametogenesis is common in several other
bivalves. Crassostrea virginica is dormant during the win-
ter in Long Island Sound and exhibits rapid gonadal
development in May and June when the water tem-
perature rises from 10°C to 12°C (Loosanorr, 1942).
In Cyprina islandica, the ocean quahog of Rhode Island,
gametogenic activity is slowed appreciably, although not
completely arrested during the cold winter months
(LoosanorFr, 1953). The hard shell clam, Mercenaria
mercenaria (LINNAEUS, 1758), also passes through a
winter period of reduced gametogenic activity in Long
Island Sound (Loosanorr, 1937a). _

It is not surprising that Ostrea edulis should develop
well and spawn in Tomales Bay, for the temperature
range is probably more favorable than in its native Euro-
pean waters. In Tomales Bay the water temperature
dropped to approximately 9°C during December 1966
and January 1967, but rose above 15°C by April 1967
(Text figures 2 and 3). This “critical temperature” (Or-
TON, 1920) was maintained or exceeded from May to
October 1967, allowing 7 months for active gonadal de-
velopment and spawning. This favorable temperature
range no doubt accounts for the early and rapid matura-
tion of the gonad of O. edulis in Tomales Bay.

Many early workers, in studying lamellibranch repro-
duction, discussed the existence of a minimum tempera-
ture below which the bivalves could not successfully
reproduce. Hopkins (1936, 1937), in his work with
Ostrea lurida CARPENTER, 1864, noted that ripening and
spawning of the oyster in British Columbia did not
occur until the water temperature reached 15 to 16°C.
Cor (1931b, 1932) found that when the water tempera-
ture reached 15°C in southern California, O. lurida
would spawn soon after. More recently, LoosANOFF &
NomeyKo (1951) showed that not only is there a mini-
mum temperature limiting reproduction, but oysters
raised in northern latitudes would spawn at different
times and different temperatures than individuals raised
in warmer, southern areas. They found that actual physi-
ological races existed in the same species grown in dif-
ferent areas. Chesapeake Bay oysters would not spawn
when transferred to the cold waters of Long Island
Sound. Crassostrea virginica from Milford, Connecticut,
however, would spawn earlier and at lower tempera-
tures when transplanted in Chesapeake Bay (LoosaNoFr
& NomMEjxko, 1951).

Although no conclusive evidence is available regarding
Pododesmus cepio in Tomales Bay, the indication is that
a temperature of 15°C or higher is necessary for the
ripening and spawning of gametes. A recent study has
been made of the seasonal gonadal changes in Crassost-
rea virginica and C’. gigas in Tomales Bay (BeErc, personal

Page 388

THE VELIGER

Vol. 11; No. 4

communication). It was found that gametogenesis oc-
curred in the spring, and that the gametes began to ripen
in May and June after the water temperature reached
15° C. Likewise, Ostrea edulis began to ripen significantly
when the water temperature rose above 15°C. This
evidence suggests that, as with the Olympia oyster, O.
lurida, 15°C may be the minimal temperature; below
this temperature P cepio and O. edulis cannot spawn in
Tomales Bay.

Salinity

The oysters were grown in wire cages suspended 1 foot
above the bottom where the winter salinity never dropped
below 22.5%. Although the surface salinities dropped
below 5%, at the oyster beds, and below 11%, in White
Gulch (Text figures 4 and 5), the development of the
gonads appeared unaffected by the lowered concentra-
tions. Pododesmus cepio occurs in the low intertidal zone,
where the briefly lowered salinity (during February
storms) could only be deleterious during extremely low

OLNED IERIE Al Niji ANSE ONNID TmENN
1966 1967 1968

Figure 4
Salinity at White Gulch, Tomales Bay, California,
from October 1966 to March 1968

tides. According to Davis & ANSELL (1958), develop-
ment and larval growth of Ostrea edulis was normal at
salinities of 22.5%, in Milford, Connecticut. Since the
average salinity at the Tomales Bay oyster beds was 27 to
32%. for most of the year, it is unlikely that extremes in
salinity adversely affected gonadal development.

Hydrogen-Ion Concentration

The hydrogen-ion concentration in both areas was re-
markably constant during the study period. The pH
ranged from 7.6 to 8.2. CaLaBrEsE & Davis (1966)
found that oysters kept at a salinity of 27%, could toler-

Surface Salinity

9 ' ‘

6 Aut \

3 = .
ONDJFMAMJJASOND J
1966 1967 1968

Figure 5

Salinity at Tomales Bay Oyster Company, California,
from October 1966 to January 1968

ate extremes in pH from 6.75 to 8.75. In most cases the
oysters developed normal straight-hinge larvae. Since
the observed pH in Tomales Bay was within the accepted
tolerance limits, it is doubtful that this factor caused any
changes in the gonadal development of Ostrea edulis and
Pododesmus cepio.

Turbidity

Excessive turbidity most likely has a greater effect on
both species, especially their larval forms, than lowered
salinities. The author observed turbid water at Tomales
Bay Oyster Company all year, with extreme conditions
at times of peak run-off from winter and late spring
storms. The White Gulch area was turbid only during
and after heavy spring rains. In many cases during the
summer spawning periods, the oyster shells would be
covered with a layer of silt and mud several millimeters
thick. Quantitative turbidity measurements made by Pa-
cific Marine Station (Water Quality Study) substantiate
these field observations. These data are available in the
recent Federal Water Pollution Control Administration
Progress Report (WP-GW-1061-02).

It seems likely that excessive turbidity did affect larval
development of Ostrea edulis in Tomales Bay. Although
the animals developed and spawned normally, no larvae
were seen by this investigator. Every specimen sampled
during the spring and summer months was examined
for the presence of larvae in the gills, but none were
found. Similarly, Bere (personal communication), in his
study of Crassostrea gigas and C. virginica, found only 4
or 5 bivalve larvae and a similar number of spat during
the summer spawning period.

Vol. 11; No. 4

THE VELIGER

Page 389

In addition to the high concentration of mud and silt
observed, a large bloom of dinoflagellates occurred
during July and August at the oyster beds. These turbid-
ity-causing organisms, together with particles of silt,
affected the oyster larvae by covering all available sur-
faces with material, thus preventing any larvae present
from settling. Moreover, the large concentrations of
plant cells either removed nutrients from the water vital
to the larvae, or released toxic metabolites causing mass
mortality. Finally, CALABRESE & Davis (1966) noted that
high turbidity could lower the pH, in some cases below
the tolerance limit of the larvae, and cause mortality.

While turbidity and related factors most likely caused
larval mortality, there was less than 10% mortality
among the Ostrea edulis adults. It is doubtful, therefore,
that turbidity affected gonadal changes in the European
oyster. In comparison, no significant mortality of Podo-
desmus cepio larvae was observed; in fact, spat were
found just a few weeks after the main spawning period in
August.

Sexuality

Hermaphroditism in mollusks, especially in bivalves, is
well known and has been thoroughly reviewed by Cor
(1943, 1944). Orton (1927, 1933) established that Ost-
rea edulis is a protandric hermaphrodite which passes
through consecutive sexual phases. Examination of O.
edulis in Tomales Bay revealed that the majority of the
animals developed as functional females early in spring
and, immediately after spawning, began producing male
gametes. Cor (1931b, 1932) has described similar sexual
phases in O. lurida and in the ship worm, Teredo navalis
(LinnaEus, 1758) (Cor, 1941).

Approximately 1% of the Pododesmus cepio examined
was hermaphroditic (Plate 60, Figure 14). In 4 of the
cases the animal had spawned as a male and still had
undischarged sperm in the tubules; at the same time
young ovocytes were developing on the follicular walls.
In another case the animal had fully developed sperm
and eggs.

Over 95% of all animals under 40 mm (average of
length and width) were male. This evidence suggests
that Pododesmus cepio is protandric. In addition, the sex
ratio of larger animals (55% ¢&, 40% 2, 4% unde-
termined) indicates that some sex reversal must occur
as the animal becomes older. As noted earlier, both P
cepio and Ostrea edulis pass through an indifferent stage
in which the sex cannot be determined. It is possible that
during this period P cepio may change sex if the ap-
propriate stimulus is present.

CONCLUSION

Seasonal gonadal changes in Ostrea edulis and Pododes-
mus cepio in Tomales Bay took place in a continuous
manner. Gametogenesis began in the late fall and con-
tinued until the ripened condition was attained in the
spring. Ostrea edulis began mass spawning in June, while
P. cepio did not spawn until July and August. Resorption
of unspawned gametes took place for varying lengths of
time in each species following spawning. Prior to the
onset of the next reproductive cycle, P cepio and O. edulis
underwent a brief period of quiescence, during which
time the gonads were generally undifferentiated.

The temperature and salinity factors were favorable
for the normal development and spawning of the imported
European oyster. However, highly turbid conditions at the
oyster beds appear to be responsible for the failure of
Ostrea edulis to propagate in Tomales Bay.

LITERATURE CITED

ATKINS, DAPHNE
1937. On the ciliary mechanisms and interrelationships of
lamellibranchs. Part III. Types of lamellibranch gills and their
food currents. Quart. Journ. Micr. Sci. 79: 375 - 421
CALABRESE, ANTHONY & Harry Cari Davis
1966. The pH tolerance of embryos and larvae of Mercena-
ria mercenaria and Crassostrea virginica. Biol. Bull. 131
(3): 427 - 436
Cor, WESLEY ROSWELL
1931. | Sexual rhythm in the California oyster (Ostrea lurida).
Sci. 74: 247 - 249
1932. Development of the gonads and the sequence of the
sexual phases in the California oyster (Ostrea lurida).
Bull. Scripps Inst. Oceanogr. Tech. Ser. 3 (6): 119 - 144
1943. Sexual differentiation in mollusks. I. Pelecypods.
Quart. Rev. Biol. 19: 85 - 97
1944. Sexual differentiation in mollusks. II. Gastropods, amphi-
neurans, scaphopods, and cephalopods. Quart. Rev. Biol.
19: 85-97
CoLe, HERBERT AUBREY
1936. | Experiments in the breeding of oysters (Ostrea edulis)
in tanks with special reference to the food of the larva and
spat. Fish Invest. London Ser. ITI, 15: 1 - 28
1941. The fecundity of Ostrea edulis. Journ. Marine Biol.
Assoc. U.K. 25: 243 - 260
1942. Primary sex-phases in Ostrea edulis. Parts Ili, IV.
Quart. Journ. Microsc. Sci. 83: 317 - 356
Davis, Harry Cart & ALAN Davip ANSELL
1962. Survival and growth of the European oyster, O. edulis,
at lowered salinities. Biol. Bull. 122 (1): 33-39
Fircu, JoHN Epcar
1953. | Common marine bivalves of California.
Fish and Game, Fish Bull. 90: 102 pp.

Calif. Dept.

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THE VELIGER

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FrizzELL, DonNALp LESLIE

1930. Pododesmus macroschisma DESHAYES. Nautilus
43 (3): 104 [for correction, see (4): 138]
Hopkins, A. E.
1936. Ecological observations on spawning and early larval

development on the Olympia oyster (Ostrea lurida).
Ecology 17 (4): 551 - 566
1937. Experimental observations on spawning, larval devel-
opment and setting in the Olympia oyster, Ostrea lurida.
Bull. U.S. Bur. Fish. 23: 439 - 503
KeEttocc, JAMES LAWRENCE
1915. Ciliary mechanisms of lamellibranchs with descriptions
of anatomy. Journ. Morph. 26 (4): 625-701
Korrinca, PIETER
1940. Experiments and observations on swarming, pelagic
life and setting of the European flat oyster, O. edulis. — Arch.
Neerl. Zool. T. V. 14: 1 - 249
1952. Recent advances in oyster biology.
Biol. 27: 266 - 308; 339 - 365
1957. | Water temperature and breeding throughout the geo-
graphical range of Ostrea edulis. Ann. Biol. 33 (1-2):
1-17
Loosanorr, Victor Lyon
1937. Seasonal gonadal changes of adult clams, Venus mer-
cenaria (L.). Biol. Bull. 72 (3): 406 - 416
1942. Seasonal gonadal changes in the adult oysters, Ostrea
virginica, of Long Island Sound. Biol. Bull. 82 (2): 195
to 206

Quart. Rev.

Loosanorr, Victor Lyon

1953. | Reproductive cycle in Cyprina islandica. Biol.
Bull. 104 (2): 146-155

1955. The European oyster in American waters. Sci.
121: 119- 121

1962. | Gametogenesis and spawning of the European oyster,
Ostrea edulis, in waters of Maine. Biol. Bull. 122 (1):

86 - 94
LoosanorrF, Victor Lyon « CHartes A. NoMEJKo

1951. Existence of physiologically-different races of oysters,

Crassostrea virginica. Biol. Bull. 101 (2): 151 - 156
Orton, Joun H.

1920. Sea temperature, breeding and distribution in marine
animals. Journ. Marine Biol. Assoc. U.K. 12: 339 - 366

1927. Sex change in the European oyster. Journ. Marine
Biol. Assoc. U. K. 14: 967 - 1045

1933. | Observations and experiments on sex change in the
European oyster (Ostrea edulis). Part III. On the fate of
the unspawned ova. Part IV. On the change from male to
female. Journ. Marine Biol. Assoc. U. K. 19: 1 - 53

PeruSko, GiLiian H.

1967. A study of the gonads of Ostrea edulis L. in relation
to its spawning cycle in the North Adriatic. Thallasia
Jugoslav. 3 (1-6): 5-10

YonceE, CHARLES MAuRICE

1962. On the primitive significance of the byssus in the Bival-
via and its effect in evolution. Journ. Mar. Biol. Assoc. U,
K., 42: 113 - 127

Vol. 11; No. 4

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Page 391

The Shell Structure and Mineralogy of Chama pellucida Broperip

BY

JOHN DAVID TAYLOR
Department of Zoology, British Museum (Natural History), London

AND

WILLIAM JAMES KENNEDY
Department of Geology and Mineralogy, Oxford

(Plates 61 to 64; 6 Text figures)

THE CALCIFIED SHELL OF THE BIVALVIA is normally built
up of two forms of calcium carbonate, aragonite and
calcite. Shells may be wholly aragonitic, or may contain
both aragonite and calcite, always in separate monomin-
eralic and structurally distinct layers.

Amongst extant bivalves, calcite is confined to 7 super-
families of the subclass Pteriomorphia (classification af
ter NEWELL, 1965) and a single member of the Het-
erodonta. This species is Chama pellucida Broverip, 1835.

Primary calcite was also present in an extinct hetero-
dont group, the rudists (Hippuritacea; KENNEDY «&
_ Taytor, 1968).

The distribution in living bivalves is:

superfamilies
Palaeotaxodonta: Nuculacea aragonite only
Nuculanacea aragonite only
Cryptodonta: Solemyacea aragonite only
Pteriomorphia: Arcacea aragonite only
Limopsacea aragonite only
Mytilacea aragonite only
or aragonite ++
calcite
Pinnacea aragonite + calcite
Pteriacea aragonite + calcite
Pectinacea aragonite + calcite
Anomiacea aragonite + calcite
Limacea aragonite + calcite
Ostreacea aragonite + calcite
Palaeoheterodonta: Unionacea aragonite only
Trigonacea aragonite only
Heterodonta: All wholly aragonite, so
superfamilies far as is known, ex-
cept for Chama pel-
lucida.
Anomalodesmata: All wholly aragonitic

The single occurrence of calcite in Chama pellucida,
outside the Pteriomorphia, is therefore of considerable in-
terest. This fact was first reported by Lowenstam (1954a,
1954b) and confirmed subsequently by him (LowEenstTam,
1963, 1964) and by our present work.

In bivalve superfamilies where calcite and aragonite
occur together in the same shell, LowEenstam (1954a,
1954b, 1963, 1964) and Dopp (1963, 1964) have shown
that in some cases the proportion of aragonite is related
to temperature. Species living in warmer water tend to
have a higher proportion of aragonite in their shells than
species living in temperate or cold waters.

Chama pellucida is a West American species ranging
from Peru to California. It thus extends into cooler waters
well outside the normal range of the Chamacea, which are
otherwise more or less confined to the tropics and sub-
tropics. LowenstaM (1954b, 1963, 1964) was thus able
to use Chama pellucida to illustrate the temperature ef
fect on mineralogy. That is, calcite appears in a species
which inhabits cooler waters than the other wholly aragon-
itic members of the superfamily. Furthermore, he illust-
rated (LowENstTAM, 1963, plate iv) the microstructure of
the shell, stating that the outer aragonitic layer of the
shell in warm water species of Chama is transformed to an
outer calcitic layer in Chama pellucida. This layer has a
distinct and different microstructure.

During a general survey of the mineralogy and micro-
structure of the Bivalvia (TayLor, KENNEDY & HALL, in
press), we have found this interpretation to be incorrect.
In view of the exceptional nature of Chama pellucida we
present this more detailed study.

METHODS

Mineralogical determinations were carried out by X-ray
diffraction on samples of the shell layers of 29 species of

Page 392

Chamacea. The microstructure of the shell was examined at
optical level by use of acetate peels prepared from pol-
ished, etched sections (methods after KUMMEL & Raup,
1965). Petrographic thin sections were also examined.
Fine structure was studied on shell interiors and on pol-
ished and E.D.T.A. etched sections with a Cambridge
Instrument Company (U. K.) ‘Stereoscan’ scanning elec-
tron microscope. Two-stage formvar, gold palladium shad-
ed replicas were studied by transmission electron micro-
scopy (techniques from Kaye, 1964).

OBSERVATIONS

The shell of most Chamacea consists of two layers, an
outer, crossed-lamellar layer and an inner, complex crossed-
lamellar layer (terminology from Boccip, 1930, TayLor,
KENNEDY & HALL, in press). These two layers are sepa-
rated by a thin layer of a blocky prismatic aragonite. This
is the pallial myostracum (OBERLING, 1964), and repre-
sents the trace of mantle attachment at the pallial line,
being deposited below the pallial muscles. It is the ‘pellucid
layer’ of Japanese workers (i.e. KopayAsHt, 1964), the
hypostracum of Jameson (1912), Lowenstam (1964)
and others, and the ‘helle Schicht’ of many workers. When
sections are cut through the muscle scars, thick pads of
similar myostracal prisms are seen, forming thick adductor
myostraca. Small areas of myostracal prisms occur else-
where in the shell, and are discussed below. These relations
are summarised in Text figure 1.

In Chama pellucida there is an additional, outer layer,
built of prismatic calcite. Within this there is a middle,
crossed-lamellar layer, and an inner, complex crossed-
lamellar layer, bordered by the trace of the pallial myos-
tracum. These relationships are summarised in Text figure
2. Chama pellucida thus possesses an additional lay-
er, not, as LowENnstAM (1964) states, a layer which

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Vol. 11; No. 4

pallial myostracum

e (RE

outer
crossed-lamellar layer

inner complex

crossed-lamellar layer

adductor pallial

myostraca myostracum

Figure 1

Distribution of shell layers in Chama macerophylla
A — section B — shell interior

replaces the outer layer of wholly aragonitic species. The
middle layer of C. pellucida is equivalent to the outer
layer of other species of Chama.

Explanation of Plate 61

Figure 1: The contact of the middle crossed-lamellar layer and the
adductor myostracum of Chama radians. Acetate peel of a radial
section. x 100

Figure 2: Radial section of the inner layer of Chama lamellosa
showing complex crossed-lamellar structure and thin sheets of ‘my-
ostracal type’ prisms. Acetate peel. X 100

Figure 3: Transverse section of the outer, prismatic calcite layer of
Chama pellucida showing the large irregularly prismatic blocks with
much finer units within. Acetate peel. X 100

Figure 4: Planar section of the outer prismatic, calcite layer of
Chama pellucida. Sub-parallel finely prismatic units and growth
bands are visible. Acetate peel. 100

Figure 5: Contact of the outer prismatic layer (above) and the
middle crossed-lamellar layer (below) in Chama pellucida. Growth
bands are continuous between layers. Crossed-lamellar structure is
seen in the lower right hand corner of the picture, the transition
zone between the layers is homogeneous and rich in organic matrix.
Acetate peel. X 100

Figure 6: The inner complex crossed-lamellar layer of Chama
pellucida showing myostracal pillars. Acetate peel. 100
Figure 7: Myostracal pillars in the inner complex crossed-lamellar
layer of Chama radians. Acetate peel of a radial section. X 100
Figure 8: Planar section through the adductor myostracum of
Chama lazarus. Acetate peel. Xx 100

Tue VELicER, Vol. 11, No. 4 [TayLor & KENNEDY] Plate 61

pa

raha > 4 t
ERR.

Figure 8

Vol. 11; No. 4

inner complex
crossed-lamellar layer

outer prismatic
calcite layer

SS Nos
WNOUGIAZ ss
x eZ

middle crossed-lamellar layer

tm

(
4
&

adductor
myostraca

ligament

Figure 2

Distribution of shell layers in Chama pellucida
A — section B — shell interior

The structure of the various parts of the shell is de-
scribed below.

STRUCTURE
OF THE
CROSSED-LAMELLAR LAYER

Conventional microscopy shows the inner surface of this
layer as a series of elongate, branching and interdigitating
lenses. These are arranged with their long axes running
concentrically, parallel to the shell margins. These lenses
are the first order lamels of Boccitp (1930). In sections,
these lamels run normal to the inner surface of the shell
layer, although often bending and turning towards the
outer surface of the shell. They often branch and inter-

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Page 393

digitate, producing a strong interlocking structure (Plate
64, Figures 20, 21).

Thin sections and peels of this layer show a very strik-
ing colour pattern, with adjacent first order lamels being
either straw or red-brown in colour.

Lamels may be up to several millimeters long, and are
usually of the order of 0.5 mm wide. Sections of the um-
bonal region show a characteristic pattern of diverging
primary lamels.

Internal structures of first order lamels are not easily
resolved at optical level. It can be seen, however, that
each first order lamel is built of second order lamels
which are inclined to the shell interior, with opposed di-
rections of inclination in adjacent first order lamels.
Each second order lamel appears to be built of smaller
laths, joined in side-to-side contact. Becke line studies
reveal that the whole of the crossed-lamellar layer is an
intergrowth of aragonite crystals in only two crystallo-
graphic orientations.

These observations are confirmed and extended by
electron microscopy (Plate 62, Figure 12). These results
are summarised in Text figure 3. Thus the whole layer
can be seen to be built of minute laths, lying parallel
within each first order lamel, and joined into sheets.
These laths are up to 1m in diameter, and some tens of

laths building up

second order lamels’

first order lamels 2
CS ~~
PR

adjacent: first

4 _ order lamels
organic matrix

Figure 3

Block diagram of crossed-lamellar structure,
based on electronmicrographs

Page 394

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Vol. 11; No. 4

microns long. Etching reveals the presence of lace-like
membranes within the crossed-lamellar layer; these cor-
respond to the proteinaceous organic matrix well known
in molluscan nacre and prisms. (GrecorE, 1967, with
references. )

STRUCTURE
OF THE
COMPLEX CROSSED-LAMELLAR LAYER

Peels and thin sections (Plate 61, Figures 2, 6, 7) show
that this layer is built up of the same basic elements as
crossed-lamellar structure, i. e., laths arranged into second
order lamels. These are not arranged into lenticular first
order lamels, but form irregular interdigitating blocks,
with lamel attitudes similar within blocks, but opposed

Civ = >
Resse
SSE

Figure 4

Block diagram of complex crossed-lamellar structure,
based on electronmicrographs

in adjacent blocks. Sections show areas of granular ap-
pearance between blocks which have a distinct lamel
structure within them, and since this pattern is seen in
sections cut in all orientations, we conclude that this layer
type is built up of second order lamels inclined in many
directions. Electronmicroscopy confirms these observa-
tions (Plate 62, Figure 9), and also shows the presence
of lace-like organic matrix within the complex crossed-
lamellar structure. This structure is shown diagrammati-
cally in Text figure 4.

STRUCTURE
OF THE
PRISMATIC LAYER

In hand specimens, this layer has a distinctive translucent,
‘pellucid’ appearance, and gives Chama pellucida its spe-
cific name. The outer layer is projected into a series of
irregular folia and squamae. At optical level (Plate 61,
Figures 3, 4, 5) this layer has a grey appearance, and is
built up of minute blade-like prisms, arranged more or
less normally to the shell interior margin at the time of
secretion. These minute prisms are variable in their atti-
tude, and are arranged into longer irregular blocks. Under
polarised light, these blocks go into extinction very irreg-
ularly, and there is thus no uniformity of orientation
within the blocks.

This structure is markedly different from that of the
calcite prismatic layer of most other bivalves, in that
thick conchiolin walls are not developed between prisms.

There is a variable relationship at the contact between
the prismatic layer and the crossed-lamellar layer. The
inner surface of the prismatic layer shows irregular cor-
rugations arranged radially from the umbo which are im-
pressions of radial musculature of the mantle. Over wide
areas of the contact there is a marked discontinuity with
a zone of fine-grained aragonite, rich in organic matrix
(Plate 64, Figures 21, 22, 23). This fills up the underlying
grooves in the corrugated surface. Elsewhere, minute
angular calcite crystals project into the outer part of
the aragonite crossed-lamellar layer (Plate 64, Figure 22).

Explanation of Plate 62

Figure 9: Scanning electronmicrograph of a fractured section of
the inner complex crossed-lamellar layer of Chama macerophylla.
Parts of three blocks of parallel laths are shown, the inclination of
laths in each block is different. 550

Figure 10: Scanning electronmicrograph of a polished, HCI-
etched section of the crossed-lamellar layer of Chama pellucida
showing sheets of fenestrate organic matrix. 1 400

Figure 11: The inner surface of a myostracal pillar in the inner
layer of Chama macerophylla. Note the irregular form of the
myostracal pillars and grooves between the prisms. Scanning elec-
tronmicrograph. X 2 200

Figure 12: Fractured section of the crossed-lamellar layer of
Chama macerophylla. The contact of two first order lamels is shown,
the second order lamels and laths of each first order lamel are in-
clined in opposed directions. Scanning electronmicrograph. X1 100

[TayLor & KENNEDY] Plate 62

THE VELIGER, Vol. 11, No. 4

°
cD
=]
3

ep

ea

Figure 12

Figure 11

Vol. 11; No. 4

The crossed-lamellar layer must thus grow over an angular
irregular prismatic surface.

Electron microscopy confirms these observations. Frac-
tured sections show the larger prismatic blocks, with
prominent transverse striations (Plate 63, Figure 16).
These are obviously built up of smaller elements. On the
inner surface of the shell layer, these blocks are not
distinct; all that can be made out is an irregular surface
of small pyramidal mounds and small granules (Plate 63,
Figure 13). These correspond to the outcrop of the smal-
ler prismatic units seen at optical level.

Etching of sections brings out details not seen at
optical level. Thus the fine prisms appear as elongate
blocks, whilst oblique sections show their outlines (Plate
63, Figure 14). Etching also reveals the presence of
organic matrix, as reticulate lace-like sheets surrounding
the prisms. This organic matrix is heavily developed at
the contact with the crossed-lamellar layer, whilst the
matrix of both layers appears to be in continuity.

STRUCTURE or tHE MYOSTRACA

The myostracal structure of all Chamacea is very similar.
At optical level it has a characteristic grey colour, strongly
contrasting with adjacent shell layers. The structure is
built of prisms, which are highly variable and irregular
in outline, with re-entrant angles (Plate 61, Figure 8).

Figure 5
Block diagram of myostracum

based on electronmicrographs
is not shown

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|

Page 395

These prisms vary greatly in size, but are always oriented
normal to the surfaces of the myostracum, with crystallo-
graphic c axes in the same direction. These relationships
are summarised in Text figure 5. On shell interiors, my-
ostracal prisms outcrop as minute irregular polygons.

Pallial and adductor myostraca are rcadily understood,
being secreted below areas of undoubted muscle attach-
ment. In the Chamacca myostracal structure is also de-
veloped as discrete sheets and pillars within the inner
shell layer (Plate 61, Figure 2). Myostracal sheets of
this type occur in some other bivalves, and we believe
they may indicate sites of temporary mantle attachment,
although conclusive evidence is lacking.

Figure 6

Block diagram of myostracal pillars.
The structure of the surrounding complex crossed-lamellar structure

Pillar-like structures (Plate 61, Figures 6 and 7), which
we propose to refer to as myostracal pillars, are developed
in most Chamacea. In peels and sections these appear as
elongate columns, with the long axis normal to the shell
interior. Serial sections reveal that these are in fact sec-
tions of continuous blocks of myostracum extending in-
wards from, and continuous with, the pallial or adductor
myostraca (Text figure 6). On shell interiors these outcrop
as minute bosses usually 0.5 to 0.1 mm across, sometimes
arranged in rows which radiate from the apex of the shell.
Plate 62, Figure 11 shows the surface of such a boss.
Here it is clear that there is a series of discrete prisms,

Page 396

separated by grooves (of uncertain origin). There are
also traces of finer structure visible within the prisms.

Histological preparations of Chama jukesu show that
the outer mantle surface is locally modified into papillae,
as the result of elongation of the mantle cells. These
papillae agree in size and shape with the myostracal pil-
lars present in the inner shell layer of that species, and
thus appear to represent the points of attachment.

TUBULES

The Chamacea all possess the remarkable shell feature
described by Oxseruine (1964) as tubules. These are
minute cylindrical perforations, usually only a few microns
in diameter, which open at the interior of the shell and
penetrate the shell layers. These are undoubtedly a pri-
mary feature of the shell, not to be confused with algal
borings or other perforations (OBERLING, op. cit.). The
distribution of tubules is very constant in the Chamacea
where they are largely confined to the inner shell layer.

At optical level tubules appear as minute, hollow, un-
branched, straight cylinders, cutting the finest observable
elements of shell structure.

At electron microscope level (Plate 64, Figures 17, 18,
19) the tubules are empty and unlined, and penetrate,
but do not seem to disturb, shell fabric. Indeed, details
of layering and fine structure are visible inside the tubule
(Plate 64, Figure 18). The opening of the tubule is a
distinct conical pit, and in some species of Chama, in-
cluding C. pellucida, the tubules are grouped in minute
oval depressions (Plate 64, Figure 17).

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Vol. 11; No. 4

We have been unable to determine the origin of tubules.
Since they occur more abundantly in older parts of the
shell, and in some species of bivalve are absent at the
margin, they may be resorptive, but histological work on
decalcified material has revealed only the ambiguous tra-
ces of mantle extensions into these holes.

DISCUSSION anp CONCLUSIONS

Chama pellucida has a shell structure which is compa-
rable to that of all other species of Chamacea in many
respects. It is, however, unique in possessing an outer,
prismatic calcite layer. The middle shell layer is equi-
valent to the outer layer of all other species; there is no
change in microstructure as LoweNnstTam (1963) has im-
plied; a new outer layer appears in C. pellucida, which
thus has a three layered, not a two layered shell.

It is also difficult to accept Chama pellucida as a wholly
cold-water species of Chama, for it ranges across the
equatorial belt.

We would therefore question previous interpretations
of the structure of Chama pellucida, and without further
evidence would doubt the validity of using it as an
example of environmental controls on shell mineralogy.

This species is, however, quite unique amongst extant
heterodonts in the development of calcite in its shell, and
the significance of this is not understood. The calcite
layer is, however, outermost, as is the calcitic shell layer
in most other bi-mineralic bivalves. In common with
Chama pellucida all other calcite bearing bivalves are
epifaunal, or are derived from epifaunal ancestors.

Explanation of Plate 63

Figure 13: Scanning electronmicrograph of the inner surface of

the outer prismatic calcite layer of Chama pellucida. The outcrop

of the minute prisms building up this layer are shown. 2 100

Figure 14: Polished, HCl-etched radial section of the outer pris-

matic layer of Chama pellucida. Scanning electronmicrograph.
X 700

Figure 15: Polished, etched section of the contact between the

middle crossed-lamellar layer (above) and the pallial myostracum
(below). Scanning electronmicrograph. 700

Figure 16: Scanning electronmicrograph of a fractured section of
the outer prismatic calcite layer of Chama pellucida. Surfaces of
larger prismatic units are visible, with distinct parallel, transverse
striations. The finer prismatic units building the larger blocks are
also visible. X 280

Explanation of Plate 64

Figure 17: Inner surface of the inner complex crossed-lamellar
layer of Chama pellucida. The openings of six tubules are visible,
four lie together in an oval depression, two are in separate circular
depressions. Scanning electronmicrograph. 270

Figure 18: Detail showing the opening of a single tubule. Note
details of fine structure within the hole and the lack of an obvious
lining. Scanning electronmicrograph. X 2750

Figure 19: Fractured section of inner complex crossed-lamellar
layer of Chama pellucida showing a tubule. Note the lack of lining
and the absence of any change of structure associated with the
tubule. Scanning electronmicrograph. X2 800

Figure 20: Radial section of the middle crossed-lamellar layer of
Chama pellucida. Bands of myostracal prisms representing the pal-
lial and adductor myostraca are seen. Acetate peel. X 100
Figure 21: Radial section of the contact between the outer calcite
prismatic layer and the middle crossed-lamellar layer of Chama
pellucida. Acetate peel. X 100

Figure 22: As above, showing details of the highly irregular contact.
The calcite prismatic layer (lower part of field) projecting into the
crossed-lamellar layer. Acetate peel. X 100

Figure 23: Radial section of the prismatic and crossed-lamellar
layer contact, showing the irregular contact and an organic rich
zone next to the prismatic layer. Thin section. X 100

[TayLor « KENNEDY] Plate 63

THE VELIGER, Vol. 11, No. 4

Figure 14

Figure 13

tT

> a = é
S => & Se gl
t ant * o es
- a ; ae
4 me A wy ps . \ “< ~
oe “ 4 a ‘
er . we Ss
N ¥ , a
¥¢ rf ‘ :
¥ wa | F > i 4
a 4 bags Ps ~
a ee re Ps 4
4 ~ * ‘
‘ # e SAS id ~~
a ¥ - a Z “ -
- f ef
2 f ,
ot ne 4° > ‘ s
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, 4 very a, id -.
: Le .
. re

es

Figure 16

Figure 15

Tue VELIGER, Vol. 11, No. 4 [TayLor & KENNEDY] Plate 64

igure

Figure 18
ee
Figure 22

Figure 23

Vol. 11; No. 4

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Page 397

An interesting comparison comes from the extinct het-
erodont group, the rudists. These are derived from wholly
aragonitic pachydonts such as Megalodon and Pachyrisma
(Cox, 1960), but on assuming a cemented habitat, and de-
veloping massive shells, calcitic outer layers appear. This
cannot, however, be taken as evidence for affinity between
Chamacea and Hippuritacea as NEwELL (1965) or YONGE
(1967) have suggested.

We also describe evidence for the local attachment of
mantle to shell interior as demonstrated by the presence
of sheets of myostracal prisms. Myostracal pillars and
papillae on the mantle surface of Chama jukesu indicate
continued attachment at localised sites with the pallial
line throughout the life of the animal in this, and by
inference, all other Chamacea.

ACKNOWLEDGMENTS

We gratefully acknowledge the help of Dr. A. Hall (King’s
College, London) with mineralogical determinations. We
are also grateful to Mr. B. Martin and the staff of the
electronmicroscope unit of the British Museum (Natural
History), and Mr. R. Holland (Department of Geology
and Mineralogy, Oxford) for technical assistance.

APPENDIX

A calcitic outer layer is also present in specimens in the
British Museum (Natural History) labelled Chama exo-
gyra ConraD, from California.

Examination of the type specimens of Chama exogyra
Conrap, 1837 and C. pellucida show that the specimens
are distinctive. Chama pellucida is rounded in outline,
‘normal’ (i. e., attached by the left valve) and has striking
translucent squamae. Chama exogyra is ‘inverse’ (at-
tached by the right valve), irregular, elongate, and lacks
conspicuous squamae.

Other specimens of Chama exogyra show rather more
conspicuous ridges and cementation can take place by
the left valve. ‘Normal’ specimens of C. exogyra closely
resemble C. pellucida and ‘inverse’ specimens of C. pel-
lucida closely resemble C’. exogyra.

The similarity of the unusual shell structure together
with the similar geographical range (YoncE, 1967) of
these two species, taken together with the problems of
inversion in the Chamacea makes us suspect that these
species may be synonymous.

We believe that there is sufficient doubt of their valid-
ity to merit a field investigation of their relationships. We
hope that this note will stimulate workers in California to
investigate the problem.

LITERATURE CITED

Boccixp, O. B.

1930. The shell structure of the mollusks. D. Kg. Danske
Vidensk. Selsk. Skr., Naturv. Math. Afd., 9 R II 2: 231 - 325;
10 figs.; plts. 1-15

Cox, Lestm REGINALD

1960. Thoughts on the classification of the Bivalvia. Proc.

Malacol. Soc. London 34 (2): 60-68
Dopp, J. R.

1963. Palaeoecological implications of shell mineralogy in two
pelecypod species. Journ. Geol. 71: 1-11

1964. Environmentally controlled variation in the shell struc-
ture of a pelecypod species. Journ. Palaeont. 38: 1065 - 1071

GréEcore, C.

1967. Sur la structure des matrices organiques des coquilles

des mollusques. Biol. Rev. 42: 653 - 688
Jameson, L. H.

1912. Studies on pearl-oysters and pearls. I. The structure of
the shell and pearls of the Ceylon pearl-oyster (Margaritifera
vulgaris SCHUMACHER): with an examination of the Cestode
Theory of pearl-production. Proc. Zool. Soc. London 1912:
260 - 358

Kaye, D.

1964. ‘Techniques for electron microscopy, pp. 1 - 381.

Blackwell Scient. Publ., Oxford
KENNEDY, WILLIAM JAMES & JOHN Davip TayLor
1968. Aragonite in rudists. Proc. Geol. Soc. London 1645:

325 - 331
Kosayasui, I.

1964. Microscopical observations on the shell structure of
Bivalvia, part I, Barbatia obtusoides (Nysvt.) Sci. Rep.
Tokyo. Kyoika Daig. Sect. C, 8: 295 - 301

KummMEL, B. & D. M. Raup
1965. | Handbook of palaeontological techniques. W. H.

Freeman, San Francisco & London, xiii+852 pp.
LowEnNsTaM, HEINnz A.
1954a. Environmental relationships of modification composi-
tions of certain carbonate-secreting marine invertebrates.
Proc. Nat. Acad. Sci. 40: 39 - 48
1954b. Factors affecting the aragonite:calcite ratios in car-
bonate-secreting marine organisms. Journ. Geol. 62: 284 - 322
1963. _ Biological problems relating to the composition and dia-
genesis of sediments. In: The earth sciences: Problems and
progress in current research: 138 - 195
1964. Coexisting calcites and aragonites from skeletal carbon-
ates of marine organisms and their strontium and magnesium
contents. In: Recent researches in the fields of hydrosphere,
atmosphere and nuclear geochemistry: 373 - 404
Matong, PG. & J. R. Dopp
1967. Temperature and salinity effects on calcification rate in
Mytilus edulis and its paleoecological implications. Lim-
nol. & Oceanogr. 12 (3): 432 - 436
NEWELL, Norman D.
1965. Classification of the Bivalvia.
2206; 3 figs.; 1 tab.

Amer. Mus. Novit.

Page 398 THE VELIGER

OBERLING, J. J.
1964. Observations on some structural features of the pelecy-

pod shell. Mitteil. Naturforsch. Gesellsch. Bern, new ser.,
20: 1 - 63; 6 plts.; 59 figs.
TayLor, JoHN Davin, WILLIAM JAMES KENNEDY & A. HALL

(in press) Shell structure and mineralogy of the Bivalvia:
(Nuculacea — Trigonacea). Bull. Brit. Mus. (Nat. Hist.)
(Zool.)

YONGE, CHARLES MAUuRICE
1967. Form, habit and evolution in the Chamidae (Bivalvia)

with reference to conditions in the rudists. Phil. Trans. Roy.

Soc., B 252: 49 - 105

Vol. 11; No. 4

—

Vol. 11; No. 4

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Page 399

Molluscan Faunas

of Pacific Coast Salt Marshes and Tidal Creeks

KEITH B. MACDONALD!

Scripps Institution of Oceanography, La Jolla, California 92037

(1 Map)

INTRODUCTION

Paciric Coast Spartina - Salicornia salt marshes, and the
tidal creeks that dissect them, contain distinctive mollus-
can faunas. The taxonomy of most of the species repre-
sented in these faunas is well documented (Frrcu, 1953;
Paumer, 1958; Keen, 1958, 1963; Hanna, 1966) and in
some cases papers describing aspects of the ecology of
individual species, or of closely related species in other
localities are available (e.g., HausEMAN, 1932; Mac-
GiniTig, 1935; SanpErR, 1950; Yonce, 1951; Meyer, 1955;
SELLMER, 1967).

These faunas deserve further study, not only because
they occupy habitats that are rapidly disappearing on the
Pacific Coast, but because the huge species populations
found at several sites may play an important role in the
cycling of nutrients and detritus (organic debris) and
thus indirectly affect offshore communities (NEWELL,
1965; CarrikerR, 1967; HepcpETH, 1967).

This report outlines the geographic distribution and
relative abundance of the species represented in these
faunas. No previous studies of this subject have been
found in the literature.

METHODS

Salt marshes and tidal creeks in 9 Pacific Coast bays and
estuaries were examined (Figure 1). At 7 of these locali-
ties single sites were sampled. Local spacial variations
were studied by sampling 2 sites at both Tomales Bay and
San Quintin Bay. Ten of the sites were sampled only once,
each sample-set being collected over a 2-3 day period.
To provide a basis for separating real latitudinal differ-

1 Present address: Department of Geology, University of Califor-
nia, Santa Barbara, California 93106

ences from yearly population fluctuations, 5 replicate
sample-sets were taken at Mission Bay, at approximately
quarterly intervals (November 1964 to July 1966).

Individual sites were selected for a minimum of pollu-
tion and freshwater runoff. Isolated marshes that could be
sampled as discreet units were preferred to artificially
defined sections of more extensive marshlands.

At each site the vegetated marsh surfaces and tidal
creeks were sampled independently. For the former, stra-
tified random sampling patterns were set up (CocHRAN,
1954). Random number tables were used either to locate
samples at random intervals paced along previously sur-
veyed relief transects, or, to select pairs of random co-
ordinates locating the samples within quadrants of a
prescribed irregular area. The molluscan data were collec-
ted from stainless steel rings enclosing an area of 200 cm’;
each sample was excavated to a depth of 1 cm.

At 9 of the sites the tidal creek samples were located
at random intervals paced along the creek banks; at Grays -
Harbor and Mission Bay these samples were collected at
fixed intervals. In most cases (cf. Table 2) the creek
bottom mollusks were sampled from 25 25 cm quadrats
excavated to a depth of approximately 25 cm. Upon
return to the laboratory each marsh or creek sample
was washed through a 1 mm mesh screen and all of its
molluscan components were sorted, identified and counted
(see Macpona_p, 1967, for additional details of sites and
sampling methods).

RESULTS ann DISCUSSION

Seventy-six mollusk species were collected during this
study; 64 of these were represented in the quantitative
samples and the remainder were picked up during recon-
naissance of the sites. In the samples, 2 species were
represented by live specimens only, 28 species by both

Page 400

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Vol. 11; No. 4

Figure 1

North American Pacific Coast,
showing the general location of the sites investigated

living and dead material and, the remaining 34 species
by empty shells only. Only the live material from the
quantitative samples is considered in this report.

The geographic distribution and relative abundance
(i. e., percentage of total individuals, per sample set) of

the mollusks represented in the live material are shown
in Tables 1 and 2. Table 3 summarizes seasonal changes in
the relative abundance of selected species in the replicate
samples taken at Mission Bay.

ABUNDANCE

In sets of 25 - 178 samples, 2 - 7 species were found in the
salt marshes and from 0-11 in the tidal creeks. The
number of species increases significantly southward in the
marsh environment (Kendall rank correlation procedure,
P < 0.05, 1-tailed). There is similar increase in the
tidal creeks, but largely because of the poor represen-
tation of mollusks at San Quintin Bay (Keen, 1962), it
is not significant (P > 0.05).

The mean density of mollusks (i. e., total individuals
per sample set/combined area of samples) and the mean
densities of separate species in both environments show
no correlation with the number of species recorded from
each marsh or creek, or with the latitude or area of the
site being sampled. This suggests that the abundance of
mollusks at a specific site is controlled by a variety of
local factors (sediment type, food supply, etc.) rather
than by regional trends of climatic or oceanographic
variables.

The distribution of individuals between species exhibits
a distinct pattern in the faunas of both environments:
90 - 100% of the individuals collected in each sample set
belonged to 2 or 3 species; any additional species present
were each represented by relatively very small numbers
of individuals. Inspection of individual sample-sets also
reveals that the abundant species were widely distributed
at each site whereas the less common species had markedly
patchy distributions. This pattern remains essentially the
same despite differences in the latitude and area of the

site and the species compositon and density of the faunas.

SPECIES COMPOSITION

The creek faunas usually contain more species and have a
more variable species composition than do those of the
marshes. In part this may reflect the more highly special-
ized fauna of the latter, a marked contrast to that of the
less extreme creek environments which contain a wide
variety of species found in other barely subtidal habitats.
Since a majority (83%) of the creek species are infaunal
and thus subject to the selective effects of substrate
(THorson, 1957; Purpy, 1964) the compositional vari-
ability between sites might also reflect the variable nature
of creek sediments. For example, at Tomales Bay shelly

Vol. 11; No. 4

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Page 401

Table 1

Salt Marsh Mollusk Faunas
Species Composition and Relative Abundance (i. e., percentage of total individuals per site)

Species 1 2 3

Gastropoda
Littorina newcombiana (HEMPHILL, 1876) SUG 2:3 0.3
eae 7) 65.8
Assiminea translucens (CARPENTER, 1864) 9.4 44.0 33.9

Phytia myosotis (DRAPARNAUD, 1801)

Batillaria zonalis (BRuGUIERE, 1792) +
Cerithidea californica (HALDEMAN, 1840)
Melampus olivaceus CARPENTER, 1857
Littorina scutulata Gou.p, 1849
Acteocina culcitella (Goutp, 1852)
Littorina planaxis (Putvipri, 1847)
Acteocina carinata (CARPENTER, 1857)

Bivalvia

Chione fluctifraga (SowErRBy, 1853)
Mytilus edulis LinNaEus, 1758
Ostrea lurida CARPENTER, 1864
Lasaea subviridis DAL, 1899

Total individuals 553 300 693
Marsh area (km?) 0.237 0.071 0.056
Number of samples 50 59 40

“Localities as in Figure 1

+ =present, < 0.1% individuals per site

* — present at site but did not occur in samples

+ = introduced species, not native to the West Coast

** — common at other sites around Grays Harbor

“ Mission Bay values obtained from combined replicate sample sets

gravels were common, at Grays Harbor only coarse clean
sands were present, at Coos Bay both soft muds rich in
plant debris and firmer muddy sands were found and at
the Humboldt Bay and San Quintin Bay sites almost
liquid muds predominated.

The geographic distribution of recurrent groups of spe-
cies supports the classification of Pacific Coast molluscan
provinces proposed by VALENTINE (1966). Within the
salt marsh faunas the Californian (27° - 34° N) and Ore-
gonian (34°-50° N) provinces are characterized by
Assiminea - Cerithidea and Assiminea - Phytia associations
respectively.

The distribution of the more abundant tidal creek
species suggests that the Oregonian Province can be fur-
ther divided into 2 sub-provinces: the Macoma- Mya
assemblage from Grays Harbor and Coos Bay, and rep-
resented by in situ empty shells at Humboldt Bay, charac-
terizing a northern sub-province; the Batillaria - Gemma
(both introduced species) assemblage from Tomales Bay

Locality '
4 5 6 7 8A 9 10 11

64.1 7.1 45.0
35.9 8.8 55.0 93.2 68.6 79.3 76.0 5.0

84.1
* 5:9 a) 28:2) 200) = ee 5
087 019) 0.3 32.4
4 -
AO. OF Be
+
27.6
- 0.5
*
*

ileil
245 309 5246 2282 8 8§=—. 2153 852 362 809
0.111 0.046 0.104 0.886 0.139 0.060 0.019 0.099
49 20 45 60 100 50 20 50

and Elkhorn Slough characterizing a southern one. The
Acteocina - Cerithidea association found in the tidal creeks
south of Point Conception characterizes the Californian
Province.

LOCAL VARIATION

Reconnaissance of different sites bordering the same bay
or estuary indicates that the faunas of adjacent marshes
or creek systems can vary considerably. Some of this vari-
ation reflects the environmental gradients characteristic
of estuaries: for example, Phytia myosotis was not found
at Westport but was common at other sites well within
Grays Harbor (Markham, Oyhut), perhaps indicating a
preference for less exposed or more brackish habitats.
Conversely, Littorina newcombiana was very rare at Ar-
cata but reached densities of 16 to 48 per m? on Samoa
Marsh near the mouth of Humboldt Bay, suggesting a

tH
3
Zz

b)

Vol. 11

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Page 402

Table 2

Tidal Creek Mollusk Faunas: Species Composition and Relative Abundance

Species Locality !
1 2 3 4 5 6 i 8 9 10 11

Gastropoda

Littorina newcombiana (HEMPHILL, 1876) 0.7

Batillaria zonalis (BRUGUIERE, 1792) + 92.2 63.9 43.0

Assiminea translucens (CARPENTER, 1864) 0.1 +

Cerithidea californica (HaLpEmMaN, 1840) * 84.4 28.2 3.5 78.0 74.4

Acteocina culcitella (Goup, 1852) 11.5 68.7 29.4 22.0

Bulla gouldiana Pirssry, 1893 0.1 +

Stphonaria brannani STEARNS, 1873 +

Nassarius tegula (REEVE, 1853) 1.4 4.42

Bittium sp. 67.1

Acteocina carinata (CARPENTER, 1857) 19.4
Bivalvia

Macoma inconspicua (BRopERIP & SOWERBY, 1829) 77.5 lee

Mya arenaria LinnaEus, 1758 4.2 71.4 0.1

Cryptomya californica (Conran, 1837) 1.4 0.1 “3

Macoma nasuta (Conran, 1837) 16.2 + 0.6 o

Laternula japonica (LiscHxKE, 1872) + 21.4

Gemma gemma (TortTEN, 1834) + 7.8 29.7 56.9

Modiolus senhousei (BENSON, 1842) + 5.3 ee 0.6

Protothaca staminea (Conrap, 1837) 0.6 2.8 0.2

Tapes japonica DESHAYES, 1853 + 0.1

Mytilus edulis LinnaEus, 1758 ae 0.1 oe

Chione undatella (SowERBy, 1835) 0.4 uP

Lyonsia gouldit Datu, 1915 +

Chione. fluctifraga (SowERBY, 1853) 0.6 1.3

Tagelus californianus (Conrap, 1837) 0.3 0.4
Total individuals 142 14 0 490 1010 3179 720 5462 681 1496 227
Creek area (km?) 0.026 0.006 0.003 0.008 0.004 0.004 0.045 0.003 0.003 0.001 0.003
Number of samples 20 29 16 18® 5° 20° 15 78 10 4 10

ee — (000.0

‘ Localities as in Figure 1

For explanation of symbols, see Table 1

B: sample size 312cm?; ©: sample size 400cm?; : sample size

200cm?; =: represented by Nassarius tegula tiarula (KIENER,
1834), a possible subspecies.

Vol. 11; No. 4

THE VELIGER

Page 403

Table 3

Mission Bay
Relative abundance of selected species in successive sample sets

Salt marsh
Assiminea translucens (CARPENTER, 1864)
Cerithidea californica (HaLpEmAN, 1840)
Acteocina culcitella (GouLp, 1852)
Melampus olivaceus CARPENTER, 1857
Tidal creek
Acteocina culcitella (CARPENTER, 1864)
Cerithidea californica (HaLtpEMAN, 1840)
Nassarius tegula (REEvE, 1853)

A B Cc D E

4.9 0 0 0.2

77.0, 45.7 66.5 85.4 61.1
19.9 50.0 30.4 13.0 35.9
1.8 Zl 0.2 0.2 Nees

A — November 1964 (marsh samples hand-picked in the field,

data found unreliable)
B — July 1965; © — October 1965;
E = July 1966

preference for less brackish environments.

Data from Tomales Bay and San Quintin Bay suggest
that differences of drainage pattern, substrate or vegeta-
tion can also account for local variability. The dominance
of Bittium in the creek fauna of one of the San Quintin
sites (10), for example, reflects the presence of dense
Zostera (Eel-grass) beds, absent from the other site (9).
Similarly, much of Millerton’s (5) diverse fauna was
collected from a network of narrow protected creeks with
soft mud bottoms, while at Walker Creek (4) exposed,
deep-water creeks with gravel bottoms were sampled.
Possibly the highly dissected nature of the Millerton marsh
also allowed Batillaria to remain closer to water sources
and successfully colonize the marsh surface while it did
not do so at either Walker Creek or Moss Landing (6).

SEASONALITY

Data from replicate samples taken at Mission Bay (‘Table
3) suggest that the relative proportions of the more abun-
dant species remain generally similar throughout the
year. Changes noted among the less common species prob-
ably reflect the sampling problems associated with rare or
accidental species ( e. g., Bulla gouldiana, Littorina plan-
axis, Siphonaria brannani), or with patchily distributed
species (c. g., Index of Dispersion indicated that all the
bivalve species were aggregated), rather than real com-
positional differences.

Seasonality was noted in Melampus olivaceus and Nas-
sarius tegula. The former exhibited hibernation behavior
similar to that described in M. bidentatus (HAUSEMAN,
1932) and was generally absent from the marsh surface

D = March 1966;

from about November through March. During this period
many individuals were found clustered together in crab
burrows 5-10cm beneath the surface. Nassarius ap-
peared to be a seasonal migrant into creek habitats rather
than a permanent resident; January, March and October
samples showed individuals to be restricted to the creek
mouth, while in July they extended well upstream. The
apparent seasonality of Acteocina in the marsh fauna
marked the establishment of small temporary populations
from larvae accidentally washed ashore.

Several new species records and range extensions have
been noted during this study. For example, Laternula is
not a native West Coast genus and the occurrence of
Laternula sp., cf. L. japonica (LiscHKe) at Pony Slough,
Coos Bay, Oregon, is the first known record of this species
from the North American Pacific Coast. Occurrences of
Modiolus senhousei (BENSON) at Elkhorn Slough and
Mission Bay represent a considerable southward extension
of the previously known West Coast range (36° to 48° N,
Hanna, 1966) of this introduced species. The occurrence
at Mission Bay of Siphonaria brannani STEARNS, previ-
ously known only from a single site near Santa Barbara
(34° N; Keen, 1937), also represents a range extension.
Several of the salt marsh gastropods, Assiminea trans-
lucens (CARPENTER), Phytia myosotis (DRAPARNAUD) and
Littorina newcombiana (HEMPHILL) were previously
known only from scattered localities (TRvon, 1865; Hemp-
HILL, 1876; BartscH, 1920; Pautson, 1957; Duccan,
1965; Hanna, 1966) ; this study indicates that in fact
these species are widely distributed on most marine
marshes between latitudes 25° to 48° N, 35° to 49° N, and
41° to 47° N, respectively.

Page 404

SUMMARY

Quantitative sampling at 11 sites between latitudes 27°
to 48° N reveals that the molluscan faunas of Pacific Coast
salt marshes and tidal creeks have a characteristic struc-
ture. At each site onc or two species are widely distributed
and very abundant; additional species are all represented
by small numbers of very patchily distributed individuals.
The creek faunas usually contain more species and have
a more variable species composition than do the marsh
faunas.

The geographic distribution of recurrent species groups
supports the classification of Pacific Coast molluscan
provinces proposed by VALENTINE (1966). There are
some indications that both environments in the Californi-
an Province (27° - 34° N) contain a greater variety of
species than do similar environments in the Oregonian
Province (34° - 50° N).

Now that the general composition of these faunas is
known, their community interrelationships and ecological
significance can be assessed through experimental studies
of their more common species.

ACKNOWLEDGMENTS

Financial support was made available through the Scripps
Institution of Oceanography and the United States Steel
Research Fund. I wish to thank E. W. Fager for his advice
and encouragement throughout this study. Identifications
of the molluscan material were confirmed by: E. P. Chace
(Natural History Museum, San Diego), A. Myra Keen
(Stanford University) , R. Stohler (University of Cali-
fornia, Berkeley) and D. W. Taylor (U.S.G.S., Menlo
Park, California).

LITERATURE CITED

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1920. The West American mollusks of the families Rissoellidae
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CarrRIKER, MELBOURNE ROMAINE
1967. Ecology of estuarine benthic invertebrates: A perspec-
tive. pp. 442-487 In: G.H.Laurr (ed.) Estuaries, Amer.
Assoc. Adv. Sci., Washington
Cocuran, W. A. et al.
1954. _— Principles of sampling.
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Journ. Amer. Stat. Assoc.

THE VELIGER

Vol. 11; No. 4

Duccan, ELEANor P.
1963. Report of non-indigenous marine shells collected in
the State of Washington. The Veliger 6 (2): 112
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Fircu, Joun Epcar
1953. Common marine hivalves of California. Calif.
Dept. Fish & Game Bull. 90: 102 pp.; illust.
Hanna, G DALias
1966. Introduced mollusks of western North America.
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Hauseman, S. A.
1932. A contribution to the ecology of the salt-marsh snail,
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HepcpPetTH, JoeL W.
1967. The sense of the meeting. pp. 707-710 In: G. H.
LaurF (ed.), Estuaries. Amer. Assoc. Adv. Sci., Washington
HEMPHILL, HENRY
1876. Description of a new California mollusc.
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Keen, A. Myra
1937. An abridged check list and bibliography of west North
American marine Mollusca. Stanford Univ. Press, Stanford,
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1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi+624 pp.; 10
colored plts.; 1700 text figs. Stanford Univ. Press, Stan-
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1962. A new West Mexican subgenus and new species of
Montacutidae (Mollusca, Pelecypoda) . Pacific Naturalist
3 (9): 321-328; 5 text figs. (16 October 1962)
1963. | Marine molluscan genera of western North America: an
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MacponaLp, Keiru B.
1967. Quantitative studies of salt marsh mollusc faunas from
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MacGinitie, Georce Esser

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1935. Ecological aspects of a California marine estuary. Amer.
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Meyer, K. O.

1955. | Naturgeschichte der Strandschnecke Ovatella myosotis

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NEWELL, R.
1965, The role of detritus in the nutrition of two marine

deposit feeders, the prosobranch Hydrobia ulvae and the bi-
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PALMER, KATHERINE VAN WINKLE
1958. Type specimens of marine mollusca described by P P
Carpenter from the West Coast (San Diego to British Colum-
bia). Memoir 76, Geol. Soc. Amer. i- viii + 1-376; plts.
1 - 35. New York, N. Y. (8 December 1958)
PauLson, Epwarp G.
1957. Taxonomy of salt marsh snail, Ovatella myosotis, in
central California. Nautilus 71 (1): 4-7; 1 fig.
Purpy, E, A.
1964. Sediments as substrates. pp. 238-271, In: J. Imsrie
« N. D. Newett (ed.), Approaches to Paleoecology. Wiley,

Vol. 11; No. 4 THE VELIGER Page 405

New York
SANDER, K.

1950. Beobachtungen zur Fortpflanzung von Assiminea gray-

ana LEACH. Arch. Molluskenk. 79: 147 - 149
SELLMER, G. P.

1967. Functional morphology and ecological life history of

the gem clam, Gemma gemma. Malacologia 5: 137 - 223
Tuorson, GUNNAR

1957. | Bottom communities. In Treatise on marine ecology
and paleoecology. J. W. Hedgpeth (ed.) Geol. Soc. Amer.
Mem. 67, 1: 461 - 534

Tryon, GeorcE WASHINGTON, Jr.

1865. Descriptions of new species of Amnicola, Pomatiopsis,
Somatogyrus, Gabbia, Hydrobia and Rissoa. Amer. Journ.
Conchology 1: 219 - 222

VALENTINE, JAMES W. .

1966. Numerical analysis of marine molluscan ranges on
the extratropical northeastern Pacific shelf. Limnol.
Oceanogr. 11: 198 - 212

YonceE, CHaRLES MAurRICE

1952. Studies on Pacific coast mollusks. ITV. Observation on
Siliqua patula Drxon and on the evolution within the Solen-
idae. Univ. Calif. Publ. Zool. 55: 421 - 438.

Page 406

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Vol. 11; No. 4

The Type of Tegula funebralis (A. ADAMS, 1855)

BY

RUDOLF STOHLER

Department of Zoology, University of California, Berkeley, California

94720

(Plate 65)

As WAS SHOWN EARLIER (STOHLER, 1964), Tegula funeb-
ralis (A. ApAMs, 1855) is a relatively variable species,
even though, at the same time, it appears to be one of
the least variable species of gastropods on the western
coast of North America. This variability seems to apply
only to two or, perhaps, three characters of the shell.
One of these variable characters is the configuration of
the base of the shell with which the paper already cited
dealt. Another variable character seems to be the general
shape of the shell. However, while the configuration of
the base shows essentially the same type and range of
variability from population to population, it would seem
that the variability of the shape is limited to certain
localities. This fact was also briefly mentioned in 1964.

In the latter part of August 1968 I had the opportunity
to examine the type specimens of Chlorostoma funebrale
A. Apams, 1855 in the collection of the British Museum
(Natural History) (Figures 1 to 3, Plate 65). It will be
noticed that one of the 3 specimens (Figure 3) is very
similar to the specimen illustrated in figure 2 of the
earlier report. Unfortunately no exact type locality was
given by Apams (1855, p. 317), only the very general
statement “Hab. California.”

After my return to Berkeley I re-examined the 596
specimens from 55 localities in the collection of the De-
partment of Zoology at the University of California,
Berkeley, with a view to measuring them and ascertaining

the height: width ratios of these shells. However, even
a casual glance at the 7 specimens shown in Figures 4 to
10 on the accompanying plate reveals the utter futility
of such an attempt. The extremely variable amount of
damage done to the apex of the shells by the responsible
fungus and other agents (see PEpparp, 1964) could not
possibly allow statistically valid measurements, calcula-
tions, and conclusions.

When, in spite of these acknowledged difficulties, I
measured the figured specimens, it was done only for the
purpose of ascertaining general trends in shell shape. The
Table includes also the measurements obtained from the
syntypes of the species in the BM(NH).

A careful examination of lots of specimens from a
single locality shows that the shells of young animals havea
lower H:W ratio, i. e., are relatively wider than older shells.
There seems to be no correlation with geographical origin
of the specimens. On the other hand, the specimens
shown in Figures 3 and 8 are so similar in appearance and
their respective H: W ratios that it becomes a great temp-
tation to conclude that they come from the same locality.
1.e., Duxbury Reef, Marin County, California. Yet the
specimens shown in Figures 1 and 2 would fit very well
into the population represented by the specimens pictured
in Figures 4 and 5. These were collected at La Jolla, San
Diego County. It seems more probable that the specimens

Explanation of Plate 65

Tegula funebralis (A. ADAMS, 1855)

Figure 1: Lectotype, Chlorostoma funebrale A. ADAMS, 1855.
Cuming Collection, British Museum (Natural History) no. 1968208a
Figure 2: Paralectotype, BM(NH) no. 1968208b

Figure 3: Paralectotype, BM(NH) no. 1968208c

Figure 4: Tegula funebralis, collected at La Jolla, California,

25 March, 1957; 30°52’N; 117°15/19” W; R, Stohler, coll.
Figure 5: Same data as for preceding figure

Figure 6: San Simeon, California, 18 August 1947; 35°39’30" N;
121°14’ W; R. Stohler, coll.

Figure 7: Same data as for preceding figure

Figure 8: Duxbury Reef, California, 30 October 1947; 37°53 N;
122°42’ W; R. Stohler, coll.

Figure 9: Drakes Estero, California, 18 July 1947; 38°02’N;
122°56’ W; R. Stohler, coll.

Figure 10: Elk, California, 24 June 1947; 39°06’ N; 123°42’30” W;

R. Stohler, coll.

Tue VELIGER, Vol. 11, No. 4 [SToHLER] Plate 65

Figure 1 Figure 2 Figure 3

{ Figure 5 | ‘Figure Go Figure 7

Figure 9 Figure 10

photographs by R. STOHLER

Vol. 11; No. 4

Table 1

Height and Width Measurements (in millimeters)
and Height: Width Ratios of Some Specimens
of Tegula funebralis

Height Width H:W
(mm) (mm)
Figure 1 39.45 34.45 LS
Figure 2 35,85 3272 eve
Figure 3 41.15 29.9 1.38
Figure 4 28.3 2029) ail
Figure 5 24.0 24.9 0.96
Figure 6 26.5 5) Hels
Figure 7 3515) 29.1 12
18.25 18.3 0.99
18) 18.6 0.85
11.1 12.8 0.87
Figure 8 43.85 31.0 eal
Figure 9 26.9 28.0 0.96
Figure 10 28.75 29M 0.97
14.4 19.4 0.74
14.0 17.8 0.79
GF2 10.8 0.85

(Note: the data for shells following those for Figures 7
and 10, respectively, were obtained from specimens from
the same locality. )

in the Cuming collection were actually obtained at or near
San Diego, rather than at the northern site.

It has been emphasized earlier (STOHLER, 1964) that
the tall and relatively slender specimens found in fairly
large numbers at Duxbury Reef can always be picked out
with ease in any general collection. A close comparison
of Figures 3, 7 and 8 will, I believe, substantiate this
claim, although possibly not as convincingly as the actual
examination of specimens in large collections would.

Despite the unusually large and relatively slender speci-
men from near San Simeon in San Luis Obispo County,

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Page 407

I still maintain that the shape observed at Duxbury Reef
is characteristic for the Tegula funebralis population there.
The specimen from San Simeon is but a single one in
a lot of 6, where the remaining 5 conform more to the
“normal” shape of the species, while the one from Dux-
bury Reef is one of a lot of 6, all of which are extremely
similar to this one.

Since the type lot of Apams’ species included 3 speci-
mens, one of which exhibited a shape relatively rarely
encountered in the field among large populations of
Tegula funebralis, I here select as the lectotype the shell
illustrated in Figure 1 on Plate 65; the other 2 specimens
are to be considered as paralectotypes. In my opinion
the lectotype exemplifies the most commonly encountered
representative of T: funebralis, while the paralectotypes
may be considered as illustrating the range of normal
variability of adult specimens.

The lectotype has been given the number 1968208a,
and the paralectotypes the numbers 1968208b « 1968208c,
respectively, at the British Museum (Natural History).

ACKNOWLEDGMENTS

It is a pleasant duty to acknowledge the gracious cour-
tesy with which the specimens and facilities at the British
Museum (Natural History) were made available to me
by Dr. Norman Tebble and Miss A. Fullick.

LITERATURE CITED

ApAMs, ARTHUR
1855. Descriptions of twenty-seven new species of shells from
the collection of Hugh Cuming, Esq. Proc. Zool. Soc. Lon-
don 22: 311 - 317 (8 May 1855)
PrpparD, MarGARET CAROLINE
1964. Shell growth and repair in the gastropod Tegula funeb-
ralis (Mollusca: Gastropoda) The Veliger 6, Supplement:
59 - 63; 3 text figs. (15 November 1964)
STOHLER, RupDOLF
1964. Studies on mollusk populations VI. - Tegula funebralis
(A. Apams, 1855) (Mollusca: Gastropoda) The Veliger
6, Supplement: 77 - 81; 5 text figs. (15 November 1964)

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Vol. 11; No. 4

The Litiorina ziczac Species Complex

BY

THOMAS V. BORKOWSKI

Department of Biological Sciences, Florida State University, Tallahassee, Florida 32306"?

AND

MARILYNN R. BORKOWSKI

Bureau of Commercial Fisheries, Tropical Atlantic Biological Laboratory, Virginia Key, Miami, Florida 33149

(Plate 66; 4 Text figures)

INTRODUCTION

Tue Littorina ziczac SPECIES COMPLEX has had a confused
taxonomic history. The group has been described and fig-
ured by several authors since GMeuin (1791) (D’Orsic-
ny, 1840, 1841; Puiriprr, 1847, 1851; MoOrcu, 1876;
Tryon, 1887; Bequaert, 1943; ABBoTtT, 1964) and each
has changed the nomenclature. Tryon placed them all
under the name Littorina ziczac but indicated three vari-
eties. BEQUAERT concurred and attributed the differences
in shape and sculpture to sexual dimorphism; however,
RopricuEz (1959) stated that he could almost always
separate the species into two forms showing different
vertical distributions and FRAENKEL (1968) found two
forms that differ in opercular structure. ABBotT (1964),
examining live material, concluded that there are two
species, Littorina ziczac (GMELIN) and L. lineolata p’Or-
BIGNY, but from evidence given below, it will readily be
seen that there are actually three species which are eco-
logically and morphologically distinct.

TAXONOMY

The three species of the Littorina ziczac species complex
are: Littorina ziczac (GMELIN, 1791), L. lineolata p’Or-

« This study is based upon a thesis submitted in partial fulfillment
of the requirements of the Masters of Sciences Degree at
Florida State University by the senior author.

2 Present address: Institute of Marine Sciences, 1 Rickenbacker
Causeway, Miami, Florida 33149

BIGNY, 1840, and L. lineata p’OrBicNy, 1841 (Plate 66).
Partial synonymies are given for each species.

Littorina ziczac (GMELIN, 1791)

Trochus ziczak (pars) CHEMNITZ, 1781, Syst. Conchyl. Cab.,
5, pit. 166, fig. 1599 [non-binomial]

Trochus ziczac GMELIN, 1791, in Syst. Nat., 13 ed., 1, prt. 6,
no. 3587

Littorina zigzag D’OrRBIGNY, 1842, In DE LA Sacra, Hist. Phys.
Pol. Nat. Cuba, Moll., Atlas, plt. 15, figs. 5 - 8

Littorina debilis Puivipri1, 1846, Proc. Zool. Soc. London for
1845, p. 140

Littorina d’Orbignyana Puiiprt, 1847, Abb. Beschr. Conch.,
2: 162; plt. 3, fig. 12

Cuemnitz in 1781 first figured Littorina ziczac, sensu
stricto, and based his description on the descriptions of
LisTER (1687) and FavANNE (1784). But the name as
given and spelled by CuEmnitz must be considered in-
correct because the International Commission of _Zoo-
logical Nomenclature ruled that the first 11 volumes of
CueEmnitz’ Conchylien Cabinet were nomenclatorially in-
valid (SCHENK & McMasters, 1948). LisTER’s descrip-
tion is pre-Linnean while FAVANNE uses non-binomial
names. Therefore, GMELIN’s description and spelling of
L. ziczac is the taxonomically valid citation for the species.
D’Orpicny (1842) apparently used the names zigzag and
ziczac interchangeably so that zigzag is to be considered
only as an emendation of ziczac.

Examination of Puiuippr’s (1851) and Tryon’s (1887)
figures of Littorina debilis and Puiiprr’s (1847) figure
of L. d’Orbignyana (which is L. zigzag p’OrBicny, 1842,
Cuba p. 210, plt. 15, figs. 5-8) shows that these are
synonyms of L. ziczac (GMELIN).

Vol. 11; No. 4

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After some practice, this species can be easily distin-
guished from the other two species by its lighter color,
smooth shell surface, microscopic spiral striations, dark
nuclear whorls, and by its generally larger size. As will be
seen later, vertical position on the shore and the presence
of 2 color bands in the aperture are also useful discrim-
inators.
frtor

Littorina lineolata p’Orpicny, 1840

Trochus ziczak (pars) CHemnirz, 1781, Syst. Conchyl. Cab.
5: plt. 166, fig. 1600 [non binomial]

Littorina lineolata p’Orsicny, 1840, Voy. Amér. Mérid., 5,
prt. 3, Moll., p. 392

Littorina lineata (pars) p’OrpicnNy, 1841, in DE LA Sacra,
Hist. Phys. Pol. Nat. Cuba, Moll., text p. 208; Atlas
(1842), plt. 14, fig. 25 (not 24, 26-27)

Littorina lineata Puiirri, 1847, Abb. Besch. Conch., 2: 162;
pit. 3, fig. 18 minor

Littorina jamaicensis C. B. ApaMs, 1850, Contr. Conch. 5: 71

Littorina floccosa Morcu, 1876, Malak. Blatter 23: 140

Littorina angustior var. fasciata Morcu, 1876, Malak. Blat-
ter 23: 139

Cuemnitz described and figured a variety of Trochus
ziczak which he did not name and GMELIN, 1791, copied
this description, but p’OrBicny named it in 1840. Litto-
‘rina jamaicensis, as redescribed by MOrcu, is lineolata
and L. floccosa Morcu, 1876, must be considered a syn-
onym of L. lineolata since M6rcu considered Trochus
ziczak, fig. 1600, and L. lineata v’Orsicny, fig. 25, as
synonyms of L. floccosa. MOrcu lists L. lineata, fig. 25,
as a synonym for L. angustior, var. fasciata.

As Tryon indicates, this species is more ovate, with
conspicuous striations. It is sometimes sharply keeled and
has but one adapical color band in the aperture. The
nuclear whorls are light colored and the columella tends
to be a light brown or lavender, as opposed to dark brown
in the other two. This species may also have a smooth
grayish appearance due to erosion and the presence of
boring fungi, but in this condition it cannot be mistaken
for L. ziczac.

Littorina lineata D’OrsIGNY, 1841

Littorina lineata p’Orsicny, 1841, in DE LA Sacra, Hist. Phys.
Pol. Nat. Cuba, Moll., text p. 208; Atlas plt. 14, figs. 24,
26-27 (not fig. 25)

Littorina carinata pD’OrBIGNY, 1842, in DE LA Sacra, Hist.
Phys. Pol. Nat. Cuba, Moll. (not Turbo carinatus SowEr-
By), Atlas, plt. 15, figs. 1-4; text p. 209

Littorina angustior Morcu, 1876, Malak. Blatter 23: 139

This species, although common and easy to find, has
often been overlooked because of its smaller size and
proximity to the other, more conspicuous species. D’OrR-

BIGNY Split it into 2 groups, keeled and unkeeled, and
named them carinata and lineata, respectively. This dis-
tinction, however, cannot be consistently made, since the
two extremes completely intergrade into one another.
Either name, then, is applicable to the species. The name
carinata, however, is a homonym to Turbo carinatus
SoweErsy, 1819, which is also a littorinid, but not the same
as Littorina carinata D’OrBIGNY (BEQUAERT, 1943). There-
fore, the name lineata p’Orpicny, 1841 is the correct
name, with the understanding that p’Orsicny’s figure 25
is lineolata. It is unfortunate that the two names lineata
and lineolata are so similar but p’OrBIGNYy made it clear
that he considered these different when he described line-
olata in 1840 and considered lineata as a variety of lineo-
lata. It is also unfortunate that Puiwiprr in 1847 rede-
scribed lineata p’OrBicNy in such a manner that lineata
Puiippi is clearly a synonym of lineolata.

Assort (1964) included Littorina lineata p’OrBicNy
within his concept of L. lineolata but has since agreed
(personal communication) that this third species is valid
and distinct. M6rcn’s and Tryon’s figures and descrip-
tions show that L. angustior is a synonym of L. lineata.

This species is exceedingly similar to Littorina lineolata,
but several characters serve to separate them easily. Lit-
torina lineata is generally smaller and darker than the
other two, and has tendencies towards flattened whorls
and straight rather than wavy or zigzag “flames.” Its
nuclear whorls are light and there are two or more color
bands in the aperture. A melanic form of the species is
found on the east coast of Florida and the Florida Keys.

DISTRIBUTION

The Littorina ziczac species complex belongs to the West
Indian fauna, but it is also found all around the Caribbe-
an and most of the Gulf of Mexico (HEpcPETH, 1953;
Bequaert, 1943). It is not found on the Florida west
coast and is absent from the Gulf coast of the United
States as far west as Galveston, Texas, where it again
becomes abundant. The complex extends as far south as
Uruguay and as far north as Bermuda. All three species
seem to occur together nearly uniformly, but collectors
have not been aware of the three species within the com-
plex and records are not always reliable.

On the Florida east coast, the three are distributed
decidedly differently and show separate limits of northern
and southern distributions (Text figure 1). The farthest
north any of the three ranges is Cape Kennedy, where
Littorina lineolata reaches its limit. Only occasional spe-
cimens of L. lineata are found north of Sebastian Inlet
and L. ziczac seldom occurs north of Jupiter Inlet. On

Page 410

THE VELIGER

Vol. 11; No. 4

Cape Kennedy .

Sebastian Inlet
Ft. Pierce Inlet

Hutchinson’s Island
Jupiter Inlet

North Palm Beach Inlet *

Boca Raton eod

Government Cut
Cape Florida %:

Key Largo

Tavernier ~ Jon

Islamorada OD
&

West Summerland Key we fon

e

Figure 1

Species Distribution between Cape Kennedy
and West Summerland Key
@ = Littorina lineolata; O = Littorina lineata;
CD = Littorina ziczac

the other hand, L. lineolata occurs only sporadically
south of Cape Florida on Key Biscayne, Miami, Florida.

These limits do not appear to be operable on the other
side of the Florida straits. Littorina ziczac is abundant at
Bermuda and L. lineolata is found throughout the Baha-
mas and the West Indies, as far south as Barbados. The
mechanisms determining the distribution patterns on the
Florida east coast are not known.

The vertical distributions of the three species are dif
ferent, although all live supratidally in the splash zone.
On the Florida coast, because of their latitudinal distri-
bution, either Littorina ziczac or L. lineolata is often ab-
sent or poorly represented. In those areas where all 3 are
well represented, the total number of animals is small or
the substrate is so irregular that adequate transects are
difficult to obtain.

Such results as have been obtained, however, indicate
that the order of representation of the 3 species on the
shore, from highest to lowest, is: Littorina lineata,

L. lineolata, and L. ziczac. The distribution of each species
overlaps those of the others somewhat and in areas where
L. lineolata is absent, i. e., the Florida Keys, there is no
gap in the distribution between L. lineata and L. ziczac.
Similarly, where L. ziczac is absent, L. lineolata extends as
far down the shore as the barnacle zone (Tetraclita spp.).

Table 1

Vertical distribution of Littorina lineata
and Littorina lineolata on beachrock at Boca Raton,
Florida, 14 August 1966

Distance Littorina

(meters) lineata lineolata ziczac
4.5 2 0 0
4.2 1 0 0
3.9 4 0 0
3.6 9 0 0
3.3 15 (0) 0
3.0 18 0 0
2.7 34 0 0
2.4 27 2 0
2.1 21 1 0
1.8 0 5 0
1.5 1 16 0
1.2 0 10 1
0.9 0 21 0
0.6 0 18 1
0.3 0 15 1
MHW 0 0 0

Table 1 indicates the distinctness of the vertical sepa-
ration of Littorina lineata and L. lineolata at Boca Raton,
Florida. The supratidal zone at Boca Raton is consider-
ably widened by wave action against a sandstone outcrop-
ping, which extends about 2 m above mid-tide level and
the range of distribution extends considerably onto the
top of the outcrop because of splash. In more sheltered
areas, since the tidal range is less than a meter along most
of the Florida coast, the vertical distributions are con-
siderably compressed. The observed pattern is the same
regardless of exposure or type of substrate (i.e., sand-
stone, coral limestone, broken concrete granite, cement
bags, etc., incorporated into jetties, sea walls, or com-
posing natural formations) (Borkowski, 1967). The
pattern is not changed when L. ziczac replaces L. lineolata.

SHELL MORPHOLOGY

The three species are statistically distinct in shell character
measurements. Less than one misidentification in 1000
is likely if shell measurements alone are used.

Vol. 11; No. 4

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Page 411

The procedures of Kim, Brown & Cook (1964, 1966)
were used to select key characters for the 3 species, those
characters with low intercharacter correlation and proba-
bility of misclassification. Shell characters for analysis
were chosen on the basis of their importance in describing
shell shape as discussed by FRETTER & GRAHAM (1962).
These were: height; height/width ratio; aperture diame-
ter; ratio of suture-to-suture whorl widths; spire angle;
angle of the parietal wall from the axis; and angle at the
keel (Text figure 2). The characters, lines/millimeter and

———

Figure 2
Measurements for Statistical Comparisons
A - height; B- width; C - aperture diameter; D - whorl width;
E - lines per millimeter

number of zigzags per whorl, were added as possible
specific discriminators. Distances were measured by sliding
scale Vernier calipers, accurate to 0.1 mm, supplemented
by a pair of small cartographer’s dividers on smaller dis-
tances, and angles were measured by a protractor accurate
to one degree.

Accuracy of measurement was checked by measuring
10 specimens 5 times for each character, each measure-
ment of a character separated by a sufficient time lapse
to minimize repetitive measurement bias.

Specimens of Littorina lineata and L. lineolata were
collected from Boca Raton beachrock, just north of the
Boca Raton Inlet, on 29 April 1966. Several small, indis-
criminately chosen areas were denuded of all animals
until approximately 1000 animals had been collected,
about 500 of each species. Littorina ziczac, s.5., was
scarce at this collecting site, so that a somewhat smaller
number had to be used, collected at various dates from
the vicinity.

Table 2

Intercharacter correlation coefficients for
Littorina lineata, Littorina lineolata and Littorina ziczac

Littorina
Variables lineata lineolata —ziczac
Whorl ratio:
spire angle . 0.9 0.8 - 7
height 0.8 -0.1 -0.3
height-width -0.7 —_ —
aperture diameter -0.7 0.0 0.4
lines/mm -0.7 0.03 -0.7
flames -0.8 -0.2 -0.8
angle parietal wall 1.0 0.1 0.3
angle at keel 0.0 -0.5 -0.7
Spire angle:
height -0.9 -0.9 -0.9
height-width 0.8 -0.2 -0.8
aperture diameter 0.7 -0.9 —0.7
lines/mm -0.3 0.4 -0.9
flames 0.8 0.2 -0.1
angle parietal wall 0.0 -0.9 -0.3
angle at keel 0.9 -0.98 -0.8
Height:
height-width 0.7 0.6 0.9
aperture diameter 0.98 1.0 0.99
lines/mm -0.9 -0.7 0.6
flames 0.97 0.97 0.8
angle parietal wall 0.9 0.3 0.5
angle at keel 0.9 0.9 0.97
Height-width:
aperture diameter 0.9 -0.1 1.0
lines/mm —0.7 -0.3 -0.2
flames 0.95 -0.4 -0.4
angle parietal wall — 0.0 0.8
angle at keel 0.6 0.8 0.8
Aperture diameter:
lines/mm -0.9 -0.7 0.6
flames 0.99 -0.8 -0.1
angle parietal wall 0.7 0.6 0.0
angle at keel 0.8 0.9 0.7
Flames:
lines/mm 0.7 -0.6 -0.1
angle parietal wall 0.7 0.6 0.3
angle at keel 0.4 0.2 0.3
Angle parietal wall:
lines /mm -0.6 0.0 0.2
angle at keel 0.8 0.95 0.4
Angle at keel:
lines/mm -0.7 —0.6 0.9

Table 2 shows intercharacter correlations for the three
species; several characters appear promising as discrim-
inators. The character pairs that may separate Littorina
ziczac from the other two are: whorl ratio-spire angle,

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Vol. 11; No. 4

height-lines/mm, aperture diameter-lines/mm, and angle
at the keel-lines/mm. For L. lineata, whorl ratio-height,
whorl ratio-aperture diameter, whorl ratio-angle at the
parietal wall, spire angle-aperture diameter, spire angle-
flames, spire angle-angle at the keel, height-width ratio-
flames, and angle of the parietal wall-lines/mm, and per-
haps whorl ratio-angle at the keel, appear promising.
But for L. lineolata, whorl ratio-lines/mm and height-
width ratio-aperture diameter are the only promising
character pairs. Spire angle-height-width ratio, spire angle
-lines/mm, aperture diameter-flames, flames-lines/mm,
and flames-angle of the parietal wall, may discriminate
among all three.

Further examination, however, indicates that height,
height-width ratio, aperture diameter, angle at the keel,
spire angle, and flames form a well-correlated unit. The 2
characters correlated to this unit and separating Littorina
ziczac from the other two are lines/mm and whorl ratio.
These are quite well correlated to one another so that
one or the other may be useful as a key character when
compared to the others on a common size scale such as
height. None of the above characters seems useful for
separating L. lineata from L. lineolata, but the lines in the
aperture, not included in the correlations because of
stable consistent differences, are very important in this
respect.

Littorina lineolata has only one color line in the aper-
ture. The other two species have 2 color lines in the aper-
ture. Littorina lineata and L. ziczac may have 3 color
lines depending on the darkness of the aperture pigment
and its ability to make the zone below the keel opaque.
Littorina lineolata seldom, if ever, shows more, or less,
than one line in the aperture.

A triangular graph (Text figure 3), using percentages

58560500 L. carinata
L. lineolata
——-— LL. ziczac

Aperture Diameter

Figure 3

Multicharacter Comparison of the Three Species

of a combined total of 3 character values, shows the good
discrimination among the 3 species when compared
against a size parameter, in this case, aperture diameter.
This result is based on data from specimens from several
collecting stations and from different times of the year,
as well as the original data, so that the result may be con-
sidered as indicative of actual species differences and not
just 3 variants of one population. The values used for
aperture diameter were restricted to the height range
6 - 12 mm to make the data comparable for the 3 species,
but the normal size range of all 3 species does not differ
considerably from this. The apparent overlap between
Littorina ziczac and L. lineata on the triangular graph is
not important if the size parameter is noted because the
values for L. ziczac are overlapping those of much smaller
L. lineata.

EGG CAPSULE DIFFERENCES

Another good species discriminator is the shape and size
of the egg capsule. All 3 species release free-floating egg
capsules of a uniform size and characteristic shape for
each species (Text figure 4).

Egg capsules were collected on several occasions by
putting freshly collected animals in a jar containing sea
water overnight. The water was then examined for cap-
sules which were measured, and drawn by aid of a camera
lucida.

The capsules of Littorina lineolata are considerably
different from those of the other 2 species. They are
large and bell-shaped while the other 2 types are small
and beehive-shaped. The absence of rings on the sides
of the L. ziczac capsule is the most reliable characteristic
in distinguishing the L. lineata and L. ziczac capsules. The
ridges on the L. lineata capsule may occasionally be less
distinct so that it closely resembles the L, ziczac capsule
in general appearance.

The capsules of Nodilittorina tuberculata MENKE are
very similar to the Littorina lineata capsules, although
less so than the figures of AsBotT (1954) would tend to
indicate. In the Nodilittorina capsules, as seen from the
side, the number of rings is 9 or 10. The L. lineata cap-
sules have only 6 or 7, and the rings are farther apart.

All 3 capsules, Littorina ziczac, L. lineata, and Nodi-
littorina, may appear concentric or spiral from the top
depending on the individual capsule and method of
lighting the capsules when observing.

The 3 Littorina capsules have been figured in the liter-
ature under the name L. ziczac. LrBour (1945) figured
the capsule of L. ziczac s.s., and Lewis (1960) figured
the capsules of L. lineolata and L. lineata, calling them

Vol. 11; No. 4

(@)
OR
1G) &

Figure 4

Egg Capsules of the Three Species
A = Littorina lineolata; B = Littorina lineata; C = Littorina ziczac
The line in each case represents one millimeter

large and small type L. ziczac capsules, respectively.
Aspotr (1954) and Marcus & Marcus (1963) fig-
ured the capsule of L. ziczac. ABpotr in 1964 used these
authors’ figures as partial evidence that 2 species exist,
but failed to distinguish between the Lewis L. lineata
capsule and the Lesour L. ziczac capsule.

CONCLUSIONS

Vertical and latitudinal distributions of the 3 species
strongly indicate that each is a distinct ecological entity.
Littorina lineolata has a more northern distribution in

THE VELIGER

Page 413

Florida, while L. ziczac is found farther south, and L.
lineata has a higher position on the shore than either.
Three distinct egg capsule types furnish further proof
of this distinctness.

Statistical testing for diagnostic characters has shown
that the characters, lines in the aperture and spiral striae
per millimeter, can distinguish among the 3 species with an
error of less than one in a thousand.

Certain differences in the operculum and the reproduc-
tive structures have also been noted. These are currently
being investigated and will be discussed in a later paper
on the comparative anatomy of the tropical western
Atlantic littorinids.

On this basis, it is evident that three species are present
within the Littorina ziczac species complex bearing the
names: Littorina lineata D’OrpicNy, 1841; L. lineolata
p Orsicny, 1840; and L. ziczac (GMELIN, 1791).

ACKNOWLEDGMENTS

We thank Dr. H. W. Wells and Dr. Wm. H. Heard for
facilities and guidance during preparation of the Master’s
Thesis. We also thank Dr. R. T: Abbott and Dr. J. Rose-
water for encouragement and taxonomic informaton, and
Dr. H.B. Moore, Dr. D. Moore and Dr. EM. Bayer
for critically reading this manuscript.

LITERATURE CITED

Assott, Ropert TUCKER
1954. Review of the Atlantic periwinkles, Nodilittorina, Echi-
ninus, and Tectarwus. Proc. U.S. Nat. Mus. 103 (3228) :

449 - 464; figs. 55 - 57

1964. Littorina ziczac (GMELIN) and L. lineolata OrpBIcNy.
Nautilus 78 (2): 65 - 66

BEQUAERT, JOSEPH CHARLES

1943. The genus Littorina in the western Atlantic.

sonia 1 (7): 1-27; 7 plts.
Borkowski, THomas V.

1967. An analysis of the systematics of the Littorina ziczac
species complex, intertidal gastropods of the West Indian fauna.
Masters Thesis, Florida State University.

Burma, B. H.

1948. Studies in quantitative paleontology. I. Some aspects of
the theory and practice of quantitative invertebrate paleon-
tology. Journ, Paleontol. 22: 725 - 761

FRETTER, VERA, & ALASTAIR GRAHAM
1962. British prosobranch molluscs, their functional anatomy
and ecology. London, Ray Soc. xvi + 755 pp.; 316 figs.
GMELIN, JOHANN FREDERICH
1791. Systema naturae per regna tria naturae
3021 - 3910.

John-

Editio

decima tertia, aucta, reformata 1 (6): Lipsia

Page 414

HEpcPETH, JoEL W.

1953. An introduction to the zoogeography of the north-
western Gulf of Mexico with reference to the invertebrate
fauna. Inst. Mar. Sci. 3: 110 - 224

Kim, K. C., B. W. Brown «& E. FE Coox

1964. A quantitative taxonomic study of the Enderleinellus
suturalis complex (Anoplura: Hoplopleuridae). Syst.
Zool. 12:134 - 148

1966. A quantitative taxonomic study of the Hoplopleura hes-
peromydia complex (Anoplura: Hoplopleuridae), with notes
@ posteriori taxonomic characters. Syst. Zool, 15: 24-45

Lesour, Marie V.
1945. The eggs and larvae of some prosobranchs from Ber-
muda. _ Proc. Zool. Soc. London 114: 462 - 489.
Lister, M.
1687. Historia conchyliorum. Edit. 3, 2.
Marcus, EvELYN & Ernst Marcus

1963. | Mesogastropoden von der Kiiste von Sao Paulo.
Abhandl. Math.-Naturwissensch. Klasse Akad. Wiss. & Lit.
1: 105 pp.

MOoNTCERVELLE, FAVANNE DE

1784. Catal. Cabinet M. de Montcervelle (Le Comte de la
Tour d’Auvergne).

Morcu, Otro ANpREas Lowson

1876. Synopsis molluscorum marinorum Indiarum occidenta-
lium. Malakozool. Blatter 26: 87 - 142

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Vol. 11; No. 4

Orsicny, ALcipe DESSALINES D’

(1841) -1853. Mollusques, in Ramon de la Sagra, Histoire phy-
sique, politique et naturelle de l’tle de Cuba. Paris (Arthus
Bertrand): 2 vols., atlas of 28 plts.

1840. Voyage dans l’Amérique méridionale
5 (3):

Puiuipr1, RUDOLF AMANDUS

1845-1851. Abbildungen und Beschreibungen neuer oder wenig

gekannter Conchylien. vols. 1 - 3
RopricuEz, G.

. . . Mollusques.

1959. The marine communities of Margarita Island, Vene-
zuela. Bul. Mar. Sci. Gulf and Carib. 9: 237-280; 26
figs.

Scuenk, E. T. « J. H. McMasrTers
1948. Procedure in taxonomy. Rev. ed. Stanford Univ. Press
93 pp. Stanford, California
STEPHENSON, T. A. « ANNE STEPHENSON
1952. Life between the tidemarks in North America. II. North-
ern Florida and the Carolinas. Journ. Ecol. 40: 1 - 49
Tryon, GEorcE WASHINGTON, Jr.
1887. | Manual of conchology, structural and systematic. IX.
Solariidae, Ianthinidae, Trichotropidae, Scalariidae, Cerithiidae,
Rissoidae, Littorinidae. 9: 1-488; 71 plts. Philadelphia.

Explanation of Plate 66

The Littorina ziczac species complex

Figures 1 & 2: Littorina lineata p’OrBIGNY, 1841
Figures 3 & 4: Littorina lineolata p’OrBIGNY, 1840

Figures 5 & 6: Littorina ziczac (GMELIN, 1791)

The lines on the figures equal one millimeter

Tue VELIcER, Vol. 11, No. 4 [Borkowsk1 & BorKxowskI] Plate 66

Figure 2

Figure 4

” P : y

Figure 5 he _.. : Figure 6

Vol. 11; No. 4

THE VELIGER

Page 415

Spawning and Development in the Trochid Gastropod

Euchelus gemmatus (Goutp, 1841) in the Hawaiian Islands

THOMAS M. DUCH

Science Department, Bennett College, Millbrook, New York 12545

(2 Text figures)

THE PURPOSE OF THIS PAPER is to describe the breeding
habits and development of the trochacean gastropod Eu-
chelus gemmatus (Goup, 1841) in the Hawaiian Islands.
There are no previous reports of spawning periodicity or
laboratory observations on the spawning behavior of a
tropical trochid.

Euchelus gemmatus is a small (2 to 4mm in height)
trochid which is common in shallow water along the shore-
lines of the Hawaiian Islands. Large populations of this

‘animal occur on the lower surfaces of dark, mottled rocks
~ in tidepools and in shallow offshore waters where there
is moderate turbulence. The snails are usually found con-
gregated in crevices and beneath algae, but they are also
able to cling to exposed rock surfaces.

METHODS

Forty animals averaging 2 to 4mm in height were collec-
ted in October, 1965, and maintained in 2 one-gallon con-
tainers, 20 in a non-aerated jug and 20 in a container in
which the turbulence of the natural habitat was simu-
lated by means of an air stone. The animals were main-
tained in both situations for a period of 2 years and
spawning and development occurred equally well among
snails in both containers.

Egg masses which were deposited on the glass walls
of the containers were removed by means of dissecting
needles and placed on microscope slides which were set
on a stand in a finger bowl. The egg masses were then
easily removed for microscopic examination and could be
replaced without injury. Daily changes of sea water
and a soft jet of air bubbles directed onto the egg
masses increased the survival rate of the developing em-
bryos by reducing the numbers of ciliates which often
congregate about the egg masses.

SPAWNING HABITS

Spawning in Euchelus gemmatus occurs from late De-
cember through April. The beginning of the spawning
period is marked by the secretion of a gelatinous coat
over the surface of the female shells; the covering remains
on the shells throughout the spawning period. Subsequent
to the appearance of the coat, pair formation occurs, a
smaller male becoming attached to the left apertural sur-

vitelline membrane

spherical enclosure

ess

oval capsule

—

120p

egg mass

_—————

10 mm

Figure 1

Euchelus gemmatus (Gou.p, 1841)
Egg capsule and egg mass

Page 416

face of the female shell. Because trochids presumably do
not copulate, it is suggested that the male’s position near
the aperture of the female shell permits gametes to pass
into the mantle cavity of the female.

During the period of observation 13 egg masses were
deposited on flat surfaces from 1 to 5cm above the sur-
face of the water; the egg masses were deposited at
twilight and at the time of high tide, with 9 of the egg
masses deposited in March at the time of “extreme”
high tide. After the initial spawning of one or two pairs
of trochids, the remaining pairs spawned with increasing
frequency.

Although it was difficult to observe egg deposition be-
cause of the initial transparency of the egg mass, it was
possible to see the general pattern. Each female moves
over a path about 15 mm in length in a counter-clockwise
direction, depositing the egg mass (Figure 1). The egg
masses are 3 to 14mm in length, 2 to 3mm wide, and
about 1 mm thick, and they are deposited in the form of
3 to 5 loops. Females with smaller shells produce masses
with fewer loops. The egg masses are flexible, jelly-like
structures, enveloped by a thin, transparent integument
which becomes translucent within an hour after depo-
sition. The wall of the mass is tubulate and several layers
of egg capsules are contained within the translucent ma-
trix except at the tapered tips of the loop.

The eggs are brown and spherical, about 50 in dia-
meter. Each egg is surrounded by a relatively homogenous
sphere of albumen-like material, that nearest the egg
semi-fluid and clear, the remainder more viscous. The
egg and semi-fluid material are in turn surrounded by the
vitteline membrane which is contained within a spherical
elastic enclosure suspended in an outer oval capsule (Fig-
ure 1). The elastic enclosure stretches slowly as the em-
bryo develops, exhibiting a fibrillar appearance. Prior to
hatching, when the envelope is ripped by the embryo, the
elastic enclosure almost fills the outer capsule. The cap-
sules, which are closely packed within the matrix of the
egg mass, are irregularly shaped because of crowding.

DEVELOPMENT

Early cleavage is initiated immediately after the egg mass
is deposited. The first 2 cleavages are equal and occur
within a few hours at a water temperature of 23°C.
Subsequent cleavages result in the formation of large
macromeres and small micromeres which envelop the
macromeres, giving rise to the blastula within 2 days. A
ciliated gastrula, which is flattened at the poles, de-
velops during the next 2 or 3 days. The eggs at the surface
of the egg mass are generally more advanced than are
those near the center of the matrix,

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Vol. 11; No. 4

The trochophore is recognizable 2 to 4 days after the
egg mass is deposited and persists within the egg capsule
for 3 to 5 days. The velar lobes are lightly pigmented
and, although ciliated, the only noticeable motion is a
slight vertical vibration. In the dorsal region a thickened
clump of cells, the shell gland, becomes visible, and by
the end of the third or fourth day this area of the embryo
is covered by a thin sheet of shell matrix. The shell
expands, occupying about # of the dorsal surface, in a
trochophore ready to metamorphose. Anterior and vent-
ral to the visceral mass are 2 large, rounded protuber-
ances which fuse, forming the foot of the veliger. As the
foot develops, statocysts form as 2 elongated depressions
behind the region of the mouth. There are 7 to 12
vibrating, calcareous grains in each statocyst, which grad-
ually “sink” into the foot and are covered by pedal tissue.
Rudiments of eyes, as 2 dark pigmented areas, are also
visible at this period.

—epipodial tentacle

shell

shell

— SSS
250p

Figure 2
Euchelus gemmatus (Goutp, 1841)
Dorsal view of the late veliger stage

The veliger becomes recognizable 3 to 9 days after
spawning, with a cap-like shell and anteriorly projecting
foot (Figure 2). There is now a series of 5 protuber-
ances: 3 small protuberances from which develop the
epipodial tentacles and 2 larger protuberances which
develop into the large anterior tentacles. Each protuber-

Vol. 11; No. 4

ance has a concentration of cilia about it, moving more
or less in a counter-clockwise direction.

The visceral hump with its cap-like shell slowly shifts
posteriorly, and torsion occurs within the next 24 hours.
The shell seems to be deposited by the mantle as a thin
translucent covering. Shortly after the translucent cover-
ing has been deposited, a shingle-like layer is deposited
and continues to develop, soon occupying the entire dorsal
surface behind the prototroch. On the posterior side of
the foot a thin functional operculum develops, which in
a short time is large enough to close the aperture of the
shell. The operculum is first laid down as a series of
“cellular” blocks which fuse and ultimately form the “true
operculum.” At the same time, the shell, which was very
thin, becomes thicker and calcareous, with a furrowed
surface.

As early as the second week of development, the veliger
is seen actively moving about in the fluid of the egg
capsule. The movement is generally limited to extending

THE VELIGER

Page 417

and quickly retracting the foot into the shell. The re-
tractive response seems to be associated with changes in
light intensity and slight external vibrations. Shortly
after, when the albumen-like material is gone, the veliger
becomes very active and rips the egg capsule with its
radula, leaving as a semi-crawling larva. Though the lar-
vae are capable of darting movement, they remain rela-
tively inactive, their movement localized to midway on
the sides of the container. The larvae grow to 2mm in
height and, as the next breeding season approaches, the
snails pair up and begin moving up and down the sides
of the container with the times of the tides, laying egg
masses in the same fashion as their parents.

ACKNOWLEDGMENT

I greatly appreciate the advice and comments of Dr.
E. Alison Kay, University of Hawaii.

Page 418

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Vol. 11; No. 4

New Records of Nudibranchs from New Jersey

BY

ROBERT E. LOVELAND, GORDON HENDLER

AND

GARY NEWKIRK !

Rutgers, the State University, New Brunswick, New Jersey 08903

IN A RECENT PAPER by FRANZ (1968), it was suggested
that the paucity of records for species of nudibranchs
occurring in New Jersey can be attributed to the absence
of systematic and diligent collecting along the coastline.
Although the general substrate of muddy-sand in New
Jersey might restrict the number of species of nudibranchs,
there are suitable outcroppings of man-made jetties and
floating wharves where nudibranchs may occur. It is
probable that new state records will occur for various
kinds of invertebrates as collecting intensity increases.
This was shown to be true for benthic, macroscopic algae
by Taytor et al. (1968). The following list represents
new information concerning the occurrence and distribu-
tion of nudibranchs along the Atlantic coast of North
America.

NUDIBRANCHIA
DENDRONOTACEA

1. Dendronotus frondosus (Ascantus, 1774), as Dendro-
notus arborescens in MINER (1950). Occurrence: Shark
River, April-May, 1968. Previous distribution, from Bay
of Fundy to Long Island Sound (Moore, 1964).

Many hundreds of this species were found in associa-
tion with Tubularia on a floating wharf. Color ranged
from nearly pure white to dark brown. ODHNER (1939)
asserts that the white form of Dendronotus is predominant
in deep water.

2. Doto coronata (GmeEuN, 1791), as Idulia coronata
in Moore (1964). Occurrence: Shark River, May, 1968.
Previous distribution, from Bay of Fundy to Long Island
Sound (Moore, 1964). Originally described from Great

' We are indebted to Dr. David R. Franz, Biological Sciences,
University of Connecticut, for criticizing this manuscript and
for confirming our identifications. This work was supported,
in part, by a grant from the Joint Investigatory Committee for
Environmental Effects on Thermal Addition in Barnegat Bay.

Egg Harbor, New Jersey, by VerRILL & SmirH (1873),
but rediscovered only recently.

DORIDACEA

3. Acanthodoris pilosa (O. EF Mier, 1776), as Doris
pilosa in ALDER & Hancock (1845 - 1855) and as Doris
bifida in VERRILL & SmirH (1873). Occurrence: Delaware
Bay, May, 1968. Previous distribution, throughout New
England (Moore, 1964). Reported, however, from Mary-
land by Marcus (1961).

A single specimen of this dorid was collected from
among unidentified encrusting ectoprocts on the back of
a large Limulus.

EOLIDACEA
4. Trinchesia aurantia (ALDER & Hancock, 1842), as
Eolis aurantia in ALDER & Hancock (1845 - 1855), as

Montagua gouldu in VERRILL & SmitH (1873), as Catri-
ona aurantia in AppotT (1954), and as Cratena aurantia
in Moore (1964). Occurrence: Shark River, April-May,
1968. Previous distribution, from New Hampshire to
Woods Hole (Moore, 1964). j

This species occurs commonly on the Connecticut shore
on Tubularia (Franz, personal communication) ; how-
ever, this is the first instance of Cratena aurantia in New
Jersey. Specimens were found in association with Dendro-
notus frondosus among thick growths of Tubularia on
floating wharves, although its food source could not be
ascertained.

5. Eubranchus pallidus (ALDER & Hancock, 1842), as
Folis picta in ALDER & Hancock (1845 - 1855) and as
Aeolis picta in Goutp & Binney (1870). Occurrence:
Shark River, February, 1967. Previous distribution, from
Bay of Fundy to Rhode Island (Moore, 1964).

This single specimen was found very low on the inter-
tidal zone of an exposed jetty, among unidentified hyd-
roids. It is interesting to note that Lomentaria orcadensis,
a rare species of boreo-arctic algae for New Jersey, was

Vol. 11; No. 4

THE VELIGER

Page 419

also found from this same collection. Franz (personal
communication) reports Eubranchus pallidus as being
commonly found on Obelia along the Connecticut shore.

6. Tergipes tergipes (ForskaL), as Eolis despecta in
Auper & Hancock (1845 - 1855), as Aeolis despecta in
GouLp & Binney (1870), as Tergipes despectus in Moore
(1964) and in Franz (1968). Occurrence: Shark River,
May, 1968; previously a single animal collected at Shark
River by Franz (1968) in November, 1961. Previous
distribution, from Bay of Fundy to Rhode Island (Moore,
1964).

A second specimen of Tergipes despectus reported for
New Jersey in 7 years qualifies this species as rare.

7. Aeolidia papillosa (Linnaeus, 1761), as Eolis papil-
losa in AtpER & Hancock (1845-1855) and as Aeolis
papillosa in Goutp & Binney (1870) and. in MINER
(1950). Occurrence: Shark River, May, 1965 - 1968;
Manasquan River, May, 1967 - 1968; also reported by
Franz (1968) for August, 1964. Previous distribution,
from Bay of Fundy to Woods Hole (Moore, 1964) ;
from Greenland to Rhode Island (Miner, 1950). Franz
(personal communication) reports this species as being
common in Connecticut.

When first collected by Franz, this species was con-
sidered quite rare for New Jersey. It has been our expe-
rience that Aeolidia is rapidly becoming established in the
estuaries of New Jersey as evidenced by its rather common
occurrence in both Shark River and Manasquan Inlet
during the spring of 1968 (Alan Schwartz, Rutgers Uni-
versity, personal communication). Specimens of 3 - 4 cm
have been taken from both localities. Both of these inlets
are characterized by water of high salinity and rock
jetties. Although our records for this nudibranch are from
the rocks, Schwartz reports these animals from within the
estuary.

DISCUSSION

It is becoming more evident that the marine fauna of
New Jersey, especially the nudibranch mollusks, repre-
sents a southern extension of the New England fauna,
as was originally suggested by Franz (1968). It is not
known, however, whether the species reported in this
paper have recently invaded New Jersey or have been
here unnoticed all along. Invasion by northern species of
algae is apparently occurring, as in the case of Codium
fragile (see Taytor, 1967). It is, therefore, possible that
the eggs of nudibranchs, or the animals themselves, are
being transported southward along the Atlantic coast
by the same mechanism that is moving the algae. Once
in New Jersey, they might then establish themselves in

the man-made, compatible habitats that are being rapidly
established in this densely populated coastal state.

The white form of Dendronotus is reported generally
from deep water (OpHNER, 1939). However, of the
specimens collected in Shark River, at least 3 out of
approximately 20 were of the white variety. All of these
specimens were on floating wharves and, therefore, never
deeper than several feet below the surface of the water.
One white specimen has been sent to Dr. D. R. Franz of
the University of Connecticut for further study.

We have included a listing of the nudibranchs reported
to date from New Jersey in Table 1.

LITERATURE CITED

ApsoTtt, RoBERT TUCKER

1954. American seashells. Princeton, New Jersey, D. van

Nostrand Co., Inc.; xiv+541 pp.; 100 text figs.; 40 plts.
ALDER, JosHUA, & ALBANY Hancock

1845 - 1855. A monograph of the British nudibranchiate mollusca
Ray Soc. London, 438 pp.

BLEAKNEY, J.S. « KantAuLono H. BaiLey

1967. Rediscovery of the salt-marsh sacoglossan Aldcria mo-
desta LovEN in Eastern Canada. Proc. Malacol. Soc.
London 37: 347 - 349

Cuameers, L. A.

1934. Studies on the organs of reproduction in the nudi-
branchiate mollusks. Bull. Amer. Mus. Nat. Hist. 66:
599 - 641

Franz, Davip R.

1967. On the taxonomy and biology of the dorid nudibranch
Doridella obscura. The Nautilus 80: 73 - 79

1968. Occurrence and distribution of New Jersey Opistho-
branchia. The Nautilus 82 (1): 7-12 (July 1968)

Gou.Lp, Aucustus AppISON & WILLIAM G. BINNEY
1870. | Report on the Invertebrata of Massachusetts. 24 ed.

vit524 pp. Boston
LowpbeEn, R.
1966. |New Jersey Mollusca checklist -— marine. The

Jersey Sheller 1: 56 - 64
Marcus, Ernst

1961. Opisthobranchia from North Carolina.

Elisha Mitchell Sci. Soc. 77 (2): 141-151
Miner, Roy WALDO

1950. ‘Field book of seashore life.

to 251; G. PR Putnam’s Sons, New York
Moore, GeorcEe M.

1964. Shell-less Opisthobranchia, pp. 153 - 164; 27 figs. In:
R.1I.Smrru (ed.) Keys to marine invertebrates of the Woods
Hole Region. Contr. 11, System.-Ecol. Program, Marine
Biol. Labor., Woods Hole, Mass.

Opune_r, Nits Hyatmar

1939. | Opisthobranchiate mollusca from the western and north-
em coasts of Norway. Kgl. Norska Vidensk. Selsk. Skr.
1: 1-93; 59 text figs.

Taytor, J. E.

1967. Codium reported from a New Jersey estuary. Bull.

Torrey Bot. Club 94: 57 - 59

Journ.

xv +888 pp.; plts. 1

Page 420

THE VELIGER

Table 1

Checklist of Nudibranchs Reported for New Jersey

Species

SACOGLOSSA

Elysia catula (Goutp, 1870)

Elysia chlorotica (Goutp, 1870)

Alderia modesta (LovEN)

NUDIBRANCHIA

Doridacea
Acanthodoris pilosa (Mier, 1776)
Doridella obscura VERRILL

Polycerella emertont VERRILL
Polycerella conyma Marcus
Tenellia fuscata (Goutp, 1870)

Dendronotacea
Doto coronata (GmEutN, 1791)

Dendronotus frondosus (ASCANIUS,
1774)
Eolidacea
Aeolidia papillosa (Linnagus, 1761)

Cratena pilata (Goutp, 1870)

Trinchesia aurantia (ALDER & Han-
cock, 1842)
Eubranchus pallidus (ALDER & Han-
cock, 1842)
Tergipes tergipes FORSKAL

Collected by

VERRILL & SMITH (1873)

F Phillips (Franz, 1968)

VERRILL & SMITH (1873)

K. Clark (Franz, 1968)

Franz (1968)

K. Clark (Franz, pers. comm., 1968)

Loveland ¢ al., 1968

VERRILL & SMITH (1873)

Franz (1967)

LowpeEN (1966) as Corambella bara-
tariae Harry

D. Dean (FRANz, 1968)

CuHamsers (1934)

Franz (1968)

CHampers (1934)

Franz (1968)

Hendler (personal communication)

VERRILL & SMITH (1873)
Loveland et al., 1968
Loveland e¢ al., 1968

Franz (1968)

Schwartz (Loveland e¢ al., 1968)
Lowpen (1966)

Franz (1968)

FE Phillips (Franz, 1968)
Loveland et al., 1968

Loveland et al., 1968

Franz (1968)

Taytor, J. E., E.T. Mout « R. E. Loveranp
1968. New and rare records of benthic marine algae from
New Jersey (in prep.)
VERRILL, Appison E. « S. I. Smiru
1873. Report upon the invertebrate animals of Vineyard
Sound with an account of the physical characters of the

region.

Rep. U.S.Comm. Fish. for 1871-1872; prt. I:

295 - 778, Washington.

Vol. 11; No. 4

Location

Great Egg Harbor
Barnegat Bay
Great Egg Harbor
Cheesequake Park
Shark River
Cheesequake Park

Delaware Bay
Great Egg Harbor
Delaware Bay
Not given

Raritan Bay
Barnegat Bay
Jarvis Sound
Barnegat Bay
Shark River
Delaware Bay

Great Egg Harbor
Shark River
Shark River

Shark River
Manasquan Inlet
Not given
Delaware Bay
Barnegat Bay
Shark River

Shark River

Shark River

Vol. 11; No. 4

THE VELIGER

Page 421

Nomenclatural Changes

for the New Species Assigned to Cratena by MACFARLAND, 1966

RICHARD A. ROLLER

1127 Seaward Street, San Luis Obispo, California 93401

(2 Text figures)

Frank Mace MacFarLANnpb’s POSTHUMOUSLY published
volume on the opisthobranchiate mollusks of the Pacific
coast of North America (1966) has been used by many
research workers who felt the need for a comprehensive
reference work on this group. The difficult task faced by
the late Olive H. MacFarland in collating her late hus-
band’s notes and drawings was mentioned by G Dallas
Hanna in his preface to the monograph, and he antici-
pated that corrections and additions might be required
. at a later date.

Burn (1968) commented on the need for Pacific coast
workers to synonymize and validate the species described
by MacFarland as new. The present contribution demon-
strates the need for the generic transfer of 7 of these new
species.

MacFar.tanp (1966) introduced 7 new aeolid species
(abronia, albocrusta, flavovulta, fulgens, rutila, spadix, and
virens) and assigned them to the genus Cratena BEeRcH,
1864. Mrs. MacFarland added a note (op. cit., p. 332)
that stated, “There is evidence among his notes and a
letter from Nils Odhner that the author was carefully
evaluating the status of the proper genus name for this
group of aeolids prior to his death in 1951. However, the
manuscript is published as left by him because he had not
seen the later publications. The paper by WinckworTH
(1941, pp. 146-149) in which Catriona is substituted
for Cratena is especially significant.”

Species of the genus C’ratena have the cleioproctic anal
position; however, all 7 of the new species of aeolids
described by MacFarland have the acleioproctic anal po-
sition. Therefore the 7 species cannot be retained in
Cratena.

Opinion no. 777 of the International Commission on
Zoological Nomenclature (1966) added Trinchesia InE-
RING, 1879 to the Official List of Generic Names. In his
proposal for stabilization of Trinchesia, LEmcueE (1964)

stated that “there is still the possibility that Catriona
could be used for a genus independently of
Trinchesia.” BurN (1964) and EpmMunps (1968) main-
tained Catriona as a valid genus separate from Trinchesia,
and several other specialists (personal communications)
have concurred with this separation.

The two genera may be characterized as follows:

Catriona Trinchesia
Acleioproct Eolidacea Acleioproct Eolidacea
Radula uniseriate with cusp Radula uniseriate with cusp
shorter than lateral as long as, or longer than,
denticles lateral denticles
Small secondary denticles No small accessory denticles
interspersed among lateral among lateral denticles of
denticles of radular tooth — radular tooth
Denticles on cutting edge of Denticles on cutting edge
jaw in form of fine bristles, of jaw not bristled
or bristled rodlets

Of the 7 new species of MacFarland, 6 have the
characteristic radular tooth shape and jaw denticulation
of Trinchesia, while the remaining one has the radular
tooth shape and jaw denticulation of Catriona (Figure
1). Therefore I propose that the following species be
assigned to Trinchesia:
abronia, albocrusta, flavovulta, fulgens, and virens,

SPHON & Lance (1968) synonymized Cratena rutila
MacFarianb, 1966 with Catriona lagunae (O’Donoc-
HUE, 1926). Since C. lagunae cannot be included in
Catriona because of its typical Trinchesia radular tooth
shape and jaw denticulation, it must be transferred to
Trinchesia as a comb. nov., Trinchesia lagunae (O’Donoc-
HUE, 1926).

As shown in Figure 1, the radular tooth shape and
denticulation of the jaw of Catena spadix MacFarLanp,
1966 are obviously different from the remaining 6 species.

Page 422

Radular Teeth

Masticatory Border of Jaw

Cratena
rutila

Cratena
fulgens

Cratena
albocrusta

Cratena
flavovulta

A

X650

Cratena
abronia

Cratena
virens

Cratena
Spadix

Figure 1

Specific differences of the species assigned to Cratena by
MacFarzanp, 1966

THE VELIGER

Vol. 11; No. 4

MacFar1anp (1966, p. 354) notes that the penial stylet
of C. spadix is not “as in the allied forms.” This species
was assigned to Catriona by EpmuNDs (1968).

SPHON & Lance (1968) provided an “abbreviated syn-
onymy” of certain of MacFarland’s 1966 species. One
of these species was Cratena spadix MAcFarLanp, 1966
which they synonymized with Cuthona alpha BaBa &
Hamatanl, 1963. The findings below support their syn-
onymy.

Lance, (1966) reported the collection of 6 specimens
of Cuthona alpha from San Diego, Newport Harbor, and
Santa Barbara, California All specimens were collected
on boat landings in protected waters. The author collected
2 specimens of C. alpha on 9 February 1968 from boat
landings in Morro Bay, California. They agreed with the
description of the holotype of C. alpha as given by BaBa
& Hamatant (1963). They also agreed with the descrip-
tion of Cratena spadix MacFarLanp.

Careful study of the descriptions of these two species
showed them to be alike in almost all of their character-
istics, but different in 2 important points: the presence
or absence of a penial stylet, and differences in the
masticatory margin of the jaw.

MacFar.anp in 1966 described the penial stylet of
Cratena spadix as a “thin-walled chitinous armature which
does not project beyond the surface of the tip as in the
allied forms.” BabA & HAMATANI (1963) described Cutho-
na alpha as a “mosaic species which has the decided rad-
ula type of Catriona on one hand, and the presumed penis
peculiarity of Cuthona on the other hand.” Cuthona was
described as having an unarmed penis. The authors ten-
tatively placed the species in the genus Cuthona ALDER &
Hancock, 1855.

It appeared to the author that the 2 species might be
synonymous, if Cuthona alpha did in fact possess a minute
non-protruding stylet. This possibility was suggested to
Dr. Baba, and upon re-examination (at high magnifica-
tion) of paratype material he found a very short non-pro-
truding stylet (personal communication).

Epmunps (1968) characterizes Catriona WINCKWORTH,
1941 as having “‘denticles on cutting edge of jaw in the
form of fine bristles . . . short, straight stylet on penis.”
He assigned Cratena spadix to the genus Catriona and
also proposed the possibility of the inclusion of Cuthona
alpha in Catriona, “if details of the preradula and jaws
were known, and if the genus were extended to include
species without a penial stylet.” The presence of a stylet
in Cuthona alpha makes the extension of the genus Catri-
ona unnecessary in order to include this particular species.

The masticatory margin of the jaw of Cratena spadix,
as described by MacFartanp (1966) was very different
from the remaining 6 species of Cratena he described

Vol. 11; No. 4

THE VELIGER

Page 423

at the same time. The margin of C. spadix was listed as
bearing “a series of transverse rod-like thickenings which
project beyond the margin as closely set blunt rodlets or
ridges laterally in contact with each other ... The
rodlets are nearly straight near the hinge but become
slightly curved as the tip is approached. At this tip, with
high magnification, the surface seems to show minute
spines.”

In unpublished notes of MacFarland, a footnote by
Mrs. MacFarland states, “With 5D magnification the sur-
face of these rodlets at the lower end show a rough sur-
face of minute spines. (Frank made one drawing).” This
drawing (Figure 2) is reproduced herein for the purpose
of augmenting MacFarLanp’s original description, since
this drawing was not included in the monograph.

Figure 2
Rodlets at lower end of masticatory margin of mandible
(from unpublished notes of F.M. MacFarianp)

Basa (personal communication) re-examined paratype
material of Cuthona alpha, and found that the denticu-
lations of the jaw edge “appeared as if they were covered
with short, thick bristles.”

Members of the genus Cuthona ALDER & HANcock,
1855, are considered to have no penial stylet. Since Cu-
thona alpha is now known to have a short, straight stylet
and bristled denticles on the jaw edge, the author pro-
poses that the species be known by the new combination,
Catriona alpha (BaBA & Hamatant, 1963).

I would like to thank Dr. Kikutar6 Baba for his kind
assistance in this study, and also Mr. Allyn G. Smith of
the California Academy of Sciences who made it possible
for me to use the unpublished notes of Dr. and Mrs. F
M. MacFarland.

LITERATURE CITED

Basa, KikuTArO & Iwao HaMaTANI
1963. A cuthonid, Cuthona alpha n. sp., with a radula of
Catriona type (Nudibranchia-Eolidoidea) . Publ. Seto Mar.
Biol. Lab. 11 (2): 339 - 343; plt. 11
Burn, RosBert
1964. Descriptions of Australian Eolidacea (Mollusca: Opis-
thobranchia). 2. The genera Nossis, Eubranchus, Trinchesia,
and Toorna. Journ. Malacol. Soc. Austral. 8: 10 - 22; figs.
1-21
1968. Archidoris odhneri (MacFarianp, 1966) comb. nov.,
with some comments on the species of the genus on the
Pacific Coast of North America. The Veliger 11 (2):
90 - 92 (1 October 1968)
EpmMunps, MaLcoLm
1968. Eolid Mollusca from Ghana, with further details of
West Atlantic species. Bull. Mar. Sci. 18 (1): 203 - 219;

figs. 1-9
Lance, JAMES RoBERT
1966. New distributional records of some Northeastern Pacific

Opisthobranchiata (Mollusca: Gastropoda) with descriptions
of two new species. The Veliger 9 (1) : 69 - 81; 12 text figs.
(1 July 1966)
Lemcuer, HENNING
1964. Proposed stabilization of the generic name Trinchesia
IneERING, 1879, and suppression under the plenary powers of
Diaphoraeolis IREDALE « O’DonocuueE, 1923 (Class Gastro-
poda). Z.N. (S.) 1106 Bull. Zool. Nomencl. 21 (1):
52 - 55
MacFarianp, Frank Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
O’DonocHuE, CHARLES HENRY
1926. _A list of the nudibranchiate mollusca recorded from the
Pacific coast of North America, with notes on their distribution.
Trans. Roy. Canad. Inst. 15 (2): 199 - 247.
Opinion 777
1966. Trinchesia InErinc, 1879 (Gastropoda), added to the
Official List of Generic Names. Bull. Zool. Nomencl.
23 (2-3): 95-97
SpHON, Gate G., Jr. « James RoBert LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif. Acad.
Sci. 36 (3): 73-84; 1 fig. (25 September 1968)
WINCKWoRTH, RONALD
1941.. The name Cratena.
24 (4) :146 - 149

Proc, Malacol. Soc. London

Page 424

THE VELIGER Vol. 11; No. 4

An Annotated List of Opisthobranchs
from San Luis Obispo County, California

BY

RICHARD A. ROLLER

1127 Seaward Street, San Luis Obispo, California 93401
AND

STEVEN J. LONG

126 Esparto Avenue, Pismo Beach, California 93449

(1 Map)

TO DATE VERY LITTLE INFORMATION has been published on
the opisthobranchs of San Luis Obispo County, Califor-
nia. MacFarianp (1925), Hanna (1951), and STEIN-
BERG (1963a) have mentioned single species collected from
the county. Lance (1961) and STEINBERG (1963b) in-
clude the county within the scope of their distribution
lists. However, most studies of California opisthobranchs
have been done in the Dillon Beach, San Francisco Bay,
and Monterey Bay areas to the north, and in the Santa
Barbara, Laguna Beach, and San Diego areas to the
south, leaving a gap in data for the central coast area.

This list is an attempt to fill that gap and to add to the
data recently reported by SPHON & LANcE (1968) for the
southern bordering county, Santa Barbara. We have tried
to continue their basic format, so that both lists may be
used conveniently by workers of the central coast area.
The present list, together with the lists of BRooksHIRE
(1968a, 1968b) form a reasonably complete catalog of
the molluscan fauna of San Luis Obispo County.

Specimens were collected during field trips made by
the authors, either jointly or separately, over a 2-year
period. The entire coastline of the county was considered
for possible collection sites, and 7 general localities were
chosen (see Map, Figure 1). The coastline north of San
Simeon is practically inaccessible except by boat, and only
sandy beaches are found south of Shell Beach. Adjacent
collecting sites of similar habitat were joined under one
locality name. Most of the specimens were collected from
the intertidal region; however, an attempt was made to
include ecologically different regions (e. g., mud flats,
boat docks) in the collections.

«County Boundary

35°40’ San Simeon
N Ria
Piedras
Blancas

\Cayucos
Pt. Estero “48; f

\e Morro Bay
p Nog

a
B San Luis Obispo
Pe Rock Avila
“<i? Shell Beach

Pt. San Luis \a Pismo Beach

an «
ie Cormeen

| i egroan

Pt. Sal \:

Pt. Purissima a

Pt. Arguello &

Pt. Conception\,.,:: Hee,
Figure 1

San Luis Obispo County, California - Collecting Stations

Vol. 11; No. 4

Of the 67 species listed, 62 species represent new rec-
ords for the county. Only Acanthodoris lutea, Aldisa san-
guinea, Bulla gouldiana, Trinchesia abronia, and Trinche-
sia lagunae have been recorded previously from the
county.

The currently known range of each species is listed;
range extensions, indicated by an asterisk (*) preceding
their names, are included for the following species:
Acanthodoris nanaimoensis, Cadlina sparsa, Doto kya,
Pleurobranchus californicus denticulatus, Trapania velox,
and Trinchesia flavovulta.

We have used the designations of SPHON & LANCE
(1968) for the relative abundance of each species, and we
must reiterate that these designations are arbitrary. No
quantitative data are published for different habitats, sea-
sonal fluctuations, or distribution within the county.

The names used for the species are the most currently
accepted ones, and are the same as those used by SPHON &
Lance (1968), except for the following changes:
Archidoris odhneri (= Austrodoris odhneri)

Catriona alpha (== Cuthona alpha)

Doridella steinbergae (= Corambella steinbergae)
Doriopsilla albopunctata (= Dendrodoris albopunctata)
Trinchesia albocrusta (== Catriona albocrusta)

_ Trinchesia lagunae (= Catriona lagunae)

Chromodoris macfarlandi (== Glossodoris macfarlandi)
Hypselodoris californiensis (== Glossodoris californiensis)
Hypselodoris porterae (== Glossodoris porterae)

A list of inferred species, which may occur along the
San Luis Obispo County coastline, follows the list of
collected species. These inferred species were reported pre-
viously in the literature.

We acknowledge the assistance of Mr. Gary McDonald
in furnishing collecting data and specimens.

LIST or COLLECTED SPECIES

Acanthodoris lutea MacFartLanp, 1925

Frequent. Intertidal to 30 feet.
Cayucos, Lion Rock, Hazard Canyon, Shell Beach.
Dillon Beach to Point Loma.

* Acanthodoris nanaimoensis O’DonNocHUuE, 1921

Rare

Shell Beach.

Vancouver Island to Shell Beach (Vancouver Island to
Moss Beach).

Acanthodoris rhodoceras COCKERELL & EvioT, 1905
Avila, Shell Beach.

THE VELIGER

Page 425

Frequent
Dillon Beach to Coronados Islands.

Aegires albopunctatus MacFar.anp, 1905

Common. Intertidal to 20 feet.

Morro Bay Docks, Hazard Canyon, Shell Beach.

Vancouver Island to Coronados Islands; Bahia de Los
Angeles

Aeolidia papillosa (Linnaeus, 1761)

Frequent

San Simeon, Cayucos, Morro Bay Docks, Hazard Canyon,
Lion Rock, Shell Beach.

Cosmopolitan.

Aglaja ocelligera (BERcH, 1894)

Rare
Morro Bay mud flats.
Alaska to Santa Barbara.

Aldisa sanguinea (Cooper, 1862)

Frequent

Cayucos, Hazard Canyon, Lion Rock, Avila, Shell Beach.
Bodega Bay to San Diego; Japan.

Ancula lentiginosa FARMER & SLOAN, 1964

Rare. On pulled wharf pilings.
Avila (Port San Luis Wharf ).
Monterey Yacht Harbor to Bahia de Los Angeles.

Ancula pacifica MacFarianp, 1905

Frequent
Hazard Canyon, Avila, Shell Beach.
Moss Beach to Point Loma.

Antsodoris nobilis (MAcFaARLAND, 1905)

Common. Intertidal to 50 feet.
Cayucos, Lion Rock, Hazard Canyon, Shell Beach.
Vancouver Island to Coronados Islands.

Antiopella barbarensis (Cooper, 1863)

Frequent
Morro Bay Docks, Hazard Canyon.
Vancouver Island to Bahia San Quintin.

Aplysia californica Cooper, 1863

Common
Morro Bay mud flats.
Humboldt Bay to Gulf of California.

Page 426

THE VELIGER

Vol. 11; No. 4

Archidoris montereyensis (Cooper, 1862)

Frequent

Cayucos, Morro Bay Docks, Hazard Canyon, Lion Rock,
Shell Beach.

Alaska to San Diego.

Archidoris odhneri (MacFartanp, 1966)
[see Burn, 1968]

Rare Subtidal from 20 feet.
Avila, Shell Beach.
Monterey Bay to Santa Barbara.

Berthella californica (Dax, 1900)

Frequent
San Simeon, Cayucos, Shell Beach.
Crescent City to San Diego.

Bulla gouldiana Pitssry, 1893

Frequent
Morro Bay mud flats.
Morro Bay to Baja California.

Cadlina flavomaculata MacFartanp, 1905

Common
Cayucos, Hazard Canyon, Lion Rock, Avila, Shell Beach.
Vancouver Island to Point Eugenia, Mexico.

Cadlina luteomarginata MAcFarLaAnp, 1966

Common
Hazard Canyon, Lion Rock, Avila, Shell Beach.
Vancouver Island to Point Eugenia, Mexico.

Cadlina modesta MacFarLanp, 1966

Common
San Simeon, Cayucos, Hazard Canyon, Avila, Shell Beach.
Monterey Bay to La Jolla.

* Cadlina sparsa (ODHNER, 1921)

Rare

Morro Bay Docks, Hazard Canyon, Avila.

Morro Bay to Juan Fernandez Island, Chile. (Santa Bar-
bara to Juan Fernandez Island, Chile).

Capellinia rustya Marcus, 1961

Frequent
Morro Bay Docks, Hazard Canyon.
San Francisco Bay to Bahia de Los Angeles.

Catriona alpha (BaBA & Hamatant, 1963)
[see RoLueEr, this issue]
Rare
Morro Bay Docks.
Monterey Bay to San Diego; Japan.

Corambe pacifica MacFarLanp & O’DonocHuE, 1929

Common. On bryozoan colonies on offshore kelp.
Morro Bay Docks.
Vancouver Island to San Diego.

Coryphella trilineata O’DonocHuE, 1921

Common

Cayucos, Morro Bay Docks, Hazard Canyon, Lion Rock.
Avila, Shell Beach.

Vancouver Island to Coronados Islands.

Dendronotus albus MacFarLanp, 1966

Frequent
Hazard Canyon, Lion Rock, Avila, Shell Beach.
Monterey Bay to Santa Barbara.

Dendronotus frondosus (Ascantus, 1774)

Common
Morro Bay Docks, Hazard Canyon, Lion Rock, Shell Beach
Cosmopolitan in northern hemisphere.

Dendronotus subramosus MacFarLanp, 1966

Common
Hazard Canyon, Shell Beach.
Humboldt Bay to Newport Bay,

Diaulula sandiegensis (Cooper, 1862)

Common

San Simeon, Cayucos, Hazard Canyon, Lion Rock, Avila,
Shell Beach.

Japan to Cape San Lucas.

Dirona albolineata CocKERELL & Eviot, 1905
Rare

Shell Beach.
Puget Sound to San Diego,

Dirona picta MacFarLanp in CocKERELL & ELioT,
1905
Common (to 100 mm long)

Morro Bay Docks, Hazard Canyon, Lion Rock, Avila,
Shell Beach.

Dillon Beach to Point Loma.

Vol. 11; No. 4

THE VELIGER

Page 427

Discodoris heathi MacFartanp, 1905

Common
Cayucos, Hazard Canyon, Lion Rock, Avila, Shell Beach.
Vancouver Island to Laguna Beach.

Doridella steinbergae (LANCE, 1962) [see FRANZ, 1967]

Common. On bryozoan colonies on offshore kelp.
Morro Bay Docks.
San Juan Islands to Coronados Islands.

Doriopsilla ? albopunctata (Cooper, 1863)

Common

San Simeon, Cayucos, Hazard Canyon, Lion Rock, Avila,
Shell Beach.

Salt Point to Point Eugenia, Mexico.

Doto amyra ? Marcus, 1961

Common
Morro Bay Docks, Hazard Canyon.
Monterey Bay to Santa Barbara.

* Doto kya Marcus, 1961

~ Common

Cayucos, Morro Bay Docks, Hazard Canyon, Shell Beach.

Moss Beach to Shell Beach. (Moss Beach to Monterey
Bay).

Flabellinopsis iodinea (Cooper, 1862)

Frequent
Hazard Canyon, Avila (Port San Luis Wharf ).
Vancouver Island to Coronados Islands.

Haminoea vesicula (Goup, 1855)

Frequent
Morro Bay mud flats, Morro Bay Docks.
Alaska to Gulf of California.

Hancockia californica MAcFarRLanp, 1923

Rare
Hazard Canyon.
Dillon Beach to Punta Abreojos, Mexico.

Hermaeina smithi Marcus, 1961

Frequent
Cayucos, Hazard Canyon, Shell Beach.
San Juan Islands to San Diego.

Hermuissenda crassicornis (EscHscHOLTz, 1831)

Common

Cayucos, Morro Bay Docks, Morro Bay mud flats, Hazard
Canyon, Lion Rock, Avila, Shell Beach.

Sitka, Alaska to Punta Eugenia, Mexico; Puertecitos,
Bahia de Los Angeles.

Hopkinsia rosacea MacFarianp, 1905

Common
Cayucos, Hazard Canyon, Lion Rock, Avila, Shell Beach.
Coos Bay to Point Loma.

Laila cockerelli MacFarianp, 1905

Rare
Lion Rock, Shell Beach.
Vancouver Island to Cape San Lucas.

Melibe leonina (Goutp, 1853)
Rare. On entangled offshore kelp fronds.
Morro Bay Docks.
Alaska to La Paz Bay, Mexico; Bahia de Los Angeles.

Navanax inermis (Cooper, 1862)
Frequent
Morro Bay mud flats.
Monterey Bay to Laguna Manuela, Mexico.

Okenia angelensis LANCE, 1966

Rare

Morro Bay Docks.

San Francisco Bay to Bahia de Los Angeles.
Onchidoris hystricina (BERGH, 1878)

Common
Morro Bay Docks, Hazard Canyon, Avila, Shell Beach.
Alaska to Point Loma.
Onchidoris spec.
Rare
Hazard Canyon, Lion Rock.
Phidiana pugnax Lance, 1962

Common
Hazard Canyon, Avila, Shell Beach.
Monterey Bay to Coronados Islands

Phyllaplysia taylori (Dat, 1900)

Common. On Zostera marina.

Page 428

Morro Bay mud flats.
San Juan Islands to San Diego Bay.

* Pleurobranchus californicus denticulatus MACFaRLAND,

1966

Rare
Avila.

Monterey Bay to Avila. (Monterey Bay).

Polycera atra MacFaruanp, 1905

Common
Morro Bay Docks, Hazard Canyon, Shell Beach.
San Francisco Bay to Coronados Islands.

Polycera hedgpethi Marcus, 1964

Frequent
Morro Bay Docks.
San Francisco Bay to Bahia de Los Angeles.

Precuthona divae Marcus, 1961

Frequent
Hazard Canyon.
Monterey Bay to Santa Barbara.

Rostanga pulchra MacFaruanp, 1905

Common

San Simeon, Cayucos, Hazard Canyon, Lion Rock, Avila,
Shell Beach.

Vancouver Island to Chile; Gulf of California.

Spurilla olviae (MaAcFaruanp, 1966)

Frequent

San Simeon, Hazard Canyon, Lion Rock, Avila, Shell
Beach.

Monterey Bay to Santa Barbara.

Stiliger fuscovittata LANcE, 1962

Frequent
Morro Bay Docks, Shell Beach.
San Juan Islands to San Diego; Bahia de Los Angeles.

* Trapania velox (CocKERELL, 1901)

Rare
Hazard Canyon, Avila.
Hazard Canyon to San Diego. (Santa Barbara to San

Diego).

Trinchesia abronia (MacFar.anp, 1966)
[see Router, this issue]

Rare
Shell Beach.
Monterey Bay to Pismo Beach.

THE VELIGER

Vol. 11; No. 4

Trinchesia albocrusta (MacFartanp, 1966)
[see RoLter, this issue]
Frequent
Morro Bay Docks, Hazard Canyon.
Monterey Bay to Santa Barbara.

* Trinchesia flavovulta (MacFartanp, 1966)

[see ROLLER, this issue]
Rare
Shell Beach.

Monterey Bay to Shell Beach. (Monterey Bay).

Trinchesia lagunae (O’Donocuug, 1926)
[see RoLLER, this issue]
Frequent
Hazard Canyon, Lion Rock, Shell Beach.
Monterey Bay to Rosarito Beach, Baja California.

Triopha carpenteri (STEARNS, 1873)

Common
Hazard Canyon, Lion Rock, Shell Beach.
Vancouver Island to San Diego; Japan.

Triopha grandis MacFartanp, 1905

Frequent
Morro Bay Docks, Avila.
Santa Cruz to Catalina Island.

Triopha maculata MacFartanp, 1905

Common

Cayucos, Morro Bay Docks, Hazard Canyon, Lion Rock,
Avila, Shell Beach.

Bodega Bay to San Diego.

Triopha spec. [same as in SPHON & LANCE, 1968]

Common
San Simeon, Cayucos, Morro Bay Docks, Hazard Canyon,
Lion Rock, Avila, Shell Beach.

Tritonia exsulans BERGH, 1894

Rare. Subtidal from 180 feet.

Dredged from off Pismo Beach.

Vancouver Island to Baja California; Japan; Manatee
Bay, Florida.

Tritonia festiva (STEARNS, 1873)

Frequent
San Simeon, Cayucos, Lion Rock, Avila, Shell Beach.
Vancouver Island to Coronados Islands.

Vol. 11; No. 4

THE VELIGER

Page 429

LIST or INFERRED SPECIES

Acanthodoris brunnea MacFaruanp, 1905
Monterey Bay to Santa Barbara.
Aplysia vaccaria WINKLER, 1955
Morro Bay to Gulf of California.
Armina californica (CoopErR, 1862)
Vancouver Island to Panama
Atagema quadrimaculata Cour, 1963
Monterey Bay to San Diego.
Chromodoris macfarlandi (CocKERELL, 1902)
Monterey Bay to Coronados Islands.
Dendronotus iris Cooper, 1862
Vancouver Island to Coronados Islands.
Elysia hedgpethi Marcus, 1961
San Juan Island to La Jolla; Bahia de Los Angeles.
Fiona pinnata EscuscHo.tz, 1831
Pelagic.
Hypselodoris californiensis (BERcH, 1879)
Monterey Bay to Coronados Islands; Gulf of California.
Hypselodoris porterae (CocKERELL, 1902)
Monterey Bay to Cedros Island.
Phylliroe bucephala PERON & LESUEUR, 1810
Pelagic, worldwide.
Platydoris macfarlandi Hanna, 1951
Pismo Beach, subtidal at 516 feet.
Thordisa bimaculata LANcE, 1966
Carmel to Isla Natividad, Mexico.
Tochuina tetraquetra (Pattas, 1788)
Alaska to Santa Barbara.
Tylodina fungina Gass, 1865
Cayucos to Todos Santos, Mexico; Guaymas, Mexico.

LITERATURE CITED

ASCANIUS, PEDER
1774. Beskrivelse over en Norsk Sneppe oget Séedyr (Mol-
luscum Amphitrite frondosa) . K. Norske Vidensk. Selskr.
Skr. 5: 153-158; plt. 3; fig. 2. [not seen]
Basa, KikuTARO & Iwao HaMATANI
1963. A cuthonid, Cuthona alpha n. sp., with a radula of
Catriona type (Nudibranchia-Eolidoidea) . Publ. Seto
Mar. Biol. Labor. 11 (2): 339 - 343; plt. 11
Bercu, Lupwic SopHus RUDOLF
1878. | Malacologische Untersuchungen. In C. SEMPER,
Reisen im Archipel der Philippinen 2 (14): 603-645; plts.
66 - 68
1879. On the nudibranchiate gastropod mollusca of the north
Pacific ocean, with special reference to those of Alaska. Proc.
Acad. Nat. Sci. Philadelphia, pt. 1: 71-132; plts. 1-8.
1880. On the nudibranchiate gastropod Mollusca of the North
Pacific Ocean, with special reference to those of Alaska. II.
Proc. Acad. Nat. Sci. Philadelphia 1; 189 - 276

Bercy, Lupwic Sopuus Rupo.F
1894. | Die Opisthobranchien.
Harvard 25 (10) :
BRooKSHIRE, JACK

Bull. Mus. Comp. Zool,
125 - 233; plts, 1-12

1968a. Mollusca of the San Luis Obispo County area. Part I.
Gastropoda. Tabulata 1 (4): 5-6 (1 April 1968)
1968b. Mollusca of the San Luis Obispo County area, Part II.

Pelecypoda and Amphineura. Tabulata 1 (5): 5-6
(1 July 1968)
Burn, Ropert
1968. Archidoris odhneri (MacFarLanp, 1966) comb. nov.,
with some comments on the species of the genus on the

Pacific Coast of North America. The Veliger 11 (2):

90 - 92 (1 October 1968)
COockERELL, THEODORE Dru ALISON
1901. Three new nudibranchs from California. Journ.

Malacol. 8: 85 - 87
1902. Three new species of Chromodoris.
19-21
CockKERELL, THEODORE Dru ALISON & CHARLES ELIOT
1905. Notes on a collection of California nudibranchs.
Journ. Malacol, 12 (3): 31-53; plts. 7, 8
Coxutrr, Cuinton L. « WEsLEY M. FARMER
1964. Additions to the nudibranch fauna of the east Pacific
and the Gulf of California. Trans. San Diego Soc. Nat.
Hist, 13 (19): 377 - 396; plts. 1-6; figs. 1-3
Cooper, JAMES GRAHAM

Nautilus 16:

1862. Some genera and species of California Mollusca.
Proc. Calif. Acad. Nat. Sci. 2: 202 - 207

1863. | On new or rare mollusca inhabiting the coast of Cali-
fornia. Proc. Calif. Acad. Nat. Sci. 3 (2): 56-60

Dati, WiLtt1AM HEALEY
1900a. On a genus (Phyllaplysia) new to the Pacific coast.
Nautilus 14 (8): 91-92
1900b. A new species of Pleurobranchus from California.
Nautilus 14 (8) : 92 - 93

1919. Descriptions of new species of mollusca from the North
Pacific Ocean in the collection of the United States National

Museum. Proc. U.S. Nat. Mus. 56: 293 - 371
EscHSCHOLTZ, JOHANN FRIEDRICH
1829-1833. Zoologischer Atlas ... Berlin [not seen]
Farmer, Wes.tey M. « ALLAN JOHN SLOAN
1964. | A new opisthobranch mollusk from La Jolla, California.
The Veliger 6 (3): 148-150; plt. 18, figs.1,2 (1 Jan. 1964)

Goutp, Aucustus ADDISON
1852-1856. U.S. Exploring Expedition. 12: Mollusca and Shells.
xv+510 pp.; Atlas (1856): 52 plts. Boston [not seen]

LancE, JAMES ROBERT

1961. A distributional list of Southern California Opistho-
branchs. The Veliger 4 (2): 64-69 (1 October 1961)
1962.a. Two new opisthobranch mollusks from Southern Cali-
fornia. The Veliger 4 (3): 155 - 159; plt. 38; 8 text figs.
(1 January 1962)
1962b. A new Stiliger and a new Corambella (Mollusca :
Opisthobranchia) from the Northwestern Pacific. The
Veliger 5 (1): 33 - 38; plt. 6; 10 text figs. (1 July 1962)
1966. New distributional records of some northeastern Pacific
Opisthobranchiata (Mollusca:Gastropoda) with descriptions
of two new species. The Veliger 9(1): 69-81; figs.
1-12 (1 July 1966)

Page 430

THE VELIGER

Vol. 11; No. 4

LINNAEUS, CAROLUS
1761. Fauna Suecica sistens animalia Suecia regni. Edit.

altera. Stockholm [not seen]
Lone, STEVEN J.
1969. Two new records of Cratena abronia MacFaruanp,

1966. The Veliger 11 (3): 281
MacFaruanp, FRANK MAcE
1905. A preliminary account of the Dorididae of Monterey
Bay, California. Proc. Biol. Soc. Washington, 18: 35 - 54
1923. The morphology of the nudibranch genus Hancockia.
Journ. Morphol. 38 (1): 65 - 104; plts. 1-6
1925 - 1926. The Acanthodorididae of the California coast.
Nautilus 39 (2): 49-65; 39 (3): 94-103; plts. 2, 3
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
MacFaruanpb, Frank Mace & CHarLes Henry O’DoNocHUE
1929. A new species of Corambe from the Pacific Coast of
North America. Proc. Calif. Acad. Sci. ser. 4, 18 (1):
1-27; plts. 1-3
Marcus, Ernst
1961. | Opisthobranch mollusks from California. The
Veliger 3 (Supplement, pt. I): 1-85; plts. 1-10. (Feb. 1, 1961)
1964. A new species of Polycera (Nudibranchia) from Cali-
fornia. The Nautilus 77 (4): 128 - 131; figs. 1-4
Opuner, Nits HjaLmar
1921. Mollusca of Juan Fernandez and Easter Island. Jn:
GC. Skotrsserc, The natural history of Juan Fernandez and
Easter Island. 3 (2): 219-254; plts. 8, 9

(1 January 1969)

WINKLER, LINDSAY ROBERT

O’DonocHuE, CHartes H.
1921. Nudibranchiate mollusca from the Vancouver Island
region. ‘Trans. Roy. Canad. Inst. 13 (1): 147-209; plts. 7-11
1926. A list of the nudibranchiate mollusca recorded from
the Pacific coast of North America, with notes on their
distribution. ‘Trans. Roy. Canad. Inst. 15 (2): 199-247.
Router, Ricuarp A.
1969a. A color variation of Aldisa sanguinea (Coorzr, 1862).
The Veliger 11 (3): 280-281; 1 fig. (1 January 1969)
1969b. Nomenclatural changes for the new species assigned to
Cratena by MacFartanp (1966). The Veliger 11 (4):
(this issue)
SpPHON, GaLeE G., Jr. & James Rosert LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif. Acad.
Sci. 36 (3): 73-84; 1 fig. (25 September 1968)
STEARNS, Rosert Epwarps CarTER
1873. Description of a new genus and two new species of
nudibranchiate mollusks from the coast of California.
Proc. Calif: Acad. Sci. 5 (1): 77-78
STEINBERG, JOAN EMILY
1963a. Notes on the opisthobranchs of the West Coast of
North America. - III. Further nomenclatorial changes in
the order Nudibranchia. The Veliger 6 (2): 63 - 67
(1 October 1963)
1963b. Notes on the opisthobranchs of the west coast of North
America. — IV. A distributional list of opisthobranchs from
Point Conception to Vancouver Island. The Veliger 6 (2) :
68 - 73 (1 October 1963)

1954. A new species of Aplysia on the southern California
coast. Bull. South. Calif: Acad. Sci. 54 (1): 5-7; plts. 2, 3

Vol. 11; No. 4

THE VELIGER

Page 431

A Note on the Range of Gastropteron pacificum

HANS BERTSCH

Santa Barbara Museum of Natural History

Santa Barbara, California '

(1 Map)

INTRODUCTION

Gastropteron pacificum BErcu, 1894, belongs to the order

Cephalaspidea, a large opisthobranch group comprising
14 families and more than 150 genera and subgenera
(Taytor & SoHL, 1962). Cephalaspideans are character-
ized by a cephalic shield, an expanded thickening of the
dorsal surface of the head. This cephalic shield may
possibly represent fused tentacle bases; in the cephalaspi-
deans the tentacles are mostly absent, although the corners
_ of the shield may be elongated into tentacle-like projec-
tions (Hyman, 1967, p. 407). Gastropteron can form a
temporary siphon by rolling the freely movable posterior
margin of the cephalic shield into a tube (BERGH,
1894, p. 201).

When actively crawling, the general body form of the
family Gastropteridae (containing the genera Gastropter-
on and Sagaminopteron) is an elongated cylindrical shape
with paired lateral parapodia (continuous with the foot)
turning up over the back, and with the prominent tail
tapering back. By flapping the expanded parapodia the
animal frequently swims through the water (Basa & ToKI-
oKA, 1965, p. 375, and HAEFELFINGER & Kress, 1967).
The radular formula of Gastropteron is 4-6:1:0-1-4-6
(BaBa & ToKioKA, loc. cit.).

Detailed descriptions and illustrations of the egg masses
and veligers of Gastropteron pacificum are given by
Hurst (1967). The adult shell (illustrated in KEEN,
1963, p. 17) is reduced, almost vestigial, and completely
enclosed within the mantle. The ground color of the adult
is a pale yellow ochre, mottled with clusters of small red
spots. A thorough description of the adult anatomy is
given by MacFartanp, (1966, pp. 2 - 6; pit. 1).

ToxkiokKA & Basa (1964) summarize the known ranges
and main references of the 5 described species of Gastro-

™ Permanent address: Franciscan School of Theology, 1712
Euclid Avenue, Berkeley, California 94709

pteron prior to their establishment of 4 new species from
Japan.

GEOGRAPHICAL DISTRIBUTION

The published range of distribution of Gastropteron

pacificum consists of a rather scattered series of localities
along the Pacific coast of western North America. Along
most of the coast it is listed as a presumed offshore species,
especially in the approximately 350 miles of coast between
Monterey Bay and Newport Bay.
An offshore, pelagic species, it is not encountered by the
tidepool collector of opisthobranchs. The relative scar-
city of G. pacificum is attested to by the majority of
negative responses the author received upon inquiries
to collectors asking if they had ever caught this species
or if they knew of any specimens in museum or private
collections. The present note correlates previously pub-
lished accounts of the range of distribution of G. paczifi-
cum with more recent data obtained by James Lance and
by the Santa Barbara Museum of Natural History from
the collecting of Pat Brophy and Shane Anderson, sub-
stantiating its occurrence along the southern California
coast.

With one exception, the literature prior to the publica-
tion of MacFar.Lanp’s (1966) monograph on the opistho-
branchs of the Pacific coast of North America gives the
range of Gastropteron pacificum as extending from the
Aleutian Islands, Alaska, to the Fuca Strait (south of
Vancouver Island, British Columbia, Canada): Bercu,
1894; Dax, 1921; O_pRoyp, 1927; Kren, 1937; BurcH,
1945; La Rocgue, 1953; Appott, 1954. SmirH & Gor-
pon (1948), however, mention the occurrence of this
cephalaspidean in Monterey Bay (dredged, rare).

More recently, STEINBERG (1963) indicates Gastropter-
on pacificum along with numerous other cephalaspideans,
as occurring between San Diego and Vancouver Island,

Page 432

Figure 1

Western Coastline of California

The localities are, from north to south: off San Francisco,
Monterey Bay, Gaviota, Santa Barbara, Ventura, Newport
Bay, and Point Loma.

but does not include specific localities. MAcFARLAND
(1966) records this species from the following localities:
off San Francisco; in Monterey Bay; Newport Bay; and
along the west coast of Central America from the Gulf
of California to the Galapagos Islands. Specimens collec-
ted in the vicinity of Friday Harbor, Washington, were
used by Hurst (1967) in her detailed study of opistho-
branch veligers. SpHON & Lance (1968) do not discuss
the occurrence of any cephalaspideans in their checklist
for Santa Barbara County.

Gaps in the range of Gastropteron pacificum occur
between Fuca Strait and San Francisco, between Monte-
rey and Newport Bays, and between Newport Bay and
the Gulf of California. The following collecting data
indicate the presence of G. pacificum in the latter two
regions:

1. One specimen; in an otter trawl off Point Loma (San
Diego), California, at approximately 100 fathoms. Jan-
uary 28, 1965. Collected by E. W. Fager of Scripps
Institution of Oceanography and party [in the collection
of James R. Lance].

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Vol. 11; No. 4

2. One specimen; backwash of the filter system of the
Undersea Gardens, Santa Barbara, California. Janu-
ary 24, 1968; collected by Shane Anderson, identified by
Richard Lee (deposited in the collection of SBMNH,
no. 25083).

3. Three specimens; Ventura, California, 85 fm. March
27, 1968. Collected by Pat Brophy, identified by Mrs.
Jacque Rogers (one specimen in the collection of SBM
NH no. 25084).

[The disposition of the remaining specimens, all col-
lected by Pat Brophy, is unknown; data obtained
through personal communication, J. R. Lance and
P. Brophy. |

4. One specimen; Rincon, California (between Ventura
and Santa Barbara), 50 fm. May 7, 1968.

5. One specimen; Santa Barbara, 105 fm. June 24, 1968.

6. One specimen; Gaviota, California, 106 fm. July 6,
1968.

7. Two specimens; Gaviota, California, 119 fm. July 29,
1968.

8. One specimen; Gaviota, California, 117 fm. July 30,
1968.

9. Two specimens; Santa Barbara, California, 130 fm.
August 15, 1968. _

10. Fourteen specimens; Santa Barbara, California, 12
miles offshore, 130 fm. August 16, 1968.

11. Ten specimens; Santa Barbara, California, 12 miles
offshore, 123 fm. August 16, 1968.

12. Three specimens; Santa Barbara, California, 12 miles
offshore, 134 fm. September 1, 1968.

The accompanying map indicates the subtidal localities
along the California coast from which Gastropteron pacif-
zcum has been collected.

With increased offshore collecting, specimens of Gastro-
pteron pacificum should be found throughout its entire
range and, hopefully, more information will be gathered
about the life history and distribution of this opistho-
branch.

ACKNOWLEDGMENTS

Sincere thanks are extended to Pat Brophy and Shane
Anderson for making available the material they col-
lected; to James R. Lance for the San Diego locality
record and his generous assistance during the preparation
of this note; to Nelson Baker and Richard S. Lee of the
Santa Barbara Museum of Natural History for allowing
me to use the material in the SBMNH;; and to the many
persons who responded to my inquiry.

Vol. 11; No. 4

THE VELIGER

Page 433

LITERATURE CITED

Asgott, Ropert TUCKER
1954. American seashells. Princeton, New. Jersey. D. van
Nostrand Co., Inc.; xiv+541 pp.; 100 figs.; 40 plts.
Basa, KikuTarO & TAKASA TOKIOKA
1965. Two more new species of Gastropteron from Japan,
with further notes on G. flavum T « B (Gastropoda: Opistho-
branchia). Publ. Seto Mar. Biol. Lab. 12 (5): 363 - 378;
pit. 25; 8 text figs.
Bercu, Lupwic SopHus RupoLeH
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 “Alba-
tross”, during 1891. XIII. Die Opisthobranchien. Bull. Mus.
Comp. Zool., Harvard Univ. 25 (10): 125 - 233; plts. 1-12
(October 1894)
Burcu, JoHN Quincy (ed.)
1945. Minutes of the Conchological Club of Southern Cali-
fornia no. 48: 6
Dai, Wituiam HEALEY
1921. | Summary of the marine shellbearing mollusks of the
northwest coast of America, from San Diego, California, to
the Polar Sea.... Bull. U. S. Nat. Mus. 112: 1 - 217; plts.
1-22 (24 February 1921)
HAEFELFINGER, Hans-Rupo.tr « A. Kress
1967. Der Schwimmvorgang bei Gastropteron rubrum (Ra-
FINESQUE, 1814) (Gastropoda, Opisthobranchiata). Rev.
Suisse Zool. 74 (3): 547 - 554; 4 figs. (December 1967)
Hurst, ANNE
1967. The egg masses and veligers of thirty Northeast Pacific
opisthobranchs. The Veliger 9 (3) : 255 - 288; plts. 26 - 38;
31 text figs. (1 January 1967)
Hyman, Lissie H.
1967. ‘The invertebrates: Mollusca I.
New York; viit792 pp.; 249 figs.

McGraw Book Co.,

Keen, A. Myra

1937. An abridged check list and bibliography of West North
American marine mollusca. Stanford Univ. Press, Stanford,
Calif. pp. 1-88

1963. | Marine molluscan genera of western North America:
An illustrated key. Stanford Univ. Press; 126 pp.; text
figs. not separately numbered (14 February 1963)

La Rocgur, AURELE

1953. Catalogue of Recent Mollusca of Canada. Nat.

Mus. Canada Bull. No. 129: ix+406 pp.
MacFarianp, Frank Mace

1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)

O.proyp, IpaA SHEPARD

1927. The marine shells of the west coast of North America.

2 (1): 297 pp.; 22 plts. Stanford Univ. Press, Stanford, Calif.
Smit, Attyn G., & MacKenziz Gorpon, Jr.

1948. The marine mollusks and brachiopods of Monterey
Bay, California, and vicinity. Proc. Calif. Acad. Sci.,
4th. Ser., 26 (8): 147-245; plts. 3-4; textfigs. 1-4

Spoon, Gate G., Jr. « James Ropert LAaNcE

1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif. Acad.
Sci. 36 (3): 73 - 84; 1 fig. (25 September 1968)

STEINBERG, JoAN EMILY

1963. Notes on the opisthobranchs of the west coast of North
America. II. The order Cephalaspidea from San Diego to
Vancouver Island. The Veliger 5 (3): 114-117

(1 January 1963)
Taytor, Dwight WILLARD & Norman F Sout

1962. An outline of gastropod classification.

gia 1 (1): 7-32
Toxrioka, TaKasa & K1KUTARO BaBA

1964. Four new species and a new genus of the family Gastro-
pterida from Japan (Gastropoda: Opisthobranchia) . Publ.
Seto Mar. Biol. Lab. 12 (3): 201 - 229; plts. 10-13; 15 text
figs.

Malacolo-

Page 434

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Vol. 11; No. 4

An Immunological Study of Pelecypod Taxonomy

LARKLYN FISHER

Department of Zoology, University of Illinois, Urbana, Illinois 61801"

(3 Text figures)

INTRODUCTION

ANIMAL RELATIONSHIPS CAN BE Studied in many different
ways. Organisms are usually classified on the basis of
similarities and differences in morphology, in physiology, in
development, and in ancestry as shown by fossil remains.
Another way that could be used to study relationship
would be to consider the amount of difference in the
types of substances present in the tissues, such as the pro-
teins present in the muscles. This method was used in this
experiment, in which muscle tissue from several species of
clams was compared by immunological means.

Immunological methods have been used in the past to
study taxonomy of vertebrates. The first extensive work
of this sort was done by Nutrati (1904) who showed
that when blood sera from many species, including man,
the great apes and the domestic cat and dog were used
as antigens and tested with rabbit antisera, the reactions
were greatest when the species from which the blood
serum was taken was more closely related to the species
from which the original antigen had been obtained for
injection into the rabbit. In NuTTALL’s study, increasing
strength of the antibody-antigen reaction was found to
follow a closer zoological relationship, and thus agree with
standard taxonomy.

More recently, serological methods have been used in
both plant and animal taxonomy, as in the work of Fair-
BROTHERS & JOHNSON (1962) with systemics of the fam-
ilies Cornaceae (Dogwood) and Nyssaceae (Sour Gum) ;
GoopMan’s (1962) immunological study of primate tax-
onomy; the serological comparison of 7 species of birds
by DuweE (1962) ; the gel diffusion tests of ungulate tax-
onomy by MaraBLE & GLENN (1962) ; and the investiga-
tion of turtle family relationships by Framr (1962).

The use of immunological techniques to study taxonom-
ic relationships is based on the assumption that just as
closely related clam species have more similarities of

™ Present address: 5421 6o' Str. N. E., Seattle, Washington 98105

gross morphology than do less closely related species, so
do they also have more similarities in their tissue proteins.
If a tissue, such as muscle, is ground up to break the cell
walls and free the proteins, it can then be used as an anti-
gen, that is, a substance to which antibodies can be
produced. When this antigen is injected into a rabbit,
the rabbit will make antibodies to the foreign proteins and
these antibodies will appear in its blood serum. If this anti-
serum is mixed with the original antigen, the antibody
and antigen molecules will attach to one another in
great numbers and form a lattice, which is more dense
than the surrounding fluid and will form a precipitate.
This reaction between the antibodies and the proteins
which make up their homologous antigen is quite spe-
cific’. Thus, if there was a precipitate between antigen
from the muscle of one clam and antiserum made against
the muscle tissue of another clam, it would show that the
two had at least one protein in common. If two such lines
of precipitate met and merged, it would show that they
were reactions to a single protein, present in both subjects.
If they did not merge, however, this would demonstrate
that while muscle tissue from each species did have a pro-
tein in common with the original antigen to which the
antiserum was made, it was not the same protein; there-
fore each of these two species of clams had one protein
in its muscle that the other one did not have, thereby
indicating a more distant taxonomic relationship. These
were the principles that were used in the present experi-
ment, to examine the amount of relationship shown
among several different species of clams.

METHODS

Seven species of clams were used in this experiment, 6 of
which occur in the fresh water streams of Illinois, and
one that is marine, from the coast of the State of Wash-
ington. The fresh water species were:

2 see Boyp, 1966, pp. 370 - 371

Vol. 11; No. 4

SPHAERIDAE
Sphaerium striatinum (Lamarck, 1818)
UNIONDAE
Unioninae
Quadrula pustulosa (LEA, 1834)
Anodontinae
Anodonta grandis Say, 1829
Lasmigona complanata (BARNES, 1823)
Lampsilinae
Lampsilis ventricosa (BARNES, 1823)
Actinonais carinata (BaRNES, 1823)
The marine clam was:
‘VENERIDAE
Saxidomus nuttalli Conrav, 1837.

The antigen to be injected was obtained from 3 species,
Lampsilis ventricosa, Quadrula pustulosa, and Lasmigona
complanata. The retractor and adductor muscles were
removed, cut into fine pieces, extracted in saline solution,
frozen and thawed several times and the same samples
were then ground in a hand grinder. The protein con-
centrations of the antigen preparations were determined
by the biuret method, using a colorimeter. Each antigen
was injected several times (see antigen schedule below)
into 2 rabbits. Injections were made intramuscularly,
with the dose divided into 4 injections, one in the prox-
imal part of each limb. At the time of the injections, 15
to 20 cc were bled from an ear of each rabbit, the clot
removed and the serum centrifuged to prevent hemolysis.
Pre-immunization (control) bleedings were taken from
each rabbit before the first injection. The schedule of
bleedings, antigen injections and their concentrations is
given below.

ANTIGEN SCHEDULE

Antigens from Quadrula pustulosa:
1. Antigen concentration — 1.68 mg/ml protein
Injected — day one: 10 ml antigen for 16.8 mg protein
and 5ml Freund’s complete adjuvant, thoroughly
mixed, into rabbits one and two.
2. Antigen concentration — 0.67 mg/ml protein
Injected — day nine: 7.5 ml antigen for 5 mg protein
and 5 ml Freund’s complete adjuvant into the same
2 rabbits.
3. Antigen concentration — 0.60 mg/ml protein
Injected — day eighteen: 14 ml antigen for 8.4 mg
protein and 6 ml Freund’s complete adjuvant into
rabbit number one only.
Antigens from Lasmigona complanata:
1. Antigen concentration — 1.0 mg/ml protein
Injected — day three: 10 ml antigen for 10 mg protein

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Page 435

and 10 ml Freund’s complete adjuvant into rabbits
three and four.

This was the only injection of Lasmigona complanata
material as no more specimens of this species could be
obtained.

Antigens from Lampsilis ventricosa:
1. Antigen concentration — 1.68 mg/ml protein
Injected — day one: 11.5 ml for 18.3 mg protein and
5.9 ml Freund’s complete adjuvant into rabbits
five and six.
2. Antigen concentration — 0.4 mg/ml protein
Injected — day nine: 7 ml for 2.8 mg protein each into
the same two rabbits.
The rabbits were bled on days one, nine, eighteen, and
thirty-three even when they were not injected.
Diffusion plates were prepared by pouring a 1%
solution of agar agar no. 3 (Oxoid Co.) onto 16 plates,
and cutting wells $ inch in diameter and 3 inch apart in
substrate $ inch deep, following the Ouchterlony proce-
dure. Antigens and antisera were placed undiluted in the
wells of the plates shown in the diagrams below. The
results with the first group of plates (Text figure 1)
indicated that dilution of the antisera would improve the
readings. Therefore each antiserum was diluted 1:1; 1:4;
1:8; 1:16; 1:32; and 1:64 and tested against its homo-
logous antigen, except for the antisera to Lasmigona com-
planata as no more antigen from this species was available.
From the results of these tests, the antisera were diluted
1:6 in the second group of plates, except that from rabbit
number six, which was diluted 1:1. All of the plates were
observed after 1, 3 and 5 days of refrigeration and the
results were recorded.

RESULTS

The patterns of precipitation on the plates are shown
on the diagrams below. These are the patterns on the fifth
day, when the plates seem to have attained their full
development.

Preimmunization (control) bleedings: the absence of
precipitation indicated that the rabbits had no antibodies
to the clam proteins at the beginning of the experiment.

DISCUSSION anp CONCLUSIONS

The results indicated that each of the clam species used
in this experiment was more closely related to some of the
remaining 6 than to others. These relationships agreed in
the main with conventional taxonomy but showed that
more distantly related groups may have a good deal more
in common than one might expect. The amount the spe-

Page 436

Plate 1 Plate 2
a/s to Quadrula pustulosa
from rabbit number one

Plate 4 Plate 5
a/s to Lampsilis ventricosa
from rabbit number six

THE VELIGER

a/s to Quadrula pustulosa
from rabbit number two

a/s to Lasmigona complanata
from rabbit number four

Vol. 11; No. 4

Plate 3
a/s to Lampsilis ventricosa
from rabbit number five

Plate 6
a/s to Lasmigona complanata
from rabbit number three

Figure 1

cies share is amplified by the fact that these animals are
not closely related to rabbits. According to LANDSTEINER
(1936) antibody formation is directed against those fea-
tures that are foreign to the animal forming the antibody;
thus much of a rabbit’s antibody forming capacity would
be directed against the major structural features not oc-
curring in rabbits, but common to all clam proteins. This
would also mean that only part of the antibodies would
distinguish the relatively minor species differences among
the clam proteins. The diagram (Figure 3) shows the simi-
larities of muscle proteins that were found in this ex-
periment. Each line represents one positive reaction, and
if 2 or more antibody-antigen systems precipitated to-

gether, it would represent more than one substance in
common in their muscle tissues. It must be noted, how-
ever, that the gel diffusion tests would not have detected
differences or similarities among clam proteins that are
also present in rabbits, as the rabbits would not have
recognized them as foreign and so would not have pro-
duced antibodies against them.

Each antigen could have shown reaction 5 or 6 times,
had all reactions been positive. The number of possible
positive reactions for each species, the number of those
that were prevented because of precipitation too close to
the well of antigen and the number of reactions that
were actually positive, are shown in Table 1.

==

Vol. 11; No. 4 THE VELIGER Page 437

Plate 1 Plate 2 Plate 3
a/s to Lampsilis ventricosa a/s to Lampsilis ventricosa a/s to Lampsilis ventricosa
from rabbit number six from rabbit number six from rabbit number six

Plate 4 Plate 5

a/s to Lasmigona complanata a/s to Lampsilis ventricosa

from rabbit number four from rabbit number five
Figure 2

Quadrula pustulosa

(Cine a) (a reaction of non-identity
(subf. Unioninae)
Saxidomus nuttalli Uj was also shown here)

(Veneridae) Lam psilis ventricosa

(Unionidae)
(subf. Lampsilinae)

Anodonta grandis
(Unionidae)
(subf. Anodontinae)

\\ //

Actinonats carinata Lasmigona carinata
(Unionidae) = ——— = _ (Unionidae)
(subf. Lampsilinae) (subf. Anodontinae)

Sphaerium striatinum
(Sphaeriidae)

Figure 3

Similarities Shown Among Muscle Proteins

Page 438 THE VELIGER Vol. 11; No. 4
Table 1 coast. The results showed 2 things: bivalves more distant-
1 ly related to one another have a good deal of similarity
Percentage of Positive Tests in their muscle proteins, and that classification of pele-
cypods by common factors in muscle proteins agrees sub-
# stantially with that based on morphology, physiology and
¥, Re Une development.
So £6 62 & o
Species ce eel FE: 3 2B By
nA a Oo Sn Ba A N
ee gs Bs go os CKNOWLEDGMENT
L ili tri 100 6 0 6 6 ‘
ines ine. 6 0 6 6 I would like to thank Dr. Nelda E. Alger for her con-
Quadrula pustulosa 100% 6 0 6 6 structive criticism of the manuscript.
Actinonais carinata 80% 6 1 5 4
Saxidomus nuttalli 80% 6 1 9) 4
Anodonta grandis 67% 6 0 6 4 LITERATURE CITED
Sphaerium striatinum 67% 6 0 6 4

The species at the top of this table showed a positive re-
action at least once with each species tested against them,
while those in the lower part of the table showed less and
less relationship with the other species.

The results of this experiment might have shown more
if several other species of bivalves could also have been
used, and if the antisera had been diluted for the first
group before being placed in the wells so that precipita-
tion would have taken place farther from the wells of
antigen. However, the results showed two things: first,
that groups considered more distantly related do have a
good deal of similarity in their muscle proteins, and,
secondly, that classification by common factors in muscle
proteins agrees substantially with the classification based
on morphology, physiology and development.

SUMMARY

The amount of relationship among 7 species of bivalves
was compared by immunological methods, using their
muscle tissue as the antigen and testing for similarity of
muscle proteins of the different species using the Ouchter-
lony technique. Six of the species were from fresh water,
representing 2 families and within one of these 3 sub-
families. The seventh species was marine from the Pacific

Boyp, WILLIAM C.

1966. Fundamentals of immunology, 4° ed., John Wiley &

Sons, Inc., New York, N. Y.; i-vii+773 pp.; 107 figs.
Duwe, ArTHuR E.

1962. An immunological comparison of seven species of birds
from four orders based on skeletal muscle antigens. In C. A.
Leone, Taxonomic biochemistry and serology. The Ronald Press
Co., New York, N. Y.; 4 pp.; 1 fig.

FarrBroTHERS, Davip E. & Marion A. JOHNSON

1962. | Comparative serological studies with the families Corna-
ceae (dogwood) and Nyssaceae (sour gum). Jn C. A. LEONE,
Taxonomic biochemistry and serology. The Ronald Press Co.,
New York, N. Y.; 16 pp.; 3 figs.

Frair, WAYNE

1962. Turtle family relationships as determined by serological
tests. In C. A. LEONE, Taxonomic biochemistry and serology,
The Ronald Press Co., New York, N. Y.; 10 pp.; 1 fig.

GoopMan, Morris

1962. Problems of primate systematics attacked by the sero-
logical study of proteins. In C. A. LEonE, Taxonomic biochem-
istry and serology. The Ronald Press Co., New York, N.Y.
20 pp.; 12 figs.

LANDSTEINER, K.

1936. The specificity of serological reactions.

mas, Springfield, Illinois
MaraB_eE, Iowa A. & W. G. GLENN

1962. Quantitative serological correspondence of ungulates by
gel diffusion. In C. A. LEonE, Taxonomic biochemistry and
serology. The Ronald Press Co., New York, N.Y.; 8 pp.; 3 figs.

NutTatLt, Grorcrk HENRY FALKINER

1904. Blood immunity and blood relationships. Cambridge

Univ. Press, Oxford; 444 pp.; 1 plt.; 4 figs.

C. C. Tho-

Vol. 11; No. 4

NOTES & NEWS

An Overlooked Subgenus and Species

from Panama

BY

A. MYRA KEEN

Department of Geology, Stanford University
Stanford, California 94305

Trichotropis gouldii A. Apams, 1857 was described in the
Proceedings of the Zoological Society of London for
1856 (p. 389, published January 10, 1857). The Latin
description, freely translated, is as follows:

Shell ovate-fusiform, very slightly umbilicate,
white, thin; spire high; with seven convex whorls
having raised lirae, the spire beautifully cancellate
and the interspaces striate with thin longitudinal la-
mellae; aperture oval, produced anteriorly, canal ob-
solete; outer lip smooth, rounded, anteriorly slightly
reflected, inner lip with simple sharp edge.

Length, 14 poll. [37 mm]. Hab. Chiriqui, Veragua
[W. Panama].

ApaMs surmised that the shell might have had a peri-
ostracum when alive, and he noted that “it differs from
the typical genus in the canal of the aperture being
almost obsolete.”

In 1861 Henry Apams (Proc. Zool. Soc. London, 1861,
p. 272) proposed the subgenus Alora, with Trichotropis
gouldi the sole species. Since then it has dropped from
notice.

Because the shell Apams described had been in the
Cuming collection, I searched for it in the collection of
the British Museum (Natural History) when I was there
in 1964 and 1965 and again in 1967 but without success.
If the shell is still in the Museum’s collection, it has
been mislabelled and the original notations obscured or
lost.

Leafing through the illustrations of known Panamic
gastropods for a possible identification, I can come up
with only one form from Panama that meets all the
requirements of Apams’ original description. This is
Recluzia insignis Pitspry & Lowe, 1932. The major dif-
ference is in size, for the Pirspry « Lowe holotype is
stated to be only 15 mm high. They even mention the thin

THE VELIGER

Page 439

periostracum that was predicted by Apams. Therefore,
I suspect that the Pmspry & Lowe species is going to
prove to be a synonym, but even if it can be retained, it
should be transferred from Recluzia, a genus in which
the shells are normally smooth, to Alora. Also, Alora
should be divorced from Trichotropis (a North Pacific
genus) and elevated to generic rank, with A. gouldii (A.
ApaMS, 1857) as type by monotypy. Sculpture and form
of the shell would suggest removal of the genus from
Janthinidae and transfer to Epitoniidae, where it might
find a place somewhere near Scalina.

Laternula Living on the Pacific Coast?

BY

A. MYRA KEEN

Department of Geology, Stanford University
Stanford, California 94305

In 1963 Miss CaTHERINE DuNLop, who was identifying
mollusks for an ecologic project being undertaken by
Portland State College students under Dr. John MacNab,
sent several small clams for determination. One lot proved
to be Laternula, a genus not native to the West Coast. The
specimens had been taken in Coos Bay, Oregon, about
150 feet from shore. There were two specimens, both
somewhat broken but with valves intact, one clearly show-
ing a lithodesma on the hinge.
Mr. Keith B. Macdonald, working in marsh channels
of Coos Bay in 1966, submitted two more specimens of
Laternula for identification. These also seem to have been
alive when taken, though the soft parts were missing by
the time I saw them. The periostracum was fresh, and one
valve showed a fragment of the lithodesma. These speci-
mens were slightly larger than the earlier two but smaller
than most adult Laternula. Comparison of the two lots
and study of the literature on Japanese Laternula (the
most likely source of importation), suggests that the
species probably is Laternula (Laternula) limicola (REEVE,
1863), a form that occurs on mud flats in central and
northern Japan. The fact that two lots were taken in
Oregon three years apart would imply that this Japanese
bivalve may be becoming established in American waters.
(Order Anomalodesmata; Superfamily Pandoracea).

Page 440

THE VELIGER

Vol. 11; No. 4

Donation by San Diego Shell Club

The Veliger Operating Fund has received another gener-
ous contribution from the San Diego Shell Club.

These donations are used, in accordance with the wishes
of the donors, to assist in paying for plates of authors who
are not in a position to carry the full burden themselves,
either because of foreign exchange restrictions by their
authorities or because of lack of access to the needed
funds. While it is the aim of The Veliger eventually to
absorb all expenses of publishing the journal, the financial
position of the California Malacozoological Society is
such that this goal is not yet achieved - in fact, far from it.
It is, therefore, an especially welcome event and we, on
our own behalf as well as that of some author(s) who
will benefit from it, express our sincere appreciation of
this continued support.

The Editor.

Important Notices

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. We wish to take this opportunity
to remind our Members that a reinstatement fee of
one dollar becomes due if membership renewals have
not been received by C. M.S., Inc. by April 15, 1969.

Sexual Dimorphism in Tegula funebralis

BY

PETER W. FRANK

Department of Biology, University of Oregon
Eugene, Oregon 97403

Durine A sTupy of behavioral and distributional differ-
ences between the sexes of Tegula funebralis (A. ADAMS,
1855), a method allowing the animals to be sexed without
killing them has been discovered. The crawling surface
of the foot is cream- to light brown-colored. The chemi-
cal basis of the pigmentation is unknown; the color sug-
gests a carotenoid. If one separates the lightest from the

darkest individuals, those with the lightest undersides are
virtually consistently males, the dark ones females (see
Table 1). Since there are a number of intermediate indi-
viduals, approximately the last quarter of any given
sample can not be separated into sexes by this means.
Moreover, the difference in pigmentation does not show
up in snails less than about 1.5cm wide, i.e., several
millimeters larger than the size at which sex may be
distinguished from the color of the gonad, once the shell
is broken.

This research is supported by grant GB 5032 of the
National Science Foundation.

‘Table 1

Number of Tegula funebralis with dark and light crawling
surfaces compared with respect to sex. Data obtained in
August 1968 from animals collected at
Cape Arago, Oregon

Surface of
Foot Sample 1 Sample 2
é 2 é g
Dark 4 48 3 19
Light 27 6 20 0
Intermediate 16 9 5 4

Spawning Notes, III. - Strombina maculosa

BY

FAY H. WOLFSON

San Diego Natural History Museum,
g San Diego, California 92115

(3 Text figures)

CoMMUNAL SPAWNING by Strombina maculosa (SowER-
BY, 1832) was observed by Glenn and Martha Vargas at
Bahia de los Angeles, Baja California during the extreme
low tides of 26 and 27 June 1968. Large numbers of
individuals, in aggregations, were depositing egg capsules
on small stones, in mussel valves and on other empty
shells. An appreciably smaller number was observed
spawning two weeks later during the lowest tides of the
next spring-tide cycle.

The capsules are deposited in rows (Figure 1) ; on the
stone collected there are 2 rows of 16 and a third of 12.

Vol. 11; No. 4

THE VELIGER

Page 441

Figure 1

Strombina maculosa (SowERBy, 1832)
Egg capsules on stone

The capsules are separated by a distance of at least 2mm
except at the base, where the flaring membranes at-
taching them to the substrate are contiguous, but not
connected.

Viewed microscopically, the puckered and opaque ap-
pearance of the capsules is seen to result from an intricate
pattern of irregularly shaped “facets” into which the
entire transparent surface is divided.

The capsules average 9.1 mm in maximum height, 7.2
mm in maximum width. Average width of the top ridge
(Figure 3) is 0.93 mm, of the side ridges, 0.69 mm. There
is no sign of an inner membrane or of a special structure
to serve as exit at hatching time. The few veligers (less
than 100 per capsule) are scattered irregularly through-

Figure 2

Strombina maculosa (SoweErRBy, 1832)
A single capsule

out (Figure 2). They average 0.187 mm in greatest diam-
eter.

Figure 3

Strombina maculosa (SoweErsy, 1832)
Detail of top of capsule, with a few of the “facets” of the top
and side ridges shown

Spawning Notes, IV.

Cerithium stercusmuscarum

BY

FAY H. WOLFSON

San Diego Natural History Museum,
San Diego, California 92115

(2 Text figures)

THE SPAWN OF Cerithium stercusmuscarum VALENCI-
ENNES, 1833 is comparatively simple. It resembles that of
the Bermuda species, C. ferrugineum Say, in that spawn-
ing occurs near the high tide line and the eggs are encased
in a gelatinous matrix within a thin-walled tube, but
differs in the form assumed by the spawn: that of C.
ferrugineum resulting in a loose coil (Lesour, 1944),
that of C. stercusmuscarum being laid down in loops
(Figure 1).

Several hundred Cerithium stercusmuscarum were con-
gregated at the edge of a tidal lagoon at La Gringa, Bahia
de los Angeles, Baja California on 23 and 31 August
1968. Water temperature was 31° C. I found the pale
yellow spawn and from 1 to 12 individuals in the process

Page 442

Figure 1

Cerithium stercusmuscarum VALENCIENNES, 1833
The egg ribbon

Figure 2

Cerithium stercusmuscarum VALENCIENNES, 1833
A segment of the ribbon

of depositing more on the underside of almost every
rock in an area of approximately 12m’ at the water's
edge.

The loops vary in orientation, height and spacing.
Although one surface of the ribbon is usually continuously
attached to the substrate, occasional loops stand free. A
small segment of a ribbon that covered 32 cm’ is shown
in Figure 1. The tube js slightly flattened and measures
0.25 mm><0.5 mm in cross-section. The eggs, averaging
0.1 mm in diameter, are very numerous, 50 or more being
packed into a 4 mm segment of the long ribbon.

The figures were drawn by Anne Acevedo of the San
Diego Natural History Museum.

LITERATURE CITED

Lesour, M. V.
1944. The eggs and larvae of some prosobranchs from Ber-
muda. Proc. Zool. Soc. London 114: 462 - 489

THE VELIGER

Vol. 11; No. 4

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Page 444

BOOKS, PERIODICALS, PAMPHLETS

Die europaischen Meeres-Gehauseschnecken

(Prosobranchia)

vom Eismeer bis Kapverden und Mittelmeer

by Dr. Frirz NorpstEck. 1968. viii+273 pp.; 1200
line drawings on 31 plates; 16 color pictures on 4 plates.
Hardcover, 48.- German marks; Gustav Fischer, publ.,
Stuttgart, West Germany.

This book will be a most welcome addition to the malaco-
logical library of museums as well as collectors in general.
Although the scope of the work is limited to shell-bearing
prosobranchs of the European seas (including the Medi-
terranean), and although it excludes the pyramidellids
(as these animals are now generally believed to be opistho-
branchs), it still covers 1220 species and 180 subspecies
and includes the descriptions of 5 new species and 7
new subspecies. While the worker who is not very con-
versant with the German language may have some diffi-
culties with the text, he will find the approximately 1200
line drawings, illustrating almost every species, of great
help.

The treatment of the species is thorough and consistent.
Not only are all major taxa succinctly defined (i. e., super-
families, families, subfamilies and genera, but in the latter
case the type species is given, as well. The treatment of
the species starts with the currently valid name, followed
by synonyms, if any. Size indications and distribution data
precede the diagnosis of the species. Color indication
(for the animal), anatomical and radular characteristics
are mentioned, where known or references to generally
available literature are listed.

In the opinion of this reviewer the unusually large store
of important information contained in a very compressed
form through the liberal use of abbreviations could be
made more easily available to the non-German worker
if a list of all abbreviations were appended, or, at least
of some of the less commonly encountered ones, such as
“idR.” which stands for “in der Regel” or, in English,
“as a rule”.

RS

The Zoological Taxa of William Healey Dall

by KENNETH J. Boss, JosEpH ROSEWATER & FLORENCE
A. RuHorFrF. U. S. National Museum Bulletin 287:
427 pp., 1968. Available from Superintendent of Docu-
ments, U.S. Government Printing Office, Washington,
D.C., 20402. Price: $2.50, paper cover.

THE VELIGER

Vol. 11; No. 4

William Healey Dall described and named a total of
5 427 taxa. Of these 5 302 belong to the mollusks.

Many of Dall’s papers were published in periodicals
that are now either very rare or difficult to find. The
authors have made a careful compilation of all names
created by Dall, omitting only the suprageneric taxa for
very good reasons.

In their introduction the authors stress that the list
was compiled with a meticulous avoidance of rendering
decisions, or even suggestions, as to the proper assignment
of the various taxa, leaving this task for the specialist to
do when they review one higher taxon or another.

Of special value to the taxonomist is the account of
Dall’s early work in which he indulged in practices that
have caused much confusion. It might be stressed here,
however, that Dall apparently adhered meticulously to
the Rules of the International Commission on Zoological
Nomenclature after the Rules were established.

The alphabetical arrangement of the taxa created by
Dall make this book easy to use. It is not possible to
overestimate the value of this work and it is no exaggera-
tion when we say that it should be in the library of
every serious student of malacology. The cost of the book
can be this low only because it is produced by a non-profit
making organization, the United States Government.

RS

The Shell.

Five Hundred Million Years of Inspired Design
by Hucu and Marcuerite Stix, and R. Tucker ABgort.
Harry N. Abrams, Inc. $25.00; approx. 200 pp.; illus.

This is a gorgeous art book that serves to present with
a dramatic layout and design the beautiful diversity of
shell characters possessed by the world of living mollusks.
The 203 illustrations, including 82 hand-tipped plates in
full color, are the creative photographic work of H.
Landshoff. Dr. R. T. Abbott, the well known malacologist,
collaborated with The Stixs, who are New York shell
dealers, to provide a scientifically accurate commentary.
This attractive, large volume is not only oriented to the
general reader of natural history, but it will be especially
welcomed by students of art and design in nature, and by
the legion of avid shell collectors.
WK. Emerson

—

Vol. 11; No. 4

THE VELIGER

Page 445

The Helmet Shells of the World (Cassidae).

Part I. - R. Tucker Assott. Indo-Pacific Mollusca,
vol. 2, no. 9, 202 pp., 187 pls. Publ. by the Department of
Mollusks, Academy of Natural Sciences of Philadelphia,
Philadelphia, Pennsylvania 19103. $14.75.

ALPHA taxonomy concerns itself with the characteriza-
tion and naming of species and is often eclipsed by the
more spectacular results of BETA and GAMMA taxonomy.
But perhaps there should be an OMEGA taxonomy, an all-
there-is-to-know level of the study of organic diversity.
ApgotT’s paper is the OMEGA approach to a family of
higher prosobranch gastropods to which have been ap-
plied over a thousand names but which constitute about
60 living species. Part 1 of this monograph considers the
genera Cassis, Cypraecassis, Phalium, and Casmaria. The
world-wide coverage of the living species and the inclu-
sive treatment of the many fossil species constitute an
outstanding contribution to malacology.

The most interesting aspect of this paper is its illustra-
tion of the dynamics of evolution. In contrast to results
of typological approaches of the past, this study reveals
the probabilistic nature of biological phenomena and ex-
poses species as natural entities in different stages of the
evolutionary process. Species possess both spatial and tem-
poral dimensions. Recent populations are the survivors
of the past and some have stabilized a genetic system
which permits little phenotypic divergence while others
are spectacularly variable.

The results of this paper show:

1. polytypic species of a wide distribution whose nearly
indistinguishable subspecies are allopatrically separated
by thousands of miles (Cypraecassis testiculus testiculus
and C. t. senegalica; Phalium granulatum, P. g. centri-
quadratum, and PR. g. undulatum) ;

2. species in the process of forming geographic isolates
(incipient subspecies) with the ultimate prospect of gen-
etic distinction and speciation (Phalium areola) ;

3. polytypic species whose rapid divergence has resulted
in a fragmentation into ‘parapatric’ subspecies with vir-
tually little isolation (C'asmaria erinaceus) ;

4. two species of Xenophalium with a broad spectrum
of ecophenotypic and intrademe variation coupled with
the possibility of introgressive hybridization.

In general, the plates are excellent. The maps are of
value for the immediate assessment of distributional data,

but they should be referred to in the text under the respec-
tive species. The information included on depth distribu-
tion, bottom type preference and probable feeding habits
is important.

This work is uneven in the treatment accorded various
taxa. Cenozoic European species and subspecies of the
mamullaris (p. 55) and rondeleti (p. 97) complexes are
discussed more thoroughly than some fossil Phalium (p.
152) and Galeodosconsia (p. 123) where the species are
only listed. The use of a key to the species of Phalium s. s.,
but not to those of other genera and subgenera is unfor-
tunate. The term ‘forma’ has no biological or evolutionary
significance, and its usage in a modern taxonomic study
is deplorable.

‘Type-information is now generally included in the syno-
nymy ; it presents mistakes of omission: where is the type
of menkrawitense BEETS?; the syntypes of Cassis pfeifferi
are in the Madrid museum according to the caption to
plate 111, but they are not mentioned in the paragraph on
types of P bisulcatum, with which pfeifferi is synonymous.

A more trenchant discussion of the genera, their phylo-
genetic relationships and distinctiveness plus a diagnostic
key to the Recent genera and subgenera should be included
in the second portion of this monograph as should an
expanded bibliography which includes all the papers cited
in the text.

The reviewer wonders how the same author who recog-
nizes the great variation in Phalium bisulcatum and syn-
thetically treats the Phalium coronadoi-wyvillei-pilsbryi
species complex can then describe a new species, P kurodai
(p. 105, pl. 87) which differs from the ecologically and
geographically sympatric P carnosum (p. 104, pl. 86) by
a mere number of knobs.

An important error in type-setting occurs on p. 200.
The remainder of the paragraph on Types and Records
for Casmaria ponderosa nipponensis is found on p. 201,
bottom of first column under C. ponderosa perryi. And
the subspecies discussed under ponderosa are placed in
the polytypic species ertnaceus in the captions on plate 14.

Dr. Abbott is to be congratulated for a substantive con-
tribution to the malacological literature. His study has
shown that to obtain a modern systematic revision of a
cosmopolitan family, research must be done on a world-
wide basis and not be constricted by the parochialism
which has characterized much systematic work.

Kenneth J. Boss

Page 446

THE VELIGER

Vol. 11; No. 4

Oceans

a monthly magazine, published by Jack C. Reynolds.

Subscription $9.- from Oceans Magazine, P. O. Box 1820,
La Jolla, California 92037.

This new magazine, started in January 1969, is devoted to
the lore of the oceans. It brings in popular language and
with numerous black-and-white and excellent color illus-
trations articles on many topics. As may be expected, the
emphasis is not on mollusks, although this group of marine
creatures is by no means neglected.

An interesting feature of the magazine is the section
entitled “Sea Squirts” - written primarily for the young
reader, although of interest also to the layman in general.
Another valuable section is the glossary appended to each
issue in which scientifically accurate definitions of the
technical terms used in the articles are given.

Advertising is restricted to non-text pages, a commend-
able arrangement. It is to be hoped that this will not be
changed when increased circulation makes the magazine
an attractive advertising medium. It would be deplorable
to have an interesting account of salmon biology broken
up by sales pitches for diving gear or underwater photo-
graphy. As it is now, the magazine can be recommended

to any one interested in any of the many thousands of
problems connected with the oceans.

RS

A Checklist of the Marine Gastropods from the
Puget Sound Region

by Tuomas Rice. Of Sea and Shore, P. O. Box 33, Port
Gamble, Washington 98364. Approximately 170 pp.;
4 maps with list of collecting stations; frontispiece.

This mimeographed checklist is divided into an alphabet-
ical list of species, occupying 138 pages; a bibliography
of 9 pages; an index of 9 pages and a systematic list of
7 pages. Under each species in the first part of the list
are given references, followed by indications of the range
of the species; in most instances the type locality is also
mentioned and usually collecting data in the Puget
Sound area complete the entry. Where the author is
aware of synonyms, he is careful to include them.

A possible flaw in the list is the fact that, as stated in
the introduction, “Not all specimens have been checked
for the validity of the record, but those interested might
contact those collectors listed in the “Collecting data” sec-
tions in reference to particular species.”

RS

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, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.

It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.

Short articles containing descriptions of new species or other 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 geo-
graphical longitudes and latitudes added.

Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS’; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A.M. U.,
U. M. E., W.S. M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION 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 invited. The column “BOOKS, PERIODICALS, and 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, not
exceeding 81” by 11”, at least double spaced and accompanied by a clear carbon
or photo 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 accommodate the pamphlet (which
measures 514” by 81%’’), 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. Jerry DonouueE, 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

Dr. E. W. Facer, Professor of Biology
Scripps Institution of Oceanography, La Jolla
University of California at San Diego

Dr. Cavet Hanp, Professor of Zoology and
Director, Bodega Marine Laboratory
University of California, Berkeley

Dr. G Datias Hanna, Curator

Department of Geology

California Academy of Sciences, San Francisco

Dr. Jorn W. Hepcretu, Resident Director

Marine Science Laboratory, Oregon State University
Newport, Oregon

Dr. Leo G. HERTLEIN,

Curator of Invertebrate Paleontology

California Academy of Sciences, San Francisco

EDITOR-IN-CHIEF

Dr. Rupotr STOHLER, Research Zoologist
University of California, Berkeley

Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology

Stanford University, Stanford, California

Dr. Victor Loosanorr, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr. Joun McGowan, Associate Professor of
Oceanography

Scripps Institution of Oceanography, La Jolla
University of California at San Diego

Dr. Frank A. Pire.xa, Professor of Zoology
University of California, Berkeley

Me. Attyn G. Smirn, Associate Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

Dr. Ratpu I. Smiru, Professor of Zoology

University of California, Berkeley

Dr. Cuarwes R. StasEx, Associate Professor .
of Zoology

Florida State University, Tallahassee, Florida
Dr. Donatp M. Witson, Professor of Biology
Department of Biological Sciences

Stanford University, Stanford, California

ASSOCIATE EDITOR

Mrs. Jean M. Cate
Los Angeles, California

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