2

ih

re
Vo

” >! ae
Oy WSU aU Ry , . .% une
ones A ae vy eg ee te Ne Ay
ey : i TM Sales

oe ate
Wha

»

Ta

Veer egy ae
vy ‘ tye

. . A
ryt atate'y bey ently: ne ’ Caen ate em

4 3 aS AORN che Gers Are 4 SNe be

4 x * " ess Suey he : exe Lh

a x A was
: oa SN > re
sk Se

SUA sy a
*¥ Lae 2

ny een
; BOS SNE A Wine” Die he
1 , +
NSM NS id
tra he 2
:

yh
*

”
' Yimad prey
as

‘

a

Fit
SUR
‘IO

IORI GAT OR,
eee

Sif =
3 TASS
-

ee

x
Ve Oto:
LaSS HES a
SP AOAS BLED LS So
i So FAS SESAS
Ko Sg Paes
z Tae

fy hee

5B $7 GOT

Ftd 7 guage ys
$09 24 res
PIII FES 9
i ee

i
of

WV Pane

AA i Hel Rik oy cee ck eT USD ty a i NS) eae
a a + W”)

IN NOILNLILSNI NVINOSHLIWS $3 iyvugi7 LIBRARI ES SMITHSONIAN _INSTITUTION NOILONLILSNI NVINOSHLIWS-
no = n = = ”
id z wy" lw z= lJ z uJ z RN uw

_ wn a ” _— we 7 < ” = n ASS d
\ x =! 0 = a UG fg =, a a NSS. t&
)< z “ z “GI 2 B aS E
LS “ “ 5 = “Uy re Sc WAL =
z 5 e 3 NK a als = 5 =
a zZ * = ee a Zz Ne
11 LIBRARIES SMITHSONIAN [fr = 1 LIBRARIES SMITHSONIAN
5 z SARA zi = 5 il
w = a. wo = wo
\y Bel = x = .
Ss z et a
@ m “m
D Zz o Zz wo
IN NOILALILSNI NVINOSHLIWS IN NOILALILSNI NVINOSHLINS
g Z : z E
5 hy. = = l iy = =
5 £5 NBL 225s
zi = ‘ed William Healey Dall S 3 =
I7 LIBRARIES SMITHSONIAD | BB lj Hil IT LIBRARIES SMITHSONIAN
2 2) = w(t Wes a . -=
g = Yu, 1 Division of Mollusks | Ue = Yin, 2
) < Ue J / \ a sectional Library...) |) 3 = iy
(ox V4 Vise, . Hi ¥ | (ox “ffl , i a
+ © Bae us - 6, | : qed pa) ce eo a =
—_ mM .*” rt | ~— am.” eS
oO = | O = 40
z _J ce cad i Fa
IN NOILNALILSNI NVINOSHLIWS N NOILNLILSNI NVINOSHLINS
Ie =i CREME CER AR EA) 2 2 QS 2
F e E az 3
ee: BRARI ES SMITHSONIAN INSTITUTION OS SSN UNTO EUINIS S2 1YVvud roel BRARI ES SMITHSONIAN _
<x XX & = < ~~ = < = /x NE = <x
“= WS 5 : ZN 3 A f'2 EM Ay ere WS = -
FEN 7 SNS § LO 3 30 hf RNS F é
ZR 3 PN 8 GSH ? 8 GU 2K & P
EN 2 EW 2 WY = 2 “iy - \: z E
= q > s S . > 4 = > = . > S
7) a iis is aL ae 7) Zz 7) ae R
IN NOILALILSNI NYINOSHLIWS Saluvadi7_LIBRARIES SMITHSONIAN _ INSTITUTION NOILNLILSNI NVINOSHLIWS
lJ a Ws WwW = rey) = lu za Reed Ww

es YM Wee st n es eae aie ne Ez: n KE m
or a NS ioe a x. tip Ag au a =u AS ox
\2 2 WKS = : iL 3 < 2K =
OU NOt aU re A OE
2 5 oe ) ey O = O a oa
an = x _) aa pas) z 4 PA x ay
11 LIBRARIES SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLINS S3/1Y¥VYUEIT LIBRARIES SMITHSONIAN
— iS and ma.
= yg ow = ow = = w

= 5 FY: E : z : 5 ae
> 5S EL = > = > 2 Was
2 — OG hi ® = x = 2 =~ Oh? ®
m ” m 2 m = m ” m
re) = w = 7) = w Zz w
IN NOIMLALILSNI NVINOSHLINS S3S1YvVudI1 LIBRARIES INSTITUTION NOILALILSNI NVINOSHILINS |
29) z n Zz s n Pals (2p) ()
= I 4 < = 777 2 = = < = i 4 =

= jy UY z = by

= Wy, zZ 2 YY. & 2 3 i yy =
8 YG? 8 ot; 2 3 2 8 Yb 3
a Ce gee yen a Be z Ee Zak Zz
>’ = > = > s >’ >
za 7p) Zz 7p) Zz ” a P24, Zz

11 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_ NWINOSHLINS S3IYWYEIT LIBRARIES SMITHSONIAN |

Wa:

x
x, \
¥

NOILNLILSNI
NOILNLILSNI

NOILNLILSNI

SMITHSONIAN INSTITUTION NOILNLILSNI

SOW,
“0 aS <P>
AMIS

LIBRARIES NVINOSHLIWS

K&
AX |

ON NOILNLILSNI

i
DL
=)

NVINOSHLINS S3I1YVYEIT

11 LIBRARIES

<EHSON/ 2
xy

=
>

wis

; TSW
Ss IN \ S,
"

<S

7

4\
A
a

Yai _ LIBRARIES
fYaiT_LIBRARIES SMITHSONIAN

yi, Sb
UTION
BB)
(a
B
S
wy

JYaGI1 LIBRARIES
UTION NOILALILSNI

UTION NOoOIin

UTION

UTION

“at

f,

.

st)
st}

Ea eh ie ret gd, ae ee a ae a een ae Ng ee ae Soe er rm te
z 5 a oe v me ?
17_LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS S31Yvu a7 LIBRARIES SMITHSONIAN
= Ww = = ep) > =
Pade WwW z tJ Fa “f re Ww z LJ «A
[op] IN (ee) 2 UY
V2 = 4 = % WX = zl = “s
2 = : : 2K 3 : <
S a o = Zz = oe Py fea) =
5 2 fe) = ro) ea je) = Z C
mA aS 4 al ee Sti Ze, an z
ON NOILNLILSNI S3Jlyvudit LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLIWS _
3 = S = S ie S eK
z bs iF
= w — w — ies] = Oo > =
2 2 = : E 2 5 a
= > ‘ = > = Ri = = \ NWR t
= 2 XW E a =. a ae MW
17_ LIBRARIES SMITHSONIAN INSTITUTION NOILMLILSNI NVINOSHLINS SSIYVUGIT LIBRARIES SMITHSONIAN
= ” Zt is, a) Zz wo ; pau tea wo ;
< = RE See = 3 s = :
BE : Ee \w i &? 2 : : |
ZA ” N n ” ~~ WS 4S) @) WE Vf 2) (2) “”) a) ¢
as Sy Oo ae Y DANS fe) YG fog ag (2) ae 2) 3
Bot S Zz BE ww 277 = Zz E Zz
7 RE Sie : : ; : :
ON NOILALILSNI_NVINOSHLINS SAIYVYAIT LIBRARIES SMITHSONIAN INSTITUTION NOILMLILSNI_NVINOSHLINS
th | x ie Ag
u a : z : “ é !
» a = & vied a =
2) & A 2 = S £5 = =
wy = = S ec S o = !
= S . = re ay 5 a je |
a3 2 - 4 aS) Zz pay Zz
17 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31yvudi7 LIBRARIES SMITHSONIAN
a = fre z ~ Are z is eS
o 2 ace) S) w Xe ° ow 2
D = Pe a = WSS = v =
ay = k S = rl
ea FH, = = > ras > E :
a - Gy a = a = 2 -
. “UL enVte INOSHLINS 7 ms = % SNI_
— NOIL NOSHLI NOILNLILSNI
on, NO SNI_ NV Ws Saldvaal LIBRARIES, _!NSTITUTION 0 be
= << = < = < NS = was
5 2 eas ie ar 2S ey ae S z 4 Zz
8 Oy = 2 Gy =X 2 : Z :
Si? n z= 7) le Mie a Wiel pend 7) :
J17_LIBRARIES _INSTITUTION NOILMILSNI_NVINOSHLINS Sa1dvudi7_LIBRARIES SMITHSONIAN
2 Ww wi = eX it Zz a ;
op) me Sash set CO rae wo ASS ais n aa Ayre
| ZL, Fey s =f + INS NS. + 7 wF Sy, ;
S «Uy Si E SM, = : = Uy,
eS a0 “U@ 3s = = . a 3 a fp |
Zz 5 is z 2] Zz ey Zz iy
ON NOILNLILSNI_NVINOSHLINS_S3IYVYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILOLILSNI_NVINOSHLINS |
z ca z ia z aie = Gig, |
= oO . — o _ Vp ise) = ies)
\5 2 Wg 5 7 5 Gy 2 5 a !
3 5 = YEE = t GY 2 = = |
J EVER QE a Ee feo S = ie Ne
2 S\\ ee m ODE nas op) : (
& Gator ne a z O z O pay
17 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NWINOSHLINS S31uvudly ONIAN
Zz ~ n = ¢ 122) =z Ww ; = wn 5
< = <= = vy <x = = =
Ae lead =] 5 GY = Zz 4
eS NA: g z Ie: : 2 2 |
EW? : 8 fe * : : 2 |
Ee . > me 7 = i", > 4
2 x 2 = = = = = ~ =

NN -
ON NOILNLILSNI LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_ NVINOSHLIWS

179) Ww = w = (ep) = {
“ : é : é : & ;
o m= zt feed Sica) feed ad {
< _< oa < SoA < Zz ssi
o~ co S ~ S x =
(am = re ae 5 = je) Wr!
a 4 2 ai z Jj 4 :
117 LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLIWS Sa!uvugi7 LIBRARIES SMITHSONIAN
fl oat a ~~ a ise >
. ow = “0 Sy ° = NS @) ¥. ay EHSONT eS |
wo is ON >» = i Fao) is o> = E nag Ds wo SS 2) 4 avis > o ss 4N = UA Ie.
SS LAD Ey fi a LAM) = Game 2 NSE Gam 2 (AM) SF UV KX

Division of Mollusks

Sectional Library

Wiel

PeELIGOER

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

Division of volusk®
Sectionc! Library

Volume 10

July 1, 1967 to April 1, 1968

Page II THE VELIGER Vol. 10; No. 4

Foreword to Volume 10

With the completion of Volume 10 we have reached a milestone on the road of the development of The Veliger.
This is, then, an occasion to take stock of the past — to review the accomplishments and to assess the failures.

Of all the goals we had set for ourselves about ten years ago, we think we have attained most: our journal has
been accepted on a world-wide basis, as evidenced not only by the ever-increasing number of subscribers and
members in all parts of the world, but also by the fact that we receive manuscripts from authors in many lands.
Our aim to produce a journal of high quality, not only in content, but also in appearance, has been achieved, we
believe, within reason. Two important failures, however, still plague us. The first and, to us, an important one is
the fact that we have not succeeded in producing an issue completely free of all typographical errors and this in
spite of extraordinary precautions we have taken, and also in spite of the unusual skill brought to the task of proof-
reading by Mrs. Jean Cate of Los Angeles. The blame for the remaining errors rests with the editor himself.

The second failure is the one that has plagued us since the first issue -— finances. But again, as in the past, we
must assume the responsibility for this: we have steadfastly refused to permit this problem to force us into seeking
aid through applying for governmental grants. That our situation is not completely untenable is due to the most
generous contributions made by a few individuals in the form of many uncounted hours of volunteer labor. Fore-
most among these volunteers are Mrs. Emily Reid whose artwork continues to lend that extra touch of beauty and
attractiveness to our journal; Mrs. Cate, who not only checks every word set in type for correct spelling but also,
with the aid of a magnificent malacological library and a sound knowledge of the field itself, often is able to pre-
vent erroneous citations; Mrs. R. Stohler who, through the years, has not only assisted with many menial tasks,
such as collating, preparing for mailing, recording and checking and double-checking, but also has encouraged the
editor in his desire to avoid asking for governmental assistance.

What measure of success we have had with the development of The Veliger is also due to many contributions
made in various ways by a large number of persons. If we single out a few for special mention, it is not to imply
that the contributions of those not listed were less important. Among those who have assisted the editor with their
technical advice are Mr. Armand Lamadrid of Los Angeles, an efficient linotype operator and mechanic as well
as a staunch friend; Mr. John Schoen and the late Earl Gustafson of the Printing Department of the University
of California in Berkeley have been ever ready with helpful advice, suggestions and real practical assistance.

Another group of men and women who have given and continue to give unstintingly of their talents and time
are the members of the Editorial Board who spend many hours reviewing the manuscripts submitted for publi-
cation and whose recommendations are the guides for the decisions by the editor. Last — most assuredly not least,
the members of the Executive Board of the California Malacozoological Society give generously of their time,
talents and substance.

Finally, the craftsmen at the Printing Department, who seem to take special pains to turn out a high quality
product deserve to be included in this list.

To all the many individuals, named and unnamed alike, we express our deeply felt gratitude, hoping at the
same time to be permitted in the future to continue taking advantage of their knowledge, skills and talents so
that The Veliger may continue to develop and, hopefully, to become self-supporting at a time not too distant in
the future.

Sincerely,
Your Editor

Vol. 10; No. 4

TABLE OF CONTENTS

A checklist of intertidal mollusks for Bahia Willard and
the southwestern portion of Bahia San Luis Gonzaga,
State of Baja California, Mexico

Heten DuSHANE & GALE G. SPHON ......... 233
A ctenostomatous ectoproct epizoic on the chiton Ischno-
chiton mertensiu
Eucene S. HELFMAN
A new cowrie species from the Philippines
CrawrForp N. Cate & FRANZ A. SCHILDER ..... 382
A new marine mollusk from Mozambique in the genus
Festilyria Prtspry & Oxsson, 1954
CGEIRTON STOKES) WEAVER | 3)..000 cleo eee + 442
A new species of Marginella from the coast of Brazil
JeEAN-JAcgues VAN Mot & BernaArD TurscuH 196

A new terebrid species with check list of Terebridae from

the Red Sea
Twita BRATCHER & RoBert D. BurcH

Aplysia vaccaria, a new host for the pinnotherid crab

Opisthopus transversus

INNEHONE CRAIG BEONDE i.e )aicie oe cele ele ls 375

A quantitative sampling of the mollusks of Batangas Bay,
Philippines

AUPANN IV Les CUNT Seiya statcsche gel ovoi</ssre.64) save clorons\e)z1 ove 83

A radula muscle preparation from the gastropod, Kelletia
kelletiit, for biochemical assays

Howarp M. FEeper & REUBEN LASKER ........ 283
A re-interpretation of the sand-pipes described by ADE-
GOKE

NJOLUNMWVMEVANS prc Aeon listen ererets elecensetete ts 174
A remarkable new cancellariid from the Philippines, with
comments on other taxa
\ NATE), (COC iss yo ceo Ben Ge eo oraN 286
A review of the living tectibranch snails of the genus
Volvulella, with descriptions of a new subgenus and
species from Texas
TAR OLDY Wi EVARRY oc tons ciel sieic visio vais vee we 133
Boiled lettuce and cress as diet supplements for certain
species of mollusks

STW VOI ERE: Rimmoeee eect me Seca, ict ta ne. Sears, 446
Coan molluscan collection at University of California,
Davis

AMESBWON/ABEN TINE 9 (rjc 0s 0c celeisie sles eaiere see 293
Effects of feeding by Armina californica on the biolumi-
nescence of Renilla koellikeri
IAN SWBERTS CH | 5 there cette svaisucrege a dale ee 440
Enzymatic defenses of certain snails against metal ions
Linpsay R. WINKLER & Lots Wonc CuI ...... 188
Epitonium (Asperiscala) billeeana (DUSHANE & Brat-
CHER, 1965) non Scalina billeeana DUSHANE &
BratcHer, 1965
Heten DuSHANE

THE VELIGER

Page ITI

Freshwater mollusks collected by the United States and
Mexican Boundary Surveys

Dic HTaWahAViOR won) ss ene ale seco ole eles 152
Invalid names in oysters

ISENINETH, S)AVBOSSie o/s -tekeraiels isin toskeca siaveictel: 447
Itinerary of the voyage of H. M. S. Blossom, 1825 to 1828

JOSEP HE ROSEWATERD A477 yr evil aetiier 350
Marine fouling and boring organisms in Monterey Harbor

EFC pEVADEREIER sacs celestial: RSS eke n iolebeee 327
Mating behavior in Mitra idae MEtvit1, 1893
EAN SMP CATE atten talcoiels canter sacle soln eee o 247

Mode of feeding and diet, and synthesis of studies on
marine pelecypods from Tomales Bay, California

DON SMIAURER En cia ctv ne iiotrtd Aetren oa eras 72
New checklist in preparation

UD Ya SieADERRV A nye stele crersia Mesto Se bin oe tsoloa eres 89
New record of Conus ebraeus in Costa Rica

J CSEPHEREHOUBRICK MERE Eran eee eer nir 292

New records of Nudibranchia (Gastropoda : Opistho-
branchia) from the central and west-central Pacific
with a description of a new species

DAVIDSKCAVOUNG © causes Wee Ca ece he tee 159

Note on the northward spreading of Mya arenaria Lin-

NAEUS in Alaska

AMES Bo GROSS 2%, uReyaaee ty, horeania acim nea ee 203
Note on the radula of Mitromica Berry, 1958

JAMES) EV MCILEAN (eterno eee ae aes 58
Notes on cephalopods from Northern California

ROBERT: Re WALMADGE 75) svn). « cistteler so cle Nomtnsierls 200

Notes on Morum dennisoni (REEVE) and related species
S. PETER DANCE & WILLIAM K. EMERSON ..... 91
Notes on the range extension of the boring clam Penitella
conradi VALENCIENNES and its occurrence in the
shell of the California mussel
STEVEN) Ey MEREDITH «214. doteioitiec eens eee 281
Observations on Hipponix conicus (SCHUMACHER, 1817)

WALTER OLIVER CERNOHORSKY ......--se0c0: 275
On the identity of Phos laevigatus A. ApAMs, 1851
WIERTAMOKeR ES MERSONSEOM See ene: 99

Preliminary observations on the feeding behavior of
Conus purpurascens Broverip, 1833
James W. NyBaKKEN
Relationship between Penitella penita (Conran, 1837)
and other organisms of the rocky shore
Joun W. Evans
Reproduction in Olivella biplicata
DA CRAIGZEDWARDS sc) ieselce siete otele eleven era cir cie ate 297
Role of snails’ disease in the biological control of Achatina
fulica Bownicu, 1822 on the Andamans
P. D. Srivastava & Y. N. SRIVASTAVA ......... 320
Semele martini (ReEEvE, 1853) of southern Brazil and
Uruguay
MicueEt A. KLAPPENBACH

Page IV

THE VELIGER

Vol. 10; No. 4

Spawning notes, I. — Hexaplex erythrostoma
Fay Henry WoLrson
Structure of the bivalve rectum. - I. Morphology
Tuomas C. JecLA & MicHaEL J. GREENBERG ... 253
Structure of the bivalve rectum. - II. Notes on cell types
and innervation.

Tuomas C. JEGLA & MICHAEL J. GREENBERG .. 314
Studies on East Australian cowries
Marta SCHILDER & FRANZ ALFRED SCHILDER ... 103

Studies on the Mytilus edulis community in Alamitos Bay,
California: — III. The effects of reduced dissolved
oxygen and chlorinity concentrations on survival and
byssus thread production

Donatp J. ReisH & JosepH L. Ayers, Jr. .... 384

Studies on the vitality of the Japanese pearl oyster Pterza
(Pinctada) martensu (DUNKER) under abnormal
conditions. - I. Oxygen uptake and shell movement
in sea water of low oxygen content

AUETUO MEIVAUTIs Fores cysregsislovcxsie| neter su arorstetarereterens 342

Taxonomic placement of Coralliophila incompta BrErry,

1960, with the proposal of a new genus, Attzliosa
WTA UINYE IK IVT So0d000000c0000800000 379

The behavioral role and the structure of the aesthetes of

chitons

JONG GK © JMG OIC COVE ON mewn cata MIRAI. UNM TE I oe 77
The cowries of the Ryukyu Islands
CRAWEORDEINEILICn CATER Cerin 13

The date of publication of K1ENER’s Mitra monograph in
the “Spécies général et iconographie des coquilles
vivantes”

WALTER OLIVER CERNOHORSKY ...........+5. 349

The egg mass and veligers of Limacina helicina Puirrs

Wine AN, JUNUNNTNDS 5 o0000d00000000000000 322

The functional morphology of Lyonsia californica Con-

RAD, 1837 (Bivalvia)

WATTERG NAR CHITIn eieicg ways etion enna arena ae 305
The generic classification of cowries
ERAN ZPACEREDESCHILDER Mn 264

The Muricidae of Fiji. Part I - Subfamilies Muricinae

and Tritonaliinae — _ plus an addendum
WALTER OLIVER CERNOHORSKY .............. 111
The Ovulidae, Pediculariidae and Triviidae of Fii
WALTER OLIVER CERNOHORSKY ..........+0- 353
The radulae of nine species of Mitridae
JEANG MUG CATE g ccrocie: scopes siiscstohorensna nko mer nets cnet 192

The radula of Zierliana woldemariu (KiENER, 1839)
Jean M. Cate
The rediscovery of Erosaria menkeana (DEsHAYES, 1863)

CRAWHORDENEILIN CATER nreeneieri incr 198
The reproductive system of the British Turridae
EDMUND SEL. SMITH ‘5 shi0iesiescr mir Pera tseene says 176

The retention of lamellibranch larvae in the Niantic
Estuary
Jounes K. Moore & Nr1son MarsHALL ..... 10
The role of behavior in determining the intertidal zona-
tion of Littorina planaxis Puiuiprt, 1847, and Littor-
ina scutulata Goutp, 1849
Cart E. Bock « RicHarp E. JOHNSON ........ 42
The shell ornament of Hysteroconcha and Hecuba (Bi-

valvia): a test case for inferential functional mor-
phology
Ropert M.. GARTER 4 S51. pio a%.c yee so ee 5S)
Two new species of British turrids
EDMUND H. SMITH) 26 cals so-so eee 1

West American mollusk types at the British Museum
(Natural History). - IV. CARPENTER’s Mazatlan col:
lection

A; Myra KEEN} gs. oes cae oe see eee 389

Western Australian cowries —

expanded report

G@RAWEORDENEIIT CATERER ener nee 212

Zonation in marine gastropods of Costa Rica and species
diversity
GERALD J... BAKUS, (o-/os0itis fi croc cee 207
AUTHOR INDEX

BaKUS; GERALD Jicetertt- sete ee een 207
BEONDEWANDEONE CRAIG EEE eee rrr 375
BERTSGH, HANS) caus 5 seins ewes ude aconearoG een 440
Bock, Cari E. & RicHarp IX. JOHNSON .......... 42
Boss, KENNETH) JAY: ot ceccits aricade eee 447
BRATCHER, TwitA & RopertT D. BurcH ........... 7
Burcu, Rosert D. see BRATCHER, TWILA & —
CARTER, ROBERT) MU 15 ois. /lere. ave euczsroers -ceneco here eee 59
CatTre, CRAWFORD NEILL ............-- 13, 198, 212
Cate, CRAWForRD N. & FRANZ ALFRED SCHILDER .. 382

CATES JEANAM IS cers ees ieee 5, 83, 192, 247
CERNOHORSKY, WALTER OLIVER ... 111, 275, 349, 353
Cut, Lois Wonc see: WINKLER, Linpsay R. & —

Dance, S. PETER & WILLIAM K. EMERSON ......... 9]
DuSHANE; HELEN) 253600). sc teen sso enone 87
DuSHane, HELEN & GALE G. SPHON ........... 233
EDWARDS) Ds GRAIG (as cides eter ererie eter ete 297
JOneT SON, WMATA I, 5 0050000000000000000 99, 379
see also Dance, S. PETER & —
EVANS) OHING Wilh ceee ter iiriarer 148, 174
FEepER, Howarp M. & REUBEN LASKER .......... 283
BYSHER( TSW: Suracdismiaxonsgeyeas torsenneniee toe eee e 446

GREENBERG, MicHAEL J. see JEGLA, THomas C, & —
(GOS, |VANERS 135 ob 00d00000000050b000000000000 203

Vol. 10; No. 4 THE VELIGER Page V
EWADERTIE 0B ey Cin tive tie sivisetevece sift oseretsit cy sis areraaierslats 327 ReisH, Donan J. & JosEpH Ayers, Jr. ........ 384
EVARRYAELAROLD Walia raya eerste aishevaisieiers et cVeleseret a'sa je 133 ROSEWATER © GJOSEDHE i ioie sca neusiatle ace «o-cassoelsr 350
UETPEMAN SEUGENE USS sreinciniea cis « erietest sien esis cl 290 ScHILDER, Maria & FRANZ ALFRED SCHILDER ..... 103
HIERTIEIN, LEO G:, .....-5..-.. (205))5(294)))(295)) SCHIEDER SURANZPALFRED) see sciatic ee elec e 264
HIOUBRICK,, JOSEPH (Reo) 23.6 ieee cise oe ons oe ne 292 see also Cate, CRAwrorp NEILL & —

Jecia, THomas C. & MicHaAEL J. GREENBERG 253, 314 see also ScHILDER, Maria & —

JouNnson, RicHarp E. see Bock, Cart E & — SMITH yD NGUIND REISS Vanve emis es caiaiens ovsiaisiave: sree 1, 176
IKSERN WAC ENIYRA)) or cejersts bisjere.o:0:e eres (90), (295), 389 SPHON, Gate G. see DuSHANE, HELEN & —
KQAPPENBAGH, MIGUEL .........0-000csecc00s- 274 Srivastava, P. D. & Y. N. SRIVASTAVA .....-.0000. 320
Lasker, ReuBEN see FEDER, Howarp M. & — Srivastava, Y.N. see Srivastava, P D.s —

MarsHALL, Netson_ see Moore, JoHNES K. & — STOHLER, RuDOLF ...... (90), (205), (294), (296)
NIPSO RE R DON Wey etic esches nna wiasiseldceie.oFiaiete so eens in Se UE orcs epoke eer eyey tae tesa emse cetera 4 (449), (450)
VICTORIAN MMJAMES IETS /el05 ayale olorsis drei e aiisievetesieie es! « 58 SPAT MADCE SROBERTE RGM Nene eer rneeeicie rin 200
IMIEREDIRH A OTEVEN Eo io ayeicie ike oats) oisle oieleis ole ore ere 281 HEAVLOR NS MOWAG HT Tra Wott eieeverettlayavccais ereisrea terre eve eacis 152
NAT ANU TIAMAT WO)e arene nied ahemesecetorseavols eevee neal aos 342 PEERRY Se UD YAS eee EY. ee yeh rai iuacausidl hearst eo eaneeraeees 89
Moore, JoHNES K. & NELSON MARSHALL ........ 10 TurscH, BERNARD see VAN Mot, JEAN-JACQUES & —
INARGHTMEVWATBERY lalvereiecis cteiare oy ciesie) seseuer oltanguotstee ¢ 305 WALENTINES AMES SW) rinih. «niauetnn micros acm ae nr 293
INVBAKEIGEN, HJAMES (We 1 viccjn cis eise tela dies cesses as 55: Van Mot, JEAN-JACQUES & BERNARD TurRSCH .... 196
ORD MAVVIETIAN EJs oni alel ceca sie cles wa arene da eyes 286 WEAVERS CLIRTONGOTOKESM se criada tie ene: 442
©) NaETAT CETe a Pe AUT ares cy aces a Siete close euekevemceuerieis 6 77 WINKLER, LinpsAy R. & Lois Wonc CuI ........ 188
BARANJAPE, MADHU J... 6. cece cece cee ete sees 322 WOLFSON WPAYSHENRY. 1afeeleceiieeonrielieieasis aise 292

Younc, Davwm K. ........

SU OS osOnC OSHC UONDe US 159

2!

Ad ii 7
“a oF
io iy » — :
cs a i
: ¥ =e
7
‘
wi
ise ‘ ny
= ee
;
i ~
a
*
el iat
Sep ry
’
-
he
'
,
t
i) \
ee
}
!
=
tir
Hi
s
ho .s
€ ‘
te P

Net)

é ay hy

le

Moll. THE

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME 10 JuLy 1, 1967 NuMBER I
CONTENTS

Two New Species of British Turrids (Plate 1; 5 Text figures)

EDMUND RLIROMUEEL corm aey Wyn tle io Mgot., say AMncy. 5). o 7 GMy ego) ede Seater vee. oe COE
The Radula of Zierliana woldemarti (KiENER, 1839) (3 Text figures)

JJSEINT IML, Ginga 2 ics > ho BO Bun nn area Mic au een a Re aera ee er 2
A New Terebrid Species with Check List of Terebridae from the Red Sea

(Mollusca : Gastropoda) (Plate 2)

A WAlAY BRATCHER IE ROBERTI BURGH) =.5 . sou sect) sc) cic: ees hecon men
The Retention of Lamellibranch Larvae in the Niantic Estuary (1 Map)

JOHNESHIC MOORE & NELSONSMIARSHALL 5 4 iG. - =) sees) 10

The Cowries of the Ryukyu Islands (Plate 3; 2 Maps)

CRAWEORDONS CATE) 2oeammr en ee on Se heme pe akg ats alle, est ES
The Role of Behavior in Determining the Intertidal Zonation of Littorina planaxis
Puuiuiprl, 1847, and Littorina scutulata Goutp, 1849 (8 Text figures)
CARE PE MBOCK & RICHARD ba JOHNSON 4500-0150 cy = lie evel ek ss ee, | 42

Preliminary Observations on the Feeding Behavior of Conus purpurascens BRODERIP,
1833 (Plate 4)

IPARUES \WOINSBAKICEN ji nkt-s eRe eich i) a a, Maier me Beene gy ee ase 5

Note on the Radula of Mitromica Brrry, 1958 (1 Text figure)
(JAMESPEONMIGIEE AND: 7 40-) SMMC CCUG ete asst ea: eagle ole yiane Sy eS, Berea GO

The Shell Ornament of Hysteroconcha and Hecuba (Bivalvia): a Test Case for
Inferential Functional Morphology (Plates 5 to 7; 2 Text figures)

ROBERT ANIC ARTER 2 0n ai. stl a ede acy Gane Seen UC Das. ar Mee a ueT le stan hO)

[Continued on Inside Front Cover]

Distributed free to Members of the California Malacozoological Society Inc.
Subscriptions (by Volume only) payable in advance to Calif. Malacozoological Soc., Inc.
Volume 10: $12.- Domestic; $12.60 in the Americas; $12.90 all other Foreign Countries.

Single copies this issue $4.25. Postage extra.
Send subscriptions to Mrs. Jean M. Cate, Manager, 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

Mode of Feeding and Diet, and Synthesis of Studies on Marine Pelecypods from
Tomales Bay, California

DON MAURER ioe 2k ee Re eg yeh PRO ers Css ey ee ea een a

The Behavioral Role and the Structure of the Aesthetes of Chitons (Plates 8, 9)
PauL OMELICH
A Quantitative Sampling of the Mollusks of Batangas Bay, Philippines
(Plates 10, 11; 1 Map; 2 Text figures)
JEAN MEICATE © 275), es Gos) Grate) cel canine ne cy ae

Epitonium (Asperiscala) billeeana (DUSHANE & BRATCHER, 1965) non Scalina
billeeana DUSHANE & BRATCHER, 1965

ELEN DUSHANE? 03" ap as, Centres et el iaree gems
NOTESU& NEWS igo ee ee io echt Sa ott un
BOOKS, PERIODICALS & PAMPHLETS .............

72

71

83

87

89

go

eS SS SSSSSSSSSSSSSSS—SSSSaaaaaaa>s
Note: The various taxa above species are indicated by the use of different type styles as

‘shown by the following examples:

ORDER, Suborder, DIVISION, Subdivision, SECTION,

SUPERFAMILY, FamILy, Subfamily, Genus, (Subgenus).
New Taxa

Vol. 10; No. 1

THE VELIGER

Page 1

Two New Species of British Turrids

BY

EDMUND H. SMITH

Department of Zoology, University of Glasgow, and the Marine Station, Millport '

(Plate 1; 5 Text figures)

THE PRESENT CLASSIFICATION of the Turridae is based
almost entirely on shell characters. The weakness of such
a Classification, however, has been clearly demonstrated
during a recent study of the turrids of the Clyde Sea Area,
Scotland in which almost identical shells were found to
“house” radically differing animals. Thus, while some spe-
cimens of “Philbertia leufroy: booth (Smitu, 1839)”
from the Clyde Sea Area possess the usual complement of
radula, poison gland and salivary glands, others, almost
indistinguishable conchologically, were found without
any trace of these structures. A second, apparently unde-
scribed, turrid from the same area was found to lack these
structures. It is considered that these findings warrant the
creation of a new genus and of two new species.

Cenodagreutes E. H. Smiru, gen. nov.

Type species: Cenodagreutes aethus spec. nov.

The shell characters are similar to those of Philbertia
(see esp. THIELE, 1931, p. 370), and as in that genus, the
operculum is absent. Internally, however, it can be distin-
guished from Philbertia, and probably from all other
genera of British turrids (E.H.Smirn, 1967), by the
absence of the radula, poison gland, and salivary glands.
Etymology: The generic name Cenodagrcutes is derived
from the Greek xevodov7is, toothless, and aypevtns, the
hunter, meaning the “toothless hunter”, alluding to the
lack of radular teeth.

Cenodagreutes aethus E.. H. Situ, spec. nov.
(Plate 1, Figures 1 and 2; Text figures | to 3)

Shell: The shell is small (7mm in length), fusiform, has
a short spire and seven to eight prominently convex whorls.

' Present address: Pacific Marine Station, Dillon Beach, California
94929.

The nuclear whorls number three and are diagonally de-
cussate. The postnuclear whorls are sculptured with very
prominent axial ribs which are crossed by thin, well defined
uniformly spaced spiral cords; on the penultimate whorl
the axial ribs number 14, the spiral cords 6. The orna-
mentation consists of very fine axial growth rugae and
small pustules covering the depressions between the spiral
cords. The suture is fine and there is no sutural shelf. The
labial aperture is ovate and the siphonal canal is short
and smooth. The sinus is shallow and occupies most of
the shoulder. The parietal and columellar calluses are
smooth and abruptly marked off from the surface orna-
mentation. The outer lip is thin.

Color: The ground color of the shell is creamy white, the
pustules are brilliant white and the spiral cords reddish-
brown. Over some of the axial ribs, however, the brown
coloration of the cords is absent.

General appearance: The foot is a translucent creamy
color, flecked on the sides and near the opening of the
rhynchodaeum with brilliant opaque white. The siphon
is uniformly cream colored.

Internal anatomy: The polyembolic proboscis (SmirH,
1967) is much shorter than that of Philbertia leufroyi
boothi (Text figure |). There are two projections (br)
on the mid-line of the floor of the rhynchodaeum (rh).
The two major introvert retractor muscles (rm) send
small bundles of fibers into each of the two small projec-
tions. The oral opening (00) is surrounded by a thin
circular muscle layer which thickens posteriad around the
antcrior ocsophagus (oc). The epithelium surrounding the
oral region and passing posteriorly for a short distance is
densely ciliated. Vhe glandular sheath (gs) which en-
circles the oral opening is short with a few thin muscle
strands leading from the fold of the sheath to the wall
of the cephalic hacmococl. After the oesophagus passes
through the nerve ring (nr) it enlarges; gland cells replace
the densely ciliatecl epithelium found in the anterior

Figure 1

Cenodagreutes aethus

Idealized diagram depicting longitudinal section through the
rhynchodaeum.

br mid-ventral bumps of the rhynchodaeum

ct circumoesophageal tensors gs glandular sheath
lr lip of the rhynchodaeum nr nerve ring oe oesophagus
00 opening of oesophagus th rhynchodaeum
rm major retractor muscles of the rhynchodaeum

rs _rhynchostome sr sphincter of the rhynchostome

region. There are no salivary glands, radular sac, or
poison gland. The intestine is covered by a thin muscular
layer and lined by ciliated cells. There is no anal gland.

Present in the male reproductive system (Text figure 2)
is a large vesicula seminalis (vs) and a connective tissue
strand (a remnant of the gonopericardial duct, gd) run-
ning from the vas deferens (vd) to the pericardial wall
(pc). An opening (mo) connects the vas deferens to the
mantle cavity and there is a convoluted prostate gland
(pg) which opens into a short penis (p).

In the female system (Text figure 3) the gonadial ovi-
duct (go) is composed of columnar gland cells and sur-
rounded by a thick muscular layer. The renal oviduct (ro)
is short, narrow, and enters the albumen gland (al)
ventrally. There is no gonopericardial duct. The albumen
gland is ciliated throughout with no subepithelial glands.
The epithelium of the albumen gland becomes continuous
with the posterior part of the capsule gland (cg). There

THE VELIGER

Vol. 10; No. 1

is no pallial oviduct. A short, constricted muscular duct
joins the anterodorsal part of the albumen gland with the
ingesting gland (ig) and functions as a receptaculum
seminis. The capsule gland has no ventral channel and
opens into a small bursa copulatrix (bc). This bursa is
muscular with a narrow lumen which opens directly into
the capsule gland. The epithelium lining the capsule gland
and extending a short way into the bursa is composed of
gland cells.

Type specimens: The holotype was collected off Farland
Point, Isle of Cumbrae, Firth of Clyde, Scotland (Lat.

T P PA
VS Cc HR P
PD
D
‘ GD MO PG
Figure 2

Cenodagreutes aethus

Diagramatic reconstruction from sections, of the male reproductive

system.
gd gonopericardial duct hr heart
mo opening from vas deferens to mantle Pp penis
pa opening of penis pc pericardial wall pg prostate gland
pd duct from prostate gland t testis

vd_ vas deferens vs vesicula seminalis

O GO IG CG
AL
BC
RO N00) GT
Figure 3

Cenodagreutes aethus

Diagramatic reconstruction from sections, of the female reproductive

system.
al albumen gland be bursa copulatrix cg capsule gland
go gonadial oviduct gt genital opening
id duct to ingesting gland ig ingesting gland

renal oviduct

la lumen of the albumen gland o ovary ro

Explanation of Plate 1

Figure 1: Cenodagreutes aethus, a view of the shell aperture.
Figure 2: Cenodagreutes aethus, a view of the labial sinus.

Figure 3: Cenodagreutes coccyginus, a view of the shell aperture.
Figure 4: Cenodagreutes coccyginus, a view of the labial sinus.

[E. H. Smiru] Plate 1

Tue VELIGER, Vol. 10, No. 1

Vol. 10; No. 1

THE VELIGER

Page 3

55° 44’ 20” N, Long. 4° 5730” W) on a bottom of stones
and mud in 20m. The holotype has been serially sectioned
and these sections, along with the broken shell, are depos-
ited at the California Academy of Sciences, Department
of Invertebrate Zoology, Type number 320. It consists of
4 microscope slides, CAS numbers 338 - 341. A paratype,
consisting of an unbroken shell is also deposited at the same
institution, Type number 321.

Etymology: The specific name aethus is derived from the
the Greek a:Owv, and refers to the red color.

Cenodagreutes coccyginus E. H. Smitu, spec. nov.

(Plate 1, Figures 3 and 4; Text figures 4 and 5)

Shell: The shell is small (54mm in length), fusiform, and
has a short spire with seven whorls which are somewhat
less convex than in Cenodagreutes aethus. The nuclear
whorls number three, and are diagonally decussate. The
postnuclear whorls are sculptured with very prominent
axial ribs which are crossed by thin, well defined and
uniformly spaced spiral cords; on the penultimate whorl
the axial ribs number 14, the spiral cords 6. The orna-
mentation consists of very fine axial growth rugae and fine
granules, in contrast to the pustules of C. aethus, which
cover the depressions between the axial ribs and spiral
cords. The suture is fine and there is no sutural shelf. The
labial aperture is ovate and the siphonal canal short and

SR RS

RM SM CT

Figure 4

Cenodagreutes coccyginus

Idealized diagram depicting longitudinal section through the
thynchodaeum.

ct circumoesophageal tensors gs glandular sheath

Ir lip of the rhynchodaeum nr nerve ring oe oesophagus

00 opening of the oesophagus th rhynchodaeum

rm major retractor muscles of the rhynchodaeum

rs_rhynchostome sm sheath muscles
sr sphincter of the rhynchostome

smooth. The labial sinus is distinct when compared to C.
aethus and occupies most of the shoulder. The parietal
and columellar calluses are smooth and abruptly marked
off from the surface ornamentation. The outer lip is
thicker than in C. aethus.

Color: The ground color of the shell is creamy white
with alternating dark reddish-brown spiral lines which
are broken occasionally by opaque white or yellowish
zones.

General appearance: The foot is translucent creamy white,
sometimes tinged with purple and covered with opaque
white spots on the sides. The siphon is uniformly white.
Internal anatomy: As in Cenodagreutes aethus a polyem-
bolic proboscis is present (Text figure 4). The projections
on the floor of the rhynchodaeum (rh) are absent and
the oral opening (00) is surrounded by a very thick
muscle layer. In contrast to C. aethus the epithelium sur-
rounding the oral region and passing posteriorly for a
short distance is cuticular. The glandular sheath (gs)
which encircles the oral opening is long with well devel-
oped muscle strands (ms) leading from the fold of the
sheath to the wall of the cephalic haemocoel. The oeso-
phagus extends into the oral sheath for some distance
forming “lips”. After the oesophagus passes through the
nerve ring (nr), it enlarges, as in C. aethus, with gland

© €O renee CG =
RO PO R | — 10
Figure 5

Cenodagreutes coccyginus

Diagramatic reconstruction from sections, of the female reproductive
system.

al albumen gland b_ distal bulb of the receptaculum seminis
be bursa copulatrix cg capsule gland
go gonadial oviduct gt genital opening
Oo ovary po pallial oviduct r receptaculum seminis
ro renal oviduct ve ventral channel

cells replacing the cuticular epithelium of the more ante-
rior region. There are no salivary glands, radular sac,
or poison gland. The intestine is ciliated and surrounded
by a thick muscular layer. There is a long, thin anal
gland which follows the intestine for some distance,
opening into its lumen near the anus.

Only female specimens of this very rare turrid have
been found. The female system (Text figure 5) is similar
to that of Cenodagreutes acthus with the following ex-
ceptions. There is a pallial oviduct (po) between the

Page 4

THE VELIGER

Vol. 10; No. 1

albumen gland (al) and capsule gland (cg). No sperm
were found in the small distal bulb (b) of the receptacu-
lum seminis nor was there any evidence of sperm ingestion.
The capsule gland has a ventral channel (vc). The bursa
copulatrix (bc) is much larger than that of C. aethus
with the bulb forming the distal end of the bursa and
lying at the anterior end of the capsule gland. This
enlargement forms a sperm filled sac. it is not lined by
glandular epithelium but by one composed of densely
ciliated cells.

Type specimens: The holotype was collected off Tan
Buoy between Great Cumbrae and Little Cumbrae Islands,
Firth of Clyde, Scotland (Lat. 55° 44’ 20” N, Long.
4° 57’30” W) on a sandy shell bottom in 17 m. The
holotype has been serially sectioned and these sections,
along with the broken shell, are deposited at the California
Academy of Sciences, Department of Invertebrate Zool-
ogy, Type number 322. It consists of 5 microscope slides,
CAS numbers 342 - 346. A paratype, consisting of an un-
broken shell is also deposited at the same institution,
Type number 323.

Etymology: The specific name coccyginus is derived from
the Greek koxkvytvos, and refers to the purple color.

Discussion: Cenodagreutes coccyginus is most easily dis-
tinguished externally from C. aethus by its darker overall
shell color and by having granules which are finer than
the pustules of C’. aethus covering the depression between
the axial ribs. Internally, the major differences lie in the
alimentary tract and the reproductive system. In C. aethus
as contrasted to C. coccyginus the projections along the
floor of the rhynchodaeum are absent, the glandular
sheath is long, a cuticular epithelium lines the anterior
part of the oesophagus and a long, thin anal gland lies
along the intestine near the anus. The reproductive system
of C. aethus differs from that of C. coccyginus by having
a pallial oviduct along with an ingesting gland, a ventral
channel in the capsule gland and a large anteriorly placed
sperm sac which is part of the bursa copulatrix.

LITERATURE CITED

SmitH, EpmMuNp Hosart
1967. | The proboscis and oesophagus of some British turrids.
Trans. Roy. Soc. Edin., in press
THIELE, JOHANNES

1929. | Handbuch der systematischen Weichtierkunde. _Jena,
Gustav Fischer, 1929 - 1935; 1154 pp.; 893 text figs.

Vol. 10; No. 1

Page 5

THE VELIGER

The Radula of Zierliana woldemarit (KiENeR, 1839)

JEAN M. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(3 Text figures)

THE GENUS Zierliana Gray, 1847 is quite different mor-
phologically from most other mitrid forms, except for the
presence of typical columellar folds. Some of its species
have such un-mitrid characters as prominent denticles
on the labrum, a distinct posterior notch or canal, and in
some cases — as in the type species, for example - a
clearly atypical shape for a mitrid. Some of the species
have whorls that are smooth as in Mitrinae [Zierliana
anthracina (REEVE, 1844), ? Z. quoyi (DESHAYEs, 1844) ],
or weakly spirally ribbed [Z. woldemari (KizNER, 1839) ].
At least one form [Z. aethiops (REEVE, 1845) ] has vexillid
axial ribbing on the body whorl, while another has a
combination of both axial and spiral ribs [Z. ziervogeliana
(GmeELIn, 1791) ]. All of these factors have led to a certain
amount of conjecture as to the correct systematic position
of the genus.

For some time I had tried to secure preserved speci-
mens for further study of the radula and soft parts, but
without success. There are only a few species in the genus,
probably not more than six or seven; most of these seem
relatively rare in collections and are seldom scen pre-
served with animal intact. Recently my mention of this

Figure 1

Zierliana woldemarii (KIENER, 1839)

taxonomic problem led Mrs. Virginia Orr Maes of the
Academy of Natural Sciences of Philadelphia to extract
and mount the radula of a specimen of Zierliana wolde-
maru from the Academy’s collection (Text figure 1). This

Figure 2

Zierliana ziervogeliana (GMELIN, 1791), type species

species is quite similar in form to the type species, Z. zier-
vogeliana (Text figure 2), and it can probably be safely
assumed that the radulae of both species would be rela-
tively similar also. Mrs. Maes sent me the slide she had
prepared, together with the two halves of the shell from
which the radula had been extracted. I am grateful to
her for her quick and practical response to my casual
mention of the problem.

As may readily be seen from the drawing (Text figure
3), the radula of Zierliana woldemarii is definitely of
vexillid type, with a multicuspid rachidian and sickle-
shaped lateral teeth, similar to the radulae of Vexillum
cadaverosum (REEVE, 1844), V. exasperatum (GMELIN,
1791) and V. semifasciatum (Lamarck, 1811). However,
it is proportionately a good deal smaller, and possesses
rounded rachidians, whereas the cusps of the other species
are sharply pointed, as illustrated by CERNOHORSKY
(1966, p. 119).

Page 6 THE VELIGER Vol. 10; No. 1

It would seem, therefore, that this evidence supports in the Vexillinae. Its morphological characters are distinct
the previously tentative placement of the genus Zierliana enough from other vexillid forms, however, to warrant
retaining Zierliana as a full genus apart from those forms

having closely similar raduiae.

The accompanying text figures were drawn by Mrs.
Sie Emily Reid of the Veliger staff.
LITERATURE CITED
el

50 p CERNOHORSKY, WALTER OLIVER
1966. A study of mitrid radulae and a tentative generic ar-
rangement of the family Mitridae (Mollusca: Gastropoda).

The Veliger 9 (2) ) : 101 - 126; 47 text figs. (1 October 1966)

Kiener, Louis Cuarzes
Zierliana woldemarii (K1ENER, 1839) 1839.

Figure 3

Spécies général et iconographie des coquilles vivantes;
Radula, Preparation by V. O. Maes, ANSP. Mitra. Rousseau, Paris; 4: 1 - 120; plts. 1 - 34

Vol. 10; No. 1

THE VELIGER

Page 7

A New Terebrid Species
with Check List of Terebridae from the Red Sea

(Mollusca : Gastropoda )

TWILA BRATCHER'

AND

R. D. BURCH’

(Plate 2)

Since 1962 THE JUNIOR AUTHOR has been studying the
terebrid fauna of the Gulf of Akabar on the Red Sea
with the cooperation of Mr. D. C. Insall, an ardent local
collector. The check list here included and the description
of a new species are the result of that study. The majority
of the specimens examined was taken by shore collecting
and diving. A few additional specimens were collected
from the dredged sea bottom when the Port of Eilat,
Israel was being constructed.

Eilat, on the Gulf of Akabar (listed on many maps
as the Gulf of Aqaba) is located at 29° 32’20” North
and 34°57’00” East. This is an area where the desert
sands and the water of the gulf mect, and there is little,
if any, rise and fall of tide. The fauna examined is entirely
Indo-Pacific and is at the northern end of its range.
Nothing seems to have been published on the Recent
terebrid species of the Red Sea.

Specimens of each species listed here remain in the
collection of Mr. Insall, and it is our understanding that
his collection will be open to students of conchology.
Arrangements for examining the collection may be made
by writing Mr. D.C. Insall at PO. Box 3079, Haifa,
Israel.

The following list is not presumed to be all-inclusive,
but it does contain all the species of Terebra encountered
by Mr. Insall and included in his own collections and in
those of other collectors of his area.

‘8121 Mulholland Terrace, Hollywood, California 90046
2 P.O. Box 133, Downey, California 90241

Terebra BRUGUIERE, 1789
(Triplostephanus) Da.x, 1908

Terebra (Triplostephanus) insalli BRATCHER & BuRCH,

spec. nov.

(Plate 2, Figures 1 to 3)

Shell: Medium sized, pale beige, slender and elongated
with 21 narrow, slightly concave whorls divided by a
convex sutural band of white. A subsutural band of pearly
tubercles joins the sutural band. Balance of the whorl
sculptured by transverse lines adjacent to the sutural and
subsutural bands plus two additional weaker spiral lines
crossed by axial lines more numerous than the tubercles
on the subsutural band. This gives the effect of three
corded bands of small almost squared, somewhat flattened
nodes, the posterior being the most prominent. Sculpture
remarkably consistent from the early whorls to the body
whorl on which the third row from the subsutural band
becomes a row of protruding nodes at the periphery. These
spiral lines are broken by somewhat obsolete axial lines.
Body whorl short. Aperture small, ovate, ending in a short
recurved canal. Columella short, recurved, with one
microscopic plication, laminated. Length 59.9mm, width
8.0mm.

Holotype: California Academy of Sciences, Department
of Geology, Type collection no. 12946 (Plate 2, Figures
1 and 3).

Paratypes: Paratype no. 1: R.D. Burch coll. no. 589;
Length 59.1mm, Width 8.0mm; nucleus missing.

Page 8 THE VELIGER

Paratype no. 2: T. Bratcher coll. no. 30001; L. 30.9mm,
W. 4.9 mm; nucleus intact (Plate 2, Figure 2).

Paratype no. 3: Insall coll. no. 7/11G; L. 59.8mm, W.
7.6mm; nucleus missing.

Paratype no. 4: Insall coll. no. 7/12; L. 82.3mm; W.
8.8mm; apex missing.

Paratype no. 5: Conchological collection, Stanford Uni-
versity; L. 50.2mm, W. 6.9mm; apex missing.
Paratype no. 6: Museum of Comparative Zoology, Har-
vard University; L. 50.3mm, W. 7.1 mm; half of nucleus
missing.

Paratype no. 7: Santa Barbara Museum of Natural His-
tory coll. no. 23729; L. 50.8mm, W. 7.4mm; apex missing.
Paratype no. 8: British Museum (Natural History) col-
lection; L. 52.4mm, W. 7.9mm; apex missing.

Type Locality: All specimens were collected on the Gulf
of Akabar, Red Sea, 29° 32’20” N, 34° 57’00” E. Only
two specimens of the type lot were live taken. Paratype no.
3 was collected at South Beach in 10 feet of water on
sand bottom, and Paratype no. 7 was taken at Coral Beach
in 6 m of water, both by divers.

DISCUSSION

This species may be separated easily from the three to
which it bears greatest resemblance: Terebra cumingit
DesuHayEs, 1857 (Plate 2, Figure 4), T: triseriata Gray,
1834 (Plate 2, Figure 5), and T: jennings R.D. Burcu,
1965 (Plate 2, Figure 6). The whorls are more numerous
and much narrower than in T: cumingi but with about the
same apical angle. Also the sculpture of T- insalli is coarser
and with fewer spiral striae than in that of 7: cumingit.
The sculpture of T: insalli, though similar to that of T.
triseriata, has a more beaded look between the sutural
bands. The whorls of T: insalli are about the same width
as those of T: trisertata. Terebra triseriata is a more slender
shell with a more acute apical angle. There is less resem-
blance to T: jennings: which has quite smooth sculpture,
wider whorls, and a longer, less recurved canal. Some of

Vol. 10; No. 1

the paratypes exhibit a double row of incised spiral lines
between the first and second row of nodulated cords
posterior to the suture.

For the sake of expediency the following subgenera are
provisionally accepted. Much work on the generic struc-
ture of this family is needed.

Genus:

Terebra BrucutéRE, 1789. Encycl. Méth., Vers, vol. 1,plt.

xv. Type species: Buccinum subulatum LINNAEUS,
1767.

Subgenera:

(Abretiella) Bartscu, 1923, Nautilus, vol. 37 (2): 61
to 63. Type species: Terebra cerithina LaMaARcK, 1822.

(Decorthastula) Oyama, 1961, Venus, vol. 21 (2): 185.
Type species: Terebra affinis Gray, 1834.

(Dimidiacus) IREDALE, 1929, Austral. Zool., vol. 5: 341.
Type species: Terebra cingulifera LaMarcK, 1822.

(Oxymeris) Dati, 1903, Proc. U.S. Nat. Mus., vol.
26: 951. Type species: Buccinum maculatum Lin-
NAEUS, 1758.

(Perirhoe) Dati, 1908, Nautilus, vol. 21: 124. Type
species: Terebra circumcincta DesHayEs, 1857.

(Strioterebrum) Sacco, 1891, Moll. terr. Terz. Pied.
e Lig. pt. 10, p. 33. Type species: Terebra basteroti
Nyst, 1843.

(Subula) ScHuMACcHER, 1817, Ess. Nouv. Syst., p. 233.
Type species: Buccinum dimidiatum LinNnaEus, 1758

(Triplostephanus) Dau, 1908, Nautilus, vol. 21: 124.
Type species: Terebra triseriata Gray, 1834.

Genus:

Hastula H. & A.ApaMs, 1853. Gen. Rec. moll., vol. 1: 225.
Type species: Buccinum strigilatum LinNAEus, 1758.

Subgenus:

(Hastulina) Oyama, 1961, Venus, vol. 21 (2): 183-184.
Type species: Terebra casta Hinps, 1843.

Explanation of Plate 2

Figure 1: Terebra insalli BRATCHER & Burcu, spec. nov. Holotype,
C.A.S. No. 12946 (x14)

Figure 2: Terebra insalli. Paratype No. 2. Bratcher collection (x 44)
Figure 3: Same shell as Figure 1, body whorl (x 2.8)
Figure 4: Terebra cumingii DEsHayEs, 1857. Body whorl. Burch
collection (x 2.8)

Figure 5: Terebra triseriata Gray, 1834. Body whorl. Burch
collection (x 2.8)

Figure 6: Terebra jenningst BurcH, 1965. Paratype No. 34. Body
whorl. Burch collection (x 2.8)

TueE VELIGER, Vol. 10, No. 1 [BRATCHER & Burcu] Plate 2

Figure 2 Figure 3

Figure 5 Figure 6

photographs by Twita BratcHER

Vol. 10; No. 1

A Check List of Terebridae
from the Gulf of Akabar, Red Sea

Terebra (Decorthastula) affinis Gray, 1844. Proc. Zool.
Soc. London f. 1843: 60; fig. Hips in Sowersy, Thes.
Conch., 1844, plt. 44, fig. 78. (non Turton, 1832).

T. (Decorihastula) columellaris Hinvs, 1844. Proc. Zool.
Soc. London f. 1843: 151; fig. Htnps in Sowersy,
Thes. Conch. 1844, plt. 44, fig. 77.

T. (Decorihastula) flavofasciata Pitspry, 1921, Proc.
Acad. Nat. Sci. Philadelphia 69: 306; plt. 2, fig. 3.

T. (Decorithastula) nebulosa SowrErsy, 1825, Tank. Cat.
append., p. 25 (non KieneEr, 1839; non Lorots, 1859).

T. (Triplostephanus) insalli BRATCHER & BURGH, spec. nov.

T. (Dimidiacus) albomarginata DrsHayrs, 1859. Proc.
Zool. Soc. London 1859: 314, no. 212; fig. REEVE, Conch.
Icon. 1860, plt. 15, fig. 65.

T. (Dimidiacus) amanda Hinps, 1844. Proc. Zool. Soc.
London, 1843: 154; fig. Hinps in Sowersy, 1844, p.
166, no. 46; plt. 45, fig. 100.

T. (Dimidiacus) consobrina Desuayers, 1857. Journ. Con-
chyl. 6: 72; plt. 3, fig. 3.

T. (Subula) areolata (Linx, 1806). Beschr. Nat. Samml.

Univ. Rostock, p. 128; fig. CHEMNitTz, Conch. Cab.,
1780 (4), plt. 153, fig. 1441 and plt. 154, fig. 1443.
(non ADAMS & REEVE, 1850).

T. (Subula) argus Hinps, 1844. Proc. Zool. Soc. London
f. 1843: 160; fig. Hinps in Sowersy, Thes. Conch.
1844, plt. 23, fig. 64.

T. (Subula) dimidiata (Linnaeus, 1758), Syst. Nat., ed.
10, p. 741, no. 420; fig. KizeNErR, Icon. Coq. Viv., 1839,
plt. 2, figs. 2, 2a.

T. (Perirhoe) babylonia Lamarck, 1822. Anim. s. Vert.
7: 287, no. 9; fig. KiENER, Icon. Coq. Viv., 1839, plt. 14
(in part), fig. 35 only.

THE VELIGER

Page 9

T. (Abretiella) cerithina Lamarck, 1822, Anim. s. Vert.
7: 288, no. 15; fig. Kiener, Icon. Cog. Viv., 1839, plt.
NN, ee, ZB)

T. (Oxymeris) crenulata (LinnakEus, 1758), Syst. Nat.,
ed. 10, p. 741, no. 416; fig. KizNER, Icon. Coq. Viv.,
1839, plt. 5, figs. 9, 9a.

T. (Oxymeris) maculata (LINNAEUS, 1758), Syst. Nat.,
ed. 10, p. 741, no. 415; fig. Krener, Icon. Cog. Viv.,
1839, plt. 1, fig. 1.

T. (Strioterebrum) textilis Hinps, 1844. Proc. Zool. Soc.
London f. 1843: 156; fig. Hinps in Sowersy, Thes.
Conch., 1844, plt. 44, fig. 73.

Hastula (Hastulina) casta (Hinps, 1844), Proc. Zool.
Soc. London f. 1843: 156; fig. Hinps in Sowersy, Thes.
Conch. 1844, plt. 44, fig. 84.

ACKNOWLEDGMENT

We wish to express our thanks and appreciation to Mr.

D. C. Insall for his generous cooperation in sending speci-

mens for this study and for donating type specimens for

distribution.

LITERATURE CITED

Burcu, Ropert DonaLp
1965. — New terebrid species from the Indo-Pacific Ocean and
from the Gulf of Mexico, with new locality records and pro-
visional list of species collected in western Australia and at
Sabah, Malaysia. The Veliger 7 (4): 241-253; plt. 31; 6
tables (1 April 1965)
Desuayes, GERARD PAUL
1857. Descriptions d’espéces nouvelles du genre Terebra.
Journ. de Conchyl. 6: 65 - 102; plts. 3-5
Gray, JoHN Epwarp
1834. Untitled. [7érebra] Proc Zool. Soc. London, Part II
(1834) : 50 - 63 (50 - 56: 26 Sept.; 57-63: 25 Nov. 1834)

Page 10

THE VELIGER Vol. 10; No. 1

The Retention of Lamellibranch Larvae in the Niantic Estuary’

BY
JOHNES K. MOORE?’
AND

NELSON MARSHALL

Graduate School of Oceanography
University of Rhode Island, Kingston, Rhode Island 02881

(1 Map)

MarSHALL & WHEELER (1965) found that phytoplankton
were most abundant in the inner stretches of the Niantic
estuary; 7. €., on the upriver side of the shoals that set off
the estuarine basin from the outer Bay (see Map). Phyto-
plankton concentrations, somewhat less than those occur-
ring in the basin and commonly different in composition,
were also found in the Bay, while numbers generally
decreased over the shoals. Conditions favoring reproduc-
tion in the inner estuary, particularly for the dinoflagellates
so numerous there, plus the general lack of dispersal from
this semi-enclosed area may account for much of the
abundance observed in the basin area.

Data gathered by the senior author on the dispersal
of common planktonic lamellibranch larvae show little or
nothing when examined for correlations with data on
light, depth, salinity and stage of tide. However, on noting
the distributions of each species on early dates of the
summer spawning season, it is clear that the larvae are
consistently most abundant in the basin and upriver sta-
tions in spite of extreme irregularities in other respects
(Table 1). In this way the larval distributions grossly
parallel those of the phytoplankton. For these planktonic
larvae it seems unlikely that this distribution is the direct
result of a concentrated spawning activity in the inner
estuary. The spawning bay scallops, Aequipecten irradians,
are concentrated on the shoals. The spawning oysters,
Crassostrea virginica, are scattered on the intertidal rocks
throughout the estuary. The shipworm, Teredo, is ubi-
quitous but may be most abundant just inshore from the

‘From work partially supported by Contract AT (30-1) 2678 with
the Atomic Energy Commission and by the National Science
Foundation.

2 Present address: Salem State College, Salem, Massachusetts 01970

inlet where there are many docks and pilings. The minute
pelecypod identified as Mysella (Rochefortia) planulata
is thought to be ubiquitous, judging from observations
of PHELps (1964) on a nearby estuary.

MarsHALL & WHEELER (1965) suggested that there
might be a differential tidal effect with the flood being
more effective than the ebb in the transport of phyto-
plankton across the shoals. This may be even more signifi-
cant in the distribution of the planktonic larvae observed
and perhaps for holoplankton as well. At the beginning
of the flood less than a foot of water covers the shoals. On
the flooding tide, waters of relatively high salinities come
in from the Bay, cross the shoals and apparently move up
estuary along the bottom. With the ebb, surface waters
from the basin cross the shoals and move seaward but may
tend to remain near the surface. Waters in the Bay, semi-
enclosed by headlands, are not immediately swept from
the area, so these surface waters may return on the follow-
ing flood. This simple effect should carry plankton well
into the estuary and tend to keep them there. It would
tend to be operative irrespective of the vertical movements
of the plankton unless they were strongly grouped toward
the surface.

The flushing of the estuary tends to counter the hydro-
graphic effect just described. Using runoff data from U.
S. Geological Survey Water supply records, it is calculated,
with the method of KetcHum (1951), that 25 days are
required for water entering from the tributaries to reach
the Bay during times of low runoff such as characterize
the spawning period. This does not seem strong enough
to counter a tidal mechanism but it may be sufficient
to account for the numbers of larvae in the Bay late
in the summer.

Vol. 10; No. 1 THE VELIGER Page 11

Table 1

The number of lamellibranch larvae per m? as sampled at four stations along the axis of the Niantic estuary
during the Summer 1963. See Map for station locations’.

Mysella Teredo Crassostrea Aequipecten
planulata navalis virginica trradians

(Stimpson, 1851) Linnaeus, 1758 (GMELIN, 1791) (Lamarck, 1819)

I ov g ao} g 4 gv g

> Sibaaenenl| aS eae] > Bs AG Bets

Date a wa OO nig) a a 21a wa eS
10 June 4 422 2 39 0 7 0 0
14 June 6 0 710 6 O62 0 0 1 0 0 7
21 June 2 232 650 6 0 18 0 0 0 0 0 2
1 July 4 32 1416 24 Oneao2 6 4 9 2 4 6916
8 July 2 88 2222 6 12S 0 0 0 11 0 0 3
15 July 54 124 9542 10 0 39 0 0 6 0 0 9
22 July 6 440 2218 56 60 39 4 4 6 0 4 22
29 July 154 304 764 22 180 56 4 20 15 2 0 21
6 August 160 112 244 143 16 10 12 16 23 0 8 0
12 August 84 148 690 42 COM al 32 2, 169 0 32 31
15 August 36 52 472 46 64 74 20 56 188 14 20 59
26 August 22 0 95 74 4 39 4 0 4 8 0 546
3 September | 58 24 31 64 12 33 16 0 5 0 0 17
18 September | 10 4 IS 0 5 0 0 0 0 0 0

3 Samples from both the surface and off the bottom were taken at the Bay, Basin and Upriver stations. Since
differences did not follow significant patterns they are averaged in this presentation. For the same reason Basin
and Upriver data are averaged.

41°22"

Page 12

67 ).
\ ey): YardsO 500
‘! \ Bi ie
oh \\y 4 Km. O 0.5
“AV
Up River \\y/)
oni Ke
ey SN
: “| 4
| 3
el EN
} :
e((t =
“i ch)
NS o
oD NS c
a
E
a
> r)
“4 ©
oes gS
in a eaey, 2
(See USCGS Chart No.214) 137 // &
JF :
ae :
ett \\ a
ee Lp
es EON =
iy Po aA E
oe 7 12° / — ©
Df SASS 5
Se Bay ~\ a
19’ ny Z \
a aes \4
Oo NIANTIC BAY ~
atl CG
NYY
“pol
n/N
“4 ( '
ont SS
a \ J i
Oy, 7 Ve
revl2 TT 10'

THE VELIGER

Vol. 10; No. 1

Table 2
2-7 values as observed along the axis of the Niantic
estuary during the lamellibranch larvae sampling,
Summer 1963. For each date and station the upper
reading is for the upper depth, the lower reading for the
lower depths. See Map for station locations and

41°22" approximate total depths.
Station

Date Bay Shoals Basin Upriver
14 June 21.528 19.738 16.505 16.209
22.279 21523 21.257
21 June 21.282 21.705 20.173 18.699
22.052 Ae syil 21.489
1 July 20.063 18973 18.074 18.166
21.408 20.317 20.477
el 8 July 21.224 «21.304 | 20} aT toatl
21.900 21.032 21.185
15 July 20.888 20.256 20.211 19.346
21.604 20.856 20.932
22 July 21.528 21.509 19.176 18.138
21.922 21.039 21.003
29 July 20.292 19.106 18.895 17.941
21.429 20.306 20.429
6 Aug 21.022 20.628 19.500 18.672
20' 21.446 20.241 20.358
12 Aug 20.632 19.837 20.269 19.748
21.560 20.712 20.844
15 Aug 21.355 20.349 20.517 19.724
21.672 21.079 21.022
26 Aug 21.782 21.011 20.207 20.308
22.044 20.462 21.017
3 Sept 21.464 21.494 21.042 20.567
22.643 21.384 21.451
i9’ 18 Sept 22.301 22.272 21.833 21.611
22.458 22.211 22.254

LITERATURE CITED

KetcuuM, B. H.
1951. | The exchanges of fresh and salt waters in tidal estuaries.
Journ. Mar. Res. 10: 18 - 38
MarsHALL, NELSON & BERNICE M. WHEELER
1965. Role of the coastal and upper estuarine waters contrib-
uting phytoplankton to the shoals of the Niantic estuary.
Ecology 46: 665 - 673
PueE tps, D. K.
1964. Functional relationships of benthos in a coastal lagoon.
Ph. D. thesis, Univ. Rhode Island Library. -

Vol. 10; No. 1

THE VELIGER

Page 13

The Cowries of the Ryukyu Islands

CRAWFORD N. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plate 3; 2 Maps)

INTRODUCTION

IN ANY DISCUSSION of the Cypraeidae of the Ryukyu
Islands it would seem well to include a brief description
of this semi-remote, curving chain of approximately 70
islands, for the area comprises roughly a 900 mile long
continuous link between the Philippine and Japanese
faunal regions.

These Islands have been known by various names. The
Japanese refer to them as the Ryukyu Retto; they also
are known as the Luchu or Loochoo Group; but, perhaps
more correctly, they should be referred to as the Nansei
or Ryukyu Islands. They form a natural broken land arc
linking the Japanese island of Kyushu with Taiwan (form-
erly Formosa). In these islands are three important sub-
divisions: the Amami, the Sakishima, and the Okinawa
groups. Altogether they represent an exposed land mass
of approximately 847 square miles.

The sovereignty of the Ryukyus has changed many
times. From a Chinese protectorate in 1372, successively,
this island group became subsidiary to both China and
Japan about 1451, eventually falling deeper under Japa-
nese influence in 1609, then finally coming under full
control of that country in 1879. As a result of World War
II the island group became subject in 1945 to the United
States military government. Civil government was returned
to the inhabitants in 1951; in 1953 the Amami Group was
returned to Japan; custody of Okinawa, the largest island,
remains still with the United States.

Most of the field work upon which this report is based
was carried out on Okinawa, which is about 65 miles
long and approximately 3 to 10 miles wide, forming an
important land barrier between the comparatively shallow
East China Sea and the deeper Philippine Sea. The
island chain lies in a northeast-southwest attitude, approx-
imately 26° 30’ North Longitude and 128° 00’ East Lati-
tude.

I have examined many specimens of the numerous
cowrie species from this area and can find very little, if

any, differences in color and form as compared with the
same species in the Philippine fauna. The shells are indeed
strikingly similar in nearly every respect. Perhaps the most
obvious differences are that some species are less abundant
in the Ryukyus and the depth ranges seem to be more
shallow in the northern islands.

Despite its proximity to the Japanese Archipelago the
island chain remains a separate ecological entity, and
appears to be a transitional area for many of the east
Asian cypraeids. Species that are quite common in the
Philippines appear here as scarce or decidedly uncommon,
even disappearing from the fauna; others appearing only
infrequently, then becoming more abundant in the more
northern Japanese islands. It should be mentioned that in
the northern Japanese islands new species have been dis-
covered fairly recently [such as Schilderia langfordi (Ku-
RoODA, 1938), S. teramachu (Kuropa, 1938), and Erosaria
guttata azumat ScHILDER, 1960] some of which may
eventually be found in Ryukyu waters.

With a study of the Philippine cowries recently con-
cluded, I am impressed by how closely the shells of the
two regions appear to resemble one another morpholog-
ically. For this reason I have referred for illustrations to
appropriate figures in some of my earlier papers such as
CaTeE, 1965, 1966. The reader is also referred to the latter
paper for a comparison of certain similar species, such as
Bistolida pallidula, B. interrupta; Pustularia cicercula, P
bistrinotata mediocris, P. tetsuakit, P globulus; Bistolida
kieneri depriesteri, B. hirundo neglecta, B. ursellus.

This study, which has been underway more or less
interruptedly for nearly 15 years, will furnish collectors
with an authentic modern list of the Ryukyu Cypraeidae.
It cannot be considered the final word about cowries in
these waters, however, but only the beginning, a basis for
future field work and discovery. Listed here are all the
species presently known to be living in these waters, all
recently collected and substantiated, and now on deposit
in various Okinawan collections or in that of the author.

TAIWAN:

e

YAEYAMA
ISLANDS ,

0;
‘ ISHIGAKI
© : A
IRIOMOTE at ISEAWE)
ISLAND— ~~.
ob

THE VELIGER Vol. 10; No. 1

SEA 2 KIKAISHIMA

ISLAND

AMAMI
ISLANDS

OKINAWAS 2oq.ci7
» e

Lo IOI

oe et PACIFIC
OCEAN

all

©
[7]

Qa

Vol. 10; No. 1

THE VELIGER

127°40' 127°50'

EAST CHINA SEA

MOTOBU

SAKIYAMA

128°10'

BAY

1E-MURA
HEDO PEN.
26°40'
MOTOBU
&> PEN.
IMBU (OLD)
IMBU (NEW)
SMUGGLER'S COVE SE
BeosG! OTABARUA SI Ye SERAGAKI \.
nye SERAGAKI 26°30
SITE 2, HIGHWAY I Sx ONNA (COVE 4)
MINAMI
PACIFIC
PUNCH BOWL N
ME TASAKI wine OCEAN
ZAMPA-MISAKI—@yesMAEDA PT.o—, ISHIZA
BOLO POINT——*>¢x
(SOUTH) Aa
BOLO POINT
(NAGAHAMA) 2
AGGURA Ct IKE! ISLAND
TORI STATIONS: HENZA -,
Se SANS) AUS shite Beet) MIYAGI ISLAND
we b
<@: KADENA Nyt
26°20" yHENZA ISLAND 090!
YABUCHI : }——— 262520
ISLAND gag’ i) MIRUCHIBISHI
; ; REEF
KUE %. , HIGA ISLAND
SUKIRAN }% & \ ares) O K | N AWA
w A Be winami-ukiparu \.
VWsweee
WHITE BEACH Fee al SAGO
x £0; | Ie
MACHINATO 1
BUCKNER ES TSUKEN ISLAND
BAY “(OFF ISLAND) |
MILES
YONABARU Ee aS Se
ee, ) 5 10
a 26°10
GUSHICHAN
j PHILIPPINE SEA
[| SW/kina REEF 127°50! 128° 128°10 128°20'

Page 16

THE VELIGER

Vol. 10; No..1

ECOLOGICAL NOTES

Because of the lack of convenient access to many other
islands, collections and reports have been, for the most
part, confined to the island of Okinawa, with only an
occasional reference to the other Ryukyu islands. Like
cowries from most localities, they are found living in coral,
under rocks, on sandy flats, and in muddy exposures,
usually here, however, in from 4 to 10 feet of water.
The distribution for each species in different localities is
noted; in few instances have they been of common
occurrence.

So that present knowledge of the cowrie populations
may not be lost, it is important to record the known
species and their distribution in these islands; especially
is this true at Okinawa. Great changes are taking place
in many of the intertidal and adjacent marine areas
because of extensive dredging operations, many sewage
systems emptying into the critical offshore waters, and
shoreline land fills that are destroying large sections of
the natural habitat; further, large quantities of sand and
coral are being removed to be made into building material
— thereby eliminating sand flats and underwater habitats;
and finally, and far from the least, native Okinawans
have become conscious of the commercial value of their
mollusk fauna, selling the shells to the military personnel.
They collect indiscriminately, unintentionally destroying
habitats and most of the sea life within their reach.

It would therefore seem of interest to mention the
present physical appearance of some of the most important
collecting areas (see Maps): — 1/ Machinato is a reef,
most accessible at a minus tide; -— 2/ Kue is an area
of sandy flats and coral tide pools protected from turbu-
lent outer water by reefs that are exposed at low tide;
— 3/ Kadena is noted for its coral reef collecting; —
4/ Zampa-Misaki is the general collecting area at the
end of the Zampa peninsula. Land’s end is known as
Bolo Point, and northeast of the point is the Punch Bowl,
an area that is exposed at a good minus tide. It should
be noted that the entire Zampa region has a very rough
sea exposure. — 5/ Onna has important reefs on which
Cypraea can be found; there are two main collecting
areas there, one just south of town, the other to the north.
Still further north, Otabaru, Smuggler’s Cove, New and
Old Imbu (Inbu) are successively important collecting
stations. — 6/ Yagaji Island used to be an excellent area
for collecting C'ypraea, especially near the bridge that
approaches from Makiya, but a tidal wave of great magni-
tude a few years ago destroyed the original bridge and
disrupted much of the shoreline; since then the molluscan
fauna has not returned. — 7/ The northeastern coastline
from Takae to Oku is very rocky and mostly inaccessible.

Hedo, at the north end of the peninsula of the same
name has been the scene of very limited collecting; the
most outstanding species from there is Ovatipsa chinensis
chinensis (GMELIN, 1791). -— 8/ Off the east-central
coast of Okinawa is an old Coast Guard station on Ikei
Island; nearby are the conchologically important Henza
and Miyagi Islands. Between them impressive sand flats
are exposed at low tide, and likewise a rocky reef becomes
accessible on the east side of Miyagi. It is in this rocky area
that the only specimen of Adusta onyx onyx (LINNAEUS,
1758) was found. - 9/ Baten was at one time a very
productive collecting station. To this day large quantities
of dead shells litter the area everywhere, possibly evidence
of the extensive dredging operations that have taken
place since World War II. There remains, however, now
a fine, flat expanse of mud, with the fauna there unknown
at present. — 10/ Further south from here are Chinen
and Gushichan where many species of Cypraea can be
found. These are important habitats, and consist of sand,
rock, and a limited reef area. Since the southern third of
Okinawa is the most densely populated part of the island,
it is to be expected that the shore in this area will be
most heavily worked over. This is especially true because
of the local concentration of military personnel; in their
effort to find relaxation they go to the beach to picnic
and collect shells, and consequently the tidal areas are
denuded of most molluscan life; however, just beyond the
intertidal zone, in deeper water, many cowries can still
be found.

EARLIER LITERATURE
ON THE AREA

Kuropa (1960) appears to have contributed the first
checklist of mollusks of the Okinawa Islands. In addition
to members of other families listed, he recorded 63 species
of Cypraea (l.c., pp. 21-23; plt. 3), including two new
species, Notadusta katsuae and Bistolida luchuana. Of
those mentioned 57 appear to be valid, some seem mis-
identified and others species unverified in these waters.
It is the purpose of this paper to list only those valid
species we have been able to substantiate as currently
living in Ryukyu waters.

The late Lloyd E. Berry of Los Angeles probably was
the originator of this work with the Ryukyu cowries.
In 1952 he received his first specimens from Mrs. Anita
Scott, then of Okinawa, and he started to accumulate
many of the shells used in this study. It was at this
time that I joined Mr. Berry in this work. Later Mr. Berry
died, Mrs. Scott returned to the United States, and, for
lack of additional material, the project was set aside.

Vol. 10; No. 1

However, in more recent years Bernice and Ernest Albert,
Peter Way, and Barbara Keily, all of Okinawa, have
joined in the field work to confirm the localities and to
substantiate the species, thus assisting me in bringing this
study to a close. The decision to ‘close’ here is arbitrary,
for the work can never stop.

During November 1966 I had the opportunity to visit
Okinawa. Through the kindness of the just mentioned
friends I was able to examine their shell collections and
determine the probable population density of a given
species from the quantities noted in each instance, as
well as to collect at several of the areas listed here and
to observe many of the cowrie species in their native
habitats.

ACKNOWLEDGMENTS

The success of such a report as this depends upon the
interest, integrity, enthusiasm, and hard work of many
individuals. I have mentioned the contributions of Lloyd
Berry and Anita Scott; I must now acknowledge those
whose field work has been the final word in the completion
of this paper: Bernice and Ernest Albert, Peter Way and
Barbara Keily - to them all I give heartfelt thanks.
To Emily Reid who drew the excellent maps, and to others
who have helped in many unseen ways I also express my
appreciation. To Jean Cate goes my deep gratitude for
continual encouragement and understanding.

A Summary of Ryukyu Cowrie Species
from Selected Localities

ZAMPA-MISAKI (BOLO POINT)

Mauritia (Leporicypraea) mappa mappa
Mauritia (Arabica) arabica asiatica

Mauritia (Arabica) maculifera

Mauritia (Arabica) scurra indica

Talparia talpa talpa

Cypraea (Lyncina) argus argus

Cypraea (Lyncina) lynx vanelli

Cypraea (Lyncina) vitellus vitellus

Cypraea (Lyncina) carneola carneola

Luria (Basilitrona) isabella rumphu
Monetaria (Ornamentaria) annulus annulus
Erosaria (Ravitrona) labrolineata labrolineata
Erosaria (Ravitrona) cernica ogasawarensis
Erosaria (Ravitrona) helvola helvola
Erosaria (Ravitrona) caputserpentis caputserpentis
Erosaria (Erosaria) poraria scarabaeus
Erosaria (Erosaria) erosa phagedaina

THE VELIGER

Erosaria (Erosaria) miliaris miliaris
Erronea (Erronea) errones errones
Erronea (Erronea) caurica caurica
Erronea (Melicerona) felina pauciguttata
Purpuradusta fimbriata marmorata
Bistolida (Blasicrura) pallidula pallidula
Bistolida (Blasicrura) teres teres

Cribraria (Ovatipsa) chinensis chinensis
Cribraria (Cribraria) cribraria orientalis

HENZA-MIYAGI

Mauritia (Arabica) arabica asiatica
Mauritia (Arabica) scurra indica
Monetaria (Monetaria) moneta rhomboides
Erosaria (Ravitrona) labrolineata labrolineata
Erosaria (Ravitrona) helvola helvola
Erosaria (Ravitrona) caputserpentis caputserpentis
Erosaria (Erosaria) erosa phagedaina
Erronea (Adusta) onyx onyx

Erronea (Erronea) errones errones
Purpuradusta gracilis japonica

Purpuradusta fimbriata marmorata

SMUGGLER’S COVE

Cypraea (Lyncina) carneola carneola
Palmadusta clandestina moniliaris
Bistolida (Blasicrura) teres teres
Bistolida (Bistolida) stolida stolida
Cribraria (Cribraria) cribraria cribraria

CHINEN

Mauritia (Arabica) arabica asiatica

Cypraea (Lyncina) vitellus vitellus

Luria (Basilitrona) isabella rumphu

Pustularia (Pustularia) cicercula cicercula
Pustularia (Ipsa) childreni samurai

Erosaria (Erosaria) erosa phagedaina

Erosaria (Ravitrona) caputserpentis caputserpentis
Staphylaea (Nuclearia) nucleus nucleus

Erronea (Erronea) ovum ovum

ONNA TIDE FLAT

Cypraea (Lyncina) lynx vanelli

Cypraea (Lyncina) vitellus vitellus
Monetaria (Monetaria) moneta rhomboides
Erosaria (Ravitrona) labrolineata labrolineata
Erosaria (Ravitrona) helvola helvola
Erosaria (Erosaria) erosa phagedaina

Page 17

Page 18 THE VELIGER

Staphylaea (Staphylaea) staphylaea staphylaea
Staphylaea (Staphylaea) limacina limacina
Erronea (Erronea) errones errones
Erronea (Erronea) cylindrica cylindrica
Notadusta punctata atomaria

Palmadusta asellus vespacea

Palmadusta clandestina moniliaris
Purpuradusta gracilis japonica
Purpuradusta fimbriata marmorata
Bistolida (Blasicrura) luchuana

Bistolida (Derstolida) hirundo neglecta
Cribraria (Cribraria) cribraria orientalis

METASAKI REEF FLAT

Talparia talpa talpa

Cypraea (Lyncina) argus argus
Cypraea (Lyncina) lynx vanelli
Cypraea (Lyncina) carneola carneola
Luria (Basiltrona) isabella rumphii
Erosaria (Erosaria) poraria scarabaeus
Staphylaea (Nuclearia) nucleus nucleus
Palmadusta asellus vespacea

MINAMI-UKIBARU

Erronea (Melicerona) felina pauciguttata
Purpuradusta gracilis japonica
Bistolida (Derstolida) kieneri depriesteri

MACHINATO REEF

Mauritia (Arabica) arabica asiatica

Cypraea (Lyncina) carneola carneola

Erosaria (Ravitrona) caputserpentis caputserpentis
Erosaria (Erosaria) erosa phagedaina

Staphylaea (Staphylaea) staphylaea staphylaea
Erronea (Erronea) errones errones

OTABARU REEF

Maurnitia (Arabica) eglantina couturieri

Erosaria (Ravitrona) caputserpentis caputserpentis
Staphylaea (Staphylaea) limacina limacina
Palmadusta clandestina moniliaris

Cribraria (Cribraria) cribraria orientalis

GUSHICHAN

Talparia talpa talpa
Luria (Basilitrona) isabella rumphi

Vol. 10; No. 1

Pustularia (Pustularia) cicercula cicercula
Staphylaea (Staphylaea) staphylaea staphylaea
Staphylaea (Nuclearia) nucleus nucleus

MIYAKO REEF

Mauritia (Leporicypraea) mappa mappa
Mauritia (Arabica) eglantina couturieri
Cypraea (Cypraea) tigris pardalis
Erronea (Erronea) cylindrica cylindrica

Locality Index

1. Ada 45. Miyako Island
3. Atsuta 47. Motobu (north)
7. Baten 48. Motobu (south)
7a. Bise 49. Nago
9. Bolo Point (South) 51. Naha
9a. Bolo Point 51a. Nakaoshi
(Nagahama) 52. Off Island
11. Buckner Bay 53. Okinawa
13. Chinen 55. Okuma
14. Cove 4 (adjacent 57. Onna (Onna Flat)
Seragaki) 57a. Onna (outer reef)
15. Gushichan 59. Otabaru
17. Hedo 61. Punchbowl
19. Henza-Miyagi 62. Ryukyu Islands
20. Henza Reef 62a. Shell Club Island
20a. Ie-Mura 63. Sakiyama Bay
21. Ikei Island 63a. Seragaki Island
23. Imbu (Inbu) -old 63b. Site 2, Highway 1
25. Imbu (Inbu)-new 65. Smuggler’s Cove
27. Iriomote Island 65a. Sony Sand Pit
29. Ishigaki Island 67. Sukiran
31. Ishiza 69. Toguchi
33. Kadena 69a. Tori Station
35. Kikaishima Island 71. Taiwan
35a. Kin Bay 73. Takae
35b. Kina Reef 75. Toyohara
37. Kue 75a. Ueno (Miyako Isld.)
39. Machinato 75b. White Beach
39a. Maeda Point 77. Yabu
41. Metasaki (Maeda saki) 79. Yaeyama Island
42. Minami 81. Yagaji
43. Minami-Ukibaru Isld. 82. Yonabaru
44, Miruchibishi Reef 83. Yontan

85. Zampa-Misaki

Vol. 10; No. 1

THE VELIGER

Page 19

adusta ..... roo PI
amarata$§ .... 24
ATED Gt GN pee ie nl da Sek oe ALR ena els 28
PATETILLU AL ice rte see ere uk ene net 28
annulus 39
arabica ... 37
arenosa § 27
CG GOS cctmetrrerrt meee eee 26
asellus 37
asiatica ... 24
CUDADOTTAD, ciecemabreatoeereaeeeeeenseerereee eps 18, 22, 36
VIRLTTL GLP ee nee ee Mee a 13
SOT ROGOEO. “crmetceeeenreecae erie 13, 20, 29
calxequina .... Beier eeeeeeate A) Wasdo)
caputserpentis . , 18, 21, 32, 38, 39
carneola 38
35
31
30
40
29

GOLEEGE” cepenrtrenncee reer
cernica ..
childrenz ...
chinensis ...
cicercula
cincta § 31
clandestina 37
GOULUTICTIMM eee LOS 23 OF
CHIDTATIA recrtrriieisennnnes 17, 18, 23, 37, 41
GTO O casstecreecenturr ser ere eeeES: 4)
CITI C peer 35
dayritiana . 39
LETC SSC rere ee he as reich cneitisnecaetenett) A
HOYIOSIOTE cecomeaerenanincererarannee 13, 18, 22, 39
Colaniin ampere er 18, 23, 24
CTOSA viene 5 18, 21, 32, 38
errones .... ON 2, ohh, SD
Lin CMe re ete. Me, LB 22 AO
fimbyiatae eens) 17,185.22) 38
flaveolas .... eee aot
LOD ULUSmme etre eee oneal 13, 20, 29
TAGISH eee MAN Ss 1225 38.140

* = synonym;

N = nomen nudum

INDEX or SPECIES

BUOY GN cc tetensd a sane Ream anunen sel aed 24
guttata .......
harmandiana §
elvolay Recenter aa 17, 21, 31
hirundo . 13, 18, 22, 37, 40
indica wu... . 17, 20, 24
LMOGELIGY a) ape ern et ean ace rest ne ene 33
intern ptammrcntrrr reset ser seen LS
WADEUA veces 17, 18, 20, 28, 38
japonica 18, 22, 38
KGS ACen ene eee OM 230
VHOGC commerce Ie, Ie}, 9292, SI)
dra eetoer tee nr a a 20, 27
VA TOVMCAEA ecsressssssrtrsississsesisinsneeie 7, il, Bl
langfordi ... iemmameee S,
Limacin aby eee mre ee nee 18, 21, 33
liste rssubrmamenn ee oon wenahien hand corti
luchuana : 22, 39
Dedric alies Stee lata Ja 210) a eee La pees ara 20
lutea ... mam Cy oul
Lymigcsreran tenes ube es Fane 17, 18, 20, 26, 38
MOAGUIU CTA nnn eee 17, 20, 38
mappa 23
POOUOG cceicecmreoercerseereee 28
marmorata 38
> Ol
mauritiana ... 25
MEALOCTIS roses ; 29
Michaclisseemn emer eee nee we eb,
MUKA oS eee emt nena te Abe ae eae nS
miliaris ... noone lawl
MONELA ooscsescee : 21, 30, 39
moniliaris ....... b 18, 22, 37
NIUUSUINE Cee eee ONES
MANGES IS pos cmct ee iors are ero 26
neglecta ..... 40
nucleus

maturata § .....

SL ENE Simeone

ON Ahearn ean tctialt acc ean tind acces
orientalis ..
OU UTI tae orca neat ctsiatiecentstceets
pallidula
Dard alice

PAUCIZ“ULALA ocerecrnennne 17, 18, 22, 35, 40
phagedaina .
PORATIG Mm
pulchella
punctate ........
punctulata § d
DN TEP OTINUSS Reet as een AR, eer cer isttinse 21
quadrimaculatapemennte ne naae 22, 38
TEGO Ore rec eeeecertmrernrer ne ncereorgneeea OF
rhinoceros ......

TROMBOIMES oevvscccssesssssssssssssrsssssivie
samurai .....

scarabaeus ....
schilderorum
SGUNU A trian ae et en ree

Staphylaca ee
stolida
sulcidentata ....
tabescens $
talpa
teramachii

Lestudinayiguen tt eee
tetsuakit pice ie Seen ee nee ee
tigrts .....
tomlini ..
ursellus .
vanelli
vespacea
victoriana
WILE tE Seer eee tnt eee sre

PAIL AG esta tae te ees Ay teers ee ae

Systematic List of the Ryukyu Cowries

Cy PRAEIDAE FLEMING, 1828 I

Hist. Brit. Anim., 330 (em.) (Edinburgh)

CyPRAEINAE STOLICZKA, 1867

Pal. Ind. (5), 2: 45

Cypraeini ScHiLpEr, 1927

Arch. Naturgesch. 91/A: 92
Mauritia TroscHEL, 1863 5)

Das Gebif§ der Schnecken 1: 205
> Type Species: Cypraea mauritiana Linnaeus, 1758 <

(Leporicypraea) IREDALE, 1930 3.
Mem. Queensld. Mus. 10 (1): 83

> Type species: Cypraea mappa LiNNaEus, 1758 <

Mauritia (Leporicypraea) mappa mappa

(Linnaeus, 1758)
Systema Naturae, ed. 10: 718

(Arabica) JoussEAUME, 1884
Naturaliste 1884: 414

> Type species: Cypraea arabica LINNAEUS, 1758 <

Mauritia (Arabica) eglantina couturieri

(VayssIERE, 1905) )
Journ. Conchyl. 53: 13; plt. 1, fig. 3

Mauritia (Arabica) arabica asiatica
SCHILDER & SCHILDER, 1939

Proc. Malacol. Soc. London 18 (4): 183

Page 20

THE VELIGER

Vol. 10; No. 1

4. Mauritia (Arabica) maculifera ScuiLvErR, 1932
Zool. Anz. 100 (7/8): 165

5. Mauritia (Arabica) scurra indica (GMELIN, 1791)
Systema Naturae, ed. 13: 3412

(Mauritia) TroscHEL, 1863
Das Gebi®& der Schnecken 1: 205

> Type Species: Cypraea mauritiana LinnaEvs, 1758 <
Systema Naturae, ed. 10: 721

6. Mauritia (Mauritia) mauritiana calxequina
(MELVILL & STANDEN, 1899)
Journ. Conchol. 9: 236

Talparia TROSCHEL, 1863
Das GebiB& der Schnecken 1: 204
> Type Species: Cypraea talpa Linnazus, 1758 <
(Talparia) TRoscHEL, 1863
7. Talparia (Talparia) talpa talpa (Linnarus, 1758)
Systema Naturae, ed. 10: 720

Cypraea Linnaeus, 1758

Systema Naturae, ed. 10: 718
(em.) Monrtrort, P. Denys be, 1810

Conchyl. Syst. 2: 630

(Cypraea) Linnaeus, 1758
> Type Species: Cypraea tigris LinNaEus, 1758 <

8. Cypraea (Cypraea) tigris pardalis Suaw, 1795
Vivar. Natur. Misc. 6: plt. 193

(Lyncina) TRoscHEL, 1863
Das GebifS der Schnecken 1: 205
> Type Species: Cypraea lynx Linnaeus, 1758 <

9. Cypraea (Lyncina) argus argus LiINNAEUus, 1758
Systema Naturae, ed. 10: 719

10. Cypraea (Lyncina) lynx vanelli Linnazus, 1758
Systema Naturae, ed. 10: 721

11. Cypraea (Lyncina) vitellus vitellus LInNaEus, 1758
Systema Naturae, ed. 10: 721

12. Cypraea (Lyncina) schilderorum IrEpAe, 1939
Austral. Zool. 9 (3): 303

13. Cypraea (Lyncina) carneola carneola LINNAEUS, 1758
Systema Naturae, ed. 10: 719

14. Cypraea (Lyncina) kuroharai (Kuropa & Habe, 1961)
Col. Illust. Shells of Japan 2: 42; plt. 19, fig. 17

Luriini ScHILDErR, 1932
Fossil. Cat. 1: Animalia, pars 55, Cypraeacea

Chelycypraea SCHILDER, 1927

(Chelycypraea) ScuipER, 1927
Arch. Naturgesch. 91A (10): 92

> Type Species: Cypraca testudinaria LinNaEus, 1758 <

15. Chelycypraea (Chelycypraea) testudinaria testudinaria
(Linnagus, 1758)
Systema Naturae, ed. 10: 719

Luria JoussEAUME, 1884

Bull. Soc. Zool. France 9: 92
(Naturaliste 1884: 414, nom. nud.)

> Type Species: Cypraea lurida Linnaeus, 1758 <
Systema Naturae, ed. 10: 720

(Basilitrona) IrREDALE, 1930
Mem. Queensid. Mus. 10 (1): 83

> Type Species: Cypraea isabella LinnaEus, 1758 <
Systema Naturae, ed. 10: 722

16. Luria (Basilitrona) isabella rumphi
SCHILDER & SCHILDER, 1938
Proc. Malacol. Soc. London 23 (3): 177

Nariini ScHILDER, 1932
Fossil. Cat. 1: Animalia, pars 55, Cypraeacea

(Pustulariini ScHiLpEeR, 1932)
Fossil. Cat. 1: Animalia, pars 55, Cypraeacea

Pustularia Swainson, 1840
Larpner’s Encycl., p. 324

(Annepona) IREDALE, 1935
Austral. Zool. 8 (2): 114

> Type Species: Pustularia mariae ScHILDER, 1927 <

17. Pustularia (Annepona) mariae (ScHILpER, 1927)
Arch. Naturgesch. 91A (10): 104

(Pustularia) Swainson, 1840
Larpner’s Encycl., p. 324

> Type Species: Cypraea cicercula LINNAEus, 1758 <

18. Pustularia (Pustularia) cicercula cicercula
(Linnaeus, 1758)
Systema Naturae, ed. 10: 725
19. Pustularia (Pustularia) bistrinotata mediocris
SCHILDER & SCHILDER, 1938
Proc. Malacol. Soc. London 23 (3): 126

20. Pustularia (Pustularia) globulus globulus
(Linnaeus, 1758) :
Systema Naturae, ed. 10: 725

Vol. 10; No. 1

(Ipsa) JoussEAuME, 1884
Naturaliste 1884: 415
> Type Species: Cypraea childreni Gray, 1825 <

21. Pustularia (Ipsa) childreni samurai ScuitveEr, 1940
Arch. Molluskenk. 72: 42

(Narinii ScHiLper, 1932)
Fossil. Cat. 1: Animalia, pars 55, Cypraeacea

Monetaria TRoscHEL, 1863
Das Gebi® der Schnecken 1: 205

(Ornamentaria) SCHILDER &
ScHILDER, 1936
Proc. Zool. Soc. London 1936: 1120

> Type Species: Cypraea annulus LinnaEus, 1758 <

22. Monetaria (Ornamentaria) annulus annulus
(Linnaeus, 1758)
Systema Naturae, ed. 10: 723

(Monetaria) TroscHEL, 1863
Das Gebi®& der Schnecken 1: 205
> Type Species: Cypraea moneta LinnaEvs, 1758 <

23. Monetaria (Monetaria) moneta rhomboides
SCHILDER & SCHILDER, 1933
Zool. Meded. Leiden 16: 163

Erosaria TROSCHEL, 1863
Das GebiB der Schnecken 1: 205
> Type Species: Cypraea erosa Linnagus, 1758 <

(Ravitrona) IREDALE, 1930
Mem. Queensld. Mus. 10 (1): 82
> Type Species: Cypraea caputserpentis LINNAEUS, 1758 <

24, Erosaria (Ravitrona) labrolineata labrolineata
(Gasxoin, 1849)
Proc. Zool. Soc. London 1849: 97

25. Erosaria (Ravitrona) cernica ogasawarensis
ScHILDER, 1944
Arkiv Zool. 36 A (2): 23

26. Erosaria (Ravitrona) helvola helvola
(LinnaEus, 1758)
Systema Naturae, ed. 10: 724

27. Erosaria (Ravitrona) caputserpentis caputserpentis
(LinnaEus, 1758)
Systema Naturae, ed. 10: 720

(Erosaria) TRoscHEL, 1863
Das GebifS der Schnecken 1: 205
> Type Species: Cypraea erosa Linnaeus, 1758 <

THE VELIGER

Page 21

28. Erosaria (Erosaria) poraria scarabaeus (Bory, 1827)
Encycl. Méth. 3: 164; atlas (VALENCIENNES MS)

78) Erosaria (Erosaria) erosa phagedaina
(MELviILL, 1888)
Mem. Manchest. Lit. Soc. 4 (1): 223; fig. 11

30. Erosaria (Erosaria) miliaris miliaris (GmeEin, 1791)
Systema Naturae, ed. 13: 3420

Staphylaea JouSSEAUME, 1884
Naturaliste 1884: 415

(Staphylaea) JouSSEAUME, 1884
> Type Species: Cypraea staphylaea Linnagus, 1758 <

31. Staphylaea (Staphylaea) staphylaea staphylaea
Systema Naturae, ed. 10: 725

32. Staphylaea (Staphylaea) limacina limacina
(Lamarck, 1810)
Ann. Mus. Nat. Hist. Paris 15: 101

(Nuclearia) JOUSSEAUME, 1884
Bull. Soc. Zool. France 9: 98

> Type Species: Cypraea nucleus Linnaeus, 1758 <

33. Staphylaea (Nuclearia) nucleus nucleus
(Linnagus, 1758)
Systema Naturae, ed. 10: 724

(Erroneini ScuiLpER, 1927)
Arch. Naturgesch. 91/A 10: 109

Erronea TROoSCHEL, 1863
Das GebiB der Schnecken 1: 205

> Type Species: Cypraea errones LinNaEus, 1758 <

(Adusta) JoussEAuME, 1884
Naturaliste 1884: 414
> Type Species: Cypraea adusta Lamarck, 1810 <
Ann. Mus. Hist. Nat. 16: 92

= Cypraea onyx Linnaeus, 1758

34. Erronea (Adusta) onyx onyx (LINNAEUS, 1758)
Systema Naturae, ed. 10: 722

(Gratiadusta) IREDALE, 1930
Mem. Queensld. Mus. 10 (1) : 82
> Type Species: Cypraea pyriformis Gray, 1824 <
Zool. Journ. 1: 371

35. Erronea (Gratiadusta) pulchella pulchella
(Swainson, 1823)
TiLLocn’s Phil. Mag. 61: 376

Page 22

THE VELIGER

Vol. 10; No. 1

(Erronea) TRoSCHEL, 1863
Das GebiB® der Schnecken 1: 205

> Type Species: Cypraea errones LinNAEuS, 1758 <

36. Erronea (Erronea) errones errones (LINNAEUS, 1758)
Systema Naturae, ed. 10: 723

37. Erronea (Erronea) ovum ovum (GmeEun, 1791)
Systema Naturae, ed. 13: 3412

38. Erronea (Erronea) cylindrica cylindrica (Born, 1778)
Index Mus. Caes. Vindob. 1: 169

39. Erronea (Erronea) caurica caurica (Linnaeus, 1758)
Systema Naturae, ed. 10: 723

(Melicerona) IrEpDALE, 1930
Mem. Queensld. Mus. 10 (1) : 83
> Type Species: Cypraea listert Gray, 1824 <

Zool. Journ. 1: 384
= Cypraea felina GMELIN, 1791

Systema Naturae, ed. 13: 3412

40. Erronea (Melicerona) felina pauciguttata
(SCHILDER & SCHILDER, 1938)
Proc. Malacol. Soc. London 23 (3): 161

Notadusta SCHILDER, 1935
Proc. Malacol. Soc. London 21 (4): 350

> Type Species: Notadusta victoriana ScHILDER, 1935 <
Proc. Malacol. Soc. London 21 (4): 350

41. Notadusta punctata atomaria (GMELIN, 1791)
Systema Naturae, ed. 13: 3412

42. Notadusta katsuae (Kuropa, 1960)
Cat. Moll. Fauna Okinawa 1960: 74; plt. 3, figs. 32 - 34

43. Notadusta musumea (Kuropa & Haze, 1961)
Color Illust. Shells Japan 2: 42; plt. 19, fig. 18

Palmadusta IREDALE, 1930
Mem. Queensld. Mus. 10 (1): 82

> Type Species: Cypraea clandestina LinNaEus, 1758 <
(Palmadusta) TREDALE, 1930

44. Palmadusta (Palmadusta) asellus vespacea
(MeEtvitt, 1905)
Journ. Conchol. 11: 192

45. Palmadusta (Palmadusta) clandestina moniliaris
(Lamarck, 1810)
Ann. Mus. Hist. Nat. Paris 16: 98

46. Palmadusta (Palmadusta) lutea lutea (GMELIN, 1791)
Systema Naturae, ed. 13: 3414

47. Palmadusta (Palmadusta) ziczac ziczac
(Linnaeus, 1758)
Systema Naturae, ed. 10: 722

(Purpuradusta) ScuiLper, 1939
Arch. Molluskenk. 71: 165

> Type Species: Cypraea fimbriata GMELIN, 1791 <
Systema Naturae, ed. 13: 3420

48. Palmadusta (Purpuradusta) gracilis japonica
(ScHILpeR, 1931)
Zool. Anz. 96: 67 - 68

49. Palmadusta (Purpuradusta) fimbriata marmorata
(ScHroTER, 1804)
Wiedem. Arch. Zool. 4 (1): 14

Bistolida CossMANN, 1920
Rev. Crit. Paléozool. 24: 83

> Type Species: Cypraea stolida LinNaEus, 1758 <

(Blasicrura) IREDALE, 1930
Mem. Queensld. Mus. 10 (1) : 84

> Type Species: Cypraea rhinoceros SOUVERBIE, 1865 <
Journ. Conchyl. 13: 156; plt. 511

50. Bistolida (Blasicrura) quadrimaculata quadrimaculata
(Gray, 1824)
Zool. Journ. 1: 376

ile Bistolida (Blasicrura) pallidula pallidula
(Gasxoin, 1849)
Proc. Zool. Soc. London 1849: 97

52. Bistolida (Blasicrura) luchuana Kuropa, 1960
Catal. Moll. Fauna Okinawa 74; plt. 3, fig. 40

53. Bistolida (Blasicrura) teres teres (GMELIN, 1791)
Systema Naturae, ed. 13: 3405

(Bistolida) CossMann, 1920
Rey. Crit. Paléozool. 24: 83

> Type Species: Cypraea stolida Linnagus, 1758 <

54. Bistolida (Bistolida) kieneri depriesteri
(ScuiLpER, 1933)
Journ. Conchol. 19: 355

DoF Bistolida (Bistolida) hirundo neglecta
(Sowersy, 1837)
Catal. Rec. Cypraeidae, London 1: 6

56. Bistolida (Bistolida) stolida stolida (Linnaeus, 1758)
Systema Naturae, ed. 10: 724

Vol. 10; No. 1

Cribraria JouUSSEAUME, 1884
Bull. Soc. Zool. France 9: 94
> Type Species: Cypraea cribraria LinNaEuS, 1758 <
Systema Naturae, ed. 10: 723

(Ovatipsa) IREDALE, 1931
Rec. Austral. Mus. 18 (4): 219
> Type Species: Cypraea chinensis GMELIN, 1791 <

By. Cribraria (Ovatipsa) chinensis chinensis
(GmeELIN, 1791)
Systema Naturae, ed. 13: 3421

(Cribraria) JouSSEAUME, 1884
Bull. Soc. Zool. France 9: 94
> Type Species: Cypraea cribraria LINNAEUS, 1758 <
Systema Naturae, ed. 10: 723

58. Cribraria (Cribraria) cribraria orientalis
SCHILDER & SCHILDER, 1940
Arch. Molluskenk. 72 (2/3): 43

DISCUSSION anp STATISTICAL NOTES

it, Mauritia (Leporicypraea) mappa mappa
(LinnaEus, 1758)

— Cate, 1966: plt. 34; figs. 7a, 7b —
Localities: 41, 45, 48, 69, 79, 85

(@)P 8 L WwW H Lip Col'
Largest shell: ky Ge) Aken} GBS)
Smallest shell: WS GAO BU she ahs

At present, this species has to be considered rare in
the Ryukyu Islands, particularly so in the littoral area of
Okinawa. Reports of its being found at the southern
islands of Miyako and Yaeyama are made occasionally;
however, only 3 other shells are presently known to have
been found at Okinawa (all at Zampa- Misaki), 2 of these

Shell measurements (in millimeters) :

L = length of shell; W = width of shell; H = height of shell;
Lip = number of teeth on outer lip; Col = number of teeth on
columella, excluding terminal ridge.

The numbers in parentheses indicate the number of specimens
examined in each case.

The abbreviations refer to the relative abundance of the species
in the Ryukyus: vr = very rare; r = rare; u = uncommon;
rc = relatively common; c = common.

being dead shells. Sometimes the local market has a speci-
men for sale, but these are thought to have been brought

oo

THE VELIGER

Page 23

from the Philippine Islands. The 2 specimens whose meas-
urements are listed above were collected in 1952, one at
Zampa- Misaki, the other at Yaeyama Island. The Ryukyu
shells, like those from Japanese waters, can usually be
easily distinguished from the Philippine shells by the pale
coloring on the base of the shell; the Okinawa shells are
mostly light beige, sometimes exhibiting a faint flush of
very pale brown, only occasionally pale pink. The south-
ern Philippine shells, in contrast, usually are some shade
of pink or rose. It is interesting to note that the base of
the Japanese shell approaches an almost off-white color
in many instances.

The shell is large, of medium weight, often more
humped; base has a hint of flatness, gently convex ab-
apically toward higher margins; terminals produced,
acutely and strongly formed; apex never completely
covered; margins thickened, evenly calloused, becoming
vaguely flanged adapically, conspicuously so on either
side in front; aperture fairly wide, almost straight; teeth
fine (for a shell of this size), numerous, and not too
heavily formed; columella broad, finely ribbed adaxially;
fossula wide, fairly deep, with an upraised central ridge
within, ribbed with teeth overall; primary shell color is
ivory brown; the dorsal covering is a curious pattern of
fine chestnut-brown, wavering, longitudinal lines; a wide
tributary mantle pattern covers a large portion of the
central dorsal surface (the species name is derived from
this unique pattern) ; large, brown, diffused spots recede
to obscurity from the sides to the base; base and terminals
pale pink-brown blush to off-white, teeth, columella,
fossula, and interstices ivory; there may be a large dark
brown blotch at the spire. It is interesting to note that
this species emits an intense red fluorescence when exposed
to short-wave ultra violet light.

De Mauritia (Arabica) eglantina couturieri
(VayssIERE, 1905) )

— Carte, 1966: plt. 35; figs. 8a, 8b —
Localities: 55, 59, 85

(6) u L W H Lip Col
Largest shell: DO Ome Ol 20:9 39 SS

Smallest shell: HOP le 3ia 20h2) Oo, ON

This species is somewhat difficult to find at Okinawa.
As far as I know the shell is presently found only at
Otubaru and Zampa- Misaki reefs, living hidden in coral
crevices.

The shell is long, narrow, cylindrically-ovate, base
somewhat flattened; terminals produced; aperture long,
narrow, wider in front; teeth numerous, fine, short;
margins rounded, lightly calloused, right side sub-angled;

Page 24

THE VELIGER

Vol. 10; No. 1

basic dorsal color greenish-grey, overlaid with an irregular
network of chestnut-brown lines; there is a wide mantle
line; numerous small (as compared with Mauritia (Ara-
bica) arabica asiatica) dark brown spots thickly decorate
sides; most spots are obscurely visible through thin callus;
base and interstices smoky beige, teeth red brown.

3 Mauritia (Arabica) arabica asiatica
SCHILDER & SCHILDER, 1939

— Cate, 1966: plt. 35: figs. 9a, 9b -
Localities: 3, 9, 13, 19, 20, 20 a, 33, 48, 51, 55, 57, 62 a, 65, 82, 85

(25) G L WwW H Lip Col

Largest shell: (OY? CLO VB) BP We
Smallest shell: SQ 22P ild@ Sil Bs

This is one of the most common cowrie species occurring
in the Ryukyu intertidal zone. It is usually found wedged
within algae-covered coral crevices, in reef pools, and
under rocks and rocky ledges. The frequency of this
species in the adjacent outer islands is comparable to that
at Okinawa. The shell varies a great deal in both size and
shape, from short and wide to elongate and narrow, ap-
proaching the shell character of Mauritia (Arabica)
eglantina couturiert. The large black spots on the heavy,
calloused margins and the teeth on the columellar lip
identify this species. Because of the similarity in general
shape and color of the shells of this species to those of
M. (A.) arabica grayana (ScuitpeEr, 1930) it has often
been confused with this species (ScHILDER, 1966, p. 7).
Mauritia arabica asiatica differs by having larger and
fewer teeth, and by being more round, less pyriform; the
elongate form of M. arabica asiatica poses no problem.
The Erythraean shell is noticeably more abrupt, as well.
The species seems to attain its largest dimensions at
Yonabaru.

One form of this species is oblong-ovate, another
stunted, short, broad, somewhat elevated, semi-cylindrical,
swollen, and solid; base weakly convex, appearing flat-
tened; apex prominent; terminals not produced, well
formed; teeth numerous, fine on columella, larger, more
widely spaced on outer lip; columella broad, channeled,
ribbed; fossula long, broad, deeply concave, ribbed, den-
ticles prominent on adaxial ridge; aperture straight, wide,
more so and constricted in front; dorsal color covering
dark, chestnut-brown, with longitudinal, closely parallel,
broken brown lines and light-grey lacunae; a light-grey
mantle line is also present; light-grey to beige margins
are thickly covered with large, black spots; terminals
grey-black to beige; base and interstices beige, teeth
red-brown.

4. Mauritia (Arabica) maculifera ScuvEr, 1932
Syn.: [Cypraea] reticulata Martyn, 1784 (non-binomial)
The Universal Conchologist 1: fig. 15. London
— Cate, 1965: plt. 8; figs. 26a, 26b —
Localities: 9, 29, 48, 85

(8) u L WwW H Lip Col
Largest shell: 59.8 37.9. 29:6 aaeenmes

Smallest shell: 415° 28.3 20:9 220

The generic name for the synonym cited above has been
placed in square brackets because Martyn did not list a
genus for this species; it became necessary to infer the
name C'ypraea from the preceding line in the text.

This species, in my experience, has the center of popu-
lation in the Hawaiian Islands. Hawaii and Japan appear
to be the northwestern limits of its living range. HaBe
(1961) listed and illustrated this species correctly; how-
ever he listed also Arabica depressa (Gray, 1824), pos-
sibly in error (plt. 19, fig. 25). From the appearance of
his illustration it would seem that his shell is a stunted
form of this species. The range of Gray’s species apparently
does not extend north of the Fiji Islands, it being a more
southern, warmer water species, ranging to Cocos Keeling
Island and elsewhere in the northeastern Indian Ocean.

The Ryukyu shells seem to be smaller on an average
than the Polynesian form. The shell is comparatively
small, solid, compact, oblong-ovate; margins heavily cal-
loused, thick, bumpy; terminals short, restricted; aperture
Narrow, curving; columella broad, semi-denticulate; fos-
sula long, wide, deep; base perceptibly swollen; teeth
short, well defined on both lip and columella; primary
shell color light beige, appearing as lacunae on dorsum;
dorsum conspicuously red-brown, with a narrow mantle
line on the right side; margins heavily speckled with large,
black spots whose dilution in the nacre give the margins,
terminals, and part of the base a dark blackish-grey cast;
areas of the base, interstices, columella, and fossula light
beige; teeth dark red-brown.

5. Mauritia (Arabica) scurra indica (GMELIN, 1791)

Syn.: Cypraea amarata Morcn, 1852
Cat. Conch. Yount, p. 114
— Care, 1966: plt. 35; figs. 10a, 10b —
Localities: 19, 33, 85

(3) u L WwW H Lip Col
Largest shell: Avie) P44 IV B18) BNE,

Smallest shell: 38.3 20.1 158 43 32

Vol. 10; No. 1

This is not a common species, found only at three
presently known locations, Zampa-Misaki, Kadena, and
the dual island locality Henza-Miyagi. There seems to
be no logical explanation for the greater number of
teeth on the lip of the smaller shell listed above. The
species is collected on algae covered dead coral outcrop-
pings and under coral boulders.

The shell is cylindrically-oblong, solid, strong; terminals
produced, heavily formed; aperture long, straight, nar-
row; teeth numerous, fine, somewhat longer on the central
base; margins barely thickened, rounded; basic dorsal
color is greyish-beige, overlaid with a network of yellow-
brown lines, broken with numerous large lacunae; mantle
line also present; margins, sides, base, interstices smoky-
brown, teeth red-brown; large brown spots decorate the
sides; terminal edges smudged with dark brown.

6. Mauritia (Mauritia) mauritiana calxequina
(MELvILL & STANDEN, 1899)

— Cate, 1966: plt 34; figs. 5a, 5b —
Localities: 20, 23, 25, 27, 33, 45, 57, 63, 65, 85

(9) u L WwW H Lip Col
Largest shell: Qe Ci S08 2 Zi
Smallest shell: C28) Od 48 w/o

This species probably seems uncommon because of its
more or less isolated habitat. The animal is restricted
to the outer surfaces of the reef and inaccessible head-
land promontories, living on the exposed rocky-slab sur-
faces facing the open ocean. At one locality on Miyako
Island it was almost impossible to walk due to the number
of these mollusks living on the substrate. It seems best
suited to living in water 6 to 25 feet deep, in pounding,
swirling surf, and even so it is difficult to find and collect.
Yet at another island to the south, Iriomote, at a minus
tide, 32 specimens were counted on a single reef-flat.

Shell medium sized, roundly-ovate, terminals incon-
spicuous, base and lip wide, concave overall; aperture
somewhat narrow, curving left sharply adapically ; margins
sharply angled, solid; teeth strong, large, long; primary
shell color very light beige-brown, appearing as lacunae
in dark red-brown dorsal surface — sides solid, deep
red-brown halfway up shell; base and teeth deep red-
brown, interstices and terminals orange beige; mantle
line sometimes present.

7. Talparia (Talparia) talpa talpa (Linnaeus, 1758)
— Cate, 1966: plt. 35; figs. 11a, 11b —
Localities: 3, 9a, 15, 35 b, 41, 63 a, 65, 85

THE VELIGER

Page 25
(i) we L W H Lip Col
Largest shell: 8215) 444) 36:13, 7,

Smallest shell: AGS? 25:2) 2018) 415) 36

Although this species has been found living at several
localities around the island of Okinawa, it is not commonly
encountered at any of them. MacNeri (1960) reported:
“fragments, internal molds, and otherwise poorly pre-
served specimens of Cypraeids were obtained from many
localities.” He suggests that Talparia talpa talpa, as indi-
cated by the appearance of the shell fragments, may have
been recovered from at least 8 localities in the Okinawan
fossil beds. One recent collection of a living specimen was
made at the edge of the reef off Gushichan; 2 or 3 beach
specimens have also been recovered at Zampa-Misaki
and Metasaki.

The shell is of medium size for the species, light weight,
cylindrically elongate; base long, fairly narrow, vaguely
convex ; terminals produced, thickly, solidly formed; aper-
ture long, narrow, nearly straight; margins thickened,
more so on the right, gently angled; teeth numerous, fine,
distinct, short; basic shell color ivory-yellow, with 4 wide
transverse bands of light chestnut-brown; interstices ligh-
ter in color, nearly off-white; wide, partially ribbed
columella yellow-brown; short, wide, deep, off-white
fossula is ribbed and denticulate adaxially.

8. Cypraea (Cypraea) tigris pardalis SHaw, 1795
— Cate, 1966: plt. 34; figs. 6a, 6b —
Localities: 27, 33, 35 a, 48, 51, 57, 57 a, 59, 63 a, 65, 65 a, 81, 85

(5) u L WwW H Lip Col
Largest shell: led) rete ON 7 70)
Smallest shell: Sie CB) GQ D7 iI9

This species is not very plentiful, being found only now
and then. It lives on coral shelves just below low tide mark.
Fishermen most often, in diving operations, bring shells
of this species in and offer them in the market place. The
Naha limestone formation has yielded a fossil fragment,
according to MacNeit (1960), of a large Cypraea that
seems related to C. tigris pardalis, but appears to differ
from it by possessing a depressed area on the right lip
just adjacent to the base of the teeth. From his excellent
figure (/.c., plt. 17, fig. 1) it is impossible to determine
whether the depressed area is there because of a loss in
shell material, or is the morphological character of a
different species. The base on the left side appears to
exhibit signs of surface attrition as well. It is conceivable
that the sloughing away of the thick marginal callus on

Page 26

THE VELIGER

Vol. 10; No. 1

either side and on the base could leave the shell’s teeth
elevated in a manner such as this figure illustrates. Frag-
mentary evidence of this marginal callus still clings to the
right side of the MacNet shell. The visible apertural teeth
seem to conform with those of the species as we know it
today. The Ryukyu C. tigris pardalis that I have examined
seem to be of smaller dimension than those seen from
the more southern Philippine area. A recent discovery
(Way) shows the species to be actually quite common
half a mile off shore at Kin Bay, in 35 to 40 feet of
water.

The shell is of medium size, medium weight, solid, sub-
pyriform; base and lip swollen, concave to aperture;
aperture wide, fairly straight ; terminals barely protruding;
margins rounded, thickened; columella unribbed adap-
ically, ribbed abapically to and including the broad,
shallow fossula; basic shell color on dorsum varies in
shades of brownish-orange, overlaid with large, diffused
black spots that continue over the sides and margins to
the base; columella marked with vague orange-brown;
base, teeth, and interstices are white.

9. Cypraea (Lyncina) argus argus LinNAEuS, 1758
— Cate, 1966: plt. 33; figs. 3a, 3b —
Localities: 1, 33, 35 a, 41, 85

(4) u L W H Lip Col
Largest shell: 80:5) 4325/5 35 Oe 4 Oe ST
Smallest shell: 6123) 30:8 2445 735) 338

The large shell, whose measurements are recorded here,
was collected in early December, 1954, at Metasaki. Live-
collected shells of this species are rare, although beach
shells in good condition are found from time to time at
this locality and at Zampa-Misaki. MacNei (1960) de-
scribes a related species from the Naha Pliocene lime
formation, Aristorides nahaensis. He says, that although
closely related to Aristorides argus argus (LINNAEUS,
1758), it differs most importantly in that the columellar
teeth do not extend into the aperture, apparently not
crossing the columella or fossula. Its length and width are
given as 40mm and 24mm, respectively. The holotype
is in the U.S. National Museum with the catalogue
number 563023.

Shell of medium size, relatively light weight, cylindri-
cally elongate, abruptly elevated front and back, central
dorsal surface almost horizontal; base and lip convexly
rounded; terminals inconspicuous; aperture wide, nearly
straight; margins lightly calloused, more so, but narrowly
restricted, on the right side, thickly enveloping the terminal
openings; teeth numerous, short, well defined; columella
broad, fossula long, becoming deeper in front, both strong-

ly ribbed with teeth; primary shell color creamy-cocoa,
overlaid dorsally with 3 wide, transverse pale cocoa-
colored bands, and over all, numerous “eyes” or ocellae;
many are thin-lined rings, others, less numerous, are
larger, thick-lined, heavier ocellae; 4 large, brown, ventral
blotches at either quarter of the base — sometimes the
right rear blotch is incomplete or missing; terminals,
base, teeth, and interstices creamy-cocoa; teeth outlined
on either side with a fine broken line.

10. Cypraea (Lyncina) lynx vanelli Linnarus, 1758
Syn: Cypraea lynx var. michaelis MEtvitL, 1905
Journ. Conchol. 11: 192
— Cate, 1966: plt. 35; figs. 12a, 12b —
Localities: 17, 33, 37, 39 a, 41, 55, 57, 65, 85

(4) u L WwW H Lip Col
Largest shell: 41.6 24:7" “OTS oe

Smallest shell: 307) 178 Sa 2S

This species is uncommonly found on the coral reefs,
living on ledges, in cavernous tunnels, and in algae covered
tide pools. The invasion of this species into these islands
seems to be of relatively recent date, as there appears to
be no reference to its being found in the lime formations
as late as the Pleistocene. Although comparing very
favorably morphologically with the shells found in the
Philippines, the Ryukyu shells seem to be generally smaller
in size and narrower, with a distinctly deeper dorsal
coloring.

Shell narrowly ovate, solid; margins thickened, more
so on the right side, somewhat angled; terminals semi-
produced, well developed; aperture narrow, only slightly
curving; columella broad, fossula shallow, both heavily
denticulate; base flattened, sloping sharply adaxially, lip
rounded; primary shell color light beige, overlaid dor-
sally with light yellow-brown, with large and small dark
brown spots irregularly dotting the upper surface, becom-
ing obscure in the marginal callus; teeth short, well
defined, fine on columella, heavier on inner edge of outer
lip; margins, base, and teeth light cream color, inter-
stices bright orange.

11. Cypraea (Lyncina) vitellus vitellus Linnaeus, 1758
— Cate, 1966: plt. 36; figs. 17a, 17b —
Localities: 1, 9, 13, 33, 37, 39, 57, 65, 81, 85

(7) rc L Ww H Lip Col
Largest shell: 389 263i e212 S el
Smallest shell: ee ass IO IE IS)

Vol. 10; No. 1

This species is relatively common, occupying crevices
and the underside of coral ledges. Most of the shells I
have examined appear to be uniformly small for the
species. The coloring is richly dark grey-brown, the smudge
of which darkens the normally beige colored base. This
species seems to be equally common at each of the
localities listed.

Shell relatively small, abruptly humped, pyriformly-
ovate, strong, solid; margins thickened, especially the
right side; terminals prominent, thickly formed; base and
lip swollen; aperture somewhat narrow, fairly straight;
teeth do not extend onto base, are semi-fine, long, crossing
the broad columella and the long, shallow fossula; teeth on
lip somewhat longer, heavier, more numerous; dorsal
color dark grey-brown, copiously marked with large and
small spots; margins fawn, with the characteristic nu-
merous, fine transverse striae, sweeping from the outer line
of the base teeth to well up onto either side of the dorsum;
teeth pale ivory, most of the base and interstices pinkish-
fawn to light grey.

12. Cypraea (Lyncina) schilderorum IrEpDALE, 1939
Syn.: Cypraea arenosa Gray, 1824
Zoolog. Journ. 1: 147; plt. 7, fig. 6; plt. 12, fig. 6
— Cate, 1965: plt. 9; figs. 31a, 31b -
Localities: 49, 51, 77

(2) r IL WwW H Lip Col
Largest shell: 308) ZB40° Wi 82 Ws)

Smallest shell: SilO 220 IGs 2 2

In the synonym cited above, Dittwyn (1823) invali-
dated this name by publishing SoLaNDER’s name arenosa
(Index Lister Hist. Conch., p. 33, 1823) and no other
names were available; a new name, therefore, became
necessary.

This species is rare in the Ryukyu Islands. The smaller
shell recorded above was collected in 1952 on the beach at
Nago. Broken beach-worn shells have been collected at
the same locality, and it is probable that SCUBA-diving
will some day produce live-collected specimens. The larger
shell, also a beach specimen, was found at Yabu. Both
shells appear to be typical representatives of the species.
MacNem (1960, p. 50) recorded a single, imperfect
specimen from the Pliocene limestone at Naha. His figures
(I. c., plt. 17, figs. 4 - 8) are well defined and clearly seem
to represent this species.

Shell ovate, perceptibly flattened, solid, heavy; termi-
nals inconspicuous, especially adapically ; aperture straight,
narrow; margins thickly calloused, sharply angled; teeth
short, fine, numerous; base generally convex; primary

THE VELIGER

Page 27

shell color beige to off-white; dorsum dark beige, over-
laid transversely with 4 somewhat narrow reddish-beige
bands; marginal beige callus covers sides well up onto
shell; conspicuous grainy texture and numerous vertical
striae in the lateral callus; portion of base, teeth, and
interstices off-white.

13. Cypraea (Lyncina) carneola carneola LinNaEus, 1758
— Cate, 1966: plt. 36; figs. 16a, 16b —
Localities: 9, 39, 41, 65, 85

(5) rc L W H Lip Col
Largest shell: 33) Olean l/l
Smallest shell: OA) WaT WP 7 DP

This species is rather uncommon, although it has been
collected at several localities. When it is found it is usually
on coral ledges of the reef adjacent to deep water. At
Zampa- Misaki it was found nestling in crevices around the
circumference of a littoral area locally referred to as the
Punchbowl. It was living under clusters of a certain species
of sea anemone that nearly covered that portion of the
reef floor. These shells, like so many other cowrie species
in the Okinawa area, are comparatively smaller in size
than those from Philippine waters. Nearly all of the speci-
mens I have observed appear to have broken lips that
subsequently have been mended in the process of continued
growth.

Shell oblong-ovate, solid, small; margins heavily cal-
loused, somewhat bumpy in fully adult shells; terminals
inconspicuous, though strong and thickly formed; base
semi-rounded, uneven; aperture fairly straight, curving
left only slightly; teeth well developed, but not lengthened
onto base or lip; columella broad, fossula long, deep, and
prominently ribbed adapically by the inner teeth; primary
shell color pale grey-blue, overlaid with 4 bands of pale
tomato-red; margins have a grainy texture of deep beige,
base and terminals somewhat lighter in color; teeth and
interstices bright violet.

14. Cypraea (Lyncina) kuroharai (Kuropa & Hane, 1961)

(Plate 3, Figures 2a, 2b)
Localities: 53, 62

(1) Vr L WwW H Lip Col
Shell measurements: Ole BUG CRgs Be Bi)

This is an extremely rare species and is supposed to
have been taken only in Ryukyu waters. Specimens that I
know of are as follows: Akibumi Teramachi, Kyoto, a live-
collected specimen and one from the Ryukyu Islands; in

Page 28

THE VELIGER

Vol. 10; No. 1

ee

another collection there is one allegedly from Kyushu
(Koshik Islands), one from Kii and two from Tosa Bay.
Kurohara has one specimen, Sugatani has one, and one
live-collected one is in the author’s collection (Cate no.
C3214) (Plate 3, Figures 2a, 2b). Scumper (1963)
mentions two specimens, one in the Habe collection
(L 42mm), and one (subfossil) in his own collection
(Schilder no. 15820). The specimen whose measurements
are listed here for comparative purposes was live-collected
in 70 fathoms by trawling off Tosa, Japan, in 1962. It
compares favorably with the holotype in mensurable
characters. Although shells of this species have been in
Japanese collections for many years, it was only recently
recognized as being different from Cypraea (Lyncina)
arenosa Gray, 1824 (=C. schilderorum IrEDALE) for
which it had previously been mistaken; it had also been
compared with C. (L.) sulcidentata Gray, 1824, by some
workers.

Shell pyriformly ovate, heavy, solid; margins thickened,
though not prominently so; terminals prominent, openings
narrow, thickly formed; base and outer lip swollen,
rounded; aperture narrow, curving left adapically; teeth
short, not extending onto base or lip, very fine, sharp;
those on columella larger, extending pointedly into aper-
ture; columella fairly broad, fossula short and deep, both
ribbed with columellar teeth; terminal ridge straight;
primary shell color beige, base lighter beige, margins
darker beige and grained; dorsum faintly marked with 4
broad, deep tan transverse color bands; lower terminal
surface, base, teeth and interstices light creamy-beige.

15. Chelycypraea (Chelycypraea) testudinaria testudinaria
(Linnaeus, 1758)
— Cate, 1966: pit. 33; figs. 4a, 4b —
Localities: 45, 47, 62, 63

(7) u L W H Lip Col

1128 564 440 45 43
10525010) 44 OmeeLD

Largest shell:
Smallest shell:

The shells used in this study were sent from Okinawa
on various dates during 1952 and 1953 without specific
locality data, the accompanying labels simply reading
“Okinawa”. This is regrettable; however, a report from
Bernice Albert of Okinawa states: “... we have one
we found with some tiger cowries the fishermen had
brought in, but they are rather vague as to where they
had found it. We have known of others brought in by
fishermen, but none by divers or collectors.” Further
exploration will be needed in this area to ascertain the
extent of the distribution of this species here.

Shell large, ponderous, solid, strong, cylindrically elon-
gate; base somewhat swollen, yet because of its size

seems flattened; terminals prominent, apex a part of
left side; aperture long, straight, wide; rear half of
columella smooth, front half and the long, wide, deeply
concave fossula ribbed with teeth; teeth on both lips
comparatively small, weakly formed, very short; margins
only barely calloused, rounded; primary shell color café-
au-lait to beige, overlaid with large, irregular areas of
brown, including many various sized large brown spots
— additionally a thick haze of fine white specks is spread
over ail; base, inner terminals, and interstices of primary
shell color, teeth light beige.

16. Luria (Basilitrona) isabella rumphu
SCHILDER & SCHILDER, 1938

— Cate, 1966: plt. 35; figs. 13a, 13b —
Localities: 9, 13, 15, 33, 41, 57, 85

(5) u L W H Lip Col
Largest shell: 26:7 14.8 1272 3omeeo
Smallest shell: 25.9 149° 1200 32a

This is not a common species, though it is relatively
well distributed throughout the island’s waters. Adult
shells are not very large and are fairly well characterized
by a distinctly wide, flattened upper shell surface tra-
versing the central dorsal area from side to side. This
species grows comparatively larger in the Philippines
and still larger in Hawaii.

Shell cylindrically ovate, narrow, solid; margins only
barely thickened; terminals blunt, weakly formed; base
area narrow, semi-flattened, lip rounded as is left mar-
ginal surface; aperture long, narrow, slightly angled ab-
apically; teeth very short, extremely fine, though well
developed, extending neither onto base nor lip; columella
long, fossula deep, smooth, adaxial edge of fossula denti-
culate; primary shell color white, overlaid dorsally with
fawn-grey; two narrow, lighter beige-colored bands divide
the upper surface transversely, and numerous irregular,
fine, black horizontal lines over all lengthwise; terminals
orange, faintly quadrimaculate with dark brown; lower
margins, base, teeth and interstices stark white.

17. Pustularia (Annepona) mariae (ScuiLpeER, 1927)
Syn.: Cypraca annulata Gray, 1825 (nom. nud.)
Zool. Journ. 1: 518

Cypraca annulata Gray, 1828
Zool. Journ. 4: 88

~ Cate, 1966: plt. 36; figs. 18a, 18b —
Localities: 7, 49, 55
This species is rare in the Ryukyu Islands as it is

at all other localities. Indications are that it lives in deep
water as it is almost never found intertidally, only occa-

THE VELIcER, Vol. 10, No. 1 [C. N. Cate] Plate 3

Figure 1 a Figure 2 a Figure 2 b

Figures 1 a, 1b:
Notadusta katsuae (Kuropa, 1960), ex C. N. Cate Coll. no. 3392; (x 2)

Figures 2a, 2b:

Cypraea (Lyncina) kuroharai (Kuropa & HaBe, 1961), ex C. N. Cate Coll. no. 3214;
(natural size)

Figures 3 a, 3b:
Notadusta musumea (KuropA & HaBeE, 1961), ex P Clover Coll. (x 14)

photographs by Takzo Susuk1

Vol. 10; No. 1 THE VELIGER Page 29
(2) z L W i li Gal 19. Pustularia (Pustularia) bistrinotata mediocris

ee ae SCHILDER & SCHILDER, 1938

Largest shell: I WO 0) BB) 1 283

Smallest shell: 11.0 74 Gye ole me 22 © COB Mielbs PE 23 ise AN 2, HOLS

TS Localities: 33, 49, 59

sionally as beach rolled shells. The two shells, whose

measurements are recorded here, and a broken fragment (19) u L W H Lip Col

of a shell were collected as dead beach specimens at Largest shell: 20459 12:9) 117, 30) 24

Okinawa by Anita Scott in 1952. Recently another dead Smallest shell: 15.7 9.4 Sui

beach shell was found at Baten, a second shell was picked
up on the beach just south of Nago. Nothing more can be
reported on this species at this time.

Shell thin, lightweight, bulbously ovate; base narrow
on either lip, peculiarly flattened; terminals barely visible;
aperture long, exceedingly narrow, curving gently; teeth
very fine, weak, poorly developed, almost obscure on
inner lip; margins not thickened, a rounding continua-
tion of the dorsum; primary shell color milk-white, upper
surface and sides thickly covered with ocellated, irregularly
shaped and sized straw-colored spots; base, teeth, and
interstices milk-white.

18. Pustularia (Pustularia) cicercula cicercula
(Linnaeus, 1758)

— Cate, 1966: plt. 36; figs. 19a, 19b —
Localities: 13, 15, 43, 75

(10) ou Wi GEE Lipet ol

Largest shell: NE) UTES) OS Bil 28)
Smallest shell: 14.5 8.8 SO 2G 2

Gushichan has been a productive locality for this
species. More recently, however, freshly dead shells have
been found at Chinen and Toyohara. I have live-collected
specimens from Minami-Ukibaru Island, northern Buck-
ner Bay, collected in 1953. MacNer (1960, p. 50; plt.
19, figs. 1, 2) seems to have mistaken this species for
Pustularia g. globulus (see discussion of this shell here
under the title of that species). It would therefore follow
that Pc. cicercula has not been found in the fossil deposits
of Okinawa.

Shell pyriformly-ovate, numerously, completely, pustu-
late dorsally ; grooved mantle line originates within small,
deep umbilicus; terminals extended, beaked, delicately
formed; aperture long, narrow, straight; margins not
thickened; base semi-inflated, convexly arched longitudi-
nally; teeth numerous, fine, distinct, extending to both
margins inclusive of terminal beaks; shell uniformly
lemon-beige, glossy though pustulate.

The largest shell whose measurements are recorded
here was picked up on the beach at Kadena, and two
others, including the small one recorded here, were col-
lected on the beach about a mile south of Nago. Other
beach-rolled shells have been found at the latter locality
and at Otabaru. I know of no live-collected specimens in
Okinawan collections. Further field work may disclose
the habitat of these mollusks.

Shell semi-ovate, globularly humped, lightweight, and
strong ; terminals extended; beaked; mantle line originates
in shallow, vague umbilicus; margins thickened, upswept,
smooth; aperture long, narrow, straight; base flattened,
convexly elevated at either end; central dorsal area
smooth (pustules absent), glossy; sides and ends of dorsal
area thickly covered with pustules; teeth numerous, fine,
distinct, extending out to either margin; primary shell
color pale yellow-beige, dorsum with a light brown blush,
with pustule crests darker brown and a large brown spot
centrally situated, as well as one at each end.

20. Pustularia (Pustularia) globulus globulus
(Linnaeus, 1758)

— Cate, 1966: plt. 37; figs. 21a, 21b —
Localities: 31, 47, 69, 83

(8) ir L WwW H Lip Col

Largest shell: PAO NB iL EE 2)
Smallest shell: leks) ER TO) SE

This species is quite rare at Okinawa. Few specimens
are to be found in local collections. The shells listed here
(Cate coll. no. C3390) were collected in the beach drift
at Ishiza by Anita Scott in 1953; the most recent collec-
tions were made by Bernice Albert at Onna tide flat in
1965.

MacNeiL (1960, p. 50; plt. 19, figs. 1, 2) records
“Pustularia cf. PR cicercula (LinNE) s. l.,” and illustrates
the shell. It seems to me that the shell shown in the figures
cited rather clearly represents the nominate species, P g.
globulus. He states, “It lacks dorsal granules, has no indi-

Page 30

cation of sulcus, and the teeth tend to become weak or
absent towards the central part of the aperture.” The
lack of dorsal granules would appear to eliminate P c.
cicercula, and the weakening or shortening of the colu-
mellar teeth rather emphatically indicate P g. globulus.
This weakening of the central columellar teeth can be
easily observed in a series of these shells, never in P c.
cicercula, and it is visible in the specimen illustrated in
Cate, 1966 (plt. 37; fig. 21b). The columellar teeth of
all other species in the genus Pustularia are long and
strong, extending to the margins; this species appears to
be the exception.

Shell pyriformly ovate, humped, bulbous; margins de-
fined, though barely perceptibly thickened, and upswept on
the right side; terminals protrude, beaked; aperture long,
narrow, curving slightly to the rear; teeth fine, numerous
centrally on columella somewhat shortened; the columella
and the deep fossula half-ribbed, half-smooth adaxially;
shell color yellow-beige, copiously spotted with medium
sized brown spots; base and interstices yellow-beige; teeth
darkened, some light brown; faint brown marks at either
quarter on the base. (It should be stressed that these color
descriptions are taken from dead, though fairly fresh
beach shells and do not represent the natural color of
live-collected shells. )

21. Pustularia (Ipsa) childreni samurai ScuiLveEr, 1940
— Cate, 1966: plt. 37; figs. 22a, 22b —
Localities: 3, 13, 49, 57, 82

(13) u L WwW H Lip Col
Largest shell: Zl Sean OG Se

Smallest shell: 16.3 10.6 85 37 24

This cowrie species throughout its Indo-Pacific range
seems always to be difficult to collect alive. The animals
from the Ryukyu Islands are no exception, as those in
most of the local collections have been gathered dead on
the beaches. Onna, Nago, and Chinen are the localities
where the species is most commonly found.

Shell rectangularly inflated, abruptly terminated at
either end, base flattened, peculiarly flanged at the side of
either terminal; terminals barely projecting adapically,
hardly more so in front; aperture narrow, nearly straight;
right margin thickened, upswept; mantle line grooved,
originating from small, shallow umbilicus; teeth numerous,
fine, sharply defined, each denticle a continuous ridge
from columellar tooth across base and over dorsum, ter-
minating as a tooth on the outer lip; shell color a deep,
rich, faintly brown honey-yellow, color darker only on
small spot at center of umbilicus.

THE VELIGER

Vol. 10; No. 1

22. Monetaria (Ornamentaria) annulus annulus
(Linnagus, 1758)

Syn.: Monetaria harmandiana RocHEBRUNE, 1884
Bull. Soc. Malacol. France 1: 90; plt. 2, fig. 5

— Cate, 1966: plt. 37; figs. 23a, 23b —
Localities: 9, 13, 17, 19, 23, 25, 33, 37, 39, 41, 65, 81, 85

(IS) G L WwW H Lip Col
Largest shell: 2716 + 20:10) 143" ieee

Smallest shell: ISO) 30)33 7.8 9 9

This is one of the most common cowrie species living
in Ryukyu tidal waters. It can be found plentifully on
nearly all tide flats and shallow reefs. Adult shells seem
to have a wide range in size variation, yet shells of a par-
ticular locality being fairly uniform; at Metasaki, for
example, they are noticeably large. Especially observable
in these shells are the numerous very fine central white
lines that traverse the median, smoky-grey dorsal sec-
tion of the shell; although perceptible in the shells
from other localities, it seems more evident here.

Shell rhomboid-ovate, flat, broad, solid; margins solidly
thick, angled, perceptibly upswept; terminals blunt, a
continuation of the margins; base swollen, lip less so,
flattened; aperture straight, widening to the front; teeth
large, strong, well defined, lengthening slightly onto base
and lip; columella smooth, fossula weak or missing entire-
ly; primary shell color creamy-beige, central dorsal sec-
tion light grey, transversely marked with numerous fine
white lines, an orange line encircles the central dorsal
area; sides, margins, terminals, base, teeth, and inter-
stices of primary shell color.

23. Monetaria (Monetaria) moneta rhomboides
SCHILDER & SCHILDER 1933

— Cate, 1966: plt. 37; figs. 24a, 24b —
Localities: 1, 3, 7, 11, 13, 15, 20, 33, 37, 39, 39a, 51, 62a, 65, 73,

75, 75a
(18) c L W H Lip Col
Largest shell: 25:9 20:0) 12.3) atom

Smallest shell: 15.0 9.0 HO. iil 1

This is a common species at Okinawa, but not as
plentiful as Monetaria annulus annulus. Like the latter
species, it can be found on most of the tide flats. The
shell’s shape indicates that this subspecies is well named.
The shells appear to be nearly identical with those from
the Philippine area.

Shell rhomboid, flattened, margins protruding promi-
nently adapically; margins thick, narrow, sharply angled;

Vol. 10; No. 1

THE VELIGER

terminals and openings thickly formed; base and lip
convexly swollen; aperture straight, broadening slightly
abapically; teeth well defined, finer on the columellar
edge than on the lip; columella smooth, fossula weak or
absent ; primary shell color off-white to light ivory, central
dorsal area yellow-green, with two distinct darker green
narrow bands traversing dorsum from side to side; sides,
terminals light ivory, becoming lighter in color on inner
half of base, teeth, and interstices.

24. Erosaria (Ravitrona) labrolineata labrolineata
(Gasxorn, 1849)

Syn.: Cypraea flaveola Gray, 1825
: Zool. Journ. 1: 502
— Cate, 1966: plt. 38; figs. 27a, 27b —

Localities: 19, 20, 41, 57, 61, 65, 82, 85

(7) u L WwW H Lip Col
Largest shell: 20.3 10.6 3:3) 9 20) 20

Smallest shell: 14.4 8.8 OSes:

This uncommon species is occasionally found on the
tide flats at low tide. The shells seem to have quite a
wide size range in adults. MacNert (1960, p. 52; plt. 2,
figs. 17, 18, 25) lists a species “Cribraria (Talostolida)
aff. C. (T.) cincta (Martin, 1899),” from the Miocene
clay of Yonabaru, Okinawa. The shell compares favorably
morphologically with Erosaria (Ravitrona) |. labrolineata,
and may indeed be this species.

A dead shell, without animal, was found in 110 feet
of water at Metasaki.

Shell pyriform to oblong-ovate, sometimes cylindrical,
base swollen; terminals prominent; margins thickened,
mostly on the crenulated right side, angled, shouldered;
aperture wide, more so abapically; teeth prominent,
medium sized, shorter in the central columella area,
larger, evenly dispersed on outer lip; columella smooth,
fossula absent; primary shell color white, overlaid dor-
sally with greyish-white to dark grey (as seen in the mantle
line), upon which is a covering of tawny, light-brown,
broken with numerous large and very small light-grey
lacunae; margins, more particularly the terminal collars,
spotted and marked with chestnut-brown; terminals, base,
teeth, and interstices white.

25. Erosaria (Ravitrona) cernica ogasawarensis
ScHILDER, 1944

Syn.: Erosaria tomlini maturata Kira, 1954
Japan. Shells Col. 1954: 39; plt. 19, fig. 11
— Cate, 1960: pit. 1; figs. 5, 5a —
Localities: 20 a, 35, 85

Page 31

(3) r L W H Lip Col
Largest shell: BIO) AO WG 22 Aly)
Smallest shell: BUA) NTs NBeff Bil AG

This species, whose type locality is given as the Ogasa-
wara Islands, is found in both Japanese and Ryukyu
waters and may be said to belong equally to these two
areas because of the proximity of the type locality to
each. Even s0, it is rare at Okinawa, only one specimen
having been found there. This specimen was collected on
the Zampa-Misaki reef flat on April 7, 1966 by Bernice
Albert. This particular specimen, the fourth I have ex-
amined (23.2 15.3. 12.0 19 15), appears to have
just attained maturity as the shell is comparatively thin
and lightweight.

Shell ovately pyriform, solid; terminals faintly pro-
duced; aperture wide, gently curving; teeth well defined,
strong; right margin thickly calloused, upswept, angled,
shouldered, pitted, left side barely thickened; primary
shell color off-white, dorsum covered with light brownish-
yellow and numerous faint off-white spots, a vague mantle
line also present; both sides thickly covered with brown
spots; terminals, base, teeth, and interstices off-white;
narrow columella smooth, shallow fossula ribbed.

26. Erosaria (Ravitrona) helvola helvola
(Linnaeus, 1758)

— Cate, 1966: plt. 38; figs. 30a, 30b —
Localities: 33, 35, 37, 39, 47, 51, 57, 61, 63 b, 65, 67, 85

(7) u L WwW Ee ae Lipye Col
Largest shell: BE MG UN) NS

Smallest shell: Dido Weofl IOAE 20) ee

Though not at all common, the species is widely
distributed. It is usually collected in reef areas, in coral
recesses and on protected ledges. The Ryukyu shells are
of average size for the species. MacNert (1960, p. 51;
plt. 17, figs. 2, 6) reports finding the species in the Plio-
cene layer of the Naha limestone.

Shell ovate, short, broad, strong, base swollen; terminals
short, well formed; margins thickened on either side,
upswept, shouldered, crenulated; aperture narrow, curv-
ing; teeth large, bold, heavily formed, extending up onto
base and lip, longer toward the rear on base; columella
narrow, smooth, fossula long, shallow, ribbed; shell color
light caramel-brown, overlaid dorsally with deep, rich
brown; a faint grey mantle line and numerous, various
sized off-white spots are exposed; terminal collars, spire
pale lavender; margins, base, teeth, and interstices rich
light caramel-brown, base slightly darker.

Page 32

27. Erosaria (Ravitrona) caputserpentis caputserpentis
(Linnaeus, 1758)

Syn.: Ravitrona caputserpentis mikado ScHILDER & SCHILDER,
1938

Proc. Malacol. Soc. London 13 (3): 135
— Cate, 1966: plt. 38; figs. 31a, 31b —
Localities: 1, 3, 9, 13, 20a, 37, 39, 39a, 51, 59, 81, 85

(30)eaac L W H Lip Col
Largest shell: 300 AOD yd 19 5

Smallest shell: 26.9 19.8 12.9 14 12

This species is plentiful, living on reef flats, seemingly
always adjacent to deep water. One specimen had a red
streak of color along one side of the shell. In my experience
the species appears to be most commonly found at Smug-
gler’s Cove. In a study of series, involving at least 25
specimens each from three localities (Japan, Philippines,
and Ryukyus), I was unable to detect any morphological
or color differences. From my observations that this species
has a more or less continuous distribution from the Sulu
Sea and Tosa Bay, I have come to the conclusion that
Ravitrona caputserpentis mikado SCHILDER & SCHILDER,
1938 must be considered a synonym of the nominate
species. MacNeix (1960, p. 51; plt. 17, figs. 2, 6) records
shells of this species from the Naha layer of Pliocene
limestone.

Shell rhomboidally-ovate, strong, heavy, wide, appear-
ing flattened; margins well developed, sharply angled,
curving, perceptibly upswept; terminals obscure, though
well formed within; base and lip somewhat convex; aper-
ture long, narrow, curving abruptly adapically; teeth
large, of equal size on each lip, pointedly so on the
apertural edge; sides and outer surface of base dark
brown, remainder of base and interstices pastel shades of
light brown and beige; central dorsal area light brown
broken with numerous off-white spots of various sizes;
a thin mantle line may be present.

28. Erosaria (Erosaria) poraria scarabaeus (Bory, 1827)
— Cate, 1966: plt. 39; figs. 32a, 32b —
Localities: 9, 41, 57, 59, 61, 77, 83, 85

(8) rc L WwW H Lip Col
Largest shell: AVA NS. WO UD. la

Smallest shell: 16.0 11.0 B38 7s 1G

While seldom collected alive, shells of this species are
quite often seen in the beach drift, especially at Zampa-
Misaki. MacNeit (1960, p. 52; plt. 18, figs. 19, 22)

THE VELIGER

Vol. 10; No. 1

appears to have recorded this species under the name
Palmadusta asellus (“Cypraea” sp. aff. “C.” asellus Lin-
NE), from the Yontan limestone of Okinawa (see species
43 below for a discussion of this identification) .

Shell ovate, longer than wide, strong; base convexly
swollen, lip curving; margins heavily calloused, upswept,
sharply angled, shouldered, and crenulated on the right
side; terminals inconspicuous; aperture narrow, nearly
straight; teeth fine, well developed, short, barely extending
onto base and lip; dorsal color medium brown, with
numerous darker lacunae having off-white centers; ter-
minals and base deep lavender, teeth and interstices pale
lavender.

29 Erosaria (Erosaria) erosa phagedaina
(MEtvitt, 1888)

— Cate, 1966: plt. 39; figs. 33a, 33b —
Localities: 3, 9 a, 13, 19, 33, 37, 39, 48, 51, 57, 62 a, 63 b, 65, 69a,

81, 82, 83, 85
(11) Cc L W H Lip Col
Largest shell: 34.6... 19.4 , 15100 SZ es
Smallest shell: D0) te jlilsil =I

This species is quite common at several localities. It is
found in coral pockets, under rocks, and under over-
hanging submerged ledges. It has been established in
these islands for many centuries; MacNeit (1960, p. 51;
plt. 19, figs. 9, 12) records this species from the Pliocene
(Naha limestone) and Pleistocene (Yontan limestone),
Okinawa. The adult shells vary in size, though only
occasionally becoming as large as the Philippine speci-
mens, Yonabaru seeming to produce the exceptions.

Shell oblong-ovate, semi-umbilicate, solid, base con-
vexly swollen; terminals produced, thickly formed; mar-
gins thickened, shouldered, and pitted on the right side,
pitting extending over and around terminal collars; teeth
coarse, well formed on outer lip, finer and shorter on the
columella, receding to mere pustules centrally; aperture
wide, straight, somewhat constricted adapically; basic
shell color ivory with tannish-grey-green covering the
dorsum; numerous off-white spots of various sizes, some
encircled by a darker ring, speckle the upper surface, a
faint mantle line is also present; base, teeth, and inter-
stices ivory; both margins have the large brown marginal
banding characteristic of this species; lateral divisions
of margin pitting crested with a short, fine brown line.

Ko

Vol. 10; No. 1

THE VELIGER

Page 33

30. Erosaria (Erosaria) miliaris miliaris (GMELIN, 1791)

Syn.: Cypraea inocellata Gray, 1828
Zool. Journ. 1: 504
— Cate, 1966: plt. 39; figs. 34a, 34b -
Localities: 57, 81, 85

(5) or ToeselW aniEone Lip.Col
Largest shell: AG Th (Oe Tis Th
Smallest shell: BD FO 158 We

The largest shell, whose measurements are recorded
here, was collected alive by Anita Scott in 1957. It is
completely adult and heavily calloused. The locality is
cited simply as “Okinawa”. Since then the species is
known to have been found in at least three localities,
most recently by Emest Albert at Yagaji, Okinawa. His
comments were: “Yagaji is sandy, with some areas sup-
porting sea grass, while at the end of one edge of the
locality it is gravelly and rocky. Although the sand flat is,
for the most part, barren of rock I found this shell living
under one near the gravel area. It seems to be of a
deeper color than those we have seen from the Philippines,
and the base is more pinkish. We searched for more
but could find none.”

Shell large, heavy, thickly calloused, pyriformly ovate;
terminals prominent, fully developed; aperture fairly
straight, wide, especially so abapically; margins heavily
calloused, upswept, angled, crenulated along both sides
and over terminal collars; teeth large, strong, short; dorsal
color pale grey-green, lightened in shade by a thin white
film of nacreous callus over all; terminals, base, teeth,
and interstices white.

31. Staphylaea (Staphylaca) staphylaca staphylaea
(LinnaEus, 1758)

— Cate, 1966: plt. 39; figs. 35a, 35b -
Localities: 15, 39, 44, 57, 62 a, 75 b, 81

(5) u L W H Lip Col
Largest shell: WES OE2 Bil AO UG
Smallest shell: 7S 10,2 G2 INQ

This species is not at all common. At Onna, on the east
(China Sea) side of Okinawa, the two specimens whose
measurements are recorded here, were found on a sand
flat under a bush-like, green, spongy algal growth, whose
base was embedded in a viscous black substance. The hab-
itat of this species seems to be restricted to this type of alga
in the Ryukyus. The shell seems to be typical in appear-
ance for the species, but generally not as large as the
Philippine specimens.

Shell comparatively small, oblong-ovate, solid, base
somewhat flattened; terminals conspicuous, sharply
formed; aperture straight, widening abapically; margins
apparent, though scarcely projecting; teeth fine, prom-
inent, extending across base and lip to margins; a fine,
white lateral surface at the margins separates the dorsum
from the base, dorsum pale grey-brown, with numerous
fine, light grey-white pustules; a grooved, lateral mantle
line bisects the right dorsum; terminals and teeth bright
burnt-orange; base and interstices off-white; columella
narrow, semi-concave; fossula long, narrow, more deeply
concave; columella and fossula ribbed adaxially with a
continuation of the base teeth.

32. Staphylaea (Staphylaea) limacina limacina
(Lamarck, 1810)

— Cate, 1966: plt. 39; figs. 36a, 36b —
Localities: 33, 44, 48, 52, 57, 59, 62 a, 75 b, 85

(7) u L W H Lip Col
Largest shell: 20:6 aS OF 20 20) eZ

Smallest shell: 20:8 llee OD i 15

This species, like the preceding one, is not very common
here. The larger shell, whose measurements are recorded
here was collected at Off Island. This is a large rock pile,
with accompanying reef, located on the outer periphery
of Buckner Bay on the Philippine Sea side of Okinawa.
The specimen was found on a +0.5 foot daytime tide,
with its purple-black mantle and the animal fully ex-
tended. The smaller shell was collected from under a rock
at the Onna tide flat at night on a +0.6 foot tide. Both
shells are typical examples of Staphylaea 1. limacina,
though of smaller size than the comparable Philippines
shells. An interesting observation regarding this species
that may clarify past confusion as to its identification is
that the shell seems to attain maturity through at least
three separate, distinct stages of development. The first
stage has a solid, smooth, dark brown shell; the middle
stage has numerous white spots on the smooth brown
dorsum; in the final stage, in completely adult shells, the
smooth white spots become white raised pustules, and a
grooved mantle line is added.

Shell elongately ovate, somewhat light in weight, base
convex; margins distinct, though not overly thickened;
terminals sharply produced; apex depressed; aperture
long, narrow, more or less straight; teeth distinct, finely
formed, short, not extending to margins, sometimes even
shorter centrally on the base; dorsal color battleship grey,
marked overall with numerous white spots which may or
may not be pustulated; a grooved mantle line usually
present; margins, base and lip are white; terminals deep

Page 34

burnt-orange, teeth and interstices pale orange, each den-
ticle peculiarly, characteristically outlined on either side
with dark burnt-orange lines; columella and shallow
fossula ribbed by extension of “base teeth.

33. Staphylaea (Nuclearia) nucleus nucleus
(Linnaeus, 1758)

— Cate, 1966: plt. 39; figs. 37 a, 37b —
Localities: 13, 15, 41, 47, 83, 85

(iO) - w L WwW H Lip Col
Largest shell: 2047 124 ON 2a eld
Smallest shell: sy NOx) 2 Pile}

An uncommon species in the Ryukyu Islands, it has
been collected at Zampa-Misaki and Metasaki. Beach
drift shells have also been found at Gushichan and Chi-
nen. MacNei (1960, p. 51; plt. 19, figs. 5, 6) reports
finding a fossil shell of this species in the Pleistocene
limestone of Yontan.

Shell of medium size, ovate, granular-pustulate, strong;
terminals produced, semi-beaked; aperture narrow, curv-
ing; base and margins thickened, the latter upswept; teeth
fine, long, sharply defined, numerous, extending from
aperture edge to either margin and up onto sides; grooved
mantle line originates from a small, weak umbilical de-
pression adapically; shell color uniform, except that dor-
sum reflects a light grey; granular surface, terminals,
margins, base, teeth, and interstices medium beige.

34. Erronea (Adusta) onyx onyx (LinNAEUS, 1758)
— Cate, 1966: plt. 40; figs. 38a, 38b —
Localities: 19, 69, 75b

(2) r L WwW H Lip Col
Largest shell: 200 BiB lsh - AO) is

Smallest shell: 325 MG WO Wo ily

This species, although occasionally found on tide flats,
must be considered rare in the Ryukyu Islands at present.
The two specimens listed here were found at Taguchi;
another shell has been reported from Henza-Miyagi. Of
this latter shell Bernice Albert says: “I found the only
one of these that I know of having been found at Okinawa.
It was stranded on a sandy tide flat here at Henza- Miya-
gi. I had the feeling it did not come from the sandy
area and decided to investigate further, but the water
toward the reef [was] so full of long-spined urchins and
the water was so shallow to swim in we had to give up
trying. We’ve looked in this area since and have never
seen another. The animal was jet black and the animal
sort of bubbly.”

THE VELIGER

Vol. 10; No. 1

Shell medium sized, pyriformly-ovate, somewhat light
in weight; terminals prominent; aperture straight, wide;
margins well calloused, angled; teeth short, finer on
inner lip, extending weakly onto broad columella, fossula
interrupting the teeth that cross it, labial teeth larger,
well formed; base swollen; dorsal coloring a succession of
various shades of light brown (the characteristic light
grey-white not present) superimposed upon one another,
exhibiting the effect of mantle action; narrow band of
the sides, margins, terminals, base, and interstices dark
brown, teeth red-brown; interior of shell off-white.

35. | Erronea (Gratiadusta) pulchella pulchella
(Swainson, 1823)

— Cate, 1966: plt. 40; figs. 40a, 40b —
Localities: 27, 39, 71

(1) r L W H Lip Col
Shell measurements: 41.3 2310 2010) eS 1ee27

During the last few years trawling operations by fisher-
men in the Taiwan-Ryukyu Islands area have confirmed
the occurrence of this species in these waters. There have
been reports of collections off Iriomote Island, though this
has not been substantiated. The specimen reported here
was trawled in 48 fathoms off the island of Miyako,
southern Ryukyu Sakishima Group; this is, to my know-
ledge, the first authenticated shell of this species from
the Ryukyu Islands.

Shell large for the species, pyriform, inflated adapi-
cally; terminals prominent, sharply formed, front almost
beaked; aperture straight in front, curving sharply left
to the rear; teeth fine, long on the base, short on outer
lip; margins barely thickened, subangled on the right;
basic dorsal color whitish-beige, with small, scattered
flecks of light-brown; white margins numerously spotted
with medium brown spots, base and interstices off-white,
teeth red-brown (cracks and condition of shell indicate
the specimen may have been dead when trawled from
the ocean bottom).

36. Erronea (Erronea) errones errones (LINNAEUS, 1758)
— Cate, 1966: plt. 40; figs. 42a, 42b —
Localities: 1, 19, 33, 39, 48, 51, 57, 65, 69 a, 75b, 81, 85

(14) c L W H Lip Col
Largest shell: 26.2 14.0 115 > 17 ale
Smallest shell: 18.3 10.0 810° 15s

— ————————————————————E—E

This species is common and widely distributed. It is
almost never found out in the open, at least in the day-
time, but is seen under rocks and in dark coral crevices.

Vol. 10; No. 1

Shell elongately ovate, sub-pyriform, strong, lightweight
in structure ; dark brown apex visible in shallow umbilicus,
base convexly swollen; terminals somewhat prominent;
margins thickened, right side particularly so; aperture
wide, becoming wider in front; teeth weak on both lips,
often nearly absent to the rear on the base; columella
perceptibly denticulate in front, fossula absent; dorsum
pale greenish-grey, tribanded, over which are numerous
fine flecks of lighter brown, while usually a larger dark
brown blotch adorns the central area; terminals, margins,
base, teeth, and interstices light to dark ivory.

37. Erronea (Erronea) ovum ovum (GMELIN, 1791)
— Cate, 1966: plt. 40; figs. 43a, 43b —
Localities: 7, 82

(5) u IL, W H Lip Col
Largest shell: 2510 ela Sil 2:27 alee

Smallest shell: 23.6 140 116 18 #416

The two shells listed here were collected at Baten,
on the southern rim of Buckner Bay. This species has
also been collected at Onna on the south (China Sea) side
of Okinawa. An interesting feature of the Ryukyu shells
is the very pale yellow coloring of the interstices; usually
it is much darker and a more obvious identifying char-
acter. The shells of this species are more pyriform than
those of Erronea e. errones, lack the brown dorsal blotch,
base and terminal coloring is darker, and the aperture
is wider, but more constricted abapically.

Shell pyriform, somewhat bulbous; terminals prominent ;
aperture straight, wide, openly constricted in front; mar-
gins calloused, semi-angled on the right side; teeth short,
fine on the inner lip, larger, heavier on outer lip; columella
smooth, fossula almost flat, ribbed; dorsal surface covered
evenly with small, irregular pale brown spots (flecks),
undercoloring off-white, creating a greenish appearance;
margins, terminals, base, and teeth light beige, interstices
very pale yellow.

38. Erronea (Erronea) cylindrica cylindrica (Born, 1778)
— Cate, 1966: pit. 41; figs. 46a, 46b —
Localities: 37, 45, 57, 75 b, 79, 85

(4) ou i We En Rip! ol
Largest shell: BB Wi WOW iy Be
Smallest shell: Beads ND BRS 16520

Although these shells are found more commonly at the
southern islands of Miyako and Yaeyama, they are also
collected on the tide flats at Onna, Kue, and Zampa-

THE VELIGER

Page 35

Misaki. The narrowness of these shells seems, more than
usual, to emphasize their length.

Shell long, narrow, cylindrical, somewhat lightweight;
terminals protrude awkwardly; aperture straight, wide,
openly constricted in front; left margin barely thickened
or not at all, and perceptibly flanged abapically; right
side narrowly thickened and angled; umbilicate, with
black apex therein; dorsal color grey, irregularly blotched
with chestnut-brown on top; large brown spots at each
terminal (quadrimaculate) ; umbilicus, terminals, mar-
gins, base, teeth, and interstices off-white; columellar
teeth long, fine, weak, heavier in front, crossing fossula;
labial teeth short, strong, widely separated.

39. Erronea (Erronea) caurica caurica (LINNAEUS, 1758)

— Cate, 1966: plt. 41; figs. 44a, 44b —
Localities: 39, 41, 57, 85

(2) r L W H Lip Col
Largest shell: of) I IG BD. 1@
Smallest shell: od = FOO NES MS. AY

This species is very scarce here, only 3 specimens at
present being known from Okinawa. Two of these were
found at Zampa-Misaki, one by a SCUBA diver in deep
water, the other at nearby Bolo Point; the third specimen
was picked up on the Onna tide flat by Ernest and Bernice
Albert.

Shell long, narrow, with rounded base; terminals prom-
inent, apex depressed; aperture wide, generally straight;
teeth strong, bold, though finer on rear half of base;
margins well calloused, more so on right side, angled and
shouldered; basic dorsal color greenish-ivory, with 3 wide
transverse bands of light brown, overall thickly flecked
with light brown; large brown spots line both sides;
margins, terminals, base, and teeth light beige, interstices
brownish-beige.

40. Erronea (Melicerona) felina pauciguttata
(SCHILDER & SCHILDER, 1938)

— Cate, 1966: plt. 41; figs. 47a, 47b —
Localities: 9, 21, 33, 41, 43, 57, 81, 85

(4) u 1b, W H Lip Col
Largest shell: 18.9 11.0 hg NE I

Smallest shell: 15.0 8.2 6.0 12 14

This species has been collected intertidally at several
locations at Okinawa; the most notable of these are
Zampa-Misaki, Onna, Metasaki, and Yagaji. It may be

Page 36

THE VELIGER

Vol. 10; No. 1

found fairly frequently at the offshore islands of Ikishima
(on the reef flats), and Minami-Ukibaru (intertidally).

Shell oblong-ovate, with a suggestion of flatness; ter-
minals obscure; aperture straight, comparatively wide;
right margin thickened, angled; base flatly concave; teeth
short, weakly developed on the rear half of base, stronger
in front, prominent on outer lip; fossula flat, ribbed;
dorsal color grey-green, overlaid with 4 (3 prominent)
broken transverse bands of darker color, with a yellow
wedge of color just to the rear of the front terminal
collar; large, dark brown spots at either side of terminals
and in a line along the right margin, scattered along the
left margin; base, lip, teeth, and interstices pale ivory.

41. Notadusta punctata atomaria (GMELIN, 1791)
— Cate, 1966: plt. 41; figs. 48a, 48b —
Localities: 39 a, 41, 57, 65, 85

(8) u L WwW H Lip Col
Largest shell: 186 11.1 3 A Zl

Smallest shell: 14.6 8.4 68 19 18

To the best of my knowledge this is a rare mollusk in
Ryukyu waters. The specimens measured were collected in
1950 on the reef just north of Onna on the west coast,
and at Metasaki in 1952 by Anita Scott. Shells also are
known to have been collected at Zampa-Misaki.

Shell generally fairly large, pyriformly-ovate, light in
weight, umbilicate; terminals produced; aperture wide,
curving; margins barely formed on the left, thickened,
minutely shouldered, and angled on the right, semi-flanged
in front; shell color off-white to ivory-beige, teeth and
terminal openings slightly darker beige; dorsum and mar-
gins numerously speckled with medium dark brown spots.

42. Notadusta katsuae (Kuropa, 1960)

(Plate 3, Figures 1a, 1b)
Localities: 53, 62

(2) r L WwW H Lip Col
Largest shell: 7AM)3) INNO) hich S10) 28)

Smallest shell: 20.1 9) SO) S124

Kuropa cites only “Okinawa” as the type locality of this
relatively newly discovered species. The fact that it was
trawled in 80m (presumably by fishermen) precludes
the fixing of an exact locality for it. Fernando Dayrit,
Diliman, Rizal, Philippines reports that practically all of
the fishing for Manila markets is done off the eastern
coast of Palawan Island, a possible general locality for this
specimen. SCHILDER (1963, p. 127) records the holotype

as in the Teramachi collection (Kyoto). The smaller shell,
whose measurements are listed here (Cate coll. no. C3392)
was taken from the stomach of a fish in a Manila fish
market, thus the exact locality for this shell is also un-
known. It is conceivable that a migrating fish transported
this specimen to the Philippine area from Japanese waters.
Dr. Schilder (zn litt.) mentions the species as coming from
the Sulu Sea also.

Shell narrow, elongately ovate, thin, lightweight though
strong; base convexly rounded, swollen; terminals beaked
delicately though strongly, sharply defined; aperture long,
narrow; inner columella narrow, smooth; fossula almost
without depression; teeth numerous, very fine, somewhat
long on lip, very short on inner edge of base; margins per-
ceptibly thickened, more so on outer lip where sub-angled,
shouldered; slightly flanged at either side in front; primary
shell color dark beige, tri-banded with a darker shade
dorsally, terminal beaks lighter color, margins covered
thickly with various sized chestnut-brown spots, narrow
base and interstices ivory-beige, teeth ivory within, chest-
nut-brown on the ridges without.

It should be emphasized that my description was made
from an ex pisce specimen, although one in excellent
condition; there is none of the dorsal spotting as seen in
Notadusta musumea.

43. Notadusta musumea (KuropA & Hane, 1961)
(Plate 3, Figures 3a, 3b)

Locality: uncertain in Ryukyu waters

(1) r L WwW H Lip Col
Shell measurements: 22:3) 1 125) OCS Oem

This is a rare species, occurring in Ryukyu and southern
Japanese waters. The specimen whose measurements are
recorded here was trawled in 50 fathoms off Kochi, Shi-
koku, Japan (in the Tosa Sea). The species has also been
obtained by trawling off Amami-O-Shima, in 40 fathoms.
However, I have been unable to get any further infor-
mation about the Ryukyu shell. Scuitper (1963) believes
this species to be synonymous with Notadusta katsuae
(Kuropa, 1960), but it appears to me to be distinct. I am
tentatively listing N. musumea with the Ryukyu fauna
until further field work brings the information on the
species here into better perspective.

Shell light-weight, strong, pyriform, shallowly umbili-
cate; base broad, convexly swollen; terminal in front
semi-beaked, obscure at rear except that right extremity
of lip is flared; aperture straight, wide; columella smooth
adapically, increasingly ribbed toward the front; fossula
long, narrow, shallow, ribbed, prominently denticulate

Vol. 10; No. 1

THE VELIGER

along either side; teeth numerous, very fine; though
distinct, interstices are shallow; teeth larger on lip, finer
and more indistinct on base; left margin rounded, scarcely
calloused, thickened, broadly upswept on right side, weak-
ly shouldered; primary shell color light ivory, with a
faint, narrow band transversely bisecting the dorsum;
margins thickly flecked with medium-brown spots, the
left side more thickly so, the dorsum is more sparingly
flecked, somewhat methodically with very small brown
spots; a large chestnut-brown spot covers a portion of
apex and umbilicus which latter is partially obscured
with opaque nacre.

44. Palmadusta (Palmadusta) asellus vespacea
(MEtvitt, 1905)
— Cate, 1966: plt. 42; figs. 50a, 50b -
Localities: 7 a, 9, 33, 41, 57, 85
(8) u L WwW H Lip Col

Largest shell: Wigs i031 0) AN A
Smallest shell: Sid) 8.1 2 I) es

This species is often found living on coral ledges and
under rocks at Zampa- Misaki, Bolo Point, and Metasaki.
The animals seem to migrate more or less seasonally;
one or two may be found on a collecting trip, then it
may be a long time before another is seen. MAcNEIL
(1960, p. 52; plt. 18, figs. 19, 22) lists and illustrates a
shell from the Yontan limestone of Okinawa that may
belong to this species, though to me it seems more probably
to be Erosaria poraria scarabaeus (Bory, 1827) (see Cate,
1966, plt. 39, figs. 32a, 32b). The shape of the shell, the
lack of teeth on the front half of the columella, and
the presence of an exaggerated umbilical area all tend to
eliminate Palmadusta asellus vespacea from consideration
here.

Shell oblong-ovate, solid, strong, umbilicate; terminals
somewhat produced; aperture fairly narrow, straight;
margins thickened, heavily so on the right side, upswept,
shouldered, angled; primary shell color white; three wide
dark brown bands traverse the dorsum from right side
to adaxial edge of columella, bands partially obscured
by thin layer of white callus on left margin and base;
umbilicus, terminals, margins, base, teeth, and interstices
white.

45. Palmadusta (Palmadusta) clandestina moniliaris
(Lamarck, 1810)

— Cate, 1966: plt. 42; figs. 51a, 51b —
Localities: 14, 23, 25, 33, 37, 57, 59, 62 a, 65, 85

Page 37

(5) u L W H Lip Col
Largest shell: 15.7 8) GS 7 ely
Smallest shell: i3%2 7.8 OOS Wye AG

This species is far from common. At Imbu it has been
collected under rocks, at Onna out on the tide flats. It has
also been obtained at Kue and Zampa- Misaki.

Shell narrowly ovate, small but strong; terminals prom-
inent ; aperture nearly straight, curving gently adapically;
margins thickened, semi-angled on the right side; teeth
small, distinct, short on outer lip, longer on the base;
primary shell color white, overlaid dorsally with light
grey-beige, bisected by a narrow transverse band of off-
white; a yellow-beige spot toward the front, overall fine
straw-colored lines mark a zigzag pattern on upper sur-
face; terminals, margins, base, teeth, and interstices white.

46. Palmadusta (Palmadusta) lutea lutea (GMELIN, 1791)
— Cate, 1966: plt. 42; figs. 54a, 54b —
Localities: 9, 53, 85

(1) r L W H Lip Col
Shell measurements: 13.5 8.0 6.2 16 14

As this paper was nearly completed, a personal commu-
nication from Peter W. Way, Okinawa, was received which
lists a species hitherto unknown in Ryukyu waters. It
states in part:

“Cypraea lutea (Gmelin) was found by myself yester-
day morning about 3:30 AM on the outer reef at Zampa-
Misaki (Bolo). To my knowledge another has not been
found on Okinawa before, at least there are none in any
of the collections that I have seen.

“I took it to the Alberts last night and we cleaned it
together. I wanted them to verify my claim, as I was by
myself at the time I found it. Naturally, we were all in
‘seventh heaven’ about finding this item.

“The tide for 27 January, 1967 was —0.5 ft. at 0154
hours. About 0430, I made one last quick sweep of the
dry outer reef and saw the lutea; it looked as if it had
just crawled out of a hole in the reef. When I came
upon the animal it was high and dry and was still moving.

“The animal: the foot was a bright orange-red; it had
minute black marks /// or lines on the dorsal side; the
mantle was a dark red (not intense) and had numerous,
simple papillae which gave it a fuzzy appearance; the
tentacles were also the same color as the foot, with black
eyes. (The mantle was not as brilliant a red as the foot.)

“Other shells found were Cypraea hirundo neglecta, C.
helvola, C. cribraria, C. arabica, C. teres, C. asellus, C.

Page 38

THE VELIGER

Vol. 10; No. 1

nucleus, C. punctata, C. maculifera, C. fimbriata, C. pora-
ria, C. isabella, C’. caputserpentis, C. carneola, C. vitellus,
C. vanelli (lynx), C. erosa, C. pallidula, Conus geogra-
phus and Conus distans. Another fellow found two Cyp-
raea punctata and one C. scurra.”

47. Palmadusta (Palmadusta) ziczac ziczac
(Linnaeus, 1758)

— Cate, 1966: plt. 42; figs. 55a, 55b —
Localities: 57, 65, 85

(1) r L W H Lip Col
Shell measurements: 11.1 10.6 86 16 15

Until about a year ago this species had been thought
extinct in Ryukyu waters. Since then one specimen has
been collected in shallow water on the reef just north of
the village of Onna; another has been found in the same
locality in deeper water by a SCUBA diver; still more
recently a specimen was collected at Bolo Point.

Shell short, wide, pyriformly ovate, umbilicate, with a
swollen base; terminals weakly produced; aperture wide,
gently curving left adapically; margin thickened only on
right side; teeth short, well defined, finer on the base;
basic dorsal color light grey, overlaid with light olive-
brown in a zigzag manner, thereby exposing the light
grey primary color; base, teeth, and interstices burnt-
orange; medium dark brown spots encircle the umbilical
area, numerous on base, less on sides; terminal edges
medium brown.

48. Palmadusta (Purpuradusta) gracilis japonica
(ScHILpDER, 1931)

— Cate, 1966: pit. 43; figs. 56a, 56b —
Localities: 19, 37, 43, 57, 75 b

(3) u L WwW H Lip Col
Largest shell: 18.0 10.6 86 16 13

Smallest shell: 15.0 912 Ue) ay

This is another uncommon species living in the inter-
tidal zones of Okinawa. It is most frequently collected on
the tide flats at Kue, Onna, and Henza- Miyagi.

Shell pyriform, semi-bulbous; terminals strongly pro-
duced; aperture wide, straight; right margin heavily
calloused, sub-angled; teeth short, fine; primary dorsal
color light grey-white, overlaid with numerous thin flecks
of light brown, a larger dorsal blotch frequently present as
well; large brown spots line both margins, both terminal
openings lined with purplish-brown; base, teeth, and
interstices ivory.

49. Palmadusta (Purpuradusta) fimbriata marmorata
(ScHROTER, 1804)

— Cate, 1966: plt. 43; figs. 58a, 58b —
Localities: 9, 19, 33, 37, 57, 65, 81, 85

GH) e LW Heel
Largest shell: 13.1 U2 59h eee

Smallest shell: 11.1 6.1 Oe is KS

This species is found commonly under rocks and in
coral at Kue, Onna, Yagaji, and Bolo Point. The shells
closely resemble the Philippine forms and may be a link
in the range of the species, which terminates in the Japa-
nese islands to the north.

Shell sub-pyriform, bluntly flattened adapically; ter-
minals obscure to the rear, less so in front; aperture
fairly wide; teeth very fine, weaker on base; right margin
thickened, angled, sub-shouldered; primary dorsal color
light grey, overlaid irregularly with pale brown, with a
wide, darker brown transverse broken band; apex and
terminals deep lavender; margins, base, teeth, and imter-
stices off-white.

50. Bistolida (Blasicrura) quadrimaculata quadrimaculata
(Gray, 1824)

— Cate, 1966: plt. 43; figs. 60a, 60b —
Localities: 39, 57, 67

(2) r L W H Lip Col

Largest shell: Zales OKO) 8.55) 1s Sas
Smallest shell: Adis 1 LOI 8:5 5 (189 aaly

This species must be considered rare until additional
field work can give us a true picture of its distribution m
these islands. One specimen was collected at Okinawa in
1955, but no records were kept of the specific locality at
the time. However, two more shells were collected by the
same worker at Machinato in 1956.

Shell long, narrow, sub-pyriform; terminals heavily
calloused, blunt; aperture wide, straight; teeth long and
fine on base, heavy and short on outer lip; primary color
off-white, dorsal area darker because of 4 wide, pale gray
transverse bands, all overlaid with minute flecks of light
yellow-brown; two dark brown spots at either end; base,
terminals, margins, teeth, and interstices off-white.

Dil. Bistolida (Blasicrura) pallidula palhidula
(Gasxoin, 1849)
— Cate, 1966: plt. 44; figs. 63a, 63b —
Localities: 41, 57, 81, 85

Vol. 10; No. 1

(2) u L WwW H Lip Col
Largest shell: Ailey Mey YD) BR ANZ
Smallest shell: 19.0 9.3 US 20) 1s}

This species lives on the reef flats, usually hidden in
rough, algae-covered crevices in about four feet of water;
it is not often found. The solidly formed shells seem to
have a thicker, more concentrated dorsal color than speci-
mens from elsewhere.

Shell cylindrically oblong, heavy, solid ; terminals heavily
calloused, weakly produced; aperture nearly straight,
narrow; teeth relatively fine, well defined; margins thickly
calloused, right side more so, sub-angled; basic color off-
white, dorsum overlaid with a thick covering of brownish-
yellow flecks (the large shell, whose measurements are
listed here, has a definite mantle line) that obscures the
4 broken transverse brown bands characteristic of this
species; terminals, base, teeth, and interstices off-white
to pale ivory.

52. Bistolida (Blasicrura) luchuana Kuropa, 1960
— Cate, 1963: plt. 15; figs. 2, 2a —
Localities: 14, 39, 45, 55, 57, 63 a, 65, 75a, 85

(23) ese L W H Lip Col

Largest shell: Zo Ne) Oe OD AD
Smallest shell: 16.1 9.4 Tod 2 Gs. ie

This is perhaps the only species endemic to the Ryukyu
Islands; as far as I have been able to determine, its range
does not extend as far as the Philippines or Japan. I have
examined over 75 specimens of this species and compared
them with Bistolida dayritiana (Cate, 1963), and am
convinced this latter species is not presently found in
Ryukyu waters, and therefore cannot be considered as part
of this molluscan fauna. At Onna B. luchuana is found
abundantly on the submerged tide flats, most often in
water just below low tide line under rocks, living in a
soft, thick-stemmed, green sponge of an unidentified spe-
cies. Bistolida luchuana, particularly at Onna, is associated
with such other cypraeid species as Ravitrona h. helvola,
R. 1. labrolineata, Erosaria erosa phagedaina, Staphylaea
5. staphylaea and S. 1. limacina. An interesting observation
is that many of the specimens of B. luchuana have jet-
black animals, while others seem to be lighter, grey. This
species has also been found at Okuma, and is never seen
exposed above water due to the receding tide, as are
species such as Monetaria moneta rhomboides, Ornament-
aria a. annulus, and Ravitrona c. caputserpentis; the usual
depth for this mollusk is from 3 to 7 feet. The species has
also been collected on the south east coast of Miyako Is-

THE VELIGER

Page 39

land (Way) ; ScHILpER (1962) reports a “‘semifossil’ shell
(22.5mm.) coming from the Sukiran dredgings off Okin-
awa (Col. McCarty, 1955).”

Shell pyriform, lightweight; terminals strongly pro-
duced; aperture fairly wide, mostly straight, curving
gently left; teeth fine, well defined, longer on rear half
of base, shorter in front, labial teeth of medium length;
margins well calloused, somewhat upswept and angled on
the right side; basic color light-grey, 4 very pale, wide
transverse bands of darker color, all overlaid with numer-
ous, fine flecks of pale yellow-brown; a faint mantle line
often present; apex depressed, dark brown; terminals,
margins, base, teeth, and interstices pale ivory.

53. Bistolida (Blasicrura) teres teres (GMELIN, 1791)

Syn.: Cypraea tabescens Dittwyn, 1817 (Sotanper MS)
Descr. Cat. Recent Shells 1: 463

Cypraea punctulata Hipa.co, 1907
Monogr. gén. Cypraea (M. Acad. Cien. Madrid 25: 484)

— Cate, 1966: plt. 44; figs. 66a, 66b —
Localities: 33, 42, 65, 71, 85

(5) u L WwW H Lip Col

Largest shell: 2 OIG Ee Se Ot. OO.
Smallest shell: 2S O-2) ml 2 O23 25

The shells whose measurements are recorded here were
collected on a shallow-water coral outcropping at Mina-
mi. Other specimens from Smuggler’s Cove, Zampa-Mi-
saki and from Taiwan would suggest a link with the
species in the Philippines and Japan.

Shell ovate to oblong-ovate, solid, heavily formed, base
swollen, apex depressed; terminals produced; aperture
fairly wide, curving slightly adapically; teeth fine, short
on base, crossing columella and long shallow fossula,
somewhat larger and longer on outer lip; margins heavily
calloused, angled and shouldered on right side, enveloping
rear terminal; primary dorsal color pale blue-grey, over-
laid with a thick pattern of light brown flecks, smudges,
and a large dark brown central blotch; widely dispersed
brown spots mark both margins; terminals, margins, base,
teeth, and interstices off-white.

Oar Bistolida (Bistolida) kieneri depriesteri
(ScHILpErR, 1933)
— Cate, 1966: plt. 44; figs. 67 a, 67 b; text fig. 1 -
Localities: 43, 53

(ger fw Ep ipa col

Shell measurements: 13.5 8.1 6:25 1S eel2

Page 40

The present specimen was found among some cowries
collected at Minami-Ukibaru reef on March 20, 1965.
At the same time and place were also found Melicerona
felina pauciguttata and Purpuradusta g. gracilis. As far
as is known this is the only record of this species at
Okinawa. It was collected from under a rock in about
four feet of water.

Shell small, sub-pyriform, somewhat flattened; termi-
nals inconspicuous; aperture straight, fairly wide; teeth
sharp, well defined, long on rear half of base, short and
restricted to columella in front, on outer lip slightly larger,
of even length, almost reaching to the margin; margin
thickened, sub-angled, callus very thin on the left side;
primary dorsal color off-white, overlaid with 3 large
dark grey areas separated by 2 irregular transverse yellow
bands; 2 large dark brown spots at each terminal (quadri-
maculate), small light brown spots dot the dorsum and
sides; terminals, base, margins, teeth, and interstices white,
except that some color pattern shows through the thin
nacre covering the base.

55. Bistolida (Bistolida) hirundo neglecta
(Sowerby, 1837)
— Cate, 1966: plt. 45; figs. 68a, 68b; text fig. 2 -
Localities: 7 a, 33, 39 a, 57, 62 a, 81, 85

(7) u L WwW H Lip Col
Largest shell: My les 94 Bil av

Smallest shell: 13.1 Wel) 6.1 195 15

Especially interesting is the fact that this species has
two distinctly different forms, noted in both Ryukyu
specimens and in others from West Australia (CaTE, 1964,
p. 19). Some are long, narrow, and delicate; others are
short, globose, and massive; strikingly different in appear-
ance. Typical dimensions of both forms are reflected in the
shell measurements recorded here. The shells seem to be
always either one size or the other, without intermediate
forms. In this instance, the larger shells were collected at
Yagaji, the smaller on the tide flats at Onna. There may
be other differences of a specific or subspecific character
between the two forms; these only a study of the soft parts
could clarify.

Shell may be relatively large or small, larger ones more
cylindrical, smaller shells more pyriform; terminals prom-
inent, apex depressed; aperture straight, narrow; margins
thickened, more so on the right side, upswept, subangled,
shouldered; teeth long, fine, numerous; columella and
fossula denticulate; dorsum grey, divided toward the rear
by a faint yellow transverse narrow band, with a larger
irregular yellow mark anteriorly, the whole loosely covered
with fine brown flecks; brown spots mark both margins, 2

THE VELIGER

Vol. 10; No. 1

larger dark brown spots at each terminal collar; terminals,

margins, base, teeth, and interstices off-white.

56. Bistolida (Bistolida) stolida stolida (LiNNAEUs, 1758)
— Cate, 1966: plt. 45; figs. 70a, 70b —

Localities: 7 a, 9a, 20a, 37, 63 a, 63b, 65, 75b, 81, 85

(55) rc L W H Lip Col
Largest shell: 28:5 154 “120 Re 22eee20
Smallest shell: EO 103 19 iG

In 1954-1955 this species must have been more plenti-
ful as I have at least 55 shells collected at Okinawa during
those years. On this basis I suggested (Cate, 1966, p. 263)
that the Ryukyu Islands seemed to be the locality of
greatest abundance for the species. Recent field experience
now indicates a far less common occurrence of this species.
Dredging in 1955 at Chatan (Sukiran) yielded many
unusually large subfossil specimens, reflecting the earlier
abundance of the species.

Shell cylindrically-rectangular, sharply angled, tapering
abruptly toward terminals; apex depressed; terminals
boldly exposed, semi-beaked; aperture gently curving
left; teeth prominent, fairly long on both lips, but short
on front half of base; margins thickened, especially so on
right side where margin is upswept, angled and should-
ered, with both margins becoming flanged near terminals;
basic dorsal color light grey to pale beige, with a large,
squarish chestnut-brown dorsal blotch; grey color bands
of irregular width and shape extend from the 4 corners
of the blotch to the margins, chestnut-brown stain covers
margins to the terminal collars; base, teeth, and inter-
stices white.

Sik Cribraria (Ovatipsa) chinensis chinensis
(GMELIN, 1791)

Syn.: Cypraea cruenta Dittwyn, 1817
Descr. Cat. Recent Shells 1: 460
-— Cate, 1966: plt. 45; figs. 71a, 71b —
Localities: 9, 17, 57, 85

(6) uo. LOW ©on “apie
Largest shell: 45.2 27:0, 19:8 19 Ri
Smallest shell: SOP 18'S 13:6.) 1920

This species has been found only at two Ryukyu locali-
ties, Zampa-Misaki and Hedo. Usually it has been col-
lected at night, although it was once found in the daytime
feeding on coral outcroppings and ledges. There seems to
be much variation in the size of the adult shells.

Vol. 10; No. 1

Shell ovate, heavy, strong, somewhat flattened; termin-
als inconspicuous, heavily formed; aperture fairly narrow,
curving gently; teeth fine, well defined on base, heavy
and bold on outer lip; right margin thickened, shouldered ;
callus covers most of right and left dorsum; dorsal area
light straw-brown, with numerous off-white lacunae and
an inconspicuous mantle line; sides, margins, most of base
and teeth light beige, covered with numerous, various
sized lavender spots; interstices bright orange.

58. Cribraria (Cribraria) cribraria orientalis
SCHILDER & SCHILDER, 1940
— Cate, 1966: plt. 45; figs. 72a, 72b —
Localities: 9a, 33, 59, 65, 85

(18) re ean aw LOH Walp) a Col

Largest shell: PESO) MAE NAO Ne} 920)
Smallest shell: 189 11.3 O33 ean Oge lS

Although several shells of this species have been col-
lected at various localities at Okinawa, it is, nevertheless,
far from common. The larger shell, whose measurements
are recorded here, was trawled off Kadena, ex Dr. T.
Shikama, probably in 1961 or 1962 (Albert).

Shell pyriform, bulbously-humped, solid, base swollen;
terminals produced; aperture wide, straight, abruptly
curving left adapically; margins thickened, more heavily
on right side, angled, shouldered, and vaguely pitted;
dorsum light yellow-brown, with numerous off-white la-
cunae of various sizes; terminals, margins, base, teeth,
and interstices pure white.

LITERATURE CITED

Cate, Crawrorp NEILL
1960. A new Hawaiian subspecies of Cypraea cernica SowER-
BY. The Veliger 3 (1): 3-7; plt. 1; figs. 1-5
(1 July 1960)

THE VELIGER

Page 41

Cate, CrawForp NEILL
1963. A new cowrie from west-central Philippines.
The Veliger 5 (4): 140 - 143; plt. 15; 1 text fig. (1 Apr. 1963)
The Veliger 7 (1):
(1 July 1964)
The Veliger 8(2): 45-61;
(1 October 1965)

1966. Philippine cowries. The Veliger 8 (4): 234 - 264;
pits. 32-45; 3 text figs.; 1 map. (1 April 1966)

1964. Western Australian cowries.
7 - 28; pit. 5; 1 map

1965. Hawaiian Cowries.
plts. 4- 10; 4 maps

Hase, TADASHIGE

1961. Coloured illustrations of the shells of Japan. Hoikusha
2: 1-183; pits. 1-66
TREDALE, Tom
1930. Queensland molluscan notes, No. 2. Mem. Qld.
Mus., 10(1): 73-88; plt. 9
Kuropa, ToKuBEI
1960. A catalog of molluscan fauna of the Okinawa Islands.

Univ. Ryukyus, 1960; pp. 1 - 106; plts. 1-3
MacNetx F. Stearns
1960. _—‘ Tertiary and Quaternary gastropods of Okinawa. Geol.
Surv. Prof. Pap. 339: 1-148; plts. 1-21
Nomura, SHICHIHEI & Kotora Hatat
1936. A note on the zoological provinces-in the Japanese Seas.
Bull. Biogeogr. Soc. Japan 6 (21): 207 - 214; plt. 13
ScHILDER, FRANZ ALFRED
1962. Note on Bistolida luchuana Kuropa.
1 (3): 39

1963. Zur Kenntni& der Cypraeidae. 6. Uber zwei seltene
Arten aus Japan. Arch. Molluskenk. 92: 123 - 130

The Cowry
(1 February 1962)

1965. The geographical distribution of cowries (Mollusca:
Gastropoda) The Veliger 7 (3): 171-183; 2 maps
(1 January 1965)

1966. The cowry Mauritia grayana Scuitper, 1930. Ha-
waiian Shell News 14 (7): 1, 8 -NS 77 -

Page 42

THE VELIGER

Vol. 10; No. 1

The Role of Behavior in Determining

the Intertidal Zonation of Littorina planaxis Puttiprt, 1847,

and Littorina scutulata GouLD, 1849

BY

CARL E. BOCK

AND

RICHARD E. JOHNSON

Department of Zoology, University of California, Berkeley, California

94720

(8 Text figures)

INTRODUCTION

Littorina planaxis Putiprt, 1847, and L. scutulata GouLp,
1849, are common upper littoral gastropods of the west
coast of North America. Littorina planaxis occurs higher
intertidally than L. scutulata, being found on the highest
rocks in the splash zone. The purpose of this study was
to investigate those behavioral factors which might limit
the vertical distribution of both species in the vicinity
of Bodega Head, California.

HABITAT PREFERENCE

Hewatt, 1937, and Ricketts « Carvin, 1952, describe
Littorina planaxis as a dominant species on the rocky
intertidal Zone I (above mean higher high tide) and L.
scutulata as characteristic of Zone II (between mean
higher high and mean lower high tides) . There is consider-
able variation in the tidal range of both species, but this
undoubtedly relates to the degree of exposure and con-
comitant wave splash in the individual study areas. Littor-
ina planaxis occurs almost exclusively on those exposed
rocks which are wetted only by splash, and which are free
of macroscopic algae (HEwatrt, op. cit.; DAHL, 1964). The
habitat of L. scutulata appears more variable. DAHL and
Hewatt report it in beds of algae and also occasionally
in beds of Balanus glandula Darwin, 1854.

In order to determine the distribution of the two
littorines at Bodega Head, we conducted counts along
two transects running from the high intertidal down

through the mid-intertidal region. This spanned the ranges
of both species. The results for adults and for young are
plotted in Figure 1. The number of individuals was
counted in each square meter along the transects, and
these counts were combined into twelve equal groupings
such that the two transects, although of different lengths,
could be represented in a single graph.

Adult Littorina planaxis clearly occupy a higher posi-
tion than L. scutulata (Figure 1). We found L. planaxis
almost exclusively on exposed rocks wetted at high tide
by splash and the larger waves. Littorina scutulata were
most abundant in beds of macroscopic algae, especially
Pelvetiopsis limitata (SETCHELL) GARDNER, 1910, but
they were also common in high tide-pools. These pools
were usually rimmed with Gigartina papillata (C.A.
AGARDH) J. G. AcArDH, 1846, and it was here that L.
scutulata congregated. The lower limits of L. scutulata
came abruptly in each transect as the relatively exposed
beds of algae gave way to deep pools or channels.

Curiously, almost no small Littorina planaxis were
found, and yet the small L. scutulata greatly outnumbered
the adults (Figure 1). Also, we found many young L.
scutulata occurring higher up than the adults of the same
species (cf. data in Figure 1). They were found commonly
on exposed rocks along with adult L. planaxis. They were
usually in small crevices in the rocks or packed into old
Balanus glandula tests, as reported by HEwatt, op. cit., for
the Monterey coast. However, in contrast to the findings
for the Monterey region, adult L. scutulata were not com-
mon in barnacle beds at Bodega.

Vol. 10; No. 1

THE VELIGER

Page 43

Adults

Number of Individuals per Square Meter

d
Z

ad NAAAAAAQAAAAAAAAARAAAAAAANNT

<«——— Mid-Intertidal ——> < Hich Intertidal >

Littorina planaxis >TV 1<— Littorina scutulata

Littorina planaxis

5
<7
2}
gq

oO

SYS. Ww np, Fe"

Baa] NAQAAQAAAAAAANAANAN NINN Be

Y
j
g
y
Z
Z
Z
Z
Z
j
]
y
j
j
Z
y
y
Z
y
j
g
q

PISSING
INWSSWIWAMA

10

<«— Mid-Intertidal ———> <— High Intertidal —

Exposed Rocks Algal Beds (1) Pools

Littorina scutulata

Adults Young Adults Young
rocks 394 21 51 1345
pools 16 1 83 259
algae 3 1 250 3011
n = 413 n = 23 T3042 Ol)
Figure 1

Distribution of Littorina in the rocky intertidal area

Three types of factors could be involved in the zonation
and habitat preferences of Littorina planaxis and L. scu-
tulata. First, the physiological tolerances of the species to
one or more physical parameters of the environment
might act as limiting factors to distribution. Second, biotic
characteristics such as predation, competition, and avail-
able food could cause one region to be preferable to
another. The next section summarizes the available data
on physical and biotic factors which may influence the
distribution of the two littorines. While these biotic and
abiotic factors must be the ultimate causes of the distribu-
tion patterns, it is possible that the behavior of the littor-
ines prevents their movement into those areas where con-

ditions would be unfavorable. Our research centered on
the third possibility and the results are presented in
“Behavioral Limits to Distribution.”

POSSIBLE ENVIRONMENTAL FACTORS
LIMITING DISTRIBUTION

Desiccation: Since Littorina scutulata does not occur as
high intertidally as L. planaxis, the possibility exists that
the upper limits of the former might be a function of
greater vulnerability to temperature or desiccation, or
both. Bow.us, 1966, studied water loss in L. planaxis and

Page 44

L. scutulata; all animals were held in dry containers at
temperatures varying from 29.4°C to 32.2°C. At 100
hours L. scutulata had begun to die and by 127 hours
70% were dead. In the same time period, none of the L.
planaxis had died, and they lost much less body water.
Littorina planaxis secretes a mucous seal between the shell
and substrate when exposed to dry conditions. We did not
observe this response in L. scutulata, but it undoubtedly
would help prevent water loss. HEwATT, op. cit., recorded
100% survival of L. planaxis above the highest wave
splash for at least 64 days. BARKMAN, 1955, gathered
similar data for many littorines and was able to correlate,
in a general way, desiccation tolerance with the degree
of periodic exposure experienced by each species.
Although it appears that Littorina planaxis is more
resistant to high temperatures and dry conditions than is
L. scutulata, we shall show that the behavior of the latter
protects it from these circumstances.
Submersion: RicKETTS & CALVIN, op. cit.,state that Lit-
torina planaxis will “ultimately drown” if held under water.
Many species of high intertidal periwinkles, including L.
planaxis, have relatively degenerate ctenidia and a vascu-
larized mantle epithelium, both of which favor oxygen ex-
change in air (Nicot, 1960). Thus drowning or at least
respiratory inefficiency under water may be a factor cont-
rolling the lower limits of distribution in L. planaxis.
Fresh Water: Since Littorina planaxis occurs higher than
L. scutulata, and since considerable rainwater can accu-
mulate in the high intertidal, it might be expected that
the former species would be more tolerant to fresh water.
In the laboratory, we found only one percent mortality
in L. planaxis after 3 days submersion in tap water
(n= 100), while 65% of the L. scutulata died in 2 days.
Seventy-five percent of these had burst from their shells,
presumably due to osmotic stress. Water circulation was
maintained during the experiment. BARKMAN, op. cit., in

THE VELIGER

Vol. 10; No. 1

a survey of other littorine studies, found that species which
occupy the upper zones tend to have greater fresh water
tolerances.

Hypersalinity: In the summer, the highest tide-pools at
Bodega Head become hypersaline between spring tides
due to evaporation. Although we have no experimental
data regarding tolerances, littorines are absent from many
of these pools. We have found Littorina scutulata common
in pools with salinities up to 104% sea water, only rarely
in those of 110%, and not at all in pools of about 140%.
Competition and Predation: The behavioral experiments
described in the following section suggest that competition
is not the immediate cause of behavioral zonation, since
both Littorina planaxis and L. scutulata selected their
respective habitats in the absence of one another. Never-
theless, this does not rule out the possibility that competi-
tion in the past has resulted in selection for such behavioral;
responses.

Littorina planaxis occurs above the predatory snail

Thais emarginata (DEsHAYES, 1839) in the intertidal,
but there is no evidence to suggest that this results from
predatory exclusion.
Food: Daut, op. cit., noticed that Littorina planaxis
occurred largely on high exposed rocks with encrustations
of microscopic algae, while L. scutulata were common at
lower levels in beds of macroscopic forms; however, it
remains unclear whether this apparent difference in food
preference is a cause or result of differences in zonation.
Dau did find that L. planaxis were unwilling or unable
to feed on the large algae unless the fronds were chopped
into small pieces; this suggests that food might be a
zonation factor, at least for this species.

Table 1 summarizes those environmental factors which
may be involved in controlling the limits of distribution
in the two littorines.

Table 1

Possible Environmental Factors Controlling the Distribution of Littorina

Upper Limits

Littorina planaxis

2. lack of algae as food above splash zone

Littorina scutulata 1. desiccation on exposed rocks

2. fresh (rain) water on exposed rocks

1. permanent desiccation above splash zone

Lower Limits

1. possible respiratory inefficiency during
submersion

2. possible lack of microscopic algae as food

3. possible predation

4. possible competition

1. possible lack of suitable foods
2. possible respiratory inefficiency during
submersion

3. hypersaline waters of highest pools
4. lack of macroscopic algae for food

ee —— — — — i —<“<_—_—<—<<<<<_.0 >> —_na—qo

Vol. 10; No. 1

BEHAVIORAL LIMITS TO DISTRIBUTION

Field Studies

In attempting to evaluate the role of behavior in the
distribution of Littorina planaxis and L. scutulata, we
carried out a field experiment at Doran Beach involving
transplantation of marked animals. Region 1 (Figure 2)
encompassed a small tide-pool rimmed with Gigartina,
and ran up about 2 m onto an exposed rocky shelf. Littor-
ina scutulata were common in and around the pool while
L. planaxis occurred on the rock face above. We placed
75 L. planaxis and 50 L. scutulata (marked with red or

Number of Individuals

3 j a
Ye
Be Yj af
ie y i
Qe
yo
Ge

J ] Littorina planaxis
| Littorina scutulata

THE VELIGER

Exposed Rocks
Algal Beds

Page 45

yellow enamel) in the pool and equal numbers up on
the rocks. This is the situation represented at day one in
Figure 2. Region 2 (Figure 3) was similar except that the
exposed rock face gave way to an extensive bed of Pelveti-
opsis instead of a tide-pool. Also, region 2 experienced a
much greater wave splash during high tides.

Since our habitat work showed that Littorina planaxis
was common on exposed rocks while L. scutulata occurred
in pools and beds of algae, we felt that the outcome of this
experiment would indicate the possible role of movement
in zonation. If the littorines segregate according to habitat,
those L. planaxis placed high should remain there, while
those placed in pools (region 1) or algae (region 2) should

Days 4 5 6

’ A: Animals marked red

B: Animals marked yellow

Figure 2

Doran Beach. Habitat selection study, region 1

Page 46

140

120

100

80

60-

Number of Individuals

40

20-

ee j
g
G
y
Y

[ ] Littorina planaxis
| Littorina scutulata

—

THE VELIGER

i)
K

Exposed Rocks
Algal Beds

Vol. 10; No. 1

LESS

XG

Ree )
Pa

D

Days 4 5

A: Animals marked red
B: Animals marked yellow

Figure 3

Doran Beach. Habitat selection study, region 2

move up to the exposed rocks. The reverse would be
expected for L. scutulata. Results (Figures 2 and 3) show
that this is essentially what happened, and we conclude
that behavioral zonation occurs.

Littorina planaxis moved up rapidly in both arcas, al-
though a small percentage was apparently swept away
before becoming established. The behavioral response of
L. scutulata was not as immediate. Also, a larger per-
centage was washed away; both those placed on exposed
rocks and those in pools or algae were lost. It is highly
significant that the downward trend of L. scutulata was
slower in region 1 than in region 2 (cf. Figures 2 and 3).

We attribute this to the combined effects of voluntary
downward movement and involuntary movement due to
heavier wave shock in the latter region. Littorina scutulata
were casily dislodged from the high rock by wave action;
they then tumbled down into the Pelvetiopsis. The res-
ponse in region | represents largely behavioral movement
alone, as this region was protected from strong wave
action.

We next conducted a series of experiments at Bodega
Head, designed to analyze in more detail the responses
of Littorina when placed in a variety of habitats. These
experiments are represented in Figures 4 through 7, and

Vol. 10; No. 1

involve distances traveled by each species in relation to
the tide cycle. Each species was tested separately in order
to prevent possible interactions. Snails were marked
with enamel and counted periodically during the trial
without being removed from the substrate.

1. Thirty Littorina planaxis and 42 L. scutulata were
placed about 35 cm above the surface of a high tide-pool
on an exposed rock (Figure 4). Although L. planaxis

501 ----0 Littorina planaxis

——® Littorina scutulata 27

30

Distance from Starting Point (cm)

- 20

Water
Level

Time (hours)

Horizontal Line: X Vertical Line: Range Rectangle: + 1 s.d.

Figure 4

Movement of Littorina placed above a high tide-pool

moved in all directions (range: -33cm to +56cm), they
averaged 16.8 cm higher on the rock after 22 hours. In
L. scutulata there was an obvious downward movement,
averaging 25.3 cm below the starting point after 24 hours.
This position was only about 7.6cm above water level; the
majority of the snails came to rest on the Gigartina around
the lip of the pool. This difference is highly significant at
the 0.01 probability level, using a two-sided “t’’-test for
differences of means, and indicates a movement of L.
scutulata down towards the pools in the high intertidal.

2. Figure 5 shows the results of a related experiment
in which 70 Littorina planaxis and 42 L. scutulata were
set in the bottom of a tide-pool. All animals in the pool
were given the value zero cm. After 48 hours, L. scutulata

THE VELIGER

Page 47

100} ----0 Littorina planaxis
— ® Littorina scutulata

50

Distance above Pool Level (cm)

Water
Level 0

A
High High Tide High High Tide

02 10 20 30 40 48
Time (hours)

Horizontal Line: ¥ Vertical Line: Range Rectangle: + 1 s.d.

Figure 5

Movement of Littorina placed in a high tide-pool

averaged 7.26 cm above pool surface, very comparable
to the position of this species in the first experiment;
during the same period, L. planaxis moved up an average
of 32 cm. Again, L. scutulata selected the pool rim, while
L. planaxis moved out on the exposed rocks. It is signifi-
cant that movements ceased when the rocks were dry
between higher high tides (Figure 5).

3. In the two preceding experiments, we demonstrated
behavioral differences between the two littorines in the
high intertidal. The final two experiments were conducted
at the lower limits of distribution. First, we placed 45
Littorina planaxis and 55 L. scutulata on a rock slope
with scattered algae. This was within the normal range of
L. scutulata. The results (Figure 6) show a significant
(to 0.01 level) upward movement of L. planaxis when
compared to the other periwinkle. We interpret this as
evidence of L. planaxis seeking its preferred, higher, inter-
tidal position. Note here that the movement occurred
between high tides rather than during them. The snails
clung firmly in place during the periods of wave shock.

4. The final experiment was carried out at a level
comparable to that of the last, but on an exposed point
subject to very heavy splash and surge; macroscopic algae
were lacking. Here both species moved up to a higher

Page 48

1254 --.-- © Littorina planaxis

——e Littorina scutulata

_ Co)
2 4
FZ
z
5
au -
pia, od a ee
Fe 29
&£
S Cie ci ENG pe
€
$
a) High
High Tide
—50 ae
0 3 10 12.0 25

Time (hours)

Horizontal Line: ¥ Vertical Line: Range Rectangle: + 1 s.d.

Figure 6

Movement of Littorina placed in a bed of macroscopic algae

position (Figure 7) ; there was no significant difference in
means at the 0.05 level. This area, then, was apparently
below the preferred habitat of either species.

We noticed in the Doran Beach study that Littorina
scutulata were more easily dislodged than L. planaxis.
Data from the last series of experiments support this and
suggest two ways in which the latter is better adapted.
First, when L. scutulata become dry on an exposed rock,
instead of fastening on with a mucous seal as L. planaxis,
they come to lie with opercula facing up and the shell
attached by only a thin strand of mucus. In this position,
L. scutulata are easily dislodged. Splash thus accounted
for a considerable proportion of the downward movement
of L. scutulata in the first experiment (Figure 4).

Second, in the two experiments concerning exposed
lower areas, 13% of 69 Littorina planaxis were swept
away while 41% of 79 L. scutulata were lost. Since these
areas were continuously moist, the snails did not withdraw
into their shells, but simply clung to the rocks. It would
appear that L. planaxis is better able to do so, although
NortH, 1954, found relative foot size to be comparable in
both specics. NortH found that greater numbers of L.

THE VELIGER

Vol. 10; No. 1

planaxis were lost from a rock which he washed through
a tide-pool. Our data, collected under more natural con-
ditions, suggest a greater resistance to wave shock in L.
planaxis — certainly an essential adaptation for life on
exposed rocks.

In summary, Littorina planaxis and L. scutulata appar-
ently occur in their respective habitats largely because of
differences in behavior.

We found Littorina planaxis will move up out of pools
or beds of macroscopic algae; thus these constitute the
lower limits of their distribution. WiLuiaMs, 1950, found
he could induce L. planaxis to move higher than they
normally occur in the field by artificially wetting the sub-
strate. This suggests that these snails will move as high as
there is wave splash. WiLLIAMs reports that L. planaxis

----- © Littorina planaxis

125

——e Littorina scutulata

100

Distance from Starting Point (cm)

4
Low High Tide

Hieh
High Tide
QO 2 4 10 20 26
Time (hours)

Horizontal Line: x Vertical Line: Range Rectangle: + 1s.d.

Figure 7

Movement of Littorina placed on a low intertidal rock

will move back down from a high position if they have
experienced a period of desiccation. Thus it may be that
populations fluctuate with the high tide levels. Littorina
scutulata avoids both high rocks and lower areas by selec-
ting pools and beds of algae in the mid-intertidal.
Evans, 1961, attributed behavioral zonation in Littor-
ina punctata FERRUSAC, 1821, to a combination of geotaxis

Vol. 10; No. 1

THE VELIGER

Page 49

and orientation to the shore. BARKMAN, op. cit., and
NEWELL, 1958a, 1958b, considered gravity, light, tidal
fluctuations, and preference for certain algae to be in-
volved in the zonation of L. obtusata Frrrusac, 1821,
and L. littorea Ferrusac, 1821. In the following section
we have attempted to evaluate the role of various environ-
mental stimuli which might affect the behavior of L.
planaxis and L. scutulata.

Laboratory Studies

The Role of Algae: In the field adult Littorina scutulata
are most abundant on Pelvetiopsis limitata, somewhat
less common on Gigartina papillata, and occur only rarely
on other species of algae. At least two explanations for
this distribution are possible. Littorina scutulata may
choose certain species of algae in preference to others or
they may choose a certain intertidal level regardless of
the type of algae present. Three sets of experiments were
set up to test the first alternative.

1. An aquarium (3353127 cm) was set up fol-
lowing the procedure of BARKMAN, op. cit., and VAN
DonceEN, 1956. There was only a shallow layer of water

on the bottom which ran from an inlet at one end to an
outlet at the other. One species of alga was placed at the
inlet with a concrete block placed on each side to prevent
the algae from washing down the aquarium (cf. diagram
in VAN DoncEN). One hundred Littorina scutulata were
placed 41.5 cm downstream from the algae. Controls were
run in which no algae were present. All trials lasted for
12 hours and each of the 4 species of algae was run twice.
The bottom of the tank was scrubbed after each run to
prevent the periwinkles from detecting the scent of algae
or their own mucus trails. Results were recorded in 4
categories: those snails which were on the algae, those
that moved upstream but were not found on the algae,
those which remained in the starting area, and those which
moved away.

Results of this experiment (Table 2) show that Littorina
scutulata moves toward Gigartina and Pelvetiopsis in
greater numbers than toward Iridophycus flaccidum Set-
CHELL & GARDNER, 1937, and toward the latter in greater
numbers than toward Prionitis lanceolata Harvey, 1853.
These differences are all significant at greater than the
0.001 level by the chi-square test. There was no significant
difference between Pelvetiopsis and Gigartina. The num-
ber moving upstream but not found on the algae remained

Table 2

Attraction of Littorina scutulata to Single Species of Algae when Placed Down-Stream from the Algae

Up-Current
on algae not on algae
Control:
trial 1 0 35
trial 2 0 4]
Total 0 76 (38% )
Pelvetiopsis limitata
trial 1 44 32
trial 2 30 50
Total 74(37%) 82(41%)
Gigartina papillata
trial 1 34 34
trial 2 32 46
Total 66(34%) 80(41% )
Inidophycus flaccidum
trial 1 14 45
trial 2 23 34
Total 37(20%) 79 (42%)
Prionitis lanceolata
irial 1 5 31
trial 2 8 77
Total 13( 6%) 108 (54% )

Central Down-Current Total
41 24 100
27 32 100
68 (34%) 56 (28% ) 200
15 99

1 19 100

9( 5%) 34(17%) 199

9 16 93

9 13 100

18( 9%) 29(15%) 193
31 7 97
13 22 92
44 (23%) 29(15%) 189
9 52 97

2 15 102

11( 5%) 67 (34%) 199

Page 50

nearly constant throughout the runs except in the Prionitis
tests. For unknown reasons the second of the two Prionitis
runs differed significantly from the first and caused this
upstream non-algae value to be high.

When no algae were present, 38% of the Littorina
scutulata moved upstream ; when algae were present, 60%
or more moved upstream except in the one run using
Prionitis.

2. The second set of experiments, similar to another
series carried out by BARKMAN, op. cit.,and VAN DoNGEN,
op.cit., was conducted on the bottom of two large tanks,
each of which had a shallow layer of water running
over the bottom. One hundred Littorina scutulata were
placed on the bottom in the center and four patches of
algae were each placed at 15cm distance form the L. scut-
ulata. At the end of 12 hours the number of animals on each
species of alga was recorded, the bottom of the tank
scrubbed, and the four algal species were rotated 90°
so that after four trials each species had been in each
position, thus eliminating any effects caused by unequal
lighting, etc., on the tank floor.

Gigartina and Pelvetiopsis attracted the greatest num-
ber of periwinkles (Table 3), Iridophycus was second,

Table 3

Selection of Algae by Littorina scutulata
When Four Species of Algae are Presented Simultaneously

dees

Se Sans ees
a, <S ~S as) we
ee ean | &
aS Ry. a =
Tank 1 21 38 16 3 20 98
12 32 44 4 6 98
9 15 36 24 11 95
32 7 28 7 15 89
subtotal 74 92 124 38 52 380
Tank 2 35 30 15 1 15 96

subtotal 91 97 70 53 84 395

Total 1G5a SS a9 4ee eS lS CMe
Percent of algae:
25.8) 2916) 30/4 4-2

and Prionitis was last. The difference between the first
three and Prionitis was significant at the 0.001 level, but

THE VELIGER

Vol. 10; No, 1

there was no significant difference between the first three.
Nonetheless the order of preference was similar to that
of the first experiment.

3. In the third set of experiments two sheets of
clean glass (22 52cm) were placed in a small aquarium
(303861cm) such that they formed a V at the
bottom; the slope of each was 37° from the horizontal.
An 8 to 10cm band of a single species of alga was held
on the upper surface of each plate 3 cm below the surface
by a thread around each plate. In the absence of the
algae, periwinkles placed under water in the V would
climb out of the water to the top of one of the glass plates.
However, with the algae present as described they would
congregate on the algae, at least temporarily, rather than
continue up the plate. Between 60 and 70 individuals
were tested separately for each of four species of algae.
After one hour the number of animals on the algae and
above it was recorded, the glass plates were scrubbed,
and fresh animals were started at the bottom.

Pelvetiopsis retained the greatest number of Littorina
scutulata beneath the water (71%), Gigartina and Irido-
phycus were second (36 and 42%), and Prionitis was
third (18%). The results are similar to the preceding
experiments except that Gigartina dropped to second
place with Iridophycus. The difference between Pelveti-
opsis and the second two is significant to the 0.001 level,
and between these two and Prionitis to the 0.01 level
by the chi-square test.

The results of these three experiments show that the
four algae are probably preferred by Littorina scutulata
in the order: Pelvetiopsis, Gigartina, Iridophycus, Prio-
nitis. On Bodega Head, L. scutulata was found on nearly
every patch of Pelvetiopsis that we observed. Gigartina
appeared to be occupied only when it was found at levels
comparable to Pelvetiopsis, or when it was lining tide-
pools. It was commonly found at lower levels and in more
exposed sites without any periwinkles on it. Iridophycus
was lower than Pelvetiopsis in the field, and L. scutulata
was present on it only in those unusual locations where
it occurred high. Prionitis was found in tide-pools and it
only rarely had adult periwinkles on it.

When Littorina scutulata were placed in an aquarium
the majority would quickly climb out of the water. Our
laboratory experiments above have shown that the pres-
ence of Pelvetiopsis in the water will keep these animals
below the water line. Gigartina, while preferred over
the remaining two species of algae when at the surface,
was not effective in keeping the periwinkles beneath the
water; this might explain why Gigartina lacks periwinkles
when it occurs low intertidally.

In order to test the effects of the interaction between
water level and algae on the position of Littorina scutulata,

Vol. 10; No. 1

THE VELIGER

Page 51

the third experiment was modified. In this case the algae
formed a 20cm band extending equally above and below
the water line. Color-marked individuals were placed in
the bottom of the V, and their locations with respect to
the water line were recorded every 1 or 2 hours, except
during 8 hours of darkness each night. These figures were
converted to an estimated time spent above or below
water line on the algae (a separate set of experiments,
testing the effect of tidal action independent of the position
of macroscopic algae, will be described later).

It was found that all individuals (15) remained on the
algae for the 75 hour duration of the study; most moved
up and down, entering and leaving the water several
times, but 3 apparently remained well above the water
line throughout the test. Littorina scutulata were above
the water 59% of the time, below 21% and at the water
line 20%. This indicates that L. scutulata retains the
tendency to move up out of the water when on Pelvetiopsis
but that they will remain in the water rather than leave
the algae.

These experiments indicate that Littorina scutulata
select both the level and the species of algae on which they
occur. BARKMAN, op. cit., found a similar situation for
L. obtusata.

On Bodega Head small (2-6mm) Littorina scutulata
occur in large numbers in tide-pools on Prionitis lanceo-
lata, but adults are rarely found in this situation. Conse-
quently we repeated the “V-glass” experiment with 77
young L. scutulata using Prionitis on the glass below water
level. The results of this study showed that 79% of the
young remained submerged for at least 1 hour on Prio-
nitis, while only 18% of the adults did so.

Since Littorina planaxis is normally found above the
level of macroscopic algae, the third experiment (V-glass)
tried on L. scutulata was rerun on L. planaxis. Whereas
Pelvetiopsis was sufficient to keep 71% of the L. scutulata
beneath the water for an hour, it retained only 17% of
69 L. planaxis tested.

Dau, op. cit., has shown that Littorina planaxis will
feed on chopped-up macroscopic algae, but rarely on
whole fronds. He suggested 3 reasons for this behavior:
(1) L. planaxis prefers a more stable substrate, (2) they
are unable to bite off pieces of whole algae, (3) “most
macroscopic algae are somehow chemically disagreeable.”
He eliminated the third possibility, since the snails did
consume the chopped-up macroscopic algae. DAHL found
that there were no obvious differences in the radulae of
the two littorines, which makes the second reason unlikely.
Therefore the type of substrate may be the stimulus to
which L. planaxis responds. Apparently L. planaxis will
continue to crawl upwards until it locates a suitable, (7. e.
stable) substrate on which to feed.

In summary, Littorina scutulata display a strong nega-

tive geotaxis while in water; this is suppressed in the
presence of certain preferred species of macroscopic algae.
Pelvetiopsis is preferred by adults and Prionitis by young.
The snails can distinguish the various species of algae and
show a definite order of preference. Negative geotaxis
continues in effect while L. scutulata is submerged; this
tends to move the periwinkles to the highest macroscopic
algae, which are usually also the preferred species. In
contrast, L. planaxis displays a strong negative geotaxis
which is not suppressed in the presence of macroscopic
algae. Its usual food is microscopic encrusting forms
occurring high intertidally.
Tidal Action: It has been shown in the laboratory that
the presence of certain species of macroscopic algae will
cause a separation in vertical location of Littorina plan-
axis and L. scutulata; however, other factors, such as
tide, may also play a role in determining the separation
seen in the field. In order to determine in the laboratory
whether fluctuating water level alone would cause a
separation of the two species, a large aquarium (113
113<179cm) was set up in which water level was peri-
odically raised and lowered. The “high tide” was 72cm
above the bottom and the “low tide” was 30cm up,
making a total range of 42 cm.

Both species were found over a wide range of heights

(Figure 8), but it will be seen that there was an immediate
separation of over two standard deviations between the
average positions of the two species. Littorina scutulata
occurred mainly around the mean “high tide” line. Littor-
ina planaxis climbed higher out of the water and the
majority was found at the top of the tank, where an
overhanging lip kept them from moving higher. Neither
species made vertical migrations with the artificial tides.
Littorina scutulata became active each time the water
reached their level, but most of the L. planaxis were a
considerable distance above the water and remained dry
and inactive.
Geotaxis: It has already been mentioned that both Littor-
ina planaxis and L. scutulata climb up out of water when
placed in an aquarium. Also we noted that certain species
of macroscopic algae will override the negative geotaxis
in L. scutulata, but that this geotaxis is still present,
acting to move the animals up to the highest algae.

Since Littorina planaxis will continue to crawl upwards
after leaving the water, it seemed likely that some environ-
mental cue must prevent them from continuing to crawl
up away from the ocean. In the field we noted that they
would not continue to move up on dry rock and seemed
to limit their activities to periods when the rocks were wet
from the spray of high tide.

Page 52

+40

+20

- 20

cm below

- 40

- 60

Littorina planaxis
E33 Littorina scutulata

-70
ul 2 3 4 5

- Tidal Cycles

Figure 8

Movement of Littorina placed on bottom of tank with fluctuating
water level

In the laboratory we found that Littorina planaxis
would climb out of water and up a warm, dry glass
surface as far as they were able to carry enough water
with them to keep the glass moist. Those animals placed
directly on a warm, dry surface closed up and did not
move down. However, those animals that moved up from
water onto a dry surface would turn and move down the
glass if a blast of hot dry air from a hair dryer were
directed toward them. A blast of cool air did not have
this effect. Heating the glass ahead of the animal from

THE VELIGER

Vol. 10; No. 1

the underside would also cause the snail to reverse direc-
tions and go down. Thus, a reversal of negative geotaxis
in L. planaxis can be elicited when an animal climbing
above water encounters unfavorable temperatures. Also,
an animal desiccated for a considerable time will move
down when rewetted. Studies by other workers have
shown that many species of Littorina exhibit reversals of
geotaxis after desiccation (BARKMAN, op.cit.; NEWELL,
op.cit.; JANSSEN, 1960; Evans, op. cit.).

Light: Since Littorina planaxis and L. scutulata occur at
different tidal levels, it seemed possible that their response
to light might be one of the controlling factors. Eighty-
seven L. planaxis and 85 L. scutulata were tested 3 times
each for light response, using the same small aquarium
and sloping glass plates which were employed in the third
of the experiments with algae. A 100 watt bulb was placed
at one end of the aquarium so that an animal climbing
the glass plates would either be going directly toward the
light or away from it. The light intensity varied from
400 foot-candles at the top of the nearest glass plate to
12 foot-candles toward the far end of the aquarium.
Littorina planaxis proved to be strongly photonegative
but L. scutulata was only slightly so (Table 4). We ob-
tained similar results using 86 young L. scutulata. Smaller
samples of each species were tested on a horizontal glass
plate in the aquarium; results were similar.

Table 4

Responses to Light
by Littorina planaxis and Littorina scutulata

Littorina' Littorina scutulata'
planaxis (young) (adult )
(adult ) (34-6mm) (10- mm)
ap AP ae 2 10 13
++4+— 1 24 14
4+ —— 13 29 32 ©
——— 71 23 26

™ Numbers of individuals falling into each of the possible categories
of response. + = movement towards, — = movement away
from light. Each symbol equals one experiment and each animal
was tested three times.

The experiments on sloping glass were rerun using
strong natural light (15000 foot-candles) shining in at
a similar angle and the results were practically identical
to those with the weaker light. All experiments were
run at water temperatures between 14 and 15°C.

The strong photonegative response of Littorina planaxis
does not prevent it from moving up in the intertidal zone.

Vol. 10; No. 1

THE VELIGER

Page 53

However, this species is found clustered in cracks and
crevices, and light may be the stimulus causing this. Such
behavior would likely be of survival value by reducing
desiccation, heating, and predation. Littorina scutulata
may show a weaker photonegative response because, in
its more protected and wet location, a strong response
would be of little value. However, the response may be
sufficient to drive some animals under the protection of
algal fronds.

Apparently most species of Littorina have distinct light
responses, but these are not always correlated in any
obvious way with behavior in the field (BARKMAN, Of. cit.;
NEWELL, op. cit.; JANSSEN, op. cit.; CHARLES, 1961a,
1961 b, 1961c).

Behavioral responses controlling distribution of Littor-
ina planaxis and L. scutulata are summarized in Table 5.

Table 5

Behavioral Responses Controlling Distribution of
Littorina planaxis and Littorina scutulata

Upper Limits Lower Limits

Littorina planaxis
Reversal of Geotaxis Negative Geotaxis

1. on reaching high 1. when under water

temperatures
2. when rewetted after 2. when on macrosco-
desiccation pic algae

Littorina scutulata
Negative Geotaxis
suppressed by pre-
ferred species of
algae

Negative Geotaxis

SUMMARY anv CONCLUSIONS

1. Littorina planaxis and L. scutulata are common
periwinkles of the rocky coast of California. Littorina
planaxis occurs very high intertidally, usually on exposed
rocks in the splash zone. Littorina scutulata is found largely
in mid-intertidal areas, frequently in beds of macroscopic
algae and pools rimmed with algae.

2. The physiology and food habits of each species are
correlated with these differences in habitat. Littorina
planaxis is more tolerant of desiccation, temperature, and
fresh water, — conditions encountered more often in
the high intertidal. Littorina scutulata readily consumes
macroscopic algae, while L. planaxis apparently will take
only microscopic encrusting forms.

3. Although tolerances to environmental conditions
ultimately must determine the distributions of Littorina
planaxis and L. scutulata, selection apparently has resulted
in behavioral responses which cause avoidance of these
limits. Littorina planaxis moves up in the field, acting
under a strong negative geotaxis. It thus congregates in
the highest portions of the splash zone. Littorina scutulata
shows a similar upshore orientation which keeps it out
of low intertidal areas, but this behavioral response is
negated by the presence of certain macroscopic algae in
the mid-intertidal.

ACKNOWLEDGMENTS

We wish to thank Dr. Ann E. Kammer and Dr. Cadet
Hand for their assistance in carrying out this investigation,
and for reading the original manuscript. We are especially
indebted to Dr. Ralph I. Smith for his help in preparing
the final version. We were both supported by fellowships
from the National Science Foundation during the course
of this study.

LITERATURE CITED

BarkKMaAN, J. J.

1955. . On the distribution and ecology of Littorina obtusata
(L.) and its subspecific units. Arch. Neerl. Zool. 11: 22 - 86

Bow us, R. Davip
1966. | Comparison of the ability to resist and withstand desic-
cation of Littorina planaxis and L. scutulata. Unpubl.

student report (Zool. S212) on file, Dept. Zool., Univ. Calif.,
Berkeley.

Cuar es, G. H.

1961 a. The orientation of Littorina species to polarized light.
Journ. Exp. Biol. 38: 189 - 202

1961 b. The mechanism of orientation of freely moving Littorina
hittoralis (L.) to polarized light. Journ. Exp. Biol. 38: 203
to 212

1961c. Orientational movements of the foot of Littorina species
in relation to the plane of vibration of polarized light. | Journ.
Exp. Biol. 38: 213 - 224

DanHL, ARTHUR LYON
1964. | Macroscopic algal foods of Littorina planaxis Philippi

and Littorina scutulata Gould (Gastropoda: Prosobranchiata).
The Veliger 7(2): 139-143. (1 October 1964)

Evans, FRANK

1961. Responses to disturbance of the periwinkle Littorina
punctata (GMELIN) on a shore in Ghana. Proc. Zool.
Soc. London 137: 393 - 402

Hewatt, WIus G.

1937. Ecological studies on selected marine intertidal commu-
nities of Monterey Bay, California. Amer. Midland Nat.
18: 161 - 206

Page 54 THE VELIGER Vol. 10; No. 1

Janssen, C.R.
1960. The influence of temperature on geotaxis and phototaxis
in Littorina obtusata (L.). Arch. Neerl. Zool. 13: 500 - 510

NEWELL, G. E.
1958a. The behaviour of Littorina littorea (L.) under natural

conditions and its relation to position on the shore. Journ.
Mar. Biol. Assoc. U.K., 37: 229 - 239

1958b. An experimental analysis of the behaviour of Littorina
littorea (L.) under natural conditions and in the laboratory.
Journ. Mar. Biol. Assoc. U. K. 37: 241 - 266
Nico, J. A. CoLin
1960. The biology of marine animals. i-xi+707 pp.;
illus. New York, N. Y. (Interscience Publishers, Inc.)
NortH, WHEELER J.
1954. Size distribution, erosive activities, and gross metabolic
efficiency of the marine intertidal snails, Littorina planaxis and
L. scutulata. Biol. Bull. 106: 185 - 197
Ricketts, Epwarp F. « Jack CaLvin
1962. Between Pacific tides. 3rd. ed., 2nd. rev. by Jorn W.
HEDGPETH. xilli+516 pp.; 135 text figs.; 46 plts.; Stanford
Univ. Press, Stanford, California.
Van Doncen, A.
1956. The preference of Littorina obtusata for Fucaceae.
Arch. Neerl. Zool. 11: 373 - 386
WILuiAMs, GEorcE C.
1950. | Movement and dispersal in Littorina planaxis. Un-
publ. student report (Zool. S212) on file, Dept. Zool., Univ.
California, Berkeley.

Vol. 10; No. 1

THE VELIGER

Page 55

Preliminary Observations on the Feeding Behavior

of Conus purpurascens BRoDERIP, 1833

JAMES NYBAKKEN

Department of Biological Sciences, California State College at Hayward, Hayward, California

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

94542

95039"

[These studies were aided in part by a contract between the Office of Naval Research, Department of the Navy,
and the University of Arizona, NR 4839 (00).]

(Plate 4)

AT LEAST TWO SPECIES of the genus Conus, Conus striatus
LInNAEuS, 1758, and C. catus Hwass in BrucutkrE, 1792,
are known to prey on living fishes (KoHN, 1956). Both
are Indo-West Pacific in distribution. Until now no species
of the West American Conus fauna has been reported in
the literature as preying on fishes. This paper reports on
the feeding of C. purpurascens Broperip, 1833 on fishes
under captive conditions.

During the summer of 1966, while I was initiating
a study of the ecology of the West American species of
Conus in the Guaymas area of western Mexico, four speci-
mens of C. purpurascens, together with representatives of
7 other West American Conus species, were collected and
brought alive to aquaria at the Moss Landing Marine
Laboratories. In an effort to maintain these specimens
alive, I contacted Dr. Alan Kohn who stated, from obser-
vations of Dr. R. T: Paine, that C’. purpurascens was a fish
eater. I therefore attempted to feed local fishes to the
Captive specimens with the aim of studying the feeding
process itself. A few preliminary results of these studies,
primarily in the form of a sequence of photographs of
the feeding process, have been obtained and are reported
here.

METHOD anp RESULTS

A one gallon aquarium was set up near a window in
my office so that illumination woud be sufficient for
photography. The aquarium had a bottom substrate of
sand and was filled with sea water having a temperature

' Contribution No. 6 from Puerto Pefasco Marine Research Station

near that of the large holding tank (about 22°C). Each
feeding sequence was either observed and notes taken or
it was the subject of a photographic record.

Only two species of fishes have as yet been used in the
experiments, Xererpes fucorum (JORDAN & GILBERT, 1880)
and Oligocottus snyderi GREELEY, 1901. Both species have
been taken without hesitation by all four captive speci-
mens of Conus purpurascens. More species of fish are
being tested.

For each observation the specimen of Conus purpur-
ascens was removed from the holding tank and placed into
the experimental aquarium. Shortly after placing the
Conus in the tank a fish was netted out of the fish holding
aquarium and placed into the experimental aquarium
with the Conus.

In general the sequence of events observed in the feeding
was similar to that outlined by KoHn (1956) for Conus
striatus.

When placed into the isolation aquarium from the
holding tank, Conus purpurascens shows little movement
for several minutes. Then the siphon is extended and
moved slowly around. If the animal is left alone in such
circumstances, after about 10 minutes it extends out from
its shell and begins to burrow down into the sand. The
usual inactive position of this species in aquaria is partially
or totally buried in the sand with only the siphon pro-
truding above the surface.

If, however, a fish is introduced into the aquarium its
presence is sensed by the Conus almost immediately.
Koun (1956) has suggested that Conus striatus detects
the presence of fish by chemoreception using the osphra-
dium. This would appear to be the mechanism acting in

Page 56

THE VELIGER

Vol. 10; No. 1

Conus purpurascens also. I tested this by adding to the
Conus tank water taken from the fish holding tank. This
prompted immediate activity on the part of two Conus
individuals indicating use of chemoreception to sense the
presence of the fish.

The animal responds to the presence of the fish by
extending its proboscis (Plate 4, Figure 2). The proboscis
is orange-red in color on the dorsal side and flesh color
on the ventral side. The orange-red color is particularly
striking in an animal which otherwise has a subdued
purple or brownish coloration. The proboscis may extend
out as far as one to one-and-a-half times the shell length.
The proboscis is waved around or else moved along the
substrate in search of the fish (Plate 4, Figure 2). Often
a combination is used. The exact method of location of the
fish by the proboscis is unknown as yet, but it appears
in some cases as if it were accidental. In no instance
observed did the proboscis follow a direct route to the fish.

Once the proboscis tip touches the fish (Plate 4, Figure
3) there is occasionally a slight retraction away from the
fish followed by a second contact. On other occasions
the initial contact is maintained. The proboscis next begins
to move slowly over the body of the fish as the cone
apparently attempts by touch to locate a soft area where
injection of the radula tooth, which is held in the tip
of the proboscis, is assured. With the exception of two
instances, in all feeding sequences observed thus far in
this study (a total of 20 observations on 4 individuals)
the proboscis tip continued to pass over the body of the
fish until it reached the ventral abdominal surface. Only
when this area had been reached was the radula tooth
injected and the proboscis contracted to pull the fish in
toward the mouth (Plate 4, Figure 4). This aspect of
the feeding sequence occurred with such rapidity it was
impossible to separate individual events. Photographs at
1/250 second were blurred (Plate 4, Figure 4).

Of the two instances where injection of the radula
tooth was not in the ventral abdominal area, one was
an injection just ahead of the caudal fin into the lateral
trunk musculature, while the other was, assumedly, into
the soft tissue of the mouth of the fish. In this latter case,

the fish observed the moving tip of the proboscis and
attacked it, probably as potential prey, taking the tip
into the buccal cavity whereupon it was immediately
stung. In both of these instances the fish were specimens
of Oligocottus snyderi.

That the radula tooth is actually injected is certain
for I have observed the actual tooth imbedded in a
specimen of Xererpes fucorum at one end and held in the
proboscis of the Conus at the other. The radula teeth are
very large (6.8 to 7.6mm in one specimen) and thus
observable in large dissected specimens of Conus purpur-
ascens without optical equipment.

On only one occasion did a fish escape after the injection
of the tooth, despite the fact that injection of the tooth
invariably caused violent thrashings by the fish. In this
instance the fish (Oligocottus snyderi) swam straight to
the surface of the water and sank immediately back to
the bottom where it died in two minutes. It was then
consumed by the cone.

When the captured fish has been brought up to the cone
via contraction of longitudinal muscles of the proboscis,
the muscular mouth surrounding the proboscis expands
greatly and engulfs the fish (Plate 4, Figures 5 to 8). This
process is usually completed within one minute. When the
fish has been thus ingested, the cone begins to burrow
into the sand.

Live fish do not seem mandatory for this species as on
two occasions when dead Oligocottus snydert were put
into the aquarium they were attacked and eaten in exactly
the same way as living fishes. However, both of the dead
fishes used in these experiments were very fresh. It is
not yet known at what time after death Conus purpur-
ascens will refuse such fish. This is in contrast to KoHN’s
(1956) findings for C. striatus.

Conus purpurascens appears to be a nocturnal feeder
as are most of the species of the genus (KoHn, 1959).
In several instances, specimens having initiated attacks on
fishes in a dimly illuminated room immediately ceased
searching and retracted into their shells when bright light,
especially daylight, was let into the room. Further study
is being continued on this aspect of behavior.

Explanation of Plate 4

Figure 1: Conus purpurascens, siphon out, and a specimen of the
fish Oligocottus snyderi which has just been placed into the tank
Figure 2: Conus purpurascens with proboscis extended actively
searching for the fish. The extended proboscis is visible below the fish
Figure 3: The proboscis of Conus purpurascens making initial
contact with the fin of the Oligocottus snyder

Figure 4: Conus purpurascens immediately after injection of the
radula tooth into the Oligocottus snydert. The blur is the thrashing
fish

Figure 5: Conus purpurascens proboscis with impaled fish con-
tracted, expanding its mouth to engulf the fish

Figure 6: Conus purpurascens beginning to engulf the captured fish

Figure 7:Conus purpurascens with fish almost completely engulfed
by the expanded mouth and buccal cavity
Figure 8: Conus purpurascens with fish completely engulfed

Tae VELIGER, Vol. 10, No. 1 [NyBAKKEN] Plate 4

Figure 1 Figure 2

Figure 3

Figure 5 Figure 6

Figure 7 Figure 8

Vol. 10; No. 1

THE VELIGER

Page 57

Although it is not known with what frequency feeding
occurs in nature in Conus purpurascens, under captive con-
ditions specimens in my aquaria have fed on two days in
succession indicating, as KoHN (1956, 1959) has reported,
that digestion in this genus is extremely rapid.

Conus purpurascens also appears to be able to survive
without food for periods of at least two months, the
length of time specimens in the author’s aquaria were
held before feeding was initiated. This time interval of
survival without food is similar to that reported for C.
californicus by SAUNDERS & WoLFson (1961). How-
ever, the author has maintained other tropical American
Conus species alive in aquaria, notably C. brunneus Woop,

1828, and C. virgatus Reeve, 1849, for five months with-
out feeding.

SUMMARY

The feeding behavior of Conus purpurascens under cap-
tive conditions is described. Conus purpurascens was ob-
served to attack and eat two species of fishes, Oligocottus
snydent and Xererpes fucorum. The feeding behavior was
the same in all instances and was similar to that reported
by Koun (1956) for C. striatus. Feeding behavior in
C. purpurascens appeared to be influenced by light.

LITERATURE CITED

Koun, ALan J.

1959. The ecology of Conus in Hawaii. Ecol. Monogr.
29: 47 - 90.
1956. _ Piscivorous gastropods of the genus Conus. Proc.
Nat. Acad. Sci. 42 (3): 168-171
SaunpeErRs, Paut R. « Fay HENry WOLFSON
1961. Food and feeding behavior in Conus californicus Hinps,

1844. The Veliger 3 (3): 73 - 76; plt. 13

(1 Jan. 1961)

Page 58

THE VELIGER

Vol. 10; No. 1

Note on the Radula of Mitromica Berry, 1958

JAMES H. McLEAN

Los Angeles County Museum of Natural History
goo Exposition Boulevard, Los Angeles, California 90007

(1 Text figure)

THE SYSTEMATIC position of the monotypic species Mitro-
mica solitaria (C. B. ADAMS, 1858) has long been a puzzle.
Although the species has four columellar plaits with the
heaviest fold in adapical position, it has other shell char-
acters not typical of Mitridae: a thickened and highly
denticulate outer lip, a suggestion of an anal sulcus, and
sharply cancellate sculpture, characters stressed by BERRY
(1958, p. 94) in his diagnosis of Mitromica. Copies of
previously published illustrations of the holotype of Mitra
solitaria C. B. ApAMs and the synonymous M. nodocan-
cellata STEARNS, 1890, are given by KEEN (1958, p. 428,
text figure 646).

The radula in the Mitridae has been well documented
and is considered a more reliable indicator of family and
subfamily relationships than the shell (CERNOHORSKY,
1966). Examination of the radula of Mitromica solitaria,
from a specimen collected at low tide, San Luis Gonzaga
Bay, Gulf of California, skillfully mounted by Mr. Peter
M. Oringer (Figure 1), shows the genus to belong to the
subfamily Vexillinae of the family Mitridae. The rachid-
ian tooth is bow-shaped and bears seven cusps, inter-
mediate in character between that of the tricuspid
rachidian of Pusta Swainson, 1840, and the multicuspid

| 25éh |

rachidian of Vexillum Ropine, 1798, figured by THIELE
(1929, pp. 337-338, text figures) and CERNOHORSKY
(1966, pp. 118-121, text figures). The lateral teeth are
sickle-shaped as in both of these genera.

LITERATURE CITED

Berry, SAMUEL STILLMAN
1958. Proposal of five new generic taxons. Leafl. in Malacol.
1 (16 ) fart. 1]: 91-98 (31 May 1958)
CERNOHORSKyY, WALTER OLIVER

1966. A study of mitrid radulae and a tentative generic ar-
rangement of the family Mitridae. The Veliger 9 (2): 101
to 126; 47 text figs. (1 October 1966)
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)
THIELE, JOHANNES
1929. Handbuch der systematischen Weichtierkunde. Erster
Teil, Loricata; Gastropoda. I: Prosobranchia (Vorderkiemer).
pp. 1-376; 470 text figs. Jena, Gustav Fischer Verlag

Tf

Figure 1

Radula of Mitromica solitaria (C.B. ADAMS).
The rachidian plate and one lateral plate are shown

Vol. 10; No. 1

THE VELIGER

Page 59

The Shell Ornament of Hysteroconcha and Hecuba (Bivalvia):

a Test Case for Inferential Functional Morphology

BY

ROBERT M. CARTER

Sedgwick Museum, Cambridge, England

(Plates 5 to 7; 2 Text figures)

For MANY YEARS zoologists have been making functional
inferences about the animals that they study, and, in the
case of vertebrate zoologists at least, this type of inference
has often been made with a fair amount of scientific
rigour. But the recent paper by Geist (1966) on the
evolution and functional significance of horn-like organs
in the mammals convincingly demonstrates that even with
such spectacularly developed and common organs as these
there is still ample room for subjective interpretation and
divergences of opinion. Unfortunately, the tendency in
invertebrate studies has been for functional interpretations
to be made merely as asides in the course of routine
systematic work; at best such interpretations are generally
loosely argued, at their worst they may even become self-
contradictory. In the field of invertebrate palaeontology
there has recently been a strong resurgence of interest in
functional, as opposed to systematic, studies, and in a
series of stimulating papers Rupwick (1961, 1964a,
1964b) has outlined a rigorous ‘paradigmatic’ method-
ology which enables at least some degrec of precision to
be reached in such interpretations. Though the method
was developed initially for fossil brachiopods, its appli-
cations are not restricted to any one group of animals,
and its implications are equally broad.

THE PARADIGMATIC METHOD

Briefly, the paradigmatic (or mechanistic) approach may
be considered to consist of four logically discrete steps:

1, Perception

The animal structure in question is examined in some
detail and compared with our knowledge of the biology
of the animal’s relatives, living and fossil, and with our
knowledge of similar structures in unrelated animal groups.
As a result of this comparison it is generally possible to

suggest more than one reasonably plausible function for
the structure.

2. Specification

It is now necessary to test these various tentative func-
tional interpretations against the different idealised struc-
tural specifications that are relevant to each. It is essential,
in so doing, to take account of the limitations imposed
by the properties of the materials involved (e. g. bone,
muscle or calcium carbonate, etc.), where these factors
are significant. One is thus led to the concept of estab-
lishing a paradigm for each postulated function. The
paradigm is “the structure that would be capable of
fulfilling the [postulated] function with the maximal
efficiency attainable under the limitations imposed by
the nature of the materials” (Rupwick, 1964a, p. 36).

3. Evaluation

By comparing the observed structure with its paradigm
for each postulated function it is possible to get some
idea of the degree of efficiency with which the observed
structure could have fulfilled each function. As Rupwick
(1964a, pp. 36-37) has already stressed, it is most
important to note that closeness of approximation of a
structure to a given paradigm only measures the degree
of possible functional efficiency of the structure for that
paradigm; it tells us nothing directly about the probability
that the function we are considering is in fact the correct
function.

4. Interpretation

Thus far the analysis has been entirely methodological,
and therefore relatively objective. However, to choose
between several competing paradigmatic analyses inevi-
tably involves a higher degree of subjectivity. It is
necessary at this point to digress slightly.

Page 60

THE VELIGER

Vol. 10; No. 1

Most animals that have been adequately studied
demonstrate a high level of integration between function
and structure. Indeed, it was presumably just this that
gave rise to the presently popular technique of viewing
animals as functional units of the same type as machines.
But it has become accepted that in writing of animals in
this way the worst crime is to use language that might
be construed as teleological in any sense. PITTENDRIGH
(1958, p. 394) has pointed out that an animal, just like
a machine, is an end-directed mechanism, and that the
slightly hysterical attempts of many writers to use non-
teleological language are based “‘on the mistaken view that
the efficiency of final causes is necessarily implied by the
simple description of an end-directed mechanism” (I. c.,
p. 393). PrrrenpricH further maintains that it is wrong
to consider, as many of us do, the animal as a system some
features of which may, or may not, be adaptive: “the
living system,” he writes, “is all adaptation insofar as
it is organized.” In addition, he takes issue with HUXLEY
for having called the organism “‘a bundle of adaptations”
because such a statement implies that organisation is an
additive phenomenon, and that discrete adaptations can
be isolated from the system.

It would seem to be a dangerous procedure, then, to
abstract one particular structural feature of an animal,
and to attempt to analyse its probable adaptive or
functional role, for “the organism’s ends” are likely to be
often “served in complex ways unamenable to simple de-
scription.” But presumably PrrTENpRIGH would be the first
to agree that it is only by initially, and tentatively, treating
the organism as a bundle of adaptations, and by attempt-
ing to analyse each of these adaptations separately that
we are in the end able to piece them all together
and approach the task of interpreting the animal as
“all adaptation.”

Some structures approach their paradigm very closely
— for example, the camouflage of many moths and stick
insects, if not corresponding 100% with a camouflage
paradigm, is certainly very close to it — but it should be
obvious that no structure will ever perfectly fulfill its
paradigm, for no animal functions at 100% efficiency.
Further, in line with the points quoted above, since all
structures are an integrated part of the adaptation of
the whole organism they are very rarely, if ever, going
to be the result of a single steady selective pressure.
Commonly we might expect a structure to be selected for
more than one specific function and hence to approach
several paradigms loosely, and none specifically. Indeed,
Rupwick (1965) hasalready described a structure (spines
on the Jurassic brachiopod Acanthothiris) that he con-
siders to have probably served two discrete functions; and

Geist (1966, p. 192), in writing of the functions of horn-
like organs in vertebrates, concludes that “cervid antlers
and bovid horns evolved convergently and function as
weapons, secondly as guards and thirdly as organs binding
opponents together during pushing and wrestling contests.”

Assuming, then, that a structure will never completely
fulfill its paradigm, in a given instance we are faced with
making comparisons between the actual structure, and the
structural paradigms of several possible functions. We
adopt for this the procedure outlined by Rupwick (1961)
in his introduction of this methodology: whichever para-
digm, of several alternatives, is adjudged to be most
closely approached by the structure in question, that para-
digm is most likely to be the paradigm of the sole correct
function, or of the dominant function of several. The
other postulated functions may then (subjectively) be con-
sidered as close enough to their paradigms to be inter-
preted as secondary functions, or alternatively, they may
be discarded.

It is important to note that it is at this interpretative
stage that a paradigmatic analysis reaches its least rigorous
point, for there is generally no quantitative way of meas-
uring how closely a structure approaches several alterna-
tive paradigms. Hence the choice between such paradigms
is certainly subjective, and must depend largely on the
preferences and opinions of the writer concerned.

THE QUESTION OF FUNCTIONALITY

The question as to whether all structures must of
necessity have a function is, of itself, outside of paradig-
matic methodology. But it is a question that cannot be
evaded when interpreting animal structures, for the
possibility must always be considered that a given struc-
ture is in fact non-functional ( 7. e. non-adaptive). “The
living system is all adaptation insofar as it is organized”:
PirTeNpRIGH would thus presumably maintain that all
major structures are adaptive, and hence have a function.
Rupwick (1964a) is primarily concerned with the logical
impossibility of proving non-functionality, but he clearly
makes the point (pp. 35, 38) that, without prejudice to
the correctness or not of a gencral principle of functionality,
unless we initially assume that a given structure is func-
tional “we shall fail even to perceive the evidence on which
any functional reconstruction must be based.”

However, these two writers’ views apart, it is clearly
impossible to assert that all structures must be functional,
except in a semantically very loose sense, for there are
at least two obvious types of morphological structure for
which such an assertion would be false:

Vol. 10; No. 1

THE VELIGER

Page 61

1. Pleiotropic genetic effects

The pleiotropic effect, whereby one gene locus may give
rise to manifold morphological consequences, has long
been known. Predictably, most of the cited examples
involve Drosophila; for instance, the mutation ‘vestigial’
in Drosophila acts primarily to reduce the wing size, but
it also makes certain bristles erect instead of horizontal,
modifies the balancers, and has noticeable effects on the
wing muscles, the shape of the spermatheca, the speed of
growth, life length and fecundity of the insect (DoBzHAN-
sky, 1941). There is no doubt that pleiotropism is a
widespread genetic phenomenon affecting all groups of
animals. Human ingenuity is such that, if pushed to it,
a discrete function could readily be invented for all the
above morphological changes; or, to put it another way, a
different selective advantage could be attributed to each
of them. But a paradigmatic analysis of any one of these
morphological effects would almost certainly be trivial,
for they all form part of a stable genotype and cannot
meaningfully be treated in isolation. That is not to say
that they are not all individually subject to natural selec-
tion, for modern evolutionary theory holds that selectively
neutral characters are very rare. Rather, it seems probable
that, though all the characters of a pleiotropic set may be
subject to individual selection, in most cases the selection
for the (or fora) dominant character swamps the effects of
the others. E

The genetic research necessary for recognizing pleio-
tropism has scarcely been started for many groups of
Recent animals, and it is all but impossible that we shall
ever be in a position to describe the genome of fossil
forms. Hence the most reasonable approach to this prob-
lem is presumably to treat well developed structures (such
as, say, elephant tusks, or the spines described later in this
paper) as truly adaptive, and assume that they are not
merely ‘side-effects’ due to pleiotropism. In the present
state of knowledge there appears to be no rigorous way
of justifying such decisions; and only experience, or
intuition, can help us make them correctly.

2. Vestigial structures

A further example of a non-functional structure is one
that is adjudged to have become obsolescent after a fairly
long phyletic history. Such vestigial structures are in-
ferred to have been functional, and hence selected for
at some former time, though at the present time they
appear to have no continuing usefulness.

The paradigmatic interpretation of vestigial structures
is also difficult, for they are capable of closely approaching
the paradigm of their ancestral function; as examples
one might quote the unerupted teeth of the baleen whales,

or the rudimentary limbs of some snakes. However, the
problem is not as difficult as with pleiotropism, for vestig-
iality usually leaves some morphological trace — in the
case of a fossilised baleen whale jawbone one would
certainly be suspicious of the apparent lack of wear on
the teeth. Nevertheless, one has no case for insisting a
priori that all vestigial structures are always going to be
recognisable as such because of their inevitably possessing
certain inconsistencies in their morphology; and this is
particularly true of fossil animals where only part of the
total morphology is preserved. The possibility of vestigi-
ality, then, must always be borne in mind in functional
studies.

PHENOTYPIC RESPONSES

The classic types of phenotypic response to the environ-
ment, such as an oyster moulding itself to the substrate or
the shells of a particular species of mollusc being thinner
in sub-saline water, either have an obvious function, or do
not require functional interpretation. But GitBerTt (1966)
has recently described a fascinating and unusual type of
phenotypic morphology in the rotifer Brachionus calyci-
florus Pattas, 1766. This species normally possesses two
pairs of short anterior spines, and a further pair located
posteriorly. If B. calyciflorus is kept in cultures of the
carnivorous rotifer Asplanchna brightwelli Hupson, 1889,
its female parthenogenetic offspring have an additional
pair of long posterolateral spines — structures that were
completely lacking in the previous generation — and there
is also an induced relative elongation of the normal spines.
Eggs from mothers grown in the conditioned medium
that are transferred to fresh medium immediately after
extrusion develop into long spined forms; conversely, eggs
from mothers grown in fresh medium and transferred
on extrusion to conditioned medium hatch into short
spined forms. Successive offspring from the same mother
exhibit decreasing spine production. As GILBERT con-
cludes: “the production of extra spines was
mediated by a factor released into the medium by Asp-
lanchna, and represents a phenotypic response of un-
doubted adaptive significance” — long spined Brachionus
being far more difficult for Asplanchna to eat.

It is presumably a moot point whether this rather
straightforward type of phenotypic response can be ex-
pected in animals higher than rotifers, but it is as well
to be aware of the existence of such a phenomenon, even
if it should presently appear to be confined to one of
the lower animal groups. It would seem likely that such
a confinement reflects our lack of knowledge of the phe-
nomenon in other phyla, rather than its true absence.

Page 62

Since a paradigmatic analysis is not concerned with
how a particular structure arose, but only with how well
adapted it is, phenotypic structures are perfectly amenable
to paradigmatic interpretation. However, should it tran-
spire that the phenomenon described above be widespread
in different animal groups, there might be a case for some
amendment of the present methodology.

SUMMARY

Though it is not possible to maintain the rigour of para-
digmatic methodology right through the functional inter-
pretation of animal structures, such a methodology is an
essential tool in any closely reasoned functional analysis.

The palaeontologist normally accepts as part of his
discipline an almost complete ignorance of the soft-part
anatomy and the behaviour patterns of the specific fossils
that he studies. Of course, he may make much use of the
current biological knowledge of extant relatives of the
animal in question, but in many cases such knowledge is
appallingly scanty. As with most types of scientific method,
the paradigmatic is dependent for its success on its initial
data — these need to be both sufficient and of good quality.
Palaeontological data are always relatively insufficient
in that soft parts are rarely fossilised.

Theoretically the neontologist is in a much stronger
position, for he should have a comprehensive knowledge
of the anatomy, ecology and ethology of the animal he is
studying. But in fact, so little work has been done on many
groups of Recent animals, apart from routine systematics,
that the neontologist is in almost exactly the same position
as the palaeontologist. This is especially so in invertebrate
groups, and particularly true of the Mollusca.

Hecuba and Hysteroconcha are two groups of living
bivalves about which very little has been published outside
of taxonomic information (and even that is scarce). The
degree of interest shown in such spectacular shells by early
European shell collectors is reflected by the voluminous

Explanation

Figure 1: Pitar (Hysteroconcha) lupanaria (Lesson, 1830), xX 2;
Recent, America; Saul Collection, Zoology Department, University
of Cambridge. The left valve of a pair; note especially the rounded
concentric ridges, raised at the anterior end of the shell.

Figure 2: Pitar (Hysteroconcha) dione (LiNNAEUs, 1758), X25
Recent, America; Sedgwick Museum Recent Mollusca Collection,
Cambridge University. In this, and the specimens figured as 3 and
5, note the characteristic position and morphology of the muricid
gastropod boring.

THE VELIGER

Vol. 10; No. 1

description of Hysteroconcha in Systema Naturae — in the
entire ‘Regnum Animale’ only Homo sapiens and Apis
mellifera (the honey bee) can record a longer entry
(DoncE, 1952, p. 87). The demand for specimens of these
shells was due mainly to their beauty of form, but their
exotic origins and relative scarcity were contributing fac-
tors; and even today it is not easy to come by specimens
with wholly intact spines. Inevitably, therefore, attention
has centred on the aesthetic appeal these shells made,
rather than on their scientific significance. It has passed
unnoticed that Hysteroconcha, known from the tropical
Americas to the Northern Americas, and Hecuba, known
from the western Indo-Pacific, have independently ac-
quired the remarkably similar morphological structures
that constitute their main attraction from the collectors’
point of view.

The unfortunate lack of published data on these shells
carries one unexpected advantage — it enables a ‘palae-
ontological type’ of paradigmatic analysis to be carried
out with which future observations on the living animal
may be compared. Thus this paper may serve as a partial
‘test case’ of paradigmatic methodology; the amount of
light thrown on the functional suggestions made herein
by future research on the living animals will be some
guide as to the reliability of the methodology, and perhaps
tell palaeontologists just how far it is possible to make
reasoned inferences from the incomplete data of dead
shells.

Hysteroconcha

In most respects Hysteroconcha is a typical member of °
the Pitarinae (Plate 5, Figure 1), possessing the charac-
teristic dentition, moderately deep pallial sinus and con-
centric ornament of that group. It differs, however, in the
possession of two remarkable sets of spines, one (the

of Plate 5

Figure 3: P. (H.) dione (LinnaEus), x 2; Recent, America; Saul
Collection.

Figures 4 to 7: P. (H.) lupanaria (LEsson), x 2; Recent, America;
Saul Collection. A beautifully preserved double-valved specimen,
photographed from varying angles to show the arrangements of the
spines. Note particularly that no protection is afforded by the
spines to guard the posterior gape against turbid sediment.

[The magnification is stated approximately for all figures]

TuHeE VELIGER, Vol. 10, No. 1 [CarTER] Plate 5

Hee:

ae

fees

ae

|

|
|

Figure 6

Figure 7

Vol. 10; No. 1

primary row) carried on the rounded posterior carina
delimiting the edge of the corcelet', and the other (the
secondary row) along the shell sector marking the edge
of the escutcheon.

ECOLOGY

Hysteroconcha is collected living on sandy beaches, and
also offshore to depths of about 13 fathoms (KEEN, 1958).
Mr. J. Q. Burch (personal communication) has collected
it living on sandy beaches near Guaymas, Sonora, Mexico.
He comments that it is common as an intertidal form
when a living ground has been located (i.e. there are
large numbers of individuals), but that localities where
it occurs are rare. Like many intertidal bivalves it remains
buried under the sand when the tide is out, rising to the
substrate surface to feed on the incoming tide. Its feeding
position I thus infer to be with the posterior spines, and
probably most of the corcelet, protruding above the sub-
strate. The anterior end of the shell is undoubtedly always
buried.

The only predator reported as actually feeding in the
field on Hysteroconcha is Polinices reclusianus (DEs-
HAYES, 1839), which lives under the sand in the same
habitat (Mr. Burch, personal communication). Other
predators that are found in the same type of ecological
station include muricid gastropods (Forreria, Hexaplex,
Muricanthus), Natica, Oliva, asteroids (Astropecten),
crabs and fish (Dr. M. Keen, Professor FE M. Bayer,
personal communication).

DETAILED MORPHOLOGY

Hysteroconcha lupanaria group

[Hysteroconcha lupanaria (LEsson, 1830) ]
This species group can be recognised by the large size
of the adult shell (up to 60mm long, excluding spines),
the possession of a large pre-spinous dissoconch (about
6mm long) and the characteristic rounded concentric
Ornament (not sharp, raised lamellae) in the centre of
the main disc.

The exact pattern of spine secretion is variable within
all species of Hysteroconcha. However, an attempt is
made below to describe the pattern that sets of spines on
the majority of animals approximate to; it must be stressed
that this involves considerable generalisation, and that

1 The term “corcelet” is used in this paper for a morphologically
differentiated area lying outside the escutcheon at the posterior
end of the shell. A further discussion of the term may be found
in CarTErR, 1967.

THE VELIGER

Page 63

it will be possible to find an animal for which just about
every statement made below is mildly incorrect.

Spines

On any one valve the first spine secreted in ontogeny is
generally of the primary row, and thereafter spines are
secreted at regular intervals alternately on the primary
and secondary spine rows. On a particular spine row,
say the primary, though the spines are definitely secreted
on a regular pattern, the distance between any two con-
secutive spines increases during life, 7. e. the ontogenetically
earlier spines are closer together than the later adult ones.

The pattern of spine arrangement between the two
valves is also variable, but generally the spines of the two
primary rows are secreted alternately, whilst those of the
two secondary rows are introduced in symmetrical pairs
at the growing edge (Plate 5, Figures 4, 5, 7). However,
not uncommonly the spines of primary and secondary
rows are both introduced as symmetrical pairs at the
growing edge. Irrespective of this, the alternating relation-
ship between spines on the primary and secondary rows
of one value is generally maintained (Plate 6, Figure 8).

The spines of the primary row rise from the valve edge
at a fairly high angle in an almost truly radial direction
(Plate 5, Figures 5, 6). However, they quickly become
directed posteriorly, at the same time bending markedly
into subparallelism with the plane of the commissure.
They may achieve a length the same as, or a little longer
than, the shell length at the time of their inception (e. g.
spine length 34mm, shell length 27mm).

The spines of the secondary row rise from the shell
edge at a very high angle (over 80°), and are slightly
directed posteriorly from a true radial direction (Plate 5,
Figures 4, 7; Plate 6, Figure 8). They also bend inwards
toward the plane of the commissure, but much less slowly
than do the spines of the primary row. As a consequence,
at about half their normal length they meet with, and
pass through, the plane containing the spines of the primary
row (Plate 5, Figure 5). The length of the secondary row
spines is normally less than the fully developed spines
of approximately the same growth stage on the primary
row, but they may grow to as much as 3 of the length
of the shell.

The spines themselves (that is, apart from their length
and orientation relative to the valve surface) are of
identical morphology in both primary and secondary spine
rows. They are rounded and smoothed on the commissural
side, and reflected at the edges so that a marked groove
or channel runs the length of the upper surface of the
spine (Plate 6, Figures 8, 10). The upper surface of the
spine carries typical shell surface growth striae; they taper
gradually to an extremely acute point. Even in the juve-
nile they are relatively thick at the base, and hence strong,

Page 64

THE VELIGER

Vol. 10; No. 1

for the size of the shell (e.g. Plate 5, Figure 5). For
instance in a shell 30mm long, the last secreted primary
spines are 2.1 mm thick at the base, tapering to a fine point
over a spine length of 33mm. In the largest adults that I
have seen the spines are generally relatively shorter than
this, but obviously extremely strong. For example in a
specimen 50mm long, the latest formed spines are only
24mm long, but are 3mm thick at the base: they there-
fore taper relatively rapidly.

Concentric lamellae

These are only developed at the anterior end of the shell
in the form of flaring, recurved, sharp concentric rings,
sometimes as much as 3mm high. As they are traced
across the main disc they grade insensibly into low, irreg-
ular, rounded concentric rings; in mature specimens they
may become completely obsolescent just anterior to the
posterior carina.

Hysteroconcha dione group

[Hysteroconcha dione (LINNAEUS, 1758) ]
This species group can be recognised by the small pre-
spinous dissoconch (about 3mm long), the corresponding-
ly smaller size of the adult shell (commonly less than
50mm long), and the possession of sharp raised concentric
lamellae over the whole of the main disc, and not just
confined to the anterior end.

Spines

The morphology of the individual spines is much the
same as in Hysteroconcha lupanaria, but the following
minor differences in arrangement and morphology appear
to be relatively consistent.
1. On any one valve the first spines secreted in ontogeny
are generally coincident in the primary and secondary
spine rows.

Explanation

Figure 8: Pitar (Hysteroconcha) lupanaria (Lesson), x 33. Same
specimen as figures 4 to 7. An enlarged view of the primary spine
row; note the comparative regularity of spine secretion with respect
to the concentric ornament; generally any two spines are separated
by 3 to 4 concentric ridges.

Figure 9: P (H.) dione (LinNAEus), x3; Recent, America; Saul
Collection. The primary (outer in this view) spine row delimits the
edge of the corcelet; inside this are two further differentiated areas,
conveniently termed the inner and outer escutcheon as both corre-
spond to the growth tracks of particular parts of the dentition.
Figure 10: P (H.) lupanaria (LEsson), x 4. Same specimen as fig.

2. The spines are relatively more numerous (compare
Plate 5, Figures 2 and 5) ; there are therefore fewer con-
centric laminae between each consecutive pair of primary
row spines.

3. The spines are relatively shorter (for example, primary
spine 14mm long in a 28mm long shell), broader at the
base (as much as 4.5mm wide in a 34mm long shell),
and more markedly flattened than in H. lupanaria (Plate
6, Figure 9).

4. Each spine is very clearly a continuation of a concentric
growth element.

5. There is a tendency for the secondary spine row to
have alternating introduction of spines on the two valves,
and for the primary row to have paired spines (Plate 6,
Figure 9).

A morphological detail especially marked in Hystero-
concha dione, but also present in H. lupanaria, is the
cessation of the secondary spine row at a particular growth
stage (Plate 6, Figure 9). This is generally when the
shell has reached a length of about 20 to 30mm. In some
specimens an initial cessation at this shell size may be
followed by a final pair of secondary row spines being
secreted after a considerable gap. For instance, in one
specimen the initial cessation came at about 20mm, but
there is another well developed pair of spines secreted
at a shell length of 38mm.

Concentric lamellae

The whole of the main disc between the carina carrying
the primary spine row and the lunule is ornamented
with spaced raised concentric lamellae, often slightly
recurved dorsally. Each lamina is about 0.6mm thick and
perhaps 2mm high; the anterior end of a lamella is
markedly flared, and the distance between the shell surface
and the lamina top may then be over 3mm (Plate 5,
Figure 1). The pre-spinous dissoconch does not have

of Plate 6

8. Note the ‘umbra’ zone dorsal to each primary row spine. This
zone carries growth lines clearly displaying that an epithelial tongue
continued to secrete the spine after it had become displaced dorsally
from the growing edge of the shell.

Figure 11: Hecuba scortum (LINNAEUS, 1758), x5; Recent, Indo-
Pacific; Saul Collection. View of the frills from a dorsal aspect
showing the shallow, though marked, radial gutters, and their
correspondence with the radial ornament.

Figure 12: Hecuba scortum (LINNAEUS), x3. Same specimen as
figure 11. View of the frills from ventral aspect. Note the structural
frill ribs — each of which corresponds to a radial gutter on the dorsal
surface of the frill (cf. fig. 11).

[The magnification is stated approximately for all figures]

Tue VELIGER, Vol. 10, No. 1 [CarTER] Plate 6
ate

Figure 8

Figure 9
Figure

Figure 11

Vol. 10; No. 1

raised lamellae, though it does have low rounded con-
centric ornament. There may also be faint concentric
ornament on the corcelet and escutcheon (Plate 6, Figure

8).

SHELL STRUCTURE

Macroscopically a section through the shell reveals the
characteristic three shell layers, each clearly marked by
its differing texture (Text figure 1). The ectostracum is

THE VELIGER

Page 65

stracum. They are clearly due to a periodic extension of
the mantle beyond its normal shell edge limits, but secre-
tion continues unchanged during these periods of mantle
extension.

The boundary between the ectostracum and the meso-
stracum is gradational; there is a gradual fading away of
the crossed lamellae into the mesostracum. The meso-
stracum itself is composed of thin subparallel laminae and
carries many conspicuous secretion traces; it is otherwise
microscopically structureless.

Figure 1

Cross section through the shell of
Lupanaria (Hysteroconcha) dione (LINNAEUS, 1758)
The three shell layers correspond to the endo-, meso- and ecto-
stracum respectively. x 6.

white, with pink tinges near the shell surface, and clearly
composed of crossed-lamellae of calcite. The mesostracum
is structureless but has a very characteristic grey ‘greasy’
appearance like that of nepheline. The endostracum is
a light grey-white, and coarsely layered.

Microscopically (Text figure 2) the ectostracum is made
up of fairly coarse crossed lamellae — a large lamella
being about 0.05 mm wide at its thickest point — which are
always at right angles to secretion traces, and thus clearly
reflect the actual direction of shell secretion. Bundles of
crossed lamellae are terminated ventrally by strong secre-
tion traces; these presumably represent growth pauses of
greater or lesser extent. The individual calcite folia can
often be traced across these secretion traces, but the fact
that this is not always so — sometimes there is a completely
new and unrelated set of folia initiated on the ventral
side of the trace — lends credence to the suggestion that
the traces represent extended growth pauses. The individ-
ual folia are noticeably finer on the inner side of the
ectostracum.

The raised concentric ridges on the shell surface, and
therefore the spines, are entirely built of ectostracal shell
material that is identical with that of the main ecto-

The endostracum coincides with the area of shell inside
the pallial line, and abuts sharply behind the dentition
(Text figure 1). It is microscopically very similar to the
mesostracum, but it is clearly separated from that layer
by the thin pallial myostracum. There do not appear to
be any shell canals [sensu Omori & Kopayasut, 1963;
tubule (OBERLING, 1964) is already in use for a part of
the stomach diverticula].

INFERRED PATTERN OF SHELL SECRETION

There is a certain rhythmical activity in the mantle
edge of Hysteroconcha, similar to that in many less striking
venerids, which results in the secretion of concentric shell
ornament. The activity takes the form of the extension of
a narrow strip of mantle beyond the general shell edge,
and its reflexion to make an angle of about 60° with the
plane of the commissure. Whilst in this relatively exposed
position outside the shell, the mantle secretes a layer of
shell about 0.6mm thick. Generally the extended mantle
strip stretches from the edge of the lunule to the primary
spine row and thus the result is raised concentric lamellae
of similar disposition.

Page 66

THE VELIGER

Vol. 10; No. 1

Figure 2

Enlarged view of the valve edge (x30) showing only meso- and

ectostracum. Note the gradational boundary between the two

layers, the fact that the concentric lamellae are constructed entirely

of ectostracum, and the fact that the crossed calcite lamellae of

the ectostracum record the actual direction of shell secretion at
any point.

Upon this basic pattern is superimposed another:
about every third mantle expansion a small tongue of
mantle at the posterior end of the raised mantle strip
Starts to expand locally, secreting as it does so. This
local expansion is presumably caused by localised mantle
cell generation. The result is the gradual building of

Explanation

Figure 13: Hecuba scortum (LinNAEuS), x 2; Recent, Indo-Pacific;
Saul Collection. Right valve of a pair, showing the general morphol-
ogy, and an unsuccessful gastropod boring.

Figure 14: H. scortum (LinnaEus), x8; Recent, Indo-Pacific;
Saul Collection. Enlarged photograph of the posterior spine row;
note the pointed nature of the spines and the presence of a dorsal
keel.

Figure 15: H. scortum (Linnaeus), x 4. Same specimen as figure
14. Muricid predation localised on the mid-ventral border (cf. figs.
2,3 and 5).

a projecting spine of calcareous material, smooth under-
neath (analogous to the normal internal shell surface),
but bearing above the typical growth lines and micro-
ornament of the external shell surface. The tongue of
mantle continues to expand, all the time secreting and
closely adhering to the spine that is resulting. The edges

of Plate 7

Figure 16: Spondylus spec., x3; Recent, unlocated; McAndrew
Collection, Zoology Department, University of Cambridge. Photo-
graph of the under side (i. e. the side facing the opposing valve) of
a major spine. All the spines on this specimen have expanded tips
that one might suggest as paradigmatic for sensory mantle protec-
tion (N. B. This is only taken as a convenient example; I am in no
way implying that the spine figured should be so interpreted) .
Figure 17: H. scortum (LINNAEuS), x 3. Same specimen as figure
15. General view of the corcelet, and spine bearing posterior ridges.
[The magnification is stated approximately for all figures]

Tue VELIGER, Vol. 10, No. 1 [CarTER] Plate 7

etree

Figure 13

w Figure 14 Figure 15

Figure 16

Figure 17

Vol. 10; No. 1

of the mantle tongue curl upwards somewhat resulting
in reflexed smooth edges to the spine (Plate 6, Figure 8).
In specimens of Hysteroconcha lupanaria these reflexed
edges may meet above the spine thus completely obliter-
ating any sign of the upper ornamented surface of the
spine. Before the spine has reached its full length the
continued general secretion of the whole mantle edge has
resulted in further growth of the shell all round the
commissure. Hence the base of the spine becomes relatively
distant from the shell edge, but it continues to be lined
with epithelial material and to grow in length as is shown
by the marked ‘umbra’ zone immediately ventral to the
spine; this umbra zone is shown by a marked re-entrant
in the growth lines (Plate 6, Figure 10).

When the growth of the spine is completed it would
seem likely that the mantle tongue responsible atrophies.
Irregularities in the concentric growth ornament on the
main disc can often be correlated with the continued
growth of spines that are no longer right on the valve edge.

FUNCTIONAL INTERPRETATION
Spines

The following functions are considered:
1. The spines might serve as devices for stabilizing the
shell in a shifting substrate.
2. The spines may bear some relationship to the feeding
mechanism or water circulation system of the animal;
this would correlate with their location precisely at the
boundary of the siphonal area.
3. The spines might be interpreted as protective cover,
or support, for sensory mantle outposts giving relatively
early warning of the presence of predators, or silt-laden
water. That the spines themselves are sensory is a possibil-
ity that must also be considered.
4. The spines might be interpreted as structural defensive
mechanisms combating predation.

Rejected Interpretations

In view of the fact that the spines are not buried when
the animal is in its normal feeding position, the first
suggestion seems unlikely to be true. However, even if the
shell were totally buried, the spines would act as rela-
tively inefficient stabilising devices. Effective stabilisers,
by virtue of their working principles, must be broad and
flat, and have a large surface area oriented at right angles
to the direction of expected movement. The long pointed
spines of Hysteroconcha do not remotely approach this
paradigm. One might make the stabiliser function more
plausible by suggesting that the spines, since as a set
they are oriented more or less parallel to the substrate,

THE VELIGER

Page 67

prevent the animal from sinking into the substrate, 7. e.
that they act as a type of “snowshoe.” But again the same
objection may be made: the spines should paradigmat-
ically be flattened structures lying along the surface of
the substrate; in fact they are pointed and project at
right angles to the surface of the substrate.

In order to have an advantageous effect on either
feeding or water circulation systems, the spines would
need to act as a set to form either a grill or a mesh for
preventing the entry of harmful particles into the in-
halent siphon (cf. Rupwick, 1961). But this hypothesis
is precluded by the fact that the spines are oriented in
such a way as to conspicuously offer no grill or mesh
protection whatsoever to the siphons or siphonal gape
(Plate 5, Figure 4).

Dealing now with the third postulate: undoubtedly the
mantle tissue lining the spines retains certain of the sensory
abilities that the normal mantle edge possesses. However,
if the main object of the exercise is to achieve a piece
of sensory mantle edge as distal as possible from the
commissure edge (the spine then being inevitably secreted,
and serving only as a protective cover for the contained
tissue), one would expect to find that the mantle outpost
comprised at least the two outer folds of the mantle
edge: the outer fold in order to have secreted the spine,
and the middle fold in order to provide a sensory outpost.
Though I have not been able to procure a preserved
animal of Hysteroconcha, by analogy with the spines of
Spondylus (which carry only outer mantle fold epithelia
to their tip) it seems unlikely that a sensory mantle
outpost was the reason for the spines on Hysteroconcha.
An additional factor bearing against this hypothesis is that
a spine which functions as a protective sheath for an
early warning mantle should paradigmatically be broad
(commensurate with its not interfering with normal feed-
ing currents), convex downwards, and might be expected
to carry some sort of an expanded tip in order to provide
for as large an area of sensory mantle as possible at a
distance from the shell edge (cf. Plate 7, Figure 16). The
spines of Hysteroconcha are narrow, concave downwards,
and extremely sharply pointed.

However, though I think it unlikely that the primary
function of the spines is as protection for the exposed
mantle tissue that secretes them, it is an inevitable con-
sequence of all spine morphology that the spines them-
selves serve at least partly as sensory structures. Undoubt-
edly the contact of any browsing predator with the distal
end of the spines would result in the immediate closure
of the shell, and hence they do provide a relatively ‘early
warning’ of potential danger. Yet one would anticipate
that if this had been their primary function, the selection
pressure responsible for their perfection would have fa-

Page 68

THE VELIGER

Vol. 10; No. 1

voured individuals with spines like that of Plate 7, Figure
16 — for such a spine is itself every bit as effective as the
pointed spines of Hysteroconcha for serving as a sensory
structure, and carries the additional advantage of acting
as an efficient protection for the mantle tissue (sensory
or secretory) that lies beneath it.

Spines as a Protective Structure

The most promising of the initially postulated functions
is that the spines serve for the protection of the animal.

To be effective as a defence structure spines would be
expected to a) guard the most vulnerable part of the
animal; b) be as long as possible commensurate with
their being sufficiently strong; c) taper to a sharp point;
and d) beso oriented as to face the direction of approach
of expected predators. The spines of Hysteroconcha ap-
proach very closely to such a structural paradigm.

On a priori grounds one would expect small sized preda-
tors to experience more difficulty when preying on
Hysteroconcha than when preying on a similar shell with-
out the posterior spines. There are two reasons for this:
firstly, the spines increase the effective size of the shell and
hence help to prevent the wholesale ingestion of the shell
by predators whose mouth size dictates the size of food
that they eat (small fish, intraorally digesting asteroids) ;
secondly, the effect of a closely spaced set of sharp spines
must have a strong deterrent value against those predators
that actually have to open the shell before eating its
contents (extraorally digesting asteroids, gastropods,
perhaps Crustacea). Such aquarium observations as I
have been able to make confirm these a prior: postulates;
both Asterias rubens (LINNAEUS, 1758) and Buccinum
undatum (LINNAEUS, 1758) seem to (subjectively) ex-
perience more difficulty in attacking baited model shells
that possess spines than they do in attacking similar shells
but without spines.

From the experiments of CHRISTENSEN (1957) it has
become obvious that virtually nothing will protect a
bivalve from predation by a determined asteroid. Curis-
TENSEN induced starved asteroids to feed through wire
mesh, and also observed that they could extrude their
stomach into prey that was situated at the bottom of
a plasticine cylinder as high as the asteroids arms were
long. However, one of the conspicuous traits of the
normal feeding of Asterias is the very close application
of the oral region to the shell edge at the point of pene-
tration of the stomach lobes; indisputably the spines of
Hysteroconcha would serve to prevent this, and therefore
lessen the efficiency with which the two valves can be
gripped and pulled apart by asteroids.

The spines must also help to protect against gastropod
attack. When feeding normally Hysteroconcha lies with

just the spines and posterior part of the shell protruding
above the substrate. A roving gastropod therefore initially
makes contact with the spines, and in order to mount the
bivalve and chip the edge off the valves (the normal
technique of gastropods like Busycon and Buccinum) it
would first have to dig the bivalve out of the substrate
in order to get at the unprotected ventral margins. It is
significant that in every case that I have seen of a Hystero-
concha which can be inferred to have been killed by
muricid gastropods, the site of shell drilling has been on
these unprotected ventral margins (Plate 5, Figures 2,
By O))e

Concentric lamellae

It is far more difficult to postulate intuitively reasonable
functions for the concentric lamellae than it is for the
spines. As the anterior end of the shell is buried in the
substrate during normal life, the only credible postulates
will be related in some way to the burying of the shell,
or to its stability when buried. One might distinguish three
possibly discrete functions:

1. The lamellae assist the animal to bury itself.

2. The lamellae help to keep the animal stable in the
substrate after it is buried.

3. The lamellae help to prevent the animal being dug
out of the substrate by its invertebrate predators.

To be adapted for the first function, the lamellae would
not be well developed at the anterior end of the shell,
where they must add to the resistance encountered by the
animal as it pushes into the sand; and they would perhaps
be best developed mid-ventrally where they might be
expected to help the “streamlining” of sand past the shell
as the animal buried itself. Yet in Hysteroconcha the frills
are always best developed at the anterior end of the shell
and may (H. lupanaria) be obsolescent mid-ventrally.

It is difficult to dispute that the lamellae might have
some stabilising effect in a sand substrate, if Hysteroconcha
be compared with a similar shell not possessing lamellae.
On the other hand, this is an extremely difficult propo-
sition to test, and it does not seem reasonable to conclude
at the moment that this is the primary function of the
lamellae.

Again, it would seem possible that the lamellae —
particularly in view of their strong recurved development
at the anterior end of the shell, and in view of the known
behaviour patterns of asteroids and gastropods — might
serve as ‘barbs’ to prevent the shell from being drawn
out of the substrate by such predators. But, to put it
mildly, this is a relatively farfetched explanation.

I find none of these solutions totally satisfactory, and
it seems likely that the correct interpretation will prove
to be another as yet un-suggested explanation. It might

Vol. 10; No. 1

THE VELIGER

be that the combination of all three suggestions made
here has some validity, but it is fair to conclude that, in
our present state of knowledge, it is impossible to be
sure of the exact function of the raised concentric lamel-
lae. Indeed, the whole question of concentric ornament
sensu lato is one that needs careful comparative study
throughout the Bivalvia. The extremes of development of
similar raised concentric lamellae in such groups as the
Australasian Bassina must surely have some adaptive
significance; exactly what significance is at the moment
obscure.

Hecuba

The adult shell is of comparable size to Hysteroconcha,
a mature individual being about 75mm long. It is of
triangular shape with a conspicuous posterior carina de-
limiting the edge of the corcelet (Plate 7, Figure 13) ;
there is a small escutcheon. Anteriorly the main disc
carries spaced raised concentric frills which cease abruptly
at the border of the large lunule. It is generally possible to
recognise an inner and an outer lunule corresponding to
the external shell tracks of the inner and outer lateral
dentition. The inner ventral valve margins are weakly
serrated, and provide a very inefficient interlock.

ECOLOGY

I have no significant information on the ecology of this
animal.

DETAILED MORPHOLOGY

All specimens that I have examined are referable to the
species Hecuba scortum (LinNAEus, 1758) ; they mostly
derive from Ceylon, India and the east African coast.

Spines

The posterior carina of this species is furnished with
a set of spines analogous to the primary spine row of
Hysteroconcha (Plate 7, Figures 13, 14, 17); there is
no trace of a further secondary spine row. Unfortunately,
the spines on the posterior carina are usually worn in the
juvenile stages, making it impossible to ascertain at what
stage they first appear. However, assuming that they were
present as early as the first concentric frills, they were
certainly present by the time the shell had reached a
length of 3mm. Thereafter the spines are introduced at
regular intervals — about one every 2 to 3mm in the adult.
In large specimens the late spines become more and more
crowded, and may be only 1 mm apart.

Page 69

The spines themselves are fairly short (a long one
measures just over 4mm), broad at the base (the same
spine is 3mm wide at the base), and taper to a beauti-
fully moulded point (Plate 7, Figure 14). They are not
symmetrical about the carina, that portion anterior to the
carina being markedly the broader (Plate 7, Fig. 13). The
upper surface of the spine carries fine shell secretion
traces, and a deep median furrow which runs out to the
very tip of the spine. Underneath they are smooth with
texture typical of internal shell surfaces, and carry a
prominent keel (corresponding with the groove on the
dorsal spine surface) that tapers away in the ‘umbra’
zone immediately ventral to each spine.

Concentric frills

The raised frills stretch from the edge of the lunule to
a point almost exactly mid-ventral on the main disc. The
interval between successive frills gradually increases with
the size of the shell. In a specimen 37 mm long, the spacing
between the two last-formed frills is 4mm. There is a
general correlation between individual spines on the ca-
rina and individual anterior frills: it is sometimes, though
by no means always, possible to trace one into the other
right across the shell (Plate 7, Figure 13).

The dorsal side of the frills carries typical concentric
secretion traces and spaced shallow radial gutters corre-
sponding to structural “frill ribs” on the ventral side of
the frills (Plate 6, Figures 11, 12). This ventral side,
like the ventral side of the spines, is texturally similar to
internal shell surfaces. The frill ribs are about 1 mm apart,
and correlated with this the shell surface between succes-
sive concentric frills carries low rounded radial ribs, each
corresponding to a structural rib on the ventral side of
the adjacent frill. These radial ribs, and also the frill
ribs, become less conspicuous towards the central part
of the disc and are not present at all on the zone between
the posterior end of the frills and the posterior carina. The
ribs are in no way related to the weak marginal inter-
locking.

FUNCTIONAL INTERPRETATION

Because of the remarkable similarity of the basic struc-
tures, most of the comments on the process of secretion
and the possible functions of the spines and concentric
lamellae on Hysteroconcha apply equally well to Hecuba.
However, there are differences and some difficulties.

Spines

The spines, though very beautifully moulded structures,
are far shorter than is typical of Hysteroconcha lupanaria.
However, of the possible functional interpretations, it is

Page 70

the defensive paradigm that the spines of Hecuba most
closely approach. Even so, there remain nagging doubts
about the relatively short length of the spines; could they,
in fact, be selectively effective?

This seems to be another case (biology is already littered
with them, for example, the problem of the development
of the vertebrate eye) where there is an intuitive difficulty
in believing that the earlier, less well-adapted manifesta-
tions of a particular structure can possibly have carried
any selective advantages. There is no doubt that a strong
case can be made out for the defensive nature of the spines
of Hysteroconcha lupanaria. But the closely related H.
dione possesses spines that are morphologically almost iden-
tical except for their much shorter length. In fact, the
spines of many H. dione are absolutely no shorter than
those of Hecuba. It would obviously be foolish to attribute
a different function to the spines of Hysteroconcha dione
and H. lupanaria, and hence it is a reasonable suggestion
that the same function will probably apply to Hecuba. The
fact is that the Hysteroconcha spines, certainly in ontogeny
and probably in phylogeny, have passed through a stage
when they were absolutely no longer than the present
adult spines of Hecuba. One should obviously be extremely
careful before attributing a different function to each on
the completely subjective judgement as to how long a
spine must be before it acts as an advantageous defensive
structure.

Concentric frills

As in the case of Hysteroconcha, beyond the general
postulate that the raised concentric frills (or laminae) are
probably in some way concerned with the burrowing of
the shell, or with its stability when buried, it is not possible
to be sure exactly what their specific function is. How-
ever, granted the general postulate, it will obviously be
advantageous for the concentric frills to be as strong as
possible; in the case of Hysteroconcha this is achieved
by having relatively thick laminae, but in Hecuba the
same problem is solved by having thin frills together with
strengthening frill ribs.

This is of considerable interest, for though it has often
been suggested that radial structures in the Bivalvia are
for strengthening the shell, it is rarely possible to demon-
strate such a suggestion plausibly. The frill ribs do appear
to be a genuine radial feature that results in structural
strengthening. The overall appearance of the concentric
frills and their frill ribs (Plate 6, Figure 12) is strikingly
similar, and perhaps structurally analogous, to the fins of
a fish. The radial ‘ribs’ on the shell surface between suc-
cessive frills are clearly the result of secretion by the same
strip of mantle that is responsible for the secretion of the
frill ribs. Further support for a strengthening hypothesis

THE VELIGER

Vol. 10; No. 1

comes from the observation that the frill ribs become
less well developed towards the centre of the main disc,
and it is in just this spot that the frills themselves are
most commonly broken. Similar frill rib structures can be
seen in other groups of bivalves (e. g. Chamidae), and
a strengthening interpretation may also be relevant in
such cases.

However, the fact that the raised frills of Hecuba
carry strengthening frill ribs does not of itself help to solve
the problem of the function of the frills. Apart from the
observation that muricid predation in Hecubza is also pref-
erentially located on the unprotected ventral borders
(Plate 7, Figure 15), it remains to conclude that a precise
diagnosis of the function of this type of concentric orna-
ment is not at present possible.

CONCLUSIONS anp SUMMARY

This paper has discussed the morphology, and suggested
functional interpretations, for ‘ornamental’ features on
two of the more spectacular Recent bivalve genera. It
could perhaps be maintained that the spines and frills of
Hecuba and Hysteroconcha have no functional signifi-
cance, but such an interpretation is too sterile. When one
considers the remarkable convergent evolution implied
by the gross homeomorphy of Hecuba and Hysteroconcha
— two unrelated genera that live on opposite sides of the
northern hemisphere — it becomes vanishingly improb-
able that their convergent structures are not in some way
adaptive, and hence functional.

However, though it is obvious that much valuable anal-
ysis can be performed on dead shells, a satisfactory inter-
pretation of many bivalvian structures can only be
reached in conjunction with detailed studies of the living
animal. Since the palaeontologist is never in a position
to garner this type of additional data, except by the use
of homological and analogical comparisons with living
forms, it will be of interest to see how many of the
essentially “palaeontological” arguments presented in this
paper can be shown to be spurious, or irrelevant, when
more is known of the modern Hecuba and Hysteroconcha.
The lack of published information on the anatomy and
habits of spectacular tropical mollusca remains as one of
the largest gaps in modern understanding of invertebrate
biology. Yet the functional interpretation of fossil and
Recent Bivalvia is critically dependent upon the amount
of information available on the biology of Recent species.
All too often the assumption is made that because an
animal is common, and its habits are locally well known,
it would be undesirable to ‘clutter up the literature’ with
the more pedestrian details of its ethology and mode of

Vol. 10; No. 1

THE VELIGER

Page 71

life. I can only repeat my view, that it is these very
pedestrian details that are so necessary for successful func-
tional analysis, and that such details are conspicuous only
by their rarity. One must hope for some fundamental
contributions to this field of research in the near future.

ACKNOWLEDGMENTS

I should like to thank Professor O. M. B. Bulman, F.R.S.,
for making available the facilities of the Sedgwick Mu-
seum, Cambridge. Dr. A. Bidder and Mr. S.P Dance
kindly gave access to collections in their care, and Dr. J. D.
Taylor generously lent me a preserved specimen of Spon-
dylus collected by him on Seychelles.

In my efforts to obtain information on the ecology of
Hysteroconcha I have been fortunate in receiving help
from many people, and I should like to thank Professor
FE M. Bayer, Drs. M. Keen and F. Haas, Mr. J. Q. Burch
and Mrs. G. Varnardo in this respect. Mr. A. C. Simpson
and Dr. D. A. Hancock kindly made available facilities
for studying the behaviour of asteroids and gastropods at
the Fisheries Laboratory, Burnham-on-Crouch. Dr. R. T.
Paine drew my attention to the interesting paper by J. J.
Gilbert; and I am particularly grateful to Drs. M. J. S.
Rudwick and R. Cowen for their critical readings of the
manuscript, and for frequent helpful advice.

This work was carried out during the tenure of
a Commonwealth Scholarship awarded by the British
Council.

LITERATURE CITED

Carter, Rosert M.

1967. On the nature and definition of the lunule, escutcheon
and corcelet in the Bivalvia. Proc. Malacol. Soc. London
74 (4): in press

CurIsTENSEN, A. M.

1957. ‘The feeding behaviour of the seastar Evasterias troschelli
STIMPSON. Limnol. Oceanogr. 2: 180 - 197

DoszHANsky, THEODOSIUS

1941. Genetics and the origin of species. 2" ed.
Dopce, HENRY
1952. A historical review of the mollusks of Linnaeus. Part 1.

The classes Loricata and Pelecypoda. Bull. Americ. Mus.

Nat. Hist. 100: 1 - 263

Geist, V.
1966. The evolution of horn-like organs. Behaviour 27:
175 - 214

Gitsert, J. J.

1966. _Rotifer ecology and embryological induction. Science
151: 1234 - 1237
KEEN, A. MyRA
1958. Sea shells of tropical West America; marine mollusks

from Lower California to Colombia.
Stanford, Calif. (Stanford Univ. Press)

OseEr.inG, J. J.

1-xi + 624 pp.; illus.

1964. | Observations on some structural features of the pelecy-
pod shell. Mitteil. Naturforsch. Gesellsch. Bern, new ser.,
20: 1 - 63
Omort, M. & J. Kopayasut
1963. On the micro-canal structures found in the shell of Arca
navicularis BRUGUIERE and Spondylus barbatus REEVE.
Venus 22: 274 - 280
PITTENDRIGH, C. S.

1958. Adaptation, natural selection and behaviour. In:
Roe & Simpson (ed.) “Behaviour and Evolution”, Yale Univ.
Press: 390 - 416

Rupwick, M. J. S.

1961. The feeding mechanism of the Permian brachiopod
Prorichthofenia. Palaeont. 3: 450 - 471

1964a. The inference of function from structure in fossils.
Brit. Journ. Phil. Sci. 15: 27 - 40

1964b. The function of zigzag deflexions in the commissures of
fossil brachiopods. Palaeont. 7: 135-171

1965. Sensory spines in the Jurassic brachiopod Acanthothiris.
Palaeont. 8: 604 - 617

Page 72

THE VELIGER

Vol. 10; No. 1

Mode of Feeding and Diet,

and Synthesis of Studies on Marine Pelecypods

from Tomales Bay, California

DON MAURER

Old Dominion College, Biology Department, Norfolk, Virginia

INTRODUCTION

THE PRESENT INVESTIGATION has a threefold purpose:
1) To describe the mode of feeding and diet of bivalves
previously unreported; 2) To provide further evidence to
interpret pelecypod-sediment associations described in an
earlier study; 3) To synthesize field and laboratory work
heretofore treated separately.

In a series of papers on marine pelecypods from Tomales
Bay, California, I have been primarily concerned with the
ecology of bivalves in relation to sediment and the
interpretation of these relationships. More specifically,
Maurer (1967) reported on the relationship between
distribution, abundance, and size of Tellina buttoni Da.u,
1900, T: salmonea (CarPENTER, 1864), Mysella tumida
(CarpPENTER, 1864), Transennella tantilla (Goutp, 1852),
and Lyonsia californica Conrab, 1837, to sediment type.
In addition, Maurer (1967 a, 1967b) described filtering
and burial experiments respectively to explain the re-
sponse of mollusks to sediment. One observation which
emerged from these experiments was that, at least for
a few of the pelecypods, the sediment might represent
an important food source, or that their mode of feeding
may be dependent on the type of substrate in which they
live. If this is true some of the pelecypod-sediment associa-
tions recognized by the author might be accounted for.

STEPHEN (1928, 1929, 1930, 1932), Horme (1949,
1961), Yonce (1949), Sanpers (1958, 1962), and
Powto (1966) have contributed information on feeding
habits of tellinids. At present there are very few data on
feeding of Tellina buttoni and Tellina salmonea, or, for
that matter, on any of the other species treated in the
study. The following discussion is restricted mainly to the
tellinids and Transennella tantilla with some brief com-
ments on Mysella tumida and Lyonsia californica.

23508

OBSERVATIONS

Siphons of Tellina buttoni are separate, long and flexible.
The exhalant siphon is slender and tapers towards the
aperture, while the inhalant siphon, which terminates
abruptly, is more muscular than the exhalant siphon. On
a conservative estimate, 7’ button may extend its siphons
four or five times the shell length which varied from 0.19
to 1.85 cm in length.

In an aquarium when Tellina buttoni burrowed into the
substrate, it would gently test the sediment with its in-
halant siphon and begin to inhale the substrate like an
elephant’s trunk. This feeding behavior of T. buttoni
compares favorably with that of British tellinids described
by YoncE (1949). Tellina buttoni has also been observed
to probe the substrate with its inhalant siphon. Suction
from its siphon disturbs sediment and causes fine matter to
be raised into suspension. The mollusk then inhales the
suspended particulate material.

During filtering experiments it was noticed that test
animals would occasionally reach up with their inhalant
siphons and take carmine particles out of suspension
(Maurer, 1967 a). Nevertheless, the inhalant siphon was
usually occupied with the bottom deposits or material
raised by its movements into suspension just above the
bottom.

During burial experiments Tellina buttoni would rarely,
if ever, form any holes in the overlying clay (MAuRER,
1967 b). And depending on the thickness of clay it would
almost never extend its siphons above the clay-water
interface. Following the burial experiments siphons of
dead T: buttoni were frequently packed with clay and test
animals that survived invariably contained some clay. For
T. button there was increased ingestion of clay with in-
creased amounts of clay used in the experiments.

Vol. 10; No. 1

THE VELIGER

Page 73

Concerning the diet of Tellina buttoni some preliminary
observations can be offered. Diatoms were extremely
abundant in outdoor aquaria where the clams were main-
tained. Upon dissection T: buttoni was found to be full of
needle-like species of Navicula. This diatom lay obliquely
in ctenidial chambers of the bivalves blocking water
circulation. Other diatoms, Coscinodiscus and Bidulphia,
were observed to be scattered throughout the alimentary
tract and stomach. It is unknown whether T: button
ingests such large quantities of diatoms in nature. An
unsuccessful attempt was made to identify stomach con-
tents of freshly preserved material aboard ship. However,
material found in feces of bivalves just after collection
consisted of quartz, biotite, quartz diorite, schist, and the
foraminifer Elphidium. An empty ostracod carapace
was found in the area of the digestive gland of one of these
mollusks. Several days after these pelecypods had been
placed in aquaria with glass beads as substrate, they were
examined and found to contain glass beads in the alimen-
tary tract. ;

Several citations to feeding of tellinids in nature can
be alluded to. For example, STEPHEN (1928) noticed
that during the spring diatoms appeared almost exclu-
sively in the gut of several species of Tellina. YONGE
(1949) reported that a species of Macoma fed on deposits
rich in diatoms. SANDERS et al (1962) listed a species of
Tellina as containing sand, diatoms, and detritus in its
stomach.

Both Tellina salmonea and T: buttoni have well devel-
oped longitudinal and transverse muscles in their siphons
and a cruciform muscle that is characteristic of members
of the Tellinacea (YonceE, 1949). The muscle is a sheath
of fine, tough fibers attached ventrally to the base of the
siphons and contributes to the flexibility and siphonal
activity of forms with long siphons. Moreover, the siphons
of Tellina salmonea are translucent, and as seen in T:
buttoni, its siphons may extend 4 or 5 times the shell
length which varied from 0.20 to 1.87 cm in length.

When Tellina salmonea was placed in aquaria without
any substrate it would almost immediately raise its siphons
up into the water. When sediment was present, its inhalant
siphon would settle upon and probe the substrate,
alternating continuously from water to substrate. For the
main part siphons of T: salmonea were characteristically
in the water as it would often function as a suspension
feeder. Duality of feeding habit exhibited by this tellinid
is even more marked than that of T. buttoni, and T.
salmonea showed a preference for suspension feeding.
Hoime (1961) has suggested a dual feeding habit for
a British tellinid as he described the disposition of its
siphons while feeding. He concluded that it may be able
to feed as a suspension feeder and is probably not restricted

to a deposit feeding habit. PoHto (1966), who worked
on Tagelus californianus (Conran, 1837), another tellinid
which occurs in Tomales Bay, stated that it was not an
active deposit feeder, but that it fed on suspended material.
These references and my observations indicate that tellin-
ids are not as restricted to a deposit feeding habit as was
previously thought.

In filtering experiments Tellina salmonea had the lowest
filtering rate (Maurer, 1967a). Yet it was commonly
observed to inhale suspended carmine particles. Depending
on the thickness of clay in the burial experiments T:
salmonea would form holes in the heavy, viscous clay
(Maurer, 1967b). This behavior was never observed in
T. buttont. Siphons of T: salmonea were rarely packed with
clay, whereas siphons of the other tellinid were generally
filled. ARMsTRONG (1965) described the difficulty en-
countered by clams covered by sand. Unless bivalves could
extend their siphons through the new cover to the surface
or maintain a sand free passage, they would perish. This
might explain in part the high mortality of 7: buttonz in
the burial experiments.

Concerning the diet of Tellina salmonea it was observed
to inhale copepods and the diatom Coscinodiscus which
was extracted from the gills of this tellinid. These diatoms
were also found in the stomach, and chains of diatoms
were noticed in the digestive tract as well. Although
T. salmonea was maintained in aquaria under conditions
similar to T: buttoni, at no time was T: salmonea observed
to be filled with diatoms or contain as much detrital
material as T: buttoni.

Only a few general comments on the mode of feeding
and diet of Transennella tantilla, Mysella tumida, and
Lyonsia californica can be offered. Siphons of Tr. tantilla
are short and free distally, but fuse proximally. Siphons
contain frills or small tentacles around the apertures and
this tentacular arrangement seems to be characteristic of
many suspension feeders. In filtering experiments 77. tan-
tilla had a relatively high filtering rate and was seen to
inhale suspended carmine and India ink particles. In
burial experiments it suffered the least amount of mor-
talities and survived the rigors of the viscous clay. The
fact that Tr. tantilla occurs intertidally to subtidally might
be accounted for by its ability to remain shut for long peri-
ods of time, and thereby circumvent the deleterious
conditions imposed by the overlying clay. The diatoms
Navicula, Bidulphia, and Coscinodiscus, which were dis-
sected from the other test species, were also found in 7?.
tantilla. In addition, Nitzschia and Melosira were dis-
covered within its gill system. These examinations were
made on specimens that had lived in the outdoor aquaria
for several weeks. Examination of specimens which had

Page 74

THE VELIGER

Vol. 10; No. 1

remained in the aquaria a couple of days showed some
glass beads, quartz, and shell fragments in their feces.

Mysella tumida was observed to feed on copepods and
diatoms and the latter were dissected from the stomach.
Upon dissection of Lyonsia californica no debris was found
in its stomach. Other than this no observations were
recorded on diet or mode of feeding of L. californica.
Kettocc (1915) briefly described and figured L. saxicola
(Barrp, 1863) and he stated that its ciliary mechanism
was very similar to that of L. californica. On this tenuous
basis it is provisionally suggested that L. californica is a
suspension feeder.

DISCUSSION

Results of field studies by Swan (1952), Pratr (1953),
Pratt & CAMPBELL (1956), and KrisTENSEN (1957) in-
dicated that size of Mya arenaria LiInNAEUus, 1758, Venus
mercenaria (LINNAEUS, 1758) and Cardium edule Lin-
NAEUS, 1758 may be strongly influenced by turbidity, rate
of sedimentation, and nature of the substrate. Moreover,
Mavrer (1967) arrived at a similar conclusion concerning
the distribution and abundance of Lyonsia californica,
Tellina buttoni, Tellina salmonea, and Mysella tumida in
Tomales Bay, California. Further it was revealed that the
average size of the latter 3 species was statistically associ-
ated with sediment particle size. Laboratory studies re-
ferred toin ALLEN’s (1963) review and especially the work
of LoosANorr and co-workers showed the deleterious
effects of increased sedimentation (organic and inorganic
particles) on respiratory and feeding activities of Mytilus
edulis LINNAEUS, 1758, Crassostrea virginica, and Venus
mercenaria. About the response of different ctenidial
types to filtering experiments it might be pointed out that
Mytilus edulis is a filibranch, Crassostrea virginica is a
pseudolamellibranch, and Venus mercenaria is a eulamelli-
branch; however, all feed as suspension feeders. Evidence
on harmful effects of increased sedimentation to bivalves
was provided by Maurer (1967a, b). Results of these
experiments on mollusks from Tomales Bay agree generally
with those cited from the literature.

Since the recognition of the pelecypod-sediment associ-
ations, one of the main objects of the investigation has
been to obtain information concerning response of mol-
lusks to sediment, in terms of food, protection, larval sites,
and turbidity. From experiments and observations the
author considers, at least for a few of the pelecypods, that
sediment as a food source may dictate mode of feeding
or that certain conditions of turbidity are related to the
deposition of particular sediment types.

On the effect of turbidity on distribution, abundance,
and size of clams it has been asserted that an increase or

inhibition of growth might be related to sediment. Pelecy-
pods that live in turbid environments waste energy in
clearing palps and gills from excessive particulate matter,
produce greater amounts of pseudofeces, and consequently
spend less time feeding. On the other hand, in a relatively
clean environment the mollusks obtain maximal nutri-
tional benefit of feeding time. Turbid conditions inhibit
growth; clean conditions foster it. Pratt & CAMPBELL
(1956) clearly stated this proposition in their study of
Venus mercenaria and LoosanorF (1962) voiced a similar
opinion. The present author considers this effect of sedi-
ment or sedimentation to be an important factor influ-
encing the pelecypod-sediment associations with which he
was concerned.

SYNTHESIS
oF FIELD ann LABORATORY STUDIES

For Tellina buttoni statistical analyses indicated that
average size was positively associated with sediment size
(Maurer, 1967). Laboratory work suggested some inter-
esting features that might account for the association on
the basis of turbidity tolerance and mode of feeding.
Filtering ability of 7: buttoni was generally high and it
has been observed to inhale suspended material. Yet even
with small concentrations used in the filtering experiments,
T: buttoni commonly ingested more ink, carmine, and clay
particles than Tellina salmonea and Transennella tantilla.
In burial experiments Tellina buttoni again ingested large
amounts of clay that it was generally unable to handle
and these amounts were almost always greater than in
the other test animals. Furthermore its production of
pseudofeces was usually greater in heavier loads of clay.
The relative amount of pseudofeces formed agreed sub-
stantially with that of other bivalves exposed to excessive
amounts of material described by Loosanorr (1962).
These mollusks were suspension feeders and had different
ctenidial structures than T. buttonz. The preference of T-
buttoni for silty-sand in light of laboratory work indicates
that this species functions chiefly as a deposit feeder. The
same evidence also indicates that there is a critical sedi-
ment size which inhibits growth. Sediment, which contains
more than 30% silt and clay, is probably deposited in a
turbid environment and is too densely packed to allow
easy use of siphons to filter, feed, and digest efficiently.

For Tellina salmonea statistical analyses showed that
average size was positively associated with sediment size
(Maurer, 1967). Laboratory work revealed some inter-
esting adaptations. As with T. buttonz, the association may
be explained on similar grounds, that is, food and tolerance
to turbidity. With T: salmonea the siphons were generally
extended above the substrate, but were occasionally seen

Vol. 10; No. 1

THE VELIGER

Page 75

to inhale bottom deposits. Contrary to this in the filtering
experiments it had the lowest filtering rate and in the
burial experiments T. salmonea was rarely full of clay.
Distribution of this tellinid in substrates that range in
particle size from silt through pebbles, particularly in the
coarse sediment, suggests a suspension feeding habit. It
was not clear whether TJ: salmonea was more sensitive
to turbid conditions or had more difficulty inhaling large
amounts of clay and silt than T: buttoni. On the basis of
its more limited distribution in coarse sediment it is sug-
gested that T: salmonea is more sensitive to turbid condi-
tions than 7: buttoni. When T: salmonea occurred in
sediment with as much as 30% silt and clay, its size was
limited and it was smaller than-individuals which lived
in sediment with little or no particulate material. In view
of laboratory studies the preference of 7: salmonea for
detritus-free, medium sand to shell-sand indicates that
this species functions as a suspension feeder in clean, coarse
sediment and as a deposit feeder in fine substrates. This
result tends to confirm suspicions raised by Maurer
(1967a) on possible dual feeding habit of 7: salmonea
and T: buttont.

For Mysella tumida statistical analyses showed that
average size was negatively associated with sediment size.
From its distribution in sediment types comparable to the
distribution of Tellina buttoni, and its single performance

in the filtering experiments, it is tentatively concluded that |

M. tumida can filter fine material better than the tellin-
ids. Studies by BALLENTINE & Morton (1956) showed
that a closely related genus was a suspension feeder which
could feed selectively on diatoms. It appears that the
distribution, abundance, and size of MM. twmida is influ-
enced by sediment size. However, its negative association
with sediment size indicates that conditions of turbidity or
sediment as a source of food do not have the same effect
on Af. tumida as they do on the tellinids.

Average size of Transennella tantilla did not show a
significant association with sediment size (Maurer, 1967).
Its ubiquitous distribution in the bay seemed to illustrate
its independence from characteristic sediment types. Lab-
oratory work did not disclose any dual feeding habit as
was noticed in the tellinids. Relatively high filtering eff-
ciency, low mortality rate and absence of clogging in
burial experiments, tentacular frill work on siphons define
Tr. tantilla as a suspension feeder. It was concluded that
sediment does not exert a controlling influence on distri-
bution, abundance, and size of Tr. tantilla as it does on
the tellinids and Mysella tumida.

Although Lyonsia californica had no significant pelecy-
pod size - sediment size associations, its maximal distribu-
tion and abundance were found to be in mud with 85%
to 100% silt and clay. It appears that L. californica is
adapted to extremely turbid conditions and heavy sedi-

mentation encountered in the southern portion of the bay.
The nature of these adaptations has not been examined,
but work on the functional morphology of its feeding
apparatus would probably explain its rather restricted
occurrence in very fine sediment.

ACKNOWLEDGMENTS

Research was carried out at the Pacific Marine Station,
Dillon Beach, California during the summer of 1964.
The writer is grateful for the aid extended by Drs. Joel W.
Hedgpeth and Edmund H. Smith, and by John Sibert
who kindly identified the diatoms for the author. In-
vestigation was supported by funds from a National
Science Foundation cooperative grant in Oceanography
made to the University of Chicago.

LITERATURE CITED

ALLEN, JoHN A.

1963. Ecology and functional iorphology of molluscs, — in
Harorp Barnes (ed.), Oceanography and marine biology, an
annual review. 1: 253 - 288. George Allen and Unwin, Ltd.,
London

ArMsTRONG, Ler R.

1965. = Burrowing limitations in Pelecypoda.

The Veliger
7 (3): 195 - 200; 4 text figs.

(1 January 1965)
BALLENTINE, Dorruy «& JouN Epwarp Morton
1956. Filtering, feeding and digestion in the lamellibranch

Lasaea rubra. Journ, Mar. Biol. Assoc. U. K. 35: 241 - 274

Hoitme, Norman A.

1949. The fauna of sand and aud banks near the mouth of
the Exe Estuary Joum. Mar. Biol. Assoc. U. K. 28:
189 - 237

1961. Notes on the mode of life of the Tellinidae (Laimelli-
branchia) . Journ. Mar. Biol. Assoc. U. K. 41: 699-703.

KEttoce, J. L.
1915. Cihary tacchanisius of lamellibranchs with descriptions

of anatomy. Journ. Morph. 26: 625 - 701

KRISTENSEN, INo\ vt

1957. Dilferences in density and growth ina cockle population
in the Dutel Wadden Sea. Arch. Neerl. Zool. 12: 351 - 4-4
LoosaNnore, Vicror b.
1962. Elects of turbidity on some larval and adult) bivalves.
Proc. Gull aud Carib. Fish. fast. lth An. Sess., 80-94
Loosanorr, Vicor b. 8 Jvames Bo ENGrt
1940. Effect of different concentrations of plankton fonns up-
on shell movements, rate of water pumping and feeding, and

fattening of oysters Anat. Rec. 84: 86

1947. Feeding of oysters im relation to density of microorgan-
Isis. Science 105: 260 - 261

Page 76

THE VELIGER

Vol. 10; No. 1

Loosanorr, Victor L. & FRaNcEs D. TOMMERS

1948. Effect of suspended silt and other substances on the rate
of feeding of oysters. Science 107: 69 - 70

Maurer, Don
1967. Pelecypod-sediment associations in Tomales Bay, Cali-
fornia. Malacologia _(in press)

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)
Ponto, Ross H.

1966. A note on the feeding behavior in Tagelus californianus

(Bivalvia : Tellinacea) . The Veliger 8 (4) : 225; 1 text fig.
(1 April 1966)
Pratt, D. 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, D. M. « D. A. CampBEeLy

1956. Environmental factors affecting Venus mercenaria.
Limnolog. « Oceanogr. 1: 2 - 17

SANDERS, Howarp L.
1958. Benthic studies in Buzzards Bay I. Animal sediment
relationships. Limnol. « Oceanogr. 3: 245 - 258
Sanpers, H. L., E. B. Goupsmire, E. L. Mitts « G. E. Hampson
1962. A study of the intertidal fauna of Barnstable Harbor,
Massachusetts. Limnol. « Oceanogr. 7: 63 - 79
STEPHEN, A. C.
1928. Notes on the biology of Tellina tenuis Da Costa.
Journ. Mar. Biol. Assoc. U. K. 15: 683 - 702
1929. _ Notes on the rate of growth of Tellina tenuis Da Costa
in the Firth of Clyde. Journ. Mar. Biol. Assoc. U. K. 16:
117 - 129
1930. Studies on the Scottish marine fauna: additional obser-
vations on the fauna of the sandy and muddy areas of the tidal
zone. Trans. Roy. Soc. Edinb. 56: 521 - 535
1932. Notes on the biology of some lamellibranchs in the Clyde
area. Journ. Mar. Biol. Assoc. U. K. 18: 51 - 68
Swan, Emery FREDERICK
1952. ‘The growth of the clam Mya arenaria as affected by the
substratum. Ecology 33 (4): 530 - 534
YonceE, CuHartes Maurice
1949. On the structure and adaptations of the Tellinacea,

deposit-feeding Eulamellibranchia. Phil. Trans. Roy. Soc.
B 234: 29 - 76

Vol. 10; No. 1

THE VELIGER

Page 77

The Behavioral Role and the Structure

of the Aesthetes of Chitons

BY

PAUL OMELICH

Electron Microscope Laboratory, University of California, Berkeley, California

94720

(Plates 8 and 9)

INTRODUCTION

Younc AMPHINEURA OF THE ORDER Polyplacophora,
commonly called chitons, have long been known to be
negatively phototactic. During hours of daylight they are
found under and between closely spaced rocks and in other
places of relative darkness. Older chitons with worn and
encrusted valves seem to be indifferent to the general
illumination. The valves of chitons contain numerous sense
organs. Due to their gross structure, these organs, called
aesthetes, have been assumed to be photoreceptors by
most investigators.

I have attempted to determine the function of the
aesthetes by electron microscopy, electrical recording of
nervous impulses, and behavioral observations.

MATERIALS anp METHODS

BEHAVIORAL

Observations regarding photoreception were made using
a beain of light from a dissecting microscope lamp nar-
rowed to an incident beam of approximately 3mm dia-
meter. The beam of light was directed onto different
areas of the animals and their movements were observed.

Mechanoreception of the aesthetes was investigated by
observing the movements of the animals while exposing
their shell surfaces to touch of dissecting needles and a
soft camel’s hair brush.

In order to test chemical sensitivity of the chiton’s dorsal
surface, two thin glass cylinders were wax-sealed to the
valves on each side of Mopalia lignosa (Goutp, 1846).
The animals were placed on a marked glass plate and
submerged in sea water so that the open end of the glass
cylinders remained above the level of the water (Plate 8,
Figure 1). Homogenates of the sea star, Pisaster ochraceus

(BranpT, 1835), and mid-tidal red and green algae
were made. The homogenates were placed in contact with
the surface of the shell by partially filling the glass cylinder
on one side of the animal. The cylinder on the other side
was always filled with unaltered sea water. Movement of
the animal, toward or away from the side exposed to the
homogenates, was observed. In addition to the homo-
genates, dilute hydrochloric acid was placed in one of
the cylinders and the animals were observed.

ELECTRICAL RECORDING

I first attempted to record the mass electrical response
from the whole animal with amplifier and oscilloscope.
The chiton was immobilized between two chambers filled
with sea water with light-shielded electrodes placed in
each. Alternate exposure to light and darkness of the dor-
sal surface of the valves produced no recognizable change
in the large slow muscle potentials continually seen. Iso-
lating the anterior valve alone between the two sea-water
bathed electrodes likewise produced no pattern related to
exposure to light. Glass capillary microelectrodes with tips
of approximately 20 diameter were placed over indi-
vidual micraesthete caps and unit recording was attempted.
Here again no impulses could be detected. Fine insulate!
steel recording clectrodes were then introduced into the
main aesthete nerve canals of broken valves and only
photoelectric effects, probably of the elctrode itself, were
observed. Due to these repeated failures at electrical re
cording from the aesthetes I did not pursue this approach
further.

ELECTRON MICROSCOPY

Mopalia lignosa (Goutp, 1846), M. hinds (Reeve,
1847), and Tonicella lineata (Woop, 1815) were the three
chiton species used throughout this work. All were ob-

Page 78

THE VELIGER

tained along the California coast from Monterey to Point
Arena. Shells were broken into fragments of less than
0.5mm* and immediately fixed in 1% osmium tetroxide
in sea water for 30 minutes to one hour at 0 to 5°C.
Following fixation they were rinsed with sea water and
decalcified in a 5% solution of ethylene-dinitrilo-tetra-
acetic acid in sea water for periods of 12 to 24 hours.
Dehydration was in a graded series of ethanol. Some of
the tissues were infiltrated with Araldite and some with
Epon. Sections were cut with both diamond and glass
knives on a Porter-Blum microtome. Both unsupporting
and Formvar-coated copper grids were used. The sections
were examined with an Akashi TRS-50EI electron micro-
scope.

Potassium permanganate in sea water and acetate-ver-
onal buffered osmium tetroxide fixatives were used, but
without success. In addition, double fixation with 1%
osmium tetroxide in sea water, before and after decalci-
fication, was done. Here the processing resulted in destruc-
tion of much fine structure. Undecalcified shell fragments,
fixed, dehydrated and embedded in hard Epon were
sectioned with a diamond knife, but the ultrathin sections
disintegrated upon the microtome trough. Thicker sections
proved to be useless.

RESULTS ano DISCUSSION

Probably all chitons with exposed valves possess organs,
called aesthetes, embedded in the shell. These organs are
apparently sensory and always occur in two sizes, megal-
aesthetes and micraesthetes. There are no aesthetes of
intermediate size. In addition to the aesthetes, some chitons
possess larger organs branching from the same intra-shell
nerves, called eyes. These eyes possess a cornea, lens and
retina (MosELEy, 1884 and 1885). None of the species of
chiton on the California coast have been reported to
possess eyes, but all species that I have observed or that
other investigators have reported on have the two types
of aesthetes.

Vol. 10; No. 1

HISTORY

MarsHALL (1869) found in the shells of chitons canals
of two sizes terminating in cup-shaped caps. He regarded
the tissue found in these canals as respiratory in function.
Van BEMMELEN (Husrecut, 1882) proposed that the
organs found in these canals were homologous with the
bundle of fibers supporting the spines of the girdle. MosE-
LEY (1885) reported that a Dr. W. B. Carpenter first
observed the perforate structure of the tegmentum of
chitons (date not specified). MosELEy examined many
alcohol-preserved chitons and found three types of organs:
those which he called eyes, and two other similar, but
much smaller structures. He named these smaller struc-
tures aesthetes. He thought the aesthetes to be organs of
touch and the eyes to be photoreceptive organs. MosELEY
followed the pathways of the tubes leading from the
eyes and aesthetes and he was of the opinion that they
terminated in the parietal (pallial, branchial) nerves.
The size of the eyes found ranged from 188u to 42 in
diameter. The arrangement of the eyes varied from an
irregular scatter to rows, either concentric or radiating
to the apex of the tegmentum.

Biumricu (1891) made a microscopic study of the
aesthetes of various chitons. He identified these organs as
nervous structures and suggested they were probably
photoreceptors. PLate (1902) did an extensive study on
the general anatomy of the chitons, including the aesthetes,
but with apparently no new discoveries regarding the
nature of these organs. Nowrkorr (1907, 1909) did
detailed anatomical work on the aesthetes and was con-
vinced that they were nerves. LELoup (1940) looked at
the chitons of the California coast and noted the occur-
rence and gross appearance of the aesthetes, but did no
microscopic work.

BEHAVIORAL OBSERVATIONS

Photoreception: Crozier & Arey (1918) found that a
shadow of a fly 6 feet away caused Chiton tuberculatus

Explanation of Plate 8

Mopalia lignosa (Goutp, 1846)

Figure 1: Chitons with glass cylinders attached for observations on
chemoreception. cyl. - glass cylinders for receiving substances to be
presented to the surface of the shell.

Figure 2: Photomicrograph of an aesthete complex. me - megal-
aesthete; mi - micraesthete; cv - clear vacuoles; s - surface of shell.

(x 590)

Figure 3: Electron micrograph of a longitudinal section through
the megalaesthete cap. tu - tubes of cap; mc - megalaesthete cone
material. (x 5500)

Figure 4: Electron micrograph of a cross section of the megal-
aesthete nerve between or below the point of micraesthete branching.
cv - clear vacuoles; nu - nucleus. (x 7500)

Tue VELIGER, Vol. 10, No. 1 [Ome icy] Plate 8

Figure 3 Figure 4

photographs by Paut OMELICH

Vol. 10; No. 1

Linnaeus, 1758, to halt motion temporarily! These au-
thors stated that a miscellaneous collection of individuals
may be caused to separate into two general size groups by
sunlight, the larger moving into, the smaller away from
lighted areas. The periphery of the girdle was found to be
sensitive to a light increase, but the reaction was not
oriented. Only a depression of the girdle to the substrate
occurred. When they amputated the girdle the smaller
chitons still oriented away from the light while the
older ones went toward it.

Heatu (1899) observed the chitons of Monterey Bay,
California and found that they remain out on their feeding
ground only when the day is foggy or dark. He observed
that Ischnochiton magdalenensis (= I. heathiana Berry,
1946) adults are found under rocks during the day and
only come out to feed at night.

Crozier (1921) found an almost perfect correlation
between the degree of erosion of chitons and the relative
illumination of the situations frequented. Uneroded chi-
tons were found to be photonegative even to moonlight.
They were photopositive or indifferent when completely
eroded. He found that the age of the animal was not
important, only the degree of erosion.

Douctas (1952) observed that the normal negative
response to light exhibited by C'yanoplax dentiens (GouLp,
1846) depends upon the presence of all or most of its
aesthetes. He removed the aesthetes of young photonega-
tive C. dentiens by eroding their valves with emery cloth.
These chitons were then often found in the direct rays
of the sun along with older, naturally eroded animals.

I exposed portions of Mopalia lignosa to illumination
from a narrow beam of light from a dissecting microscope
lamp. A direct beam to any part of the girdle produced a
clamping of the girdle to the substrate similar to that
observed by other investigators. Illumination to one side
of the valves only caused the animals to move away from
the light. Illumination of either the anterior or posterior
ends likewise produced a negative taxis.

It is remotely possible that the actual photoreceptors
of chitons are not contained within the shell, but are
located in the tissue beneath, the light being transmitted
through the dense calcium carbonate of the valves. But
this seems very improbable due to the relatively low inten-
sities of light necessary to produce an oriented response.
When only diffuse sunlight is allowed to enter through a
slot into a blackened wooden box containing chitons,
orientation is obtained (AREY & Crozier, 1919) and the
animals orient even to moonlight (Crozigr, 1921). It
seems likely, therefore, that photoreception does occur
in the shell of chiton. Because the aesthetes are the only
nervous structures found within the shells it is quite
probable that they are photoreceptive.

THE VELIGER

Page 79

Mechanoreception: Mose.ey (1885) believed that the
aesthetes were tactile organs. He thought this on the basis
of the slightly protruding caps which he observed on the
aesthetes. AREY & Crozier (op. cit.) state regarding the
aesthetes, “Definite evidence as to their functional signi-
ficance has been completely lacking.” They did compre-
hensive tactile, chemical and light behavioral work on
Chiton tuberculatus. They concluded that in this species
“There are no tactile receptors in the shell plates.” But
regarding another kind of chiton they observed minute
projecting hairs on the valves, and they stated (without
describing the methods used), “We find that the teg-
mentum of Ischnochiton purpurascens (ADAMS, 1845) is
very sensitive to touch.”

I tested Mopalia lignosa, M. hindsii and Tonicella
lineata for mechanoreception. Lightly dragging a steel
dissecting needle across the surface of the shells con-
sistently produced a clamping reaction of the girdle to the
substrate. But brushing with a camel’s hair brush produced
no effect. It seems likely that the response found as a
result of touching with the hard steel needle is probably a
result of shell distortion or vibration, or both, being trans-
mitted to the foot and girdle where no doubt mechanore-
ceptors are found.

My longitudinal sections of the megalaesthete caps show
them often slightly elevated above the surface of the shell
(Plate 8, Figure 3). The micraesthete caps are consistently
depressed. Moserey (1885) and Briumricn (1891)
showed both the megal- and micraesthetes slightly ele-
vated from the surface. Nowikorr (1909) saw them
level with the shell. The position of the caps higher
than that of the shell surface is not a prerequisite
to mechanoreception, but touch receptors firmly fixed on
their peripheries to the hard calcium carbonate of the
shell and not protruding above it would most likely operate
by recording minute distortions of the shell as do campani-
form sensilla of the insect cuticle. But this does not seem
likely due to the extreme hardness of the valves.

While this investigation did not conclusively prove that
either the megalaesthetes or the micraesthctes are or-
gans of touch, it scems to me that it is very unlikely that
they function in this way.

Chemoreception: BarNawevv (1960) found that in their
natural habitat several species of chitons of the same types
which I used cat food in the following order of total
quantity:

1. Algae (red and brown)
. Diatoms
. Bryozoa
. Hydroids
. Barnacles
. Sponges and mollusks

DM OS © PO

Page 80

THE VELIGER

Vol. 10; No. 1

FepER (1959) studied the food of the starfish Pzsaster
ochraceus (BRANDT, 1835) and stated that Pisaster was
observed to feed primarily on gastropods and chitons.

Thinking that exposure of the surface of the valves of

chitons to extracts of their most favored food and their
common predator might reveal the possession of chemo-
receptive function of the aesthetes, I applied homogenates
of algae and starfish to the surface of one side of the shell
only, one substance at a time (Plate 8, Figure 1). The
glass cylinders containing the materials protruded above
the surface of the water so that none of the homogenates
could reach any other receptor areas of the animals. The
cylinder opposite the food- or predator-filled one was
filled to the same level with sea water from the chiton’s
habitat tank. Exposure of the shells to algae, starfish
extract and dilute hydrochloric acid did not produce a
visible response. Then I dropped several drops of the
algae onto the girdle on one side only. Again there was no
response. But when I dropped the starfish extract onto the
mantle the animals initiated motion anteriorly and slightly
away from the exposed side. Dilute hydrochloric acid
evoked the same responses as the starfish extract, only
more pronounced. These experiments were repeated a
sufficient number of times to justify the conclusion that
it is improbable that the aesthetes are chemoreceptive
organs for the materials tested.
Microscopy: Invertebrate photoreceptors are much more
diverse than vertebrate ones. They comprise several dif-
ferent structural types with only a few common features.
Electron microscope studies have generally concentrated
on the “higher” invertebrate phyla. Much work on the
arthropod ommatidia and the cephalopod retina has been
done. Aside from the Cephalopoda little work has been
done on mollusks. I think that this investigation is the first
attempt to ascertain the fine structure of the presumed
photoreceptors in the Amphineura. The aesthetes of chi-
tons are thought to be photoreceptors only because of
behavior and gross structure. Therefore I prepared these
organs for electron microscopy.

The invertebrate photoreceptor region is generally com-
posed of cells containing organelles called rhabdomeres.
These rhabdomeres in mollusks and in many other inver-
tebrates contain pigment granules. In addition, the rhab-
domeres are composed of an array of tubules and microvilli

probably derived from cell membranes. The average
rhabdomere tubule is approximately 600 A in diameter
(Moony, 1964). Eakin (1963) adds the generalizations
that many mitochondria are situated near the light sensory
apparatus and that an axon-like fiber leads from the
basal end of the receptor cell. Eakin (op. cit. ) states that
in the annelid-arthropod-mollusk complex a fibrillar and
centriolar assembly like that of the vertebrate photore-
ceptor cilium has not been seen. Mitter (1958) found an
exception to this generalization in the ocellus of the
scallop Pecten irradians Lamarck, 1819. Here he found
ciliary filaments leading into the bases of the lamellae.

Miiter (1960) described the tubular units of rhabdo-
meres in general as varying in diameter from 0.04 -0.12y.
He looked at Pecten ocelli (1958) and described globular
appendages that are derived from cilia. Irregular matted
microvilli extended from the photoreceptor cell. The dia-
meter of these microvilli tubules was approximately 0.07.

Eakin (1963) studied the eye of the garden snail
Helix aspersa Mier, 1776, and found the photorecep-
tor cell studded with microvilli, each about 0.1 in dia-
meter. The microvilli are radially arranged parallel to the
long axis of the cell. They extend as much as 12y to the
surface of a large structureless lens. No ciliary or centriolar
apparatus was found.

Wo ken (1958) looked at the retinal structure of
Octopus and Sepia. He found the retina made up of
rhabdomes analogous to those of the arthropod eyes.
Each rhabdome consists of four visual units radially
arranged. These units contained regularly arranged tu-
bules approximately 0.05 in diameter. A central space
containing pigment cells with screening pigment granules
separated the rhabdomes. ZoNANA (1961) saw Sepia
microvilli of an average diameter of 0.1y.

In this investigation light microscopy showed that the
general arrangement of the aesthetes of Mopalia lignosa
is similar to that found in chitons by other researchers
in that two types of organs are present, the smaller micr-
aesthetes branching from the megalaesthete (Plate 8,
Figure 2).

Electron microscopy revealed some new detail. The
megalaesthete cap in longitudinal section (Plate 8, Figure
3) shows a regular tubular honeycomb extending from the
surface of the shell almost to the center cavity. This

Explanation of Plate 9

Mopalia lignosa (GouLp, 1846)

Figure 5: Electron micrograph of a longitudinal section of the
aesthete nerve below the branching of the micraesthetes. nf - neuro-
fibrils; pi - pigment granules. (x 20000)

Figure 6: Electron micrograph of a longitudinal section of the
aesthete nerve farther down than that of Figure 5. nf - neurofibrils;

pi - pigment granules. (x 22000)

Tue VELIcER, Vol. 10, No. 1 [OmeE.icH] Plate 9

photographs by Paut OmELICH

Vol. 10; No. 1

cavity under the cap is partially filled with a mass of what
appears to be tubules extending from a central body which
I called the megalaesthete cone because thick sections
and light photomicrographs show the outline of this
material as conical in shape. A finely structured layer
seems to line the cavity. The micraesthete caps were
similar in configuration, but smaller. They did, however,
generally appear recessed from the shell surface in contrast
to the slight protrusion of the megalaesthetes. MosELEY
(1885) saw both the megalaesthetes and the micraesthetes
loosely occupying their canals and slightly protruding
above the shell. Perhaps the dilute nitric acid he used for
decalcification of the shell was the cause of this. BLuM-
ricH (1891) and Nowikorr (1909) saw the caps conti-
guous with the shell surface.

The tubular appearing structures wthin the aesthete
caps are approximately 0.06p in diameter. This is slightly
larger than the smallest diameter of microvilli, 0.04-0.12y,
as reported by Miter (1960), but smaller than the
0.25. microvilli of the polychaete Nereis (Eakin, 1963).
There seems to be a lack of protoplasmic connection
between the cap itself and the megalaesthete cone material
below. This cone with its tubules seems most likely to be
the photoreceptive site, the microvilli radiating out into
the cavity where light transmitted through the passages
in the cap can be recorded. The cone tubules are slightly
smaller than the cap passages and still within the range of
invertebrate microvilli. Another possibility is that in better
preserved material the tubules of the cone could be seen
to continue out into the passages of the cap where the
photoreceptive site would be even closer to the surface
of the shell. Since no such connections were seen, this
seems unlikely. But it is possible that the decalcification
process destroyed connections to the cap passages. A less
harsh method of removing the calcium carbonate should
confirm this.

The cap material appears to be hard in nature and
not primarily protoplasmic. It may be simply a special
configuration of shell material which both protects the
underlying sense organ and provides a light guide to
increase sensitivity over what it would be if the receptors
were embedded in ordinary shell.

Cross sections of the megalaesthete nerves show large
clear vacuoles near the center (Plate 8, Figure 4). These
vacuoles are also seen in photomicrographs (Plate 8,
Figure 2) (BLumricn, 1891, Nowrkorr, 1909). These
vacuoles seem to be present in the nerve only at the point
of micraesthete branching. Their function is unknown.

Longitudinal sectioning of the megalaesthete nerve
reveals clumps of dense bodies, especially on the periphery

THE VELIGER

Page 81

(Plate 9, Figures 5 and 6). Perhaps these are gliosomes
within the cytoplasm of neuroglial cells. They appear
similar to those found in the cockroach (Pirea, NisHIoKA
& Bern, 1962). But until the fine structure of these dense
bodies can be seen, I shall call them only pigment granules.
Neurofibrils are characteristically seen in the presumed
axons from the aesthetes.

SUMMARY

Chitons possess organs in their shells which are appar-
ently sensory. These organs, called aesthetes, seem to be
photoreceptors. Behavioral tests indicate that it is very
improbable that the aesthetes act as mechano- or chemo-
receptors. Electrical recording from these organs did not
give positive results.

Analysis of electron micrographs indicates that struc-
tures somewhat analogous to the microvilli of invertebrate
photoreceptors do exist. A tubular structure, located in
the caps of the aesthetes, was found. I believe that these
tubes must either act as chambers for photoreceptive
tubules similar to microvilli or as channels for transmission
of light to receptor organs below. Since protoplasmic con-
nections to the caps were not identified, it would seem
that the photoreceptive material is located just under the
cap. Tubules suggestive of microvilli were found there.

ACKNOWLEDGMENT

Much of this study was done in the Department of
Zoology, University of California, Berkeley, under the
guidance of Dr. Donald M. Wilson. I am indebted to him
for his help. The photographic work of Mr. Philip
Spencer of the Electron Microscope Laboratory is very
much appreciated.

LITERATURE CITED

Arey, Lesuig B. « W. J. Crozier
1919. The sensory responses of Chiton.

29: 157 - 260

BaRNAWELL, Eart B.

Journ. Exp. Zool.
(April 1919)

1960. The carnivorous habit among the Polyplacophora.
The Veliger 2 (4) : 85 - 88; plt. 19 (1 April 1960)
BLumRIcH, JOSEPH

1891. Das Integuiment der Chitonen.
Zool. 52: 404 - 476; plts. 23 - 30; 1 text fig.

Zeitschr. Wissen.

(21 August 1891)

Page 82

Crozier, W. J.

1920. Note on the photic sensitivity of the chitons. Amer.
Naturalist 54: 376 - 380 (July-August 1920)

Amer. Naturalist 55:
(May-June 1921)

1921. | Homing behavior in chiton.
276 - 281; 1 fig.
Crozier, W. J. & Lesiie B. AREY
1918. On the significance of the reaction to shading in chiton.
Amer. Journ. Physiol. 46: 487 - 492 (1 August 1918)
Douctas, LEE

1952. The photic responses of some chitons of Monterey Bay
and vicinity. Adv. Marine Invert. Nat. Hist. Reprt., 45.
Zool. Dept., Univ. Calif. Berkeley (unpubl. student report)

Eakin, RicHARD MARSHALL

1963. Lines of evolution of photoreceptors. In: General
physiology of cell specialization, D. Mazia and A. Tyter, eds.,
393 - 425, McGraw-Hill, New York
FepER, Howarp M.
1959. The food of the starfish, Pisaster ochraceus, along the
California coast. Ecology 40 (4): 721-724; 2 text figs.
(October 1959)

Heatu, Haroip

1899. The development of Ischnochiton. Zool. Jahrb., Abt.
Anat. Ontog. 12: 567 - 656; pits. 31-35; 5 text figs.
(22 July 1899)
Husrecut, A. A. W.
1882. A contribution to the morphology of the Amphineura.

Quart. Journ. Micr. Sci. 22: 212 - 228; 11 figs. (April 1882)
LeLoup, E.
1940. Caractéres anatomiques de certains chitons de la Céte

Californienne. Mém. Mus. Hist. Nat. Belg., Serv. 2,
Fasc. 17: 1-41; 94 figs. (30 April 1940)

MarsHALL, W.

1869. Note sur l’histoire naturelle des chitons. Arch. Néer-

land. sci. exactes et nat. 4: 328 - 341; plt. 5

THE VELIGER

Vol. 10; No. 1

Mitier, Wituiam H.

1958. Derivatives of cilia in the distal sense cells of the retina of
Pecten. Journ. Biophys. and Biochem. Cytol. 4 (2) : 227 to
228; plts. 120, 121 (25 March 1958)

1960. The cell. Vol. 4, chapt. 6, pp. 325 - 364; 19 figs. Eds. J.
BracHet &« A. E. Mirsky. vii, ix, xi-xv+511 pp.; illus.
Academic Press, New York

Moopy, M. FE

1964. Photoreceptor organelles in animals.

43 - 86; plt. 1; 6 text figs.
Mose tey, H. N.

1884. On the presence of eyes and other sense-organs in the
shells of the Chitonidae. Ann. Mag. Nat. Hist., ser. 5, 14:
141 - 147 (August 1884)

1885. On the presence of eyes in the shells of certain Chitoni-
dae, and on the structure of these organs. Quart. Journ.
Micr. Sci. 25: 37-60; plts. 4-6 (January 1885)

Nowrkorr, M.

Biol. Rev. 39:
(February 1964)

1907. Uber die Riickensinnesorgane der Placophoren nebst
einigen Bemerkungen iber die Schale derselben. Zeitschr.
Wiss. Zool. 88: 153 - 186; plts. 10, 11; 4 text figs.

(29 October 1907)

1909. Uber die intrapigmentaéren Augen der Placophoren.
Zeitschr. Wiss. Zool. 93: 668 - 680; plt. 29; 2 text figs.

(12 October 1909)
Pipa, RupotpH L., Ricuarp S. NisHioKa « Howarp ALLAN BERN

1962. Studies on the hexapod nervous system. V. The ultra-
structure of cockroach gliosomes. Journ. Ultrastr. Res. 6:
164-170; 8 text figs. (April 1962)

WOLKEN, JEROME J.

1958. _ Retinal structure. Mollusc cephalopods: Octopus, Sepia.
Journ. Biophys. and Biochem. Cytol. 4 (6): 835 - 838; plts.
420, 421; 1 text fig. (25 November 1958)

ZoNANA, Howarp V.

1961. Fine structure of the squid retina.
Hopkins Hosp. 109 (5): 185 - 205; 14 text figs.

Bull. Johns
(Nov. 61)

PHILIPPINE
SEA

| MANILA

MANILA
BAY

vy Z 25 MILES
CORREGIDOR 1.7” te

Va
TAAL .
1BAUAN

BALAYAN A \ TABAYAS
Ine BAY : ELS/
= a
MARICABAN
fo I. SD BATANGAS
: BAY

MINDORO

SIBUYAN
SEA

Vol. 10; No. 1

THE VELIGER

Page 83

A Quantitative Sampling of the Mollusks

of Batangas Bay, Philippines

BY

JEAN M. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plates 10 and 11; 1 Map; 2 Text figures)

ON A RECENT TRIP TO THE PHILIPPINES we had an
opportunity to visit one of the world’s most noteworthy
collecting areas for mitrids, Batangas Bay in southern
Luzon. We are greatly indebted to Mr. Fernando Dayrit
for having arranged a day-trip to this area so that Mr.
Cate and I could see it and sample the collecting there.

Two experienced divers employed by Mr. Dayrit (Plate
10, Figure 1) preceded us to the collecting grounds, and
worked there for a total of three full days. With the excep-
tion of miscellaneous crabs, octopus and the like which
they kept for food, we brought home with us their entire
three-day catch. While we had expected to find a large
number of mitrids in this material, as Batangas Bay is
known to be one of the most productive areas in the world
for this group, we were impressed by the large quantities
and the diversity of other forms as well. This prompted
me to put together the following checklist to record the
various forms found, and to indicate their relative abun-
dance within a comparatively limited area at that par-
ticular time. It should be stressed that this list is not
intended to be a complete catalog of all mollusks that have
been or are to be found in Batangas Bay, but merely of
those collected during the three days in mid-November
1966 by the two expert divers.

Batangas Bay, situated about 100 kilometers south of
Manila, is reached over a good paved road (see dotted
line on Map). Our destination was Bauan Town; more
specifically, the area in front of the CalTex refinery,
where the company personnel graciously permitted us to
park on the fenced and guarded property and to use the
convenient supply of fresh water.

Batangas Bay is about 16 km wide at its mouth, and
some 12 km long from the refinery area southward to
the open sea; its depth (according to a hydrographic

chart) ranges to approximately 140 fathoms, though it
seems fairly shallow near the shore and a long pier had to
be built to accommodate oil tankers and other oceangoing
vessels (see Plate 10, Figure 2). The collectors were free-
diving in about 4 to 5 fathoms, approximately a mile or
two south-southwest of the refinery (see Map).

The substrate is an extremely fine silty black mud
several inches deep at the point where we attempted to
collect in knee-deep water; it is probably of much the
same type over most of the bay, judging by the material
found clinging to the specimens brought in by the divers.
To our disappointment at first, we found that the shallow
water near the refinery was polluted by oil seepage, and
the shells we were able to collect without swimming out
into deeper, cleaner water were all dead. Furthermore,
the nature of the substrate made it very difficult to see
bottom after the silt had been stirred up by our wading,
for great lingering clouds of suspended material obscured
the view of any mollusk tracks or even of the bleached,
white dead shells. However, our disappointment was soon
forgotten when we saw the first sackful of mollusks spread
out for our inspection by the divers; it is a tribute to
their skill that they were able to locate so many small
items that would ordinarily have been extremely difficult
to find in the soft mud. Subsequent dives resulted in an
array of specimens to satisfy the most exacting collector
(see Plate 10, Figure 3): cones, terebras, turrids, bursas,
cassis, harpas, strombids and many other forms were dis-
played before us, in addition to 23 species of Mitra and
Vexillum. In all, 903 specimens were collected, repre-
senting 114 species in 33 families. I have listed them all
below in systematic order.

The Mitridae provided the greatest number of species,
with a total of 23; however, a number of these were

Page 84 THE VELIGER Vol. 10; No. 1
Table 1 represented by only a few specimens, so that the total
of 152 mitrid shells came as a poor second to the Conidae,
Number of which had the second highest number of species (16),
Species Specimens but an extremely high number of specimens (355).
GASTROPODA The group of mitrids contained a few surprises for me
= in that certain species usually considered relatively com-
Melaniidae 2 4 :
Thiaridae 1 1 mon were represented by only one or two specimens (e. g.,
Tarbinidae 9 9 Mitra filaris [1], M. granatina [1 dead] M. clathrus [2]) ;
Architectonicidae 2 4
Cerithiidae 2 41 (CATS) AOIDIA
Xenophoridae 1 1 es
: iaridae
Strombidae 3 4 Turbinidae
Cypraeidae 3 9 Architectonicidae
Naticidae 4 5 Cerithiidae
Cassididae 3 4 Xenophoridae
Bursidae 1 25 Strombidae
Tonnidae 1 5 Cypseiee
Ficidae 1 2 ee
Cassididae
Muricidae Z 3 Biretioe
Columbellidae 3 4 Tonnidae
Cymatiidae 1 9 Ficidae
Buccinidae 5 50 Muricidae
Nassariidae 2 4 Columbellidae
: ae Cymatiidae
Fasciolariidae 2 4 Bees
Olividae 3 34 Nassariidae
Mitridae 23 152 Fasciolariidae
Harpidae 1 2, Olividae
Cancellariidae 3 4 Mitridae
Turridae 7 118 Harpidae_
Conidae 16 355 Sere
Terebridae 10 39 Conidae
Atyidae 1 2 Terebridae
Ellobiidae 1 1 Atyidae
106 888 Ellobiidae
SCAPHOPODA
SCAPHOPODA Dentaliidae
Dentaliidae 2 6 PELECYPODA
PELECYPODA Seaaee
penenips ESS SEE eneridae
Carditidae 1 2 Tellinidae
Veneridae 1 1 Psammobiidae
Tellinidae 1 1 5 ue 2 payee yo!
Psammobiidae 2 5
5 9 Figure 1
Totals 113 903

Illustrating the number of species in each family-group

Explanation of Plate 11

Figures 1a, 1b: Mitra suturata REEVE, 1845 (x 33) Figures 5a, 5b: Vextllum rectilateralis (SowERBY, 1874) (x 23 )
Figures 2a, 2b: Vexillum coccincum (Reeve, 1844) (x §) Figures 6a, 6b: Vexillum rusticum (REEVE, 1845) (x 3)
Figures 3a, 3b: Vexillum ligatum (A. AvaMs, 1853) (x 33) Vigures 7a, 7b: JVenillum spicatum (REEVE, 1845) (x 34)
Figures 4a, 4b: Vexillum marmorea (A. Avams, 1853) (x 32) Figures 8a, 8b: Vexillum vulpecula (LINNAEUS, 1758) (x 14)

Figures 9a, 9b: Pusia spec. indet. (x 5)

THE VELIGER, Vol. 10, No. 1 [J. M. Care] Plate 10

boat

Figure 1 Figure 2

TL BIG 6S REE LIE EIR

“Ge 5
¥
\¢ 2

Figure 3

Figure 1: Divers from Samar who collected the shells in this study
Figure 2: General view of Batangas Bay
Figure 3: Results of the first dive made by the two divers in Figure 1

photographs by Jean M. Carte

Tue VELIGER, Vol. 10, No. 1 [J. M. Cate] Plate 11

ae ea BE is hte CO BAS

Figure 1 a Figure 1 b Figure 2 a

Figure 7 a Figure 7 b = Figure 8 a i Figure 8b Figure 9 a

Vol. 10; No. 1

Table 2
Batangas Bay Mitridae included in the survey
Mitrinae:
1. Mitra (Cancilla) clathrus (GMELIN, 1791)

Ue

8.

THE VELIGER

(see CERNOHORSKY, 1965, plt. 14, fig. 25)

. Mitra (Cancilla) filaris (Linnatus, 1758)
(see CERNOHORSKY, 1965, plt. 15, figs. 33, 33 a)
. Mitra (Cancilla) flammea Quoy & Gaimarp, 1833

(see CERNOHORSKY, 1965, plt. 15, fig. 29)

. Mitra (Cancilla) flammigera REEVE, 1844

(see CERNOHORSKY, 1965, plt. 15, fig. 30)

. Mitra (Cancilla) granatina LaMarck, 1811

(see CERNOHORSKY, 1965, plt. 14, fig. 23)

. Mitra strigillata SowErRBy, 1874

(see CERNOHORSKY, 1965, plt. 15, fig. 31)
Mitra suturata REEveE, 1845

(see Plate 11, Figures la, 1b)
Mitra variegata (GMELIN, 1791)

(see CERNOHORSKY, 1965, plt. 14, figs. 24, 24a)

Vexillinae:

).

10.

IM,

W

13.

14.

Dior

De

Vexillum acupictum (REEVE, 1844)

(see CERNOHORSKY, 1965, plt. 19, fig. 81)
Vexillum amanda (Reeve, 1845)

(see CERNOHORSKY, 1965, plt. 18, fig. 73)
Vexillum coccineum (REEvE, 1844)

(see Plate 11, Figures 2a, 2b)
Vexillum ligatum (A. Adams, 1853)

(see Plate 11, Figures 3a, 3b)
Vexillum marmorea (A. ApamMs, 1853)
see Plate 11, Figures 4a, 4b)
Vexillum obeliscus (REEVE, 1844)

see CERNOHORSKY, 1965, plt. 19, fig. 80)

. Vexillum radix (Sowersy, 1874)

see CERNOHORSKY, 1965, plt. 20, fig. 96)

. Vexillum rectilateralis (SowErBy, 1874)

see Plate 11, Figures 5a, 5b)

. Vexillum rusticum (ReEEve, 1845)

see Plate 11, Figures 6a, 6b)

. Vexillum semisculptum (ApAMS & REEVE, 185())

see CERNOHORSKY, 1965, plt. 18, fig. 67)

. Vexillum spicatum (REEvE, 1845)

see Plate 11, Figures 7a, 7b)
Vexillum todilla (MicHEts, 1848)

(see CERNOHORSKY, 1965, plt. 21, fig. 104)
. Vexillum verrucosum (REEVE, 1845)

(see CERNOHORSKY, 1965, plt. 19, fig. 84)
Vexillum vulpecula (Linnaeus, 1758)
(see Plate 11, Figures 8a, 8b)

Pusia spec. indet.

(see Plate 11, Figures 9a, 9b)

Number of
Specimens

2
1

11

3

20

152

Page 85

Page 86

GASTROPODA
Melaniidae
Thiaridae
Turbinidae
Architectonicidae
Cerithiidae
Xenophoridae
Strombidae
Cypraeidae
Naticidae
Cassididae
Bursidae
Tonnidae
Ficidae
Muricidae
Columbellidae
Cymatiidae
Buccinidae
Nassariidae
Fasciolariidae
Olividae
Mitridae
Harpidae
Cancellariidae
Turridae
Conidae
Terebridae
Atyidae
Ellobiidae

SCAPHOPODA
Dentaliidae

PELECYPODA
Carditidae
Veneridae
Tellinidae
Psammobiidae

THE VELIGER

Vol. 10; No. 1

Figure 2

Illustrating the number of specimens in each family-group

conversely, relatively uncommon species were taken in
greater numbers (e. g., Vexillum marmorea [4], V. todilla
[5], Vv verrucosum [4]), and there were two forms that
were entirely new to me, despite the fact that I already
had a large amount of Batangas Bay material in my
collection before visiting the area.

Table 1 indicates the number of specimens for each
of the 33 molluscan families in the study; Text figure 1
illustrates the relative abundance of the various family-
groups of shells found, and Text figure 2 shows the number
of specimens.

Table 2 is a list of the mitrid species contained in this
collection, the Mitridae being the only group for which
I feel qualified to make specific identifications with
reasonable accuracy. The reader is referred to CERNOo-
HorSKY (1965) for illustrations of certain of these; other-
wise the shells are illustrated here on Plate 11.

At least one specimen of each mollusk species included
in the study will be preserved in a special-collection unit,
so that subsequent work may be done on various groups
as time permits, or as interested workers may wish to
examine certain spccics.

I wish to thank Mr. Dayrit again for having made this
trip possible, and Dr. Myra Keen for assistance in deter-
mining gencra unfamiliar to me. The text figures and map
are by Mrs. Emily Reid of The Veliger staff.

LITERATURE CITED

CERNOHORSKy, WALTER OLIVER
1965. The Mitridae of Fiji (Mollusca: Gastropoda). The

Veliger 8 (2): 70-160; plts. 13-23; 11 text figs.; 1 map;
2 tables; index (1 October 1965)

Vol. 10; No. 1

THE VELIGER

Page 87

Epitonium (Asperiscala) billeeana (DUSHANE & BRATCHER, 1965)

non Scalina billeeana DuSHANE & BraATCHER, 1965

BY

HELEN DUSHANE'

SINCE THE PUBLICATION of the description of Scalina
billeeana DUSHANE & BraTCHER, 1965, it has been sug-
gested that the species was erroneously placed in the
genus Scalina. CLENCH & TURNER (1951, p. 287) regard
Scalina as a subgenus of Amaea while KEEN (1958, p.
278) treats Scalina as a full genus. Regardless of the course
adopted, the species in question, Scalina billeeana, shows
no particular affinity to either of these two generic taxa.

We are provisionally placing Scalina billeeana in the
genus Epitonium, subgenus Asperiscala DE Boury, 1909,
which is characterized by fine spiral striae. The species
“Scalina” billeeana differs from any known species of
Amaea or Scalina in having an open umbilicus. It also
differs in having a lighter, more fragile shell. The
complete number of spiral ribs appears at a very early
stage. In Epitonium billeeana the numerous spiral ribs
are present on the third whorl with little appreciable
increase in number on subsequent whorls, while Scalina
ferminiana Dat, 1908 (p. 316; plt. 8, fig. 8) shows
one strong spiral rib on the first three whorls, adding
more spiral ribs in the following whorls until the final
number is reached.

Another difference is the habitat. Epitonium billeeana
occurs in shallow water and is parasitic on the coral
Tubastrea tenuilamellosa (MiLNeE-Epwarps & Hare,
1848), while Scalina species are dredged on soft, silty
mud bottoms.

In general proportions, Epitonium billeeana resembles
E. (Asperiscala) frielei (Dati, 1889) (CLENCH & TuR-
NER, 1852, p. 301, plt. 139), which also has an umbilicate
shell with fine axial and spiral striae. Epitonium frielei
is stated to be a flat white shell, while the living animal of
E. billeeana is bright yellow and the shell is yellow tinted
with a thin brown periostracum. Also, E. frielei has fewer
costae and its spiral ribs are weaker; consequently, the
relationship between the two species is probably super-

* 15012 El Soneto Drive, Whittier, California 90605

ficial. In habitat, E. frielei differs in having been taken in
depths of 63 to 135 fathoms off the coasts of North
Carolina and Florida.

Epitonium billeeana cannot be reconciled with any of
the genera currently used for Eastern Pacific or Western
Atlantic Epitoniidae but the author is reluctant to create
a new genus for it until there has been a complete study
made of the family names on a world-wide basis and the
ecological relationships are better known.

Since the publication of the original description of
Epitonium (Asperiscala) billeeana, additional localities
for the species have been reported:

Bahia de las Animas, Gulf of California, Mexico.
Several live specimens with egg masses taken by
diving at 5 to 30 feet, September 1965, Don Wobber,
Hillsborough, California. — Reported as common.

Punta Colorado, Guaymas area, Sonora, Mexico. From
siftings taken at extreme low tide, 1 beach specimen,
November 1965, S.S. Berry.

Guaymas area, Sonora, Mexico. Dredged at 40 feet,
one live specimen, Mrs. Camden Ernest, San Anto-
nio, Texas.

Cape San Lucas, Baja California del Sur, Mexico.
Five live specimens on coral taken by diving at 10
feet, April 3, 1966, James H. McLean. Size record:
long. 24.6mm, lat. 15.3 mm.

Manzanillo, Mexico. Five crab specimens taken by
dredging at 17 fathoms, December, 1965, Laura
Shy, Westminster, California.

This species can be expected to occur throughout the
entire Panamic area in shallow water.

ACKNOWLEDGMENT

For critical reading of the manuscript and helpful
suggestions I am indebted to Dr. James H. McLean.

THE VELIGER

Page 88

LITERATURE CITED

Ciencu, Witu1aM JAMEs & RutH Dixon TurRNER
1951. The genus Epitonium in the Western Atlantic (Part I).
Johnsonia 2 (30) : 249 - 288; plts. 108 - 130. Mus. Comp. Zool.,
Harvard Univ. (28 September 1951)

1952. The genera Epitonium (part II), Depresstscala, Cylind-
riscala, Nystiella and Solutiscala in the Western Atlantic.
Johnsonia 2 (31): 289 - 356; plts. 131 - 177. Mus. Comp. Zool.
Harvard Univ. (23 July 1952)

Dati, WILLIAM HEALEY

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)

DuSuane, HELEN « Twita BrRATCHER
1965. A new Scalina from the Gulf of California. The
Veliger 8 (2): 160-161; plt. 24 (1 October 1965)
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)

Vol. 10; No. 1

Vol. 10; No. 1

NOTES & NEWS

New Checklist in Preparation

BY

JUDY S. TERRY

Department of Geology, Stanford University
Stanford, California 94305

A CHECKLIST oF Recent marine prosobranch gastropods
and pelecypods is being prepared for the U. S. Geological
Survey with special emphasis on range and ecologic data
to be used as a basis for biogeographic and paleoecologic
interpretations of fossil assemblages. Depth, temperature,
and kind of substrate are being recorded for live speci-
mens or dead material which is likely to have lived
recently in the habitat from which it was taken. For
each species the following information will be given:
reference to the original description and a good figure,
catalogue number and repository of the type specimen,
type locality, geographic range, the quality of material on
which the range is based, ecologic data and a list of
synonyms.

Ecologic and range data are expected to come from
the literature, material in existing collections, specimens
taken in dredging operations in connection with other
U.S. Geological Survey, Bureau of Commercial Fisheries
and state Fish and Game projects, and from collections
and observations made at numerous marine stations along
the coast of western North America. Range extensions and
other pertinent information not reported in the literature
are welcomed.

U.M.E. to Meet

It has just been announced that the third European
Malacological Congress will take place in Vienna, Austria
from September 3 through September 6, 1968. All mala-
cologists are invited to attend. Detailed information may
be obtained by writing to Dr. Oliver E. Paget, Museum
of Natural History, Burgring 7, A-1014, Vienna, Austria.
Inquiries should reach Dr. Paget as early as possible, but
not later than August 31, 1967.

THE VELIGER

Page 89

Great Generosity of
SAN DIEGO SHELL CLUB

The San Diego Shell Club a few weeks ago made a
generous donation to The Veliger. This contribution has
been used by the Editor of The Veliger to defray a part of
the cost of the magnificent plates accompanying the paper
on Hysteroconcha and Hecuba in this issue. We wish to
express our appreciation as well as that of the author
to the San Diego Shell Club.

CauirorniIA Matacozoo.tocicaL Society, Inc.
announces:

Backnumbers of

THE VELIGER

and other publications

Volumes 1 through 7: out of print
Volume 8: $14.- Volume 9: $22.-
Supplement to Volume 3: $3.-* plus $-.35 mailing charge

[Opisthobranch Mollusks of California
by Prof. Ernst Marcus]

Supplement to Volume 6: $4.-* plus $-.35 mailing charge
P p §

[Biology of Tegula funebralis (A. ApaMs), edited by
Abbott, Blinks, Phillips and Stohler]

Supplement to Volume 7: $2.-* plus $-.35 mailing charge

[Glossary of A Thousand-and-One Terms used in
Conchology, compiled by Wintrrep H. Arnoxp]

Items marked with * are subject to sales tax in the State
of California; residents of that State please add the
appropriate amount to their remittances.

Prices subject to change without notice.
Send orders with remittance to:

Mrs. Jean M. Cate, Manager, 12719 San Vicente Boule-
vard, Los Angeles, Calif. 90049. Please, make checks
payable to C. M. S., Ine
Shipments of material ordered are generally made within
two wecks after receipt of remittance.
Backnumbers of the current volume will be mailed to new
subscribers, as well as to those who renew late, on the
first working day of the month following receipt of the
remittance. The same policy applies to new members.

Subscription to Volume 10: $12.- domestic; $12.60 in
Canada, Mexico, Central and South America;
$ 12.90 all other foreign countries.

Page 90

BOOKS, PERIODICALS, PAMPHLETS

The molluscan families Speightiidae and Turridae:
an evaluation of the valid taxa both Recent and fossil,
with lists of characteristic species.

A. W. PowELL

Bulletin of the Auckland Institute and Museum, no. 5,
184 pp., 23 pls., 179 text figures. November 1, 1966.

Some 549 generic and subgeneric taxa are listed in this
long-needed work, with citation of original reference
and type species of each and with notes on distribution.
The Turridae and Speightiidae have diverged from Con-
idae and Terebridae, and a graphic frontispiece points up
the relationships of the evolving groups since Mesozoic
time. The introduction, an essay on morphology, is remi-
niscent of the one in a monograph on New Zealand
Turridae published by Powell in 1942 but with added
new material. He now has much more radular evidence
evidence available; in fact, he figures 169 radulae, 126 of
these for the first time. Although there are other impor-
tant anatomical features that are reviewed, the classifi-
cation of the turrids still must be based primarily on
shell characters, for these are all that we have for the
fossil forms. Insofar as possible, Dr. Powell has illustrated
all the type species of valid genera. As in his other papers,
the line drawings and photographs are of excellent qua-
lity. An alphabetical index to generic and specific names
is a useful feature of the work. This monograph will be
immensely helpful to anyone who has to try to identify
turrids — a puzzling group at best — or to deal with the
problem of their classification.

MK

Shell structure of patelloid and bellerophontoid gastro-
pods.

CopELAND MacC.LintTock

Peabody Museum of Natural History, Yale University,
Bull. 22, 132 pp., 40 pls., 128 text figs. February 21, 1967

Conventionally, the molluscan shell is described as being
3-layered, but MacClintock shows that there actually may
be up to 6 distinct layers in some of the limpets. A large
part of the material on which his studies are based — al-
though this cannot be stated in the title — is from the West
Coast. The layering in limpet shells comprises four types
of structure — prismatic, foliated, and lamellar in two
degrees of complexity. Numerous combinations are pos-
sible and MacClintock recognizes 17 for the species

THE VELIGER

Vol. 10; No. 1

“groups” that he has studied. Most of his species groups
seem consonant with groups that modern workers recog-
nize on the basis of soft parts, although there are some
discrepancies. The study of shell structure may well pro-
vide a new tool to aid in the much-needed revision of
West American limpets.

One advantage of a structural analysis is that when
surface detail and coloring are lacking (as in most fossil
limpets), a clue to placement may still be possible. Mac-
Clintock demonstrates this application by a study of the
Paleozoic bellerophontids.

Illustrations in this work are of exceptional quality —
text figures that are superbly drafted and photographic
plates showing thin sections of shells under high magnifi-
cation. The text is well written. MacClintock has made
what could have been a forbidding technical paper into
one that even the uninitiated can understand, and it
should serve as a model for later investigations.

MK

Valid zoological names of the Portland Catalogue

Haratp A. REHDER. Proc. U.S. Nat. Mus., vol. 121
(3579) ; 52 pp.; 1967.

In this very important paper REHDER points out that
the names which are valid must be credited to Lightfoot,
not to Solander or Humphrey. There are 62 available
names out of a total of 111 proposed molluscan names;
these are listed together with many pertinent data. A
systematic list together with many synonyms is appended.
Taxonomists will be grateful for this painstaking piece of
work.

RS

How to clean sea shells

EUGENE BERGERON

Privately printed. Pp. 1-19 (Obtainable from the author,
P.O. Box 1236, Balboa, Canal Zone; price $1.50).

The relative advantages and disadvantages of 5 methods
of removing soft parts from the inside of the shell and
incrustations from the exterior are reviewed, and suggcs-
tions are made as to the best methods to be applied for
various groups of mollusks, notably gastropods, pelecy-
pods and chitons. The cover design on the pamphlet is a
photograph that illustrates the effectiveness of one of the
recommended methods in removing the soft parts intact;
posed alongside the shells from which they had been
extracted, the snail bodies look like replicas.

MK

THE VELIGER is open to original papers pertaining to any problem
concerned with mollusks.

This is meant to make facilities available for publication of original
articles from a wide field of endeavor. Papers dealing with anatomical,
cytological, distributional, ecological, histological, morphological, phys-
iological, taxonomic, etc., aspects of marine, freshwater or terrestrial
mollusks from any region, will be considered. Even topics only indi-
rectly concerned with mollusks may be acceptable. In the unlikely event
that space considerations make limitations necessary, papers dealing
with mollusks from the Pacific region will be given priority. However,
in this case the term “Pacific region” is to be most liberally interpreted.

It is the editorial policy to preserve the individualistic writing style of
the author; therefore any editorial changes in a manuscript will be sub-
mitted to the author for his approval, before going to press.

Short articles containing descriptions of new species or lesser taxa will
be given preferential treatment in the speed of publication provided
that arrangements have been made by the author for depositing the
holotype with a recognized public Museum. Museum numbers of the
type specimens must be included in the manuscript. Type localities
must be defined as accurately as possible, with geographical longitudes
and latitudes added.

Short original papers, not exceeding 500 words, will be published in
the column “NOTES & NEWS”; in this column will also appear notices
of meetings of the American Malacological Union, as well as news items
which are deemed of interest to our subscribers in general. Articles on
“METHODS & TECHNIQUES” will be considered for publication in
another column, provided that the information is complete and tech-
niques and methods are capable of duplication by anyone carefully fol-
lowing the description given. Such articles should be mainly original
and deal with collecting, preparing, maintaining, studying, photo-
graphing, etc., of mollusks or other invertebrates. A third column, en-
titled “INFORMATION DESK,” will contain articles dealing with any
problem pertaining to collecting, identifying, etc., in short, problems
encountered by our readers. In contrast to other contributions, articles
in this column do not necessarily contain new and original materials.
Questions to the editor, which can be answered in this column, are in-
vited. The column “BOOKS, PERIODICALS, PAMPHLETS?” will
attempt to bring reviews of new publications to the attention of our
readers. Also, new timely articles may be listed by title only, if this is
deemed expedient.

Manuscripts should be typed in final form on a high grade white
paper, 8/2” by 11”, double spaced and accompanied by a carbon copy.

EDITORIAL BOARD

Dr. Donatp P. Azsorrt, 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. Jor, W. Hepcretu, Resident Director

Dr. LEo G. HERTLEIN,
Curator of Invertebrate Paleontology

California Academy of Sciences, San Francisco

Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology

Stanford University, Stanford, California

Dr. Victor LoosanorrF, 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. Pite.xa, Professor of Zoology
University of California, Berkeley

Mr. Attyn G. Siru, Associate Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

Dr. Ratpu I. Smiru, Professor of Zoology

Marine Science Laboratory, Oregon State University and Chairman, Department of Zoology

Newport, Oregon

University of California, Berkeley

Dr. Cuartes R. Stasex, Assistant Curator
Depariment of Invertebrate Zoology

California Academy of Sciences, San Francisco

EDITOR-IN-CHIEF

Dr. RupoLtF STOHLER, Research Zoologist
University of California, Berkeley

ASSOCIATE EDITOR

Mrs. Jean M. Cate
Los Angeles, California

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME 10 OcToBER 1, 1967 NUMBER 2
CONTENTS
Notes on Morum dennisoni (REEVE) and Related Species (Gastropoda : Tonnacea).
(Plate 12)
See EE LERE DANCE dca WILTIAM OK. EMERSON) ogee eo es) wey ge OF
On the Identity of Phos laevigatus A. ApaMs, 1851 (Mollusca : Gastropoda).
(Plate 13)
NVALETA NG KGE MERS ONGH sm retmanreyiaisy, ob Zeetiisl iis) We) fush yey sie sh) ai ar eaaew Se se eOO
Studies on East Australian Cowries. (9 Tables)
WIARTAS SCHILDER| & FRANZ ALERED SGHILDER). . © 5 = . . « . “e-. 2 + 103

The Muricidae of Fiji (Mollusca: Gastropoda) Part I — Subfamilics Muricinae
and Tritonaliinae. (Plates 14 and 15; 10 Text figures; 1 Map) plus
an Addendum with Text figure 11

MATERERS OFTIVERS CERNOHORS Ke Maen en ny) las eee on es) oes stoner

A Review of the Living Tectibranch Snails of the genus Volvulclla, with Descriptions
of a New Subgenus and Species from Texas. (21 Text figures)
EIGROUD MV VOB ELARR Yeates ee ccm situate eka tears in. ts ee) ay adits oo OS

Relationship between Penitella penita (Conrap, 1837) and Other Organisms of
the Rocky Shore. (Plate 16)

OLINPVGLVANS sr cueserng ie eirr wor enn tr a nen.) foc awe slat? wo ek AG

Freshwater Mollusks Collected by the United States and Mexican Boundary Surveys.
DD WAGER WW set AyIc OR We ve MeeitS ase uRtn ENE hoe os pe Ge es THD

New Records of Nudibranchia (Gastropoda : Opisthobranchia) from the Central
and West-Central Pacific with a Description of a New Species.
(18 Text figures)

DAVID EI a OUNG wearyermine, Tints. Weft inc, Not mene oMealn aS oyu ets 6 L5G

[Continued on Inside Front Cover]

Distributed free to Members of the California Malacozoological Society Inc.
Subscriptions (by Volume only) payable in advance to Calif. Malacozoological Soc., Inc.
Volume 10: $12.- Domestic; $12.60 in the Americas; $12.90 all other Foreign Countries.

Single copies this issue $4.85. Postage extra.
Send subscriptions to Mrs. Jean M. Cate, Manager, 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

A Re-Interpretation of the Sand-Pipes Described by ADEGOKE. (Plate 17)
JOHN W. EVANS: (.oSi%1c Sygate se) Siu ae ghee on oe eh tien a a

The Reproductive System of the British Turridae (Gastropoda : Toxoglossa) .
(Plate 18; 16 Text figures)

EpMUND Hs SMIrit: 41) 5 Dees) 6. er Mos secs. ta tere Uae
Enzymatic Defenses of Certain Snails Against Metal Ions. (1 Text figure)
LINDSAYGR], WINKLERY& 201s) WONG) Gry 2) 065 rs euler ry

The Radulae of Nine Species of Mitridae (Mollusca : Gastropoda)
(Plate 19; 9 Text figures)

JEAN) Mo G@arte: coe), es cha Tm pepe eraser siete tag ae c etCee
A New Species of Marginella from the Coast of Brazil (1 Text figure)
Jean-Jacques Van Mot « BERNARD TuRscH std. Sirois ahh te, Da! Go te Rea ES
The Rediscovery of Erosaria menkeana (Drsuayes, 1863) (Mollusca : Gastropoda)
(Plate 20)
GRAWFORD NS@ATE (0 ie Se ee ee oe rea ano tc en

Notes on Cephalopods from Northern California.
Ropert R. TAEMADGE 0°. <3. Seas 1S Spicy pieces) eye ee

NOTES & NEWS. .005. 0 or cay), con 1-P Pep yee nen cere eee
Note on the Northward Spreading of Mya arenaria LINNAEus in Alaska.
James B. Gross

BOOKS; PERIODICALS (& 7 PAMPENCE IES wees.) slater tt a a eels

ES
Note: The various taxa above species are indicated by the use of different type styles as
‘shown by the following examples:

ORDER, Suborder, DIVISION, Subdivision, SECTION,
SUPERFAMILY, FAMILY, Subfamily, Genus, (Subgenus).
New Taxa

Vol. 10; No. 2

THE VELIGER

Page 91

Notes on Morum dennisoni (RrEvE) and Related Species

(Gastropoda : Tonnacea )

S. PETER DANCE

The Manchester Museum, The University, Manchester 13, United Kingdom

WILLIAM K. EMERSON

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

(Plate 12)

THE PURPOSES OF THIS PAPER are threefold: (I) to
identify and re-describe the holotype of Oniscia dennisoni
REEVE, a rare and poorly known species from the West
Indies; (II) to provide an annotated list of the names
that have been applied to living representatives of the
genus Morum; (III) to provide an annotated list of the
fossil representatives of the genus Morum reported from
the New World.

Anew name, Morum (Cancellomorum) watsont DANCE
& Emerson, is proposed herein to replace Morum cithara
(Watson, 1881), which is a secondary homonym of
Morum cythara (Broccut, 1814).

I. The Holotype
of Oniscia dennisoni REEVE, 1842

REEVE’s taxon was based on a single shell in the collection
of John Dennison (?d. 1864), a leading British shell
collector (REEvE, 1842, p. 212, pl. 253, figs. 5, 6). No
locality information was provided. In the following year
he published a description of the same shell, giving the
measurements, “Long; 2. lat. 1 1/10 poll.” (Reeve, 1843,
page 91).

In 1858, Commandant Beau listed the species in a
catalogue of the shells of Guadeloupe, West Indics (Brau,
1858) ; subsequently Henrik Kress (1864, p. 35) cited
Beav’s record in a catalogue of West Indian marine shells.
Neither Beau nor Kress indicated the ownership of the
specimens on which their respective records were based.

At the auction of the Dennison collection in 1865 two
specimens were sold on the fourth day and are listed in
the sale-catalogue as follows:

p. 33, Lot 665 “ONISCIA DENNISONI. This, and
the companion lot 753, are exquisitely beautiful shells,
in the finest possible condition, and it is supposed that
no other specimens are at present known.” (Anon.,
1865).

p. 37, Lot 753 “ONISCIA DENNISONI, this and
the other specimen are probably all that are at present
known” (Anon., 1865). Lot 665 was purchased for

17 pounds and Lot 753 for 18 pounds; the names of the
respective purchasers are unknown to us (DaANceE, 1966,
p. 210). In 1873, a specimen was sold at the auction of
the collection formed by Thomas Norris (d. 1852) of
Preston, Lancashire, and is listed in the sale-catalogue
(of the second or “remaining” portion of the collection)
as follows:

p. 17, Lot 757 “ONISCIA DENNISONI, a shell of
extreme rarity” (Anon. 1873).

Neither the purchaser nor the price obtained are known
to us (DANcE, 1966, p. 212). In 1876, one was sold at
the auction of the collection formed by the Dutchman
H.C.R. van Lennep (1820-1879) and is listed in the
sale-catalogue as follows:

p. 12, Lot 161 “ONISCIA DENNISONI a great
rarity (SoweErBy, 1876).

Again neither the purchaser nor the price obtained are
known to us (DANcE, 1966, p. 193).

Page 92

The few other shells of this species recorded in literature
were collected after 1870 and must be dismissed from this
inquiry. '

On the basis of this admittedly scanty information, we
consider that the holotype must be one of three shells; two
of these are now in the British Museum (Natural History)
and the other is preserved in the American Museum of
Natural History. The smaller of the two in the British
Museum once formed part of the collection of Hugh
Cuming (1791-1865), acquired in its entirety by that in-
stitution in 1866 (DANcE, 1966, p. 167). It is labelled
merely “dennisoni” and measures 31.7 mm in length and
23.0mm in width. The larger shell once belonged to Mrs.
de Burgh (d. 1881), a well-known collector whose shells,
incorporated into the collection of V. W. MacAndrew
(d. 1940), were acquired by the British Museum after
MacAndrew’s death (Dance, 1966, p. 213). This shell
too is labelled, by Mrs. de Burgh, merely “dennisonz” and
measures 50.0mm in length and 30.5mm in width. The
shell in the American Museum of Natural History once
belonged to the English collector John Calvert, known for
his magpie collecting activities in the late nineteenth and
early twentieth centuries. In 1939, Calvert’s collection of
shells and other natural objects was bought by an Amer-
ican dealer and offered for sale in New York City
(D’Artiztio, 1950, p. 2). Among specimens from that
collection obtained by Mr. Anthony D’Attilio and gener-
ously presented to the American Museum of Natural
History were the type specimen of Murex centrifuga
Hinps (Emerson, 1960) and a specimen of Morum
dennisoni (D’Artitio, 1955; Emerson, 1967). The latter
is undocumented and measures 53.5mm in height and
30.5 mm in width.

The illustrations in the Conchologia Systematica
(REEvE, 1842, pl. 253, figs. 5, 6) and the measurements
given subsequently by ReEve (1843) are based on the
same shell and are sufficient to eliminate the Cuming

THE VELIGER

Vol. 10; No. 2°

specimen (Brit. Mus. No. 19671) as a contender for
holotype status. The de Burgh and Calvert specimens, on
the other hand, both approximate to the illustrations and
measurements. We suggest that these were the two shells
sold at the auction of Dennison’s collection; that one
came into Norris’ possession (actually to a nephew, Tho-
mas Norris [?d. 1873], who acquired the collection after
his uncle’s death) and one (possibly the same one) into
van Lennep’s; and that de Burgh and Calvert acquired
one each. de Burgh probably acquired her specimen
directly from the Dennison sale, Calvert his specimen from
the van Lennep sale (de Burgh was collecting shells
prior to the Dennison sale while Calvert seems to have
become active rather later in the century). In 1919, J.C.
Melvill (1845-1929) commented on the two Dennison
specimens and said: “Type in the Mus. Brit.? ... The
specimen in the British Museum is, I presume, one of the
two specimens then sold; what I believe to have been the
other was offered by auction at Deventer, Holland, in
July, 1876, at the sale of the Rocters van Lennep Collec-
tion, but I am ignorant of its destination” (MELvILL,
1919, p. 72). We have already shown that the Cuming:
specimen (the only one in the British Museum when.
MELVILL’s paper was published) cannot be the holotype. *

Calvert’s shell (Plate 12, Figure 7) is a well-preserved,
specimen, but it is slightly larger than the measurements.
given by REEvE (1842). It also possesses weaker plications
on the shoulder of the body whorl and has a wider outer:
lip than that of the specimen figured by REEVE (cf.
Figure 6 with 7 of Plate 12). On the other hand, the de

' Tn a footnote to the original description in Conchologia System-
atica REEVE, (1842: 212) says: “we have been informed that
there is another specimen in existence but in very poor condition.”
Although we have not been able to determine the identity of this
“second” specimen, it cannot, in any case, be considered a
paratype because it was not used in the preparation of the
description.

Explanation of Plate 12

Figures 1 to 3: Morum strombiformis (REEVE, 1842)

Figure 1: copy of original figure of holotype of Oniscia strombi-
formis Reeve, after Rerve, 1842, pl. 253, fig. 1; the width of the
parietal callus is inaccurately shown in ReEeEve’s illustration, cf.
Figure 1 with Figure 2. Figure 2: Cuming specimen, holotype, in
British Museum (Natural History), No. 1966723, no locality, photo-
graphs courtesy Dr. R. T. Abbott. Figure 3: immature specimen, in
California Academy of Sciences, Loc. 1900, Cartagena Bay, Colom-
bia, found dead on the beach, F M. Anderson, collector. Figures
approximately x 2.

Figure 4: Morum oniscus (LINNAEUS, 1767), juvenile specimen,
in American Museum of Natural History, No. 112483, from St.

Croix, Virgin Islands, G. Nowell-Usticke, collector; x 2. Note the
more inflated form, shorter spire, wider parietal callus, and coarser
sculpture than in a specimen of about the same size and number of
whorls of Morum strombiformis (cf. Figure 3). baie
Figures 5 to 7: Morum dennisoni (REEVE, 1842).
Figure 5: de Burgh specimen, holotype of Oniscia dennisoni REEVE,
in British Museum (Natural History), No. 19601620, no locality.
Figure 6: copy of original figure of holotype, after REEVE, 1842, pl.
253, figs. 5, 6. Figure 7: Calvert specimen, ex D’Attilio collection,
in American Museum of Natural History, No. 128181, no locality.
Figures approximately x 2.

[Dance « Emerson] Plate 12

Tue VELIGER, Vol. 10, No. 2

Figure 3

Figure 7

Figure 6

Vol. 10; No. 2

Burgh specimen, which is sparingly perforated in its upper
whorls by a boring organism, has the same measurements
as Reeve’s figure, and it appears to be morphologically
more like REEve’s original illustrations (cf. Figure 5 with
6 of Plate 12). Consequently, we consider the de Burgh
shell to be the holotype of Oniscia dennisoni REEVE, 1842.

Morum (Cancellomorum) dennisoni (REEVE, 1842)
(Plate 12, Figures 5 to 7)

Oniscia dennisoni REEVE, 1842: 211, pl. 253, figs. 5, 6,
no locality; Reeve, 1843: 91, no locality; REEveE, 1849,
Oniscia pl. 1, fig. 6, no locality; Beau, 1858: 7, Guade-
loupe, Lesser Antilles; Kress, 1864: 35, repeats BEAU’s
record.

Oniscidia dennisonit REEVE, DA, 1889: 20, 231, Arrow-
smith Bank, Yucatan, Mexico, in 130 fathoms, coral (U.
S.N.M. 93742).

Morum dennisoni REEvE, MELviti, 1919: 72; CLENcH
&« AppoTT, 1943: 5, 6, pl. 4, fig. 5, Sandy Bay, Barbados,
Lesser Antilles, in 75-100 fathoms, a poorly preserved
specimen that may have been ingested by a fish (M.C. Z.
124699) DANcE, 1966: 11935 2105 2125 214, ply 22c¢
(figured specimen ex de Burgh Collection, B.M.N.H.
19601620).

Morum (Cancellomorum) dennisoni REEVE, EMERSON,
1967: 290, pl. 39, figs. 1a, 1b (specimen ex Calvert Collec-
tion, A.M.N.H. 128181).

Type locality: Guadeloupe Island, Lesser Antilles, desig-
nated by Maury (1922 :119) and CLencu & ApporrT
(1943: 6), on the basis of the record of Beau (1858).

Type depository: Holotype, British Museum (Natural
History), No. 19601620, ex de Burgh Collection.

Description of holotype: Shell triangularly ovate, imper-
forate, solid and strong; whorls seven, shouldered, and
formed below the shoulder angle. Spire depressed, slightly
produced. Whorls longitudinally compressed to form
blade-like plications, cancellated with numerous narrow
ribs forming small knobs which are longer and more
produced on the shoulder of each whorl. Interspaces
between nodose ridges with numerous axial threads crossed
by fewer spiral threads. Parictal shield broadly expanded,
thick, minutely pustulate; pustules weakly lincate. Outer
lip reflected, thickened, denticulate along entire inner
margin with irregular ridges. Base color whitish with 4
spiral bands of light brown on the body whorl and with
irregular patches of brown on the spire, on the outer
margin of the outer lip, and on the nodose ridges of the
body whorl. Parietal shield a vivid reddish brown (blood-
red on fresh specimens, fide Reeve, 1849), pustules
whitish. Measurements 50.5mm in length, 30.5mm_ in
width. Operculum not scen.

THE VELIGER

Page 93

New records (specimens not previously reported in the
literature) :

S.S.E. of Freeport, Texas, in 34 fathoms, by shrimp net,
Allen Knight, in 1962; one specimen, dead, faded and
iron stained in collection of Mildred Tate; another dead
specimen taken at the same time.

Off Pensacola, Florida, 30° 02’N, 86°55’ W, 64 fms.,
dredged by Bureau of Commercial Fisheries R/V Oregon,
station 353, May 25, 1951, one dead specimen, mud
bottom. *

Off Tarpon Springs, Florida, 28° 15’N, 84°53’ W, 60
fathoms, dredged by Bureau of Commercial Fisheries R/V
Oregon, station 919, March 11, 1954, one live specimen,
coral bottom, temperature 65° F. ?

Off Strangers Cay, Bahamas, 27° 08’ N, 77° 52’ W, 160
fms., dredged by Bureau of Commercial Fisheries M/V
Silver Bay, station 3474, October 25, 1961, one specimen,
coral bottom. ”

Off Matthew Town, Great Inagua Island, 20°54’ N,
73° 37’ W, 88 fms., dredged by Bureau of Commercial
Fisheries M/V Silver Bay, station 3502, November 5,
1961, two dead specimens, coral bottom. *

On eastern edge of Quita Suefio Bank, off Nicaragua,
14° 05’ N, 81° 21’ W, 100 fms., dredged by Bureau of Com-
mercial Fisheries R/V Oregon, station 4928, June 8, 1964,
two dead specimens, coral bottom. ?

Off Pina, Panama, 9° 19’ N, 80° 25’ W, 75 fms., trawled
by Bureau of Commercial Fisheries R/V Oregon, station
3587, May 29, 1962, one dead specimen, mud bottom. ?

Off Punta San Blas, Panama, 9° 38’N, 78°57’ W, 65
fms., dredged by Bureau of Commercial Fisheries R/V
Oregon, station 5735, October 19, 1965, one live speci-
men, rocky bottom. ?

Off Punta Guaniquilla, Puerto Rico, 18° 03’ N, 67° 27’
W, 165 fms., dredged by Bureau of Commercial Fisheries
R/V Oregon, station 2643, October 5, 1959, one dead
specimen, muddy bottom. ”

West of St. Martin Island, 18° 14’N, 63° 20’W, 76
fms., dredged by Bureau of Commercial Fisheries R/V
Oregon, station 5919, February 25, 1966, one live specimen,
sponge bottom. ”

Off Calibishie, Dominica, 15° 37’ N, 61° 22’ W, 34 fms.,
dredged by Bureau of Commercial Fisheries R/V Oregon,
station 5920, March 3, 1966, two live specimens, sponge
bottom. *

Off Anse Quanery, Dominica, 15° 25’ N, 61° 12’ W, 60
fms., dredged by Bureau of Commercial Fisheries R/V
Oregon, station 5933, March 5, 1966, two live specimens,
rocky bottom. ”

2 Deposited in the collection of the U.S. Bureau of Commercial
Fisheries, Pascagoula, Mississippi, teste Harvey R. Bullis, Jr.

Page 94

Barbados, obtained by the Smithsonian-University of
Iowa Expedition, in 1918; one beach specimen from sta-
tion 411 (U.S.N.M., No. 459829), one fragmental spe-
cimen, station 483, 90-100 fms. (U.S.N.M., No.
459830).

Off east coast of Barbados, 13° 02’N, 59° 34’ W, 123
fms., dredged by Bureau of Commercial Fisheries R/V
Oregon, station 5015, September 20, 1964, one specimen,
rocky bottom. ”

96 miles N of Georgetown, Guiana [British Guiana]
08° 10.9’N, 57°48’ W, 53-60 fms., dredged by R/V
Chain, station 35, cruise 35, R. W. Foster, April 28, 1963;
one specimen with animal (M. C. Z., Harvard University,
No. 262149).

Northeast of Paramaribo, Surinam, trawled by fisher-

men and purchased from them by Mr. T: H. Munyan,
March, 1965; one live specimen in the collection of Mr.
Munyan.
Range: Gulf of Mexico: off Freeport, Texas, Pensacola
and Tarpon Springs, Florida, and Yucatan, Mexico; off
the Bahama Islands: Strangers Cay and Great Inagua
Island; Caribbean Sea: off Puerto Rico, Lesser Antilles,
Nicaragua, and Panama; and off Guiana and Surinam, in
the western Atlantic, Living specimens taken in 34 to 76
fathoms (median of 60 fathoms), on coral, rocky, and
sponge substrates.

II. Annotated List of Names
that have been applied
to Living Representatives of the Genus
Morum Roopine, 1798

Morum (sensu stricto)

Type species: Morum purpureum Rooine, 1798 [= M.
oniscus (LINNAEUS, 1767) ], by M.

Strombus oniscus LINNAEUS, 1767, Syst. Nat., ed. 12:
1210 [West Indies]. Type material presumed lost (fide
Dance, 1967, p. 22).

Cypraea conoidea Scopout, 1786, Delic. Flor. Faun.
Insubr., prt. 2, p. 78, plt. 24, fig. 3 [= M. oniscus (Lin-
NAEUS) fide CLENCH & ABzorTT, 1943, p. 4].

Morum purpureum Roopinc, 1798, Mus. Bolt., p. 53
[= M. oniscus (LinNaEvs) }.

“Oniscia tuberculosa G. B. SowErBy,” 1* of name, 1824,
The Genera of Recent and fossil shells, Oniscia, p. 2
[nomen nudum].

Oniscia tuberculosa “SowERBY” REEVE, 1842, Conch.
system., 2: 211, Oniscia plt. 253, figs. 2 - 4 [eastern Paci-
fic]. Type material presumed lost. Type species of Plesi-
oniscia P. FiscHER, 1884, by M.

THE VELIGER

Vol. 10; No. 2

Oniscia triseriata MENKE, 1830, Syn. method. moll.
gen. ... , Ed. 2, p. 64 [= M. oniscus (Linnagus) fide
CLENCH & ApsorTT, 1943, p. 4].

Oniscia lamarckii Desuayes, 1844, in LaMarck, Hist.
nat. anim. sans vert., Ed. 2, 10: 12 [=M. oniscus (Lin-
NAEuS) ]. Not “Cassidaria (Oniscia, SowERBY) lamar-
kit” [sic] Lesson, 1840, Rev. Zool. Paris 3: 212, from
“Australia,” an unrecognizable taxon according to MEL-
vILL (1919, p. 70). The type of Lesson’s taxon is not in
the collection of the Muséum National d’Histoire Natu-
relle, Paris, teste Dr. Bernard Salvat (zn litt.) .

Morum xanthostoma A. Apams, 1854, Proc. Zool. Soc.
London for 1853: 174 [= M. tuberculosum (REEVE) ].
Lectotype (here selected) in British Museum (Natural
History), No. 1966721, ex H. Cuming collection.

(Herculea) Haney, in H.&A.
ApaMs, 1858

Type species: Oniscia ponderosa HANLEY, 1858, by M.

Oniscia ponderosa Hantey, 1858, Proc. Zool. Soc. Lon-
don, p. 255, plt. 42, figs. 9, 10 [western Pacific]. Lectotype
(here selected) in the British Museum (Natural History),
No. 1966724, ex. H. Cuming collection. We do not agree
with Tryon (1885, p. 282) that Morum exquisita
(ApaMs & REEVE) is the same species; neither does it
resemble M. ponderosa, which is characterized by a
non-papillate parietal callus, sufficiently to be placed in
the same subgenus.

(Cancellomorum) EMERSON &
O xp, 1963

Type species: Morum grande (A. Apams, 1855), by O.D.

Until recently, the cancellate species of Morum were
commonly referred to either “Oniscidia” (SwaAINsoNn,
1840, p. 299) or to “Onimusiro” (Kira, 1955, p. 43; 1959,
p. 51), neither of which is an available name. As WoopRING
(1959, p. 202) has pointed out, “Oniscidia” is based on
an apparent lapsus made by Swainson for Oniscia
Sowersy, 1824, a fact first noted by HERRMANNSEN in
1847. Oniscia, together with Lambidium Linx, 1807,
Ersina Gray, 1847, and Plesioniscia FiscHER, 1884, is a
junior synonym of Morum (s.s.) Ropinc, 1798. The
name “Onimusiro” was first used by Kira (1955) in a
caption for a figure of “Morum (Onimusiro) grande.”
Kira (1955; 1959) did not present a diagnosis nor did he
designate a type species for “Onimusiro.” The Internation-
al Code of Zoological Nomenclature, however, requires
a definite fixation of a type species for genus-group names
proposed after 1930 (Art. 13b), and new names published
after 1930 must be accompanied by a statement purporting
to give characters differentiating the taxon (Art. 13, a, i).

Vol. 10; No. 2

Thus, “Onimusiro” was not properly proposed, and we
have not found a validation of the name in the litera-
ture, unless HaBe (1964, p. 64) inadvertently validated
it as a replacement name when he noted that “Onimusiro”
substitutes Oniscidia SwaINson, which was stated to be
a misspelling of Oniscia Sowersy. If his statement con-
stitutes a validation, Onimusiro would date from Habe,
1964; consequently it would be referable to the synonymy
of Cancellomorum EMERSON & OLD, 1963: 18.

Oniscia cancellata G. B. SowerBy, 1" of name, 1824,
The Genera of Recent and fossil shells, Oniscia figs. 1 - 3
[western Pacific]. Type material presumed lost.

Oniscia strombiformis REEvE, 1842, Conch. system. 2:
210, plt. 253, fig. 1 [West Indies]. Holotype in the British
Museum (Natural History), No. 1966723, ex H. Cuming
collection. Provenance not stated in the original descrip-
tion, but it is cited from “Honduras” by REEvE (1849)
and Me vit (1919, p. 71). CLENcH & AspotTT (1943, p.
5), in a review of the western Atlantic species of the
genus, suggested that the type specimen may have been
obtained by Cuming in the Philippine Islands. As a
result of the present study, a specimen with definite
locality data was obtained on loan through the kind
offices of Dr. Leo G. Hertlein of the California Academy
of Sciences. It was found washed up on the beach at
Cartagena Bay, Colombia, by Dr. F M. Anderson in 1915.
The specimen is worn, but agrees well with the description
of REEveE’s taxon. It has a shorter spire and a less mature
outer lip than the type specimen (compare Figures 1
and 2 with Figure 3 of Plate 12 in the present paper).

Oniscia dennisoni REEVE, 1842, Conch. system. 2: 212,
plt. 253, figs. 5, 6 [western Atlantic]. Holotype in the
British Museum (Natural History), No. 19601620. See
Part I of the present paper.

Oniscia exquisita ADAMS & REEVE, 1850, in A. ADAMs,
The zoology of the voyage of H.M.S. Samarang . . .,
Mollusca p. 35, plt. 5, figs. 3a, 3b [western Pacific]. The
depository of the holotype was not known to MELVILL
(1919, p. 72) and is not known to us.

Oniscia grandis A. ApaMs, 1855, Proc. Zool. Soc. Lon-
don for 1853, p. 185 [western Pacific]. Lectotype in the
British Museum (Natural History), No. 1966726, figured
by Yen (1942, plt. 17, fig. 104).

Oniscia cithara Watson, 1881, Journ. Linn. Soc. Zool.,
15: 266; 1886, Rprt. Sci. Res. Voy. H.M.S. Challenger,
15: 410, plt. 34, fig. 6 [western Pacific]. Holotype in the
British Museum (Natural History), No. 1887.2.0.1262.
The unique type specimen is not fully grown. Not “Onis-
cia cithara. Buccinum Cythara Broccnr’ G. B. Sowersy,
1* of name, 1824, The Genera of Recent and fossil shells,
Oniscia p. 2, fig. 5 [== Buccinum cythara Broccin, 1814,

THE VELIGER

Page 95

Conchiol. foss. Subappennina . .
species of Morum.

Oniscia macandrewi G. B. SowErBy, 3" of name, 1889,
Proc. Zool. Soc. London for 1888: 567, plt. 28, figs. 1, 2
[western Pacific]. Holotype in the Melvill-Tomlin collec-
tion, National Museum of Wales, Cardiff, U. K.

Morum praeclarum MELvILL, 1919, Proc. Malacol. Soc.
London 13: 69, text fig. [? western Pacific]. Holotype in
the Melvill-Tomlin collection, National Museum of Wales,
Cardiff, U. K.

Oniscidia bruuni Powett, 1958, Rec. Auckland Inst.
Mus. 5: 80, plt. 11, fig. 5 [western Pacific]. Holotype in
the Zoological Museum, Copenhagen, Denmark.

Morum (Onimusiro) teramachii Kuropa « Hass, in
T. Haze, Coloured illustr. shells Japan II, 1961, Appen.
p. 15, plt. 20, fig. 3 [western Pacific]. Holotype in the
National Science Museum, Tokyo, Japan. No. 38900.

Morum (Onimusiro) uchiyamai Kuropa & Hase, in
T. Hane, Coloured illust. shells Japan II,, 1961, Appen. p.
16, plt. 20, fig. 5 [western Pacific]. Holotype in personal
collection of Prof. Kawamura, Tokyo, Japan.

Morum (Cancellomorum) matthewsi EMERson, 1967,
The Veliger 9: 290; plt. 39, figs. 2-4 [western Atlantic].
Holotype in the American Museum of Natural History,
No. 129201.

., p. 330], an extinct

Morum (Cancellomorum) watsoni DANCE & EMERSON,

new name for Oniscia cithara Watson, 1881, not Oniscia
cythara (Broccut, 1814). Under the provisions of Articles
59b and 58 (9) of the International Code of Zoological
Nomenclature Watson’s taxon is a secondary homonym
and must be replaced (Art. 60b). Vide supra: Oniscia
cithara Watson, 1881.

“Morum sobrinus A. ADAMS” may be a manuscript
name that is referable to one of the Recent cancellate
species, although we have not found a citation to this
taxon in the primary literature. This name appears in a
recently published popular handbook. It is also on a
label accompanying a specimen of Morum macandrewi
(Sowersy) that was recently received from Japan and is
in the collection of the American Museum. It seems more
likely, however, that “Morum sobrinus” is a lapsus for
Murex sobrinus of ARTHUR ApAMs (1863, p. 370) from
Japan.

(Pulchroniscia) Garrarp, 1961
Type species: Pulchroniscia delecta GARRARD, 1961, by M?

Fixation of the type species of “Pulchroniscia delecta gen.
et sp. nov.” by monotypy is open to question. The formula,
“gen. n., sp. n.,” was not acceptable for fixing generic type
specics after 1930 by the provisions of Art. 68(a) (i) of

Page 96

the International Code of Zoological Nomenclature. This
taxon is tentatively placed in the genus, in the absence of
knowledge of the soft parts.

Pulchroniscia delecta GARRARD, 1961, Journ. Malacol.
Soc. Austral. no. 5, p. 16, plt. 1, figs. 9a, 9b [western
Pacific]. Holotype in the Australian Museum No. 63343.
The description was based on a single specimen, and no
further specimens have been found, teste T: A. Garrard
(in litt., April 1967).

III. Annotated List
of the Fossil Representatives of the
Genus Morum Ropine, 1798,
reported from the New World

Morum (sensu stricto)

Morum oniscus (Linnaeus), Gass, 1881, Journ. Acad.
Nat. Sci. Philadelphia, ser. 2, p. 357, [late] Pliocene, Moin
Hill, Costa Rica. [Geological range: Pliocene to Recent].

Morum floridana [sic] Tucker & Witson, 1933, Bull.
Amer. Paleont. 18 (66): 71; plt. 10, figs. 3-5, “Pliocene,”
Prairie Creek, Florida. The shell of this taxon is stated
to differ from that of M. oniscus (LINNAEUS) in having
a higher spire and by possessing more axial ribs.

Morum tuberculosum “SowrErBy” (REEVE), JORDAN,
1936, Contrib. Dept. Geol. Stanford Univ. 1: 114, Pleisto-
cene, Magdalena Bay, Baja California, Mexico; PALMER
& HeERTLEIN, 1936, Bull. South. Calif. Acad. Sci. 35: 68,
Pleistocene, Oaxaca, Mexico; HERTLEIN & STRONG, 1939,
Proc. Calif. Acad. Sci. 33: 370, Pleistocene, Isla San Sal-
vador, Galapagos Islands, Ecuador. [Geological range:
Pleistocene to Recent].

(Cancellomorum) EMERSON &
Oxp, 1963

Oniscia scotlandica TRECHMANN, 1925, Geol. Mag. 62:
491, plt. 24, figs. 18a, 18b, Scotland Beds, Barbados;
middle Eocene according to W. P. Woodring (in litt., June
16, 1966).

Morum (“Oniscidia”) species Wooprinc, 1959, U.S.
Geol. Surv. Prof. Paper 306-B: 202, 203, plt. 25, figs. 11,
17, mid-Eocene, Gatuncillo formation, Panama Canal
Zone. [Record based on a small fragment].

Morum caracoli (ANDERSON, 1938), Geol. Soc. Amer.
Spec. Papers, no. 16, p. 19, pl. 1, fig. 5, Eocene, Colombia,
as “Athleta (Volutospira) caracolt.” CLARK, in CLARK &
DuruHaM, 1946, Geol. Soc. Amer., Mem. 16: 35, pl. 21.
According to Wooprinc (1959, p. 203), ANDERSON’s
taxon “. . .is an earlicr name for M. corrugatum, if not
also for M. chiraense.”

Morum (Herculea) corrugatum Ciark, in CLARK &
Duruam, 1946, Geol. Soc. Amer., Mem. 16: 34, pl. 21,

THE VELIGER

Vol. 10; No. 2

figs. 5, 21, late Eocene of Colombia (see remarks for M.
caracolt).

Morum (“Oniscidia”’) cf. M. antiquum (Bayan),
Wooprinc, 1959, U.S. Geol. Surv. Prof. Paper 306-B:
203, pl. 25, figs. 12, 13, late Eocene or early Oligocene,
Bohio(?) formation, Gatun Lake area, Panama. Woop-
RING records an incomplete, strongly cancellate, low spired
specimen, which apparently represents an unnamed spe-
cics, and he compares it with BayAN’s taxon, a species
described from Eocene deposits in Italy.

Morum peruvianum Otsson, 1931, Bull. Amer. Paleont.
17 (63): 95, pl. 17, figs. 5, 7, late Eocene or carly Oligo-
cene of Peru. Stated to be related to M. harpulum
(ConraD).

Morum chiraense Ousson, 1931, Bull. Amer. Paleont.
17, no. 63, p. 96, pl. 17, figs. 6, 8, late Eocene or early
Oligocene of Peru (sce remarks for M. caracolt).

Oniscia harpula Conrap, 1847, Proc. Acad. Nat. Sci.
Philadelphia 3: 288; 1848, Journ. Acad. Nat. Sci. Phila-
delphia, ser. 2, 1: 119, pl. 12, fig. 6, [Oligocene], vicinity
of Vicksburg, Mississippi.

Oniscia domingensis SowErBy, 1850, Quart. Journ.
Geol. Sci. London 6, pt. 1, p. 47, pl. 10, fig. 3, “Santo
Domingo;” Pitssry, 1922, Proc. Acad. Nat. Sci. Philadel-
phia 73: 363 [Miocene, Dominican Republic].

Oniscia coxi TREGHMANN, 1935, Geol. Mag. 72 (12):
543, pl. 21, fig. 17, Grand Bay Beds, “L. Miocene = Bur-
digalian(?) == Cercado or Baitoa beds of Haiti,’ Carriacou
Islands, West Indics.

Morum chipolanum “DatL” Maury, 1925, Brazil Serv.
Geol. Mineral. Monogr. 4: 115, 617, pl. 4, fig. 4, early Mio-
cene, [Chipola Formation], Florida; GarpNrr, 1947, U.S.
Geol. Surv. Prof. Paper 142-H: 538, pl. 54, fig. 18, as
“Morum chipolanum (Dati, ms.) GARDNER, 0. sp.,”
Chipola formation, Calhoun County, Florida.

Morum (“Oniscidia’) cf. M. chipolanum Mauoury,
Wooprinc, 1959, U.S. Geol. Surv. Prof. Paper 306-B:
203, carly Miocene, Culebra formation, Gaillard Cut,
Panama Canal Zone. Wooprine records a small, incom-
plete specimen, with strongly developed axial ribs and
axial lamellac, that is reminiscent of M. chipolanum.

Morum chipolanum tampanum MawnsFieLp, 1937,
Florida Dept. Conserv., Geol. Bull. 15: 141, carly Miocene,
‘Tampa limestone, Florida; type specimen figured by DALL,
1915, U.S. Nat. Mus. Bull. 90, pl. 12, fig. 28, as “Morum
domingense SoweRsy.” Not Sowersy, 1850.

Morum harrist Maury, 1925, Brazil Serv. Geol. Miner-
al., Monogr. 4: 115, pl. 4, fig. 14, early Miocene, Rio
Pirabas, Para, Brazil.

Morum (Oniscidia) obricnae OLsson & Petit, 1964,
Bull. Amer. Paleont. 47 (212) : 555, 556, pl. 80, figs. 8, 8a

Vol. 10; No. 2

(not “Pl. 83, figs. 8, 8a”), Pliocene, Caloosahatchee marl,
Fort Denaud, Lee County, Florida.

Morum macgintyi [sic] Maxwe.u SmirH, 1937, Nau-
tilus 51 (2): 67, 68, pl. 6, fig. 12, Pliocene, Clewiston,
Florida; Otsson «& Petit, 1964, Bull. Amer. Paleont. 47
(217): 556, 574, pl. 80, fig. 7 [holotype refigured; cited
from “? Unit A”’].

Tertiary representatives of Morum that are apparently
referable to the subgenus Cancellomorum also are known
from Europe, India, Indonesia, New Zealand, and Japan.

ACKNOWLEDGMENTS

We should like to acknowledge our indebtedness to the
following individuals for courtesies of various kinds: Dr.
R. Tucker Abbott of the Academy of Natural Sciences of
Philadelphia, Pennsylvania; Dr. C. O. van Regteren Altena
of the Rijksmuseum Van Natuurlijke Historie, Leiden;
Dr. FE. M. Bayer of the Institute of Marine Sciences, Uni-
versity of Miami, Florida; Dr. Bernard Salvat, Muséum
National d’Histoire Naturelle, Paris; Dr. Kenneth J. Boss
of the Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts; Mr. & Mrs. Riley
Black of Fort Myers, Florida; Mr. Harvey R. Bullis, Jr.,
of the U.S. Bureau of Commercial Fisheries, Exploratory
Fishing and Gear Research Base, Pascagoula, Mississippi;
Dr. ‘Tadashige Habe of the National Science Museum of
Tokyo; Dr. H. E. Coomans of the Zoological Museum,
Amsterdam; Dr. Leo G. Hertlein of the California Acad-
emy of Sciences, San Francisco; Mr. Thomas L. McGinty
of Boynton Beach, Florida; Mr. Thomas H. Munyan of
Atlantic City, New Jersey; Mr. Joseph H. Peck, Jr. of the
Museum of Paleontology, University of California, Ber-
keley; Mrs. Mildred Tate of Lake Jackson, Texas; Mr.
Norman Tebble and Mr. John FE Peake of the Department
of Zoology, British Museum (Natural History), London;
Mr. Gordon Nowell-Usticke of Christiansted, St. Croix,
U.S. Virgin Islands; Dr. T: E. Pulley of the Houston
Museum of Natural Science, Texas; Dr. Joseph Rosewater
of the U.S. National Museum, Smithsonian Institution,
Washington, D. C.; and Dr. Wendell P. Woodring of the
U.S. Geological Survey, Washington, D. C.

Mr. William E. Old, Jr. and Miss Rae Weinstein of
the American Museum of Natural History kindly provided
technical assistance.

LITERATURE CITED

ApAMS, ARTHUR

1863. | On the specics of Muricinae found in Japan. Proc.
Zool. Soc. London for 1862: 370 - 376

THE VELIGER

Page 97

ANONYMOUS

1865. The celebrated Dennison collection of shells. A catalogue
of this choice, valuable and extensive collection of shells, being
one of the finest ever offered to public competition, etc. London

ANONYMOUS

1873. A catalogue of the valuable collection of shells
formed by the late T. Norris, Esq., of Howick House, Preston.
(In 2 parts: “First Portion,” Lots 1 - 430; “Remaining Por-
tion,” Lots 431 - 908). London
Beau, M.

1858. Catalogue des coquilles recueillies 4 la Guadeloupe et
ses dépendances. Rev. Coloniale for Dec. 1857, 27 pp. [not
seen]

CLencu, WILLIAM JAMEs & RopertT TUCKER ABBOTT

1943. The genera Cypraecassis, Morum, Sconsia, and Dolium

in the Western Atlantic. Johnsonia 1 (9): 8 pp.; 4 plts.
DALL, WILLIAM HEALEY

1889. Reports on the results of dredging, . in the Gulf of
Mexico (1877-78) and in the Caribbean Sea (1879-80), by

the U. S. Coast Survey Steamer “Blake”, . . . Report on the
Mollusca, pt. 2, Gastropoda and Scaphopoda. Bull. Mus.
Comp. Zool. 18 (2) : 1 - 492; plts. 1 - 40
Dance, S. PETER
1966. Shell collecting: an illustrated history. Univ. Calif.

Press, Berkeley & Los Angeles, 344 pp.; 35 plts.
1967. Report on the Linnaean shell collection.
Soc. London 178: 1 - 24; plts. 1-10

Proc. Linn.

D’Attitio, ANTHONY
1950. | Some notes on the great Calvert shell collection. New
York Shell Club notes, no. 2, pp. 2, 3

1955. | Some rarer shells (2). Oniscia dennisoni. New York

Shell Club notes, no. 14, p. 30
EMERSON, WILLIAM KEITH
1960. | Remarks on some eastern Pacific muricid gastropods.
Amer. Mus. Novitates no. 2009: 15 pp.; 7 figs.

1967. A new species of Morum from Brazil, with remarks on
related species (Gastropoda: Tonnacea). The Veliger 9 (3) :
289 - 292; plt. 39; 1 text fig. ( 1 January 1967)

Emerson, WituiaM Kerry & WILLIAM ERwoop OLp, Jr.

1963. Results of the Puritan-American Museum of Natural
History expedition to western Mexico. 19. The Recent mol-
lusks: Gastropoda, Strombacea, Tonnacea and Cymatiacea.
Amer. Mus. Novit. 2153: 38 pp; 28 figs.

Fiscuer, Pau, Henri
1880-1887. Manuel de conchyliologie et de paléontologie con-
chyliologique ou histoire naturelle des mollusques vivants et
fossiles ... Paris (EF Savy) pp. i- xxiv + 1 - 1369; plts.
1-23; 1138 text figures

GarrarD, THomas A.
1961. Mollusca collected by M.V. “Challenge” off the east

coast of Australia. Journ. Malac. Soc. Austral. 5: 2-37;
pits. 1-2 (November 1961)

Page 98

HaseE, TADASHIGE
1964. Shells of the western Pacific in colour. | Osaka, vol. II,
233 pp.; 66 plts.
HANLEy, SYLvANUS Cuar_es T. in Henry & ARTHUR ADAMS
[1854 -]1858. The genera of Recent Mollusca. London, vol. 2,
661 pp.

Kira, TETSUAKI

1955. Coloured illustrations of the shells of Japan. Osaka,
vili+ 203 pp.; illus.
1959. Coloured illustrations of the shells of Japan. Rev. ed.,
Hoikusha, Osaka, 1: 1 - 239; 71 plts.
Kress, Henrik J.
1864. | The West-Indian marine shells with some remarks.

Printed by W. Laub’s widow & Chr. Jorgensen, Nykjobing,
Falster, Denmark, pp. 1 - 137
Maury, CaRLoTTa JOAQUINA

1922. Recent Mollusca of the Gulf of Mexico and Pleistocene
and Pliocene species from the Gulf states, prt. 2. Bull.

Amer. Paleont. 9 (38): 30-172
MELvILL, JAMES Cosmo
1919. Description of Morum praeclarum, sp. nov., with remarks

on the Recent species of the genus.
London 13 (3, 4):
(5, 6): 145

ReEEvE, LovELL Aucustus

Proc. Malac. Soc.
69-72; 1 text fig.; Addendum, Ibid. 13

1842. | Conchologia systematica or complete system of concho-
logy. London, Paris, Berlin and New York, 2.

THE VELIGER

Vol. 10; No. 2

1843. Descriptions of two new species of Oniscia, a genus of
pectinibranchiate mollusks. Proc. Zool. Soc. London for
1842, prt. 10 (114): 91 (January 1843)

1849. Conchologia iconica: illustrations of the shells of mollus-
cous animals. London 5: Monograph of the genus Oniscia,
2 pp.; 1 pit.

Ropinc, PETER FRIEDRICH

1798. Museum Boltenianum ...: pars secunda continens con-
chylia sive testacea univalvia, bivalvia & multivalvia.
Hamburg (Johan Cristi Trappii) pp. i-vii+1- 109

Sowersy, GrorcE BrettinciaM, (3"° of name)

1876. ‘The celebrated collection of shells formed by Mr. H. C.
Roeters van Lennep. A catalogue of this valuable and extensive

collection of shells, etc. London.
SwAINSON, WILLIAM
1840. _A treatise on malacology; or the natural classification of

shells and shell-fish. London, pp. 1-419

Tryon, GeorceE WASHINGTON, Jr.

1885. Manual of conchology.
pp. 268 - 283; plts. 1 - 10

Wooprinc, WENDELL PHILLIPS

Philadelphia, ser. 1, 7

1959. Geology and paleontology of Canal Zone and adjoining
parts of Panama. Geology and descriptions of Tertiary mollusks
(Gastropoda: Trochidae to Turitellidae). U.S.Geol. Surv.
Prof. Paper no. 306 -B: 147 - 239; plts. 24 - 38

Yen, TENG-CHIEN

1942. _A review of Chinese gastropods in the British Museum.
Proc. Malacol. Soc. London 24: 170 - 289; plts. 11 - 28

Vol. 10; No. 2

THE VELIGER

Page 99

On the Identity of Phos laeoigatus A. ADAMS, 1851

(Mollusca : Gastropoda)

WILLIAM K. EMERSON

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

(Plate 13)

INTRODUCTION

Phos laevigatus was briefly described by ARTHUR ADAMS
(1851) on the basis of a single specimen which apparently
was in the collection of Hugh Cuming. The specimen was
stated to have been obtained at the Cape of Good Hope.
As was the case for most of the numerous new taxa that were
proposed by ArTHUR ADAms, no illustrations were pro-
vided and the description was not very diagnostic. G. B.
Sowersy, 2 (1859) subsequently figured a dorsal view
of the type specimen and Tryon (1881) copied this figure.
An illustration of an apertural view of the holotype is pre-
sented for the first time in the present paper (Plate 13,
Figure 1). Both Sowersy and Tryon repeated ADAMS’
locality citation. Additional specimens apparently have not
been referred to Phos lacvigatus since the appearance of
the original description of this taxon more than one
hundred years ago.

As a result of the present study, evidence is presented
to demonstrate that Phos lacvigatus A. ApaMs (1851)
actually occurs in the tropical eastern Pacific Ocean, where
it is currently known as “Phos” chelonia Datu (1917), a
taxon described from off the Galapagos Islands.

DISCUSSION

The problem of the identity of these taxa first came to my
attention when I received from Mrs. Carmen Angermeyer
three specimens of “Phos” that she had collected in the
Galapagos Islands. On examination, I found the two
smaller specimens (Plate 13, Figure 5) to be referable to
Datv’s “Phos” chelonia, but the larger, obviously mature

specimen (Plate 13, Figure 7) appeared to be referable to
“Phos” laevigatus, which had been reported from the Cape
of Good Hope.

Through the kind offices of Mr. Norman Tebble of the
British Museum (Natural History) and Dr. Joseph Rose-
water of the U.S. National Museum, I obtained from
their respective institutions, a photograph of the apertural
view of Apams’ type and the loan of the type specimens of
Datv’s taxon. Our largest Galapagan specimen proved to
be a live-taken example of “Phos” laevigatus, and our
smaller Galapagan specimens were found to be fresh
specimens of “Phos” chelonia, which was determined to
be based on immature specimens that are referable to
“Phos” laevigatus. The availability, however, of ADAMs’
name for the Galapagan species would appear to be
challenged by the Fifty Year Rule. Under Article 23 (b) of
the International Code of Zoological Nomenclature, “A
name that has remained unused as a senior synonym in
the primary literature for more than fifty years is to be
considered a forgotten name (nomen oblitum).” The
code requires that, if such a name is discovered after 1960,
it should be referred to the Commission to be placed on
either the Official Index of Rejected Names or the Official
List of Preserved Names. Under the peculiar circum-
stances of the present case, I prefer to consider ADAMs’
laevigatus an earlier valid name for the nominal species,
Phos chelonia Dati. ADAMS’ taxon would be without
question the first available name for this species, if the
Rule of Priority were followed. '

' At the time of this writing, the “Fifty-Year Rule” (Article 23b)

was under suspension, and this matter was to be considered at
the next meeting of the International Commission on Zoological
Nomenclature (Bull. Zool. Nomencl., vol. 23, p. 260, 1966).

Page 100

SYSTEMATIC TREATMENT

BuccinIDAE

Metaphos Ousson, 1964

Type species: Phos chelonia Dati, 1917 [= Metaphos
laevigatus (A. ApAms, 1851)], by O.D. Recent, Gala-
pagos Islands, in 11 to 40 fathoms.

Other referred species are: Metaphos cocosensis (DALL,
1896), Gulf of California and off the Tres Marias Islands,
Mexico (Parker, 1964), and off Cocos Island in the Gulf
of Panama (Recent); M. pacificus Otsson, 1964, M.
scillus Otsson, 1964, and M. calathus Ousson, 1964,
all from the Esmeraldas formation in northwestern Ecua-
dor (early Pliocene?) . Metaphos cocosensis is questionably
reported by Pitspry & Oxtsson (1941) from the Jama
formation in western Ecuador (Pliocene), and it is re-
corded from Pleistocene deposits on Albemarle Island of
the Galapagos Islands by DaLt & OcHSNER (1928).
Remarks: As pointed out by Otsson (1964), none of the
nearly 50 Phos-like species that are known from the
Cenozoic faunas of the New World is referable to the
genus Phos of Montrort, 1810. The type of Phos (sensu
stricto) is Murex senticosus LinNaEus, 1758, a Recent
species occurring in the Indo-Pacific faunal province.

The shells of the species assigned to Metaphos resemble
those of the genus Cymatophos Pitspry & OLsson (1941),
but they differ notably by the possession of a larger and
more strongly sculptured nucleus. The type species of
Cymatophos is Cymatophos galeris Pitspry & OLsson,
1941, a Pliocene species from the Jama formation in
western Ecuador.

THE VELIGER

Vol. 10; No. 2

Metaphos laevigatus (A. ADAMS, 1851)
(Plate 13)

Phos laevigatus A. Abas, 1851, p. 155, “Hab. Promontorium Bonae
Spei.” Sowersy, 1859, Phos no. 10, pl. 221, fig. 6 [dorsal view of
type], Cape of Good Hope. Tryon, 1881, p. 217, pl. 83, fig. 499
[copy of SowERsy’s figure].

Phos chelonia Datu, 1917, p. 578. “Dredged at the Galapagos
Islands in 40 fathoms.” Stronc « Lowe, 1936, p. 310, pl. 22, fig. 3
[holotype].

Type localities: of laevigatus, not Cape of Good Hope,
here corrected to the Galapagos Islands; of chelonia, U.S.
Fish Commission Station 2813, 01° 21’00” S, 89° 40’ 15”
W, off the Galapagos Islands, in 40 fathoms, coarse sand.

Type depositories: of Jaevigatus, British Museum (Nat-
ural History), holotype no. 1966128, here figured, Plate
13, Figure 1; of chelonia, U.S. National Museum, holo-
type no. 194961, here figured, Plate 13, Figures 3, 4, and
paratypes no. 637989, 12 specimens, including 5 frag-
mental specimens.

Descriptions: As was pointed out by Stronc & Lowe
(1936, p. 308) in their review of the “West American
species of the genus Phos,” the characters of the nuclear
whorls, presence or absence of a columellar keel, and
dentition of the outer lip are specific characters, whereas
the color, number of ribs, and details of sculpture are more
variable characters. The present study serves to corroborate
their findings.

The holotype of Jaevigatus is not well preserved, lacking
the nuclear whorls, and is approximately 38mm in length
(Plate 13, Figure 1). Our largest Galapagan specimen is
a well-preserved, live-taken individual with 54 post-nuc-

Explanation of Plate 13

Metaphos laevigatus (A. Apams)

Figure 1: Holotype of Phos lacvigatus A. ADAMS, in British Museum
(Natural History), No. 1966128; about x 2.

Figure 2: Copy of figure of holotype of Phos lacvigatus A. ADAMS
after SowErRBY, 1859, pl. 221, fig. 6; x 2.

Figures 3, 4: Holotype of Phos chelonia Datu, in U.S. National
Museum, No. 194961; x 2. Figure 3, photographed with incandescent
light, Figure 4 photographed with ultraviolet light.

Figure 5: Juvenile specimen of Metaphos laevigatus (A. ApAMsS),
left Figure photographed with incandescent light, right Figure photo-
graphed with ultraviolet light, dredged off Barrington Island,
Galapagos Islands, in 14 fathoms, together with the specimen illus-
trated below as Figure 7, by the Angermeyers, American Museum

of Natural History No. 135503; x 2.

Figure 6: Juvenile specimen, paratype of Phos chelonia DALt, left
Figure photographed with incandescent light, right Figure photo-
graphed with ultraviolet light, U.S. National Museum No. 637989;
X 4.

pee 7,8: Mature specimens of Metaphos laevigatus (A. ADAMs) ;
x 2. Figure 7, dredged in same haul with the specimen illustrated
above as Figure 5, American Museum of Natural History, No.
135504. Figure 8, dredged in Academy Bay, Indefatigable Island,
in 24.5 fathoms, deposited in the collection of Mrs. DeRoy. Early
whorls of the specimen clearly show the characters of juvenile
paratypic specimens of Phos chelonia DaLt, cf. Figures 6 and 8.

[EMERSON] Plate 13

THE VELIGER, Vol. 10, No. 2

Vol. 10; No. 2

lear and 34 nuclear whorls and the shell measures 36.4 mm
in length (Plate 13, Figure 7). The nuclear whorls of the
latter specimen are spirally sulcate with weak axial riblets.
The first three post-nuclear whorls have strong axial ribs
that are crossed by fine spiral lamellae; the remaining
whorls have fewer but coarser axial ribs and finer and
more numcrous spiral riblets. The base of the body whorl
has low, spiral ridges. In the last two whorls, the axial
ribs terminate below the sutures and the intervening
sutural regions have spiral riblets. The color of the shell is
basically whitish buff, but is tinged with purplish brown
on the body whorl, on the anterior canal and the basal
spiral riblets. Weakly mottled, rusty brown stains occur
in the sutural areas of the early whorls. The aperture is
whitish within with a tinge of purple. Operculum is semi-
crescentic in outline and possesses an apical nucleus.

All the type specimens of chelonia are dcead-collected,
rather bleached specimens, with only a suggestion of the
original color patterns. Photographed under ultraviolet
light (cf. Figures 3, 4, 6), the fluorescent patterns of the
pigments remaining on the surface of the type specimens
are evident, but these patterns are reverscd, 7.c. the white
areas appear as dark areas and the dark areas appear as
white areas. The holotype, the largest specimen in the type
lot, isa complete specimen with 44 post-nuclear whorls and
3% nuclear whorls and measures only 24.5mm in length
(Plate 13, Figures 3, 4). The sculpture of the holotype of
chelonia, which possesses one less whorl than our largest
Galapagan specimen, agrccs in all details with the orna-
mentation of each corresponding whorl of the holotype of
laevigatus and our Galapagan specimens (cf. Figures 3,
4 with Figures 1, 7, 8). The body whorl of chelonia has the
ridge of the siphonal fasciole tinged with purple as does
the holotype of laevigatus and our largest Galapagan
specimen (cf. Figure 3 with Figures 7, 8). The brownish
stains that occur in the sutural regions in the well-pre-
served Galapagan specimens, although not preserved in
the holotype of chelonia, are preserved in some of DALL’s
paratypes.

Non-typological specimens examined, all from off the
Galapagos Islands:

Off Barrington Island [Isla Santa Fe], in 13 to 20 fath-
oms, Angermeyer lIeg., 1 specimen. 27.6mm in length,
Calif. Acad. Sci. no. 39280.

Off Barrington Island, in about 14 fathoms, 3 live-
collected specimens from the gray sand, in the same dredge
haul, Angermceyer lIeg., February 1965, 1 specimen, 36.4

THE VELIGER

Page 101

mm in length (Plate 13, Figure 7), Amer. Mus. Nat. Hist.
no. 135503; 1 specimen, 15.3mm in length (Plate 13,
Figure 5), Amer. Mus. Nat. Hist. no. 135504; 1 specimen,
16.2mm in length, Amer. Mus. Nat. Hist. no. 135502.

Off Jervis Island [Isla Rabida], in about 11 fathoms,
DeRoy leg., April 1966, 2 dead specimens, 26.6 and 14.1
mm in length, respectively, Amer. Mus. Nat. Hist. no.
135501.

Academy Bay, Indefatigable Island [Isla Santa Cruz], in
24.5 fathoms, sand, DeRoy leg., December 1966, 1 live-
collected specimen, 42.6mm in length (Plate 13, Figure
8), DeRoy Collection.

Near the Galapagos Islands, 0° 2130’ N, 89° 37/45”
W, in 20 fathoms, coral sand, U.S. Fish Commission
Steamer Albatross Station 2812, April 1887, 2 dead speci-
mens, 15.6 and 12.9 mm in length, respectively, U. S. Natl.
Mus. no. 206704.

After the text for this manuscript was completed, we
received from Mrs. Jacqueline DeRoy three additional
specimens, including a large, fully mature, extremely
well-preserved specimen that was dredged alive from
Academy Bay, Indefatigable Island (see Plate 13, Figure
8). In this specimen, the largest I have seen, the early
whorls preserve the conchological characters and the color
pattern that are typical for juvenile paratypic specimens ‘
of Da.w’s taxon (cf. Plate 13, Figures 6 and 8). On the
other hand, the later whorls of Mrs. DeRoy’s large speci-
men are typical for the characters of ADAMs’ holotype of
Phos laevigatus (cf. Figures 1, 2 with Figure 8).

On the basis of the material now available for study, I
must conclude that Datw’s chelonia was based on juvenile
specimens which are referable to Apams’ laevigatus.
Therefore, Phos chelonia Dau, 1917, is a junior subjective
synonym of Phos laevigatus A. ADAMS, 1851, which is
in fact a constituent of the Panamic faunal province.

ACKNOWLEDGMENTS

In addition to Mesdames Carmen Angermeyer and
Jacqueline DeRoy of Academy Bay, Isla Santa Cruz,
Galapagos Islands, I am indebted to the following indi-
viduals for courtesies of various kinds: Mr. Norman Tebble
of the British Museum (Natural History); Dr. Joseph
Rosewater of the U.S. National Museum; Dr. Leo George
Hertlein of the California Academy of Sciences; Dr.
Edward C. Wilson of the San Diego Natural History
Muscum; and Mr. William E. Old, Jr. and Miss Rae
Weinstein of the American Museum of Natural History.

Page 102

LITERATURE CITED

ApaMs, ARTHUR
1851. A monograph of Phos, a genus of gasteropodous Mollusca.
Proc. Zool. Soc. London 18 for 1850: 152-155 (28 Feb. 1851)
DALL, WILLIAM HEALEY
1896. Diagnoses of new species of mollusks from the west coast
of America. Proc. U.S. Nat. Mus. 18: 7 - 20
1917. | Summary of the mollusks of the family Alectrionidae of
the west coast of America. Proc. U. S. Nat. Mus. 51: 575 - 579
Dati, WittiaM HeaLey « WASHINGTON HENRY OCHSNER

1928. Tertiary and Pleistocene Mollusca from the Galapagos
Islands. Proc. Calif. Acad. Sci., ser. 4, 17 (4): 89 - 136;
pits. 2-7

LInNAEuS, CAROLUS
1758. Systema naturae per regna tria naturae editio
decima, reformata 1 [Regnum animale]. Stockholm (Laurentii
Salvii) pp. 1 - 824+1- iii
Montrort, PrerrE DENYS DE
1810. Conchyliologie systématique, et classification méthodique
des coquilles; ... 2 [Coquilles univalves, non cloisonnées]
Paris (F Schoell) pp. 1-676+1 - 16; illustr.

THE VELIGER

Vol. 10; No. 2

Ousson, Axe ADOLF

1964. | Neogene mollusks from northwestern Ecuador. Ithaca,

New York (Paleo. Res. Inst.) pp. 1 - 256; plts. 1 - 38 (28 Oct.)
Parker, Rosert H.

1964. | 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 « AxEL ADoLF OLSsoNn

1941. A Pliocene fauna from Western Ecuador.
Nat. Sci. Philadelphia 93: 1 - 79; plts. 1 - 19

Proc. Acad.
(9 Sept. 1941)

SowErBy, GEORGE BRETTINGHAM (2° of name)

1859. Thesaurus conchyliorum, or monographs of genera of
shells. London 3: plt. 19, Monograph of the genus Phos,
Montfort, pp. 89-96; plts. 221, 222.

Srronc, ARCHIBALD McC.ure « HERBERT NELSON LOWE

1936. | West American species of the genus Phos.

Diego Soc. Nat. Hist. 8 (22): 305-320; plt. 22

Tryon, GEorcE WASHINGTON, Jr.

Trans. San

1881. Manual of Conchology, ser. 1, vol. 3, Sub-Family
Photinae.

3: 215 - 220; plts. 83, 84

Vol. 10; No. 2

THE VELIGER

Page 103

Studies on East Australian Cowries

MARIA SCHILDER

AND

FRANZ ALFRED SCHILDER

University of Halle, German Democratic Republic

DuRING THESE LAST YEARS we have examined 8300 cowrie
shells coming from 170 localities in the area between
Cooktown, Kenn Reef (ScHiLper, 1966) and Sydney:
this multitude of cowries with exact locality data allows
a special study on the cowrie fauna of East Australia in
the true sense, as we have excluded the Torres Straits,
southern New South Wales, and the Lord Howe Islands.

The investigated area comprises the areas QUE 47q, 47c, 47b and
the northern half of 47s in ScuitpEr, 1965, page 175.

We are much indebted to the following malacologists
some of whom presented or lent us large series of cowries
collected in restricted localities: W. O. Cernohorsky, Vatu-
koula (Fiji), Miss D. Constantine, Cowes (Victoria),
*C. Coucom, Yeppoon (Queensland), R. J. Griffiths, Port
Macquarie (New South Wales), Mrs. E. D. Harton, Coff’s
Harbour (N.S. W.), *W Hart-Smith, North Sydney (N.
S. W.), *G. Houston, Mackay (Qld.), *W. Krause, Avoca
Beach (N.S. W.), *Mrs. M. Lee, Maroochydore (Qld.),
*Mrs. K. M. Matcott, Dromana (Victoria), *A. Schele-
choff, Brisbane (Qld.), S. R. Shadlow, Mermaid Beach
(Qld.), R. Summers, Petaluma (California), *Mrs. E.
Tautorat, Hayman Island (Qld.), *K. Uetz, Vienna
(Austria). Collectors distinguished with an asterisk (*)
have supplied us with the ten local sets each exceeding
250 shells which will be treated in Table 3.

Total Frequency of Species

Table 1 comprises the following columns:

1 — total number of examined specimens coming from
East Australia;

2 — number of specimens represented in 10 large scts
from restricted localities, cach set including at least
250 shells;

3 = name of genus and spccics (or prospecies) according
to ScHILDER, 1967';

' The generic name Cribraria JoussEAuME, 1884 is preoccupied by

4 — column 1 (East Australia) expressed in per cent of
the total 8300;

5 = column 2 (10 large sets) expressed in per cent of the
total 5936.

In columns indicating the percentage of shells, 7. e. in
the columns 4 and 5 of Table 1 and in all columns of Tables
3 to 6

the figure 0 indicates “less than 4%,”

the sign — indicates total absence, and

figures printed in boldface italics indicate “charac-
teristic species,” 2. e. the most common species the total
number of which just exceeds 50% of the collected speci-
mens.

The six “characteristic species,” 7. e. the most frequent
species the total number of which just exceeds 50% of
all East Australian cowries collected, are identical with
those represented in the total of the 10 largest sets, and
even the percentage of specimens is very similar: their
percentage among the total of 8300 shells (67 species)
and among the 10 sets containing 5936 shells (63 species)
is illustrated in Table 2.

Though this conformity may be explained by the great
influence of the 10 sets over the total (of which the sets
constitute 72% ), the Table 2 shows that the sum of the 10
scts can be considered as a good representative of the East
Australian cowrie fauna.

Frequency in Different Localities

The 10 large sets each exceeding 250 specimens scem to
represent random samples from restricted localities. The

Cribraria GMELIN, 1792 and Persoon, 1794; though the latter genus
is now considered to belong to plants (Myxomycetes), its members
often have been treated as animals (“Mycetozoa”) before 1884 so
that Stranp, 1929 (Acta Univ. Latviensis 20: 8) was correct to
rename the cowrie genus by the name Cribrarula (see International
Rules of Zoological Nomenclature, 1958, art. 2b).

THE VELIGER Vol. 10; No. 2

Page 104
Table 1
1 2 3 4 5
Mauritia
3 2 mappa (LinnagEus, 1758) 0 O
275 230 eglantina (Ductos, 1833) 3 4
242 115 arabica (LiNNAEUuS, 1758) 3 2
4 3 mauritiana (LinnaEus, 1758) 0 0
6 2 scurra (GMELIN, 1791) 0 O
Talparia
20 19 talpa (LinNaAEus, 1758) 0 O
Cypraea
18} il tigris LINNAEUS, 1758 0 0
Lyncina
11 8 argus (LinnaAEus, 1758) 0 O
224 173 lynx (LinnaEus, 1758) 3 8
278 216 vitellus (LinNAEus, 1758) 3. 4
197 164 carneola (LiNNAEUuS, 1758) 2 3
Chelycypraca
1 - testudinaria (LINNAEuS, 1758) 0 -
Luria
Sz isabella (Linnaeus, 1758) ane
Pustularia
1 - mariae SCHILDER, 1927 QO -
20 20 globulus (Linnaeus, 1758) 0 Oo
30 29 margarita (Dittwyn, 1817) Oo 1
81 78 cicercula (LinNAEus, 1758) ify, Sail
35 1 bistrinotata SCHILDER & 0 O
ScHILpER, 1937
Monetaria
385 281 annulus (LinNAEuS, 1758) |
214 200 moneta (LINNAEUS, 1758) Oo 43
Erosaria
242 199 labrolineata (Gasxo1n, 1849) 3. 4
ile) al tomlini ScuiLpErR, 1930 0 O
13 8 helvola (Linnaeus, 1758) 0 O
346 270 caputserpentis (LINNAEUS, 1758) ARO)
7 4 poraria (Linnaeus, 1758) 0 O
568 444 erosa (LINNAEUuS, 1758) ZL 38
2 2 miliaris (GMELIN, 1791) 0 O
12 7 eburnea (Barnes, 1824) 0 O
Staphylaea
173 147 staphylaca (LinnaEus, 1758) 2 8
78 71 limacina (Lamarck, 1810) it tl
Nuclearia
83 73 nucleus (LinNAEuS, 1758) it il
Schilderia :
4 4 moretonensis SCHILDER, 1965 0 O
1 1 quecnslandica SCHILDER, 1966 0 0
Umbilia
5 1 hesitata (IREDALE, 1916) 0 O
Erronea
25 6 walkeri (SowERBY, 1832) 0 O
5 - pyriformis (Gray, 1824) 0 -
10 10 coucomi ScHILDER, 1964 0 O
1117 658 xanthodon (SoweErsy, 1832) 13 11
161 104 subviridis (REEVE, 1835) 2 2

1 4
Erronea (continued)
1 - ovum (GmEtIn, 1791) 0
1157 747 errones (LINNAEUS, 1758) 14
64 29 cylindrica (Born, 1778) 1
251 208 caurica (LinNagEus, 1758) 3
218 137 listert (Gray, 1824) 3
Notadusta
16 4 punctata (Linnaeus, 1771) 0
2 1 martini (ScHEPMAN, 1907) 0
1 1 hartsmitht ScHILDER, 1967 0
Palmadusta
by) &) asellus (LINNAEUS, 1758) 1
195 166 clandestina (Linnaeus, 1767) 2
6 1 saulae (Gasxoin, 1843) 0
2 1 contaminata (SoweErsy, 1832) 0
47 34 humphreysii (Gray, 1825) 1
3 2 ziczac (LinNaEus, 1758) 0
Purpuradusta
727 534 gracilis (Gaskotn, 1849) 9
28 89-24 hammondae (IrEDALE, 1939) 0
29 «17 minoridens (MeEtvitz, 1901) 0
8 3 microdon (Gray, 1828) 0
Blasicrura
By) 27/ quadrimaculata (Gray, 1824) 1
138 128 pallidula (Gasxotn, 1849) 2
25 19 teres (GMELIN, 1791) 0
Bistolida
31 12 kieneri (Hipatco, 1906) (0)
64 45 hirundo (Linnaeus, 1758) 1
5 2 ursellus (GMELIN, 1791) 0
144 90 stolida (Linnaeus, 1758) 2
Ovatipsa
6 4 chinensis (GMELIN, 1791) 0
Cribrarula
329 cribraria (LINNAEUS, 1758) 0
3 1 catholicorum SCHILDER & 0
ScuILper, 1938
- Table 2
most frequent species total
Erronea
errones (LINNAEUuS, 1758) 14%
xanthodon (SoweErsBy, 1832) 13%
Purpuradusta
gracilis (GaskoIn, 1849) 9%
Erosaria
erosa (LINNAEUS, 1758) 7%
Monetaria
annulus (LiNNAEvs, 1758) 5%
Erosaria
caputserpentis (LINNAEUS, 1758) 4%
sum of six species 52%

2 3

Noh OnN !

10 sets

12%
11%

97%
8%
5%

5%
50%

Vol. 10; No. 2

localities have been arranged from North to South and
are designated by the letters Q to Z as follows:
OQ = Holborn Island (leg. Coucom, 1965): 528 shells;
R = Hayman Island (leg. Uetz, 1959 and ‘Tautorat,
1963): 731 shells;
S = Penrith Island (/eg. Houston, 1962):
(see SCHILDER & Houston, 1964) ;
T = Middle Island (leg. Coucom, 1964 and 1965):
399 shells;
U = Humpy Island (leg. Coucom, 1963 and 1964):
664 shells;
V = One Tree Island (leg. Coucom, 1966): 1155 shells
(see CoucoM & ScHILpER, 1967) ;
W = Maroochydore (leg. Lee, 1963) : 453 shells;

472 shells

THE VELIGER

Page 105

X = Mooloolaba (leg. Matcott, 1963): 349 shells;

Y = Moreton Bay (leg. Schelechoff, mostly 1964) : 930

shells;

Z = Avoca Beach (leg. Krause, 1962-1966) and Sydney

area (leg. Hart-Smith, 1966): 255 shells.

The locality Q is situated North off Bowen; R is the northern
border of the Whitsunday Group North of Mackay, S is its southern
border; T and U are in the Keppel Bay off Yeppoon rather close
to the coast (both are reefs bordering the Great Keppel Island in
the North and the South, only 10 km apart), whereas V is 70 km
off Cape Capricom in the Barrier Reef; W and X represent prac-
tically the same beach South-East of Gympie; Y refers chiefly to
the West coast of Stradbroke Island; Z comprises several beaches
North of Sydney from Norah Heads to Port Jackson.

Table 3 shows the similarities and the differences be-

Table 3
species QR SBS RWW Ww ws Woz

Mauritia

eglantina (Ductos, 1833) Sel eee aoe ae ms ll OA =

arabica (LINNAEUS, 1758) (0) a ee eS Re eres elie) Lane) Ui) aes
Lyncina

lynx (Linnaeus, 1758) Ie TD Se Ss Bye TO We

vitellus (LINNAEUS, 1758) 2 LOMA ESR Se oe Ble al i Oe h7

carneola (LINNAEUus, 1758) Cee EURO HON ee ee. = 27/e lene lee be regall
Luria

isabella (LINNAEUS, 1758) 6.73) em ON IE ee Oa
Pustularia

margarita (Dittwyn, 1817) De er fee ame Meet ee esi ice

cicercula (LinNAEuS, 1758) 6 4 - - 0 0 - = = =
Monetaria

annulus (LINNAEUS, 1758) LO SOS ON tie ae eee ao

moneta (LINNAEUvS, 1758) Ol 10} eey7h oA eee eee 2 Bel
Erosaria

labrolineata (GasxoIn, 1849) PD Se Se Ope Aa 84 6 5

caputserpentis (LinNAEUS, 1758) O 1 5 0 - 7 9 7 3 29

erosa (LINNAEUS, 1758) Gh CI) 2 G7 3 8 1276
Staphylaea

staphylaea (LinnaEus, 1758) OF Bie ON 2ae Se a0) —

limacina (Lamarck, 1810) Od Ss sto OU OO GT =
Nuclearia

nucleus (LINNAEUS, 1758) eee ey OF a
Erronea

xanthodon (Sowersy, 1832) S = = @ OM Ali GB 3

subviridis (REEVE, 1835) ee ee cla Fern el baile ee 5)

errones (LinNaAEus, 1758) 2 Wl A 6D We 8B § 2 6B il

caurica (LINNAEuS, 1758) Suede pediment. all) BOY Oi 3D

listeri (Gray, 1824) One Og Okan Ol UO i hod
Palmadusta

clandestina (LiNNAEus, 1767) ORM eee ome OV (Sire G) mon tS) 66

humphreysit (Gray, 1825) Soh oe eae Ole wD eiOe 0
Purpuradusta

gracilis (Gaskotn, 1849) 3 2 O13 7 O Al Sl iG 2
Blasicrura

quadrimaculata (Gray, 1824) ae al - = = = = = =

pallidula (Gasxotn, 1849) 6S) BORK 16 a Pe
Bistolida

stolida (LINNAEus, 1758) 2

Page 106

tween the cowrie faunas collected at these 10 localities
Q to Z. The figures indicate the percentage of species in
each set. To facilitate comparison of the sets, the number
of species has been restricted to 27 so that all rare species
not exceeding 3% in any set have been omitted; but in
the recorded species all data have been indicated even if
the figures do not reach 4%; of course, the sums of these
10 columns never equal 100%. The figures for the “char-
acteristic species” have been printed in boldface italics.

The figures entered in Table 3 seem to be rather hap-
hazard as in most sets different cowries are the most
frequent species. However, there are some cases of sur-
prising regional regularity: thus, for instance, some species
(Mauritia arabica, Monetaria annulus, Erosaria erosa, Er-
ronea caurica) seem to be equally distributed in East
Australia; other species (M. eglantina, Lyncina lynx, Pus-
tularia margarita + cicercula, Nuclearia nucleus, Erronea
errones, Blasicrura quadrimaculata) occur chiefly in the
northern localities, two species (Erronea xanthodon, Pur-
puradusta gracilis) prefer the central area, while other
species (Erosaria caputserpentis, Staphylaea staphylaea, S.
limacina) become more frequent in the South; Lyncina
vitellus and Erosaria labrolineata could be classified as
bicentric. However, these conclusions obtained from 10
populations only seem to be premature.

At any rate, there is some similarity among adjacent
localities; in T and U Erronea xanthodon and E. errones
are prevalent, in W and X Purpuradusta gracilis and E.
xanthodon, and even in R and S E. errones, Mauritia eg-
lantina, Lyncina lynx.

The southernmost area Z with predominance of Erosaria caput-
serpentis (race caputanguis Puruipri), Lyncina vitellus and E. erosa
(race pulchella CozN) seems to be very different from the other
East Australian areas: the cowrie fauna is poor, as the other tropical
species become rare and the South Australian species usually do not
extend as far as the Sydney area; besides, we have observed that an
unusually large percentage of beach shells is not fully grown so

that unfavorable conditions may cause the premature death of the
animals.

Frequency in Split Populations

The significance of the local differences illustrated by
Table 3 may be checked by splitting some sets into two
natural halves:

The set U (Humpy Island) is composed of two collec-
tions made in two subsequent years: 1963 (292 mostly
fresh shells) and 1964 (372 mostly beach shells). The
percentage of specimens collected in 1963 vs. those collec-
ted in 1964 (1963 : 1964) is illustrated in Table 4:

The figures exceeding 3% are very similar in both years,
and the total absence of rare species in one year is only
accidental; the 3 most frequent species are identical.

THE VELIGER

Vol. 10; No. 2

Table 4
U_ [Humpy Island]

Mauritia

eglantina (Ductos, 1833) 3 aol

arabica (LINNAEUS, 1758) 4: 2
Luria

isabella (LINNAEUuS, 1758) O: -
Pustularia

cicercula (LINNAEUS, 1758) =" 3710

bistrinotata SCHILDER & 0: -

ScuiLper, 1937

Erosaria

labrolineata (Gasxotn, 1849) 0: -

erosa (LINNAEUS, 1758) Die
Staphylaea

staphylaea (LinnakEus, 1758) -: 0
Erronea

xanthodon (SoweErsy, 1832) 46 : 45

subviridis (REEVE, 1835) eo id)

errones (LINNAEUS, 1758) 26 : 27

cylindrica (Born, 1778) -: 0

caurica (LInNAEus, 1758) 4: 4
Palmadusta

clandestina (LinNAEuS, 1767) -: 0
Purpuradusta

gracilis (Gasxoin, 1849) 6: 8
Bistolida

hirundo (LinnagEvs, 1758) 0: -

stolida (LINNAEus, 1758) =s eel
Cribrarula

cribraria (LINNAEuS, 1758) -: 0

Both collections practically correspond each to the other.

The figures of collecting set T (Middle Island) in 1964
(295 shells) and in 1965 (104 shells) are illustrated in
Table 5 in the same way:

Table 5

T [Middle Island]

Mauritia

eglantina (Ductos, 1833) 28 ea

arabica (LINNAEuS, 1758) 6: -
Monetaria

annulus (LinNAEuS, 1758) 12
Erosaria

caputserpentis (LINNAEUS, 1758) =a

erosa (LinNaEus, 1758) 2. ctabe
Erronea

xanthodon (SoweErsy, 1832) 34 : 35

subviridis (REEVE, 1835) =e 3

errones (LinNAEuS, 1758) 44 : 32

caurica (LinNAEus, 1758) ue te
Purpuradusta

gracilis (GasKoIn, 1849) 10 : 22
Bistolida

stolida (LinNaAEus, 1758) =e 8

Vol. 10; No. 2 THE VELIGER Page 107

Probably on account of the rather small number of Table 6
shells collected in 1965, there is a less evident correspond-
ence, as even the most common species is different (E7ro- V_ [One Tree Island]
nea errones : E. xanthodon), and Mauritia arabica, rep- We
resented by 6% in 1964, was totally absent in 1965, while eglantina (Ductos, 1833) One
E. subviridis and Bistolida stolida were represented by 3% arabica (Linnazus, 1758) De 7
in 1965 only. Talparia

Set V (One Tree Island) is composed of two parts, both A talpa (Linnagus, 1758) a cael
collected in August 1966; we received 648 fine shells, Eee

: tigris LINNAEUS, 1758 OF:
mostly collected alive, and 507 badly worn beach speci- Lyneina
mens. As emphasized in a previous paper (Coucom «& lynx (Linnagus, 1758) Be G
ScHILpER, 1967) these faunas are very different (see Table vitellus (LinNaEus, 1758) 4: 7
6: the first figure indicates per cent of the fresh specimens, carneola (Linnagus, 1758) Se wl2
the second figure those of the beach shells). . Luria
s isabella (LINNAEUS, 1758) - 3
: te nt : Pustularia

Here not a single species is “characteristic” in both cicercula (LinNaEus, 1758) a)
parts: all species which must be classified as characteristic Monetaria
among the fresh shells are less frequent to rare among the annulus (LinNaEus, 1758) 12: 2
beach shells, and vice versa, and even one characteristic a moneta (Linnazus, 1758) 2; 1

5 5 . LTOSATIA
beach shell (Erosaria labrolineata) is totally absent from IckictPocatin (Cherry LD) Beecrrs
among the fresh specimens. These numerical differences Rival (xessissrae, EG) He ae
point to great ecological differences between the habitat caputserpentis (LINNAEUS) 1758) Bie ZU
of the live taken specimens and that of the dead shells poraria (Linnagus, 1758) 0: 0
washed ashore. erosa (LINNAEUS, 1758) 4: 9
Staphylaea
Discussion of Frequency ee Ee 3 5 0
imacina AMARCK, ee
The total frequency of East Australian cowries indicated Nuclearia
by Table 1 can be regarded as genuine to a certain extent, nucleus (LinNAEvs, 1758) EB
though intensive trawling in deep waters may prove some LBinkarae
: xanthodon (SoweErsy, 1832) - 0
very rare species to be less uncommon. i
aa ree! errones (LINNAEUS, 1758) TS 2:

However, the frequency of species in each set cannot caurica (Linnakus, 1758) 10: 2
be taken for granted as genuine though in some sets it listeri (Gray, 1824) iGe i
may be so, especially in sets composed of beach shells only Notadusta
(U 1964, W, X, Z). In other cases the percentage of punctata (Linnagvs, 1771) ail!)
species may be greatly influenced by the accidental con- Palmadusta
ane . : asellus (LINNAEUS, 1758) OR sae!

itions of the area searched by the collector. For investi-

i Pepe iend (Ss H 1964 clandestina (LinNaEus, 1767) 67 =
gations on Penrith Island (ScH1LpER & Houston, ) frolic (Chuse, 123) aber
and on One ‘Tree Island (Coucom & ScHILDER, 1967) Purpuradusta
have shown that each cowrie species prefers certain tidal gracilis (GasxKotn, 1849) Oe i
zones or even restricted places on the shore, and it is minoridens (MEtviL1, 1901) ORs 0
mere chance from which place the collector gets his set Blasicrura
of living cowries, whereas dead shells washed ashore come i uve Cee ek : z

. y ELIN a

from various zones and thus scem to represent the average ae Pata
fauna in a better way. hirundo (Linnagus, 1758) 4g) 3
These facts confirm the studies on populations of living and fossil stolida (Linnaeus, 1758) O29

cowries published 25 years ago (ScHILDER, 1942). Ovatipsa
: chinensis (GMELIN, 1791) - 0
The Size of the Shells Cribrarula

: ¢ 5 cribraria (LINNAEUS, 1758) is @
The length of cowrie shells is a character influenced by aoe usibaot oi SORE C Om =

individual, sexual, ecological, and geographical factors;
the average length of specimens composing a local popu-

ScuHILper, 1938

Page 108

lation depends on its environment, but the average length
of the sum of all populations living in larger areas evidently
points to geographical characters.

Maria SCHILDER (1967) has computed and tabulated
the average length of the cowrie species and its “usual
variation,” 7. e. the limits of 3 of specimens approaching
the median. These standard figures concerning shells
coming from the whole range of distribution of each
species have been repeated with regard to East Australian
species in column 3 of Table 7.

Table 7 contains the following 5 columns:

1 = number of East Australian populations containing
more than 10 specimens of the species, plus 1 if
the sum of single specimens coming from other
localities exceeds 10;

2 = name of the species (see column 3 of Table 1) ;

3 = lower limit, median, and upper limit of “usual
variation” in length (in mm) of the species in its
whole range (M. Scuitper, 1967).

4 — the same for East Australian shells, expressed by the
mean of the populations the number of which is
indicated in column 1;

= tendency and mathematical significance of differ-

ences in median length of the species generally
(column 3) and that of East Australian shells (col-
umn 4) : small letters indicate that the East Austral-
ian shells are smaller, CAPITALS indicate that they
are larger than the shells coming from everywhere;
the probability of the difference has been classified in
four degrees:

a, A= both medians are within the range of usual
variation of the other column (e. g. Mauritia eglan-
tina): the difference is probably caused by chance;

b, B = one median only is within this range, the other
is outside the other range (e. g. Luria isabella) : the
difference may be real;

c, C = both medians are outside the range of the
other column, but the ranges themselves cross each
other (e.g. Monetaria annulus): the difference
is significant ;

d, D = the ranges of variation indicated in column 3
and 4 exclude each other (e. g. Purpuradusta minor-
idens): the difference is very significant.

Table 7
1 2 3 4 5
Mauritia
7 eglantina 44.51.58 45.49.54 a
4 arabica 37.44.56 55.59.64 C
Tal paria
1 talpa 52.62.72 52.61.71 a

THE VELIGER

Vol. 10; No. 2

1 2 3 4 5
Cypraea
1 tigris 67.81.103 73.89.95 A
Lyncina
1 argus 59.70.83 69.77.84 A
5 lynx 30.35.41 37.41.45 B
6 vitellus 36.44.54 4148.52 <A
4 carneola 24.28.34 32.38.45 C
Luria
4 isabella 20.25.29 24.30.34 B
Pustularia
1 globulus 12.16.19 17.19.20 B
1 margarita 11.12.15 13.17.19 C
2 cicercula 14.16.19 16.17.19 A
2 bistrinotata 13.15.18 15.17.18 <A
Monetaria
10 annulus 16.19.22 21.24.26 C
6 moneta 16.20.25 23.25.27 C
Erosaria
7 labrolineata 12.14.18 18.20.23 C
1 tomlini 12.18.24 19.26.29 C
1 helvola 17.20.23 19.23.27 A
10 caputserpentis 25.29.32 29.31.34 A
13. erosa 25.30.35 29.33.38 <A
1 eburnea 33.38.44 35.37.40 a
Staphylaea
5 staphylaea 11.15.19 17.19.22 B
2 limacina 19.24.28 23.26.30 A
Nuclearia
2 nucleus 16.19.23 22.23.25 B
Erronea
1 walkeri 20.23.28 22.26.28 A
14 xanthodon 23.26.29 23.26.29
3 subviridis 26.30.35 26.30.34 .
17 errones 19.23.28 22.26.29 A
3 cylindrica 24.28.33 24.27.31 a
8 caurica 29.35.41 35.39.42 A
4 listeri 12.15.18 17.19.21 CG
Notadusta
1 punctata 9.11.14 10.13.15 A
Palmadusta
2 asellus 13.15.17 15.17.21 A
7 clandestina 11.14.17 14.16.17 A
2 lutea 12.15.19 15.16.17 A
Purpuradusta
12 gracilis 14.16.19 15.17.19 A
1 hammondae 12.14.15 13.14.16 =
2 minoridens 7. 8.9 10.11.12 D
Blasicrura
2 quadrimaculata 17.20.23 19.20.22 =
4 pallidula 15.16.19 17.20.22 B
2 teres 21.24.29 26.30.34 C
Bistolida
1 kieneri 11.13.18 12.14.16 A
2 hirundo 12.15.17 15.17.19 A
4 stolida 21.24.29 21.22.26 a
Cribrarula
1 cribraria 17.21.26 20.22.25 A

Vol. 10; No. 2

THE VELIGER

The frequency of differences entered in column 5 of
Table 7 is in 43 species as follows (Erronea xanthodon and
E. subviridis must be omitted, as the former is totally
restricted to East Australia and most typical E. subviridis
(s. str.) examined also came from this region only) :

dmace Dia AG BC; | D
ee Pot 20) bee “Ord

In East Australia no cowrie species is really smaller than
in other regions, and 5 species only are possibly smaller;
most species are distinctly larger, 16 species even more or
less significantly larger than usual. This fact may be
caused by the general tendency in many cowrie species to
grow larger in peripheral regions than in the central
areas: (SCHILDER, 1961, GrirriTtHs, 1964), especially in
colder zones (ScHILDER, 1964).

species.

The Size in Different Populations

Table 8 shows the following columns:

1 = the name of the species represented by a sufficient
number of shells in at least 2 localities;

2 = the median length and the usual variation of East
Australian specimens as indicated in column 4 of
Table 7;

Q to Z = the 10 sets enumerated in Table 3; each of
these 10 columns shows the median length of the
shells collected in the set and the letter indicating the
significance of differences as explained above (Table
7, column 5), but here comparing the local sizes with
those observed in East Australia generally (column

Dy,

The mean of negative and positive differences between
the local sizes of various species and those in the whole
region of East Australia shows that several populations

(S, W, Y, Z) are typical; in R, V, X, and especially
in Q (Holborne Island) the shells are rather small, while
they are rather large in T (Middle Island) and very large
in U (Humpy Island). However, in each locality the
cowrie species differ much in relative size: so, for instance,
in U (Humpy Island) Mauritia eglantina and M. arabica
are very large, but Erronea subviridis is rather small, and
in X (Mooloolaba) Erosaria caputserpentis is small, but
Erronea errones is very large.

The divisible sets T, U, and V treated above in Tables
4 to 6 show interesting differences between the parts.
Table 9 enumerates species represented in sufficient num-
bers in both parts indicated in the 3 headlines, and
shows the median and the usual variation in each part,
followed by the letter indicating the significance of the
difference of the first column if compared with the second.

Page 109
Table 9

U [Humpy Island] 1963 1964
Erosaria

erosa (LinNAEus, 1758) 34.36.38 33.34.40 A
Erronea

xanthodon (Sowersy, 1832) 25.28.30 20:28:30

subviridis (REEVE, 1835) 27.29.31 25.28.29 A

errones (LinNaEus, 1758) 26.28.31 2512013

caurica (LiNNAEUus, 1758) 40.43.48 37.41.47 A
Purpuradusta

gracilis (Gasxorn, 1849) 17.18.20 18.19.20 a

T [Middle Island] 1964 1965
Erronea

xanthodon (Sowersy, 1832) 24.26.29 23.25.29 A

errones (LINNAEUS, 1758) 26.28.30 23.26.29 A
Purpuradusta

gracilis (GasKo1n, 1849) 17.18.19 15.17.18 A

V [One Tree Island] living beach
Mauritia

eglantina (Ductos, 1833) 46.53.58 41.46.52 B

arabica (LINNAEuS, 1758) 52.56.62 44.48.54 C
Lyncina

vitellus (L1NNAEus, 1758) 37.39.44 36.42.50 a

carneola (LinnaEus, 1758) 33.35.40 29.34.39 A
Erosaria

caputserpentis (LinNaEusS, 1758) 32.34.36 28.31.34 B

erosa (LiINNaEus, 1758) 34.36.38 26.31.34 C
Erronea

errones (LinNaEus, 1758) 22.25.27 21.24.27 A

caurica (LiNNaEus, 1758) 36.38.41 37.40.43 a

The letters indicating the differences show that in
Humpy Island (U) the length of species is practically
identical in the two subsequent years, while in Middle
Island (T) the fresh specimens of 1964 are altogether
slightly larger than in 1965, and in One Tree Island (V)
the live collected specimens of many species are distinctly
larger than the beach shells, but only 2 species are
possibly smaller (see Coucom « Scuitper, 1967)! One
will observe that this sequence agrees with that of differ-
ences in frequency of species in these sets (see Tables 4

to 6).

Results

Our extensive studies in East Australian cowries show that
both the relative frequency of species and the average
size of shells in this region can be approximately estimated
by the mean of 10 sets of at least 250 shells, the sets being
collected in different parts of the region with partly
different environments. Several widely distributed species
evidently differ in frequency in East Australia from that

Page 110

THE VELIGER

Vol. 10; No. 2

Table 8

Mauritia

eglantina (Ductos, 1833) 45.49.54 | 48 a

arabica (LINNAEUS, 1758) 55.59.64
Lyncina
lynx (LinnaEus, 1758) 37.41.45
vitellus (LinNAEus, 1758) 41.48.52 .
carneola (LiNnNAEUuS, 1758) 32.38.45 | 34 a
Luria
isabella (LINNAEUS, 1758) 24.30.34 | 29 a
Monetaria
annulus (LINNAEUs, 1758) 21.24.26
moneta (LINNAEUS, 1758) 23.25.27
Erosaria
labrolineata (Gasxotn, 1849) 18.20.23 /C
caputserpentis (LINNAEUS, 1758) 29.31.34

erosa (LINNAEuS, 1758) 29.33.38 | 28 b

Staphylaca
staphylaea (LinnaAEus, 1758) 17.19.22
Erronea
xanthodon (Sowersy, 1832) 23.26.29
subviridis (REEVE, 1835) 26.30.34

22.26.29 18 d
24.27.31 28 A
35.39.42 35 c¢

errones (LINNAEus, 1758)
cylindrica (Born, 1778)
caurica (LinNaAEus, 1758)

listert (Gray, 1824) 17.19.21
Palmadusta

clandestina (LINNAEUS, 1767) 14.16.17
Purpuradusta

gracilis (Gasxorn, 1849) 15.17.19 16 a
Blasicrura

pallidula (Gasxotn, 1849) 17.20.22 | 18 a
Bistolida

stolida (LinNAEus, 1758)

21.22.26 | : : ria ‘ :

in other regions, and the size of most species is larger in
East Australia than in the tropics.

These 10 sets, however, differ each from the other both
with regard to relative frequency of species and to average
size of specimens; these differences seem to be partly
caused by mere chance, but in both ways certain regional
tendencies cannot be denied. There are similarities in ad-
jacent localities and great differences between collections
made in the same locality at different times or in different
conditions.

These results seem to be rather poor and necd confirma-
tion by similar collections in other regions. Nevertheless,
we think our provisional investigations worth publishing.

LITERATURE CITED

Coucom, Cepric, FRANz ALFRED SCHILDER & MARIA SCHILDER
1967. | The cowries of Once Tree Island. Keppel Bay Tidings
6 (1): in press

GrirritHs, R. JoHN

1964. Size variation in Cypraca xanthodon. The Cowry

1 (6) : 82 - 83 (18 January 1964)
ScHILper, FRANZ ALFRED

1942. Statistische Untersuchungen an Populationen. Arch.

Molluskenk. 74 (4): 129 - 142 (15 July 1942)

1961. _A statistical study in cowries: the size of Mauritia ara-

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)
Keppel Bay Tidings
(December 1966)

bica (LinNaAEus). The Veliger 4 (1):

1966. Cowries from the Kenn Reef.
4) (@) 3 3
ScHILDER, FraANz ALFRED & MARIA SCHILDER
1964. Latitudinal differences in size of East Australian cowries.
The Cowry 1 (7): 100-101 (1 November 1964)
ScHILDER, FRANZ ALFRED, Marta 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)

Vol. 10; No. 2

THE VELIGER

Page 111

The Muricidae of Fiji

(Mollusca : Gastropoda )

Part I

Subfamilies Muricinae and Tritonaliinae

Plus an Addendum with Text figure 11

BY

WALTER OLIVER CERNOHORSKY

Vatukoula, Fiji Islands

(Plates 14, 15; 10 Text figures; 1 Map)

INDEX or SPECIES
(* denotes synonym or homonym)

TOLahy Seen e te eee ean eT OR

CALORGITO ® Prcrramceem tres tree eee 119 haustellum (Murex) ommuosmssnisennenise 116
aculeatus (Chicoreus) 117 incarnata * 121
CLO I I he eeetente er inflatus *

kurodai *

*
MUBESCENS iat nce t oa ate ene,
rufolirata ~ ....
sandwicensis* .,

120

115
128

anatomica (Homalocantha) ees 128 laciniatus (CHicOreUs) wecsmuoseosenenissmees 119 SAULAE versie 122
brevicula (Favartia) seu 126 Lae Ue chassasicatsttavemnengnrctnmnesaiits 116 QOUTAO™ 5 repentant arrears CL)
brunneus (CHiCOrvEUS) crcwisenrisnnssnnnes 117 longicaudus * 116 Saulin (Ghecoreus) mam enna 122

Cancellata™ vivcessun

Longmant ® rerecsun

sawlii* ......

122

CAPUCINUS (CHICOTEUS) veererersrnannmen 118 martinetana (Homalocantha) ecco. 129 SCOLOPACEUS * eeicsiseee: man Hu)
” carneolus (Chicoreus) ensinisnnsinnanne 119 MaATtINIANUS REEVE” veces L1G SOLO PONE recreate enn ene eT TIS
GUETSIGS =" crepes arene anon microphyllus (Chicoreus) Lennispin aig pena eanvnntn eae bana 115
convolutum (Phyllocoma) rumen 127 miliaris (Vitularia) cscececeemeenn LetragOna (FAvartid) crececsisisussiusneen 126
CHASSIS PIN tga ne ne re 115 MONO ON a eres arene era mene Mere en COPIA CRIES” eraceesremrectrcecerenenm percent 119

crassispira” ...
denudata* .
despectus” .

Pada r
nigrispinosus ... rember
NOdulifera (POt7iCTIA) eeccsosnisorssiseiersinsisicn
ONAZTINA © vireirseissi

brapan (Mittie2)) perenne eee net
trialatus var. *
tribulus (Murex)

115

were, HOY
. 115

CTO penta tee eat ener POGIG) screen crate ra eterna ene ea SG OGOOU OG arenes rir rrtaner te 124
UDI SONA? acre racers src eee ert OCOGOG © comemmremesimmmeeremrmcxoremtrernmsm 1S) tripterus (Pterynotus) .... mas PY
elongatus (Pterynotus) 123 IBOUTLETL iene a ccc smere tees rte ae 120 triqueter (Pterynotus) ceecmmusmnnnmnn I24
ETYthrOstoMa * reece 116 PUM PUN Aiokerinnten Ae inaueniae peice ieaes 128 CDOT CONATA ~ coneremreccicerstemmeroren armen HOLS
fenestratus” ... we 129 raciniatus * ysSvetaa ae see re 119 uncinarius * .. 123
LCL ULO Sangre tee een ee ean 124 TAMOSUS (CHICOTEUS) reccserssstireienssininieren 121 variegata * 124
LONG OSUS sere nit ee ee ee 121 TATIS PIN Gia Preece aan Oe een ee 115 CLO OUS * ccctramrmtecremmmeneniartee nae HOLS
recurvirostris (MUrex) eccocsnsn 129

THIS IS THE SEVENTH PART in the series of faunal mo-
nographs dealing with the marine mollusca of the Fiji
Islands.

In accordance with past procedures only species col-
lected by resident collectors and the author are listed as
verified records. Fiji specimens preserved in Museums
and private collections outside the Fiji Islands shall be

taken into consideration at a later stage, provided that
collecting data arc reliable.

Species recorded from Fiji have a wide-ranging distri-
bution in the Indo-Pacific, living in an area extending
from the Red Sea to the Tuamotu Archipelago. Members
of the two subfamilies discussed below are confined within
tropical to temperate waters between Latitudes 40° N to

Page 112 THE VELIGER Vol. 10; No. 2
So

e

uate

‘ VANUA LEV
BYadua _— - EvU

v Ra Naculo
22 GF
Ie} fe ae pale se
ne)
op e g Noviti
4 © &
Nananu-! RA
'*) Waya yy;
OWoya Loilai vee ng Ellington Wharf
o 5 SRakiraki r Vitilevu Boy
7A Vomo 3 aa
he :
One LAUTOKA
MAMANUCA
G p °Mana
Heo “AMalolo ‘Lodoni S
NADI TAILEVUGa5;° 5
ae VITI LEVU 2
Momi ?: Re
Cl .BAU
18° = os atCoba—P
Noel ea mq SIGATOKA ac WY, BEg- sNasilai
Korotogo \ ~~ “SUVA :Nukulau
Sovi Bay ere
Korolevu
Nadroga reef
if BEOA
Yanuca & S
VATULELE
| S
A?
4
i °
| AY Dravuni
| ab Ces
K - -Bulia
fone
vF ™ KADAVU
19° VUNE ws
ne “a
:
any, Naurul® ~ , GILBERT
© — NEw ho Ire
DR > e PHOENIN's
NEWS yes r4 1s
GUINEA OV othe 4 “
De RIT log v ELLICEIS
~ ee “eo ‘sea curls . tien *
&, ; oO? Ss SAMOA
~ Bye Ss als OD. 5
Of A
New &; :
HEBRIDES QO” Vv 3. IJ It
n NEWS) *o povady* pee
CALEDOMA ie
QUEENSLAND %

PISA aC alee iG,
AUSTRALIA

NEW SOUTH

WALES 2Y Qe FA

zenthno Th NOT To SEALE

178° Longitude East 179°

Scale of Miles

0246810 20 30 40
a _ s —_ ee)

Vol. 10; No. 2

40°S. The area between Southern Japan-Philippines-
Northeast Australia (Longitude 120° E to 150° E) is the
richest in muricine species. About 35 muricine species
have been recorded from Japan and Australia, and there
is an appreciable fall-off in the number of species living
to the West and East of this central Indo-Pacific region:
South Africa 7 species, East Africa 9 and Aden about 10
species. In the easterly direction the number of species
declines to 16 in Fiji, 10 in the Society Islands-Tuansotu
Archipelago region and less than half a dozen species in
the Hawaiian Islands. Other groups of Mollusca, particu-
larly tropical gastropod forms, attain their optimum dis-
tributional densities within this geographical area.

For notes on the geography of the Fiji Islands and
other pertinent data see CERNOHORSKy (1964).

THE ANIMAL

The animal’s foot is moderate in size but powerful, the
mantle is thin and generally of the same colour as the
foot. The siphon is simple, tentacles are moderately short
and stubby, and simple eyes are positioned on the outside
of the thickencd base of the tentacles. Sexes are separate,
and the penis which is situated behind the right tentacle
is thickened at the base but slender, whip-like and curved
at the distal end; the size of the male verge will vary from
14% to 20% of shell-length.

The proboscis is long and slender and the radular
ribbon of specimens examined varied from 4mm to 24mm
in length. The radula, which is of the rhachiglossate type,
contains about 100 to 300 rows of teeth, each row con-
sisting of a multicuspid rhachidian and simple unicuspid
and curved laterals. The first dozen rows of teeth are
greatly worn, and broken cusps may often persist for
another dozen rows. In the subfamily Muricinae the rha-
chidian has either 3 larger main cusps and 2 intermediate
denticles or 3 large cusps only. In the Tritonaliinac the
rhachidian is equipped with 3 to 5 main cusps and with
from 3 to 6 small, vertically incised secondary denticles.

The operculum is generally brown to yellowish-brown
in colour, corncous, and attached to the foot by the scab-
rous part of the interior opercular lining, while a semicircle
portion is smooth and glazed. The nucleus is cither basal,
latero-basal or off-central in Muricinae; in the Tritonali-
inae the nucleus is mostly dextro-lateral or occasionally
latero-basal. As pointed out by Voxes (1964) no grcat
reliance can be placed on the opercular nucleus in the
determination of a subfamilial position, since members of
other muricid subfamilics possess both types of opercula.

Muricidae are active predators and borers; they do not
confine thcir hole-drilling to any particular group of mol-

THE VELIGER

Page 113

lusks but prey on almost all species of gastropods and
pelecypods. The holes bored by muricids into the shells
of other molluscs are perfectly circular on the outer rim,
with the widest diameter at this part, and then slightly
converge towards the interior of the shell. The break-
through point is less perfect and somewhat irregular.
Because of their geometrical outline, these muricid holes
have on occasion been mistaken for man-made holes and
classed as artifacts by archacological excavators.

An accessory boring organ located near the anterior
part of the foot has been reported by Carriker (1961) in
some species of Muricidae. Although some writers have
attributed a dissolving or softening effect to the secretion
of the accessory salivary glands, the actual boring process
appears to be mainly mechanical. The secretion of a
proteolytic enzyme during boring in Muricidae (Man-
souR-BEek, 1934) undoubtedly would assist during the
boring process as a lubricant and absorber of small frag-
ments of calcium carbonate. GraHaM (1941) reports the
secretion of the accessory salivary gland of the muricid
Nucella lapillus (LinNAEus, 1758) to have a pH of 6, and
to have no dissolving effect on calcium carbonate. A
similar secretion was observed by the author in the cyp-
raeid Mauritiana eglantina (Ductos, 1833), which is
an algae- and deposit-feeder, and not a borer. The
muricid species Hexaplex pomum (GMELIN, 1791) does
not have accessory salivary glands (Carriker, 1961),
and the boring process is therefore purely mechanical
in this species. In view of the group’s anatomical features
and feeding habits, the Muricacea are regarded to be
in a more primitive stage of development than the two
other stenoglossan superfamilics Buccinacea and Voluta-
cea (FRETTER & GRAHAM, 1962).

SHELL anp HABITAT

Spccics of the genus Murex s. str. are especially vulnerable
to predators due to the moderate thickness of the shell and
wide open aperture; the prominent sharp spines may have
developed in this group as a compensatory defence feature.
Species of the subgenus Haustellum ScHuUMACHER, 1817,
have retained the wide-open apertures, but the shell has
become thicker and heavier and the spines have become
degraded or obsolete. In Chicoreus Montrort, 1810 shells
are rather solid and spines have changed into shorter
fronds. In the large species of Chicoreus and Phyllonctus
Swainson, 1833, the aperture is still wide open, but the
shells are so large and solid that it would prove difficult
for predators to dislodge the shell or gain access to the
apertural opening when the animal is reposing on the
substratum with the aperture downward. The smalier

Page 114

species of Chicoreus, still retaining their solid shell, have
a reduced apertural size which compensates for their
smaller size and lighter weight. In the Pterynotus-Naquetia
group species have lost the spines or fronds, but have
become smaller and more slender in form, a distinct
advantage in coral-reef environment, and their apertural
opening is reduced. Living species of Muricinae show
therefore a certain correlation of compensating characters
of shell-weight and thickness, size, degree of prominence
of spines and apertural size.

In Fiji specimens examined, the colour and size of the
shell, apertural colouring, size and number of spines or
fronds, sculpture, columellar denticulation were found to
be subject to ecologic and individual variation; a similar
degree of variation has been reported by Wu (1965). The
general form of the shell, number of varices, labial den-
ticulation and length of siphonal canal were less prone
to vary, and surprisingly consistent in many species.

Fijian Muricidae are always associated with coral reefs;
the majority of species flourish in muddy-sand environ-
ment, while species like Chicoreus ramosus (LINNAEUS,
1758) favour a clean sand environment, and Pterynotus
triqueter (Born, 1778) occurs in both clean and muddy
sand localities. Most species inhabit the intertidal zone
while only a few species live in deeper water. Some inter-
tidal species, e. g. Chicoreus carneolus (ROpING, 1798)
can exist under the most adverse conditions and even
flourish. During heavy floods on the Island of Viti Levu,
whole molluscan populations of coastal reefs were
completely destroyed due to the inundation of fresh water
in sheltered bays. Chicoreus carneolus, however, survived
without any ill effects or reduction in numbers.

TAXONOMY

A recent account of the taxonomy of muricids and supra-
specific arrangement of the subfamilies Muricinae and
Tritonaliinae has been published by Vokes (1964), and
no duplication is intended here. In Muricidae as in other
groups of Mollusca, supraspecific taxonomy has received
more attention than taxonomy on the specific level. The
taxonomic works of Ropinc (1798) and Linx (1807),
both binomial catalogues containing numerous generic and
specific names, have been brought to the attention of
malacologists early in this century. Yet in the last 50 years
little attention has been directed to the elucidation of
new taxa contained in these works. Efforts have been
made, commendable ones, to be sure, to elucidate the true
authorship of works published anonymously, while the

THE VELIGER

Vol. 10; No. 2

actual contents have largely remained ignored. Conse-
quently many Lamarckian names adopted by subsequent
writers continue to remain in use, names which should
have been replaced by valid prior ones.

Valid but forgotten names which have been rushed into
print before 1960 are taxonomically acceptable, while
such names introduced into literature after 1960 qualify
as nomina oblita (art. 23b, Code of ICZN, 1964). Many
zoologists feel that insufficient time has been granted to
the elucidation of names introduced in works which have
come to the attention of workers in recent years. In view
of the current dispute about art. 23b of the Code of IZCN
(1964) nomina oblita which have been simply rejected
by the International Commission on Zoological Nomen-
clature, must be suppressed under the plenary powers (in
litt., Assist. ICZN, 10 October 1966). Matters are further
complicated in so far that a name which has been used
in primary literature on the subject in the last 50 years is
taxonomically valid, whereas a name appearing in second-
ary literature is not. No definition on what constitutes
primary or secondary zoological literature has been offered
by the International Commission on Zoological Nomen-
clature, and the interpretation therefore rests with every
individual worker.

The three R6piNG names re-introduced here into muri-
cid literature may after all not qualify as nomina oblita,
as they could have been mentioned in the literature on
the subject in the last 50 years. In view of the taxonomic
problems involved in this particular case, it was thought
advisable to accept the prior names; in one instance the
existing name is a possible homonym, while in another
instance a later but equally new name would have to be
substituted.

ACKNOWLEDGMENTS

I would like to record my thanks to Dr. A. W.B. Powell,
Auckland Institute and Museum, for the free access to the
Institute’s reference library and for other facilities made
available to me while at the Museum. To Drs. H.A.
Rehder and J. Rosewater, Smithsonian Institution, U.S.
National Museum, I am grateful for references and assist-
ance supplied in connection with this paper.

The assistance of Fiji collectors has as always been much
appreciated, and thanks are due in particular to Mr. G.
Broescl, Vatukoula; Mr. & Mrs. R.F Browne, Nausori:
Mrs. J. Hill, Suva; Mrs. A. Jameson, Lautoka; Mr. A.
Jennings, Nadi Airport, and Mr. & Mrs. F Freitag, Suva.

Vol. 10; No. 2

THE VELIGER

Page 115

NEOGASTROPODA
MvuRICACEA

MurRIcIDAE

Muricinae
Murex Linnaeus, 1758

Murex Linnaeus, 1758, Syst. Nat., ed. 10, p. 746 — Type species
by SD (Gray, 1847) Murex tribulus Linnazus, 1758

Characters: Shell moderately light in weight, spire eleva-
ted, whorls convex; spines slender and pointed, somewhat
curved, closed, alternating between short and long. Sculp-
ture consists of 3 varices per whorl, plain or gemmate
axial cords, axial riblets in interstices and axial ribs on
earlier whorls. Aperture wide and open, roundly ovate,
columella smooth, edge of labial lip bluntly denticulate;
siphonal canal generally slender and long, spinose, and
with a narrow central slit on the ventral side. Operculum
with a basal or sinistro-lateral nucleus. Rhachidians of
radula with 3 large main cusps and 2 smaller intermediate
denticles, laterals simple and unicuspid.

Discussion: Vokes (1964) in her revision of the group
cited Murex tribulus LinNAEuS as the type species of
Murex s. str. by subsequent designation of MonTForT
(1810) as Murex pecten MontTFortT. MontTrort (op. cit.,
p. 619) did not cite M. tribulus Linnaeus but rather M.
pecten as the type of Murex, and his illustration on the
opposite page as well as his figure references represent
Aranea triremis Perry, 1811 (=Murex pecten Licut-
Foot, 1786), a similar but distinct species. MontrFort’s
type designation is invalid, since his M. pecten is not syn-
onymous with M. tribulus LinNAgus, nor was M. pecten
one of the originally included nominal species (art. 69(a)
(i), Code of ICZN, 1964).

Murex pecten Montrort, 1810, although an earlier
name for the species M. triremis (Perry, 1811), is a
primary homonym of M. tribulus var. pecten LicHTFooT,
1786 (p. 188, no. 4001 — based on Rumpuius [1705], t.
26, fig. 3). The cited figure is the “Dubbelde Spinnekop”
of Rumpuius, and is the same species as M. triremis
(Perry) and M. tenuispina LaMarck, 1822. Murex pec-
ten LicHTFooT, 1786, should therefore replace PERRY’s
later M. triremis.

(Murex s. str.)

Murex trapa Rovine, 1798
(Plate 14, Figure 1)
1798. Murex trapa Rovinc, Mus. Bolten., p. 145, no. i817 (ref.
Martini, 1777, 3, t.113, figs. 1055, 1056)

1822. Murex rarispina Lamarck, Hist. nat. anim. sans vert., 7:
ref. MarTINI, op. cit., fig. 1056)

1845. Murex martinianus REEvE, Conch. Icon., 3, pl. 18, sp. 72
(ref. Martin1, of. cit., fig. 1056) [non Preirrer, 1840]
1965. Murex trapa Ropinc, Sui-KueE1 Wu, Bull. Inst. Zool. Acad.

Sinica 4: 98, fig. 13 (radula)

Shell: Shell light in weight, base colour creamy-white,
spiral cords on body whorl white or creamy-white, inter-
stices rusty-brown. Teleoconch with 7 whorls, protoconch
with 13 - 2 smooth brown nuclear whorls. Sculptured with
smooth or gemmate spiral cords, 3 varices at every whorl
and close-set axial riblets in interstices. Spines moderately
long, sharp, closed and somewhat recurved; primary spines
number from 3 - 6 on the labial varix and from 4-5 on
the siphonal canal. Aperture wide and oval, labial lip
with 11 - 16 denticles, columella white and smooth, deep
interior of aperture brown. Siphonal canal long, slender
and spinose, slit on ventral side open. Operculum with a
basal nucleus.

Size: 60mm to 110mm.

Type locality: None. The specimens illustrated by Mar-
TINI (1777) were reported from Amboina and Tranque-
bar. “Amboina, Indonesia” is designated as type locality.
Habitat: In muddy sand, in shallow water. Not uncommon
in Southeast Viti Levu (Lodoni jetty), but rare elsewhere.
Distribution: North and South Viti Levu. — From Indo-
nesia to Japan, East Australia and the Fiji Islands.
Discussion: The species is similar to Murex tribulus Lin-
NAEUS, but differs in characters of creamy-white spines
and spiral cords, brown interstices, somewhat fewer and
shorter spines, coarser sculpture on the penultimate and
earlier whorls, and an irregular siphonal canal.

Murex tribulus Linnaeus, 1758
(Plate 14, Figure 2)

1758. Murex tribulus Linnaeus (pars), Syst. Nat., ed. 10, p.
746, no. 444 (first ref. CocumNa, 1616, t. 60, fig. 6)

1817. Murex scolopax Dittwyn, Descr. cat. rec. shells 2: 681
(ref. MartInt, 1777, 3: t. 113, fig. 1052 and CuEemnirz, 1795,
11, t. 189, figs. 1819, 1820)

1822. Murex ternispina Lamarck, Hist. nat. anim. sans vert. 7:
158 (no figures cited)

1822. Murex crassispina Lamarck, Hist. nat. anim. sans vert. 7:
157 (MarrTINI, op. cit., t. 113, figs. 1052-1054 and Cuem-
NITZ, op. cit., t. 189, figs. 1819, 1820)

1845: Murex nigrispinosus REEvE, Conch. Icon., 3, pl. 20, sp. 79

1913: Murex (Tribulus) ternispina LAMARCK, SCHEPMAN, Sib.
Exped. 49d: 341

1913. ? Murex (Tribulus) ternispina var. rufolirata SCHEPMAN,
Siboga Exped. 49 d: 342

1957. Murex crassispira [sic] LaMarck, DoncE, Bull. Amer. Mus.
Nat. Hist. 113 (2) : 82-83

Shell: Uniformly creamy-white to brown in colour. Teleo-
conch with 7 - 8 whorls, protoconch with 2 smooth, glassy-
brown nuclear whorls. Sculptured with clathrate spiral

Page 116

cords, 3 varices per whorl, and close-set and gemmate
axial ridges in interstices. Spines long, sharp, closed
and recurved; primary spines number from 6-8 on the
labial varix and from 4-6 on the siphonal canal. Aper-
ture wide and oval, labial lip with 10 - 15 denticles, some
of which are paired; columella white and smooth, anal
notch obsolete, deep interior of aperture brown. Siphonal
canal long, slender, straight and spinose. Operculum with
a basal nucleus.

Size: 70mm to 120mm.

Type locality: O. Asiae. (Java, Indonesia).

Habitat: in clean and muddy sand and coral rubble sub-
stratum, from 1 - 7 fathoms. Moderately common.
Distribution: Throughout the Fiji Islands. - From the
Red Sea and the Gulf of Oman to Japan, East Australia
and the Fiji Islands.

Discussion: From the 7 figures cited by LinnAEus (1758),
4 figures depict Murex tribulus auctt., and 3 figures repre-
sent M. pecten LightFooT. Lamarck (1822) cited Mar-
TINI’s figures 1052-1054 for his M. crassispina. These
figures represent M. tribulus and were cited for this species
by both R6épinc (1798) and Linx (1807); figure 1052
was cited by Dittwyn (1817) for M. scolopax.

Murex nigrispinosus Reeve, a variant of M. tribulus
with black-tipped spines, appears to be restricted to the
Western Pacific and does not occur in Fiji. Neither does
M. pecten Licutroot, which differs from M. tribulus in
having more numerous and geometrically arranged spines;
the spines on the siphonal canal are wider apart in M,
tribulus, having a spacing of ca. 4.5mm to 6.5 mm, where-
as in M. pecten the width between the spines is only 31mm
to 4mm.

The length of spines in Murex tribulus is variable: the
longest spine, which is generally situated on the dorsal
varix on the body whorl, measured from 12mm to 50mm
in specimens examined.

(Haustellum) Sct UMACHER: 1817

Haustellum ScHUMACHER, 1817, Essai Nouv. Syst., p. 213 — Type
species by T Murex haustellum Linnazus, 1758

Characters: Shell solid and globular, spire elevated, whorls
convex and with a presutural keel, spines absent or few
and thornlike. Whorls number 8, varices 3, early whorls
sculptured with axial riblets and spiral striae, later whorls
striate and nodulose. Aperture wide and oval, elevated,
labial lip obsoletely denticulate, labrum plicate, columella
smooth or plicate, anal notch prominent and formed like
an inverted (2. Siphonal canal slender and long, slit on
ventral surface open. Operculum with an off-central
nucleus.

THE VELIGER

Vol. 10; No. 2

Murex (Haustellum) haustellum Linnakzus, 1758
(Plate 14, Figure 3)

1758. Murex haustellum Linnagus, Syst. Nat., ed. 10, p. 746, no. _
443 (first ref. BuoNANNI, 1684, 3, t. 268)

1798. ? Murex scolopaceus Ropinc, Mus. Bolten., p. 144, no.
1813 (ref. FAVANNE, 1780, t. 38, fig. B2)
1811.. Aranea denudata Perry, Conchology, pl. 45, fig. 1 (non
Triplex denudata Perry, 1811, pl. 7 = Chicoreus sp.)
1817. Haustellum laeve ScHuMAcHER, Essai Nouv. Syst., p. 213
1840. Murex erythrostoma Swainson, Treat. Malac., p. 296 (non
M. erythrostomus Swainson, 1831)

1891. Murex haustellum var. longicaudus BAKER, Proc. Acad.
Nat. Sci. Philadelphia, p. 56 (non Woop, 1828)

1964. Murex (Haustellum) kurodai Sui1kama, Venus: Japan.
Journ. Malac., 23 (1): 35, pl. 3, figs. 1, 2

Shell: Shell solid and globular, creamy-white to fawn in
colour, variously maculated with brown. Teleoconch of
8 convex whorls, protoconch missing in specimens exam-
ined. Early whorls sculptured with axial ribs and spiral
striae, later whorls nodulose, nodules most prominent on
presutural keel ; varices prominent, numbering 3 per whorl,
spines absent or few and thorn-like. Aperture wide and
oval, elevated, labial lip with small denticles, labrum with
short plicae. Columella creamy-white to orange, smooth
or plicate; interior of aperture white, anal notch distinct
and moderately deep. Siphonal canal long, thickened at
base but tapering to distal end, slit on ventral surface
narrow.

Size: 80mm to 125mm.

Type locality: O. Asiae.

Habitat: On sand and coral-rubble substratum, in 2 to
3 fathoms. Rare.

Distribution: North and South Viti Levu and Yasawa
group. — From the Red Sea to Japan, East Australia
and the Fiji Islands.

Chicoreus MontTForT, 1810

Chicoreus Montrorr, 1810, Conchyl. Syst., 2: 611 — Type species —
by OD Murex ramosus LINNAEUS, 1758

Characters: Shell heavy and solid, spire elevated, whorls
convex; spines frondose and prominent, closed or open,
sometimes recurved. Sculptured with spiral striae, axial
riblets and nodules, 3 varices per whorl and nodes between
varices. Aperture wide and oval in the type-species, but
less large in smaller species. Labial lip dentate, columella
smooth or occasionally denticulate, calloused, anal notch
prominent. Siphonal canal broad, only moderately long,
slit on ventral surface open.

Operculum with a sub-basal or basal nucleus. Radula
similar to that of Murex s.str. but the base of the rhachidi-
ans is gencrally less broad and somewhat higher than in

Vol. 10; No. 2

Murex; the central main cusp is longer than the flanking
main cusps, the 2 intermediate denticles are either moder-
ately large or small.

Discussion: For remarks on the type-species see under
Chicoreus ramosus (LINNAEUS).

Chicoreus aculeatus (LAMARcK, 1822)
(Plate 14, Figure 5; Text figure 1)

1822. Murex aculeatus Lamarck, Hist. Nat. anim. sans vert., 7:
163 (no figures cited; non Aranea aculeata Perry, 1811 =
Murex s. str.)

Shell: Shell light in weight, reddish-brown in colour,
varices and siphonal canal occasionally tinged with laven-
der. Teleoconch of 6 to 7 whorls, protoconch with 14 to 2
smooth and mammillate nuclear whorls. Sculptured with
strong and weak finely beaded spiral ridges, 3 varices per
whorl and nodes between varices. The frondose spines are
very short or obsolete on the first three mature whorls,
but moderately produced on later whorls; the spines num-
ber from 5 to 8 on the labial varix and from 2 to 4 on
the siphonal canal. Aperture small and oviform, rose-
violet or lavender in colour, deep interior bluish; labial
lip with 10 to 15 denticles, columella smooth, anal notch
distinct. Siphonal canal irregular and moderately pro-
duced, recurved; slit on ventral side open.

| 100 p j

Figure 1

Half-Row of Radular Teeth of Chicoreus aculeatus (LAMARCK)
Fiji Islands

Radula: The radula is of the Chicoreus type, with 3
main cusps and 2 intermediate denticles on the rhachi-
dian; rhachidians are light brown in colour, laterals
translucent white. Length of ribbon 4.6mm, width 0.14
millimeters in a shell 25.0mm in length; 164 rows of
teeth (-++ 26 nascentes) were counted in this example
(Text figure 1).

Size: 22mm to 38mm.

Type locality: None. (“L’Océan Indien,” Kiener, 1843).
Habitat: On sand and coral-rubble substratum in 10 to
15 fathoms. Rare.

THE VELIGER

Page 117

Distribution: Mamanuca group, Fiji Islands.) — From
the Philippines to Japan and the Fiji Islands.
Discussion: Murex aculeatus LAMARCK is not a secondary
homonym of Aranca aculcata Perry, 1811 (pl. 46, fig. 2)
which is a Bolinus species of the genus Murex s. str.

Chicoreus brunneus (Linx, 1807)

(Plate 41, Figure 6; Text figure 2)

1807. Purpura brunnea Linx (pars), Beschr. Nat.-“Samml. Univ.
Rost., 3te Abth., p. 121 (ref. Martini, 3, t. 105, figs. 990,
991 only)

1811. Triplex rubicunda Perry, Conchology, pl. 6, fig. 4 (non
Purpura rubicunda Rovinc, 1798; nec Murex rubicunda
Perry, 1811, pl. 54)

1822. Murex adustus LaMarck, Hist. nat. anim. sans vert., 7:
161 (ref. MartINI, of. cit., figs. 990, 991)

1854. Murex despectus A. ApaMs, Proc. Zool. Soc. London for
1853: 71

Shell: Shell solid and heavy, generally blackish-brown in
colour, blackish cords appearing on a lighter background;
spire elevated, whorls convex. Teleoconch consisting of 6
whorls, protoconch of 2 nuclear whorls which are calcified
in adult specimens; juvenile shells have a protoconch of
2 smooth, light-brown nuclear whorls. Sculptured with
strong spiral cords and intermediate spiral ridges, 3 varices
per whorl (4 on early whorls) and a prominent node

| 100 p |

Figure 2

Chicoreus brunncus (LINK)
a. Half-Row of Radular Teeth b. Penis
Fiji Islands

c. Operculum

Page 118

between varices. The frondose spines are thick, heavily
foliated and close-set, central labial fronds generally re-
curved. Aperture moderate in size, roundly oviform, rosy-
red at edges, deep interior white; labial lip elevated,
omamented with 13 to 23 irregular denticles, columella
smooth, anal notch distinct. Siphonal canal broad and
short, slit on ventral side open.

Radula: The radula (Text figure 2) is of the Chicoreus
type with 3 main cusps and 2 intermediate denticles on
rhachidian. The radular ribbon is 7.1mm long and 0.24
millimeters wide in a shell 42.0mm in length; the first
dozen rows of teeth are badly worn, cusps on rhachidians
worn down to the base. Fully-formed rows of teeth num-
ber 187 (+ 26 nascentes). Operculum with a basal
nucleus.

Size: 25mm to 85mm.

Type locality: None. Martini (1777) mentioned Banda,
Amboina and Taekang Besi as localities for the species.
“Banda Island, Indonesia” is designated as type locality.
Habitat: Under coral rocks, on clean sand or coral sub-
stratum, in shallow water. Moderately common.
Distribution: Throughout the Fiji Islands. — From East
Africa to Japan, Australia and Samoa.

Discussion: Linx’s Purpura brunnea is a composite spe-
cies, based on Martinr’s figures 990, 991, 993, and 994.
The first 2 figures represent P brunnea Linx which is the
Murex adustus of Lamarck and authors. The latter 2
figures, however, depict M. capucinus Ropinc and La-
MARCK. TOMLIN & WINCKWoRTH (1936) arrived at the
same conclusion in their analysis of Linx’s species. The
type figures are therefore here restricted to Martin1,
1777, 3, t. 105, figs. 990 and 991.

Corton (1956) included the Queensland species Murex
australiensis A. ApAMs, 1853 and the New Caledonian
M. huttoniae WricuT, 1878 in the synonymy of M. adustus
Lamarck, E. A. SmirH (1897) compared the holotype of
M. penchinati Crosse, 1861 in the British Museum (Nat-
ural History) with New Caledonian specimens of ©.
huttoniae WricutT, and found them to be identical in
every respect.

THE VELIGER

Vol. 10; No. 2

Chicoreus capucinus (R6pinc, 1798) -
(Plate 14, Figure 7)

1791. Murex ramosus var. y GMELIN (pars), Syst. Nat., ed. 13,
p. 3528, no. 13 (non Linnagus, 1758) ;
1798. Purpura capucina Rovinc, Mus. Bolten., p. 143, no. 1797
(ref. Tour d’Auverg., [FavaNNE, 1784], 1073, and Martini,

1777, 3, t. 105, fig. 994)
1807. Purpura brunnea Linx (pars), Beschr. Nat.-Sammil. Univ.
Rost., p. 121 (ref. Martini, of. cit., figs. 993, 994 only)
1822. Murex capucinus Lamarck, Hist. nat. anim. sans vert. 7:
164 (ref. CHEMNiTz, 1795, 11, t, 192, figs. 1849, 1850 — spec.
juv., fide LAMarcK)

1914. Murex permaestus HEDLEY, Proc. Linn. Soc. New Sth.
Wales, 39: 745 (nom. nov. pro M. capucinus auctt.)

Shell: Shell heavy and solid, uniformly blackish-brown in
colour, whorls convex. Teleoconch consists of 6 whorls,
protoconch of 14 nuclear whorls. Sculptured with prom-
inent spiral cords, intermediate lirae, axial riblets and
striae; 3 varices per whorl and 2 axial nodes between
varices; varices bluntly foliated. Aperture oviform, tinged
with brown, labial lip with 14 to 17 prominent denticles;
columella smooth, anal notch distinct. Siphonal canal
broad, moderately short and slightly recurved, slit on
ventral side open. Operculum with a basal nucleus.

Size: 45 mm to 60mm.

Type locality: None. (‘““East Indies” fide Martini, 1777).
Habitat: Under coral rocks on muddy sand substratum,
in shallow water. Rare.

Distribution: North Viti Levu. — From the Philippine
Islands to North Australia and the Fiji Islands.
Discussion: The Martini (1777) figures 993 and 994
represent dorsal views of a dark brown shell resembling
Murex capucinus of authors. The figured specimens were
said to come from the East Indies and Martini further
remarks that the specimen depicted in fig. 994 has slightly
more frondose varices than the specimen illustrated in fig.
993. PFEIFFER (1840) assigns the figured species to M.
capucinus Lamarck. R6pinc (1798) clearly separated
the frondose species of Chicorcus by placing them in the
section “Frondosae — Acstige” while specics of Naquetia

Explanation of Plate 14

Figure 1: Murex trapa ROpiNc. x 0.8

Figure 2: Murex tribulus LINNAEUS. x 0.7

Figure 3: Murex (Haustellum) haustellum LinNaEus. x 0.6
Figure 4: Phyllocoma convolutum (BRODERIP). x 2.6

Figure 5: Chicoreus aculeatus (LLAMARCK). x 1.4

Figure 6: Chicoreus brunneus (Linx). x 0.6

Figure 7: Chicoreus capucinus (ROpInc). x06

Figure 8: Chicoreus carneolus (RopiNc). x 0.6

Figure 9: Chicoreus laciniatus (SowERBY). =X 1.3

Figure 9a: Chicoreus laciniatus (SowERBY). = X_:1.3,

Figure 9b: Chicorcus laciniatus (SowzERBY). x 13

Figure 10: Chicoreus microphyllus (LAMARCK). x 1.4 & 1.2

Figure 11: Chicoreus ramosus (LINNAEUS). x 0.2

THE VELIGER, Vol. 10, No. 2 [CeERNOHORSKY] Plate 14

Figure 1 Figure 2 Figure 3 Figure 4

Figure 9 a Figure 9 b Figure 10

photographs by W. O. CernoHorsky

Vol. 10; No. 2

Page 119

THE VELIGER

including Purpura capucina were placed under “Optusae
— Abgestumpfte.” R6pinc (op. cit.) appended a rather
important diagnostic sentence after P. capucina “Ketten-
hérner mit stumpfen Nahten,” a feature which identifies
P. capucina in conjunction with the cited figures and Mar-
TINI's text.

Recent writers assign Murex capucinus to Naquetia
JoussEaumE, 1880, in the genus Pterynotus Swainson.
H.«A.Apams (1853) placed the species in Chicoreus
Montrort, and this placing appears to be quite appro-
priate. The general features of the shell of the species
resemble Chicoreus more than Naquetia or Pterynotus;
varices are neither wing-like nor compressed, and blunt
varices are evident in other Chicoreus species; the anal
notch is distinct in Chicoreus but shallow or obsolete in
Pterynotus or Naquetia.

Chicoreus carneolus (Rop1nc, 1798)
(Plate 14, Figure 8)

1791. Murex ramosus GMELIN (pars), Syst. Nat., ed. 13, p. 3528,
no. 13 (non Linnaeus, 1758)

1798. Purpura carneola Répvinc, Mus. Bolten., p. 142, no. 1792
(ref. Martini, 1777, 3, t. 106, figs. 995, 996)

1807. Purpura elongata Linx, Beschr. Nat.-Samml. Univ. Rost.,
p. 121 (ref. MartInI, op. cit., figs. 995, 996, 997) [non Murex
elongatus LicutFoot, 1786 = Pterynotus sp.]

1811. Triplex abortiva Perry, Conchology, pl. 6, fig. 5

1841. Murex torrefactus SowerBy, Conch. Illust., Murex, figs.
110, 111

1841. Murex torrefactus SowErBy, Proc. Zool Soc. London for
1840; pt. 8: 141

1966. Murex (Chicoreus) torrefactus SowERBY, CERNOHORSKY,
The Veliger 8 (4): 231-233, 6 text figs. (radula and egg-
capsules)

Shell: Shell solid and heavy, more slender than that of
Chicoreus brunneus (Link), uniformly dark brown in
colour, spire elevated, whorls convex. Teleoconch of 7

whorls, protoconch with 2 orange-brown, smooth and bul-
_ bous nuclear whorls which are calcified in large specimens.
Sculptured with gemmate spiral cords and close-set inter-
mediate spiral ridges, 3 varices per whorl, and 1 to 2
nodes betwen varices. The frondose spines are moderately
short on early whorls, but prominent on the labial varix
and siphonal canal; this feature, however, is variable,
and in some specimens the varical fronds are quite short.
The labial varix has generally 5 primary and 5 secondary
fronds, and the siphonal canal 3 frondose spines on the
average. Aperture oviform, cream, orange or bluish-white
in colour, deep interior white. Labial lip with 10 to 15
sharp denticles, columella smooth or occasionally with 1
to 3 denticles, anal notch distinct. Siphonal canal moder-
ately produced, broad, slightly recurved, slit on ventral
side open.

Size: 25mm to 85mm.

Type locality: None. The species has been reported from
“Ostindien” by Martini (1777), and “Indonesia” is here
designated as type locality.

Habitat: Under coral, on muddy sand substratum in
shallow water. Common.

Distribution: Throughout the Fiji Islands. — From East
Africa to Japan, East Australia and the Tuamotu Archi-
pelago.

Discussion: The Martini figure 995 cited by R6pInc
(1798) depicts the species Murex torrefactus SoWERBY
and of authors. Figure 996 on the same plate is an imma-
ture specimen of the same species. The additional figure
997 cited by Linx (1807) is a white C’hicoreus species
with yellowish varices and is undeterminable.

It is unfortunate that an older and prior name has to
be re-introduced into the literature in replacement for
Murex torrefactus Sowersy. Although both Purpura car-
neola Répinc and Triplex abortiva Perry could qualify
as nomina oblita under the 50-year rule of the Code of
ICZN (1964; art. 23b), Purpura elongata Link does not.
ToMLIN & WinckworTH (1936) drew attention to Linx’s
P. elongata which they rightly synonymized with Murex
torrefactus SowERBy. Even though P elongata Linx is
neither a primary nor a secondary homonym of Murex
elongatus LicHTFooT (one is a Chicoreus species, the other
a Pterynotus species), secondary homonymy could be
introduced by writers assigning both species to Murex s. lat.
In this particular case I thought it advisable to re-intro-
duce R6pino’s prior name.

Chicoreus laciniatus (SowERBY, 1841)
(Plate 14, Figures 9, 9a, 9b; Text figure 3)

1841. Murex laciniatus SowErBy, Conch. Illust., Murex, fig. 59
(non DesHayes & Mi_nE-Epwarps, 1843)

1960. Murex (Chicoreus) raciniatus (sic) SowERBY, AZUMA,
Cat. Moll. Okin. Japan., p. 33, pl. 4, fig. 10

Shell: Shell moderately solid, orange-brown to greyish-
brown in colour, spire elevated, whorls convex. Teleoconch
of 6 to 7 whorls, protoconch of 14 smooth nuclear whorls
which are generally calcified in mature specimens. Sculp-
tured with strong spiral cords, 3 varices per whorl, 2
nodes between varices and axial ribs on earlier whorls.
The frondose spines are short or very short, numbering
from 7 to 9 on the labial varix and 2 to 3 on the siphonal
canal. Aperture moderately large, oviform, mauve in col-
our; labial lip with 12 to 17 denticles, columella smooth,
anal notch moderately distinct. Siphonal canal moderately
short and broad, slit on ventral side open. Operculum with
a basal nucleus.

Radula: The radula (Text figure 3) is of the Chicoreus
type, rhachidians with 3 main cusps and 2 only slightly

Page 120

smaller intermediate cusps. The radular ribbon is 4.6mm
long and 0.22 mm wide in a shell 39mm in length. The
ribbon is translucent white and contains 182 rows of
teeth (+ 7 nascentes).

100 p

Figure 3
Chicoreus laciniatus (SOWERBY)

a. Half-Row of Radular Teeth
Fiji Islands

b. Operculum

Size: 32mm to 45mm.
Type locality: None. (“Les cétes de lisle Aroé, mers du
Japon,” fide Kiener, 1843).
Habitat: Under coral, on sand and coral-rubble substra-
tum from 0 to 5 fathoms. Rare.
Distribution: Mamanuca group, West off Viti Levu. —
Philippine Islands, Japan.
Discussion: Fiji specimens have always a rose-purple or
mauve coloured aperture, and conform in this feature
with Sowerpy’s illustration. Japanese specimens of Murex
laciniatus, as figured by AzuMaA (1960), Hase (1961)
and D’Attitio (1966) show the aperture to be cither
white or cream.

The varical fronds are short or even blunt in this specics,
a feature which may prompted some writers to assign
this species to Naquetia JoussEAuME. The species Chico-
reus microphyllus (LAMARCK) has also degencrate fronds
or spines, as do some specimens of C. carneolus (R6pING) ;
it would appear that the length of fronds or spines is a
rather variable feature and an unrcliable character for
generic assignment.

THE VELIGER

Vol. 10; No. 2

Chicoreus microphyllus (Lamarck, 1816)
(Plate 14,Figure 10; Text figure 4)

1816. Murex microphyllus Lamarck, Tabl. Encycl. Meth., p. 4,
pl. 415, fig. 5

1833. ? Murex rubescens BroveEriP, Proc. Zool. Soc. London for
1832, pt. 1: 174

1881. Chicoreus poirieri JoussEAUME, Le Naturaliste, 2 (42) :
349 (New Caledonia)

Shell: Shell moderately solid, white to cream in colour,
ornamented with blackish-brown cords and patches on
varices. Telcoconch of 7 whorls, protoconch with 1}
smooth nuclear whorls. Sculptured with strong spiral
cords, fine intermediate spiral lirac and 3 varices per
whorl; varical fronds degenerate, appearing as short foli-
ations, numbering from 5 to 7 on the labial varix and from
2 to 3 on the siphonal canal. Aperture modcrately small,
cream in colour, labial lip with 10 to 15 denticles, colum-
ella with 13 to 15 small denticles, anal notch prominent.
Siphonal canal moderately slender, slit on ventral side
open. Operculum (Text figure 4) orange-brown and with
a basal nucleus.

Figure 4

Operculum of Chicorcus microphyllus (LAMARCK)

Size: 30mm to 43mm.

Type locality: None. (“Tahiti,” fide BropErip, 1833).
Habitat: Under coral rock, on clean sand substratum,
from 0 to 3 fathoms. Rare.

Distribution: North Viti Levu. — From the Seychelles
Islands to Japan, East Australia and the Society Islands.
Discussion: Chicoreus microphyllus differs from other
members of the genus in features of blunt varices and
denticulate columella. These two charactcrs seem to be
rather variable and appear in specics of the genera Ghico-
reus, Pterynotus and Haustellum.

Chicoreus ramosus (LINNAEUS, 1758)
(Plate 14, Figure 11; Text figure 5)

1758. Murcx ramosus LinNAEuS (pars), Syst. Nat., ed. 10, p.
747, no. 488 (Rumputus, 1705, t. 26, fig. A — first correct ref.)

Vol. 10; No. 2

1798. Purpura ramosa var. a Répinc, Mus. Bolten., p. 142, no.
1786 (ref. Martin1, 1777, 3, t. 103, figs. 981, 982)
1798. Purpura incarnata Rovinc, Mus. Bolten., p. 142, no. 1791
(ref. MartINI, of. cit., t. 102, figs. 980 & t. 103, fig. 981)
1822. Murex inflatus Lamarck, Hist. nat. anim. sans vert., 7:
160 (ref. Martini, of. cit., figs. 980, 981)

1825. Murex monodon (pars) Sowersy, Cat. shells coll. Tank.,
App. p. 19 (ref. Martint, of. cit., fig. 980 only)

1852. Murex frondosus Morcu, Cat. conch. Yoldi, 1: 97 (non
Triplex frondosus Perry, 1811 = Chicoreus sp.)

Shell: Shell large and solid, white or creamy-white in
colour, ornamented with brown spiral lines and occasional
brown patches near sutures. Teleoconch consists of 6 to 7
whorls, protoconch generally calcified. Sculptured with a
few strong spiral cords, numerous fine spiral lirae and 1 to
2 nodes between varices; varices number 3 per whorl or
4 varices on earlier whorls. Varical fronds moderately
produced, foliated, open and recurved; labial varix with
6 to 10 fronds, siphonal canal with 2 to 3. Aperture large,

100
eae |

Figure 5

Chicoreus ramosus (LINNAEUS)
a. Half-Row of Radular Teeth
Fiji Islands

b. Operculum

roundly ovate, rosy-red on periphery, white in interior.
Labial lip with 11 to 17 denticles, fifth denticle anteriorly
of the siphonal canal larger than others; columella smooth,
anal notch prominent. Siphonal canal broad and recurved,
slit on ventral side open. Operculum with sub-basal
nucleus.

THE VELIGER

Page 121

Radula: Radular ribbon (Text figure 5) is 24.0mm long
and 0.80mm wide in an animal with a shell 216mm in
length. The ribbon contains 215 rows of teeth (++ 13 nas-
centes) ; rhachidians equipped with 3 main cusps and 2
rather small intermediate denticles,

Size: 100mm to 230mm.

Type locality: In Sinu Persico, Jamaica; (latter locality
erroneous).

Habitat: On coral reefs, on clean sand substratum, from
0 to 5 fathoms. Moderately common.

Distribution: Throughout the Fiji Islands. — From the
Red Sea to Japan, East Australia and Samoa.
Discussion: Murex ramosus LINNAEUS is such a badly
conceived species that it is questionable whether it can be
retained in muricid nomenclature either as a species or a
type species. From the 16 figures cited by Linnazus
(1758) for M. ramosus, only 2 or possibly 3 figures depict
M. ramosus auctt.; the citations cover 7 different species.
Gme.in (1791) and Dittwyn (1817) adopted the same
confused synonymy. Répinc (1798) cited MARTINI
(1777), figures 981 and 982 for Purpura ramosa var. a;
the figures cited represent Murex ramosus auctt. and M.
brevifrons LaMarckK, 1822 respectively. Purpura ramosa
Linx, 1807 is identical with R6p1Ne’s species. The M.
ramosus of MontrFort, 1810, generally cited as the type of
Chicoreus Montrort, 1810 is M. brevifrons LaMaRcK.
Murex inflatus Lamarck, 1822 is M. ramosus auctt. and
is an objective synonym of P incarnata Répinc, 1798 and
a primary homonym of M. inflatus Broccuy 1814.

Murex ramosus of authors will have to remain a species
without a holotype since the Linnean collection contains
2 syntypes of M. ramosus LINNAEUS, one of which is the
species M/. pomum GMELIN, 1791 and the other M. brun-
neus (Linx, 1807) ; both specimens appear to be genuine
syntypes as both were marked in Linnaeus’ handwriting
for M. ramosus (fide Dopce, 1957). Neither of the two
syntypes would qualify for a lectotype selection of M.
ramosus auctt., and a re-introduction of Purpura incarnata
R6pine, 1798 for the large Indo-Pacific Chicoreus species
may prove to be the most acceptable course in the
circumstances.

On the basis of cited figures, Murex monodon SowErRBy,
1825 is M. ramosus LINNAEUS in part only. SowERBy
(1825) refers to “Martini [1777], Conch. Cab., 3, t. 105,
figs. 987, 980” for M. monodon; figure 987 is M. monodon,
figure 980 is M. ramosus. However, figure 980 does not
occur on plate 105 but on plate 102, and it is therefore
probable that figure 980 is a transcription error for figure
988 on plate 105, which is indeed M. monodon. Purpura
cornucervi R6pinGc, 1798, was based on the same figures
987 and 988, and has priority.

Page 122

Chicoreus saulit (SowERBy, 1841)
(Plate 15, Figure 12)

1841. Murex saulit SowErsy, Conch. Illust., Murex, fig. 77

1841. Murex saulii Sowersy, Proc. Zool. Soc. London for 1840,
pt. 8: 141

1853. Murex (Chicoreus) sauliae (sic) SowErBy, H. « A. ADAMS,
Gen. Rec. shells, 1: 73

1896. Murex saulae (sic) SowErRsy, Croucu, Proc. Malac. Soc.
London, 2 (3): 135

1959. Murex (Chicoreus) sawlii (sic) Sowersy, Kira, Col. illust.
shells Japan, 1: 189, pl. 69, fig. 1

Shell: Shell moderately light in weight, orange-brown in
colour, ornamented with dark brown spiral cords. Teleo-
conch consists of 7 to 8 whorls, protoconch with 14 brown
and smooth nuclear whorls. Sculptured with spiral cords
and fine, gemmate intermediate spiral lirae, 1 to 2 nodes
between varices which number 3 per whorl. Varical fronds
moderately produced, foliated and open, numbering from
4 to 6 on the labial varix and 2 to 3 on the siphonal canal.
Aperture moderately large, white in colour, occasionally
with a light rosy tinge; labial lip with 13 to 18 denticles,
columella smooth, but occasionally with a single denticle
near the siphonal canal; anal notch prominent. Siphonal
canal moderately slender and produced, slit on ventral
side open.

Size: 64mm to 75mm.

Type locality: Insulam Capul, Philippinarum.
Habitat: Dredged in 8 fathoms on coral-rubble substra-
tum. Rare.

Distribution: South Viti Levu. — From the Philippine
Islands to Japan, East Australia and the Fiji Islands.
Discussion: As is evident from the synonymy, this specific
name is frequently misspelled, due no doubt to SowErBy’s
unusual latinization of a patronymic name. Miss Jane
Saul had 8 species named in her honour which were
spelled either “saulae” or “sauliae.’ SowERBy named 2
of these, one which he spelled Murex saulit while he
spelled the other Marginella sauliae Sowrrsy, 1846, al-
though both species originated from the same collection.

Pterynotus Swainson, 1833

Pterynotus Swainson, 1833, Zool. Illust., ser. 2, pl. 100 — Type
species by SD (Swarnson, 1833) Murex pinnatus Swain-
son, 1822 = Pterynotus alatus (R6p1NG, 1798)

Characters: Shell light in weight, slender, spire high,
whorls convex. Sculptured with numerous fine spiral cords,
axial riblets in interstices, thin, compressed and incised
varices, 3 varices per whorl and nodes between varices;
varical spines or fronds absent in the type species, but
may appear as curved hook-like appendages in other mem-
bers of the genus. Aperture small, elongate-ovate, labial

THE VELIGER

Vol. 10; No. 2

lip denticulate, columella generally smooth, anal notch
weak. Siphonal canal slender, produced and recurved,
slit on ventral side open.

Operculum with a basal nucleus. The rhachidians of -

the radula are equipped with 3 large, almost equal-sized
main cusps, while intermediate denticles are absent (fide
Hase in VoKEs, 1964).
Discussion: The type species Murex pinnatus SwAINSON
was no stranger to 18th century iconographers. MARTINI
(1777) describes in detail his “gefliigelte dreieckige Pur-
purschnecke” and figures the species on plate 111, figures
1036 and 1037. These figures are unquestionably Murex
pinnatus and were cited by ROopinc (1798) for Purpura
alata (“The white winged Purpura snail” [transl.]). The
synonymy of M. pinnatus SwAINSoN appears to be as
follows:

1791. Murex ramosus var. e (pars) GMELIN, Syst. Nat., ed. 13,
p. 3528, no. 13 (non Linnagus, 1758)

1798. Purpura alata Rovinc, Mus. Bolten., p. 144, no. 1085 (ref.
Martint, 1777, 3, t. 111, figs. 1036, 1037)

1822. Murex pinnatus Swainson, Cat. coll. Bligh, App. p. 17
(non Triplex pinnata Perry, 1811 = Pterynotus sp.)

1840. Murex martinianus PFEIFFER, Krit. Reg. Mart. & Chemn.,
p. vii (ref. MarTINI, of. cit., figs. 1036, 1037) [non REEVE,
1845]

PFEIFFER (1840) in his critical analysis of MARTINI &
CHEMNITz’s “Conchylien-Cabinet” remarks that he could
elucidate the species illustrated in the first four volumes
through the study of Martini’s original specimens; the
labels were marked in Martini’s own handwriting and
were in the Museum of Pfeiffer’s brother-in-law, Mr.
Hermann Nathusius of Hundisburg. Pfeiffer considered
the species figured by Martini (op. cit.) in figures 1036
and 1037 as undescribed, overlooking R6p1ne’s prior de-
scription. DesHAyES & Mi_tnE-Epwarps (1843) as well
as DuNKER (1882) refer the Martini figures 1036 and
1037 to the synonymy of Murex pinnatus Swainson.

Not only is Pterynotus alatus (ROpING) an earlier name
for Murex pinnatus, but SwAINSON’s species name may
well be a secondary homonym of Triplex pinnata Perry,
1811 (plate 7, figure 5). PErrRy’s figure represents a white
Pterynotus species which has been synonymized, possibly
incorrectly, with M. tripterus Born, 1778 by DESHAYES
& Mitne-Epwarps (1843, p. 578). Murex phyllopterus
Lamarck, 1822, described from an unknown locality and
without figure citations, is almost certainly M. pinnatus;
the description fits the species perfectly.

Murex alatus GmELIn, 1791 (based on CHEMNITZ,
1780, 4, t. 159, figs. 1503, 1504) does not pre-occupy Pur-
pura alata Ropinc, 1798, as GMELIN’s species is a
clavinine turrid better known as Pleurotoma crenulata
Lamarck, 1822.

Vol. 10; No. 2

THE VELIGER

Page 123

No type locality has been cited by Répinc (1798) for
his Purpura alata; however, Martini (1777, p. 349)
reports the species from the coast of Coromandel, ‘Tran-
quebar [= Tranquebar, southeast coast of India].

Pterynotus elongatus (LicutrFoor, 1786)
(Plate 15, Figure 13)

1786. Murex elongatus Licutroot, Cat. Port. Mus., p. 65, no.
1479 (ref. FavaNNeE, 1780, pl. 79, fig. H)

1791. Murex ramosus var. e (pars) GMELIN, Syst. Nat., ed. 13,
p. 3528, no. 13 (non Linnaeus, 1758)

1798. Purpura draco Répvinc, Mus. Bolten., p. 144, no. 1809 (ref.
Martin1, 1777, 3, fig. 1033)
1822. Murex uncinarius Lamarck, Hist. nat. anim. sans vert., 7:
166 (ref. Martint, op. cit., t. 111, figs. 1034 ?, 1035 ?)
1842. Murex clavus Krengr, Spéc. Gen. Icon. Coq. Viv., Murex,
pp. 111-112, pl. 37, figs. 2, 2 (ref. Martint, of. cit., t. 111,
figs. 1033, 1034, 1035) [non MicueEtortTi, 1841)

1967. Murex elongatus SoLANDER, Cross, Hawai. Shell News,
15 (1): 1, 2 figs. (animal)

Shell: Shell light in weight, white or creamy-white in
colour throughout; spire very long and slender, whorls
convex. Teleoconch consisting of 7 whorls, protoconch cal-
cified in adult specimens. Sculptured with spiral striae
which are close-set and prominent on varices, granulose
spiral cords at base and 3 wing-like, thin and compressed
varices at every whorl; earlier whorls axially ribbed
between varices. Fronds or spines generally absent, but
some specimens do have a few curved, open and hook-like
varical appendages. Aperture small, elongate-ovate, white
or creamy-white, sometimes with a pinkish tinge; labial
lip with 15 to 20 denticles, columella elevated and with
2 to 4 denticles situated near the siphonal canal, anal
notch obsolete. Siphonal canal moderately broad and
short, slit on ventral side open.

Size: 55mm to 70mm.

Type locality: None. “Ostindien, Batavia” was mentioned
as locality by Martini (1777) and “Batavia, Indonesia” is
here selected as type locality.

Habitat: On coral and clean sand substratum, from 3 to
5 fathoms. Rare.

Distribution: Throughout the Fiji Islands. — From the
Red Sea to Japan, North Australia and the Hawaiian
Islands.

Discussion: Martin's figure 1033 was based on a speci-
men from the Bolten collection (fide Martin, 1777, p.
349) ; the same specimen was later named Purpura draco
Ro6pinc, 1798, and the same figure 1033 has been referred
to by Kiener (1843) for his Murex clavus. Martint (op.
cit.) figured another specimen of M. elongatus on the
same plate in figures 1034 and 1035. The figured specimen,
which appears to be beach-worn (labial lip is missing)

has been described in detail by Martini; the author
mentioned the three-sided appearance of the shell and the
3 compressed varices as well as the shell’s fragile texture.
He went on to point out that the actual shell was larger
than depicted in the figure (ca. 56.0mm). These 2 figures,
however, were cited with a query for M. uncinarius
Lamarck, 1822, for which a size of 11 lignes (ca. 25.0
millimeters) was given. On the basis of LaMarcx’s figure
citations, M. uncinarius is unquestionably conspecific with
M. elongatus LicutFooT; his description also agrees with
the latter species, with the exception of the phrase “albido-
fulva” and the small size of LAMaRcK’s specimen. Murex
uncinarius KIENER, 1842 and of authors is a small muricid
species from South Africa, and is the type species of
Poropteron JoUSSEAUME, 1880. This is once again a case
where cited figures are in conflict with the subsequent
interpretation of the species and possibly the holotype. The
only possibility of evaluating M. uncinarius LAMARCK is
to figure and re-describe the holotype, and expunge the
cited Martini figures from the synonymy, provided that
LAMARCK’s species is indeed conspecific with KIENER’s
M. uncinarius. Another alternative would be to accept
the subsequent junior synonym M. capensis SOWERBY,
1841 (Conch. Illust., fig. 76). SowEerBy’s M. mitraeformis
(1841, fig. 75) has figure priority, but is a homonym of
M. mitraeformis Broccut, 1814.

Murex clavus MicHeEtotTt1, 1841 need not be replaced
on account of the earlier M/. clava GMELIN, 1791. The Latin
words clava and clavus are nouns of feminine and mascu-
line gender respectively, and the rule of adjectival species-
group names does not apply in this case.

Kiener’s plates to Murex were issued in 1842 (fide
SHERBORN & Woopwarp, 1901, p. 217) and the text in
1843. The plates contain the specific names at the lower
margin and K1EnEr’s Murex species have been established
as from the date of issue of the plates.

(Naquetia) JoussEAUME, 1880

Naquetia JoussEAuME, 1880, Le Naturaliste, 2 (42) : 335 — Type
species by OD Murex triqueter Born, 1778

Characters: Shell solid, moderately slender, spire elevated,
whorls convex. Sculptured with coarse spiral cords, axial
ribs and 3 varices per whorl; varices are not wing-like
and compressed as in Pterynotus, and flat flanges are
evident on the labial varix. Aperture oviform, labial lip
denticulate, columella smooth or denticulate, anal notch
shallow or obsolete. Siphonal canal broad and moderately
short.

Operculum with a basal nucleus. The rhachidians of
the radula are broad and short, with only 3 cusps of

Page 124

THE VELIGER

Vol. 10; No. 2

ne, TN EN ET

which the central cusp is twice as long as the side-cusps;
intermediate cusps are degenerate, indicated only as verti-
cally incised wrinkles in some specimens, or are absent
altogether in other specimens.

Discussion: In shell and radula characters, this group is
intermediate between Chicoreus MontFort and Ptery-
notus Swainson. The radula, although similar in charac-
ters to Pterynotus (vide Hase in Vokes, 1964), differs
sufficiently to separate the group at least subgenerically.
The 3 main cusps in Pterynotus are large and almost of
equal size, and rather similar in pattern to the radula of
Typhis tosaensis AzuMA, 1960 (p. 99, text fig. 2). In
Naquetia, however, the main central cusp is twice as long
as the flanking main cusps, and remnants of the small
intermediate denticles may occasionally be seen in some
specimens.

Pterynotus (Naquetia) tripterus (Born, 1778)
(Plate 15, Figure 14)

1778. Murex tripterus Born, Ind. rer. nat. Mus. Caes. Vindob.,
p. 287

1780. Murex tripterus Born, Test. Mus. Ces. Vindob., p. 291, t.
t. 10, figs. 18, 19

1834. ? Murex trialatus var. SowERBY, Conch. Illust., Murex, fig.
54 only

Shell: Shell solid and heavy, dirty-white or light fawn in
colour throughout. Teleoconch consists of 5 to 6 whorls,
protoconch with 2 calcified nuclear whorls. Sculptured
with spiral ridges, 3 varices per whorl and a single node
between varices; labial varix large and compressed. Aper-
ture elongate-oviform, flesh or cream in colour, labial
lip with 6 to 8 white denticles and small accessory den-
ticles; columella with 7 to 10 white denticles, anal notch
obsolete. Siphonal canal broad and short, slit on ventral
side open. Operculum orange-brown and with a sub-basal
nucleus.

Size: 30mm to 60mm.

Type locality: Coasts of Batavia.

Habitat: Under coral rocks, on muddy sand substratum,
in shallow water. Rare.
Distribution: West Viti Levu. —
Japan and the Fiji Islands.
Discussion: DesHayes & MiLNe-Epwarps (1843, p. 578)

From Indonesia to

place Triplex pinnata Perry, 1811 in the synonymy of
Murex tripterus Born. Prerry’s figure does not quite
resemble M. tripterus, unless it be a very worn specimen.

Pterynotus (Naquetia) triqueter (Born, 1778)
(Plate 15, Figure 15; Text figure 6)

1778. Murex triqueter Born, Ind. rer. nat. Mus. Caes. Vindob.
p. 288

1780. Murex triqueter Born, Test. Mus. Caes. Vindob., p. 291,
t. 11, figs. 1, 2

1791. Murex ramosus var. € (pars) GMELIN, Syst. Nat., ed. 13,
p. 3528, no. 13 (non Linnagus, 1758)

1798. Purpura variegata Rovinc, Mus. Bolten., p. 143, no. 1799
(ref. Martin1, 1777, 3, t. 111, fig. 1038)

1798. Purpura cancellata Ropinc, Mus. Bolten., p. 143, no. 1801
(ref. MarTInt, of. cit., t. 111, fig. 1038)

1811. Triplex flexuosa PERRY, Concheleay pl. 7, fig. 1

1816. Murex trigonulus Lamarck, Tabl. Encycl. Méth., p. 5, pl.
417, figs. 4a, 4b

1907. Murex (Chicoreus) triqueter var. amanuensis COUTURIER,
Journ. Conchyl., 55: 142

Shell: Shell moderately light in weight but solid, spire
elevated; uniformly creamy-white or light brown, macu-
lated with dark brown on varices and labial lip. Teleo-
conch consists of 5 to 6 whorls, protoconch with 2 smooth
brown nuclear whorls. Sculptured with spiral cords and
intermediate spiral ridges, 3 varices per whorl and 2 to 3
axial ribs between varices; varices foliated and somewhat
compressed anteriorly. Aperture moderately small, ovi-
form and white, labial lip with 12 to 16 denticles;
columella smooth, occasionally with a denticle near the
siphonal canal, anal notch obsolete. Siphonal canal broad
and short, or moderately slender and slightly produced,
slit on ventral side open. Operculum with a basal nucleus.
Radula: The radula is of the Pterynotus type, with 3
main cusps only, but occasionally with extremely weak
vestiges of intermediate denticles which do not protrude
past the top of the plate. Length of radular ribbon
6.3mm, width 0.31 mm in an animal with a shell 58mm
in length; the ribbon is white and numbers 102 rows of
teeth (+ 6 nascentes) (sce Text figure 6).

Size: 40mm to 70mm.

Type locality: None. (“Tranquebar, Ostindien,’ MartTINI,
TD)

Explanation of Plate 15

Figure 12: Chicoreus saulii (SowERBy). x 0.7
Figure 13: Pterynotus clongatus (LicutFoor). x 0.9
Figure 14: Pterynotus triptcrus (Born). x 0.9
Figure 15: Pterynotus triqueter (Born). x 0.9

Figure 20:

Figure 16: Homalocantha anatomica (PERRY). x 0.9
Figure 17: Poirieria nodulifera (SowERBY). Xx 0.9
Figure 18: Vitularia miliaris (GMELIN). x 1.9 & 0.8
Figure 19: Favartia brevicula (SOWERBY). 2.0

Favartia tetragona (BRopERIP) . xX 1.5

Tue VELIGER, Vol. 10, No. 2 [CERNOHORSKY] Plate 15

Figure 12 Figure 13 Figure 14 Figure 15 Figure 16

Figure 17 Figure 18

|
|
Figure 19 Figure 20
|
|

photographs by W. O. CernoHorsky

Vol. 10; No. 2

b

Figure 6
Pterynotus triqueter (Born)
a. Half-Row of Radular Teeth
Fiji Islands

b. Operculum

Habitat: Under coral rocks, on clean and muddy sand
substratum, in shallow water. Uncommon.
Distribution: Throughout the Fiji Islands. — From East
Africa to Japan, East Australia and the Tuamotu Archi-
pelago.
Discussion: The type figures of Murex trigonulus La-
MARCK, 1816, clearly depict M. triqueter Born; six years
later Lamarck himself placed the species in the synonymy
of M. triqueter Born.

Murex triqueter amanuensis CouTuRIER appears to be
a smaller, more slender and elongated variant which
occurs sporadically in Fiji and the Philippine Islands, and
is of no racial significance.

Poirieria JOUSSEAUME, 1880
(Text figure 7)

Poirieria JoUSSEAUME, 1880, Le Naturaliste, 2 (42): 335 — Type
species by OD Murex zelandicus Quoy « Gatmarp, 1833

Characters: Shell light in weight, spire elevated, whorls
angulate, varical spines moderately long and open, but
becoming progressively shorter towards early whorls.
Sculptured with weak spiral ridges and 5 to 6 varices per
whorl. Aperture wide and oval, labial lip with foliated
depressions, columella smooth and calloused, anal notch
obsolete. Siphonal canal moderately long, open and re-
curved. Operculum with a basal nucleus.

THE VELIGER

Page 125

Discussion: The radula of the type species Poirieria zelan-
dica is figured here (Text figure 7). The radula was
extracted from a specimen collected off Mayor Island,
Bay of Plenty, New Zealand in 1926, and the slide was
made available by Dr. A. W. B. Powell. The radular ribbon
measured 6.7 mm in length and 0.24 mm in width, length
of shell unknown; the radular ribbon contained 182 rows
of teeth, and no nascentes were visible.

| 100 p }

oj

Figure 7
Poirteria zelandica (Quoy « GAIMARD)
a. Half-Row of Radular Teeth
New Zealand

b. Operculum

Poirieria nodulifera (SOWERBY, 1841)
(Plate 15, Figure 17)

1841. Murex noduliferus Sowersy, Conch. Illust., Murex, fig.
101

1841. Murex noduliferus SowErsy, Proc. Zool. Soc. London for
1840, pt. 8: 147

Shell: Shell small and light in weight, spire elevated;
creamy-white to yellowish in colour, irregularly lined with
brown, varical spines and siphonal canal stained brown.
Sculptured with spiral ridges, 6 to 7 varices on body
whorl and 7 to 9 varices on earlier whorls; varical spines
prominent and open, numbering from 4 to 6 on the labial
varix and from 0 to 1 on the siphonal canal. Aperture
oviform, white or creamy-yellow, labial lip with 5 to 7
denticles; columella smooth and calloused, occasionally

Page 126

with 0 to 4 denticles near the siphonal canal, anal notch
obsolete. Siphonal canal slender, moderately long, open
and recurved.

Size: 20mm to 32mm.

Type locality: Insulam Masbate [Philippine Islands].
Habitat: On sand and coral-rubble substratum in 8
fathoms. Moderately rare.

Distribution: North and South Viti Levu. — From the
Philippine Islands to Japan, New Guinea and the Fiji
Islands.

Discussion: SH1kAMA (1963) assigned the species to the
subfamily Drupinae, genus Drupa Ropinc, 1798, sub-
genus Morula ScHUMACHER, 1817, but I believe that a
placing of this species in Potrieria is more appropriate.
This species has an elevated spire, not unlike the type
species of Mfurexsul IrREDALE, 1915, and the denticulate
labial lip is a character evident in Muricopsis Bucquoy,
DAUTZENBERG & DoLtFrus, 1882. The number of varices,
produced and open varical spines and features of siphonal
canal are characters associating the species with Poirieria
JoUSSEAUME.

Favartia JoUSSEAUME, 1880

Favartia JoUSSEAUME, 1880, Le Naturaliste, 2 (42): 335 — Type
species by OD Murex breviculus SowerBy, 1834

Characters: Shell small and solid, teleoconch with about
5 whorls, protoconch with 14 bulbous nuclear whorls.
Sculptured with strong spiral cords and descending axial
ridges which appear on varices as deep pits; varices strong
and rounded, numbering 4 per whorl. Aperture roundly
ovate, elevated, labial lip scalloped, columella smooth,
anal notch absent. Siphonal canal short, recurved and
either slightly open or completely closed in adult specimens.
Operculum is muricine, orange-brown in colour and
with a basal nucleus. The radula type is muricine(?) (fide
THIELE, 1929).
Discussion: Voxes (1964) placed Favartia in the genus
Aspella and remarked that the group was intermediate
between Muricinae and Tritonaliinae on account of the
operculum which is muricine and a radula which is closer
to Tritonaliinae. The operculum is undoubtedly muricine,
and the radula of Aspella was described as muricine by
THIELE (op. cit.), 1. e. “rhachidian of radula with trian-
gular central cusp and two smaller denticles between the
three main cusps” [transl.]. Favartia, however, may not
be related to Aspella at all, especially if the radulae of the
two genera could be compared. Favartia, and especially
the species F tetragona (Broprrip), bears a resemblance
to Nothotyphis FLEm1nG, 1962, in features of shell appear-
ance, size, latticed sculpture, guttered spines and com-
pletely closed canal.

THE VELIGER

Vol. 10; No. 2

Favartia brevicula (SowERBY, 1834)
(Plate 15, Figure 19; Text figure 8)

1834. Murex breviculus Sowursy, Conch. Illust., Murex, fig. 37 .
1841. Murex breviculus SowErby, Proc. Zool. Soc. London for
1840, pt. 8: 146

Shell: Shell small and solid, dirty-white in colour, whorls
convex. Teleoconch consisting of 5 whorls, protoconch
with 14 bulbous calcified nuclear whorls. Sculptured with
5 to 6 strong spiral cords which are intersected by des-
cending axial ridges which are especially prominent on
varices; varices generally blunt and rounded, numbering
4 per whorl, and ornamented with 1 to 2 short spines.
Aperture roundly-ovate, elevated, white, deep interior
purplish; labial lip with 6 to 11 scalloped denticles, colu-
mella white, calloused and smooth, anal notch absent.
Siphonal canal short and slender, slightly open or com-

Figure 8

Operculum of Favartia brevicula (SOWERBY)

pletely closed in adult specimens, and recurved almost at
90° towards the dorsum. Operculum orange-brown and
with a basal nucleus (Text figure 8).

Size: 15mm to 30mm.

Type locality: None.

Habitat: Under coral rocks, on muddy sand and coral
substratum in shallow water. Moderately common.
Distribution: Throughout the Fiji Islands. — From the
Philippine Islands to Japan, East Australia and Tonga
Islands.

Discussion: The siphonal canal is more slender and sharply
recurved in Favartia brevicula than in F tetragona. The
siphonal canal is either slightly open or completely closed
in adult specimens.

Favartia tetragona (Broperip, 1833)
(Plate 15, Figure 20)
1833. Murex tetragonus Broperir, Proc. Zool. Soc. London for
1832, pt. 1: 174

1834. Murex tetragonus Broperirp, SowERBy, Conch. Illust.,
Murex, fig. 25

Vol. 10; No. 2

Shell: Shell small and solid, white in colour throughout.
Teleoconch consisting of 4 whorls, protoconch of 14 nuc-
lear whorls. Sculptured with 8 to 10 spiral cords, obsolete
and irregular axial riblets and pittings in interstices of
cords; varices number 4 per whorl, and are large and com-
pressed and bear remnants of somewhat curved guttered
spines. Aperture small and oviform, white, interior laven-
der with dark brown bands. Labial varix broad and
flaring, labial lip ornamented with 7 to 8 raised and
scalloped denticles, columella smooth, anal notch absent.
Siphonal canal broad, short, completely closed and only
slightly recurved for a short distance. Operculum with a
basal nucleus; the animal is yellowish-brown in colour.
Size: 20mm to 36mm.

Type locality: None.

Habitat: Under coral rocks on clean sand and coral rubble
substratum, from 0 to 5 fathoms. Rare.

Distribution: Mamanuca group. West off Viti Levu. —- ?
Discussion: This species differs from Favartia brevicula in
features of form, sculpture, aperture and anterior canal;
the latter is broad and short and lacks the tube-like,
turned over extension of the canal of F brevicula.

Phyllocoma TAPPARONE-CANEEFRL 1881

Phyllocoma TappaRoNnE-CanErrt, 1881, Ann. Soc. Malac. Belg.
Mém., 15: 44 — Type species by SD (WeENz, 1941) Triton
convolutus BRopvErRIP, 1833

Characters: Shell small, ranellid in appearance, spire
higher than aperture, whorls subulate and convex, early
whorls fenestrate. Sculptured with strong spiral cords,
axial ridges and 2 plain but prominent varices at every
whorl. Labial lip calloused, glazed and smooth, anal notch
shallow; siphonal canal short, open and recurved.

Discussion: The genus is assigned to the Muricinae only
tentatively. THIELE (1929) describes and figures the radu-
la of Phyllocoma as having a rhachidian with almost
equal-sized 3 main cusps; the operculum is said to have
a basal nucleus. A tricuspid rhachidian of the radula
may be found in Muricinae, Typhinae and Rapaninae,
and an operculum with a basal nucleus is present in the
first 2 subfamilies; in shell morphology the genus appears
to stand alone. Shell morphology, however, is in certain
cases of little assistance in a supraspecific division of a
group. In the past, toxoglossate turrids were classed with
stenoglossate mitrids on account of the similarity in shell
morphology. It would be reasonable to assume that a
particular radula type is peculiar to a group of species due
to their evolutionary process through time from a common
ancestral stock, and not only because of divergence in
food requirement. It is possible, however, that the radula
type of Phyllocoma described by THIELE (op. cit.) may

THE VELIGER

Page 127

really belong to Galfridus IREDALE, 1924 and may after
all be appreciably different from Phyllocoma s. str. Until
more information is known about the anatomy of the
type species, the subfamilial position of Phyllocoma is
incertae sedis.

Phyllocoma convolutum (Broverip, 1833)
(Plate 14, Figure 4)

1833. Triton convolutus BropeErip, Proc. Zool. Soc. London for
1832, pt. 1: 7

Shell: Shell small and solid, ivory-white or fawn in colour,
spire high, whorls subulate and convex. Teleoconch con-
sists of about 8 whorls apart from protoconch which was
missing in specimens examined. Sculptured with promi-
nent raised spiral ridges which number from 18 to 20 on
the body whorl and 8 to 10 on the penultimate whorl;
spiral ridges closely axially striate, terminating as close-set
spiral grooves at the base. The first 5 whorls are fenestrate
in appearance, with axial and spiral ridges intersecting.
Aperture large and oval, ivory-white in colour, labial lip
slightly elevated and ornamented with 20 to 23 denticles
which extend towards the interior of the aperture; colum-
ella heavily calloused, smooth and porcellaneous, anal
notch moderately distinct, interior of aperture light brown.
Siphonal canal short and open, moderately recurved to-
wards the dorsum.

Size: 20mm to 26mm.

Type locality: None.

Habitat: On sand and coral rubble substratum in 15
fathoms. Rare.

Distribution: South Viti Levu and Mamanuca group,
West off Viti Levu. — From the Philippine Islands to
Japan, North Australia and the Fiji Islands.

Tritonaliinae

Homalocantha Morcu, 1852

Homalocantha Morcu, 1852, Cat. Conch. Yoldi, 1: 95 — Type
species by M Murex scorpio LiNNAEus, 1758

Characters: Shell moderately small, solid, spire low,
whorls angulate and convex. Teleoconch consisting of 4 to
5 whorls, protoconch of 1 to 14 nuclear whorls. Sculptured
with strong and distant spiral cords, fine intermediate
spiral striae, 5 to 6 varices on body whorl and 5 to 7
varices on penultimate whorl. Varical fronds moderately
long, T-shaped and open at distal end, numbering ca. 2
to 6 on the labial varix and 1 to 3 on the siphonal canal.
The webbing between labial digitations is prominent in the
type species but can be absent in other members of the
genus. Aperture small, roundly ovate, labial lip with

Page 128

THE VELIGER

Vol. 10; No. 2

scalloped denticles, columella smooth, anal notch absent.
Siphonal canal moderatcly broad and long, closed part
of the way, but open and recurved at end.

Homalocantha anatomica (PErRRy, 1811)
(Plate 15, Figure 16; Text figure 9)

1811. Hexaplex anatomica Perry, Conchology, pl. 8, fig. 2

1823. Murex rota MaweE, Linn. Syst. Conch., p. 131, pl. 26, fig. 3

1921. Murex pele Puspry, Proc. Acad. Nat. Sci. Philadelphia

for 1920: 318, pl. 12, figs. 29, 30

Shell: Shell moderately light in weight, solid, generally
dirty-white in colour throughout. Whorls angulate, num-
bering 4 apart from protoconch which is calcified in adult
specimens. Sculptured with 2 prominent cords on the
body whorl and fine small cords at the labial varix; varices
number from 5 to 6 per whorl, varical digitations prom-
inent, compressed and open. Aperture small, raised, white
in colour, labial varix with 2 primary digitations; labial
lip with irregular scalloped denticles, columella smooth,

Figure 9

Operculum of Homalocantha anatomica (PERRY)

anal notch absent. Siphonal canal moderately long, closed
for part of the way but open for about 3 of its total
length towards the distal end.

Size: 34mm to 56mm.

Type locality: East Indies.

Habitat: Under coral rocks, on sand and coral substratum,
in 0 to 2 fathoms. Moderately rare.

Distribution: Throughout the Fiji Islands. — From the
Red Sea to Japan, Fiji and the Hawaiian Islands.
Discussion: D’Attitio (1964, 1964a) specifically differ-
entiated Homalocantha anatomica and H. pele on the
basis of main cord characters. He described and figured
the Indian Ocean H. anatomica (from the Red Sea, Mo-
zambique and Zanzibar), showing 3 main primary cords
emanating from the body whorl and terminating in
frondose digitations. ‘The Hawaiian and Japanese H. pele
were shown to possess only 2 main cords. All Fiji speci-
mens collected to date are sculptured with only 2 main
cords on the body whorl, while all other cords are second-
ary and degencrate. No comparison could be made with

Indian Ocean H. anatomica; however, should these
differences in main cord characters really be stable, then
a subspccific separation of the Indian Ocean H. anatomica
(Red Sea to Philippine Islands) and the Pacific H. ana-
tomica pele (Japan to New Guinea, Fiji and the Hawaiian
Islands) would be warranted. For further discussion on
the species see REHDER (1964).

Vitularia Swainson, 1840

Vitularia Swainson, 1840, Treat. Malac., p. 297 — Type species
by M Vitularia tuberculata Swainson, 1840 = Murex mili-
aris GMELIN, 1791

Characters: Shell solid, moderate in size, whorls depressed,
body whorl keeled; whorls number 4 to 5 apart from
protoconch of 1 to 14 nuclear whorls. Sculptured with 7
to 9 oblique and nodulose varices, spiral ridges and occa-
sionally scabrous granules. Aperture elongate-ovate, labial
lip denticulate, columella sinuous, depressed and smooth,
anal notch obsolete. Siphonal canal short and open,
slightly recurved. Operculum with a dextro-lateral nucleus.
Discussion: Swainson (1840) applied the spelling Vitu-
lina on page 64 and Vitularia on page 297. Gray (1847,
p. 134) adopted Vitularia as the correct spelling from
SwaInson’s multiple original spellings.

IREDALE (1929) established the genus Transtrafer for
the new species T: longmani IREDALE, 1929, and in way
of explanation mentioned the “striking resemblance” of
his new species to the “American Murex vitulinus La-
MARCK.” This is obviously a case of mistaken identity, as
M. vitulinus Lamarck (= M. miliaris GMELIN) is an
Indo-Pacific species, and conspecific with T. longmani
IREDALE. Several writers have passed comments on the
casual way in which IREDALE erected his new molluscan
genera, and a detailed discussion on their nomenclatural
validity can be found in Sotem (1964). Transtrafer is just
another of IREDALE’s undefined generic groups, which
would have qualified as a nomen nudum had it been
described only 6 months later. Australian workers would
certainly render malacological science a great service by
re-describing IREDALE’s several hundred generic groups
and thousand odd species.

Vitularia miliaris (GmeEtNn, 1791)
(Plate 15, Figure 18; Text figure 10)

1791. Murex miliaris Gmr.in, Syst. Nat., ed. 13, p. 3536, no. 39
(ref. Martini, 1777, 3: 303, vign. 36, figs. 1-5 & CHEMNITZ,
1788, 10, t. 161, figs. 1532-1535)

1798. Purpura onagrina Roévinc, Mus. Bolten., p. 139, no. 1751
to 1753 (ref. sup. cit.)

1816. Murex vitulinus Lamarck, Tabl. Encycl. Meéth., p. 5, pl.
419, figs. 1a, lb, 7a, 7b -

Vol. 10; No. 2

1840. Vitularia tuberculata Swainson, Treat. Malac., p. 297

1843. Murex purpura DESHAYES & MILNE-Epwarps, Hist. nat.
anim. sans vert., 2éme edit., 9: 595 (nom. nov pro M. vitu-
linus Lamarck, 1816)

1861. Vitularia sandwicensis PEASE, Proc. Zool. Soc. London for
1860: 397 (spec. juv.)

1929. Transtrafer longmant IREDALE, Mem. Queensld. Mus., 9
(3) : 290, pl. 31, fig. 10 (shell), fig. 11 (protoconch)

Shell: Shell moderately heavy and solid, dirty-grey to
orange-brown in colour, occasionally ornamented with 1
to 3 rows of dark brown spots on varices on the body
whorl. Teleoconch consisting of 4 to 5 whorls, protoconch
of 14 nuclear whorls. Early whorls flattened, body whorl
with a peripheral keel and 7 to 8 oblique varices; sculp-
tured with tightly packed spiral ridges or scabrous small
granules. Aperture elongate, edge of aperture variable
in colour, either bluish-white, creamy-yellow or reddish-
orange, occasionally with 1 to 3 dark brown bars on labial
lip; deep interior of aperture generally white. Labial lip
angulate, ornamented with 7 to 13 denticles, columella

Figure 10

Operculum of Vitularia miliaris (GMELIN)

calloused, sinuous and smooth, anal notch obsolete. Sipho-
nal canal short and open and only slightly recurved.
Operculum (Text figure 10) with a dextro-lateral nucleus.
Size: 15mm to 63mm.

Type locality: None.

Habitat: Under coral rocks on muddy sand substratum,
in shallow water; juvenile specimens in sand and weed
pockets of coral reefs. Moderately common.
Distribution: Throughout the Fiji Islands. - From the
Seychelles Islands to Japan, East Australia and the
Hawaiian Islands.

Discussion: Vitularia miliaris is a variable species, espe-
cially in colour and sculpture. LAMARCK’s Murex vitulinus
is conspecific with M. miliaris Gmein. In LaMaArRck’s
1822 work, M. vitellinus bears the same synonymy as
M. miliaris GMELIN and Purpura onagrina RGpING.

THE VELIGER

Page 129

UNCONFIRMED REPORTS

Homalocantha martinetana (R6p1nc, 1798)

1798. Purpura martinetana Répinc, Mus. Bolten., p. 141, no.
1778 (ref. CHEmnitz, 1788, 10, t. 161, figs. 1536, 1537)
1817. Murex fenestratus Dituwyn, Descr. cat. rec. shells, 2: 716
(ref. CHEMNITZ, of. cit., figs. 1536, 1537 & FavaNneE, 1780,
t. 35, fig. C1)

1963. Murex (Homalocantha) fenestratus DitLwyN, SHIKAMA,
Sel. shells world, 1: 72, pl. 56, fig. 5)

This species is known from only one very worn specimen
found in shell debris at Caboni Beach, North Viti Levu.
The digitations on the labial varix are completely worn
away, but the deep pits between the axial ribs are still
visible.

The species has been reported from the Red Sea,
Japan and Okinawa.

FOSSIL RECORD

Murex cf. M. recurvirostris Broperip, 1833

1934. Murex (Murex) aff. recurvirostris BropErtP, Lapp, Bull.
Bern. P. Bishop Mus., 119: 224-226, pl. 40, figs. 3-5

Lapp (1934) reported the fossil species from stations
59 and 165 (banks of Wailoa river near Nasogo, Viti
Levu, elevation 995 feet) from deposits which are prob-
ably Lower Miocene (fide Lapp, 1966). This species
does bear a resemblance to the Recent species M. recurvi-
rostris BRopeRIP from the West Indies, and especially to
the subspecies M. recurvirostris rubidus F. C. Baker,
1897. It differs only in the slightly more numerous axial
ribs and denticulate columella. There is no similar species
living in Fiji today.

SUMMARY

The majority of the Fijian molluscs live within the littoral
zone, while only a few species are confined to deeper
water. Of the enumerated muricid species, 70% live in
the intertidal region and only 30% inhabit the sub-littoral
zone.

The total number of species recorded in the various
muricid genera is as follows:

Page 130
Murex 2
(Haustellum ) 1
Chicoreus 8
Pterynotus 1
(Naquetia) 2
Poirieria 1
Phyllocoma 1
Favartia 2
Homalocantha 1
Vitularia 1
Total 20 species
Addendum

Just before going to press we received the following para-
graphs from the author. We are pleased to be able to
include them here.

Prior to the manuscript’s going to press a specimen of
Homalocantha anatomica (Perry, 1811) was collected by
the author; the radula was extracted and examined. The
specimen had 3 main primary cords emanating from the
body whorl and terminating in T-shaped digits; as both
three-corded and two-corded specimens are found in the
Fiji Islands, the Pacific H. pele (Prrspry, 1921) cannot
be separated from the nominal species H. anatomica.
Radula: Rhachidians with 3 broad main cusps, interme-
diate denticles lacking; central cusp longer than side cusps.
Length of radular ribbon 6.1mm, width 0.16mm in an
animal with a shell 43.0mm long. The ribbon is trans-

\ 100p. j

SE

?

Figure 11

Half Row of Radular Teeth of Homalocantha anatomica (PERRY)
Fiji Islands

lucent white and is comprised of 295 rows (+17 nascen-
tes) of teeth; the teeth in the anterior rows are greatly
worn. The penis is light fawn in colour, 3.1 mm in length
and very similar to the penis of Chicoreus brunneus (Linx,
1807) and C. carneolus (R6pinc, 1798).

Animal: Sole of foot light creamy-brown in colour, dorsum
of foot white. Tentacles slender, moderately short, broad
at base, white in colour; siphon whitish, eyes very small,
black in colour.

THE VELIGER

Vol. 10; No. 2

The radula of Homalocantha anatomica is muricid, and
the genus therefore should be removed from Tritonaliinae
and placed in the subfamily Muricinae. Homalocantha
should be placed near Pterynotus Swainson, 1833 and
Naquetia JOUSSEAUME, 1880, as these genera encompass
species with tricuspid rhachidians without the intermedi-
ate denticles.

LITERATURE CITED

Apvams, Henry « ARTHUR ADAMS
1853[1853-1858]. The genera of Recent Mollusca; arranged accord-
ding to their organization. London (John van Voorst) 1: ito
xl; 1 - 484
Azuma, Masao
1960. A catalogue of the shell-bearing Mollusca of Okinoshiina,
Kashiwajima and the adjacent area (Tosa province) Shikoku,
Japan. pp. 1 - 102; 1-17; plts. 1-5 (20 March 1960)
BaRNARD, KEPPEL HARCOURT
1959. Contributions to the knowledge of South African marine
Mollusca, Part II :Gastropoda: Prosobranchiata: Rhachiglossa.
Ann. South Afric. Mus. 45 (1): 1-237; 52 figs. (June)
Broperip, WILLIAM JOHN
1833. No title. Proc. Zool. Soc. London for 1832, prt. 2:
173 - 181 (January 1833)
RuONANNI, FILIPPO
1684. — Recreatio mentis et oculi in observatione animalium tes-
taceorum curiosis naturae inspectoribus. Roma, prts. 1 - 3;
139 plts.
CarrikER, M. R.
1961. | Comparative functional morphology of boring mecha-
nisms in gastropods. Amer. Zool. 1; 263 - 266; 1 fig.
CERNOHORSKY, WALTER OLIVER
1964. The Cypraeidae of Fiji (Mollusca:Gastropoda).
The Veliger 6 (4): 177 - 201; plts. 21-26; 1 Text fig.; 1 map
(1 April 1964)
1966. The radula, egg capsules and young of Murex (Chico-
reus) torrefactus Sowrrsy (Mollusca: Gastropoda). The
Veliger 8 (4): 231-233; 6 text figs. (1 April 1966)
CHEMNITZ, JOHANN HiERONYMUS
1780 - 1795. Neues systematisches Conchylien-Cabinet (conti-
nuation of F H.W. Martini) Niirnberg, vols. 4- 11
1780 4: 1 - 344; plts. 122 - 159; 1788 10: 1 - 376; pits. 137
to 173; 1795 11: 1-310; plts. 174 - 213
Cotton, BERNARD CHARLES
1956. No. 8: Family Muricidae. Roy. Soc. South Austral.
Malac. Sect., 6 pp. (unpaginated); 2 plts.
D’Attitio, ANTHONY
1964. Some observations on Homalocantha anatomica and
Homalocantha pele. Hawaiian Shell News 12 (11): 2
(September 1964)

Vol. 10; No. 2

D’Attitio, ANTHONY
1964a. A prime differentiating characteristic between species of
Homalocantha. Hawaiian Shell News 12 (12) : 3; figs. 1-7
(October 1964)
1966. Notes on the Japanese species of the family Muricidae.
Hawaiian Shell News 14 (7) : 4-5; figs. 1-5 (May 1966)
DavuTzEnBERG, PHILIPPE & L.-J. BoucE
1933. Les mollusques testacés marins des établissements Fran-
cais de l’Océanie. Journ. Conchyl. 77 (2): 145 - 326
(25 July 1933)
Deswayes, Gérarp Pau & HENrt MitNE-EDwARDS
1843. Histoire naturelle des animaux sans vertébres. 2. édit.
Paris, 9: 1 - 728
Dittwyn, LEwis WESTON
1817. A descriptive catalogue of Recent shells, arranged
according to the Linnaean method; ... London (John «
Arthur Arch) pp. i-xii+1-1092+41-29 (unnumbered index) ;
plts. 1-5 [1: i-xii + 1-580; plts. 1-5 & 2: 581-1092 + 1-29]
Dopcrt, HENRY
1957. A historical review of the mollusks of Linnaeus. Part 5.
The genus Murex of the class Gastropoda. _—_ Bull. Amer. Mus.
Nat. Hist. 113 (2) : 73 - 224 (30 September 1957)
Dotirus, Gustave FE & PHitippE DAUTZENBERG
1932. Les mollusques de Fabius Columna.
76: 283 - 333; 15 plts.

DunkKER, GUILIELMO

Journ. Conchyl.

1882. Index molluscorum maris Japonici. Cassellis, pp.
1 - 301; plts. 1 - 16

FAVANNE, JACQUES DE MONTCERVELLE DE & JACQUES GUILLAUME DF
FAVANNE DE MoNTCERVELLE

1780. La conchyliologie, ou histoire naturelle des coquilles.
3. édit. [of A. J.D. d’Argenville’s “L’histoire naturelle . . . ”].
Paris, 3 vols.

FAavANNE, JACQUES GUILLAUME DE MONTCERVELLE

1784. Catalogue systématique et raisonné, . . . cidevant 4 M.

le C. de la Tour d’Auvergne. Paris; 9 plts.
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
decima tertia, aucta, reformata 1 (6): 3021 - 3910
Murex: 3524 - 3565

GrawaM, ALASTAIR

1941. | The oesophagus of the stenoglossan Prosobranchs.

Proc. Roy. Soc. Edinburgh 61: 737 - 761
Gray, JoHn Epwarp
1847. _A list of the genera of Recent Mollusca, their synonyma

and types. Proc. Zool. Soc. London 17 (178): 129-219
(30 November 1847)

Editio
Lipsia.

Hasse, TADASHIGE
1961. Coloured illustrations of the shells of Japan.
2: 1-183; plts. 1-66

Hoikusha

THE VELIGER

Page 131

IREDALE, Tom

1929. | Queensland molluscan notes, no. 1.
Mus. 9 (3): 261 - 297; plts. 30-31

Krener, Louts CHarres

Mem. Queensid.
(29 June 1929)

1842-1843. Spécies général et iconographie des coquilles vivantes;
genre Rocher. Paris 7: 1 - 130; plts. 1-47 [plates issued
1842; text issued 1843]

Lapp, Harry S.

1934. Geology of Viti Levu, Fiji. Bernice P. Bishop Mus.
Bull. 119: i- iii; 1-263; plts. 1-44; maps, text figs.

1966. | Chitons and Gastropods (Haliotidae through Adeorbi-
dae) from the Western Pacific Islands. Geol. Surv. Prof.
Pap. 531: i-iv; 1-98; plts. 1-16; maps

Lamarck, JEAN-BApTisTE PrerrE ANTOINE DE MONET DE

1816. ‘Tableau encyclopédique et méthodique des trois régnes
de la nature. Paris, pp. 1-16; plts. 391 - 488

1822. Histoire naturelle des animaux sans vertébres, 7 [Mol-
lusques]. Paris (“chez l’auteur, au jardin du Roi”) pp. 1-711

(August 1822)
Licutroot, JoHNn

1786. A catalogue of the Portland Museum, lately the property
of the Duchess Dowager of Portland, deceased ... . London.

pp. 1- vili+ 1 -194

Link, HErnricn FRriepRicH

1806 - 1808. Beschreibung der Naturalien-Sammlung der Uni-
versitat zu Rostock, Rostock. 6 parts in 1 vol., some
paginated separately [Mollusca: part 2: 82-100 — 29 March
1807; part 3: 101-160 - 17 May 1807; part 4: 6-23 — 25
December 1807; part 6: 33-37 — 5 June 1808).

LINNAEus, CaRoLus

1758. | Systema naturae per regna tria naturae. Ed. 10, refor-
mata. Holmiae 1 (1): 1 - 824

Mansour-Bek, J. J.

1934. Uber die proteolytischen Enzyme von Murex anguliferus
Lamk. Zeitsch. vergl. Physiol. 20: 343 - 369

MartTINI, FrrepRicH HEINRICH WILHELM
1777. | Neues systematisches Conchylien-Cabinet.
3: 1 - 434; plts. 66 - 121

Montrort, PrerrE DENYS DE

Niirnberg,

1810. Conchyliologie systématique et classification métho-
dique des coquilles. Paris, 2 vols.
1: i- Ixxxvii + 1-409; 2: 1-676 + 1-16
Perry, GEORGE

pee Lon=
(April 1811)

1811. | Conchology, or the natural history of shells:
don (Wm. Miller) pp. 1-4; plts. 1-61
PFEIFFER, Louris
1840. Kritisches Register zu Martini und Chemnitz, Systema-
tisches Conchylien Cabinet. Kassel, i- viii; 1-112
Pmssry, HENry Aucustus
1921. Marine mollusks of Hawaii; VIII-XIII. Proc. Acad.
Nat. Sci. Philadelphia for 1920; 72: 296 - 328; plt. 12; 11 text
figs. (January 1921)

Page 132

THE VELIGER

Vol. 10; No. 2

Reeve, Lovett AuGuUsTUS

1845-1846. Conchologia Iconica; monograph of the genus Murex.
London, plts. 1 - 36 (April 1845 - April 1846)

REHDER, Harap A.

1964. Notes on Hawaiian Muricidae.
12 (6): 4-5; figs. 1-8

Ropinc, PETER FRIEDRICH

Hawaiian Shell News
(April 1964)

1798. | Museum Boltenianum sive catalogus cimeliorum; pars
secunda Conchylia. Hamburg, i- viii; 1 - 199
RumpuHius, GEorcE EvERHARD
1705. | D’Amboinsche Rariteitkamer. Amsterdam, plts. 1-60

ScHEPMAN, MatrHeus MartTINUS
1913. The Prosobranchia of the Siboga Expedition. Part IV.
Rhachiglossa. Siboga Exped. 49d: 247 - 363; plts. 18 - 24
SHERBORN, CHartes Davis & BERNHARD BARHAM WooDWARD
1901. Notes on the dates of publication of the parts of Kiener’s
“Spécies général et iconographie des coquilles vivantes,” etc.
(1834-80). Proc. Malac. Soc. London 4 (5): 216-219
(25 July 1901)
Suixama, Toxo [& M. Horixosu!]
1968. Selected shells of the world illustrated in colours.
Tokyo, 1: 1-154; 102 col. plts.; 211 text figs.
SHOPLAND, E. R.
1902. List of marine shells collected in the neighbourhood of
Aden between 1892 and 1901. Proc. Malacol. Soc. London
5 (2): 171 - 182 (1 July 1902)
SmirH, Epcar ALBERT
1897. Notes on some type-specimens in the British Museum.
Proc. Malacol. Soc. London 2 (5): 229 - 232 (July 1897)
SoLtem, ALAN
1964. Amimopina, an Australian enid land snail. The Veli-
ger 6 (3): 115-120; 4 text figs. (1 January 1964)
WENZ, WILHELM

Sowersy, GrorcE BreTTINcHAM (first of name)

1825. A catalogue of the shells contained in the collection of
the late Earl of Tankerville, arranged according to the La-
marckian conchological system; together with an appendix,
containing descriptions of many new species, ... London
(E. J. Stirling) pp. i- vii+ 1-92; Appendix: i - xxxiv; plts.
1-8

Sowersy, GEorcE BrETTINGHAM (second of name)

1834-1841. Conchological illustrations. Murex. London, prts.
58-67 (30 May 1834 to 1 October 1834); prts. 187-199 (1
January 1841 to February 1841)

1841. No title. Proc. Zool. Soc. London for 1840, prt. 8:
135 - 151 (July 1841)

Srott, Norman Rupotpg, et al.

1964. International Code of Zoological Nomenclature adopted
by the XV International Congress of Zoology, ed. 2.
London (Internat. Trust f. Zool. Nomencl.) pp. i- xvii+
1-176; 5 appendices & Glossary

Swarnson, WILLIAM
1840. A treatise on malacology, or the natural classification
of shells and shellfish. London, pp. 1 - 419; text figs.
THIELE, JOHANNES

1929. Handbuch der systematischen Weichtierkunde. Erster
Teil, Loricata; Gastropoda. I: Prosobranchia (Vorderkiemer).
pp. 1-376; 470 text figs. Jena, Gustav Fischer Verlag
(Muricidae pp. 287 - 301)

Tomun, JoHN Reap LE BrocKTon « RONALD WINCKWORTH

1936. An index to the species of Mollusca in the Beschreibung
of H. F Linx. Proc Malacol. Soc. London 22 (1). 27 - 48
(14 March 1936)

VoKeEs, Emity H.

1964. | Supraspecific groups in the subfamilies Muricinae and
Tritonaliinae (Gastropoda : Muricidae). Malacologia 2
(1): 1-41; text figs.

1943. Handbuch der Palaozoologie; Gastropoda, Prosobranchia.

Berlin, 6 (1):
Wu, Sui-KvueE1

1 - 1639; 4211 text figs.

1965. Studies of the radulae of Taiwan muricid gastropods.
Bull. Inst. Zool. Acad. Sinica 4: 95 - 106; 35 text figs.

Vol. 10; No. 2

THE VELIGER

Page 133

A Review of the Living Tectibranch Snails
of the Genus Voloulella,

with Descriptions of a New Subgenus and Species from Texas

HAROLD W. HARRY

A&M Marine Laboratory, Galveston, Texas 77550 and Rice University, Houston, Texas 77001

(21 Text figures)

THIS STUDY BEGAN with an attempt to identify some
specimens of Volvulella from Galveston, Texas, which
proved to belong to an undescribed species more similar
to one from the west coast of North America than
to any hitherto known from the Atlantic. It is proposed
as the type of a new subgenus. Snails of this genus
live subtidally and to several hundred meters depth in
tropical and warm temperate oceans throughout the
world. They are local in occurrence and rarely abundant.
It is not surprising therefore that the limits of species
variation are poorly understood. There has also been con-
fusion of the generic nomenclature, and the family to
which they belong is still a moot question. The most
recent monograph of the genus is by Pirssry (1893).
Monographic treatment of the whole genus has not here
been attempted, since sufficient material was not avail-
able. Effort was concentrated on differentiating the species
on American shores. A synonymy is given of the species
of the eastern Atlantic, probably representing only one
biological species, and a list of the nominal species of the
western Pacific and Indian Oceans is appended, with
comments. Five species named by Dati (1919) from the
west coast and one from the east coast (DALL, 1927) were
never illustrated. Opportunity to study this problem at
the U.S. National Museum allowed me to make camera
lucida drawings of the type specimens of DaLt’s nominal
species. Measurements were made with an ocular micro-
meter, and are possibly more exact than those given by
Dau. Most of Datu’s type material was series of badly
worn specimens, from which he evidently made composite
descriptions. Lectotypes are here designated for each lot.

I am obliged to the Bureau of Commercial Fisheries
of the U.S. Fish and Wildlife Service, Galveston, and
especially to Messrs. Robert Temple, Clarence Fisher and
Charles Guice for the material from the western Gulf of

Mexico. The many courtesies of Dr. Harald Rehder, Dr.
J.P E.Morrison and Dr. Joseph Rosewater, while I
worked at the U.S. National Museum, are gratefully
acknowledged. This study was supported in part by
National Science Foundation Grant GB 2753.

This genus has sometimes been placed in a separate
family (Volvulidae, Locarp, 1892; Rhizoridae DELL,
1956). After an extensive study of the anatomy of Volvu-
lella persimilis from Brazil, Marcus & Marcus (1960)
concluded that tentatively this genus should be placed in
the Retusidae.

Volvulella Newton, 1891

Voluula A. ApaMs in SoweERBy’s Thesaurus Conchyliorum 2: 555.
Type by SD of A. Apams, 1862, Ann. and Mag. Nat. Hist.
Ser. 3, vol. 9, p. 154, V. acuminata (BrucuIERE) (= Bulla
acuminata BrucuterE, 1792). Not Volvula GisTEL, 1848,
Naturgesch. Thierr. f. Héhere Schulen, p. viii (Diptera).

Volvulella Newton, 1891, British Oligocene and Eocene Mollusca,
p. 268. Substitute name for Voluula A. Apams, 1850, hence
with same type species.

Rhizorus MontFort, 1810 of authors, not of MonTForrt.

ARTHUR ApAms (1850) did not designate a type from
among the 5 species which he included in Volvula, pro-
posed as a subgenus of Bulla, nor was there otherwise
intrinsic fixation of the type. MENKE (1854) declared
Voluula Apams to be a synonym of Rhizorus MONTFORT,
citing as type V. acuminata, which species he admitted he
did not have. Since the two generic concepts are not the
same, his designation of type can not be valid for Volvula
Apams. H. « A. Apams (1854) also denied the identity
of these two concepts. Their choice of the Australian
species, V. rostrata ADAMS (1850) as an “example” of the
genus was not construed by them or any subsequent

Page 134

author to be per se a designation of the type species, but it
may have led Bucguoy, DauTzENBERG & DotFus (1886)
to designate that species as type. In 1862 A. Apams
designated the first species of his monograph of 1850 as
type, the European V. acuminata BrucuizRE. PILsBRY
designated the same type in 1893, but neglected to indicate
the basis for his type selection. All subsequent authors
have erroneously adopted the type designation of Bucguoy
et al.

Newton (1891) renamed Apams’ concept, thinking
Voluula was a homonym of Volvulus Oxen, 1815. By
modern concepts of nomenclature these are not homo-
nyms, since they differ by gender as shown by the endings.
Therefore Pitspry (1893) rejected NEwTon’s substitute
name, but he overlooked the genus Volvula GisTEL, 1848,
which does invalidate Voluula Apams, 1850.

Several authors have continued to use Voluula A. ADAMS
for this group of snails (NickLks, 1950; Pruvot-Fot,
1954), although Grant & Gare (1931) gave the valid
argument for accepting Volvulella.

The genus Rhizorus MontFort, 1810 (Conch. Syst. 2:
339, and plate on opposite page) has been considered by
several authors during the nineteenth century as applying
to this group of snails, but most rejected it as representing
some other genus or as a result of their subjective nomen-
clatural conservatism (H. « A. ApaMs, 1854; JEFFREYS,
1867; Pitspry, 1893; Dati, 1889). The single species
which Montrort included in his genus, and designated as
type, Rhizorus adelaidis, was described as being as big
as a grain of millet. It came from the island of Elba,
Italy, but was thought probably also present in England.
He refers to a figure in SoLDANI, a work not available
to me, nor evidently to most writers who have studied

ACE TY

Lf

Uf

Figure 1

Tracing of figure of “Rhizorus adelaidis” from volume 1, page 338,
Montrort, 1810

THE VELIGER

Vol. 10; No. 2

the question. To show that Montrort’s genus can
scarcely apply to Volvulella, a tracing of his illustration of
R. adelaidis is here reproduced (Figure 1). It has the
apex broadly truncate, scarcely tapering, and widely
umbilicate; the aperture extends well above the apex,
and is not curved toward the columellar axis there. All
of these characters contrast with Volvulella, which, though
some variants may be spineless and minutely umbilicate,
have the aperture curved distinctly if only slightly toward
the shell axis. Moreover, there are much better candidates
for MonTForT’s concept, from the area he indicated, than
the single species of Volvulella which is generally recog-
nized from there. His concept may apply to something
like “C'ylichna conulus DesHayes” as described and
figured by Forpes & Haney (1851, 3: 517; 4: pl. 114c,
figure 7). Gray (1847) early associated MontTForT’s
genus with such forms.

Several major works of recent years have adopted
Rhizorus MontrFort for this genus (e.g. WINCKWORTH,
1932; Appott, 1954; PoweLt, 1962). Keen (1947) and
PaLMER (1958) have protested its use in this respect.

Snails of the typical subgenus have small shells (up to
9mm long) which are tapering, ovate or subcylindrical.
They are completely without intrinsic color, being light
grey and translucent when taken alive, but soon turning
snow white and opaque after “weathering” in nature.
The aperture is narrow, elongate, rounded basally and
extended the entire height of the shell. There is no colum-
ellar lamella or other apertural dentition. At its apical
end the aperture bends over the apex of the shell, and the
parietal wall here forms a spine which completely covers
the apex, so that only the final whorl is visible. This,
together with the non-determinate growth of the shell,
makes it difficult to recognize juvenile shells as such,
unless one has a large series available. The apical end of
the aperture is almost or entirely as far advanced in the
direction of growth as the rest of the outer lip. It is
not deeply sinuate, and consequently extends as high as
the tip of the spine in apertural view. There is a thin,
colorless periostracum which is soon destroyed after death.
Spiral sculpture of fine, incised lines, rather widely spaced,
occurs at both ends of the shell (macro-spirals) in most
species, and most (all?) have finer, closely set, wavy lines
incised over the middle part of the whorl (micro-spirals).
The micro-spirals are almost limited to the periostracum,
and rarely evident on the even slightly weathered speci-
mens. The shells are always imperforate basally, but a
slight clevation of the narrow columellar lip from the
whorl defines a furrow and minute pseudo-umbilicus.

Most characters show considerable variation within a
species. The apical extent of the aperture and its bending

Vol. 10; No. 2

toward the columellar axis may vary even within speci-
mens from one locality. This results in the spine varying
from long and tapering to short and blunt, or in extreme
cases, being replaced by a minute umbilicus at the apex.
In worn shells the spine seems to undergo similar degrada-
tion, even if it were long and acute in the living animal.
The spiral lines are variable in unworn shells of a species
from one locality, and weathering rapidly degrades them.
The basal lines are stronger and more persistent than the
apical ones, and both are generally more so than the
micro-spirals. The form may vary from oval to more
elongate, perhaps within the growth of a single individual.
But the shape of the shell seems to vary less than other
characters within a species, and therefore is the most use-
ful criterion at the species level.

Few species have special features which aid in identi-
fication, but examples are: the unusual thickness of the
shell in Volvulella paupercula and V. catharia; the trans-
verse riblets near the apex of V. recta; the tendency of the
shells of V. panamica and V. texasiana to be stained with
iron, and the peculiar apical sinus in the lip of these
species; the apical ridge in V. texasiana.

At least 4 distinct groups of species are living today.
The eastern Atlantic may have only one of the 4, but the
western Atlantic, eastern Pacific and Indo-Pacific areas
all contain very similar representatives of the other 3
species groups. Just how closely related these analogous
populations are from the standpoint of taxonomy can not
be definitely decided at present. The analogous species of
each region differ so little, and then chiefly in their form,
that a comparison of specimens from the several areas is
the only convincing demonstration of their distinctness. Al-
though only one species is generally recognized from the
eastern Atlantic (Bucguoy et al., 1882; Nick&s, 1950;
Pruvort-For, 1954), other species have occasionally been
recognized from there (DALL, 1889; Pitssry, 1893). The
following table summarizes the analogous species of the
Atlantic and eastern Pacific Oceans. As my study of the
populations of the Indian and western Pacific Occans is
based almost entirely on the literature, I have not included
them in this table, but appended a list of the nominal
species below.

Eastern Atlantic Western Atlantic
Subgenus Volvulella s. s.

Eastern Pacific

V. acuminata V. persimilis V. cylindrica
none? V, recta V. californica
none? V. paupercula V. catharia

Subgenus Paravolvulella, new subgenus

none? V. texasiana V. panamica

THE VELIGER

Page 135

Species of the Eastern Atlantic

Volvulella acuminata (Brucutére, 1792)

Bulla acuminata Brucutkre, 1792, Encycl. Méth. vol. 1, prt. 2, p.
376, no. 9. Not figured. Type locality: not specified; evidently
the Mediterranean, near northwestern Italy

Volvula cylindrica E. A. Smitu, 1872, Proc. Zool. Soc. London p.
738, plt. 75, fig. 29. Not V. cylindrica CarPENTER, 1864

Volvula smithit Pitspry, 1893, Trvon’s Man. Conch. 15: 233-234,
plt. 26, fig. 65. New name for V. cylindrica Smiru, 1871

Voluula acuminata var. brevis Pitspry, 1893, Tryon’s Man. Conch.

15: 235, plt. 60, fig. 11. Type locality: Northern Europe,
Mcditerranean

Volvula suavis TutELeE, 1925, Wiss. Ergeb. d. deutsch. Tiefsee Exp.
17 (2): 238, plt. 31, fig. 20. Type locality: 16° 26.5’S. Lat.,
11° 41.5’ E. Long. (off Angola, Africa). He compared the
single shell found with V. acuminata

There is variation in shape of this species, at least
among the several lots from northern Europe of the Jeff-
reys collection in the U.S. National Museum. Larger
specimens tend to be more elongate and more slender
than smaller ones, which are more oval. The spine is
elongate and acutely tapering. Completely spineless speci-
mens were not found among the fresh, non-worn shells.
Macrospiral lines occur at both ends, and faint micro-
spirals are evident in many fresh shells. The eastern
Atlantic shells always seem slightly more inflated than
Volvulella persimilis, although the difference is very small,
and requires actual comparison of specimens to demon-
strate it adequately.

The range of Volvulella acuminata extends from
Norway along the coast of Europe, including the Medi-
terrancan, and south along the African coast to Angola
(Nickiés, 1950). Pruvot-For (1954) lists it also from
the Gulf of Suez at the head of the Red Sea, and indicates
V. oxytata (BusH) and V. persimilis (M6rcH) may be
only varictics. She thought this species might be circum-
terrestrial, as did MEtvitt (1906).

Figure 2 (height 3.94mm, diameter 1.69mm) was
drawn from one of 4 specimens on a card (USNM
175142) noting one of these shells was drawn as figure 1
of plate 93 of Jerrreys’ “British Conchology.” The exact
locality of the specimens was not indicated, but it is
presumably the British Isles. Figure 3 (height 4.06mm,
diam. 1.25mm, USNM 175146) is one of a large lot
from St. Magnus Bay, Shetland Islands. It illustrates a
more cylindrical variant, with Icss acute spine. The speci-
men had been bored 3 times by some predaceous snail.

Page 136

Species of the Western Atlantic

Volvulella persimilis (M6rcu, 1875)

Volvula persimilis M6rcu, 1875, Malak. Blatter 22: 179. Not figured.
Type locality: ““M. Antil.” (Antilles). 1900, DauTzENBERG,
Méem. Soc. Zool. France 13: 155 - 156; plt. 9, fig. 10

Volvula oxytata Busu, 1885, Trans. Conn. Acad. Sci. 6 (2): 468;
pit. 45, fig. 12. Type locality: off Cape Hatteras, N. Car. in
7 to 17 fathoms

Volvulella mérchi Datu, 1927, Proc. U.S. Nat. Mus. 70: 22. Not
figured. Type locality: off Georgia, U.S. Fish. Comm. Sta.
2415, 440 fathoms

Volvula ischnatracta Pitspry, 1930. Proc. Acad. Nat. Sci. Philadel-
phia 82: 301; text fig. 1, p. 302. Type locality: Andros Bank,
west of Middle Bight, about 12 miles within the western edge
of the bank, in 34 fathoms (Bahamas)

This species is the western Atlantic counterpart of
Volvulella acuminata, from which it is differentiated only
by subtle differences of curvature of the whorl. The differ-
ence seems to be consistent when shells from the two sides
of the ocean are compared side by side. The sculpture
on both species is the same; prominent, broadly spaced
spiral lines at both ends of the shell, and finer spiral lines
between. To recognize V. persimilis as merely a subspecics
of V. acuminata might be equally acceptable. Volvulella
persimilis may not extend north of North Carolina, and
thus the ranges of the two species do not, as presently
known, meet in the North Atlantic. Numerous lots of V.
persimilis are present in the U, S. National Museum from
off Beaufort, North Carolina, and also from along the
Florida Keys (J.B. Henderson) as well as smaller lots
from Puerto Rico and Cuba. They range in depth from 15
to 209 fathoms. Marcus & Marcus (1960) described
the anatomy of specimens from Brazil. Shells from Beau-
fort are long spined forms. Figure 4 (USNM 35871,
height 3.50mm, diam. 1.25mm) is drawn from a spcci-
men collected by the U.S. Fish Commission, Station 2112,
in 15 fathoms off Cape Hatteras, North Carolina. It was
sent to Dall by Bush. Whereas it may not be a paratype, it
is at least an authentically identified specimen of what
Busu named Volvula oxytata, from the type locality. The
terminal spiral lines are present on this specimen, but finer
spiral lines on the middle of the shell were not seen,
possibly due to slight weathering.

In the more southern part of the range of this species,
there are specimens with shorter, blunted spines, and even
no spine but a small apical umbilicus. Such forms occur
in the same lots with acutely spined shells, but tend to
be more abundant than the latter. It is probably intrinsic
variation, rather than a common characteristic regularly
evoked by a special environment. It was on such speci-
mens that the species names Volvulella moerchi Dati and
Volvula ischnatracta Pitssry were based.

THE VELIGER

Vol. 10; No. 2

Mo6rcn’s original description of Voluvula persimilis was
based on a single specimen received from Krebs from the
Antilles. His name is applied to the present species in
part through a process of elimination, because his descrip-
tion best fits this species of those known to occur in the
western Atlantic, and because he adequately differentiated
it from the only other common shallow water species of
the area (V recta Morcw). The case for recognizing
Mo6rcn’s name is much better than that for recognizing
V. acuminata (BrucurrRE). The original description
[bracketed words inserted] of V. persimilis (M6rcH) and
my translation follow:

“Differt a Volvula angustata A. Av. (Thes. XV 1850 p. 596 n
121, f 153) t. spiraliter subtilissime striata sub lente vix detegen-
da, solidiore; columella valida obliqua, plica crassa recta. Differt
a praecedente [V. recta “p’Ors.” Morcu -= V. acuta v’Ors.] t.
subcylindrica. Long 44mm; diam. 1? mm. Hab. M. Antill. (Krebs)
spm. unicum.”

“Differs from Voluvula angustata A. ApaMS (SowERBY’s The-
saurus Conchyliorum vol. 15, 1850, p. 596, No. 21, fig. 153) in
that the shell is more solid, and spirally, very finely striate, the
striae scarcely detectable under a lens; columella strongly ob-
lique, with a coarse, straight lamella. It differs from the pre-
ceding [V. recta “p’OrB.” M6rcu = V. acuta p’Ors.] in that the
shell is subcylindrical. Length 44mm, diameter 13mm. Habitat

M. [major? minor?] Antilles, a single specimen received from
Krebs.”

The figure of Volvulella angustata (A. ApAMs) (from
the Philippines) referred to fits very well the brevi-spinous
forms of this species in the West Indies. The aperture
extends to the tip of the spine, thus differentiating it
from V. texasiana, and the cylindrical form differentiates
it from the bulbous V. recta and V. paupercula.

DauTzENBERG (1900) redescribed and figured Volvul-
ella persimilis from shells obtained at 10 fathoms off
Venczuela. Although he argues that this species is distinct
from V. oxytata (Bus), the described and figured shells
are quite within the range of variation of the species as
here understood. Marcus & Marcus (1960) also thought
these two names are synonyms.

Figure 5 is a drawing of the single specimen on which
Volvulella moerchi Daux, 1927 was based (USNM 108268,
U.S. Fish Commission, Station 2415). The shell is very
worn, with the spine completcly broken off and the outer
lip badly damaged. It is chalky and opaque, showing no
sculpture: height 3.68 mm, diam. 1.55mm (Da tz stated:
Height 3.5; diameter 1.25mm.). This specimen may have
been accidentally present in the 440 fathom station where
it was taken. Dati made no mention of depth in his
description, but “440 fms.” is on the label. He noted it
“Gs nearest V. persimilis MOrcu but is smaller and not
spirally striated.”

Vol. 10; No. 2

THE VELIGER

Page 137

Figure 2

Figure 3

Figure 5

Figure 4

Figure 2: Volvulella acuminata (BRUGUIERE). 3.94 mm high
Figure 3: Volvulella acuminata (BRucuIiRE). England. 4.06 mm high
Figure 4: Volvulella persimilis (MOrcH) (authentic specimen of
Volvulella oxytata (BusH) ; 3.50mm high)

Figure 5: Holotype of Volvulella moerchi Datu (= V. persimilis (MO6rcH) )
3.68 mm high

Pitspry (1930) may have had only a single specimen
available to describe Volvulella ischnatracta. It is com-
pletely spineless (broken?), but has an elongate form
characteristic of this species, and spiral lines at both ends.

Volvulella recta (MOrcn, 1875)

Bulla acuta p’Orpicny, 1842. Mollusca, in Sacra, Hist. . . . Cuba
1: 126; plt. 4, figs. 17 - 20. Type locality not specified; cited
from Cuba, Jamaica, Guadaloupe and Martinique and
“toutes les Antilles.’ Non Bulla acuta GraTELouP, 1828,
Bull. Hist. Nat. Soc. Linn. Bordeaux 2 (9): 87

Volvula recta “pD’OrBicNy” Morcu, 1875. Malak. Blatter 22: 179.
Not figured. This fortuitous lapsus calami merely cites the
page and illustration in p’Orpicny, 1842 of Bulla acuta
p’Orsicny, but meets all requirements for rectifying p’Or-
BIGNY’s homonym

Voluula minuta Busu, 1885. Trans. Conn. Acad. Sci. 6: 469; plt. 45,
fig. 11. Type locality: off Cape Hatteras, N. Car., in 14 to
16 fathoms

Volvula bushii Dati, 1889. Bull. Mus. Comp. Zool. 18: 51. Not
figured. Type locality: Station 2602, 36 miles S. 4W. from
Cape Hatteras, N.C., in 124 fathoms, sand. 1925, Dati,
Proc. U.S. Nat. Mus. 66: 31; plt. 25, fig. 3

This, the first species in the genus to be described from
the New World, had one of the most succinct descriptions
and better illustrations of any from that area. The follow-
ing is a free translation of p’OrBIGNyY’s description in
Trench.

“The shell is oblong, bright, thin, swollen in the middle, thin-
ning at the extremities, marked in front by several spiral striations
and behind by several others, a little in front of the extremity,
which is transversely striate. Spire entirely enclosed, without
umbilicus, replaced by a long, sharp prolongation. Aperture very
straight, a little sinuate, prolonged behind, enlarged in front;
columella a little projecting, separated by a groove from the um-
bilicus and forming a slight ridge. Color, uniformly white. Length
2mm. Diam. 2 mm.”

He specifically called attention in both the Latin and
French descriptions and in the discussion to the delicate,
transverse, apical striae which are unique to this species.
But he referred to them as longitudinal, using the term
transverse for what is here termed spiral. The transverse
striae are around the apical third of the shell and extend
onto the base of the spine. They are very closely spaced,

Page 138

THE VELIGER

Vol. 10; No. 2

regular, and somewhat stronger than growth lines. The
greatest diameter of this species is above the midpoint of
the length of the shell, in contrast to all others of the East
Coast. The spine may be short and blunt, but the shell is
never as solid in structure as that of Volvulella paupercula.

The U.S. National Museum has specimens from Cape
Hatteras, North Carolina, and numerous ones from the
Florida Keys. It extends into the Gulf of Mexico at least

Figure 6

to Mobile, Alabama. These lots range in depth from 12
to 124 fathoms.

Figure 11 (USNM 44773, U.S. Fish Commission Sta-
tion 2113, 15 fathoms, off Cape Hatteras, North Caro-
lina) is drawn from a single shell of Voluula minuta Busu
which she sent to DALL. It is scarcely worn, and agrees
closely with Volvulella recta, both in the shape and in
having transverse striae at the apex. Spiral striae are

Figure 7

Figure 9

Figure 8

Figure 6: Volvulclla cylindrica (CARPENTER). 6.19mm high
Figure 7: Lectotype of Volvulella coopert DALL
(= V. cylindrica (CarPENTER) ). 6.45mm_ high
Figure 8: Lectotype of Volvulella callicera DALL
(= V. cylindrica (CarrENnTER) ). 3.06mm high
Figure 9: Volvulella cylindrica (CARPENTER). 7.88mm high

Vol. 10; No. 2

Figure 11

Figure 10

THE VELIGER

Page 139

Figure 12

Figure 10: Lectotype of Volvula bushii Datu (= Volvulella recta (MOrcu)).
4.31 mm high

Figure 11: Volvulella minuta (BusH) (= V. recta (MOrcu) ).

2.02mm high

Figure 12: Lectotype of Volvulella californica DaLL. 4.44mm high

strong at the apical end, less pronounced basally, and none
could be detected on the midregion of the shell. It may
be juvenile. Height 2.02mm, diameter 0.92 mm.

Figure 10 (USNM 95301, off Cape Hatteras, North
Carolina) is drawn from the larger of two specimens
which are presumably cotypes of Volvula bushit Dat,
1889. It agrees most closely with the dimensions cited in
the original description (Dati, 1889) “Lon. 4.6; Lat. 2.3
mm,” and is here designated lectotype. He originally
_cited 6 specimens; there are no locality data with this lot,
but the catalogue number is that which he cited in figuring
the species (Dat, 1925).

Measurements of the two shells are:

Height Diameter
Lectotype: 4.31 2.06
Paratype: 3/40) 1.56

Both shells are opaque, white, and slightly worn. They
are quite characteristic of Volvulella recta in shape, having
a very long, acute spine, and transverse striae at the apical
end. Spiral striae are present at both ends, and micro-
spirals are evident in the middle of the shell.

Volvulella paupercula (Watson, 1883)

Cylichna (Volvula) paupercula Watson, 1883. Journ. Linn. Soc.
London 17: 325. Not figured. Type locality: Lat. 18° 38’ 30”
N., Long. 65° 05’ 30” W, North of Culebra Island, St. Tho-
mas, West Indies. 390 fathoms. 1886, Watson, Challenger
Report, Zoology, 15: 669-670; plt. 50, fig. 5

Volvula aspinosa Datu, 1889. Bull. Mus. Comp. Zool. 18: 51. Not
figured. Type locality: “Off the North Carolina Coast in 18
to 168 fms., sand Straits of Florida, 150-200 fms.”
1925, Dati, Proc. U.S. Nat. Mus. 66: 31; plt. 25, fig. 5

This specics is somewhat more inflated than Volvulella
recta. Larger specimens have the greatest diameter at the
midpoint of the length, rather than above it. Transverse
striae at the apical end are absent, although more widely
spaced growth lines extending from apex to base are
gencrally present. Coarse spiral striae are prominent at
both ends, and fine spiral lines are often well shown in
fresh shells. The spine is small, but usually absent. In the
latter case the aperture docs not quite reach the columella,
although it bends strongly toward it at the apical end. A
distinct but minute apical umbilicus is present in the

Page 140

aspinous specimens. The shell is unusually thick for the
genus. This may be a species which is reverting to a
spineless condition. The general shape and sculpture, and
presence of a distinct, short spine in some specimens,
suggest it is truly a Volvulella rather than a Cylichna.

The species is well represented in the collection of the
U.S. National Museum by material dredged by J. B. Hen-
derson in 1917, on the yacht Folis, along the Florida Keys
and off Barbados. The range in depth is from 75 to 190
fathoms.

Figure 15 (USNM 95305, off Cape Hatteras, North
Carolina) was drawn from the larger of two cotypes of
Volvula aspinosa Dati, and is here designated the
lectotype.

Figure 13

Figure 15

THE VELIGER

Vol. 10; No. 2

Measurements of the two shells are:

Height Diameter
Lectotype: 3.63 2.06
Paratype: 3.06 1.75

Dati (1925) cited USNM 95302 as the type lot, but
I think that was a misprint. The type locality should be
restricted to North Carolina. He gave the dimensions as
“Long. 4.0; Lat. 2.0 mm.” (Dat, 1889).

Figures 16 (height 3.63mm, diameter 1.94mm) and
17 (height 2.94mm, diameter 1.50mm) are drawn from
specimens (USNM 500381) from 94 fathoms off Barba-
dos, and show somewhat slimmer forms of this species.

Although Volvulella paupercula (WATSON) was de-
scribed from shells taken at 390 fathoms, and thus much

Figure 14

Figure 16

Figure 17

Figure 13: Volvulella paupercula (Watson). Tracing of figure 5 of plate 50,
Figure 14: Lectotype of Volvulella catharia DaLL. 2.75mm high
Figure 15: Lectotype of Voluvula aspinosa DaLL
(= Volvulella paupercula (Watson) ). 3.63mm_ high
Figure 16: Volvulella paupercula (Watson). Barbados. 3.63 mm high
Figure 17: Volvulella paupercula (Watson). Same lot as Figure 16. 2.94mm high

Vol. 10; No. 2

deeper than any examined in the present study, the
excellent description and detailed figure of it leaves little
doubt that this is what Datt later named Volvula aspinosa.
Watson’s figure has the sculpture more strongly indicated
than it actually is. A tracing of his figure (without the
sculpture) is here reproduced in Figure 13. The dimen-
sions given by WATSON, converted from inches, are: height
1.55 mm, diameter 0.75 mm.

(Paravolvulella) Harry, subgen. nov.

Type: Volvulella (Paravolvulella) texasiana Harry, spec.
nov.

This subgenus differs from the typical one in the more
cylindrical form of the shell, with bluntly rounded rather
than gently tapering shoulder; and especially in the deep,
rounded sinus of the apical end of the outer lip, which
appears in apertural view not to extend farther posteriorly
than the base of the spine, rather than to its tip. However,
the parietal lip does curve upward medial to the sinus to
form the spine, which is short but acute.

There is a tendency for the species of this subgenus to
be colored brown by iron stain (extrinsic coloration) in
fresh specimens. This is unusual in marine mollusks. At
Galveston, only a few of the off-shore species exhibit this
phenomenon, whereas many others living under the same
conditions are free of it.

Volvulella (Paravolvulella) texasiana Harry, spec. nov.
(Figure 21)

Shell small, cylindrical, translucent, grey, flecked irreg-
ularly with opaque, white areas (growth interruptions) ,
the ends washed with reddish brown (iron stain). Aper-
ture as long as the single visible whorl, but exceeded
slightly by the tip of the spine. Side of whorl and aperture
flattened, or in larger shells very slightly constricted by a
vague, broad, medial sulcus. The diameter of the shell is
the same at both ends in larger shells, but in very young
shells the outline is inflated, suboval, with the greater
diameter in the middle. Apical and basal ends gently
curved. The spine is short, though usually broken, with a
minute, bilobed appearance when seen in profile, and a
minute, deep pit in the spine when seen from above. There
is a narrow spiral band of roughened shell material
around the base of the spine, its outer edge slightly above
the shell surface. This tapers out along the upper, truncate
part of the outer lip. Outer lip roundly quadrate above,
its inner end meeting the columellar axis at a right angle;
it is thin, acute throughout its length. The edge of the
outer lip forward of the posterior sinus is almost in one

THE VELIGER

Page 141

plane, but slightly more advanced in the direction of
growth toward the base. Basal part of the lip strongly
arched, not as advanced in the direction of growth as the
outer. Columellar lip short, narrow, thickened, adnate, and
without a lamella. No umbilicus, nor indentation in the
umbilical area.

There are a few minute, closely and equally spaced
spiral lines on the base, and similar, less prominent ones
near the apical spine. Much fainter, wavy ones cover the
middle part of the shell. These can only be seen at higher
magnification.

Holotype: The holotype is USNM molluscan collection
No. 678000, Height: 3.94mm; Diameter 0.75 mm.
Type locality: Southeast of the base of the Bolivar Pen-
insula (east of Galveston, Texas), about 10 miles offshore,
at 7 fathoms, Long. 94° 23’ W, Lat. 29° 22’N. Two speci-
mens were dredged there in December, 1965, and 4 more
at the same place in January 1966, but no shells of this
species were found in dredgings I examined from 54
other stations west and south of there, ranging from 3 to
18 fathoms. However, Mr. Charles Guice kindly provided
specimens from three other stations (among 20 he exam-
ined) in the area. These yielded only one or two live
animals per station. Two stations were at 7, and one at 9
to 10 fathoms. The species is evidently sparse and local.

Volvulella texasiana differs from V. panamica in its
smaller size, in the slight medial constriction, and particu-
larly in having a ridge around the spine. I could not find
such a ridge in any specimens of V. panamica in the U. S.
National Museum.

Species from the Eastern Pacific

The species of the west coast of North America have
been most recently reviewed by J. Q. BurcH (1947), with
many locality and depth records. He gave two keys, taken
from the notebook of the late Dr. Strong, which rely
chiefly on presence of spiral sculpture and height-diameter
ratios of the shell.

Volvulella cylindrica (CARPENTER, 1864)

Volvula cylindrica CARPENTER, 1864. Reprt. Brit. Assoc. Ady. Sci.
1863, p. 537 and 647. Not figured. Type locality: Santa Bar-
bara, California. 1958, Parmer, Geol. Soc. Amer. Memoir
76: 240; plt. 25, figs. 1, 2 (holotype, Redpath Museum, No.
2364; extensive references)

Volvulella cooperi Dati, 1919. Proc. U.S. Nat. Mus. 56: 297 - 298.
Not figured. Type locality: Scammon Lagoon, Lower Califor-
nia

Volvulella callicera Dati, 1919. Proc. U.S. Nat. Mus. 56: 299. Not
figured. Type locality: U.S. Fish Comm. Sta. 2813, off Gala-
pagos Islands, in 40 fathoms, coral sand

Page 142

Volvulella lowci Srronc « Hertiein, 1937. Proc. Calif. Acad. Sci.
ser. 4, 22: 164-165; plt. 35, fig. 2. Type locality: Puerto
Escondido, Gulf of California. 1939, Srronc « HERTLEIN,
Allan Hancock Pacific Exped. 2 (12): 190; plt. 18, fig. 1
(specimen from Panama)

This is an elongate specics closely related to Voluvulella
acuminata and V. persimilis, but it grows larger than
either, and the sides are somewhat more flattened than in
those species. The apical end of the aperture is not sinuate
as in V. panamica, and it extends to the top of the spine
in the same plane as the latter. The apical end of the
shell tapers more gently than in that species. Broadly
spaced spiral striae are present at both ends, those at
the apical end being less prominent and more easily
worn off. Finer, more closely spaced spiral lines are present
on the midpart of the whorl of at least some non-worn
specimens.

The spine is variable in length, but tends to be short
with a broad base. Figure 6 (USNM 212655, largest of
4 specimens; height 6.19mm, diameter 2.00mm) was
dredged from 5} fathoms off Lower California, and has
a more tapering spine than the holotype figured by
Parmer (1958). The range of this species is generally
cited as Vancouver Island to the Gulf of California

THE VELIGER

Vol. 10; No. 2

(Parmer, 1958), but the U.S. National Museum has a
lot of 2 shells (USNM 509036) from Panama, collected
by Zetek, which are quite typical of this species.

Although Grant & GALE (1931) and others have recog-
nized Volvulella cooperi DAL. as distinct, I think it is
only an aspinous variant of this species. Figure 7 (USNM
105501, Scammon Lagoon, Lower California) was drawn
from the next to smallest of 4 cotypes of V. coopert DALL.
All 4 are badly worn. Weak spiral lines are present on
the bases of 2, but none could be found elsewhere. DALL
evidently gave the measurements of the largest shell (9.5
mm long, diameter 3.6mm) but both larger specimens
are so badly worn as to make them poor choices for a
type specimen. The one illustrated (Figure 7) is here
selected as lectotype. It is only slightly worn, but the
lip is badly broken.

Measurements of the cotypes are:

Height Diameter
Paratype: 9.68 3.61
Paratype: 7.61 2.58
Lectotype: 6.45 2232
Paratype: S28) 2.06

Figure 8 (USNM 194176b) is drawn from the smallest
of 3 cotypes (here designated lectotype) of Volvulella

Figure 18 Figure 19 Figure 20 Figure 21
Figure 18: Lectotype of Volvulella panamica Dau. 4.25mm high
Figure 19: Side view of the same shell as in Figure 18.
Figure 20: Paratype of Volvulella tenuissima WILLETT. 4.5mm high

Figure 21: Holotype of Volvulella texasiana Harry, spec. nov. 3.94mm high

Vol. 10; No. 2

callicera Dau. All are bleached, with sand grains lodged
in the aperture and the upper part of the lip slightly
broken. But the sculpture is evident and quite typical:
broadly spaced spiral lines at both ends, with finer, closely
spaced lines in the middle. Dati (1919) gave the meas-
urements: length 3.5mm, diameter 1 mm.

My measurements of largest and smallest cotypes are:

Height Diameter
Paratype: 4.31 1.38
Lectotype: 3.06 0.94

Figure 9 (USNM 267574, Head of Conception Bay,
Gulf of California) is drawn from a specimen set aside
as “figured type” of a species to be named by Bartscu.
I can not find that he ever published it, although a
manuscript name was present on several lots in the U.S.
National Museum. They are a mixture of Volvulella cylin-
drica and V. panamica, all from the Gulf of California.
The dimensions of the specimen drawn are: height 7.88
mm, diameter 1.69 mm.

I have seen no authentic specimens of Voluulella lower
Strone & HeErTLEIN, but judging from the figures and
description, it is a poorly spined form of V. cylindrica.

Volvulella californica Dati, 1919

Volvulella californica Dau, 1919. Proc. U.S. Nat. Mus. 56: 299 -300.
Not figured. Type locality: off Santa Rosa Island, California,
in 53 fathoms, sandy mud. 1927, Otproyp, Marine shells
west coast, 2 (1): 34; pl. 2, fig. 11 [poor]

Figure 12 (USNM 211303) is drawn from the larger
(here designated lectotype) of 2 complete cotypes. A
third cotype has the body whorl badly broken. The speci-
mens are slightly weathered, so that there is no trace of
finer spiral lines which may be present on unworn speci-
mens. About 8 widely spaced spiral lines are present on
the base of the lectotype, which shows none near the
apex. Apical spirals are present in the complete paratype,
however. The profile of the shell is broadly arched, with
the greatest diameter in the middle. The spine is of
moderate length. This may be the West Coast counterpart
of Volvulella recta, but it lacks any trace of transverse
striae near the apex. It is distinctly more cylindrical than
that species. The only specimens I found in the U.S.
National Museum which could certainly be referred to
this species were those of the type series. Possibly this is
only a growth stage of V. cylindrica. However, BurcH
(1947) cites it from 6 stations, between Santa Cruz,
California, and Todos Santos Bay, Lower California,
ranging in depth from 30 to 298 fathoms. Measurements
of 2 non-broken cotypes (cf. Dati, 1919, who stated
“length 4; diameter 1.7mm’) are:

THE VELIGER

Page 143
Height Diameter
Lectotype: 4,44 2.00
Paratype: 3.63 1.56

Volvulella catharia Dauu, 1919

Volvulella catharia Dax, 1919. Proc. U.S. Nat. Mus. 56: 298. Not
figured. Type locality: U.S. Fish. Comm. Sta. 2794 in Pana-
ma Bay, in 62 fathoms, sand

Figure 14 (USNM 211784) is a drawing of the slightly
more globose of the 2 non-broken of 4 cotypes, all of
which are about the same size and slightly worn. It is here
designated lectotype. It measures: height 2.75 mm, di-
ameter 1.44mm (Da tt, 1919, stated: “length of shell,
2.75; diameter, 1.8mm’). This is the West Coast counter-
part of Volvulella paupercula (Watson), and the limited
series at the U.S. National Museum (only one other lot,
194976a, Galapagos Islands, 40 fathoms) leaves room for
doubt about whether it is distinct from that species. It
may not grow as large as the eastern one. A few widely
spaced spiral lines are present on the base, but none
could be seen on the apex, nor over the midregion of the
shell (worn off?). It is very thick shelled, like its eastern
relative, and there is no spine, but a minute, rimate
umbilicus at the apex.

Volvulella (Paravolvulella) panamica Datt, 1919

Volvulella panamica Datt, 1919. Proc. U.S. Nat. Mus. 56: 298. Not
figured. Type locality: Panama Bay at Sta. 2799, in 294
fathoms. 1937, Srronc « HERTLEIN, Proc. Calif. Acad. Sci.
ser. 4, 22: 164; plt. 35, fig. 3

Volvulella tenuissima WiLLETT, 1944. Bull. So. Calif. Acad. Sci. 43:
71-72; pit. 4. fig. 1. Type locality: Off Redondo, California,
in 75 fathoms

Figures 18 and 19 are drawn from the single shell in
a separate vial of USNM 212654, height 4.25mm, di-
ameter 1.56mm (Dati stated: “length 4.25, diameter
1.75mm’’) which is here designated lectotype of Volvul-
ella panamica Dat. Another vial with the same catalogue
number contains a large series of specimens, all slightly
weathered. The lectotype has traces of strong spiral lines
at both ends, and finer, closely spaced ones over the
middle. This is the most distinctly cylindrical of the West
Coast species, with the apical end abruptly rounded, the
spine short and blunt, and the apical end of the lip deeply
sinuate by not being as advanced in the direction of
growth as the rest of the lip. The apical part of the lip
appears truncate, not extending to the tip of the spine,
although the parietal wall actually forms the latter
(Figure 19).

Page 144

Figure 20 is drawn from the larger of 2 perfect (of 3)
paratypes of Volvulella tenuissima Wiwietr (USNM
573516, height 4.50mm, diameter 1.63mm). The speci-
mens were evidently collected alive, and show pronounced
sculpture, with brown iron stain on the ends. The spiral
lines on the ends are very closely spaced, but I suspect
this varies within the species. It is notable that WILLETT
(1944) compared his new species only with V. cylindrica
and did not mention V. panamica. This species ranges
from off Redondo, California, to Panama, and is abund-
dantly represented in the U. S. National Museum from the
Gulf of California. There is no apical spiral ridge in
any of the numerous specimens seen of V. panamica, such
as that present in V. texasiana.

Species of the Indian
and Western Pacific Oceans

Listed below in alphabetical order are all trivial names
I have found of living species of this genus in the western
Pacific and Indian Oceans. This includes 3 which have
been attributed to the genus, but probably belong else-
where, and one which was described as Cylichna which
probably is a Volvulella. Further studies on this genus in
the Indo-Pacific area should review all those described
as Cylichna, for possibly others originally so allocated
belong here. Populations very similar to V. acuminata are
widely distributed in these oceans. Whether they are
specifically distinct from that species is a moot question.
But one of them, V. pia, was described from the southern
tip of Africa and might as well be relegated to the Atlantic
as to the Indo-Pacific fauna. The other 3 species groups
of American shores are found in these oceans also. I
have made a guess regarding the Atlantic analogue of
each nominal species listed below. The species described
by A. Apams in 1862 are particularly difficult to recognize,
being described in a few lines of Latin, not figured, and
with no measurements. Some authors have named new
species if they found them at a distance, or of an age,
remote from any species previously described. Insufficient
specimens from the Indo-Pacific have prohibited decisions
on the validity of these, but I have pointed out such
opinions on synonymy as have been found.

acutaeformis YOKOYAMA, 1922 (Volvula), p. 26, pl. 1,
fig. 9. Type locality: fossil, from Shito, Japan (of V.
recta group; he compared it to V. minuta BusH). Kuropa
& Hase (1952) attributed it to the recent fauna of Japan,
and Hase (1954) declared it a synonym of V. ovulina
ADAMS.

angustata A. Apams, 1850 (Bulla (Volvula)), p. 596,
pl. 125, fig. 153. Type locality: Cagayan, Mindanao (Phil-
ippines), 25 fathoms (of V. acuminata group).

THE VELIGER

Vol. 10; No. 2

aomoriensis Nomura, 1939 (Rhizorus), p. 26, pl. 2,
figs. 9a, 9b. Type locality: Japan, numerous localities
cited, 30 fathoms (of V. acuminata group). HaBe (1955)
called this a synonym of V. radiola ADAMs.

artiaperta YAMAKAWA, 1911 (Volvula), p. 50, pl. 11,
figs. 33 - 36. Type locality: fossil, Japan [original not seen].
Kuropa & Habe, 1952, cited this living, and Hane, 1955,
said it is a synonym of V. eburnea ADAMS.

attenuata A. ApAms, 1862 (Volvula), p. 155, not
figured. Type locality: Tsu-Sima (Japan), 26 fathoms.
(Probably of V. recta group). !

compacta MELVILL, 1906 (Volvula), p. 79, pl. 7, fig. 26.
Type locality: not indicated, evidently Persian Gulf or
Gulf of Oman. No depth given (of V. paupercula group).

cylindrella A. ApaMs, 1862 (Volvula), p. 155, not
figured. Type locality: Mino-Sima (Japan), 63 fathoms
(of V. acuminata group).

eburnea A. Apams, 1850 (Bulla (Voluula)), p. 597,
pl. 125, fig. 155. Type locality: China Sea. Kuropa &
Hase (1954, pl. 2, fig. 11) provided a photograph of a
specimen supposedly identified by ADAMS, now in the
Redpath Museum (probably of V. recta group).

exilis THIELE, 1925 (Volvula), p. 316, pl. 31, fig. 25.
Type locality: Padang, Sumatra (a poorly spined specimen
of the V. acuminata group).

flavotincta Martens, 1903 (Volvula), pp. 130-131, pl.
5, fig. 21. Type locality: East Africa, (Valdivia) station
270, in Gulf of Aden, 1840m deep (possibly of the V.
paupercula group).

fortis Turere, 1925 (Volvula), p. 238, pl. 31, fig. 23.
Type locality: 0° 30.2’N Lat., 97°59.7’E Long., 132m
depth, “Nias-Siid Kanal” (South of Sumatra) (probably
of V. recta group; it has transverse ridges at the base of
the spine).

granulum Primirrt, 1851 (Bulla), pp. 63-64; not figured.
Type locality: China. This is probably the “granulum
Puiuippr’ listed under Voluula by H. & A. ADAMs (1854:
14). I agree with Pirspry (1893: 309), who placed it
in Cylichna.

kinokuniana Hae, 1946 (Rhizorus), p. 186, not figured.
The description is in Japanese. KuropA & HaseE (1954, pp.
8-9, pl. 2, fig. 15) illustrated a shell in the Redpath
Museum, supposedly received from A. ADAMs, under the
unpublished name Voluula solidula A. Apams. They de-
clared this is Volvuella [sic] kinokuniana Hase and note
the species is from “deep bottoms of Tosa Bay, Shikoku,
and also off Wakayama Pref., Honshu (Japan)” (of V.
paupercula group).

lenis TuteLe, 1925 (Volvula), p. 238, pl. 31, fig. 22.
Type locality: 0° 39.2’S Lat., 98°52.3’E Long., 750m

Vol. 10; No. 2

THE VELIGER

Page 145

depth, near Siberoet Island (South of Sumatra) (of V.
paupercula group).

mecyntea ME vit, 1912 (Cylichna), p. 253, pl. 11,
fig. 15. Type locality: Persian Gulf, Ormara. The single
specimen was spineless, but seems to be a Volvulella of
the V. texasiana group.

minutissima THIELE, 1925 (Volvula), p. 316, pl. 31,
fig. 24. Type locality: (J.c., p. 254) : Padang, Sumatra (of
V. recta group; it has transverse ridges at the base of the
spine).

mucronata Puruipri, 1849 (Bulla), p. 22, not figured.
Type locality: Red Sea at Aden. This is probably the
mucronata PuitipPi listed by H. & A. ApAMs (1854: 14)
under Volvula, and cited by A. ApaMs (1862). I agree
with Pitssry (1893: 191), who placed it in Tornatina.

nesentus FINLAY, 1926 (Rhizorus), p. 438; new name
for Voluulella reflexa SutTER, 1913 (p. 529, pl. 23, fig. 2;
Recent of New Zealand, non Cylichna refleca Hutton,
1886, fossil, Miocene of New Zealand).

opalina A. ApaAms, 1862 (Volvula), p. 154, not figured.
Type locality: Mino-Sima, 63 fathoms (Japan) (possibly
of V. acuminata group).

ovulina A. Apams, 1862 (Volvula), p. 155, not figured.
Type locality: Mino-Sima, 63 fathoms (Japan) (probably
of V. recta group).

parata IREDALE, 1936 (Volvulella), p. 332. New name
for a figured specimen Hepiey (1903: 394, fig. 109) had
identified as V. rostrata A. ApaMs. Det (1956) argued
convincingly that this is merely a synonym of V. rostrata.

pia Tuiere, 1925 (Volvula), p. 238, pl. 31, fig. 21.
Type locality: 34°51’S Lat., 19° 37.8’ E Long., 80m,
near Cape Agulhas (Southern tip of Africa) (similar to
V. acuminata, of which it may be a synonym).

radiola A. ApaMs, 1862 (Volvula), p. 155, not figured.
Type locality: Tabu-Sima, 25 fathoms (Japan). The de-
scription suggests it belongs to the V. acuminata group, but
Kuropa & Habe (1954, pl. 2, figs. 3 and 4) published a
photograph of a shell in the Redpath Museum supposedly
from A. Apams, which resembles the V. texasiana group.

rostrata A. ADAMS, 1850 (Bulla (Volvula)), p. 596, pl.
125, fig. 154. Type locality: Port Lincoln (Australia)
(probably of V. recta group).

spectabilis A. ADAMS, 1862 (Volvula) p. 154, not fig-
ured. Type locality: Tabu-Sima, 25 fathoms (Japan)
(probably V. acuminata group).

striatula A. Apams, 1850 (Bulla (Volvula)), p. 597, pl.

125, fig. 156. Type locality: China Sea (possibly of V.
texasiana group).

sulcata Watson, 1883 (Cylichna (Volvula)), p. 326,
not figured. Type locality: Torres Strait, North of Aus-
tralia. Watson, 1886, p. 670, pl. 50, fig. 6 (of V. recta
group).

tokunagai MixtyaMA, 1927 (Volvulella acuminata sub-
spec.), pp. 144-145, not figured. New name for the Japa-
nese Pleistocene fossil Cylichna acuta “D’OrB.” Toxu-
naGA, 1906, Journ. Coll. Sci. Univ. Tokyo 21 (2): 32, pl.
2, fig. 13, not Bulla acuta p’OrBicNy. Nomura (1939:
26) elevated this subspecies to specific rank, and cited it
living at several localities in Japan. Nomura’s figures
(J. c., pl. 2, figs. 8a, 8b) show it belongs to the group of
V. recta.

tragula Hepiey, 1903 (Volvula), p. 395, text fig. 110.
Type locality: 63-75 fathoms, off Port Kembla, Australia
(resembles V. recta; there are coarse spiral ridges at the
base of the spine shown in the figure but not mentioned
in the text).

truncata DELL, 1956 (Volvulella), p. 146, pl. 20, figs.
211, 212. Type locality: 260 fathoms, Chatham Islands,
New Zealand (possibly of V. texasiana group).

SUMMARY

Volvulella, a genus of minute snails, is world wide in
distribution in the tropics and warm temperate waters,
ranging from a few to many meters depth. They are rare
and local, with few distinctive characters to distinguish
species. Four species groups are recognized, with analogous
species in the various major oceans. Possibly only one
species group occurs in the eastern Atlantic, but all 4
groups are represented in the western Atlantic and eastern
Pacific, and in the Indo-Pacific area. Volvulella NEwTon,
1891, is accepted as the correct name of this genus, of
which Bulla acuminata BrucuiERE is the type species by
designation of A. ApamMs, 1862. Paravolvulella, new sub-
genus is established with V. (RP) texasiana, spec. nov.,
designated as type. Six unfigured type specimens of nom-
inal species of Dati from the east and west coast of
North America are illustrated. A catalogue of 30 nominal
species from the Indo-Pacific area indicates to which of
the 4 recognized species groups each may belong.

LITERATURE CITED

Axspott, ROBERT TUCKER

1954. | American seashells. Princeton, New Jersey. D. van
Nostrand Co., Inc.; xiv + 541 pp.; 100 figs.; 40 plts.

Page 146

THE VELIGER

Vol. 10; No. 2

ApamMs, ARTHUR
1850. Monograph of the family Bullidae in G. B. Sowersy,
Thesaurus Conch. 2: 553 - 608; plts. 119 - 125
1862. On some new spccies of Cylichnidae, Bullidae and
Phylinidae from the seas of China and Japan. Ann. Mag.
Nat. Hist. ser. 3, 9: 150 - 161
Apams, Henry, « ARTHUR ADAMS
1853-1858. The genera of Recent Mollusca, arranged according
to their organization. | London; 1: vi-xl, 1-484; 2: 1-661: 3:
pits. 1 - 138
(pertinent data are on page 14 of vol. 2, published 1854)
BRUGUIERE, JEAN-GUILLAUME
1789-1797. Histoire naturelle des vers.
1(1): 1-344; 1(2): 345-757. Paris.
(relevant data appeared in 1892)
Bucguoy, Eucine, PHitippE DAUTZENBERG & GUSTAVE FREDERIC
Do.irus
1882-1898. Les mollusques marins du Roussillon.
Balliére et Fils; 2 vols.
(fasc. 13, containing Volvula, appeared 1886)

Encycl. Méthod.

Paris, J. B.

Burcu, JoHN Quincy
1945. (Review of West Coast Volvulella) Minutes Conch. Club
So. Calif. 47: 17 - 20; plt. 2, figs. 17 - 19 (mimeographed)
Busu, KATHERINE JEANNETTE
1885. Additions to the shallow water mollusca of Cape Hat-
teras, N.C. dredged by the U.S. Fish Commission steamer
“Albatross” in 1883 and 1884. Trans. Connecticut Acad.
Sci. 6: 453 - 480; plt. 45
CARPENTER, PHILIP PEARSALL
1864. | Supplementary report on the present state of our know-
ledge with regard to the Mollusca of the west coast of North
America. Rept. Brit. Assoc. Adv. Sci. for 1863: 517 - 686
(August 1864)
(not seen; relevant material quoted in PaLMER, 1958)
Dati, Witti1am HEALEY
1889. __ Reports on the results of dredging
Mexico and in the Caribbean Sea by the U.S. Coast
survey steamer “Blake” Part 2, Gastropoda and Scapho-
poda. Bull. Mus. Comp. Zool. 18: 1 - 492; plts. 10 - 40
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
1925. _ Illustrations of unfigured types of shells in the collection
of the United States National Museum. Proc. U.S. Nat.
Mus. 66: 1 - 41; plts. 1 - 36
1927. Small shells from dredgings off the southeast coast of
the United States by the United States Fisheries Steamer
“Albatross” in 1885 and 1886. Proc. U. S. Nat. Mus. 70:
1 - 134
DAUTZENBERG, PHILIPPE
1900. Croisiéres du yacht Chazalie dans l’Atlantique. Mollus-
ques. Mém. Soc. Zool. France 13: 145 - 256; plts. 9, 10
Deux, R. K.
1956. The archibenthal Mollusca of New Zealand. Dom.
Mus. Bull. Wellington, No. 18, 235 pp., 27 plts.
Forses, EpwArD & SYLVANUS HANLEY
1848-1853. A history of British Mollusca and their shells. Van
Vorst, London; 4 vols.; illustr.

in the Gulf of

Grant, Utyssss S., IV. « Hoyr RopNey GALE

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)
Gray, Joun 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)

Hasse, TADASHIGE

1946. On some species of tectibranchiate Mollusca found in
Japan. Japan. Journ. Malacol. 14 (5-8): 183-190
(in Japanese).

1954. Report on the Mollusca chiefly collected by the S.S.
S6y6-Maru of the Imperial fisheries experimental station on the
continental shelf bordering Japan during the years 1922-1930.
Part I: Cephalaspidea. Publ. Seto Mar. Biol. Lab. 3 (3) :
301 - 318; plt. 38

1955. A list of the cephalaspid Opisthobranchia of Japan.
Bull. Biogeogr. Soc. Japan vols. 16-19 [sic]: 54 - 79; plt. 4

HEDLEY, CHARLES

1903. Scientific results of the trawling expedition of H. M. C. S.
“Thetis” off the coast of New South Wales in February and
March 1898. Mollusca, Part II. Scaphopoda and Gastropoda.
Mem. Austral. Mus. 4 (6): 327 - 402; plts. 36 - 38

IREDALE, TOM

1936. Australian molluscan notes, No. 2

Mus. 19: 267 - 340; plts. 20 - 24
JEFFREYS, JoHN Gwyn

1862 - 1869. British conchology, or an account of the Mollusca
which now inhabit the British Isles and the surrounding seas.
London, 5 vols., illustr.

[pertinent part in vol. 4 (1867), p. 410 ff. and vol. 5. plt. 93 (1869) ]
KEEN, A. Myra

1945. (Note on use of Rhizorus MontForT, 1810 for Volvulella
Newton, 1891). Min. Conch. Club So. Calif. 47: 17 - 18
(mimeographed)

Kuropa, TokuBrEI & TADASHIGE HABE

1952. | Check list and bibliography of the Recent marine Mol-
lusca of Japan. Tokyo, pp. 1 - 210 (4 April 1952)

1954. On some Japanese Mollusca described by A. ADAMs,
whose specimens are deposited in the Redpath Museum of
Canada (No. 1). Venus 18 (1): 1-16; 2 plts.

Locarp, ARNOULD

1892. Les coquilles marines des cétes de France.

et Fils, Paris, 384 pp.
MakryaMa, J.

1927. Molluscan fauna of the lower part of the Kakegawa
Series. Mem. College Sci. Kyoto Imp. Univ. Ser. B, 3 (1):
1 - 147; plts. 1-6

Marcus, EvELYN & Ernst Marcus

1960. | Opisthobranchs from American Atlantic warm waters.

Bull. Mar. Sci. Gulf and Caribb. 10 (2): 129 - 203
Martens, EpuARD CARL VON

1903. Die beschalten Gastropoden der deutschen Tiefsee-Ex-
pedition 1898-1899. A: Systematisch-geographischer Teil:
1 - 146; plts. 1-5; 1 text fig. in Wissenschaftl. Erg. d. deutsch.
Tiefsee Exped. “Valdivia” 1898 - 1899

Rec. Austral.

Bailliére

Vol. 10; No. 2

ME tvitt, James Cosmo M.

1906. Descriptions of thirty-one gastropoda and one scapho-
pod from the Persian Gulf and Gulf of Oman, dredged by Mr.
FW. Townsend 1902 - 1904. Proc. Malacol. Soc. London
7 (2): 69 - 80; plts. 7-8

1912. Descriptions of thirty-three new species of gastropoda
from the Persian Gulf, Gulf of Oman and North Arabian Sea.
Proc. Malacol. Soc. London 19: 240 - 254; plts. 11 - 12

MENKE, Kart THEODOR

1854. Zur Familie Bullacea und deren Gattungen und Arten.

Malakozool. Blatter 1: 33 - 48
Montrort, Pierre DENYS DE

1810. Conchyliologie systematique, et classification méthodique
des coquilles; vol. 2: 676 pages. Paris (vol. 1: 410 pp.,
was published in 1808)

Mércu, Orro AnprEaAs Lowson

1875. Synopsis molluscorum marinorum Indiarum occidenta-
lium. Malakozool. Blatter 22: 142 - 184
Newton, R. B.

1891. Systematic list of the Frederick E. Edwards collection of
British Oligocene and Eocene Mollusca in the British Museum
(Natural History). British Mus., 365 pp.

Nickires, Maurice

1950. Mollusques testacés marins de la céte occidentale d’Af-
rique. Man. Ouest-Afr., 2: i-x+1-269; 459 text figs.
Paris, Lechevalier

Nomura, SHICHIHEI

1939. Notes on some Opisthobranchiata based upon the
collection of the Hoon Kai Museum, chiefly collected from
northeast Honsyu, Japan. Japan. Journ. Geol. Geogr.
(1938) 16 (1-2): 11-27; plts. 2-3

O.proyp, IpaA SHEPARD

1927. Marine shells of the West Coast of North America.

Stanford Univ. Press, 2 vols.
p Orpicny, ALcIDE DESSALINES

1842-1853. Mollusques. In Histoire physique, politique ct natu-
relle de l’ile de Cuba (French ed.), Ramon de la Sagra, ed.
Part 2

(Atlas with named figures published in 1842)
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)

Puiuipr1, RuDoLF AMANDUS

1849. Centuria tertia Testaceorum novorum. Zeitschr. f.
Malakozool. 6 (2): 17 - 26
1851. | Centuria quarta Testeaceorum novorum. Zeitschr. f.

Malakozool. 8 (4): 49 - 64
Pitspry, HENRy AuGuSTUS
1893. Tectibranchiata, in Tryon’s Manual of Conch. 15: 436
pp.; 61 plts. Acad. Nat. Sci. Philadelphia. (the genus
Volvula A. Avams is included on pp. 233 - 242; plts. 26 and 60)

THE VELIGER

Page 147

Pitssry, Henry Aucustus
1930. List of land and freshwater mollusks collected on
Andros, Bahamas. Proc. Acad. Nat. Sci. Philadelphia 82:
289 - 302; plt. 22
PoweE.LL, ARTHUR WILLIAM BADEN
1962. Shells of New Zealand.
land, 4™ ed., 203 pp.; 36 plts.
Pruvot-Fo., ALICE
1954. | Mollusques opisthobranches.
to 460; 1 plt.; 173 text figs.
SmitH, Epcar ALBERT
1872. _ A list of species of shells from West Africa, with descrip-
tions of those hitherto undescribed. Proc. Zool. Soc. Lon-
don for 1871: 727 - 739; plt. 75 (April 1872)
Srronc, ARCHIBALD McCuure & LEo GeorcE HERTLEIN
1937. The Templeton Crocker Expedition of the California Aca-
demy of Sciences, 1932. No. 35. New species of Recent mollusks
from the coast of western North America. Proc. Calif. Acad.
Sci., ser. 4, 22 (6): 159 - 178; plts. 34 - 35
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
Suter, Henry
1913. | Manual of New Zealand Mollusca.
Wellington, New Zealand, 1 - 1120; atlas of plates
THIELE, JOHANNES
1925. | Gastropoden der deutschen Tiefsee-Expedition. II. Teil.
Wiss. Erg. d. deutsch. Tiefsee Exp. “Valdivia” 1898 - 1899.
17 (2): 1-348; 34 plts.
Watson, R. B.
1883. Mollusca of the “Challenger” Expedition. Part 19.
Journ. Linn. Soc. London, Zoology 17: 319 - 346
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.
WILLETT, GEORGE
1944. | New species of mollusks from Redondo, Calif. Bull.
So. Calif. Acad. Sci. 43: 71 - 73
WINCKworTH, RONALD
1932. The British marine Mollusca.
19 (7): 211 - 252
YAMAKAWaA, G.

Whitcombe & Tombs, Auck-

Faune de France 58: 1

MacKay,

Journ. Conch. London

1911. Descriptions of some fossil opisthobranchiata from the
diluvial deposits of Japan. Journ. Geol. Soc. Tokyo 18 (211) :
38 - 45, 47 - 52; plts. 10 - 11 (not seen; information from Kuro-
DA & Habe, 1952 and Hane, 1955)
YoxoyaMa, M.
1922. Fossils from upper Musashino of Kasusa and Shimosa.
Journ. Coll. Sci. Imp. Univ. Tokyo 44 (1): 1-200; plts. 1-17

Page 148

THE VELIGER

Vol. 10; No. 2

Relationship between Rnitella penita (Conrap, 1837)

and Other Organisms of the Rocky Shore

BY

JOHN W. EVANS

Department of Biology

Memorial University of Newfoundland, St. John’s, Newfoundland, Canada

(Plate 16)

Evans (1949) anp Lewis (1953), IN THEIR STUDIES of
the English coast, refer to the intertidal animals which live
in holes, crevices and other areas, where they are protected
from the sun and from desiccation, as the “cryptofauna.”

KtHne tt (1951) suggested the following terminology
to describe the fauna of hard marine bottoms: animals
living on the surface of rock occupy the epilithion, those
partially embedded occupy the mesolithion, and those
wholly embedded occupy the endolithion. Since these
terms carry more information than ‘cryptofauna’, they
will be used in this paper. The endolithic community is
that group of animals inhabiting the endolithion.

The cryptofauna of Evans (1949) and Lewis (1953)
included a wide assemblage of relatively delicate, mostly
filter feeding animals, such as: hydroids, anemones, tuni-
cates, sponges and bryozoans. These authors observe that
the cryptofauna is present primarily in areas of exposed
limestone which is readily attacked by borers, such as the
clam Hiatella sp., the polychaete Polydora ciliata (JoHN-
sToNn, 1838) and the sponge Cliona celata Grant. The
rough surface produced by these borers serves to hold water
during low tide periods and provide homes for nestlers.
The large burrows of Hiatella were usually occupied by
a variety of anemones.

Penitella penita is the most numerous and most widely
distributed rock borer along the eastern Pacific coast
(Coan, 1964; Turner, 1955). It is found both subtidally
and intertidally, on exposed coasts and in protected bays
wherever rock of suitable hardness and composition is
available (Lioyp, 1897). The boring activity of P penita
is primarily responsible for developing the endolithion as
a possible habitat. The conical holes drilled by this
animal form dwellings for a large number of nestling
animals which move into the empty burrows after the
pholads’ death.

The following observations were made in the course of
a general ecological study of Penitella penita. The
principal area of study was on two intertidal benches
near Fossil Point in Coos Bay, Oregon. Observations
were also made on animals collected from South Jetty,
a jetty protecting the entrance to Coos Bay, and from
the north side of Cape Blanco, Oregon.

Relationship between
Penitella penita and Organisms of
the Epilithion

Once the young Penitella penita is well established in
the rock, there appears to be little interaction between it
and organisms of the epilithion, since pholads derive their
food and oxygen from the overlying water. However, sur-
face encrustations appear to control to a certain extent
the success of settlement.

Settlement on cleared surfaces facing approximately
southeast was compared with surfaces facing northeast
(Table 1). The former surfaces quickly developed a heavy
coating of algac, presumably from the greater exposure
to light, whereas the latter surfaces remained relatively
free of algae but received a heavy settlement of Balanus
crenatus Brucutére, 1789. The density of Penitella
penita was greater on the northeast barnacle-encrusted
surfaces than on the algac-coated surfaces. This indicates
that conditions suitable for settlement of pholads are simi-
lar to those for B. crenatus. It also seems likely that either
algal cover or high light intensity inhibits settlement of
both barnacles and pholads. (Sce NAGABHUSHANAM,
1959 c, 1960; Istam et al. 1951 on taxis of marine larvae. )

Vol. 10; No. 2

Table 1

Settlement Density as Related to Surface Direction

y ge ye

ac a oO a
- A ae eal FE *
© n> ao ood ~ @
@ 4 so S eS =o
Fe ee be fe Se,
iS Zo. ara ZS is <A.
50 44. x DISS O
70 3 x 24 4.3
190 21 x 62) 8 Hho
350 16 64 4.6
800 353 < 8 44.1
800 157 x 8 19.6

Interference

Between Barnacles
and Newly Settled Pholads

A certain amount of interference occurs during, and for
a short time after settlement between young pholads and
barnacles. If a pholad enters the rock close to newly-
settled barnacles, it runs a risk of having its entrance
occluded by the laterally expanding base of the barnacle.
The pholad siphon, however, appears to be able to dissolve
the edge of the barnacle and thus distort its symmetry
(see Plate 16).

Examples of barnacles completely occluding burrow
entrances were also found. The enclosed pholad was, of
course, dead. It is not known whether the barnacle
covered the entrance hole before or after the death of
the clam.

Botula californiensis (PHitippt, 1847), inhabiting the
mesolithion, interferes with Penitella penita by settling in
burrow entrances or boring laterally into burrows. The
interference caused is sometimes enough to kill P penita.

The only animal that has been observed to prey on
Penitella penita at both Fossil Point and South Jetty is
the flatworm Stylochus sp. PEARSE & WHARTON (1938)
report that S. inimicus (Patomst, 1931) is a predator of
oysters. Stylochus can enter remarkably narrow pholad
holes. For example, a flatworm about 32mm by 16mm

THE VELIGER

Page 149

was found inside a burrow, the entrance of which was
only 1.8mm in diameter. Stylochus consumes the flesh of
Penitella and leaves the valves in place. It often lays eggs
on the inside of the burrow and valves.

The empty burrows left after the death of pholads are
filled by a number of nestling animals, which make up
the remainder of the endolithic community.

At Fossil Point the empty burrows become filled with
sand and mud. Most of the silt-filled burrows are occupied
by a terebellid worm, Thelepus sp., and its commensal
scale worm, Halosydna brevisetosa KinBERG, 1855. Thele-
pus appears to extract CaCO; from the pholad valves
and deposit at least part of it as a chalky layer on
the inside of its parchment burrow. The valves of the
dead pholad are gradually dissolved completely.

Occasionally bivalves, such as Tresus nuttallii (Conran,
1837), Petricola carditoides Conrap, 1837, Macoma
nasuta (Conrap, 1837) and ZIrus lamellifer (Conran,
1837) are found nestling in pholad burrows at Fossil Point.
Appicott (1963) found fossil Tresus nuttallii nestling in
Penitella penita burrows in rock of late Pleistocene age.

Just as the epifauna on exposed rocky shores is richer
and more varied than in the muddy bays, so too the
endolithic community in rocks exposed to the open ocean
is more diverse.

Empty burrows exposed to the open ocean are very
seldom mud or sand filled. No attempt was made to
compile a complete list of organisms occupying this
habitat, but the obvious forms were collected and identi-
fied (Table 2). On the open coast, 25 taxa were observed
as compared with 9 at Fossil Point.

DISCUSSION

The position of the pholad in the littoral community can
be compared to that of the woodpecker in the forest com-
munity. Both are capable of boring holes in substrates
that cannot be worked by other members of the commu-
nity. In both cases the burrow is used only once, by its
originator, and is thercafter left open for a variety of
hole or crevice dwellers. In the absence of marine rock
borers the endolithic community will not develop. The
variety of animal specics whether epi-, meso-, or endolithic,
on the rocky shore is greatly enhanced by the presence of
borers.

Page 150

THE VELIGER Vol.

Table 2

Nestlers Inhabiting Vacated Pholad Burrows

Open Fossil
Coast Point Major Taxa
Coelenterata
>< x< Annelida
x x
x
x
x
x
x
x
x
x
x Sipunculoidea
x
S< Crustacea
x
x
x
x Mollusca
x
x
x
x
x x
x
x
x x
x
x
x Urochordata

Species
Anthopleura artemisia (PickERING in Dana, 1848)
Diadumene leucolena (VERRILL, 1866)
Thelepus sp.
Halosydna brevisetosa KinBErRG, 1877
Serpula sp.
Eupolymnia heterobranchia (JOHNSON)
Ramex sp.
Pista elongata Moore, 1909
Schizobranchia sp.
Distylia sp.
Idanthersus sp.
Demonax sp.
Phascolosoma agassizit KEFERSTEIN, 1866
Dendrostomum pyroides CHAMBERLIN, 1919
Pachycheles rudis Stimpson, 1859
Oedignathus inermis (Stimpson, 1860)
Spirontocaris palpator (OwEN, 1839)
Betacus harfordi (Kincstey, 1878)
Crepidula nummaria Goutp, 1846
Trus lamellifer (Conrab, 1837)
Trimusculus reticulatus (SOWERBY, 1835)
Tresus nuttallii (Conran, 1837)
Petricola carditoides Conrab, 1837
Macoma nasuta (Conrap, 1837)
Protothaca staminea (Conrap, 1837)
Entodesma saxicola (Bairp, 1863)
Saxicava sp.
Kellia suborbicularis (Montacu, 1804)
Botula californiensis (Putvippi, 1847)
Pyura haustor (Stimpson, 1864)

LITERATURE CITED

AppicoTT, WARREN O.

Coan, EuGENE ViIcToR

10; No. 2

1964. The Mollusca of the Santa Barbara County area. Part
I.—Pelecypoda and Scaphopoda. The Veliger 7(1): 29-33.

1963. An unusual occurrence of Tresus nuttalli (ConraD). (1 July 1964)
(Mollusca: Pelecypoda) The Veliger 5 (4): 143-145;
3 Text figs. (1 April 1963) Conran, TiMoTHy ABBOTT
Crapp, Wituam E « Roman KEnkK 1837. | Descriptions of new marine shells from Upper California,
1963. Marine borers: an annotated bibliography. Office Nav. collected By Thomas Nuttall, Esq. Journ. Acad. Nat. Sci.
Res., Dept. Navy, Washington, D.C., pp. 1136 Philadelphia 7: 227 - 268; plts. 17-20 (21 November 1837)

Explanation of Plate 16

The distorted edge of the central barnacle is caused by the young
pholad siphon which emerged from the hole at the right. The siphon
may have the ability to dissolve CaCO,

THE VELIGER, Vol. 10, No. 2 [Evans] Plate 16

Vol. 10; No. 2

THE VELIGER

Evans, R. C.
1949, The intertidal ecology of rocky shores in South Pem-
brokeshire. Journ. Ecol. 37: 120 - 139
IsHAM, LAwreENcE B., FREDERICK GEORGE, WALTON SMITH &
V. SPRINGER
1951. Marine borer attack in relation to conditions of illumi-
nation. Bull. Mar. Sci. Gulf and Caribb. 1 (1): 46 - 63
KUHNELT, WILHELM
1951. Contributions 4 la connaissance de l’endofaune des sols
marins durs. Année Biol. (3) 27 (7): 513 - 523 (fide CLapp
&« Kenx, 1963)
Lewis, J. R.
1953. | The ecology of rocky shores around Anglesey. Proc.
Zool. Soc. London 123: 481 - 549
Lioyp, Francis ERNEST
1897. On the mechanisms in certain lamellibranch boring
Mollusca. Trans. New York Acad. Sci. 16: 307 - 316
NAGABHUSHANAM, R.
1959. Settlement of marine borers (Mollusca) in the Visakha-

patnam Harbour in relation to light and gravity. Zool.
Soc. India 30 - 31
1960. A note on the inhibition of marine woodboring molluscs

by heavy fouling accumulation. Sci. and Cult. 26: 127-128
PearRSE, ARTHUR SPERRY & G. W. WHARTON
1938. The oyster “leech” Stylochus inimicus (PALOMBI) asso-
ciated with oysters on the coasts of Florida. Ecol. Monogr.
8 (4): 605 - 655
Turner, RutH Dixon

1955. The family Pholadidae in the Western Atlantic and
Eastern Pacific. Part IZ — Martesiinae, Jouannetiinae and
Xylophaginae. Johnsonia 3 (34): 65-160; plts. 35-93.

Page 151

Page 152

THE VELIGER

Vol. 10; No. 2

Freshwater Mollusks Collected
by the United States and Mexican Boundary Surveys

DWIGHT W. TAYLOR

Department of Zoology
Arizona State University, Tempe, Arizona 85281

TRACING THE TYPE LOCALITIES of several freshwater mol-
lusks led to a review of collections made by the two
commissions that surveyed the boundary between the Unit-
ed States and Mexico, in 1849 - 1855 and 1891 - 1894.
The present paper summarizes the species collected, relo-
cates the localities as precisely as possible, and corrects
previously published identifications. I have emphasized
study of specimens from west of the continental divide,
but the summary of published information concerning all
of the collections is intended to be complete.

The mollusks collected during the first boundary survey
have never been treated as a group before. Indeed, mala-
cological literature gives no indication that they are a
result of that survey. Dati (1896) reported on collections
by the second survey, but as his paper omitted some speci-
mens and gives erroneous localities for others I have
thought it worthwhile to review collections by both bound-
ary expeditions.

A general account of the boundary surveys, as well as
other early explorations in the west, was published by
Wattace (1955). This summary is briefer and more
accessible than the official reports of the boundary com-
missions. It also provides historical background and com-
ments on the numerous lighter aspects — strange as it
may seem — of these surveys.

Geographic coordinates in the form “NH 14-5 C-5”
designate a 15-minute quadrangle as explained more fully
by Taytor (1966a: 28). The letters and first number
(NH 14) are part of the system of reference of the
“International Map of the World, 1 : 1000000” published
under auspices of the United Nations Cartographic Office ;
they designate a quadrangle 4 degrees of latitude by 6
degrees of longitude. “NH 14-5” designates a subdivision,
1 degree by 2 degrees, in the system used by the U.S.
Army Map Service and U.S. Geological Survey for maps
at scale 1 : 250000.

ABBREVIATIONS

Names of Institutions housing specimens cited herein have
been abbreviated as ANSP, Academy of Natural Sciences
of Philadelphia, Pennsylvania; MCZ, Museum of Com-
parative Zoology, Cambridge, Massachusetts; and USNM,
U.S. National Museum, Washington, D. C.

First Boundary Survey (1849 to 1855)

One of the series of Commissioners of the United States
and Mexican Boundary Survey was J. R. Bartlett, who
published a voluminous account (BARTLETT, 1854) of his
travels together with a map showing his route. Dr. T. H.
Webb, Surgeon and Secretary to the survey party, collected
mollusks at localities I have discussed below. The survey
was completed by W. H. Emory, whose official account
(Emory, 1857) includes no mention of mollusks except in
the report by Parry (1857), and no indication that any
mollusks were collected.

BarTLeETT’s account (1854) is precise enough to permit
relocation of several molluscan type localities. These are
listed in the order of collection.

1. Llano River, Texas. Type locality of Lampsilis brac-
teata (Goutp, 1855b) and Quadrula petrina (GouLp,
1855 b). Collected October 18, 1850 (BARTLETT, 1854, v. 1,
p. 64), probably not far from the present crossing of U. S.
highway 87 southeast of Mason, Mason County, Texas
(NH 14-5 C-5).

Jounson (1964) illustrated type material of the two
species from this locality: plt. 31, fig. 4, holotype (MCZ
169291) of Quadrula petrina; and plt. 31, fig. 5, lecto-
type (USNM 84966) of Lampsilis bracteata.

2. Comanche Creek, Texas. Type locality of Anodonta
imbecillis horda Goutp (1855b). Collected October 18,

Vol. 10; No. 2

1850 (Bartett, 1854, v. 1, p. 65), 9 miles from the
preceding locality, close to the present site of Mason,
Mason County, Texas (NH 14-5 C-5).

Jounson (1964: 88) selected as lectotype the speci-
men (British Museum, uncatalogued) illustrated by
SowerBy (1867, plt. 18, fig. 66) as Anodon hordeum.
Subsequently the measured holotype has been found in
Gould’s collection at the U.S. National Museum (USNM
uncatalogued) by R. I. Johnson (personal communication,
March 1967).

3. Pecos River, Texas. One of two original localities of
Physa humerosa Goutp (1855 a). Bartlett’s party crossed
the Pecos October 29, 1850, and then traveled northward
along the west side until November 5 (BarTLETT, 1854, v.
1, pp. 93 - 109). Horse-head Crossing is a plausible site
of collection; it is near the bridge on U. S. highway 67-385
over the Pecos, where the river forms the boundary
between Crockett and Pecos Counties, Texas (NH 13-3
A-2).

Collections made by Dr. Webb in Arizona, Baja Cali-
fornia, and California might have been made en route
to San Diego, California, in late 1851 and early 1852, or
(more probably) on the return journey eastward in 1852.
The report by Wess to BarTLetT (1854, v. 2, pp. 1-5),
and Bartuett’s (1854, v. 2, p. 550) statement about
Webb’s collections, indicate loss of baggage and physical
hardships on the westward trip.

4. Colorado Desert or Cienega Grande, Baja California
and California. Type locality of Biomphalaria gracilenta
(Goutp, 1855a), the type specimen collected by Webb.
Gou p also cited Webb as one of the collectors of original
material of Physa humerosa, Planorbis ammon, and Try-
onia protea, but whether Webb or W. P. Blake collected the
type specimen is not determinable and does not matter in
this case. Bartlett and Webb crossed the Colorado Desert
to Fort Yuma June 5 to 9, 1852 (BartLett, 1854, v. 2,
pp. 129-152).

5. “River Gila, near San Diego” (seemingly one local-
ity) was cited by Goutp (1862: 216) as the source of
Physa virgata. “River Gila, and near San Diego” (evi-
dently two localities) was the original form (GouLp,
1855a). I have found only one San Diego near the Gila
River; it was on the route traveled by Dr. Webb and is
probably the type locality. The name was given by Father
Garcés November 12, 1775, to a group of Indian rancher-
ias that Coues (1900, v. 1, p. 117) identified as being
most likely at Kenyon’s. This was a stage station on the
overland mail route, about in the NEi T.5 S., R. 7 W,
Maricopa County, Arizona (NI 12-7 A-4).

If San Diego is taken as a second locality, additional to
the Gila River, there are two possible identities. One is
the city in southern California, where Bartlett and Webb

THE VELIGER

Page 153

stayed in the spring of 1852. The other was on the Rio
Grande in New Mexico, upstream from El Paso. Dr. Webb
crossed the river here at least three times: in the spring
of 1851, en route to and from the copper mines at Santa
Rita del Cobre (Bart Lett, 1854, v. 1, pp. 178 - 181), and
on April 27, 1851, when the Survey party as a whole
traveled westward (Bart Lett, 1854, v. 1, p. 215).

Jounson (1964, plt. 44, fig. 5) illustrated his lectotype
(MCZ 72995) of Physa virgata. I have examined one
serics of paratypes (USNM 27 966), but not the specimen
(ANSP 17 244a) identified by H. B. BaKer (1964) as the
holotype.

6. Bartlett mentioned collecting mollusks only once, and
strangely the specimens seem never to have been recorded
in scientific writings, nor has anyone else collected mol-
lusks there. The entry is for August 6, 1852, 8 miles west of
Janos, Chihuahua: “In the afternoon we crossed a fine
clear stream, thirty to forty feet wide and about two deep,
occasionally expanding into ponds twice that depth, and
encamped on the opposite bank where there was excellent
grass. This stream is a branch of the river which passes
Janos and Correlitos, the latter being known both as the
San Miguel and as the Casas Grandes River. Like other
streams in Mexico, it takes the names of the several towns it
passes. In it we caught a sufficient number of trout to
give us all a meal. We also collected specimens of unios,
which abound here ...” (BARTLETT, 1854, v.2, p. 337).
The locality is shown on the American Geographical Soci-
cty’s Map of Hispanic America, 1: 1000000, Sonora
shect (1937) as Arroyo del Salto del Ojo (NH 12-6 D-2).

7. Ojo Caliente, Chihuahua. “Cochliopa” chihuahua
(Pitspry, 1928) was described from 4 shells found on
an herbarium sheet of Myriophyllum heterophyllum in the
Academy of Natural Sciences of Philadelphia. The speci-
mens were collected by George Thurber, October, 1852.
Thurber was botanist on the first Boundary Survey (Bart-
LETT, 1854, v. 1, p. 47), the party camping at Ojo Caliente
on October 16, 1852 (Bartiert, 1854, v. 2, p. 410). The
narrative gives the location as 12 miles south of Carrizal
and a mile north of the “river Carmen” (now Rio del
Carmel); these distances and the map by BarTLett
(1854) are consistent with the place shown as Ojo Cali-
ente de Santa Rosa on the American Geographical Soci-
cty’s Map of Hispanic Amcrica, 1 : 1000000, Chihuahua
sheet (1934) (NH 13-4 B-3).

For the convenience of anyone trying to relocate the
type locality on the ground, I quote Bartlett’s description:

“Ojo Caliente is a spring which rises from the plain
about one hundred and fifty yards from the base of a
rocky hill. Its temperature is nearly the same as that of
the atmosphere. A small pool about one hundred and
twenty feet in circuit, and from three to four deep, is here

Page 154

formed, with a sandy bottom, from which warm water
bubbles up in many places; this water has an outlet
through a small creek into the river Carmen, in which
creek some fish were taken and preserved for specimens”
(BARTLETT, 1854, v. 2, p. 410).

8. Puntiagudo, Nuevo Leon. The type locality of Unio-
merus tetralasmus manubius (Goutp, 1855b) is “Chihua-
hua, 60 miles from Camp Ringgold.” According to the
itinerary this locality would have been about 30 miles
southwest of Mier, Tamaulipas, at the village of Puntia-
gudo where Bartlett and Webb passed December 18, 1852.
BarTLeTT (1854, v. 2, p. 507) noted “A small stream
passes here, one of the tributaries of the Alcantra.”

Neither of the names Alcantra or Puntiagudo appears
on modern maps. The map by RoEMER (1935; first pub-
lished in 1849 and dating to 1845-1847) shows the “Ac-
lantro R.” flowing into the Rio Grande near Mier,
evidently the Rio Alamo of modern usage.

Puntiagudo is identified by WistizENus (1848: 78) as
“a burnt village on a creek, which is one of the headwaters
of the Alamo,” 15 miles northeast of Cerralvo, Nuevo
Leon, and 30 miles southwest of Mier, Tamaulipas. This
description and WIsLIZENUS’ map show that the site of
Puntiagudo is about 3 miles northeast of the modern town
General Trevifio, Nuevo Leon, on the Rio Agualeguas,
a tributary of Rio Alamo (NG 14-5 B-6).

Jounson (1964, plt. 32, fig. 5) illustrated the holotype
(MCZ 169447) of Uniomerus tetralasmus manubius.

9. Sphaertum nobile (GouLp, 1855b) was described
from “near San Pedro, California,” a locality known now to
be in error. A possible locality might seem to be the San
Pedro River drainage of southern Arizona, crossed by
Bartlett and Webb several times. Yet a paratype of S.
nobile (USNM 11592) seems to me to be S. striatinum
(Lamarck) and not S. triangulare (Say), the only sim-
ilar species in Arizona. Hence a “San Pedro” farther east
is most likely, perhaps San Pedro Springs at San Antonio,
Texas, where Bartlett and Webb camped September 27
to October 9, 1850 (BARTLETT, 1854, v. 1, pp. 38 - 48).

I found no explicit statement about the scope or final
disposition of Webb’s collection of mollusks. BARTLETT
(1854, v. 1, p. viii) mentioned that zoological collections
were divided between the Smithsonian Institution and
Boston. The Tenth Annual Report of the Smithsonian
Institution, for 1855 (p. 47) mentions “The fruits of the
travels of Dr. Thos. Webb, in the more western portions
of northern Mexico,” but does not refer explicitly to shells.
The Proceedings of the Boston Society of Natural History
for December 21, 1853 (4: 395) referred to the “collection
of Dr. Webb, now on deposit with the Society,” from
which the shells were referred to Dr. Gould. Whether
these were all of Webb’s collections, and whether addition-

THE VELIGER

Vol. 10; No. 2

al locality data will turn up, depends on unpublished
sources, if any.

Second Boundary Survey (1891 to 1894)

During the second Boundary Survey mollusks were collec-
ted by E. A. Mearns, who like T:H. Webb was medical
officer to the party. Unlike Webb, Mearns (1907) wrote
an exemplary account of his itinerary and collecting local-
ities. This work is of value because of its detailed descrip-
tion of places that have been modified subsequently by
agriculture and irrigation development. DA.u’s (1896)
report on the mollusks mislocates some localities, or locates
them less precisely than Mearns. In the following list
the mollusks are discussed by Mearns’ collecting stations,
in east-west sequence. In all cases quotations describing
the localities are from Mearns’ account. The citation of
USNM catalogue numbers indicates I have examined the
specimens, the lack of such citation that I have not been
able to find the specimens in USNM collections. Revised
identifications of the species, and occurrence by locality,
are listed in Table 1.

Mearns station 1 (NI 14-12 C-6, D-6). Fort Worth,
Tarrant County, Texas. “Birds and shells were collected
on the South Fork of Trinity River January 30 and 31,
1892, by Mearns and Holzner.” Datu (1896: 368) re-
corded Limnaea bulimoides LEA (USNM 130185) and
Physa mexicana Puiuipp1 (USNM 130225). The latter
set is labeled “Rio Grande, Ft. Worth, Texas,” giving rise
to the suspicion that this series is from El Paso (Mearns
station 5).

Mearns station 2 (NH 14-7 B-2). Fort Clark, Kinney
County, Texas. “The post is located at the head of Las
Moras Creek, a wooded stream encircling it on three
sides....”” Mearns made a few collections in 1892 and
1893, but most of the mollusks were collected in 1897-
1898 after the report by Dat (1896) had been published.
The following list of species was published by MEARNS
(IGO72 77) s

Limnaea columella (Say)

Limnaea humilis Say

Physa osculans HALDEMAN

Planorbis liebmanni DUNKER

Planorbis tumidus PFEIFFER

Planorbis trivolvis Say

Planorbis bicarinatus Say

Valvata guatemalensis MoRELET

(USNM 151544)
(USNM 218 412)
(USNM 218 424)

(USNM 151533)
(USNM 151532,
218425, 218426)
Amnicola peracuta WALKER

Amanicola sp.

Sphaerium elevatum HALDEMAN

Vol. 10; No. 2

THE VELIGER

Page 155

Table 1

Freshwater Mollusks Collected by E. A. Mearns during the Second United States and Mexican Boundary Survey.

Mearns Localities

1 2 5 8 24 25 46 58 66 67 68 73 76 77 81 91 92 219

x

Bakerilymnaea techella (HaLpEMAN, 1867)
Physa virgata Goutp, 1855 a
Elliptio popet (Lea, 1857)
Lampsilis anodontoides (LE, 1830)
Cyrtonaias tampicoensis berlandieri (LEA, 1857)
Carunculina parva compressa (Simpson, 1900)
Carunculina parva mearnsi (Simpson, 1900)
Carunculina parva texasensis (LEA, 1857)

_ Megalonaias gigantea (BaRNES, 1823)
Quadrula couchiana (Lea, 1860)
Anodonta imbecillis horda Goutp, 1855b
Sphaerium partumeium (Say, 1822)
Sphaerium striatinum (Lamarck, 1818)
Pisidium compressum PriME, 1852
Cincinnatia peracuta (PitsBry & WALKER, 1889)
Cochliopina riograndensis (Piuspry & Ferriss, 1906)
“Amnicola” sp.
Fossaria obrussa (Say, 1825)
Pseudosuccinea columella (Say, 1817)
Biomphalaria liebmanni (DuNxKER, 1850)
Helisoma anceps (MENKE, 1830)
Planorbella trivolvis (Say, 1816)
Planorbella tenuis (DUNKER, 1850)
“Bythinella” palomasensis Piuspry, 1895
Anodonta dejecta Lewis, 1875
Sphaerium triangulare (Say, 1829)
Anodonta californiensis Lea, 1852
Physa humerosa Goutp, 1855 a
Biomphalaria gracilenta (Goutp, 1855 a)
Tryonia protea (GouLp, 1855 a)

XK OS 8 OK OS OS OS OS HK KN KOON ON OS ON OOK

x
BOS XS Ks x x x xX XxX
x x XK
x
x x X X x x xX X x
x
Xx
x
x x xX x x
x x
x
x

Sphaerium solidulum Prime (USNM 151541,
216217, 216 218, 216219)
Pisidium compressum PRIME
Lampsilis anodontoides LEA
Lampsilis texasensis LEA (USNM 151 536, 151 540)
Lampsilis texasensis compressus SIMPSON
(USNM 151539, 152059)
Lampsilis berlandieri Lea (USNM 151 548)
Lampsilis mearnsi Simpson (USNM 151549,
308 846)
Anodonta imbecillis Say
Unio popeti LEA
Quadrula undulata BARNES
Quadrula couchiana LEA (USNM 308 855)
From the earlier collections by Mearns, 1892 - 1893,
Dat (1896: 368 - 371) recorded Physa mexicana Putt-

(USNM 308 943)

ippt from Fort Clark (USNM 130 226), Planorbis
tumidus PreirreR from Las Moras Creek (USNM
130 228), Unio couchianus LEa without locality (USNM
130 207, 152058), Unio undulatus BARNES from Kinney
County, and Unio popeii from Kinney County (USNM
130175, 151538). From the description of his collecting
by Mearns (1907) I suppose that all of these specimens
came from nearby Fort Clark, in spite of the discrepant
published data. C.'T. Simpson (1900) described Lampsilis
mearnsi from Mearns’ collections around Fort Clark.
Two labels with a series of paratypes of Lampsilis
mearnst (USNM 151549) provide discrepant locality
data. One label says “Ft. Clark, Texas,” the other “Elm
Cr., 24 mi. S. Ft. Clark, Texas.” The label of USNM
308 846 says “from pool, 20 mi. S. of Ft. Clark, Texas,”
with a note on the back quoting Mearns: “These are

THE VELIGER

Page 156

topotypes, collected at the same time and place as the
types.”

The specimens of Cochliopina riograndensis (PILSBRY
& Ferriss) recorded by Taytor (1966b: 177) from
Kinney County, Texas, without more precise data probably
come from Mearns’ collection at Fort Clark.

Mearns station 5 (NH 13-1 D-2). El Paso, El Paso
County, Texas. Collections were made by Mearns in Feb-
ruary, March, and November, 1892, and June, 1893.
Mearns (1907: 80) noted that “Fresh-water mollusks
are numerous in the Rio Grande.” Dati (1896: 368-369)
recorded Limnaea bulimoides LEA (USNM_ 130186,
130235) and Planorbis tumidus PreirFER (USNM
130239) from the Rio Grande “near El Paso.” BAKER
(1945, plt. 98, fig. 25; plt. 99, figs. 9, 10) illustrated some
of the “Planorbis tumidus” as Helisoma tenue sinuosum
(Bonnet). I suspect the record by BAKER (1911: 216) of
Galba bulimoides techella from the Rio Grande, near El
Paso, is based on Mearns’ specimens.

Mearns station 8 (NH 13-1 D-7). “Palomas Lakes,
Mimbres Valley, Chihuahua, Mexico. One mile south of
Monument No. 21.” Collections were made from April 7
to 15, 1892. Dax (1896: 368-370) recorded Physa mexi-
cana Puteri (USNM 130183), Bythinella palomasensis
Pitssry (USNM 130016) and Pisidium compressum
Prime (specimens not found). Pitspry (1895) described
B. palomasensis from these specimens, the label of which
gives the locality as “Mimbres valley near boundary mon-
ument 19.” Monument 21 is in the W3 sec. 18, T. 29 S.,
R. 7 W, and monument 19 in the Ed of sec. 14 of the
same township, 4.6 miles to the east. The types of B. palo-
masensis were not alive when collected, for the shells are
white and bleached. They are perhaps fossil, or at least
stream-drifted.

Mearns station 24 (NH 12-3 B-5). Hall’s Ranch, Gua-
dalupe Canyon, Sonora. “Camp was made at Monument
No. 73, in the canyon.” The locality is 2100 feet west,
900 feet south of the northeast corner sec. 21, T. 24 S., R.
32 E., Cochise County, Arizona. Mearns camped here
in July and August, 1892; and August and October, 1893.
Dati (1896: 368) recorded Physa mexicana PHILIPPI
(USNM 130182) from Guadalupe Canyon, New Mexico.

Mearns station 25 (NH 12-3 B-6). San Bernardino
Ranch, Cochise County, Arizona. “Camp was made at
Monument No. 77, in a mesquite flat between the San
Bernardino Springs and the neighboring San Bernardino
River.” Mearns’ collections seemingly came from the Rio
San Bernardino on both sides of the boundary. The river
crosses the boundary line in the San Bernardino grant, in
what would be the NW3 sec. 23, T. 24 S., R. 30 E., Cochise
County, Arizona. Mearns camped here in July and Sep-
tember, 1892; and in August and October, 1893.

Vol. 10; No. 2

Dati (1896) recorded several species from “San Ber-
nardino River,” variously quoted by him in Arizona, New
Mexico, and northern Mexico: Physa mexicana PHILIPPI
(USNM 130219), Planorbis tumidus PreirreR (USNM ©
130 230, 130 233), Sphaerium solidulum Prime (USNM
130 236), Pisidium compressum Prime (USNM 130241),
and Anodonta dejecta LEwis (USNM 130208).

Victor Sterki applied the manuscript name Sphaerium
eminens to this collection of Sphaertum. No formal de-
scription has been published but Brooks « HERRINGTON
(1944: 95) cited S. emznens in the synonymy of S. stria-
tinum Lamarck. In my opinion these specimens are S.
triangulare (Say). Simpson (1893) described Anodonta
mearnsiana (a synonym of A.dejecta) from this collection.

The Physa (USNM 130219) is identified as Physa
virgata in a comprehensive sense, but the series is not
typical and might come from an unusual habitat, or
represent another species. Some of the smaller specimens
are usual P virgata in shape, but most are short-spired
with an expanded body whorl.

Mearns station 46 (NI 12-11 A-4). Tucson, Pima
County, Arizona. Mearns collected both living and stream-
drifted specimens in the Santa Cruz River near Tucson,
in October, November, and December, 1893. From the
river drift “near Tucson” Dati (1896: 368-369) recorded
Limnaea desidiosa Say and Aplexa hypnorum LINNAEUS,
and from the Santa Cruz River near Tucson, DALL
(1896: 368-369, 373) listed Physa mexicana PHILIPPI
(USNM 130220), Planorbis tumidus PreirFER (USNM
130 237), and Anodonta dejecta Lewis (USNM 130172,
“Tucson;’”” USNM 130 180, “Santa Cruz River, near Tuc-
son”). STEARNS (1901: 289) suggested the “Aplexa hyp-
norum” might prove to be one of the Mexican species
now classified in Physa (Stenophysa), but I consider the
single specimen (USNM 130188) is only an elongate
Physa virgata. Some of the “Planorbis tumidus” (USNM
130237) were illustrated by Baker (1945, plt. 98, figs.
9-13; plt. 99, figs. 4-8) as Helisoma tenue sinuosum
(Bonnet). The record of Limnaea desidiosa was assigned
by Baxer (1911: 272) to Galba obrussa (Say). A lot
of Pisidium compressum PrimME (USNM 130 243) was not
mentioned by Datu (1896).

Mearns station 58 (NH 12-1 D 5). “Rancho de Agua
Dulce, Sonoyta River, Sonora, Mexico. This place is
about 6.4 kilometers (4 miles) southwest of Monument
No. 173.” Mearns collected here in January and February,
1894. Dati. (1896: 369) recorded Planorbis tumidus
PFEIFFER from “Sonoyta River, northern Mexico.” The
specimens are USNM 130229, one of which was illus-
trated by Baker (1945, plt. 98, fig. 14) as Helisoma
tenue sinuosum (BONNET), from “Soroyta” Creek, Sonora.

Vol. 10; No. 2

THE VELIGER

Page 157

Mearns station 66 (NI 11-12 D-2). Gila River at Gila
City (now Dome), Yuma County, Arizona. “Along the
Gila River are numerous sloughs, bordered with cat-tail,
tule, cane, sedge, and rush.” A series of Physa virgata
Goutp (USNM 187481) collected by Mearns during
March 1 to 5, 1894, was not mentioned by Datu (1896).

Mearns station 67 (NI 11-12 C-3 A). Yuma, Yuma
County, Arizona. “This station is on the left (east) bank
of the Colorado River, at the mouth of the Gila,” where
Mearns collected in March and April, 1894. Dati (1896:
369) recorded Planorbis tumidus PrerirFER (USNM
130 232, 130234).

Mearns station 68 (NI 11-12 B-4). Monument No. 204.
“This camp was beside a laguna of the Colorado River,
at the east edge of the bottom land,” sec. 12, T. 11 S., R.
25 W, Yuma County, Arizona. Mearns collected here
during March, 1894. Dati (1896: 373) recorded Ano-
donta dejecta Lewis from “Colorado River, near the Mex-
ican boundary.” The specimens (USNM 130171) are
labeled monument 204. A series of Physa virgata GouLD
(USNM 130224) from this collection was overlooked by
Dat (1896).

Mearns station 73 (NH 11-3 D-4). “Left (east) bank
of the Colorado River, opposite the mouth of Hardy
River, Sonora, Mexico.” Mearns collected mollusks here
during March, 1894. Datu (1896: 373) recorded Ano-
donta dejecta Lewis from “mouth of Colorado River”
(USNM 128801).

Mearns station 76 (NI 11-12 C-5). Seven Wells, Baja
California. “This station is 8 kilometers (5 miles) south
of Monument No. 213.” Mearns collected mollusks here
during April, 1894. Seven Wells was a former watering
place on the wagon road between Yuma, Arizona, and San
Diego, California. It is shown on the U.S. Geological
Survey “Reconnaissance map of the Salton Sink, Califor-
nia” (1908), scale 1 : 500000. Dati (1896: 368-369)
recorded Planorbis tumidus PFEIFFER (USNM 130238)
and Physa mexicana Putuippi (USNM 130 217, subfossil? )
from Seven Wells, Arizona.

Mearns station 77 (NI 11-12 C-5). Gardners Laguna,
Baja California. “Station about 10 kilometers (6 miles)
south of Monument No. 216.” Mearns collected here
during April, 1894. This was evidently one of a series of
ponds and sloughs along the course of the Alamo River
at that time. Dati (1896: 369) recorded Planorbis tumi-
dus PFEIFFER (USNM 128952).

Mearns station 81 (NI 11-12 C-7 or D-7). Laguna
Station, Imperial County, California. “Station about 11
kilometers (7 miles) north of Monument No. 224,” where
Mearns camped May 3 to 6, 1894. This locality was a

station on New River, about sec. 25, T. 16 S., R. 12 E.
Dart (1896: 369, 373) recorded Planorbis tumidus Preir-
FER (specimens not found), Anodonta dejecta LEwis
(USNM 128812) and A. californiensis Lea (USNM
128817)

Mearns station 91 (NI 11-11 C-2). “Thomas Cameron’s
Ranch, San Diego County, Calfornia, 13 kilometers (8
miles) nearly north of Monument No. 240.” Mearns
collected here June 21 to 23, 1894. Cameron Valley is
traversed by La Posta Creek in secs. 9, 10, and 16, T.17S.,
R. 5 E., evidently the source of the specimens (USNM
130218) that Dart (1896: 368) recorded as Physa
mexicana PHILIPPI.

Mearns station 92 (NI 11-11 D-2). “Campbell’s Ranch,
at Laguna Mountains (Coast Range), San Diego County,
California, 31 kilometers (19 miles) north of Monument
No. 240.” Mearns collected here June 9 to 21, 1894. Dati
(1896: 368-369) recorded as from “Laguna, 20 miles
north of Campo” Physa mexicana Pumiper (USNM
130223) and Planorbis liebmannii DUNKER (specimens
not found). The locality is evidently either Big Laguna
Lake or Little Laguna Lake, both in sec. 10, T. 15 S.,
R. 5 E.

Monument 219 (NI 11-12 C-6B). This monument on
the international boundary is in sec. 15, T. 17 S., R. 15 E.,
Imperial County, California, about 34 miles east of Cal-
exico. Mearns assigned no station number. The specimens
are all fossils from the surface of the desert, and were
recorded by Dati (1896: 369, 373, 376) as Planorbis
tumidus PFEIFFER (specimens not found), Anodonta cali-
forniensis LEA (USNM 128802), Physa humerosa Goutp
(specimens not surely found), and Amanicola protea
Goutp. The series of Physa from this locality might be
USNM 130 184, a set of subfossil P humerosa labeled as
collected by Dr. Mearns but without locality data.

A series of subfossil specimens of Physa humerosa
Goutp (USNM 130450) is labeled as coming from “Salt
Creck, W. side of Colorado Desert,” collected by Mearns.
This collection is not from any of Mearns’ (1907)
numbered stations.

ACKNOWLEDGMENTS

For the opportunity to study and search for specimens in
the U.S. National Muscum, I am indebted to Joseph
Rosewater, Curator of the Division of Mollusks. Richard
I. Johnson, Associate in the Department of Mollusks,
Muscum of Comparative Zoology, Harvard University,
reviewed the paper and provided unpublished information
on type specimens.

Page 158

THE VELIGER

Vol. 10; No. 2

LITERATURE CITED

BaKER, FRANK COLLINS
1911. The Lymnaeidae of North and Middle America, Recent
and fossil. | Spec. Publ. Chicago Acad. Sci. 3: i-xvi+1 - 539;
plts. 1-58
1945. The molluscan family Planorbidae; collation, revision,
and additions by Harley Jones Van Cleave. | Urbana (Univ.
Illinois) i- xxxvi+1-530; plts. 1-141
Baker, Horace BurriINGTON
1964. Type land snails in the Academy of Natural Sciences of
Philadelphia. Part III. Limnophile and thalassophile Pulmo-

nata. Part IV. Land and fresh-water Prosobranchia. Proc.
Acad. Nat. Sci. Philadelphia 116: 149-193 (20 Oct. 1964)
BaRTLETT, JOHN RUSSELL

1854. Personal narrative of explorations and incidents in Texas,
New Mexico, California, Sonora, and Chihuahua, connected
with the United States and Mexican Boundary Commission,
during the years 1850, °51, 52, and °53. New York (D.
Appleton & Co.) ; 2 vols., map and illus.

Brooks, STANLEY TRUMAN & Harry BiccGAR HERRINGTON

1944. The Sphaeriidae, a preliminary survey. The Nautilus
57: 93 - 97
CoveEs, ELLIoTT
1900. On the trail of a Spanish pioneer; the diary and

itinerary of Francisco Garcés (Missionary Priest) in his travels
through Sonora, Arizona, and California, 1775-1776
New York (Francis P. Harper), 2 vols.
Dai, WILLIAM HEALEY
1896. Report on the mollusks collected by the International
Boundary Commission of the United States and Mexico, 1892-
1894. Proc. U.S. Nat. Mus. 19: 333 - 379; plts. 31 - 33

Emory, WILLIAM HEMSLEY

1857. | Report on the United States and Mexican boundary
survey U.S. 347 Congress, 1* session, Sen. Exec.
Doc. 108, House Exec. Doc. 135, 1 (1): i-xvi+1- 258; maps
and illus.

GouLp, Aucustus ADDISON

1855a. New species of land and fresh-water shells from western
(N.) America. Proc. Boston Soc. Nat. Hist. 5: 127 - 130

1855 b. [Descriptions of shells]. Proc. Boston Soc. Nat. Hist.
5: 228 - 229

1862. Otia conchologica. Boston (Gould & Lincoln) : 1 to
256

Jounson, RicHarp IRwIN
1964. The Recent Mollusca of Augustus Addison Gould.
Bull. U.S. Nat. Mus. 239: 1 - 182; plts. 1-45
(28 July 1964)

Mearns, Epcar ALEXANDER

1907. Mammals of the Mexican boundary of the United
States; a descriptive catalogue of the species of mammals
occurring in that region, with a general summary of the natural
history, and a list of trees. Bull. U.S. Nat. Mus. 56:
i-xv+1 - 530; plts. 1-13

Parry, CHARLES CHRISTOPHER

1857. Physical and geological description of the country from
the initial point on the Pacific to the junction of the Gila and
Colorado. In W.H. Emory, Report on the United States
and Mexican boundary survey U.S. 34™ Congress,

1ST session, Sen. Exec. Doc. 108, House Exec. Doc. 135, 1 (2) :
78 - 99

Pitspry, Henry AucusTus
1895. A new Mexican Bythinella.
1928. Mexican mollusks.
phia 80: 115-117
RoEMER, FERDINAND

1935. Texas, with particular reference to German immigration
and the physical appearance of the country. Transl. from
the German by Oswald Mueller. San Antonio, Texas
(Standard Printing Co.) : i-xii+1-301; map

The Nautilus 9: 68 - 69
Proc. Acad. Nat. Sci. Philadel-

Simpson, CHARLES TORREY
1893. Anew Anodonta. The Nautilus 6: 134 - 135
1900. New and unfigured Unionidae. Proc. Acad. Nat. Sci.
Philadelphia 52: 74 - 86; plts. 1-5

SoweErBy, GrorGE BRETTINGHAM

1867 - 1870. Monograph of the genus Anodon.
Aucustus Reeve, Conchologia Iconica 17.
Reeve). 37 col. plts. and sheets of text

In LovE.Lu
London (L. A.

STEARNS, RoBerT EDWARD CaRTER

1901. The fossil fresh-water shells of the Colorado Desert,
their distribution, environment, and variation. Proc. U.S.
Nat. Mus. 24: 271 - 299; plts. 19 - 24

TayLor, DwicHt WILLARD
1966a. Summary of North American Blancan nonmarine mol-
lusks. Malacologia 4: 1 - 172
1966b. A remarkable snail fauna from Coahuila, México.
The Veliger 9 (2): 152 - 228; plts. 8- 19; 25 text figs.
(1 October 1966)

WALLACE, Epwarp S.
1955. The great reconnaissance.
(Little, Brown & Co.) : 1 - 288
WISLIZENUS, FREDERICK ADOLPHUS
1848. Memoir of a tour to northern Mexico, connected with

Col. Doniphan’s expedition, in 1846 and 1847. U.S. 30™
Congress, 15T session, Misc. Doc. 26: 1-141; maps

Boston and Toronto

Vol. 10; No. 2

THE VELIGER

Page 159

New Records of Nudibranchia

(Gastropoda : Opisthobranchia : Nudibranchia )
from the Central and West-Central Pacific

with a Description of a New Species

DAVID K. YOUNG

Department of Zoology
University of Hawaii, Honolulu, Hawaii 96822"?

(18 Text figures)

INTRODUCTION

INFORMATION ON THE nudibranchiate opisthobranchs of
the Central and West-Central Pacific consists chiefly of
morphological studies on preserved specimens collected
during exploratory expeditions to the Pacific Islands. A
recent example is the taxonomic study of Marcus (1965)
on Micronesian opisthobranchs in which 28 species of
nudibranchs are identified, 7 of which are new.

The purpose of this paper is to present new records and
biological notes of nudibranchs collected from Palmyra
Atoll, Johnston Island, and Eniwetok Atoll. Palmyra
Atoll, the northernmost of the Line Islands, is located at
5°51’N and 162° 04’W. Johnston Island, which is lo-
cated at 16° 15’N and 169° 30’W, is unusual in that it
has a marginal reef only at one side and that it comprises
the only land mass in the triangular area formed by the
Hawaiian Islands 450 miles to the northeast, the Line
Islands 700 miles to the southeast and the Marshall
Islands 1300 miles to the west. Eniwetok Atoll, one of the
westernmost of the Marshall Islands, is located at 11° 30’
N and 162° 15’E.

‘ Contribution No. 272, Hawaii Institute of Marine Biology,
Honolulu, Hawaii. Work supported by the AEC Eniwetok
Marine Biological Laboratory and a Marine Science Graduate
Research Fellowship from the Bureau of Commercial Fisheries.
Based on a doctoral dissertation submitted to the Graduate
School, University of Hawaii, Honolulu, Hawaii.

2 Present address: Systematics-Ecology Program, Marine Biological
Laboratory, Woods Hole, Massachusetts 02543

Nudibranchs have not previously been reported from
Palmyra Atoll and Johnston Island, whereas Marcus &
Burcu (1965) report 8 species of nudibranchs from Eni-
wetok Atoll, 2 of which are new. One specimen of Hexa-
branchus marginatus (Quoy « Gaimarp, 1832), identified
by Marcus (1965) from a 1946 collection of J.P. E.
Morrison, in addition to these 8 species, makes a total
of 9 nudibranchs previously reported from Eniwetok.

ACKNOWLEDGMENTS

Nudibranchs were collected from Eniwetok Atoll between
June 21 and July 9, 1965 under the auspices of the AEC
Eniwetok Marine Biological Laboratory and the director,
Dr. Robert W. Hiatt (University of Hawaii), to whom the
author gives grateful acknowledgment for financial sup-
port. Specimens from Palmyra Atoll and Johnston Island
were collected in June, 1962 and August, 1965 by the
Hawaii Institute of Marine Biology and were given to the
author by Dr. E. Alison Kay (University of Hawaii) and
Mr. William van Heukelem (University of Hawaii).
Special thanks are due Dr. E. Alison Kay for her interest
and aid in all aspects of this study. Appreciation is also
duc Dr. Gerald J. Bakus (University of Southern Cali-
fornia) for his identification of the sponges in this study.
The author appreciates the comments and criticisms of
the manuscript by Dr. Melbourne R. Carriker, Dr.
Victor A. Zullo, Dr. Barry A. Wade and Dr. Thomas J. M.
Schopf (all of the Systematics-Ecology Program, Marine
Biological Laboratory).

Page 160

COLLECTIONS ann METHODS

Except for one specimen collected from Igurin Island,
Eniwetok Atoll, two specimens from Palmyra Atoll and
onc specimen from Johnston Island, the nudibranchs dis-
cussed in this paper are from three locations on Eniwetok
Atoll: (1) a subtidal flat at the northern lagoonward side
of Chinimi Island, (2) a tidepool on the northern lagoon-
ward side of Eniwetok Island, and (3) the quarry on the
northern seaward reef of Eniwetok Island. These three
sites represent a type of shallow water environment rela-
tively rare on Eniwetok Atoll: rock areas comparatively
free of sand and carbonate sediment and with a free
exchange of water at all stages of the tide.

Collections were made from the intertidal zone to a
depth of 5 meters. Species of Doridacea from Hawaii
referred to in this paper were collected from the islands of
Oahu and Kauai between 1962 and 1966. Descriptions,
figures and ecological notes will be given for these and
other species in a paper by Kay & Younc (in prepa-
ration).

Descriptions, measurements and drawings of external
morphological features were made from live specimens.
Observations of feeding, when possible, were made from
animals held in laboratory aquaria. Internal anatomical
examination was performed on specimens rclaxed by
refrigeration, fixed in 5% formalin and stored in 70%
ethyl alcohol. Animals are described and figured where
these data are not adequately provided in the literature.

SPECIES

DoRIDACEA

DoriDIDAE

Doriopsis viridis PEASE, 1861
(Text figures 1 to 3)

Doriopsis viridis PEASE, 1861: 244 - 245; Pease, 1871: 301, plt. 19,
figs. la, 1b, le

Guyonia viridis (Pease), Rispec, 1928 (in part): 106 (variété
verte not variété bleu)

Doriopsis viridis (PEASE), RisBrc, 1953 (in part) : 45 (vancété verte
not varicté bleu)

Description: Doriopsis viridis has an oblong body that is
flattened dorso-ventrally (Figure 1). No specimens were
found greater than 8mm in length and 3mm in width.
The mantle, foot, rhinophores and branchiac are olive
green to bronze green. The mantle, which flares around
the foot, is firm and has small spiculate pustules resemb-
ling “granules” over the dorsum. Along the mid-dorsum,
elongate epidermal spicules are arranged diagonally in

THE VELIGER

Vol. 10; No. 2

Figure 1

Doriopsis viridis
Dorsal view of adult specimen

approximately alternate series. The 7 to 8 branchiae are
simply pinnate and retractile into the transversely posi-
tioned, crescent shaped branchial sheath at the posterior
end of the mantle. The rhinophores are rod-like with 6 to
9-lamellae and are retractile into separate sheaths.

The radular formula of 2 specimens, 6 and 8mm long, is

26-30 x 24-260: 24-26 (Figure 2). The radular teeth are
hamate, non-denticulate and range in size from 46 to a
maximal length of 119 in an 8mm specimen. The largest
tecth are centrally positioned within each row. No jaws
are present. Ducts in the female portion of the reproduc-
tive system (Figure 3, ud, v) are parallel in arrangement.
The spermatocyst (sc) is ovate-oblong and smaller than
the spherical spermatheca (st).
Habits: Four specimens were collected on the lagoon side
of the northern end of Chinimi Island, Eniwetok Atoll.
Specimens were found eating the orange sponge, Prianos
phlox pp LAauBENFELS, 1954, which occurred in abun-
dance on the undersides of rocks at the collecting site.
Examination of feces revealed spicules of the same sponge
and laboratory feeding tests suggested that specimens of
Doriopsis viridis fed exclusively on P. phlox.

A 7mm specimen produced a light yellow, ribbon-like
egg mass, 60mm long and 1.5mm wide, with 3 whorls.
The egg mass contained approximately 72 ova per mm’,
Each ovum was enclosed by a capsule 150m to 162 in
diameter. The larvac hatch as free swimming veligers.
Remarks: Two color “varicties” of Doriopsis viridis are

Vol. 10; No. 2

THE VELIGER

aD:

Page 161

1

010mm

Figure 2

Doriopsis viridis
Lateral view of the left half row of radular teeth

1 = innermost lateral

proposed (Rispec, 1928, 1953) and Doriopsis pecten
CoLiincwoop, 1881, the blue variety, is synonymized with
D. viridis Pease, 1861, the green variety (ALLAN, 1947;
Basa & HaMaTANI, 1961). The two species are similar in
many external and internal morphological characters, but
when specimens of similar size are compared, specimens
of D. viridis have larger eggs, larger radular teeth and
fewer teeth per radular row than specimens of D. pecten.
The species are further distinguished by different feeding
habits (see below).

Specimens of Doriopsis viridis are reported from Tahiti
(Pease, 1861, 1871) and New Caledonia (Rissec, 1928,
1953).

Figure 3

Doriopsis viridis
Lateral view of the female duct system
sc = spermatocyst st = spermatheca ud = uterine duct
v = vaginal duct

26 = outermost lateral

Doriopsis pecten (CoLLINGwoop, 1881)

Description: The animal is figured and described by
CoLLincwoop (1881), ALLAN (1947) and Baba & Hama-
TANI (1961). The radular formula is given by BaBA &
Hamatani (1961). Hamatani (1961) provides a photo-
graph of specimens in association with encrusting blue
sponge colonies and descriptions of eggs and veliger larvae.
Synonymies are given and additional features of the exter-
nal and internal morphology of Doriopsis pecten are
described and figured by Kay & Younc (in preparation).
Habits: Four specimens (6mm to 11mm long) were
collected at Eniwetok Atoll. One specimen was found in
the quarry on Eniwetok Island in 2m depth and the
other specimens were collected from 0.5m depth on the
northern lagoonward side of Chinimi Island. They were
scen feeding on the blue sponge, Terpios aploos pE Lau-
BENFELS, 1954, which was fairly abundant under rocks
and in crevices in dead coral at both collecting sites.
Spicules of 7: aploos were found in the feces and gut
contents of freshly collected specimens. Specimens of
Doriopsis pecten in Hawaiian waters (unpublished data)
were also found to specifically eat T: aploos.
Remarks: The occurrence of mutually exclusive feeding
habits is confirmatory ecologic evidence that Doriopsis
pecten and D. viridis are distinct species and that the blue
coloration of D. pecten is specifically diagnostic.
Specimens of Doriopsis pecten are reported from Tai-
wan (CoLiincwoop, 1881), Japan (BABA & HAMATANI,
1961), New South Wales (ALLAN, 1947), and are found
in Hawaii (Kay & YouNG, in preparation) in addition to
New Caledonia (RisBEc, 1928, 1953), where they co-occur
with specimens of D. viridis.

Page 162

Hypselodoris tryoni (GARRETT, 1873)
(Text figures 4 to 7)

Goniodoris tryont GARRETT, 1873: 232; plt. 4
Chromodoris tryont (GarreTT), Bercu, 1877: 490; Bercu, 1884:
69

Description: Hypselodoris tryoni has an elongate smooth
body up to 60mm in length and 18mm in width (Figure
4). The convex mantle is expanded anteriorly as a hood,

Figure 4

Hypselodoris tryoni
Dorsal view of adult specimen

58

= = ome
0.10 mm

THE VELIGER

Vol. 10; No. 2

rounded posteriorly and held above the foot. The foot is
elongate, tapered and projected posteriorly beyond the
mantle edge. The background color of the mantle is cream
or light brown. The mantle is bordered and spotted with
purple maculae each of which is surrounded by a white
areola in turn encircled by a pale fawn ring. The foot is
white, margined with lilac and covered with spots similar
to those on the mantle. The rhinophores are rod-like, elon-
gate, finely lamellate, retractile into widely separated
sheaths, and brown with a white medial streak anteriorly.
The 14 to 16 branchial plumes, which encircle the posteri-
orly positioned anus, increase in arborization and decrease
in size posteriorly. They are erect, vibratory and white with
red-brown margins. The rhachides are quadrangular in
cross section and retractile into a common cavity.

The radular formula of a 60mm specimen is 94 x
58-058 (Figure 5). The innermost radular tooth is simply

Figure 6

Hypselodoris tryont
Elements of the buccal armature

bicuspid and 50 long. Within each row, denticulation
increases laterally to a maximum of six outer denticles and
tooth length increases centrally to 83. The buccal arma-

Figure 5

Hypselodoris tryont
Lateral view of the left half row of radular teeth

1 = innermost lateral

58 = outermost lateral

Vol. 10; No. 2 THE VELIGER Page 163

Figure 7

Hypsclodoris tryoni
Lateral view of the genital mass with an offset of a medial view of the female duct system
= ampulla a+m = albumin and mucous gland od = oviduct pr = prostate gland
ed = ejaculatory duct hd = hermaphroditic duct st = spermatheca

vd = vas deferens

sc spermatocyst
ud = uterine duct Vv = vaginal duct

Page 164

THE VELIGER

Vol. 10; No. 2

ture, which ranges in length from 45 to 50u, has singly
hooked tips and long sculptured bases (Figure 6).

A common orifice (Figure 7) is shared by the ejacula-
tory duct (ed), vaginal duct (v) and oviduct (od). No
cirral hooks line the ejaculatory duct. The elongate pros-
tate gland (pr) partially envelops the spermatheca (st).
The vaginal duct is short and non-convolute and passes
directly into the spherical spermatheca. A short duct passes
from the spermathecal end of the vaginal duct into the
small pyriform spermatocyst (sc). A long, tightly con-
volute uterine duct (ud) joins the junction of the prostatic
and ampullary ducts at the albumin and mucous gland
(a-+m).

Habits: Two specimens were found on dead coral in the
lagoon of Palmyra Atoll. The structure and arrangement
of the digestive system indicates that this species is a
rasping sponge-feeder. The food was not determined.
Remarks: On the basis of specimens collected in Zanzibar,
Exiot (1904) suggested that the dorid Chromodoris aure-
opurpurea CoLtincwoop, 1881, from the China coast is
a variety of Hypselodoris tryoni. However, C. aureopur-
purea has characters which are sufficiently distinctive
so that this species should be regarded separately from H.
tryoni as originally suggested by CoLLINGwoop.

Specimens of Hypselodoris tryoni are reported from the
Society Islands (Garrett, 1873) and the Philippines
(BercH, 1877).

Hypselodoris kayae Younc, new spccies

(Text figures 8 to 11)

Description: Hypselodoris kayae has a smooth, elongate
body with maximal dimensions of 10mm length and 3mm
width (Figure 8). The mantle is convex and held above

21

Figure 8

Hypsclodoris kayae
Dorsal view of adult specimen

the foot. The elongate, white foot tapers and extends
posteriorly to the mantle, which is light lilac, edged with
irregularly spaced rose red spots. The rhinophores are
rod-like, white with rose red tips and retractile into closely
spaccd sheaths; there arc 8 to 9 lamellae. The 7 branchiae
are in a circlet about the posterior mid-dorsal anus; they
are simply pinnate, white, tipped with rose red, smaller
posteriorly and retractile into a common cavity.

The radular formula of a 10 mm specimen is 28 x 21:0:21
(Figure 9). The innermost radular tooth, 25 long, is bi-

1

0.050mm

Figure 9

Hypselodoris kayac
Lateral view of the Icft half row of radular tecth

1 = innermost lateral

21 = outermost lateral

Vol. 10; No. 2

cuspid with a single outer denticle. The teeth increase
in denticulation and size towards the center within each
row to a maximal number of 4 outer denticles and a
length of 46. The buccal armature consists of small bifid
hooks, 12 to 16 long, with broad sculptured bases (Figure
10).

B
<<

y

I eae |
0.025 mm

Figure 10

Hypselodoris kayae
Elements of the buccal armature

The ejaculatory duct (ed), vaginal duct (v) and ovi-
duct (od) have separate, external orifices (Figure 11).

THE VELIGER

Page 165

No cirral hooks line the ejaculatory duct. The ejaculatory
duct, vas deferens (vd) and prostate gland (pr) are
loosely convolute and merge imperceptibly. The vaginal
duct joins the uterine duct (ud) and forms a common
vaginal-uterine duct which in turn leads to the junction
of the spermatheca (st) and spermatocyst (sc). The
oblong spermatocyst connects directly with the spherical
spermatheca. The long, loosely convolute uterine duct
passes from the vaginal duct and merges with the ampul-
lary and prostatic ducts at the albumin and mucous
gland (a-+-m).
Habits: Two specimens, 6mm and 10mm long, were col-
lected from underneath a rock at 0.5m depth on the
northern lagoonward side of Chinimi Island, Eniwetok
Atoll. They were found eating a rose-red sponge, Aplysilla
glacialis DE LAUBENFELS, 1951, which was a common
encrusting sponge upon which the dorids were inconspic-
uous. Spicules of this sponge were recovered from the
feces of both specimens. The specimens also ate A. glacialis
in laboratory aquaria.

An egg mass produced by the 10mm specimen was a
doubly whorled white ribbon, 15mm long and 1.7mm
wide. The ova, 75y1 in diameter, were individually enclosed

Figure 11

Hypselodoris kayac
Lateral view of the genital mass

st
a+m
nega PR TRE eT ene
a ampulla a+m = albumin and mucous gland

ed ejaculatory duct hd = hermaphroditic duct

od = oviduct
st = spermathcca

pr = prostate gland
ud = uterine duct

sc = spermatocyst
v= vaginal duct

vd = vas deferens

Page 166

in capsules, 100 in diameter. There were approximately
320 ova per mm’ of egg mass. The larvae hatch as free
swimming veligers.

Type locality: Lagoon, north side of Chinimi Island,
Eniwetok Atoll, Marshall Islands, 11° 30’ N. 162° 15’ E.
Type material: The holotype and the paratype (slide of
radular tecth) are deposited in the Bishop Museum,
Honolulu, Hawaii, where they are registered as No. 8918.
The species is named for Dr. E. Alison Kay.

Diagnosis: Hypselodoris kayae is easily distinguished from
other species of Hypselodoris by the light lilac mantle
edged with rose red spots, the white foot, and the white
rhinophores and branchiae tipped with rose red.
Remarks: Following OpHNER (1957) this new species is
placed in the genus Hypselodoris Stimpson, 1855 on the
basis of the predominantly bicuspidate condition of the
radular teeth. Further characters shared by H. kayae, H.
tryoni and 5 species of Hypselodoris from Hawaii (Kay &
Younc, in preparation) are the broad, sculptured bases
of the buccal armature and the united condition of the
uterine and vaginal ducts at the spermatheca.

Chromodoris geometrica RisBEC, 1928
(Text figures 12 to 15)

Chromodoris geometrica RisBxc, 1928: 148 - 151, fig. 41, plt. VI, 10;
Rispec, 1953: 70; figs. 31, 32
Glossodoris geometrica (RisBEc) , ALLAN, 1947: 441; fig. 14, plt. 41

Description: The single specimen of Chromodoris geo-
metrica collected has an oblong body, 22mm long and 8
mm wide (Figure 12). The mantle flares about the foot,
expands anteriorly as a hood and narrows posteriorly as
a rounded projection. The foot broadens antcriorly into
two lateral projections and tapers posteriorly beyond the

28

0.10 mm

THE VELIGER

Vol. 10; No. 2

Figure 12

Chromodoris geometrica
Dorsal view of adult specimen

margin of the mantle which is white and fluted at the edge.
Rays of purple radiate peripherally from two medial,
parallel, purple bands, which in turn join anteriorly to the
rhinophores, posteriorly to the branchiae and centrally at
mid-body length. Enclosed by the purple reticulations are
white pustules upon a light yellow background. The foot
is white ventrally, light yellow dorsally and bordered with
a purple margin. The rhinophores are fusiform, elongate,
finely lamellate and retractile into closely spaced sheaths.
The peduncles are white and the lamellae are yellow. The

Figure 13
Chromodoris geomcetrica
Lateral view of the left half row of radular tecth

1 = innermost lateral

28 = outermost lateral

Vol. 10; No. 2

7 branchiae are simply pinnate with yellow rachides and
transparently white pinnae; they form a circlet about the
anus and decrease in size posteriorly. The rachides are
quadrangular in cross section and retractile into a common
cavity, but they are not vibratile as in Hypsclodoris tryoni.
The radular formula of a 22 mm specimen is 43 x 28:0:28
(Figure 13). The radular teeth are unicuspid with 2 to 9
outer denticles and range from 53 to 79u in length.
Length and denticulation of the teeth increase centrally
within each row. The outermost teeth are denticulate only
at their tips. The buccal armature consists of simple bifid
hooks, 32 to 39 long, with smooth bases (Figure 14).
The ejaculatory duct (ed), vaginal duct (v) and ovi-
duct (od) have separate external openings (Figure 15).
There are no cirral hooks in the ejaculatory duct. The
prostate gland (pr) is long and tightly convoluted. The
vaginal duct merges in a common duct with the uterine
and spermatocystic duct which in turn passes into the

THE VELIGER

Page 167

( ease
0.025mm
Figure 14

Chromodoris geometrica
Elements of the buccal armature

spherical spermatheca (st). The spermatocyst (sc) is pyri-
form and approximately half the size of the spermatheca.

Figure 15

Chromodoris geomcetrica

Lateral view of the genital mass

= ampulla a+m = albumin and mucous gland
ed = ejaculatory duct hd = hermaphroditic duct

pr
= oviduct pr = prostate gland sc = spermatocyst
st = spermatheca ud = uterine duct v= vaginal duct

vd = vas deferens

Page 168

THE VELIGER

Vol. 10; No. 2

The uterine duct (ud) enters the albumin and mucous
gland (a-+-m) separately from the ampullary and pros-
tatic ducts.

Habits: One specimen was found under an intertidal
rock on the northern lagoonward side of Chinimi Island,
Eniwetok Atoll. The food was not determined but the
structure and form of the digestive tract suggest that this
species is a rasping sponge-feeder.

Remarks: The occurrence of this specimen at Eniwetok
Atoll is the northernmost record for Chromodoris geomet-
rica, which was previously reported only from New Cale-
donia (Rispec, 1928, 1953) and New South Wales
(AttAN, 1947).

Chromodoris lilacina (Goutp, 1852)

Doris lilacina Gou.p, 1852: 297

Doris amabilis KELAART, 1859: 294 - 295

Chromodoris porcata BERGH, 1888: 831 - 833

Chromodoris amabilis (KELAART), Ex1oT, 1906: 642, plt. XLII, fig. 1
Glossodoris amabilis (KELAART), PruvoT-Fot, 1951: 84

Description: External morphology and coloration of one
specimen are described by Goutp (1852). The repro-
ductive system and the radular teeth in addition to
further descriptions of external morphology are presented
and figured by Kay « Younc (in preparation).
Habits: Two specimens (8 and 14mm long) were collected
on separate occasions from 0.5m of water at the lagoon
side of the northern end of Chinimi Island, Eniwetok
Atoll. The food was not determined. Specimens of Chro-
modoris lilacina in Hawaii feed on Mycale maunakea
DE LAUBENFELS, 1951, a sponge not reported from Eni-
wetok (unpublished data).
Remarks: The species described by Ketaart (1859) as
Doris amabilis and suggested by Pruvot-Fot (1951) to
be a glossodorid, probably is synonymous with Chromo-
doris lilacina. Exrot (1906) figured the original of Ket-
AART and suggested that it may be a chromodorid. ELioT
further suggested the synonymy of C’. porcata (BERGH,
1888) with C. amabilis.

Specimens of Chromodoris lilacina are reported from
Ceylon (Kexaart, 1859), Mauritius (Bercy, 1888) and
Hawaii (Gou Lp, 1852).

Chromodoris albopustulosa (PEASE, 1860)

Doris albopustulosa PEasE, 1860: 30

Chromodoris albopustulosa (PEASE), BERcH, 1884: 70

Glossodoris albopustulosa (PEASE), PRuvot-Fot, 1947: 108; Pruvot-
Fo, 1951: 83

Descripiton: The external characters of Chromodoris
albopustulosa are described by Pease (1860). External
morphology, reproductive system, and radular teeth are

further described and figured by Kay « Younc (in prepa-
ration).

Habits: One 10mm specimen was collected under a rock
in 2m of water in the quarry on Eniwetok Island. The
structure and form of the alimentary tract suggest that
this specics is a rasping sponge-feeder, although the food
was not determined.

Remarks: The occurrence of this specimen at Eniwetok
Atoll is the only definite record of Chromodoris albopus-
tulosa outside of the Hawaiian Islands (PEAsE, 1860) as
Bercu (1884) merely records it from the Pacific.

Jorunna tomentosa (Cuvier, 1804)

Description: A synonymy is given and external and inter-
nal characters are discussed by Pruvot-Fot (1954). A
specimen is figured by Pruvot-Fot (1953). The repro-
ductive system with its characteristic stylet is figured by
BercH (1880) and Pruvot-Fot (1954). Additional de-
scriptions of external and internal morphological charac-
ters are given and the radular teeth and the reproductive
system are figured by Kay & Younc (in preparation) from
specimens collected from the Hawaiian Islands.
Habits: One 18mm specimen of Jorunna tomentosa was
collected under a rock in 0.5m of water on the northern
lagoonward side of Chinimi Island. This species is reported
to be a sponge-feeder in Europe (MILter, 1961; THomp-
son, 1964) and Hawaii.

Remarks: Jorunna tomentosa is reported by PRuvot-Fo.
(1954) to be distributed from 65° N latitude in Scandi-
navia down the coasts of Great Britain and France and
into the Mediterranean Sea. This species is the only dorid
collected which is not restricted in distribution to the
Indo-West-Pacific faunal region. The occurrence of speci-
mens of J. tomentosa in European waters as well as in
Pacific waters suggests a cosmopolitan distribution.

DENDRODORIDIDAE

Dendrodoris nigra (Stimpson, 1855)

Descriptions A synonymy is given and external morpho-
logical characters of adult specimens are described by
Marcus & Burcu (1965). External and internal mor-
phological characters of this common Indo-West-Pacific
dorid throughout different stages of its growth are
described and figured by Kay & Younc (in preparation).
Habits: Twelve specimens of Dendrodoris nigra (4 to 30
mm long) were collected during the 3 week period: 9
from the quarry on Eniwetok Island and 3 from the
northern lagoonward side of Chinimi Island. All specimens
were black with white spots except for the smallest which

Vol. 10; No. 2

was orange and similar in coloration to juvenile specimens
of D. nigra in Hawaii. One specimen (12mm preserved
length) was also collected on a head of Porolithon sp.
inside the fringing reef at Johnston Island in August,
1965.
Remarks: Although there were no direct observations of
feeding, fluorescence-microscopical examinations of mate-
rial from the gut and feces indicated a complete absence of
organic matter of plant origin with the exception of several
errant diatoms. Lacking radular teeth, these specimens are
probably sucking sponge-feeders as are those in Hawaii.
Dendrodoris nigra is not only one of the most abundant
dorid species in Hawaii and at Eniwetok, but also one of
the most widely distributed and frequently recorded dorids
in the Indo-West-Pacific. This is the only species of dorid
collected which has been previously recorded from Eni-
wetok. Marcus « Burcu (1965) report 13 specimens of
D. nigra collected on Parry and Japtan Islands, Eniwetok
Atoll, during Spring 1960.

PoLYCERIDAE

Gymnodoris citrina (BERGH, 1877)
(Text figures 16 to 18)

Trevelyana citrina BErcH, 1877: 440 - 443; plt. 56, figs. 18 - 25

Description: Gymnodoris citrina has a limaciform body
measuring 6 to 24mm long (Figure 16). The dorsum,
which is cream or light yellow with small orange tipped
pustules, is sparsely scattered with irregularly shaped
epidermal spicules which are 120u to 250 in length and
pointed at one or both ends. Anteriorly there is a broad
cephalic hood edged with orange tipped serrations. In
dorsal view the lobiform oral tentacles project anterolat-
erally beyond the edge of the cephalic hood, the genital
orifice protrudes from the right side of the body immedi-
ately anterior to the level of the branchiae and the body
tapers posteriorly with the elongate foot. The rhinophores,
positioned laterally in the cephalic hood, are club-like in
shape, bear 15 to 20 light yellow lamellae and may be
tipped with orange. The 8 to 9 branchiae, in a horseshoe-
shaped placement about the mid-dorsal anus, are simply
pinnate, flattened laterally and smaller posteriorly; they
are light yellow and often tipped with orange. Character-
istically, but macroscopically visible only in large speci-
mens, is a slightly raised V-shaped area edged with
shallow, orange tipped serrations projecting and tapering
posteriorly from the level of the branchiae to the posterior
one-third of the body.

The radular formula of 3 specimens (6, 10, and 24
mm) is 10-18 x 14-30:0-14-30 (Figure 17). The first
lateral tooth is simply hamate and larger than the succes-

THE VELIGER

Page 169

Figure 16

Gymnodoris citrina
Dorsal view of adult specimen

sive teeth in cach row, varying from 142u to 390 long in
6mm and 24mm specimens respectively. The second
lateral tooth is awl-shaped and broadly based, whereas
the following teeth are successively narrower and shorter.
At the buccal lip are paired unarmored cuticular
thickenings.

Three ovate, orange-red ovotestes lie anteroventral to
the digestive gland. Components of the genital mass were
not discernible in the specimens dissected, excepting the
long and tightly convoluted male duct system. Numerous
cirral hooks, 74 to 8.5u long, line the lumen of the
ejaculatory duct (Figure 18).

Habits: Six specimens were collected at Eniwetok Atoll.
One was found in the quarry on Eniwetok Island and 5
on the northern lagoonward side of Chinimi Island under
rocks in 0.5m of water. An egg mass containing late stage
veliger larvae, 150 to 162 in diameter, with an opistho-
branch larval shell type 1 (terminology according to
Tuompson, 1961) was the only identifiable food found in
the midgut of one specimen of Gymnodoris citrina collec-
ted in the field. One conclusion resulting from the category
of this larval shell type is that the veligers are not of
aeolid origin, as aeolids have a different type of larval

Page 170

THE VELIGER

Vol. 10; No. 2

0.10 mm

Figure 17

Gymnodoris citrina
Lateral view of right half row of radular teeth

1 = innermost lateral

2 = second lateral

14 = outermost lateral

shell from that of other opisthobranchs. A 10mm speci-
men devoured a 6mm specimen in the laboratory but this
occurrence was probably induced by unnaturally crowded
conditions in the aquarium.

Remarks: Since the name Trevelyana (== Gymnodoris)
citrina was first proposed by Bercu (1877) for an 11mm
specimen from the Palau Islands, it has been put into the
synonymy of Gymnodoris bicolor (ALDER & HANcock,
1866) by a number of workers (e. g. Rispec, 1953;
MacNag, 1958; Basa, 1960). Comparison of internal and
external characters of G. bicolor and G. citrina indicates
that the two species are distinct.

oe:

a Sa

0.010 mm

Figure 18

Gymnodoris citrina
Cirral hooks

Much of the difficulty and confusion which have arisen
concerning Gymnodoris bicolor and G. citrina are appar-
ently because the first lateral tooth is larger in relation to
the succeeding outer lateral teeth in both species. A com-
parison also reveals that the innermost laterals of G. citrina
are much larger than those of G. bicolor in similar sized
specimens. In fact, the 120m length of the innermost
lateral tooth in a 22mm specimen of G. bicolor from
Hawaii (Kay « Younc, in preparation) is even shorter
than the 142u length of the similar tooth in a 6mm
specimen of G. citrina from Eniwetok.

The broad base and awl-shaped cusp of the second
lateral tooth is a useful specific character for Gymnodoris
citrina as stated by BercH (1877). The 180m length of
the innermost lateral tooth from the 11mm specimen of
Bercu also agrees favorably with the 165 length of the
similar tooth from a 10mm specimen in this study.

Rispec (1928) described two species of Trevelyana
(= Gymnodoris), T. perlucens and T. suggens, from New
Caledonia. Both were later put into the synonymy of T:
bicolor (Rispec, 1953). In the latter work, RisBec de-
scribes “un nouvel exemplaire,” literally “a new speci-
men,” which he states is intermediary in certain characters
to the T. perlucens and T. suggens “forms” of T. bicolor.
This new specimen described by RisBec (1953)- exhibits
many diagnostic characters of G. citrina. Among the
common ones are: “... papilles oranges disposées en V
en arriére des branchies ect les encadrant ... plaques

Vol. 10; No. 2 THE VELIGER Page 171
cornées incolores situées en avant de la rotella ..° La 1™ AEOLIDACEA

latérale a une cuspide de 0.24mm de long ... La glande

génitale est formée de trois masses arrondies d’un rouge FAVoRINDAE

foncé ... le canal déférent est plus long.” (Rispec, 1953,

pp. 100-101). The specimen figured by RisBec (1953,
fig. 57) bears a greater resemblance to G. citrina than to
G. bicolor.

Although only one specimen of Gymnodoris citrina has
been reported from the Palau Islands (BERcH, 1877), the
species is likely to be further distributed throughout the
Indo-West-Pacific faunal region. Because past species de-
scriptions have not always been sufficient to enable one
to distinguish between G. citrina and G. bicolor, the
distributions of both species remain uncertain.

VAYSSIEREIDAE

Okadaia elegans Basa, 1930

Description: Synonymies are given and certain aspects of
the biology of Okadaia elegans, including descriptions of
the external and internal morphology, are discussed by
Basa (1937).

Habits: Numerous specimens of Okadaia elegans were
found in the shallow tide pool at the northern lagoonward
side of Eniwetok Island. These minute limaciform dorids
with a maximal length of 4mm were found feeding on
spirorbid polychaetes on the undersides of rocks in the
area. In penetrating the calcareous exoskeleton of a spir-
orbid polychaete, O. elegans bores a hole through the tube
with its radular teeth.

The egg masses of Okadaia elegans were deposited as
flattened gelatinous masses on the undersides of rocks at
the collecting site and on the sides of aquaria in the
laboratory. Juvenile forms, 0.55mm to 0.70mm long,
emerge from individual capsules contained in each egg
mass within 10 days after oviposition at 26° to 27° C and
feed upon spirorbid polychaetes in a similar manner as
the adult forms.

Remarks: Previously reported from Japan (Basa, 1930,
1931, 1937) and commonly found in Hawaii, this species
is likely to be found throughout the Indo-West-Pacific
faunal area if a careful examination is made of substrata
colonized by spirorbid polychaetes. Specimens of Okadaia
elegans are likely to be overlooked because of their small
sizes. Species closely allied to O. elegans have been
reported from New Caledonia (RisBec, 1928), New Zea-
land (Ratpu, 1944), and Formosa (CoLLincwoon, 1881).

Herviella mietta Marcus & Burcu, 1965

Description: Marcus & Burcu (1965) discuss external
and internal morphological characters, and figure an
adult specimen, a radular tooth, a jaw and a penial stylet
in their original description of Herviella mietta from 16
specimens from Eniwetok.

Habits and Remarks: Twelve specimens of Herviella
muetta (5mm to 10mm long) were collected intertidally
on the northern lagoonward side of Eniwetok Island, the
same location where 13 specimens were collected in 1960
(Marcus & Burcn, 1965). A single 8mm specimen was
also collected intertidally under a rock on the western
lagoon side of Igurin Island, the only nudibranch collected
at this island.

Numerous egg masses of Herviella mietta were depos-
ited in the collecting area on the undersides of rocks and
even on the shells of specimens of the prosobranch, Ceri-
thium sejunctum IrEDALE, 1929. The larvae pass the
veliger stage within the egg mass and each emerges in
the crawling stage with the larval shell yet attached
within 6 days after oviposition at 28° to 29°C. Within
24 hours after hatching the larval shells are dropped. The
juvenile forms bear 2 simple cerata.

Unreported in the description of the species is the
feeding of Herviella mietta on the eggs of the prosobranch
gastropod Cerithium sejunctum. This association is likely
to be a seasonal occurrence during the spawning of C.
sejunctum and the aeolid is probably a hydroid-feeder
during the remainder of the year. Additional evidence
that another food is eaten by these aeolids is that newly
hatched juveniles of H. mietta do not feed on the eggs
of C. sejunctum. Variation from a hydroid diet by aeolids
is reported in England for Calma glaucoides (ALDER &
Hancock, 1855) which feeds on demersal fish eggs
(Evans, 1922; Rowett, 1946) and in Hawaii where an
unidentified species of aeolid feeds on the eggs of the
opisthobranch gastropod Aplysia juliana Quoy & Gat-
MARD, 1832 (E. Alison Kay, personal communication).

Specimens of Herviella mietta were the only nudi-
branchs eaten by a butterfly fish (Chaetodon auriga)
which was kept in a large sea water tank in the Eniwetok
Marine Biological Laboratory. All other nudibranchs
collected were rejected after they were initially taken into
the mouth of the fish. The nudibranchs were not visibly
harmed by such treatment and thereafter were not taken
in by the fish after an initial investigation. Specimens of
H. mietta were readily eaten by the fish on every occasion,

Page 172

even when the fish was fed to apparent satiation with
other food. This is one of the few occasions that possible
predation on aeolid nudibranchs by fish has been demon-
strated (see Epmunps, 1966).

Herviella claror Burn, 1963

Description: Marcus & Burcu (1965) expanded on the
description of Herviella claror Burn, 1963 and figured an
adult specimen, a radular tooth and a jaw on the basis of
3 specimens collected from Eniwetok.

Habits and Remarks: Two specimens (13mm and 15mm
long) of Herviclla claror were found in the quarry on
Eniwetok Island not far from the north end of the same
island where 3 specimens were collected in 1960 (Marcus
& Burcu 1965). This species was described from a spcci-
men collected in New South Wales (Burn, 1963).

SUMMARY

Nine new geographic records, 3 confirmatory geographic
records and one new species, Hypsclodoris kayae, repre-
senting 2 superfamilies and 5 families of the Nudibranchia
are reported from 3 atolls (Eniwetok, Palmyra and
Johnston Island) in the Central and West-Central Pacific.
With one exception of a presumed cosmopolitan species,
Jorunna tomentosa, all the nudibranchs are limited in
distribution to the Indo-West-Pacific faunal region. Six
species co-occur in Hawaii and Eniwetok.

Only 1 of the 7 species of Doridacea previously reported
from Eniwetok was found. This difference probably
reflects the seasonality in occurrence of dorids in shallow
water. The 2 species of Aeolidacea previously reported
from Eniwetok are verified.

The foods of 3 sponge-feeders, Doriopsis viridis, D. pec-
ten, and Hypselodoris kayae, and one polychacte feeder,
Okadaia elegans, are reported. Two occurrences of preda-
tion by nudibranchs on egg masses of gastropods are
given. It is verified that specimens of Herviella mietta feed
on egg masses of a prosobranch gastropod and suggested
that specimens of Gymnodoris citrina feed on egg masses
of an unknown opisthobranch gastropod.

Three general types of larval development are shown
by the nudibranchs collected. The larvae of Doriopsis
viridis and Hypselodoris kayae emerge from egg masses as
free swimming veligers. The larvae of Herviella mietta
hatch as crawling larvae that drop their shells within 24
hours. The larval stages of Okadaia elegans are completed
within egg masses from which emerge juveniles that are
capable of adult feeding habits.

THE VELIGER

Vol. 10; No. 2

LITERATURE CITED

ALpER, JosHuUA & ALBANY HANcock

1855. A monograph of the British nudibranchiate Mollusca.
Ray Soc. London, 438 pp.

1866. Notice of a collection of nudibranchiate Mollusca made
in India by Walter Eliot, Esq., with descriptions of several new
genera and species. Trans. Zool. Soc. London 5: 113 - 145

ALLAN, Joyce K.

1947. | Nudibranchia from the Clarence River Heads, North
Coast, N.S. W. Rec. Austral. Mus. 21: 432 - 463

Basa, KikuTARO

1930. Studies on Japanese nudibranchs. 2. _- Venus 2: 47 - 50

1931. A noteworthy gill-less holohepatic nudibranch Okadaia
elegans Basa, with reference to its internal anatomy. Annot.
Zool. Japon. 13: 63 - 83

1937. Contribution to the knowledge of a nudibranch, Okadaia
elegans Basa. Japan. Journ. Zool. 7: 147 - 190

1960. The gencra Gymnodoris and Nembrotha from Japan
(Nudibranchia - Polyceridae) . Pub. Seto Mar. Biol. Lab.
8: 71-74
Basa, KrkuTARO & Iwao HaMaTANI
1961. On two species of Doriopsis (Syn. Ctenodoris) from
Japan (Nudibranchia - Dorididae). Pub. Seto Mar. Biol.
Lab. 9: 63 - 65

Bercy, Lupwic SopHus Rupo.teu
1877. | Malakologische Untersuchungen. in (C. SEMPER,
Reisen im Archipel der Philippinen 2 (11): 377 - 645; plts. 54
to 57
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

1884. Report on the Nudibranchia.
Challenger Exped. 10: 1 - 154; plts. 1 - 14

1888. | Malakologische Untersuchungen. in C. SEMPER,
Reisen im Archipel der Philippinen. 2 (16): 831 - 833

Rep. Sci. Results

Burn, ROBERT
1963. Descriptions of Australian Eolidacea (1. The genera
Catriona and Hervicella). Journ. Malac. Soc. Austral. 7:
12-20
CoLLINGwoop, CUTHBERT
1881. | On some new species of nudibranchiate Mollusca from
the eastern seas. Trans. Linn. Soc. London 2: 123 - 140
Cuvier, Georcrs LEopoLp CHreETIEN FD.

1804. | Mémoire sur le genre Doris. Ann. Mus. Nat. Hist.

Natur. 4: 447 - 473

DE LAUBENFELS, Max WALKER

1951. The sponges of the Island of Hawaii. Pacific Sci.
5: 256 - 271
1954. The sponges of the West-Central Pacific. Oregon
State Monogr. 7: 306 pp.
Epmunps, Matco_m
1966. Protective mechanisms in the Eolidacea (Mollusca Nudi-

branchia) . Journ. Linn. Soc. London 46: 27 - 73

Vol. 10; No. 2

Exot, Crarzes N. E.

1904. On some nudibranchs from East Africa and Zanzibar. IV.
Proc. Zool. Soc. London 1904 (vol. 1) : 380 - 406

1906. On the nudibranchs of southern India and Ceylon,
with special reference to the drawings by Kelaart and the
collections belonging to Alder_and Hancock preserved in the
Hancock Museum at Newcastle-on-Tyne. Proc. Zool. Soc.
London 1906 (vol. 2): 636 - 691

Evans, T. J.

1922. Calma glaucoides: A study 2 adaptation.

Journ. Micr. Sci. 66: 439 - 455
Garrett, ANDREW G.

1873. Description of a new species of Goniodoris. Proc.

Acad. Nat. Sci. Philadelphia 25: 232
Goutp, Aucustus ADDISON

1852. Mollusca and shells. Vol. 12. United States Exploring
Expedition etc. U.S. Govt., Philadelphia, 510 pp.

Quart.

HamatanI, Iwao
1961. Notes on the veligers of Japanese opisthobranchs (4).
Publ. Seto Marine Biol. Lab. 9 (2): 353 - 361
KELAART, Epwarp FREDRICK
1859. Descriptions of new and little-known species of Ceylonese
nudibranchiate mollusks. Ann. Mag. Nat. Hist. 3: 291-304,
488 - 496
MacnaeE, WILLIAM

1958. The families Polyceridae and Goniodorididae (Mollusca,
Nudibranchia) in southern Africa. Trans. Roy. Soc. South

Africa 35: 341 - 372
“W—.

Marcus, Ernst
1965. Some opisthobranchia from Micronesia.

3: 263 - 286

Marcus, Ernst « JoHN Bayarp Burcu

Malacologia

1965. Marine euthyneuran Gastropoda from Eniwetok Atoll,
Western Pacific. Malacologia 3: 235 - 262
MiLuer, MicHaer C.

1961. Distribution and food of the nudibranchiate Mollusca
of the South of the Isle of Man. Journ. Anim. Ecol. 30:

95 - 116
Opune_r, Nirs Hyatmar SS
1957. Chromodoris contra Glossodoris, a systematic-nomen-
clatorial controversy. Proc. Malacol. Soc. London 32: 250
to
Pease, WILLIAM HarPER .
1860. Descriptions of new species of Mollusca from the
Sandwich Islands in the collection of Hugh Cuming. Proc.

Zool. Soc. London 28: 18 - 36

THE VELIGER

Page 173

Pease, WILLIAM Harper
1861. Descriptions of new species of Mollusca from the Pacific
Islands. Proc. Zool. Soc. London for 1861: 242 - 247
(September 1861)
1871. Descriptions of nudibranchiate Mollusca, inhabiting
Polynesia. Amer. Journ. Conch. 6: 299 - 305
Pruvot-Fo., ALIcE
1947. Les Opisthobranches de W. Harper Pease, Révision.
Journ. de Conchyl. 87: 96 - 114

1951. Révision du genre Glossodoris EHRENBERG.
Conchyl. 91: 76 - 164

Journ. de

1953. Etude de quelques opisthobranches de la céte Atlantique
du Maroc et du Sénégal. Trav. Inst. Sci. Chér. 5: 1 - 93

1954. Mollusques opisthobranches. Faune de France.
Paris, Lechevalier. 58: 460 pp.; 173 figs.; 1 plt.

Quoy, Jean RENE ConsTANT, & JosEPH PauL GAIMARD
1832-1834. Voyage de la Corvette l’Astrolabe exécuté par ordre du
Roi, pendant les années 1826-1829, sous le commandement de
H. J. Dumont d’Urville; Mollusques. Zoologie, Paris. 2:
95 plates

Ravpu, Patricia M.

1944. Pellibranchus cinnabareus, a new genus and species of
non-pelagic nudibranch of the family Phyllirhoidae.
Roy. Soc. New Zealand 74: 24 - 31

RIsBEc, JEAN
1928. Contributions 4 létude des nudibranches Néo-Calé-
doniens. Faune Colon. Frang. 2(1): 1-238; plts. 1-16
1953. | Mollusques nudibranches de la Nouvelle Calédonie.
Faune l’union Frang. 15: 1 - 189

Rowetrt, HELEN G. Q.

1946. A comparison of the feeding mechanisms of Calma

glaucoides and Nebaliopsis typica. Journ. Mar. Biol. Assoc.
U. K. 26: 352 - 357

Trans.

Stimpson, WILLIAM
1855. Descriptions of some new marine invertebrates from the
Chinese and Japanese Seas. Proc. Acad. Nat. Sci. Phila-
delphia 7 (10): 375 - 384
TuHompson, 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

1964. Grazing and the life cycles of British nudibranchs.
Brit. Ecol. Soc. Symp. 4: Grazing in terrestrial and marine
environments, ed. D.J.Crisp. Blackwell, Oxford: 275 - 297

Page 174

THE VELIGER

Vol. 10; No. 2

A Re-Interpretation of the Sand-Pipes Described by ApEGOoKE

BY

JOHN W. EVANS

Department of Biology

Memorial University of Newfoundland, St. John’s, Newfoundland, Canada

(Plate 17)

ADEGOKE (1966) DESCRIBED A NUMBER Of silicified sand
and mud-filled pholad burrows in Upper Miocene strata
from San Luis Obispo County, California. The rock into
which the burrows were bored belongs to the Monterey
Formation and is described by ADEGOKE as a “hard, dull,
greenish-brown, siliceous and cherty mudstone.” The Mon-
terey Formation is overlain by a “whitish-gray, fine-
grained and tuffaceous silty sandstone” of the Pismo
Formation, which fills the burrows, forming the “sand-
pipes.”

As I have recently been engaged in the study of the
ecology of living pholads, I would like to discuss his
findings and to offer some alternative interpretations.

ADEGOKE tentatively assigned the fossil pholad to the
genus Chaceia (?). While on the available evidence this
identification cannot be ruled out, it should be pointed out
that the characters used to identify the genus are not
definitive.

One of the principal characters used was the short
sub-spherical outline of the valves. Unfortunately, ApE-
GOKE does not give a numerical value for the ratio: valve
length to depth. Measurements made by TurNER (1955)
show that adult Chaceia ovoidea (Gout tp, 1851) with a
length to depth ratio range of 1.46 - 1.88 could be
confused with a number of the Pacific Coast pholads on
the basis of this character. This is especially truce of
Penitella fitchti TURNER, 1955, with a ratio ranging from
1.1 to 1.9 and Zirfaea pilsbry: Lowe, 1931, with ratios
from 1.5 to 2.1 (Turner, 1954). In the vicinity of Coos
Bay, Oregon, adult Penitella penita (Conran, 1837) have
been collected by the author with a length to depth ratio
ranging from 1.55 to 2.56. Young, actively boring animals

are significantly shorter than the adults, with the ratio
ranging all the way down to 1.0 in the newly settled
forms.

Another character used by ADEGOKE to identify Chaceia
was the thinness of the walls of the valves. Because valve
thickness is a highly variable characteristic even within a
single species, it is not a useful characteristic for identifying
pholad genera. The valve thickness of Penitella penita
varies from very thin in soft rock to very thick in hard
rock (Evans: A, in press).

The presence of a pedal gape is characteristic of all
young Pholadidae. The callum which partially or wholly
closes the pedal gape of adult Martesiinae is usually very
delicate and is the first portion of the shell to dissolve or
break after death. Thus the numerous holes in the anterior
region of the burrows could either be explained by the
absence of a callum or by a broken callum.

The observation that “the faint lamellar ribs are not
unlike those of Chaccia” docs not help to identify the
fossils because the lamellar ribs of all Martesiinae are
basically very similar. On the other hand, within a single
species (Penitella penita) the morphology of the lamellar
ribs can vary considerably depending on the type of sub-
strate inhabited by the animal (Evans: A, in press and B,
in preparation).

ADEGOKE described two types of burrows: elongate
conical burrows with tapering necks and bulbous bottoms
(similar to Figure 1, Plate 17) and small spherical bur-
rows with or without short slender necks (similar to Figure
72, exe i17))).

Most pholads drill conical burrows, similar to the first
type. With the present state of our knowledge about pholad

Explanation of Plate 17

Figure 1: Penitella penita valves and cast of burrow

Figure 2: Nettastomella rostrata valves and cast of burrow

Note: Base of burrow incomplete in both cases

TuHE VELIGER, Vol. 10, No. 2 [Evans] Plate 17

Figure 1

Figure 2 Scale 33:1

Vol. 10; No. 2

THE VELIGER

Page 175

burrow morphology, no clue as to the former occupant
can be obtained from this information alone.

With respect to the spherical slender-necked burrows
(ApEGOKE, plate 21, figures 8 and 5) a more informed
guess can be made about the identity of the original
occupant. This burrow shape is very different from the
usual pholad burrow and has only been observed by the
author in one species, Nettastomella rostrata (VALENCI-
ENNES, 1846). The cast of a burrow of a living N. rostrata
from near Fossil Point in Coos Bay, Oregon, is illustrated
in Plate 17, Figure 2. Note the similarity both of size and
shape to the burrows shown by AbEGOKE. It is almost
certain that the same species did not form both of the
burrow types described by Aprcoxe. The burrow of
Penitella penita varies somewhat with age and signifi-
cantly with differences in substrate hardness; nevertheless
the conical shape is always retained (EvANs, in prepar-
ation C). An examination of figure 9, plate 21 of ApE-
GOKE’s paper will show that there is a conical burrow on
the left side of the hand specimen which is considerably
smaller than either of the spherical burrows in figures 5
and 8; thus his contention that the spherical burrows are
formed by young animals is open to question.

ADEGOKE states that Chaceia ovoidea, Penitella penita,
Penitella gabbi (Tryon, 1863), Parapholas californica
(Conrap, 1837) are all capable of burrowing into ex-
tremely hard rock and even into concrete. TURNER (1955)
states that C. ovoidea lives in soft shale, that Parapholas
californica is found in clay, shale and soft friable stone;
she does not say in what kind of rock Penitella gabbi is
found. Penitella gabbi is quite common in the soft sand-
stone rock of the Empire Formation near Fossil Point in
Coos Bay, Oregon, but is not found in harder rocks nearby
which are, however, bored by Penitella penita. Only P.
penita of all Pacific Coast pholads is capable of boring
extremely hard rocks.

The rock into which these burrows are drilled is reported
to be hard, siliceous and cherty. If, as ADEGOKE seems
to imply, the rock in the late Miocene period was as hard
as it is today, only Penitella penita or an extinct but
equally tenacious borer could have made these burrows.
However, there is considerable evidence to suggest that
at the time of the boring, the Monterey Formation rock
was a relatively soft shale with a high silica content
probably, as BRAMLETTE (1946) suggests, in the form of
diatom skeletons. BRAMLETTE (op. cit.) claims that the
present hard cherty character of the rock is due to the
solution and subsequent redeposition of diatomaceous silica.

ADEGOKE’s description of the dull glassy nature of the
burrows and their contents supports this suggestion.

The thinness of the valves is another indication of the
softness of the rock at the time of burrowing. In my
studies of living Penitella penita, animals with thin-walled
valves were always associated with soft rock.

If my claim is correct that the spherical burrows were
formed by Nettastomella rostrata, this is further evidence
that the rock in late Miocene was very soft. TURNER
(1955) states that N. rostrata is found in soft shale and
mud shale. In the Coos Bay region this species was found
in considerable numbers in very soft shale dredged from
deep water and more rarely collected intertidally in very
friable sandstone.

If the identity of the pholad which formed the conical
burrows could be established with confidence, the relative
hardness of the rock at the time it was being bored could
be estimated by analysis of burrow shape (EvANs, in prep-
aration, C) and this could be compared with its present
hardness. A method for estimating the relative hardness of
sedimentary rock is described in another paper (EvANs,
in preparation, C).

LITERATURE CITED

ADEGOKE, OLUWAFEYISOLA SYLVESTER
1966. Silicified sand-pipes belonging to Chaceia (?) (Phola-
didae: Martesiinae) from the Late Miocene of California.
The Veliger 9 (2): 233 - 235; plt. 21 (1 October 1966)
BRAMLETTE, Mitton NunN
1946. ‘The Monterey formation of California and the origin
of its siliceous rocks, U.S. Geol. Surv. Prof. Paper 212: 57
pp.; 19 plts.
Evans, JoHN WILLIAM
A in press. Factors modifying the morphology of the rock
boring clam, Penitclla penita. Proc. Malacol. Soc. London
[11 MS pp.; 4 plts.; 6 figs.]
B in preparation. Growth rate of the rock boring clam Penitella
penita (Conran, 1837). 14 MS pp.; 4 plts.; 7 figs.; 4 tables
C in preparation. The effect of rock hardness and other factors
on the shape of the burrow of the rock-boring clam, Penitella
penita. 10 MS pp.; 1 fig.; 2 tables
Turner, RutH Drxon
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

Page 176

THE VELIGER

Vol. 10; No. 2

The Reproductive System of the British Turridae

(GASTROPODA : TOXOGLOSSA)

EDMUND H. SMITH

Department of Zoology, University of Glasgow, and the Marine Station, Millport '

(Plate 18; 16 Text figures)

Tuat THE Neocastropops exhibit a similar organization
in their reproductive systems has been well illustrated in
the families Muricidaec, Buccinidae and Nassariidae (FRET-
TER, 1941; Jonansson, 1942, 1957; FRETTER & GRAHAM,
1962). Few species have been described from the families
Conidae, Terebridae, Olividac, Columbellidae and Turri-
dae (Rispec, 1955; Marcus, 1959, 1960, 1962; Rosin-
son, 1960) and the aim of the present paper is to provide
a more detailed account of the reproductive system of
the Turridae than has been published hitherto. The
species examined were Hacdropleura septangularis (Mon-
TAGU, 1803); Philbertia Icufroyi boothi (Smiru, 1839) ;
Cenodagreutes aethus E.H. Smirn, 1967; Cenodagreutes
coccyginus E.H.Smitu, 1967; Mangelia attenuata
(Montacu, 1803); Lora trevelliana (Turton, 1834) ;
Lora turricula (Montacu, 1803). This last species is of
particular interest since it exhibits a form of protandric
consecutive sexuality which has not been previously re-
corded in the Prosobranchia.

METHODS

The animals were removed from their shells, relaxed in
propylene phenoxytol (Owen, 1955) and fixed in Bouin’s
or Gilson’s fluid. The fixed material was sectioned and
stained with Alcian blue, hacmalum and eosin. Heiden-
hain’s iron haematoxylin and Mallory’s triple stains were
used for the study of the glandular parts of the repro-
ductive system.

MALE REPRODUCTIVE SYSTEM

In the typical male reproductive system (Figure 1) the
testis (t) shares part of the visceral mass with the digestive
diverticula (dd) which may even be partly displaced

' Present address: Pacific Marine Station, Dillon Beach, California
94929.

during the height of the reproductive period. The testicular
duct leaves the testis and progresses along the columella
muscle to enter the prostate gland (pg) at the posterior
end of the mantle cavity. This duct can be divided into
the vesicula seminalis which is the portion nearest the
testis and the vas deferens (vd) which is the narrow part
entering the prostate gland. The vesicula seminalis is dis-

Figure 1

Typical Male System

an — anus ct — ctenidium dd - digestive diverticula
f — foot fm — floor of mantle cavity i-— intestine
kd — kidney os — osphradium Pp — penis
pg — prostate gland pa — opening of penis t — testis

sh — siphon te — tentacles vd — vas deferens

Vol. 10; No. 2

THE VELIGER

Page 177

tended with sperm during the breeding season and some-
times functions in the ingestion of excess sperm. In some
species a gonopericardial duct or a connective tissue rem-
nant of the duct joins the vas deferens with the peri-
cardium. The convoluted prostate gland lies on the left
side of the mantle cavity (fm) and in some species in-
complete fusion of the two lobes of the gland leaves an
opening into the mantle cavity. A duct leads from the
prostate to the base of the penis (p) and then continues
along the length of the penis to open at its tip. The
penis itself lies along the left side of the mantle cavity
and in some species actually curves to the right along the
posterior limit of the cavity.

Haedropleura septangularis

The thin wall of the long, folded vesicula seminalis of
Haedropleura septangularis (Figure 2, vs) is composed of

pa P
¥ vs
aL a =
Figure 2

Haedropleura septangularis

Diagrammatic Reconstruction from Sections of the Male System
Ip —lumen of prostate gland pa — opening of penis
pd — duct from prostate t — testis

vd — vas deferens

Pp — penis
pg — prostate gland
vs —vesicula seminalis

a low epithelium surrounded by connective tissue; no
sperm ingestion was found. The vas deferens (vd), as it
leaves the vesicula seminalis (vs) is lined by glandular
cells which contain small, yellow particles concentrated
along their apical borders. No gonopericardial duct nor
any remnant of one could be found. After the vas deferens
leaves the area of the pericardium it narrows, becomes
heavily ciliated, and is surrounded by a thick muscular
layer. The long prostate gland (pg) which extends from
the area of the kidney, through the ventral floor of the
cephalic haemocoele, to the penis (p), is convoluted with
an indistinct midventral line of fusion between its two
lobes; no opening into the mantle cavity could be found.
The ciliated epithelium lining the lumen of the gland (Ip)
is underlain by a layer of circular muscles and also present
are subepithelial gland cells of two types: mucous cells
which form the dorsal portion of the gland and clusters
of large, darkly staining cells which form the ventral

part. These latter cells contain refractile inclusions
throughout their cytoplasm and open into the lumen by
long ducts which pass through the circular muscles and
between the ciliated cells.

The convoluted duct from the prostate gland (pg) to
the tip of the penis (pa) is ciliated and surrounded by a
thin, muscular layer. The large penis (p) reaches far
into the mantle cavity with its tip turning to the right
along the posterior limit. The spermiduct does not open
directly at the tip of the penis but into a shallow cavity
formed by the inpocketing of the two muscle layers. The
penis is covered by a layer of cuticular cells interspaced
by scattered mucous cells.

In view of the fact that most of the male turrids
studied exhibit a similar pattern in their reproductive
systems, the following species will be briefly compared
to Haedropleura septangularis.

Philbertia leufroyi boothi

In contrast to Haedropleura septangularis the vesicula
seminalis of Philbertia leufroyi boothi (Figure 3, vs) is
covered by a thick layer of circular muscle and lined with
a glandular epithelium which is engaged in ingesting
sperm. The sperm is attached to the apical pole of the

pa p

vs pd
vd ps

Figure 3
Philbertia leufroyi boothi
Diagrammatic Reconstruction from Sections of the Male System
p — penis
pd — duct from prostate
vd — vas deferens

mo-—opening from vas deferens to mantle
pa — opening of penis
pg — prostate gland t — testis

vs — vesicula seminalis

ingesting cells where it is engulfed and deposited in small
vacuoles. The epithelium of the vas deferens (vd) is com-
posed of short, ciliated, columnar cells resting on a thick
basement membrane and the muscular layer below this
membrane is thinner than that surrounding the vesicula
seminalis. An opening into the mantle cavity which some-
times occurs when there is incomplete fusion of the lobes
of the prostate gland is, in this species, at the end of a
short, ciliated duct (mo) from the vas deferens (vd)
where the vas deferens joins the prostate gland (pg). The

Page 178

histology of the prostate gland (pg) is similar to that of
H. septangularis although there are no mucous cells and
no line of fusion between the two lobes.

Cenodagreutes aethus

The vesicula seminalis (Figure 4, vs) of Cenodagreutes
aethus is not ciliated. The area near the testis (t) is
glandular, while the remaining portion of the vesicula
seminalis was distended with sperm, making it difficult
to ascertain the nature of the epithelium. The vas deferens

t pe hr pa
vs p
pd
vd gd mo PS
Figure 4

Cenodagreutcs aethus

Diagrammatic Reconstruction from Sections of the Male System

gd — gonopericardial duct hr — heart
mo-opening from vas deferens to mantle Pp — penis
pa — opening of penis pc — pericardial wall
pd — duct from prostate pg — prostate gland t — testis

vd —vas deferens vs — vesicula seminalis

(vd) is narrow and convoluted with an epithelium com-
posed of ciliated, cuboidal cells that rest on a thick layer
of circular muscle. Contrary to the preceding turrids there
is a remnant of the gonopericardial duct (gd) in the form
of a short, thick strand of connective tissue which joins
the duct to the pericardial wall (pc). As in Philbertia
leufroyi boothi a ciliated muscle duct (mo) opens into
the mantle cavity from the vas deferens (vd). Whcreas
the prostate gland (pg) is similar to those in the previously
described turrids, the densely ciliated duct (pd) to the tip
of the penis (p) differs in being muscular; also the penis
is round, and not dorsoventrally flattened.

Cenodagreutes coccyginus

The few specimens collected of this rare turrid were all
females.

Mangelia attenuata

As was the case with Cenodagreutes coccyginus, all the
specimens collected were females.

THE VELIGER

Vol. 10; No. 2

Lora trevelliana

The vesicula seminalis (Figure 5, vs) is glandular and
composed of cells with yellow granules of different sizes
and shapes scattered throughout the cytoplasm. The densely

pe hr pa Pp

vd gd Ps

Figure 5
Lora trevelliana

Diagrammatic Reconstruction from Sections of the Male System

gd — gonopericardial duct hr — heart p-— penis
pa — opening of penis pc — pericardial wall
pd — duct from prostate pg — prostate gland t — testis

vd —vas deferens vs — vesicula seminalis

ciliated vas deferens (vd) is narrow and gives off a long
gonopericardial duct (gd) which opens into the peri-
cardium (pc) through a ciliated funnel. The prostate
gland (pg) is composed of the usual three cell types
although the gland cells are not subepithelial. As in Haed-
ropleura septangularis there is no opening from the pros-
tate to the mantle cavity. The duct (pd) from the prostate
(pg) to the penis (p) is glandular and composed of large,
vacuolated cells interspaced by small, ciliated cells. This
type of epithelium continues along the large penis almost
to the tip where it becomes heavily ciliated and not
glandular. A sphincter surrounds the opening of the duct
at the tip of the penis.

FEMALE REPRODUCTIVE SYSTEM

In the typical female reproductive system (Figure 6)
the ovary (0) shares part of the visceral mass with the
digestive diverticula (dd). A gonadial oviduct (go) leaves
the ovary and runs along the inner surface of the viscera
next to the columella muscle to join the renal portion of
the oviduct (ro) which passes close to the pericardium
and in some species gives off a duct to the pericardial
cavity. This region of the oviduct enters a large, convoluted
albumen gland (al) which opens in turn into the capsule
gland (cg) through a short pallial oviduct. In some species

Vol. 10; No. 2

Figure 6

Typical Female System
an — anus
ct — ctenidium

al—albumen gland
cg —capsule gland

be — bursa copulatrix
dd — digestive diverticula

f — foot fm — floor of mantle cavity hr — heart
i—intestine ig—ingesting gland kd — kidney O — ovary
op-—operculum os—osphradium _ sh ~ siphon te — tentacles

a receptaculum seminis and ingesting gland (ig) lie be-
tween the albumen gland and capsule gland. A duct
connects the receptaculum with the pallial oviduct. The
capsule gland is the largest organ of the female system
and lies along the Icft side of the mantle cavity. This
gland opens into the bursa copulatrix (bc) which in turn
opens into the mantle cavity slightly anterior to the anal
opening (an).

Haedropleura septangularis

The gonadial oviduct (Figure 7, go) is composed of
columnar gland cells with a lightly staining basal ergasto-
plasm and is surrounded by a thick muscular layer. No
evidence of egg yolk absorption was found (FRETTER,
1941).

The renal oviduct (ro) is short, narrow, and enters the
the albumen gland (al) ventrally. There is no gonoperi-
cardial duct.

The large albumen gland has a ciliated lumen (la) that
is partially divided medially by a thin septum (sp) which
extends halfway to the floor from its dorsal wall. The
entire gland is surrounded by a thick muscular layer. The

THE VELIGER

Page 179

anterior end of the gland opens into the capsule gland

(cg) through a very short, ciliated pallial oviduct (po).
The large ingesting gland (ig) lies between the albumen

(al) and capsule glands (cg) and extends for a short

go sp 8 cg
fo) sr be
ro al la P° be gt
Figure 7

Haedropleura septangularis
Diagrammatic Reconstruction from Sections of the Female System
al — albumen gland be — bursa copulatrix cg —capsule gland
go—gonadial oviduct gt-genital opening _ ig — ingesting gland
la—lumen of albumen gland 0 - ovary po — pallial oviduct
ro—renal oviduct sp — septum sr —receptaculum seminis

distance over the dorsal surface of the latter. The gland
is not attached to the pallial oviduct (po) but to the
anterior end of the albumen gland (al) by a long, narrow,
ciliated duct which acts as a receptaculum seminis (sr).
This fact would indicate that the most likely place for
fertilization would be in the albumen gland (al) and not
in the pallial oviduct (po). The lumen of the ingesting
gland is filled with oriented sperm cells arranged in con-
voluted rows and not attached to the walls. Some sperm
ingestion does occur, but the epithelium is so thin that
the amount of sperm ingested, when compared to the vast
amount of sperm present in the lumen, must be small.

The capsule gland (cg), although somewhat more com-
plex in its staining properties than those of the muricids
and buccinids, is essentially similar in its morphology to
that described by FretTeR & GraHaAM (1962) for Thais
lapillus.

An elongated bursa copulatrix (bc) runs dorsad along
the anterior end of the capsule gland (cg) and curves
slightly posteriad at the distal end of the capsule gland.
The bursa is lined anteriorly by a mucoid epithelium inter-
spaced by small, ciliated cells. Further to the posterior the
epithelium changes to a low, ciliated, cuboidal layer. Ori-
ented sperm were attached to this ciliated epithelium, while
unoriented sperm, mixed with mucus, filled the distal end of
the bursa. The entire bursa copulatrix is covered by a thick
layer of muscle.

Philbertia leufroyi boothi

The gonadial oviduct (Figure 8, go) from the ovary to
the albumen gland (al) is short and surrounded by a thin

Page 180

connective tissue layer; the lumen is lined by tall columnar
cells which stain lightly with haemalum. The renal oviduct
(ro) is also extremely short and enters the large, slightly
elongated albumen gland (al) through a narrow opening
that does not possess a sphincte:.

ig cg cg as
go sr
oO
be gt
To al i Ic
Figure 8

Philbertia leufroyi booth

Diagrammatic Reconstruction from Sections of the Female System
al — albumen gland as — anterior sperm sac
be — bursa copulatrix go — gonadial oviduct

gt— genital gland ig — ingesting gland
la—lumen of albumen gland Ic—lumen of capsule gland
0 — ovary ro — renal oviduct sr — receptaculum seminis

cg — capsule gland

The lumen of the albumen gland (al) is narrow, ciliated
and lacks the septum present in Haedropleura septangu-
laris. The dorsal half of the albumen gland is composed of
clusters of mucous cells which open into the lumen through
long protoplasmic processes. Lining the lumen (ls) and
interspaced between the ciliated cells are other gland cells
which stain deeply in haemalum. The ventral part of the
albumen gland is of the same cellular organization except
that the subepithelial gland clusters are not mucoid, but
stain with haemalum and eosin.

As in Haedropleura septangularis a short, straight duct
connects the ingesting gland (ig) with the anterodorsal
wall of the albumen gland (al). The ciliated lumen of
the duct is partially divided near its junction with the
albumen gland by two opposing lateral folds. A sharp line
of demarcation divides the epithelium of the duct from
that of the ingesting gland. Although no oriented sperm
were found in the duct (sr), the possibility that it acts
as a receptaculum seminis cannot be ruled out.

The ingesting gland is composed of very large (200p to

THE VELIGER

Vol. 10; No. 2

270 long and 50u to 120m wide) gland cells (Plate 18,
Figure 1). These cells contain ingested sperm cells (is)
which are in the process of disintegration. The cytoplasm
surrounding the vacuoles containing the disintegrating
sperm is filled with yellow granules which accumulate
at the apical pole of the cell. A deeply staining, fibrous
ergastoplasm (ep) forms a dark basal border while large,
misshapen nuclei (pn) up to 75y long, lie basally near
the corners of the cells. The pronounced nuclear poly-
morphism may be due to endomitosis with endopolyploidy
which is common in some invertebrates (GABE & ARvVy,
1961). Partly disintegrated sperm (s) were found in the
lumen of the gland.

The staining properties and ciliary currents of the
capsule gland are not as complex as those of Haedropleura
septangularis or those of the muricids and buccinids
(FRETTER, 1941; personal observation).

In the albumen gland (al) lateral ciliary currents
(Figure 9) beat into a strong central current which flows

Figure 9

Philbertia leufroyi booth

Diagram of the Ciliary Currents in the Capsule Gland
al — albumen gland cg — capsule gland ro — renal oviduct
sO — opening to receptaculum seminis sr — receptaculum seminis

directly into the capsule gland (cg). Here anteriorly
directed currents beat along the dorsal lips of the lobes
and along the ventral suture while anteroventral currents
beat over the middle of each lobe towards the main vent-
ral current. The complex currents, which beat in opposite
directions, found in the muricids and buccinids, are absent.

Since the ventral channel of the capsule gland is absent,
the small bursa copulatrix (bc) opens directly into the

Explanation of Plate 18

Figure 1

Philbertia leufroyi boothi

Cross Section of the Ingesting Gland
cg—capsule gland ep—basal ergastoplasm —_is— ingested sperm
kd — kidney Ic —lumen of capsule gland
pn — polymorphic nuclei

s— sperm sr —receptaculum seminis

Figure 2
Philbertia leufroyi boothi

Cross Section of the Anterior Sperm Sac
cg —capsule gland es — epithelium of the anterior sperm sac
i — intestine s— sperm

THE VELIGER, Vol. 10, No. 2 [SmiTH] Plate 18

fie m

2

300p.

Figure 2

photographs by E. H. Smita

Vol. 10; No. 2

THE VELIGER

Page 181

lumen of the capsule gland (Ic). The tall epithelium lining
the bursa contains ciliated cells with granular inclusions
which are actively secreting into the lumen.

In contrast to Haedropleura septangularis a short, non-
ciliated duct enters the dorsal surface of the bursa (bc)
from a large, round “anterior sperm sac” (as) which lies
near the anterior end of the capsule gland (cg). This
sac is filled with oriented sperm (Plate 18, Figure 2)
with their heads pointing towards the walls (es) of the
sac but not in actual contact with the epithelium which
consists of long, thin cells filled with granules and small
vacuoles. No evidence of sperm ingestion could be found.
A somewhat similar “sac,” termed a “terminal pouch,”
was described by Marcus (1960) in Hastula cinerea,
but, whereas in this species it is situated anteriad to the
bursa copulatrix, in Philbertia leufroyi booth: it lies dor-
sally between the bursa and the capsule gland.

Cenodagreutes aethus

The gonadial and renal portions (Figure 10, go, ro)
of the oviduct are similar to those of Haedropleura sept-
angularis. The albumen gland (al) is ciliated throughout,
but contrary to the preceding turrids, there are no subepi-
thelial glands. Instead, large mucous cells interspaced with

3 cg
1g
o i al | | be
To la id gt
Figure 10

Cenodagreutes acthus

Diagrammatic Reconstruction from Sections of the Female System
be — bursa copulatrix cg — capsule gland
gt—genital opening _ig — ingesting gland
la—lumen of albumen gland
ro — renal oviduct

al — albumen gland

go — gonadial oviduct

id—duct to ingesting gland
0 — ovary

ciliated cells form the dorsal wall of the gland while the
much thinner ventral wall has only scattered mucous cells.
The epithelium of the albumen gland (al) becomes con-
tinuous with the posterior part of the capsule gland (cg) ;
a slight constriction rather than a duct marks the boundary
between the two glands. Therefore, the pallial oviduct is
absent.

A short, constricted, muscular duct joins the antero-
dorsal part of the albumen gland with the ingesting
gland (ig) and functions as a receptaculum seminis (sr).
The lumen of the ingesting gland is lined with gland cells
possessing large, polymorphic nuclei that exhibit chroma-

ticity when stained with haemalum. The cells rest on a
thin basement membrane which is in turn surrounded by
a layer of connective tissue.

The capsule gland (cg) is similar to that of Haedro-
pleura septangularis, although its staining properties are
more complex. The bursa copulatrix (bc) is muscular with
a narrow lumen which opens dircctly into the capsule
gland as in Philbertia leuroyi boothi. The epithelium lining
the capsule gland and extending a short way into the
bursa is composed of gland cells which stain deeply in
haemalum. This glandular epithelium rapidly gives way
to a simple cuboidal layer of cells surrounded by thick
(30 to 60), circular muscles.

Cenodagreutes coccyginus

The female system of Cenodagreutes coccyginus is very
similar to that of C. aethus, with some exceptions (Figure
11). There is a pallial oviduct (po) between the albumen
gland and the capsule gland (cg).

i b po cg

be

ro al sr be

gt
Figure 11

Cenodagreutes coccyginus

Diagrammatic Reconstruction from Sections of the Female System
al—albumen gland b — bulb be — bursa copulatrix
cg — capsule gland go — gonadial oviduct gt — genital opening

0 — ovary po — pallial oviduct ro—renal oviduct
sr —receptaculum seminis

No sperm were found in the small distal bulb of the
receptaculum seminis (sr), nor was there any evidence
of sperm ingestion. However, the possibility of sperm
ingestion cannot be completely ruled out. The low epi-
thelium lining the terminal bulb of the receptaculum semi-
nis is composed of eosinophilic cells with irregularly shaped
granules throughout the cytoplasm. The general histolog-
ical and morphological data indicate that both bulb and
duct function as a receptaculum seminis.

The bursa copulatrix (bc) is much larger than that of
Cenodagreutes aethus with the bulb forming the distal
end of the bursa and lying at the anterior end of the
capsule gland (cg). The bulb, although it has become
part of the bursa and functions in a different manner,
lies in the same position as the “anterior sperm sac” of
Philbertia leufroyi boothi. It is not lined by glandular
cpithelium, but by one composed of densely ciliated, cub-
oidal cells which rest on a thin layer of circular muscle.

Page 182

Mangelia attenuata

The gonadial oviduct (Figure 12, go) is comprised of
tall, glandular cells which are, for the most part, chromato-
phobic and in which there is a concentration of granules
near the apical pole that stains with Alcian blue. In

Figure 12
Mangelia attenuata

Diagrammatic Reconstruction from Sections of the Female System
al - albumen gland b—bulb bd — duct to bulb
cg — capsule gland go — gonadial oviduct gt — genital opening
la—lumen of albumen gland Ic—lumen of capsule gland
0 - ovary po — pallial oviduct ro-—renal oviduct

sp — septum ve — ventral channel

contrast to the previously described turrids, a strong
sphincter divides the renal oviduct (ro) from the albumen
gland (al). This is ciliated throughout and as in both
species of Cenodagreutes there are no subepithelial glands.
The entire gland and the long pallial oviduct (po) con-
necting it with the capsule gland (cg) are covered by
a thick sheet of muscle. The duct is lined by columnar
cells bearing dense cilia.

Branching from the pallial oviduct is a long, muscular
duct (bd) with a densely ciliated lumen similar to that
of the pallial oviduct. This straight duct leads to a very
large, elongated bulb (b) which lies over the dorsal
surface of the albumen gland and is not branched as in
Mangelia brachystoma (Rosinson, 1960). The cells lining
the bulb exhibit different stages of intracellular elabora-
tion, culminating in the extrusion of a voluminous globule
and they resemble the ingesting type of cell found in
Philbertia leufroyi boothi in having large, polymorphic
nuclei, although neither the duct nor the bulb contained
sperm in any of the specimens sectioned and the nature
of the epithelium does not suggest an ingesting function.

The capsule gland shows the same staining complexity
found in both species of Cenodagreutes. As in Mangelia
brachystoma (Roxpinson, 1960) no separate bursa copu-
latrix (bc) is present, but a large, ventral channel (vc),
which is separated from the capsule gland by a septum
(sp), extends posteriorly past the gonopore (gt) for some
distance. The penis is inserted into this channel instead of
a separate chamber as in the preceding turrids. The
septum, which separates the anterior part of the ventral

THE VELIGER

Vol. 10; No. 2

channel from the capsule gland, disappears posteriorly so
that the channel is marked only by a slight constriction.

Lora trevelliana

Whereas in most of the turrids studied the gonadial
oviduct (Figure 13, go) is long, in this species it is short
and ciliated and, in addition, the cells have a vacuolated
cytoplasm with small granules concentrated around the
cell walls. The long albumen gland is similar to that of
both species of Cenodagreutes with some slight staining
differences; the gland cells are not subepithelial. The
elongated ingesting gland (ig) extends posteriorly over the
dorsal surface of the albumen gland (al) and the cells
lining the duct to the ingesting gland are densely ciliated,

2 8° ig po cg

Tro al sr be

Figure 13
Lora trevelliana
Diagrammatic Reconstruction from Sections of the Female System
al — albumen gland be — bursa copulatrix cg — capsule gland
go—gonadial oviduct gt—genital opening __ ig — ingesting gland
0 — ovary po — pallial oviduct ro— renal oviduct
sr — receptaculum seminis

while the epithelium lining the gland itself is composed of
large gland cells actively engaged in sperm ingestion. Ori-
ented sperm cells were found attached to the walls of the
duct to the ingesting gland, showing that the duct functions
as a receptaculum seminis (sr).

The capsule gland resembles that of Philbertia leufroyt
boothi in the complexity of its staining reactions. The
bursa copulatrix (bc) is an incapacious, elongated struc-
ture with an epithclium comprising scattered mucous cells,
ciliated cells and large gland cells.

HERMAPHRODITIC SYSTEM

Lora turricula

In a number of specimens of this species both male and
female systems were present in the same individual. It
was thought best to deal with both the male and the
female systems at the same time in order to facilitate
discussion of the hermaphroditic condition.

Vol. 10; No. 2

The male system is quite normal and follows a pattern
similar to that described for Lora trevelliana (Figure 14).
The testis (t) shows no evidence of sexual transformation
such as the presence of oocytes around the periphery, as
found in Crepidula fornicata (Cor, 1942). The vesicula

pe hr pa p

pd

Vs vd &P pd

Figure 14
Lora turricula
Diagrammatic Reconstruction from Sections of the Male System
gp —connective tissue remnant of gonopericardial duct
hr — heart Pp — penis pa — opening of penis
pc — pericardial wall pd - duct from prostate t — testis
vd — vas deferens vs — vesicula seminalis

seminalis (vs) is well developed and as in Philbertia leu-
froyi boothi, it is not ciliated but composed of gland cells,
which are engaged in sperm ingestion near the testis.
These cells rest on a basement membrane which is sur-
rounded by a thin layer of connective tissue. The vas
deferens (vd) is convoluted and made up of densely
ciliated, cuboidal cells resting upon a thin basement mem-
brane covered by a thin layer of muscle. There is a
remnant of the gonopericardial duct (gp) attaching the
vas deferens (vd) to the wall of the pericardium (pc).
There is no prostate gland nor opening into the mantle
cavity. The ciliated epithelium of the vas deferens changes
abruptly, near the posterior end of the kidney, to a mucoid
epithelium containing small, ciliated cells wedged between
large mucous cells. This type of glandular spermiduct (pd)
continues to the tip of the penis as in Lora trevelliana.

sr pa p

fo) To b

Figure 15

Diagrammatic Reconstruction from Sections of the Female System
with the nearly complete Male System

al- albumen gland b—bulb be - bursa copulatrix
cg — capsule gland go — gonadial oviduct gt — genital opening
0 - ovary Pp — penis pa — opening of penis

pd —duct from prostate po-pallial oviduct —_ ro — renal oviduct

sr — receptaculum seminis

THE VELIGER

Page 183

A number of predominantly female specimens had large,
round penes (Figure 15). In these specimens there was
no trace of testis and only a small portion of the ovary (0)
contained oocytes. The remainder of the last two visceral
whorls was filled with mature, normal eggs. No evidence
of a renal oviduct nor of a gonopericardial duct could be
found. The thin gonadial oviduct (go) enters the albumen
gland (al) directly, with no sphincter between it and
the gland. It is composed of a non-glandular epithelium
surrounded by a thin muscular layer.

The albumen gland (al) is ciliated and possesses sub-
epithelial cells which are of two kinds. Those of the dorsal
wall color deeply with Alcian blue, while those of the
ventral wall stain with haemalum. The albumen gland
opens into the capsule gland (cg) through a short, ciliated
pallial oviduct (po); a thin layer of muscle surrounds
the duct.

The ciliated ventral channel leads posteriorly from the
bursa copulatrix (bc) along the ventral part of the capsule
gland (cg) to join the pallial oviduct (po).

A short, ciliated duct connects the elongated recepta-
culum seminis (sr) to the pallial oviduct (po). The bulb
is lined with a very thin (1.3), non-glandular epithelium.
No evidence of sperm ingestion could be found, and indeed
the thin epithelium does not seem to be of the type capable
of ingesting sperm. The duct and receptaculum seminis
are surrounded by a thin (4) muscle layer.

The well developed capsule gland (cg) is similar to that
of Lora trevelliana, except that the three histological areas
present in other species react differently to the same stain.
The bursa copulatrix (bc) is very muscular with a large,
ciliated lumen; gland cells are absent.

The male system, which is present in these same female
specimens, consists of a normal penis (p) and glandular
spermiduct (pd), which terminates in a sperm filled bulb
near the albumen gland (al) of the female. Unoriented
sperm was found along the spermiduct almost as far as
the penis tip (pa).

e b pd sr pa
Can Sig
pd
go
ro al po cg be gt

Figure 16

Diagrammatic Reconstruction from Sections of the Female System
with Vestiges of the Male System

al—albumen gland b—bulb be — bursa copulatrix
cg — capsule gland go — gonadial oviduct gt —genital opening
0 — ovary p — penis pa — opening of penis

pd-duct from prostate po-—pallial oviduct ro — renal oviduct

sr —receptaculum seminis

Page 184

THE VELIGER

Vol. 10; No. 2

In one of the female specimens (Figure 16) the male
system had atrophied so that only the terminal bulb (b),
near the albumen gland (al) of the female, and a short
portion of the glandular spermiduct remained. The penis
was very reduced with no sperm present in any part of
the male system.

PEDAL GLANDS

Within the neogastropods the pedal glands are not
entirely engaged in lubricating the foot during locomotion.
They may undertake such diverse functions as moulding
and attaching egg capsules or aiding in the boring of
holes in the shell or hard exoskeleton of the prey.

The anterior pedal mucous glands open into a ciliated,
sagittal canal in the turrids studied. In many cases these
subepithelial glands do not react with Alcian blue nor
give any indication of the presence of an acid mucopoly-
saccharide. In Philbertia leufroyi and Mangelia attenuata
scattered mucous clusters occur mixed with darker, blue
staining cells on both sides of the sagittal canal. In Ceno-
dagreutes aethus and C. coccyginus the anterior pedal
glands give three staining reactions. Some gland clusters
stain with haemalum, some with eosin and haemalum, and
a few others with Alcian blue.

“Sole glands” is a collective term applied by FRETTER
& GRAHAM (1962) to include all the glands which pour
secretions onto the surface of the sole, including goblet
cells, which lie within the epithelium and subepithelial
glands. In Philbertia leufroyi booth a large mass of sub-
epithelial glands fills much of the mesopodium. The gland
cell clusters stain intensely with Alcian blue and open
between the ciliated epithelial cells by long ducts onto
the surface of the foot. In Mangelia attenuata the epi-
thelium covering the sole is composed of eosinophilic cells
interspaced by ciliated cells, mucous cells and subepithelial
glands being absent. The posterior pedal mucous gland,
which occurs in some rissoids and triphorids, is absent.
The ventral pedal gland present in female muricids and
buccinids and concerned with the final moulding and
attachment of the egg capsules, is totally absent.

DISCUSSION

The neogastropod reproductive system has been con-
sidered uniformly dioecious and it is, therefore, surprising
to find that Lora turricula exhibits a form of hermaphro-
ditism. In the mesogastropod genera, which exhibit true
protandric consecutive sexuality, such as Calyptraca, Cre-
pidula and Crucibulum, each individual functions as one
sex when young (male) and as the opposite sex when

older. The histological basis of this type of sexuality
depends upon the presence of an ambisexual gonad (Cos,
1944). In addition, the male genital system is recon-
structed into an oviduct, seminal receptaculum and vagina
during the transition from the male phase to the female
phase. This is not the case with L. turricula, since there
is no ambisexual gonad, and the male and female repro-
ductive systems are found together in the same individual.
In L. turricula the gonads undergo a similar cycle to that
described by Yonce (1962) for Trichotropis cancellata,
where the gonads are undifferentiated during the transi-
tion from the male to the female phase. However, L.
turricula differs from the above gastropods and from T.:
borealis (GRAHAM, 1956) in having two separate and
complete reproductive systems present in the same indi-
vidual, although the male system is no longer connected
to the gonads and gradually atrophies during the change
to the female phase, eventually leaving a separate, func-
tional female system. This type of hermaphroditism is
similar to that found in the pulmonates and opisthobranchs
except that both sexual systems are not complete at the
same time. It is conceivable that some specimens of L.
turricula could act as functional hermaphrodites during
the brief transitional phase when there is still sperm in
the male duct and a well developed penis still present.
The possibility of self-fertilization, as suggested by YONGE
(1962) for T: cancellata, where the penis remains through-
out the life of the female, seems remote in L. turricula.

In Patella vulgata, which exhibits alternative sexuality,
Orton, SouTHwaARD & Dopp (1956) found, in a single
population, that 90% of the small specimens, 16 - 25mm
in shell length, were males. Those with shells 40mm long
were male and female in equal numbers, while 60 - 70%
of the larger animals, with shells 60mm in length or
longer, were female. Earlier Orton (1919, 1928) sug-
gested that the young population contained two kinds of
male-phase individuals, true males and temporary males
(protandric females). Due to the difficulty of locating
Lora turricula, only a small number could be collected.
This sample consisted of 15 individuals with an average
shell length of 14.8mm (Table 1). The sample can be
broken into 3 males, 5 females, and 7 in various stages of
the transitional phase. Contrary to what Orton found,
this small sample can not be broken into distinct sex phases
based on shell size. The results from this sample seem to
indicate that the population may consist of two sexual
phases: a pure male phase and a temporary male phase
(protandric hermaphrodite), later becoming female.

Populations of Lora turricula are probably restricted to
small geographic areas by a number of complex factors,
including specialized feeding habits and particle size of
the substrate. Therefore, this form of hermaphroditism
would be of direct importance to such insular populations

Vol. 10; No. 2

THE VELIGER

Page 185

Table 1

Occurrence and Condition of the Reproductive Systems
in 15 Specimens of Lora turricula Collected from the
Clyde Sea Area, Scotland, During June, 1964.

Z
:
g

OOANODULON &
—
oO
o

cs
o
1, 40 40 1 1 10 GF QQ, 40 10 1] QA

Condition of reproductive system

well developed male and female systems

typical male, no trace of female system

very reduced penis present

well developed male and female. systems, no testis present
well developed female, no male system present

well developed female, no male system present

typical male, no trace of female system

very reduced penis present

typical male, no trace of female system

well developed female, no male system present

well developed male and female systems, no testis present
very reduced penis present

well developed female, no male system present

well developed female, no male system present

well developed male and female systems, no testis present

by assuring the presence of both males and females. Two
causal factors have been indicated in the evolution of the
hermaphroditic state, genetic change and environmental
conditions (FRETTER & GRAHAM, 1964). Obviously, a
genetic change has taken place in the isolated population
studied and evidently hermaphroditism has not been se-
lected against by the environmental conditions. Of course,
a number of widely separated populations will have to
be sampled in order to show that all populations of L.
turricula exhibit hermaphroditism.

In some Stenoglossa FRETTER (1941) found that sperm
ingestion, and in some species egg yolk ingestion, takes
place in an ingesting gland located between the albumen
and the capsule glands. The convoluted duct, which runs
from the ingesting gland to the pallial oviduct, acts as a
receptaculum seminis with oriented sperm attached along
its walls. JoHANNSON (1957), from his work with Nassa-
rius pygmaea and N. incrassatus, doubted that sperm in-
gestion occurred in all Stenoglossa which possess a gland in
this same position. He also stated that it was not possible to
rule out sperm ingestion in these species because of the
simplicity of the epithelium lining the gland or because,
at the time of sectioning, the gland cells did not contain
ingested sperm. Sperm ingestion was not found in the
turrid species Mangelia attenuata and Lora trevelliana.
The histological nature of the epithelium and the condi-
tion of the sperm within the gland indicate that it is used
for sperm storage as is the duct leading to the gland. This

case agrees with JOHANNSON’s observations with both the
gland and duct functioning as a receptaculum seminis.

In the turrids the ingesting gland and the receptaculum
seminis are located in a proximal position which at first
suggests a homology with the receptaculum of other gastro-
pods, as postulated by FRETTER (1941) for the Stenoglossa.
However, JoHANNSON (1957) has suggested that the
ingesting gland corresponds to a proximal or inner bursa
copulatrix and not a receptaculum. Although the turrids
studied do not offer exception to FreTTeER’s theory, it
would be misleading not to mention Drupa nodulosa
(E. H. Smiru, in preparation) which supports JoHANN-
son’s theory since there is in this species both an ingesting
gland and a separate receptaculum seminis. In addition,
Marcus (1962) has found that in some species of Colum-
bellidae the pericardial cavity acts as a receptaculum
seminis with no sperm ingestion, while in other species
the long duct from the pallial oviduct may act as a
receptaculum seminis or in some cases as an ingesting
gland. From this information on the Columbellidae, it
is apparent that the position of the ingesting gland varies
in different families of the neogastropods and it is im-
possible to postulate a general homology for the gland that
applies to the whole group.

In some Stenoglossa and Toxoglossa an “anterior sperm
sac” is found. This bulb lies dorsally above the bursa copu-
latrix and opens into it through a duct or muscular
opening. It occurs in the turrid Philbertia leufroyi boothi,

Page 186

THE VELIGER

Vol. 10; No. 2

the terebrid Hastula cinerea (Marcus, 1960), and in
some Columbellidae (Marcus, 1962) and Muricidae
(personal observation). This bulb usually contains vast
amounts of oriented sperm cells which are attached to the
wall or lie in rows that are unattached. In the case of the
Columbellidae it takes on the function of sperm ingesting
and opens into the anteroventral portion of the bursa copu-
latrix. However, in P leufroy: boothi there is no sperm
ingestion and the “anterior sperm sac”’ acts as a receptac-
ulum seminis in the absence of the usual one between
the albumen and capsule glands.

In some species of Stenoglossa and Toxoglossa there is
a duct leading from the prostate gland to the mantle
cavity instead of the slit caused by the incomplete fusion
of the two lobes of the prostate gland found in some
muricids. In two turrid species, Mangelia brachystoma
(Ropinson, 1960) and Cenodagreutes aethus, a duct
leads to the mantle cavity from the pallial portion of the
vas deferens instead of from the prostate gland. FRETTER
(1941) suggested that these well developed ducts which
connect the spermiduct to the mantle cavity, may function
in the removal of excess sperm though no definite evidence
was presented. There is little doubt that the ducts in these
two turrids are functional rather than being mere rem-
nants of evolution, although there is no indication what
the function could be.

The pedal glands, commonly found in the muricids and
buccinids, are absent in the turrids, and yet the egg
capsules, which are formed, are similar to those of the
muricid Trophon truncatus. The laying of one egg capsule
of Cenodagreutes aethus was observed, but owing to the
small size of the snail, it was difficult to follow the entire
event. The capsule seemed to be moved to the sole of the
foot in the usual manner exhibited by the muricids and
buccinids. With the absence of the ventral pedal gland
the capsule gland may take over its function. Perhaps the
more complex staining properties of the capsule gland in
many of the turrids is an indication of the presence of
added chemical substances that take the place of the
secretion from the ventral pedal gland.

It is difficult to establish any phylogenetic relationships
between the Turridae and other neogastropod families, or
between the turrid species themselves. ScHILDER (1947)
has placed the origin of the Toxoglossa in the upper Cre-
taceous or Paleocene, while PoweEtt (1942) believes that
the origin of the Turridae took place at least in the carly
part of the Cretaceous, if not much earlier. The presence
of free-swimming larvae (LEBour, 1937; THorson, 1946)
indicates a primitive condition of the Turridae, as com-
pared to the Stenoglossa, where the tendency is to suppress
free-living larvae. On the other hand, the complete fusion
of the prostate gland in the male, the general lack of

gonopericardial connectives, and the occurrence of the
hermaphroditic state, suggest a more advanced position.
The paleontological record seems to indicate that the
Turridae arose early from a common neogastropod stem.
This early origin would allow time to develop specialized
trends while retaining some of the more primitive features.
The morphology of the turrid reproductive system is
similar in many respects to that found in the Buccinidae
and Muricidae, and does suggest parallel evolution of the
Stenoglossa.

SUMMARY

The functional morphology of the reproductive system in
seven species of British turrids is described. One species,
Lora turricula, is a protandrous hermaphrodite. The form
of hermaphroditism described differs from that found
in the mesogastropods in not having an ambisexual gonad
and by possessing two separate reproductive systems,
neither derived from the other, in the same individual;
this latter characteristic is also present in the functional
hermaphroditism of pulmonates and opisthobranchs. The
possible cause and benefit of this type of reproduction is
discussed. The function of the ingesting gland and recep-
taculum seminis is discussed and it is suggested that the
homology betwecn the ingesting gland of the Stenoglossa
and the reccptaculum seminis of other neogastropods can
only be made in certain cases and can not be applied to
the group as a whole. The pedal glands and the formation
of the cgg capsules in the turrids is described. It is con-
cluded that, although no definite phylogenetic conclusions
can be reached, the morphology of the turrid reproductive
system suggests a parallel evolution to the Stenoglossa.

ACKNOWLEDGMENTS

I would like to thank Professor C.M. Yonge, C.B.E.,
ER.S., for his help and encouragement during the course
of this investigation. I would especially like to thank
Professor G. Owen and Dr. J. Bowden for their help. I
would also like to thank Dr. C. H. Mortimer, ER.S.,
Director of the Marine Station, Millport, and his staff
for the research facilities made available and their help
during my stay. Dr. J. Pearce and Dr. K. Ockelmann sent
specimens of Lora trevelliana from Denmark, which were
of great help to the study. This investigation was supported
(in part) by a U.S. Public Health Service fellowship
(1-FI-GM-21, 563-01) from the National Institute of
General Medical Sciences.

Vol. 10; No. 2

LITERATURE CITED

Coz, WEsLEY R.

1942. The reproductive organs of the prosobranch mollusk
Crepidula onyx and their transformation during the change
from male to female phase. Journ. Morph. 70: 501 - 512

1944, Sexual differentiation in mollusks. II. Gastropods, amphi-
neurans, scaphopods, and cephalopods. Quart. Rev. Biol.
19: 85-97
FRETTER, VERA
1941. The genital ducts of some British stenoglossan proso-
branchs. Journ. mar. biol. Assoc. U. K. 25: 173 - 211
FRETTER, VERA, & ALASTAIR GRAHAM
1962. British prosobranch molluscs, their functional anatomy
and ecology. London, Ray Soc. xvi+755 pp.; 317 figs.

1964. Reproduction. In: Physiology of Mollusca, ed. K. M.
Wizzur & C. M. Yonce, Vol. 1. Acad. Press, New York, pp. 127
to 164

GaseE, M. « L. Arvy
1961. Gland Cells.

GraHaM, ALASTAIR

Chapt. 1 in “The Cell, 3” = Acad. Press

1954. The anatomy of the prosobranch Trichotropis borealis
BropveErip & SOWERBY, and the systematic position of the Capuli-
dae. Journ. mar. biol. Assoc. U. K. 33: 129 - 144

Jouansson, J.

1942. Von diaulen Geschlechtsapparaten bei den Prosobranchi-

ern. Ark. Zool. 34: 1 - 10

1957. Notes on the littorinacean and stenoglossan genital organs,
and a comparison with the Rissoacea. Zool. Bidr. Uppsala
32: 81-91

Lesour, Marie V.

1937. The eggs and larvae of the British prosobranchs with
special reference to those living in the plankton. Journ.
mar. biol. Assoc. U.K. 22: 105 - 166

Marcus, Ernst & EvELINE Marcus

1959. Studies on Olividae. Bol. Fac. Filos. Cienc. 8. Paulo,
Zool. 22: 99 - 188

1960. On Hastula cinerea.
Zool. 23: 25 - 66

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

THE VELIGER

Page 187

Marcus, Ernst & EvELINE Marcus
1962. Studies on Columbellidae.
S. Paulo, Zool. 24: 335 - 384
Orton, Joun H.
1919. | Sex phenomena in the common limpet (Patella vulgata).
Nature 104: 373 - 374

1928. Observations on Patella vulgata. Part I. Sex-phenomena,
breeding and shell-growth. Journ. mar. biol. Assoc. U. K.
15; 851 - 862
Orton, Joun H., A. J. SourHwarp & J. M. Dopp

1956. Studies on the biology of limpets II. The breeding of
Patella vulgata L. in Britain. Journ. mar. biol. Assoc. U. K.
35: 149-176

OwEN, GaRETH

Bol. Fac. Filos, Cienc.

1955. | Use of propylene phenoxetol as a relaxing agent.
Nature 175 (4453) : 434 (5 March 1955)
PoweEL., ARTHUR WILLIAM BADEN
1942. The New Zealand Recent and fossil mollusca of the
family Turridae. Bull. Auckland Inst. Mus. 2: 1 - 180
RisBEc, JEAN

1955. Considérations sur l’'anatomie comparée et la classification
des gastéropodes prosobranches. Journ. Conchyl. (4) 48
[95] (2): 45-82; 22 text figs. (30 October 1955)

RosINsON, ELIZABETH

1960. Observations on the toxoglossan gastropod Mangelia
brachystoma (Puriprt). Proc. Zool. Soc. London 135:
319 - 338

ScHILDER, FRANZ ALFRED
1947. Die Zahl der Prosobranchier in Vergangenheit und
Gegenwart. Arch. Molluskenk. 76: 37 - 44
TuHorson, GUNNAR

1946. | Reproduction and larval development of Danish marine
bottom invertebrates, with special reference to the planktonic
larvae in the Sound (@resund) . Medd. Komm. Havunder-
sog., Kbh., Ser. Plankton, 4 (1): 1-523

YoncE, CHARLES MAuRICE

1962. On the biology of the mesogastropod Trichotropis can-
cellata Hinps, a benthic indicator species. Biol. Bull. 122:
160 - 181

Page 188

THE VELIGER

Vol. 10; No. 2

Enzymatic Defenses of Certain Snails Against Metal Ions

BY

LINDSAY R. WINKLER

AND

LOIS WONG CHI

College of the Desert, Palm Desert, California 92260
and California State College Dominguez Hills, Gardena, California '

INTRODUCTION

LITTLE STUDY HAS BEEN GIVEN to the defenses of snails
against heavy metal ions even though it has long been
known on a pragmatic basis that aquatic mollusks are
highly sensitive to copper ions. Furthermore, copper ions
have been used with some success against aquatic snails
as a means of control. Since some snails appear to be more
resistant to copper used as a control a study to assay
possible defenses in one of these more highly resistant
species should be of some general interest to snail physio-
logy.

The shell is the first line of snail defense which protects
the greater part of the otherwise poorly covered body from
membrane-contact with the external environment. If the
snail is an operculum-bearing prosobranch the ability to
isolate itself from the surrounding environment is in-
creased. Any ability to metabolize anaerobically further
extends the period during which the snail can remain
self-imprisoned during ionic crises in the surrounding en-
vironment and thus enhances the probability of ultimate
survival.

During a previous study using Oncomelania formosana
(Pitspry & Hrrase, 1905) as the experimental animal it
was determined (WINKLER & Cut, 1964) that certain
cells free in the hemocoel picked up copper during chronic
poisoning by very low levels of copper added to the
environmental bath provided for these amphibious proso-
branch snails. During the study it was further observed
that snail squashes exposed to copper solution turned blue.
Immersing live de-shelled snails in copper solutions simi-
larly caused light blue areas to appear on the surface of
the mantle, midbody and even on areas of the foot and
digestive gland. The possibility that this represented a

: This investigation was supported in part by grant No. AI-02705-07
from the National Institutes of Health. Part of this study was
conducted at Immaculate Heart College, Los Angeles, California.

defensive reaction of protective value to the snail served
as the take-off point for the present study.

MATERIALS anp METHODS

Snails used in this study were obtained from stock dishes
of Oncomelania formosana (PitsBry & Hirase, 1905)
reared and maintained in our laboratory as previously
described (WaGNER & Cut, 1959). In order to remove the
protection provided by the shell and operculum and to
produce a degree of uniformity in the results all snails
were de-shelled prior to use. The living snails were placed
in 1% copper sulfate for the empirically chosen time peri-
ods of 3, 10, and 30 minutes, and 1, 3, 5 and 7 hours. This
concentration was chosen to speed results since de-shelled
snails do not live more than 24 hours. Immediately after
exposure snails were killed and fixed in acetone at deep-
freeze temperatures after which they were allowed to
return to room temperature for storage. Snails destined
for histochemical study were removed from acetone to
two changes of tertiary butanol for a total of two hours,
and then to benzene with one change for a total time of
15 minutes. From the last benzene the snails were trans-
ferred to about 5 ml of molten paraffin for 25 hours,
followed by conventional embedding procedures. This
abbreviated procedure produced very adequate results
with the small amount of tissue found in these snails.
After serial sectioning and subsequent mounting on glass
slides the sections were either left unstained for direct
observation and chemical study of the blue colored objects,
or were treated with the rubeanic acid histochemical re-
agent for demonstration of micro amounts of copper using
a modified method similar to that of UzMan (1956). Eosin
was used as counterstain. The carbonic anhydrase method
of Gomort (1952) employing cobalt was used on de-shelled
acetone-fixed snails for confirmation of the presence of
this enzyme. Snails were killed in subzero acetone, then

Vol. 10; No. 2

THE VELIGER

treated with the reagents, after which they were sectioned
and mounted. Parallel procedures using copper were car-
ried out for comparative purposes.

Quantitative determination of the rate of concen-
tration of copper in the snail was made by exposing 6
de-shelled snails for each of the above mentioned time
intervals. These were then rinsed and stored in cold
acetone. After storage the acetone was removed with a
minimum of 4 hour air evaporation. A modified form of
the procedure of GuBLER ef al. (1952) was used for
spectrophotometric determination of copper. Individual
snails, after being weighed to 4 decimal points on a Mettler
H-15 balance, were placed in the courvette, after which
10 drops of 2N HCl were added and left to stand for 20
minutes. The solution was then made slightly alkaline with
4 drops of 1:3 ammonium hydroxide and the level was
brought to a total of 3.2 ml with de-ionized water. After
setting the 100% transmittance reading of the Spectronic
20 with this solution, 3 drops of diethyldithiocarbonate in
0.1% solution was added and the resultant reading was
recorded from the photometer at 440 w. If this transmit-
tance reading was below 10 on the scale, an additional 3.2
ml of de-ionized distilled water was added and a second
more accurate reading was obtained by halving the sub-
sequent reading as indicated on the spectrophotometer.
The copper present was then read in y of copper from a
curve previously prepared, using the same procedure on
graded dilutions made froma standard copper solution. The
addition of sodium pyrophosphate and citrate recom-
mended by the original writer was omitted, since the
control snails gave a uniform reading of under one y of
copper-equivalent of the combined iron and copper impu-
rities. The results in y per snail were finally converted to
y of copper per milligram of snail tissue and plotted
against time (Figure la).

In order to determine the rate of copper concentration
by carbonic anhydrase when the protective, living processes
of the snail were removed, snails were killed by immersion
in subzero acetone which did not deactivate the enzyme
systems, and were then stored at deepfreeze temperatures
for at least 12 hours. They were then exposed to the
copper bath for the same time intervals as used in the
previous experiments. After removal each was stored in
acetone until it was quantitated by the procedure previous-
ly described (Figure 1b).

RESULTS

The most significant observation noted on the sectioned
material was the presence of large numbers of birds-egg-
blue crystals which were effervescent when weak hydro-
chloric acid was introduced under the cover slip. This,

° I 2 3 4 5 6 7

Time in hours

Figure 1
Curves showing rate of copper uptake by Oncomelania formosana.
Curve a is that of de-shelled but living snails; 5 is the result when
the snails were killed by storage in cold acetone — a procedure which
but slightly reduced the enzyme activity. Each point represents
averages for 6 and 4 snails on the two curves respectively. The
copper levels of individual living snails of curve a were wider in
range than those of the dead snails b, especially in the flat portion

of the curve.

combined with their color, identified them as CuCOs.
These crystals were found covering the surfaces of the
mantle, central body wall and, to a more limited extent,
the foot and digestive gland. Furthermore, they also oc-
curred in the peripheral tissucs in specimens exposed for
the longer time periods. Associated with these crystals
were numbers of the free, spherical cells described in the
previous paper ; however, in these unstained sections many
showed the same light blue color as the crystal formations.
In addition, careful examination revealed the presence of
one to many crystals contained in most of those showing
the blue color. The Gomorr (1953) carbonic anhydrase
activity test established these cells along with certain areas
of the body wall, specifically those covering loose connec-
tive tissue, as the principal sites of activity of that enzyme.

The copper penetration observed in the rubeanic acid-
stained sections of snails exposed only briefly was pre-
dominantly in the midbody region with very little in the
foot or in the digestive gland despite the very thin walls
of the latter. Under limited exposure periods very few
copper carbonate crystals were observed. Few of the pro-
tective cells (p-cells) were noted peripherally but these
increased in number with time as did also the formation
of copper carbonate.

Page 190

THE VELIGER

Vol. 10; No. 2

The chief loci of copper carbonate as well as of clumped
p-cells were in loose connective tissue, usually immediately
below the thin outer walls. It was difficult to separate
the crystal formation from the clumps of these p-cells.
Particularly was this true under the high sensitivity of
the rubeanic acid test which tended to blur the clumps
of cells. Unstained slides were distinctly superior under
these conditions.

As time of exposure progressed concentration rose in
the peripheral connective tissue of the mid-body area.
Only in exposures above 3 hours was there considerable
invasion in other than peripheral tissues of the foot and
digestive gland.

The extent of the immobilization of soluble copper and
its concentration into insoluble copper carbonate was
studied using snails killed in subzero acetone in order to
retain enzyme activity. When the amount of copper fixed
in each snail was quantitated, averaged and graphed
(Figure 1b) it was observed to rise sharply to 16y per
mgm in those with 10 minutes exposure. Thereafter the
curve sharply leveled, then continued its rise at a gentler
slope terminating at 27y in snails of 7 hour exposure.
When the living processes were superimposed by not killing
the snails till after copper exposure (Figure la), the
curve rose considerably less precipitously to a little over
10y of copper per mgm during the first half hour. This
was followed by 44 hours in which little change was
observed but in which considerably more variability oc-
curred between individuals of the same cxposure incre-
ment. The resistance of the living process appeared to
break down somewhere between 5 and 7 hours, allowing
a rapid rise to approximately the same level as that of
the snails exposed after having been killed.

DISCUSSION anp CONCLUSIONS

The observed formation of insoluble CuCO:; seems to be
a part of the snails’ defenses rather than only a byproduct
of shell formation since the centers of carbonic anhydrase
activity revealed by the Gomori test were principally the
p-cells which were widely spread throughout the snail
body. Copper used in place of cobalt in a parallel pro-
cedure also revealed one factor that appears in distinct
contrast between the two ions. While cobalt localized
itself in the sites of carbonic acid activity with very little
attaching to the tissues, copper attached itsclf extensively
on all tissues This is not the result of greater sensitivity of
the test as the same “‘sulfide procedure” for detecting the
presence of cobalt was used for the copper in this proce-
dure. One exception to the above statement was the
strong affinity for the primary reproductive organ evident

by heavy differential staining of the latter tissue when
using the cobalt procedure.

The p-cells exist in various sizes and may be observed
thus in the various clumps where they occur. When iron
was substituted for the copper, the p-cells were not masked
by staining of surrounding areas. It was then possible to
locate what appeared to be the sites of origin of the
p-cells in pockets of the loose connective tissue. In these
pockets large numbers of miniature p-cells of homogeneous
sizes were observed. These were interpreted to be devel-
oping p-cells.

When iron or calcium was substituted for copper, the
results were less definitive. In both cases the cells gave
the stain characteristic of the ion involved, but crystal
formation did not occur. Calcium was not noted bound to
the tissues, but iron, as was the case also with copper, was
observed extensively on the surface of all the tissues.

The curves which compare the concentrations of copper
in the snail emphasize the modifying influence of the
living process and the effectiveness of the active transport
system at keeping the system relatively free of copper
even under the extremes of trauma and high copper con-
centration. What does find its way into the hemolymph
of the snail is picked up by the p-cells.

The abrupt leveling off of the rise in snails without the
living process at 16y (Figure 1b) probably results from
depletion of carbon dioxide from the environment. The
remaining even slope may represent the diffusion of carbon
dioxide into the solution from the surrounding environ-
ment, perhaps augmented as the result of the binding of
copper by the dead tissues.

The less steep rise during the first hour apparent in the
curve of the living snails (Figure 1a) apparently repre-
sents the formation of crystals on the surface and in the
peripheral p-cells. Copper penetration then remains static
until sometime after the 5" hour. The rise after the 5™
hour would logically seem to be the result of the internal
protein binding or the depositing of copper crystals in
arcas previously unavailable because of the resistance of
the living process.

The inspection of the graphs offers little comfort to
mollusciciders. The resistance here observed against these
high concentrations of copper used on snails having al-
ready been deprived of their first line of defense would
scem to indicate their ability to withstand relatively long
sieges of the lower concentrations normally used as mol-
luscicides by snails possessing all their faculties.

It is not known how far the process of crystal formation
can actually protect these snails from death but the com-
bination of an impermeable shell with functional opercu-
lum, a considerable anacrobic capability and the enzymatic

Vol. 10; No. 2

deactivation of copper seem to make this prosobranch
snail relatively immune to ordinary acute ionic attacks
which might be lethal to less highly favored snails.

LITERATURE CITED

Guster, C. J., M. E.Lanry, HELEN ASHENBRUCKER, G. E. CarTH-
WRIGHT, & M. M. WINTHROBE

1952. Studies on copper metabolism. I. A method for deter-

mination of copper in whole blood, red blood cells, and plasma.

Journ. Biol. Chem. 196: 209 - 220

THE VELIGER

Page 191

Gomort, GEORGE

1953. Microscopic histochemistry. Univ. Chicago Press,
Chicago
Uzman, L. Lauut

1956. Histochemical localization of copper with rubeanic
acid. Laborat. Invest. 5: 299 - 305

Wacne_r, Epwarp D. & Lois Wone Cui

1959. | Methods on the rearing of the snail Oncomelania.
Suppl. Trans. Amer. Microsc. Soc. 78: 421 - 423

WINKLER, Linpsay RosBert « Lots Wonc Cur

1964. Defensive mechanisms of the Schistosome snail host
Oncomelania formosana against copper sulfate in adult and
egg stages.

Amer. Journ. Trop. Med. & Hyg. 13: 897 - 902

Page 192

THE VELIGER

Vol. 10; No. 2

The Radulae of Nine Species of Mitridae

(Mollusca : Gastropoda )

BY

JEAN M. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plate 19; 9 Text figures)

INTRODUCTION

IN RECENT YEARS more and more attention is being given
to the study of mitrid radulae. CERNoHoRSKy (1966)
has provided the largest and most comprehensive recent
work, covering about 50 species; even so, the total of
known, accurately identified mitrid radulae still stands at
only around 100 species. Helpful as this is to workers in
this group, we have to admit that it is only a beginning,
in a family comprised possibly of some 500 to 600 Recent
species. It has been seen that our new knowledge of
many of the radulae is negating the earlier generic and
subgeneric placements of certain species whose taxonomic
assignments were based on shell morphology alone. Con-
sequently, it is evident that at some future date the
Mitridae will need to be thoroughly revised, with totally
new concepts of the genera and subgenera different from
those we know now. However, with such a slow trickle of
information as is presently becoming available, it would
seem to me premature to attempt to make such drastic
changes at this time, for the radula patterns of approxi-
mately 85% of the species are still unknown. It may well
be that the radula will prove to be of greater (or, possibly,
lesser!) importance than is accepted today. Certainly the
morphological characters can not be totally ignored, for it
is seldom indeed that live-collected material becomes
available for radula study and it may be many ycars
before a relatively complete knowledge of all the species
can be obtained. It seems to me that the conchologists
must, therefore, continue to arrange their collections in
groupings of “look-alike” shells; i. e., as nearly as possible
in the recognized genera and subgenera, however arbitrary
and artificial a system this may be according to the
radular characters.

Acasc in point is Strigatella columbellaeformis (KIENER,
1839). The morphological features of this species fit the
description of the typical Strgatella so well as to be
almost amusingly “supertypical.” Its features seem an
exaggerated version of SwAINSON’s description: “Size

very small; spire thick, obtuse; outer lip thickened, and
often reflected in the middle; aperture smooth.” CERNo-
HORSKY (1966a) has shown that the radula pattern of
this species renders its placement in Sérzgatella incorrect,
and that it should be assigned more correctly to Mitra
s.s. It comes down, then, to the very practical con-
sideration of how museum curators and private collectors
shall curate their collections, with such apparently
“typical” forms really belonging, according to their
radular characters, alongside morphologically com-
pletely dissimilar forms. This problem is negligible
in small collections, but with a collection of Mitridae
numbering into the hundreds of species and thousands of
specimens it becomes important to have the similar species
grouped together.

If a student were, for example, to bring to a large
museum for identification a typical small, drab, brown
strigatelliform shell, the most logical place to seek its
identity would be among the Strigatellas. Unless the
curator happened to be a specialist in Mitridae he would
be more than likely to overlook a comparison of the un-
known form with “Mitra” columbellaeformis which had
been placed, in accordance with its radular pattern,
together with such typical mitrid forms as the showy,
red-spotted, elongate and pointed Mitra mitra (LINNAE-
us, 1758), and not among the Strigatellas. Only by a
slow, painstaking comparison with every species in the
Mitra collection could an identification be made under
this system, whereas the smaller sub-grouping of all shells
having Strigatella characters would normally make it casy
to locate the name within a relatively short time.

It seems that the whole question may already have
resolved into an argument between the supporters of the
shell-morphology school of thought and the champions of
the radula pattern, in somewhat the same way Cypraca
collectors have their “lumpers and splitters’ — though
for slightly different reasons. Both may be correct in their
different views; who can say, at this early stage, which has

Vol. 10; No. 2

THE VELIGER

Page 193

the ultimate single answer? I believe we must continue to
strive for the solution by continuing to study additional
radula material, but not to make the final decision until
vastly larger numbers of species have been studied. At
the present time only about 15% of the mitrid radulae
are known; it would seem to me that nearer 90% should be
known before the radical revision can be undertaken that
will decide the question once and for all.

To add to our slowly increasing series of radula studies
in Mitridae, I offer here illustrations of the shells and
radulae of nine species of Mitra and Vexillum. Seven of
these are either previously unrecorded or at least not
specifically verified through references to adequate illus-
trations. Two radulae (Mitra idae MeEtvity, 1893 and
Strigatella tristis (BrRopERIP, 1836) ) have recently been
figured by CerNoHorSKy (1966) but without accompa-
nying illustrations of the shells. My work corroborates
his findings on these two species.

MATERIAL anp METHODS

The radula preparations were made by me during the
past three years, whenever appropriate fresh material
became available: the radulae were extracted through the
use of potassium hydroxide solutions, dehydrated in al-
cohol series, stained with Acetocarmine and mounted in
Euparal on glass slides. The text figures were prepared for
publication by Mrs. Emily Reid.

OBSERVATIONS

Study of these radulae resulted in no unexpected changes
of assignment of the species to their respective genera,
with the exception of Mitra (Cancilla) verrucosa (REEVE,
1845) (see Plate 19, Figure 7). CeERNoHorRsky (1965)
tentatively placed this species in Vexillum, but while its
morphology does present a conflicting set of characters, its
placement in Cancilla is justified on morphological grounds
as well as radula characters. It is a puzzling species, for
the reason that it bears spiral rows of prominent nodules
throughout; from one viewpoint it could be said that the
nodules are aligned in axial rows (making it eligible for
assignment to Vexillum), whereas from another viewpoint
the nodules could be considered to be arranged spirally
in concentric rows. After a good deal of study when I
first added this species to my collection several years ago,
I concluded that the axial alignment was more or less
accidental, and decided to place it among the Cancillas
on the basis of the concentricity of the rows of nodules;
the radula now supports this decision. The nodules are an
atypical feature of most species of Cancilla, but the con-

centric sculpture is a character that places it without
question in that subgenus.

io Mitra idae MELvILL, 1893
(Plate 19, Figure 1)

Animal: Foot, body, siphon and tentacles pure porcelain-
white. Eyes situated at base of tentacles. Sexes separate.

Figure 1
Half-Row of Radula of Mitra idae MELVILL
Specimen collected in 50 feet off Point Loma, San Diego, California
by Glen Bickford

2. Mitra belcheri Hinps, 1832
(Plate 19, Figure 2)

Animal: Unknown (received in dried-out condition).

Figure 2
Half-Row of Radula of Mitra belcheri Hinps
Specimen trawled in deep water off Guaymas, West Mexico
by shrimp fishermen

3. Mitra zaca Stronc, HANNA & HERTLEIN, 1933
(Plate 19, Figure 3)

Animal: Unknown (received in dried-out condition).

Figure 3
Half-Row of Radula of Mitra zaca Stronc, HANNA & HERTLEIN
Specimen trawled in deep water off Guaymas, West Mexico
by shrimp fishermen

4. Mitra terebralis LAMaRcK, 1811
(Plate 19, Figure 4)
(synonyms: Mitra incompta Licutroot, 1786; M.
tessellata Martyn, 1786)
Animal: Top of foot mottled with tan. Base of foot cream,
tinged with reddish brown. Tentacles pale tan tipped with

Page 194

THE VELIGER

Vol. 10; No. 2

white. Siphon banded proximally from the tip with white,
brown and light tan, each color grading into the next
(C.S. Weaver, pers. comm.). Length 80.9mm, width
20.8 mm. leg. C. S. Weaver, Oahu, Hawaii, 1961.

Figure 4
Half-Row of Radula of Mitra terebralis LAMARCK
Specimen described above.

5. Mitra aurora Donrn, 1860
(Plate 19, Figure 5)

Animal: Unknown.

ee es

Figure 5
Half-Row of Radula of Mitra aurora DoHRN
Specimen collected in 15 feet of water, in sand under dead coral
off Barber’s Point, Oahu, Hawaii; leg. C. S.Weaver

6. Strigatella tristis (BRODERIP, 1836)

(Plate 19, Figure 6)
Animal: Foot, body, siphon and tentacles creamy-beige;
eyes situated at base of tentacles.

Figure 6
Half-Row of Radula of Strigatella tristis (BRODERIP)
Specimen collected at Puertecitos, Baja California, Mexico;
leg. Fay Wolfson

Ce Mitra (Cancilla) verrucosa REEVE, 1845
(Plate 19, Figure 7)

Animal; Unknown.

50 p

Figure 7
Radular Row of Mitra (Cancilla) verrucosa REEVE
Specimen collected at Mauban, Quezon, Philippines;
E G. Dayrit, don.

8. Mitra (Cancilla) hindsti Rerve, 1844
(Plate 19, Figure 8)

Animal: Unknown.

Figure 8
Half-Row of Radula of Mitra (Cancilla) hindsti REEVE
Specimen collected in 20 fathoms off Bacochibampo Bay,
West Mexico. leg. Gale G. Sphon, Jr.

Q. Vexillum taeniatum (LAMARCK, 1811)
(Plate 19, Figure 9)

Animal: Unknown.

(ne

5° B

Figure 9
Radular Row of Vexillum taeniatum (LAMARCK)

Specimen collected by A. Jennings at Akuilau Island, Fiji
and kept in his aquarium until it died. He then generously sent it
to me intact so that I could obtain the radula without damaging the

shell; the animal had completely decomposed. This is the

specimen figured on Plate 19, Figure 9.
Length 50.9 mm, width 10.9 mm.

Explanation of Plate 19

Figure 1: Mitra idae MELVILL, 1893 (x 14)

Figure 2: Mitra belcheri Hinps, 1832 (x 4)

Figure 3: Mitra zaca Stronc, HANNA & HERTLEIN, 1933 (x 4)

Figure 4: Mitra terebralis LAMarRcK, 1811 (x 2)
Figure 9:

Figure 5: Mitra aurora DourN, 1860 (x 2)

Figure 6: Strigatella tristis (BRopERIP, 1836) (x 2)
Figure 7: Mitra (Cancilla) verrucosa REEVE, 1845 (x3)
Figure 8: Mitra (Cancilla) hindsii REEVE, 1844 (x 23)

Vexillum taeniatum (LaMaRcK, 1811) (x)

[J. M. Cate] Plate 19

Figure 8

photographs by JEAN M. Cate

Vol. 10; No. 2 THE VELIGER

LITERATURE CITED

CrrNonorsky, WALTER OLIVER
1965. The Mitridae of Fiji (Mollusca:Gastropoda). The
Veliger 8 (2): 70-160; plts. 13-23; 11 text figs; 1 map;
2 tables; index (1 October 1965)
1966. A study of mitrid radulae and a tentative generic ar-
rangement of the family Mitridae. The Veliger 9 (2): 101
to 126; 47 text figs. (1 October 1966)
1966a. Die Radula Platten einiger wenig bekannter Prosobran-
chier (Mollusca - Gastropoda). | Arch. Molluskenk. 95 (5-6) :
275 - 277; 3 text figs. (22 December 1966)

ff
(gg
on

Page 195

Page 196 THE VELIGER Vol. 10; No. 2

A New Species of Marginella from the Coast of Brazil

BY

JEAN-JACQUES VAN MOL

Laboratoire de Zoologie Systématique, Université Libre de Bruxelles, Belgique
AND

BERNARD TURSCH

Laboratoire de Chimie Organique, E. P, Université Libre de Bruxelles, Belgique

(1 Text figure)

THIs IS THE SECOND in a projected series of papers describ- Dimensions:

ing new species of mollusks from Brazil, the first having (in millimeters) Number

appeared in Volume 7, No. 1 of The Veliger (Turscu «& Length Width of Whorls

PrerRET, 1964). Holotype 39.4 23.8 4.5
Paratype A 45.2 Zee 5
Paratype B 40.2 24.0 4.5

CTENOBRANCHIATA

Type locality: The holotype and paratype B were obtained
from the stomachs of fishes locally known under the name
VOLUTACEA of “pacamao,” caught off Fortaleza, State of Ceara,

MaArGINELLIDAE

Marginella Lamarck, 1799

Marginella (Prunum) matthewsi van Mo. & TurRscH
spec. nov.

(Figure 1)

Description: Shell large for the family, rather thin but
solid, light orange-brown with numerous fine yellowish
spiral bands. Surface glossy. Whorls about 5, including
teleoconch. Aperture ivory-white, long and rather narrow,
extending the entire length of the shell. Four prominent
columellar plications, the uppermost being separated from
a long glazed parictal callus by a well-defined sulcus. The
parietal callus is thickened at the posterior extremity of
the aperture. Outer lip thickened, showing on its inner
side a row of numerous rather indistinct denticulations on
all its length. Spire flat, nuclear whorls apparent, suture
distinct as a white line. Figure 1

Vol. 10; No. 2

THE VELIGER

Brazil, in about 20 fathoms. Nature of the bottom: numer-
ous coral heads on sand. Paratype A is from Acarau, State
of Ceara, also ex pisce.

Repositories: Holotype, Stanford University Paleo. Type
Collection no. 9856. Paratype A, in Tursch collection.
Paratype B in American Museum of Natural History, New
York.

Discussion: This species is somewhat similar to the well-
known Marginella bullata LinNAzvs, 1758, from which it
is easily separated by the following characters:

Marginella bullata Marginella matthews
Colour: rosy-gray orange-brown
Spire: covered by glassy layer apparent, suture distinct
Parietal callus: absent present
Interior of outer lip: nearly smooth numerous small teeth
Dorsal side of orange-brown, creamy, subcarinated,
labial thickening: rounded profile characteristically

expanded at anterior end

To our knowledge, Marginella matthewsi is not sym-
patric with M. bullata, whose distribution center is in the
State of Bahia. This new species is dedicated to Mr. Henry
Matthews, British Vice-Consul in Fortaleza, whose kind
and competent help greatly facilitated our collecting in
Ceara.

Page 197

Page 198

THE VELIGER

Vol. 10; No. 2

The Rediscovery of Erosaria menkeana (DESHAYES, 1863)

(Mollusca : Gastropoda)

BY

CRAWFORD N. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plate 20)

CONSIDERABLE REFERENCE has already been made in the
literature to a cowrie specics that apparently remains
clouded in synonymy. It is likely that earlier authors
lacked access to an obviously rare species for comparison.
I am certain that the shells illustrated in this paper
definitely appear to be unrelated to one another, and
should now be given adequate consideration, with one of
them, Erosaria menkeana (DrESHAYES, 1863) being re-
garded as a valid species once more.

The genus Evosaria TroscHEL, 1863, with which I
associate this species, usually has a characteristic morpho-
logy, a broad, flattened shell, with colored spotting. There-
fore, shells of Erosaria follow this general description:
shell ovate to sub-pyriform, not umbilicate, usually thickly
margined, heavily calloused; tuberculate or smooth, with
coarse teeth that do not cross the base and do not reach
the margins; usually flattened, spreading; terminals only
slightly produced, or in some cases not at all; shell may be
marginally spotted, spotted over all, blotched, obscurely
banded, or peripherally ringed with color. It would seem
E. menkeana agrees well with this generic description.

Erosaria (Erosaria) menkeana (DrEsHAYES, 1863)
(Plate 20, Figure 1, 1a)

Conch. ile Réunion, Annexe E (Moll.): 139; plt. 13, figs. 21 - 22
Syn.: Cypraca modesta (OWEN) SoweErBy (2%”), 1870
Thesaurus Conchyl. 4: 11; plt. 322, figs. 333 - 334

Locality: Mauritius (Long. 20° 18’S; Lat. 57° 36’E).
Shell Measurements: Length 20.0mm, width 13.0mm,
height 9.0mm; __ labial teeth 16; columellar tecth 13.

This species is clearly and unmistakably illustrated in
Sowersy (2°?) (1870). With the discovery of a live-col-
lected specimen, it seems to me this heretofore obscure
species should be accepted as valid. Toward this end I
offer the following illustrations and redescription.

Shell small for the genus, elongately ovate, broad,
flattened, solid, strong, thickly formed; dorsum trans-
versely rounded, laterally flattened centrally due to two
abrupt contour angles that divide the shell into three
equal parts; base and outer lip convex from margins to
aperture; terminal strongly produced abapically, less so
adapically ; aperture straight, narrow; both columella and
fossula dentate; fossula shallow, nearly obsolete; teeth
large, strong, well defined on both outer lip and base,
extending halfway to margins; margins thick, rounded,
indistinct, barely angularly shouldered above; primary
shell color milk-white; dorsum glossy, thickly flecked with
numcrous very fine chestnut-brown spots, becoming fewer
in number and minutely larger on margins, part of base
and outer lip; very faintly quadrimaculate at terminals; a
narrow portion of base, outer lip, teeth, and interstices
white.

This shell was sent to me for identification by an
Australian correspondent. The locality given was Mau-
ritius in the Indian Ocean; at the present time this locality
remains unverified. In making a search of the literature,
I found that Sowersy (2*’) (1870, plt. 322, figs. 333, 334)
provided drawings almost identical with the shell at hand.
With the aid of these hand-colored illustrations one has
little difficulty in identifying this shell as a species long
thought to be synonymous with Bistolida owen owen
(Sowersy [1°], 1837); Sowersy (2°) (1870, plt. 323,
figs. 366 - 371) ; this paper (Plate 20, Figure 2). It may be
of interest to note that ReEve (1846) and Kiener (1845)
do not list the latter species even though it had been
almost ten years since SowERBy (2°) described it.

In the text Sowersy (2”) (1870, p. 11, species 27)
discussed the species under consideration under the name
Cypraca menkeana OWEN (==Cypraea menkeana Des-
HAYES, 1863). When comparing this shell with Bistolida
owenii owenti he said, “... it does not well agree.” He
then stated further, “... the shell which Mr. Hugh Owen

[C. N. Cate] Plate 20

Figure 1
Erosaria menkeana (DESHAYES, 1863)

Left Side Right Side

Figure 1a
Erosaria menkeana (DESHAYES, 1863) (x 24)

Bistolida owenit owenii Bistolida oweni vasta
(SowerBy, 1837) (x34) (SCHILDER & SCHILDER. 1938) (x 24)

photographs by C. N. Cate

Vol. 10; No. 2

names modesta ... seems to fulfill all the conditions to
identify it with menkeana.”

As I compared this recently collected shell with the
Sowersy (2) drawings of “Cypraea menkeana” it
became increasingly apparent that the two are almost
identical, almost as if the original drawings were made
from this specimen. It is now evident that this specimen
disproves the uniqueness of the Sowersy shell, and sup-
ports the validity of this species.

I have compared the shell with Bistolida owenii owenii
and with the allopatric form B. owenii vasta (SCHILDER
& SCHILDER, 1938) (see Plate 20, Figure 3), the former
from Mauritius and the latter from Durban, South Africa.
The morphological distinction between them seems com-
plete, with menkeana even probably belonging in a
different genus, Erosaria, to which I have assigned it.

Sowersy (2*) illustrated “Cypraea menkeana” twice:
on plate 327, figure 512, a shell from Borneo (a question-
able locality since the species involved in this discussion
are now known to occur only in the western Indian
Ocean), and plate 322, figures 333, 334 (ventral and
dorsal aspects), a shell from ile Réunion, located approx-
imately 200 miles west of Mauritius. However, these two
figures clearly seem to illustrate different species, figure
512 depicting Bistolida owenti owenn and only figures 333
and 334 in fact representing Evosaria menkeana.

WEINKAUFF (1881) recognized the species in question,
using the name “Cypraea menkeana, non Sowerby,” to
identify it. He illustrated the species on plate 13 (figures
9 and 12). It is interesting to note that neither of
WEINKAUFF'S drawings agree with one another in outline
of form, although it would appear that an effort had been
made to copy Sowerby’s figures. Ignoring the poor outline
the ventral aspect agrees well, but the dorsal view is
incomplete, showing what appears to be a decorticated
upper shell surface.

Hipatco (1906) mentioned this species as a synonym,
and listed it as a sixth variety of “Cypraea owentt Gray”
[sic]. He interpreted Sowerby’s use of names for the species
as “Cypraea menkeana var. modesta.”

Tryon (1885) said, “From a careful comparison of
specimens of this shell [Bistolida oweni oweniz] with those
of C. menkeana Desh. (PI. 4, figs. 69, 70, from Deshayes;
Pl. 4, fig. 58; Pl. 5, fig. 73, from Sowerby), both of which
were received from Mr. Sowerby, I do not hesitate to
place the latter as a synonym. The only noticeable differ-
ence is in the marginal dots, which in menkeana sometimes
run together giving a solid brown appearance. Specimens
of C. owenii [sic] received from Mr. J. F Bailey, of Mel-
bourne, Victoria (Australia), are rather elongate, yellowish,
and have a faded appearance, but decorticated shells are
fawn-colored.”

THE VELIGER

Page 199

In an assessment of Tryon’s opinion in regard to these
shells it is important to bear in mind that DesHayes’
type specimen, or one similar to it, and SowErBy’s shell
used for his Thesaurus illustrations conceivably could be
in color and morphological agreement with each other.
Even though Tryon’s illustrations of Bistolida owenii
owenu are poor and hardly representative of the species,
it is fairly easy to note a difference between the species
as he separated them for purposes of comparison.

Most recently Erosaria menkeana is listed in ScHILDER
& SCHILDER (1938, p. 166, species 104A), and again in
ScHILDER (1941, p. 99) as Bistolida (Derstolida) menke-
ana, citing it as a synonym of B. (D.) owenii owenit.
However, with this rare cowrie species now available for
study, it seems likely the taxon will stand as a valid species.

LITERATURE CITED

DesHAYES, GERARD PAuL
1863. Maillard (L.), Annexe E (Mollusques), p. 139; plt., 13,
figs. 21-22 (Not. fle Réunion) (not seen)
Hipatco, Joaquin GonzaLez
1906 - 1907. Monografia de las especies vivientes del género
Cy praea. Mem. Acad. Cienc. Madrid 25: 1 - 240 (1906) ;
Kiener, Louis Caries
1845. Spécies général et iconographie des coquilles vivantes
genre Cypraea. 1: 1-165; plts. 1-57 Paris, J.-B.
Bailliére
Reeve, LovELL Aucustus
1845-1846. Conchologia Iconica, 3: Monograph of the genus Cyp-
raca, 16 plts; 1846: 11 plts.
ScHILpER, Franz ALFRED
1941. | Verwandtschaft und Verbreitung der Cypraeacea.
Arch. Mollusenk. 73 (2/3): 57-120; 2 plts.
SCHILDER, FRANZ ALFRED, & MARIA SCHILDER
1938-1939. | Prodrome of a monograph on living Cypraeidae.
Proc. Malacol. Soc. London, 23(3 - 4): 119-231.
SoweErBy, GeorcE BRETTINGHAM (1‘T of name)

1837. Catalogue of recent Cypraeidae. London; 1: 6 (not seen)

SowersBy, GEorGE BRETTINGHAM (2%? of name)

1870. Thesaurus Conchyliorum 4: 1 - 58; 37 plts. London

Roserts, S. RAYMOND
1885. | Monograph of the Cypraeidae. In: Tryon, GEORGE
WasuINGTON, Jr., Manual of Conchology 7: 153 - 198; plts.
1-23; Philadelphia
WEINKAUFF, HEINRICH CONRAD
1881. Die Gattungen Cypraca und Ovula. Conchylien-Ca-
binet (2nd. ed.), Martini « CHEMNiTz. Niirnberg, pp. 1 to
230; plts. 1-53A

Page 200

THE VELIGER

Vol. 10; No. 2

Notes on Cephalopods from Northern California

BY

ROBERT R. TALMADGE

Field Associate, Department of Invertebrate Zoology,
California Academy of Sciences, San Francisco, California 94118, and Eureka, California

In 1912 S. Strr-MAN BERRY PUBLISHED a paper on the
cephalopods of western North America, a painstaking and
comprehensive study, long out of print, but still the only
accurate account available today, covering this interesting
class of mollusks in the designated area. Ropson’s (1929-
1931) two part monographic work was limited to the
Octopoda, and Pickrorp (1964) presented a detailed
study of a single polymorphic species of octopod. There
have been a number of taxonomic and distributional
notes published, but as far as I could learn, there has
been no detailed study covering any one portion of
the California coast.

The following brief notes are presented not as a taxo-
nomic discussion, but rather to bring together new and
additional information on the basic ecology, distribution
and, to some extent, the physical characteristics of several
cephalopods found in two degrees of Latitude on the
extreme northern coast of California (Lat. 39° 30’N to
41°30’N). All of the data are based upon specimens
deposited in the collection of the California Academy of
Sciences or in my own collection. The specimens for the
most part were obtained by the commercial dragboat
fleet operating out of Humboldt Bay, California.

At this time I wish to express my appreciation to the
men who have so willingly brought me the specimens for
many of the present records. I also wish to express my
appreciation to the staff of the California Academy of
Sciences for assistance given me while I was at the Acad-
emy and for the permission granted to include data from
the Academy Collection.

COMMERCIAL FISHERIES

There are no commercial fisheries as such for either squid
or octopus on Humboldt Bay. I have noted squid, Loligo
opalescens Berry, in one of the local fish markcts, but
learned that these had been shipped from Monterey,
California. Squid used for bait by the crab fishermen is
brought from Monterey and recently also from Japan. A
few octopus are brought ashore by the fishermen for
display in a local commercial aquarium, for specimens, or
as an epicurean item. Although a few squids have been

caught off this coast, the numbers are small and not
sufficient to support a bait fishery.

DAMAGED anv DISTORTED SPECIMENS

The often torn or mangled condition of many benthic
cephalopods may be easily understood if a person observes
the operation of capture. All cephalopods have soft bodies,
but many of the benthic species have a body so soft that
the tissues are gelatin-like in texture. This is possibly
useful to the animal as it moves voluntarily from layer to
layer in the depths. It would allow the pressure to equalize
both within and without the animal tissues. However,
when the cephalopod is caught in the fishing net, mixed
in with perhaps several tons of fish, and then rapidly
brought to the surface, it is difficult or probably impossible
for the animal to equalize the internal pressure, and a
nearly explosive decompression occurs in the cephalopods
as it does in the fish brought from the same depths. The
ultra-soft tissues then are torn easily by the net, by pro-
jections on deck, or among the fishes themselves.

Distortion occurs from such pressure release as well as
from several other factors. In order to keep this distortion
to a minimum, special jars of preservatives were placed on
a number of the Otter Trawlers so that all of the specimens
would be field-preserved in a similar preserving medium.
This was also necessary in order to harden the gelatin-like
tissue of the deeper benthic species so that they would
be available for later study.

COLORATION

Reference to the coloration of cephalopods is, at best,
rather uncertain. The species not only are able to change
color at will, but vary considerably from life to death,
and from death to a preserved state. The preservatives
themselves may affect coloration, alcohol acting as a
bleaching agent, to name one cxample. Therefore, the
preservatives furnished to the vessels actually were a
mixture, and every effort was made to secure any speci-
mens brought in within a week in order to obtain a better
understanding of the coloration. In addition, comparison

Vol. 10; No. 2

with living animals was made whenever possible. One
important factor was noted: in life the animals are trans-
lucent or nearly transparent, with the colored chromato-
phores standing out in a starkly contrasting manner; at
death this translucent body became opaque and when
preserved turned a dead white. In some species the rusty
browns became darker and in others lighter. But most
very old specimens examined were a dead opaque white.

SPECIES

At the present time there are 10 species of cephalopods
in the collections of the California Academy of Sciences
and in the Talmadge Collection which were obtained
from off this region in northern California. Since many
species are pelagic, it is quite possible that additional
species will be found in time. The following is a list of the
species represented in the collections mentioned.

Octopus dofleini martini Pickrorp, 1964

I use this taxon, but the species is also known as Octopus
punctatus Gass, 1862; O. hongkongensis auct., non HoyLe,
1885; O. apollyon auct., et (BErry, s.1.), non (BERRY,
s.s.), 1912. As Pickrorp has probably worked with
more specimens of this, the largest known species, than
any other worker, I follow her diagnosis. Berry (1912)

was perhaps the first worker to hint at an eastern : western .
Pacific separation and proposed the name O. appolyon/

for the western Pacific population, but restricted the type
to an Alaskan specimen. Pickrorp agreed with BERRY
and designated O. dofleini apollyon Brrry as the Alaskan
subspecies. It is evident that Berry realized the situation,
but lacked sufficient material to complete his diagnosis.

In northern California the species is found from the
deep intertidal down to around 40 fathoms, usually in a
hard mud or rocky substratum. This is one of the “hard”
bodied species and sets firmly in preservatives. In life the
species is usually tan or grey, darker on the dorsum, and
when placed in preservatives will often turn to a purplish
brown. Although it is reported to attain great size, (BERRY,
1912, quoting DAL1 as stating that the species will reach a
spread of 28 feet at Sitka) I have never seen one that
would exceed 18 feet when spread on a dock.

Octopus californicus (BERRY, 1911)

The “Anna W”, one of the larger dragboats on Humboldt
Bay, has brought me several specimens taken north of
Eureka in less than 100 fathoms on a sandy bottom. The
captain noted that the coarse nodes present on preserved
specimens were not noticeable in life, but became quite
apparent when the animal was placed in preservatives.
The coloration is a dark rich reddish brown, and becomes

THE VELIGER

Page 201

somewhat lighter in preservatives. Both sexes of this small
species were present, and the hectocotylized tip of the
third right arm of the male appeared to be detached in
several specimens, A closer examination revealed that
where the missing tip had been, a minute growth was
forming, probably a regeneration of the male portion of
the arm.

Octopus leioderma (Brrry, 1911)

This is a small, smooth, “soft” octopus usually taken on
a muddy sand substratum in about 50 fathoms of water.
Usually the animals are considerably distorted and dam-
aged, as they are found in the narrow crevices of the
deck grates and about the scuppers. There is a definite
constriction between the body and the head, the head and
the arms, and another, less pronounced ridge or rim on
the horizontal plane of the body, which resembles a
mold-mark on a plastic toy. In coloration the animal is
a pinkish-tan, livid, and when placed in preservatives it
darkens to a brownish-red, lighter on the ventral surface.
Both males and females were obtained from the “Winga”
and the “Anna W”.

Opisthoteuthis californicus Berry, 1949

The type locality for this interesting and non-octopod ap-
pearing species is off Humboldt Bay and the majority of
the known specimens has been obtained from off this
portion of the California coast. The species may attain
over 2 feet in diameter, and in life looks like a translucent
jelly-fish veined with rust on the dorsal surface; ventrally
the animal is entirely rust color. The minute fins on the
dorsal surface are puzzling structures. The body is ex-
tremely soft and the name the fishermen use, “jelly-fish-
octopus,” seems to me to be more appropriate than the
official name “flapjack devilfish.” Most of the specimens
were taken in excess of 300 fathoms, but it is known to
occur at times in depths of only about 100 fathoms. All
material in my collection came from a very soft mud sub-
stratum and was obtained for me by the “Ina,” “Flicker”
and “Anna W”.

Loligo opalescens Berry, 1911

A few specimens taken off Redding Rock, north of Eureka,
have reached me through the “Anna W,” fishing in about
100 fathoms. None of the specimens exceeded 200 mm in
length; they were taken singly, not in a school of squid.
In life the animal is truly opalescent, a pinkish-tan, trans-
lucent, with the chromatophores standing out in stark
contrast. In death the body is opaque and when placed in
preservatives, the body turns white with the red chromato-
phores turning dull brown.

Page 202

Meleagroteuthis hoylei Purrer, 1900

A single, slightly mangled specimen (a female), 118mm
in length, including the arms but not the tentacles, was
caught in the net of the “Flicker” in a depth of over 300
fathoms. This species is well figured by Berry, 1912, who
records it from Monterey, California. There are 2 very
small specimens, bleached white, in the collection of the
California Academy of Sciences; they were taken on the
Cordell Bank, Marin County, California. The recent speci-
men was a dark purple in life with small tan, grain-like
nodules on the dorsal surface, especially on the body, and
until the specimen was placed into preservative, these
grain-like markings were actually extruded and could be
felt with the fingers of one’s hand. In preservative, the
color has changed to a rich purple with tiny tan grain-like
markings on the body, but the small nodes now no longer
may be noted.

Galiteuthis pyllura Berry, 1911

The dragboat “Jna” obtained the rather diagnostic “leaf”
tail of a specimen of this species amid the debris left on
deck after a drag. The captain and crew noting that this
was animal tissue new to them, saved the bit of flesh by
freezing it in their refrigerator. In life or when found, the
tissue was nearly translucent with dark reddish macula-
tions, which in freezing turned an opaque dead white with
dark red markings. Based upon Berry’s measurements of
the type specimen, the Humboldt specimen must have been
350mm long, much larger than Berry (1912) recorded.
There are some specimens, again bleached, from the Cor-
dell Bank, in the collection of the California Academy of
Sciences.

Meroteuthis robusta (VERRILL, 1876)

The giant squid, which is reputed to attain a length of
50 feet, has been taken by dragboats off Eureka, California.
For obvious reasons, only a major research center has the
necessary facilities to store preserved specimens. I have
seen no fresh or living examples, but have examined some
on docks at various times; I also have examined the
preserved specimens in the California Academy of Sciences.

me
RS

PUT

pT tnentittas

MI

THE VELIGER

Vol. 10; No. 2

The dragboat fishermen have no love for these creatures
as their size makes them difficult to handle and they often
damage the nets. In coloration the preserved specimens
remind me of a dark Loligo opalescens.

Rossia pacifica Berry, 1911

“Short-bodied squid” is the vernacular name for this small,
ultra-soft-bodied cuttlefish. It is usually taken in about
100 fathoms, on a sandy mud to mud bottom; it is trans-
lucent grey on the dorsal surface and greyish white on
the ventral surface. It is badly mangled when mixed in
with fish. It is perhaps more common than noted as the
small size and very soft body allow this animal to wash
between the deck grates and out the scuppers without
being seen.

Gonatopsis sp.

There are 2 specimens of this eight armed squid in the
collection of the California Academy of Sciences, taken
by the “City of Eureka” in 200 fathoms of water off
Eureka. To some extent these preserved animals resemble
the giant squid, but they are much smaller; the 2 body fins
are quite strong and the coloration is the same brownish
with dark brown maculations.

LITERATURE CITED

Berry, SAMUEL STILLMAN
1912. A review of the Cephalopoda of western North America.
Bull. Bur. Comm. Fisheries 1 (XXX), Doc. 761: 267 - 336;
plts. 32 - 56; with text figs.

Pickrorp, GRACE E.

1964. Octopus dofleint (WULKER). Bull. Bingham
Oceanogr. Coll., Peabody Mus. Nat. Hist. 19 (1): 5-70

Rosson, G. C.
1929. A monograph of the Recent Cephalopoda (based upon

the collections in the British Museum (Natural History) ).
Part I: v-xi. 1 - 236; plts. 1-7

1931. A monograph of the Recent Cephalopoda (based upon
the collections in the British Museum (Natural History) ).
Part 2: vii- xi, 1 - 359; plts. 1-6

Vol. 10; No. 2

THE VELIGER

Page 203

NOTES & NEWS

Note on the
Northward Spreading of

Mya arenarta Linnaeus in Alaska

BY

JAMES B. GROSS'

Douglas Marine Station
P.O. Box 185, Douglas, Alaska 99824

Mya arenaria (Linnaeus, 1758) is apparently spreading
northward along the Pacific shores of North America in
a dispersal similar to that which occurred in northern
Europe in the 17" Century.

In recent field trips to determine its distribution in
nearby waters, the author found it living in sparse to dense
concentrations in favorable mud and sand flats along
Stephens Passage, the northern reaches of Chatham Strait
and the Lynn Canal, all in southeast Alaska. It is expected
that further distribution and population studies will reveal
that Alaska has acquired a sizeable and heretofore un-
known harvestable asset.

At present the earliest evidence of its arrival in these
waters comes from a single pair of valves in the collections
of the Bureau of Commercial Fisheries Biological Labo-
ratory at Auke Bay, Alaska. This specimen was collected
by J. J. Gonor in Oliver Inlet, Stephens Passage, on April
22, 1963. The specimen was in its 7" year of growth,
placing the earliest known date of arrival here as 1956.

The Earthquake Study Team, under the direction of
Dr. G Dallas Hanna in 1965 collected Mya arenaria shells
in Prince William Sound. Specimens of this collection were
loaned to the author by the California Academy of
Sciences. The oldest of these valves, one collected at
Orca Bay, was in its 6" year. Presuming that it was
cast on the beach during the convulsions of the earthquake
of March 27, 1964, an arrival date of 1958 can be
estimated for that area.

1 Contribution No. 34 from the Institute of Marine Science,
University of Alaska.

It should be noted that the California Academy of
Sciences specimens are labelled Mya japonica (Jay).
Collections of shells of the representative subgenus from
widespread areas and now in the author’s possession,
show ranges of variability tending to invalidate any
division into separate spccics in spite of attempts by others
to do so, However, since so much more oyster spat was
brought to our West Coast from Japan than from our own
eastern shores, it is quite possible that Alaska populations
are descendants of either Japanese or American stock,
or an admixture of both.

The author hopes to continue investigations into the
origin, distribution, and population dynamics of Mya
arenaria in Alaska and make further taxonomic studies.
Valid evidence that it is a relict of past times is also being
sought but to date no such evidence has been encountered.

{Epvrror’s Note: A very short time before going to press with the
current issue we learned of the sudden death of Mr. Gross. We
extend our sincere condolences to his relatives and many friends.}

JX, Wil, 1).

Pacific Division

THE TwentieTH ANNUAL MEETING of the American
Malacological Union — Pacific Division was held from
June 28 to July 1, 1967, at Asilomar Conference Grounds,
Pacific Grove, California. The Conchological Club of
Southern California acted as host for the meeting.

The Pacific Division Award of Honor was presented to
Mr. Attyn G. Smiru, California Academy of Sciences,
San Francisco. The presentation was made by Dr. S. S.
Berry. There was also a Chairman’s Award to Rose and
Joun Q. Burcu. Dr. Anne Hurst from the University of
Reading, England, was the principal speaker at the ban-
quet, which was held this year at the Outrigger on
Monterey’s well-known Cannery Row.

Officers for the coming year were elected. They are:
Chairman: Mrs. Fay Wolfson (San Diego)

First Vice Chairman: Dr. Bruce Campbell (Lynwood)

Second Vice Chairman: Dr. Dwight W. Taylor, Tempe
(Arizona )

Secretary: Mrs. Barbara Good, (San Diego)

Treasurer: Mrs. Helen DuShane (Whittier)

Mentor-Parliamentarian: Dr. Rudolf Stohler (Berkeley)

Page 204

CauiFoRNIA MALACOZOOLOGICAL SocIETy, Inc.
announces:

Backnumbers of

THE VELIGER

and other publications

Volumes 1 through 7: out of print —
Volume 8: $14.- Volume 9: $22.-
Supplement to Volume 3: $3.-* plus $-.35 mailing charge

[Opisthobranch Mollusks of California
by Prof. Ernst Marcus]

Supplement to Volume 6: $4.-* plus $-.35 mailing charge

[Biology of Tegula funebralis (A. ApAMs), edited by
Abbott, Blinks, Phillips and Stohler]

Supplement to Volume 7: $2.-* plus $-.35 mailing charge

[Glossary of A Thousand-and-One Terms used in
Conchology, compiled by Win1FreD H. ARNOLD]

Items marked with * are subject to sales tax in the State
of California; residents of that State please add the
appropriate amount to their remittances.

Prices subject to change without notice.

Send orders with remittance to:

Mrs. Jean M. Cate, Manager, 12719 San Vicente Boule-

vard, Los Angeles, Calif. 90049. Please, make checks
payable to C. M. S., Ine.

Shipments of material ordered are generally made within
two weeks after receipt of remittance.

Backnumbers of the current volume will be mailed to new
subscribers, as well as to those who renew late, on the
first working day of the month following receipt of the
remittance. The same policy applies to new members.

Subscription to Volume 10: $12.- domestic; $12.60 in
Canada, Mexico, Central and South America;
$ 12.90 all other foreign countries.

Affiliate Membership in the C.M.S., Inc. is $6.- for
the fiscal year July 1, 1967 to June 30, 1968. Postage for
members living in Canada, Mexico, Central and South
America 60 cents, for members living in any other foreign
country 90 cents additional. Membership open to indi-
viduals only - no institutional memberships. Please,
send for membership application forms to the Manager
or the Editor.

THE VELIGER

Vol. 10; No. 2

At a Regular Membership Meeting of the CatirorNiA
MatacozootocicaL Society, Inc. the following policies
were adopted by unanimous vote:

There will be an initiation fee of $2.- (in addition to
the annual dues) for persons joining the Society on or
after January 1, 1967.

Members receive The Veliger free of further charges and
are entitled to purchase one copy of any supplement pub-
lished during the current membership year at a special
discount (to be determined for each supplement).

Membership renewals are due on or before April 15
each year. If renewal payments are made after April 15
but before March 15 of the following year, there will be
a re-instatement fee of $1.-. Members whose dues pay-
ments (including the re-instatement fee) have not been
received by the latter date, will be dropped from the rolls
of the Society. They may rejoin by paying a new initiation
fee. The volume(s) published during the time a member
was in arrears may be purchased, if still available, at the
regular full volume price plus applicable handling charges.

THE CALIFORNIA MALACOZOOLOGICAL SOCIETY, Inc.

is a non-profit educational corporation (Articles of In-
corporation No. 463389 were filed January 6, 1964 in
the office of the Secretary of State). The Society publishes
a scientific quarterly, the VELIGER. Donations to the
Socicty are used to pay a part of the production costs and
thus to keep the subscription rate at a minimum. Donors
may designate the Fund to which their contribution is
to be credited: Operating Fund (available for current
production) ; Savings Fund (available only for specified
purposes, such as publication of especially long and signi-
ficant papers); Endowment Fund (the income from
which is available. The principal is irrevocably dedicated
to scientific and educational purposes). Unassigned dona-
tions will be used according to greatest need.

Contributions to the C. M.S., Inc. are deductible by *
donors as provided in section 170 of the Internal Revenue
Code (for Federal income tax purposes). Bequests, lega-
cies, gifts, devices are deductible for Federal estate and
gift tax purposes under section 2055, 2106, and 2522 of
the Code. The Treasurer of the C. M. S., Inc. will issue
suitable receipts which may be used by Donors to substan-
tiate their respective tax deductions.

Vol. 10; No. 2

THE VELIGER

Page 205

BOOKS, PERIODICALS, PAMPHLETS

Late Pleistocene Marine Paleoecology and
Zoogeography in Central California

by Warren O. Appicorr. Geological Survey Professional
Paper 523-C, 21 pp.; 4 pls.; 6 figs. in text. U. S. Govern-
ment Printing Office, Washington, D. C., 1966, price 35cts.

In this excellent paper which contains a wealth of
detailed data, a comparison is made of Pleistocene faunae
with those of Recent time in an area extending roughly
from Canada to Lower California. Conclusions are drawn
as to depth of water and other ecological factors obtaining
during the Late Pleistocene.

RS

Mollusca [from below 4000 meters in the Atlantic]

by N. H. OpHNER- Reports of the Swedish Deep-Sea
Expedition 1947 - 1948. Edited by Hans Pettersson, Vol.
2, Zoology, Fascic. 4, Zoology No. 22, pp. 367 - 400, pls.
1 and 2, figs. 1 - 12 in text, November, 1960. (Goteborgs
Kungl. Vetenskaps- och Vitterhets-Samhialle).

This report contains a discussion of 11 species, 8 pelecy-
pods and 3 gastropods; 7 species are described as new.
These mollusks were dredged in the Atlantic Ocean at
depths varying from 4430to 5987m from 4 areas: the

Romanche Deep; off the coast of Venezuela; north of the
West Indies; and between the Azores and Ireland. The
known species, fossil and Recent, of Vesicomya, Poromya,
and Pleurotomella, are listed.

Species described as new are: Portlandia (Yoldiella)
subcircularis, Poromya romanchensis, Cuspidaria nybelini,
Kellyella goest, Scissurella josephinae, Benthonella? kul-
lenbergi, and Pleurotomella heterogramma.

LGH

Brachiopoda and Mollusca

by J. Hop—E MacpuHerson. Memoirs of the National Mu-
seum No. 27. pp. 199 - 384; 2 charts; 5 plates; 3 figures
in text; 3 tables. 2 November 1966.

This is, in essence, a report on the Port Phillip Survey
of 1957 - 1963. Although the Brachiopods are mentioned
in the title of the paper, only one species is mentioned and
a general remark is made to the effect that brachiopods
were conspicuously absent from Port Phillip. The Amphi-
neura are represented with about 20 species, the gastropods
with about 120 species, the Bivalvia with over 40 species,
the cephalopods with 12 species. However, this is not a
simple enumeration of species found; Miss Macpherson
has provided amplifications of the original descriptions
where this seemed to be desirable. Moreover, a list of
species reported from Port Phillip but not obtained during
the survey of 1957-1963 is also given.

RS

Page 206 —

THE VELIGER

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 Society, Inc. Editor: Rudolf Stohler. Owner:
California Malacozoological Soicety, Inc., a non-profit, educational corporation.
Bondholders, mortgagees, and security holders: none.

(signed) R. Srouier, Editor.

Vol. 10; No. 2

ee

VELIGER

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME 10 January 1, 1968 NuMBER 3
ConTENTS

Zonation in Marine Gastropods of Costa Rica and Species Diversity.
CER AUDI pe NCU Si tarmac ear Men mes ot gourd ts ote atic 6: as fe we e207

Western Australian Cowries, —- A Second, Revised, and Expanded Report.
(Plates 21 to 34; 5Maps)
CRAWRORDE NM GALE amen tierce ernie =) oo ie sed fcute ed. way om es iets e. sie OTD

A Checklist of Intertidal Mollusks for Bahia Willard and the Southwestern Portion
of Bahia San Luis Gonzaga, State of Baja California, Mexico.
(Plate 35; 1 Map)

EIERENE DU OHANE:& | GATEIG. SPHON] 0.) 2) sp si «| o)o4 elis4.e ss ee 6 233
Mating Behavior in Mitra idae MELVvILL, 1893. (8 Text figures)

[BASIMG (CAG8R og) G26 eS TiQU ONL Gh SOAS) NO MCN Ones ee acne Rerun ae Hiee rls Gnalesae by)
Structure of the Bivalve Rectum. - I. Morphology

(Plates 36 to 40; 1 Text figure)
AMTOMAS Ca EGLAV& MICHAEL J. GREENBERG).2) = |) ¢ 302 = 6 2s ss 253

The Generic Classification of Cowries.
ERANZBEATERED, SCEMIEDER@ ey loe ler ia WE il. «pars aeeia dee icy Jie ciuh uiFebgtel isle aet-tor le, cap BOA,

Semele martinii (REEVE, 1853) of Southern Brazil and Uruguay.
IMD@UIEE, Ay ROU NE ENING G6. i big. o 6 onnon 0 lol on tonug Gl uommonei7/.)

Observations on Hipponix conicus (SCHUMACHER, 1817).
(Plate 41; 3 Text figures)

Wartrens OrnvER)| GERNOHORSKY =) sy te vi u-r sins ees wien sce We =) 275
[Continued on Inside Front Cover]
.
om +n Distributed free to Members of the California Malacozoological Society Inc.
2 w Subscriptions (by Volume only) payable in advance to Calif. Malacozoological Soc., Inc.
ge oe Volume 10: $12.- Domestic; $12.60 in the Americas; $12.90 all other Foreign Countries.
= ~ 4 Single copies this issue $9.00. Postage extra.
oO 4 Send subscriptions to Mrs. Jean M. Cate, Manager, 12719 San Vicente Boulevard,
i. m

? Los Angeles, California 90049. Address all other correspondence to Dr. R. STOHLER, Editor,
W, Department of Zoology, University of California, Berkeley, California 94720.
a Second Class Postage paid at Berkeley, California

Contents — Continued

Notes on the Range Extension of the Boring Clam Penitella conrad
(VALENCIENNES) and its Occurrence in the Shell of the California Mussel,
(Plate 42)

STEVEN EB. MEREDITH 20 een is ce ec er eevee es . 281

A Radula Muscle Preparation from the Gastropod, Kellettia kelletu, for Biochemical
Assays. (1 Text figure)

Howarp M. FEpER & REUBEN LASKER bo 6 BSR

A Remarkable New Cancellariid from the Philippines, with Comments on Related
Taxa. (Plate 43; 2 Text figures)
OAT a) JRO) A Anl) ROAM RMEECEIS Ss 66 5 0 0 6 0 coo o Hae

A Ctenostomatous Ectoproct Epizoic on the Chiton Ischnochiton mertensit.
(2 Text figures)

EUGENES, EREERMAN?) -c.006) 30 vee elie a ee fel coherent

Saco 6- 2E0
NOTES. & NEWS) 55) ee 50 Si ob ce seco oo ee
New Record of Conus ebracus in Costa Rica. JosEpH R. Housrick
Spawning Notes, I. — Hexaplex erythrostoma. (1 Text figure)

Fay Henry WOLFSON
COAN Molluscan Collection at University of California, Davis.
James W. VALENTINE

BOOKS, PERIODICALS & PAMPHLETS . ......+. +. + + + + + 294

Note: The various taxa above species are indicated by the use of different type styles as
‘shown by the following examples:

ORDER, Suborder, DIVISION, Subdivision, SECTION,
SUPERFAMILY, Famity, Subfamily, Genus, (Subgenus).
New Taxa

Vol. 10; No. 3

THE VELIGER

Page 207

Zonation in Marine Gastropods of Costa Rica

and Species Diversity

BY

GERALD J. BAKUS

Allan Hancock Foundation, University of Southern California, Los Angeles, California 90007

Durinc JuLy anp Aucust of 1962 I participated in the
second Advanced Science Seminar in Tropical Ecology
offered at the University of Costa Rica, San Jose, Costa
Rica by the University of Southern California through a
grant (G-21935) from the United States National Science
Foundation. One of the requirements of the program was
to initiate a limited research project. A comparative
study was made of littoral (intertidal) zonation of gastro-
pod mollusks on the Pacific and Atlantic coasts of Costa
Rica. The littoral zone was divided into thirds for con-
venience, representing high, middle and low portions.
Pacific coast zonation on rocky shores was studied at
three localities: Punta Caldera near Mata de Limon,
Puerto Quepos and Punta Catedral, abbreviated below as
ML, PQ and PG, respectively. These regions are charac-
terized by having a mixed tide with a daily flux of 2.6m
to 3.3m (8 to 10 feet). Mata de Limon (within the Golfo
de Nicoya) has a coastline with mangroves, considcrable
mud by runoff from a local river and turbid water. The
site of collection and observation (Punta Caldera) was
reached by boat and located about 1.5km southwest of
Caldera. The shoreline is rocky and relatively barren.
Brachyuran and anomuran crabs are abundant as is one
of three species of barnacles living there. The bivalve
mollusk Brachidontes puntarenensis is especially numerous.
Benthic algae are low in diversity and standing crop. Puerto
Quepos and especially Punta Catedral have exposed rocky
shorelines. The latter region is reminiscent of rugged por-
tions of the coast of central and northern California but
the epibiota is sparse excepting brachyuran and anomuran
crabs and gastropod mollusks. Long stretches of sandy
beaches are found nearby and olives (Olivella) are abun-
dant there. A composite portrayal of zonation among the
more conspicuous gastropods is presented as follows: The
supralittoral zone is occupied by Littorina conspersa at
Mata de Limén, found as high as 10m above the 0-tide
level. No supralittoral littorines were observed at Puerto
Quepos or Punta Catedral. The high littoral zone contains
L. conspersa (ML, PQ, PC), L. aspera (PQ, PC), Nerita

scabricosta ornata (ML, PQ, PC), Purpura patula pansa
(PQ, PC), PR columellaris (PQ, PC), and Siphon-
aria gigas (PQ, PC). The mid-littoral zone is character-
ized by L. conspersa (ML, PQ, PC), L. aspera (PQ, PC),
N.s.ornata (ML, PQ, PC), S. gigas (PQ, PC), and
Fissurella virescens (PQ, PC) with Opeatostoma pseudo-
don (PQ, PC), Tegula pellisserpentis (PQ, PC), Thais
melones (PQ, PC) and Acanthina brevidentata (ML, PQ,
PC) in the lowermost levels. The low littoral contains
O. pseudodon (PQ, PC), Tegula pellisserpentis (PQ, PC),
Thais melones (PQ, PC), A. brevidentata (ML, PQ, PC),
F. virescens (PQ, PC), Neosimnia sp. (on a pink gorgonian
near PC), Conus nux (PC), Natica chemnitzii (ML),
Polinices otis (ML) and with Thais biserialis (ML) in
the lowermost levels. Atlantic coast zonation on old up-
lifted limestone was studied at three localities: Portete,
Puerto Limon, and Isla Uvita, abbreviated below as P,
L, and U, respectively. These coastal areas are character-
ized by having a diurnal tide with a daily flux of about 1m
(3 fect). Portete, located about 5km north of Puerto
Limon, has a small bay which is rather rich in benthic
algae and flowering plants. Caulerpa, Thalassia and the
sea urchin Diadema are common in shallow waters. Live
corals grow in pockets near the mouth of the bay. Puerto
Limon has an uplifted shelf of old exposed limestone
pavement pocked by numerous shallow depressions con-
taining abundant bivalved mollusks (Isognomon radiatus
and Isognomon cf. I. bicolor). This is especially charac-
teristic of the shoreline north of the town. Benthic algae
are diverse and relatively abundant. Numerous echinoids,
ophiuroids, sca anemones, brachyuran and anomuran
crabs, gastropod mollusks and several species of live corals
occur in this region. Isla Uvita is located about 1 km off-
shore from the town of Puerto Limén. The windward side
is rocky and precipitous; the crab Grapsus grapsus is a
rapid moving and common representative there. The
leeward side contains a fringing reef with a sandy lagoon
between the island and the reef. Gastropod zonation,
discussed below, was studied on the leeward side.

Page 208

Thousands of small black snails (Cerithtum variabile)
are found on the sand in the Thalassia beds within the
lagoon. A composite portrayal of zonation among the more
conspicuous gastropods is presented as follows: The sup-
ralittoral zone contains Tectarius muricatus (P, L) which
is found as high as 2m above the 0-tide level. Littorina
ziczac (P, L) occurs in the lowermost levels of this zone.
The high littoral zone contains L. ziczac (P, L) and L.
nebulosa (U). The mid-littoral zone is characterized by
Purpura patula (U), Thais haemastoma floridana (P),
Littorina meleagris (L), Conus mus (L, U), Nerita ful-
gurans (P), N. tessellata (P), and with Cittarium pica
(L, U) in the lower levels. The low littoral zone contains
Cittarium pica (L, U), N. fulgurans (P), Neritina virgi-
nea (P) and Leucozonia nassa (L). A list of all mollusks
collected alive as well as shells is presented below in the
appendix. Specimens taken are now located in the Los
Angeles County Museum of Natural History under the
care of Dr. James H. McLean, Curator of Invertebrate
Zoology.

DISCUSSION

The supralittoral and high littoral zones on both Pacific
and Atlantic coasts are dominated by Littorinidae. Pur-
pura, Nerita and Siphonaria seemed to be missing from
these zones on the Atlantic coast. It is common to find
littorines predominating in the high littoral and supra-
littoral zones in part because of their resistance to desic-
cation. Ros—EWATER (1963) reviewed the literature on
desiccation in littorines. Tectarius muricatus lived for 18
months in a box. Littorina ziczac lived for 7 months with
only infrequent wetting. Both Pacific and Atlantic coasts
of Costa Rica samples contain supralittoral and littoral
gastropods in which roughly half are carnivores and half
herbivores. Visual observations indicated that Pacific coast
stations supported about twice the standing crop of gastro-
pods as that of the Atlantic coast but quantitative informa-
tion is needed for verification. If this is a true measure of
standing crop a paradox is encountered because the
diversity and standing crop of benthic algae are very low
along the Pacific coast; it is one of the poorest regions for
benthic algac in the world (Dr. E. Yale Dawson, personal
communication). Some of the reasons for differences in
benthic algae along the two coasts are discussed by Daw-
son (1962). However, primary production by benthic
algae must be reasonably high on the Pacific coast, despite
high turbidity in many areas, in order to account for the
apparently greater standing crop of gastropods. It may be
that benthic algae seem to be depauperate on the Pacific
Coast because of high intensity grazing by invertebrates

THE VELIGER

Vol. 10; No. 3

and perhaps fishes. This same phenomenon is character-
istic of many other tropical regions (Bakus, 1966a and in
press).

The pioneer taxonomic work on the mollusks of Pacific
Central America is that of Prrssry & Lowe (1932). Com-
prehensive information on Pacific Coast mollusks is found
in Keen (1958) and on Caribbean mollusks in WARMKE
& AppotT (1961). Recently, Parne (1966) studied briefly
the feeding relationships between two small carnivorous
gastropods, a mytilid (probably Brachidontes) and bar-
nacles near Mata di Limon, Costa Rica, and compared
them with similar observations in the northern Gulf of
California and at Mukkaw Bay, Washington. The work of
PatneE (1966) and observations by the present author
agree in that the local species diversity of rocky littoral
and supralittoral mollusks of the Pacific Coast of Costa
Rica is low. This leads to the interesting question of why
does local species diversity appear to be low in Costa Rica,
slightly greater in Washington and high in the northern
Gulf of California. Paine (1966) presented a logical
hypothesis supported in part by field data. He stated (op.
cit., p. 65): “Local species diversity is directly related to
the efficiency with which predators prevent the mono-
polization of the major environmental requisites of one
species.” Patne did not consider the importance of the
local substratum on the settling of larvae and spores, in
order that predators have something with which to “pre-
vent” monopolization. Gipson (1966), from studies on
foraminiferal diversity in Mississippi Sound, Gulf of
Mexico, concluded that regions of high diversity are asso-
ciated with stable physico-chemical environments. Both
Paine and Gigsson have contributed to ecological theory
but their viewpoints are restrictive. For example, if Grp-
son’s hypothesis were generally true, then the rocky and
sandstone intertidal region of the northern Gulf of Cali-
fornia (where water temperatures vary between 15°C
and 31°C during the year, where there is a relatively
long history of runoff from the Colorado River, where a
tidal flux of up to about 7m (22 feet) occurs daily, and
where the ambient air temperature may reach well over
38°C (100° F) in July-August and approach freezing
(0° C) in December-January) should have a relatively low
species diversity. Contrarily, the northern half of the Gulf
of California contains a high diversity of littoral animals
(GarTH, BELTRAN, & SavaGE, 1960; McLean, 1961;
Paine, 1966), and the Gulf of California may support the
greatest diversity of shallow water invertebrates in the
world (Parker, 1964). DUSHANE & PoorMAN (1967) re-
ported over 1000 species of mollusks from Guaymas, Sono-
ra, Mexico. Studies conducted by students and the present
author on the fauna of the region of Pefiasco, Sonora,
Mexico (northern Gulf of California) have indicated

Vol. 10; No. 3

that local species diversity of littoral invertebrates is
often high.

Gipson (1966) made the generalization that in the
distribution of marine benthon, species variety is lowest
near shore and greatest in deeper portions of the continen-
tal shelf. However, the diversity of scleractinian corals,
coral reef fishes and even temperate latitude crabs (Dr.
John Garth, personal communication) is usually greatest
in shallow waters. This probably holds true for many other
animal groups, especially those associated with sclerac-
tinian corals. More significantly, both Grsson and PAINE
overlooked the very important aspect of geological history.
For example, the northern Gulf of California has a unique
history. It has received organisms from the Pacific Ocean
by way of openings in the present Baja Peninsula (see
DurHaAM «& ALLISON, pp. 47-91 in GarTH, BELTRAN &
SavacE, 1960), it has received other organisms from the
Caribbean (see GarTH, BELTRAN & SAVAGE, 1960 and
Bakus, 1966b: 423), and several migrations of surface
water isotherms have occurred along the coast of southern
California and Baja California (Huss, 1960, 1961;
Banpy, 1967). The northern Gulf of California has a
complex biota consisting of elements from the cold tem-
perate, warm temperate, subtropical and tropical regions.
Moreover, it has a moderately high degree of endemism
(GarTH, BELTRAN & SAvaGE, 1960) and curious summer
and winter algal communities (DAwson, 1944: 200).
Much of the Pacific coast of Costa Rica has considerably
less diversity. The rocky littoral zone contains tropical
forms that must resist high air temperatures and desicca-
tion, excessive turbidity and frequent exposure to fresh-
water during part of the year, and probably periodic
effects of upwelling. On the other hand, it is likely that
the effects of predaceous, grazing and browsing fishes
on the littoral biota are more pronounced in Costa Rica
than in the northern Gulf of California (Bakus, 1966a).

In conclusion, more information, particularly of a
quantitative nature, is needed to better understand inter-
tidal zonation in Central America. Hypotheses that deal
with species diversity in the marine environment, should
be considered only as parts of an enormously complex
group of variables that change with time.

ACKNOWLEDGMENT

I would like to thank the following persons: Jay M.
Savage for introducing me to tropical biology; the late
E. Yale Dawson for introducing me to benthic marine
algae and for his companionship on field trips throughout
Costa Rica; W. Jake Houk for a collection of shells from
Mata de Limon, Costa Rica; James H. McLean for veri-
fications, corrections and additions on my identifications

THE VELIGER

Page 209

of marine mollusks taken in Costa Rica; Manuel M.
Murillo for a translation of a summary into Spanish; and
James H. McLean, Los Angeles County Museum of
Natural History, Jay M. Savage, Kristian Fauchald,
Manuel M. Murillo, Amada Reimer, Department of
Biological Sciences, University of Southern California,
and Orville L. Bandy, Department of Geology, University
of Southern California, for suggestions and criticisms.

APPENDIX

List of mollusks collected in Costa Rica, all of which are
on deposit in the Los Angeles County Museum of
Natural History

A. Pacific Coast:

1. Punta Caldera, near Caldera, Costa Rica: live mollusks,
littoral and supralittoral, collected by G. Bakus,
20 July 1962.

Acanthina brevidentata (Woop, 1828)

Anachis sp.

Anachis sp.

Brachidontes puntarenensis (PirsBry & Lowe, 1932)
Littorina conspersa Pui.iprt, 1847 2
Natica chemnitzii PFEIFFER, 1840

Nerita scabricosta ornata SOwERBY, 1823

Olivella zanoeta (Ductos, 1835)

Polinices otis (BRODERIP & SOWERBY, 1829)

Thais biserialis (BLAINVILLE, 1832)

Thais triangularis (BLAINVILLE, 1832)

Tricolia mazatlanica (STRONG, 1928)

ee aE STREP ODN NE

2. Mata de Limon, Costa Rica: shells cast up on beach, all
but one inhabited by hermit crabs, all but one
collected by Jake Houk, 21 July 1962.

Acanthina brevidentata (Woop, 1828)

Anachis fluctuata (Sowersy, 1832)

Cantharus ringens (REEVE, 1846)

Cantharus vibex (BropErip, 1833)

Cassis centiquadrata (VALENCIENNES, 1832)

Cerithidea pulchra (C. B. Apams, 1852)

Cerithidea valida (C. B. ApaMs, 1852)

Cymatium gibbosum (Bropverip, 1833)

Cymatium vestitum (Hinps, 1844)

Pitar lupanaria (Lesson, 1830), collected by G. Bakus,
20 July 1962

Rhinocoryne humboldti (VALENCIENNES, 1832)

Tegula byroniana (Woop, 1828)

Thais triangularis (BLAINVILLE, 1832)

Turritella sp. (juvenile)

Ree POOF NOK oO

[oe le Dos

Cypraea spurca acicularis GMELIN, 1791

Page 210 THE VELIGER Vol. 10; No. 3
3. Puerto Quepos, Costa Rica: live mollusks, littoral and Littorina nebulosa (Lamarck, 1822) 11
supralittoral, collected by G. Bakus, 20 July 1962. Littorina ziczac (GMELIN, 1791) 1
; ; Nerita fulgurans GMEtin, 1791 3
Acanthina brevidentata (Woop, 1828) a Nerita tessellata GMEuin, 1791 1
tee aspera PuILippl, ne F Fs Neritina virginea (LinNagus, 1758) 15
See Aas BL fae 5d 1893 4 Tectarius muricatus (LinNaAEus, 1758) 3
erita scabricosta ornata SOWERBY . .
, Thais h t

Opeatostoma pseudodon (Burrow, 1815) 2 MABE as foridana(CONnA ee ;
Planaxis planicostatus Sowersy, 1825 1 8. Portete, Costa Rica: shells cast up on beach, collected

Purpura columellaris (LaMaRcK, 1822) 2 by G. Bakus, 4 August 1962.

Purpura patula pansa Goutp, 1853 (?1852) 2 (

Siphonaria gigas SowERByY, 1825 I Charonia variegata (LAMaRCK, 1816) 1
Tegula pellisserpentis (Woop, 1828) 3 Cittarium pica (LinNaEuS, 1758) 1
Thais melones (Ductos, 1832) 5 Columbella mercatoria (Linnagus, 1758) 1
1
1

4. Puerto Quepos, Costa Rica: shells cast up on beach,
collected by G. Bakus, 20 July 1962.

Opeatostoma pseudodon (Burrow, 1815) 1
Purpura patula pansa Goutp, 1853 (?1852) 1
Pyrene labiosa (SowERBY, 1822) 2
Siphonaria gigas SowERBY, 1825 3
Tegula panamensis (PHt.ipPt, 1849) 1
Tegula pellisserpentis (Woop, 1828) 1

5. Near Punta Catedral (14km south), Costa Rica, live
mollusks, littoral and shallow sublittoral, collec-
ted by G. Bakus, 21 July 1962.

Conus nux Broperip, 1833 1
Fissurella virescens SowERBY, 1835 1
Neosimnia sp. (on pink gorgonian) 1

6. Near Punta Catedral (14km south), Costa Rica: shells
cast up on beach, collected by G. Bakus, 21 July
1962.

Conus purpurascens SowERBY, 1833

Cypraea arabicula LAMaRcK, 1811

Cypraea cervinetta KienrEr, 1843

Cypraea roberts: Hwwatco, 1906

Fissurella virescens SOWERBY, 1835

Oliva spicata (R6p1NG, 1798)

Strombus peruvianus Swainson, 1823

Trigoniocardia guanacastensis (HERTLEIN & STRONG,
1947) valvesonly 3

RSP One OD >

B. Atlantic Coast:

7. Portete, Costa Rica: live mollusks, littoral and supra-
littoral, collected by G. Bakus, 4 August 1962.

Anadara ovalis (BrucutirRE, 1789) 1
Isognomon radiatus (ANTON, 1839)
Leucozonia nassa (GMELIN, 1791)

wo —

Strombus raninus GMELIN, 1791

9. Puerto Limén, Costa Rica: live mollusks, littoral and
supralittoral, collected by G. Bakus, 7 July 1962.

Cittaritum pica (LinNaEus, 1758)

Conus mus Hwass, 1792

Isognomon cf. I. bicolor (C. B. ApaMs, 1845)
Isognomon radiatus (ANTON, 1839)
Leucozonia nassa (GMELIN, 1791)

Littorina meleagris (PotiEz & MicHaup, 1838)
Littorina ziczac (GmEuIN, 1791)

Modulus modulus (Linnaeus, 1758)

Tectarius muricatus (LINNAEUS, 1758)

Vasum muricatum (Born, 1778)

tl el tl ee On Oe

10. Puerto Limén, Costa Rica: shells cast up on beach,
collected by G. Bakus, 7 July 1962.

Arcopagia fausta (PULTENEY, 1799)
Cerithium eburneum Brucutkre, 1792
Columbella mercatoria (LINNAEUS, 1758)
Conus mus Hwass, 1792

Crassispira sp.

Isognomon sp.

Littorina ziczac (GMELIN, 1791)

Nerita fulgurans GMEun, 1791

Nerita tessellata GMELIN, 1791

Tectarius muricatus (LinNAEuS, 1758)

ee No

11. Puerto Limén, Costa Rica: mollusks in shallow sub-
littoral, collected by G. Bakus, 8 July 1962.

Vasum muricatum (Born, 1778) 2

12. Isla Uvita, Costa Rica: live mollusks, littoral and sup-
ralittoral, collected by G. Bakus, 8 July 1962.

Cerithium litteratum (Born, 1778)
Cerithium variabile C. B. ApaMs, 1852

ND bo

Vol. 10; No. 3

THE VELIGER

Page 211

Cittarium pica (LinNAEUuS, 1758)
Codakia orbicularis (LinNAEvs, 1758)
Conus mus Hwass, 1792

Leucozonia nassa (GMELIN, 1791)
Littorina nebulosa (LAMaRCK, 1822)
Purpura patula (Linnaeus, 1758)

eS ee ed

13. Isla Uvita, Costa Rica: shells cast up on beach, collec-
ted by G. Bakus, 8 July 1962.

Cittarium pica (LinNAEuS, 1758)
Conus mus Hwass, 1792

Cymatium pileare (LinNaEus, 1758)
Littorina nebulosa (Lamarck, 1822)

Ne ND PO

LITERATURE CITED

Baxus, GERALD J.
1966a. Some relationships of fishes to benthic organisms on
coral reefs. Nature 210 (5033) : 280 - 284

1966b. Marine poeciloscleridan sponges of the San Juan Archi-
pelago, Washington. Journ. Zool. London 149: 415 - 531

(in press) The feeding habits of fishes and primary production

at Eniwetok, Marshall Islands. Micronesica
Banpy, Orvitte L.

1967. Problems of Tertiary foraminiferal and radiolarian zona-
tion, circum-Pacific area. pp. 95-102 in: Hatat, K. (ed.)
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

Dawson, E. YALE

1944. The marine algae of the Gulf of California. Allan
Hancock Pacif. Exped. 8 (10): 1-452

1962. Additions to the marine flora of Costa Rica and Nica-
Tagua. Pacif. Nat. 3 (13): 375 - 395

DuSwane, HELEN & Roy PooRMAN

1967. A checklist of mollusks for Guaymas, Sonora, Mexico.
The Veliger 9 (4): 413-441; 1 map (1 April 1967)

GartH, J.S., E. Bettran « Jay M. Savace (Chairmen)
1960. Symposium: The biogeography of Baja California and
adjacent seas. Syst. Zool. 9 (2-4): 47-91, 93-231
Grsson, L. B.
1966. Some unifying characteristics of species diversity.
Contrib. Cushman Found. Foram. Res. 17 (4): 117 - 124
Hugss, Cari
1960. Quaternary paleoclimatology of the Pacific coast of North
America, pp. 105-112 in: O.E. Sette « J.D. Isaacs (eds.),

Part 2. Symposium on the changing Pacific Ocean in 1957 and
1958. Invest. Reprt. Calif. Co-op. Ocean Fish. 7: 13 - 217

1961. Some highlights from the natural radiocarbon datings of

La Jolla laboratory. Science, N. Y. 134: 1430
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)

McLgan, James H.

1961. Marine mollusks from Los Angeles Bay, Gulf of Califor-

nia. Trans. San Diego Soc. Nat. Hist. 12 (28): 449 - 476
Paine, Ropert T.

1966. Food web complexity and spccies diversity. Amer.

Natur. 100 (910): 65-75
PARKER, Rospert 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

Pirspry, Henry Aucustus & Herbert N. Lowe

1932. | West Mexican and Central American mollusks collected
by H. N. Lowe, 1929-31. — Proc. Acad. Nat. Sci. Philadelphia
84: 33 - 144; 7 figs.; plts. 1-17; 2 photogr. (21 May 1932)

RosEWATER, JOSEPH

1963. Resistance to desiccation in dormancy by Tectarius muri-

catus. The Nautilus 76 (3): 111
WarMKE, GERMAINE L. & RopErT 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.

Page 212

THE VELIGER _

Vol. 10; No. 3

Western Australian Cowries

A Second, Revised, and Expanded Report

BY

CRAWFORD N. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(Plates 21 to 34; 5 Maps)

INTRODUCTION

SEVERAL IMPORTANT FACTORS have made this sequel to my
1964 report advisable. First is the need to record subse-
quent information gathered in the area; new range
extensions are given, with special reference to the arca
including Port Hedland, Onslow, the outer shore of North
West Cape from Yardie Creek south to Dongara; more-
over, this report includes information gathered on a recent
collecting expedition to the Abrolhos Islands for the ex-
press purpose of learning the extent of cypraeid popula-
tions there and the relationships of the island fauna to
that of the mainland.

Secondly, the new generic arrangement of SctiItLDER
(1965, 1966), in which the species are listed in order
from the most primitive to the most specialized forms,
has been adopted; note should be taken, therefore, that
the taxonomic arrangement used here will supersede that
of Cate, 1964. A limited number of species has been
shifted into different genera, and subspecific recognition
_ has been accorded a West Australian form of Cypraca
leviathan SCHILDER & SCHILDER, 1937, Zoila friendit (Gray,
1831), and Palmadusta saulae (GasKo1n, 1843).

Thirdly, photographs have been added to illustrate all
the species of cowries presently known to inhabit West
Australian waters, from Cape Leeuwin in the south to
Port Darwin in the north. These illustrations will assist in
identification, and the revised text will offer a uniform
taxonomic approach to the species and their generic rela-
tionships.

The locality data as shown in Cate, 1964, will not be
repeated in the new text, unless they are incidental to the
listing of a newly recorded species in a previously reported
area. It has also been necessary to omit latitude and longi-
tude readings in most instances, since many of the remote
collecting stations do not appear on maps. In these cases

appropriate bearings in miles will be given in relation to
better known localities.

I was able to examine a great number of cowric species
in the South Australian Museum, Adelaide, and in the
West Australian Muscum, Perth, as well as in many
private West Australian collections. I found that most of
the shells fell within the size-range of the largest and
smallest shell dimensions recorded in Cate, 1964. It there-
fore seemed appropriate to take an average of their sizes
for usc in this overall study, as a median of shell dimension.
The five figures given in parentheses are, in order: length,
width, and height (in millimeters), number of teeth on
outer lip, and on the columellar lip. The numbers in
parentheses in the text under individual species discussions
indicate the locality references. The list of localities is
amended to include a South Australian locality where a
West Australian species has been recently discovered, or
overlaps in a natural range extension.

THE HOUTMAN ABROLHOS ISLANDS

The Houtman Abrolhos Islands Group, often referred to
as the “Houtman Rocks,” is an archipelago lying approxi-
mately 49 miles off the west coast of Australia in the
Indian Ocean, almost due west of Geraldton. Named for
its discoverer, Cornelis de Houtman, it consists of three
separate groups of islands lying approximately parallel
to the mainland, between the latitudes 28° 15’ and
29° 01’ South.

Little has heretofore been known about the mollusca
living in these remote island waters; our limited knowledge
of the fauna ts necessarily based upon the shells found
in fishermen’s crayfish pots, and upon visits to the islands
by Archer Whitworth of Geraldton in 1964 and a recent
trip of my own.

Vol. 10; No. 3

During October, 1966 (it was Spring in Australia) I
visited the Houtman Abrolhos Islands, personally collect-
ing at nearly every island of the southern Pelsart group
and of the middle Easter group. The cowrie populations
living in the intertidal zones were sampled and repre-
sentative lots of each available species were sent home
for study. In earlier years, Archer Whitworth, Geraldton,
and James “Jimmy” Seabrook of Perth have made
repeated trips to the islands, the former working the
littoral and adjacent shallow water, the latter, in the
motor vessel “Lancelin” of the Fisheries Department,
Perth, using baited crayfish pots and limited dredging in
the islands and other mainland offshore reef areas. It is
interesting to note that many of the common mainland
shore cowries are absent, as far as we have been able to
determine from the Abrolhos fauna. The ecological con-
ditions seem much the same in many instances, although
the islands have, generally speaking, much rougher water
and are more exposed to the storm force of the normal
southerly winds. The island waters are perhaps more
varied as to depth, condition of food, food source, and
substrate (there appears to be a noticeable scarcity of
algae; a large portion of the coral is living, thus being
devoid of marine plants) and in the distribution of sand,
mud, and dead coral. Shallow water -_ scarcely ever
more than 30 fathoms deep — and the combination of
many reef patterns in and out of the water, plus the end-
less broad, shallow-water, spongy, white coral sand flats,
and minimal coral and rock habitats, must surely affect
the distribution of many of the mainland mollusks.

Aside from the collectors already mentioned, the follow-
ing persons should also be listed: Alec Gilbertson, Max
Cramer, George Barker, Martin van der Oord, W. Hems-
ley, Joseph Burton, Terry Butterworth, O. Hewitt of
Geraldton; F I. “Sonny” Healy, Dongara; and Edward
Nickles of Mandurah. These and many others have con-
tributed in one way or another with specimens and infor-
mation, all of which have been used in this study. For the
most part these men are cray fishermen, and are to be
credited for having discovered and made available such
rare deep water species as Bernaya catei ScHILpER, 1963,
Zoila marginata (Gasxo1n, 1849), Z. venusta (SowERBY,
1846), the northern form of Z. friendii (Gray, 1831), Z.
rosselli Cotton, 1948, and the round northern form of
Cypraea reevei SowERBY, 1832. These rare species and
others are found feeding on the bait (often fish, dried
meat, but more often than not on dried split lamb’s heads
and other dried bone material) set in the crayfish pots.

The Wallabi Group to the north (see map) includes
North Island, East and West Wallabi Islands, with North
East Reef, Morning Reef, Noon Reef, and Evening Reef

THE VELIGER

Page 213

providing the most important collecting areas in this
section.

To the south of Middle Channel is the central (or
middle) Easter Group, made up of numerous small
islands, barely 8 to 10 feet above high water level, the
largest and most important of which is Rat Island. It
was at one of the fishermen’s huts here that we made our
headquarters during our stay at the Houtman Abrolhos
Islands. Of this group Little North and Sandy Islands are
the most recent of the many growing reefs to develop a
permanent footing above high water mark.

The southernmost link of this coral island chain is the
Pelsart Group (named after Francisco Pelsart, Captain of
the Dutch trading vessel Batavia, wrecked in these islands
on June 4, 1629), also referred to as the Southern Group,
with its main island bearing the same name. The Zeewyk
Channel separates this group from the Easter Group
complex. Made up of a combination of reefs, partially
submerged jetties, interior landlocked lagoons that rise
and fall with the tide, and atoll-like sand spits, it provides
excellent collecting stations, the most important of which
is at Wreck Point at the west end of mile-long Pelsart
Island. Otherwise, the best areas are Mangrove Island,
Hummock Island, Square Island, White Banks, and King
Reef, the latter becoming awash at high tide and accessible
only at low tide, and then for only a short time.

While Zoila friendu vercoi ScHILDER, 1930 is almost
surely a South Australian subspecies of Z. friendit (Gray,
1831), there are labels in Museum collections and else-
where indicating a questionable incidence for these shells
in West Australian waters. I am omitting them here as
not being a part of the western fauna, but it seems perti-
nent to remark on them briefly at this time (see Plate 23,
Figure 11, which illustrates this form). Cate coll. no.
Ci W706 (83.8 Sail B90 25 iO) == (Qa) 2 ito, © Sa00
(Js 49.2 Bode 2 O)) s= (BA) eo nm, C Se0il BS
44.0 34.3. 22 5) = (59a) are additional specimens
for comparative statistics.

Although geographically adjacent to Zoila friendii frien-
du of Western Australia, Z. friendit vercot may be con-
sidered as being an intra-population variant. WILSON &
Summers (1966) may be correct in their appraisal of
these shells. Among other proposals, they suggest placing
Z. f. vercoi into synonymy with Z. f. friendiit as being
“either clinical or subject to non-geographical, inter-loca-
lity variation.” This may possibly be true. One way or the
other, the south Australian form Z. f. vercoi does possess
apertural and teeth arrangement so characteristic of Z.
f. friendii, linking them very closely to one another.

I have examined 11 specimens of this subspecies, in
the South Australian Museum, the West Australian Mu-
seum, and in my own collection. The answer is always the

Page 214 THE VELIGER Vol. 10; No. 3

COLLIER ~W
COCKATOO I~, < BAY

BUCCANEER ARCHIPELAGO -:.:<
CAPE LEVEQUE —

PENDER BAY i Des,
BEAGLE BAY Y¢" ING SOUND

LACEPEDE Is/ &

X ~ QUONDONG

GANTHEAUME PT, (ENTRANCE PT.)$y. BROOME
ROEBUCK BAY

LA GRANGE BAY—% i» GRance
CAPE BOSSUT

DAMPIER ARCHIPELAGO
ENDERBY |.
ANGEL I. DIXON CErUee

e

“06 PORT SAMSON
MONTEBELLO IS cape © S

PRESTONY ~' ROEBOURNE

BARROW ")
I.

ARLIE |,
THEVENARD I>y
LONG |.
NORTH WEST CAPE ¢@
VLAMING HEAD ~f/ _Tey MOUTH GULF

3
YARDIE CREEK ey CEARMOUTH

NORWEGIAN BAYS; <4

PT. CLOATES -“\: a
MAUD'S LANDING

—

PT, QUOBBA
KOKS |..

i a «WESTERN | AUSTRALIA

DIRK

HARTOG |.
SOUTH PASSAGEG

STEEP PT. 5)
FALSE ENTRANCE

\

HOUTMAN *
ABROLHOS *

» GERALDTON
ISLANDS a

i /BASILC'S |. :
wy *DONGARA
{ FR
TURTLE DOVE* elie hots
SHOALS eo 0
GREEN HEADY: |,
JURIEN BAY —\

INDIAN CERVANTES ae

LANCELIN |)

OCEAN SORRENTO BEACHY. -
PERTH
ROTTNEST |. g PT EREMANTIE

‘
GARDEN 1./4%"'
WOODMAN'S PT. MANDURAH

¥, BINNINGUP
DUNSBOROUGH v BUNBURY

CAPE NATURALISTE !_ SeausseLTON
YALLINGUP

CANAL ROCKS~**% ELLENSBROOK
MARGARET RIVER .

ALBANY
CAPE LEEUWIN lpg: pei’ DENMARK

PT. DENTRECASTEAUX

Vol. 10; No. 3

THE VELIGER

Page 215

same; the mollusks from the southwest coast localities do
vary remarkably in morphological development. A student
could say there is a distinct difference in these shells, and
could maintain there is some point of geographical sepa-
ration between the southern and western populations in
the area between Cape Naturaliste and Cape Leeuwin,
though it is a narrow one and open to possible question.
The subspecies, Zola friendi vercoi, does retain many of
the shell characters of the western nominate species.
These would include the familiar apertural appearance,
the number of teeth (almost never more than 6 to 10 on
the abapical columella), and their arrangement. Particu-
larly unifying these two variants are the limited and
knobby, comparatively rudimentary teeth on the abapical
quarter of the columellar lip, adjacent to the fossula; the
remaining three fourths of the columella, adapically, is
normally smooth and without teeth. However, these shells
are distinctive enough, so that for purposes of distinguish-

at. WESTERN
& AUSTRALIA

‘tg NORTH ISLAND

WALL ABI
¥ . GROUP

_MID REEF é
CHAMPION \*.
BAY

\

3 A li PELSART GERALDTON.

GROUP

——S—)
5 MILES

ing the south southwest Australian shells I am tentatively
retaining the available name for this ‘end of cline’ group
as a South Australian subspecies (see Table 1 for compara-
tive statistics).

ACKNOWLEDGMENT

As is always the case in a work like this, many persons
have given generously of their help in countless tangible
and intangible ways; to them all I express my thanks. In
addition to those mentioned elsewhere in this paper for

their contributions of shells and information toward this
work, I wish to express my gratitude to Molly and Robert
Gedling, to Iris and Robert Thomas, both families (the
latter in the earlier days of this study) light keepers at
Vlaming Head, North West Cape. To Brian Kember,
Port Hedland, to Theodore Crake, Broome, to Mrs. D.
Clarke, Onslow, to Ned Harrold, Victoria Park, Perth, to
Dr. Helene Laws, South Australian Museum, Adelaide,
and to Dr. Barry Wilson, West Australian Museum, Perth,
for permission to inspect the collections in their care;
to Mrs. Emily Reid for the excellent maps; to Takeo
Susuki for assistance with photography; to Archer Whit-
worth, a dear friend of long standing living in Geraldton;
and to Jean Cate for assistance in many helpful ways.

LOCALITY INDEX

Due to the wild, uninhabited nature of much of the
present coastline of West Australia, distances given, and
many of the indicated localities, are only approximate.
Even so, they are reasonably correct as to landfall, and
truly represent the localities and ranges of the Cypraeidae
as we know them today.

Abrolhos Islands
Airlie Island, 16 miles NNE of Onslow
. Albany, South Australia
Albany, Frenchman’s Bay
Alec Gilbertson Island, Easter group, Abrolhos Islands
Alec Finlay Island (the nook), Abrolhos Islands
Anchor Island, 5 miles off-shore, 22 miles NW of Onslow
Angel Island, Dampier Archipelago
Anttila Island, Abrolhos Islands
Arch Whitworth Island, Abrolhos Islands
Ashburton Island, approximately 11 miles W of Onslow
Busselton, Geographe Bay
Barrow Island, 60 miles N of Onslow
Basilc’s Island, Abrolhos Islands
Bathurst Island, 60 miles NE of Darwin
Bathurst Point, Rottnest Island
Bernier Island, Shark Bay
Binningup, approximately 15 miles N of Bunbury

iS)

Oe Be ee eee
SY) Go NiO Sse © 9 Fo Si Go Gu cS Sw

21. Black Island, Abrolhos Islands

22. Bossut, southern point of La Grange Bay
23. Bowes River, 28 miles N of Geraldton
24. Broome, Roebuck Bay

25. Broome, Cocoanut Well

25a. Bunbury

Bunker Bay, 14 miles E of Cape Naturaliste

Capel, 17 miles S of Bunbury

Cape Bossut

Calgadup Brook, near Margaret River S of Cape Naturaliste
30a. Cape Jervis, South Australia

Cape Leeuwin

Cape Naturaliste

Oo NO PO PO
SOR

oo oO
nN —

Page 216

Cape Preston, Onslow
Cape Villaret, 30 miles N of Broome
. Carnac Island, just N of Garden Island
Carnarvon
Cervantes Island

37. Chabjuwardoo Bay, S of Point Cloates

38. Coburn

39. Cockburn Sound, S of Fremantle

40. Cockburn Sound, Pamelia Bank off Woodman’s Point
41. Cockatoo Island, Buccaneer Archipelago

42. Cottesloe Beach, Perth

43. Cowrie Creek, 35 miles S of Port Hedland

44. Cambridge Gulf, Wyndham, 250 miles S of Darwin
45. Darwin

46. Delambre Island, Dampier Archipelago

47. Direction Island, 7 miles NE of Onslow

48. Dirk Hartog Island, adjacent to Shark Bay

49. Dixon Island, off Port Samson

52. Dongara

s Man, ts
“ANSE =.

iN

seose AS

way
a
“Pp

> SS], 5

SESS =:
Es,

Ha

WALLABI
_ GROUP

“oo. EAST WALLABI |.

"each NOON REEF
MIDDLE CHANNEL

Dongara, Leander Reef

Dongara, Turtle Dove Shoal sce 156

54. Dorre Island, S of Bernicr Island, Shark Bay

57. Eagle Nest Island, Abrolhos Islands

58. Eagle Nest Island (south island), Abrolhos Islands
59. Enderby Island, Dampier Archipelago

THE VELIGER

63.
64.

62a.

Vol. 10; No. 3

. Esperance, South Australia ’

Exmouth Gulf
Fremantle

. Fremantle, off entrance to Swan River in 30 feet, on sponge

Far Eastern Islands (3), Abrolhos Islands

beazwerm
™g WHITWORTH
et ea

Rey Je

260M ¢

a

= Piaitsenrson
ie: ate
PASSAGE |.

ae Fi R s y
‘ UK

i
. yrs |

ae

Bite
x

%
pes
5 Rata! nee
fScut

ee © U

I.
UTANTTILA)
= HELSINKI

OR FINN I-

LITTLE 4 Ds a
aaa: Huey ae

HBERT a
a? EAGLES NEST I.

gfe ex 50. EAGLES NEST |.
“WHITE |.
woooee sen ¥” re LITTLE WHITE |.

“ BLACK I.

EASTER
GROUP

SOUTHERN SANDY I.

ZEEWYK | CHANNEL

False Entrance, 12 miles SSE of Steep Point, outer Denham
peninsula

Flat Reef, west end of West Wallabi Island, Abrolhos Islands

“Five Mile Beach,” 5 miles S of Vlaming Head, N. W. Cape

Garden Island

Geelvink Channel, Abrolhos Islands

Geographe Bay

Geraldton

Glenroy Station, 10 miles S of Onslow

Good Friday Bay, N of Rat Island, Abrolhos Islands

. Green Head, approximately 140 miles N of Perth
. Grindal Island, near Port Lincoln, South Australia

Gun Island, Abrolhos Islands
Helsinki (Finn) Island, Abrolhos Islands
Hopctown Beach, south coast of West Australia
Juricn Bay, N of Hill River, 75 miles S of Dongara
Juricn Bay, 3 miles off North Head

King Sound

King Sound, Disaster Bay

King Sound, Dugong Bay

Koks Island, off N tip of Bernier Island, Shark Bay
Lancelin Island, adjacent to Cervantes Island
Learmouth, 30 miles S of Vlaming Head in Exmouth Gulf
Ledge Point, 80 miles N of Perth (30 fathoms)
Leighton Beach, adjacent to Perth

Leo’s Island (Anttila), Abrolhos Islands

Little Italy Island, Abrolhos Islands

Little North Island, Abrolhos Islands

Little White Island, Abrolhos Islands

Locker Island, 30 miles SW of Onslow

Long Island, 32 miles W of Onslow

Ludlow, 8 miles N of Busselton

Vol. 10; No. 3

THE VELIGER

108.
109.
110.
111.
112.
113.
114.
115.
116.
72
117a.
118.
119.
120.
121.
12la.
124.
125.
126.
130.
131.
132.
133.
134.
135.

Mandurah, approximately 35 miles S of Fremantle

Maud’s Landing, approximately 135 miles N of Carnarvon
Middle Channel, Abrolhos Islands

Middle Island, Abrolhos Islands

Moore River, approximately 55 miles N of Perth

Ningaloo Woolshed, 200 miles N of Carnarvon

Nickol Bay, 25 miles S of Roebourne

North Island, Wallabi Group, Abrolhos Islands

. North Island, Wallabi Group, SW Bank

Norwegian Bay, Whaling Station N of Point Cloates
North West Cape
Quondong, 35 miles N of Broome

ZEEWYK CHANNEL

gi

See

PELSART
GROUP

‘m7, AM
y, WRECK,
z. PT. <* WRECK I.

Onslow

Onslow, 10 miles south of —

Passage Island, Abrolhos Islands

Pelsart Island, Abrolhos Islands

Pender Bay, approximately 150 miles N of Broome
Pender Bay, Bell Point

Point Locker, 50 miles S of Onslow

Point Murrat, Exmouth Gulf

Point Samson, 9 miles N of Roebourne

Point Cloates, 75 miles S of Viaming Head Light, N. W. Cape
Port Darwin

Port Denison Beach, 15 miles SW of Dongara
Port Hedland

Port Lincoln, South Australia

Price Point, 40 miles N of Broome

Quindalup, Geographe Bay

Quondong, 35 miles N of Broome

Quobba Point, 40 miles N of Carnarvon

Port Hedland, Finicane Island, $ mile off-shore
Rat Island, Abrolhos Islands

Rottnest Island, 12 miles NW of Fremantle
Rottnest Island, 5 miles NW of -, in 75 fathoms
Rottnest Island, Bathurst Point

Rottnest Island, Ricey Beach

Round Island, 2 miles E of Long Island

Page 217
136. Roebourne, Tonymia
137. Roebuck Bay, Middle Bank
140. Sandy Island (north), Abrolhos Islands
141. Sandy Island (south), Abrolhos Islands
142. Shark Bay
143. Shark Bay, False Entrance
144. Shark Bay, South Passage
145. Sorrento Beach (Sorrento Reef )
146. Sydney Liddon Island, Abrolhos Islands
147. Snag Island, 100 miles S of Geraldton
150. Tautibiddi Well, approximately 10 miles S of Vlaming Head,
N. W. Cape
151. Taylor’s Island, 20 miles S of Port Lincoln, South Australia
152. “The Flats,’ oceanward, S of Pelsart Island, in 35 fathoms,
Abrolhos Islands
153. Thevenard Island, 15 miles NW of Onslow
154. Thompson’s Bay, NW side of Rottnest Island
155. ‘Torloise Island, 15 miles W of Onslow
156. Turtle Dove ReefShoal, 37 miles WSW of Dongara
157. Twin Island, 9 miles ENE of Onslow
159. WVlaming Head, North West Cape
160. Vlaming Head, North West Cape, 4 miles SW of —
162. Wallabi Island (east), Abrolhos Islands
163. Wallabi Island( west), south side, Abrolhos Islands
164. Wedge Island, near Port Lincoln, South Australia
165. West Bank, 6 miles WNW of North Island, Abrolhos Islands
166. West Lewis Island, Dampier Archipelago
167. White Island, Abrolhos Islands
168. Windy Harbor, Cape D’Entrecasteaux, SW Australia
169. Woodcock Island, Abrolhos Islands
170. Wooded Island, Abrolhos Islands
171. Wreck Point, W-end of Pelsart Island, Abrolhos Islands
172. Woodman’s Point, 11 miles S of Fremantle
175. Yallingup, approximately 157 miles S of Fremantle
176. Yardie Creek, approximately 7 miles S of Vlaming Head,
N. W. Cape
177. Yardie Creek, 11 miles S of N. W. Cape
178. Yardie Creek, 20 miles S of N. W. Cape
180. Zeewyk Channel, Abrolhos Islands
INDEX or SPECIES
GUO comonomer POH OUOCODS creme Ay PRO
GIVGUST ALC ree na mee 229) cicercula . 219, 227
annulus... 219, 227| citrinicolor 219, 228
CHAGUAD — ccerocteneetcersreceemcrrces 218, 226] clandestina , 221, 230
ANGUS ee ene 2G) 220|\CONUPLOTt mem 2PA8)
ascllus _.... 220, 230] continens . 220, 229
DiCOLOT, | oN a tesn cman 229] contraria .. 220, 229
bizonata 221, 231] coxi ware 220, 229
blacsa : M2 20523 0|Cre kets 221, 230, 231
brevidentatay pasennseee Qo 232|Worbraia serene 221, 232
brunnescens . 218, 226] cylindrica .. 220, 230
cameroni © QQL, 232] AAMPLETENSIS eeeesrsessee 221, 231
COPULSETPCNEIS cccrrseenee ZI, 228) AECUPLENS oertrsrirsreseriririees 218, 225
COTMEONA etricscnaennranene: 219, 227| diversa . 220, 228
Catena ne eee, 213, 218, 221| dorsalis . 220, 229
caurica .. u. 220, 230| eglantina .. 218, 226
COIMICE nnn 219,228 Episema . 224

Page 218 THE VELIGER Vol. 10; No. 3

CVOSA cesssrsssssnisiaieis 2UQ, 220, 228|| PAVAAIIS scecrersceerirmrsrernsnes 218, 226 1. Bernaya (Bernaya) catet SCHILDER, 1963

errones 220, 229 perconfusa seri Sreeenten 218, 225 The Veliger 5 (4): 127

facifer .. 220, 229] piperita 220, 229

ROLLE he cere ee ee eee 221, 232| poraria 219, 228 P

EUING vovsusisnreenmnnnnnnnn 220, 230) pulicaria 220, 229 Zoila ee d

fimbriata . 221, 231] purissima 220, 228 Sih 2 Be : 414, nom. nud.)

LICE Sten ed ner eee 2210 RD 1] OF 711 Sameer ecmetarr 220, 229 ST wd ae Zool. France 9: 89; 1884

PHCNAI —evenssrsmninsene 212, 213, 214) quadrimaculata nmr 221, 231 ype species: Cypraca friendii Gray, 1831 <
sn 218, 222, 223, 224) TEEVEL wenn 213, 219, 226, 227 2. Zoila friendii friendii (Gray, 1831)

gedlingae .. . 219, 227| reticulum . 219, 228 Zool. Misc. 1: 35

globulus © 21Q, 227| THINOCETOS vrcrrmumnarercecnenes 22K padi =

Tacilis meinen eee 221, 231| rhomboides — erimnenunns 219, 227 3. Zoila fricndi jeaniana Cate, subspec. nov.

PAMMONA|AE erersrsrssesrsnesn 221, 231] rossellt_...... 213, 218, 225 J

helvola oo PUG, POLY! POLITE ccemsecerecceireca 219, 227 4. Zoila venusta (SowERBY, 1846)

hilda ve QZ2T, 2ZT) SQULAE erresrsesen 212, 221, 230, 231 Proc. Linn. Soc. London, prt. 1: 314

hirundo we 221, 232) siasiensis . 230, 231 . .

histrio 226, 228] SIMULANS rercrrstcrscrsieeniniresn 221, 231 2 ZaNe Hance Seer bee for 1848: 91
isabella vane 21Q, 227| smithz .. 220, 229 cae ee aaa ‘

jeaniana 218, 222, 223) SOTTCMEEMSIS arsccrsriesrnnmenniemeen 224 6. Zoila rosselli Cotton, 1948

kenyonae . 219, 228) sowerbyana 220, 230 Trans. Roy. Soc. So. Austral. 72 (1): 30; plt. 1

kieneri . 221, 231) staphylaca . 220, 229 ‘ Megs

labrolineata 219, 228] stolida . 221, 232 7. Zoila decipiens (E. A. SmirH, 1880)

leviathan 219, 227| subviridis a 220, 229 Proc. Zool. Soc. London for 1880: 482; plt. 48, fig. 8

limacina 220, 229| talpa . 218, 226

Listeria eee reece sees 220] teres 221, 231 Mauritia TRoscHEL, 1863
Lid LO Hatcheries 224. Das Gebi® der Schnecken 1: 205
lutea . 221, 231) thersites 222, 223 > Type Species: Cypraca mauritiana Linnaeus, 1758 <

lynx QNOW220|NtClCU pr atmmaertener a2 21 y2 Oi Systema Naturac, ed. 10: 721
AACOUS ONG ctccreccensarccoercrceomert20 226) ligris . 219, 226 ‘

marginata .... 213, 218, 224, 225] turdus 220, 228 (Arabica) JOUSSEAUME, 1884

mauritiana... vo 218] ursellus . 221, 232 : : Naturaliste 1884: 414
MEMIA reserves 218} vanelli een 218220 > Type species: Cypraca arabica Linnarus, 1758 <
MelOillin cetces cee henteere 220, 230) VENUSEA riers 213, 218, 223, 224 Systema Naturae, ed. 10: 718
AOUOS | scarommnncercremeceneoce 220, 228] VETCOL eremmnen 213, 214, 222, 223 BAG : 6

Fags SE | BiG, aex|| abradbiaallnr | BG, B® 8. Mauritia (Arabica) eglantina perconfusa

nucleus co PAD, PA)! WCAG cxcrccreacrecmsieise 219, 226 TREDALE, 1935 ,

NUGALA recen alot ate ARS 230 cla) hereunto 220, 229 Austral. Zoologist 8 (2): 108
ONYX eaten ea eens 2 20 Scene - 218, 226, 228 O), Mamata (Arabica) arabica brunnescens Cate, 1964
OVUT rrnnrnnnrninsennn 220, 229) w PROPANE cerca re 221, 232 The Veliger 7 (1): 24; plt. 5, figs. 3a, 3b
pallidilay eens: DP, AN AGHA crommomcenrnttosncrcnc 221, 231

SYSTEMATIC LIST

CyPRAEIDAE FLEMING, 1828
Hist. Brit. Anim., 330 (em.) (Edinburgh)

CyPRAEORBINAE SCHILDER, 1939
Arch. Molluskenk. 71: 165

Bernayini ScuiLper, 1927
Arch. Naturgesch. 91/A 10: 88

Bernaya JoUSSEAUME, 1884
(Naturaliste 1884: 414, nom. nud.)
Bull. Soc. Zool. France 9: 88; 1884

(Bernaya) JousSEAUME, 1884
> Type species: Cypraca media Destuayes, 1835 <
Descr. coq. foss. envir. Paris 2, 723, 95: 37 - 38

10. Mauritia (Arabica) histrio westralis (IREDALE, 1935)
Austral. Zoologist 8 (2): 108

Talparia TRoscHEL, 1863
Das Gebif& der Schnecken 1: 204
> Type Species: Cypraca talpa Linnazus, 1758 <

it, Talparia talpa talpa (Linnaeus, 1758)
Systema Naturae, ed. 10: 720

Cypraca Linnagvs, 1758
Systema Naturac, ed. 10: 718

(cm.) Montrort, P. Denys bE, 1810
Conchyl. Syst. 2: 630

> Type Species: Cypraca tigris Linnazus, 1758 <
(Cypraea) LINNAEUS, 1758

12. Cypraca (Cypraca) tigris pardalis Suaw, 1795
Vivar. Natur. Misc. 6: plt. 193

Vol. 10; No. 3

THE VELIGER

Page 219

(Lyncina) TroscHEL, 1863
Das Gebi® der Schnecken 1: 205
> Type Species: Cypraea lynx LinnAEus, 1758 <
Systema Naturae, ed. 10: 721

13. Cypraea (Lyncina) argus argus LINNAEUS, 1758
Systema Naturae, ed. 10: 719

14. Cypraea (Lyncina) lynx vanelli Linnarus, 1758
Systema Naturae, ed. 10: 720

15. Cypraea (Lyncina) vitellus vitellus LINNAEUS, 1758
Systema Naturae, ed. 10: 721

16. Cypraea (Lyncina) reevet SowERBy, 1832

Conch. Illustr. (London) fig. 52

7, pee (Lyncina) carneola carneola LINNAEUS, 1758
Systema Naturae, ed. 10: 719

18. Cypraea (Lyncina) leviathan gedlingae Cate,
subspec. nov.

Luria JoussEAUME, 1884
Bull. Soc. Zool. France 9: 92
(Naturaliste 1884: 414, nom. nud.)
> Type Species: Cypraca lurida Linnazus, 1758 <
Systema Naturae, ed. 10: 720

(Basilitrona) TrREDALE, 1930
Mem. Queensld. Mus. 10 (1): 83

> Type Species: Cypraca isabella Linnazus, 1758 <
Systema Naturae, ed. 10: 722

IG), Luria (Basilitrona) isabella rumphiu
ScHILDER & SCHILDER, 1938
Proc. Malacol. Soc. London 23 (3 - 4): 177

NarmnaeE ScHILDER, 1932
Foss. Cat. 1: Animalia, pars 55, Cypraeacea. 149

(Pustulariini ScHILpER, 1932)
Foss. Cat. 1: Animalia, pars 55, Cypraeacea. 149

Pustularia Swainson, 1840
Larpner’s Encycl., p. 324

(Pustularia) Swainson, 1840
Larpner’s Encycl., p. 324
> Type Species: Cypraea cicercula LinnaEus, 1758 <
Systema Naturae, ed. 10: 725

20. Pustularia (Pustularia) cicercula cicercula

(Linnaeus, 1758)
Systema Naturae, ed. 10: 725

Zl. Pustularia (Pustularia) globulus globulus
(Linnaeus, 1758)
Systema Naturae, ed. 10: 725

Nariini ScHILpER, 1932
Foss. Cat. 1: Animalia, pars 55, Cypraeacea: 159

Monetaria TroscHEL, 1863
Das Gebi®§ der Schnecken 1: 205
> Type Species: Cypraea moneta Linnagus, 1758 <
Systema Naturae, ed. 10: 723

(Ornamentaria) SCHILDER &
SCHILDER, 1936
Proc. Zool. Soc. London 1936: 1120

> Type Species: Cypraea annulus LinnaEus, 1758 <
Systema Naturae, ed. 10: 723

22. Monetaria (Ornamentaria) annulus annulus
(LinnagEus, 1758)
Systema Naturae, ed. 10: 723

(Monetaria) TroscHet, 1863
Das Gebif& der Schnecken 1: 205

23. Monetaria (Monetaria) moneta rhomboides
SCHILDER & SCHILDER, 1933
Zool. Meded. Leiden 16: 163

Erosaria TROSCHEL, 1863
Das Gebif& der Schnecken 1: 205
> Type Species: Cypraea erosa LinNaEus, 1758 <
Systema Naturae, ed. 10: 723

(Ravitrona) IrEDALE, 1930
Mem. Queensld. Mus. 10 (1): 82
> Type Species: Cypraca caputserpentis LINNAEUS, 1758 <
Systema Naturae, ed. 10: 720

24. Erosaria (Ravitrona) labrolineata labrolineata
(Gaskorn, 1849)
Proc. Zool]. Soc. London for 1848: 97

25. Erosaria (Ravitrona) cernica viridicolor (Cate, 1962)
The Veliger 4 (4): 175; plt. 40, figs. 1-9

26. Erosaria (Ravitrona) helvola citrinicolor
TREDALE, 1935
Austral. Zoologist 8 (2): 116

27. Erosaria (Ravitrona) caputserpentis reticulum
(GmeEtIn, 1791)
Systema Naturae, ed. 13: 3407

28. Erosaria (Ravitrona) caputserpentis kenyonae
SCHILDER & SCHILDER, 1938
Proc. Malac. Soc. London 23 (3): 136
ibid. 3: 77, fig. 2

(Erosaria) TROSCHEL, 1863
Das GebifS der Schnecken 1: 205

> Type Species: Cypraea erosa Linnaeus, 1758 <
Systema Naturae, ed. 10: 723

29. Erosaria (Erosaria) poraria poraria
(LinnAEus, 1758)
Systema Naturae, ed. 10: 724

Page 220

THE VELIGER

Vol. 10; No. 3

30. Erosaria (Erosaria) erosa purissima

(VREDENBURG, 1919)
Journ. Asiat. Soc. Bengal 15: 143

31. Erosaria (Erosaria) miliaris diversa (Kenyon, 1902)
Journ. Conch. 10: 184

32. Erosaria (Erosaria) turdus turdus (Lamarck, 1810)
Ann. Mus. Hist. Nat. Paris 6: 74

Staphylaea JoussEAUME, 1884
Naturaliste 1884: 415

(Staphylaea) JousSEAUME, 1884
> Type Species: Cypraca staphylaea Linnazus, 1758 <
Systema Naturae, ed. 10: 725

33. Staphylaea (Staphylaea) staphylaea staphylaea
(LinnaEus, 1758)
Systema Naturae, ed. 10: 725

34. Staphylaea (Staphylaea) limacina facifer
(IREDALE, 1935)
Austral. Zool. 8 (2): 119; plt. 8, fig. 6

(Nuclearia) JouSSEAUME, 1884
Bull. Soc. Zool. France 9: 98
(Naturaliste 1884: 415, nom. nud.)
> Type Species: Cypraea nucleus Linnazus, 1758 <
Systema Naturae, ed. 10: 724

2h), Staphylaea (Nuclearia) nucleus nucleus

(Linnaeus, 1758)
Systema Naturae, ed. 10: 724

Notocypraea ScHILDER, 1927
Arch. Naturgesch. 91/A 10: 110
> Type species: Cypraca piperita Gray, 1825 <
Zool. Journ. 1: 498 (SoLanpER MS)

(Guttacypraea) IREDALE, 1935
Austral. Zool. 8 (2): 134

> Type species: Cypraca pulicaria REEVE, 1846 <
Conch. Icon. sp. 84, plt. 17, f. 84

36. Notocypraea (Guttacypraea) pulicaria (REEVE, 1846)
Conch. Icon. 3: Cypraea, fig. 84

(Notocypraea) ScuiLper, 1927
> Type species: Cypraea piperita Gray, 1825 <
Zool. Journ. 1: 498 (SoLanpER MS)

37. Notocypraea (Notocypraeca) piperita (Gray, 1825)
Zool. Journ. 1: 498 (SoLanpER MS)

CyPRAEOVULINAE SCHILDER, 1930
Proc. Malacol. Soc. London 19: 120

Erroneini ScHILDER, 1927
Arch. Naturgesch. 91/A 10: 109

Erronea TRoscHEL, 1863
Das GebiB der Schnecken 1: 205
> Type Species: Cypraea errones LINNAEUS, 1758 <

(Adusta) JouSSEAUME, 1884
Naturaliste 1884: 414
> Type Species: Cypraea adusta LAMarcx, 1810 <
Ann. Mus. Hist. Nat. 16: 92
= Cypraea onyx LinnarEus, 1758
Systema Naturae, ed. 10: 722

38. Erronea (Adusta) subviridis dorsalis
SCHILDER & SCHILDER, 1938
Proc. Malacol. Soc. London 23 (3): 149

39. Erronea (Adusta) pyriformis smithi (SowERBY, 1881)
Proc. Zool. Soc. London for 1881: 638

40. Erronea (Adusta) walken continens (IrEDALE, 1935)
Austral. Zoologist 8 (2): 127

(Erronea) TrRoscHEL, 1863
Das Gebif& der Schnecken 1: 205
> Type Species: Cypraeca errones LinNAEuS, 1758 <
Systema Naturae, ed. 10: 723

41. Erronea (Erronea) ovum ovum (GMELIN, 1791)
Systema Naturae, ed. 13: 3412

42. Erronea (Erronea) errones coxi (BRAzIER, 1872)
Proc. Zool. Soc. London for 1872: 617

43. Erronea (Erronea) cylindrica sowerbyana
ScHILDER, 1932
Foss. Cat. 1: Animalia, pars 55, Cypraeacea, 192

44. Erronea (Erronea) caurica blaesa IREDALE, 1939
Austral. Zoologist 9 (3): 322

(Melicerona) IREDALE, 1930
Mem. Queensld. Mus. 10 (1) : 83
> Type Species: Cypraca listerti Gray, 1824 <
Zool. Journ. 1: 384
= Cypraea felina GmEin, 1791
Systema Naturae, ed. 13: 3412

45. Erronea (Melicerona) felina melvillt (Hmatco, 1906)
Mem. Acad. Cienc. Madrid 25: 180

Palmadusta IREDALE, 1930
Mem. Queensld. Mus. 10 (1): 82

(Palmadusta) TrepAteE, 1930
> Type species: Cypraca clandestina LinnaEus, 17/67 <
Systema Naturae, ed. 12: 1177

46. Palmadusta (Palmadusta) asellus asellus

(Linnaeus, 1758)
Systema Naturae, ed. 10: 722

Vol. 10; No. 3

47. Palmadusta (Palmadusta) clandestina clandestina
(Linnaeus, 1767)
Systema Naturae, ed. 12: 1177

48. Palmadusta (Palmadusta) saulae crakei Cate,
subspec. nov.

49, Palmadusta (Palmadusta) lutea bizonata

TREDALE, 1935
Austral. Zoologist 8 (2): 126

50. Palmadusta (Palmadusta) ziczac ziczac

(Linnaeus, 1758)
Systema Naturae, ed. 10: 722

(Purpuradusta) ScuiLpvER, 1939
» Arch. Molluskenk, 71: 165
> Type Species: Cypraea fimbriata Gein, 1791 <
Systema Naturae, ed. 13: 3420

Silke Palmadusta (Purpuradusta) gracilis hilda
(IREDALE, 1939)
Austral. Zoologist 9 (3): 312

52. Palmadusta (Purpuradusta) fimbriata fimbriata
(GMELIN, 1791)
Systema Naturae, ed. 13: 3420

53. Palmadusta (Purpuradusta) hammondae dampierensis
SCHILDER & CERNOHORSKY, 1965
The Veliger 7 (4): 225; plt. 29, figs. 1, 2

Bistolida CossMann, 1920
Rev. Crit. Paléozool. 24: 83
> Type Species: Cypraea stolida LinnaEus, 1758 <
Systema Naturae, ed. 10: 724

(Blasicrura) IrREDALE, 1930
Mem. Queensld. Mus. 10 (1): 84
> Type Species: Cypraea rhinoceros Souversiz, 1865 <
Journ. Conchyl. 13: 156; plt. 511
= Cypraea pallidula Gasxoin, 1849
Proc. Zool. Soc. London for 1848: 97 (Mar. ’49)

54. Bistolida (Blasicrura) quadrimaculata thielei
SCHILDER & SCHILDER, 1938
Proc. Malacol. Soc. London 23 (3): 164

Do: Bistolida (Blasicrura) pallidula simulans
SCHILDER & SCHILDER, 1940
Arch. Molluskenk. 72: 42

56. Bistolida (Blasicrura) teres teres (GMELIN, 1791)
Systema Naturae, ed. 13: 3405

(Derstolida) IrEDALE, 1935
Austral. Zoologist 8 (2): 121
> Type species: Derstolida fluctuans IREDALE, 1935 <
Austral. Zoologist 8 (2): 121
= Cypraea brevidentata SowErRBy, 1870
Thesaur. Conch. (4): 11; fig. 325

THE VELIGER

Page 221

57. Bistolida (Derstolida) kienert kieneri (Hwatco, 1906)
Mem. Acad. Cienc. Madrid 25: 177

58. Bistolida (Derstolida) hirundo cameroni
(IREDALE, 1939)
Austral. Zool. 9 (3): 314; plt. 28, figs, 29 - 31

59. Bistolida (Derstolida) ursellus ursellus (GMELIN, 1791)
Systema Naturae, ed. 13: 3411

(Bistolida) CossMANN, 1920
Rev. Crit. Paléozool. 24: 83
> Type Species: Cypraca stolida Linnazus, 1758 <
Systema Naturae, ed. 10: 724

60. Bistolida (Bistolida) stolida stolida (Linnaeus, 1758)
Systema Naturae, ed. 10: 724

61. Bistolida (Bistolida) brevidentata (SowERBy, 1870)
Thes. Conch. 4 (30): 11; plt. 30, figs. 325 - 326

Cribrarula StRAND, 1929
Acta Univ. Latv. 20: 8
Syn.: Cribraria JoussEauME, 1884 (twice preoccupied)
Bull. Soc. Zool. France 9: 94
> Type Species: Cypraca cribraria Linnaeus, 1758 <
Systema Naturae, ed. 10: 723

(Ovatipsa) IREDALE, 1931
Rec. Austral. Mus. 18 (4): 219
> Type Species: Cypraca chinensis GMELIN, 1791 <
Systema Naturae, ed. 13: 3421

62. Cribrarula (Ovatipsa) chinensis whitworthi
(Cate, 1964)
The Veliger 7 (1): 20; plt. 5, figs. 2a, 2b

(Cribrarula) StRAND, 1929
> Type species: Cypraca cribraria LINNAEuS, 1758 <
Systema Naturae, ed. 10: 723

63. Cribrarula (Cribrarula) cribraria fallax
(E. A. Smiru, 1881)
Ann. Mag. Nat. Hist., ser. 5, 8: 441

DISCUSSION orf tHE SPECIES

1. Bernaya (Bernaya) cate: ScutiLpeErR, 1963
(Plate 21, Figure 1)

Locality: 163
(JS. GSAS SS) 23 7)
Cate (1964, p. 21, no. 41)

This is a unique specimen, collected by a crayfisherman
during the autumn of 1960, on a western beach of West
Wallabi Island, Houtman Abrolhos Islands. The holo-
type is no. 12756 in the California Academy of Sciences,
Department of Geology Type collection.

Page 222

2 Zoila friendi friendii (Gray, 1831)
(Plate 24, Figure 12)

Localities; 18, 19, 25a, 28, 31, 35, 38, 54, 61a, 65, 83, 95,
98, 99, 118, 121a, 142, 143, 145, 172
(God 23.8 273 22 ()
Cate (1964, p. 23, no. 47)

A specimen was collected in 10 feet of water on brown
sponge, mile off Sorrento Beach (145) ; collectors: B. R.
Wilson and R. W. George; January 10, 1962; Cate coll.
no. C 3404.

Four animals (2 of them subadult) were collected in
12 feet of water, living on orange sponge and Pinna shells;
Cockburn, off Woodmans Point on the Parmelia Bank;
collector: N. Mills, St. James Park, West Australia; Feb-
ruary 1964; Cate coll. no. C 3405.

Much has been written about these shells, including the
broad forms occurring in southwestern Australia east of
Cape Leeuwin. However, it is interesting to note that
Heptey (1916) did not mention this species while listing
the mollusks of Western Australia.

In recent years a concerted effort has been made to
ascertain the exact number of cypraeid species now living
in the West Australian coastal waters. Concurrently, a
study of the ranges of their occurrence has been carried
on as well. Pertinent to this report has been the considera-
tion of this particular species and its development and
living aspects, as we were able to find them. It was noted
that Zoila friendii and its variations (excluding the sub-
species Z. f. thersites (Gasxkotn, 1849) and its color variant
named contraria by IREDALE in 1935, appears to range
from Esperance, in southwest Australia, northward along
the west coast to an obscure point to the north of Sorrento
Beach.

It should be noted that the largest specimens of Zozla
friend friendi (Cate coll. no. C 3404: 99.5 51.5 38.3
28 7) seem to be found in shallow water (10 feet) about
# mile offshore at Sorrento Beach. The specimens exhibit
no hint of change in shell form at this locality where the
species approaches the northern end of its range. Shells
here possess the typical rudimentary abapical columellar
teeth, which are large and stubby.

Recent discovery has brought to light a new cowrie
form living in the deep waters west of Shark Bay. In
1965 the brothers William and Wilfred Poole, fishing out
of Fremantle, were trawling in the area west of the Dorre-
Bernier-Koks Island chain (25° 00’ S Long.; 113° 08’ E
Lat.), approximately 40 miles west of Carnarvon. Along
with the outer peninsula of Denham Sound and Dirk
Hartog Island, these islands form the western perimeter
of Shark Bay. It is said the Poole brothers found 10 shells
in about 60 fathoms of water. It is further believed other
specimens have since been taken from this area. WILSON

THE VELIGER

Vol. 10; No. 3

& SuMMERS (1966) list specimens as having been taken
“off Geraldton,” and from False Entrance (False Entrance
is 12 miles south of Steep Point, which marks the southern
shoreline of South Passage; South Passage is the waterway
separating the mainland and Dirk Hartog Island).

On examination, the Dorre-Bernier-Koks Island shells
appear to be morphologically distinct from those found at
Sorrento Beach. I have compared them with 31 shells in
my collection, among which are specimens representing
most of the known localities for this species on the south-
west coast and east into South Australia. In addition, I
was able to examine the shells in both the South Australian
Museum, Adelaide, and the West Australian Museum,
Perth. Except for the obvious morphological change as
observed in the South Australian Zoila friendii vercoi
(Plate 23, Figure 11), the species elsewhere was note-
worthy for its normally uniform shell shape and apertural
count and arrangement. Even so, despite this broader
shell growth in the south coast shells, they are all typical
Z f. friendu in general overall shape and dentition.

The Dorre-Bernier-Koks Island shells, on the other
hand, are not typical, in my opinion. I have examined 6
of these shells and find them to be distinctly different
from any other allopatric form in the Zoila friendii species
complex. The differences will be discussed further in the
following subspecies.

8); Zoila friendii jeaniana Carte, subspec. nov.
(Plate 24, Figure 13)

Localities: 62a, 68, 71a, 83
Shell large, strong, lightweight, humped, globular-ovate;
base sloping inward from outer margins; lip base flat,
columellar base perceptibly convex; terminals prominent,
thin-sided, sharply edged, and more thickly and roundly
formed in front; margins acutely angled, only thinly cal-
loused, vertically broad, with granular texture; aperture
straight, curving abruptly left adapically; teeth numerous,
medium in length, strong, well defined, particularly on
abaxial margin of fossula; fossula deep, without denticles,
milk-white in color; primary shell color on dorsum light
grey, with approximately three narrow white transverse
bands, all of which can be seen through an irregular outer
layer of light chestnut-brown, which becomes an irregular,
darker color immediately above the lateral margin; broad
margins are off-white, with faint touches of beige, loosely
marked otherwise with large, dark brown spots; base
dark chestnut-brown, with same coloring extending over
half the length of teeth and interstices; other half of
teeth and interstices off-white.

Zoila fricndit jeaniana differs from Z. f. friendi (Gray,
1831) in being a shorter, broader, more humped, more
globular-ovate shell; in having a full complement of

Tue VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 21

Figure 1
Bernaya catei ScutLver, 1963 (holotype)

Figure 2
Zoila episema IREDALE, 1939 (holotype)
= Zoila venusta (SOWERBY, 1846)

photographs by C. N. Cate

Tue VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 22

Figure 3
Zoila venusta (SoweRBy, 1846) x1
(Color Variant)

Figure 4 Figure 5
Zoila venusta (SowERBY, 1846) x4 Zoila venusta (SowERBy, 1846)
Geographe Bay Binningup

photographs by C. N. Catz

Vol. 10; No. 3

teeth on both lip and columella; in having broad off-white
lateral margins with large brown spots; in having, in some
specimens, an abruptly curved adapical terminal opening;
in having an unusual color pattern in the dorsal nacre.

Zoila friend jeaniana differs from Z. friendii thersites
(Gasxorn, 1849), a subspecies restricted almost entirely
to South Australia, by a more pyriform, globular-ovate
shell; by the base and part of the teeth being all brown;
by the broad off-white spotted margins; and by the more
numerous teeth on the apertural edges.

The holotype will be deposited in the West Australian
Museum, where it will bear the catalogue number
WAM 1320-67. This new form of Zoila friendii has been
named in honor of Mrs. Jean Cate, Los Angeles, whose
contributions to malacology and its literature are well
known.

A further consideration of consequence is a geographi-
cal separation between the living areas of the new form
and the nominate species, Zoida friendi friendii from
Sorrento Beach of approximately 500 miles. It seems
probable that somewhere between these two localities
there may be an overlapping in the ranges of the two sub-
species. This possibility can be seen in the specimen listed

THE VELIGER

Page 223

as hypotype no. 5 (85.8 52.0 43.0 31 29) from Green
Head, which is about 100 miles north of Sorrento Beach.
It possesses some morphological affinity to Z. f. friendit,
yet at the same time displaying more of the shell charac-
ters of Z. f. jeaniana, suggesting that the Green Head area
could be the point of change in the species.

The specimen from Green Head seems to be analogous
to the pale color forms of Zoila venusta and Z. friendi
thersites (the color form of the latter subspecies known as
“Z. contraria” IREDALE, 1935). The pale form of Z. f.
jeamiana is a pale offwhite with pale beige, irregular
markings, shading to a more intense pale orange color on
either side of both terminals; the base is milk-white, and
the teeth are numerous.

Since having completed this report 3 additional speci-
mens have come to hand; one was trawled in deep water
off Quobba Point. Jack Allen, a fisherman, of Babbage
Island off Carnarvon, was the collector (June 1967),
and the specimen was a gift to me from Mr. George Barker
of Geraldton. This shell (68.5 45.5 38.3 32 27), Cate
coll. no. C 3504, will be identified as hypotype no. 2. The
other two animals were trawled at 30 fathoms, 4 miles
west of Quobba Point, July 1, 1967 (collector not deter-

Table 1

Measurements (in millimeters) and other data, for comparison

Lip Col. Specimen
Species Length Width Height Teeth Teeth Locality Ident. Disposition of Shells
Zoila friendit jeaniana
subspec. nov.
75.7 50.0 40.7 33 29 83 Holotype West Australian Museum no. WAM 1320-67
75.6 50.5 41.4 32 iets 83 Paratype 1 Ned Harrold coll., Perth
(subadult) 67.4 45.3 39.0 28 26 83 Paratype 2 Anthony Kalnins, Mayfields, W. A.

73.1 47.6 39.0 28 22 83 Paratype 3 Theodore Gurr, Carlisle, W. A.
69.3 45.2 36.1 28 21 83 Paratype 4 Theodore Gurr, Carlisle, W. A.
85.8 52.0 43.0 31 29 Tia Hypotype 1 Ned Harrold, Victoria Park, W. A.

Zoila friendu friendu
99.3 51.6 38.3 27 7 145 Hypotype 1 Cate coll. no. G 3404
70.9 41.0 34.0 27 7 172 Hypotype 2 Cate coll. no. C 3497
72.4 37.4 29.5 21 6 172 Hypotype 3 Cate coll. no. G 3497
73.8 38.6 31.1 25 8 6la Hypotype 4 Cate coll. no. C 1712
73.4 38.0 31.0 27 7 6la Hypotype 5 Cate coll. no. C 1712
80.0 40.0 31.0 25 7 12la Hypotype 6 Cate coll. no. C 3498
78.9 41.8 34.5 27 12 95 Hypotype 7 Cate coll. no. C 1711
68.5 36.0 28.6 24 11 95 Hypotype 8 Cate coll. no. C 1711
92.6 49.9 37.0 30 10 25a Hypotype 9 Cate coll. no. CG 3499
78.0 41.0 32.8 25 9 25a Hypotype 10 Cate coll. no. C 3499

Zoila friendii vercoi
83.3 53.1 39.0 26 10 2a Hypotype 1 Cate coll. no. C 1706
76.8 49.2 35.4 24 9 59a Hypotype 2 Cate coll. no. C 3500
73.8 44.0 34.3 2 5 59a Hypotype 3 Cate coll. no. C 3501

Page 224

mined). These are from the Ned Harrold collection, Vic-
toria Park, West Australia. They are the hypotypes no. 3
(68.0 46.8 35.7 29 27) and no. 4 (63.9 42.0 33.1
33 26).

The 9 shells of this new taxon that I have seen give a
convincing indication that it should be considered a dis-
tinct species rather than only a subspecies; but until we
know more about the animal’s soft parts and radula and
its still unknown habitat and geographical distribution,
it seems best to take a conservative approach and consider
it a subspecies of Zoila friendi.

4, Zoila venusta (SowERBy, 1846)
(Plate 21 Figure 2 to Plate 23, Figure 10)

Syn.: C. venusta var. SowerBy, 1846 = Cypraca
thatcheri Cox, 1869
Z. episema IREDALE, 1939
Z. sorrentensis SCHILDER, 1963
Localities: 14, 18, 20, 36, 39, 40, 42, 52, 65, 67, 71, 78,
79, 88, 97, 101, 145, 154, 163

(id 48.8 Zit! 25 9)

CaTE (1964, p. 22, no. 42)

Three specimens were found in 8 to 30 feet of water
at Sorrento Beach (145), living on green algae under
reef ledges, by N. Mills, St. James Park, West Australia
in March 1963; Cate coll. no. C 3409. Another specimen,
Cate coll. no. C 3410, was removed from a crayfish pot
set in 19 fathoms, 5 miles west of Rat Island (130) by
Joseph Burton, Geraldton, in August 1964. Still another
specimen, Cate coll. no. C 3406, was collected by fisher-
man Gordon McAulley, Geraldton, in January 1965 from
a crayfish pot set in 28 fathoms about 22 miles west of
Dongara (52). Two specimens were collected in 20 fect
of water with SCUBA gear, 12 miles south of Mandurah
(97) in 1961; Cate coll. no. C 3408. One animal was
found living in a marine cave at 10 fathoms off Binningup
(20) in January 1964; ex. Edward Nickles, Mandurah;
Cate coll. no. 3194, Yet another animal was found on the
ceiling in a limestone cave in 30 feet of water at Rottnest
Island (131), adjacent to the wreck of the ship Macedon;
kelp and other seaweed was present; the collector was
William Hill, Rottnest Island; October 1963; Cate coll.
no. C 3023.

In Cate (1962), after examination of many specimens
of Zoila venusta, and observing the gradual change in
shell size and structure throughout its range, I was con-
vinced that there was but one species involved, but with
different local ecological variations to be scen at progres-
sive localities. Shells in my collection, and those recently
seen in the South Australian Museum, the West Australian
Museum, and in numerous private collections (one of
which contained approximately 40 specimens from the
Sorrento Beach-Reef area!) in West Australia, appear

THE VELIGER

Vol. 10; No. 3

very definitely to have close affinity, all seeming to possess
a common specics character. The morphological change
commences immediately, starting at Geographe Bay (see
Plate 22, Figure 4), becoming obvious in the arca of
Binningup (Plate 22, Figure 5), then Mandurah (Plate
23, Figure 6), Rottnest Island (Plate 23, Figure 7), and
attaining the greatest change in the waters just north of
Fremantle, Sorrento Beach (Plate 23, Figure 8) to Jurien
Bay (Plate 23, Figure 9), then receding back to “normal”
size and shape at Dongara, Geraldton, Abrolhos Islands
(Plate 23, Figure 10), and north to the Dampier Islands.

The point of recession from the Sorrento Beach-Reef
area variation seems to commence in the waters of Jurien
Bay, and noticeably so at Dongara. It is true, the shells
from north of Geraldton appear more bulbous, darker in
color, and with a pinched, often-times narrower base and
aperture (sce Plate 23, Figure 10). It would seem that
none of the various changes in the shell, including the
pale pinkish-beige to off-white variants (Plate 22, Figure
3) collected at Cervantes Island and Geographe Bay,
are sufficient to justify considering them as more than
localized variants. This appears to be particularly true
because of the continuous range of the species, with no
distinct natural barriers to interbreeding.

I have examined the pale colored shell variants collec-
ted at Cervantes Island (1) and Geographe Bay (3), one
of these 3 in my own collection (no. C 3502: 73.2 48.3
38.8 25 13) that was trawled in 40 fathoms of water
in 1950 by an unidentified Dutch fisherman. Mr. A. R.
Whitworth obtained this shell from Arthur Bassett of
Denham, Shark Bay, in 1960. The Cervantes specimen is
in the Ned Harrold collection, Victoria Park; the other 2
of the 3 specimens from Geographe Bay are in the George
Barker collection, Geraldton, and the Trevor Sutcliffe col-
lection (see ScHILDER, 1966), Mount Yokine, Western
Australia.

oO: Zoila marginata (Gasko1n, 1849)
(Plate 24, Figure 14)

Localities: 2a, 3, 8, 30a, 34a, 36, 52, 53, 57, 67, 68, 71,
71b, 78, 83, 89, 118, 147, 151, 152, 162, 163, 180

(Gifs SO 229 27 22)

Cate (1964, p. 23, no. 49)

A specimen was removed from a crayfish pot set at 28
fathoms 7 miles south of Long Island (94), ex Max
Cramer, Geraldton; October 1953; Cate coll. no. C 2516.
Another animal was removed from a crayfish pot set in
22 fathoms off Dongara (52) in September 1963. The
collector was fisherman F I. “Sonny” Healy, Dongara and
the shell is in his collection. Still another specimen was
found in a crayfish pot that had been set in 20 fathoms
off Leander Reef (53), WSW of Dongara, collected by
fisherman Edward Seabrook, Fisheries Department, Perth.

Tue VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 23

Figure 6
Zoila venusta (SoweErRBy, 1846) Zoila venusta (SOWERBY, 1846)
Mandurah Rottnest Island

Figure 8 Figure 9
Zola venusta (SowERBY, 1846) x¥% Zoila venusta (SowERBY, 1846) x#%
Sorrento Reef Jurien Bay

Figure 10 Figure 11
Zoila venusta (SowERBY, 1846) Zotla friendi vercoi SCHILDER, 1930
Rat Island, Abrolhos Albany, South Australia

photographs by C. N. Cate

Tue VELIcER, Vol. 10, No. 3 [C. N. Cate] Plate 24

Figure 12 Figure 13
Zoila friendii friendit (Gray, 1831) Zoila friendi jeaniana subspec. nov., x 4
Geographe Bay Koks Island, West Australia

Figure 14 Figure 15
Zoila marginata (GaAsKoIN, 1849) Zoila rosselli Corton, 1948
Long Island, Abrolhos Rat Island, Abrolhos

eeeters e

te.

Figure 16
Zoila decipiens (E. A. SmituH, 1880) x1 Maunitia eglantina perconfusa IREDALE, 1935 x#%#
Gourdon Bay, West Australia Roebuck Bay, West Australia

photographs by C. N. Catz

THE VELIcER, Vol. 10, No. 3 [C. N. Cate] Plate 25

Figure 18 Figure 19
Mauritia arabica brunnescens CaTE, 1964 Maurnitia histrio westralis (IREDALE, 1935)
Roebuck Bay, West Australia Roebuck Bay, West Australia

Figure 20 Figure 21
Talparia talpa talpa (LINNAEUS, 1758) Cypraea tigris pardalis SHAw, 1795

Yardie Creek, North West Cape Roebuck Bay, West Australia

zh.

Figure 22 Figure 23
Cypraea argus argus LINNAEUS, 1758 Cypraea lynx vanelli LINNAEUS, 1758 XI
Point Maud Vlaming Head, North West Cape

photographs by C. N. Cate

Vol. 10; No. 3

DWE SVELIGER

Page 225

The shell is in the Cate collection, no. C 3400.

Another specimen was found in a crayfish pot set at
24 fathoms, 3 miles east of Rat Island in Good Friday
Bay (71), Abrolhos Islands; collector: Alec Gilbertson,
Geraldton, 21 November 1957; Cate coll no. C 3395. A
broken shell and a subadult shell, both without animal,
were dredged from 28 fathoms, 16 miles WSW of Gerald-
ton in November 1964 by Alec Gilbertson; the shells are
in his collection.

In comparing the shells from Western Australia with
those from South Australia, one notices that the southern
specimens are more frail, with lighter brown spots more
sparscly arranged in fewer and more shallow concavities;
another noticeable feature is the pale pinkish color within
the shells themselves, while all those I have seen from
western waters have white interiors. Additionally, the base
of the southern shells is flat, while the west coast shells
appcar to have a more swollen, convex base.

In 1961 I referred to Zoila marginata (Cate coll. no.
C 906), citing Albany, Southwest Australia as its locality.
At that time the knowledge concerning this specics was
limited to that of the holotype in the British Muscum,
whose original label shows “New Holland” as its source.
Specimens that came to light after 1961 were all from
the area west of Geraldton, West Australia. The Albany
locality for the specimen mentioned above, therefore,
seemed to be doubtful. This was especially true with no
collecting stations then known south of the Abrolhos
Islands (see Map). In October 1966, Dr. Helene M. Laws
of the South Australian Muscum mentioned to me that 3
specimens of Z. marginata had recently been collected in
South Australian waters: at Cape Jarvis, Taylor Island,
and at a station near Grindel Island. A detailed report is
now documented in Laws (1966). It is reasonable to
surmise, therefore, that Albany may have been the home
port of an carly fishing flect and that one of the fishermen
may have taken the hypotype in a trawl or crayfish pot.

6. Zoila rosselli Cotton, 1948
(Plate 24, Figure 15)

Localitics: 19, 36, 42, 48, 52, 53, 75, 87, 90, 104, 118,

163, 170

(Sd4e Bo 209 2) 23)

CaTeE (1964, p. 22, no. 45)

Once of several specimens was dredged in mud and coral
rubble adjacent to North Wharf, Fremantle (61). This
is one of the Rossell paratypes (Cate coll. no. C 1351).
A specimen was found in approximatcly 6 inches of water,
crawling on algac-covercd substrate at the southwest
corner of Wooded Island (170), Abrolhos Islands, in
1963; cx A. R. Whitworth, Geraldton; Cate coll. no.

C 3288. An animal was removed from a crayfish pot set
in 18 fathoms at Leander Recf (53), off Dongara, in the
spring of 1965; collector was EI. Healy, Dongara; Cate
coll. no. C 3290. Another specimen was removed from a
crayfish pot set in 22 fathoms, 25 miles southwest of
Dongara. The shell is only half-grown, with a razor-edge
thin lip; this is the only bulla-stage specimen of Zoila
rosselli that I have seen; in this form it is indeed rare; the
collector was EI. Healy of Dongara; January 3, 1965;
Cate coll. no. C 3289. An additional specimen was re-
moved from a crayfish pot at Ledge Point (87), set in 30
fathoms in July, 1965; ex A. R. Whitworth, Geraldton;
Cate coll. no. C3401. One animal was collected inter-
tidally on algae-covered coral at the west end of Rat
Island (130), Abrolhos Islands; ex A.R. Whitworth,
Geraldton, 1965; Cate coll. no. C 3402. The most recently
collected specimen was removed from a crayfish pot set
in 80 fathoms (this is exceptionally deep for craypots) off
the southwest bank at North Island (104a). It was ad-
hering to animal bait (cow hock); William Thompson,
Geraldton; April 1967.

Ds Zoila decipiens (E. A. Smirn, 1880)
(Plate 24, Figure 16)

Localities: 7, 29, 44, 60, 80, 119

(SD) BU B30 22 ils)

Cate (1964, p. 22, no. 45)

Locality records of the South Australian Muscum list
these additional collecting stations for Zoila decipicns:
Cossack, near Port Walcott; ‘King (George) Sound’; and
Fremantle (H. Rossell, So. Austral. Mus. no. 11636). This
latter locality is considerably farther south than the
hitherto southern-most authenticated record known to me,
namcly, Learmouth at Wapet Creck in Exmouth Gulf
(86). This shell was picked up on the beach after Cyclone
“Katic,” Easter Sunday 1964 by Mrs. Molly Gedling,
one of the keepers of the Viaming Head Light; Cate coll.
no. C 3112. Otherwise, the most northern record we have
been able to authenticate is a specimen collected at Wynd-
ham, Cambridge Gulf, 250 miles south of Darwin (44) ;
Cate coll. no. C 3399,

8. Mauritia (Arabica) eglantina perconfusa
TIREDALE, 1935

(Plate 24, Figure 17)
Localitics: 10, 37, 43, 46, 47, 49, 54, 93, 98, 108, 111, 117,
IIE), 6 EO SB, aS, UG TZ, 17/83
(C22 30-2, SOD 83 31))
Cate (1964, p. 24, no. 53)
[For a discussion of this specics sec: Cater, 1964]

Page 226

9. Mauritia (Arabica) arabica brunnescens Cate, 1964
(Plate 25, Figure 18)

Localities: 10, 15, 108, 176
(61.7 36.7 30.1 28 24)
Cate (1964, p. 24, no. 54)

[For a discussion of this species see: CaTE, 1964]

10. Mauritia (Arabica) histrio westralis (IREDALE, 1935)
(Plate 25, Figure 19)

Localities: 24, 45, 60, 115

(102 459 B52 30. 22)

Cate (1964, p. 25, no. 55)

Weaver (1960) listed “Mauritia maculifera ScuIL-
DER,” beach collected on the east side of Long Island. This
collecting station is just north of the land end of North
West Cape on the outer perimeter of Exmouth Gulf. Based
on our present knowledge of the West Australian cowries,
and upon a subsequent personal conversation with Mr.
Weaver, it seems likely the shells found were specimens
of M. histrio westralis. There can be little doubt of the
close relationship between these two species, but one can
observe the smaller, shorter, and more numerous teeth
and lack of the central, though diffused, coloring on the
base, both of which clearly distinguish this western
species.

iil. Talparia talpa talpa (Linnaeus, 1758)
(Plate 25, Figure 20)

Localities: 60, 64, 153, 159,176
(68.0 37.9 33.3 44 44)
Cate (1964, p. 24, no. 52)

[For a discussion of this species see: CaTE, 1964]

12. Cypraea (Cypraea) tigris pardalis SHaw, 1795
(Plate 25, Figure 21)

Localities: 2, 68, 102, 108, 109, 114, 119, 124, 142, 162,
163, 170, 176

(89.1 60.4 47.0 27 22)

CaTE (1964, p. 26, no. 56)

A specimen (111.2 73.2 57.1 24 24) was found in
approximately one foot of water on a portion of the reef
extending out from the southwest shore of West Wallabi
Island (163).It was collected by William McWade of
Geraldton in July 1966; Cate coll. no. C 3417. Another
specimen was found at Wooded Island (170), Abrolhos
Islands, in 3 feet of water on algae-covered coral, by A. R.
Whitworth, Geraldton in the spring of 1965; Cate coll.
no. C 3418.

THE VELIGER

Vol. 10; No. 3

13. Cypraea (Lyncina) argus argus LinNAEvs, 1758
(Plate 25, Figure 22)

Localities: 44, 69, 70, 98, 106, 109, 159, 160, 176

(91.3 49.6 37.7 43 40)

Cate (1964, p. 24, no. 51)

A specimen was found by Mrs. Alan Nicol, Carnarvon,
at Glenroy (69), 10 miles south of Onslow, in the beach
drift; Cate coll. no. C 3396. In June 1964, a dead, broken
shell was found in a tide pool 4 miles southwest of
Vlaming Head Light (160), North West Cape, by Mrs.
Molly Gedling; Cate coll. no. C 3053. Another specimen
was found on the beach just south of Vlaming Head Light
after a heavy southwest wind storm in August, 1964, also
by Mrs. Gedling; Cate coll. no. C 3107.

14. Cypraea (Lyncina) lynx vanelli Linnazus, 1758
(Plate 25, Figure 23)

Localities: 2, 15, 24, 37, 43, 44, 48, 59, 63, 86, 108, 114,

117, 119, 142, 150, 153, 163, 176, 178

(4097 24:1) 20:6) 26 919)

Cate (1964, p. 26, no. 57)

The shells of this species exhibit great variation in
size, color, and shape, wherever it occurs. In some locali-
ties this variability is more pronounced than in others;
the base, teeth, aperture, and interstices are the most
constant shell characters.

15. Cypraca (Lyncina) vitellus vitellus Linnazus, 1758
(Plate 26, Figure 24)

Localities: 2) 6; 8;10) 16; 35547552) 5/7) G2NGSaa 2 oe
86; 89; 90; 94; 100; 104; 108) 10 Tae oracle
150, 153, 163, 170, 176, 178

(49.7 31.4 27.8 26 23)
Cate (1964, p. 26, no. 58)
[For a discussion of this species see: Cate, 1964]

16. Cypraca (Lyncina) reevei SowErsy, 1832
(Plate 26, Figure 25)

Localities: 18, 20, 32, 36, 39, 40, 52, 53, 68, 88, 156, 163,

175

(B42 21-9) 1857229522)

CaTeE (1964, p. 26, no. 59)

A specimen was found in a crayfish pot set in 15
fathoms adjacent to Turtle Dove Shoal (156), WSW of
Dongara, ex James Seabrook, Fisheries vessel Lancelin,
Perth, March 1964; Cate coll. no. C 3397. Another animal
was collected intertidally under a rock on the north ap-
proach to Cape Naturaliste; B. R. Wilson collected it in
November 1965; Cate coll. no. C 3398.

Tue VELIGER, Vol. 10, No. 3 [C. N. Care] Plate 26

Figure 24 Figure 25
Cypraea vitellus vitellus LINNAEUS, 1758 Cypraca reevei SOWERBY, 1832 x2
Broome, Roebuck Bay [normal form] Geographe Bay

4
ae:
~

a}
“Fa)
an
.)
e
>
Ps

Figure 26 Figure 27
Cypraca reevei SowERBy, 1832 Cypraea carneola carneola LinNazus, 1758
[spherical form] Dongara Vlaming Head, North West Cape

i
Figure 28 Figure 29
Cypraca leviathan gedlingae subspec. nov. Luria isabella rumphii ScuiLpER & SCHILDER, 1938 x 3
Five Mile Beach, North West Cape Broome, Roebuck Bay

photographs by C. N. Catz

Tue VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 27

Figure 30 Figure 31
Pustularia cicercula cicercula (LINNAEUS, 1758) x3 Pustularia globulus globulus (LINNAEUS, 1758) x3
Vlaming Head, North West Cape Yardie Creek, North West Cape

Figure 32 Figure 33
Monetaria annulus annulus (LINNAEUS, 1758) x2 Monetaria moneta rhomboides SCHILDER & SCHILDER,
1

Vlaming Head, North West Cape 1933 Xz
Broome, Roebuck Bay

Figure 34 Figure 35
Erosaria 1. labrolineata (GasKoIN, 1849) Erosaria cernica viridicolor (CATE, 1962) x2
Exmouth Gulf Vlaming Head, North West Cape

photographs by C. N. Catz

TuHE VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 28

Figure 36 Figure 37
Erosaria caputserpentis reticulum (GMELIN, 1791) x3 Lrosaria caputserpentis kenyonae (SCHILDER &

Yardie Creek, North West Cape SCHILDER, 1938) x7
Quobba Point

Figure 38 Figure 39
Erosaria helvola citrinicolor IREDALE, 1935 x2 Erosaria poraria poraria (LINNAEUS, 1758) x3
Vlaming Head, North West Cape Vlaming Head, North West Cape

Figure 40 Figure 41
Erosaria erosa purissima (VREDENBERG, 1919) x{ Erosaria miliaris diversa (KENYON, 1902) x2
Broome, Roebuck Bay Light House Point, Broome

photographs by C. N. Cate

Vol. 10; No. 3

THE VELIGER

Page 227

It should be noted that Cypraca reever undergoes a
very striking morphological change in shell form from
Jurien Bay northward. In the Dongara area particularly,
the shell loses its familiar clongately-ovate shape, becoming
almost spherical (Plate 26, Figure 26). The shell is
arresting in appearance, with the rear terminal flattened
out and the front terminal hardly discernible; this, and its
bulbously-humped appearance give it a nearly round con-
figuration. Cate coll. no. C 3397 and no. C 3411.

17. Cypraea (Lyncina) carneola carneola LINNAEUS, 1758
(Plate 26, Figure 27)

MOcaAlitiese2 Os Oy Oy los 305.30) 44.047) 04, 917,60) 63;
Sone9) 1085 1105 11, 114, 119) 142° 150; 153; 156; 157,
163, 176

(290 U2 Ses 23) 22)
Cate (1964, p. 27, no. 60)
A specimen was found under dead coral at Alec Finlay’s

Island (5), Abrolhos Islands, by Terry Butterworth of

Geraldton, in November 1964; Cate coll. no. C 3416.

18. Cypraea (Lyncina) leviathan gedlingae CaTE
subspcec. nov.
(Plate 26, Figure 28)

Localities: 64, 106, 159, 176, 177

(64.9 38.7 32.5 33 27)

Shell large, long, heavy, massive, cylindrically-ovate,
dorsally elevated; base and outer lip rounded, convex,
solid, of uneven contour, bumpy; terminals large, thick,
ponderous; aperture wide, straight, curving gently left
adapically; teeth short, thick, distant, strong on outer lip,
finer, longer, crossing columella and fossula to adaxial
edge; fossula short, wide, deepening toward the front;
margins thick, heavily calloused, unevenly swollen, weakly
shouldered; primary dorsal color pale tomato-red, four-
banded; margins, terminals and base beige, except that
the upper margins and, to a lesser extent, the sides are
granular appearing darker beige; beige callus over and
around terminals noticeably broadened; teeth and inter-
stices medium lavender.

The holotype (Plate 26, Figure 28) was collected at a
remote place on the outer coast of the North West Cape
peninsula, locally called the “Five Mile Beach” (approx-
imately 22°06’ S Lat., 114° 01’ E Long.), the distance
being measured southward from the Vlaming Head Light.
The statistical details of the holotype arc: length 80.6mm,
width 45.3 mm, height 39.6 mm, 36 tecth on lip and 33 on
the columella. The holotype will be deposited in the Type
Collection of the West Australian Muscum, Perth, West-
ern Australia, where it will bear the catalogue number
WAM 334-67.

Other specimens were collected between the above
location and Yardie Creek (176), the most southerly
locality where one additional shell was found. This new
subspecies has been named for Mrs. Molly Gedling, Rott-
nest Island Main Light, Western Australia. Mrs. Gedling
with her husband Robert, formerly keepers of the Vlaming
Head Light, collected on the beaches and reefs of both
the inner and outer coasts of the North West Cape. They
are largely responsible for the information we now have
concerning cowries in this area.

119} Luria (Basilitrona) isabella rumphi
SCHILDER & SCHILDER, 1938
(Plate 26, Figure 29)
Localities: 44, 47, 59, 107, 119, 153, 159
(233122392 0!ome2 One 20)
Cate (1964, p. 24, no. 50)
[For a discussion of this species see: CaTE, 1964]

20. Pustularna (Pustularia) cicercula cicercula
(LinnagEus, 1758)
(Plate 27, Figure 30)
Localities: 47, 153, 159, 176
(76 W2gb Nil 27 23
Cate (1964, p. 11, no. 1)
[For a discussion of this species see: Cate, 1964]

Zale Pustularia (Pustularia) globulus globulus
(Linnaeus, 1758)
(Plate 27, Figure 31)

Localities: 60, 159, 176
(TAS Goll Goll 27 iB)
Cate (1964, p. 12, no. 2)
[For a discussion of this species see: CaTE, 1964]

22. Monctaria (Ornamentaria) annulus annulus
(Linnaeus, 1758)
(Plate 27, Figure 32)
Localities: 10; 15, 22; 25, 36, 47, 93, 104, 108, 119;.153,
176
(2354 Ol Simla Sinn)
Cate (1964, p. 15, no. 14)
[For a discussion of this species see: Cate, 1964]

23. Monctaria (Monetaria) moneta rhomboides
SCHILDER & SCHILDER, 1933
(Plate 27, Figure 33)

Localities $4, @, Qe INO), IS, Zi, HO, BH, 428), 227, D2, Te, (G0.
O2MOS i 2 7OO OER SO al OOM OA a OS st ONmItelemtn 9:

Page 228

141, 146, 153, 163, 169, 171, 175, 176
(Bila) Bilge} USS) 1) 122))
Cate (1964, p. 15, no. 15)

[For a discussion of this species see: CATE, 1964]

24. Erosaria (Ravitrona) labrolineata labrolineata
(Gasxoin, 1849)
(Plate 27, Figure 34)

Localities: 24, 44, 94, 119, 176
(lege 10.8 @d iy ies)
CaTE (1964, p. 12, no. 6)

[For a discussion of this species see: Cate, 1964]

25. Erosaria (Ravitrona) cernica viridicolor (CATE, 1962)
(Plate 27, Figure 35)

Wocealities:9 365.53, 715 G5, 104 Ie 142 e150 53156;
159, 162, 163, 164, 165, 176
(2254 ZOO SaaS)
Cate (1964, p. 13, no. 7)
Three specimens were found on the beach at Vlaming
Head (159), North West Cape, on Easter Sunday 1964,
after “Cyclone Katie” by Mrs. Molly Gedling. Cate coll.
no. C 3419. Specimens were removed from a crayfish pot
set in “deep” water off North Island (104), Wallabi
Group, Abrolhos Islands, by W. Hemsley of Geraldton, in
May, 1964. Cate coll. no. C 3413. Another specimen was
collected in a crayfish pot in 12 fathoms, west of Cervantes
Island (36). This shell is in the Ned Harrold collection,
Victoria Park, West Australia.

26. Erosaria (Ravitrona) helvola citrinicolor
IREDALE, 1935
(Plate 28, Figure 38)

Localities: 4, 8, 15, 36, 52, 57, 62, 68, 72, 75, 89, 91, 92,
he Sey MO, Mois}, Tah 2 Male) AYO), 58}, G8), 18, 170,
171, 176

(See WAS)” GS ig 1183)
Cate (1964, p. 13, no. 8)
[For a discussion of this specics sec: Catr, 1964]

27. Erosaria (Ravitrona) caputserpentis reticulum
(GmeEtIn, 1791)
(Plate 28, Figure 36)
Localities: 2, 6, 10, 15, 47, 93, 94, 108, 112, 119, 153, 155,
159
(G20 Peo NOs Alle) 312))
Cate (1964, p. 13, no. 9)
[For a discussion of this species see: Cate, 1964]

THE VELIGER

Vol. 10; No. 3

28. Erosaria (Ravitrona) caputserpentis kenyonae
SCHILDER & SCHILDER, 1938
(Plate 28, Figure 37)

Localities: 4, 8, 9, 16, 21, 35, 36, 37, 43, 52, 57, 58, 62, 68,
72, 75, 86, 89, 90, 91, 92, 100, 104, 105, 110, 111, 140,
141, 146, 150; 159; 163; 167; 169, 170) 171) vamos 7,

(A 228 iil i718)
Cate (1964, p. 13, no. 10)

[For a discussion of this species see: CATE, 1964]

29. Erosaria (Erosaria) poraria poraria
(Linnaeus, 1758)
(Plate 28, Figure 39)

Localities: 60, 159, 176
(164 10.8 83 19 13)
Cate (1964, p. 14, no. 11)

[For a discussion of this species see: CaTE, 1964]

30. Erosaria (Erosaria) erosa purissima
(VREDENBURG, 1919)
(Plate 28, Figure 40)

Localities: 44, 104, 108, 119, 163, 176, 178
(41.4 258 186 18 14)
Cate (1964, p. 14, no. 12)

[For a discussion of this species see: Cate, 1964]

31. Erosaria (Erosaria) miliaris diversa (Kenyon, 1902)
(Plate 28, Figure 41)

Localities: 33, 47, 108, 119, 153
(Bibl 22 iGO) Bil 118)))
Cate (1964, p. 14, no. 13)

[For a discussion of this species see: Cate, 1964]

32. Erosaria (Erosaria) turdus turdus (LAMarRcK, 1810)
(Plate 29, Figure 42)

Locality: 119

(Cou ZS) We iS) iS)

This specimen (Cate coll. no. C 3421) was picked up
in the beach rubble on Lancelin Island (85) by Bernard
E. Bardwell, Broome, in 1924. The species appears not to
have been previously reported from West Australia, and
this must remain a questionable species record until more
definite proof of its occurrence here can be established. A
second specimen of this species from West Australia is in
the collection of Anthony Kalnins, Perth. His specimen
(40.2 29.1 19.7 18 15) is reputed to have been col-
lected in the same general area of Port Hedland (119).
With the Erythraean Mauritia histrio (Gmeuin, 1791)
(as M. histrio westralis (IREDALE, 1935)) ranging into

THE VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 29

Figure 42 Figure 43
Erosaria turdus turdus (LamarcK, 1810) x Staphylaea st. staphylaea (LINNAEUS, 1758) x3
Lancelin Island Vlaming Head, North West Cape

Figure 44 Figure 45
Staphylaea limacina facifer (IREDALE, 1935) x2 Staphylaea nucleus nucleus (LINNAEUS, 1758)
Vlaming Head, North West Cape Viaming Head, North West Cape

Figure 46 Figure 47
Notocypraea pulicaria (REEVE, 1846) X 3 Notocypraea piperita (Gray, 1825)
Busselton, Geographe Bay Cape Naturaliste

photographs by C. N. Catz

Tue VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 30

Figure 48
Erronea subviridis dorsalis (SCHILDER & SCHILDER,
1938) x2
Broome, Roebuck Bay

Figure 49
Erronea pyriformis smithi (SOwERBY, 1881)
Disaster Bay, King Sound

Figure 50
Erronea walkeri continens (IREDALE, 1935) x2 Erronea ovum ovum (GMELIN, 1791) x14
Broome, Roebuck Bay Light House Point, Broome

Figure 52 Figure 53
Erronea errones coxi (BRAzIER, 1872) x1} Erronea cylindrica sowerbyana ScHILDER, 1932 x 12
Weedong, Pender Bay Broome, Roebuck Bay

photographs by C. N. Cats

THE VELIGER, Vol. 10, No. 3

Figure 54
Erronea caurica blaesa IREDALE, 1939
Tautibiddi Well, North West Cape

Figure 56
Palmadusta asellus asellus (LINNAEUS, 1758) x3
Old Onslow

Figure 58
Palmadusta lutea bizonata IREDALE, 1935
Vlaming Head, North West Cape

photographs by C. N. Cate

[C. N. Cate] Plate 31

Figure 55
Erronea felina melvilli (Hwatco, 1906)
Yardie Creek, North West Cape

Figure 57
Palmadusta cl. clandestina (LINNAEUS, 1767)
Broome, Roebuck Bay

Figure 59
Palmadusta ziczac ziczac (LINNAEUS, 1758)
Five Mile Beach, North West Cape

2S)

83

Vol. 10; No. 3

West Australian waters it seems conceivable that this
species could establish itself in these waters as well.

33. Staphylaea (Staphylaea) staphylaca staphylaea
(Linnaeus, 1758)
(Plate 29, Figure 43)

Localities: 52, 58, 159, 176
(155° Oze Ws) Bil ZO)
CaTE (1964, p. 12, no. 3)

[For a discussion of this species see: Carr, 1964]

34, Staphylaea (Staphylaea) limacina facifer
(IREDALE, 1935)
(Plate 29, Figure 44)

Localities: ove, (Sh, Mls}, IT, BS), G8), a7
(159 G2 Ted 19" il)
CaTE (1964, p. 12, no. 4)

[For a discussion of this species see: Cate, 1964]

35. Staphylaea (Nuclearia) nucleus nucleus
(Linnaeus, 1758)
(Plate 29, Figure 45)

Localities: 60, 108, 159, 176
(Be WAS IIs) 22) ills)
Cate (1964, p. 12, no. 5)

[For a discussion of this species see: CaTE, 1964]

36. Notocypraea (Guttacypraca) pulicaria (REEVE, 1846)
(Plate 29, Figure 46)

Localities: 31, 42, 88, 131, 168
(18.6 10.6 8.2 25 24)
Cate (1964, p. 17, no. 23)

[For a discussion of this species sec: CATE, 1964]

37. Notocypraea (Notocypraca) piperita (Gray, 1825)
(Plate 29, Figure 47)

Localities: 3, 26, 30, 32, 39, 67, 76

(2ZIQ Wil GS MW) a7)

Generally speaking, this is a South Australian specics
and is not often found in west southwest Australian watcrs.
The South Australian Muscum, Adelaide (Bernard C.
Cotton Collection), however, records it living as far
north on the west coast as Fremantle. I have 4 specimens
collected in Bunker Bay (26), 4 mile cast of Cape Natu-
raliste (Cate coll. no. C 838) from the B. E. Bardwell
collection, Broome.

Many of the recognized spccics in the complex genus
Notocypraca have undoubtedly arisen from a common an-
cestor, the most noticcably divergent forms being N. puli-
caria and N. piperita. There has been much spcculation

THE VELIGER

Rage 229

concerning the other similar species in this genus. It would
appear that separating them becomes little more than a
question of convenicnce-splitting in many cases. Especially
in the N. angustata (GMELIN, 1791), N. comptoni (Gray,
1847), N. bicolor (Gasxorn, 1849), etc. species complex,
one usually has little difficulty in sorting out many varia-
tions to be found in a group of mixed specimens. However,
N. pulicaria and N. piperita are easily separated from
other members of the genus because of their readily
recognizable shape and prominent color markings. These
latter two species are essentially South Australian but do
range into west southwest Australian waters.

38. Erronea (Adusta) subviridis dorsalis
SCHILDER & SCHILDER, 1938
(Plate 30, Figure 48)

Localities: 2, 8, 23, 33, 47, 89, 104, 108, 111, 118 ,119,

153, 163, 176

(2. Idd USS) 19 6)

Cate (1964, p. 15, no. 16)

A specimen was taken from a crayfish pot set in “deep”
watcr off North Island (104), Abrolhos Islands. O.
Hewitt of Geraldton was the collector in 1963. Cate coll.
no. C 3414.

39. Erronea (Adusta) pyriformis smithi (SowerRBy, 1881)
(Plate 30, Figure 49)
Localities: 41, 44, 45, 81, 112, 113
(Adkss isigs} iI} iS) 31))
Cate (1964, p. 15, no. 17)
[For a discussion of this species see: Cate, 1964]

40. Erronca (Adusta) walker continens (IREDALE, 1935)
(Plate 30, Figure 50)
Localities: 24, 121
(23.9) Wad) WN Bil NG)
Cate (1964, p. 15, no. 18)

[For a discussion of this species see: CATE, 1964]

41. Erronnea (Erronea) ovum ovum (GMELIN, 1791)
(Plate 30, Figure 51)

Localities: 17, 44, 49, 60, 110
(26.3 15.5 13.0 16 15)
Cate (1964, p. 16, no. 19)

[For a discussion of this species see: Cate, 1964]
42. Erronea (Erronea) crrones coxi (BRAziER, 1872)
(Plate 30, Figure 52)

JLoealltsess Ga, Bis 433, 0, G8, NOB, Moe wiles Wily, IONS)
(J20) MBI Weis iG Ik)

Page 230

Cate (1964, p. 16, no. 20)
[For a discussion of this species see: Cate, 1964]

43, Erronca (Erronea) cylindrica sowerbyana
ScHILDER, 1932
(Plate 30, Figure 53)

Localities: 10, 47, 105, 153, 176
(ARQ) Wabi} 128 ils 20)
Cate (1964, p. 16, no. 21)

[For a discussion of this species see: Care, 1964]

44. Erronea (Erronea) caurica blaesa IREDALE, 1939
(Plate 31, Figure 54)

Localities: 10, 14, 33, 37, 43, 47, 73, 80, 93, 108, 111, 113,

i, UNO), WAS, LO), 153, 155, 168 7, W7O, ia

(Grlg3 BSatk Af) Ilks} 15))

Cate (1964, p. 16, no. 22)

Specimens were found at Inner (West) Beach (171),
one mile north of Wreck Point, which is located at the
west end of Pelsart Island, Abrolhos Islands. Collector:
James Seabrook, Fisheries vessel Lancelin, March 1965.
Cate coll. no. C 3415.

45. Erronea (Melicerona) felina melvilli (Hmauco, 1906)
(Plate 31, Figure 55)

Locality: 176
Shell data: Length 16.7, width 9.8, height 7.3mm; num-
ber of teeth on lip - 13, on columella - 13.

As far as we have been able to ascertain, this species is
very rare in West Australia. A single dead specimen was
found wedged in the bottom of a tide pool at Yardie
Creek (176), approximately 7 miles south of Vlaming
Head Light House, North West Cape — the only spceci-
men of Erronea felina melvilli known to me from West
Australia. Even though the shell was dead-collected, it
would seem probable that the species lives in this arca.
The locality is so remote, so sparsely settled that the possi-
bility of its having been artificially introduced scems
unlikely. However, until living specimens are collected in
the area I am listing it only tentatively in the West
Australian fauna.

46. Palmadusta (Palmadusta) asellus asellus
(Linnaeus, 1758)
(Plate 31, Figure 56)

Localities: 24, 60, 108, 119, 159, 160
(U7 ON} BB) yes)
Cate (1964, p. 17, no, 24)

[For a discussion of this species see: Cate, 1964]

THE VELIGER

Vol. 10; No. 3

47. Palmadusta (Palmadusta) clandestina clandestina
(LinnagEus, 1767)

(Plate 31, Figure 57)

Localities: 8, 15, 33, 37, 43, 47, 98, 108, 111, 117, 119,
VEZ ISS G2 ello Seale/G
(Ue) lS Os 28). 15)
Cate (1964, p. 17, no. 25)
[For a discussion of this species see: CaTE, 1964]

48. Palmadusta (Palmadusta) saulae crakei Cate,
subspec. nov.
(Plate 34, Figures 73a, 73b, 73c)

Localities: 117a, 119, 124

(AKO) ISA Wiley 22 iq)

Cate (1964, p. 17, no. 26)

Shell large, broad, narrowing quickly abapically, bul-
bously ovate, umbilicate; teeth short, well defined on
outer lip, intermittently so on columella, but extending
across fossula ; front and rear terminals noticeably rostrate;
right margin thick, left side uncalloused, curving smoothly
from dorsum to base; primary shell color pearl-grey, with
large central brown dorsal blotch that covers at least 20%
of upper surface, the rest of dorsum thickly sprinkled with
minute brown spots; margins copiously flecked with larger
brown spots, with a brown spire blotch; base mostly devoid
of spotting, of a lighter basic grey color, as are teeth;
terminal openings, columella, fossula, and interstices bright
orange-yellow; foot and mantle brilliant orange, marked
with fawn-colored spots.

Palmadusta saulae crakei differs from P. saulae nugata
in having a larger, broader, heavier shell; by the orange
interstices (PR s. nugata normally does not have color in
the interstices) ; by larger and heavier teeth; and by larger
and more prominent chestnut spotting at the margins. It
differs from P saulae siasiensis Cate, 1960 (Plate 34,
Figure 75) by a larger, wider, heavier shell; by a wider,
more curving aperture; by heavier, more well-defined
teeth; and by the larger, more numerous dorsal spots.

This new molluscan form is restricted, as presently
known, to the area from Port Hedland (119) north to
Port Darwin (117a). The specimen illustrated here was
collected at Quondong, a popular fishing area 35 miles
north of Broome, which is here designated the type loca-
lity (17°59% S Lat; 122° 14° E Long). Its range ais
distinctly separated geographically from its nearest allo-
patric relative, Palmadusta saulae nugata IREDALE, 1935,
from the Queensland coast, eastern Australia..

This new subspecies is presently known from 3 speci-
mens: one from Port Darwin, the Port Hedland shell, and
the specimen discussed in this paper. There is also an un-

THE VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 32

|
|

Figure 60 F igure 61
Palmadusta gracilis hilda (IREDALE, 1939) x3 Palmadusta fimbriata fimbriata (GMELIN, 1791) x4
Cape Preston, Onslow Vlaming Head, North West Cape

Figure 62 Figure 63

Palmadusta hammondae dampierensis ScHILDER & —Bistolida quadrimaculata thielet (SCHILDER &
CERNOHORSKY, 1965 x4 SCHILDER, 1938) x2}

Port Hedland Sunday Island, King Soun

$

Figure 65
Bistolida teres teres (GMELIN, 1791) x 14
Broome, Roebuck Bay

Bistolida pallidula simulans ScHILDER & SCHILDER,
1940 x2}
Broome, Roebuck Bay
photographs by C. N. Cate

THE VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 33

Figure 66
Bistolida kieneri kienert (Hipatco, 1906) Bistolida hirundo cameroni IrEDALE, 19389 x3
Broome, Roebuck Bay Broome, Roebuck Bay

“def

oy re

Figure 68 Figure 69
Bistolida ursellus ursellus (GMELIN, 1791) Bistolida stolida stolida (LINNAEUS, 1758) x 2
Broome, Roebuck Bay Five Mile Beach, North West Cape

Figure 70 Figure 71
Bistolida brevidentata (SowERBY, 1870) Cribrarula chinensis whitworthi (CatE, 1964)
Cable Beach, Broome Vlaming Head, North West Cape

photographs by C. N. Catz

THE VELIGER, Vol. 10, No. 3 [C. N. Cate] Plate 34

Figure 73 a
Cribrarula cribraria fallax (E. A. SMITH, 1881) x2 Palmadusta saulae crakei subspec. nov.
Broome, Roebuck Bay Broome, Roebuck Bay

Figure 73b oe y Figure 13.¢
Palmadusta saulae crakei subspec. nov. ¢ Palmadusta saulae crakei subspec. nov.
Left Lateral Aspect Right Lateral Aspect

{

Figure 74 Figure 75
Palmadusta saulae saulae (Gasxorn, 1843) x14 Palmadusta saulae siasiensis (CATE, 1960)
Manila Bay, Philippines Siasi Island, Sulu Sea

photographs by C. N. Cate

Vol. 10; No. 3

THE VELIGER

Page 231

verified shell in a Broome collection. A detailed description
will be found in Cate (1964), with other pertinent infor-
mation.

For comparative purposes I have added illustrations of
Palmadusta saulae saulae (GAskoin, 1843) (Plate 34,
Figure 74) and P saulae siasiensis (Plate 34, Figure 75) to
more clearly illustrate the differences to be found in these
shells. Palmadusta saulae crakei differs from the typical
Philippine P. s. saulae by being a larger, broader, heavier
shell; by having more abundant and larger chestnut-brown
spots on the dorsum as well as a larger dorsal color blotch;
by having the terminals less protruding, more massive, and
without color on the terminal collars; by the absence of
the large brown spots usually found on the margins; by
the paler orange in the interstices; and by the aperture,
which curves more at cither end of the shell.

The holotype will be deposited at the West Australian
Museum, Perth, where it will bear the catalogue number
WAM 1321-67. This subspecies is named in honor of
Theodore Crake, who probably found the first animal of
this taxon living in Western Australian waters. Mr. Crake
has also assisted to a great degree in the study of the
cowries occurring in the Broome area and its environs.

Ag. Palmadusta (Palmadusta) lutea bizonata
TREDALE, 1935
(Plate 31, Figure 58)

Localities: 33, 47, 108, 119, 176
(16.5 9D G8 is 117)
Cate (1964, p. 18, no. 27)

[For a discussion of this species see: Cate, 1964]

50. Palmadusta (Palmadusta) ziczac ziczac
(Linnaeus, 1758)
(Plate 31, Figure 59)

Localities: 119, 157, 159, 176
(IGA) OF dod S183)
CaTeE (1964, p. 18, no. 28)

[For a discussion of this species see: Cate, 1964]

51. Palmadusta (Purpuradusta) gracilis hilda
(IREDALE, 1939)
(Plate 32, Figure 60)

Macalities: ly) 335.395) 5/1, 49,1005.93,90, 1035 108, 117;
IG), 138}, WS). 7
(U2 WO Zee) . iS) lés))
CaTeE (1964, p. 18, no. 29)

[For a discussion of this specics sec: Cate, 1964]

52, Palmadusta (Purpuradusta) fimbriata fimbriata
(GmeELIn, 1791)
(Plate 32, Figure 61)

Localities: 58, 119, 121, 159
(HAIL 3) Bail 113) 1@))
CaTE (1964, p. 18, no. 30)
[For a discussion of this species see: Cate, 1964]

53. Palmadusta (Purpuradusta) hammondae dampierensis
SCHILDER & CERNOHORSKY, 1965
(Plate 32, Figure 62)

Localities: 46, 57, 119, 126, 166, 176
(IDB CH Weil Gs 1G)
Cate (1964, p. 18, no. 31)

[For a discussion of this species see: Cate, 1964]

54. Bistolida (Blasicrura) quadrimaculata thielei
SCHILDER & SCHILDER, 1938
(Plate 32, Figure 63)

Localitics: 82, 121
(AV9) 1s WO il G)
Cate (1964, p. 19, no. 32)

[For a discussion of this species see: Cater, 1964]

55), Bistolida (Blasicrura) pallidula simulans
SCHILDER & SCHILDER, 1940
(Plate 32, Figure 64)

Localities: 108, 116, 119
(CASH) OZ Gil Pil a7)
Cate (1964, p. 19, no. 33)
[For a discussion of this species see: Cate, 1964]

56. Bistolida (Blasicrura) teres teres (GMELIN, 1791)
(Plate 32, Figure 65)
Localities: 8, 24, 59, 89, 98, 108, 111, 142, 150, 159, 163,
176
(Zi USL 125) De 5)
Cate (1964, p. 20, no. 38)

[For a discussion of this species sce: Cate, 1964]

57. Bistolida (Derstolida) kienert kieneri (Hwatco, 1906)
(Plate 33, Figure 66)
Localities: 24, 176
This species is rare in West Australia, with only 3
specimens collected so far, to my knowledge. For this

Largest shell (176): 16.7 9.8 Lesa. oe lS
Medium shell (24): 14.3 8.5 Oe UG. al
Smallest shell (24): 14.2 8.3 66 15 14

reason statistical data of all three shells are given here.
The larger shell (176) was collected by Mrs. Molly
Gedling of Vlaming Head Light, North West Cape, at
Yardie Creek in September 1964. One of the other 2

Page 232

shells was collected at Gantheaume Point, Broome (24),
in 1956 by Archer Whitworth, Broome; and the last shell
was collected by Theodore Crake, Broome, at Entrance
Point, Broome, in 1964. This latter animal was found
living among Bistolida hirundo cameroni (IREDALE, 1939).

58. Bistolida (Derstolida) hirundo cameron
(IREDALE, 1939)
(Plate 33, Figure 67)

Localities: 15, 47, 108, 119, 153, 176
(GAS Oss hes 218) 115)
Cate (1964, p. 19, no. 34)

[For a discussion of this species see: Cate, 1964]

59. Bistolida (Derstolida) ursellus ursellus (GMELIN, 1791)
(Plate 33, Figure 68)

Locality: 24
(Ui GO Bo is 7
Cate (1964, p. 19, no. 35)

[For a discussion of this species sec: Cater, 1964]

60. Bistolida (Bistolida) stolida stolida (LinNakus, 1758)
(Plate 33, Figure 69)

Localities: 46, 60, 119, 159
(Zid 195 129) 2B. 18)
Cate (1964, p. 20, no. 36)

[For a discussion of this specics sec: Cate, 1964]

61. Bistolida (Bistolida) brcvidentata (SowrrBy, 1870)
(Plate 33, Figure 70)

Localities: 24, 119, 176

(ZA WBS) MONS) 15) 1133)

Cate (1964, p. 20, no. 37)

This species is clearly separable from Bistolida s. stolida
because of the total absence of the lateral marks at cach
quarter of the shell. In its dorsal aspect the central color
blotch is usually more conscrvative and rounder in ap-
pearance; the teeth are shorter and more rounded as well.
This taxon is here given full specific status since it
occupies the same habitat in West Australia as B. 5. stolida,
and therefore should not be regarded as subspccific to it.

62. Cribrarula' (Ovatipsa) chinensis whitworthi
(Cate, 1964)
(Plate 33, Figure 71)
Localities: 36, 53, 89, 163, 176
(G25 WY ion! WG 14s)
Cate (1964, p. 20, no. 39)

1 see SCHILDER & SCHILDER, 1967

THE VELIGER

Vol. 10; No. 3

A specimen was taken from a crayfish pot set on a coral
bottom in 65 fathoms, 10 miles west northwest of West
Wallabi Island, Abrolhos Islands by Martin van der Oord
of Geraldton in June 1964. Cate coll. no. C 2907.

63. Cribrarula (Cribrarula) cribraria fallax
(E. A. Smiru, 1881)
(Plate 34, Figure 72)

Localities: 36, 52, 66, 68, 91, 111, 118, 119, 144, 159, 163,

176, 180

(AS. UG 12s} 33.117)

Cate (1964, p. 21, no. 40)

A specimen was removed from a crayfish pot set at 23
fathoms 4 mile cast of the first island south of Little
North Island (91), Abrolhos Islands by Alec Gilbertson of
Geraldton, in June 1964. Cate coll. no. C 2943.

LITERATURE CITED

Cate, Crawrorp NEILL
1961. ~~ Rediscovery of Cypraca marginata Gasxon, 1848.
The Veliger, 3 (3): 76 - 78; plt. 14; figs. 1-4. (1 Jan. 1961)
1962. | Comparison of two rare cowrie species (Gastropoda)
The Veliger 5 (1) : 6 - 14; plts. 1-4; 2 text figs. (1 July 1962)
1964. | Western Australian cowries. The Veliger 7 (1):
7 - 29; pit. 5; 1 map (1 July 1964)
Hepiry, CHARLES
1916. A preliminary index of the Mollusca of Western Aus-
tralia. Journ. Proc. Roy. Soc. West Austral. 1 (1914-1915) :
1 - 246; 16 plts. pub. August 1915, volume dated: Perth 1916
Laws, HELENE
1966. Zoila marginata (Mollusca, Cypracidac) in South Aus-
tralia. Rec. So. Austral. Mus. 15 (2): 251 - 256; figs. 1-6
(23 December 1966)
ScHILDER, FRANZ ALFRED
1965. The geographical distribution of cowries (Mollusca:

Gastropoda) The Veliger 7 (3): 171 - 183; 2 maps
(1 January 1965)
1966. = Zoila venusta and ce pisema. Hawaiian Shell News

14 (8): 5; N.S. 78; fig’d.
1966. = Zola friendii (Grav) in Shark Bay.
News 14 (10): 4; N.S. 80; fig’d.
Seyitprr, Franz ALFRED «& Marta SCHILDER
1966. ‘The size of ninety-five thousand cowries.
8 (4): 208-215
Scuiper, Maria, & Franz ALFRED SCGHILDER
1967. Studies on East Australian cowries.
10 (2): 103-110; 9 tables
Weaver, CLIFTON STokES
1960. Hawaiian scientific expedition finds rare Western Aus-
tralian volutes. Hawaiian Shell News 8 (12: 1 and 3; N.S.

(June 1966)
Hawaiian Shell
(August 1966)

The Veliger
(1 April 1966)

The Veliger
(1 October 1967)

10; fig’d. (October 1960)
Wiison, Barry R. & Ray SUMMERS
1966. Variation in the Zoila fricndi [sic] (GRAY) species com-

plex in South-Western Australia. Journ. Malacol. Soc.
Austral. 9: 2 - 24; plts. 1-4; text figs. 1 - 4 (28 Jan. 1966)

Vol. 10; No. 3

THE VELIGER

Page 233

A Checklist of Intertidal Mollusks for Bahia Willard

and the Southwestern Portion of Bahia San Luis Gonzaga

State of Baja California, Mexico

BY

HELEN DUSHANE'

AND

GALE G. SPHON’

Conchological Club of Southern California, Los Angeles, California 90007

(Plate 35; 1 Map)

INTRODUCTION

Tue Bays oF WILLARD AND SAN Luis GoNzaGA are
located in the State of Baja California, Mexico, on the
Gulf of California (Lat. 29° 48’ N, Long. 114° 24’ W),
200 km south of the International Boundary. In time, with
improved road conditions, it is probable that this area
will become a favorite locality for collectors. For this
reason we consider it desirable to present a checklist of
the intertidal mollusks that have been found there.

This list is not intended to be all inclusive, but presents
a preliminary survey indicating the species obtained in a
partially explored area. It is based on the results of a joint
field trip made by the authors from February 1 to 6, 1966
during which 310 species were collected. Mrs. Faye B.
Howard collected mollusks at both Willard and Gonzaga
Bays during the month of May of 1957, 1958, 1959, and
1960. Her collection was made available to us. Dr. Homer
King and the junior author collected specimens in Janu-
ary 1960; a report of the specimens they obtained is
included here. Dr. James H. McLean collected on Febru-
ary 2, 1966 at Gonzaga Bay. His list of mollusks collected
is included in this paper.

™ 15012 El Soneto Drive, Whittier, California 90605
2 Malacologist, Santa Barbara Muscum of Natural History, Santa
Barbara, California 93105

The nudibranch fauna of the entire Panamic province
is very poorly known. Accordingly, a determined effort
was made to collect these animals and to transport them
alive to La Jolla, California, where they were given to
James R. Lance for photographing and _ identifying
(Lance, 1966).

The fauna of the Bays of Gonzaga and Willard has
not been as intensively collected as that of some of the
other areas in the Gulf of California, such as Punta
Penasco (Lowe, 1935), Guaymas (DUSHANE & PooRMAN,
1967), Puertecitos (DUSHANE, 1962), Bahfa de los An-
geles (McLean, 1961) and Isla Espiritu Santo (KEEN,
1964).

PREVIOUS COLLECTING in tHe BAYS
oF WILLARD ano GONZAGA

Historically, the first collectors in this area were the
aborigines who are reported to have worn “pearls, berries,
white round shells from small snails, and pieces of shell
and mother of pearl” (DuNnNeE, 1952). The earliest histo-
rian, Clavigero, 1788 (1937 transl.) , reporting on physical
conditions in Baja California prior to the expulsion of
the Jesuits in 1768, tells “Of the testaceous there are
mussels, murex shellfish, mother-of-pearl, and many other
kinds of periwinkles, mollusks and oysters.”

Coan (1965) reports on the shells of an Indian kitchen
midden with a table of mollusks from the midden. He

Page 234

shows a photograph of shell designs superimposed on the
sand dunes. However, one should be most careful in
reporting shell figures supposedly made by Indians. Usu-
ally the aborigines were too busy obtaining their daily
food to spend time making figures in the sand. The Jesuit
priests who were the first Caucasians to live in these
regions made no report of such designs (ENGELHARDT,
1929)

Three records of molluscan collecting in the vicinity of
Willard and San Luis Gonzaga Bays appear in the litera-
ture: (1) The 1921 expedition of the California Academy
of Sciences to the Gulf of California anchored in San Luis
Gonzaga Bay, dredged there and at San Luis Island to the
west. A general account of this expedition was given by
StEvin (1923). Mollusks taken during this investigation
have been partially recorded by BAKER, HANNA & STRONG
(1928): Pyramidellidae; Baker (1926): Triphoridae;
Baker & Hanna (1927): Opisthobranchiata; Baker,
Hanna & Stronc (1938): Cerithiopsidae, Cerithiidae
and Cyclostrematidae. (2) The Allan Hancock Pacific
Expeditions of 1937 and 1940 collected off Willard Point,
at Willard Island, and in Gonzaga Bay. Fraser (1943)
listed 8 dredging stations and one shore station, but the
mollusks remain largely unworked. However, 21 speci-
mens of Dentalium hancocki EMERSON were reported by
Emerson (1956) from one dredge haul in Gonzaga Bay.
(3) The 1957 expedition of the Puritan-American Muse-
um of Natural History reported 2 dredging stations, one
diving and one shore station within Gonzaga Bay. A general
account of this expedition was give by EMERSON (1958).

Shore collected specimens are the main concern of
this report, but several dredging records are worthy of
mention. Cantharus bilirata (REEVE) (fragments only),
Cancellaria obesa Sowersy, Clavus melea (Datt),
Daphnella allemani (BartscH), D. crebriforma (SHASKY
& CAMPBELL) and Trigonostoma campbelli SHasky were
dredged in 50m off Punta Final, January 1, 1961 by
Shasky, Campbell and Sphon (SHasxy, 1961). On the
south side of Willard Island in Gonzaga Bay, in 50m,
Anatina cyprinus (Woop) was taken by the same trio
(CamMpBELL, 1961). CampBett (1964) reports 3 speci-
mens of Terebra adairensis CAMPBELL from Gonzaga Bay.
DononveE (1966) reports 2 specimens of Titvia myrae
CampBELL dredged off Punta Final. The senior author of
this paper dredged specimens of Acmaea semirubida
Dati, Diodora pusilla Berry, Coralliophila incompta
Berry, Pleuroliria picta (ReEvE), Aspella baker: Hert-
LEIN & STRONG, Clavus aeginus DAL, Triphora hanna
Baker, 7. oweni Baker, and Metaxia convexa (CARPEN-
TER) off San Luis Island in October 1966 and December
1966 in 11 m.

THE VELIGER

Vol. 10; No. 3

OCEANOGRAPHIC CONSIDERATIONS

Very little is known concerning meteorological and oceano-
graphic conditions at Gonzaga and Willard Bays. No year
around water temperature data are available (RODEN &
Groves, 1959: p. 11), but local inhabitants claim the air
temperature rarely goes above 100° in the hot months of
July and August (personal communication Alfonsina
Urquidez V.). The lowest temperatures are about 40°F
Temperature ranges were corroborated by José Samano
Sanchez, Servicio Federal, Forestal y de la Fauno, who is
the conservation inspector for this area. Pacific storms
create violent winds at Gonzaga Bay, less so at the adjacent
Willard Bay. Gonzaga Bay is separated from Willard Bay
to the west by a sand spit extending at low tide to Willard
Island. This small island, together with the mountains to
the west, serve to protect Willard Bay from the wind.
Gonzaga Bay is an open indentation in a low lying valley,
exposed to winds. The floor of Gonzaga Bay is sandy with
many subtidal sand bars, while that of Willard Bay is
silty and muddy, with a vast expanse of mud flat at low
tide. At high tide a depth of 14 feet (ca. 4m) of water
joins Willard and Gonzaga Bays. The tidal range is about
4 to 5m. The sea floor outside the bays slopes gradually
to a depth of 400 fathoms (approximately 729 m). A
small archipelago of 6 islands stretches westward from
Gonzaga Bay, the largest and most southerly of which is
Isla San Luis, 5 miles offshore. Around these islands, known
as Las Islas Encantadas, there are swift currents. The cur-
rent on the west side of the chain runs counter to that on
the east side. Yet the main current of the Gulf of California
on the west shore runs south (Berry, 1954: p. 24; RopEN,
1958: pp. 24 and 33). These factors create a maelstrom
which probably is influential in mixing the waters within
Gonzaga and Willard Bays. Since extensive evaporation,
which increases salinity, occurs in the shallow protected
bays of Bahia Concepcion, Bahia San Felipe, and Bahia
Adair (RopEN & Groves, 1959: p. 16) one would expect
the same to be true at Bahias Willard and Gonzaga.

FAUNAL RELATIONSHIPS

This checklist records 310 species of mollusks, of which
102 are pelecypods, 196 are gastropods, and 12 are chitons.
Of this number 7 are doubtfully identified (“cf”) and
6 are identified only to genus. This report extends the
known range northward for specimens of 32 species
(range data from Kren, 1958, and McLean, 1961).
These are Barbatia lurida, B. gradata, Anadara cepoides,
Noetia reversa, Glycymeris tessellata, Mytella speciosa,
Lithophaga attenuata, Gregariella denticulata, Diplodon-

Vol. 10; No. 3

THE VELIGER

Page 235

ta caelata, Lasaea subviridis, Chama frondosa mexicana,
Pseudochama panamensis, Pitar consanguineus, Apoly-
metis cognata, Cyathodonta lucasana, C. undulata, Peta-
loconchus complicatus, P. flavescens, Sinum noyesi, Deci-
pifus gracilis, Anachis taeniata, Nassarina pammicra,
Engina reevei, Clathrodrillia maura, Crassispira aterrima,
C. monilifera, Mangelia finitima, M. subdiaphana, Tena-
turris carissima, Terebra albocincta, Acteocina carinata,
Pyramidella panamensis. The northernmost locality for
each of these is mentioned in the checklist. One species,
Lamellaria sharonae WiLuETT, 1939 (type locality: Ana-
heim Bay, Orange County, California) was taken on the
south side of Willard Island. Two southward extensions of
range in the Gulf of California are specimens of Terebra
berryi, reported by DuSHANE (1962) from Puertecitos,
and Melampus mousleyi (type locality, Bahia de Adair,
Sonora, Mexico).

Although the molluscan fauna is predominantly Pan-
amic, some members of the Californian province are
represented in the northern Gulf. PARKER (1963: p. 124)
presents a hypothesis attempting to explain the presence
of California province mollusks in the Gulf of California.
He considers it possible that the cold water fauna of
California entered the Gulf of California area during the
colder parts of the Pleistocene and during a time when
the sea level in the Gulf was at least 100m lower than at
present. The lowered sea level left a narrow continental
shelf along which mollusks might migrate northward into
the Gulf. Specimens of the following species occurring in
both the Californian and Panamic provinces are also to
be found at Gonzaga and Willard Bays: Chione californi-
ensis, Transennella tantilla, Sphenia fragilis, Polinices
reclusianus, Lamellaria sharonae, Hermissenda crassicor-
nis, Spurilla chromosoma, and Rostanga pulchra.

The faunal element restricted to the northern and north-
western shores of the Gulf is less well known but includes:
Acmaea strongiana, Nomaeopelta dalliana, Cantharus
macrospira, Turritella anactor, Terebra berryi,T. dushanae,
Recluzia palmer, and Acanthochitona exquisita. There are
also some peculiarities in faunal distribution on the western
side of the Gulf of California. Anachis varia and Parame-
taria duponti are to be found from San Felipe south to
Gonzaga Bay, but are not reported by McLean (1961)
from Los Angeles Bay. Specimens of Nassarina pammicra
reported by McLEAN (op. cit.) from Los Angeles Bay as
a range extension northward from Nicaragua have also
been collected by DUSHANE (1964) at Puertccitos as well
as at Gonzaga Bay. There are a few species which occur in
a very limited area such as Terebra berryi and T. dushanae
(type locality for both: Puertecitos). The former occurs
living at Gonzaga Bay but has not been found at San
Felipe to the north or at Los Angeles Bay to the south. The
latter species occurs uncommonly at Agua de Chale, 24

miles north of Puertecitos; one beach specimen has been
taken at Gonzaga Bay; it is unrecorded at Los Angeles Bay.
Strombina dorsata occurs at both Gonzaga Bay and at
Puertecitos but is unrecorded from Los Angeles Bay.
Mitra erythrogramma, usually taken by dredging, is fairly
common intertidally at Willard Bay.

A considerable number of species common to the
eastern shore of the Gulf of California is apparently absent
from Gonzaga and Willard Bays. These include the fol-
lowing species, which we have collected at cither Guay-
mas or Punta Penasco, or both: Astraea unguis, Turritella
gonostoma, Mitra lens, Agaronia testacea, Cypraea arabi-
cula, Thais kiosquiformis, Thais speciosa, Purpura patula
pansa, Conus brunneus, C’. purpurascens and C. virgatus.
These are essentially the same species McLEan (1961)
also reported to be absent from Los Angeles Bay.

SYSTEMATIC ACCOUNT

The following format is adopted:

1. The order in the checklist, the nomenclature, and the
specics number is that given by KEEN (1958). Re-
ferences to species listed by KEEN may be located in
her bibliography. References to species proposed since
1958 are included in the present paper.

2. The habitat and relative abundance of species found
at Bahias Gonzaga and Willard are given. The species
referred to as beach shells were not found living by us.
The bathymetric ranges given by Keen (1958) in-
dicate many of these species live in deeper water.

3. The collecting stations referred to in the list by num-
bers are shown on the accompanying Map.

4. Range extensions are indicated by asterisks (*) fol-
lowing the Keen numbers. The area from which the
range is extended follows the collector’s initial.

5. The following collectors are designated by initials:

JoscpheandiEvelensDuShane en D
Rayen Bun blow ancl ie stirs: ster, aire acse ames cn the, Baa H
FLOM Rain ous ce ee ak ed iene eit K
JarmeseRegleance metre ais Man etal Skee) eee aes 1G,
ames Mccann.) te eee ee M
Gale S phonics: tere antes teen hon JEON S

The specimens collected by Howard, King and Sphon
are in the Santa Barbara Museum of Natural History. The
DuShanes and Lance maintain private collections. Speci-
mens collected by McLean arc in the Los Angeles County
Museum of Natural History.

ACKNOWLEDGMENTS

We wish to express our gratitude to Drs. Robert Robertson
of the Academy of Natural Sciences, Philadelphia for

Page 236

making type comparisons; James H. McLean of the Los
Angeles County Museum of Natural History for critical
evaluation of the manuscript; Myra Keen of Stanford
University for suggestions and criticisms; Mr. Joseph
DuShane for his triple rdle as driver, fellow collector and
companion. Several people have contributed identifica-
tions: Mr. Peter Oringer, Caecidae; Dr. Donald Shasky,
Turridae; Mr. Allyn G. Smith, Polyplacophora; Mr.
James R. Lance, Nudibranchiata. To all these people we
are indebted. Credit for the map is due Mrs. Emily Reid
and for the photographs Dr. Leo G. Hertlein, Mr. Allyn
G. Smith, Mrs. Jean M. Cate, Mr. Perfecto Mary, and
the Los Angeles County Museum of Natural History.

ECOLOGICAL NOTES on
FOURTEEN COLLECTING AREAS

(see Map)

1. A low lying drainage channel bordered by Salicor-
nia: two species of Melampus are in the mud at the
roots of these plants.

2. A flat mud covered rock shelf : Crassispira nymphia
and C. pluto are abundant. Large specimens of
Lithophaga attenuata have bored into the rock.

3. Silty mud flats in the southern half of Willard Bay:
Laevicardium elatum, Oliva spp., Conus spp., Nassa-
rlus spp., Polinices spp., and Cassis ientiquadrata
are common. Heterodonax bimaculata is abundant in
the upper littoral zone.

4. Rocky shore line with turnable rocks: Barbatia,
Lima, Crucibulum, Murex, and Strombus are com-
mon.

5. Rocky shore in an exposed area: only the hardier
forms are found, such as Acmaea, chitons, and Acan-
thina. Specimens of Cypraea cervinetta have been
taken here. Coralliophila costata and Modulus dis-
culus are on the rocks.

6. The largest of a six-island archipelago. Limited
dredging here has corroborated the report of the
1921 expedition of the California Academy of
Sciences to Isla San Luis.

7. A sheltered cove interspersed with boulders and sandy
beaches: Pecten circularis and Lima tectrica are
under and around the rocks. Ostrea palmula is
attached to the rocks in the high intertidal zone.

8. The south side of Willard Island: this is a protected,
rocky shore line with small to medium sized rocks
broken by a few flat mud areas. The underside of
rocks is an excellent location for nudibranchs, chitons,
and Mitra solitaria. Pteropurpura erinaccoides and
Cymatium gibbosum cling to the rocks at low tide.

THE VELIGER

10.

il

12%

36

37

Vol. 10; No. 3

Sand spit connecting the shore of Gonzaga Bay to
Willard Bay and Willard Island: at low tide the spit
is exposed, making it possible to walk to the island.
At high tide a depth of 4m of water has been repor-
ted. The outgoing tides create shifting sand bars
where Terebra spp. and Conus ximenes are half
buried. This is also the area where Tivela stultorum
has been introduced (SHasky, 1961; DuSHane,
1966).

The south side of Willard Island, east of the sand
spit and fronting on Gonzaga Bay: a rocky precipitous
shore line with rough water. Opalia diadema, Tena-
turris carissima and Mitra solitaria are under rocks.
Acmaea turveri are attached to rocks in the upper
littoral, while A. strongiana and Nomaeopelta dalli-
ana are on the rocks in the mid-tide zone.

A steep beach which is actually a continuation of
the sand spit: not particularly productive, although
Laevicardium elatum and Dosinia ponderosa are
common.

A rock reef area with tide pools: the tide pools
occur in the upper intertidal zone. Crucibulum imbri-
catum, Anadara multicostata, and Haminoea strongi
are common.

. A rocky headland with medium size rocks: Cyp-

raea annettae, Arcopsis solida, and Isognomon chem-
nitzianus are under and attached to rocks.

. Asand bar area parallel to numbers 12 and 13: this is

the low intertidal zone with extensive sand bars.
Cassis centiquadrata, Oliva spicata, and Terebra va-
riegata are abundant.

PELECYPODA

Arca mutabilis (SowerBy, 1833). Common (5),
under rocks (H, K, S).

Arca pacifica (SowERBy, 1833). Uncommon (4, 5,
7), adhering to underside of rock ledge, low inter-
tidal zone (H, K).

38* Barbatia lurida (SowerBy, 1833). Uncommon (10),

39

attached to underside of rocks; Espiritu Santo Is-
land, Gulf of California (K, M).

Barbatia baily: (Bartscu, 1931). Rare (10), beach
specimen (H).

40* Barbatia gradata (BRopERiIP & SowErRBY, 1829). Un-

43

44

common (5), attached to underside of rocks;
Scammon’s Lagoon, Lower California to Peru (K).
Barbatia reeveana (Orpicny, 1846). Common (4,
5, 7, 8), attached between rocks (D, H, K, M, S).
Barbatia illota (SowrRBy, 1833). Uncommon (4, 8,
10), attached to rocks (D, H, K).

Vol. 10; No. 3 THE VELIGER Page 237

SS

Isla San Luis

Bahia San Luis Gonzaga

Punta Final

MAP SHOWING LOCALITIES

Numbers refer to Collecting Stations
SRECH — Rocky Areas
1 — Mud Reef
=") — Sand Flats and Bars

— Landing Field for Small Aircraft
— Simple Accommodations for Eating and Sleeping

Page 238

46

57

60*

69*

72

74

75

76

dg

83

86*

88

90

Sis

95

S97

101

107

111

115

124

126

Arcopsis solida (SowERBY, 1833). Common (5, 13),
attached to underside of rocks (D, H, K, M, S).
Anadara multicostata (SowERBY, 1833). Common
(4, 12, 13), between rocks in low intertidal zone
(D, K, S).

Anadara cepoides (Reeve, 1844). Rare, beach spe-
cimen; Cerralvo Island, Gulf of California to
Panama (H).

Noetia reversa (SowERBy, 1833). Rare (3); beach
specimen; Concepcion Bay, Gulf of California to
Peru (K).

Glycymeris bicolor (REEvE, 1843). Rare (3, 10),
lying free in sandy runnels (D, H, M).

Glycymeris gigantea (REEvE, 1843). Rare alive,
beach valves common (3), partially buried on silty
mud flats (H, K, S).

Glycymeris maculata (BropEriP, 1832). Common
(3, 9, 11, 14), partially buried on sand bars (D, H,
K, M, S).

Glycymeris multicostata (SOwERBY, 1833). Uncom-
mon (3), partially buried on sand flats (H, K).
Glycymeris tessellata (SowERBY, 1833). Rare (3),
partially buried on silty mud flats (S).

Hormomya adamsiana (DunkeER, 1857). Common
(7, 8), attached to underside of rocks (H, S).
Mytella speciosa (Rerve, 1857). Rare (11), buried
in sandy runnel; Magdalena Bay to Peru (D).
Crenella divaricata (OrsIGNY, 1853). Rare; beach
specimen (H).

Lithophaga aristata (Dittwyn, 1817). Common
(5), boring in rock (H, K).

Lithophaga attenuata (DesuayeEs, 1836). Common
(2, 5), boring in mud-covered rock ledge; Costa
Rica to Chile (D, K, S).

Lithophaga spatiosa (CARPENTER, 1856). Common
(2), boring in mud-covered rock ledge (S).
Gregariella denticulata (Datu, 1871). Common
(8), boring in soft rock; Acapulco, Mexico (H).
Modiolus capax (Conrap, 1837). Common (2), on
mud-covered rock ledge (S).

Pteria sterna (GouLp, 1851). Rare; beach specimen
(H).

Pinna rugosa Sowersy, 1855. Uncommon (4, 5),
among rocks, low intertidal zone (K).

Isognomon chemnitzianus (OrpicNy, 1853). Com-
mon (7, 8, 10, 13), attached to underside of rocks
(H, K, M, S).

Ostrea palmula CarPENTER, 1857. Common (4, 7),
attached to rocks in the high intertidal zone (K, S).
Pecten vogdesi ArNoLD, 1906. Rare living, beach
valves common (3), on sand and gravel flats (D, H).

THE VELIGER

128

132

137

140

141

146

147

152

162

163

186

200

205,

206

210

22

213

214

217

Done

236

238

Vol. 10; No. 3

Aequipecten palmeri (Dat, 1897). Rare; beach
specimen (H).

Aequipecten circularis (SOwERBY, 1835). Uncom-
mon (5, 7, 8), around and under rocks intertidally
(D, H, K, S).

Lyropecten subnodosus (SowERBY, 1835). Rare (5,
7), among small rocks (H, K, S).

Lima tetrica Goutp, 1851. Rare (4, 7), under rocks
(D, H, K).

Lima hemphilli HERTLEIN & STRONG, 1946. Rare
(4, 7), under rocks (H, K).

Spondylus calcifer CARPENTER, 1857. Uncommon
(5), on rocks (K).

Plicatula anomioides KEEN, 1958. Uncommon (5),
under rocks (H, K).

Anomia peruviana Orpicny, 1846. Rare (4), on
rocks (K).

Cardita affinis SowERBY, 1833. Common (3, 4, 10),
under rocks and on silty mud flats (D, H, K, M, S).
Cardita crassicostata (SowERBy, 1825). Rare (7,
8), among rocks on Willard Island, intertidally (D,
K, S).

Lucina lingualis CarpENTER, 1864. Rare; valves
only (H).

Codakia distinguenda (Tryon, 1872). Uncommon
(5), in very shallow water at low tide (H, K).
Ctena mexicana (Dati, 1901). Common (14), ly-
ing on sand bars (H, S). 3
Dwwalinga eburnea (Reeve, 1850). Uncommon
(14), partially buried on sand bars (H).
Diplodonta subquadrata CarPENTER, 1856. Com-
mon (10), among rocks (M, S).

Diplodonta sericata (REEVE, 1850). Common (3),
on mud flats at edge of sting ray basins (D, H, S).
Diplodonta caelata (Reeve, 1850). Rare (2), in
pholad holes (H).

Diplodonta semirugosa Dati, 1899. Rare (3), on
mud flats (D, S).

Aligena nucea Datu, 1913. Rare; single worn valve
(H).

Lasaea subviridis Dati, 1899. Rare; in the coils of
a vermetid; Shelter Cove, California to Cape San
Lucas, Lower California (H).

Solecardia eburnea Conran, 1849. Uncommon (14),
on sand bars (D).

Chama buddiana C. B. Apams, 1852. Common (4,
5, 7), attached to rocks (H, K, S).

240a*Chama frondosa mexicana CARPENTER, 1857. Un-

243

common (2), on mud covered rock reef; Guaymas
to Panama and Ecuador (D, H, S).

Chama venosa ReEve, 1847. Uncommon (8), on
rocks (H).

Vol. 10; No. 3 THE VELIGER Page 239
247* Pseudochama panamensis (REEVE, 1847). Uncom- 347 Petricola denticulata Sowrrsy, 1834. Common (2),
mon (5), on rocks; Panama (H, K). boring in soft, rocky ledges (D, H, K).
248 Pseudochama saavedrai HERTLEIN & STRONG, 1946. 355 Mactra dolabriformis (Conran, 1867). Uncommon
Uncommon (4), on rocks (K). (14), partially buried on sand bars (S).
252 Trachycardiwm panamense (Sowersy,1833).Com- 408 Tellina pristiphora Dau, 1900. Rare; beach speci-
mon (3, 14), on silty mud flats and sand bars (D, H, mens (K).
K, S). 422* Apolymetis cognata (Pitspry & VANATTA, 1902).
255 Papyridea aspersa (SOWERBY, 1833). Common (3), Rare; beach specimens; Nicaragua (H).
on silty mud flats (D, H, K, S). 423 Apolymetis cognata clarki DurHAM, 1950. Rare;
256 ‘Trigoniocardia granifera (BRODERIP & SOWERBY, beach specimens (H).
ed er ( is [pene y lat elo tats aes (0S) 450 Donax gracilis Hantey, 1845. Common (14), par-
258 Trigoniocardia biangulata (BRODERIP & SOWERBY, : :
& tially buried on sand bars (D, S).
1829). Common (3, 11, 12, 13), on sand flats and 455 D :
onax punctatostriatus HANLEY, 1843. Uncommon
along sand beach (D, H, K, S). 14 b _
262 Laevicardium elatum (Sowersy, 1833). Common (14), on sand bars (H).
(3, 10, 11 12, 13, 14), rolling in on low tide and 466 Gari regularis (CARPENTER, 1864). Uncommon
partially buried in sand (D, H, K, M, S). (11), sand beach (D, H, S).
263 Laevicardium elenense (SowERBy, 1840 [? 1841]). 467 Heterodonax bimaculatus (LinnaEus, 1758). Com-
Common (3, 14), on silty mud flats and on sand mon (3, 10, 12, 13), high tide zone, sand beach (D,
bars (D, H, K, S). H, K, M, S).
280 Transennella puella (CARPENTER, 1864). Uncom- 4/1 Tagelus californianus (Conran, 1837). Uncommon
mon (8, 9, 10), sand flats (M, S). (3), silty mud flats (K).
282 Transennella tantilla (Goutp, 1853). Uncommon; 475 Tagelus politus (CARPENTER, 1857). Rare (3), on
under rocks (S). silty mud flats (D, H).
284* Pitar consanguineus (C.B.ApaAms, 1852). Rare; 477 Semele bicolor (C. B. ApaMs, 1852). Rare; beach
valves only; Port Guatulco, Mexico (H). specimen (H).
286 Pitar newcombianus (Gass, 1865). Uncommon 481 Semele flavescens (Goutp, 1851). Uncommon (10),
(3), on silty mud flats (H). among rocks (H, M).
303 Megapitaria squalida (Sowersy, 1835). Common 483 Semele guaymasensis Pirspry & Lowe, 1932. Rare
(3, 9, 10, 13), partially buried on sand bars (D, H, (8), around rocks (S).
K, M, S). 489 Semele pacifica Datx, 1915. Rare; beach specimens
305 Dosinia dunkeri (Purviprr, 1844). Uncommon (14), (H).
partially buried in sand bars (H, K). 508 Cumingia lamellosa Sowrrsy, 1833. Uncommon (6,
306 Dosinia ponderosa (Gray, 1838). Common (14), 12, 13), among rocks (D, H, S).
partially buried hi sand bars (D, H, K, 8). 512 Solen mexicanus Datt, 1899. Uncommon (3), silty
318 Chione californiensis (Bropertp, 1835). Common mud flats (H).
(3,72) ,onvsilty mud fatsand sand bars)(D,M). 521 Sphenia fragilis (H. « A. AvaMs, 1854) (teste
319 Chione compta (Broperip, 1835). Uncommon (3), Keen). Rare (8), under rocks (S).
penteely paricd on aes se eel otal sD) 523 Corbula bicarinata SowErBy, 1833. Uncommon (7),
320 Chione guatulcoensis HERTLEIN & STRONG, 1948.
Uncommon; on silty mud flats (D). mae woes (D),9).
391 Chione oraitleivalie (Sowsrsy, 1835), Uncommon 542 Hnatella arctica (LINNAEUuS, 1767). Common (4, 5,
(11, 12), on sand beach (D, H). 7, 8), under rocks (H, K, S). :
328 Chione purpurissata Daur, 1902, Rare (9, 10), °/8 Zyonsia gouldi Datt, 1915. Rare (9), partially
partially buried on sand beach (D, M). buried on sand spit (H).
335 Chione picta Wiett, 1944. Uncommon (8), a- 590 Thracia squamosa CARPENTER, 1856. Rare (2), in
mong rocks (S). pholad holes (H).
337 Anomalocardia tumens (VeERRILL, 1870). Rare (9, 993 Cyathodonta dubiosa Dati, 1915. Rare; beach spe-
11), on sand beach (D, H). cimen (H).
340 Protothaca grata (Say, 1831). Common (3), silty 594 Cyathodonta lucasana Dati, 1915. Rare; beach
mud flats (K). specimen (H).
341 Protothaca asperrima (SowersBy, 1835). Uncom- 595 Cyathodonta undulata Conran, 1849. Rare; beach

mon (3), silty mud flats (S).

specimen (H).

Page 240

GASTROPODA

10 Acmaea semirubida Datr, 1914. Rare (3), inter-
tidally, attached to dead Cardita crassicostata
(SowErsy, 1825) ; dredged 11m, common (6), (D).

12a Acmaea strongiana HERTLEIN, 1958. Common (8,
10), on rocks at midtide zone (D, H, M, S).

14 Acmaea turveri HERTLEIN & STRONG, 1951. Com-
mon (8), on rocks in high intertidal zone (D, H, S).

16 Nomaeopelta dalliana (Pitspry, 1891). Common
(7, 8), on rocks at midtide zone (D, H, K, S).

18 Nomaeopelta stanfordiana (Berry, 1957). Rare (8,
10), beach specimens (H, M).

Lucapinella milleri Berry, 1959 (p. 109). Uncom-
mon (8), attached to underside of rocks (D, H).

30 Diodora alta (C. B. AbAMs, 1852). Uncommon (7,
8), under rocks (D, H, S).

32 Diodora inaequalis (SowERBy, 1835). Common (5,
7, 8, 10), under rocks (D, H, K, M, S).

32a Diodora digueti Masitte, 1895. Uncommon (5, 7,
8), under rocks (D).

34 Diodora saturnalis (CARPENTER, 1864). Rare; beach
specimen (H).

Diodora pusilla BERRY, 1959 (p. 109).Common (6),
dredged 7 fms. (D). a

38 Calliostoma cf. C. angelenum Lowe, 1935. Rare;

beach specimen (H).

THE VELIGER

Vol. 10; No. 3

Tegula sp. Under rocks (8, 10), common (H, M,S).

54 Tegula mariana Datu, 1919. Common (8, 10),
under and around rocks (H, M).

58 Tegula rugosa (A. Apas, 1853). Common (5, 7,
8, 10, 12, 13), on rocks (D, H, K, M, S).

64 Turbo fluctuosus Woop, 1828. Uncommon (8, 10),
among rocks (H, K, M).

68 Turbo squamiger Reeve, 1843. Uncommon (6),
dredged in 11 m (D).
Tricolia typica (DALL, 1908).Common (6), dredged
in 11m (D).

81 Nerita scabricosta LAMARCK, 1822. Rare (5), on
rocks (H).

82 Nerita funiculata Menke, 1851. Common (5, 8,
10), on rocks in high tide area (H, M, S).

83 Neritina luteofasciata Miter, 1879. Uncommon
(3), in runnels on silty mud flat (H).

87 Niso excolpa Bartscu, 1917. Beach specimen (D).

90 Epitonium acapulcanum Dati, 1917. Uncommon
(7), on Cardita crassicostata (SowERBY, 1825) (D).

124 Epitonium oerstedianum HERTLEIN & STRONG, 1951.
Beach specimen (H).

158 Opalia diadema (Sowersy, 1832). Uncommon
(10),under rocks on west side of Willard Island
(D, H).

175a Littorina dubiosa penicillata CARPENTER, 1864.
Common (7), on rocks (H).

Explanation of Plate 35

Figure 1a, 1b: Cyclostrema spicert BAKER, HANNA & STRONG, 1938.
Holotype, California Academy of Sciences Paleo. Type Collection
5462 (x 15.5). Greatest diameter 3.97mm; least diam. 2.93 mm;
alt. 2.2mm. Type locality: Coyote Bay, Concepcion Bay, Lower
California, Mexico, in about 2 fathoms.

Figure 2: Terebra berryi CAMPBELL, 1961. Holotype: Calif. Acad.
Sci. Paleo. Type Coll. 12352 (x2). Length 32mm; width 7mm.
Type locality: Puertecitos, Lower California, Mexico.

Figure 3: Triphora hannai BAKER, 1926. Holotype, Calif. Acad.
Sci. Paleo. Type Coll. 2136 (x10). Length 8.17mm; diam. 2.43
mm. Type locality: San Francisco Island, Gulf of California, Mexico.
Figure 4: Lamellaria sharonae WittetTT, 1939. Holotype: Los
Angeles County Mus. Nat. Hist. 1059 (x 3.5). Max. diam. 5.5mm;
alt. 7.4mm. Type locality: Anaheim Bay, Orange County, Cali-
fornia.

Figure 5: Diodora pusilla Berry, 1959. Holotype: Stanford
Univ. Paleo. Type Coll. 8587 (x3). Long. 3.8mm; lat. 2.5mm;
alt. 3.6mm. Type locality: 6-10 fathoms, off Acapulco, Guerrero,
Mexico.

Figure 6: Retusa gonzagensis BAKER & HANNA, 1927. Holotype:
Calif. Acad. Sci. Paleo. Type Coll. 2519 (x9). Length 2.9mm; diam.
1.6mm. Type locality: San Luis Gonzaga Bay, Gulf of California,
Mexico.

Figure 7: Lucapinella milleri Berry, 1959. Holotype: Stanford
Univ. Paleo Type Coll. 8588 (x3.5). Long. 8.6mm; lat. 4.6mm;
alt. 1.4mm; diam. of foramen at center 1.3mm. Type locality:
Puertecitos, Lower California, Mexico.

Figure 8: Melampus mousleyi Berry, 1964. Holotype: Stanford
Univ. Paleo. Type Coll. 9503 (x3). Alt. 1omm; diam. 5.4mm.
Type locality: Cholla Cove, Bahia de Adair, Sonora, Mexico.
Figure 9a, gb: Aspella bakeri HERTLEIN & STRONG, 1951. Holo-
type: Calif. Acad. Sci. Paleo. Type Coll. 5893 (x 1.3). Length
18mm; diam. 8mm. Type locality: Agua Verde Bay, Gulf of
California, Mexico.

Figure 10: Epitonium oerstedianum HERTLEIN & STRONG, 1951.
Holotype: Calif. Acad. Sci. Paleo Type Coll. 9622 (x8). Length
6.5mm; diam. 4.2mm. Type locality: Off San Domingo Point,
Santa Inez Bay, Gulf of California, Mexico.

Figure 11: Coralliophila incompta Berry. 1960. DuShane Coll.
Length 23mm; diam. 14mm. Holotype: Berry Coll. 18 768;
length 33.5mm; diam. 20.3mm. Type locality: Puerto Refugio, —
Angel de la Guarda Island, Gulf of California, Mexico. ee
Photo Credits: Figures 1a, 1b, 2, 3, 6, ga, gb, 10 — California
Academy of Sciences. Figure 4 - Mr. Allyn G. Smith, California
Academy of Sciences. Figures 5, 7 — Los Angeles County Museum
of Natural History. Figure 8 - Perfecto Mary, Stanford Univer-
sity. Figure 11 -— Mrs. Jean M. Cate, Los Angeles.

Tue VELIGER, Vol. 10, No. 3 [DuSHANE & SpHON] Plate 35

Figure 4 Figure 5

Figure 9 a Figure 10 Figure 11

Vol. 10; No. 3

* Cyclostrema spiceri BAKER, HANNA & STRONG,

198

99

1938 (p. 234). Beach specimen (10) ; Concepcion
Bay, Gulf of California (M).

Truncatella bairdiana (C. B. Avams, 1852) (p.
437). Uncommon (8, 10), under rocks on west side
of Willard Island (D, H, M, S).

Rissoina burragei BartscuH, 1915 (p. 28), Uncom-
mon (8, 10), under rocks; Los Angeles Bay, Gulf
of California (McLgan, 1961), (D, M,S).
Rissoina barthelowi Bartscu, 1915 (p. 28).
Dredged in 11m (6); Concepcion Bay, Gulf of
California and Guaymas, Mexico (DUSHANE &
PoorMAN, 1967), (D).

Rissoina stricta (MENKE, 1850) (p. 161). Dredged
in 11m (6) ; Guaymas, Mexico (DUSHANE & Poor-
MAN, 1967), (D).

Rissoina zeltneri DEFoLIN, 1867. Rare, Beach speci-
mens; Panama (H).

Alvania monserratensis BAKER, HANNA & STRONG,
1930 (p. 27). Beach specimens; Monserrate Island,
Gulf of California (H).

Barleeia polychroma (pEFoutn, 1870). Rare (8),
under rocks; Panama (H).

Turritella leucostoma VALENCIENNES, 1832. Rare
(11), beach specimens (H).

Vermicularia pellucida eburnea (REEVE, 1842).
Common (7,8, 10,11, 12,13),under rocks (D, H,S)
Heliacus bicanaliculatus (VALENCIENNES, 1832).
Uncommon (3, 5), on muddy sand reef and among
compound ascidians in low intertidal zone (D, H,
K).

Caecum sp. Rare, attached to live Cardita crassicos-
tata (SoweErRBy, 1825), (D).

Elephantulum liratocincta (CARPENTER, 1857) (p.
317). Uncommon (8), in sand; Panama (D, S).
Elephantanellum heptagonum (CARPENTER, 1857)
(p. 319). Uncommon (8), in sand; Bahia de Los
Angeles, Gulf of California (D, S).

Modulus cerodes (A. Apams, 1851). Rare (3, 8,
12), on silty mud flats and among rocks (D, H, K).
Modulus disculus (Puripr1, 1846). Common (3, 5,
8, 12, 13), on silty mud flats and among rocks (D,
H, K, M, S).

Vermetus centiquadrus VALENCIENNES, 1846 (teste
Keren). Uncommon (8), under rocks (H).
Petaloconchus cf. P complicatus Dati, 1908. Com-
mon (5), on rock; Cocos Island, Panama (H).

200a*Petaloconchus flavescens (CARPENTER, 1857). Com-

201

mon (5), on rocks; Guaymas to Mazatlan, Mexico
(Jal).

Vermetus indentatus (CARPENTER, 1856), (teste
KEEN). Common (5), on rocks (H).

203

204

208

2A

215

216

THE VELIGER

*

220

224

227

229

233

240

241

*

Page 241

Petaloconchus macrophragma CARPENTER, 1865.
Beach specimen (H).

Serpulorbis cf. S. eruciformis (Moércu, 1862). Rare
(8), on rocks (H).

Vermetus tripsycha Pitssry & Lowe, 1932. Uncom-
mon (8), on rocks (H).

Cerithium sculptum Sowersy, 1855. Common (4,
5, 10), around rocks (H, K, M).

Cerithium stercusmuscarum VALENCIENNES, 1833.
Common (10), around rocks (H).

Cerithium uncinatum (GMELIN, 1791). Rare (8),
around rocks in low intertidal zone (D).

Triphora hannai Baker, 1926 (p. 225). Dredged in
11m (6) ; San Francisco Island, Gulf of California
(D).

Triphora owent Baker, 1926 (p. 232). Dredged in
11m (6); San Francisco Island, Gulf of California
(D).

Triphora panamensis (BartscH, 1907) (p. 249).
Rare (8), under rocks; Panama (S).

Metaxia convexa (CARPENTER, 1857) (p. 444).
Dredged in 11m (6), (D).

Seila assimilata (C. B. Apams, 1852) (p. 374). Un-
common (8), under rocks (H, S).

Alaba interruptelineata Pitspry & Lowe, 1932 (p.
81).Uncommon (8,10),under rocks (see McLEAN
1961; DuSHANE 1962), (D, M, S).

Alaba supralirata CARPENTER, 1857 (p. 366). Rare
(8), under rocks; Bahia de Los Angeles, Gulf of
California (H).

Alabina diomedeae (Bartscu, 1911) (p. 413). Un-
common (7, 10), under rocks (D, M, S).

Alabina monicensis (BARTSCH, 1911) (p. 409). Un-
common (7, 10), under rocks; Santa Monica, Cali-
fornia (S).

Alabina tenuisculpta (CARPENTER, 1864) (p. 517).
Uncommon (7, 10), under rocks (S).

Cerithidea mazatlanica CARPENTER, 1857. Common
(1), under Salicornia bushes on mud flats (D, H, K,
M, S).

Rhinocoryne humboldti (VALENCIENNES, 1832).
Rare, beach specimens (H).

Hipponix pilosus (DESHAYES, 1832). Uncommon
(5), on rocks (H).

Hipponix serratus CARPENTER, 1857. Rare (10),
beach specimens (H, M).

Calyptraca mamullaris Bropverip, 1834. Uncommon
(3), beach specimens (H).

Crepidula arenata (BropErip, 1834). Common (8),
attached to rocks (H, S).

Crepidula excavata (Broperip, 1834). Common
(7,8), attached to rocks and shells (S).

Page 242

242

243

245

248

252

254

258

266

272

274

275

276*

Crepidula incurva (Broperip, 1834). Common (3,
10, 13), living on other shells (H, S).

Crepidula lessonii (Broperip, 1834). Uncommon
(8, 12), under rocks (S).

Crepidula onyx SoweRBy, 1824. Common (3, 5,
10, 13), on beach shells (H, K, M, S).

Crepidula striolata Menke, 1851. Common (5),
living on other shells (H, K).

Crucibulum scutellatum (Woop, 1828). Common
(3, 4, 12, 13), attached to rocks (D, H, K, S).
Crucibulum spinosum (SoweErBy, 1824). Common
(4,7, 10, 12,13), attached to rocks (D,H,K,M,S).
Natica chemnitzii PFEIFFER, 1840. Common (3, 12,
14), on silty mud flats and sand bars (D, H, K, S).
Polinices bifasciatus (Gray, 1834). Common (3, 10,
14), on silty mud flats and sand bars (D, H, K, M,
S).

Polinices uber (VALENCIENNES, 1832) Common (3,
10, 14), on silty mud flats and sand bars (D, H, K,
M, S).

Polinices reclusianus (DEsHAYES, 1839). Common
(3, 4, 9, 14), on silty mud flats and sand bars (D,
K, S).

Sinum debile (Goutp, 1853). Rare (3), partially
buried on silty mud flat (S).

Sinum noyestt DALL, 1903. Rare (3), partially bur-
ied on silty mud flat; Nicaragua to Panama (S).
Lamellaria inflata (C. B. ApAms, 1852). Common
(8), under rocks (D, S).
Lamellaria sharonae WiueETT, 1939 (p. 123). Rare
(8), under rocks; Anaheim Bay, Orange County,
California (S). Original spelling L. “sharoni”’ emen-
ded by Burcu (1946) to L. sharonae.

Cypraea cervinetta KiENER, 1843. Under rocks (5),
[collected by Phillips, Santa Barbara, November,
1966].

Cypraea annettae Dati, 1909. Common (4, 5, 7,
8, 13), attached to underside of rocks (D, H, K, S).
Erato columbella MrenKeE, 1847. Uncommon (8,
10), attached to underside of rocks (D, H, M, S).
Trivia californiana (Gray, 1828). Uncommon (5,
7, 8), under and on rocks (H, K, S).

Trivia radians (LAMarcK, 1810). Beach specimens
(5, 12), (K).

Trivia solandri (SowErBy, 1832). Uncommon (5, 7,
8), under and on rocks (H, K, S).

Jenneria pustulata (Licutroot, 1786). Uncommon
(5, 8), under rocks (H, K, 8).

Strombus galeatus Swainson, 1823. Uncommon
(4, 8), in sand next to rocks (H, K, S).

Strombus gracilior SowerBy, 1825. Uncommon (3,
4, 11, 12), partially buried in sand (D, H, K, S).

THE VELIGER

309

315

324

335

339

344

348

37m

387

388

398

404

409

411

412

424

433%

Vol. 10; No. 3

Strombus granulatus Swainson, 1822. Common (3,
4, 7, 10), on silty mud flats and around rocks (D, H,
K, M, S).
Cassis centiquadrata (VALENCIENNES, 1832). Un-
common (3, 14), on silty mud flats and sand bars
(D, H, K, S).
Cymatium gibbosum (Bropverip, 1833). Uncommon
in February, common in late spring and early sum-
mer (4, 5, 7, 8, 12, 13), on rocks (D, H, K, S).
Murex elenensis DAL, 1909. Uncommon (3, 4, 5),
on rocks (H, K, S).
Hexaplex erythrostomus (Swatnson, 1831). Un-
common in February, common in May (3, 4, 5),
on silty mud flats and around rocks (D, H, K, S).
Muricanthus nigritus (Puiviprr, 1845). Uncommon
in February, common in May (3, 5), on tidal flats
and among rocks (D, H, K, M, S).
Pteropurpura erinaceoides (VALENCIENNES, 1832).
[see Emerson, 1964, p. 5]. Common (5, 7, 8, 12,
13), on top and sides of rocks (D, H, K, S).
Aspella bakert HERTLEIN & STRONG, 1951. Dredged
in 11m (6); Agua Verde Bay, Gulf of California
(D).
Coralliophila costata (BLAINVILLE, 1832). Uncom-
mon (2, 5, 12, 13), on top of rocks (D, K).
Coralliophila hindsu (CARPENTER, 1857). Rare (5),
on rocks (H).
Coralliophila incompta Berry, 1960. Uncommon
(6); 11m (D). .
Thais biserialis (BLAINVILLE, 1832). Common (4,
5, 7503 12.013)- on rocks) (Ds Ee s)e
Acanthina angelica 1. OtpRoyp, 1918. Common (4,
5, 7, 8, 10, 12, 13), on rocks at high tide level (D,
K, M, S).
Acanthina tuberculata (SowERBY, 1835). Common
(4, 5, 7, 8, 10, 12, 13), on rocks (D, H, K, M, S).
Morula ferruginosa (REEVE, 1846). Common (7,
8, 10, 12, 13), intertidally under rocks (D, H, M,
S).
Morula lugubris (C. B. ApaMs, 1852). Uncommon
(8), intertidally under rocks (H).
Decipifus gracilis McLean, 1959. Uncommon (8),
under rocks; Bahia de los Angeles, Baja California
(D, S).
ae spec. nov. Uncommon (8), under rocks
D, S).
a coronata (SowerBy, 1832). Uncommon
(5, 8, 10, 13), under rocks; dredged in 11m, com-
mon (6), (D, H, K, M, S).
Anachis cf. A. taeniata Pupp, 1846. Rare, beach
specimens; west Mexican coast (H).

Vol.

10; No. 3

THE VELIGER

Page 243

437

444

464

478

482

484

486

490*

494

508

512

515

543

549

Anachis hilli Piuspry & Lowe, 1932. Rare (8),
under rocks on the west side of Willard Island (D).
Anachis nigricans (Sowersy, 1844). Uncommon
(5, 8), under rocks (H, K).

Anachis varia (SoweErBy, 1832). Uncommon (5,
8, 13), under rocks (D, H, K).

Mitrella dorma BakrErR, HANNA & StrRONG, 1938.
Uncommon (8), intertidally under rocks (D, H).
Mitrella lalage Pitssry « Lowe, 1932. Uncommon
(8), intertidally under rocks (S).

Mitrella guttata (SowERBy, 1832). [see Howarp,
1963]. Common (4, 5, 7, 8), intertidally under
rocks (D, H, M).

Mitrella santabarbarensis (CARPENTER, in GOULD &
CarpPENTER, 1857). Rare (10), intertidally under
rocks (M).

Nassarina pammicra Pitspry & Lowe, 1932. Un-
common (8), clustered under rocks at sides of
drainage channels; Bahia de Los Angeles, Gulf of
California (D, S).

Parametaria duponti (KieNeER, 1849-1850). Un-
common (5, 8), on rocks (H, K).

Pyrene aureomexicana Howarp, 1963. Common (8,
10, 12, 13), under rocks (D, H, M, S).

Strombina dorsata (SowERBY, 1832). Rare (2), on
sandy mud reef (D, S).

Strombina gibberula (SowErRBy, 1832). Rare (2),
on mud covered rock reef (S).

Strombina maculosa (SowERBy, 1832). Common
(3, 4, 10), on silty sand flats (D, H, K, M, S).
Cantharus macrospira (Berry, 1957) (?) as “So-
lenosteira anomala.’ Common (3), on silty sand
flats (D, H, K, S).

Engina maura (Sowersy, 1832). Rare, beach spe-
cimen (H).

550a*Engina reevet Tryon, 1883. Rare (8), on rocks;

551

567

587

595

610

612

618

Southern Gulf of California, possibly south to Pan-
ama (H).

Engina solida (Dat, 1917). Rare, beach specimen
(D).

Melongena patula (BRopERIP & SOWERBY, 1829).
Rare (3), juvenile on silty mud flats (S).
Nassarius versicolor (C. B. ApAMs, 1852). Common
(10, 14), on sand bars (D, H, M, S).

Nassarius tiarula (KieNER, 1841). Common (10,
14), on sand bars (D, H, M, S).

Fusinus dupetitthouars: (KiENER, 1846). Rare,
beach specimen (H).

Fusinus ambustus (Goutp, 1853). Common (2, 7,
8), on mud covered rock ledge (D, H, S).

Fusinus felipensis Lowe, 1935. Uncommon (8, 10,
13), under rocks (D, M, S).

666

669

807

818*

Olwa incrassata (Licurroor, 1786). Common (3,
4, 14), partially buricd in sand (D, H, K, S).
Oliva spicata (R6p1NG, 1798). Common (3, 4, 14),
partially buried in sand (D, H, K, S).

Oliva undatella Lamarck, 1810. Common (11, 12,
13, 14), partially buried in sand bars (D, S).
Olivella dama (Woop, 1828, ex Mawe MS.).
Common (9, 10, 11, 12, 13, 14), partially buried
in sand bars (D, H, K, M, S).

Olivella fletcherae Berry, 1958. Common (11, 12,
13, 14), partially buried in sand bars (D, K, M, S).
Olwella zanoeta (Ductos, 1835). Uncommon (3,
12, 13), partially buried in sand flats (K).

Mitra solitaria C. B. Apams, 1858. Uncommon (8,
10), under rocks (D, H, M, S).

Mitra tristis BropErip, 1836. Common (2,5, 12,13),
on mud covered rock reef (D, H, K, M, S).
Mitra erythrogramma Tomutn, 1931. Uncommon
(2), partially buried on sandy, mud covered rock
ledge (D, S).

Lyra cumingu (Broperip, 1832). Uncommon (3,
4, 12, 13), on mud covered rock shelf (D, H, K,
M,S).

Volvarina taeniolata (MoOrcu, 1860). Formerly
Marginella californica Tomutn, 1916 [see Coan &
Rotu, 1966]. Common (4, 5, 7, 8, 13), under
noOcksi (DS Ey Ke),

Cancellaria obesa SowerBy, 1832. Rare (3, 12),
beach specimens (H, K).

Cancellaria cassidiformis SowErBy, 1832. Rare (3,
12), beach specimens (K, S).

Trigonostoma goniostoma (Sowersy, 1832). Un-
common (2, 5, 13), on mud covered rock shelf (D,
lah IS IME Se

Knefastia funiculata (Kiener, 1839-1840, ex Va-
LENCIENNES MS.). Uncommon, beach specimen
(H).

Knefastia olivacea (Sowrersy, 1833). Rare (8)
among rubble rocks (H).

Knefastia tuberculifera (BRoperie & SOWERBY,
1829). Rare (2), on mud covered rock ledge (S).
Clavus aeginus (Daux, 1919). Rare (6), dredged in
11m; Agua Verde Bay, Gulf of California (D).
Clavus ianthe (Da.r, 1919). Rare, beach specimen
(S).

Clathrodrillia maura (SowErBy, 1834). Rare, beach
specimen; Mazatlan, Mexico (D).
Clathrodrillia aenone (Dati, 1919). Rare (2)
rocky reef (S).

Crassispira flavonodosa Pitspry & Lowe, 1932.
Rare (7), under rocks (D).

Crassispira cf. C. monilifera (CARPENTER, 1857).

>

on

>]

Page 244

982

983

984

Common (2, 8), on muddy rocky reef and under
rocks; Mazatlan, Mexico to Panama (D, S).
Crassispira nymphia Pitspry & Lowe, 1932. Com-
mon (2), on mud covered rocky reef (D, H, M, S).
Crassispira pluto Puspry & Lowe, 1932. Common
(2), on mud covered rocky reef (D, S).

Mangelia finitima (Pitspry & Lowe, 1932). Rare
(8, 10), under rocks; Guaymas, Mexico [see Du-
SHANE & PoorMAN, 1967) (D, H, M, S).
Mangelia subdiaphana CarrENTER, 1864. Rare (8),
under rocks (S).

Mangelia antiochroa Pitssry & Lowe, 1932. Rare,
beach specimen (H).

Philbertia scammoni Datu, 1919. Rare (8), under
rocks; Scammon’s Lagoon, west coast of Lower
California (S).

Tenaturris carissima (Pitspry & Lowe, 1932).
Rare, beach specimen; Manzanillo, Mexico (M).
Tenaturris nereis (Pitspry & Lowe, 1932). Rare
(8), under rocks (D, H, M, S).

Hormospira maculosa (SowersBy, 1834). Uncom-
mon (2), on silty mud reef (D, S).

Pleuroliria picta (Rerve, 1843, ex Beck MS.).
Dredged in 11m (6), (D).

Conus princeps LINNAEUS, 1758. Beach specimen
(K).

Conus ximenes Gray, 1839. Common (3, 14), par-
tially buried in sand flats (D, H, K, M, S).
Conus mahogani REEVE, 1843. Uncommon (9), on
sand flats (D, S).

Conus regularis SowERBy, 1833. Rare (3), on silty
mud flats (K).

Terebra albocincta (CARPENTER, 1857). Uncom-
mon (9), sand bar next to Willard Island (coll.
DuShane December 1964, Puertecitos) (D, H, K,
S).

Terebra armillata Hinps, 1844. Uncommon (9), on
sand bars (D, S).

Terebra variegata Gray, 1834. Common (3, 14),
on silty mud flats and sand bars (D, K, S).

Terebra berryi CAMPBELL, 1961 (p. 26). Uncom-
mon (9), on sand bar; Puertecitos, Gulf of Califor-
nia [see DUSHANE, 1962] (D, S).

Terebra dushanae CAMPBELL, 1964 (p. 135). Rare,
beach specimen (H).

Bulla gouldiana Pitssry, 1895. Uncommon (12),
in tide pools (D, H, M).

Bulla punctulata A. ADAMs, in Sowersy, 1850. Un-
common, beach specimens (H).

Haminoea angelensis BAKER & HANNA, 1927. Beach
specimen (H).

THE VELIGER

986

992

993

995

1007

Vol. 10; No. 3

Haminoea strongi BAKER & Hanna, 1927. Common
(12), in small tide pools around rocks (D, H, S).
Acteocina angustior BAKER & Hanna, 1927. Rare
(8), under rocks (S).

Acteocina carinata (CARPENTER, 1857). Uncom-
mon (8), under rocks; Guaymas, Mexico (Du-
SHANE & PoorMANn, 1967) (S).

Acteocina infrequens (C. B. Apams, 1852). Un-
common (8), under rocks (S).

Retusa gonzagensis BAKER & HANNA, 1927 (p. 131).
Uncommon (8), under rocks [type locality] (S).
Pyramidella panamensis DALL & Bartscu, 1909.
Rare, beach specimen (D).

Aplysia sp. Rare (8), among rocks (L).
Berthellina sp. Uncommon (8), under rocks (L).
Tridachiella diomedea (BercH, 1894) (p. 125).
Uncommon (8), under rocks (L).

Rostanga pulchra MacFartanp, 1905 (p. 35). Un-
common (8), under rocks (L).

Acanthodoris spec. nov. Uncommon (8), under
rocks (L).

Hermissenda crassicornis (EscHscHoLtz, 1831)
(p. 15). Common (8), under rocks (L).

Spurilla chromosoma CockERELL & Euiot, 1905
(p. 31). Common (8), under rocks (L).
Melampus mousleyi Berry, 1964 (p. 152). Com-
mon (1), in mud under Salicornia bushes; Cholla
Cove, Bahia de Adair, Sonora, Mexico (D, S).

1023*Mclampus tabogensis C. B. ApaMs, 1852. Rare (1),

1025

28

39)

in mud under Salicornia bushes; Guaymas, Mexico
(DuSHANE & Poorman, 1967) (D).

Pedipes liratus BinNEy, 1860. Rare, beach speci-
mens (D, H).

AMPHINEURA

Chiton virgulatus SowerBy, 1840. Common (8, 10),
under rocks (H, M, S).

Acanthochitona cf. A. arragonites (CARPENTER,
1857). Rare (8), under rocks (H).

Acanthochitona exquisita (Pitspry, 1893). Com-
mon (7, 8), under rocks (S).

Ischnochiton tridentatus Pitspry, 1893. Rare (8),
under rocks (H).

Callistochiton gabbi Prrssry, 1893. Uncommon
(8), under rocks (H).

Callistochiton infortunatus Pitspry, 1893. Rare (7,
8), under rocks (H, S).

Lepidozona clathrata (Reeve, 1847). Rare (7, 8),
under rocks (H, S).

Vol. 10; No. 3

41 Lepidozona serrata (CARPENTER, 1864). Rare (7,
8), under rocks (H, S).

42 Lepidozona subtilis Berry, 1956. Common (8, 10),
under rocks (H, M).

44 Stenoplax limaciformis (SowERBY, 1832). Common
(7, 8), under rocks (H, S) .

45 Stenoplax magdalenensis (Hinps, 1845). Common
(7, 8), under rocks (H, S).

46 Stenoplax mariposa (Datu, 1919). Common (8,
10), under rocks (H, M).

47 Stenoplax conspicua sonorana Berry, 1956. Com-
mon (8, 10), under rocks (H, M).

LITERATURE CITED

ApAMs, CuHar.LEs BAKER
1852. Catalogue of shells collected at Panama, with notes on
synonymy, station and habitat, .. . Ann. Lyc. Nat. Hist.
New York 5: 229 - 296 (June) ; 297 - 549 [p. 549 unnumbered]
(July 1852)
Baker, FREDERICK
1926. Mollusca of the family Triphoridae.
Acad. Sci., ser. 4; 15 (6) :223 - 239; plt. 24
Baker, FREDERICK & G DaLttas 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 DaLttas 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)
1930. Some Rissoid Mollusca from the Gulf of California.
Proc. Calif. Acad. Sci., ser. 4; 19 (4): 23.- 40; plt. 1; 4 text figs.
(15 July 1930)
1938. Some Mollusca of the families Cerithiopsidae, Ceri-
thiidae and Cyclostrematidae from the Gulf of California and
adjacent waters. Proc. Calif. Acad. Sci., ser. 4; 23 (15):
217 - 244; plts. 17 - 23 (24 May 1938)
BartscH, PAuL
1907. The west American mollusks of the genus Triphoris.
Proc. U. S. Nat. Mus. 33 (1569) : 249 - 262; 1 plt.
(12 December 1907)
1911. The Recent and fossil mollusks of the genus Alabina from
the west coast of America. Proc. U. S. Nat. Mus. 39 (1790) :
409 - 418; plts. 61, 62 (13 January 1911)
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)
1915. The Recent and fossil mollusks of the genus Rissoina
from the west coast of America. Proc. U.S. Nat. Mus.
49 (2094): 33-62; plts. 28 - 33 (24 July 1915)
Bercy, Lupwic SopHus Rupo.F
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

Proc. Calif.
(26 April 1926)

THE VELIGER

Page 245

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)
BERRY, SAMUEL STILLMAN

1958. Is the Colorado River an efficient barrier to mollusks
distribution? Amer. Malacol. Union Bull. 24: 24

1959. Notices of new eastern Pacific Mollusca - III.

Leaflets in Malac. 1 (18): 107 - 114 (29 July 1959)

1964. Notices of new eastern Pacific Mollusca — VI. Leaflets
in Malacology 1 (24): 147 - 154 (29 July 1964)

Burcu, Joun Q. (Editor)

1944 to 1946. Distributional list of the west American marine
mollusks from San Diego, California to the Polar Sea. Conch.
Club S. Calif., minutes; 2 parts; pagination by issue; plts. 1 - 3.

CampPBELL, G. BRucE

1961. Four new Panainic gastropods. The Veliger 4 (1):
25 - 28; pit. 5 (1 July 1961)

1961. Range extension of Anatina cyprinus (Woop, 1828).
The Veliger 4 (2): 115; 3 text figs. (1 October 1961)

1964. New terebrid species from the eastern Pacific (Mollusca:
Gastropoda). The Veliger 6 (3): 132-138; plt. 17

(1 January 1964)
CarPENTER, PuHitip PEARSALL

1857. 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)

1864. | Supplementary report on the present state of our know-
ledge with regard to the Mollusca of the west coast of North
America. Rept. Brit. Assoc. Adv. Sci. for 1863: 517 - 686

(August 1864)
CravicEro, Don Francisco Javier, S. J.

1788. Storia della California (transl. fr. Italian by Sara E.
Lake « A.A.Gray, 1937). 413 pp. Stanford Univ. Press,
Stanford, California

Coan, EucENE Victor

1965. Kitchen midden mollusks of San Luis Gonzaga Bay.

The Veliger 7 (4) : 216 - 218; plt. 28; 1 table (1 April 1965)
Coan, EucEene Victor « Barry Rot

1966. The West American Marginellidae. The Veliger 8 (4) :

276 - 299; plts. 48-51; 5 text figs. (1 April 1966)
CockerELL, THEODORE Drew ALIson « CHARLES ELIOT

1905. Notes on a collection of California nudibranchs.

Journ. Malacol. 12 (3): 31-53; plts. 7, 8
Dati, WiLit1AM HEALEY

1919. Descriptions of new species of mollusks of the family
Turritidae from the west coast of America and adjacent regions.
Proc. U.S. Nat. Mus. 56 (2288): 1 - 86; 24 plts.

DonouuE, JERRY

1966. The range of Trivia myrae CAMPBELL. The Veliger
9 (1): 35-36; 1 map (1 July 1966)

1967. Additional remarks on the range of Trivia myrae Camp-
BELL. The Veliger 9 (3): 355 (1 January 1967)

Dunne, Peter Masten, S. J.

1952. Black robes in Lower California.

Berkeley, California, 540 pp.; 1 end map
DuSuane, HEeLen

1962. A checklist of mollusks for Puertecitos, Baja California,
Mexico. The Veliger 5 (1): 39-50; 1map (1 July 1962)

1966. __ Erroneous range extension for Tivela stultorum (Mawe,
1823). The Veliger 9 (1) :86 - 87 (1 July 1966)

Univ. Calif. Press,

Page 246

DuSuan_E, HELEN & Roy PooRMAN

1967. A checklist of mollusks for Guaymas, Sonora, Mexico.

The Veliger 9 (4) : 413 - 441; 1 map (1 April 1967)
Emerson, WILLIAM KEITH

1956. A new scaphopod mollusk, Dentalium ( Tesseracme)
hancocki, from the eastern Pacific. Amer. Mus. Novit. No.
1878: 7 pp., 1 fig. (28 September 1956)

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)

1964. A new name for Murex rhyssus Dax, 1919 (Mollusca:
Gastropoda). The Veliger 7 (1): 5-7 (1 July 1964)

ENGELHARDT, Fr. ZEPHYRIN, O. FE M.

1929. The Missions and Missionaries of California, vol. 1
Lower California. Mission Santa Barbara, Santa Barbara,
California, 784 pp.

Esc#iscHOLTZ, JOHANN FRIEDRICH

1829-1833. Zoologischer Atlas. Heft 4 (1831) Berlin.
Foun, ALEXANDRE GUILLAUME LEOPOLD DE
1867. Les Méléagrinicoles. Espéces nouvelles. 74 pp.; 6 plts.

1867-1871. Fonds de la mer. Etude internationale sur les parti-
cularités nouvelles des régions sous marines, etc. vol. 1: 316
pp.; 22 plts. [The work was originally issued at Bordeaux and
ascribed to “MM. Berchon, de Folin et Perrier.’’]

Fraser, C. McLEAn

1943. General account of the scientific work of the Velero
III in the eastern Pacific, 1931-41. Part III 1 (3): 259 - 431;
115 charts. Univ. South. Calif. Press, Los Angeles, Calif.

Howarb, Faye B.

1963. Notes on a Mitrella (Mollusca:Gastropoda) from the
Gulf of California. The Veliger 5 (4): 159 - 160.

(1 April 1963)

1963. Description of a new Pyrene from Mexico (Mollusca -

Gastropoda). Sta. Barbara Mus. Nat. Hist., Occ. Papers

7: 1-12; 2 pits. (May 1963)
KEEN, A. Myra

1958. Sea shells of tropical West America; marine mollusks

from Lower California to Colombia.
Stanford, Calif. (Stanford Univ. Press)
1961. A proposed reclassification of the gastropod family

i-xi + 624 pp.; illus.

Vermetidae. Bull. Brit. Mus. (Nat. Hist.) Zool. 7 (3):
15 - 214; 1 plt.; 33 figs.
1964. A quantitative analysis of molluscan collections from

Proc. Calif.
(1 July 1964)

Isla Espiritu Santo, Baja California, Mexico.
Acad. Sci., ser. 4, 30 (9) : 175 - 206; figs. 1 - 4
Lance, 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)
Lowe, HERBERT NELSON
1935. | New marine Mollusca from West Mexico, together with
a list of shells collected at Punta Penasco, Sonora, Mexico.

THE VELIGER

Vol. 10; No. 3

Trans. San Diego Soc. Nat. Hist. 8 (6): 15 - 34; plts. 1-4
(March 1935)
MacFarianp, Frank Mace

1905. A preliminary account of the Dorididae of Monterey

Bay, California. Proc. Biol. Soc. Washington, 18: 35 - 54
McLEan, James Howarp

1959. A new marine gastropod from West Mexico. The
Nautilus 73 (1): 9- 12; plt. 4 (1 July 1959)

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)

MENKE, Kart THEODOR

1850. | Conchylien von Mazatlan, mit kritischen Anmerkungen.

Zeitschr. Malakozool. 7: 161 - 173
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 « Hersert N. Lowe

1932. | West Mexican and Central American mollusks collec-
ted by H. N. Lowe 1929-31. Proc. Acad. Nat. Sci. Philadel-
phia 84: 33 - 144; 6 figs.; plts. 1-17; 2 photographs

(21 May 1932)
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)
RopeNn, 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)

Suasky, Donan R.

1961. | New deep water mollusks from the Gulf of California.
The Veliger 4 (1): 18-21; plt. 4, figs. 1-10 (1 July 1961)

1961. Notes on rare and little known Panamic mollusks. The
Veliger 4 (1) : 22-24; plt. 4, figs. 11-16 (1 July 1961)

SuHASKY, DONALD R. & G. BRUCE CAMPBELL

1964. | New and otherwise interesting species of mollusks from
Guaymas, Sonora, Mexico. The Veliger 7 (2): 114 - 120;
plts. 21, 22; 1 text fig.; 1 map (1 October 1964)

SLEVIN, JosEpH 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)

Spon, Gate G., Jr.

1960. Range extensions of two Panamic gastropods. The

Veliger 3 (1): 31 (1 July 1960)
STRONG, ARCHIBALD MCCLuRE

1938. | New species of west American shells. Proc. Calif.

Acad. Sci., ser. 4, 23 (14) : 203 - 216; plts. 15, 16 (24 May °38)
WILLETT, GEORGE

1939. Description of a new mollusk from California. The

Nautilus 52 (4): 123 - 124; plt. 9, figs. 1, la, 1b (1 Apr. 39)

lo

Vol. 10; No. 3

MESVELIGER

Page 247

Mating Behavior in Mitra 1dae MELVILL, 1893

BY

JEAN M. CATE

12719 San Vicente Boulevard, Los Angeles, California 90049

(8 Text figures)

INTRODUCTION

THE OBSERVATIONS SET FORTH in this paper are the
indirect result of an experiment to learn the feeding habits
and food preferences of Mitra (Atrimitra) idae MELVILL,
1893 (see Text figure 1). The intended study of foods
and feeding in M. idae was a complete failure; during the

Figure 1

Mitra (Atrimitra idae) MELvILL, 1893

approximately 8 months of experimentation with various
kinds of foods thought likely to tempt the animals,
apparently no feeding whatsoever took place. However,
the animals seemed unaffected by their apparent long fast
and were as active after months of presumed starvation
as when they were first collected, although they had not
grown during that time, as evidenced by periodic meas-
urements. i
Concurrent with the intended experiment, careful notes
were taken on the mating behavior of these animals

whenever it was observed, and it is these notes that are
the basis for this paper.

MATERIALS ann METHODS

A 15-gallon marine aquarium was set up on an outdoor
patio with a northern exposure. Though completely open
to the north, the porch was protected on the other three
sides by the walls of the U-shaped house, and was covered
by a translucent whitish plastic roof which admitted
filtered sunlight on bright days.

Local coarse beach sand, sterilized in an oven at 400° F
for one hour, comprised the bottom layer in the tank; this
was covered by a layer of finer sterile sand to a total
depth of about 3 inches. Sea water drawn from a marine
well was filtered through a double layer of heavy filter
paper before being added. These precautions were taken
in order to prevent any unknown food from obscuring
the results of the feeding experiment; in other words,
they were the control for the unsuccessful experiment, and
were not planned as a part of the mating-behavior obser-
vations.

Two filtering systems were used, one a subsurface
“Miracle” filter with two outlets, and the other an out-
side charcoal-and-fiberglass filter with one outlet; in
addition two airstones were used to ensure adequate aera-
tion; all this equipment worked off one commercial aquar-
lum pump. A 50-watt heater with a submerged ther-
mostat helped to regulate the temperature; a constant
temperature of 12° to 15° C was thus maintained through
the winter months, this being the average thermic range
in the area where the animals were collected. Approxi-
mately the same temperatures were maintained in summer
through the use of frozen plastic-jelly “Scotch Ice”
blocks. All filters and aeration systems were in operation
for 36 hours before the animals were introduced into the
tank.

Page 248

Twenty-six animals, ranging in size from 35.0mm to
56.9 mm, were placed in the aquarium February 24, 1965.
They had been collected by SCUBA divers 10 days pre-
viously, in 23 feet of water off Little Dume Point, near
Malibu, southern California, and kept in a large holding
tank while the experimental aquarium was made ready.
The habitat at the collecting grounds was a rocky reef
with scattered patches of sand.

All animals were code-marked with an electric drill,
following the procedure described by SToHLER (1962).
The marks were later painted with white “Snopake” for
easy identification during observation. Shell measurements
were recorded in the hope that some rate of growth
could be ascertained by the time the experiment ended.
While the whitened marks did prove helpful in distin-
guishing the animals from one another during observation.
no discernible growth increment in any animal was noted
when the experiment was discontinued approximately 8
months later; this fact further seemed to verify that the
animals had ingested no food during that period. Most
of the animals exuded a purple fluid when they were
subjected to work with the dental drill.

OBSERVATIONS

Although copulation was observed on numerous occasions,
the general behavioral pattern was about the same each
time; therefore this will be a composite report of the
several occasions when mating was observed.

Once or twice I noted attempts at copulation being
made at the water line near the filter’s air outlet at the
upper edge of the aquarium, but the usual place was the
sandy floor of the aquarium.

In every instance the female was the larger of two
coupled animals. The two most active mollusks were “No.
4,” a male whose shell measured 43.45mm by 14.50mm,
and “No. 8,” a female with a shell measuring 56.90 mm
by 19.60mm — the latter being the largest of all the
experimental animals. These same two individuals seemed
to prefer one another to the other animals; while they
were not mutually exclusive, my notes show a repetitive
pattern referring to “No. 4” and “No. 8” on several
occasions. This pattern does not occur among any other
specified individuals. Other coupling pairs, while generally
smaller, averaged the same as far as the male-female size
ratio is concerned.

All animals seemed to be more or less generally active
or dormant at the same times; frequently all would be
buried completely out of sight in the substrate for about
a week, then their siphons would appear above the sur-

THE VELIGER

Vol. 10; No. 3

face of the sand for a day or two, and finally all of the
animals would climb to every part of the aquarium at
once — the most popular place evidently being the area
nearest the air outlets at the waterline near the top of
the tank. Here they were out of the sea water, but within
the spray-zone of the aerator. This was especially inter-
esting behavior in animals usually found in greatest
abundance subtidally (see Text figure 2).

Figure 2

Bottom of foot of Mitra idac in normal crawling position, as viewed
through glass wall of aquarium

It was frequently observed that the periods of greatest
activity, especially sex activity, corresponded within min-
utes to a high tide in Los Angeles Harbor, despite the
fact that the animals had been collected from a subtidal
habitat. There seemed to be no correlation between the
active or dormant periods and the introduction of food
items; the animals simply continued whatever they had
been doing when food was put into the tank, ignoring the
food completely or moving in an other direction to avoid
contact with it.

PRELIMINARY SEX ACTIVITY

The first unusual behavior to be noted, other than the
normal moving around in the aquarium, was the exuda-
tion of a cloud of clear mucus from one of the larger
animals, which later proved to be a female. When occur-
ring up high in the tank at the waterline, this appeared
as a mass of tiny clear bubbles trapped in a cloud of
clear, colorless mucus roughly the same length as the
animal’s shell, but sagging somewhat heavily in the water
like a small plastic bag full of carbonated water (see Text
figure 3). Other, smaller animals quickly gathered around
this area, evidently drawn by some powerful attractant;
these later proved to be the male individuals. On a few
occasions I deliberately moved the male animals as far
away as possible from the “bubble-sac;” without excep-
tion they responded by moving rapidly back toward the
mucus sac (and incidentally toward the female individ-

Vol. 10; No. 3

THE VELIGER

Page 249

top of aquarium

“ys -— bubble sac

Figure 3

Typical “bubble sac” of female Mitra idae seen clinging
to aquarium wall at water line

ual), often even taking shorter routes by crawling up and
along suspended air-hoses (like walking a tightrope) rath-
er than to take the slower but more normal route of
crossing the sand floor and climbing the glass walls of
the aquarium. No animals were ever seen to climb along
the air-hoses at any time other than when a female was
exuding a “bubble-sac.” Whatever the attractant was, it
seemed to disperse very rapidly to all corners of the
aquarium, probably through the currents set up by the
filter- and aeration-systems.

Figure 4

Copulatory positions as viewed from above
Left, female; Right, male

In the upper corners of the tank at the points where
the air outlets emerged and the animals frequently con-
gregated, observations were not easily made due to the
position of the tank against a house-wall and the presence
of a metal rim on the tank which obscured my view of
the animals. It is possible that copulation did sometimes
occur at this location (see Text figure 4), but with less
success than on the sandy floor of the tank or beneath
the substrate. Some animals were seen to drop from the
waterline to the floor at times, possibly from the weight of
several individuals clinging to one female. At such times
no further sexual activity was noted.

At times females could be seen exuding the “bubble
sac” while at rest on the substrate. The first noticeable
action would be for the female to stand erect, with foot
fully extended and shell raised high so that a large por-
tion of the body above the foot could be seen — appar-
ently an unusual position except at times of pre-sexual
activity (see Text figure 5). This upper body area seemed

Figure 5

Typical female of Mitra idac in what is apparently a “courtship”
position. Note the swollen body, elevated shell, and extended siphon

tumid, swollen and more translucent than the solid milk-
white color usually observed in an animal at rest; it also
exhibited a pale pinkish color that seemed almost a blush
originating deep inside the body.

It is possible that the attractant material would first be
released at about this stage, for it was noted that several
of the other animals would become active, extending their
siphons far longer than normal, actively searching and
testing the water -— both males and females. Males
moved fairly rapidly toward the performing individual.
After remaining in this position for a period of about
5 to 10 minutes, the female would turn the right edge
of her foot toward the nearest male, and curl it upward
with a twisting, rolling motion that brought a small mass
of loosely agglutinated sand with it. It seems that sand-
grains adhere to the “bubble sac” when they come in
contact with it. This twisting action usually continued for
about another 10 minutes or so, until the female would

Page 250

THE VELIGER

Vol. 10; No. 3

sometimes be buried under a self-made sand pile; fre-
quently only a small area of black shell would be visible
under the loosely-packed sand mass.

COPULATION

At this point the male would align its shell in a position
approximately parallel with that of the female, and on
the female’s right side. The female’s shell would be under-
neath and to the left, with the aperture toward that of
the male; the male’s foot would be fully extended toward
his left side, providing full contact between the outer lip
of the female’s shell and with the left anterior portion
of the male’s foot about opposite the adapical end of
the female’s shell aperture. The verge would appear rather
rapidly from a point directly posterior to the head and
eyes of the male, probe two or three times under the left
side of the female’s shell above the foot, and finally come
to a stop. The verge emerged from the anterior end of
the male’s body, on the right side so that the right eye-
stalk was usually in contact with it throughout the copu-
latory act. The verge usually measured somewhat shorter
in length than the animal’s siphon; it had an extremely
swollen, more or less shapeless appearance, and was
about twice as thick in diameter as the siphon; it was a
pale, translucent bluish-white color (see Text figure 6).

Figure 6

Relative positions of male and female Mitra idae during copulation

v: verge s: siphon sg: sand grains adhering to “bubble sac”

f: foot of male m: shell of male

sf: shell of female

es: eye stalk

Upon first contact with the male’s verge, the female
quickly withdrew her body a short way into the shell,
then partly extended it again. After insertion of the verge,
the female continued her twisting motion, turning further
onto her back and bringing up more of the agglutinated
sand to partially cover both mollusks, somewhat like a
blanket. At about this time, both siphons gradually be-
came shorter and less active, resuming their normal ap-

pearance. After remaining in this position for about an
hour to an hour and a half, the verge was quickly with-
drawn and could be observed as a small white fleshy flap

Figure 7

Post-copulatory positions of male and female Mitra idae
sf: shell of female
ff: foot of female, starting to right itself

m: shell of male v: verge _ es: eye stalk

s: siphon

under the male’s shell as he raised and lowered it a few
times preparatory to moving away (see Text figure 7).
The translucent appearance and bluish color had disap-
peared, and the verge was the same color as the foot and
body of normal adults.

ADDITIONAL OBSERVATIONS

It was evident that mating sometimes occurs beneath
the sand substrate, as well as on the surface, and occasion-
ally at or above the waterline of an aquarium. On one
occasion I carefully moved a partially buried mating
pair to the surface for easier observation, whereupon the
male quickly discharged into the water rather large
amounts of what may have been sperm, in a stream of
fairly solid whitish flakes which soon disintegrated and
were carried off by the water current. On this occasion,
copulation was interrupted and the animals separated of
their own accord. At other times, however, I was able to
move mating animals to a better vantage point without
disturbing the copulatory procedure.

On another occasion it was noted that two males were
trying to mate with the same female, one of the males
being atop the surface of the sand as noted above, and
other buried beneath it. It seems improbable that the
mating act of the third specimen was completed at
this time.

The presence of living Crepidula onyx SowErRBy, 1824
on the shell of a female Mitra idae (a not unusual occur-
rence) seemed a deterrent to copulation, although it did
not completely prevent it. The male Mitra would, while

Vol. 10; No. 3

THE VELIGER

Page 251

“feeling” a female’s shell with its siphon, quickly with-
draw it when coming in contact with the Crepidula’s
shell, evidently being able to sense the difference between
the two kinds of shell.

TERMINATION or THE EXPERIMENT

The experiments were terminated and the animals still
remaining alive were put into alcohol in November 1965,
when an unidentified red-colored bacterial growth in-
vaded the filter systems and the entire aquarium, com-
pletely covering the floor and sides, and the animals as
well. For a time the bacterial growth seemed controllable
through the use of medicinal penicillin. It was difficult,
however, to regulate the amount of the drug correctly,
and in our attempts to strike a balance between barely
controlling the obnoxious growth and giving too much
penicillin to the animals (several of the Mitra died on
one or two occasions when the drug was used), we found
it impossible to continue.

EGGS

No eggs were ever laid by any female Mitra idae in the
aquarium during the course of the experiment, despite
the numerous copulatory acts observed during the several
months of the experiment.

In the field, observations made by SCUBA divers (Dr.
R. C. Fay, personal communication) revealed that Mitra
idae were laying eggs during July and August, 1965. These
eggs were laid on a shell substrate at a depth of 25 feet
in the open ocean off southern California, in approxi-
mately the same area where the experimental animals
had been collected.

A year or two earlier, an unsuccesful attempt had been
made to hatch egg capsules of Mitra idae in a laboratory
at the University of California, Berkeley. At that time, a
specimen of Mitra idae collected by R. Ames in 30 feet
of water at Mission Point, Carmel (Monterey County),
was maintained in a jar of sea water in a cold room at
14°C. An airstone was kept in the jar for aeration, but
no food was offered at any time during the nearly 3
months the animal was under observation. Approximate-
ly one and a half months after capture, the animal depos-
ited a large egg mass on the side of the jar (see Text
figure 8). About one month later, one additional egg
capsule was deposited. Although at this time motion could
be observed in the first-laid capsules, indicating embryonic

Figure 8

Egg capsule of Mitra idae

life in the unhatched veligers, none of the eggs in any of
the capsules ever hatched.

CONCLUSIONS

During the course of eight or nine months, observations
were made on the mating behavior of 26 animals of
Mitra idae MEtvi.v. Based on these observations, it seems
evident that the following conclusions may be made:

1. Males are able to locate females by some sensory
means when attracted by a substance emitted by the
female in a mucus “bubble sac.”

2. The attractant provided by the female is evidently
dispersed by means of water currents.

3. Females use their muscular foot and the “bubble sac”
in a twisting motion to pick up sand from the substrate,
ultimately nearly covering themselves and sometimes the
mate as well with loosely agglutinated sand.

4. Observations made during this experiment confirm
the fact that Mitra idae is a dioecious species, and that
fertilization is internal.

5. There seems to be a relationship between the timing of
mating action in Mitra idae and the exact hour of
local high tides even when the animals are a considerable
distance from the ocean.

Page 252

6. Mitra idae is able to subsist in a healthy condition
without food for a period of at least 8 to 9 months.

7. Either the state of captivity or a lack of food, or both,
may have inhibited the production of fertile eggs and the
growth of the animals.

ACKNOWLEDGMENTS

Many persons provided help and encouragement toward
the carrying out of this experiment; to all of them I
wish to express my gratitude. Crawford Cate was helpful
and encouraging in many ways; Dr. Rudolf Stohler
assisted in setting up of the original experiment, including
the code-marking of the animals, and offered several

THE VELIGER

Vol. 10; No. 3

suggestions for possible foods; Dr. and Mrs. Rimmon C.
Fay were extremely helpful in providing animals, various
kinds of marine organisms which we tried as food, and
fresh sea water, as well as personal observations on these
animals in their natural habitat; Saul L. Fox, M. D., very
kindly donated the medicinal penicillin, and Mrs. Emily
Reid adapted Kodachrome slides into the excellent text
figures illustrating the experiment.

LITERATURE CITED

STOHLER, RUDOLF

1962. Preliminary report on growth studies in Olivella bipli-
cata. The Veliger 4 (3): 150-151; plt. 36 (1 Jan. 1961)

Vol. 10; No. 3

THE VELIGER

Page 253

Structure of the Bivalve Rectum

I. Morphology

THOMAS C. JEGLA

Department of Biology, Kenyon College, Gambier, Ohio 43022

AND

MICHAEL J. GREENBERG

Department of Biological Sciences, Florida State University, Tallahassee, Florida 32306

(Plates 36 to 40; 1 Text figure)

INTRODUCTION

THE INTESTINE OF BIVALVES, after leaving the stomach,
loops through the visceral mass and is succeeded by the
rectum, which courses posteriorly, usually through the
pericardium and ventricle, over the posterior adductor,
and finally opens into the exhalant current. The rectum
is usually attached to the pericardium and ventricle only
at its entrance and exit sites in these tissues.

Several investigators, in studies of the digestive tract of
bivalve mollusks, have devoted some attention to the mor-
phology and histology of the rectum. These include DAKIN
(1909), Pecten maximus (LinnaEus, 1758); GuTHEIL
(1912), Anodonta cellensis ScurRGTER, 1779; YoNGE
(1923), Mya arenaria LINNAEUS, 1758; YoNcE (1926),
Ostrea edulis LinNaEuS, 1758; WuiTE (1937), Mytilus
edulis LinNAEus, 1758; YoncE (1941), seven species of
protobranchs; and GattsorF (1964), Crassostrea virgin-
ica (GMELIN, 1791). Although primarily interested in the
histology and physiology of the heart, MotLey (1933) did
discuss the trans-ventricular region of the fresh-water bi-
valve rectum. ARAKAWA (1963, 1965), in his studies on the
shape and constitution of molluscan faeces, also looked at
the histology of the rectum. Finally, Puituis (1966) and
Nystrom (1967), in the course of physiological studies
of the Tapes watlingi IREDALE, 1958 and Spisula solidis-
sima (Dittwyn, 1817) rectums, briefly examined histo-
logy. These studies all clearly show that the construction
of the rectum varies markedly in the bivalve mollusks.

This report describes the morphology of the rectum of
at least one species in each of the orders of the class
Bivalvia; 23 families are represented. While the variation
is large, familial similarities in rectal construction have
been found.

MATERIALS anno METHODS

The rectum was removed from medium and large indi-
viduals, sometimes with the ventricle around it, and fixed
in aqueous Bouin’s fluid; if the clam was very small, the
entire animal was fixed. After the usual histological pro-
cedures, the 7-10 micron thick sections were dyed either
with Masson’s trichrome stain (HuMason, 1965), Mal-
lory’s triple connective tissue stain (Guyer, 1953), or
with Ehrlich’s hematoxylin with eosin as a counterstain.
Masson’s, with a very short treatment in the fast green,
proved to be most effective for differentiating muscle and
connective tissue fibers in molluscan tissues.

RESULTS

Species of bivalves differ in the cross-sectional morpho-
logy of their rectums. Variations in three major character-
istics occur:

(1) Bore or shape of the lumen. The lumen is sur-
rounded by a single layer of columnar epithelial cells,

Page 254

the surface area of which is increased by differential
size of the epithelial cells, or by simple folding at various
points around the lumen, or by the occurrence of one or
more prominent typhlosoles.

(2) Thickness of the wall. A basement membrane,
composed of connective tissue fibers that are stained by
fast green or aniline blue, lines the basal portions of the
epithelial cells. Peripheral to the basement membrane, the
tissue of the rectal wall varies considerably in thickness
and composition.

(3) Composition and arrangement of tissue elements
in the wall. In many species this tissue is thinner than the
layer of columnar epithelium and is composed essentially
of connective tissue fibers; but in over 50% of the species
studied, it is as much as 19 times the thickness of the
columnar epithelial layer. In rectums with substantial
walls, connective tissue is the basic component; although
muscle tissue is usually present, it varies considerably in
density and orientation. These characteristics will be
discussed in greater detail below.

While the morphology of the rectum remains uniform
from intestine to anus in many species, in others there is
a progressive change of structure. Therefore, most inter-
specific comparisons in this paper are made at a standard
position, homologous in most of the animals studied:
namely, at the level of the auriculo-ventricular valves
in the trans-cardiac region of the rectum.

1. Bore or shape of the lumen

While the shape of the lumen is variable in bivalve
mollusks, generally there is uniformity within a family.
Some species have an entirely smooth-bored rectum; all
the columnar epithelial cells are of the same size (Text
figure la; Plate 36, Figure 1 and Plate 38, Figure 5). In
the majority of species studied, the smoothness of the
bore is interrupted by ridges and furrows, folds, typhlo-
soles, or some combination of these (see diagrams in
Text figure 1). A differential in size of the epithelial cells
results in ridges where the cells are largest, and furrows
where the cells are smallest (Text figure 1b and Plate 37,
Figure 3). Folding of the rectal wall also occurs (Text
figure lc and Plate 36, Figure 2). A typhlosole, as we
distinguish it, is a projection of connective tissue, with
its covering epithelium, into the lumen (Text figures 1d
and le) ; the structure extends for most of the length of
the rectum. A typhlosole may be small (Plate 36, Figure
2) or voluminous (Plate 39), single (Plate 39) or multiple
(Plate 40, Figure 9). Whatever their condition, typhlosoles
considerably increase the surface area of the epithelial
lining of the lumen.

THE VELIGER

Vol. 10; No. 3

The smooth-bored condition may occur among the
Protobranchia, Heterodonta and Adapedonta. While it is

Figure 1
Diagrammatic Cross-Sections of Bivalve Rectums,
Illustrating Variation in the Bore or Shape of the Lumen
a - Smooth. Epithelial lining of more or less uniform height.
b — Variation in height of epithelium producing ridges and furrows.

c — Folds in the epithelium. d — Small typhlosoles. Peripheral
wall tissue, as well as epithelium, is folded into the lumen.
e — Single, large typhlosole.

frequently found in such small species as Nucula and
Donax variabilis Say, 1822, there is certainly no relation-
ship between the size of the animal, the diameter of the
lumen, or the extent of irregularity of the rectal wall.
For example, a smooth-bored rectum occurs in the large
species Mya arenaria (Plate 38, Figure 5), Chlamys hindsi

Vol. 10; No. 3

CarpPENTER, 1864 and Cyrtopleura costata (LINNAEUS,
1758). The rectum of these animals has a large lumen
and an overall diameter of 1.5-2.0mm, whereas the
rectal diameter in the small species is less than 0.3mm
(see Mactra, Plate 36, Figure 1). The smoothness of the
bore is also unrelated to the thickness of the rectal wall;
compare thin-walled Mactra (Plate 36, Figure 1) with
thick-walled Mya (Plate 38, Figure 5). According to
YonceE (1923) the rectum of Mya arenaria has its colum-
nar epithelium thrown into longitudinal folds, but his
illustration (see his figure 24) is of a region very near
to the anus. Epithelial folds are commonly found near
the anus in rectums where no folds existed in the cardiac
region; we have observed this in Chlamys and Macoma,
and Ga.ttsorF (1964) observed it in Crassostrea.

Ridges and furrows along the entire inside perimeter
of the rectum were observed in Acila castrensis (Hinps,
1843), Lucina floridana Conran, 1833, Tridacna maxi-
ma (Ropine, 1798) and Cardiomya oldroydi (DALL in
Oxproyp, 1924). The cells vary in height by factors of
1-5, but the absolute size differences vary widely and
depend on the species. Often, in combination with differ-
ential cell size, folding of the epithelium occurs, and
apparently is found only in those species having a thin-
walled rectum. We have observed it in Mytilus edulis
(Plate 37, Figure 3) and Brachidontes exustus (LINNAEUS,
1758).

Typhlosoles, alone, invade the rectal lumen in many
species in the families Arcidae, Pinnidae, Cardiidae and
Unionidae. One or more typhlosoles were observed in
the Vulsellidae (von HArFrner, 1958), and one large
bifurcated typhlosole occurs in the Ostreidae (YoNGE,
1926; Gattsorr, 1964). With respect to lumen volume,
the typhlosoles are small in the Cardiidae, but many of
them occur on the perimeter of the lumen (Plate 40,
Figure 9). The Arcidae and Pinnidae have a typhlosole
that is intermediate in size (Plate 39, Figure 8). A single
typhlosole reaches the ultimate in relative size in the fresh-
water family Unionidae where it may reduce the lumen
to a slit (Plate 39, Figure 7).

Combinations of ridges and furrows and one or more
typhlosoles occur. Among these are Mytilus californianus
Conrab, 1837, Solen sicarius Gouin, 1850 (Plate 36,
Figure 2) and all the species of Veneridae we have studied
[Mercenaria mercenaria (LINNAEUS, 1758) (Plate 38,
Figure 6), M. campechiensis (GmeEtIN, 1791), Chione
cancellata (LiInNAEUuS, 1767) and Macrocallista nimbosa
(Licutroot, 1786) and Tapes watlingi (see PHILLIs,
1966) |. The typhlosoles in these species may be large
or small, while, in ridges and furrows, the size of the
columnar cells varies from a factor of two in Chione and

THE VELIGER

Page 255

Macrocallista to five or six in Mercenaria. The smallest
typhlosoles are narrow and have a very thin core of
connective tissue, but the largest are very broad-based
and have a massive core of connective tissue and some
muscle cells.

2. Thickness of the wall of the rectum

In five orders of Bivalvia, four or more species were
sampled (Table 1). Within the Protobranchia, Anisomy-
aria and Schizodonta, only a small variation in thickness
of the rectal wall was observed. Greater differences occur
in the Adapedonta and the large order Heterodonta
(Table 2). However, the order notwithstanding, variation
is minimal within the families from which we have studied
more than one species. The 40 species discussed in this
report can be conveniently divided into three types with
regard to the thickness of the rectal wall peripheral to
the basement membrane of the columnar cells:

(a) epithelial walls

(b) thin walls

(c) thick walls.

(a) Epithelial walls. In species with epithelial rectal
walls, the columnar cells and their basement membrane
account for 90-100% of the wall of the rectum (Plate
36, Figure 1; Plate 37, Figure 4). This condition was
found in all four species of Protobranchia, several Hetero-
donta, and in the lone species of the Anomalodesmata
and Septibranchia. Epithelial walls are prevalent among
the small species but are not necessarily highly correlated
with a small lumen nor with epithelial cell height. A
scarcity of tissue external to the basement membrane is
observed in the large rectum of Macoma nasuta (ConrRaD,
1837) (Plate 37, Figure 4) and in the small rectum of
Mactra fragilis GMELIN, 1791 (Plate 36, Figure 1); the
lumen is 70 times larger in Macoma than in Mactra.
Also, the columnar cells are six times taller in Macoma
than in Mactra.

(b) Thin walls. In those species with a thin-walled
rectum, the average height of the columnar cells and
their basement membrane account for 40-80% of the
rectal wall; the tissue peripheral to the basement mem-
brane is about as thick as the height of the columnar cells
and basement membrane combined (Plate 37, Figure 3).
We have found the thin-walled condition in all three
families of the Anisomyaria and it is apparently a char-
acteristic of this order. Also half the families studied in
both the Heterodonta and Adapedonta have thin walls.
Again, thinness of wall and lumen volume are not corre-
lated.

Page 256

THE VELIGER Vol. 10; No. 3

Table 1

Systematic Distribution of the Bivalves used in this Study.
Classification after THmELE (1935), Appotr (1954) and Morton « YoncE (1964)

SUBCLASS
ORDER FAMILY Genus and species
PROTOBRANCHIA
NucULIDAE Nucula sp.; Acila castrensis (Hinps, 1843)
NUCULANIDAE Nuculana hamata (CarpEenTER, 1864) ; Yoldia limatula (Say, 1831)
LAMELLIBRANCHIA
TAXODONTA ARCIDAE Noetia ponderosa (Say, 1822)
ANISOMYARIA MyTILIDAE Brachidontes exustus (LINNAEUS, 1758) ; Mytilus edulis LINNAEUS,
1758; Mytilus californtianus Conrap, 1837
PINNIDAE Atrina rigida (LicutrFoot, 1786) ; A. serrata SowERBy, 1825
PECTINIDAE Chlamys hastata hericia (Goutp, 1850); C. hindsi CarRPENTER,
1864
HETERODONTA CorBICULIDAE Polymesoda caroliniana (Bosc, 1802)
SPHAERIIDAE Sphaerium sp.
LucINnIDAE Lucina floridana Conran, 1833
CaRDIIDAE Dinocardium robustum (Licutroot, 1786); Serripes groenlandi-
cus (BRuGurERE, 1789) ; Clinocardium nuttalli (Conrap, 1837)
TRIDACNIDAE Tridacna maxima (Roéoine, 1798)
VENERIDAE Mercenaria mercenaria (LinnaEus, 1758); M. campechiensis
(GmeE.tn, 1791) ; Chione cancellata (Linnaeus, 1767) ; Macro-
callista nimbosa (LicHTFoor, 1786)
MactrRIDAE Mactra fragilis GMELIN, 1791; Spisula solidissima (DiLLWYN,
1817) ; Rangia cuneata (Gray, 1837)
TELLINIDAE Macoma nasuta (Conran, 1837)
DoNAcIDAE Donax variabilis Say, 1822
SANGUINOLARIDAE Tagelus gibbus (SPENGLER, 1794)
SCHIZODONTA UNIONIDAE Actinonaias carinata (BARNES, 1823) ; Cyclonaias tuberculata (Ra-
FINESQUE, 1820); Quadrula quadrula (RAFINESQUE, 1820) ;
Anodonta cataracta Say, 1817; Amblema plicata RAFINESQUE,
1820
ADAPEDONTA SOLENIDAE Solen sicarius Goutp, 1850
HIATELLIDAE Panope generosa Goutp, 1850
My1paE Mya arenaria LINNAEUS, 1758
PHOLADIDAE Cyrtopleura costata (LINNAEUusS, 1758)
ANOMALODESMATA LYONSMDAE Lyonsia striata (Montacu, 1815)
SEPTIBRANCHIA
CusPIDARIDAE Cardiomya oldroydi (Datu in Oxproyp, 1924)
Explanation of Plate 36
Variability in Rectums of Similar Small Size
Figure 1: Mactra fragilis. Epithelial wall; smooth bore.
Figure 2: Solen sicarius. Thin wall; bore interrupted by epithelial

folds. Note small typhlosole in upper left.
C —- columnar epithelium
H -— heart muscle

L - lumen
W -— tissue of the wall

THE VELIGER, Vol. 10, No. 3 [JEcLA & GREENBERG] Plate 36

Vol. 10; No. 3

Thickness of the Rectal Wall and Height of the Columnar
Epithelial Cells in Relation to Size of the Species and its

(Size of marine species calculated from Axpsort, 1954)

Nucula sp.
Acila castrensis
Nuculana hamata
Yoldia limatula
Noetia ponderosa
Brachidontes exustus
Mytilus edulis
Mytilus californianus
Atrina rigida
Atrina serrata
Chlamys hastata hericia
Chlamys hindsi
Sphaerium sp.
Lucina floridana
Dinocardium robustum
Serripes groenlandicus
Clinocardium nuttalli
Tridacna maxima (small)
Mercenaria mercenaria
Mercenaria campechiensis
Chione cancellata
Macrocallista nimbosa
Mactra fragilis (small)
Spisula solidissima
Macoma nasuta
Donax variabilis
Tagelus gibbus
Actinonaias carinata
very small
very large
Cyclonaias tuberculata
Quadrula quadrula
Anodonta cataracta
Amblema plicata
Solen sicarius
Panope generosa
Mya arenaria
Cyrtopleura costata
Lyonsia striata
Cardiomya oldroydi

THE VELIGER

Table 2

Rectum.

Length of
animal (mm)

the rectum ()1)’

Diameter of

and basement membrane (,)

Height of columnar cells

the wall (n)

Thickness of

epithelium : wall

Ratio

Page 257

Page 258

(c) Thick walls. A rectum is thick-walled when the
average height of the columnar cells and their basement
membrane is less than 40% of the thickness of the wall.
A thick wall is characteristic of Mya arenaria (Plate 38,
Figure 5), the Veneridae (Plate 38, Figure 6), Cardiidae
(Plate 40, Figure 9), and the Unionidae. Many of the
species in these families are large bivalves, but not only
the large bivalves have a thick-walled rectum. The thick-
walled rectum bears no apparent relationship to gauge or
shape of the lumen. Thus, the lumen is large and smooth
in Mya (Plate 38, Figure 5); large and interrupted by
typhlosoles in the Veneridae (Plate 38, Figure 6) and
Cardiidae (Plate 40, Figure 9) ; and nearly obliterated by
the huge typhlosole in the Unionidae (Plate 39, Figure 7).

3. Composition and arrangement of tissue elements

Conspicuous tissue elements in bivalve rectums are colla-
gen fibers, fibroblast cells, phagocytes, smooth muscle
cells, and sometimes vesicular cells. Layering may occur.
The nature of the tissue layers, and of their component
elements, is closely related to the thickness of the rectal
wall external to the basement membrane. Not only is
there great similarity within families, but also within some
orders.

In those species with epithelial walls, what little outer
tissue is present usually consists of a few collagen fibers
and perhaps a few fibroblasts and muscle cells.

Many species with a thin-walled rectum are exemplified
by the condition observed in the family Pinnidae (Plate
39, Figure 8) ; the wall of the rectum consists of a dense
layer of collagen fibers, fibroblast cells and scattered
muscle cells. However, the peripheral portion of the rectal
wall of the Pectinidae and Mytilus californianus is com-
posed almost entirely of vesicular cells; collagen fibers
and muscle cells are densely packed immediately adjacent
to the basement membrane but also are scattered through
the vesicular layer. In addition the Pectinidae may have
visceral organs closely attached to the rectum and they
progress with it for some distance through the pericardium

THE VELIGER

Vol. 10; No. 3

and ventricle. The digestive gland is especially prominent
in Chlamys hindsi, while in C. hastata hericia (GouLp,
1850) both digestive gland and gonad are present. KEL-
Locc (1890) also found gonad investing most of the
length of the rectum of the oyster, Crassostrea virginica.
Spisula solidissima, a heterodont bivalve, has the central
half of the wall heavily infiltrated with longitudinal,
circular, and oblique muscle fibers and the peripheral half
primarily of collagen fibers.

In those species with a thick-walled rectum, the basic
component of the wall is connective tissue. In all cases
a dense layer of collagen fibers and fibroblasts is found
adjacent to the basement membrane; there are also muscle
cells in this layer in the Veneridae and Cardiidae (Plate
40). Going peripherally the wall differs, in the various
families, primarily with regard to the density and arrange-
ment of muscle. A thin, medial layer of circular and
oblique muscle is found in the Unionidae. Venerids have
a medial layer of dense longitudinal muscle fibers (Plate
38, Figure 6). In Mya arenaria (Plate 38, Figure 5) and
the Cardiidae (Plate 40, Figure 9), the dense concentra-
tion of circular and oblique muscle occurs in a peripheral
layer.
In most species the core tissue of the typhlosole is
essentially that of the connective tissue layer adjacent
to the basement membrane, but in some species, more than
one layer of the wall may extend into the typhlosole
(Plate 40, Figure 10). The large unionid typhlosole has
a dense filling of collagen fibers, fibroblasts and muscle
cells which fold in from the medial layer. In addition,
numcrous large blood sinuses course throughout the fibrous
layer (Plate 39, Figure 7).

DISCUSSION

The distinguishing morphological features — shape of the
lumen, thickness of the rectal wall, occurrence of the
various tissuc elements, and orientation of muscle fibers
— which characterize the rectums of bivalve mollusks

Explanation of Plate 37

Variability in Rectal Structure

Figure 3:

Mytilus edulis. Characteristic shape of anisomyarian

rectum. Thin wall; ridges and furrows. The two large ventral ridges
are the remnant of the intestinal typhlosolar folds.

Figure 4: Macoma nasuta. Large rectum with epithelial wall and
smooth bore; compare with Figure 1, Plate 36. Note folds on

ventral side.
C — columnar epithelium
H — heart muscle

L — lumen
W — tissue of the wall

Tue VELIcER, Vol. 10, No. 3 [JECLA « GREENBERG] Plate 37

he
ae

rain ae 2

Figure 3

7

Figure 4

Vol. 10; No. 3

THE VELIGER

Page 259

have been summarized, by families, in Table 3. These
features are consistent enough to enable one to distinguish
between families. Some similarities also appear at the
ordinal level; these are discussed below:

Among the Protobranchia (Nuculidae and Nuculanidae)
the structure of the rectum is only slightly variable. It
is smooth bored and has an epithelial type wall. An illus-
tration of Nucula nucleus (LINNAEUS, 1758) in PELSE-
NEER (1906) and descriptions of Malletia by YONGE
(1941) indicate that these species do not differ materially
in pattern from those we have observed.

Rectums in the order Anisomyaria [Pecten maximus
(Dakin, 1909) ; Lima inflata Lamarck, 1819 (von Stup-
NITZ, 1931) ; Ostrea edulis (YoncE, 1926) ; Mytilus edulis
(Wuire, 1937); Vulsella (von HAFFNER, 1958) ; Crass-
ostrea virginica (GatTtsorF, 1964) ; and species listed in
Table 2] are basically similar. All have a relatively thin
wall; the muscle fibers are few in number, and are usually
not organized into layers or bundles. Also, in Mytilus cali-
fornianus and the Pectinidae visceral mass tissue (vesicular
storage cells, gonad, or digestive gland) extends, with the
rectum into the trans-ventricular region. This has also
been observed in Crassostrea virginica by KELLoce (1890)
and therefore it may be a general morphological charac-
teristic of the order. On the other hand, this extension
of the viscera may be a seasonal phenomenon associated
with expansion of the gonad in the breeding season.

The continuous change in rectal structure, from intes-
tine to anus, is conspicuous in the Anisomyaria. One of
these changes is the reduction and decrease in height of
the well-developed intestinal typhlosole folds to two broad
ridges with a mid-ventral furrow (Plate 37, Figure 3).
The large, bifurcated Atrina rectal typhlosole (Plate 39,
Figure 8) is clearly homologous to the same structure in
Mytilus intestine. Finally, in most species, the rectum is
flattened to a crescentic shape.

We have observed five genera in the order Schizodonta,
all in the family Unionidae. There are no significant
differences among the species in structure of the rectum,
and the rectum is strikingly different from that of any
other species in any other order of bivalves that has been
examined. The one unmistakable feature common to all
unionids is the very large typhlosole which reduces the
lumen to a slit, and which persists for the entire length of
the rectum. Also characteristic is the extensive blood sinus
system permeating the connective tissue core of the typhlo-
sole. Previous studies have included the genus Anodonta
(Vocr « Yunc, 1888, and GutuHem, 1912) and the
genera Tritogonia, Lasmagonia, Elliptio, Fusconaia, Lam-
psilis and Megalonaias (Mot ey, 1933).

While species in the large order Heterodonta exhibit
a variety of rectal structures, we have observed consider-
able intra-family similarity. Furthermore, reports in the
literature support this. Outstanding characteristics of the
order as a whole do exist. Large animals, especially in
some families, show a heavy muscular development,
usually circular, in a robust rectal wall. The occurrence
of many small typhlosoles is another common feature (see
Table 3).

Although the number of species available for compari-
son is small, the degree of variation in rectal form in the
order Adapedonta seems to approach that of the hetero-
donts. Thus, while Mya and Cyrtopleura are without
significant sculpturing of the columnar epithelium both
differences of columnar cell size and typhlosoles occur
in Solen and Panope. Again, the wall is thick and heavily
muscular in Mya but relatively thin in Cyrtopleura and
Solen. In all, the rectum is essentially circular in trans-
verse section.

We have, thus, found only two orders of Bivalvia whose
rectal structure is characteristic of that, and no other,
order: Anisomyaria and Schizodonta. According to Cox
(1960), the modern Arcacea and the superfamilies of the
Anisomyaria arose from a common cyrtodontid ancestral
stock in the early Ordovician. However, based on the
morphology of the gills and stomach and on the fossil
record there is some question as to whether the Mytilacea
share this ancestry or arose from the Modiolopsidae,
another early Ordovician fossil family; this evidence is
also summarized by Cox. The unique form of the aniso-
myarian rectum, and its occurrence, although modified,
in an arcid confirms the idea of common ancestry. A
morphological feature which occurs in such highly evolved
animals as the Ostreacea, Pinnacea, and Mytilacea, must
be highly conservative. Since we find no evidence of its
existence in the Protobranchia or Heterodonta (although
it would be interesting to examine the primitive Astarti-
dae) we assume that the anisomyarian rectum arose after
the divergence of the cyrtodont stock from that of the
above subclasses.

StasEK (1963) has produced a phylogenetic scheme
correlating his studies on the ctenidium-palp association,
the stomach types of PurcHon (1963), and the fossil
record. The distribution of the anisomyarian rectum is
identical to that of the stomach type. Namely, all the aniso-
myarians except the Pectinacea (and the Anomiacea
which we haven't studied) have stomach type 3, which
is also found nowhere else. The Pectinacea have a type 4
stomach and also, in this group, the usual rectal structure
cannot be recognized. The ctenidium-palp association

i}

Page 260

THE VELIGER

Table 3

Vol. 10; No. 3

Summary of the Structure of the Rectum in some Families of Bivalvia

Ratio
epithelium:
FAMILY Shape of Lumen wall thickness Tissue Elements
NUCULIDAE smooth or small difference in cc size 0.9-1.0 none or ct
NucuLANIDAE nearly smooth 0.9-1.0 none or ct
ARCIDAE large typhlosole 0.58 mostly ct, cm; some ve
MYyTILIDAE ridges and furrows, folding of wall or 0.54-0.62 mostly ct; a few cm, Im;
a typhlosole may have layer of vc
PINNIDAE 1 large typhlosole 0.45 mostly ct; a few Im, om
PECTINIDAE smooth or small typhlosoles 0.42-0.54 some ct, cm, Im; visceral
mass tissues
SPHAERIIDAE essentially smooth 0.85 ct
CorBICULIDAE essentially smooth 0.85 ct; a few cm
LUCINIDAE small difference in cc 0.65 dense layer of ct
CaRDIIDAE ridges and furrows, and many typhlo- 0.05-0.22 dense layer of ct, a few cm,
soles om; dense, thick layer
cm, Im
TRIDACNIDAE small difference in cc 0.61 mostly ct; a few cm
VENERIDAE ridges and furrows, and many typhlo- 0.22-0.35 dense layer of ct, some cm,
soles Im; dense layer ct, Im; layer
ct, some Im
MactRIDAE smooth or small difference in cc 0.50-0.91 some ct or layer of cm, om,
Im and dense layer ct, some
Im
TELLINIDAE smooth to a small difference in cc 0.91 mostly ct
DoNACIDAE smooth 0.71 ct
SANGUINOLARIIDAE smooth 0.75 ct, a few cm
UNIONIDAE 1 large typhlosole 0.16-0.35 dense layer ct; layer of cm,
om; layer ct with cm
SOLENIDAE ridges and furrows and 1 typhlosole 0.68 dense ct, some cm
HIATELLIDAE ridges and furrows and several typhlo- 0.38 ct, some cm
soles
MyIDAE essentially smooth 0.22 layer of ct; dense layer cm,
some ct, om
PHOLADIDAE smooth 0.50 ct, some cm
LYONSIDAE smooth 1.0 none
CusPIDARIIDAE smooth to a small difference in cc 0.94 ct

Im — longitudinal muscle cells
om — oblique muscle cells
ve — vesicular cells

ce — height of columnar cells
cm — circular muscle cells
ct — connective tissue

Explanation of Plate 38

Variability in Thick-Walled Rectums of Similar Large Size

Figure 5: Mya arenaria. Smooth bore. Note dense circular muscle
layer in periphery of the wall.
Figure 6: Mercenaria mercenaria. Many small typhlosoles as well
as ridges and furrows. This is a characteristic of venerid rectums.
Note thick layer of longitudinal muscle, and paucity of circular
muscle.
C — columnar epithelium
CM = circular muscle L — lumen
CT - connective tissue LM - longitudinal muscle
W - tissue of the wall

H — heart muscle

THE V. ‘
HE VELIGER, Vol. 10, No. 3 [JecLa « GREENBERG] Plate 38

a ee

sts a rstehiay

Vol. 10; No. 3

THE VELIGER

Page 261

(STAsEK, 1963) distribution is dissimilar; apparently the
most primitive Category I arose in the Cambrian and
divergence began very early.

Other morphological features of the Anisomyaria and
Arcidae are well known (e. g. Morton & YONGE, 1964).
In addition, one of us (M.J.G., unpublished) has ob-
served that, in the Mytilidae, Ostreidae, Pectinidae, and
Pteriidae, the auricles are glandular and communicate
with each other posteriorly and ventral to the ventricle.
This condition was not found in the Pinnidae.

The uniform rectal structure of the Unionidae, taken
together with their freshwater habitat suggests that this
rectum might be an adaptation to the hyposmotic environ-
ment. In particular, the presence of sinuses suggests that
the typhlosole might function in osmoregulation. PicKEN
(1937) and Potts (1954) found that blood was filtered
through the heart to the pericardial cavity. Possibly filtra-
tion also occurs into the rectum facilitated by the blood
sinuses in the typhlosole. The typhlosole epithelium, in
contrast to the remaining rectal lining, stains intensely
(Plate 39, Figure 7). This characteristic also suggests
some unique function of this region. However, the rectums
of other fresh- and brackish water bivalves have been
studied: Polymesoda caroliniana (Bosc, 1802) and
Sphaerium sp., both from the superfamily Sphaeriacea,
and Rangia cuneata (Gray, 1837), a mactrid. In all of
these species, the rectum was thin-walled, relatively
smooth, and lacked, altogether, a typhlosole. Thus, while
the unionid rectum might have evolved as an environ-
mental adaptation, this has happened only once. Although
Rangia cuneata is subjected to considerable osmotic stress,
the rectum resembles closely those of Spisula solidissima
and Mactra fragilis, the other marine members of the
family studied.

The important, mandatory function of the rectum is
preparation and propulsion of faecal material. Therefore
the shape and composition of the faeces are useful indica-
tors of the functional morphology of the rectum. In a
recent consideration of molluscan faeces, KORNICKER
(1962) has outlined the structure and distribution of
five major kinds of bivalve faecal pellets and has sug-
gested evolutionary trends connecting them. His generali-
zations are borne out by ArAKAWwa’s later description of
the faeces from a number of species (1963; 1965).

Protobranch faeces are always pellets which are broken-
off pieces of sculptured rods. In the Anisomyaria, 60%
of the species studied had either sculptured or ribbonlike
faecal pellets. The ribbonlike faeces are also sculptured
and have ventrally curved edges. They are, of course, casts
of the unique flattened rectums previously described as
characteristic of this order. Heterodont faeces are usually

either unsculptured or shapeless, and oval pellets appear
only in this order. KornicKER points out that the sedi-
ments in sculptured faeces are segregated by particle size
in different portions of the faecal rod, whereas the compo-
sition of unsculptured, shapeless, and oval pellets is uni-
form. He concludes that sculptured faeces with segregated
sediments are primitive, while unsculptured, oval, and
shapeless faeces, with their uniform composition, are
derived forms.

Sculpturing and particle distribution are largely depen-
dent upon the mechanism by which faeces are propelled
through the rectum. If propulsion is by means of ciliary
tracts, with little muscle involvement, then there can be
no gross mechanical activity accompanying the passage of
the faecal rod. Consequently, the distribution of sedi-
ments determined by the ciliary sorting systems of the
stomach and gut will be undisturbed; similarly, the surface
sculpturing of the rod will be preserved. On the other
hand, a muscular rectum would probably obliterate sur-
face sculpturing, and also thoroughly mix the sediment.
If the faeces are of appropriate consistency, and if the
peristaltic contractions are sufficiently powerful, the gut
contents will be molded into a string of oval pellets. The
distribution of muscle among bivalve rectums is well
correlated with the distribution of sculptured and un-
sculptured faeces. Muscular rectums are found mainly
among the Heterodonta. Protobranch and anisomyarian
rectums have, characteristically, thin walls with little
muscle.

What is the functional significance of the interfamily
differences in rectal structure? A reasonable supposition
is that they should be related in some way to feeding and
digestive mechanisms. The possibility is best examined in
the Heterodonta, since the rectum in this order is mostly
notable for its variability from family to family. Thus, one
might expect to find some relationship between rectal
structure and stomach-type (PurcHoNn, 1963), gill-palp
association (STASEK, 1963), habitat, or diet. None was
observed. Thus, while subtle species specific differences in
rectal physiology occur, and while they must be related to
structural differences, probably at the submicroscopic
level (GREENBERG, 1966), there are no such relationships
at the level of gross histology. The rectum in Bivalvia is,
therefore, a non-adaptive structure which has undergone
little change since the divergence of the families.

ACKNOWLEDGMENTS

We arc indebtcd to the staffs of the marine laboratories at
Friday Harbor (University of Washington) and Alligator

Page 262

Harbor (Florida State University) for their kind coopera-
tion and to Mrs. J. Goldstein and D. Nelson for their
technical assistance.

This study was supported by Research Grant HE-06291
from the National Heart Institute, U.S. PH.S. Additional
funds were obtained from the University of Illinois
Research Council and the Illinois Marine Biological
Association.

LITERATURE CITED

Assott, RoBERT TUCKER
1954. American seashells. Princeton, New Jersey. D. van
Nostrand Co., Inc.; xiv+541 pp.; 100 figs.; 40 plts.
ARAKAWA, KoHMAN Y.
1963. Studies on the molluscan faeces (I).
Biol. Lab. 11: 185 - 208
1965. Studies on the molluscan faeces (II).
Biol. Lab. 13: 1-21; plts. 1-6
Cox, Leste REGINALD
1960. | Thoughts on the classification of the Bivalvia. _ Proc,

Publ. Seto Mar.

Publ. Seto Mar.

Malacol. Soc. London 34 (2): 60-88
Dakin, WILLIAM JOHN
1909. Pecten. L.M.B.C. Memoirs XVII. The

Univ. Press of Liverpool. 1 - 136; plts. 1-9

THE VELIGER

Vol. 10; No. 3

Ga.tsorF, Paut S.
1964. The American oyster Crassostrea virginica GMELIN.
Fishery Bull. Fish & Wildlife Serv., U.S. Bur. Comm. Fish. 64:
480 pp.; 400 figs.
GREENBERG, MIcHAEL JOHN
1966. Species specific effect of acetylcholine on bivalve rectums.
Science 154: 1015 - 1017
Guyver, MICHAEL FREDERIC
1953. Animal micrology.
GuUTHEIL, Fritz
1912. Uber den Darmkanal und die Mitteldarmdriise von
Anodonta cellensis Scuror. Zeitschr. fiir wissenschaftl.
Zool. 99: 444 - 538
von Harrner, KoNSTANTINE
1957, Untersuchungen tiber die Anatomie der in Schwammen
lebenden Muschel Vulsella. Zool. Jahr., Abt. Anat.
Ontog. d. Tiere 76: 371 - 422
Humason, GrRETCHEN Lyon
1962. | Animal tissue techniques.
KELLOGG, JAMES LAWRENCE
1890. A contribution to our knowledge of the morphology of

lamellibranchiate mollusks. Bull. U.S. Fish. Comm. X: 389
to 486

Kornick_Er, Louts S.

1962. Evolutionary trends among mollusk fecal pellets.
Journ. Paleontol. 36: 829 - 834

Univ. Chicago Press

Freeman, San Francisco

Explanation of Plate 39

Rectums with Voluminous Typhlosoles

Figure 7: Actinonaias carinata. Characteristic schizodont rectum.
Note large vascular channels, lined by endothelium, in the typhlo-

sole.

Figure 8: Atrina rigida. Note modified anisomyarian rectum; com-

pare with Figure 3, Plate 37.
CG — columnar epithelium
CH — vascular channels
H — heart muscle

L - lumen
T — typhlosole
W - tissue of the wall

Explanation of Plate 40

Solid Typhlosoles in the Rectum of Dinocardium robustum

Figure 9:

Low magnification photograph. Note thick wall with

extensive, dense musculature in the periphery of the wall, and
many moderately large, solid typhlosoles.

Figure 10:

Details of structure of typhlosole. Note penetration

of connective tissue fibers and muscle cells into the typhlosole, and
lack of vascular channels; compare with Plate 39, Figure 7.

C — columnar epithelium

CTF - connective tissue fibers

H — heart muscle

L — lumen
M — muscle cell
T — typhlosole

W — tissue of the wall

Tue VELIcER, Vol. 10, No. 3 [JEcLA & GREENBERG] Plate 39

Figure 8

Tue VELIGER, Vol. 10, No. 3 [JEcLA « GREENBERG] Plate 40

Figure 10

Vol. 10; No. 3

THE VELIGER

Page 263

Morton, Joun Epwarp & CHARLES MauricE YONGE

1964. Classification and structure of the Mollusca. In: Phys-
iology of Mollusca (ed.: Kart Mitton Witpur « CHARLES
Maurice YoncE) 1: 1-58. Academic Press New York

Mot Ley, Huriey LEE
1933. Histology of the fresh-water mussel heart with reference
to its physiological reactions. Journ. Morphol. 54: 415 - 427
Nystrom, RicHarp ALAN
1967. | Spontaneous activity of clam intestinal muscle.
Comp. Biochem. Physiol. 21: 601 - 610
PELSENEER, PAUL

1906. Mollusca. In: A treatise on Zoology (ed.: E. Ray

LANKESTER) part V: 1-355 Black, London
Puiuis, J. W.

1966. Regulation of rectal movements in Tapes watlingi.

Comp. Biochem. Physiol. 17: 53 - 70
PickEN, LAuRENCE E. R.

1937. | The mechanism of urine formation in invertebrates. II.
The excretory mechanism in certain Mollusca. Journ. Exp.
Biol. 14: 20 - 34

Ports, WiLL1aM TayLor WINDLE

1954. The rate of urine production of Anodonta cygnea.

Journ. Exp. Biol. 31: 614-617
Purcuon, R. DENISON

1963. Phylogenetic classification of the Bivalvia, with special
reference to the Septibranchia. Proc. Malacol. Soc. London
35: 71 - 80

STAsEK, CHARLES ROBERT

1963. Synopsis and discussion of the association of ctenidia and
labial palps in the bivalved Mollusca. The Veliger 6 (2) :
91-97; 5 text figs. (1 October 1963)

VON STUDNITZ, G.

1931. Die Morphologie und Anatomie von Lima inflata, der
Feilenmuschel, nebst biologischen Untersuchungen an Lima
hians Gmel. Zool. Jahr., Abt. Anat. Ontog. Tiere 53:
199 - 316

THIELE, JOHANNES
1935. Handbuch der systematischen Weichtierkunde, Part 2:
779 - 1154. Gustav Fischer, Jena
Voct, Cart «& EMILE YUNG
1888. Traité d’anatomie comparée pratique.
Waite, KaTHLeEEN M.

1937. Mytilus. L. M. B.C. Memoirs XXXI. The

Univ. Press of Liverpool. 1-117; plts. 1-10
Yonce, Cuartes Maurice

1923. Studies on the comparative physiology of digestion. I.
The mechanism of feeding, digestion, and assimilation in the
lamellibranch Mya. Journ. Soc. Exp. Biol. 1: 15 - 63

1926. Structure and physiology of the organs of feeding and
digestion in Ostrea edulis. Journ. Mar. Biol. Assoc. U.K.
14: 295 - 386

1941. | The protobranchiate Mollusca: a functional interpreta-
tion of their structure and evolution. Phil. Trans. Roy. Soc.
London B230: 79 - 147

Reinwald, Paris

Page 264

THE VELIGER

Vol. 10; No. 3

The Generic Classification of Cowries

BY

FRANZ ALFRED SCHILDER

University of Halle, German Democratic Republic

IN A PREVIOUS PAPER (SCHILDER, 1966) I have published
a survey of the superfamilies, families, subfamilies, and
tribus (infrafamilies) of the “true” cowries (Cypraeidae)
and of the closely allied groups of mollusca, the “false”
cowries: Triviidae, Pediculariidae, and Ovulidae. In the
present paper this survey will be expanded to a list of
all genera and subgenera belonging to these higher taxa,
with their type species and synonyms.
A similar list has been published about a quarter of a
century ago (ScuiLper, 1941: 65-68), but since that time
many new generic names have been established. The main

morphological differences of allied taxa have been discussed
two years earlier (SCHILDER, 1939).

In the present list the names of SUPERFAMILIES,
Famities, Subfamilies and (Tribus) have been distin-
guished by type styles, indicating the taxonomic category;
generic and (subgeneric) names have been enumerated as
coordinated names in a taxonomic order thought to be a
phylogenetic order also; but to the (subgenera) the genus
to which they should be united has been added in square
brackets [ ]. Synonyms have been listed in chronological
order.

The nine subfamilies have been distinguished by the
digits 1 to 9; a second digit indicates the tribus; these bold
face figures facilitate the spotting of names enumerated
in the alphabetical index.

In the systematic list the following symbols have been
used:

+ the dagger designates extinct taxa

|| names invalid by homonymy

/ (preceding the name) not valid, as established by
an invalid author or in an invalid way, and never
adopted in a valid way

(separating the name and the author) wrong inter-
pretation by adopting a wrong type species
objective synonym by identity of the type species
subjective synonym, as the different type species
now are supposed to belong to the same subgenus
or indivisible genus

misspelled generic names, invalid authors, and
names of tribus have been put in parentheses

—
—

[...] opinions of the author of the present paper have
been put in square brackets

The type species of genera and subgenera have been
distinguished as follows:

M monotypical

TT _ type species by absolute tautonymy

OD type species by original designation

SD type species. by subsequent designation with the
author added

ex. example only, not valid type species

The meaning of other abbreviations is as follows:

> < type species cited between these symbols

err. errore, name misspelled by mistake

em. name emended by a later author on purpose

pro name replacing another name, mostly on account of
homonymy

nomen nudum (invalid)

nomen oblitum: not cited for past 50 years (invalid)
used as a synonym only (invalid)

manuscript name only (invalid)

nud.

obl.

syn.

MS

The two papers of JoussEAUME both published in 1884

should be distinguished: 1884a (Naturaliste, p. 414) undoubt-

edly is prior to 1884b (Bull. Soc. Zool. France 9: 81), as

1884a has been published February 15 1884, while 1884b has

been read only before the Society on February 12 and therefore

has been printed much later. Nevertheless, concerning 4 differ-

ences in spelling of new names (Cribraria, Mauxiena, Trona,

Zonaria) I agree with all later writers in adopting the names

of 1884b as valid, since 1884a is a popular excerpt only which
contains many errors in spelling of species names.

SYSTEMATIC LIST

TRIVIACEA TROSCHEL, 1863

TrIVODAE TROSCHEL, 1863

1 Eratoinae ScHiLpErR, 1927

+ (Johnstrupiini) Scuiper, 1939

{Johnstrupia Ravn, 1933
>M fJohnstrupia tfaxensis Ravn, 1933 <

Vol. 10; No. 3

+Eratodium Koroskov, 1955
>M Erato (?)tmedius Isazva, 1933 <
[possibly no genus of Triviacea at all]

12 } (Eratotriviini) ScHILDER, 1936

}Eratotrivia Sacco, 1894
> OD Cypraea ||tcrenata Desuayes, 1835 =
}Eratotrivia tcrenularis ScHILDER, 1927 <

13 (Eratoini) Scuitper, 1927
{Proterato ScHILDER, 1927
> OD Erato tneozelanica Suter, 1917 <
Sulcerato Fintay, 1930 [Proterato sub-
genus]
> OD Erato fillota Tate, 1890 <
[=] Lachryma (Humpurey, MS,
SowErRBy, 1837 syn.) Cotton & Gop-
FREY, 1932
> OD Erato (trifasciata HumMpHREY, MS =)
Erato lachryma SowErRBy, 1832 <
Eratoena IrEDALE, 1935 [Proterato
subgenus]
> OD Ovulum corrugatum Hinps, 1844 <
Cypraeerato SCHILDER, 1932 [Proterato
subgenus |
> OD Erato bimaculata Tate, 1878 <
[=] Lachryma Irepate, 1931 nud.
>M Lachryma bisinventa IREDALE, 1931 <
+Archierato ScHILDER, 1932
> OD Erato (Erato) +pyrulata Tate, 1890 <
Eratopsis HORNES & AUINGER, 1880
[Erato subgenus]
>M Eratopsis tbarrandei HORNES & AUINGER, 1880 <
Erato Risso, 1826
> M Marginella tcypraeola Broccut, 1814

Hespererato ScuiLper, 1932
> OD Ovulum vitellinum Hinps, 1844 <

2 Triviinae TROSCHEL, 1863

21 (Triviellini) ScHILpER, 1939
}Willungia Powe, 1938
> OD fWillungia ttasmanica PowE Lu, 1938 <
+Semitrivia CossMANN, 1903
> OD Trivia terugata Tate, 1890 <
Fossatrivia IREDALE, 1931
>M Trivia caelatura Hepiey, 1918 <
+Nototrivia ScHILpER, 1932
> M, SD Scuirver 1932 Foss. Cat.: Cypraea (Trivia)
tavellanoides McCoy, 1867 <
Triviella JoUSSEAUME, 1884a
> SD JousszaumE, 1884b: Cypraea |loniscus LAMaRcK,
1810 = Cypraea aperta Swainson, 1822 <
+Prototrivia Scuitper, 1941
>M Cypraea (Trivia) twetherellii Epwarps, 1854 <

THE VELIGER

Page 265

22 (Triviini) TroscHEL, 1863

Trivia BroperiP, 1837
> SD Gray, 1847: Cypraea europaca Montacu, 1808 =
Cypraea monacha Costa, 1778 <

(em.) Trivea Swainson, 1840
[=] || Coccinella (Sowrrsy, 1823
syn.) HERRMANNSEN, 1846
> OD Cypraca arctica Monracu, 1803 <
Sulcotrivia ScuiLpeER, 1933 [Trivia
subgenus ]
> OD Cypraca tdimidiata Bronn, 1831 <
23 (Pusulini) ScuiLpEr, 1936
Pseudotrivia ScHILDER, 1936
> OD Trivia sibogae ScHEpMaN, 1909 <
Ellatrivia IREDALE, 1931 [Niveria sub-
genus |
> OD Triviella merces TREDALE, 1924 <
Cleotrivia IREDALE, 1930 [Niveria sub-
genus]
> OD Cypraca pilula Kiener, 1843 <

Niveria JouSSEAUME, 1884a
> M, SD JousseaumE, 1884b: Cypraca ||nivea Gray, MS =
Sowersy, 1832 = Trivia nix ScuHiLpER, 1922 <
Trivirostra JOUSSEAUME, 1884a, 1884b
> SD Roserts, 1885: Cypraea oryza LAMaRcK, 1810 <
Dolichupis IREDALE, 1930 [Pusula sub-
genus]
> OD Cypraca producta Gasxoin, 1836 <
[=] Trivellona IrEDALE, 1931
>M Trivellona excelsa IREDALE, 1931 <
Pusula JOUSSEAUME, 1884a, 1884b
> SD Roserts, 1885: Cypraca radians LaAMarck, 1810 <
(err.) Pustula Roperts, 1885
(err.) Vusula CossMann, 1896

30 PEDICULARIDAE ADAMS & Apams, 1854
+Semicypraea ScHILpEr, 1936
> OD Cypraea koninckiti Rovautt, 1850 <
+Cypracogemmula VREDENBURG, 1920
>M Trivia ||tscabriuscula Kornen, 1890 =
Cypraca tliliputana ScurtpER, 1922 <
Pediculariona IREDALE, 1935 [Pedicu-
laria subgenus]
>M Pedicularia stylasteris HEDLEY, 1903 <
?{==] Dentiora Prase, 1862
> M Dentiora rubida PEasE, 1862 = [possibly]
Pedicularia pacifica PEASE, 1865 <
Pediculariella Tureve, 1925 [Pedicu-
laria subgenus]
> OD Pedicularia californica NEwcoms, 1864 <
Pedicularia Swainson, 1840
>M Pedicularia sicula Swainson, 1840 <

Page 266

==|| Thyreus Pupp, 1844
>M Thyreus paradoxus Puiuipri, 1844 =
Pedicularia sicula Swainson, 1840 <

CyPRAEACEA Gray, 1824

CyPRAEIDAE Gray, 1824

4 Bernayinae ScHILDER, 1927

41 (fArchicypraeini) ScHILpeER, 1927
[=] (}Mandolinini) ScuitpEr, 1936

}Palaeocypraea SCHILDER, 1928
> OD }Cypracacites }spiratus ScHLOTHEIM, 1820 <
[=]||+Palaeocypraea Sayn, 1932 [nov.

genus]!
> SD Scuirper, 1936: Cypraea (}Palacocypraea) ||tantiqua
Sayn, 1932 = +Palaeocypraca }+primigenia SCHILDER,

1936 <
(==}Porcellanites (WALLER, 1747;
ARGENVILLE, 1757; SCHROTER,
1782, 1783), ScHLoTHEIM, 1813
nud.
>M +FPorcellanites tseelandicus ScHLtoTHEm, 1813 nud.,
which is = }+Cypraeacites tspiratus ScHLOTHEIM, 1820
according to Bronn, 1848 <

}Pustulariopsis ScuitpER, 1941 [}Pa-
laeocypraea subgenus]
> OD Cypraea + pustulifera Parona, 1909 <
tArchicypraea SCHILDER, 1926
> OD Cypraea tlioyi Bayan, 1870 <
{+Mandolina JoussEAUME, 1884a
>M, SD JousszaumeE, 1884b: Cypraea ||tgibbosa

Borson, 1820 = Cypraea ({Mandolina) ||{gibbosa
Borson, 1820 var. pergibba Sacco, 1894 <

42 (Bernayini) ScuitpEr, 1927
[=] (Cypraeorbini) ScuiLpeEr, 1927
[=] (Zoilini) IREDALE, 1935

Protocypraea ScHILDER, 1927 [Berna-

ya subgenus]
> OD tEocypraea torbignyana VREDENBURG, 1920 <

Bernaya JOUSSEAUME, 1884a
> SD JousseaumE, 1884b: Cypraea {media Desuayes, 1835 <
(em.) Bernayia Cossmann, 1889

+Barycypraea ScuiLpeEr, 1927
> OD Cypraea (Aricia) tcaputviperae Martin, 1899 <

+A frocypraea ScuiLvER, 1932 [possibly
no Cypraeidae at all]
> OD Cypraea tchubbi Rennie, 1930 <

Zoila JOUSSEAUME, 1884a
> SD JoussEaumE, 1884b: Cypraca scottii BRoDERIP,
1831 = Cypraea friendit Gray, 1831 <

THE VELIGER

Vol. 10; No. 3

+Gigantocypraea ScuitpER, 1927 [Zo-
ila subgenus]
> OD Cypraea tgigas McCoy, 1867 <

+Cypraeorbis Conran, 1865
>M Cypraca tsphaeroides Conran, 1847 <

Akleistostoma GarDNER, 1948 [}Sipho-
cypraea subgenus]

> OD Cypraea tcarolinensis Conran, 1841 <

[=] Muracypraea Wooprtne, 1957
> OD Cypraea mus LinnagEus, 1758 <

?[=] +Cypraeactaeon Wurrte, 1887

> M fCypracactaeon tpennai [em.] Wuite, 1887

[figs. 1 and 2 only, according to ScuitpEr, 1927] <
+Siphocypraca HEILprin, 1887

> M tSiphocypraca tproblematica Hemprin, 1887 <

43 ({Gisortiini) ScHiLpErR, 1927

{Megalocypraca ScHILDER, 1927 [}+Gi-
sortia subgenus]
> OD +Gisortia ({Megalocypraca) tovumstruthionis
SCHILDER, 1927 <
[=]}Pseudogisortia ScumpER, 1941
> OD }Gisortia tbenedicti VREDENBURG, (1920) 1927 <

{Gisortia JouSSEAUME, 1884a
> SD Joussraume, 1884b: Ovula tgisortiana Passy, 1859 <

{Vicetia Fastant, 1905
> M Ovula thantkeni LEFEvRE, 1878 <

5 Cypraecinae Gray, 1824

51 (Cypraeini) Gray, 1824
[=] (Porcellanini) Roserts, 1870
[=] (Talpariini) Scuitper, 1936
[=] (Mauritiini) SreapMan «& Cotton,

1946

{+Miolyncina ScHILpER, 1932
> OD Cypraca tsubovum Orsicny, 1852 <

Trona JOUSSEAUME, 1884b
> OD Cypraca stercoraria LINNAEUS, 1758 <
(err.) Etrona JouSSEAUME, 1884a
ex. Cypraea stercoraria LINNAEUS,
1758
[=]||+Basterotia JoussEaumME, 1884b
> OD Cypraca tleporina Lamarck, 1810 <
(err.) +Bastorotia JOUSSEAUME, 1884a
ex. Cypraea {leporina LAMARCK,
1810
= }Cavicypraca CossMANN, 1896
pro || + Basterotia JousSEAUME,
1884b

Macrocypraea SCHILDER, 1930
> OD Cypraca exanthema Linnazus, 1767 =

Vol. 10; No. 3

Cypraea zebra LINNAEUS, 1758 <
(= Cypraea/Lamarck, 1801 ex.
Cypraea exanthema LINNAEUS, _
1767)
(= Cypraea/JoussEAuME, 1884b
SD JoussEAuME, 1884b: Cypraea
exanthema LinnagEus, 1767)
(= Erythraea (BarReEvier, 1714;
Sowersy, 1839 syn.) MoOrcu,
1877; /ScHILDER, 1924)
pro || Tigris TRoscHEL, 1863
therefore
> SD Scuiiper, 1924: Cypraea exanthema LinnagEus, 1767 =
Cypraea zebra LinnakEus, 1758 not valid <

Leporicypraea IREDALE, 1930 [Mauri-
tia subgenus]
> OD Cypraea mappa Linnaeus, 1758 <
(= Cypraca/LamMarck, 1799 ex.
Cypraea mappa LINNAEUS, 1758)
(= Cypraea/CossMAnn, 1903: SD
CossMann, 1903: Cypraea map-
pa Linnaeus, 1758)

Arabica JoussEaAuME, 1884a [Mauri-

tia subgenus]
> TT, SD JoussEaumE, 1884b: Cypraea arabica LINNAEUS,
1758 <

(err.) Arabia JoussEAUME, 1886
(=Cypraea/Swatnson, 1840, ex.,
SD Scuirper, 1924: Cypraea ara-
bica LinnaEvs, 1758)

Mauritia TRoscHEL, 1863
> SD Cossmann, 1903: Cypraea mauritiana LINNAEUS,
1758 <
= Maurina JoussEAuME, 1884a
>M Cypraea mauritiana Linnazus, 1758 <
= Mauxiena JoussEAUME, 1884b
[err. for Maurina?]
> M, OD Cypraea mauritiana LinNAEuS, 1758 <
= Etronsa JoussEAUME, 1886 [err.
for Etrona?]
>M Cypraea mauritiana LinnaEus, 1758 <
(= Cypraea/Cossmann, 1889, SD
CossMANN, 1889: Cypraea mau-
ritiana LINNAEUS, 1758)
(== Peribolus (ApaNson, 1757)
BLAINVILLE, 1824 (obl.) M Potan
ApANSON, 1757 ==Cypraea mau-
rittiana LINNAEUS, 1758 juvenis
[adopted as generic name by Ku-
RoDA, 1960 and Azuma, 1960])

THE VELIGER

Page 267

Talparia TROoSCHEL, 1863
> M, SD Scuiper, 1924: Cypraca talpa LinNAEus, 1758 <
(= Cypraea/MEtviti, 1888, SD
MELvILL, 1888: C'ypraca talpa
Linnaeus, 1758)

Cypraea Linnaeus (1740), 1758
> SD Montrorrt, 1810: Cypraca tigris LINNAEUS, 1758 <
(err.) C'yproea SERRES, 1822
(err.) Cipraea CutajeE, 1827
(em.) Cyprea Costa, 1776, ABEL,
1787, Montrort, 1810
(em.) ||Cypria Jerrreys, 1867
[=] Cypriarius DumeriL, 1806 pro
Cypraea LINNAEUS, 1758
(em.) Cypraearius Froriep, 1806
=||Tigris Troscuer, 1863 [preoccu-
pied by (Linnaeus, 1735) OKEN,
1816
> TT, SD Cossmann, 1903: Cypraea tigris Linnazus, 1758 <
= Erythraea (Barrevurr, 1714) So-
WERBY, 1839 syn.) MOrcu, 1877
pro Tigris TRoscHEL, 1863 [nec
Kein, 1753]
(err.) (Aerythraea ADANSON, 1757)
= Vulgusella JousseauME, 1884a
> SD JoussEaumE, 1884b: Cypraea tigris Linnarus, 1758 <
(err.) Vulpicella CossMANnN, 1889,
1896, OppENHEIM, 1894 [correc-
ted by Cossmann, 1906]
[=] Pantherinaria Sacco, 1894
> OD Cypraca pantherina SoLanper, 1786 <
Lyncina TroscHEL, 1863
> SD Tryon, 1883: Cypraca lynx Linnaeus, 1758 <
[=]]| Porcellana (Rumpuius, 1705)
Linck, 1783, Roserts, 1870
> SD Jousszaume, 1884b: Cypraca argus Linnagus, 1758 <
[=] Ponda JoussEAuME, 1884a
> SD JoussEaumE, 1884b: Cypraca achatina (SoLAN-
DER) Perry 1811 = Cypraca ventriculus LAMARCK,
1810 <
([=] Lyncina/ScuiLpEr, 1924)
> SD Scuitper, 1924: Cypraea carneola Linnaeus, 1758 <
[==] Prolyncina ScutLpErR, 1927
> OD Cypraea reevei Sowersy, 1832 <
[=] Callistocypraca ScuiLpER, 1927
> OD Cypraca aurantium (Martyn, 1784 =) GMELIN,
1791 <
[=] Arestorides IREDALE, 1930
> OD Cypraea argus LinaEus, 1758 <
(err.) Aristorides Cate, 1967
[=] Mystaponda IREDALE, 1930
> OD Cypraca vitellus Linnagus, 1758 <

Page 268

THE VELIGER

Vol. 10; No. 3

52 (Luriini) ScuitpEr, 1932

+Jousseaumea Sacco, 1894 [em. by
ScHILper, 1927 and ICZN Opin.
673, 1963]
> OD Cypraea tsublyncoides OrBicny, 1852 [= Cypraea
tdiluviana Gray, 1824] <

(=tJousseaumia Sacco, 1894, original
spelling)
{Fossacypraea SCHILDER, 1939 [Jous-
seaumea subgenus]
> OD Cypraea thieroglyphica ScuiLvER, 1923 <

Chelycypraea ScuILpER, 1927
> OD Cypraea testudinaria LinNAEus, 1758 <

Luria JoussEAuME, 1884a
> SD JoussEaumE, 1884b: Cypraea lurida LinNaEus, 1758 <
[=] Tessellata JousSEAUME, 1884a
[em. by JoussEAuME, 1884b]
> TT, M Cypraea tessellata Swainson, 1822 <
[=] Bastltrona IrREDALE, 1930
> OD Cypraea isabella LinNAEus, 1758 <

6 Erroneinae SCHILDER, 1927
61 (Zonariini) ScuiLpErR, 1932

+Prozonarina ScuiLpER, 1941 [}Zona-
rina subgenus |
> OD Cypraea tbrocchii Desuayes, 1844 <

+Zonarina Sacco, 1894
> OD Cypraea t pinguis/Sacco, 1894 [nec GRATELOUP,
1845] = tlongovulina Sacco, 1894 <
Schilderia Tomuin, 1930
pro|| Globulina Cerruti, 1911; OD
Cypraea tutriculata LAMARCK,
1810
= || Globulina Crruut, 1911

>M Globulina tinfernot Crrutit, 1911 = Cypraea tutriculata
Lamarck, 1810 juvenis <

Zonaria JOUSSEAUME, 1884b
> OD Cypraea zonata (CHEMNITZ, 1788) Lamarck,
1810 [= Cypraea zonaria GmELin, 1791] <
(err.) Zonatia JOUSSEAUME, 1884a
ex. Cypraea zonata (CHEMNITZ, 1788) [= zonaria
Gmeun, 1791]

[=] Pseudozonaria ScuitpeEr, 1927
> OD Cypraea arabicula LaMarcx, 1810 <
Neobernaya ScHILvER, 1927 [Zonaria

subgenus |
> OD Cypraea spadicea Swainson, 1823 <

62 (Cypraeovulini) ScuitpeEr, 1927
[=] (Umbiliini) Scuitper, 1932

}Notoluponia ScuiLvER, 1935
> OD {Notoluponia tmurraviana telegantior ScutLpER, 1935 <

Luponia Broverip, 1837 [Cypraeovula
subgenus]
> M Cypraea algoensis Gray, 1825 <
(err.) Lupina Bronn, 1853
(err.) Luponaria WEINKAUFF, 1881
(err.) Luponica WEINKAUFF, 1881
(err.) Luperia OPPENHEIM, 1901

[=] Gaskoinia Roserts, 1870
> M Cypraea edentula Gray, 1825 <

Cypracovula Gray, 1824
> M, SD Gray, 1828: Cypraca capensis Gray, 1828 <
(err.) Cypreovula FLEMING, 1828
(err.) Cyprovula Gray, 1828, 1840,
1847
(em.) Cypraeovulum SowrrBy, 1842
(em.) Cypraeova Swainson, 1840

(em.) Cypraeovum Scuauruss, 1869

Guttacypraea IREDALE, 1935 [Notocyp-
raea subgenus]
> OD Cypraca pulicaria REEVE, 1846 <

Notocypraea ScHILpER, 1927
> OD Cypraea piperita Gray, 1825 <
[=] Thelxinovum TREDALE, 1931
> OD Thelxinovum mélleri Treva, 1931 [= Noto-
cypraca emblema IREDALE, 1931] <

{+Rhynchocypraea CossMann, 1898
[Umbilia subgenus]
> OD Cypraca (Luponia) tleptorhyncha McCoy, 1877 <

Umbilia JoussEAUME, 1884a
> M, SD Joussraume, 1884b: Cypraea || umbilicata
Sowersy, 1825 = Cypraca hesitata IREDALE, 1916 <
[=]t Rhynchocypraea /CossMann,
1903
> SD Cossmann, 1903: +Rhynchocypraca tloxorhyncha
[sic] /Cossmann, 1903 fig. [= Umbilia +maccoyi
Scuitper, 1932 nec Cypraca ttoxorhyncha Tate, 1890] <

}Palliocypraea CossMann, 1906 [Um-

bilia subgenus]
> OD Cypraca (Aricia) tgastroplax McCoy, 1875 <

63 (Erroneini) ScHiLpeEr, 1927
[=] (Adustini) SteapMan & Corton, 1946

Gratiadusta IrEDALE, 1930 [Erronea
subgenus]
> OD Cypraca pyriformis Gray, 1824 <
=TIpserronea IREDALE, 1935
> OD Ipserronea problematica IrEpALE, 1935 = Cypraca
pyriformis Gray, 1824 juvenis [according to
TREDALE, 1939] <
[=] Ficadusta Hase & Kosuce, 1966
> OD Cypraea pulchella Swainson, 1823 <

Vol. 10; No. 3

Adusta JousSEAUME, 1884a [Erronea

subgenus]
> TT, SD JoussEaumeE, 1884b: Cypraca adusta
(CHEmMnitTz, 1788) = Cypraea adusta LAMARCK,

1810 <
[=] Solvadusta IrEDALE, 1935
> OD Gratiadusta vaticina IREDALE, 1931 <

Erronea TROSCHEL, 1863
>M, SD JoussEaumeE, 1884b: Cypraca errones
Linnaeus, 1758 <
[=] Palangerosa IREDALE, 1930
> OD Cypraea cylindrica Born, 1778 <

Melicerona IrREDALE, 1930 [Erronea

subgenus]
> OD Cypraea listeri Gray, 1824 <

Notadusta ScuiLpeEr, 1935
> OD Notadusta tvictoriana ScuHitvER, 1935 <

Palmadusta IREDALE, 1930
> OD Cypraea clandestina LinnarEus, 1767 <
[=] Evenaria IrEDALE, 1930
> OD Cypraea asellus Linnazus, 1758 <
(err.) Evanaria STEADMAN & COTTON,
1943 [always]

Purpuradusta ScHILDER, 1939 [No-
vember] [Palmadusta subgenus]
> OD Cypraea fimbriata GMEtin, 1791 <
[=] Opponaria IrREDALE, 1939 [De-

cember|
> OD Cypraea minoridens MEtvitx, 1901 [“minoridens
series” | <
[=]Cupinota IrepatE, 1939 ([De-
cember]

> OD Cypraea macula Ancas, 1867 <

Blasicrura IREDALE, 1930 [Bistolida
subgenus]
> OD Cypraea rhinoceros SouvERBIE, 1865 <
[=] Eclogavena IREDALE, 1930
> OD Cypraea coxeni Cox, 1873 <
[==] Talostolida IrEDALE, 1931
> OD Cypraea teres GmEtin, 1791 <

Bistolida CossMANnN, 1920
pro || Stolida JoussEAUME,
1884b
= || Stolida JoussEAuME, 1884a
> TT, SD JoussEaumeE, 1884b: Cypraea stolida
LinnaEus, 1758 <
= Derstolida IREDALE, 1935
> OD Derstolida fluctuans IrEpaALE, 1935 [= Cypraca
stolida LINNAEUS, 1758] <<

Ovatipsa IREDALE, 1931
> OD Cypraca chinensis GMEuin, 1791 <

THE VELIGER

Page 269

Cribrarula STRAND, 1929
pro || Cribraria JoussEAUME,
1884b
(err.) Criraria JouUSSEAUME, 1884a
> TT Cypraca criraria [err. pro cribraria] LinNarus, 1758 <
= || Cribraria JoussraumeE, 1884b
> TT, OD Cypraea cribraria Linnarus, 1758 <
= Nivigena IREDALE, 1930
> M, OD Nivigena melwardi Irepate, 1930 [= variety
of Cypraeca cribraria LinnaEus, 1758] <

7 Erosariinae ScHILpDER, 1924

71 (Pustulariini) Scumper, 1932
[=] (Cypraeacitini) ScuitpEr, 1930

[=] (Austrocypraeini) IREDALE, 1935

[=] (Conocypraeini) ScuiLver, 1936

Annepona IREDALE, 1935 [Pustularia
subgenus }
> OD Cypraca || annulata Gray, 1828 = Pustularia
mariae SCHILDER, 1927 <
(=Epona/ (WeEINKAuFF, 1881)
IREDALE, 1939)
> SD Wernxaurr, 1881: Cypraca || annulata Gray,
1828 according to IREpALE, 1939 to be proved [in
WEINKAUFF, 1881 no type species; || annulata Gray,
1828 is the first named species only] <

Pustularia Swainson, 1840

> M Cypraca cicercula Linnacus, 1758 <
(err.) Postularia Coccont, 1873
= Epona ApaMs & Apams, 1854

> SD ME vixt, 1888: Cypraca cicercula GMELIN, 1791
= Linnaeus, 1758 <
(err.) Eponia Rossiter, 1882
(err.) ||Globularia Sowersy, 1842, ex.
Cypraca globulus LINNAEUS,
1758

Ipsa JOUSSEAUME, 1884a, b [Pustula-
ria subgenus]
>M Cypraca childreni [em.] Gray, 1825 <
{Austrocypraca CossMANN, 1903
> OD Cypraca (Luponia) tcontusa McCoy, 1877 <

+Proadusta Sacco, 1894
> OD Cypraca tsplendens /Sacco, 1894[nec GRATELOUP,
1827] = tdenticulina Sacco, 1894 <
[=] tConocypraca OprpENHEIM, 1901
> SD Scuirper, 1924: Cypraca tpcrsona OpPpENHEIM,
1901 <
[=] tCypracacites (ScHLOTHEIM,
1820: not available according to
IRZN art. 56b) Scuiper, 1931

> SD Scuwpver, 1924: ¢Cypracacites tinflatus /Scuiot-
HEIM, 1820 = Cypraca tmcycri BortcER, 1883 <

Page 270

(err.) +Cyprecites Orzicny, 1850
(err.) tCypraeites BOTTGER, 1883

Propustularia ScHILDER, 1927 [}{Pro-
adusta subgenus]
> OD Cypraea surinamensis Perry, 1811 pro Cypraea
bicallosa Gray, 1831 <

72 (Erosariini) ScHILDER, 1924
[=] (Nariini) ScuiLpEr, 1932
[==] (Staphylaeini) IREDALE, 1935
Monetaria TRoScHEL, 1863
> SD Roserts, 1885: Cypraea moneta LINNAEUS, 1758 <
=f Thoracium (Rumputus, 1705)
Linnaeus, 1758 syn. [comprises
Monetaria and Ornamentaria|
= ||Aricia Broperip, 1837
> SD Fiscuer, 1884: Cypraea moneta LINNAEUS, 1758 <
(err.) Ariza DEsHayYEsS, 1865
(err.) Mercatoria MELvILL, 1888

Ornamentaria SCHILDER & SCHILDER in
ScHILDER, 1936 [Monetaria
subgenus]
> OD Cypraea annulus LinnaEus, 1758 <
==|| Aricia/MEtvitx, 1888
>SD MEtvitt, 1888: Cypraea annulus LinnaEus, 1758 <

Naria BroveEriP, 1837
> M Cypraea irrorata Gray, 1828 <

Paulonaria IrEDALE, 1930 [Erosaria

subgenus]
> OD Cypraea beckii Gasxoin, 1836 <

Erosaria TROSCHEL, 1863
> SD JoussEauME, 1884b: Cypraea erosa LINNAEUS, 1758 <
[=]]| Ocellaria We1nKaAurF, 1881
> SD Scuitper, 1927: Cypraea spurca Linnagus, 1758 <
[=] Ravitrona IREDALE, 1930
> OD Cypraca caputserpentis LINNAEUS, 1758 <
[==] Pertsserosa IREDALE, 1930
> OD Perisserosa brocktoni IREDALE, 1930 = Cypraea
guttata GMELIN, 1791 <
[=] Albacypraea STEADMAN & CotT-
TON, 1946
> OD Cypraea eburnea Barnes, 1824 <

Staphylaea JOUSSEAUME, 1884a
> TT, SD Jousseaume, 1884b: Cypraea staphylaea
LinnagEus, 1758 <
[==] Purperosa IrREDALE, 1935
> OD Purperosa facifer IREDALE, 1935 [= Cypraea
limacina LAMaRcK, 1810} <
[=] Eustaphylaea STEADMAN & CorT-
TON, 1946
> OD Cypraea semiplota MicuHEts, 1845 <

THE VELIGER

Vol. 10; No. 3

Nucleolaria Oyama, 1959
> pro Nuclearia JoussEAUME, 1884b, OD Cypraea nucleus
Linnaeus, 1758 <

==||Nuclearia JoussEAUME, 1884a
> SD JousszaumE, 1884b: Cypraca nucleus LinnaEus, 1758 <

OvuLipak FLEMING, 1828
= [em.]AMPHIPERATIDAE ADAMS & ApAMs, 1854

8 Eocypraeinae ScHiLper, 1924

81 (Eocypraeini) ScHiLpeEr, 1924
[==] (Sulcocypraeini) ScuiipeEr, 1932
[=] (Pseudocypraeini) STEADMAN & Cor-
TON, 1943

{Eocypraea CossMANnN, 1903
> OD Cypraea tinflata Lamarck, 1802 <

+Oxycypraea SCHILDER, 1927 [}Eocyp-
raea subgenus]
> OD Ovula tdelphinoides Cossmann, 1886 <

tApiocypraea ScHILDER, 1927
> OD Cypraca tmichaudiana GRraTELoup, 1847 <

+Eschatocypraea ScHILDER, 1966 [7A-
piocypraea subgenus]
> OD }Eschatocypraea tbalcombica ScuitpEr, 1966 <

Pseudocypraea ScHILDER, 1927
> OD Cypraea adamsonii SowErsy, 1832 <

{Sulcocypraea Conran, 1865
>M Cypraea tlintea Conran, 1847 <

+S phaerocypraea ScuHipErR, 1927
> OD Cypraea tbowerbankii SowerBy, 1850 <
[=] +Marginocypraea INcRAM, 1947
> OD }Marginocypraea +paraguana INcram, 1947
[= tSphaerocypraca twegeneri SCHILDER, 1939] <

82 (Cyproglobinini) ScHiLpER, 1932
[=] (Jenneriini) TureLe, 1929

+Cyproglobina Grecorio, 1880
> M, SD Cossmann, 1903: Cypraca (tCyproglobina)
tparvulorbis Grecorio, 1880 <

(em.) +Cypraeoglobina CosSMANN,
1903

+Luponovula Sacco, 1894 [;+Cyproglo-
bina subgenus]
> OD Cypraca t proserpinae Bayan, 1870 [= Cypraca
trugosa Broperip, 1827] <
+Cypracopsis SCHILDER, 1936 [+Cypro-
globina subgenus]
> OD fCypracopsis tvandervlerki ScuttpER, 1936 <

{Eotrivia ScHILDER, 1924 [{Cypropte-
rina subgenus]
> OD Cypraca (Trivia) tbouryi CossMaNN, 1889 <

Vol. 10; No. 3

TRE VELIGER

Page 271

+Cypropterina Grecorio, 1880
> M Cypraea (tCypropterina) }ceciliae Grecorio, 1880 <

{Cypracotrivia VREDENBURG, 1920
[tC ypropterina subgenus]
> OD Cypraea tduclosiana BAsTEROT, 1825 <
Jenneria JoUSSEAUME, 1884a [}Cyp-
ropterina subgenus |
> SD JousszaumeE, 1884b: Cypraea pustulata LAMARCK,
1810 [= pustulata SoLANDER, 1786] <
(= Pustularia/ MELVILL, 1888)
> SD ME vit, 1888: Cypraea pustulata Lamarck, 1810 <

+ Transovula Grecorio, 1880
>M, SD Cossmann, 1903: }Transovula tscheffert
Grecorio, 1880 <

B35, (+Cypraediini) ScuLpEr, 1927

{Cypraedia Swainson, 1840
>M {Cypraedia tcancellata Swainson, 1840 [= Cypraea
elegans Sowrrsy, 1823] <

(err.) +Cypraeadia SowErBy, 1842
(err.) ¢Cypraeidia TaTE, 1892
—=tCypraeoides Acassiz, 1846 (pro
+Cypraedia Swainson, 1840)
[=] tProtocypraedia ScHILDER, 1927
> OD Cypraea (+Cypraedia) tconigera Martin, 1914 <
;Eucypraedia Scuitper, 1939 [+Cyp-
raedia subgenus]
> OD Cypraea tsulcosa Lamarck, 1802 <

}Eovolva ScHILpER, 1932
> OD Amphiperas tnigeriensis NEwTon, 1922 <

9 Ovulinae FLemine, 1828

91 (Ovulini) FLemine, 1828
= Amphiperasini ADAMS & ApAMs, 1854
= (em.) Amphiperatini WinckworTH,
1929

Prionovolva IREDALE, 1930
> OD Ovulum breve SowrErsy, 1828 <

Diminovula IREDALE, 1930 [Primovula
subgenus]
> OD Diminovula vere punctata IREDALE, 1930 <
[==] Margovula IrEpALE, 1935
> OD Ovulum pyriforme SowERBy, 1828 <

Pseudosimnia Scuitper, 1927 [Prim-

ovula subgenus]
> OD Bulla carnea Poet, 1789 <

(err.) Pseudosinarica ARRECGROS,
1958

Primovula TuieEe, 1925
> OD Amphiperas beckeri Sowersy, 1900 <

[=] Dentivolva Haze, 1961
> OD Ovulum dorsuosum Hinps, 1844 <

Prosimnia ScuitvErR, 1927 [Primovula
subgenus |
> OD Ovula semperi We1nKAurF, 1881 [= Ovulum
coarctatum ApaMs & REEVE, 1848] <

Procalpurnus Tutere, 1929 [Calpur-

nus subgenus]
> OD Ovula lactea Lamarck, 1810 <

Calpurnus Montrort, 1810
> OD Bulla verrucosa LinnaEus, 1758 <

= Cypraella Swanson, 1840
> M Bulla verrucosa LinnaEus, 1758 <

(err.) Cypraela Sowerby, 1842
Ovula BrucutzreE, 1789

> SD Lamarck, 1801: Ovula oviformis Lamarck, 1801
[= Bulla ovum Linnaeus, 1758] <

(em.) Ovulus Montrort, 1810

(em.) Ovulum Sowersy, 1828

(em.) Ovularia Linx, 1830

(err.) Anula GrirritH, 1834
—=/Semiporcellana Costa, 1776 (obl.,

ApAMS & ADAMS, 1854, syn.)
> SD Scuitper, this paper: Bulla ovum Linnaeus, 1758 <

=/Liciuum (Humpurey, 1797)
ApAMs & Apams, 1854, syn.
> SD Scuixper, this paper: Bulla ovum Linnaeus, 1758 <
= Amphiperas (Gronow, 1781, not
valid according to ICZN opinion
261) HERRMANNSEN, 1846; A-
DAMS & ADAMS, 1854
> SD Herrmannsen, 1846: Bulla ovum Linnagus, 1758 <
(err.) Amphiceras Gray, 1847
[=] Parlictum IrEDALE, 1935
' > OD Ovula costellata Lamarck, 1810 <

92 (Simniini) ScHiLpeEr, 1927
[=] (Volvini) Scxuitper, 1932
Simnia Risso, 1826
> SD Gray, 1847: Simnia nicaccnsis Risso, 1826 <
(err.) Scymnia Risso, 1826 (partim)
(err.) || Stmia Leacu, 1847
[=] 7Calpurna FLeminc, 1828
> M Orcula fleathesi J. SowErsy, 1825 <
[=] Neosimnia FiscueEr, 1884
> OD Bulla spelta Linnaeus, 1758 <
Cyphoma Ropine, 1798
> M Bulla gibbosa Linnaeus, 1758 <
(err.)||Cyphonia Gray, 1847
= Ultimus Montrort, 1810
> OD Bulla gibbosa LinnaEus, 1758 <

Page 272

= Binvoluta SCHLUTER, 1838

> M Ovula gibbosa Lamarck, 1810 [= Bulla gibbosa
LINNAEUS, 1758] <

= Carinea Swainson, 1840

>M Bulla gibbosa Linnaeus, 1758 <

Pellasimnia IREDALE, 1931

> OD Ovulum angasi REEvE, 1865 <

Phenacovolva IREDALE, 1930 [Volva
subgenus]
> OD Phenacovolva nectarea IREDALE, 1930 [= Radius
brevirostris SCHUMACHER, 1817] <
—|| Radius ScHuMacuHER, 1817
>M Radius brevirostris ScoHUMACHER, 1817 <
(err._||Voluula Prrspry, 1895

> TT Bulla volva Linnaeus, 1758 <

(err.) Volvula Pitspry, 1895
— Radius Mon Trort, 1810

> OD Bulla volva Linnaeus, 1758 <

==/ Birostris Fasricius, 1823 nud.,
ScHILpER, 1932 syn.
= Birostra Swainson, 1840

> M Bulla volva Linnaeus, 1758 <

The bold face figures refer to the corresponding bold face numbers

of the tribus in the Systematic List

Adusta 63
Aerythraea 51
Afrocypraea 42
Akleistostoma 42
Albacypraea 72
Amphiceras 91
Amphiperas 91
Annepona 71
Anula 91

A piocy praca 81
Arabia 51
Arabica 51
Archicypraea 41
Archierato 13
Arestorides 51
Aristorides 51
Aria 72

|| Aricia 72, 72
Austrocypraca 71
Barycy praca 42
Basilitrona 52
||Basterotia 51
Bastorotia 51
Bernaya 42
Bernayia 42
Binvoluta 92
Birostra 92
Birostris 92
Bistolida 63
Blasicrura 63
Callistocypraea 51

Calpurna 92
Calpurnus 91
Carinea 92
Cavicypraca 51
Chelycypraea 52
Cipraca 51
Cleotrivia 23
||Coccinella 22
Conocypraea 71
||Cribraria 63
Cribrarula 63
Criraria 63
Cupinota 63
Cyphoma 92
||Cyphonia 92
Cypraea 51 (7 times)
Cypraeacites 71
Cypraeactaeon 42
Cypraeadia 83
Cypraearius 51
Cy pracdia 83
Cypracerato 13
Cy pracidia 83
Cypraeites 71
Cypraela 91
Cypraella 91
Cypracogemmula 30
Cypraeoglobina 82
Cypracoides 83
Cy pracopsis 82
Cypracorbis 42

THE VELIGER

Cypracotrivia 82
Cypraeova 62
Cypraeovula 62
Cypracovulum 62
Cypracovum 62
Cyprea 51
Cyprecites 71
||Cypria 51
Cypreovula 62
Cypriarius 51
Cyproca 51
Cyproglobina 82
Cypropterina 82
Cyprovula 62
Dentiora 30
Dentivolva 91
Derstolida 63
Diminovula 91
Dolichupis 23
Eclogavena 63
Ellatrivia 23
Eocypraca 81
Eotrivia 82
Eovolva 83
Epona 71, 71
Eponia 71

Erato 13
Evatodium 11
Eratoena 13
Erato psis 13
Eratotyivia 12
Erosatia 72
Erronca 63
Erythraca 51, 51
Eschatocypraca 81
Etrona 51
Etronsa 51
Eucypracdia 83
Eustaphylaea 72
Evanaria 63
Evenaria 63
Ficadusta 63
Fossacypraca 52
Fossatrivia 21
Gaskoinia 62
Gigantocypraca 42
Gisortia 43
||Globularia 71
||Globulina 61
Gratiadusta 63
Guttacypraea 62
Hespercrato 13
Ipsa 71

Jenneria 82
Ipserronea 63
Johnstrupia 11
Jousscaumca 52
Jousseaumia 52
Lachryma 13, 13
Leporicypraea 51

Vol. 10; No. 3

Licium 91
Luperia 62
Lupina 62
Luponaria 62
Luponia 62
Luponica 62
Luponovula 82
Luria 52
Lyncina 51, 51
Macrocypraea 51
Mandolina 41
Marginocypraca 81
Margovula 91
Maurina 51
Mauritia 51
Mauxiena 51
Megalocypraea 43
Melicerona 63
Mercatoria 72
Miolyncina 51
Monetaria 72
Muracypraea 42
Mystaponda 51
Naria 72
Neobernaya 61
Neosimnia 92
Niveria 23
Nivigena 63
Notadusta 63
Notocypraea 62
Notoluponia 62
Nototrivia 21
||Nuclearia 72
Nucleolaria 72
||Ocellaria 72

O pponaria 63
Ornamentaria 72
Ovatipsa 63
Ovula 91
Ovularia 91
Ovulum 91
Ovulus 91
Oxycypraca 81
Palacocypraca 41
||Palacocypraca 41
Palangerosa 63
Palliocypraca 62
Palmadusta 63
Pantherinaria 51
Parlicium 91
Paulonaria 72
Pedicularia 30
Pediculariella 30
Pediculariona 30
Pellasimnia 92
Peribolus 51
Perisserosa 72
Phenacovolva 92
Ponda 51
Porcellana 51

Vol. 10; No. 3

Porcellanites 41
Postularia 71
Primovula 91
Prionovolva 91
Proadusta 71
Procalpurnus 91
Prolyncina 51
Propustularia 71
Prosimnia 91
Proterato 13
Protocypraea 42
Protocypraedia 83
Prototrivia 21
Prozonarina 61
Pseudocypraea 81
Pseudogisortia 43
Pseudosimnia 91
Pseudosinarica 91
Pseudotrivia 23
Pseudozonaria 61
Purperosa 72
Purpuradusta 63
Pustula 23
Pustularia 71, 82

Pustulariopsis 41
Pusula 23
Radius 92
||Radius 92
Ravitrona 72

Rhynchocypraea 62, 62

Schilderia 61
Scymnia 92
Semicypraea 30
Semiporcellana 91
Semitrivia 21
||Simia 92
Simnia 92
Siphocypraea 42
Solvadusta 63
Sphaerocypraea 81
Staphylaea 72
||Stolida 63
Sulcerato 13
Sulcocypraea 81
Sulcotrivia 22
Talostolida 63
Talparia 51

|| Tessellata 52

THE VELIGER

Thelxinovum 62
Thoracium 72
|| Thyreus 30
Tigris 51
Transovula 82
Trivea 22
Trivellona 23
Trivia 22
Triviella 21
Trivirostra 23
Trona 51
Ultimus 92

Page 273

Umbilia 62
Vicetia 43
Volva 92

|| Voluula 92
Vulgusella 51
Vulpicella 51
Vusula 23
Willungia 21
Zoila 42
Zonaria 61
Zonarina 61
Zonatia 61

LITERATURE CITED

ScuHILper, Franz ALFRED

1939. Die Genera der Cypraeacea. Arch. Molluskenk. 71

(5/6): 165 - 201; 2 plts.

(1 November 1939)

1941. Verwandtschaft und Verbreitung der Cypraeacea.
Arch. Molluskenk. 73 (2/3) : 57 - 120; 2 plts. (15 May 1941)

1966. The higher taxa of cowries and their allies. The

Veliger 9 (1): 31-35

(1 July 1966)

Page 274

THE VELIGER

Vol. 10; No. 3

Semele martinit (REEVE, 1853) of Southern Brazil and Uruguay

MIGUEL A. KLAPPENBACH

Museo Nacional de Historia Natural

Montevideo — Uruguay

Semele martini WAS PRESENTED and read by ARTHUR
Apams to the Zoological Society of London on July 12,
1853, but it was not published in the Proceedings until
a year later, in July, 1854 (for 1853: p. 97). Meanwhile,
it was cited, described and figured by REEVE as Amphi-
desma martinii in November, 1853 (Conch. Icon., spec.
43, pl. 6, fig. 43). It may be noted that REEVE used two
“?’s, whereas ADAMS used one in the trivial name. In spite
of the fact that Reeve credited the species to ADAMS and
made reference to the presentation by ApAmMs in the Society
in July, 1853, this species must take authorship of REEVE,
according to the Rules of Nomenclature. Since then, this
species has not been often noted in the malacological
literature. We find only a listing of it by PAETEL (1890:
p. 63) and a brief footnote by Lamy (1913: p. 354).
LANGE DE Morretes cited it later in his Addenda (1954:
p. 43). He had previously not mentioned this species in his
Catalog (1949). It is evident that LancE DE MorretTes
was not acquainted with it. However, the brief original de-
scription and the figure are quite adequate, and allow us
to identify the species.

Recently, this species was rediscovered and redescribed
under the name Semele aurora TURSCH & PiERRET (1964:
p. 35). The description by these authors coincides with the
one by Reever, and the type locality is the same, Rio de
Janeiro, Brazil. This is a rare offshore species that probably
does not reach the beach, which explains the scant know-
ledge we have about it. Semele martinii, restricted ap-
parently to the Rio de Janeiro area, ranges south to off
the Rio de la Plata. The Museo Nacional de Historia Na-
tural of Montevideo, Uruguay, has specimens (single
valves) from Maldonado Bay and Lobos Island, Uruguay.
They are rather eroded, but identifiable. They reach a
larger size than the specimens from Rio de Janeiro.

In the collection of the Academy of Natural Sciences of
Philadelphia (no. 313078) there is a specimen, left valve,
from La Paloma, Depto. Rocha, Uruguay, in the same
condition. Length, 56.0mm; height, 48.2 mm. Also in the
collection of the Academy there is a live-taken specimen
from Raza Island, off Rio de Janeiro, Brazil, in 30
fathoms, which measures: length, 40.0mm; height, 33.6
mm; width, 14.2mm; the light-brown periostracum is
more visible on the anterior and posterior ends. In this
specimen it is possible to see the irregular and strong con-

centric folds, often bifurcated, mentioned by REEVE in his
original description. These bifurcated folds, among other
characters, allow us to separate easily this species from the
others recorded from the South American Atlantic coast,
namely: Semele proficua (PULTENEY, 1799), S. purpur-
ascens (GMELIN, 1791), S. cancellata (SowErRBy, 1833),
S. (Semelina) nuculoides (Conran, 1841) and S. casali
DoELLo-Jurabo, 1949.

Semele uruguayensis Piissry, cited by some authors, is
the result of the confusion between Abra uruguayensis
Pitspry, 1897 (the types of which are in the ANSP, lot
no. 70493), and the species that DorLto-Jurapo recently
described. Semele casali has, according to its author
(1949: p. 1), a range that extends along the Uruguayan
coast, south to San Matias Gulf in Argentina. However,
in the collection of the ANSP (no. 182030) we found a
left valve of this species, in good condition, from Cara-
pegus, State of Espirito Santo, Brazil, which extends its
distribution to the north approximately 2500 kilometers.

LITERATURE CITED

ADAMS, ARTHUR
1854. Descriptions of new species of Semele, Rhizochilus,
Plotia and Tiara in the Cumingian collection. Proc. Zool.
Soc. London (1853) 21: 94 - 99 (25 July 1854)
DoELLo-Jurabo, M.

1949. Dos nuevas especies de Bivalvos marinos. Com.
Zool. Mus. Hist. Nat. Montevideo 3 (57): 1-8; plt. 1
Lamy, E.
1913. Revision des Scrobiculariidae vivants du Muséum d’His-
toire Naturelle de Paris. Journ. de Conchyl. 61: 243 - 368;
plt. 8
LANGE DE MorretEs, F.
1949. Ensaio de catalogo dos moluscos do Brazil. Arq.
Mus. Paranaense 7: 5-216 Curitiba, Parana, Brasil
1954. | Adenda e corrigenda ao ensaio de catalogo dos moluscos
do Brasil. Arq. Mus. Paranaense 10: 37-76, Curitiba,

Parana, Brasil
PaETEL, FRIEDRICH
1890. | Catalog der Conchylien-Sammlung. Die Acephalen und
die Brachiopoden. 3: 1-256
REEvE, Lovett Aucustus
1853. | Conchologia Iconica 8: Monograph of the genus Am-
phidesma; pits. 1-8; 53 sps.
Turscu, BERNARD & JEAN PIERRET
1964. | New species of mollusks from the coast of Brazil.
The Veliger 7 (1): 35-37; 5 text figs. (1 July 1964)

VS
JONG Lot

Vol. 10; No. 3

THE VELIGER

Page 275

Observations on Hipponix conicus (SCHUMACHER, 1817)

WALTER OLIVER CERNOHORSKY

Vatukoula, Fiji Islands

(Plate 41; 3 Text figures)

SPECIMENS OF THE PARASITIC protandrous hermaphrodite
Hipponix conicus (SCHUMACHER, 1817) utilized in this
study were collected in Fiji, Tonga and the Niue Islands.
The specimens occurred as an associated male-female pair
in all three localities, parasitic on Conus mustelinus Hwass
in Brucuikre, 1792, Mitra coffea SCHUBERT & WAGNER,
1829, Strigatella decurtata (RerveE, 1844) and Bursa gra-
nularis (ROpiNG, 1798). The four hermaphroditic speci-
mens were attached directly to Mitra coffea, M. stictica
(Linx, 1807), Conus striatus Linnarus, 1758 and C.
omaria Hwass in BruGulkrE, 1792.

All specimens were collected in shallow water, from
0 to 5 feet in depth. The specimens examined were as
shown in Table 1.

Fijian specimens of Hipponix conicus were examined in
the living state, while the Tongan and Niue Island speci-
mens were received preserved in spirits. A post mortem
change in preserved specimens was a uniform brown dis-
coloration of all soft parts. The six female specimens, all
twice as large as the small males, and the four hermaphro-
ditic specimens were attached to the shell-surface of their
host ; the seven small males were sedentary on the females.

TAXONOMY

Hipponix DEFRANCE. 1819

1817. Amalthea ScuuMacueER, Essai nouv. syst., pp. 56, 181.—Type
species by SD (Gray, 1847) A. conica ScHuMACHER, 1817
(non Amalthea RaFINESQUE, 1815)

1819. Hipponix DEFRANCE, Journ. Physique et Chim. 8: 217—Type
species by OD “H. mitrata GMELIN” (error for mitrula GMELIN,
1791) = Hipponix antiquatus (LinNAEus, 1767)

1819. Hipponyx [sic] DEFRANGE, BLAINVILLE, Bull. Sci. Soc. Philom.,

p.

1841. Sabia Gray, Syn. Cont. Brit. Mus. ed. 43, p. 126 [nom. nud.]

1842. Sabia Gray, Syn. Cont. Brit. Mus. ed. 44, p. 90 [nom. nud.]

1847. Sabia Gray, Proc. Zool. Soc. London, p. 157 — Type species
by OD Amalthea conica ScHuMACHER, 1817

1906. Malluvium Mevviti, Proc. Malac. Soc. London 7: 81 - 84
Type species by OD Capulus lissus E. A. SMiru, 1894

Table 1
2
a a
Se : 5
Oy Host g
iat BA ok eae species Position 3
1 20.5 ¢ + C. mustelinus apex of spire F
2 Wi). Oi eis i granularis body whorl F
3 11.3 ¢ + S.decurtata body whorl N
fe OV/ae etm Viencoviea labial lip T
59 101 2 + M.coffea penult whorl T
OS one ee Mitcofiea antepen. whorl T
7 94 $¢ — C. striatus spire F
8 80 ¢ — C.omaria spire F
9 703 So = OWS SGhIEG penult whorl N
10 76 ¢ — M.coffea labial lip AD
11 49 $ — H.conicus 2 dextral side N
12 49 $¢ — H.conicus ¢ dextral side FE
13 44 $¢ — H.conicus 2? dextral side a
14 42 ¢ — H.conicus 2? dextral side F
15 3.7 oS: —= Ai. conicus ¢ dextrallside aw
16 3.33 8 — H.conicus ? dextral side T
17 22 $3 — H.conicus 2 dextral side F

' F — Fiji; N - Niue Island; T — Tonga Island

Mipponix conicus (ScHUMACHER, 1817)

1817. Amalthea conica ScHuMACHER, Essai nouv. syst. p. 181, plt.
21, figs 4a, 4b, 4c

1819, Patella australis Lamarck, Anim. sans vert. 6: 335

1830. ? Pileopsis paleacea MENKE, Syn. meth. Moll., p. 147

SCHUMACHER, 1817, established Amalthea for the spe-
cies A. conica SCHUMACHER, 1817 and A. maxima Scuu-
MACHER, 1817 [==Capulus ungaricus (Linnaeus, 1758) ].
Capulus ungaricus (LinnaEuS) has already been appro-
priated as the type-species of Capulus Montrort, 1810,

Page 276

and Amalthea conica was subsequently designated by
Gray (1847, p. 157) as the type species of Amalthea.
ScHUMACHER’s genus name, however, is a homonym of
Amalthea RAFINESQUE, 1815, and therefore not available.

The Drerrance reference appears to be a scarce journal
which is not available to me and is lacking in the Austral-
ian and Auckland Museums; entries have been compiled
from SHERBORN (1902 - 1933) and E. A. Smiru (1906).
SmirH, who obviously consulted the original reference,
states that “Hipponix mitrata GMELIN,” which presum-
ably is a lapsus for H. mitrula (GMELIN), was designated
as the type-species of Hipponix by DrEFRANCE in the
original publication. Hipponix mitrula (GMELIN) is syn-
onymous with H. antiquatus (LINNAEUS). “Hipponyx”
BLAINVILLE is only a misspelling of Hipponix DEFRANCE
in a later publication (vide synonymy). Since the type-
species of Hipponix has been originally designated, Gray’s
subsequent designation (op. cit., p. 157) of Patella cornu-
copiae Lamarck, 1803 (Eocene fossil from the Paris
Basin) as the type species of Hipponix is invalid.

Gray (op. cit., p. 157) established the genus Sabia
through publication in synonymy of Amalthea ScHUMA-
CHER; the type-species therefore is also A. conica ScHu-
MACHER by original designation. Gray himself quoted
Sabia as of 1833, but this reference appears untraceable;
on two occasions Sabia appeared as a nomen nudum.

ME tv, 1906, established Malluvium as a subgenus of
Amalthea SCHUMACHER for the deep-water species Capu-
lus lissus E. A. Smiru, from the Bay of Bengal. The author
laid stress on the absence of radiating sculpture of C.
lissus, which he stated is present in all specimens of
either Amalthea ScHUMACHER or Capulus MontTFortT.
SMITH (op. cit.) regarded MELviLi’s Malluvium as “pre-
mature if not unnecessary,’ and correctly pointed out
that the presence or absence of a concentric or radial
sculpture in this group of species is hardly sufficient for
the erection of a new subgenus. SmMiTH’s contention must
be supported in view of the presence of smooth and
heavily ribbed or concentrically lamellated specimens of
Hipponix conicus in even the short series of specimens
used in this study. Tureve, 1903, figured the radula of
“Hipponix lissus,” which is basically similar to the radula
of H. conicus, and included it in the family Capulidae.
E. A. SmitH (op. cit.) remarked that Professor Gwatkin
studied the radulae of Capulus lissus and pronounced
them to be nearest Amalthea SCHUMACHER.

Taxonomy on the specific level appears to be equally
confused in view of the great variability of conchological
characters, i.e. colour, form and sculpture of Hipponix
conicus; several currently acceptable specific names may

THE VELIGER

Vol. 10; No. 3

prove to be synonyms in a detailed comparison survey.
E. A. SmirH (op. cit.) synonymized Patella australis La-
MARCK with Hipponix conicus, and regarded LAMARCK’S
name to be of prior date without mentioning the year of
authorship. I was unable to trace Patella australis La-
MARCK prior to 1819. THrELEe, 1925, equates H. acutus
Quoy & Gamarp, 1835 with H. conicus (SCHUMACHER).

There is no unanimity of opinion among malacologists
as to a supragencric classification of the group of related
hermaphroditic taenioglossate gastropods. THIELE, 1929,
placed the cap-shaped, non-operculate parasitic and herm-
aphroditic Amaltheidae (= Hipponicidae) together with
operculate, spirally coiled and sometimes viviparous Fossa-
ridae, and operculate and dioecious Vanikoroidae in the
superfamily Amaltheacea (== Hipponicacea). Families
with similar shells, anatomy, oviposition and sexes to the
Hipponicidae, e.g. Capulidae and Calyptraeidae, were
placed by TureELe with operculate and dioecious Tricho-
tropidac and operculate Amaplocamidae in the super-
family Calyptraeacea.

Hipponicidae show a considerable similarity to Capu-
lidae: they are similar in shell-form, having a high-peaked,
cap-shaped shell which lacks an internal apical plate.
Both are sedentary parasitic or semi-parasitic hermaphro-
dites, with the females retaining their brood within the
shell and veligers spawning as echinospiral veligers. ‘Their
radulae arc so similar that in other groups of gastropods
they would not be considered to exceed generic level.
Specics of the hipponicid genus Cheilea Mopegr, 1793,
are more dissimilar to Hipponix than the latter is to
Capulus. Cheilea species are parasitic on rocks, the ani-
mal’s proboscis is slender and longer and lacks the side-flaps
near the mouth; the rhachidians of the radula are wing-
shaped, and the interior of the shell has an horse-shoe
shaped, moderately long and shelly plate extension; fe-
males retain their spawn within the shell, and males are
often found attached to the females.

The difference between Capulidae and Hipponicidae
appears to be mainly in the mode of feeding and host
selection. Capulus is reported to be a ciliary feeder and
gencrally prefers bivalves for hosts. SCHEPMAN, 1909, how-
ever, reports C’. danicli Crosse, 1858, attached to “Gyri-
neum cuspidatum Reeve.” These differences are few and
doubtfully exceed family or superfamily level of classifi-
cation. The present classification appears to be a reten-
tion of an orthodox taxonomic arrangement and not a
superfamilial separation based on tangible evidence. Hip-
ponix conicus would be more appropriately placed some-
where close to Capulidae, within the superfamily Calyp-
tracacea, and not in the Hipponicacea.

Vol. 10; No. 3

THE SHELL AND POSITION OF ATTACHMENT

The species Hipponix conicus is rather variable in form,
colour and sculpture. The outline of the shell margin
generally corresponds with the shell-surface of its host.
Some specimens are high and cap-shaped, with a curved
and obsoletely coiled apex which protrudes slightly over
the posterior shell margin; other specimens are oval,
elliptical, round or irregularly pentagonal in outline from
a ventral view. The apex can be displaced as much as 4
of the shell-length towards the centre, and viewed in
profile some specimens appear rather depressed. Shells
are sculptured with 15 - 34 flattened radial ribs, while in
some small males ribs are completely absent. The con-
centric lamellations and coiled apex are generally obsolete
in female shells, but quite distinct in some males; the
radial ribs form a scalloped shell-margin. The exterior of
the shell is either white, dirty grey, partly white and
reddish-brown or white with reddish-brown rays. The
interior of the shell is either pure white, white with orange
muscle-scars, white with brown stains towards the apex
or bluish-white with traces of brown muscle scars. Females
are twice as large as the males, and hermaphroditic speci-
mens are only slightly smaller than the females.

In all specimens examined, the females and hermaphro-
dites were attached directly to their hosts while the small
males were sedentary on the females. The female will oc-
cupy almost any position on the host’s shell-surface, with
the exception of the area on the dorsal centre axis. The
small males were fastened to the dextral side of the female

100 p

THE VELIGER

Page 277

shell, and close to the shell margin, with the apex facing in
the general direction of the female’s apical region. Promi-
nent scars are left on the host which correspond closely to
the internal anatomy of the parasite. The scalloped shell
margin is clearly indented in the host’s shell-surface, the
surrounding area is shallow and oval and rises to a small
elevated plateau which has a central depression for the
reception of the foot. The same type of scars are visible
on the female Hipponix where the males were attached.

Restricted locomotion is evident in the species prior to
reaching the required position of final immobility; one
small male was somewhat too far away from the female’s
shell margin and subsequently moved into position, leaving
a well-defined track in the female’s shell-surface. HARTLEY
(1958) discussed locomotion of Hipponix conicus and
illustrated tracks of the species on the dorsal surface of
Haliotis ruber Leacu, 1815. Once in position, it appears
that the animal becomes immobile and temporarily sed-
entary. There is no evidence of hole-boring in Hipponix
conicus; the small hole visible near the anterior margin
of the labial lip on Mitra coffea is irregular, and has been
caused through the softening process of the animal’s body
during deposition on a thin part of the host’s shell.

THE ANIMAL

The animal’s foot is small, disc-like, and fawn in colour;
the mantle and horse-shoe shaped muscle are yellow in
colour. The head is long when extended, thick and brown-
ish-grey in colour; two short, curved and pointed flaps

Figure 1

Hipponix conicus (SCHUMACHER)
a. Half row of radular teeth of female specimen No. 1
b. Rhachidian of radula of male specimen No. 14
Fiji Islands

Page 278

are situated on each side of the animal’s head. The head
moves steadily from side to side and at times exposes the
proboscis, displaying the whole length of the radular
ribbon, vertically to the axis of the body, from the roof
of the mouth to the floor of the mouth. Tentacles are
long, moderately stout, yellow in colour, and either later-
ally streaked with purplish-brown or fully dark purple at
the distal end. Eyes are small and simple and placed
dorsally on the body near the junction of the tentacles
and close to the tentacle’s margin. The penis, where ob-
served, was either moderately short or rudimentary, and
cream in colour. An operculum is absent.

THE RADULA

The radula is taenioglossate, short and broad in the
female, translucent-white in colour, formula 2-1:1-1- 2.
Rhachidians are broader than they are long, and equipped
with a long central cusp which extends past the plate-
margin; four smaller accessory cusps are positioned on
either side of the main cusp. Lateral teeth are very broad,
with a large main cusp and eight accessory denticles on
the cutting edge of the main cusp. Inner marginals have
six denticles, outer marginals about 5 denticles on the
cutting edge of the main cusp. The female’s radular ribbon
measured 1.70mm in length, 0.74mm in width in an
animal with a shell 20.5mm in length; the ribbon con-
tained 32 fully formed rows of teeth and 6 nascentes.
Considerable wear on the teeth was evident in the front
rows and persisted to the 12" anterior row. The consid-
erable wear on the teeth and rather large proportion of
nascentes to fully formed teeth would indicate frequent
use of the radula on hard substratum.

The radulae of small males differ in minor but signifi-
cant points from those of the females. In one male the
radular ribbon measured 1.12 mm in length and 0.26mm
in width in an animal with a shell 4.2mm in length;
radulae of other males differ only slightly in proportion.
The rhachidians of the radula have a slightly shorter
central cusp and only three accessory denticles, while the
laterals have only seven denticles on the cutting edge. The

THE VELIGER

Vol. 10; No. 3

marginal pegs on rhachidians are longer and more promi-
nent, and the ribbon contains about 53 rows of fully
formed teeth and only 1 to 2 nascentes.

The radular ribbon of the males is longer in proportion
to shell-length than that of the female (male: 37.5% of
length; female 12% of length), and the ribbon is more

proboscis

tentacle

eye

Figure 2

Hipponix conicus (SCHUMACHER)

Dorsal view of anterior part of animal

slender in proportion to total ribbon-length (male: 23%;
female: 43%). No wear whatsoever was evident in the
male radulae and the cusps of the first anterior row were
as sharp as those in subsequent rows; the number of
nascentes is reduced accordingly.

EGG CAPSULES AND VELIGERS

Egg-capsules are retained within the parent’s body and
are attached to the underside of the animal. Six females
contained spawn, while only brood-sacs were present in
hermaphroditic specimens. The female specimen from Fiji
contained 36 clongated, vase-shaped capsules which are
translucent-cream in colour; each capsule contained on
the average 300 ova, which would be about 11000 ova in
the spawn. Capsules were attached by fine but strong
filaments to a central brood-sac, which contained round
ova of varying sizes, measuring from 15 uw - 30 u. Eggs in
the capsules were in different stages of development, some
having reached an early veliger stage, while others were
in the trochophore phase. Both trochophores and veligers
measured from 120 - 150 wu. Early veligers had a cepha-

Explanation of Plate 41

Figure 1: Ventral view of living animal of Hipponix conicus
(ScHUMACHER): view of female specimen No. 1, showing egg
capsules, tentacles and extended head. x 4.0

Figure 2: Female Hipponix conicus specimen No. 1 carrying male
specimen No. 12, in situ on Conus mustelinus Hwass in BRUGUIERE

X 1.3

Figure 3: Female Hipponix conicus specimen No. 4 carrying male
specimen No. 13, and hermaphroditic specimen No. 10 attached to
labial lip margin of Mitra coffea SCHUBERT & WAGNER. x 2.5

Figure 4: Etched out shell-scars on Mitra coffea after removal of
Hipponix conicus specimens. x 2.6

Figure 5: Dorsal and ventral view of female Hipponix conicus
specimen No. 5 carrying male specimen No. 15. x 2.5

Figure 6: Lateral view of specimens No. 5 and No. 15. x 2.5
Figure 7: Dorsal and ventral view of female Hipponix conicus
specimen No. 1 carrying male specimen No. 12. x 2.1

Figure 9: Lateral view of specimens No. 1 and No. 12. x2.1

THE VELIGER, Vol. 10, No. 3 [CeRNOHORSKY] Plate 41

Figure 2

Figure 3 Figure 4 Figure 5

Figure 6 Figure 7 Figure 8

photographs by W. O. CernoHorsky

Vol. 10; No. 3

THE VELIGER

Page 279

Figure 3

Cheilea equestris (LINNAEUS)

a. Half row of radular teeth of a female specimen
b. Rhachidian of radula of a male specimen parasitic on the female
Fiji Islands

lopedal mass of transparent cells and a transparent embry-
onic shell. Some of the liberated veligers were free-
swimming and vigorously active, while others were station-
ary but rotating freely within the fluid.

FEEDING HABITS

The actual feeding process was not observed and deduc-
tions can be made only from evidence on hand. The
unworn radula of the male shows that he is not a detritus
feeder and does not abrade on hard substrata. He does
not only position himself on the female’s dextral marginal
edge solely for the purpose of fertilization but possibly
also for feeding purposes. The male either subsists on
particles of food gathered by the female or even feeds on
her faecal pellets which are regularly discharged and man-
euvered to the same marginal edge where the male is
reposing.

FAECES

The gut contained loose faeces as well as formed faecal
pellets. During examination of the living animal, faecal
pellets in groups of 4-5 were discharged by the female
in about 20 minute intervals; pellets are “0O”-shaped,
greenish-brown and about 1.0mm long. They contain
mucus and a predominance of solids, e.g. coral, sand

grains, algae, some fragments of radiolarians and some
unidentified regular and slender spicules. In one specimen
a rather large and cuboidal piece of sand grain was
reposing in the gut.

DERMATOPTIC PERCEPTION

The living animal was turned over onto its shell, with the
internal organs facing the light source, but eyes well
shielded from the light source by the interposing body.
The animal’s head was moving freely, but it violently
retracted its head extension as soon as the impact of
the shadow fell on its body. It was thought at first that
the closeness of the hand may have been sensed by other
means than dermatoptical ones, and a sheet of clear
glass was placed between the light source and brought
close to the animal’s body; it failed to elicit a quick with-
drawal of the head. The variation of light intensity may
therefore be perceived by the animal through the skin
alone or may be sensed by minute eye-sensors (as in
Chitonidae) as yet not recognized.

SUMMARY

Hipponix conicus (SCHUMACHER) is a parasitic protand-
rous hermaphroditic mesogastropod which is not host-
specific. Shells are small, about 2mm to 21 mm in length,

Page 280

generally cup-shaped or conforming in shape to the shell
substratum of its host; shells are rather colorless, smooth
or sculptured with radial ribs or occasionally concentric
rings. The radula is taenioglossate. Males and females
occur in associated pairs with the females parasitic on a
gastropod and the males attached to the female Hipponix.
Evidence suggests that the species begins its first sexual
phase as a small male fastened to the female’s dextral shell
margin for fertilization and possibly faecal pellet feeding.
The species then gradually passes into the hermaphroditic
stage when it grows larger, intermediate in size between
males and females, and settles directly on the main host.
In the final female stage the species is completely seden-
tary and has developed a larger shell, capable of accommo-
dating the brood-sac and spawn which is retained within
its body. Veligers are free-swimming and echinospira.

Hipponix conicus has a wide distribution ranging from
East Africa through the Indo-Pacific to Japan, the Hawai-
ian Islands and the Tuamotu Archipelago.

ACKNOWLEDGMENTS

I would like to thank Dr. A. W.B. Powell, Auckland
Institute and Museum, and Dr. D. F McMichael, Aus-
tralian Museum, Sydney, for their assistance with literature
references. The preserved material received from Mr. H.
C. Gay, Nuku’alofa, Tonga and Mr. N. McDowall, Niue
Island, has been much appreciated.

LITERATURE CITED

Gray, Joun Epwarp
1847. _ A list of the genera of Recent Mollusca, their synonyma
and types. Proc. Zool. Soc. London 17 (178): 129 - 219
(30 November 1847)

THE VELIGER

Vol. 10; No. 3

Hartley, THELMA
1958. | Movement by commensal Hipponix conicus (ScHUMA-
CHER). Journ. Malac. Soc. Austral. 2: 37; plt. 8
(November 1958)
MELVILL, JaMEs Cosmo
1906. Capulus lissus SM1TH, as type of a proposed new subgenus
(Malluvium) of Amalthea ScHuMAcHER. Proc. Malac. Soc.
London 7 (2): 81 - 84; 4 text figs. (29 June 1906)
ScHEPMAN, MaTTHEUS MaRINUS
1909. The Prosobranchia of the Siboga Expedition. Part II.
Taenioglossa and Ptenoglossa. Siboga Exped. 49b: 110 - 231;
plts. 10 - 16
SCHUMACHER, CHRETIEN FREDERIC
1817. Essai d’un nouveau systéme des habitations des vers
testacés. Copenhague, pp. 1 - 287; plts. 1 - 22
SHERBORN, CHARLES Davies
1902 - 1933. Index animalium.
to 7056; i- cxlvii; 1 - 1098

SmitH, Epcar ALBERT

London, Sect. 2: i - cxxxvi; 1

1906. Note on the subgenus Malluvium MELVvILL. Proc.
Malac. Soc. London 7 (3) : 122 - 123 (5 October 1906)

THIELE, JOHANNES
1903. Die beschalten Gastropoden der deutschen Tiefsee-

Expedition 1898-1899. B: Anatomisch-systematische Unter-
suchungen einiger Gastropoden: 7: 149 - 179; plts. 6-9

1925. Gastropoden der deutschen Tiefsee-Expedition. II. Teil.
Wiss. Erg. d. deutsch. Tiefsee Exp. “Valdivia” 1898 - 1899.
17 (2): 1-382; plts. 13-46; 31 text figs.

1929. Handbuch der systematischen Weichtierkunde.

Jena, Gustav Fischer, 1929-1935; 1- 1154; 893 text figs.
London, Ser. B. 234: 29 - 76

YONGE, CHARLES MAURICE

1953. | Observations on Hipponix antiquatus (LINNAEUS).
Proc. Calif. Acad. Sci. ser. 4, 28 (1): 1-24; 9 figs.
(15 July 1953)

Vol. 10; No. 3

THE VELIGER

Page 281

Notes on the Range Extension

of the Boring Clam Penitella conrad: VALENCIENNES

and its Occurrence in the Shell of the California Mussel

BY

STEVEN E. MEREDITH

Department of Zoology, Oregon State University, Corvallis, Oregon 97331

and

Oregon State University Marine Science Center, Newport, Oregon 97047

(Plate 42)

THE STUDY REPORTED ON herein was conducted at the
Haystack Rock area, Cannon Beach, Clatsop County,
Oregon, Latitude 45° 53’04” N, Longitude 123° 58’ W.
Investigations were carried on seasonally throughout 1964
as a part of the studies for a master’s thesis at the Univer-
sity of Puget Sound, Tacoma, Washington, and again in
the summer of 1966, while the writer was in residence at
the Oregon State University Marine Science Center,
Newport, Oregon.

The range of Penitella conradi VALENCIENNES, 1846
was previously reported to be from Bahia San Bartolomé,
Baja California, Mexico, to Gualala, Mendocino County,
California (Turner, 1955). This species has not previ-
ously been reported in Oregon, and this report represents
a considerable extension of its northern range limit.
TurNER reported finding P. conradi in the shells of various
mollusks, including Mytilus californianus Conrap, 1837,
in California. This species is very commonly found infest-
ing species of Haliotis (abalones) and has been commonly
referred to as the ““Haliotis borer.”

The mussel beds at Cannon Beach attain a maximum
vertical width of approximately 7 feet on the rock faces,
and a check of several hundred Mytilus revealed that
Penitella is restricted to the lowest 2 feet of the beds. A
thorough examination of 113 mussels showed that the
percentage of infestation was approximately 20%. The
number of borers per host ranged from 1 to 11, with an
average of 3.8.

Penitella conradi invades the mussel when quite young
and grows considerably as it bores slowly into the valve,
thus creating a conically-shaped burrow. The entry holes
range in size from less than 1mm to 2mm in diameter
and are often difficult to detect, especially if the mussel
valves are heavily fouled with such organisms as barnacles,
tubiculous polychaetes, etc; therefore, a fast and reliable
method of determining whether or not the borer is present
is to open the host, remove the soft tissues, and examine
the inner surface of the valves. The presence of the in-
vader is evidenced by patches of black, leathery conchy-
lin, laid down in response to the boring activities (Plate
42, Figure 1). In many cases the conchyolar patch is
surrounded by a thickened, warped area of the valve.
Occasionally, where there is multiple infestation in a
limited area of the valve, warping may be so severe as to
nearly occlude the mantle cavity; however, such mussels
appear otherwise normal.

Removal of the borer from its burrow intact may be
accomplished with the careful aid of a hammer and chisel
by fracturing the host’s valve immediately adjacent to the
burrow; this will usually lay open the burrow without
harming the borer, which can then be removed gently with
forceps. In one instance this procedure fortunately revealed
both an immature and an adult stage of Penitella (Plate
42, Figure 2), the former being larger than the latter in this
case. The immature stages of P conradi and other pho-
lads are characterized by gaping valves and a large,

Page 282

THE VELIGER

muscular foot, both vital in the boring process. Upon
attaining the adult stages, the animal ceases boring,
becomes sessile, and the area between the valves becomes
closed off by a thin, calcareous layer known as a callum.
This striking change in morphology during the life cycle
of the animal has lead to considerable confusion in the
taxonomy of pholads in the past, e.g., the immature,
boring stage of P conradi was previously described and
well known as Navaea subglobosa Gray, 1851.

LITERATURE CITED

Turner, RutH Dixon
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.

Vol. 10; No. 3

Tue VELIGER, Vol. 10, No. 3 [MeEREpDITH] Plate 42

Figure 1: Interior of the valve of Mytilus californianus, showing
the dark patches of conchyolin laid down in response to the boring
activity of Penitella conradi, which lie immediately behind
the patches (x 1.5)

Figure 2: A cross section of the valve of Mytilus californianus,
showing the two major stages in the life cycle of Penitella conrad.
In this instance the immature stage individual is the larger. The
gaping, toothed valves of the boring (immature) individual are
obvious, as is the callum of the mature stage, closing off the
anterior end of the sessile (adult) individual (x 5)

Vol. 10; No. 3

THE VELIGER

Page 283

A Radula Muscle Preparation from the Gastropod,

Kelletia kelletur, for Biochemical Assays

HOWARD M. FEDER
Hartnell College, Salinas, California 93901

AND

REUBEN LASKER

United States Bureau of Commercial Fisheries

California Current Resources Laboratory, La Jolla, California 92037

(1 Text figure)

THE MUSCLE BUNDLES, the odontophore muscle complex
or radula muscle, which move the molluscan radula have
been physiologically examined in isolated preparations
from two species of carnivorous gastropods: Busycon
canaliculatum (LINNAEUS, 1758) (Hitz, 1956, 1958), a
snail from the east coast of North America, and Buccinum
undatum Linnaeus, 1758 (FANGE & Mattison, 1957,
1958), a snail common to European waters. These authors
found that the muscle contracts under the influence of
acetylcholine and is relaxed by 5-hydroxytryptamine and
that rhythmic contractions of the muscle occur when
acetylcholine and 5-hydroxytryptamine are added together.
FANGE (1963) and Feper «& Arvipsson (1967) showed
that aqueous extracts of the sea stars Henricia sanguino-
lenta (Mixer, 1776), Marthasterias glacialis (LINNAEUS,
1758) and Asterias rubens LinnaEus, 1758 also induce
strong contractions in the isolated radula muscle of Buc-
cinum undatum. Because of the usefulness of this kind of
radula muscle preparation for bioassay we sought one from
a gastropod of the American west coast. This paper
reports on the preparation and use of the radula muscle
from the carnivorous snail Kelletia kelletii (ForBrs, 1850)
for biochemical assays.

Kelletia kelletii were collected by means of SCUBA at
a depth of about 20 m on sandy ocean bottom, near the
Scripps Institution of Oceanography, La Jolla, California.
The bathymetric range of K. kelletii is known to extend
from 18 to 62m (Assort, 1954). Some of the Kelletia
were subsequently shipped to Hartnell College and used in
experiments there. The radula muscle was prepared and

mounted by a modification of the techniques of FANGE &
Mattison (1958), FANce (1963), Mattison & Arvips-
SON (1966), and FEDER « Arvipsson (1967) for isolating
the radula muscle complex of Buccinum undatum. The
final Buccinum muscle preparation included part of the
proximal proboscis sheath which served for suspension of
the muscle. Kelletia radula muscles prepared in this man-
ner tended to behave erratically and showed a decreased
sensitivity as compared to Buccinum radula muscles. This
fact necessitated that the Kelletia radula muscle and the
inner epithelial lining of the proboscis sheath be teased
free of the sheath; the resulting preparation was more
responsive and less erratic. The preparation was then sus-
pended directly from the epithelial lining to the recording
lever (Figure 1). The chamber was a plastic 30 cc syringe
with the outlet enlarged to permit a rapid outflow when
it was flushed. A lever with an arm 18cm long and
counterweighted with 2.0 g was used. The writing stylus
was a fine glass rod of capillary size with a rounded tip.
The muscle was mounted to the lever at its proximal end
by a cotton thread through the epithelial tissue and to the
glass rod at its oral end by a small loop from the radula
sheath tissue. During perfusion sea water was either re-
moved by aspiration or allowed to overflow the chamber.
All other details of the experimental setup were the same
as those described for radula muscle preparations of Buc-
cinum undatum by FEDER & ArviIpSSON (1967).
Specimens of the sea star Pisaster ochraceus (BRANDT,
1835) were taken either from pilings off the pier at Scripps
Institution of Oceanography, or from rocky reefs off La

Page 284

THE VELIGER

Vol. 10; No. 3

Jolla, and from intertidal areas in Monterey Bay, Califor-
nia. The sea star Pycnopodia helianthoides (BRANDT, 1835)
was collected in intertidal areas in Monterey Bay. Extracts
were obtained by the same procedures used by FEDER &
Arvipsson (1967) for the preparation of Marthasterias
glacialis test solutions. Living sea stars were washed in
tap water, frozen, and then thawed; the resultant cloudy
exudate was collected and centrifuged. The clear super-
natant fluid was used in all tests.

String to Lever

Water Level

Proximal End

Glass Rod

Radula Muscle
Sea Water

Test Chamber
Oral End

Modified Syringe Outlet
mo = Sea Water
ee

ro OOOO OY VOL OO GOO 0 © OY

Air
aS

Rubber Tubing
Pinch Clamp

Drain

Figure 1

Diagram of the mounted radula muscle preparation

The Kelletia radula muscle contracted in the presence
of acetylcholine of which the final concentration was
5 & 10% g/ml. Extracts of Pisaster ochraceus also caused
the radula muscle of Kelletia to contract; the strength of
the contractions varied roughly with amount of extract
added. After addition of acetylcholine or P ochraceus
extract the contraction could be reversed immediately by

3 < 107 g/ml (final concentration) of 5-hydroxytrypt-
amine. Strong, rhythmical contractions followed the initial
relaxation. Similar responses of Kelletia muscle to Pycno-
podia extracts were found in preliminary experiments.
Responses, however, were always of greater magnitude
to Pycnopodia than to Pisaster ochraceus, whether as con-
tractions in response to the extracts or rhythmic contrac-
tions in response to the extracts after addition of 5-hydr-
oxytryptamine. The amplitude of muscle contraction after
additon of Pycnopodia extract was 2 to 3 times greater
than that caused by addition of identical volumes of
Pisaster ochraceus extract.

Sensitivity of muscles varied with time; in general,
older preparations, 24-48 hours old maintained in a
water bath at 10° C, were more sensitive. In addition,
muscle preparations responded only when a certain
“threshold” concentration (40 microliters in a final volume
of 30 ml of sea water) of Pisaster extract was reached. An
addition of sea star extract 5 times the threshold amount
(i.e. 200 microliters in a final volume of 30 ml of sea
water) frequently caused an extremely strong contracture
from which the muscle often did not recover when the
sea water in the bath chamber was renewed. Thus, like
Buccinum muscle preparations, responses were quantitative
only within a narrow range of extract additions.

The successful isolation of the radula muscle complex
of Kelletia kelletu and its subsequent use in assay increases
the number of available preparations of this type to three;
Busycon canaliculatum and Buccinum undatum represent
the other two species. The Kelletia preparation is very
similar to that of Buccinum, and the responses of the
radula muscles of the two species also are similar. It is
probable that other carnivorous snails have radula muscles
suitable for physiological and pharmacological studies.

ACKNOWLEDGMENTS

This work was supported by a National Science Foundation
Research Participation Grant to one of us (H. M. F),
and was accomplished at the Bureau of Commercial
Fisheries, Fishery-Oceanography Center, La Jolla, Cali-
fornia. We thank Mr. Daniel Tam of Hartnell College
and Mr. William Vlymen of the Scripps Institution of
Oceanography for their very able technical assistance, and
Mr. George Crozier of the Scripps Institution of Oceano-
graphy and Mr. Robert Bowers of the Scripps Tuna Ocea-
nography Research Program for collection of experimental
animals.

Vol. 10; No. 3 THE VELIGER Page 285

LITERATURE CITED

Assott, Ropert TUCKER

1954. American seashells. Princeton, New Jersey. D. van
Nostrand Co., Inc.; xiv + 541 pp.; 100 figs.; 40 plts.

FANGE, RAGNAR
1963. Toxic factors in starfishes. Sarsia 10: 19 - 21
FANGE, RAGNAR & ArTUR MATTISSON

1957. Rhythmical activity of a smooth muscle rich in haematin
compounds. Swedish conference on cell research. Ark. f. Zool.
11: 112-113

1958. Studies on the physiology of the radula muscle of Buc-
cinum undatum. Acta Zoologica 39: 53 - 64
FEDER, Howarp M. « J. Arvipsson

1967. Studies on a sea star (Marthasterias glacialis) extract
causing avoidance reactions and radula muscle contractions in
a gastropod (Buccinum undatum). Ark. Zool. 19: 369 - 379

Hitt, Rosert BENJAMIN

1956. Regulation of rhythmic activity in two types of red

muscle in Busycon canaliculatum. Anat. Rec. 125: 119

1958. ‘The effects of certain neurohumors and of other drugs
on the ventricle and radula protractor of Busycon canalicu-
latum and on the ventricle of Strombus gigas. Biol. Bull.
115: 471 - 482

Mattisson, ARTUR & J. ARVIDSSON

1967. Some effects of electrical stimulation and exogenous
metabolites on the contractile activity and the ultrastructure
of the radula-muscle of Buccinum undatum. Zeitschr. fiir

Zellforsch. 73: 37 - 55

Page 286

THE VELIGER

Vol. 10; No. 3

A Remarkable New Cancellariid from the Philippines,

with Comments on Other Taxa

(Gastropoda : Volutacea )

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 43; 2 Text figures)

SOME TIME AGO Mr. and Mrs. Clifford Ames, who reside in
the Philippines, submitted to us a specimen of a cancel-
lariid which had been taken by Mr. Mario Mercado
during trawling operations off Marinduque Island, Phil-
ippines. This distinctive gastropod apparently is an unde-
scribed species. It does not seem to have been reported by
ScHEPMAN (1908-1913) from the collections of the
“Siboga” trawled in Indonesian waters, nor is it repre-
sented in the extensive deep water dredgings of the
“Albatross” in Philippine waters, which are now housed
in the U.S. National Museum, Smithsonian Institution.
The species is named in honor of the collector, Mario
Mercado.

Scalptia JouSSEAUME, 1887

Type species: Scalptia obliquata LaAMarck, 1822, by OD.

Wenz (1938) considered Scalptia to be a subgenus of
Trigonostoma BLAINVILLE, 1827, of which the type species
is Delphinula trigonostoma Lamarck, 1822, by M and
tautonymy. More recent workers, notably Hape (1961a
and 1961 b; 1964), Habe & Kosuce (1966) have accorded
Scalptia full generic recognition.

Scalptia mercadoi OLD, spec. nov.
(Plate 43, Figures 1 to 3; Text figures 1 and 2)

Description of Holotype: (Plate 43, Figure 1) Shell ovate,
with an extended spire forming an angle of 50°. Aperture
less than one half the length of the shell. Shell 1 \easuring
34+ mm in length; 19mm in width. Shell of 64 whorls.
Nucleus of about 24 whorls (partially missing in the
holotype). Nuclear area and first two post-embryonic

whorls translucent, due to cleaning. Body whorl ornamen-
ted with 9 large raised ribs. Ribs rather narrowly attached,
excavated on both leading and trailing edges. Excavation
deeper on leading edge. Penultimate whorl with 10 raised
ribs. Ribs truncate posteriorly, and sutures deeply chan-
neled. Sutures bridged by posterior margin of old varices,
which connect with the varix obliquely above (see Text
figure 1).

Yep

Figure 1

Scalptia mercadoi OLD, spec. nov.

Enlargement of body whorl and penultimate whorl of holotype
showing detail of sutures; x 4
Drawing courtesy of Mr. Anthony D’Attilio

Ribs on body whorl ornamented with raised, spiral
threads, commonly 22 in number and with one or more
weaker threads between. Interstices of ribs also with
transverse threads, from 22 to 24 in number. Interstices

Vol. 10; No. 3

also ornamented with fainter axial sculpturing, imparting
a delicate cancellate effect. Profile of rib strongly convex,
thickest near the trailing edge, and convexity diminishing
towards the leading edge.

Umbilicus deep and profound, entered by anterior at-
tachments of ribs.

Aperture elliptically ovate, with 8 raised spiral cords on
outer wall. Exterior ribs are visible as axial lines. Posterior
canal with one strong, grooved rib. Three spiral plaits on
columella, of which the uppermost plait is the strongest.

Color of shell light orange. Body whorl ornamented
with 2 conspicuous bands of light purplish-brown, one at
periphery, the other between the periphery and the ante-
rior angulation of the ribs. A third, fainter band of pig-
mentation is indicated at posterior or uppermost angula-
tions of ribs. Aperture bluish-white, shading to light
orange on the columella.

From a portion of the partially decomposed animal,
the holotype was determined to be a male.

Periostracum unknown .

Type locality: Trawled in 175 fathoms, off Gasan,
Marinduque Island, Philippines. November 2, 1965.
Type depository: American Museum of Natural History,
no. 138276.

Descriptions of paratypes: Paratype “A.” (Plate 43,
Figure 3). Shell of 35.4mm in height, 18.5 mm in width.
Shell of 84 whorls. Nucleus intact, and consisting of 24
whorls. Nucleus polished and tan in color. Body whorl
with 10 ribs, more crowded than those of the body whorl
of the holotype. It was collected in Tayabas Bay, Quezon,
Philippines, on November 14, 1966. Paratype “A” in
collection of Mr.and Mrs. Crawford N. Cate, no. CA 107,
ex Romeo Lumawig.

Paratype ““B” (Plate 43, Figure 2) measures 26mm in
height and 16.4mm in width. The coloration is darker
than in either of the other specimens, the transverse
bands being of a dark purplish-brown, and the orange
pigmentation is more livid. The interstices are suffused
with solid purplish-brown, the transverse banding being
faintly discernible as darker coloration. The umbilicus is
almost completely closed. The columella is colored a pale
orange shading to dark purplish-brown in the area of
the plaits. It was trawled in 120 fathoms, Tayabas Bay,
Luzon, Philippines, in 1966. The radula of this specimen
was extracted and sketched by Mr. Masao Azuma (see
Text figure 2). Paratype “B” in the collection of Mr.
Victor Dan, of Manila.

Additional, non-typological specimens are in the col-
lection of Mr. F G. Dayrit of Manila (figured by Dayrir,
1967, p. 4, fig. 4).

Remarks: The new species differs markedly from its con-
geners by its larger size, ornate ribbing, and coloring.

THE VELIGER

Page 287

Scalptia mercadoi is perhaps closest in relationship to S.
textilis (KiENER, 1841), and to S. scalariformis (La-

Figure 2

Scalptia mercadoi OLD, spec. nov.

Enlargement of radula (approximately x 480)
Length about 3mm, width 2. Drawing courtesy of
Mr. Anthony D’Attilio, from a sketch by Mr. Masao Azuma

Page 288

MARCK, 1822). KreneEr’s species also occurs in the Phil-
ippines, but it inhabits shallower water than does S.
mercadoi. LaMARCK (1822) briefly described Cancellaria
scalariformis without citing figures or localities. SoWERBY
(1833) stated, “{Lamarck’s] C. scalariformis is unknown
to me.” DesHayes & Mitne-Epwarps (1843) repeated
Lamarck’s description, citing figures of KreENER (1841).
These figures seem to be referable to specimens in the
American Museum which appear to be Lamarck’s taxon.

Other figures and descriptions of related taxa which
somewhat resemble the new species, are those of Cancel-
laria funiculata Hinns, 1844 and of C. lyrata ADAMS &
Reeve, 1848 (non Broccut, 1814, nor Borson, 1820).
Cancellaria funiculata was described as having been taken
in 7 fathoms, Gulf of Magdalena [Mexico]. The locality
of C. lyrata was given as “China Seas,” which REEVE
(1856) and Loppecke (1887) believed to be erroneous.
Reeve, Loppecke, and Crosse (1861) considered C.
lyrata of ADAMS & REEVE to be synonymous with Cancel-
laria funiculata Hinps. KEEN (1958) questioned the Pan-
amic occurrence of Trigonostoma funiculata (Hinps),
but this sentence has been deleted from the 1960 edition,
probably due to the examination of material with reliable
locality data from the Panamic marine province. The
American Museum has 8 specimens taken in 30 fathoms,
by the Templeton Crocker Expedition at Manzanillo,
Mexico in 1938. These specimens appear to be typical
specimens of Hind’s funiculata.

In recent years, both of these specific taxa also have
been applied to shells occurring in the western Pacific.
Kuropa & HABE (1952), however, queried the occurrence
of Trigonostoma funiculata (Hinps) in Japanese waters.
Hase (196la) proposed the generic name Nipponaphera
with N. lyrata (ADAMS & REEVE) as the type species.
Haze (1961b, 1964) repeated the same figure (as in
1961a), but employed Hind’s taxon, N. funiculata.
Oyama (1963) also figured a Japanese specimen under
this name. Japanese shells which have been referred to
N. funiculata (Hinps) are probably specimens of N.
iwaotaku Hae, 1961, though Haze (1961 b) cited sculp-

THE VELIGER

Vol. 10; No. 3

tural differences between his species and shells which had
been identified as N. funiculata from Japanese waters.

The taxon Cancellaria lyrata also has been employed
for a species from the Tertiary of Europe. Broccut, 1814,
had described the fossil from the Piedmont district of
Italy as Voluta lyrata. Borson (1820) subsequently used
the combination Cancellaria lyrata for Brocchi’s species
(fide Sacco, 1894). Most workers have placed Brocchi’s
lyrata in the genus (or subgenus) Sveliia JouSSEAUME,
1887, although Strausz (1966) recently referred the
fossil species to the subgenus Calcarata JouSSEAUME,
1887, which he placed in Cancellaria (s.1.).

In the present century, the taxon Sveltia lyrata
(Broccu1) also has been applied to a Recent species that
is rather rarely trawled off western Africa. DAUTZENBERG
& FiscHER (1906) cited a specimen from 628 m near
Maio, Cape Verde; ApAM & KNupsEN (1955) cited 4
examples from 145m, 210m, and 230m off Cabinda;
MarcHe-MarcHap (1958) off Cape Verde, GraHam
(1966) cited material from 170 - 200 fathoms, in the Gulf
of Guinea; and Barnarp (1958, 1959) off South West
Africa. BARNARD questioned the use of Brocchi’s taxon
for the Recent mollusk, It is unlikely that the Recent
species and the Mio-Pliocene fossil from northern Italy
and Central Europe are conspecific.

ACKNOWLEDGMENTS

I wish to thank the following people for assistance of
varous kinds: Mr. and Mrs. Clifford Ames, Mr. Victor
Dan, and Mr. Mario Mercado, of the Philippines; Mr.
Masao Azuma of Nishinomiya, Japan; Dr. C. A. Fleming
of Wellington, New Zealand; Mr. «& Mrs. Crawford
N. Cate of Los Angeles, California; Mr. Richard Petit
of Ocean Drive Beach, South Carolina.

I am particularly indebted to my colleagues at the
American Museum of Natural History, Dr. William K.
Emerson, Mr. Anthony D’Attilio, and Mrs. Margaret
Richards for various courtesies.

Explanation of Plate 43

Scalptia mercadoi OLp, spec. nov.

Figure 1: Holotype (AMNH no. 138276) ; trawled in 175 fathoms,
off Gasan, Marinduque Island, Philippines (x 2). c: Basal view
of holotype (x 2)

Figure 2:

Paratype “B” (Victor Dan collection); trawled in

120 fathoms, Tayabas Bay, Quezon, Philippines. (x 2)

Figure 3:

Paratype “A” (Crawford and Jean Cate collection) ;

Tayabas Bay, Quezon, Philippines, a: view of apex (approximately
x 10); b and c (x 2)
Photographs courtesy of American Museum of Natural History

Tue VELIGER, Vol. 10, No. 3 [Op] Plate 43

Figure 2 a Figure 2 b

Figure 1 c

Figure 3 c

Vol. 10; No. 3

LITERATURE CITED

Apam, WILLIAM & JORGEN KNUDSEN
1955. Note sur quelques espéces de mollusques marins nou-
veaux ou peu connus de |’Afrique Occidentale. Inst. Roy.
Sci. Nat. Belgique 31 (61): 1-24; 2 plts.
Apams, ARTHUR & Lovett Aucustus REEVE in A. ADAMS
1848. The zoology of the voyage of H. M.S. Samarang, under
the command of Captain Sir E. Belcher ... Mollusca. 1 - 87;
plts. 1 - 24
BarNaRD, KEpPELL HARCOURT
1958. The radula of Cancellaria.
24 (7): 243-244
1959. Contributions to the knowledge of South African marine
Mollusca, Part II :Gastropoda: Prosobranchiata: Rhachiglossa.
Ann. South Afric. Mus. 45 (1): 1-237; 52 figs. (June)
BLaInvitLE, Henri Marre DucrotTay DE
- 1827. Manuel de malacologie et de conchyliologie.
649 - 664; pits. 1 - 87
BorsoNn, STEFANO
1820. Saggio di orittografia Piemontese.
Sci. Torino 25: 180 - 229
[not seen]
Broccut, GIovANNI BATTISTA
1814. | Conchiologia fossile Subapennina con osservazioni geo-

Milano (Dalla
1-56 +i-Ixxx +1-240; 2: 241-712;

Journ. of Conch.

Mem. Real. Accad.

logiche sugli Apennini e sula suolo adiacente.
Stamp. Reale) 1:

pits. 1- 16
[not seen]
1843. | Conchiologia fossile Subapennina con osservazioni geo-
logiche sugli Apennini e sula suolo adiacente. 2%? ed.
Crosse, H.
1861. Etude sur le genre Cancellaire, suivie du catalogue des
espéces vivantes et fossiles actuellement connues. Journ.

de Conchyl. 9: 220 - 256
DauTZENBERG, PHILIPPE & HENRI FISCHER
1906. Mollusques provenant des dragages effectués a l’ouest
de l’Afrique. In Résultats des campagnes scientifiques du
Prince de Monaco 32: 3 - 125; plts. 1-5
Dayrit, FERNANDO G.
1967. in Hawaiian Shell News 15 (3): 3, 4
Desuayes, Gérarp Paut « HENRt MitNE-EDWARDS
1843. Histoire naturelle des animaux sans vertébres. 2. édit.
Paris, 9: 1-728
GraHAM, ALASTAIR
1966. The foregut of some marginellid and cancellariid pro-
sobranchs. Stud. Trop. Oceanogr. Miami 4 (1): 134-151
Hasse, TADASHIGE
1961a. Coloured illustrations of the shells of Japan II. 1 to
183; 66 colored plts.
1961b. Description of four new cancellariid species, with a list
of the Japanese species of the family Cancellariidae. Venus
21 (4): 431-441; plts. 23, 24
1964. Shells of the western Pacific in color.
233 pp.; 66 plts.
Hase, TADASHIGE & SADAO KosucE
1966. Shells of the world in colour, vol. II: The tropical
Pacific

Osaka, vol. II,

THE VELIGER

Page 289

Hinps, Ricuarp BrinsLey
1844-45. The zoology of the voyage of H. M.S. Sulphur
Mollusca, pt. 1, pp. 1-24, pls. 1-7 (July, 1844) ; pt. 2, pp. 25-48,
pls. 8-14 (Oct. 1844) ; pt. 3, pp. 49-72, pls. 15-21 (Jan. 1845)
[exact dates from a copy with original wrappers in the library
of the California Academy of Sciences].
JoussEaumE, F. P.
1887. La famille des Cancellariidae. Le Naturaliste “9”
[th year; second ser., first year ]: 155-157; 192 - 194; 213-214;
221 - 223
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)
1960. Sea shells of tropical west America, 2% printing.
Kiener, Louis CuHaries
1841. | Genre Cancellaire. “vol. 6”:
Fide SHERBORN & Woopwarp, 1901
Kuropa, TokusBeI & TADASHIGE HABE
1952. Check list and bibliography of the Recent marine mol-
lusca of Japan. pp. 1-210; 2 maps. Tokyo, Japan.
(4 April 1952)
Lamarck, JEAN-BaptisTe PrrrrE 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)

1-44; plts 1-9

L6BBECKE, T.

[1881-] 1887. Das Genus Cancellaria [Admete by W. Koper] in

Systematisches Conchylien-Cabinet, 4 (4): 1-108; plts. 1-24
REEveE, Lovett Aucustus

1856. | Conchologia iconica; or illustrations of the shells of
molluscous animals. Monograph of the genus Cancellaria.
vol. 10; 18 plates

Sacco, FEDERICO

1894. I molluschi . del Piemonte e della Liguria (16).

Cancellariidae. 1-78, 3 plts.
ScHEPMAN, MattrHeus Marinus

1908-1913. The Prosobranchia, Pulmonata and Opisthobranchia
Tectibranchiata of the Siboga-Expedition. In Siboga-Expeditie
49: 1-494; plts. 1 - 32

SHaw, H.O.N.

1909. On the dates of issue of Sowerby’s ‘“Conchological

Illustrations.” Proc. Malacol. Soc. London 8: 333 - 340
SHERBORN, CHARLES Davies & BERNHARD BARHAM WooDWARD

1901. __ Notes on the dates of publication of the parts of Kiener’s
“Spécies général et iconographie des coquilles vivantes,” etc.
(1834-80). Proc. Malac. Soc. London 4 (5): 216-219

(25 July 1901)
Sowerby, GrorcE BRETTINGHAM, Jr.

1832 - 1833 [- 1841] The conchological illustrations. Cancellaria
[parts 9- 13 appeared from November 30, 1832 to January 4,
1833. Catalogue, January 11, 1833]. Fide SHaw, 1909

Strausz, LAsz_6

1966. Die Miozan-Mediterranen Gastropoden Ungarns.

Budapest: 7 - 692 (incl. plts.)
WENz, WILHELM

1943. | Handbuch der Paldozoologie (O.H. Schindewolf, ed.)
6(1) (allgem. Teil und Prosobr.). Teil 6 (=Lief. 8) : 1201-1506;
text figs. 3417-4211. (October 1943) Berlin (Gebr. Born-
traeger Verl.)

Page 290

THE VELIGER

Vol. 10; No. 3

A Ctenostomatous Ectoproct Epizoic on the Chiton

Ischnochiton mertensi1

BY

EUGENE S. HELFMAN

Department of Zoology, University of California, Berkeley, California

94720

(2 Text figures)

DurING THE SUMMER Of 1966 I collected 167 chitons of
18 species; collections were made both intertidally and
subtidally in the region of Bodega Bay, Sonoma County,
California. On three individuals of one species, Ischno-
chiton mertcnsi (MippENporrr, 1846), stolonifcrous ecto-
procts of the species Farella clongata (vAN BENEDEN,
1845) were found attached to the ventral surface of the
girdle, while all other chitons observed were free from
any similar attachment. A review of the available litera-
ture has not revealed any descriptions of an ectoproct-
amphincuran relationship of the nature discussed here.

Ospurn, 1963, described the habitat of Farclla elon-
gata as intertidal, where the species is said to form a dense,
mat-like network of colonies on the substrate. O’DoNnoc-
HUE & O’Donocnue, 1923, found it on Pinnixa species,
but failed to give any details. As found on Ischnochiton
mertensu, the ectoproct colony consists of several 1.0mm
zooecia arising from one or a few clongated stolons. A
stolon with its branching polypides may extend half the
length of a 4.0 cm long chiton, giving off other stolons at
various placcs along the girdle (Figure 1). The organ of
attachment is the stolon, the polypides being free from
the substrate.

The sites of attachment of the cctoprocts varied on the
three chitons affected. The longest stolons and greatest
number of polypides were found in the slight depression
of the ventral girdle tissue bordering the pallial groove
(Figure 2). Branches of these stolons and some individual
polypides were located on the flat portion of the ventral
girdle, while isolated polypides were found along the
edge of the girdle on two of the three chitons. The ap-
proximate number of cctoprocts on cach chiton was 100,
62, and 20. No individuals of Farella elongata were found
on the dorsal surface of the chitons, nor were any other
ectoprocts found on the ventral surfaces.

Several of the polypides that bordered the pallial cavity
projected their tentacles into the chiton’s incurrent groove

I cm

Figure 1

Ventral aspect of Ischnochiton mertensii (MippENpoRFF, 1846)

Stippled areas show the gencral configuration of approximately

100 attached ectoprocts of the species Farella clongata (van BENE-
bEN, 1845). From a photograph of the live animals x3

and contacted the ctenidial leaflets. These ectoprocts fil-
tered out and ingested particles flowing in the current of
the mantle cavity. The tentacles in contact with the
ctenidial leaflets projected outward from the ectoproct’s

Vol. 10; No. 3

THE VELIGER

Page 291

aperture like a cone, with cilia on the inward-directed
surfaces of the 14 tentacles The distal tips of the tenta-
cles touched the ctenidial leaflets, and particles were seen
to pass ventrally along the cilia of the leaflet, onto the
ciliated inner surface of the tentacles and toward the
mouth of the ectoproct, at which time the tentacles would
be quickly retracted and the particles ingested or rejected.

Figure 2

Enlargement of the inset in Figure 1
The zooecia of Farella elongata, with the apertures of the five
polypides oriented toward the pallial cavity on the left side of the
chiton. From a photograph of the live animals x 25

The literature revealed two instances of branchial cavity
attachment by organisms living in a way similar to that
of Farella elongata. OsBURN, 1953, reported that the
ectoproct Triticella elongata had been found living in the
gill chambers of the pinnotherid crab Scleroplax granulata.
SOULE & SOULE, 1965, reported two species of loxosomatid
entoprocts epizoic upon the gill filaments of a mantis
shrimp from Southern California.

Daves, 1966, stated that many ectoprocts “may be
found as epibionts on other animals, but the associations
are not usually specific;’ and ApEcoKE, 1967, reported

that two opposing schools of thought had developed, one
regarding ectoproct-molluscan associations to be fortui-
tous, the other maintaining that ectoproct species are very
specific in their choice of substrate. A statement on the
possible specificity of the present association will have
to await more extensive collection and observation.

ACKNOWLEDGMENTS

I would like to thank Professor Cadet Hand of the Depart-
ment of Zoology, University of California, Berkeley, and
Professor Charles R. Stasek, Department of Biological
Science, Florida State University, for their suggestions
and criticisms. Professor John D. Soule of the Allan
Hancock Foundation, University of Southern California,
identified the ectoproct from slides prepared by Mr. Lloyd
Austin of the Department of Zoology, University of Cali-
fornia, Berkeley.

LITERATURE CITED

ADEGOKE, OLUWAFEYISOLA SYLVESTER
1967. Bryozoan - mollusk relationships.
298 - 300; plt. 40

Dates, R. PHILuip
1966. Symbiosis in marine organisms. pp. 299 - 322 in S. Mark
Henry (cd.) Symbiosis, Vol. 1: Associations of microorganisms,
plants, and marine organisms. 478 pp., illus. Acad. Press,
New York

O’DonocuHvuE, Cuartes Henry & Erste O’DoNoGHUE
1923. A preliminary list of the Polyzoa (Bryozoa) from the
Vancouver Island region. Contrib. Canad. Biol. Fish., n.s.
1: 143 - 201; plts. 1-4
Ossurn, RayMmonp C.
1953. Bryozoa of the Pacific coast of America, part 3, Cyclo-

stomata, Ctenostomata, Entoprocta, and addenda. Allan
Hancock Pacific Exped. 14 (3): 613 - 841; plts. 65 - 82

Sou.e, Dorotuy FE « Joun D. Sous
1965. Two new species of Loxosomcella, Entoprocta, epizoic on
crustacea. Allan Hancock Found. Publ. Occas. Paper no.
29: 1-19; frontispiece, figs. 1 - 2

The Veliger 9 (3) :
(1 January 1967)

Page 292

NOTES & NEWS

New Record of Conus ebraeus
in Costa Rica

BY

JOSEPH R. HOUBRICK

Department of Science and Mathematics
Saint Leo College, Saint Leo, Florida 33574

Conus cbracus LINNAEuS, 1758, an Indo-Pacific cone, is
known to range in the eastern Pacific to Clipperton Island
and also to the Galapagos Islands. Keen (1958) and
Hanna (1963) both stated that Mr. Ted Dranga found
it living in Costa Rica. There seem to be no other
published records of C’. ebraeus from the Central Ameri-
can coast. On March 18, 1965 I collected two living
specimens of C. ebraeus LINNAEUS under rocks in the
intertidal zone one mile south of Playas del Coco, Guana-
caste Province, Costa Rica. These specimens have been
deposited in the Museum of Comparative Zoology, Har-
vard University.

LITERATURE CITED

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

Spawning Notes, I.

Hexaplex erythrostomus

BY

FAY HENRY WOLFSON

Associate Curator of Conchology
San Diego Museum of Natural History, San Diego
California 92115

(1 Text figure)

Hexaplex erythrostomus (Swatnson, 1831) was encoun-
tered only once during intensive collecting (11 and 5
days, respectively) in June and July 1967 in Bahia de los
Angeles, Baja California: on July 20, I came upon a

THE VELIGER

Vol. 10; No. 3

group of seven individuals in the process of depositing
egg masses on rock substrates. Local residents inform me
that this was the beginning of an annual spawning migra-
tion from the depths by this species. Two egg masses,
each within a single mussel valve, were also found on
the reef.

In September, specimens ranging from 1” to 4” were
present throughout the bay, 4 egg masses, all deposited
in mussel valves were found, and an individual was ob-
served spawning on a small rock while two others were
copulating beside it.

One irregularly shaped egg mass measured 100mm by
70mm by 40mm (greatest dimensions) and consisted of
approximately 400 curved, tongue-shaped capsules, the
base of each attached in an irregular manner to one or
two underlying capsules. A few short stalks cement the
mass to the substrate. Individual capsules are roughly of
the same size (see Figure 1), averaging 6mm in length

Figure 1

and 1mm in breadth; average width is 6mm at the base
of the “tongue,” 3mm at the tip. The upper surface
bears a groove with rounded edges, the lower is smooth
and convex. The base color is creamy white, when fresh,
but some capsules (generally those at the base of the
mass) are tinged with varying amounts of pink, ranging
from the shade characteristic of the species to magenta.
The eggs exhibit the same range of pigmentation, which
may be indicative of the degree of development. There
are over 100 eggs per capsule.

The mass apparently serves as a symbiotic habitat: a
lcm amphipod, an 8mm Hormomya and several cope-
pods were lodged between the capsules of the masses
collected.

Jone INE, 1).

The thirty-third annual mecting of the Amcrican Malaco-
logical Union was held at Ottawa, Ontario, Canada July
31** to August 5™, 1967. It was the best attended mecting
in AMU history as 163 persons listened to a record

Vol. 10; No. 3

THE VELIGER

Page 293

number of papers and enjoyed such added features as an
all day field trip to an Ottawa River site, a banquet as
guests of the National Museum of Canada and a trip to
Montreal and its famed Expo 6/7.
Officers elected to serve until July, 1968 are:
President — ARTHUR H. CxarKE, Jr.
Vice-president -— JosEPH ROSEWATER
Second Vice-president — Fay WoLFSON
Secretary — Marcaret C. TESKEY
Treasurer — Mrs. H. B. BAKER
Publications Editor — M. Karu JAcoBson
Councillors-at-Large — DorotTHy BEETLE, Harorp D.
Murray, Davin H. STANSBERY, DAN STEGER.
The 1968 meeting will be held July 15 to 19 at Corpus
Christi, Texas, at which time six Texas Shell Clubs will
join ranks to act as official hosts to AMU members, their
families and friends. Pacific Division members especially
are urged to attend. Detailed notices will be mailed in
April, 1968.
M. C. Teskey

COAN Molluscan Collection

at University of California, Davis

BY
JAMES W. VALENTINE

Department of Geology, University of California
Davis, California 95616

A collection of marine Mollusca from the eastern Pacific
Ocean, assembled over many years by Eugene V. Coan,
has been acquired by the Department of Geology,
University of California, Davis. It is desirable to call
attention to the present location of this collection, for
it contains specimens that form the basis of published
records (however, the shells from Santa Barbara County
recorded by Coan, 1964, are not included). At this
writing the Coan material at Davis is being catalogued
and will be available for consultation soon.

With the acquisition of the Coan collection the Depart-
ment of Geology at Davis has embarked upon a long-
range program of building and maintaining a permanent
reference (research) collection of Recent and fossil in-
vertebrates and protists. The collection will be continually
expanded and lodged, under the care of a professional
curator, in a geology building now being designed. Storage
of type material and of special collections will be pro-
vided for. We welcome the use of the collection by
students of the Invertebrata and Protista.

LITERATURE CITED

Coan, EucENE VicToR
1964. | The Mollusca of the Santa Barbara County area. Part
I.—Pelecypoda and Scaphopoda. The Veliger 7(1): 29-33.
(1 July 1964)

CALIFORNIA
MALACOZOOLOGICAL SocIETY, Inc.

is a non-profit educational corporation (Articles of In-
corporation No. 463389 were filed January 6, 1964 in
the office of the Secretary of State). The Society publishes
a scientific quarterly, the VELIGER. Donations to the
Society are used to pay a part of the production costs and
thus to keep the subscription rate at a minimum. Donors
may designate the Fund to which their contribution is
to be credited: Operating Fund (available for current
production) ; Savings Fund (available only for specified
purposes, such as publication of especially long and signi-
ficant papers); Endowment Fund (the income from
which is available. The principal is irrevocably dedicated
to scientific and educational purposes). Unassigned dona-
tions will be used according to greatest need.

Contributions to the C.M.S., Inc. are deductible by
donors as provided in section 170 of the Internal Revenue
Code (for Federal income tax purposes). Bequests, lega-
cies, gifts, devices are deductible for Federal estate and
gift tax purposes under section 2055, 2106, and 2522 of
the Code. The Treasurer of the C. M. S., Inc. will issue
suitable receipts which may be used by Donors to substan-
tiate their respective tax deductions.

CauiFoRNIA MatacozooLocicaL Society, Inc.
announces:

Backnumbers of

THE VELIGER

and other publications
Volume 9: $22.-

Supplement to Volume 3: $6.-* plus $-.35 handling sharge
[Opisthobranch Mollusks of California
by Prof. Ernst Marcus]

Supplement to Volume 6: $4.-* plus $-.35 mailing charge
[Biology of Tegula funebralis (A. ApaMs), edited by
Abbott, Blinks, Phillips and Stohler]

Volumes 1 through 8: out of print

Page 294

Supplement to Volume 7: $2.-* plus $-.35 mailing charge

[Glossary of A Thousand-and-One Terms used in
Conchology, compiled by Wintrrep H. ARNOLD]

Items marked with * are subject to sales tax in the State
of California; residents of that State please add the
appropriate amount to their remittances.

Prices subject to change without notice.

Send orders with remittance to:

Mrs. Jean M. Cate, Manager, 12719 San Vicente Boule-

vard, Los Angeles, Calif. 90049. Please, make checks
payable to C. M.S., Inc.

Shipments of material ordered are generally made within
two weeks after receipt of remittance.

Backnumbers of the current volume will be mailed to new
subscribers, as well as to those who renew late, on the
first working day of the month following receipt of the
remittance. The same policy applies to new members.

Subscription to Volume 10: $12.- domestic; $12.60 in
Canada, Mexico, Central and South America;
$ 12.90 all other foreign countries.
Subscriptions to Volume 10 accepted until March 15, 1968.

Subscription to Volume 11: $18.- domestic; $18.80 in
Canada, Mexico, Central and South America;
$19.20 all other foreign coutries.

Affiliate Membership in the C. M.S., Inc. is $6.- for
the fiscal year July 1, 1967 to June 30, 1968. Postage for
members living in Canada, Mexico, Central and South
America 60 cents, for members living in any other foreign
country 90 cents additional. Membership open to indi-
viduals only - no institutional memberships. Please,
send for membership application forms to the Manager
or the Editor.

At a Regular Membership Meeting of the CALirorNIA
MatacozooLocicaL Society, Inc. the following policies
were adopted by unanimous vote:

There will be an initiation fee of $2.- (in addition to
the annual dues) for persons joining the Society on or
after January 1, 1967.

Members receive The Veliger free of further charges and
are entitled to purchase one copy of any supplement pub-
lished during the current membership year at a special
discount (to be determined for each supplement).

THE VELIGER

Vol. 10; No. 3

Membership renewals are due on or before April 15
each year. If renewal payments are made after April 15
but before March 15 of the following year, there will be
a re-instatement fee of $1.-. Members whose dues pay-
ments (including the re-instatement fee) have not been
received by the latter date, will be dropped from the rolls
of the Society. They may rejoin by paying a new initiation
fee. The volume(s) published during the time a member
was in arrears may be purchased, if still available, at the
regular full volume price plus applicable handling charges.

BOOKS, PERIODICALS, PAMPHLETS

Cretaceous Thyasira from the Western Interior
of North America

by Erte G. KAurrMan. Smithsonian Miscellaneous Col-
lections, Volume 152, No. 1, Smithsonian Publication
4695; pp. 1 to 159; plates 1 too, text) figures ironic:
June 30, 1967.

This paper deals especially with species of Thyasira of
Cretaceous age. Seven species and 10 subspecies from the
western interior of North America. The geologic occur-
rences and relationships of these forms are described
in detail.

Pages 12 to 49 contain much information concerning
the anatomy, morphology, variation, ecology, biotic asso-
ciation and general biology of various living species of
this genus. The author mentions that most of the living
species of Thyasira occur in the deep inner or outer
sublittoral and bathyal environments of cool to cold
water, on a bottom of soft mud lacking coarse clastic
material.

LGH

Notes on Hydrobia totteni

by Grorce M. Davis. “Venus”, the Japanese Journal of
Malacology, vol. 25, pp. 27 to 42,6 text figures. June 1966.

The author points up the characters based on which this
species is considered specifically different from its closely
related European ally, Hydrobia ventricosa. The excellent
illustrations accompanying the discussion deal mostly with
the morphology of the eastern American species. What
appears to be a complete synonymy accompanies the
article.

RS

Vol. 10; No. 3

THE VELIGER

Page 295

Marwick’s Illustrations of New Zealand Shells,
with a checklist of New Zealand Cenozoic Mollusca

by C. A. Femina. New Zealand Department of Scienti-
fic and Industrial Research, Bulletin 173, pages 1 to 456,
frontispiece (portrait), and 145 plates; December, 1966.
R. E. Owen, government printer, Wellington, New Zea-
land. Price 45 shillings ($4.50).

This fine handbook of the New Zealand Geological
Survey was prepared in honor of John A. Marwick
of that organization, on his seventy fifth birthday. The
major divisions of the volume are: introduction; check-
list of New Zealand Cenozoic mollusca; biblography;
plates with explanations, references, localities, geologic
ages; and an index. The line drawings of shells illustrated
by Marwick in various publications (but not including
those originally published in halftone) are republished,
and in addition, a few drawings by others.

This very useful work, such as we have become accus-
tomed to expect from Dr. Fleming, will be a great aid to
all workers who have occasion to refer to the Cenozoic
mollusks of New Zealand. The checklist, bibliography,
the many illustrations (1753 figures), as well as an index
to families, genera, and species, all combine to furnish
not only a valuable handbook but also a suitable tribute
to John A. Marwick.

LGH

Van Nostrand’s Standard Catalog of Shells

by J. L. Wacner & R. Tucker Assott. D. Van Nostrand
Co., Princeton, N. J., second edition, October 12, 1967. —
$5.95.

The content of the new edition of this work is basically
as described in my review of the first edition (The Veliger,
vol. 7, no. 4, p. 255, April 1, 1965); thus, all of my
earlier critical comments still apply. Some new material
has been added to the Catalog, such as a list of the
Marginellidae, lists of selected genera in Tonnacea, Muri-
cacea, and Buccinacea, Neritacea, Tridacnacea, and a
few others. The new catalog now runs to 303 pages. To
the “Quick Lists” two new areas have been added — the
British Isles and South Australia. Some of the inequities
in suggested values in these lists have been smoothed: I
was relieved to find that the rarer abalones now are
valued higher than commoner ones and that Haloconcha
reflexa, an Arctic lacunid not yet in the Stanford collec-
tion, has gone up from 12 cents to 80 cents. On the other
hand, Mysella pedroana, a very ordinary-looking small
commensal clam, remains at 3 dollars, with or without
data. Hyalina californica in the “Quick List” is valued at

4 cents (3 cents if dataless) ; in the Check List the name
is cited as a synonym of Volvarina taeniolata and the
value cited as 15 cents (10 cents without data).

The World Size Records now form a separate section,
with a list running to eight pages of presumed records.
Although not stated in the explanation of the list, the
1950-1959 records were culled from lists in the Minutes
of the Conchological Club of Southern California. The
implication is, rather, that these records were submitted
especially to the present editors of this compilation. It
is disconcerting, therefore, to find “A. M. Keen” listed
as owner of a Clinocardium specimen that actually is and
has been for the last twenty years or more in the Stanford
University collection. This was made clear at the time the
record was submitted, in 1950, to the Conchological Club
editor. Others of the “owners” in the 1950 records are no
longer living. This column in the list, therefore, should be
differently captioned.

As indicated in the previous review, it is probable that
the most useful part of this catalog will prove to be the
check lists of genera and families, even though the total
is yet far from being complete and comprises only some of
the more showy molluscan groups.

MK

Chitons and Gastropods
(Haliotidae through Adeorbidae)
from the Western Pacific Islands

by Harry S. Lapp. U.S. Geological Survey Professional
Paper 531, 98 pp., 16 pls., 1966

About 200 molluscan species and subspecies are discussed
in this initial report on late Tertiary and Quaternary
horizons in seven Island groups of the western Pacific.
Three new subgenera are proposed: Vitiastraca, subgenus
of Astraca (family Turbinidae) ; Subditotectarius, sub-
genus of Tectarius (family Littorinidae) -— both from
the lower Miocene; and Ailinzebina, subgenus of Zebina
(family Rissoidac), living, Marshall Islands. Some 75
specics and subspccies are described as new and illustrated
by photographs of exceptional quality.

The introduction has a useful review of known records
of Cenozoic fossil mollusks from this region, an area in
which relatively little collecting has been done. Material
is yet too scanty to justify generalizations on migration
or dispersal routes, but the occurrence of a genus hither-
to known only in southern Baja California - Haplo-
cochhas — and another — Arene — that is largely
confined to tropical Amcrica, suggests that movement
from cast to west may at times have been possible.

MK

THE VELIGER

Page 296

Pliocene Fossils from Rancho El Refugio,
Baja California, and Cerralvo Island, Mexico

by Leo Georcr Hert ein. Proc. Calif. Acad. Sci. vol. 30,
no. 14, pp. 265 to 284, figs. 1 to 17. 30 Nov. 1966

Two new subspecies, one of Ostrea and one of Chlamys,
are described. A summary of earlier work on the fossil
fauna of the two localities is given. In analyzing the
lists of species recorded, the author finds that the Recent
fauna in the adjacent waters is very similar.

RS

Dimensions and Shapes of Larvae
of some Marine Bivalve Mollusks

by Victor L. Loosanorr, Harry C. Davis « Pau E.
CuHantey. Malacologia, vol. 4, no. 2, pp. 351 to 435; 61
text figs. (halftones and graphs).

Based on recent developments in methods of rearing
bivalve mollusk larvae, developed by Dr. Loosanoff and
his co-workers, the authors report on the observations on

Vol. 10; No. 3

20 different species. Larvae were grown past metamor-
phosis from eggs or recently released larvae of known
parents. Data concerning each of the 20 species studied
are given. A number of problems in identification, par-
ticularly of closely related forms are discussed. Helpful
suggestions for improved methods of identification are
given.

RS

Der Status zweier von Peter Merian beschriebener
Arten von Gastropoden

by Peter Junc. Verhandl. Basler Naturforsch. Gesellsch.,
vol. 77, pp. 76 to 81; 8 text figs. | June 30, 1966.

Two fossil species of Gastropods, obtained in the year
1843 from Jamaica and Antigua respectively, were de-
scribed by MeEriAn in 1844 as Terebellum procerum and
Scalaria melanoidea. The latter taxon was a nomen dubi-
um, while the former is apparently a valid species. In
1914, Brown & Pitssry described Hemisinus antiguensis,
using the material that Merian had had.

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, distributional, ecological, histological, morphological, phys-
iological, taxonomic, etc., aspects of marine, freshwater or terrestrial
mollusks from any region, will be considered. Even topics only indi-
rectly concerned with mollusks may be acceptable. In the unlikely event
that space considerations make limitations necessary, papers dealing
with mollusks from the Pacific region will be given priority. However,
in this case the term “Pacific region” is to be most liberally interpreted.

It is the editorial policy to preserve the individualistic writing style of
the author; therefore any editorial changes in a manuscript will be sub-
mitted to the author for his approval, before going to press.

Short articles containing descriptions of new species or lesser taxa will
be given preferential treatment in the speed of publication provided
that arrangements have been made by the author for depositing the
holotype with a recognized public Museum. Museum numbers of the
type specimens must be included in the manuscript. Type localities
must be defined as accurately as possible, with geographical longitudes
and latitudes added.

Short original papers, not exceeding 500 words, will be published in
the column “NOTES & NEWS’; in this column will also appear notices
of meetings of the American Malacological Union, as well as news items
which are deemed of interest to our subscribers in general. Articles on
“METHODS & TECHNIQUES?” will be considered for publication in
another column, provided that the information is complete and tech-
niques and methods are capable of duplication by anyone carefully fol-
lowing the description given. Such articles should be mainly original
and deal with collecting, preparing, maintaining, studying, photo-
graphing, etc., of mollusks or other invertebrates. A third column, en-
titled “INFORMATION DESK,” will contain articles dealing with any
problem pertaining to collecting, identifying, etc., in short, problems
encountered by our readers. In contrast to other contributions, articles
in this column do not necessarily contain new and original materials.
Questions to the editor, which can be answered in this column, are in-
vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will
attempt to bring reviews of new publications to the attention of our
readers. Also, new timely articles may be listed by title only, if this is
deemed expedient.

Manuscripts should be typed in final form on a high grade white
paper, 812” by 11’, double spaced and accompanied by a carbon copy.

EDITORIAL BOARD

Dr. Donatp P. Assort, Professor of Biology

Hopkins Marine Station of Stanford University

Dr. Jerry DononueE, 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 Datitas Hanna, Curator

Department of Geology

California Academy of Sciences, San Francisco

Dr. Joe, W. Hevcretu, Resident Director

Dr. Leo G. HERTLEIN,
Curator of Invertebrate Paleontology

California Academy of Sciences, San Francisco

Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology

Stanford University, Stanford, California

Dr. Victor LoosanorrF, 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

Mr. Attyn G. Smiru, Associate Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

Dr. Ratpu I. Smitu, Professor of Zoology

Marine Science Laboratory, Oregon State University and Chairman, Department of Zoology

Newport, Oregon

University of California, Berkeley

Dr. Cuarres R. STAseEK, Assistant Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

EDITOR-IN-CHIEF

Dr. RupotF STOHLER, Research Zoologist
University of California, Berkeley

ASSOCIATE EDITOR

Mrs. Jean M. Cate
Los Angeles, California

(75

be THE

A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.

Berkeley, California

VOLUME 10 APRIL 1, 1968 NuMBER 4
CoNnTENTS
Reproduction in Olivella biplicata (Plate 44; 3 Text figures)
DRC RATC HED WARDS 20 r Metcaee ine otter Tsar ct U ct ver eal wll ala heh rive Valet) alia aeOOM

The Functional Morphology of Lyonsia californica Conran, 1837 (Bivalvia)
(7 Text figures)

NVAIRERMINARG EDD cot sii seston omen ieeWran utr). sere (es Sop te io). (ol vel tel ore vaitey i vies GOH
Structure of the Bivalve Rectum. — II. Notes on Cell Types and Innervation
(Plates 45 to 48)
DHomAs C: JEGEA & MIcHAEL J.GREENBERG =. . 2. s: «+ - » «© «© » « «+ 314

Role of Snails’ Disease in the Biological Control of Achatina fulica Bownicu, 1822
on the Andamans

PED) SORIVAS TAVAUS YG) NE ORIVAS TAVAU@ oie ii) ool (etimietur elt 7 (oil el uetuje elon cat te) 20
The Egg Mass and Veligers of Limacina helicina Puipps (2 Text figures)
IVIAD ENE Nee ARANI TAPE Metin: soc ey! soflie el pelt sc ce) atl) ier yo) col te) wr ominlatcnapathelgeripa

Marine Fouling and Boring Organisms in Monterey Harbor
(Plate 49; 3 Text figures)
PO MELADERTAE eet oe ey tah tro ce cell bau a), 6.) o's, Miele tet oel Wiehe wom Mco ten yey

Studies on the Vitality of the Japanese Pearl Oyster, Pteria (Pinctada) martensu
(DunKER) under Abnormal Conditions — I. Oxygen Uptake and Shell
Movement in Sea Water of Low Oxygen Content (5 Text figures)

ME TLUOM NCW AUT ee ne yarrow eres ale leas, | ore einen ien seoinel set 4e
The Date of Publication of K1ieNER’s Mitra Monograph in the “Spécies général et
iconographie des coquilles vivantes.”
WsTttERe @MIVEROERNOHORS Kiva | orer- ca leged eu nol eine eon lel 6) =) pe 2349

[Continued on Inside Front Cover]

Distributed free to Members of the California Malacozoological Society Inc.
Subscriptions (by Volume only) payable in advance to Calif. Malacozoological Soc., Inc.
Volume 11: $18.- Domestic; $18.80 in the Americas; $19.20 in all other Foreign Countries

Single copies this issue $10.-. Postage extra.
Send subscriptions to Mrs. Jean M. Cate, Manager, 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

ContTENTts — Continued

Itinerary of the Voyage of H. M.S. Blossom, 1825 to 1828
Josepm RoSEWATER.\(.00.5< Ase; ss cae ee eee
The Ovulidae, Pediculariidae and Triviidae of Fiji (Mollusca: Gastropoda)
(Plates 50 to 52; 5 Text figures; 1 Map)
Watrer Oniver: CERNOHORSKY, |: 7. hcg eee ene

Aplysia vaccaria, a New Host for the Pinnotherid Crab Opisthopus transversus
(2Text figures)
ANTHON ‘GRAIG: BEONDE |. (Sie segs eo Hae ee cece ets

Taxonomic Placement of Coralliophila incompta Berry, 1960, With the Proposal of

a New Genus, Attiliosa (Gastropoda: Muricacea) (Plate 53; 5 Text
figures)
WIcrianm K. EMERSON: = 2 95900515 seco) eee on ota ects ato eS
A New Cowrie Species from the Philippines. (Plate 54)

CRAWFORD NEILL CaTE & FRANZ ALFRED SCHILDER . .. . « « « « « «

Studies on the Mytilus edulis Community in Alamitos Bay, California: - III. The
Effects of Reduced Dissolved Oxygen and Chlorinity Concentrations on
Survival and Byssal Thread Production (2 Text figures)

Dona.p J. ReisH & JosepH L. Ayers, Jr. .
West American Mollusk Types at the British Museum (Natural History), IV.
CarPENTER’s Mazatlan Collection (Plates 55 to 59; 171 Text figures)
A. Myra KEEN
Effects of Feeding by Armina californica on the Bioluminescence of Renilla koellikeri
Hans BertscH
A New Marine Mollusk from Mozambique in the Genus Festilyria Pitspry & OLSSON,
1954 (Gastropoda : Volutidae) (Plate 60; 1 Map)
CuiFToN STOKES WEAVER .
MEDHODS & TECHNIQUES 35) 25) a

Boiled Lettuce and Cress as Diet Supplements for Certain Species of
Mollusks T. W. FisHER

NOTES & NEWS
BOOKS, PERIODICALS & PAMPHLETS

. 350

- 353

- 375

- 379

. 382

- 384

- 389

. 440

- 442
. 446

- 447
- 449

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. 10; No. 4

THE VELIGER

Page 297

Reproduction in Olivella biplicata

BY

D. CRAIG EDWARDS

Department of Zoology, University of Chicago

Department of Oceanography, Scripps Institution of Oceanography '

(Plate 44; 3 Text figures)

IN CONJUNCTION with more general studies of Olivella
biplicata (SoweERBy, 1825) considerable information has
been obtained on its reproduction. These observations
cover aspects of sexuality, the courtship and mating
behavior, including documentation of a nonrandom
mating pattern, evidence of year-round mating and settle-
ment, a fecundity count, and the first description of the
egg capsule, larva, and form of early development.
Field work was done at various sites on the Oregon and
California coasts; laboratory investigations were done at
the Oregon Institute of Marine Biology, Charleston,
Oregon, and at Scripps Institution of Oceanography,
La Jolla, California.

GONOCHORISM

Olivella biplicata is dioecious with internal fertilization,
but sexual dimorphism is not conspicuous. Sex is most
easily determined from the positions of individuals in
courting pairs. Alternatively presence or absence of a
penis, shape of the ventral pedal gland, gonad color, or
form of the genital tracts may be used. Although presence
of a penis is decisive, absence may indicate either a
female or a male that has lost sex characters owing to
trematode infection, in which case a small, easily over-
looked rudiment may remain. Females possess an incon-
spicuous, cup-shaped pedal gland for molding and
attaching egg capsules, whereas males have a_ longi-
tudinal, probably glandular (see later) slit at this site.
The mature testis is orangeish, the ovary yellowish, but
the difference is slight and is lacking in immature or
parasitized animals. The anatomy of the genital tracts is
given in Marcus & Marcus (1959a) and Ze (1955).

Unlike most prosobranchs (Comrort, 1957; FRETTER
& GRAHAM, 1964; Gattsorr, 1961; AspoTtt, 1954;
Ropertson, 1959), male Olivella biplicata grow faster

' Present address: Department of Zoology, University of Massa-
chusetts, Amherst, Massachusetts 01002.

(FRANK & Epwarps, unpubl. data) and are larger than
females. (Size data hercinafter are shell lengths from
the siphonal canal to the apex.) The sexual size dimorph-
ism was demonstrated by measuring and sexing the
mature snails taken in total-count transect sampling across
Olivella-occupied beaches (‘lable 1). In addition, in
experiments regarding size as a treatment, randomly se-
lected “Large” snails (22.0-25.0mm) were nearly all
males, whereas most of the “Medium” snails (17.0 to
20.0mm) were females. Finally, males taken in courting
pairs were significantly larger than their female partners
(Table 2; Figures 1 and 2). Thus shell size and the
spacing of growth lines provide weak sex indicators:
lines are close together on slow-growing female shells,
which rarely exceed 23mm, but further apart on male
shells, which are often large (up to 30mm).

Table 1

Mean lengths (X), standard deviations (SD), and numbers (N) of
mature snails of cach sex taken in transect sampling at two beach
sites on the Oregon coast in 1964.

Site as Males oe Females 1
X(mm) SD N X(mm) SD N (t-test)
Yaquina Bay 19.9 la? 7 NGO), 4} OO!
Coos Bay 19.84 267 52 Ns shal 8 OO

Both Stouter (1959-1960) and I (several sets of
measurements) find that, in contrast to some gastropods
(FRETTER & Grattam, 1964), female Olizella biplicata
shells are no more tumid than male ones.

Among mollusks generally, and including Oliella
verreauxtt (Ductos, 1857) (Marcus & Marcus, 1959a
and b), females tend to be more numerous than males.
As the disparity is lacking in young animals and increases
with age, it is attributed to carlicr death in males (FRET-
TER & GRAHAM, 1904). Comrort (1957) also reports

Page 298

THE VELIGER

Vol. 10; No. 4

greater longevity for females. These findings could, how-
ever, be due to mistaken sex identifications with para-
sitized animals, wrongly equating age with size, or samp-
ling bias favoring capture of larger individuals. Sex ratios
for O. biplicata can be read from the N columns in Table
1. The ratio was approximately 1:1 for both whole
populations and by beach levels within them. Of the 52
males (vs. 58 females) taken at Coos Bay less than half
(23) had a functional penis -— often it was absent.
For the effective sex ratio to be unity, the probability of
sexual impairment from parasitic infection would have
to be the same for both sexes. At Yaquina Bay, where
design forbade dissections, 67 males, 45 females, and 16
“unknowns” were taken. Most “unknowns” would be
females, since their sex characters are hard to detect and
positive identifications were required.

Olivella biplicata of both sexes mature sexually at
about the 16mm size: these are the smallest snails taken
in numbers in courting pairs (Figures 1 and 2). Since
16mm probably represents about 1 year’s growth and O.
biplicata’s longevity likely exceeds 10 years (FRANK &
Epwarps, unpubl. data; StoHLER, 1962, and personal
communication), about 10% or less of the life span is
passed in immaturity. Prosobranchs generally exhibit no
genetically set post-reproductive period, though heavy
trematode infections can have the same effect.

MATING BEHAVIOR

Courtship in Olivella biplicata is initiated by the male’s
extending his propodium and grasping the female’s shell
at or near its apex. A tandem, courting pair results, with
the female in front, the male behind (Plate 44). While
joined, the pair alternately moves about and sits quietly;
the female may feed. A characteristic activity of courting
snails is a forward-and-back rocking of the body on the
large foot. This motion is more common in females, who
initiate it, than in males, but at times the partners rock
together in unison. Whether this activity is a necessary
prelude to copulation is not known, but ZeLL (1955)
believes it is one step in a reciprocal reaction chain like
those demonstrated in the courting sequences of many
animals (TINBERGEN, 1953).

Although Marcus & Marcus (1959b) found no
evidence for it in Olivella verreauxu, the bond between
courting O. biplicata is a sticky mucus, strands of which
adhere to the shell apices of females from pairs. This ad-
hesive is apparently secreted by the male’s pedal gland,
which may, in courting males, be filled or covered with
thick mucus. This connective can support the weight of
the male when the female is lifted from the substrate,

and the coupling ismaintained whena pair isrolled about
by beach surf. In a shifting medium like sand an attach-
ment between courting individuals likely helps ensure
fertilization (see PEARSE et al., 1942).

Pairings may persist for a long time: one couple was
joined continuously for at least 31 hours, and another
paired repeatedly for 3 days. During courtship the male
extends his penis along the right side of the female
shell and, when successful (many attempts fail; males
are very sensitive to disturbances at this vulnerable
stage), into her mantle cavity. Males have sometimes
advanced on the right side of the females’ shells by
this stage. During intromission, which on several occa-
sions lasted about 10 minutes, the two snails are firmly
locked together. Copulation may occur repeatedly be-
tween a single pair, or individuals may mate with several
partners in a few days time.

Mate-finding in Olivella biplicata seems to be based
on males’ reactions to short-lived chemicals left in female
mucus trails, contact chemosensory responses, and pos-
sibly tactile cues, but not on distance chemoreception.
Although these snails often travel for a distance in each
others’ sand trails (probably a tactile response), males
are unable to use direct trail following to locate distant
females. But a male coming upon a very fresh track of
a sexually-ready female (usually less than 15cm away)
turns into the track, accelerates, makes contact, and
attempts to pair. Evidence for a chemical stimulus in
female mucus trails is furnished by occasional obser-
vations of triplets consisting of a regular courting pair
with a second male behind the first, all linked in tan-
dem. No pairings of males alone were observed. (One
new male bit the courting male, who turned aside, the
former taking the female!) Males who paired repeated-
ly, however, showed surprisingly little ability to relocate
their partner when pairing was interrupted: although
lying just behind the female, they often moved off in
an inappropriate direction, sometimes attempting un-
successfully to attach to another nearby snail. Once
(San Diego Flood Control Channel; 3 August 1966) a
male Olivella baetica CARPENTER, 1864, was found
courting a female O. biplicata, suggesting that initially
sex may be more readily discriminated than species.
During courtship, males often curl the lateral edges of
their propodia anteriorly under the females, contacting
the latter’s metapodia; females sometimes extend their
proboscides to their shell apices and the male propodia
and occasionally bite the latter, terminating courtship.
Females may also curl the posterior edge of the meta-
podium dorsally, preventing male coupling - though
one male bit this barrier and later formed a pair. These

Vol. 10; No. 4

contacts would permit contact chemosensory responses
and further discrimination of partners.

Distance chemoreception in mate-finding was tested
on a large (91cm by 290cm) outdoor water table fitted
with a median longitudinal divider (115cm) at the
water inlet end. On one side sea water passed through
an enclosure of females from pairs and on the other
through an empty enclosure. Males from pairs were
released at the downstream end of the table, but showed
no tendency to move preferentially toward the females.
The high densities in natural Olivella biplicata popula-
tions and the snails’ considerable mobility should pro-
duce frequent contacts between sexually ready individuals
without special means of mate-finding at a distance
being required.

On the gently sloping, protected sand beaches where
Olivella biplicata usually lives, this snail develops a size
class distribution with large animals higher on the shore,
smaller lower (Epwarps, 1965). The resulting graded
separation of reproductives and immatures should fur-
ther aid mate-finding efficiency.

Measurements of individuals in courting pairs showed
that 1) males were significantly larger than their female
partners, as expected from the sex difference in growth
rate (Table 2 and see above) and 2) mating is non-
random, larger males pairing with larger females, smaller
males with smaller females (Figures 1 and 2). The best
evidence for the mating pattern is given in Figure 1,
since at the Charleston lagoon the lengths of 210 nearest

THE VELIGER

Page 299

neighbors of 97 pairs were found to be independent of
the courting females’ sizes: the regression slope b—=(.067
was not statistically significant (data furnished by Peter
W. Frank). Apparently lagoon channel shifts and cur-
rents do not permit a size class distribution to be
developed at this site. Inspection of Figure 1 suggests
large males mate less selectively than small males, which
court only smaller females. Figure 2, on the other hand,
presents data from sites where segregation of Olivella
biplicata by size category has been demonstrated (Yaqui-
na Bay, Coos Bay, and Duxbury Reef [Epwarps, 1965})
or indicated (Monterey Harbor [RryNo.ps, 1948] and
San Diego Flood Control Channel [F Wolfson, cited in
STOHLER, 1959-1960}). Although the relationship be-
tween male and female sizes is not statistically significant
for any of these small samples, the trend is evident: the
regression line for the grouped data (N=82) is
Y= 13-223 0444 OX

slope b statistically significant at the 0.01 level. The
steeper slope for these samples, compared to the Charles-
ton lagoon value (b==0.350), may be due to the size
class distribution. Also at Duxbury Reef, where only
larger animals were taken in pairs (Figure 2), the
nonselectivity of large males is clear (nonsignificant
slope b==-0.343).

Although differences in activity cycles — large snails
are active at night, whereas small ones may be active in
the light (Epwarps, 1965) -— could account for non-
random mating, the nonselectivity of large males sug-

Table 2

Differences in lengths of males and females taken in courting pairs
at six sites on the Oregon and California coasts.

Pairs Pairs with Mean Difference
Site Date (N) oS> ee 3o-? (mm) SD p*
Yaquina Bay June-July
(Tide pool stream) 1964 18 17 2.88 2.53 0.001
Coos Bay July-Aug.
(Charleston lagoon) 1963 197 150 1.85 2.53 <0.001
Coos Bay June-July
(Beach) 1964 20 15 1.94 2.77 0.01
Duxbury Reef® Oct.-Jan.
1964-65 22 18 2.51 2.43 0.001
Monterey Harbor® Nov.
1964 6 3 0.22 1.96 0.80
San Diego Flood Oct.-June
Control Channel 1965-66 16 14 1.91 2.31 0.01

4 tested by paired comparison t-test
® STOHLER (1959, 1960) found females larger than males in all of
11 pairs taken here. No explanation for the disagreement is a-

vailable except for sampling error
© no large snails were taken in pairs here (cf. Text figure 2), pos-
sibly owing to trematode infections

Page 300

THE VELIGER

Vol. 10; No. 4

gests another explanation. Since in mating a male must
extend his penis most of the length of the female’s shell
and into the mantle cavity, probably small males simply

Length of Male (mm)

15 16 17 18 19 20 21 22 23 24 25

Length of Female (mm)

Figure 1

The relationship between lengths of males and females taken in
197 courting pairs at the tidal lagoon, Charleston, Oregon, July to
August 1963. Single pairs are indicated by a solid circle, two pairs
by a circle with a cross through it. The regression line, fitted by the
least squares method, is Y= 14.815 +0.350 X. Slope b is statistically
significant at the 0.001 level. (Data collected by Peter W. Frank)

cannot accommodate larger females. A suggestion that
small males are at a disadvantage in courting comes from a
comparison between mean sizes of mature animals taken
in total count sampling and those from courting pairs:
on the Coos Bay beach, the only site for which such data
are available, courting males were somewhat larger than
mature males over-all (20.78mm vs. 19.84mm; P=
0.10), whereas female sizes were very similar (18.84mm
vs. 18.45mm; P=—0.50). Another test would be a com-
parison of the mating success of large females isolated
with large vs. small males. If small males are inferior
mating partners, the size class distribution of Olivella
biplicata would promote efficient pairing both by sepa-
rating mature snails from immatures and by placing
large matures with large, small with small. The more
rapid growth of males may also be related to their

mating effectiveness. Since pairing data came from
courting snails, many of whom had likely not yet copu-
lated, apparently either or both sexes can recognize and
select, probably by chemosensory means (see above),
partners of suitable sizes before actual mating attempts.

YEAR-ROUND MATING
AND SETTLEMENT

Three lines of evidence indicate that Olivella biplicata
both breeds and spawns all year. (1) SToHLER (1959-
1960) and I have both observed courting pairs in every
month. Numbers mating at a given time seem to be due
more to as yet undetermined local, short-term conditions
than to annual changes: large numbers mate in every
season, though on successive spring tide series pairs may
be abundant or scarce. (A complicating factor is that
pairs are easier to find in dim light and quiet water.)
(2) The smallest, youngest snails taken by sieving (4 to

26
24
22
20
18
16

Length of Male (mm)

15 17 19 QI
Length of Female (mm)

23 2515 17 19 21 23 25

Figure 2

The relationship between lengths of males and females taken in
courting pairs at five sites on the Oregon and California coasts.
(A) Yaquina Bay (44° 37’ N Latitude)

(B) Coos Bay, beach (43° 21’ N Latitude)

(C) Duxbury Reef (37° 54’ N Latitude)

(D) Monterey Harbor (36° 36’ N Latitude)

(E) San Diego Flood Control Channel (32° 45’ N Latitude)
Sampling dates are given in Table 2

Tue VE.IcER, Vol. 10, No. 4 [Epwarps] Plate 44

+5 4 4
ols

tin ;
* i “ee
(or J « %

A courting pair of Olivella biplicata with the female in front
and the male following

photograph by Joun W. Evans

Wol lO Now

THE VELIGER

Page 301

5mm) entered the Coos Bay beach population through-
out the year and occurred at every locality on every
sampling date. (3) Size frequency distributions for
populations showed no breaks that would indicate year
classes. Because small animals do not cease growing in
winter as mature snails do (Epwarps, unpubl.), the lack

Figure 3

(A) The egg capsule of Olivella biplicata; three quarters view from
above. (B) The same; diagrammatic cross section. (C) The veliger
larva of Oliwella biplicata shortly after hatching. These sketches of
semi-transparent materials are not accurate in details.
(Drawings by Susan Ahrend)

of year class modes cannot be accounted for on the basis
of mating and spawning only occurring during periods
of growth for all snails. Whether individuals, in contrast
to populations, exhibit reproductive cycles is not known.

Using the occurrence of mating pairs or of very small
animals at different times and places, GirrorpD & GirrorD
(1942, 1944, 1948) proposed a complicated series of
mating seasons for Olivella biplicata. Their data equally
well support year-round reproduction.

Year-round reproduction occurs in a number of ma-
rine gastropods (Moore, 1938; RickeTrs & CaLvin,
1962; THorson, 1950). In Olivella biplicata continuous
breeding may be related to the genus’ neotropical origins,
there being some evidence that tropical forms enjoy
longer breeding seasons than temperate ones (THorson,
1950). However, O. mutica (Say, 1822) in Florida ap-
parently spawns for only about one month in spring
(Paring, 1962), and O. fulgurata’s (A. ADAMS & REEVE)
spawning season in Japan is May to July (Hane, 1960).

Unless having new young enter the population all year
is crucial for Olivella biplicata’s success, the factor
thought to limit northward ranges of species, and hence
faunal provinces, viz. the continuous period that sea
temperatures meet requirements for reproduction and
early growth (Hatt, 1964), does not seem applicable
here. Yet O. biplicata’s species range - Magdalena
Bay, Baja California, Mexico (25° N Lat.) to lower
Vancouver Island (49° N Lat.) (KEEN, 1937) — close-
ly fits the limits of the Oregonian plus the Californian
shallow water, marine faunal provinces (KEEN, 1958;
HA, 1964) ”.

EGG CAPSULE, FECUNDITY
AND TYPE or EARLY DEVELOPMENT

Olivella biplicata’s egg cases and form of development
were found by collecting courting females in the San
Diego Flood Control Channel and maintaining them at
Scripps Institution of Oceanography in aquaria provided
with running sea water (14° to 15° C), a sand substrate,
and possible ovipositing sites. Egg capsules were subse-
quently found on various bivalve shells, empty O. bipli-
cata shells, glass slides, and glass and plastic dishes, but
not on the mother shells as reported for Japanese species
(Haze, 1960; Hortkosut, unpubl.). As in O. mutica,
O. pusilla and O. verreauxii, empty shells are likely
the substrate used by O. biplicata in nature. Capsules
are attached separately and preferentially in depressions
or grooves (c. g. on Donax) ; their dispersion pattern on

21. McT. Cowan (1964, personal communication) has, however,
found empty shells of Olivella biplicata as far north as Queen
Charlotte Island.

Page 302

shells is markedly contagious, and on a smooth surface
several may be placed contiguously.

The egg capsule (Figures 3a and 3b) consists of a
dome-shaped cap with an indistinct median suture (not
shown in the Figures) above a broader, slightly irregular
base. Capsules are small (0.5mm basal diameter; 0.3mm
lumen diameter), transparent, and unsculptured. A fis-
sion line borders the convex cap, which often comes free
of the base at hatching. The capsule appears very similar
to that figured for Olivella fulgurata from Japan (Amio,
1963), but differs slightly from that of O. verreauxu
from Brazil (Marcus « Marcus, 1959b), whose cap
bears sculptured ridges, and considerably from those of
O. mutica (PAINE, 1962) and O. pusilla (Marrat, 1871)
(PERRY & SCHWENGEL, 1955) from Florida, whose caps
are more apical with a ridged border.

Although early development was not studied, Olivella
biplicata would be good material for such work. Each
transparent egg capsule contains one clearly visible white
egg or embryo without “nurse eggs.” Cleavage and lar-
val stages are distinctly observable. The snails will ovi-
posit on glass slides, and the time of spawning can be
known precisely by exposing clean slides tomated females
for short time intervals. Finally, year-round reproduction
means eggs and larvae would be available in large
numbers all year.

Numbers for eggs laid incapsulesare generally modest
compared to those broadcast into the sea. Olivella further
provides for its young by placing them one to a capsule.
To determine fecundity, courting females from the field
were placed in separate clear plastic dishes provided
with screen tops and running sea water (14° to 15°C).
The containers were periodically examined for capsules,
the females being transferred to clean dishes. One effort
was successful. By total-count a 20.6mm female depos-
ited 4236 egg capsules in 46.8 days (June 22 to August
8, 1966) ; mean per day rate was 90.5 (range 60.6 - 171.4
for 3 censuses). This is a minimum estimate, since
spawning may have ceased due to depletion of the
gonad, decline in the female’s vigor under artificial con-
ditions (occasionally algal food was supplied), or lack
of opportunity for repeated mating. The female may
also have mated and spawned before capture. This count
lies among previously reported values for capsule-spawn-
ing snails (PELSENEER, 1935). Size specific fecundity
data would be valuable, as they might indicate the
adaptive value of large size, slow growth, and long life
in O. biplicata (cf. McLaren, 1966).

Development time in the capsule was highly variable.
Many young hatched by day 10, but others were still
alive in their capsules after 28 days. All capsules were
kept in 14° to 15° C circulating sea water, but the late-

THE VELIGER

Vol. 10; No. 4

hatching animals were subjected to occasional warming
during microscope observations.

The young emerge as veligers (Figure 3c). The shell
is nearly transparent and unsculptured; it is a barely
skewed helix of ca. 14 whorls and a maximum diameter
of 0.2-0.3mm. The large, bilobate velum has a light
brown band of pigment near its borders and bears eye
spots on slight prominences anteriorly. The body is
nearly colorless, and internal organs, especially the vis-
ibly beating heart, are easily seen. The operculum is clear
and scalelike.

The veligers are apparently nonpelagic. Rather they
swim, anterior (eye spots) end down, at the bottom of their
containers, opening and closing their velum lobes against
the substrate. Occasionally they swim in a tight vertical
loop, but return quickly to the bottom. The larvae appear
to be examining the substrate. When clean sand was
added to a dish, the animal opened and closed its velum
around several grains, then swam on. Possibly these lar-
vae never join the plankton; but, perhaps responding
negatively to light and positively to gravity (several tests
proved inconclusive), swim at the bottom until suitable
cues for metamorphosis — presumably ones associated
with lower shore situations — are detected. Balanced
sex ratios across beach levels show females do not move
down shore to spawn. The larval behavior would explain
how the young, hatching higher on the shore where
adults live, are able to cross wide expanses of beach and
reach the lower shore levels where physiological stresses
are within their ranges of tolerance (Epwarps, 1965).

No larvae metamorphosed - or developed appre-
ciably -— when maintained in bare glass or plastic
dishes, or in ones provided with sand. Isolated individ-
uals died after 3-4 days. Presumably stimuli required
for metamorphosis were missing from the laboratory set-
up. Aggregations of very small snails occur on the shore,
suggesting either that settlement is stimulated by irreg-
ular features of the environment or presence of settled
young, or that metamorphosed young show strong posi-
tive responses to one another.

Since close relatives of Olivella biplicata from warmer
water exhibit either direct development — O. fulgurata
(Amio, 1963), O. spreta Goutp, 1860 (Horikosui, un-
publ.), O. mutica (Paine, 1962), and Ancilla sp. (NaTa-
RAJAN, 1957) — ora short, nonpelagic larval life — O.
verreauxtt (Marcus & Marcus, 1959b) —- O. bipli-
cata would have been expected to show direct develop-
ment in cool West Coast waters (THorson, 1950). Long
breeding seasons are also often linked to direct develop-
ment (THoRSON, op. cit.). Yet the mixed occurrence,
even within genera, of species with free-swimming veli-
gers and others with direct development is well-known

Vol. 10; No. 4 THE VELIGER Page 303
among Prosobranchs (THorson, op. cit.). A larval stage Hatt, Crarence A., Jr.

in the life history of O. biplicata appears to be an adap- 1964. Shallow-water marine climates and molluscan provinces.
tation to the particular ecology of this species; it provides Ecology 45: 226 - 234

young an opportunity to attain suitable beach levels in Keen, A. Myra

a species exhibiting a marked segregation by size class.

ACKNOWLEDGMENTS

I am deeply indebted to Dr. Peter W. Frank for kindly
making available the data on nonrandom mating at the
Charleston lagoon, for providing facilities for part of
this work, and for many valuable discussions. I am also
grateful to Dr. Thomas Park, Dr. Rudolf Stohler, and
Dr. Gunnar Thorson for advice and encouragement and
to Dr. E. W. Fager for providing facilities. This work was
supported by National Science Foundation Predoctoral
and Postdoctoral Fellowships. Dr. Frank’s contribution
was supported by National Science Foundation grant

GB 977.

LITERATURE CITED

AssotT, RoBERT TUCKER

1954. American seashells. Princeton, New Jersey. D. van
Nostrand Co., Inc.; xiv+541 pp.; 100 figs.; 40 plts.
Amio, M.

1963. A comparative embryology of marine gastropods, with
ecological considerations. Journ. Shimonoseki Univ. of
Fisheries 12 (2, 3): 229 - 358

Comrort, ALEXANDER

1957. The duration of life in molluscs.

London 32: 219 - 241

Epwarps, Daas Craic
1965. Distribution patterns within natural populations of
Olivella biplicata, their underlying behavioral mechanisms, and
their ecological significance. Univ. of Chicago Ph. D.
Thesis. And: Bull. Ecol. Soc. Amer. 46: 168

FRETTER, VERA, & ALASTAIR GRAHAM
1964. Reproduction. In: Physiology of Mollusca, ed. K. M.
Wizpur « C. M. Yonce, Vol. 1. Acad. Press, New York, pp. 127
to 164

GattsorF, PauL SIMon

1961. Physiology of reproduction in molluscs.
1: 273 - 289

GirrorD, DELILA SaRA GIFFEN & Epwarp WINSLOW GIFFORD
1942. Color variation in Olivella biplicata in various localities.
The Nautilus 56 (2): 43 - 48

1944. Californian Olivellas.
1948. Oregonian Ollivellas.

Hase, TADASHIGE
1960. Egg masses and egg capsules of some Japanese marine
prosobranchiate gastropods. Bull. Mar. Biol. Sta. Asamushi

10: 121 - 126

Proc. Malacol. Soc.

Amer. Zool.

The Nautilus 57 (3): 73 - 80.
The Nautilus 62 (2): 51 - 54.

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)

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)

Marcus, EvaLine & Ernst Marcus

1959a. Studies on Olividae. Univ. Sao Paulo Fac. Filos.
Cienc. 'e Letras. Bol. Zool. 22: 99 - 188

1959b. On the reproduction of Olivella. Univ. Sao Paulo
Fac. Filos. Cienc. e Letras. Bol. Zool. 22: 189 - 199

McLaren, IAN A.

1966. Adaptive significance of large size and long life of the
chaetognath Sagitta elegans in the Arctic.
852 - 855

Moore, Hinary B.

1938. The biology of Purpura lapillus. Part II. Growth. Part
III. Life history and relation to environmental factors.
Journ. Mar. Biol. Assoc. U.K. 23: 57 - 74

NatTaraJAN, A. V.

Ecology 47:

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

PaInE, RoBErT TREAT
1962. Reproduction of Olivella mutica.
75 (4) :139 - 142; 1 text fig.

Pearse, A.S., Harotp J. HumMm « Georcr W. WHARTON
1942. Ecology of sand beaches at Beaufort, N. C. Ecol.
Monogr. 12: 135 - 190

The Nautilus

PELSENEER, PAUL
1935. Essai d’éthologie zoologique d’aprés létude des mol-
lusques. Acad. Roy. Belg. Classe sci. publ. Fondat. Agathon
de Potter 1: 1 - 662

Perry, Louise M. & JEANNE SANDERSON SCHWENGEL
1955. Marine shells of the western coast of Florida.
Res. Inst., Ithaca, N. Y.

Paleont.

REYNOLDs, Harotp C,
1948. Notes on the feeding and food habits of the gastropod
Olivella biplicata (SowERBy) at Monterey Harbor. Unpubl.
Student rept. Invert. Zool., Univ. Calif:, Berkeley

Ricketts, Epwarp I. & Jack CALVIN
1962. Between Pacific tides. 3rd. ed., 2nd. rev. by Jorn W.
HepopetH. xili+516 pp.; 135 text figs.; 46 plts.; Stanford
Univ. Press, Stanford, California.
ROBERTSON, ROBERT
1959. Observations on the spawn and veligers of conchs
(Strombus) in the Bahamas. Proc. Malacol. Soc. London
33: 164-171

Page 304

THE VELIGER

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 Vcliger 4 (3): 150-151; plt. 36 (1 Jan. 1961)

Tuorson, GUNNAR
1950. Reproductive and larval ecology of marine bottom in-

vertebrates. Biol. Rev. 25 (1): 1-45.
TINBERGEN, NIKO
1953. Social behaviour in animals. Methuen, London

ZELL, CLARACE PLUMB Bock
1955. |The morphology and general histology of the repro-
ductive system of Olivella biplicata (SowERBY), with a brief
description of mating behavior. Univ. Calif., Berkeley,
M.A. Thesis

Vol. 10; No. 4

Vol. 10; No. 4

THE VELIGER

Page 305

The Functional Morphology of Lyonsia californica Conrap, 1837

(Bivalvia )

WALTER NARCHI

Faculdade de Filosofia, Ciéncias e Letras

Universidade de Sao Paulo, Sao Paulo, Brasil

(7 Text figures)

INTRODUCTION

THE FAMILY LyoNsmpAE comprises 3 genera: Lyonsia,
Mytilimeria and Entodesma (AxsorttT, 1954, p. 468). The
last is considered by THIELE (1963, p. 936) as a subgenus
of Lyonsia. Species of Lyonsia are widely distributed in
the northern hemisphere including the west coast of
Europe and the Atlantic and Pacific coasts of North
America.

Lyonsia californica Conran, 1837, occurs on the Pacific
coast of North America from Puget Sound to Lower Cali-
fornia (ABBoTT, 1954, p. 468). According to KEEN (1937,
p. 21) it ranges from 24° to 56° North Latitude. Isolated
references to this species were found in the papers of
Arxins (1937), Kettocc (1915) and YoncE (1952),
but there is no special research dealing with all aspects
of the functional morphology of this species.

This paper is concerned with observations on the
structure, ciliary currents of feeding and digestion, and
other functional adaptations of Lyonsia californica. The
animals were collected at Lawson’s Flat and White Gulch
in Tomales Bay, Marin County, California; the observa-
tions on live specimens were made at Pacific Marine
Station, Dillon Beach, California.

SHELL

The shell is about 3 times as long as it is broad and
moderately obese in the anterior part where the shell
valves are rounded, while the posterior end is laterally
compressed. It is thin, fragile and almost transparent.
The umbo is situated approximately one-third from the
anterior end; the beak area is swollen. The shell is made
largely of nacreous material and fractures easily. BocciLp
(1930, p. 291) noted that the shell valves consisted of a
nacreous substance between two layers of fine prismatic
structure.

The valves are covered by a thick layer of periostra-
cum which covers the margin of the shell; weak, radial,
dark lines run through the periostracum. Dorsally the two
shell valves are joined along their entire length by peri-
ostracum; it covers the base of the siphons and the ventral
surface of the fused mantle edges. This feature is con-
sidered as primitive by YoncE (1952, p. 446). In the
living animal sand grains are found attached everywhere
on the shell but adhere more firmly at the base of the
siphons and to the ventral periostracum as in Entodesma
saxicola Bairp, 1863 (YoncE, 1952, p. 440). The dense
accumulation of sand grains around the base of the
siphons fills the posterior gape when the siphons are
withdrawn.

There is a lithodesma under the hinge attached to the
ligament which is very similar to that of Entodesma saxi-
cola (YONGE, 1952, p. 446).

The margins of both valves are poorly calcified in the
siphonal region as in Pandora inaequivalvis (LINNAEUS,
1758) (ALLEN, 1954, p. 474). The valves are slightly
inaequivalve. When closed, the periostracum of the free
margin of the left valve folds over and fits against the
outer marginal region of the right valve.

The shell of the largest animal encountercd in this
study measured 38mm in length, 16mm in height and
12mm in width.

MANTLE

The mantle edges and periostracal grooves are fused
ventrally (Figure 1, mef). There is a pedal gape (pg)
and a smaller pallial aperture (a:), which opens about
5mm from the base of the inhalant siphon (in). The
fourth aperture in Lutraria and Thracia lics closer to the
inhalant siphon than in Lyonsia californica and in Ento-
desma saxicola (ATKINS, 1937, p. 444). According to
KELLoce’s figures of E. saxicola and Mytilimeria nuttallii

Page 306

Conrap, 1837 (1915, p. 654; figs. 22, 26, 27, 28) the
fourth aperture of the mantle is just posterior to the
region in which the rejecta brought by the mantle cur-
rents collect. While KeLLoce (op. cit., p. 656) did not
note the significance of this opening, YoNcE (1952, p.
447) thought that it was a purely structural feature,
possibly a result of the manner in which the mantle
edges fuse during development.

Figure 1

Lyonsia californica CONRAD
ventral aspect of the animal, showing the pedal gape and foot with
byssal groove and the fourth aperture
a, — 4» aperture bf — byssus fold f — foot
in — inhalant siphon mef — mantle edges fused
pg — pedal gape

On both sides of the fourth aperture fusion is complete.
The muscles cross the ventral region from one lobe to the
other. In Ensvs siliqua and E. arcuatus actual tissue fusion
occurs only posteriorly to the fourth aperture (ATKINS,
1937, p. 431). Thracia villosiuscula has a fourth aper-
ture near the posterior end of the animal just below the
inhalant siphon and exhibits a true tissue fusion (ATKINS,
op. cit., p. 431).

SIPHONS

The siphons are transparent, separated and short (Figure
2). The basal region is covered with periostracum in
which sand grains are incorporated. The cavity of the
basal region represents a posterior extension of the
mantle cavity (YoNncE, 1952, p. 441). The base of each
siphon is surrounded with many small tentacles. The
aperture of the inhalant siphon is fringed with 9 simple
tentacles. The exhalant siphon consists of a tapering
tubular membrane without any tentacles between the
aperture and the common outer ring. This arrangement
is similar to that in Lyonsia hyalina as described by
Morse (1919, p. 161).

Regularly distributed black spots are found on both
siphons; on the exhalant siphon the spots are concen-

THE VELIGER

Vol. 10; No. 4

trated in the lower part, forming a dark ring. Many spots
are present on the tentacles of the inhalant siphon.

As in Pandora, the siphons are not capable of great
extension, and for this reason the animal is morc or less

0, wont ye

“Apis Sly
SA Gea)

Figure 2
Appearance of fully extended siphons
projecting out of the shell valves

limited to the epifaunal regions of the substrate (Boss «
Me_rRILL, 1965, p. 183).

As in Glossus humanus (LINNAEUS, 1758) specimens
placed in a dish withdrew their siphons when a hand was
placed lightly on the table on which the jar stood; this
sensitivity is evidence that the animal normally inhabits
quiet waters (OweEN, 1953, p. 92). The exhalant siphon
contracts by lateral constriction of its sides forming a
figure 8 when seen from above; this is followed by the
infolding of the siphonal edges.

THE MANTLE CAVITY

The disposition of the organs in the mantle cavity (Figure
3) is revealed after removal of the left shell valve and
mantle lobe. The slightly smaller anterior adductor muscle
(aam) and the larger posterior adductor muscle (pam)
are situated in a line parallel with the dorsal margin.
The important features of the different organs will be
outlined below.

THE CTENIDIA

The outer demibranch consists only of a direct lamella,
which is upturned and attached near the dorsal surface
(Figure 3). The inner demibranch is broad and reflected.
Ripewoop (1903, p. 150), Ketioce (1915, p. 656) and
Atkins (1937, p. 351) noted that in allied species the
same phenomenon occurs. The lamellae are deeply pli-

Vol. 10; No. 4

THE VELIGER Page 307

Figure 3

Figure 3

Lyonsia californica CONRAD
mantle cavity viewed from left side after removal of the left shell
valve and mantle lobe

a, — 4 aperture aam — anterior adductor muscle

ap — anal papilla co — lithodesma
ex — exhalant siphon f — foot

d — digestive diverticula
id — inner demibranch

ilp — inner labial palp in — inhalant siphon k — kidney

Figure 4

The labial palps of the left side. Shell length 19 mm

aam — anterior adductor muscle
ilp — inner labial palp

id — inner demibranch
olp — outer labial palp

1 — ligament olp — outer labial palp Ov — ovary
pam — posterior adductor muscle pe — pericardium
pg — pedal gape pr — periostracum r — rectum
prm — posterior retractor muscle t — testis v — ventricle

cated and heterorhabdic. As in Lyonsia norwegica, 13 to
17 filaments form a plica, while the apical 5 are more
strongly developed than the others. As with the remaining
filaments, the apical filaments have a middle tract of
long, coarse frontal cilia, bordered on each filament by
tracts of fine, short cilia.

At the free edge of the inner demibranch there is a
deep marginal groove (Figure 4). Ciliary currents were
studied by the application of carmine, aquadag, and fine
carborundum in minute quantities. The ctenidia are sim-
ilar to those of Lyonsia norwegica (ATKINS, 1937, p. 350),
Mytilimeria nuttalli (KELLocc, 1915, p. 656) and Ento-
desma saxicola (KELLOGG, op. cit., p. 659) and fall into
Atkins’ Type E (ATKINS, op. cit., p. 409).

Ciliary currents move downward to the margin of the
outer demibranch and continue in the same direction on

Page 308

the descending lamellae of the inner demibranch to its
margin. On the two exposed faces of the ctenidium,
particles are driven ventrally, therefore, while on the
inner lamella of the inner demibranch, the movements of
the particles are also ventral.

Along the marginal groove there is an oral current
moving material on the ventral margin of the inner demi-
branch to be taken on the palp folds and passed to the
mouth (Figure 4).

The ciliary mechanism on the ctenidia of Lyonsia cali-
fornica is similar to that of L. norwegica, described by
Atkins (1937, p. 347).

Frontal cilia extend around the ventral ends of the
filaments; they occur on both sides of each filament and
beat toward the axial groove.

Latero-frontal cilia (10 in length) are situated on a
slight ridge. The lateral cilia are very similar to those of
Lyonsia norwegica.

With the ctenidium deeply plicated, the marginal food
groove has scalloped sides. The marginal groove in Lyon-
sia californica is deep and similar to that of L. norwegica
described by ATKINS (op. cit., p. 350).

Fine particles reach the principal filaments and adja-
cent filaments within the plical grooves. The principal
filaments end at the bottom of the marginal groove.
Particles carried in the principal filaments and the adja-
cent filaments come to the same deep channel.

Coarse particles are kept from entering the narrow
interplical space and are carried on the plical crests or
passed directly off the ctenidia, or are passed toward the
mouth to fall on the mantle where they are removed.

The animal could contract the ctenidia, covering the
principal filaments, and thus exposing a reduced surface
to the current being directed to the mouth.

The ventral tips of the anterior filaments of the inner
demibranch are inserted and fused to a distal oral groove
and fall in Category II of the association of ctenidia and
labial palps (Stasex, 1963, p. 91).

THE LABIAL PALPS

The labial palps are large, active and continually coiling.
Each palp lamella bears 20 ridges on the inner surface.
The inner demibranch of the ctenidium projects deeply
between the palps. Particles form a narrow stream that
moves in the oral groove, and are subjected to selection
by the folds of the palps, eventually passing over them to
the mouth. If material is not removed, it continues for-
ward and is carried into the proximal oral groove.
Material bound in mucus which passed to the palp from
the inner demibranch was rapidly transported anteriorly

THE VELIGER

Vol. 10; No. 4

and ventrally, being rejected at the free ventral margin
of the palp.

The following ciliary currents may be recognized:

(a) On the floor of the groove between adjacent folds,
particles are driven to the ventral border of the palp
and from there moved to the tip of the palp where they
are rejected (Figure 5a, 5b).

(b) On the crests of the folds and on the superficial
slopes, particles are carried forward from fold to fold
toward the mouth (Figure 5a, 5b).

(c) On the crests of the folds toward the dorsal border
of the palp, particles are moved to the dorsal part of
the labial palp where they form a resorting current.

In Lyonsia californica there is one main acceptance
current, one rejection current and one dorsally directed
resorting current (Figure 5).

Under normal conditions particles of carmine and car-
borundum are carried rapidly forward and downward
over the folds, traveling over the folds without being

Figure 5

a — Section of the labial palps; @ indicates the position of a
proximal current; x indicates the position of a distal current;
arrows show the dircction of an oral current
b - Diagrammatic representation of the ciliary mechanisms on the
folded surface of the labial palps

Vol. 10; No. 4

deflected dorsally by the resorting current. However, some
are taken on the crests of the ridges and transported dor-
sally by the dorsal resorting current. Particles are accu-
mulated at the base of the palp and, with particles from
the ctenidial margin, are passed into the lateral oral
groove, then to the mouth where they are ingested.

Very few particles were carried in the deepest part of
the grooves to be rejected by the free ventral border of
the palp, as observed in Petricola pholadiformis La-
MARCK, 1818 (PurcHoN, 1955, p. 264).

THE VISCERAL MASS AND THE FOOT

The visceral mass of Lyonsia californica is very similar to
that of L. norwegica, as described by PELSENEER (1891,
p. 210). The digestive diverticula are placed anteriorly
and ventrally, with the kidney posterior to the pericardi-
um. This species is hermaphroditic, possessing two large
pairs of gonads each with its own duct. The ovary
occupies the dorsal region of the visceral mass and is
surrounded by the testis which fills the ventral region of
the digestive diverticula. Observations were made during
the process of spawning, which occurred in April. Great
numbers of eggs left the exhalant siphon followed in the
same animal by a discharge of sperm. This alternation
between the discharge of eggs and sperm continued for
a few minutes. Fertilization occurs in the sea water as in
Entodesma saxicola and Mytilimeria nuttalli (YONGE,
TO52 pyre)

The foot is slender, and, when fully distended, is half
the length of the shell. There is a well developed byssus
gland with a groove extending along the ventral side of
the foot almost to its tip. The construction of byssus
threads by adult animals was observed in the laboratory.
YoncGE (1952, p. 446) states that the foot and pedal gape
are small in Lyonsia norwegica. Lyonsia californica is
capable of burrowing. A specimen 20mm long was
observed to bury itself in 25 minutes. Once buried, the
animal remains in the same position for a long time,
which is indicated by the algal growth on the posterior
end of the shell valves.

MUSCULATURE

The anterior adductor muscle of Lyonsia californica is
smaller than the posterior adductor muscle, and both are
located on the antero-posterior axis of the body.

The anterior pedal retractor muscle arises a short
distance posterior to the anterior adductor, is poorly de-
veloped and its fibers spread out immediately below the

THE VELIGER

Page 309

epidermis on the antero-dorsal surface of the visceral
mass.

The posterior retractor muscle arises anteriorly to the
posterior adductor muscle, is well developed and inserts
at the postero-dorsal part of the visceral mass.

Transverse muscle fibers arise in the epithelium of the
visceral mass and pass inward either to the walls of the
alimentary canal or transversely across the visceral mass
to the opposite side.

THE ALIMENTARY CANAL
GENERAL STRUCTURE

The alimentary canal (Figure 6) was dissected from
specimens that had been relaxed with magnesium chloride
and preserved in alcohol. The internal structures of the
stomach were studied in living animals. The stomach was

Figure 6

General dissection of the alimentary canal
ap — anal papilla arm — anterior retractor muscle
cssi — combined style sac and mid-gut f — foot k — kidney
mg — mid gut mo — mouth oe — oesophagus
omg — origin of the mid-gut at the base of the style sac
pam — posterior adductor muscle
prm -— posterior retractor muscle

r — rectum
st — stomach

opened by a mid-dorsal incision through the roof, and
the right side of the stomach was drawn downward
(Figure 7). The ciliary currents were investigated with
the aid of carmine, aquadag and fine carborundum
particles. The nomenclature used by GraHam (1949),
Owen (1953), PurcHon (1955) and Rew (1965) has
been followed.

The mouth (mo) opens into a long oesophagus (oe)
that enters at the anterior part of the stomach (st). The
lumen of the oesophagus is dorso-ventrally compressed.
The stomach consists of two parts: a globular anterior
region and a posterior elongated region. The combined

Page 310 THE VELIGER Vol. 10; No. 4

Figure 7

The interior of the stomach, seen from the right side, after opening
by making an incision in the right wall

af — anterior fold at — acceptance tract oe — oesophagus p — ciliated pad pf — posterior fold
cd, — left collecting duct cd, — right collecting duct rt — rejection tract sa — sorting area
dg —- dorsal groove dh —- dorsal hood gs — gastric shield sa, — posterior sorting arca ss — style sac

ig — intestinal groove Ip — left pouch mt — minor typhlosole ty — major typhlosole

Vol. 10; No. 4

THE VELIGER

Page 311

style-sac and intestine open into the posterior and ventral
region of the stomach. The intestine is convoluted in the
region anterior to the stomach and then passes backward,
proceeding posteriorly and dorsally over the kidney and
the posterior adductor muscle to open at the anus.

STRUCTURE OF THE STOMACH

Lyonsia californica has a stomach of Type 4 as defined by
Purcuon (1958, p. 488). The minor typhlosole (mt)
ends where the anterior globular part of the stomach
begins, so that the intestinal groove (ig) running across
the floor of this region has no fold along it on the right
side, though its left margin is bordered by the extension of
the major typhlosole (ty). The major typhlosole is thick,
large, and extends across the opening of the mid-gut to
come in contact with the minor typhlosole. As in other
bivalves which have stomach Type 4, the major typhlosole
and the intestinal groove pass forward over the floor of
the stomach, curving gradually to the left. The major
typhlosole does not possess a tongue, nor does it enter any
of the orifices of the ducts from the digestive diverticula
on the anterior right side of the stomach (PurcHon,
1958, p. 489).

The dorsal hood (dh) is large and directed anteriorly
to the left. It opens posteriorly on the left wall of the
stomach and arches forward above the oesophagus and
opens into the stomach anteriorly. The ventral wall of
the stomach is covered with a finely ridged sorting area,
posteriorly bordering on the intestinal groove which dis-
appears, on the right side of the major typhlosole, into
the intestine. The sorting area is composed of a regular
series of alternately long and short folds, separated from
one another by grooves with diverse ciliary currents
setting up a sorting mechanism. The posterior sorting
area (sa:), which forms a Type A sorting mechanism
(Rem, 1965, p. 159), is well developed, extending dor-
sally over the right wall of the stomach and on to the
right wall of the dorsal hood. The posterior margin of
the sorting area is formed by a well developed fold (pf ),
while the anterior margin is bounded by the rejection
ciliary tract (rt) which empties into the intestinal groove
(ig) on the floor of the stomach at the point where the
minor typhlosole ends. The anterior margin of the rejec-
tion tract is formed by a second fold with several small
parallel ridges (af ).

The gastric shield (gs) is well developed and attached
to the left wall of the stomach by the teeth extending into
the apertures of the left pouch (lp) and the dorsal hood.
The left pouch is a blind hollow into which the anterior
border of the gastric shield fits. This is similar to Pandora

inaequivalvis (PurcHON, 1958, p. 509). The left pouch
of the Lyonsiidae falls into Section III, Group B of
Dinamant (1967, p. 262).

In the ventral part of the stomach wall there is a fan-
shaped sorting area (sa), with a series of folds and ridges
that have the same arrangement as the posterior sorting
area, and drains particles into the intestinal groove. This
well developed area extending from the mid-gut pene-
trates the two large apertures, one on the left anterior wall
and the other on the right anterior wall. A similar condi-
tion exists in Pandora. The aperture of the left wall of
the stomach was termed by ALLEN the “left caecum.” In
Lyonsia californica there are 17 to 20 folds, while in P
maequivalvis there are 10 to 15. ALLEN (1954, p. 480)
described this region as the “lateral rejection grooves.”
All ducts of the digestive diverticula open into the two
larger apertures. Dissection of the main collecting ducts
(cd) revealed the apertures of the ducts which open into
them. Five ducts from the digestive diverticula open
into the left aperture and 3 of them open into the right
aperture of the stomach.

Particles accepted by the palps are carried through the
oesophagus and carried to the roof of the stomach by the
dorsal groove (dg), where they are caught by the tip of
the crystalline style and directed to the posterior sorting
area by its revolving action. The crystalline style in Lyon-
sia californica is large (7 mm long in a specimen of 19mm
overall length) and rotates in a clockwise direction when
viewed from above. The particles in the posterior sorting
area are directed against the ridges of the anterior fold.
From here they are carried by the revolving style to the
ridged surface of the posterior sorting area, where heavier
particles are carried by the cilia of the grooves to the re-
jection tract (rt) and to the mid-gut by the intestinal
groove (ig). The finer particles are conveyed dorsally
from crest to crest across the ridges. Only the lighter par-
ticles are retained in the distal region of the dorsal hood,
where they are included in the mass at the tip of the
crystalline style. The acceptance tract (at) conveys par-
ticles along the roof of the dorsal hood and over the
posterior wall of the stomach to the mid-gut.

The ciliary currents of the dorsal region convey fine
particles from the tip of the style to the acceptance tract.
Particles in the sorting area (sa) are conveyed in the
grooves, by cilia, to the intestinal groove. Cilia on the
crests of the folds beat from the right to the left, carrying
particles to the left region of the stomach. Here, coarse
particles are rejected in the grooves, while fine material
passes across this region on the crests of adjacent ridges
by ciliary action.

Particles that do not enter the digestive ducts are
carried towards the oesophageal region of the stomach

Page 312

where they join others that come from the oesophagus,
and then both are carried to the base of the dorsal hood
by the dorsal groove. The dorsal groove was noted by
ALLEN (1948, p. 480), but not by PurcHon (1959, p. 509)
in Pandora inaequivalvis.

The stomach of Lyonsia californica is similar to that
of Pandora inaequivalvis. The single duct from the diges-
tive diverticula which ALLEN (1954) established as en-
tering the middle of the floor of the stomach and which
he erroneously interpreted as “right caecum” (PuRCHON,
1958, p. 508), and not seen by PurcHoN in the same
species, is not present in Lyonsia californica.

DISCUSSION

Lyonsia californica is adapted to live in a substratum of
soft mud, and is found on muddy beaches. Algal growth
confined to the siphonal region indicates that the animal
buries itself in such a way that only the posterior part of
the shell and siphons are exposed. The siphons are sensi-
tive, and it is not probable that large quantities of mate-
rial are normally taken into the pallial cavity. The siphons
are short and never extend more than a few millimeters
beyond the end of the shell.

The species has a well developed foot used to dig, and
to orientate the animal when it digs.

The animal lives in quiet waters with little disturbance
of the bottom deposits so that the water contains little
suspended material. The presence of well developed sor-
ting areas in the stomach is evidently desirable to reject
large particles or excessive quantities of small particles
(PurcHon, 1960, p. 487).

The stomach is very similar to that of Pandora inacqui-
valvis, which was studied by ALLEN (1954) and PurcHON
(1958). In Lyonsia californica there is no simple duct
from the digestive diverticula that PurcHoN named
DDD: and which ALLEN (1954, p. 480) interpreted as the
right caecum and established as entering the floor of the
stomach and communicating with the intestinal groove
by a special groove. PurcHON (op. cit., p. 508) was
unable to find this duct in the animals that he studied but
he inserted it later, as he said, in the figure of the stomach
of P inaequivalvis.

In Lyonsia californica the major typhlosole accompa-
nied by the intestinal groove passes toward the openings
of the collecting ducts and curves across the opening of
the mid-gut to come into contact with the minor typhlo-
sole. PurcHoNn believed that in Pandora inaequivalvis the
major typhlosole and the intestinal groove terminate at
the posterior border of the mouth of the collecting duct,
which is situated on the left anterior wall of the stomach. .

THE VELIGER

Vol. 10; No. 4

As Purcuon found in Pandora inaequivalvis, the diges-
tive diverticula do not open into the left pouch.

A groove on the roof of the stomach and in which cilia
beat backward and convey particles toward the dorsal
hood, exists in Lyonsia californica, and this was also found
in Pandora inaequivalvis by ALLEN. It was not seen by
Purcuon when he studied the same species of Pandora.

The stomach of Lyonsia californica resembles in gen-
eral the stomach of Anodonta cygnea (GraHAm, 1949, p.
745), but is much more similar to that of Pandora
inaequwwalvis.

PurcHon (1958, p. 488 and 1960, p. 432) made an
intensive study of the stomach in Bivalvia and found in
22 families the stomach that he defined as Number 4.
The family Lyonsiidae is now the 23" family known to
have this type of stomach.

Of 4 families in the super-family Pandoracea of the
sub-order of Anomalodesmacea (Axssott, 1954, p. 468)
we know that 2 have stomach type Number 4, Pandoridae
and now Lyonsiidae. Future studies will prove if in the re-
maining 2 families, Periplomatidae and Thraciidae, the
stomachs are of the same type as defined by PurcHon,
proving that the form of the stomach is an indication of
the phylogeny of the bivalves and not a consequence of
the adaptations to different environments.

SUMMARY

Lyonsia californica Conrad occurs in muddy substrates
on the Pacific coast of North Amcrica from Puget Sound
to Lower California. The calcarcous shell is thin and
largely nacreous. The periostracum is very thick and sand
grains adhere to it. A lithodesma is present and dorsally
the shell valves are united by the periostracum. The
mantle edges are fused except in the region of the small
fourth pallial aperture and along the relatively well de-
veloped pedal gape.

The anatomy and functioning of the stomach is de-
scribed in detail. It belongs to type Number 4 as defined
by PurcHon (1958, p. 488). The stomach of Lyonsia
californica is similar to that of Pandora inacquivalvis.

ACKNOWLEDGMENTS

I wish to express my thanks to Dr. Edmund H. Smith,
Director of the Pacific Marine Station, Dillon Beach,
California, for the use of the facilities and laboratorics.
To the staff of the Marine Station my thanks are due for
assistance in collecting.

I also wish to thank Dr. Charles R. Stasek for his
valuable suggestions and constructive criticism of this

paper.

Vol. 10; No. 4 THE VELIGER Page 313
This work was made possible by a grant from the Morse, Epwarp SYLVESTER
Department of the Interior (W P01061) and by financial 1919. | Observations on living lamellibranchs of New England.

help of the Coordenagao do Aperfeicgoamento de Pessoal
de Nivel Superior (CAPES).

LITERATURE CITED

Assott, Rospert TUCKER

1954. American seashells. Princeton, New Jersey. D. van

Nostrand Co., Inc.; xiv + 541 pp.; 100 figs.; 40 plts.
ALLEN, JoHN A.

1954. On the structure and adaptations of Pandora inaequi-
valuis and P. pinna. Quart. Journ. Micr. Sci. 95 (4): 473
to 482

ATKINS, DAPHNE

1937. On the ciliary mechanisms and interrelationships of
lamellibranchs. Part II. Sorting devices on the gills. Quart.
Journ. Micr. Sci. 79: 339 - 373

Part III. Types of lamellibranch gills and their food currents.
Quart. Journ. Micr. Sci. 79: 375 - 421
Part IV. Cuticular fusion with special reference to the 4‘
aperture in certain lamellibranchs. Quart. Journ. Micr.
Sci. 79: 423 - 444
Bgccitp, O. B.

1930. The shell structure of the mollusks. D. Kgl. Danske
Vidensk. Selsk. Skr., Naturn. Math. Afd., 9R II 2: 231 - 325;
pits. 1-15

Boss, KENETH J. & ARTHUR S. MERRILL

1965. The family Pandoridae in the western Atlantic.

sonia 4 (44): 181-215
DinaMaANnl, P.

1967. Variation in the stomach structure of the Bivalvia.

Malacologia 5 (2) : 225 -268
GraHaM, ALASTAIR

1949. The molluscan stomach.

61: 737 - 778
KEEN, A. MyRA
1937. An abridged check list and bibliography of West North

American marine mollusca. Stanford Univ. Press, Stanford,
Calif. pp. 1 to 88.

Kextoce, J. L.

John-

Trans. Roy. Soc. Edinb.

1915. Ciliary mechanisms of lamellibranchs with descriptions
of anatomy. Journ. Morph. 26 (4): 625-701

Proc. Boston Soc. Nat. Hist. 35: 139 - 196
Owen, GarETH

1953. On the biology of Glossus humanus (L.) (Isocardia
cor Lam.). Journ. Mar. Biol. Assoc. U. K. 32: 85 - 106

PELSENEER, PAUL
1891. Contribution 4 I’étude des lamellibranches. Arch.
Biol. 11: 147 - 312.
Purcuon, R. DENISON

1955. The functional morphology of the rock-boring lamelli-
branch Petricola pholadiformis, Lam. Journ. Mar. Biol.
Assoc. U. K. 34: 257 - 278

1955. The structure and function of the British Pholadidae.
Proc. Zool. Soc. London 124: 859 - 911

1957. The stomach in the Filibranchia and Pseudolamellibran-
chia. Proc. Zool. Soc. London 129: 27 - 60

1958. The stomach in the Eulamellibranchia; Stomach Type
IV. Proc. Zool. Soc. London 131: 487 - 525

1959. Phylogenetic classification of the Lamellibranchia, with
special reference to the Protobranchia.
London 33 (5): 224 - 230

1960. The stomach in the Eulamellibranchia; Stomach Types

Proc. Malacol. Soc.

IV and V. Proc. Zool. Soc. London 135 (3): 431 - 489
Rem, R. G. B.
1965. The structure and function of the stomach in bivalve
molluscs. Journ. Zool. 147: 156 - 184

RipEwoop, W. G.
1903. On the structure of the gills of the Lamellibranchia.
Phil. Trans. Roy. Soc. London (B) 195: 147 - 284
STAseEK, CHARLES ROBERT
1963. Synopsis and discussion of the association of ctenidia and
labial palps in the bivalved Mollusca. The Veliger 6 (2) :
91-97; 5 text figs. (1 October 1963)
THIELE, JOHANNES ;
1963. | Handbuch der systematischen Weichtierkunde, Teil 3.
Classis Bivalvia. i-v + 779-1154 pp.
YoncE, CHarLes Maurice
1952. Structure and adaptation in Entodesma saxicola (Baird)
and Mytilimeria nuttallii Conrad, with a discussion on evolution
within the family Lyonsiidae (Eulamellibranchia). Studies on

Pacific mollusks I1V- VI. — Univ. Calif. Publ. Zool. 55 (9-11):
421 - 454

Page 314

TMENVERIGER

Vol. 10; No. 4

Structure of the Bivalve Rectum

II. Notes on Cell Types and Innervation

THOMAS C. JEGLA

Department of Biology, Kenyon College
Gambier, Ohio 43022

MICHAEL J. GREENBERG

Department of Biological Sciences
Florida State University, Tallahassee, Florida 32306

(Plates 45 to 48)

INTRODUCTION

RECENTLY, THERE HAS BEEN increasing interest in the
pharmacology (GREENBERG & JEGLA, 1963; GREENBERG,
1966; Puts, 1966) and physiology (Prosser, Nyst-
roM & Nacai, 1965; Nystrom, 1967) of the bivalve
rectum as an example of an invertebrate organ with
smooth muscle. While these studies have all been con-
cerned with the innervation of the nonstriated muscle of
the rectum, the histology of this structure has been but
briefly noted (GREENBERG & JEGLA, 1962; PuiLuis, 1966).

Although the structure of molluscan muscles has been
intensively studied (see Hanson & Lowy, 1960, 1965;
Hoyte, 1964), the bulk of these investigations have
concerned skeletal muscles such as shell adductors and
byssus retractors. Consequently, with the exception of
work on heart (reviewed by Hitt & WELSH, 1966;
NisBET & PLuMMER, 1966), visceral musculature has
largely been ignored.

We have shown that the gross histology of bivalve
rectums varies markedly from species to species (JEGLA

& GREENBERG, 1968). Also, GREENBERG (1966) described
striking pharmacological differences between the acetyl-
choline responses of two intrageneric species. In the
present study we have looked for specific differences in the
cellular elements which might be correlated with gross
structural or pharmacological differences. In addition
we have made observations of the pattern of innervation
of the rectal musculature of some spccics. Finally, we
have compared the structure of the rectal muscle fibers
with those of the heart and other molluscan muscles.

MATERIALS anp METHODS

The species of bivalve mollusks and methods used in this
study were identical to those previously reported (JEGLA
& GREENBERG, 1968). In addition, for observation of
nerves, we have used a modification of Bodian’s silver
staining technique as well as the method of KoELLE
(1951) for acetylcholinesterase.

Explanation of Plate 45

Figure 1: A transverse section through the wall of a Mya arenaria
rectum showing the columnar epithelium, a layer of collagen fibers
and circular muscle arranged in bundles.

C —- columnar epithelium

CTF -— connective tissue fibers

M — muscle cell

Figure 2: Quadrula quadrula (RaFINESQUE, 1820). Muscle is ar-
ranged in bundles and embedded in a network of connective tissue.
Figure 3: Amblema plicata RAFINESQUE, 1820. Connective tissue
in the typhlosole.

CM - circular muscle
F — fibroblast cell
P — phagocyte

Tue VELIGER, Vol. 10, No. 4 [JEcLA « GREENBERG] Plate 45

i)
bo SMG
t
’ 7!

Figure 1

Vol. 10; No. 4

RESULTS

The wall of the bivalve rectum is composed of an inner
columnar epithelium, connective tissue, and muscle layers
(Figure 1). The epithelium is separated from the more
peripheral elements by a basement membrane. These
elements, as well as innervation of the muscle are dis-
cussed below.

Epithelial Cells

Details of the columnar epithelial cells lining the
digestive tract lumen in Anodonta cellensis ScHROTER,
1779 (see GuTHEM, 1912) and Ostrea edulis LINNAEUS,
1758 (see YoncE, 1926) have been described. Only a
few additional comparative observations will be given
here. These cells are much higher than wide but their
average height varies depending, not only on the species,
but also on the size of the individual animal. The
smallest cells were found in Nucula (7.54) and the
largest in Mytilus californianus Conrad, 1837 (115p).
In a very small Actinonaias carinata (BARNES, 1823) the
average cell height was about 26m, while in a large
specimen of the same species, it was about 74. In some
species, height of the cells may vary greatly around the
periphery of the lumen; this is observed in conjunction
with ridges and furrows, and with typhlosoles. In Mytilus
edulis LiINNAEuS, 1758 the columnar cells are twice as
large on the ventral side of the rectum as they are on the
dorsal side (see Part I, plate 37, fig. 3). The reverse
situation occurs in Actinonaias (see Part I, plate 39,
fig. 7) and other unionids. Furthermore, the cytoplasm
of the small cells bordering the unionid typhlosole stains
more intensely than that of the taller cells on the oppo-
site side of the rectum.

Daxin (1909) thought that cilia are poorly developed
in the rectum of Pecten maximus (Linnaeus, 1758),
and Wuirte (1937) reported that cilia are long but
scantily developed in the rectum of Mytilus edulis. How-
ever, rectal cilia were fairly abundant in all the species
we studied. The mean length of the cilia in most species
was between 6u and 12. Conspicuously smaller cilia
were observed in Nucula and Dinocardium robustum
(Licutroot, 1786) [about 4], and larger cilia in
Chlamys and Mya arenaria Linnagus, 1758 (15u to
16).

Mucous cells and phagocytes occur in the columnar
epithelial layer and were described by YonceE (1926) in
Ostrea, and by GutHett (1912) in Anodonta. The fib-
rous nature of the underlying basement membrane is
easily observed in Atrina rigida (LicHTFoot, 1786)
(Plate 46, Figure 4). There is some small interspecific
variation in thickness of this membrane but in all species

THE VELIGER

Page 315

it is thin (lu to 5u). The outer surface of the rectum is
covered by a very thin epithelium which is observed
only with difficulty.

Connective Tissue Fibers and Cells

The numerous, randomly oriented fibers in the rectum
have the staining properties of collagen (Plate 45, Plate
46, Fig. 4). They are considerably smaller than the
average muscle fiber, and are less than 1p in diameter.
Their length is not known because they are contorted and
their ends could not be observed in histological sections.
This feature is shared with the collagen fibers in verte-
brate connective tissue. A reticulum of very small fibers
is frequently observed around the individual muscle
fibers.

The prominent cellular elements of the connective
tissue are phagocytes, fibroblasts, and vesicular cells.
Phagocytes are numerous among the columnar cells,
throughout the wall of the rectum and in the circulation.
These cells were described extensively by GuTHEIL
(1912) and Yonce (1926). While the amount of cyto-
plasm is variable, the nucleus is spherical, small (2.5u
to 4, in diameter), and hyperchromatic (Plate 45, Fig.
3) ; these characteristics serve to distinguish a phagocyte
from any other cell in the rectum.

Fibroblast cells are a universal and common component
of the connective tissue. They are fusiform or stellate-
shaped, with fibrous processes extending from the cell
body. The fusiform type is 2.51 by 4 to 12 in size; it has
one or two fibrous extensions and the nucleus occupies
most of the cell body (Plate 45, Figure 3). The stellate
cells average 54 by 7; they have three or more long
fibrous extensions and a small spherical nucleus. Fibro-
blasts are best observed in typhlosoles, since they are
filled, principally, with fibrous connective tissue.

The rectums of a few species have vesicular cells that
are spherical to oval in shape. These are the largest con-
nective tissue cells in the rectum (up to 134 by 40y);
they resemble those cells surrounding the digestive tract
in the visceral mass. Gencrally, vesicular cells are empty
after fixation; the sparse cytoplasm and small nucleus
are located centrifugally.

Muscle

At the light microscope level all of the muscle fibers
in the bivalve rectum are nonstriated. They are spherical
or oval in transverse section (Plate 46, Figure 5), and
have a diameter of 1 to 6; the larger diameters may
be the result of contraction. The fibers are very long, at
least 170, but we were unable to determine their exact
length. The muscle cells are uninucleate; the nucleus is
large, oval, and with its major axis parallel to the longi-

Page 316

THE VELIGER

Vol. 10; No. 4

tudinal axis of the fiber (Plate 46, Figure 5). In all
species the muscle nuclei can be distinguished from those
of fibroblasts by the larger size and lighter staining
quality of the former.

The fibers are arranged in bundles in Mya (Plate 45,
Figure 1) and the family Unionidae (Plate 45, Figure 2),
but there is little indication of such grouping in Cardi-
idae or Mactridae which have the muscle fibers packed
closely together throughout the area of their occurrence
in the rectum. In those species with a scarcity of muscle,
and in the Veneridae, the muscle cells are generally not
organized into bundles but lie embedded as individual
units in the meshwork of connective tissue fibers and
cells (Plate 47, Figure 6).

Hanson & Lowy (1957) distinguished three kinds of
molluscan nonstriated muscle on the basis of the distri-
bution, in the cell, of myofibrils, the course (e. g. helical
or straight) of the myofibrils, and the appearance of
“contraction bands” during excessive shortening of the
muscle. We have examined these characteristics.

Myofibrils are not restricted to the periphery of the
fiber in the rectum. While this is especially clear in
transverse sections of Dinocardium muscle (Plate 46,
Figure 5), an even distribution of fibrils in the sarco-
plasm has been found in all of the species examined.

Rectumsof Mercenaria mercenaria (LINNAEUS, 1758),
fixed either while tied at an extended length, or while in
5-hydroxytryptamine contracture, were compared. In the
stretched rectum, the smallest fibers have the classic ap-
pearance of vertebrate smooth muscle; no myofibrils are
visible. But the larger fibers have longitudinal fibrils
lying parallel to the long axis; they were not helically
arranged. We were unable to determine the length of
these fibrils. In Mercenaria rectums highly contracted by
5-hydroxytryptamine, many of the fibers appear to have
contraction bands which cross each other at angles of
up to 20° to the fiber axis, producing the classical double-
oblique, or diamond lattice pattern. Many of our prep-
arations from other species also showed this pattern

following fixation or 5-hydroxytryptamine contracture
(Plate 46, Figure 5).

Comparison of Heart and Rectal Muscle

The nonstriated muscle fibers of the heart of Mercena-
ria are different from those of the rectum in that the
myofibrils are peripherally disposed (Plate 47, Figure
7). This difference in the distribution is probably not due
to artifact since both tissues were necessarily fixed, em-
bedded and stained together. However, NisBeT & PLuM-
MER (1966) examined the fine structure of the heart of
achatinid snails and found that the contractile elements
were centrally disposed in the muscle cells, occupying
about 2 of the cross-sectional area.

The myofibrils of the Mercenaria heart appear to spi-
ral and helical “contraction bands” were observed Plate
47, Figure 8). Often the fibrils are observed to cross at
acute angles, producing the classic diamond lattice pat-
tern of Marceau (1905, 1909). A reticulum of very fine
connective tissue fibers surrounding individual muscle
fibers was often observed. Larger collagen fibers and
fibroblast cells are found between the heart muscle cells.
Nuclei are centrally placed and appear to be more nu-
merous in heart muscle; perhaps the fibers are shorter,
or syncytial as was proposed for the Mercenaria heart
(LovELAND, 1963).

Nervous Tissue

Intrinsic nerve fibers and cells have not often been
demonstrated in bivalve muscles. Brick (1914) illus-
trated small nerve fibers ending in motor end plates on
muscle cells. BowpEN (1958), using the cholinesterase
technique of KoeLie (1951), demonstrated small nerve
fibers and motor end plate-like structures in the fast
part of the adductor muscles of Anodonta and Unio; in
other muscles studied, only free terminations were ob-
served. BowbeEN also reported ganglion cell-like structures
and nerve fibers in the slow parts of bivalve adductors.
Puitus (1966) observed small nerve fibers and a gan-
glion cell in the rectum of Tapes watling: IREDALE, 1958.

Occasionally we have scen ganglion cells and nerve
fibers in the rectum of Dinocardium. The nerve fibers
bifurcate near their ends and terminate on different

Explanation of Plate 46

Figure 4: Atrina rigida (LicHTFoot, 1786). Shows the fibrous

nature of the basement membrane.

Figure 5: Dinocardium robustum (LicHTFoor, 1786). Muscle cells

of the rectum in longitudinal and transverse section. Note the
BM - basement membrane

distribution of myofibrils in the transverse section and the occur-
rence of the diamond lattice pattern in the longitudinal sections
of muscle cells.

C - columnar epithelium

N — nucleus

[JEGLA & GREENBERG] Plate 46

Tue VELIGER, Vol. 10, No. 4

bila!

ea ig

Vol. 10; No. 4

THE VELIGER

Page 317

muscle cells. Nervous tissue was also demonstrated in
Mercenaria mercenaria rectum with silver staining and
the cholinesterase technique of Korette (1951). Appar-
ently a nerve plexus lies at the base of the epithelial cells
lining the lumen; fine nerve fibers run transversely (peri-
pherally) from the plexus (Plate 48, Figure 9). Longitu-
dinal collateral fibers are given off by these radial nerves
which finally branch and terminate on muscle fibers
(Plate 48, Figure 10). In addition, larger nerve fibers
were also found running parallel to longitudinal muscle,
then branching, with the small arborizations coursing
transversely to end, again, on individual muscle fibers.
Occasionally we have observed small unipolar and bipo-
lar ganglion cells (2 by 54) with fine, branching fibers.
We have infrequently observed end plate-like structures
in Mercenaria and Dinocardium rectums. Mostly, the
nerve-muscle junctions were of the en passage type with
only a small increase in density of staining where the
nerve fiber lay athwart the muscle cell.

DISCUSSION

The structure of the bivalve rectum, in each species, is
different from that of the more anterior intestinal portion
of the gut (Voct « Yunc, 1888; GuTuHeEiIL, 1912;
YoncE, 1926; Gattsorr, 1964). For example, the mus-
culature is usually better developed or, at the least, a
denser network of connective tissue is present in the
rectum. Also, the number of cilia in the rectum has been
reported as being smaller than in the intestine. The
physiological significance of such differences is not clear,
but there have been suggestions.

GuTHEIL (1912) believed that longitudinal and cir-
cular muscle layers in the rectum of Anodonta cellensis,
which are absent in the intestine, provide support for the
rectum where it hangs free in the pericardial and ventri-
cular cavities. However, this hypothesis would appear to
be contradicted by the fact that many species, regardless
of rectal size, have little or no tissue peripheral to the
basement membrane (e.g., Nucula and Macoma).

Probably no peristalsis occurs in the intestine; material
is moved along the tract by the ciliated columnar epi-
thelium. GuTHEIL noted (1912) that, while contractions
in the intestine would be impeded by the surrounding
tissue of the visceral mass, peristaltic action might occur
in the freely suspended, muscled, scantily ciliated rectum.
In fact, spontaneous contractions of in vitro prepara-
tions of rectums have been observed in Mercenaria mer-
cenaria by GREENBERG & JEGLA (1963), Tapes watlingi
by Puituis (1966), Spisula solidissima (Dittwyn, 1817)
by Prosser, Nystrom & Nacar (1965) and Nystrom
(1967) and in other bivalves (see GREENBERG, 1966).

Prosser et al also observed spontaneous contractions
and associated electrical activity in in vivo preparations
of the Spisula rectum.

The histological architecture of the bivalve rectum
supports the view (Prosser et al, 1965) that conduction
in this organ is by way of nerves rather than muscle. In
most muscular bivalve rectums there is a large amount
of connective tissue in which the muscle cells are embed-
ded. Even in those species where muscle bundles are
found (Mya and Unionidae), there are usually many
individual fibers scattered through the collagen fiber net-
work. Such an arrangement of the muscle appears to
preclude electrical conduction from cell to cell such as
occurs in the vertebrates and Echinodermata (PRossER
et al, 1965). Furthermore, our observations of silver-
impregnated preparations of Mercenaria rectum indicate
that while longitudinal nerve fibers are present, they are
small. And Prosser, Nystrom & Nacar (1965) found
the conduction velocity in the Spisula solidissima rectum
(8 -9cm/sec at 20°C) to correspond to conduction in
such small, non-myelinated nerve fibers.

A wide array of physiological and pharmacological
experiments, with rectums from a variety of bivalve
species, have led to two broad predictions concerning the
pattern of innervation of the musculature (GREENBERG
& JecLA, 1963; Prosser et al, 1965; Puituis, 1966):
Firstly, there ought to be ganglion cells present. These
have been demonstrated by Puitiis (1966), and we
have now scen them in Mercenaria and Dinocardium.
Secondly, there should be multiple innervation of the
muscle cells. In fact, Prosser ct al (1965) proposed that
inhibitory as well as both slow and fast excitatory fibers
innervate the rectum of Spisula solidissima. While we
have not observed innervation of muscle cells by more
than one nerve fiber, only a portion of a particular
muscle fiber appears in any one histological section.
Furthermore, since nerve fibers are difficult to observe,
multiple innervation is not ruled out.

Recently, BuRNstTocK, GREENBERG, KirBy & WILLIS
(1967) studied the rectum of the chiton, Poncroplax.
They were able to correlate the clectrophysiology of
muscle with its innervation. Thus, the intestine, which
acts like “unitary” smooth muscle (sec Burnstock, Hot-
MAN & Prosser, 1963), was markedly less densely inncr-
vated than the rectum which had the properties of a
“multiunit” muscle. So far, no such corrclation between
structure and function has been made with bivalve intes-
tinal muscle.

The muscle fibers in bivalve rectums appear to belong
to Hanson & Lowy’s (1957) second of three major
types of molluscan smooth muscle distinguishable in the
light microscope; namely, a smooth muscle having the

Page 318

classical appearance at physiological lengths and oblique
or helical bands at excessively short lengths. In relaxed
or stretched rectums the muscle fibers of bivalve molluscs
are without visible fibrils or have fibrils paralleling the
long axis of the fiber, whereas in highly contracted rec-
tums dense bands appear in the fibers which give the
appearance of a diamond lattice pattern. The myo-fibrils
do not appear to be helically wound around the periphery
as they are in the heart, but are dispersed throughout
the core of the fiber.

ACKNOWLEDGMENTS

We are indebted to the staffs of the marine laboratories
at Friday Harbor (University of Washington) and Alli-
gator Harbor (Florida State University) for their kind
cooperation and to Mrs. J. Goldstein and D. Nelson for
their technical assistance.

This study was supported by Research Grant HE-06291
from the National Heart Institute, U.S. P.H.S. Addi-
tional funds were obtained from the University of Illinois
Research Council and the Illinois Marine Biological
Association.

LITERATURE CITED

Bowpen, J.

1958. The structure and innervation of lamellibranch muscle.
In: International Review of Cytology (ed. Bourne, Geoffrey
Howard and James F Danielli) 7: 295 - 335. Acad. Press, New
York, N. Y.

Brick, A.

1914. Die Muskulatur von Anodonta cellensis ScHROT.

Zeitschr. wi. Zool. 110: 481 - 619

THE VELIGER

Vol. 10; No. 4

Burnstock, GrorrrEY, MICHAEL JoHN GREENBERG, S. KIRBY &
A. G. WILLIs
1967. An electrophysiological study of visceral smooth muscle
and its innervation in a mollusc, Poneroplax albida. Comp.
Biochem. Physiol. [in press]
Burnstock, Grorrrey, M. E. HoLMAN & CLirrorp Lapp PRossER

1963. _Electrophysiology of smooth muscle. Physiol.
Rev. 43: 482 - 527
Dakin, WILLIAM JOHN

1909. Pecten. L.M.B.C. Memoirs XVII. The

Univ. Press of Liverpool. 1 - 136; plts. 1-9
GattsorF, PauL SIMON
1964. The American oyster Crassostrea virginica GMELIN.
Fishery Bull. Fish & Wildlife Serv., U.S. Bur. Comm. Fish. 64:
480 pp.; 400 figs.
GREENBERG, MICHAEL JOHN
1966. Species specific effect of acetylcholine on bivalve rectums.
Science 154: 1015 - 1017
GREENBERG, MICHAEL JOHN & THoMAS CyriL JEGLA
1962. Histology of the bivalve rectum. Amer. Zool. 2: 526
1963. The action of 5-hydroxytryptamine and acetylcholine
on the rectum of the venus clam, Mercenaria mercenaria.
Comp. Biochem. Physiol. 14: 275 - 290
GuTHEIL, Fritz
1912. Uber den Darmkanal und die Mitteldarmdriise von
Anodonta cellensis Scuror. Zeitschr. fiir wissenschaftl.
Zool. 99: 444 - 538
Hanson, JEAN & J. Lowy
1957. Structure of smooth muscles. Nature 180: 906 - 909
1960, Structure and function of the contractile apparatus in
the muscles of invertebrate animals. In: Muscle (ed.
Geoffrey Howard Bourne) 1: 265 - 335. Acad. Press, New
York, N. Y.
1965. Molecular basis of contractility in muscle. Brit.
Med. Bull. 21: 264 - 271
Hitt, Ropert BENJAMIN & JoHN HENRY WELSH
1966. Heart, circulation, and blood cells. In: Physiology of
Mollusca (ed. Karl Milton Wilbur and Charles Maurice
Yonge). 2: 126-174. Acad. Press, New York, N. Y.

Explanation of Plate 47

Mercenaria mercenaria (LINNAEUS, 1758)

Figure 6: Muscle cells in the rectum lie embedded as individual
units in the meshwork of connective tissue.
Figure 7: Myofibrils in heart muscle cells cut in transverse section.

CB — helical “contraction bands”

Note the peripheral distribution of the fibrils.
Figure 8: Heart muscle cells in longitudinal section. Note the
“contraction bands.”

M —- muscle cell

Explanation of Plate 48

Nervous tissue in the rectum of Mercenaria mercenaria
(Linnaeus, 1758)

Figure 9: Note the small nerve fibers coursing transversely over the
muscle cells.

F — fibroblast cell

M — muscle cell

Figure 10: Note the branching nerve fiber and the termination of
a nerve fiber on a muscle cell.

NF — nerve fiber

Tue VELIcER, Vol. 10, No. 4 [JEcLA & GREENBERG] Plate 47

Tue VEuicER, Vol. 10, No. 4 [JeEcLa « GREENBERG] Plate 48

wich ™ A ae bet 6 pL 2

Figure 9

Figure 10

Vol. 10; No. 4

THE VELIGER

Page 319

Hoye, GRAHAM
1964. Muscle and neuromuscular physiology. In: Physio-
logy of Mollusca (ed. Karl Milton Wilbur and Charles
Maurice Yonge) 1: 313-351. Acad. Press, New York, N. Y.
Jecita, THomas Cyrit « MicHAEL JoHN GREENBERG
1968. Structure of the bivalve rectum. — I. Morphology.
The Veliger 10 (3): 253 - 263; plts. 36-40; 1 text fig.
(1 January 1968)
KoELLE, GeorcE BRAMPTON
1951. The elimination of enzymatic diffusion artifacts in the
histochemical localization of cholinesterases and a survey of
their cellular distributions. Journ. Pharmacol. Exp. Ther.

103: 153 - 171
LovELAND, RoseErtT E.
1963. Some aspects of the cardioregulation system in Mer-
cenaria (Venus) mercenaria. Proc. XVI Int. Congr. Zool.
2: 106

Marceau, F.
1905. Recherches sur la structure du coeur chez les mollusques.
Arch. Anat. Microscop. 7: 495 - 588
1909. _ Recherches sur la morphologie, Phistologie, et la physi-
ologie comparées des muscles adducteurs des mollusques acé-

phales. Arch. Zool. Exp. et Gen. 2: 295 - 469

Nisset, R. H. « J. M. PLUMMER
1966. Further studies on the fine structure of the heart of

Achatinidae. Proc. Malacol. Soc. London 37: 199 - 208
Nystrom, RicHAarp ALAN
1967. | Spontaneous activity of clam intestinal muscle.

Comp. Biochem. Physiol. 21: 601 - 610

Puiuus, J. W.
1966. Regulation of rectal movements in Tapes watlingi.
Comp. Biochem. Physiol. 17: 909 - 928
Prosser, CiirForp Lapp, RicHarp ALAN Nystrom « T. Nacat

1965. Electrical and mechanical activity in intestinal muscle
of several invertebrate animals. Comp. Biochem. Physiol.
14: 53 - 70
VoctT, Cart & EMILE YUNG
1888. Traité d’anatomie comparée pratique. Reinwald, Paris

WuitE, KATHLEEN M.
1937. Mytilus. L. M. B.C. Memoirs XXXI. The
Univ. Press of Liverpool. 1-117; plts. 1-10
YoncE, CHARLES MaAurRICE
1926. Structure and physiology of the organs of feeding and
digestion in Ostrea edulis. Journ. Mar. Biol. Assoc. U.K.
14: 295 - 386

Page 320

THE VELIGER

Vol. 10; No. 4

Role of Snails’ Disease in the Biological Control

of Achatina fulica Bowpicu, 1822 in the Andamans

PD. SRIVASTAVA

AND

Y. N. SRIVASTAVA

Division of Entomology

Indian Agricultural Research Institute, New Delhi- 12. India

A LEUCODERMIA-LIKE DISEASE of the giant African snail
was observed to cause heavy mortality in our laboratory
culture in 1965 (Srivastava, 1966). Since then experi-
ments have been in progress to see if it could be made use
of to control field populations. Vaco (1955) has studied
10 different diseases of several species of snails in France
and has also reported the mortality of as many as 50) 000
snails in “snail parks.”” Mrap (1959) is tempted to believe
that diseases “successfully hold most snail populations in
check and produce a major effect in population fluctua-
tions.” He has also predicted that in course of time a
number of snails’ diseases would be found and reported
upon. However, enough data to show the definite effect
of diseases on the populations of snails are lacking.

In the course of the survey of the giant African snail
and its natural enemies on the Andaman Islands since
1965, some snails, dead and dying with disease, were ob-
served in the laboratory culture as well as in the field
population. The diseased snails were inactive, suspended

and finally stopped feeding. This confirmed the existence
of the disease in the snail on the Andamans. The diseased
snails collected from the ficld as well as picked up from
the laboratory culture were placed in a field cage of
the size of 10’x10’x10’, ovcrpopulated with normal
snails and having an excess of moisture, the two re-
quirements for the development of disease in snails. This
resulted in mass mortality in the ficld cage, proving con-
clusively that the discase can spread through contact.
Diseased snails were crushed and their aqueous extracts
sprayed over healthy snails in the field cage to sce if
this technique would help in spreading the discase. This
was repeated twice and the results are tabulated in
Table 1.

From Table 1 it is clear that within about a week of the
date of spraying the discased snails’ extract over the
healthy oncs, all the snails diced. This method was then
adopted to spread the disease in the field population of
snails in three heavily infested localities; it was successful

Table 1

Field cage trials with diseased snail’s extract spray

Number of Total number

Sl. No. diseased Date of Date of of healthy
snails spraying observation snails

crushed introduced
1 6 9V1I1967. 13VI 1967 1650
16 VI 1967 0
18 VI 1967 0
2 2 19 VI 1967 23 VI 1967 180
25 VI 1967 0

Number of | Number of Remarks

dead living % imor-

snails snails tality

1350 300 81.8 No mortality was ob-
220 80 73.3 served in the snails kept

under similar conditions

ep v ae except that they were not
127 53 70.5

sprayed with diseased
snail’s extract

53 0 100

Vol. 10; No. 4 THE VELIGER Page 321
Table 2
Field trials with diseased snails’ extract spray
Sl. No. Date of Date of
spraying observation Locality Result Remarks
il 21 V1 1967 23 VI 1967 Humphri No mortality due to disease ob- Soon after spraying it rained hea-
Ganj served or reported vily on 21 VI 1967
27 VI 1967 Humphri No mortality observed or On 23 VI 1967 it rained heavily
Ganj reported throughout
29 VI 1967 Humphri No mortality observed or
Ganj reported
2. 27.V1 1967 29VI 1967 Manglutan Population of snails reported less No rain soon after spraying on
27 VI 1967
3 VII1 1967 Manglutan Heavy mortality, & foul smell
emitted from dead snails
3 29V1I 1967 3 VII 1967 Maymyo Population much less No rain soon after spraying on
27 VI 1967
in spreading the disease in the field population of two of ACKNOWLEDGMENTS

these localities (Table 2). At Humphriganj, spraying was
followed by heavy showers which probably did not give a
chance to the causative organism of the disease to become
established there.

Meap (1956) has reported high incidence (35 to 68% )
of disease in snails, but of low gastropod host specificity,
meaning thereby that the disease of one species of snail
may cause the disease in another species of snail.
Boycott & OLpHAM (1938) have reported the disease of
Helix aspersa Mi.ier, 1776, to be contagious. MuMA
(1954) has reported a disease in the tree snail Drymaeus
dormant (BinNEy, 1857) and considers it to be of “pos-
sible bacterial origin.” The reproductive potential of the
introduced predators is very low, which is to some extent
a disadvantage. Whereas in the case of disease it is not so.
Meap (1955) says that the reproductive capacity of
disease is so great (unlike that of predators) that it can
produce a catastrophic effect quickly upon the snail popu-
lation. Present investigations go to show that the disease
can be a potential agent in the control of snails; and one
way of spreading disease in the field population of snails
is to spray aqueous extract of diseased snails over the field
population.

SUMMARY

Disease of the giant African snail has now been detected
in the culture as well as in the field population of this
species on the Andamans.

Aqueous extract of diseased snails, when sprayed over
healthy populations of the giant African snail, is capable
of spreading the disease in them and finally causing
mortality in about a week.

The authors wish to express their gratitude to Drs. S.
Pradhan and N. C. Pant, Head of the Division and Pro-
fessor of Entomology, respectively, for the facilities ex-
tended and the interest taken in the progress of the work.
The authors are also grateful to the Director of Agricul-
ture, Port Blair, and his staff, more especially Sri Bhagat
Singh, for the active cooperation and assistance given to
the authors at Port Blair.

LITERATURE CITED

Boycott, A. E. « C. OLDHAM
1938. A contagious disease of Helix aspersa.
Soc. London 23 (2): 92-96
GuosE, KrisHNA CHANDRA
1960. A monograph on the giant land snail Achatina (Liss-
achatina) fulica fulica Bownicu. Unpubl. Thesis City Col-
lege, Calcutta
Meap, ALBERT RAYMOND
1955. The giant African (Kalutara) snail in Ceylon.
Ceylon Forester N. S. 2 (1): 47-50
1956. Disease in the giant African snail Achatina fulica Bow-
DICH. Science 123 (3208): 1130-1131
1959. Increasing complexity in the problem of the giant
African snail. Journ. Colo. Wio. Acad. Sci. 4 (11): 51-52
Muma, M. H.
1954, Predators and parasites of the citrus tree snail. Cit-
rus Mag. 16 (10): 8-9
Srivastava, P. D.

Proc. Malacol.

1966. Leucodermia-like disease in the culture of giant African
snail Achatina fulica Bownicu. Indian Journ. Entom. 28
(3): 412 - 413

Voco, C.

1955. quoted by ALBertT RAyMonp Mean, 1961 in “The
giant African snail, a problem in economic malacology.”
Univ. Chicago Press, pp. 1-257; 15 figs; 1 map; 1 chart

Page 322

THE VELIGER

Vol. 10; No. 4

The Egg Mass and Veligers of Limacina helicina Puipps

BY

MADHU A. PARANJAPE

Department of Oceanography, University of Washington

Seattle, Washington

98105

(2 Text figures)

INTRODUCTION

IN THE SUMMER OF 1966, while at the Friday Harbor
Laboratories, Friday Harbor, Washington, I had an op-
portunity to observe the spawn and the veligers of an
euthecosomatous pteropod, Limacina helicina (Purrs,
1774), in the laboratory. In view of the widespread
occurrence of this genus throughout the world oceans,
and in particular of this species in the polar and subpolar
seas, it is rather surprising that there should be so little
information on the breeding of these animals. More than
100 years ago, VAN BENEDEN (1839) described the struc-
ture of the reproductive system of L. arctica (L. helicina)
and PELSENEER (1888) gave a detailed account of the
morphology and anatomy of this species. Massy (1920)
observed many spawning specimens in the material col-
lected in the Antarctic Ocean. She described the spawn-
mass as a transparent, glutinous material connecting many
white, oblong bodies, each of which averaged 132 by
88. Besides this brief note, there are apparently no
other observations on the spawn and veligers of this
species. Thus, the present observations are of some interest.

McGowan (1963), in an extensive study of the distri-
bution of Limacina helicina in the subarctic North Pacific,
recognized two morphologically dissimilar, well-defined
varieties: Variety A restricted to the Northwest Pacific,
and Variety B limited to the Northeast Pacific. Where
the boundaries of the distributions of these two varieties
meet, intergrades AB were found. The present observa-
tions were made on Variety B.

MATERIALS ann METHODS

The animals were collected with a 1.5m diameter ring-
net, in Saanich Inlet, British Columbia, Canada. The net
was towed vertically from 100m to the surface at night
and the contents of the net were poured and diluted in
several polyethylene trays. The animals were sorted out

immediately and were placed in 32-02 glass jars with
screw caps, which were immersed in tanks with running
sea water. In the laboratory, the animals were maintained
in similar containers with unfiltered sea water to which a
mixture of Platymonas sp. and Skeletonema costatum
(GREVILLE) CLEve, 1878 was added. The water was
changed once every two days. The temperature of the
water was about 13°C (+ 2°C).

After two days in the laboratory, the majority of the
animals spawned, perhaps in response to the higher tem-
perature of the laboratory sea water. Copulation and the
actual act of spawning were not observed. Most animals
released egg ribbons which were found floating in the
containers. In few animals, the egg ribbons were still
attached to the body and the animals appeared to be
having considerable difficulty in swimming. Instead of
smooth, swift strokes of the wings, very awkward flap-
ping movement was observed. In such cases, the egg
ribbons were carefully removed from the animals. Few
animals spawned again after about 20 days. The egg
ribbons were observed under the microscope and their
linear dimensions and the total number of eggs were
estimated. The egg ribbons were then transferred to 6-oz
finger bowls containing glass-fiber-filtered sea water. Thir-
ty mg of streptomycin sulfate per liter was added to the
water to control the bacterial growth. The finger bowls
were placed on the water table with running sea water.

After the eggs had hatched, the veliger larvae were
divided into two groups. One group was given unfiltered
sea water, while the other group was maintained in the
filtered sea water, supplemented with Platymonas sp.
Periodically, a few larvae were removed from each group
and were anesthetized partially with 1% chloretone or
1% magnesium sulphate, on a depression slide. Drawings

' Contribution No. 445 from the Department of Oceanography,
University of Washington, Seattle, Washington 98105

Vol. 10; No. 4

THE VELIGER

Page 323

were made with the aid of either a Whipple disc or a
camera lucida.

OBSERVATIONS
Adults:

Most animals were healthy for about 22 to 28 days in
the laboratory. The animals actively swam up to the sur-
face of the water in a broad spiral path and passively
sank to the bottom, by a method very similar to that of
Limacina retroversa FLEMING, 1823 (Morton, 1954).
Healthy specimens were almost transparent except for
the deep-brown gonads. Unhealthy specimens soon ac-
quired a dark-brown color around the edges of the wings
and the visceral mass, became inactive, and died.

The genus Limacina is known to bea filter-feeder (Mor-
TON, 1954). The gut contents of the specimens preserved
in the field showed a few remains of diatoms, Skeleto-
nema costatum, Chaetoceros spp., and Thalassiosira spp.;
but dinoflagellates Prorocentrum sp., Oxytoxum sp.,
Gymnodinium sp., Ceratium sp., and tintinnids were most
abundant. Unfortunately, phytoplankton samples from
the Saanich Inlet, at the time of collection of animals,
were not taken. Therefore, it is not known whether the
dominance of dinoflagellates in the food was because of
their abundance in nature or was due to selective feeding.
Skeletonema costatum was most abundant in the guts of
animals fed in the laboratory. In any case, the second
spawning indicates that the animals were getting ade-
quate food in the laboratory.

Spawn:

The eggs of Limacina helicina were embedded in a
thin, gelatinous ribbon, 3-4mm long and 1-2mm wide.
The matrix was colorless and transparent but the eggs
were slightly yellow, and were fairly closely spaced. The
egg diameter was 95-100 in the longest dimension, while
the diameter of the ovum was 75. Because of the trans-
parency and thinness of the ribbon, it was possible to
estimate the egg number fairly accurately. Five complete
egg ribbons and 4 egg-ribbon fragments were measured,
their areas were estimated and the total number of eggs
was counted. The mean egg-number per mm? was 80 and
and the total egg-number per spawn was 500 to 700
(Table 1). The second spawn contained the same number
of eggs.

Fox (1875) has described the spawns of several species
of pteropods. The eggs are generally embedded in a

Table 1

Egg-Mass of Limacina helicina Puirrs

Length Width Total Area Eggs/mm? Total Number
(mm) (mm) (mm?) of Eggs
3.72 2.05 7.61 80.3 611
4.09 2.08 8.52 83.1 708
3.42 1.71 5.86 79.4 465
3.59 1.51 5.41 92.1 498
3.66 1.79 6.54 82.8 542
2.33" 1.86 4.32 76.1
3.06F 1.24 3.80 88.8
Dssied 2.05 4.80 73.4
2.46F 1.69 4.15 78.0

Averages: 81.5 564.8

¥ Fragments of the egg-mass.

gelatinous, transparent ribbon, or in a beaded string, or
in a pea-pod-shaped case. The length of the egg ribbon
varies from 2 to 10-50mm(in Cavolinia tridentata (For-
SKAL, 1773)), while 20 to 725 eggs are produced with
each spawn. A small number of eggs per spawn is usually
associated with large-sized eggs. Limacina retroversa pro-
duces a 2mm-long egg ribbon containing approximately
300 eggs (calculated from figure 1 in Lesour, 1932).
Most pteropods have a pelagic larval stage in the life
history, but L. helicoides Jerrreys, 1877 is known to be
a viviparous species (BoNNEvIE, 1913).

Microscopic examination of the crushed gonadial tis-
sues of Limacina helicina showed ova of various diame-
ters. This observation and the second spawning, 20 days
after the first, suggest that this species apparently does
not spawn out after a short period of intensive reproduc-
tive activity. Hsato (1939) observed that L. retroversa
lays few eggs at a time and the female reproductive cells
of various ages are found in the ovotestis of the animal.
For (1875) also noted such a protracted spawning in
other species of pteropods.

Development:

No effort was made to study the details of the embryo-
genesis. For’s (1875) information on the embryogenesis
and organogenesis of pteropods has been summarized
recently by Raven (1958).

The embryos began developing immediately (Figure
1a, 1b, 1c). Toward the end of the first day, they started
actively rotating inside the egg membrane. Unlike Lima-

Page 324 THE VELIGER Vol. 10; No. 4

Figure 1

a, b, c: Stages in the development of the egg.
d, ec: Egg, just before hatching.
Scale bar is 25 u

Vol. 10; No. 4

DG

CM

THE VELIGER

\)
\
YK,

Page 325

vc

Vv

Figure 2

Seven-day-old veliger larva.
Scale har is 25

BM —- Buccal Mass
K - Kidney

CM - Columellar Muscles
O — Operculum

F — Foot

S -— Stomach V - Velum

VM - Velar Cilia

cina retroversa (LEBouR, 1932), a cap-like, symmetrical
and smooth shell developed before hatching (Figure 1d,
le). Hatching occurred at the end of the second day.
The newly hatched veliger larvae were about 75u in
diameter, with a bilobed velum, a small tongue-shaped,
ciliated foot, and a thin, long, transparent operculum.

The larvae were very active and soon after hatching
congregated near the water surface or on the lighted side
of the container. The apical mass contained all the in-
ternal organs. By the fourth day their shell diameter
increased to about 105 and the internal organs were
clearly visible. By the seventh or eighth day, the shell

Page 326

THE VELIGER

Vol. 10; No. 4

became slightly purple, and faint horizontal striations
could be recognized under proper illumination. Besides
an increase in size, no basic changes in the internal organs
were apparent. Figure 2 shows the darkly pigmented
kidney and some parts of the digestive system that were
recognizable. Larvae attained a mean diameter of 200u
by the 30" day. The foot then had enlarged, and cilia of
the velum had become stronger and longer. The opercu-
lum was still present, and larvae were still positively
phototactic. When the veliger of L. retroversa becomes
0.32mm in diameter, from the sides of the foot, two
lappets develop, which grow to form the wings of the
adult. As these lappets grow in size, the velum and the
foot diminish (LreBour, 1932). No such signs of meta-
morphosis were seen even after 30 days in L. helicina.
For lack of time, the experiment was terminated at the
end of 30 days.

There was no difference in the growth of larvae placed
in unfiltered sea water dominated by Skeletonema costa-
tum, and of those grown in the filtered sea water supple-
mented with Platymonas sp. Larvae at all stages fed well
and the food cells could be easily seen in the alimentary
canal. An effort was made to induce metamorphosis by
subjecting the larvae to a sudden temperature change.
On the 25" day, the temperature of one batch of larvae
was raised to 18°C for 12 hours and that of another was
dropped to 4°C for the same length of time. All the
larvae survived the temperature change but the process
of metamorphosis was not initiated. Prior to or after the
temperature change, no larval shell showed any signs of
vertical striations, which are characteristic of L. helicina

Variety A (McGowan, 1963).

SUMMARY

The euthecosomatous pteropod Limacina helicina pro-
duces a 3-4mm long cgg ribbon containing 500 - 700
eggs. The adults spawn morc than once and apparently
their reproductive activity is not limited to a short period.
Development of vcligcr larvac up to 30 days is described
and some observations on the food of the adult animals
are recorded.

ACKNOWLEDGMENT

This work was supported in part by U.S. Public Health
Service Grant No. WP-00633-02 to Drs. K. Banse and
G. C. Anderson, and in part by National Science Founda-
tion Grant No. GB-3360 to Drs. K. Banse and Y. Komaki.
Facilities of the Friday Harbor Laboratories were kindly
made available by Dr. R. L. Fernald. Dr. K. Banse
critically read the manuscript.

LITERATURE CITED

BonNeEvVIE, KRISTINE
1913. Pteropoda. Sci. Reprt. “Michael Sars” N. Atlantic
Deep-sea Exped. 1910, 3 (1): 1-69

Fou, H.

1875. Etudes sur le développement des mollusques. I. Sur le
développement des pteropodes. Arch. Zool. Expér. Gén.,
4 (1): 1-214

Hsiao, S.C. T.

1939. ‘The reproductive system and spermatogenesis of Lina-
cina (Spiratella) retroversa (FLEM.) . Biol. Bull. 76 (1):
7-25

Lexour, M. V.
1932. Limacina retroversa in Plymouth waters. Journ.

Mar. Biol. Assoc. U. K. 18 (1): 123 - 126
McGowan, JouN A.

1963. Geographical variation in Limacina helicina in the

North Pacific. Syst. Assoc. Publ. 5: 109 - 128
Massy, A. L.

1920. Mollusca. Part III. Eupteropoda (Pteropoda Thecoso-
mata) and Pterota (Pteropoda Gymnosomata). Brit. Ant-
arctic (“Terra Nova”) Exped. 1910. Zool. 2 (9) : 203 - 232

Morton, Joun E.

1954. The biology of Limacina retroversa.

Biol. Assoc. U. K. 33 (2) : 297 - 312
PELSENEER, PAUL

1888. | Report on the Pteropoda collected by H. M.S. “Chal-
lenger”’ during the years 1873 - 1876. Part III. Anatomy.
Sci. Reprt. Challenger, Zoology 23 (66) : 1 - 97

Raven, C. P.

1958. Morphogenesis: the analysis of molluscan development.

Pergamon Press, London, 311 pp.
Van BENEDEN, P. J.

1839. Mémoire sur la Limacina arctica.

Acad. Roy. Sci. Belles Lettres Brux. 14: 1 - 14

Journ. Mar.

Nouv. Méin.

Vol. 10; No. 4

THE VELIGER

Page 327

Marine Fouling Organisms in Monterey Harbor

E. C. HADERLIE

Department of Meteorology and Oceanography
Naval Postgraduate School, Monterey, California 93940

(Plate 49; 3 Text figures)

INTRODUCTION

THE PRESENCE of fouling and boring organisms in a
harbor is not only of interest to the marine biologist but
is of considerable economic importance to harbor engi-
neers and maintenance personnel and to boat owners. On
the Pacific Coast of North America relatively few pub-
lished studies on fouling organisms have been made.
These include investigations in Washington at Admiralty
Inlet (DE Parma, 1966) and Friday Harbor (JoHNSON &
Mirter, 1935) and in California at Oakland (GRAHAM &
Gay, 1945), Newport Harbor (ScHEER, 1945) and sev-
eral studies in the La Jolla-San Diego area (Coe, 1932;
Cor & ALLEN, 1937; ALEEM, 1957). The classic study of
boring organisms on the Pacific Coast was published by
Kororw & MILier (1927).

Until recently no systematic investigation of marine
fouling and boring organisms had been made in Monterey
Bay, California. MomMsEN (1966) and MiLter (1966)
carried out two short term studies in Monterey Harbor,
and the present paper presents the results of a full year’s
subsequent observations in the same area. The object of
the study was to obtain information concerning the kinds
of organisms that settle on or burrow into test panels
exposed in the water at various depths, to learn something
of the season of attachment or settling, to note any cor-
relation between settling of organisms and the tempera-
ture or salinity or both of the water, to determine choice
of substrate for individual organisms, to measure the rate
of growth of the dominant forms and to study seasonal
progression over the year. Other workers have found
great variability from year to year in the results obtained
from such studies, and the present investigation is planned
to continue for several years. This contribution is there-
fore to be viewed as an interim report on the first year of
study. Additional systematic observations are being made
simultaneously at various places in open water in Monte-
rey Bay down to depths of 100 feet in order to compare

results obtained with those from the harbor. These studies
will be reported in due time.

The author wishes to acknowledge the following col-
leagues for help in identification of various fouling organ-
isms: Dr. E. N. Kozloff (Suctorians), Dr. Z. M.Arnold
and Mr. Jack Gougé (Foraminiferans), Mr. G. E Kelso
(juvenile barnacles), Dr. I. A. Abbott (Algae) and Dr.
D. P Abbott (Ascidians). Mr. J. R. Lance assisted in re-
solving nomenclatural problems with opisthobranchs.
Acknowledgment is also due my wife, Mrs. A. E. Hader-
lie, for assistance in the laboratory work, and to the Office
of Naval Research for financial support.

AREA or STUDY

The site chosen for this study was near the outer end of
Monterey Municipal Wharf No. 2, about 1000 m from
the shore (Figure 1). This wharf was constructed in 1926
and is supported by concrete piles about 8 to 10 feet
apart and has rows of creosoted wooden fender piles
along each side. All of the piles are covered with a luxu-
riant growth of attached organisms dominated by acorn
barnacles [Balanus nubilis Darwin, 1854; B. glandula
Darwin, 1854; B. crenatus Brucutkre, 1789; B. tintin-
nabulum (Linnaeus, 1758) ]; anemones [Metridium se-
mle (LINNAEUS, 1767); Anthopleura xanthogrammica
(BranpT, 1835); Corynactis californica CARLGREN,
1936]; bay mussels [Mytilus edulis Linnazus, 1758];
and numerous hydroids, bryozoans, polychaetes and tuni-
cates. The depth of the water at the test site is about 21
feet at mean low tide. The maximum tidal fluctuation is
8 feet. The test site is at the outer edge of the harbor
and pollution is not excessive; the water is often so clear
that the sandy bottom can be seen easily. Throughout
the period of study surface water temperature was re-
corded and salinity determinations were made every two
weeks (Figure 2). Most of the test panels were exposed
under the eastern edge of the wharf. The water in this

Page 328

THE VELIGER

Vol. 10; No. 4

Outer Harbor

Figure 1

Map of Monterey Harbor, showing test sites under Municipal
Wharf No. 2 (test sites indicated by arrows)

area receives very little direct sunlight and then only in
the morning. One group of panels, to test for the effect
of more light, was placed on the western edge of the
wharf where the water receives direct sunlight for several
hours in the afternoon.

METHODS

In their short term studies of fouling organisms in
Monterey Harbor, both Mommsen (1966) and MILLER
(1966) tried a variety of test panels such as glass, ply-
wood, stainless steel, masonite, fiberglass, and aluminum
coated with Teflon. They both found that 4 inch douglas
fir plywood was the best all-round collector, and this

material had the added advantage of collecting wood-
boring organisms. In the present study standardized 8
inch by 10 inch panels of $ inch marine grade douglas fir
plywood were used as the primary collecting surfaces.
Some workers (PoMERAT & Weiss, 1946) have reported
that asbestos board was the most promising material for
maximum attachment of foulers. To test and compare
asbestos board with wood in Monterey Harbor one set of
collecting panels consisted of asbestos board of the same
size as the plywood panels with one smooth and one
roughened side (Johns-Manville Colorlith).

The collecting panels were supported vertically in
racks similar to those employed by Coz & ALLEN (1937)
for the last 3 years of their study. All of the submerged
racks were constructed of stainless steel (see Plate 49)
and each held six 8 inch by 10 inch panels spaced at 3
inch intervals. One rack designed to float on the surface
regardless of tidal level was made of $ inch redwood
slats. All racks were supported by a bridle and were held
in position by 3/32 inch stainless steel bathythermograph
wire. The racks were free to swing and rotate in the slight
tidal currents found at the site.

A total of six racks was employed in this study. Five
of the racks contained plywood panels, one contained
asbestos board; they were positioned as follows:

I. Floating rack. Constructed of wood and tethered
with counterweights so that it floated with the top
edge of the plywood collecting panels at the water’s
surface.

II. Intertidal rack. Positioned about 4 feet above the
lowest low tide level. The plywood panels in this
rack were submerged approximately one half the
time and exposed to air the other half.

III. Shallow rack. Located 1 foot below lowest tide lev-
el; plywood panels always submerged.

IV. Deep rack. Positioned 14 feet below lowest tide
level and about 7 feet off the bottom. Plywood
panels.

V. Deep asbestos rack. This rack contained the asbestos
board panels and was positioned at the same depth
and close to the “deep rack” containing plywood
panels.

VI. Lighted rack. This rack of plywood panels was po-
sitioned 1 foot below lowest tide level on the western
side of the wharf where the panels were exposed
to afternoon sunlight.

The period covered by this study was from October 1,
1966 to October 1, 1967. Of the six panels in each of
the racks, there were four that were left in place for 3
months (Panel 3P), 6 months (Panel 6P), 9 months
(Panel 9P) and 12 months (Panel 12P) respectively.
These were designated as Long Term panels. The re-

Vol. 10; No. 4

THE VELIGER

Page 329

700
35.0

34.6
34.2
33.8

33-4

SS SSSA Se ees

Figure 2

Biweekly morning temperatures and salinities at test site for the year October 1, 1966 to October 1, 1967

maining two panels in each rack were designated Short
Term panels (T panels): one was put in on the first of
each month and retrieved for study on the first of the
next month; the second was put in on the fifteenth of
each month and retrieved a month later. These short term
panels therefore overlapped one another and made pos-
sible more exact determination of settling time. A total
of 162 panels were exposed and examined during the
year. Two of the panels from the 3 month series were
lost. The length of exposure of the various panels in
each rack is shown diagrammatically in Figure 3.

In many of the previous studies on fouling organisms
on this coast, investigators have found it expedient to

scrape the fouling organisms off the panels and preserve
the entire lot in formalin or alcohol for later sorting and
identification. In a preliminary pilot study carried out
two years ago in Monterey Harbor this technique proved
to make identification very difficult and often impossible,
especially for small, delicate, recently-settled forms.
Throughout this study, therefore, the retrieved panels
were placed in tubs of sea water or in aquaria with
running sea water immediately after removal from the
racks and were examined as soon as possible, usually
within 3 hours of being removed from the sea. For
examination each panel was placed in a pan of sea water
and the entire surface surveyed with a binocular stereo-

Page 330

THE VELIGER

Vol. 10; No. 4

Interval | Interval of Expose | Interval of Expose

Cee eee
| fot [o [africa tal s[ spats:

Figure 3

Diagram illustrating period of exposure for test panels
from October 1, 1966 to October 1, 1967

T=Short term panels P=Long term panels

scopic microscope (7x to 30x magnification). This made
it possible to see and record very small living organisms
such as some of the protozoans and newly settled stages
of bryozoans and barnacles. This technique not only
made identification easier but made it possible to observe

the behavior of the organisms, determine where on the
panel they tended to settle, their size, relative abundance,
what their associates were, etc. Proper identification of
the fouling organisms, especially the very young stages,
was constantly a problem, but with experience and the
help of specialists it became easier. In some cases panels
with juvenile organisms of questionable identity were
kept in aquaria or were returned to the original racks for
additional periods during which time the organisms in
question matured enough to make identification possible.
Representative organisms were then removed from the
panel and preserved for later study and reference. The
entire wooden panel of the short term series was air dried
and retained for reference. In the case of the panels in
the long term series that had been in the water from three
months up to a full year, a careful study was made to
determine which organisms were dominant, their size and
shape and the percentage of the surface each species
covered. After examination, one of the flat surfaces of
each of these panels was scraped clean of attached or-
ganisms and the scrapings were oven dried at 100°C
until the weight was constant. The results are expressed as
grams of dry weight tissue per panel side (80 square
inches or 5122 cm’). This provides a rough statistical
measure of concentration and an index to productivity
of the environment for the specific time period of expo-
sure (see bottom line Table 3). In many cases, however,
the older panels had obviously suffered loss of part of the
fouling growth due to periodic buffeting of the racks
against piles when wave action was severe. The panel with
the remaining organisms intact on one side was air dried
and saved for reference.

THE FOULING COMMUNITY

I. Discussion of Organisms
Settling on Short Term Panels

During the year of observation a total of 70 different
kinds of organisms, identified at least to genus, were
found on the short term panels. Some organisms were
found to have settled during every month of the year,
indicating either a continuous breeding cycle or larval
forms that can live free for some time before settling.
Others settled only seasonally. The truly sessile organisms
(primary foulers) received the most attention and a
serious attempt was made to identify to species all such
forms, but many of the more easily identified motile
organisms (secondary foulers) that were found on the
panels, were also recorded. Numbers of diatoms, tiny
flatworms, nematodes, small errant polychaetes, minute
snails, mites, free-swimming copepods and crustacean

Vol. 10; No. 4

THE VELIGER

Page 331

larval forms were often observed but few of these were
identified.

Table 1 is an example of how the data were collected
and recorded for one common fouling organism, Balanus
crenatus BrucutkrE, 1789. Similar tables were prepared
for each of the 70 organisms. For bryozoans, hydroids,
etc., numbers of discrete colonies were counted rather
than individuals.

Table 1

The number of barnacles (Balanus crenatus) that settled on
various panels throughout the year. A table similar to this was
prepared for each of the 70 species encountered

ye 8g 85 Fs Ss .% oa
fa] 1 Gl ey el 5 é¢

ag 8G 22 82 Ba Be 38

(aT 20

QT iL 2

eu 1

4-T 1 2

=a Terie” 63

6-T 1 2

qa 1-3

8-T te 2 1 4

9-T 4 4

10-T 30 150 100 160 6

ils Tees 2one20) Bue, 10) lS

en J WM 5 8

13-T 3 50 2

14-T 4 12 WM 2

(G70 2) eM Scere

16-T OK 1 9a

Ves 1 HDS 2

=e 20 10

19-T 5 iC

20-T 9 DD) 8 GS aD

la 2 BW) 1. 9 M0) Glen

eam Tee Oe 10 10

Bot 6 15 10

Total
panels fouled: 9 20 18 14 11 11

The data from the entire observation period are sum-
marized in Table 2. This table lists the identified organ-
isms, the numbers of individual test panels where the
organisms settled, and the period of the year when the
organisms occupied the panels. Also indicated are those
panels that carried the greatest number of individuals
or colonies of organisms, and at the bottom of the table
the number of species collected by each panel.

As can be seen from an examination of Table 2, the
most common fouling organisms encountered on the short
term panels were encrusting cheilostomate and cyclosto-
mate bryozoans, calcareous serpulid worms, the acorn
barnacle Balanus crenatus, foraminiferans, ciliates, the
nestling clam Hiatella arctica and the opisthobranch Her-
missenda crassicornis. The following discussion will bricfly
review the most common organisms of each of the major
groups listed in Table 2.

Algae:

None of the short term panels ever had an extensive
growth of algae. After one month in the water all panels
usually had a slippery surface film that probably consisted
of microorganisms too small to be seen, and on many of
the panels in racks near the surface, chains of colonial
diatoms were observed but not identified. On the rack
suspended just below tide level on the western side of
the wharf (the lighted rack) where afternoon light was
greatest, it was expected that the panels would collect
algae and during the summer months small sprigs of
green and red algae were often found. Ulva sp., Polysi-
phonia pacifica and Herposiphonia parva were the only
plants identified. In all cases the growth was very sparse.
On one panel, numerous fans of the colonial diatom
Licmophora sp. were observed.

Protozoa:

Three species of foraminiferans were observed on vari-
ous panels. The most common by far was Cornuspira
lajollaensis. It was found throughout the year on panels
at all depths and positions, but was especially abundant
on the intertidal and floating panels. The next most
abundant protozoan was the large pink or reddish ciliate
Folliculina sp. which again was found throughout the
year and often in such numbers as to make the panel
look dark to the naked eye. The suctorian Ephelota gem-
mipara was sometimes seen as a small forest on the sur-
face of the panels and especially as a fuzzy growth on the
clam Hiatella arctica and on hydroids. Ciliated swarmers
were seen leaving Ephelota during February. The stalked
colonial ciliate Zoothamnium sp. was fairly common and
was most clearly seen on the roughened edge of the
panels.

Porifera:

The only sponge that was common on the short term
panels was Leucoselenia eleanor. In the time available
this sponge never was able to form dense masses of
anastomosing tubes, but consisted of a few short tubes

Page 332 THE VELIGER Vol. 10; No. 4
Table 2
List of Species, and of Panels where Organisms Settled in Short Term Series
Number of test panels on which | Total panels | Interval test panels were occupied
Species organisms settled! occupied | (October, 1 1966 - October 1, 1967)
aS DE vA

Algae:

Licmophora sp. 1 September

Ulva sp. 8 January - October

Polysiphonia pacifica HOLLENBERG, 1942 4 July - September

Herposiphonia parva HOLLENBERG, 1943 4 July - September
Protozoa:

Cornuspira lajollaensis Ucuto, 1960 11 October - October

Rosalina columbiensis (CusuMan, 1925) 2 June - October

Spirillina revertens (RHUMBLER, 1906) 2 July - September

Zoothamnium sp. 7 October - October

Folliculina sp. 15 October - October

Vorticella sp. 0 September

Ephelota gemmipara (Hartwic, 1876) 1 October - October

Porifera:

Leuconia heathi (Urpan, 1905)
Leucosolenia eleanor Ursan, 1905
Coelenterata:

Obelia spp.

Hydractinia sp.

Syncoryne mirabilis (Acassiz, 1862)
Campanularia sp.

Anthopleura sp.
Platyhelminthes:

Stylochoplana gracilis HEATH & McGrecor, 1912
Notoplana acticola (Boone, 1929)
Thysanozoon californicum Hyman, 1953
Stylochus tripartitis Hyman, 1953
Stylochus sp.
Nemertea:

Emplectonema gracile (JoHNSTON, 1837)
Ectoprocta (Bryozoa):

Bowerbankia gracilis O DONOGHUE, 1926
Filicrisia franciscana (RoperTsoNn, 1910)
Crisia sp.

Crisulipora occidentalis Ropertson, 1910
Tubulipora pacifica Ropertson, 1910
Bugula neritina (LinNAEvs, 1758)
Bugula californica Ropertson, 1905
Hippothoa hyalina (Linnaeus, 1767)
Lyrula hippocrepis (Hincxs, 1882)
Scrupocellaria californica TRasK, 1857
Scruparia ambigua (p’Orsicny, 1841)
Holoporella brunnea (Hincks, 1884)
Entoprocta:

Barenisia gracilis (Lomax, 1886)
Annelida:

Nereis sp.

Platynereis agassizi (EHLERS, 1868)
Pectinaria californiensis HartTMAN, 1941
Halosydna brevisetosa KinBErc, 1855

ao

9 5 9 0 1 1
0 1 1 0 0 0 2
5 7 8 3 0 1 24
1 0 0 0 0 0 1
0 0 0 1 1 0 2
7 7 4 5 9 i 44
0 2 1 0 1 6 10
1 0 1 1 1 0 4
0 0 2 0 0 0 2
1 0 1 0 0 0 2
0 0 3 0 0 0 3
© i Ww 7 6 9 56
4 1 1 4 1 4 15
1 0 0 0 0 1 2
1 1 1 10 7 2 22
17 2 2 22 wy 2 83
8 4 1 1G. al 9 49
4 0 0 3 4 6 17
Ml 310 1 Bi 2B we 106
13 8 8 12 3 i) 65
0 0 0 0 0 2 2
17 2 i al 54
13 7 4p je) ies) 76

Oo
oO
Oo
oO
Oo
=
_

or O°
- OO =
=- © © ©
ooro
nooo
- Ont OC
are oe

January - July
February - October

October - October
May

November - October
July

November - December

October - October

February - October

March; May - June; September
June

March; July

March - May

January - October
May - October

June - August
November - October
October - October
January - October
November - January; May - October
October - October
November - October
May

October - October
November - October

January

August

April - July

August

March; August - October

Vol. 10; No. 4

THE VELIGER

Page 333

Table 2 [Continued]

Species
Lawn
Annelida (continued)
Sabellaria cementarium Moore, 1906
Spirorbis spirillum (Linnaeus, 1758)
Serpula sp.
Polydora brachycephala Hartman, 1936
Thelepus sp.
Arthropoda:
Balanus crenatus BRuGUIERE, 1789
Balanus glandula Darwin, 1854
Chthamalus sp.
Idothea sp.
Limnoria quadripunctata Hottuuts, 1949
Caprella sp.
Corophium insidiosum Crawrorp, 1937

Nh
SN a a)
S) I= aS es) ©)
(o) le (e8) (te) (>)

ouwmnoro vo
=
UuUnoorr,.&

Mollusca:

Pododesmus cepio (Gray, 1850)

Mytilus edulis Linnagus, 1758

Pecten sp.

Hiatella arctica (LinNaEus, 1771)

Bankia setacea (TRYon, 1863)

Hermissenda crassicornis (ESCHSCHOLTZz, 1831) i
Doto kya Marcus, 1961

Dendronotus frondosus (Ascantus, 1774)
Acanthodoris brunnea MacFaruanp, 1905

Triopha grandis MacFaruanp, 1905

Cratena albocrusta MAcFaRLAND, 1966

Polycera atra MacFarxanp, 1905

Acmaea sp.

Mopalia sp.

Echinodermata:

Strongylocentrotus sp.

Eupentacta quinquesemita (SELENKA, 1867) 0
Pisaster sp. 1

=

OCOWrFrFOOWNONONO Ur
—_

—
Orr OoOrFWOrFrONOCOYWHO CO

COrFNNOOCOOCOCOOD KY NN

We)
ooo
oo ©

Total species collected by each type panel: aks) Geil ae

organisms scttled'

S250 Di A
0 1 0 1 March
2, 2S UP 105 October - October
1} 118} 59 October - October
1 5 11 February - August
0 0 1 1 August
gh IT alah 83 October - October
0 0 0 2 June; August
1 0 0 4 July - September
0 1 0 2 February; May
19 10 0 60 October - October
6 3 5 26 November; February; May - October
0 0 1 6 March - July; August
5 3 1 12 October; July - September
1 2 0 19 October; January; July - September
0 0 2 3 July - September
it 113} 9 63 October - October
2 0 0 2 February - April
13 1@ 68 October - October
0 0 0 1 May
1 0 1 8 January - July
0 0 0 il June
2 0 0 4 December; February; April
0 3 0 6 January; May; August
2 1 0 6 February - August
0 0 0 4 March - July
1 0 0 1 May
7 8 8 49 January
0 0 1 1 June
1 2 2 6 June - September
39° 41 41

‘Panel designations are as follows: L=Lighted Panel

I=Intertidal Panel

F=Floating Panel

S=Shallow Panel
D=Deep Panel
A=<Asbestos Panel

2 Bold face figures indicate the group of panels with the greatest number of individuals or colonies

tightly pressed on the panel surface. A few small speci-
mens of Leuconia heathi were occasionally observed,
especially on the deep panels.

Coelenterata:

In most fouling studies, investigators have reported
hydroids as among the most common organisms. This
was not found to be true under the Monterey wharf.
Obelia sp. was found throughout the year, but was usually
a very sparse growth except for a few of the intertidal

panels where it formed a flourishing colony with stalks
2cm high. Maximum growth was in May, and medusae
were released at that time. The only other hydroid found
in any significant number was Syncoryne mirabilis and it
was found as a very sparse growth except on a few
floating panels in June. Small anemones (Anthopleura
sp.) were found on only two occasions on the short term
panels. It is surprising that more anemones did not settle
on the panels, for several species are found growing in
great numbers on nearby piles.

Page 334

TE MVEEIGER

Vol. 10; No. 4

Platyhelminthes:

Flatworms were common secondary foulers on many
of the panels, but some of the worms were so minute as
to make identification impossible. The most consistent
and conspicuous form was the small cuneate Stylocho-
plana gracilis which was present all year. It was abundant
on the deep panels, especially those of asbestos board,
and was always seen gliding in and among the bryozoans
and barnacles. An interesting flatworm encountered only
rarely was Thysanozoon californicum. This small worm
has a papillated dorsal surface and its color and texture
often closely matched the bryozoan colonies (Holoporella
brunnea) among which it lived.

Nemertea:

Small specimens of Emplectonema gracile were found
on two panels in the floating rack in March and April.
This was the only nemertean found on the short term
panels.

Ectoprocta (Bryozoa):

During the period of this study, bryozoans clearly
constituted the dominant fouling organisms on the short
term panels in respect to numbers of species involved,
number of panels fouled and numbers of individual colo-
nies. The single species that was dominant on all the
submerged panels throughout the year was Hippothoa
hyalina. This organism forms a circular colony that is
hyaline and appears as delicate silvery patches on test
surfaces. On many of the shallow and deep panels there
were hundreds of colonies up to 5.0mm in diameter on
each side of the pancl. Next in importance was Tubuli-
pora pacifica. This cyclostomate bryozoan forms a colony
that when small is a fan-shaped mat of calcareous tubes
with the open ends pointing upward. On the short term
panels these colonics never were more than 1 -2mm
in diameter, but on deep wooden and asbestos panels in
June there were hundreds of colonics which made the
test surface feel rough like sandpaper. Tubulipora was
found on the floating and intertidal panels only in Au-
gust and September, but on others it occurred throughout
the year. Two other encrusting bryozoans that were a
conspicuous part of the fouling community throughout
most of the year were Lyrula hippocrepis and Holo-
porella brunnea. These colonies were often colored yellow
or orange and after being on the panel one month or less
were up to 4 or 5mm across. One large Lyrula colony,
however, attained a diameter of 10mm in a month or
less.

Erect bryozoans were less common on the short term

panels than the encrusting forms, yet many of these had
formed small colonies. Scruparia ambigua was never
abundant on any one panel, but it was found throughout
the year on all submerged panels. The site of growth of
Scruparia was most often on the roughened edge of the
panel and it was never more than a few millimeters high.
Bowerbankia gracilis was found from January through
September and was represented on panels at all depths.
The creeping stolons of Bowerbankia often gave rise to
only a few zooecia, but in some cases on intertidal panels
in February and again from May to July a thick furry
coating was found. Bugula neritina is easily recognized by
its purplish color, and was found as small sprigs from
January to October. Crisulipora occidentalis was the only
other bryozoan that occurred with any regularity. It was
found throughout most of the year on many of the sub-
merged panels, but was most common on the shallow
ones.

Entoprocta:

The only entoproct observed during the year of study
was Barentsia gracilis; it was seen as a small colony only
once on an asbestos panel in February.

Annelida:

Spirorbis spirillum was second only to the bryozoan
Hippothoa hyalina in number of panels fouled and it
settled on the surfaces throughout the year. It was least
common on the floating and intertidal panels, but was
abundant on all of the submerged ones, both wood and
asbestos. On some panels, over 50 tubes per side were not
uncommon. Most of the tubes were tightly spiraled and
averaged 0.5mm in diameter but some were up to
1.0mm. Both sinistral and dextral tubes in about equal
numbers were represented. The twisted but not coiled
tubes of another serpulid (Serpula sp.) were also found
throughout the year, but these were less abundant than
Spirorbis, both as individuals and in terms of number of
panels settled on. The maximum growth of Serpula sp.
in one month was up to 6mm in tube length. The only
other annelid that occurred regularly was the tube-
building Polydora brachycephala. The tubes built of silt
and debris were fairly common, especially on the asbestos
panels. Many tiny errant polychaetes were observed on
nearly every test surface, but none of these were identi-
fied.

Arthropoda:

As was to be expected, the dominant arthropod foulers
were acorn barnacles. On the piles of the wharf Balanus

Vol. 10; No. 4

glandula, B. crenatus, B. tintinnabulum and B. nubilis are
common, yet, except for a few specimens of B. glandula
and Chthamalus sp., B. crenatus was the only barnacle
that was found as a significant fouler on the test surfaces.
Throughout the entire year and on panels at all clepths,
small specimens of B. crenatus could be found. Many
were recently settled forms 0.1 mm across, but others were
up to 2.5mm after being on the panel one month or less.
As a rule any one panel had only a few barnacles, but
enough to show that B. crenatus settles throughout the
year (see Table 1). The period of maximum scttling was
from mid-February through March. This was also found
by Mommsen (1966) to be the period of maximum
settling of B. crenatus. It is interesting to note that during
late February and carly March the maximum settling of
up to 25 barnacles per square inch was on the floating,
shallow and deep wooden panels, but in late March the
maximum settling was on the intertidal panels. In late
May and early June another period of intense settling of
B. crenatus occurred on the floating pancls and in late
July and August a heavy scttling on the shallow panels.

In a survey in the same area during the previous sum-
mer, Mitter (1966) found massive settling (up to 50
per square inch) of Balanus crenatus in late June and
early July, and the deep panels were most heavily fouled.
In contrast to this, the present study found practically no
barnacles settling on the deep panels during late June
and July, and only a few on the intertidal and floating
panels. The explanation of this might be found in differ-
ences in water temperature. In MILier’s study the tem-
perature fluctuated between 12.5°C and 14.0° C during
this period, whereas in 1967 the temperature varied be-
tween 13.5° C and 15.8° C (Figure 2).

In terms of choice of color of substrate, no significant
differences were found in the rate or season of scttling of
Balanus crenatus on the dark asbestos panels and the
lighter wooden panels at 14 feet depth. On the upper
panels subject to more light, however, it was found that
the larvae tended to settle most abundantly on darker
parts of the wood and on dark lines drawn on the wood.

As will be noted later, Balanus tintinnabulum scttled
on the long term panels and on the stainless stecl racks,
but this barnacle was never scen on the short term panels.
Perhaps this species, like some other fouling organisms,
requires the presence of living organisms on a surface
before it will settle.

Other arthropods recorded on the short term pancls
were secondary foulers. Various copepods were found on
practically every test surface, but none were identified.
The skeleton shrimp, Caprella sp., and the tube-building
amphipod Corophium insidiosum were occasionally pres-
ent as was the isopod Idothea sp. Mites and crustacean

THE VELIGER

Page 335

larvae were common and abundant and the wood boring
gribble Limnoria quadripunctata was often seen crawling
over the surface or in shallow burrows in the wood.

Mollusca:

The attached mollusk most often encountered on the
short term panels was the nestling clam Hiatella arctica.
This white, rectangular clam was found throughout the
year on panels at all depths, but the greatest number of
individuals was on the floating panels. Most of the ani-
mals were very small (2mm and less) and were usually
seen crecping over the test surface, but larger clams
(10mm and more) were often attached by byssal threads.
The clams were most common in depressions in the wood
surfaces.

Small numbers of the bay mussel, A/ytilus edulis, were
also found on wooden panels at all depths throughout
most of the year. Most of these were small and mobile,
but others were up to 10mm long and attached. Never
were there more than 4 or 5 animals on a panel. Podo-
desmus cepio was also found attached to or moving over
the pancls. These were invariably very small.

Opisthobranchs were encountered on the test surfaces
throughout the year. By far the most common and abun-
dant species was Hermissenda crassicornis. Each panel
occupied normally carried from 1 to 5 animals, and egg
masses of Hermissenda were found during every month
of the year. Dendronotus frondosus was sccn only on a
few panels, but when seen it was very abundant. Cratena
albocrusta and Polycera atra, although not scen often,
were abundant on certain panels. Other opisthobranchs
listed in Table 2 were found only rarely. Except for
Hermussenda crassicornis, which appeared on panels re-
gardless of what othcr organisms were present, most of
the nudibranchs found were on panels which supported
colonics of Obelia or other hydroids.

Other mollusks encountered on the temporary pancls
were gastropods of a varicty of forms but invariably
small and none were identified. Also found one or more
times were tiny limpets (Acmaca sp.) and very small

chitons (Mopalia sp.).
Echinodermata:

From January through September, small sea urchins
(Strongylocentrotus sp.) averaging 0.5mm in diameter
were frequently encountered in small numbers on pancls
at all depths. Tiny asteroids (Pisaster sp.) appeared on
a few panels from June to September and a single speci-
men of the sea cucumber Eupentacta quinquesemita was
seen in June.

Page 336

Chordata (Tunicata) :

As will be seen later, 3 species of tunicates settled on
the long term panels and on the stainless steel and wooden
racks. Yet, no tunicates were ever observed on the short
term panels. It is possible that these animals are not
pioneers in the fouling community, but settle only after
the surface has been “prepared” by other organisms.

II. Depth and Surface Preference

At the bottom of Table 2 is a summation of the total
number of species of fouling and boring organisms collec-
ted by each category of test panel. There are only slight
differences in the total number of species each collected.
As has been pointed out in the above discussion of the
organisms that settled on the short term panels, some def-
initely preferred one depth to another, but the preference
varied from species to species. One of the purposes of this
study was to determine the type of surface that would
collect the maximum number of fouling organisms, and
to get data relative to this problem, panels of 2 different
materials (plywood and asbestos board) were submerged
to identical depths in the same area to test their relative
effectiveness as collectors. As can be seen from Table 2,
there was no difference in the number of species col-
lected by these different materials, and in settling most
foulers did not seem to distinguish between the two.
Some organisms, however (e. g. Folliculina sp.), seemed
to prefer plywood to asbestos board, whereas the reverse
was true for others (e.g. Lyrula hippocrepis). The as-
bestos board had one smooth and one rough surface but
these differences had no effect on rate of settling of
foulers except that there was a tendency for more barna-
cles (Balanus crenatus) to settle on the rough surface than
on the smooth, and encrusting bryozoa seemed to prefer
the smooth surface. These differences, however, were
only slight.

III. Discussion of Organisms
Settling on Long Term Panels

Table 3 lists the fouling and boring organisms observed
on the long term panels. As can be seen, the list is
similar to that of the organisms on the short term
panels (Table 2). It is a longer list, however, containing
81 kinds of organisms identified at least to genus, and
several organisms were found in fair numbers on the long
term panels that were never encountered on panels in
the water for only one month. As SHEER (1945) and
others have pointed out, this is to be expected, for many

THE VELIGER

Vol. 10; No. 4

organisms settle on surfaces only when these surfaces
have been colonized by pioneering foulers.

The long term panels presented many difficulties in
analysis. Very often one or two species of bryozoans, for
example, were the dominant organisms at the time the
panels were removed for study, and these had overgrown
and completely smothered earlier populations of barna-
cles and serpulid worms. In the case of panels practically
covered with a foliacious growth of erect bryozoans, it
was impossible to locate all the organisms living in and
around the bryozoans. It was obvious that very often the
growth of organisms had become so heavy on a panel
that large slabs of fouling growth had broken off and the
area was being recolonized by other forms. This explains
why the weight of the fouling mass on a 9-month panel
might be less than on one of 6 months (see Table 3,
bottom). As before, each of the major groups and domi-
nant organisms will be discussed briefly.

Algae:

Five species of green and red algae were found on the
long term panels in the lighted rack. Even after 12
months the growth of algae was very sparse and did not
constitute a dominant part of the fouling mass. Diatoms
were observed attached to bryozoans and other fouling
animals but none of these were identified.

Protozoa:

It was very difficult to detect protozoans in the mass
of multicellular foulers, yet essentially the same forms
were found as on the short term series. Cornuspira lajol-
laensis was the dominant foraminiferan, and on the 12-
month floating panel was found in thousands. Folliculina
sp. was the most commonly encountered ciliate.

Porifera:

On the long term panels Leucosolenia eleanor appeared
as large masses of anastomosing tubes, and was often
covered with silt and debris. At one time or another it
was found on panels at all depths and was also abundant
on the floating and intertidal racks. Leuconia heath
was a common sponge on the submerged panels and
racks but was not found on those in the intertidal or the
floating series. On some of the 12-month panels single
specimens of this sponge were often up to 8cm long, but
most were globular and about 3cm in diameter. Rhabdo-
dermella nuttingi was found only on 9-month and 12-
month test surfaces. It was never abundant, and was
found in small clumps, the individuals reaching a maxi-
mum length of 5cm.

Tue VELIGER, Vol. 10, No. 4 [HADERLIE] Plate 49

wR
AN

\\

aN \

Photograph of test racks and panels

Left, rack and 6 panels in place ready for exposure. Right, deep

rack following removal from water after 12 months exposure.

The 12 month panel and the last temporary panel (23-T) still in
place. Fouling growth primarily Crisulipora occidentalis.

Vol. 10; No. 4

Coelenterata:

The only consistent hydroid fouler on the long term
panels was Obelia sp. It was most abundant on the inter-
tidal panels, and on the deep asbestos panels. The ane-
mone Corynactis californica was found in small numbers
on one asbestos panel at 9 months and on one deep wood
panel at 12 months and was found attached to the wood
bottom plates of the intertidal, shallow and deep racks.
This anemone is common on the wharf pilings, but was
never encountered except in these few instances cited.

Platyhelminthes:

Essentially the same group of flatworms inhabited the
long term and short term panels. Much larger worms,
however, especially Stylochus tripartitis, were found on
the long term surfaces, and in all cases the flatworms
were more abundant on the older, more heavily fouled
surfaces.

Nemertea:

The three species of nemerteans found on the long
term series were few in number but large in size. One
Emplectonema gracile attained a length of over 30cm.
These worms often occupied open burrows that had been
excavated by Limnoria, or dead shells of barnacles.

Ectoprocta (Bryozoa) :

The bryozoans again were the dominant organisms on
all of the long term panels and on most of the racks. On
the submerged panels and racks Crisulipora occidentalis
was often so dominant as to make the surface appear to
be covered with nothing else (see Plate 49). The thick-
ness of the Crisulipora growth often exceeded 5cm, and
on the deep panels this growth made up more than three
fourths of the weight of the entire fouling community.
Crisia sp., Filicrisia franciscana and Bugula neritina of-
ten grew abundantly with Crisulipora, but never were
any of these dominant. Under the Crisulipora growth,
and encrusting the surface of all submerged panels of 6
months age and older and all racks was Lyrula hippocre-
pis. Holoporella brunnea and massive fans of Tubulipora
pacifica were also found in abundance. Lyrula was in all
cases, however, the dominant encrusting bryozoan.

On the floating panels and those in the intertidal po-
sition, Crisulipora was usually completely replaced by
Bowerbankia gracilis as the dominant erect bryozoan,
and many panels and the racks in these positions had a
thick furry growth of Bowerbankia. Under this blanket,
however, was the ubiquitous Lyrula hippocrepis and,
except on the intertidal and floating panels, Holoporella

THE VELIGER

Page 337

brunnea. The encrusting bryozoan Hippothoa hyalina,
the most conspicuous organism on practically all of the
short term panels, was found on a few panels of the 3-
month and 6-month series, but after this was not seen
on the panels as a living organism. The dead remains of
Hippothoa were often seen on 9- and 12-month panels
under a smothering encrustation of Lyrula hippocrepis.

Annelida:

The dominant annelids were again, as before, the ser-
pulid worms Spirorbis spirillum and Serpula sp. They
were found on all panels except those in the intertidal
series, but on the older panels were often smothered by
Lyrula growth. On some of the 12-month panels, Serpula
sp. attained a length of 8cm and up to 0.5cm in dia-
meter.

Arthropoda:

Balanus crenatus was fairly common on only the inter-
tidal panels at 3 months, but at 6 months this barnacle
was very abundant on all except the deep wooden and
asbestos panels. From then on, as the bryozoan growth
increased, the barnacles declined. Under the layer of
Lyrula hippocrepis could be seen the dead shells of many
B. crenatus. Balanus tintinnabulum was never abundant,
but on many of the panels except the deep ones from 1
to 5 large B. tintinnabulum could be found. The largest
of these from the 12-month panels were 3 cm in diameter.
A small number of B. glandula was found on the 12-
month panels of the intertidal series, and a few were also
observed on the stainless steel rack of the intertidal
series and on the wooden floating rack.

Mollusca:

The dominant mollusk on the long term panels was,
as on the short term ones, the nudibranch Hermissenda
crassicornis. The dominant attached mollusk was Hiatella
arctica. This nestling clam was found embedded in the
fouling growth of panels at all depths, but was most
abundant and attained the largest size on the floating
panels. On some of these, and in crevices in the floating
rack, after 12 months this mollusk reached a length of
3cm and had siphons up to 5cm long.

Mytilus edulis is a common wharf piling dweller yet
is was found only in small numbers on the test panels and
racks during this study. Never were more than 5 mussels
found on any one panel, but they were sometimes of
large size. Several mussels attached to the floating rack
were 5cm long when the rack was removed after 12
months.

Page 338 THE VELIGER Vol. 10; No. 4

Table 3

List of Species, and of Panels where Organisms Settled in Long Term Series
October 1, 1966 to October 1, 1967

6 months 9 months 12 months
L IF S D A‘! LI F S D-A EVE eSseaDRA

3 months
leer SmeDaeAN

Algae:

Ulva sp. 2

Giffordia sandriana (ZANARDINI) HaMEL, 1939

Polysiphonia pacifica HoLLENBERG, 1942 72

Pterosiphonia dendroidea (MONTAGNE) 1

FALKENBERG, 1901

Brachioglossum woodii (AcaRDH) Ky.in, 1924 1
Protozoa:

Cornuspira lajollaensis Ucuio, 1960 ND. oD) 2 2 2 2 2 8) Be @ 2
Rosalina columbiensis (CusHMAN, 1925) ND ND, 2B 2
Zoothamnium sp. ND ae 2 74 ee
Folliculina sp. ND, wy 2) @ @ 3 3 3 3 2
Ephelota gemmipara (Hartwic, 1876) SD) }y BD 3
Porifera:

Leuconia heathi (Urpan, 1905) ND mil il 2 il By i 2 3) 8 2
Leucosolenia eleanor Ursan, 1905 NP) NONE liars anita 2 1 ie Ga ae | L cles tees 2:
Rhabdodermella nuttingi Ursan, 1902 ND nD 1 1 aa 1
Coelenterata:

Obelia spp. Be 2 g)) BD 2 BB e 3 2 1
Syncoryne mirabilis (Acassiz, 1862) oe) uD 1 1
Hydractinia sp. ND xD 1
Corynactis californica CARLGREN, 1936 ND np 1 1
Platyhelminthes:

Stylochoplana gracilis HEATH & McGrecor, 1912 | %? BD 1 1 1 1 1
Notoplana acticola (Boone, 1929) AD a) AM ab | 1
Thysanozoon californicum Hyman, 1953 ND ND 1

Stylochus tripartitis Hyman, 1953 BD BD) >?
Nemertea:

Micrura verrilli Cor, 1901 aD oy 1 1 1
Emplectonema gracile (JoHNSTON, 1837) aD) aD 1 1 1
Tetrastemma nigrifrons Cor, 1904 ND, ND i i al 1 1
Ectoprocta (Bryozoa) :

Bowerbankia gracilis ODonocuHuE, 1926 ND ND 3 3 3 B33 312 3 2 2 2
Filicrisia franciscana (Ropertson, 1910) aw) Bw) 1 1 1 it ih yl 1s
Crisia sp. ND xP tor 1 1 1
Crisulipora occidentalis Ropertson, 1910 ND me) 2 33) 313) 3 3°38 8 | 3 eo Be 3
Tubulipora pacifica Ropertson, 1910 sD m2 3 8) 3 2 2 3 2 ai 2 2) 22 Be?
Bugula neritina (LinnaEus, 1758) ND mm jt 1 fey it i il 1 2k leat
Bugula californica Ropertson, 1905 xD sil sh al yi Aga 1 1
Hippothoa hyalina (Linnaeus, 1767) xD, ND 2 2

Lyrula hippocrepis (Hincxs, 1882) aD eee 8) Ss A PSs B Gild BoB 8 3B FZ
Scrupocellaria californica Trask, 1857 BD 3D 2 1 iL | 2 242:
Scruparia ambigua (p’Orsicny, 1841) aD SD 1 1 1 1 1 1
Holoporella brunnea (Hincxs, 1884) ND D3 2 Qi 8 3 2 8] 2 2 || 3 2 2
Annelida:

Nereis vexillosa Grupe, 1851 wD aD 1

Nereis sp. uD SD el
Platynereis agassizi (EHLERS, 1868) ND ND 2

Halosydna brevisetosa KinBERG, 1855 ND ND 1 3 3

Vol. 10; No. 4 THE VELIGER Page 339
Table 3 [Continued }
3 months 6 months 9 months 12 months
El BS DAE 1 Bb Ss DA Lrirs DA
Annelida (continued)
Pseudopotamilla ocellata Moore, 1905 ND ND 1 1
Sabella sp. ND xB i) Al
Spirorbis spirillum (Linnaeus, 1758) Be) mB 2B) 2 1 Pea 2 2 Bl 2 2 2B 2
Serpula sp. ND, D2 2.2 2 bine 2 2B D\ 2 2
Polydora brachycephala Hartman, 1936 ND) aD 1 1 3°38 8 2 2 1 1
Arthropoda:
Balanus crenatus BRuGuIERE, 1789 NOMS ND Wee Ss MS iet Be A 3 2 2p 3 3
Balanus glandula Darwin, 1854 sD nD 1
Balanus tintinnabulum (Linnaeus, 1758) ND ND ak al 1 1 fe elit
Chthamalus sp. ND ND 1 1
Limnoria quadripunctata Hottuuts, 1949 ND Il ehDant open! 3 93 3 3 2 33 3 3 38 3 3 8
Corophium insidiosum Crawrorp, 1937 ND ND 2
Caprella sp. ND a) 3 2 Zee
Cancer antennarius Stimpson, 1856 xD Bw) 1 1
Pachygrapsus crassipes RANDALL, 1839 ND uD 1
Pugettia producta (RANDALL, 1839) ND ob) 1
Loxorhynchus crispatus Stimpson, 1875 MD, ND 1 1
Mollusca:
Pododesmus cepio (Gray, 1850) ND ND 2 yD
Mytilus edulis LinNAtus, 1758 AD ap) 1 til 1 il 2
Pecten sp. BB By) 1 1
Hiatella arctica (LinNAEus, 1771) No BD 2 3 2 2 PB 273, 2 2 2 D 2 @
Bankia setacea (Tryon, 1863) SD ND 1 i Gl al 1
Hermissenda crassicornis (EScHscHOLTz, 1831) | 1 ™ 1 1 1] 1 1 241421 «41 1 il il Iola Il
Doto kya Marcus, 1961 a i BB
Dendronotus frondosus (Ascantus, 1774) a) ND) 1 1 1
Acanthodoris brunnea MacFarvanp, 1905 aD) ND) 1
Triopha grandis MacFarvanp, 1905 ND Dil 1 1 1
Cratena albocrusta MacFartanp, 1966 ND ND 1
Polycera atra MacFarvanp, 1905 ND ND 1 1 1
Aegires albopunctatus MacFarvanp, 1905 aD ND 1 1 1
Dirona picta MacFartanp, 1905 sD uD 1
Acmaea sp. aD ND 1 1
Ischnochiton sp. AD, ND 1
Echinodermata
Strongylocentrotus purpuratus (Stimpson, 1857) | %? NB 1 2 t i 2 1
Strongylocentrotus sp. ND sD 1 1 1
Ophiotrix spiculata LEConTrE, 1851 ND ND 1
Pycnopodia helianthoidces (Branpvt, 1835) ND ND 1
Pisaster sp. xD ND 1 tll
Eupentacta quinquescmita (SELENKA, 1867) AD aD) 1
Chordata (Tunicata) :
Aplidium solidum (RitTER & Forsytu, 1917) aD BE 1 2
Ascidia ceratodes (HUNTSMAN, 1912) ND ND 1 1 ih al | (ee Leeel eetee 1

Styela montereyensis (Dax, 1872)
Styela truncata Rirrer, 1901

Weight of dried organisms scraped froin
one side of 8 inch by 10 inch panel

3 0¢ 2%
3 CZ'8Z
8 Ch Eh

For explanation of symbols, see next page

Page 340

THE VELIGER

Vol. 10; No. 4

Explanation of symbols used in Table 3

1 Symbols used at head of columns indicate:
L=Lighted panels;
I= Intertidal panels;
F=Floating panels;

2 Symbols in columns indicate:
xp =No data

S=Shallow panels;
D=Deep wooden panels;
A=<Asbestos panels

1=species present in numbers from 1 to 10 individuals or colonies per panel side
2=species present in numbers from 11 to 20 individuals or colonies per panel side
3 =species present in numbers upward from 20 individuals or colonies per panel side

In addition to Hermissenda crassicornis, many species
of nudibranchs were observed on the long term panels,
but always in small numbers.

Echinodermata:

The only echinoderms to appear regularly on the test
surfaces were small green sea urchins. In the older panels
these became purple and could be identified as Strongy-
locentrotus purpuratus. They were especially abundant
on the floating rack and panels.

Chordata (Tunicata):

No tunicates were observed on the short term panels
nor on those in the water for 3 months. On older panels,
however, 4 species of tunicates were encountered. Asci-
dia ceretodes was on many of the panels of 6 months age
and older and at one time or another was found on panels
at all depths. It was never a dominant organism, and
reached a maximum size at 12 months of about 3cm
across.

Aplidium (formerly Amaroucium) solidum made an
inconspicuous first appearance on a deep asbestos panel
at 9 months. At 12 months this tunicate was found in
large numbers attached to all the racks and to panels
in the intertidal, floating and shallow positions. On the
intertidal rack and the 12-month intertidal panel, A.
solidum was clearly the dominant organism, forming
large massive slabs up to 10cm across.

Styela truncata and S. montcreyensis were found in
small numbers on several of the 12-month pancls and
racks. On the floating panel and rack S. montercyensis
reached a maximum length of 8cm.

WOOD BORING ORGANISMS

In addition to a study of the fouling organisms found in
Monterey Harbor, one of the purposes of this investi-
gation was to check on the occurrence of wood boring
animals. The presence of these destructive organisms in
a harbor is usually of considerably more economic im-

portance than that of foulers. Two species of borers were
found, the shipworm Bankia sctacca and the gribble Lim-
noria quadripunctata. Some of the gribbles were small
and lacked the characteristic tubercles on the telson as
found in L. quadripunctata, so it is possible a second spe-
cies of Limnoria is also present in the harbor.

Bankia setacea was found on only two of the short
term plywood panels from the shallow series that were in
the water between February 15 and April 1, 1967. In
both cases the borers were very small and had just begun
to excavate the wood. The first large shipworms were
found on one deep panel of the long term series at 6
months. There were only 3 borers in the wood, but they
were up to 5cm long. None were found in the 9-month
panels, but at 12 months large Bankia up to 8cm long
were found in the superficial layers of the plywood in
the lighted, intertidal, floating and deep positions. In
each panel there were usually 3 to 5 shipworms. The
heaviest concentration of Bankia encountered was not in
the panels, but in a board of 1 inch by 6 inch redwood
that had served as a proiective bottom in the stainless
steel lighted rack. When removed after 12 months in the
water this piece of wood was heavily fouled with bryo-
zoans and looked sound externally, but it was absolutely
riddled by large B. setacea up to 10 cm long and 0.7 cm
in diameter. It is doubtful if there was any room in the
wood for more shipworms, and the redwood crumbled
when handled. Yet, the redwood rack that supported the
floating panels appeared to be free of Bankza.

Limnoria quadripunctata was found on practically all
the wooden short term panels throughout the year. Usu-
ally only from 1 to 5 animals were found on any one
panel and they were often either wandering over the sur-
face or at work in very shallow burrows. The panels in
the shallow rack were most commonly attacked and sup-
ported the largest numbers of individual gribbles. On
the long term panels, Limnoria was not found in great
numbers until after the wood had been in the water
6 months or longer, but on these older panels, especially
the 12-month series, the outer lamellae of the plywood
were often completely riddled with burrows containing

Vol. 10; No. 4

THE VELIGER

Page 341

both adult and juvenile Limnoria. It was interesting to
note, however, that the encrusting bryozoans gave pro-
tection to the wood from gribble attack. Wherever exten-
sive, unbroken layers of Lyrula hippocrepis covered the
wood surface, no Limnoria burrows were found. In addi-
tion to the panels, the redwood floating rack carried
gribbles in the superficial wood layers.

SUMMARY

1. A series of 162 plywood and asbestos board panels
were exposed in the water under Monterey Municipal
Wharf No. 2 at various positions and water depths from
October 1, 1966 to October 1, 1967.

2. Short term panels (exposed for 1 month) made it
possible to determine season of settling of fouling and
boring organisms, while long term panels (exposed for
from 3 to 12 months) made it possible to measure rate
of growth and determine the dominant foulers.

3. On short term panels, a total of 70 species of
animals and plants settled on the surface, while on the
long term panels 81 species were recorded.

4, With few exceptions on both the short term and
long term panels, encrusting and erect bryozoans were
the dominant fouling organisms. These included Hippo-
thoa hyalina, Lyrula hippocrepis, and Crisulipora occi-
dentalis.

5. Balanus crenatus was the dominant barnacle and
was found on panels at all positions throughout the year.

6. Calcareous serpulid worms, Spirorbis spirillum and
Serpula sp., were the most common annelids found.

7. Water temperature and salinity records were kept
throughout the year but no clear correlation between
these physical factors and the rate of fouling was noted
with the possible exception of the influence of a gener-
ally higher temperature in late June and early July,
1967, on the settling at depths of Balanus crenatus.

8. Plywood and asbestos board, when exposed at 14
feet depth, collect essentially the same organisms and at
the same rate.

9. The wood borers Bankia setacea and Limnoria
quadripunctata attacked the plywood, especially in the
long term panels.

LITERATURE CITED

ALEEM, ANWER ABDEL
1957. | Succession of marine fouling organisms on test panels
immersed in deep water at La Jolla, California. Hydro-
biologia 2 (1): 40-58
Cor, WesLeY RoswELL
1932. Season of attachment and rate of growth of sedentary
marine organisms at the pier of the Scripps Institution of

Oceanography, La Jolla, California. Bull. Scripps Inst.
Oceano. Tech. Ser. 3 (3): 37 - 86

Cor, WEsLEY RosWELL & WINFRED EMory ALLEN
1937. Growth of sedentary marine organisms on experimental
blocks and plates for nine successive years at the pier of the

Scripps Institution of Oceanography. Bull. Scripps Inst.

Oceano. Tech. Ser. 4 (4): 101 - 136
DePatma, JouN R.
1966. A study of the marine fouling and boring organisms at

Admiralty Inlet, Washington.
Inf. Man. Rept. 0-6-66: 1 - 23
GraHaM, HERBERT WILLIAM & HELEN Gay
1945. Season of attachment and growth of sedentary
marine organisms at Oakland, California. Ecol. 26 (4):
375 - 386
Jounson, Martin Wicco « Ropert CUNNINGHAM MILLER
1935. The seasonal settlement of shipworms, barnacles and
other wharf-pile organisms at Friday Harbor, Washington.
Univ. Wash. Publ. Oceano. 2 (5): 1-18
Koro, Cuartes ATwoop & RoBperT CUNNINGHAM MILLER
1927. Biological section, pp. 188-343 In: Marine borers
and their relation to marine construction on the Pacific Coast.
Final report, San Francisco Bay Marine Piling Committee

U.S. Naval Oceanogr. Office

MILier, THomas LERoy
1966. Marine fouling organisms in Monterey Harbor, Cali-
fornia. Unpubl. Master’s Thes. U.S. Naval Postgrad.
School, Monterey, Calif. (October 1966)
MommseEN, Durwarp BELMONT
1966. A study of marine fouling in Monterey Harbor.
Unpub. Master’s Thes. U.S. Naval Postgrad. School, Monte-

rey, Calif. (May 1966)
PoMERAT, CHARLES Marc & CHARLES MANUEL WEISS
1946. The influence of texture and composition of surface
on the attachment of sedentary marine organisms. Biol.
Bull. 91: 57 - 65

ScHeEErR, BRADLEY Titus

1948. The development of marine fouling communities.
Biol. Bull. 89 (1): 103 - 121

Page 342

THE VELIGER

Vol. 10; No. 4

Studies on the Vitality of the Japanese Pearl Oyster

Pteria (Pinctada) martensi (DUNKER)

under Abnormal Conditions - I. Oxygen Uptake

and Shell Movement in Sea Water of Low Oxygen Content

TETUO MIYAUTI

The Laboratory of Takashima Pearl Farm, Inc.
Sasebo, Nagasaki Prefecture, Japan

(5 Text figures)

IN RECENT YEARS, the productivity of the sea water of
pearl culture farms has decreased and, from time to time,
shown low oxygen content.

In order to obtain further knowledge of the tolerance
of oysters for this extreme environmental condition, the
effects of low oxygen content of sea water on the rate of
oxygen uptake and shell movement of Japanese pcarl
oysters have been studied and the results are reported in
the present paper.

MATERIALS anp METHODS

Japanese pearl oysters, Pteria (Pinctada) martensi
(DuNKER), were reared in a pearl farm in Sasebo Bay,
Nagasaki Prefecture. The shell length of the oysters was
6.0 to 7.3cm and the age was 3 years.

Prior to the experiment the oysters were kept in a
laboratory tank with running sea water for 2 days. The
shells were cleaned to avoid errors due to the respiration
of small organisms attached to their surface.

The apparatus used in this investigation, shown in
Figures 1 and 2, was a modification of the type used by
Hatt (1929) for determination of oxygen uptake of
fishes at different oxygen contents. One oyster was placed
in a metabolism jar containing ca. 1.5 liters of sea water.
Temperature fluctuation was kept at a minimum (20°
+-1°C) by keeping the jar in a water tank (A), which
was supplied by filtered running water. The rate of
water flow from the supply tank into the metabolism jar
was kept constant (15 liters per hour) in order to avoid

the effect of current velocity which was reported by
MryauTr & Irie (1965, 1966a). The oxygen content of
the water in the jar was controlled by varying the number
of Mytilus edulis LinNagus, 1758, in the supply tank and
the volume of inflowing water from the storage tank into
the supply tank. To avoid the effect of daily rhythm
(Mort, 1948a), oxygen uptake was measured once for
every one hour period from 10 a. m. to 3 p. m.

In order to observe shell movement, the right valve
of the oyster was fastened with alpha cyanoacrylate to a
stone sinker placed at the bottom of the metabolism jar.
A glass fibre thread attached to the outer margin of the
left valve was linked to the lever of a kymograph (Figure
2), placed over the tank (A), and the shell movement was
recorded on a weekly kymograph.

Determination of dissolved oxygen was made by the
Winkler method. Experiments were carried out during
October and November of 1965.

RESULTS

Oxygen Uptake: The rate of oxygen uptake (cc/g dry
tissue/hour) was obtained from determinations of the
dissolved oxygen in the sea water flowing in and out of
the metabolism jar.

The results of tests made on 5 oysters are shown in
Table 1 and Figure 3. The rate of oxygen uptake re-
mained nearly constant when the oxygen content of the
sea water was maintained above 1.5cc per liter. Below
1.5cc of oxygen per liter, the uptake dropped abruptly.

Vol. 10; No. 4 THE VELIGER Page 343

Figure 1
Apparatus used for the determination of oxygen uptake
A: water tank supplied with running sea watcr; B: metabolism jar; F: Mytilus edulis; G, G: supply tube; H: stop cock;
C: supply tank; D: water sample tank; E: pearl oyster; I, I’: tube for taking samples of water

Figure 2

Apparatus used for the kymograph record of shell movement

A: water tank supplied with running sca water; B: metabolism jar; F: Mytilus edulis; G, G’: supply tube; H: stop cock;
C: supply tank; D: water sample tank; E: pearl oyster; I, I’: tube for taking samples of water
J: kymograph; K: stone sinker; L: liquid paraffin

Page 344

THE VELIGER

Vol. 10; No. 4

Table 1

Relation between the rate of oxygen uptake of the pearl oysters
and the oxygen content of the environment

Oxygen

content Oxygen uptake (cc/g dry tissue /hr)

(cc/L) A B Cc D E_ Average
5.00 — 4.51 0.541 0.485 0.455 0401 0.505 0.4774
4.50 — 4.01 0.537. 0.499 0.445 0.379 0.488 0.4696
4.00 — 3.51 0.514 — 0.457 0.415 0.519 0.4762
3.50 — 3.01 0.526 0.495 0.450 0.375 — 0.4615
3.00 — 2.51 0.526 0.450 0.499 0.369 0.499 0.4586
2.50 — 2.01 0.491 0.451 — 0.419 — 0.4537
2.00 — 1.51 0.491 0.405 0.445 0.315 0432 0.4176
1.50 — 1.01 0.298 0.098 0.279 0.086 0.217 0.1956
1.00 —0.51 — 0.023 0.032 0.066 0.115 0.0590
0.50 —0.15 — 0.011 0.014 0.024 0.009 0.0145

Date of experiment: October 22 to November 9, 1965
Cl: 18.01 to 18.45%,

Oxygen Uptake (cc/g dry tissue/hour)

4 3 2 I (0)
Dissolved Oxygen of the Environment (cc/1)

Figure 3
Effect of low oxygen content on the oxygen uptake rate
of the pearl oysters

As shown in Table 1, the rate of oxygen uptake was
0.372 - 0.541 cc per g dry tissue weight’ per hour when
the amount of oxygen of the sea water was above 1.5cc

per liter. Morr (1948 a) found it to be 0.029 - 0.068 cc per
g wet tissue weight’ per hour; Mort (1948b), 0.02 - 0.05
cc per g wet weight per hour; Sawano (1950), 0.074 to
0.1447cc per g wet weight per hour; UEmorto e¢ al.
(1964), 0.022 - 0.087 cc per g wet weight per hour;
MryauT! & Irie (1966b), 0.3 - 0.6cc per g dry weight
per hour. These differences in value of the rate of oxygen
uptake may be due to differences in experimental con-
ditions, such as water temperature, age of oysters, experi-
mental season, physiological condition of oysters and other
parameters.

BERKELEY (1923) reports that the presence of the

crystalline style in certain molluscs is related to anaerobic
respiration. He observed that a mollusc maintained under
anaerobic condition for 8 days lost the crystalline style
entirely. On the other hand, Nozawa (1929) found that
the crystalline styles of Ostrea circumpicta disappear
after anaerobic respiration, though not completely. In this
experiment, the crystalline style diminished in size after
48 hours (but did not disappear entirely), under 0.5 cc
oxygen content per liter of sea water.
Shell Movement: A representative kymograph record of
shell movement under normal conditions during this ex-
perimental period is shown in Figure 4. Figure 5 shows
shell movement at different levels of oxygen content. The
results were obtained from 4 oysters.

Under normal conditions, the degree of valve opening
showed periodic changes (daily rhythm), and the fre-
quency of opening and closing was rclatively low (Figure
4). If, however, the oxygen content was reduced to about
1.5 cc per liter, the frequency of opening and closing was
much higher, and the degree of valve opening was
greater. When the oxygen content was reduced to about
0.5cc or less per liter, the daily rhythm of shell move-
ment disappeared.

All oysters gradually regained their normal vitality
when they were returned to a pearl farm after 3 to 7
days’ experiments.

DISCUSSION

The present experiments show that the rate of oxygen
uptake and shcll movement of the Japanese pearl oyster
are independent of the oxygen content of the environ-
ment, when the latter is above 1.5 cc per liter.

It has been known that low oxygen content of the
environment influences the physiological activity of the
Japanese pearl oyster. According to Sawano (1950) and

10 g wet tissue weight without shell is equivalent to about 1.2 to
1.3 g dry tissue weight without shell.

Vol. 10; No. 4 THE VELIGER Page 345

Nov. 4 Nov. 6

Figure 4
Kymograph record showing the shell movement of pearl oyster

in normal sea water

White and black bands at the bottom show daytime (6:00 - 18:00) and night time (18:00—6:00). The writing pen is at the lowermost
position when shell is closed.

a i i | : /

Dissolved Oxygen (cc/1)
bho f£ wo

Dissolved Oxygen (cc/1)
i) » of Oo

Figure 5 -— continucd next page

<

Page 346

THE VELIGER

Vol. 10; No. 4

Dissolved Oxygen (cc/I)

Dissolved Oxygen (cc/1)

if WOM f,
If WE Wf |

Figure 5

The effect of low oxygen content on the shell movement of
the pearl oysters. The base line is marked in two-hour intervals.

Uemoto et al. (1964), the rate of oxygen uptake of the
pearl oyster was almost directly proportional to the oxy-
gen content of the sea water. On the other hand, Mort
(1948b) and Miyautr & Irte (1966b) found that the
rate stayed constant when the amount of oxygen of the
sea water was above ().5cc per liter and 1.5 - 1.0ce per
liter, but it decreased considerably when the amount was
lower than that level. The results of the former two
studies differ from those of the present experiment; the
latter are roughly similar to the present results. These
differences in the rate of oxygen uptake may be due to
differences in the experimental methods used. In the
previous investigations, the water surface was covered with
liquid paraffin. Consequently, the oxygen content was
reduced continually as the oyster removed oxygen from

the water. Therefore, the results of such experiments do
not accurately represent oxygen uptake at one particular
oxygen content level. The method used in the present
experiment, however, removes this difficulty, and fairly
consistent results were obtained. In addition, the results
of the present experiment are in agreement with those
found for Ostrea circumpicta (Nozawa, 1929), O. edulis
(GaLtsorr & WauppLe, 1930), Ostrea gigas (IsHma,
1935), Pecten grandies, P. irradians (VAN Dam, 1954),
Mytilus edulis (Bruce, 1926; RorrHauwe, 1958), and
Anodonta cygnea (Hers, 1943).

Low oxygen content also affects shell movement.
MryautT! & Irte (1966b) reported that when the oxygen
content was reduced to 1.5-1.0cc per liter or less, the
frequency of shell movement was much higher and the

Vol. 10; No. 4

THF VELIGER

degree of opening of the valves was greater; when oxygen
was reduced to 0.5cc or less per liter, the degree of
opening of the valves was slightly smaller; and when it
was reduced to 0.2 cc or less per liter, the valves eventually
gape open after closure for several ten-minute periods.
The minor difference in shell movement between the
above results and the present results may be ascribed to
the use of a different method. In the previous investiga-
tion, the water surface was covered with liquid paraffin,
so that the oysters may be affected by accumulated metab-
olites in addition to the low oxygen content.

The results of the present experiments do not necessar-
ily indicate the response of the oyster as a whole animal.
However, oxygen uptake and shell movement are impor-
tant physiological factors controlling the vitality of the
oyster; therefore these results will offer some fundamental
data for the living conditions of oysters and will give an
indication of the tolerance to low oxygen tension.

The most reasonable conclusion is that the pearl oysters
are affected by low oxygen content only when the latter
is 1.5cc per liter, or less.

SUMMARY

1. The effects of oxygen content on oxygen uptake and
shell movement of the Japanese pearl oyster were studied.
2. The rate of oxygen uptake of this animal remained
relatively constant until the oxygen content of the medi-
um was decreased to 1.5cc per liter.

3. The shell movement was generally normal until the
oxygen content of the medium was decreased to 1.5 cc
per liter.

4. Below 1.5cc per liter, the rate of oxygen uptake
dropped to a low level, and the shell movement was
abnormal.

ACKNOWLEDGMENTS

The author wishes to express his sincere thanks to Prof.
Yaichiro Okada, ‘Tokai University, Prof. Sadayoshi
Miyake, Kyushu University, and Assistant Prof. Tadashi
Tsujii, Prefectural University of Mie, for their valuable
suggestions and criticisms. It is a pleasure to record here
a debt of gratitude to Dr. Norimitsu Watabe, Duke Uni-
versity, for his kindness in correcting the manuscript.

LITERATURE CITED

BERKELEY, C.
1923. On the crystalline style as a possible factor in the
anaerobic respiration of certain mollusks.

Zool. 37: 447 - 488

Journ. Exp.

Page 347
Bruce, J. R.

1926. The respiratory exchanges of the mussel (Mytilus edu-
lis). Biochem. Journ. 20: 829 - 846 [cited by EF Guiretti,
(1966) ]

Hatt, E G.
1929. The influence of varying oxygen tensions upon the rate

of oxygen consumption in marine fishes.
Physiol. 88: 212 - 218

Amer. Journ.

Hers, M. J.
1943. Relation entre respiration et circulation chez Anodonta
cygnea L. Ann. Soc. Roy. Zool. Belg. 74: 45 - 54 [cited by
F Guiretti (1966) |
Isuipa, S.

1935. On the oxygen consumption in the oyster, Ostrea gigas
Thunberg under various conditions. Sci. Reprt. Tohoku
Imp. Univ. 10: 619 - 638 (in English)

GatsorFF, Paut Simon & D. V. WuprLe
1930. | Oxygen consumption of normal and green oysters.
Bull. U.S. Bur. Fisheries 46: 489 - 508

GuireEtTTI, F
1966. Respiration. In: Physiology of Mollusca (ed. Karl
Milton Wilbur and Charles Maurice Yonge) 2: 201 - 203.
Acad. Press, New York, N. Y.

Mryaut!, TeTuo «& H. Irie
1965. Relation between pearl oysters (Pteria martensii) and
the current-velocity of the environmental waters. Bull.
Fish. Nagasaki Univ. 19: 56-6+ (in Japanese with English
sununary )

1966a. Supplemental report on the relation between pearl
oysters (Pteria martcnsit) and current-velocity of environ-
mental waters. Effects of the velocity of the environmental
waters on the motion of shell opening and shutting and shell
regeneration. Bull. Fish. Nagasaki Univ. 20: 22-28 (in
Japanese with English summary)

1966b.

movement of the pearl oyster, Pécria martensii, in the sea water
Bull. Fish. Nagasaki Univ. 21: 139-144
(in Japanese with English summary)

Mort, S.
1948a. Daily rhythmic activities of “Martens” pearl-oysters
(Pinctada martensii (DUNKER) ). Venus 15 (1-4): 46-51
(in Japanese)
1948b. On the respiration of Pinctada martensii (DUNKER) in
the sea water of low oxygen tension (a preliminary notc).

Venus 15 (1-4): 52-54 (in Japanesc)

Nozawa, A.
1929. The normal and abnormal respiration in the oyster,
Ostrea circumpicta Pils. Sci. Reprt. Tohoku Imp. Univ.
Biol. 4: 315 - 325 (in English)

RotTHAuwE, H. W.

1958. | Untersuchungen zur Atmungsphysiologic und Osmo-
regulation bei Mytilus cdulis mit einem kurzen Anhang iiber
die Blutkonzentration von Dreissensia polymorpha in Ab-
hangigkeit von Elektrolytgehalt des AuRenmediums. Ver-
Off. Inst. Meeresforsch. 5: 143-159 [cited by E Guirerri
(1966) ]

Relation between the oxygen consumption and _ shell

of low oxygen tension.

Page 348

THE VELIGER

Vol. 10; No. 4

Sawano, E.

1950. Shinzyugai no Chityu-yoshoku-ho no Kenkyu (Studies
on pond culture of pearl oyster). Suisankenkyu Kaiho 3: 48
to 57 (in Japanese)

Uemoto, H., M. WataBe « M. Matsuo

1964. Physiological studies on the nuclear insertion operation
of pearl oyster. VII. On the decreases of oxygen consumption
of oysters which had been kept under controlled state in the

physiological conditions. Bull. Nat. Pearl Res. Lab. 9:
1121-1127 (in Japanese)
Van Daw, L.

1954. On the respiration in scallops (Lamellibranchia).
Biol. Bull. 107: 192 - 202 [cited by F Gutretti (1966) |

Vol. 10; No. 4

THE VELIGER

Page 349

The Date of Publication of Kirner’s Mitra Monograph in the

““Spécies général et iconographie des coquilles vivantes”’

BY

WALTER OLIVER CERNOHORSKY

Vatukoula, Fiji Islands

THE INTERPRETATION OF THE DATES of issue of KIENER’s
“ivraisons” of molluscan monographs has caused some
difficulty to malacologists in view of the irregular appear-
ance of the undated fascicles. SHERBORN & WooDWARD
(1901), realizing the need for a stabilized interpretation
of publication dates, collated K1ENER’s monographs as to
the date of issue of text and plates. As in many instances
the plates appeared prior to the text, KIENER’s new
species names are available from the date of issue of
the plates which contain specific names on plate legends.
SHERBORN & Woopwarbd’s collatory work (op. cit.) was
based on external evidence as to the date of appearance
of the fascicles; the authors realized that many gaps
remain to be filled.

According to SHERBORN & Woopwarp (op. cit.) the
plates to Mitra and Voluta were supposed to have been
issued in fascicle 34, which they date from 22 April
1839; the complete set of plates and text of the mono-
graphs on Mitra and Voluta were said to have appeared
in fascicles 34-47 during 1839. The text to Mitra and
the plates to Voluta most probably were issued during
1839; however, there is evidence that the Mitra plates
appeared already in 1838.

Evidence as to an earlier publication date of the
Mitra plates may be found in Epuarp ANTon’s “Ver-
zeichnif&g der Conchylien” (1839), a work which is gen-
erally considered to antedate KiENER’s Mitra monograph.
A perusal of this publication shows, however, that ANn-
TON’s “Verzeichnif{” postdates KiENER’s Mitra mono-
graph. The title page of the “Verzeichnif&” bears the date
1839, ANToN’s “Vorrede” (page iii) is dated 3 August
1838, and the last page, 110, bears the imprint “Im
Druck beendigt den 9. Oktober 1838,” meaning that by
that date printing was completed. ANTON (page x) lists
the titles of works used in his study, and acknowledged
receipt of KieNnerR’s 32 fascicles dated 1834 - 38 as fol-
lows: Thracia, Pyramidella, Tornatella, Scalaria, Delphi-
nula, Solarium, Rotella, Mitra, Struthiolaria, Purpura,
Eburna, Terebra, Buccinum, Harpa, Dolium, Cassidaria,
Cassis and Marginella. In the text ANToN freely quotes

Krener’s species and figures of Mitra, the highest figure
number cited from KiENeER being 105. It is evident that
prior to the completion of printing of the “Verzeichnif,,”
i.e. 9 October 1838, ANToN received the complete set
of Mitra plates (32 plates out of a total of 34 being
cited), which must have appeared in or prior to fascicle
32. It appears, therefore, that the records of the “Biblio-
graphie de la France” as cited by SHERBORN & Woop-
WARD (op. cit.) were correct after all when acknowledg-
ing receipt of KrENER’s “‘livraisons 26 4 36” by December
1838. SHERBORN & Woopwakrb suspected that this should
have read 20 - 30.

It is evident that KrENEr’s Mitra monograph, apart
from the text, should date from 1838, and that it ante-
dates ANToN’s “Verzeichnif.”” Since there is no evidence
in AnTon’s publication that the Mitra text appeared
already in 1838, SHERBORN & Woopwarp’s date of 1839
should be tentatively retained for the text alone. The
species M. cancellarioides ANTON, which replaces M.
nodosa SwAINSON, 1823 (non Borson, 1820), has prior-
ity over M. tuberculata Kiener, which first appeared on
plate 27, figure 87 as M. fraga (non Quoy & GarmarD,
1833; nec Kiener, plt. 9, fig. 26). KieENER changed his
homonymous M. fraga to M. tuberculata in the text to
Mitra under the heading “Table des Espéces des Mitres”
on page 119, which dates from 1839.

LITERATURE CITED

ANTON, HerMANN Epuarpb

1839. VerzeichniS der Conchylien welche sich in der Samm-
lung von Hermann Eduard Anton befinden. Halle, pp.
i- xvi; 1-110
Krener, Lours CHarLes
1839. Spécies général ct iconographie des coquilles vivantes;

Genre Mitra. Rousseau, Paris [3]: 1-120; plts. 1 - 34
(text: 1839; plates: 1838)

SHERBORN, CHARLES Davies « BERNHARD BARHAM WoopwarRD
1901. Notes on the dates of publication of the parts of Kiener’s
“Spécies général et iconographie des coquilles vivantes,” etc.
(1834-80). Proc. Malac. Soc. London 4 (5): 216-219

(25 July 1901)

Page 350

THE VELIGER

Vol. 10; No. 4

Itinerary of the Voyage of H. M.S. Blossom, 1825 to 1828

BY

JOSEPH ROSEWATER

Division of Mollusks, United States National Museum Smithsonian Institution, Washington, D. C.

His Majesty’s sHip Blossom under the command of
Captain F W.Beechey sailed to the Pacific and Arctic
Oceans in the year 1825 and remained in that area until
1828 to provide support for an expedition mounted by
the British for the purpose of discovering a northwest
passage. The ship was scheduled to meet the expedition
north of Bering Strait some time during the months of
July, August or September of the years 1826 or 1827,
and she was ordered to explore, chart and to collect
natural history specimens during the balance of the time.
Upon her return to England the natural history speci-
mens were distributed to various naturalists for study.
The mollusks were entrusted to J. E. Gray and his report
on them appeared after numerous delays (Gray, 1839;
see also general introduction by F W. BEECHEy, pp. vii,
viii). In his introduction to the section on Mollusca,
Gray remarked that a portion of the Blossom collections,
mostly that collected by Captain Belcher and presented
by him to the Zoological Society, was described by
BRoDERIP & SOWERBY (1829). Gray mentioned that he
was including in his report additional material collected
by others from some of the same areas that had been
visited by the Blossom, but since he seldom named a
collector or a precise locality, there is little basis for
considering material to be from a source other than that
voyage.

Unfortunately, naturalists of that day in many cascs
refrained from citing an exact locality for a species they
were describing. Probably this was not entirely duc to
carelessness on their part, but often was the fault of
collectors who neglected to label specimens properly.
Today, this would be as inexcusable error, but at that
time it probably seemed just not that important.

While working with the report on the Blossom Mollus-
ca, in an attempt to identify certain Indo-Pacific marine
mollusks, I found the quality of the locality data to vary
considerably. Over 100 new species of mollusks are de-
scribed in the report and locality information ranges in

20560

precision from none at all to “Pacific” to relatively excel-
lent data such as “Icy Cape.” It will be noted that the
section completed by G.B.SowerBy (pp. 143 - 155)
often contains considerably better locality information
than that which precedes it written by Gray (pp. 103 to
142). During a brief visit to the British Museum (N. H.)
in 1963 I examined some of the types of species described
by Gray in his report and found that the museum labels
lacked additional locality data.

In many groups of mollusks it is of considerable impor-
tance to ascertain with fair precision the locality from
which a particular species was described because similar-
appearing but distinct species often live in widely sepa-
rated regions. For this reason type localities are often
designated in order to localize a taxon and to remove it
from geographical limbo. To provide some basis for type
locality designations in a similar case, CHAMBERLAIN
(1960) prepared an itinerary of the voyage of the Venus.
I have prepared the following itinerary of the voyage of
the Blossom, and I hope it will obviate the necessity for
future workers to search the cruise narrative to discover
appropriate type localities.

BreecHeEy (1831) prepared an elaborate narrative of
the voyage of the Blossom containing rather complete
and fascinating descriptions of the places and peoples
visited. Events are related chronologically, but precise
dates and places are buried in the text. In order to
extract these for an itinerary it was necessary to read the
narrative in some detail, a not unpleasant task. Where
locality names differ appreciably from those recognized
today, I have used either the modern name or indicated

it in parentheses. The dates of visitation associated with
each locality are not always certain and in some cases

had to be left blank. It is often difficult to be sure from
the cruise narrative that the ship actually stopped at a
locality or whether it merely observed the position of an
island or port while sailing past.

Vol. 10; No. 4 THE VELIGER Page 351
Arrival Nihau, Hawaii June 1-2
and/or Petropavlovsk, Bay of Awatska

Departure Kamchatka, U.S.S.R. June 28 to
Localities Dates July 5

Departed from Spithead, England May 19, 1825 Off Bering Island,

Santa Cruz de Tenerife, Canary Ids. June 2-5 Commander Islands, U.S.S.R. July 10

Rio de Janeiro, Brazil July 11-13 Off St. Lawrence Island, Alaska July 16

Rounded Cape Horn September 16 Off King Island, Alaska July 19

Isla Mocha, Chile October 6 Diomede Islands,

Talcahuano and Concepcién, Chile October 8-20 Bering Strait, U.S.S.R. July 20

Valparaiso October 27-29 Schismareff Inlet, Cape Prince

Off Sala-y-Gémez November 15 of Wales, Alaska July 21

Basten Island Nov. 17-18? Kotzebue Sound, Alaska July 22-23

Ducie Island November 29 Chamisso Island, Mouth of

Henderson Island December 3 Eschscholtz Bay, Alaska July 25-30

Pitcairn Island December 5 - 21 Cape Thompson, Alaska August 2

Oeno Island December 23 Point Franklin, Alaska August 15

Gambier Islands Dec. 28, 1825 Between Icy Cape and Cape

Jan. 13, 1826 Beaufort, Alaska August 20

Lord Hood’s Island Point Hope, Alaska August 26

(Marutea, Tuamotus) January 14 Return to Chamisso Island August 28
Clermont Tonnere Explorations of Kotzebue Sound
(Reao, Tuamotus) January 18 and vicinity while waiting for
Serle Island Captain Franklin and shore party
(Pukarua, Tuamotus) January 21 (September 10; return of Barge
Whitsunday Island from north to 71° 23’31”N and
(Pinaki, Tuamotus) January 22 156° 21°30” W) August 29 to
Queen Charlotte’s Island October 14
(Nukutavake, Tuamotus) January 23 King Island, Alaska October 16
Lagoon Island St. Paul, Pribilof Islands, Alaska October 21
(Vahitahi, Tuamotus) January 24 Unimak, Aleutian Islands, Alaska October 22

November 7 to
December 28
January 1-5, 1827

January 26 to

Egmont Island San Francisco, California
(Vairaatea, Tuamotus)
Barrow Island

(Vanavana, Tuamotus)

January 25?
Monterey, California
Honolulu, Hawaii

Carysfort Island March 3
(Tureia, Tuamotus) Left Hawaii March 4
Osnaburgh or Matilda South of Wake Island March 15

(Mururoa, Tuamotus) February Assumption (Asuncion, Marianas) March 25
Byam Martin Island Bashee Islands (Batanes Islands) April 7

(Ahunui, Tuamotus) Macao April 10-30
Gloucester Island Naha, Okinawa, Ryukyu Ids. May 18-25

(Paraoa, ‘Tuamotus) Bonin Islands June 8
Bow Island Petropavlovsk, Bay of Awatska

(Hao, Tuamotus) Feb. 15-20 Kamchatka, U.S.S. R. July 3-20
Tahiti, Society Islands March 18 to Bering Island,

April 26 Commander Islands, U.S.S.R. July 22

Oahu, Hawaii May 19-31 St. Lawrence Island, Alaska August 1

Page 352 THE VELIGER Vol. 10; No. 4
King Island, Alaska August 2 Acapulco, Mexico March 13-18
Chamisso Island, Kotzebue Sound, Crossed Equator March 29

Alaska (Lieut. Belcher takes Barge Valparaiso, Chile April 29 - May 20

north to explore Icy Cape) August 5 Coquimbo, Chile May 23 - June 3
Cape Krusenstern, Alaska August 25 Cape Horn June 30
Chamisso Island, Kotzebue Sound, Rio de Janeiro, Brazil July 21

Alaska August 26 Spithead, England October (prior
Cape York, Alaska August 31 to 12", 1828)
Exploring north to vicinity of Icy

Cape and southward to Port LITERATURE CITED

Clarence and return to Kotzebue

Sound waiting for Capt. Franklin Brecuey, FW.

and shore party October 6 1831. Narrative of a voyage to the Pacific and Beering’s
Daesthare asta Sreake eradl fa when Strait to cooperate with the Polar Expeditions: performed in

PES ELS peer! His Majesty’s Ship Blossom, under the coninand of Captain

of St. Lawrence Island, Alaska October 6-7 FE W. Becchey, R.N. in the years 1825, 26, 27, 28. Quarto
St. Paul, Pribilof Islands October 12 ed., Colburn & Bentley, London; publ. in 2 prts.: prt. 1, xxiii+
Unimak Island, Aleutian Islands October 14 pp. 1-392; prt. 2, viiit+ pp. 393 - 742. American Octavo ed.,

Monterey, California
San Francisco, California

Off Cape St. Lucas, Baja
California, Mexico

Off Tres Marias Islands, Mexico

San Blas, Mexico

Mazatlan, Mexico

Exploration of coast between
Mazatlan and San Blas
Left San Blas, Mexico

October 29 to
November 17

November 18 to
December 3

December 13
December 14
December 16

February 3 to
Feb. 7, 1828

February 7
March 8

1832, Carey & Lea, Philadelphia, vi+493 pp. [lacks appen-
dix and illustrations]
Broperip, WILLIAM JOHN & GEORGE BRETTINGHAM SOWERBY

1829. | Observations on new or interesting Mollusca contained,
for the most part in the Museum of the Zoological Society.
Zool. Journ. 4: 359 - 379; 5: 46 - 51

Cuampberuin, J. L.

1960. Voyage of the Venus.
68

The Nautilus 74 (2): 65 to

Gray, JoHN Epwarp & GrorGE BRETTINGHAM SOWERBY
1839. | Molluscous animals and their shells im The Zoology
of Captain Beechey’s voyage in His Majesty’s Ship
Blossom. ... H. G. Bohn, London, pp. 101 - 155; plts. 33 to
44 [by J. E. Gray; continued by G. B. SowErBy]

Vol. 10; No. 4

THE VELIGER

Page 353

The Ovulidae, Pediculariidae and Triviidae of Fiji

(Mollusca : Gastropoda)

WALTER OLIVER CERNOHORSKY

Vatukoula, Fiji Islands

(Plates 50 to 52; 5 Text figures; 1 Map)

INTRODUCTION

THIS IS THE EIGHTH PART in the series of faunal mono-
graphs dealing with the marine mollusca of the Fiji
Islands.

Unlike the Cypraeidae, species of Ovulidae, Pediculari-
idae and Triviidae are moderately rare in Fiji; the minute
size and often inaccessible habitat are responsible for their
rare occurrence in local collections.

All the species recorded from the Fiji Islands have a
Pacific or Indo-Pacific distribution, and the majority of
species live within the area of the East African coast and
the Tuamotu Archipelago.

In this, as in the previous monographs, only species
collected by resident collectors and the author are listed as
verified records. For notes on the geography of the Fiji
Islands and other pertinent data see CERNOHOoRSKy, 1964.

TAXONOMY

The higher categories of Cypraeacea have been discussed
in a recent publication by ScHiLpER (1966). Despite the
non-binding recommendation by the International Com-
mission on Zoological Nomenclature that the ending
“-oidea” be employed for Superfamilies, most malaco-
logists have accepted the ending “-acea” since the publi-
cation of THIELE’s ‘Handbuch der systematischen Weich-
tierkunde’ (1929). The taxon Cypraeacea has been
introduced by ScuitpER (1927) and was accepted by
THIELE (op. cit.) and most subsequent authors. The
superfamilial term Cypraeoidea has been used by IREDALE
(1935) and Scuitper (1939, 1941). Cypraeoidea has,
however, been recently re-introduced into literature by
TREDALE & McMicuaer (1962), and the suffix ‘-acea’
has been retained in this publication for some subclasses
of molluscs. In view of the non-binding recommendation
regarding term endings for superfamilies in zoology, a

standardization of such suffixes by malacologists would be
welcome.

The family Lamellariidae has been removed from the
Cypraeacea by previous authors and placed in the super-
family Lamellariacea; the families Cypraeidae, Ovulidae,
Pediculariidae, and Triviidae have been retained in the
Cypraeacea. In a broad classification of all mollusca, the
erection of a new superfamily Triviacea for the reception
of ‘Triviidae and Pediculariidae, would not appear to
contribute to a clearer definition of relationships between
these allied families. A continuous upgrading of higher
categories of Cypracacea by specialists may finally result
in monofamilial superfamilies if further divisions will
prove imperative. Since the ranking of higher taxonomic
categories is largely subjective, a retention of Triviidae
and Pediculariidae in the Cypraeacea may appear ortho-
dox to the specialist, but would not tend to obscure
existing phylogenetic relationships between the families;
it would also be more consistent with our present classi-
fication of higher categories of other gastropod groups.

Species of Ovulidae have been assigned to various
generic groups since the start of organized nomenclature:
Bulla Linnaeus, 1758; “Amphipceras’ Gronovius, 1781
(non binomial) ; Ovula BrucutERE, 1789; Volva and Cy-
phoma Ropine, 1798; and Radius ScHUMACHER, 1817.
Montrort (1810), who had a decided preference for
gencric names of masculine gender changed Ovula to
Ovulus and also established Calpurnus and Ultimus,
while SowerBy (1828) introduced Ovulum, which he
considered properly latinized. The genera Ovula and
Ovulum remaincd in use for the remainder of the 19"
century together with Simnia Risso, 1826, Radius
SCHUMACHER and Neosimnia Fiscuer, 1884. In the early
part of the 20" century “Amphiperas” was re-introduced
and several new genus-group names were proposed by
THIELE (1925, 1929), ScumpEr (1927) and IREDALE
(USS OMS SIF 1935))e

Page 354

THE VELIGER

Vol. 10; No. 4

Living species of Pediculariidae had a less eventful

taxonomic history, due mainly to the small number of
species and the somewhat belated appearance of the first
Recent species, Pedicularia sicula Swainson, 1840, in
molluscan literature. The authors H. and A. Apams
(1854) erected the family Pediculariidae and the genera
Pediculariella Tuer, 1925 and Pediculariona (IREDALE,
1935, nud.) ScuitpER, 1939 were subsequently estab-
lished. Most authors considered Pediculariidae worthy
of familial rank, but ScHiLDER (op. cit.) combined all
species in the subfamily Pediculariinae within the family
Ovulidae; in 1966, however, he distinguished Ovulidae
from Pediculariidae and transferred the latter family to
Triviacea.
As far as the Triviidae are concerned, authors were unable
to reach agreement as to generic placement of the species.
Species of Triviinae and Eratoinae were assigned to Cyp-
raeca LINNAEUS, 1758, Bulla Linnagus, 1758, Voluta
Linnaeus, 1758, Marginella Lamarck, 1799, Colum-
bella LamMarcK, 1799, and other lesser known genera.
The introduction of the new genera Erato Risso, 1826,
Lachryma Sowersy [1832], (introduced inadvertently),
Trivia Bropverip, 1836, Triviella, Trivirostra, Niveria and
Pusula JouSSEAUME, 1884, did not appreciably change
the taxonomic concept of this group; species of Trivi-
inae continued to be assigned to the Cypraeidae, and
Eratoinae to Marginellidae. ScuHiLpER (1927) thoroughly
revised the group according to phylogeny, and IREDALE
(1930, 1931, 1935) contributed to the subdivision of the
family by establishing several new genus-group names
whose validity is seriously questioned by practising mala-
cologists.

IREDALE’s GENUS -GROUP NAMES

Several adverse comments have been passed by recent
malacologists and zoologists on the validity of genus- and
species-group names proposed by IrEDALE (Zier, 1960;
Sotem, 1964). Whatever the taxonomic shortcomings of
TREDALE’s generic diagnoses, their brevity and casualness
appear to be the product of taxonomic procedures preva-
lent among malacologists at the time, in contrast to

present-day systematics. It is unfortunate that authors
continue to use genus-group names proposed by IREDALE,
which in accordance with the rules of the Code of the
ICZN (1964) are nomina nuda. Although a common-
sense interpretation of the Code of the ICZN appears
advisable at times, the rules governing the availability
of zoological names are unequivocal: a name can be
either valid or invalid, but not tentatively valid.

The usage of Iredalean genus-group names in this
monograph made it advisable to test each scientific name
established by Iredale between 1930 and 1935 as to its
taxonomic availability. It became apparent that Iredale’s
names fall into three categories: 1) names which are
technically and scientifically valid; 2) names which are
technically nomina nuda; and 3) names which are
technically valid but scientifically nomina dubia.

The majority of Iredale’s names proposed prior to 1931
satisfy the requirements of art. 12 in conjunction with
art. 16(v) or 16(vi) of the Code of the ICZN; their
definition may not be scientifically adequate, but they are
certainly available. Names established after 1930 are
either valid, invalid or nomina dubia; the first must be
accepted, the second rejected; it is the nomina dubia
which will pose a taxonomic problem.

A list of genus-group and specific names proposed by
TREDALE (1931, 1935), which qualify as nomina nuda
(art. 13, ICZN) is appended. In all cases these genus-
group names have been diagnosed by ScuiLpER (1939)
and validated by ScuILpeErR as author:

Ellatrivia (IREDALE, 1931, nud.) ScutpER, 1939

1931. Ellatrivia IREDALE, Rec. Aust. Mus. 18: 221 (nom. nud.)
1935. Ellatrivia IREDALE, Aust. Zool. 8: 100 (nom. nud.)
1935. Ellatrivia ScuitpER, Proc. Malacol. Soc. London, 21:
330 (nom. nud. — no type fixation, art. 13(b) of ICZN)
1939. Ellatrivia ScurtpER, Arch. Molluskenk. 71: 173, fig. 26
(type spec. by OD Triviella merces IREDALE, 1924)

Fossatrivia (IrEDALE, 1931, nud.) Sctmper, 1939

1931. Fossatrivia Irepavr, Rec. Aust. Mus., 18: 222 (nom. nud.)
1935. Fossatrivia IREDALE, Aust. Zool., 8: 100 (nom. nud.)
1939. Fossatrivia ScuitperR, Arch. Molluskenk., 71: 173 (type
spec. by OD Trivia caelatura HEvLEy, 1918)

Explanation of Plate 50

Figure 1: Ovula ovum (LinNakus). Fiji. (x 0.5)

Figure 2: Ovula costcllata (LAMaRCcK). Fiji. (x 1.0)

Figures 3and3a: Calpurnus verrucosus (LINNAEUS). Fiji. (x 2.0)
Figure 4: Calpurnus lacteus (Lamarck). Fiji. (x 2.5)

Figure 5: Primovula punctata (Ductos). Fiji. (x 3.0)

Figure 6: Prionovolva fruticum (REEvE). Fiji. (x 2.0)

Figure 7: Prosimnia coarctata (ADAMS & REEVE). Fiji. (x 3.5)
Figure 8: Primovula striatula (SowERBY). Fiji. (x 5.0)

Figure 9: Volva volva (LinNatus). Fiji. (x 0.6)

Figure ga: Volva volva (Linnafus). Fiji, juvenile specimen (x 1.0)

Figure 10: Phenacovolva birostris (LINNAEUS). Fiji. (x 2.5)

Tue VELIGER, Vol. 10, No. 4 [CERNOHORSKY] Plate 50

Figure 4 Figure 5 Figure 6

Figure 7 Figure 8 Figure 9 Figure 9 a Figure 10

photographs by W. O. CerNoHORSKY

Vol. 10; No. 4

THE VELIGER

Page 355

Volva volva cumulata (IrEDALE, 1931, nud.) IREDALE,
1935
1931. Volva volva cumulata IrEpALE, Rec. Aust. Mus., 18:
222 (nom. nud.)
1932. Volua cumulata ScuiprER, Proc. Malacol. Soc. London,
20: 56 (nom. nud.)
1935. Volva volva cumulata IREDALE, Aust. Zool. 8: 104

Pellasimnia (IrEDALE, 1931, nud.) ScHILpER, 1939

1931. Pellasimnia IrEDALE, Rec. Aust. Mus., 18: 222 (nom.
nud.)

1932. Pellasimnia ScuiLpER, Proc. Malacol. Soc. London, 20:
54 (in synonymy of Neosimnia FiscueEr, 1884 —no type
fixation, art. 13(b) of ICZN)

1935. Pellasimnia IrEDALE, Aust. Zool. 8: 104 (nom. nud.)

- 1939. Pellasimnia Scutver, Arch. Molluskenk., 71: 195 (type
spec. by OD Ovulum angasi REEVE, 1865)

Margovula (IrEDALE, 1935, nud.) ScHILDER, 1939
1935. Margovula IrepaLE, Aust. Zool. 8: 103 (nom. nud.)
1939. Margovula Scuitper, Arch. Molluskenk. 71: 200 (pub-

lished in synonymy of Diminovula IrEDALE, 1930 - type
spec. by OD M. pyriformis (SowERBy, 1828) )
1941. Margovula Scuttporr, Arch. Molluskenk. 73: 68

Margovula appears to have been validated by Scuit-
DER (1939) through publication in synonymy (art. 11(d)
of ICZN) and subsequent adoption by ScuiLpErR (1941)
as a senior synonym of Diminovula IREDALE, 1930.

Pediculariona IREDALE, 1935 (p. 101) was diagnosed as
follows: “The elevated sculptured spire later buried in
the shell separates this [Pediculariona] distinctly, and
there are representatives of this in Queensland.” This
inadequate diagnosis could be interpreted as a statement
purporting to give characters differentiating the taxon,
but by the same token it is this particular diagnostic
character which places it in the synonymy of Pedicularia
Swainson, 1840.

METHODS

The nomenclature of diagnostic characters of Ovulidae
follows ScHILDER (1932, p. 57, fig. 20). The number of
dorsal ribs quoted for species of Triviidae consists of
the total of centrally-placed ribs on both sides of the
dorsal groove or dorsal centre; the loop-like ribs confined
within the outlets of the extremities have been excluded
from the count. Only mature specimens have been utilized
for morphometric measurements.
The following abbreviations are uscd in this paper:

L = Length of shell expressed in millimeters (mm)
W = Width of shell expressed in % of length

H = Height of shell expressed in % of length

LT = Number of labial teeth

ACKNOWLEDGMENT

I would like to express my appreciation to Mr. D. Heppell,
Royal Scottish Museum, Edinburgh; Drs. H. Rehder
and J. Rosewater, Smithsonian Institution, U. S. National
Museum, Washington; Dr. FA. Schilder, Halle, and Mr.
N. Tebble, British Muscum (Natural History), London,
for their assistance with reference material and other
information. The permission to publish photographs of
the types of Ovulum margarita SowErsy, received from
the Trustees of the British Muscum (Natural History),
London, is gratefully acknowledged.

I wish to thank the following collectors for the loan of
study material: Mrs. M. Ashton, Proserpine; Mr. R. FE
Browne, Nausori; Mr. J. Candrics, Suva; Mrs. FE Fitz-
maurice, Kurrimine Beach; Mr. & Mrs. F Freitag, Suva;
Mr. & Mrs. A. Jameson, Lautoka; Mr. A. Jennings,
Auckland; and Mrs. M. Wood, Nadi.

INDEX or SPECIES

(* denotes synonym or homonym)

CLC) S eee eS OAM C ONT COLUT ane SARS TARR GD Et ensuite reset oats aN Fe 363
adamsonit (Pseudocy prada) crcccucscreove 365 costellata (Ovula ED 5 Onl |pep MLA IVC SU eehee cree ereh crane So age We See 364
PF CLALIT GTO ape cee re eee 365 | * cristalline ...... . 362 hordacea\ (Urvinostra))) osacneccennoetoss 368
* alba SCHUMACHER .. Zh Ob | ca GLUIL ULE Cy wma a he reer tery SOSmlie dy alin ane ee ieee morn ee
* alba SOWERBY ....... 367 | * cygnea ....... . 358 | * imperialis oh
CHV RTOS ecertceater ttre ree eps eae ene B35 Ou Peel CALE CL Oe eeresies meer tances eter eee eed Oia [Mes ICIUSE CED) nance rece Ren te ohm son ee ee
birostris (Phenacovolva) recmescnrrneenein GYOG) || CLGTTAOTES secccreaesecmmmnentcemmmmmicraseemm GIS |) CGM OTTO cceeceeereterererttteee aerator
PS OMCULNOSE 1 Spammer ee edgari (Trivirostra) ; PNUTLECIITC ALC eee
Me COM AUC UL AN erate arrc cases cvcissceenr ieee BCX CCS rane cents 370 Konoensisn(diiuinostra)) mame
PEN CO PETS IS rede ctres Satan catteaeh sesame natin PHEHG (HOOT) erercrrereceeecerececrirrcr 367 lacteus)|(Galpurnus) ences
COT PEILLCT Imre nes fruticum (Prionovolva) . PER LETIVUTICC perenne
coarctata (Prosimnia) gracilis (Phenacovolva) Margarita (PrimMovula) cecmsscssisnisnrsrneres
SENG OUI D a) inner ee re er elaine BR On ee nmonand of memes eae eae eens SOM lb Me MINOT, RAY: iealamsciensncncsceasmnein ee
PMC OMI SAL reat terseenssce etre trnitarrsatienets GAO] || FACUOIRLICE - eceeccireereeteen erro SO 7m le niNOT SCHED MAN petreeatre arene

ee Deliiculayee semistriata

pellucidula (Trivirostra) sem pent
Belay A 1 Harr eater tec ea eem rere 5s sempieri
philippinarum (Phenacovolva) cco 364. smithi

producta (Trivirostra) econ striata LAMARCK

Trivirostra species
turnert

SIU CVC PUTIGLAUO resntancncerernn tase

Page 356 THE VELIGER Vol. 10; No. 4
PANG! bess tenucn encanta a PM eSulcata valiant eee
* natalensis ... * sulcifera (Lachryma) ....
* nectarea * rubida ® SUTADAJENSIS vecseerosesine
Keli TAL been eet eerste PRS CIUCCIUGIUGTISUS meet tcneete » BOS || 9 COORD crrccecrasscrnarcaner
oryza (Trivirostra) * sandwichensis 2 SOS | LODEEOTAE srarroscrmememrco
* oryzoidea GOGH TMISCOWE ccraecerecxrasrncecasrn 2 HORDUI OY: cxcsxvasciaseccseree
EMOUUfONIISIEetrnte * schmeltiana * tortile
ovum (Ovula) ....... Bere * schmeltziana * tremeza ..
pacifica (Pedicularia) ....... * schneider . © CUI. cceresserarm
%
*
*
*
*
*

punctata (Primovula) cecccccencne striata SHIKAMA

POT Dy BINGE Osea etek adetrs Seeeaniatenra eee ne es

MESOGASTROPODA

CYPRAEACEA

OvuuDAE FLEMING, 1828 (nom. corr.
OvuLADAE)

1854. AMPHIPERASIDAE H. « A. Apams, Gen. Rec. Moll., 1: 269
1929. AMPHIPERATIDAE WINKWoRTH, Proc. Malacol. Soc. London,
18: 298

The shells of Ovulidae are pyriform or fusiform in shape,
uni-coloured, spotted or banded, and the cxtremities are
either short or produced; shells are smooth or transversely
striate and some species have a dorsal carina. The labial
side is margined, columellar side rounded, labial lip with
or without denticles, columella gencrally edentulous.

The animals are similar to the Cypracidae, i.e. they
possess a foot, mantle, tentacles, cyes, and a siphon; the
papillae, however, are gencrally shorter. The anatomy of
the Ovulidae has been described by TimeLe (1929) and
SCHILDER (1932, 1939, 1944).

The radula is taenioglossate with 7 tecth per row;
Recent species of Eocypraeinac have an additional edent-
ulous connecting plate. Rhachidians are variable, but
generally have a large central cusp and often side-den-
ticles; laterals are large, with a beak-like main cusp and
generally side-denticles on the cutting edge. Marginals are
fairly uniform throughout the family and the inner
marginals are smaller than the outer ones; flabellae are
numerous, slender, extended, recurved and bifid or some-
times trifid at the distal end. SctmLpER (1932) considered
the anatomy and radula features of Ovulidae to be rather
similar among the genera and subgenera, and his subdivi-
sion has therefore been restricted to differences in shell
morphology. Although the marginals are basically similar

striatula (Primovula) ..
GOWIISIOTOS. racsortmen memrmermannececcr

verrucosus (Calpurnus
UOlWaN QVOLUG) ren nent eee

throughout the family, the rhachidians and sometimes
the laterals exhibit appreciable differences between spe-
cics of different genera; on the other hand, however, the
radular features of some species of Ovulinae may re-
semble those of Simniinae.

The majority of Recent ovulids lives in the Indo-Pacific
region, and only a few spccies live in other major faunal
regions.

Some Ovulidae share the same habitat with cypraeids,
while others are associated with octocorals and sea-fans.
The food-requirements of tropical ovulids are not known;
however, food is possibly extracted from the same host-
coral on which they live.

Ovulinae FLEmine, 1828

1929. Amphiperasinac Turetr, Handb. syst. Weichtierk., p. 270
1932. Amphiperatinae SermtpEr, Proc. Malacol. Soc. London,

20: 46

The subfamily contains a group of species with pyriform,
inflated shells with short extremities, a plain, spotted or
banded dorsum, which is cither smooth, transversely
striate or carinate. Columellar teeth are absent, labial
tecth developed and the antcrior terminal ridge on the
columella is prominent; the fossula is smooth and broad.
The rhachidians and lateral teeth of the radula are vari-
able, marginals with numerous slender flabellae.

Species of Ovulinac are distributed in all major Oceans,
but specics of the genus Pseudosimnia ScuiLpErR, 1927,
are restricted to the Mediterranean and the Eastern
Atlantic.

Specics of Ovulinac inhabit crevices of coral rocks, or
occasionally live in folds or at the base of soft coral or
on gorgonians.

Vol. 10; No. 4 THE VELIGER Page 357
eee WBYadua :
% .Nacula M
2&2 gi
5 Se
5 O
20 ae Noviti e
4 2
Noanonu-! RA
Waya v
| OWaya Loailai oo wellington Wharf
| °° — A Rokiraki litilevu Bay a
; wt Makogai
6 “2 °
Bo LAUTOKA lokaya
MAMANUC a . {i poe
G Pp ee ey , J OVALAU
of odoni YF

19°

TAILEVUsaee

LEV U “ie
Momi fe > BAU
“ a Pe Vi cba F
o 7 Br
NatadolaNee. SIGATOKA -- _ aNasilai
Cu i, Cage ee :Nukulau
- Korotogo
Sov Bay
Korolevu
Nadroga reef
{ £9 BEOA
Yanuca
(varucece E
pe
5°
ph
av _#Dravuni
A 19) 70 ;
K - -Bulia
{9 Ono i
os 2 . KADAVU

x NEw <* 5 LovattY Is
D) CALEDONIA oa
Seen rec PARC ac) ENT IG
AUSTRALIA ‘| ans
“f - NORFOLK |.

ey O56 VEAL AN™
j ve ee iN See Seelt

a _QABERT

PHOENIX |e.

HEBRIDES
.

Vv au le

Scale of Miles

0246810 20 30

178°

ia eit det

Longitude East 179°

Page 358

THE VELIGER

Vol. 10; No. 4

Ovula Brucuieére, 1789

Ovula BrucutkreE, 1789, Hist. nat. vers, Index, p. 15. Type species
by SD (Lamarck, 1801) Ovula oviformis Lamarck, 1801 = Bulla
ovum Linnaeus, 1758.

1781. “Amphiperas” Gronovius, Ind. Zooph. Gron., p. 293
(non binom.)

1798. Ovula Lamarcx, Tabl. Encycl. Méth., plts. 357-358 (no
type selected)

1810. Ovulus MontrFort, Conch. Syst., 2: 634 (nom. nov. pro
Ovula Brucutére, 1789)

1828. Ovulum Sowersy, Zool. Journ. 4: 145 (nom. nov. pro
Ovula Brucutére, 1789) :

1935. Parlictum Irepae, Aust. Zool., 8: 101 (Type species by
OD Ovula costellata Lamarck, 1811)

The genus contains the two largest species of the family
Ovulidae; their shells are smooth or obsoletely carinate,
the interior is dark in colour, the columella is edentulous
and labial teeth are irregular. Only two Recent Indo-
Pacific species are known, and fossil species have so far
not been recorded.

Discussion: The genus Parlictum IREDALE has been estab-
lished for Ovula costellata LAMARCK on the basis of a
difference in animal features and the formation of the
posterior canal. Differences in animal features are specific
characters and the diagnostic difference in the formation
of the posterior canal appears to be of insufficient taxo-
nomic importance to warrant the erection of a new
monotypic genus.

Ovula ovum (Linnaeus, 1758)
(Plate 50, Figure 1)

1758. Bulla ovum Linnaeus, Syst. Nat., ed. 10, p. 725, no. 327

1798. Volva cygnea Rovinc, Mus. Bolt., p. 21, no. 255

1811. Ovula oviformis Lamarck, Ann. Mus. Hist. Nat., 16: 110

1817. Ovula alba ScuumaAcHER, Ess. nouv. syst., p. 258

1828. Ovulum ovum var. pygmaca SowerBy, Zool. Journ..
4: 149

1905. Amphiperas ovum (Linnaeus), Bercu, Siboga Exp.,
50: plt. 5, fig. 21 (animal)

Shell: Large and ovate, extremities produced, porcellanous
white, interior orange-brown; dorsum either moderately
smooth or with 1 - 4 fine transverse carinate lines. Labial
lip convex, teeth irregular and often bifid, numbering
from 35 to 45; columella edentulous, second funiculum
strong and projecting, fossula absent.

L: 65 - 110mm; W: 58 - 64%

Type Locality: Oceano Asiatico (“Amboina,” IREDALE, .
1935))e

Habitat: On soft coral, from 1 - 4 fathoms. Common.

Distribution: Throughout the Fiji Islands. -— From

Madagascar through the tropical Indo-Pacific to the

Tuamotu Archipelago.

Ovula costellata Lamarck, 1811
(Plate 50, Figure 2)

1811. Ovula costellata Lamarck, Ann. Mus. Hist. Nat. 16: 110

1817. Ovula imperialis Dittwyn, Descr. cat. rec. shells 1: 470

1822. Ovula angulosa Lamarck, Anim. sans vert.. 7: 367

1829. Ovula columba ScuuBeERt & WAGNER, Conch. Cab., 12:
116; plt. 228, figs. 4043, 4044

Shell: Pyriform and inflated, white in colour, centre of
dorsum occasionally pink, interior violet; dorsum with an
obsolete dorsal carina, transverse carinate lines and striae
at extremities. Labial lip convex, teeth irregular, num-
bering from 19 to 23; columella edentulous, occasionally
with 3-4 obsolete columellar teeth, anterior terminal
ridge obsolete, fossula absent.

L: 38-42mm; W: 62-65%

Type Locality: L’Océan des Grandes Indes (“Friendly
Isles,’ Martyn, 1886) [= Tonga Islands].

Habitat: On soft coral, from 1-4 fathoms. —- _ Rare.
Distribution: Southwest Viti Levu. — From East Africa
through the tropical Indo-Pacific to Japan and the Tonga
Islands.

Calpurnus Montrorrt, 1810

Calpurnus Montrort, 1810, Conch. Syst., 2: 638. Type species by
OD Bulla verrucosa LiNNaEus, 1758.
1840. Cypraella Swainson, Treat. Malac., p. 325 (Type spe-
cies by M Bulla verrucosa LinNaEus, 1758)

Shells of the genus are moderately small, angulately pyri-
form, smooth or striate, and white in colour; the dorsal
carina is prominent, obsolete or absent, extremities with
or without a yellow-ringed wart-like knob. Columella is
edentulous, labial lip prominently denticulate.

The radula (fide ScHEPMAN, 1909) has rhachidians
which carry a massive central cusp which protrudes over
the concave plate margin; one small accessory denticle is

Explanation of Plate 51

Figure 11: Phenacovolva gracilis (ADAMS & REEVE). Fiji. (x 2.5)
Figure 12: Phenacovolva philippinarum (Sowerby). Fiji. (x 1.3)
Figure 13: Host coral of Phenacovolva philippinarum. Fiji. (x 1.0)
Figure 14: Pseudocypraea adamsonii (SowERBY). Fiji. (x 5.0)

Figure 19:

Figure 15: Trivirostra oryza (LAMaRCK). Fiji. (x 4.0)

Figure 16: Trivirostra species (= ? T: scabriuscula Gray). Fiji. (x 5)
Figure 17: Trivirostra edgari (SHaw). Fiji. (x 4.5)

Figure 18: Trivirostra exigua (Gray). Fiji. (x 6.5)

Trivirostra hordacea (KiENER). Fiji. (x 8.0)

Tue VE.icER, Vol. 10, No. 4 [CeRNOHORSKY] Plate 51

Figure 11 Figure 12 Figure 13 Figure 14

Figure 15 Figure 16

Figure 17 Figure 19

photographs by W. O. CeRNOHORSKY

Vol. 10; No. 4

positioned at either side of the cusp; laterals are simple
and curved, and a small denticle is positioned centrally on
the main cusp; marginals are similar to those of other
groups of Ovulidae.

Species of Calpurnus are associated with soft coral.

Calpurnus (Calpurnus) verrucosus (LINNAEUS, 1758)
(Plate 50, Figures 3, 3a)

1758. Bulla verrucosa LLNNAEUS, Syst. Nat., ed. 10, p. 726,
no. 330

1798. Volva perla Ropinc, Mus. Bolten., p. 22, no. 260

1909. Calpurnus verrucosus (LINNAEUS) SCHEPMAN, Siboga
Exp., 49b: 146; plt. 15, fig. 10 (radula)

- 1935. Calpurnus verrucosus (LINNAEUS) IREDALE, Aust.

Zool., 8: 102; plt. 8, figs. 4, 4a (animal)

Shell: Angulately ovate and humped, white in colour,
extremities flushed with light rose-purple, ornamented
with 2 prominent orange-ringed wart-like tubercles. Dor-
sum with a prominent central transverse carina and
numerous fine striae; sides minutely granulose, granules
extending partly towards dorsum. Aperture dilated ante-
riorly, labial lip flattened and with 17-23 prominent
teeth which extend towards the margins; columella eden-
tulous, columellar anterior ridge indistinct, fossula shallow
and smooth.
Animal: Foot and siphon are white in colour, mantle is
smooth, white to greyish-white, spotted with small pink
spots over entire mantle surface.
L: 18-27mm; W: 59-67%
Type Locality: In India Orientali (“Amboina,” IREDALE,
ISB).
Habitat: In folds and at the base of soft coral, in shallow
and deeper water. ~ Moderately common.
Distribution: Throughout the Fiji Islands. -— From
Madagascar through the tropical Indo-Pacific to Japan
and the Fiji Islands.
Discussion: In the majority of Fiji specimens examined,
the anterior extremity is partly recurved towards the
aperture.

(Procalpurnus) THIEE, 1929

Procalpurnus Tutete, 1929, Handb. syst. Weichtierk., p. 272. Type
species by M Ovula lactea Lamarck, 1811.

The subgenus, which is monotypic, contains a species
which lacks the wart-like nodes at extremities and central
dorsal carina. The rhachidians of the radula of the type
species of Procalpurnus are also dissimilar to those of the
type species of Calpurnus.

THE VELIGER

Page 359

Calpurnus (Procalpurnus) lactcus (LaMarRcK, 1811)
(Plate 50, Figure 4)

1811. Ovula lactea LAMarcx, Ann. Mus. Hist. Nat., 16: 111

1863. Amphiperas scmistriata PEASE, Proc. Zool. Soc. London
for 1862: 241 (“Pacific Islands”)

1935. Procalpurnus lactcus (LAMARCK), IREDALE, Aust. Zool.
8: 103; pl. 8, figs. 3, 3a (animal)

1965. Amphiperas semistriata PEASE, Kay, Bull. Brit. Mus.
(Nat. Hist.) Zool. Suppl. 1: 74; plt. 12, figs. 5, 6 (lecto-
type)

Shell: Elongate-ovate to broadly ovate, white in colour;
the dorsal carina is obsolete in most specimens, but some
specimens show 1 or 2 weak carinae. Dorsum with 20 - 50
transverse striae which are distributed along the entire
length of the shell but may become obsolete centrally and
confined to the extremities. Labial lip is flattened, teeth

“A gage
oY

DP apr

Figure 1

Calpurnus (Procalpurnus) lacteus (LAMARCK)

Fiji Islands
a — Half row of radular teeth
b — Rhachidian of radula (ventral view) c — (dorsal view)

100 p

We

Figure 2

Calpurnus (Procalpurnus) lacteus (LAMARCK)

Pearl Reef, Queensland, Australia
Rhachidian and lateral teeth of radula

Page 360

THE VELIGER

Vol. 10; No. 4

prominent, numbering from 20 to 31; columella edentu-
lous, anterior columellar terminal ridge prominent and
oblique, fossula smooth.

Animal: Foot blackish-grey, mantle smooth, jet-black
and ornamented with small white spots; tentacles black,
siphon black and finely ciliated at the distal end.
Radula: The odontophore is 4.6mm long and 0.68mm
wide in an animal with a shell 17.4mm in length; fully
formed rows of teeth number 138 (plus 14 nascentes)
and early rows of teeth are greatly worn. The rhachidian
is small, 0.09 mm wide, with a long central cusp and 3 - 9
side-cusps; laterals are large, with a massive main cusp
and 3 - 4 side denticles. Inner marginals have ca. 18 and
outer marginals ca. 38 flabellae which are curved and
bifid at the distal end.

A specimen from Queensland had a radula 3.9mm
long and 0.7 mm wide and a shell 17.8mm in length; the
rhachidians were 0.1 mm wide and the ribbon contained
132 rows (plus 9 nascentes) of teeth.

L: 12- 19mm; W: 54 - 63%

Type Locality: Timor.

Habitat: At the base of soft coral, from 0-4 fathoms.
Uncommon.

Distribution: Throughout the Fiji Islands. -— From

the Red Sea through the tropical Indo-Pacific to the

Society and Hawaiian Islands.

Prosimnia SciuLpER, 1927

Prosimnia Scuivper, 1927, Arch. Naturgesch. 91A: 77. Type species
by OD Ovula semper’ WEINKAUFF, 1881 = Ovulum coarc-
tatum ApdAMS & REEVE, 1818.

The shell of the type species is clongate-cylindrical, the
dorsum is carinate and spirally striate, labial lip densely
denticulate, columellar lip with short denticles on axis,
fossula narrow and denticulate.

The radula of the type species has been described by
ScHILDER (1944, p. 32) and was said to be somewhat
similar to the radula of Pscudosimnia carnea (PoireET,
1789) as figured by Tire (1929, fig. 287). The radula
of Primovula becker (SowrrBy, 1900) differs appreci-
ably from that of Prosimnia coarctata (vide BARNARD,
19637 ps oO Migs Ore).

Prosimnia coarclata (AbAMS & REEVE, 1848)
(Plate 50, igure 7)

1848. Ovulum coarctatum ADAMS & REEVE, Zool. Voy. Sam.,
p. 21; plt. 6, figs. 2a, 2b

1881. Ovula sempicri WetnkaurF, Conch. Cab., ed. 2, 5: 190;
plt. 48, figs. 14, 15

1882. Ovula sempert WEINKAUFF, Jahrb. Deut. Malakol. Ge-
sellschaft, p. 174 (nom. corr.)

194-4. Primovula coarctata (ADAMS & REEVE), SCHILDER,
Ark. Zool., 36A: 32 (descr. radula)

Shell: Small, slender and cylindrical, orange-brown or
orange-fawn in colour, labial margin and labial lip bright
yellowish-orange. Dorsum sculptured with a prominent
clevated dorsal keel and occasionally 1 - 2 smaller keels;
transverse striae vary in thickness and number from 55 to
70 and are intersected by longitudinal growth striae.
Labial lip flattened and curved, teeth sharply sculptured
but irregular and numbering from 43 to 53, partly ex-
tending over the shell-margin; columella denticulate al-
most along entire length to funiculum, teeth small, num-
bering from 45 to 50; anterior columellar terminal ridge
is absent, first funiculum is crenulate, second funiculum
prominent, fossula narrow and projecting, smooth and
denticulate.
Animal: Foot light orange, mantle orange to red, siphon
and tentacles reddish-orange, eyes black.
L: 11-13mm; W: 31 - 36%
Type Locality: Straits of Sunda, near Java.
Habitat: On red gorgonian coral, from 2-3 fathoms.
Moderately rare.
Distribution: South and West Viti Levu. — From Mada-
gascar through the tropical Indo-Pacific to Japan and the
Fiji Islands.

Primovula Tutee, 1925

Primovula Tine e, 1925, Kickentuat, Handb. Zool., 5: 88. Type
species by OD Amphipceras beckerii Sowersy, 1900.
1961. Dentiovula Hae, Col. illust. shells Japan, 2: 41; plt. 19,
fig. 1 [Type species by M D. dorsuosa (Hinps, 1844) ]
1961. Dentivolva Hae, Col. illus. shells Japan, 2: App. p. 14
[Type species by OD D. dorsuosa (Hinps, 1844) ]
1963. Dentiovula Hase, Su1kama, Sel. shells world, 1: 45; plt.
32, fig. 15 (first reviser)

Shells are small, cylindrical-fusiform, occasionally with a
light-coloured central band, a dorsal carina and striae;
labial lip flattened, prominently or irregularly denticulate,
columella edentulous, labial margin often serrate.

The radula (fide BarNnarp, 1963) has horizontally
egg-shaped rhachidians with a short and stout central
cusp and 3-4 side denticles; laterals are moderately
slender with a main cusp and 4 - 5 accessory denticles on
cutting edge; marginals similar to those of other genera
of Ovulidae.

Discussion: Hase (1961) established Dentiovula on page
41 and later on, on Appendix page 14 described Denti-

Vol. 10; No. 4

THE VELIGER

Page 361

volva in Japanese. In accordance with art. 32b of the
Code of ICZN (1964), Dentiovula is the correct spelling
as selected by SH1kaMA (1963), who appears to be the
first reviser. The new genus name, however, seems
superfluous.

Primovula (Primovula) striatula (SowERBy, 1828)
(Plate 50, Figure 8)

1828. Ovulum striatulum Sowersy, Zool. Journ., 4: 155

1830. Ovulum striatulum SoweErBy, Spec. Conch., Ovulum,
1: 7, fig. 38

1848. Ovulum dentatum ApaMs & REEvE, Zool. Voy. Sam., p.
21; plt.6, figs.4a,4b (non Ovula dentata Fiscuerr, 1807)

1930. Prosimnia renovata IrREDALE, Mem. Qld. Mus., 10: 85
(nom. nov. pro Ovula dentata ApaAMs & Reeve, 1848)

1932. Prosimnia verconis Corton & GODFREY, South Aust. Nat.
13: 46; plt. 1, fig. 15

1963. Primovula striata (sic) SowERBY, SHIKAMA, Sel. shells
world, 1: 45

1963. Neosimnia tinctura GARRARD, Journ. Malacol. Soc. Aust.
7: 45; pit. 7, figs. 5, 6

Shell: Small, cylindrically-fusiform, dark pink in colour
with one white central transverse band and pale pink
margins and extremities; the dorsum is sculptured with
50 - 60 fine transverse striae and a dorsal carina which
may be obsolete. Labial lip is flattened, labial teeth some-
times obsolete anteriorly but reaching margins posteriorly,
numbering from 25 to 28; in some specimens the labial
teeth are very large and reach as far as the outlets. Anteri-
or columellar terminal ridge is obsolete, first funiculum
with 3-5 denticles, second funiculum pronounced or in-
distinct, interior of columella finely striate, fossula exca-
vate and smooth.
L: 7-9mm; W: 35 - 39%
Type Locality: Ad littora Oceani Indici; East Indies
{= Indonesia].
Habitat: Dredged from 15 - 18 fathoms on coral. Rare.
Distribution: West off Viti Levu. -— From the Persian
Gulf through the tropical Indo-Pacific to Japan and the
Fiji Islands.
Discussion: Fiji specimens are more slender in comparison
with specimens from other regions (W: 41 - 49%). The
species is extremely variable in colour, some specimens
being purplish-pink or purple; for further notes on the
species see SCHILDER, 1964.

(Diminovula) IrREDALE, 1930

Diminovula IrEDALE, 1930, Mem. Qld. Mus., 10: 85. Type species
by OD D. verepunctata IrEpDALE, 1930 = Ovula punctata
Duc os, 1831

Ai
eA

1935. Margovula IrEpALE, Aust. Zool., 8: 103 (nom. nud.)
1939. Margovula Scuitprr, Arch. Molluskenk., 71: 200 (Type
species by OD M. pyriformis (SowERBy, 1828) )

Species of the subgenus are characterized by their small
pyriform shells and somewhat produced extremities; the
dorsum is smooth or striate, unicoloured or spotted, labial
teeth are developed, columellar teeth obsolete; first funi-
culum distinct, crenulate or smooth, fossula smooth and
concave.

The radula of the type species was not available, but
the radula of Diminovula bimaculata (A. Apams, 1855)
from Queensland was examined; this species has essen-
tially similar morphological features as the type species,
and has been assigned by Scumper (1941) and other
authors to Diminovula. The radular ribbon was 6.6mm
long and 0.9mm wide in an animal with a shell 13.5mm
in length; the ribbon contained 111 rows of teeth (plus
12 nascentes) and early rows of teeth were greatly worn.
The rhachidian is small, 0.17mm wide, with a long
central cusp and 2 - 3 cusps at either side; laterals large
and unicuspid, inner marginals slender and with ca. 12
flabellae, outer marginals broad and with 21 - 23 flabellae.

The radula is basically similar to the radula of Ovula
angasi Rerve, 1865, the type species of Pellasimnia (vide
ScHEPMAN, 1909), in the subfamily Simniinae; it also
resembles the radula of Primovula beckerii (SowERBy,
1900) in the subfamily Ovulidae. The laterals are unicus-
pid in Diminovula bimaculata, but denticulate on the
cutting edge of the main cusp in Pellasimna angasi and
Primovula becker; this feature, however, is a variable
characteristic in other taenioglossate radulae, e.g. in
Cymatiidae and Bursidae.

——

pg

Pape

Figure 3
Primovula (Diminovula) bimaculata (A. ADAMS)

a — Half row of radular teeth
b — Distal end of flabella (enlarged)

Page 362

THE VELIGER

Vol. 10; No. 4

Primovula (Diminovula) punctata (Ductos, 1831)
(Plate 50, Figure 5)

1831. Ovula punctata Ductos, Mag. de Zool., 1: 7; plt. 7,
fig. 1

1930. Diminovula verepunctata TREDALE, Mem. Qld. Mus.,
10: 85

1932. Primovula (Diminovula) cristallina (K1ENER), ScHIL-
DER, Proc. Malacol. Soc. London, 20: 51; plt. 3, figs. 2, 3,
4 (? non Krener, 1843)

Shell: Small and pyriform, extremities slightly produced;
white to creamy-white in colour, generally ornamented
with 3 pairs of white spots on dorsum. Some specimens
have additional orange transverse lines descending onto
the margins from the anterior and posterior pair of
spots, and an orange line follows the periphery of the
labial margin; extremities are tinged with light yellow.
The dorsum is sculptured with 25-60 fine transverse
striae; labial lip is convex, labial teeth are irregular but
prominent, numbering from 19 to 28. The anterior ter-
minal ridge is well developed, columella edentulous, first
funiculum projecting and rounded at end, sculptured with
4-7 irregular denticles; first posterior outlet is shallow,
second funiculum and outlet are obsolete. Fossula is con-
cave and smooth, interior of columella smooth or striate.
L: 6-11mm; W: 57-65%
Type Locality: I. Bourbon [= Réunion Island, Indian
Ocean]
Habitat: Under coral rocks, in shallow and deeper water.
Moderately rare.
Distribution: North and Southwest Viti Levu. — From
East Africa through the tropical Indo-Pacific to Japan
and the Fiji Islands.
Discussion: The Primovula cristallina of ScHILDER
(1932) is conspecific with P punctata. The holotype of
Ovula cristallina KiENER, preserved in the Paris Museum,
is a worn and calcified Diminovula species (fide ScHIL-
DER, 1932) ; in view of the bad preservation of the holo-
type it is doubful whether Ovula cristallina is a good
species or conspecific with Primovula punctata.

Prionovolva IREDALE, 1930

Prionovolva IrEDALE, 1930. Mem. Qld. Mus., 10: 85. Type species
by OD Ovulum breve SoweErBy, 1828
1941. Prionovola (sic) ScuimtpER, Arch. Molluskenk. 73: 117
1964. Prionovula (sic) CERNOHORSKyY, The Veliger 6: 200

The genus contains ovate to pyriform shells which are
either banded or unicoloured; the dorsum is smooth, but
spirally striate at extremities, labial teeth are irregular
but prominent, columella is edentulous. The genus differs

from Primovula in the prominently projecting smooth
funiculum and columellar carina.

Prionovolva fruticum (REEVE, 1865)
(Plate 50, Figure 6)

1865. Ovulum fruticum Reeve, Conch. Icon., Ovulum, 15,
plt. 4, sp. 16a, 16b

Shell: Elongate-ovate, white to pinkish-white in colour,
ornamented with 3 - 4 dark pink transverse bands; mar-
gins and base are white. The dorsum lacks a dorsal
carina, and is sculptured with 8-30 transverse striae
which are generally confined to the extremities; labial
lip is flattened, teeth sharply sculptured, numbering from
19 to 25; some of the labial teeth (ca. 2-7) extend
towards the margin which becomes serrate. Aperture
dilated anteriorly, columella edentulous and with a sharp
columellar carina, anterior columellar terminal ridge
prominent; first funiculum prominent, smooth and pro-
jecting, second funiculum absent, fossula short and
concave.
L: 10- 15mm; W: 52-59%
Type Locality: Malacca.
Habitat: Dredged from 10 - 18 fathoms on coral.
Moderately rare.
Distribution: West Viti Levu. -— From East Africa
through the tropical Indo-Pacific to Japan and the Fiji
Islands.
Discussion: The dorsal spiral striae are generally confined
to the extremities, but may extend across the dorsum,
becoming somewhat obsolete centrally. One specimen
showed 3 distinct dorsal transverse breaks, while another
was rather angulate and had orange extremities and a
marginal peripheral line.

Simniinae ScHILDER, 1927

1932. Volvinae ScuitpeEr, Proc. Malacol. Soc. London, 20: 47
(as Volvini)

1956. Volvinae ALLAN, Cowry shells of world seas, p. 126

1958. Volvinae Corron, Journ. Malacol. Soc. Austral., 2: 11

Species of Simniinae have elongate-fusiform and de-
pressed shells with produced extremities, shells are uni-
coloured, occasionally with a median band, smooth or
transversely striate; a dorsal carina is obsolete or absent.
The columella lacks the anterior terminal ridge, labial
teeth are obsolete or absent, columella edentulous, fossula
either narrow or absent. The radula is similar to that of
the Ovulinae.

Species of Simniinae are associated with reef-dwelling
gorgonians and alcyonarians, and sometimes adopt the

Vol. 10; No. 4

THE VELIGER

Page 363

colour of the host coral; the majority of species is
confined to deeper water.

Volva Ropine, 1798

Volva Ropinc, 1798, Mus. Bolten., p. 21. Type species by T Bulla
volva LinNAEuS, 1758.
1810. Radius Montrort, Conch. Syst., 2: 626 (Type species
by OD Bulla volva Linnatus, 1758)
1840. Birostra Swainson, Treat. Malacol., p. 325 (Type spe-
cies by OD Bulla volva LinnaEus, 1758)

Shells are pyriform, inflated, extremities greatly produced,
dorsum spirally striate; labial lip convex, labial teeth
absent or obsolete, columella edentulous, fossula absent.

The radula of the type species (fide BARNARD, 1963)
has roundly pentagonal rhachidians with a moderately
short central cusp and a small accessory denticle at either
side; the laterals have a large main cusp and one side
cusp on the cutting edge, and a large and slender basal
peg; marginals comb-like and similar to those of other
genera of Ovulidae.

Volva volva (Linnaeus, 1758)
(Plate 50, Figures 9, 9a)

1758. Bulla volua Linnaeus, Syst. Nat., ed. 10, p. 725, no. 328

1798. Volva textoria ROp1nc, Mus. Bolten., p. 22, no. 259

1811. Ovula striata LamMarcx, Ann. Mus. Hist. Nat., 16: 113
(“Coast of Brazil” — erroneous)

1931. Volva volva cumulata TrEDALE, Rec. Aust. Mus., 18: 222
(nom. nud.)

1935. Volva volva cumulata IrEDALE, Aust. Zool., 8: 104

1937. Volva volva surabajensis ScuiLpErR, Ing. Ned. Indie, 4:
205 (fossil from Soerabaja)

1941. Volva (Volva) volva lemurica ScuitpER, Arch. Mollus-
kenk. 73: 110 (nom. nov. pro V, volva var. MELVILL, 1909)

1961. Volva volva habei Oyama, Venus : Japan. Journ. Mala-
col., 21: 288; text figs. 3, 4

1963. Volua volva (LinNAEUS), BarNarD, Ann. South Afr.
Mus., 47 (1): 56; fig. 6d (radula)

Shell: Large and fusiform, inflated centrally, extremities
produced into long and slender canals; pinkish-white in
colour, margins white, interior of aperture yellowish-
brown. Sculptured with ca. 80 transverse striae which
become obsolete centrally on dorsum; outer lip convex,
ornamented with ca. 14 very obsolete blunt denticles,
columella edentulous. Anterior terminal ridge, first funi-
culum and posterior outlet absent, second funiculum
projecting, fossula absent.

Juvenile shells are white in colour, fragile, extremities
are greatly produced and slender, and the transverse
striae appear as distinct elevated cords on the dorsum.

L: 77-85mm; W: 28 - 30%

Type Locality: Ad Jamaicam (Error) [“Ceylon,”’ Ire-
DALE, 1935]

Habitat: In 15 fathoms, on coral rubble and sand sub-

stratum. Rare.

Distribution: North and West Viti Levu. — From East

Africa through the tropical Indo-Pacific to Japan and the

Fiji Islands.

Phenacovolva IrEDALE, 1930

Phenacovolva Irepaue, 1930, Mem. Qld. Mus., 10: 85. Type species
by OD P. nectarea IrREDALE, 1930 = Bulla birostris LINNAEUS,
1767

1817. Radius ScHuMACHER, Ess. nouv. syst.. p. 259 (Type spe-
cies by M R. brevirostris ScHuMACHER, 1817 = Bulla
birostris LiNNAEUS, 1767) [non Montrort, 1810]

1956. Phenacolepas (sic) IREDALE, ALLAN, Cowry shells of
world seas, p. 134 [non Pirssry, 1891]

Shells are moderately small, slender and fusiform, with
shorter extremities than in Volva s. str.; dorsum smooth
or striate, unicoloured or with a light coloured median
band. Labial and columellar lips edentulous, fossula
absent.

Theradulaof Phenacovolva aurantia (SowErRBy, 1889)
[=? RP birostris (Linnaeus, 1767) or P sowerbyana
(WernkAurF, 1881)] as figured by BarnarD (1963), is
dissimilar to the radula of Volva s. str. The rhachidians
are triangular, the central cusp is solid but short and
flanked by moderately solid side cusps; the laterals and
marginals are similar to those of Volva volva (LINNAEUS).

Phenacovolva contains a compact group of Recent
species which appear to be separable from Volva on the
basis of shell morphology and radula features.

Phenacovolva birostris (LINNAEUS, 1767)
(Plate 50, Figure 10)

1767. Bulla birostris LinNaEus, Syst. Nat., ed. 12, p. 1182, no.
371

1817. Radius brevirostris ScHUMACHER, Ess. nouv. syst., p. 259

1855. Volva rosea A. ApAMs, Proc. Zool. Soc. London for
1854: 130; plt. 28, fig. 9

1930. Phenacovolva nectarea IREDALE, Mem. Qld. Mus. 10:
85; plt. 9, fig. 6

Shell: Fusiform and rostrate, dark pink in colour, orna-
mented with a white median transverse band; margins
white, extremities brown. Dorsum without a carina, sculp-
tured with ca. 50 transverse striae across dorsum; aper-
ture dilated anteriorly. Labial lip flattened and curved,
teeth very obsolete and numbering ca. 20 in the Fiji
specimen; anterior columellar terminal ridge absent, first
funiculum obsolete, second funiculum projecting and

Page 364

THE VELIGER

Vol. 10; No. 4

sculptured with 4 denticles. Interior of columella finely
striate, second posterior outlet prominent, fossula absent.
L: 20.0mm; W: 22%

Type Locality: Ad Javam.

Habitat: Dredged in 15 fathoms on coral. — Rare.
Distribution: From Indonesia to Japan and the Fiji
Islands.

Discussion: ScHILDER (1966a) pointed out that the type
specimen of Linnaeus’ Bulla birostris is the same species
as Radius brevirostris SCHUMACHER, and that the Volva
birostris auctt. is the species V. longirostrata (SOWERBY,
1828). The presence of both species among the types of
Bulla birostris in the Linnean collection was responsible
for the confusion.

The species is known from Fiji from one complete and
one partially broken specimen; both, however, are more
slender than the usual specimens of Phenacovolva birost-
ris, and are also striate across the dorsum along its entire
length.

Phenacovolva gracilis (ADAMS & REEVE, 1848)
(Plate 51, Figure 11)

1848. Ovulum gracile ApaMsS & REEvE, Zool. Voy. Sam., p.
22; pit. 6, figs. lla, 11 b, 11c

Shell: Fusiform and rostrate, pinkish-white in colour,

generally with a white median transverse band; an orange

coloured line encircles the margins. Dorsum sculptured

with ca. 70 transverse striae which become somewhat

obsolete centrally. Labial lip convex and curved, orna-

mented with 15 very obsolete denticles; first funiculum

absent, second funiculum projecting, sculptured with 5

denticles; interior of columella finely striate, fossula

absent.

L: 20.0mm; W: ? 26%

Type Locality: East coast of Borneo.

Habitat: Dredged in 15 fathoms on coral rubble substrate.
Rare.

Distribution: West off Viti Levu. — From Indonesia to

Japan and the Fiji Islands.

Discussion: A small portion of the extremities in the Fiji

specimen is missing and the width ratio of 26% appears

therefore somewhat inflated. This species is rather similar
to Phenacovolva birostris and differs mainly in colouring,
the convex labial lip and more dilated aperture. SCHILDER
(1932) states that the labial lip of P gracilis is edentulous,
but in the Fiji specimen the labial denticles are obsolete,
but can nevertheless be counted.

(Pellasimnia) (IREDALE, 1931, nud.)
ScHILDER, 1939

Pellasimnia ScuitpeR, 1939, Arch. Molluskenk. 71: 195. Type
species by OD Ovulum angasi REEvE, 1865
1931. Pellasimnia IrEDALE, Rec. Aust. Mus., 18: 222 (nom.
nud.)
1939. Pellasimnia ScuitperR, Arch. Molluskenk., 71: 195 (first
valid description)

Shells of the subgenus are fusiform with pointed extremi-
ties, smooth and lack a dorsal carina; dorsum is unicol-
oured, extremities with or without dark stains. Labial lip
reflexed, teeth generally obsolete, columella edentulous,
fossula narrow.

The radula of the type species (fide ScHEPMAN, 1909)
differs in features from those of other genera of Simniinae.
The rhachidians are somewhat quadrate, the central cusp
is broad and long, and is flanked by twoaccessory denticles
at either side.

Phenacovolva (Pellasimnia) philippinarum
(SowersBy, 1849)

(Plate 51, Figure 12)

1849. Ovulum philippinarum SoweErsy, Proc. Zool. Soc. Lon-
don, 16: 136

1877. Volva carpenteri DuNKER, Malakol. Blatter, 24: 75

1877. Volua adamsii DuNxKER, Malakol. Blatter, 24: 75

1882. Radius carpenteri DuNKER, Ind. Moll. Mar. Jap., p.
102; plt. 13, figs. 1, 2

1882. Radius adamsii DunxER, Ind. Moll. Mar. Jap., p. 102;
pit. 13, figs. 3, 4

1889. Ovulum (Birostra) haynesi SowErsy, Journ. Lin. Soc.
London, 20: 397; pit. 25, figs. 1, 2

1909. Amphiperas (Radius) philippinarum (SoweERBy),
ScHEPMAN, Siboga Exp., 49b: 144; plt. 15, fig. 6 (radula)

Explanation of Plate 52

Figures 20 and 20a: Trivirostra producta (Gasxoin). Fiji.
(x 3.2 and 3.0, respectively)
Figure 21: Trivirostra pellucidula (REEVE). Fiji. (x 5.0)
Figure 22: Lachryma sulcifera (SowERBY). Fiji, granulose speci-
men. (x 8.0)
Figure 22a: Lachryma sulcifera (SowErsy). Fiji, smooth speci-
men (x 8.0)

Figure 23: Primovula margarita (SowERBY). Holotype, B. M. N. H.
Reg. No. 1967613/1. Length: 13.8mm (holed).

Figure 24: Primovula margarita (SowERBY). Paratype B. M.N.H.
Reg. No. 1967613/2. Length 8.2mm

Figure 25: Primovula margarita (SowERBY). Paratype B. M.N.H.
Reg. No. 1967613/3. Length 12.1 mm (holed).

Tue VELIGER, Vol. 10, No. 4 [CeRNoHOoRSKY] Plate 52

Figure 20 Figure 20 a Figure 21

Figure 21 a Figure 22 Figure 22 a Figure 23

Figure 24 Figure 25

photographs by W. O. CERNOHORSKY

Vol. 10; No. 4

Shell: Elongate-fusiform, striate at extremities, dorsal

carina absent; dorsal colour salmon-pink, labial margin

light pink, basal colour pinkish-white, extremities dark

brown. Labial lip flattened and smooth, anteriorly some-

what recurved, anterior columellar terminal ridge and

first funiculum absent, second funiculum moderately pro-

jecting, second posterior outlet prominent, fossula absent.

L: 33.0mm; W: 21%

Type Locality: Philippines.

Habitat: On small, black-spotted coral, in shallow water.
Rare.

Distribution: South Viti Levu. -

Japan and the Fiji Islands.

From Indonesia to

Eocypraeinae ScHILDER, 1924

1929. Jenneriinae THIELE, Handb. syst. Weichtierk., p. 269
1939. Sulcocypraeinae ScuiLpeER, Arch. Molluskenk., 71: 191

Species of Eocypraeinae are characterized by pyriform
to ovate shells which are either smooth, transversely
ribbed or pustulose; teeth on both lips are well developed,
aperture is narrow, columella is ungrooved and the fossula
is smooth.

The radulae of living species differ appreciably from
those of other families of Cypraeacea. The laterals are
broad, with numerous long and slender cusps; marginals
are slender and have fewer cusps than have species of
Ovulinae. There is an additional smaller edentulous plate
which connects the laterals with the marginals; a similar
connecting plate is present in the radula of Pediculariidae.

The subfamily contains numerous fossil species but only
2 living relics, i. e. the Indo-Pacific Pseudocypraea adam-
soni (SoweErBy, 1832) and the West American C'ypro-
pterina pustulata (Licutroot, 1786). Living species of
Eocypraeinae inhabit crevices and the underside of coral
rocks.

Pseudocypraea SCHILDER, 1927

Pseudocypraea ScuiLvErR, 1927, Arch. Naturgesch., 91A: 71. Type
species by OD Cypraca adamsonii SowERBy, 1832

The genus is monotypic and contains only the one Recent
Indo-Pacific species.

Pseudocypraea adamsoni (SOWERBY, 1832)
(Plate 51, Figure 14)

1832. Cypraca adamsonii SowErBy, Conch. Illust., Cat. Cyp.,
p- 11, figs. 7, 7a

1879. Cypraeovula adamsonii Gray, GaRRETT, Journ. Conch.,
2: 121 (animal description)

THE VELIGER

Page 365

1929, Pseudocypraea adamsonii (SowERBy), TuH1ELE, Handb.
syst. Weichtierk., p. 270, fig. 285 (radula)

1961. Pseudocypraea adansoni (sic) (SowERBy), Hae, Col.
illust. shells Japan, 2: 41; plt. 19, fig. 2

Shell: Small and pyriform, whitish in colour, ornamented
with 4-5 brown spots on dorsum which are generally
arranged in 3 zones, and orange-brown transverse streaks
on labial margin. Dorsum sculptured with 35 - 40 coarse
and flat dorsal transverse cords. Columellar margin
rounded, aperture narrow, base convex, labial teeth
strong and produced to margin, numbering from 19 to
23; columellar teeth thickened at aperture, extending to
margin, numbering from 18 to 20, interior of columella
striate.

L: 8- 10mm; W: 58 - 62%

Type Locality: Mauritius.

Habitat: Under coral rocks, in shallow water. — Rare.
Distribution: South Viti Levu. — From Mauritius
through the tropical Indo-Pacific to Japan, Fiji and the
Tuamotu Archipelago.

Discussion: ScHiLper (1941) does not report the species
from the Indian Ocean. I have seen a specimen from
Mauritius (coll. E. Couacaud) which appears to confirm
the correctness of SoweErBy’s type locality.

PEepicuLARMDAE H. «& A. Apams, 1854

The family contains a group of species with small cap-
shaped, unicoloured shells with a flaring margin, striate
dorsum with minutely granulose interstices, sharply edged
lips and a projecting or covered spire. Juvenile shells are,
however, subglobular with a projecting spire and denti-
culate lips.

The radula is similar to that of Eocypraeinae (Ovuli-
dae) ; the rhachidians are roughly rectangular, the central
cusp is moderately long and flanked by 4 - 7 side denticles;
laterals are strong, main cusps large, with 3 - 4 denticles
on the cutting edge; marginals are slender and have 2 - 4
finger-like cusps and a shorter claw-like cusp at either
side.

Pediculariidae are sessile on stylasterid corals, and their
shell-margins correspond to the general outline of the
host coral on which they live, and also approximate it in
colouring. Representatives of Pediculariidae are found in
all major seas.

Discussion: Swainson (1840) established Pedicularia
with the Mediterranean P sicula Swainson, 1840, as
type species. Pediculariella THIELE, 1925, was created for
the Californian species Pedicularia californica NEwcoms,
1864. IREDALE (1935) introduced Pediculariona for the
Australian species Pedicularia stylasteris Hepitry, 1903,

Page 366

THE VELIGER

Vol. 10; No. 4

because the shells “superficially differ.” Pediculariidae
contain only 10 species, some being possible synonyms;
they are so similar that some species are being separated
on the basis of the number of striations. SCHILDER (1944)
found the radula of the Pacific Pedicularia pacifica
Pease, 1865, to be similar to the radula of the Mediterra-
nean P. sicula Swatnson, as figured in THIELE (1929).
The shells, anatomy and radula features of the 10
Recent species of Pediculariidae are all so basically alike
that one would tend to question the taxonomic value of
2 genera created on a geographic basis alone. So far
only one fossil species has been recorded: PR. deshayesiana
SEGUENZA, 1865.

Pedicularia Swatnson, 1840

Pedicularia Swainson, 1840, Treat. Malac., pp. 244, 357. Type
species by M P. sicula Swatnson, 1840

1844. Thyreus Pupp, Enum. Moll. Sic. 2: 92 (Type species
by M T: paradoxus Puiuiprt, 1844 = Pedicularia sicula
Swarnson, 1840)

1863. ? Dentiora PrasE, Proc. Zool. Soc. London, p. 240
(Type species by M D. rubida Pease, 1863 =
Pedicularia pacifica PEASE, 1865)

1925. Pediculariella Tu1ELE, KUcKENTHAL, Handb. Zool., 5:
88(Type species by OD Pedicularia californica NEw-
coms, 1865)

1935. Pediculariona IrEDALE, Aust. Zool., 8: 101 (Type species
by OD Pedicularia stylasteris Heptey, 1903 = P. paci-
fica PEASE, 1865)

spec. juv.

The genus contains only 9-10 species which have a
world-wide distribution. One cannot describe morphologi-
cal or anatomical differences between genera or subgenera
where such do not exist. ScHILDER (1939) in his diagnosis
of genera of Pediculariidae comments: “Unterschiede
unsicher; Anatomie abweichend.”

Pedicularia pacifica PEASE, 1865
(Text figure 4)

1863. ? Dentiora rubida Prase, Proc. Zool. Soc. London for
1862: 240 (spec. juv.?)

1865. Pedicularia pacifica Preasr, Proc. Zool. Soc. London,
p. 516

1868. Pedicularia pacifica PEAsE, Amer. Journ. Conch., 4:
96; plt. 11, figs. 17, 18

1903. Pedicularia stylasteris HEDLEY, Mem. Aust. Mus., 4:
342, figs. 69, 70

1944. Pedicularia pacifica PEASE, ScuitpEr, Ark. Zool. 36A:
29 - 31 (animal, radula, veligers)

1965. Pedicularia pacifica Pease, Kay, Bull. Brit. Mus. (Nat.
Hist.) Zool. Suppl., 1: 84; plt. 14, figs. 13, 14 (lectotype
6x3 mm)

Shell: Small, irregularly ovate or oblong-ovate, reddish-
pink to purple in colour. Dorsum sculptured with numer-
ous concentric ribs and intersecting striae which are
alternately coarse and fine. Aperture wide open, margins
sharply edged, teeth obsolete in adult specimens; spire
visible and clathrate or concealed.

L: 5.5 - 6.2mm.

Figure 4

Pedicularia pacifica PEASE
(after ScHILDER, 1931)

Type Locality: Central Pacific (Apaian Island, PEase,
1868) [=Abaiang Island, Gilbert Islands]

Habitat: Unknown (sessile on coral, fide ScHtLpER, 1931).
Discussion: The species is known from only 2 beach-worn
and partly broken specimens; being unsuitable for photo-
graphy, a drawing of the species is given from SCHILDER,
IQBil,

The species has occasionally been reported from the
Hawaiian Islands, but according to Kay (1965) it does
not live there.

TrIvIDAE TROSCHEL, 1863

1927. ERATOIDAE ScHILpER, Arch. Naturgesch. 91: 1 (as Eratoinae)
1932. EraTomarE SCHILDER, Proc. Malacol. Soc. London, 20: 46

Shells of Triviidae are moderately small or very small,
ovate, round and inflated, sometimes biconical, unicol-
oured, spotted or banded; they are smooth or sculptured
with transverse ribs, granules and a dorsal groove. The
spire is covered or projecting, aperture wide or narrow,
lips denticulate.

The animals of Eratoinae have an open siphon, ie.
grooved, whereas animals of the Triviinae have a closed,
tube-like siphon; in other respects they approximate the
animals of Cypraeidae. The rhachidians of the radula
are either quadrate or trapezoidal, with a larger central
cusp and up to a dozen side denticles; laterals are hook-

Vol. 10; No. 4

THE VELIGER

Page 367

like, simple, and with or without denticles on the cutting
edge; marginals are claw-like, inner marginal occasionally
with a small denticle.

Triviidae have a world wide distribution and range

farther into colder waters than the Ovulidae. Tropical
Triviidae share the same habitat with the Cypraeidae;
they are reef-dwellers and live in crevices or on the under-
side of coral blocks.
Discussion: ScHILDER (1933) pointed out that the anato-
my of Eratoinae hardly differs from that of the Triviinae,
and that the radulae are almost identical. All indications
are that Eratoinae and Triviinae are closely related, the
Eratoinae being the more primitive group of Triviidae
from which Recent Triviinae have diverged.

Triviinae TrRoscHEL, 1863

1932. Eratoinae ScHILpeEr, Proc. Malacol. Soc. London, 20: 46

Species of Triviinae have ovate or round, inflated shells
which are unicoloured or spotted; the transverse dorsal
ribs, which may or may not be interrupted by a dorsal
groove, extend onto the base and into the aperture; the
dorsal ribs are occasionally noduled near the dorsal
groove. The spire is generally covered in adult specimens,
aperture is linear and sometimes wide, and both lips are
denticulate; the columella generally extends deep into the
aperture.

Triviinae have a world wide distribution and live in the
intertidal zone and in deeper water.

Tnvirostra JOUSSEAUME, 1884

Trivirostra JOUSSEAUME, 1884, Bull. Soc. Zool. France, 9: 100. Type
species by SD (RosBErts in Tryon, 1885) Cypraea scabrius-
cula Gray, 1827 = ? Cypraea oryza Lamarck, 1811

Shells small, generally ovate, white in colour; trans-
versely ribbed, dorsal groove moderately short, interstices
of ribs granulose. Sides rounded, aperture narrow, fossula
broad, columella intruding deeply into the aperture.

The radula of Trivirostra differs in some features from
the radula of Trivia Broperip, 1837 [type species T.
monacha (pA Costa, 1778) ]. The rhachidians of Trivia
are quadrate, but are trapezoidal in Trivirostra, and the
basal pegs are more prominent. The laterals of Trivirostra
have small accessory denticles on the cutting edge, similar
to the laterals of Trivia arctica (PULTENEY, 1799), where-
as in Trivia monacha the laterals lack the small denticles;
the marginal teeth of Trivirostra are simple and hook-like,
and the inner marginal may on occasion have a very small
denticle.

Species of the genus inhabit crevices of coral rocks,

and are frequently found on marine benches with a
maximum of algal matting.

Trivirostra (Trivirostra) edgari (SHaw, 1909)
(Plate 51, Figure 17)

1849. Cypraea grando Gasxown, Proc. Zool. Soc. London, 16:
96 (non Potrez & Micuaup, 1838)

1909. Trivia edgari SHaw, Proc. Malacol. Soc. London, 8: 310
(nom. nov. pro Cypraca grando Gasxotn, 1849)

1917. Trivia oryza Opuner, Svens. Akad. Handl. 52: 53; plt.
2, figs. 52, 53; text fig. 12 (shell and radula of Trivirostra
oryzoidea IREDALE, 1935) [non Trivia oryza Lamarck,
1811]

1935. Trivirostra oryzoidea IREDALE, Aust. Zool., 8: 101 (nom.
nov. pro Trivia oryza ODHNER, 1917)

1944. Trivirostra edgari insularum ScuitpErR, Ark. Zool., 36A:
14 (descr. radula)

1944. Trivirostra edgari tomlini ScuitpER, Ark. Zool., 36A: 14

Shell: Small, broadly ovate, extremities produced and
broad ; white in colour throughout. Sculptured with coarse
and elevated transverse ribs and a broad but short dorsal
groove which does not interrupt the 23 - 26 dorsal ribs;
interstices of ribs are minutely crenulate. Aperture al-
most central, labial lip with 20 - 23 denticles, columellar
lip with 16-19 denticles; the interior of the columella
extends in a flatly convex arc, and the fossula is moder-
ately narrow and concave and does not protrude towards
the labial wall as in Trivirostra oryza.
L: 5.9-6.6mm; W: 73-79%; H: 66 - 69%
Type Locality: Manilla (Manila, Philippine Islands).
Habitat: Under coral rocks, in shallow water.
Moderately rare.
Distribution: South Viti Levu, Vanua Levu and Taveuni.
— From East Africa through the tropical Indo-Pacific to
Japan, Hawaii and the Tuamotu Archipelago.
Discussion: Fijian specimens of Trivirostra edgari are
rather similar in dimensions, teeth and dorsal rib count
to the East African race T: edgari tomlini ScHILDER.

Trwirostra (Trivirostra) exigua (Gray, 1831)
(Plate 51, Figure 18)

1831. Cypraea exigua Gray, Zool. Misc., 1: 35

1833. Cypraea tremeza Ductos, Mag. de Zool., plt. 25

1845. Cypraea gemmula Goutp, Proc. Boston Soc. Nat. Hist.,
23 OT)

1868. Trivia corrugata PEASE, Amer. Journ. Conch., 4: 95; plt.
11, figs. 14, 15

1914. Trivia exigua var. alba Sowerby, Proc. Malacol. Soc.
London, 11: 10

1923. Trivia exigua (Gray) VaysstireE, Ann. Mus. Hist. Nat.
Marseilles, 18: 86 - 87; plt. 14, figs. 209 - 212 (anatomy)

1933. Trivirostra exigua hyalina Scuttprr, Zool. Anz., 102: 290

Page 368

THE VELIGER

Vol. 10; No. 4

1964. Cypraea (Trivia) gemmula Goutp, Jounson, U.S.
Nat. Mus. Bull. No. 239: 82; plt. 7, fig. 10 (lectotype)

Shell: Small, roundly pyriform, extremities produced;
pinkish-white in colour, ornamented with 2 - 5 pink spots
on dorsum, dorsal ribs occasionally pink in colour. Sculp-
tured with 16-29 coarse dorsal ribs and an impressed
dorsal groove which does not separate the ribs; the ribs
are sometimes thickened near the margins of the dorsal
groove and interstices of ribs are finely crenulate.
Aperture is slightly offcentral, labial teeth number from
20 to 23, columellar teeth from 18 to 19; fossula is
concave, but extends only slightly towards the labial wall.
L: 4.2-4.5mm; W: 66-71%; H: 60-63%
Type Locality: None (“New South Wales,’ Sowersy,
[1832]).
Habitat: Under coral rocks, in shallow water.
Moderately rare.
Distribution: South Viti Levu. —-
tropical Pacific.

Throughout the

Trivirostra (Trivirostra) hordacea (KiENER, 1843)
(Plate 51, Figure 19)

1827. ? Cypraea scabriuscula var. minor Gray, Zool. Journ.,
3: 364

1843. Cypraea hordacea KiENER, Spéc. gén. icon. coq. viv., p.
149; pl. 54, figs. 5, 5a

1845. Cypraea insecta MicHELs, Proc. Boston Soc. Nat. Hist.,
2: 24

1870. ? Cypraea sandwichensis Sowrrsy, Thes. Conch., 4: 47,
expl. to plt. 35

1870. ? Cypraea sandvichensis SowERBy, Thes. Conch., 4: 57

1912. Trivia desirabilis TREDALE, Proc. Malacol. Soc. London,
10: 226; plt. 9, figs. 8, 9 (Kermadec Island)

1932. Trivirostra hordacea (KIENER), SCHILDER, Zool. Anz.,
100: 226; fig. 2 (radula)

Shell: Small, subcylindrical and elongate, extremities
hardly produced, dorsum depressed; white in colour
throughout. Sculptured with fine and numerous trans-
verse ribs which number from 26 to 32; the dorsal groove
is long and interrupts the dorsal ribs; interstices of ribs
are finely crenulate. Aperture is offcentral, outer lip
moderately narrow and with 22-23 labial teeth; the
columella is rather broad and sculptured with 19 - 21
teeth. Fossula broad and concave, only slightly extending
towards the labial wall, posterior of columellar lip some-
what truncated.

L: 3.8-4.1mm; W: 64-66%; H: 52-58%

Type Locality: Mers de l’Inde, les cétes de l’ile Bourbon

[= Réunion Island].

Habitat: Under coral rocks, in shallow water. — Rare.
Distribution: North and South Viti Levu. — From East

Africa through the tropical Indo-Pacific to Hawaii and
the Tuamotu Archipelago.

Tnivirostra (Trivirostra) oryza (LAMarRcK, 1811)
(Plate 51, Figure 15)

1811. Cypraea oryza Lamarck, Ann. Mus. Hist. Nat., 16: 104

1817. Cypraea nivea Dittwyn, Desc. cat. Rec. shells, 1: 466

1817. ? Cypraea sulcata var. 8 Dittwyn, Desc. cat.. Rec.
shells, 1: 466

1827. ? Cypraea scabriuscula Gray, Zool. Journ., 3: 364

1843. Cypraea intermedia KiENER, Spéc. gén. icon. coq. viv., p.
143; plt. 52, figs. 2, 2a (mon Gray, 1824)

1908. Trivia oryza (LaMARCK), BercH, Semp. Reise Arch.
Philipp., 9: 144; plt. 11, figs. 27 - 37 (anatomy)

1909. Trivia oryza forma minor SCHEPMAN, Siboga Exped.,
49b: 137 (non Gray, 1827)

1932. Trivirostra oryza triticum ScutLpErR, Foss. Cat., 1/55:
101

1932. Trivirostra oryza turneri SCHILDER, Foss. Cat., 1/55: 101

Shell: Small and ovate, extremities generally blunt but
sometimes moderately produced; white in colour through-
out. Dorsum is ornamented with close-set fine transverse
ribs, numbering from 30 to 45; the dorsal groove is
either moderately deep or shallow and, in the majority of
specimens examined, does not separate the dorsal ribs.
Sides are rounded, base convex, aperture narrow, labial
lip fairly straight and sculptured with 19 - 26 denticles.
Interior of columella curved, fossula broad and concave,
and extending towards the labial wall.

Poe

Figure 5
Trivirostra oryza (LAMARCK)
Fiji Islands
Half row of radular teeth

Animal: Sole and dorsum of foot brown, finely veined
with white; mantle light grey, flecked with blackish-
brown, becoming dark brown towards the mantle margin.
Papillae few, short and fringed, yellow in colour. The
siphon is cylindrical, closed and tubular, yellow in colour
and papillate on the sides. Tentacles are short and blunt at -

Vol. 10; No. 4

distal end, brown in colour, flecked with white; eyes

black and ringed with white.

Radula: Radular ribbon white, 3.9mm long and 0.57 mm

wide in an animal with a shell 8.0mm in length; the

ribbon contains 49 rows (plus 4 nascentes) of teeth.

Rhachidians are trapezoidal, 0.13 mm wide and 0.06mm

long, central cusp is long and flanked by 3 or 4 small

denticles; base of plate with a large basal peg on either

side; laterals are large and have ca. 5 denticles on the

cutting edge; marginals simple and curved.

L: 5- 10mm; W: 66-78%; H: 57-62%

Type Locality: Timor.

Habitat: On reefs, under coral rocks, in shallow water.
_ Common.

Distribution: Throughout the Fiji Islands. — From the

Gulf of Oman through the tropical Indo-Pacific to the

Tuamotu Archipelago. ;

Discussion: The species is extremely variable and diagnos-

tic characters show a rather long range of variation.

Well over 100 specimens were measured, and in ca. 90%

of them the labial teeth were 1.2 - 1.3 times as numerous

as the columellar teeth.

Trivirostra (Trivirostra) species

(Plate 51, Figure 16)

Shell: Similar to Trivirostra oryza (LAMARCK), but differs
in being more roundly pyriform with produced extremi-
ties and a shallow dorsal groove which does not interrupt
the dorsal ribs; the aperture is almost central, both colu-
mellar and labial lips are curved, and are sculptured with
28 and 22 denticles respectively. The fossula is extremely
broad, concave and coarsely ribbed, and extends deeply
towards the labial wall.

L: 8.3mm; W: 72%; H: 60%; dorsal ribs: 38
Habitat: Under coral rocks, in shallow water. — Rare.
Distribution: South Viti Levu. - ? Indonesia.
Discussion: The species is known from one live-collected
specimen which seems to be separable from Trivirostra
oryza. It resembles the species T: scabriuscula (Gray),
as of authors, and is depicted under this name in several
monographs on the family.

(Dolichupis) IREDALE, 1930

Dolichupis IrEDALE, 1930, Mem. Qld. Mus., 10: 83. Type species
by OD Cypraea producta Gasxoin, 1836
1931. Trivellona IrEDALE, Rec. Aust. Mus., 18: 221 (Type spe-
cies by M T: excelsa IREDALE, 1931 = Cypraea producta
Gasxoin, 1836)

THE VELIGER

Page 369

Shells are small, ovate, white in colour, extremities pro-
duced, sides broad and somewhat angulate; the dorsal
ribs are coarse and prominent, interstices granulose,
dorsal groove is absent.

Discussion: Dolichupis was established by IREDALE (1930)
for “the forms with produced extremities.” This short
diagnosis is misleading for a generic assignment, in view
of the occurrence of both produced and blunted extremi-
ties in the same species. SCHILDER (1939) treated Dolich-
upis as a subgenus of Pusula JoussEauME, 1884. The
type species of Dolichupis, however, seems morphologi-
cally closer to Trivirostra than to Pusula radians (La-
MARCK, 1811), the type species of Pusula; the latter
species is brownish with lighter coloured dorsal ribs which
are pustulose and widely spaced, and the extremities are
blunter than in Dolichupis. Dolichupis differs from Trivi-
rostra mainly in the more produced extremities, broad
and angulate sides and obsolete dorsal groove; all other
characters are variable between species of the two groups.

Trwirostra (Dolichupis) pellucidula (REEvE, 1846)
(Plate 52, Figures 21, 21a)

1846. Cypraea pellucidula Rerve, Conch. Icon., Cypraea, plt.
26, sp. 153. (March 1846)

1846. Cypraca pellucidula Gasxown, Proc. Zool. Soc. London,
p. 23 (May 1846)

1881. Cypraca pellicula (sic) WretnKaurF, Conch. Cab., ed.
2, p. 161; pl. 43, figs. 1, 4

1932. Trivirostra pellucidula natalensis SCHILDER, Foss. Cat.,
1/55: 102

Shell: Small, roundly pyriform, extremities produced;
white in colour throughout. Sculptured with 26 - 30 fine
dorsal ribs, dorsal groove is absent or very obsolete,
interstices minutely crenulate; sides are rounded, aperture
slightly off-central and curved. Labial teeth number from
26 to 28, columellar teeth from 20 to 22; fossula is very
broad and concave and extends towards the labial wall.
L: 6.0-6.8mm; W: 69-71%; H: 58-60%
Type Locality: South Pacific.
Habitat: In 15 fathoms, on broken coral substratum.
Rare.
Distribution: West off Viti Levu. — From East Africa
through the tropical Indo-Pacific to Japan, Hawaii and
the Tuamotu Archipelago.
Discussion: ScuitpER (1941) placed Trivirostra pelluci-
dula in the subgenus Trivirostra s. str.; if the subgenus
Dolichupis is to contain triviid species with produced
extremities and an obsolete dorsal groove, then the species
T. pellucidula clearly belongs to Dolichupis.

Page 370

GaskoIN is generally credited with the authorship of
Trivirostra pellucidula, but ReEve’s publication of the
species name has 2 months priority over GasKOIN’s.

Trivirostra (Dolichupis) producta (GasxKoin, 1836)
(Plate 52, Figures 20, 20a)

1836. Cypraea producta Gasxoin, Proc. Zool. Soc. London
for 1835: 200

1837. Cypraea producta GasKkoIn, SowERBY, Conch. IIlust.,
fig. 155 (holotype)

1871. Trivia candidula Ancas, Proc. Zool. Soc. London, p. 94
(non GasxoIn, 1836)

1923. Trivia producta (GasKoIn), VayssizrE, Ann. Mus.
Hist. Nat. Marseilles, 18: 88-89; plt. 9, figs. 115-121
(anatomy)

1931. Trivellona excelsa IREDALE, Rec. Aust. Mus., 18: 221;
plt. 14, figs. 13, 14

Shell: Small, ovate and humped, sides broadened and
slightly angulate; white in colour throughout. Sculptured
with 20 - 26 strong, elevated and sharply cut transverse
ribs which cross the centre of the dorsum and descend
towards the margins; a dorsal groove is absent, but oc-
casionally faintly impressed, interstices of ribs broad and
crenulate. Base flattened, aperture almost central and of
medium width, labial lip broad and with 22 - 24 denticles;
columella with 19 - 23 denticles, interstices of ribs with
small axial riblets. Posterior outlet sharply cut, fossula
very broad, concave and extending towards the labial
wall.
Animal: Sole of foot white, dorsum of foot translucent
white, ornamented with iridescent white spots; mantle
white, papillae tufted, cream-coloured, and extending
half-way up the mantle; the mantle margins are speckled
with grey. Tentacles creamy-white; siphon greyish-white,
flecked with darker grey and almost as long as the shell
itself.
L: 10- 15mm; W: 70-73%; H: 54-56%
Type Locality: None (“Unsang, East coast of Borneo,”
fide ADAMS & REEVE, 1848).
Habitat: In 15 - 16 fathoms, on coral rubble substratum.
Moderately rare.
Distribution: West off Viti Levu. — From Indonesia to
the Tuamotu Archipelago.
Discussion: The holotype of Trivellona excelsa IREDALE,
1931, preserved in the Australian Museum, Sydney, is a
large (18.0mm), dead-collected, anteriorly somewhat
worn specimen of Trivirostra producta (Gasxkotn).
Sowersy (1870) reports the species from Agulhas
Bank; if this record is correct, then the species’ range
extends as far westward as South Africa.

THE VELIGER

Vol. 10; No. 4

Eratoinae ScuiLpEr, 1927

Species of this subfamily differ from Triviinae in having
biconical, marginellid shells with a projecting and broad
spire and an angulate aperture. Shells are unicoloured or
banded, dorsally grooved, smooth, granulose or ribbed;
shell outlets are wide and shallow, aperture narrow, and
both lips are denticulate.

Recent species of Eratoinae have been recorded from
the Indo-Pacific, Europe, West Indies and the west coast
of America. Tropical Eratoinae are reef-dwellers, and
inhabit crevices of the underside of coral blocks.

Lachryma Sowersy [1832]

Lachryma Sowersy, [1832], Conch. Illust., Cat. Cyp., p. 15. Type
species by T Erato lachryma Sowersy, 1832
[1832]. [Erato] lachryma Gray, SoweErRBy. Lachryma trifasci-
ata Humpurey, Mss. Conch. Illust., fig. 48
1927. Proterato ScuitpER, Arch. Naturgesch., 91A: 57 (Type
species by OD Erato neozelanica Suter, 1917)
1931. Lachryma IrEpALE, Rec. Aust. Mus., 18: 223 (in com-
bination Lachryma bisinventa IREDALE, 1931)
1935. Lachryma SoweErsy, IREDALE, Aust. Zool., 8: 97
1935. Eratoena IrREDALE, Aust. Zool., 8: 97 (Type species by
OD Ovulum corrugatum Hinps, 1845 = Erato sulci-
fera SowErRBy, 1832)
1958. Lachryma SowerBy, Cotton, Journ. Malacol. Soc.
Aust., No. 2: 11

Shells are very small, biconical, smooth or granulose, spire
broad and often projecting, dorsal groove prominent or
obsolete, aperture narrow, lips denticulate, posterior colu-
mellar denticles occasionally obsolete.

Discussion: Lachryma SowerBy [1832] will have to re-
place Proterato ScuHiLvER, 1927. Lachryma was published
by Sowerby [1832] in the synonymy of Erato lachryma
Sowersy, 1832, ex HumMpurey MS. Lachryma has been
treated as an available name with its original date and
authorship prior to 1961 (art. 11d, Code of ICZN, 1964)
and has been adopted as the name of a taxon by IREDALE
(1935) and Cotrron (1958).

The date of publication of the “Catalogue of Cyprae-
adae” by SowerBy appcars to be in dispute. SCHILDER
(1933) quotes 1837 as the publication date, whereas
IREDALE cites the 16" November 1832. SHaw (1909 a)
seems to favour the latter date, as a part of the “Cata-
logue of Cypraeadae” (? pp. 1-8) has definitely been
issued with part 7 of the Conchological Illustrations (9
November 1832). The remaining pages 9 - 18 probably
appeared with part 8 (prior to 30 November 1832).
Malacologists not accepting 1832 as the publication date
would have to date Lachryma from 1837, the date of

Vol. 10; No. 4

THE VELIGER

Page 371

appearance of the “Index to the Cypraeadae” with part
131 (15 November 1837).

TREDALE (1935) established Eratoena for Recent trop-
ical species of Eratoinae with sculptured shells; he
pointed out that although the sculpture can become “sub-
obsolete,” the species can be distinguished by apertural
characters, which he did not specify. When IREDALE
mentioned apertural differences between Lachryma and
Eratoena, he probably meant the difference in denticula-
tionof the columellar lip. In Lachryma lachryma (SowErR-
By) the posterior columellar denticles are obsolete, but
they can be prominent along the whole length of the
columella in Lachryma sulcifera (SowERBY) or can be
obsolete posteriorly. This feature is rather variable, and
ScHILDER (1933) states that specimens of L. callosa
(ApaMs & REEveE, 1848) with obsolete or prominent pos-
terior columellar teeth are represented in about equal
numbers. Since Eratoena has been based on variable
specific characters, it is relegated to the synonymy of
Lachryma Sowersy, following ScHILDER (1939, but not
1941).

Proterato SCHILDER could questionably be retained as
a subgenus of Lachryma for the New Zealand Miocene
fossil Erato neozelanica SuTER, 1917, which is rather
globular, smooth, with a conical spire, obsolete posterior
columellar teeth, and has a somewhat wider aperture.

Lachryma sulcifera (SowERBy, 1832)
(Plate 52, Figures 22, 22a)

1832. Erato sulcifera SowERsBy, Conch. Illust., Cat. Cyp., p.
15; fig. 46

1845. Ovulum corrugatum Hinps, Zool. Voy. Sulphur, p. 47;
pit. 16, figs. 5, 6

1859. ? Erato nana Sowersy, Thes. Conch., 3: 82; plt. 219,
figs. 12, 18

1867. Erato schmeltziana Crossr, Journ. Conch., 15: 301;
pit. 11, fig. 5 (Fiji Islands)

1933. Proterato (Proterato) sulcifera capensis SCHILDER, Proc.
Malacol. Soc. London, 20: 248; fig. 22

1933. Proterato (Proterato) sulcifera smitht ScutLveEr, Proc.
Malacol. Soc. London, 20: 248; fig. 24

1933. Erato (Cypraeerato) schneideri ScuitpER, Zool. Anz.,
102: 294, 296

1933. Erato schmeltiana (sic) Crosse, DAUTZENBERG &
Bouce, Journ. de Conchyl., 77: 294

Shell: Marginellid in shape, spire elevated, apex blunt,
labial lip moderately broad, roundly angulate near shoul-
der; greenish or light greenish-fawn in colour, rarely
greenish-white, occasionally banded with 1-3 bright
green or greenish-brown transverse bands which may be
broken up into blotches. Anterior extremity pinkish-brown
or orange-brown. Sculptured with a decp or shallow

dorsal groove, numerous small granules which may co-
alesce into short or long axial riblets; some specimens are
smooth and only granulose at the shoulder and spire.
Labial lip with 15 - 21 denticles, which are either short
or produced to the margin; columellar terminal ridge is
transversely striate, columellar teeth are small denticles
which may become obsolete posteriorly, and number from
8 to 18. Aperture is narrow, fossula either narrow or
broad.
Animal: Foot long and slender, pointed posteriorly,
cream in colour, finely spotted with brown at the margins.
Mantle creamy-yellow, spotted with small brown spots
and a few irregular and large brown zones. Siphon open,
creamy-yellow, minutely spotted with brown; tentacles
moderately long, slender, translucent creamy-white, finely
spotted with brown; eyes black.
L: 2.8-4.3mm; W: 61 - 68%
Type Locality: Cape of Good Hope.
Habitat: Under coral rocks, in shallow water.
Uncommon.
Distribution: Throughout the Fiji Islands. -— From
the Red Sea through the tropical Indo-Pacific to the
Tuamotu Archipelago.
Discussion: Tropical species of Eratoinae are highly
variable in colour and sculpture; what may be considered
a specific diagnostic character may well be only an eco-
phenotypic variation. BARNARD (1963) observed that
specimens of Lachryma sulcifera from South Africa are
either smooth, granulose, axially ribbed or granulose only
posteriorly, that the dorsal sulcus may be short or even
absent and that the colour is variable. A similar variation
has been observed in Fiji specimens. SCHILDER (1933)
considers L. corrugata (Htnps) as a race of L. sulcifera,
but admits that intermediate shells have been recorded
and that in some localities both races seem to live together.
This certainly holds true for Fiji, where the corrugata
form lives side by side with typical L. sulcifera.

The difficulty of interpretation of diagnostic characters
of Eratoinae is best documented by the taxon Lachryma
sulcifera schneideri (SCHILDER) which was originally
described as a full species in the genus Erato Risso, 1826,
subgenus Cypracerato ScHILpER, 1932. It was reduced to
the rank of a subspecies the same year and transferred to
Proterato s. str. In 1941 Lachryma schneidert was once
again considered to be a good spccies and was assigned to
the subgenus Sulcerato Fintay, 1930, while Lachryma
sulcifera was transferred to the subgenus Eratoena IrRE-
DALE, 1935. The majority of geographical races of La-
chryma sulcifera have been based on diagnostic characters
(c.g. absent dorsal groove, granules becoming riblets,
granules confined to spire or base), which can be observed

Page 372

in populations of L. sulcifera from a confined geograph-
ical region.

UNCONFIRMED REPORTS

Primovula (Diminovula) margarita (SowERBy, 1828)
(Plate 52, Figures 23, 24, 25)

1828. Ovulum margarita SowErsy, Zool. Journ., 4: 150

1828. Lictum margarita SowERByY (ex HumpHREY MS), Zool.
Journ, 4: 150, in synonymy of Ovulum margarita
Sowerby, 1828)

1830. Ovulum margarita SowERBy, Spec. Conch., Ovulum,
1: 4; figs. 19, 20

1849. Ovulum umbilicatum SoweErBy, Proc. Zool. Soc. Lon-
don, 16: 135

One very worn specimen collected in the Fiji Islands
could possibly be this species, but since it is partially
broken, the identification is not at all certain.

Type Locality: In Insulis quas “Friendly” vocamus, in
Mari Pacifico [= Tonga Islands].

Dimensions of type series:

L: 8.2-13.8mm; W: 65-70%; LT: 22-29; crenula-
tions on first funiculum: 3 - 6.

Discussion: SowerBy (1828) described the species from
several specimens in the Humphrey collection and re-
marked that all specimens bar one were pierced and
strung by natives. The type series of Ovulum margarita
preserved in the British Museum (Natural History) con-
sists of 5 specimens; 4 of these have been holed above
the labial margin, and the smallest specimen is complete
(N. Tebble, in litt.). Sowersy’s dimensions of the holo-
type given in the original publication were length 11/20
poll. (= 13.97 mm) and width 4/10 poll.(—= 10.16mm) ;
these dimensions closely correspond with the holotype,
which is the largest specimen in the series. The dimensions
of the type series of O. margarita are as follows:
Holotype: B. M. N. H. Reg. No. 1967613/1. L: 13.8mm;
W: 9.5mm; LT: ca. 29; first funiculum with ca. 3
crenulations.

Paratype: B. M. N. H. Reg. No. 1967613/2. L: 8.2mm;
W: 5.5mm; LT: ca. 28; (the only complete specimen,
slightly immature).

Paratype: B. M. N. H. Reg. No. 1967613/3. L: 12.1mm;
W: 8.5mm; LT: ca. 23; first funiculum with ca. 6 cren-
ulations.

THE VELIGER

Vol. 10; No. 4

Paratype: B. M. N. H. Reg. No. 1967613/4. L: 8.8mm;
W: 6.2mm; LT: ca. 22; first funiculum with ca. 4 crenu-
lations.
Paratype: B. M. N. H. Reg. No. 1967613/5. L: 11.5mm;
W: 7.5mm; LT: ca. 24; first funiculum with ca. 3 crenu-
lations.

The species is pyriform, inflated and generally white in
colour; the dorsum is finely transversely striate, and a
dorsal carina is either absent or feebly pronounced. Labial
teeth are strong but irregular, the first funiculum is prom-
inent and crenulate, the posterior outlet is distinct, poste-
rior extremity calloused and recurved and the fossula is
concave.

WeinKaurF (1881) also reported this species from
the Tonga Islands, and ScuiLpeR (1941) lists the distri-
bution for this species as ranging from Indonesia to the
Philippines, Bismarck Archipelago and the Tonga Islands.

FOSSIL RECORDS

Trivirostra (Trvirostra) koroensis (Lapp, 1934)

1934. Trivia (Trivia) koroensis Lapp, Bernice P. Bishop Mus.
Bull., 119: 220; plt. 39, figs. 9, 10

The species was described from late Tertiary (Pliocene)
deposits at station 160 (Walu Bay, Suva, few feet above
sea-level). The length was given as 3.9-4.7mm, the
width (calculated) as 68 - 69% of length, and the height
as 62% of length. The holotype of Trivirostra koroensis
has 27 labial and 23 columellar teeth and ca. 32 - 34
dorsal ribs (counted from type figure).

The species resembles Trivirostra hordacea (KiENER,
1843), but has somewhat more numerous teeth on both
lips.

SUMMARY

Further records of species of the 3 families treated in this
monograph are only to be expected. In the case of rare
species only diligent collecting over a great number of
years can present a true record of the actual existence of
species of any molluscan family in a certain geographical
region.

The total number of species recorded in the various
genera of the 3 families is as follows:

Vol: 10; No. 4

THE VELIGER

Page 373

OvuLDAE

Ovula-
Calpurnus
Prosimnia
Primovula
Prionovolva
Volva
Phenacovolva
Pseudocypraea

me OF Re e& ND ND DO

Total

—
iss)

PEDICULARIDAE

Pedicularia 1

TRIVUDAE

Trivirostra 7
Lachryma

Total 8

LITERATURE CITED

ApaMs, ARTHUR &« Lovett, Aucustus REEVE
1848-1850. The zoology of the voyage of H.M.S.Samarang,
under the command of Captain Sir Edward Belcher. Mollusca.
London, prts. 1-3: 1-87; 24 plts.
Barnarp, KeppeL Harcourt

1963. Contributions to the knowledge of South African marine
Mollusca. Part III. Gastropoda: Prosobranchiata: Taenioglossa.
Ann. South Afr. Mus. 47 (1): 1- 199; 37 text figs.

CERNOHORSKY, WALTER OLIVER
1964. The Cypraeidae of Fiji (Mollusca: Gastropoda).
The Veliger 6 (4): 177-201; plts. 21-26; 1 Text fig.; 1 map
(1 April 1964)
Corton, BERNARD CHARLES
1958. Western Australian cowries.
Austral. 2: 8-15; plts. 2-5

GarreETT, ANDREW

Journ. Malac. Soc.
(November 1958)

1879. Annotated catalogue of the species of Cypraeidae col-
lected in the S. Sea Islands. Journ. Conch. 2: 105 - 128
(April 1879)
Hase, TADASHIGE
1961. Coloured illustrations of the shells of Japan.
2: i- xii; 1- 183; App. pp. 1 - 42; plts. 1 - 66; text figs.
(20 April 1961)

Osaka,

Hipatco, Joaquin GonzaLEz
1906 - 1907. Monografia de las especies vivientes del género
Cypraea. Mem. Rea. Acad. Cienc. 25: 1-240 (August
1906) ; 25: 241 - 588 (August 1907)
TREDALE, Tom

1930. Queensland molluscan notes, No. 2. Mem. Qld.
Mus., 10 (1): 73 - 88; plt. 9 (28 August 1930)

IREDALE, Tom

1931. Australian molluscan notes. no. 1. Rec. Austral. Mus.
18 (4): 201 - 235; plts. 22 - 25 (29 June 1931)
1935. Australian cowries, Part I. Austral. Zoologist

8 (2): 96-135; plts. 8, 9
IREDALE, Tom & Donatp F McMicHaev
1962. _A reference list of the marine Mollusca of New South
Wales. Austral. Mus. (Sydney), Mem. 11: 1 - 109
(30 May 1962)
Kay, E. ALISON
1965. | Marine molluscs in the Cuming collection, British Mu-
seum (Natural History) described by William Harper Pease.
Bull. Brit. Mus. (Nat. Hist), Zool. Suppl. 1: 1 - 96; plts. 1 - 14
(December 1965)
Kiener, Lours Cuar.es
1833-1844. Spécies général et iconographie des coquilles vivantes.
Genre Ovula. Paris, 1: 1 - 26; plts. 1-6 (for collation see
SHERBORN & Woopwarp, 1901)
Lamarck, JEAN-BapTiste PreRRE ANTOINE DE MONET DE
1798. Tableau encyclopédique et méthodique des trois régnes
de la nature. Paris, plts. 287-390 [22 September 1798]

1801. Systéme des animaux sans vertébres, ou tableau général
des classes, des ordres et des genres de ces animaux;
vili+432 pp. Paris (chez l’auteur & Deterville)
Martyn, THOMAS

1784-1786. The universal conchologist.
plts.

London, 4 vols., 160

Montrort, Pierre DENYS DE
1810. | Conchyliologie systématique et classification métho-
dique des coquilles. Paris, 2 vols.
1: i- Ixxxvii + 1-409; 2: 1-676 + 1-16
Pease, WILLIAM HarPER
1868. Descriptions of marine gasteropodae, inhabiting Poly-
nesia. Amer. Journ. Conch. 4: 71-80; 91-102; 4 plts.
Peize, A. J.
1925. The differentiation as species of the two forms of
British Trivia. Proc. Malacol. Soc. London 16 (4): 195
to 197; 2 text figs. (9 April 1925)
Ray, HartsH CHANDRA
1949. Revision of Cypraeacea in the collection of the Zoolo-
gical Survey of India. Part I. The families Triviidae, Eratoidae
and Pediculariidae. Rec. Ind. Mus. 46 (1-4): 183 - 213
(April 1949)
REEvE, LovELL Aucustus
1845. | Conchologia Iconica vol. 3: Monograph of the genus
Cy praca. London, plts. 1-27 (Nov. 1845-March 1846)
Ropinc, PETER FRIEDRICH
1798. Museum Boltenianum sive catalogus cimeliorum; pars
secunda Conchylia. Hamburg, i - viii; 1 - 199
(10 September 1798)
ScHEPMAN, MatrHeus Marinus
1909. The Prosobranchia of the Siboga Expedition. Part II.
Taenioglossa and Ptenoglossa. Siboga Exped. 49b: 110 - 231;
plts. 10 - 16

Page 374

ScuiLper, Franz ALFRED
1922. Contributions to the knowledge of the genera Cypraea

and Trivia. Proc. Malac. Soc. London 15 (2-3): 98 - 122
(December. 1922)
1927. Revision der Cypraeacea (Moll. Gastr.) Arch.
Naturgesch. 91A: 1 - 171
1931. Revision of the subfamily Pediculariinae. Journ.
Conch. 19 (6): 165 - 169; plt. 6 (December 1931)
1932. The living species of the Amphiperatinae. Proc.
Malacol. Soc. London 20 (1): 46 - 64; plts. 3-5 (March ’32)
1933. | Monograph of the subfamily Eratoinae. Proc.

Malacol. Soc. London 20 (5): 244 - 283; 85 text figs.
(July 1933)

1939. Die Genera der Cypraeacea. Arch. Molluskenk.
71 (5/6): 165-201; plt. 8 (1 November 1939)

1941. | Verwandtschaft und Verbreitung der Cypraeacea.
Arch. Molluskenk. 73 (2-3): 57-120; 2 plts.
(15 May 1941)

1964. Primovula striatula tinctura (GARRARD). Hawaiian
Shell News 12 (11): 4; 2 figs.; chart (September 1964)

1966a. The higher taxa of cowries and their allies. The
Veliger 9 (1): 31-35 (1 July 1966)

1966b. LinNaEus’ type specimens of cowries. The Veliger

9 (2): 91-100 (1 October 1966)
ScHILDER, Franz ALFRED & Maria SCHILDER
1944. | Westpazifische Cypraeacea von den Forschungsreisen des

Prof. Dr. Sixten Bock.
11-32 authored by F A. ScHILpEr only)

SHaw, H.O.N.
1909. Notes on the genera Cypraea and Trivia, Proc.

Malacol. Soc. London 8 (5): 288 - 313; plts. 12-13; text figs.
(10 August 1909)

1909a. On the dates of issue of Sowerby’s “Conchological
Illustrations,” from the copy preserved in the Radcliffe Library,

Oxford. Proc. Malacol. Soc. London 8 (6): 333 - 340
(5 October 1909)

Swerporn, Davies Cuar.es & BERNARD BARHAM WoopWARD
1901. Notes on the dates of publication of the parts of
Kiener’s “Species général et iconographie des coquilles vi-
vantes,” etc. (1834-80). Proc. Malacol. Soc. London 4 (5) :
216 - 219 (25 July 1901)

Arkiv Zool. 36A (2): 1-32 (pp.
(11 Sept. 1944)

THE VELIGER

Vol. 10; No. 4

Smit, Epcar ALBERT
1910. Notes on the genus Erato, with a list of the known
Recent species. Proc. Malacol. Soc. London 9 (1): 13 - 22
(31 March 1910)

SoLem, ALAN

The Veli-
(1 January 1964)

1964. Amimopina, an Australian enid land snail.
ger 6 (3): 115 - 120; 4 text figs.
Sowersy, GrorcE BRETTINGHAM

1828. On the Recent species of the genus Ovulum. Zool.
Journ. 4: 145 - 162 (October 1828)

1830. Monograph of the genus Ovulum. Species Con-
chyliorum, London 1 (1): 1-10; 2 plts.; figs. 1-58
(November 1830)

Sowerby, GrorcE BRETTINGHAM, Jr.

1832-1837. Conchological illustrations. London, prts. 1-8
(28 September, 1832 to November 1832) ; prts. 101 - 131 (30
July 1836 to 15 November 1837)

1870. Thesaurus conchyliorum. Monograph of the genus
Cypraea. London, 4: 1 - 58; plts. 1 - 37

Stott, NorMANn Rupo.tpu, et al.

1964. International Code of Zoological Nomenclature adopted
by the XV International Congress of Zoology, ed. 2. —
London (Internat. Trust f. Zool. Nomencl.) pp. i- xvii+
1-176; 5 appendices & Glossary

Swainson, WILLIAM
1840. _A treatise on malacology; or the natural classification of
shells and shell-fish. London, i- viiit 1 - 419; 130 text figs.
(May 1840)

THIELE, JOHANNES
1929. Handbuch der systematischen Weichtierkunde.
Jena, Gustav Fischer, 1929-1935; 1- 1154; 893 text figs.
(Cypraeidae: pp. 267 - 277)

WeInKaurF, Heinrich ConraD
1881. Die Gattungen Cypraea und Ovula. Conchylien-Ca-
binet (2nd. ed.), Martini & CHEMNITZ. Niirnberg, pp. 1 to
230; pits. 1-53A

ZitcuH, ADOLF

1960. | Gastropoda Euthyneura. Handb. Paldozool., Band
6, Teil 2, Lief. 4: 601-834; figs. 2112-2515.

Vol. 10; No. 4

THE VELIGER

Page 375

Aplysia vaccaria, a New Host for the Pinnotherid Crab

Opisthopus transversus

ANTHON CRAIG BEONDE

Department of Biology

California State College at Long Beach, Long Beach, California 90804

(2 Text figures)

On Aprit 2, 1967, Two sPEcIMENS of Aplysia vaccaria
WINKLER, 1955, were collected at Corona del Mar State
Beach, California, for anatomical studies. From each A.
vaccaria a specimen of the pinnotherid crab Opisthopus
transversus RATHBUN, 1893, was obtained. Both sexes
were obtained. The carapace widths were 10.4mm and
10.5mm for the male and female respectively. Further
collecting at Corona del Mar State Beach, and examina-
tion of 15 A.vaccaria and 29 A. californica Cooper, 1863,
did not yield any additional specimens.

The two specimens of Opisthopus transversus were
originally observed leaving the pallial cavities of Aplysia
vaccaria while the mollusks were being prepared for dis-
section. Their significance was not realized at the time
and no observations were made on any possible damage
they may have done to the pallial cavity or ctenidium.

Pearce (1966) observed extensive ctenidial erosion
in Mytilus edulis LinNAEuS, 1758, associated with the
pinnotherid crab Fabia subquadrata Dana, 1851. Mac-
Ginitie & MacGrnitie (1949) and Pearce (1966) both
observed that Fabia fed upon the food string produced
by the ctenidia in Mytilus and that while feeding upon
this string would occasionally eat sections of the ctenid-
ium. McDermott (1962) had made similar observations
with Pinnotheres ostreum Say, 1817, where he found that
the ctenidial edge in Anomia showed swelling and per-
foration accompanied in some cases with polyp erosion.

Numerous hosts for Opisthopus transversus have been
recorded, although it is seldom mentioned in the literature
and rarely encountered. Table 1 gives the recorded hosts
along with the respective references.

What may prove to be of more interest is the great
number of hosts with which Opisthopus transversus has
been associated, but that it also chooses to live with
Aplysia, an animal that to date has no recorded commen-
sal associations, or predators, except possibly Anthopleura

(WINKLER & Titton, 1962). It was not until 1962 that
VicENTE described the first known parasite from Aplysia:
a trematode metacercaria parasitic upon the nerve ganglia.

In its pelecypod hosts, Opisthopus transversus remains

Figure 1

Dorsal View of Aplysia vaccaria
C —- Ctenidium M - Mantle P — Parapodium
RPC —- Right Pallial Cavity

Page 376

THE VELIGER

Vol. 10; No. 4

Table 1

Recorded Hosts for the Pinnotherid Crab Opisthopus transversus and the Respective Literature References.

Mollusca
AMPHINEURA
Amicula stelleri (MmprEnporFF, 1847)
(= Cryptochiton stelleri)
GASTROPODA
Megathura crenulata (SowERBy, 1835)

Ricketts & Carvin, 1939; MacGinitie & MacGiniTie, 1949

Ratusun, 1893; 1904; 1918; WeymMoutH, 1910; Scumirtt, 1921;

Ricketts & Carvin, 1939; MacGinitie « MacGiniTiz, 1949

Astraea undosa (Woon, 1828)

Polinices lewisii (Goutp, 1847)

Navanax inermis (Cooprr, 1862)

Bulla gouldiana Pitssry, 1895

Aplysia vaccaria WINKLER, 1955
PELECYPODA

Mytilus edulis Linnaeus, 1758

Pholas sp.

Sanguinolaria nuttallii Conrav, 1837

Schizothaerus nuttallii (Conrap, 1837)

(= Tresus nuttallii Conran, 1837)

Zirfaea sp. :

Platyodon sp.

Modiolus sp.

Megapitaria squalida (SowERBY, 1835)

Echinodermata

HOLOTHUROIDEA

Stichopus californicus (Stimpson, 1857)

ScumitT, 1921"

MacGinitigE & MacGinitie, 1949

MacGinitig & MacGrnitie, 1949

RickeTTs & Carvin, 1939; MacGinitre & MacGrnirig, 1949

Ratusun, 1904; 1918; Scumirt, 1921

Ratueun, 1904; 1918; Ricketts & Catvin, 1939
MacGinitTrE, 1935; MacGinitiz & MacGinitig, 1949
ScumitT, 1921'; Ricketts & Cavin, 1939; MacGIniTIE &

MacGinitie, 1949

MacGinitiz & MacGinitig, 1949
MacGinitizE & MacGinitie, 1949
MacGinitiz & MacGinitiz, 1949
GarTH, 1967?

WeymoutTH, 1910; Ratusun, 1918; Scumirt, 1921, RickETTS &

Carvin, 1939; MacGinitie & MacGinirtie, 1949

* Scumitt (1921) notes that Mr. E. P. Chace of Los Angeles collected
specimens with Astraea undosa and Schizothaerus nuttallu.

2

Dr. Garth in a personal communication (1967) mentions that only

two specimens are in the Allan Hancock collection. One found with
Megapitaria squalida, was collected at Laguna San Ignacio, Baja
California, by Mr. R. L. Eberhart of the California Department of
Fish and Game. The other was collected with Megathura crenulata

at Santa Monica, California.

safe within the confines of , and relies upon the host to col-
lect its food. However, with the gastropod, amphineuran,
and holothuroidean hosts it becomes evident that O.
transversus is forced occasionally to leave the protected
confines and forage for food. Aplysia does not offer the
advantage of being a ctenidial feeder: the ctenidium
serves purely a respiratory function.

Figure 1 represents a dorsal view of the pallial cavity
of Aplysia vaccaria and depicts the large area between
the mantle shelf on the left and the overhanging para-
podial flap on the right, an area in which the crabs could

freely move. Aplysia vaccaria, unlike Aplysia californica,
usually keeps the parapodial flaps closed over the pallial
cavity. The parapodia in A. vaccaria are thick and mus-
cular while those in A. californica are thin and fleshy,
which, when the animal is active, hang loosely out into
the water, thereby exposing the pallial cavity. Figure 2
depicts what remains of the “left” pallial cavity that
would have ancestrally contained the left ctenidium. This
is a small cavity found beneath the overhanging mantle

with its enclosed shell. There is a small aperture to the

pallial cavity at the base of the ctenidium. In a specimen

—

Vol. 10; No. 4

THE VELIGER

Page 377

of A. vaccaria, 10 inches long, this cavity is large enough
to allow entrance of the index finger with ample room
to spare. Since this area is well shut off from the out-
side and protected by the shell overhead it would be
the most likely place to find Opisthopus transversus.
Generally pinnotherid crabs display host specificity, but
there are recorded instances where more than one host
may be utilized, or where more than one host may be
involved in the life cycle of a single species. McDERMOTT
(1962, in reviewing CHRISTENSEN, 1958: On the life
history and biology of Pinnotheres pisum) has reported
that Pinnotheres pisum, a European species, may utilize
two mollusks in completing its life history. He also referred
to P ostreum, a species commonly found in the pelecypod
Anomua. It is evident that P ostreum first invades Mytilus
in the fall, where it seeks shelter, and matures to the hard
stage in winter at which time it is no longer found. Both
sexes possibly leave the hosts and seek other mollusks
(i. e. Anomia) in which the female may grow to maturity.
This may partially explain the recorded hosts for O pistho-
pus transversus or it may be simply that O. transversus
has only recently evolved a commensal relationship and

Figure 2

Cross-Sectional View of Aplysia vaccaria
C — Ctenidium F — Foot GC — Gut Cavity
LPC — “Left” Pallial Cavity ©M- Mantle P — Parapodium
RPC — Right Pallial Cavity S — Shell

is presently experimenting with many possible hosts. The
choice of a host may therefore be dependent upon three
simple factors: 1. the size of the crab itself, 2. the
host’s ability to supply or bring it within close proximity

of a food supply, and 3. the protection afforded by the
host animal.

Opisthopus transversus is known to range as far north
as Monterey, California (RaATHBUN, 1904) and as far
south as San Felipe, Baja California (GLASSELL, 1935).
NININGER (1918) reported having dredged specimens
from a depth as great as 40 metres off Laguna Beach,
California. Aplysia vaccaria ranges from Morro Bay, Cal-
ifornia, to Bahia de Los Angeles, Baja California (LANCE,
1967), totally within the known range of O. transversus.

ACKNOWLEDGMENT

The writer is indebted to Mr. Roby Ward Renshaw,
California State College at Long Beach, for his invaluable
assistance in preparing and proof reading this paper.

Addendum at time of proof-reading:

The following additions to Table 1, due to the original oversight
of
Hopkins, Tuomas S. « Tuomas B. ScANLAND

1964. Host relations of a pinnotherid crab Opisthopus trans-

versus RATHBUN. Bull. So. Calif Acad. Sci. 63: 175 - 180
should be made:

Opisthopus transversus was found commensally with 4 known
hosts: Megathura crenulata, Astraca undosa, Bulla gouldiana, and
Schizothaerus nuttallii [=Tresus nuttallii] - and 6 new hosts: the
polychaete Chaetopterus variopedatus (RentER, 1804), the mol-
lusks Zirfaea pilsbryi Lowe, 1931, Hinnites multirugosus (GALE,
1928), Trachycardium robustum and the holothurians Parasticho-
pus parvimensis (CLarK, 1913), and Molpadia arenicola (Stimp-
son, 1857).

LITERATURE CITED

GLASSELL, STEVE
1935. | New or little known crabs from the Pacific coasts of
northern Mexico. San Diego Soc. Nat. Hist. Trans. 8 (14) :
91 - 106
LancE, JAMES ROBERT
1967. Northern and southern range extensions of Aplysia vac-
caria_ (Gastropoda : Opisthobranchia). The Veliger 9 (4) :
412 (1 April 1967)
MacGinitie, Georce EBER
1935. Ecological aspects of a California marine estuary. Amer.
Midl. Natur., 16 (5) : 629 - 765
MacGinitige, GeorcE Eser & Nettie MacGinitie
1949. Natural history of marine animals.
McGraw-Hill, New York.

McDermott, JoHN JosEPH

473 pp.; illus.

1962. Incidence and host parasite relations of pinnotherid
crabs. Proc. first Natl. Coastal Shall. Water Res. Conf.
pp. 162 - 164

Page 378 THE VELIGER

NININGER, Harvey HarLtow

1918. | Crabs taken at Laguna Beach in the summer of 1916.
Journ. Entomol. Zool., Pomona College 10: 36

PEARCE, JACK
1966. The biology of the mussel crab, Fabia subquadrata,
from the waters of the San Juan Archipelago, Washington.
Pacific Sci. 20 (1): 3-35
RatHusun, Mary
1893. Descriptions of new genera and species of crabs from
the west coast of North America and the Sandwich Islands.
Proc. U.S. Nat. Mus. 16: 252

1904. | Decapod Crustacea of the northwest coast of North
America. Harriman Alaskan Exped. 10: 188

1918. | Grapsoid crabs of America. U.S. Nat. Mus. Bull.
97: 461

Ricketts, Epwarp FE « Jack CaLvIN
1939. Between Pacific tides. 3rd ed.; Stanford Univ. Press,
516 pp.

Scumitt, WaLpo LaSALLe

1921. |The marine decapod Crustacea of California, with spe-
cial reference to the decapod Crustacea collected by the United
States Bureau of Fisheries steamer “Albatross” in connection
with the biological survey of San Francisco Bay during the
years 1912-1913. Univ. Calif: Publ. Zool. 23: 1-470;
pits. 1-50; 165 text figs.

VICENTE, N.

1962. Premiéres observations sur un trematode parasite dans
le systéme nerveux d’ Aplysia fasciata. Recl. Trav. Stat.
mar., Endoume, 41: 303 - 305

WEYMOUTH, FRANK W.

1910. Synopsis of the true crabs (Brachyura) of Monterey

Bay, California. Stanford Univ. Publ., Univ. Ser. 4: 61
WINKLER, LINDSAY ROBERT & BERNARD E. TILTON

1962. Predation on the California sea hare, Aplysia califor-

nica, by the solitary great green sea anemone Anthopleura

xanthogrammica, and the effect of sea hare toxin and acetyl-
choline on anemone muscle. Pacif. Sci. 16 (3): 286 - 290

Vol. 10; No. 4

Vol. 10; No. 4

THE VELIGER

Page 379

Taxonomic Placement of Coralliophila incompta Berry, 1960,

With the Proposal of a New Genus, Attiliosa

(Gastropoda : Muricacea )

5 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 53; 5 Text figures)

DurING THE PROCESS of sorting the mollusks obtained
by the Puritan-American Museum Expedition to western
Mexico of 1957, two specimens of an apparently new
species of stenoglossid gastropod were found in a dredge
sample from off Isla Angel de la Guarda. The concho-
logical and opercular characters of these specimens sug-
gested to us at the time taxonomic placement near the
genus Cantharus, in the Buccinidae. These specimens,
along with other unidentified material from this vast
collection, were set aside for future study. Shortly there-
after, Coralliophila incompta was described by BERRY
(1960) from specimens that also were dredged off Isla
Angel de la Guarda, in the Gulf of California. The
Puritan specimens, together with additional specimens
in our collection taken by Mexican fishermen operating
out of Guaymas, Sonora, are conspecific with BERRy’s
taxon for which only an abbreviated diagnosis, lacking
an illustration, has been published. The holotype of
Berry’s taxon is figured herein, courtesy of Dr. James
H. McLean.

te EL we

Figure 1

Radular rachidian tooth of Cronia amygdala (KtENER),
type species of Cronia H. & A. Apams; greatly enlarged, dorsal
view, after Cooke (1919, fig. 33).

My colleague, Anthony D’Attilio, who is an ardent
student of the Magilidae, recently expressed his opinion
that Berry’s incompta was not referable to the genus
Coralliophila. Mr. D’Attilio submitted three live-taken
specimens to Mr. Masao Azuma of Nishinomiya, Japan,
who kindly provided us with a description and drawing
of the radulae that he extracted from the specimens.
Inasmuch as the shell of zncompta is not characteristic
of Coralliophila and a radula is not known to occur in
the Magilidae [Coralliophilidae], the placement of BEr-

(dull,

Figure 3
Figure 2

Figures 2, 3
Radular teeth of Aftiliosa incompta (BERRY), type species of
Attiliosa, new genus; greatly enlarged, dorsal view.
Figure 2: Lateral tooth. Figure 3: Rachidian tooth
(Drawings courtesy of Mr. Azuma)

Ry’s taxon in that family would appear to be untenable.
The radula was found to be similar to those possessed by
species in the genus Cronia H. & A. Apams (1853) of
the muricacid subfamily Thaisinae (see Text Figures 2
and 3). The type species of C’ronia is Purpura amygdala
KIENER (1836), by monotypy (see Text Figure 1).

Page 380

THE VELIGER

The placement of “Coralliophila” incompta BErry in
the subfamily Thaisinae near the genus Cronza, necessi-
tates the proposal of a new generic name:

Attiliosa EMERSON, new genus

Diagnosis: A thaisinid with a semi-ovate shell of mod-
erate size; spire acuminated; aperture large with spiral
lirations within the outer lip, and with small lirations
anteriorly placed on the inner lip; anterior canal rela-
tively long; base with siphonal fasciole. Radula with 3
major and 2 lesser cusps on central teeth (Text Figure 3).
Operculum “purpuroid,” 7. e. with a lateral nucleus and
a prominent marginal inner callus (Text Figures 4 and 5).

Type Species: Aftiliosa incompta (Berry, 1960).

Remarks: The type species of Aftiliosa is characterized
by the possession of prominent lirations within the aper-
ture on the outer lip, by the development of a broad
siphonal fasciole, and by having an extended, somewhat
recurved siphonal canal.

It is my pleasure to name this new taxon in honor of
my valued friend, Mr. Anthony D’Attilio.

Figure 4

Figures 4, 5

Operculum of Attiliosa incompta (BERRY) ; greatly enlarged.
Figure 4: Inner surface. Figure 5: Outer surface.

Vol. 10; No. 4

Attiliosa incompta (Berry, 1960)
(Plate 53, Figures 1 - 5; Text Figures 2 - 5)

Coralliophila incompta Berry, 1960, pp. 119 - 120;
type locality: “20 mi. off Puerto Refugio, Isla Angel de
la Guarda, Baja California,’ Mexico; B. W. Walker, 20
to 21 Apr. 1953; holotype here illustrated, Plate 53,
Figure 1. DuSHANE & Poorman, 1967, p. 429, Guaymas,
Sonora, Mexico, dredged in 10 - 45 fathoms, on rock and
gravel bottom.

Additional Records: Off southeast end of Isla Angel de la
Guarda, Mexico, 29° 01’00” N, 113° 07’00” W, dredged
in 15-17 fathoms, on sand? bottom, 1 live-taken speci-
men, May 20, 1957 (Puritan expedition, station 167;
Emerson, 1958), A. M. N.H. No. 77128.

Off southeast end of Angel de la Guarda Island,
Mexico, 29° 01’30” N, 113°07’20” W, dredged in 16
to 17 fathoms, on sand?, 1 live-taken specimen, May 20,
1957 (Puritan expedition station 168; EmErson, 1958),
A. M.N.H. No. 77143.

Gulf of California, Mexico, trawled by fishermen,
cruising out of Guaymas, Sonora, 6 live-taken specimens,
ex. Purdy collection, A. M.N.H.No. 138277; 4 speci-
mens, Germer collection.

Holotype deposited in Stanford University Paleontology
Type Collection No. 9512; ex collection of S.S. Berry,
No. 18768.

ACKNOWLEDGMENTS

In addition to Messrs. Azuma and D’Attilio, I am greatly
indebted to the following individuals for courtesies of
various kinds: Mrs. Helen DuShane of Whittier, Cali-
fornia, Mrs. John Germer of Maywood, New Jersey,
Dr. James H. McLean of the Los Angeles County Muse-
um of Natural History, Mr. and Mrs. Ben Purdy of
San Diego, California, and Mr. William E. Old, Jr. of
the American Museum of Natural History.

Mr. A. A. Olsson of Coral Gables, Florida kindly read
the manuscript.

Explanation of Plate 53

Attiliosa incompta (BERRY, 1960)

Figure 1: Holotype (x1)

Figure 2: Mature specimen from Puritan Station 167 (American
Museum of Natural History No. 77128); (x2)

Figure 3: Juvenile specimen, with immature lip, Gulf of Califor-
nia (American Museum of Natural History No. 138277); (x2)

Figures 4, 5: Mature specimen from Puritan Station 168 (Ameri-
can Museum of Natural History No. 77143); (x2). Note
prominent siphonal fasciole and lirations extending within the
aperture in Figure 4. Exterior of shell largely covered with foreign
matter in Figure 5.

Tue VE IcER, Vol. 10, No. 4 [Emerson] Plate 53

Holotype
Coralliophila..incompta

Figure 3

Figure 5

Vol. 10; No. 4 THE VELIGER Page 381

LITERATURE CITED

ApaMs, Henry & ARTHUR ADAMS
1853-[1854]. The genera of Recent Mollusca. London, vol. 1
[pp. 1-256, 1853; pp. 257 - 484, 1854]

BERRY, SAMUEL STILLMAN
1960. Notices of new eastern Pacific Mollusca — IV.
Leafl. in Malacol. 1 (19): 115 - 122 (31 December 1960)

Cooxg, A. H.
1919. The radula in Thais, Drupa, Morula, Concholepas,
Cronia, Iopas, and the allied genera. Proc. Malacol. Soc.
London 13 (3, 4): 90-110; 38 text figs. (August 1918)

DuSwHan_E, HELEN & Roy PoorRMAN
1967. A checklist of mollusks for Guaymas, Sonora, Mexico.
The Veliger 9 (4): 413-441; 1 map (1 April 1967)

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; 25 pp.; 9 figs. (22 July 1958)

Kriener, Louis CHARLES
1836. | Spécies géneral et iconographie des coquilles vivantes.
Genre Pourpre. Paris, 8: 1 - 151; plts. 1 - 46

Page 382

THE VELIGER

Vol. 10; No. 4

A New Cowrie Species from the Philippines

CRAWFORD N. CATE
12719 San Vicente Boulevard, Los Angeles, California 90049

FRANZ ALFRED SCHILDER

University of Halle, German Democratic Republic

(Plate 54)

Durine Aucust 1967, Mr. Fernando G. Dayrit, Univer-
sity of the Philippines Village, Diliman, Rizal, Philippines,
received a curious small cowrie which had been collected
alive off Laminusa Island, Siasi Group, in the south-
western Philippines (see Cate, 1966; map, p. 237). The
small, slender shell superficially recalls Notadusta rabaul-
ensis SCHILDER, 1964, but some of its shell characters,
especially the reduced fossula and the dorsal blotch, prove
it to be related to Erronea pallida (Gray, 1824) and more
particularly to its south-eastern race E. p. insulicola
SCHILDER & SCHILDER, 1938. The differences, however, are
so striking that we think the shell belongs to a distinct
species that is new to science; we propose to call it:

Erronea stohleri CATE & SCHILDER, spec. nov.

Holotype: The unique holotype of Erronea stohleri is
15.1 mm long, 8.2 mm broad, and 6.9mm high, and shows
18 labial and 17 columellar teeth (the two terminal ridges
excluded, but a small intercalated columellar tooth in-
cluded); therefore the characters, as tabulated by M.
SCHILDER (1967, p. 373) read for this species as follows:
13) D4 Orillia ps:

The adult shell is elongate pyriform, only slightly cal-
lous, with the spire projecting from a shallow umbilicus;
the equally narrow right margin is separated by a rather
indistinct dorsal rim, the left margin is regularly convex;
the terminal impressions are distinct in front, but obsolete
behind; the base is rather flattened, impressed in front,
inner lip slightly convex behind with its posterior beak
acuminate and bent to the left; the aperture is narrow,
distinctly widened in front and distinctly bent behind; the
labial teeth are short even in the declivous anterior ex-

tremity; the terminal tooth is composed of 2 long, slender
convergent ridges the anterior of which borders the out-
let; it is well separated from the columellar teeth, the
anterior of which are much coarser than the short central
ones and the rather oblique posterior ones (the last tooth
lies on the base of the posterior beak the inner margin of
which is smooth) ; the fossula is much reduced, as the
three strong anterior columellar teeth project towards the
aperture and cross the vertical plane plate as far as its
inner margin, forming three continuous vertical ribs with-
out any swelling interiorly; the columella shows no longi-
tudinal sulcus, it is constricted in front and crossed here
by vertical ribs similar to those of the fossular plate, but
ventricose behind with traces of interior denticles termi-
nating the obsolete transversal ribs.

Dorsum grey, without transversal zones, irregularly
freckled with often confusely confluent fulvous-brown
specks, and adorned with a large square fulvous-brown
central blotch; margins and base whitish; outer margin
with six small chestnut spots which do not extend to the
base, left margin with four obsolete spots in front only;
extremities suffused with pale greyish fulvous, anterior
extremity with two blackish brown spots which are re-
stricted to the dorsal area, posterior extremity practically
unspotted; teeth white without any traces of colored
striae; spire pale fulvous.

Type Locality: The animal was collected alive in ap-
proximately two fathoms of water off Laminusa Island.

Type Repository: The holotype is deposited in the Cali-
fornia Academy of Sciences Geology Department Type
collection, where it bears the number 13101. The holotype
is figured on Plate 54.

[CaTE & SCHILDER] Plate 54

Erronea stohleri CATE & SCHILDER

Lateral, dorsal and ventral views of the holotype

photographs by TrosTEL

Vol. 10; No. 4

THE VELIGER

Page 383

Discussion: Size and shape of the holotype of Erronea
stohleri recall to mind Notadusta rabaulensis ScHILDER,
1964, fig. 1; see also 1966, figs. 1-3), but E. stohleri
differs from it by the coarser anterior columellar teeth
which cross the narrow vertical fossula without any im-
pression instead of being compressed in the central part
along the fossula and thickened within as a row of inner
denticles. Besides, in E. stohleri the dorsum is not zonate,
it shows irregularly confluent specks instead of regularly
distributed round spots, and a large central blotch never
observed in any species of Notadusta; the scarce lateral
spots do not extend to the base, and the teeth of both lips
are far less numerous.

These and other characters agree with Evronea pallida
insulicola SCHILDER & SCHILDER (1938, p. 148) ; however,
E. stohleri is much smaller, more elongate, less callous so
that the right margin is vertical and the base flattened,
with the hind top of the inner lip more acute, and with
finer and more numerous teeth; the features of the fossula
agree with those of E. p. insulicola, whereas in typical E.
p. pallida (Gray, 1824) the anterior columellar teeth
project far more so that they become rather semicircular
within, not reaching the inner border of the fossular plate.

There is a remarkable parallelism of the morphological
characters in relation to the geographical distribution:
The central Evronea pallida insulicola, which lives on the
north coast of Java, seems to be intermediate morpho-
logically between the eastern, small, delicate E. stohleri
from the Sulu Sea (southern Philippines), and the west-

tai ty 4 =
a an §

ern callous E. p. pallida, which is distributed from the
Gulf of Thailand via Singapore and Burma to India and
the Persian Gulf. The common ancestor, E. vredenburgi
ScHILDER (1927) with a distinctly concave real fossula
is restricted to the south coast of Java from the Sunda
Strait to Bali.

This species is named in honor of Dr. Rudolf Stohler
of the University of California, Berkeley, California, in
recognition of his tireless efforts in promoting Malaco-
logy, as the author of many valuable papers on mollusca
and as the founder and editor of “The Veliger.”

LITERATURE CITED

Cate, CRawFrorp NEILL
1966. Philippine cowries.
pits. 32-45; 3 text figs.;

ScHILDER, FRANz ALFRED
1964. | Zur Kenntnis der Cypraeidae: 7. Eine neue Notadusta
aus Melanesien. Arch. Molluskenk. 93 (3/4): 141 - 144;
2 figs. (20 June 1964)

1966. Rediscovery of a unique cowry. Hawaiian Shell

News (n. s.) 6: figs. 1-3 (July 1966)
SCHILDER, FRANZ ALFRED, & MARIA SCHILDER

1938 - 1939.

ScHILDER, Maria

1967. Length, breadth, and dentition in living cowries.
Veliger 9 (4) : 369-376; 1 diagram

Proc. Malacol. Soc. London, 23(3 - 4):

The Veliger 8 (4): 234 - 264;
1 map. (1 April 1966)

Prodrome of a monograph on living Cypraeidae.

The
(1 April 1967)
119 - 231.

a .

Page 384

THE VELIGER

Vol. 10; No. 4

Studies on the Mytilus edulis Community

in Alamitos Bay, California
III. The Effects of
Reduced Dissolved Oxygen and Chlorinity Concentrations

on Survival and Byssus Thread Formation

BY

DONALD J. REISH

AND

JOSEPH L. AYERS, Jr.

Department of Biology
California State College at Long Beach, Long Beach, California 90804

(2 Text figures)

DurRING THE couRSE of studying the ecology of the
Mytilus edulis LinnaEus, 1758, community in Alamitos
Bay, California (ReisH, 1963, 1964a, 1964b) the senior
author was interested in learning what biological or phys-
ical factors would limit or destroy the population. No
animal was or has subsequently been observed to prey
on M. edulis in this area. Estinc et al (1964) reported
that crabs fed upon M. edulis in the laboratory and in
suspended sea cages in English waters. An occasional
crab, Hemigrapsus oregonensis (DANA, 1851), has been
taken from mussel beds in Alamitos Bay, but it is not
known whether or not it will feed upon it. No empty M.
edulis shells have been observed with circular holes pres-
ent as a result of radular drilling by snails.

A mass mortality of Mytilus edulis and associated or-
ganisms occurred in Alamitos Bay in 1962 following an
extensive red tide bloom of the dinoflagellate Gonyaulax
polyhedra Stein (RetsH, 1963). It was not known if the
mussels were killed directly by the accumulation of
toxic material in the body or indirectly by the decrease of
dissolved oxygen in the water.

The chlorinity of the sea water throughout the bay is
that of normal sea water (19.2%,). Decreases in chlorin-

ity in Alamitos Bay occur only following rains as the
result of run-off from the surrounding area (STONE &
ReisH, 1965) ; however, the chlorinity returns to normal
levels rapidly. It is not known whether or not these re-
duced chlorinities in Alamitos Bay are lethal to Mytilus
edulis.

Two additional causes of community destruction may
be cited. Large clumps of Mytilus edulis on floating docks
may be dislodged by their own weight or by heavy waves
and fall to the bottom. Dead mussel shells have been taken
in bottom samples beneath these docks, but, of course,
the cause of death was unknown. People utilize mussels as
fishing bait.

Of the different possible causes of destruction to the
mussels discussed above, two lend themselves to labora-
tory experimentation: reduced chlorinities and dissolved
oxygen concentrations. The initial purpose, therefore, of
this investigation was to determine the lethal limits of
these two environmental factors on Mytilus edulis under
controlled laboratory conditions. During the course of
conducting the preliminary experiments, it was discovered
that the production of byssal threads, which were used
for attachment to the flask, was related to varying con-

Vol. 10; No. 4
100
go
80
70

60

Average Number of Byssal Threads Produced and Per Cent Survival
ra

% of Survival and
Number of Threads

THE VELIGER

Page 385

12 14 16 19
Chlorinity in %,

Figure 1

Survival and average number of byssal threads produced by Mytilus edulis under reduced concentrations of chlorinity during
a 14 day period at 15°C to 16°C

centrations of chlorinity and dissolved oxygen. Therefore,
it was possible not only to investigate the lethal concen-
trations of dissolved oxygen and chlorinity but also to
ascertain intermediate levels of the condition of the ani-
mal by counting the number of byssal threads produced
during the experimental period.

MATERIALS anp METHODS

Specimens of Mytilus edulis were collected from the boat
floats in Alamitos Bay marina. They were cleaned of
fouling organisms from the surface of the shells and all
their byssal threads were cut. Only specimens measuring
from 15 to 20mm in width were used. Preliminary studies
indicated these smaller sized mussels produced a greater
number of byssal threads per unit time than larger ones,

and since the gonads were immature, no spawning would
occur during the course of the experiment. The speci-
mens were then placed in aquaria in a cold bath with
water temperatures of 15°C to 16°C. Specimens were
left in the aquaria for 3 days. The specimens used in the
experiments were cleaned again and the byssal threads
cut. One specimen was placed in an Erlenmeyer flask
with 150ml of filtered sea water. Ten specimens were
used for each concentration of dissolved oxygen and
chlorinity. The animals were not fed during the course
of the experiment.

Decreased chlorinities were made by adding an appro-
priate amount of distilled water. Chlorinities were de-
termined by titrating with silver nitrate as outlined by
Barnes (1959). The following chlorinity concentrations
were used: 19.0 (control), 16.0, 14.0, 12.0, 10.0 and
8.0%. . The top was closed with a rubber stopper.

Page 386

THE VELIGER

Vol. 10; No. 4

Decreased dissolved oxygen concentrations were ob-
tained by flushing the overlying air in the Erlenmeyer
flask with nitrogen gas through hypodermic needles which
pass through the rubber stopper according to the proce-
dure outlined by REisH & RicHarps (1966). Three addi-
tional flasks, each with a mussel, were used for dissolved
oxygen measurement during the course of the experi-
ment. The dissolved oxygen concentration was determined
with an electrode following the procedure described by
REISH & RicHarps (op. cit.). The dissolved oxygen values
given in Figure 2 were determined by averaging the
values measured within a series. These dissolved oxygen
values were 4.5 (control), 2.1, 0.9, 0.6, 0.3, and 0.2 ppm.

The Erlenmeyer flasks containing the mussels were
placed in the cold bath which maintained the tempera-
ture at 15°C to 16°C. Specimens were examined daily

go ry

80

70

60

50

40

go

Average Number of Byssal Threads Produced

20

2

for death and to count byssal threads. The byssal threads
were attached along the sides of the flasks by the mussels.

Death was ascertained during the course of the exper-
iment by noting the failure of the shell to close. When
this condition occurred, the specimen was left another
day as a check to see whether or not tissue decomposition
had commenced. At the termination of the experiments,
which ran for 14 days, the condition of the remaining
specimens was determined.

DATA

The data for the effects of reduced chlorinities on sur-
vival and byssal thread production are summarized in
Figure 1. It can be seen from this figure that Mytilus

3 4 5

Dissolved O: (ppm)

Figure 2

The average number of byssal threads produced by Mytilus edulis under reduced concentrations of dissolved oxygen during a 14 day
period at 15°C to 16°C

Vol. 10; No. 4

THE VELIGER

Page 387

edulis is capable of living for 14 days in chlorinities as
low as 10.0%. The 14 day median tolerance level (TLm)
is approximately 8.6%. However, these data are of inter-
est when compared to the production of byssal threads
during this 14 day period. The number of threads formed
decreased at chlorinities between 14%, and 16%. The

number of byssal threads continued to decrease to 10%. —

No threads were secreted by these mussels at 8%, but
30% of the specimens were still alive at the end of the
14 day period.

The relationship between the number of byssal threads
produced and the dissolved oxygen concentration is given
in Figure 2. Nearly all animals survived at all concentra-
tions of dissolved oxygen. The difference between the
number of threads produced at 0.9 ppm and 0.6 ppm
is striking: 80 per animal per 14 day period as compared
to 9.

DISCUSSION

Small specimens of Mytilus edulis from Alamitos Bay are
capable of surviving low concentrations of chlorinity
and dissolved oxygen for a 14 day period at 15°C to 16°C
under laboratory conditions. While the majority of the
test organisms survived the reduced concentrations of
chlorinity and dissolved oxygen, the specimens in the
lower concentrations produced fewer byssal threads. The
response of M. edulis in these lowered concentrations
was similar; the two shells remained tightly closed most
of the time. The shells of specimens in the controls of
higher concentrations would be open.

Apparently Mytilus edulis is affected physiologically,
as measured by byssal thread production, by chlorinities
between 14%, and 16%,, but is capable of surviving much
lower levels. Chlorinities as low as 2.7%, have been
measured in Alamitos Bay (STonE « ReisH, 1965).
Since there are no rivers emptying into Alamitos Bay,
recovery to normal chlorinities is rapid following the
fresh water run-off from rain. If the rain falls over a week
or more, which may occur some years, then the effects
of this run-off could become an important survival factor
to a population of M. edulis. It is of interest to note that
populations of M. edulis occur in chlorinities of 8%. in
the Baltic Sea (ScHLIEPER, 1955).

The survival of Mytilus edulis in a reduced dissolved
oxygen environment under laboratory conditions has not
been studied previously. During a red tide bloom of the
dinoflagellate Gonyaulax polyhedra in Alamitos Bay in
1962 (RetsH, 1963), the dissolved oxygen content of the
water was lowered as a result of the decomposition of
dead protozoans and other organisms. Dissolved oxygen

concentrations as low as 0.1 ppm were measured, but it
was unknown how long this low level persisted.

The relationship of byssal thread production in Mytilus
edulis to altered environmental conditions has not been
observed in the past. It seems logical to assume that the
fewer number of threads secreted would reflect a lower
metabolic rate. A reduced metabolic rate, as measured by
oxygen consumption, was measured in mussels living in
chlorinities below their range of tolerance (Bouxin, as
reviewed by Kinng, 1964). Similar decreases in meta-
bolic rate employing other measures of activity, but with
a similar experimental procedure, have been noted in
polychaetes (ReisH, 1966) and in the wood-boring iso-
pods Limnoria (ANDERSON & ReIsH, 1967). Four species
of polychaetes, Nereis grubei (Kinserc, 1866), Dorvillea
articulata (HarTMAN, 1938), Neanthes arenaceodentata
(Moore, 1903), and Capitella capitata (Fasricius,
1780), failed to eat algae under sublethal concentrations
of dissolved oxygen. Fecal pellet production in Limnoria
tripunctata (MeEnzirs, 1951), L. quadripunctata (Hout-
HuIs, 1949), and L. lignorum (RatHKE, 1799), was
directly related to the dissolved oxygen concentration.
The daily egestion rate decreased markedly in dissolved
oxygen concentrations of less than 2.0 ppm.

It is apparent from these data that Mytilus edulis can
withstand lowered chlorinities and dissolved oxygen con-
centrations for extended periods of time. The shells
remain closed under these altered conditions. Correla-
ting these data with field conditions, it is evident that
if lowered environmental conditions extend for only a
short period of time, then M. edulis would be able to
withstand these conditions by keeping its shells closed.
If, on the other hand, reduced dissolved oxygen concen-
trations or chlorinities extend over a longer period of
time, such as a week or two or more, then the survival
factor for M. edulis would become critical. Since some
knowledge of the limiting values of dissolved oxygen
and chlorinity on M. edulis are now known under labo-
ratory conditions, it would be of particular value to study
more precisely the survival of M. edulis under adverse
conditions in the field.

SUMMARY

1. The survival of Mytilus edulis under reduced concen-
trations of dissolved oxygen and chlorinity at 15°C to
16°C was studied under laboratory conditions.

. The 14 day TL» value for chlorinity was 8.6%c.

3. Nearly all specimens of Mytilus edulis lived for 14

days in reduced dissolved oxygen concentrations as low
as 0.2 ppm.

NO

Page 388

THE VELIGER

La

Vol. 10; No. 4

4. The number of byssal threads produced was found
to be directly related to concentration of chlorinity
and dissolved oxygen. The number of threads pro-
duced decreased sharply between 14.0%, and 16.0%,
chlorinity and 0.6 ppm and 0.9 ppm dissolved oxy-
gen.

LITERATURE CITED

ANDERSON, JAcK W. & Donatp J. REISH
1967. The effects of varied dissolved oxygen concentrations
and temperature on the wood-boring genus Limnoria. Mar.
Biol. 1: 56 - 59
BarnEs, HAROLD

1959. Apparatus and methods of oceanography, part one:
chemical. Intersci. Publ. Inc., New York, 341 pp.

Esiine, F. J., J. A. Kircuinc, L. Muntz « M. C. Taytor

1964. The ecology of Lough, Inc. (Ireland). XIII. Experi-
mental observations of the destruction of Mytilus edulis and
Nucella lapillus by crabs. Journ. Animal Ecol. 33: 73 - 82

KINNE, Otto
1964. The effects of temperature and salinity on marine and
brackish water animals. II. Salinity and temperature, salinity
combinations. In: Oceanogr. Mar. Biol. Ann. Rev., edited by
H. Barnes, George Allen and Unwin Ltd., London, 2: 281 - 339

RetsH, Donatp J.

1963. | Mass mortality of marine organisms attributed to the
“red tide” in southern California. Calif. Fish & Game 49:
265 - 270

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)

1966. Relationship of polychaetes to varying dissolved oxy-

gen concentrations. Third Internat. Conf. Water Pollut.
Res., Munich, sect. 3, paper 10, pp. 1-10

ReisH, Donatp J. & THomas L. RicHarps

1966. A technique for studying the effect of varying con-
centrations of dissolved oxygen on aquatic organisms. Air
and Water Pollut. Int. journ. 10: 69-71

ScHLIEPER, C.

1955. | Uber die physiologischen Wirkungen des Brackwassers
(nach Versuchen an der Miesmuschel Mytilus edulis) .
Kiel. Meeresforsch. 11: 22 - 33

Stone, ALFRED N. & Donatp J. REISH

1965. The effect of fresh-water run-off on a population of
estuarine polychaetous annelids. Bull. So. Calif: Acad. Sci.
64: 111-119

Vol. 10; No. 4

THE VELIGER

Page 389

West American Mollusk Types

at the British Museum (Natural History)

IV. CarpentTer’s Mazatlan Collection

A. MYRA KEEN
Department of Geology, Stanford University, Stanford, California 94305

(Plates 55 to 59; 171 Text figures)

IT Is NOW MORE THAN a century since the publication of
Philip Carpenter’s “Catalogue of the Mazatlan’ shells in
the British Museum collected by Frederick Reigen.” This
was a pioneering work, for Carpenter was in the forefront
of those who saw potential benefits from the study of
geographic distribution. He realized that here, for the
first time, was a reasonably complete collection from a
single locality (or at least a limited area), and he did
his best to examine it in detail. He enumerated nearly 700
species of mollusks from this West Mexican station and
described some 255 kinds as new. Unfortunately, the cata-
logue was published without illustrations, and the 60
plates of camera lucida drawings of the smaller forms
that he did prepare remained (with minor exceptions) in
manuscript until recently, when the Paleontological Re-
search Institution undertook their reproduction (BRANN,
1966). These plates did not include figures of the larger
shells, of which there were nearly a hundred species.
BRANN cites published photographs of 6 of these types,
and a few others were figured during the 1860’s by the
iconographers Reeve and Sowerby.

Carpenter’s catalogue was the second of his 3 major
publications that have been fundamental to all later stu-
dies of West American mollusks. However, a century had
to elapse before a systematic survey of the actual type
specimens that he worked with was initiated. To Dr.
Katherine Palmer we owe a lasting debt for having per-
severed in the task of hunting out those types that had
been deposited at institutions other than the British Mu-

1 Technically, the Mexican place-name Mazatlan should be
written — as pronounced — with the final syllable accented.
However, to be consistent with Carpenter’s usage, I have
omitted the accent mark throughout this paper. The word
“mazatlan,” an Amerind term, means “place above the corn-
fields.”

seum. All of us had assumed that she would monograph
in similar fashion the British Museum material. There-
fore, when I went to England for study in 1964, I had no
intention of intruding upon her domain of research. The
drawings and photographs of Carpenter material that I
then made were meant to be for my own use. However,
now that Dr. Palmer (in Brann, 1966, p.3) has indicated
that with a trilogy of publications completed she plans
no further research on the subject, I realize that my work
may supplement the available printed material and can
supply, at long last, figures of virtually all of the non-
microscopic types. It can also provide some data on sizes
of the specimens and an attempt to interpret the species
in terms of our modern classification.

One point that I made in a foreword to Brann’s paper
perhaps should be repeated here, that although Carpen-
ter referred throughout his published catalogue to the
“Liverpool Collection,’ the material was not at any time
under the care of the Liverpool Museum. Rather, Liver-
pool was the docking area to which the major part of
Reigen’s collection was shipped for sale. Reigen sent an-
other smaller fraction to Le Havre, France. Sylvanus
Hanley purchased a part of the latter stock, and some of
these specimens later found their way to the British Mu-
seum. The part of the “Liverpool Collection” that Car-
penter bought was worked over by him, then split into
duplicate series, of which the first and best was offered
to the British Muscum (Natural History) under certain
conditions -— that the collection remain intact, that
it be made available to qualified students, and that it
remain as he had arranged and mounted it on glass
slides numbered to correspond with the entries in his
printed “Catalogue, a copy of which went with the col-
lection. ‘These conditions were accepted, and the collec-
tion has remained as a unit.

Page 390

Some of my initially incomplete observations were
supplemented during two later brief trips to the British
Museum, but there still remain some unanswered ques-
tions for future research. The reader, to make full use of
the present report, should have at hand the work by
Brann, and a copy of Carpenter’s “Catalogue,” now made
available through a reprint edition by the Paleontological
Research Institution. Carpenter’s drawings having in-
cluded details of sculpture, it would be duplication of
effort if I did the same; I have therefore reduced mine
to little more than simple outlines that show the shape
of the shells under standard orientation (Carpenter often
tilted specimens to bring out apertural features or hide
defects). It is gratifying to find how closely, in general,
our two sets of camera lucida sketches agree. There has
been a tendency to distrust Carpenter’s figures because
some of the few that were published looked so amateur-
ish. They prove, however, not grossly inaccurate, though
perhaps a little distorted, and this gives us more confi-
dence in his drawing of type specimens now lost or
deteriorated.

Several problems emerge as one examines the material
and the literature about it. First, the date of publication
of the “Catalogue.’’ Many workers, including my younger
self, interpreted the date at the end of each printed sig-
nature as the date of publication of a separate part.
IREDALE (Proc. Malac. Soc. London, vol. 12, p. 36, 1916)
argued that the date of the entire work must be post-June
1857 for the official edition and later for the Warrington
edition issued by Carpenter. In 1961, when I was pub-
lishing a paper in the British Museum (N.H.) Bulletin
series, I was reluctantly obliged to accept IREDALE’s con-
clusion, for the editor could find no evidence in the British
Museum archives of any publication of the work as parts
and I could produce no contemporary testimony on the
point. Only recently did I notice a statement by Carpen-
ter that should have settled the question long ago. In the
bibliographic list introducing a reprint of his papers,
CaRPENTER (1872, p. xi) cited the “Catalogue of the
Reigen Collection, the first edition with preface arranged
by Dr. J. E.Gray .. ” and the “Second edition, with
author’s preface, accompanying duplicate collections of
the shells, published simultaneously.” Thus, the publi-
cation date for both editions must be accepted as not
earlier than June, 1857.

A second problem has to do with the area covered by
the collection. Did all of it come from the immediate
vicinity of Mazatlan? Mr. A. M. Strong (Bull. Southern
Calif. Acad. Sci., vol. 48, prt. 2, p. 73, 1949) has noticed
(and my own observations are in harmony with his) that
the microscopic shells seem to match up with those now
taken on the west side of the Gulf of California, notably

THE VELIGER

Vol. 10; No. 4

in the La Paz area. However, Major Rich, who met
Reigen in Mazatlan, had evidence that Reigen did much
local collecting, for he recorded that Reigen had so filled
his house with decomposing mollusks that the neighbors
appealed to the police (CARPENTER, 1864, p. 540). The
part of the collection that is now in the British Museum
looks to be mainly live-taken, certainly not beach-worn.
Carpenter indicated that the microscopic forms were in
crevices in large Spondylus and Chama. He lamented
how few of these there were, for before he got to Liver-
pool, most of the large shells had been sold to an inn-
keeper who wanted them for decoration; it was a bitter
disappointment to Carpenter to learn that the washings
from these were discarded, thus destroying the chance for
the study of the countless small attached and nestling
specimens. Whether the large shells had all been gathered
at Mazatlan or whether they had been brought there from
elsewhere in the Gulf is a problem for further research.
Lime for mortar used in building was gotten from shells
— Carpenter commented on this (1857b, p. 153). We
therefore need to know something more about the distri-
bution, past and present, of large Spondylus and Chama
and the amount of commercial exploitation.

Precise measurements of the types turned out to be
more of a problem than I had anticipated when I set out
to photograph each larger specimen with a millimeter
scale in the picture and to calibrate each microscope-lens
combination by reading the resulting magnification of a
millimeter scale. In his preface Carpenter stated that all
his measurements were in inches and decimal parts. Later
(1865 a, p. 133), he described the inch as 2.53 cm. We
normally take it as 2.54cm or 25.4mm, but this is not a
critical difference. As soon as one begins converting his
decimal parts to millimeters one realizes that he was
dealing with very small shells, and it is remarkable that
he was able to read his scale as accurately as he did.
However, when I checked my readings of dimensions
against his I found some disharmonies. Fortunately, I had
both photographed and drawn a number of the types,
and this gave a basis for a comparative analysis that
showed my camera lucida readings to be about 10% too
high and his to be slightly low. Because my millimeter
scale was thinner than the glass slides, I would have
changed focus (and therefore image size) between taking
the scale reading and making the drawing. If he used the
same method for getting readings and if his scale was
thicker (which is likely, as plastic was not then available),
this would account for the average 15% discrepancy be-
tween his readings of size and mine. In any case the dis-
crepancy is not of a magnitude to be of real consequence
in making identifications, for it is within the range of
expected variations in size of specimens. In a few cases ~

Vol. 10; No. 4

Carpenter seems to have misread his scale, however, or
there is a typographical error in his text. I have called
attention to these in my discussions of species.

A minor problem has to do with the correct citation
of some of the specific names. Carpenter adopted the
practice — later followed by Dall in Bulletin 112 of the
U.S. National Museum -—- of citing subgeneric names
as if they were generic, making adjectival specific names
agree in gender with the subgeneric rather than the ge-
neric name. This has sometimes been a source of con-
fusion in the citations by later authors, notably for the
genera Terebra, Turbonilla, and Odostomia.

FORMAT

Carpenter’s catalogue was arranged according to a sys-
tem, but the sequence of family and ordinal groups was
markedly different from modern usage. He ranked chitons,
for example, as prosobranch gastropods. Once he had
arrived at his classification, he numbered the species and
also the mounts or tablets, and he saw no need for further
indexing. Maintaining the original material according to
his scheme is convenient and desirable in the museums
where it is housed, and the Brann publication was obliged
to follow the same pattern. For the present report it
seems preferable to attempt a fresh start. I have re-
arranged the species into modern systematic order insofar
as possible, with an alphabetical index for cross-refer-
encing. Each entry is given a paragraph number (in bold
face numerals). The name is cited exactly as Carpenter
spelled it (even to disagreement of the adjectival endings),
with page reference (unless otherwise noted) to the Maz-
atlan Catalogue. Following an equality sign is my inter-
pretation of the correct modern assignment. Tablet
numbers are as listed by Carpenter, with number of
specimens and their present type status. The entry
in square brackets is the size given by Carpenter
(longest dimension, whether height or diameter), con-
verted to millimeters. The Carpenter figures now avail-
able are cited as “BrANN,” without repeating the date.
My commentary includes notes on present condition of
the specimens, on morphology, on nomenclatural prob-
lems, and on figures by other authors.

It may seem that in the inventory of type specimens I
have been rather too liberal in the use of the term
“syntype.” The natural assumption on the part of authors
who figured or cited Carpenter’s material has been that
when a single specimen was on a Carpenter mount in
the British Museum it would automatically be the holo-
type, and any other distributed material would be para-
type. However, Carpenter did not select type specimens

THE VELIGER

Page 391

in the modern sense. He only said “the specimen” when
he had but a single shell. Thus, we must consider his type
lots, when there were two or more shells, syntypic. It is
desirable that the British Museum specimens be desig-
nated as lectotypes if they are in a good state of preser-
vation, and I have done this for many lots.

ACKNOWLEDGMENTS

A fellowship from the John Simon Guggenheim Founda-
tion provided my travel expenses during 1964 and 1965,
for which I am grateful. I wish also to thank the officials
and staff of the British Museum (Natural History) for
the privilege of use of the collections. Words of high
praise are due to the curators whose care of the material
during the last century has kept so much of the Carpen-
ter collection in prime condition, this in spite of two
world wars and numerous museum-cabinet transfers.

I am indebted for helpful advice and suggestions es-
pecially to Leo G. Hertlein, Ronald Ives, Allyn G. Smith,
Judith Terry, and Spencer Thorpe; three photographs
are from color slides made in 1962 by Elaine Reeves (now
Mrs. Francois Padovani); my camera lucida sketches
were turned into finished line drawings by Perfecto Mary,
artist-technician at Stanford, who was of great assistance
to me in the preparation of the plates. To all of these
persons and to the many others who helped in less evident
ways, my thanks.

PELECYPODA

ARCIDAE

1. Byssoarca vespertilio, p. 140
= Barbatia lurida (Sowersy, 1833)
(Plate 55, Figure 1)
Tablet 651, holotype. [33 mm].
No distinctive characters seem to justify this as a separate
form.

2. Arca bifrons, p. 134
= Anadara (Cunearca) bifrons (CARPENTER, 1857)
(Plate 55; Figures 18 a-c)

Tablet 631, fragments only. [43 mm].

Two syntypes from the Cuming Collection, now in the
British Museum’s Type Collection, are illustrated here;
the larger may be the one figured by Otsson (1961, plt.
9, fig. 3b) as “type”; it is here chosen as lectotype.

Page 392

MyTILIDAE

3. Mytilus multiformis, p. 118
= Brachidontes (Scolimytilus) multiformis (CARPENTER,
1857)
(Plate 55, Figures 5 a-b)

Tablets 540 - 552, numerous syntypes. [11.5 mm]. BRANN:
pit. 15, fig. 168 (also Oxsson, 1961, plt. 17, fig. 11, with
selection as lectotype).

Carpenter listed 13 tablets of syntypes; specimens from
543 and 551 are figured here; no. 546 is attached to a
specimen of the next species.

4, Mytilus palliopunctatus, p. 118 (as of DUNKER)
—=Choromytilus palliopunctatus (CARPENTER, 1857)
(Plate 55, Figures 4a-b)
Tablets 528 - 539 (539 here figured). [90 mm]. Brann:
pit. 15, fig. 167 (juveniles from tablet 528).
Although the specific name was credited to Dunker, he
had not validated it; thus it is to be attributed to Carpen-
ter.

5. Modiola brasiliensis var. mutabilis, p. 122
=Mytella guyanensis (LAMaRCK, 1819)

(Plate 55, Figures 20a-b)

Tablets 559 - 565 (tablet 559, 2 adolescent syntypes fig-
ured), 11 syntypes. [63 mm]. Brann: plt. 15, fig. 171-b
(juvenile from tablet 565).

If the West American form ever proves distinct from
that of the Atlantic, Carpenter's specific name would be
available for use.

6. Lithophagus aristatus gracilior, p. 129
= Lithophaga aristata (Dittwyn, 1817)

(Plate 55, Figure 9)
Tablet 601, 7 valves, syntypes [29mm]. Brann: plt. 16,
fig. 176.

None of the valves is now complete. However, they col-
lectively show that the form falls within the range of
variation of the species. Carpenter’s enlarged drawing
exaggerates the dorsal angle.

7. Lithophagus aristatus tumidior, p. 129
= Lithophaga aristata (Dittwyn, 1817)

(Plate 55, Figures 8a-b)

Tablets 602 - 603, 7 syntypes. [39mm]. BRANN: plt. 16,
fig. 176 (juvenile, enlarged, from tablet 602).

Tablet 603 has one large specimen (here figured), very
tumid, the incrustation wide as well as long; two smaller
specimens are of the more usual form of L. aristata.

THE VELIGER

Vol. 10; No. 4

8. Lithophagus calyculatus, p. 124
= Lithophaga (Stumpiella) calyculata (CARPENTER,
1857)
(Plate 55, Figures 16 a-b)

Tablet 571, holotype. [9.2 mm]. Brann: plt. 16, fig. 174.
Soot-Ryen (1955, p. 93; plt. 10, figs. 61-63) seems
to have identified this form correctly.

9. Lithophagus (Leiosolenus) spatiosus, p. 130
= Lithophaga (Leiosolenus) spatiosa (CARPENTER, 1857)
(Plate 55, Figures 7 a-b)
Tablet “605” [606*], syntype. [38mm].

Carpenter (p. 131) cites tablet 605 as having a young
shell and a fragment; on p. 550 he states that tablet 605
contains the original specimen presented by R. Derby-
shire. A specimen with the stated dimensions is in the
collection, the number on the tablet being “606*.”

Soot-Ryen (1955, plt. 10, fig. 59) has figured similar
material. Carpenter’s shell appears more slender because
it has curled in drying and the ventral margin has bent
inward.

10. Crenella coarctata, p. 123 (ex DUNKER MS)
= Gregariella coarctata (CARPENTER, 1857)
(Text figure 1)

Tablets 566 - 567, originally 6 syntypes. [4.5mm].
BraANN: plt. 15, fig. 172.

Carpenter credits the name to Dunker, who had not
validated it. On tablet 566 are 3 juvenile pairs about 2
mm long. One loose semi-adult 4mm long may be from
tablet 567. The other specimens are missing, represented
only by glue smears on the mount. The small shell re-
sembles G. chenui as to outline, with 15 radial ribs.

ISOGNOMONTIDAE

11. Isognomon janus, p. 151
=TIsognomon janus CARPENTER, 1857
(Plate 55, Figure 10)

Tablets 689 - 690, 4 syntypes. [28 mm, central area only].
Brann: plt. 17, fig. 206 (juvenile, tablet 689).

As the legend on tablet 690 states, “Jsognomon janus
on I. chemnitzianum” and there is but a single specimen
on the mount, I assumed that the J. janus syntype was
lost, but I photographed the tablet anyway. Later study
of the picture showed that not only does the specimen
have the dimensions cited by Carpenter but it has only
5 ligamental sockets, whereas I. chemnitzianum has 6 to
12. Therefore the shell in my figure evidently is one that
had been on J. chemnitzianum before mounting and is a
syntype of Carpenter’s species.

Vol. 10; No. 4

THE VELIGER

OSTREIDAE

12. Ostrea conchaphila, p. 161
= Ostrea conchaphila CarPENTER, 1857

Tablets 715 - 734, numerous syntypes. [26mm]. Tablet
728 here selected as lectotype.

The lectotype was figured by HERTLEIN & STRONG,
(1955, plt. 3, figs. 29 - 30) as “holotype.” Part of the syn-
type lot was figured by SowerBy in REEVE, 1871 (Conch.
Icon., vol. 18, plt. 28, sp. 69).

13. Ostrea palmula, p. 163
= Ostrea palmula CarRPENTER, 1857
(Plate 55, Figure 6)

Tablets 735 - 737; largest syntype, no. 737, here selected
as lectotype [58 mm].

The lectotype here selected was figured by HERTLEIN &
Stronc, 1946 (Zoologica, vol. 31, p. 76; plt. 1, fig. 14)
as “holotype.”

SPONDYLIDAE

14. Spondylus calcifer, p. 152
—=Spondylus calcifer CARPENTER, 1857

Tablets 692 - 699 (692, juvenile, with query). [About 180
mm].

Authors have identified this species correctly. None of
the syntype specimens is in good enough condition to
photograph, for they consist mostly of broken and cut
fragments, what was left after Carpenter had extracted
the borers and nestlers from the surface layers.

15. Plicatula penicillata, p. 155
= Plicatula penicillata CARPENTER, 1857
(Plate 55, Figures 2a-b)

Tablet 701, two syntypes. [28mm].

The smaller syntype shows brown pencilling on the outer
layer; the larger is white, triangular, fine-ribbed, attached
to a specimen of Crepidula aculeata. The form was fig-
ured by Sowersy in ReEveE (1873, Conch. Icon., vol. 19,
pit. 1, fig. 3).

ANOMIDAE

16. Placunanomia pernoides, p. 164
= Pododesmus (Tedinia) pernoides (Gray, 1853)
(Plate 55, Figures 14 a-c)
Tablets 745 - 748, two syntypes, complete, two odd valves,
one broken, all on one mount, of which the largest, no.
748, may be taken as lectotype. [46mm].
If, as seems possible, this form proves distinct from
Gray’s, a new name will be needed, for Carpenter’s spe-

Page 393

cies, named as new, seems to be conspecific with Gray’s.
The difference in outline between Mexican and Califor-
nian material may be only a matter of difference in
habitat, for the Californian specimens, favoring pholad
holes, are distorted by the cramped quarters, whereas
Mazatlan specimens, not having borers as associates, have
had to attach in more open sites.

17. Placunanomia claviculata, p. 166
= Anomia peruviana Orzicny, 1846
(Plate 55, Figures 15 a-b)
Tablet 750, syntype, both valves. [37mm]. BRANN: plt.
17, fig. 218.

This seems tobea thin white formof Anomia peruviana
grown on a smooth surface that had been encrusted with
a few small serpulid worm tubes. As the anomiid grew
out over the tubes a strong trace was left on the shell, and
the supposed “clavicle” of the lower valve seems to me to
be only a reflection of such a tube. Carpenter’s dimen-
sions must have been taken from the odd valve he men-
tions, for they are a third larger and are wider for the
length than the mounted pair.

ASTARTIDAE

18. Gouldia varians, p. 83
== Crassinella varians (CARPENTER, 1857)
(Text figure 2)
Tablets 415 - 419, 22 syntypes (none now on 418). [2.2
mm]. Brann: plt. 9, fig. 117.
Shells small, triangular, nearly smooth, with a brown
stripe or spot posteriorly.

CarDITIDAE (?)

19. Cardium lucinoides, p. 96
?==Cardita sp.
Text figure 3)

Tablet 463, holotype. [1.0mm]. BRANN: plt. 11, fig. 135
[““136” on plate].

The minute shell, a right valve, is probably too young
for positive determination. There are 12 ribs with beaded
sculpture and a tinge of brown color toward their ends;
the hinge is not well preserved, so that details as to den-
tition are unclear, but it does not seem to be a cardiid.

CorBICULIDAE

20. Cyrena olivacea, p. 114
— Polymesoda (Egeta) olivacea (CARPENTER, 1857)
(Plate 55, Figures 12 a-c)

Page 394

THE VELIGER

Vol. 10; No. 4

Tablets 505 - 511, 17 syntypes (2 here figured from tablet
509). [60 mm].

There is some variation in outline among the syntypes
as well as in size.

PisiopAE (SPHAERUDAE of authors)

21. Cycladella papyracea CarPENTER, 1865b, p. 270
=Sphaerium sp., cf. S. trigonare (Say, 1829)
(Plate 55, Figures 3 a-b)
Tablet 119 (fragments, as cited by CARPENTER, 1865;
now nearly disintegrated).

The type locality is Mazatlan, but the genus and species
was not described in the Mazatlan Catalogue. Carpenter
tentatively assigned the genus C’'ycladella to KELLODAE.
The holotype had been among the Reigen material pur-
chased by S. Hanley, and it came to the British Museum
from H. Harvey, Esq.;it is registered as no. 1907.12.30.
117. I am indebted to Mr. Peter Dance, then of the British
Museum staff, for the reallocation of the form as non-
marine; he ably demonstrated that hinge and sculpture
match those of specimens of Sphaerium from Mexico,
and Dr. Dwight Taylor suggested that Say’s species is the
most likely candidate for comparison. The generic name
Cycladella therefore may be removed from marine lists,
for it falls as a synonym of Sphaertum Scopou, 1777. The
specific name is not apt to prove useful either.

BERNARDINIDAE

22. Circe margarita, p. 81
= Bernardina margarita (CARPENTER, 1857)
(Text figure 4)

Tablet 412, one odd valve and 3 entire syntypes (the
largest here selected as lectotype). [1.8mm]. Brann:
plt. 9, fig. 114.

Microscope examination of the type lot in 1964 corrects
my hand-lens assignment to Lasaea (KEEN, 1958, p. 622).
The hinge is close to that of Bernardina bakeri Dat,
1910, type of the genus, from southern California. This
extends the range of the genus into the Panamic province,
and material in the Stanford University collection taken
by diving at the Tres Marias Islands and Banderas Bay
in 1965 extend it further. The form proves to be ovovivi- —
parous, the shell small, concentrically ridged, white to
cinnamon brown variously tinged and rayed with pink,
especially near the beak.

23. Circe subtrigona p. 82
= Halodakra subtrigona (CARPENTER, 1857)

Tablet 413, 4 syntypes. [2.8 mm]. Brann: plt. 9, fig. 115.

Haas (1945, Fieldiana, Zoology, vol. 31, no. 2, pp. 4
to 5), discussing a “paratype” at Chicago Museum
pointed out that on account of the sunken ligament this
form is neither a venerid nor petricolid as had been

Explanation of Text figures 1 to 22

Note: Stated lengths are camera lucida readings and may be as

much as 1/10 too high.

Figure 1: Crenella coarctata. Syntype, interior left valve. Length,
4mm (x7).

Figure 2: Gouldia varians. Syntypes, a) exterior; b) interior.
Height, 2.7mm (x 8).

Figure 3: Cardium lucinoides. Holotype, interior of right valve.
Length, 1.4mm (x13).

Figure 4: Circe margarita. Lectotype. a) interior, left valve; b)
interior, right valve; c) enlarged sketch. Length, 1.5mm (x15).

Figure 5: Lucina mazatlanica. Syntypes. a) interior, left valve
(xg); b) interior, right valve (x8); c) exterior, right valve
(x5). Length, 2mm to 4mm.

Figure 6: Lepton dionaeum. Holotype, interior of left valve.
Length, 1.8mm (x12).

Figure 7: Lepton umbonatum. Syntypes, a) sketch of recrystallized
adult left valve (xg); b) and c), interior of juvenile shell (x 8).
Length of adult, 2.6mm; of small pair, 1.8mm.

Figure 8: Montacuta chalcedonica. Holotype. a) enlarged sketch of
hinge, right valve; b) interior, right valve (x17); c) exterior
(x 21). Length, 1.0mm.

Figure 9: Montacuta elliptica. Syntype, right valve interior. Length,
4.0mm (x6).

Figure 10: Montacuta obtusa. Holotype. a) right valve; b) left
valve. Length, 1.7mm (x 1.3).

Figure 11: Montacuta subquadrata. Syntype. Left valve. Length,
3.8mm (x6).

Figure 12: Pythina sublaevis. Syntype. Left valve. a) exterior
(x9); b) interior (x12). Length, 2.2mm.

Figure 13: Cardium alabastrum. Holotype, exterior, left valve.
Length, 3.0mm (x 7).

Figure 14: Cardium rotundatum. Holotype, interior, right valve.
Length, 1.8mm (x 13).

Figure 15: Tapes squamosa. Lectotype, exterior, left valve. Length,
2.8mm (xg).

Figure 16: Rupellaria exarata. Syntype, interior, right valve.
Length, 5mm (x5).

Figure 17: Rupellaria linguafelis. Syntype. a) hinge, right valve;
b) interior, left valve. Length, 3.1 mm (x8); c) exterior, right
valve. Length, 4mm (x6).

Figure 18: Naranio scobina. a) Holotype, left valve. Length,
3.5mm (x5); b) hypotype, right valve. Length, 4mm (x6);
c) enlarged hinges.

Figure 19: Tellina donacilla. Holotype, right valve. Length 4mm
(x7).

Figure 20: Tellina lamellata. Syntype. a) exterior. Length, 3.5 mm
(x6); b) enlarged sketch of hinge.

Figure 21: Tellina regularis. Holotype, right valve. Length 4mm
(x7).

Figure 22: Sphaenia fragilis. Three syntypes, left valves, showing -
resilifer. Length of largest, 8mm (x 3.5).

Vol. 10; No. 4 THE VELIGER Page 395

« Saget

Page 396

thought, and he suggested semelid affinities. OLsson
(1961, p. 319; plt. 27, figs. 1-1c) proposed a generic
name for it, Halodakra, but he was uncertain as to family
placement. Although superficially Bernardina and Halo-
dakra are not strikingly similar, they do have enough in
common so that I have allocated them to the family
BERNARDINIDAE (KEEN, 1963, p. 91).

The species ranges from Baja California to Peru and is
readily recognizable by its chevron-shaped color mark-
ings, well shown in Olsson’s figures; both Carpenter and
Olsson have good figures of the hinge.

LucINIDAE

24. Lucina excavata, p. 98
= Lucina (Here) excavata CARPENTER, 1857
(Plate 56, Figure 23)
Tablet 468, originally 2 syntypes. [10.4mm]. Brann: plt.
12, fig. 140.
Only a single valve remains. It is worn but recognizable
and has been correctly identified by modern workers.

25. Lucina prolongata, p. 100
= Lucina (Cavilinga) prolongata CARPENTER, 1857
(Plate 56, Figures 22 a-d)
Tablet 474, 7 syntypes. [4mm]. Brann: plt. 12, fig. 145.
The species seems to have been correctly identified by
modern workers. Carpenter’s drawing also has been re-
produced by Orsson (1961, plt. 31, fig. 10).

26. Lucina mazatlanica, p. 99

= Lucina (Parvilucina) mazatlanica CarPENTER, 1857

(Plate 56, Figures 29 a-b; Text figures 5 a-c)

Tablet 472, originally 15 syntypes (now 6 pairs, 2 odd
valves). [4mm]. Brann: plt. 12, fig. 144.

The shell is thin, with a lunule longer in most syntypes
than is shown in the figure in my book (Kren, 1958, fig.
193), but the form seems to have been correctly identi-

fied. Orsson (1961, plt. 31, fig. 3) reproduced Carpen-
ter’s drawing.

27. Lucina pectinata, p. 98 (non C. B. Apams, 1847)
= Ctena mexicana (Dat, 1901)
(Plate 56, Figure 21)
Tablet 470, holotype. [12.5 mm].
The single valve is somewhat chipped but in recog-
nizable state.

THE VELIGER

Vol. 10; No. 4

DIPLODONTIDAE

28. Diplodonta ?semiaspera, var. discrepans, p. 103
= Diplodonta discrepans CARPENTER, 1857
(Plate 56, Figures 30 a-d)
Tablet 481, holotype. [7.9 mm]. Brann: plt. 12, fig. 150b.
The single specimen is young but not juvenile. Car-
penter’s drawing was reproduced by Oxsson (1961, plt.
32, fig. 7a), who considers the species distinct and extends
the range south to Colombia. Carpenter was hesitant
about describing this form on account of the variability
he noted in a cluster of diplodontas found within a single
cavity, a mixture of what might otherwise be assigned to
several distinct species. The largest specimen on tablet
479, his typical “D. semiaspera,” is punctate and shows
the wide hinge and sunken ligament of what has recently
been separated out as a new genus and species, Pegma-
pex phoebe Berry, 1960 (figured by Brann on plt. 12,
fig. 150 and by Oxsson from Carpenter’s manuscript
drawings on plt. 32, fig. 3d).

LEPTONACEA

29. Lepton clementinum, p. 110
= ?Mysella clementina (CaRPENTER, 1857)

Tablet 498, holotype. [0.9mm]. Brann: plt. 14, fig. 157.

The holotype is now missing from the tablet, one of
the few that is irrevocably lost. Identification must de-
pend, therefore, on Carpenter’s camera lucida drawing,
which fortunately includes both the exterior and the
interior views.

30. Lepton dionaeum, p. 111
= ?Mysella dionaea (CARPENTER, 1857)
(Text figure 6)
Tablet 499, holotype. [1.5 mm]. Brann: plt. 14, fig. 158.
The holotype consists of one valve, in poor condition;
no trace of pallial line or muscle scars is now visible.

31. Lepton umbonatum, p. 111
= ?Mysella umbonata (CaRPENTER, 1857)
(Text figure 7)

Tablet 500. [2.0mm]. Brann: plt. 14, fig. 159.

Of the two specimens originally on the mount, the
adult is now encrusted with crystals of calcite; a pair of
young, later added (CARPENTER, 1865 b, p. 269), are in
good condition but too immature to be of any help in
interpreting morphology. Thus, Carpenter’s figure is the
only reliable clue.

Vol. 10; No. 4

32. Montacuta chalcedonica, p. 531
?— Kellia chalcedonica (CARPENTER, 1857)
(Text figure 8)

Tablet 2529, holotype. [0.7 mm]. Brann: plt. 10, fig. 694.

The single valve is a nepionic shell, thin and white, the
hinge showing two teeth. Carpenter’s drawing must be
taken as accurate and indicates that already the valve
had been damaged along the lower margin.

33. Montacuta elliptica, p. 113
?—=Lepton ellipticum (CARPENTER, 1857)
(Text figure 9)

Tablet 502, 2 syntypes. [3.3 mm]. Brann: plt. 10, fig. 161.
The shell is so thin that the sculpture, which is of fine
incremental lines, shows through to the inside.

34. Montacuta obtusa CARPENTER, 1865 b, p 270
= Bornia (?) obtusa (CARPENTER, 1857)
(Text figure 10)

Tablet 2530, syntype. [1.5mm]. Brann: plt. 14, fig.
[706].

In 1857 (p. 550) Carpenter cited tablet 2530 as
“Crenella sp. ind.” and mentioned a “hairy epidermis on
the posterior part where it lies in radiating lines, while it
is concentric on the anterior, decussated towards the
middle,” evidently an erroneous reference intended for
his Gregariella coarctata. My notes on the true tablet
no. 2530 are: “Exterior smooth, somewhat iridescent or
lustrous due to the coating of glue; there are faint radial
and concentric striae near the ventral margin.”

35. Montacuta subquadrata, p. 113
see no. 55

36. Pythina sublaevis, p. 112
= Pythinella sublaevis (CARPENTER, 1857)
(Text figure 12)

Tablet 501, 2 syntypes. [2.0mm]. Brann: plt. 14, fig.
160; figure also reproduced by Otsson, 1961, p.238; plt.
83, fig. 12.

One valve of the type material is now badly recrystal-
lized and broken; the drawing is from the hinge of the
other, which is intact. The form has recently been recog-
nized and figured from Panama (Otsson, 1961, plt. 36,
fig. 11).

37. Lasea trigonalis, p. 109
= Orobitella trigonalis (CARPENTER, 1857)
(Plate 56, Figures 28 a-c)
Tablet 496, 2 syntypes, one odd valve. [7.6 mm]. Brann:
pit. 14, fig. 155.
One pair of valves is more trigonal than the other.

THE VELIGER

Page 397

All valves lack the periostracum that probably was pres-
ent. In shape the shell is somewhat similar to Orobitella
(Isorobitella) singularis KEEN, 1962 from San Quintin
Bay, Baja California, but with a less massive hinge. The
pair of valves in Figure 28 a-b (Plate 56) is here selected
as lectotype.

38. Lasea oblonga, p. 109
= Orobitella oblonga (CARPENTER, 1857)
(Plate 56, Figure 24)
Tablet 497, holotype. [5.1 mm]. Brann: plt. 13, fig. 156.
The single valve still shows a remnant of periostra-
cum. The species is close to Sportella stearnsii Dai, 1899
in outline but less elongate.

CHAMIDAE

39. Chama ?frondosa var. fornicata, p. 89
= Chama frondosa mexicana CARPENTER, 1857
(Plate 56, Figures 31 a-b, 35 a-b)
Tablets 439 - 443, 6 syntypes.

Only two of the several syntype specimens are complete
enough and in good enough condition to provide recog-
nizable figures. The variety seems not to be morphologi-
cally valid, for the only constant feature Carpenter cited
was presence of an ashy periostracum.

40. Chama ?frondosa var. mexicana, p. 87

== Chama frondosa mexicana CARPENTER, 1857
Tablets 425 - 438, about 30 syntypes. [140 mm]. Brann:
pit. 10, fig. 121.

Most of the specimens were less than perfect when
collected. There has been no problem in recognition of
the form from Carpenter’s description. Sixteen juvenile
shells are on tablets 425 - 430.

CARDIIDAE

41. Cardium alabastrum, p. 99
= Trigoniocardia granifera (BRODERIP & SOwERBY, 1829)
(Text figure 13)
Tablet 461, holotype. [3.0 mm]. Brann: plt. 11, fig. 133.
The shell is juvenile, with ribs beaded on the anterior
slope and with interspaces between ribs concentrically
striate.
42. Cardium rotundatum, p. 531
= Trachycardium (Mexicardium) panamense (SowERBY,
1833)
(Text figure 14)
Tablet 2522, holotype. [1.6 mm]. Brann: plt. 11, fig. 687.
The valves of this juvenile shell had come loose from

Page 398

THE VELIGER

Vol. 10; No. 4

the mount. Although slightly recrystallized, the shell is
not yet seriously damaged, and surface details can still be
made out.

VENERACEA

43. Trigona humilis, p. 57
= Transennella tantilla humilis (CARPENTER, 1857)
(Plate 55, Figures 17 a-c)
Tablet 244, 6 syntypes. [4.0 mm].

This is the southern form of Transennella tantilla
(Goutp, 1853), from which it differs by being smaller,
longer, smoother, and more brightly colored. The largest
specimen (Plate 55, Figures 17 a-b) here chosen as lecto-
type, shows clearly the oblique grooving of the inner
ventral margin.

44. Dosinia annae, p. 61
= Dosinia annae CarPENTER, 1857
“— Dosinia (Dosinia) semiobliterata DESHAYES, 1853”
teste FISCHER-PIETTE & DELMas, 1967
(Plate 56, Figures 36 a-b)
Tablet 258, 2 syntypes, “A young and a full-grown speci-
men.” [62 mm].
Authors seem to have interpreted the species correctly.
An additional immature shell has subsequently been ad-
ded to the lot; as Carpenter gave dimensions only for

the adult, the uncertainty as to which was added may be
evaded by selecting the full-grown (the largest) speci-
men as lectotype.

In a work received while this paper was in press,
FiscHER-PIETTE & Detmas (Mém. Mus. Nat. d’Hist.
Nat., n.s., sér. A, Zool., vol. 47, fasc. 1, pp. 69 - 70,
1967), revising the genus Dosinia, have shown that the
supposed Australian Dosinia semiobliterata DESHAYES,
1853 actually seems to be D. annae with an erroneous
locality. The subgenus Dosinia does not occur in Austral-
ia but is confined to the tropical American region. Under
Article 23 (b) of the International Code of Zoological
Nomenclature, this would qualify as a nomen oblitum
that should be brought to the attention of the Interna-
tional Commission. FiscHER-PIETTE & DELMas (plt. 11,
figs. 4-6) figure the holotype of the Deshayes species;
it does indeed very closely match my photographs of
the type material of D. annae.

45. Clementia gracillima, p. 54
nomen dubium
Tablet 216. [3.3 mm].

The specimen has recrystallized and is now unrecog-
nizable. Unfortunately, Carpenter did not make a draw-
ing, and his brief description gives little real clue as to
the morphology.

Explanation of Plate 55

All specimens are in the British Museum (Natural History), Mollusca Section. They were photographed in place on glass mounts.

Generic and specific names are as cited by CaRPENTER.

Figure 1: Byssoarca vespertilio. Holotype. a) exterior; b) interior.
Length, 34mm (x1).

Figure 2: Plicatula penicillata. Syntype (smaller of two specimens).
Length, 23mm (x1).

Figure 3: Cycladella papyracea. Holotype. a) interior; b) exterior.
Photograph by Elaine Reeves. Length, 3.5mm (x8).

Figure 4: Mytilus palliopunctatus. Interior of one syntype, a small
adult. Length, 65 mm (xo.7).

Figure 5: Mytilus multiformis. Two paralectotypes. a) length
1omm (x3); b) length 12mm (x3).

Figure 6: Ostrea palmula. Lectotype (here selected) ; interior of
right valve. Length, 60 mm (xo0.7).

Figure 7: Leiosolenus spatiosus. Syntype, adult. a) exterior;
b) interior. Length, 38mm (x1).

Figure 8: Lithophagus aristatus tumidior. Syntype, interior. a) left
valve; b) right valve. Length 4omm (x1).

Figure 9: Lithophagus aristatus gracilior. Interior of one of 7
broken syntype valves. Length (incomplete), 15mm (x3).

Figure 10: Isognomon janus. Syntype. Maximum diagonal length,
47mm (x1).

Figure 11: Naranio scobina. a) Holotype, left valve. Length, 3.5

mm; b) Hypotype, right valve. Length, 4.5mm (x 3.8).

Figure 12: Cyrena olivacea. Two syntypes. a), b), interior, left
valve and right valve. Length, 35mm; c) exterior, left valve.
Length, 50mm (x0.6).

Figure 13: Tapes squamosa. Syntype (here selected as lectotype) ,
left valve. Length, 2.9mm. a) (x4); b) (x7).

Figure 14: Placunanomia pernoides. Lectotype (here selected).
a, b) right and left valves, interior. Length, 52mm; c) Para-
lectotype, interior, left valve. Length, 52 mm (xo.6).

Figure 15: Placunanomia claviculata. Syntype, maximum length
27mm (x1.1).

Figure 16: Lithophagus calyculatus. Interior of holotype. Length
8.4mm (x5).

Figure 17: Trigona humilis. a, b) lectotype, interior of paired
valves. Length, 4mm; c) lectoparatype. Length, 3.2 mm (x 4.2).

Figure 18: Arca bifrons. Two syntypes, Cuming collection. a) here
chosen as lectotype, exterior. Length, 43mm; b, c) interior.
Length, 33 mm (xo.6).

Figure 19: Venus cortezi. Holotype, from Guaymas, Mexico.
Length, 61 mm (x0.6).

Figure 20: Modiola mutabilis. Two syntypes, sub-adult. a) interior,
left valve. Length, 44mm; b) exterior, right valve. Length 42
mm (x1).

Tue VELIcER, Vol. 10, No. 4 [KEEN] Plate 55

Vol. 10; No. 4

46. Tapes squamosa, p. 78
= Timoclea (Glycydonta) squamosa (CARPENTER,
1857)
(Plate 55, Figures 13 a-b; Text figure 15)

Tablet 372, 3 valves. [2.3 mm]. Brann: plt. 9, fig. 111.
Otsson (1961, p. 309) recognizes this small chionid
as ranging from southern California to Peru. After restu-
dying Carpenter’s type and comparing my figures with
material very like it in the Stanford collection, from the
La Paz area, I am obliged to revise my previous sugges-
tion (KEEN, 1966, Occ. Pap. Calif. Acad. Sci. no. 59, pp.
9-10) and to admit that Venus troglodytes Morcu,
1861 is a synonym of Tapes squamosa. However, I feel
that there are two forms on the West Coast, Timoclea
picta (WiLLeTT, 1944), ranging north from Magdalena
Bay and averaging more than 6mm in length, with
somewhat more subdued ribbing, and Timoclea squamosa,
ranging from the Gulf of California southward to Peru,
averaging less than 6 mm in length, with stronger scales
on the concentric ribs. Otsson has objected to the use
of the generic name Timoclea because the type species
has a nearly smooth escutcheon; perhaps we should there-
fore adopt the subgeneric allocation to Glycydonta Cor-
TON, 1936 ( based on Venus marica LINNAEUS, 1758),
the type of which has strong scales on the posterior slope.
Carpenter’s largest syntype is here selected as lectotype.

PETRICOLIDAE

47. Rupellaria exarata, p. 20
=Petricola (Petricola) exarata (CARPENTER, 1857)
(Text figure 16)

Tablet 73, 4 syntypes. [5.3 mm]. Brann: plt. 3, fig. 28.

Radial ribbing is somewhat developed, and the fine
zigzag pattern is not as apparent as in the next species.
There are a few brown color spots. One specimen was
nestling in a barnacle test. Coan in 1962 (The Veliger,
vol. 5, no. 2, p. 92) reported some specimens of this
form taken near Mazatlan in 1961.

48. Rupellaria linguafelis, p. 20
= Petricola (Petricola) linguafelis (CARPENTER, 1857)
(Text figure 17)
Tablet 72, 5 syntypes. [4.0 mm]. Brann: plt. 2, fig. 27.
The specimens are juvenile, with microscopically fine
zigzag striae. With growth the shell tends to become more
elongate.
49. Naranio scobina, p. 529
= Petricola (Petricola) linguafelis (CARPENTER, 1857)
(Plate 55, Figures 11 a-b; Text figures 18 a-b)

Tablet 2516, holotype and hypotype. [3.3mm]. Brann:
pit. 4, fig. 680.

THE VELIGER

Page 399

When he described the species, Carpenter had but a
single valve, the left valve. Later he found a larger
opposite (right) valve, which he added to the mount in
1858 (Carpenter, 1865b, p. 269). His drawing, which
therefore must have been made after 1858, shows both
specimens, the hinge of the left (holotype), and the full
interior of the right (hypotype). There seems no good
basis for separation of the form as distinct.

TELLINIDAE

50. Tellina donacilla, p. 34
?== Tellina (Moerella) donacilla CARPENTER, 1857
(Text figure 19)

Tablet 116, holotype, 1 valve. [3.5 mm]. Brann: plt. 6,
fig. 53.

The shell looks like a young Tellina amianta DAL,
1900, but the posterior end is shorter and the lateral tooth
stronger ; no trace of the pallial line could be made out.

DOoNACIDAE

51. Donax ?punctatostriatus caelatus, p. 46
= Donax punctatostriatus HANLEY, 1843
(Plate 56, Figure 34)
Tablet 168, 3 syntypes; 168*, two syntypes. [35 mm].
This form does not seem separable from Hanley’s
species.

52. Donax culminatus, p. 43
= Donax carinatus Haney, 1843
(Plate 56, Figures 33 a-b)
Tablet 139, 2 syntypes, the smaller, from Mazatlan, here —
selected as lectotype. [24 mm].

Carpenter considered this a variant of Donax carinatus
because the ridge setting off the posterior slope seemed
sharper and the radial striae seemed more crowded and
granular; later authors have not considered these as valid
distinguishing characters.

GaRIDAE

53. Solecurtus politus, p. 27
= Tagelus (Mesopleura) politus (CARPENTER, 1857)
(Plate 56, Figure 27)
Tablet 90, 1 syntype. [37 mm].
Authors seem to have identified the species correctly.
The right valve is now broken; Carpenter implied that
the shell had been complete.

54. Solecurtus violascens, p. 27, footnote
= Tagelus (Tagelus) violascens (CARPENTER, 1857
(Plate 56, Figure 37)

Page 400

THE VELIGER

Vol. 10; No. 4

Tablet 90, holotype. [83 mm]. Brann: plt. 17, lower
right.

The original specimen was from the Cuming collec-
tion with an indefinite locality, “Southwest Mexico.” All
that remains now is a somewhat broken right valve.
Fortunately, growth lines are well developed and enable
the reconstruction of the correct outline. The pallial sinus
reaches only just slightly past the intersection of a line
drawn vertically through the beaks; Carpenter’s drawing
shows it not quite reaching this line. The pallial sinus in
Tagelus californianus (Conrab, 1837) is still shorter and
the shell proportionately longer. The interior of the right
valve of Carpenter’s specimen was figured by PALMER
(1963, plt. 64, fig.1), registry no. 1857.6.4.2531.

SEMELIDAE

55. Montacuta subquadrata, p. 113
= Semelina subquadrata (CARPENTER, 1857)

(Text figure 11)

Tablet 503, 2 syntypes, one broken. [3.5mm]. Brann:
pit. 10, fig. 162.

The shell is worn, with a chipped edge; the sculpture
is strongly concentric; interior markings (pallial line and
muscle scars) not being visible, Carpenter was unaware
of the deep pallial sinus that is evident in fresh material.
Otsson (1961, p. 375; plt. 66, fig. 11) has figured the
exterior sculpture and has discussed morphology and
distribution.

56a. Tellina lamellata, p. 37
= Semele sp., juvenile
(Text figure 20)

Tablet 121, 3 valves. [3.3 mm].

The figure in KEEN (1958, fig. 388) is fairly correct as
to outline except that Carpenter’s syntypes are longer
anteriorly. The hinge shows the internal ligament that is
characteristic of Semele. The shell is probably too young
for a positive determination of what Semele is repre-
sented. Studies of growth series might give clues.

96 b. Tellina regularis, p. 36
= Semele sp., juvenile
(Text figure 21)

Tablet 120, 2 syntypes, one a fragment. [1.8mm]. BRANN:
plt. 7, fig. 57.

This resembles the “7? /amellata in hinge and outline
but has only fine concentric lines. My drawing indicates
a larger size than Carpenter’s cited dimensions.

57. Sphaenia fragilis, p. 24, “n.s.”
=Sphenia fragilis (H. « A. ADaAMs, 1854)
(Text figure 22)

Tablets 80 - 82, about 15 syntypes. [8 mm]. Brann: plt.
6, fig. 35.

The several syntypes show variations in size and out-
line. Authors have correctly identified the form. Unfor-
tunately, Carpenter chose the same specific name as had
H. and A. Apams (Genera of Shells, vol. 2, p. 368; plt.

Explanation of Text figures 23 to 50

Note: Stated lengths are camera lucida readings and may be as

much as 1/10 too high.

Figure 23: Dentalium liratum. Lectotype. Length, 7mm (x8).

Figure 24: Dentalium corrugatum. Holotype. Length, 1.5mm
(x19).

Figure 25: Rimula mazatlanica. Syntype. Length, 4mm (x 8).

Figure 26: Scissurella rimuloides. Holotype. Diameter, 0.9 mm
(x22). a) apertural view; b)and c) apical and lateral views,
sketches, not to scale.

Figure 27: Liotia carinata. Holotype. Diameter, 1.6mm (x19).

Figure 28: Liotia striulata. Holotype. Diameter, 1.1mm (x21).

Figure 29: “Phasianella perforata Puuipri.” Paralectotype of
Tricolia mazatlanica (STRONG, 1928). Diameter, 2.6mm (x9).

Figure 30: Phasianella perforata striulata. Syntype. Diameter,
1.8mm (x15).

Figure 31: Lunatia tenuilirata. Syntype. Diameter, 1.6mm (x 14).

Figure 32: Leiostraca linearis. Holotype. Length, 2.2mm (x14).

Figure 33: Leiostraca producta. Holotype. Length, 3.8mm (x 10).

Figure 34: Leiostraca iota retexta. Syntype. Length, 2.6mm (x 12).

Figure 35: Leiostraca distorta yod. Syntype. Length, 2.6mm (x 12).

Figure 36: Mucronalia involuta. Holotype. Length, 3.2 mm (x 10).

Figure 37: Aclis tumens. Holotype. Length, 1.6mm (x 19).

Figure 38: Scalaria raricostata. Holotype. Length, 3.5 mm (x 10).

Figure 39: Scalaria suprastriata. Syntype. Length, 12mm (x 2.8).

Figure 40: Cirsotrema funiculata. Syntype. Length, 14.5 mm
(x 2.6).

Figure 41: Vitrinella orbis. Holotype. Diameter, 1 mm (x 23).

Figure 42: Viitrinella bifrontia. Syntype. a) apertural view; b)
base. Diameter, 1 mm (x 23).

Figure 43: Vitrinella coronata. Syntype. Diameter, 1mm (x 23).

Figure 44: Vitrinella lirulata. Holotype. Diameter, 2mm (x 16).

Figure 45: Vitrinella perparva nodosa. Holotype. Diameter, 1 mm
(x 20).

Figure 46: Vitrinella ornata. Holotype. a) apertural view; b) base.
Diameter, 1mm (x 23).

Figure 47: Vitrinella tenuisculpta. Holotype. Diameter, 1 mm
(x 23).

Figure 48: Vanicoro cryptophila. Lectotype. Diameter, 2mm (x 15).

Figure 49: Vitrinela planospirata. Holotype (fragment). Radius
3.2mm (x11).

Figure 50: Vitrinella cincta. Holotype. Diameter, 1 mm (x 20).

THE VELIGER Page 401

Vol. 10; No. 4

Page 402

THE VELIGER

Vol. 10; No. 4

97, figs. 3, 3a) for their Tyleria fragilis. Carpenter had
been skeptical of their figure and was at pains to study
their type specimen, as he had seen Sphenias with some-
what similar features. He concluded (p. 527) that the
form was correctly illustrated and was unique. I concur
that the figure is accurate, but I feel certain that their
specimen is pathologic. For some reason, shell material
was either not deposited or was resorbed, leaving a few
islands of solid calcium carbonate embedded in what is
mostly periostracum. In shape, size, and hinge, the shell
is strikingly similar to Carpenter’s syntypes, as I could
prove by comparing the two lots side by side. If my
interpretation is correct, the generic name Tjleria falls as
a synonym of Sphenia, and the specific name has priority
from the Adamses, 1854.

CorBULIDAE

58. Corbula pustulosa, p. 22
= Corbula nasuta SoweErsBy, 1833

(Plate 56, Figure 25)
Tablet 77, 1 syntype (here selected as lectotype). [4 mm].
Brann: plt. 4, fig. 32.

The lectotype shows clearly the radial rows of small
pustules that are the hallmark of juvenile Corbula nasu-
ta. Carpenter’s other syntype was from the Cuming
collection, from “Panama and San Blas.”

PHOLADIDAE

59. Martesia intercalata, p. 13
= Penitella conradi VALENCIENNES, 1846

Tablet 41 (sketches).

The type material consisted of two specimens in the
Hanley collection, of which Carpenter made drawings.
The shells are no longer available. Ruth Turner has re-
produced the drawings (Johnsonia, vol. 3, no. 34, p. 75,
plt. 72, figs. 1-2, 1955).

SCAPHOPODA

DENTALIDAE

60. Dentalium liratum, p. 188
= Dentalium semipolitum Broperte & SOWERBY, 1829

(Text figure 23)

Tablet 879. [6 mm]. BRANN: plt. 19, fig. 244.

Carpenter cited a “perfect specimen” (here selected as
lectotype), a small specimen, and a fragment. There are
two juvenile shells on the tablet, one of which is probably
a caecid. Pitspry & SHarP (Manual of Conchology, vol.
17, p. 92, 1897) suggested the synonymy here accepted.

Explanation of Plate 56

All specimens are in the British Museum (Natural History), Mollusca Section. They were photographed in place on glass mounts.

Generic and specific names are as cited by CarPENTER.

Figure 21: Lucina pectinata. Holotype, left valve. Length, 13 mm
(x3).

Figure 22: Lucina prolongata. Four unmatched syntype valves.
a, d) interior, left valve; b) interior, right valve; c) exterior, left
valve. Length, 3mm to 3.5mm (x3).

Figure 23: Lucina excavata. Syntype, exterior, seen through glass
mount (the central circular area is glue). Length 10 mm (x 3.5).

Figure 24: Lasea oblonga. Holotype, left valve, interior. Length,
5-7mm (x 4.5).

Figure 25: Corbula pustulosa. Exterior, left valve, lectotype (here
selected). Length, 4.2 mm (x 4.5).

Figure 26: Diplodonta subquadrata. Syntype. a) interior, left
valve; b) exterior, right valve. Length 26.3 mm (x 1.3).

Figure 27: Solecurtus politus. Syntype, left valve. a) exterior;
b) interior. Length, 32mm (x1).

Figure 28: Lasea trigonalis. a) interior, left valve; b) interior, right
valve, lectotype (here selected). Length, 7.5mm (x3). c) ex-
terior of paralectotype, seen through glass mount (central shiny
area is glue). Length, 7.6mm (x 4.6).

Figure 29: Lucina mazatlanica. Two syntypes. a) interior, right

valve. Length, 3.2mm; b) interior, left valve. Length 2mm
(x10).

Figure 30: Diplodonta discrepans. Holotype. a) exterior, right
valve; b) interior, right valve; c) exterior, left valve; d) interior,
left valve. Length, 8mm (x3).

Figures 31, 35: Chama frondosa fornicata. Syntypes, sub-adults.
31a) interior, left valve; 31 b) exterior, right valve; maximum
length, 78mm (x0.6); 35a) interior, left valve; 35b) interior,
right valve; maximum length, 45mm (x1).

Figure 32: Thracia squamosa. Holotype, Cuming collection. a) in-
terior, right valve;b) exterior, left valve. Length, 28mm (x1.1).

Figure 33: Donax culminatus. Lectotype (here selected), exterior,
right valve. Length, 25 mm (x0.8). b) Hypotype, interior, left
valve of a pair from an unknown Central American locality.
Length, 37mm (x0.8).

Figure 34: Donax caelatus. Exterior, left valve of a syntype, largest
of three specimens. Length, 37 mm (x 1.1).

Figure 36: Dosinia annae. Lectotype (here selected). a) interior,
right valve; b) interior, left valve. Length, 60mm (xo0.5).

Figure 37: Solecurtus violascens. Holotype, with Carpenter’s label,
from “Southwest Mexico.” Length, 83mm (x 0.9).

THE VELIGER, Vol. 10, No. 4 [KEEN] Plate 56

itd

SOE Mes
1 lay, Cte fase £9,

Si)

Vol. 10; No. 4

61. Dentalium corrugatum CarPENTER, 1857, p. 189
(non Hupé in Gay, 1854)
= Dentalium semipolitum Broperip & SowERBY, 1829
(Text figure 24)

Tablet 881, holotype. [1.2 mm]. Brann: plt. 19, fig. 246.

The shell is probably too young for positive deter-
mination, but as the name is preoccupied, synonymizing
seems the best course.

62. Dentalium ?pretiosum “Nutt. (teste Hinds) ,” p. 189
nomen nudum

Tablet 882.

The citation is based on a specimen too worn and
broken for determination. Had Carpenter validated the
name by giving a description, it would have taken pre-
cedence over Sowerby’s usage of 1860.

GASTROPODA

FISSURELLIDAE

63. Fissurella alba, p. 218 (non Puiuprt, 1845)
= Fissurella gemmata MENKE, 1847
(Plate 57, Figures 38 a-b)

Tablets 1058 - 1068, 29 syntypes (tablet 1059 photo-
graphed). [41 mm].

Carpenter had some misgivings about the identity of
MENKE's species, but later authors have generally accep-
ted the synonymy. Menke’s type material, however, has
not been studied.

64. Fissurella nigrocincta CARPENTER, 1856a, p. 234
(cited 1857b, p. 217, as “n:s.”)
= Fissurella nigrocincta CARPENTER, 1856
(Plate 57, Figures 39 a-b)
Tablet 1056, 2 juvenile syntypes. [19 mm]. Brann: plt.
19, fig. 274.

The species was described from adult material in the
Cuming collection, locality not stated, shown in my photo-
‘graph. Carpenter cited the locality in 1857 as Mazatlan,
but the label says “California.” The figure in BRANN is
of a juvenile shell on tablet 1056, which Carpenter re-
ferred to this species with some doubt. Sowerby published
an illustration of the adult in 1862 (Thesaurus Conch.,
vol. 3, plt. 239, figs. 61 - 63).

65. Fissurella spongiosa, p. 219
=Fissurella nigrocincta CARPENTER, 1856
(Plate 57, Figures 40 a-b)

Tablet 1070, syntype, here selected as lectotype. [18 mm].

THE VELIGER

Page 403

The shell is immature and worn, which accounts for
the texture.

66. Rimula mazatlanica, p. 222
== Diodora sp., juvenile
(Text figure 25)
Tablet 1080, 3 syntypes. [3.1 mm]. BRANwN: plt. 22, fig.
281.

The species has also been figured by Sowerby in Reeve
(Conch. Icon., vol. 19, plt. 1, sp. 5, 1873). The callus on
the inside of the apex has the truncation of Diodora,
and there are a few color spots on the shell; also, the
sculpture near the margin is like that of Diodora. As
Berry (1964, Leaflets in Malacology, no. 24, p. 148) has
surmised, this form represents the growth stage in which
the orifice has not yet absorbed the pointed apex.

ScISSURELLIDAE

67. Scissurella rimuloides CARPENTER, 1865 b, p. 271
= Sinezona rimuloides (CARPENTER, 1865)
(Text figures 26 a - c)
Tablet 2532, holotype. [0.7 mm]. Brann: plt. 22, fig.
698.

The species seems to have been correctly identified by
authors.

TROCHIDAE

68. Trochus macandreae, p. 232
—Calliostoma macandreae (CARPENTER, 1857)
(Plate 57, Figure 41)

Tablet 1129, 1 syntype and some fragments. [8.7 mm].

The photographed specimen is here selected as lecto-
type. A specimen from the Cuming collection was figured
by Reeve, 1863 (Conch. Icon., vol. 14, plt. 7, figs. 50-51),
as Ziziphinus macandreae.

69. Omphalius globulus, p. 236
= Tegula (Agathistoma) globulus (CarPENTER, 1857)
(Plate 57, Figures 52 a-b)
Tablet 1145, 1 syntype, here selected as lectotype. [8.4
mm].
There have been some differences of opinion among
collectors as to the identity of this form.
70. Omphalius ?rugosus var. rufotinctus, p. 233
— Tegula (Omphalius) rugosa (A. Apams, 1853)
(Plate 57, Figures 45 a-b)
Tablets 1130 - 1131, 3 syntypes. [30 mm]
The 3 syntypes of Chlorostoma rugosum A. ApaMs,
1853 are also in the type collection of the British Muse-
um, with the locality “China.” Because of this incorrect

Page 404

and indefinite label, Carpenter described the Mazatlan
form as a new variety. I regard the two lots as conspecific
and consider Adam’s locality to be in error.

LIoTODAE

71. Liotia carinata, p. 248
= Arene carinata (CARPENTER, 1857)

(Text figure 27)

Tablet 1164, holotype. [1.1 mm]. Brann: plt. 25, fig.
313

As Carpenter noted, the shell is immature, but the
sculpture is well developed. The spire is so depressed that
it does not show in the apertural view of the shell.
Strone (1934, Trans. San. Diego Soc. Nat. Hist., vol. 7,
no. 37, p. 440; plt. 28, figs. 1 - 3) was probably not justi-
fied in using a figure of the holotype of Liotia lurida
Dat, 1913 under the name of L. carinata. His figure
of another specimen (ibid., plt. 31, figs. 1-3) is of a
shell with a tabulate but not flattened spire. Dall’s species
should be reinstated as distinct. Carpenter’s form has not
been recorded again unequivocally, but some specimens
in the Stanford University collection from the La Paz
area approach his figure in flatness of spire.

72. Liotia striulata, p. 248
= Arene striulata (CARPENTER, 1857)
(Text figure 28)
Tablet 1165, holotype. [0.7mm] Brann: plt. 26, fig.
314.
Carpenter’s shell has an unusually thick apertural mar-
gin. The spire is higher than in Arene carinata. His
drawing is somewhat tilted, and he exaggerated the spi-

THE VELIGER

Vol. 10; No. 4

ral sculpture. Part of the lip thickening seems to be a
build-up of the mounting glue. One specimen in the
Stanford University collection, dredged off La Paz,
matches the figure fairly well as to size and shape, but
it has a thinner apertural margin and does not have the
sculpture so markedly developed.

73. Liotia c-b-adamsii, p. 248

Tablet 1166, holotype. [0.6 mm]. Brann: plt. 26, fig. 315.

The holotype is now completely disintegrated by chem-
ical alteration. Carpenter’s drawing suggests the apical
whorls of a juvenile shell that evidently is not liotiid.
For all practical purposes it must be regarded as a
species dubia. In a way this may be well, because under
the new International Rules of Zoological Nomenclature,
the name must be written without hyphens, which results
in an unpronounceable vocable, cbadamsii.

PHASIANELLIDAE

74. “Phasianella perforata Puuiprt,” p. 224 (not of
Puiuipri, 1845)

= Tricolia mazatlanica (Stronc, 1928), “new name”
(Plate 57, Figure 46; Text figure 29)

Tablet 1084, 5 syntypes. [3.3mm]. Brann: pl. 22, fig.
283.

The type of Philippi’s species was from Peru. StRoNG
(1928, p. 198) reprinted Carpenter’s description and
cited a figure in REEvE (Conch. Icon., vol. 13; plt. 6,
fig. 17, 1862), of a specimen in the Cuming collection
from Mazatlan and Panama. A specimen with an oper-
culum, mentioned by Carpenter, is now the only good
specimen of the Reigen lot, and it is here selected as
lectotype of Strong's species.

Explanation of Text figures 51 to 74

Note: Stated lengths are camera lucida readings and may be as

much as 1/10 too high.

Figure 51: Vitrinella decussata. Syntype. Diameter, 1.3 mm (x 16).

Figure 52: Vitrinella monile. Syntype. Diameter, 1.6mm (x17

Figure 53: Vitrinella monilifera. Syntype. Diameter, 1.2mm (x 20

Figure 54: Vitrinella annulata. Holotype. Diameter, 1.6mm (x 17).

Figure 55: Ethalia carinata. Syntype. Diameter, 1mm (x 19).

Figure 56: Vitrinella carinulata. Holotype. Diameter, 1 mm (x 16).

Figure 57: Ethalia pyricallosa. Holotype. Diameter, 1.4mm; a)
apertural view (x 23); b) base (x 18).

Figure 58: Teinostoma amplectans. Holotype. Diameter, 3.3 mm
(x9).

Figure 59: Globulus amplectans. Syntype. Diameter, 1mm; a)
apertural view (x 23); b) base (x 27).

Figure 60: Globulus lirulata. Syntype. a) apertural view; b) basal
view. Diameter, 1mm (x 23).

Figure 61: Globulus pallidula. Holotype. Diameter 1mm (x 25).

.
Ve

Figure 62: Teinostoma substriatum. Syntype. Diameter, 1.2mm;
a) apertural view, (x19); b) basal view (x20).

Figure 63: Globulus sulcatus. Syntype. Diameter, 1mm (x 28).

Figure 64: Globulus tumens. Syntype. a) apertural view; b) basal
view. Diameter, 1.6mm (x15).

Figure 65: Vitrinella bifilata. Syntype. Diameter, 1.5mm (x 19).

Figure 66: Vitrinella naticoides. Syntype. Diameter, 1.2mm (x 20).

Figure 67: Vitrinella subquadrata. Syntype. Diameter, 1.2mm
(x 19).

Figure 68: Alvania tumida. Syntype. Diameter, 1.6mm (x 10).

Figure 69: Aclis fusiformis. Syntype. Length, 1.8mm; a) aper-
tural view; b) detail of initial apical whorl from back of shell
(x 18).

Figure 70: Jeffreysia bifasciata. Syntype. Length, 1.6mm (x17).

Figure 71: Jeffreysia tumens. Syntype. Length, 1.6mm (x17).

Figure 72: Jeffreysia alderi. Syntype. Length, 2mm (x 18).

Figure 73: Rissoina woodwardit. Syntype. Length, 3.6mm (x8). ~

Figure 74: Rissoa lirata. Syntype. Length, 4mm (x8).

Page 405

@ @) OG eas
Gy FO On° TCS
FE hO®D VG
G8 OHO

57b

THE VELIGER

Vol. 10; No. 4

Page 406

THE VELIGER

Vol. 10; No. 4

75. Phasianella ?perforata, var. striulata, p. 225
= Tricolia striulata (CARPENTER, 1857)
(Text figure 30)
Tablet 1085, 1 syntype, registry no. 1857.6.4.1085. [2.2
mm]. BRANN: plt. 22, fig. 283b.

Strone (1928, p. 198), in proposing the name Phast-
anella mazatlanica as a replacement for Carpenter’s mis-
identification, seems to have overlooked Carpenter’s
varietal name. If, as seems possible, this form is not
separable from the one on tablet 1084 (the only cited
difference being less apparent sculpture), then Strong’s
T. mazatlanica will fall into synonymy, having been junior
to T. striulata less than 50 years required for the latter
to become a nomen oblitum.

76. Phasianella compta “Goutp MSS,” p. 225
?Phasianella, s. l., sp.
Tablet 1086. Brann: plt. 22, fig. 284.

It is probable that Carpenter’s Mazatlan specimen is
misidentified, for Tricolia compta (Goutp, 1855) is a
Californian form. A note with the specimen, by Dr. R.
Robertson, 1962, suggests that this is a ballast shell from
the Indo-Pacific or Caribbean and is unidentifiable.

77. Lunatia tenuilirata, p. 451
= Tricolia tenuilirata (CARPENTER, 1857)
(Text figure 31)
Tablet 2052, syntype. [1.4 mm]. Brann: plt. 50, fig. 572.

Carpenter mentions two specimens, a broken older and
a perfect young shell and says the mount contains the
latter. However, the one on the tablet has an incomplete
aperture and the dimensions are closer to those he cites for
the larger specimen. This form seems very close to Tri-
colia substriata (CARPENTER, 1864), from the Californi-
an area, and only minor differences can be seen between
a photograph of a Californian shell (Strone, 1928, p.
195; plt. 10, fig. 11) and the camera lucida drawings by
Carpenter and myself. Possibly the record of T: substriata
at San José Island, Gulf of California, cited by Strone,
should be considered as of T. tenuilirata instead. The
main differences seem to be that the Mazatlan syntype
has stronger developed color banding of white spots (not
shown in Carpenter’s drawing) and a heavier inner lip
on the aperture.

PHENACOLEPADIDAE

78. Scutellina navicelloides, p. 211
= Phenacolepas osculans (C. B. ApAMs, 1852)
(Plate 57, Figures 44 a-b)

Tablet 1016, holotype. [5.6mm]. Brann: plt. 21, fig.
269.

The apex of the holotype is broken away. Outline,
however, and sculpture seem to confirm synonymy with
Adams’ species.

Explanation of Plate 57

All specimens are in the British Museum (Natural History), Mollusca Section. They were photographed in place on glass mounts.

Generic and specific names are as cited by CARPENTER.

Figure 38: Fissurella alba. a) interior; b) exterior; two syntypes.
Length of larger, 36mm (x1).

Figure 39: Fissurella nigrocincta. Two of 3 syntypes, Cuming col-
lection. a) exterior; b) interior. Length of largest specimen in
lot, 18mm (x1.5).

Figure 40: Fissurella spongiosa. Lectotype (here selected). a) ex-
terior, seen through glass; b) interior. Length, 17 mm; a)(x 2.2);
b) (x 1.1).

Figure 41: Trochus macandreae. Lectotype (here selected). Dia-
meter, 9mm (x4).

Figure 42: Hipponyx planatus. Two syntypes, Cuming collection,
from Panama. a) interior, b) exterior of specimen on which
from Panama. a) interior; b) exterior of specimen on which
Carpenter cited dimensions. Length, 18mm (x 1.3).

Figure 43: Hipponyx serratus. Two syntypes. a) exterior; b) in-
terior. Diameter (larger syntype). 23mm (x1).

Figure 44: Scutellina navicelloides. Holotype. a) interior (x 3.3) ;
b) exterior, seen through glass mount (central area that appears
flattened is the mounting glue) (x5.2). Length, 5.5 mm.

Figure 45: Omphalius rufotinctus. Two syntypes. a) diameter,
18mm; b) diameter, 24mm (x1).

Figure 46: “Phasianella perforata Puitipri.” Lectotype (here se-
lected) of P. mazatlanica Stronc, 1928. Height, 2.5mm (x8).

Figure 47: Cerithidea mazatlanica. Four syntypes. Length of
largest, 28mm (x1).

Figure 48: Cerithium alboliratum. Lectotype (here selected) .
Height, 4.0mm (x7).

Figure 49: Scalaria raricostata. Holotype. Length, 3.3mm (x13).

Figure 50: Cirsotrema funiculata. a) Lectotype (here selected),
largest of 3 specimens from Panama, Cuming collection. Length,
21mm (x1.9). b) Hypolectotype, from Mazatlan. Length, 16
mm (x 2.4).

Figure 51: Cerithium mediolaeve. Holotype: Length, 25.3 mm
(x 1.7).

Figure 52: Omphalius globulus. Lectotype (here selected). a)
basal view; b) apertural view. Diameter, 8.5mm (x4).

Figure 53: Galerus fuscus. Holotype, from “Gulf of California.”
Diameter, 15.5mm (x 1.1).

Figure 54: Litorina philippi. Syntype. Length, 11mm (x3).

Figure 55: Trochita ventricosa. Holotype. a) side view; b) basal
view. Diameter, 19 mm (x1).

Figure 56: Crepidula bilobata. Exterior of a syntype. Diameter
gmm (x4).

THE VELIGER, Vol. 10, No. 4 [KEEN] Plate 57

Vol. 10; No. 4

THE VELIGER

Page 407

EULIMIDAE

79. Leiostraca linearis, p. 440
— Balcis linearis (CARPENTER, 1857)
(Text figure 32)
Tablet 2025, holotype. [1.8mm]. Brann: plt. 48, fig.
554.

Bartscu (1917, p. 310; plt. 36, fig. 4) did not repro-
duce the original figure, as he did with other of Carpen-
ter’s Mazatlan forms, but figured instead a specimen
from off Baja California. His identification seems to be
correct.

80. Leiostraca producta CARPENTER, 1864
= Balcis producta (CARPENTER, 1864)

(Text figure 33)

Tablet 2022, holotype. [3.1 mm]. BRANN: plt. 48, fig.
Solr

Although he did not publish a formal description until
1865, Carpenter introduced the name earlier with an
acceptable indication (1864 a, p. 357), in connection
with discussions of the C. B. Adams types, when he de-
cided that his identification of Mazatlan material with
a Panama form had been unjustified. Carpenter's draw-
ing seems distorted. Having made a separate one of my
own from the type, I cannot reconcile the two unless he
rotated the shell on account of the broken outer lip and
attempted a restoration. The body whorl seems wider
than in any other Panamic Balcis in proportion to the
height of the spire.

81. Levostraca Piota var. retexta, p. 440
= Balcis retexta (CARPENTER, 1857)

(Text figure 34)

Tablet 2026, 1 syntype. [2.2mm]. Brann: plt. 48, fig.
50:

BartscuH (1917, p. 317; plt. 38, fig. 1) reprinted the
original discussion of Carpenter and published the cam-
era lucida drawing now available in Brann’s plates. My
own drawing of the type differs in minor details but in
general confirms Carpenter’s accuracy.

82. Leiostraca ?distorta, var. yod, p. 441
= Balcis yod (CarpENTER, 1857)
(Text figure 35)

Tablet 2027, 4 syntypes. [2.0mm]. Brann: plt. 48, fig.
556.

Again, Bartscu (1917, p. 330; plt. 40, fig. 9) pub-
lished a Carpenter drawing, not having recognized any
specimens of this form. My own drawing of the largest
syntype differs only in the rendering of the inner lip.

83. Mucronalia involuta CARPENTER, 1865b, p. 273
= ?Eulima involuta (CARPENTER, 1865)
(Text figure 36)
Tablet 2021, holotype. [2.7 mm]. Brann: plt. 48, fig.
550.

In his monograph on melanellid mollusks, BArTscH
(1917, p. 297) stated that as this form has a tilted apex,
it is an Odostomia. However, he had not cited it as such
in the pyramidellid monograph (Dati & Bartscu, 1909),
at which time it was stated that he had studied Carpen-
ter’s types at the British Museum. Thus, this species was
omitted in both of the accounts and had not been figured
until the publication of Carpenter’s plates. My observa-
tions confirm Carpenter’s statement that there is no co-
lumellar fold. I could not see the apical whorls as hetero-
strophic. The holotype is very worn and is brown stained.
Its general shape, the form of the outer lip, the lack of
a columellar fold, and the apparently simple apex seem
to point to eulimid rather than pyramidellid affinities.

ACLIDIDAE

84. Aclis tumens, p. 438
= ?Aclis tumens CARPENTER, 1857
(Text figure 37)

Tablet 2017, holotype. [1.3mm]. Brann: plt. 48, fig.
546.

Placement of the genus Aclis and the family AcLipipaE
is unsettled. The shells resemble pyramidellids in many
ways but lack a columellar fold; the apex is normal. They
do not have the shiny texture of the Eutimiparg, and the
whorls tend to be more inflated. My notes on Aclis tumens
indicate that the apex is minute, sunken but apparently
dextral, the back of the shell seeming to show faint ©
spiral sculpture but the front smooth. The holotype has a
large break.

EPIroNUDAE

85. Scalaria raricostata, p. 447 [non Woop, 1828]
—=Epitonium (?Nitidiscala) carpentert (TAPPARONE-
Canerri, 1876)
(Plate 57, Figure 49; Text figure 38)

Tablet 2040, holotype. [3.1mm]. Brann: plt. 50, fig.
568.

The holotype has a slightly worn look and is greyish in
color. Axial sculpture does not appear until the fourth
apical whorl. The allocation to Epitonium (Punctiscala)
in Keen, 1958 (p. 276), following Stronc in BurcH,
(Min. Conch. Club S. Calif. no. 52, p. 20, 1945) was

Page 408

ill-advised, for there is no evidence of spiral punctations
between the low varices, of which there are 7 per whorl.

Brann (1966, p. 17) cites Carpenter’s specific name
as available because the name given by LAMaARcK in 1822
that had been considered by some authors to preoccupy
Carpenter’s was spelled S. raricosta. However, Woop in
1828 used the spelling raricostata for what is evidently
another species in the genus.

86. Scalaria suprastriata, p. 446
=Epitonium (Nitidiscala) suprastriatum (CARPEN-
TER, 1857)

(Plate 59, Figure 101; Text figure 39)

Tablet 2037, 1 syntype. [11 mm].

Although the edges of the varices on the type (which
is stored in the British Museum’s Type Collection) are
somewhat chipped, the shell is otherwise in good condi-
tion. The dimensions seem close to those of Epitonium
roberti Dax, 1917.

87. Scalaria (Cirsotrema) funiculata, p. 447
= Opbalia diadema funiculata (CARPENTER, 1857)

(Plate 57, Figure 50; Text figure 40)

Tablet 2041, 1 syntype, with broken aperture. [14.5 mm].
Three syntypes from Panama, Cuming collection.

In the annotated copy of his Catalogue, left at the
British Museum, Carpenter had correctly reassigned this
species to Opalia. The largest specimen of the 3 from
Panama is more slender and high-spired than the other
syntypes. Carpenter was not convinced of a distinction

THE VELIGER

Vol. 10; No. 4

of the species from the Ecuadorean O. diadema (SoweEr-
BY, 1832). Because the largest Panama specimen is the
only one differing enough in outline possibly to justify
separation of the northern form, it is here selected as
lectotype (Plate 57, Figure 50a).

The ribs in this form seem more distinct than in the
Californian Opalia crenimarginata (Dat, 1917), which
it resembles.

JANTHINIDAE

88. Ianthina striulata, p. 185
= Janthina janthina (LinnatEvus, 1758)

(Plate 59, Figure 79)
Tablets 868 - 876, 37 adults (tablet 872 photographed).
[20 mm].

There seems no basis for separating Pacific and At-
lantic forms of this pelagic species. REEvE (1858, Conch.
Icon., vol. 11, plt. 2, sp. 6) cited the Carpenter name
under synonymy of Janthina fragilis LaMarck, 1801.

89. Ianthina striulata, var. contorta, p. 186
= Janthina janthina (Linnaeus, 1758)
(Plate 59, Figure 80)

Tablet 877, 2 syntypes (only one now on the tablet).
[Size not given. ].

Synonymy with the Atlantic form seems justifiable.
REEvE (1858, Conch. Icon., vol. 11, plt. 4, sp. 19) figured
a specimen from Mazatlan said to be in the Cuming
collection.

Explanation of Text figures 75 to 104

Note: Stated lengths are camera lucida readings and may be as

much as 1/10 too high.

Figure 75: Caecum clathratum. Syntype. Length, 2.9mm (x14).

Figure 76: Caecum compactum. Syntype. Length, 2.8mm (x15).

Figure 77: Caecum elongatum. Syntype. Length, 3mm (x 10).

Figure 78: Caecum farcimen. Syntypes. Length, a) 1.7mm (x 17);
b) 1.6mm (x 16).

Figure 79: Caecum quadratum. Syntypes. Length, a), b) 2mm
(x 16).

Figure 80: Caecum semilaeve. Syntype. Length, 3.2mm (x11).

Figure 81: Caecum subimpressum. Syntype. Length, 3.4 mm (x 10).

Figure 82: Caecum undatum. Syntype. Length, 2mm (x17).

Figure 83: Caecum heptagonum. Holotype, incomplete, side and
sectional views. Diameter, 0.8mm (x17).

Figure 84: Caecum liratocinctum. Syntypes. Length, a) 4mm
(x13); b) 3mm (x14).

Figure 85: Caecum subconicum. Syntypes. Length, a), b) 2.4mm
(x 10).

Figure 86: Caecum subobsoletum. Syntype. Length, 3mm (x 14).

Figure 87: Caecum tenuiliratum. Holotype. Length, 2.8mm (x 14).

Figure 88: Caecum abnormale. Syntype. Length, 1.6mm (x17).

Figure 89: Caecum insculptum. Syntypes. a) Length, 1.5 mm
(x10); b) Length, 4mm (x10).
Figure 90: Caecum obtusum. Syntype. Length, 3.9mm (x12).
Figure 91: Caecum subspirale. Syntype. Length, 4.6mm (x Io).
Figure 92: Caecum dextroversum. Syntypes. a) Length, 2.4mm
(x14); b) Length, 2mm (x15).
Figure 93: Caecum reversum. Holotype. Length, 1.6mm (x17).
Figure 94: Caecum teres. Syntype. Length, 3.2mm (x 14).
Figure 95: Caecum corrugulatum. Holotype. Length, 2.2 mm (x 17).
Figure 96: Bivonia albida. Syntype. Length of coil, 1omm (x7).
Figure 97: Cerithium alboliratum. Lectotype. Length, 4mm (x 7).
Figure 98: Cerithiopsis albonodosa. Syntype. Length, 1.7mm
(x 14).
Figure 99: Cerithiopsis cerea. Holotype. Length, 3mm (x12).
Figure 100: Cerithiopsis pupiformis. Syntypes. a) juvenile; b)
broken adult. Length of each, 2.2mm (x14).
Figure 101: Cerithiopsis sorex. Syntype. Length, 2mm (x 14).
Figure 102: Cerithiopsis tuberculoides. Syntypes. a) Length, 2.4mm
(x13); b) Length, 4.6mm (x8).
Figure 103: Cerithiopsis convexa. Holotype. Length, 5.6mm (x 7).
Figure 104: Cerithiopsis decussata. Holotype. Length, 5.2mm

(x7).

THE VELIGER Page 409

Vol. 10; No. 4

Page 410

THE VELIGER

Vol. 10; No. 4

90. Ianthina decollata, “nom. prov.” p. 187
= Janthina prolongata BLAINVILLE, 1822
(Plate 59, Figures 78 a-c)
Tablet 878, 3 syntypes. [26 mm].
Carpenter had justifiable misgivings about the distinct-
ness of this form.

LITTORINIDAE

91. Litorina philippiu, p. 349
= Littorina dubiosa philippu CARPENTER, 1857
(Plate 57, Figure 54)

Tablets 1671 - 1685, 109 syntypes (tablet 1676 photo-
graphed. [10 mm].

The numerous variants were arranged by Carpenter on
15 slides, of which one was photographed that he de-
scribed as “zigzag at base.”

VITRINELLIDAE

92. Vitrinella orbis, p. 247
=Cyclostremella orbis (CARPENTER, 1857)

(Text figure 41)

Tablet 1163, 1 syntype. [0.8mm]. Brann: plt. 25, fig.
S122,

The shell is nearly smooth but shows one thin carina
at the periphery and a fainter one above and below.
Pitspry & Otsson (1952, p. 84) suggest the plausible
allocation to C'yclostremella.

93. Vitrinella bifrontia, p. 242
= Cyclostremiscus bifrontia (CARPENTER, 1857)
(Text figure 42)
Tablet 1154, 1 syntype. [1.3mm]. Brann: plt. 24, fig.
303.
The allocation by Pirspry & Otsson (1952, p. 62), on
the basis of another syntype, seems justified.

94. Vitrinella coronata, p. 244
= Cyclostremiscus coronatus (CARPENTER, 1857)

(Text figure 43)
Tablet 1157, 2 syntypes, the larger figured. [1.4 mm].
Brann: plt. 24, fig. 306.
The allocation by Pirspry & Oxtsson (1952, p. 62)
seems correct.

95. Vitrinella lirulata, p. 241
= C'yclostremiscus lirulatus (CARPENTER, 1857)
(Text figure 44)
Tablet 1151, holotype. [2.1 mm]. Brann: plt. 23, fig. 300.
The species was allocated to Cyclostremiscus by Pits-
BRY & OLsson (1952, p. 83). When studying the holotype
I noted that it was nearest in form to their Cyclostremis-
cus nummus from Panama but lacked any peripheral
carination.
96. Vitrinella perparva, var. nodosa, p. 243
= C'yclostremiscus nodosus (CARPENTER, 1857)
(Text figure 45)

Tablet 1155, holotype. [0.8mm]. Brann: plt. 24, fig. 304.

Explanation of Plate 58

All specimens are in the British Museum (Natural History), Mollusca Section. They were photographed in place on glass mounts.

Generic and specific names are as cited by CaRPENTER.
Figure 57: Oliva intertincta. Syntype, back view. Length, 20mm
(x2).
Figure 58: Litiopa divisa. Syntype, from “Cape San Francisco.”
Length, 2.7mm (x10).
Figure 59: Nassa nodulifera. Holotype. Length, 16mm (x 1.9).
Figure 60: Nassa crebristriata. Holotype. Length, 13.5 mm (x 2.7).
Figure 61: Nassa acuta. Syntype. Length, 20 mm (x 1.6).
Figure 62: Mangelia subangulata. Holotype. Length, 4.3mm
(x 6.5).
Figure 63: Murex lividus. Two syntypes. a) Length, 44mm; b)
Length, 40mm (x1).
Figure 64: Muricidea indentata. Holotype. Length, 34 mm (x 1.7).
Figure 65: Pisania elata. Two hypotypes, without definite locality,
in British Museum collection as “P. elata, var.” a) Length, 34mm;
b) Length, 37 mm (x1).
Figure 77: Drillia cerithoidea

Figure 66: Erato panamensis. Syntype, from Panama. Length,
75mm (x 4.4).

Figure 67: Melampus olivaceus. Syntype. Length, 15mm (x3).

Figure 68: Siphonaria aequilirata. Holotype. Length, 17mm (x 2.4).

Figure 69: Siphonaria palmata. Two syntypes. a) Length, 21 mm;
b) Length, 29mm (x1.3).

Figure 70: Terebra albocincta. Longest of 5 syntypes on tablet
1828. Length, 42mm (x1.5).

Figure 71: Yerebra hindsii. Two syntypes. a) Apertural view.
Length, 31.5 mm; b) Back view. (x1.8).

Figure 72: Terebra subnodosa. Syntype. Length, 33 mm (x 1.8).

Figure 73: Terebra rufocinerea. Syntype. Length,.31 mm (x 1.8).

Figure 74: Clathurella aurea. Holotype. Length, 14.5 mm (x 3.5).

Figure 75: Drillia punctatostriata. Holotype from Panama. Length,
19mm (x 3.5).

Figure 76: Drillia monilifera. Holotype. Length, 15.4 mm (x 4.2).

. Syntype. Length, 16mm (x 4.2).

[KEEN] Plate 58

Tue VELIGER, Vol. 10, No. 4

Vol. 10; No. 4

The allocation by Pirspry & Oxsson (1952, p. 63)
seems justified, but their figure of a supposed paratype
may be of a specimen that is not conspecific, for the
peripheral nodes are relatively larger than in the holotype,
as shown both by my drawing and that of Carpenter.

97. Vitrinella ornata CARPENTER, 1864 b, p. 271
= C'yclostremiscus ornatus (CARPENTER, 1864)
(Text figure 46)
Tablet 2533, holotype. [0.9mm]. BrRANN: plt. 25, fig.
2535.

Although they did not have a figure for guidance,
Pitspry & Orsson (1952, p. 82) seem to have allocated
this correctly. Carpenter evidently labeled his figure
2535” by error. He described no. 2535 as only a frag-
ment of a shell, whereas the figure given is of a complete
specimen and the drawing matches well the specimen
mounted on tablet no. 2533.

98. Vitrinella tenuisculpta CarPENTER, 1864b, p. 271
= Cyclostremiscus tenuisculptus (CARPENTER, 1864)

(Text figure 47)

Tablet 2534, holotype [0.7mm]. Brann: plt. 25, fig.
Zao"

Pirspry & Oxrsson (1952, p. 83) seem to have allo-
cated the form correctly. Carpenter’s drawing somewhat
exaggerates the dorsal curve and hence the outline of
the shell. My own drawing suggests a more lenticular
form.

99. Vitrinella trigonata, p. 244
=Cyclostremiscus trigonatus (CARPENTER, 1857)

Tablet 1156, hypotype. [0.6mm]. Brann: plt. 24, fig.
305, as Vitrinella exigua C. B. Apams, 1852.

Carpenter’s specific name could be rejected as having
been proposed in synonymy, for it was a name he had
used in manuscript and merely mentioned in his discus-
sion of Vitrinella exigua. Pirspry & Otsson (1945, pp.
268 - 269), however, consider that Adams’ species is a
secondary homonym of Delphinula exigua Pui.iprt, 1845,
which also seems to be a Cyclostremiscus. Thus, they
reject the name exigua and substitute trigonata of Car-
penter. Although names proposed in synonymy are to be
rejected under Article 11 of the International Code of
Zoological Nomenclature, this Article has been modified
in the second edition of the Code to provide that if such
a name has, prior to 1961, been treated as available, it
may be accepted, with its original date and authorship.
Thus, V. trigonata may be utilized. The holotype, of
course, remains that of Adams, not the figured specimen
on tablet 1156.

THE VELIGER

Page 411

100. Vanicoro cryptophila, p. 262
= Macromphalina cryptophila (CARPENTER, 1857)
(Text figure 48)

Tablets 1313 - 1315. [4 mm]. Brann: plt. 28, fig. 330.

Carpenter’s drawings show well the tilted apex (which
is brown in color) and the sculpture of Macromphalina.
I had the impression when observing some of the juvenile
specimens under the microscope that one or two might be
Vanikoro. 1 am therefore designating the adult in tablet
1314, shown also in my drawing, as the lectotype. The
adult in tablet 1315, which will thus become a lectopara-
type, is in place in a crevice of a Spondylus fragment; it
shows fine spiral sculpture in the body whorl.

101. Vitrinella planospirata, p. 246
= ?Miralabrum planospiratum (CARPENTER, 1857)
(Text figure 49)

Tablet 1162, holotype. [ca. 5mm]. Brann: plt. 24, fig.
elite

The shell is represented by a fragment only, and Car-
penter’s attempt at a restoration is highly fanciful. The
sculpture and size suggest Miralabrum Pitspry & OLS-
son (1945, p. 276), the type of which, also incomplete,
is from Ecuador (M. unicornis). A specimen of Maira-
labrum has been found by Faye Howard at Cape San
Lucas that may represent Carpenter’s species; a photo-
graph of this shell is given here for comparison (Plate
59, Figure 102).°’The size (diameter 4.7 mm) compares
well with Carpenter’s estimate of probable size of his
holotype.

102. Vitrinella cincta, p. 245
=Lydiphnis (Cymatopteryx) cincta (CARPENTER,
1857)
(Text figure 50)
Tablet 1159, holotype. [0.8mm]. BRANN: plt. 25, fig. 308.
Pirspry & Orsson (1952, p. 83) suggest that this
species should be allocated to Cyclostremiscus, but study-
ing the holotype and comparing it with their figures I
noted that it was closest to their figure of Lydiphnis
strongi. Therefore, I revise their assignment.

103. Vitrinella decussata, p. 239
== Parviturboides decussatus (CARPENTER, 1857)
(Text figure 51)
Tablet 1148, 6 syntypes. [1.1 mm]. BRANN: plt. 23, fig.
297.
Pirspry & Otsson (1952, p. 68; plt. 11, figs. 3, 3a-b)
have correctly reallocated this species; their figure is of

a “paratype” [7. e., syntype].

Page 412

104. Vitrinella monile, p. 240
— Parviturboides monile (CARPENTER, 1857)
(Text figure 52)
Tablet 1149, 4 syntypes. [1.4mm]. Brann: plt. 23, fig.
298.

Carpenter’s drawing as published by Brann confirms the
allocation to Parviturboides made by Pitspry & OLSSON
(1952, p. 84). My own drawing may not be entirely
accurate as to outline, due to my faulty adjustment of
the camera lucida device, and his is definitely tilted.

105. Vitrinella monilifera, p. 240
— ?Parviturboides monilifer (CARPENTER, 1857)

(Text figure 53)

Tablet 1150, 1 syntype. [1 mm]. Brann: plt. 23, fig. 299.
The allocation to Parviturboides has been suggested by
Pitspry & Oxsson (1952, p. 84). My notation when
studying the syntype was that it might be a juvenile Sol-
ariorbis. The lenticular outline would be more in harmony
with such an allocation, but the sculpture and evenly
curved inner lip are reminiscent of Parviturboides.

106. Vitrinella annulata, p. 245
= Solariorbis (Systellomphalus) annulatus (CARPEN-
TER, 1857)
(Text figure 54)

Tablet 1158, holotype. [1.3mm]. Brann: plt. 25, fig.
307.

It is doubtful if Pmspry « Orsson (1952, p. 83) had
an adequate figure of Carpenter’s material, else they
could not have suggested an assignment to Cyclostremis-
cus for this smooth-spired form. It closely resembles
the Ecuadorean Pliocene species that is the type of
Systellomphalus but differs in the arrangement of the
basal cords. Prrssry & Oxsson described Systellompha-

THE VELIGER

Vol. 10; No. 4

lus as a genus in 1941 but by 1952 (p. 51) demoted it to
a species-group rank. However, it seems to me to have
sufficient distinctness to be usefully retained as a sub-
genus under Solariorbis.

107. Globulus (Ethalia) carinata, p. 252
= ?Solariorbis carinatus (CARPENTER, 1857)
(Text figure 55)
Tablet 1172, 1 syntype. [0.9mm]. Brann: plt. 28, fig.
321.

With some hesitation I accept the allocation to Solari-
orbis of Pitspry & Otsson (1952, p. 83). When I was
examining Carpenter’s specimen and comparing it with
the Pilsbry and Olsson figures, I concluded that it best
matched that of Cyclostremiscus tricarinatus (C.B.
ApaMs, 1852) but that it had a weaker upper keel, a
lower spire, and a shallower umbilicus.

108. Vitrinella carinulata, p. 246
. =Solariorbis carinulatus (CARPENTER, 1857)
(Text figure 56)
Tablet 1160, holotype. [0.8mm]. Brann: plt. 25, fig.
309.

The figure by Pirspry & Oxsson (1952, plt. 13, figs.
4a, 4b) is a good representation of the form, but the
specimen (now at Harvard) cannot be a paratype, for
Carpenter cited only a single example. My drawing is a
little distorted obliquely, due to my faulty adjustment of
the camera lucida attachment.

109. Globulus (Ethalia) pyricallosa, p. 251
—Solariorbis fyricallosus (CARPENTER, 1857)
(Text figure 57)
Tablet 1169, holotype. [1.2mm]. Brann: plt. 26, fig.
318.

Explanation of Text figures 105 to 128

Note: Stated lengths are camera lucida readings and may be as
much as 1/10 too high.
Figure 105: Alaba laguncula. Holotype. Sketch (about x 22).
Figure 106: Alaba mutans. Holotype. Length, 2.6mm (x11).
Figure 107: Alaba scalata. Holotype. Length, 1 mm (x 23).
Figure 108: Alaba terebralis. Holotype. Length, 1.8mm (x 19).
Figure 109: Alaba violacea. Holotype. Length, 1.8mm (x17).
Figure 110: Alvania effusa. Holotype. Length, 3.6mm (x 8).
Figure 111: Alvania excurvata. Syntype. Length, 3.6mm (x8).
Figure 112: Hipponyx planatus. Syntypes. a) Diameter, 1.8mm
(x8); b) Diameter, 5mm (x 4.5).
Figure 113: Fossarus tuberosus. Syntype. Length, 1.5mm (x 23).
Figure 114: Crepidula bilobata. Syntype. Diameter, 4mm (x 8).
Figure 115: Anachis gaskoini. After Carpenter’s sketch of holo-
type. Length, 6mm (x6).
Figure 116: Anachis serrata. Syntype. Length, 39mm (x49).
Figure 117: Fusus apertus. Syntype. Length, 4.6mm (x7).

Figure 118: Fusus tumens. Syntype. Length, 5mm (x7).

Figure 119: Marginella margaritula. Lectotype. a) Length, 2.2 mm
(x11); b) and c), paralectotypes, apertural and back views.
Length, 2.1mm (x11).

Figure 120: Marginella polita. a) Paralectotype. Length, 0.9 mm
(x 25); b) lectotype. Length, 1.1 mm (x 26); c) detail of ante-
rior part of aperture, tilted forward.

Figure 121: Drillia cerithoidea. Syntype. Length, 15mm (x 2.3).

Figure 122: Mangelia sulcata. Holotype. Length, 5.6mm (x 7).

Figure 123: Mangclia subangulata. Holotype. Length, 4.3 mm (x 7).

Figure 124: Clathurella aurea. Holotype. Length, 15mm (x 2.5)-

Figure 125: Haminea cymbiformis. Holotype. Length, 2.7 mm
(x 13).

Figure 126: Bulla exarata. Syntype. Length, 3.8mm (x 10).

Figure 127: Smaragdinella thecaphora. Holotype. Length, 2.4mm _
(x 15).

Figure 128: Tornatina carinata. Syntype. Length, 3.2 mm (x 13).

Vol. 10; No. 4 THE VELIGER Page 413

Page 414

THE VELIGER

Vol. 10; No. 4

The allocation to Solariorbis by Pitssry & OLssoN,
(1952, p. 82) seems justifiable.

110. Teinostoma amplectans, p. 253
= Teinostoma (Pseudorotella) amplecians CARPEN-
TER, 1857
(Text figure 58)

Tablet 1174, holotype. [3mm]. Brann: plt. 26, fig. 323
(also figured by SowerBy in Reeve, 1874, Conch. Icon.,
vol. 19, plt. 1, figs. la-c, from a specimen in the British
Museum).

The figure of Teinostoma americanum Pitssry & OLS-
son, 1945 (plt. 23, fig. 3), a southern variant of this
species, is a good representation of the form except for
the slightly smaller size. In 1952 (p. 41) they allocated
both forms to the subgenus Pseudorotella.

111. Globulus (Ethalia) amplectans, p. 253
—=?Teinostoma amplectans CARPENTER, 1857

(Text figure 59)

Tablet 1173, 1 syntype. [1 mm]. Brann: plt. 26 fig. 322.

Pirspry & Orsson (1952, p. 82) allocated this to Sol-
ariorbis, probably not having seen a good figure. Both
Carpenter’s drawing as published by BRANN and my own
suggest the strong umbilical callus of Teinostoma. Car-
penter himself was not sure but that this might be the

young of T: amplectans — hence his choice of the same
specific name for both. The shell is translucent and rela-
tively thick; my notes suggest that it may be a young of
T. lirulatum.

112. Globulus (Ethalia) lirulata, p. 251
= Teinostoma lirulatum (CARPENTER, 1857)
(Text figure 60)

Tablet 1170, 1 syntype. [1 mm]. Brann: plt. 26, fig. 319.

As with the previous form, Pirssry & Otsson (1952,
p. 82) allocated this to Solariorbis. The callus seems to
me to be too well developed for that. I noted when
examining the holotype that the closest figure in the Prts-
BRY & OLSSON monographs was their Teznostoma ecua-
dorianum (1952, plt. 3, fig. 5), but T: lirulatum has, as
the specific name suggests, fine spiral sculpture.

113. Globulus (Ethalia) pallidula, p. 252
= ?Teinostoma pallidulum (CARPENTER, 1857)
(Text figure 61)

Tablet 1171, holotype. [1.1 mm]. Brann: plt. 26, fig.
320.

The large aperture and relatively small diameter of
this shell are peculiar. Prrspry & Otsson (1952, p. 82)
considered it to be a Solariorbis, but it seems to me that
the outline is closer to that of a Teinostoma, such as T.

Explanation of Plate 59

All specimens are in the British Museum (Natural History), Mollusca Section. They were photographed in place on glass mounts.

Generic and specific names are as cited by CARPENTER.

Figure 78: Janthina decollata. Three syntypes. Diameter of largest,
25mm (x1.1).

Figure 79: Janthina striulata. Syntype. Diameter, 17mm (x 1.6).

Figure 80: Janthina contorta. Syntype. Diameter, 21mm (x 1.1).

Figure 81: Sistrum rufonotatum. Three syntypes, from Cape San
Lucas, Baja California. Length of largest, 121mm (x 2).

Figure 82: Anachis albonodosa. Syntype. From a color photograph
by Elaine Reeves. Length, 3mm (x 10).

Figure 83: Anachis pachyderma. Two syntypes. Length of larger,
16mm (x 2.4).

Figure 84: Columbella cervinetta. Holotype. Length, 7.8 mm
(x 5.5).

Figure 85: Columbella obsolcta. Syntype. Length, 6mm (x5,.5).

Figure 86: Anachis gaskoini. Hypotype, from Peru. Length, 6.5
mm (x 5.5).

Figure 87: Anachis serrata. Two syntypes. a) Apertural view;
Length, 3.1mm; b) Back view (specimen here sclected as lecto-
type) ; Length, 3.5mm. From a color photograph by Elaine
Reeves. (x8).

Figure 88: Olivella aureocincta. Two syntypes. a) Apertural view;
b) Back view. Length, 8.3mm (x 3.5).

Figure 89: Olivella glandinaria. Two syntypes, from California,
Nuttall collection. Length of larger, 25 mm (x 1.1).

Figure 90: Pisania acquilirata. Holotype. Length, 25.5 mm (x 1.6).

Figure 91: Lophyrus striatosquamosus. Holotype. Length, 4.5 mm
(x 8).

Figure 92: Acanthochites arragonites. Syntype (here selected as
lectotype). Length, 3.0mm (x11).

Figure 93: Lepidopleurus macandrcac. Holotype. Length, 3.3 mm
(x11).

Figure 94: Lepidopleurus beanit. Syntype (here selected as Iecto-
type). Length, 6mm (x7).
Figure 95: Lepidopleurus bullatus. Syntype (here selected as lecto-
type). Length, 4.5 min (x8).
Figure 96: Lepidopleurus calciferus. Holotype. Length, 3.3 mm
(x 10).

Figure 97: Lepidopleurus clathratus. Holotype. Length, 4.4 mm
(x 8).

Figure 98: Drillia albovallosa. Holotype. Length, 1omm (x 4.2).

Figure 99: Drillia hanlcyi. Holotype. Length, 10mm (x 4).

Figure 100: Drillia albonodosa. Lectotype (here selected). Length,
13mm (x 3.6).

Figure 101: Scalaria suprastriata. Syntype. Length, 11.5 mm (x 4).

Figure 102: Miralabrum sp., cf. ?M. planospirata (CARPENTER).
Hypotype from Cape San Lucas, Baja California, collected by
Faye Howard. Diameter, 4.7 mm (x8).

Figure 103: Chiton flavescens. Syntype. Length about 6mm (x 10).

Figure 104: Tonicia forbcsit. Syntype (here selected as lectotype).
Length, 19mm (x 2.4).

[KEEN] Plate 59

THE VELIGER, Vol. 10, No. 4

Vol. 10; No. 4

THE VELIGER

Page 415

rarum PitsBry & Oxsson, 1945 (plt. 23, fig. 2). There is
no umbilical chink, and the shell has a fine punctate
spiral sculpture. The small size suggests immaturity.

114. Teinostoma substriatum, p. 254
= Teinostoma substriatum CARPENTER, 1857

(Text figure 62)

Tablet 1175, syntype. [1.1 mm]. Brann: plt. 26, fig. 324
(figured also by Sowerby in Reeve, 1874 (Conch. Icon.,
vol. 19, plt. 1, fig. 4, from a specimen in the British Mu-
seum).

Pitssry & Oxtsson (1952, p. 43, plt. 11, fig. 2) have
illustrated a “paratype” (actually a syntype) that, al-
though an imperfect specimen, seems to be conspecific
with the specimen in the British Museum. The umbilical
callus is less well developed than in other species, but
there is no chink at its margin.

115. Globulus sulcatus, p. 250
= Teinostoma sulcatum (CarPENTER, 1857)
(Text figure 63 )

Tablet 1168-b, 1 syntype. [0.7 mm]. Brann: plt. 26, fig.
317-b.

Carpenter published this name provisionally and did
not cite a separate tablet number, indicating that he had
mounted one of two specimens on the card with the type
of his Globulus tumens. The British Museum specimen
now becomes secondary, for Pirssry « Orsson (1952, p.
44; plt. 3, fig. 3) selected the other of Carpenter’s two
specimens as lectotype (citing it, however, as “‘holotype”).
It was at that time in the collection of the New York
State Museum but has since been transferred to the Mu-
seum of Comparative Zoology at Harvard University on
permanent loan. Their figure accords well with my sketch
of the British Museum specimen. Whether the furrow on
the callus has any morphological significance remains to
be seen.

116. ?Globulus tumens, p. 250
== Teinostoma tumens (CARPENTER, 1857)

(Text figure 64 a-b)
Tablet 1168, 1 syntype. [ — ]. BRANN: plt. 26, fig. 317.
Pirssry & Orsson (1952, p. 43) correctly allocated
this to Teinostoma.

117. Vitrinella bifilata, p. 241
= Vitrinella bifilata CARPENTER, 1857

(Text figure 65)

Tablet 1153, 2 syntypes. [1.4mm]. Brann: plt. 24, fig.
302.

This has been figured from a “paratype” [2. e. syntype]
in the New York State Museum collection (now on per-
manent loan to Harvard University) by Pitspry « OLs-
Sonn (1952 pn /osiplt. ll, figs l))

118. Vitrinella naticoides, p. 246
= ?Vitrinella naticoides CARPENTER, 1857
(Text figure 66)
Tablet 1161, 1 syntype. [1.2 mm]. Brann: plt. 25, fig.
310.

Pitspry & Orsson (1952, p. 73; plt. 11, fig. 4) illus-
trated a “paratype” that was in the New York State
Museum collection ( now at Harvard University on per-
manent loan). It is similar in form to the syntype at the
British Museum. On the basis of the spiral cord that
makes the inner lip of the aperture of uneven width, I
should have supposed this would be a Solariorbis. They,
however, retain it in Vitrinella with a query.

119. Vitrinella subquadrata, p. 241
= Vitrinella subquadrata CARPENTER, 1857
(Text figure 67)

Tablet 1152, 4 syntypes. [1 mm]. Brann: plt. 23, fig. 301.

Pirspry & Oisson (1952, p. 76; plt. 11, fig. 5) have
figured a “paratype” [i.e.syntype] that was then in the
New York State Museum( now at Harvard University
on permanent loan). The figure accords well with that of
the best syntype drawn by Carpenter. My own drawing
is a freehand sketch intended only to show general fea-
tures.

120. Vitrinella clathrata, p. 238
Invalid name
This name was given in the synonymy of Vitrinella
parva C. B. Apams, 1852. It was a manuscript name that
Carpenter abandoned when he studied Adams’ Panamic
material, and it falls as a name proposed in synonymy.

RISSOIDAE

121. Alvania tumida, p. 360
= Alvania (Alvinia) tumida CarPENTER, 1857
(Text figure 68)

Tablet 1711, 1 syntype. [1.4mm]. Brann: plt. 39, fig.
esa

BartscH (1911d, p. 361; plt. 32, fig. 2) has given a
figure of a specimen from the U.S. National Museum
collection that seems to be correctly identified.

122. Alaba mutans, p. 369
= ?Alvania (Lapsigyrus) mutans (CARPENTER, 1857)
(Text figure 106)

Tablet 1729, holotype. [2.2 mm]. Brann: plt. 40, fig. 431.

The back of the body whorl is broken away in the
holotype, which explains Carpenter’s choice of side-view
for the drawing. The spiral ribs are not so strong as he
would make them, and there is a channeled suture, as
shown in my drawing, though he does not indicate it. The

Page 416

THE VELIGER

Vol. 10; No. 4

apex is slightly sunken. I had hoped that this would
provide a name that could be salvaged for the “Alvania
livata” of authors, but direct comparison of specimens
ruled this out. No nodes develop above the suture, the
aperture is more oblique, and the spire is more tapering.
The shell is closer to the type species of Lapsigyrus BERRY,
1958 (Leaflets in Malacology, vol. 1, no. 16, p. 92), a
Pleistocene form from Magdalena Bay — Alvania cont-
rerast JORDAN, 1936 (Contributions, Department of
Geology, Stanford University, vol. 1, no. 4, p. 160; plt.
19, fig. 9). As compared to Jordan’s figure, Carpenter's
shell is smaller, the aperture is shorter, and the spiral
sculpture is finer. If these differences prove to be consist-
ent for the Recent form (the mention by Berry of fine
sculpture in his report of specimens from the Sonoran
coast suggests such a possibility), Carpenter’s specific
name may indeed have utility.

123. Aclis fusiformis, p. 437
= ?Onoba fusiformis (CARPENTER, 1857)

(Text figure 69)

Tablet 2016, 2 syntypes. [1.6mm]. BRANN: plt. 48, fig.
545.

At first glance one would take this for a pyramidellid,
for the apex is sunken and folded over at the back as if
heterostrophic. Bartsch evidently rejected it from Pyrami-
dellidae, however, when he examined the types at the
British Museum. Although the columella is twisted, it
does not really bear a fold. The sculpture is pitted, in the
manner of Acteon, but it lacks other characters of that

genus. The outline seems nearest to that of the rissoid
genus Onoba, and there is a first record of that group in
the West American fauna as O. fortis Pitspry « OLSsoN,
1941 (Proc. Acad. Nat. Sci. Philadelphia, vol. 93, p. 45;
plt. 8, fig. 3), from the Pliocene of Ecuador. Several spe-
cimens from beach drift taken at Salinas, Ecuador, match
their figure well, so that the species may be regarded
as still living there. These specimens, in the Stanford Uni-
versity collection, have the apical whorls in good condi-
tion; the apex is acute, not sunken as in Carpenter’s
Mazatlan specimens. I therefore make the allocation
with considerable doubt.

RISSOELLIDAE

124. Jeffreysia bifasciata, p. 362
= Rissoella bifasciata (CARPENTER, 1857)
(Text figure 70)

Tablet 1716, 3 syntypes. [1.4 mm]. Brann: plt. 39, fig.
419.

The color bands show up well, especially on the back
of the largest syntype and on the two smaller shells in
apertural view. BartscH (1920, p. 162; plt. 12, fig. 2)
has figured a specimen he calls “type” (probably a syn-
type).

I have collected live specimens intertidally in gravelly
sand at Cape San Lucas. RoBertson (1962, Notulae
Naturae, Acad. Nat. Sci. Philadelphia, no. 352, p. 1) has
shown that the generic name Rissoclla M. E. Gray,

Explanation of Text figures 129 to 158

Note: Stated lengths are camera lucida readings and may be as
much as 1/10 too high.
Figure 129: Odosiomia lamellata. Syntype. Length (incomplete) ,
2mm (x14).
Figure 130: Odostomia subsulcata. Syntype. Length, 1.4 mm (x 19).
Figure 131: Odostomia vallata. Syntype. Length, 1.6mm (x17).
Figure 132: Odostomia convexa. Syntype. Length, 2.3 mm (
Figure 133: Odostomia effusa. Holotype. Length, 2.6mm (x12).
Figure 134: Odostomia fasciata. Syntype. Length, 2.5mm (
Figure 135: Odostomia nodosa. Syntype. Length, 4.9mm (
Figure 136: Odostomia oblonga. Syntype. Length, 4.9mm (x 7).
Figure 137: Odostomia ovata. Syntype. Length, 4.9mm (x7).
Figure 138: Odostomia ovulum. Syntype. Length, 1.7mm (x 16).
Figure 139: Odostomia reigeni. Holotype. Length, 1.7mm (x15).
Figure 140: Odostomia rotundata. Syntype. Length, 2.6 mm (x 12).
Figure 141: Odostomia telescopium. Syntype. Length, 4mm (x9).
Figure 142: Odostomia lacunata. Syntype. Length, 1.4 mm (x 19).
Figure 143: Odostomia tenuis. Syntype. Length, 1.8mm (x 19).
Figure 144: Chemnitzia intermedia. Syntype. Length, 1.6mm
(x 19).
Figure 145: Odostomia photis. Syntype. Length, 1.4 mm (x 20).

Figure 146: Odostomia quinquccincta. Syntype. Length, 1.8mm
(x 19).

Figure 147:Odostomia clausiliformis. Syntype. Length, 3.5 mm
(x10).

Figure 148: Odostomia sublirulata. Holotype. Length, 2.3 mm
(x14).

Figure 149: Odostomia ziziphina. Holotype. Length, 1 mm (x 23).

Figure 150: Odostomia armata. Syntype. Length, 2.5 mm (tilted)
(x11).

Figure 151: Odostomia exarata. Hypotype. Length, 2.9mm (x11).

Figure 152: Odostomia mammillata. Holotype. Length, 1.4mm
(x 18).

Figure 153: Odostomia scalariformis. Syntype. Length, 3 mm
(x11).

Figure 154: Fossarus maculosa. Syntype. Length, 4mm (x8).

Figure 155: Chemnitzia muricata. Syntype. Length, 2.8inm
(x14).

Figure 156: Chemnitzia subangulata. Syntype. Length, 2.6mm
(x 14).

Figure 157: Chemnitzia cancellata. Holotype. Length, 1.2 mm
(x 26).

Figure 158: Chemnitzia paucilirata. Holotype. Length, 2.7 mm

(x14).

Vol. 10; No. 4 THE VELIGER Page 417

Page 418

March 1850, takes precedence over Jeffreysia FoRBES &
Han ey, May, 1850.

125. Jeffreysia tumens, p. 363
= Rissoella tumens (CARPENTER, 1857)
(Text figure 71)
Tablet 1719, 2 syntypes. [1.2 mm]. Brann: plt. 39, fig.
421.

The specimen figured by Bartscu (1920, pp. 160- 161;
plt. 12, fig. 1) seems to be correctly identified. The shell
figured by Baker, Hanna, & STRONG (1930, p. 36; plt. 1,
fig. 13) has a more pointed apex and a small umbilicus.

126. ?Jeffreysia alderi, p. 362
= Barleeia alderi (CARPENTER, 1857)
(Text figure 72)

Tablet 1718, 3 syntypes. [1.7 mm]. Brann: plt. 39, fig.
420.

The allocation to Barleeia was made by BartscH
(1920, p. 175; plt. 12, fig. 6). Baker, HANNA, & STRONG
(1930, p. 38) also have given a figure of a specimen,
neither illustration being of type material.

RISSOINIDAE

127. Rissoina woodwardi, p. 357
= Rissoina woodwardii CARPENTER, 1857
(Text figure 73)

Tablets 1706 - 1707, 13 syntypes. [3.1 mm]. Brann: plt.
39, fig. 410 (also figured by Sowerby in REEvE 1878
(Conch. Icon., vol. 20, Rissoa pit. 11, fig. 104).

In his manuscript plates (published by BRANN) Car-
penter omitted the numeral for tablet 1707, which should
be below the figure to the right of “410” on plate 39.
There seem to be no problems with this form, of which
Carpenter had 15 syntypes. There are now only 13 entire
shells and two fragments. Carpenter cited one unusually
large specimen as length 4.1mm. My drawing of the
largest indicated 3.6 mm.

128. ?Rissoa lirata, p. 358
= ?Rissoina (s.l.) lirata (CARPENTER, 1857)
(Text figure 74)

Tablet 1708, 2 syntypes. [4 mm]. Brann: plt. 39, fig. 411.

Instead of publishing Carpenter's manuscript drawing,
as he did for other Mazatlan forms, Bartscu (1911, p.
338; plt. 29, fig. 3) illustrated a specimen from the U.S.
National Museum collection, and his figure has been the
basis for the misidentification of a relatively common
form as “Alvania lirata CARPENTER.” Carpenter described
and his figure shows an operculum with an apophysis,
like that of Rissoina; the operculum is still in place on
the tablet. In his errata (page 552) he transferred the

THE VELIGER

Vol. 10; No. 4

species to Barleeia. The figured syntype (here chosen as
lectotype) is more slender and tapering than “A. lirata”
of authors, with regular spiral riblets crossed by a few
low, angular axial ribs; there are faint color bands, and
the suture is smoothly appressed, not channeled or bor-
dered by nodes. A specimen in the California Academy
of Sciences collection from San Francisquito Bay, Gulf
of California, matches the figure of Carpenter’s type
well.

If this form is correctly to be assigned to Rissoina it
makes the fifth of a group of Gulf of California species
having spiral sculpture predominating. The others listed
by Baker, HANNA, & STRONG (1930, p. 34) are: R. la-
pazana Bartscu, 1915; R. kelseyi DaLtL & BartTscH,
1902; R. berryi and R. stephensae BAKER, HANNA &
STRONG, 1930. The group may prove to deserve a sepa-
rate generic name or may turn out to be closer to Duala.
At present we have insufficient knowledge about the
type species of Diala.

CAECIDAE

129. Caecum clathratum, p. 322
= Caecum clathratum CarPENTER, 1857
(Text figure 75)
Tablet 1528, 4 syntypes. [2.5mm]. Brann: plt. 34, fig.
369.
The largest of 4 syntypes is illustrated here.

130. Caecum ?quadratum, var. compactum, p. 322
— Caecum compactum CarPENTER, 1857
(Text figure 76)
Tablet 1530, 7 syntypes. [ - ].

I have drawn the syntype that is third from the right
on the tablet. Carpenter did not prepare a camera lucida
drawing, not having been convinced of the validity of
the form as separate. The syntypes comprise a growth
series.

131. Caecum elongatum, p. 319
= Caecum elongatum CarPENTER, 1857
(Text figure 77)
Tablet 1525, 4 syntypes. [2.6 mm]. Brann: plt. 33, fig.
366.
My drawing is of the largest of 4 specimens on the
tablet.

132. Caecum farcimen, p. 326
= ?Caecum farcimen CaRPENTER, 1857
(Text figure 78)
Tablet 1544, 4 syntypes. [1.6 mm]. Brann: plt. 36, fig.
3/3:

Vol. 10; No. 4

There are 2 adult syntypes. Annulations are present
though faint. Carpenter assigned this to his “section”
Fartulum, which otherwise is made up of smooth forms.

133. Caecum quadratum, p. 322
= Caecum quadratum CarPENTER, 1857
(Text figure 79 a-b)

Tablet 1529, 7 syntypes. [1.8mm]. Brann: plt. 35, fig.
370.

I have drawn the two largest among the seven. There
are about 17 somewhat flattened annulations, only the
terminal ones being sketched in in my drawings.

134. Caecum ?elongatum, var. semilaeve, p. 319
; = Caecum semilaeve CARPENTER, 1857
(Text figure 80)
Tablet 1526, 2 syntypes. [ — ].

I am illustrating the larger of the syntypes; the annu-
lations are weak and irregularly developed. Carpenter did
not prepare a camera lucida drawing for this form, as he
was not confident of its validity.

135. Caecum subimpressum, p. 320
=Caecum subimpressum CARPENTER, 1857

(Text figure 81)
Tablet 1527, 7 syntypes. [3 mm]. Brann: plt. 34, fig. 367.
My figure is of the largest syntype. The annulations
are strong and recurved near the aperture.

136. Caecum undatum, p. 323
— Caecum undatum CarPENTER, 1857

(Text figure 82)

Tablets 1531 - 1538. [1.8 mm]. Brann: plts. 35, 36, fig.
Sale

Of the numerous syntypes, I have selected for my
drawing the second from the right on tablet 1535. The
shell resembles that of Fartulum but with low ribs, its
surface otherwise shiny and smooth.

137. Caecum heptagonum, p. 319
= Elephantanellum heptagonum (CarPENTER, 1857)
(Text figures 83 a-b)
Tablet 1524, holotype. [0.5 mm]. Brann: plt. 32, fig. 365.
This is the type species of the genus Elephantanellum
BartscuH, 1921. A figure of a better-preserved specimen
has been given by StRoNG & HERTLEIN (1939, p. 226; plt.
20, fig. 9) from Panama.

138. Caecum liratocinctum, p. 317
= Elephantanellum liratocinctum (CARPENTER, 1857)
(Text figures 84 a-b)
Tablets 1518 - 1519, 8 syntypes. [3.7 mm]. BRANN: plts.
32, 33, fig. 364.

THE VELIGER

Page 419

My drawings are of the largest specimens on both tab-
lets.

139. Caecum liratocinctum, var. subconicum, p. 318
= Elephantanellum subconicum (CARPENTER, 1857)
(Text figure 85)
Tablet 1522, 2 syntypes. [-]. Brann: plt. 33, fig. 364
(part).

It may well be that this variety and the next are within
the range of variation of the species. There are two speci-
mens, one broken near the aperture, the other mounted
with its convex side down.

140. Caecum liratocinctum, var. subobsoletum, p. 318
=Elephantanellum subobsoletum (CarPENTER, 1857)
(Text figure 86)

Tablet 1521, 2 syntypes. [-]. BRANN: plt. 33, fig. 364
(part).

141. Caecum liratocinctum, var. tenuiliratum, p. 318
= Elephantanellum tenuiliratum (CARPENTER, 1857)
(Text figure 87)
Tablet 1520, holotype. [-]. Brann: plt. 33, fig. 364
(part).
One is tempted to transfer this form to Elephantulum,

for there is little evidence of transverse sculpture. The
longitudinal ribs are fine and sharply cut.

142. Caecum abnormale, p. 316
= Elephantulum abnormale (CarPENTER, 1857)
(Text figure 88)

Tablet 1516, 1 syntype. [1.2 mm]. Brann: plt. 32, fig.
362.

143. Caecum insculptum, p. 315
= Elephantulum insculptum (CarpeNnTER, 1857)
(Text figures 89 a-b)
Tablet 1514, 2 syntypes. [3.3 mm]. BRANN: plt. 32, fig.
360.

The longitudinal ribs are well spaced on both syntypes,
one of which Carpenter regarded as “old,” the other
“young.”

144. Caecum obtusum, p. 317
= Elephantulum obtusum (CARPENTER, 1857)
(Text figure go)
Tablet 1517, 2 syntypes. [3.2 mm]. Brann: plt. 32, fig.
363.

I have drawn the larger of the two syntypes. The ribs

have a worn look and are faint, more like internal lirae.

Page 420

145. Caecum subspirale, p. 315
= Elephantulum subspirale (CARPENTER, 1857)
(Text figure 91)
Tablet 1515, 4 syntypes. [4 mm]. Brann: plt. 32, fig. 361.
Of the syntypes mentioned by Carpenter I have drawn
the adult. The longitudinal ribs are numerous and with
narrow interspaces.

146. Caecum dextroversum, p. 328
= Fartulum dextroversum (CARPENTER, 1857)
(Text figure 92)
Tablet 1548, 9 syntypes. [2.3mm]. Brann: plt. 37, fig.
376.
I have drawn the two syntypes at the right of the
tablet, one being an adult, the other juvenile.

147. Caecum glabriforme, p. 327

—=Fartulum glabriforme (CARPENTER, 1857)
Tablet 1546, originally 2 syntypes. [1.8 mm]. BRANN: plt.
37, fig. 374.

The young shell mentioned by Carpenter is now mis-
sing, and the adult has so deteriorated that it is unrecog-
nizable. The species will have to be interpreted on the
basis of Carpenter’s drawing and description.

148. Caecum reversum, p. 329
= Fartulum reversum (CARPENTER, 1857)
(Text figure 93)
Tablet 1549, holotype. [1.2 mm]. Brann: plt. 37, fig.
STi
The specimen is glued to the tablet in such a way that
the outline is hard to draw accurately.
149. Caecum teres, p. 329
= ?Fartulum teres (CARPENTER, 1857)
(Text figure 94)

Tablet 1550, 3 syntypes. [3 mm]. BRANN: plt. 37, fig. 378.
Of the three syntypes mentioned by Carpenter, one is

THE VELIGER

Vol. 10; No. 4

now unrecognizable on account of chemical deteriora-
tion; another is partly decorticated. The remaining one
is the largest. It shows banding, perhaps from wear,
which Carpenter well characterized as “mottling.” There
are also some faint and irregular growth lines and a few
weak longitudinal lines. One wonders if, when the shell
was fresh, it may not have been recognizable as an Ele-
phantanellum rather than a Fartulum, which Carpenter
thought it might be.

150. Caecum corrugulatum, p. 327
= ?Micranellum corrugulatum (CARPENTER, 1857)
(Text figure 95)

Tablet 1547, holotype. [1.9mm]. Brann: plt. 37, fiz.
37D

The specimen is now an indeterminate broken tube
filled with sediment. I suggest an allocation to Micranel-
lum on the basis of Carpenter’s drawing and description.

VERMETIDAE

151. Siphonium (Aletes) ?centiquadrus imbricatus, p. 302
= ?Vermetus sp., indet.

Tablet 1484, 1 syntype (?holotype). [Length, 15mm;
diameter of aperture, 3 mm].

Aware on my later visits to the British Museum that I
had earlier concentrated on the Vermetidae, I was less
rigorous in checking these slides, and thus I overlooked
the fact that neither Carpenter nor I had sketched the
specimen on slide 1484. I have therefore only the notes I
made in 1958 when I decided that the shell is too im-
mature for generic allocation. The coil has 2 to 3 whorls
and is 15 mm in diameter; the sculpture is evenly striate
rather than imbricate, as in Serpulorbis. However, were
this to be allocated to Serpulorbis it would jeopardize
the name of the well-known S. imbricatus (DUNKER,
1860) from Japan. Because young vermetids of less than
3 whorls cannot be assigned to a genus unless the nuclear

Explanation of Textfigures 159 to 171

Note: Stated lengths are camera lucida readings and may be as

much as 1/10 too high.

Figure 159: Eulimella obsoleta. Holotype. Length, 1.6mm (x 19).

Figure 160: Chemnitzia flavescens. Holotype. Length, 3.3 mm
(x11).

Figure 161: Chemnitzia gracillima. Syntype. Length, 3.4mm
(x11).

Figure 162: Odostomia indentata. Syntype. Length, 4.6 mm (x 7.5).

Figure 163: Chemnitzia tenuilirata. Syntypes. Length, a) 3mm
(incomplete) (x12); b) 2mm (x16).

Figure 164: Chemnitzia terebralis. Holotype. Length, 2.8mm
(x12).

Figure 165: Chemnitzia unifasciata. Holotype. Length, 2.6mm
(x 12).

Figure 166: Chemnitzia c-b-adamsii. Syntype. Length, 4.6mm
(x 7.5).

Figure 167: Chemnitzia undata. Syntype. Length, 2mm (x 16).

Figure 168: Chemnitzia prolongata. Syntype. Length, 6.4mm
(x 7.5).

Figure 169: Melampus olivaceus. Syntype. Length, 12.5 mm
(CAD)

Figure 170: Nacella peltoides. Hypotype. Length, 3.7mm (x 7).

Figure 171: Litiopa divisa. Syntype. Length, 2.7mm (x11).

Vol. 10; No. 4 THE VELIGER Page 421

Page 422

THE VELIGER

Vol. 10; No. 4

whorls and soft parts are preserved, it seems best to
regard this variety as indeterminate.

Carpenter cited two specimens, one in good condition,
the other worn. As he says that “Tablet 1484 contains the
characteristic specimen,” one might interpret this as holo-
type selection.

152. Bivonia contorta, p. 305
= Vermetus (Thylaeodus) contortus (CARPENTER,

1857)

Tablets 1489 - 1493, several syntypes. [23 mm].
The specimen on tablet 1490 was figured as lectotype
by Keen (1961, p. 201; plt. 55, fig. 3).

153. Bivonia contorta, var. indentata, p. 307
= Vermetus (Thylaeodus) indentatus (CARPENTER,
1857)
Tablet 1494, 3 syntypes. [13 mm].
The best of 3 specimens on this tablet was figured as
lectotype by Keen (1961, p. 202; plt. 55, fig. 4).

154. Bivonia albida, p. 307
Indeterminate vermetid
(Text figure 96)
Tablet 1495, 3 syntypes. [5.5 mm]. Brann: plt. 31, fig.
356.

Carpenter’s drawing and my own of the nuclear whorls
agree well. He thought that one of the 3 syntypes was
mature, but I do not find it identifiable. Several of the
tropical West American vermetids have juvenile shells
with this mode of coiling.

155. Petaloconchus macrophragma, p. 309, “n.s.”; Car-
PENTER, 1857 a [March], p. 313
= Petaloconchus (Macrophragma) macrophragma

CarRPENTER, 1857

Tablets 1499 - 1506. Brann: plt. 30, fig. 359.

Carpenter’s drawing is of the shell on tablet 1499,
showing the internal laminae. A specimen on tablet 1500
was selected as lectotype by Keen (1961, p. 205; plt. 55,
fig. 2). There is a typographical error in BRANN’s citation
of the registry number, which is 57.6.4.1500 (not 1494).
Carpenter validated the specific name prior to the pub-
lication of the Mazatlan Catalogue but based both de-
scriptions on the same type material.

CERITHIDAE

156. Cerithium ?famelicum, var. mediolaeve, p. 334
= Cerithium mediolaeve CARPENTER, 1857

(Plate 57, Figure 51)
Tablet 1587, holotype. [25 mm].

Because Cerithium famelicum C. B. Avams, 1852, has
been ranked as a synonym of C. uncinatum (GMELIN,
1791), Carpenter’s unillustrated variety has also been
synonymized. However, the photograph shows that it is
not only a smoother shell than C’. uncinatum, with sub-
dued color markings, but it is much more slender. The
recurved canal and placement of sculptural elements
relates it more closely to C. nicaraguense Pitspry & Lowe,
1932, from which it differs in its slightly smaller size and
more slender outline. Judging by the holotype, I conclude
that this may be a distinct species and one to be watched
for in West Mexican material.

157. Cerithium alboliratum p. 336,
= ?Cerithium maculosum KienEr, 1841

(Plate 57, Figure 48; Text figure 97)

Tablet 1591, 4 syntypes. [3.1 mm]. Brann: plt. 38, fig.
385.

Carpenter’s drawings illustrate 2 of the 4 syntypes. My
drawing and photograph are of the only one now on the
tablet that is unquestionably a Certthium, and this spe-
cimen is here selected as lectotype. BAKER, HANNA, &
Strone (1938, p. 225; plt. 17, fig. 7) seem to have recog-
nized the form correctly from Carpenter’s description and
from a figure by REEvE, 1866 (Conch. Icon., vol. 15, plt.
16, sp. 109) of a specimen said to be in the Cuming col-
lection, British Museum. I am not convinced that this is
a distinct species, for it seems to me to be the juvenile
tip of C. maculosum. The other syntypes on the tablet
are, I think, specimens of Alabina and Bittium.

158. Cerithiopsis ?tuberculoides, var. albonodosa, p. 443
= Cerithiopsis tuberculoides CARPENTER, 1857
(Text figure 98)
Tablet 2029, 1 juvenile syntype and fragments. [ — ].
Brann: plt. 48, fig. 557-b.
This seems to be a color variant and a juvenile shell
otherwise identical to Cerithiopsis tuberculoides. Car-
penter cited no measurements.

159. Cerithiopsis cerea, p. 443
= Cerithiopsis cerea CARPENTER, 1857

(Text figure 99)

Tablet 2030, holotype. [2.4 mm]. Brann: plt. 48, fig. 558.

As with other of Carpenter’s species of Cerithiopsis,
BartscH (1911b, p. 333; plt. 37, fig. 6) published the
same camera lucida sketch later republished by BRANN,
and he reprinted Carpenter’s description without com-
ment. It is obvious that he was not able to recognize the
species in the material he had available. Carpenter’s
drawing is distorted as to outline.

Vol. 10; No. 4

THE VELIGER

Page 423

160. Cerithiopsis pupiformis, p. 443
—Cerithiopsis pupiformis CARPENTER, 1857
(Text figures 100 a-b)

Tablet 2031, 2 syntypes. [1.8mm]. BRANN: plt. 48, fig.
559.

One wonders whether Carpenter had correctly matched
the two specimens he mounted as syntypes, the apex of
a juvenile shell and the last few whorls of an adult.
Drawn to a common scale, as I have done them, they do
not seem to match in contour. The young shell has strong
sculpture, the upper of 3 threads being weakest. BarTscH,
as mentioned above, merely reprinted Carpenter’s de-
scription (op. cit., p. 337; plt. 38, figs. 1, 5).

161. Cerithiopsis sorex, p. 444

= ?Cerithiopsis sorex CARPENTER, 1857
(Text figure 101)

Tablet 2032, 1 syntype. [1.6mm]. Brann: plt. 49, fig.
560.

Carpenter’s drawing does not show, nor does he men-
tion, the slot-like posterior notch, which gives the aper-
ture in the holotype the semblance of a nassariid. I have
not found this structure so well developed in actual spe-
cimens. One of what appears otherwise to be Cerithiopsis
sorex, in the Stanford University collection, from near
La Paz, Baja California, has a newly-mended outer lip
that has not reached its full development; also, the
pointed apex is broken off. A species from Panama, C.
eiseni STRONG & HERTLEIN, (1939, p. 216; plt. 20, fig. 6)
seems to have the same contours of the later whorls, but
it, too, lacks apical whorls in the figured holotype. Another
similar form is C. perrini HERTLEIN & STRONG, 1951
(Zoologica, vol. 36, no. 5, p. 106; plt. 7, fig. 6) from Port
Guatulco, Mexico.

162. Cerithiopsis tuberculoides, p. 442
= Cerithiopsis tuberculoides CARPENTER, 1857
(Text figures 102 a-b)

Tablet 2028, 2 syntypes. [1.4mm; 4mm]. Brann: plt.
48, fig. 557.

The younger of the two syntypes was chosen by Car-
penter for his drawing. The apex in this has 3 smooth
whorls. The adult has a broken aperture; its apex is
somewhat imperfect but the remainder of the shell shows
the sculpture well, 3 beaded cords per whorl equal in
strength.

163. Cerithiopsis convexa, p. 444
= Metaxia convexa (CARPENTER, 1857)
(Text figures 103 a-b)
Tablet 2033, holotype. [4.5 mm]. Brann: plt. 49, fig. 561.

Authors have been able to recognize this species
correctly.

164. Cerithiopsis decussata, p. 445
= Bittium decussatum (CARPENTER, 1857)
(Text figure 104)

Tablet 2034, holotype. [4mm]. Brann: plt. 49, fig. 562.
BartscH (1911 c, p. 409; plt. 52, fig. 2) transferred
the species to Bitttwm, reprinted Carpenter’s description,
and published his manuscript drawing. It is evident that
Bartsch did not have material at hand.
[Note: Carpenter has been criticized for having bestowed
names on fragmentary and unrecognizable material. His
usage of Alaba as a generic term is especially open to
such criticism. Because all but one of the species belong
elsewhere and so many are unrecognizable, I have listed
all of his named Alabas here, alphabetically instead of
systematically. Possibly some of the forms can be matched
with fresh material eventually. One can hope that the
names will prove useful and do not displace later names
based on more complete specimens. |

165. Alaba alabastrites, p. 368
a)

Tablet 1726, holotype. [1.4 mm]. Brann: plt. 40, fig. 428.

The holotype is only the apical tip of a shell, the
nucleus and 3 whorls of a slender form that could be
turrid, columbellid, or rissoinid; therefore, it seems gener-
ically and specifically indeterminate.

166. Alaba conica, p. 368
?—=Crerithium adustum Kiener, 1841

Tablet 1728. [2 mm]. Brann: plt. 40, fig. 430.

The species must be recognized, if it can be at all, from
Carpenter’s description and drawing and from 3 syntypes
in other collections, for the one placed in the British Mu-
seum collection is now missing. At some time the end of
the vial had broken and the specimen, having come un-
glued, had lost out. I found no trace of it in the box.

167. ?Alaba laguncula, p. 369
?Iselica sp.
(Text figure 105)
Tablet 1730, holotype. [1.1 mm]. Brann: plt. 40, fig.
432.

The specimen consists of the nuclear whorls and part
of one spire whorl; the apex seems to be partially im-
mersed or tilted, and as the sculpture suggests Iselica,
the form may be a pyramidellid. My drawing is only a
sketch to show outline and is not to scale.

Alaba mutans, p. 369. See no. 122, under Rissoidae.

168. Alaba scalata, p. 368
?Epitonium sp., juvenile
(Text figure 107)
Tablet 1727, holotype. [0.9 mm]. Brann: plt. 43, fig. 429.

Page 424

TES VEEICER

Vol. 10; No. 4

The juvenile shell probably is specifically and subge-
nerically indeterminate.

169. Alaba supralirata, p. 366
= Alaba supralirata CARPENTER, 1857

Tablet 1723, 3 syntypes. [4.8mm]. Brann: plt. 43, fig.
425.

This is Carpenter’s only Alaba that really qualifies for
inclusion. The smallest specimen has 6 whorls, and the
apex is well shown in Carpenter’s drawing; the middle-
sized shell shows the early post-apical sculpture that is
portrayed by Bartscu (1910, pp. 153 - 156, figs. 1-2).
The largest specimen has obscure sculpture but strong
varices, its terminal whorl shown in Carpenter’s figure.

Bartsch has quoted E. A. Smith’s criticism of Carpen-
ter’s overnaming in this genus.

170. Alaba terebralis, p. 367
?Fulima sp.
(Text figure 108)

Tablet 1725, holotype. [1.7 mm]. Brann: plt. 40, fig.
A2ie

The tapering and loosely coiled apex that Carpenter
shows now has chipped away to a single whorl that
appears obliquely twisted.

171. Alaba violacea, p. 367
?Eulima sp.
(Text figure 109)
Tablet 1724, holotype. [1.5mm]. Brann: plt. 40, fig.
426.
The shell is broken and one must guess at the true
outline.

172. Alvania effusa, p. 359
= Alabina effusa (CARPENTER, 1857)
(Text figure 110)

Tablet 1710, holotype. [3 mm]. Brann: plt. 39, fig. 413.

BartscH (1911 d, p. 358; plt. 32, fig. 5), reproduced
Carpenter’s drawing and continued this in Alvania rather
than Alabina, which he also reviewed (1911 a). It seems to
me to have all the characters of Alabina, for no other
Alvania is so slender and tall.

173. ?Alvania excurvata, p. 359

= Alabina excurvata (CARPENTER, 1857)
(Text figure 111)

Tablet 1709, 8 syntypes. [3 mm]. Brann: plt. 39, fig. 412.
This is a prior name for Alabina diomedeae Bartscu,
1911 (1911 a, p. 413; plt. 62, fig. 1), an abundant small
mollusk in the Gulf of California. One wonders that
Bartsch failed to recognize the form from the Carpenter
drawing, which he had available. Under the Article 23(b)

of the International Code of Zoological Nomenclature,
Bartsch’s specific name might be declared a nomen ob-
litum, for it has been a junior synonym more than the
requisite 50 years, but at present the International Com-
mission is at an impasse on the proper procedure for
implementing this rule.

PoTAMIDIDAE

174. Cerithidea ?varicosa SowERBY, var. mazatlanica, p.
344
= Cerithidea mazatlanica CARPENTER, 1857
(Plate 57, Figures 47 a-d)
Tablets 1628 - 1637, 33 syntypes (4 on tablet 1630 fig-
ured). [25 mm].

Authors have had no problems with the identification
of this form. A specimen from the Cuming collection was
figured by Sowerby in REEvE, 1866 (Conch. Icon., vol.
15, Cerithidea, plt. 1, sp. 8).

HiprponicDAE

175. Hipponyx serratus, “n.s.”, p. 296
= Hipponix serratus (CARPENTER, 1856)
(Plate 57, Figures 43 a-b)
Tablets 1462 - 1468, 12 syntypes. [25mm]. BRANN: plt.
31, fig. 346.

Two syntypes on tablet 1465 are figured here. The
species name was validated earlier (CARPENTER, 1856 b,
p. 3) based on other British Museum as well as Reigen
material.

176. Hipponyx planatus, p. 298
= Hipponix planatus (CARPENTER, 1857)
(Plate 57, Figures 42 a-b; Text figures 112 a-b)

Tablet 1470, 2 syntypes. [1.5mm - 5mm]. Brann: plt.
31, fig. 348.

The Mazatlan material consisted of immature speci-
mens. One lot of adult shells from Panama (Cuming
collection) was also cited; it is now in the Type Collec-
tion, British Museum, registry no. 1966625. [19 mm].

FOSSARIDAE

177. Fossarus angulatus, p. 354

= Fossarus angulatus CARPENTER, 1857
Tablet 1701, 1 syntype. [1.8mm]. Brann: plt. 38, fig.
405.

The syntype originally retained at the British Museum
is now almost entirely disintegrated; there appear to have
been 2 carinae, a wide columellar lip, and a large aper-
ture with a thickened rim, but Carpenter’s drawing is -
the only real clue to identity.

Vol. 10; No. 4

178. Fossarus tuberosus, p. 354
= Fossarus tuberosus CARPENTER, 1857

(Text figure 113)

Tablet 1700, 2 syntypes. [1.1 mm]. Brann: plt. 38, fig.
404.

Carpenter’s drawing should enable the identification
of this form. The smaller syntype shows 2 brown nuclear
whorls and 4 carinae on the body whorl; the larger has
3 brown nuclear whorls and fine spiral intercalary ribs.

CALYPTRAEIDAE

179. Trochita ventricosa, p. 264
_ =Calyptraea (Trochita) spirata ventricosa (Car-
PENTER, 1857)
(Plate 57, Figures 55 a-b)
Tablet 1316, holotype. [19 mm].
The holotype is glued to the mount at an angle diffi-
cult for proper photography.

CREPIDULIDAE

180. Crepidula >dorsata Brod., var. bilobata, p. 273
?—=Crepidula aculeata (GMELIN, 1791)
(Plate 57, Figure 56; Text figure 114)

Tablet 1354, 5 syntypes. [9.5mm]. Brann: plt. 28, fig.
336.

Carpenter’s drawings are of the 4 smallest syntypes.
My photograph shows the exterior of the largest. Al-
though covered by an incrustation, it shows a pattern of
large and small spines as in Crepidula aculeata. Carpen-
ter describes the deck as brown; perhaps it was, in fresh
material, but to my eyes it is now white. All the speci-
mens were immature. Possibly the name may have some
use if a brown-decked variant of C. aculeata comes to
light.

MuricipAE

181. Murex ?recurvirostris lividus, p. 519
= Murex (Murex) lividus CARPENTER, 1857
(Plate 58, Figures 63 a-b)
Tablets 2467 - 2480, 23 syntypes (specimens from tablet
2477 here figured). [52 mm].

This form seems to be morphologically distinct from
Murex recurvirostris BRoDERIP, 1833, in which spines are
obsolescent, and M. elenensis Dat, 1919, in which they
are numerous. In M. lividus spines are sparsely present
on spire and canal.

THE VELIGER

Page 425

182. Muricidea ?erinaceoides, var. indentata, p. 527
= Aspella indentata (CARPENTER, 1857)
( Plate 58, Figure 64)

Tablet 2510, holotype. [34 mm].

A few months after describing Aspella perplexa KEEN,
1958, I had opportunity to see Carpenter’s holotype and
— perhaps too hastily — decided my species must fall as
a synonym. Now with a photograph of the type and more
Mexican material, I suspect our Aspellas all need re-
study.

CoLUMBELLIDAE

183. Anachis albonodosa, p. 512
= Anachis albonodosa CARPENTER, 1857
(Plate 59, Figure 82)

Tablet 2432, syntype. [3.3 mm]. BRANN: plt. 56, fig. 654.

Axial sculpture begins on the fourth whorl of the
otherwise smooth spire and fades out on the base of the
body whorl; the ribs are smooth and end above in white
nodes. The aperture is contracted and the outer lip thin,
perhaps immature. I have not as yet found among the
numerous available specimens of Anachis of the West
Mexican area in collections I have studied one that I
could positively identify as A. albonodosa. Perhaps now
that figures are published, collectors may be able to
recognize the form.

184. Anachis gaskoini, p. 510
= Anachis gaskoini CARPENTER, 1857
(Plate 59, Figure 86; Text figure 115)

Tablet 2430 (sketch). [6 mm].

This form was described by Putuipr1 in 1846 as Colum-
bella taeniata, a name preoccupied by C. taeniata Linx,
1807. Before Carpenter had accidentally smashed the
type specimen of a Mazatlan form his friend Gaskoin
had pronounced a new species, he had made a sketch,
which is reproduced in facsimile here. Although poor, it
is recognizable as the C. taeniata Puitipri of authors and
is reinforced both by Carpenter’s label on a specimen in
the British Museum collection from Callao, Peru, registry
number 79.2.26.109 (see Plate 59, Figure 86) and by
his synonymizing of the two names later (CARPENTER,
1865 b, p. 273). Dati in 1918 named Anachis bartschit,
which has remained unfigured, but photographs of the
type lot show it to be identical to Carpenter’s species.
Baker, Hanna, & STRONG (1938, Proc. Calif. Acad. Sci.,
ser. 4, vol. 23, no. 16, p. 249; plt. 24, fig. 11), working
from the description alone, figured a specimen as A.
bartschu that is a yet-undescribed species. Unfortunately,

Page 426

I copied their figure (KEEN, 1958, fig. 421). GranT &
Gate (1931, p. 688; plt. 26, fig. 47) attempted to illust-
rate A. gaskoini, but their figure turns out to be of a
variant that also is unnamed. Again — unfortunately —
I copied the figure (Keen, 1958, fig. 433). I plan a
review of some of these West Coast Columbellidae to
clarify the nomenclature. The true A. gaskoini is white,
with golden-brown spiral lines and a few dark brown
dots. ‘The species is not uncommon but up to now has not
been adequately illustrated.

185. Anachis nigrofusca, p. 509
= Anachis nigrofusca CARPENTER, 1857

Tablet 2427, 2 syntypes. [10 mm].

Authors have identified this species correctly. The syn-
types show spiral striae between axial ribs more distinctly
than in the specimen figured in KEEN, 1958 (p. 383; fig.
445) but the identity is obvious.

186. Anachis ?costellata, var. pachyderma, p. 507
?—= Anachis scalarina (SowErRBy, 1832)
(Plate 59, Figures 83 a-b)
Tablets 2422 - 2423, 7 syntypes. [17 mm]. Brann: plt.
56, fig. [646b] (operculum).

Carpenter described this as a variant of Anachis costel-
lata, but it seems closer to A. scalarina; perhaps this
northern form may prove subspecifically distinct because
of the periostracum and smaller size.

187. Anachis rufotincta, p. 511
= Anachis diminuta (C. B. ApaMs, 1852)

Tablet 2431, 3 syntypes. [3.5mm]. Brann: plt. 56, fig.
653.

Looking at the type material my judgment was that
these were only faded specimens of Anachis diminuta,
a wide-ranging Panamic species.

188. Anachis serrata, p. 509
= Anachis (?Glyptanachis) serrata CARPENTER, 1857
(Plate 59, Figures 87; Text figure 116)

Tablet 2428, 3 syntypes, all juvenile, the largest here se-
lected as lectotype. [3.3 mm]. BRANN: plt. 56, fig. 650.

The two fragments on the tablet, mentioned by Car-
penter, are of other species, and only the 3 specimens at
the left are of this form. The relationships are still in
doubt, for it may prove to be a Nassarina rather than an
Anachis. I plan further work on the West American
columbellids.

189. Columbella cervinetta, p. 493
= M*itrella baccata (Gasxoin, 1852)
(Plate 59, Figure 84)
Tablet 2360, holotype. [6.9 mm]. Brann: plt. 55, fig. 618.

Carpenter was evidently unfamiliar with Gaskoin’s
species. His figure shows well the axial ribbing on the

THE VELIGER

Vol. 10; No. 4

spire that distinguishes this and another as-yet-unnamed
Gulf Mitrella.

190. Columbella cervinetta obsoleta, p. 493
= Mitrella baccata (Gasxotn, 1852)
(Plate 59, Figure 85)
Tablet 2361, syntype. [—]. BRaNN: plt. 55, fig. 618-b.
I can see no reason for separating this form, merely a
worn and faded shell.

BuccinmAE

191. Fusus apertus, p. 504
?==Cantharus biliratus (RrEve, 1846)
(Text figure 117)
Tablet 2414, 3 incomplete syntypes. [2.5 mm]. Brann:
pit. 53, fig. 641.

The best specimen shows 4 brown bands and nodose
sculpture, suggesting young Cantharus. Specimens of C.
biliratus from Guaymas, collected by D. R. Shasky, match
it well as to details of apical sculpture.

192. Pisania aequilirata, p. 515
== Cantharus elegans (GrirrirH & PipcEon, 1834)
(Plate 59, Figure go)
Tablet 2451, holotype. [25 mm].
Authors have correctly recognized this from Carpen-
ter’s description.

NASSARIIDAE

193. Nassa pagodus, var. acuta, p. 497 [non Say, 1822]
?—=Nassarius pagodus (REEVE, 1846), ?n. subsp.
(Plate 58, Figure 61)
Tablet 2394, 1 syntype. [21.5 mm].

The shell has finer spiral sculpture than typical Nassa-
rius pagodus, and the axial ribs are fewer. If this is
consistent, perhaps the form may have validity as a sub-
species, but a new name will be required, Carpenter’s
having been preoccupied.

194. Nassa crebristriata, p. 499
= Nassarius versicolor (C. B. ApaMs, 1852)
(Plate 58, Figure 60)
Tablet 2402, holotype. [13.8 mm].
This seems to be a slender form of the variable Nassa-
rius versicolor.
195. Nassa ?tegula, var. nodulifera, p. 496

ex Puiuipr1, MS
—=Nassarius luteostoma (BRODERIP & SOWERBY, 1829)

(Plate 58, Figure 59)
Tablet 2393, holotype. [16.2 mm].
As Carpenter suspected, this is a needless name.

Vol. 10; No. 4

THE VELIGER

Page 427

FUSINIDAE (?)

196. Fusus tumens, p. 503
?— Fusinus cinereus (REEVE, 1847)

(Text figure 118)
Tablet 2413, 2 syntypes. [4 mm]. Brann: plt. 53, fig. 640.
The shell is whitish, banded at the suture and on the
base with brown. It appears to be a juvenile Fusinus.

OLIVIDAE

197. Oliva intertincta, p. 465
= Oliva spicata Rovinc, 1798
(Plate 58, Figure 57)
Tablet 2121, 3 syntypes. [20 mm].
The two larger syntypes are in good condition; the
smallest is partially decorticated.

198. Olivella ?petiolita, var. aureocincta, p. 470
= Olivella aureocincta CARPENTER, 1857

(Plate 59, Figures 88 a-b)

Tablet 2186, 8 syntypes. [9.5 mm].

All of these syntypes are beginning to show deteriora-
tion of the surface layer, but neatsfoot oil restores to
some extent the color pattern, especially in the best-
preserved two that are figured here. The shell is near
Olivella dama (Woop, 1828) in outline but of smaller
size. The color pattern is undulating to zigzag. When
the shells were fresh, according to Carpenter’s descrip-
tion, there were two spiral color bands of a golden hue.
This form probably is the basis for the records of the
Caribbean O. petiolita (Ductos, 1835) in the tropical
West American fauna.

MarGINELLIDAE

199. Marginella margaritula, p. 462
= Cypraeolina margaritula (CARPENTER, 1857)
(Text figure 119)
Tablet 2109, 7 syntypes (2 adult). [1.8mm]. Brann:
plt. 49, fig. 589.
The larger adult specimen was chosen as lectotype by
Coan & Rotu (1966, The Veliger, vol. 8, no. 4, p. 294).

200. Marginella polita, p. 462
= Kogomea polita (CaRPENTER, 1857)
Text figures 120 a-c
Tablet 2108, 3 syntypes. [0.8 mm]. Brann: plt. 49, fig.
588.
Coan & RotH (1966, The Veliger, vol. 8, no. 4, p.
293) have selected the largest specimen as lectotype.

TURRIDAE

201. Drillia albonodosa, p. 397
= Crassispira albonodosa (CARPENTER, 1857)
(Plate 59, Figure 100)

Tablet 1901, 1 syntype (here chosen as lectotype). [13
mm].

This seems to be a distinct species. It comes closest to
Clathrodrillia jaculum (Pirspry & Lowe, 1932) as to
sculpture and outline, but its sooty color puts it in Crassi-
Spira.

202. Drillia albovallosa, p. 396
= Crassispira rudis (SowERBY, 1834)
(Plate 59, Figure 98)

Tablet 1900, holotype. [10.8 mm].

Someone (probably E.A.Smith, possibly J. R. leB.
Tomlin) has put the notation, “D. rudis, jun.” on the
tablet. A juvenile topotype specimen of the latter in the
Stanford University collection matches well my photo-
graph of Carpenter’s type; so also does a juvenile speci-
men from Mazatlan. Adults show a greater convexity of
outline near the base of the body whorl that changes the
appearance, and this probably influenced Carpenter in
his decision that the form was distinct.

203. Drillia cerithoidea, p. 394
= Crassispira cerithoidea (CARPENTER, 1857)
(Plate 58, Figure 77; Text figure 121)

Tablet 1897, 1 syntype. [16mm].

This may be a prior name for the Crassispira pluto
Pitspry & Lowe, 1932. Comparison of material collec-
ted and identified by Lowe with Carpenter’s type showed
only minor differences, such as slightly coarser nodes on .
the spire of C. pluto in some specimens.

204. Drillia hanleyi, p. 398
= Crassispira hanleyi (CARPENTER, 1857)

(Plate 59, Figure 99)
Tablet 1907, holotype. [9.8 mm].
This seems to be distinct. Although close to Crassispira
ericana HERTLEIN & STRONG, 1951, it is proportionately
wider and shorter.

205. Drillia aterrima, var. melchersi (MENKE, 1851),
ja, BE
= Crassispira aterrima (SowErRBy, 1834)

Tablets 1891 - 1896. [18 mm].

Although he used the same format that he did for
proposal of new varieties, Carpenter included in his
synonymy a reference to Menke’s proposal of the name.
The name is available for subdivision of the somewhat

Page 428

variable Crassispira aterrima, but Menke’s type would
need study.

206. Drillia monilifera, p. 395
—Crassispira monilifera (CARPENTER, 1857)
(Plate 58, Figure 76)
Tablet 1899, holotype. [16.4 mm].

Crassispira nymphia Pitspry & Lowe, 1932, is very
close to this. Comparison of specimens collected and
identified by Lowe with Carpenter’s type showed only
that the peripheral nodes in C. nymphia are coarser and
fewer. The nodes are yellow-orange, and the spiral
threads on the pillar beaded with yellow.

207. Mangelia sulcata CARPENTER, 1865b, p. 272
= Mangelia (Kurtzina) sulcata CARPENTER, 1865

(Text figure 122)

Tablet 2538, holotype. [5 mm]. Brann: plt. 41, fig. 702.
The shell resembles Mangelia (Kurtzina) cymatias

Pitssry & Lowe, 1932, but the axial ribs are wider; the

upper part of the whorls shows a faint brown band.

208. Mangelia ?acuticostata, var. subangulata, p. 400
—= Mangelia (Kurtzina) subangulata (CARPENTER,
1857)
(Plate 58, Figure 62; Text figure 123)

Tablet 1914, holotype. [3.5 mm]. Brann: plt. 41, fig. 473.

This also resembles Mangelia (Kurtzina) cymatias
Pitspry « Lowe but has one more axial rib per whorl.
The axial ribs are more sinuous, and there are faint
spirals showing between them. The anterior canal of the
holotype is a little broken. There is a discrepancy of
nearly a millimeter between my measurement of length
and Carpenter’s.

209. Clathurella aurea, p. 400
— Clathurella aurea CARPENTER, 1857

(Plate 58, Figure 74; Text figure 124)

Tablet 1913, holotype. [15.8 mm].

The nodes on the spiral ribs are more elongate than in
Clathurella (Lioglyphostoma) armstrong: HERTLEIN &
Stronc, 1955, which otherwise has a strong resemblance
to Carpenter’s species. The outer lip is denticulate with-
in; one wonders whether this is a reliable character for
differentiating between Clathurella s.s. and Lioglypho-
stoma. The outline of this and other West Coast turrids
now classed as Lioglyphostoma is closer to that of the
type of Glyphostomops Bartscu, 1934, from deep water
in the Caribbean than to the type of Lioglyphostoma,
also Caribbean.

THE VELIGER

Vol. 10; No. 4

TEREBRIDAE

210. Terebra (Myurella) albocincta, p. 384
= Terebra variegata Gray, 1834
(Plate 58, Figure 70)
Tablets 1828 - 1835, 23 syntypes (tablet 1828 photo-
graphed). [41 mm].

A note on the back of the mount, by G. K. Robson
says, “Designated type by me for Dr. Bartsch, 25/5/23,”
but neither Robson nor Bartsch published a lectotype
selection. CAMPBELL (1964, p. 137) has concluded, on
the basis of supposed paratype and of topotype material,
that this form is not separable from the widely distributed
Terebra variegata. The type lot supports his judgment;
Carpenter, however, felt that 7: armillata Hinps was
closer.

211. Terebra (Myurella) hindsii, p. 385
= Terebra variegata Gray, 1834
(Plate 58, Figures 71 a-b)
Tablet 1836, 2 syntypes. [32 mm].

As the photograph here given of the type material
supports the conclusion by CAMPBELL (1964, p. 137), we
must synonymize this as part of the Terebra variegata
complex. It is not the form from the outer coast of
Baja California identified as T: hindsit by GRANT & GALE
(1931, p. 469), which is now to be known as T: tiarella
DesHayeEs, 1857. Therefore, figures 955 and 964 in
KEEN, 1958, are incorrectly captioned.

212. Terebra (Myurella) rufocinerea, p. 386
= Terebra variegata Gray, 1834
(Plate 58, Figure 73)
Tablet 1838, 2 syntypes. [32 mm].
This, too, seems indistinguishable from Terebra varie-

gata. It is a form with the axial ribs slightly stronger and
straighter.

213. Terebra (Myurella) subnodosa, p. 386
= Terebra intertincta Hinps, 1844

(Plate 58, Figure 72)

Tablet 1837, 1 syntype. [32 mm].

Although, following authors, I synonymized this with
Terebra albocincta (see Kren, 1958, p. 490), the photo-
graph of the syntype shows a more stubby shell with
decided nodes on the base, a hallmark of the species T.
intertincta. Carpenter’s specific name has one month’s
priority over T: marginata DesHayes, 1857.

Vol. 10; No. 4

THE VELIGER

Page 429

BULLDAE

214. Haminea cymbiformis, p. 174
—= Haminoea cymbiformis CARPENTER, 1857
(Text figure 125)

Tablet 793, holotype. [1.8 mm]. Brann: plt. 19, fig. 229.

Carpenter’s restoration of the outline for the broken
outer lip in this minute juvenile shell differs somewhat
from mine. The apex is a shallow pit; there are fine and
somewhat sinuate growth lines. The shell is thin and
greenish-yellow in color.

215. Bulla exarata, p. 173
= Atys exarata (CARPENTER, 1857)
(Text figure 126)

Tablet 791, 1 syntype. [3.2 mm]. BRANN: plt. 19, fig. 227.

Carpenter’s drawing does not show the irregular break
in the front of the body whorl. The shell is brown, but
this may be a stain. The columellar lip has no umbilical
chink and no fold. There are 4 thin brown punctate
spiral lines near the apex, a smooth band below this,
then two wider lines, with 10 somewhat more close-set
lines toward the base. Though near Atys chimera BAKER
& Hanna, 1927, this may prove to be a good species.

JULODAE

216. Smaragdinella thecaphora, p. 533
= Julia thecaphora (CarPENTER, 1857)
(Text figure 127)

Tablet 2527, holotype. [2 mm]. Brann: plt. 18, fig. 692.
Oxsson, 1961 (Panama-Pacific Pelecypoda, p. 142)
has pointed out that Carpenter’s name has priority over
Julia equatorialis Ptitspry & Oxsson, 1944. I suspect
that the supposed J. exquisita Goutp, 1862, reported by
A. D. Howard from Baja California also is this species.

ACTEOCINIDAE

217. Tornatina carinata, p. 171
= Acteocina carinata (CARPENTER, 1857)
(Text figure 128)

Tablet 784, 5 syntypes. [2.8mm]. Brann: plt. 19, fig.
223.

Carpenter’s drawing does not show a hole in the body
whorl made by a carnivorous snail. The adult, shown
both in Carpenter's drawing and mine, does not exhibit
as clearly as do the younger syntypes the double ridge on
the spire that is characteristic of the species; the young
are relatively shorter and lower spired.

PyRAMIDELLIDAE

In the introduction to their comprehensive review of
West American Pyramidellidae, Dati & BartscH (1909,
p. 2) state that Bartsch had opportunity while in Britain
to study Carpenter’s Mazatlan types. I shall therefore
rely upon their allocations for these species. They also had
available and used as figures some of the manuscript
drawings that have now been published in full by Brann
(1966). I shall cite BRANN’s plates rather than the earlier
version of Dati & BartscH. Carpenter cited the generic
names in his introduction to the discussion of the family
but in the body of the text combined the specific names
with subgeneric names. Brann correctly lists the generic
and specific combination, but Dall and Bartsch utilize
the subgeneric-specific name combination. When Car-
penter's allocation is in complete agreement with that of
Dall and Bartsch, I shall, to save space and avoid repe-
tition, abbreviate generic and subgeneric names to the
initial letters.

218. Odostomia lamellata, p. 411
?— Pyramidella (Longchaeus) mazatlanica Dau. &
Bartscu, 1909
(Text figure 129)
Tablet 1954, 2 fragments. [2.5 mm]. Brann: plt. 42, fig.
489.

Datu & BartscH (1909, p. 24) cite this form as of
uncertain standing and suggest that it may the young of
their new species. Even the larger fragment is too broken
for positive determination.

219. Odostomia subsulcata, p. 411
?Pyramidella (Longchaeus) sp.
(Text figure 130)

Tablet 1955, 2 syntypes. [1.1 mm]. Brann: plt. 43, fig.
490.

I can add nothing to Bartsch’s comment that the speci-
mens are too young and worn for specific determination.

220. Odostomia vallata, p. 411
Pyramidella (Longchaeus) sp.
(Text figure 131)

Tablet 1956, 3 syntypes. [1.4mm]. Brann: plt. 43, fig.
ASIF

According to my notes, my drawing is of the best of
the 3 specimens and there is only a faint carina. Carpen-
ter’s drawing shows a strong one. In any case, the shell
seems to be unidentifiable; it is juvenile.

Page 430

221. Odostomia (Chrysallida) convexa, p. 424
— Odostomia (Besla) convexa CARPENTER, 1857

(Text figure 132)
Tablet 1984, 1 syntype. [2 mm]. Brann: plt. 45, fig. 514.
This species is type of the subgenus Besla Dati &
BartscH, 1904.
222. Odostomia (Chrysallida) effusa p. 422
=O. (C.) effusa CARPENTER, 1857
(Text figure 133)
Tablet 1980, holotype. [2.2 mm]. Brann: plt. 45, fig. 510.
A supposed younger specimen glued to the tablet by
Carpenter seems to be an Alvania, cf. A. tumida CaRPEN-
TER. I regard the larger shell as a holotype because
Carpenter cites it as “the specimen.”
223. Odostomia (Chrysallida) fasciata, p. 423
=O. (C.) fasciata CARPENTER, 1857
(Text figure 134)
Tablet 1981, 5 syntypes. [2.1 mm]. Brann: plt. 45, fig.
Silt,
The interspaces appear to be punctate, and the nodes
on the spire are well developed.
224. Odostomia (Chrysallida) nodosa, p. 417
— QO. (C.) nodosa CARPENTER, 1857
(Text figure 135)
Tablet 1969, 2 syntypes. [4.1 mm]. Brann: plt. 44, fig.
504.
One of the syntypes is juvenile.
225. Odostomia (Chrysallida) oblonga CARPENTER, 1857,
p. 418 (non Maccituivray, 1848)
=O. (C.) benthina Datu & BartscH, 1909, new name
(Text figure 136)
Tablet 1971, 2 syntypes. [3.8 mm]. Brann: plt. 44, fig.
506.
One of the syntypes is juvenile.
226. Odostomia (Chrysallida) ovata, p. 417
=O. (C.) ovata CarPENTER, 1857
(Text figure 137)
Tablet 1968, 3 syntypes. [3.8 mm]. Brann: plt. 44, fig.
503.

227. Odostomia (Chrysallida) ovulum, p. 423 (non Par-
thenia ovulum LE, 1845)
=O. (C.) oonisca DALL & Bartscu, 1909, new name
(Text figure 138)
Tablet 1982, 9 syntypes. [1.4mm]. BRANN: plt. 45, fig.
Bil,
The sculpture is nodose, the basal furrows crenulate.

THE VELIGER

Vol. 10; No. 4

Brann, p. 70, erroneously records the replacement name
as Odostomia benthina.
228. Odostomia (Chrysallida) reigeni, p. 422
=O. (C.) reigeni CARPENTER, 1857
(Text figure 139)
Tablet 1979, holotype. [1.4 mm]. Brann: plt. 44, fig. 509.

229. Odostomia (Chrysallida) rotundata p. 418
=O. (C.) rotundata CarPENTER, 1857
(Text figure 140)
Tablet 1970, 3 syntypes. [2.2mm]. Brann: plt. 44, fig.
505.
230. Odostomia (Chrysallida) telescopium, p. 421
=O. (C.) telescopium CarPENTER, 1857
(Text figure 141)
Tablet 1978, 4 syntypes. [3.2 mm]. Brann: plt. 44, fig.
508.
231. Odostomia (Parthenia) lacunata, p. 414
= Odostomia (Egila) lacunata CARPENTER, 1857
(Text figure 142)
Tablet 1964, 1 syntype. [1 mm]. Brann: plt. 43, fig. 499.
232. Odostomia (Odostomia) tenuis, p. 412
= Odostomia (Evalea) tenuis CARPENTER, 1857
(Text figure 143)
Tablet 1958, 1 syntype. [1.6mm]. Brann: plt. 43, fig.
493.
The columellar plait can be seen only faintly by tilting
the shell.
233. Chemnitzia (Dunkeria) intermedia, p. 435
== Odostomia (Evalina) intermedia (CarPENTER, 1857)
(Text figure 144)
Tablet 2010, 1 syntype. [1.4mm]. Brann: plt. 45, fig.
539)
The sculpture is entirely spiral except for some wrinkles
below the suture.
234. Odostomia (Chrysallida) photis, p. 425
= Odostomia (Haldra) photis CARPENTER, 1857
(Text figure 145)
Tablet 1985, 1 syntype. [1.2 mm]. Brann: plt. 45, fig.
515.
235. Odostomia (Parthenia) quinquecincta, p. 414

= Odostomia (Ividella) quinquecincta CARPENTER,
1857

(Text figure 146)

Tablet 1963, 1 syntype. [1.6mm]. Brann: plt. 43, fig.
498.

Vol. 10; No. 4

THE VELIGER

Page 431

236. Odostomia (?Chrysallida) clausiliformis, p. 426
= Odostomia (Lysacme) clausiliformis CARPENTER,
1857
(Text figure 147)

Tablet 1987, 1 syntype. [3.8mm]. Brann: plt. 45, fig.
Bylo

There is a suggestion of basal cords. The shell is decor-
ticated, with a broken outer lip. The mounted operculum

(if it is one) could hardly have come from so worn a
shell.

237. Odostomia (Odostomia) sublirulata, p. 410
= Odostomia (Menestho) sublirulata CARPENTER, 1857
(Text figure 148)

Tablet 1952, holotype. [2mm]. Brann: plt. 42, fig. 487.
Lirations on the body whorl of the type are faint.

238. Odostomia (Parthenia) ziziphina, p. 416
= Odostomia (Menestho) ziziphina CarPENTER, 1857
(Text figure 149)
Tablet 1967, holotype. [0.7 mm]. Brann: plt. 43, fig. 502.

239. Odostomia (Parthenia) armata, p. 415
= Odostomia (Miralda) armata CarPENTER, 1857
(Text figure 150)

Tablet 1965, 1 syntype. [2.5mm]. BRANN: plt. 43, fig.
500.

The shell cannot be oriented in a normal position for
drawing or photographing without removal from the
mount, which I did not attempt.

240. Odostomia (Parthenia) exarata, p. 415
= Odostomia (Miralda) exarata CARPENTER, 1857
(Text figure 151)

Tablet 1966, 1 hypotype. [2.2 mm]. Brann: plt. 42, fig.
501, holotype [6.3 mm.”’}.

Carpenter’s description was prepared from the speci-
men figured in his drawing. Later he stated (1865 b, p.
274) that he had substituted a much finer shell, which is
the one shown in my figure. Only a glue smear remains
on the slide to show where the original (holotype) had
been, and he did not indicate what had been done with
it. Perhaps it may be the specimen reported by PALMER
(1951, p. 55) at McGill University (Redpath Museum),
or it may be in one of the other institutions holding Car-
penterian material. Evidently his stated dimensions are
in error — length, 6.3, width 0.8mm — for this is at
variance with his words, “‘stout little shell.” I measure
the hypotype as length 2.7 mm, width 1.8mm, which
is more in accord with his diagnosis.

241. Odostomia (Odostomia) mammillata, p. 412
=O. (O.) mammillata CarPENTER, 1857
(Text figure 152)
Tablet 1957, holotype. [1.1 mm]. Brann: plt. 43, fig.
492.

The columellar fold, somewhat obscured by sand
grains, is small but present. Later, CARPENTER (1865 b,
p. 272) thought the form might be Diala paupercula
(C. B. Apams, 1852), but Dart & Bartscu (1909, p.
233) restored it to Odostomia without comment.

242. Odostomia (Parthenia) scalariformis, p. 413
= Odostomia (Salassia) scalariformis CARPENTER, 1857
(Text figure 153)
Tablet 1962, 1 syntype. [2.5mm]. Brann: plt. 42, fig.
AQT:

Dat & BartscH (1909, p. 135) cite the length as 5
mm. Baker, HANNA, & StronG (1928, p. 227), on the
basis of a California Academy specimen, state it as
2.5mm. I measured the British Museum syntype as 2.7
mm, using a vernier caliper, as 3.0mm with the camera
lucida. In any case, the type is well under 5 mm in length.
243. Fossarus (Isapis) maculosa, p. 355

=TIselica maculosa (CARPENTER, 1857)
(Text figure 154)
Tablet 1702, 2 syntypes. [3 mm]. BrRANN: plt. 39, fig.
406.

The apex of the larger syntype is white, flattened, ap-
parently heterostrophic. The smaller syntype is so dete-
riorated as to be unrecognizable.

244. Chemnitzia (Chemnitzia) muricata, p. 428
= Turbonilla (Chemnitzia) muricata (CARPENTER,
1857)
(Text figure 155)
Tablet 1993, 2 syntypes. [2.2 mm]. BRANN: plt. 46, fig.
O22s
245. Chemnitzia (Dunkeria) subangulata, p. 434
== Turbonilla (Bartschella) subangulata (CARPENTER,
1857)
(Text figure 156)

Tablet 2008, 2 syntypes (one juvenile). [2.8mm]. BRANN:
plt. 47, fig. 537.

246. Chemnitzia (Dunkcria) cancellata, p. 435 (non
Orpicny, 1842)
= Turbonilla (Dunkeria) sp.
(Text figure 157)

Tablet 2009, holotype (juvenile). [1 mm]. Brann: plt.
45, fig. 538.

Page 432

Dall and Bartsch failed to catch this homonymy because
they had cited the species under the subgeneric rather
than the generic name. It is doubtful whether the form
can be recognized from so young a type. In any case, the
homonym should not be renamed but rather a new species
described, based on adult material, should growth series
become available that would link Carpenter’s type with
adult specimens in need of a name.

247. Chemnitzia (Dunkeria) paucilirata, p. 434
= Turbonilla (Dunkeria) paucilirata (CARPENTER,
1857)
(Text figure 158)
Tablet 2007, holotype. [2.1 mm]. Brann: plt. 47, fig. 536.
The specimen is, as Carpenter’s drawing shows, broken
and worn. It may, however, be recognizable.

248. ?Eulimella obsoleta, p. 436
= Turbonilla (Ptycheulimella) obsoleta (CARPENTER,
1857)
(Text figure 159)

Tablet 2011, holotype. [1.5 mm]. Brann: plt. 47, fig. 540.

The specimen is worn smooth, the apex gone, the
aperture incomplete. I question whether it is recogniz-
able, although Dall and Bartsch seemed confident as to
its allocation.

249. Chemnitzia flavescens, p. 432
= Turbonilla (Pyrgiscus) flavescens (CARPENTER, 1857)

(Text figure 160)

Tablet 2003, holotype. [2.8 mm]. BRANN: plt. 46, fig. 532.

Spiral threads are minute, the axial ribs sinuous. The
anterior part of the aperture in the type is broken, as
Carpenter’s drawing shows.

250. Chemnitza gracillima, p. 431
= Turbonilla (Pyrgiscus) gracillima (CARPENTER,
1857)
(Text figure 161)

Tablet 2001, 1 syntype. [2.8mm]. Brann: plt. 46, fig.
530.
251. Odostomia (Chrysallida) indentata, p. 425
= Turbonilla (Pyrgiscus) indentata (CARPENTER,
1857)
(Text figure 162)

Tablet 1986, 1 syntype. [4 mm]. BrRANw: plt. 45, fig. 516.

The shell is brown, not white, its interspaces with fine
spiral threads. The allocation of Dall and Bartsch to
Turbonilla seems justified.

252. Chemnitzia tenuilirata, p. 433
= Turbonilla (Pyrgiscus) tenuilirata (CARPENTER,
1857)

(Text figures 163 a-b)

THE VELIGER

Vol. 10; No. 4

Tablet 2005, 2 syntypes. [1.6mm]. Brann: plt. 47, fig.
534.

Dati « BartscH (1909) did not cite this form and
apparently regarded it as a synonym of Turbonilla tenu-
icula (see discussion under the next species). The axial
ribs are prominent, the spirals obsolete on the upper part
of the whorls.

253. Chemnitzia terebralis, p. 432
= Turbonilla (Pyrgiscus) terebralis (CARPENTER, 1857)
(Text figure 164)

Tablet 2004, holotype. [2.4 mm]. Brann: plt. 46, fig. 533.

Da. & BartscH (1909, p. 92) place this in the syno-
nymy of the California species Turbonilla (Pyrgiscus)
tenuicula (GouLp, 1853), a variable form, which they
do not otherwise cite as from Gulf of California localities.
Until this form can be shown definitely to range south
of Baja California, it seems best to regard T: terebralis
and the closely related T: tenuilirata and T: untfasciata
either as separate species or as only subspecifically distinct
from each other. In T: terebralis the intersections of the
fine spiral and distinct axial ribs are well marked, almost
beaded.

254. Chemnitzia unifasciata, p. 433
= Turbonilla (Pyrgiscus) unifasciata (CARPENTER,
1857)
(Text figure 165)

Tablet 2006, holotype. [2.3mm]. Brann: plt. 47, fig.
535.

The outer lip of the holotype is broken, the shell worn
and polished by abrasion. A yellow color band shows
through the shell; spiral sculpture is faint. (See discussion
under the two entries above).

255. Chemnitzia c-b-adamsi, p. 427
= Turbonilla (Strioturbonilla) c-b-adamsti_ (CarPEN-
TER, 1857)
(Text figure 166)

Tablet 1990, 1 syntype. [3.8mm]. Brann: plt. 42, fig.
519.

As with Liotia c-b-adamsu, I am disregarding the
requirement in Article 32 (c) of the International Code
of Zoological Nomenclature and retaining the hyphens,
for otherwise the specific name is unpronounceable.
256. Chemnitzia undata, p. 431

= Turbonilla (Strioturbonilla) undata (CARPENTER,
1857)
(Text figure 167)
Tablet 2002, 1 syntype. [1.6mm]. Brann: plt. 45, fig.
531.

The aperture in this specimen is broken, and the shell

seems too young and worn to be identifiable; the sculp-

Vol. 10; No. 4

ture can be seen, only faintly, by looking through the
shell, which is semi-transparent.

257. Chemnitzia prolongata, p. 429
= Turbonilla (Turbonilla) prolongata (CARPENTER,
1857)
(Text figure 168)
Tablet 1995, 1 syntype. [5.5mm]. Brann: plt. 45, fig.
p24:

Spiral sculpture is faint, especially on the early whorls.

258. Chemanitzia gibbosa, p. 430
BY

Tablet 1996, 2 syntypes, both fragmentary. [6.7 mm].
Brann: plt. 46, fig. 525.

The larger specimen seems to me to be a rissoinid too
worn for determination. The smaller is a columbellid,
perhaps an Aesopus. The apex is wanting, and there is no
sign of a columellar fold. DALL & BartscH (1909, p. 61;
plt. 6, fig. 2) allocated the form to Turbonilla (Pyrgo-
lampros) but indicated that such an identification was
problematical.

ELLOBODAE

259. Melampus olivaceus, p. 178
= Melampus olivaceus CARPENTER, 1857
(Plate 58, Figure 67; Text figure 169)
Tablets 812 - 814, 17 syntypes (7 from tablet 812 photo-
graphed and 1 from tablet 814 drawn). [16 mm].
Authors have had no problem identifying this species.

SIPHONARIIDAE

260. Siphonaria aequilirata, p. 184
= Siphonaria maura maura SowERBY, 1835
(Plate 58, Figure 68)
Tablet 866, holotype. [21.5 mm].
Carpenter seems not to have been aware of the
amount of variability in this species.

261. Siphonaria lecanium palmata, p. 183
=Siphonaria maura palmata CARPENTER, 1857
(Plate 58, Figure 69)
Tablets 836 - 845, 56 syntypes (tablet 840 photographed).
[25 mm].
Authors have identified the form correctly.
262. Nacella peltoides CARPENTER, 1864b, p. 474
= Williamia peltoides (CARPENTER, 1864)
(Text figure 170)
Tablet 944, hypotype. [ — ]. Brann: plt. 21, fig. 262.

THE VELIGER

Page 433

This was cited on page 202 of the Mazatlan Catalogue
as “Nacella, sp. ind.” Later the species was described on
the basis of better material from Cape San Lucas, Baja
California. PALMER (1958, p. 259; plt. 25, figs. 15 - 16;
and 1963, p. 367) has discussed and figured type
material.

AMPHINEURA

The following notes are a summary of material as it
is now available. I have turned to chiton specialists for
advice on placement in modern terms, notably to Mr.
Allyn G. Smith and Mr. Spencer Thorpe. Mr. Thorpe,
after studying my photographs and notes, decided to con-
tribute a separate commentary, to be submitted for pub-
lication at a future date.

263. Lophyrus striatosquamosus, p. 192
(Plate 59, Figure 91)
Tablet 897, holotype. [4.4 mm]. Brann: plt. 18, fig. 250.

The surface is finely punctate to scaly; the girdle is not
preserved.

264. Tonicia forbesii, p. 193
= Tonicia forbesii CARPENTER, 1857
(Plate 59, Figure 104)
Tablet 898, 1 syntype (here chosen as lectotype). [20
mm].

Authors seem to have recognized this form correctly;
it was figured by Pirssry (1893, Manual of Conch., vol.
14, p. 196; plt. 45, figs. 69-72) under the preoccupied
name of “T: crenulata (SowERBy, 1832).”

265. Acanthochites arragonites, p. 198
= Acanthochitona arragonites (CARPENTER, 1857)
(Plate 59, Figure 92)
Tablet 907, 1 syntype, 4 odd valves. [4.1 mm]. Brann:
pit. 21, fig. 258.

The articulated specimen (here chosen as lectotype)
shows the bristle tufts well, but the drying girdle has
arched it so much that it does not photograph well. There
is a color pattern of pink and brown spots on white, and
the margins of the girdle look scalloped. Patmer (1963,
pit. 68, fig. 1) has figured a specimen from Cape San
Lucas labelled by Carpenter.

266. Lepidopleurus macandrei, p. 195
=TIschnochiton macandreae (CARPENTER, 1857)
(Plate 59, Figure 93)

Tablet 904, holotype. [3.3 mm]. Brann: plt. 20, fig. 255.
The spelling of the specific name was emended to

Page 434

macandreae by Carpenter on page 550. The specimen
was donated by Mr. Thomas Nuttall, and the collecting
locality is not positively Mazatlan. Carpenter assumed
it to be specifically identical to one from Mazatlan that
was smashed during examination. His drawing shows 10
slits in the head valve, but I thought I could count 12.
The girdle is wide and laps over the valves, almost con-
cealing one of them.

267. Lepidopleurus beanii, p. 197
=Chaetopleura beanii (CARPENTER, 1857)
(Plate 59, Figure 94)
Tablet 905, 1 syntype (here chosen as lectotype), 2
valves. [5.9mm]. Brann: plt. 20, fig. 256.

There are a few girdle setae on the entire specimen,
rather long and curved. The mucro is sunken. Possibly
the 2 odd valves are not specifically identical to the entire
specimen; both are smoother and differently sculptured.

268. Lepidopleurus bullatus, p. 195
= Chaetopleura bullata (CARPENTER, 1857)
(Plate 59, Figure 95)
Tablet 902, 1 syntype, 2 odd valves. [4.4mm]. Brann:
pit. 20, fig. 254.

An articulated specimen, here chosen as lectotype, has
parts of 2 valves broken out, and the 2 odd valves mounted
separately may not be the same form. Girdle scales are
minute.

269. Lepidopleurus bullatus calciferus, p. 195
=Chaetopleura calcifera (CARPENTER, 1857)
(Plate 59, Figure 96)
Tablet 903, holotype. BRANN: plt. 20, fig. 254-b.

The type, somewhat contracted, is about 3 mm long.
The girdle is not well preserved. The form is doubtfully
distinct.

270. Chiton flavescens, p. 198
?==Chaetopleura flavescens (CARPENTER, 1857)
(Plate 59, Figure 103)
Tablet 906, 3 syntypes, 2 odd valves. [4.1 mm]. BRANN:
joke, Al, tee, 2d

The largest specimen shows 9 to 10 slits on the head
valve; a smaller is attached within a dead Crepidula,
and there is one subadult, badly contracted by the drying
girdle.

271. Lepidopleurus clathratus, p. 195
?== Lepidozona subclathrata (Pitssry, 1892)

(Plate 59, Figure 97)
Tablet 901, holotype. [4.4 mm]. Brann: plt. 20, fig. 253.
As Carpenter’s drawing shows, the girdle is wide, with

THE VELIGER

Vol. 10; No. 4

fine scales; ribs are beaded. If this is a Lepidozona, as
seems probable, the name must fall as a secondary homo-
nym of L. clathrata (ReEve, 1847) ; Pilsbry has provided
a replacement name.

ADDENDUM

Carpenter described a number of other Panamic prov-
ince mollusks in addition to those in the Reigen collection.
Most have been discussed already by Parmer (1958,
1963), but as some of the types that were thought to be
lost now have come to light, I can add a few more
observations on these as well as on some Californian
material.

272. Diplodonta subquadrata CARPENTER, 1856
_ ==Diplodonta subquadrata CARPENTER, 1856
(Plate 56, Figure 26)

Proc. Zool. Soc. London for 1855, p. 230.

This was described from Mazatlan but was not in the
Reigen material. Authors seem to have identified the
species correctly in spite of lack of illustration. PALMER
(1958, p. 85) has discussed the form briefly. There are
two syntypes in the British Museum collection, registry
number 196384.

The generic name Diplodonta Bronn, 1831 has been
rejected by many authors because Taras Risso, 1826 has
priority, but according to CHAvAN, the type species of
the latter is indeterminate.

273. Venus cortezi CARPENTER, 1864
—=Chione (Chionista) cortezi (CARPENTER, 1864)
(Plate 55, Figure 19)

Report British Assoc. Adv. Sci. for 1863, p. 570. Type
locality, Guaymas, Mexico.

Carpenter’s proposal of the name comes uncomfort-
ably close to being a nomen nudum or a name proposed
in synonymy, but as it had been adopted prior to 1961
and seems to be useful (KEEN, 1958, p. 142; fig. 322),
I would argue for its acceptance.

274. Thracia squamosa CARPENTER, 1856
= Thracia squamosa CARPENTER, 1856
(Plate 56, Figure 32)

Proc. Zool. Soc. London for 1855, p. 229.

Parmer (1963, p. 320) has reprinted the original de-
scription. Carpenter’s holotype, from the Cuming collec-
tion, type locality Mazatlan, was figured by Reeve in
1859 (Conch. Icon., vol. 14, plt. 3, fig. 16). I reproduced

Vol. 10; No. 4

this figure (KEEN, 1958, fig. 590). Two valves are in the
Type Collection of the British Museum, registry no.
1966570; the right valve is somewhat broken.

275. Litiopa divisa CARPENTER, 1856
?—= Litiopa melanostoma Rane, 1829
(Plate 58, Figure 58; Text figure 171)

Proc. Zool. Soc. London for 1855, p. 234. “Cape San
Francisco.” [3.3 mm].

PatmerR (1963, p. 339) has reprinted the original de-
scription. She points out that Carpenter had misgivings
on the locality, which is apparently eastern Pacific but
could have been in California or in Ecuador. The form
is doubtfully distinct from Litiopa melanostoma, known
in both the Atlantic and Pacific as a pelagic gastropod.
There are now only 3 of the original 6 specimens on the
mount. The apical 2 whorls are white, the next with fine
axial riblets, the body whorl with faint spiral threads,
and there is some variation in color and outline. The best
specimen is here illustrated.

276. Galerus ?sinensis, var. fuscus CARPENTER, 1856
= Calyptraea sp.
(Plate 57, Figure 53)

Proc. Zool. Soc. London for 1855, p. 233.

Carpenter’s original description is brief: “G. sinensis,
t. rufo-fusca, huc et illuc lineis declivis instructa.” He
compares this to the British Calyptraea sinensis but be-
cause of the slightly darker color suggests the subspecific
name fuscus. The holotype, which was in the Cuming
collection marked “Gulf of California,” might not have
come from there. The registry number in the British
Museum Type Collection is 1966636. If the shell actually
came from West Mexican waters, one would have to con-
sider the name a synonym of Calyptraea mamillaris
Broperip, 1834, for there seem no clearcut points of
distinction other than the brown coloration that suggests
it is indeed C. sinensis.

277. Erato ?maugeriae panamensis CARPENTER, 1856
= Erato (Hespererato) panamensis CaRPENTER, 1856
(Plate 58, Figures 66 a-b)

Proc. Zool. Soc. London for 1856, p. 162.

PatMER (1963, p. 345) has reprinted the original de-
scription. Three specimens that seem to me unquestion-
ably to be the syntypes are in the British Museum col-
lection. They do not carry a Carpenter label, but they
are from Panama and have on the back of the mount,
in the handwriting of E.A.Smith, the page reference
to the Carpenter description. The specimen figured here

THE VELIGER

Page 435

was illustrated by SowerBy, 1859 (Thesaurus Conchy]l.,
vol. 3, plt. 219, fig. 3) as “Erato sulcifera, var.” The
form has also been labelled E. laevis, var., and E. mau-
geriae. It seems to be distinct from E. maugeriae SowErR-
BY, 1832, of the West Indies, in having a lower spire and
more inflated body whorl. Although it has not again
been recorded in the West Coast literature, the species
may be valid. It is close to E. columbella Menke, 1847
but more uniformly colored, the outer lip not white. As
I have pointed out recently (KEEN, 1966, Occ. Papers,
Calif. Acad. Sci., no. 59, p. 23), this is probably a senior
synonym of E. marginata Morcu, 1860, which, too, was
compared to FE. maugeriae.

278. Sistrum (?ochrostoma, var.) rufonotatum CaRrPEN-
TER, 1864
= Morula rufonotata (CARPENTER, 1864)
(Plate 59, Figures 81 a-c)

Ann. Mag. Nat. Hist., ser. 3, vol. 14, p. 48. Type
locality, Cape San Lucas, Baja California.

PaLMER (1963, p. 347) has reprinted the original de-
scription. Her search for the holotype at the U. S.
National Museum was unsuccessful. There are, in the
British Museum’s collection, 3 specimens on a glass mount
labelled in Carpenter’s handwriting, part of some mate-
rial that had been sent by the Smithsonian Institution
during the 1860’s on an exchange arrangement. These
are therefore syntypes; their registry number is 1967576.
The species seems to be distinct, characterized by a row
of reddish dots below the peripheral cords. Dr. Donald
Shasky has collected similar specimens on the Tres Marias
Islands and the adjacent mainland of West Mexico.

279. Pisania elata CARPENTER, 1864
= Fusinus (Aptyxis) cinereus (REEvE, 1847)
(Plate 58, Figures 65 a-b)

Ann. Mag. Nat. Hist., ser. 3, vol. 14, p. 49.

The type specimen should be at the U.S. National
Museum but has not been detected. PALMER (1963, p.
354) reprinted Carpenter’s description and figured a
badly encrusted specimen from his collection at McGill
University. In the British Museum there is one small
specimen from Cape San Lucas, received from the Smith-
sonian Institution, that may well be a syntype. There are
4 lots labelled “Pisania elata CARPENTER,” all from the
Cuming collection, either without locality or merely as
from “California.” All these specimens are good Fusinus
cinereus; 2 are figured here. Perhaps the missing type
may still be at the U.S. National Museum under a
changed label.

Page 436

280: Olivella glandinaria CarPENTER, 1857, ex Nuttall
MS
= Olivella biplicata (SowrErBy, 1825)
(Plate 59, Figures 89 a-b)

Proc. Zool. Soc. London for 1856, p. 227. Type locality,
California.

There are 2 specimens in the Nuttall collection at the
British Museum with a Carpenter label. Although he was
apparently aware that the form was not distinct, he made
a gesture of tribute for Nuttall’s work by salvaging in
part a name that Nuttall had chosen, Glandinaria cali-
fornica. The types are well within the range of variation
of Sowerby’s species; registry number, 1861.5.20.54.

281. Drillia punctatostriata CARPENTER, 1856
= Crassispira punctatostriata (CARPENTER, 1856)
(Plate 58, Figure 75)

Proc. Zool. Soc. London for 1856, p. 164. Type locality,
Panama.

PatMeER (1963, p. 361; plt. 68, figs. 5, 6) has reprinted
the original description and has given a figure of the
holotype in the British Museum. I am adding a figure
that is larger and shows more details of sculpture. There

THE VELIGER

Vol. 10; No. 4

is a typographical error in PaLMEr’s plate explanation
that makes the shell seem minute — “x 7” should read,
x28

This species unfortunately was omitted from my book
(KEEN, 1958), for the species was at that time unfigured
and its status seemed uncertain. Now it proves to be so
similar in form to Crassispira solitaria PrtsBry & Lowe,
1932, from Mazatlan, that I see no means of separation.
It may be, therefore, that the range of C’. punctatostriata
should be considered as the southern part of the Gulf of
California to Panama.

282. Melampus bridges CARPENTER, 1856
= Tralia bridgesu (CARPENTER, 1856)

Proc. Zool. Soc. London, pt. 24, p. 161. Type locality,
Panama.

Parmer (1963, p. 366; plt. 67, figs. 14, 15) has
reprinted the original description and figured a speci-
men from the British Museum collection as “holotype,”
registry no. 196312. As Carpenter states there were 3
specimens in the lot, this must be a syntype, which is
here selected as lectotype. Under modern allocation the
form is not a Melampus but a Tralua.

INDEX

Numbers refer to paragraphs, not to pagination or to CARPENTER Catalogue.

A

abnormale, Caecum 142
Acanthochites, 265

Aclis, 84, 123

aculeata, Crepidula 180
acuta, Nassa, 193
aequilirata, Pisania. 192
aequilirata, Siphonaria, 260
Alaba, 122, 165-171
alabastrites, Alaba, 165
alabastrum, Cardium, 41
alba, Fissurella, 63

albida, Bivonia, 154
albocincta, Terebra, 210
alboliratum, Cerithium, 157
albonodosa, Anachis, 183
albonodosa, Cerithiopsis, 158
albonodosa, Drillia, 201
albovallosa, Drillia, 202
alderi, Jeffreysia, 125
Alvania, 121, 172-173
amplectans, Globulus, 111
amplectans, Teinostoma, 110
Anachis, 183-188
angulatus, Fossarus, 177
annae, Dosinia, 44

annulata, Vitrinella, 106
apertus, Fusus, 191

Arca, 2

aristata, Lithophaga, 6

armata, Odostomia, 239
arragonites, Acanthochites, 265
aterrima, Crassispira, 205
aurea, Clathurella, 209
aureocincta, Olivella, 198

B

baccata, Mitrella, 189, 190
bartschi, Anachis, 184
beanii, Lepidopleurus, 267
benthina, Odostomia, 225
bifasciata, Jeffreysia, 124
bifilata, Vitrinella, 117
bifrons, Arca, 2

bifrontia, Vitrinella, 93
biliratus, Cantharus, 191
bilobata, Crepidula, 180
biplicata, Olivella, 280
Bivonia, 152-154

bridgesti, Melampus, 282
Bulla, 215

bullatus, Lepidopleurus, 268

C
Caecum, 129-150

caelatus, Donax, 51

calcifer, Spondylus, 14
calciferus, Lepidopleurus, 269
calyculatus, Lithophagus, 8
cancellata, Chemnitzia, 246
Cardium, 19, 41, 42
carinata, Globulus, 107
carinata, Liotia, 71

carinata, Tornatina, 217
carinatus, Donax, 52
carinulata, Vitrinella, 108
carpenteri, Epitonium, 85
c-b-adamsti, Chemnitzia, 255
c-b-adamsii, Liotia, 73
cerea, Cerithiopsis, 158
Cerithidea, 174

Cerithiopsis, 158-164
Cerithium, 156-157
cerithoidea, Drillia, 203
cervinetta, Columbella, 189
chalcedonica, Montacuta, 32
Chama, 39-40

Chemnitzia, 233, 244-247, 249-

250, 252-258

Chiton, 270

cincta, Vitrinella, 102
cinereus, Fusinus, 196, 279
Circe, 22, 23

Cirsotrema, 87

clathratum, Caecum, 128
clathratus, Lepidopleurus, 271
Clathurella, 209
clausiliformis, Odostomia, 236
claviculata, Placunanomia, 17
Clementia, 45

clementinum, Lepton, 29
coarctata, Crenella, 10
Columbella, 189, 190
compactum, Caecum, 130
compta, Phasianella, 76
conchaphila, Ostrea, 12
conica, Alaba, 166

contorta, Bivonia, 152
contorta, Ianthina, 89
conradt, Penitella, 59
convexa, Cerithiopsis, 163
convexa, Odostomia, 221
Corbula, 58

coronata, Vitrinella, 94
corrugatum, Dentalium, 61
corrugulatum, Caecum, 150

Vol. 10; No. 4

cortezi, Venus, 273
crebristriata, Nassa, 194
Crenella, 10

Crepidula, 180

cryptophila, Vanicoro, 100
culminatus, Donax, 52
Cycladella, 21
cymbiformis, Haminea, 214
Cyrena, 20

D

decollata, Ianthina, 90
decussata, Cerithiopsis, 164
decussata, Vitrinella, 103
Dentalium, 60-62
dextroversum, Caecum, 146
diminuta, Anachis, 187
dionaeum, Lepton, 30
Diplodonta, 28
discrepans, Diplodonta, 28
divisa, Litiopa, 275
donacilla, Tellina, 53
Donax, 51, 52

Dosinia, 44

Drillia, 201-206

E

effusa, Odostomia, 222
elata, Pisania, 279
elegans, Cantharus, 192
elliptica, Montacuta, 33
elongatum, Caecum, 130
Ethalia, 107, 109, 111-113
Eulimella, 248

exarata, Bulla, 215
exarata, Odostomia, 240
exarata, Rupellaria, 47
excavata, Lucina, 24
excurvata, Alvania, 173

F

farcimen, Caecum, 132
fasciata, Odostomia, 223
Fissurella, 63-65
flavescens, Chemnitzia, 249
flavescens, Chiton, 270
forbesii, Tonicia, 264
fornicata, Chama, 39
Fossarus, 177, 178, 243
fragilis, Sphaenia, 57
funiculata, Scalaria, 87
fuscus, Galerus, 276
fusiformis, Aclis, 123
Fusus, 191, 196

G

gaskoini, Anachis 184
gemmata, Fissurella, 63
gibbosa, Chemnitzia, 258
glabriforme, Caecum, 147
glandinaria, Olivella, 280
Globulus, 111-113, 115, 116

THE VELIGER

globulus, Omphalius, 69
Gouldia, 18

gracilior, Lithophagus, 6
gracillima, Chemnitzia, 250
gracillima, Clementia, 45
granifera, Trigoniocardia, 41
guayanensis, Mytella, 5

H

Haminoea, 214

hanleyi, Drillia, 204
heptagonum, Caecum, 137
hindsii, Terebra, 211
Hipponix, 175, 176
humilis, Trigona, 43

I

Tanthina, 88-90
imbricatus, Siphonium, 151
indentata, Bivonia, 153
indentata, Muricidea, 182
indentata, Odostomia, 251
insculptum, Caecum, 143
intercalata, Martesia, 59
intermedia, Chemnitzia, 233
intertincta, Oliva, 197
intertincta, Terebra, 213
involuta, Mucronalia, 83
Isognomon, 11

J

janus, Isognomon, 11
Janthina, 88-90

janthina, Janthina, 88, 89
Jeffreysia, 124-126

L

lacunata, Odostomia, 231
laguncula, Alaba, 167
lamellata, Odostomia, 218
lamellata, Tellina, 56a
Lasea, 37-38

Leiostraca, 79-82
Lepidopleurus, 266-269, 271
Lepton, 29-31

linearis, Leiostraca, 79
linguafelis, Rupellaria, 48, 49
Liotia, 71-73

lirata, Rissoa, 128
liratocinctum, Caecum, 138
liratum, Dentalium, 60
lirulata, Globulus, 112
lirulata, Vitrinella, 95
Lithophaga, 6

Littorina, 91

lividus, Murex, 181
Lophyrus, 293

Lucina, 24-27

lucinoides, Cardium, 19
Lunatia, 77

lurida, Barbatia, 1
luteostoma, Nassarius, 195

M

macandreae, Lepidopleurus, 266
macandreae, Trochus, 68
macandret, Lepidopleurus, 266
macrophragma, Petaloconchus, 155
maculosa, Fossarus, 243
maculosa, Iselica, 243
maculosum, Cerithium, 156
mammillata, Odostomia, 241
Mangelia, 207, 208
margarita, Circe, 22
margaritula, Marginella, 199
Marginella, 199, 200
Martesia, 59

maura, Siphonaria, 261
mazatlanica, Cerithidea, 174
mazatlanica, Lucina, 26
mazatlanica, Pyramidella, 218
mazatlanica, Rimula, 66
mazatlanica, Tricolia, 74
mediolaeve, Cerithium, 156
Melampus, 259

melanostoma, Litiopa, 275
melchersi, Drillia, 205
mexicana, Chama, 39, 40
mexicana, Ctena, 27
Modiola, 5

Modiolus, 5

monile, Vitrinella, 104
monilifera, Drillia, 206
monilifera, Vitrinella, 105
Montacuta, 32-35
Mucronalia, 83

multiformis, Mytilus, 3
Murex, 181

muricata, Chemnitzia, 244
Muricidea, 182

mutabilis, Modiola, 5

mutans, Alaba, 122

Mytilus, 3

N

Nacella, 262

Naranio, 49

Nassa, 193-195

nasuta, Corbula, 58
naticoides, Vitrinella, 118
navicelloides, Scutellina, 78
nigrocincta, Fissurella, 64
nigrofusca, Anachis, 185
nodosa, Odostomia, 224
nodosa, Vitrinella, 96
nodulifera, Nassa, 195

O

oblonga, Lasea, 38
oblonga, Odostomia, 225
obsoleta, Columbella, 190
obsoleta, Eulimella, 298
obtusa, Montacuta, 34
obtusum, Caecum, 144

Page 437

Odostomia, 218-232, 234-242, 251
Oliva, 197

olivacea, Cyrena, 20
olivaceus, Melampus, 259
Olivella, 198

Omphalius, 69, 70
oonisca, Odostomia, 227
orbis, Vitrinella, 92
ornata, Vitrinella, 97
osculans, Phenacolepas, 78
Ostrea, 12

ovata, Odostomia, 226
ovulum, Odostomia, 227

P

pachyderma, Anachis, 186
pagodus, Nassarius, 193
pallidula, Globulus, 113
palliopunctatus, Mytilus, 4
palmata, Siphonaria, 261
palmula, Ostrea, 13
panamense, Trachycardium, 42
panamensis, Erato, 277
papyracea, Cycladella, 21
paucilirata, Chemnitzia, 247
pectinata, Lucina, 27
peltoides, Nacella, 262
penicillata, Plicatula, 15
perforata, Phasianella, 74
pernoides, Placunanomia, 16
peruviana, Anomia, 17
Petaloconchus, 155
Phasianella, 74-76

philip pi, Litorina, 91
phoebe, Pegmapex, 28
photis, Odostomia, 234
Pisania, 192

planatus, Hipponyx, 176
planospirata, Vitrinella, 101
Plicatula, 15

polita, Marginella, 200
politus, Solecurtus, 53
pretiosum, Dentalium, 62
producta, Leiostraca, 80
prolongata, Chemnitzia, 257
prolongata, Janthina, 90
prolongata, Lucina, 25
punctatostriata, Drillia, 281
punctatostriatus, Donax, 51
pupiformis, Cerithiopsis, 160
pustulosa, Corbula, 58
pyricallosa, Globulus, 109
Pythina, 36

Q

quadratum, Caecum, 133
quinquecincta, Odostomia, 235

R

raricostata, Scalaria, 85
regularis, Tellina, 56b
reigent, Odostomia, 228

Page 438

THE VELIGER

Vol. 10; No. 4

retexta, Leiostraca, 81
reversum, Caecum, 148
Rimula, 66

rimuloides, Scissurella, 67
Rissoa, 128

Rissoina, 127

rotundata, Odostomia, 229
rotundatum, Cardium, 42
rufocinerea, Terebra, 212
rufonotatum, Sistrum, 278
rufotincta, Anachis, 187
rufotinctus, Omphalius, 70
rugosa, Tegula, 70
Rupellaria, 47-48

S
Scalaria, 85-86

scalarina, Anachis, 186
scalariformis, Odostomia, 242
scalata, Alaba, 168
Scissurella, 67

scobina, Naranio, 49
Scutellina, 78

Semelina, 55

semilaeve, Caecum, 134
semiobliterata, Dosinia, 44
semipolitum, Dentalium, 60, 61
serrata, Anachis, 188
serratus, Hipponyx, 175
Siphonaria, 260-261
Siphonium, 151

Smaragdinella, 216
Solecurtus, 53-54

sorex, Cerithiopsis, 161
spatiosus, Leiosolenus, 9
Sphaenia, 57

Sphenia, 57

Spondylus, 14

spongiosa, Fissurella, 65
squamosa, Tapes, 46
squamosa, Thracia, 274
striatosquamosus, Lophyrus, 263
striulata, Ianthina, 88
striulata, Liotia, 72
striulata, Phasianella, 75
subangulata, Chemnitzia, 245
subangulata, Mangelia, 208
subconicum, Caecum, 139
subimpressum, Caecum, 135
sublaevis, Pythina, 36
sublirulata, Odostomia, 237
subnodosa, Terebra, 213
subobsoletum, Caecum, 140
subquadrata, Diplodonta, 272
subquadrata, Montacuta, 55
subquadrata, Vitrinella, 119
subspirale, Caecum, 145
substriatum, Teinostoma, 144
subsulcata, Odostomia, 219
subtrigona, Circe, 23
sulcata, Mangelia, 207
sulcatus, Globulus, 115

LITERATURE CITED

Baker, FREDERICK, G DaLtas 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

1930.

(29 June 1928)

Some Rissoid Mollusca from the Gulf of California.

Proc. Calif. Acad. Sci., ser. 4; 19 (4): 23 - 40; plt. 1; 4 text figs.

1938.

(15 July 1930)

Some Mollusca of the families Cerithiopsidae, Ceri-

thiidae and Cyclostrematidae from the Gulf of California and

adjacent waters.
217 - 244; plts. 17 - 23
BARTSCH, PAUL
1910.

Proc. Calif. Acad. Sci., ser. 4; 23 (15):

(24 May 1938)

The West American mollusks of the genus Alaba.

Proc. U.S. Nat. Mus. 39 (1781): 153-156; text figs. 1-4

(25 October 1910)

191la. The Recent and fossil mollusks of the genus Alabina from

the west coast of America.
409 - 418; plts. 61, 62

Proc. U. S. Nat. Mus. 39 (1790) :

(13 January 1911)

1911b. The Recent and fossil mollusks of the genus Cerithi-

opsis from the west coast of America.

Mus. 40 (1823) : 327 - 367; plts. 36 - 41

1911c.

from the west coast of America.
(1826): 383 - 414; plts. 51-58

Proc. U.S. Nat.
(8 May 1911)

The Recent and fossil mollusks of the genus Bittium

Proc. U.S. Nat. Mus. 40
(12 May 1911)

supralirata, Alaba, 169

suprastriata, Scalaria, 86

T

tantilla, Transennella, 43
Tapes, 46

Teinostoma, 110, 114
telescopium, Odostomia, 230
Tellina, 50, 55, 56
tenuilirata, Chemnitzia, 252
tenuilirata, Lunatia, 77
tenuiliratum, Caecum, 141
tenuis, Odostomia, 232
tenuisculpta, Vitrinella, 98
Terebra, 210-213

terebralis, Alaba, 170
terebralis, Chemnitzia, 253
teres, Caecum, 149
thecaphora, Smaragdinella, 216
Tonicia, 264

Tornatina, 217

Trigona, 43

trigonalis, Lasea, 37
trigonare, Sphaerium, 21
trigonata, Vitrinella, 99
Trochita, 179

Trochus, 68

tuberculoides, Cerithiopsis, 162
tuberosus, Fossarus, 178
tumens, Aclis, 84

tumens, Fusus, 196

BartscH, PAUL

tumens, Globulus, 116
tumens, Jeffreysia, 125
tumida, Alvania, 121
tumidior, Lithophagus, 7

U

umbonatum, Lepton, 31
undata, Chemnitzia, 256
undatum, Caecum, 136
unifasciata, Chemnitzia, 254

Vv

vallata, Odostomia, 220
Vanicoro, 100
varians, Gouldia, 18
variegata, Terebra, 210, 211, 212
ventricosa, Trochita, 179
versicolor, Nassarius, 194
vespertilio, Byssoarca, 1
violacea, Alaba, 171
violascens, Solecurtus, 54
Vitrinella, 92-99, 101-106, 108,
117-120

Ww
woodwardii, Rissoina, 127

Y
yod, Leiostraca, 82

Z
ziziphina, Odostomia, 238

1911d. The Recent and fossil mollusks of the genus Alvania

from the west coast of America.
(1863) : 333 - 362; plts. 29 - 32

1915.

1917.

Proc. U.S. Nat. Mus. 41
(15 November 1911)

The Recent and fossil mollusks of the genus Rissoina
from the west coast of America.
49 (2094): 33-62; plts. 28-33

A monograph of West American melanellid mollusks.

Proc. U.S. Nat. Mus.
(24 July 1915)

Proc. U.S. Nat. Mus. 53 (2207): 295-356; plts. 34-49

1920.

(13 August 1917)

The West American mollusks of the families Rissoellidae
and Synceratidae, and the rissoid genus Barleeia.

Proc. U.S.

Nat. Mus. 58 (2331): 159-176; plts. 12, 13

BERRY, SAMUEL STILLMAN
1960.

Brann, Doris C.
1966.

Ithaca, N. Y.
CAMPBELL, G. Bruce
1964.
Gastropoda).

(9 November 1920)

Notices of new eastern Pacific Mollusca — IV. Leafl.
in Malacol. 1 (15): 115-122

(31 December 1960)

Illustrations to “Catalogue of the Collection of Mazat-
lan Shells” by Philip P Carpenter.
111 pp.; 60 plts.

Paleont. Res. Instit.
(1 April 1966)

New terebrid species from the eastern Pacific (Mollusca:
The Veliger 6 (3): 132-138; plt. 1?

(1 January 1964)

Vol. 10; No. 4 THE VELIGER Page 439
CarRPENTER, Puirip PEARSALL Keen, A. Myra
1856a. Descriptions of (supposed) new species and varieties of 1958. Sea shells of tropical West America; marine mollusks

shells, from the Californian and West Mexican coasts, princi-
pally in the collection of H. Cuming. Proc. Zool. Soc.
London, prt. 23: 228 - 235 (5 and 23 February 1856)
1856b. Notes on the species of Hipponyx inhabiting the Amer-
ican coasts, with descriptions of new species. Proc. Zool.
Soc. London, part 24 (for 1856): 3-5 (16 June 1856)
1856c. Description of new species of shells collected by Mr. T.
Bridges in the Bay of Panama and its vicinity, in the collection
of Hugh Cuming, Esq. Proc. Zool. Soc. London, part 24
(for 1856): 159 - 166 (11 November 1856)
1857a. First steps toward a monograph of the Recent species
of Petaloconchus, a genus of Vermetidae. Proc. Zool.
Soc. London, part 24 (for 1856): 313-317; figs. 1-8
: (10 March 1857)
1857b. Catalogue of the collection of Mazatlan shells in the
British Museum collected by Frederick Reigen. London:
Brit. Mus.: i-iv+ix-xvi+552 pp. (with preface by J. E.
Gray) (post-June 1857). [Warrington edition, with author’s
preface, pp. v- viii, published simultaneously for distribution
with duplicate collections.]
1864a. Review of Prof. C.B. Adams “Catalogue of the shells
of Panama,” from the type specimens. Proc. Zool. Soc.
London for 1863: 339 - 369 (April 1864)
1864b. Diagnoses of new forms of mollusks collected at Cape
St. Lucas, Lower California, by Mr. J. Xantus. Ann. Mag.
Nat. Hist., ser. 3; 13: 311-315 (April 1864) ; 474-479 (June
1864) ; 14: 45-49 (July 1864).
1864c. Supplementary report on the present state of our know-
ledge with regard to the Mollusca of the West Coast of North
America. Rep. Brit. Assoc. Adv. Sci. for 1863, pp. 517
to 686, publ. August 1864
1865a. Diagnoses des mollusques nouveaux provenant de Cali-
fornie et faisant partie du Musée de I’Institution Smithsonienne.
Journ. de Conchyl]. 13[(3) 5](2): 129-149 (4 April 1865)
1865b. Diagnoses of new species and a new genus of mollusks,
from the Reigen Mazatlan Collection; with an account of
additional specimens presented to the British Museum. Proc.
Zool. Soc. London for 1865: 268 - 272 (June 1865)
1872. The mollusks of western North America, embracing
the second report made to the British Association on this
subject, with other papers; reprinted by permission, with a
general index. Smithson. Misc. Coll. no. 252: xii+325+
index, 121 pp.
DALL, WILLIAM HEALEY, & PAUL BARTSCH
1909. A monograph of the West American pyramidellid mol-
lusks. U.S. Nat. Mus. Bull. 68: 1 - 258; 30 plts.
(13 December 1909)
Grant, Utysses S., IV. « Hoyt Ropney Gate
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)
Hertvein, Leo Grorce & ARCHIBALD McC.iure StRoNnG
1955. | Marine mollusks collected during the “Askoy” Expedi-
tion to Panama, Colombia, and Ecuador in 1941. Bull.
Amer. Mus. Nat. Hist. 107 [article 2]: 159-318; plts. 1-3
(28 November 1955)

from Lower California to Colombia.
colored plts.; 1700 text figs. Stanford Univ. Press, Stan-
ford. Calif. (5 December 1958)

1961. A proposed reclassification of the gastropod family
Vermetidae. Bull. Brit. Mus. (Nat. Hist.) Zool. 7 (3):
183 - 213; plts. 53-54; 33 text figs. (February 1961)

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 illustrated key. Stanford Univ. Press; 126 pp.; text
figs. not separately numbered (14 February 1963)

Otsson, 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)

i-xi+624 pp.; 10

PaLMER, KATHERINE VAN WINKLE
1951. Catalog of the first duplicate series of the Reigen Collec-
tion of Mazatlan shells in the State Museum at Albany, New
York. N. Y. State Mus. Bull. 342:1-79;1plt. (Jan. 1951)
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)
1963. Type specimens of marine mollusca described by P. P.
Carpenter from the west coast of Mexico and Panama.
Bull. Amer. Paleo. 46 (211): 285-400; plts. 58 - 70
(22 October 1963)
Pirspry, Henry Aucustus & AxEL ADOLF OLSSON
1945; 1952. Vitrinellidae and similar gastropods of the Panamic
province. Acad. Nat. Sci. Philadelphia; I: 97: 249 - 278;
plts. 22-30 (27 December 1945); II: 104: 35-99; plts.
2-13 (10 September 1952)
REEvE, LovELL AUGUSTUS
1843 to 1878. Conchologia Iconica: or, illustrations of the shells
of molluscous animals. London, vols. 1 - 20, continued by
G. B. Sowersy 2%, beginning with monograph on Pyramidella
in vol. 15, 1865
Soot-RYEN, TRON
1955. A report on the family Mytilidae (Pelecypoda). Allan
Hancock Pacific Expeditions, vol. 20, no. 1 (Univ. S. Calif.
Press, Los Angeles), 175 pp., 10 plts., 78 text figs.
(10 November 1955)
STRONG, ARCHIBALD McCLurE
1928. West American Mollusca of the genus Phasianella.
Proc. Calif. Acad. Sci., ser 4, 17 (4): 187-203; plt. 10
(22 June 1928)
Stronc, ArcHr1BALD McC.iure « Leo Grorcre 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)

Page 440

THE VELIGER

Vol. 10; No. 4

Effect of Feeding by Armina caltfornica

on the Bioluminescence of Renilla koellikeri

BY

HANS BERTSCH

Franciscan Theological Seminary, Old Mission, Santa Barbara, California 93105

THE BIOLUMINESCENCE of the sea pansy (Renilla koelli-
keri PFEFFER, 1886) has been extensively studied in the
past two decades (Harvey, 1952; JoHNson, 1955; Cor-
MIER, 1961; etc.). Most of the published information
treats of chemical analyses of the bioluminescent sub-
stance produced by Renilla when stimulated. Observa-
tions on the behavior of the living animal were performed
to supplement the chemical information, and indicated
that Renilla exhibits a general pattern of yellow-green
bioluminescence that is confined to the rachis, but is able
to be elicited by stimulating any part of the animal. The
pattern varies between a local glow, a wave passing
across the entire rachis, or a general throb, depending on
the intensity of the stimulation. Under the control of
the nerve net, the reaction shows an apparent pattern of
facilitation, with increased stimulation causing increased
bioluminescence. Waves simultaneously elicited from two
different parts of the Renilla will dissipate when the
nerve impulses meet, rather than continuing across the
entire rachis.

The slime that bioluminesces is apparently produced
by the animal even when the response is inhibited by an
external light source. Such a light source will inhibit
that part of the rachis that is exposed to it without
affecting the other side of the rachis, indicating a factor
other than nerve net conduction in the production of
the bioluminous response. Recovery of inhibited bio-
luminescence follows the Bunsen-Roscoe Law.

Observers have mentioned (LANcE, 1961; RickETTs
& Carvin, 1962) that Armina californica (CoopErR,
1862) will feed on Renilla, but I was unable to find in
the literature any details about the feeding patterns.
Therefore I conducted a group of experiments to de-
termine whether the feeding of Armina on the Renilla
would effect any special behavioral responses on the part
of the prey.

First, general observations were made of their inter-
specific behavior; second, a special experiment was per-
formed to determine whether the feeding of Armina
would cause a bioluminescent reaction on the part of the

Renilla; and, finally, a simple experiment was devised to
determine whether the body juices of the Armina would
produce an inhibitory or an enhancing effect on the wave
of bioluminescence.

METHOD

For the first experiment, the method was just general
observation of Armina feeding on Renilla. The animals
were in a tank through which fresh sea water was con-
stantly running.

The second experiment was to test the hypothesis that
the Renilla would give a bioluminous response when the
Armina feeds on it. One animal of each kind was placed
in a bowl with the other. The bowl contained sea water
and a sandy bottom to better simulate their natural en-
vironment. Observations were made in the dark, with
sufficient light only to see the position and behavior of
the two animals, but not enough to inhibit the biolumi-
nescence of the Renilla. Control experiments were also
performed, with a Renilla alone in a bowl with sand and
sea water, or with Renilla and a non-feeding Armina
together.

The third experiment, to test the hypothesis that fluid
from the Armina would inhibit the bioluminescence of
the Renilla, involved macerating one Armina in a blender
with 15 ml of sea water to aid the grinding process. This
suspension was placed on the rachis of a Renilla with an
eye dropper. Visual observation in a darkened environ-
ment would then determine if there were any differences
between the part with the Armina suspension on it, and
the part without. A stimulating probe induced the wave
response and differences would be judged according to
deviations from the normal bioluminescent pattern.

RESULTS

The general behavior of the Renilla when Armina feeds
upon it involves a retraction of the anthocodia and mus-

Vol. 10; No. 4

cular contractions of the rachis. The bioluminescent res-
ponse was tested for only in the second experiment and
will be discussed there.

Under the laboratory conditions (both with a sand
bottom and with only a glass bottom to the sea water
tank), the Armina ate the anthocodia, the rachis, and the
tip of the peduncle when exposed. Further experiments
should be performed to determine preference habits of
the Armina for these various parts of the Renilla, and
how the Armina senses the presence of the Renilla.

The results of the second experiment were that every
time the wave response was elicited from the Renilla, the
Armina was feeding on it. There were times during the
15 trials when the Armina was ingesting the Renilla, but
without stimulating the wave response. Often the flashes
would occur in rapid sequences of 6, 8, 10 or more waves.
The Armina exhibited no avoidance behavior, as far as
could be told from unaided visual observation. The Ar-
mina was also observed moving across the rachis, but
without feeding. At these times there were no wave pat-
terns elicited from the Renilla, although the rachis of the
animal was curling and folding extensively.

The results of the third experiment failed to substanti-
ate the hypothesis. No visual differences were observed
between that part of the rachis covered with the macer-
ated Armina and that part which was not covered by the
preparation. This result did not vary, no matter how
thoroughly the rachis was covered by the suspension.

DISCUSSION

There is a stimulation threshold below which the Renilla
will not exhibit a bioluminescent response. Certain behav-
ioral patterns of an Armina fall beneath this level
(crawling across the rachis, righting itself after having
fallen onto the Renilla, burrowing under the animal).
The only observed behavior of the Armina that caused
the Renilla to bioluminesce was feeding on any part of
the animal.

Continued identical stimulation can possibly result in
a raising of the threshold level necessary to produce the
wave response. This would explain why a number of
times the Armina was observed to be apparently eating,
but without eliciting the wave response. Any theory that
the bioluminous reaction serves as a defense mechanism
would have to account for this lack of behavioral response.

From the third experiment one must conclude (tenta-
tively) that the Armina possesses no chemical substance
that will inhibit the bioluminescence. The hypothesis had
been that such a chemical may exist as a narcotizing
agent. More refined measurements of the biochemistry
of Armina could result in either a verification or a nulli-

THE VELIGER

Page 441

fication of the conclusion drawn from this segment of
the experiment.

SUMMARY

When Armina feeds on Renilla, it will elicit a biolumin-
escent response which other behavior patterns will not
evoke. The body fluid of Armina seems to have no effect
on the bioluminescence. Other reactions of the Renilla
include muscular contractions of the rachis and retrac-
tion of the anthocodia. Armina will feed on any part of
the Renilla.

ACKNOWLEDGMENTS

The research was carried out at the Marine Laboratory,
University of California, Santa Barbara, during the sum-
mer of 1967. Mr. Tony Barnes and Mr. Al Eggleston of
the University, and Dr. Jack Tomlinson provided facili-
ties, experimental specimens, and research assistance. I
wish also to thank Mrs. Jacque Rodgers, Mr. Nelson
Baker, and especially Mr. Gale Sphon of the Santa Bar-
bara Museum of Natural History for reading the manu-
script and suggesting possible research methods, and Mr.
Bob Flather of Santa Barbara for assistance in the exper-
imenting laboratory.

LITERATURE CITED

Cooper, JAMES GRAHAM
1862. Some genera and species of California Mollusca.
Proc. Calif. Acad. Nat. Sci. 2: 202 - 207
Cormier, MILTON
1961. Biochemistry of Renilla reniformis luminescence. pp. 274
to 293. In: William McElroy and Bentley Glass (ed.): A
Symposium on light and life. Baltimore, John Hopkins Press,
xu+942 pp.; illust.
Harvey, E. NEwTon
1952. _ Bioluminescence.
xvit+649 pp.; 187 figs.
Jounson, Frank H. (ed.)
1955. The luminescence of biological systems.
ton, D.C. (AAAS) xiv+452 pp.; illust.
LancE, JAMES ROBERT
1961. A distributional list of Southern California opistho-
branchs. The Veliger 4 (2): 64-69 (1 October 1961)
PFEFFER, GEORG
1886. Neue Pennatuliden des Hamburger Naturhistorischen
Museums. Jahrb. Wiss. Anst. Ham. III, Beilage Jahresber.
Naturhist. Mus. Hamburg 1885: 53 - 61 [not seen]
Ricketts, Epwarp F. & Jack CALvIN
1962. Between Pacific tides. Stanford Univ. Press, rev. ed.
by Joet W. HepcretuH. 516 pp. Stanford, California
xiit516 pp.; 135 figs.; 46 plts.

Acad. Press, Inc., New York, N. Y.

Washing-

Page 442

THE VELIGER

Vol. 10; No. 4

A New Marine Mollusk from Mozambique

in the Genus Festilyria Pttspry & OLsson, 1954

(GASTROPODA : VOLUTIDAE)

CLIFTON STOKES WEAVER

Hawaiian Malacological Society

2777 Kalakaua Avenue, Honolulu, Hawaii 96815

(Plate 60; 1 Map)

In 1966 I optatinep for Mr. John duPont, a shell which
had been dredged in 40 fathoms of water off Boa Paz,
Mozambique, Portuguese East Africa. The specimen was
empty but in excellent condition (see Plate 60, Figures 1
and 2). Subsequently, I learned of the existence of a
second conspecific shell dredged alive in about 12 fathoms
of water 150 miles northeast of Lourenco Marques, a
town only a few miles south of Boa Paz (see Map). This
specimen, with animal preserved, is in the collection of
the Department of Mollusks, Museum of Comparative
Zoology, Harvard University, Cambridge, Massachusetts.
These 2 specimens seem to belong to a hitherto unknown
species.

The new species appears to be related to both Festi-
lyria ponsonbyi (E. A. Smitu, 1901) (see Plate 60, Fig-
ures 3 and 4) and EF festiva (Lamarck, 1811) (Plate 60,
Figures 5 and 6). It is unfortunate that the animals of
these 2 species are unavailable for study. However, there
are sufficiently strong morphological differences between
the 3 taxa to warrant specific separation.

In my arrangement of subfamilial and generic classi-
fication I follow Pitssry « Oxsson, 1954.

Fulgorarinae Pirspry « Orsson, 1954

1954. Fulgorarinae Prtssry & Oxsson, Bull. Amer. Pale-
ont. 35 (152): 16

Festilyria PrtsBrY & Otsson, 1954

1954. Festilyria Pitspry & Oxtsson, Bull. Amer. Paleont.
35 (152): 24

Type Species: Voluta festiva Lamarck, 1811, by OD.
Recent, Africa.

Distribution: In addition to the new taxon being de-
scribed below, there are 3 other Recent species in the
genus, all of which inhabit moderately deep water off
East Africa. They are Festilyria africana (REEVE, 1856),
FE. festiva (LaMarck, 1811), and F ponsonbyi (E. A.
SmitH, 1901). Two species, F africana and EF. ponsonbyi,
are known almost exclusively from specimens removed
from the stomachs of fishes.

Diagnosis: Shells medium to large, solid, ovate or fusi-
form. Spire high or low, turreted, or with rounded shoul-
ders; apex blunt. Protoconch pupiform, medium to large,
smooth. Teleoconch sculpture consists of shoulder nodules
or sharp tubercles which may or may not continue from
suture to suture as low, rounded axial ribs. Aperture
semi-ovate. Outer lip simple, slightly thickened, bevelled.
Columella with 3 rather strong anterior plaits (the second
being strongest), followed by several weak plaits covering
4 to } parietal area. A black callus or blotch may or may
not be present at upper and lower end of columella.
Siphonal notch narrow, deep; fasciole weak, without de-
fining ridge. Periostracum absent; horny operculum pres-
ent (Plate 60, Figure 7).

Remarks: Pitspry « Otsson (1954) stated that a fas-
ciole was absent in the type species, Festilyria festa.
This must have been a Japsus on their part as I have
examined a large adult specimen of F festiva at the
American Museum of Natural History, New York City,
and observed a distinct fasciole. Mr. John duPont con-
firmed this observation when he examined several speci-
mens of F festiva at the British Museum (Natural His-
tory), London. Apparently the lack of a defining fasciole
ridge caused this error. Furthermore, some books have

Vol. 10; No. 4 THE VELIGER Page 443

INDIAN
OCEAN

i I “I TAMATAVE

MALAGAS!Y
© REPUBLIC

20°

CAP SAINTE MARIE

t 5
) =: LOURENCO MARQUES
sS ORANILE De INHACA

30°

SO. AFRICA

LIM 1D) 4 i OEE Ais
35° pe ales 45°

Page 444

drawings of F festiva that show a distinct fasciole (K1-
NER, 1839, plt. 22, fig. 2, dorsal view) and some show
none at all (ReEve, 1849, plt. 12, fig. 28b). This, no
doubt, has also contributed to the error.

Festilyria duponti WEAVER, spec. nov.

(Plate 60, Figures 1 and 2)

Type Specimens: Holotype, Delaware Museum of Nat-
ural History No. 13706; Paratype, Museum of Com-
parative Zoology, No. 262150.

Type Locality: In 40 fathoms of water off Boa Paz,
Mozambique, Portuguese East Africa.

Range: From waters off Boa Paz to a point 150 miles
northeast of Lourengo Marques (24° 64’S; 34°50’
E).

Habitat: The only known living specimen (Paratype)
was dredged in 12 fathoms; substrate unknown.

Dimensions: Holotype length 124.4mm, maximum dia-
meter 56.5 mm, aperture length 72.6mm, maximum dia-
meter of 2™* protoconch whorl 3.5mm.

Description: Shell moderately large for genus, solid,
elongate-fusiform. Spire high, turreted, with rounded
apex. Protoconch pupiform, rather large, of 24 smooth,
deeply sutured, flesh-colored whorls. Teleoconch of about
54 strongly turreted and sculptured whorls. Sculpture on
early whorls consists of axial ribs, 15 such ribs on pen-
ultimate whorl of holotype, diminishing to blunt shoulder
nodules on later whorls. On early teleoconch whorls
axial ribs are crossed by many closely spaced revolving
lirae which disappear on later whorls. At anterior tip
of adult body whorl are 8 or more strong revolving lirae
bordering the fasciole. Aperture wide, semi-ovate, about
3 total length of shell. Outer lip simple, slightly thickened,
bevelled. Columella arched, with 3 rather strong anterior
plaits (the second being strongest), followed posteriorly

THE VELIGER

Vol. 10; No. 4

by 4 weaker plaits to midpoint on columella. Siphonal
notch narrow, deep; fasciole weak. A black callus at pos-
terior junction of outer lip with parictal wall and a black
blotch at anterior end of columella. Base color flesh, pro-
fusely overlaid with revolving zones of pinkish-brown
blotches; the narrow, pale, intermediate areas crossed by
numerous, short, curved, irregular pinkish-brown lines.
Inner edge of outer lip spotted with black where revolving
pinkish-brown lines terminate. A large horny operculum
is present.

Animal and Radula: Top of broadly expanded foot
creamy-white with numerous closely spaced reddish-brown
lines radiating outwards. These lines often split and ana-
stomose. Siphon and tentacles encircled by narrow bands
of reddish-brown.

I have not had the opportunity to study the anatomy

of the soft parts of the paratype at Harvard.
Remarks: Before comparing Festilyria duponti with its
closest relative, F festiva, the following remarks should
be noted: The holotype of F festiva illustrated in this
paper is a juvenile shell and I have no photograph of
an adult specimen. Therefore, I have referred to REEVE
(1849, plt. 12, fig. 28a) in making comparisons with a
fully adult shell, as well as referring to my notes on the
adult shell I studied at the American Museum of Natural
History in New York.

Festilyria duponti differs from F festiva as follows:
Shell is smaller, lighter in weight, and more attenuated in
outline; strongly shouldered and turreted whorls are in
sharp contrast to gently sloping non-turreted whorls of
FE. festiva; ribs do not run from suture to suture (other
than on first 2 teleoconch whorls) ; protoconch is smaller
and has 1 less whorl.

Compared to Festilyria ponsonbyi, E duponti exhibits
the following differences: It has ribs which terminate
posteriorly as low nodules rather than the ribless pointed
tubercles of FE ponsonbyi; protoconch is twice as large;
shell is larger and more attenuated; no black callus or
blotch appears on the parietal area of F ponsonbyi.

Explanation of Plate 60

Figures 1 and 2: Festilyria duponti WEAVER, spec. nov. Holotype,
ex DMNH no. 13706; dredged dead in 40 fathoms off Boa Paz,
Mozambique. Height 124.4mm; maximum diameter 56.5 mm;
photographs by Clifton Weaver.

Figures 3 and 4: Festilyria ponsonbyi (E.A.Smitru, 1901), ex
duPont collection, from the stomach of a fish caught off Durban,
Natal, Republic of South Africa. Height 100.8 mm; maximum dia-
meter 51.2mm; photographs by Clifton Weaver.

Figures 5 and 6: Festilyria festiva (LaMaRcK, 1811). Holotype, ex
MNHN, general collection of Volutidae No. 57; “Africa;” a juve-

nile specimen; height 71 mm, maximum diameter 31. 5mm; photo-
graphs courtesy Mr. H. Chevallier, MNHN.

Figure 7: Operculum from paratype of Festilyria duponti WEAVER,
spec. nov., ex MCZ no. 262150; animal dredged alive in 12 fathoms,
150 miles northeast of Lourengo Marques, Mozambique; photo-
graph courtesy Dr. Ruth Turner, MCZ.

Explanation of abbreviations used: DMNH=Delaware Museum
of Natural History, Greenville, Delaware. MCZ=Museum of
Comparative Zoology, Harvard University, Cambridge, Massachu-
setts. MNHN= Muséum National d’Histoire Naturelle, Paris,
France.

Tue VE IcER, Vol. 10, No. 4 [WeEavER] Plate 60

/f

’

Figure 5

Figure 2

Figure 3 < Figure 4

Figure 7

Vol. 10; No. 4 THE VELIGER Page 445

It gives me pleasure to name this beautiful species in
honor of my friend and associate, Mr. John E. duPont,
whose interest in the family Volutidae is well known.

LITERATURE CITED

Krener, Lours CHARLES
1834-1841. Spécies général et iconographie des coquilles vivantes
: genre volute 3: 1-69; plts. 1-52 (1839)
Lamarck, JEAN-BapTisTe PreERRE 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)

Pitspry, Henry Aucustus « AxEL ADOLF OLssoNn

1954. Systems of the Volutidac. Bull. Amer. Paleont. 35
(152): 271 - 306; plts. 25 - 28 (7 September 1954)
REEvE, LovELL Aucustus
1856. Descriptions of three new volutes from the collections
of the Hon. Mrs. Cathcart and Mr. Cuming. Proc. Zool.
Soc. London 24: 2-3; figs. 1, 2 (June 1856)
SmitH, Epcar ALBERT
1901. A list of the known forms of Volutidae from South
Africa, with descriptions of two new species of Voluta from
Natal. Proc. Malacol. Soc. London, 4 (6): 231 - 235

(October 1901)

Page 446

THE VELIGER

Vol. 10; No. 4

METHODS & TECHNIQUES

Boiled Lettuce and Cress
as Diet Supplements
for Certain Species of Mollusks

BY

T. W. FISHER

Department of Biological Control

University of California, Riverside, California 92502
AN ISOLATED BIT of information such as this normally
would be included in a more extensive biological study;
however, the boiled leaf technique has produced such
excellent results during the past several months it seems
worthy of bringing to the attention of Veliger readers. No
reference to this specific technique has been found in a
hurried search of the literature (KincsTON, 1966; KRULL,
1937a, b). Nonetheless, it is doubtful that other workers
have not chanced on this disarmingly simple, yet highly
successful, means of fecding certain species of terrestrial
and fresh water mollusks.

METHOD or PREPARATION

Coarse outside leaves of romaine and head lettuce and
mature large-leafed water cress, Nasturtium officinale
R.BR., have been utilized. The unaltered plant material
as whole leaves — no shredding is necessary — _ is
immersed about 10 seconds in boiling water and then
placed in cold tap water for immediate use or storage.
With weekly water changes a supply of boiled cress can
be kept (in water) at 40°F for several weeks. Boiled let-
tuce does not hold up as well and is best used within one
week. Depending on the number of snails to be fed,
entire or fragmented lettuce leaves can be placed with the
snails in amounts dictated by experiment to be consumed
in 48 hours. Boiled cress sinks and boiled lettuce floats.
Consequently, the latter is brought within reach of ben-
thic snails if it is weighted down with small rocks.

REARING CONTAINERS

In the laboratory aquatic snails are reared in 48 ounce
covered plastic containers (Bay, 1967) and 5-gallon aqua-
ria which are equipped with sub-sand filters and air
bubblers. Larger numbers are reared in a greenhouse in

wooden tanks measuring approximately 3 ft by 5 ft of
surface by 6 inches deep with a very feeble flow-through
of water, and with a layer of sand/gravel/soil on the
bottom. Succinea is maintained in a large covered rigid
plastic food container with a layer of soil which is covered
with decaying grasses. In the laboratory Helix and Limax
are maintained in 5-gallon aquaria with soil. A large col-
ony of Helix is maintained in a lathhouse in a screen-
covered pit with sides of concrete blocks.

MOLLUSKS REARED

Species which respond well to the boiled cress lettuce diet
are the following:

HyprosnpaeE: Fontelicella californiensis GREGG & TAYLOR
seem to prefer boiled cress, but readily accept boiled let-
tuce. The following mollusks have been fed mainly boiled
lettuce: LyMNAEWAE: Stagnicola proxima (palustris nut-
talliana (LEA) ); PLANoRBIDAE: Helisoma tenue califor-
niense FE C. Baker; PuysiwaE: Physa virgata Goup, P
gyrina Say. Terrestrial species which readily consume this
diet are: SucciNnEwAE: Succinea californiensis FISCHER &
Crosse; Limaciwae: Limax flavus Linnarus; HE ict-
DAE: Helix aspersa MU uuer. It is well known that H.
aspersa feeds on raw lettuce leaves as well.

DISCUSSION

Apparently, boiling breaks down the plant tissues, thus
permitting immediate feeding even by very early juvenile
mollusks. In the aquaria and larger wooden culture tanks
previously mentioned raw lettuce usually lies in the water
for two days before newly hatched snails will touch it,
and they do not swarm on it until the leaf is in a state
of decay one or two days later. A cooked leaf, on the
other hand, is attacked at once and may be skeletonized
in 4 hours if snails are abundant. In 24 hours usually only
the coarsest vascular tissue remains.

Investigation into a better snail feeding technique was
prompted by the need for a constant supply of small
juvenile snails which are used as hosts for first instar lar-
vae of marsh flies (Diptera: Sciomyzidae). After a few
hours in snail cultures bits of boiled lettuce or cress can
be torn away with juvenile snails still attached and trans-
ferred en masse to the fly cultures.

An interesting side observation has been the suitability
of this diet for various other fresh water invertebrates.
These include, among others, Ostracoda, Amphipoda
(Gammarus spp.), Eubranchiopoda, and planaria. These
animals frequently attach themselves to the boiled plant
material and it is assumed they are feeding on it.

The foregoing listing of mollusks includes all of the
species, representing 7 families, that have been reared on

Vol. 10; No. 4

the boiled lettuce diet. It is hoped that this diet will find
broader use and acceptance and that it will make possible
the laboratory rearing of species whose biologies are now
incompletely understood.

An extension of this technique might apply to the
study of small cryptic terrestrial mollusks. Since it is
often stated that many land mollusks are “humus feeders,”
the boiled leaf technique using leaves from their environ-
ment may serve as a short cut to “humus.”

If close control of mollusk cultures is required, boiling
of vegetation eliminates contaminants such as extraneous
fungi, arthropods, mollusks, etc. This method should not
be construed as a definitive, or complete, diet for mol-
lusks, but rather as a very useful supplement to the epi-
phyton normally available, miscellaneous decaying or
green plant material and shell building materials such as
calcium carbonate, limestone or shell.

LITERATURE CITED

Bay, Ernest C.
1967. An inexpensive filter aquarium for rearing and experi-
menting with aquatic invertebrates. Turtox News 45 (6):
146 - 148
Kineston, NEwTon
1966. Observations on the laboratory rearing of terrestrial
molluscs. Amer. Midland Natur. 76 (2) : 528 - 532
Kru, WENDELL
1937. (a) Rearing aquatic snails, pp. 523 - 526; (b) Rearing
terrestrial snails, pp. 526-527. In: Culture methods for in-
vertebrate animals. J. G. Needham, ed. Comstock Publ. Co.,
New York, N. Y.

NOTES & NEWS

A.M. U.

Pacific Division

The Executive Board of the American Malacological
Union — Pacific Division will meet at Asilomar Con-
ference Grounds, Pacific Grove, California on Thursday,
June 20, 1968 at 1:30 p.m. The annual Business Meeting
of the AMU-PD will follow immediately at 2:00 p.m.
For reasons beyond the control of the elected officers it
was impossible to organize any other program for the
1968 conference.

THE VELIGER

Page 447

W.S. M.

The first Annual Meeting of the Western Society of
Malacologists will be held at the conference grounds at
Asilomar State Park, Pacific Grove, California June 19
to 22, 1968. Scientific papers, symposia on related prob-
lems, and exhibits will be presented in the various fields
related to the study of malacology and invertebrate
zoology.

All persons interested in malacology and conchology
are cordially invited to attend, and participate in, this
historic meeting. 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, CA 90720.

Invalid Names in Oysters

BY

KENNETH JAY BOSS

Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts 02138

Recently 8 new names for species in the family Ostreidae
(Mollusca; Bivalvia) were introduced into the literature
by GitperT RANson (1967. Les espéces d’huitres vivant
actuellement dans le monde, définies par leurs coquilles
larvaires ou prodissoconques. Etude des collections de
quelques-uns des grands musées d’histoire naturelle.
Rev. Trav. Inst. Péches marit., 31 (2): 127-199, figs. 1-25;
(3): 205-274, figs. 26-55).

Although these so-called new species are accompanied
by “illustrations” - line drawings or photomicrographs
of prodissoconchs at magnifications of 200 x and above,
they are not validly introduced or available since they do
not satisfy Article 13 (a) of the International Code of
Zoological Nomenclature (1964)’ which states:

. a name published after 1930 must be either

(i) accompanied by a statement that purports to
give characters differentiating the taxon; or

(ii) accompanied by a definite bibliographic ref-
erence to such a statement; or

Page 448

THE VELIGER

Vol. 10; No. 4

(iii) proposed expressly as a replacement for a pre-
existing available name.
Ranson has not fulfilled any of these criteria. Neither
a written description, nor a comparison with related spe-
cies, nor an indication of type-locality, nor a citation of
type material are given. The names, consequently, are not
nomenclatorially available, and no zoologist should cite
them. Since they are unavailable there is not even any
need to place these names on the Official Index of Re-
jected and Invalid Specific Names in Zoology.

CALIFORNIA MALACOZOOLOGICAL Society, Inc.

Important Notice!

Because of drastic changes in Postal Regulations, which
went into effect on January 7, 1968, we will no longer be
able to include in our January issue an envelope as a
reminder that dues are payable. Other changes affect the
cost of returning and remailing of undelivered copies of
the journal. In addition, the mailing rate has been in-
creased and will be increased annually on January 1
each year for the next several years -— provided that
there is not some extra-ordinary increase made at some
other date. Changing the addresses also has recently been
increased. After a very careful study of the whole prob-
lem, we find that a change of address alone costs us just
one cent less than a dollar; return and remailing of
copies because of the addressee having moved causes us
an expenditure of a minimum of one dollar (depending
on the weight of the particular issue and the distance of
the original address). Since our charges reflect less than
the actual cost of producing the magazine, we cannot
any longer absorb the costs for these extraneous services.
We are forced to ask that with each change of address
our members and subscribers send along one dollar. For
re-mailing a returned copy we are asking a reimbursement
of $2.-, a fee which will include the change of address.
As we are not charging for the time consumed in render-
ing the service, and as we are not breaking even on the
charges for returning and remailing, we earnestly re-
quest that we be informed at least four weeks before the
change of address becomes effective. If that is impossible,
then we urge that arrangements be made with the local
postoffice for forwarding all second class mail - a service
which, in view of all other charges, is very economical.

CALIFORNIA
MALACOZOOLOGICAL Society, Inc.

is a non-profit educational corporation (Articles of In-
corporation No. 463389 were filed January 6, 1964 in
the office of the Secretary of State). The Society publishes
a scientific quarterly, the VELIGER. Donations to the
Society are used to pay a part of the production costs and
thus to keep the subscription rate at a minimum. Donors
may designate the Fund to which their contribution is
to be credited: Operating Fund (available for current
production) ; Savings Fund (available only for specified
purposes, such as publication of especially long and signi-
ficant papers); Endowment Fund (the income from
which is available. The principal is irrevocably dedicated
to scientific and educational purposes). Unassigned dona-
tions will be used according to greatest need.

Contributions to the C.M.S., Inc. are deductible by
donors as provided in section 170 of the Internal Revenue
Code (for Federal income tax purposes). Bequests, lega-
cies, gifts, devices are deductible for Federal estate and
gift tax purposes under section 2055, 2106, and 2522 of
the Code. The ‘Treasurer of the C. M. S., Inc. will issue
suitable receipts which may be used by Donors to substan-
tiate their respective tax deductions.

CALIFORNIA MALACOZOOLOGICAL SocrEty, Inc.
announces:

Backnumbers of

THE VELIGER

and other publications

Volumes 1 through 8: out of print Volume 9: $22.-
Supplement to Volume 3: $6.-* plus $-.35 handling charge
[Opisthobranch Mollusks of California
by Prof. Ernst Marcus]

Supplement to Volume 6: $4.-* plus $-.35 mailing charge
[Biology of Tegula funebralis (A. ApaMs), edited by
Abbott, Blinks, Phillips and Stohler]
Supplement to Volume 7: $2.-* plus $-.35 mailing charge
[Glossary of A Thousand-and-Onc Terms used in
Conchology, compiled by Wintrrep H. ArNoLp]

Items marked with * are subject to sales tax in the State
of California; residents of that State please add the
appropriate amount to their remittances.

Prices subject to change without notice.

Vol. 10; No. 4

Send orders with remittance to:

Mrs. Jean M. Cate, Manager, 12719 San Vicente Boule-

vard, Los Angeles, Calif. 90049. Please, make checks
payable to C. M.S., Inc.

Shipments of material ordered are generally made within
two weeks after receipt of remittance.

Backnumbers of the current volume will be mailed to new

subscribers, as well as to those who renew late, on the

first working day of the month following receipt of the
remittance. The same policy applies to new members.

Subscription to Volume 10: $12.- domestic; $12.60 in
Canada, Mexico, Central and South America;
$12.90 all other foreign countries.
Subscriptions to Volume 10 accepted until March 15, 1968.

Subscription to Volume 11: $18.- domestic; $18.80 in
Canada, Mexico, Central and South America;
$19.20 all other foreign coutries.

Affiliate Membership in the C. M.S., Inc. is $6.- for
the fiscal year July 1, 1967 to June 30, 1968. Postage for
members living in Canada, Mexico, Central and South
America 80 cents, for members living in any other foreign
country $1.20 additional. Membership open to indi-
viduals only - no institutional memberships. Please,
send for membership application forms to the Manager
or the Editor.

At a Regular Membership Meeting of the CALIFORNIA
MatacozootocicaL Society, Inc. the following policies
were adopted by unanimous vote:

There will be an initiation fee of $2.- (in addition to
the annual dues) for persons joining the Society on or
after January 1, 1967.

Members receive The Veliger free of further charges and
are entitled to purchase one copy of any supplement pub-
lished during the current membership year at a special
discount (to be determined for each supplement) .

Membership renewals are due on or before April 15
each year. If renewal payments are made after April 15
but before March 15 of the following year, there will be
a re-instatement fee of $1.-. Members whose dues pay-
ments (including the re-instatement fee) have not been
received by the latter date, will be dropped from the rolls
of the Society. They may rejoin by paying a new initiation
fee. The volume(s) published during the time a member

THE VELIGER

Page 449

was in arrears may be purchased, if still available, at the
regular full volume price plus applicable handling charges.

BOOKS, PERIODICALS, PAMPHLETS

Marine Shells of the Pacific

by Water O. CernonHorsky. Pacific Publ. Pty. Ltd.
Sydney, Australia. $6.50 Austr., > $9.50 U.S. 248 pp;
60 halftone plates; 20 text figs.

This book, while primarily based on the molluscan
fauna of Fiji, will be very useful to the collector of shells
from a much wider area than that island group. The
numerous excellent photographs will greatly assist in the
identification of species belonging to the families Bursi-
dae, Colubrariidae, Cypraeidae, Ovulidae, Triviidae, Mu-
ricidae, Mitridae, Olividae, Terebridae, and Conidae. It
would appear that a second and, possibly third, volume in
a series may be coming forth, which, if of the same
high quality as the present volume, would certainly be
welcomed by a large public.

In addition to the taxonomic treatment of the many
species, several other chapters add to the value of the
book. We especially like the one on the photography of
shells as the detailed instructions will enable even a
beginner in the new art to obtain satisfactory results.
Other portions which no doubt will be appreciated
widely are the ones dealing with collecting techniques,
the removal of radulae for study, and a short glossary. -
RS

Les Protoconques ou Coquilles Larvaires des Cyprées

by Gitpert Ranson. Mém. Mus. Nat. Hist. Nat., ser. A,
47 (2): 93 - 126; 39 halftone plates. Paris, 1967.

On the basis of outstandingly fine photographs of 149
specimens (cypraeid shells apparently ground down to
expose the protoconch) and the examination by the
author of an amazingly large number of specimens, Dr.
Ranson comes to the conclusion that the majority of
species belong to but a single genus, in which Kay in
1957 concurred. In the copy graciously sent to your editor
there is a handwritten list of 6 species which could not
be examined because the collection of the Paris Museum
contains only a single specimen of each. Therefore Ranson
does, by implication, not decide at this time that they

Page 450

Tine VEEIGER

should be included in the genus Cypraea. The 6 species
are: Cypraea teulerei, C. pulicaria, C. euclia, C. saulae,
C. gracilis and C. labrolineata.

It would appear to this reviewer that the illustrations
lend themselves, if taken by themselves and without con-
sideration of other characters, such as shell, radula, soft
parts, for a deviating interpretation, one which, in fact,
would support a different approach. It seems to allow a
logical grouping into what might be at least subgenera,
although in many instances the differences seem to be of
sufficient magnitude to justify generic separation.

If nothing else, this paper should stimulate a lively
debate between the less conservative interpreters of the
genus Cypraea (or, should we say ‘Supergenus’?) and the
(possibly) ultra-conservative interpreters.

RS

The Proboscis and Oesophagus of some British Turrids

by Epmunp H. Smiru. Trans. Roy. Soc. Edinburgh 67
(1): 1-21; 17 text figs. 1967.

On the basis of the examination of a number of rep-
resentatives of various genera of turrids, the author de-
scribes two new types of proboscides, calling one type A
or intraembolic and the other type B or polyembolic.

RS

The Neogastropod Stomach, with Notes on
the Digestive Diverticula and Intestine.

by Epmunp H. Smiru. Trans. Roy. Soc. Edinburgh 67
(2): 24-42; 11 text figs. 1967.

Vol. 10; No. 4

THE VELIGER is open to original papers pertaining to any problem
concerned with mollusks.

This is meant to make facilities available for publication of original
articles from a wide field of endeavor. Papers dealing with anatomical,
cytological, distributional, ecological, histological, morphological, phys-
iological, taxonomic, etc., aspects of marine, freshwater or terrestrial
mollusks from any region, will be considered. Even topics only indi-
rectly concerned with mollusks may be acceptable. In the unlikely event
that space considerations make limitations necessary, papers dealing
with mollusks from the Pacific region will be given priority. However,
in this case the term “Pacific region” is to be most liberally interpreted.

It is the editorial policy to preserve the individualistic writing style of
the author; therefore any editorial changes in a manuscript will be sub-
mitted to the author for his approval, before going to press.

Short articles containing descriptions of new species or lesser taxa will
be given preferential treatment in the speed of publication provided
that arrangements have been made by the author for depositing the
holotype with a recognized public Museum. Museum numbers of the
type specimens must be included in the manuscript. Type localities
must be defined as accurately as possible, with geographical longitudes
and latitudes added.

Short original papers, not exceeding 500 words, will be published in
the column “NOTES & NEWS”; in this column will also appear notices
of meetings of the American Malacological Union, as well as news items
which are deemed of interest to our subscribers in general. Articles on
“METHODS & TECHNIQUES?” will be considered for publication in
another column, provided that the information is complete and tech-
niques and methods are capable of duplication by anyone carefully fol-
lowing the description given. Such articles should be mainly original
and deal with collecting, preparing, maintaining, studying, photo-
graphing, etc., of mollusks or other invertebrates. A third column, en-
titled “INFORMATION DESK,” will contain articles dealing with any
problem pertaining to collecting, identifying, etc., in short, problems
encountered by our readers. In contrast to other contributions, articles
in this column do not necessarily contain new and original materials.
Questions to the editor, which can be answered in this column, are in-
vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will
attempt to bring reviews of new publications to the attention of our
readers. Also, new timely articles may be listed by title only, if this is
deemed expedient.

Manuscripts should be typed in final form on a high grade white
paper, 812” by 11”, double spaced and accompanied by a carbon copy.

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 Datias Hanna, Curator

Department of Geology

California Academy of Sciences, San Francisco
Dr. Joe, W. Hepcpetn, 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 STouier, 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

Mr. Atiyn G. Smiru, Associate Curator
Department of Invertebrate Zoology

California Academy of Sciences, San Francisco

Dr. RaupH I. Smiru, Professor of Zoology

and Chairman, Department of Zoology

University of California, Berkeley

Dr. Cuarzes R. STASEK, Associate Professor

of Zoology

Florida State University, ‘Tallahassee, Florida

Dr. Donato W. Witson, Professor of Biology
Department of Biological Sciences

Stanford University, Stanford, California

ASSOCIATE EDITOR

Mrs. JEAN M. Cate
Los Angeles, California

Ye w — ow oS — w — Ll fp w —_——
re E 2 a 5 2 E Gy > =
ju = > NE > = 7 f= 2a
Wb) ix E 2 \XAY = a = UY fl: Oe re
f a ” = Ww on ae beeing iene ony ne
a = in ase z= a , z m Zz
= — — w —ae
SHLINS SaluvUaIT LIBRARIES, SMITHSONIAN INSTITUTION NOILMJILSNI_NVINOSHLINS saiuvaai7 OI
ES ae = < = tt < S <
o = tf, = =a
: z 3 ‘ 2 Wy? : 5
8 2 3 2 8 Whe? g 2
= = S = Zh, YD fo Wan = 2 ea
= A Sa = SS 2 = = =i
SONIAN. INSTITUTION NOILOLILSNI_NVINOSHLINS SA1YVYGIT LIBRARIES SMITHSONIAN INSTITUTION Ni
ty, 2 us i Mi 5 tl ye til
Gy,“ z = z = e fe wz |
O fase < 2 < a < a < |
foi GS oc oc Cc ow = oe
va a a) = mo a al = coll
g 5 2 3 2 5) S ="
—_ a
SHLINS SAIMVYAIT LIBRARIES SMITHSONIAN INSTITUTION NOILMLILSNI NVINOSHLINS S314¥V¥aI7_ LI
oie rc z c z NCE oe c
ay ce ze z |
aS — w ae 2M — w = Joop
Xe = Pe) EN yee rad 5 ee 5 a
Oe > rif * F SF - > |
SE z a) a : a - 2a
2 m 9° 2 5 7
7 = on ; — — (22) 5 = q
SONIAN NOILALILSNI NVINOSHLINS S3INVUGIT LIBRARIES SMITHSONIAN INSTITUTION WN
z Z z 2 ZU RG Hen iee = =
z = 2 yy ae Ws = = =|
FE 2 c 2 iy = \. Z E 24
SHLIWS S31uvudly SMITHSONIAN INSTITUTION NOILMJILSNI_NVINOSHLIWS S31¥v¥g17_E
3 wn 2 o s Sa =a
>= ® = «2 w 6X # o
Nh ie = a5 a < a NGA =
= = e a 0 a) Ye a
si S a; (i 2 i S lee So
. J —J a 4 q
SONIAN” INSTITUTION ~ NOILMLILSNITNVINOSHLINS S31NVYaI7_ LIBRARIES SMITHSONIAN INSTITUTION _N
. ag z es S a S y ie a
y, ree (e} oo 4
LiL oo = jes] > — w = VY ow —
Vy, 2 : > Ne 5 : & Gy = =
ZY > = SS Mire > = Lo& K |
fi ® = 2 NS - % a a =i
OT Ne D ae = a a
m 2 a) Ye rm 2 m zy
SHLWS S31YvYGIT_ LIBRARIES, SMITHSONIAN INSTITUTION NOILNLILSNI_ NVINOSHLINS Saiuvua 1a JCF
= o Zz \ = = = YY
: XY 3 § WN: ig 5 Yeh
fo) Q \ SS 2 = JA IW (2) ReaD (e) i Sau
Bi, = = Wr 2: Ss =
2 ee Gaels a 2
SONIAN INSTITUTION NOILMLILSNI_ NVINOSHLINS S3IYVYEIT LIBRARIES SMITHSONIAN INSTITUTION
— ar w ey
Yi, © = = Uiy,?
Vly fier a < Di jae
a : eG lis
es —_— mo. =
e) ie) ZS e)
Ze = J) z

SHLINS SAIYVYaIT INSTITUTION NOILALILSNI NVINOSHLINS SS!1YVYdtI

INSTITUTION NOILNLILSN!

SJINVYUGIT LIBRARIES

INSTITUTION

NOILALILSNI NVINOSHLINS S3IYVYdE!IT LIBRARIES SMITHSONIAN INSTITUTION

INSTITUTION
Saluvudi
INSTITUTION

ISONIAN INSTITUTION

NVINOSHLINS S3IYVHYEIT LIBRARIES SMITHSONIAN
NVINOSHLINS S31YVUGIT LIBRARIES SMITHSONIAN

NVINOSHLIWS
Re
N NS
SMITHSONIAN
4:
La
NVINOSHLIWS

SMITHSONIAN

SMITHSONIAN
SMITHSONIAN

E (8 Geta J- ff“ | gE z} & \ WR ;
2 S Wass m CNS - eit m CeIn osI OS Z was’ m vy ; 2 Z I
— (2) = 9) = (2p) rye I (
ZINSTITUTION NOILNLILSNI_ NVINOSHLINS, SAI uvu a! LIBRARIES SMITHSONIAN INSTITUTION
< NN = VY yy < = ZN = < _ :
z x 4 Ys fj z = z \. = z \ :
re) \S Sn My O ac fo) : 3 Oo WS
= AS 2, GH Bos Ze i ‘N ~~ z E \ “SS
= S\N . S Me = S , = » WS > = Ny
nee w) es (7) us ;
NVINOSHLINS SA1YVUSIT_LIBRARIES INSTITUTION NOILALILSNI NVINOSHLINS S3lu¥vudl
| tu Zz a Zz if 2 hf ,
= a = o Be a = |
< a 0 = re = ce
ie cS 2 Gc e S oe |
je) ood o.? = faa) = co
a S 3 S in = af 3
SMITHSONIAN INSTITUTION | NOILNLILSNI SaluVudiT_ LIBRARIES SMITHSONIAN_ INSTITUTION
oe i z= oe J
fp? : : : ; 2 e
ee) A aw
> = > 4 > > a" >
a = eo) i 7 eet eS)
— Li aa = ws - =
i g ; : ; : ;
— (¢p) yi = on —_ (ep)
S3IYVYEGIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI
w za ond w i. nee wm = w
= y Hes = < = < =
=F “6 4 Zz a z z
i pig 2 z S z S z
e) MWY: ae re) 3e O ? sc oO 4
zZ li PN Z iE Z E Ea
| dia _ 2 2 ie cei -
SMITHSONIAN INSTITUTION NOILOLILSNI_NVINOSHLIWS SJINVUGIT LIBRARIES SMITHSONIAN _ INSTITUTION
2 ay S So ie Z au Fe
(op) SON 4
“i x WX = “ a *S
a < a WA < = < 3 a
= F et hae, Vale = e 5
2 5 = ae S$ Se =
ae hi —_J =
~Saluvualy LIBRARIES SMITHSONIAN INSTITUTION | NOILALILSNI-_NVINOSHLIWS_
Ge \
3 eS = 4ily ay re S =
= > =) G7 Yi > =) > =
= a = Vel; “2 = 0 =
io G4 eae et) anes LS =
Z m 2 m 2 i Zz
— w a w — =
SMITHSONIAN INSTITUTION NOILNLILSNI_ NVINOSHLINS Sa1Yvu a! ey BRARIES |
WE Ae a A: :
=2 Ne = Siok ars | z= = Zz
6 ASG =r YY; oO = 2 , NN = ro)
2 WS YG 2 & 2 SN 8 2
BAS Y far = 2 E Wy 2 E
MY Soe ee SAS >
AL eae 2 et ee
SJIYVYSIT LIBRARIES INSTITUTION NOILALILSNI_ NVINOSHLIWS Saluvy9gitq
n” = (2p) Ss n Sr eR oP) 3
us ee Lu 2 Lu (a) uy
oe @ —_ Lf — on ee = :
oc a tx ea ox = LYK oc ,
Wics = < ap a x c ~ NS <x |
oc oc ox Ww & |
ee = mo. = a 5 XS a |
Gi 2 oT 2 ins Fe ae ay
INSTITUTION NOILONLILSNI NVINOSHLINS S3I1YVYdIT LIBRARIES SMITHSONIAN INSTITUTION
a z Ne le is z i
Las S ® So 2 Ef See eee
hf » e > = > Puff *
; n = (a9) es o = o
| ee lYVYdIT LIBRARIES SMITHSONIAN pNOLLNLILSNI NVINOSHLINS |S 3 buvudit
: Se = ef CW = < = é =
PAR 2 EQ 2 5 z é 2 fh fy
By 2 We 2 Z a 2 ae,
m2 E \. 2 = 2 F 2 “yy
me 2 a ae - 2 i 2
- n v”) a ” -

INSTITUTION

NOLLALILSNI

be
=
z
S
\&2)
be
=
=
it
in

F i My, '
in ae wk ied if x ‘ ry f rere ’
eg hdd fod oll 4 dk AF cae fobs each 3 ae
‘ fi ened a ye Ane ’ ‘ re
hatin Mh Me : fue Vas Pe ‘ ‘ ‘ i
We fomregn st aCe TW, mee
y LALLA CRO Len ed 1 ' wa 1
ror des oy Ware feta on “astey t . F
: ete ae eee te ' ‘ , A Fit
CAM ted en git 4 tt ‘ ‘ : ' |
nitiaad hes cepts : :
dt Mir rhe rae ‘ mf ae ;
4 re i ys fa vA F
Wee of Miho P A me fl jae.
ah tee Khe hh 4h weetee
4 ’ ' "e ‘ di ’ ' 4
[i ‘ ‘ ‘ ’ jt '
oe et ay
, ‘
. ‘ ‘
45,45 Ro ' \
’ m4 ‘
\ i : ‘
Janse hk A aca i
RAL ; ;
rene att M ‘ , bey Net wa ‘
TRIES VV tis : ieua
Peat vay ar hak it \ : ‘
wa cet By i tot ;
rs tS, 4,09,8 NON t 1
IAAL us JUROR itt {
Seeks \ : ; i)
WA RA wi 4 4 a Ved Can ety ’ i
Peon ic Ue ae vA \ wa 1 ‘
Nira Fea y Ves . ab Pig wha ’
Sy teh a) ( ‘
es a PUN HT SS AM AVN, 1 ‘
4 we ed '
Ans Vy teh, ‘

Se tty,

aay i

Wie tks
Renan .
DON
i Leh May

\

iy 4

ea
mY

Ae eth, a8)
Boke We WW
N"4

ewe ys
VL aay
A

DS Oak Ys CR

wher Stee
ae

Wye ay 4

yey at 4

MAES ASL HIN Ua ha
Wy 4.

LK TA

sa Ak

aN!

ST

Heh
a A

‘GN,
CST RAL
Ayn ) wel he
RA Bs ap Ga ACY

Wibeose
Vit at Wy)
Deets ‘ie Re irae
AR PERLE en
Ml CN

. ’ i doe ts
aN bt
SAN, eaditary

av * VA ke
See Ae At
Hy &
Bain 4

ee asarasi

eT ad

i waits
Aneta Ge) 8 SOR tn
ah, sti Ki Ae FPO We RI aia

ml Se AEN MR

yA {4 Wy ficial
ty Seon ve
on, Ve bi, %
neal, Mie i AN

" Dan)
Monit he

sa
Hoey ae

: Womb ger 8 Sod a bey
DC Waa AC OD
Wh Ad ak th Yea Skea Ga Ta bak
; ‘

‘iia
IE

Rf yal
VAS,
(yh

1 ae aS
Re DOE
De ey
FAA OL ey

DO MR

ik

ag ate Th AOL CA OA f

“oy ; fh ates vat Vinsiva

oh ROE ” hy) ways ‘ ’

aaah ARSE Bay Oh ek 5 aye oA a
waa a 4 taste DANN NS

4h

\ !
EVUD NE REY

SMITHSONIAN INS

WIAA

088 00838 8357