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VELIGER
A Quarterly published by
NORTHERN CALIFORNIA MALACOZOOLOGICAL CLUB
Berkeley, California
Volume 4
July 1, 1961 to April 1, 1962
Page II THE VELIGER Volume 4
Foreword to Volume Four
Practically everything said in the Foreword to the two preceding volumes could be repeated,
word for word, except, perhaps, that we must admit that our hopes of a brighter financial picture
at the end of this year did not materialize. Of course, we realize that we are to blame for this
ourselves, because we have continued experimenting with the format and have added new features.
We think that it was worth the effort. Possibly we will be accused of being conceited, but never-
theless we are proud of the growth of The Veliger.
As before, we have enjoyed the wholehearted cooperation of many individuals who have, with-
out exception, gone beyond the call of duty to assist us in our efforts to make this a better journal.
The extremely time-consuming job of typing the copy was again performed skillfully by Mrs.
Heidi S. Norskog. Mrs. Emily Reid has donated uncounted hours for the excellent art work re-
produced in these pages. The letterpress work again was contributed by your editor in his "spare
time", and his family deserves recognition for bearing with the absences for many extra hours as
well as for patience with the delay caused by this activity in performing the handyman tasks
around the house. Also, the special care given to the production by the employees of the Printing
Department deserves a word of praise; if we mention just a few by name, it is not to neglect
the others who should be included also: Earl Gustafson, Charles Peckham, Louis Rengel, and
John Schoen.
Perhaps the most striking advance we have made this year was the addition of color plates.
This was possible, in part, because of the generosity of certain individuals who contributed toward
the cost of the experiment. But the extraordinary detail shown in the reproductions is due solely
to the unusual skill and devoted care of Dr. and Mrs. G Dallas Hanna and Mr. Maurice Giles who
have produced these plates for us as an accommodation and on an experimental basis. To them
we wish to extend a special word ot thanks.
One step forward, we think, will be overlooked by most. That step is the reduction in the
number of typographical errors that still pass the watchful eye of the editor. This improvement
is due entirely to the extreme care given the reading of the copy by Mrs. Jean Cate and to her
goes Our appreciation. The index has again been under the continued care of Dr. Phyllis Kutsky.
And as before, there are many other persons who have contributed, in one way or another, to
the current volume. Among these are the members of the Editorial Board, the Officers and Mem-
bers of the Northern California Malacozoological Club, as well as many others, too numerous to
be listed by name. To all, named and unnamed, go the continued thanks of
Your Editor.
Berkeley, California, April 1, 1962.
Volume 4
THE VELIGER
Page III
TABLE OF CONTENTS
About copyright.
Rudolf Stohler . . cnt coe VN
A discussion of Gear & regina (GeaeHios 1825)
and related species, with description of a new
subspecies.
Meame Vins Cares (rie ieee 6 elon 200
A distributional list of Southern California
opisthobranchs.
Janae IRs IbEMCGs fos Go. fee 64
A living fossil.
lynemy 12, CmAce oo 6 oe seein 162
A new commensal polyclad fuera Panama.
ingisawmiGl Ii, Samy 5 56 6 6 6 oo oF WY
A new Dampierian SERtene
Crawford N. Cate. . Ocoee ree MES)
A new deep-water Anadara een the Gulf of
California.
Brace Canypoell, o 6 6 oo oo 0 0 Ube
Anew doridid nudibranch from Torquay, Vic-
toria.
ROSH: IWIN S 6 dg o -0 © o 9 55
A new method of determining the accuracy of
geotactic orientation of the snail Helix as-
persa Miller.
Donald R. Bower . . 5 oo lo A
A new subgenus and species of eeea’ inhabiting
barnacle from the Gulf of California.
Wicior Ao ‘Aulllote “6 Salo 6 ~o 6 Jo wl
A new Vexillum (Mitridae) from the Philippine
Islands.
Yeain Mi, GA 5 Bi Oe A pOeliton Meron, 4 woes
A new whale barnacle ee Late Pleistocene
deposits at San Quintin Bay, Baja California.
Wiiei@r Ao ACOs 6 6 « 6 »o oo) o 1 de
Another statistical study in size of cowries,.
ING My S@ouGlem 5 oo 6 06 © 6. 6 107
A preliminary report on spawning and related
phenomena in California chitons.
Spencer IR, Worse, Yeo 6 6 oo o OWA
Arion ater (Linnaeus) in California.
AMlsinn. (Gig Stools oi 5 Go oe ZS
A statistical study in cowries: The size of Mau-
ritia arabica (Linnaeus).
Ng Ako S@oullleler 5G 6 56 «6 oo 9 15
A study of food choices of two opisthobranchs,
Rostanga pulchra McFarland and Archidoris
montereyensis (Cooper).
lBimouillsye JE 5. COOKS Gg. 6. oi 46 o- ao. ao a. GE:
A study of the reproductive cycle in the Califor-
nia Acmaeidae (Gastropoda), Part Ill.
lnlaieicyy 1G, iMreynelsranain, WL 5 5 ¢ 41
A study of the reproductive cycle in the Califor-
nia Acmaeidae (Gastropoda). Part IV.
Harry K. Fritchman, II . 134
Busycoptus (B.) canaliculatus in San Francisco
Bay.
ixyaclolki Swelnee 4 5 6. o 6 6 6. 4 @blil
California brown cowrie in Central California.
AMUN (Gio, Sau a9 15 6) 6 16 6. 6 o. Gils
Ciliary currents in the mantle cavity of species
of Acmaea.
Go IMI, Wome. 6 Oi Ot tO Os lls 04 aces ad be)
Color photography of lang marine mollusks,
AMiee@el Ay IBNIEerR 56 16 5 6 47
Egg-laying in Fusitriton onguomencile (Redfield).
Faye B. Howard... so 6 GO)
Eucrairia nom. nov. for Drcaaahcllin Burn.
NODE IMEI. oo @ oo 0 6 oo 0 oo Bil
Four new Panamic gastropods.
Bruce Campbell. . . eels a PINS:
Limestone boring by the Tayi ‘Upahersneee.
INj@serageyo IMi, Veloso 6 56 Go 6 o 123
Manometric measurements of respiratory ac-
tivity in Tegula funebralis.
Yarnes Ik, IMIGILGER 6 «6 6c .o0 o, 6 o AGI
Name change in Mitra.
Jee Ml, CAE o 6G o «a o ao a)
Narcotizing and fixing Opisenoneanene!
laeronwinel Isl, Sell 5 6 50 oo 0 0 56 Be
New deep water mollusks from the Gulf of Cali-
fornia.
Donald R. Shasky . . . 0 oso J
New name for Strombus geanulaturs subsp. acutus
Durham, 1950, not Perry, 1811.
Yo Wihyae IDoweloen ~ 5 5c 0. 6. 9 GUS
Nomenclatural notes on some West American
mollusks with proposal of a new species
name.
Pio Mise. IS@GN 5 96 og 6. 6 178
Notes on cleaning mollusks.
PU Go Sreotde 5 6 9 6 so ae ages At SZEIG
Notes on rare and little hae Panamic mol-
lusks.
Donald R. Shasky . . . ° o | 6A
Notes on the Mitridae of the Eee Pacific, I.
Mitra fultoni E. A. Smith.
GalleiGesphon,y Jima tau 32
Notes on the opisthobranchs be the test Gonst
of North America. I. Nomenclatural changes
in the order Nudibranchia (Southern Califor-
nia).
Joam 13, Sieimnery 5 o o «0 «0 oo 0 BY
Observations of the effect of diet on shell col-
Oration in the Red Abalone, Haliotis rufescens
Swainson.
DAWG! IL, ILeneNion 5 6 5 6 29
Observations on the biology on Hermaeina
smithi, a sacoglossan opisthobranch from the
West Coast of North America.
Nie AUieek GrOMO Ut ye ser fers Wels arze ey Me Paris ts 85
Page IV
THE VELIGER
Volume 4
Observations on three species of Vexillum (Gas-
tropoda).
lig@am Mio Gat® 26 o 0 «0 6 @ oo og AWS
On certain littoral species of Octolasmis (Cir-
ripedia, Thoracica) symbiotic with decapod
Crustacea from Australia, Hawaii, and Japan.
Wiwahiaion AN\o INGEN 56 Cl Cl Cl Cli YY
On Cypraea tigris schilderiana Cate.
Aingom Kay o oa o Bee ont Se eee tie)
On the identifications of age Pacific Mitra.
V@aia Mi, GAB 6 6 6 athe! ite Maan eP ELS 2
On the systematic place of Cyprian mus.
ING IMispre, ISG 5 oo Say wale eee wed Ovt
Preliminary report on erowth studies in Oli-
vella biplicata.
Roclolkt Groller 5 og -o 6» 0 0 9 o ISO
Range extension for Tenaturris nereis (Pilsbry
and Lowe, 1932).
Helen DuShane . . 0 0 0 ta o BY
Range extension for Teoian elsiae Howard and
Sphon.
IPAWEe IBo IblowasGl 56 56 o 60 «0 o
Range extension of Anatina cyprinus (Wood,
1828).
rece Caynsoel, 5 6 a ees NUS
Recent uses of non- hoe works.
R. Tucker Abbott . at cee cits C 213
Reinstatement of the specific name Macoma in-
quinata (Deshayes).
AS Mynrakeeni 0). 0 0 161
Relationship of living eet to ell cavity vol-
ume in Helix aspersa.
Andrea Herzberg & Fred Herzberg 112
Remarks on a variation in Cypraea annettae
Dall, 1909. re aa
GrANMORGC ING GAS 5 oc 6 oc : 112
Revision of some Hawaiian mitrid species.
Yea Mio CAO «o o 6 o 6 140
Revision of the Clavagellacea.
liee Anderson Smith . . . 5 9 Nor
The fresh water clam Pisidium moemipert (Han-
nibal),
ID) iio WWElSAone We inl, Iso Ini@imreinyettorl 5 9 ies)
The genus Bankia Gray (Pelecypoda) in the Oli-
gocene of Washington.
J. Wyatt Durham & Victor A. Zullo . .1
The W. Mack Chiton Collection.
AMMSio, (Gio Sissi og G 9 0 5 5 «wi
Herrington, H. B.
Gig Wasplose, ID)o Wo We ==
Hertllein,, Weol Ga ere GDS) ape Ca
Herzberg, Andrea & Fred Herzberg . . 197
Hodgkin, Norman M. Siete oo 8
Howard, Faye Bs .*.s .« =. «© (5 oeOnmuao
Kay,,-Allis6én 9... 40 bo) a, 7 nee oe oe
Keen, A. Myra
(117), (118), 161, (164), (166),
ILAINCG, VARAGS Bo o o 0 0 a oo 0 Oth
ibeyeaincom, IDewGl in 56 56 6 ao 0 0 9
WieILeAm, Jamm@S Il, 56 o 0 06 oo « oa
INGwaaoeve, Wollitema NG 56° 6 5 6 o oo « YW)
Schilder, F. A. sitet. U: is Aida eh Les 15, 107, 199
Shasky, Donald R. too ecme Ley Ze
Smith, Allyn G.
. (116), (165), 214, 215, 216,
Smith, Edmund H. ee ec mea Se, 5)
Smith, Wee Anderson... 2) es) etnr) aceon
Sphon, :Gale:G., Jars 4) ohone Ie ee
Siresiaoyryy, Mern IN5 5 6 o » a 6 0 Sl
(218), (219), (220)
Stohler, Rudolf (53), 150, 162, (164), 211, 217
Taylor, Dwight W. & H. B. Herrington . 129
Thorpe, Spencer R., Jr. Se Gl be, 6 COZ
May SGI ee oo Ss bon a 6 6 UY
Fialilioy » Vaiietor WAtey io) wll etelatcpiiiees Io Se
cf. also: Durham, J. Wyatt & —
Volume 4
THE VELIGER
Page V
JOUR IRL AN AL! Ja
In the following list, compiled by Mrs. Crawford N. Cate, the
first column refers to the page, the second column to the column on the
page, the third column to the line in which the error is found; in the
fourth column the error is repeated, and in the fifth column the correct
word is given.
from the top of the page.
19
12
Bl
63
70
78
83
84
90
98
104
132
153
211
ZU
Z5
NN eS N
1
32 Essay
Plate 3 explanation;
hypotpye
Plate 4 explanation, figure 15:
Turitella
8 Monserrate
27 regards
20 Owens
14 from bottom:
Haliotis rufescens
25 from bottom:
1841
Unless otherwise stated, the count of the lines is made
Essai
hypotype
Turritella
Montserrate
regard
Owen
Haliotis rufescens
1840
bottom line of textfigure explanation:
localty
14 from bottom:
Mitra fultoni
12 reef
25 from bottom:
A. scabra
5 Eucrairia
Inside front cover, bottom line:
Sufamily
bottom line:
L. O. Yates
30 luracola
2 from bottom:
V. compressum
Figure explanation:
Bardiagrams
32 Fort Douglas
Textfigure 7 explanation:
appearanc
(Pruvot-Fol):
Etudes
19-21
Plate 31, righthand column:
Ballenas, Bay
32 W ood)
12 Anadara
8 Cantrell
14 from bottom:
cowry
1 cowry
locality
Mitra fultoni
reef-associated
A. scabra
Eucrairia
Subfamily
Gee Vaitels
luracola
V. compressum
Bar Diagrams
Port Douglas
appearance
Etudes
Delete
Ballenas Bay,
W ood,
Anadara
Cantrell
cowrie
cowrie
=
THE ye
VELIGER &
A Quarterly published by
NORTHERN CALIFORNIA MALACOZOOLOGICAL CLUB
Berkeley, California
VOLUME 4 JuLy 1, 1961 NUMBER I
CONTENTS
The Genus Bankia Gray (Pelecypoda) in the i of ee (3 nee
J. Wyatt DurHAM & Victor A. ZULLO . . 3a : : I
A New Vexillum (Mitridae) from the ae ieee (Plates iy 8 di TOCEGES)
JEAN M. Cate. . . cathe Se ee eae it) eben wer
What is Anatina anatina? 5 Testfigures)
A. Myra KEEN. . : 5G)
A New Whale Barnacle an ee BM occas Deven at wee Ouints Bay, Baja
California (Plate 3)
Victor A. ZULLO Ch esets Dae Ear An weg tae BU ar 18
A Statistical Study in Cowries: The Size of Mauritia arabica (anna 26 Testi )
F. A. ScHILDER aoe : ears
New Deep Water Mollusks A ie Gulf of California (Plate 4, Fie I- 5
HTD) NVA BRS EVA'S Val spe tee ig rl is i compan pee Woo Lachey eneea ne rlinnce Nags Maha cr | se ah Gee cerruae cer GEO
Notes on Rare and Little Known Panamic Mollusks (Plate 4, Figures 11-16)
Donatp R. SHASKY . . . abe tee catia tad ipcrc ce WBaNr al aly coer na FMM bear Nes aD
Four New Panamic Gastropods (Plat oh
G. Bruce CAMPBELL Ma 3 ; Seth Oly
Observations of the Effect of Diet on Shell Calcio in ine. Red ie acs
Haliotis rufescens Swainson (Plate 2
DAVID ME ICETGHMTONI eee eee nelle Wea een ore lee bel ncventale acPalpe ORY. 20.0)
Notes on the Mitridae of the Eastern Pacific I - Mitra fulton E. A. SmitH
(Plate 7; 1 ae
GALE G. SPHON, JR. : oh ot babs eh aaa ame aOR Re eat ae heen a
On Cypraea tigris schilderiana CATE (Plate 8; 2 Textfigures)
ALISON Kay ee rete yak metre te a (lien lk Mapu hanndase acoavam ae enc no eo
[Continued on Inside Front Cover]
Subscriptions payable in advance to N. C. M. C.
$3.50 per year ($4.00 to Foreign Countries)
$1.25 for single copies.
Send subscriptions to: Mrs. PHOEBE Batcu, Treasurer, 1150 Brown Avenue, Lafayette,
California. Address all other correspondence to: Dr. R. STOHLER, Editor, Department of
Zoology, University of California, Berkeley 4, California.
CONTENTS
[CONTINUED |
A Study of the Reproductive Cycle in the California Acmaeidae (Gastropoda).
Part III. (Plates 9 to 14)
Harry K. Frircuman, II ee
Color Photography of Living Marine Mollusks
ALFRED A. BLAKER awe ee
NOTES & NEWS: Here LETS oy hinh Tice. Ga, es 28
Range Extension for Trivia elsiag HOWARD & SPHON. Faye B. Howarp.
Range Extension for Tenaturris nereis (PitsBry & Lowe). HeLen DuSHANE.
Name Change in Mitra. Jean M. Cate. :
Eucrairia nom. nov. for Drepaniella BURN. ROBERT BurRN.
METHODS & TECHNIQUES Dg Woe
Narcotizing and Fixing Opisthobranchs. Epmunp H. Smit.
BOOKS, PERIODICALS, PAMPHLETS
41
47
50
52
33
Vol. 4; No. 1
THE VELIGER
Page 1
The Genus Bankia Gray (Pelecypoda)
in the Oligocene of Washington
by
J. Wyatr DuruaM and Vicror A, ZULLO
Museum of Paleontology, University of California, Berkeley 4, California
A contribution from the University of California Museum of Paleontology, Berkeley, California
(3 Textfigures)
The family Teredinidae, including the genera
Teredo Linnaeus and Bankia Gray, is a diverse
and widely distributed group in Recent seas.
However, the number of fossil species that have
been described are relatively few, even though
the genus Teredo is known from deposits as old
as Jurassic and the genus Bankia is recorded
from deposits of Paleocene age. The teredinids
are not uncommon in the fossil record, but
their presence is usually indicated only by bur-
rows with calcareous linings in fossil wood,
rather than by the shell or pallets. To separate
Teredo from Bankia it is necessary to examine
the siphonal pallets. These pallets, which seal
the aperture of the burrow in case of need, con-
sist of a calcareous stalk and blade and are
covered by periostracum. In the genus Bankia
the blade of the pallet is composed of a series
of nested cones. In comparison the blade of the
pallet of Teredo is entire. Previously, the or-
namentation and shape of the shell have been
used to delimit living and fossil species, but
these features have been shown to be environ-
mentally controlled and not constant for a spe-
cies. More recently (i.e., Clench and Turner,
1946), the character of the periostracum cover-
ing the siphonal pallets has been considered
Significant at the specific level, but the perios-
tracum is not readily preserved, and, conse-
quently, fossil species must be defined on other
characters. The morphology of the calcareous
parts of the siphonal pallets appears to be of
some value in specific determination, but de-
tailed illustrations of this part of the pallet are
seldom available, even for Recent species.
Therefore, it is with some reservation that the
following species is described. However, as
this discovery represents, insofar as is known,
the only fossil record of the genus Bankia in
the eastern Pacific, and as the specimens are
not readily comparable with any known species,
Recent or fossil, the description of this species
appears justified.
Several shells and pallets of this new species
of Bankia were collected from the middle Oli-
gocene Lincoln formation near Porter, Wash-
ington. The specimens are preserved in un-
filled burrows in a fragment of wood (Morus ?)
embedded in a concretion about 160 mm. long
and 90 mm. wide. Most of the material is
coated to varying degrees with calcite, but the
details of one pallet are exceptionally well pre-
served (text fig. 1). This new species is des-
cribed as follows:
Family TEREDINIDAE LartreILte, 1825
Genus Bankia Gray, 1840
Banka lincolnensis DURHAM & ZULLO, spec. nov.
(Figures 1, 2, and 3)
Description: Shell similar to that of other
species of the genus Bankia, equivalve, strong-
ly convex, gaping widely anteriorly; exterior of
shell not visible; interior of shell covered with
thin coating of calcite; lobe a little more than
one-third length of anterior margin of disc;
auricle placed centrally on posterior margin of
disc, not produced posteriorly; apophysis flat—
tened dorsoventrally, extending ventrally be-
yond center of shell; pallets consisting of series
of closely-spaced cones; calcareous part of
each cone funnel-shaped, semi-circular in
cross-section, with inner margin higher than
outer margin; inner margin nearly straight;
outer margin slightly concave on either side of
low, median convexity; lateral edges of cal-
careous part of each cone produced into long,
narrow, vertically extended "'awns"; stalk of
pallet long, length about five times width of
proximal cone, circular in outline.
Dimensions: Holotype UCMP no. 34'672;
length of burrow 56 mm.; maximum width of
burrow 6 mm.; width of distal cone of pallet 2
Paratype UCMP no. 34'675; height of
length of shell 5 mm.
mm.
shell 5 mm.;
Page 2
Fig. 1. Bankia lincolnensis DURHAM & ZULLO, spec. nov.
Holotype, UCMP no. 34672, x 9.6
outer view of incomplete pallet (secondary deposits
of calcite only partially indicated).
Holotype: University of California, Museum
of Paleontology no. 34'672.
Paratypes: University of California, Museum
of Paleontology nos. 34'673, 34'674, 34'675.
The specific name "lincolnensis'' is derived
from the Lincoln formation in which the speci-
mens occurred.
Occurrence: University of California, Muse-
um of Paleontology locality A-8724, middle
Oligocene Lincoln formation, ''type Porter"! lo-
cality. Southeast quarter of the northeast quar-
ter of Section 28, T. 17N., R. 5 W. Twenty-
foot stratigraphic interval along cliffs on north-
east side of State Highway 9 for a distance of
one-half mile immediately southeast of Porter
Creek,
THE VELIGER
Fig. 2. Bankia lincolnensis DURHAM & ZULLO, spec. nov.
Paratype, UCMP no. 34673, x 19.2
paired pallets heavily coated with secondary deposits.
Vol. 4; No. 1
Vol. 4; No. 1
THE VELIGER
Page 3
Fig. 3. Bankia lincolnensis DURHAM & ZULLO, spec. nov.
Paratype, UCMP no. 34674, x 9.6
outer view of series of cones partially
embedded in matrix.
Discussion: The pallets were compared with
those of 1) the Recent eastern Pacific species
Bankia setacea (Tryon) and B. zeteki Bartsch;
2) the Recent western Atlantic species (after
Clench and Turner, 1946) B. gouldi Bartsch, B.
caribbea Clench and Turner, B. katherinae
Clench and Turner, B. destructa Clench and
Turner, B. fimbriatula Moll and Roch, B. fos-
teri Clench and Turner, and B. cieba Clench
and Turner; 3) the Recent western Pacific spe-
cies B. australis Calman and B. debenhami
Iredale, and the New Zealand Oligocene species
B. turneri Powell and Bartrum; and 4) the Eu-
ropean fossils figured by Moll (1942, pl. 24).
The vertically projected ''awns'' developed on
the calcareous part of the cone of the pallet
serve to distinguish Bankia lincolnensis from
the above mentioned species. Bankia lincoln-
ensis most closely resembles species of the
subgenus Neobankia Bartsch in the character of
the pallets.
Teredid borings have been reported (as Te-
redo sp.) from both Cretaceous and Tertiary
formations on the Pacific Coast of North Amer-
ica including the following: Cretaceous, Santa
Ana Mountains, Southern California (Packard,
1916, p. 147); Paleocene, Martinez formation,
California (Dickerson, 1911, p. 173; 1914a, pp.
73, 78, 82, 86, 101; 1914b, p. 295); Paleocene,
Meganos formation, California (Clark and
Woodford, 1927, p. 103); late Eocene, Tejon
formation, California (Dickerson, 1916); late
Eocene, Cowlitz formation, Washington (Wea-
ver, 1916, p. 24); Oligocene, San Lorenzo for-
mation, California (Clark, 1918, p. 162); mid-
dle Oligocene, ''Porter", Washington (Van
Winkle, 1918, p. 77); middle Oligocene, Van-
couver Island, British Columbia (Merriam,
1896, p. 104); early Miocene, Vaqueros forma-
tion, California (Loel and Corey, 1932, p. 234);
early Miocene, Sooke formation, Vancouver
Island, British Columbia (Clark and Arnold,
1923, p. 156); late Miocene, ''Santa Margarita''
formation, California (Nomland, 1917, p. 302).
Personal observation indicates that fossil te-
redinids are more common on the Pacific
Coast than is indicated by the records listed
above, but they have largely been ignored.
Literature Cited
Clark, B. L.
1918. The San Lorenzo series of middle California.
Univ. Calif. Publ. Bull. Dept. Geol., vol. 11, pp.
45-234, textfigs. 1-4, pls. 3-24.
Clark, B. L., & R. Arnold
1923. Fauna of the Sook Formation, Vancouver Island.
Univ. Calif. Publ. Bull. Dept. Geol. Sci., vol. 14,
pp. 123-234, pls. 15-42.
Clark, B. L., & A. O. Woodford
1927. The geology and paleontology of the type section of
the Meganos formation (lower middle Eocene) of
California. Univ. Calif. Publ. Bull. Dept. Geol. Sci.,
vol. 17, pp. 63-142, pls. 14-22, 1 map.
Clench, W. J., & R. D. Turner
1946. The genus Bankia in the western Atlantic.
sonia, vol. 2, no. 19, pp. 1-28, pls. 1-16.
Dickerson, R. E.
1911. The stratigraphic and faunal relations of the Mar-
tinez formationto the Chicoand Tejonnorth of Mount
Diablo. Univ. Calif. Publ. Bull. Dept. Geol., vol. 6,
pp. 171-177.
John-
’
.1914a. Fauna of the Martinez Eocene of California. Univ.
Calif. Publ. Bull. Dept. Geol., vol. 8, pp. 61-180,
pls. 6-18.
1914b. The Martinez Eocene and associated formations
at Rock Creek on the western border of the Mohave
Desert area. Univ. Calif. Publ. Bull. Dept. Geol.,
vol. 8, pp. 289-298, textfig. 1.
1916. Stratigraphy and fauna of the Tejon Eocene of Cal-
ifornia. Univ. Calif. Publ. Bull. Dept. Geol., vol.
9, pp. 363-524, pls. 36-46.
Loel, W., & W. H. Corey
1932. The Vaqueros formation, lower Miocene of Cali-
fornia. I, Paleontology. Univ. Calif. Publ. Dept.
Geol. Sci., vol. 22, pp. 31-410, pls. 4-65, 2 maps.
Page 4 THE VELIGER Vol. 4; No. 1
Merriam, J. C. Packard, E. L.
1896. Note on two Tertiary faunas from the BEES of ERE 1916. Faunal studies in the Cretaceous of the Santa Ana
southern coast of Vancouver Island. Univ. Calif. Mountains of Southern California. Univ. Calif. Publ.
Bull. Dept. Geol., vol. 2, pp. 101-108.
Moll, F.
1942. Die fossilen Terediniden und ihre Beziehung zu
den rezenten Arten. Paleontographica, Bd. 94, Abt.
A, Lief. 3-6, pp. 134-153, pls. 24-26.
Nomland, J. O.
1917. Fauna of the Santa Margarita beds in the north Co-
alinga regionof California. Univ. Calif. Publ. Bull.
Dept. Geol., vol. 10, pp. 293-326, textfigs. 1-2, pls.
14-20.
Bull. Dept. Geol., vol. 9, pp.
Van Winkle, K. E. H.
1918. Paleontology of the Oligocene of the Chehalis Val-
ley, Washington. Univ. Washington Publ. Geol., vol.
1, pp. 69-97, pls. 6-7.
Weaver, C. E.
1916. Tertiary faunal horizons of western Washington.
Univ. Washington Publ. Geol., vol. 1, pp. 1-67, pls
1-5.
137-159, 1 map.
A New Vexillum (Mitridae) from the Philippine Islands
by
” Jean M. Cate
Conchological Club of Southern California, Los Angeles 7, California
(Plates 1 and 2 and one Textfigure)
About a year ago I received a shipment of
Mitra species from various parts of the Philip-
pine Archipelago; among these was a specimen
from Balabac Island labelled Vexillum regina
(Sowerby, 1825) which aroused my curiosity be-
cause its color seemed to differ from that of
other specimens I had seen. Further investi-
gation brought out that there were several ad-
ditional differences between this specimen and
the typical V. regina: first, it was not sharply
shouldered; further, the surface sculpture was
far less coarse than in Sowerby's original fig-
ure of V. regina, and the color pattern was ar-
ranged in a different manner. In the course of
working out a solution as to what this species
might be, additional problems arose involving
other closely related species; these, however,
will be dealt with in a subsequent paper.
A careful search through all the known
monographs and many other papers on Mitridae
failed to turn up anything resembling the Bala-
bac species. It therefore seemed likely that
this was an undescribed taxon, but as nearly
100 years had passed since the last comprehen-
sive monograph had been compiled for the fam-
ily Mitridae, this was not an adequate basis for
describing a new species. Consequently, in ad-
dition to perusal of all available literature, as
thorough a search as possible was made among
many of the larger museums and private col-
lections to make certain that this species was
indeed new and hitherto undescribed. This
search turned up two specimens, also labelled
Vexillum regina (Sowerby, 1825), which match
my specimen fairly closely; both are from pri-
vate collections and were collected in the
southern Philippines.
Wherever it was not possible to visit col-
lections personally, Ektachrome color trans-
parencies of the dorsal and ventral aspects (4
actual size) were submitted for comparison
with other collections; without exception these
transparencies were returned promptly and
with a notation to the effect that nothing similar
existed in that particular collection. Most of
the responses stated that it superficially re-
sembled Vexillum regina (Sowerby, 1825). Re-
grettably, due to a change in personnel taking
place at the time of this search, the Mollusca
Section of the British Museum (Natural History)
was not able to compare the photographs with
the specimens in its collection; it is felt, how-
ever, that the remaining institutional and pri-
vate collections which participated represent a
good cross-section of the material available in
this Vexillum complex. For the prompt and
courteous response so willingly given in all in-
stances, I wish to express my gratitude to the
following persons and institutions cooperating:
Dr. William J. Clench and Dr. Ruth D. Turner,
Museum of Comparative Zoology, Harvard Uni-
versity; Dr. Alan Solem, Chicago Natural His-
tory Museum; Mr. William Old, Jr., New York
Museum of Natural History; Dr. Myra Keen,
Stanford University; Dr. Leo G. Hertlein, Cali-
Vol. 4; No. 1
THE VELIGER
Page 5
fornia Academy of Sciences; Dr. Joseph Rose-
water, U. S. National Museum; Dr. Robert Ro-
bertson, Academy of Natural Sciences of Phil-
adelphia; Mr. and Mrs. E. P. Chace, San Diego
Museum of Natural History; Mr. George Kana-
koff and Miss Joan Troesch, Los Angeles
County Museum; Mr. Fernando Dayrit, Nation-
al Museum, Manila; Mr. and Mrs. John Q.
Burch; Mr. and Mrs. F. K. Hadley; Mr. An-
thony d'Attilio; Mr. James Bailey; Mrs. Mary
Saul; Mrs. Z.W. Craine; Dr. Howard R. Hill;
SOU ll Cla is
Sle A
Mr. Ditlev Thaanum, and Mr. E.W. Ulrich.
With the likelihood established that a new
species was involved, further correspondence
was indicated in order to obtain as much eco-
Mr. Fernando
Dayrit of Manila, who sent the original ship-
ment of shells containing the specimen in ques-
tion, has kindly furnished from his personal
experiences and observations much additional
information pertaining to the Balabac region.
logical information as possible.
ea
AXISIGAN 1. ?~
Sonne le
Cape Melville
BAM MATEVAIC. oe Al 1
Page 6
THE VELIGER
Vol. 4; No. 1
The island of Balabac (see map), just north
of Borneo, is very sparsely populated by a
tribe known as Melebuganon Moros, numbering
altogether not over 2,000 individuals. These
people differ from the less isolated and better-
known Moros of the Sulu Archipelago in several
ways: they live in shacks at the edge of the
forest instead of in stilt-houses built high above
the water; they confine themselves to the rela-
tively small area of Balabac, seldom venturing
away from the island, whereas the Moros of the
Sulu Sea islands are more inclined to be sea-
farers. The Melebuganons prefer dry land un-
der all circumstances to any contact with wa-
ter; therefore they do no diving of any sort and
the shells they collect for food or for barter
are always taken at low tide in water no more
than knee-deep.
The favorite and most profitable collecting
locality for shells at Balabac is a small beach
to the northwest of the lighthouse at Cape Mel-
ville; the shell of the new species was collected
here on the reef, on the patches of sand between
coral heads. Fringing coral reefs border the
shoreline at this point, and at low tide a wide
area is exposed, giving easy access to the reef-
dwelling mollusks. Offshore, in the Balabac
Strait which separates the island from British
North Borneo, there is a swift current, particu-
larly during the change of tides, making naviga-
tion especially hazardous in view of the sub-
merged coral reefs. It is quite possible that this
condition is responsible for the natives' reluc-
tance to venture further afield, especially when
one realizes the nearest land was until recently
inhabited by the head-hunting savages of North
Borneo. On the other hand, a personal letter
from Mrs. Mary Saul of Kudat, North Borneo
(January 1961) states that the more adventure-
ous natives of her country frequently cross the
Balabac Strait (a distance of only a few miles)
and collect shells — quite possibly on this
Same beach at Cape Melville. For about six
months Mrs. Saul has been watching for addi-
tional specimens of the new species among the
material brought in by her collectors for barter
at Kudat, but to date she has not seena shell
Similar to the one discussed here,
The shallow coral reefs and warm tropical
water at Cape Melville furnish a suitable en-
vironment for many molluscan species. Among
these Mr. Dayrit mentions the following: Conus
nobilis Linnaeus, 1758 (''seems to be found only
in this area"), C. stramineus Lamarck, 1811, (Ge
pica Adams & Reeve, 1848, C. nocturnus So-
lander, 1786, C. omaria Bruguiére, 1792; Poli-
nices fluctuatus (Sowerby, 1825); Rhinoclavus
fasciatus (Bruguiére, 1792); assorted Terebras
and many others. Mitra species also itound liv-
ing on this reef include M, tigrina A. Adams,
1851, M. dactylus Lamarck, 1811, M. stigma-
taria Lamarck, 1811, M. episcopalis (Linnaeus)
Gmelin, 1790, M. papalis (Linnaeus) Petiver,
1767, M. pontificalis Lamarck, 1811, M. punc-
ticulata Lamarck, 1811, M. filaris (Linnaeus,
1771), Vexillum plicarium (Linnaeus, 1758), V.
corrugatum (Lamarck, 1811), and V. vulpecula
(Linnaeus, 1758). Vexillum regina (Sowerby,
1825), V. vittatum (Swainson, 1821), and V.
taeniatum (Lamarck, 1811) are also mentioned by
Mr. Dayrit as coming from Cape Melville, but
as there is presently so much confusion regard-
ing their identification, and since they are in-
cluded in the complex group under consideration
in the separate study mentioned above, I believe
it unwise to include them in the present faunal
list.
VOLUTACEA
MITRIDAE
Subfamily
VEXILLINAE
Genus
Vexillum ROpinc, 1798
Vexillum coloscopulus J. Cate, spec. nov.
(Plate 1, Figures 1, 2)
Shell long, straight, slender, fusiform,
somewhat turriculate; spire longer than the last
whorl. Protoconch lacking; teleoconch consisting
of 11 slightly convex abutting whorls; sutures
impressed, shoulders rounded. Axial sculpture
of low, fairly sharp collabral costae (about 13
on penultimate whorl) which tend to become
obsolete near outer lip; costae not regularly
aligned between sutures. Spiral ornament of
low cords, rounded, crenulated and narrow be-
low the sutures, flattened into slightly wider
bands at the periphery, again becoming rounded,
narrower and faintly granulose at lower part of
neck; all spiral cords separated by smooth,
shallow, impressed striae. Aperture straight,
siphonal canal slightly recurved; labrum thin,
simple, about 12 faint lirae within. Parietal
ridge present; columella straight, inductura
restricted adaperturally; three strong oblique
adapical columellar folds and one faint ante-
rior columellar fold. Peristome discontinuous.
Siphonal fasciole weakly produced, helicocone
nonumbilicate.
THE VELIGER, Vol. 4, No. 1 [J. Cate] Plate 1
Vexillum coloscopulus J. Cate, spec. nov.
Dorsal and Ventral Aspects of Holotype
Vol. 4; No. 1
THE VELIGER
Page 7
Color of holotype dark brick-red (Maerz &
Paul Dictionary of Color, 1950 Rev. Edit., Plate
8, L-9, Domingo Brown) with one narrow dirty-
white band centrally placed on adapical whorls,
bordered on each side by a blackish zone that
occupies most of the remainder of each whorl.
White band composed of three flat spiral
cords separated by narrow, smooth, shallowly
incised striae; its bordering black zones, ap-
proximately twice as wide, composed of five
cords. Color at sutures orange-brown where
successive whorls overlap just below the black
zone; with two wide brick-red zones, interrupt-
ed by an additional black zone, on the neck of
the shell; the brick-red of adapical band (M &
P, pl. 8, L-9) lightening adaxially to orange-red
(M & P, pl. 6, B-12, Gypsy). Final abapical
band slightly paler (M & P, pl. 7, L-12, Cal-
dera), also lightening to Gypsy Brown at colu -
mella. Color of aperture ivory, darkening
slightly toward labrum. Columella and folds
yellowish-orange. Animal of the species un-
known.
Measurements of holotype: Height, 71.4
mm.; maximum diameter, 15.8 mm.; length of
aperture, 34.6 mm.
The type locality of Vexillum coloscopulus
is here designated as Cape Melville, Balabac,
Philippine Islands (7°30' North Latitude, 117°
00' East Longitude).
The holotype will be deposited in the Geol-
ogy Department Type Collection, California
Academy of Sciences, San Francisco, Califor-
nia, where it will bear the catalog number
12'363.
The specific name is derived from the
combination of two Latin words most closely
approximating a description of the mollusk's
habitat. According to Cassell's Latin Diction-
ary, 1959 Revised Edition, the Latin word
scopulus means ''dangerous rocks in the water",
or reef; colare means ''to dwell''; therefore the
combination coloscopulus signifies a reef-
dweller. As the word coloscopulus is a mas -
culine noun in apposition, no change in ending
can be made.
Comparison of Holotype with Hypotypes:
two similar specimens, kindly loaned for this
study, are designated as hypotypes rather than
paratypes since they are not from the type lo-
cality.
(Plate 2, fig. 2) Collection of Mr.
and Mrs, John Q. Burch, Los An-
geles, California. Locality: Sulu
Archipelago.
Hypotype 1.
The
Hypotype 2. (Plate 2, fig. 3) Collection of Mr.
Anthony d'Attilio, Valley Stream,
New York. Locality: Zamboanga,
Philippine Islands.
Table 1: Measurements of Holotype and Hypotypes
(in millimeters)
Height Maximum Length of
Diameter Aperture
Holotype 71.4 15.8 34.6
Hypotype 1 68.1 17.6 35-5
Hypotype 2 55-3 15-5 28.9
All three specimens agree very closely in ~
general shape and proportions, the holotype,
however, being the most slender. Morphologi-
cally they are all very similar, even to being
decollate, as are all available specimens of the
other species in the complex which includes
Vexillum regina.
Hypotype 2 matches the holotype exactly in
the distribution of its color pattern, the narrow
white band being placed in the center of each
whorl and the proportions of each color-zone
matching those of the holotype as well. Hypo-
type 1 has a somewhat more coarse, blurred
appearance than either of the other two speci-
mens and is possibly a shell from an older ani-
mal; its black zones are better defined as black
bands, narrower than in the other two speci-
mens and revealing more of the brick-red color
between. In Hypotype 1 the narrow white band
is not so centrally placed on the apical whorls;
in fact, in the first five whorls this white band
abuts the lower suture and no black zone is ap-
parent there. It does approach the central po-
sition more closely, however, on the penulti—
mate and antepenultimate whorls, whereas in
other species of this complex the white zone is
always seen adjoining the sutures abapically.
The color of all three specimens differs very
little; the holotype is the darkest in color, but
it is possible that a certain amount of perios-
tracum remains; it was deemed better not to
attempt a further cleaning in order to ascertain
whether an epidermis was present. It is likely
that further cleaning might render the white
band a purer white but would not otherwise
change the specimen's appearance materially.
Hypotype 2, the smallest of the three shells, is
the lightest in color, its reddish bands more
nearly approaching a creamy-yellow shade.
Both hypotypes possess a heavier labral edge
than the holotype, which is generally more del-
icate in appearance than the others.
Page 8
THE VELIGER
Vol. 4; No. 1
Comparison of Vexillum coloscopulus with V.
regina (Sowerby, 1825): The following six spe-
cimens of Vexillum regina were used in this
comparative study, all approximately the same
size as the holotype of the new species:
1. Collection of Anthony d'Attilio, New York;
locality, Zanzibar
2. Collection of E. W. Ulrich, Long Beach,
California; locality, Andaman Islands
3. Collection of James Bailey, Los Angeles,
California; locality, China
4. Collection of Los Angeles County Museum,
No. A2'777, Los Angeles, California; lo-
cality, Moluccas
5 and 6. Collection of Stanford University
Paleontology Department, lot No. 1'660; lo-
cality, Moluccas
LEY:
Vexillum coloscopulus and V. regina re-
semble one another only superficially, upon
close inspection. Both species possess spires
longer than the last whorl, although this charac-
teristic is somewhat more pronounced in VY. re-
gina. Both species possess four columellar
folds, arranged in a similar fashion. Other-
wise, except for the faintly similar coloration
and arrangement of color pattern, the features
they possess in cOmmon are generic characters
and could apply equally to many of the other
species of Vexillum; i.e., axial costae, pro-
duced spire, spiral ornament, and so on.
Vexillum coloscopulus differs from V. re-
gina (Sowerby, 1825), its most closely related
described ‘species, in the following ways, tabu-
lated for easier comparison;
Table 2: Comparison of Characters
Characters compared Vexillum regina (SOWERBY) Vexallum coloscopulus spec. nov.
angular, gradate
extremely coarse, elevated
Sutural ramp
Axial costae
Spiral ornament
Spire
Aperture
Parietal lip
Neck of shell
Pseudumbilicus
Canal
Labral lirae
coarse, rough
turriculate
constricted
well defined
faintly produced
sharply recurved
roughly granulate
about 3 raised lirae
rounded
smooth, somewhat flattened
smooth
fusiform
straight
lacking
nearly smooth
lacking
slightly recurved
about 12 faint lirae
brick-red
Color
White band on last whorl
White band placement
White band bordered by
White band ornamentation
bright orange, or blackish
one wide, one narrow
contiguous to sutures
narrow black lines
central yellow thread
one narrow band only
central on adapical whorls
wide black zones
lacking
Aperture color white
Acknowledgment
I should like to express my sincere appre-
ciation to all those who aided me in so many
ways in the preparation of this paper: first,
Mr. Fernando Dayrit of Manila, who sent the
holotype specimen and provided background in-
formation on the ecology of the new species;
Dr. Myra Keen, Mr. George Kanakoff, Mr.
Tony d'Attilio, Mrs, John Q. Burch, and Mr. E.
W. Ulrich for generously allowing me the use
of comparative material; Dr. Rudolf Stohler
for his unfailing kindness and helpful sugges-
tions, and particularly Crawford Cate for his
patience and helpfulness in countless ways.
cream to dark ivory
The map of the type locality was furnished
by Mr. Fernando Dayrit and adapted for use as
a textfigure by Mrs. Emily Reid of the Veliger
staff.
The terminology employed in the descrip-
tion of the new species was derived from the
1960 Treatise on Invertebrate Paleontology,
Part I, Mollusca 1, pp. 106-135.
The photographs of Vexillum regina were
furnished through the courtesy of Stanford Uni-
versity. All other photographs are by Pierson,
Oswald and Pierson. The color reproduction
for Plate 1 was done by the California Academy
of Sciences from Kodachrome slides.
my
Tue VELIGER, Vol. 4, No. 1 [J. CaTE] Plate 2
Figure 2 a Figure 3 a
Figure 1 b Figure 2 b Figure 3 b Figure 4 b
Figures 1, 2, 3: Vexillum coloscopulus J. CATE, spec. nov.
Figure 1: Holotype. Figure 2: Hypotype No. 1. Figure 3: Hypotype No. 2.
Figure 4: Vexillum regina (SOWERBY, 1825)
(a: dorsal aspects; b: ventral aspects)
Vol. 4; No. 1
THE VELIGER
Page 9
What is Anatina anatina?
by
A. Myra KEEN
Stanford University, California
(Five Textfigures)
The generic name Anatina (literally, pertain-
ing to a duck) has been applied in molluscan lit-
erature to two unrelated bivalve groups that
look alike because they have a posterior gape
surrounded by an extension of the shell shaped
a little like a duck's beak. Lamarck used the
French equivalent, ''Anatine', as early as 1809,
but he did not Latinize it until 1818 (Hist. Anim.
sans Vertébres, vol. 5, p. 462). Tautonymy
would fix the type species — Solen anatinus
Linnaeus, 1758, a nacreous-shelled form re-
lated to Periploma. In 1817 Schumacher had
proposed the name Anatina for Mactra anatina
Spengler, 1802, a mactrid clam with a thin, por-
celaneous shell. As tautonymy was frowned
upon in those days, Schumacher renamed the
species Anatina pellucida and provided an illus-
tration. Some attempt has been made to rescue
Lamarck's usage by dating it from 1816, when
Bosc cited Anatina Lamarck in proper Latin
fashion (Nouvelle Dictionnaire d'Histoire Natu~
relle, vol. 1, p. 492), but this falls short of
validation because the only species mentioned,
"le solen canard", is a nomen nudum and also
in the vernacular. Even if one could salvage
Lamarck's Anatina in this way, one still would
find obstacles preventing its use. Rodding in
1798 had proposed the name Laternula, which
has for type Solen anatinus Linnaeus (by subse-
quent designation of Gray, 1847). Also, Auri-
scalpium Megerle von Miihlfeldt, 1811, is based
on the same species. Thus, if the name
Anatina is to survive at all, it must be as of
Schumacher's usage, for a genus in Mactridae.
The substitute name Labiosa Miller, 1832 (ex
Schmidt MS), has been favored by some authors,
a name proposed when Anatina Schumacher was
considered homonymous. Letting both uses of
Anatina lapse and accepting Labiosa still would
not solve the problem of interpreting the type
species, for the type of Labiosa necessarily is
Mactra anatina Spengler. Let us, therefore,
examine this form.
Mactra anatina was briefly described by
Spengler thus: ''Testa diaphana, transverse
striata, vulva hiante, ano planato.'' There was
a longer description in Danish but no figure.
The length was stated to be azlspbreadthi2saiy,
The shell was said to be a rare form, from
South America. Fifteen years later, Schu-
macher, who lived in Denmark, mentioned in
the introduction to his Essai (pp. 16-17) that,
writing at his residence in the country, he had
to rely on his own private collection but that he
had opportunity to compare his material with
that of Spengler in Copenhagen, especially for
such groups as Mactra, on which Spengler had
published. The implication is, then, that Schu-
macher's figure of Anatina pellucida (see fig.
la-b) illustrates either Spengler's holotype or a
shell very much like it. An inquiry addressed
to Dr. Henning Lemche brought the reply that
Spengler's material still is in Copenhagen, now
at the Universitetets Zoologiske Museum. Dr.
Lemche (letter dated April 13, 1961) resolves
the problem of identifying the holotype bya
tracing of the outline of the shell (see fig. 2)
and by quoting the label, which he says is in
Mérch's handwriting: ''M. anatina Spengler/
Anatina pellucida Sch.: 125. VIII, f. 1. Origi-
nal.'' The traced outline compares almost
exactly in size and shape to the figure given by
Schumacher (length, 41 mm.). It is in the pro-
Mnabna pdllucida.
Figure 1: Reproduction of Spengler’s original figure of
Anatina pellucida.
Page 10
THE VELIGER
Vol. 4; No. 1
Figure 2: Tracing of outline (courtesy of Dr. H.
Lemche) of the holotype of Anatina anatina (SPENGLER).
Length, 41 mm.
Figure 3: Outline of a shell from Florida of compar-
able size (Stanford University collection).
Figure 4 (Inner figure): Outline of right valve,
specimen from off Guaymas, Mexico (Stanford
University collection). Length, 35 mi.
Figure 5 (Outer figure): ‘Tracing of outline of
Mactra cyprinus, holotype, as figured by Wilkins, 1957.
Length, 70 mm.
portion of 33 to 23, and one can thus infer that
Spengler's ''T'' was a measure of length ap-
proximately one-half inch long. Spengler's
Mactra anatina and Schumacher's new name for
it seem in the main to have been ignored by
other nineteenth century workers. Dall, how-
ever, in 1894 (p. 41), without giving evidence
for his statement, cited the species as occur-
ring in West Mexico. He listed as synonyms
"Lutraria papyracea (Lam.) Sowerby, 1824" and
Mactra cyprinus Wood, 1828. Accepting Dall's
identification, I utilized the name Anatina ana-
tina, taking as illustration Reeve's excellent
figure of Wood's Mactra cyprinus (Keen, 1958,
p. 159, fig. 363). When first described this
latter shell was thought to have come from Pe-
ru, but Reeve felt uncertain as to the locality
and merely said, ''Hab.---?"'. It is fortunate
that I overlooked a discussion of the Cracherode
collection by Wilkins (1957, p. 164), where the
holotype of this form was refigured, with the
conclusion that it is conspecific with Labiosa
lineata (Say) and was actually from the Carib-
bean. The specimen is in the collection of the
British Museum (Natural History) (see fig. 5).
It had been purchased in 1797 by the Rey.
Cracherode (for whom the abalone Haliotis
cracherodii was named), having formerly come
from the Calonne collection. Wilkins con-
curred in the judgment of previous British Mu-
seum curators, suchas E. A. Smith, that the
differences in outline between this specimen
and a normal L. lineata were a matter of rela-
tive maturity and also the result of an injury
that had given this shell an unusually wide gape
— in other words, that the degree of difference
could be within the range of variation of the
species. Wilkins, of course, had no reason to
explore synonymic history prior to 1822, the
date of Say's name lineata.
So long as no authentically-located West
American specimens were available, one had
no basis for questioning the conclusions of
workers like E.A.Smith, Dall, or Wilkins,
even though these conclusions were not in com-
plete agreement. The existence of a problem
came to light in the summer of 1960 when three
fragmentary specimens of an Anatina (two un-
matched opposite valves and the upper halves
of two matched valves) were dredged by the
Ariel Cruise at two stations in the Gulf of Cali-
fornia — off Cabo Haro, near Guaymas, depth
15-25 fathoms, and off Carmen Island, 15-20
fathoms. These specimens corresponded al-
most exactly in size and outline with the illus-
tration of Anatina anatina in my book, that fig-
ure having been reproduced at approximately
one-half the size of Reeve's original. We there-
fore identified the find on the spot as Anatina
anatina. Later, when I had returned to Stan-
ford and had compared the specimen with Schu-
macher's figure, 1 be came increasingly per-
plexed and now feel obliged to raise the ques-
tion of whether authors have not been in error
as to the synonymies for both the Caribbean and
the West Coast species of Anatina.
Literature aside, a comparison of 14 avail-
able East Coast specimens (for a sample out-
line, see fig. 3), from Florida and Brazil, with
the newly-found West American material re-
veals the following consistent differences:
Vol. 4; No. 1
WEST COAST
No radial sculpture, except for the rib bound-
ing the siphonal area
Concentric sculpture weak, fine
Umbones broad, not pointed
Anterior dorsal margin smooth, even
Posterior end longer,53 percent of total length
Posterior dorsal margin angular
Posterior gape wide
Ventral margin nearly straight
Proportion of height to length, 68 percent
These differences are sufficient to distinguish
the two species even if there were no demon-
strable geographic separation, being consistent,
with little variation among the available speci-
mens.. Re-examining the type illustrations one
sees that all of the characteristics listed above
for the West Coast form are evident in the pho-
tograph of the holotype of Mactra cyprinus
Wood published by Wilkins. The only real dif-
ference between this specimen and the Gulf of
California material (see fig. 4, inner outline) is
in size, for Wood's specimen is about twice as
long as the others which possibly may be juve-
niles. To my mind, this is good evidence that
Wood's specimen came from the West Coast.
Whether it was actually from Peru remains to
be seen, but surely it must have been from the
Panamic province. Support for this argument
came while the present paper was in prepara-
tion: Olsson (1961) figures a single somewhat
broken valve he had collected at Santa Elena,
Ecuador. It is 42 mm. long, thus slightly
larger than the specimens from the Gulf of Ca-
lifornia. It exhibits all of the ''West Coast'"'
characteristics listed above.
Schumacher's illustration does not show the
exterior sculpture, but otherwise all the char-
acteristics cited under ''East Coast'' above ap-
ply to this shell. I would therefore suggest that
the following revised synonymy be considered.
It represents, of course, a radical departure
from established convention. It is offered with
a query, in the hope of stimulating further in-
vestigations.
THE VELIGER
Page 11
EAST COAST
Fine radial striae and wrinkles, especially in
front of rib
Concentric sculpture apparent, coarse
Umbones narrow, pointed
Anterior dorsal margin flared, set off by a ra-
dial depression
Posterior end shorter, 47 percent of length
Posterior dorsal margin curved
Posterior gape moderate
Ventral margin evenly curved
Proportion of height to length, 72 percent
Anatina anatina (Spengler, 1802)
Mactra anatina Spengler. Skrifter af Naturh. -
Selsk., Bd. 5, Heft 2, p. 126. ''South Ameri-
ca''; Mérch, 1870. Malak. Bl., vol. 17, p. 124,
Anatina pellucida Schumacher, 1817.
test., p. 125, pl. 8, fig. 1.
Lutraria lineata Say, 1822. Jour. Acad. Nat.
Sci. Philadelphia, vol. 2, pt.2, p. 310. South-
eastern United States.
Lutraria sp. Sowerby, 1824. Genera of shells,
pt. 24, pl.7. Loc. --? (Compared to but not
identified with L. papyracea Lamarck, a
composite species).
Mactra recurva Wood, 1828. Index Test., Suppl.,
p. 4, pl. 1, Mactra, fig.2. South Carolina.
"Mactra nuttallii Conrad" of Reeve, 1854.
Conch. Icon., vol. 8, Mactra, fig. 125 (not of
Conrad, which is a Schizothaerus).
Ess. vers
Geographic distribution: East Coast of the
United States, from New Jersey southward; Gulf
of Mexico; northern South America to Brazil.
Anatina cyprinus (Wood, 1828)
Mactra cyprinus Wood. Index Test., Suppl.,
p. 4, pl. 1, Mactra, fig.1. "Peru"'.
————— Reeve, 1854. Conch. Icon.,
Mactra, sp. 37.
"Labiosa anatina (Spengler)" of Dall, 1894.
Nautilus, vol. 8, no. 4, p. 41. 'West Coast of
Mexico",
vol. 8,
of KLamiy, 1918 (in part). Jour. —
Conchyl., vol. 63, no. 4, p. 349. (With exten-
sive synonymy and citation of two specimens
Page 12
THE VELIGER
Vol. 4; No. 1
ee SSeS
in the Paris Museum, presumably from the
West Coast but without exact locality.)
"TLabiosa lineata (Say)"' of Wilkins, 1957. Bull.
British Mus. (Nat. Hist.), Historical Series,
vol. 1, no. 4, p. 164, pl. 24, fig. 9. (Refigured
holotype of Mactra cyprinus.)
"Anatina anatina (Spengler)'' of Keen, 1958. Sea
Shells of Tropical West America, p. 158, fig.
363. (Reproduction of Reeve's figure.)
"TLabiosa anatina (Spengler)'' of Olsson, 1961.
Mollusca of the Tropical Eastern Pacific, p.
333, pl. 57, figs. 3, 3a. Santa Elena, Ecuador.
Geographic distribution: Gulf of California,
from off Guaymas and Carmen Island, to Santa
Elena, Ecuador, possibly to Peru.
In a review of Spengler's collection, Mérch
(1870) gives evidence that further strengthens
the case for Spengler's holotype having been an
East American shell as he says that Spengler's
principal sources for material were the Danish
colonies in the East and West Indies and else-
where. There were no Danish colonies on the
West Coast of the Americas. Of course, Speng-
ler could have purchased imported shells, as
did other wealthy. collectors. Even if we accept
the premise that his holotype was from the East
Coast, the puzzle remains as to what collector,
prior to 1800, could have found so rare a shell
as Anatina cyprinus (and especially so large an
individual as the holotype) in Peru or anywhere
else along the West Coast. The Calonne col-
lection was worldwide in scope and did contain
other rare items from western South America.
We must assume, therefore, that some enter-
prising early sailor conveyed it back to Europe,
where it has graced three collections and sur-
vived two World Wars.
Acknowledgment
I wish to thank Dr. Henning Lemche of the
University at Copenhagen for his prompt and
effective reply to my inquiry about Spengler's
type specimens, Dr. Leo G. Hertlein of the
California Academy of Sciences for consulting
a reference unavailable to me, and my col-
leagues who have encouraged me to write down
these observations instead of merely talking
about them. Also, I should not forget to thank
the members of the Ariel Cruise, who made the
discovery of these specimens in the Gulf of
California possible.
Literature Cited
Dall, William Healey
1894. Synopsis of the Mactridae of North America.
Nautilus 8 (4): 25-28; (5): 39-43.
Keen, A. Myra
1958. SeashellsoftropicalWestAmerica. xit+624 PP.»
illus. Stanford Univ. Press, Stanford, Calif.
Lamy, E.
1917-1918.
d'Histoire Naturelle de Paris.
(3): 173-275; (4): 291-411;
Mérch, O. A. L.
1870. Ubersicht der von Lorentz Spengler beschrieben-
en Conchylien. Malakozool. Blater 17: 99-124.
Olsson, A. A.
1961. Mollusks of the tropical eastern Pacific.
Ithaca, N. Y. 572 pp., 86 pls.
Schumacher, H. C. F.
1817. Essay d'un nouveau systéme des habitations de
vers testacés. Copenhagen. iv+287 pp., 22 pls.
Spengler, Lorenz ;
1802. Beskrivelse over det toskallede Conchylie-slaegt
Mactra. Skr.Nat.Selsk., Copenhagen, 5(2): 92-128.
Wilkins, G. L.
1957. The Cracherode tell collection.
Mus, (Nat. Hist.), Historical Ser.,
pls 20-25.
Révision des Mactridae vivants du Muséum
Jour. Conchyl., 63
2 pls.
1g 3h
Bull. British
1 (4): 121-184,
Vol. 4; No. 1
THE VELIGER
Page 13
A New Whale Barnacle from Late Pleistocene Deposits
at San Quintin Bay, Baja California
by
Vicror A, ZULLO
Department of Paleontology, University of California, Berkeley 4, California
(Plate 3)
Two isolated compartmental plates of a new
species of the whale barnacle Cryptolepas Dall,
1872, were collected from late Pleistocene de -
posits at San Quintin Bay, on the Pacific Coast
of Baja California. This new species repre-
_ sents the only known fossil record of the genus
Cryptolepas, which until now has included only
the living North Pacific species, C. rhachian-
ecti Dall, 1872. The new species closely re-
sembles C, rhachianecti, but the lamellar ribs
of the shell of the former terminate distally in
T-shaped flanges which form an outer wall (pl.
2, figs. 3, 4). Im this respect, the new species
resembles the genus Coronula Lamarck, 1802.
The whale barnacles including the genera
Coronula, Cryptolepas, Tubicinella Lamarck,
1802, and Xenobalanus Steenstrup, 1851, form a
morphologically allied group within the sub-
family Coronulinae Leach. All of the species
of these genera live partially or wholly embed-
ded in the skin of cetaceans, as opposed to the
platylepadid group of coronulines which have a
diversity of hosts, but have never been found
on cetaceans, The whale barnacles can be dis-
tinguished morphologically from other coronu-
lines by the presence of an oral hood covering
the cirri, and by the absence of the internal
midribs, median parietal sulci, and parietal
furrows which characterize the platylepadid-
group.
Within their own group, the whale barnacles
have attained a great deal of morphological di-
versity, apparently as a result of their individ-
ual habitats. The fossil record of the coronu-
lines, hitherto restricted to species of the ge-
nus Coronula, is sparse, but the world-wide
distribution of these fossils in deposits no older
than late Miocene indicates that the whale bar-
nacles have achieved their present diver sifica-
tion in a relatively short period of time.
Apparently Cryptolepas, Tubicinella, and Xe-
nobalanus developed from Coronula, but until
now, no intermediate forms have been found to
substantiate this hypothesis. The lack of fossil
specimens of these genera is evidently a result
of the fragile nature of their shells. The shell.
of Coronula, which is only partially embedded
in the skin of the whale and therefore exposed
to the surrounding environment, is tough and
stout, and difficult to disarticulate or break.
However, the shells of Cryptolepas, Tubicinel-
la, and Xenobalanus, which are totally buried
in the whale's skin and are not exposed to the
surrounding environment are friable and easily
disarticulated or broken and would probably be
reduced to fragments before being preserved,
This theory is demonstrated by the new fossil
species whose shell, although closely resem-
bling that of Cryptolepas rhachianecti in most
features, is structurally more similar to the
shell of Coronula.
Pilsbry (1916, p.279) must have envisioned
such an intermediate form as represented by
the San Quintin specimens in his statement con-
cerning the origin of Cryptolepas rhachianecti:
"With the exception of the grooves of the
sheath, nearly all of the differences from
Coronula are degenerative changes appar-
ently correlated with the protected station
of the animal embedded in the skin of the
host. An outer wall is no longer needed.
The branches of the ribs, which in Coro-
nula serve as buttresses, have become
short and extremely variable."'
Pilsbry also noted that among the species of
Coronula, Cryptolepas rhachianecti most close-
ly resembles Coronula (Cetopirus) complanata
(Mérch, 1852) in its depressed exterior outline
and cylindric body chamber, and he concluded
Page 14
THE VELIGER
Vol. 4; No. 1
that Cryptolepas had been derived from such an
ancestor. The discovery of compartmental
plates of Coronula complanata in late Pliocene
deposits at Cape Blanco, Oregon, supports
Pilsbry's conclusion, although there are no au-
thenticated Recent records of this species in
the North Pacific.
The new fossil species of Cryptolepas, there-
fore, appears to be an intermediate, having all
the characteristics of Cryptolepas, but retain-
ing the outer wall of its Coronulid ancestor.
Family BALANIDAE Gray, emended
Subfamily CORONULINAE Leacu, emended
Genus Cryptolepas DALL
Cryptolepas Dall, 1872, Proc. Calif. Acad.
Sci., vol. 4, p. 300; Pilsbry, 1916, U. S. Nat.
Mus. Bull. 93, p. 278; Kruger, 1940, in Bronn's
Klass. u. Ordn. des Tierreichs, Bd. 5, Abt.1,
Buch 3, Teil 3, p. 453.
Cryptolepas rhachianecti Dall
(by monotypy). Living on the California gray
whale, Rhachianectis glaucus Cope. Subsequent
authors (e.g., Pilsbry, 1916, p. 279; Cornwall,
1955, p. 44) have not retained the first "h'' in
the spelling of the specific name.
Type species:
Range: Late Pleistocene, Baja California;
Recent, North Pacific.
Cryptolepas murata ZULLO, spec. nov.
Dimensions: Holotype Paratype
UCMP no. UCMP no.
34676 34677
Height: 10.3 mm. 12.1 mm.
Width of ribs: 8.4 mm. 10.1 mm.
Width of base of ribs: 11.1 mm. 12.5 mm.
Description: Two worn lateral compart-
mental plates; six parietal ribs per compart-
mental plate formed by four complete radial la-
mellar folds and two sutural half-folds; ribs
unbranched or with one or two short, free-
ending branches and terminating in T-shaped
flanges forming a more or less complete outer
wall; ribs composed of outer and imer lamellae;
inner lamellae separated by closely-spaced,
longitudinal septa, forming longitudinal tubes;
Ornamentation of ribs consisting of fine,
closely-spaced, vertical striae crossed by dis-
tantly-spaced growth wrinkles; growth wrinkles
prominent inapical half and fading in basal half;
sheath transversely grooved, three-fourths
height of shell; basal edge of sheath not pro-
jected, but nearly flush with lamellar folds be-
low; sutural edges of radii intricately crenu-
lated; opercular plates unknown.
The specific name is derived from the Latin
"muratus'' signifying ''walled".
Holotype: University of California, Museum
of Paleontology, no. 34'676 (pl. 3, figs. 1, 2,
3); paratype: UCMP no. 34'677 (pl. 3, figs. 4, 5).
Occurrence: UCMP locality no. A-8677.
Late Pleistocene, San Quintin Bay, Baja Cali-
fornia. One mile south of San Quintin pier at
base of cliffs. This locality is identical to that
described by Jordan (1924, p. 243). Jordan
listed 225 species of invertebrates from this
locality, including 13 not known to be living and
several whose northern limits are presently
south of this locality. It was concluded that this
fauna could be correlated with the late Pleisto-
cene warm water San Pedro fauna of Southern
California.
Discussion: Cryptolepas murata differs from
the living C. rhachianecti in the presence of an
outer wall formed by T-shaped flanges at the
ends of the ribs, and in the less complex
branching of the ribs. Pilsbry (1916, p. 280)
noted that a few individuals of C. rhachianecti
he examined had one or more compartments
with only three or four loops of the inner wall,
and with rib branches parallel with and close to
the upper sides of the ribs, forming a second-
ary, incomplete ''outer wall'’. The terminal
flanges of C. murata, however, are homologous
with the terminal flanges forming the outer wall
in the genus Coronula.
Darwin (1854, p. 413) observed that the shell
of Coronula complanata is more completely
buried in the skin of the whale than are the
shells of other species of Coronula. Probably
Cryptolepas murata was also deeply buried, ap-
proaching the habit of C. rhachianecti. The
body chamber was probably cylindric with its
base equal in size to its orifice. The exterior
outline of the shell would therefore be approxi-
mately that of C. rhachianecti (pl. 3, figs. 6, 7).
Literature Cited
Cornwall, I. E.
1955. The barnacles of British Columbia. Brit. Co-
lumbia Prov. Mus., Dept. Educ., Handbook no. 7,
pp. 1-69, figs. 1-9.
Darwin, C.
1854. A monograph on the sub-class Cirripedia, Balan-
idae, Verrucidae. Ray Soc. London, 684 p., 11] figs.,
30 pls.
Jordan, E. K.
1924. Expedition to Guadalupe Island, Mexico, in 1922.
No. 4. Molluscan fauna of the Pleistocene of San
Quintin Bay, Lower California. Proc. Calif. Acad.
Sci., fourth ser., 15 (7): 241-255, textfig. 1, pl. 25.
Pilsbry, H. A.
1916. The sessile barnacles (Cirripedia) contained in
the collections of the U. S. National Museum; including
a monograph of the American species. U. S. Nat.
Mus. Bull. 93, xi, 366 p., figs. 1-99, pls. 1-76.
Tue VELIGER, Vol. 4, No. 1 [ZuLto] Plate 3
Be ac
Figure 1
Figure 6 Figure 7
Figures 1 to 5: Cryptolepas murata ZULLO, spec. nov., x 3
Figure 1: interior view of holotype UCMP no. 34676; Figure 2: outer
wall of the same; Figure 3: basal view of the same, showing terminal
T - shaped flanges; Figure 4: basal view of paratype UCMP no. 34677,
showing branching lamellar ribs; Figure 5: sutural edge of radius of
holotype UCMP no. 34676. Figures 6 and 7: Cryptolepas rhachianecti
DaLL, hypotpye UCMP no. 34678. Figure 6: basal view, x 1.4.
Figure 7: oral view, x 0.9.
: ae
rte oot
prance
o “
OR > cep oneal pa
Vol. 4; No. 1
AE WEEIGER
Page 15
A Statistical Study in Cowries:
The Size of Mauritia arabica (Linnagvs)
by
F. A. SCHILDER
University of Halle (Saale), Germany
(2 Textfigures)
Mauritia arabica (Linnaeus) ranges from
the Red Sea and Natal to Japan and the Samoan
Islands, if we include the well separable East
African race immanis Schilder & Schilder, as
well as the distinct species M. grayana Schilder
which replaces M. arabica in the Red Sea and
in the Persian Gulf. The other allied species,
however, as M. eglantina (Duclos), histrio,
(Gmelin), maculifera Schilder, depressa (Gray),
etc., will not be discussed in the present paper
as they belong to other superspecies (Schilder,
1947).
For forty years we have accumulated ac-
curate notes on more than one hundred thousand
cowry shells examined by us personally; they
include almost 3,000 Mauritia arabica coming
from about 350 localities.
We have stated the exact length of each
shell in tenths of a millimeter (using a vernier);
in this paper, however, the mean length of the
shells coming from each locality or geographi-
cal area has been expressed in millimeters,
As the standard deviation in these series of
shells generally is about 4 to 7 mm., the mean
error of their average length mostly is about
+1mm., never exceeding +2 mm.
The following list contains 55 habitats from
which we have examineda significant number
of Mauritia arabica personally; there are a few
specific localities from which we received
plenty of shells, and numerous extended areas
containing several adjacent localities from
which the number of specimens was too small
to be treated separately. The geographical
names are preceded by the average length of
shells in millimeters, and they are followed by
three figures: the first figure indicates the
number of examined specimens divided by ten,
s0 that, e.g., 3 is equal to about 25 to 35 shells;
while the second figure indicates the number of
different localities included in the area (inde-
pendent collectors said to have collected at the
same ''locality'' have been treated as different
localities also). The average length of speci-
mens living in each area can be estimated more
accurately by a few shells coming from several
localities than by numerous shells coming from
one locality only in which the size of the shells
may be influenced by an unusual environment.
The third figure (in parentheses) indicates the
average temperature (in degrees Centigrade) of
the surface of the sea in the coldest month
(February or August).
Mauritia grayana:
52 Aqaba — Ras Benas 2/10 (21°)
44 Jidda — Assab 2/6 (25°)
42 "Red Sea'' (no locality) 5/8 (25°)
40 Perim — Berbera — Obbia 3/11 (25°)
49 Aden 2/9 (23°)
59 Muscat — Persia 2/4 (21°)
63 Karachi 3/5 (21°)
43 Seychelles — Mauritius (The occur-
rence needs confirmation, see Schilder
& Schilder, 1939; Allan, 1956.) 2/5 (23°)
Mauritia arabica immanis:
74 Mogadishu — Delagoa Bay 3/11 (24°)
13 Wereul 1/5) (23>)
68 Madagascar 2/12 (23°)
66 Réunion -- Rodriguez 2/9 (22°)
72 Seychelles 1/3 (25°)
Mauritia arabica arabica:
59 Bombay — Malpé 1/2 (25°)
47 C. Comorin — Pamban — Galle 2/6
(26°)
58 "Ceylon'' (no locality) 5/5 (27°)
56 Trincomali 5/4 (27°)
58 Madras -- Waltair 1/5 (26°)
55 Mergui Archipelago 1/2 (27°)
47 Penang (Griffiths, 1956) 5/9 (28°)
52 Andaman Islands 6/8 (277)
45 Atjeh — Nias — Oosthaven 7/16 (28°)
44 Labuan — Wijnkoopsbay 1/4 (27°)
Page 16
THE VELIGER
Vol. 4; No. 1
42
40
46
47
50
50
54
58
53
45
44
36
39
44
43
45
61
55
42
41
45
44
38
40
43
bys}
56
55
49
Tjilaut Eureun (Schilder and Schilder,
1934) 54/1 (27°)
Tjilatjap — Sumbawa 2/7 (27°)
Tiger Islands — Macassar — Kutei 2/6
(27°)
Northcoast of Java 6/9 (27°)
Belitong — Singapore 1/6 (27°)
Siam Gulf — Pakhoi 1/7 (23°)
Hong Kong — Amoy 2/6 (14°)
Tokyo — Shikoku 1/3 (13°)
Ryukyu Islands — Taiwan 2/8 (20°)
Philippine Islands 4/12 (27°)
Sangi Islands — Mapia Island 1/2 (27°)
Ternate 7/1 (27°)
Menado 1/2 (27°)
Busak (N. W. Minahassa) 2/1 (27°)
Oly <> Ew = Banca S/I2 (27°)
Kaimana — Kei — Aru 3/4 (26°)
Port Essington — Broome 1/2 (25°)
Sydney — Torres Straits 1/7 (20°)
Geelvink Bay — Huon Gulf 2/7 (28°)
Purdy Islands, Admiralty Islands 2/2
(28°)
N ew Britain (Schilder and Schilder,
1937): Bitokara 3/2 (28°)
id.: Ulamona 4/2 (28°)
id.: Mope — Iltishuk 46/6 (28°)
ide: (Karlei i/iN(2e°)
Solomon Islands — Santa Cruz Islands
4/8 (28°)
New Caledonia 4/16 (23°)
Fiji Islands -- Tonga Islands 2/5 (24°)
Samoan Islands? ? (Schilder, 1958) 3/1
(Zit)
Samoan Islands 2/7 (27°)
48 Wallis Island — Marshall Islands 2/7
(28°)
49 Palau Islands — YapIsland 2/6 (27°)
49 Guam, Marianas Islands 1/4 (26°)
According to this list, the average length
of Mauritia arabica varies in various areas
from 36 mm. (Ternate) to 78 mm. (Natal). We
can afford a general view of these figures, if
we reduce them into classes differing from each
other by 5 mm. (e.g., class 40 embraces 38 to
42 mm., class 45 embraces 43 to 47 mm., etc.).
Then we express these classes by visually im-
pressible signs so that darker signs and tri-
angles indicate larger shells than plain and
round signs; we enter them on a map (fig. 1)
from which we can learn the following interest-
ing facts:
1, The smallest Mauritia arabica (classes 40
and 45 mm.) inhabit all areas between the
Solomon Islands, the Philippine Islands,
and Western Sumatra; this central zone
can be indistinctly traced as far as to the
Southern Red Sea.
2. On the Northern and Southern border of this
equatorial zone the average size becomes
larger (Bergmann's rule, Schilder, 1956);
the gradual increasing of size in Mauritia
arabica towards the polar confines of its
distribution can be followed most distinctly
from Singapore to Japan and from Berbera
in both directions towards Aqaba and Kara-
chi.
040 ©45 @50 955 @60 A65 A7O A775 AsO
Figure 1
Vol. 4; No. 1
THE VELIGER :
Page 17
3. In addition, there is also a distinct increas-
ing of size from the Malayan and Melane-
sian region towards the Eastern and West-
ern borders of the habitat of Mauritia ara-
bica arabica, i.e., towards Polynesia and
India, so that the central zone around the
Moluccas with small M. arabica becomes
totally encircled by a zone with larger
ones.
4. The North Western Mauritia grayana gen-
erally agrees in size with the Eastern M.
arabica arabica, but the South Eastern race
of the latter, M. arabica immanis, is ex-
tremely large; its size gradually increases
from the Mascarene Islands to the African
coast, and attains its maximum on the
South Western border (Natal).
5. The North West Australian Mauritia arabica
also seem to be much larger than one would
expect, but the material available is too
scanty for a definite statement; most Maur-
itia coming from this region belong to M.
eglantina (perconfusa) and M. histrio
(westralis).
These facts can also be shown by plotting
the size against the winter temperature of the
areas (fig. 2). The general ecological influence
increasing the size in colder waters is modified
by the probably genetical enlargement of shells
gray | imm.|
ae E we |
Figure 2
in the central Pacific (nameless) and in the In-
dian race (dilacerata Schilder and Schilder)
which becomes far surpassed in the East Afri-
can Mauritia arabica immanis. Even in M.
grayana there seems to bea racial difference
between the Western and the Eastern popula-
tions {see Schilder and Schilder, 1939; the mean
size of M. grayana from the Red Sea (Aqaba to
Berbera) and from the Persian Gulf (to Karachi)
is 43.8 + 0.74 mm. and 61.8+1.32 mm., respec-
tively; the difference is significant (P < 0.001)}.
Annex. The correlation between the length
of the shells and their relative breadth (i. e., the
maximum breadth expressed in percent of the
length) may be shown by the following table con-
cerning 154 adult Mauritia arabica from Tjilaut
Eureun:
Length
30 35 40 45 50 55 60 65
es
ee ly 5) Sh gl
DEE RGOR Wal AO SNS Sree Nee
MOM oli er UO san one eee ey) ee
CHCA Mer Ti. ONO Bip eee 12 at
GN are Pe. <6) a ee
8 BS = 8 2 Tt eS SoS se
oe Ny Neale) = Ge sh geo ee
The correlation coefficient between these
classes has been computed at r=+0.104 +0.080
so that no correlation can be proved: broad
shells generally occur among small specimens
about as frequently as among large ones.
Literature Cited
Allan, Joyce
1956. Cowry shells of the world seas. Melbourne.
x + 170 pp., 15 pits.
Griffiths, R. J.
1956. Cypraea in north-west Malaya. Journ. Conch.,
London, 24 (3): 85-90; pl.
Schilder, Franz Alfred
1947. Die Cypraeaceaim Lichte der Formenkreislehre.
Arch. Molluskenkunde, 76 (4-6): 169-189.
3; 2 textfigs.
1956. Die Bergmannsche Regel bei Porzellanschnecken,
Zool, Anz. Suppl. vol. 20: 410-414.
1958. Eine fast unbekannte Porzellanschnecke der Ha-
waii - Inseln.
Veréff, Uberseemus.
Schilder, F. A.. & M. Schilder
Bremen (A) 3: 32-38.
1934. Thirteen thousand Cypraeidae from South Java.
Proc. Malac. Soc., 21 (3): 199-213.
1939. Prodrome of a monograph on living Cypraeidae.
Proc. Malac. Soc. 23 (4): 181-231.
Schilder, M, & F. A. Schilder
1937. Die Cypraeidae des Bismarck-Archipels.
Anzeiger 119: 177-194.
Zool.
Page 18
REV ELIGER
Vol. 4; No. 1
New Deep Water Mollusks from the Gulf of California
DONALD R. SHASKY
Conchological Club of Southern California, Los Angeles 7, California
(Plate 4, Figures 1 to 10)
The Ariel Expedition in the Gulf of California
during late August and early September 1960
provided collectors of the Conchological Club
of Southern California an opportunity to secure
first hand deep water shells seldom seen in
private collections. bye
Although all of the material is not yet worked
up, it can be safely estimated that 12 to 15 new
species were obtained. Several papers dealing
with new species are now in preparation by
other members of the expedition.
It was discovered during the preparation of
this paper that some of the museum collections
on our west coast already had specimens of
some of the species described below, either un-
named or confused with similar species. Dr.
Leo Hertlein generously provided me with ma-
terial from the California Academy of Sciences
collection which greatly assisted in describing
some of these species.
In addition to Dr. Hertlein, I wish to thank
Dr. A. Myra Keen for her suggestions in the
preparation of this manuscript, Dr. G. Bruce
Campbell, Joe and Helen DuShane, Dr. Homer
King, and Captain Xavier Mendoza for the loan
of specimens, John Q.and Rose Burch for their
encouragement and the privilege of spending
many hours in their fine library, and last but
not least my wife, Ruth. The photography is
by Elwyn Spaulding.
Emarginula velascoensis SHasky, Spec. nov.
(Plate 4, Figures 1 to 3)
Shell small, oval, white; anterior slope
strongly convex; posterior slope concave below
apex; apex small, of one and one-half whorls,
placed about two-thirds of the way down the
shell; radial sculpture of approximately 24 pri-
mary ribs and numerous secondary ribs which
intersect with concentric cords to form square
pits; concentric cords become increasingly
stronger toward the periphery so that they are
of nearly equal strength with the primary axial
ribs at the margin; fissure narrow, about one-
sixth the length of the anterior slope; anal fas-
ciole laminated, with lamellae about equal in
number to the concentric cords; fasciole mark-
ed internally by a tiny ridge; interior of shell
probably glossy in living specimens; measure-
ments of holotype: length 3.9 mm., width 2.8
mm., height 2.1 mm.; measurements of para—
type: length 5.3 mm., width 3.7 mm., height
2.7 mm.
Holotype: Stanford University Paleontological
Type Collection No. 8'619
Paratype: Shasky Collection
Type locality: in 40-80 fathoms, off the south-
west end of Isla Montserrate, Gulf of Cali-
fornia, Lat. 25°39"Ni; Long, lil Osuive.
September 1, 1960. Ariel Expedition
Explanation of Plate 4
Figure 1: Emarginula velascoensis SHAsKy, spec. nov. Dorsal view of holotype; x 4.5.
Figure 3: Sideview of paratype; x 2.5.
Figure 5: Trigonostoma campbelli Suasky, spec. nov. Holotype, x 2.
ventral view; x 4.5.
Holotype, x 2.
funculatum (Hinps). Guaymas, Sonora, Mexico; x 1.5
Holotype, x 2. Figure 9: Paratype, x 2.
slightly enlarged.
Figure 12: Thyca callista Berry. Pt. Diablo, Baja California, Mexico. x 2.
Figure 15: Turitella sanguinea REEVE. Guaymas, Sonora,
Figure 16: Cantharus biliratus (REEVE). Montserrate Island, Baja California, Mexico. x 2.
(DesHayEs). Gulf of Fonseca, El Salvador. x 1.
Mexico. x 2.5.
Figure 2: Same specimen,
Figure 4: Cancellaria strongi SHASKY, spec. nov.
Figure 6: Trigonostoma
Figures 7 and 8: Arielia mitriformis SHASKY, spec. nov.
Figure 10: Clathrodrillia bicarinata SHasky, spec. nov. Holotype,
Figure 11: Diodora pusilla Berry. Montserrate Island, Baja California, Mexico. x 2.5.
Figures 13 and 14: Sinum grayi
Tue VELIGER, Vol. 4, No. 1 [SHasky] Plate 4
Figure 13 Figure 14 Figure 15 Figure 16
Ga
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Vol. 4; No. 1
THE VELIGER
Page 19
Comparison and Discussion: This species is
closest to Emarginula phrixodes Dall, 1927,
from the tropical Western Atlantic. KE. velas-
coensis differs in having a longer, narrower
fissure, fewer radial ribs, and proportionally
fewer lamellae of the anal fasciole. It should
be noted, however, that Dall apparently record-
ed the number of primary axial ribs of a single
Sagittal diameter, while the number of primary
ribs described for E. velascoensis is the total
at the circumference.
I take pleasure in naming the first tropical
Eastern Pacific representative of the genus for
Sefior Miguel Velasco Pechir of the Mexican
Fish Commission. When the permits for the
Ariel Expedition seemed impossible to obtain,
his intervention and help turned a very dark
hour into a most rewarding week.
Cancellaria strong: SHasky, spec. nov.
(Plate 4, Figure 4)
Shell small, olive brown, with a slightly
darker, thin, somewhat scaly periostracum,
which is tufted at the shoulder where the axial
and spiral sculpture intersect; whorls, includ-
ing nucleus, eight; nucleus smooth, blunt, pol-
ished, of one and one-half whorls; subsequent
whorls strongly shouldered and tabulate; axial
ribs stronger than the spiral cords; axial ribs
12 to 13 forming prominent nodes at the inter-
sections with the spiral cords; shoulder nodes
spinose; spiral cords between the shoulder and
the suture much weaker than those on the whorl;
aperture subtrigonal; white in the adult, brown
in immature specimens; outer lip thickened,
with about 14 lirations in fully mature individu-
als; inner lip thinly calloused and somewhat
reflected back over a small umbilicus; colu-
mella bent toward outer lip; columellar plaits
three, the inferior the strongest; canal short;
length of holotype 17.7 mm.; maximum diameter
10.7 mm.
Holotype: California Academy of Sciences Pale-
ontology Type Collection, No. 12'348
Hypotypes: Stanford University Paleontological
Type Collection, Conchological Col-
lection of the San Diego Natural His-
tory Museum, Campbell collection,
DuShane collection, King collection,
Schowalter collection, Mendoza col-
lection, and Shasky collection
Type locality: in 33-55 fathoms, off Point
Arena, Baja California. Crocker-Beebe
Expedition, 1936.
Hypotype stations;
1, in 20-55 fathoms, off Cabo Haro, Guaymas,
Sonora, Mexico, Lat. 27°50'N., Long. 110°
55'W., December 27 and 31, 1959. Collec-
tors — B. Campbell, X. Mendoza, T. Scho-
walter, D. Shasky, and I. Thompson
2. in 20-40 fathoms, off Montserrate Island,
Gulf of California, Lat. 25°35'N.; Long.
111°05' W., September 1, 1960. Ariel Ex-
pedition
3. in 50-90 fathoms, off Partida and Espiritu
Santo Islands, Gulf of California, Lat. 24°
32'N.; Long. 110°26'W., August 30 and 31,
1960. Ariel Expedition
Comparison and discussion: Of the known West
American species this is closest to Cancellaria
(Admete?) californica Dall, 1908, from which it
differs in having fewer axial ribs, stronger
shoulders, a narrower umbilicus, a pronounced
tufting of the periostracum, anda definite
canal.
Since a comparison of Cancellaria strongi
with the figure of C, elata Hinds, 1843, might
lead to some confusion, mention should also be
made of this species. Cancellaria elata was
described from a single dead specimen dredged
in Panama Bay and has not been reported sub-
sequently. Cancellaria strongi would seem to
differ in being chunkier, lacking the fine spiral
sculpture, having fainter columellar plaits and
a straight canal. Hinds failed to mention a pe-
riostracum and it is assumed that if one were
present in living individuals, it was eroded
away in his specimen.
It would appear that Cancellaria californica
and C. strongi belong to a subgenus, not yet
described, that is intermediate between Admete
and Trigonostoma. As Dall pointed out in the
discussion of C. californica, "It has the aspect
of an Admete, in spite of the presence of an
umbilicus, but is perhaps only a delicate form
of Trigonostoma. "'
That these two species do not belong in the
genus Admete seems certain. Their relation-
ship to Trigonostoma remains to be evaluated.
This species is named in honor of the late A.
M. Strong, who recognized it as a new species
but failed to provide a name and complete de-
scription. The holotype was mentioned in his
notes on Cancellariidae, published after his
death, as Cancellaria (Cancellaria) sp.
Page 20
THE VELIGER
Vol. 4; No. 1
Trigonostoma campbelli Suasxy, spec. nov.
(Plate 4, Figure 5)
Shell small, acutely turreted, dark brown;
whorls six, including one and one-half smooth,
glossy nuclear whorls; whorls pinched at base
so that succeeding whorls override the previ-
ous whorl and immerse the suture; axial ribs
six, rounded at the shoulder, crossed by fine
primary and secondary spiral threads; aper-
ture trigonal, brown within; outer lip finely li-
rate; columella two-plaited with the superior
plait stronger; columella angled toward outer
lip; inner lip calloused and reflected back to
form a deep umbilicus; canal short, open;
length of holotype 16.0 mm.; maximum diame-
ter 9.3 mm.
Stanford University Paleontological
Type Collection No. 8'620.
Holotype:
Campbell collection, Mendoza col-
lection, and Shasky collection
Paratypes:
Conchological Collection of the San
Diego Natural History Museum and
Shasky: collection
Hypotypes:
Type locality: in 30-50 fathoms, off Cabo Haro,
Guaymas, Sonora, Mexico, Lat. 27°50'N.;
Long. 110°55'W., December 14, 1958, and
December 27 and 31, 1959. Collectors —
B. Campbell, X. Mendoza, T. Schowalter,
and D. Shasky
Hypotype stations:
1. in 10 fathoms, Puerto Pefiasco, Sonora,
Mexico. February 1934. Collector — H.
N. Lowe
2. in 20 fathoms, off Punta Final, San Luis
Gonzaga Bay, Baja California, Mexico,
Lat. 29°47'N., Long. 114°18'W, January
1, 1961. Collectors — B. Campbell, G.
Sphon, and D. Shasky
Comparison and discussion: Trigonostoma
campbelli is close to T. funiculatum (Hinds,
1843) (see Plate 4, figure 6) but differs from it
in its dark brown color, being more acutely
turreted, the sutures more deeply impressed
and immersed, the aperture more trigonal, and
the two-plaited columella bent more toward the
outer lip.
Despite these differences the two are easily
confused, and it was not until I collected both
species from off Cabo Haro, Guaymas, Sonora,
that their differences were recognized.
Lowe (1935) reported collecting Trigonosto-
ma funiculatum at Puerto Pefiasco and Guay-
mas, Sonora, and Manzanillo, Colima. Exam-
ination of his material, however, reveals that
the specimen from Puerto Pefiasco is T. camp-
belli.
Ilalso reported Trigonostoma funiculatum
(Shasky, 1960), but the material reported is the
type material for T. campbelli.
This fine species is named in honor of my
close friend and colleague, Dr. G. Bruce
Campbell, whose enthusiasm for diving and
dredging for shells has and will continue to be
a stimulant to all West Coast conchologists.
Artelia Suasky, gen. nov.
Shell small, fusiform, reticulately sculp-
tured; nucleus smooth, planorboid, of one and
one-half whorls; aperture long, narrow, about
one-half length of shell; anal notch abrupt,
shallow, with no subsutural callous pad; colu-
mella two-plaited; outer lip lirate within; type
species Arielia mitriformis.
As much as one dislikes adding another ge-
nus to the already overcrowded turrid nomen-
clature, the type species did not seem to fit into
any of the recognized genera.
Arielia mitriformis SHasKy, spec. nov.
(Plate 4, Figures 7 to g)
Shell small, fusiform, with one and one-half
smooth, white, planorboid nuclear whorls fol-
lowed by seven sharply reticulate whorls; su-
ture inconspicuous with a strong subsutural bi-
carinate band; axial sculpture of fine ribs which
form small nodes with the spiral cords, leav-
ing shallow rectangular depressions; axial ribs
18 to 19 on the body whorl; spiral sculpture, in
addition to the subsutural band, of three cords
which arise wea kly but become increasingly
stronger until they reach the antepenultimate
whorl where they are of equal strength with the
axial ribs; spiral cords increase to 18 or 19 on
the body whorl; aperture long, narrow, about
one-half the length of the shell; anal notch open,
shallow, and lacking a subsutural callous pad;
outer lip thin, lirate within; columella two-
plaited, the superior plait much stronger; inner
lip not calloused; canal short, open; color pre-
sumably tawny white banded with brown; length
of holotype 12.5 mm., width 4.5 mm.
Holotype: Stanford University Paleontological
Type Collection No. 8'621
Paratypes: California Academy of Sciences
Paleontology Type Collection,
Campbell collection, DuShane col-
Vol. 4; No. 1
THE VELIGER
Page 21
lection, Shasky collection, and Sphon
collection
Type locality: in 40-90 fathoms, off Islas Par-
tida and Espiritu Santo, Gulf of California,
Lat. 24°32'N.; Long. 110°26'W. August 30
and 31,1960. Ariel Expedition
Comparison and discussion: Arielia mitrifor-
mis has the general outline of a Daphnella, but
the columellar plaits preclude its inclusion with
that group.
It has been suggested that it might fall into
the genus Zetekia, but Dall proposed this for a
much smaller shell with a short and propor-
tionally more open aperture, and four or five
lirations on the columella.
Because of the columellar plaits, Arielia mi-
triformis also has the appearance of a small
Mitra, and during the hasty initial sorting
aboard ship it was included with specimens of
Mitridae.
Clathrodrillia (Carinodrillia) bicarinata
SHASKY, spec. nov.
(Plate 4, Figure 10)
Shell medium size, turreted, with 15 whorls,
three of which form a smooth convex nucleus;
color mocca brown with dark reddish brown
spiral bands between the spiral cords; spiral
cords two, increasing to three or four, on later
whorls; the two primary spiral cords form a
double undulating keel as they intersect with the
axial ribs; later whorls with one or two lesser
spiral cords except the body whorl which has
12 to 15 spiral cords separated by fine spiral
threads; axial sculpture of seven to nine sharp
ribs; aperture long, narrow, bluish white with-
anal sulcus rounded, with a moderately
heavy subsutural callus; outer lip thin, crenu-
late, and gently curved, smooth within; pillar
relatively straight; canal open; length of holo-
type 48 mm.; length of aperture 19 mm.; maxi-
mum diameter 15 mm.
in;
Stanford University Paleontological
Type Collection No. 8'622
Holotype:
Paratypes: Burch collection, Campbell collec-
tion, DuShane collection, King col-
lection, Rogers collection, Scho-
walter collection, Shasky collec-
tion, and Sphon collection
Type locality: in 45-90 fathoms, off Islas Par-
tida and Espfritu Santo, Gulf of California,
Lat. 24°32'N.; Long. 110°26'W. August 30
and 31, 1960. Ariel Expedition
Hypotype stations:
1. in 20-120 fathoms, off Cabo Haro, Guay-
mas, Sonora, Mexico, Lat. 27°50'N.;
Long. 110°55'W. September 2, 1960. Ariel
Expedition
2. in 12-25 fathoms, off Loreto, Baja Califor-
nia, Lat. 26°01'N.; Long. 111°18'W. Au-
gust 29,1960. Ariel Expedition
Comparison and discussion: This is the larg-
est of the Carinodrillias so far described. That
it should escape description until this late date
is difficult to understand since it seems to be
distributed widely in the southern half of the
Gulf of California.
Of the previously described species, from
the Panamic region, it is closest to Clathro-
drillia (Carinodrillia) alcestis (Dall, 1919), from
which it differs in its larger size, the double
keeled sculpture, the curved outer lip, and the
spiral color banding.
Literature Cited
Dall, 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 inthe Gulf of California.
XIV. The Mollusca and Brachiopoda. Bull. Mus.
Comp. Zool., Harvard, 43(6): 205-487, pls. 1-22.
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:
1-86, pls. 1-24.
1927. Diagnosis of undescribednew species of mollusks
in the collection of the United States National Mu-
seum. Proc. U. S. Nat Mus., 70(no. 2668): 1-11.
Hinds, Richard Brinsley
1844. The zoology of the voyage of H. M. S. Sulphur,
under the command of Capt. Sir Edward Belcher.
London. Mollusca, vol. 2: 1-72, pls. 1-21.
Keen, A, Myra
1958. SeashellsoftropicalWestAmerica. xi, 624 pp.,
illus. Stanford Univ. Press, Stanford, Calif.
Lowe, H.N.
1935. Newmarine mollusca fromwest Mexico, together
with a list of shells collected at Punta Penasco,
Sonora, Mexico. Trans. San Diego Soc. Nat. Hist.
8 (6): 15-34, pls. 1-4.
Perez Farfante, I.
1947. The genera Zeidora, Nesta, Emarginula, Rimu-
la, and Puncturella in the western Atlantic. John-
‘sonia, 2 (24): 93-148, pls. 41-64.
Pilsbry, H. A., & H. N. Lowe
1932. West Mexican and central American mollusks
collected by H. N. Lowe, 1929-1931. Proc. Acad.
Nat. Sci. Phila., 84: 33-144, pls. 1-17, 7 text figs.
Shasky, Donald R.
1960. Deep water collecting off Guaymas,
The Veliger, 3 (1): 22-23.
Strong, A. M.
1954. A review of the eastern Pacific species in the
Mexico.
molluscan family Cancellariidae. Minutes Conch.
Club Southern Calif., no. 135: 7 tono. 136: 23.
Page 22
THE VELIGER
Vol. 4; No. 1
Notes on Rare and Little Known Panamic Mollusks
DONALD R. SHASKY
Conchological Club of Southern California, Los Angeles 7, California
(Plate 4, Figures 11 to 16)
During the last five years I have engaged in
extensive collecting in the Gulf of California,
especially along the coast of northern Baja Ca-
lifornia. Almost every trip has brought to light
one Or more species previously reported only
from the type locality, frequently 1,000 — 3,000
miles south of the stations to be discussed here.
These trips are usually in company with other
collectors who will be mentioned in the text as
various species and records are considered.
I have also had the opportunity to examine
several lots of shells from the Gulf of Fonseca,
El Salvador, and from the Chiapas and Oaxaca
coasts of Mexico. These were sent to me
through the courtesy of Captain Xavier Mendoza
currently of Salina Cruz, Mexico. In the Salva-
dor material were several examples of a spe-
cies previously considered to range from Peru
to Chile.
Mention is also made of a common California
bivalve recently discovered in the Gulf of Cali-
fornia.
Lioberus salvadoricus (Hertlein & Strong, 1946)
This small mytilid was originally described
from Costa Rica but has more recently been
reported from along the Sonora coast of Mexico.
Station: in 6-8 fathoms, five miles north of
Punta San Felipe, Baja California. Lat. 31°
7'N.; Long. 114°46'W. March 1959. B.
Campbell and D. Shasky.
The bottom in which we took this species is
a thick mud that quickly clogs the dredge. It
seems to be one of the few species living in this
area, although valves of Aequipecten palmeri
(Dall, 1897) were quite common.
Tivela stultorum Mawe, 1823
It is well known that in the northern Gulf of
California there exists a cold water fauna that
compares in many ways with that of the Pacific
Coast of California and northern Baja Califor-
nia. To list here species that are common to
both areas is not within the scope of this paper.
It is mentioned merely to add another species
common to both regions. Tivela stultorum is
reported by Fitch to range only as far south as
Magdalena Bay, but Oldroyd lists Socorro Is-
land as its southern limit.
Station: intertidally at southwest end of Wil-
lard Island, San Luis Gonzaga Bay, Baja
California. Lat. 29°49'N.; Long. 114°24' W.
January 1, 1961. G. Sphon and D. Shasky.
Approximately 20 specimens were uncovered
in a very small area. These ranged in size
from 43 to 6 inches. Although it has been sug-
gested that these might have been introduced
here, it is difficult to reason why this rather
isolated location in an already semi-isolated
area would be chosen.
Diodora pusilla Berry, 1959 (Plate 4, fig. 11)
This small off-shore form has only recently
been described from off Acapulco, Mexico. It
is figured here for the first time.
Stations:
1. in 40-80 fathoms, off the southwest end of
Isla Montserrate, Gulf of California. Lat.
25°39'N.; Long. 111°05'W. September 1,
1960. Ariel Expedition, D. Shasky.
2. in 20 fathoms, off Punta Final, San Luis
Gonzaga Bay, Baja California. Lat. 29°47'
N.; Long. 114°18'W. January 1961. B.
Campbell, G. Sphon, and D. Shasky.
Thyca callista Berry, 1959 (Plate 4, fig. 12)
This very interesting form, parasitic on the
starfish Phataria unifascialis Gray, was first
taken by Leonard Bessom in 1954 while diving
at San Carlos Bay, Guaymas, Sonora, Mexico.
I collected a single specimen in an adjacent
area in 1958.
Stations:
1. Mazatlan, Sinaloa, Mexico. December
1959. James McLean.
2. in 23 fathoms, off Punta Diablo, Baja Cali-
fornia. Lat. 24°18'N.; Long. 110°19'W.
August 31,1960. D. Shasky.
Vol. 4; No. 1
THE VELIGER
Page 23
LEW
All specimens of this species taken to date
have been off-shore in depths of 6-20 feet, al-
though the starfish host is frequently seen at
extreme low tides.
Mr. Bessom and I estimate that Thyca cal-
lista occurs once on every 1,000-1,500 starfish
examined.
Sinum grayi (Deshayes, 1843) (Plate 4, figs. 13, 14)
This magnificent deep water species has been
reported only once north of Peru, and then it
was supposed to have been taken living at San
Pedro, California. I have grave doubts con-
cerning the accuracy of the San Pedro record.
Stations:
‘1. in 20-40 fathoms, Gulf of Fonseca, El Sal-
wadorwe fat. 13015"N.;) Long. 87° 45'W.
October 1960. X. Mendoza.
2. specimens brought in by the Guaymas
shrimp fleet have recently been acquired
by C. Zimmerman, R. Burch, and Dr. M.
Keen. It is assumed these were all taken
within the territorial waters of Mexico.
Lamellaria inflata (C. B. Adams, 1852)
Range: Panama
Stations:
Ihe aire 1-13 fathoms, in siftings, Puertecitos,
Baja California. Lat. 30°25'N.; Long. 114°
39'W. June 1960. D. Shasky.
Z. in 10 fathoms, La Paz Bay, Baja Califor-
nia. Lat. 24°12'N.; Long. 110°22'W. Ariel
Expedition. G. Sphon.
Mr. Sphon's specimen was living when col-
lected. He reports that the animal is light yel-
low marked with brown.
Turritella sanguinea Reeve, 1849 (Plate 4, fig. 15)
This species does not seem to have been rec-
ognized since Reeve's original description.
The type locality was cited as California.
Station: in 20-40 fathoms, off Cabo Haro,
Guaymas, Sonora, Mexico. Lat. 27°50'N,;
Long. 110°55'W. December 28 and 31,1959.
B. Campbell and D. Shasky.
Our specimens match Reeve's description
and figure except that he described the color as
blood red while our six shells tend to be brown-
ish red.
We had this mixed in the same lot with Tur-
ritella leucostoma Valenciennes, 1832, until we
noted the more inflated early whorls. With this
in mind, separation was relatively simple.
Typhis lowei Pilsbry, 1931
Previously reported only as far north as the
coast of Oaxaca, Mexico.
Stations:
1. crab specimen, under a rock, about five
feet below the surface. Puerto Ballandra,
Isla Carmen, Guif of California. Lat. 26°
O1'N.; Long. 111°11'W. August 29, 1960.
D. Shasky.
2. in20-40 fathoms, off Loreto, Baja Califor-
nia. Lat. 26°01'N.; Long. 111°18'W. Au-
gust 29, 1960. Ariel Expedition. G. Sphon.
Cantharus bilirata(Reeve, 1846) (Plate 4, fig. 16)
Previous range: Galapagos and Viti (Fiji) Is-
lands.
Stations:
1. in 40-80 fathoms, off Isla Montserrate,
Gulf of California. Lat. 25°39'N.; Long.
111°05'W. September 1,1960. Ariel Expe-
dition. B. Campbell.
2. in 20 fathoms, off Punta Final, San Luis
Gonzaga Bay, Baja California. Lat. 29°47'
N.; Long. 114°18'W. January 1, 1961. B.
Campbell, D. Shasky, and G. Sphon. (Spe-
cimens taken here were fragments only.)
Nassarius howardae Chace, 1958
Recently described from San Felipe, Baja
California.
Station: in 10-15 fathoms, off Cabo Haro,
Guaymas, Sonora, Mexico. Lat. 27°53'N.;
Long. 110°50'W. December 14, 1958. D.
Shasky.
Although not mentioned in the description,
this species is closely related to Nassarius ca-
tallus (Dall, 1908). Most apparent differences
in these two species are the more acute apex,
solid buff-white color and flaring outer lip in
N. howardae.
Cancellaria obesa Sowerby, 1832
Although this is not an uncommon off-shore
form, it is included here to record it as having
been collected alive intertidally.
Station: intertidally, bumping in sand at
night, one cove south of Puertecitos, Baja
California. Lat. 30°25'N.; Long. 114°39'W.
November 1959 (G. Sphon); January 1961 (D.
Shasky); and February 1961 (W. Barber).
Page 24
THE VELIGER
Vol. 4; No. 1
Daphnella allemani (Bartsch, 1918)
Previous record: Taboga Island, Panama.
Stations:
1. in20-40 fathoms, off Loreto, Baja Califor-
nia. Lat. 26°01'N.; Long. 111°18'W. Au-
gust 29,1960. Ariel Expedition. D.Shasky.
2. in 20 fathoms, off Punta Final, San Luis
Gonzaga Bay, Baja California. Lat. 29°47'
N.; Long. 114°18'W. January 1, 1961. B.
Campbell, G. Sphon, and D. Shasky.
Daphnella mazatlanica Pilsbry and Lowe, 1932
This species is taken occasionally at Guay-
mas, but this seems to be the first record in
the northern Gulf of California.
Station: intertidally, under a rock, Willard
Island, San Luis Gonzaga Bay, Baja Cali-
fornia. Lat. 29°49'N.; Long.'114°24' Ww.
January 1,1961. Ruth Shasky.
Clavus melea (Dall, 1919)
This seems to have been recorded only from
Panama.
Stations:
1. in20-40 fathoms, off Loreto, Baja Califor-
nia. Lat.26°01'N.; Long. 111°18'W. Au-
gust 29, 1960. Ariel Expedition. D. Shasky.
2. in 20 fathoms, off Punta Final, San Luis
Gonzaga Bay, Baja California. Lat. 29°47'
N.; Long. 114°18'W. January 1, 1961. B.
Campbell, D. Shasky, and G. Sphon.
Mangelia finitima (Pilsbry and Lowe, 1932)
Unreported since taken by Lowe in Nicaragua.
Stations:
1. crab specimen, under a rock, 6-10 feet
below the surface, Saladita Bay, Guaymas,
Sonora, Mexico. June 1958. D. Shasky.
2. crab specimen, under a rock, intertidally,
Puertecitos, Baja California. April 25,
1959. D. Shasky.
Tenaturris burchi (Hertlein and Strong, 1951)
Type locality: 45 fathoms off Arena Bank,
Gulf of California.
Stations:
1, in20-40 fathoms, off Cabo Haro, Guaymas,
Sonora, Mexico. Lat. 27°50'N.; Long. 110°
55'W. December 27 and 31,1959. D. Shasky.
2. crab specimen, under a rock, 4-5 feet be-
low surface, Puerto Ballandra, Isla Car-
men, Gulf of California. Lat. 26°01'N.;
Long. 111°11'W. August 29,1960. D. Shas-
ky.
Tenaturris carissima (Pilsbry and Lowe, 1932)
Originally described from Manzanillo, Coli-
ma, Mexico, it has recently been dredged off
Isla Carmen, Gulf of California.
Station: crab specimens under rocks, inter-
tidally, San Luis Gonzaga Bay, Baja Cali-
fornia. December 30 and 31, 1960. B.
Campbell and D. Shasky.
Tenaturris nereis (Pilsbry and Lowe, 1932)
Type locality: San Juan del Sur, Nicaragua.
This spécies is quite common in the Gulf of
California as crab specimens, intertidally, un-
der rocks. I have taken this species at the fol-
lowing areas: Puertecitos and San Luis Gonza-
ga Bay, Baja California, and Puerto Pefiasco
and Guaymas, Sonora, Mexico. It has been ta-
ken by H. DuShane at San Felipe, Baja Califor-
nia.
Literature Cited
Berry, S.S.
1959. Notices on new eastern Pacific Mollusca - III.
Leaflets in Malacol. 1 (18).
Chace, E. P.
1958. A newmollusk from San Felipe, Baja California.
Trans. San Diego Soc. Nat. Hist., 12(20): 333-334,
one fig.
Dall, William 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:
1-86, pls. 1-24.
Emerson, William K., & Elton L. Puffer
1957. Recent mollusks of the 1940 "E. W. Scripps"
cruise to the Gulf of California. Am. Mus. Novit.
no. 1825: 1-57, 2 figs., 1 table.
Fitch, John E.
1953. Common marine bivalves of California. Calif.
Dept. Fish & Game, Fish Bull. No. 90; 106 pp.,
63 figs.
Keen, A. Myra
1958. Seashells oftropicalWestAmerica. xi, 624 pp.,
illus. Stanford Univ. Press, Stanford, Calif.
Oldroyd, Ida Shepard
1924. The marine shells of the west coast of North
America. Stanford Univ. Publ. Geol. Sci., vol. 1,
247 pp, 57 pls.
Pilsbry, H. A, & H. N. Lowe
1932. West Mexican and central American mollusks
collected by H. N. Lowe, 1929-31. Proc. Acad.
Nat. Sci. Phila. 84; 33-144, 6 figs., pls. 1-17.
Reeve, L.
1846.
1849. Ibid.
Tryon, G. W., Jr.
1886. Manual of conchology, vol. 8.
Turner, R. D.
1956. The eastern Pacific marine mollusks described
by C.B. Adams. Occ. Pap. Mollusks, Mus, Comp,
Zool., Harvard, 2 (20): 21-135.
Conchologia Iconica. Buccinum. London.
Turritella. London.
Philadelphia.
Vol. 4; No. 1
THE VELIGER
Page 25
Four New Panamic Gastropods
by
G. BRUCE CAMPBELL
Conchological Club of Southern California, Los Angeles 7, California
(Plate 5)
As areas of the Gulf of California and Pana-
mic region become more accessible and as
dredging operations become more frequent,
mollusks new to science are certain to appear.
There will be extensions of ranges, and doubtful
species of early authors may be rediscovered.
The four new species described in this paper
were collected in this region during the past
three years, obtained by shore collecting at low
tide, from the trawling of the Ariel Expedition,
from the nets of the shrimp fleets, and by
dredging from my small outboard boat.
Genus Trivia Broverip, 1837
Subgenus Pusula Jousszaume, 1884
Trivia (Pusula) myrae Camppe.i, spec. nov.
(Plate 5, Figures 1 to 3)
DESCRIPTION:
The shell is minute, ovately globular, and
dark brownish purple with the right side and
extremities thickened and margined but not
quite as rounded as the columellar portion. The
shell is moderately produced at the extremi-
ties, anda fine, shallow, dorsal sulcus is
crossed by eight ribs without interruption.
There are 22 ribs on each side, nine of which
are interrupted or intercalary and correspond
to 15 sharp labial teeth, with 14 similar teeth
extending over the columella and internal lobe.
As the ribs enter the dorsal sulcus, the color
is lighter, giving the impression of very slight
beading. The ribs are narrow and sharp, equal-
ing about one-half the width of the interspaces,
which are filled with minute granulations that
extend up on the sides but not to the crests of
the ribs. The spire is completely obscured.
Holotype: length 4.8 mm.; width 3.6 mm.; height
2.8mm. Paratype I: length 4.5 mm.; width
3.4mm.; height 2.9 mm. Paratype II: length
3.9 mm.; width 3 mm.; height 2.7 mm. Holo-
type: Stanford University Paleontological Type
Collection No. 8'529; Paratype I: private col-
lection of Dr. Donald Shasky. Paratype II: in
my private collection.
TYPE LOCALITY:
The holotype and the two paratypes were
trawled off Loreto in the channel between Lo-
reto, Baja California, and Carmen Island, Gulf
of California, on the Ariel Expedition at an ap-
proximate depth of 25 fathoms on August 29,
1960. Lat. 26°01'N.; Long. 111°18'W.
HYPOTYPES:
Seven additional specimens were trawled off
Monserrate Island, Gulf of California, on the
Ariel Expedition at a depth of 40-80 fathoms on
September 1,1960. Two specimens were
dredged off Punta Final, 10 miles south of San
Luis Gonzaga Bay, Baja California, in approxi-
mately 30 fathoms by Dr. Donald Shasky, Mr.
Gale Sphon, and myself. :
Discussion
So far this species appears to be limited to
the eastern shore of Baja California at moder-
ate depths, 25-80 fathoms. Extensive dredging
at Puerto Pefiasco, Guaymas, Mazatldn, Salina
Cruz and El Salvador, all on the mainland,
has failed to produce this small Trivia. Con-
siderable time has been spent reviewing the 23
or more species of Trivia that have been re-
ported from the Eastern Pacific, and with the
help of Dr. Keen, it was determined that there
is no valid name to which this small species
can be assigned. Of the recognized species of
Panamic Trivia, T. myrae resembles T. ato-
maria Dall, 1902, which was dredged in Pana-
ma Bay at a depth of 18 fathoms. Trivia ato-
maria belongs to the subgenus Cleotrivia, which
differs from Pusula in that the rib ends are not
beaded on either side of the dorsal sulcus.
Trivia myrae is further separated from T.
atomaria by having produced extremities,
slightly beaded ribs, and more numerous ribs.
In comparison with T. sanguinea (Sowerby,
1832), T. myrae is much smaller, has fewer
ribs, is much more globular, and has more
produced extremities.
This new species of Trivia is named in honor
of Dr. Myra Keen for her participation in the
Ariel Expedition during which time this spe-
cies was discovered, and for the many hours
that she has devoted in helping me with prob-
lems regarding mollusks.
Page 26
THE VELIGER
Vol. 4; No. 1
Genus Nassarina: Dat, 1889
Subgenus Zanassarina Pirssry « Lowe, 1932
Nassarina (Zanassarina) anitae
CAMPBELL, Spec. nov.
(Plate 5, Figure 4)
DESCRIPTION:
The shell is fusiform, basically brown with a
peripheral orange-brown band that colors the
interspaces as well as alternately coloring the
fourth node on one axial rib and the third and
fifth nodes on the next; the remaining nodes are
white. The protoconch consists of three coni-
cal, convex whorls, followed by six subsequent
whorls with two weak, subsutural spiral cords
and three remaining strong spiral: cords that
intersect the 12 straight axial ribs which are
narrower than the interspaces, forming more
prominent nodes. The ribs of the convex whorls
are obtusely angled by the third spiral cord.
There are 13 additional spiral cords on the
base of the body whorl, the last 12 being gen-
erally brown, which continue under the colu-
mellar callus corresponding to 10 weak plica-
tions on the columella. The siphonal canal is
not differentiated, slightly recurved and deep.
The anal sinus is shallow and the siphonal fas-
ciole is not discernible. There are five denti-
cles within the outer lip, the first more promi-
nent than the remaining four. Length 10 mm.,;
width 4.5 mm.
Holotype: Stanford University Paleontologi-
cal Type Collection No. 8'530.
Paratypes: To be deposited in the U. S. Na-
tional Museum, California Academy af Sciences,
Academy of Natural Sciences of Philadelphia,
and the private collections of Dr. Donald Shas-
ky, Mr. Mark Rogers, Mr. Gale Sphon, Mr.
Todd Schowalter, and myself.
TYPE LOCALITY:
The holotype and seven paratypes were
trawled off Cabo Haro, Guaymas, Mexico, by
the shrimp boat, ''General Yafiez"', in 30 fathoms
by Captain Xavier Mendoza, Dr. Donald Shasky,
Mr. Todd Schowalter, Mr. Ivan Thompson, and
myself in December of 1959. Lat. 27°50'N.;
Long. 110°55'W.
Sixteen additional paratypes were trawled off
Cabo Haro, Guaymas, Mexico, during the Ari-
el Expedition in 10-25 fathoms on August 28,
1960. Eight hypotypes were collected by Dr.
Shasky under rocks at low tide on the north
shore of Bacochibampo Bay, Guaymas, Mexico,
in December of 1958.
Discussion
This species seems related to Nassarina xeno
Pilsbry and Lowe, 1932, and N. poecila Pilsbry
and Lowe, 1932. The somewhat flattened sub-
sutural region, slightly obtuse-angled ribs, usu-
ally 12 per whorl, five spiral cords, and absent
siphonal fasciole distinguish it from N. xeno,
which has 10 to 12 convex ribs, small scattered
black spots, broad, rounded anal sinus, six
spiral cords and weakly developed siphonal
fasciole, and from N. poecila, which has black-
ish brown blotches above the periphery on part
of the ribs, seven spiral cords of which the
subsutural cord is larger, and only 10 ribs.
It differs from Nassarina atella Pilsbry and
Lowe, 1932, which has seven spiral cords, the
five peripheral ones being stronger, and anal
sinus separated by a callus from the suture.
Nassarina pammicra Pilsbry and Lowe, 1932,
is a slender black shell with more numerous
small diminishing ribs which, on the last whorl,
scarcely reach the suture.
This new species is named in honor of my
wife, Anita Campbell, who participated in the
trip aboard the ''General Yafiez"'.
Genus Terebra Brucurére, 1789
Subgenus Strivterebrum Sacco, 1891
Terebra (Strioterebrum) berry
CAMPBELL, Spec. nov.
(Plate 5, Figures 5, 6)
DESCRIPTION:
The shell is medium sized, slender, of light
cream with brown spots between the subsutural
nodes and brownish mottling of the whorls with
exception of the protoconch and very early
whorls, which are practically black. The four
dark, glassy whorls of the protoconch are fol-
lowed by 15 subsequent whorls. The early
whorls have very narrow, slightly curved axial
ribs, 14 in number, with scarcely any indica-
tion of a subsutural band. On later whorls the
axial ribs are prominent, quite curved, and
about equal to the interspaces and number about
16. There are three to four spiral grooves in
the interspaces, usually not crossing the ribs,
totaling 18 to 20 down over the base of the body
whorl. The general surface is microscopically
striolate, and the slightly convex axial ribs are
mildly depressed or on an equal level with the
subsutural band to form straight sides. The
aperture is elongate and passes below into a
short, open, slightly recurved canal with the
Vol. 4; No. 1
THE VELIGER
Page 27
siphonal fasciole convex, bounded by a weak
cord, and the columella straight with one weak
plication. Length 32 mm.; width 7 mm.
Holotype: California Academy of Sciences,
Department of Geology, Type Collection No.
12'352.
Paratype I: Private collection of Mrs. Helen
DuShane. Length 23 mm.; width 5.5mm. This
specimen displays a common variation in Tere-
bridae by having 21 axial ribs, but in all other
respects it agrees with the holotype.
Paratype II: Private collection of Mr. Mark
Rogers. Length 27 mm.; width 6 mm.
Paratype III: Private collection of Dr. S.
Stillman Berry.
eS O GAT LTR:
The holotype was collected at Puertecitos,
Baja California, crawling in the sand at a very
low tide by the DuShanes in April, 1958. Para-
type I, collected by myself on March 3, 1958,
Paratype II, collected by Mr. Rogers on April
11, 1960, and Paratype III in the Berry collec-
tion were taken at the same locality. Lat. 30°
25'N.; Long. 114°39'W.
Discussion
There isno little confusion in the genus Ter-
ebra with regards to the Panamic area. I have
collected more than 80 names from the litera-
ture for the 37 species as listed in ''Sea Shells
of Tropical West America''". Fortunately, a
good share of these names can safely be placed
in synonymy. It is this background that causes
some hesitance in describing this and the fol-
lowing species. The Terebra collections of the
San Diego Museum of Natural History, Los An-
geles County Museum, Stanford University, and
the California Academy of Sciences were stud-
ied, and through the generosity of the réspec—
tive curators, I was able to make color slides
of each species at Stanford University and at
the California Academy of Sciences. These
have proved invaluable.
The first specimen of Terebra berryi was
collected by the author along with numerous T.
variegata Gray, 1834, at Puertecitos, Baja Ca-
lifornia. It was easily separated from T. vari-
egata, but was regarded as a ''freak''. In Oc-
tober of 1960 the Terebra was shown to Dr.
Berry who promptly produced a single speci-
men from his unnamed material that had been
collected at Puertecitos. Friends who had col-
lected in this area were then asked if they knew
of any similar shells, and Mrs. DuShane had
the one chosen as the holotype, and Mr. Rogers
had another, both collected at Puertecitos.
Terebra berryi can be compared to the most
common species found at Puertecitos, T. vari-
egata which has a prominent subsutural band
set out like a collar, a well developed siphonal
fasciole that is concave, and axial ribs that
are flat and almost straight. Terebra armilla-
ta Hinds, 1844, is also found there, but its sub-
sutural band is even more prominent and the
shell is deep brown. Another species found at
Puertecitos is T. glauca Hinds, 1844, It is dark
gray-brown with its axial ribs distinctly beaded.
Terebra berryi differs from these in that the
subsutural band is slightly depressed, the axial
ribs are convex and curved, and the siphonal
fasciole is less developed and convex. Terebra
berryi can be separated most easily by exami-
nation of the protoconch and early whorls which
differ uniquely from the other species as is
seen in figures 6,9, 10. After comparison with
descriptions, pictures, and specimens of other
Panamic species, it was felt that T. berryi
should be described as new.
This new species is named in honor of Dr.S,
Stillman Berry for his great contributions to
the knowledge of malacology and for his gener-
osity of time as well as material.
Terebra (Strioterebrum )) ninfae
CAMPBELL, Spec. nov.
(Plate 5, Figures 7, 8)
DESCRIPTION:
The shell is minute, medium to dark red-
brown with a tan peripheral band. The three
and one-half glassy whorls of the protoconch
are followed by six whorls of the teleoconch.
The first two whorls following the protoconch
are transparent white with brownish subsutural
bands. There are 12 convex, narrow, arched,
axial ribs with sharp, round, beaded nodes on
the subsutural band. On the subsequent whorls
the axial ribs develop into a row of whitish pe-
ripheral nodes axially lengthened, narrower
than the interspaces, and there is a continuous
series of fine incised spiral grooves, two in the
interspaces of the band and eight on the rest of
the whorl between subsutural bands, intersect-
ing the ribs, continuing down over the body
whorl to total 16. The general surface is cov-
ered with microscopic striulae with the aper-
ture elongate, canal open and recurved, and the
columella is bent to the left with a single plica-
tion weakly entering the aperture. Length6.2
mm.; width 2 mm.
Page 28
GEM EGER
Vol. 4; No. 1
Holotype: California Academy of Sciences,
Department of Geology, Type Collection No,
23.53%
Paratypes: To be deposited in the Academy
of Natural Sciences of Philadelphia, and in the
private collections of Captain Mendoza, Dr.
Shasky, and myself.
TYPE LOCALITY:
The holotype and three paratypes were
trawled by the shrimp boat, ''Cameronera No.
20", in 15-20 fathoms while working in the area
between Puerto Madero and 30 miles north of
the Guatemala border, Chiapas, Mexico, in
January, 1961. Seven additional paratypes were
trawled from this area by the same boat in 14-
20 fathoms during March, 1961. Specimens
supplied by Captain Mendoza. Lat. 14°55'N.;
Long. 92°15'W. i
Discussion
Due to the small size of the first four shells
that were received, I thought that they were the
young of one of the larger nodulosed Terebra.
Using microscopy, the protoconchs and early
whorls were compared with T. tuberculosa
Hinds, 1844 (Pl. 5, fig. 14), and T. cracilenta Li,
1930 (Pl. 5, fig. 12), but T. ninfae was found to
be very different as seen in figures 8, 11-14.
Several weeks later seven more specimens of
T.ninfae were received from Captain Mendoza,
and they too were uniformly of the same small
size.
Terebra ninfae resembles T. roperi Pilsbry
and Lowe, 1932 (Pl. 5, fig. 12), except that T.
roperi is much larger, light brown with a dark
base and protoconch, has traces of two im-
pressed spiral lines in the concavity of the
whorls, and a concave siphonal fasciole, while
T.ninfae is dark red-brown with a clear glas-
sy protoconch and early whorls, and a convex
siphonal fasciole. Terebra ninfae differs from
T. bridgesi Dall, 1908 (Pl. 5, fig. 11), by having
a peripheral row of nodes that are in line with
the nodes on the subsutural band while T.
bridgesi is purple with a broad, white periph-
eral band and has the sutural band distinctly set
off by a strongly constricted sulcus with the
ribs on the band alternating with the ribs on the
whorl.
This new species is named in honor of Sra.
Ninfa Mendoza, wife of Captain Xavier Men-
doza.
Acknowledgment
Appreciation is expressed to Mr. and Mrs.
John Q. Burch for the kind use of their library;
to Drs. Hanna and Hertlein and the California
Academy of Sciences, Dr. S. Stillman Berry,
Dr. Donald Shasky, Mr. Mark Rogers, and Cap-
tain Xavier Mendoza for making study material
available; to Dr. Myra Keen who has helpfully
provided suggestions regarding historical and
taxonomic problems, and to the several other
persons and institutions mentioned in the text.
Literature Cited
Dall, William Healey
1902. New species of Pacific coast shells.
16 (4): 43-44.
Nautilus
1908. Report on the dredging operations off the west
coast of central America to the Galapagos, to the
west coast of Mexico, andin the Gulf of California.
XIV. The Mollusca and Brachiopoda. Bull. Mus.
Comp. Zool., Harvard, 43 (6): 205-487, pls. 1-22.
Gray, J. E.
1834. Enumeration of the species of the genus Terebra,
with characters of many hitherto undescribed.
Proc. Zool. Soc. London, 2: 57-63.
Hinds, Richard Brinsley
1844, Descriptions of new shells, collected during the
voyage of the Sulphur, and in Mr. Cuming's late
visit to the Philippines. Proc. Zool. Soc. London,
11; 149-168.
Keen, A. Myra
1958. SeashellsoftropicalWest America. xi+624 pp.,
illus. Stanford Univ. Press, Stanford, Calif.
Pilsbry, H. A. :
1931. The Miocene and Recent mollusca of Panama
Bay. Proc. Acad. Nat. Sci. Phila., 83: 427-440,
pl. 41, 5 textfigs.
Pilsbry, H. A, & H. N. Lowe
1932. West Mexican and central American mollusks
collected by H. N. Lowe, 1929—31. Proc. Acad.
Nat. Sci. Phila., 84: 33-144, pls. 1-17, 7 textfigs.
Explanation of Plate 5
Figure 1: Trivia myrae CAMPBELL, spec. nov. Dorsal view of holotype.
Figure 4: Nassarina anitae CAMPBELL spec. nov. Holotype.
Figure 5; Terebra berryi CAMPBELL, spec. nov. Holotype.
Figure 7: Terebra ninfae CAMPBELL, spec. nov. Holotype.
Figure 3: Same specimen, side view.
Figure 2: Same specimen, ventral view.
Figure 6: Drawing of protoconch of the holotype.
Figure 8: Drawing of protoconch of the holotype.
Figure 9: Terebra armillata Hinps, 1844. Drawing of protoconch. Figure 10: Terebra variegata GRAY, 1834.
Drawing of protoconch.
Figure 11: Terebra bridgesi DALL, 1908. Drawing of protoconch. Figure 12: Terebra
cracilenta Li, 1930. Drawing of protoconch. Figure 13: Terebra roperi Putspry & Lowe, 1932. Drawing of pro-
toconch. Figure 14: Terebra tuberculosa Hinps 1844. Drawing of protoconch.
(See text for actual measurements)
Tue VELIGER, Vol. 4, No. 1 [CAMPBELL] Plate 5
Figure 1 Figure 2
Figure 7
Figure 13 al Figure 14
Figure 9 Figure im - Figure 12
Vol. 4; No. 1
THE VELIGER
Page 29
Observations of the Effect of Diet on Shell Coloration in
the Red Abalone, Haliotis rufescens Swainson
by
Davip L. LEIGHTON
(Contribution from the Scripps Institution of Oceanography, New Series)
University of California,
La Jolla, California
(Plate 6)
The color of the ostracal shell layer of the
red abalone, Haliotis rufescens Swainson, 1822,
is typically a ''dull brick red'' (Cox, 1960). In
many specimens the continuity of the general
red aspect may be interrupted by zones of pink,
white or green coinciding in position with the
growth lines. Variation of shell color may be
extreme; the writer has collected many speci-
mens with predominantly or purely white shells.
The feature of variable shell color occurs, toa
greater or lesser degree, in all California aba-
lones and has been reported to occur also in
Japanese forms (Ino, 1952). Study has been
made of the effect of diet on shell coloration in
the topshell, Turbo cornutus (Solander, 1788),
by the same investigator (Ino, 1949, 1958).
Coloration of shells in mollusks has in sev-
eral other instances been linked with diet. Per-
haps most notable of these is the demonstration
that a diet of Mytilus edulis Linnaeus, 1758, is
accompanied by secretion of brown shell in
Purpura lapillus Linnaeus, 1758, and that feed-
ing upon barnacles (Balanus balanoides and
Chthamalus stellatus) results in deposition of
white shell material (Moore, 1936). Purpura
is, however, a carnivorous mollusk while the
abalones and topshells are herbivores.
The red abalone was selected for the present
study because the range of colors displayed in
shells is appreciable. Also minute, juvenile
and young adult individuals were easily obtained
in the required numbers.
Valuable advice given by Dr. Denis L. Fox,
Mr. John E. Fitch, and Dr. Wheeler J. North
is gratefully acknowledged.
Materials & Methods
Specimens of Haliotis rufescens were col-
lected, using SCUBA {Self Contained Underwa-
ter Breathing Apparatus}, at depths ranging
from 40 to 80 feet in waters near La Jolla, Ca-
lifornia. Greatest numbers of small specimens
(1.1 to 25.0 mm. in length) were found within
holdfasts of the giant kelp, Macrocystis pyri -
fera (Linnaeus) Agardh, 1771. Larger individ-
uals were commonly taken from beneath rocks.
Kelp plants torn from the bottom and washed
ashore by storms often yielded minute abalones
in large numbers. As many as 23 specimens
were retrieved from a single beached holdfast.
Interestingly, the abalones collected from hold-
fasts were largely of the species H. rufescens,
though occasionally H. corrugata Gray, 1828,
and H. assimilis Dall, 1878, were taken. Only
among specimens less than 3 mm. long was
there any confusion as to species. Identifica-
tion of smaller individuals was made by com-
parison with a carefully graded series of juve-
nile shells of each locally occurring abalone
species.
Sizes of experimental animals ranged from 4
to 120 mm. in shell length. The smallest were
maintained in one-quart polyethylene containers
perforated with numerous holes } inch in diam-
eter and submerged in a wooden frame rack in
circulating aerated sea water. The largest ani-
mals were held in 100-gallon concrete tanks
similarly supplied with sea water. Low light
intensities were maintained in the laboratory at
all times precluding adventitious growth of al-
gae.
Food consisted primarily of a variety of spe-
cies of marine algae representing the brown,
red and green algal classes. On several occa-
sions such foreign materials as boiled potato,
carrot, and yam were given.
Feeding experiments were conducted through-
out one year during which time over 50 abalone
were held for varying periods of time on re-
stricted diets.
Results
Shells of juvenile Haliotis rufescens collected
from holdfasts.of Macrocystis were purely
white, pale green or, when found in holdfasts
with epiphytically growing red algae, were par-
Page 30
THE VELIGER
Vol. 4; No. 1
tially red. Predominantly red-shelled speci-
mens were collected about rocks which sup-
ported red algae.
In the laboratory a wide variety of red algal
species (Rhodophyta) were provided as food
with the consistent result that red pigmented
shell was deposited. Pink color appeared in
newly deposited shell after red algae had been
given together with brown algae (Phaeophyta).
When brown algae, green algae (Chlorophyta), or
a number of foreign foods (e. g., potato or yam)
had been given, the shell formed was either
white, cream, or green, but never red. Re-
sults of all feeding experiments are given tabu-
lar summary below (see Table I).
Diets of brown algae, Macrocystis and Lam-
inaria, were found to result in either white or
Table 1:
green color in the newly formed shell. Other
brown algae (e.g., Egregia and Eisenia), when
ingested, influenced the formation of cream
colored shell. Pelvetia fastigiata (Agardh) De
Toni, 1895, gave origin to olive-green shell
coloration.
Individuals fed alternately red and brown al-
gae for one-month periods displayed similar
growth rates on either diet. Shells of these
abalones show alternate banding of red and
white. It is also noteworthy that growth was
substantial in abalone fed either potato or yam.
Light green shell was produced on occasions
when those diets were provided.
The multiple-colored specimen shown in the
accompanying plate (Lower, right) is one of nu-
Coloration of shell (ostracum)
secreted by Haliotis rufescens on restricted diets
Diet Color of Shell Number of
Produced Observations
Rhodophyta (Red Algae)
Pterocladia pyramidale red 12
Plocamium pacificum. red 20
Gelidium purpurascens red 6
Gelidium nudifrons red 5
Gigartina spinosa red 25
Gigartina californica red 7
Gigartina canaliculata red 8
Phaeophyta (Brown Algae)
Macrocystis pyrifera white or pale green 44
Laminaria farlowu green 18
Egregia laevigata white or cream 8
Eisenia arborea cream
Pelvetia fastigiata olive green 5
Chlorophyta (Green Algae)
Ulva sp. pale green 8
Miscellaneous
Potato, boiled pale green 6
Yam, boiled pale green 6
Carrot, boiled cream 4
Agar cream 4
Explanation of Plate 6
Figure 1: ” Normally‘‘ pigmented juvenile Haliotis rufescens collected from a kelp bed with a mixed benthic
flora. Figure 2: White phase abalone collected from a holdfast of the giant kelp Macrocystis pyrifera.
Figure 3: | Multiple-colored specimen with bands formed as a result of restricted feeding in the laboratory.
For convenience, diets and their respective color bands are listed in reverse order of their deposition: Evsenia
(shell margin), cream; Pterocladia, red; Potato, green; Miscellaneous red algae, red; Potato, pale green;
Gigartina, red; Macrocystis, pale green. For additional explanation of this specimen, see text.
(The scale adjacent to the specimens represents ten millimeters)
THE VELIGER, Vol. 4, No. 1 [LeicHTon] Plate 6
Figure 1
Figure 2 Figure 3
Vol. 4; No. 1
THE VELIGER
Page 31
merous abalone which were maintained in the
laboratory for as long as six months. When
collected from beneath a rock on June 4, 1960,
in the Point Loma Kelp Bed, it was 18.5 mm.
long and had a predominantly red shell. When
the animal was removed from the shell on Jan-
uary 15,1961, it was 37.5 mm. long and its
shell displayed bands of several colors. Initi-
ally, Macrocystis and Egregia were provided
for a period of three months. The two minor
red bands at the center of the shell were formed
during brief feedings of Plocamium. Subse-
quently, it was supplied the foods listed in the
explanation of the plate.
Microscopic epiphytes were found to contain
substances influencing the formation of red
shell. These small filamentous plants were
probably red algae. A future study designed to
demonstrate the effects of ingestion of sessile
diatoms and other microflora on shell pigmen-
tation will provide an answer to the question.
In all feeding experiments of the present study
care was taken to select fronds of algae which
were free of epiphytic growths.
Discussion
White shell is secreted by Turbo cornutus,
when restricted to a diet of the brown alga Ei-
senia bicyclis (Kjellman) Setchell, 1887, a Ja -
panese form similar to our local E. arborea
Areschoug, 1876, (Ino, 1949). Shell of 'normal"
color was secreted when both E. bicyclis anda
coralline alga (Rhodophyta) were given. A diet
consisting of a species of Sargassum influenced
the production of white shell, while a species of
Gelidium yielded a mixture of green-black and
brown color. MHaliotis rufescens, similarly,
has been found in the present study to secrete
cream colored shell when fed Eisenia and red
shell when fed Gelidium.
The red pigment of shells of Haliotis rufes-
cens has been given the name rufescine (Dhéré
and Baumeler, 1930).. Chemically it is a bilin,
or member of a biochemical category of pig-
ments including bilirubin and biliverdin, pig-
ments of bile. The relationship of rufescine to
phycoerythrin (the red pigment of red algae and
also a bilin) has yet to be demonstrated. The
nature of the other pigments of abalone shells
must also be ascertained through careful bio-
chemical analyses. These studies will be un-
dertaken by the writer in the near future.
Ecologically there may exist a considerable
utility in the type of study described in this pa-
per, for a dietary history may be obtained from
examination of the pigments in a given shell.
Food habits may be indicated by the presence
and arrangement of pigments. In abalones,
which do not, as a rule, move about extensively
(Cox, 1960) and which normally live in excess
of a decade, the sequence of color bands ina
number of specimens may indicate trends in
floral succession occurring over periods of
several years in their home area, A series of
eight Haliotis rufescens was collected from
the Point Loma Kelp Bed which shows impress-
ive similarity of shell color arrangement and
extent. The known kelp history of the area
agrees well with the observed pigment patterns.
Additional support is given this hypothesis by the
findings of almost purely white adult H. rufes-
cens by commercial abalone divers in areas
rich in brown algae (primarily the giant kelp,
Macrocystis) slightly south of Point Conception,
and of ''blood red'' specimens from the red alga
beds north of Point Conception (B. Owens, per-
sonal communication).
Conclusions
Pigmentation of the ostracum of the red aba-
lone reflects the diet in nature. Feeding expe-
riments performed in this study indicate that
the source of red pigments may be found in the
red algal foods. Shells lacking the typical red
coloration are formed by abalone living in
areas or niches with brown or green algae, but
which lack red algae. A minor quantity of red
algae in the diet may be detected as a diluted
red pigment in the shell. Since red algae will
usually form at least a minor portion of the
diet of Haliotis rufescens in nature, the most
prevalent color of the ostracum of H. rufescens
is red. Color sequences in shells of this spe-
cies may be used as a key to botanical succes-
sion in the home area of the respective ani-
mals.
Summary
Common collection of Haliotis rufescens with
little or no red shell pigmentation from areas
devoid of red algae, but rich in brown algae,
suggested that shell pigmentation was related
to the foods available. These observations
prompted an investigation of the effect of re-
stricted diet on shell pigmentation. Through
controlled feeding in the laboratory it was con-
sistently demonstrated that a primary source
of red shell pigment is redalgae. Feedings of
brown or green algae resulted in secretion of
shell of white or green color, but never red.
The ecological and biochemical significance
of these observations is discussed.
Page 32 THE VELIGER Vol. 4; No. 1
Literature Cited 1952. Biological studies on propagationof Japanese aba-
Cox, K. W. lone (Genus Haliotis). Tokai Reg. Fisheries Res.
1960. Review of the abalone of California. Calif. Fish Jase,
and Game, 46: 381-406, 9 figs.
Dhéré, C., & C. Baumeler
1930, Recherches sur la rufescine, pigment de la co-
quille de 1l'Haliotis rufescens. Arch. Int. Physiol.
348 2),
Ino, Takashi
1940. The effect of food on growth and coloration of the
topshell, (Turbo cornutus Solander). J. Mar. Res.
8: 1-5.
Lab. Tokyo, Japan.
1958. Ecological studies of the topshell, Turbo cornutus
(Solander). Bull. Tokai Reg. Fisheries Res. Lab.
Tokyo, Japan, 22: 33-36.
H. B.
The biology of Purpura lapillus. I. Shell variation
J. Mar. Biol. Ass. U. K.
Moore,
1936.
in relation to environment.
21: 61-89.
Notes on the Mitridae of the Eaciem Pactae
I - Mitra fulton: E. A. SmitH
by
GALE G. SPHON, JR.
Conchological Club of Southern California, Los Angeles 7, California
(Plate 7 and one Textfigure)
Introduction
This is the first of a series of papers to be
presented on the family Mitridae as it occurs
in the Eastern Pacific area. It is my plan to
take the four subgenera occurring in the area,
Mitromica, Scabricola, Strigatella, and Tiara,
separately, beginning with the subgenus Striga-
tella, and discuss each of the species involved
in a series of papers, one paper to each spe-
cies.
Mistory S Classification
The family Mitridae was segregated from the
Linnaean Volutidae by Réding (1798). The ge-
nus Mitra is usually incorrectly attributed to
Lamarck (1799), but as Réding's work precedes
that of Lamarck by one year, he is the author
for the family and the genus under the rules of
priority.
Mitra is an extremely large genus that has
over a thousand specific and subspecific names
assigned toit. Some authors have elevated sub-
genera to generic rank; there may be merit in
this in the case of those subgenera which are
well defined. This makes it possible to demon-
strate, by the use of nomenclature, the closer
relationship among the species of a subgenus.
In other words, it would allow for one more
subdivision which might be desirable in some,
but not all, cases. If this system of classifica-
tion were followed to its logical conclusion, the
end result would bea large number of mono-
typic genera and subgenera.
It is my feeling
that the whole group is closely enough knit to
use the single generic name Mitra; this is the
classification which will be followed in this
series of papers.
Description
The present paper deals with one of the rarer
species of the subgenus Strigatella Swainson,
1841. Mitra fultoni was described by E.A.
Smith (Ann. Mag. Nat. Hist., S. 6, vol. 9, 1892)
and as far as I can ascertain, this is one of
those few species which have no known syno-
nyms.
Smith gives a good diagnosis in Latin and then
goes on to complete his description with the
English equivalent. Only the English descrip-
tion is quoted here:
"This species is well characterized by the
punctate sulci, the punctures falling in reg-
ular longitudinal rows, through which pass
well-marked impressed lines of growth. It
has, I believe, been confounded with M. ori-
entalis, Gray, by some conchologists; but
from that species it may be sufficiently
distinguished by the above-mentioned fea-
ture and the difference of form. The whorls
are more convex, the epidermis blacker,
and the fine spiral striae which adorn the
surface of that species are scarcely indi-
cated in the present form."
Mitra fultoni has a thin, smooth, shiny black
periostracum which is often worn, revealing
the brown shell underneath. The interior of
SAN MARTIN ISLAND &
Vol. 4; No. 1
THE VELIGER
Page 33
the shell is white in the adult, while the younger
specimens show brown through the white enam-=
el. The soft parts of the animal are white.
This holds true for all members of the genus
examined by me so far.
Identification @ Relationship
For identification of the species the main
features are the punctate sulci mentioned by
Smith in his description. These sulci appear
not only spirally as he mentions, but also axi-
ally. The over-all appearance is similar toa
checkerboard with the small pits in the corners
of the squares. The pits occur even on the base
of the nuclear whorls. Other Eastern Pacific
species of Mitra which have this peculiar pit-
ting are M. lens Wood, 1828; M. orientalis
Gray, 1834; M. idae Melvill, 1893; and M. mon-
tereyi Berry, 1920.
Mitra fultoni can be distinguished from M.
lens by the former's smooth appearance. The
pitting in M. lens is limited to the area between
the coarse, flattened axial ribs, the pits are
o POTRERO CANYON
2 PLAYA DEL REY
© NEWPORT BAY
QPUERTECITOS
®PUNTA FINAL
SAN BARTOLOME BAY &*
WHALE ROCK WK SANTA INEZ BAY
MAGDALENA BAYS
LEGEND
© FOSSIL LOCALITIES
@ RECENT LOCALITIES
@ TYPE LOCALTY (PT. ABREQJOS)
twice the size of those occurring in M. fultoni,
and are not nearly as numerous. In M. orien-
talis, M. idae, and M. montereyi the pitting oc-
curs only on the spire and upper part of the
body whorl whereas in M. fultoni the pitting
covers not only the spire but the entire body
whorl as well.
Mitra fultoni is one species of a complex
(including M. idae, M. orientalis, and M. mon-
tereyi) within the subgenus Strigatella. The
Peruvian species, M. orientalis, is the only
other member of this complex whose range
may possibly extend into the Panamic area.
Both M. idae and M. montereyi are members
of the Californian province, exclusively. The
shape of M. montereyi and its slight pitting,
which must be seen with a good hand lens or a
microscope, suggest that M. fultoni and M.
montereyi are closely related members of this
particular group within the subgenus. Perhaps
when further research on the radular structure,
cytology and genetics has been done on the spe-
cies involved, it will be possible to make more
definite statements on the relationships within
this complex.
COCOS ISLAND ©
Page 34
THE VELIGER
Vol. 4; No. 1
Type Locality G Range
Smith gave the type locality for Mitra fultoni
as Point Abreojos, Lower California, Mexico,
Latitude 27° N., Longitude 113° W., on the Pa-
cific side of the peninsula. Williamson (1906)
quotes Dall as saying that he had seen no spe-
cimens from north of San Diego; in Burch (1945)
S. Stillman Berry is reported as having taken
specimens at San Diego.
One specimen in the California Academy of
Sciences (no. 17'749 Cal. Acad.) extends the
range to Santa Inez Bay in the Gulf of Califor -
nia. I have dredged one dead specimen (Sphon
Collection, no. 504) and a fragment of another
from Punta Final, near Bahfa San Luis Gonza-
ga. I have also taken a living specimen at
Puertecitos (Sphon Collection, no. 503), which
extends the range still farther up the eastern
side of Baja California. Another specimen (no.
23'077 Cal. Acad.) extends the range south to
Cocos Island, approximately 300 miles off the
coast of Central America. As far as it is pos-
sible to ascertain, there are no records, even
vague ones, to place the species on the main-
land of Mexico or Central America. It is on the
basis of the specimens mentioned that the range
extensions are recorded.
The majority of specimens I examined have
come from the type locality. This is in the
area where the greatest overlap between the
Californian and Panamic faunas occurs; since
there are far more records for the species oc-
curring in the more northern Californian prov-
ince, the species probably should be considered
as a member of that province and merely a
"straggler'' in the Panamic area. At least it
should be so considered until more specimens
from the Panamic area come to light.
Fossil Record
The fossil records of Mitra fultoni are of no
help in assigning it to a particular province.
There are three localities in Southern Califor-
nia from which fossils of the species are known:
Valentine (1956) reports it from Potrero Can-
yon in the Pacific Palisades area of Los Angel-
es County; Kanakoff and Emerson (1959) from
the Newport Bay Mesa area in Orange County;
and George Willett from the Lincoln Avenue
deposit in the Playa del Rey area of Los Angel-
es County. These three records are from warm
water, Upper Pleistocene deposits and area
mixture of what are now the Californian and
Panamic faunas.
Type Specimen
In the original description of the species,
Smith gives the dimensions of the holotype as;
"Long. 39 mm., diam. 13; aperture 194 long;
5 lat.'' This would indicate that the type is an
adult shell but not quite the maximum size as
there are larger specimens in the Stanford
University collection.
Smith figured the species with a line drawing
and the only other illustration of the species
which I can locate is in a paper by Mrs. Burton
Williamson (1906). Smith did not say where
the type was deposited; however, it is now in
the British Museum (Natural History), B. M.
(N. H.) Reg. No. 92.2.2.38.
Summary
Mitra fultoni E. A. Smith, 1892, is a rare
species which belongs to the subgenus Striga-
tella and to the Mitra orientalis-idae complex
within the subgenus. It is, however, quite dis-
tinct from all other members of that complex
due to the distinct pitting which occurs in both
axial and spiral lines on the spire as wellas
the entire body whorl.
The known Recent range of the species is
along the Pacific Coast from San Diego, Califor-
nia, to the type locality into the Gulf of Califor-
nia to Santa Inez Bay, Punta Final, and Puerte-
citos on the eastern side of Baja California,
and further south to Cocos Island. The species
is not known to occur on the mainland of Mexi-
co or in Central America. Probably it should
be considered as a member of the Californian
faunal province even though there are a few
records of its occurrence in the Panamic prov-
ince.
Specimens Examined
(Recent and Fossil)
1, 15 specimens from Potrero Canyon, Upper
Pleistocene, Los Angeles County, Califor-
nia. UCLA Geology Collection (Sp. Cat.
10'078).
2. 12 specimens from Lincoln Avenue deposit,
Upper Pleistocene, Playa del Rey, Los An-
geles County, California. los Angeles
County Museum Collection (Loc. #59 S.
218, Willett Collection).
3. 4 specimens from Newport Bay Mesa, Up-
per Pleistocene, Orange County, Califor-
nia. Los Angeles County Museum (Loc,
#66-2),
Explanation of Plate 7
Figures 1 to 3: Mitra fultom E. A. Smiru, 1892
(a: dorsal aspects - b: ventral aspects; all figures x 2.5)
Figure 1: Puertecitos, Baja California, Mexico (Sphon collection no. 503)
Figures 2 and 3: Whale Rock, near Point Abreojos, Baja California, Mexico
(Cate collection no. M - 46)
Tue VELIGER, Vol. 4, No. 1 [Spoon] Plate 7
Figure 2 a
Vol. 4; No. 1
THE VELIGER
Page 35
4.
10.
Mile
12,
13.
14,
I5)5
‘California, Mexico.
1 specimen from San Martin Island, Baja
California, Mexico. California Academy
of Sciences (24'041 Cal. Acad.).
3 specimens from San Bartolome Bay
(Turtle Bay), Baja California, Mexico. Los
Angeles County Museum (Willett Collec-
tion).
2 specimens from Whale Rock area (near
Point Abreojos), Baja California, Mexico.
Cate Collection (46).
1 specimen from Point Abreojos, Baja Ca-
lifornia, Mexico. Los Angeles County Mu-
seum (Willett Collection, A.375).
4 specimens from Point Abreojos, Baja
California, Mexico. California Academy
of Sciences (367'002 Cal. Acad.).
1 specimen from Point Abreojos, Baja Ca-
lifornia, Mexico. California Academy of
Sciences (24'065 Cal. Acad.).
10 specimens from Point Abreojos, Baja
Stanford University
(4'543). (It is believed that these speci-
mens may be part of the original collection
of the species.)
1 specimen from Magdalena Bay, Baja Ca-
lifornia, Mexico. California Academy of
Sciences (24'063 Cal. Acad.).
1 specimen dredged from 35 fathoms from
Santa Inez Bay, Baja California, Mexico.
California Acad emy of Sciences (17'749
Cal. Acad.).
1 specimen and a fragment dredged from
50 fathoms off Punta Final (near San Luis
Gonzaga Bay), Baja California, Mexico.
Sphon Collection (504).
l specimen from Puertecitos, Baja Cali-
fornia, Mexico, Sphon Collection (503).
{Note: This number should also appear in
the explanation of Plate 7. —-Ed.}
1 specimen from Cocos Island, off Costa
Rica. California Academy of Sciences
(23'077 Cal. Acad.).
Additional Records
Three specimens from Point Abreojos,
Baja California, Mexico. American Muse-
um of Natural History, New York. (Arnold
Constable Collection).
. Two specimens from Point Abreojos, Baja
American Museum of
(I. S. Oldroyd
California, Mexico.
Natural History, New York.
Collection).
One specimen from Point Abreojos, Baja
California, Mexico. British Museum (Na-
tural History). Holotype. B. M. (N.H.)
Reg. No. 92.2.2.38.
Acknowledgment
I wish to extend my thanks to Miss Jane Tur-
ner of the Botany Department at the University
of California at Los Angeles for her excellent
chart. Mr. S. P. Dance of the British Museum
(Natural History) was kind enough to confirm
the location of the holotype. Mr. Todd Scho -
walter took the photographs.
My deepest thanks are extended to Mr. and
Mrs. E. P. Chace of the San Diego Natural His-
tory Museum for making their own collection
and that of the Museum available for me to use
and for furnishing some much needed litera-
ture. I also wish to thank Dr. Myra Keen of
Stanford University for her kind help and for
making the Stanford University collection avail-
able for study. The staff at the California
Academy of Sciences: Drs. G. Dallas Hanna and
Leo G. Hertlein and Mr. Allyn G. Smith, were
particularly kind in giving their time and help
in making specimens available. Dr. William K.
Emerson of the American Museum of Natural
History, New York, furnished some locality
records as did Mr. George Kanakoff of the Los
Angeles County Museum.
Many private individuals have helped me by
furnishing specimens, literature, references,
and suggestions. Among these are Mr. and
Mrs. Crawford N. Cate, Dr. Rudolf Stohler
Mr. and Mrs. John Q. Burch, Mr. and Mrs.
Joseph DuShane, and Mr.. Mark Rogers. Last-
ly, I wish to give special thanks to one person
in particular, Mrs. Faye Howard, without
whose help, advice, and criticism this paper
would not have been completed.
Literature Cited
Burch, John Q. (ed.)
1945. Minutes of the Conchological Club of Southern Cal-
ifornia, No. 49: 32.
Grant, U.S., IV, & H. R. Gale
1931. Catalogue of the marine Pliocene and Pleistocene
mollusca of California and adjacent regions. Mem.
San Diego Soc. Nat. Hist., vol. 1, 1036 pp., 15 text-
figs., 32 plts.
Kanakoff, G. P, & W. K. Emerson
1959. Late Pleistocene invertebrates of the Newport Bay
area, California. Los Angeles County Mus., Contr.
in Sci. 31: 1-47, 5 figs., 3 tables.
Keen, A. Myra
1958. Sea shells of tropical West America. xi, 624p.,
illus. Stanford Univ. Press, Stanford, Calif.
Linnaeus, C.
1758. Systema naturae per regna tria naturae. Editio
decima, reformata.
Lamarck, J. B. P. A.
1799. Prodrome d'une nouvelle classification des co-
quilles. Mem. Soc. Hist. Nat. Paris, 1: 63-91.
Melvill, J. C.
1893. Description of a new species of Mitra.
chologist 2(6): 140.
The Con-
Page 36 THE VELIGER Vol. 4; No. 1
Valentine, J. W.
1956. Upper Pleistocene mollusca from Potrero Canyon,
Pacific Palisades, California. Trans. San DiegoSoc.
Nat. Hist. 12(10): 197.
Oldroyd, I. S.
1927. Marine shells of the west coast of North America.
Stanford Univ. Press, vol. 2 (1): 297 pp., 29 plts.
Réding, P. F. sls
1798. Museum Boltenianum. Hamburg. viii + 199 pp. Williamson, Martha Burton
Smith, E. A. 1905. Some west American shells - including a new var-
iety of Corbula luteola Cpr. and two new varieties of
gastropods. Bull. Southern Calif. Acad. Sci. 4; 123
to 124.
1906. West American Mitridae - north of Cape St. Lucas,
Lower California. Proc. Biol. Soc. Washington,
19: 193-197.
1892. Descriptions of new species of shells from Mauri-
tius and California. Ann. Mag. Nat. Hist., ser. 6,
9: 255-256.
Swainson, W.
1840. A treatiseon malacology. London, viii + 419 pp.
130 text figs.
On Cypraea tigris schilderiana Cate
by
ALISON Kay
Department of General Science, University of Hawaii, Honolulu, Hawaii
(This paper is Contribution No. 156 from the Hawaii Marine Laboratory)
(Plate 8 and 2 Textfigures)
Cypraea tigris schilderiana from the Ha- Figure 1 (see Plate 8), a length-frequency
waiian Islands has recently been described by histogram, shows that whereas specimens of
Cate (1961b). The purpose of this paper is to Cypraea tigris throughout the Pacific range in
contribute further measurements and records length between 51 and 147 mm., the majority of
of C. tigris both in the Hawaiian Islands and Hawaiian examples is longer than 100 mm., and
elsewhere in the Pacific, and to suggest three the majority of those from other areas of the
matters which merit further consideration: 1) Pacific, with the exception of Johnston Island,
the possible existence of C. tigris schilderiana is less than 100 mm. in length. Of the 403 in-
outside the Hawaiian Islands; 2) the difference dividuals measured, only two specimens from
in habitat exhibited by C. tigris in the Hawaiian the Hawaiian Islands were less than 95 mm. in
Islands and elsewhere in the Pacific; and 3) the length, and only nine from other Pacific areas
variability in size of the subspecies within the excluding Johnston Island were greater than 95
Hawaiian chain of islands. mm. in length. The mean length of the Hawaii-
Shell measurements of Cypraea tigris were an epechnens ca sute woe agen the mean
; 5a f. ee : length of those from other Pacific areas ex-
obtained from material in the Bernice P. Bishop ;
; i cluding Johnston Island was 77 mm.
Museum in Honolulu, and from numerous pri-
vate collections in the State of Hawaii. A de- The difference in size between specimens
tailed geographical analysis of the specimens of Cypraea tigris from the Hawaiian Islands and
examined is summarized in Table 1, Measure- Johnston Island, and those from elsewhere in
ments of length, breadth, and height were made the Pacific is further emphasized by compari-
to the nearest millimeter with vernier calipers, sons of breadth and height (Table 1).
and the means and standard deviations for these
measurements calculated. The shells were The figures for the coefficient of differ-
separated into three groups: 1) those from the ence for the three linear dimensions of 1.9 for
Hawaiian Islands; 2) those from Johnston JIs- length, 1.9 for breadth, and 2.0 for height indi-
land; and 3) those from other Pacific areas. cate that 95 percent of the Hawaiian population
The 75 percent rule for subspecies as set forth is different from the Pacific population (exclud-
by Mayr, Linsley, and Usinger (1953) was ap- ing Johnston Island) for these dimensions. Fol-
plied by use of calculations of the coefficient of lowing Mayr, Linsley, and Usinger (1935), the
difference. Hawaiian population, considered only from the
Tue Veticrr, Vol. 4, No. 1 [Kay] Plate 8
Marshall Islands
Line Islands
Fiji
Society Islands
Samoa
Caroline Islands
a
[|
E
x3)
o
[on
wn
a
{e)
ro
oO
E
Z
Marianas Islands
Philippines
Z
Johnston
Hawaiian Islands
go
Length in millimeters
Figure 1: Length-frequency histogram showing the variations in size of Cypraea tigris
in the Hawaiian Islands and other Pacific areas
Ae
j a: ty i ie >
| ’ 7 . rae
Vol. 4; No. 1
THE VELIGER
Page 37
Table 1: Geographical and Statistical Summary
S. D?
No. of Length
Locality Specimens | Range! | Mean!
Pacific 205 51-103
Marshall Is. 58- 96
Line Is. 79- 95
Samoa 64-100
Fiji 60- 96
Marianas Is. 59- 93
Caroline Is 51- go
Society Is. 74-103
Philippines 59- 90
1 Measurements in millimeters
standpoint of linear dimensions, may be sepa-
rated as a subspecies. No statistically signifi-
cant differences were found for calculations of
length/breadth and length/height ratios.
It is apparent from the data (Table 1 and
fig. 1, Plate 8) that samples of Cypraea tigris
from Johnston Island, which is 450 miles south
of the Hawaiian Islands, fall within the size
range of the Hawaiian specimens. The mean
dimensions of the Johnston Island specimens
are, however, slightly smaller than those of
the Hawaiian sample. Because of the small
sample of Johnston Island material, statistical
tests for this group have not been included.
However, it should be pointed out that the shells
from Johnston Island resemble those from the
Hawaiian Islands in color and lack a marginal
callus, characters which Cate (1961 b) considers
distinctive of the Hawaiian subspecies. An hy-
pothesis accounting for the occurrence of a
population of C. tigris at Johnston Island simi-
lar to the Hawaiian population has been put for-
ward elsewhere (Kay, manuscript in press).
A survey of the literature and discussions
with shell collectors in Hawaii disclose that the
habitat of Cypraea tigris in the Hawaiian Islands
is in contrast to its habitat elsewhere in the
Pacific. In the Hawaiian Islands C, tigris is
collected from depths of eight to ten feet and
deeper (with one or two records of collection at
depths of four to eight feet) where it is found
beneath dead coral and on basalt boulders and
spits. In other areas of the Pacific the species
has been described as occurring exposed on the
surface of large coral heads at Biak in the
Marshall Islands (Bayr and Neurohr, 1946), in
association with living coral heads on reefs be-
tween the tide marks in American Samoa (In-
gram, 1939), and on various lagoon and seaward
reefs in the Pacific (Demond, 1957).
Range!
Breadth
Mean!
Height
S. D2 | Range! | Mean!
80
Ud
2 Standard deviation
Specimens of Cypraea tigris in the Hawaii-
an Islands vary in size with both depth and lo-
cale. Cate (196la) has noted that ''the smaller
shells seem to come from the shallower locali-
ties in the southern end of the range, medium-
sized ones from the intermediate depths of mid-
range, and the largest known in the world from
deep-water Oahu stations.'' Data confirming
Cate's observations are presented in figs. 2
and 3. Although measurements for specimens
from the Island of Hawaii are few and incon-
clusive due to the unavailability of records at
the present time, the data from the islands of
Kauai, Oahu, and Maui indicate a tendency for
the larger specimens to be more abundant
around the northern islands, with the exception
of the extremely large specimens which were
collected from deep waters off Oahu. It is also
apparent that depth of collection varies with the
islands; individuals from Kauai, the northern-
most of the main islands in the Hawaiian chain,
having been found at depths of 25 to 40 feet,
while the islands to the south yielded collections
from progressively shallower water.
Two specimens of Cypraea tigris have been
examined from islands to the north of Kauai. It
is noteworthy that both, one from Midway and
the other from Kure, fall into the size range of
material from the Pacific, being 63 mm. and
75 mm. in length, respectively. They also ex-
hibit the callus and color pattern characteristic
of the Pacific type. The dimensions of these
specimens have not been included in the analy-
sis of data, because of their apparent similar-
ity to individuals from the other Pacific areas.
Another interesting feature concerning Cy-
praea tigris in the Hawaiian Islands is the
shortage of collections of juveniles. There are
only three known juveniles from Hawaii, all
three collected by Dr. C. M. Burgess. Other
Page 38 THE VELIGER Vol. 4; No. 1
Kauai
5
= Oahu
oo
A
2
QO,
a
we)
a)
-Q
g Maui
Z 5
Hawaii
5
80 go 100 110 120 130 140 150
(measurements in millimeters)
Figure 2: Length-frequency histogram showing the relationship between length and island for Cypraea tigris
in the Hawauan Islands
shell collectors with wide collecting experience
have remarked that they have been unable to
find juveniles of this species, although they are
well known for other species of Cypraea.
The history of Cypraea tigris in the Hawa-
iian Islands is sparse, Cate (1961 a) has noted
that Schilder (1933) considers early records of
Garrett (1879), Baldwin (1898), and Hidalgo
(1906) doubtful, and Cate regards the first au-
thentic record to be that of a specimen collected
in 1929 and recorded by Harris (1935), Cate's
(196la) reference to a record cited by Martens
and Langkavel (1871) is apparently an error;
the reference quoted refers to Terebra tigrina
from the Kingsmill Islands. Martens and Lang-
kavel (1871) do not refer to C. tigris from the
Hawaiian Islands.
Two early reports of Cypraea tigris in the
Hawaiian Islands have recently come to the au-
thor's attention. The naturalist on the Portland
and Dixon voyage to Hawaii mentions "',., beau-
tiful shells such as Cypraea tigrina.,,'' from
the Hawaiian Islands (Dixon, 1789). Another
record is that of W, H, Pease, who described
a specimen of "'...Cypraea tigris from your
island (Hawaii) five inches in length and over
nine inches around the thickest part...'' in a
letter to Andrew Garrett dated March 5, 1857
(Manuscript Collection, B, P. Bishop Museum,
Honolulu). As the dimensions mentioned by
Pease fall well within the size range of the Ha-
waiian specimens, it would appear that C. tigris
had been collected in the Hawaiian Islands prior
to the 20th century, although Pease notes in the
same letter, ''...I never obtained but few.!!
While there are no records of Cypraea ti-
gris in the Hawaiian archaeological collections
of the B. P. Bishop Museum or among fossils
from the late Pleistocene, there is one fossil
example of C, tigris which has been tentatively
dated as Recent. The shell was dredged from
not more than four feet below the surface of a
reef near Kaaawa, Oahu, in 1948 (R. Gage, per-
sonal communication), Ostergaard (personal
communication) observes that the specimen
cannot be assigned an age comparable to that of
the fossiliferous limestone of Oahu (usually
considered Late Pleistocene) because of its po-
sition below the surface of the reef,
Vol. 4; No. 1
THE VELIGER
Page 39
um _
5
q
oO
g
ra)
5 5
6
5
E
7,
5
go 100
110
8 to 20 feet
20 to 40 feet
70 to 100 feet
120 130
140 150
(measurements in millimeters )
Figure 3: Length- frequency histogram showing the relationship between length and depth of occurrence of
Cypraea tigris in the Hawaiian Islands
Conclusions
The data presented here, although some-
what scanty, are of interest not only with re-
spect to studies of Cypraea, but as materials
for speciation studies of marine invertebrates.
It is well known that isolated areas and those
near the periphery of a range are frequently in-
habited by populations with distinctive biologi-
cal characteristics. The existence of a dis -
tinctive population of C, tigris in the Hawaiian
Islands, an area isolated from other Pacific is-
lands by expanses of deep water and forming
the eastern periphery of the range of C. tigris
in the Indo-West Pacific is, therefore, not sur-
prising. That the distinctive features of peri-
pheral populations are correlated with ecologi-
cal factors is also a generalization, but there
is little information on the effects of ecological
factors. We are particularly handicapped by
the dearth of information and lack of experi-
mental studies concerning early stages in life
history, These data accumulated for adult C.
tigris are, nevertheless, useful in that they
focus on two major questions: 1) how are we to
account for the large size of C. tigris in the
Hawaiian Islands? and 2) is the size difference
exhibited by the Hawaiian population genotypic
or phenotypic?
Increased size in various animals has long
been associated with low temperatures (Berg-
mann's Rule), Since surface temperatures of
Hawaiian waters are at least 1.5°C. cooler than
the lowest surface water temperature of other
areas from which we haye records of C, tigris
(Sverdrup, et al., 1946), temperature must be
considered as a possible factor affecting the
size of individuals in the Hawaiian population of
C, tigris, But the picture is complicated by
the apparent presence of the Pacific type of C.
tigris at Midway and Kure which have the low-
est surface water temperatures in the Hawaiian
chain, and of the Hawaiian type at Johnston Is-
land which has a higher mean annual surface
water temperature than that of the Hawaiian Is-
lands (Sverdrup, et al., 1946), The occurrence
of specimens in relatively deep water in the
Hawaiian Islands compared with their existence
in shallow water elsewhere in the Pacific, and
variation in depth at which the species is found
within the Hawaiian Islands is another compli-
cating factor. Is there a relationship between
water temperature and depth which would ac-
count for the observations? Or is there a fac-
tor associated with a different habitat? The
Hawaiian Islands are high islands with reef
formation and constitution considerably differ -
ent from the atolls elsewhere in the Pacific.
Are there differences in food and feeding
habits? Is larval population density a factor af-
fecting adult size?
The question as to whether the differences
in size of Cypraea tigris are phenotypic or ge-
notypic is equally provocative. It has been
pointed out that slight geographical variation
has been demonstrated in many marine inverte-
brates and the suggestion has been made that
this sort of variation is a response to local en-
vironmental conditions (Mayr, 1954). The yari-
ation in size and depth of occurrence exhibited
Page 40
THE VELIGER
Vol. 4; No. 1
nn ————
by C. tigris within the Hawaiian Islands is pro-
bably best considered an example of this type
Is the presence of the Hawaiian
type of C.tigris at Johnston Island, where there
isa higher mean annual surface water temper-
ature and an atoll-type environment, indicative
of a genotypic difference when the pronounced
difference in size of the individuals in the Ha-
waiian population is considered?
of variation.
It is evident that there are no simple an-
swers to the two questions posed above. That
we are able to enumerate some of the factors
and speculate on their possible effects is en-
couraging. With the further accumulation of
ecological data and emphasis on experimental
work we may in time contribute further infor-
mation toward an elucidation of the factors af-
fecting speciation in marine invertebrates.
Acknowledgment
I would like to acknowledge the patience
and help of the many collectors in Hawaii who
have contributed their collections of Cypraea
tigris for measurement and their observations
for analysis.
Yoshio Kondo of the B. P. Bishop Museum in
Honolulu, to Dr. C. M. Burgess, Mr. and Mrs.
Arch Harrison, Mr. Harold Jewell, and Mr.
and Mrs. James Dennis of Honolulu, and to Mr.
John Duarte of Kauai.
Literature Cited
Baldwin,
1898.
DyDs
Hawaiian Cypraeidae. Nautilus, 11; 123.
I am particularly grateful to Dr.
Bayer, F. M., & G. N. Neurohr
1946. Life on a tropical Pacific reef. Mollusca, 1:
107-112.
Cate, Crawford N.
1961la. Redescription of Cypraea tigris lyncichroa Mel-
vill, 1888. The Veliger, 3 (3): 66-69, pl. 11.
1961b. Description of a new Hawaiian subspecies of
Cypraea tigris (Linnaeus, 1758). The Veliger 3(4):
107-108, pl. 19.
Demond, Joan
1957. Micronesian reef gastropods. Pacif. Sci. 11:
226-275.
Dixon, G.
1789. -A voyage roundthe world..... in the ''King
George" and "'Queen Charlotte." London.
Garrett, A.
1879. Annotated catalogue of the species of Cypraeidae
collected in the S. Sea Islands. Journ. Conch., 2:
105-128.
Harris, W.
1935. Cypraea tigris Linné in the Hawaiian Islands.
Nautilus, 49: 39-41.
Hidalgo, J. G.
1906-1907. Monografia de las especies vivientes del
genero Cypraea. Mem. Ac. Cienc. Madrid, 25:
1-588.
Ingram, W. M.
1939. Cypraeidae from American Samoa with notes on
species from Palmyra Island. Nautilus, 52: 103
to 105.
Martens, E. von, & B. Langkavel
1871. Stidsee - Conchylien: Donum Bismarckianum.
Berlin.
Mayr, E.
1954. Geographic speciation in tropical Echinoids.
Evol., 8: 1 - 18.
Mayr, E., E. G. Linsley, & R. L. Usinger
1953. Methods and principles of systematic zoology.
ix + 328 pp., 44 textfigs. McGraw-Hill, New York.
Schilder, F. A.
1933. Cypraeacea from Hawaii. B. P. Bishop Mus. Occ.
Pap., 10 (3): 3-22.
Sverdrup, H. U., M. W. Johnson, & R. H. Fleming
1946. The Oceans, their physics, chemistry, and gen-
eral biology. x + 1087 pp. New York.
Vol. 4; No. 1
THE VELIGER
Page 41
A Study of the Reproductive Cycle in the
California Acmaeidae (Gastropoda)
Part III
by
Harry K. FrircHman II
(From the Department of Zoology, University of California, Berkeley, California,
and the Department of Biology, Boise Junior College, Boise, Idaho)
(Plates 9 to 14)
Subgenus GCOLLISELLA DaLt, 1871
Acmaea pelta EscHSCHOLTZ, 1833
Ecology: This species was studied exclusive-
ly from the rocky point south of Rockaway
Beach, San Mateo County, California (37°30'25"
N.; 122°30'W.). The animals are common
there, but are not as numerous as Acmaea scu-
tum Eschscholtz, 1833. They seem to be about
equally distributed between zone 2 and zone 3,
and appear to require macroscopic algae upon
which to browse. Acmaea pelta is somewhat
more eurytopic than A. scutum and is often
found high on the rocks among tufts of Endo-
cladia and on rocks which are quite rough and
have considerable barnacle growth on them.
Acmaea scutum seems to prefer smoother sur-
faces and to avoid barnacles. There is alsoa
behavioral difference in that A. scutum seeks
moist, shady spots when the tide is out and may
migrate onto the under surfaces. Acmaea pelta
appears to be less sensitive to desiccation,
perhaps due to its increased shell height and
reduced aperture and may remain high on the
boulder surface in the full heat of the sun.
Collections: — These were begun on September
25, 1949, and continued to April 16,1952. At
the outset and until mid-September, 1950, the
sample consisted of about 25 animals. It be-
came apparent that this rate of collecting would
eventually deplete the population, and the num-
ber was reduced to 10 animals per collection
beginning September 26, 1950, and continuing at
this rate until the study was completed. A total
of 934 animals was studied; 483 males, 422 fe-
males, 27 indeterminate, and 2 immature.
Results: — By comparing (Plate 9) the spawning
record of Acmaea pelta with that of A. scutum
(Fritchman, 1961), it will be seen that there is
a striking similarity between the two. Both
species were studied over the same period of
time and from the same habitat. During this
interval A. scutum definitely spawned 12 times
and A. pelta 11 times. The spawnings of both
were restricted, for the most part, to the fall,
winter, and spring months. Despite this, how-
ever, there are only three times when the pe-
riods of spawning of the species coincide:
January 22 to February 5, 1950, March 28 to
April 11, 1950, and April 15 to April 28, 1951.
In addition to these, there are three other times
in which the spawning periods are adjacent in
time to one another. Those of A, pelta are Oc-
tober 9 to October 22, 1949; March 12 to March
28, 1950; and September 26 to October 14, 1950.
Thus, of the 11 spawnings of A. pelta, six ei-
ther coincide with or else precede or follow by
a maximum of only two weeks corresponding
spawnings by the other common limpet in zone
2, A. scutum.
Analysis of Environmental Conditions Coincident
with Spawning:
As explained for Acmaea scutum, only nine
spawning periods can be evaluated with refer-
ence to tidal and lunar phases, five being asso-
ciated with full moons (January 22, 1950, March
28, 1950, June 18, 1950, March 18, 1951, and
April 15, 1951) and four with new moons (Octo-
ber 9, 1949, February 5, 1950, March 12, 1950,
and September 26, 1950). It is thus impossible
to assess the role, if any, played by the moon
in the spawning of this species. As previously
noted with several other limpet populations, the
summer months seem to inhibit the reproduc-
tive activity of this species, only the minor
spawning of June 18 to July 2, 1950, occurring
in this period of high temperature. Since A.
pelta has been found to spawn at water temper-
atures’ ranging from 48.5°F. (January 22 to Feb-
Page 42
THE VELIGER
Vol. 4; No. 1
eee
ruary 5, 1950) to 60.0°F. (September 26 to Oc-
tober 14, 1950), or over the entire yearly mean
range of the surface water temperatures, it is
doubtful that the spawnings are initiated by cri-
tical temperature levels. However, tempera-
ture levels may, as indicated before, play a
role in rate of gonad redevelopment. During the
fall and winter of 1949 we again see a retarded
development of the gonads of A. pelta. The fall
months of 1950 and 1951, both of which were
warmer than 1949, each have two spawnings
during that period, while 1949 has but one. It
may be supposed that the spawning expected for
November, 1949, was postponed until January,
1950. If this is assumed, then a general yearly
cycle for A. pelta would appear to be as fol-
lows: one or two spawnings in the period of
January through April, this depending on the
mean temperatures of the winter:months (if
warm, as in 1950-1951 and 1951-1952 then pos-
sibly two, one in January and one or two close
together in March and April); a reduced repro-
ductive activity from May through August fol-
lowed by a spawning in late September and Oc-
tober and another about two months later in
November. Thus a total of three, or possibly
four, spawnings occur throughout the year.
Acmaea limatula moerchu Dau, 1879
Ecology: — This species does not occur in
numbers large enough to be studied north of
Monterey, California, except for a restricted
population found in Tomales Bay, Marin County,
California, where it is the dominant limpet, and
because of certain modifications of the shell
and its markings, it has been given the rank of
a subspecies, Acmaea limatula moerchii Dall.
While the open coast A. limatula have a low
shell which is predominantly yellow in color
and lacking in brown markings, the subspecies
moerchii has a high shell similar to that of A.
digitalis Eschscholtz, 1833, and is strongly
marked with brown. Both the animals from the
open coast and bay have the dorsal surface of
the foot pigmented with black but the degree of
this pigmentation is reduced in the bay form
and is rather grayish. In the bay this limpet is
very eurytopic and can be found from the upper
limit of the barnacle, Balanus, down to the low-
est limit uncovered by the tide. The east side
of the bay which receives the brunt of the wind
and waves from the northwest does not have
large growths of algae, probably because of the
constant deposition of silt. The only alga pres-
ent at the sites of collection was small amounts
of Ulva, the collections being made at about the
level of zone 2. The animals are scrapers of
the bare rock surfaces and ingest large amounts
of the mud which has settled on them.
Collections: — These were made from three
points located at distances of 0.9, 1.5, and 1.9
miles south of the post office of the village of
Marshalls (38°09'45'"'N.; 122°53'30"W.). Col-
lections from the most southern of these areas
were begun on October 10, 1950, from the cen-
tral area on December 31, 1950, and from the
northern area on March 25,1951. Each sample
consisted of 10 animals from an area and a to-
tal of 434 was examined; 156 males, 204 fe-
males, and 74 indeterminate. Collections were
made at monthly intervals, the final sample be-
ing taken on April 29, 1952. The graph on Plate
10 has been constructed using the information
gained from the study of limpets from all three
areas, there being little, if any, variation in
the reproductive cycles among residents of the
three populations,
Results: It seems apparent (Plate 10) that
this species is following a yearly cycle which is
different from any encountered thus far in the
study. However, the gonad of this species does
not develop the extreme turgor which charac-
terizes the ripe condition of many of the other
species and for this reason it is difficult to de-
termine when the gonad has progressed beyond
the partially ripe stage. These conditions were
not included in the plot since the spawned and
indeterminate conditions are readily noted and
provide a much more dependable basis for es-
tablishing the reproductive cycle. The collec-
tion of this species began on October 10, 1950,
at a time when the animals were just beginning
to spawn, as is evidenced by the 40 percent of
the first sample which were in the spawned
condition. The next collection, made on Novem-
ber 5, showed that approximately 50 percent
were indeterminate and it is seen that this con-
dition prevailed until December 31. By late
January, 1951, most of the animals were in the
process of redevelopment of the gonad, and this
condition prevailed until the next spawning sea-
son which began in September and reached its
peak in October. Here again indeterminate ani-
mals appear following the completion of spawn-
ing. Acmaea limatula moerchii thus spawns
during the time of year when mean sea and air
temperatures are at their maxima, a fact that
correlates with the southern distribution of the
species. The attainment of these temperature
levels undoubtedly determines the period of re-
production in this species. Possible stimuli to
spawning cannot be speculated upon since col-
lections were made only at monthly intervals.
Acmaea asmi_ (MippENDoRFF, 1847)
Ecology: This stenotopic limpet is restricted
to the shell of the trochoid gastropod, Tegula
Tue Veuicer, Vol. 4, No. 1 [FRITCHMAN] Plate g
Acmaea pelta EscuscHoxtz, 1833
Rockaway Beach
J of sample ripe
Y of sample spawned
% of sample of indeterminate sex
spawning period
laboratory spawning
THe Vewicer, Vol. 4, No. 1
Acmaea limatula moerchii Dati, 1879
Tomales Bay
Q% of sample sexually determinate
Q% of sample spawned
Vv spawning period
¥ laboratory spawning
[FRitcHMaN] Plate 10
Vol. 4; No. :
THE VELIGER
Page 43
funebralis (C. B. Adams, 1854), a common zone
2 resident. This small limpet gathers its en-
tire food supply from the diatoms and other
microscopic plant life which are present on the
Tegula shell. Casual observation of such shells
shows that they are singularly barren of vege-
table life and are all more or less eroded well
into the prismatic layer. If the fecal pellets of
the limpet are examined, it will be seen that
they are purplish in color due to the large per-
centage of shell particles which have been
rasped from the purple-black host. The poor
source of food and the zone 2 location in the in-
tertidal, which restricts feeding time and al-
lows considerable periods of exposure to heat-
ing and drying, would appear to combine to
render this unique habitat a rather unfavorable
one for a limpet, but the species seems quite
successful,
Collections: Acmaea asmi has been collected
from a rocky shelf which extends toward the
sea and lies just to the north of the mouth of
Sunshine Creek, a small stream found at Moss
Beach, San Mateo County, California (37°31'30"
N.; 122°31'W.). In this area Tegula funebralis
is found in abundance, congregated in masses
when the tide is out. The procedure was to col-
lect all of the A. asmi present on the Tegula
along this shelf at each collecting period. This
usually provided between 20-25 animals and, of
course, means that the area must be replen-
ished with Acmaea-bearing Tegula every two
weeks, Most of the immigrating Tegula prob-
ably came from the flat sandstone rocks to the
north of the shelf, this area having moderate
quantities of various algae growing upon it (Pe-
trocelis, Endocladia, Gigartina, Iridophycus,
and Cladophora). Since the creek runs not far
to the south, replenishment from that area is
unlikely,
Only the adult animals of this species have
been examined, and because of the fixed size
that the animals can attain, the thickness of the
gonad can be used, in combination with that of
degree of turgor, as an indication of ripeness.
An adult animal which is ripe will have a gonad
showing extreme turgor and which will be 1.5
to 2.0 mm. in thickness. Adult animals with
gonads of 1.0 mm. or less are considered par-
tially ripe or partially spawned, while spawned
animals are readily identified by the thinness of
the gonad and the small residual quantities of
eggs and sperm. The study of this species was
begun on November 5, 1949, and continued until
April 16, 1952. However, during the first eight
months, considerable difficulty was experienced
in identifying the condition of the animals and
for this reason these data have been discarded.
Plate 11, then, has been prepared from the data
of the collections of July 2, 1950, to April 16,
1952. A total of 851 animals was examined;
391 males, 446 females, 12 indeterminate, and
2 immature,
Results: Acmaea asmi from this area of col-
lection appears (Plate 11) to have a reproduc-
tive cycle which includes two spawnings per
year, one in the spring in March or April, and
another in the fall in September or October.
Following the spring spawning, the gonads are
redeveloped and remain in a ripe or partially
ripe condition during the summer, much as was
indicated for A. scutum. However, subsequent
to the fall spawning period, this species passes
into a rather strange condition, which extends
until the following March or April. During this
time the gonads are all much as they were at
the conclusion of the fall spawning, practically
none of them being ripe and a variable percent-
age spawned, but most of the animals possess
gonads of 0.5to 1.0 mm. in thickness amd
with moderate amounts of eggs and sperm.
This latent period during the winter period is
unusual since at that time all of the other spe-
ciesand populations studied are very active re-
productively. This spawning pattern, like that
found for A. limatula moerchii, reflects A. as-
mi's southern distribution and greater toler-
ance for and dependence upon high temperature.
Analysis of Environmental Conditions Coincident
with Spawning:
It is impossible to evaluate the effects, if any,
which lunar and tidal periodicity have on this
species because of the occurrence of periods of
both new and full moons during the interval
when spawnings are known to have taken place.
As regards temperature, little can be said here
since the temperatures at which reproductive
activity begins in March and April, 52° to 53°
F., is several degrees below that at which it
ends, in the cases recorded, 57° to 60°F. It is
thus difficult to designate any temperature as a
critical one as has been done for several of the
other species.
Acmaea digitalis EscHscHOLTz, 1833
(Rockaway Beach Population)
This species was studied from two
localities and these will be considered sepa-
rately. The first of these is a large concrete
breakwater which stands at the tip of the rocky
point which forms the southern boundary of the
community of Rockaway Beach, California.
This structure extends for about 100 feet ina
Ecology: —
Page 44
THE VELIGER
Vol. 4; No. 1
north-south direction and stands about 12 to 15
feet high. It is anchored in the center to a huge
boulder which divides it into north and south
portions. It was from the lee side of this north
section that extensive collections of Acmaea
digitalis were made. The upper surface of the
breakwater is never submerged although it is
dashed by the surf at high tide. The leeward
side furnishes a very uniform environment for
the large numbers of A. digitalis which live
upon it. The animals are completely protected
from the direct beat of the surf and receive
their water as it runs down the vertical face of
the concrete. They receive uniform illumina-
tion from the east and south and are exposed
twice daily to desiccation. The only source of
food is that of microscopic plant life left by the
surf and, during the winter, what algal film can
proliferate. Throughout the summer months,
the vertical faces appear to be burned bare of
any living algae. Theonly other animals in this
habitat consist of a few Littorina scutulata
Gould, 1849, L. planaxis Philippi, 1847, and
small barnacles, probably Balanus glandula
Darwin, 1854. During the summer this habitat
undergoes extreme heating and drying during
the occasional long periods of low tide. Collec-
tions made at such times find the limpets ina
state of extreme desiccation, the bodies of the
animals being shrunken and brown within their
shells, the foot being held to the substrate only
by a thin film of dried mucus. This degree of
desiccation is caused, to an extent, by the ver-
tical surfaces of the breakwater which almost
immediately drain and dry when the last wave
of the descending tide has passed over the
structure.
Collections: — These were begun from this habi-
tat in February, 1949, and continued to April
16, 1952. A totalof 9'902 animals was collected
during this period at intervals of two weeks;
3'415 males, 3'365 females, 3'084 indetermin-
ate, and 38 immature. The size of the samples
varied, Initially, the sample consisted of ap-
proximately 250 animals. This practice was
continued until February, 1950, at which time
the number was reduced to about 100 per
sample, A final reduction was made in Octo-
ber, 1951, to 50 animals per collection.
Results; — Two gonad conditions have been
plotted (Plate 12): percent of the sample which
was ripe and percent of the sample which was
of a determinate sex. The two features which
appear most evident from the plot of the repro-
ductive cycle of this population are the single
major spawning which occurs during the late
winter or spring months and the indeterminate
condition which is present during the summer
months. The major spawning depletes the go-
nads completely except for a few residual eggs
and sperm masses which remain for one to
several months and permit the sex of the ani-
mal to be ascertained. Sooner or later these
genital products are resorbed and the gonad
becomes indeterminate. The major spawning
may be preceded by a partial spawning in which
all or only part of the animals participate.
Such spawnings occurred in mid-January of
1951 and between late February and early
March, 1951, After such spawnings, the gonad
redevelops to a ripe condition prior to the ma-
jor spawning. During the winter months when
the gonads are developing, indeed, even before
they have attained a fully ripe condition, spon-
taneous spawnings occur in the storage jars in
the laboratory. Such occurred eight times during
the winter of 1950-1951 and five times during
the winter of 1951-1952. Accurate analysis of
the condition of the population is seriously in-
terfered with by this type of spawning and one
of two alternatives may be used to express the
data. One can assume that the population was
either ripe or was approaching that condition
and plot the sample as being ripe. This prac-
tice was followed for the 1950-1951 data. Or
one can plot the actual condition of the sample
when it was examined regardless of the degree
of spawning which had taken place as was done
in 1951-1952. The former method is preferred
when the population is known to have reached a
fully ripe condition prior to the first spontane-
ous spawning. There will usually be a few ani-
mals which will not have spawned and will, by
the ripe condition of their gonads, give an indi-
cation of the status of the population at the time
of collection. However, if the development of
the gonad has been retarded asit was in the fall
of 1951 and spontaneous spawning occurs prior
to the fully ripe condition, then it is impossible
to estimate what the natural condition of the
population is at the time of collection. In such
a case it is perhaps preferable to plot the data
as it actually appears at the time of examina-
tion. This accounts for the low percentages of
ripe animals which appear during the first two
months of 1952.
Analysis of Environmental Conditions Coincident
with Spawning:
Of the five spawning periods which can be ana-
lyzed relative to the influences of lunar and ti-
dal factors, namely April 24 to May 9, 1949
May 9 to May 23, 1949, March 18 to April 1,
1951, April 1 to April 15, 1951, and March 31to
April 16, 1952, three are associated with full
moons and two with new moons. The maximal
ae
Vol. 4; No. 1
THE VELIGER
Page 45
tidal ranges of these periods are 6.1, 8.0, 6.5,
5.8, and 6.5 feet. The only spawning period
which was accurately reduced to an interval of
seven days was that of February 5 to 12, 1950.
This was a complete spawning so there could
be no possibility of misinterpretation when the
animals were examined. Here the tidal fluctu-
ations were only moderate beginning with 5.8
on February 5 and continuing as follows: 5.0,
4.5, 5.0, 5.5, 6.0, 6.5, and 6.8 feet on Febru-
ary 12. Furthermore, this period does not fall
under the influence of either a full or a new
moon. Thus, there is no reason to suppose that
the spawning periods of this population of high
intertidal limpets are correlated with lunar pe-
riodicity and the attendant high tides. The sur-
face water temperatures at which spawnings
occurred are as follows: 1949, 55.0°F.; 1950,
49.5°F.; partial spawning 1951, 50.5°F.; com-
plete spawning 1951, 52.0°F.; partial spawning
1952, 53.0°F.; and complete spawning 1952,
53eo B,
Acmaea digitalis EscuscHo.rz, 1833
Ecology: — (Moss Beach Population) There are to
be found at Moss Beach, San Mateo County,
California, a series of large stone ledges lo-
cated about 100 yards north of Sunshine Creek.
These rocks are from eight to ten feet high and
arevery soft sandstone. They are seldom com-
pletely submerged but are wetted daily by the
wash of the tide. The broad horizontal surfaces
of the tops of these rocks have several small
tide pools which support heavy growths of cor-
alline algae. The top three or four feet of the
vertical sides of the rocks support large colo-
nies of limpets, Many occur in small depres-
sions in the rock, either singly or in clusters
of several per depression. Of these limpets,
Acmaea digitalis is the most abundant, but A.
scabra is also very common. The rock at this
height supports a moderate growth of green al-
gae, Ulva being the predominant type. In addi-
tion to the macroscopic algae, there is a film
of microscopic plant life covering the rock.
The porous rock allows these plants to thrive
at a height which would be impossible, were the
rock more dense, since the substrate is always
-damp. The limpets in feeding scrape up large
amounts of sand and with it the attached plant
life. Food is plentiful and, in addition, desic-
cation is reduced by the nature of the rock. All
of the limpets were collected from the northern
side of the northernmost of these rocks, thus
assuring that they were taken from the most
favorable area of this habitat which, because of
its continued dampness, corresponds roughly
to lower zone 2 or upper zone 3.
Collections; — These were begun in February,
1951, and extended to April, 1952. They were
made every two weeks, each sample consisting
of 15 animals. A total of 416 was examined;
247 males, 156 females, 5 indeterminate, and
8 parasitized.
Results: — The most striking difference (Plate
13) between the reproductive cycles of this pop-
ulation and that of the breakwater is the lack of
an indeterminate period during the summer. Of
the 416 examined, only five were indeterminate
because of a complete spawning. Eight others
were found to be indeterminate due to heavy in-
festations of trematode sporocysts and cerca-
riae. The analysis of this population was more
difficult because of its tendency to spawn dur-
ing storage. This fact itself indicates the in-
creased reproductive potential of this popula-
tion as compared with that of the less favorable
breakwater. These spontaneous spawnings took
place throughout the year and resulted in nor-
mal trochophore larvae. The best indication of
what was taking place in the population in its
natural habitat was gained by recording the in-
tensity of the laboratory spawnings and await-
ing a sample which did not spawn and which,
upon examination, showed itself to consist of
recently spawned animals. Plate 13 indicates
the condition of the sample when examined and
also the extent of the laboratory spawning. By
correlating these two pieces of information, the
times of natural spawning may be approximated.
An example of the method used is seen in rela-
tion to the collections of April 1 and April 15,
1951, The first of these spawned heavily as is
indicated by the designation +3. This sample
is believed to have been ripe when collected.
The other sample showed only one ripe animal,
the entire group being either completely or
partially spawned. It is thus presumed that the
population spawned between these dates. On the
basis of this type of analysis, it is believed that
the population spawned three times during 1951.
This is considered, however, to be a conserva-
tive estimate. When the spawning reaction is
as easily provoked as it is in this population, it
is not at all improbable that minor, undetected
spawnings may occur frequently and serve to
augment the major spawnings.
The study of this population dramatically
illustrates the effect which extrinsic environ-
mental factors can have upon the reproductive
capacity of these limpets. Both the Moss Beach
and the Rockaway Breakwater populations are
unquestionably Acmaea digitalis and in both
habitats the limpets are abundant. The differ-
ence wrought in these animals by the envi-
ronment is alteration ad the reproductive potential
Page 46
THE VELIGER
Vol. 4; No. 1
Analysis of Environmental Conditions Coincident
with Spawning:
Of the four spawnings recorded for this popula-
tion of Acmaea digitalis, two coincide with
spawnings of the breakwater group, these being
in April, 1951 and 1952. The other two spawn-
ings occurred in June to July and in December
when the breakwater population was either in-
determinate or in the process of gonad rede-
velopment. Two of the periods were in the
presence of full moons (June 1951 and March to
April 1952) and two in new moons (April 1951,
and June to July 1951). The tidal ranges were
5.8, 6.5, 6.6, and 8.8 feet for the spawning pe-
riods. Sea temperatures ranged from 52.2°F.
to 57.4°F., the latter temperature being 2.4°F.
higher than the highest temperature at which
the breakwater group spawned.
dig
Acmaea scabra (GOULD, 1846)
Ecology: — This species was studied from the
horizontal surfaces of the Rockaway Breakwa -
ter. The top of this structure is about four feet
wide and tends to retain water to a greater ex
tent than that of the vertical surfaces from
which Acmaea digitalis was collected. There
is an increased concentration of barnacles,
Balanus glandula, and moderate numbers of
Mytilus californianus Conrad, 1837, along the
seaward edge. Plant life is sparse, but some
Endocladia is present. Although the radiation
on this horizontal surface is probably more in-
tense and of longer duration than on the verti-
cal surfaces, the retention of water on the top
reduces the degree of desiccation experienced.
It appears that each A. scabra has a home scar
on the rock from which it goes forth to feed and
to which it returns when the tide recedes. Such
a depression in the concrete allows the limpet
to retain water in its shell and mantle cavities
more effectively than A, digitalis which has no
home scar and usually retreats to a shaded
crevice when the tide is out.
Collections: — These were begun on September
25, 1949, and continued at approximately two
week intervals until March 31, 1952. A total of
822 animals was examined: 414 females, 373
males, 33 indeterminate, and 2 immature. Al-
though this is a sizeable number of specimens
to be taken from a rather restricted area, and
although the species managed to maintain its
numbers despite the heavy collections, this
habitat is not considered to be an optimal one
for this animal. At no time, even at the time
that collections were begun, was the species
very numerous and some search was always
necessary to obtain the desired number. In ad-
dition, the animals seldom attained the degree
of gonad development which is associated with
a condition of ripeness, or at least the degree
found in animals from a lower position in the
intertidal zone. This tendency for only moder-
ate gonad development interfered, to an extent,
with a clear-cut analysis of the reproductive
cycle of this population. For this reason, the
trend of gonad redevelopment was misinter-
preted for the first year's study, and conse -
quently no ripe animals are plotted for this pe-
riod in Plate 14. Subsequent to this, only the
categories of spawned and ripe have been plot-
ted, with percentages of indeterminate when-
ever this condition occurred. During the peri-
od of April through August, 1950, collections
were too infrequent and specimens too few to
be considered reliable and thus this section of
the graph is incomplete.
Results; — On the basis of data obtained from
this study, it seems probable (Plate 14) that Ac -
maea scabra, in this habitat, spawns three
times per year, in the late winter or early
spring, in the early summer, and in late fall.
There is no extensive period of indeterminacy
in the summer as is seen for A.digitalis. This
difference may be partially related tothe some-
what more favorable habitat inhabited by A.
scabra as described above, but, as will be dis-
cussed later, probably also is correlated with
the greater tolerance of A. scabra to high tem-
peratures. That A. scabra will become indeter-
minate in the summer months when it inhabits
a very high intertidal position is shown by a
brief examination of a population living on the
high granite rocks near the Hopkins Marine
Station, Pacific Grove, California. This study
was made in July, 1949, and showed that the
limpets were all indeterminate, while those
taken from zone 2 amidst the Ulva were all re-
productively active. Here, then, is another ex-
ample of the effect of the environment on two
ecologically different populations of a single
eurytopic species.
Analysis of Environmental Conditions Coincident
with Spawning:
Of the six spawnings recorded for Acmaea
scabra, two coincide exactly with spawnings of
A. digitalis: January, 1951, and February, 1952
A third in mid-February, 1950, very nearly co-
incides with A. digitalis in the early part of the
month. During the two periods of coincidental
spawnings, it seems not unlikely that the two
species were responding toa similar stimulus
and, considering the fact, as will be shown la-
ter, that the two limpets evidently differ in
THE Venicrr, Vol. 4, No. 1
1950
[FRivcHMan]
D
pee tr dh daft
Aug. Sep. Oct.
Nov.
Dec.
May
Acmaea asmi (Mipvenporrr, 1847)
Moss Beach
% of sample ripe
----- % of sample spawned
O & of sample of indeterminate sex
Vv spawning period
¥v laboratory spawning
Plate 11
Tue VELIGER, Vol. 4, No. 1 [FRITCHMAN] Plate 12
100 we a
10)
80
Acmaea digitalis EscuscHoitz, 1833
Rockaway Breakwater
% of sample ripe
% of sample of determinate sex
spawning period
laboratory spawning
=
Tue Ve.icer, Vol. 4, No. 1 [FRITCHMAN] Plate 13
Jan.
1952
Acmaea digitalis EscuscHoLtz, 1833
Moss Beach
+ % of sample ripe
© ------ % of sample spawned
Vv spawning period
¥v laboratory spawning
THe Vevicer, Vol. 4, No. 1 [FRitcHman] Plate 14
+
O
a)
O---Q----O---- O---O---©---O---O---O--
Aug. Sep. Oct. Nov. Dec.
Jan.
1951
Acmaea scabra (Goutp, 1846)
Rockaway Breakwater
+ %, of sample ripe
©----- & of sample spawned
oO % of sample of indeterminate sex
Vv spawning period
v laboratory spawning
Vol. 4; No. 1
THE VELIGER
Page 47
their temperature requirements, this stimulus
must probably be related to tidal action or lu-
nar cycle. However, these influences cannot be
accurately defined because of the appearance of
both new and full moon phases during these pe-
riods. The maximum tidal ranges during which
A. scabra spawned were 6. 6 feet (June to July,
1951), 7.3 feet (February, 1950), and 8.7 feet
(July, 1951). As with A. digitalis, these are
quite inconclusive. The sea water tempera-
tures at which spawnings occurred vary from a
near minimum for this latitude (e.g., 50.0°F.
in February, 1950) to 57.5°F. during June and
July, 1951. This range is appreciably more
than that found for the A. digitalis population on
the breakwater.
Literature Cited
Fritchman, Harry K., 1.
1961. is
gradually added. The relaxed specimens are
then placed in Gilson's fixative for about six
hours. Dorididae can be narcotized in MgCl?
and killed with boiling Susa's fixative. Aplysi-
idae may be narcotized with chloral hydrate (a
teaspoon to a medium-sized beaker of water)
and placed in liquid of Perényi to harden.
When either Susa's or Gilson's fixatives are
used, all traces of the corrosive sublimate
must be removed from the opisthobranchs before
they are stored in alcohol. This can be accom-
plished by adding tincture of iodine to 70 per-
cent alcohol containing the specimens. Enough
iodine should be added to color the alcohol a
port wine shade. The specimens should be
placed in changes of fresh alcohol until all
traces of the iodine are removed. Do not use
metal instruments with either of the above men-
tioned fixatives; use a paint brush mounted in
plastic, or a pipette. For permanent storage 70
percent alcohol should be used. Glycerine can
be added in regions of hot weather or when the
specimens are to remain packed for some time.
This will prevent complete desiccation and de-
composition of the specimen in case of evapo-
ration.
Corrosive sublimate (mercuric chloride, bi-
chloride of mercury), as referred to in this
article, is a saturated solution in distilled wa-
ter (about a 7 percent solution). Both Susa's
and Gilson's fixatives must be filtered after
three days. A list of formulas for the fixatives
used in this article is given below.
Susa's Fixative:
HgCl2 45.0 gm.
NaCl 5.0 gm.
Distilled water 800 cc.
Trichloracetic acid 20 gm.
Acetic acid (glacial) 40 cc.
Formalin (40 percent) 200 cc.
Gilson's mercuro-nitric mixture;
Corrosive sublimate 5.0 gm.
Nitric acid 4 cc.
Glacial acetic acid N ee.
Alcohol (70 percent) (313) (Xp
Distilled water 200 cc.
Liquid of Perényi:
Nitric acid (10 percent) 4 parts
Alcohol (70 percent) 3 parts
Chromic acid (0.5 percent) 3 parts
Vol. 4; No. 1
THE VELIGER
Page 53
Books, Periodicals, Pamphlets
A PROPOSED RECLASSIFICATION
OF THE GASTROPOD FAMILY
VERMETIDAE
by A. Myra Keen
Bull. British Museum (Nat. Hist.)
Zoology, vol. 7, No. 3, pp. 183-214
pl. 54, 33 text figs. London, Feb., 1961
It is most fortunate that the very difficult
family at last has found such masterly treat-
ment; many of the obscure and puzzling points
are now cleared up. The student of marine in-
tertidal ecology as well as the shell collector
will be greatly benefited by this study.
However, we cannot pass by lightly a cir-
cumstance which we must deplore most serious -
ly. For reasons known probably only unto the
Trustees of the British Museum, this paper is
copyrighted. While generally speaking a copy-
right may be a desirable thing to protect the
rights of the creator of a literary work, it
seems completely out of place in scientific
writings. We readily grant that copyrights are
justified in certain scientific works, such as
keys and many of the compilative faunal sur-
veys. However, a ticklish question is raised by
the procedure of the Trustees of the British
Museum. In this paper Dr. Keen describes a
new genus. Since we have not had time to cor-
respond with the Trustees to obtain permission
to quote from the paper for the purposes of this
review, we are not able to list thenew genus!
It will also mean that every future worker who
has occasion to use the paper — and there will
be many of these — will have to obtain permis-
sion from the Trustees to quote the new genus
name. We wonder if the International Commis-
Sion on Nomenclature will have to rule that the
name in question is not available? And, if this
copyright will stand, does it mean that all future
authors should be "'protected'' in a similar fa-
shion? We especially deplore this occurrence
because it happened to Dr. Keen, a most gen-
erous scientific worker who would never have
consented to this, we are convinced, had she
had a voice in the matter. This brings to mind
that not long ago in a periodical, a copyright
had been obtained but when the problems con-
nected therewith were pointed out to the author
concerned, the copyright was rescinded
promptly and retroactively. Let us hope that
the Trustees of the British Museum will follow
an equally sensible course.
RS
PELECYPODS FROM TRISTAN DA CUNHA
by T. Soot-Ryen
Results Norweg. Scientif. Exped. to
Tristan da Cunha 1937-1938, No. 49.
47 pp., 3pls., and 9 text figs. Oslo, 1960.
This is another of the thorough papers by
Dr. Soot-Ryen. In it he describes the following
new species: Philobrya sivertseni, Ph. insu-
laris, Verticipronus tristanensis, Cyclopecten
(Cyclochlamys) perplexus, Notolepton atlanti-
cum, and Rochefortula variabilis.
RS
WEST AMERICAN SPECIES
OF THE BIVALVED GASTROPOD GENUS
BERTHELINIA
by A. Myra Keen and Allyn G. Smith
Proc. Calif. Acad. Sciences, 4th Ser.,
vol. 303) no, 25 pp. 47-66, 33 figs, 1
color plate. March 20, 1961.
Ever since the exciting discovery that a
heretofore ''typical'' clam was really a snail,
much interest has been centered on this group
of small gastropods. In the present paper an
account is given of the search made for repre-
sentatives of this group in the southern Gulf of
California, and of the success of the search.
' When one regards the colored plate, one begins
to understand why this amazing form has been
overlooked for so long by even very experienced
collectors.
RS
PECTINIDES DU NEOGENE
DE LA HONGARIE
ET LEUR IMPORTANCE
BIOSTRATIGRAPHIQUE
by I. Csepreghy-Meznerics
Mémoires de la Société Géologique de
France, Nouvelle Série, Tome XXXIX,
Feuilles 15 4 18, planches XXXVIII 4
LXXII, Mémoire No. 92, pp. 1 4 58,
planches I 4 XXXV, 1960.
The scope and presentation of this mono-
graph, dealing with the Pectinidae of the late
Tertiary of Hungary, is in general similar to
the fine European monographs of this family of
pelecypods by Deperet and Roman, Roger, and
Ugolini. Seventy-one species and subspecies
are described, all except one are illustrated;
their relationship to other species in the Medi-
terranean basin is discussed as well as their
geologic and geographic range and facies oc-
currences, Conclusions concerning the bio-
stratigraphic and paleogeographic significance
of the species are included.
Page 54
THE: VELIGER
Vol. 4; No. 1
The species and subspecies are placed in
four genera and one subgenus: Pecten (10),
Flabellipecten (8), Amussium (3), Chlamys (48),
subgenus Camptonectes (2). The following are
described as new: Pecten fotensis, n.s., P.
promontorensis, n.s., Flabellipecten telegdi-
rothi, n.s., Chlamys scabrella hungarica,
n. subsp., C. biaense, n.s., C. rakosense, n.s.,
C. darnoensis, n.s., C. agriensis, n.s., C.
palmata bipartita, n. subsp.
This paper, beautifully illustrated, is a fine
addition to the paleontological literature of the
Mediterranean basin. LGH
SEA SHELLS
OF TROPICAL WEST AMERICA
by A. Myra Keen
Second Printing. xi + 624 pages, over
1700 illustrations. Stanford University
Press. $12.50.
In keeping with the modest approach of
Stanford University Press, this new edition of
the very valuable work by Dr. Keen is calleda
"second printing" yet there are quite a few
changes incorporated, as well as all the com-
ments and corrections of errata which were
listed separately on the ''Errata'' and ''Supple-
mentary Errata''’. There are several new fig-
ures and additional figures, as well as a few
name changes.
This ''second printing'' is again a must for
the student of the mollusks of the west coast, as
well as for the amateur shell collector, be he a
beginner or an advanced connoisseur. How im-
portant the contribution of Dr. Keen is through
the first edition (pardon us, the first Printing)
of this splendid work may be assessed through
the number of papers that have been appearing
since the book was published; the research in
the area covered by the book has certainly been
stimulated and the result of these investigations
will, in due time, necessitate another ''Print-
ing" with further additions. In many collections
that we have seen in the past number of months,
we find labels which bear the ''K number"! of the
shell species, i.e., the number preceding the
name in Dr. Keen's book.
It would appear that ''Keen number''! may
come to mean to shell collectors what ''Scott
number'' means to the stamp collectors, a
quick, simple, reliable way of communication
among the devotees.
Perhaps we should have mentioned that the
"second printing'' is on a whiter paper than the
previous edition, which, to us at least, is more
agreeable.
RS
REVISION OF
TORNATELLINIDAE AND ACHATINELLIDAE
(GASTROPODA, PULMONATA)
by C. Montague Cooke, Jr., and Yoshio Kondo
Bull. 221, Bernice P. Bishop Museum.
December 30, 1960.
This comprehensive work was received too
late for an adequate review. We expect to re-
port on it in our next issue.
Ed.
CARIBBEAN SEASHELLS
A GUIDE TO THE MARINE MOLLUSKS
OF PUERTO RICO AND OTHER WEST
INDIAN ISLANDS, BERMUDA AND
THE LOWER FLORIDA KEYS
by Germaine L. Warmke
Curator of Mollusks
Institute of Marine Biology
University of Puerto Rico
and R. Tucker Abbott
Academy of Natural Sciences
of Philadelphia
This eagerly awaited volume describes
800 species of shells, most of which
are not found in any other book on sea-
shells. The superb photographs on 44
plates (some in full color) and the many
line drawings make this an indispens-
able reference book for serious students
and hobbyists alike. The unique feature
of this new book is an extensive guide
to shelling areas of the Caribbean.
Each major island, from Bermuda to
Trinidad and from Cozumel to St,
Croix, is conchologically described,
with travel hints, when and where to
collect, and how best to clean and ship
home specimens.
($8.95 postpaid)
JQB
PRELIMINARY STUDIES ON THE VENOM
OF THE MARINE SNAIL CONUS
by A.J. Kohn, P. R. Saunders, and S,. Wiener
Ann. New York Acad. Sci., vol. 90,
article 3, pp. 706-725, Nov. 17, 1960,
A well documented study of the venom and
its effect upon a variety of other animals. Cone
species studied were Conus striatus, C, tex-
tile, C. aulicus, and C, marmoreus.
RS
THE VELIGER is open to original papers pertaining to any problem
concerned with mollusks from the Pacific Region.
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EDITORIAL BOARD
Dr. Donatp P. Asport, Associate Professor of Biology
Hopkins Marine Station of Stanford University
Dr. J. Wyatr Duruam, Professor of Paleontology
University of California, Berkeley
Dr. E. W. Facer, Associate Professor of Biology
Scripps Institution of Oceanography, University of
California, La Jolla
Dr. Cavet HAnp, Associate Professor of Zoology
University of California, Berkeley
Dr. G. DAtitas Hanna, Curator, Department of Geology
California Academy of Sciences, San Francisco
Dr. LEo G. HERTLEIN, Associate Curator, Department of Geology
California Academy of Sciences, San Francisco
Dr. Myra Keen, Associate Professor of Paleontology and
Curator of Concholog y
Stanford University
Dr. Frank A. PitEvka, Professor of Zoology
University of California, Berkeley
Mr. Atiyn G. Smit, Research Malacologist
California Academy of Sciences, San Francisco
Dr. Ravpw I. Situ, Professor of Zoology
University of California, Berkeley
EDITroR
Dr. Rupoir STOHLER, Associate Research Zoologist
University of California, Berkeley
A Quarterly published by
NORTHERN CALIFORNIA MALACOZOOLOGICAL CLUB
Berkeley, California
VOLUME 4 OcTOBER I, 1961 NUMBER 2
CONTENTS
A New Doridid Nudibranch from ae Victoria (Plate 1 5
RoBERT BuRN rs ; SRO CNet mie eis
Notes on the Onna of the West Guan of North pees
I. Nomenclatural re in the Order Nudibranchia (Southern California)
Joan E. STEINBERG . . Sane WORM ot ue oe Wenig!
A Distributional List of Southern California Oni ebraneh:
James R. Lance. . siti cer ca hey sired: ae ee?
A New Commensal polvelad fon ee (Plate 16)
Epmunp H. SmirH oe 6 : SOA ASS ery glen = Tesi BES 7SEAC SLO)
A New Subgenus and Species of Coral- ue pane
from the Gulf of California (Plate 17, 2 seer
Victor A. ZULLO eee. BORIC ; Pe Sano ge Ne SN Os cea
A Discussion of Vexzllum regina Cone 182 5 and rece Species,
With Description of a New Subspecies (Plates 18, 19, 20; 1 Textfigure )
Jai IME, MCRAE Nt ESI Seibel ur Maer Cumin mes te ie gre ae mie may)
Observations on the Biology of Hermaeina smithi, a Sacoglossan Opisthobranch
from the West Coast of North America (13 Textfigures)
J. J. Gonor st Mine times buon 6 . 5 6 OS
On Certain Littoral Species of Relient (Cirripedia, TRB) Senioue with
Decapod Crustacea from Australia, Hawaii, and Japan (Plates 21, 22, 23)
WitiramM A. NEWMAN. .. ae ran CVS HREM MES BERS ER EVE Rohn n, ay tmenn Mea ee A fxC0\0)
Another Statistical Study in Size of C Cowries
F. A. SCHILDER Sei se sneer UM pen aiae! aN ivae UO
Remarks on a Variation in Gyrus annettae Daa, 1909 (Plate 24)
CRAWFORD N. CaTE ee en aid Mee ea ers, AAS Ook hon wea ee eS
[Continued on Inside Front Cover ]
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Send subscriptions to: Mrs. PHOEBE Batcu, Treasurer, 1150 Brown Avenue, Lafayette,
California. Address all other correspondence to: Dr. R. STOHLER, Editor, Department of
Zoology, University of California, Berkeley 4, California.
Application to Mail at Second Class Postage Rates is pending at Berkeley, California
CONTENTS
[ CONTINUED |
NOTES & NEWS |. icici ee ee Er ee ee
Range Extension of Anatina cyprinus (Woop, 1828) BrucE CAMPBELL
About Copyright
BOOKS, PERIODICALS (& PAMIPEIE BAS oye yce =) eee) cnn eet
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, Sufamily Genus, Subgenus.
Vol. 4; No. 2
THE VELIGER
Page 55
A New Doridid Nudibranch from Torquay, Victoria
by
ROBERT BURN
34 Autumn Street, Geelong West, Victoria, Australia
(Plate 15)
The genus Glossodoris Ehrenberg, 1831, is
already represented in Victorian waters by five
species (Burn, 1957, pp. 16-18). The new spe-
cies described below differs in colouring from
all of these species with one exception but can
be separated from that species, G. haliclona
Burn (loc. cit., p.17), by the very different
shape of its radular teeth.
According to Odhner (1957) the generic
name Glossodoris should not be used, as the
type species G, xantholeuca Ehrenberg, 1831,
has never been anatomically examined. In
place of Glossodoris Odhner proposed to use
two names, based upon the rather simple divi-
sion of the species assigned to the genus ac-
cording to the shape of the radular teeth. For
those species with the teeth hamate and denti-
culate, he proposed to use Chromodoris Alder
and Hancock, 1855; and for those species which
have tri- and bi-cuspid teeth he proposed to use
the genus Hypselodoris Stimpson, 1855. How-
ever, until the question of the anatomy and ra-
dula are satisfactorily known in the type of
Glossodoris with the probability of one of Odh-
ner's suggested genera becoming a synonym, I
prefer to use the older generic name.
Glossodoris arbuta BURN, spec. nov.
(Plate 15, Figures 1 and 2)
The single specimen (holotype) measured
alive 22 mm. long and 5 mm. broad. Along ei-
ther lateral margin of the mantle there are four
or five deep indentations, each corresponding
to a colour patch on the dorsal surface. The
mantle is very little wider than the foot. The
foot is grooved for its full length, anteriorly
the lateral corners are expanded and form
sharp corners, posteriorly the tail is narrow
and rounded. The oral tentacles are short and
digitiform. The rhinophores have 12 lamellae.
The branchiae number 12 simple plumes, they
encircle the white-rimmed anus.
The radular formula is 24x 40.0.40. The
teeth are all simply hamate, the cusp of each is
unequally bifid. Except for the marginals each
tooth has a longer basal plate than cusp.
The body colour is bright pink of a nearly
strawberry hue, the rhinophores and branchiae
are transparent pale red. The foot is outlined
with a row of obscure white dots. About each
indentation of the mantle margin is a large
rounded patch of yellow; in the midst of these
patches is a small number of minute bright
red flecks.
Type locality: Point Danger, Torquay, Vic-
toria. Long. 144°19' East, lat. 38°20' South. (1
specimen, 30 March, 1959, collected by the
writer.) It was found crawling on brown sea-
weed at extreme low tide level.
The specific name arbutus has been chosen
in allusion to the wild strawberry tree or Ar-
butus tree, the colour of which is present on the
new species.
Remarks: Three or four pink coloured
species of Glossodoris are recorded from the
Australasian region, and the new one must be
compared with each of them. The Victorian G.
haliclona Burn, 1957 (p. 17) does not have the
marginal yellow patches on the mantle, the ra-
dula has half as many rows of teeth, and the
teeth are denticulate instead of bifid as in the
present species. The South Australian G. epi-
curia (Basedow and Hedley, 1905 [p. 153, pl. 7,
figs. 1-3]), is larger, has five branchiae as
compared with 12 in G. arbuta, the rhinophores
carry more lamellae, the foot corners and oral
tentacles extend beyond the anterior mantle,
and the radular formula is smaller according
to the original description while the individual
teeth are denticulate. It is probable that the
Page 56
Glossodoris sp. of Allan, 1947 (p. 445) is synon-
ymous with Chromodoris (= Glossodoris) aus-
tralis Risbec, 1928 (p. 143, pl.7, fig.8), but as
the description of the former is inadequate and
lacks any mention of the radula, the two must
be maintained as separate species. As both
species have darker red spotting on the mantle
and the latter has denticulate radular teeth,
neither can be identified with G. arbuta.
According to the arguments of Odhner
(1957, p.252), Glossodoris arbuta would be
classified as a Hypselodoris.
The holotype has been presented to the Na-
tional Museum of Victoria, Melbourne, where
it is registered as F 21,272.
The radulae of the three following Victori-
an species of Glossodoris have been examined
for comparison with G. arbuta. As none of
these have previously been fully ‘described or
figured from Victorian material, the discrep-
ancies are here made good.
Glossodoris victoriae Burn, 1957 (p.16)
Plate 15, figure 3. The radular formula is
36 x 37.0.37. The first lateral is narrow and
denticulate on each side of the cusp, one denti-
cle is on the median side and four denticles on
the marginal side. The subsequent teeth have
five denticles on the marginal side. This spe-
cies is referable to Chromodoris according to
the arguments of Odhner (1957, p. 252).
Glossodoris tasmaniensis (Bergh, 1905),
(Burn, 1957, p. 17) Plate 15, figure 4. The ra-
dular formula is 48 x 40.0.40. The first later-
al is very broad, the cusp has two denticles on
the median side and three on the marginal side.
The subsequent teeth each have three denticles
on the marginal side, the marginal teeth have
one or two denticles whilst the outermost tooth
appears to be bifid at the tip. The exceptional-
ly broad first lateral tooth is a little separated
from its neighbour, and for these conditions,
i.e., broad lateral tooth and separation from
other teeth in half row of radula, the genus
Noumea Risbec (1928, p. 165) was proposed. In
every other way the teeth are typical of Chrom-
THE VELIGER
Vol. 4; No. 2
odoris (Odhner, 1957, p. 252). Pruvot-Fol (1951,
p. 147) gives the radular formula of G. tasma-
niensis as 50 x 150 (= 50 x 75.0.75), which is
considerably more teeth per half row than in
the present material.
Glossodoris haliclona Burn, 1957 (p.17)
Plate 15, figure 5. The radular formula is
12 x 30.0.30. The first lateral is denticulate on
both sides, the median side has one denticle and
the marginal side, five. The subsequent teeth
each have five denticles, that nearest the tip of
the cusp is largest and is more like a bifurca-
tion of the cusp than a denticle. The marginal
teeth are small and dumpy, each with five den-
ticles. The elements of the labial disk are
shallowly curved with one end bifid. This ra-
dula approaches closest to G. hilaris (Bergh)
as figured by Baba (1953, p.210, fig. 6J), al-
though the labial elements are very dissimilar.
As with the previous two species, this one is
referable to Chromodoris (Odhner, 1957, p. 252).
Literature Cited
Allan, J.
1947.
North Coast, New South Wales.
21 (8): 443-463, pls. 41-43,
Baba, Kikutar8
1953.
genus Glossodoris from Japan.
Biol. Lab., 3 (2): 205-211.
Basedow, H., & C. Hedley
1905. South Australian nudibranchs, and an enumera-
tion of the known Australian species. Trans. Roy.
Soc. So. Aust., 29: 135-160, pls. 1-3.
Burn, Robert
1957. On some opisthobranchia from Victoria. J,
Malac. Soc. Aust., 1: 11-26, pls. 1-3.
Odhner, Nils
1957.
nomenclaturial controversy.
London 32 (6): 250-253.
Pruvot-Fol, Alice
1951, Révision du genre Glossodoris Ehrenberg.
Jour. Conchyliol. 91: 76-164.
Risbec, Jean
1928. Contribution a l'étude des nudibranches Néo-
Calédoniens. Faun. Colon. Frang 2 (1): 1-328,
pls. 1-16.
Publ. Seto Mar.
Proc. Malacol. Soc.
Explanation of Plate 15
Figure 1: Glossodoris arbuta Burn, spec. nov. Dorsal view, x 4%,
Figure 2: Two radular teeth of the same
[a - an inner lateral, b - a marginal tooth] Figure 3: Glossodoris victoriae BuRN. A half row of radular teeth
[a - inner lateral, b - side view of a lateral tooth, c - near marginal tooth, d - marginal tooth] Figure 4: Glos-
sodorts tasmaniensis (BERGH). A half row of radular teeth [a - inner lateral, b - side view of a lateral, c - near
marginal, d - marginal tooth] Figure 5:
Glossodoris haliclona Burn. A half row of radular teeth and a labial
element [a - labial element, b - side view of inner lateral, c - side view of lateral,
d - side view of marginal tooth]
Nudibranchia from the Clarence River Heads,
Rec. Aust. Mus.,
Three new species and two new records of the
Chromodoris contra Glossodoris, a systematic-
at
B Plate 1
Tue VELIGER, Vol. 4, No. 2 [Burn ] Plate 15
Figure 2
ee AA /
Figure 1 Figure 3
a
Figure 4
Burn, del,
Vol. 4; No. 2
THE VELIGER
Page 57
Notes on the Opisthobranchs
of the West Coast of North America
I. Nomenclatural Changes in the Order Nudibranchia
(Southern California)
Joan E. STEINBERG
850 38th Avenue, San Francisco 21, California
{Editor's Note: In the work preliminary to the preparation
of the key to the opisthobranch mollusks of the west coast of
North America, Miss Joan Steinberg has encountereda num-
ber of problems which she considers, quite rightly, should
be dealt with before the key is completed. The present ar-
ticle is concerned with the first few of these; two other pa-
pers in this issue may be regarded as also contributing to
the solution of some of these problems. - Since there are
several serious gaps in our knowledge of this interesting
group, gaps concerning assignment of certain species toa
particular genus, as well as some others, we think that Miss
Steinberg's approach is correct and that our readers will be
glad to wait a little longer than originally anticipated for the
completion of the keys which will, it is hoped, stimulate in-
tensive further research in this group. }
The recent paper by Marcus (1961) on Opis-
thobranch Mollusks from California has con-
tributed greatly to our knowledge of this group
on the West Coast. However, it is evident that
much more work is necessary, especially in
those areas where little, if any, systematic
collecting has been done, before we will really
begin to appreciate the wealth of opisthobranchs
in our fauna. For example, the only truly com-
prehensive list of nudibranchs from the West
Coast of North America was prepared by
O'Donoghue in 1926. It includes references and
synonyms for all species known up to that
time. However, O'Donoghue's work was com-
piled largely from the literature, and it has be-
come apparent that a re-examination of a num-
ber of species is necessary in order to deter-
mine their validity or their exact systematic
positions.
Earlier this year I had several opportuni-
ties to discuss systematic problems in the Or-
der Nudibranchia with Mr. James R. Lance
(Scripps Institution of Oceanography, La Jolla,
California). Of primary concern to us were
certain species recorded from Southern Cali-
fornia, the systematic status of which was open
to question. This paper is the result of our
mutual and separate investigations. Mr. Lance
has incorporated the nomenclatural changes
proposed herein in his forthcoming list of the
Opisthobranchs from Southern California. {Ed.
Note: See the following paper by J. Lance.}
Iam deeply grateful to Mr. Lance for his
wholehearted cooperation with my attempts to
achieve some standardization of nomenclature
for our West Coast opisthobranch fauna.
Primarily, only nudibranchs occurring in
Southern California were considered for this
paper, although the ranges of many of these ex-
tend far northward. Therefore, in certain in-
stances, it has been necessary to examine and
compare specimens and descriptions of animals
from Washington and Vancouver Island, as well
as from Northern California, in order to clari-
fy the issues at hand. I would like to thank the
staff and students at both the Friday Harbor
Laboratories (University of Washington) and
the Hopkins Marine Station (Stanford Universi-
ty) for their assistance. Dr. Leo Hertlein (Ge-
ology Department) and the library staff of the
California Academy of Sciences were most
helpful to me in locating literature.
In 1926, O'Donoghue listed three species of
dorid nudibranchs belonging to the Family
Dendrodorididae from the West Coast of North
America:
Dendrodoris fulva (MacFarland, 1905)
Dendrodoris vidua (Bergh, 1878)
Doriopsilla albopunctata (Cooper, 1863)
Page 58
THE VELIGER
Vol. 4; No. 2
Doriopsilla albopunctata (Cooper, 1863) was
first described as Doris (7?) albopunctata from
specimens obtained in deep water near Santa
Barbara and from rocks at low water on Santa
Catalina Island. In 1870, Cooper published ad-
ditional distributional information on his spe-
cies, stating that its range was from Baulines
(sic., Bolinas) Bay to San Diego and that it was
rare under stones at low water at Santa Cruz.
Orcutt (1885) and Yates (1890), in faunal lists
of mollusks from San Diego and Santa Barbara,
respectively, also record Doris albopunctata
but their records are based purely on Cooper's
works. Orcutt has been the only author to uti-
lize the information in Cooper's later paper,
and it has been overlooked by subsequent work-
ers.
In 1905, Cockerell and Eliot described a
new species, Doridopsis reticulata, from San
Pedro, which they suggested was probably iden-
tical with Cooper's form. O'Donoghue (1922a)
later synonymized Cockerell and Eliot's spe-
cies with Cooper's and, following Eliot's (1906)
subsequent work on Cockerell's material, as-
signed it to the genus Doriopsilla.
However, also in 1905, MacFarland de-
scribed a dendrodorid from Monterey Bay which
he called Doriopsis fulva and which was later
(O'Donoghue, 1926) referred to the genus Den-
drodoris Ehrenberg, 1831. MacFarland, in his
original description stated: "It is very possible
that this species is identical with the Doris al-
bopunctata of Cooper ...''. He concludes, how-
ever, that, aside from similarity in coloration,
the brief description contained points which
were at variance with his material and that, in
the absence of Cooper's type specimens, iden-
tification was very difficult.
One point which led Mr. Lance and me to
consider the possibility that perhaps we were
dealing with two separate species, Dendrodoris
fulva in Central California and Doriopsilla al-
bopunctata in Southern California, was that the
animals from the Monterey area are nearly al-
ways bright yellow, whereas the animals in
the San Diego region vary from yellow in the
very small animals to a warm brown, tending
to yellow near the edge of the notum in the
larger individuals. In the latter the white
glands which both MacFarland and Cooper men-
tion are quite conspicuous, especially against
the darker background, whereas they must
sometimes be looked for carefully in living
animals from Monterey.
In July, 1961, 1 collected a nurnber of spe-
cimens from Mission Point and Point Pinos on
the Monterey Peninsula which matched Mac-
Farland's description and color plate (1906) al-
most exactly. In addition, I also collected the
darkest forms I could find. One specimen from
Mission Point was quite orange in general ap-
pearance but, under a dissecting microscope,
proved to have the white glands typical of Mac-
Farland's species, Furthermore, it bore a
great many brown dots dorsally which gave it a
dusky orange color. Examination of more typi-
cal specimens showed that there is a great
variation in the presence of these brown dots.
Several of the specimens at either extreme of
the color range for the Monterey forms (from
bright yellow to dusky yellow) and also varying
in size were taken alive by air to LaJolla. The
next morning a large number of the larger-
darker and smaller-lighter forms was collected
at Point Loma. After careful examination of
the coloration of animals from both localities,
it was concluded that, although the southern
forms may get much darker dorsally as the
animal grows larger, we are only dealing with
a color variation which varies with latitude.
Several other nudibranchs on this coast exhibit
a similar darkening in color depending on the
latitude. [A good case in point is Diaulula
sandiegensis (Cooper, 1862). Animals in the
Vancouver Island and San Juan Island regions
are much darker than animals from San Diego
or Monterey, although darker specimens may
occasionally be collected in Central Califor-
nia. }
The yellow color of small specimens from
both localities does not differ at all. As far as
could be determined by dissection, the repro-
ductive systems of both fitted MacFarland's
(1906) description, and the central nervous sys-
tems, as well as the anterior parts of the di-
gestive systems, were identical with that de-
_ scribed by Eliot (1906) for Doriopsilla reticu-
lata. It is my opinion that the two are identical
and that Cooper's specific name takes prece-
dence over MacFarland's.
Having established the synonymy of the two
forms, the question arises as to which generic
name should be used. Dendrodoris is distin-
guished from Doriopsilla by having the buccal
ganglia situated some distance from the central
nerve ring in a bend in the esophagus but joined
to the central ganglia by a pair of long commis-
sures, In Doriopsilla, the buccal ganglia are
located immediately behind the central nerve
ring, the two ganglia being joined by a short
commissure, Although, as Eliot points out, it
Vol. 4; No. 2
THE VELIGER
Page 59
is difficult to determine the position of the buc-
cal ganglia in relation to the rest of the nerve
ring (except by sectioning), it was possible in
the specimens I examined to determine that the
position of the buccal ganglia was not as it is in
Dendrodoris. The '"strands' referred to by
Eliot were first thought to be the long commis-
sures to the buccal ganglia, but a more careful
examination revealed their true nature.
Pruvot-Fol (1954), on the basis of the con-
dition of the central nervous system, retains
Doriopsilla as a distinct genus; however, Baba
apparently considers it to be a subgenus of Den-
drodoris (e.g., see Baba, 1949), Considering
the fact that the central nervous systems of
most of the numerous Dendrodorids which have
been thus far described have not been investi-
gated, I think that, until a thorough revision of
the family has been undertaken on a worldwide
basis, it is best to retain our species in the
genus Dendrodoris belonging to the subgenus
Doriopsilla. Until it can be shown that the con-
dition of the central nervous system may be
successfully used in separating genera in this
very difficult family, I do not consider it wise
to maintain Doriopsilla as generically distinct
from Dendrodoris.
The precedence of Dendrodoris over other
names has been thoroughly discussed (O'Dono-
ghue, 1926, and Pruvot-Fol, 1954) and will not
be repeated here.
I cannot agree with Eliot (1907) that Mac-
Farland's Dendrodoris fulva (now D. albopunc-
tata) is synonymous with Dendrodoris citrina
(Cheeseman, 1880) from New Zealand. Apart
from the great geographical separation of the
two species, the two differ morphologically.
Eliot describes the notum of D. citrina as
"covered with numerous well-developed tuber-
cules of rather irregular shape and size, and
sometimes confluent.'' The notum of D. albo-
punctata is only minutely tuberculate. Further-
more, the buccal ganglia in D. citrina are some
distance from the central nerve ring whereas,
as has been pointed out, they are located di-
rectly behind the central nerve ring in D. albo-
punctata.
Cockerell and Eliot (1905) also described a
dendrodorid from La Jolla as Doridopsis vidua
(?) Bergh, 1878. The specimens had been sent
to Eliot by Cockerell in California (as had his
specimens of Dendrodoris albopunctata) and the
latter worker believed his material to be a new
Species. Eliot suggested that, if the species
were new, it should take Cockerell's manu-
script name, Doridopsis nigromaculat - Coc-
kerell later (1908) listed this form as Doridop-
sis nigromaculata C & E(vidua Bergh, var. (?}
thus suggesting that he was not in full agree-
ment with Eliot's determination.
The only other mention in the literature of
this species from California, apart from O'Do-
noghue's later comments and lists (1922a, 1926,
1927), is ina list by Kelsey (1907) of mollusks
collected in San Diego. It is apparent from
Kelsey's list that the nudibranch records, at
least, were obtained from the literature. I am
informed by Mr. Lance that nothing which fits
Cockerell and Eliot's description has ever been
seen by hirn in over ten years of intensive col-
lecting in intertidal areas, as well as in deep
water, in the San Diego area.
Eliot never commented again on this spe-
cies, and it is my opinion that, in view of the
great geographical separation of the type local-
ity of Dendrodoris vidua (Tahiti) and Cockerell
and Eliot's material [later referred to the ge-
nus Dendrodoris by O'Donoghue (1926)], it is
most probable that, if specimens which can be
referred to Cockerell and Eliot's description
are eventually found, it will be shown that they
are not synonymous with Bergh's species. For
that reason I support Cockerell's contention
that the name of this species should be Dendro-
doris nigromaculata (Cockerell and Eliot, 1905).
Doris (Asteronotus) alabastrina (Cooper,
1862) is known from only one specimen collect-
ed by Cooper under stones at San Diego Bay.
The description is very brief: ''Alabaster white,
opaque, form depressed-oval, dorsal tentacles
short, acute, branchiae of twelve simple rays
expanding in the posterior fifth of the body.
Length, four tenth in., breadth, three tenths of
an inch.'' This species was later assigned to
the genus Aldisa Bergh, 1878, by O'Donoghue
(1926) who commented, "As near as can be
judged from the meager description given by
Cooper, this animal... probably belongs to the
genus Aldisa, as his Doris (Asteronotus) san—
guinea is properly Aldisa sanguinea.'' I cannot
agree that such a decision is warranted.
Nothing which could properly be ascribed
to Cooper's species has ever been collected by
Mr. Lance in the San Diego area, and none of
the characters described by Cooper can be con-
sidered truly diagnostic. I therefore propose
that Doris (Asteronotus) alabastrina Cooper,
1862, be treated as a nomen dubium.
Cabrilla occidentalis Fewkes, 1889, the
Page 60
THE VELIGER
Vol. 4; No. 2
type species on which Fewkes based his genus
Cabrilla Fewkes, 1889, is definitely allied to the
genus Triopha Bergh, 1880, as O'Donoghue
(1926) suggested. It is known from a single
specimen obtained by Fewkes on a buoy chain
in Prisoner's Harbor, Santa Cruz Island. The
illustration which Fewkes provides does not
really resemble any of the described species of
Triopha, nor does the description of the color
(greenish brown covered with light green spots).
As no description of the radula or of the inter-
nal anatomy was included in the description, it
is impossible to decide this form's exact sys-
tematic position. I suggest, therefore, that
Cabrilla occidentalis Fewkes, 1889, be consid-
ered a nomen dubium.
The species comprising the genus Triopha
need to be studied more thoroughly in order to
determine how many species actually exist on
our coast. Marcus (1961) lists seven species
and tabulates the radular characteristics of
each. An eighth species, Triopha catalinae
(Cooper, 1863), the radula of which is unknown,
is discussed below.
A ninth species, omitted from Marcus'
list, is Triopha elioti O'Donoghue, 1921, de-
scribed from the Vancouver Island region.
O'Donoghue believed his species to be identical
with a Triopha sp. described by Cockerell and
Eliot in 1905. In 1922, O'Donoghue (1922b) dis-
covered that Cockerell in 1908 had given the
name Triopha aurantiaca to his material from
San Pedro, California. O'Donoghue then ap-
plied Cockerell's name to his specimens.
O'Donoghue was quite specific in stating that
his animals were white with orange or red
markings and compared this coloration to T.
carpenteri (Stearns, 1873) which is white to yel-
lowish white with orange and red markings. He
apparently ignored completely the fact that
Cockerell, in naming his species, stated clear-
ly that his specimens were orange.
In the museum at the University of Wash-
ington laboratory at Friday Harbor, Washing-
ton, are two specimens belonging to the genus
Triopha. I had the opportunity to examine them
briefly in August, 1960, and found that the
smaller specimen (approximately 40 mm.),
which was grayish white in formalin, had a ra-
dula which was identical with that described for
T. elioti, The number of rows in the radula
was not counted but the radula formula was
8-9.4.2.2.4.8-9.
In order to determine the original color of
this animal, I contacted the collector, Mr. Mi-
chael Marsh (Department of Zoology, Univer-
sity of California, Berkeley) who kindly pro-
vided me with the information that the colora-
tion was like Triopha carpenteri.
The second specimen referred to above
was collected during my stay at the Laboratory.
It measured over 150 mm. in length. The col-
or was yellowish with many dark brown flecks
scattered over the notum, producing a dirty
yellow. The dorso-lateral processes were
orange-red, and there were more velar proces-
ses than MacFarland (1906) figures for Triopha
carpenteri. Again, the number of rows in the
radula was not counted, but the radular formula
was 9.9-10.2.2.9-10.9, This fits the lower
limit for the number of lateral and marginal
teeth recorded for T. carpenteri.
As no orange Triophas are known from the
Friday Harbor or Vancouver Island regions, it
is evident that O'Donoghue's specimens are not
referable to Cockerell's Triopha aurantiaca,
although the radulae are similar. Further in-
vestigation will be necessary in order to deter-
mine the relationship of T. elioti to T. carpen-
teri.
As was mentioned above, the radula of
Triopha catalinae (Cooper, 1863) is not known.
The color described for this form resembles -
that of T. carpenteri and T. elioti as well as
that of T. scrippsiana Cockerell, 1915. Triopha
catalinae was first described from Santa Catali-
na Island, and later Cooper listed its range as
being from Baulines(sic., Bolinas) Bay to Cata-
lina Island, stating also that it was rare on
stones at Santa Cruz. This range approximates
the range for T. carpenteri which is known
from Bodega Bay to Laguna Beach. Cooper's
description is quite brief and offers no charac-
ters by which his species may be definitely al-
lied to T. carpenteri. Moreover, his descrip-
tion could easily be applied to T. elioti or T.
scrippsiana, although neither are known to oc-
cur within the range of T. catalinae. Marcus
- (1961) suggests that T. catalinae may have pri-
ority over T. carpenteri or T. aurantiaca
(which he believed to be identical with T. eli-
oti). I do not consider Cooper's description as
adequate enough to permit definite identification
of T. catalinae or to allow it to be compared
with the other three species in this genus which
it appears to resemble. I propose, therefore,
that Triopha catalinae (Cooper, 1863) be con-
sidered a nomen dubium.
The only orange Triophid occurring in
Southern California may possibly be Triopha
Vol. 4; No. 2
THE VELIGER
Page 61
aurantiaca if Cockerell's species from La Jolla
may be shown to be distinct from T. maculata
MacFarland, 1905. Mr. Lance informed me
that, although the specimens of Triopha in
Southern California greatly resemble immature
T. maculata, he has never seen anything re-
sembling a mature T. maculata in either inter-
tidal or deep water collections. Triopha macu-
lata was recorded by O'Donoghue in 1927 as
occurring at Laguna Beach, but he states that
his specimens ''... were orange or tawny in
color, and covered with few or many small
white spots. The processes on the sides of the
dorsum and the oral veil, the tips of the
branchiae and rhinophores were orange red."
He describes the radula as having four to five
lateral teeth and seven to eight marginal teeth
on a side in each row, but figures only the first
three lateral teeth and nine marginal teeth, the
outermost one being quite small. It is, there-
fore, quite difficult to compare his description
with T. maculata as it is known in Central Ca-
lifornia.
In July, 1961, I took with me to La Jolla
several living specimens of Triopha maculata
collected at Point Pinos on the Monterey Penin-
sula. The largest of these, about 25 mm. in
length, was beginning to show the darker color
typical of mature forms. The dots on its notum
and sides were white. The other animals were
uniformly orange-red. Ten specimens of the
southern Triopha were collected the next morn-
ing at Point Loma and the material from both
collections was compared.
Nine of the animals from Point Loma were
small (averaging about 10 mm.) and were near-
ly identical in color and in size with the small
specimen from Point Pinos, except that several
of the Point Loma animals had grayish dorso-
lateral processes which were only tipped with
orange-red.
The tenth specimen from Point Loma was
about 35 mm. in length. The ground color was
pale orange with white spots and darker dusky
orange dorso-lateral and velar processes and
branchiae. The notum and sides were speckled
with tiny brown flecks everywhere between the
white spots. These brown flecks were much
lighter than the brown flecks which caused the
darker color in the largest specimen from
Point Pinos.
All of the specimens from both localities
were identical morphologically except for the
velar processes which varied in number from
nine to 13. All bore five dorso-lateral proc-
esses Ona side, and all had four small tuber-
cles arranged longitudinally down the midline
of the notum as MacFarland figures for a ma-
ture Triopha maculata (1906, pl. XVIII, fig. 18).
Of the four small specimens from Point
Loma whose radulae were examined, all had
three hamate lateral teeth on a side in the com-
plete rows and a fourth lateral tooth which had
a poorly developed hook. The largest specimen
had five hamate lateral teeth.
All of the specimens from Point Pinos had
four lateral teeth ona side, the outermost la-
teral being well developed.
In view of the obvious absence of speci-
mens from Southern California which are
clearly referable to MacFarland's Triopha
maculata as he described and figured it, I can-
not now synonymize the Southern California
form with that which occurs in Central Califor-
nia. It seems obvious that much additional
work is necessary in order to separate clearly
the species in this genus.
Marcus (1961) has pointed out that the
number of lateral teeth increases and the num-
ber of marginal teeth decreases as the animal
gets larger and describes the origin of the spu-
rious rachidian teeth. I believe, as his discus-
sion seems to point out, that this can happen
only to a certain extent ina given species. In
considering the genus as a whole, it appears
that the radula may be of only secondary im-
portance in identifying species. I suggest that
some other set of characters must be selected
for use in separating species in this genus.
It is apparent from O'Donoghue's list (1926)
that he was unaware of the exact nature of La-
teribranchia festiva Stearns, 1873, as he placed
it with the phanerobranch dorids. It is correct-
ly placed in the genus Tritonia by Marcus (1961)
and has been known by workers on this coast
variously as Duvaucelia or Sphaerostoma fes-
tiva since 1927 when Johnson and Snook pub-
lished a short description and a colored figure
of Tritonia festiva, presumably placing this
species in its correct systematic position on
the advice of Dr. MacFarland [see also Smith
and Gordon, 1948; Steinberg (in Light, et al.),
1954].
The coloration of Tritonia festiva varies
in California from completely translucent white
to dull orange (occasionally light brown) and
translucent white with opaque white markings.
The variation of color does not seem to be con-
sistent within a given geographical range. A
Page 62
THE VELIGER
Vol. 4; No. 2
careful comparison of the description of
Sphaerostoma undulata O'Donoghue, 1924, with
a number of specimens from both Northern and
Southern California reveals no differences
other than that O'Donoghue encountered no spe-
cimens which exhibited the orange coloration
commonly seen in California. The rachidian
teeth of the largest animal in my collections (35
mm. in length, preserved, from Moss Beach,
San Mateo County, California) resemble those
described and figured by O'Donoghue for a spe-
cimen 48 mm. in length. The lateral cusps of
the teeth are very much reduced. However, in
a preserved 10 mm. long animal from the same
locality, the lateral cusps of the rachidian teeth
are much more prominent. I cannot find any
characters which distinguish S. undulata from
T. festiva and therefore synonymize the two,
with T. festiva (Stearns, 1873) taking priority.
In treating the nudibranchs from Southern
California, several comments on some of the
species described by Guernsey (1912) from La-
guna Beach must be included.
a. Her Chromodoris sp. is, as O'Donoghue
(1926) suggested, Glossodoris californien-
sis (Bergh, 1879).
b. Mr. Lance informs me that specimens
which conform to the illustration and to the
coloration described for her Genus and
Species (?) occasionally occur in great
numbers on the kelp of the San Diego re-
gion. He considers them to belong to the
genus Polycera Cuvier, 1816, and is in the
process of describing this form as a new
species.
c. Guernsey's Doriopsis fulva MacFarland
and Doris sp. are both Dendrodoris albo-
punctata (Cooper, 1863).
d. O'Donoghue (1926) synonymized her Cu-
thonia (sic.) sp. with a species described
by him (1922b) from Vancouver Island as
Cuthona concinna (Alder and Hancock, 1843)
without comparative material from both
localities. It will be necessary to obtain
specimens from both areas for comparison
before the exact systematic position of
each may be ascertained.
e. Hervia sp. ? Guernsey, 1912, was given the
name Hervia lagunae by O'Donoghue (1926).
The genus Hervia Bergh, 1871, is now con-
sidered to be a synonym of Facelina Alder
and Hancock, 1855 (see MacNae, 1954).
Guernsey's description does not permit her
species to be assigned to any of the genera
to which other species, previously consid-
ered to belong to the genus Hervia, are now
allocated. As O'Donoghue's name for this
form was based only on Guernsey's de-
scription, I suggest that the name Hervia
lagunae O'Donoghue, 1926, be considered a
nomen dubium.
f. Spurilla sp. Guernsey, 1912, was more
completely described by O'Donoghue (1927)
as Eolidina orientalis. This species has
subsequently been synonymized, rightly,
with Spurilla chromosoma Cockerell and
Eliot, 1905, by Marcus (1961).
Neither the description of Facelina stearn-
si Cockerell, 1901, in the original description
of material from San Pedro nor in the subse-
quent description by O'Donoghue (1927) of a
form from Laguna Beach which he somewhat
tentatively considered to be Cockerell's spe-
cies, give any characters by which these spe-
cimens may be assigned definitely to the genus
Facelina, as it is now defined. The same holds
true for Facelina hiltoni O'Donoghue, 1927. Ad-
ditional collecting in the San Pedro and Laguna
Beach regions may reveal specimens which
may be referable to either or both of these
species but, in such an event, further research
will be necessary in order to determine their
exact systematic position.
Coryphella cooperi Cockerell, 1901, ap-
pears to belong to the genus Coryphella and is
compared to other species of the genus by Mar-
cus (1961). However, it will be necessary to
collect further in the San Pedro area, in the
hope of obtaining specimens comparable to
Cockerell's description, before this species
may be compared fully with others in the ge-
nus.
Literature Cited
Baba, Kikutar6
1949. Opisthobranchia of Sagami Bay collected by His
Majesty, the Emperor of Japan. 4+2+194+7 pp.,
pls. 1-50. Iwanami Shoten, Tokyo.
Bergh, (Ludwig S.) Rudolph
1878. Malakologische Untersuchungen, Band 2. In:
C. Semper, Reisen im Archipel der Philippinen.
Zweiter Teil: Wissenschaftliche Resultate, Heft 14:
603-645, i-l, ple. 66-68.
1879. Onthe nudibranchiate gasteropod mollusca of the
North Pacific Ocean. Part I. Proc. Acad. Nat.
Sci. Phila. 71-132, pls. 1-8.
1880. Idem, Part II. Ibid.: 40-127, pls. 9-16.
Cockerell, Theodore Dru Alison
1901. Three new nudibranchs from California.
Malacol. 8 (3): 85-87.
Jour.
THE VELIGER
Vol. 4; No. 2 Page 63
1908. Mollusca of LaJolla, California. Nautilus 21(9): O'Donoghue, Charles H.
106-107. 1921. Nudibranchiate mollusca from the Vancouver
—— Island region. Trans. Roy. Canad. Inst. 13: 147
1915. The nudibranchiate genus Triopha in California. to 210, pls. 7-11.
Jour. Ent. Zool. Pomona Coll. 7 (4): 228-229, 2
textfigs.
Cockerell, T. D. A. and Charles Eliot
1905. Noteson a collection of Californian nudibranchs.
Jour. Malacol. 12 (3): 31-53, pls. 7-8.
Cooper, James Graham
1862. Some generaand species of California mollusca.
Proc. Calif. Acad. Nat. Sci. 2: 202-207.
1863. On new or rare mollusca inhabiting the Coast of
California. Proc. Calif. Acad. Nat. Sci. 3: 56-60.
1870. Notes on mollusca of Monterey Bay, California.
Amer. Jour. Conchol. 6: 42-70.
Eliot, Charles N. E.
1906. The genus Doriopsilla Bergh. Jour. Conch. 11:
366-367.
1907. Nudibranchs from New Zealand and the Falk-
land Islands. Proc. Malacol. Soc. London 7: 327
to 361, pl. 28.
Fewkes, J. Walter
1889, New invertebrata from the coast of California.
Bull. Essex Inst. 21: 99-146.
Guernsey, Mabel
1912. Some of the mollusca of Laguna Beach.
na Coll. Rep. Marine Lab. 1: 68-82, figs. 32-43.
Johnson, Myrtle E., & H. J. Snook
1927. Seashore animals of the Pacific coast.
illus. New York, MacMillan.
Kelsey, F. W.
1907. Mollusksand brachiopods collected in San Diego,
California. San Diego Soc. Nat. Hist. 1(2); 31-55.
MacFarland, Frank Mace
1905. A preliminary account of the Doriidae of Monte-
rey Bay, California. Proc. Biol. Soc. Washington
18: 35-54.
,
1906.
California, and vicinity.
for 1905, 25: 109-151.
MacNae, William
1954. On the status of the generic names of the nudi-
branch genera Catriona, Cratena, Hervia, Rizzo-
lia and Trinchesia. Proc. Malacol. Soc. London
31 (2): 52-55.
Marcus, Ernst
1961. Opisthobranch mollusks from California.
Veliger 3 (Suppl., pt. I): 1-84, pls. 1-10.
659 pp.,
Opisthobranchiate mollusca from Monterey Bay,
Bull. U. S. Bur. Fish.
Pomo- .
1922a Notes on the taxonomy of nudibranchiate mollus-
ca from the Pacific coast of North America. Proc.
Malacol. Soc. London 15: 133-150.
,
1922b Notes on the nudibranchiate mollusca from the
Vancouver Island region. III. Trans. Roy. Canad.
Inst. 14: 145-167, pls. 5-6.
.
1924, Notes on the nudibranchiate mollusca from the
Vancouver Island region. 1V. Trans. Roy. Canad.
Inst. 15 (1): 1-33, pls. 1-2.
i]
1926. A list of the nudibranchiate mollusca recorded
from the Pacific coast of North America, with
notes on their distribution. Trans. Roy. Canad.
Inst. 15 (2): 199-247.
1927. Notes on a collection of nudibranchs from Lagu-
na Beach, California. Jour. Ent. Zool. Pomona
Coll. 19: 77-119, pls. 1-3.
Orcutt, Charles Russell
1885. Notes on the mollusks of the vivinity of San
Diego, California, and Todos Santos Bay, Lower
California. Proc. U.S. Nat. Mus. 8: 534-552, pl.
1.
Pruvot-Fol, Alice
1954. Mollusques opisthobranches. Faune de France,
58: 460 pp., 173 textfigs., 1 pl. Paul Lechevalier,
Paris.
Smith, Allyn G, & MacKenzie Gordon, Jr.
1948. The marine mollusks and brachiopods of Monte-
rey Bay, California, and vicinity. Proc. Calif.
Acad. Sci., ser. 4, 26 (8): 147-245, pls. 3-4.
Stearns, Robert E. C.
1873. Descriptions of anew genusand two new species
of.nudibranchiate mollusks from the coast of Cali-
fornia. Proc. Calif. Acad. Sci. 5 (1): 77-78.
Steinberg, Joan E.
1954. Key to the more common opisthobranchs. In:
Light, S. F., Ralph I. Smith, et al. Intertidal In-
vertebrates of the central California coast. 264
to 270. Univ. Calif. Press, Berkeley.
Yates, L. O.
1890. The mollusca of Santa Barbara County. Bull.
Santa Barbara Soc. Nat. Hist. 1 (2): 37-45.
Page 64
THE VELIGER
Vol. 4; No. 2
A Distributional List of Southern California Opisthobranchs
by
JAMES R. LANCE
Scripps Institution of Oceanography, La Jolla, California
The opisthobranch fauna inhabiting the in-
tertidal regions along the coast of California
has become comparatively well known. The
early species descriptions of Cooper (1862,
1863), Bergh (1879, 1894), and Cockerell (1901-
1915), while of a rather cursory nature, form
the basis upon which more precise investiga-
tions by later workers have been carried out.
Outstanding among these reports are the excel-
lent anatomical accounts by MacFarland (1905-
1929) of Central California forms. The publi-
cation of a distributional list of all the known
species from the Pacific Coast of North Amer-
ica up to 1926 (O'Donoghue) has also contribut-
ed greatly to our knowledge of these mollusks.
Of more recent date is a list by Smith and
Gordon (1948) of species from the Monterey
area, anda key by Steinberg (1954) to the Cen-
tral California forms. Marcus (1961) has de-
scribed 12 new species from this coast and
provided anatomical details for 38 others.
In view of additional data obtained by vari-
ous observers and from my own collections
along the coasts of California and Baja Califor-
nia, Mexico, during the past several years, it
seems appropriate to offer a synopsis of the
bathymetric and geographic ranges of Southern
California opisthobranchs as they are now
known. Miss Joan Steinberg has kindly pro-
vided me with information regarding the north-
ern ranges of many species, and for bathyme-
tric observations I am indebted to the late Con-
rad Limbaugh, to Mrs. Limbaugh, and to Mrs.
Homer Rydell.
The marine province of Southern Califor-
nia is here geographically defined as that re-
gion between Point Conception and the Mexican
border. Excluding the cephalaspideans and
pteropods, all the regional species for which
there exist descriptions sufficient to permit
identification are included. In view of the cha-
otic taxonomic state of the genus Triopha, it
was deemed advisable to include the names of
all the species reported from Southern Califor-
nia although probably only two distinct forms
occur.
The systematic arrangement here followed
is that of Odhner (1939). For convenience, the
habitats are divided into (1) bays, lagoons, and
estuaries with their associated boat landings,
pilings, wharfs, buoys, etc.; (2) intertidal; (3)
subtidal; and (4) the kelp canopy which, within
the limits of these observations, consists ex-
clusively of the giant brown kelp, Macrocystis
pyrifera, and grows in 20-100 feet of water.
The quantitative observations are based on
some 140 field trips over the past 10 years,
principally to Mission and San Diego Bays,
Point Loma, the Coronados Islands, and the
kelp beds off La Jolla and San Diego.
The degrees of abundance are arbitrarily
designated as (1) common, which, for example,
in the case of Corambella sp., may number as
many as 140 individuals per square foot of kelp
surface; (2) uncommon, such as Crimora cone-
ja, one or two specimens of which may be ob-
served every second or third trip to Point
Loma; and (3) rare, indicating that very few
individuals have ever been collected.
Examples of prey-predator associations
such as that which exists between Armina cali-
fornica and the sea pansy, Renilla koellikeri,
are noted. The substrata for the intertidal and
subtidal habitats are rocky unless stated other-
wise.
The new ranges are followed by the old in
parentheses. A species name preceded by a
single asterisk (*) indicates that I have never
collected that form. Two asterisks (**) are
used to signify new species whose descriptions
will shortly appear in this journal. A list of
positions for the geographic points mentioned
is included.
ANASPIDEA
APLYSIIDAE
Aplysiinae
THE VELIGER
Vol. 4; No. 2
ALASKA
Sitka MO SUIN GS De 20 Wi
BRITISH COLUMBIA
Nanaimo, Vancouver
Island 49° 10'N 123° 56'W
CALIFORNIA
Humboldt Bay 40° 45'N 124° 14'W
Gualala 38° 48'N 123° 30'W
Bodega Bay 38° 18'N 123°03'W
Dillon Beach 38° 14" N) 1225 58" W
Tomales Bay 38° 14'N 122°59'W
San Francisco Bay 37° 47"N 122° 27'W
Moss Beach 37° 32'N 122° 31'W
Santa Cruz 36° 58'N 122°01'W
Pacific Grove 36 38UNP a2 55) Ww
Monterey 36° 37'N 121° 53'W
Cayucos 35°26'N 120° 54'W
Pismo Beach 35° 09'N 120° 38'W
Point Conception 34°27'N 120° 28'W
Santa Barbara 34°25'N 119° 41'W
Santa Cruz Island 34° O1'N 119° 41'W
San Pedro 33° 44'N 118° 16'W
Corona del Mar 33° 36'N 117°54'W
Newport Bay 33°36'N 117°54'W
Laguna Beach 33° 32'N 117° 44'W
Catalina Island 33°29'N 118° 36'W
Doheny Beach S32 Nalin 39) Ww.
La Jolla 32° 52'N 117° 15'W
San Diego SO CNS eS RA
Point Loma 32° 40'N 117° 14'W
MEXICO
Los Coronados Islands 32°24'N 117° 14'W
Bahia Todos Santos 31°52'N 116° 38'W
Bahia de Los Angeles 28° 55'N 113° 32'W
Isla Cedros 28° 22"N 115° 12' WwW
Punta Eugenia 727] NDING MNES EN
La Paz 24° 10'N 110° 19'W
Cabo San Lucas 22°52'N 109°53'W
FLORIDA
Manatee Bay 2730 New 82035) Wi
CHILE
Isla de Chiloé 41°52"S 73° 50'W
Aplysia californica Coorer, 1863.
Common intertidally and subtidally to 40
feet. Bodega Bay to the Gulf of California.
* Aplysia juliana Quoy & GAIMARD, 1832.
Cosmopolitan in warm seas. Coast of Cali-
fornia.
Aplysia vaccaria WINKLER, 1955.
Common intertidally. San Pedro to Point
Loma (San Pedro to Doheny Beach).
Page 65
Dolabriferinae
Phyllaplysia zostericola McCautey, 1960.
Common in bays and lagoons on the eel-
grass, Zostera marina. San Juan Island to San
Diego Bay (San Juan Island to Newport Bay).
NOTASPIDEA
Umbraculacea
TYLODINIDAE
Tylodina fungina Gass, 1865.
Locally abundant intertidally and in bays
on the yellow sponge, Verongia thiona. Cayucos
to Todos Santos Bay (Santa Barbara Island to
San Diego).
Pleurobranchacea
PLEUROBRANCHIDAE
* Pleurobranchus californicus DaLL, 1900.
Crescent City to San Diego (San Pedro to
San Diego).
* Pleurobranchus digueti ROCHEBRUNE, 1895.
San Pedro to the Gulf of California.
SACOGLOSSA
Elysiacea
ELYSIIDAE
Elysia hedgpethi Marcus, 1961,
Seasonally common intertidally, July
through September. Tomales Bay to La Jolla.
HERMAEIDAE
Hermaeinae
Hermaeina smithi Marcus, 1961.
Seasonally common intertidally,
through June.
(Tomales Bay).
January
San Juan Island to San Diego
NUDIBRANCHIA
Doridacea
EUDORIDACEA
Cryptobranchia
DorIDIDAE
Glossodoridinae
Cadlina flavomaculata MACFARLAND, 1905.
Uncommon intertidally and subtidally to 65
feet. Vancouver Island to Point Eugenia (Van-
couver Island to San Diego).
Page 66
THE VELIGER
Vol. 4; No. 2
Tn ee
Cadlina marginata MACFAaRLAND, 1905.
Common subtidally to 150 feet; not known
from the intertidal. Vancouver Island to Point
Eugenia (Vancouver Island to San Diego).
Cadlina sparsa (ODHNER, 1921).
Rare intertidally and subtidally to 131 feet.
San Diego to the Juan Fernandez Islands, Chile.
** Cadlina sp.
Rare subtidally to 140 feet.
Coronados Islands.
Glossodoris californiensis (BERGH, 1879).
La Jolla to the
Uncommon intertidally and subtidally to
100 feet. Monterey to the Coronados Islands
(Monterey to San Diego).
Glossodoris macfarlandi (COCKERELL 1902),
Uncommon intertidally at the southern end
of its range; rare subtidally to 30 feet in the
north. Monterey to the Coronados Islands (San
Pedro to San Diego).
Glossodoris porterae (COCKERELL, 1902).
Uncommon intertidally. Monterey to Ce-
dros Island (Monterey to San Diego).
SUBFAMILY incertae sedis
* G lossodoridiformia alba O'DONOGHUE, 1927.
Intertidal. Laguna Beach.
Thorunninae
Rostanga pulchra MACFARLAND, 1905.
Uncommon intertidally and subtidally to 60
feet. Vancouver Island to Chile; Japan.
* Aldisa sanguinea (Cooper, 1862).
Rare intertidally.
ego; Japan.
Bodega Bay to San Di-
Archidoridinae
Archidoris montereyensis (COOPER, 1862).
Rare intertidally and uncommon subtidally
to 150 feet. Alaska to San Diego.
Discodoridinae
Anisodoris nobilis (MACFARLAND, 1905).
Common on bay boat landings and pilings
and subtidally to 100 feet; rare intertidally.
Vancouver Island to the Coronados Islands
(Vancouver Island to Laguna Beach).
Diaulula sandiegensis (CoopER, 1862),
Uncommon intertidally; common subtidally
to 120 feet. Japan to Cape San Lucas (Japan to
San Diego).
* Discodoris heathi MACFARLAND, 1905.
Rare intertidally. Vancouver Island to La-
guna Beach.
PLATYDORIDINAE
* Platydoris macfarlandi HANNA, 1951.
Subtidal to about 516 feet. Pismo Beach.
Phanerobranchia
NONSUCTORIA
NoToporRIDIDAE
Aegires albopunctatus MACFARLAND, 1905
Seasonally common intertidally, March
through August; common subtidally to 100 feet.
Vancouver Island to the Coronados Islands
(Vancouver Island to San Diego).
POLYCERIDAE
Polycera atra MACFARLAND, 1905.
Seasonally common in bays on boat land-
ings and pilings, April through August; rare in-
tertidally and subtidally. San Francisco Bay to
the Coronados Islands (Monterey to San Diego).
** Polycera sp.
Seasonally common on offshore kelp, June
through September. Laguna Beach to the Coro-
nados Islands.
Laila cockerelli MACFaRLAND, 1905.
Sporadically common intertidally in the
spring, at other times absent; uncommon sub-
tidally to 110 feet. Vancouver Island to Cape
San Lucas (Vancouver Island to San Diego).
TRIOPHIDAE
* Triopha aurantiaca CocKERELL, 1908.
Laguna Beach to La Jolla.
Triopha carpenteri (STEARNS, 1873).
Rare subtidally to 80 feet. Dillon Beach
to San Diego; Japan (Dillon Beach to Laguna
Beach; Japan).
* Triopha catalinae (CooreR, 1863).
Santa Cruz to Catalina Island.
* Triopha grandis MAaAcFAarLanb, 1905.
Monterey to Laguna Beach.
Triopha maculata MACFARLAND, 1905.
Seasonally common intertidally; rare sub-
tidally to 60 feet. Bodega Bay to San Diego.
* Triopha scrippsiana COCKERELL, 1915.
La Jolla.
Vol. 4; No. 2 THE VELIGER Page 67
Crimora coneja Marcus, 1961. * Tritonia exsulans BERGH, 1894.
ace OR HAUCEUCONI]s LEG Care Intertidal and subtidal to 1'020 feet. Coast
SUCTORIA
ONCHIDORIDIDAE
Acanthodoris lutea MACFARLAND, 1925.
Very rare intertidally in the southern part
of its range; seasonally abundant at Moss
Beach. Moss Beach to Point Loma (Moss
Beach to Cayucos).
Acanthodoris rhodoceras COCKERELL & ELIor, 1905.
Seasonally common intertidally, April
through July; rare subtidally to 60 feet. Dillon
Beach to the Coronados Islands (Dillon Beach
to San Diego).
GONIODORIDAE
Hopkinsia rosacea MACFARLAND, 1905.
Common intertidally; rare subtidally to 20
feet. Eureka to Point Loma (Gualala to Point
Loma).
Ancula pacifica MacFaRLanD, 1905.
Very rare intertidally. Moss Beach to
Point Loma (Monterey to La Jolla).
Trapania velox (COCKERELL, 1901).
Seasonally common on bay boat landings
and pilings, July through October. San Fran-
cisco Bay to San Diego Bay (La Jolla).
CoRrAMBIDAE
Corambe pacifica MacFaRLanD & O’DONOGHUE, 1929.
Seasonally common on colonies of the
bryozoan, Membranipora serrilamella, growing
on offshore kelp, January through September;
at other times absent. Vancouver Island to
Point Eugenia (Vancouver Island to Monterey).
** Corambella sp.
This species shares the same habitat with
the above form. Although their seasonal ap-
pearance coincides, Corambe pacifica seems to
reach its maximum population density in March
and April while Corambella sp. is most abun-
dant during June and July. Vancouver Island to
the Coronados Islands.
POROSTOMATA
DENDRODORIDAE
Dendrodoris albopunctata (Cooper, 1863).
Common intertidally and subtidally to 150
feet. Monterey to Point Eugenia (Monterey to
Point Loma).
Dendronotacea
TRITONIIDAE
of Japan to Baja California (26° 14'N; 113° 13!
W); Manatee Bay, Florida.
Tritonia festiva (STEARNS, 1873),
Uncommon intertidally and subtidally to 80
feet. Vancouver Island to the Coronados Is—
lands (Tomales Point to Corona del Mar).
* Tritonia palmeri Cooper, 1862.
San Pedro.
*Tritoniopsis aurantia MATTOX, 1955.
Catalina Island.
HANCOCKIIDAE
Hancockia californica MACFARLAND, 1923.
Uncommon intertidally at the northern end
of its range, rare on floating kelp at the south-
ern extremity; unknown from any intermediate
point south of Monterey. Dillon Beach to Baja
California (26° 43'N; 114° 29.5'W) (Monterey).
DENDRONOTIDAE
Dendronotus frondosus (AsCANIUS, 1774).
Rare intertidally and subtidally to 1'312
feet; uncommon on offshore kelp. Cosmopolitan
in the Northern Hemisphere.
Dendronotus iris Cooper, 1863.
Rare subtidally to 85 feet. Vancouver Is-
land to the Coronados Islands (Vancouver Island
to Santa Barbara).
PHYLLIROIDAE
Cephalopyge trematoides (CHUNN, 1889).
Rare; pelagic.
ally in plankton tows.
Usually collected accident-
Cosmopolitan.
TETHYIDAE
Melibe leonina (GouLp, 1853).
Seasonally common on offshore kelp,
March through September; at other times rare
or absent. Alaska to La Paz Bay (Alaska to La
Jolla).
Arminacea
EUARMINACEA
ARMINIDAE
Armina californica (Coorer, 1862).
Locally common subtidally on sandy and
muddy bottoms to 289 feet in association with
the sea pansy, Renilla koellikeri, upon which it
feeds; rare intertidally. Vancouver Island to
Panama.
Page 68 THE VELIGER Vol. 4; No. 2
PACHYGNATHA California, intertidal and subtidal to 120 feet.
ANTIOPELLIDAE Also common in bays and lagoons on mud flats,
Janolus barbarensis (Cooper, 1863).
Seasonally common intertidally and in bays
and lagoons, May through August. Santa Bar-
bara to Point Loma (Santa Barbara to La Jolla).
DIRONIDAE
* Dirona albolineata COcKERELL & ELIOT, 1905.
Rare intertidally and subtidally to 115 feet.
Vancouver Island to Laguna Beach.
Dirona picta CockERELL & ELIOT, 1905.
Seasonally common intertidally, June
through August; at other times absent. Dillon
Beach to Point Loma (Dillon Beach to La Jolla).
Eolidacea
PLEUROPROCTA
CoryYPHELLIDAE
* Coryphella cooperi COCKERELL, 1901.
San Pedro.
Coryphella piunca Marcus, 1961.
Uncommon intertidally and on boat land-
ings; seasonally common on offshore kelp,
April through June. Dillon Beach to the Coro-
nados Islands (Dillon Beach to Point Pinos).
FLABELLINIDAE
Flabellina iodinea (Cooper, 1862).
Uncommon intertidally; common subtidally
to 120 feet. Vancouver Island to the Coronados
Islands (Vancouver Island to San Diego).
ACLEIOPROCTA
EUBRANCHIDAE
Capellinia rustya Marcus, 1961.
Seasonally common on offshore kelp, Fe-
bruary through August. San Francisco Bay to
Bahia de Los Angeles (Monterey).
FIONIDAE
Fiona pinnata EscuscHowrz, 1831.
Seasonally uncommon on floating wood, al-
gae, buoys, etc., April through August. If pres-
ent at all, there are often large numbers of in-
dividuals on the floating object. Cosmopolitan.
CLEIOPROCTA
FACELINIDAE
Hermissenda crassicornis (ESCHSCHOLTZ, 1831).
The most common nudibranch in Southern
boat landings, and pilings. Sitka, Alaska, to
Point Eugenia (Sitka, Alaska, to San Diego).
PHIDIANIDAE
** Phidiana Sp.
Uncommon intertidally; common subtidally
to 120 feet. Monterey to the Coronados Islands.
AEOLIDIIDAE
Aeolidia papillosa (LINNAEUS, 1761).
Seasonally rare intertidally and subtidally
to 2'493 feet, February through May. Cosmo-
politan.
SPURILLIDAE
Spurilla chromosoma CockERELL & ELIOT, 1905.
Rare intertidally. San Pedro to Point
Loma.
Literature Cited
Bergh, (Ludwig S.) Rudolph
1879. Onthenudibranchiate gasteropod mollusca of the
North Pacific ocean. Partl. Proc. Acad. Nat.
Sci. Phila. 71-132, pls. 1-8.
1894. Die Opisthobranchien.
Harvard, 25 (10): 125-233, pls.
Cockerell, Theodore Dru Alison
190la. Three new nudibranchs from California.
Malacol. 8 (3): 85-87.
Bull. Mus. Comp. Zool.
1-12.
Jour.
1901lb. Notes on two Californian nudibranchs. Jour.
Malacol. 8 (3): 121-122.
1901c. A new Tethys (ritteri) from California.
lus, 15: 90-91.
1908. Mollusca of LaJolla, California. Nautilus 21 (9):
106-107.
1915. The nudibranchiate genus Triopha in California.
Jour. Ent. Zool. Pomona Coll. 7 (4): 228-229, 2
textfigs.
Cockerell, T. D. A., & Charles Eliot
1905. Noteson acollection of Californian nudibranchs.
Jour. Malacol. 12 (3): 31-53. pls. 7-8.
MacFarland, Frank Mace
1905. A preliminary account of the Doriidae of Monte-
rey Bay, California. Proc. Biol. Soc. Washington
18: 35-54.
1906.
California, and vicinity.
for 1905, 25: 109-151.
Opisthobranchiate mollusca from Monterey Bay,
Bull. U. S. Bur. Fish.
1912.
Suppl.,
——
0 ’
1923.
cockia.
The nudibranch family Dironidae. Zool. Jahrb.
15: 515-536, pls. 30-32.
The morphology of the nudibranch genus Han-
Jour. Morphol. 38: 65-104, pls. 1-6.
Vol. 4; No. 2
THE VELIGER
Page 69
1924. Expedition of the California Academy of Sciences
to the Gulf of Californiain 1921. Opisthobranchiate
Proc. Calif. Acad. Sci., ser. 4, 13 (25):
10-12.
mollusca.
389-420, pls.
—- ——
1925-1926. The Acanthodorididae of the California
coast. Nautilus 39 (2, 3): 49-65, 94-103, pls. 2, 3.
Marcus, Ernst
1961. Opisthobranch mollusks from California.
Veliger 3 (Suppl., pt. I): 1-84, pls. 1-10.
Odhner, Nils
1939. Opisthobranchiate mollusca from the western
and northern coasts of Norway. Kgl. Norske Vi-
densk. Selsk. Skr. No. 1, 93 pp.
O'Donoghue, Charles H.
1926. A list of the nudibranchiate mollusca recorded
from the Pacific coast of North America, with
notes on their distribution. Trans. Roy. Canad.
Inst. 15 (2): 199-247.
Smith, Allyn G, & MacKenzie Gordon, Jr.
1948. The marine mollusksand brachiopods of Monte-
rey Bay, California, and vicinity. Proc. Calif.
Acad. Sci., ser. 4, 26 (8): 147-245, pls. 3, 4.
Steinberg, Joan E. 0
1954. Key to the more common opisthobranchs. In:
Light, S. F. Ralph I. Smith, et. al. Intertidal In-
vertebrates of the central California coast.
Univ. Calif. Press, Berkeley. pp. 264-270.
A New Commensal Polyclad from Panama
by
EDMUND H. SMITH
Pacific Marine Station, Dillon Beach, California
(Plate 16)
While collecting prosobranch gastropods
along the sea wall in Balboa Park, Panama City,
Panama (9°0'N., 79°30'W.), an acotylean poly-
clad was found in the mantle cavity of Nerita
(Ritena) scabricosta ornata Sowerby, 1823. The
classification of the snail was determined by
comparison with the text and figures of A. Myra
Keen (1958, p. 266, figs. 81,8la). The gastro-
pods were lodged in rock fissures in the supra-
littoral zone. The polyclads were seen in the
shell aperture when the Nerita were removed
from the substratum. When the snails were
disturbed, the flatworms returned to the man-
tle cavity before the operculum closed the shell
aperture. Some of the gastropods had as many
as eight to ten large polyclads within the man-
tle cavity. As stated in an earlier paper (Smith,
1960, p. 385), the micro-plankton or detritus
which enter the mantle cavity could hardly
serve as food for a carnivorous polyclad. Since
the polyclads were observed outside of the
feeding gastropods, it would seem that the po-
lyclads only used the snail's mantle cavity as a
retreat after feeding and for the protection of-
fered by a closed operculum.
Hoploplana luracola EK. H. SMITH, spec. nov.
The living form is elongate, becoming
slightly circular after preservation (Plate 16,
figure 1). The largest preserved specimen
measured 6.0 mm. by 3.0 mm. However, most
of the animals are of about the same size (av-
erage of six measured specimens, 5.5 mm. by
3.0 mm.). The dorsal surface is smooth with a
pair of short tentacles anteriorly.
The over-all color of the polyclad is light
brown with slightly darker dots covering all of
the dorsal surface. A dark brown stripe runs
the length of the animal, mid-dorsally.
The tentacular eyes occur in groups around
the bases of the tentacles and some appear to
occur within the tentacles (Plate 16, figure 3).
Each group consists of 14 to 16 tentacular eyes.
The cerebral eyes number 19 to 33 in each
cluster and form two irregular rows on oppo-
site sides of the mid-dorsal line. The eye clus-
ters extend both anteriorly and posteriorly
from the tentacles.
Page 70
THE VELIGER
Vol. 4; No. 2
In cleared specimens, eyes, ruffled phar-
ynx, spermiducal bulbs, and uteri can be seen
(Plate 16, figure 2).
The copulatory apparatus (Plate 16, figure
4) lies immediately behind the ruffled pharynx.
The spermiducal bulbs are slightly muscular,
lie ventrally and nearly transverse to the lon-
gitudinal axis of the body. The thin walled ejac-
ulatory duct leaves the spermiducal bulb ven-
trally, coursing dorsally, and turns anteriorly
to meet the male copulatory apparatus. The
duct enters the prostatic vesicle from the ven-
tral side. The stylet is straight and occupies
nearly the entire length of the male antrum
which is well defined and quite large.
The female genital pore lies directly be-
hind the male opening and opens into a simple
vagina which curves posteriorly. Scattered ce-
ment glands surround the vaginal region near
the female gonopore.
Occurrence: Southern middle Panama, Pa-
cific coast, Balboa Park. Panama City, in the
pallial cavity of Nerita (Ritena) scabricosta or-
nata at low water-level; December 1960. 25
specimens. Holotype: One whole mount depos-
ited in the American Museum of Natural Histo-
ry, New York, A.M.N.H. Cat. No. 501. Para-
types are in the U. S. National Museum, Wash-
ington, D. C. and the author's collection.
The word luracola is a Latin substantive in
apposition and means sack dweller. The name
was chosen to denote the fact that the polyclad
lives within the mantle cavity of the snail.
Discussion of Hoploplana luracola
Since the original five species assigned to
the genus Hoploplana by Bock (1913), there have
‘been eight more added including one from the
Pacific coast of North America. Inasmuch as
the copulatory apparatus is very similar
throughout the genus (Hyman, 1953, p. 346), ex-
ternal characters become important taxonomi-
cally. Hoploplana luracola is distinguished
from its closest geographic neighbor H. cali-
fornica Hyman, 1953, by lacking a papillate dor-
sal surface and having cerebral eyes extending
behind the tentacles and a longer stylet. The
nearest morphologically related species seems
to be H. deanna Kato, 1939, from Mutsu Bay,
Japan. This species can be distinguished from
H. luracola by the different arrangement of the
cerebral eye clusters and the number of tenta—
cular eyes.
Literature Cited
Bock, S.
1913. Studien tiber Polycladen.
2: 31-344, pls. 3-10.
Hyman, Libby H.
Zool. Bidr. Uppsala,
1953. The polyclad flatworms of the Pacific coast of
North America. Bull. Amer. Mus. Nat. Hist.
100 (2): 265-392.
Kato, K.
1939. The Polyclads of Mutsu Bay. Sci. Rep. Tohoku
Univ. ser. 4, 14 (4): 143-153, pls. 8-9.
Keen, A. Myra
1958. Sea shells of tropical west America; marine
mollusks from Lower California to Colombia.
Stanford, Calif., Stanford Univ. Press; xi + 624
pp., illus.
Smith, Edmund Hobart
1960. Ona new polyclad commensal of prosobranchs.
An. Acad. Brasil. Cienc. 32 (3, 4): 385-390, pl. 1.
Explanation of Plate 16
Haploplana luracola E. SmirH, spec. nov.
clarified worm.
Figure 1: Dorsal view of living worm.
Figure 3: Antero-dorsal part of body, showing eye patterns.
Figure 2: Dorsal view of
Figure 4: Diagram of copu-
latory apparatus (reconstructed).
a-male antrum, c - cement gland, d - pharynx, e¢ - ejaculatory duct, f - female genital pore, g - granule
glands, h - spermiducal bulb, i - eyes, j - sperm, k - uteri, | - dark brown dorsal area, m - male genital
pore, o - lighter brown dots, p - prostatic vesicle, s - stylet, t - tentacle, v - vagina, y - sperm duct.
THE VELIGER, Vol. 4, No. 2
Figure 1 Figure 2
—— =
LLL
a pees 72
Fase y ey:
nit!
rs
3
=
SSS
oo
= TT
. i wo
Pas ae
Yj esc ooe
im y
Figure 4
Surrn, del.
[SMITH] Plate 16
Vol. 4; No. 2
THE VELIGER
Page 71
A New Subgenus and Species of Coral-Inhabiting Barnacle
from the Gulf of California
by
Vicror A. ZULLO
Department of Paleontology, University of California, Berkeley 4, California
(Plate 17, 2 Textfigures)
The balanomorph barnacle genera Creusia
Leach, 1817, and Pyrgoma Leach, 1817, which
occur embedded in the coralla of anthozoan and
milleporine corals, have traditionally been
considered derivatives of Balanus DaCosta,
1778, through loss and concrescence of various
compartmental plates of the shell. The grada-
tional aspect of the series Balanus (shell with
six compartmental plates)-Creusia (shell with
four compartmental plates)-Pyrgoma (shell
formed of a single plate) has been recognized
for some time. Darwin (1854, p. 375) was re-
luctant to consider Creusia apart from Pyrgo-
'ma stating that"had not this genus already been
adopted by several authors, I should not, I
think, myself have formed it. .'', Elsewhere
he (Darwin, loc. cit., p. 359) observed that ''the
subgenus Creusia is closely, perhaps too
closely, allied to Pyrgoma"'. Species variously
referred to Creusia and Pyrgoma are known
which are characterized by an intermediate
stage in the concrescence of the compartmental
plates and appear to form ''connecting links"
between typical Creusia and typical Pyrgoma.
Recently, on the basis of these intermediate
forms, Brooks and Ross (1960, p. 361) conclud-
ed that Darwin's uncertainty was justified and
synonymized Creusia with Pyrgoma.
Withers (1926, 1929, 1935) has discussed
the phylogeny of these coral-inhabiting barna-
cles in respect to their fossil record and con-
cluded that ''Pyrgoma is, no doubt, a derivative
from Creusia, but certain Miocene forms of
similar coral-living barnacles have six com-
partments like Balanus (Withers, 1929, p. 560),
so that in this little group alone there is a tran-
sition from forms having six compartments to
a single shell" (Withers, 1935, p. 38). Evidence
of the transition from forms having six com-
partmental plates to those having four, how-
ever, has been lacking. Apparently the Creu-
sia condition was derived through the loss of
the carinolaterals of the Balanus ancestor (Wi-
thers, 1935, p.38). However, the six-plated
Miocene species, Balanus duvergieri (de Ales-
sandri), discussed by Withers (1929, p. 560) is
a Balanus in all respects but habitat and no
form is known that exhibits any indication of
reduction of the carinolaterals toward the
Creusia condition. Withers (1935, p. 38) pre-
sents Acasta sporillus Darwin as an example
of an intermediate form in which ''the carino-
laterals do not reach to the basis and appear to
be on their way to being crowded out''. How-
ever, the loss of carinolaterals in A. sporillus
need not necessarily have a direct relationship
to the development of Creusia from Balanus.
The discovery of a new coral-inhabiting
barnacle in the Gulf of California lends added
strength to the hypothesis of the derivation of
the Creusia condition from a Balanus ancestor.
Both fossil and living specimens of this barna-
cle have been found embedded in coralla of the
hermatypic stony coral Porites californica Ver-
rill from several localities throughout the Gulf
(textfigure 2). This barnacle is creusoid in all
respects (e.g., the cup-shaped to sub-cylindric
basis, the exterior ornamentation of the parie-
ties, the extension of the sheath nearly to the
basal margin and the continuous lower edge of
the sheath, the plate-like nature of the internal
ribs or longitudinal septa, and the creusoid na-
ture of the opercular valves) except that it has
a shell composed of six plates as in the genus
Balanus. For this reason a new supraspecific
taxon, here considered as a subgenus of Bala-
nus, is erected for this species.
Page 72
THE VELIGER
Vol. 4; No, 2
Cirripedia
THORACICA Darwin
Balanomorpha PILSBRY
BALANIDAE GRAY
Balaninae Gray
Balanus DA Costa
Hexacreusia ZULLO, subgen. nov.
DIAGNOSIS
Compartmental plates six; shell depressed;
parieties solid; radii broad, solid, with thick,
septate edges; sheath extending nearly to basal
margin; lower edge of sheath free, continuous;
basis calcareous, solid, cup-shaped to sub-
cylindric; opercular plates as in Creusia; ap-
pendages and mouth parts balanoid.
TYPE SPECIES
Balanus (Hexacreusia) durhami Zullo, spec.
nov.
ADDITIONAL SPECIES
Possibly Balanus duvergieri (de Alessan-
dri).
GEOLOGIC RANGE
? Early Miocene (Balanus duvergieri); late
Pliocene to Recent (Balanus durhami).
HABITAT
Living in the coenosteum of the coral
Porites.
REMARKS
This new subgenus is created to include a
species which in every respect, excepting the
number of compartmental plates in the shell
wall, could be assigned to the genus Creusia.
At least some forms included within Creusia
appear to have developed from an hexacreusoid
ancestry. The creusoid shell morphology ei-
ther resulted through the loss of the carinolat-
erals or through their concrescence with the
laterals or, possibly, the carina. The available
evidence, although negative, favors the loss of
the carinolaterals, as there have been no rec-
ords, either extant or fossil, of intermediate
stages of fusion between the six- and four-plate
condition.
The conclusions reached by Brooks and
Ross (1960) regarding the classificatory signi-
ficance of the presence of separate compart-
mental plates or sutures in the shell wall as the
distinguishing feature in the separation of
Creusia from Pyrgoma could also be extended
to Hexacreusia. However, where they regard
the presence of intermediate stages in the de-
velopment of the concrescence of the shell as
indicating that the genera Pyrgoma and Creusia
should not be separated, I would interpret these
intermediate stages in development as indicat-
ing a phylogenetic relationship between the two:
Creusia with the shell of four, unfused com-
partmental plates giving rise to Pyrgoma with
a shell composed of fused plates. Any closely-
linked, phylogenetic series should be expected
to include forms of intermediate nature, but
these forms do not govern the validity of recog-
nizing and delimiting categories (in this case
the end members) within the series.
This latter concept is exemplified by He-
xacreusia. In synonymizing Creusia with Pyr-
goma, Brooks and Ross (1960, p. 361) modified
the diagnosis of Pyrgoma "to include all the
obligate, epizoic, tetramerous barnacles with
their basis invaginated in the corallum of live
corals". Application of the criteria used by
Brooks and Ross (loc. cit., pp. 359-362) to the
relationship of Pyrgoma to Balanus in view of
the new six-plate, creusoid species here de-
scribed would result in the suppression of Pyr-
goma inasmuch as these genera differ only by
the number of compartmental plates in the shell
wall.
Presently, Hexacreusia is definitely repre-
sented only by the type species. However, the
specimens from the Miocene of Bordeaux,
which had been assigned by Withers (1929, p.
560) to Balanus (Balanus) duvergieri (de Ales-
sandri) are possibly referable to Hexacreusia.
Withers' assignment of this species to the sub-
genus Balanus was based on the irregularly
porous nature of the parieties of some of the
specimens. The parieties of the remaining
specimens were noted as being solid. These
pores are probably homologous with those in
some individuals of Creusia described by Dar-
win (1854, p. 377) and interpreted to represent
spaces left by the incomplete filling of the in-
ternal rib interspaces, and are not, therefore,
homologous with the parietal tubes in the sub-
genus Balanus. Withers (1929, pp. 565-566)
considered Balanus duvergieri as representing
an early stage in the development of Creusia.
Vol. 4; No. 2
THE VELIGER
Page 73
Balanus (Hexacreusia) durhami ZuLLO, spec. nov.
(Textfigure 1: Plate 17)
DIMENSIONS
of Holotype, University of California Museum
of Paleontology (hereafter referred to by the
abbreviation UCMP) No. 34'689, height (ex-
cluding elongate basis) 0.7 mm., carinorostral
diameter of orifice 0.9 mm., carinorostral di-
ameter of base 3.0 mm. The height of the shell
including the elongate basis may be as much as
4.5 mm.
DESCRIPTION
Shell small, depressed conic; orifice small,
subtrigonal to diamond-shaped, not toothed;
parieties solid; surface of parieties pink in
color, somewhat regularly ribbed; ribs project
from base of compartmental plates, giving stel-
late appearance to shell; radii broad, glossy,
lighter pink in color than parieties or white,
vertically striate; summits of radii horizontal;
sutural edges of radii thick, with primary and
secondary denticulae; alae thin; sheath long,
extending nearly to basal margin, lower edge
free, continuous; interior of parieties ribbed;
ribs plate-like in appearance, continuing un-
derneath edge of sheath; basis cup-shaped or
conical, up to three and one-half times as long
as height of compartmental plates, solid, radi-
ally ribbed; ribs and alternating furrows of
basis correspond to ribs and furrows of parie-
ties; ribs on interior of basis finely beaded.
Scutum white with pink-tinged, beaked
apex; tergal part reflexed at right angles to
rest of plate; exterior ornamented by thin,
closely-spaced growth lines; irregular, longi-
tudinal furrow divides exterior of plate in posi-
tion of (adductor ?) ridge on interior; basal
margin sinuous, longer than tergal margin; oc-
cludent margin straight, toothed; articular
ridge high, almost as long as tergal.margin;
articular furrow narrow, shallow; thin ridge
(adductor ?) extends from middle of scutum to
basal margin and continues along basal margin
to basioccludent angle; shelf continuous with
occludent margin and occludent side of the ar-
ticular ridge projects out over half the interior
of the scutum leaving a deep pocket beneath it;
free (basal) edge of shelf sinuous; shelf widest
in apical part, narrowing towards and termi-
nating in basioccludent angle; pit for depressor
muscle small, deep, situated on edge of basal
Margin at point of reflection of tergal part of
scutum; ''rostral tooth'’' suggested by intersec-
tion of occludent shelf and basal margin at ba-
sioccludent angle.
Figure 1:
durham: ZULLO, spec. nov., x 25.
paratype UCMP no. 34691;
paratype UCMP no. 34690;
paratype UCMP no. 34691;
paratype UCMP no. 34690.
Opercular plates of Balanus (Hexacreusia)
a-exterior of tergum,
b-exterior of scutum,
c-interior of tergum,
d -interior of scutum,
Tergum white in color, tinged pink at apex;
carinal part of tergum curving inwardly; arti-
cular ridge low, thin; articular furrow broad,
shallow; depressor muscle crests well devel-
oped; scutal and carinal margins straight; spur
broad, one-half width of basal margin, situated
close to, but differentiated from, the basiscutal
angle; spur long, length approximating width;
end of spur broadly rounded; spur furrow not
sharply differentiated.
Labrum deeply notched with three promi-
nent, blunt, triangular teeth and one or two
smaller teeth on each side.
Palpi with numerous, closely clustered,
short, curved, pectinate spines arranged in two
parallel rows along superior margin; a few long
spines situated on inner margin.
Mandibles with five teeth including inferior
angle; first, second, and third teeth large, dis-
tinct; second and third teeth bifid; fourth and
fifth teeth smaller than preceding; fourth tooth
bifid; fifth tooth trifid; distance between first
and second tooth one and one-half times greater
than distance between second and third tooth;
Page 74
THE VELIGER
Vol. 4; No. 2
first, second, and third teeth occupying nearly
three-fourths entire cutting-edge; inferior
margin of mandible and posterior surface of
mandible hirsute.
Inner maxillae with irregular, nearly
straight cutting edge; rudimentary notch located
below uppermost two large spines; two smaili
spines situated in notch; five large spines with
a few smaller spines situated below notch; low-
ermost two of the large spines as large as the
large spines above notch; large spines followed
by approximately five small spines; edge below
inferior angle bearing several small spines.
Outer maxillae of two lobes (as figured by
Hoek, 1913, pl. 27, fig. 8; and Nilsson-Canteil,
1921, textfig. 79e), bearing numerous pectinate
bristles.
Rami of Cirrus I unequal, densely setose;
outer ramus about one-third again as long as
inner ramus, curved posteriorly in opposing
direction from all other rami.
® late Pliocene
® Pleistocene
@ Recent
Figure 2: Late Cenozoic distribution of Balanus (Hexa-
creusia) durhami ZuLLo, spec. nov. embedded in Porites
californica VERRILL in the Gulf of California.
1; northernmost occurrence of Porites californica
(CAS locality 36745)
Rami of Cirrus II similar in structure to
those of Cirrus I, densely setose, unequal, with
outer ramus being longer.
Cirrus III more similar in structure to
Cirri I and II than to Cirri IV to VI; rami of
Cirrus III not as densely setose as those of Cir-
ri I and II, subequal with outer ramus longer;
articles 1 through 6 on outer ramus (counting
distally from pedicel) bearing teeth on posteri-
or and upper margins.
Cirri IV through VI with subequal rami,
inner rami slightly longer than outer rami; in-
termediate segments of rami bearing one to
three pairs of setae on posterior margins.
The number of articles on the individual
cirri (paratype California Academy of Sciences
No. 12'372) are as follows:
Cirrus: I Il. TL. IVG Sg Vea
Outer Ramus: 10 7 7, 10> s13as
Inner Ramus: 6 5 6 10° 14 Sg
HOLOTYPE
UCMP No. 34'689, from UCMP locality A-
3652. Paratypes: UCMP Nos. 34'690, 34'691,
from UCMP locality A-3640; UCMP No. 34'692,
from UCMP locality A-3642; California Acade-
my of Sciences No. 12'372, from CAS locality
36'745.
OCCURRENCE
(Textfigure 2)
Late Pliocene, Gulf of California, UCMP
localities A-3519, A-3534; Pleistocene, Gulf of
California, UCMP localities A-3525, A-3547,
A-3582, A-3584, A-3596; Recent, Gulf of Cali-
fornia, UCMP localities A-3640, A-3642, A-
3646, A-3652, A-3653, A-3654, A-3663, Cali-
fornia Academy of Sciences locality 36'745.
RANGE
Late Pliocene to Recent, Guli of California.
REMARKS
Balanus durhami differs from all known
species of Balanus and Creusia in the posses-
sion of an occludent shelf on the interior of the
scutum (textfigure ld). The tergum somewhat
resembles that of Creusia spinulosa var. bre-
viterga Hiro (1938, p. 397) but can be distin-
guished by its longer, narrower spur (textfigures
la, lc). The opercular plates of B. durhami do
not appear to be as variable in form as those
described by Hiro (1938).
Tue VELIGER, Vol. 4, No. 2 [ZuLLo] Plate 17
Figure 6 Figure 7 Figure 8
Balanus ( Hexacreusia) durhami ZULLO, subgen. et spec. nov.
Figure 1: Holotype UCMP no. 34680, x 2.5 (center specimen); Figure 2: Para-
type UCMP no. 34692 and associated individuals embedded in the coenosteum of
Porites californica VERRILL from UCMP locality A-3642, x1. Figures 3 to 8: Mouth
parts and appendages of paratype CAS no. 12 372, x 70. Figure 3: Labrum (teeth
partially obscured); Figure 4: Right mandible; Figure 5: Right inner maxilla;
Figure 6: Right outer maxilla; Figure 7: Right labial palp; Figure 8: Right
Cirrus ITI (outer ramus above).
ZULLO, phot.
Vol. 4; No. 2
THE VELIGER
Page 75
The mouth parts and appendages of species
of Creusia and Pyrgoma have seldom been fi-
gured. In the case of Balanus durhami the
mouth parts and appendages of only a single
specimen (paratype California Academy of Sci-
ences No. 12'372, Plate 17, figures 3-8) were
available as the corals from other localities
had been cleaned in sodium hypochlorite, de-
stroying the bodies of the barnacles. Compari-
son of the mouth parts and appendages of B.
durhami with figured specimens of Creusia and
Pyrgoma failed to uncover any significant dif-
ference, except in the possession of teeth on the
third cirrus of B. durhami.
Balanus durhami resembles species of the
subgenus Armatobalanus and some species of
the subgenus Balanus in bearing teeth on the
cirri. However, the teeth are borne only on the
anterior margins of segments of the outer
ramus of Cirrus III in B. durhami, whereas
teeth are found only on Cirrus IV in species of
Armatobalanus, and on both Cirri III and IV in
B. trigonus Darwin and B. perforatus Brugui-
gre.
teeth on the basal seven segments of Cirrus IV
in specimens of Pyrgoma anglicum Sowerby
from the Moroccan coast.
Balanus durhami is found embedded in the
coenosteum of the coral Porites californica
Verrill. The parieties of large individuals are
completely encrusted by the coenosteum; a
small hole marks the position of the buried ori-
fice. The presence of B. durhami is usually
indicated by a slightly raised mound on the sur-
face of the corallum. This mound is bounded
by a fracture zone where the overlying coenos-
teum has been fractured by the upward growth
of the barnacle. This fracture zone corres-
ponds to the basal margin of the parieties.
This species is named in honor of Dr. J.
Wyatt Durham of the Department of Paleon-
tology, University of California.
Broch (1927, p. 30) noted the presence of
I would like to thank Mr. William A. New-
man of the Department of Zoology, University
of California, for his advice in the preparation
of this paper and for the dissection of the mouth
parts and appendages of Balanus durhami. The
illustrations of the opercular plates were pre-
pared by May Blos.
Literature Cited
Broch, H.
1927. Studies on Moroccan cirripeds (Atlantic coast).
Bull. Soc. Sci. Nat. Maroc. 7: 11-38, pls. 1-4,
textfigs. 1-10.
Brooks, H. K., & A. Ross
1960. Pyrgoma prefloridanum, anew species of cirri-
ped from the Caloosahatchee marl (Pleistocene) of
Florida. Crustaceana 1 (4): 353-365, pls. 5, 6,
textfigs. 1-4.
Darwin, Charles
1854. A monograph onthe sub-class Cirripedia. Bala-
nidae, Verrucidae. Ray Soc. London, pp. 1-684,
figs. 1-11, pls. 1-30.
Hiro, F.
1938. Studies on the animals inhabiting reef corals, II.
Cirripeds of the genera Creusia and Pyrgoma.
Palao Trop. Biol. Stat. Stud. no. 3: 391-416, figs.
Vay ple) ale
Hoek, P. P. C.
1913. The cirripedia of the Siboga-Expedition, B. Cir-
ripedia sessilia, Siboga-Exped. Monog. 31b: xxv
+ 129-275, figs. 1, 2, pls. 11-27.
Nilsson-Cantell, C. A,
1921. Cirripedien Studien. Zur Kenntnis der Biolo-
gie, Anatomie und Systematik dieser Gruppe.
Zool. Bidrag fr. Uppsala 7: 75-395, textfigs. 1 to
89, pls. 1-3.
Withers, T. H.
1926. Barnacles of the Creusia - Pyrgoma type from
the Pleistocene of Barbados. Ann. Mag. Nat. Hist.,
Ber Dy Ws NeO, fle Ih sts Wo
1929. The phylogeny of the cirripedes Creusia and
Pyrgoma. Ann. Mag. Nat. Hist., ser. 10, 4: 559
to 566, pls. 10, 11.
1935. Catalogue of fossil cirripedia in the Department
of Geology, II, Cretaceous. Brit. Mus. (Nat. Hist. )
London, i-xvi, 1-434, figs. 1-64, pls. 1-50.
Page 76
THE VELIGER
Vol. 4; No. 2
A Discussion of Vexillum regina (Sowersy, 1825) and Related Species,
With Description of a New Subspecies
by
JEAN M. CATE
Conchological Club of Southern California, Los Angeles 7, California
(Plates 18, 19, 20; 1 Textfigure)
In the study necessary to ascertain that
Vexillum coloscopulus J. Cate, 1961, was a hith-
erto undescribed species, I realized through an
analyeis of the works of many authors that at
least five additional closely related species
were included in the same complex; .all are
similar enough to have been either confused
with one another or placed in synonymy. How-
ever, a careful diagnosis of the most outstand- —
ing characters of each species brought out that
these six forms are confused chiefly because of
their similarity of color, whereas the morpho-
logical differences, considered separately, pro-
vide a sufficient basis for separating them into
six different categories.
It became apparent that a further study of
the entire group was necessary in order to
clarify the confusion encountered at every turn;
1 shall attempt here to outline the results of
this study and illustrate the identifying features
of each species in the hope that other collectors
who have been similarly puzzled by this group
may be able to identify their specimens without
question,
The first species encountered in the study
was Vexillum regina (Sowerby, 1825), as this
was the name incorrectly applied to the speci-
men which is now the holotype of V. coloscopu-
lus J. Cate, 1961. The next species to be con-
sidered in the course of comparing these shells
was V. compressum (Sowerby, 1874), which at
first glance appears to be a dwarf form of Vv.
regina. Two additional species, V
(Lamarck, 1811) and V. vittatum (Swainson,
1821), which are perhaps the two most difficult
to separate, were included in the study because
of their similarity in color and pattern and
their superficial resemblance to V. regina.
The final species, from the type locality of V.
coloscopulus, is an intermediate form between
that species and V. regina; this had been erro-
neously figured by Reeve, Sowerby, and Tryon
as Mitra melongena Lamarck, 1811. This last
form will be described here as a new subspe-
cies of V. regina.
The important illustrated works on Mitri-
dae are unfortunately few; the most complete of
these, in chronological order, are by Kiener
(1839); Kuster (1841); Reeve (1844-45); Chenu
(1860); Sowerby (1874); and Tryon (1882). There
are also several helpful papers on the Mitridae
of various Indo-Pacific localities, for example,
Dautzenberg and Bouge (1922, 1933), and Daut-~
zenberg (1935), but the approach in these pa-
pers is more in the nature of annotated geo-
graphical faunal lists rather than a complete
monograph of the family, and consequently they
do not include all the species under discussion
here. Nevertheless, these are the most modern
works on the group at the present time and pro-
vide much useful information. Other than the
above-mentioned references and three or four
important papers describing many new species
but without illustrations, the only sources of
information available on the worldwide Mitri-
dae are scattered locality records or occasion-
al descriptions of new species.
These works, therefore, formed the basis
of the present study. Although numerous other
references were consulted, those listed here
were the only ones to contribute any informa-
tion pertaining to the differences or similari-
ties among the six species.
Vexillum regina (Sowerby, 1825)
(Plate 18, figures la, lb; Plate 19, figure 1)
Type locality: China Seas
The original citation of Vexillum regina
presents no technical description of any sort.
In Genera of Shells Sowerby (1825) figured the
Vol. 4; No. 2
ventral aspect of this species as one of eight
typical representatives of the genus Mitra; the
only mention of its name appears in the caption
to Plate 250 (op. cit.), merely as ''4. Mitra Re-
gina''". This original figure, however, is an ex-
cellent illustration of the species and leaves no
doubt as to its identity.
Kiener (loc. cit.) published fairly good like-
nesses of both dorsal and ventral views, al-
though with some exaggeration as to the shell's
specific characters; he included, however, a
very complete description and cited the type
locality as ''China Seas".
Kiister (loc. cit.) figured Vexillum regina
rather well as compared with many of the other
species in his ''Conchylien-Cabinet", the chief
fault with his illustration being a somewhat
foreshortened last whorl which makes the spire
appear exaggeratedly high. Kister's descrip-
tion is complete enough and cites the type lo-
cality as ''The Chinese and Indian Oceans".
Reeve (loc. cit.) published only one view
(this time the dorsal side) of the species, slight-
ly more obese than normal but reasonably typi-
cal and well colored; his brief description is
adequate except for ignoring the coarse, heavy
surface ornament which is one of the species'
most distinguishing features. Reeve correctly
pointed out that Vexillum regina could be dis-
tinguished from V. taeniatum (Lamarck) ''by
the angular structure of the ribs near the su-
tures",
Chenu (loc. cit.) included a good represen-
tation of the species, correctly identified.
Nearly fifty years after the original publi-
cation of the species, Sowerby figured the ven-
tral aspect of Vexillum regina in Thesaurus
Conchyliorum, this time using a different, more
perfect specimen for his illustration which re-
veals more brilliant colors than the original
figure. He dismissed the species with only a
brief reference to Genera of Shells and still
furnished no written description but added the
type locality as ''Moluccas"'.
Tryon (loc. cit.) merely copied the second
Sowerby figure, described the colors of the
stripes on the shell, and mentioned its size and
locality.
Vexillum regina may be readily recognized
by its heavy, coarse sculpture, its slender, tur-
riculate form with attenuated spire, its gradate
sutural ramp, angular outlines, and constricted
lip. This species has been recorded from Zan-
zibar and from the Andaman Islands, in addition
to the Moluccas.
THE VELIGER
Page 77
Vexillum compressum (Sowerby, 1874)
(Plate 18, figures 2a, 2b; Plate 19, figure 2)
Type locality: Moluccas
This species is encountered in the works
of Chenu and Reeve prior to its description by
Sowerby in 1874, though it is not recognized as
distinct. Chenu indicated two different species
under the name Vexillum taeniatum (Lamarck);
one of these is V. compressum, the other a
typical V. taeniatum. Reeve considered V.
compressum merely the young state of V. taen-
iatum, though remarking at the same time upon
its different form and recurved base; he based
his judgment on the similarity of color and on
its being found in the same locality. Tryon al-
so included this species with V. taeniatum as a
narrower, juvenile form.
Sowerby first recognized this small, rare
form as a separate species, stating "it is much
narrower and more attenuated and laterally
compressed than M. regina, narrowed and re-
curved anteriorly." Examination of actual spe-
cimens of Vexillum compressum gives validity
to Sowerby's convictions; the shells are almost
like miniatures of V. regina, though a close in-
spection reveals the following differences: V.
compressum is adult at about one-half to two-
thirds the size of a typical adult specimen of V.
regina; unlike that of V. regina, the spire of V.
compressum is shorter than the last whorl, and
there are many more raised labral lirae than
in the typical V. regina. Its sutural ramp is
more rounded, its early whorls are compara-
tively larger than those of V. regina despite its
smaller size, the axial costae of the adapical
whorls are smooth, not nodose, and finally the
siphonal canal is more acutely curved in V.
compressum and inclined ina different direc-
tion.
The Kiener and Kiister monographs do not
refer to the species.
Vexillum compressum has been recorded
from the Philippine Islands, although the rec-
ords are somewhat ambiguous in the light of its
atum. Specimens included in the present study,
however, were collected at Subic Bay, Davao,
and Mindoro in the Philippines, giving s0me
credence to the earlier records.
Vexillum coloscopulus J. Cate, 1961
(Plate 18, figures 3a, 3b; Plate 19, figure 3)
Type locality: Balabac Island, Philippines
This species differs from Vexillum regina
in the following ways: the surface ornament is
smoother and is continuous over the axial cos-
Page 78
THE VELIGER
Vol. 4; No. 2
tae; the sutural ramp is rounded instead of
gradate; the shape is more fusiform than tur-
riculate; the aperture is straight, not constrict-
ed; there is no pseudumbilicus; there are sev-
eral faint labral lirae instead of only a few
raised ones, and the pattern arrangement and
color are different. There is no central thread
on the white band, the black areas are wide
zones rather than narrow borders outlining the
white band as in V. regina, and the white band,
narrower than in the other compared species,
is centrally placed on the adapical whorls.
No reference or illustration for this spe-
cies was found in the early literature. It is
presently known from only four specimens, the
holotype from Balabac, two hypotypes which
were collected in the Sulu Archipelago and at
Zamboanga, and an additional specimen whose
locality is given only as "Philippine Islands".
Of the three remaining species in the com-
plex, the first rough scrutiny made it possible
to divide the available specimens, on the basis
of spire ornament, into two main groups. Cer-
tain of the specimens were seen to have many
very closely-spaced, well-defined axial costae
on the upper whorls, and fewer, flatter costae
on the last whorl; the others possessed approx-
imately the same number of axial costae on the
last whorl as on the upper ones, and these were
all equally sharply defined. After the two
groups had been separated, it was interesting
to note that the specimens with closely-spaced
adapical costae had all been collected in
Queensland, Australia, while the shells in the
second group were all from the southern Phil-
ippines, mainly from Balabac Island or the ad-
jacent Sulu Sea. It is the Philippine group
which I consider a subspecies of Vexillum re-
Further study after the first separation re-
vealed additional morphological differences be-
tween the two groups; the Queensland shells
were more obese, their spires proportionately
shorter and less turriculate, their axial costae
less pronounced and less regularly spaced,
their colors less variable, and so on.
The next step was to identify the Queens-
land species; close study of this large group
brought out two subgroups which could be sepa-
rated from one another, first, by the interstitial
striae present in one but lacking in the other;
also by the more obese form and by the flatter,
more numerous axial costae of one of these. It
was then apparent that these two Queensland
forms are the species respectively described
as Vexillum taeniatum (Lamarck, 1811) and V.
' vittatum (Swainson, 1821).
Vexillum vittatum
has been considered a synonym of V. taeniatum
by some workers, yet the diagnostic morpholo-
gical characters of both seem clearly separa-
ble. The question remained as to whether these
separating characters are sufficiently strong
and numerous to define separate species,
Vexillum taeniatum (Lamarck, 1811)
(Plate 18, figures 4a, 4b; Plate 19, figure 4)
Type locality: Indian Ocean
Lamarck's original description of Vexillum
taeniatum is somewhat sketchy, though his ref-
erence to a figure in the Encyclopédie Mé-
thodique and one phrase in his description def-
initely separate this species from the others:
this is his reference to "petites c6tes longitu-
dinales, obtuses, fréquentes, peu élevées, et de
stries transverses qui ne paroissent bien qu'
entre les cétes''. This is the only species of
the six under consideration which possesses
transverse interstitial striae; in the other spe-
cies the spiral ornament is continuous, even
though in V. regina this ornament is nodose
where it intersects the costae. Therefore, one
of the two Queensland species can be definitely
identified as V. taeniatum. All of the available
study specimens which I have assigned to this
species agree closely with the original descrip-
tion, and with the figure in the Encyclopédie
Méthodique referred to by Lamarck. Other
separating features include a more obese form,
more closely-spaced, flatter axial costae on
the last whorl, and a rounded sutural ramp.
Kiener illustrates dorsal and ventral views
of what may have been Lamarck's holotype of
Vexillum taeniatum, clearly showing emphasis
on the obese form and interstitial striae which
are its most important diagnostic features.
Reeve's two illustrations labelled Vexillurm
taeéniatum leave much to be desired: one (Conch.
Icon., fig. 52a) represents a fine specimen of V.
compressum (Sowerby), the other (ibid., fig.
_52b) adequately indicates the general shape of
V. taeniatum but fails to show the interstitial
striae. Reeve considered V. compressum
merely a juvenile example of V, taeniatum, ap-
parently overlooking the several morphological
differences between these two species,
Kuster illustrated this species with exag«
gerated drawings but fortunately emphasized
the important characters.
Chenu followed Reeve's error in consider-
ing Vexillum taeniatum and V. compressum
synonymous; he illustrated both species very
TuHE VELIGER, Vol. 4, No. 2 [ J. Cate] Plate 18
Fig. 1 a Sz, QE) Fig. 3a Fig. 4a Fig. 5 a
Vexillum r. V. compressum V. coloscopulus V. taeniatum V. vittatum
regina
Fig. 6 a
V. r. filiareginae
Pierson, Oswap, & Pierson, photo. except Figures 1a & 1b: Courtesy of Stanford University
Tue VELIGER, Vol. 4, No. 2 [ J. CATE] Plate 19
| = aaecae -
see
,eee4
Figure 1 Figure 2 Figure 3
Vexillum r. regina (SOWERBY) Vexillum compressum (SOWERBY) Vexillum coloscopulus J. CATE
Figure 4 Figure 5 Figure 6
Vexillum taeniatum (LAMARCK) Vexillum vittatum (SwAtnson) Vexillum r. filiareginae subsp. nov.
Photographic enlargements showing distinctive sculpture
Pierson, OswALD, & PIERSON, photo.
Vol. 4; No. 2
THE VELIGER
Page 79
well but applied the name V. taeniatum in both
instances.
Sowerby's Thesaurus Conchyliorum is the
only monograph to illustrate four of the dis-
cussed species in accordance with their origi-
nal descriptions; two of these four are Sowerby
species, but the excellent color plates also
clearly point out the differences between Vexil-
lum taeniatum and V. vittatum.
Tryon's contribution will be covered under
the section on Vexillum vittatum; he considered
all the species in this complex synonymous,
with the exception of V. regina.
Vexillum taeniatum has been recorded
from Madagascar; Bombay; Moluccas; Mindoro,
Masbate, and Cebu in the Philippines; Maurice
Island, and Queensland.
Vexillum vittatum (Swainson, 1821)
(Plate 18, figures 5a, 5b; Plate 19, figure 5)
Type locality: Pacific Ocean
Swainson's original description of this spe-
cies mentions "interstices with slender, crowd-
ed, transverse grooves", but the accompanying
poorly-colored figure of the holotype does not
indicate these, nor are they visible in two out
of three of Reeve's poor figures, in Kiener's,
Kister's, Sowerby's, nor in Dautzenberg's
(1935) excellent illustration, the only actual
photograph of this species to appear in the lit-
erature. All these monographers represent
Vexillum vittatum with continuous spiral sculp-
ture over the high points of the costae.
Kister's stylized.ventral view presents a
recognizable illustration of Vexillum vittatum
and his description is adequate.
Tryon placed Vexillum vittatum in synony-
my with V. taeniatum, along with V. compres-
sum (Sowerby), V. coccineum (Reeve), and V.
taylorianum (Sowerby), copying the obviously
different type figures of all these species but
asserting that they are identical.
Dautzenberg (1935) nearly arrived at the
solution to the problem of separating Vexillum
vittatum from V.taeniatum, saying "if it is true
that these species resemble one another by the
disposition of their pattern and their coloration,
they differ considerably by the form and the
sculpture; M. vittata having more angled
whorls, the latter whor)] larger; the axial folds
are much heavier and more projecting on the
last whorls.'' However, he then included in his
illustration of V. vittatum both that species and
V. taeniatum under the same name, thereby
confusing the issue even more than before. In-
stead of illustrating the species V. vittatum as
he intended, his figures afford a fine compari-
son of V. vittatum (Dautzenberg, 1935, Pl. 4,
fig. 3) with V. taeniatum (ibid., Pl. 4, fig. 4).
On the same plate Dautzenberg also illustrated
V. coccineum (Reeve, 1844) under the question-
able combined name of V. taeniatum coccineum
(Reeve) —a combination difficult to understand
when both names apply to separate, established
species, thereby further complicating the prob-
lem of identifying the typical V. taeniatum.
The basic differences between Vexillum
vittatum and V. taeniatum are few but easily
enough defined if one disregards the similarity
of color and pattern, as may be seen by the ac-
companying black-and-white photographs on
Plate 19, figures 4 and 5. While the adapical
whorls of both species indicate interstitial
striae, in V. vittatum these striae become con-
tinuous on the last whorl (sometimes on the
penultimate whorl as well) and are not inter-
rupted by the axial costae. Vexillum vittatum
is more shouldered and the base is more angu-
late than in V. taeniatum. The closely spaced
adapical costae become more distant with ma-
turity in V. vittatum, reducing in number from
approximately 20 on the antepenultimate whorl
to about six on the last whorl; in V. taeniatum
the number of costae remains more or less
constant on all the whorls. The tendency among
the specimens studied seems to be for the spire
length to be about equal to the last whorl in V.
vittatum, slightly shorter than the last whorl in
V. taeniatum, though this characteristic cannot
be considered definitive in view of the small
number of specimens studied.
The lack of spiral ornament just below the
sutures which is so apparent in the enlarged
photograph (Plate 19, figure 5) should be disre-
garded as a diagnostic character as it is not
constant among the specimens studied, nor is
it as prominent on the photographed specimen
as the photograph indicates.
With the above five species adequately
identified, a sixth group remained which pos-
sessed certain traits common to several of the
others but still did not fit all of the character-
istics of any one of them, nor was it encount-
ered in the literature except as a wrongly la-
belled figure. Since a fairly large sample (34
specimens) of this particular group was avail-
Page 80
able for study, all consistently exhibiting the
same morphological features exclusive of col-
or, this form is considered a separate taxon
and is here described as a new subspecies.
VOLUTACEA
MITRIDAE
Vexillinae
Vexillum RODING, 1798
Vexillum regina (SOWERBY, 1825)
Vexillum regina filiareginae J. CATE, subspec. nov.
(Plate 18, figures 6a, 6b; Plate 19, figure 6;
Plate 20, figures 1a - 10a, 1b - rob)
Shell long, straight, slender, turriculate-
fusiform; spire about as long as the last whorl.
Protoconch lacking; (three to four postnuclear
whorls eroded by acid in the holotype); teleo-
conch of about nine straight- sided oblique abut-
ting whorls; shoulders obtusely rounded. Axial
sculpture of raised equidistant collabral costae
(about 12 on the penultimate whorl) whose ridges
and valleys are of about equal depth, forming a
regular, zigzag pattern when viewed from the
apex; costae not regularly aligned between su-
tures; proportionately the same number of cos-
tae on adapical whorls as on last whorl. Spiral
ornament of low cords, rounded, crenulate, nar-
row and crowded below the sutures, flattened
into wider bands at periphery, again becoming
rounded, narrower and wrinkled at lower part
of neck; all spiral cords separated by smooth,
shallow impressed striae and crossed by faint,
irregularly spaced orthocline rugae. Aperture
straight, siphonal canal slightly recurved; la-
brum thickened, constricted (adapical edge
chipped in holotype); number of labral lirae
variable (usually three to six raised lirae and
several faint abapical lirae present). Parietal
ridge present; columella straight; three to five
oblique columellar folds, greatly diminishing
in strength abapically; peristome continuous,
pseudumbilicus and siphonal fasciole faintly
produced.
Color of holotype white, with four narrow
black spiral bands on last whorl; neck of shell
below abapical band orange (Maerz & Paul Dic-
tionary of Color, Ist Edition, 1930; Plate 12,
J-12). Color of paratypes variable; numbers
1-4 and 10-13 similar to holotype, but neck and
one narrow spiral band sometimes dark orange
to dark gray. Paratypes 5-8 and 14-32 pre-
dominantly orange-brown, with one wide and
one narrow white spiral band on last whorl;
Paratype 9 is black with two white bands. In
all specimens the wide white spiral band is
contiguous to abapical sutures and approxi-
mately six cords wide; the narrower band, about
THE VELIGER
Vol. 4; No. 2
‘half as wide, is always immediately above the
adapical columellar fold adaxially and borders
the upper part of the neck; a central yellow
thread present on wide white band in about half
the specimens. Aperture color white to cream;
color pattern visible through translucent la-
brum.
Animal of the species unknown.
Measurements of the holotype: Height, 66.4
mm.; Greatest diameter, 18.1 mm.; Length of
aperture, 34.9 mm.
The type locality of Vexillum regina filia-
reginae is here designated as Cape Melville,
Balabac, Philippine Islands (7° North Latitude,
117° East Longitude).
The name filiareginae means ''daughter of
the queen", or princess — a name considered
suitable for a species closely related to Vexil-
lum regina. Since V, regina filiareginae has so
long been confused “with the older, more famil-
iar V. regina, it was made a subspecies of this
form rather than of the new species V. colo-
scopulus, to which it is as closely related.
When the preliminary study for this paper
was nearing completion and the photographs of
the first ten specimens listed below had been
taken, an additional shipment of 22 newly col-
lected specimens of Vexillum regina filiareginae
from Balabac was sent to me by Mr. Fernando
Dayrit of Manila. Still later one important ad-
ditional specimen was discovered in the collec-
tion of Mr. and Mrs. John Q. Burch, this shell
having been unavailable earlier. The Burch
specimen is identical with the orange-colored
paratypes from Balabac, particularly resem-
bling Paratype 6 (Plate 20, figures 7a, 7b). Its
locality, however, is reported as Mozambique,
East Africa — approximately 8° of latitude
southward and 77° of longitude westward, or a
distance of roughly 5,500 statute miles ina
southwesterly direction from the type locality.
This locality record would seem to establish a
tentative range for the subspecies from Mo-
zambique to Zamboanga. An even wider range
may come to light as more specimens are re-
ported.
The latter 23 specimens were included in
the study and designated as paratypes, though
they do not appear in the photograph on Plate 20
as they were received too late for processing.
The most variable of the six taxa under
discussion, Vexillum regina filiareginae com-
bines some of the morphological features of V.
coloscopulus and some of those of V. regina
and may be a link between these two Spe cies
[ J. Cave] Plate 20
THE VELIGER, Vol. 4, No. 2
Figure 1 Figure 2 Figure 3 Figure 4
Holotype Paratype 1 Paratype 2 Paratype 3
Figure 6 Figure 7
Paratype 6
Figure 5
Paratype 5
Paratype 4
Figure 10
Paratype 9
Figure 9
Paratype 8
Figure 8
Paratype 7
Holotype and nine Paratypes of Vexillum regina filiareginae subspec. nov
Pierson, OswaLp, & PIERSON, photo
Vol. 4; No. 2
THE VELIGER
Page 81
although itself possessing enough constant
morphological characters to be considered a
separate taxon. Its greatest variability lies in
its color; except for a slight fluctuation in obe-
sity and a certain amount of flattening-out of
the ribs on the last whorl in some instances
(apparently always in the orange-colored spe-
cimens rather than in the predominantly white
ones), its other features seem well established.
It appears to be the least rare of the six mem-
bers of its complex.
In arrangement of its color pattern, the
new subspecies resembles Vexillum taeniatum
and V. vittatum; the wide white band is contig-
uous to the abapical sutures, bordered in most
instances by narrow black lines instead of wide
black zones as in V. coloscopulus, and fre-
quently (but not always) with a central yellow
thread.
The color ranges from predominantly white
with gray and black areas, as in the holotype
and paratypes 1-4 (Plate 20, figures 1-5) and
10-13, through various shades, amounts, and
combinations of orange; one specimen (para-
type 9, Plate 20, figure 10) exhibits a reversed
arrangement of the holotype's color and is
largely black with white zones. Paratypes 8
(Plate 20, figure 9), 14, 15, 17, 18, and 19 reveal
a tendency toward dark blackish-orange zones
rather than the more sharply defined narrow
black bordering lines of the other specimens;
these are the most closely related to Vexillum
coloscopulus but remain distinct from that spe-
cies because of the much wider white band, the
presence of two white bands on the last whorl,
by the sharper and more frequent axial costae
on the last whorl, and by the placement of the
wide white band adjacent to the abapical su-
tures.
A central yellow thread is present in about
half the specimens studied, while the others
have simply a plain white band without adorn-
ment.
Vexillum r. filiareginae has a slightly more
gradate sutural ramp than V. coloscopulus,
though it is considerably less angled than in V.
regina. Its spiral ornament is smoother than
that of V. regina and consists mostly of flat,
continuous spiral cords similar to those of V.
coloscopulus., The chief distinguishing charac-
teristic of V. r. filiareginae (especially true of
the holotype and paratypes 1, 2, 4, 10-13) is its
regular, closely-spaced axial costae which for
the most part do not diminish as they approach
the labrum and maintain proportionately the
same number on each whorl.
Synonymy:
1844 Mitra melongena Reeve (pars) not La-
marck, 1811. (Conch. Icon., fig. 47c)
1874 Mitra melongena Sowerby (pars) not
Lamarck, 1811. (Thes. Conch., fig. 132)
1882 Mitra melongena Tryon (pars) not La-
marck, 1811. (Man. Conch., Pl. 49, fig. 401)
It is impossible for the present subspecies
to be the same as Mitra melongena Lamarck,
1811, since according to the original descrip-
tion M. melongena is a smaller shell, 38-40mm.
long instead of 55-67 mm.; its color is de-
scribed as similar to M. lyrata Lamarck, 1811,
which is basically a grayish-brown; and most
importantly, Vexillum regina filiareginae has
no resemblance to the illustration cited by La-
marck as the type figure of M. melongena (En-
cyclopédie Méthodique, Pl. 373, fig.9). Reeve
apparently based his conclusions solely on that
part of Lamarck's written description of M.
melongena which refers to a "whitish back-
ground, ornamented with several transverse
zones, some of a very brownish-red, others a
livid tawny color".
Sowerby, and later Tryon, copied Reeve's
erroneous figure, thus compounding the error
and confusing several generations of concholo-
gists; each of these authors illustrated three
obviously different species under the name of
Mitra melongena, only one of which in each
case approaches a similarity to Lamarck's
type figure.
Relative importance of some diagnostic characters
used to separate the species
Some of these morphological features are
more reliable than others in identifying certain
Mitrid species; their relative importance here
is based on observation of the most outstanding
features of all the subgenera and over 300 rec-
Ognized species which are presently available
to me for study.
The relative width and placement of the
pattern-stripes appear to be constant in those
species where they are present, such as Vexil-
lum caffrum (Linnaeus, 1758); Vv. gruneri
(Reeve, 1844); V. plicarium (Linnaeus, 1758); V.
lyratum (Lamarck, 1811); several species of
Imbricaria, Swainsonia, etc. It therefore seems
reasonable to assume that this character will
be a constant factor in the species discussed
here also, and it is one of the bases for sepa-
rating V.coloscopulus from the other members
of the complex.
Page 82
THE VELIGER
Vol. 4; No. 2
Table showing measurements (in millimeters) of the holotype and paratypes, the collecting localities and
present distribution of these specimens
Vexillum regina filiareginae J. CATE, subspec. nov.
Length of
Length | Width | Aperture
Locality
Holotype 66.4 18.1 34.9 | Balabac
Paratype 1 64.6 15.4 33.3 | Balabac
Paratype 2 55-4 15.0 29.7 | Balabac
Paratype 3 56.2 15-4 29.8 | Balabac
Paratype 4 62.7 16.3 30.7 | Balabac
Paratype 5 62.3 16.7 32.5 | Laminusa
Paratype 6 67.4 17.5 34.6 | Zamboanga
Paratype 7 65.2 15-4 31.5 | Zamboanga
Paratype 8 57.0 15.0 29.0 | Balabac
Paratypeg | 62.9 | 16.9 31.6 | Balabac
Paratype 10 | 54.9 14.6 29.6 | Balabac
Paratype 11 | 60.8 15.8 30.9 | Balabac
Paratype 12 | 57.4 14.1 27.8 | Balabac
Paratype 13 | 60.2 14.9 29.9 | Balabac
Paratype 14 | 57.3 14.2 29.4 | Balabac
Paratype 15 | 60.7 14.8 30.1 Balabac
Paratype 16 | 57.0 13.8 28.3 | Balabac
Paratype 17 | 64.3 15.6 31.6 | Balabac
Paratype 18 | 67.0 17.3 32.6 | Balabac
Paratype 19 | 71.5 17.9 34.6 |Balabac
Paratype 20 | 70.8 17.6 36.0 |Balabac
Paratype 21 | 66.9 15.9 32.7 |Mozambique
Paratypes 22 to 26 Balabac
Paratypes 27 to 28 Siasi, Sulu
Paratypes 29 to 32 (juveniles ) Balabac
Stanford Univ. Paleo. Dept. Type Coll., No. 8623
Calif. Acad. Sci. Geol. Dept. Type Coll., No. 12 366
C. N. Cate Collection
A. d’Attilio Collection
F. Dayrit Collection
C. N. Cate Collection
A. d’Attilio Collection
A. d’Attilio Collection
F. Dayrit Collection
C. N. Cate Collection
F. Dayrit Collection
F. Dayrit Collection
F. Dayrit Collection
F. Dayrit Collection
C. N. Cate Collection
‘|G. N. Cate Collection
C. N. Cate Collection
C. N. Cate Collection
GC. N. Cate Collection
F. Dayrit Collection
F. Dayrit Collection
J. & R. Burch Collection
C. N. Cate Collection
C. N. Cate Collection
C. N. Cate Collection
The following paratypes will be deposited as indicated:
Paratype 22
Paratype 23
Paratype 24
The relative amounts and various kinds of
surface ornament and the angle of the sutural
ramp are also reliable characters for species
separation.
The two least reliable diagnostic charac-
ters are the number of columellar folds and the
color of the shell. The columellar fold-count,
long supposed to be a valid reference point for
Mitrid species, is not a constant factor. When
large series of shells of the same species are
examined, the number of folds is seen to vary
from half a fold to several folds; therefore, this
character should be used only on the generic or
subgeneric level to indicate a general trend.
Among species and subspecies it is not suffi-
ciently constant to be relied upon. Also, since
the strength of these folds may vary somewhat
from one specimen to the next, it has been de-
monstrated that the count will be interpreted
U. S. National Museum
Museum of Comparative Zoology
British Museum (Natural History)
differently by different workers. A weak an-
terior fold may be counted by some, disregard-
ed entirely by others.
Color is an extremely variable feature
among the Mitridae and must be one of the last
physical characters to be considered in identi-
fying a species. This is more true of some
species than others, but until one knows which
ones may be relied upon to be constant, color
should be avoided as a diagnostic feature.
Conclusions
It is my belief that the six forms discussed
in this paper are separate, well-defined taxa,
heretofore confused through the similarity of
color and of pattern arrangement, with Vexil-
lum taeniatum and V. vittatum the most closely
‘Table 1
Chart Showing Distinguishing Characters of Vexellum regina (SOWERBY, 1825)
and Related Forms
Diagnostic Characters
Spire
Shell shape
Labrum
Labral lirae
Siphonal canal
Columellar folds
Pseudumbilicus
Axial costae
Sutural ramp
Parietal lip
Spiral ornament
Spiral sculpture
Neck of shell
Adult shell length
Peristome
Shell color
White bands
White band placement
White band bordered by:
White band ornamentation
Aperture color
V. regina
longer than last whorl
slender turriculate
constricted
about 3, raised
sharply recurved
4
faintly produced
extremely coarse, elevated,
nodulose
angular, gradate
well defined
coarse, rough
continuous
roughly granulate
long: 75 mm.
entire
bright orange, or blackish
one wide, one narrow
contiguous to abapical sutures
narrow black lines
central yellow thread
white
V. 1. filiareginae
about as long as last whorl
slender fusiform, sides straight
constricted
variable; usually 3-6 raised lirae
and several faint abapical lirae
slightly recurved
3 to5
usually faintly produced
widely spaced, well defined,
smooth
moderately angled
variable
smooth
continuous
nearly smooth
57 to 68 mm.
entire
variable
one or two
contiguous to abapical sutures
narrow black lines
variable; some with yellow
thread, some without
white to cream
longer than last whorl
straight
about 12 faint lirae
slightly recurved
4
lacking
smooth, somewhat flattened
rounded
lacking
smooth
continuous
nearly smooth
long: 55 to 80 mm.
discontinuous
brick-red
one narrow band only
central on adapical whorls
wide black zones
lacking
cream to dark ivory
slender fusiform, sides convex
V. coloscopulus V. compressum V. taentatum V. vittatum
shorter than last whorl
slender fusiform - turriculate
constricted
12 to 15 raised lirae
sharply recurved
4
faintly produced
coarse, elevated, nodulose
rounded
moderately well defined
coarse
continuous
roughly granose
short: 48 mm.
discontinuous
orange
one
contiguous to abapical sutures
narrow black lines
yellow thread present
white
shorter than last whorl
obese fusiform, sides convex
constricted
about 12 raised lirae
very slightly recurved
4to5
very faintly produced
close, faintly raised, fewer on last
whorl than on spire
smoothly sloping, hardly defined
variable
smooth
transverse interstitial striae
wrinkled
65 to 70 mm.
entire
orange
one white, one pale orange
contiguous to abapical sutures
narrow black lines
yellow thread present
white
shorter than last whorl
slender fusiform, sides angled
constricted
more widely spaced, strong,
about 6 to 9 lirae
nearly straight
4to5
faintly produced
few, heavy, more projecting
slightly angled
variable
smooth
continuous
wrinkled
45 to 65 mm.
entire
orange
one white, one pale orange
contiguous to abapical sutures
narrow black lines
yellow thread present
white
Vol. 4; No. 2
THE VELIGER
Page 83
related and most difficult tosseparate. The re-
lative sizes of adult specimens and differences
in spiral ornament separate V. regina from V.
compressum; surface ornament and pattern
placement define V. coloscopulus; axial costae
differences make Vv. r. filiareginae distinct;
transverse interstitial striae set V. taeniatum
apart; and V. vittatum is characterized by a
shorter spire, wider shoulders, and a less at-
tenuate last whorl.
Using a goniometer, the spire angles of all
six forms were measured, as it was hoped an
additional basis for separating them could be
established in this way. The imbalance in num-
bers of specimens of each species makes such
a study somewhat ineffective, as there were
available at the time the measurements were
made only two specimens of Vexillum com-
pressum and V. coloscopulus, respectively;
four each of V. taeniatum and V. vittatum; five
of V. regina; and 29 of V. r. filiareginae, the
others having been returned to their owners.
A comparison of such figures would be incon-
clusive except as a general indication of the
trend. However, as shown in the accompany—
ing bar diagram (textfigure 1), it will be seen
that while the angle measurements overlapped
in some of the species, the mean spire angle of
each is relatively distinct from the others,
with V. compressum having the most acute-
ly angled spire, and V. vittatum the most ob-
tuse.
No work has been recorded on the anato-
mies or radulae of any of the six species.
For easy identification of the various
forms, I offer the following brief key based on
morphological differences exclusive of color,
and an expanded chart (Table 1) tabulating all
the characteristics of each species.
V. compressum
V. regina
V. coloscopulus
V. filareginge
V. taenatum
V. vittatum
Figure 1:
1, Costae interrupted by interstitial striae;
shell obese . . 7.» taeniatum
Costae not interrupted by interstitial
ales Bacihl piece 5 56 46 5 oo 6G
2. White spiral band wide, contiguous to ab-
ajolcel SUNOS «o oo 6 6 © 0 © of
White spiral band narrow, centered on ad-
apical whorls - + « » coloscopulus
3. Regular, sharply angled costae oO
Widely spaced, infrequent costae. . .
O10 8) 6. & Bip o oo oRBtHeE ON
A eAxialicostae noduloseln i.) 6D
Axial costae smooth . r. filiareginae
5. Adult shell short (48 mm.), sutures round-
Gl Gy io. sh6 - . compressum
Adult shell Tone (75 mm. “Np sutures with
Gradatesrampw jer a eo regina
The rarity of these shells is probably part-
ly responsible for the fact that the confusion in
their identities has not been corrected earlier;
it is seldom, indeed, that all six of these mag-
nificent species may be seen at one time for
comparative work.
I was extremely fortunate in having access
to loaned specimens of those species lacking in
my own collection. For the courtesies extended
in this regard I am indebted to Dr. Myra Keen,
the late Dr. Howard R. Hill, Mr. George Kana-
koff, Mr. Anthony d'Attilio, Mr. and Mrs. John
Q. Burch, Mr. E. W. Ulrich, Mr. James Bailey,
and Mr. Fernando Dayrit. A special word of
appreciation is also directed to Dr. Rudolf
Stohler who translated the references by Kis-
ter. The long hours necessary for the prepara-
tion of this paper would not have been available
without the patience and helpfulness of Craw-
ford Cate in many practical ways, for which I
am deeply grateful.
MIEETELELMEATMETEMEEMENEEMIEEEMEEEEEOEEMEEEEEEEEEEEEEE LS
LLLALLLLLLLAAALMMMAAAALALL ALANNA AANA AAA Ahhh hh
CZLLILLALLAAAAMM LAA ANNA
Bardiagrams showing the relationships among various forms under discussion based upon
measurements of apical angles.
Page 84
THE VELIGER
Vol. 4; No. 2
Numer of Locality
Specimens
Vexillum regina (Sowerby, 1825)
2 Moluccas
Zanzibar
Moluccas
Andaman Islands
China
Madagascar
Philippine Islands
Moluccas
Madagascar
Madagascar
Philippine Islands
=e & oO eS NY Be & eS
Vexillum compressum (Sowerby, 1874)
1 Lubang, Mindoro, P. I.
2 Davao, Philippines
1 Philippine Islands —
2 Moluccas
] locality unknown
1 Subic Bay, Philippines
1 Philippine Islands
Vexillum coloscopulus J. Cate, 1961
1 (type) Balabac, Philippines
1 Sulu Archipelago
1 Zamboanga, P. I.
1 Philippine Islands
Vexillum taeniatum (Lamarck, 1811)
1 Cooktown, Australia
1 Cooktown, Australia
1 North Queensland
1 Fort Douglas, Australia
2 Gubbins Reef, Queensland
2 Queen's Beach, Bowen,
Queensland
Queensland
—J
Vexillum vittatum (Swainson, 1821)
1 _ Cooktown, Australia
1 North Queensland
2 Gubbins Reef, Queensland
1 Masbate
10 Moluccas
Vexillum r. filiareginae J. Cate, subspec. nov.
2 Zamboanga, P. I.
8 Balabac, Philippines
1 Balabac, Philippines
18 Balabac, Philippines
3 Laminusa, Siasi, Sulu
1 Mozambique, East Africa
1 Philippine Islands
In the Collection of
Stanford University, No. 1'660
Anthony d'Attilio, Valley Stream, New York
Los Angeles County Museum, No. A2'777
E. W. Ulrich, Long Beach, California
James Bailey, Los Angeles, California
C. N. Cate, Los Angeles, California
American Museum of Natural History
American Museum of Natural History
American Museum of Natural History
Academy of Natural Sciences of Philadelphia
EF. K. Hadley, West Newton, Massachusetts
George P. Kanakoff, Los Angeles, California
Howard R. Hill, Los Angeles, California
American Museum of Natural History
American Museum of Natural History
American Museum of Natural History
Academy of Natural Sciences of Philadelphia
F. K. Hadley
Calif. Acad. Sciences, Geol. Dept. Type Coll.
No. 12'363, San Francisco, California
Mr. and Mrs. John Q. Burch, Los Angeles
Anthony d'Attilio
F. K. Hadley
E. W. Ulrich
James Bailey
Howard R. Hill
C. N. Cate
C. N. Cate
Mr. and Mrs. John Q. Burch
F. K. Hadley
E. W. Ulrich
Cc. N. Cate
Cc. N. Cate
American Museum of Natural History
American Museum of Natural History
Anthony d'Attilio
Fernando G. Dayrit, Manila, Philippines
Anthony d'Attilio
Cc. N. Cate
Cc. N. Cate
Mr. and Mrs. John Q. Burch
F. K. Hadley
Vol. 4; No. 2
THE VELIGER
Page 85
ee ee ee ae ——————————— ase
The photographs of Vexillum regina (Plate
18, figures la, lb) were furnished through the
courtesy of Stanford University. The remain-
ing photographs are by Pierson, Oswald and
Pierson.
An unexpected trip east after this manu-
script had been completed made it possible to
visit the collections of Mr. and Mrs. F. K.
Hadley, the American Museum of Natural His-
tory, and the Academy of Natural Sciences of
Philadelphia. All specimens of the six dis-
cussed species in these collections were mea-
sured and recorded as part of the study, and I
am grateful to Dr. William K. Emerson, Mr.
William E. Old, Jr., Virginia Orr, and Mr. and
Mrs. Hadley for their kindness in providing
this opportunity.
Literature Cited
Cate, Jean M.
1961. A new Vexillum (Mitridae) from the Philippine
Islands. Veliger 4 (1): 4-8, pls. 1, 2, 1 textfig.
Chenus, J. C.
1859. Manuel de Conchyliologie. I: 1-507. Paris.
Dautzenberg, Ph.
1935. Résultats scientifiques du voyage aux Indes Ori-
entales Néerlandaises. 2(17) Gastéropodes marins.
2. Famille Mitridae 44-208, pls. 2-4.
Dautzenberg, Ph., & L.-J. Bouge
1922. Mitridés de la Nouvelle Calédonie et de ses dé-
pendances. Jour. Conchyl. 67: Famille Mitridae,
83-259, pl. Il.
ene ge
1933.
ments frangais de l'Océanie.
Famille Mitridae, 152-199.
Encyclopédie Méthodique
Les mollusques testacées marins des établisse-
Jour. Conchyl. 77:
1797. Vers, coquilles, mollusques et zoophytes. pls.
189-390.
Kiener, L. C.
1839. Iconographie des coquilles vivantes. Vol. 4,
Mitra. pp. 1-120, pls. 1-34, Paris.
Kuster, H. C.
1841. Die Familie der Walzenschnecken (Volutacea
Menke). Nurnberg. Mitra: pp. 33-150, pls. 7-17e.
Lamarck, J.-B. P.
1811. Annales du Muséum d'Histoire Naturelle, vol. 17.
Mitra: 195-222. Paris.
Reeve, Lovell Augustus
1844-1845. Conchologia Iconica.
genus Mitra. pls. 1-39.
Sowerby, George Brettingham
1874. Thesaurus conchyliorum. Vol. 4, Monograph of
the genus Mitra: 1-46, pls. 1-28.
Sowerby, James, & George Brettingham Sowerby
Monograph of the
1825. The genera of recent and fossil shells. Vol. II:
159-279. London.
Swainson, William
1822. Zoological illustrations, vol. 2, pls. 9- 119.
London.
Tryon, George W., Jr.
1882. Manual of conchology, 4(Mitridae): 106-200; pls.
32-58. Philadelphia.
Observations on the Biology of Hermaeina smithi, a Sacoglossan
Opisthobranch from the West Coast of North America
by
J. J. GONoR
Department of Zoology and Friday Harbor Marine Laboratory, University of Washington
(13 Textfigures)
Introduction
This work concerns a member of the Opis-
thobranch fauna of the west coast of North
America, Hermaeina smithi Marcus 1961 (Or-
der Sacoglossa), probably the most common
sacoglossan of this coast and certainly widely
distributed. The Sacoglossa are very poorly
known on the coasts of North America, and the
major purpose of this study is to present vari-
ous aspects of the biology of this common form
which aid in understanding the principal fea-
tures of its organization and its relationships
with the habitat. The species was described
from a single specimen, so that it is also in-
tended to serve as a description of the salient
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THE VELIGER
Vol. 4; No. 2
features of the species. Moreover, in none of
the ten described species in the genus has the
internal anatomy been studied adequately. It is
hoped that this description will enable the place
of the genus within the sacoglossa to be evalu-
ated at a later time when adequate information
is available about other members of the very
difficult Stiliger-Hermaea group.
This study began as a comparison between
the feeding biology of this typical form and the
more specialized Olea hansineensis Agersborg,
but it soon became evident that more informa-
tion had been gathered than is usually available
about members of this poorly known group so
that it was considered worthwhile to present
this information separately. A similar study of
Olea is being prepared.
I would like to gratefully acknowledge the
support given to me by the Department of Zool-
ogy and the Friday Harbor Laboratories of the
University of Washington. Part of this work
was done during the tenure of a National Sci-
ence Foundation graduate fellowship. The kind-
ness of Joan E. Steinberg, James R. Lance, and
W. Patrick Milburn made the examination of
material from California localities possible.
Habitat
Hermaeina smithi Marcus, 1961, has been
repeatedly found intertidally on San Juan Island,
Washington, in Argyll Lagoon and in Garrison,
Wescott, and Mitchell Bays. These bays are
very similar and the first two share a common
mouth, while the third is located near the other
two. Both these bays and the smaller Lagoon
are quiet, shallow mud-bottomed embayments,
with extensive meadows of Zostera from the
lowest intertidal level toward the center of the
bays, and with bare mud flats in the higher in-
tertidal zones. In summer the mud flats are
often partially covered by patches of green al-
gae, principally Ulva and Enteromorpha. It is
in and near such algal mats at approximately 0
tide level that this sacoglossan may be found,
at times in large numbers.
In California Hermaeina smithi has been
found intertidally in upper Newport Bay by W.
P. Milburn; in Tomales Bay (type locality;
Marcus, 1961) and Bodega Bay by J. E. Stein-
berg. These bays generally resemble the mud-
dy bays of San Juan Island. However, it has
also been collected by J. R. Lance from small
rocky pools among large boulders on the ex-
posed beach at La Jolla, California, and by
Miss Steinberg from high pools at Duxbury
Reef, Bolinas, California, so that it cannot be
said to be restricted in habitat entirely to the
muddy bays of the Pacific coast. It has not
been seen in dredgings in the San Juan Islands.
Adults and great numbers of egg masses
have been found in the Enteromorpha tangles on
San Juan Island from early April through Sep-
tember from 1954 through 1959. The animals
are easily maintained in the laboratory on sec-
tions of the Enteromorpha mat. Examination
demonstrated that the masses of Enteromorpha
had finer filaments intermingled with the
strands of the principal alga. These were iden-
tified as Rhizoclonium and Urospora spp.
Tests were undertaken to determine the food
species of the opisthobranch. Animals were
removed from the algal mat and kept in clean
dishes with running water for two days. After
this period of starvation the animals were in
good condition, actively moving about and mak-
ing "seeking" motions with the head. When
placed in dishes containing algae, these animals
were quickly attracted to the filaments. Feeding
began immediately on Rhizoclonium and Uro-
spora (both Cladophorales-Cladophoraceae) but
Enteromorpha (Ulvales-Ulvaceae) was never
fed upon, even after prolonged starvation. In-
dividuals that had rejected Enteromorpha read-
ily fed upon Rhizoclonium. Both Urospora and
Rhizoclonium have a thallus of slender, septate
filaments, and the cells of the Urospora spe-
cies used are of greater diameter than those of
the Rhizoclonium. The tubular, multicellular
thallus of Enteromorpha varies in diameter,
with some strands of the same width as those
of Rhizoclonium. The effect of filament diam-
eter on food selection was investigated. Equal
numbers of filaments of Enteromorpha and
Rhizoclonium of the same width were selected,
washed, and placed together in a container with
circulating sea water. Starved animals were
added, and it was observed that the sacoglos-
sans quickly ''examined"' the filaments with the
oral lobes and mouth region (never the rhino-
phores) and selected the Rhizoclonium for feed-
ing. Microscope observations revealed that the
animals would "examine" the filaments of En-
teromorpha and even occasionally clasp them,
just as they would the filaments of Rhizocloni-
um but would always quickly release the En-
teromorpha and wander away. Feeding is de-
scribed later. [Mr. J. R. Lance has observed
this species feeding on Chaetomorpha sp. (Cla-
dophoraceae) in southern California (personal
communication). }
Vol. 4; No. 2
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Page 87
Habitus
This species is a slender form, which va-
ries in size at maturity, but most individuals
are about 10 mm. long, with some attaining 15
mm. Figure 1 depicts a living animal crawling
on a flat surface. The rhinophores are held out
laterally at a slight angle to the substrate. The
anterior margins of the foot are extended on the
surface and can be seen dorsally. The plump
cerata are held curved dorsally over the back,
at times interdigitating, but when the animal is
disturbed they are extended out straight, giving
a "bushy" appearance. To the unaided eye most
individuals appear uniformly black except for
the whitish rhinophore and cerata tips. The in-
tensity of pigmentation of all parts of the body
varies greatly in the local populations, and ex-
amination of large numbers of individuals re-
veals much variation. Some individuals are in-
tensely pigmented so that even the white areas
appear dark grey. Other individuals are so
lightly pigmented that the animals appear
brownish or even yellowish due to the color of
the gonad. In many of the very lightly pigment-
ed animals the usual pigment pattern becomes
further obscured since the cream-colored
areas do not contrast strongly with the darkly
pigmented areas, just as it is obscured in dark
forms by the light areas being invaded by dark
pigment. The pattern described here is that of
individuals in which the contrast is most. dis-
tinct.
External Morphology
The cerata are not scattered but arise
from two well-defined dorso-lateral zones.
They are roughly arranged in obliquely trans-
verse rows, with the smaller cerata located
anteriorly, ventro-laterally, and posteriorly, a
typical arrangement in this Opisthobranch
group. The cerata number increases with size
of individual, but the usual number is from 8 to
15 oblique rows, with two, three, or even four
cerata in each row. A 10 mm. long mature
specimen may have only about 16 cerata on each
side, while a large specimen may have about
40, including many very small ones. In life the
cerata are spindle shaped, abruptly widening
close to the base and tapering distally toward
the blunt point (figure 7). The side held against
the back and against other cerata is somewhat
flattened so that the larger cerata, about 2 to
25 mm. long, are ovoid in cross section and
may be somewhat flattened toward the ends. In
preserved material the cerata seldom retain
life-like proportions; they usually change toa
slender, evenly tapering form, circular in
In life the cerata are contrac-
tile, with well-developed longitudinal muscle
strands arranged in a loose cylinder around the
digestive diverticulum (see figure 7B).
cross section.
The cerata tips are light in color, usually
creamy-white while the lateral cerata surface,
that usually exposed to view, is black. Lightly
pigmented cerata show minute white dots, es-
pecially clustered at the tip (figure7 B); these
can be identified in sections as large mucus
gland cells. The proximal half of the inner,
hidden surface (figure 7A) is creamy-white and
shows the dendritic pattern of vessels as de-
scribed for Hermaeina orientalis, H. nigra, and
H. toyamana by Baba (1949, 1959). These veins
are not easily seen in preserved material and
may also be difficult to detect in many lightly
pigmented live animals.
Other structures visible on the dorsal sur-
face are the very slightly elevated anus, locat-
ed just to the right of midline, near the second
and third cerata, a black spot near it, and the
renal aperture. In intensely pigmented individ-
uals, the back is solid black, and these struc-
tures cannot readily be found. In many animals
the black pigment of the back is broken up by
irregular unpigmented streaks, and in these, a
clear area is usually present around the anus
and the renal pore. The black spot seen near
the anus is the location, just under the surface,
of a small vesicle which has an intensely pig-
mented wall and a short duct opening to the
surface well away from the anus so that this
structure could not contribute a solidifying se-
cretion to the feces. The renal aperture is
tiny but can often be located in the center of a
clear spot to the right of the heart area at the
level of the fifth or sixth cerata row (figure 1).
The foot is prolonged posteriorly so that it
extends behind as a slender tail (figure 1). It
is widest anteriorly, and this end often shows
slight bilobation in a crawling animal (figure 2)
(see description of feeding). The anterior end
of the foot projects forward under the head and
between this anterior margin of the foot and the
head above is a shallow median pit. Into this
pit above the end of the foot opens a large num-
ber of small whitish foot glands (figure 4 fg).
In the living animal the foot is not sharply set
off from the sides by any groove or ridge but
rather by color. The foot is much lighter than
the sides. The foot margin is often a pure
cream color, while the center of the sole is
usually streaked and marbled by black pigment
Page 88
THE VELIGER
Vol. 4; No. 2
(figure 2). This is quite variable; in a few in-
dividuals the sole was virtually pure cream
color, but in most there is some streaking
vaguely arranged into two parallel longitudinal
bands down the foot reminiscent of the bold
pattern on the foot of Hermaeina minor (Baba,
1959).
The non-pigmented areas of the head pro-
duce a characteristic "masked" pattern. Dor-
sally, the eyes are surrounded by ovoid clear
areas, and this zone extends anteriorly up along
the lateral edges of the rhinophores (set off by
dotted lines in figure 1). Paralleling this is a
similar non-pigmented strip on the anterior
rhinophore edge, leaving a black strip running
up the middle. This black strip widens across
the width of the rhinophore near the tip, which
is cream colored. The trough-like underside
of the rhinophore is not pigmented. The width
of the rhinophore is constant almost to the
rounded tip.
The rhinophores are auriculate, with both
the median and the lateral borders rolled in
slightly toward the middle. These margins are
mobile and may be unrolled and the flat surface
presented anteriorly. The antero-median bor-
Figure 1: Dorsal view of living animal, crawling
Ventral view of head of living animal
(black pigment shaded)
Figure 2:
rl - rhinophoral lobe
ol - oral lobe
der is extended as a rhinophoral lobe (figure 2,
r 1) about one-third of its length from the
base. This extension is folded across the rhi-
nophore and extends past the postero-lateral
border so that it is visible dorsally (see figure
1).
The mouth is directed downward on the
ventral surface of the head. It is flanked by
two highly mobile lips, or oral lobes (figure 2,
or 1), which extend tab-like from near the rhi-
nophore bases downward. These oral lobes
are usually extended downward as in figure 2
and are repeatedly touched to the substrate as
the animal crawls. The lobes are extended
directly anterior and their median edges in-
curved in feeding (figure 3). The sides of the
lobes are cream colored, usually with a median
black streak down their length. Their ventral
edges and part of their anterior faces are also
cream colored, while their posterior faces are
black. The ventral surface of the head thus
displays an unpigmented pattern extending onto
the rhinophore lobes, around the mouth, and on
the oral lobes, which is indicated in figure 2.
Feeding
The animals crawl onto an algal filament
and, grasping it with the anterior margin of the
foot and the oral lobes, begin to feed (figure 3).
No sticky secretion, such as observed by Fret-
ter (1941) in feeding Elysia, was seen. The
oral lobes are extended anteriorly and their in-
ner margins clasped across the filament. The
Vol. 4; No. 2
THE VELIGER
Page 89
Ventral view of head of feeding animal
(note slits in cells)
Figure 3:
anterior margin of the foot is extended in a bi-
lobate fashion and these lobes also grasp the
filament. This holds the filament securely and
bends it in toward the mouth. An algal cell is
slit with the radula and the contents quickly
evacuated; the animal then moves to the next
cell of the filament. The action of the radula
could be seen in animals feeding on Rhizoclo-
nium. The radula is extended outward and
back, pressed against the cell, then pulled in-
ward and forward, producing a fine slit (see
cells in figure 3) in the cell wall. Observation
of details was difficult, but it appeared that
only one tooth was used in the cutting action.
This general mode of feeding is the typical
sacoglossan method and has been previously
described by Rao (1937) for Stiliger gopalai, by
Fretter (1941) and MacNae (1954) for Elysia,
by Gascoigne (1956) for Limapontiids, and by
Gonor (1961) for Lobiger serradifalci. In addi-
tion, both MacNae and Gascoigne observed a
grasping action of the foot margin and the lat-
eral lips similar to that described here.
Internal Anatomy
Features of external form and color serve
to differentiate species of Sacoglossa, although
sometimes imperfectly, but are of little aid in
grouping species or understanding relations
within the group. The major features of inter-
nal anatomy are given here as an effort toward
these ends. Both dissections and serial sec-
tions were examined, but histology will not be
given except where necessary.
Figure 4, a ventral view of a dissection,
indicates the topographic relations of the major
internal structures. In this figure the gonad is
displaced to the right and many of the fine tu-
bules of the albumen gland are removed.
The slit-like mouth was described above.
In dissections the stout buccal bulb (figures 4,
5, and 6) can be seen to be surrounded at its
oral end by a mass of small bulbular multicel-
lular buccal glands (figures 4, 5, b g) with
short necks that open ventrally into the oral
cavity. The buccal bulb is lined inside by a
smooth cuticle. The muscle of its wall is not
striated as is that of Hermaea dendridica
(Fretter, 1941). From the side and ventral view
the well-developed ascus (figure 5, as, figure
4) can be seen to be almost as large as the buc-
cal mass proper. The posterior edge of the
ascus is keel-like so that the posterior end ap-
pears pointed. There are a pair of stomato-
al g
st
Figure 4: Dissection, ventral view
alg - albumen gland sg - salivary gland
es - esophagus fg - foot glands
in - intestine bg - buccal glands
st - stomach dd - digestive diverticulum in a
hd - hermaphroditic duct c - ceras
g - gonad p-g - pre-glandular portion
of digestive diverticulum
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THE VELIGER
Vol. 4; No. 2
Figure 5 Figure 6
Figure 5: Lateral view of Buccal bulb
as - ascus
sg g - stomatogastric ganglion
es - esophagus
Figure 6: Dorsal view of Buccal bulb
bg - buccal glands
sa - salivary ampulla
sd -_ salivary duct
gastric ganglia (figure 5, sg g) closely attached
to the posterior and ventral surface of the buc-
cal mass. Two slender salivary glands (figure
4, s g) extend from the posterior end of the
buccal mass, near the esophagus, through the
nerve ring. They widen posteriorly but remain .
strap-like and do not branch. The ciliated sal-
ivary ducts (figure 6, s d) do not directly en-
ter the bulb but on its dorsal surface join the
very short ducts of two small, rounded vesi-
cles, or salivary ampullae (figure 6, s a) with
thin, muscular walls. These salivary ampullae
are apparently reservoirs which can forcibly
eject their contents into the posterior parts of
the buccal bulb. Cyerce elegans has a similar
arrangement of salivary glands and ampullae
(Hoffmann, 1938, p. 1065), and Limapontia has
similar salivary vesicles on the salivary duct
itself (Gascoigne, 1956).
The long and slender esophagus arises
dorsally from the buccal bulb (figures 4, 5, es)
and extends posteriorly for about one-fourth
the length of the animal before joining the Y-
shaped stomach area near the place where the
intestine (figure 4, in) leaves it. About mid-
way along the length of the esophagus is located
a small diverticulum (see figure 4) which ex-
tends antero-dorsally. Its structure is not dif-
ferent from that of the rest of the esophagus.
The stomach area gives rise to two nonglandu-
lar arms (figure 4, p-g, preglandular portion
of digestive glands) extending laterally, each to
bifurcate into an anterior and posterior exten-
sion of the brown digestive gland, running dor-
sally under the cerata rows. The digestive
diverticula (figure 4, d d) come off from these
extensions and run into the cerata (figure 4, c)
where they are moderately lobate or branched
(figure 7B). The diverticula narrow as they
pass through the body wall, and there is pres-
ent here a sphincter reminiscent of that de-
scribed by Marcus (1959) for the same situation
in Hermaeina brattstroemi.
The intestine runs antero-dorsally from
the stomach to open as the anus dorsally, and
just to the right of the midline, posterior to the
level of the first few cerata. The lining epithe-
lium of the intestine is ciliated and thrown into
folds but is nonglandular, unlike that of Her-
maea dendridica (Fretter, 1941).
The older portion of the radular ribbon re-
tains its organization and can be seen coiled in
the ascus, making up to three turns (figure 5,
as). As the ribbon moves into the ascus a cu-
ticular membrane is laid down around it so that
the radula is enclosed within a very thin, tubu-
lar casing, indicated in figure 8C. This casing
does not dissolve in potassium hydroxide and
can be seen in radular preparations, rather
closely applied to the teeth. The figure of the
radula of Phyllobranchopsis enteromorphae
given by Cockerell and Eliot (1905) shows a
similar membrane. The coiling of the old por-
tion of the radula is not simply due to the rib-
Figure 7: Cerata
A Median side, appearanc of surface in life
B Median side, internal structure
Vol. 4; No. 2
THE VELIGER
Page 9!
SSS?
bon being mechanically forced into the confines
of the ascus. Radulae isolated from both fresh
and preserved material by maceration in a
weak detergent solution and freed from all tis-
sue and membranes still exhibit coiling. All
parts of these radulae are very springy and
recoil tightly when straightened with needles.
The coiling seems inherent in the basal ribbon
for the ascending arm of the radula, not yet
coiled in the ascus and held almost straight by
tissue, also bends into a tight curve when freed
from the tissue.
The number of teeth varies with the size of
individual. Most local animals are about 10
mm. long and have tooth numbers ranging from
33 to 49, but a large specimen had a radula of
88 teeth. These numbers include the first four
or five teeth, which are very small and rudi-
mentary, consisting mostly of the flat basal
portion. The maximum length of unused teeth
in the usual radulae varied from 125 to 170
microns.
The base of the radular tooth is rectangu-
lar, with thickened lateral edges and a slightly
excavated surface attached to the basal ribbon.
From the base rises the crown or middle por-
tion of the tooth, which is thickened down the
center. The middle portion bears distally the
rounded tooth tip, a very thin, discoidal lamina
which is the principal cutting portion, judging
from the extensive wear it shows in old teeth.
The middle portion of the tooth bears, on its
leading (cutting) edge, two serrate lamina flank-
ing a longitudinal excavation in the face of the
tooth into which fits a humped keel on the back
of the preceding tooth in the series. The
sharply pointed denticulations of the serrate
edge are directed slightly inward. They show
only occasional wear in old teeth. These den-
ticulations are tallest in the center of the lam-
inae and evenly spaced. Near the middle of the
largest teeth the denticulations are about 2.5 to
3 microns high and spaced about 5 or 6 teeth
per 10 microns of lamina edge, with 2 micron
8paces between their tips. This is twice the
number given by Marcus for denticles in the
type specimen, otherwise his description and
figure is identical to the one here. However, a
radula with 50 teeth, from a 7.5 mm. long (live)
specimen from La Jolla, California, had 3 den-
ticulations per 10 microns of edge at mid-
length of the serrate lamina, in teeth 150 mi-
crons long, corresponding to the figure of Mar-
cus (1961) for the type specimen. A specimen
from the type locality (Tomales Bay) 11 mm.
long (preserved) with a radula of 40 teeth had 3
to 4 denticles per 10 microns length in teeth
170 microns in heighth. Differences between
other details of the teeth of the California ma-
terial and the teeth of the Washington material
are no greater than differences between teeth
in the same radula or between two Washington
individuals. This minute difference is the only
one noted between the morphology of California
specimens and local ones and is not considered
of specific importance.
The predominant feature of the back of the
tooth is a large median humped ridge, slightly
excavated at the sides, with a rounded edge.
Distally, between this ridge and the tooth tip is
a prominent indentation into which fits the tip
of the succeeding tooth. There is a pair of low
laminae running on either side of the large me-
dian keel, which start as faint lines near the
middle of the ridge and rise slowly as they pro-
ceed distally. They are widest on either side
of the indentation in the tooth back and end on
the tooth tip. The denticulate edges of the suc-
ceeding tooth fit closely inside these laminae,
further interlocking the teeth. The close inter-
locking of the teeth and the tension of the basal
ribbon would seem to afford the necessary ri-
gidity to hold the tooth being used in position
while in action.
The cutting tip of the teeth shows marked
effects of wear; the thin, broadly ovoid tip of
unused teeth wears down so that the used teeth
have a truncated end. The teeth within a radu-
la show considerable differences in the amount
of wear to which they have been subjected.
Smaller and medium sized teeth are relatively
more severely worn down than larger ones in
the same radula, and the cutting edge of some
of the smaller teeth may be completely worn
off. Comparison of large and small radulae
reveals the same type of difference between
teeth in analogous positions. Figure 8 demon-
strates the differences between unused (A, Al)
and used (B, Bl) teeth. The difference between
the amount of wear of larger teeth and smaller
teeth of the same radula can be seen by com-
paring A with C, while the comparison of A and
Al indicates the difference between used teeth
of small and large radulae, respectively.
The dusky yellow gonad, the largest inter-
nal structure, lies ventral to the other viscer.
(figure 4, g). The gonad is elongate, extend
ing at maximum development from the far pos-
terior limit of the haemocoele at the base of
the tail anteriorly to more than three fourths
the length of the body. It is composed of many
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Vol. 4; No. 2
A
well separated ovoid lobules which are joined
only by ducts. Each lobule produces both eggs
and sperm. The small collecting ducts from
each lobule join a narrow duct that runs through
the center of the gonad. From the latter duct,
about at the middle of the gonad, runs the com-
mon hermaphroditic duct (figures 4, 9, h d).
The hermaphroditic duct is narrow at its ori-
Figure 8: Teeth from small (A to C)
and large (A1 to B1) radulae
(camera lucida)
A Unused tooth from small radula
B Used tooth, same radula (first preceding tooth in use)
C Used tooth, same radula (9th preceding tooth in use)
A1 Unused tooth from large radula
Bi Used tooth, same radula (first preceding tooth in use)
gin but quickly enlarges, and the tightly coiled
portion is swollen to form a sperm storage or-
gan (ampulla). Shortly before joining the other
reproductive structures, the hermaphroditic
duct again narrows to a fine tube which, like
the ampulla region, has a ciliated epithelium,
but the muscle layer in its wall is thicker than
that in the ampulla. The male portion of the
rest of the genital tract will be described first.
The vas deferens (or efferent duct) (figure 9,
v d) arises from the hermaphroditic duct very
near the common junction of the latter and sev-
eral other structures. The first portion of the
vas deferens is a fine tube leading to a yellow,
multilobulate prostate gland (figure 9, pr g)
which is appended to the vas deferens by a
short duct leading from its cavities. After the
junction of the prostate gland and vas deferens,
the latter is a larger, ciliated tube with muscu-
lar walls. It runs anteriorly to enter the penis
(figure 10, v d) without change in structure.
The male atrium, or eversible penis sac (fi-
gure 10, w p s wall, penis sac) opens to the ex-
terior on the right side, below and just poster-
ior to the level of the eye, as the male aper-
ture. When the penis is extended (figure 9, p)
Figure 9: Genital System (Dissection)
alg - albumen gland prg - prostate gland
ald - albumen duct s - spermatheca
ov - oviduct hd ~- hermaphroditic duct
mg - mucus gland vd -_ vas deferens
aff - afferent spermathecal duct p - penis
eff - efferent spermathecal duct
Vol. 4; No. 2
THE VELIGER
Page 93
it appears as a cone-shaped base surmounted
by the small, unarmed muscular penis proper,
which projects straight out. The penis is a
barb-shaped structure when everted, with a
blunt, tapered tip or penial tentacle (figure 10,
t) and a small, blunt backward projecting arm
bearing the orifice of the vas deferens (figure
10, o v d). When the penis is retracted the
conical penis base is inverted (figure 10,
c p b). When the penis is extended, the penial
tentacle becomes filled with body fluid and dis-
tended straight out (figure 10B). There is a
single female orifice (figure 9, 9) just posterior
to the male aperture, rather than separate va-
ginal and uterine openings. The female orifice
leads into a short, wide female atrium (figure
9, at) which is partially partitioned by folds
into uterine and nidamental portions. The tru-
ly common portion of the atrium is nonglandu-
lar and ciliated. Leading from the anterior
side of the female atrium is the large ciliated
afferent duct (figure 9, aff) of the spermathe-
ca. The spermatheca (figure 9, sp) is a large,
spherical brownish organ and, when filled with
sperm and secretion of the prostate, may be
enormously enlarged and form a bulge in the
right side of the body. The epithelium of the
spermathecal wall is glandular except for a
small area on the postero-ventral portion.
Here it is ciliated and in the center of this area
is located the opening of a short, thin ciliated
duct (figure 9, eff, efferent duct of spermathe-
ca). This duct contains sperm apparently col-
lected from the spermathecal contents. It leads
to the area of junction of the albumen gland
duct (figure 9, al d), the distal oviduct (figure
9, ov), and the hermaphroditic duct (figure 9,
h d). This is probably the site of fertiliza-
tion. This common female section is ciliated
and has a thick muscular wall, which could act
as a valve to regulate the entry of eggs from
the hermaphroditic duct and exclude the ani-
mal's own sperm. The ciliated nonglandular
albumen duct and an efferent oviduct are con-
nected near each other, at the posterior end of
the chamber, opposite to the hermaphroditic
duct. The albumen duct extends posteriorly a
short distance, where it receives at once the
two branches of the albumen gland (figure 9,
al g). The albumen gland is a very extensive
structure (figure 4, al g), apparently basically
composed of two bilateral portions which rami-
fy and branch so extensively that they cannot be
separated except at their junctions with the
common albumen duct. The extensive dendritic
branches of the albumen gland are mostly lo-
cated dorsally and to the sides, under the heart
and kidney but above the other structures. The
Penis and duct
Figure 10:
(total preparation, camera lucida)
A Completely retracted penis
B End of penis, extended state
wps -
cpb -
wall of penis sac (male atrium)
involuted conical penis base
vd -_ vas deferens
ovd ~-_ orifice of vas deferens
t - penial tentacle
fine translucent tan tubules composing the
gland extend into all parts of the haemocoele,
between the organs, even into the head, but do
not enter the cerata or rhinophores.
A large, ciliated oviduct (figure 9, ov)
leaves the common female chamber and runs
anteriorly toward the female atrium on the
right. A short portion of its end connected to
the chamber is muscular, while most of its
length contains mucus gland cells. The oviduct
joins the large, pale yellow mucus gland (figure
9, m g) just posterior to the junction of the
latter with the female atrium. The oviduct
opens into the base of a crease in the wall of
the mucus gland which runs in the median side
of that gland posteriorly to its tip. This crease
partially partitions off a nonglandular ciliated
track of the mucus gland in a manner strikingly
similar to the way the mucus gland of bulloid
tectibranchs is partitioned. Figure 11 depicts
a diagrammatic transverse section of the mu-
cus gland. Apparently eggs are carried in this
ascending groove (figure 11, asc) to the tip of
the gland, and then pass back down anteriorly
THE VELIGER
Vol. 4; No. 2
Figure 11:
mucus gland (mucus cells indicated by shading)
Diagram of transverse section of
asc - ascending ciliated channel
des - descending channel .
through the descending glandular section (figure
11, des), propelled by the heavy ciliation of
the latter region, to emerge from the female
aperture enclosed in the mucus layers of the
egg mass.
Individuals spawn repeatedly in the labora-
tory, attaching the epg masses to any of the al-
gae in the Enteromorpha mat as in the field but
also to the sides of the containers. The egg
masses are elongate, flattened ribbons, usually
about 12 to 20 mm. long, attached by one flat
side and usually curved into a ''C'' shape. They
are about 1.5 mm. wide, the width of about 20
egg capsules and about one-half as deep. The
closely packed eggs are lemon-yellow when
laid, later becoming paler as development pro-
ceeds. They are very much like the egg mass-
es described for many other Sacoglossans (see
Rao, 1937; Rasmussen, 1951; Baba and Hamata-
ni, 1952; and Hamatani, 1960), and for that rea-
son they are not described in detail or figured
here. The larvae emerge as free-swimming
veligers and do not settle in dishes in the lab-
oratory, so apparently have a long planktonic
feeding phase.
The relative size and position of the gang-
lia of the central nervous system is frequently
of use in comparing different species of opis-
thobranchs and in assessing relationships. To
provide information for this purpose, a brief
description of the ganglia will be given here,
omitting all considerations of the nerves aris-
ing from them. The nerves are largely omitted
in figures 12 and 13 for clarity. The terminol-
ogy used is that of Russell (1929).
The cerebropleural ganglia (figure 12, cp)
are ovoid as seen from above, with their long
axes running antero-posteriorly. The pedal
ganglia (figure 12, pe) are also ovoid, but their
long axes are at right angles to those of the
pair above. The cerebropleurals extend anter-
iorly a little further than the pedals, and both
pairs of ganglia are well separated, joined by
distinct commissures so that they lie to either
side rather than directly above and below the
esophagus. The connectives joining the cere-
bropleurals to the pedals are quite short and
broad. The statocysts (figure 13, st) are lo-
cated on the posterior ventral surface of the
pedals and project above that surface. The
stomatogastric ganglia (figure 5, sg g) are not
closely united to the cerebropleurals but joined
to them by a commissure as in Elysia.
On the left side a small sub-intestinal
ganglion (figure 12, sbi) can be seen between
the left cerebropleural and the large abdominal
ganglion, joined to each by a short connective.
The abdominal ganglion (figure 12, ab) is only
slightly smaller than the pedal ganglion. It is
displaced somewhat to the left of midline, On
the right the supraintestinal ganglion (figure 12,
su-i) can be seen to be joined to the right ce-
rebropleural and to the abdominal by short,
equal connectives. The supraintestinal gangli-
on has appended to it a small satellite ganglion
(figure 12, sat) of undetermined homology,
which in part may represent the right parietal
su-i
ab uh
Figure 12: Central nervous system, dorsal view
(total preparation, camera lucida)
cp - cerebropleural ganglion
pe - pedal ganglion
sbi - sub-intestinal ganglion
ab - abdominal ganglion
su-i - supra-intestinal ganglion
sat - satellite ganglion
Vol. 4; No. 2
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Page 95
ganglion of the tectibranchs. It gives rise toa
single small nerve that runs directly posterior
but which could not be traced to its ending.
In general, the central nervous system of
this species is notable for the lack of fusion
and the distinctiveness of the ganglia. Except
for the small satellite on the supraintestinal, it
is remarkably like the nervous system of Ely-
sia viridis described by Russell (1929). It dif-
fers from that of Stiliger gopalai (Rao, 1937,
textfig. 1) and other Stiliger species (Hoffmann,
1936, p. 757) as well as from Hermaea dendri-
tica (Hoffmann, 1936, p. 755, Fig. 530C, from
Vayssiére, 1888), in that the small sub-intesti-
nal ganglion is distinct and not fused into the
abdominal. Apparently this fusion has occurred
independently several times in the Sacoglossan
group, since the seven ganglia are distinct in
otherwise diverse forms such as Elysia viridis
(Russell, 1929), Cyerce iheringi (= Lobifera
crystallina) (Pelseneer, 1894), Oxynoe olivacea
(Hoffmann, 1936, p. 756), and in the present
case.
The pericardial bulge is not prominent on
the dorsal surface but low and gently rising.
The pericardial cavity and heart are, however,
large, occupying the middle of the dorsal sur-
face starting at a point about one-third of the
body length from the head, just anterior to the
renal pore. The kidney is an extensive flat sac
extending under most of the dorsal surface, ly-
ing just under the dorsal epithelium but not ap-
plied to it. Its walls consist of a simple, non-
folded epithelium. The anterior end of the kid-
ney is at the level where the intestine emerges
from the stomach. It is narrow and mid-dorsal
here; posteriorly it widens, and in front of the
pericardial cavity it extends across the width
Figure 13: Ventral view of central nervous system
(camera lucida)
st - statocyst on pedal ganglion
of the back. It is narrow at the level of the
heart, placed almost completely to the right of
the pericardial cavity but overlaps it slightly.
The renal pore leads from the anterior part of
the narrowed portion. More posteriorly, where
the kidney is closely applied to the side and
part of the dorsal surface of the pericardial
membrane, is found the renopericardial aper-
ture. The kidney is widened and again mid-
dorsal behind the heart. It narrows and ends
just anterior to the tail region. The kidney, in
partially surrounding the heart, resembles that
of Stiliger gopalai (Rao, 1937) but is more ex-
tensive anteriorly than in that species.
Discussion
Ecology: The habitat of this species on the
coast of California and Washington was dis-
cussed earlier. It is becoming firmly estab-
lished that the Sacoglossa are typically re-
stricted, as it has been shown here for Her-
maeina smithi, in habitat and feeding to one or
a few closely related species of algae. MacNae
(1954) has discussed the association of many
Elysia species with Codium (Siphonales); Gas-
coigne (1956) has established experimentally
that two species of Limapontiids are restricted
to Cladophora species, while a third is re-
stricted to Vaucheria (Siphonales); Rao (1937)
found that Stiliger gopalai will feed only on
Chaetomorpha; and Gonor (1961) has reviewed
the association of the Oxynoids with Caulerpa
species (Siphonales). This type of information
can greatly aid faunistic and zoogeographical
investigations and may in time, through clari-
fying ecological specializations, assist in the
understanding of the interrelationships within
the Sacoglossa. For example, knowledge of the
relation of Bosellia mimetica, a cryptic spe-
cies, with the alga Halimeda tuna (Siphonales)
enabled Portmann (1958a, 1958b) to rediscover
this form and greatly extend its known range;
and the association of the Tamanovalvidae with
Caulerpa species has permitted the collection
of this very interesting group in Australia
(Burn, 1960a, 1960b) and Baja California (Smith,
1961).. The constancy of such associations is
borne out by the findings of Marcus and Marcus
(1956) and Hand and Steinberg (1955) which de-
monstrate that Alderia uda and the well known
Alderia modesta occupy identical habitats (mud
flats, on Vaucheria) in widely separated parts
of the range of this genus (Europe, Brazil, Ca-
lifornia, and Washington).
Systematics: Ten species of the genus
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Vol. 4; No. 2
Hermaeina have been described to date; these
are:
1. Hermaeina maculata Trinchese, 1874; from
the west Mediterranean coast of Europe.
2. H.(Phyllobranchopsis) enteromorphae
(Cockerell and Eliot, 1905); from San Pe-
dro, California.
3. H. orientalis Baba, 1949; from Sagami and
Suruga Bays, Japan.
4. H. nigra Baba, 1949; from Sagami and To-
yama Bays, Japan.
5. H. formosae Pruvot-Fol, 1953; from the
Atlantic coast of Morocco, North Africa.
6. H. sinusmensalis MacNae, 1954; from Ta-
ble Bay, near Capetown, South Africa.
7. H. minor Baba, 1959; from Toyama Bay,
Japan.
8. H. toyamana Baba,1959; from Toyama Bay,
Japan.
9. H. brattstroemii Marcus, 1959; from
Northern Chile.
10. H. smithi Marcus, 1961; described from
Tomales Bay, California.
With the possible exception of the fir st,
none of these species has been described in
sufficient detail to be distinguished completely
and with certainty from all of the others, yet
the resemblances are such that at least some
are probably synonymous. All lots of material
examined in this study agreed in the details of
internal anatomy, especially the genital sys-
tem, so these species will be discussed in the
light of the variation shown to exist in the ex-
ternal appearance of Hermaeina smithi. It is
probable that the material described by Eliot
(in Cockerell and Eliot, 1905) was of the species
treated here; this, however, cannot be estab-
lished definitely, and it seems best to regard
this name as a nomen dubium. Eliot carefully
stressed in three places that his four speci-
mens were badly macerated both externally and
internally and that important distinguishing
characters could not be made out with certain-
ty. The only character that might allow iden-
tification is the description that the cerata
were flattened. Their shape was not further
described, and the very poor figure is useless.
The abundant material examined here occa-
sionally showed flattened cerata tips, and in
badly preserved specimens other distortions as
well, and it seems likely that material in the
condition described by Eliot would be likewise
distorted. Moreover, in starved animals the
cerata become more flattened and ovoid as the
digestive diverticula regress. The figure given
by Eliot of the radula is obviously of a Her-
maeina, but it is of the worn portion only and
cannot be used to distinguish the species.
The other species from this Pacific coast,
Hermaeina brattstroemi, was also described
from a single small preserved specimen, and
unfortunately, much of its anatomy is unknown,
since it was sexually immature. It cannot be
completely distinguished from the present ma-
terial on external appearance. Contracted,
preserved specimens of H. smithi often re-
semble the figures Marcus (1959, figs. 21-23,
p. 113) has given, in that the cerata and the la-
bial lobes may be much contracted and the rhi-
nophores very shortened so that their natural
shape is lost; they come to be short and wid-
ened distally, with the rhinophoral lobe quite
obscure. The foot is identical, and the range of
variation in pigment intensity and pattern in-
clude the condition of Marcus' specimen. The
careful drawing of two apparently unused radu-
lar teeth (Marcus, 1959, fig. 27, p. 114) does not
show any features which would distinguish
these teeth from some of the material exam-
ined in this study, and indeed resembles the fi-
gure later provided for H. smithi. However,
Marcus states that the earlier teeth were not in
order in the ascus, a feature not shown by any
of the other known species. Two other features
might distinguish this southern form. Figure
24, p. 113, of Marcus (1959) indicates that the
digestive diverticula only bulge out at regular
intervals. In the present material these struc-
tures varied from showing slight, irregular lo-
bulation in young and starved animals to ex-
tensive, rather regular branching of the type
indicated by fig. 34, Pl. 2 of Marcus, 1961. The
penis described for the specimen of H. bratt-
stroemi consisted of a simple straight tubular
structure. If this organ was completely differ-
entiated in this immature specimen, then it
alone will serve to differentiate the two spe-
cies.
Hermaeina sinusmensalis MacNae was not
sufficiently described to permit it to be dis-
tinguished from other dark species with any
certainty. The description of the radula per-
mits assignment to the genus only; it is pre-
sumed that the structure of the teeth was mis-
interpreted because of the close overlapping
and that they are similar in detail to those of
other species, with two denticulate lamellae on
the cutting face. Unfortunately, the size and
shape of the terminal disc cannot be used alone
as a specific character because it is subject to
wear; indeed, it differs slightly within the un-
used teeth of the same radula in H. smithi,
The material examined is identified as
Hermaeina smithi chiefly on the basis of the
morphology of the penis as given by Marcus
(1961) since this proved to be completely con-
Vol. 4; No. 2
THE VELIGER
Page 97
stant while other characters given proved to be
quite variable or of generic level only. The
form of this structure is usually species-spe-
‘cific in Opisthobranchs and is used here in se-
lecting a name for the material studied even
though the’ structure is not described for most
of the species.
It is difficult to evaluate the four species
described from Japan by Baba, since he pro-
vided no information about internal anatomy.
Fortunately, however, Baba was able to exam-
ine and accurately figure living material. All
four species show certain close resemblances
to the present material, at least indicating that
this genus is a well defined group of closely
related species and further emphasizing the
marine faunal connections between Japan and
the American Pacific Coast (Baba, 1957; Mar-
cus, 1961). These common characters have
been utilized in an attempt to re-define the ge-
neric concept.
The two light-colored species of Baba
seem distinct from Hermaeina smithi. Her-
maeina orientalis has strongly rolled rhino-
phores with flaring tips and no rhinophoral
lobes, thus differing from H. smithi. The
black-tipped cerata seem more pointed as well.
Baba only tentatively separated the other light
form, H. toyamana, from H. orientalis because
of difference in pigment intensity, and The Gifs
likely that these specific names are synonyms.
Because of the highly variable nature of
the black color and the variation in tooth and
cerata number with age and size, it is more
difficult to separate Hermaeina smithi from the
two black Japanese species, H. nigra and H.
minor. According to the very brief description,
H. minor differs from the preceding two in that
it is said to have no ceratal veins and, judging
from the figure (Baba, 1959, figs. 7, 7a), ap-
parently no rhinophoral lobes on the anterior
rhinophore margins. The description of H. ni-
gra greatly resembles the specimens OnE
smithi in external appearance, except that the
three specimens upon which H. nigra was based
were larger (12-30 mm. long) and had more
ceratal rows (25). The figure (Baba, 1949, p.
34, textfig. 23-B) of an apparently unused ra-
dular tooth of H. nigra differs from the appear-
ance of the unused teeth of H. smithi in outline
of the tooth tip and in relative height and may
prove significant if constant.
The original description of Hermaeina for-
mosa is not available to me at present, but ap-
parently this species is similar to H. maculata
in the distinctive color stripes on the cerata
and also in the structure of the radula (Pruvot-
Fol, 1954, p. 190). These purple stripes differ-
entiate these two species from H. smithi and
the other known Pacific forms.
The two earlier attempts of MacNae (1954)
and Pruvot-Fol (1954) at defining the generic
concept uniting the species described under
Hermaeina may be extended now that more in-
formation is available.
Hermaeina Trinchese 1874
Type: Hermaeina maculata Trinchese, 1874
(?=Aplysiopsys elegans Deshayes, 1834-1858)
Rhinophores auriculate, with the anterior
(median) border often prolonged into a rhino-
phoral lobe which overlaps the posterior bor-
der below. Distinct, short oral lobes present.
Anterior free margin of the foot set off by a
groove behind the oral lobes. Cerata fusiform,
ovoid in cross section, usually with venation on
the mesial surface, and branched hepatic di-
verticulum (but no branches of the albumen or
prostate glands) within. Anus anterior near the
first cerata and placed medianly, or slightly to
the right. The radula coiled in the ascus, with
old teeth in order; tooth form (described earli-
er) quite characteristic of the genus, but not at
the specific level. Genital system diaulic, pe-
nis unarmed. Often with black or brown pig-
mentation showing a general pattern, such as
clear areas around the eyes and two streaks
down the foot, which, however, is held in com-
mon with other Styligerids.
Hermaeina may be separated from the ge-
nera of the ''family'' Polybranchiidae (sensus
Pruvot-Fol, 1954) possessing cerata on many
points of internal and external anatomy. The
most important of these are the dorso-median
anus, cerata with hepatic branches, unbranched
rhinophores, lack of a muscular crop ora large
esophageal diverticulum and various details of
the genital tract (compare Caliphylla mediter-
ranea, Marcus, 1958) such as lack of peneal
style and the single spermatheca.
The arrangement of the supraspecific ca-
tegories in the Styligerid group is highly arti-
ficial and remains in a confused state uninflu-
enced by evolutionary concepts or newer taxo-
nomic ideas reflected in the systematics of
better known groups. This can readily be at-
tributed to the lack of good studies of anatomy.
The resulting disorder is well described by
Pruvot-Fol (1954, pp. 180 ff.). This situation
makes systematic comparison of the genera
impossible since they are poorly defined. A
Page 98
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Vol. 4; No. 2
LS
few comparisons have been made in the de-
scriptions of internal anatomy whenever these
have been pertinent. Some remarks, support-
ing the separate status of Hermaeina, about the
critical genital system may be made since
through the careful work of Marcus and Rao
there are available recent accounts of five spe-
cies of Stiliger (s.s.) and one of Hermaea
coirala.
The genital systems of Hermaeina smithi
and Hermaea coirala are similar since both
are diaulic, with an unarmed penis. But Her-
maea coirala has both a spermatocyst and a
spermatheca and the latter structure does not
have separate efferent and afferent ducts as
does Hermaeina smithi. Hermaea dendritica
differs even more in that it is triaulic, pos-
sesses a peneal stylet and has branches of the
albumen gland in the cerata. The reproductive
system of Hermaeina smithi differs from this
system in Stiliger talis, S. fuscatus, and S. va-
nellus mainly in that these species are triaulic,
have peneal stylets, and the albumen gland joins
the mucus gland directly. But in addition, it
differs from S. talis and S. gopalai in the loca-
tion of connections of the female organs.
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1961. Opisthobranch mollusks from California.
Veliger 3 (Suppl., pt. I): 1-84, pls. 1-10.
Marcus, Eveline, & Ernst Marcus
1956. On two sacoglossan slugs from Brazil.
Mus. Novitates No. 1796: 1-21.
Portmann, Adolf
1958a. Uber zwei wenig bekannte Ascoglossa des Mit-
telmeers (Gast. Opisthobranchia). Rev Suisse
Zool. (65 (25): 405-41).
s
1958b. Bosellia mimetica Trinchese, Opisthobranche
retrouvé en Méditerranée. Vie et Milieu 9: 74-80.
Pruvot-Fol, Alice
1953. Etudes de quelques opisthobranches de la c6te
Atlantique de Maroc et du Sénégal. Trav. Inst.
Cherifien 5 (1952): 1-105, pls. 1-3.
Amer.
1954. Mollusques opisthobranches. Faune de France,
58: 460 pp., 173 textfigs., 1 pl. Paul Lechevalier,
Paris.
Rao, K. Virabhadra
1937. Structure, habits and early development of a
new species of Stiliger Ehrenberg. Rec. Indian
Mus. 39: 435-464, pls. 7-9.
Rasmussen, Erik
1951. Faunistic and biological notes on marine inver-
tebrates Il. The eggs and larvae of some Danish
marine gastropods. Vidensk. Medd. Dansk Naturh.
Foren. 113: 201-249.
Russell, Lilian
1929. The comparative morphology of the elysioid and
aeolidioid nervous system. Proc. Zool. Soc. Lon-
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Smith, Allyn G.
1961. Notes on the habitat of Berthelinia sp. nov. from
the vicinity of La Paz, Baja California. Veliger 3
(3): 81-82.
Vol. 4; No. 2
THE VELIGER
Page 99
On Certain Littoral Species of Octolasmis (Cirripedia, Thoracica)
Symbiotic with
Decapod Crustacea from Australia, Hawaii, and Japan
by
WiLutiamM A. NEwMAN
Department of Zoology, University of California, Berkeley 4, California
(Plates 21, 22, and 23)
This paper concerns the reinstatement of
Octolasmis neptuni (MacDonald), long held in
synonymy with O. lowei (Darwin). Australian
and Japanese subspecies are designated. The
paper is also concerned with a remarkable new
species of Octolasmis from Hawaii. An analy-
sis of the ontogeny and adult morphology sug-
gests a new interpretation of the origin and
phylogeny of this and related forms is required.
POECILASMATIDAE NILSSON-CANTELL, 1921
[TRILAsMATIDAE Nisson-CANTELL. 1934]
Octolasmis GRaAy, 1825
Octolasmis PILSBRY. 1907
When Darwin (1851) described the small
pedunculate barnacle Octolasmis lowei from the
gills of a spinous crab from Madeira, he was
quite unaware that numerous comparable forms
were to be found on large crustacea in all warm
seas, and in the depths of the ocean as well.
Had he foreseen this, he probably would have
recognized that the reduction in external arma-
ment of this type of barnacle in response to the
protection afforded by the host might not only
express more than ordinary variability within a
species, but that the trend would achieve com-
parable degrees and form of disarmament in
different species. Had he suspected such
trends, he probably would have stressed ar-
thropodal rather than conchological character-
istics in the description of O. lowei for the fu-
ture was to see a number of otherwise distinct
species confused with it.
Subsequent taxonomists have also tended to
rely on conchological characteristics in their
diagnoses of species in this group, and many
doubtful as well as good species were described
during the five or so decades following Darwin.
During this period of expansion, workers began
to suspect the sorts of variability found in the
group, and many of the previously described
species were placed in synonymy. Octolasmis
lowei acquired ten junior synonyms primarily
due to the efforts of Annandale (1909), Barnard
(1924), and Nilsson-Cantell (1927). In some
cases the synonymies were rather indiscrimi-
nately placed, being based on external appear-
ance alone and were thus not well founded
(Newman, 1960a).
Resistance to this trend was inaugurated in
the brilliant analysis of the ''Octolasmis lowei
complex" by Hiro (1937) in which he clearly
demonstrated that the deep water form, O.
aymonini (Lessona & Tapparone-Canefri), was
distinct from the littoral form, O. lowei sensu
Darwin, and that O. geryonophila Pilsbry, also
from deep water, was probably closely related
to it. He further set order to the complex by
dividing the remaining species into two series:
forma lowei and forma neptuni. However, this
system has not been followed in later system-
atic considerations of the group.
I have been able to confirm the opinion of
Hiro (1937) regarding the close affinities of
Octolasmis geryonophila and O. aymonini; in
fact, it is my opinion that they must be consid-
ered the same species (Newman, 1961), It now
affords me pleasure to announce that not only
is O. lowei forma neptuni distinct from his
forma lowei as he proposed, but that O. nep-
tuni (MacDonald, 1869), in accordance with the
present state of our knowledge, must be con-
sidered a distinct species, as originally de-
scribed. This conclusion has been reached
through a study of MacDonald's form from the
type locality and type host,
A supplementary description of Octolasmis
Page 100
THE VELIGER
Vol. 4; No. 2
neptuni is given here, and Australian and Ja-
Ppanese subspecies are designated.
Octolasmis (Octolasmis) neptunt
(MacDonaLpD, 1869)
Distribution: Indo-West-Pacific; Durban,
South Africa (Barnard, 1924) and Suez (Gruvel,
1905), to Moreton Bay and Sydney, Australia,
and the ? Islands of the South West Pacific
(MacDonald, 1869), to the Seto Inland Sea, Japan
(Hiro, 1937). ?Caribbean (Pearse, 1932). Oc-
curring on gills, occasionally on mouth parts,
of Brachyura, generally portunids.
Diagnosis: Basal segment of scutum nar-
row, needle-like, usually not reaching carinal
fork; tergum broad, semilunar or quadrangular
with small occludent projection (Hiro, 1937).
Supplementary diagnosis: Basal arm of tergum
often chitinous and/or vestigial, situated ap-
proximately one fourth the length of the capitu-
lum above the basal fork of the carina. Surface
of Capitulum and peduncle studded with minute
transparent closely spaced sharp or blunt
spines (not hair-like bristles or amber beads).
Aperture without inner margin of minute denti-
cles. Labrum with 15 to 21 sharp closely
spaced teeth; mandible with fifth tooth conspic-
uous (not rudimentary); penis with apex ob-
liquely truncate, without pronounced apical lan-
guet, supporting terminal comb of relatively
short bristles; clothed sparsely throughout with
soft setae arranged more or less at random or
in ill-defined rows.
Octolasmis (Octolasmis) neptuni neptunt
(MacDonaLp, 1869)
(Plate 21)
1869 Parodolepas neptuni MacDonald: 442
1894 Dichelaspis neptuni (MacDonald), Aurivillius: 19
1900a Dichelaspis vaillanti Gruvel: 2
1900b Dichelaspis vaillanti Gruvel: 109
1900b Dichelaspis neptuni (MacDonald), Gruvel: 110
1905 Dichelaspis peptuni (MacDonald), Gruvel: 127
1905 Dichelaspie vaillanti Gruvel: 128
1906 Dichelaspis vaillanti Gruvel, Annandale: 45
1908 Dichelaspie vaillanti Gruvel, Annandale: pl. 4, fig. 6
1909 Dichelaspis 6inuata in part. Annandale: 12) (non D.
Sinuata Aurivilliue, 1894)
1924 Octolaémis neptuni (MacDonald) in part. Barnard: 60
1927 Octolasmis lowei (Darwin) in part. Nilsson-Cantell:
766
1931 Octolasmis lowei (Darwin) in part. Broch: 129
1932 ?Dichelaspies sinuata Aurivillius, Pearse: ) 10 (identi-
fied by P. J. Vissher)
1937 Octolasmis lowei forma neptuni Hiro: 426
1938 Octolasmis lowei (Darwin) in part. Nileson-Cantell:11
1960a Octolasmis lowei forma neptuni Hiro, Newman: 108
1960b Octolasmis lowei forma neptuni Hiro, Newman: 10
1961 Octolasmis lowei (Darwin) in part. Causey: 5)
Locality: Moreton Bay and Sydney, Australia,
on the gills of Neptunus pelagicus (Linnaeus);
the islands of the South-West Pacific (accord-
ing to MacDonald). Eighteen specimens were
collected from the gills of N. pelagicus from
Moreton Bay, Queensland, Australia, by Dr.
Cadet Hand, March 26, 1960. Five of these
specimens have been sent to Dr. Huzio Utinomi,
Seto Marine Biological Laboratory, Japan.
Neotype: U.S.N.M. Cat. No. 107'011.
Type locality: Moreton Bay; approximately 27°
10'S, Lat., 153° E, Long.
Diagnosis: Capitulum and peduncle trans-
lucent, surface studded with minute sharp
spines (Plate 21, figure 1). Color (in alcohol),
pink. Valves 5 in number; arms not overlap-
ping; basal arm of scutum traversing capitulum
approximately one fourth the length of the ca-
pitulum above the capitulo-peduncular junction;
distal portions of basal arms of scuta and ca-
rina chitinous rather than calcareous. Labrum
with 15 to 21 closely set minute conical teeth.
Rami of Cirrus I equal, each ultimate article
supporting 5 or 6 strong plumose spines and a
few simple shorter setae (Plate 21, figure 7).
Penis with soft setae scattered, or arranged in
linear groups of 2,3, or 4 over the surface;
apex without distinct languet, supporting a few
short blunt spines and a transverse row of
about 7 longer soft setae of about equal length
(Plate 21, figure 9).
Supplementary Description: Capitulum
ovoid or somewhat pear-shaped, laterally com-
pressed, translucent, tinted pink in most spe-
cimens; occludent margin, from basi-scutal
angle to apex, nearly straight; orifice somewhat
flaring, without inner border of small denti-
cles; carinal margin broadly convex (Plate 21,
figures 1,2). Outline of growth increments as
chitinous areas, approximating the original ex-
tent of the valves, clearly visible in young spe-
Explanation of Plate 21
Octolasmis neptuni neptuni (MACDONALD, 1869)
Figure 1: neotype; Figure 2: young specimen;
Figure 3: spines on peduncle;
Figure 6: inner maxilla;
Figures 3 through g: armament and appendages of neotype;
Figure 4: spines on lateral surface of capitulum; Figure 5: mandible;
Figure 7: last three articles of rami of Cirrus I;
Figure 8: caudal appendage;
Figure 9: terminal portion of penis.
Pun V oi 7
THe VELIGER, Vol. 4, No. 2 {NeEwMAn] Plate 21
) }
2 MM
Figure 5
)
O.05¥™
Figure 4
Figure 6
fo)
0.05 MM
Figure 9
Figure 7
Newman, del.
THE ViEvicer, Vol. 4, No. 2 [Newman] Plate 22
Figure 4 Q@___—°3™
Figure 7 Figure 8
Newman, del.
Vol. 4; No. 2
THE VELIGER
Page 101
SS
cimens, becoming less conspicuous with
growth, essentially lost in mature forms except
in tergal region. Surface studded with minute
sharp spines which are transparent, never am-
ber ("minute points or pseudo-cellular struc-
tures'' in MacDonald's account), spines best
seen along capitular margins or along folded
edge of torn mantle (Plate 21, figures 3, 4).
Measurements taken from four specimens
(in millimeters):
Range Mean
Capitulum
Length: 1.72 - 2.25 2.08
Depth: 1.27 - 1.84 MENT)
Width: 0.69 - 1.15 0.97
Peduncle
Length: 0.92 - 1.95 1,32
Av. Diam.: 0.53 - 0.78 0.66
Total Length: 2.87 - 4.13 3.40%
*Three millimeters in MacDonald's account.
Valves 5, much reduced. Scutum of 2
arms; occludent arm calcified, straight or a
little convex, not overlapping tergum; basal
arm reduced, chitinous rather than calcareous
for most of its length (''rudimentary'" according
to MacDonald), traversing capitulum approxi-
mately one fourth the length of the capitulum
above the capitulo-peduncular junction, point-
ing at or somewhat below the middle of the ca-
Tina. Basi-scutal angle acute, ranging between
50° and 76° (mean 64°, from paired measure-
ments of 18 specimens; approximately 58° in
MacDonald's figure). Tergum variable in form;
moon or saddle-shaped; occludent arm reduced,
nearly lacking in some specimens. Carina not
extending up between terga; forked basally; a
small point extending below basi-carinal angle;
basal arms of carinal fork for most part chiti-
nous rather than calcareous, not overlapping
basal arms of scuta.
Peduncle variable in length, depending on
position on host and state of contraction; color
pink, becoming maroon basally (color or tinting
comparable to color of gills and membranes of
host); studded with minute closely spaced stout
sharp spines.
Labrum bullate; bullate portion marked by
fine widely spaced scales over general surface
and a few short soft setae in front of crest
area; crest supporting 15 to 21 rather closely-
spaced small conical teeth (13 visible in Mac-
Donald's figure, rest obscured?). Palps spa-
tulate, supporting numerous long setae along
upper margin and inner angle; lower margin
and surface covered with small ctenoid scales.
Mandible with 5 teeth, third, fourth, and fifth
tooth bifid, second tooth occasionally bifid; fifth
tooth, although smaller than fourth, not rudi-
mentary, Occasionally trifid; inferior angle
cleft, forming 2, sometimes 3, hyaline spines;
superior margin supporting several pairs of
soft spines; inferior margin supporting numer-
ous strong spines (Plate 21, figure 5). Inner
maxilla with cutting edge nearly straight, di-
vided into 2 portions by a small notch; portion
above notch supporting 3 strong spines; notch
with 2 spinules; portion below notch with 6 or 7
spines and a few spinules; superior margin
with numerous pairs of soft spines equally
spaced over at least half the entire length; in-
ferior margin supporting 3 to 5 pairs of stiff
spinules (Plate 21, figure 6). Outer maxilla
rhomboid, corners rounded; superior margin
supporting about 7 slender long curved spines
and a few spinules; inner margin with numer-
ous spinules.
Cirrus I with equal rami; rami one half
length of rami of Cirrus II; outer ramus about
three fourths the width of inner ramus; both
rami clothed with dense soft setae; ultimate
articles supporting 5 or 6 strong plumose
spines and a few soft setae (Plate 21, figure 7).
Cirri II through VI ctenopod, essentially equal
in length and with equal or nearly equal rami;
posterior margins of pedicles and some proxi-
mal articles supporting small distally directed
scales. Inner curvature of intermediate arti-
cles of Cirrus VI supporting 7 pairs of setae,
the number gradually becoming less in more
distal articles; greater curvature supporting
about 5 long setae ina clump at each articula-
tion. Counts for the neotype are given below,
Explanation of Plate 22
Octolasmis indubia NEWMAN, spec. nov.
Figure 1: holotype;
Figures 2 and 3: valves of a paratype;
Figure 2: anterior and dorsal view of carina;
Figure 3: disarticulated valves - lateral view of carina, interior view of tergum and scutum; Figures 4 through 9:
trophi and appendages of holotype;
Figure 4: labrum and palps;
Figure 5: outer maxilla; Figure 6: mandible;
Figure 7: intermediate articles of outer ramus of Cirrus IV; Figure 8: intermediate articles of outer ramus of
Cirrus VI;
Figure 9: caudal appendage and pedicle of Cirrus VI.
Page 102
THE VELIGER
Vol. 4; No. 2
variations noted in other specimens being
placed parenthetically:
II!
Cirrus I Il LV Viet Val
Outer Ramus 6 10 9(10) 9 9 9
6(7) 10(9) 9 CS pate) 9
Inner Ramus
Caudal appendage uniarticulate, nearly as
long as pedicle of Cirrus VI, supporting apical
tuft of about 10 long strong setae, the longest of
which equals the length of the entire appendage
(Plate 21, figure 8).
Penis moderately long, surface supporting
a few scattered soft bristles occurring singly
or in linear groups of 2, 3, or 4; tapering grad-
ually for the first three quarters and then ab -
ruptly to a smaller diameter in distal quarter
of length; basal surface clothed with closely
spaced distally directed scales, central portion
with smooth but irregular surface, distal third
(except apical region) finely folded; apex with-
out distinct terminal languet, supporting a few
short spines followed by a transverse row of
about 7 longer soft obliquely placed bristles of
nearly equal length (Plate 21, figure 9).
Discussion
The species Octolasmis neptuni (Mac-
Donald, 1868), reinstated here, was originally
described from the gills of Neptunus pelagicus
(Linnaeus) from Moreton Bay, Queensland,
Australia. It was placed in synonymy with O.
lowei (Darwin, 1851) by Nilsson-Cantell (1927)
while working on specimens from the gills of a
xanthid crab, Pseudocarcinus gigas (Lamarck),
from Tasmania. It is my opinion that although
Nilsson-Cantell's form is O. lowei s.1., it is
not O. neptuni as he supposed. It does not
compare favorably with the material described
here, which originated from the type locality
and the type host and is clearly O. neptuni as
Originally described.
In addition to the Australian localities,
MacDonald reports and describes a comparable
barnacle from a swimming crab from Fiji.
From the description and accompanying text, it
appears that he is not at all convinced that the
Fijian form is identical with those
Moreton Bay. Although one can only guess, it
is my opinion that MacDonald's Fijian form is
not Octolasmis neptuni, This opinion rests on
MacDonald's own reluctance to confirm the
identity and on the fact that the fifth tooth of the
mandible is rudimentary and the rami of the
seen at
first cirri are grossly unequal. For these rea-
sons the locality records for this species in the
islands of the south west Pacific have been
questioned.
Octolasmis (Octolasmis) neptuni hirot
Newman, subspec. nov.
1937 Octolasmie Jowei forma neptuni in part. Hiro; 426; fig.
13 E-H, fig. 15 A-G.
Locality: Seto Inland Sea, Japan, on gills
and occasionally mouth parts of Neptunus tri-
tuberculatus Miers and on gills of Charybdis
japonica (Milne- Edwards).
Holotype: U.S.N.M. Cat. No. 107'308; Hukuya-
ma, Seto Inland Sea (34° 30'N. Lat.;133° 22' E.
Long.), on gills of C. japonica, kindly sent me
by Dr. Huzio Utinomi.
Diagnosis: The present subspecies is de-
scribed by Hiro (1937) from Japan, and it dif-
fers little from the typical form from Austral-
ia. The differences are considered diagnostic
for the Japanese subspecies and are briefly
described here: Color (in alcohol), opaque
white; mature specimens with minute transpa-
rent closely spaced extremely low blunt spines
clothing capitulum and peduncle. In the speci-
mens on hand the basal arm of the scutum is
for the most part calcareous rather than almost
entirely chitinous as it is in the typical form.
In any event, it ranges from needle-like to
nearly absent, and this is normal for the spe-
cies.
The following description is of an interest-
ing species of Octolasmis from the mouth parts
of a macrurous decapod crustacean from Ha-
waii. The valves, and armature of the mouth
parts and cirri, are quite different from the
aforementioned species. An understanding of
these differences is crucial to an appreciation
of the unique position occupied by this and re-
lated forms. For this reason, a consideration
of their significance follows rather than pre-
ceeds the general description.
Octolasmis (Octolasmis) indubia NEWMAN,
spec. nov.
(Plate 22)
Locality: Kaneohe Bay, Oahu, Hawaii (Co-
conut Island; approximately 21° 27'N, Lat.; 157°
47'W.Long.). Numerous specimens from the
Vol. 4; No. 2
THE VELIGER
Page 103
mouth parts of a single specimen of Scyllarides
squamosus (Milne-Edwards), the gills of which
were infected with Octolasmis lowei (Darwin);
collected by Stephen A. Wainwright.
Holotype: U.S.N.M. Cat. No. 107'310; para-
types: U.S.N.M. Cat. No.107'311, 107'312.
Diagnosis: Capitulum ovoid, laterally flat-
tened, of 5 valves separated from one another
by a narrow border of mantle wall (Plate 22,
figure 1), Carina narrow, extending up between
terga, terminating basally ina bifurcate knob.
Scutum of 2 arms connected by an uncalcified
portion; occludent arm spatulate, extending into
notch of tergum; basal arm broadly expanded,
with internal broad flat platform. Border of
scutum in line with occludent margin of capitu-
lum below primordial valve, forming a broad
curve with the basal margin. Labrum with nu-
merous closely spaced large sharp teeth, palps
relatively small (Plate 22, figure 4). Chaeto-
taxis of Cirrus 1V acanthopod (Plate 22, figure
7). Mandible with 4 sharp teeth, each tooth
with strong basal spines; inferior angle devel-
oped as a sharp tooth, often as large as the
fourth tooth (Plate 22, figure 6). Inner maxilla
with 3 strong spines above and approximately 6
spines below the deep notch, second spine in
lower group strongest (Plate 22, figure 5).
Description: Five valves, appearing as 7
due to incomplete calcification of scuta above
rostral angle (Plate 22, figures 1-3). Carina
narrow, extending up between terga, terminat-
ing proximally in a bifurcate knob, calcification
not extending below primordial valve. Tergum
broad, rounded, tending to form 3 angles; ca-
rinal angle projecting between carina and basal
arm of scutum; median angle extending over
occludent arm of scutum; median and occludent
angles forming notch receiving distal portion
of occludent arm of scutum. In young speci-
mens, terga somewhat rectangular, although
indentation receiving occludent arm of scutum
already developed. With growth, portion be-
tween carinal and median angles of tergum
broadens and the scutal indentation becomes
reduced, Occludent arm of scutum spatulate,
connected to basal arm by a yellowish ligament
overlying primordial valve. About three
fourths of the primordial valve extends onto the
occludent arm of the scutum in large individu-
als. Basal arm of scutum broadly expanded,
rounded; interior portion, near rostral angle,
thickened and elevated as a broad flat platform.
Platforms of scuta are fused, rendering basal
segments rigid, preventing lateral compression
of the animal. (Valves can be readily separated
by corrosion of cementing substance with five
percent sodium hypochlorite.) Ligament al-
lows movement of the occludent arm and per-
mits closure of the aperture. The nearly com-
plete armored condition seen in mature speci-
mens is attained by slow ontogenetic incre-
ments, during which the area occupied by the
valves becomes greater and greater. This
process is quite unusual and its postulated
significance is taken up in the discussion be-
low.
Peduncle: Smooth, without chitinous
spines, knobs, beads, or calcareous inclusions;
length apparently dependent upon postion of
barnacle on host.
Measurements for four individuals (in mil-
limeters):
Capitulum
Height Width Depth
1.2 0.4 0.9
1.4 0.6 1.0
1.8 0.8 1,3
2.0 0.8 1,4
Peduncle
Length Average Diameter
3.5 0.35
2.5 0.3
3.5 0.5
3.5 0.5
Labrum bullate, crest supporting approx-
imately 21 strong sharp teeth. Palps oblong,
sparsely setose along the superior margin and
inner angles; situated well toward lateral mar-
gins of labrum (Plate 22, figure 4). Mandible
with 5 sharp teeth not including inferior angle;
first, second, third, and fourth teeth supporting
2 to4 strong spines at their bases (not pectinate
in the ordinary sense); inferior angle forming a
sharp tooth which may be equal to or slightly
less well developed than the fourth tooth (Plate
22, figure 6). Inner maxilla with 3 strong
spines above and 7 strong spines below deep
notch, second spine in the lower group the
strongest; 3 pairs of setae along upper margin
appear to be a constant characteristic (Plate
22, figure 5). Outer maxilla broadly rounded,
clothed sparsely with short bristles and long
slender spines or setae along the superior and
inner margins.
Cirri relatively short and stout. Cirrus |
with unequal, Cirrus II with subequal, and Cir-
ri III through VI with equal rami. Pedicles of
Cirrus 1 originate at some distance forward of
Cirrus II. Outer ramus of Cirrus I about two-
thirds as long and two-thirds as wide as inner
Page 104
THE VELIGER
Vol. 4; No. 2
a el
ramus; clothed with numerous long relatively
soft setae; terminal article supporting tuft of
about 3 stout spines; inner ramus clothed with
numerous stout setae. Several proximal arti-
cles of rami of Cirrus I with indistinct or fused
sutures. Cirrus IV with articulations of fourth
and fifth articles of rami supporting about 4
stout claw-like spines on the greater curvature
and 3 to 4 slender long spines on the lesser
curvature. Fourth and fifth articles of Cirrus
VI supporting approximately 4 slender spines
on greater and Z or 3 slender spines on lesser
curvature at the articulations (cf. Plate 22, fi-
gure 8 — acanthopod; figure 9 — lasiopod).
Pedicles of all cirri, except the first, clothed
with minute ctenoid elevations or scales (Plate
22, figure 9).
Counts of articles of the rami are given
below for two spei
with minute ctenoid elevations or scales (Plate
22, figure 9).
Counts of articles of the rami are given
below for two specimens. Although Cirrus I is
6-6, three segments in each are fused. The
first set is for the holotype.
Cirrus I TD) Ot SOEs F We Nad
1. Outer Ramus 6 8 7 8 7 7
Inner Ramus 6 7 8 8 7 6
2. Outer Ramus 6 8 8 8 8 8
Inner Ramus 6 6 9 8 8 8
Caudal appendage as long or slightly longer
than the first article of pedicle of the Cirrus
VI; margins roughened by scales; apex support-
ing a tuft of 4to 6 long slender spines (Plate
22, figure 9). Penis smooth, tapering gradual-
ly through its length; clothed sparsely with mi-
nute setae not symmetrically arranged; termi-
nating abruptly with a tuft of terminal sensory
hairs, apical or just below a rounded terminal
languet.
Discussion
The species described here has been as-
signed to the subgenus Octolasmis because the
valves and their primordia are most compara-
ble to those of other members of this group;
that is, the terga are notched to receive the
distal ends of the occludent arms of the scuta,
and the primordial valves of the scuta lie ina
line along the occludent margin of the capitu-
lum. The latter character is the most undis-
puted diagnostic feature for the subgenus, and
its form clearly separates this species from
species of Temnaspis, which is currently
somewhat enigmatic (Broch, 1932; and 1947;
Nilsson-Cantell, 1934; Stubbings, 1936).
The following species of the Poecilasmati-
dae are known to me from Hawaii:
1. Trilasmis (Temnaspis) fissum hawaiense(Pilsbry, 1928)
1899 Trilasmis fissum Weltner
Trilasmis (Trilasmis) eburneum Hinds, 1844
Trilasmis (PoeciJasma) kaempferi (Darwin, 1851)
Octolasmis (Octolasmis) lowei (Darwin, 1851)
Octolasmis (Dichelaspis) hawaiense (Pilsbry, 1907)
Megalasma (Megalasma) minus Annandale, 1906
1907 Poecilasma bellum Pilsbry
a nS
Of the Hawaiian forms, the present spe-
cies is most similar to Trilasmis (Temnaspis)
fissum hawaiense. These two barnacles are
comparable in size and habitat but are basical-
ly different in the development of the primor-
dial valve of the scutum and the degree of ar-
mament of the capitulum. Trilasmis (T.) fis-
sum hawaiense is more fully covered by its
valves, the scutal umbo is "rotated" basally,
the distal portions of the scutal arms are acute
rather than rounded, the tergum is consider-
ably more reduced and is not notched to receive
the occludent arm of the scutum, and the carina
terminates in an "obtuse tooth and a distinct
heel" rather than a bifurcate knob. Internally,
the scutal platform is basal, corresponding to
the rotation of the umbo, rather than being si-
tuated along the occludent margin. The mouth
trophi are similar in both species, however,
the fourth rather than the second spine below
the notch of the inner maxilla is strongest in
Pilsbry's form, the mandible has but 4 teeth,
including the inferior angle, and the teeth are
simple, lacking spines or pectinations. The
chaetotaxis and mouth trophi of the new species
appear quite similar to those described for
Octolasmis clavula Hiro (1936) from Japanese
waters, especially in regard to the details of
the mandible. However, there are 50 many
differences in the form of the valves that fur-
ther comparison seems unnecessary. A com-
parison with O. tridens (Aurivillius, 1894) and
its Caribbean allies (forms all having ctenopod
cirri) seems superfluous. At this time I see no
close affinities of the new species with any
form known to me. The barnacle clearly
shares characters with members of the genera
Trilasmis, Temnaspis (if considered valid),
and Octolasmis. As Broch (1931) has pointed
out, the group is in need of revision.
The nearly fully armored condition seen in
mature specimens of this species is achieved
Vol. 4; No. 2
THE VELIGER
Page 105
through slow ontogenetic increments in the
valves, particularly in the basal arm of the
scutum, at a rate greater than the rate of in-
crease in the size of the capitulum. By this
sort of differential development, partially ar-
mored juveniles become fully armored some-
what later in life than do related free-living
species in which juveniles are essentially as
fully armored as adults. The same process is
seen to occur in Octolasmis tridens (Aurivil-
lius), (Nilsson-Cantell, 1934), and in O. cor
(Aurivillius), (Newman, 1960b). This process
suggests that species in this family with the
scutum split into two parts have descended
from forms in which the valves were much re-
duced, the split-valve being a product of re-
armament. This suggestion is believed to be
quite plausible for it not only explains their
somewhat aberrant form, but it also explains
the vestige of a scutal suture seen in such spe-
cies as Trilasmis eburneum Hinds and occa-
sionally in T. crassum sensu Darwin (1851).
These two species have apparently completed
the rearmament process, the vestigial, fully-
fused, non-functional suture testifying to a
split-valve ancestry, being acquired through
relatively unarmored forms living in protected
environments.
This argument is contrary to the concept
of the prototype of split-valve forms in which
Pilsbry (1911) envisions a fully armored ances-
tral type with a split-valve already developed,
becoming reduced in protected environments.
The same argument is the reverse of that of
Broch (1947), which is essentially Pilsbry's
concept in more detail, without additional evi-
dence. What Pilsbry and Broch fail to take into
consideration is the functional aspect of the
split-valve; that is, what purpose does it serve
the barnacle, especially when the occludent arm
is immovably fused to the basal arm? They
also fail to consider what possible path selec-
tion might take to achieve such an unusual
structural arrangement.
In fully armored species, without a split-
valve, the scuta are hinged to one another as
they are in nearly naked species of Octolas-
mis. In this way the valves can be opened and
closed, and when closed, being fully approxi-
mate at their edges, they prevent crushing of
the soft animal within. Forms with greatly re-
duced valves are relatively easy to crush, but
they generally survive in environments where
crushing is not likely to occur. If rearmament
were gradually selected for, in response to
more exposed conditions, the barnacle would
still be subject to crushing until the valves be-
came fully approximate. In such partially ar-
mored forms, this problem is overcome by a
fusion of the hinge area of the basal scutal
arms, which, being the strongest and overlying
the body of the animal when withdrawn, offer
fair protection. However, this necessitates
having the occludent arms, each connected to a
basal arm by an uncalcified ligament, free to
be drawn together by the scutoral adductor
muscle, closing the aperture. In this way, par-
tially armored species are protected against
crushing, yet retain the facility of closing. In
exploiting more exposed environments, fuller
armament being selected for, the valves would
become fully approximate. Fusion of the basal
scutal arms is no longer a requirement but ac-
tually a disadvantage, for the occludent arms
must remain free of the basal arms in order to
continue to close the aperture. This arrange-
ment sacrifices a considerable degree of ri-
gidity. In overcoming this final complication in
rearmament, the fusion platforms could become
transformed into a hinge, with the simultane-
ous fusion of the occludent arm to the basal
arm of the scutum, leaving the vestigial suture
mentioned above. Indeed, the peculiar hinge-
like platforms of Trilasmis fissum (Darwin)
and the peculiar hinge of T. eburneum Hinds
are more readily understood as being derived
in this way than from the ordinary hinge of
more remote fully armored ancestral forms.
Following this explanation for the origin of
the split scutal valve, it appears most likely
that split-valve forms have descended from
relatively naked ancestors inhabiting the gill
chambers of decapod Crustacea (Plate 23).
This is not as unlikely as it may at first ap-
pear, for itis a fact that relatively unarmored
species which inhabit gills are often found oc-
curring around the exhalant area of gill cham-
bers and along the basal portions of the last
maxillipeds of the host (Annandale, 1909). In
this position they are at a disadvantage as re-
gards mechanical injury, but they conceivably
may be rewarded by fragments of food drifting
away from the food being fed upon by the host.
To this end a ctenopod feeding mechanism
would be satisfactory, for barnacles could car-
ry on normal setose feeding when not receiving
food escaping the host. If the protection af-
forded by fuller armor were attained in forms
tending to settle near the mouth field, there
would be a. better chance of survival. Thus,
they could occupy positions further out on the
mouth parts, closer to the new source of food.
If the rather delicate feeding mechanism were
gradually replaced with stouter clawed append-
Page 106
THE VELIGER
Vol. 4; No. 2
ages, the barnacle might be able to capture
larger particles of food or even rasp directly
at the food being manipulated by the host. It is
a fact that most split-valve forms occur on the
mouth parts, primarily the maxillipeds of
macrurous forms, and it is a fact that where
the ontogeny of these forms is known, the split-
valve is seen to be acquired through slow onto-
genetic increments and differential growth. It
may be simply a coincidence that most species
in this position also have very peculiar, strong,
stout clawed cirri and unusually spiny mouth
parts which appear poorly adapted to setose
feeding, but I prefer to suggest that these
structures are correlated with the unique habi-
tat in which the barnacle, having descended
from forms inhabiting gill chambers, has
moved forward into the mouth field, enabling it
to share the food of the host.
That such species are the descendants of
more naked ancestors and have come to occupy
such an unusual niche by an indirect route
through the gill chambers of the host is sug-
gested not only because it would explain the
split-valve as a result of rearmament, but also
because a more gradual transitional process
could hardly be imagined in achieving such a
remarkable and precarious position on the host
and in attaining the highly modified feeding
mechanism correlated with it.
Acknowledgment
Specimens of Octolasmis (Octolasmis),
Trilasmis (Trilasmis), T.(Poecilasma), and
Megalasma (Megalasma) studied in conjunction
with this paper were obtained from a number
of individuals to whom I am deeply indebted.
The specimens originated from widely sepa-
rated localities, and together they allow one to
gain a fuller concept of the family than could
otherwise be obtained. I am grateful to Dr.
Thomas E. Bowman for the loan of specimens
fic.
from the Caribbean, to Dr. Fenner A. Chace,
Jr., for the loan of specimens from the Society
Islands, to Dr. Cadet Hand for specimens from
New Zealand and Australia, to Dr. Arthur G.
Humes for specimens from the Caribbean, East
Africa, and New Guinea, to Arnold Ross for
specimens from the Caribbean, to Dr. H. G,
Stubbings for specimens from West Africa, to
Stephen A. Wainwright for specimens from Ha-
waii, to Dr. Huzio Utinomi for specimens from
the Seto Inland Sea, and to Victor A. Zullo for
specimens from Shoal Guyot, South East Paci-
I would also like to thank Dr. Hand for his
reading and criticism of the manuscript.
It is currently my wish to revise the fam-
ily Poecilasmatidae and I solicit specimens
from any quarter of the world for this purpose.
Literature Cited
Annandale, N.
1906a. Natural history notes from the R. I. M. S. 'In-
vestigator''. Ser. III, No. 12. Preliminary report
on the Indian stalked barnacles. Ann. Mag. Nat.
Hist. 7th ser. 17 (100, Art. 53): 389-400.
1906b. Natural history notes from the R. I. M.S. "In- .
veStigator". Ser. III, No. 13. Two new barnacles
dredged in 1905-06. Ibid. 18 (103, Art. 10): 44-47,
2 textfigs.
1908.
Ship ' Investigator"’.
ii, pls. 3-5; Calcutta.
Illustrations of the zoology of the R. I. M. S.
Crustacea entomostraca, Ft.
1909. An account of the Indian cirripedia pedunculata.
Mem. Ind. Mus. 2 (2):61-137, 11 textfigs., pls. 6
and 7.
Aurivillius, C.
1894. Studien Uber Cirripedien. Svensk. Vet. Ak.
Handl. 26 (7): 4-107, 9 pis.
Barnard, K.
1924. Contribution to the crustacean fauna of South
Africa. 7. Cirripedia. Ann. S. Afr. Mus. 20 (1):
103 pp., 1 pl.
Explanation of Plate 23
Schematic drawing indicating certain types of barnacles associated with gills and mouth parts of decapod crustacea.
From right to left: Octolasmis lowei (DARWIN), usually on ‘‘soft’” gills of macrurans, and Octolasmis cor (AURIVILLIUs),
usually on the “hard” gilled brachyuran, Scylla serrata (FORSKAL); Octolasmis indubia NEWMAN, spec. nov. and Trilasmis
fissum (Darwin), both usually on last maxillipeds of macrura. While acanthopod types are quite location-specific,
lasiopod and ctenopod types are not, their degree of armament apparently determining where they will survive.
‘The lasiopod type (PitsBry, 1911) is an intermediate condition between ctenopod and acanthopod types. Split-valve
forms can be acanthopod, lasiopod or ctenopod, while unarmored forms are always ctenopod as are armored forms
that do not have a split-valve ancestry.
Tue VELIGER, Vol. 4, No. 2 [Newman] Plate 23
TAG ie ou Octolasmis indubia | Octolasmis cor Octolasmis lowei
Acanthopod Types Ctenopod ‘T’ypes
Newman, del.
Vol. 4; No. 2 THE VELIGER Page 107
Broch, H. —_,
1931. Indomalayan Cirripedia. Papers from Dr. Th. 1961. Notes on certain species of Octolasmis (Octo-
Mortensen's Pacific expedition 1914-16. 56. Vid-
densk. Meddel. Dansk.naturh. Foren. 91; 142 pp.,
41 textfigs.
—
1947. Cirripedes from Indochinese shallow - waters.
Arch. utg. Norske Vidensk. - Akad. Oslo. I. Mat. -
Naturv. Kl. No. 7: 32 pp., 8 textfigs.
Causey, D.
1961. The barnacle genus Octolasmis in the Gulf of
Mexico. Turtox News 39 (2): 50-55, 17 textfigs.
Darwin, Charles
1851. A monograph on the sub-class Cirripedia. I.
Lepadidae. Ray Soc. London,
to 3, pls. 1-10.
Gruvel, A.
1900a. Note sur deux espéces nouvelles du genre ''Dich-
elaspis''". Proc. Verb. Soc. Sci. phys. nat. Bor
deaux, 1899-1900: 2.
1900b. Sur quelques nouvelles espéces appartenant au
genre Dichelaspis Darwin. Bull. Mus. Hist. nat.
Paris, No. 3: 109-111
pp. 1-400, figs. 1
1905. Monographie des cirripédes ou thécostracés.
pp. 1-472, 472 textfigs. Mason, Paris.
Hiro, F.
1936. Descriptions of three new species of Cirripedia
from Japan. Bull. Soc. Japan 6 (23):
221-230, 3 textfigs.
1937. Studies on cirripedian fauna of Japan II. Cirri-
peds found in the vicinity of the Seto Marine Biolo-
gical Laboratory. Mem. Col. Sci. Kyoto Imp.
Univ. Ser. B, 12 (3, art. 17): 385-478, 43 textfigs.
MacDonald, J. ‘
1869. Onan apparently new genus of minute parasitic
Cirriped, between Lepas and Dichelaspis. Proc.
Zool. Soc. London, 440-444, pls. 33, 34; 1 textfig.
Newman, William A.
1960a. Octolasmis californiana, spec. nov., a pedun-
culate barnacle from the gills of the California spi-
ny lobster. Veliger 3 (1): 9-11, pl. 2.
Biogeogr.
—_—_,
1960b,
Pacific, including two new forms.
(2): 100-116, 6 textfigs.
Five pedunculate cirripeds from the western
Crustaceana ]}
lasmis) from deep sea crustacea. Crustaceana 2
(4): in press.
Nilsson-Cantell, C. A.
1921. Cirripedien Studien. Zur Kenntniss der Biolo-
gie, Anatomie und Systematik dieser Gruppe.
Zool. Bidrag fr. Uppsala 7: 75-395, textfigs. 1 to
89, pls. 1 - 3.
1927. Some barnacles in the British Museum (Nat.
Hist. ). Proc. Zool. Soc. London Pt. 3: 743-790,
19, textfigs.
1934. Indomalayan cirripeds in the Raffles Museum,
Singapore. Bull. Raffles Mus. (9): 42-73, 7 textf
ply:
1938. Cirripedes from the Indian Ocean. Mem. Ind.
Mus. 13 (1): 81 pp., 28 textfigs., pls. 1-3.
Pearse, A.
1932. Observations on parasites and commensals
found associated with crustacea and fishes at Dry
Tortugas, Florida. Pap. Tortugas Lab. Carnegie
Inst. Wash. 28 (6): 103-115.
Pilsbry, Henry A.
1907a. Hawaiian cirripedia. U.S. Bull. Bur. Fish.
26: 181-190, pls. 4, 5, 4 textfigs.
1907b. The barnacles (cirripedia) contained in the col-
lections of the U. S. National Museum. Bull. U. S.
Nat. Mus. 60: 1-122, pls. 1-76, textfigs. 1-99.
===
1911.
Sci.
Remarks on new cirripeds. Proc. Acad. Nat.
Phila. 63: 170-173, 3 textfigs.
1928. Littoral barnacles of the Hawaiian islands and
Japan. Proc. Acad. Nat. Sci. Phila. 79: 305-317,
pls. 24-26, 4 textfigs.
Stubbings, H.
1936. Cirripedia. Brit. Mus. (Nat. Hist.), John
Murray Expedition 1933-34, Sci. Report 4(1): 1-70,
30 textfigs.
Weltner, W.
1899. Ergebni&®e einer Reise nach dem Pacific (Schau-
insland 1896-97). Cirripedien. Zool. Jahrb. Abt.
System. 2: 441-447.
Another Statistical Study in Size of Cowries
by
F. A. ScHILDER
University of Halle (Saale), East Germany
The length of the shells of each cowry spe-
cies varies considerably, even in specimens
coming from a limited area. The frequencies
of the classes, distributed according to a bi-
nomial curve, show a positive skewing so that
the summit of the curve approaches nearer to
the class of the smallest shell than to that of
the largest specimen.
Page 108
THE VELIGER
These usual features may be illustrated by
1'555 Luria isabella (Linnaeus) coming from
the beach of Tjilaut Eureun, South Java, the
length of which varies from 15 to 40 mm. in
adult shells. In the following table the size has
been reduced to classes of 3 mm. (e.g., 15 =
13.5 to 16.5 mm.); the number of specimens be-
longing to every class has been expressed in
percent of 1'555 (the sign 0 indicates less than
0.5 percent; the sign o expresses less than 0.1
percent):
Nye US = S727 Gy RNS SYNE esi 7i ZX)
mm.
L Nw 30 @ Ww 7 1 0 fo) %
The sum of shells coming from the whole
area of distribution of a species supplies a si-
milar curve, if the numbers of specimens pre-
served at each locality are rather similar.
Thus, e.g., the distribution of 8'169 Monetaria
annulus coming from 425 localities between
Natal and Polynesia is as follows:
(x (a) (R}— I
10 13. 16 19 34 mm.
0 LO 83. 32 NY © 1 0 fo) %
The minimum length is 10.3 mm., the maxi-
mum 33.7 mm.; ninety percent of these shells
vary from 14 to 24 mm. only.
The differences in size observed in vari-
ous populations can be environmental or ra-
cial. The local modifications may be caused by
the depth of the sea in which the population
lives, by differences in the motion of the sea or
in the quantity and quality of food: so we learned
from the collections carefully made by R. S.
Benton that most specimens living on the coral
reefs of Mombasa Island are distinctly smaller
than the specimens of the same species col-
lected among the sea weed on a sandy and mud-
dy bottom at Shimoni a few miles off. In the
same way, Barry Wilson stated (personal com-
munication) that in Geographe Bay the adult
specimens of Zoila friendii (Gray) collected be-
tween Quindalup and Dunsborough (about a mile
apart) are stunted: they live "in an enclosed
area in only three to six feet of water cut off
from the deeper water offshore by a wide and
very shallow sand bar.'' He has ''never seena
small adult specimen outside the bank, nor a
large specimen inside the bank'’. He suspects
"the difference may be a physiological one"!
(quoted from Wilson's letter, dated 6th Febru-
ary, 1959). Therefore, the curve of variation in
79 Z. friendii (the Eastern vercoi Schilder in-
Vol. 4; No. 2
cluded) shows two summits:
40 50 60 70 80 90 100 mm.
2n5) 14 9 29 24 19 2.5 %
There are, however, also racial differ-
ences in size, if the average length of most
populations living in a continuous large region
Significantly differs from the usual size ob-
served in other parts of the world. Thus, in my
previous statistical study, published in The
Veliger (Schilder, 1961), I have shown that in
Mauritia arabica (Linnaeus), the South West -
ern* race immanis Schilder-Schilder distinctly
differs by its larger size from M. arabica liv-
ing between Ceylon and the Pacific, though the
limits of variation overlap; this fact may be il-
lustrated by the following table, in which the
size of 2'140 M. arabica and 86 M. immanis has
been expressed in percent:
20 30 40 50 60 70 80 90 mm.
arabica:
oO ish _———
1960. Mantle cavity, habits, and habitat in the blind
limpet, Lepeta concentrica Middendorff. Proc.
Calif. Acad. Sci., 31: 103-110.
the Mytilid Lithophaga
by
NorMAN M. HopGkIN
Department of Zoology, University of California, Berkeley 4, California
(Plates 25, 26, 27; 3 Textfigures)
Lithophaga (synonym: Lithodomus), one of
the rock-boring members of the family Mytili-
dae, occurs throughout the world in tropic and
temperate seas. The genus was erected by
Roding in 1789 and is characterized by its bor-
ing habits. It is found in stone, chalk, coral,
and Spondylus shells, according to Reeve (1858).
I have noted it in lime-cemented sandstone,
calcareous shale, and Chama shells. Sayre
(1931), Hanley (1844), Ihering (1900), and other
authors have specifically mentioned calcareous
material as a substrate for Lithophaga, while
Berry (1907), Amemiya (1933), and Haas (1942)
have reported finding Lithophaga plumula in
non-calcareous material,
Carazzi (1903), List (1902), Kiihnelt (1930),
Otter (1937), and Yonge (1955) have all recog-
nized the possibility that Lithophaga may bore
into limestone substrates with the aid of an
Page 124
THE VELIGER
Vol. 4; No. 3
acid secretion from the mantle tissue or at
least by some chemical means. Of the investi-
gators that have reported Lithophaga in non-
calcareous material, Haas (1942) is the only
author specifically to contradict the acid-boring
theories. He does not offer an explanation for
boring, but does state categorically that it can-
not be chemical and it therefore must be me-
chanical. A summary of the findings of the
above-mentioned authors appears in the next
section.
It has been the purpose of this study to de-
termine whether Lithophaga plumula bores into
rock by a chemical or a mechanical process,
The conclusion of this work is that L. plumula
kelseyi does bore by a chemical action on the
rock substrate, but it is beyond the scope of
this paper to analyze the chemical mechanisms
involved.
HISTORICAL THEORIES OF BORING
BY LITHOPHAGA
Carazzi (1903) was the first worker to pub-
lish a theory of boring by Lithophaga. He de-
scribed three glands in the mussel which he
claimed were acid-secreting: a gland in the
visceral mass of the animal which he called
“protacid gland", and two "'deutacid glands",
One of these ''deutacid'’ glands is anterior to
the hinge, just dorsal to the anterior adductor
muscle, and is called the ''anterior acid gland".
The other gland is called the "posterior acid
gland''and is immediately posterior to the hinge
in the extreme dorsal position adjacent to the
internal surface of the shell valves.
List (1902) has also suggested that these
mussels bore by acid secretions from the area
of the fused inner mantle lobes. He described
two types of gland cells: basophils with hyaline
content and granulated acidophils.
Pelseneer (1911) redescribed the same
glands that Carazzi and List had seen and em-
phasized that these glands occur in the Mytili-
dae. Pelseneer also described a second pallial
gland in Lithophaga gracilis which, he said, is
absent in other species of Lithophaga. Pelse-
neer claimed that this unpaired gland may have
something to do with boring and, in fact, called
it an ''acid"’ gland. Yonge (1955) dismissed its
connection with boring by saying that this gland
secreted into the mantle cavity and so the acid,
if there were such, would never come in con-
tact with the rock substrate. Yonge (1955) did
not find this gland in the Californian L. plumu-
la.
the shell.
In a long and general discussion Kihnelt
(1930) reviewed the history of the theories of
boring by the pelecypods and brought it up to
date with his own observations. Kiihnelt recog-
nized two mechanisms by which pelecypods
bore into their substrates: mechanical and
chemical. Most of the paper dealt with chemi-
cal borings as exemplified by Lithophaga. Ktih-
nelt's experimental work showed that Lithopha-
ga does bore chemically. He put a piece of
Sepia shell in a tube and allowed one of his spe-
cimens to press its anterior mantle up against
There were marks of erosion in a
few days, although Kithnelt had not seen any
movement by the mussel. He was also able to
detect actual boring of from two to three milli-
meters in pre-bored Sepia shells in a period
of 14 days. Kihnelt put some Lithophaga in
paraffin-lined glass test tubes but was unable
to detect any free acid in these tubes over a
period of eight days. He also attempted to re-
peat Carazzi's experiments in which the latter
had shown that the so-called "'acid gland" tissue
turned blue litmus paper red. Kiihnelt found
that this reaction was not limited to the so-
called "glandular" tissue. The mantle, gill,
and foot also turned blue litmus paper red.
Kihnelt further stated that all experiments to
prove the existence of acid in the glands were
negative, although he did not describe his ex-
periments. He was convinced that the boring is
effected chemically and that the agent which
causes the solution does not come from any
specific source in the animal.
Otter (1937) studied several species of
Lithophaga at Low Isles on the Great Barrier
Reef. He included Lithophaga and probably
Modiolus as. chemical borers among the Mol-
lusca in a general article on rock-destroying
organisms. He felt that the rock is dissolved
away by an acid but stated that no acid had been
identified. He suggested hydrochloric acid as
the most likely but also considered an organic
acid or mixture of acids. Otter cited Duerden
(1902) as saying that most boring algae utilize
carbonic acid from respiration and Carazzi
(1892) as saying that carbonic acid is also used
by Lithophaga for boring. Otter objected to the
idea that free acid is secreted into the burrow
because if it were, the calcareous deposits on
the shell and burrow would be dissolved. He
wrote (l.c., p. 336), "It appears that the acid
secretion is only applied directly to those sur-
faces of the burrow in contact with the free
edges of the siphons and mantle when these are
protruded, the acid secretion being immediate-
ly neutralized by the rock before it can come
Vol. 4; No. 3
THE VELIGER
Page 125
into contact with the shell or any other region
of the burrow."
Yonge (1955) made observations on Litho-
phaga plumula (Hanley, 1844) at Pacific Grove,
California. He actually saw a specimen of
Lithophaga extrude its anterior mantle tissue
and place it in intimate contact with the rock at
the head of a broken burrow. Yonge was unable
to detect any free acid on the surface of the
fused inner lobes of the mantle. He did confirm
the glandular areas, both anterior and posteri-
or, that were described by Carazzi, List, and
Pelseneer. Yonge postulated an acid mucous
secretion from these glands. The anterior
gland would do the original softening and the
posterior gland would widen the burrow as the
animal grows in size. Yonge saw cilia on the
protruded anterior mantle tissue and suggested
that these cilia carry the dissolved rock and
mucus ventrally and then posteriorly into the
mantle cavity. This dissolved material then
passes posteriorly and out the dorsal surface
of the inhalant siphon, which is the usual tract
for the removal of pseudofeces,.
Three authors, Berry (1907), Amemiya
(1933), and Haas (1942), have reported finding
Lithophaga plumula in non-calcareous sub-
strates. Berry claims to have found L. plumula
in hard blue clay dredged from 12 fathoms in
Monterey Bay. The Lithophaga were found with
other pelecypods which are known to bore me-
chanically, such as Botula, Parapholas califor-
nica (Conrad, 1837), and three species of
Pholadidae. Amemiya (1933) reported finding
Lithophaga in tuffaceous mudstone. He did not
test the mudstone for calcium carbonate con-
tent. Haas (1942) collected L. plumula in lime-
cemented sandstone and in non-calcareous ar-
gillaceous shale, both at LaJolla and at Pacific
Grove, California. The first two authors men-
tioned did not comment on the methods of bor-
ing. Haas (1942) offered no theories on chemical
or mechanical boring except to state that the
boring could not be chemical. The reason he
gave for this statement was that he had found
Lithophaga in non-calcareous shale along with
Irus lamellifer (Conrad, 1837), Botula califor-
niensis Philippi, 1847, and some pholads. If
this be true, it would suggest that Lithophaga
can bore mechanically.
MATERIALS AND METHODS
Field and laboratory studies were made at
the Kerckhoff Marine Laboratory, Corona Del
Mar, California. Further laboratory experi-
ments were conducted at the University of Cali-
fornia, Berkeley, California. The specimens
used for these studies were collected at Car-
penteria, California, and at Corona Del Mar.
The specimens were identified by Dr. L. G.
Hertlein of the California Academy of Sciences
as Lithophaga plumula kelseyi Hertlein and
Strong, 1946. This subspecies is found from
San Diego, California, to Duxbury Reef, imme-
diately north of San Francisco Bay andis shown
in Plate 25, figures 1, 2, 3, and 4.
Two separate experiments were set up for
the observation of Lithophaga in the laboratory
in both artificial and natural burrows. In the
first experiment, which will be identified as
Experiment No. 1, ten of the mussels were left
on the bottom of a five-gallon tank of aerated
sea water. No burrows were provided. Three
others were put into Pyrex test tubes of ap-
proximately the same diameter as the original
burrows, and five were put into holes bored in
limestone rock with a tungsten-carbide drill
bit. The shape of the bottom of the holes in the
limestone is shown by the dotted line across
the bottom of the hole in textfigure 1. The pur-
pose of this experiment was to observe the ani-
mals and to determine if they could live for a
protracted period in the laboratory.
The second experiment (No.2) was a con-
trolled experiment in which freshly collected
specimens of Lithophaga plumula kelseyi were
placed in holes bored in the same limestone as
that used in Experiment No.1, and in a non-
calcareous mudstone. The limestone had a
g
Figure 1: Cross section of a typical artificial burrow in
limestone and mudstone showing extent of boring
activity by Lithophaga
Page 126
hardness equal to fluorite (number 4 on the
hardness scale), The mudstone had a hardness
of 3, equivalent to calcite. The mudstone was
obtained from Duxbury Reef, Bolinas, Califor-
nia, and is a rock in which numerous mechani-
cally boring mollusks such as Botula, Platyo-
don, and Pholadidea are found, Lithophaga were
placed in the holes in the two pieces of rock,
and the rocks were put into a five-gallon aquar-
ium filled with sea water which was kept aerat-
ed at all times. Experiment No.1 lasted one
year, followed by Experiment No.2 which was
continued for six months,
The purpose of Experiment No.2 was to
compare the boring abilities of Lithophaga ina
rock which can be attacked and dissolved by
chemical action, namely, calcium carbonate or
limestone; and in a rock that cannot be affected
by an acid or by chemical activity, viz., non-
calcareous mudstone. Additional observations
were made on the rotation of the animals in the
mudstone and limestone burrows.
Miscellaneous observations were made on
animals from both of the above-mentioned ex-
periments in regard to deposits on the burrow
walls. The anterior end of one of the natural
burrows was broken off, and the activities of
the anterior mantle tissue of the enclosed spe-
cimen was observed.
OBSERVATIONS AND RESULTS
Lithophaga Burrows in Nature
In nature Lithophaga burrows are found in
carbonate-cemented sandstone and calcareous
shale. The inside of the burrow is a little long-
er than the animal, which allows it to move
back and forth by the action of the byssal re-
tractor muscles. When the animal is in its ex-
treme posterior position, the ends of the
"plume'' are level with the plane of the entrance
of the burrow (textfig.2). The aperture is
small and is usually found with a mass of white
limy material surrounding it and extending it
above the rock surface for a distance of about
two millimeters. This white material fizzed
THE VELIGER
Vol. 4; No. 3
and produced CO, gas when it was treated with
dilute hydrochloric acid. The cation was not
determined but was presumed to be calcium.
The sandstone burrows were variably lined
with a white substance which also proved to be
a carbonate and was thus assumed to be calci-
um carbonate. In all cases the posterior third
was lined and in some cases observed the en-
tire burrow was lined, as in Plate 25, figure 5.
= WET,
Vi Aye
=
\
EGON
Lyd
ger {jor
w ———
Figure 2: Lithophaga in its most posterior position (top)
and most anterior position (bottom) in its natural rock
burrow.
Experimental Results
The main purpose of Experiment No. 1 was
to determine the length of time Lithophaga
could live under laboratory conditions in a five-
gallon tank of sea water both inside and outside
the confining burrows. The specimens in the
artificially bored limestone burrows, in the test
tubes, and on the bottom of the tank all lived
through a period of one year. No attempt was
made to feed the animals, and the water was not
changed. It was noted, however, that there was
a fairly uniform algal growth on the rocks and
on the walls of the tank, indicating that free-
Explanation of Plate 25
Figures 1 to 4:
Figure 1: Ventral view. Figure 2: Dorsal view.
Figure 5:
Lithophaga plumula kelsey.
Figure 3: Lateral view. Figure 4: Lateral view.
A longitudinal section of a Lithophaga burrow in lime-cemented sandstone showing complete
calcareous lining.
Tue VELIGER, Vol. 4, No. 3 [Hopexn] Plate 25
re
pct,
va
ogee
Hopexw, photo,
THE VELIGER, Vol. 4, No. 3 [ Hopckn | Plate 26
Bottom of the burrow of Textfigure 3. The dark area in the center is the erosion caused by
the specimen of Lithophaga that was unable to rotate. Notice the symmetry
across the dorso-ventral axis.
Hopoxrn, phote.
Vol. 4; No. 3
THE VELIGER
Page 127
swimming algal gametes were probably avail-
able for food.
The three groups of Lithophaga all depos-
ited a material assumed to be calcium carbo-
nate. The specimens on the bottom of the tank
placed their deposits flat on the glass bottom
adjacent to their ventral surface where they had
attached themselves by their byssal threads.
The observation that led directly to Exper-
iment No.2 was that the specimens of Litho-
phaga left in the limestone holes all continued
to bore into the rock. After the animals were
removed, symmetrical, concave depressions
were seen at the bottom of the holes as shown
in textfigure 1. The volume of material re-
moved by the two largest Lithophaga (about 6
cm. in length) was measured and found to be
0.12 cm3 in one case and 0.17 cm? in the other
case.
The next point to determine was whether
Lithophaga could elongate artificially-bored
holes in a non-calcareous mudstone under labo-
ratory conditions similar to those in Experi-
ment No.1. Experiment No. 2 was set up with
a block of mudstone from Duxbury Reef, anda
new piece of the same limestone that was used
in Experiment No.1 was used as a control.
The result of Experiment No. 2 was that all
four specimens of Lithophaga continued to bore
into the limestone but none of the three individ-
uals in the mudstone had any effect whatsoever
on that rock. No erosion could be detected,
even microscopically, on the bottom of the
mudstone holes. The bottoms of the holes in
the mudstone remained as they had been bored
by the drill, as shown by the dotted line in text-
figure 1. The volume of material removed
from the limestone by three of the Lithophaga
was measured and found to be 0.18, 0.23, and
0.12 cubic centimeters, respectively. The
fourth Lithophaga burrow was not measured be-
cause of accidental fracture of the rock. This
particular specimen was slightly larger than
the others and asa result was not able to rotate
in its burrow as did the others. The shape of
the excavation caused by this particular animal
is significant as proof against a mechanical
theory of boring and is discussed at the end of
this section and in the conclusion.
Rotation of the Lithophaga specimens in
their burrows was observed during Experiment
No. 2 and checked by the actual byssal attach-
ments at the conclusion of the experiment. Both
groups rotated very little, but it was noted at
the end of the experiment that the animals in
the mudstone as a group rotated less than those
in the limestong, In natural burrows the points
of attachment around the inside are quite regu-
lar, indicating that the animal in nature does
not seem to favor any one position.
One of the Lithophaga in the limestone of
Experiment No.2 was too large to rotate free-
ly. It was able to gape a very slight amount,
however, and this gape was sufficient to allow
anterior mantle tissue to flow out and up against
the head end of the artificial burrow. The ex-
cavation was symmetrical about the projection
of the dorso-ventral axis on the head end of the
burrow, but it was not radially symmetrical.
The holes eroded by the Lithophaga which were
free to rotate in both Experiments 1 and 2 were
very nearly radially symmetrical. Textfigure
3 and Plate 26 show the shape of the hole erod-
ed by the individual which was stuck in its hole,
Records of the rotation of all the animals were
kept throughout the experiment. At no time
was the specimen which made the boring shown
in textfigure 3 and Plate 26 seen to rotate. As
further evidence against rotation by this animal
the points of byssal attachment were checked
after the animal was removed, Only one small
area was covered heavily by the byssus im-
plantations, proving that the animal kept its
dorso-ventral orientation constant during the
experiment.
Figure 3: View of artificial burrow in limestone in
which the animal was unable to rotate. The plane of
the break in the rock is almost in the plane of symmetry
of excavation.
Page 128
THE VELIGER
Vol. 4; No. 3
Miscellaneous Observations
A specimen of Lithophaga plumula kelseyi
in its natural burrow was placed in the tank
with Experiment No. 1 and kept alive for a year.
The anterior tip of the burrow was broken off.
From time to time mantle tissue would extend
out of the broken end and expand. The appear-
ance of the mantle tissue as shown in Plate 27,
figures 1 and 2, would seem to indicate that if
the burrow end had been intact the tissue would
have made intimate contact with the rock sur-
face. In the case of the animal that could not
rotate, it is apparent that some sort of chemi-
cal action is responsible for the peculiar shape
of the erosion caused by that animal. Consid-
ering the shape of the eroded area in question
and the fact that sea water is a highly buffered
medium, intimate contact of the mantle tissue
with the rock may be a necessary part.of the
boring process,
The deposits of calcareous material on the
walls of the burrows may furnish a clue to the
mechanisms involved in boring by Lithophaga
plumula kelseyi. The deposits on the walls of
the glass test tubes in Experiment No. 1 and the
mudstone in Experiment No.2 were chalky in
appearance and very fragile. There was a
small amount of deposition compared to the de-
posits on the walls of the artificially-bored
limestone burrows. The deposits on the lime-
stone burrows were shiny, very hard, and me-
chanically strong. The deposit on the wall of
the natural burrow deserves some comment.
In the case of the small burrows, only the pos-
terior half or third was covered with the cal-
The largest burrows were
completely lined with the material. A possible
explanation for this is that the small and pre-
careous substance.
sumably young animals are still growing and,
consequently, must continue to elongate their
homes. It may also be assumed, therefore, that
the largest specimens are the adults and as
such no longer need to bore. The adults then
lay down a continuous covering over the entire
inner surface of the burrow.
Conclusions
The direct evidence from the boring activ-
ity of Lithophaga plumula kelseyi as compared
in limestone and mudstone would seem to indi-
cate that this species bores by some sort of
chemical rather than mechanical means, This
evidence is supported by the observation that
the specimen that was stuck in its burrow and
could not rotate did actually dissolve limestone
and in so doing made an oddly shaped excava-
tion that could not have been made by any sort
of rotation.
The argument against a mechanical method
of boring is as follows: The animals in the
mudstone were able to rotate and did rotate but
had no abrasive effect on the burrow bottoms.
The specimen which was stuck in the limestone
burrow and was not able to rotate did make an
excavation. It is not known whether this dis-
solving of the limestone was caused by an acid
mucus secretion by the mantle or by a more
complex activity such as ion exchange between
the mantle tissue and the calcareous rock.
Acknowledgment
I wish to acknowledge the help of the fol-
lowing people in collecting specimens, provid-
ing research facilities, identification of the
animals, and criticizing the paper: Dr. D. Da-
venport, Mrs. G. E. MacGinitie, Dr. ©) Ho:
Hand, Dr. R. 1. Smith, Dr. J. W. Durham, and
Delain Gepilertlerns
Literature Cited
Amemiya, I. Y., & O. Amemiya
1923. Note on the habitat of rock-boring molluscs on
the coast of central Japan. Proc. Imp. Acad. (To-
kyo) 9: 120-123.
Berry, S. Stillman
1907. Molluscan fauna of Monterey Bay,
Nautilus 21] (2): 17-21.
Carazzi, D.
1892.
dei Datteri (Lithodomus dactylus).
sci. nat. Anno 3: 279-297.
California.
La perforazione delle rocce calcaree per opera
Atti. Soc. Li-
gust.
1903. Contribute all'istologia e alla fisiologia dei La-
mellibranchi. Int. Monat. Anat, Physiol., 20: 57
to 86.
Duerden, J. E.
1902. Boring algae as agents in the disintegration of
corals. Bull. Amer. Mus. Nat. Hist. New York,
16: 323-332.
Explanation of Plate 27
Figure 1:
Anterior mantle tissue of Lithophaga plumula kelseyi when extended, looking posteriorly.
Figure 2: Lateral view of extended anterior mantle tissue of the same animal.
Tue VELIGER, Vol. 4, No. 3 [Hopcxin] Plate 27
PG a 1 : 5 ‘
Figure 2
Hopexw, photo,
Vol. 4; No. 3
THE VELIGER
Page 129
Haas, Fritz
(1942. The habits of life of some west coast bivalves.
Nautilus 55; 109-113.
Hanley, S.
1844, Description of new species of Mytilacea, & C.
Proc. Zool, Soc. London, p. 17.
Hertlein, Leo G,, & A. M. Strong
1946. Mollusks from the west coastof Mexico and Cent-
ral America. Zoologica, 31: 53-79.
Ihering, H. von
1900. On the South American species of Mytilidae. P.
Malacol. Soc. London, 4: 84-98.
Kuhnelt, W.
1930. Bohrmuschelstudien, 1. Paleobiologica 3: 53 to
91.
List, Th.
1902, Die Mytiliden des Golfes von Neapel und der an-
grenzenden Meeresabschnitte. Fauna & Flora Gol-
fes von Neapel, Monogr. 27.
Otter, G. W.
1937, Rock destroying organisms in relation to coral
reefs, Sci, Repts. Gr. Barrier Reef Exp. 1: 323
to 352.
Pelseneer, P.
1911, Les lamellibranches de l'expédition du Siboga.
Siboga-Expeditie, Leiden, Monogr. 53a: 1-125,
Reeve, Lovell Augustus
1858, Conchologia Iconica. Monograph of the genus
Lithodomus.
Sayre, A. N.
The fauna of the Drum limestone of Kansas and
Kansas Univ. Sci. Bull. 19 (2):
1931.
western Missouri.
102-202; pls. 1-21.
Yonge, C. M.
1955, Adaptations to rock boring in Botula and Lithopha-
ga (Lamellibranchia, Mytilidae) with a discussion
on the evolution of this habit. Quart. J. Micr.Sci.
96: 383-410.
The Fresh Water Clam Pisidium tremperi (HANNIBAL)
D. W. TAYLOR & H. B. HERRINGTON
U. S. Geological Survey, Washington, D. C., and Westbrook, Ontario, Canada
(Plate 28)
(Publication authorized by the Director, U. S. Geological Survey)
Only one species of freshwater clam has
been considered restricted to southern Califor-
nia. Pisidium tremperi (Hannibal) has so far
been known only from Bluff Lake, San Bernar-
dino Mountains. We have examined the type lot
and refer P. tremperi to the synonymy of the
widespread P. obtusale Pfeiffer. The species
is known nowhere else in the State, however,
and its disjunct occurrence in southern Cali-
fornia is parallel to that of other locally re-
stricted species.
SYNONY MY
Corneocyclas sp. nov.: Hannibal, 1912a,
p. 42.
Corneocyclas Tremperi, n. sp.: Hanni-
bal, 1912b, p. 137, 210, pl. 7, fig. 22.
Pisidium trempéri Hannibal: Sterki,
1916, p. 471.
Pisidium tremperi (Hannibal): Gregg,
1947, no. 70, p. 18.
ORIGINAL DESCRIPTION
"Shell minute, globular-trigonal, beaks
anterior, broad and somewhat elevated, hinge
much reduced; habitat marshy lakes. Length
1.4, altitude 1.3 mm., depth of valves 1 mm.
"Mojave System.
"Bluff Lake Cienaga, San Bernardino
Mountains, California (H. Hannibal).
"Named after Dr. R. H. Tremper, the first
conchologist to visit this portion of the San
Bernardino Mountains" (Hannibal, 1912b, p, 137).
This inadequate description is not usefully
supplemented by the poor illustration.
Page 130
THE VELIGER
Vol. 4; No. 3
IDENTITY
Considering the inadequate description by
Hannibal, one cannot blame later writers for
misinterpreting Pisidium tremperi. Sterki
(1916, p. 471) thought it was close to P. ashmu-
ni, a synonym of P. casertanum (Poli). Gregg
(1947, no. 70, p. 18) thought P. tremperi was
probably the young of P. ashmuni.
We have examined the following material:
4827S. S. Berry collection. Lower end of
Bear Lake, alt.6700 ft., San Bernardino Moun-
tains. S.S. Berry, August 1908. Identified as
Pisidium rotundatum Prime by Victor Sterki,
January 26, 1920.
4824 S. S. Berry collection. Swamp at
Bluff Lake, alt. 7550 ft., San Bernardino Moun-
tains. S.S, Berry, August 1908. Identified as
Pisidium rotundatum Prime by Victor Sterki,
January 26, 1920.
5805 Stanford University Paleontological
Type Collection. Bluff Lake Cienaga, San Ber-
nardino Mountains. Harold Hannibal. Type and
three others.
All these specimens are closely similar
and easily referable to the form rotundatum of
Pisidium obtusale Pfeiffer. They have the
characteristic oval shape, fine striae, small
size, short hinge, and low rounded, smooth
beaks. The cusps on the lateral teeth are sharp
on top as in P. obtusale, not blunt. The pseu-
docallus at the proximal end of the posterior
sulcus is weak, and the shells are not as large
as usual P, obtusale, but these are trivial dif-
ferences.
The type (Plate 28, figures 1 and 2) mea-
sures as follows: Length 1.8 mm., breadth 1.3
mm., height 15mm. The paired valves were
not separated for fear of breakage, but the spe-
cimen is so close to the others in external fea-
tures that one cannot reasonably doubt they are
all one species.
Our measurements of the type do not agree
well with those published by Hannibal, It is
possible but not certain that we have erred in
simply accepting the identification as type on
the labels. If this specimen is not the type,
then the identity of Pisidium tremperi remains
It is a synonym of either P. caser-
tanum or P. obtusale, in our opinion.
uncertain.
ZOOGEOGRAPHY
Although we have deprived southern Cali-
fornia of an endemic species, the isolated oc-
currence there of Pisidium obtusale retains
considerable zoogeographic interest. The dis-
junct distribution is shown to be of historical
significance by the occurrence in the San Ber-
nardino Mountains of seven other species of
mollusks also isolated there.
FRESHWATER CLAMS, SPHAERIIDAE
Pisidium obtusale Pfeiffer. In California
known only from Bear Lake and Bluff Lake, San
Bernardino Mountains.
Sphaerium lacustre (Miller). In southern
California known only from Bluff Lake, San
Bernardino Mountains (Hannibal, 1912a, p. 42;
Gregg, 1947, no. 69, p. 12, as Musculium ray-
mondi).
Sphaerium hanhami Sterki. In southern
California known only from Bear Lake, San
Bernardino Mountains (Gregg, 1947, no. 69, p.
13).
FRESHWATER SNAILS, VALVATIDAE
Valvata humeralis californica Pilsbry. In
southern California known living only from
Bear Lake and Bluff Lake, San Bernardino
Mountains (Hannibal, 1912a, p. 35). It is re-
corded by Woodring and others (1946, p. 65)
from the upper Pleistocene of the Palos Verdes
Hills.
LYMNAEIDAE
Stagnicola sp. Whatever may be the spe-
cies recorded by Hannibal (1912a, p. 42) and
Berry (1909, pp. 76-77) as Lymnaea palustris
(Miller), it is known in southern California only
from Bluff Lake and Bear Lake, San Bernardi-
no Mountains.
PLANORBIDAE
Menetus centervillensis (Tryon). In south-
ern California known only from Bear Lake, San
Bernardino Mountains (Baker, 1945, p. 478;
Hannibal, 1912a, p. 42,as Segmentina dilatata).
LAND SNAILS, PUPILLIDAE
Vertigo modesta form microphasma Berry.
THE VELIGER, Vol. 4, No. 3 [TAYLOR & HERRINGTON | Plate 28
Elinor
Sere’
Figure 1
Figure 2
Pisidium obtusale PFEIFFER form rotundatum PRIME (type specimen of P. tremperi HANNIBAL), xX 27.
Bluff Lake, San Bernardino Mountains, San Bernardino County, California.
Vol. 4; No. 3
THE VELIGER
Page 131
Known only from the San Bernardino Mountains
(Berry, 1919; Pilsbry, 1948, p. 987). —
Vertigo occidentalis Sterki. Known only
from the San Bernardino Mountains (Pilsbry,
1948, p. 993).
Hannibal (1912a, pp. 40-42) accounted for
the aquatic species in the Pacific Coast drain-
age by stream capture of former Mojave head-
waters. Pleistocene uplift of the San Bernar-
dino and San Gabriel Mountains, and the recent
origin of the present Mojave stream courses
are documented geologically (Noble, 1954;
Bowen, 1954). From this point of view, Hanni-
bal's interpretation is plausible. Another zoo-
geographic question remains unanswered,
however: Why are these local populations of
mollusks (some differentiated, some not) in the
San Bernardino Mountains? Their isolation and
differentiation may considerably antedate re-
cent drainage changes. The occurrence of
these species in the coastal drainage may there-
fore be a relatively minor geographic phenom-
enon superimposed on the older biological fea-
ture of local isolation.
In the case of Valvata, one can say that the
history of the species in southern California is
more complicated than appears at first sight.
Although the only known living occurrences are
in the Bear Lake area of the headwaters of the
Santa Ana in the San Bernardino Mountains,
this snail is known as an upper Pleistocene fos-
sil from the Palos Verdes Sand in what is now
Los Angeles River drainage (Woodring and
others, 1946, p. 65). Evidently the restriction
of Valvata in southern California to the San
Bernardino Mountains is a relatively recent
feature of its distribution. Probably its local
occurrence is due to the disappearance of other
formerly suitable habitats. Whether Pisidium
obtusale is analogous to the Valvata in this re-
spect, one cannot Say.
Acknowledgment
We are grateful to S. S. Berry, Redlands,
California, and to A. Myra Keen, Stanford Uni-
versity, Stanford, California, for loan of the
specimens which form the basis of this note.
Literature Cited
Baker, F, C,
1945. The molluscan family Planorbidae.
nois Press. xxxvi + 530 pp., frontis.
Berry, S. Stillman
Univ. Illi-
1909. The known mollusca of San Bernardino County,
California, Nautilus 23; 73-79.
SS,
1919, Three new alpine Vertigos from California.
Nautilus 33: 48-52; figs.
Bowen, O. E., Jr.
1954. Geology and mineral deposits of Barstow quad-
rangle, San Bernardino County, California. Calif.
Div. Mines Bull. 165: 7-185, pls. 1-9.
Gregg, Wendell O.
1947. The fresh water mollusca of California, includ-
ing a few forms found in adjoining areas, Conch.
Club Southern Calif., Minutes 67: 3-21; 69: 3-18 ;
70: 14-20.
Hannibal, Harold
1912a. The aquatic mollusc of Southern California.
Southern Calif..Acad. Sci. Bull. 11: 18-46, frontis.
1912b. A synopsis of the Recent and Tertiary freshwa-
ter mollusca of the Californian Province, based
upon an ontogenetic classification. Malacol. Soc.
London Proc., 10: 112-211, pls. 5-8.
Noble, L. F.
1954,
to Cajon Pass, California.
170, chap. 4: 37-48, pl. 5.
Pilsbry, Henry A.
1948. Land mollusca of North America (north of Mex-
ico), v. 2, pt. 2. Acad. Nat. Sci, Philadelphia
Monogr. 3, v. 2, pt. 2: i-xlvii, 521-1113.
Sterki, Victor
1916. A preliminary catalogue of the North American
Sphaeriidae. Carnegie Mus. Annals 10: 429-477.
Woodring, W. P., M. N. Bramlette, & W. S. W. Kew
1946, Geology and paleontology of Palos Verdes Hills,
California. U.S. Geol. Survey Prof. Paper 207,
v + 145 pp., 37 pls.
1-8.
The San Andreas fault zone from Soledad Pass
Calif. Div. Mines Bull.
Page 132
THE VELIGER
Vol. 4; No. 3
On the Identifications of Five Pacific Mitra
by
JEAN M. CaTE
Conchological Club of Southern California, Los Angeles 7, California
(Plate 29)
A great deal of confusion exists concerning
the species generally identified as Mitra lugu-
bris Swainson, 1822, M. coronata Lamarck, 1811
(non Helbling, 1779), M. tiarella A. Adams,
1851, and M. lugubris honoluluensis Pilsbry,
1920.
It has been pointed out (Dautzenberg and
Bouge, 1923) that the name Mitra coronata La-
marck, 1811 is preoccupied by Voluta coronata
Helbling, 1779, which applies to a species
Swainson later named Mitra mucronata. If
these two species were classified in the same
genus, the Lamarck name would be a secondary
homonym, but since under modern practice M.
coronata Lamarck is retained in the genus Mi-
tra, and M. mucronata Swainson (Voluta coro-
nata Helbling) is placed in Vexillum, and since
no valid replacement name has been proposed
for the presumed junior homonym, it seems
possible under the International Rules for both
names to be retained — as Mitra coronata La-
marck, 1811, and Vexillum coronatum (Helbling,
1779) — at least until the long-awaited new
Code is published and the matter of secondary
homonymy is clarified.
Mitra coronata and M. lugubris were in-
correctly illustrated by Sowerby (1874) and by
Tryon (1882); only one of Sowerby's three fi-
gures of M. lugubris accurately illustrates the
species, and his drawings purporting to repre-
sent M. coronata are equally confused. Of the
eight figures labelled M. coronata by Tryon,
only one is referable to the species, although
two of his figures of M. lugubris in reality de-
pict M. coronata. None of Tryon's "lugubris"'
figures is actually that species. Tryon consid-
ered M. tiarella a synonym of M. coronata;
Sowerby figured it accurately on two occasions,
though again referring in another case to an
identical species as M. coronata. These erro-
neous identifications in two widely used mono-
graphs may account for some of the incorrect
labels so frequently seen in collections. Daut-
zenberg and Bouge (1923) also considered M.
coronata and M. tiarella synonymous, basing
this decision mostly on the size differential
seen in various specimens. It is true that M.
tiarella is extremely variable in size; in my
collection there are perfectly typical adult spe-
cimens ranging from 12.4 mm. to 31.8 mm.,
with all intermediate sizes well represented.
There are other important morphological dif-
ferences between these two species, which are
enumerated below.
A list of references to some of the valid
illustrations will be found at the end of this pa-
per, as well as new photographs of typical spe-
cimens of these species and two additional re-
lated forms, M. aurora Dohrn, 1861 (Plate 29,
fig. 4) and M. floridula Sowerby, 1874 (Plate 29,
fig. 5).
The original figure of Mitra coronata re-
ferred to by Lamarck(Encyclopédie Méthodique,
Pl, 371, figs. 6a, 6b) and subsequent illustra-
tions by Kiener, Kuster, Reeve, Wood) and
Chenu represent a species similar to M. lugu-
Explanation of Plate 29
Figure 1: Mitra lugubris Swatnson, 1822. Queensland, Australia (right-hand specimen courtesy of Helen DuShane)
Figure 2: Mitra coronata LAMARCK, 1811. Hawaii.
46 797.
Figure 4: Mitra aurora Donen, 1861. Hawaii.
Figure 3: Mitra tiarella A. ADAMS, 1851. Hawaii. ANSP No.
Figure 5: Mitra floridula SowERBy, 1874. Okinawa.
Photographs (except Figure 3) by Victor Duran, Scientific Photographic Laboratory, University of California.
Figure 3 photograph by Perfecto Mary, courtesy of Stanford University. All photographs twice natural size.
All specimens from Cate collection unless otherwise noted.
Tre VELIGER, Vol. 4, No. 3 [CaTE] Plate 29
Figure 1 Figure 3
Figure 2 Figure 4
Figure 5
Vol. 4; No. 3
THE VELIGER
Page 133
eS
bris in some ways: its color is dark brown, or
brownish-orange in faded specimens, and it is
crowned with a wide white crenulated zone be-
low the sutures. Otherwise, it differs from M.
lugubris as follows: it is a slimmer shell,
more ovate than pyriform; its aperture is shin-
ing and white instead of dull brown; its lipis
effuse instead of following the pyriform outline;
but most of all, the true M. coronata lacks the
white base which is the most striking feature of
M. lugubris. This white base is visible from
the dorsal side of the anterior tip of the shell;
on the ventral side it extends to the adapical
fold of the columella. The remainder of the
columella is brown, as is the labral side of the
aperture except where the white subsutural
band may be seen from inside.
Mitra tiarella A. Adams, 1851 (M. lugubris
honoluluensis Pilsbry, 1920) differs from M.
lugubris and M. coronata in that it exhibits a
narrow pale band below the sutures, and its
crenulations are white on a brown background,
there being no solid white subsutural zone.
Correct Illustrations of the Above Species:
Mitra lugubris Swainson, 1822 (see Plate 29,
figure 1):
1822 Swainson, William. Zoological Illus-
trations, Vol. 2, Pl. 66, upper and
lower figures.
1839 Kiener, L. C. Icon. Coq. Viv., Mi-
tra, Pl. 30, fig. 100.
1841 Kitster, H. C. Conchylien-Cabinet,
Mitra, Pl. 17e, fig. 1.
1844 Reeve, Lovell A. Conch. Icon., Mi-
Gra, IPL, NO, tie, 7G
1856 Wood, W. Index Testaceologicus,
Supplement Pl. 3, fig. l2a.
1874 Sowerby, G. B. Thes. Conch., Mi-
tra, Pl. 13, fig. 199 (only).
Mitra coronata Lamarck, 1811 (see Plate 29,
figure 2):
Encyclopédie Méthodique, Pl. 371,
figs. 6a, 6b.
1839 Kiener, L. C. Icon. Coq. Viv., Mi-
tra, Pl. 18, fig. 60a (only).
1841 Kuster, H. C. Conchylien-Cabinet,
Mitra, Pl. 26, figs. 5, 6.
1844 Reeve, Lovell A. Conch. Icon., Mi-
tra, Pl. 14, figs. 104a, 104b.
1856 Wood, W. Index Testaceologicus,
Voluta, Pl. 21, fig. 146a.
1860 Chenu, J. C. Man. Conchyl., Vol. 1,
p. 193, fig. 904.
1874 Sowerby, G. B. Thes. Conch., Mi-
tra, Pl. 13, figs. 200, 201, 220.
1882 Tryon, George W., Jr. Man. Conch.,
Vol. 4, Mitridae; Pl. 44, figs. 284,
285.
1946 Edmondson, C. H. Reef and Shore
Fauna of Hawaii, p. 127, fig. h (as M.
lugubris).
Mitra tiarella A. Adams, 1851 (see Plate 29,
figure 3):
1874 Sowerby, G. B. Thes. Conch., Mi-
eel, IPL, Sy tale, Gop iP, ws, ies, Qe.
217.
Pilsbry, Henry A. Proc. Acad. Nat.
scij Phila. Vole m2 pla l2,) fign 16 (a's
M. lugubris honoluluensis).
1920
Other species in this group include Mitra
aurora Dohrn, 1861 (Plate 29, figure 4), and M.
floridula Sowerby, 1874 (Plate 29, figure 5).
Mitra aurora resembles M. tiarella, but is gen-
erally smoother, especially in the middle of the
last whorl; it is, additionally, dotted and speck-
led with white, marked with large irregular
white blotches below a white subsutural band,
and is a deep rusty-red color in live-taken spe-
cimens. It is known from Hawaii, the Cook
Islands, Tahiti, the Paumotus and the Philip-
pines.
Mitra aurora has frequently been associ-
ated with M. coronata as a subspecies, but
since it maintains constant morphological char-
acteristics throughout its range, I prefer to
restore it to its original rank as a full species.
Even if it were a subspecies, it would have to
be assigned to M. tiarella instead of M. coro-
nata.
Mitra floridula Sowerby, 1874 resembles-
M. aurora but is larger, more ventricose, with
broad spiral ribs and deeply punctured spiral
grooves. It exhibits a much coarser appear-
ance than any of the other species discussed
here. It is recorded from Japan, the Ryukyus,
and Mauritius.
Acknowledgment
I wish to thank Dr. Myra Keen for her kind
cooperation in furnishing the exact Helbling
reference, which was only alluded to in Daut-
zenberg and Bouge (1922). Dr. Keen's many
important contributions tomalacology aremore
than equalled by her gracious helpfulness to
students and fellow workers.
Page 134 THE VELIGER Vol. 4; No. 3
Literature Cited cain es
Chenu, J. C. 1841. Die Familie der Walzenschnecken (Volutacea
1859. Manuel de Conchyliologie. I: 1-507. Paris. Menke). Nirnberg. Mitra: pp. 33-150, pls. 7-17e.
Dautzenberg, Ph., & L.-J. Bouge
1922. Mitridésde la Nouvelle Calédonie et de ses dé-
pendances. Jour. Conchyl. 67: Famille Mitridae,
83-259, pl. II.
Edmondson, Charles Howard
1946. Reef and shore fauna of Hawaii. B. P. Bishop
Mus. Spec. Publ. 22, iii + 381 pp., figs. 1-223,
(family Mitridae pp. 128-130, fig. 58 e- i).
Encyclopédie Méthodique.
1797. Vers, coquilles, mollusques et zoophytes.
189-390. ‘
Helbling auf Hirzenfeld, Sebastian Georg
1779. "Beitrdge zur Kenntnis neuer und seltener Con-
chylien.'"' Abhandlungen einer Privatgesellschaft
in Béhmen (Ceska spoleénost nauk, Prague) 4: 102
to 131, 4 pls.
Kiener, L. C.
1839. Iconographie des coquilles vivantes. Vol. 4,
Mitra. 1-120, pls. 1-34. Paris.
pls.
Pilsbry, Henry A.
1920. Marine mollusks of Hawaii.
Sci. Phila., 72: 309-318, pl. 12.
Reeve, Lovell Augustus
1844-1845. Conchologia Iconica.
genus Mitra. pls. 1-39.
Sowerby, George Brettingham
1874. Thesaurus conchyliorum. Vol. 4, Monograph of
the genus Mitra: 1-46, pls. 1-28.
Swainson, William
1820-1821. Zoologicalillustrations, Conchology, Series
1, vols. 1& 2, pts. 1-3, 72 pls.
Tryon, George W., Jr.
1882. Manual of conchology, 4(Mitridae): 106-200; pls.
32-58. Philadelphia.
Wood, W., & Sylvanus Hanley
1856. Index testaceologicus, an illustrated catalogue
of British and foreign shells. 1-234; pls. 1
to 38, suppl. pls. 1-8.
Proc. Acad. Nat.
Monograph of the
i-xx,
A Study of the Reproductive Cycle in the
California Acmaeidae (Gastropoda)
Part IV
by
Harry K. Frircuman II
(From the Department of Zoology, University of California, Berkeley, California,
and the Department of Biology,
(Plates 30,
Discussion
In Parts I-III (Fritchman, 196la, b, c) of
this paper the reproductive periods of eleven
species of the limpet Acmaea Eschscholtz 1833
have been reported for the latitude of San Fran-
cisco, California. An effort will now be made
to correlate these periods with the latitudinal
distribution of the species.
The effects of temperature on the breeding
and distribution of marine invertebrates have
long been noted and discussed. From his stud-
ies of the invertebrate fauna at Plymouth, Or-
ton (1920) concluded that the breeding season
seemed to be limited by apparently constant
maximum or minimum temperatures, or both,
Boise Junior College, Boise, Idaho)
31 and 32)
which seem to be physiological constants for
the species. In addition, he believes that the
greatest influence of temperature is at the
maximal or minimal temperature for the local-
ity investigated. Runnstrom (1927) working at
Bergen showed that the ranges of temperature
in which the development of larvae was possible
were correlated with the origins of the different
faunae producing the larvae, Thus, the northern
forms had a much lower range of temperature
for development than did the southern faunal
components. In addition, he found that the ear-
liest developmental stages were most affected
by the temperatures, the larvae and the adults
becoming increasingly less sensitive to tem-
perature conditions. Further work by Runn-
strom (1929) demonstrated that breeding sea-
Vol. 4; No. 3
THE VELIGER Page 135
ee oh TS
sons of the North Sea invertebrates were cor-
related with their geographic origins, the
southern forms being summer breeders in the
North Sea; the northern forms, winter breeders.
In a later paper, Runnstrom (1936) concludes
that stenothermic animals can spread only as
far as the reproductive adaptations to tempera-
ture permit, the limiting temperature being that
at which the early developmental stages suc-
cumb. There will, of course, be a fringe of
greatest adult extension which is maintained by
immigrant larvae, but where successful repro-
duction cannot occur. The species can be ex-
tended beyond these temperature boundaries
only by the development of physiological races
adapted to different temperature conditions.
Hutchins (1947) briefly reviews the chief
contributions to the knowledge of temperature
and distribution of marine organisms. He pro-
poses that four types of temperature zonation
can be recognized which account for the distri-
bution of marine life. The limits of these
distributions are maximal or minimal tempera-
tures at which (1) the adults succumb; (2) re-
production becomes impossible; or (3) and (4)
combinations of (1) and (2), one boundary being
fixed by adult survival, the other by failure to
reproduce. He has applied this system with
considerable success to the worldwide distri-
bution of Mytilus edulis and Balanus balanoides
and to the distribution of the fauna of the east-
ern coast of the United States.
Hutchins and Scharff (1947) have plotted
calendar month isotheres and isocrymes for
intervals of five degrees Fahrenheit. Such iso-
theres and isocrymes are isotherms which con-
nect points of maximal and minimal monthly
temperatures. In these compilations nominal
calendar months are the basis of calculations
and not the coldest or warmest 30-day periods.
These authors, therefore, refer to these iso-
therms as calendar month isotheres and calen-
dar month isocrymes. Using these data the
reproductive seasons of Acmaea were studied
with attention to the geographic ranges of the
various species.
The species ranges used are largely those
of Test (1937), amended where information is
available. Two species, Acmaea digitalis Esch-
scholtz, 1833, and A. asmi (Middendorf, 1847),
are listed by her as occurring on the Revilla-
gigedos Islands, Mexico, although no reports
of expeditions thereto substantiate this (Strong
and Hanna, 1930). The southern limits for these
species are, therefore, given as the points of
their most southern continental occurrence.
The southern limit of A. triangularis (Carpen-
ter, 1864) is vaguely given by Test as the Gulf
of California, while Keen (1937) specifically
lists it as 33°N. Latitude, which is here
adopted. On the basis of specimens in the col-
lection of the California Academy of Sciences,
the range of A. insessa (Hinds, 1842) is extend-
ed to Cape San Lucas, Lower California. The
range of A. mitra Eschscholtz, 1833, has re-
cently been extended by Stohler (1959) to Point
Santo Tomas, Lower California, or to the lati-
tude of 31° 35'N. The animals here, however,
are subtidal in 40 to 150 feet of water. Ac-
maea funiculata (Carpenter, 1864), being sub-
tidal, is omitted as is also A. paradigitalis
Fritchman, 1960, for which the range limits are
presently uncertain. In many cases the north-
ern limits of ranges are listed as the Aleutian
Islands or other islands of the Alaskan Penin-
sula, the latitude of which may be 55° or less.
In such cases the northern latitudinal limit will
be given as 60°N. Latitude, the most northern
point of the southern coast of Alaska. Plate 30
shows the ranges of the California acmaeids in
degrees of North Latitude; Plates 31 and 32
show the ranges in relation to isotheres and
isocrymes as set up by Hutchins and Scharff
(op. cit.). When the ends of the ranges do not
coincide with specific isotheres or isocrymes,
these limiting temperatures are approximated
by interpolation with latitude.
Acmaea persona Eschscholtz, 1833
Geographic range: Shumagin Island and
Yakutat, Alaska, to Monterey, Califor-
nia.
Latitudinal range:
Isothere range: 50°- 60° F.
Isocryme range: 35°-55° F,
(sO) S377 IN], Tee:
(10°)
(20°)
Although this high intertidal species has an
extensive geographic range, its range in rela-
tion to maximal temperatures encountered is
quite short (Plate 31). The reproductive cycle
of this species correlates well with its geo-
graphic range as a northern form. At the lati-
tude of study it is reproductively active from
mid-October through March or April and sexu-
ally indeterminate from May to mid-October.
Although the upswing in gonad activity begins in
October, it is not until November that large
percentages of ripe animals are found. Simi-
larly, it is not until late April or early May that
significant percentages of indeterminate animals
are found (Plate 15, PartII). It would appear, then,
that the temperatures encountered in April and
November are critical ones for this species.
Page 136
THE VELIGER
Vol. 4; No. 3
As indicated in Table I, Part I, the mean water
temperature for April is 54.7° F. and that for
November, 55.9°F. Thus, 55°F. is adopted
as a critical temperature for Acmaea persona.
It is important to note that, despite Runn-
strom's (1927) demonstration that adults are
less sensitive to temperature than larval forms,
it is the adult sensitivity which determines the
reproductive period of Acmaea persona, since
the gonads are completely nonfunctional during
the summer months. Thus, although the larvae
may be very sensitive to temperatures in ex-
cess of 55° F. or thereabouts, it is never a
question of their susceptibility to temperature
that determines the southern limit of the range
since no larvae are produced above 55° F. If
this is taken as the maximum at which A. per-
sona spawns and this value is compared with
the species range on the isocryme plate, it is
seen that the southern range is terminated at
the 55° isocryme. Acmaea persona, then, ap-
pears to be restricted in its southern distribu-
tion by the maximum temperature at which
reproduction is possible, this being a function
of the adult physiology.
Acmaea fenestrata cribraria Carpenter, 1857
Geographic range: Coal Harbor, Unga Is-
land, Alaska, to Cayucos, San Luis Obis-
po County, California.
Latitudinal range: 60°- 35.5° N. Lat.
Isothere range: 50°-60° F. (10°)
Isocryme range: 35°-56.5° F. (21.5°)
As was the case with Acmaea persona, the
period of reproductive activity of this species
can be correlated directly with its northern
distribution. Plate 16, Part II, shows that re-
productively active animals are found only from
September through April and during the sum-
mer the population is either spawned or sexu-
ally indeterminate. The upswing in August and
September represents partially ripe animals,
fully ripe forms not appearing until October and
November. This pattern of activity does not
bear close relation to sea temperatures pre-
vailing during the assumed critical months of
April and August, 54.7° and 59.8° respectively.
However this may be, the redevelopment of the
gonads begins during the period of the maximum
yearly temperature, at least two months prior
to that seen in A. persona. This difference
may result from the different intertidal posi-
tions of the two species, since A. fenestrata
cribraria occurs only in zone two and lower
while A. persona is strictly a zone one animal
facing slightly increased heating and desicca-
tion. Or the reproductive difference between
the two species may reflect the somewhat
greater warm water tolerance of A. f. cribra-
ria. Unlike A. persona, a correlation between
the maximum temperature at which spawning
occurs and the distribution of the subspecies in
relation to isocrymes cannot be made. The
data indicate that 54° F. must be about the max-
imum spawning temperature for A. f. cribraria
at the latitude of study. Yet, as shown on Plate
32, its range extends almost to the 57° iso-
cryme producing a discrepancy of 3° F. It ap-
pears, nevertheless, that this subspecies is re-
stricted in its southern distribution by the
maximal temperature at which the adult repro-
ductive mechanism can operate.
Acmaea scutum Eschscholtz, 1833
Geographic range: Aleutian Islands to San
Pedro, California
Latitudinal range: 60°- 34° N. Lat.
50°-65.5° F. (15.5°)
357s R37 IP, (23°)
Isothere range:
Isocryme range:
The population of this moderately euryto-
pic limpet is reproductively active throughout
the year (Plate 17, Part II). The gonads rede-
velop subsequent to each spawning and no sexu-
ally indeterminate period is found. There is,
however, a tendency toward a summer latent
period at which time the gonad is fully develop-
ed but spawnings are reduced or stopped. In
the absence of data on this species from more
southern points, it is difficult to suggest what
effect increasing temperature has on its repro-
duction. Does it, at the southern end of its
range, become sexually indeterminate or does
it develop gametes without shedding them?
Orton (op. cit.) believes that the production of
eggs capable of being fertilized is sufficient
evidence of breeding activity. Thorson (1946)
disagrees with this and points out that in cer-
tain regions the temperatures are such that the
animals can ripen their gonads but spawning
will not occur. As will be shown later, this is
presumably the case for two species of Acmaea
investigated and may also be true for A. scut-
um. It is interesting to note that in 1949, 1950,
and 1951 the first fall spawning occurred ata
time when water temperature was at or very
near its maximum. The larvae are thus prob-
ably able to survive at sea temperatures near
60° F. and the range of A. scutum in relation to
isocrymes substantiates this, since the range
of the species terminates at the 58° isocryme.
Acmaea mitra Eschscholtz, 1833
Geographic range: Chirikoff Island, Alas-
ka, to Punta Santo Tomas, Lower Cali-
fornia
Latitudinal range: 60°- 31° 35'N. Lat.
[FritcHMAN] Plate 30
THE VELIGER, Vol. 4, No. 3
aq, souXsoosy yjuoy repugen
Da90aDg “Ww
Rme—=|— oc =
D.iqvas 7
— ase ere
ees ee Z se
ees Cee ae)
Eis aa
aesd Pisce a
California Aeaadlae
Ranges in Latitude
noysoual fy
|
sympa “y
J, soxypos] Uo, repuryen
%
apniney yon saeig9q
alifornia
Conception,
California
Ballenas, Bay
Cape San Lucas
California
California
Aleutian Islands
Halfmoon Bay
Baja C
[FRITCHMAN] Plate 31
Cayucos,
(ee ee |
2 gg
Eee ee a
an see ee
wusD “Wy
smignysut *
I
[oe ees
pupigua {Wy (aad
California Acmaeidae
Ranges in Isotheres
°
~
A. Sef9y OST YWUOPY Iepuzery)
Tut Vewicer, Vol. 4, No. 3
Baja California
Vol. 4; No. 3
THE VELIGER nee
ee BE TSI
50°- 65.5° F.
35°- 58° F.
(15.5°)
(23°)
Isothere range:
Isocryme range:
On Plate 10, Part I, it can be seen that this
species is a winter breeder, spawning when the
sea temperature is at or near its minimum.
The population studied shows a slow redevelop-
ment of the gonads and a fully ripe condition
may be reached as early as July, although
spawning may not occur until December or Jan-
uary. The maximal temperature at which
spawning occurred was in February, 1952, when
the surface waters were about 53.5° F. If this
value is tentatively taken as the upper limit for
spawning and is compared with the isocryme at
the southern end of the range (Plate 32), itis
seen that the latter figure is approximately
58° F., or 4.5° in excess of the postulated max-
imal temperature. This discrepancy is proba-
bly explained by the fact that Acmaea mitra be-
comes subtidal somewhere on the central or
southern California coast and thus may remain
in temperature ranges suitable to it.
It is interesting to observe that, in Acmaea
mitra, the sensitive point in its reproductive
physiology is its spawning mechanism and the
activity of the gonad seems to be unimpaired
unless the slow redevelopment after spawning
is so regarded. From the above discussion, it
seems certain that A, mitra is restricted in its
southern distribution by the maximal tempera-
ture at which spawning can occur.
Acmaea pelta Eschscholtz, 1833
Geographic range: Aleutian Islands to San
Diego, California
Latitudinal range:
Isothere range: 50°- 66° F.
Isocryme range: 35°-59° F.
60°- 33° N. Lat.
(16°)
(24°)
This eurytopic species is reproductively
active throughout the year and is known to spawn
at sea temperatures from 48.5° F. to 60.0° F.
(Plate 9, Part III). Furthermore, there is no
long summer period during which the gonads
are ripe but unspawned as is found in Acmaea
scutum. This may be related to the slightly
greater southern range of A.pelta, which itself
seems to be uncertain since Keen (1937) lists
the species from 19° N. Lat. and Test (1937),
while giving San Diego as a southern termina-
tion, suggests that it may extend even further.
When a species is reproductively active
throughout. the year, it is difficult to account
for the distribution in terms of temperatures
encountered since no critical temperatures
have been defined. The best that can be done
with Acmaea pelta is to accept tentatively the
highest known spawning temperature, 60.0° F,
and compare the species range with this parti-
cular isocryme, If a relationship is present,
the two may be expected to coincide within a
few degrees. Plate 32 shows that the range of
this species terminates at the 59° F isocryme,
only 1° from the highest determined spawning
temperature. It may thus be assumed that A.
pelta, like other northern species, is restricted
to the south by the maximal temperature at
which it can spawn. Whether the gonad shows
a summer indeterminate phase in the more
southern latitudes is not known.
Acmaea digitalis Eschscholtz, 1833
Geographic range: Aleutian Islands to Gua-
dalupe Island, Mexico
Latitudinal range: 60°- 30° N. Lat.
50°-69° F. (19°)
35°-61° FF. (26°)
Isothere range:
Isocryme range:
Disagreement exists concerning the south-
ern limits of this species which is listed by
Keen (1937) as 19° N. Lat. and by Test (1937)
as Guadalupe Island, Mexico. Specimens in the
collection of the California Academy of Sciences
from Guadalupe Island are unmistakably Ac-
maea digitalis, and this is the most southern
point from which they have recorded the spe-
cies. It does not occur on Socorro Island as
listed by Dali (1921).
The data on reproduction from the popula-
tion at Moss Beach (Plate 13, Part III) suggest
that the species is active at this latitude
throughout the year. The maximum tempera-
ture at which spawning occurred was in June-
July, 1951, the mean temperature for the peri-
od being 56° F. The isocryme for this temper-
ature does not even closely agree with the
southern limit of the range, the 56° isocryme
falling approximately at 36°N. Lat. or just
south of Monterey, California. This continuous
breeding is to be expected when the area of
study lies near the center of the range of tem-
perature tolerance of a species as is the case
for Acmaea digitalis.
The data from the Rockaway Breakwater
population of Acmaea digitalis (Plate 12, Part
III) are perhaps more reliable because interpo-
lation, as explained previously, was not neces-
sary to localize spawning periods. The mini-
mum temperature at which spawning occurred
was 49.5° F. during February, 1950. This value
is approximately the isothere for the northern
limit of the species. Despite the extreme con-
Page 138
ditions of temperature and desiccation which
the breakwater population must endure, there
can be no doubt that it is an adequate habitat
for the adult limpets. Were this not so, the
continual collections which were made there
over a period of three years and which totaled
more than 9'000 individuals would certainly
have drastically reduced the population. This
did not happen, and thus the settlement of lar-
vae and their growth must have been continuous.
It seems that the extreme environmental con-
ditions affected primarily the activity of the
gonads and resulted in the sexually indetermi-
nate summer phase. If it be assumed that this
condition is realized throughout the most south-
ern ranges of this species, it is clear that the
time of development of the adult gonad restricts
breeding times to the period most favorable to
the larvae. Presumably the ultimate response
of this species, like A. persona, to increasing
temperatures would be a nonfunctioning gonad.
Acmaea limatula Carpenter, 1864
Geographic range: Tomales Bay, Califor-
nia, to Tres Marias Islands, Mexico
Latitudinal range: 38°-21° N. Lat.
The open coast range terminates approxi-
mately at Santa Cruz, California, a dis-
tribution for which the following temper-
ature ranges are given.
Isothere range: 60°-80° F.
Isocryme range: 55°-75° F.
(20°)
(20°)
This southern species is given a southern
limit by Keen (1937) of 24° N. Lat. which is
near the southern tip of Lower California.
However, Strong and Hanna (1930) found it on
the Tres Marias Islands, and the California
Academy of Sciences possesses an unmistakable
specimen from Socorro Island of the Revilla-
gigedos. The southern range is therefore ex-
tended to 21° N. Lat. According to Test (1937)
the species is numerous at Pacific Grove, Ca-
lifornia, but, except for an occasional speci-
men, is not found north of there or Santa Cruz
on the open coast. Pockets of warm water do
exist which support the species, one of which is
Tomales Bay. Keen (1937) lists the northern
limit as Puget Sound, but it is not reported by
Curtiss (1941) nor has the author collected it
despite extensive collecting there and in the San
Juan Islands.
In Tomales Bay the so-called subspecies
Acmaea limatula moerchii Dall, 1878, spawns
once a year during September at a time when
water temperatures are at a maximum of about
60° F. on the open coast (Plate 10, Part III).
THE VELIGER
Vol. 4; No. 3
This is followed by a period when spawned and
sexually indeterminate animals are found and
when the gonad is slowly reformed. This cycle
correlates well with the distribution of the spe-
cies, since it terminates just northward of the
60° F. isothere (Plate 31). It appears that this
species is limited in its northward distribution
by the temperature required for its spawning.
The gonad can redevelop even during the cold-
est part of the year as shown by the decrease
in the numbers of indeterminate animals during
the winter months. But, as previously indicat-
ed, the gonads of this population do not develop
the extreme turgor seen in most of the other
species suggesting that the temperatures do not
favor maximal reproductive activity.
Acmaea asmi (Middendorf, 1847)
Geographic range: Sitka, Alaska, to San
Quintin Bay, Lower California
Latitudinal range: 57°- 30° N. Lat.
Isothere range: 55°-69° F. (14°)
Isocryme range: 38°-61° F. (23°)
Acmaea asmi, as would be expected of a
species with a southern distribution, is repro-
ductively active during a period from April to
September or October and then enters a period
of reproductive latency until the following
March when gonad redevelopment begins (Plate
11, Part III). The minimum water temperature
at which it is known to have spawned at Moss
Beach is 52°F. in April, 1951. This figure
agrees reasonably well with the 55° F. isothere
which forms the northern limit of the species
near Sitka. The reproductive cycle of this spe-
cies appears to be controlled as much by the
effect of low temperature on the gonad as by
the requirement for a certain spawning temper-
ature since, unlike A. limatula, its gonad is
nonfunctional during the winter months. It
would be interesting to know if the animals be-
come sexually indeterminate during the winter
in Alaskan waters.
Acmaea scabra (Gould, 1846)
Geographic range: unknown point in North-
ern California or southern Oregon to
Cape San Lucas, Lower California
Latitudinal range: unknown point near 42°
to 23> N, Lat.
Isothere range: 58° (California-Oregon
border) to 80° F. (22°)
Isocryme range: 50° (California-Oregon
border) to 68° F. (18°)
This species is listed by Keen (1937) as
Santo Tomas,
Baja California
Cape San Lucas,
Baja California
Revillagigedos
Halfmoon Bay
California
» Aleutian Islands
Point
Sitka, Alaska
® Vancouver, B. C)
[FritcHman] Plate 32
sf —[
aed ee eae
Se. Lee ee
DTP Sars
Pe ae es
oe ee syiqujsua “y
a See ee
er ee ere
eevaaeenn pupsgua “fy ESaal Apousoual JER,
Seed cdese Sa,
3 & S
J, Sesy}OST YUOJ, FepuateD
Oo. = em we ee
Ranges in Isotheres
oO
ss)
ine)
7
oO
E
1S)
<
(99)
=
S
bet
o
o—
—
isy}
12)
THE Veticrr, Vol. 4, No. 3
Vol. 4; No. 3
THE VELIGER Page 139
a OD
extending to Puget Sound. Like Acmaea lima-
tula, it is not reported by Curtiss (op. cit.) nor
have I found it in these waters. Furthermore,
it has been rarely, or doubtfully, reported from
the vicinity of Coos Bay, Oregon. Hence, the
California-Oregon border is taken asa tenta-
tive northern limit.
Acmaea scabra does not show any repro-
ductive phenomenon which can be correlated
with a critical temperature. Consequently, few
conclusions can be drawn between its repro-
duction and distribution. It is, at this latitude,
a year around breeder (Plate 14, Part III). This
is quite strange considering that the area of
study lies probably within 2° F. of the northern
limit of the species in isotheres. It would be
expected that it would be a late summer breeder
like A. limatula while, in actuality, it spawns at
sea temperatures ranging from 50° to 57° F.
However, two items are worth noting. One is
that A. scabra did not develop the extreme tur-
gor of the gonad which characterizes the typi-
cally ripe limpet, a condition also seen in A.
limatula, and which may be correlated with the
colder waters of Central California. The sec-
ond is that A. scabra from the Rockaway Break-
water does not undergo a summer indeterminate
period as does A. digitalis which may reflect
the adaptation of A. scabra to the higher tem-
peratures of its southern distribution. How-
ever, it must be admitted that A. scabra, be-
cause of its possession of a home scar on the
rock and its inclination toward horizontal sur-
faces, would not receive the severe desiccation
of A. digitalis.
Acmaea insessa (Hinds, 1842)
Geographic range: Sitka, Alaska, to Cape
San Lucas, Lower California
Latitudinal range: 57°- 23° N. Lat.
Isothere range: 55°- 80° F. (25°)
Isocryme range: 38°-68° F. (30°)
It is unfortunate that the reproductive data
on this very eurythermal southern species are
rather confused as a result of this limpet's
tendency to spawn readily. Such data as are
available indicate that spawning occurs through-
out the year, spawnings having been recorded
from November, 1949; January and February,
1950; April and May, 1950; and in July, 1950
(Part II). Spawnings in the laboratory have oc-
curred at temperatures ranging from 10° C.to
19°C. (50°-66° F.), the latter value being at
least 6° F. in excess of sea temperatures nor-
mally encountered at this latitude. Plate 31
shows that the area of study lies well to the
north of the midpoint of the isothere range of
the species, a fact that cannot be reconciled
with its presumed continuous breeding which,
theoretically, should be confined to the summer.
As with Acmaea scabra, the lack of an estab-
lished critical temperature prevents establish-
ment of relationships between reproduction and
distribution.
Summary
The above analysis reveals that the repro-
duction of the acmaeids is affected in several
ways as the various species approach the ends
of their geographic ranges.
1, The production of gametes may be entirely
suspended at unfavorable periods of the
year. For northern forms like Acmaea
persona, A. fenestrata cribraria, and A.
digitalis, this occurs during the summer
months. In A. asmi, a southern species,
gonad activity is curtailed during the win-
ter. In these cases the physiology of gam-
etogenesis appears to be the factor limiting
the species range, since at more extreme
latitudes the gonad would presumably be-
come entirely nonfunctional.
2. Another limiting response is that of the
spawning reactions of the adult. In such
species as Acmaea mitra and A. limatula,
and perhaps also A. pelta and A. scutum,
the limits of the range are correlated with
the spawning temperature required by the
species. Thus, although the gonad may be
fully ripe, as is the case with A. mitra,
spawning may not occur because of lack of
proper water temperature stimuli.
3. A third response is the reduced activity of
the gonad of southern forms in more north-
ern latitudes. This has been observed in
Acmaea scabra and A. limatula where the
turgor, characteristic of the ripe gonad,
is seldom attained.
The data accumulated here all suggest that
the ranges of the limpets studied are deter-
mined by the failure of the reproductive mech-
anism and that death of the adults is probably
not involved except, perhaps, in extreme north-
ern waters where freezing may occur.
Acknowledgment
The author desires to express his appreci-
ation for the many helpful suggestions made by
Drs. J. W. Durham, Frank Pitelka, and Ralph
I, Smith, all of whom read the original manu-
script. I also would like to thank Dr. Willard
Hartman for his help and encouragement and
Drs. R. Stohler and Cadet Hand for guidance in
its publication.
Page 140
Literature Cited
Curtiss, Ruby M.
1941. An ecological and taxonomic survey of the Ac-
maeidae of the Northwest Pacific area. M. S. the-
sis, Univ. of Washington.
Dall, William Healey
1921. Summary of the marine shellbearing mollus
of the northwest coast of America, from San Diego,
California, to the Polar Sea, mostly contained in
the collection of the United States National Museum,
with illustrations of hitherto unfigured species.
U.S. Nat. Mus. Bull, 112: pp. 1-217, pls. 1-22.
Fritchman, Harry K.
196la. A study of the reproductive cycle in the Califor-
nia Acmaeidae (Gastropoda). Part I. Veliger 3(3):
57-63.
1961b. A study of the reproductive cycle in the Califor-
nia Acmaeidae (Gastropoda). Part II. Veliger 3(4):
95-101. °
1961c. A study of the reproductive cycle in the Califor-
nia Acmaeidae (Gastropoda). Part Ill. Veliger 4(1):
41-47.
Grant, Avery R.
1937. A systematic revision of the genus Acmaea
Eschscholtz, including consideration of ecology
and speciation. Ph. D. thesis, Univ. California,
Berkeley.
Hutchins, L. W.
1947. The bases for temperature zonation in geogra-
phical distribution. Ecol. Monogr. 17 (3): 325-335.
Hutchins, L. W., & M. Scharff
1947, Maximum and minimum monthly mean sea
temperatures charted from the "World atlas of sea
surface temperatures". J. Mar. Res. 6 (3); 264 to
268.
THE VELIGER
Vol. 4; No. 3
Keen, A. Myra
1937. An abridged check list and bibliography of west
North American marine mollusca. Stanford Univ.
Press. 87 pp.
Orton, J. H.
1920. Sea temperature, breeding and distribution in
marine mammals. J. Mar. Biol. Assoc. N. S. 12:
339-366.
Runnstrom, S.
1927. Uber die Thermopathi der Fortpflanzung und
Entwicklung mariner Tiere in Beziehung zu ihrer
geographischen Verbreitung. Bergens Mus. Aar-
bok, Nat. Rekke 2: 1-67.
——, ——
1929. Weitere Studien uber die Temperaturanpas sung
der Fortpflanzung und Entwicklung mariner Tiere.
Bergens Mus. Aarbok, Nat. Rekke 3: 1-46.
———,,
1936. Die Anpassung der Fortpflanzung und Entwick-
lung mariner Tiere an die Temperaturverhdltnisse
verschiedener Verbreitungsgebiete. Bergens Mus.
Aarbok, Nat. Rekke 10: 1-36.
Stohler, Rudolf
1959. Range extensions of some west N. A. marines.
Nautilus 72 (4): 127-130.
Strong, A. M., & G. Dallas Hanna
1930a. Marine mollusca of the Revillagigedos Islands,
Mexico, Proc. Calif. Acad. Sci., 4th ser., 19(2):'
7-12.
—p — & .
1930b. Marine mollusca of the Tres Marias Islands,
Mexico. Proc. Calif. Acad. Sci., 4th ser., 19(3):
13-22.
Test, A. R. see Grant, Avery R.
Thorson, Gunnar
1946, Reproduction and larval development of Danish
marine bottom invertebrates. Medd. Komm. Dan.
Fisk. -og Havunders., ser. Plankton, 4: 523 pp.
Revision of Some Hawaiian Mitrid Species
by
JEAN M. CaTE
Conchological Club of Southern California, Los Angeles 7, California
(Plates 33, 34 & 35)
There are several Mitrids from Hawaiian
waters which have proved difficult to identify,
particularly some of the deep-water species
dredged during the 1959 Pele Expedition. The
research necessary to identify some of these
forms has brought to light certain problems of
synonymy which, to the best of my knowledge,
have never been discussed.
The only readily available illustrated ref-
erences pertaining specifically to the Hawaiian
marine gastropod fauna are Pilsbry (1920), Ed-
mondson (1946), Morris (1952) and Tinker
(1958); none of these works covers all of the
Hawaiian species. The checklist of Hawaiian
Mitra published in the Hawaiian Shell News
(1956) indicates about 100 species recorded
from Hawaii, and a second list shows 22 addi-
tional species which are undescribed Dall man-
uscript names, In the four works cited, Ed-
mondson illustrates five of these 100 species,
Morris eight, and Tinker 33, although Tinker
has also included illustrations of several un-
identified species.
Vol. 4; No. 3
THE VELIGER
Page 141
EE aN Ee ee ee gn
I found on further investigation that some
of the Pilsbry Hawaiian species, supposedly
endemic, resembled very closely certain long-
known Indo-Pacific species. The Pilsbry type
figures are rather unsatisfactory, and it was
not possible, based on them alone, to work out
a definite answer as to whether all of the Pils-
bry names are valid or merely synonyms of
earlier-named species. It was therefore my
good fortune when a vacation in the northern
California area last spring coincided with the
loan of some of the Pilsbry types from the
Academy of Natural Sciences of Philadelphia to
the Paleontology Department of Stanford Uni-
versity. At that time I had the privilege of ex-
amining this material at first hand and the op-
portunity of comparing it with similar species
in the Stanford Collection. I am greatly indebt-
ed to Dr. Myra Keen for directing my attention
to this material, for offering the use of the
Stanford Paleontology Department's collection
and library, and for providing me with excellent
photographs of many of the Pilsbry type speci-
mens through the facilities of her department.
The photographs made it possible for me
to pursue the study of these 13 species after
returning home, comparing them at leisure
with similar specimens in my own collection
and with the literature pertaining to worldwide
Mitridae. I found that certain of the species
were undoubtedly synonyms, while others were
valid and some of these apparently endemic to
Hawaii. I shall report my findings on each of
the 13 species separately here, following the
same order in which they appeared in the orig-
inal publication:
Mitra kamehameha
Mitra thaanumiana
Mitra ostergaardi
Mitra lugubris honoluluensis
Mitra ticaonica vagans
Mitra olivellaeformis
Mitra langfordi
Mitra emersoni
Mitra waikikiensis
Vexillum thaanumi
Vexillum xenium
. Vexillum micra
13. Vexillum turben kanaka
DANANARWNYE
See
SS —
Ln)
To explain some points to be raised later,
I should like to mention that I have had the ad-
ditional good fortune to have on loan examples
of each of the species in the very large and well
catalogued Mitra collection of Mr. Ditlev Thaa-
num of Honolulu. This valuable collection,
certain maps and charts of remote Pacific
atolls, and his catalog lists have very gener-
ously been made available to me by Mr. Thaa-
num to use in my current work with the Mi-
tridae, and have proved invaluable in identifying
some species not in my own collection. Mr.
Thaanum's field notes provide the only ecologi-
cal information available on some of the spe-
cies,
Many other species of Hawaiian Mitra have
been sent me, both on loan and as gifts, by Mr.
Clifton S. Weaver of Honolulu; among these are
many which were dredged during the Pele Ex-
pedition of 1959, Still others have been sent by
several of the enthusiastic Hawaiian collectors
including Evelyn Gage, R. P. Gage, Jr., and
Elizabeth Harrison. To all these kind people I
wish to express my gratitude for their willing-
ness to help, both with specimens and informa-
tion.
The collection of Professor Jens M. Os-
tergaard of Mountain View, California, was ex-
amined in the course of the research for this
paper, as his collection contains paratype and
topotype material of some of the Pilsbry spe-
cies, and was a part of the nucleus for Dr.
Pilsbry's work.
With the single exception of Mitra olivel-
laeformis, one of the more common of the 13
species, the types of all of the Pilsbry Hawaii-
an species discussed here were collected by
either Ostergaard, Thaanum or D. B. Lang-
ford. Mr. Langford was Mr. Thaanum's close
personal friend and fellow collector for over 50
years; his collected material is now in the Thaa-
num Collection which, together with the Os-
tergaard Collection, probably constitutes the
bulk of the remaining Pilsbry type material not
presently in the Academy of Natural Sciences
of Philadelphia.
After the preliminary manuscript for this
paper had been completed, an unexpected trip
to the east coast made it possible to verify my
views by examining the collections at the Amer-
ican Museum of Natural History, the Academy
of Natural Sciences of Philadelphia, and the
United States National Museum. This trip
broadened the scope of the study by giving me
the opportunity to see additional specimens of
the 13 species and comparative material, as
well as to record further locality data; without
exception, the findings remained the same as
my original conclusions. I am grateful to Mr.
William E. Old, Jr., Miss Virginia Orr and
Dr. J. P. E. Morrison of these institutions for
expediting my work during the brief stops at
each place.
Page 142
While at the National Museum in Washing-
ton, I was able to see the 22 unpublished Hawa-
iian Mitra species of W. H. Dall; some of these
were found to be synonymous with the Pilsbry
species and are so recorded here. Pending
validation, the Dall manuscript names listed in
the Hawaiian Shell News are nomina nuda and
are presently unavailable.
1, Mitra kamehameha Pilsbry, 1920 = Mitra
ustulata Reeve, 1844
[Plate 33, figure 1; holotype of M. kame-
hameha]
This species was described froma single
decollate subfossil specimen from the dredger
dump at Honolulu Harbor. Pilsbry mentioned
its similarity to Mitra ustulata, the chief dif-
ference being in the lengths of the apertures,
which he described as "less than half the total
length of ustulata, more than half in kameha-
meha'"', As may be seen in the accompanying
photograph of the type specimen of M. kameha-
meha, the aperture and spire would be of about
equal lengths in that specimen if the spire were
complete. This photograph closely matches the
Reeve type figure of M. ustulata (Conch. Icon.,
sp. 89), the Sowerby figure of that species
(Thes. Conch., fig. 227), and the Tryon figure
(Man. Conch., Pl. 35, fig. 51).
Specimens of Mitra ustulata from the Viti
Isles, Aden and the Paumotus in the American
Museum of Natural History match it also, ex-
cept that the surface sculpturing of a few of
these specimens is rougher and deeper than in
the Pilsbry holotype of M. kamehameha; this
might be explained by the fact that the subfossil
shell is smooth through being worn.
A juvenile subfossil specimen of this spe-
cies was collected by R. P. Gage, Jr. in 1960.
THE VELIGER
Vol. 4; No. 3
Three specimens in the Academy of Natural
Sciences of Philadelphia were collected by Dr.
C. M. Burgess in 25 to 50 feet of water, inside
coral heads at Nanakuli, Oahu. Mitra ustulata
is rare in Hawaii, and to the best of my know-
ledge is represented in collections there large-
ly by dead specimens,
Mitra ustulata somewhat resembles M. ab-
batis Dillwyn, 1817 and M. ignobilis Reeve,
1844 which are also found uncommonly in Ha-
waii. They differ in that M. ustulata is propor-
tionately longer, slimmer, and smoother, with
M. abbatis bearing strong spiral sculpture, a
spire considerably longer than the aperture,
and a more obese body whorl, Mitra ignobilis
is intermediate between the other two species
in its sculpture; that is, ignobilis is more
strongly sculptured than ustulata but less so
than abbatis; the spire is shorter than the last
whorl, and the last whorl is more ventricose.
The colors and markings of these species are
similar.
2. Mitra thaanumiana Pilsbry, 1920 = Mitra
coffea Schubert and Wagner, 1829 (Mitra
fulva Swainson, 1832)
[Plate 33, figure 2; type specimen of M.
thaanumiana]
There is little doubt that these two species
are synonymous; Pilsbry's types and written
description of Mitra thaanumiana are identical
with Schubert and Wagner's type figure and de-
scription and Swainson's figure of M. fulva, as
well as with examples of M. coffea from vari-
ous parts of the Pacific.
Mitra ambigua Swainson, 1832 and M. ful-
va Swainson, 1832 have been considered syn-
onymous by some authors, which could explain
the error in identification frequently encoun-
Explanation of Plate 33
Figure 1: Mitra ustulata REEVE 1844. Ventral and dorsal aspects of the holotype of Mitra kamehameha Pirssry, 1920
(ANSP No. 46 753, height 59.4 mm.) Figure 2: Mitra coffea ScHuBERT & WaGNER, 1829. Holotype and paratype
of Mitra thaanumiana Pitssry, 1920 (ANSP No. 46810, height of left-hand specimen 53.4 mm.) Figure 3: Mitra
ostergaardi Pitssry, 1920. Ventral and dorsal aspects of the holotype (ANSP No. 46 770, height 43.5 mm.)
Figure 4: Mitra tiarella A. ADAMS, 1851. Holotype and paratype of Mitra lugubris honoluluensis PitsBRY, 1920
(ANSP No. 46 797, height of left-hand specimen 23.0 mm.) Figure 5: Mitra ticaonica REEVE, 1844. Holotype and
paratype of Mitra ticaonica vagans Pitspry, 1920 (ANSP No. 46 790, height of right-hand specimen 25.0 mm.)
(Figures 1 to 5 photographs by Perfecto Mary, courtesy of Stanford University)
Figure 6: Mitra olivaeformis Swainson, 1821. Dorsal aspects of typical variants (left to right): 1. Resembles the
unfigured punctate holotype of Mitra olivellaeformis Pitssry, 1920. From Kauai. 2. A typical smooth specimen, from
Oahu. 3. With produced and punctate spire, from Okinawa. 4. A slender specimen, from the Caroline Islands.
5. An obese specimen, from Kauai. 6. With mucronate spire, from Kauai.
(Photograph by Victor Duran, Scientific Photographic Laboratory, University of California; twice natural size.)
(Specimens 1 to 4 ex Cate Collection; 5 & 6 ex Thaanum Collection.)
Tue VELIGER, Vol. 4, No. 3 [Cate] Plate 33
Figure 2
Vol. 4; No. 3 THE VELIGER Page 143
a ere AB ENTAY
tered in connection with these species. The
original figures of both appear on the same page
(Swainson, Zoological Illustrations, Part 2,
Plate 30) and are obviously separable, M. am-
bigua having a wide whitish band a short dis-
tance below the sutures and stronger punctate
spiral sculpture, besides being more attenuate
anteriorly; M.fulva is a smoother shell with no
color pattern. However, the name coffea was
applied to this species three years earlier than
Swainson's name and thus has priority.
The error in identification and its applica-
tion to the present species is seen in a note in
the species catalog of Mr. Ditlev Thaanum,
listing this entry:
"Mitra ambigua Swainson, #1519. Keau-
kaha, Hilo, Hawaii. Th., Coll., '03. Un-
der rocks. Rare. Id. by Dr. W. H. Dall
as Mitra ambigua Swainson. This is type
lot of M. thaanumiana Pilsbry."'
Mitra coffea is chestnut brown inside and
out, with five or six white folds on the columel-
la and many white denticles on the outer lip.
The spiral sculpturing is weakly punctate
throughout; the lip is constricted above, effuse
below; the canal is short and sharply recurved
abaxially. Its range includes Madagascar, the
Philippines, Marianas, Guam, Samoa, Tahiti
and Polynesia. Considered rare in Hawaii, it
has been collected in 10 to 15 feet of water off
Waikiki, Oahu (Dranga and Thaanum), and un-
der rocks at Keaukaha Ponds, Hawaii (Thaa-
num).
3, Mitra ostergaardi Pilsbry, 1920
[Plate 33, figure 3; syntypes of M. oster-
gaardi; Plate 35, figure c; ex. Cate coll.]
_Pilsbry erred in comparing this species
“with Mitra ambigua; he apparently accepted
Tryon's synonymizing of that species with M.
fulva Swainson (= M. coffea Schubert and Wag-
ner, 1829). It differs, however, in its slimmer
and more cylindrical form, its pure white aper-
ture, and a smooth adapical edge to the labrum
(whereas that of M. coffea is denticulate for its
entire length). Mitra ostergaardi has fewer
punctate spiral grooves than M. coffea, strong-
est on the upper whorls, shoulder and neck of
the shell, with the periphery nearly smooth.
The types of Mitra ostergaardi are faded
subfossil specimens from the dredger dump at
Honolulu, and therefore differ to a certain ex-
tent from live-collected specimens. Freshly
collected shells are a deep chocolate brown
with a faint indication of a paler narrow subsu-
tural band. Some have tiny speckles of pure
white on the lower half of the last whorl. The
aperture is constricted above, effuse at the
base, as in M. coffea; there are five columel-
lar folds.
Living specimens have been collected in
recent years in deep water at Oahu, Hawaii: in
30 to 40 feet of water under dead coral, Kailua
Bay (C. S. Weaver), dredged in 100 feet on a
sand and coral rubble substrate off Waianae
(Pele Expedition), in 25 to 35 feet on top of the
reef at Nanakuli (C. M. Burgess, ANSP speci-
mens), in 45 feet in coral heads off Makua (C.
M. Burgess, AMNH specimens). Mitra oster-
gaardi has also been reported in dredgings
from the lagoon on the north side of Kwajalein
Atoll in the Marshall Islands in 1943 (R. V.
Dietrich).
The type of Mitra ostergaardi is at the
Academy of Natural Sciences of Philadelphia.
Mitra pararhodia Dall (MS.)is probably the
Same species. USNM specimens bearing this
label were collected at Sand Island, Midway, by
Bartsch in 1907.
4, Mitra lugubris honoluluensis Pilsbry, 1920
= Mitra tiarella A. Adams, 1851
[Plate 33, figure 4; type specimens of M.1,
honoluluensis]
This subspecies was erroneously assigned
by Pilsbry to Mitra lugubris Swainson, 1822. It
is more closely related to M. coronata La-
marck, 1811 (non Helbling, 1779), a species
similar to M. lugubris in some ways and fre~
quently confused with it.
Mitra tiarella A. Adams, 1851 was first
figured by Sowerby (1874); it differs from M.
coronata in that its crenulations are lightly
tipped with white instead of occupying a wide,
solid-white subsutural zone, and M. tiarella
bears also a narrow pale band below the crenu-
lations which does not appear in M. coronata.
In Hawaii Mitra tiarella has been collected
off Maili, Oahu (Thaanum), in 20 to 30 fathoms
off Waikiki, Oahu (Thaanum), under dead coral
in 30 to 40 feet (C. S. Weaver), dredged in 100
feet in coral rubble and sand off Waianae, Oahu
(Pele Expedition), and many subfossil speci-
mens are known from the dredger dump in Ho-
nolulu. The species is also known from the Sulu
Sea (F. Dayrit), Japan (Thaanum), and the Ryu-
kyu Islands (Thaanum, A. Scott). With so much
confusion regarding the identification of speci-
mens labelled Mitra coronata and Mitra lugu-
Page 144
THE VELIGER
Vol. 4; No. 3
bris, due possibly to errors in Sowerby (1874)
and Tryon (1882) (see page 132), I prefer not to
list further locality records beyond those spe-
cimens which I have personally seen and regard
as unquestionably M. tiarella.
Mitra assimilis Pease, 1867 is very simi-
lar, but Pease in his description says ''there is
not the slightest evidence of crenation at the
sutures'' and his holotype will have to be stud-
ied before a decision may be made as to whe-
ther it is conspecific with M. tiarella.
Mitra crassula Dall (MS.) is the same spe-
cies (USNM No. 339'899).
5. Mitra ticaonica vagans Pilsbry,'1920 = Mi-
tra ticaonica Reeve, 1844
[Plate 33, figure 5; type specimens of M.t.
vagans]
Pilsbry's basis for separating his subspe-
cies from the typical Philippine Mitra ticaonica
was that the Hawaiian specimens were smooth-
er on the last whorl and less deeply grooved on
the spire. After examining many museum spe-
cimens from various other points in the Pacific
from Samoa to Tawi-Tawi (including the type
locality, the island of Ticao), I believe the two
geographical forms are not separable morpho-
logically. The subspecies was described from
two worn subfossil shells which could be ex-
pected to exhibit less sculpturing than live-col-
lected specimens; any other discernible differ-
ences appear to be merely normal variations
within the species. Some specimens are short-
er and more obese than others and a few show
more pronounced sculpturing, but these differ-
ences appear throughout the range of the spe-
cies with no more emphasis on one locality
than another.
Live-collected specimens taken by Thaa-
num on the reefs at Kewalo, Oahu are choco-
late brown and have fine spiral striae through-
out, these becoming somewhat more coarse at
the neck of the shell. The aperture is also dark
brown, with four white columellar folds and a
strong, pale brown nodule in the throat. Speci-
mens are frequently so encrusted that only the
aperture may be seen,
Mitra pupiformis Dall (MS,) is the same
species (USNM No, 338'028).
6, Mitra olivellaeformis Pilsbry, 1920 = Mitra
olivaeformis Swainson, 1821
[Plate 33, figure 6; ex. Cate coll.]
Since Pilsbry's species, admittedly very
similar to Mitra olivaeformis, is found living
in the same populations with that species in
Hawaii and elsewhere throughout the Pacific, it
would appear that M. olivellaeformis is not de-
serving of specific rank. Its only difference
lies in the presence of punctate spiral sculpture
on the last whorl which is not usually present in
the typical olivaeformis. This punctate charac-
ter is frequently seen in other Mitrid species;
in M. pontificalis Lamarck, 1811 and M. pa-
palis Lamarck, 1811, for example, the upper
whorls are entirely punctate while the last
whorl is smooth. Therefore, young specimens
would appear punctate while adults of the same
species would not. Mitra olivaeformis seems
to be one of the more variable of Mitrid spe-
cies, and the punctate sculpture noted by Pils-
bry merely one of its variations.
Other even more obvious differences within
the species may be seen, with large series to
study. Some specimens have a very strongly
produced spire, while others are mucronate; in
some the upper whorls are swollen and bulbous,
while the usual form of an average specimen
has sloping shoulders. The fact that all forms,
smooth and punctate, produced and mucronate,
swollen and gradate, slender and obese, are
found in the same populations within the range
of the species seems ample reason to consider
them merely intrapopulation variants and to re-
tain the original name Mitra olivaeformis for
all of the variations. Even if Pilsbry's M. oli-
vellaeformis were considered a separate taxon,
it should be made a subspecies of M. olivae-
formis, as its differences are too minor to jus-
tify full species rank for this form.
The photographs on Plate 33 (figure 6) il-
lustrate some of the extremes mentioned above,
The first shell resembles Pilsbry's unfigured
type of Mitra olivellaeformis, while the second
is a typical smooth specimen.
Mitra olivaeformis is recorded from local-
ities throughout the entire central Pacific area,
reported by Garrett (1880) as ''very abundant
and gregarious in sand inside the reefs". It has
been collected in depths ranging from one or
two feet to 30 fathoms (off Waikiki, Oahu; Thaa-
num).
7. Mitra langfordi Pilsbry, 1920= Mitra peasei
Dohrn, 1860
[Plate 34, figure 1; syntypes of M. lang-
fordi]
Pilsbry compared this species with Mitra
granatina Lamarck, with M. gracilis Reeve, M.
Vol. 4; No. 3
THE VELIGER
Page 145
Se
filosa Born, M. bernardiana Philippi, and M.
circula Kiener, but unfortunately overlooked
Dohrn's M. peasei, which is identical with Pils-
bry's type specimens of M. langfordi.
Mitra peasei is a white shell with a buff
periostracum. Its aperture is china-white, with
a pink tinge in young specimens. It is encircled
with slender, rather beaded brown spiral
cords, the brown color on these sometimes in-
terrupted. There is fine cancellate sculpture
in the interstices, caused by the intersection of
a fine median spiral cord with equally fine axi-
al impressed lines. The surface of the shell
frequently shows irregular brown clouding, es-
pecially in young specimens. The protoconch
is rosy-pink, conical-multispiral, deviated, ho-
meostrophic, and unlike many Mitrid species,
the protoconch persists in most specimens into
adulthood.
Two notations in Mr. Thaanum's catalog on
the page devoted to Mitra peasei are of interest
here; the first refers to a set of shells collect-
ed in Maui:
"No. 1710, e. Id. by Dr. W. H. Dall, 1920,
as Mitra peasei Dohrn."
The second note reads as follows:
"No. 1710. Honolulu, Oahu, harbor en-
trance in 6-10 fathoms of water. D. B. L.,
Coll., 1915. This is type lot of Pilsbry's
Mitra langfordi."'
With patronymic names being applied to
new species so freely today in honor of persons
who have made no real contribution to malacol-
ogy, it is regrettable that the name of D. B.
Langford should be lost in this case through an
oversight. His carefully documented collections
throughout the Pacific area over a period of
some 50 years contributed greatly to the sci-
ence.
The type locality of Mitra peaseiis Aus-
tralia.
8. Mitra emersoni Pilsbry, 1920
[Plate 34, figure 2; type specimens of M.
emersoni; Plate 35, figure d; ex. Cate coll.]
"close-
This species, as Pilsbry stated, is
ly related to Mitra crenifer (sic) Lamarck" (=
M. clathrus Gmelin, 1790). His basis for sepa-
rating the Hawaiian form is that it is ''smaller,
the vertical impressed lines about twice as far
apart". The relative size is a poor criterion
for establishing a new species; however, it may
be readily seen with a much larger series than
Pilsbry had available that there is an appreci-
able difference in its proportions from those of
the typical M. clathrus from Japan and the
Philippines. Mitra emersoni is a slimmer spe-
cies with a more produced spire, averaging
nearly two millimeters narrower at the peri-
phery than M. clathrus specimens of compara-
ble length.
Additional differences include the further-
apart vertical impressed lines noted by Pils-
bry, a fainter and less distinct though very
similar color pattern, and the prominence and
beaded appearance of the raised spiral bands.
In Mitra clathrus the surface is uniformly can-
cellate; in M. emersoni the interstices between
the spiral bands exhibit additional, finer orna-
mentation.
The color of Mitra emersoni is white under
a buff periostracum; nucleus and base are pink,
with a pink blush extending onto the columella
in fresh specimens. The interior of the aper-
ture is white, the pattern and intermittent sec-
tions of the spiral threads chestnut brown.
Mitra emersoni is a deep-water species,
recorded by Thaanum in from 25 to 75 feet off
Laniupoko Camp and Mala Bay, West Maui; off
Waikiki, Oahu in 20 to 30 fathoms; in Waialua
Bay, Oahu in 12 to 15 fathoms; at the entrance
to Pearl Harbor in 6 to 10 fathoms (type lot);
and in 150 feet in sand off the entrance to Pearl
Harbor during the Pele Expedition. Further
records in the U. S. National Museum indicate
its occurrence at Koloa, Kauai; at Pearl and
Hermes Reef; and at Sand Island, Midway.
The type specimens of Mitra emersoni are
at the Academy of Natural Sciences of Philadel-
phia.
9. Mitra waikikiensis Pilsbry, 1920
[Plate 34, figure 3; type specimens of M.
waikikiensis; Plate 35, figure a; ex. Cate
coll, ]
Mitra waikikiensis is a distinctive species,
easily separable from any other Miters in the
Hawaiian fauna. It is characterized by its no-
dulose sculpture throughout, and ornamented
with two brown bands from which the glossy
white nodules protrude. In specimens still re-
taining the periostracum, the nodules are very
dark blackish-brown, as is the protoconch; the
remainder of the periostracum is a cinnamon-
brown color, The nodules are arranged in rows
Page 146
in a very precise manner, suggesting the nodu-
lose sculpture seen on the parietal wall of Dis-
torsio anus (Linnaeus, 1758).
Pilsbry's species was described from two
specimens dredged by D. B. Langford off Wai-
kiki in 35 to 50 fathoms. Both are small shells,
the larger of the two measuring only 12.5 mm.
In addition to the type specimens, I have seen
nine of comparable size in the Thaanum Col-
lection and approximately 35 live-taken and
dead specimens which were dredged during the
Pele Expedition. Except for eight very large
shells dredged in 60 fathoms in Keehi Lagoon,
Oahu, all of these were also approximately the
same small size. The Keehi shells, one of
which is illustrated on Plate 35 (fig. a), are
twice as large as the type specimens, ranging
from 23 mm. to 29 mm. in length.
With over 45 specimens at hand, it was
possible to make certain observations about the
species which were not noted by Pilsbry in his
original description. In addition to the unusual-
ly large size of the eight examples noted above,
other observations include a variability in the
number of columellar folds; Pilsbry mentioned
three folds, the lower one quite small. The
majority of shells in the series studied possess
four folds, but one is seen to have five and only
four have three folds. The protoconch is coni-
cal, glossy-white except as noted in those spe-
cimens still retaining the epidermis, and con-
sists of about De; to 3 whorls.
Mitra waikikiensis is apparently a deep-
water species; the shallowest record among my
notes is off Mala Bay, West Maui, dredged by
Thaanum in 25 to 75 feet; otherwise, it is re-
corded only from depths of 100 to 360 feet on
sand, or a sand and coral rubble substrate
(Pele Expedition), and from 60 to 500 feet
(Thaanum).
THE VELIGER
Vol. 4; No. 3
Mitra waikikiensis bears a striking re-
semblance to the type figure and description of
M. loricata Reeve, 1844 (Conch. Icon., sp. 174)
and to Sowerby's figure of the same species
(Thes, Conch., Plate 10, fig. 148), but since I
have not seen examples of typical M. loricata
it seems best at this time not to conclude that
the two species are synonymous.
The type specimens of Mitra waikikiensis
are in the Academy of Natural Sciences of Phi-
ladelphia.
Mitra colpophila Dall (MS.) is the same
species (USNM No, 337'991).
10, Vexillum thaanumi Pilsbry, 1920
[Plate 34, figure 4; ex. Cate coll.; Plate
35, figure e; ex. Cate coll. ]
One of the most beautiful of all the Mitrid
species, Vexillum thaanumi is apparently known
only from Hawaii, and of rare occurrence even
there. It is a deep-water form, most of the
known specimens having been dredged in from
25 to 300 feet of water. The type lot was
dredged off Waikiki, Oahu, in 200 to 300 feet by
D. B. Langford in 1916; V. thaanumi has also
been collected in 4 to 12 fathoms off Mt. Lihau;
in 28 to 43 fathoms in the Auau Channel; in
Waialua Bay; off West Maui; at the entrance to
Pearl Harbor, and in Keehi Lagoon.
The photographs of this species on Plate
34 do not illustrate the type specimens, which
are somewhat encrusted and unattractive. The
shells figured here were collected during the
Pele Expedition in from 100 to 150 feet at the
entrance to Pearl Harbor, Oahu, and are from
the Cate’Collection.
Pilsbry's original description of this spe-
cies is as follows:
Explanation of Plate 34
Figure 1: Mitra peasei Dourn, 1860. Ventral and dorsal aspects of syntypes of Mitra lang fordi Prssry, 1920 (ANSP
No. 46 805, height of left-hand specimen 35.3 mm.) Figure 2: Mitra emersoni Pirspry, 1920. Holotype and para-
type (ANSP No. 46 804, height of left-hand specimen 29.4 mm.) Figure 3: Mitra waikikiensis Pispry, 1920. Syn-
types (ANSP No. 46 788, height of left-hand specimen 12.5 mm.) Figure 4: Vexzllum thaanumi Pirssry, 1920. Ex
Cate collection (height of left-hand specimen 23.3 mm.) Figure 5: Vexillum xenium Picssry, 1920. Type and para-
type (ANSP No. 116983, height of left-hand specimen 18.0 mm.) Figure 6: Vexillum micra Pirspry, 1920. Type
figure of holotype and paratype (ANSP No. 116986, height of left-hand specimen 7.5 mm.) Figure 7: Vexillum
lurben (REEVE, 1844). Ventral and dorsal aspects of the holotype of Vexillum (Idiochila) turben kanaka Pussry, 1920
(ANSP No. 46 763, height 23.0 mm.) Figure 8: Vexillum turben (REEVE, 1844). Typical specimen from Mauritius.
(Figures 1 to 3 and 5 to 8, photographs by Perfecto Mary, furnished through the courtesy of Stanford University)
(Figure 4, photograph by Victor Duran, Scientific Photographic Laboratory, University of California.)
THE VELIGER, Vol. 4, No. 3 [Cate] Plate 34
Figure 2
Vol. 4; No. 3
THE VELIGER
Page 147
"Vexillum thaanumi, n. sp.
"The shell is fusiform, rather slender,
white, with a cinnamon brown band below
the periphery. At the periphery there are
narrow vinaceous or brownish spots be-
tween the ribs, surmounted by a continu-
ous, cinnamon line. On the penult whorl
this line is median.
"Sculpture of vertical ribs weakening to-
wards the base, the intervals with short
impressed lines in a spiral direction; on
the last whorl there are 11 ribs and about
19 spirals, exclusive of those on the si-
phonal fasciole. On the penult whorl there
are 10 or 11 impressions in an interval.
"The aperture is slightly pink tinted
within, lirate in the throat. Five columel-
lar plaits, the lower one very small.
"Length 25.3, diameter 8, aperture 12
mm; 10 whorls.
"Off Waikiki, Oahu, in 200-300 feet. D.
B. Langford.
"Mitra interstriata Sowerby (Thes.
Conch. fig. 392) resembles this species
somewhat in color, but it is wider, con-
tracted more above the more prominent si-
phonal fasciole, and has a wider aperture."!
The largest specimen I have seen (in the
U. S. National Museum Collection) measures
31.6 mm. in length; the smallest adult shell, in
the same collection, is just under 8 mm. long.
The type specimens are in the Academy of
Natural Sciences of Philadelphia.
11, Vexillum xenium Pilsbry, 1920
[Plate 34, figure 5; type specimens of V.
xenium; Plate 35, figure b; ex. Weaver
coll. ]
Vexillum xenium is rare in collections and
is presently known only from Hawaii. A few
live specimens were taken during the Pele Ex-
pedition, dredged on a sand and coral substrate
at 17 fathoms at the entrance to Keehi Lagoon,
Oahu.
Vexillum xenium is said to resemble V.
approximata (Pease, 1860), but is a distinct spe-
cies,according to Dr. Alison Kay of the Univer-
sity of Hawaii (personal communication) who
has seen and photographed the unfigured Pease
holotype.
The original description of Vexillum xeni-
um follows:
"Vexillum xenium n. sp.
"The shell is fusiform, white with a
chestnut band traversed by several paler
spiral lines, below the periphery, two or
three paler interrupted lines above it on
the summits of the ribs only, and a few
widely spaced blackish-brown spots below
the suture, on the ends of some of the ribs.
The first three whorls are also deep brown.
Sculpture of smooth, longitudinal ribs, 22
on the last whorl, equal to their interstices,
the latter marked with short impressions
in spiral series, 6 on the penult whorl in
each interval; base spirally grooved over
ribs and intervals forming about 4 spiral
series of tubercles. Two obliquely spiral
cords are more prominent just above the
siphonal fasciole. Aperture shorter than
the spire, the throat with 9 thin beaded li-
rae. Columella with 5 thin plaits.
"Length 18, diameter 7.5, aperture 8.4
mm., 10 whorls.
"Off Waikiki, Oahu, 25-50 fathoms. D.
B. Langford.
"Turricula approxima (sic) Pease (P, Z.
S. 1860) is described as convexly angulated
at the sutures and with 4 plaits; it seems
therefore to be a different species."'
The type specimens of Vexillum xenium are
in the Academy of Natural Sciences of Philadel-
phia.
12. Vexillum micra Pilsbry, 1920
[Plate 34, figure 6; type specimens of V.
micra |
Vexillum micra is a tiny species, just over
+ inch long. It is rare in collections, possibly
because of its small size and deep-water habi-
tat.
This species is not represented on the col-
or plate (Plate 35) which illustrates the other
five valid Pilsbry species from Hawaii, due to
its unavailability for photographing in color.
However, the black-and-white photograph on
Plate 34 (figure 6) which was furnished through
the courtesy of Stanford University, is the type
figure of Vexillum micra, as no previous illus-
tration has been published.
Page 148
THE VELIGER
Vol. 4; No. 3
Pilsbry's original description follows:
"Vexillum micra, n. sp.
"The shell is fusiform, vinaceous tawny
with a band at the periphery and another at
the base of burnt umber. Sculpture of
many vertical rounded ribs, about 30 on the
last whorl, the intervals with wide, low
spiral cords separated by impressed lines,
of which there are 4 on the penult whorl;
base with spiral cords. Aperture colored
like the outside. Columella with four
plaits. The embryonic shell is long-conic,
of about 34 smooth whorls.
"Length 6.5, diameter 2.7, aperture 3
mm.; 53 post-embryonic whorls,
"Off Waikiki, Oahu, in 15-50 fms.
Langford.
D. B.
"One specimen is a little stouter and
lacks the lower brown band."
The type specimens are in the Academy of
Natural Sciences of Philadelphia.
13. Vexillum (Idiochila) turben kanaka Pilsbry,
1920 = Vexillum (I.) turben (Reeve, 1844)
[Plate 34, figure 7: a type specimen of V.t.
kanaka; figure 8: typical V. turben (Reeve)]
This subspecies was described from three
subfossil dredger-dump shells, after apparent-
ly having been compared with only one other
example of Vexillum turben (Reeve, 1844). To
quote Pilsbry's description, ''on comparison
with a specimen of Reeve's species they show
certain differences of shape which seem to be
of racial significance". The
Plate 34 include one of the Pilsbry types (figure
7) and a second ANSP specimen of V. turben
illustrations on
from Mauritius (figure 8). The only outstand-
ing difference between the two seems to be in
size, the Honolulu specimen being considerably
larger than that from Mauritius and exhibiting
proportionately larger dimensions throughout.
Museum specimens from Mauritius, Lubang,
Maeiret and Midway were examined in New
York, Philadelphia and Washington and com-
pared with several Hawaiian examples, and no
morphological differences could be discerned
which would justify subspecific rank for the
Hawaiian specimens.
Vexillum turben is the type species of Pils-
bry's sectionIdiochila, a small section contain-
ing only one or two additional species, one of
these as yet undescribed. Vexillum turben is a
very distinctive species bearing little resem-
blance to most other forms of Mitra. It is
obesely pyriform, closely ribbed, finely stri-
ated in the interstices, with deep sutures and
strong spiral grooves at the base. Its labrum
is closely and finely lirate, the columella bears
five or six strong folds and a parietal callus.
The abapical edge of the outer lipis evidently a
weak point in this species, as all the specimens
I have seen give the appearance of having been
broken away at this point; Reeve mentions this
characteristic also in the original description.
The typical color of a living specimen is un-
known to me, as I have seen only pale yellow
subfossil dredger-dump shells in Hawaiian col-
lections and similarly faded specimens from
other parts of its range in museum collections.
The type locality is the Philippine Islands. It is
a rare species worldwide.
To sum up the conclusions derived from
this study, I list here what I consider the pres-
ent status of the 13 species described from Ha-
waii by Pilsbry in 1920:
Explanation of Plate 35
Five of the Six Valid Species of Mitra and Vexillum Described by Pitssry in 1920 as from Hawaii
a: Mitra waikikiensis Pitssry, 1920.
Dredged in 60 fathoms on sand, Keehi Lagoon, Oahu (Pele Expedition,
August 1959). Height 27.2 mm. Cate Collection No. HD 754.
b: Vexillum xenium Pitssry, 1920. .
Endemic to Hawaii. Dredged in 17 fathoms in sand and coral. Keehi
Lagoon, Oahu (Pele Expedition, April 1959). Height 23.0 mm. ex Weaver Collection.
c: Mitra ostergaardi PitsBRy, 1920.
Fell out of broken-up dead coral in 65 to 70 feet. Makua, Oahu. Height
28.7 mm. leg. C. S. Weaver, October 1958. Cate Collection No. H 164.
d: Mitra emersoni Pitssry, 1920.
Dredged in 25 fathoms in sand, off entrance to Pearl Harbor, Oahu (Pele
Expedition, April 1959). Height 24.7 mm. Cate Collection No. HD 501.
a
Vexillum thaanumi PitsBry, 1920.
Endemic to Hawaii. Dredged in 16 to 25 fathoms in sand, entrance to
Pearl Harbor, Oahu (Pele Expedition, April 1959). Height 22.5 mm. Cate Collection No. HD 489.
THE VELIGER, Vol. 4, No. 3 [J. CaTE] Plate 35
Kodachromes by Victor DuRAN
ADAA
wo
Vol. 4; No. 3
THE VELIGER
Page 149
Mitra kamehameha Pilsbry, 1920
2. Mitra thaanumiana Pilsbry, 1920
3. Mitra ostergaardi Pilsbry, 1920
4, Mitra lugubris honoluluensis Pilsbry, 1920
5. Mitra ticaonica vagans Pilsbry, 1920
6. Mitra olivellaeformis Pilsbry, 1920
7. Mitra langfordi Pilsbry, 1920
8. Mitra emersoni Pilsbry, 1920
9. Mitra waikikiensis Pilsbry, 1920
Vexillum thaanumi Pilsbry, 1920
Vexillum xenium Pilsbry, 1920
Vexillum micra Pilsbry, 1920
Vexillum turben kanaka Pilsbry, 1920
MERELY soulel sa
= Mitra ustulata Reeve, 1844
coffea Schubert and Wagner, 1829
fulva Swainson, 1832
pararhodia Dall (MS.)
tiarella A. Adams, 1851
= Mitra crassula Dall (MS.)
= Mitra ticaonica Reeve, 1844
pupiformis Dall (MS.)
olivaeformis Swainson, 1821
= Mitra peasei Dohrn, 1860
Valid
Probably valid; cf. Mitra loricata Reeve,
1844
Valid; endemic
Valid; endemic
Valid; endemic
= Vexillum turben (Reeve, 1844)
Acknowledgment
I wish to express my deep appreciation
once again to all those persons and institutions
mentioned in the early part of this paper; also,
to Dr. Theodore Downs and Miss Joan Troesch
of the Los Angeles Museum for the loan of per-
tinent material, to Dr. Harald Rehder of the U.
S. National Museum for permission to cite the
Dall manuscript names, and especially to
Crawford Cate for his willing and invaluable
help with the more tedious aspects of the study.
Literature Cited
Adams, Arthur
1851. Descriptions of fity-two new species of the ge-
nus Mitra from the Cumingian collection. Proc.
Zool. Soc. London, Pt. 19; 132-14).
Dietrich, Richard V., & Percy A. Morris
Nautilus 67 (1): 16.
1953. Mollusks from Kwajalein.
Edmondson, Charles Howard.
1946. Reef and shore fauna of Hawaii. B. P. Bishop
Mus. Spec. Publ. 22, iii + 381 pp., figs. 1-223,
(family Mitridae: pp. 128-130, fig. 58 e- i).
Garrett, Andrew
1880. Catalogue of the Polynesian Mitridae.
3: 1-73.
J. Conch.
Hawaiian Shell News
1956. Check list of Hawaiian Miters. H.S.N. 4 (4): 38
to 39.
1959. Check list of Hawaiian gastropods, Mitridae.
H.S.N. 11 (7): suppl. pp. 23-28.
Martini, Friedrich Heinrich Wilhelm, & J. H. Chemnitz
1829. Neues systematisches Conchylien Cabinet, fort-
gesetzt von Gotthilf Heinrich von Schubert und Jo-
hann A. Wagner. Vol. 12, Abtlg. 1. Nurnberg,
Morris, Percy A.
1952. A field guide to shells of the Pacific coast and
Hawaii; Mitridae pp. 194-196, pl. 39.
Pilsbry, Henry A.
1920. Marine mollusks of Hawaii.
Sci. Phila., 72: 309-318, pl. 12.
Reeve, Lovell Augustus :
1844-1845. Conchologia Iconica.
genus Mitra. pls. 1-39.
Sowerby, George Brettingham
1874. Thesaurus conchyliorum. Vol. 4, Monograph of
the genus Mitra: 1-46, pls. 1-28.
Swainson, William
Proc. Acad. Nat.
Monograph of the
1820-1821. Zoologicalillustrations, Conchology, Series
1, vols. 1& 2, pts. 1-3, 72 pls. ,
—,——
1829-1833. Zoological illustrations, the shells, Series
2.
Tinker, Spencer Wilkie
1958. Pacific sea shells (rev. ed.).
family. pp. 140-160, illus.
Tryon, George W., Jr.
1882. Manualofconchology, 4(Mitridae): 106-200; pla
32-58. Philadelphia.
The Miter shell
Page 150
THE VELIGER
Vol. 4; No. 3
Preliminary Report on Growth Studies in Olivella biplicata
by
RUDOLF STOHLER
Department of Zoology, University of California, Berkeley 4, California
(Plate 36)
While making observations on Olivella bi-
plicata (Sowerby, 1825) for the studies reported
on previously (Stohler, 1952, 1960), it became
apparent that an estimate of the life span of this
species was not possible on the basis of distinct
size classes of the shells. As was indicated
(op. cit., 1960), Olivella exhibited mating behav-
ior at every low-tide period when observations
were possible, and thus distinct and chrono-
logically separable size classes could not be
expected. However, the growth rate, as well as
the life span, was of interest to me. A possible
approach to the study of these two problems
presented itself when the populations of Olivel-
la were seen in the Flood Control Channel in
San Diego. The following facts seemed to make
these particular populations especially suitable
for an experimental approach to the problem.
The Flood Control Channel is closed to the
ocean, at least for the major part of the year,
by an extensive sand bank, yet in the study area
the water changes with each tidal cycle, al-
though the change is somewhat retarded. .The
water flows through a dike along the north side
of the area. This dike forms the boundary be-
tween the entrance to Mission Bay and the Flood
Control Channel and consists of large quarried
rocks relatively loosely piled upon each other.
Furthermore, adjacent to these large rocks
there is a relatively extensive area consisting
of small boulders and gravel, thus effectively
preventing the Olivella living in the deeper sand
bed from escaping, yet permitting free ex-
change of the water. That the ecological condi-
tions in the area are suitable for the animals is
borne out by the fact that the populations seem
to be thriving.
It will be seen, then, that it is possible to
collect some of the animals living in this re-
stricted area, measure them, mark them,
return them to their usual habitat with a fair
probability of being able to recapture a per-
and
centage of the marked animals at a later date.
It need hardly be mentioned that this approach
would not be feasible in the usual habitats where
Olivella biplicata occurs along the California
coast, namely, on the open shore.
The problem, however, was to find a suit-
able method of marking the shells. The first
approach was to file an area near the apex of
the shell so that a flat surface was obtained on
which numbers were written with a crow-quill
pen and India ink. In a preliminary test, 100
individuals were marked in this manner and re-
leased in 1958. This method of marking, how-
ever, had to be abandoned because experience
showed that the numbers became illegible after
approximately six months.
A second experiment consisted in applying
a pressure-sensitive tape on which had been
printed numbers with a specially prepared
printing ink. For further protection of the
printed number, adhesive cellulose tape was
placed over this patch. This method proved to
be unsuitable within less than three months; the
pressure-sensitive tape was lost in the water.
It was unfortunate that individually mea-
sured and marked specimens could not be ob-
served by the use of one or the other of these
methods. As an alternative method, it was then
decided to collect a large number of individuals,
measure them, and pick out a particular size
class, mark all individuals of that selected size
class, and release all captured animals. The
mark consisted of a notch filed into the whorl
just above the body whorl, opposite of the aper-
ture. Care had to be exercised in preventing
the file from cutting through the wall of the
shell, thus injuring and exposing the living ani-
mal,
On July 25, 1959, 242 Olivella biplicata,
from 11.0 to 13.0millimeters, were marked in
[StoHteR] Plate 36
Tue VELIGER, Vol. 4, No. 3
Biers
F igure I
Figure 2
BLakER, photo.
Vol. 4; No. 3
this fashion and released. On January 9, 1960,
six of these marked animals were recovered
and measured. It was found that they had at-
tained a length of 16.3 to 23.4 millimeters. The
return of only approximately three percent, in
spite of prolonged and careful search, was con-
sidered insufficient to allow making any conclu-
sions of value as to the growth rate of this spe-
cies, A much larger number of marked animals
was desired, but the mechanical difficulties of
marking them adequately with a triangular file
was a serious handicap. As a further experi-
ment, another group of Olivella was collected
in the Flood Control Channel area on July 16,
1960. Of approximately 1'500 specimens, 266
were selected in the size class 16.0 to 18.0 mil-
limeters. The marks were made with a dental
drill, and while the shell was still dry from the
friction exerted by the ,action of the drill, the
groove was covered with India ink. These ani-
mals were then returned as usual.
On November 6, 1960, 964 animals were
collected in the Flood Control Channel; of these,
five were clearly marked ones, but included
one dead shell. The measurements ranged from
18.6 to 20.25 millimeters. On January 28, 196l,
a total of 754 specimens was collected, includ-
ing seven marked ones. The sizes of these
ranged from 18.8 to 21.9 millimeters. On April
22, 1961, of 1'925 shells collected, four were of
the marked group, with a size range of 17.7 to
20.9 millimeters.
From these preliminary results, it seemed
to become apparent that Olivella is a relatively
slowly growing gastropod, and further, it
seemed plausible that a shell of 30 millimeters
or over in length might indicate a life span of
several years, However, I felt that these ex-
periments were not revealing enough, inasmuch
as they gave no information as to the possibility
of different growth rates at different stages of
development. Such information could be ob-
tained, of course, with individually marked
shells; that is, shells whose exact measurement
is known initially and which could be positively
identified on subsequent recapture. However,
as indicated above, all efforts in this direction
were unsuccessful, The use of the dental drill
for marking the shells seemed to offer a possi-
ble way out of the difficulty. On July 3, 1961,
1'350 specimens of Olivella ranging in size
from 14.0 to 24.0 millimeters were collected at
Solano Beach. On July 4 a total of 2'612 indi-
viduals ranging over the same measurements
were obtained from the Flood Control Channel;
all these shells were separated into size class-
THE VELIGER
Page 151
es of two millimeter range, i.e., 14.1 to 16.0
millimeters, etc., with one exception: the group
from 22.1 mm. on up was divided into two
classes, namely, 22.1 to 23.0 and 23.1 to 24.0
mm. All 3'967 individuals (1'035 marked and
2'932 unmarked) were liberated in the experi-
mental area in the Flood Control Channel.
The technique employed may be best ex-
plained by referring to Figures 1] and 2, Plate
36. The handpiece of the dental drill was se-
curely fastened in a clamp and the shell could
be pressed against the rotating drill, either
pushing it up at right angles to the longitudinal
axis (figure 1) or along an extension of the lon-
gitudinal axis (figure 2). By a combination of
marks made, it was possible to distinguish the
different size classes.
It is planned to endeavor a recapturing of
these Olivellas at three-month intervals,
it is hoped that the large number of marked
specimens, together with the diversity of size
classes marked, will make possible a reason-
able appraisal of growth rate and life span of
Olivella biplicata.
and
Acknowledgment
This study could not be successful without
the unusually generous and wholehearted coop-
eration of several people whose contributions I
wish to acknowledge here: Mr. and Mrs. Alan
H. Wolfson of San Diego, their son Arthur, and,
occasionally, various of their friends have
spent (and plan to spend in the future) many
hours in diving for the Olivella. Dr. E. W.
Fager generously made available to me the fa-
cilities of his laboratory at Scripps Institution
of Oceanography in LaJolla, where the animals
were kept alive while out of their natural habi-
tat during the days of measuring and marking.
Literature Cited
Stohler, Rudolf
1952. Studies on mollusk populations II. Nautilus 65
(5); 135-137.
——s
1960. Studies on mollusk populations: IV. Nautilus 73
(2, 3): 66-72 and 95-103; figs. 1-3.
Page 152
THE VELIGER
Vol. 4; No. 3
A New Deep-Water Anadara from the Gulf of California
by
BRUCE CAMPBELL
Conchological Club of Southern California, Los Angeles 7, California
(Plate 37, 1 Textfigure)
During the month of June, 1959, while
spending a week diving at Guaymas, Mexico, I
had occasion to visit several times my friend
Captain Xavier Mendoza who was at that time
manager for Productos Marinos, the largest
shrimp fishing organization in Guaymas. On
the day I returned home he gave me, among
several other things, a live specimen of an
Anadara that had just come off one of the
shrimp boats. That fall it was sent to Dr. Myra
Keen who reported that she knew of no other
Anadara with such a felt-like periostracum. As
there was no precise locality data available at
the time, the shell was held for confirmation of
its occurrence which has since been gained
through the dredging of odd valves at several
stations off Cabo Haro, Guaymas, Mexico.
ARCACEA
ARCIDAE
Anadarinae
Anadara GRAY, 1847
(Scapharca) Gray, 1847
Anadara (Scapharca) hyphalopilema CAMPBELL, spec. nov.
(Plate 37, Figures 1 to 8)
Shell large, thin, strongly and evenly con-
vex, markedly inflated, with the anterior end
roundly sloping ventrally; posterior end round-
ed and obliquely produced at an angle of about
75 degrees; color white internally, externally
white except for a brownish stain over the dor-
sal two-thirds of both valves. Height equal
length, with the highest portion at the posterior
third; umbones full, passing forward into small
prosogyrate beaks located at the anterior third;
posterior dorsal extremity slightly flared and
the anterior dorsal extremity unusually short-
ened; both valves with 41 squarish ribs that are
proximally solid and distally sculptured with
four to five radial riblets; narrower interspaces
concentrically sculptured by small cords about
0.75 mm. apart that slightly involve the ribs,
giving an impression of corrugations; inequi-
valved with left valve slightly overlapping right;
interiorly, the ventral margin deeply fluted by
the ends of the ribs; hinge long, straight, nar-
row with a continuous series of short fine con-
verging teeth that become larger at the extrem-
ities; midpoint of hinge less than 1 mm. thick;
cardinal area wide, covered completely by the
ligament, traversed by three to four slightly
arched lines; periostracum dark brown, with a
fine velvety texture and up to 2 mm. thick over
the posterior end.
Measurements of the holotype: Length 60
mm., height 60 mm., diameter 55 mm.
Holotype: Stanford University Paleontolo-
gical Type Collection No. 8'629.
Paratypes: A paratype will be deposited in
the U. S. National Museum, Campbell collec-
tion, Mendoza collection, and Shasky collection.
Explanation of Plate 37
Anadara (Scapharca) hyphalopilema CAMPBELL, spec. nov.
Figure 1: Exterior right valve of Paratype 1.
of left valve of Paratype 5.
type.
Figure 2: Posterior view of Holotype.
Figure 4: Interior of left valve of Holotype.
Figure 6: Interior of right valve of Paratype 4.
Figure 3: Exterior
Figure 5: Dorsal view of Holo-
Figure 7: Exterior of right valve of Holotype.
Figure 8: Exterior of left valve of Holotype.
Tue VELIGER, Vol. 4, No. 3 [CampBELL] Plate 37
CamMPBELL, photo.
Vol. 4; No. 3
THE VELIGER
Page 153
Table 1: Comparison of Holotype and Paratypes
Length Height |
Holotype
Holotype
Paratype 1
Paratype 2
Paratype 3
Paratype 4
Paratype 5
Paratype 6
Type locality: The holotype was brought in
by the shrimp boats that were at that time
working in the vicinity of Cabo Haro, Guaymas,
Sonora, Mexico. Lat. 27°50'N., Long. 40° 55!
W. June, 1959. The paratypes were trawled off
Cabo Haro, in about 50 fathoms, December 27
and 31, 1959. Collectors: B. Campbell, X. Men-
doza, T. Schowalter, and D. Shasky.
The specific name is derived from the
combination of two Greek words in order to
signify a unique anatomical feature and the
habitat of the mollusk. The Greek adjective
hyphalos means ''under the sea''; pilema is the
Greek noun for "'felt''. The combination hypha-
lopilema indicates a species with a soft perios-
tracum that resides offshore. As hyphalopile-
ma is a neuter noun in apposition, no change in
ending can be made.
Semidiameter | Rib number
(measurements in millimeters)
Semidiameter: Height
(in percent)
Discussion
The most distinctive feature that separates
this species from any of the other recent east-
ern Pacific Anadara is the soft felt-like cover-
ing displayed by the holotype and one paratype,
which resembles more the periostracum of a
Noetia with close-packed, overlapping scales,
arranged like thatch; in addition, A. hyphalopi-
lema is finely carpeted with hair. This is one
of the largest members of the subgenus Scaph-
arca that has been described from the Panamic
region. One paratype is 78 mm. in length. The
largest Scapharca is A. cepoides (Reeve, 1844)
[Olsson (1961) records a specimen with a length
of 90 mm.] with which the paratypes were con-
fused, but A. cepoides is broader and less in-
flated (see Table 2) with 32 to 35 smooth ribs
Table 2: Comparison of Species
Length | Height
(measurements in millimeters)
Anadara hyphalopilema
CAMPBELL, sp. nov.
Holotype
Paratypes (aver.)
Arca hopkinsi PitsBry
& OLssON, 1941
Type
Anadara cepoides
(REEVE, 1844)
{in Olsson, 1961]
{in Olsson, 1961]
SUPTC no. 5332
right valve
{in Keen, 1958]
Oldroyd coll. no. 1150
right valve
1 based on semidiameter
Rib number
Diameter: Length Height: Length
(in percent)
2 based on semidiameter of left valve
Page 154
and olive-brown or olive-green periostracum,
and the posterior end is not as obliquely pro-
duced, The umbonal areas are more subcentral
than those of A. hyphalopilema which are locat-
ed at the anterior third.
The species to which Anadara hyphalopile-
ma is closely related is Arca (Scapharca) hop-
kinsi Pilsbry and Olsson, 1941. This species
was described from the Canoa formation of the
Pliocene from Western Ecuador. Comment was
made in the description: ''This fine, large spe-
cies is not closely related to any living or fos-
sil ark known to us from this region.'' There
are several points of difference separating A.
hyphalopilema from Arca hopkinsi. Anadara
hyphalopilema is smaller and more inflated with
a proportionally greater height (see Table 2),
and somewhat differently shaped (see Textfigure
1). It also has 41 ribs slightly corrugated con-
centrically, 43 in two paratypes, compared with
38 radially sculptured ribs in Arca hopkinsi.
The anterior extremity is not nearly as promi-
nent as that of Arca hopkinsi.
Figure 1: Outer, tracing of Arca hopkinsi Pirssry &
OLsson, 1941 (from their plate); inner, tracing of
Anadara hyphalopilema CAMPBELL, Spec. nov.,
from paratype 3
THE VELIGER
Vol. 4; No. 3
So far as known, Anadara hyphalopilema
has only been collected from the Guaymas area
in deep water. A possible explanation why this
species has not come to light sooner is that the
Mexican fishermen would have no reason to
save dead valves coming up in the shrimp nets,
and only when a live specimen appears — such
as the holotype — would it be salvaged, as a
food item.
Acknowledgment
I wish to thank Dr. Myra Keen for her as-
sistance and helpful suggestions, and also Dr.
Donald Shasky for the loan of the unmatched
valves used as paratypes.
Literature Cited
Hertlein, Leo George, & A. M. Strong
1943. Eastern Pacific expeditions of the New York
Zoological Society. Mollusks from the west coast
of Mexico and Central America. Zoologica pt. 2,
28: 149-168, pl. 1.
Keen, A. Myra.
1958. Sea shells of tropical west America; marine
mollusks from Lower California to Colombia.
Stanford, Calif., Stanford Univ. Press; xi + 624
pp., illus.
Lowe, Herbert N.
1935. New marine mollusca from West Mexico, togeth-
er with a list of shells collected at Punta Penasco,
Sonora, Mexico. Trans. San Diego Soc. Nat. Hist.
7 (6): 15-34, pls. 1-4.
Olsson, Axel A.
1961. Mollusks of the tropical eastern Pacific, par-
ticularly fromthe southern half of the Panamic Pa-
cific faunal province (Panama to Peru). Part I.
Panamic- Pacific pelecypoda. Paleont. Res. Inst.
Ithaca, New York. pp. 1-574, pls. 1-86.
Pilsbry, Henry.A., & Axel A, Olsson
1941. A Pliocene fauna from Western Ecuador.
Acad. Nat. Sci. Phila. 93: 1-79, pls. 1-19.
Reinhart, P. W.
1943. Mesozoic and Cenozoic Arcidae from the Pacific
slope of North America. Spec. Paper Geol. Soc.
Amer., No. 47: i-xi, 1-117, pls. 1-15.
Rost, Helen
1955. A report on the family Arcidae. Allan Hancock
Pacific Exped. 20 (2): 177-249, pls. 11-16, text-
figs. 79-95. ‘
Proc,
Vol. 4; No. 3
THE VELIGER
Page 155
Two New Opisthobranch Mollusks from Southern California
by
James R. LANCE
Scripps Institution of Oceanography, University of California, La Jolla, California
(Plate 38, 8 Textfigures)
The coastal waters of Southern California
abound in numbers and species of opisthobranch
mollusks and those forms regularly inhabiting
or seasonally occurring in the intertidal regions
have become relatively well known. A distri-
butional list of all the opisthobranchs (excluding
the pteropods and cephalaspideans) occurring
between Point Conception, California, and the
Mexican border for which there have been pub-
lished descriptions sufficient to permit reiden-
tification, has been compiled by Lance (1961),
Of the 65 species listed only two (Platydoris
macfarlandi Hanna, 1951, and Tritoniopsis au-
rantia Mattox, 1955) are known exclusively from
subtidal regions. Consideration of this fact by
itself might lead to the conclusion that the local
opisthobranch fauna is almost completely repre-
sented in the intertidal zone. Such, however, is
not the case, and it merely reflects the degree
of limitation to which our observations have
been subject. Recent investigations incorpo-
rating self-contained diving gear at depths of
30 to 200 feet have shown that certain local
rocky areas are far richer in these mollusks
than are comparable regions of the intertidal.
Thus it seems probable that knowledge of the
total opisthobranch fauna along this as well as
any other coastal region is in reality quite in-
complete since the vast majority of specimens
have been collected only in regions uncovered
by the tides.
Several undescribed conspicuous forms of
opisthobranchs have recently been found in the
subtidal regions of the San Diego, California,
area and it now seems appropriate to offer de-
scriptions of two such species.
The systematic concepts of Odhner (1939)
. who achieved order out of chaos for the higher
taxa of Opisthobranchiata are almost univer-
sally accepted. Those arrangements are here
followed.
Iam very grateful to the Scripps Institution
of Oceanography, Marine Life Research Pro-
gram, for providing funds for the color plate.
Also, I appreciate Miss Joan Steinberg's kind-
ness in supplying me with specimens of Phidi-
ana from Central California.
NUDIBRANCHIA
Doridacea
EUDORIDACEA
Cryptobranchia
Doripiwar
Glossodoridinac
Cadlina limbaughi LANCE, spec. nov.
(Plate 38, Textfigures 1 to 3)
Synonymy: Cadlina spec., Lance, 1961
The body in general shape is very similar
to other species of Cadlina described from the
Pacific Coast of North America. The largest
of four specimens measured 27 mm, long and
11 mm, broad when alive; the smallest was 11
mm, long and 5 mm. broad. Thus it would
seem that the length-width ratio changes as the
animals mature. The notum is elongated with
parallel lateral margins which are slightly
more rounded in front than behind. The median
arch slopes rapidly to leave a free flat margin
laterally and anteriorly. The notum covers the
entire body in the smallest specimen, but in
larger individuals the foot extends for a short
distance beyond the posterior brim. It is warty
and somewhat gritty to the touch due to a net-
work of spicules, some of which project freely
in bundles above the surface.
The entire body is white with highly con-
trasting rhinophores and branchiae which ap-
pear black. Closer examination shows that the
‘clavi of the rhinophores are very dark reddish
brown with the distal two-thirds of the branchi-
ae being a lighter shade of the same color, The
most posterior branch of the branchial circlet
lacks this dark pigment in all four specimens.
The rhinophore stalks are white. Numerous
Page 156
THE VELIGER
Vol. 4; No. 3
Opaque white subepidermal glands occur scat-
tered over the notum. These are somewhat
larger around the lateral margins and smaller
and less numerous at the anterior and posteri-
or ends. The dorsal and ventral surfaces of
the foot and the sides of the body are sparsely
sprinkled with minute black dots which are ab-
sent on that portion of the foot extending beyond
the notum. In the smallest specimen these dots
were much reduced in number.
The foot is rather narrow, slightly expand-
ed anteriorly, and shallowly bilabiate. The
lower lip is thick, the upper lip thinner, not
notched in the middle. The tail is bluntly
rounded. :
The head is distinct with thick, triangular,
auriform tentacles grooved on their outer mar-
gins (textfig. 1). Pronounced, retractile, rhi-
nophores with deeply perfoliate clavi composed
of 11 folds in the smallest individual and 18 in
the largest, are borneon short stalks. The
sheaths of the rhinophores are low and smooth.
Figure 1: Cadlina limbaughi LANCE, spec. nov.
Antero-ventral view
There are six bipinnate and tripinnate re-
tractile branchial filaments joined at their
bases in a crescent in the smallest specimen
and seven to eight in the others. The most an-
terior filament is the largest. The conical anal
papilla is low, fleshy, and surrounded by the
crescentric branchial base. The genital aper-
ture is located about a third of the way back on
the right side of the body.
The central elements of the labial arma-
ture (textfig. 2) are slender, closely set, and
terminally bifid. Those of the margins are ir-
regularly curved and smooth on the ends. An
individual 27 mm. in length had the dental for-
mula 102 x 44-1-44, The rachidian tooth (text-
fig. 3b) is composed of four to six blunt denti-
cles of nearly equal size. Most of the first lat-
uit
Nae
A B
Figure 2: Cadlina limbaughi LANcE, spec. nov.
Elements of the labial armature
A: marginal elements
B: central elements
erals (textfig. 3c) have three inner denticles, a
large central spine, and six outer denticles.
The outward laterals become elongated, and the
30th (textfig. 3a) has a large central cusp with
14 small denticles. The outermost laterals
(textfig. 3d) are shortened, denticulate, and lack
the prominent central cusp.
Type locality: La Jolla, California. Lat.
32°52'N., Long. 117°15'W. Further distribu-
tion: Los Coronados Islands, Mexico, 32° 24!
N., 117° 14' W. Bathymetric range: Subtidal,
50 to 140 feet. Unknown from the intertidal.
aie O77
Figure 3: Cadlina limbaughi LaNcE, spec. nov.
Teeth of the radula
A: Thirtieth and thirty-first lateral teeth
B: Rachidian plate
C: First lateral
D: Outermost lateral
THE VELIGER, Vol. 4, No. 3 [Lance] Plate 38
Phidiana pugnax LANCE, spec. nov.
Kodachromes by J. R. LANcE
: a
Aes
NO ae
Vol. 4; No. 3
THE VELIGER Page 157
a EO a SOREL
The specific name limbaughi was chosen to
honor the late Conrad Limbaugh who was the
first to collect this and many other subtidal
species of opisthobranchs, and Mrs. Nan Lim-
baugh whose interest in this group has resulted
in the acquisition of previously unknown bathy-
metric distributions for many forms.
The holotype is deposited at the California
Academy of Sciences where it is registered as
Paleo. Type Coll. No. 12'396; it will be incor-
porated into the Frank Mace MacFarland Me-
morial Collection of Nudibranchs.
Remarks: Four other species of Cadlina
have been recorded from the Pacific Coast of
North America. MacFarland (1906, p. 128) has
compared the details of the radula for C. paci-
fica, C. marginata, and C. flavomaculata; and
Marcus (1961, p. 15) has “described and figured
that of the fourth species, C. sparsa. The fol-
lowing list offers distinguishing characteristics
of the color patterns of each species and will
serve to readily identify living animals.
1, Cadlina pacifica Bergh, 1880
Body bluish white; rhinophores yellow, at
least in the preserved state; nine tripinnate
gills; known only from Bergh's original
material from the Aleutian Islands, Alaska.
2. Cadlina flavomaculata MacFarland, 1905
Body dead white to yellowish white; rhino-
Pphores dark brown; six to eleven or more
yellow spots in a row on either side of the
notum; ten to twelve unipinnate and bipin-
nate gills; Vancouver Island, British Co-
lumbia, to Point Eugenia, Mexico.
3. Cadlina marginata MacFarland, 1905
The largest Cadlina so far described from
this coast; up to 80 mm. long and 40 mm.
broad; body white to yellowish white cov-
ered with low tubercles each tipped with
bright yellow; a narrow band of yellow
around the border of the notum; six bipin-
nate gills; Vancouver Island, British Co-
lumbia, to Point Eugenia, Mexico.
4. Cadlina sparsa (Odhner, 1921)
Body pale yellowish white to light tan with
the rhinophores and gills a slightly darker
shade of the same color; an uneven row of
small black or dark brown spots often with
yellow or orange centers present all around
the notum or groups of similar spots on
either side of the body; 12 unipinnate gills;
San Diego, California, to the Juan Fernan-
dez Islands, Chile.
5. Cadlina limbaughi spec. nov., Lance
Body white with opaque white spots scat-
tered over the notum; rhinophores very
dark reddish brown; six bipinnate and tri-
pinnate gills of the same color but a slight-
ly lighter shade; San Diego, California, to
Los Coronados Islands, Mexico.
NUDIBRANCHIA
Eolidacea
CLEIOPROCTA
FACELINIDAE
Facelininae
Phidiana pugnax LANCE, spec. nov.
(Plate 38, Textfigures 4 to 8)
Synonymy: Phidiana spec., Lance, 1961
The largest of 36 specimens collected
measured 63 mm. long, 10 mm. broad, and 9
mm. high when actively crawling. The speci-
men (holotype) upon which this description is
based was of average size and measured 37 mm.
long, 6 mm. broad, and 6 mm. high when alive.
The general shape of the body is eolidiform; the
sides nearly vertical with their sharply defined
dorso-lateral margins running parallel to the
blunt tail. The dorsal surface is very slightly
rounded.
The body is translucent white through which
the internal organs may be seen faintly. A line
of intense red-orange color starts on the ante-
rior side at about the midpoint of one head ten-
tacle, runs across the front of the head, and
terminates at the same point on the other ten-
tacle. A spot of this color occurs on the oppo-
site side of each tentacle. The stalk of the rhi-
nophore is white, the proximal half of the clavus
is red-orange, and the distal half of a highly
contrasting cream color. The liver diverticula
show through the cerata causing them to appear
black. The larger, more median cerata in most
of the groups are usually colored a brilliant pink
for their distal half although this may be re-
placed by a cream coloration. A pink spot with-
out defined borders is often present on the head
just anterior to the rhinophores. An irregular
line of opaque white spots runs along either side
of the body about midway up from the foot, and
a similar one, often consisting of fewer spots or
totally lacking, occurs on the midline of the dor-
sal surface starting between the rhinophores
and running posteriorly to the tip of the tail
where it appears on a distinct caudal crest.
The foot is edged with opaque white and most of
Page 158
Figure 4: Phidiana pugnax LANCE, spec. nov.
Antero-ventral view
the cerata have a spot of the same color about
three-fourths of the way up on their anterio-
lateral surface.
The foot (textfig.4) is rounded anteriorly,
bilabiate, not expanded, and extends beyond the
sides of the body for about the distal third of its
length.
The cerata start just below and slightly
posterior to the rhinophorial attachments and
occur in six major groups, the first three of
which are easily distinguishable. Each group is
B
Figure 5: Phidiana pugnax LANCE, spec. nov.
A: Inner side of mandible
B: Detail of the masticatory process
THE VELIGER
Vol. 4; No. 3
composed of numerous oblique rows attached
along a narrow dorso-lateral line of the body.
For the most part the back is free,
The head tentacles are large, measuring
13 mm. long in a 60 mm. individual. The rhi-
nophores are nonretractile and robust with
about 11 complete folds and 11 intercalary ones
on the posterior side.
The jaws (textfig.5) are horn colored and
provided with a prominent masticatory process
which bears a single row of 14 denticles. The
Figure 6: Phidiana pugnax LANCE, spec. nov.
Tooth of the radula
radula consists of a single row of teeth, 19 in
number. Each tooth is of the angular horse-
shoe shape and bears a prominent median spine
flanked by six to seven smaller lateral spines
which decrease in size towards the tip (textfig.
6). Only eight of the 19 teeth were symmetri-
cal, i.e., having either six or seven denticles
on both sides of the same tooth.
The genital papilla is located on the right
side of the body about halfway up from the foot
and below the lateral-most cerata of the poste-
rior oblique row of the first major group. The
male pore lies on the tip of a papilla and the fe-
male opening occurs as a slit immediately an-
terior. The large penis (textfig. 7) is white and
bears a black hook just: proximal to the apex.
In living animals the hook appears to be located
at the apex.
The anal papilla (textfig. 8) is a prominent
cylinder and lies far back near the right dorso-
lateral line between the third and fourth oblique
rows of the third major group of cerata.
Type locality: Point Loma, San Diego, Cal-
ifornia. Lat. 32° 40'N., Long. 117°14'W. Fur-
ther distribution: Pacific Grove, California, 36°
Vol. 4; No. 3
THE VELIGER
Page 159
Figure 7: Phidiana pugnax LANCE, spec. nov.
A: penis
B: detail of the hook
.
we OOH ya yoy,
Figure 8: Phidiana pugnax LANCE, spec. nov.
Lateral view of anal papilla within
the third group of cerata
f = foot s = side of body
38'N., 121° 55'W.; Los Coronados Islands, Me-
xico, 32° 24'N., 117°14'W. Bathymetric range:
Low intertidal to 120 feet.
The specific name pugnax was chosen to
call attention to the pugnacious nature of this
form which it often exhibits by attacking and
dismembering other eolids upon accidental con-
tact and by the violent waving motions of the
cerata which occur when the animal is dis-
turbed.
The holotype is deposited at the California
Academy of Sciences, San Francisco, Califor-
nia, where it is registeredas Paleo, Type Coll.
No. 12'397. It will be incorporated into the
Frank Mace MacFarland Memorial Nudibranch
Collection. Paratypes are available from the
author,
Literature Cited
Bergh, (Ludwig S.) Rudolph
1880. Onthe nudibranchiate gasteropod molliusca of the
North Pacific Ocean. Part I. Proc. Acad. Nat.
Sci. Phila. ; 71-132, pls. 1-8.
Hanna, G. Dallas
1951. Anew West American nudibranch mollusk.
Nautilus 65 (1): 1-3.
Lance, James R.
1961. A distributional list of Southern California opis-
thobranchs. Veliger 4 (2): 64-69.
MacFarland, Frank Mace
1906. Opisthobranchiate mollusca from Monterey Bay,
California, and vicinity. Bull. U. S. Bur. Fish.
for 1905, 25: 109-151.
Marcus, Ernst
1961. Opisthobranch mollusks from California.
Veliger 3 (Suppl., pt. I): 1-84, pls. 1-10.
Mattox, N. T.
1955. Studies on the Opisthobranchiata: I. A new spe-
cies of the genus Tritoniopsis from Southern Cal-
ifornia, Bull. So. Calif. Acad. Sci. 54(Pt. I): 8
to 13, pls. 4-5.
Odhner, Nils
1939. Opisthobranchiate mollusca from the western
and northern coasts of Norway. Kgl. Norske Vi-
densk. Selsk. Skr. No. 1, 93 pp.
Page 160
THE VELIGER
Vol. 4; No. 3
Egg-Laying in Fusitriton oregonensis (REDFIELD)
by
FAYE B. HOwARD
Research Associate in Conchology
Santa Barbara Museum of Natural History, Santa Barbara, California
(Plate 39)
Early in the morning of June 19, 1959,
while on the way from the village of Baranoff at
Warm Springs Bay, on Baranoff Island's east
coast, around the north end of the island to Sit-
ka on the west coast, a stop was made at a
small charted but unnamed island just offshore
from the larger Baranoff Island. This small
island is washed on the eastern side by Chath-
am Strait and on the north by Peril Strait (lat.
57° 20'N. and long. 134° 50'W.).
The tide was approximately -3.0 feet. The
word approximately is used advisedly because,
in all the inland passages, there is great vari-
ation from the printed tide tables. Wind direc-
tion may hasten or delay the time quoted — may
hurry or hold back the water —thus changing
the expected level.
Among all the shells observed that morn-
ing, the one that attracted the most attention
was Fusitriton oregonensis (Redfield, 1848). In
the small cove where the boat had been beached,
about 18 specimens were observed. The beach
was composed of sand and clean stones set
All the shells seen were
in a very narrow depth zone — not over 12
fairly close together.
inches variation — some just under the surface
Eggs were being depos-
ited on medium-sized stones, that is,
from six to ten inches in diameter.
every case the cluster of eggs was facing the
zenith. When this was not true, the mass was
on a smaller stone that could possibly have
been shifted by the tidal action.
and some just above.
stones
In nearly
The pictures (Plate 39, figures 1 and 2)
were taken after the stone with the spawning
female had been lifted-to the bow of the boat for
closer observation. This change of position in
no way disturbed the egg-laying process.
The female was moving, egg by egg, in a coun-
ter clockwise direction as the eggs were laid.
She would deposit one egg, proceed the precise
distance, hesitate for perhaps 20 seconds and
deposit another, and so on around her spiral
path. After she finished this process, she re-
versed her direction and, as can be seen in fi-
gure 2, extended her body across the egg mass
and in drawing back left a heavy mucus sheet
over the whole group of eggs. Other females
that had not been disturbed were also watched.
They, at the end of the mucus spreading stage,
were seen proceeding directly toward deeper
water.
No copulation was observed; but a male,
picked up within a few inches of a spawning fe-
male, showed some evidence of excitement. By
a slight tug on the lip side of the operculum, he
could be stimulated to extend the copulatory
organ. He responded to this stimulus many
times. This may suggest that mating possibly
takes place very shortly before the eggs are to
be laid.
A few egg clusters were seen at about the
spot where the previous low tide probably had
been. They were slightly higher on the beach
and were more or less obscured by sand that
had washed over them.
Naturally, one tide's worth of observation
does not answer very many of the questions that
are bound to occur to the observer. Chatham
Strait, at this collecting station, is about six
miles wide and drops off to over 400 fathoms
about a mile offshore. Do these shells ordi-
narily live much deeper and come up to spawn?
(They were not taken at any other station during
17 days of collecting.) Does the sun have a part
in the incubation process? (The shape of the
egg mass and its aiming at the position of the
noon sun for best exposure suggests this possi-
bility.) Further, the fact that.the eggs were
laid early in the series of minus tides, thus in-
suring daily exposure to sun and air for sever-
al days at the time of low tide, gives added
support to this idea. Is the mucus covering for
protection during this exposure? Only more
field work can add to the slender store of in-
formation now available.
THE VELIGER, Vol. 4, No. 3 [Howarp] Plate 39
Figure 1
Kodachromes by FayE Howarp
—
is
Vol. 4; No. 3
THE VELIGER
Page 161
Notes & News
Reinstatement
of the Specific Name
Macoma inquinata (Desuayes)
by
A. Myra KEEN
Stanford University, California
In 1934, Dr. A. E. Salisbury (Proc. Malac.
Soc. London, vol. 21, pt. 2, pp. 74-91, pls. 9-14)
published figures of the exteriors for the holo-
types of Tellina inquinata Deshayes, 1855 (type
locality, ''Columbia'', which is interpreted as
Columbia River, Oregon) and T. irus Hanley,
1845 (type locality, ''Guinea?''), which are in
the collections of the British Museum (Natural
History). He concluded that the two forms are
conspecific. Hinge illustrations not being avail-
able, we had no basis for questioning his deci-
sion as to identity, although there were small
dissimilarities in outline that might or might
not be significant. Later, someone from Japan
must have studied the British Museum materi-
al, for in 1952 Habe (Genera of Japanese Shells,
Pelecypoda 3, p. 218) proposed the generic
name Heteromacoma, with Hanley's species as
type. He cited Fragilia yantaiensis Crosse and
Debeaux, 1863 (also known as GaSstrana) in the
synonymy, as wellas ''T,"' inquinata Deshayes.
Revising the Tellinidae for the ''Treatise
on Invertebrate Paleontology,''I have been
faced with the resolution of this discrepancy.
Comparing specimens of ''G. yantaiensis" with
Salisbury's figures, I found that the outline
matched much better than for any West Ameri-
can specimens at hand. I then made up a list of
the hinge characteristics separating the West
Coast and Japanese forms and asked for a re-
examination of the two holotypes in this light.
Mr. Peter Dance, of the Mollusca Section, Bri-
tish Museum (N.H.), now reports (letter dated
October 26, 1961) that the hinges reveal all the
differences I had enumerated, the hinge of ''T."'
irus being relatively wider and heavier, with
stronger teeth than in ''T.'' inquinata. The right
valve of "T." irus has a pseudo-lunule that
makes the margin sinuate, whereas it is smooth
in the latter. Thus, to bring our terminology
into line with that already adopted by Japanese
workers, we must revive the name Macoma in-
quinata (Deshayes) as the correct one for the
West Coast shell that we have been citing in re-
cent years as M. irus. This species is a true
Macoma; its range is from Bering Strait to the
Los Angeles Area. The range of Heteromaco-
ma irus (Hanley) outside Japan is a matter for
further study: this form may or may not be
identical with the true Fragilia yantaiensis from
the Chinese coast.
On the Systematic Place of
Cypraea mus
by
A. Myra KEEN
Stanford University, California
In a discussion of sizes in cypraeid species
recently, Dr. F. Schilder (The Veliger, vol. 4,
no. 2, pp. 107-112, October, 1961) on p. 109 cited
this form as Siphocypraea mus (Linnaeus).
Probably he has difficulty at his laboratory in
East Germany keeping abreast of American lit-
erature; hence, “he has evidently overlooked
Woodring's discussion of the species and pro-
posal of the subgeneric name Muracypraea for
it (The Nautilus, vol. 70, no. 3, pp. 88-89, Janu-
ary, 1957). As Woodring showed, the type of
Siphocypraea is S. problematica (Heilprin, 1887),
from the Pliocene of Florida, a unique form of
slender outline, with a very deep spiral channel
at the apical end. The group of Muracypraea,
of which Cypraea mus is the type, originated
during Miocene time in the Caribbean, with a
number of fossil forms that occur at widely
separated localities, such as Venezuela, Flori-
da, and the Atlantic Coast of Panama. Cypraea
(M.) mus is the sole modern survivor of a group
characterized by a rather chunky, triangular,
heavily-callused shell, with low, irregular
ridges (not a channel) in the apical area.
Two recent publications of Dr. Schilder's
that may escape notice are cited below. Both
mention Pacific species.
Schilder, F. A.
Zur Kenntnis der Cypraeidae. No. 2. Ar-
chiv fiir Molluskenkunde, Bd. 89, no. 4/6,
pp. 185-192, pls. 14-15, 1960,
Schilder, F. A., & M. Schilder
Zur Kenntnis der Cypraeidae. No. 3. Lange
und Reihenzahl der Radula. Ibid., Bd. 90,
no. 1/3, pp. 33-42, Marchl, 1961.
Page 162
A Living Fossil
EMERY P. CHACE
Natural History Museum, San Diego, California
The only known living specimen of Nassa-
rius delosi (Woodring, 1946) was reported on in
1957 (Chace, 1957) although it had been previ-
ously recorded as N. californiana (Conrad,
1856). A second living specimen of this rare
species was found by Mrs. Charlene Neeb at a
low tide in July, 1961, on a sandflat on the shore
of Mission Bay, San Diego. As Mrs. Neeb did
not recognize the animal, she brought it to the
Natural History Museum of San Diego for iden-
tification. It proved to be N. delosi, thus be-
coming the second known Recent specimen. The
sculpture of the specimen in questionis very
similar to that shown in Bulletin 112 (Dall, 1921)
as N. californiana and for which Woodring pro-
posed the name N. delosi. This second speci-
men is 28 mm. in height and has a diameter of
16 mm.; it is deposited in the Natural History
Museum of San Diego, where it bears the lot
number 42'927, It may be worth mentioning
that this species, though extremely rare as a
living form, is rather common in the Pleisto-
cene exposure of the Los Angeles Area.
Literature Cuted
Chace, Emery P.
1957. Nassa delosi Woodring.
Dall, William Healey
1921. Summary of the marine shellbearing mollusks
of the northwest coast of America, from San Diego,
California, to the Polar Sea, mostly contained in
the collection of the United States National Museum,
with illustrations of hitherto unfigured species.
U. S. Nat. Mus. Bull. 112: pp. 1-217, pls. 1-22.
Woodring, W. P., M. N. Bramlette, & W. S. W. Kew
1946. Geology and paleontology of Palos Verdes Hills,
California. U.S. Geol. Survey Prof. Paper 207,
v + 145 pp., 37 pls.
Nautilus 70 (3): 108.
AME CUR ee De
Asilomar, Pacific Grove, California
June 27-30, 1962
The 1962 annual meeting of the Pacific Di-
vision, American Malacological Union, will be
held at Asilomar, Pacific Grove, California, on
June 27 through 30, inclusive. For details con-
cerning reservations, contact Mrs. Lucille
THE VELIGER
Vol. 4; No. 3
Zellers, 714 Elm Street, El Cerrito, California;
for details concerning presentation of papers,
contact Robert R. Talmadge, Box 71, Willow
Creek, California; for details on displays, con-
tact Mr. Gale Sphon, 1109 Cacique Street, Apt.
E, Santa Barbara, California.
RRT
Information Desk
What’s the Difference?
Genotype - Phenotype
by
R. STOHLER
Department of Zoology
University of California, Berkeley 4, California
The two terms in our subtitle have been
used by students of genetics to designate the
genetic constitution of an individual and the vis-
ible appearance of an individual, respectively.
Unfortunately, the word genotype has been used
by various authors in taxonomic papers to indi-
cate what we now call the type species of a ge-
nus. Both terms, it should be emphasized, are
not a part and parcel of the taxonomist's vocab-
ulary regarding type specimens of one sort or
another. Yet in recent literature of a system-
atic nature, these two terms, used either in
their noun or in their adjective form, are ap-
pearing more and more frequently. The con-
cepts for which the terms stand are, of course,
of importance to the systematist at least as
much as to the geneticist. We will endeavor
here to explain the meanings of these terms and
point up some of the ramifications of import-
ance to the student of taxonomy.
Any individual (plant or animal) inherits a
series of characteristics from its ancestors in
a very orderly fashion. It is, of course, not the
purpose of this discussion to elucidate the me-
chanisms involved, nor to enumerate the con-
sequences of the facts of inheritance. It is suf-
ficient to mention that hereditary traits include
many factors known and understood, as well as
many more as yet unknown or only partially un-
derstood. It is also known that the genotype —
the genetic equipment — of an individual is un-
alterably determined at the moment the egg is
fertilized; it is also understood that the outward
appearance (the phenotype) of an organism is
Vol. 4; No. 3
THE VELIGER
Page 163
governed by the genotype. This, at first glance,
might imply that the phenotype is also unalter-
ably determined at the time of fertilization. In
a great number of cases this holds true, but
there are many instances known where the phe-
notype may be affected in one way or another.
While such characters as hair color or eye
color in human beings or mammals are easily
understood, there are other traits that are not
as readily recognized as being inherited by
means of the same mechanisms. We have come
to realize that, to mention one example, sta-
ture in human beings may be affected by varia-
tions in nutrition during critical growth peri-
ods; thus a child of relatively short parents may
develop into a tall individual if properly fed
during the growing period,yet at the same time,
another individual may be properly fed and still
remain short. This would simply indicate that
the first individual had genotypically the ability
to become tall if properly fed and the ability to
remain short if improperly fed, while the sec-
ond individual genotypically did not have this
ability to respond in the same manner to the
environmental effect. We might express this
same situation by saying that both individuals
are phenotypically of short stature, assuming
the first individual was improperly nourished,
but genotypically they are still different from
each other, one having inherited the possibility
of responding in different ways to environmental
factors, while the other individual did not in-
herit such an ability.
From the foregoing it becomes clear, then,
that many organisms may inherit the ability of
responding to a variety of outside factors —
factors situated in the environment in which the
individual lives; we might call such individuals,
for convenience in our discussion, ''plastic
forms'', Other species not having inherited the
ability of response to the environment by pro-
ducing a different phenotype, by the same token,
may be called ''set forms",
The student of mollusk shells may become
aware of these differences when he studies and
compares the shells of one and the same spee
cies obtained from various points of a wide
range of distribution. He will find, probably,
that in some species there is no difference rec-
ognizable, no matter from where the shells were
obtained. This would be the case in a "set"
species. In other species, on the other hand, it
may be possible for the experienced collector
to look at a given shell and know the exact geo-
graphical locality from which it came (''plastic"
species). It seems unnecessary to stress that
many apparently ''set'' species of mollusks may
possess a much greater range of distribution
than is at present known, and having at the
same time genotypically the ability to respond
to. different environmental conditions by pro-
ducing different shell characteristics, thus ac-
tually being ''plastic'’. In such a case, repre-
sentatives obtained from scattered areas of the
entire range of distribution may be mistakenly
considered as different taxonomic entities.
Only experimental transplantations of groups of
individuals from one area into the other could
reveal whether they are individuals of a ''plas-
tic'' or ''set'' species. The assumption that they
are part of a'"'plastic'’ species would gain
strength, possibly, if specimens could be col-
lected from all intermediate areas of the known
localities.
From the foregoing, two observable facts
can possibly be understood. First, it is an al-
most logical conclusion that a ''set'' species
would have, generally speaking, a much more
limited range of distribution than a "'plastic"'
species may possess, simply because of the in-
ability to produce different phenotypes in dif-
ferent environments, since it may be assumed,
again generally speaking, that the different phe-
notypes of ''plastic'' species are better equipped
to cope with the conditions in the particular en-
vironment in which they developed, although the
possibility that there is no advantage (but also
no disadvantage!) accruing to such variable
forms must also be borne in mind. Secondly,
that''plastic'' species occupying large ranges of
distribution have, once again generally speak-
ing, acquired long lists of synonyms.
It is, however, also clear that the terms
phenotypic and genotypic cannot be used cor-
rectly as truly alternative terms, since both
"plastic" and "'set'' phenotypes are determined
by the particular genotypes. If a student wishes
to bring out the differences between a ''plastic"’
and a ''set'' species, he may have to coin new
terms (such as we have done here) to designate
unmistakably, unequivocally, and logically what
he really means. It is obviously inaccurate to
state that a certain character in a particular
species is phenotypic while another one is ge-
notypic, when actually the student means that
one character is a ''plastic'' character while
the other one is a "'set'' character,
Page 164
THE VELIGER
Vol. 4; No. 3
Books, Periodicals, Pamphlets
HOW TO KNOW THE AMERICAN
MARINE SHELLS
by R. Tucker Abbott
Signet Key Book, 1961. 75¢.
This fine little book should prove very help-
ful for all beginners in the field of conchology.
Much valuable information is contained in the
222 pages and the color illustrations are, in
part, magnificent. West coast species are well
represented, which up to now has been rather
unusual in books written and published in the
East. We would take exception only to two points
in the whole work, one being what Dr. Abbott is
saying about our journal, The Veliger. In a per-
sonal letter to your editor, sent before the
booklet reached the market, Dr. Abbott indi-
cated that he was aware that his statement no
longer held true, and he explained the lapsus as
due to the time necessary in the production of a
work such as this. The other point with which
we cannot wholeheartedly agree is Dr. Abbott's
consistent habit of translating the scientific
names and giving them ina manner as if they
were generally accepted vernacular names.
This practice in many instances yields a ''com-
mon'' name which is more awkward and more
difficult to remember than the real scientific
name. Aside from these minor points, we feel
that this book can be recommended without re-
servation and will fill a long-felt need —a
source to which the many beginners can be re-
ferred when they inquire of shell club secre-
taries about sheli collecting.
RS
JOURNAL DE CONCHYLIOLOGIE
Vol. 101, No. 1, for January 1, 1961.
"Malformations et colorations spécifiques
chez plusieurs Cypraea de la Nouvelle-Calédo-
nie et de la Polynésie frangaise'', L. J. Bouge.
MK
PRINCIPLES
OF ANIMAL TAXONOMY
by George Gaylord Simpson
Columbia University Biological Series,
No. XX, 1961. $6.—.
The author is Alexander Agassiz Professor
of Vertebrate Paleontology at the Museum of
Comparative Zoology, Harvard University. He
has been a member of the staff of the American
Museum of Natural History and on the faculty of
Columbia University. He is the author of The
Meaning of Evolution; Horses; The Major Fea-
tures of Evolution; Life, An Introduction to Bio-
logy; and Quantitative Zoology.
This book provides a detailed introduction
to the fundamental principles of animal taxono-
my. It is the only general study available in
English on the principles underlying the classi-
fication of animals, and the only modern treat-
ment of the subject in any language,
The author defines taxonomy as ''the theo-
retical study of classification, including its
bases, principles, procedures, and rules". He
begins by considering order in nature and dis-
cusses the scientific ordering responsible for
the modern classification of organisms. He
examines the kinds of taxonomic evidence avail-
able to the scientist and discusses the historical
development of taxonomy from Linnaeus to the
present, including the relationship of taxonomy
and evolution. Special chapters are devoted to
the lower and the higher taxonomic categories.
JQB
THORACIC CIRRIPEDIA
OF THE GULF OF CALIFORNIA
by Dora Priaulx Henry
Univ. Washington Publ. Oceanography,
vol. 4, no. 4, pp. 135-215, pls. I-V. De-
cember 27, 1960. University of Wash-
ington Press. Seattle. $1.—.
This paper is based ona collection of bar-
nacles collected by Walter J. Eyerdam in the
vicinity of Guaymas, Sonora, Mexico, in Janu-
Vol. 4; No. 3
THE VELIGER
Page 165
ary, 1959. A total of 14 species and subspecies
are covered, including Balanus eyerdami, new
species. Dr. Henry includes a useful table (pp.
146-147) of the 21 species of thoracic cirripedia
reported from the Gulf of California with their
localities, habitats, and distributional ranges.
The plates contain excellent photographs of Ba-
lanus eyerdami, new species, and its charac-
teristic component parts. Dr. Henry's paper is
an important contribution to knowledge of the
invertebrate fauna of the Gulf and is a neces-
sary reference for students of the barnacle
group.
AGS
2
SHELLS
OF THE NEW YORK CITY AREA
by Morris K. Jacobson
and William K. Emerson
Argonaut Books, Inc., 1961. $4.—.
A handbook of the land, fresh water, and
marine mollusks ranging from Cape Cod to Cape
May, with 150 drawings by Anthony d'Attilio.
Over 140 species of shells are described, and
each is illustrated by an exquisitely executed
line drawing.
Several unique features enhance the useful-
ness of this handbook.
1, A map of the area, keyed to the principal
collecting localities.
2. A listing of all the shells to be found in each
major collecting area, cross-referenced to the
pages on which the shells are described.
3. A listing of all the shells according to their
scientific arrangement by class and subclass,
order and suborder, superfamily, family, and
subfamily, also cross-referenced to the text of
the book,
Another feature that will be welcomed by
collectors is the fact that the illustration for
every shell is located within one page of its de-
scription. The combination of an informal and
entertaining style with authoritative technical
data makes this book a ''must'' for both the be-
ginning nature student and the advanced collec-
tor or malacologist.
JQB
CARIBBEAN SEASHELLS
A Guide to the Marine Mollusks
of Puerto Rico and Other
West Indian Islands, Bermuda,
and the Lower Florida Keys
by Germaine L. Warmke
and R. Tucker Abbott
Livingston Publishing Company. 196l.
$8.95.
This book is a unique shelling guide and au-
thoritative identification book designed for both
the amateur conchologist traveling in the West
Indies and for students of Caribbean marine
biology. Over 800 species are accurately clas-
sified and described. Forty-four plates of
photographs (some in full color), habitat notes,
and geographical ranges simplify finding and
naming the shells of this tropical area. Although
primarily designed for students and research
workers at Puerto Rico's Institute of Marine
Biology, the book includes most of the seashells
of the Lower Florida Keys, Bermuda, the Ba-
hamas, and the Lesser Antilles.
A series of distributional maps at the end
of the book gives the reader a useful picture of
what species are most likely to be found in any
one area of the tropical Western Atlantic region.
JQB
EAST AFRICAN SLUGS
OF THE FAMILY UROCYCLIDAE, |
THE GENUS TRICHOTOXON
by B. Verdcourt and R. Polhill
Journal of the East Africa Natural His-
tory Society, Special Supplement No. 7,
36 pp., 42 textfig. April 1961. $1.50.
This is a scholarly paper, illustrating and
discussing the anatomy of the species, their
habits, and includes a well-prepared key to the
species.
JQB
Page 166
THE VELIGER
Vol. 4; No. 3
VENUS:
THE JAPANESE JOURNAL
OF MALACOLOGY
Vol. 21, No. 3, August, 1961.
Molluscan shells from Southern Kii. T.
Kuroda and K6édé It6.
Latisipho: Notes on two species of the ge-
nus Latisipho. T. Habe.
Three new gastropods from Japan. T. Habe.
On Japanese Terebrid fossils and a few
new facts of the family. K. Oyama.
Review of nomenclature of Japanese shells
(4). K. Oyama.
On three new species of Notoacmea (sic).
T. Kira.
Descriptions of six new species of Japanese
marine gastropoda. M. Azuma.
Solariella nektonica, sp. nov. Description
with special reference to its swimming
T. Okutani.
behaviour.
On the family Triphoridae (Gastropoda)
from Amami Islands. S. Kosuge.
MK
MOLLUSCS, AN INTRODUCTION
TO THEIR FORM AND FUNCTIONS
by J. E. Morton
This scholarly book was previously re-
viewed in The Veliger, Vol. 2, No. 3, p. 68. Itis
now available paperbound from Harper & Bros.,
Inc. $1.40.
JQB
THE VELIGER is open to original papers pertaining to any problem
concerned with mollusks from the Pacific Region.
This is meant to make facilities available for publication of articles
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of marine, fresh water or terrestial mollusks from any region bordering
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Short original papers, not exceeding 500 words, will be published in
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EDITORIAL BOARD
Dr. Donatp P. Assort, Assoctate Professor of Biology
Hopkins Marine Station of Stanford University
Dr. J. Wyatr Duruam, Professor of Paleontology
University of California, Berkeley
Dr. E. W. FAGER, Associate Professor of Biology
Scripps Institution of Oceanography, University of
California, La Jolla
Dr. Cavet Hann, Associate Professor of Koolog y
University of California, Berkeley
Dr. G. Datias Hanna, Curator, Department of Geolog y
California Academy of Sciences, San Francisco
Dr. Joe, W. Hepepetn, Professor of Zoology
Director of the Pacific Marine Station, Dillon Beach
Dr. Leo G. HERTLEIN, Associate Curator, Department of Geology
California Academy of Sciences, San Francisco
Dr. Myra KEEN, Associate Professor of Paleontology and
Curator of Concholog y
Stanford University
Dr. Frank PITELKA, Professor of Coolog y
University of California, Berkeley
Mr. ALLyn G. Situ, Associate Curator, Department
of Invertebrate Zoology
California Academy of Sciences, San Francisco
Dr. Racpeu I. Suir, Professor of Koology
University of California, Berkeley
EDITOR
Dr. RUDOLF STOHLER, Associate Research Coologist
University of California, Berkeley
A Quarterly published by
NORTHERN CALIFORNIA MALACOZOOLOGICAL CLUB
Berkeley, California
VOLUME 4 April 1, 1962 NUMBER 4
CONTENTS
*
Revision of the Clavagellacea (9 Textfigures )
ELEM ANDERSON GOMUBE mires ee TL ON vol be nic rst Saha ie aes vey) Rede ee LO
A New Dampierian Cypraca (Plate 40; 1 Textfigure)
CRIA VE ORD MINER CARE Mrmr ler camOns . lad ay ey LES nee we wee ie a le ES
Nomenclatural Notes on some West American Mollusks, with Proposal of a
New Species Name
Nem LS RVAGE BEN nen as Sth ROC TIES nae WEY el nce og ck cng ees eB
A New Method of Determining the Accuracy of Geotactic Orientation of
the Snail Helix aspersa MULLER (3 Textfigures)
DONALD EROBBOWER Bn rehire can Lar pte a) te eter airmen Motos keene ee OOM
Observations on Three Species of Vexillum (Gastropoda )
(Plates 41 to 44; 2 Textfigures)
BREAN [eg AGRE Gain Ric eco eect an) Geo utaei Sl epticaeksg yim set sng <2! aap oimie: se POA
Manometric Measurements of Respiratory Activity in Tegula funebralis
JASMISS JE [e IMICIEIE ANS oc ek See ee ic dle? ad. oh el allaieatua ie lb cee ee real gareteil emia re acre (0)
A Study of Food Choices of Two Opisthobranchs, Rostanga pulchra MCFARLAND
and Archidoris montereyensis (COOPER) (4 Textfigures)
Emity F. Coox RO eee Wer SR eR Me se suse Gen ules ten as yi oe ae AOA.
Relationship of Living Weight to Shell Cavity Volume in Helix aspersa
(Plate 45; 2 Textfigures)
ANDREAGHERZBERGICCHERED IERZBERGI Gri) ee) Geile el et 6 TO
[Continued on Inside Front Cover |
Subscriptions payable in advance to N. C. M. C.
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Send subscriptions to: Mrs. PHorsE BaLcu, Treasurer, 1150 Brown Avenue, Lafayette,
California. Address all other correspondence to: Dr. R. STOHLER, Editor, Department of
Zoology, University of California, Berkeley 4, California.
Second-Class Postage paid at Berkeley, California
CONTENTS
[ CONTINUED ]
Type Localities
F. A. ScHILDER SHAT a OMe veh Mato Res
e . e ° ° e
A Preliminary Report on Spawning and Related Phenomena in
California Chitons (3 Textfigures)
SPENCER R. THORPE, JR.
NOTES & NEWS
Busycotypus (B.) canaliculatus in San Francisco Bay. Rupoir STOHLER
Recent Uses of Non-binomial Works. R. Tucker ABBOTT
New Name for Strombus granulatus subsp. acutus DURHAM, 1950, not PERRY, 1811.
J. Wyatr DurHaM
The W. Mack Chiton Collection. ALLyn G. SmitH
California Brown Cowrie in Central California. ALtyn G. SmirH
Arion ater (LINNAEUS) in California. ALLYn G. SmitH
METHODS & TECHNIQUES
Notes on Cleaning Mollusks. Attyn G. Smitu
INFORMATION DESK a ea er ee Pt nea te Ms
What’s the Difference? Wolotype - Paratype - Syntype - Hypotype. R. STOHLER
BOOKS, PERIODICALS & PAMPHLETS
- 199
- 202
- 2ir
. 216
F any
. 218
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).
Vol. 4; No. 4
THE VELIGER
Page 167
Revision of the Clavagellacea
BY
LEE ANDERSON SMITH
Department of Geology, Stanford University, California
(g Textfigures)
The restudy of this unusual and interesting
pelecypod group was undertaken as a revision
for the ''Treatise on Invertebrate Paleontology’,
in which a résumé of the taxa and types of the
superfamily will appear. It seems advisable,
however, to publish a more complete account at
an earlier date. Two type designations are con-
tained herein which, it is hoped, will help sta-
bilize the nomenclature of the group.
I have adopted the style used in the
"Treatise on Invertebrate Paleontology" for
nomenclatural notation. The style is brief and
therefore conserves space but may not be self-
explanatory to all readers. Readers having ac-
cess to any of the ''Treatise'' volumes may refer
to the discussion of nomenclatural notation
given in the front of each.
During the investigation, it was necessary
to place as many species as possible in their
respective genera and subgenera. They are
listed here as briefly as possible after the de-
scription of each generic or subgeneric taxon.
If they could not be placed in a subgenus or do
not actually belong in the family, they are given
immediately after the description of the genus,
(s.1.). Names cited and questioned are from
references not available to me, and their sys-
tematic placement is based on information giv-
en by earlier authors.
The study was made possible through ac-
cess to the library and museum facilities of
Stanford University, the California Academy of
Sciences, and the University of California at
Berkeley. Grateful acknowledgment is extended
to Dr. Myra Keen for continuous advice, assist-
ance, and encouragement during the investiga-
tion and for critically reading the final manu-
script. However, the writer accepts full
responsibility for any errors of transcription
or of judgment.
CLAVAGELLACEA Orsicny, 1844
nom. trans., THIELE, 1934, ex CLAVAGELLIDAE
Shell nacreous, free when young, degener-
ate in adult; with one or both valves at least
partially imbedded in an_elongate calcareous
tube, an adaptation for burrowing; hinge-plate
wanting; ligament external. U. Cret. (Turoni-
an)-Rec.
Characters of the hinge, material of the
shell, and gill structures of the animal suggest
a systematic position for the Clavagellacea be-
tween the Pandoracea and the Poromyacea.,
Eames (1957) proposed Kitsoniidae as a
family that he provisionally placed in the Clava-
gellacea. The Eocene form from Nigeria, upon
which Eames! ''family'’ is based, is definitely
not a clavagellid. Characters of the hinge, lack
of nacreous shell material, and general config-
uration of the shell strongly suggest a position
in the Arcticacea.
CLAVAGELLIDAE ORBIGNY, 1844
[ = AsPERGILLIDAE GRAY, 1858 ]
Tube anteriorly rounded or discoid and
smooth or fringed with simple or branching tu-
bules, or partially closed by a calcareous disc
having several pedal foramina. U. Cret. (Tu-
ronian)-Rec.
Ecology and Anatomy
An excellent account of the anatomical fea-
tures of Penicillus, s. s., (=Aspergillum) was
given by Lacaze-Duthiers (1870, 1883) from the
study of preserved specimens. The lack of
firsthand observations of living specimens has
Page 168
THE VELIGER
Vol. 4; No. 4
misled earlier authors concerning the living
position and habits of the animal. As late as
1958, Morton stated that Penicillus (= Brechites)
lies horizontally on the sea floor, Earlier re-
ports even suggested burial in a vertical posi-
tion with the perforated disc exposed above the
substrate, an impossible suggestion consider-
ing pelecypod functional morphology. The re-
verse and true position was described by H. &
A. Adams (1858).
Living specimens of Penicillus penis (Lin-
naeus) have been observed in Java, and Purchon
(1956, 1959) gives an excellent account of the
musculature, movements, and ciliary feeding
and cleansing mechanisms in the mantle cavity.
Purchon (1959) stated that the specimens were
found ''a little above the low-water mark of
spring tides, in an area of muddy sand sparsely
colonised by the marine flowering plant, Enhalus
sp.'’ The second live specimen found "was lying
vertically in the substratum with the posterior
circular orifice of the shell projecting slightly
above the surface of the soil; .. . the siphonal
process was expanded and the inhalant and ex-
halant apertures'' were visible close to the up-
per end of the tube. ‘''When disturbed, a small
jet of water was emitted in the usual fashion and
the siphonal process was withdrawn from view.'!
In the laboratory he observed muscular
contractions that occurred irregularly and
sometimes resulted in posteriorward, some-
times anteriorward, movements of the anterior
pallial septum. Purchon (1959) suggested that
"these movements cause water to be drawn into
the shell from the substratum through the an-
terior perforated disc of the shell’ [tube]. He
believed it to be a mechanism for becoming
more deeply imbedded in the substrate and ''may
also provide a subsidiary source of food mate-
Tealeyl ly
As with other burrowing forms, the siphons
are highly developed and generally the largest
part of the animal. The calcareous tube is se-
creted around the siphons and fused with one or
both of the valves near the anterior end. Clava-
gella, s.1., may reach three feet in length and
two inches in diameter, but most species of
clavagellids are much smaller, averaging
probably not more than three to four inches in
length. Some species are only a few millime-
ters in length.
Several present-day species are found liv-
ing intertidally or in shallow subtidal areas, and
the sediments and faunas associated with fossil
How -
small and rare survivors belonging to
forms suggest shallow marine habitats.
ever,
Clavagella, s.s., are reported from depths ex-
ceeding 100 fathoms in Australasian seas.
Habit, Morphology and Taxonomy
Members of the family live either buried in
sand or sandy—mud, or attached to or within
openings in a solid substratum. The three
modes of life do not directly account for the
three-fold subdivisions of the family, but they
certainly control subgeneric groupings.
Forms living in a soft substrate, e. g.,
Penicillus, s.l., and Stirpulina, are able to
The
forms restricted to openings in a solid sub-
strate, e.g., Bryopa and Dacosta, are forced to
conform to their rocky habitat. A three-fold
division, purely based on mode of life, would
place Clavagella, s. s., and Stirpulina with the
penicillids, an arrangement with which charac-
ters of the shell and soft parts would not agree.
achieve and maintain greater symmetry.
The separation of Humphreyia as a third
generic taxon may be questioned, but Humph-
reyia appears to be intermediate between Cla-
vagella and Penicillus in both morphology and
habit. In Clavagella, s.l., only one valve
merges with the tube, and most of the valve is
visible externally even in the adult. In Peni-
cillus, s.l., both valves merge early with the
calcareous tube and only the umbonal portion of
the valve is visible externally in the adult,
In Humphreyia, according to Gray (1858),
the young becomes attached to a rock by one
valve, and both valves merge into a single
curved plate, which forms most of the bag-like
cavity. A large part of the opposite valve is
visible in the adult. If Gray's observations are
factual, Humphreyia has both valves merging
with the tube as in Penicillus, s.1., but has one
adult valve exposed externally as in Clavagella,
s.l. I have noticed in the Stanford collection
juvenile specimens of clavagellids attached by
one valve to one side of an opening in rock,
which they presumably would have filled had
they grown to adulthood. Therefore, although
both valves merge with the tube, the mode of
attachment and life habits of Humphreyia sug-
gest a closer alliance with Clavagella, s.1., than
with Penicillus, s.1l., with which it has most of-
ten been placed.
The geologic history of the group suggests
also that Humphreyia may be a Recent branch
of the clavagellid line. Clavagella, s.1., is the
oldest group and Humphreyia the youngest; but
Penicillus, s.1., was clearly separated by Late
]
Vol. 4; No. 4
THE VELIGER
Page 169
Oligocene, and it seems unlikely that a ''primi-
tive penicillid'' should have survived since that
separation undetected in the geologic record.
It seems even less likely that a modern and true
penicillid should revert to the habits and gro-
tesque form of Humphreyia.
Geographic and Geologic Distribution
Clavagella, s. s., is apparently the most
primitive group, with questionable records from
strata older than the Turonian of Cretaceous in
Europe and Africa. The suggestion is consist-
ent with the shell and tube morphology, as Cla-
vagella, s.s., has the appearance least altered
from that of a normal burrowing pelecypod. The
siphonal portion of the tube is*smaller in diam-
eter than the shell, and the adult valves still
dominate the anterior portion. .
Two distinct groups of clavagellids are re-
ported from strata of Late Cretaceous (Turoni-
an) age in Europe, North America, and in India.
The existence of two distinct lines at that time
suggests earlier beginnings for the group, per-
haps during Early Cretaceous. The Late Cre-
taceous Clavagella, s.s., and Stirpulina are
both found in Europe and before the end of Cre-
taceous Clavagella, s.s., appeared in Asia and
Stirpulina in North America. The exact syn-
chroneity of these occurrences is not known.
The two groups may have developed independ-
ently in the Pacific and Atlantic areas, but it
seems more likely that they branched from a
Single stock in the European area.
Stirpulina, which did not live beyond Cre-
taceous in North America, survived with Clava-
gella, s.S., until Late Pliocene in Europe,
Clavagella, s.s., may have a small surviving
representative in the moderately deep waters
_off Australia {C. (C.) multangularis Tate}, and
Stirpulina is represented in the Recent faunas
by C.(S.) ramosa Dunker in Japanese waters,
ee
The earliest records of Bryopa are from
strata of Late Oligocene (Aquitanian) age in
France. There are several survivors in the
Mediterranean and a few in Indo-Pacific and
Australasian provinces.
Dacosta and Humphreyia are known only
from living forms in Australia, and neither
group is well established.
Penicillus, s.s., is recorded from Late
Oligocene strata of N. W. Borneo, Miocene of N.
Borneo, Pliocene over the Indo-Pacific area,
and Pleistocene of the Indo-Malay and Philip-
pine regions. Though rare, it is reported from
a large part of the Western Pacific and Indo-
Pacific areas today.
Pseudobrechites is the only penicillid
group reported from European strata, and only
a Single species has been recorded. The speci-
mens were collected from Late Oligocene strata
of France, and no other penicillids have been
reported from nearer to Europe than the pres-
ent Red Sea.
Foegia is an entirely Pacific-Indo-Pacific
form. Earliest records are from Miocene and
Pliocene strata of Sumatra, Formosa, and Ja-
pan. It is presently living in the Indo-Pacific
and Australasian areas,
Warnea is reportedly living in the Red Sea
and in Australasian and Japanese waters. The
earliest records are from Pliocene strata of
Japan.
The Clavagellacea is at present primarily
an Indo-Pacific-Australasian group. Original
development in the European area is suggested,
and the group has apparently since declined and
moved eastward. All of the penicillids have
geologic records from at least Pliocene, and
only Dacosta and Humphreyia are Recent devel-
opments, probably of the clavagellid line.
Generic, Subgeneric, and Specific Taxa
Clavagella Lamarck, 1818
[* C. echinata; SD CuiLpren, 1823]
[=Bacilia Gray, 1858, ex Valenciennes MS
(obj.) Clavigella, spelling error] One valve
never merging with the tube, and both adduct-
ors persistent in the adult. U. Cret. (Turoni-
an)-Rec.
C. (?) cenomaniana Orbigny, 1844, p. 157
[nom.nud.] Cret., Fr.
C. (?) dubia Muenster, 1835, p.435 [nom,
nad. ]
C. (?) ligeriensis Orbigny, 1844, p. 233 [nom.,
FKL |) (Casting 1dse
C. (?) altavillae Aradas & Calcara, 1843, p.
221; (not seen)
C. (?) dalpiazi Venzo, 1941; (not seen) Olig.,
Italy
. (?) lodoiska Caillat, 1835, p. 237 (not seen)
. (?) lybica Parona, 1923, p.51; (not seen)
Us Cxret:, Afr.
. (?) prisca Goldfuss, 1840, p. 285;(not seen)
. (?) zebuensis Broderip; (not seen) Rec.
Philippines
la 1a
IO.10
Page 170
Clavagella (Clavagella)
Siphonal end of tube simple; tube free, elon-
gate, clavate, compressed and symmetrical
in shape; with irregular spine-like tubules on
the anterior portion of the tube. U. Cret.
(Turonian)-Rec., Eu.-India-Australas. —Fig.
1. C. (C.) echinata, M.-U. Eoc. (Lut.-Bart.),
Paris Basin.
C. (C.) brocchii Lamarck, 1818, p.432, U.
Plioc., Italy
C. (C.) brongniarti Deshayes, 1824, p.11, U.
IDOVEnn 1D aR.
C. (C.) cretacea Orbigny, 1845, p. 300, Cret.,
Fr. .
C. (C.) cristata Lamarck, 1818, p.432, M.
IWOCo, I iP,
C. (C.) echinata Lamarck, 1818, p. 432, M.-
Wie IZOCs, IPI,
€: (2? C:) exigua Zittel, 1865, py lO”; Ue Gret.,
Austria
C. (C.) lagenula Deshayes (not seen) U. Eoc.,
1D ep
C. (?C.) lamarcki Deshayes (not seen) M.
ITXC, IPG,
C. (C.) multangularis Tate, 1886, Rec., Aus-
tral.
C. (?C.) primigenia Deshayes (not seen) M.
Paleoc., Fr.
C. (C.) semisulcata Forbes, 1846, p. 139, U.
Cret., India
C. (?C.) tibialis (Lamarck, 1818), p. 432 [Fis-
tulana] Eoc., Fr.
higure 1
Clavagella (Clavagella)
Clavagella (Surpulina) SroviczKa, 1870
|* C. coronata DesHayes, 1824]
[=Styrpulina, spelling error. May prove to be
synonym of Tubolana BIVONA-BERNARDI,
1832 (type-species: T. digitata=Aspergillum
bacillaris DESHAYES, 1830; Tubulana, spell-
THE VELIGER
Vol. 4; No. 4
ing error)} Siphonal end of tube periodically
expanded, anterior end with tubules formed
only in terminal corona; tube long with a more
or less distinct anterior slit. U. Cret. (Tu-
ronian)-Rec., N. Am.-Eu.-Afr.-Asia —Fig. 2,
C. (S.) coronata, U. Eoc. (Bart.), Paris Ba-
sin.
7
HG (S.) armata Morton, 1833, p. 129, U. Cret.,
N. Jersey
C. (S.) aspergillum Bronn, 1828, p.5 [=C.
(S.) bacillaum (Brocchi, 1814)]
C. (S.) bacillum (Brocchi, 1814), p.273 [Te-
redo], Mioc., Austria
C. (S.) bacillum bacillaris (Deshayes, 1830),
Bo 2I95 IPMIOCs, Pie,
. (?S.) clavata (Roemer) Orbigny, 1850, p.
233, ?Cret., Germany
. (S.) caillati Deshayes (? MS) M. Eoc., Fr,
. (?S.) cornigera Schafhaeutl, 1863, p. 179,
U. Cret., Bavaria [both valves may be
free = ? pre-Clavagella]
. (S.) coronata Deshayes, 1824, p. 8, U. Eoc.,
Fr,
C. (S.) digitata (Bivona-Bernardi, 1832), p.
56, ?Plioc., Italy [=C.(S.) bacillum bacil-
laris (Deshayes, 1830) ]
C. (?S.) elegans Miller (not. seen) U. Cret.,
Germany
C. (S.) goldfussi Philippi, 1846, p. 44, L. Olig.,
Germany
C. (S.) maniculata (Philippi, 1836), p. 1,
? Plioc., ?Italy [? = C. (S.) bacillum bacil-
laris (Deshayes, 1830)]
G. (S.) oblita Michelotti, 186l, p. 53) (U.Olip:
(Tongrian), Hungary-Egypt
C. (S.) ramosa Dunker, 1882, p. 172, Rec.,
Japan
[Gil@ea@
IQ
Figure 2
Clavagella (Stirpulina)
Vol. 4; No. 4
Clavagella (Bryopa) Gray, 1847
[*C. aperta Sowersy, 1823]
[= Tiria GREGORIO, 1886 (C. aperta Sowerby;
herein desig., Smith, 1962, obj.)] Siphonal end
of tube periodically expanded, anterior end
smooth; with small short tubules through the
tube around the valves. U.Olig. (Aquitanian) -
Rees, Medit.-Indo-Pac. > Pig. 3, @: (B;)iaper-
ta, Rec., Malta.
C. (?B.) angulata Philippi, 1844, p. 2, Rec.,
Sicily [?=C.(B.) melitensis Broderip, 1835]
C. (B.) aperta Sowerby, 1823, XIII, Figs. 1-4,
Rec., Medit.
C. (?B.) astraeicola Jouss. (not seen)
C. (B.) balanorum Scacchi, 1844, p.4, Rec.,
Medit. [?=C. (B.) aperta *Sowerby, 1823]
Cc. (B.) brochoni Benoist, 1877, p.313, U.
Olig., Fr.
C. (B.) laqueata Sowerby (in Reeve), 1873,
Rec., ? Medit.
C. (B.) lata Deshayes, 1839, p.25, Rec., Ind.
O.-Austral.
C. (B.) melitensis Broderip, 1835, p. 116,
Rec., Medit.
C. (B.) philippiana Sowerby (in Reeve), 1873
[ex Desh. MS] Rec., Medit.
. (?B.) senilis Jouss. (not seen)
(B.) sicula Chiaje, 1830, pl.83, Rec., Me-
dit. [?=C. (B.) aperta Sowerby, 1823]
C. (?B.) socialis Jouss. (not seen)
10 |)
WOES
eS
(x 0.8)
Figure 3
Clavagella (Bryopa)
THE VELIGER
Page 171
Clavagella (Dacosta) Gray, 1858 (? recte Dacostaia)
[*C. australis SowERBY, 1829, ex SrurcHBuRY MS]
Siphonal end of tube not expanded, anterior
end smooth, rounded; with small, very short
tubules through the tube around the valves.
Rec. Wie Pac. =o big. 4. ©. (D-) auisitra lis,
Rec., Austral.
Cc. (D.) australis Sowerby, 1829, app. p. 3,
Rec., Austral,
Figure 4
Clavagella (Dacosta)
Humphreyia Gray, 1858
[* Aspergillum stranger A. ADams, 1854]
[=Humphreysia, spelling error] Tube twisted
and irregularly square in cross-section, both
valves united into a single plate forming most
of anterior bag-like cavity. Rec., W. Pac.
~mig. 5. H. strangei (A. ADAMS), Rec,
Austral.
H. coxi Brazier, 1872, p. 23, Rec. Austral,
H. strangei (A. Adams, 1852), p. 91, Rec.,
Austral.
Page 172
(x 1.2)
Figure 5
Humphreyia
Penicillus BruGuiiReE, 1789
[* P. javanus = * Serpula penis LINNAEUS 1758;
SD Hapse, 1952|
[=Brechites GUETTARD, 1770 (non-binom.);
Penecilli DACOSTA, 1776 (vernacular); Pene-
cillus, Penicellus, spelling errors; Verpa
ROEDING, 1798 (obj.); Aquaria PERRY, 1811
(A. radiata, herein desig., Smith, 1962, obj.);
Bunodus BLAINVILLE, 1817 (nom. nud.) ex
GUETTARD (non-binom.); Arytene OKEN,
1815 (obj.; rejected IC ZN, 1956); Arytaena
OKEN, 1817 (obj.; Arythaena, Arytene, spell-
ing errors); Aspergillum LAMARCK, 1818
(obj.; Adspergillum, Aspergillium, Aspergil-
lus, spelling errors); Clepsydra SCHUMACH-
ER, 1817 (obj.; Clepydra, spelling error)]
Both valves merging with tube; tube circular
in cross-section; anterior adductor degener-
ate, posterior adductor absent in adult. U.
Olig. (Aquitanian)-Rec.
"Aspergillum cretaceum'’ Rominger, 1847, p.
659 [nom.nud.] [?=Clavagella (Stirpulina)
Aspergillum javanum of Authors=P. (P.) penis
(Linnaeus)
Aspergillum sparsus Sowerby =P. (P.) penis
(Linnaeus)
THE VELIGER
Vol. 4; No. 4
Penicillus annulatus Lamarck, 1816 [?= Poly-
chaete Ann.]
Penicillus capitatus Lamarck, 1816 [?= Poly-
chaete Ann.]
Penicillus kobeltianus (Lébbeck, 1879), p. 95
[nom. nud.]
P. (?) listeri (Gray, 1825), p. 135 (not seen)
Penicillus phoenix Lamarck, 1816 [?= Poly-
chaete Ann.]
Penicillus (Penicillus)
Siphonal end of tube simple; anterior end
fringed with single row of distinct simple tu-
bules; anterior disc with a slit. U. Olig.
(Aquitanian) -Rec., Indo-Pac.-Austral.— Fig. 6,
P. (P.) penis (LINNAEUS), Rec., Singapore
P. (P.) annulatus (Thiele, 1934), p.943, ex
Deshayes ?MS
P. (P.) annulus (Gray, 1858),
Deshayes MS, Pleist.,
pines
P. (P.) clavatus (Chenu, 1843), p.4, Hab.-?
P. (P.) coronatus (Sieverts, 1934), p. 267
Plioc., Indo-Pac.
P. (P.) dichotomus (Chenu, 1843), p. 3, Plioc.,
Java
P. (P.) disjunctus Sowerby in Reeve, 1860, sp.
12, ex Deshayes MS, U. Plioc., Indo-Malay
P. (P.) incrassatus (Chenu, 1843), p. 4, ?Rec.,
Austral,
P. (P.) ornatus (Chenu, 1843), p.4 [=P. (P.)
tuberculatus (Chenu, 1843)]
P. (P.) penis (Linnaeus, 1758), p. 788, Rec.,
Ind. O.
P. (P.) pulcher Sowerby in Reeve, 1860, ex
Deshayes MS, Rec., Singapore
P. (P.) pulcher fossilis (Sieverts, 1934), p.
269, Mioc., N. Borneo
P. (P.) radiatus (Perry, 1811), pl. 52,
[=P. (P.) penis (Linnaeus, 1758)]
P. (P.) radix Gray (not seen) ex Deshayes
MS, U. Plioc., Indo-Malay
P, (P.) recluzianus (Chenu, 1843), p.4 [=P.
(P.) incrassatus (Chenu, 1843)}
P. (P.) semifimbriatus (Chenu, 1843), p. 4,
Rec., ?Red Sea
P. (P.) strangulatus (Chenu, 1843), p. 3, Rec.,
_ Austral.
P. (P.) tuberculatus (Chenu, 1843), p. 3, Rec.,
Moluccas
P. (?P.) venustulus (Beets, 1942), p. 230,
Mioc., N. W. Borneo
Pa Slie mex
Indo- Malay-Philip-
Hab.-?
Vol. 4; No. 4 THE VELIGER Page 173
Penicillus (Foegia) Gray, 1847
[“Aspergillum novaezelandiae” (= * P. novaezelandiae
BruGuIERE, 1789)|
Umbo almost covered with swollen promi-
nence; fringe indistinct, formed like slit in
disc, of short thick tubules. U. Olig. (Aqui-
tanian)-Rec., Indo-Pac.-W. Pac. —Fig.8. P.
(F.) novaezelandiae BRUGUIERE, Rec., Aus-
tralas,
P. (F.) agglutinans Lamarck, 1818), p. 430,
Rec., Indo-Pac.
P. (F.) cumingianus (Chenu, 1843), p. 3, Rec.,
Austral,
P. (F.) giganteus (Sowerby, 1888), p.290,
Mioc.-Plioc., Sumatra, Formosa, Japan;
Rec., Japan
P. (2F.) imbricatus (Perry, 1811), pl. 52,
Hab.-?
P, (E.) novaehollandiae (Chenu, 1843), p. 4,
Rec., New Holland [=P. (F.) agglutinans
(Lamarck, 1818)]
P. (F.) novaezelandiae Bruguiére, 1789, p.
Figure 6 129, Rec., Austral.
Penicillus (Penicillus ) P. (F.) novaezelandiae incertus (Chenu, 1843),
p. 4, Rec., Austral.
P. (F.) philippiensis (Chenu, 1843), p. 3, Rec.,
Penicillus .(Pseudobrechites) MAGNE, 194! PHI pone
[* Aspergillum leognanum HoENINGHAUS, 1827] P, (F.) zebuensis (Chenu, 1843), p. 3, Rec.,
: Philippines [=P. (F.) agglutinans (Lamarck,
Similar to P. (Penicillus), but fringe tubules 1818)] 0 Neb oY isle
shorter, less distinct, and anterior disc with-
out central slit. U. Olig. (Aquitanian), Eu.
—Fipg.7. P. (P.) leognanus (HEONINGHAUS,
1827), p.4, U. Olig., Fr.
Figure 7 Figure 8
Penicillus (Pseudobrechites ) Penicillus (Foegia)
Page 174
THE VELIGER
Vol. 4; No. 4
Penicillus (Warnea) Gray, 1858
[* Aspergillum australe CHENU, 1843; SD Sro.icz-
KA, 1871|
Tube cylindrical, siphonal end with series of
plaited ruffles; fringe distinct, of a single
series of thick simple tubules. Plioc.-Rec.,
Red Sea-Australas.-Japan — Fig. 9, P. (W.)
australis (Chenu), Rec., Australas.
P. (W.) australis (Chenu, 1843), p.3, Rec.,
Australas.
P. (W.) delessertianus (Chenu, 1843), p. 3,
Rec., Red Sea
P. (W.) vaginiferus (Lamarck, 1818), p. 430,
Rec., Red Sea
P. (W.) yokoyamai (Shikama, 1955), Plioc.-
Rec., Japan
(x 0.8)
Figure g
Penicillus ( Warnea)
Literature Cited
Adams, H., & A. Adams
1858. The genera of recent mollusca.
Van Voorst,
Bruguitre, M,
1789. Encyclop. Méthod, (Vers), vol. 1 (xv): 126-130,
genus 33,
Eames, F. E.
1957. Eocene mollusks from Nigeria (a revision).
Bull. Brit. Mus. (Nat. Hist.), 3 (2): 68-69,
Gray, J. E.
1847,
types.
Vol, 2: 649-650,
London,
Genera of recent mollusca, their synonyma and
Proc, Zool, London 15: 129-219.
Soc.
1858.
and Humphreyidae,
Bis
On the families Aspergillidae, Gastrochaenidae,
Proc, Zool. Soc. London 26:
Lacaze-Duthiers, H. de
1870. Sur l'organisation de l'arrosoir, Aspergillum
javanicum. C, R. Acad. Sci, Paris 70; 268-271,
1883. Morphologie des acéphales, premier mémoire,
Anatomie de l'arrosoir. Arch, Zool, Exp, Gen.
2 (1): 1-68.
Lamarck, J.-B. P.
1818. Animaux sans vert@bres, VI: 428-432.
Morton, John E,
1958. Molluscs. Hutchinson, London. p, 195.
Purchon, R. D.
1956. A note on the biology of Brechites penis (L. )
(Lamellibranchia). J. Linn. Soc. 43: 43-54,
1960.
(L. ) (Lamellibranchia),
don, 34 (1): 19-23.
A further note on the biology of Brechites penis
Proc. Malac. Soc, Lon -
Vol. 4; No. 4
THE VELIGER
Page 175
A New Dampierian Cypraea
BY
CRAWFORD N. CATE
Conchological Club of Southern California, Los Angeles 7, California
(Plate 40; 1 Textfigure)
During the summer of 1960, Mr. A. R. Whit-
worth of Carnarvon, Western Australia, sent me
three specimens of an unknown Cypraea from
Vlaming Head, Northwest Cape, Western Aus-
tralia, asking if they might belong to the sub-
species Cypraea cernica tomlini Schilder, 1930.
Mr. Bernard C. Cotton of the South Australian
Museum had identified one of these shells as C.
cernica Sowerby, 1870, referring to the illus-
trations of that species in Cotton and Steadman
(1946, p. 522) and Allan (1956, p. 94).
communication to Mrs. T. Hartley, May 1960.)
(Personal
To me, the shells were readily enough rec-
Ognizable as related to Cypraea cernica, though
they seemed to differ in a general way from
that species and from typical C. c. tomlini;
moreover, the type locality of C. c. tomlini
(Lifu, New Caledonia) is far enough removed
from Western Australia to give me further rea-
son for hesitating to put either name on the un-
known shells, Consequently, I asked Mr.
Whitworth for additional shells to study; he sent
me the remaining six in his collection. None of
the shells were live-taken (all having been
picked up on the beach after storms), but most
are in reasonably good to excellent condition,
Cypraea cernica at present comprises four
geographical races. Schilder (1938: p. 223, Map
1) has defined certain geographical provinces
and regions which have rather definite limits.
The "'races'' are groups of individuals of one
species separated by zones of nonoccurrence.
Cypraea cernica's four races (C. c. cernica, C.
c. tomlini, C. c.ogasawarensis, and C. c.mari-
elae) are located in the following Schilderian
Melanesian,
geographical regions: MLemurian,
Japanese, and Hawaiian, respectively.
Since the Western Australian form shows
certain morphological and color differences
from Cypraea cernica s.s., and since its typical
locality is in a completely different geographi-
cal area (the Dampierian Region) separated by
zones to the north and south in which no forms
of C,cernica are presently known to exist, it is
my opinion that the Western Australian speci-
mens belong to a new subspecies,
Cypraea (Erosaria) cernica viridicolor CATE, subspec. nov.
Shell solid, humped, pyriformly ovate, nar-
rowing abapically; base and sides rounded, right
and left margins excurvate, callous; inductura
distinctly and evenly pitted above marginal
edge; aperture fairly straight, narrow, curving
sharply left adapically; both terminals produced,
abapical terminal more so, Teeth fine, well de-
fined, centrally short on columella, lengthening
obliquely on the labial area adapically; outer
labial teeth heavier, longer, covering most of
outer lip except at central marginal edge; ter-
minal ridge extending across fossula; first four
or five teeth extending across and terminating
Dorsal
smooth, glossy, light olive-green,
prominently on adaxial edge of fossula,
inductura
very generously covered with irregularly sized
small white spots; white mantle line traverses
length of upper right dorsum; base, teeth, inter -
stices white; numerous large and small chest-
nut spots on upper marginal surface, ocellating
some white spots, and continuing over terminal
collars,
In Cypraea cernica viridicolor the shell is
more flattened and less humped than in C.c.
cernica, though approaching it more nearly than
in the other subspecies; the base from the left
margin to the columella is narrower and
straighter; the teeth are finer, shorter, and less
elevated, particularly on the outer lip; on the
fossula they are less numerous and weaker.
Cypraea c. viridicolor differs in being larger,
broader in relation to its length, flatter and
more solid in general. Its color is a lighter
greenish-beige, with the lateral spots much
larger, more numerous, and more distinct.
The name viridicolor stems from the Latin
viridis, meaning green, and color, meaning
hued, in reference to the peculiar greenish hue.
The type locality of Cypraea cernica viridi-
color is Vlaming Head, Northwest Cape, West-
ern Australia (21° 50'S. Lat., 114° 10'E,Long.).
Knowledge of its range is limited at this time,
with only two known collecting stations, Vlaming
Head and Quobba Point, approximately 40 miles
Page 176 THE VELIGER Vol. 4; No. 4
north of Carnarvon. recognized number of geographical races of
Cypraea cernica to five, two others being of
doubtful standing (C. c. percomis Iredale, 1931,
and C. c. prodiga Iredale, 1939). Starting in the
westernmost region with the typical species and
working in a more or less counter-clockwise
direction, these races may be enumerated as
Addition of the new subspecies brings the follows (see map, textfigure 1):
The holotype will be deposited in the Pale-
ontological Type Collection at Stanford Univer -
sity, Stanford, California (No. 9'506). Paratypes
and hypotype are in the respective collections of
C. N. Cate and A. R. Whitworth.
Subspecies Region Type Locality
Cypraea cernica cernica SOWERBY, 1870 Lemurian Mauritius
Cypraea cernica viridicolor subspec. nov. Dampierian Vlaming Head, NW Cape
Cypraea cernica tomlini SCHILDER, 1930 Melanesian Lifu, New Caledonia
Cypraea cernica marielae CATE, 1960 Hawaiian Maui, Hawaii
Cypraea cernica ogasawarensis SCHILDER, 1945 Japanese Bonin Islands
sh GASAWARA
PACIFIC OCEAN
AFRICA
\
“o \
!
|
, INDIAN
vy \OCE AN af
¢ ay
Be N.W. CAPE "4p .
VLAMING HEAD Ep a
\ 0777 AUSTRALIA -:3
‘2. prediee 0 Mey
Explanation of Plate 40
Cypraea cernica viridicolor C. CATE, subspec. nov.
Figure 1: Holotype; Figure 2: Hypotype No. 1; Figure 3: Paratype No. 3; Figure 4: Paratype No. 4;
Figure 5: Paratype No. 5; Figure 6: Paratype No. 6; Figure 7: Paratype No. 7; Figure 8: Paratype No. 1;
Figure g: Paratype No. 2. Hypotype No. 1 collected just north of Carnarvon, all others at Viaming Head;
Paratypes 1 and 2 are subfossil specimens. Photos by Takeo Susuki. (x 2)
[C. Cate] Plate 40
igure 9
Ax
Nee S mR ‘ 4 . E ee
Tue VELIGER, Vol. 4, No. 4
Takeo Susukl, photo.
Vol. 4; No. 4
THE VELIGER
Page 177
Table 1:
Measurements (in millimeters) of Types of Cypraea cernica viridicolor CATE, subspec. nov.
Vlaming Head Specimens
Length | Width | Height
Paratype 1 t+
Holotype
Paratype 2 +
Paratype 3
Paratype 4 *
Paratype 5
Paratype 6 * 17.5
Paratype 7 16.0
18.0
Outer Lip | Columellar Lip
Denticles
Location
C. N. Cate Collection
Stanford University Paleo.
Type Coll. No. 9506
C. N. Cate Collection
. Cate Collection
. Cate Collection
. Cate Collection
Cate Collection
Cate Collection
Quobba Point Specimen
Hypotype 1
{ Dead, subfossil specimen
Average length 20.4 mm.
Just before this issue went to press, a par-
cel was received from Mr. Whitworth contain-
ing 18 additional specimens of this subspecies.
All 18 were collected in beach drift at Vlaming
Head, North West Cape. The presence of a total
of 27 specimens, matching the morphological
characters mentioned above in all respects,
further corroborates my conclusions, I regret
that the statistics relating to the recently col-
lected specimens must be omitted here but
would like to note that one of these speci-
mens is larger than Paratype 1 listed in Table
1, measuring 31.0 mm. long, 20.1 mm. wide,
and 16.3 mm. high.
Acknowledgment
I wish to thank my friend and co-worker
Mr.Arch Whitworth of Carnarvon, Western Aus-
tralia, for his very helpful assistance and for
his generosity in sending specimens for study,
requesting that only one be returned to him;
also, to Jean Cate, whose actual work on this
paper was scarcely less than my own, I want to
express my devotion and grateful appreciation.
[25-9 17.2 | 12.8 17
* Subadult specimen
Average number of teeth: outer lip, 18; columellar, 15.
[ A. R. Whitworth Collection
Literature Cited
Allan, Joyce
1956. Cowry shells of world seas. pp. 1-170, pls. 1
to 15. Melbourne.
Cate, Crawford N.
1960. A new Hawaiian subspecies of Cypraea cernica
Sowerby. The Veliger 3 (1): 3-7; pl. 1.
Cotton, B. C., & W. K. Steadman
1946. Key to the classification of the cowries.
So. Austr. Mus., 8 (3): 503-530, pls. 8-13.
Iredale, Tom
1931. Australian molluscan notes.
18 (4): 217.
’
1939. Australian cowries, Pt. II.
9 (3): 297-323, pls. 27-29.
Franz Alfred
1930. Remarks on type specimens of some recent Cyp-
raeidae. Proc. Mal, Soc. Lond,, 19 (1): 49-58.
Franz Alfred, & Maria Schilder
Prodrome of a monograph on living Cypraeidae.
Proc. Mal. Soc. Lond., 23(3): 119-231, 16 textfigs.
===, Se :
1945, Westpazifische Cypraeacea von den Forschungs-
reisen des Prof, Dr, Sixten Bock. Ark. Zool.
Stockholm 36 A (2): 1-32.
Rec,
Rec. Austr. Mus.,,
Austral, Zoolog.,
Schilder,
Schilder,
1938.
Page 178
THE VELIGER
Vol. 4; No. 4
Nomenclatural Notes on some West American Mollusks,
with Proposal of a New Species Name
BY
A. MYRA KEEN
Department of Geology, Stanford University, California
A. Concerning Apolymetis
The type species of Apolymetis Salisbury,
1934, is Tellina meyeri Dunker, 1846, a rare
Oriental form. The original illustration shows
a lenticular shell with a very long narrow liga-
ment and regular concentric ribs. The only
points of resemblance to West American spe-
cies are a furrow down the posterior slope, a
large pallial sinus, and muscle scars unequal in
size. While revising the Tellinidae for the
"Treatise on Invertebrate Paleontology", 1 came
to the reluctant conclusion that the type species
of Apolymetis is so unlike all American species
that even if one made use of available subgeneric
names for grouping, the result would be incon-
gruous, Dr. Olsson in his ''Mollusks of the
Tropical Eastern Pacific’! (1961, pp. 410-414)
has suggested a lead by elevating Psammotreta
Dall, 1900, to generic rank. This would seem to
be a more plausible central group around which
to arrange certain West Coast forms now called
in part Apolymetis, in part Macoma. Olsson
prefers to recognize two separate genera, Flo-
rimetis and Psammotreta, but my feeling is that
more than two groups can be discerned, and to
call each a full genus would be to subdivide
more finely than we do in other pelecypod fam-
My rearrangement, then, would be:
ilies.
Genus Psammotreta Dall, 1900
Type species, Tellina aurora Hanley, 1844;
by original designation. Elongate to trigonal;
ligament sunken; posterior end weakly to
strongly bent.
Subgenus Psammotreta, s.s.
Elongate to ovate, smooth, flexure weak.
(P.) aurora (Hanley, 1844)
. (P.) mazatlanica (Deshayes, 1855)
. (P.) pacis (Pilsbry and Lowe, 1932) (Nota
synonym, as Olsson concluded.)
- (P.) plebeia (Hanley, 1844)
I'd It 9
ind
Subgenus Pseudometis Lamy, 1918
Type species, Tellina truncata Philippi,
1843 (non Linnaeus, 1767) = T. praerupta Salis-
bury, 1934; by subsequent designation, Salis-
bury, 1934.
More trigonal than P. (Psammotreta); flex-
ure weak.
P. (P.) dombei (Hanley, 1844)
P. (P.) grandis (Hanley, 1844)
P. (P.) gubernaculum (Hanley, 1844)
Subgenus Florimetis Olsson and Harbison, 1953
Type species, Tellina intastriata Say, 1826;
by original designation.
With subquadrate outline; flexure strong;
anterior end somewhat inflated.
. (F.) asthenodon (Pilsbry and Lowe, 1932)
. (E.) biangulata (Carpenter, 1856)
. (F.) cognata (Pilsbry and Vanatta, 1902)
(F.) cognata clarki (Durham, 1950)
OO hd Id
B. Tresus versus Schizothaerus
For the past 40 years or more, Tresus
Gray, 1853, has been considered unavailable, a
homonym. I am grateful to Dr. J. Lockwood
Chamberlin of the United States Bureau of Com-
mercial Fisheries, Washington, D.C., for hav-
ing pointed out to me that this is not true. Thus,
my revision of the Mactracea for the "Treatise
on Invertebrate Paleontology" is spared the er-
ror of accepting Tresus Walckenaer, 1833, as
valid in Arachnida even though two modern no-
menclators cite it (Sherborn's "Index Animali-
um'! adds a query), Consulting Walckenaer's
work, I found a French vernacular, 'Trésus",
that, moreover, is a nomen nudum, without cited
species or description. This leaves Gray's
Tresus available in Mollusca. Some question
has arisen as to whether it preceded Conrad's
Vol. 4; No. 4
THE VELIGER
Page 179
SS Te
Schizothaerus, but the evidence seems clear
that it has priority: Tresus Gray, Jan, 1853
(Anns Mag. Nat. Hist., ser. 2, vol. 11, p. 42;
type species, by monotypy, Lutraria maxima
Middendorff, 1849 [non Jonas, 1844] = L. nut-
talli Conrad, 1837) versus Schizothaerus Con-
rad, Feb. 7, 1853 (Proc. Acad. Nat. Sci. Phila-
delphia, vol. 6, p. 199; type species, by mono-
typy, L. nuttalli). Thus, the latter falls as a
synonym.
Although Tresus is an unfamiliar name to
younger workers, it represents a return to the
usage of more than a half-century, and one ad-
vantage of it is that it is both easy to spell and
easy to pronounce,
.
C. A New Species Name in EPITONIIDAE
My attention has been called by Mr. Richard
E. Petit of Ocean Drive Beach, South Carolina,
‘to the fact that the name Epitonium (Nitidiscala)
apiculatum Dall, 1917 (Proc. U.S. Nat. Mus., vol.
54, p. 480) — gastropod species no. 106 in ''Sea
Shells of Tropical West America'' — is a homo-
nym of Scala apiculata Dall, 1889 (Bull. Mus.
Comp. Zool., Harvard, vol. 18, p. 310), which
has been cited and figured as Epitonium (As-
periscala) apiculatum (Dall) by Clench and
Turner, 1952 (Johnsonia, vol. 2, no. 31, p. 290,
pl. 132, figs. 1-2). As this is a secondary rather
than a primary homonym, the name for the West
Coast form could stand were Asperiscala and
Nitidiscala treated as separate genera, for the
original name combinations were not identical:
Scala vs. Epitonium. However, as current prac-
tice does not favor such a fine subdivision of
these epitoniids, we are obliged to supply a new
name for the West American form.
The holotype of Epitonium apiculatum Dall,
1917, has not been figured. It was stated to be
in the collection of the United States National
Museum. Our concept of the species has been
based upon some figures published by Baker,
Hanna, and Strong in 1930 (Proc. Calif. Acad.
Scimuser 4, vol. 19,0p. Sil, pl. 3,, figs. 4-6) of
specimens they identified by use of Dall's de-
scription. Not having seen the holotype (which
evidently is an immature specimen) or any il-
lustrations of it, I prefer not to replace the
homonym but rather to bestow a name upon the
form figured by Baker, Hanna, and Strong.
Therefore, I propose Epitonium (Nitidiscala)
bakhanstranum Keen, sp. nov., this specific
name being an adjective coined by taking the
first three letters in each surname. To com-
memorate the work of this team is a pleasure,
for their joint paper on Epitoniidae is only one
of a series of very useful systematic reviews
on Panamic province molluscan groups.
A brief but adequate description of Epitoni-
um (Nitidiscala) bakhanstranum was given by
Baker, Hanna, and Strong (op.cit., p. 52). The
species is distinguished from the other Niti-
discalas with eight varices by the angular or
spinose shoulders of the varices, the slender
outline, the numerous whorls, and the angle at
which the varices diagonally ascend the spire.
The type locality is Carmen Island, Gulf of Cal-
ifornia, at the Salt Works. Height of the holo-
type is 9.2 mm., diameter, 3.3 mm. It is in the
Paleontological Type Collection, California
Academy of Sciences, No. 4'763; figured by
Baker, Hanna, and Strong (op.cit., pl. 3, fig. 4)
and refigured by Keen, 1958 (op. cit., p. 273, fig.
106). Hypotypes, Nos. 4'764 and 4'765 (Plate 3,
Figures 5-6) are from the La Paz area, Gulf of
California, as are some specimens in the Stan-
ford collection.
D. Names in L. Oken’s “Lehrbuch”?
The International Commission on Zoologi-
cal Nomenclature voted in 1956 (Opinion 417) to
place '"Okens Lehrbuch der Naturgeschichte,
vol. 3 (Zoology)'' on the Official Index of Reject-
ed Works. There seems to be no ground for
questioning the decision that this is a non-
binomial work, which means that all generic
names proposed therein have no status in no-
menclature as of the dates 1815-1816 and can
be validated only by a correct later usage. As
no summary list of Oken's names was published
by the Commission, except for those in certain
vertebrate groups, malacologists who do not
have access to a copy of Oken's work have been
slow to realize how many and which molluscan
names now have no standing. Three such have
recently come to my attention.
The name Arcinella Oken, 1815, is a syno-
nym in Carditidae, so that its loss there is un-
important. However, it no longer preoccupies
Arcinella Schumacher, 1817, of which the type
(by absolute tautonymy) is Chama arcinella Lin-
naeus, 1758 — the species that also is type of
Echinochama Fischer, 1887. Therefore, Echi-
nochama californica Dall, 1903, now becomes
Arcinella californica (Dall) and the Caribbean
E. arcinella becomes Arcinella arcinella (Lin-
naeus).
The second necessary change, called to my
Page 180
THE VELIGER
Vol. 4; No. 4
attention by Dr. Joshua L. Baily, involves the
name Irus Oken, 1815, in Veneracea, the type of
which has been accepted as Donax irus Linna-
eus, 1758, by tautonymy. This generic name
could have been credited to Gray, 1847, with
that species as type, but unfortunately Oken
himself used it in 1821 in his ''Naturgeschichte
fiir Schulen"', p. 647, with two species, neither
of which is veneracean; one is the type species
of Pandora Chemnitz, 1795 (Tellina inaequival-
vis) Gmelin (=Solen inaequivalvis Linnaeus,
1758), the other Mytilus rugosus, a doubtful
species of Hiatella, probably a variant of H.
arctica (Linnaeus, 1758). The former, ''Tellina
inaequivalvis", is here designated as type of
Irus Oken, 1821, which becomes a synonym of
Pandora, reverting to the usage of Blainville,
Deshayes, and other nineteenth-century writ-
ers. This leaves the veneracean group without
a name unless, (a) one petitions the International
Commission for protection of Irus Oken, 1815
— a procedure that on the average requires
approximately five years — or, (b) elevates one
of the named subgenera of the group to generic
rank (these would take precedence over any new
name that might be proposed). Three such
names are available: Notopaphia Oliver, 1923,
the type species from New Zealand, Venerupis
elegans Deshayes, 1854; Notirus Finlay, 1928,
type species, Venerupis reflexa Gray, 1843,
also from New Zealand; and Paphonotia Hertlein
and Strong, 1948, type species, Petricola
elliptica Sowerby, 1834, from Tropical West
America. The type of Notopaphia is elongate,
with fine concentric ribs anda sinuous ventral
margin. Perhaps this group might well be ele-
vated to generic rank. Notirus would then be
available as the generic name for the Irus
group. Notirus, s.s., would include those ovate-
quadrate forms with fine to coarse concentric
ribs, and Paphonotia would include those with
stronger radial sculpture. The genus ranges in
time from Oligocene to Recent in Europe but in
the Recent is mainly distributed in the Pacific,
with only one species, Donax irus Linnaeus, in
the Mediterranean. The Californian species
presently known as Irus lamellifer (Conrad,
1837) probably should be considered to fall in
Notirus, s.s., at least until such time as some-
one makes a thorough review of all known spe-
cies in the group.
The generic name Clathrus Oken, 1815, in
Epitoniidae, with Turbo clathrus [Linnaeus,
1758] as type by tautonymy, was (according to Dr.
Robert Robertson, in litt.) validated by Oken in
1821.
Literature Cited
Baker, Fred, G Dallas Hanna, & A. M. Strong
1930. Some mollusca of the family Epitoniidae from
the Gulf of California. Proc, Calif. Acad. Sci.,
ser. 4, 19 (5): 41-56, pls. 2-3.
Finlay, H. J.
1928. The recent mollusca of the Chatham Islands,
Trans. Proc. New Zealand Inst., 59: 232-286, pls.
28-43,
Hertlein, LeoG., & A. M. Strong
1948, Mollusks from the West Coast of Mexico and
Central America, pt. 6. Zoologica 33; 163 - 198,
pls. 1-2, :
International Commission on Zoological Nomenclature
1956, Opinion 417, Rejection for nomenclatorial pur-
poses of volume 3 (Zoologie) of the work by Lorenz
Oken entitled Okens Lehrbuch der Naturgeschichte,
published in 1815-1816. Opinions and Declarations
rendered by the I. C. Z. N., 14 (1): 1-42.
Keen, A. Myra
1958. Sea shells of tropical west America; marine
mollusks from Lower California to Colombia.
Stanford, Calif., Stanford Univ. Press; xi + 624
pp., illus.
Oliver, W. R. B.
1923. Notes on New Zealand pelecypods.
acol, Soc. London, 15: 179-188.
Olsson, Axel A,
1961. Mollusks of the tropical eastern Pacific, par-
ticularly fromthe southernhalf of the Panamic Pa-
cific faunal province (Panama to Peru). PartlI.
Panamic-Pacific pelecypoda. Paleontol, Res. Inst,
Ithaca, N. Y. pp. 1-574, pls. 1-86.
Proc, Mal-
Vol. 4; No. 4
THE VELIGER
Page 181
A New Method of Determining the Accuracy of Geotactic
Orientation of the Snail Helix aspersa MULLER
BY
DONALD R. BOWER
Department of Biology, San Francisco State College, San Francisco, California
(3 Textfigures)
Helix aspersa Miller progresses upward
when placed upon inclined surfaces. Thus, this
animal strongly demonstrates negative geotac-
tic orientation. Some of the*work dealing with
geotactic orientation has recently been reviewed
by Carthy (1958). The accuracy of orientation
was determined by the magnitude of the proba-
ble error of each animal's angle of orientation
(Crozier and Pincus, 1927) but not directly by
mathematical equations. Geotactic orientation
of H. aspersa was found by Cole (1927) to be
controlled by the equality of tensions produced
within the proprioceptors in the body muscula-
ture by gravity. The accuracy of geotactic ori-
entation of H. nemoralis has been found by
Hoagland and Crozier (1929) to decline in pro-
portion to an increase of the angle at which the
surface is inclined from the horizontal. They
also found that when this snail progressed on
either a vertical or a horizontal surface, the
longitudinal axis of the snail's body was paral-
lel to the longitudinal axis of the shell, but when
this alignment was changed by inclining the sur-
face, the snail turned until the two axes were
again parallel.
Equations have been used to describe di-
rectly the orientation paths of rats (Crozier and
Pincus, 1926, 1927), mice (Crozier and Oxnard,
1927), slugs (Wolf, 1927), and snails (Hoagland
and Crozier, 1929). The determination of an
animal's accuracy of geotactic orientation di-
rectly by mathematical equations shows that
this type of behavior is orderly and can be quan-
tified. This is essential if behavior patterns
are to be compared and if behavior patterns are
to have taxonomic value.
The purpose of this study is to develop a
series of mathematical equations that can be
used to describe quantitatively the accuracy of
geotactic orientation of an animal, and to deter-
mine the geotactic orientational accuracy of
Helix aspersa by these equations. The first
equation will describe the animal's true orien-
tation path in terms of the angle between the
animal and the horizontal, and the second equa-
tion will determine the animal's accuracy of
orientation in terms of the difference in degrees
between the animal's true orientation path, or
true bearing, and the angle of inclination of the
surface,
Materials & Methods
Specimens of Helix aspersa were collected
in Richmond, Contra Costa County, California,
from 24 September, 1960, to 29 May, 1961. The
animals were maintained on oatmeal, calcium
carbonate, and lettuce. Before being used for
experimental purposes, each snail was confined
for a period of between two and three days ina
quart jar with a substrate of half an inch of
small rock covered with one and one-half inches
of moist soil.
An apparatus was developed to determine
the angular deviations of the snail's path from
the vertical on the surface (see Figure l).
Figure 1: Diagram showing inclined surface and
expressions used in finding the degree of accuracy of
geotactic orientation: a = angle of inclination of the
surface; B = the angular deviation; » = angle between
the snail’s true path of orientation and the horizontal.
Method of scoring angular deviation: O - D = path
of snail; O - E = path of snail projected onto horizontal
surface; O - F = line on the surface perpendicular to
the horizontal when the surface ts inclined go°.
Page 182
THE VELIGER
Vol. 4; No. 4
Pieces of brown paper, 24 inches square, were
attached to a piece of cardboard the same size
and inclined at angles of 0°, 15°, 30°, 45°, 60°, 75°,
and 90° from the horizontal. The paper served
as a substrate on which the snails moved.
To track the snail's path of progression,
blue ink powder was dusted onto a piece of
brown paper which was used as a master sheet.
By pressing the master sheet on other sheets
of paper, enough ink powder was transferred
from the master sheet to these duplicates to
show a snail's track easily on the latter. Ten
duplicate sheets can be prepared without addi-
tional ink powder.
Each snail was placed upon the substrate at
its center with no uniformity as to the position
of the snail. At this time the snail was with-
drawn into its shell, and the surface was ina
After the snail had become
attached on the substrate, but before it could
extend itself, the surface was raised. While
performing the preceding steps, a 25-watt red
photographic safe-light bulb was used for illu-
mination at a distance of ten feet. The animals
were allowed to move for 30 minutes in total
darkness. The room temperature (20.0° to
23.7° C.) was recorded at the beginning of each
trial. To minimize adaptation, conditioning, or
learning that may occur with repeated testing,
no snail was allowed to score more than once.
horizontal position.
One hundred Helix aspersa were recorded
singly at each of the angles of inclination for
seven angles. The angular deviations from the
vertical on the surface (Figure 1, O-F) for each
slope of the surface were scored and their mean
calculated. These were plotted against the
angle of inclination of the surface. Each angu-
lar deviation was scored at a distance of nine
inches from the beginning of each run. This
distance was obtained by the chance selection of
a number from 1 through 12. The standard de-
viations and the range of each set of angular
deviations were also computed.
Geotactic accuracy was determined by
solving the following equation: a - w = degree of
accuracy, where ais the angle of inclination of
the surface, and w is the angle between the
snail's true orientation path and the horizontal
(see Figure 1). The true orientation path of the
snail was determined by solving the following
equation: sin w = sin a cos B, where B is the
mean of the angular deviation.
For each angle that the surface was in-
clined, the degree of accuracy was calculated
by using the equations presented above. The
degree of accuracy was plotted against the angle
of inclination of the plane.
Results
The results of the study are summarized in
Table 1, which shows the mean of the angular
deviations, the range of the angular deviations,
the value of w, and the degree of accuracy of
geotactic orientation for each angle of inclina-
tion of the surface.
For all the angles at which the surface was
inclined between 15° and 75°, the degree of ac-
curacy remained nearly constant with a value of
approximately 2°; however, the degree of accu-
racy decreased to a value of 8° 53' when the
surface was inclined at an angle of 90°. Since
in the control situation (0°) there was no verti-
cal component, it was not possible to score an-
gular deviations. In this case the snails moved
in an apparently random manner.
Figure 2, in which the means of the angular
deviations were plotted against the angle of in-
clination of the surface (except at 90°), shows
that as the angle of inclination of the plane in-
creased, the mean of the angular deviations de-
creased. The standard deviations and the range
of each set of angular deviations at each angle
that the surface was inclined are also shown in
Figure 2.
Table 1:
Mean and range of the angular deviations, values of w, and degree of accuracy for each angle of
inclination of the surface
Angle of inclination | Mean of angular deviations
of surface in degrees
15 28°21’
30 20°56’
45 15°38"
60 11°45!
75 T4u
go 8°53/
Range of angular deviations
Value of w
Degree of accuracy
o (control snails) moved in an apparently random manner
Vol: 4; No. 4 THE VELIGER Page 183
= range 9
50
8
45 =
g 40 ; 3
‘S <
A 35 S
cS : Bo
ea 3° mean e
Ss Q
20 ;
15 ,
10 lO} 2093040) 1508 CONN 70) (8onGo
Inclination of Surface
5
Figure 3: Degree of accuracy at each inclination of
HOME 203040) On OO ON GOGO Ute eurtace
Inclination of Surface
Figure 2: Mean angular deviation, standard deviation
and range of each set of angular deviations at each
angle of inclination of the surface. At each inclination
of the surface the vertical line indicates the total range
of the angular deviations; the broad portion of the line
indicates the standard deviation on each side of the
mean; and the crossbar, the mean.
Figure 3, in which the degree of accuracy
has been plotted against the angle that the sur-
face was inclined, shows, in graphic form, the
relation between the degree of accuracy of ori-
entation and the angle of inclination of the sur-
face.
Discusston
When accuracy of geotactic orientation is
determined by the equation a-w, the orientation
of Helix aspersa has been found to be consist-
ently accurate if progression of the animal
takes place on a surface inclined between the
angles of 15° and 75° from the horizontal; but
on a surface inclined at an angle of 90° from the
horizontal, the orientation of this snail is rela-
tively inaccurate. These results tend to dis-
agree with those of earlier workers who found
that orientational accuracy of various animals
decreased in direct proportion to log sin a or to
sin a when accuracy was determined in terms of
the magnitude of the probable error of each an-
imal's angle of orientation(Crozier and Pincus,
NAT)
The snail's true orientation path is a func-
tion of the snail's angular deviation at a given
angle of inclination of the surface. The degree
of accuracy is a function of the snail's true ori-~
entation path; therefore, these equations, a - w
= degree of accuracy, and sinw= sinacos B,
can validly be used to determine the accuracy
of a snail's geotactic orientation. The degree
of geotactic accuracy measures a snail's re-
sponse to gravity.
Summary
The geotactic accuracy of 700 specimens of
the pulmonate snail, Helix aspersa Miiller, was
determined. One hundred determinations were
made at each of the following angles: 0°, 15°
30°, 45°, 60°, 75°, and 90°. Geotactic accuracy
was determined by the equation: a - w = degree
of accuracy, where ais the angle of inclination
of the surface and w is the angle between the
snail's true orientation path and the horizontal.
w, in turn, was derived by the equation:
sin w =
sin a cos B, where B is the mean of the angular
deviations.
Geotactic accuracy had a value of about 2°
when a snail was orienting upon a surface in-
clined between the angles of 15° and 75° to the
horizontal but had a value of 8° 53'on a vertical
surface.
Acknowledgment
The author wishes to express his gratitude
to Dr. Jack T. Tomlinson for his guidance and
critical reading of the manuscript; also for the
helpful suggestions of Dr. Robert E, Berrend,
Page 184
THE VELIGER
Vol. 4; No. 4
ee
Dr. Walter J. Coppock, and others; and to Mr.
William Ho for assistance with the illustra-
tions.
Literature Cited
Carthy, J. D.
1958.
MacMillan Company, N. Y.
W. J., & T. T. Oxnard
Geotropic orientation of young mice.
141-149,
Crozier,
1927.
Gen. Physiol. 11}:
An introduction to the behavior of invertebrates,
Journ.
Crozier, W. J., & G. Pincus
1926. The geotropic conduct of young rats.
Gen. Physiol. 10; 257-269,
recat Renner arr & ’
1927. Geotropic orientation of young rats,
Gen. Physiol. 10: 519-524,
Hoagland, H., & W. J. Crozier
1929. Geotactic excitation in Helix.
Physiol. 15; 15-28.
Wolf, E.
1927.
iol.
Journ,
Journ,
Journ, Gen,
Geotropism of Agriolimax. Journ, Gen, Phys-
10: 757-765.
Observations on Three Species of Vexzllum (Gastropoda)
BY
JEAN M. CATE
Conchological Club of Southern California, Los Angeles 7, California
(Plates 41 to 44; 2 Textfigures)
It is a truism that with only a few speci-
mens froma given area, separate species may
be easily recognized, but with additional mate-
rial the lines of definition become less distinct
and the supposedly separable ''species'' appear
less so. This basic maxim of conchologists has
again proved itself in the case of three mitrid
forms all taken at one time in Philippine fishing
The three ''separate species’ re-
cently collected in a series of five trawled spe-
cimens were described as from widely separated
type localities (see map, Textfigure 1) and have
heretofore been considered distinct.
trawl-nets.
Mr. Fernando Dayrit of Manila has from
time to time sent me material for identifica-
tion. About a year agol published an illustrated
note (The Veliger, 3(4): 105-107) on a form that
I determined as Vexillum utravis (Melwvill, 1925).
Soon thereafter, two additional forms were col-
lected and sent to me that I tentatively identi-
fied as V. formosense (Sowerby, 1890) and V.
minahassae (Schepman, 1907), respectively. The
determination of V. minahassae was based in
part on the original description and type figure
and in part on a subsequent illustration and dis-
cussion of the species (Schepman, 1911). (Later
in the course of the present study, Miss G. E.
de Groot of the Rijksmuseum van Geologie en
Mineralogie in Leiden kindly furnished photo-
graphs of the holotype, which are reproduced
here on Plate 41.) In the case of V. formosense,
the identification was made not only from the
original description and figure but also bya di-
rect comparison (August 1961) with five speci-
mens from the type locality of that species in
the collection of the Academy of Natural Sci-
ences of Philadelphia (ANSP No. 251122. Anpin,
Explanation of Plate 41
Figure 1: Typefigure of Vexillum_ formosense (SowERBY, 1890) (x 1.5)
Figure 2: Holotype of Vexillum utravis (MELVILL, 1925)
Figure 3: Holotype of Vexillum minahassae (SCHEPMAN, 1907)
(x 1.5) Photo, courtesy of National Museum of Wales.
(x 3) Photo, courtesy of Rijksmuseum van Geologie
en Mineralogie.
Figure 4: Subfossil specimen of Vexillum minahassae (SCHEPMAN, 1907) dredged in 15 fathoms, Arafura Sea (Siboga
Expedition). (x 1.5)
Photo, courtesy of Zoological Museum of Amsterdam.
THE VELIGER, Vol. 4, No. 4 [ J. Cate] Plate 41
Pay
: NG" Z
© National Museum of Wales
Figure 1 Figure 2
© Zoological Museum of Amsterdam
© Rijksmuseum van Geologie en Mineralogie
Figure 3 Figure 4
Vol. 4; No. 4
THE VELIGER Page 185
RC,
8
MINDO
259
\S Q °
yx ¥ 130
PS v
CELEBES
SEA
Minahassa
2 Peninsulas™
: Kajoeragi
BANDA SEA
Figure 1: Map showing type localities of:
(a) Vexillum formosense (SOwERBY, 1890)
(b) Vexillum utravis (MELVILL, 1925)
(c) Vexillum minahassae (SCHEPMAN, 1907)
Formosa; ex Habe, Japan). These five speci-
mens seem identical in form with one of mine;
other Philippine specimens considered in the
present study are shorter and more obese,
though still falling within a range of variability
that may reasonably be expected. All three
species have been considered of rare occur-
rence, but during the past year V. utravis has
been collected in fair numbers at several local-
ities. With the exception of the ANSP shells al-
ready mentioned, however, and two specimens
of V. utravis misidentified as V. melongena
(Lamarck, 1811) in another large collection, I
have seen few of them in United States collec-
tions other than my own, and they are seldom
cited in the literature.
Vexillum formosense was described and
figured from three specimens collected off For-
mosa; Sowerby (1890) compared it with V. caf-
frum (Linnaeus, 1758), saying its whorls were
more rounded and the body whorl much shorter
in proportion to the spire. It should be noted
here that Sowerby concluded his remarks on his
new species by saying, ''Three specimens from
the Island of Formosa, all similarly marked,
and differing but little in form.'' That he noted
any difference in form, however slight, is im-
portant, as will be pointed out later. Because
of departmental reorganization currently taking
place at the British Museum (Natural History),
it is, unfortunately, not possible to obtain pho-
tographs of Sowerby's holotype, which is pre-
sumably at that institution. Consequently, the
rather poor type figure from the Journal of the
Linnean Society is here reproduced (see Plate
41, Figure 1).
Vexillum minahassae (see Plate 41, Figure
3) was described and figured by Schepman (1907)
from fossil or subfossil material collected on
Celebes. Apparently, the type lot consisted of
at least one complete specimen and several
fragments; it is not clear from his discussion
whether Schepman had more material than this,
One additional example, dredged in mud in 27
meters in the Arafura Sea (Elat, Great Kei Is-
land, Moluccas) was later reported and figured
by Schepman (1911, p.280, Pl. 12, Fig.7). The
color shown in this figure represents the typical
faded orange-red color of subfossil specimens,
whereas the Philippine shells before me are
live-taken, with fresh brown coloring. Although
the photograph of the fossil holotype of V.mina-
hassae seems to indicate a color-pattern simi-
lar to V. formosense, there is no mention of
Page 186
THE VELIGER
Vol. 4; No. 4
color in the original description. Schepman's
discussion of the Recent specimen collected on
the Siboga Expedition (Plate 41, Figure 4), how-
ever, mentions '"'...the shell is white, with a
rather broad orange-brown band below the su-
ture, another at the periphery, only partly visi-
ble on the upper whorls and a third at the base,
occupying also the canal, though less clearly.
The spirals of the base and partly the ribs, es-
pecially their upper part, are more or less
white."
Vexillum utravis (see Plate 41, Figure 2)
was described and figured from a single speci-
men of unknown locality; at that time Melvill
(1925) mentioned its possible relationship to V.
formosense, although obviously considering it
worthy of specific separation. He further sug-
gested that it might be a hybrid between V. caf-
frum (Linnaeus, 1758) and V. melongena
Lamarck, 1811). The latter two species are
compared briefly in J. Cate, 1961.
The morphological differences between
Vexillum utravis and V. formosense consist of
the relative obesity of V. formosense as com-
pared with the slender shape of V. utravis and
the ratio of spire-height to shell-length; this
character is extremely variable in V. formo-
sense, though in most of those I have seen the
spires are shorter than the last whorl. In typi-
cal V. utravis the spire is usually longer than
the last whorl or about equal to it. Color is
identical in both species, though there is some
variability in color-pattern among the slender
specimens that have been assigned to V. utravis
(see Plate 44, Figure 7), The most import-
ant difference between the two seems to be the
obesity of V. formosense.
A close relationship between Vexillum for-
mosense and V. minahassae is very apparent,
the chief difference between them being the
complete reversal of the color-pattern; V. for-
mosense is predominantly brown with white
spiral bands, while V. minahassae is mostly
white with brown bands.
is somewhat more obese in this instance also,
The sculpturing of both species is very similar,
Vexillum formosense
at the same time matching that of the third spe-
These similarities puzzled
me and were the basis for considering my iden-
cies, V, utravis.
tifications tentative, although each of the three
forms seems unquestionably to match the re-
spective descriptions and figures. The surface
Ornament is evidently the only truly constant
character in the morphology of the three spe-
cies. All three fall within the small vexillid
group having the dorsum smooth, axial costae
on the ventral surface or on the spire only, the
last whorl or two being entirely smooth in adult
specimens.
During the summer of 1961, Mr. Dayrit
mailed for identification five additional speci-
mens that had been trawled in the same area as
the pair of Vexillum minahassae sent a year
previously; that is, at Naval, Leyte. It was as-
tonishing to see that of the five shells taken at
one time, One specimen appeared to be a good
example of V. formosense, though unusually
short and obese (and possibly not fully mature),
three could be identified as V. minahassae, and
the fifth was somewhat intermediate between
these two but at the same time with a strong re-
semblance to the typical V. utravis (see Plate
43, Figures 1-5).
Receipt of this remarkable series led me to
request the loan of additional material, Mr.
Dayrit very kindly responded with a large group
of mixed specimens which had all been trawled
in Maqueda Bay during 1961. The box contained
five reasonably typical examples of Vexillum
minahassae and 32 adult and juvenile examples
of V. utravis; no specimens of V. formosense
appeared in this shipment. Mr. James E. Nor-
ton, also of Manila, sent several shells from
his collection that had been taken by divers in
from two to five fathoms in three different lo-
calities: Baler, Quezon; Masbate; and Batangas
Bay. I wish here to express my deep apprecia-
tion for the willing cooperation of both gentle-
men, as any opinion I might have formed without
their additional shells would necessarily have
been based on a study-group too small for a fair
evaluation of the species.
{It should perhaps be mentioned at this time that the
loaned material from both collections contained also sever-
al examples of an indeterminate species possessing certain
characters common to the other three under discussion,
The major part of the Norton shipment, in fact, consisted
of shells of this unidentified species, a few of these exhibit-
ing a strong resemblance to Vexillum melongena (Lamarck),
Explanation of Plate 42
Figure 1: Vexillum formosense (SoweRBy, 1890). Typical specimen resembling ANSP shells from the type locality.
Trawled in 20 to 40 fathoms, Tayabas Bay, Quezon, P. I.,
1960; ex Cate Collection. Figure 2: Vexillum formosense
(SowerBy, 1890). Specimen resembling V. minahassae (SCHEPMAN). Trawled in 20 to 30 fathoms, Naval, Leyte, P. L.,
1961; ex Cate Collection. Note similarity in form to V. formosense (fig. 1); the only difference is the arrangement of
color pattern.
Figure 3: Vexillum formosense (SowerBy, 1890). Series showing entirely ribbed specimens
before starting smooth last whorl.
THE VELIGER, Vol. 4, No. 4 [ J. Cate] Plate 42
Takeo SusukL, photo.
Vol. 4; No. 4
THE VELIGER
Page 187
12i°
“& Manila
124°
PHILIPPINE SEA
14°
Batangas Bay Ald...
= | /Maqueda Bay
oe
Figure 2: Map showing Philippine collecting areas discussed
although with some differences, The unknown form has been
seen from time to time in other collections and in earlier
shipments fromthe Philippines, but its identityhas remained
in question. It seems to be sufficiently distinct from the V.
formosense - utravis - minahassae group to allow for disre-
garding it as a part of the current study, though it should
not be overlooked as possibly belonging to a larger group of
species closely related to these. Further work is indicated
before it will be possible to place this speciesin its correct
position with relation to other Vexillidae. }
The Philippine localities where examples
of the three forms were collected encompass a
limited area, within approximately three and
one-half degrees of longitude and about two and
one-half degrees of latitude, or roughly within a
"circle'’ whose ''radius'"' fluctuates between 90
and 125 miles. Typical specimens of all three
species were taken in Carigara Bay, Maqueda
Bay, Ragay Gulf, Tayabas Bay, Batangas Bay,
Placer, and lastly, the locality where all three
forms were taken together, Naval, Leyte (see
map, Textfigure 2). According to Mr. Dayrit,
the fishermen who have collected these shells
do most of their trawling in Carigara or Ma-
queda Bays, going to the other localities only
when the weather is bad or if the catch has been
poor in these two places, The nature of the
equipment used limits their trawling operations
to depths of 20 to 40 fathoms; the three species
were all taken at these depths on a mud sub-
strate. Other mollusks collected at the same
time in these areas include Tibia fusus (Linna-
eus, 1758), T. powisi (Petit, 1842), Ficus gracil-
is (Sowerby, 1825), various species of Tonna and
Turris, and others.
Outside the Philippine area, good examples
of Vexillum utravis have been obtained in beach-
drift along the shore of Nadi Bay, Fiji, by Mr.
A. Jennings. This species seems to occur in
reasonably large numbers there, as Mr. Jen-
nings reported (March 1961) having taken ap-
proximately 40 dead specimens on one collecting
trip. Only one living example has been report-
ed at the present time, however (dredged in 10
feet of water on a weedy mud bottom in Nadi
Bay, December 1961), and the Fiji material is
mentioned here only to point out the occurrence
of the species in at least one area remote from
the Philippine Islands (a distance of approxi-
mately 4'300 statute miles). Much further study
Page 188
THE VELIGER
Vol. 4; No. 4
is needed of the Fiji specimens, and it is hoped
that living populations will be discovered there
in the near future. It will be of great value if
collectors in other areas will report any occur -
rences, with full collecting data wherever pos-
sible, of any of the species illustrated in this
paper, in order to help establish the extremes
of range for these hitherto little-known forms.
In frequency of occurrence, Vexillum utra-
vis is relatively the most common of the three
forms and is collected in fairly large numbers
at several locations; it has also been taken in
shallower water (two to five fathoms) by divers,
as seems not to be the case with either V. for-
mosense or V. minahassae. Vexillum formo-
sense is the second most frequently collected
form, and V.minahassae the least often encoun-
tered in the areas mentioned (F. Dayrit, per-
sonal communication).
Most of the specimens of Vexillum formo-
sense are entirely smooth on the last two
whorls, while a few are ribbed ventrally and
smooth dorsally as in typical apparently adult
V. utravis (see Plate 44, Figures 4-6). The
possibility exists that the specimens of V. for-
mosense with completely smooth last whorls
represent animals more mature and more fully
developed than those whose shells are ribbed
only on the ventral surfaces. There is obvious-
ly a point where the costae disappear altogether,
in this case at approximately the tenth whorl
(out of a total of 12). In V. utravis, of which I
had a far greater number of specimens to study,
the smooth dorsums (that is, the final whorls)
seem to occur at about the eighth, ninth, or
tenth volutions. A limited few are seen with en-
tirely smooth final whorls, though these seem
to be less mature than the smooth examples of
V. formosense, as the outer lips are thin and
underdeveloped, and the canals are less attenu-
ate. It is possible that these specimens were
collected while undergoing a period of growth
when the outer lip had not yet thickened; this
stage has been observed in specimens of vari-
ous size-groups in other mitrid species; there-
fore, the presence of a thin outer lip does not
necessarily indicate a completely juvenile spe-
cimen.
It is interesting to note that the majority of
living specimens of Vexillum utravis I have
seen have been collected at precisely the stage
of transition from the ribbed to the smooth
state. Out of 43 specimens studied, 28 were
ribbed on the ventral side and smooth dorsally;
nine were ribbed all around and appeared to be
juveniles (see Plate 42, Figure 3), and six were
entirely smooth on the final whorls though not
yet fully mature as to the outer lip. One possi-
ble explanation for the imbalance in these sta-
tistics might be that the more mature (i.e.,
smooth-shelled) animals may seek a slightly
different environment, such as a greater depth
of water, which would be beyond the ordinarily
limited trawling-depths of the fishermen who
have taken most of the known partly-ribbed spe-
cimens. In each of the three species there ap-
pears to be a strong tendency to become rather
exaggeratedly elongate with advancing maturity.
Juveniles frequently appear relatively more
obese in proportion to their length, and as the
number of whorls increases, the canal becomes
far more attenuate than formerly, bringing
about a change in the spire-height/shell-length
ratio. In other words, as the mollusk grows, the
increase in length is more pronounced in the
canal than elsewhere, resulting ina relatively
shorter spire and a longer body whorl than in
younger specimens.
Measurements of shell length, greatest
diameter, aperture length, and spire angle were
taken for all specimens of each of the three
species, but these provided no helpful data ex-
cept to prove that the proportions and angles of
all three forms areas variable as their mor-
phological characters; no other conclusions
could be drawn from any of several approaches.
The measurements are not included here, as
they provide no conclusive data. The only no-
ticeable trend resulting from this part of the
study is that, in general, the spire-angle tended
to decrease with increase of shell length in
Vexillum formosense and V. utravis, while this
was not evident in V. minahassae.
Since the Philippine shells were all trawled
in depths from 20 to 40 fathoms, and the 1911
Schepman record of Vexillum minahassae indi-
cates a depth of approximately 15 fathoms in the
Arafura Sea, it would seem that all three forms
live in relatively deep water as a rule and that
the mud substrate is an ecological feature com-
mon to all. The surface ornament is nearly
Explanation of Plate 43
Vexillum formosense (SOWERBY, 1890)
Series of five specimens collected together at Naval, Leyte, P. I., 1961. |
Figure 1 resembles typical but obese Vexillum formosense (SowERBY); Figures 2 to 4 resemble typical V. minahassae
(ScHepmaNn); Figure 5 shows a tendency toward typical V. utravis (MELVILL).
[ J. Care] Plate 43
Tue VELIGER, Vol. 4, No. 4
WOE
TAKEO Susuxl, photo.
Vol. 4; No. 4
identical in each, there is a pronounced variabil-
ity in shape among otherwise typical examples
of V. formosense collected within a narrow
range, and the reported collecting stations of
all three forms are in very close proximity. On
the basis of all these factors considered togeth-
er, and particularly taking into account not only
the evidence of close association of the three
forms collected within the same area at one
time, but also the strong suggestion that the
transition between partly ribbed and entirely
smooth specimens occurs at approximately the
Same growth stage in all ''species",, the conclu-
sion that these forms constitute only one very
variable species is almost inescapable, the
many variations in color and shape possibly
being the result of some unknown factor or
factors of ecology. Vexillum formosense,
therefore, would represent the northernmost
phenotype within its range, V. minahassae the
southern extreme, and V. utravis is intermedi-
ate, its typical locality being apparently mid-
way between the other two.
The possibility that hybridization of two al-
lopatric subspecies may be taking place here is
not excluded; however, until further locality
records for large numbers of specimens be-
come available both to the north and to the south
of the presently known collecting stations, it
would be premature to draw a conclusion, Con-
versely, it might be considered equally possible
that through reproductive isolation, separate
species are presently evolving.
The additional possibility that sexual di-
morphism may account for the variability
among the three forms should not be overlooked,
though the preponderance of the "'utravis'' form
suggests that this is not the case. If sex differ-
ences were the chief basis for the morphologi-
cal differences, it is likely that there would be
a more equal balance in numbers among the
different phenotypes. The present ratio of 43
utravis to 3 formosense would probably not ap-
ply to the relative distribution of males and fe-
males; as a hypothetical example, it seems an
unlikely assumption that we would find 43
"“utravis-type females''and only 3''formosense-
type males" (or vice versa) within a represen-
tative sampling of the population. It is hoped
that a thorough anatomical study can be under-
taken in the near future, to make possible a
comparison of the physical features of the va-
rious animals.
A brief list of questions relating to Vexil-
lum formosense was sent to Dr. Tadashige
Habe in Japan, in an effort to ascertain whether
he had noted any similarities to the other two
THE VELIGER
PT Nene ee aE eee ee Se SSE se ee al
Page 189
species discussed here, either in morphology
Dr. Habe very kindly responded
to my queries with several interesting observa-
or in ecology.
tions. I am very grateful to him for his atten-
tion to my problem and wish to express my
thanks for his prompt and helpful reply. A color
illustration of V. formosense has been included
in Habe (1961; Pl. 34, fig. 18).
According to Dr. Habe's letter of Decem-
ber 30, 1961, Vexillum formosense appears to
be rather common off Formosa. It is his belief
that it is probably collected in about 20 to 50
meters (10 to 25 fathoms) on a fine sand sub-
strate; it has been recorded from Okinawa
(Kuroda); and Dr. Habe suggests the northern
limits of the species may be the Amami Group
of the Ryukyu Archipelago (28°15'N. Lat., 129°
15'E. Long.). He had no data as to other mol-
lusks actually collected with V. formosense but
stated that he had received certain other spe-
cies within the same shipments, which suggests
that they might all have been collected together;
these included V. melongena (Lamarck), Bary-
spira mammilla (Sowerby), Niotha clathrus (La-
marck), Ficus ficus (Linnaeus), Bezoardicella
decussata (Linnaeus), Murex trapa (Réding),
Murex rectirostris Sowerby, Hindsia sinensis
(Sowerby), etc.
It is Dr. Habe's conclusion that Vexillum
utravis and V. minahassae are not conspecific
with V. formosense, but closely related to V.
melongena (Lamarck) from Formosa, '',..which
has more numerous costae on the surface".
the light of the very recent evidence of several
Philippine specimens of both V. utravis and V.
minahassae also exhibiting entirely smooth last
whorls, as in V.formosense (see Plate 44, Fig-
ures 4-6), my own conclusions remain as stat-
ed; however, the observations from Japan lend
emphasis to my earlier statement that further
work should be done regarding the relationships
of other similar species, including V. melon-
gena in particular. This species or a very
closely related form seems to be associated
with the others discussed here, in Fiji and the
Philippines as well as Japan.
In
The different forms, taken separately, are
easily enough accepted as three distinct spe-
cies, but if all three can be collected within the
confines of a single small bay, they must exist,
at least in that area, as a single population and
not as the remotely situated, separate species
they were formerly thought to be.
If the different forms are to be combined
as one very variable species, the earliest valid
name should apply. It is my suggestion, there-
Page 190
fore, that Vexillum minahassae (Schepman,
1907) and V.utravis (Melvill, 1925) be consid-
ered conspecific with V. formosense (Sowerby,
1890). If the theory of newly-evolving species
is accepted, V.minahassae and V.utravis could
be considered subspecies of V. formosense as
a possible temporary solution until additional
work may be done which can establish without
further doubt the correct taxonomic position of
these similar species. However, the evidence
of overlapping populations, among other consid-
erations, seems to support the conclusion that
the three forms all belong to one variable spe-
In that case, an amendation of the origi-
nal description of V. formosense is necessary
in order to include the variants within the new
interpretation of the species.
cies.
Vexillum formosense (Sowerby, 1890), amend.
Shell spindle-shaped, spire elongate, acute;
whorls approximately 11, convex, separated by
impressed sutures. First 8 or 9 whorls ribbed
longitudinally, interstices between riblets fine-
ly spirally striate. Last 2 or 3 whorls very
finely spirally striate but not ribbed; rounded,
swollen at periphery; neck of shell spirally
granulose, contracted, somewhat attenuate.
Ratio of aperture length to spire height varia-
ble, aperture longer than spire in mature spe-
cimens, about equal in subadults. Peristome
entire; pseudumbilicus present; parietal callus
prominent; canal produced, recurved abaxially.
Labrum sinuate in adult specimens, otherwise
simple; finely lirate within. Columella with 3
or 4 plaits, the upper two somewhat grooved and
flattened, the surface between plaits showing
granulose structure underlying thin coating of
nacre.
Shell color variable; typical specimens
dark brown (Maerz & Paul Dictionary of Color,
2nd Edition, 1950: Pl. 8, C9, London Smoke) to
light brown (Pl. 7, A 11, Vandyke Brown) with
one white band on each upper whorl, usually two
on periphery of last whorl. Some specimens
reversed in color, being basically white with
brown bands. Aperture, columellar folds and
parietal callus white, granulose neck of shell
whitish.
THE VELIGER
Vol. 4; No. 4
Length of holotype 50 mm., greatest diam-
eter 16 mm., aperture length 24 mm., and aper-
ture width 4s mm.
Type locality: Island of Formosa (24° N,.
Wates i 2) leElsonee)is
Acknowledgment
In the study reported on in the foregoing
pages, I have received assistance from many
persons. My thanks are expressed here to all
of them; first, to Mr. Dayrit for providing not
only large numbers of shells for study, but
much background information as well; also, to
Mr. James Norton and Mr. A. Jennings for ad-
ditional specimens and information. Dr. Rudolf
Stohler very kindly provided the necessary
Schepman and de Man translations, as wellas
fundamental information in molluscan zoology;
Mr. George Kanakoff translated Sowerby's Lat-
in description; Miss Virginia Orr directed me
to pertinent material at the Academy of Natural
Sciences of Philadelphia; Dr. Myra Keen fur-
nished welcome criticism of the preliminary
manuscript; and Crawford Cate, as always,
assisted in countless ways. Mr. Colin Mathe-
son of the National Museum of Wales provided
the photographs of Melvill's holotype; Miss G.
E. de Groot of the Rijksmuseum van Geologie
en Mineralogie in Leiden furnished photographs
of Schepman's fossil holotype; and Mrs. W.S.S.
Van der Feen-v. B. Jutting of the Zoological
Museum at Amsterdam sent photographs of the
Recent specimen of Vexillum minahassae col-
lected on the Siboga Expedition. Dr. Tadashige
Habe's important information should not be
overlooked in this acknowledgment of the many
kindnesses performed by all these people, nor
should the help of Mrs. Emily Reid of The Veli-
ger staff, who adapted the complicated maps
into textfigures. Unless otherwise noted, the
photographs are by Takeo Susuki. It is difficult
to find words adequately to express the grati-
tude due to everyone mentioned.
Literature Cited
Cate, Jean M.
1961. Vexillum utravis (Melvill, 1925) trawledin Phil-
ippine waters. The Veliger 3 (4): 105-107; pl. 18,
1 textfig.
Explanation of Plate 44
Vexillum formosense (SOWERBY, 1890)
Series showing variability in form and pattern. Note presence of only one white band in some specimens.
Figure 1: a juvenile specimen, entirely ribbed. Figures 2, 3: in transitional stage, half ribbed, half smooth.
Figures 4, 5, 6: entirely smooth on last whorls. Figure 7: series showing variability in color pattern; a to c, “uéravis”
form; d to f, “minahassae” form. All specimens of Plates 43 and 44, and of Plate 42, fig. 3 ex Dayrit Collection,
trawled in 20 to 30 fathoms, Maqueda Bay, Samar, P. I., 1961.
THE VELIGER, Vol. 4, No. 4 [ J. Cate] Plate 44
Figure 7 a
TAREO Susukl, photo,
Vol. 4; No. 4
THE VELIGER
Page 191
de Man, J. G.
1904, Beschreibung einiger brachyurer Krebse aus
posttertidren Schichten der Minahassa, Celebes.
Samml. Geol. Reichs-Mus. Leiden, Ser. 1, vol. 7:
254-278; pls. 9, 10.
Habe, Tadashige
1961.
pp. 1-183; 66 pls. Osaka, Japan.
Melvill, James Cosmo
1925. Descriptions of nine new species of Mitridae.
Proc. Mal. Soc. London, 16 (5): 215-219, pl. 10,
Colored illustrations of the shells of Japan (II).
Schepman, M. M.
1907, Mollusken aus posttertidren Schichten von Cele-
bes. Samml, Geol, Reichs-Mus. Leiden, Ser. 1,
vol, 8: 153-203, pls. 10-13.
1911. The Prosobranchia of the Siboga Expedition, pt.
4, Mitra: 266-290; pls. 18, 19, 22.
Sowerby, George Brettingham
1890, Descriptions of fourteen new species of shells.
Journ. Linn. Soc., 20: 395-400, pl. 25,
Manometric Measurements of Respiratory Activity
in Tegula funebralis
BY
JAMES H. McLEAN
Department of Biological Sciences, Stanford University, California
Tegula funebralis (A. Adams, 1855) is an
abundant gastropod of the middle and upper in-
tertidal zones on partially protected rocky
shores in central California (Galli and Giese,
-1959). It is uncommon on the exposed outer
coast and does not occur in subtidal zones, Pop-
ulations of Tegula are covered by the tide at
least once during the daily tidal cycle. Individ-
uals may also remain submerged in tide pools
where temperature and salinity fluctuate.
In order to assess some of the limiting con-
ditions in the environment of Tegula funebralis,
I made brief laboratory measurements of oxy-
gen consumption under varying conditions of
temperature, salinity, and exposure to air.
I wish to thank Dr. Arthur C. Giese for
helpful suggestions.
Materials G Methods
The Tegula were collected on rocks in the
upper tidal zone near Hopkins Marine Station,
Pacific Grove, California, and maintained in
laboratory aquaria supplied with running sea-
water. Snails used in each experiment were
collected at the same time and subjected to the
Same conditions. Oxygen consumptions were
measured in a series of six Warburg manome-
ters, using immature snails with shell diame-
ters not exceeding 14 mm., the largest size
which could be fitted into the manometer ves-
sels. Each vessel was calibrated for use with
three snails submerged in five ml. of filtered
seawater. A 0.1 ml. drop of 30 percent potas-
sium hydroxide in the sidearm of the vessel was
used as the carbon dioxide absorbent. The ves-
sels were gently agitated to the same extent
during each run.
Size constants were determined with 18
snails, 10 to 14 mm. in diameter. The wet-
shell weight of the 18 snails was 19.69 grams,
and the wet-animal weight after removal of the
shell was 2.30 grams. Dry weight of the snail
tissue was determined with an analytical bal-
The dry weight of the snails
used in each experiment was therefore calcu-
lated as 0.058 of the wet-shell weight. During
each experiment weight corrections were made,
and oxygen consumption of the three snails in
each vessel was calculated and averaged. Oxy-
ance as 621 mg.
gen consumptions are given as microliters of
oxygen per milligram of dry weight.
Experimental Results
1. Respiration while Submerged and Exposed
to: Air.
In its high intertidal environment Tegula
Page 192
THE VELIGER
Vol. 4; No. 4
a
funebralis spends long periods exposed to air.
The respiratory activity of a series of snails
was measured in three ways: while kept moist;
submerged; and after five hours exposure to
air. All runs were made at 19°C. during Aug-
ust, 1961, using 15 snails collected eight days
previously. In the first experiment a thin film
of water covered the snails, which remained at-
tached to the walls of the vessels. A second
series of readings was made 18 hours later with
the same snails in 5 ml. of seawater per vessel.
A third series of readings was taken with no
water in the vessels, after the snails had been
left to dry on paper towels at room tempera-
ture for five hours. The snails were able to at-
tach to the walls of the vessels in this condi-
tion. Results are shown in Table l.
Table 1:
Oxygen consumption (microliters per mg. dry weight) of
Tegula while wet, submerged and after 5 hours drying.
2 Respiration under Varying Conditions of
Salinity.
Salinities in tidepools in which Tegula often
occur may fluctuate with the seasons. Snails in
this series of experiments were held submerged
in beakers containing seawater of different con-
centrations for 24 hours before the readings
were made. Readings at each concentration
were taken simultaneously in order to minimize
The six vessels contained
three snails at each of six different concentra-
tions. The experiments were then repeated with
different snails to verify data based on only
three individuals at each concentration. Four
experiments with four different series of snails
were made: two experiments with concentra-
tions ranging from 100 percent to 50 percent
seawater and two experiments with concentra-
tions ranging from 100 percent to 150 percent
seawater. This series of experiments was con-
ducted during November, 1961, on snails col-
other variables.
Ciena | lected two to four days previously. Results are
(min.) Wet Submerged After Drying shown in Table 2.
|_ 30 Des | tO SO ei 05 Aneel Dee SEE The data show that respiration is relatively
60 0.77 _ 993 | 9-49 normal at salinities ranging from 80 percent to
of | LOS) igs | bo 130 percent seawater, while tapering off at con-
The data indicate that maximum respira-
tion occurs while the animal is submerged. Al-
though Tegula funebralis is strictly an intertidal
animal, daily exposure to air is not necessary
for survival. Twenty-two large specimens were
kept submerged under glass dishes in an aqua-
rium supplied with running seawater. All re-
mained alive and active for over 40 days at the
tops of inverted glass dishes. Thus, it appears
that a negative geotropic response maintains
Tegula in the intertidal zone.
centrations of 70, 60, and 50 percent, andat
140 and 150 percent seawater. Respiration at
50 and 150 percent seawater was negligible.
Snails at these concentrations did not attach to
the vessel walls during the run. Kept at the
same concentrations, they did not survive be-
yond one week. Snails in 60 percent seawater
died within two weeks. Snails in 70 to 140 per-
cent seawater survived longer than three weeks.
These results suggest that Tegula can accom-
modate to a relatively wide range of salinities
(80 to 130 percent) with little loss of activity.
Table 2:
Oxygen consumption (microliters per mg. dry weight) of Tegula at salinities ranging from 50% to 150% seawater.
Figures are given for the original and the repeated run of each experiment.
Salinity i 100% 90% 80% 60%
go min 0.29|0.36 0.31|0.39 0.31|0.35 00.4|0.17 0.02 |0.03
60 min 0.56|0.75 0.57|0.68 0.55|0.69 0.13|0.22 0.04|0.05
go min 0.87| 1.02 0.90|0.96 0.81|1.01 0.18|0.27 0.07|0.09
120 min 1.14|1 29 1.18] 1.22 0.99| 1.32 0.21|0.31 0.07|0.12
Salinity [ 100% 110% 120% 140% [
30 min 0.20|0.30 0.29|0.32 0.25|0.27 0.10|0.30 0.03|0.06
60 min 0.43|0.52 0.50|0.60 0.49|0.50 0.55 0.16|0.53 0.04|0.09
go min 0.62|0.80 0.72|0.91 0.66|0.74 0.82 0.20|0.70 0.06|0.14
120 min 0.95| 1.06 1.10|1.12 0.86|1.01 0.25|0.81 0.10|0.15
Vol. 4; No. 4
THE VELIGER
Page 193
3. Respiration under Varying Temperatures.
A series of 18 snails was run for one hour
at temperatures of 11°, 19°, 27°, and 35° C.
during August, 1961. One hour was allowed for
equilibration between 11° and 19°, two hours
between 19° and 27°, and four hours between
27° and 35°. Average oxygen consumption of
the 18 snails at each temperature is shown in
Table 3.
Table 3:
Oxygen consumption (microliters per mg. dry weight) of
Tegula at temperatures ranging from 11° to 35° C.
Between 11° and 27° the respiratory rate
appeared to increase directly with temperature.
The upper limit of respiratory efficiency was
reached between 27° and 35°, for the rate at
35° fell off to slightly less than the rate at 19°
4. Effect of Starvation upon the Respiratory
Rate.
The oxygen consumption of six animals col-
lected 20 days previously was compared to the
oxygen consumption of nine animals collected
one day previously (August, 1961). The results
are shown in Table 4.
Table 4:
Oxygen consumption (microliters per mg. dry weight)
of Tegula, starved and fresh
Starved animals respired at slightly more
than half the rate of the freshly collected ani-
mals. These results emphasize the importance,
in experiments of this kind, of using animals
collected at the same time and subjected to the
same conditions.
Conclusions
These short determinations have indicated
that respiratory activity of Tegula varies not
unexpectedly with conditions of exposure, salin-
ity, temperature, and starvation. Basic rates
of oxygen consumption are comparable to those
given for molluscan tissues by Zeuthen (1947).
Additional factors which have been shown to af-
fect respiratory activities of gastropods are
seasonal variations (Berg, Lumbye and Ockel-
mann, 1958) and tidal and diurnal rhythms (San-
deen, Stephens and Brown, 1954). The shortness
of the manometric determinations used here
served to obscure rhythms that might have been
present. Respiratory activity may also be ex-
pected to vary with the intertidal height from
which the snails are collected. Manometric de-
terminations of oxygen consumption, therefore,
are useful only with snails collected at the same
time and measured in the manometer simul-
taneously.
Literature Cited
Berg, K., J. Lumbye, & K. W. Ockelmann
1958. Seasonal and experimental variations of the oxy-
gen consumption of the limpet Ancylus fluviatilis
(O. F, Miller), Journ. Exp. Biol. 35: 43-75,
Galli, D. R., & A. C. Giese
1959. Carbohydrate digestion in a herbivorous snail,
Tegula funebralis, Journ. Exp. Zool. 140: 415 to
440,
Sandeen, M. I., G. C. Stephens, & F. A. Brown, Jr.
1954, Persistent daily and tidal rhythms of oxygen
consumption in two species of marine snails.
Physiol. Zool. 27: 350-356.
Zeuthen, Erik
1947. Body sizeand metabolic rate in the animal king-
dom with special reference to the marine micro-
fauna. Compt. rend, Lab. Carlsberg, Sér. chim.
26 (3): 16-161,
Page 194
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Vol. 4; No. 4
A Study of Food Choices of Two Opisthobranchs,
Rostanga pulchra McFartanp and Archidoris montereyensis (Cooper)
BY
EMILY F. COOK
Department of Zoology, University of California, Berkeley 4, California
(4 Textfigures)
Several of the dorid nudibranchs of the
west coast of North America feed on sponges
which they reseimble in color and texture. Ros-
tanga pulchra MacFarland, a bright red nudi-
branch, 1.0 to 1.5 cm. long, feeds on an en-
crusting red sponge, Ophlitaspongia pennata
Lambe, while Archidoris montereyensis (Coop-
er), a large yellow nudibranch, 6.0 to 10.0 cm.
long, commonly feeds on a yellow sponge, Hali-
chondria panicea (Pallas). Both the nudibranchs
and the sponges on which they feed are found
intertidally along rocky coasts.
In the Friday Harbor region (San Juan Is-
land, Washington), where this work was done,
the sponge, Ophlitaspongia pennata, is repre-
sented by the varietal form, O. pennata var.
In order to distin-
guish var. californiana measurements of the
spicules must be made. Although this was not
done, Mr. Gerald J. Bakus, who has worked on
the sponges of the area, has assured me that the
odds are about 100 to 1 that it will be the vari-
etal form, since he has not found the typical
form itself in the San Juan Archipelago.
californiana de Laubenfels.
In the field Rostanga pulchra is often found
on red sponge and is seldom found far away
from it. The sponge encrustations are 2 to 3
mm. thick and vary in area from 2 or 3 sq. cm.
to over 100 sq. cm. It is scattered in its dis-
tribution along the coast.
Archidoris montereyensis is commonly
found on Halichondria panicea but may be found
where sponges appear to be absent. Halichon-
dria panicea is a branching yellow sponge of a
fairly loose composition, often found in large
clumps. In the literature it is often referred to
as the ''bread crumb'' sponge.
The purpose of this paper is to report on
some aspects of feeding of Rostanga pulchra
and Archidoris montereyensis,
This work was done at the Friday Harbor
Laboratories, University of Washington, parti-
ally supported by N.S. F. Grant No. G-7045. I
would like to express my appreciation to Dr.
Gunnar Thorson for his many helpful sugges-
tions during the course of this work, and to Dr.
Cadet Hand for his criticism of the manuscript.
Materials & Methods
Ie
Rostanga pulchra to determine factors import-
ant in location of food. A plastic bowl 10 inches
in diameter was set up so that two currents
could be directed into it. An overflow was
placed between the two incoming currents (Fig-
ure 1). The bowl was covered with black rubber
Three sets of experiments were done with
— plastic bag
with sponge
Figure 1
Apparatus used to test the response of Rostanga pulchra
to food and current. The arrows indicate the direction
of the current.
Vol. 4; No. 4
THE VELIGER
Page 195
on the outside. During all experiments the bowl
was covered with heavy cardboard to eliminate
light. Screw clamps were used to adjust the
flow of incoming water.
except (a), 12 animals were started in a watch
Each
The cover was
For each experiment,
glass midway between the two currents.
experiment lasted 30 minutes.
lifted at 10-minute intervals to observe the
progress of the animals.
(a) Two experiments, using 12 and 13 animals,
were done to test the response of Rostanga
pulchra to current. No sponges were used. The
two currents were adjusted so that one was very
weak, and the other was much stronger. For
the second experiment the currents were re-
versed.
(b) Six experiments were done to test the abil-
ity of Rostanga pulchra to find Ophlitaspongia
Plastic bags which contained O, pen-
pennata.
nata and Halichondria panicea were tied in the
incurrent water. The currents were adjusted to
equal size. Both currents were small to reduce
swirling of water in the dish. The sponges were
switched from one current to the other between
experiments.
(c) In order to compare the response to cur-
rent and to the sponge, an experiment similar to
(a) was set up with Ophlitaspongia pennata in the
weak current.
Il. Since Archidoris montereyensis is much
larger than Rostanga pulchra, a larger appara-
tus was required, A tank 16 by 20 inches was
set up similarly to that used for R. pulchra.
Responses to food and current were tested as
described in I(a) and I(b).
strong | ~ ? | slight
current current
Figure 2
Distribution of Rostanga pulchra in response to current
Ophlitaspongia
pennata
Halichondria
: Gi panicea
Ty
on ,
Figure 3
Distribution of Rostanga pulchra given a choice between
Ophlitaspongia pennata and Halichondria panicea
Ill, Both nudibranchs were placed on different
sponges to determine which they would eat. The
sponges used were Halichondria panicea, Oph-
litaspongia pennata, Syringella amphispicula,
Aplysilla glacialis, and an unidentified suber-
itid.
Results
I(a) The results of experiments to determine
the response of Rostanga pulchra to current in-
dicate that it will move towards the current.
The distribution of animals in the apparatus at
the end of the experiment is shown in Figure 2,
The specimen near the overflow was the first to
leave the starting dish and it moved towards
the current. Of a total of 25 animals 66 percent
moved towards the current, 34 percent showed
no detectable response.
I(b) The results of food selection experiments
indicate that, in the laboratory, Rostanga pulchra
of chemical sense. The results of six experi-
ments are summarized in Figure 3. Of the 72
animals used, 8 were floating in the surface
film at the end of the experiments and are not
included. Of the remaining 64 animals, 76.7
percent showed a definite movement towards
the current from O, pennata.
I(c) When the current was strong on One side of
the apparatus and the red sponge was in the
other current, the animals at first oriented into
the current. At the end of the experiment, 83.3
percent of the 12 animals used had moved to-
wards the current from the sponge (Figure 4).
Page 196
Ophlitaspongia
pennata
and slight
current
strong
current
Figure 4
Distribution of Rostanga pulchra given a choice between
current and OpAlitaspongia pennata
II.
sponse to current nor to the current from Hali-
chondria panicea.
Archidoris montereyensis showed no re-
Ill. Of the sponges tested Rostanga pulchra
would eat only Ophlitaspongia pennata and Ar-
chidaris Mienre Re yeMmsis would eat only Hali-
Discussion
The results of these experiments indicate
that Rostanga pulchra is able to find Ophlita-
spongia pennata by chemotaxis. This chemo-
taxis probably helps the animal find its food in
nature.
Although only Ophlitaspongia pennata was
used in these experiments, there are other red
sponges in the area. None were collected dur-
ing this study. Occasional reports are found of
Rostanga pulchra on red sponges other than O.
pennata; however, they do not indicate whether
or not the nudibranch was actually feeding on
the sponge. Doran (1951) found R. pulchra on
O, pennata and Esperiopsis originalis. Bakus
(personal communication) finds R, pulchra
mostly on O, pennata but has found it twice on
Polycamia karykina. De Laubenfels (1927) re-
ports that R. pulchra eats O, pennata, Acarnus
erithacus, P. karykina, and ‘Isociona lithophoe-
Whatever the relationship between R, pul-
it seems to have
nix,
chra and these other sponges,
a definite association with O. pennata.
feed on red sponges. In eae Britain C. M.
THE VELIGER
Vol. 4; No. 4
Yonge (1949) reports that R. rufescens feeds on
Ophlitaspongia. Flattely and Watton (1922), also
in Great Britain, state that R. coccinea lives on
Microciona altrasanguinea.
Archidoris montereyensis fed only on Hali-
chondria panicea in the laboratory but it “does
not seem to have a definite association with the
It is not as consistently found with the
sponge.
sponge in nature as is Rostanga. Doran (1951)
dissected specimens of A. montereyensis and
He found cells of Ma-
crocystis sp, and sponge spicules. Other spe-
cies of Archidoris have been reported to feed on
sponges. Forrest (1952) states that A. stelli-
fera has a preference for Stylotella columella.
Miller (1961) found A. pseudoargus feeding on
H. panicea and Tethya aurantia. There are oth-
er dorid nudibranchs which eat sponges. Aldisa
sanguinea (Cooper) is found on O. pennata
(Doran, 1951). Miller (1961) found Jorunna to-
mentosa on H. panicea. In the laboratory the
author observed Diaulula sandiegensis feeding
on Haliclona sp.
examined gut contents.
Summary
Rostanga pulchra has a positive rheotaxis.
Under experimental conditions it is able to find
Ophlitaspongia pennata by chemotaxis. It ap-
pears to be quite specific in its association with
this sponge.
Archidoris montereyensis showed no rheo-
taxis or chemotaxis. Although it fed only on
Halichondria panicea in the laboratory, it seems
clear that it does feed on other things.
Literature Cited
de Laubenfels, M. W.
1927. The red sponges of Monterey Peninsula, Cali-
fornia. Ann. Mag. Nat. Hist. (ser. 9)19: 258 to
266.
Doran, D, J.
1951. Observations on the distribution and feeding
habits of nudibranchs, Zool, 112. Stud, reports
Dept. Zool,, Univ, California, Berkeley.
Flattely, F. W., & C, L. Watton
1922. The biology of the sea-shore, MacMillan Co.,
New York,
Forrest, J, E.
1953, On the feeding habits and the morphology and
mode of functioning of thealimentary canalin some
littoral nudibranchiate mollusca. Proc. Linn, Soc,
London 164: 225-235.
Miller, M. C.
1961. Distribution and food of the nudibranchiate mol-
lusca of the south of the Isle of Man. Journ, Ani-
mal Ecol. 30; 95-116.
Yonge, C. M.
1949. The sea shore, Collins, London.
Vol. 4; No. 4
THE VELIGER
Page 197
Relationship of Living Weight to Shell Cavity Volume
in Helix aspersa
BY
ANDREA HERZBERG & FRED HERZBERG
5818 Jumilla Avenue, Woodland Hills, California
(Plate 45; 2 Textfigures)
It has been suggested that the shell of the
terrestrial snail Helix aspersa may be consid-
ered a probable biologic kymograph. Increases
in the volume of the shell over a given time pe-
riod can be determined and provide a quantita-
tive measure of the growth of the animal during
that period. The effect on growth of various
environmental factors such as cold (Herzberg
and Herzberg, 1960) can thus be easily studied.
The technique by which the shell cavity volume
is determined is a simple one, but it is time-
consuming. The weight of the animal, however,
is easily and quickly obtained. It has been shown
that in H. pomatia and Zebrina detrita the cube
of the shell width, the height of the shell, and
the living weight of the animal are proportional
to each other (Kienle, 1957), and it is known that
in Concholepas conchopas the volume of the
shell is directly proportional to the weight of
the animal (Schwabe, 1959). The purpose of this
study is to discover whether such a proportion-
ality exists in H. aspersa and, if so, whether it
is exact enough so that the weight of the living
animal can be utilized to obtain the volume of
the shell cavity or, in other words, whether the
specific gravity of H. aspersa is constant.
Materials & Methods
One hundred and three specimens of Helix
aspersa were collected from the wet soil of a
garden in Woodland Hills, California, during the
early evening hours of July 5, 1961, and July 12,
1961, Each animal was placed overnight in an
individual compartment, measuring 3 x 3 x 3
cm., in an almost airtight plastic box and per-
mitted to seal off against one wall of its com-
partment, The animals were then weighed and
sacrificed by immersion in boiling water. Their
shell cavity volumes were then determined, us-
ing a technique described earlier (Herzberg and
Herzberg, 1960) and illustrated in Plate 45.
Results
The weight and the shell cavity volume
were found to be roughly proportional, as shown
in Textfig. 1, with an average specific gravity of
1.12. However, the specific gravities fall into
a random distribution ranging from 0.81 to 1.42,
as shown in Textfig. 2. The range of specific
gravities bears no relationship to the size of
the animal, so that two animals of either simi-
lar weight or similar shell cavity volume may
have widely differing specific gravities.
Figure 1: Graph showing relationship of live animal
weight to shell cavity volume.
1 2 3 4 5 6 7 8
grams
Discussion
It is not possible to multiply the weight of a
specimen of Helix aspersa by a constant factor
to determine the shell cavity volume, since the
specific gravity of the animal is not a constant,
Page 198
THE VELIGER
Number of
10) Snails
9
8
0.99 1.00 PIO ile 1.40
1.30
Specific Gravity
Figure 2: Graph showing random distribution of
specific gravities of Helix aspersa.
It is interesting to note that the range of specif-
ic gravities is great, including specific gravi-
ties of less than one. The extent of the range
may be due in part to the variations in hydration
which have previously been noted to cause large
variations in weight in individual shells within
short time periods (Howes and Wells, 1934). Ail
snails which we have observed to estivate, hi-
bernate, or otherwise seal off, withdraw far into
their shells. There is, therefore, a potentially
large difference between shell cavity volume
and soft tissue volume. In those snails which
have an especially low specific gravity, it may
be that this difference has been exaggerated.
For example, a snail which has sealed itself off,
slowly but progressively loses water and de-
creases in body weight (Howes and Wells, 1934)
and therefore in specific gravity. The random
collection of snails in this study may well have
included some snails which had recently under-
gone such changes as well as some snails which
were quite well hydrated and thus had very high
specific gravities. Moreover, even snails kept
under constant conditions may show large fluc-
tuations in weight during short time periods
Vol. 4; No. 4
(Wells, 1944). Such variations may be due in
part to changes in hydration, to previous depo-
sition of large numbers of eggs, as well as to
Snails showing a
specific gravity of less than one may be ac-
other, unknown, variables.
counted for by the presence of air in the shell
cavity, the amount of air varying with the extent
of withdrawal of the soft tissues into the shell.
Likewise, the amount of air present in the lungs
may be a factor in animals with low specific
gravities.
Summary
The specific gravity of Helix aspersa is not
constant but was found, in the 103 animals ex-
amined, to vary between 0.81 and 1.42 in a ran-
dom distribution, with an average specific
gravity of 1.12. It is not possible to multiply the
living weight of a snail of this species by a con-
stant factor to determine the shell cavity vol-
ume, and other methods must be employed to
ascertain shell cavity volume when size in-
creases in H. aspersa shell are used as an ex-
perimental criterion. Possible reasons for the
wide range of specific gravities are discussed.
Literature Cited
Herzberg, Fred, & Andrea Herzberg
1960. Theeffect of coldon the growth of Helix aspersa.
Journ. Exp. Zool., 145: 191-196.
Howes, N. H., & G. P. Wells
1934, The water relations of snails and slugs. Journ,
Exp. Biol., 11]: 327.
Kienle, M.
1957, Uber Beziehungen zwischen Kérpergrésse, Lun-
gengrésseund Energiekonsum bei Pulmonaten, ins-
besondere Helix pomatia L, und Zebrina detrita
Mull. Zeitschr. Vergl. Physiol., 40: 440-450,
Schwabe, G. H.
1959. Biometrische Daten zur Schale von Concholepas
choncopas (Bruguiére) (Moll., Muridae) an der chi-
lenischen Kiste und ihr é6kologischer Indicatorwert.
Internat. Rev. Ges. Hydrobiol., 44: 449-462.
Wells, G. P,
1944, The water relations of snails and slugs, III.
Factors determining activity in Helix pomatia L.
Journ. Exp. Biol., 20: 79-87,
Explanation of Plate 45
‘Yechnique for preparing shells to determine shell cavity volume.
Figure 1: Removing soft tissues from shell by forceps rotated in a counter-clockwise direction to avoid shell break-
age. Figure 2: Wax poured into washed and dried shell held with apex down. Wax used of specific gravity of 0.9.
Figure 3: Shell filled with wax at second pouring, showing shrinkage of wax after cooling. Figure 4: Final appli-
cation of wax with hot metal spatula used to remove excess wax to rim of aperture.
[Herzperc & Herzperc] Plate 45
THE VELIGER, Vol. 4, No. 4
‘ «
KO KY
KS CW
SIRS KW
KK ‘
KS <
SY
\
HERZBERG & HERZBERG, photo.
Vol. 4; No. 4
THE VELIGER
Page 199
Type Localities
BY
F. A. SCHILDER
University of Halle (Saale), German Democratic Republic
The type locality is a geographical term
designating the area in which the holotype of a
named lower taxon (species, subspecies = race,
morphe, variety, etc.) lived.
The International Rules of Zoological No-
menclature do not concern themselves with this
important, sometimes even decisive, addition
to the description of a newly established taxon,
though there are several analogies between the
rules concerning the names, descriptions, and
types of animals and the practice concerning
their type localities as compiled in this paper.
An author establishing a new lower taxon
by publication of its name and description should
also indicate, as exactly as possible, the local-
ity from which the holotype of the taxon is said
to come.
This original designation of the type local-
ity must be adopted by all later writers (see
Note 1), except if it can be proven that the indi-
cation of habitat was evidently erroneous, by the
following reasons:
1, The holotype really came from another lo-
cality than originally indicated: then, the previ-
ous type locality must be rejected (Note 2), even
if the taxon may later on be proved to occur
also in the locality erroneously named.
2. The exact description or figure of the holo-
type proves that it indeed belongs to the species
recognized by later writers but evidently toa
geographical race or variety restricted to an
area which does not include the indicated local-
ity (Note 3).
3. The species does not occur at all in the in-
dicated geographical area, as can be proved by
the fact that no living specimens have been col-
lected later even in adjacent areas, and their
having ever lived there is most improbable be-
cause of zoogeographical reasons (Note 4).
If there is no original designation of type
locality at all, or if the indication of habitat is
evidently too vague, Or even quite wrong, the
mistake should be corrected by a subsequent
designation as follows:
1, The revising author may establish a new
type locality, if the original description of a
lower taxon does not contain any indication at
all(Note 5), or if the original designation is ev-
idently erroneous (Note 6).
2. The revising author may select a type lo-
cality, if two or more habitats have been named
originally (Note 7).
3. The revising author may restrict an origi-
nal type locality, if the geographical term used
as habitat of the holotype is too vague with re-
gard to the whole geographical range of the
taxon (Note 8).
ad 1. The first correct subsequent desig-
nation of a new type locality is to be adopted by
later writers in the same way as a correct ori-
ginal designation, provided that the locality
chosen is the habitat of a real specimen of the
species (Note 9) agreeing with the original de-
scription and figures in all characters distin-
guishing allied species, races, varieties, stages,
etc. (Note 10); otherwise, the subsequent desig -
nation should be rejected and replaced by a
second subsequent designation with the same
prospect to become final, and so on (Note 11).
ad 2. In selecting a type locality from sev-
eral simultaneous correct indications of habitat,
no word priority should be decisive, but rather
the probability that the original author has ex-
amined real specimens from one locality only
(which should be selected) and added the remain-
ing ones from papers consulted, or personal
communications received (Note 12); if no local-
ity can be preferred by such reasons, that indi-
cated habitat should be selected as type locality
which is situated most centrally in the whole
area of distribution of the taxon (Note 13), The
selected type locality is to be adopted by later
writers, even if these recommendations have
not been followed.
Page 200
THE VELIGER
Vol. 4; No. 4
ad 3. In restricting a type locality, the new
type locality must surely lie within the limits of
the original one (Note 14), best-in the centre of
the area inhabited by the taxon. The restriction
of a type area by the original author of the taxon
himself ina later paper, in which the holotype
should be regarded as
even if another writer has re-
is indicated exactly,
valid, I think,
stricted it in another way in the meantime (Note
15).
ities in a too particular way, but such hyper-
It is not advisable to restrict type local-
exact indications are not allowed to be general-
ized later on (Note 16).
If a preoccupied name becomes replaced by
a name new to science without indication of type
locality, the type locality of the invalid name
should be adopted (Note 17), even if the second
author has quoted figures of specimens coming
from other localities merely to explain the
meaning of the taxon by additional illustrations;
if, however, the second author expressly estab-
lished another type locality for his new name,
this locality should be considered valid.
There is some difference between the mere
mentioning of the locality where the just estab-
lished taxon has been collected, and the explicit
But as the
latter term is rather modern in taxonomy and
formal calling of it "type locality".
unknown to ancient writers, no first correct or
restricting indication of habitat should be re-
jected by the only reason that it lacks the term
"type locality" (Note 18).
These proposed general rules may be illus-
trated by some examples in cowries (Cypraei-
dae):
(1) Linnaeus, 1758, has indicated Mauritius
as habitat of Erosaria caputserpentis (Linnaeus,
1758), adopting the locality named by Lister,
1688; therefore, Iredale, 1935, and Steadman &
Cotton, 1946, were right in adopting Mauritius
as type locality expressly; the habitat Tahiti of
Sowerby, 1837, however, must be rejected as
type locality, as it has been indicated later than
Mauritius of Linnaeus, 1758. —— But Iredale (1935)
was wrong in rejecting "in Maldivis"' indicated
as habitat of Palmadusta asellus by Linnaeus,
1758, in the same way and in establishing Am-
boina as type locality on account of Linnaeus'
quoting Rumphius'
(1705) in his synonymy list, the more so as
Amboinsche Rariteitkamer
many shells described by Rumphius do not live
in Amboina at all. Maldive Island
must be restored.
Therefore,
(2) Schilder, 1927, indicated Palawan as
misreading the original label ''Palabuan"; this
error has been corrected by Schilder, 1929, the
more so as E, vredenburgi lives on the south-
west coast of Java only. Steadman & Cotton,
1946, have erroneously indicated Nusa Kam-
bangan as type locality (which is about 300 kilo-
meters east of Palabuan), only for word priority
in our ''Prodrome" of 1938.
(3) Lamarck, 1810, described Erronea
adusta from this term
usually means Indonesia where the bluish white
typical E, onyx (Linnaeus, 1758) lives; but as
Lamarck clearly described the chestnut race
living from East Africa to Tenasserim only,
Steadman & Cotton, 1946, were right to replace
the type locality by Zanzibar.
"Oceanus Asiaticus'"!:
(4) Gaskoin, 1849, described Cribraria cri-
bellum from the Mediterranean, but it really
occurs in the Mascarene Islands only; there-
fore, Weinkauff, 1881, corrected the habitat into
Mauritius which has been adopted by Steadman
& Cotton, 1946, as type locality.
(5) Linnaeus, 1758, Gmelin, 1791, and La-
marck, 1810, described Erosaria ocellata (Lin-
naeus, 1758) without indicating any habitat; Gray,
1825, mentioned the "Indian Ocean"! (from Mar-
tini, 1769) and China (from Humphreys, 1797),
but E. ocellata does not live in the latter coun-
try; Sowerby, 1837, figured E. ocellata from
Ceylon, and as it is rather frequent in this is-
land, Steadman & Cotton, 1946, fixed Ceylon as
type locality also formally.
(6)
ventriculus from New Holland;
Lamarck, 1810, described Cypraea
as it does not
occur in Australian waters at all, and as it has
been mentioned from Annaa Island by Sowerby,
1837, first, Steadman & Cotton, 1946, were right
to list Annaa as type locality.
(7) When describing Luria isabella, Lin-
naeus, 1758, named two islands as habitat:
Mauritius and Madagascar; as Reeve, 1845, re-
peated Mauritius only, Ladd, 1945, was right to
select Mauritius as type locality; however, the
type locality ''Amboina" of Iredale, 1935, must
be rejected, as it is not included formally in
Linnaeus! original description of the species.
(8) Gray, 1828, described Naria irrorata
from the ''South Sea'! generally, but Sowerby,
1837, indicated Elizabeth Island as habitat; this
island should be regarded as type locality
though Iredale, 1935, and Steadman & Cotton,
1946, retained "Pacific Is."" and ''Polynesia’
respectively, an area which covers almost a
greater part of the globe than the range of N.
irrorata in its entirety.
Vol. 4; No. 4
THE VELIGER
Page 201
ee eee ne NT aI ENP NR OU Ltn eNO Ra
(9) Linnaeus, 1758, described Erosaria
poraria without habitat; Lamarck, 1810, quoted
Senegal; and Gray, 1825, added Jamaica (from
Martini, 1769); but both habitats are wrong as E,
poraria is restricted to the Indo-Pacific Ocean;
Sowerby, 1837, was the first to name pos-
sible localities, Ceylon and Pacific Ocean.
Nevertheless, Iredale, 1935, established Ambo-
ina as type locality, but it must be rejected as
E. poraria never has been collected at Amboina
nor in adjacent islands: it spreads from the
Eastern Pacific Islands (race E. p. scarabaeus
Bory, 1827) and from the western Indian Ocean
(E. poraria s. str.) to the outer borders of Indo-
nesia only, and it does not occur between the
southern coast of Java and North West New
Guinea (Skroe). Therefore, Iredale's designa-
tion of the type locality of E. poraria must be
replaced by Ceylon, as Schilder & Schilder, 1938,
restricted E. poraria to the Indian race.
(10) Gray, 1825, described Palmadusta
humphreysii without habitat; Iredale, 1939, des-
ignated Amboina as type locality, though
Schilder & Schilder, 1938, had shown that this
clearly separable race of P, lutea Gronow, 1781,
is restricted to the area between Torres Strait,
Tonga, and Sydney, whereas P, lutea s. str. oc-
curs from Malaysia to Japan; therefore, if spe-
cimens of P, lutea (s. lat.) should be found in
Amboina in future (they have not yet been found
in the southern Moluccas at all!), they undoubt-
edly would belong to P, lutea and not to PB, hum-
phreysii. Therefore, I designate Lifu as type
locality, from which Melvill & Standen, 1895,
received P, "lutea var. humphreysii" and from
which I possess a specimen myself (ex coll.
Hervier).
(11) Reeve, 1835, described Erronea sub-
viridis without habitat; in 1845 he described and
figured a shell from Dupuch's Island as E. sub-
viridis, which does not agree with the original
description but represents the West Australian
race E, s. dorsalis Schilder & Schilder, 1938;
the more eastern typical E. subviridis has been
first figured by Sowerby, 1870, from New Cale-
donia; therefore, New Caledonia should be
treated as type locality of E. subviridis (instead
of North Queensland designated as type locality
by Iredale, 1935), and Dupuch's Island as that of
E.
s. dorsalis.
(12) Iredale, 1939, was right in restricting
the type locality of Blasicrura kieneri schneid-
eri Schilder & Schilder, 1938 (originally de-
scribed from Melanesia and East Australia) to
New Britain, supposing that we had received
specimens from P, J. Schneider who collected
there; later on, Schilder, 1958, named Ulamona
in New Britain as habitat of the holotype.
(13) Reeve, 1845, established Palmadusta
diluculum with the erroneous habitat Philippine
Islands; according to Schilder & Schilder, 1938,
it spreads from ''Natal to Zanzibar’'. Steadman
& Cotton, 1946, selected Natal as type locality
evidently by word priority, though P, diluculum
is more common farther north than on its sou-
thernmost border; the selected
type locality Natal must be retained.
nevertheless,
(14) Sowerby, 1832, named no habitat of
Cribraria cumingii, but Gray, 1833, added'’Raie
tea", which was designated as type locality
(""Raietea"’) by Iredale, 1935; Steadman & Cotton,
1946, however, incorrectly quoted Tahiti as type
locality which is a better known island in the
Society Islands, but which belongs to another
group of islands more than 200 kilometers off,
(15) Palmadusta punctata iredalei Schilder
& Schilder, 1938, was originally established with
the range from ''S, Melanesia to Manokwari,
Queensland, Tonga, and Samoa’’; the figure of a
shell from Lindeman Island (Queensland) pub-
lished by Iredale, 1935, was quoted among the
illustrations of the new race. Steadman & Cot-
ton, 1946, established Lindeman Island as type
locality, but Schilder, 1958, designated a shell
from Mope (New Britain) as holotype, which was
examined personally in 1938, whereas the iden-
tity of the Lindeman shell should be regarded
as not demonstrable. Therefore, Mope should
be retained as type locality, I think.
(16) The holotype of || Erronea sophiae
(Brazier, 1876) has been described from Makeira
Harbour at San Christoval, Solomon Islands;
Iredale, 1935, and Steadman & Cotton, 1946,
were not justified in generalizing the exact ori-
ginal indication into the ''type localities'’ San
Christoval Islands and even Solomon Islands,
respectively.
(17) Schilder, 1932, rechristened the invalid
name Mauritia reticulata (Martyn, 1784) into M.
maculifera, without naming a type locality; but
as Martyn's shell is said to come from the
Friendly Islands, this habitat (=Tonga Island)
should be retained for M, maculifera too,
(18) Gray, 1824, described Erronea pyri-
formis without habitat, but in 1828 he added New
Holland, whereas Sowerby, 1837, and Reeve,
1845, both quoted Ceylon, Iredale, 1935, accept-
ed Ceylon because it 'is more likely'' and ex-
pressly designated Ceylon as type locality in
1939; but E. pyriformis lives also in Queensland
Page 202
THE VELIGER |
Vol. 4; No. 4
(where it seems to be more frequent than in In-
dia: Sowerby, 1870, Iredale, 1939, Schilders'
collection); therefore, the original habitat (New
Holland) must be restored, but it should be re-
stricted to "Queensland", as Gray's description
fits to the East Australian E. pyriformis (s.
str.) but not to the West Australian race E, p.
smithi Sowerby, 1881.
A revised list of the type localities of all
living Cypraeidae will be published in another
paper.
A Preliminary Report on Spawning and Related Phenomena
in California Chitons
BY
SPENCER R. THORPE, Jr.
116 Ramona Avenue, El Cerrito, California
(3 Textfigures)
The California Coast has one of the richest
associations of chiton species found in the
world, the total number of species occurring
here being exceeded only by that of the Austral-
ian chiton fauna. In the diversity of the groups
making up the fauna and the large number of
unique species present, the California chiton
fauna perhaps surpasses that of any other area.
Although they present a rich field for investiga-
tion, our California chitons have been the sub-
jects of relatively little biological research. In
the area of breeding habits, almost all our
knowledge comes from four papers by Heath
(1899, 1905, 1907, and 1912).
The present paper reports observations [|
have made during the period from 1956 to mid-
1961, Fourteen species of chitons have been
observed to release gametes under laboratory
As far as I can determine, there
have been no previous reports in the literature
concerning the breeding activities of 11 of the
14 species, The larvae of six species have de-
veloped for limited periods of time in the labo-
ratory, but I have been unable to carry the
larvae of any species through to maturity. Evi-
dence is also presented indicating that for two
conditions.
species the time of gamete release is correlat-
ed to the tidal cycle.
Method
My procedure for collecting and handling
The animals are pried
off rocks with a dull paring knife and placed in
collecting jars filled with sea water. The jars
are kept closed and placed ina closed collect-
ing bag while I am in the field and when return-
ing from the field. Unless injured, the chitons
will almost invariably uncurl and attach them-
selves to the sides of the collecting jars. Nei-
ther individuals nor species are ordinarily
segregated following collecting.
chitons is very simple.
The sea water in the collecting jars is
changed just before leaving the field. In the
laboratory the chitons may be left in the col-
lecting jars with the lids off or may be manual-
ly transferred into flat, open pyrex dishes. In
either case fresh sea water is generally used.
Gamete release has occurred while the an-
imals were in the closed collecting jars and
while being transported between the field and
laboratory; in open collecting jars in the labo-
ratory before and after the sea water has been
changed; and in open pyrex dishes. Gamete re-
lease has occurred in the laboratory when the
chitons were in total darkness, and also when
Vol. 4; No. 4
THE VELIGER
Page 203
a ca Sse he eS
they were exposed to rather bright electric
lights. There is no reason to believe that one,
or a combination, of these laboratory variables
induces gamete release,
Whole mount preparations were made for
studying the larval development. Bouin's solu-
tion and 5 percent formaldehyde in sea water
The formaldehyde so-
lution gives good preservation of the calcareous
It also
partially dissolves the egg cases in a matter of
a few days.
were used as fixatives.
structures in larvae over a week old.
Grenacher's borax carmine and Heiden-
hain's iron hematoxylin stains were used. I ob-
tained the best results with the carmine stain,
particularly in the early larval stages where
the larvae are still surrounded by their egg
cases. The cytoplasm of the larval cells is
quite receptive to the hematoxylin stain and
does not destain easily.
Data
Tables 1 and 2 summarize my observations
of gamete release. The date following the lo-
cality is that on which the specimens were col-
lected. The specimens were collected not more
than one and one-half hours before nor more
than two hours following the low low tide. The
"Next High Tide’ is the low high tide following
the low low tide. The ''Next Low Tide'' is the
high low tide following the low high tide. All
times after 2400 hours are on the date following
collecting of the specimens in the field. Some
of the localities shown in the tables are not
listed in the tide tables. In such instances an
estimate of the high and low tides has been
made by extrapolation. The estimates are be-
lieved to be accurate to within 15 minutes in all
cases.
1 The water was over-cooled, and the speci-
mens behaved peculiarly.
2 The specimen was collected ina tide pool
high in the intertidal zone, an unusual habi-
tat for this species.
3 Observations were made by Mr. Daryl
Sweeney.
4 The specimens released gametes while
isolated in individual jars.
4a The specimen was the only one of its spe-
cies collected on that date.
4b All specimens collected on this date were
females.
5 The start of gamete release was not ob-
served.
6 The specimen was killed before gamete re-
lease had ceased,
7 The eggs or the sperm were seen, but gam-
ete release was not observed.
8 The water became so cloudy that it was
very difficult to determine when any one in-
dividual began or ceased to release gam-
etes.
Egg laying occurred between 0400 and 0800,
October 29, 1957.
Egy laying occurred between 0040 and 0830,
Hebruany 265) lS.
9c Egg laying occurred between 0215 and 0730,
February 15, 1957.
fertilized although no cloudiness from
9b
Some of the eggs were
sperm discharge was detectable in the wa-
ter.
9d Gamete release occurred between 1730 and
1915, December 26, 1956.
9e Gamete releasc occurred between 1400 and
NZOO, Ajspeiil MC). WET
10 The five males started to release sperm at
various times between 2130 and 2330. The
exact time at which any individual started
was impossible to determine due to the
cloudiness ot the water.
11 More than one male may have released
sperm.
12 The specimen was still releasing gametes
at 0130, October 15, 1961.
13. Gamete release occurred between 0130 and
0930, October 15, 1961.
Observations
The bulk of my Observations has been
made on various species of the genus Mopalia,
and the following descriptions are based pri-
marily upon observations of this genus. Except
where specifically noted, the descriptions apply
equally well to males of the genera Ischnochi-
ton, Chaetopleura, and Placiphorella, and to the
larvae of Ischnochiton (Lepidozona) californien-
sis (Berry, 1931) through about the third day of
development.
Gamete Release
Female chitons appear to be loosely at-
tached to the substrate during the first part of
spawning. They are generally quiescent during
this period but quite frequently will move sev-
eral inches during the later part of spawning.
The behavior of the males is less predictable.
Most frequently they will remain stationary the
entire time they are releasing sperm, but on
Page 204 THE VELIGER Vol. 4; No. 4
Table 1: Mopaliidae
Gamete Shedding
Time Time
Location Date 2 oe Start Finish
Mopalia ciliata (SOWERBY, 1840)
South Side, Pigeon Point’
30 Nov. 1956 | 1
0250
South Side, Pigeon Pott’ 30 Nov. 1956 I 2320 0250
North Side, San Pedro Point’ | 25 Feb. 1957 I 2000 0235
North Side, San Pedro Point® | 25 Feb. 1957 | 1 2100 0235
North Side, San Pedro Pome | 25 Feb. 1957 I 2130 0235
North Side, San Pedro Point | 27 Feb. 1957 | 2 2200 0400
Pescadero Point’ 21 Mar. 1957 I 1815 2030
Sausalito’ 25 Sepy O57 I 0230 0726
Sausalito’ 22 Oct. 1957 I 2100 0430
Sausalito’ 2 (Ole, WOR | 1 2245 0430
North Side, San Pedro Point’ | 25 Oct. 1957 | 1 2400 0615
Mission Poimt 21 Feb. 1960 | 1 2100? 2400
Franklin Pom 22 Feb. 1960 I 2130 0110
Franklin Point’ 27 Feb. 1960 I 2330 0530
‘Tiburon’ 20 Nov. 1960 | 1 2130? 0645
‘Tomales Bay’ 11 Mar. 1961 I 1900? 0130 5
‘Tomales Bay’ 11 Mar. 1961 I 1900? 0130 5, 6
‘Tomales Bay’ 11 Mar. 1961 I 2030 0130 6
Aquatic Park, San Francisco” | 15 July 1961 I 1530 2020
Mopalia, spec. nov.
Marina, San Francisco” 30 Apr. 1961 1130? 1400 1300 1815 5
Marina, San Francisco’ 13 May 1961 1530 1700 1215 1735
Marina, San Francisco” 13 May 10961 1800 IQI5 1215 1735
Marina, San Francisco’ 13 May 1061 1800 1915 1215 1735
Marina, San Francisco” 13 May 10961 1900 2010 1215 1735
Marina, San Francisco* 14 May 1961 1200 1215 1310 1815
Marina, San Francisco” 14 May 1961 1900 2010 1310 1815
Aquatic Park, San Francisco’ | 15 July 1961 1930 2200 1530 2020
Aquatic Park, San Francisco” | 15 July 1961 1930 2200 1530 2020
Aquatic Park, San Francisco’ | 15 July 1961 2230 2300 1530 2020
Aquatic Park, San Francisco’ | 15 July 1961 2200 2300 1530 2020
Mopalia lowe: (Pitssry, 1918)
‘Tomales Bay’ 11 Mar. 1961
‘Tomales Bay’ 3 June 1961
Mopalia porifera (PitssBry, 1892)
Pescadero Point’ 10 Nov. 1958} 2 2130 2200 2230 0330
Pescadero Point’ 10 Nov. 1958 2 2145 2300? 2230 e338 8
Pescadero Point’ 10 Nov. 1958 3? 2200? 2400 2230 0330 8
Pescadero Point’ 5 Mar. 1959| 1 2425 2450 2050 0205
Pescadero Point’ 5 Mar. 1959| 1 0125 0210 2050 0205
Pescadero Point’ 5 Mar. 1959 I 2345 2415 2050 0205
Pescadero Point’ 14 Oct. 1961 I 2400 2425 0315 0740
Pescadero Point® 14 Oct. 1961 I 2430 0100 0315 0740
Pescadero Point’ 14 Oct. 1961 I 2445 O105 0315 0740
Pescadero Point’ 14 Oct. 1961 I O105 ? 0315 0740 13
Pescadero Point’ 14 Oct. 1961 I ? ? 0315 0740 7 14
Franklin Pome 12 Mar. 1960} 2 2230? 2320 2235 0420 5
Marin County *= San Francisco County © San Mateo County
Monterey County * Los Angeles County
Vol. 4; No. 4 THE VELIGER Page 205
Table 1: Mopaliidae (continued)
Gamcete Pi Tide
Time Time Next Next
Location Date Start Finish High Low Note
Mopalia hindsi (REEVE, 1847)
San Pedro Point’ 25 Oct. 1957 2345 0130 0615 3514510
Moss Beach’ 28 Oct. 1957 ga 0315 0730 3> 4, 7
Mopalia imporcata (CARPENTER in Pitssry, 1892)
Franklin Point’ 26 Feb. 1961 2120 0220 4b
Franklin Point’ 26 Feb. 1961 2120 0220 4b, 6
Franklin Point’ 26 Feb. 1961 2120 0220 4b, 6
Mopalia muscosa (GouLb, 1846)
Aquatic Park, San Francisco" | 15 July 1961 1500? 1730 1530. 2020
Aquatic Park, San Francisco” | 15 July 1961 I 1530? 1730 1530 2020
Mopalia lignosa (GouLD, 1846)
North Side, San Pedro Point’ | 27 Feb. 5 I 2300 2400 0350
North Side, San Pedro Pot’ |27 Feb. I 2400 2440? 0350
North Side, San Pedro Point" }27 Feb. ; I gb gb 0350
Muir Beach’ 12 Mar. r 2000 2100 0220
Bolinas Point’ 13 Mar. I 2000 2110 0320
Placiphorella velata (CARPENTER in Pitssry, 1892)
| Mission Point | 28 Sep. 1959 | | I | 2100? il 2240
Table 2: Other Chiton Species
Gamete Shedding Tide
Time Time Next Next
Location Date Chel anee Start Finish High Low Note
Ischnochiton radians (CARPENTER in Pitssry, 1892)
North Side, San Pedro Point’ | 14 Feb. 1957] 1? gc gc 2325 0455 7
North Side, San Pedro Point’ | 14 Feb. 1957 ? gc gc 2325 0455 Ts 10|
Ischnochiton mertensi (M1pDENDOREF, 1846)
|Ano Nuevo Bay’ | nG lao, CRW [ 2200? | 2315 | 2215 | 0330 | 5 |
Ischnochiton regularis (CARPENTER in PiLsBry, 1892)
| Mission Point’ 21 Feb. 1960 | 7 5 | 2130 | 2400 a 1915 | 2345 [ 8, 11 |
Ischnochiton californiensis BERRY, 1931
North Side, Resort Point 26 Dec. 1956] 1 gd 1800 2235 7
North Side, Resort Point’ 26 Dec. 1956 Hf : gd 1800 2235 7, 12
White's Point® 30 Dec. 1956 Die 21300 2230 2130 0200 5, 8
Chaetopleura gemma (CARPENTER in Pitspry, 1892)
‘Tomales Bay' | 3 June 1961 ine De le 1530? ail 1615 1740 | 2240 5s 8 |
Tonicella lineata (Woop, 1815)
[North Side, San Pedro Point’ | 19 Apr. 1957 ee I ge |
ge | 1600 a 2030 ia 7 |
‘= Marin County *— San Francisco County
*— Monterey County
*— San Mateo County
*— Los Angeles County
Page 206
other occasions I have seen them move inter-
mittently and, more rarely, move continuously
throughout the entire time they are releasing
sperm,
The ejection of eggs and sperm from the
body cavity seems to be aided by rhythmic
movements of the foot. This may explain why
the females seem loosely attached to the sub-
strate at the beginning of spawning.
The
eggs are first ejected into the mantle cavity
Egg laying is a two-stage process,
where several hundred may accumulate on each
side of the foot.
orly, presumably by ciliary currents, and
emerge through an upraised portion of the hind-
most part of the girdle. Frequently, the eggs
streams'', one from each side
The eggs are carried posteri-
emerge as two ''
of the mantle cavity. As they first emerge from
the mantle cavity, the eggs are propelled at
considerable speed but slow down almost im-
mediately and seldom travel more than two to
three centimeters. The eggs accumulate in a
pile behind the female.
Initially, the sperm is discharged in spurts
and has a coagulated appearance, as if enclosed
The sperm disperses very slowly at
As the male continues to release
in mucus,
this stage.
sperm, the discharge becomes more continuous
and the product no longer appears coagulated.
Dispersion of the sperm at this time is more
rapid. In all cases which | have observed, the
male extends the lateral edges of his foot to
cover the mantle cavity in the area of the geni-
tal opening. It has not been possible to deter-
mine whether the sperm accumulates in the
mantle cavity of male chitons as the eggs do in
the mantle cavities of the females.
I have gained the impression that once
gamete release has begun, rather drastic con-
ditions are required to cause the chiton to
cease, A very strong light which gives off con-
siderable heat will cause interruption of spawn-
ing, if the light is placed close to the chiton. On
the other hand, I have removed both males and
females, in the process of releasing gametes,
from collecting jars and transferred them into
dishes filled with sea water, where gamete re-
lease continued with little or no interruption.
On other occasions, female chitons have fallen
off the sides of a jar or dish in which they were
spawning, been turned over (as they had landed
foot uppermost), and resumed spawning without
interruption. One male Mopalia ciliata (Sower-
by, 1840) released sperm for over half an hour
while partially curled upon his back, Finally, in
an experiment, the water ina jar in which two
THE VELIGER
Vol. 4; No. 4
male Ischnochiton (Lepidozona) californiensis
were releasing sperm was violently agitated.
The water was emptied, replaced, and once
again violently agitated. The water was again
emptied from the jar but this time was not re-
When the
jar was refilled, the chitons were almost im-
mediately releasing sperm again, and 1 doubt if
any interruption of sperm release had occurred.
placed for a period of three minutes.
Sexual Products
The eggs of Mopalia appear to be shed in-
dividually without any visible trace of an albu-
minous sheath or envelope. However, the eggs
do not disperse in sea water as easily as one
would expect. When eggs are fixed in formalde-
hyde and sea water soon after they have been
shed, they seem to be enclosed in an amor-
phous, slightly translucent mass, Individual
eggs can be removed from this enclosing mass
only with difficulty. This mass enclosing the
eggs is most noticeable with unfertilized eggs
but gradually disappears or disperses 12 to 15
The exact nature of
this ''envelope'' is unknown,
hours after fertilization.
Ali chiton eggs I have seen are spherical in
shape. Each egg is enclosed in its own individ-
ual egg case. Numerous spine-like or plate-
like processes project from the surface of the
egg cases, giving the cases the appearance of
transparent, The
projections of the egg cases of Ischnochiton
radians (Carpenter in Pilsbry, 1893) are spine-
like. Those of the egg cases of I. mertensi
(Middendorff, 1846) are plate-like and may be
either wavy or curled. The projections of the
egg cases of the species of the genus Mopalia
are intermediate, between those of I. radians
and I. (L.) mertensi.
short-spined sea urchins.
The eggs of the species of Mopalia, Nuttal-
lina, Cyanoplax, Katharina, Cryptochiton, and
Placiphorella are a light grey-green. Those of
N. californica (Reeve, 1847) and M. lignosa
(Gould, 1846) are a brighter green. The eggs of
the species of the subgenera Lepidozona and
Stenoplax of the genus Ischnochiton are pinkish
or tawny-buff in color. Ischnochiton (S.) fallax
(Carpenter in Pilsbry, 1892) is an exception
among the Ischnochitons, the eggs of this spe-
cies being green in color.
The average diameters of the unfertilized
eggs of several species of chitons are listed in
Table 3. Measurements were made on eggs
preserved in formaldehyde and sea water and
do not include the egg case.
Vol. 4; No. 4 THE VELIGER Page 207
Table 3 closed ina ?visible mucus secretion. Accord-
; ing to Grave (1932), the eggs of Chaetopleura
Diameter apiculata are also enclosed in a mucus secre-
Species in microns| tion, and Christiansen (1954) describes a simi-
170
180
200
200
200
Mopalta porifera (Pitssry, 1892)
Mopalia imporcata (CARPENTER in Pitspry, 1892)
Mopalia ciliata (SOwERBY, 1840)
Mopalia spec. nov.
Mopalia lowe (Pitspry, 1918)
Mopalia lignosa (GouLp, 1846) 240
Ischnochiton mertensi (MiappbENDOREV, 1846) 200
Ischnochiton radians (CARPENTER in Pitssry, 1893) 180
Heath (1899) reports that the eggs of Isch-
nochiton (Stenoplax) magdalenensis (Hinds, 1844)
[=1. (Stenoradsia) heathiana Berry, 1946] aver-
It sitincl ae
curious that the eggs of various species of Mo-
age about 400 microns in diameter.
palia are so much smaller than those of I, (S.)
heathiana, when it is considered that the latter
species has a free-swimming larval stage that
lasts less than 5 percent as long as the free-
swimming stage of Mopalia larvae, One would
expect that the species with the longer free-
swimming stage would require more stored
food and therefore possess the larger eggs.
Larval Development
The development of Mopalia ciliata follows
quite closely that of Lepidopleurus asellus
(Spengler) as described by Christiansen (1954)
and that of Chaetopleura apiculata (Say) as de-
scribed by Grave (1932) in general pattern and
the timing of development, The larval develop-
ment of M. ciliata differs from that of Ischno-
chiton (Stenoradsia) heathiana as described by
Heath (1899) in having more rapid initial cleav-
age after the egg is fertilized, in emerging from
the egg case at a much earlier stage of devel-
opment, and in possessing a much longer free-
swimming larval stage. Table 4 summarizes
some of the principal events in the larval de-
velopment of M. ciliata in chronological order.
Discusston
All of the ten species considered in this
paper which have laid eggs can be described as
free spawning for, although as I have indicated
in the case of Mopalia ciliata, some mucus is
present, the eggs can be dispersed fairly easily
in sea water. This would also seem to be the
case for Ischnochiton (Lepidozona) cooperi
(Carpenter in Pilsbry, 1892) as described by
Heath (1905). Heath(1905) describes the eggs of
Katharina tunicata (Wood, 1815) as being en-
lar condition for the eggs of Lepidopleurus
asellus. The greatest development of a mucus
or albumen secretion surrounding the eggs oc-
cording to Heath (1899), the eggs are enclosed
in long albuminous sheaths, which are of suffi-
cient strength to hold the eggs together for sev-
eral days after spawning.
Both Nuttallina thomasi (Pilsbry, 1898) and
Trachydermon (= Cyanoplax) raymondi (Pilsbry,
1894) do not release their eggs into the sur-
rounding water but retain them in the mantle
cavity until after the larvae have gone through
metamorphosis (Heath, 1905). The latter spe-
cies is unique in being the only one thus far de-
scribed which is hermaphroditic (Heath, 1907).
I began to suspect rather early that there
might be some correlation between the time
that both sexes of Mopalia ciliata released
gametes and the tidal cycle. Figure 1 shows the
The ref-
erence point for this species is the low high tide
Figure 3 shows the
gamete release times for Mopalia spec. nov.
(see Thorpe, 1961), The reference point for ‘this
species is the next high low tide following col-
gamete release times for M. ciliata.
occurring after collection.
lection, Figure 2 represents the average spring
tide tidal cycle since the times between the
successive high and low tides do not remain
constant throughout a month or a year, and the
data cover a period of several years. The low
high tide for the day was selected as the refer-
ence point, which introduces a maximum error
of not more than 5 percent in the times shown
between the low low tide and the low high tide
and a maximum error of not more than 9 per-
cent in the times shown between the low high
tide and the high low tide.
times for each species are in the same order
The gamete release
and represent the same individuals shown in
Table 1, except that the last 15 individuals in
Figure | and the last four individuals in Figure
3 represent recent observations which were not
included in Table 1.
Although not perfect in detail, there seems
to be a reasonably good correlation between the
time that individuals of Mopalia ciliata release
gametes and the low high tide for the day. Mo-
palia spec. nov, follows a different pattern, The
females apparently spawn on a low tide, while
the males show a less predictable pattern. The
difference shown by the two species is of par-
ticular interest because they are believed to be
very closely related.
Page 208
THE VELIGER
Vol. 4; No. 4
Table 4:
The development of Mopalia ciliata (SOWERBY, 1840)
Time After
Fertilization
Fertilization,
Features of Development or Behavior
Typical for the Time Period Shown
Polar body minute and transparent.
First cleavage occurs.
3- 6 hours
Third and subsequent cleavages are somewhat more rapid in the mi-
cromeres, Macromeres tend to be slightly larger than micromeres.
hours
Gastrulation beginning.
and beating at about 12 hours.
Cilia of velum encircle larvae and beat in wave-like motions,
The first cilia of the velum are developed
Gas-
Development to
this stage temperature dependent, i. e., larvae developed at 12-15° C,
above normal ocean temperature emerge from egg cases in 12 to 24
hours; those developed at 3-5° C, above normal emerge at 24 hours.
Larvae developing at normal ocean temperature emerge in 36 to 42
Apical cilia develop just prior to emergence from the egg
Larvae are free swimming in aerated water; in non-aerated water
Photonegative.
swimming may be ina
loose spiral, with the larvae rotating rapidly, or in a straight line
The anlagen of the ocelli and the valves begin to develop towards the
end of the fourth day. The anterior valves are the first to be appar-
Larvae becoming elongated in the antero-posterior axis and
The predominant movement towards
All eight valves and CaCO 3 spi-
cules of the girdle present on the eighth day. End of free swimming
18-24 hours
trulation apparently complete at 24 hours.
24-48 hours Larvae emerge from egg cases during this period.
hours.
case.
2- 4 days
they remain at the bottom and move only slightly.
Larvae most active at the end of this period;
without rotation.
4- 5 days
ent.
flattened dorso-ventrally.
5- 8 days Larvae noticeably less active.
the end of this period is creeping.
stage.
8-16 days
tarded by unfavorable conditions.
"Metamorphosis" complete at 16 days where the larvae are not re-
The anus has developed. Ocelli
still present. Valves are still covered by epithelium. Velum and api-
cal cilia have been lost,
Anlagen of the radula develop during this
period. When examined under a microscope, the larvae attach to the
slide by posterior portion of the foot, which portion in stained pre-
parations has a glandular appearance.
Brewin (1942) has shown that Cryptocon-
chus porosus (Burrow, 1815) releases gametes
every 15 days for a period of 2 to 23 months,
and spawning is apparently correlated with the
phases of the moon. Gamete release by C. por-
osus occurred regularly during the middle of
the day and was not correlated with the tidal cy-
cle. Brewin was also able to show that the re-
sults obtained in the laboratory coincided with
the dates and times of gamete release by the
animals in their natural habitat.
Christiansen (1954) and Heath (1905) have
reported that egg laying in chitons occurs only
after males have released sperm. Their state-
ments were apparently based upon small num-
bers of observations. From experiments with
Mopalia lignosa and Ischnochiton heathiana,
Vol. 4; No. 4
THE VELIGER
Page 209
Heath (1905) concluded that sperm or some hor-
monal product released by the male chiton was
necessary if spawning was to occur.
On the other hand, Grave (1932) states that
Chaetopleura apiculata females will spawn when
isolated from the males. Brewin (1942) ob-
served that females of Cryptoconchus porosus
would spawn prior to sperm release by the
males of the same species. In the first case
Grave was working with a fairly large number
of animals, while Brewin's observations were
made over a prolonged period of time. The re-
sults obtained by Mr. Daryl Sweeney and myself
are shown in Table 5.
Start of spawning not observed
7 Tray Sita ream poeecessary anny errors 7; | aes ee ] an Tae
-1: Afopalia ciliata (SowreRBy, 1840) | |
at
eae
| ——
face
1 Eee
ieee ner
i eae! aN]
| | =» |
while still spawning ed CaANK | |
| Bi] | ata
be paeie of spawning not observed | ea |
| | |
| ‘ | | f | |
|
t i
from larval development
i
; |
|
|
|
qt tine of spawning estimated
|
| High Low Tide
\. Low Low Tide }~
Figure 2
Sa ea
Figure 3: Mopalia, subspec. nov.
each.division = one hour
Page 210
THE VELIGER
Vol. 4; No. 4
Table 5:
Females Spawning
before males began to|when isolated from the
release sperm or when| males for periods of 3
Species Total| males did not release to 6 hours
sperm
Mopalia porifera (PILsBRyY, 1892) 4
Mopalia imporcata (CARPENTER in Pitssry, 1892) 3
Mopalia ciliata (SowERBy, 1840) 5 2
Mopalia spec. nov. I
Mopalia hindsi (REEVE, 1847) 2
Mopalia lowe (Pitssry, 1918) t
Mopalia lignosa (GouLp, 1846) . {
Ischnochiton mertensi (MIDDENDORFF, 1846) I
Totals:
The ten females not accounted for in columns 3 and 4 are those which spawned after one or more males had released sperm
It seems to me that the evidence, although
admittedly rather scanty, fails to support the
hypothesis that the release of sperm is the im-
mediate cause for spawning. The effects of
sperm release, or the effect of some as yet un-
determined male hormone upon the females
over longer periods of time, is unknown.
Summary
Observations of gamete release by 14 spe-
cies of California chitons are reported. Details
of behavior during breeding and of the develop-
ment of the larvae are given for Mopalia ciliata.
Gamete release in this and one other species
shows correlation with the tidal cycle.
Acknowledgment
I am most grateful to Mr. Daryl Sweeney
for permission to use some of his data in this
paper.
part thereof,
They form an integral and important
My thanks also to Mr. Allyn G.
Smith for his many helpful suggestions and en-
couragement.
Literature Cited
Berry, S, Stillman
1946, A re-examination of the chiton Stenoplax mag-
dalenensis (Hinds), with the description of a new
Species. Proc. Malacol, Soc. London 26 (6): 161
to 166, :
Brewin, B. I.
1942, The breeding habits of Cryptoconchus porosus
(Burrow). Trans. Proc. Roy. Soc, N. Zealand
72 (2): 186-190.
Christiansen, M. E,
1954, The life history of Lepidopleurus asellus (Speng-
ler)(Placophora). Nytt Mag. Zool, 2: 52-72.
Grave, B. H.
1932, The embryology and life history of Chaetopleu-
ra apiculata. Journ. Morph. 54 (1): 153-160,
Heath, H.
1899. The development of Ischnochiton. G. Fischer,
Jena, 1-90, pls. 1-5. EE LN ie.
=a)
1905. The breeding habits of chitons of the California
coast. Zool, Anz. 29 (12): 390-393,
1907, The gonad in certain species of chitons, Zool,
Anz. 32(1): 10-12.
Heath, H., & R. M. Higley
1912. The development of the gonad and gonoducts in
two species of chitons. Biol. Bull. 22 (2): 95-97.
Pilsbry, Henry A.
1892. Manual of Conchology, 14. Philadelphia. pp. |
to 350, pls. 1-68.
1893, Manual of Conchology, 15. Philadelphia. pp, |
to 133, pls. 1-17.
Trachydermon raymondi in British Columbia.
Nautilus 8: 57.
Thorpe, Spencer R., Jr.
1961, The species of Mopalia ciliata. Abstr. in Ann.
Rept. Amer. Malacol. Union, Bull. 28: 40,
Vol. 4; No. 4
Notes & News
Busycotypus (B.) canaliculatus
in San Francisco Bay
BY
RUDOLF STOHLER
Department of Zoology
University of California, Berkeley 4, California
Although the channeled whelk has been
found sporadically in San Francisco Bay pre-
sumably ever since 1938, it is only within
recent years that it seems to have become rel-
atively abundant, at least in certain localities.
The first observation is attributed by Dr. Leo
G. Hertlein of the California Academy of Sci-
ences to an unnamed collector who is alleged to
have obtained living specimens of this species
while dredging for 'oyster shell", a dietary
supplement for chickens, in 1938. This report
is quoted both in Puffer and Emerson (1954) and
in Hollister (1958), from a personal communi-
cation by Dr. Hertlein. However, the earliest
recorded specimens (three) in the collection of
the California Academy were obtained by Rogers
on December 20, 1950, ''at about the foot of Gil-
man Street!’ in San Francisco (2 on map, text-
fig. 1). Four additional specimens in the same
collection were obtained by C. H. Roof at Coy-
ote Point (4 on map, textfig. 1) in San Mateo
County, on March 27, 1954, In the collection of
the Geology Department at Stanford University
is a specimen which was dredged off Bay Farm
Island (1 on map, textfig. 1), Alameda County,
by P. J. Gambetta in February 1948. This
seems to be the earliest record of the species
in San Francisco Bay, as far as I have been
able to ascertain. It seems logical to suggest,
therefore, that the year 1938 is either a typo-
graphical error or an error of memory and that
actually the year 1948 represents the first oc-
currence of the species. In the collection of the
Department of Zoology at the University of Cal-
ifornia in Berkeley, there are additional rec:
ords, as follows:
A young specimen from Point Bluff (3 on map,
textfig. 1), Marin County, collected by Earl
Barnawell on June 14, 1953.
A mature specimen snagged with a fishhook
off Belmont Slough (5 on map, textfig. 1), San
Mateo County, by H. A. Dalton on June 29,
1958.
THE VELIGER
Page 211
A mature specimen brought in alive by Mr.
Charles Barry, from Alameda (7 on map,
textfig. 1), Alameda County, in May 1960.
While all the instances of observations rec-
orded thus far concern only one or very few
individuals, thereare now available also obser-
vations on larger numbers of individuals.
On December 22, 1958, Miss Laura Cantrel of
Oakland collected 26 specimens at the foot of
San Mateo Bridge(6 on map, textfig. 1), San
Mateo County. During the year 1961 two mem-
bers of the Northern California Malacozoologi-
cal Club, Mrs. Wanda Martin of Albany and
Mrs. Verna Wegner of El Cerrito, collected
over 100 specimens, ranging from very small to
apparently fully mature specimens, near Ala-
meda (7 on map, textfig. 1) in Alameda County.
These two collectors also picked up several
strings of egg capsules. On November 16, 196],
Mrs. Martin brought two living whelks and one
egg string to the Department of Zoology for ex-
hibit in the hall aquaria. The string, about 22
inches long when gently extended (but still
somewhat coiled) consists of 98 typical egg cap-
sules. Fromthese capsules about 40 to 50
young have emerged within the first two weeks
in the aquarium.
Only one or two additional young specimens
have been observed between December 1, 1961,
. San
~ Francisco
Figure I
Map of the central and southern portions of
San Francisco Bay, showing collecting stations of
Busycotypus (Busycotypus) canaliculatus (LINNAEUS, 1758)
1: Bay Farm Island; 2: foot of Gilman Street,
San Francisco; 3: Point Bluff; 4: Coyote
Point; 5: Belmont Slough; 6: foot of San
Mateo Bridge: 7: Alameda
(broken line is - 28 foot contour)
Page 212
1962.
and January 13,
Through the generosity and cooperation of
Dr. Leo G. Hertlein, Dr. Myra Keen, Mrs.
Martin, and Mrs. Cantrell, I have been able to
measure a considerable sample of all speci-
mens collected. In Table 1 I have summarized
the results.
There are several interesting points to be
Busyco-
typus canaliculatus (Linnaeus, 1758) is reported
observed from the map and the table.
from the east coast of the United States as oc-
curring in shallow waters. Yet there are three
distinct areas within San Francisco Bay where
Busycotypus has been obtained, namely, the west
shore and the east shore, respectively, of the
southern portion of the Bay, and the west shore
of the northern portion of the Bay. These three
portions are separated from each other by rel-
atively deep channels, The San Francisco Bay
Pollution Investigation project under the direc-
tion of Mr. R. A. Wagner has carried out nu-
merous dredgings in these channels, particu-
larly in the southern portion, but no living or
dead specimens of Busycotypus have been re-
covered (personal communication from Mr,
This poses a puzzle regarding the
Possibilities
Wagner).
distribution of the species.
coming to mindare: separate introductions;
accidental transport of egg strings or young
specimens by logs floating across the Bay.
Since the young are fairly large when they
emerge from the egg capsule —i.e., about a
quarter of an inch in greatest length — and are
not free swimming, the disrupted distribution
cannot be explained by the migrationof a ''free-
swimming" larval stage.
Reports on Busycotypus from the east coast
of the United States indicate a maximum length
of 73 inches. The largest specimen in Mrs,
THE VELIGER
Vol. 4; No. 4
Martin's collection, measuring 185 mm., just
about equals this maximum length. It seems
interesting to note the more or less gradual in-
crease in the maximum length observed over
the years.
It is apparent that Busycotypus canalicu-
latus is to be regarded as well established in
San Francisco Bay, and if our amateur collec-
tors do not eradicate the species by over-
collecting, this may prove a welcome addition
to the Bay fauna. There are no common native
shallow water species which equal Busycotypus
in size, except Polinices lewisii (Gould, 1847).
Therefore, Busycotypus may be most welcome
as dissection material for the many classes in
elementary zoology taught around San Francis-
co Bay.
Acknowledgment
The generous cooperation of Drs. Leo G.
Hertlein and A. Myra Keen, who made the col-
lections under their care available to me, and
of Mrs. Wanda Martin and Miss Laura Cantrell
is gratefully acknowledged.
Literature Cited
Hollister, S. GC.
1958.
Part I,
pls. 8-18.
Puffer, Elton L., & William K. Emerson
1954. Catalogue and notes on the gastropod genus
Busycon, Biol. Soc. Wash. 67: 115-150.
A review of the genus Busycon and its allies -
Paleontographica Amer. 4 (28): 59-126
EaWoce
Table 1:
Measurements (in millimeters) of Specimens of Busycotypus ( Busycotypus) canaliculatus (LINNAEUS, 1758)
from San Francisco Bay
a == = ———————— Fa oh >
Number of Locality
Date Collection Specimens Smallest Largest (see Map)
1948 Feb. Stanford University 1 96.6 Bay Farm Island
1950 Dec. 20 | Cal. Acad. Sci. 3 64.4 102.0 San Francisco
1953 June 14 | U. C. Zoology I 36.8 Point Bluff
1954 Mar. 27 | Cal. Acad. Sci. 4 105.1 161.8 Coyote Point
1958 June 29 | U. C. Zoology I 165.0 Belmont Slough
1958 Dec. 22 | Cantrell 26 30.1 136.5 San Mateo Bridge
igso0 May U. C. Zoology I 140.0 Alameda
1961 Martin over 100 34.0 185.0 Alameda
Vol. 4; No. 4
Recent Uses
of Non-binomial Works
BY
R. PUCGKER ABBOTT
Academy of Natural Sciences of Philadelphia
The erroneous use and implied adoption of
early, non-binomial works on mollusks by re-
cent authors have prompted this note of warning
(see Jean Cate,
Although there are many nomenclatorially
invalid works which employ descriptive polyno-
mials, I think four are worthy of mention be-
cause some workers are likely to use them in
the future.
The eleven volumes of the Neues Systema-
tisches Conchylien-Cabinet (1769-1795) written
by Martini and Chemnitz were declared non-
binomial and invalid nomenclatorially by Opin-
ion 184 of the International Commission on
Zoological Nomenclature in 1944. Evidently is-
sued as part of volume 10 was an index, the
"Namen Register'' (1788), which was assembled
by J. S. Schr6ter. Although not specifically
mentioned in Opinion 184, this index is merely
an alphabetically arranged list of vernacular
and Chemnitzian descriptive Latinnames, Such
entriesas Buccinum ex sanguine adspersum and
Bulla achatina sinistrorsa speak for the Index's
unavailability. Cate(1960, p.49) used the name
Mitra nigra (Schréter, 1788), but the earliest
valid usage appears to be Mitra nigra (Gmelin,
1791). It might be mentioned that Pfeiffer's
"Kritisches Register'' (Kassel, 1840, 112 pp.) is
binomial.
Another non-binomial work, deceiving at
first glance, is G. Karsten's Museum Leske-
anum, vol. 1, 320 pp., 3 pls., Leipzig, 1789.
the main, Karsten follows the Linnaean binomial
system and gives good descriptions and figure
references for species now credited to Gmelin,
1791. Fortunately, for the sake of stability, a
few polynomials appear, a fact which I believe
renders the entire work unavailable. Examples
are: (p.152) Mya Vulsella minor Chemnitz;
(p. 173) Arca Rhomboidalis J. Orient. Chemn.;
and (p. 186) Pinna haud ignobilis Chemn. His
describedand figured Nerita reticulata is there-
fore invalid, although it is listed in Sherborn's
Index Animalium for 1758 to 1800, p. 825.
In
Quite recently (T. A. Garrard, 1961, p. 32)
THE VELIGER
1960, and T. A. Garrard, 1961),
Page 213
erroneously credited the authorship of the ge-
nus Turris to P, L. S. Miller, 1766. Miller au-
thored in 1766 a folio edition of Knorr's Deliciae
Naturae Selectae — Naturalien - Cabinet, Nirn-
berg, vol. 1, 132 pp., with colored plates on
mollusks, The molluscan names on page 129 in
the explanations to the plates are binomial and
would normally replace some well-known names
in Cypraea, Mitra, Fortunately,
Miller was merely quoting Rumphius! 1705 pre-
Linnaean names, and, if one looks on page 128,
Voluta, etc.
one can find such polynomials as Sertularia pen-
nata folliculis bidentatis and Corallium acar-
baricum nigrum ramosum,
Miller's name valid, the type species would be
Incidentally, were
by monotypy and not by original designationas
stated by Garrard. I believe the earliest author
for Turris is Réding, 1798.
The fourth non - binomial work worth men-
tioning is the once-controversial Index to Gro-
novius' Zoophylacium Gronovianum published
by F. C. Meuschen in 1781. It was rejected in
Opinion 261 on August 10, 1954, by the Interna-
tional Commission on Zoological Nomenclature,
Literature Cited
Cate, Jean M.
1960. Range extension and synonymy for Mitra nigra
(Schréter, 1788). The Veliger 3 (2): 49-51,
Garrard) i VA.
1961. Mollusca collected by M. V. "Challenge" off the
East Coast of Australia. Journ. Malacol. Soc.
Austral. 5: 2-36.
New Name for
Strombus granulatus subsp. acutus
DuruaM, 1950, not Perry, 1811
BY
J. WYATT DURHAM
Department of Paleontology
University of California, Berkeley 4, California
Dr. Robert Robertson of the Academy of
Natural Sciences of Philadelphia has kindly
called my attention to the fact that my Strombus
granulatus subsp. acutus (1950, Geol. Soc. Amer-
ica, Mem, 43, pt. 2, p. 118, pl.27, figs. 1,2, and
5) from the Pliocene and Pleistocene of the Gulf
of California, is a homonym of Strombus acutus
Perry, 1811. Accordingly, my acutus is herein
renamed cortezianus, after the ''Sea of Cortez".
Page 214
The W. Mack
Chiton Collection
BY
ALLYN G. SMITH
California Academy of Sciences, San Francisco 18, California
A brief account of the Wilfred Mack col-
lection of chitons, now preserved in part in the
Mollusk and Invertebrate Zoology Collections
at the California Academy of Sciences, may be
of some value for future reference. Moreover,
the
was made have some interesting aspects.
circumstances under which the collection
Mr. Wilfred Mack, a garden specialist in
Pacific Grove, California, became interested in
collecting chitons through friendship with the
Rev. Elwood B. Hunter of Pacific Grove who
had been collecting chitons in the Monterey
area for some time and had become a specialist
in knowing where and how to collect them. Al-
though a beginner with no knowledge of chiton
ecology, Mr. Mack soon became an expert col-
lector himself under the tutelage of Elwood
Hunter.
Mr. Mack was an enthusiastic collector —
just how enthusiastic becomes evident on read-
ing the notes he kept of his collecting activities.
This formsa particularly valuable record which
beginning conchologists all too often do not take
the pains to prepare. These notes were pen-
written in a 5 x 8 inch spiral - bound notebook,
They begin (somewhat edited) as follows:
"), On 12/12/40, Thursday — Day clear —
low at 3:05 PM. Became interested in col-
lecting chitons after seeing Elwood Hunter's
collection. First trip with Elwood to rocks
about 0,3 mile south of Pt. Joe [on the] 17-
Mile Drive [Monterey Peninsula near Asi-
lomar]. Tide about 0,3 ft. Have beginner's
luck collecting: Mopalia ciliata, Tonicella
lineata, Lepidozona mertensi(a lovely red-
purple), Ischnochiton regularis (a perfect
blue [rare color phase] and many others),
Chaetopleura gemma (orange), Stenoplax
heathiana (in quantity), one Basiliochiton
heathi (red and green, Cyanoplax dentiens,
Mopalia muscosa, Rocks granite — quite
encrusted generally,'!
THE VELIGER
Vol. 4; No. 4
Chiton names in the above account have
been changed to modern equivalents as Mr.
Mack was not familiar at the time with chiton
taxonomy or the correct spelling of specific
names, This is unimportant and certainly no
criticism of his excellent job of note-taking.
Mr. Mack's notes cover a total of 69 sepa-
rate entries, beginning 12 December, 1940, and
ending around 10 July,1941, Just how persistent
he was is attested by his almost continuous col-
lecting efforts during moderate to low tide pe-
riods, day after day, rainor shine, and in heavy
Methods used
were the usual ones for collecting chitons, es-
surf as well as in calm weather.
pecially small ones, by turning rocks and in -
A great
deal of wading was done, occasionally out to
shoulder depth in the cold water of the Monterey
area, in order to bring in rocks from below the
intertidal zone,
lecting in the area myself over past years, I
can vouch for the degree of hardihood this type
of collecting demands without benefit of modern-
vestigating their undersides carefully.
Having done considerable col-
day swim-suits worn by SCUBA divers!
Both Mack and Hunter had their favorite
hunting spots in the area, especially along the
western flank of the Monterey peninsula from
Pt. Pinos (including the famous Great Tide Pool
at Lighthouse Point) to Fan Shell Beach and Cy-
press Point. Another favorite spot was what
they called the ''Carmelite Intrusion'', open sea-
shore area extending for some distance along the
shore of Carmel Bay beginning at the south end
of the long Carmel sand beach. A few visits
were made to the area between Yankee Point to
the mouth of Malpaso Creek, south of Point
Lobos, in the area now called the ''Carmel Ri-
viera.'' One entry reads as follows:
"60. May 22-23, Chas. Jones brings me
in 2 rocks — and one on 23rd — taken about 3
miles out of slightly south of Pt. Joe at be-
tween 60-70 fathoms (300-400 ft.), rocks
catching on [fishermen's set-line] hooks.
10 specimens (one white) mostly the same
apparently. "'
The chitons from these rocksare of unusua-
al interest as thereare possibly twoormore
new species among them, illustrating the value
of getting the cooperation of friendly fishermen.
Here is another interesting entry:
"23, Sat. Jan.25— Tide, -0.8 and plenty
low. Do not have much time so go just in-
side gate of lighthouse reservation, WOW
— find one black specimen with white stripe
Vol. 4; No. 4
THE VELIGER
Page 215
ee aN ds ee a ee lO
down back, slightly mottled. [possibly a
color phase of Ischnochiton radians]; 7 Cal-
listochiton palmulatus and mirabilis; 4 Pla-
ciphorella velata; 2 Mopalia ciliata; several
nice Lepidozona mertensi, including one
deep purple [color phase]; a couple of Isch-
nochiton
regularis; several Stenoplax
heathiana; Nuttallina californica all over
rocks; and 1 blue Lschnochiton radians
(deep blue). — Weather threatening but ocean
not rough, ''
Mack finally lost interest in collecting and
turned over his entire collection, mostly all
still preserved wet, to the California Academy
They could not be
worked on immediately for lack of time, but in
March 1948 all specimens (a few had dried up)
were washed for several days in running water
of Sciences on 5 July, 1947.
to leach out the tormalin and then transferred
to alcohol.
the formalin, many specimens are still in good
In spite of the unfortunate effect of
condition and have been placed in the collection
of the Department of Invertebrate Zoology at the
Academy where they are available, along with
the collecting notes, for subsequent reference
and research,
California Brown Cowrie
in Central California
BY
ALLYN G. SMITH
California Academy of Sciences, San Francisco 18, California
Mr. Robert P, Sikora, University of Cali-
fornia Zoology student supplies information to
the effect that the California Brown Cowry (Cy-
praea spadicea Swainson) was taken alive in the
summer of 1961 by members of the Marin Skin
Divers Club. According to the report, about 20
specimens were collected with SCUBA equip-
ment in a depth of 80 feet on a rock off Cypress
Point, Monterey County, California. One of
these specimens, measuring 57.7 mm. in length,
35.8 mm. in width, and 29.3 mm. in height, has
been placed in the collection of the California
Academy of Sciences through the generosity of
Mr. Al, Giddings, a member of the Marin Club,
This specimen isa rather light-colored one and
is normal in all respects.
The finding of this cowry alive in an area
considerably to the north of its published range
is notable, It confirms an old record of a living
specimen found at Pacific Grove in 1910 or
thereabouts by Nettie (Mrs. Charles S.) Facken-
thall, which has been cited several times in the
literature. The location of this particular shell,
formerly in Mrs, Fackenthall's collection, is
now not known.
Arion ater (Linnaets)
in California
BY
ALLYN G. SMITH
California Academy of Sciences. San Francisco 18, California
The large, black European slug, Arion ater
(Linnaeus), has so far been reported from the
West Coast only in Washington and Oregon. It
now becomes an unfortunate duty to record that
this garden pest has reached California.
California specimens were collected by
field men of the Del Norte County Department
of Agriculture in nurseries in Crescent City
and vicinity, and inland five miles northwest of
Gasquet, Del Norte County.
They were forwarded to the California
Academy of Sciences for identification by Mr.
L. J. Garrett, Agricultural Commissioner of
Del Norte County, whose vigilance in the appar -
ently early discovery ot this new and potentially
injurious agricultural and garden pest is to be
commended. One can only hope that the Del
Norte County infestation can be eradicated be-
fore the species spreads farther south in the
State.
Arion ater reaches an expanded length of
4.5 to 5 inches. It may be recognized rather
easily by the rugose mantle and the coarse,
elongated tubercles over the back and sides of
the body. Like other Arions and the greenhouse
slug [Milax gagates (Draparnaud)], it contracts
into a semicircular shape when inactive or dis-
turbed. An excellent figure may be found in
Pilsbry (1948, p. 668, fig. 365). There are sev-
eral color phases, the most prevalent being
black or brownish.
My first collecting experience with this
species was on October 9 and 10, 1943, in the
Page 216
THE VELIGER
Vol. 4; No. 4
i a ee
garden of Mr. Linsley W. Ross, 12223 Eighth
Avenue, Seattle NW, when a total of 19 were
taken in moist to rainy weather conditions,
Many more were crawling among garden plants.
There were two color phases:
1, Back and mantle reddish-brown. Foot-
fringe a brilliant terra - cotta, crossed by
alternate thin black lines and wider black
stripes. Occasional dark blotches on the
mantle occurred on some specimens of this
lighter colored phase. Sole light colored.
2. Back and mantle dark chocolate brown,
Mantle with occasional small, variable-
sized, black blotches but not prominent
enough to take away from the generally
dark, unicolored aspect. Foot-fringe not
differently colored from the back, with the
same type of black striping described for
the lighter phase. Striping of the foot-
fringe extends around and under the sole
for 2 or 3 mm. on some specimens. Sole
slate colored.
These colors fade out almost entirely after
specimens have been preserved in alcohol for a
day or so. Mr. Ross stated he first recognized
this large slug as a garden pest in Seattle about
1940 because of its depredations on bearded
iris and succulents.
Pilsbry (1948, p. 670) includes an occur-
rence of Arion ater in Portland, Oregon, in 1946
(B. G. Thompson, July 9, 1946).
Methods ct Techniques
Notes on Cleaning Mollusks
BY
ALLYN.G. SMITH
California Academy of Sciences, San Francisco 18, California
After reviewing recently the excellent sug-
gestions contained in the Second Edition of the
AMU's "How to Collect Shells”, it occurred to
me to record several cleaning methods that
have been used successfully at the California
Academy of Sciences that may be helpful to
some shell collectors.
1. There is on the market a supersonic device
with the trade name ''Sonblaster''. While rather
expensive (it costs around $120.- for the unit),
it is the only equipment used so far that will
clean many kinds of shells, The container is
filled with water, which is agitated by a power
unit transmitting high-frequency sound waves.
The power is adjustable. Hands or fingers are
not affected except for a slight tingling sensa-
tion. For larger shells we merely dip the shell
to be cleaned into the agitated water several
times with the result that all loose dirt and de-
tritus is literally shaken off, falling to the bot-
tom of the container in a cloud. Tiny shells we
place in a small water-filled glass tube and dip
this in and out several times. The method is
especially fast and efficient for cleaning shells
with a heavy periostracum which one wishes to
preserve intact; it is excellent for cleaning the
girdles of chitons, especially Mopalias and oth-
ers with hairy or spiculose decoration; and it
does a beautiful job on the sutures of small land
species like Vertigo and Gastrocopta as wellas
"cleaning their teeth'' if they have any within the
apertures. For more solid, heavy shells, full
power is needed, but for more delicate speci-
mens reduced power is recommended to pre-
vent shattering. (Incidentally, this equipment
cleans eyeglasses and all sorts of small parts
and gadgets, being an excellent remover of
grease as well as of dirt.)
2. Leslie Hubricht's method (p.77) of pre-
serving slugs by anesthetizing and killing in
water with five to ten percent chloretone in so-
lution also works well with some marine spe-
cies before final preservation. This is even
better than the old method of "killing by drown-
ing'’ in fresh water, which has been used on
specimens of Onchidella with fully expanded
specimens as a result, This same method also
works well on Velutina and Lamellaria when the
shell is to be preserved with the fully expanded
animal.
3. A strong household bleach is a useful
cleaning agent, if used judiciously. I have found
that cleaning freshwater mollusks, such as Go-
niobasis, Fluminicola, Amnicola, and Hydrobia
with a bleach used at full strength for one or
two minutes only will remove all adhering algal
or other extraneous detritus, leaving clean,
bright, shining shells that are a credit to any
collection. Leaving in strong bleach too long,
however, will remove the periostracum, an un-
desirable result unless there is good reason for
removing it purposely to expose the outer shell
layer. Preliminary trial to determine just how
long to leave specimens in the bleach solution is
recommended. Unfortunately, this method will
not work on old, long-dried-out freshwater spe-
cimens; it is effective only on shells fairly re-
cently collected. Killing in alcohol prior to
cleaning with bleach seems not to inhibit this
cleaning process,
Vol. 4; No. 4
THE VELIGER
Page 217
Information Desk
What's the Difference?
Holotype - Paratype - Syntype - Hypotype
BY
R. STOHLER
Department of Zoology
University of California, Berkeley 4, California
In the 200 years that have passed since the
first species of plants and animals were de-
scribed by Linnaeus, a great many designations
Some of
"pri-
types while others are concerned with
for type material have been invented.
these refer to what may be classed as
mary"
"Secondary'' types.
Primary type material would include all the
specimens which were used by the original au-
thor in preparing his original description. Sec-
ondary type material, on the other hand, would
encompass those specimens which were used by
other authors— and, of course, possibly even by
the original author at a subsequent date—to ei-
ther amplify or emend the original description,
or to replace the original type specimen(s) if
lost or destroyed. To the first group should be
counted the holotype, the paratype(s), the syn-
type(s), and, under certain circumstances, the
hypotype(s), while the second includes the neo-
type(s), the lectotype(s), and others.
The holotype is defined as the single speci-
men taken as ''THE TYPE"! by the original au-
thor of a species or subspecies. The paratype
is a specimen or one of several specimens
which were used by the original author as the
basis of a new species or subspecies, in addi-
tion to the holotype. A syntype is one of several
Specimens of equal rank used in the original
description without, however, being singled out
as "holotype’'; the word "'cotype” is, fundament-
ally, a synonym of syntype; it is no longer used.
A hypotype, finally, is a described, listed or
figured specimen whether or not it is included
in the discussion of the new taxon.
Early authors were rather lax in their atti-
tude toward type specimens. It was not an un-
common practice to replace the original type
specimen with a better ''type’' specimen, when
it became available. Also, it was a fairly fre-
quent practice for a museum to exchange type
material, retaining one or two specimens of a
given species, Today, when we are aware of the
many difficulties attendant upon inadequate do-
cumentation, there is no excuse for less than
the utmost care in selecting and preserving type
specimens. This is true even where a species
may have been found to be invalid for one of
several possible reasons. However, the discov-
ery of the so-called sibling species has added
further strength to the need for care, Sibling -
species are morphologically identical with each
other, or at least so nearly so that even fairly
careful examination does not reveal the fact
that they are different species; yet sibling-spe-
cies are reproductively isolated in spite of the
great similarity of the adult individuals. Often,
too, sibling-species may occur in the same io-
cality and it is not impossible that they might
even occupy the same habitat. From this it be-
comes evident that the conscientious taxonomist
must base his description of a new taxon upon a
Single specimen—the holotype. This specimen
No
matter what discoveries may be made at a later
thus becomes actually the name-bearer.
time, the holotype remains the ultimate author -
ity regarding that particular species and its
name. It is not impossible that even with great
care exercised in the examination of the type
population, a sibling species might be inadvert-
ently drawn in and included in the description.
Later students will have the task of separating
out the specimens which belong to the one, the
original species, and the specimens properly
assigned to the sibling-species. If the original
author did not select a "holotype", there would
be uncertainty as to which is the original spe-
cies and which is the sibling-species, which lat-
ter must, of course, be given a different name.
There! seems to be a growing trend to
include as part of the description of a new spe-
cies as full an appraisal as possible of the var-
iability in the Original population. This is
actually most desirable, although not always
possible. All specimens from this particular
population become paratypes, except for the one
select specimen, the holotype. The paratypes,
as pointed out above, may, however, include
specimens of a different species. But this pos-
sibility is more or less implied by the very fact
that these specimens are designated as para-
types. Sometimes it is possible for an author
to include in his appraisal of the variability of
the new species material other than the original
group collected at the type locality. Many au-
Page 218
thors call such specimens also paratypes. This
is, to our way of thinking, unfortunate since it
does not clearly distinguish between the non-
holotype specimens from the type locality and
the non-holotype specimens from other places.
Since it is possible that paratypes encompass
sibling-species—and we refer here to paratypes
from the type locality ——it is even more proba-
ble that specimens from other localities may
include sibling-species. Therefore, it
only fitting that such subordinate
seems
"paratypes"
be clearly distinguished in the original descrip-
The term "hypotype"'’ seems to fit the re-
to be no
ruling by the International Commission on Zo-
tion.
quirements well. And there seems
ological Nomenclature against this use of the
term which allows a clear separation of speci-
mens with different probabilities of uncertain-
ty as to proper identity. If there is nothing
more to recommend this differentiation than the
fact that it may be of assistance to future work-
ers, we think it sufficient justification to use
the term ''hypotype'' in this sense. The defini-
tion of the paratype would then necessarily in-
clude the specification that it must come from
the type locality while the hypotype does not.
Books, Periodicals, Pamphlets
THE GIANT AFRICAN SNAIL —
A PROBLEM
IN ECONOMIC MALACOLOGY
by Albert R. Mead
Professor of Zoology
University of Arizona
University of Chicago Press, 257 pp.,
15 photographic illustr. November 28,
196). $7250.
This book is unique. It is the only one of
any scope dealing with the growing economic
problems caused by land snails in general, and
by the Giant African Snail in particular. There
are good and timely reasons for such a refer-
ence work, which is the first in any language
assembling knowledge of the economic effect of
land mollusks, both snails and slugs. For this
field of biological study, the author uses the
term "Economic Malacology".
The Giant African Snail is a growing men-
ace to be reckoned with. This five to six inch
monster, while not a champion for size among
land snails, is an "exceedingly hardy, tenacious,
THE VELIGER
Vol. 4; No. 4
variable and adaptable molluscan pest with a
high reproductive potential and remarkably few
natural enemies". Once started it is practically
impossible to eradicate, and most man-devised
methods for its control have not met with any
signal success. The spread of this snail pest
during World War II, including its build-up on
the Hawaiian Islands, its fantastic ability to re-
produce causing population explosions in the
species, and its consequent depredations result-
ing from its omnivorous food habits all have
served to create a ''Giant African Snail Prob-
lem" of primary importance to the world.
Much has been written about the Giant Af-
rican Snail in the world press and in scientific
and agricultural journals. In recent years it
has been the subject of considerable research
and has led to the expenditure of much money
to determine its present and potential economic
danger and to develop successful means of con-
trol, Mead brings all of this scattered informa-
tion together in organized form. This is a task
for which he alone is preeminently qualified,
having been personally associated with the Gi-
ant African Snail Problem for more than ten
years. He has traveled many thousands of
miles to gather firsthand data, has investigated
the possible use of the snail as a food for peo-
ple and animals, andis now studying a means of
control by infecting it with a specific virus dis-
ease.
The book opens with a well-documented
chapter on the present wide dispersal of the Gi-
ant African Snail, mainly by man, from its ori-
ginal home in East Africa. It continues with
chapters on the factors favoring dispersal and
survival, on its economic status as an agri-
cultural pest, and on the various methods of
control—chemical, mechanical, biological, leg-
islative, and last, but by no means least, its
control through human use as a possible food
for poultry and livestock. There is an exceed-
ingly interesting chapter on the phenomenon of
decline following population explosion, the
causes of which are not thoroughly understood
and which could well be a subject for future in-
tensive biological investigation leading, per-
haps, to more effective control measures. The
bibliography at the end of the book is a verita-
ble gold mine of source information, covering
over 40 pages, including 563 author listings and
881 separate titles.
Mead's work serves to bring into full focus
the various attempts to control other snail and
slug pests, with their successes and more fre-
quent failures, together with the dangers inher-
ent in approaching control problems without
sound scientific research by qualified experts
Vol. 4; No. 4
THE VELIGER
Page 219
to determine the possible future effects of the
means used, The fact that the State of Califor-
nia alone has spent over half a million dollars
to date on the unsuccessful eradication of three
exotic snail species points to the importance of
the whole snail pest problem and its potential
effect on man's future food supply.
This book is highly recommended as re-
quired reading for all workers in the field of
economic entomology and other types of agri-
cultural pest control. Conchologists and mala-
cologists interested in this very practical side
of their special field will welcome it as an im-
portant contribution. In fact, all zoologists and
biologists interested in the broad ecological and
other problems created by the introduction of
exotic species as exemplified by the lesson pro-
vided by the Giant African Snail will find it a
source of valuable information and ideas,
AGS
FOUR SPECIES OF CHITONS
FROM THE PANAMIC PROVINCE
(MOLLUSCA: POLYPLACOPHORA)
by Allyn G. Smith
Gabitee Acadei Sci, Hourth Sex.;
No. 4, pp. 81-90, pls. 8-9.
August 31, 1961.
Proc.
Vol. 30,
Four species of chitons, two of which are
described as new, are discussed in this paper.
Three are from western Mexico, one from Pan-
Nuttallina crossata
anne, Wa So@ewes evees
Berry (= N. mexicana Pilsbry in Pilsbry and
Lowe), Chaetopleura (Pallochiton) euryplex
Berry (= C, raripustulosa Pilsbry in Pilsbry
and Lowe), Ischnochiton colimensis A. G. Smith,
new species (=I. lowei Pilsbry in Pilsbry and
Lowe), and Acanthochitona tabogensis A. G.
Smith, new species (= A. panamensis Pilsbry in
Pilsbry and Lowe). This work adequately dis-
poses of the four nomina nuda of Pilsbry, 1932.
The discussions include the known occurrences
of the species and a comparison of them with
similar forms. The four species are beautiful-
ly illustrated in color on two plates.
LGH
LARGE TEREBRAS (MOLLUSCA)
FROM THE EASTERN PACIFIC
by G Dallas Hanna and Leo G, Hertlein
California Academy of Sciences
Proc Calatiavcads ioCi.. HoOUTth) Sexe,
Vol. 30, No. 3, pp. 67-80, Pls. 6-7.
August 31, 1961.
This important paper brings the taxonomy
of the larger Terebras of the West Coast up-to-
date and is a valuable addition to our knowledge
of the family Terebridae.
discussed in detail with a thoroughness for
which the authors are noted.
Five species are
These are:
T. lingualis Hinds: Upper Gulf of Califor-
nia to northern Ecuador, in 13-110 m. (7-
60 fms.).
T. ornata Gray: Gulf of California (Santa
Margarita Island to Puerto Pefiasco) south
to Panama and the Galapagos Islands, in 9-
143 m. (5-80 fms.).
T. robusta Hinds:
Rio Esmeralda, Ecuador, and the Galapagos
Islands, in 7-33 m. (8-14 fms.).
Guaymas, Mexico, to
T. strigata Sowerby: Gulf of California
(Magdalena Bay to Puerto Escondido) and
south to Paita, Peru; Sucorro [sland; Gala-
pagos Islands; in 18-22 m.(10-12 fms.) and
perhaps deeper.
‘I. dumbauldi, n. sp.:
ma.
Type locality: Pana -
Included is a key to these species, which
will be useful in identification, together witha
list of Terebras that have at times been re-
ferred to the foregoing species.
Outstanding features are the two three-
color-process plates illustrating each species.
These are the work of Dr. and Mrs. Hanna,
with printing done in the Calitornia Academy's
Shop. These plates exemplify the great advan-
tage of the use of color illustrations for scien-
tific work,
AGS
Page 220
CONTRIBUTION A UN CATALOGO
DE LOS MOLOSCOS
GASTEROPODOS CHILENOS
DE AGUA DULCE
by J. Stuardo
Gayana-Zoologia, No. 1, pp. 1-32, text-
figs. (distributional maps) 1-5, 10
drawings of shells. 1960.
Gayana is a new publication by the Instituto
Central de Biologia, Universidad de Conceptién,
Casilla 301, Conceptién, Chile, of which the di-
rector is Hugo Barrales. There are two de-
partments: Botany, headed by Mario Ricardi;
and Zoology, headed by José Stuardo. The pub-
lication (in Spanish) is named for Claudio Gay
(1800-1873),
Ciudadano Honorario de Chile"', whose portrait
"eminente Naturalista Francés y
appears in the frontispiece.
The present paper is a welcome contribu-
tion, bringing together scattered information on
the ranges of Chilean freshwater gastropods in
the form of a catalogue of species with refer-
ences to the original publication of each species
name. This fills a distinct need, for, as Stuardo
points cut, there are no existing lists of Chilean
land or freshwater mollusks. Two lists dealing
with marine mollusks of the area are cited: one
by Dall (1910, Report ona collection of shells
from Peru, Proc. U.S. Nat. Mus., vol. 37); the
THE VELIGER
Vol. 4; No. 4
other by Carcelles and Williamson (1951, Cata-
logo de los moluscos marinos de la Provincia
Magelldnica, Rev. Inst. Nat. Inv. Ciencias Natu-
rales, Zool., vol. 2, no. 5). Now, there is a need
for distributional catalogues of land gastropods
and freshwater pelecypods, as well as a com-
plete list of Chilean marine mollusks, which
Gayana hopes to supply.
AGS
GASTROPODIA
Vol. 1, Nos. 4-5. Dr. Glenn R. Webb,
Ed. (Route 3, Box 361, Conway, South
Carolina). Issued November 27, 1961.
Contains the following articles by Dr.
Webb:
"Studies on the Sexology and Development
of the Genitalia of Glyptostoma gabrielense
Pilsbry"' (pp. 29-30).
"The Phylogeny of American land Snails
with Emphasis on the Polygridae, Arionidae,
and Ammonitellidae" (pp. 31-52, pls. 15-23). To
be continued.
AGS
THE VELIGER is open to original papers pertaining to any problem
concerned with mollusks from the Pacific Region.
This is meant to make facilities available for publication of articles
from a wide field of endeavor. Papers dealing with ecological, morpho-
logical, anatomical, physiological, distributional, taxonomic, etc. aspects
of marine, fresh water or terrestial mollusks from any region bordering
on or situated within the Pacific Ocean, will be considered. Even topics
only indirectly concerned with mollusks may be acceptable.
It is the editorial policy to preserve the individualistic writing style of
the author; therefore any editorial changes in a manuscript will be 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, photograph-
ing, etc., of mollusks or other invertebrates. A third column, entitled
“INFORMATION DESK,” will contain articles dealing with any prob-
lem pertaining to collecting, identifying, etc., in short, problems en-
countered 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 , 814” by 11”, double spaced and accompanied by a carbon copy.
EDITORIAL BOARD
Dr. Donarp P. Assort, Associate Professor of Biolog y
Hopkins Marine Station of Stanford University
Dr. J. Wyatt Duruam, Professor of Paleontology
University of California, Berkeley
Dr. E. W. Facer, Associate Professor of Biology
Scripps Institution of Oceanography, University of
California, La Jolla
Dr. Capet Hann, Associate Professor of Koology
University of California, Berkeley
Dr. G. Datias Hanna, Curator, Department of Geology
California Academy of Sciences, San Francisco
Dr. Joet W. Hepcretn, Professor of Zoology
Director of the Pacific Marine Station, Dillon Beach
Dr. Leo G. HERTLEIN, Associate Curator, Department of Geology
California Academy of Sciences, San Francisco
Dr. Myra KEEN, Associate Professor of Paleontology and
Curator of Concholog y
Stanford University
Dr. FRANK PiTELKA, Professor of Zoology
University of California, Berkeley
Mr. A.LLyn G. Smiru, Associate Curator, Department
of Invertebrate Zoology
California Academy of Sciences, San Francisco
Dr. Ravpu I. Smitu, Professor of Zoology
University of California, Berkeley
EDITOR
Dr. RupDoLF STOHLER, Associate Research Koologist
University of California, Berkeley
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